Volume 9 Number 2 2014

*s

AUSTROBAILEYA

A Journal of Plant Systematics

and

Conservation Biology

Queensland Herbarium

Department of Science, Information Technology, Innovation and the Arts

Queensland

Government

Volume 9

Number 2 2014

A Journal of Plant Systematics

and

Conservation Biology

Queensland Herbarium

Department of Science, Information Technology, Innovation and the Arts

Queensland

Government

Editorial Committee

P.I.Forster (editor)

P.D.Bostock (technical advisor)

G.P.Guymer (technical advisor)

Graphic Design

Will Smith

Desktop Publishing

Aniceta Cardoza

Austrobaileya

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

Vol. 9, No. 1 was published on 17 December 2013

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ISSN 0155-4131

© Queensland Herbarium 2014

Web site: http://www.qld.gov.au/environment/plants-animals/plants/herbarium/

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

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

conservation biology and botanical history), with special emphasis on taxa from Queensland.

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

of the Queensland Herbarium, Department of Science, Information Technology, Innovation and

the Arts.

Austrobaileya 9(2): 155-320 (2014)

Contents

A taxonomic revision of Diospyros L. (Ebenaceae) in Australia

L.W. Jessup .155-197

Diploglottis alaticarpa W.E.Cooper (Sapindaceae), a new species from

Queensland’s Wet Tropics

W.E.Cooper . 198-202

Ptilotus senarius A.R.Bean (Amaranthaceae), a new species from northern

Queensland

A.R.Bean . 203-206

Diversity on a tropical sky island: two new species of Plectranthus L.Herit.

(Lamiaceae) from the Hann Tableland, northeast Queensland

P. I. Forster . 207-215

Four new Queensland species allied to Solarium ellipticum R.Br. (Solanaceae)

A.R.Bean .216-228

Systematics of Tephrosia Pers. (Fabaceae: Millettiae) in Queensland: 1. A

summary of the classification of the genus, with the recognition of two

new species allied to T. varians (F.M.Bailey) C.TWhite

L.Pedley . 229-243

C.T. White’s botanical survey and collections from Papua in 1918

A.R.Bean . 244-262

The botanical collections of Ebenezer Cowley

J.L.Dowe . 263-278

Plectranthus acariformis PI.Forst. and P. geminatus PI.Forst. (Lamiaceae):

new species from south-east Queensland

P. I. Forster . 279-291

Six new species of Bonamia Thouars. from northern Australia

R.W. Johnson . 292-310

Pluchea tenuis A.R.Bean (Asteraceae: Plucheinae), a new species from Cape

York Peninsula, Queensland

A.R.Bean .311-313

New combinations for Senegalia Raf. and Vachellia Wight & Arn. species

(Mimosaceae) that occur in Australia

L.Pedley .314-315

Lectotypification of F.M.Bailey names in Conyza (Asteraceae), Diplanthera

(Bignoniaceae), Pygeum (Rosaceae), Rhaphidophora (Araceae) and

Tetracera (Dilleniaceae) based on E.Cowley collections

P.I.Forster & J.L.Dowe .316-318

Aeschynomene micrantha (Poir.) DC. is a synonym of A. brevifolia L.f. ex

Poir.

A. E. Holland .319-320

A taxonomic revision of Diospyros L. (Ebenaceae) in Australia

L.W. Jessup

Summary

Jessup, L .W. (2014). A taxonomic revision of Diospyros L. (Ebenaceae) in Australia. Austrobaileya

9(2): 155-197. The Australian taxa of the genus Diospyros are revised. Twenty two species (21 native

and 1 naturalised) are recognised. Six species are described as new: Diospyros granitica Jessup,

D. peninsularis Jessup, D. pluviatilis Jessup, D. rheophila Jessup, D. uvida Jessup and D. yandina

Jessup and new combinations are made for D. hemicycloides (F.Muell. ex Benth.) Jessup (based

on Maba hemicycloides F.Muell. ex Benth) and D. laurina (R.Br.) Jessup (based on Maba laurina

R.Br.). Descriptions, distribution maps, illustrations and identification keys based on male or female

material are provided.

Key Words: Ebenaceae, Diospyros, Diospyros areolifolia, Diospyros australis , Diospyros

calycantha, Diospyros compacta, Diospyros fasciculosa, Diospyros geminata, Diospyros granitica ,

Diospyros hebecarpa, Diospyros hemicycloides, Diospyros humilis, Diospyros kaki, Diospyros

laurina, Diospyros littorea, Diospyros mabacea, Diospyros maritima, Diospyros peninsularis,

Diospyros pentamera, Diospyros pluviatilis, Diospyros rheophila, Diospyros rugosula, Diospyros

uvida, Diospyros yandina, Australia flora, taxonomy, new species, identification key, distribution

maps

Laurence W. Jessup, Queensland Herbarium, Department of Science, Information Technology,

Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066,

Australia, Email: Laurence.Jessup@dsitia.qld.gov.au

Introduction

Diospyros L. is a pantropical and subtropical

genus of over 500 species (Frodin 2004;

Wallnofer 2001, 2004). Robert Brown (1810)

was the first to describe Australian species

under the genera Diospyros L., Maba

J.R.Forst. & G.Forst. and Cargillia R.Br.

Bentham (1868) continued to recognise

these three genera although Mueller (1867)

had transferred several species described by

Brown and himself under Maba and Cargillia

to Diospyros. Hiern (1873) and Bailey (1900)

continued to recognise the genus Maba but not

Cargillia. Bakhuizen van den Brink (1936-

1955) produced a multilevel classification of

Diospyros recognising Maba and Cargillia as

subgenera along with three other subgenera

not represented in Australia. He also placed

several Australian species under the large

D. ferrea (Willd.) Bakh. group of taxa.

Kostermans (1977) considered D. ferrea to be

confined to India and Ceylon (Sri Lanka) and

reinstated several Australian taxa to species

under Diospyros. Smith (1981) followed

Accepted for publication 4 August 2014

Kostermans and rejected the broad concept of

D. ferrea in his treatment of Diospyros in Fiji.

Attempts at infrageneric classification

since Bakhuizen van den Brink (1936-1955)

have been proposed by White (1980) and Singh

(2005) but molecular studies of Duangjai et

al. (2006, 2009) reveal 11 well-resolved major

clades in Diospyros and indicate that all earlier

infra-generic classifications are artificial. The

authors acknowledge that although their study

is the most extensive to date for Diospyros, the

sampling is still too limited to propose with

confidence a new infrageneric classification.

Materials and methods

This revision is based on herbarium

collections in BM, BRI, CANB, CNS

(previously QRS), E, K, L, MEL and NSW

and some field observations by the author. All

specimens cited have been seen by the author.

Descriptions of flowers were prepared from

material preserved in FAA or 70% alcohol and

glycerol or reconstituted by briefly boiling in

water. The descriptions of fruit were prepared

from both dried and alcohol preserved

material. Common abbreviations in the

specimen citations are FR (Forest Reserve),

156

LA (Logging Area), NP/NPR (National Park/

National Park Reserve), R (Reserve), SFR/

SF (State Forest Reserve/State Forest), TR

(Timber Reserve). Electronic images of type

specimens at the JSTOR website are indicated

as “online image!”.

Taxonomy

Diospyros L, Sp. PI. 1057 (1753). Type

species: Diospyros lotus L. (lecto: fide Britton

& Brown 1913: 720).

Maba J.R.Forst. & G.Forst., Char. Gen. PI. 61,

t. 61 (1775); ed. 2:121, t. 61 (1776); ed. 3:61, t.61

(1776); Diospyros L. section Maba (J.R.Forst.

& G.Forst.) Fliern (as section Ferreola nom.

illegit); section Maba (J.R.Forst. & G.Forst.)

Singh, Monogr. on Indian Diospyros

(Persimmon, Ebony) Ebenaceae 23 (2005),

comb, superfl. Type species: Maba elliptica

J.R.Forst. & G.Forst.

Cargillia R.Br., Prodr. 526 (1810); Diospyros

section Cargillia Hiern, Trans. Cambridge

Philos. Soc. 12: 146,155 (1873). Type species:

Diospyros cargillia F.Muell. nom. inval. nom.

nud.

Diospyros subgenus Cargillia (R.Br.) Bakh.,

Gard. Bull. Straits Settlem. 7: 162 (1933).

Type species: not designated.

Trees or shrubs, commonly dioecious,

rarely monoecious, andromonoecious or

polygamomonoecious. Indumentum of

simple hairs or 2-armed with one arm very

short (submedifixed). Wood commonly

hard and dark, latex absent. Leaves simple,

alternate, usually distichous, entire, often

with nectaries on underside, petiolate,

exstipulate. Inflorescence axillary,

determinate, cymose in males, often solitary

flowers in females. Flowers actinomorphic,

Austrobaileya 9(2): 155-197 (2014)

hypogynous, unisexual, bracteate. Staminate

flowers with vestigial ovary. Pistillate

flowers with or without staminodes. Calyx

gamosepalous, persistent, often accrescent

in fruit, 3-6-lobed, lobes valvate. Corolla

gamopetalous, 3-6-lobed, lobes imbricate or

contorted. Stamens adnate to base of corolla

tube or attached to receptacle, isomerous,

diplostemonous or polystemonous; when

isomerous alternate with corolla lobes, when

diplostemonous then in 2 cycles; anthers

usually linear-lanceolate or oblong; longer

than filaments; filaments free or connate at

base, often connate in pairs alternating with

singles; glabrous or variously pubescent;

anthers bilocular, dehiscent by longitudinal

slits or these sometimes initially resembling

apical pores; pollen grains tricolporate.

Gynoecium of 2-8-connate biovulate

carpels as indicated by the number of styles

or stigmas, each carpel with a longitudinal

false secondary septum and then each locule

appearing 1-ovulate, or without a false septum

and 2-ovulate, with apical-axile placentation;

ovules pendulous, oblong, anatropous with

4, 6, 8, 10, 12, 14 or 16 per ovary. Ovary

commonly pubescent, sometimes glabrous;

styles distinct or connate, sometimes obscure;

stigmas often bifid. Fruit a leathery or fleshy

berry, pubescent, glabrate or glabrous; seeds

1—10(—16), pendulous, usually oblong and

laterally compressed, often segment shaped,

with thin testa and abundant, hard, often

ruminate endosperm; embryo straight or

curved, cotyledons foliaceous.

A genus of over 500 species worldwide, in

Asia, tropical Africa, South America, tropical

and subtropical regions of Australia and

Pacific islands. In Australia a genus of one

naturalised and 21 native species, 13 endemic.

Key to Australian species of Diospyros using male flowers

1 Flowers 4-6-merous; filaments adnate to corolla tube.2

1. Flowers mostly 3-merous (rarely 4 or 5-merous); filaments not adnate to corolla tube . . 12

2 Filaments glabrous or with a few short appressed hairs.3

2. Filaments pubescent or puberulent, hairs generally not appressed.8

Jessup, Diospyros in Australia 157

3 Petiole and midvein glabrous; petiole nearly flat above; calyx lobed to less

than one quarter of its length; lobes 0.4—1(—1.5) mm long; corolla tube

6-8 mm long.4

3. Petiole and midvein not glabrous; petiole channelled adaxially; calyx

lobed to two thirds of its length; lobes 1-2.5 mm long; corolla tube

1-3 mm long.6

4 Stamens less than 12; free part of filaments and base of anthers and

connectives with short appressed hairs.3. D. uvida

4. Stamens 12 or more; filaments and connectives glabrous or with a few scattered hairs... 5

5 Inflorescence with 1-3 flowers; hairs on pedicel and corolla white to grey 1. D. calycantha

5. Inflorescence with 3-10 flowers; hairs on pedicel and corolla pale to

orange-brown.2. D. hebecarpa

6 Secondary veins 4 or 5 pairs; corolla outside more or less glabrous;

pistillode glabrous.4. D. rugosula

6. Secondary veins 9-15 pairs; corolla outside pubescent; pistillode

pubescent.7

7 Twig hairs pale brown; secondary veins abaxially prominently raised;

corolla tube 2-3 mm long; corolla lobed from one half to two thirds its

length; pistillode 1-1.5 mm long.22. D. mabacea

7. Twig hairs white or grey; secondary veins abaxially flush or indistinct;

corolla tube 1—1.5 mm long; corolla lobed to at least two thirds its

length; pistillode 2.5-3.5 mm long.20. D. australis

8 Filaments with long dense hairs, 0.5-1 times length of anthers.9

8. Filament hairs c. 0.1 mm long, much shorter than anthers.10

9 Lamina discolorous, usually light grey-green below; apex acute,

sometimes acuminate or obtuse; filaments 0.5-1.5 mm long; anthers

linear to lanceolate, 2-3 mm long.21. D. pentamera

9. Lamina concolorous or slightly discolorous; apex mostly acuminate;

filaments 2.5-2.75 mm long; anthers ovate, 1-1.25 mm long.19. D. granitica

10 Lamina hairs on underside mostly erect, curved or sometimes tortuous,

more dense along the veins; midvein pubescent adaxially; anthers 4-5

mm long.7. *D. kaki

10. Lamina either glabrous or the hairs when present sparse, appressed and

straight, mostly confined to the main veins abaxially, midvein glabrous

adaxially; anthers less than 3 mm long.11

11 Lamina glands (nectaries) on underside scattered along basal half of

lamina, less than 0.5 mm across; each inflorescence with mostly 10-30

flowers.6. D. fasciculosa

11. Lamina glands on underside l(-2) on each side of midvein

confined to extreme base of lamina, more than 1.5 mm across;

inflorescence with less than 10 flowers.5. D. maritima

12 Rheophytic tree, lamina 0.8-1.7 cm wide; 4-5 times longer than wide;

obtuse at apex; restricted to Roaring Meg Ck catchment, NE Qld .... 18. D. rheophila

12. Not rheophytic; lamina length frequently less than 4 times width, apex various.13

158

Austrobaileya 9 ( 2 ): 155-197 ( 2014 )

13 Lamina when dry with prominent reticulate venation, nearly as prominent

as secondary veins.14

13. Lamina when dry with relatively obscure reticulate venation compared to

the secondary veins.15

14. Twigs usually with relatively persistent short erect hairs (sometimes

not evident in southern populations) and scattered longer appressed

submedifixed hairs.8. D. compacta

14 Twigs glabrescent, without erect hairs, only sparse appressed submedifixed

hairs.9. D. areolifolia

15 Stamens (2—)3-5.16

15. Stamens mostly 6-9.18

16 Midvein raised above.16. D. peninsularis

16. Midvein not raised above.17

17 Lamina mostly 1-3 cm wide; secondary veins 15-30 pairs; petiole 2-3

mm long, strigose; inflorescence mostly axillary; corolla tube 4-4.5

mm long.15. D. yandina

17. Lamina mostly 3-5.5 cm wide; secondary veins 7-15 pairs; petiole 3-5

mm long; inflorescence cauline, ramal and axillary; corolla tube 1.5-2

mm long.17. D. pluviatilis

18 Corolla tube 8-11 mm long; stamens 7-8.7 mm long; filaments 3.5-4 mm

long, anthers 4-4.5 mm long.13. D. laurina

18. Corolla tube to 4 mm long; stamens to 3.5 mm long, filaments to 2 mm

long, anthers to 2.5 mm long.19

19 Youngest twigs with more or less persistent minute erect hairs as well as

caducous appressed hairs .20

19. Youngest twigs glabrous or with caducous appressed hairs, minute erect

hairs absent.21

20 Lamina elliptic, oblanceolate or lanceolate, 3.5-10 cm long, acute or

acuminate at apex; margins sometimes undulate, inflorescence bracts

suborbicular, 0.25-0.3 mm long; pistillode 1-2.5 mm long.14. D. hemicycloides

20. Lamina obovate, rarely elliptic, rarely more than 3.5 cm long, obtuse or

rounded at apex; margins often recurved, inflorescence bracts obovate,

1.5-2 mm long; pistillode 0.5-1 mm long. 10. D. humilis

21 Secondary veins 5-8 pairs. Inflorescence frequently with more than 7

flowers; inside of calyx with appressed hairs.11. D. geminata

21. Secondary veins 10-15 pairs. Inflorescence with up to 7 flowers; inside of

calyx glabrous.12. D. littorea

Key to Australian Diospyros species using fruit and female flowers

1 Staminodes present. Flowers 4 or 5 (or 6)-merous. Ovary (3-)4 or 8-locular

with 1 or 2 ovules per locule, a total of (6) or 8 ovules; fruiting calyx

4-6-lobed.2

1. Staminodes absent. Flowers 3-merous (rarely 4 or 5-merous); ovary

(2-)3(-6)-locular, with (1) or 2 ovules per locule, a total of 4 or 6 ovules;

fruiting calyx 3-lobed (rarely 4 or 5-lobed).12

Jessup, Diospyros in Australia 159

2 Lamina glabrous on underside before full expansion.3

2. Lamina puberulous to pubescent on underside, glabrescent at full expansion.6

3 Flowers 3-10 on each axis, the calyx lobes 2-2.5 mm long; corolla

lobes 2.5-3 mm long; corolla tube 2-2.5 mm long; fruiting calyx tube

glabrous inside; margin of calyx tube not elevated above base of lobes,

fruit several in each axil or sometimes solitary.6. D. fasciculosa

3. Flowers solitary; calyx lobes 4.5-11 mm long; corolla lobes 9-10.5 mm

long; corolla tube 4-6.5 mm long; fruiting calyx tube pubescent inside;

margin of calyx tube more or less elevated above base of lobes, fruit

solitary.4

4 Corolla lobes less than 5 mm long; ovary and style glabrous or with few

hairs; fruit 15-18 mm diameter, glabrous.1. D. calycantha

4. Corolla lobes more than 5 mm long. Ovary and style densely

pilose or pubescent; fruit more than 20 mm diameter, pubescent.6

5 Corolla lobes 5.5-6.5 mm long, fruit 30-40 mm diameter, pubescent with

erect pale reddish brown hairs.2. D. hebecarpa

5. Corolla lobes 9-10.5 mm long, fruit 22-25 mm diameter, appressed

pubescent with white or pale brown hairs.3. D. uvida

6 Fruiting calyx lobes and tube cupular; the calyx appressed to base of fruit

throughout; fruit 9-15 mm wide.7

6. Fruiting calyx tube cupular and appressed to base of fruit, the lobes

spreading or reflexed, or whole calyx flat or reflexed; fruit

15-35(-90) mm wide.9

7 Lamina hairs, on abaxial surface minute, less than 0.25 mm long; pistillode

c. 2 mm long; staminodes glabrous; ovary 2.5-3 mm long.20. D. australis

7. Lamina hairs, on abaxial surface more than 0.25 mm long; pistillode

0.75-1.5 mm long; staminodes with long dense hairs at apex; ovary

1.5-2 mm long.8

8 Lamina concolorous, dull green or drying brown, apex acuminate; calyx

lobes on fruit indistinct or broadly deflate; 0-2 mm long.19. D. granitica

8. Lamina discolorous, light green beneath, apex acute or obtuse; calyx

lobes on fruit distinctly deflate, 2-4 mm long.21. D. pentamera

9 Fruiting calyx lobes 2-6 mm long.10

9. Fruiting calyx lobes 6-20 mm long.11

10 Twig sericeous; lamina tip acuminate or acute; lamina glands sparse;

secondary veins indistinct or slightly depressed above; calyx lobes on

fruit oblong.22. D. mabacea

10. Twig glabrous or strigose; lamina tip obtuse or rounded; lamina glands

1-3 near base; secondary veins raised above; calyx lobes on fruit deflate

or indistinct.5. D. maritima

11 Petiole 2-6 mm long; flower 5-6 mm long; style 0.4-0.6 mm long; calyx

tube on fruit up to 3 mm long, fruit 17-25 mm long.4. D. rugosula

11. Petiole 10-25 mm long; flower 15-25 mm long; style 3-4 mm long; calyx

tube on fruit 7-10 mm long, fruit 30-40 mm long.7. *D. kaki

160

Austrobaileya 9 ( 2 ): 155-197 ( 2014 )

12 Corolla tube 7-8 mm long, lobes 4-5 mm long; fruiting calyx becoming

indurated (thickened and woody), densely appressed sericeous or

pubescent outside.13. D. laurina

12. Corolla tube up to 4.5 mm long, lobes up to 3 mm long; fruiting calyx thin,

not indurated, sparsely appressed puberulous, glabrescent outside .13

13 Flowers usually in pairs or threes, only a few may be solitary.14

13. Flowers mostly solitary in each axil.16

14 Fruiting calyx lobed for more than half its length, more or less flat, not

cupular.17. D. pluviatilis

14. Fruiting calyx lobes much shorter than the cupular tube.15

15 Ovary glabrous, fruiting calyx lobes spreading or reflexed.6. D. fasciculosa

15. Ovary sericeous, fruiting calyx lobes appressed to the fruit, not spreading

or reflexed.11. D. geminata

16 Rheophytic tree, lamina 0.8-1.7 cm wide; 4-5 times longer than wide;

obtuse at apex; restricted to Roaring Meg Creek catchment, NE Qld . . 18. D. rheophila

16. Not rheophytic, lamina length frequently less than 4 times width; more

widespread distribution .17

17 Lamina when dry with prominent reticulate venation on both surfaces,

nearly as prominent as secondar veins.18

17. Lamina when dry with relatively obscure reticulate venation compared to

the secondary veins.19

18 Twigs usually with relatively persistent short erect hairs (sometimes

not evident in southern populations) and scattered longer

appressed submedifixed hairs.8. D. compacta

18. Twigs glabrescent, without erect hairs, only appressed submedifixed hairs

that soon fall.9. D. areolifolia

19 Midvein raised on adaxial surface of lamina .16. D. peninsularis

19. Midvein not raised on adaxial surface of lamina.20

20 Ovary glabrescent or glabrous except for a few hairs around style.21

20. Ovary sericeous .22

21 Young stems and petioles with sparse appressed submedifixed hairs, erect

hairs absent; peduncle of fruit 2-3 mm long.12. D. littorea

21. Young stems and petiole with dense short erect hairs and scattered

appressed hairs; peduncle of fruit 0.5-1 mm long.10. D. humilis

22 Twig indumentum glabrescent; secondary veins 5-12 pairs; flowers

3-5 mm long; corolla tube 2.5-3 mm long peduncle 3-7.5 mm long 14. D. hemicycloides

22. Twig indumentum persistent; secondary veins 15-30 pairs; flowers 5-6

mm long; corolla tube 3-7 mm long, peduncle c. 1 mm long.15. D. yandina

1. Diospyros calycantha O. Schwarz, Repert.

Spec. Nov. Regni Veg. 24: 93 (1927). Type

citation: [Northern Territory.] “Port Darwin,

Jervais road, dry jungle, F.A.K. Bleeser 156,

160, 161, 638 ” (syn: Bf). Type: Northern

Territory. 3.2 km S of U.D.P. Falls, 2 October

1986, M.J. Clark 698 (neo [here designated]:

BRI); isoneo: DNA, n.v).

Tree to 15 m, sometimes andromonoecious.

Twigs with basifixed appressed hairs, that

soon fall, and minute, more persistent dense

or sparse erect hairs. Leaves: petiole 2-6 mm

161

Jessup, Diospyros in Australia

long; lamina elliptic, 4.5-15 cm long, (1—)2.5—

6.5 cm wide; base cuneate; glands mostly 3-7

scattered on each side of midvein on basal

half of lamina, rarely absent; margins flat;

apex obtuse or acute, glabrous above and

below; secondary veins mostly 9-14 pairs,

not prominent, tertiary veins inconspicuous.

Male inflorescence shortly branched, to 2

mm long, mostly with 3-6 flowers; pedicels

to 1.5 mm long, grey pubescent; calyx

cupular, 2.5-27 mm long, shortly 4-lobed,

lightly appressed pubescent inside. Corolla

tube 6-8 mm long, lobes 4(-5), ovate or

oblong, recurved, 3-4 mm long, with a few

scattered grey hairs outside, glabrous inside.

Stamens 14-16, unequal, 1.5-5.5 mm long,

glabrous, filaments connate at base mostly

in pairs and adnate to base of corolla tube;

anthers linear, 1.2-1.6 mm long, connective

produced; pistillode subglobose, hirsute

at apex. Female flowers mostly solitary,

peduncle 3-4 mm long; bracts and bracteoles

persistent; calyx tube 3.5-4 mm long, lobes 4,

rhomboid, depressed ovate or semiorbicular,

mucronulate or not, spreading, 5-8 mm long,

7-8 mm wide, glabrous. Corolla tube 6-6.5

mm long, tubular but narrowed towards base,

lobes 4, ovate, recurved, 3.5-4.5 mm long,

apex acute. Staminodes mostly 6 or 7. Ovary

ovoid, with some appressed hairs at base,

nearly glabrous towards style, 8-locular with

1 ovule per locule, style c. 3 mm long, stigmas

shortly bilobed. Fruiting calyx tube cupular,

appressed to base of fruit, lobes spreading

or recurved, oblong, 6-8 mm long, glabrous;

tube 3-4 mm long with an elevated rim,

inside pubescent. Fruit depressed globose,

or globose, 15-18 mm diameter, glabrous,

orange to red, 5-7-seeded; seeds 10-14 mm

long, 3-8 mm wide. Fig. 1A-F.

Additional selected specimens examined : Western

Australia. 6 km S of Mining Camp in Crusher Vine

Thicket, Mitchell Plateau, N Kimberley, Jan 1989,

Kenneally KFK10845 & Hyland (DNA, PERTH).

Northern Territory. Maxwell Creek, Melville Island,

Jan 1990, Russell-Smith 8169 & Lucas (BRI); Hanguana

jungle, Melville Island, Jan 1988, Russell-Smith 4601

(DNA); Cobourg Peninsula, Wurgurlu Bay, Oct 1987,

Russell-Smith 3661 & Lucas (BRI); Ginger Palmer’s

camp, Gunn Point, Oct 1990, Russell-Smith 8345 &

Brock (BRI, MEL); Daly River, Mar 1989, Brock 725

& Russell-Smith (BRI); East Alligator River, Aug 1973,

Must 1147 (BRI, CANB, NSW); Lightning Dreaming,

Arnhem Land, Feb 1984, Dunlop 6594 & Wightman

(BRI, CANB, NSW); Nourlangie Ranger Station, May

1980, Craven 5668 (BRI, CANB, L, MEL); Head of

Koolpin Creek, West Arnhem Land, May 1978, Webb

& Tracey 12926 (BRI, CANB). Queensland. Cook

District: Batavia Downs, 3.1 km east of the western

boundary fence on the Mission River track, Oct 1989,

Neldner 2812 & Clarkson (BRI); Magnificent Creek,

10.8 km E of Rutland Plains, May 1992, Clarkson 9444

& Neldner (BRI); Shelfa Crossing, Mitchell River, Alice

Mitchell Rivers NP, 27 km NE of Kowanyama, Aug

1994, Myles HUM1078 et al. (BRI); Mungkan Kandju

NP, 2.5 km SW of Jabaroo Outstation, Coen River,

Nov 2008, Forster P1F34605 & McDonald (BRI); 13

km N of junction of Archer and Coen Rivers, Jun 1993,

Neldner 4073 (BRI); Archer Bend NP, 120 km WNW of

Coen, Jun 1994, Fell DGF4373 & Buck (BRI); Archer

River, Wenlock - Coen Road, Jul 1930, Brass 19727

(BRI); Archer River, just upstream from the Peninsula

Development Road, Apr 1991, Clarkson 8965 & Neldner

(BRI).

Distribution and habitat : Diospyros

calycantha occurs in north Queensland (Cape

York Peninsula and the Torres Strait Islands),

in the Northern Territory, mostly north of

Katherine, and in the Kimberley, northern

Western Australia (Map 1). It is found mostly

in semi-deciduous notophyll vine forest and

gallery forest. The species has also been

recorded from Papua New Guinea.

Phenology : Flowers have been recorded from

January to September, and fruit from March to

October.

Notes: The syntypes for Diospyros calycantha

in the Berlin herbarium were destroyed

during World War 2. A neotype has therefore

been selected.

2. Diospyros hebecarpa A.Cunn. ex Benth.,

FI. Austral. 4: 286 (1868). Type: Queensland.

[Cook District:] Cape York, s.dat., W. Hill

127 (syn: K); Endeavour River, June 1819, A.

Cunningham 308 (syn: K; isosyn: BM); East

Coast [Australia], in 1820, A. Cunningham

s.n. (syn: BM).

Illustration: Cooper & Cooper (2004: 155).

Tree to 20 m. Twigs with a few sparse appressed

hairs, glabrescent, soon glabrous. Leaves:

petiole 2-7 mm long, glabrous; lamina broadly

ovate, oblong, or elliptic, 5-13 cm long, 2.5-5.5

cm wide; base shortly attenuate; glands few

or several, then not confined to basal half of

lamina; margins flat; apex obtuse or scarcely

162

Austrobaileya 9(2): 155-197 (2014)

Fig. 1. A-F: Diospyros calycantha. A. male flower buds *3. B. male flower x3. C. dissected male petals showing

stamens x4. D. female flower x3. E. dissected female flower showing ovary x4. F. fruit x2. G: D. hebecarpa. G. fruit

xl. Fl-K: D. uvida. H. female flower x2.1. dissected female corolla x4. J. stamens from male flower x8. K. fruit xl. A

from Russell-Smith 8169 & Lucas (BRI); B & C from Neldner 4073 (BRI); D & E from Russell-Smith 4601 (DNA); F

from Clarkson 9444 & Neldner (BRI); G from Hyland //2//(BRI); H & I from Jessup GJM379 et al. (BRI); J from Jago

7032 (BRI); K from Cooper WWC1895 (BRI).

163

Jessup, Diospyros in Australia

acuminate; slightly discolorous and glabrescent

to glabrous below; secondary veins 6-10 pairs,

slightly raised above and below, basal pair

of secondary veins sometimes longer and

more acute than more distal veins, reticulate

veins above and below slightly raised. Male

inflorescence axes 3-11 mm long, with 3-10

flowers; pedicel hairs golden-brown. Calyx

tube 3-3.5 mm long shallowly 4-lobed, lobes

up to 0.5 mm long, acute, inside puberulous.

Corolla tube 6-7 mm long, lobes 4, ovate to

oblong; 2.5-3.5 mm long; outside appressed

puberulous in middle of lobe with pale golden-

brown hairs. Stamens 12-18,3.5-6.5 mm long,

glabrous or with a few hairs on the connective;

filaments single or connate in pairs or threes

and adnate to base of corolla tube, 1.5-4 mm

long, glabrous; anthers linear, 2-3 mm long;

pistillode 1-1.5 mm long, pubescent. Female

flowers solitary, peduncle 4-6 mm long.

Calyx tube 3.5-5 mm long; outside sparsely

puberulous, inside sericeous, 4(-5)-lobed;

lobes ovate, depressed ovate or elliptic-oblong,

7.5-8.5 mm long; 3-6 mm wide, spreading,

glabrescent, apex acuminate, acute or obtuse.

Corolla tube 5-6 mm long, glabrous inside,

lobes 4, 5.5-6.5 mm long, apex acute, outside

puberulous in middle of lobes. Staminodes 7

or 8. Ovary 3-4 mm long, densely pilose with

erect hairs, 8-locular, ovules 1 per locule;

style 2-3 mm long, pilose. Fruiting calyx tube

cupular and appressed to base of fruit, 8-9 mm

long, inside pubescent, lobes oblong, depressed

ovate or broadly obovate, rounded or apiculate,

reflexed, 7-10 mm long, 9-13 mm wide,

margins recurved, glabrescent or glabrous.

Fruit solitary, globose or depressed globose,

30-45 mm diameter, yellow, orange or red,

pubescent with pale erect hairs; seeds 13-15

mm long. Fig. 1G.

Additional selected specimens examined : Queensland.

Cook District: Dauan Island, Mt Cornwallis, Oct 1981,

Clarkson 3916 (BRI); Somerset, Cape York Peninsula,

May 1962, Webb & Tracey 6110 (BRI); 15 km from

the main Bamaga to Jardine River Road on the track

east to Ussher Point, Sep 1985, Clarkson 6230 (BRI);

Restoration Island, Aug 1965, Gittins 1060 (BRI); NPR

8, Parish of Weymouth, Oct 1981, Hyland 11211 (BRI),

Jan 1982, Hyland 11561 (BRI); Rocky River on eastern

foothills of Mcllwraith Range, Oct 1969, Webb & Tracey

9469 (BRI); Hope Vale Mission Reserve, Aug 1978,

Kants 1940 (BRI, L); Portion 5, Parish of Cook, 15° 32’S,

145° 13’E, Dec 1988, Hyland25647RFK (BRI); SFR310,

Goldsborough LA, Jul 1980, Hyland 10541 (BRI), Oct

1998, Hyland 16088 (BRI); SFR 933, Trinity, Little Pine

LA, Dec 1989, Hyland25724RFK (BRI). North Kennedy

District: Just S of Dittmer township, upper Kelsey

Creek, WSW of Proserpine, Aug 1993, McDonald5568 &

Bean (BRI); Airlie Beach, Sep 1992, Batianoff9209302

(BRI); Conway NP, Mt Rooper walking track, Nov 1985,

Warrian CW943 (BRI). South Kennedy District: R.60

Ossa, Cape Hillsborough, May 1975, Hyland 4273RFK

(BRI); Hidden Valley Road, Cape Hillsborough, 10 km

SE of Seaforth, Apr 1988, Thompson 138 (BRI); Skull

Knob, St Helens Beach, 10 km NE of Calen, Jul 1994,

Batianoff 94073 & Dillewaard (BRI); NW section of

Mt Beatrice, Eungella NP, s. dat., Pearson SP474 (BRI);

Dolphin Heads, Mackay, Sep 1994, Batianoff 94092 &

Saltman (BRI).

Distribution and habitat : Diospyros

hebecarpa occurs in northern Queensland,

from Torres Strait to Rollingstone, NW of

Townsville, and from N of Proserpine to

Mackay (Map 2). It also occurs in Papua New

Guinea and many parts of Malesia. It is found

in several types of vineforest.

Phenology : Flowers have been recorded from

December to April, and fruit from June to

December.

3. Diospyros uvida Jessup sp. nov.; resembling

D. hebecarpa , but differing in having male

flowers with 9 stamens and these with short

appressed hairs on the free part of the filaments

(12-18 stamens and glabrous filaments in D.

hebecarpa)., and female flowers with corolla

lobes 9-10.5 mm long (5.5-6.5 mm long in

D. hebecarpa ), and fruit developing to 22-

25 mm diameter and appressed pubescent

(30-45 mm diameter and pubescent with

erect hairs in D. hebecarpa) and with fruiting

calyx lobes spreading and 12-14 mm long

(reflexed and 7-10 mm long in D. hebecarpa).

Typus: Queensland. Cook District: Opposite

Burton’s dairy farm, end of Towalla Road, SE

ofMalanda, 29 October 2003 ,A. FordAF4188

& J. Holmes (holo: BRI).

Diospyros sp. Swipers LA (B.Hyland

1984RFK); Queensland Herbarium database;

Hyland et al. (2003).

Diospyros sp. (Swipers Flat); Cooper &

Cooper (2004: 156).

Tree to 10 m, often flowering as a shrub. Twigs

soon glabrous. Leaves: petiole 5-8 mm long,

glabrous, lamina elliptic or oblanceolate, 6-12

164

mm long, 2.5-6.5 cm wide; base cuneate;

glands few or if several, then not confined to

basal half of lamina; margins flat; apex shortly

and bluntly acuminate, glabrous both sides

before full expansion; secondary veins mostly

5-9 pairs, slightly raised above and below,

basal pair of secondary veins sometimes

longer and more acute than more distal veins,

reticulate veins above and below slightly

raised. Male inflorescence axes 3-10 mm long,

branched or not, few or multiple axes crowded

in a leaf axil, with up to 6 flowers maturing

together; pedicels with scattered pale brown

hairs; calyx cupular to obconical, glabrescent,

tube 2.5 mm long, inside sparsely puberulous,

lobes 4, obtuse, 0.5-0.7 mm long. Corolla tube

7-7.5 mm long, lobes 4, oblong, 4.3-47 mm

long, acute or obtuse, a few appressed hairs

present in middle of lobe on outside, otherwise

glabrous. Stamens 9 (2+2+3+2), 3.5-5 mm

long, free part of filaments and base of anthers

and connectives with short appressed hairs,

basal part of filaments connate for about 1

mm, glabrous; anthers narrowly oblong or

narrowly triangular, acute, connective not

visible at the apex; pistillode c. 0.8 mm long,

pubescent. Female flowers solitary, peduncle

2-4 mm long. Calyx tube 3.5-4 mm long,

appressed puberulous outside and inside,

4-lobed, lobes broadly ovate or obovate,

acuminate, 9-11 mm long, 6.5-8 mm wide,

glabrescent or glabrous both surfaces. Corolla

tube 4-5 mm long, glabrous inside, appressed

puberulous towards lobes outside, lobes 4,

narrowly ovate, 9-10.5 mm long, recurved at

anthesis, apex acute, outside puberulous in

middle of lobes. Staminodes 7 or 8. Ovary 3-4

mm long, densely appressed pubescent with

white or pale brown hairs, 8-locular, ovules

1 per locule, style 2-2.5 mm long, pubescent.

Fruiting calyx tube cupular and appressed to

base of fruit, 5-8 mm long, inside pubescent,

lobes broadly ovate, acuminate, spreading, 12-

14 mm long, 10-15 mm wide, glabrescent or

glabrous. Fruit solitary, subglobose, apiculate

at style remnant, 22-25 mm diameter, sparsely

appressed pubescent with white or pale brown

hairs, glabrescent, reddish black; seeds 13-14

mm long. Fig. 1H-K.

Austrobaileya 9(2): 155-197 (2014)

Additional selected specimens examined : Queensland.

Cook District: Eastern slope of Mt Sorrow, Feb 1993,

Weiblen 207 (BRI); Clacherty Road, c. 4.5 km NNE of

Julatten, Oct 2007, Jago 7032 (BRI); Foothills, Thornton

Peak, Sep 1937, Brass & White 301 (BRI); Baileys

Creek, N of Daintree River, in 1962, Webb & Tracey

6496 (BRI); Daintree River, Dec 1929, Kajewski 1449

(BRI); Mossman River Gorge, Feb 1932, Brass 2131

(BRI); Intake, Mossman, Sep 1948, Smith 3970 (BRI);

Rex Range, c. 2.8 km from Mossman - Julatten Road, c.

9 km NE of Julatten, Dec 1988, Jessup 876 et al. (BRI);

Rex Range, Mar 1991, Sankowsky 1252 & Sankowsky

(BRI); Formerly TR55, c. 300 m along snig track, near

Whyanbeel, Sep 2000, Ford AF2436 (BRI); Creek

behind Karnak, tributary of Whyanbeel Creek, Daintree

NP, Nov 1996, Jago 4160 (BRI); SFR 310, Swipers

FA, Oct 1968, Hyland 1984RFK (BRI) & Hyland 1997

(BRI); Boonjee FA near Bartle Frere track, 0.8 km S

Bobbin Bobbin Falls, 5.4 km NE Boonjee, Nov 1988,

Jessup GJM379 et al. (BRI); SFR 194, c. 600 m NE of

tower, Fonglands Gap, off McKell Road, Jul 2001, Ford

AF2906 (BRI); Towalla Road, Topaz, Aug 2004, Cooper

WWC1895 (BRI).

Distribution and habitat : Diospyros uvida

occurs in north east Queensland from Mt

Sorrow near Cape Tribulation to near Julatten

and also in Wooroonooran NP and adjacent

areas on the Atherton Tableland (Map 3)

mostly in mesophyll vine forest.

Phenology : Flowers have been recorded from

October to December, and fruit from January

to August.

Etymology : The species epithet is from Latin

uvidus meaning wet and humid and refers to

the Wet Tropics rainforest habitat where the

species grows.

4. Diospyros rugosula R.Br., Prodr.

526 (1810). Type: [Northern Territory]

Carpentaria, Groote Eylandt, 15 January

1803, R. Brown iter Austral. 2827 (holo: BM;

iso: K).

Diospyros rugulosa R.Br. ( orthogr. error) in

A.DC., Prodr. 8: 229 (1844).

Diospyros bundeyana Kosterm., Blumea 23:

454 (1977). Type: Northern Territory. W.

Arnhem land, Mt Bundey, 21 July 1971, MM

van Balgooy & N. Byrnes 1284 (holo: L).

Diospyros cordifolia auct. non Roxb.;

Bentham (1868: 286).

165

Jessup, Diospyros in Australia

Diospyros montana auct. non Roxb.;

Bakhuizen van den Brink (1936: 265-270).

Deciduous shrub or tree to 7 m. Twigs

pubescent with pale brown hairs, glabrescent.

Leaves: petiole 2-6.5 mm long; channelled

above, pubescent; lamina chartaceous, elliptic,

obovate, or ovate, 2-9(-14) cm long, 2-4(-

5.5) cm wide; base cuneate; glands sparse;

margins flat or undulate; apex obtuse, acute

or acuminate; pubescent with erect curved

hairs, glabrescent; secondary veins 4-5 pairs.

Male inflorescence axes up to 1 cm long;

bracteate, mostly 2 or 3 flowered; peduncle,

pedicel and outside of calyx pubescent with

erect hairs; calyx tube 1-1.5 mm long, lobes 4,

ovate, c. 2 mm long, inside glabrous. Corolla

contorted; lobes 4; oblong to ovate; 5-6 mm

long; tube 2-2.5 mm long, outside glabrous

except a few hairs on margin. Stamens

(14—)16, subequal, 4-5 mm long; filaments

adnate to corolla tube in pairs one above the

other or shortly connate, 0.3-0.8 mm long,

glabrous; anthers narrowly oblong-lanceolate,

acute, 2.5-3.2 mm long, with scattered short

hairs; pistillode subglobular, less than 1 mm,

glabrous. Female flowers solitary, peduncle

3.5-5 mm long, bearing 2 foliaceous bracts.

Calyx 4-lobed; 5.5-7 mm long, tube 2-2.5

mm long, lobes broadly ovate or suborbicular,

3.5-5 mm long; outside and inside pubescent.

Corolla urceolate, contorted, tube 2.5-3 mm

long, lobes 4, ovate, obtuse, 4.2-4.8 mm long,

ciliate on apical margin, otherwise glabrous.

Staminodes 4. Ovary pyramidal and shallowly

lobed, 3-3.5 mm long; glabrous; 8-locular;

with 1 ovule per locule; styles 4, connate only

at base, c. 1.5 mm long, bifurcating at the

stigmatic apex and with scattered erect hairs.

Fruiting calyx tube up to 3 mm long, lobes

reflexed, oblong or narrowly ovate, 6-9 mm

long. Fruit globose or subglobose; 17-25 mm

long, 25-30 mm wide, glabrous, 2-8-seeded;

seeds segment-shaped, rugulose, up to 15 mm

long. Fig. 2A-G.

Additional selected specimens examined : Western

Australia. Karrakatta Bay, E of Cape Leveque, Dampier

Peninsula, SW Kimberley Coast, Jun 1982, Kenneally

8534 (PERTH); Port Warrender, N. Kimberley, Jun

1974, Beard 7032 (PERTH); Walsh Point on W side of

Port Warrender off Admiralty Gulf, Jun 1985, Fryxell

et al. 4756 (CANB). Northern Territory. Port Darwin,

in 1883, Holtz 378 (MEL); Port Darwin, in 1884, Holtz

435 (MEL); Inverell Bay, 6 km W of Nhulunbuy, Nov

1989, Forster PIF5955 (BRI, MEL); Flinders Peninsula,

Dec 1987, Dunlop 7446 (BRI); Gunn Point, Nov 1989,

Forster PIF5923 & Russell-Smith (BRI); Mission Hole,

Elizabeth Downs Station, May 1984, Rankin 2906 (BRI);

10 km S Cannon Hill, Nov 1983, Russell-Smith 840

(BRI); Mataranka Falls, Roper River, Oct 1988, Russell-

Smith 6195 & Lucas (BRI); Wagait Road, 10 km N of

Finnis River Crossing, Oct 1984, Wightman 1679 (BRI);

East Alligator River, 1 mile [1.6 km] E of crossing, Jul

1972, Byrnes 2721 (CANB, K); Victoria River, Dec

1855, Mueller s.n. (MEL); 10 km SW of Timber Creek

township, Victoria River Region, May 1990, Menkhorst

1023 (MEL); 40 km NE from Numbulwar, Nov 1987,

Russell-Smith 4239 & Lucas (BRI); 16 Mile Caves

Reserve, Katherine, Oct 1977, Parker 1152 (CANB);

Towercast Area, 16 Mile Caves Reserve south of

Katherine, May 1978, Webb & Tracey 12300 (BRI); 8

km WSW of Borroloola, May 1987, Latz 10440 (BRI);

Carpentaria Hwy 15 km from Daly Waters, Apr 1993,

Egan 2061 (BRI); Fitzgerald Range 11 km N of Victoria

River Downs Station, Mar 1989, Russell-Smith 7640 &

Lucas (BRI).

Distribution and habitat: Diospyros rugosula

occurs from Cape Leveque to the Anjo

Peninsula, northern Western Australia and is

scattered throughout the Northern Territory,

north of around 16°20’S latitude (Map 3). It

is found in monsoon forest, semi-deciduous

microphyll or notophyll vine thicket, fringing

mangroves or in the vicinity of waterways. It

probably also occurs in Timor and Indonesia.

Phenology: Flowers have been recorded from

October to December, and fruit from October

to June.

Notes: This species was combined with

Diospyros cordifolia Roxb. by Bentham

(1868) and also with D. montana Roxb. by

Bakhuizen van den Brink (1936) but was

described as a distinct species by Kostermans

(1977) who compared it with D. cordifolia and

was apparently unaware of Brown’s name for

the same taxon (Brown 1810). In D. montana

and D. cordifolia the stamen filaments are

connate forming a common filament 1-2

mm long whereas in D. rugosula there is

practically no common filament, the stamens

attach to the corolla tube one above the other.

The pistillode is globular and slightly lobed

and lacks the apical style remnants found in D.

montana and D. cordifolia. It shares the same

number of staminodes (4) in the female flower

as D. montana whereas in D. cordifolia there

are 8-12 staminodes. The calyx is smaller and

166

Austrobaileya 9(2): 155-197 (2014)

Fig. 2. A-G: Diospyros rugosula. A. male flower x4. B. dissected male corolla showing stamens x4. C. dissected

female corolla showing staminodes x4. D. female flower x4. E. dissected male hypanthium showing pistillode x4. F.

dissected ovary x6. G. fruit xl.5. H-J: D. marUima. H. male flower bud x4; I. dissected male corolla showing stamens

x6; J. fruit xl.5. K-N: D. fasciculosa. K. male flower x8. L. dissected male flower showing stamens and pistillode x8.

M. female flower x6. N. infructescence xl. A, B, E from Russell-Smith 840 (BRI); C, D, F from Forster PIF5923 &

Russell-Smith (BRI); G from Webb & Tracey 12300 (BRI); H & I from Martin 90 (BRI); J from Weber 10006 (BRI); K

& L from Fell DGF3396 et al. (BRI); M from Brass 19969 (BRI); N from Fell DGF3995 et al. (BRI).

167

Jessup, Diospyros in Australia

less deeply lobed than in these two species and

the styles have scattered erect hairs (glabrous

in D. montana and D. cordifolia).

5. Diospyros maritima Blume, Bijdr. FI. Ned.

Ind. 669 (1826). Type citation: “ad littora

australie Javae insulae”. Type: possible type at

P, online image!

Cargillia laxa R.Br., Prodr. 526 (1810);

Diospyros laxa (R.Br.) F.M.Bailey, Syn.

Queensl. FI. 299 (1883), nom. illegit. non

Teijsm. & Binnend. (1855). Type: Northern

Territory. Gulf of Carpentaria opposite

Groote Eylandt, 10 February 1803, R. Brown

iter Austral. 2829 (holo: BM; iso: K).

Cargillia megalocarpa F.Muell., Fragm. 5:

163 (1866). Type: Northern Territory. Escape

Cliffs, Arnhem Land, in Nov/Dec 1865, John

McKinlay & Charles Hulls s.n. (syn: MEL

232960 & 232961).

Diospyros nitens W.Fitzg., The Western Mail

21 (1066) 10 (2 Jun. 1906), nom. inval., J. Proc.

Roy. Soc. Western Australia 3: 192 (1918).

Type: [Western Australia.] Devils Pass,

Napier Ranges, May 1905, W.V. Fitzgerald 614

(syn: PERTH 01598880); Synnott Range near

Sprigg River, August 1905, W.V. Fitzgerald

1321 (syn: PERTH 02890143 & 02890151;

isosyn: NSW926658, online image!).

Tree to 25 m. Twigs with appressed hairs,

glabrescent. Leaves: petiole 5-14 mm long;

lamina coriaceous, oblong to ovate, elliptic

or lanceolate, 5.5-30 cm long, 2-12 cm wide;

base cuneate or rounded; 1 or sometimes

2 glands on each side of midvein at base of

lamina; margins often recurved; apex obtuse

or rounded; glabrescent to nearly glabrous;

midvein depressed above, secondary veins

7-15 pairs. Male inflorescence axes up to

2 mm long with 3-8 flowers; calyx mostly

4-lobed, appressed pubescent, tube 2.5-3 mm

long, lobes 1-1.5 mm long, glands sometimes

present. Corolla appressed pubescent or

sericeous outside, glabrous inside, tube

5.5-6.5 mm long, lobes 4, 4.5-6 mm long.

Stamens 16—18(—20), 3-5 mm long, in two

whorls, filaments 1-1.5 mm long, connate

in pairs and adnate to the tube, pubescent

near anther; anthers linear, 1.8-2.6 mm long,

connectives usually with some appressed

hairs. Female flowers solitary, peduncle c. 2

mm long, calyx tube 3-5 mm long; lobes 4,

2-2.5 mm long, acute, appressed pubescent

outside, densely sericeous inside. Corolla tube

5-7 mm long, lobes 4, ovate-oblong, acute

or acuminate, 6-9 mm long, 3-5 mm wide,

staminodes 4-10, 3.5 mm long. Ovary ovoid,

ferruginous pubescent, 3-4 mm diameter,

8-locular, locules uniovulate, style 2-4 mm

long, puberulous, stigmas lobed. Fruiting

calyx patelliform, appressed to base of fruit.

Fruit depressed globose, 15-20 mm long, 20-

30 mm wide, glabrescent, black, 1-6-seeded;

seeds 8-10 mm long. Fig. 2H-J.

Additional selected specimens examined: Western

Australia. Chimney Rocks, (Emerian Point), NE of

Dampierland, Feb 1986, Martin 90 (BRI); loc. cit ., Mar

1986, Martin 91 (BRI); Surveyors Creek Road turnoff

between Mitchell Plateau mining camp and Point.

Warrender Road, May 1981, Tracey 15115 (BRI); Neville

Creek, off Calder River, Eastern Walcott Inlet, May

1983, Milewski 174 (BRI); Harding Range N of Eastern

Walcott Inlet, May 1983, Fell 40 (BRI); Hidden Island,

Buccaneer Archipelago, Jun 1982, Kenneally 8403

(BRI); Carson Escarpment; gully below Wonjarring

Falls, 4 km NNE of Face Point adjacent Drysdale River

NP boundary, Jun 1984, Forbes 2314 (BRI). Northern

Territory. Cape Hotham, 1.5 km S, Jun 1988, Russell-

Smith 5446 & Lucas (BRI); West Alligator Head, Jun

1988, Russell-Smith 5622 & Lucas (BRI); New Year

Island, Jul 1992, Leach 3132 (BRI); 5 km SSE of De

Courcy Head at Arafura Sea, May 1988, Munir 6052

(BRI); Arnhem Land, c.15 km SE from Cape Cockburn,

Jun 1988, Weber 10006 (BRI); Banjo Beach, Melville

Island, Jan 1966, Stocker GS35 (BRI); Warangaya,

Elcho Island, Sep 1987, Russell-Smith 3273 & Lucas

(BRI); Wessel Islands, Sep 1972, Latz 3259 (BRI);

Groote Eylandt, Bartalumba Bay, Jul 1972, Dunlop 2640

(CANB). Queensland. Cook District: Perry Island,

Feb 1991, Card PERU (BRI); Evans Bay, 26 km NE of

Bamaga, Feb 1994, Fell DGF3906 et al. (BRI); Mclvor

River Crossing, Cooktown - Starke Road, Tracey

14437 (BRI); Stephens Island, slope east of beach; east

of Cowley Beach, Sep 2010, Ford AF5773 & Bradford

(BRI).

Distribution and habitat : Diospyros

maritima occurs from near Derby, northern

Western Australia, to Groote Eylandt, Northern

Territory, and from northwest Cape York

Peninsula and Torres Strait to near Cowley

Beach, northeast Queensland (Map 4). It also

occurs in Indonesia, Timor and Papua New

Guinea. The species is found in coastal and

subcoastal vineforest, near mangroves and on

coral cays.

168

Phenology : Flowers have been recorded in

January and February, and fruit from March to

November.

6. Diospyros fasciculosa (F.Muell.) F.Muell,

Austral. Veg. 35 (1866); Maba fasciculosa

F.Muell., Fragm. 5: 163 (1866); Ebenus

fasciculosa (F.Muell.) Kuntze, Revis. Gen. PI.

2: 408 (1891). Type: [Queensland. Moreton

District:] Brisbane River, December 1856,

F. Mueller s.n. (lecto [here designated]: MEL

92392).

Maba laxiflora Benth., FI. Austral. 4: 290

(1868). Type: Queensland. [Port Curtis

District:] Rockhampton, s.dat., P. O’Shanesy

277 (holo: MEL 92396).

Maba ruminata Hiern, Trans. Cambridge

Philos. Soc. 12: 135 (1873). Type: New

Caledonia, in 1861, J.F. Deplanche 311 (holo:

P, online image!).

Diospyros sp. (Claudie River D.G.Fell

DGF3042); Jessup (1997, 2002, 2007, 2010).

Tree to 20 m. Twigs glabrescent or glabrous.

Leaves: petiole 6-10 mm long; ± flat

above, glabrous; lamina elliptic, ovate or

oblanceolate, 6-19 cm long, 2.5-7 cm wide;

base cuneate; glands less than 0.5 mm across,

sparsely scattered; margins recurved; tip

acute, acuminate or rounded; above and

below glabrous; midvein on upper surface

flush, raised, or depressed, secondary veins

6-10 pairs, above and below slightly raised,

tertiary veins above and below slightly raised,

flush or indistinct, above sparsely reticulate,

or reticulation obscure. Male inflorescence

axes usually 3-12 in axillary fascicles, each

axis branched with mostly 3-10 flowers,

appressed puberulous, glabrescent. Calyx

shallowly 3 or 4-lobed, 1.2-1.5 mm long,

glabrescent. Corolla campanulate, tube

2-2.5 mm long, glabrous, lobes 3 or 4,

semicircular or depressed obovate, 1.5-2 mm

long. Stamens 8-16(-20); 1.8-2.5 mm long;

filaments connate at base in pairs and adnate

to tube at base, 1-1.2 mm long, puberulous

with erect or antrorse hairs; anthers narrowly

ovate, 1.2-1.5 mm long, connective glabrous

or with a few hairs; pistillode 0.5-0.75 mm

long, glabrous. Female flowers 3-10 in a

Austrobaileya 9(2): 155-197 (2014)

fascicle, 5-10 mm long; pedicel 1.75-4 mm

long. Calyx 3 or 4-lobed; lobes 2-2.5 mm

long; tip rounded; tube 1.2-1.5 mm long;

outside puberulous; inside glabrous. Corolla

lobes 3 or 4, rounded, 2.5-3 mm long with

a median line of hairs; tube 2-2.5 mm long.

Staminodes 0-4. Ovary 2-3 mm long,

glabrous, 6-locular with 1 ovule per locule,

tapering to connate styles and 2 or 3 shortly

lobed stigmas. Fruiting calyx tube cupular

and appressed to base of fruit, 5-7 mm long,

lobes spreading or reflexed, 4-6 mm long.

Fruit broadly ellipsoid or subglobose; 13-15

mm long, 10-15 mm wide, apex and base

rounded, glabrous, 4-6-seeded; seeds 10-11

mm long. Fig. 2K-N.

Additional selected specimens examined : Queensland.

Cook District: Frangipani Beach Scrub, 0.7 km S of

Cape York, 26.4 km NE of Bamaga, Feb 1994, Fell

DGF3995 et al. (BRI); Stony Point N of Pascoe River,

Cape York Peninsula, Nov 1977, Webb & Tracey

13857 (BRI); Turrel Hill, 10 km WSW of Nesbit River

mouth, 51.6 km N of Silver Plains Homestead, Cape

York Peninsula, Aug 1993, Fell DGF3396 et al. (BRI);

Nesbit River, Sep 1973, Hyland 6831 (BRI); Bonanza

Creek, Peach River, Aug 1948, Brass 19969 (BRI); Mt

Stuckey Area NW Starcke Station, Sep 1974, Tracey

14292 (BRI). North Kennedy District: SE foothills of

Mt Dryander, Mar 1969, Smith s.n. (BRI [AQ410106]);

Adjacent to Impulse Creek, Apr 1999, McDonald 6683 &

Squires (BRI). Port Curtis District: Olsens Capricorn

Caverns, The Caves, Mar 1988, Vavryn 60 (BRI); Curtis

Island, Dec 1984, Gibson 685 (BRI); Essendean Bridge,

crossing of Baffle Creek, between Berajondo and Agnes

Water, Jan 2008, Forster PIF33230 et al. (BRI); SF 53,

Dan Dan Scrub, Dec 1987, Gibson 985 (BRI). Burnett

District: Coongara Rock, 11 km SE of Coalstoun Lakes,

SF 1344, Nov 2002, Forster PIF29110 (BRI). Wide Bay

District: Dundowran Beach, Apr 2003, Sankowsky 1994

& Sankowsky (BRI); Mt Walsh NP, Palm Valley, Coast

Range, Nov 2008, Forster PIF34508 (BRI); Tinana

Creek, 7 km ENE of Tiaro, Jan 2005, Forster P1F30537

et al. (BRI). Moreton District: Mt Eerwah, 4 km W

of Eumundi, Sep 1984, Sharpe 3578 (BRI, NSW); S of

Summit, Little Mt Brisbane, Nov 1984, Guymer 1903

& Dillewaard (BRI); Moggill FR, Nov 2005, Halford

Q8626 et al. (BRI). New South Wales. North Coast:

Woody Head, 3 miles [4.8 km] N of Iluka, Nov 1966,

McGillivray 2673 et al. (BRI, NSW).

Distribution and habitat : Diospyros

fasciculosa occurs on eastern Cape York

Peninsula north of Starcke NP, and from

Mt Dryander, central Queensland to Iluka,

northeast New South Wales (Map 4). It also

occurs in New Caledonia, Fiji and Eastern Java.

The species is commonly found in riverine,

169

Jessup, Diospyros in Australia

coastal and other types of lowland notophyll

vineforest.

Phenology: Flowers have been recorded from

August to December and fruit from November

to July.

Notes: Singh (2005) nominated the Mueller

collection at MEL as lectotype. However,

there is no evidence that he saw any of the

syntypes in MEL and as the Mueller collection

from the Brisbane River is mounted on two

sheets MEL 92392 and MEL 92394, with two

separate labels, I have chosen the former as

a subsequent lectotypification as allowed

under Art. 9.17 of the International Code of

Nomenclature (Melbourne Code).

7. *Diospyros kaki Thunberg, Nova Acta

Regiae Soc. Sci. Upsal. 3: 208 (1780). Type:

not designated.

Tree to 6 m, deciduous. Twigs pubescent or

nearly glabrous. Leaves: petiole 10-25 mm

long; lamina broadly elliptic to suborbicular

or ovate, 5-12 cm long, 2.5-10 cm wide;

base cuneate to rounded or shortly decurrent;

glands scattered along length of lamina,

sometimes few or absent; margins flat; apex

obtuse to shortly acuminate; pubescent below,

mostly on main veins, midvein pubescent

above; midvein depressed above, secondary

veins 5-7 pairs. Male inflorescence axes 5-8

mm long; in 3-5-flowered cymes. Calyx tube

1.2-1.5 mm long, lobes 4, 4.5-5 mm long,

3- 4 mm wide, pubescent outside and inside;

corolla tube urceolate, 6-7 mm long, lobes

4, ovate, recurved, 3-4 mm long, glabrous

or with a ciliate margin. Stamens 14-24,

filaments connate at base and adnate to

base of corolla tube, 1-1.5 mm long; anthers

oblong, 4-5 mm long, top of filaments and

connectives appressed pubescent; pistillode

disc-like, 0.7 mm diameter, glabrous. Female

flowers solitary, pedicel 10-12 mm long. Calyx

4- lobed, tube 4-5 mm long lobes 10-13 mm

long, acute, pubescent both sides, glabrescent;

corolla 10-15 mm long, 4-lobed, lobes ovate,

recurved, 4.5-5 mm long, Staminodes 8-16.

Ovary 3-5 mm long, sericeous or glabrous,

8-locular with 1 ovule per locule, style 4-fid,

3-4 mm long, stigmas 2-fid. Fruiting calyx

reflexed, 30-40 mm diameter. Fruit depressed

globose to broadly ovoid, 30-40 mm long,

20-85 mm wide, glabrous or nearly so, yellow

to orange; seeds 18-25 mm long.

Selected specimens examined : Queensland. Wide Bay

District: Glenbar Road, SF 57 St Mary’s, c. 18 km W of

Tiaro, Aug 2003, Watts s.n. (BRI [AQ762450]). Moreton

District: Yandina Creek, 2.5 km E of Valdora and c.

6 km NE of Yandina on Yandina Creek Road, Sharpe

4530 (BRI); SF E of Clear Mountain Road, Cashmere;

tributary of Four Mile Creek, Apr 2011, Phillips 2153 &

Phillips (BRI); Brisbane Valley Rail Trail: Mt Hallen -

Esk Section, Mar 2012, Phillips 2247 & Phillips (BRI);

Kobble Creek (most northerly branch), Dec 2008,

Phillips 1898 & Phillips (BRI); Paradise Road, Pallara,

c. 15 km S of Brisbane GPO, Nov 2003, Bean 21147

(BRI).

Distribution and habitat: Diospyros kaki

is native to China and possibly Japan and

is widely cultivated for its edible fruit. It is

naturalised in southeast Queensland, from near

Tiaro to south of Brisbane (Map 5). Clumps of

trees sometimes persist in abandoned gardens.

Phenology: Flowers have been recorded in

October and fruit in March.

8. Diospyros compacta (R.Br.) Kosterm.,

Blumea 23: 454 (1977); Maba compacta

R.Br., Prodr. 528 (1810); Ebenus compacta

(R.Br.) Kuntze, Revis. Gen. PI. 2: 408 (1891);

Diospyros ferrea var. compacta (R.Br.) Fosb.,

Brittonia 40: 61 (1988). Type: [Northern

Territory.] North Coast Island y2 [Pobassoo

Island], 18 February 1803, R. Brown iter

Austral. 2830 (holo: BM; iso: K).

Maba reticulata R.Br., Prodr. 528 (1810);

Ebenus reticulata (R.Br.) Kuntze, Revis. Gen.

PI. 2: 408 (1891), non Diospyros reticulata

Willd. (1805); Diospyros ferrea var. reticulata

(R.Br.) Balch., Bull. Jard. Bot. Buitenzorg ser.

3,15: 58, 64 (1937). Type: [Queensland. Cook

District:] Prince of Wales Island e [Good’s

Island], 2 November 1802, R. Brown iter

Austral. 2831 (holo: BM; iso: K).

Maba interstans F.Muell., Fragm. 5: 163

(1866). Type citation: “Ad sinum Rockingham

Bay, Dali achy'\ fide Bentham (1868: 291).

Type: not located.

Diospyros sp. (Mt White PI.Forster PIF14415);

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

Diospyros sp. (Mt White); Cooper & Cooper

(2004: 156).

170

Illustration : Cooper & Cooper (2004: 154).

Shrub or tree to 15 m. Twigs usually with

erect hairs c. 0.1 mm long and appressed

submedifixed hairs 0.3-0.4 mm long,

glabrescent. Leaves: petiole (1.7—) 3-5(-8)

mm long, flat above, with similar indumentum

to twigs. Lamina often coriaceous, obovate,

oblong, or elliptic, (1.5-) 3-8 (-12) cm long,

(0.8-) 2-4 (-8.5) cm wide; base cuneate;

glands 1-4 mostly in basal half, or absent

from some leaves; margins slightly to

strongly recurved; apex rounded, obtuse or

emarginate; glabrescent or soon glabrous;

midvein depressed above, secondary veins

4-7 pairs, raised above, sometimes difficult to

distinguish from higher order veins, reticulate

veins raised above on dried material. Male

inflorescence axes up to 3 mm long with

(1—)3(—5) flowers, puberulous. Male flower

pedicel to 0.5 mm long, puberulous with grey

or pale brown hairs. Flower 3.5-4 mm long.

Calyx tube 1.5-2.5 mm long, lobes 3(-4), 1-2

mm long, sparsely puberulous, glabrescent

outside, glabrous or with a few hairs inside,

tip rounded, or obtuse. Corolla tube 2.4-4

mm long, lobes 3, contorted, broadly ovate,

1-1.5 mm long, upper part of tube and lobes

appressed pubescent outside, the hairs shorter

where the lobes overlap in bud, glabrous

inside. Stamens (4-) 6-9, solitary or up to

3 singles alternating with up to 3 pairs, the

filaments of a pair connate only at base;

filaments not adnate to the corolla tube,

1.2-1.5 mm long, glabrous; anthers linear

to lanceolate, 1.5-2 mm long; pistillode up

to 1.5 mm long, pubescent. Female flowers

mostly solitary; peduncle 0.5-2 mm long,

calyx 3(-4)-lobed, tube to 1 mm long, lobes

obtuse or rounded, 1-1.5 mm long, sparsely

puberulous, glabrescent outside, sericeous

inside. Corolla tube 2-4 mm long, sericeous

outside, glabrous inside, lobes 3 or 4, broadly

ovate, rounded, 1.5-2 mm long, sericeous

outside, glabrous inside. Staminodes absent.

Ovary 2.5-2.8 mm long, sericeous, 3-locular;

ovules 2 per locule; style 0.3-0.5 mm long;

pubescent, stigmas shortly lobed. Fruiting

calyx recurved or spreading, lobes rounded or

indistinct. Fruit solitary, globose or depressed

globose, 9-14 mm long, 12-14 mm wide, soon

Austrobaileya 9(2): 155-197 (2014)

becoming glabrous, 1-4-seeded; seeds 5-6.5

mm long. Fig. 3A-G.

Additional selected specimens examined: Northern

Territory. Woolaning - Channel Point Road, N of

Daly River, May 1978, Webb & Tracey 12732 (BRI);

NE coast of Cape Van Diemen, Melville Island, May

1978, Webb & Tracey 12730 (BRI); Darwin River,

Nov 1974, Parker 560 (BRI); Darwin, Oct 1946, Blake

17325 (BRI); Point Stewart, Oct 1983, Russell-Smith

813 (BRI); Black Jungle, Nov 1982, Wightman 302 &

Dunlop (BRI, CANB, MEL, NSW); Black Jungle, Feb

1986, Wightman 2552 & Clark (BRI, CANB); Cobourg

Peninsula, Wurgurlu Bay, Oct 1987, Russell-Smith 3659

& Lucas (BRI); Gove, Cape Wirawawoi, Feb 1998,

Wightman 4110 (BRI); Mouth of Angurugu River,

Groote Eylandt, Mar 1988, Russell-Smith 5162 & Lucas

(BRI); Koolatong River, at junction with Maidjung

River, Oct 1996, Cowie 7370 (BRI). Queensland. Cook

District: Boydong Island, Dec 1987, Clarkson 7437

(BRI); Stoney Point, N of Pascoe River, Cape York

Peninsula, Nov 1977, Webb & Tracey 13859 (BRI); TR

14, Sep 1975, Hyland 3290RFK (BRI, NSW); Mt White,

Coen, Dec 1993, Forster P1F14415 (BRI); Youngmans

Crossing, Pascoe River, Nov 1977, Tracey 14170 (BRI);

Palfrey Island, near Lizard Island, Oct 1988, Batianoff

10305 (BRI); Cook’s Lookout, Lizard Island, Sep 1988,

Batianoff 10214 (BRI); Green Island, Coral Cay, E of

Cairns, Aug 1993, Jago 3016 (BRI). North Kennedy

District: Conway NP, Shute Harbour, about 35 km NE

of Proserpine, Nov 1985, Sharpe 4135 (BRI); Track

to Swamp Bay, Conway Range NP, about 25 km NE

of Proserpine, Nov 1985, Sharpe 4108 (BRI). South

Kennedy District: Turtle Bay, Carlisle Island, 35 km N

[of] Mackay, Sep 1986, Sharpe 4391 & Batianoff (BRI);

Mt Bassett, Mackay, Mar 1993, Batianoff 9303439 et al.

(BRI). Port Curtis District: Near homestead. Middle

Percy Island, Nov 1989, Batianoff11798 et al. (BRI).

Distribution and habitat: Diospyros

compacta occurs in the Northern Territory

from the Daly River to Groote Eylandt and

in Queensland from Torres Strait and Cape

York Peninsula to Middle Percy Island, central

coastal Queensland (Map 5). It is found in

semi-deciduous vine thicket and riparian

forest, frequently on coastal sands and near

mangroves.

Phenology: Flowers have been recorded from

August to November and fruit from November

to March.

Notes: Specimens with very narrow oblong

leaves with strongly recurved margins are

found in the deciduous vine thickets of Cape

York Peninsula and in coastal areas exposed

to wind shearing. These were placed in BRI

under Diospyros sp. (Mt White PI.Forster

PIF14415). This variation probably contributed

Jessup, Diospyros in Australia

171

Fig. 3. A-G: Diospyros compacta. A. male flower *4. B. dissected corolla, male flower x4. C. dissected stamens

and pistillode, male flower x8. D. female flower x4. E. dissected corolla, female flower x4. F. dissected ovary, female

flower x8. G. fruit x3. H-K: D. humilis. H. male flower x8.1. dissected male flower, showing corolla and stamens x8.

J. female flower x8. K. dissected ovary, female flower xi2. L-N: D. geminata. L. male flower x8. M. female flower

x8. N. fruit x3. A-C from Jago 3016 (BRI); D-F from Sharpe 4135 (BRI); G from Wightman 4110 (BRI); H, I from

Hyland 13692 (BRI); J, K from White 12487 (BRI); F from Jessup 919B (BRI); M from Jessup 919A (BRI); N from

Specht & Salt W531 (BRI).

172

to Brown (1810) describing Maba comp acta

based on specimens from Pobassoo Island,

NT (“North Coast Island Y2”) and Maba

reticulata based on specimens from Goods

Island, Qld (“Prince of Wales Islands e”).

Two taxa were also recognised by Bentham

(1868) and by Bailey (1900) but extensive

collecting since then has served only to blur

any previously perceived boundaries between

them. Differences in the calyx noted by

the above two authors can be attributed to

the degree of maturity of the fruit. Despite

searches by myself and others the type of

Maba interstans F.Muell. has not been found.

9. Diospyros areolifolia Kosterm., Blumea

23: 452 (1977). Type: Papua New Guinea.

Western District: Fly River area, Tarara,

Wassi Kussa River, 7 January 1936, L.J. Brass

8738 (holo: L n.v.; iso: A, online image!, BRI).

Small or large tree, once recorded with 90

cm gbh. Twigs with sparse appressed hairs,

soon glabrous. Leaves: petiole 3-4 mm long

and flat above, with scattered short erect hairs

above and appressed hairs below, glabrescent;

lamina obovate or oblong-elliptic, (2.5-)4-

6(-8) cm long 1.5-3 cm wide; base cuneate

or shortly attenuate; glands 1 or 2 on each

side of midvein on basal half of lamina below;

margins flat; apex obtuse; glabrous above and

below or with a few scattered appressed hairs

around the midvein below; midvein depressed

above, secondary veins 4-8 pairs, hardly

distinguishable from the prominent reticulate

venation on both surfaces. Male inflorescence

axes 2.5-3.5 mm long, pubescent, with

mostly erect pale brown hairs c. 0.1 mm

long, glabrescent. Male flowers not seen.

Female flowers not seen. Fruit peduncle 1

mm long; fruiting calyx shallowly cupular,

appressed to base of fruit throughout, 2-3

mm long, 5-7 mm diameter, glabrescent

outside, appressed pubescent inside, lobes

3, very short and rounded or depressed

triangular. Fruit globose, c. 1 mm diameter

(immature), appressed pubescent towards

style, glabrescent, persistent style c. 1.2 mm

long.

Additional specimen examined : Queensland. Cook

District: Wasp Creek area near Lockerbie, Cape York

Peninsula, Nov 1962, Hyland 2517 (BRI).

Austrobaileya 9(2): 155-197 (2014)

Distribution and habitat: The species

occurs in Queensland on northern Cape York

Peninsula (Map 5) and in Papua New Guinea.

Note: This species appears to be closely

related to Diospyros compacta. The type

bears only immature fruit and the Queensland

specimen appears to be male but has only

peduncles lacking flowers. Additional

collections are needed to confirm its status as

a distinct species.

10. Diospyros humilis (R.Br.) F.Muell.,

Austral. Veg. 35 (1867); Maba humilis R.Br.,

Prodr. 527 (1810); Ebenus humilis (R.Br.)

Kuntze, Revis. Gen. PI. 2: 408 (1891);

Diospyros ferrea var. humilis (R.Br.) Bakh.,

Bull. Jard. Bot. Buitenzorg ser. 3, 15: 57-62

(1937). Type: [Queensland. Port Curtis

District:] Upper Head, Broadsound, [12 & 13

September 1802], R. Brown iter Austral. 2834

(holo: BM).

Maba obovata R.Br., Prodr., 527 (1810);

Ebenus obovata (R.Br.) Kuntze, Revis. Gen.

PI. 2: 408 (1891); Maba ovata, orth. var.

F.Muell., Hooker’s J. Bot. Kew Gard. Misc.

8: 326 (1856). Type: [Northern Territory.]

Carpentaria Island a, b, c, 17-28 November

1802, R. Brown iter Austral. 2835 (holo: BM).

Illustration: Cooper & Cooper (2004: 155).

Shrub or tree to 10 m. Twigs with caducous

appressed hairs and persistent short erect

hairs. Leaves: petiole 0.9-5 mm long,

puberulous with erect hairs; lamina obovate,

rarely elliptic, 0.8-3(-5.4) cm long, 0.5-3 cm

wide; base cuneate; glands up to 10 on each

side of midvein, mostly in basal half; margins

flat; apex rounded or obtuse; glabrescent or

glabrous above, with scattered appressed

hairs below, midvein above puberulous

with erect hairs; midvein above depressed,

secondary veins 4-10 pairs, reticulate veins

not raised above. Male inflorescence axes

1-4 mm long; with (2-)3(-4) flowers. Male

flower pedicel 0-0.5 mm long, pubescent.

Calyx tube 1.5-2 mm long, lobes 3, 0.7-1.5

mm long, triangular, acute or obtuse, glabrous

or with few hairs on lobes inside, appressed

pubescent outside. Corolla tube 2-3 mm long;

lobes 3, ovate to triangular, 1.5-2 mm long,

upper part of tube and middle of lobes densely

173

Jessup, Diospyros in Australia

appressed sericeous outside, glabrous inside.

Stamens mostly 9, 3 singles and 3 pairs with

filaments connate at base, 2.5-3 mm long,

filaments not adnate to tube, 0.75-1.5 mm

long, glabrous; anthers linear, 1.4-2 mm

long; pistillode c. 0.5 mm long, pubescent.

Female flowers solitary, peduncle to 0.5 mm

long. Calyx tube 1.7-2.4 mm long, 3-lobed,

lobes triangular, obtuse, 1-1.5 mm long,

indumentum as in males. Corolla tube 1.75-3

mm long, lobes 3, broadly ovate, obtuse to

rounded, 1.5-2 mm long, indumentum as in

males. Staminodes absent. Ovary c. 1.8 mm

long, glabrescent except for some persisting

hairs around the style, 3-locular, ovules 2 per

locule, style up to 0.5 mm long, pubescent,

stigmas bifid. Fruiting calyx cupular,

appressed to base of fruit throughout, 4-5

mm long with rounded or obtuse lobes. Fruit

ellipsoid to nearly globular, 10-15 mm long,

7-9 mm wide, yellow or orange, glabrous or

nearly so, 1-3-seeded; seeds 7-9 mm long.

Fig. 3H-K.

Additional selected specimens examined : Western

Australia. Cape Leveque, Apr 1988, Dunlop 7826

(BRI); N of remote weather station, 29 km N of mining

camp, Mitchell Plateau, N. Kimberley, s. dat ., Kenneally

8554 (BRI). Northern Territory. Arnhem Land; mouth

of King River, Oct 1992, Cowie 3110 (BRI); Cutta

Cutta Caves, Oct 1988, Russell-Smith 6169 & Lucas

(BRI); Bremer Island, Jul 1992, Leach 2993 (BRI).

Queensland. Cook District: 4 km N of Edward River

Community, Dec 1979, Clarke 1091 (BRI); Brooklyn,

on track to Pennyweight Yards to junction with McLeod

River, Nov 2008, Jensen 1695 &. Stanton (BRI);

Brooklyn Nature Refuge, near Mt Carbine, Nov 2006,

McDonald KRM5936 et al. (BRI). Burke District:

Between Tully and Massacre Inlets, Carpentaria region,

Aug 1988, Hyland 13575 (BRI); Bowthorn Station,

34.6 km NNW of Bowthorn homestead beside Hedleys

Creek, Jul 2006, Thompson WES752 & Hogan (BRI);

Woodu (Muwera) between Nyuldorg and Thabugan

Point, Mornington Island, Sep 1981, Fosberg 62083

(BRI); Karumba, Aug 1943, Blake 15129 (BRI) &

15129A (BRI); 18 miles [28.8 km] NW of Normanton

township, Aug 1953, Perry 3959 (BRI, CANB). North

Kennedy District: 40 Mile Scrub, Nov 1988, Hyland

13692 (BRI) & 13693 (BRI). South Kennedy District:

3 km (direct) NW of haul road overpass, near Newlands

coal mine, WNW of Glenden, Jun 2009, Bean 29019

(BRI). Leichhardt District: Dry Creek Valley, eastern

side of Ka Ka Mundi section, Carnarvon NP, Aug 1990,

McDonald 4621 & Bean (BRI); Nathan Gorge, 23 km

SW of Cracow, Cabbagetree Creek, Apr 1991, Telford

11055 & Rudd (BRI). Port Curtis District: Marmor,

Nov 1943, White 12487 (BRI). Burnett District:

Goodnight Scrub NP, Aug 2009, Jessup 5260 & Bell

(BRI). Wide Bay District: Emu Creek Road, between

Dallarnil & Degilbo, Sep 1999, Forster PIF24915 (BRI).

Maranoa District: Western end of Eumina land, 2

km E of junction with the Orallo Road and adjacent to

Stanhope Downs, 44 km by road NW of Roma along the

Orallo Road, Oct 1996, Thomas s.n. (BRI [AQ651179]).

Darling Downs District. 14 km S of Bunya Mountains

on road to Dalby, Aug 1972, Sharpe 137 (BRI); McEwan

SF, Stoneleigh, off Young road, c. 5 km N of Pittsworth,

Jul 2011, Menkins ILM0536 (BRI).

Distribution and habitat : Diospyros humilis

occurs across northern Australia, from

the Dampier Peninsula, northern Western

Australia, Northern Territory, and from Torres

Strait to Pittsworth southwest of Toowoomba,

southeast Queensland (Map 6). The species is

also found in Timor and Papua New Guinea.

It occurs in rainforest, open woodland or

deciduous, semi-deciduous microphyll

or notophyll vine thicket, on sandy soils,

sandstone, limestone karst, or soil derived from

basalt.

Phenology : Flowers have been recorded from

October to January and fruit from February to

December.

Note : Bentham (1868) placed Maba obovata

in synonymy under M. humilis referring to the

fact that Brown’s specimens of the former had

male flowers while specimens of the latter bore

fruit. Numerous subsequent collections have

revealed some minor differences in leaf size

and fruit shape across the range of specimens

from eastern Queensland to Western Australia

but I am inclined to continue to recognise just

the one taxon.

11. Diospyros geminata (R.Br.) F.Muell.,

Austral. Veg. 35 (1867); Maba geminata

R.Br., Prodr. 527 (1810); Ebenus geminata

(R.Br.) Kuntze, Revis. Gen. PI. 2: 408 (1891);

Diospyros ferrea var. geminata (R.Br.) Bakh.,

Bull. Jard. Bot. Buitenzorg ser. 3, 15: 58,

64 (1937). Type: [Queensland. Port Curtis

District:] Keppel Bay Broad Sound, Thirsty

Sound, Broad Sound, [August - September

1802], R. Brown iter Austral. 2833 (syn: BM,

K).

Shrub or tree rarely more than 12 m high.

Twigs with very few appressed hairs,

glabrescent. Leaves: petiole 2.5-5 mm long,

glabrescent; lamina obovate, or elliptic, 3-7.5

cm long, 1-5 cm wide; base attenuate; glands

174

mostly 1-5 on each side of midvein in basal

half; margins sometimes recurved; apex

obtuse or rounded; glabrescent or glabrous;

midvein slightly depressed above, secondary

veins 5-8 pairs, reticulate veins scarcely

raised above. Male flowers fasciculate on

several condensed axes, with mostly 7-20

flowers in each axil. Flower sessile, calyx

tube 1.4-2 mm long, lobes 3(—4), 0.6-1 mm

long, appressed puberulous, glabrescent

outside, appressed pubescent inside. Corolla

tube 2.5-37 mm long, lobes mostly 3, oblong

or ovate, 2-2.5 mm long, inside glabrous,

outside appressed pubescent. Stamens 6-9,

2.5- 3.5 mm long, filaments not adnate to

tube, glabrous, 1.5-2 mm long; anthers linear,

1.5- 2 mm long; pistillode 0.5-1 mm long.

Female flowers fasciculate, sessile, 2-4 on a

very short common peduncle. Calyx 3-lobed,

tube 1.3-1.8 mm long; lobes 0.8-1.1 mm long,

obtuse or rounded, appressed puberulous,

sparsely outside, densely inside. Corolla

tube 2.5-3.5 mm long, lobes 3, 2-2.5 mm

long, obtuse, outside appressed pubescent.

Staminodes absent. Ovary 1.5-2 mm long,

sericeous, 3-locular; ovules 2 per locule;

style 0.4-0.6 mm long, pubescent. Fruiting

calyx cupular, lobes appressed to base of

fruit throughout. Fruit usually 1 or 2(-4)

on a common peduncle, broadly ellipsoid

or subglobose, 10-11 mm long, 6-7(-10)

mm wide, glabrescent with some appressed

hairs remaining around and below the style

remnant, 1-3-seeded; seeds 6-7 mm long.

Fig. 3L-N.

Additional selected specimens examined : Queensland.

Cook District: Pennefather River, Nov 2002, Kemp

TH6155 & Kutt (BRI); Lake Patricia, Weipa, Dec 1993,

Forster PIF14404 (BRI); 8.5 kmNW of Weipa Mission,

Jul 1974, Specht & Salt W531 (BRI). North Kennedy

District: Castle Hill, Townsville, Feb 1992, Bean 4064

(BRI); Tuckers Range, Jul 1993, Fensham 953 (BRI);

Woodwark Bay, Apr 1991, Forster PIF8243 & Bean

(BRI); Mingela Bluff, Jan 1992, Forster P1F9439 &

Bean (BRI); Mount Louisa West, 30 km SW of Home

Hill, Aug 1991, Bean 3523 (BRI). South Kennedy

District: N end of Ten Mile Beach, 5 km S of Midge

Point, Repulse Bay, Jun 1994, Batianoff 9406106 (BRI);

Keswick Island, Victor Bay Gully, Sep 1996, Batianoff

960965 et al. (BRI); Homestead Bay, St Bees Island,

Apr 1989, Batianoff 11263 (BRI). Leichhardt District:

Lake Elphinstone, s. dat ., Dietrich 1654 (BRI, MEL).

Port Curtis District: Yeppoon, Keppel Bay, Sep 1931,

White 8126 (BRI); Dan Dan Scrub SF, Boyne Range, Jan

2009, Forster P1F33359 et al. (BRI). Burnett District:

Austrobaileya 9(2): 155-197 (2014)

Goodnight Scrub, c. 65 km SW of Bundaberg, Jun 1957,

Smith 9836 (BRI, L). Wide Bay District: Stony Creek,

4 km E of Didcot, Oct 1990, Forster PIF7514 (BRI);

loc. cit. , Nov 1993, Telford 11970 (BRI); NW base of

Mt Boogooramunya, SF 648, Jan 1989, Forster PIF4903

(BRI); Near Imbil, Jun 1947, Smith & Webb 3132 (BRI,

L). Darling Downs District: Northern foothills of

Bunya Mountains, 51 km from Kingaroy on Dalby

Road, Nov 1984, Rodd 4223 (BRI). Moreton District:

Indooroopilly, near Brisbane River, 1.5 km upstream

from Walter Taylor Bridge, Jan 1993, Jessup 919A &

919B (BRI); Palen Creek SF near Mt Lindesay, Nov

1993, Grimshaw G97 (BRI).

Distribution and habitat : Diospyros

geminata occurs in eastern Queensland from

Cape York Peninsula to near Mt Lindesay

(Map 7), and is also in Papua New Guinea. It

is found in microphyll and notophyll vine forest

or vine thicket.

Phenology : Flowers have been recorded from

December to August and fruit from February

to December.

12. Diospyros littorea (R.Br.) Kosterm.,

Blumea 23: 461 (1977); Maba littorea R.Br.,

Prodr. 527 (1810); Maba buxifolia var. littorea

(R.Br.) Hiern, Nova Guinea (Botanique) 8

(1909); Diospyros ferrea var. littorea (R.Br.)

Bakh., Bull. Jard. Bot. Buitenzorg ser. 3, 15:

434 (1941). Type: [Northern Territory.] N

Coast Bay No 3, Point y2, 3 March 1803, R.

Brown iter Austral. 2832 (holo: BM).

Maba buxifolia auct. non (Rottb.) Pers.;

Hiern, Trans. Cambridge Philos. Soc. 12:

116-117 (1873).

Diospyros ferrea var. littorea f. laurina

(R.Br.) Bakh., Bull. Jard. Bot. Buitenzorg

ser. 3, 15: 434, 441 (1941) excluding type.

Misapplied name.

Tree to 10 m. Twigs with appressed medifixed

hairs, glabrescent. Leaves: petiole 2-6 mm

long, glabrescent; lamina narrowly obovate,

oblanceolate or elliptic, 2.5-11.5 cm long, 1.5-

5 cm wide; base cuneate; glands usually 1-3

on each side of midvein near base; margins

often undulate or recurved; apex rounded

or retuse; glabrous above, with scattered

appressed hairs below, glabrescent; midvein

above depressed, secondary veins 10-15

pairs. Male inflorescence axes branched, up

to 5 mm long, pubescent, with 3-7 flowers

in each axil, pedicel to 1.5 mm long. Calyx

175

Jessup, Diospyros in Australia

tube 0.75-1.25 mm long, lobes 3, 2-2.5 mm

long, obtuse or acute, calyx outside sparsely

appressed pubescent, inside glabrous. Corolla

tube 3.5-4 mm long, lobes 3, narrowly

ovate or oblong; 2-3 mm long, upper part of

tube and middle of lobes densely appressed

sericeous outside, glabrous inside. Stamens

6-9, 3-3.5 mm long; filaments not adnate

to tube; glabrous; 1-1.2 mm long; anthers

linear, 2-2.5 mm long; pistillode c. 0.5 mm

long, pubescent. Female flowers solitary on a

peduncle c. 2 mm long. Calyx tube 2.5-3 mm

long, 3-lobed, lobes broadly triangular, acute

or obtuse, 1.3-1.8 mm long, indumentum as in

males. Corolla tube 2.5-3.5 mm long, lobes 3,

broadly ovate, obtuse to rounded, 2.5-3 mm

long, indumentum as in males. Staminodes

absent. Ovary 1.7-2 mm long, glabrous,

3-locular, ovules 2 per locule, style 0.5-1

mm long, sparsely pubescent at base. Fruiting

calyx cupular, appressed to base of fruit

throughout, 4-4.5 mm long with depressed

triangular or rounded lobes. Fruit ellipsoid or

oblong, sometimes subglobose, 11-16.8 mm

long, 7-14 mm wide, apex rounded or flat,

glabrous, yellow or red, up to 6-seeded; seeds

to 10 mm long. Fig. 4A-E.

Additional selected specimens examined: Western

Australia. Camp on unnamed tributary of Prince

Regent River, arising 19 km SE of the mouth, Jun 1984,

Kenneally 8920 (PERTH). Northern Territory. North¬

east coast of Cape Van Diemen, Melville Island, May

1978, Webb & Tracey 12741 (BRI, CANB); Maxwell

Creek, Melville Island, Jan 1990, Russell-Smith 8168

& Lucas (BRI); Adelaide River, Daly River Road,

Nov 1982, Dunlop 6244 & Wightman (BRI); Adelaide

River, Feb 1979, Rankin 1752 (BRI, L); 30 km NE of

Numbulwar, Nov 1987, Russell-Smith 4231 & Lucas

(BRI); Black Jungle, Oct 1990, Brock 736 (DNA); NE

Arnhem Land, 2 km S of Lake Peter John, Feb 1988,

Russell-Smith 4984 & Lucas (BRI); Yirrkala, Gove

Peninsula, Oct 1983, Wightman 810 (BRI); Latram

Range, Gove Peninsula, Feb 1988, Russell-Smith 4984

& Lucas (BRI); Macarthur River mouth, 14 km SE of

Pelican Spit, Jan 1989, Russell-Smith 6785 & Lucas

(BRI, MEL). Queensland. Cook District: Saibai Island,

Oct 2007, Fell DGF8655 & Stanton (BRI); Olive River,

Nov 1978, Stirling 516 (BRI); Quintil Creek, Lockhart

River Aboriginal Reserve, Nov 1977, Tracey 14598

(BRI); Normanby River, Aug 1979, Duke AIMS674

(BRI); Batavia [Wenlock] River, in 1888, Milman s.n.

(BRI [AQ183600]); Claudie River, Aug 1979, Duke

AIM655 (BRI); Marrett River, Princess Charlotte Bay,

May 1979, El sol 666 & Stanley (BRI).

Distribution and habitat : Diospyros littorea

occurs in the Prince Regent River area, WA, in

the NT from the Daly River to the Macarthur

River and in Queensland from Torres Strait

to Princess Charlotte Bay (Map 7) in coastal

and estuarine vine forest and thickets fringing

mangroves. It also occurs from Thailand to

Papua New Guinea.

Phenology : Flowers have been recorded from

June to December, and fruit throughout the

year.

Note : Bakhuizen van den Brink (1941: 434,

440) included Australia in the distribution

of D. ferrea var. littorea f. lamponga

(Miq.) Bakh., without citing any Australian

specimens but also apparently suggesting this

form may not be distinct from typical Maba

littorea R.Br. [“ ? an Maba littorea typica

(i8ion

13. Diospyros laurina (R.Br.) Jessup comb,

nov.; Maba laurina R.Br., Prodr., 527 (1810);

Ebenus laurina (R.Br.) Kuntze, Revis. Gen.

PI. 2: 408 (1891). Type: [Queensland. South

Kennedy District:] Cumberland Islands, 17

October 1802, R. Brown iter Austral. 2836

(holo: BM; iso: E, K).

non Diospyros laurina Massalongo, nom.

nud ., see Notes below).

Diospyros cupulosa (F.Muell.) F.Muell.,

Austral. Veg. 35 (1867); Maba cupulosa

F.Muell., Fragm. 5: 164 (1866). Type:

[Queensland.] Rockingham Bay, s.dat., J.

Dallachy s.n. (lecto [here designated]: K

000792604, consisting of two mounted

branchlets with fruit attached and loose fruit

in the fragment packet with M. cupulosa

written thereon in Mueller’s hand).

Diospyros sericocarpa F.Muell., Austral.

Veg. 35 (1867); Maba sericocarpa F.Muell.,

Fragm. 5: 164 (1866). Type: [Queensland.]

Rockingham Bay, s.dat., J. Dallachy s.n.

(lecto [here designated]: K 000792603, top

left hand mounted specimen and loose fruits

in the bottom left fragment packet with M.

sericocarpa written thereon in Mueller’s

hand).

Maba rufa auct. wowLabill.; Hiern(1873: 114);

Bailey (1900: 963, 1913: 306); Bakhuizen van

176

den Brink (1936: 444-445); neque D. rufa

King & Gamble, J. Asiat. Soc. Bengal, Part 2,

Nat. Hist. 74 (1): 228 (1906).

Illustration : Cooper & Cooper (2004: 154).

Tree to 20 m, often flowering as a shrub.

Twigs with dense appressed basifixed reddish-

brown hairs, glabrescent. Leaves: petiole

3-8 mm long, glabrescent; lamina elliptic or

oblanceolate, 6-13 cm long, 2-5 cm wide; base

acute or obtusely cuneate; glands mostly 1-3

on each side of midvein in basal half; margins

flat; apex acute, shortly acuminate or rounded;

at first with dense appressed hairs both sides,

soon glabrescent; midvein above depressed,

secondary veins 6-10 pairs, often obscure.

Male inflorescence axes branched, 1.5-5 mm

long, appressed pubescent, usually with 3-12

flowers. Calyx tube 4.5-5.5 mm long, lobes

3, 0.5-1 mm long, obtuse, calyx appressed

pubescent outside, glabrous inside. Corolla

tube 8-11 mm long; lobes 3, broadly ovate,

2-3 mm long, corolla appressed pubescent

outside, glabrous inside. Stamens 9; 7-8.7 mm

long; filaments not adnate to tube, 3.5-4 mm

long, glabrous; anthers linear, 4-4.5 mm long;

pistillode 1-1.5 mm long, pubescent. Female

flowers l(-3) in each axil, pedicel to 2 mm

long; calyx tube 4.5-5 mm long, appressed

pubescent inside and outside, 3(-4)-lobed,

lobes 2.5-3 mm long, obtuse or rounded.

Corolla tube 7-8 mm long, lobes 3, narrowly

ovate, acute, 4-5 mm long, indumentum as in

males; staminodes absent; ovary 4-4.5 mm

long, coarsely sericeous, 3-locular, ovules 2

per locule, style 1-1.5 mm long, pubescent.

Fruiting calyx indurated, accrescent, 14-17

mm long, appressed to base of fruit with

lobes 3-6 mm long, slightly recurving, inside

sericeous, outside reddish-brown pubescent.

Fruit ellipsoid to subglobose, 15—18(—20)

mm long, 13-15 mm wide, pubescent, up to

6-seeded; seeds 6.5-7.5 mm long. Fig. 4F-L.

Additional selected specimens examined : Queensland.

Cook District: 5 km east along Captain Billy Landing

road, off Heathlands to Bamaga Road, Cape York

Peninsula, Jun 2008, Forster PIF33747 & McDonald

(BRI); NPR 8 Parish of Weymouth, Oct 1981, Hyland

11204 (BRI); TR 14 (Mcllwraith Range - Leo Creek

Road), Sep 1975, Hyland 8442 (BRI); Rocky River, Sep

1971, Hyland 5474 (BRI, L); Head of Temple Creek,

Cape Melville NP, May 1994, Fell DGF4340A (BRI);

Oliver Creek, Jul 1997, Hyland 15749 (BRI); Daintree

Austrobaileya 9(2): 155-197 (2014)

River, Dec 1929, Kajewski 1448 (BRI); Mossman River

Gorge, Feb 1932, Brass 2136 (BRI); Churchill Creek,

Churchill LA, SF 143, Jul 1995, Forster 17210 & Figg

(BRI); SFR 1073, Saddle Mt, near Kuranda, Nov 1966,

Berry NQNC14812 (BRI); Smithfield, Saddle LA, Jan

1982, Hyland 11493 (BRI); SF 607 Freshwater Creek, 7

km along road to Copperlode Dam, Oct 2001, Forster

PIF27564 et al. (BRI); Bridle Creek, c. 12 miles [19.2

km] SE of Mareeba, Nov 1973, Hartley 14149 & Hyland

(BRI); SFR 310, Gadgarra, Goldsboro ugh LA, Mar 1993,

Hyland 25826RFK (BRI); SFR 933, Trinity, Little Pine

LA, Oct 1988, Hyland 25587RFK (BRI); Lake Barrine,

Atherton Tableland, Jul 1929, Kajewski 1154 (BRI);

Malanda, Aug 1943, Blake 15175 (BRI); Palmerston NP,

west of Crawford Lookout, Jan 1993, Bean 5414 (BRI).

North Kennedy District: Alcock FR, rafting access

point no. 9, 5.2 km from Tully River camping area, Feb

2002, Ford AF3280 & Holmes (BRI); South Pinnacle,

c. 30 km SW of Townsville, Oct 1998, Gumming 17904

(BRI); Gregory Creek at junction of Gregory River,

Nov 1987, Perry 2 (BRI); Gregory Creek, c. 2 km NE

of Gregory and 15 km N of Proserpine, Nov 1985,

Sharpe & Perry s.n. (BRI [AQ423717]). South Kennedy

District: Scawfell Island NP, 50 km ENE of Mackay,

Nov 1986, Batianoff 6221 & Krieger (BRI); Cut Creek

at base of Eton Range, SF 652-658, Mackay, Oct 1986,

Ritchie 49 (BRI).

Distribution and habitat : Diospyros laurina

occurs in northern Queensland from northern

Cape York Peninsula to south of Mackay (Map

8) in mesophyll and notophyll vineforest and

wet sclerophyll forest. It also occurs in Papua

New Guinea.

Phenology : Flowers have been recorded from

September to November, and fruit from July to

February.

Notes : Massalongo (1859) included in a list

of fossil species the following: “ Diospyros

laurina Massal. Chiavon.” This name is a

nomen nudum leaving the way open for a

new combination to be made based on Maba

laurina R.Br. which predates M. cupulosa

F.Muell.

Mueller (1866: 164) described Maba

cupulosa and M. sericocarpa sequentially

and both from fruiting material collected by

J. Dallachy from Rockingham Bay. The only

Dallachy specimens with these two names

written in Mueller’s hand and with fruiting

material are the sheets at Kew cited above and

these are selected as the lectotypes of these two

names. The other branchlets mounted on K

000792603 have male flower buds and cannot

be considered part of the gathering described

Jessup, Diospyros in Australia

177

Fig. 4. A-E: Diospyros littorea. A. male flower x4. B. dissected stamens and pistillode, male flower x8. C. female

flower x4. D. dissected ovary, female flower x8. E. fruit x2. F-L: D. laurina. F. male flower x3. G. dissected corolla,

male flower x3. H. dissected stamens and pistillode, male flower x4.1, female flower x4. j. dissected corolla, female

flower x4. K. dissected ovary, female flower x4. L. fruit xl.5. M-Q: D. hemicycloides. M. male flower x4. N. dissected

stamens and pistillode, male flower x8. O. female flower x4. p. dissected ovary and stamens of possible hermaphrodite

flower x8. Q. fruit x4. A, B from Dunlop 6244 & Wightman (BRI); C, D from Brock 736 (DNA); E from Duke AIMS674

(BRI); F-H from Perry 2 (BRI); I-K from Sharpe & Perry s.n. (BRI [AQ423717]); L from Forster PIF27564 et al.

(BRI); M&N from Hyland 11362 (BRI); O & P from Hyland 25238RFK (BRI); Q from Forster PIF14336 (BRI).

178

by Mueller. The reference by Bailey (1900:

963) to a specimen of M. sericocarpa from

Eumundi is incorrect and most likely refers to

D. yandina.

14. Diospyros hemicycloides (F.Muell.

ex Benth.) Jessup comb, nov.; Maba

hemicycloides F.Muell. ex Benth., FI. Austral.

4: 290 (1868); Ebenus hemicyclodes (Benth.)

Kuntze, Revis. Gen. Pl. 2: 408 (1891). Type:

[Queensland.] Rockingham Bay, 13 August

1866, J. Dali achy s.n. (syn: MEF 92395;

isosyn: K 000792767); Mackay River [Tully

River], “Habit of Hemicyclia” in Mueller’s

hand, 11 August 1866, [. J. Dallachy s.n.] (syn:

MEF 233341A).

Diospyros sp. Q2 (Tinaroo Range J.G. Tracey

13936); Jessup (1994, 1997, 2002).

Diospyros sp. (Mt Fewis F.S. Smith 10107);

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

Diospyros sp. (Mt Fewis); Cooper & Cooper

(2004: 156).

Illustration : Cooper & Cooper (2004: 156),

as D. sp. (Mt Fewis).

Tree to 18 m, often flowering as a shrub.

Twigs with appressed submedifixed

pale brown hairs and minute erect hairs,

glabrescent. Feaves: petiole 1.5-5 mm long,

glabrescent; lamina elliptic, oblanceolate or

lanceolate, 3.5-10 cm long, 1-3 cm wide; base

cuneate or shortly attenuate; glands small

and sparse, up to 5 on each side of midvein

below; margins sometimes undulate, apex

shortly acuminate or acute; glabrous above,

glabrescent below; midvein on upper surface

depressed, secondary veins 7-11 pairs. Male

inflorescence axes 2-9 mm long, with 3(-4)

flowers, appressed pubescent, glabrescent.

Calyx tube 1.5-2 mm long, lobes 3, 1.5-2

mm long, obtuse or scarcely acute, calyx

glabrescent outside, glabrous inside. Corolla

tube 2.5-3.5 mm long, lobes 3(-4), ovate or

triangular with sides incurved, 3-4 mm long,

pubescent outside, glabrous inside. Stamens

7-9, 2-2.5 mm long, filaments not adnate

to tube, glabrous, 1.5-1.8 mm long; anthers

linear, 1-1.2 mm long; pistillode l(-2) mm

long, pubescent, sometimes with developed

styles. Female flowers usually solitary, rarely

Austrobaileya 9(2): 155-197 (2014)

a 3-flowered cyme, peduncle 3-7.5 mm long,

glabrescent, pedicel to 1 mm long. Calyx tube

2-2.5 mm long, lobes 3, connate in bud, 2-2.5

mm long, obtuse or acute, glabrescent outside,

glabrous inside. Corolla tube 2.5-3 mm long

lobes 3, triangular, acute with sides incurved,

2-2.5 mm long, pubescent outside, glabrous

inside. Staminodes absent. Ovary 2.2-2.5

mm long, sericeous, 3-locular, ovules 2 per

locule, style 0.7-1 mm long, with appressed

hairs at base, stigmas bifid. Fruiting calyx

appressed to base of fruit, 4 mm long, fruit

globose or broadly obovoid-ellipsoid, 13-16

mm long, 11-14 mm wide, with scattered

appressed hairs, glabrescent; seeds 8-10 mm

long. Fig. 4M-Q.

Additional selected specimens examined : Queensland.

Cook District: Cedar Bay NP, Mt Finnigan summit area,

Horans Creek, Oct 1999, Forster PIF25062 & Booth

(BRI); Kanawarra, Carbine LA, Nov 1987, Hyland25238

RFK (BRI); Riflemead, Carbine LA, Oct 1988, Hyland

25556 RFK (BRI); North Mary LA, [SFJR143 Mt. Lewis,

Sep 1973, Sanderson 335 (BRI); On Mt Lewis Road,

18 km from junction with Mareeba - Mossman Road,

Oct 1987, Foreman 1833 (BRI); SF 143, 15.5 km along

Mt Lewis Road, Oct 1999, Forster PIF25118 & Booth

(BRI); Mt Lewis Road, Aug 1957, Smith 10107 (BRI);

Daintree NP, Mt Sorrow track razorback, 4 km W of

Cape Tribulation, Dec 1997, Forster P1F22004 et al.

(BRI); Davies Creek, Aug 1954, Smith 5267 (BRI); SFR

185 Danbulla, Tinaroo, Aug 1981, Tracey 13936 (BRI);

SF 185 Danbulla, Tinaroo LA, 7.5 km past western

boundary of forestry grid, Dec 1993, Forster PIF14336

(BRI). Tinaroo Range, Jun 1970, Mori arty 310 (BRI);

Mt Bartle Frere, 1.8 km WSW of Bobbin Bobbin Falls,

4.4 km NNE of Boonjee, Nov 1988, Jessup GJM1131 et

al. (BRI); SF 194 Mt Baldy, 8 km from Rifle Range end,

Jun 1996, Forster PIF19223 et al. (BRI); Hugh Nelson

Range, Nov 1958, Smith 10504 (BRI); The Crater NP,

Atherton Plateau, Sep 1981, Fosberg 61674 (BRI). North

Kennedy District: Portion 205, Parish of Herberton,

Keoghs Scrub, Dec 1991, Gray 5400 (BRI); Tully Falls

NP, 7.6 km along Tully Falls Road from Charmillan

Creek bridge, Dec 2007, FordAF5196 & Metcalfe (BRI);

SFR 605 Luff LA, Dec 1981, Hyland 11362 (BRI); 27

km south along Culpa road, SF 605 Koombooloomba,

May 2002, Forster PIF28775 & Booth (BRI); Yuccabine

Creek SF 344, Kirrama LA, 28 km NW of Cardwell, Oct

1988, Jessup GJM2241 et al. (BRI), Jessup GJM2368

(BRI); Northern slope of Mt Thorn, SF 461 Meunga LA,

23.1 km W of Cardwell, Oct 1988, Jessup GJM 2271 et

al. (BRI); Coast Range [adjacent to Rockingham Bay],

Sep 1867, [U Dallachy ] (MEL 244448).

Distribution and habitat : Diospyros

hemicycloides is endemic to northeast

Queensland from Mt Finnigan to the Kirrama

Range (Map 9) and occurs in simple to

179

Jessup, Diospyros in Australia

complex notophyll vine forest on soils derived

from granite.

Phenology : Flowers and fruit have been

recorded from November and December.

Note : Flowers of Hyland 25238RFK were

observed to sometimes have an enlarged

ovary and stamens with slightly smaller

anthers. Further collecting is required to

determine if some plants of this species are

andromonoecious or monoecious.

15. Diospyros yandina Jessup sp. nov.;

resembling D. hemicycloides but differing in

the more numerous secondary veins (15-30

pairs compared to 5-12), in the shorter male

peduncle (c. 1 mm compared to 2-9 mm)

and the fewer stamens (3-5 compared to

7-9). It differs from D. cupulosa in the fewer

stamens (9 in D. cupulosa ) and in the smaller,

thinner, non-accrescent fruiting calyx.

Typus: Queensland. Wide Bay District: Kin

Kin Creek, beside track following boundary

of Cooloola National Park, August 1981, L.W.

Jessup 430 (holo: BRI).

Diospyros ellipticifolia f. australiensis Bakh.,

Bull. Jard. Bot. Buitenzorg ser. 3,15: 430, 436

(1941). Type: Queensland. Moreton District:

Blackall Range, November 1916, C.T. White

s.n. (holo: BRI [AQ183530]; iso: NSW).

Diospyros major var. ebenus f. australiensis

Bakh., Bull. Jard. Bot. Buitenzorg ser. 3,

15: 429 (1941). Type: Queensland. Moreton

District: Blackall Range, November 1916,

C.T. White s.n. (holo: BRI [AQ183530]; iso:

NSW).

Diospyros sp. (Blackall Range C.T.White

AQ183530); Jessup (2013).

Maba sericocarpa auct. non F.Muell.; Baker

(1899: 441-442), neque Bailey (1900: 963) pro

parte , “Rockingham Bay, Dali achy”.

Diospyros sp.l; Stanley & Ross (1986: 280,

2002 reprint: 280).

Shrub or rarely a tree to 10 m. Twigs with

appressed basifixed pale brown hairs and

minute erect hairs, glabrescent. Leaves:

petiole 2-3 mm long, glabrescent; lamina

elliptic, narrowly oblong-ovate or lanceolate,

3—8(—11) cm long, 1-3 cm wide; base cuneate

or rounded; glands small and sparse, up to

5 on each side of midvein below; margins

often undulate, slightly recurved; apex

acute or shortly acuminate; glabrous above,

sparsely appressed pubescent, glabrescent

below; midvein on upper surface depressed,

secondary veins mostly 9-18 pairs, sometimes

difficult to distinguish from higher order

veins. Male inflorescence peduncle c. 1

mm long, in the axils of reduced or normal

leaves on new season growth, pubescent,

with mostly 3 flowers, sessile or pedicels to

0.5 mm, pubescent. Calyx tube 1.5-2.5 mm

long, lobes 3, mostly connate at first then

1-2 mm long, mostly acute, calyx appressed

pubescent outside, glabrous inside. Corolla

tube 4-4.5 mm long, lobes 3, ovate, 2-3

mm long, appressed pubescent or coarsely

sericeous outside in the middle of the

lobes with shorter indumentum between,

glabrous inside. Stamens 3-5, 4-4.5 mm

long, filaments not adnate to tube, glabrous,

1.3-1.8 mm long; anthers linear, 1.8-2.2 mm

long; pistillode c. 1 mm long, pubescent.

Female flowers solitary, peduncle c. 1 mm

long, glabrescent, bract and bracteoles 2-2.5

mm long, pedicel 0-0.4 mm long. Calyx and

corolla similar to males. Staminodes absent.

Ovary 1.7-2 mm long, sericeous, 3-locular,

ovules 2 per locule, style 0.25-0.5 mm long,

appressed puberulous, stigmas bifid. Fruiting

calyx distinctly trilobed, 7-9 mm diameter,

only shortly appressed to base of fruit, the

lobes spreading or recurved, fruit globose

or depressed globose, 9-12 mm long, 11-15

mm wide, with scattered appressed hairs,

glabrescent, crimson, 1-5-seeded; seeds 6-7

mm long. Fig. 5A-C.

Additional selected specimens examined : Queensland:

Wide Bay District. Ramsays Scrub, Cooloola, May 1964,

Webb & Tracey 6348 (BRI); Mt Mothar, near Gympie,

Oct 1977, Webb & Tracey 11253 (BRI); Cooloola NP, road

to Harry’s Hut, Oct 1982, McDonald 3772 & Williams

(BRI); Road to Harry’s Hut, NE of Kin Kin, Sep 1993,

Bean 6467 (BRI); Lake Cootharaba, s.dat.. Keys 75

(BRI); Kin Kin, s.dat., Francis s.n. (BRI [AQ183541]);

Mt Cooroy, 4 km E of Cooroy, Apr 1986, Sharpe 4322

6 Guymer (BRI). Moreton District. Mt Eerwah, 4

km W of Eumundi, Dec 1984, Sharpe 3624 (BRI, L,

MEL, NSW); Wappa Falls, South Maroochy River, c. 6

km SW of Yandina, Oct 1986, Sharpe 4525 & Windolf

(BRI); Wappa Falls, Maroochy River, May 1959, Smith

10545 (BRI); Yandina, Mar 1891, Simmonds s.n. (BRI

[AQ183536]); 1 km S of Wappa Dam, SW of Yandina,

180

Austrobaileya 9(2): 155-197 (2014)

Fig. 5. A-C: Diospyros yandina. A. male flower x6. B. dissected corolla, stamens and pistillode *8. C. fruit <2.

D-J: Diospyros peninsularis. D. male flower x4. E. corolla, male flower x6. F. stamens and pistillode, male flower

x6. G. female flower x6. H. dissected ovary, female flower x6.1. dissected corolla, female flower x6. J. fruit x3. K-P:

Diospyrospluviatilis. K. male flower x6. L. dissected male flower showing stamens and pistillode x6. M. female flower

x6. N. dissected corolla, female flower x6. O. dissected ovary, female flower x6. P. fruit x2. A, B from Bird s.n. (BRI

[AQ396282]); C from Jessup 430 (BRI); D-F from Forster PIF6331(BRl ); G-I from Forster PIF5442 (BRI); J from

Williams 85195 (BRI); K, L from Gray 852 (BRI); M-0 from Jessup 708 (BRI); P from Tracey 15551 (BRI).

181

Jessup, Diospyros in Australia

Dec 1994, Bean 8131 (BRI); Nambour - roadside park

to N of town, Dec 1984, Bird s.n. (BRI [AQ396282]);

Tuckers Creek, roadside rest area on highway just N of

Nambour Dec 1980, Jessup 272 (BRI); northern tributary

of Petrie Creek, northern outskirts of Nambour, picnic

area, just N of major intersection of Bli Bli - Mapleton

Road, Dec 1986, Beesley 889 & Ollerenshaw (NSW);

Andrew Doig Park, Murray Grey Drive, Maroochy Shire

Council, 5 km WNW of Nambour, Dec 2006, Forster

PIF32363 et al (BRI); William Doig Park, Murray

Grey Drive, Maroochy Shire Council, 5 km WNW of

Nambour, Dec 2006, Forster PIF32389 et al. (BRI);

Murray Grey Drive, Dulong, west of Nambour, Dec

1993, Bean 7212 (BRI); Buderim Mt, Apr 1916, White

s.n. (BRI [AQ183529]). New South Wales. North Coast:

Tumbulgum, Feb 1897, Baeuerlen 1889 (BRI), Oct 1897,

Baeuerlen s.n. (BRI [AQ625613]), Dec 1897, Baeuerlen

s.n. (NSW), Jan 1898, Baeuerlen s.n. (NSW).

Distribution and habitat: Diospyrosyandina

occurs from the Great Sandy NP, Cooloola

section to Buderim Mt in southeast Queensland

and in the Tweed River valley, NSW (Map 9),

mostly in lowland notophyll vineforest.

Phenology: Flowers have been recorded from

December to February and fruit from October

to May.

Etymology: The species epithet

commemorates the township of Yandina,

Queensland where the earliest known

collection of the species was made by J.H.

Simmonds on 1 March 1891.

16. Diospyros peninsularis Jessup sp. nov.;

resembling D. yandina , D. hemicycloides and

D. laurina but differing in the midvein being

raised on the upper surface of the lamina (not

sunken), and differing from D. yandina in

the fewer secondary veins (7-11 compared

to 11-20), in the longer stamens (5-6 mm

compared to 4-4.5 mm), and the longer style

(c. 2 mm compared to less than 0.5 mm)

and from D. hemicycloides in the fewer and

longer stamens (up to 3 compared to 7-9,

and 5-6 mm compared to 2-2.5 mm). It also

differs from D. laurina in the much smaller

and thinner calyx in both sexes and in fruit.

Typus: Queensland. Cook District: N bank

of Pascoe River approx. 1 km inland from

river mouth, 21 November 1977, J.G. Tracey

14396 (holo: BRI, iso: distribuendi ).

Diospyros sp. (Kuranda L.J. Webb+ 7265A);

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

Diospyros sp. (Kuranda); Cooper & Cooper

(2004: 156).

Diospyros sp. Bamaga (B.P.Hyland

2517) at http://keys.trin.org.au: 8080/

key-server/data/0e0f0504-0103-430d-

8004-060d07080d04/media/Html/taxon/

Diospyros_sp._Bamaga_(B.P.Hyland_2517).

htm with respect to the description, images,

and all specimens except the reference

specimen. The reference specimen Hyland

2517 is Diospyros areolifolia.

Diospyros sp. (Bamaga); Cooper & Cooper

(2004: 156).

Shrub or tree to 10 m. Twigs with appressed or

inclined submedifixed pale brown coarse hairs

and minute erect hairs, glabrescent. Leaves:

petiole 2.5-4 mm long, glabrescent; lamina

elliptic, ovate or lanceolate, 3-10 cm long,

1-3 cm wide; base cuneate or rounded; glands

small and sparse; margins slightly recurved

and undulate; apex acute with a blunt tip; with

scattered to sparse appressed or inclined hairs

on both sides, glabrescent; midvein on upper

surface raised, secondary veins mostly 7-11

pairs. Male inflorescence axes c. 1 mm long,

1- several fasciculate in leaf axils, pubescent,

each usually terminating with 3 flowers,

sessile or on pedicels to 1 mm, pubescent.

Calyx tube 1-2.5 mm long, lobes 3, sometimes

4, connate at first then 1-2 mm long, acute,

calyx appressed pubescent outside, hairs

shorter and more sparse inside. Corolla tube

5-6 mm long, lobes mostly 3, ovate, 1.5-2

mm long, appressed pubescent or coarsely

sericeous outside, glabrous inside. Stamens

(2) or 3, 5-6 mm long, filaments not adnate to

tube, glabrous, 2.5-3 mm long; anthers linear,

2.5-3 mm long; pistillode 0.5-0.8 mm long,

pubescent. Female flowers solitary, peduncle

0.5-0.7 mm long, pubescent, bract and

bracteoles 0.8-1 mm long, pedicel up to 0.2

mm long. Calyx tube 1.3-1.5 mm long, lobes

3(-4), broadly ovate, obtuse, 1-1.3 mm long,

indumentum as in males. Corolla tube 4-4.5

mm long, lobes 3 or 4, navicular-ovate, 1.4-1.6

mm long, coarsely sericeous outside, glabrous

inside. Staminodes absent. Ovary 2 mm long,

coarsely sericeous, 2 or 3-locular, ovules 2 per

locule, styles ± completely connate, to 2 mm

long, stigmas 2 or 3. Fruiting calyx usually

182

flattened or recurved from base of fruit. Fruit

globose or depressed globose, 8-10 mm long,

9-12 mm wide, glabrescent, reddish brown or

orange, style remnant and surrounding hairs

sometimes persisting, 1-4-seeded; seeds 5-8

mm long. Fig. 5D-J.

Additional selected specimens examined: Queensland.

Cook District: Moa [Island] in Torres Strait, May 2003,

Wannan 2904 & Toh (BRI); Lockerbie (abandoned),

Lockerbie Scrub, 5 km from old homestead site, N, on Cape

York Road, Sep 1985, Williams 85195 (BRI); Lockerbie

Scrub, 3 km past Lockerbie Homestead, Feb 1990, Forster

P1F6331 (BRI); 47 km from the Cape York Road on the

track to Ussher Point, Jul 1992, Clarkson 9675 & Neldner

(BRI); Restoration Island, Apr 1995, Le Cussan 303

(BRI); Stoney Point, N of Pascoe River, Nov 1977, Tracey

14154 (BRI); Pascoe River mouth, north bank, 33.8 km

NNW of Lockhart River, Bromley Holding, North East

Cape York peninsula, Apr 1993, Fell DGF3154 & Butcher

(BRI); Between Iron Range and Portland Roads, Oct 1972,

Dockrill 557 (BRI, L); Iron Range, Oct 1983, Sankowsky

267 & Sankowsky (BRI); Head of Swamp Creek, Table

Range, 11.9 km S of Lockhart River community, Apr

1994, Fell DGF4295 & Daunt (BRI); Junie Creek, Oct

1972, Dockrill 574 (BRI); Alligator Creek, Oct 1972,

Dockrill 588 (BRI, L); Mcllwraith Range, c. 11 miles

[17.6 km] ENE of Coen, Oct 1962, Smith 11784 (BRI);

33.2 km E by road of Maloney’s Springs, 73.2 km E by

road of Moreton Telegraph Station, Jun 1989, Forster

PIF5442 (BRI); c. 6 km SSW of Cape Flattery township.

May 1990, Clarkson 8628 & Neldner (BRI); Abor. Res.

1, between Mclvor River & Cape Flattery, Nov 1972,

Hyland 6538 (BRI, L); Bridge Creek Holding (proposed

NP), upper Bridge Creek catchment; NW of Cooktown,

May 2010, Forster PIF36683 & Thomas (BRI); Carrol

Creek road crossing, NW of Hopevale mission, Dec 1984,

Jessup 647 (BRI, L).

Distribution and habitat : Diospyros

peninsularis occurs on Cape York Peninsula,

Queensland from near Cooktown to

Torres Strait (Map 9), in several forms of

semideciduous notophyll vine forest and

araucarian vine thicket.

Phenology : Flowers have been recorded

from February to July, and fruit from July to

December.

Etymology : The species epithet refers to Cape

York Peninsula where the majority of the

specimens have been collected.

Notes : The species described here is the

same taxon as the reference specimen cited

in the phrase name Diospyros sp. (Kuranda

L.J. Webb+ 7265A) but the locality details

(Smithfield-Black Mt road via Kuranda)

Austrobaileya 9(2): 155-197 (2014)

provided on the label of the reference

specimen are doubtful as no other specimens

of this species have been found south of

Carrol Creek near Cooktown.

17. Diospyros pluviatilis Jessup sp. nov.;

resembling D. yandina but differing in the

mostly larger leaves (3-5.5 cm wide compared

to 1-3 cm wide), the more prominent

secondary veins, more numerous flowers in

the inflorescence and the persistent remnants

of inflorescences on older branchlets. It

differs from D. hemicycloides in the mostly

more numerous flowers in each inflorescence

and in the fewer stamens (3 or 4 compared

to 7-9). Typus: Queensland. Cook District:

Souita Falls - Middle Brook Creek Road, SE

of Millaa Millaa, 11 November 1992, J.G.

Tracey 15551 (holo: BRI; iso: distribuendi ).

Diospyros sp. (Millaa Millaa L.W. Jessup

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

Diospyros sp. (Millaa Millaa); Cooper &

Cooper (2004: 156).

Illustration : Cooper & Cooper (2004: 156),

as D. sp. (Millaa Millaa).

Shrub to 3 m. Twigs with appressed sub-

basifixed hairs and short erect basifixed hairs,

glabrescent. Leaves: petiole 3-5 mm long,

glabrescent; lamina elliptic or lanceolate,

8-14 cm long, 3-5.5 cm wide; base cuneate

or rounded; glands small and sparsely

scattered on undersurface; margins flat; apex

shortly acuminate; adaxial surface glabrous,

abaxial surface glabrescent; midvein above

depressed, secondary veins 9-13 pairs

forming prominent loops well within the

lamina margin. Male inflorescences cauline,

fasciculate, forming clusters of 5-20 flowers;

peduncles with persistent overlapping bracts

and peduncles persistent on older branchlets.

Calyx tube 1-1.5 mm long, lobes (2-)3, 1-2

mm long, bluntly acute, calyx appressed

pubescent outside, glabrous inside. Corolla

tube 1.5-2 mm long, lobes 3, triangular,

1.5-2 mm long, appressed pubescent outside,

glabrous inside. Stamens 3 or 4, 2.5-3

mm long, filaments attached below base of

pistillode, not adnate to corolla tube, glabrous,

0.75-1 mm long; anthers linear, 1.7-2 mm

long; pistillode c. 0.6 mm long, pubescent.

183

Jessup, Diospyros in Australia

Female flowers 1-3 together, peduncles

1-1.5 mm long with persistent overlapping

bracts and the peduncles persistent on older

branchlets. Calyx tube 1.2-2.4 mm long,

lobes (2—)3, connate in bud, 2-2.5 mm long,

bluntly acute, calyx appressed pubescent

outside, glabrous inside. Corolla tube 2-2.5

mm long, lobes 3, triangular, 2.4-2.6 mm

long, appressed pubescent outside, glabrous

inside. Staminodes absent. Ovary 1.8-2 mm

long, sericeous, 3-locular, ovules 2 per locule,

style 0.5-0.75 mm long, with appressed hairs

at base, stigmas bifid. Fruiting calyx trilobed,

5-8 mm diameter, scarcely appressed to base

of fruit, the lobes spreading or recurved, fruit

globose or broadly ellipsoid, 10-15 mm long,

9-13 mm wide, with scattered appressed

hairs, glabrescent; seeds 8-10 mm long. Fig.

5K-P.

Additional selected specimens examined : Queensland.

Cook District: NPR 133, Daintree, above tributary of

Mackenzie Creek, WNW of Mt Hutchinson, Sep 2002,

Ford AF3601 & Holmes (BRI); S of junction of E & W

Mulgrave Rivers, SFR 310, Goldfield LA, 20.1 km SSE

of Little Mulgrave township, Nov 1988, Jessup GJM1663

et al. (BRI); NE slopes of Bartle Frere, in 1995, Hunter

JH2151 (BRI); TR1230, Boonjee LA, Nov 1977, Gray

772 (BRI, CNS); loc. cit ., Jan 1978 Gray 852 (BRI,

CNS); Mt Bartle Frere, 1.8 km WSW of Bobbin Bobbin

Falls, 4.4 km NNE Boonjee, Nov 1988, Jessup GJM1036

et al. (BRI); Stockwellia track, Wooroonooran NP, Dec

1998, Forster PIF24054 et al. (BRI); SFR 755 Gosschalk

LA, May 1976, Hyland 8795 (BRI, CNS); The Boulders,

North Babinda Creek, W of Babinda, Dec 1984, Jessup

708 (BRI); SFR 755, Barong LA, Francis Range, Dec

1984, Jessup 715 & 716 (BRI); 1.4 km SE of Cooroo Peak

at the head of Culla Creek 14 km NW of South Johnstone,

Oct 1988, Jessup GJM2551 et al. (BRI); Junction of

Duffer Creek & Johnstone River, Jul 1992, Tucker &

Sankowsky s.n. (BRI [AQ547363]); SF 756 Mt Father

Clancy, May 2000, Forster PIF25722 & Booth (BRI);

North Johnstone River, near Palmerston NP, Nov 1982,

Jessup 517 (BRI); Palmerston NP, North Johnstone River,

Dec 1984, Jessup 741 (BRI); Palmerston NP, Tchupalla

Falls track, Nov 1982, Jessup 515 (BRI); Kaaru LA, SW

corner 14.5 km SSE of Millaa Millaa, Oct 1988, Jessup

GJM2016 et al. (BRI); SFR 756 Jordon, Lower Downey

LA, Dec 1991, Gray 5388 (BRI, CNS), Gray 5389 (BRI,

CNS), Gray 5394 (BRI, CNS); Elinjia LA, 7.4 km NE of

Millaa Millaa, Oct 1988, Jessup GJM2130 et al. (BRI);

Gregory Falls, Lower Palmerston via Innisfail, in 1962,

Webb & Tracey 6680B (BRI); Mena Creek, Oct 1995,

Gleed s.n. (BRI [AQ585232]); Liverpool Creek W of

Silkwood, Dec 1984, Jessup 738 (BRI).

Distribution and habitat : Diospyros

pluviatilis is endemic to northeast Queensland,

occurring from near Mt Hutchinson north of the

Daintree River and in the area approximately

bounded by the upper reaches of the Mulgrave

River, Bellenden Ker township, Mena Creek,

Liverpool Creek and Millaa Millaa (Map 10),

in complex mesophyll or notophyll vineforest,

on alluvium or basalt soil.

Phenology : Flowers have been recorded from

October to February, and fruit from October,

November and May.

Etymology : The species epithet is a Latin

adjective meaning ‘relating to rain’ and refers

to the natural distribution of this species being

within areas of the Wet Tropics recording

some of the highest rainfall averages in

Australia.

18. Diospyros rheophila Jessup sp. nov.; it

differs from all other Australian species in

its rheophytic habit and leaf lamina length

4-6 times width. Typus: Queensland. Cook

District: Timber Reserve 165, Baird Logging

Area, 22 September 1980, B. Hyland 10623

(holo: BRI).

Diospyros sp. (Baird L.A. B.Hyland 9374);

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

Diospyros sp. (Baird LA); Cooper & Cooper

(2004: 156).

Shrub or tree to 5 m. Twigs with appressed

submedifixed hyaline or pale brown hairs,

glabrescent. Leaves: petiole 3-4 mm long,

flat above, glabrescent; lamina narrowly

oblanceolate or narrowly elliptic, 4.3-8 cm

long, 1.1-L4(-1.7) cm wide; base cuneate;

glands 2-6 on basal quarter of lamina below;

margins slightly recurved; apex rounded

or bluntly acute; glabrous above, sparsely

appressed pubescent with submedifixed hairs

below, glabrescent; midvein more or less flush

above, secondary veins 5-10 pairs, often

difficult to distinguish from higher order

veins. Male flowers not seen. Female flowers

solitary, peduncle 1-2 mm long, pubescent,

with caducous bracts. Calyx 3.5-4 mm long,

tube 1.5-2 mm long, lobes 3, connate at first

then 1.5-2 mm long, mostly obtuse, calyx

appressed pubescent, glabrescent outside,

appressed pubescent inside. Corolla tube

2-3 mm long, lobes 3, ovate, 2-2.5 mm long,

appressed pubescent or coarsely sericeous

184

outside in the middle of the lobes, glabrous

below and inside. Staminodes absent. Ovary

1.7-2 mm long, sericeous, 3-locular, ovules 2

per locule, style 0.75-1 mm long, pubescent

at base, stigmas bifid. Fruit not seen. Fig.

6A&B.

Additional selected specimens examined : Queensland.

Cook District: TR165, Alexandra LA, Jun 1977, Hyland

9374 (BRI); TR 106, Parish of Noah, Baird LA, Roaring

Meg Creek, Jul 1997, Hyland 26021RFK (BRI).

Distribution and habitat : Diospyros

rheophila is endemic to northeast Queensland

growing as a rheophyte amongst rocks and

boulders in notophyll vineforest along Roaring

Meg Creek, north of Daintree (Map 10).

Phenology : The species flowers in September.

Etymology : The specific epithet is from Greek

rheophilus , loving rivers, in reference to the

apparent preferred habitat of the species.

19. Diospyros granitica Jessup sp. nov.; allied

to D. pentamera but differing in the leaf lamina

not discolorous and papillate below and the

fruiting calyx fully appressed to the base of

the fruit with inconspicuous lobes compared

to the fruiting calyx with lobes 2-4 mm long

and often recurved in D. pentamera. Type:

Queensland. Cook District: State Forest

Reserve 143, Kanawarra, Carbine Logging

Area, 21 December 1988, B. Hyland 13792

(holo: BRI [2 sheets]).

Diospyros sp. (Mt. Spurgeon C.T. White

10677); Jessup (1994,1997,2002,2007,2010).

Diospyros sp. (Mt Spurgeon); Cooper &

Cooper (2004: 156).

Tree to 15 m. Twigs with appressed long and

short basifixed pale brown hairs, glabrescent.

Leaves: petiole 3-6 mm long, glabrescent;

lamina elliptic, 5-7(-9) cm long, 2-3.8 cm

wide; base shortly attenuate; glands mostly

3-8 on basal quarter of lamina below; margins

slightly recurved near base; apex acuminate;

sparsely appressed pubescent, glabrescent

both sides; midvein on upper surface

depressed, secondary veins 7-11 pairs,

sometimes indistinct, highest order veins ±

parallel. Male inflorescence axes solitary or

several in axillary fascicles, 1.5-4 mm long,

each with 3-5 flowers, appressed pubescent.

Austrobaileya 9(2): 155-197 (2014)

Calyx tube 1-1.5 mm long, lobes 4 or 5, 1-1.2

mm long, acute or obtuse; calyx appressed

pubescent outside and at base inside. Corolla

tube 1.5-1.7 mm long, lobes 4 or 5, broadly

ovate, 2.5-27 mm long, appressed sericeous

outside except margins, appressed puberulous

inside. Stamens mostly (15—)16(—17), 3.5-4

mm long, filaments connate at base in pairs

and adnate to corolla tube at base, 1.8-2 mm

long, pilose just below the anther, the longer

hairs as long as the anthers; anthers narrowly

ovate, acuminate, 1-1.3 mm long; pistillode

c. 1 mm long, puberulous towards apex.

Female flowers often 3 in each axil, each on

a pubescent peduncle 2-4 mm long, bracts

c. 1 mm long, caducous. Calyx tube 2.5 mm

long, lobes 4 or 5, 1 mm long, acute or obtuse;

calyx appressed sericeous outside and inside,

puberulous along lobe margins. Corolla tube

c. 1.5 mm long, glabrous outside and inside,

lobes 4 or 5,2-2.5 mm long. Staminodes 6-10,

like stamens with vestigial anthers, filaments

not paired. Ovary 2.5 mm long, sericeous,

3-5-locular, ovules 2 per locule, styles 3-5,

1.5-1.7 mm long, distally free and glabrous

just below the lobed stigmas, otherwise

appressed pubescent. Fruiting calyx cupular,

8-9 mm diameter, appressed to base of fruit,

lobes broadly triangular, indistinct or scarcely

visible; fruit obovoid, ellipsoid or subglobose,

10-18 mm long, 10-13 mm wide, glabrescent;

mostly 1-seeded; seeds 9-10 mm long. Fig.

6C-H.

Additional selected specimens examined : Queensland.

Cook District: Mt Spurgeon, Sep 1937, White 10677

(BRI); loc. cit., Nov 1985, Godwin C2923 (BRI); Near

Schillers Hut, Mt Spurgeon, Sep 1972, Webb & Tracey

11779 (BRI); 32.5 km along Mt Lewis Road from

Mossman - Mt Molloy Road, Dec 1989, Jessup GJD3366

etal. (BRI); Mt Lewis Road, S Mary LA, 16 kmNNW Mt

Molloy, Nov 1988, Jessup GJM1554 et al. (BRI), Jessup

GJM1497 etal. (BRI); SFR 143, Riflemead, Carbine LA,

Oct 1988, Hyland 13587 (BRI), Dec 1988, Hyland 13798

(BRI), Nov 1987, Hyland 25237RFK (BRI), Dec 1988,

Hyland 25628RFK & 25627RFK (BRI); SFR 143, Parish

of Riflemead, Carbine LA, Jul 1988, Gray 4896 (BRI);

SFR 143, Kanawarra, Carbine LA, Dec 1988, Hyland

13792 (BRI); Mt Lewis, Oct 1971, Webb & Tracey

10530 (BRI); Mt Bartle Frere, Oct 1891, Johnson s.n.

(MEL 233340); loc. cit., Jan 1891, Johnson s.n. (MEL

233339); Upper Russell River, Jan 1891, Johnson s.n.

(MEL233337); Mt Bartle Frere, in 1892, Johnson s.n.

(BRI [AQ519813]); Wooroonooran NP, East Mulgrave

River, Nov 2000, Forster P1F26441 et al. (BRI); E Bartle

Frere, Nov 1994, Hunter JH1840 (BRI).

Jessup, Diospyros in Australia

185

Fig. 6. A, B: Diospyros rheophila. A. female flower *6. B. dissected ovary, female flower x6. C-H: D. granitica. C.

male flower *6. D. dissected male corolla and stamens x8. E. female flower *4. F. dissected female corolla showing

staminodes x4. G. dissected ovary, female flower x6. H. fruit x2. I-N: D. australis. I. male flower in late bud x6.

J. dissected male flower showing stamens and pistillode x8. K: female flower x4. L. dissected corolla showing

staminodes, female flower x4. M. dissected ovary, female flower x4. N. fruit x2. A, B from Hyland 10623 (BRI); C &

D from Hyland 25628RFK (BRI); E-G from Hyland 13792 (BRI); H from Hyland 13587 (BRI); I from Volck s. n. (BRI

[AQ13088]); J from Hoy 162 (BRI); K from Stanley 78179 & Ross (BRI); L & M from Grimshaw G286 & Franks (BRI);

N from Gibson TOI491 (BRI).

186

Distribution and habitat : Diospyros

granitica is endemic to northeast Queensland

occurring between Mt Spurgeon and Mt Lewis

and also on the slopes of Mt Bartle Frere (Map

11). It is found in notophyll and microphyll

vine-fern forest on granite.

Phenology : Flowers have been recorded in

November and December and fruit from July

to December.

Etymology : The specific epithet refers to the

granite-derived soils in which this species

grows.

20. Diospyros australis (R.Br.) Hiern,

Trans. Cambridge Philos. Soc. 12: 30 (1873);

Cargillia australis R.Br., Prodr. 527 (1810).

Type: New South Wales. Port Jackson,

Hawkesbury, Hunter’s River, s.dat., R. Brown

iter Austral. 2828 (syn: BM, E, K).

Maba cargillia F.Muell., Fragm. 5: 162 (1866),

pro syn. nom. inval., nom. nud.

Diospyros cargillea F.Muell., Austral. Veg. 35

(1867) nom. inval., nom. nud.

Diospyros cargillia F.Muell. ex Hiern nom.

illeg., nom. alt:, Hiern, Trans. Cambridge

Philos. Soc. 12: 77, 155, 246, 290, 292 (1873).

Annona microcarpa Jacq., Fragm. Bot. 40,

t.44 (1800-1809); Diospyros microcarpa

(Jacq.) Giirke (1890), nom. illegit., non D.

microcarpa Span. (1836) etnonD. microcarpa

Siebold (1844).

Tree to 30 m. Twigs appressed pubescent

with basifixed hairs. Leaves: petiole 3-8 mm

long, channelled above, appressed pubescent;

lamina oblong, elliptic or narrowly ovate,

3-12 cm long, 1-3.5 cm wide; base cuneate;

glands 1—8(—16) on basal half of lamina

below; margins sometimes recurved; apex

often obtuse or rounded; glabrescent and

soon glabrous above, appressed pubescent

and minutely papillate below appearing

dull pale green; midvein on upper surface

shallowly depressed; secondary veins 7-11

pairs, indistinct below. Male inflorescence

axes solitary or several in axillary fascicles,

1.5-4 mm long, each with 3-5 flowers,

appressed pubescent. Calyx tube 2.5-3 mm

long, lobes 4, 0.5-0.7 mm long, obtuse; calyx

Austrobaileya 9(2): 155-197 (2014)

appressed pubescent outside, glabrous inside.

Corolla tube 1.5-1.6 mm long, lobes 4, ovate

or oblong, 4-5 mm long, appressed pubescent

outside, glabrous inside. Stamens 12-16, 3-4

mm long, filaments connate at base in pairs

and adnate to corolla tube at base, 1-2 mm

long, glabrous; anthers narrowly lanceolate,

2-2.5 mm long, glabrous; pistillode c. 2 mm

long, pubescent, with glabrous rudimentary

styles. Female flowers 1-3 in each axil, each

on a pubescent peduncle to 2.5 mm long,

bracts navicular, 2-3 mm long, caducous.

Calyx tube 2.5-3 mm long, lobes 4, 1-1.3

mm long, acute or obtuse; calyx appressed

pubescent outside and inside on lobes. Corolla

tube 1.5-1.7 mm long, glabrous outside and

inside, lobes 4-6, oblong, 3.5-4.5 mm long.

Staminodes 8-15, like stamens with sterile

anthers, filaments not paired. Ovary 2.5-3

mm long, sericeous, 4-locular, ovules 2 per

locule, style 1.5-1.7 mm long, appressed

pubescent but glabrous just below the lobed

stigmas. Fruiting calyx cupular, 10-12 mm

diameter, appressed to base of fruit, lobes

broadly triangular, c. 2 mm long; fruit ovoid

or subglobose, 11-14 mm long, 9-10 mm

wide, appressed pubescent; mostly with 1

seed; seeds 7-8 mm long. Fig. 6I-N.

Additional selected specimens examined : Queensland.

Cook District: Candlenut Scrub, SF 144 Mt Windsor

Tableland, Nov 1997, Forster PIF21883 et al. (BRI);

Danbulla, R185, Oct 1953, Volck s.n. (BRI [AQ13088]);

SFR 194, Sylvia LA, NE of Walsh Falls, Sep 1998, Ford

2104 (BRI). North Kennedy District: Mount Aberdeen

NP, 40 km inland from Bowen, Mar 1989, Fell DF1822

(BRI). South Kennedy District: Dairymple Heights and

vicinity, Sep-Nov 1947, Clemens s.n. BRI [AQ183475]).

Port Curtis District: 2.5 km SW of Raglan, R146,

Horrigan Creek, Mar 1989, Gibson T01491 (BRI); 20 km

from Agnes Water, S of Gladstone, Nov 1978, Stanley

78179 & Ross (BRI). Leichhardt District: Buckland

Creek, Carnarvon Range, s.dat., Jensen s.n. (BRI

[AQ183462]). Maranoa District: Carnarvon NP, Mount

Moffatt Section, gully to the north of Marlong Plain,

Nov 1996, Eddie 19 (BRI). Burnett District: Cania

Gorge NP, along track to Dripping Rock, Oct 1999,

Halford Q3858 (BRI); Mt Walsh NP, track to Coongara

Rock, 15 km SSW of Biggenden, Coast Range, Nov

2007, Forster PIF33148 (BRI); 6 km NNW of Coalstoun

Lakes, Apr 1991, Forster PIF7848 (BRI); Bunya

Mountains, between Noolers Lookout & Mt Kiangarow,

Dec 1954, Smith 6256 (BRI). Wide Bay District: 1 km

SW of Booyal, Childers, Nov 1987, Forster PIF3294

(BRI); Jimna Range Road, Kilcoy to Goomeri, north

of Jimna township, Dec 1993, Grimshaw G286 &

Franks (BRI). Darling Downs District: Mt Colliery

187

Jessup, Diospyros in Australia

to Gambubal Road, western fall of the Main Range,

NE of Killarney, Jan 2012, Forster PIF38565 & Leiper

(BRI). Moreton District: Riverside Drive near Ipswich

Mining Museum, Raymonds Hill near Coal Creek, Mar

1994, Grimshaw G514 (BRI); Black Rock Creek scrub,

13 km S of Boonah, Mar 1991, Forster PIF7815 (BRI).

New South Wales. North Coast: Nymboida River,

near Bibiranga Road, 11 km S of Nymboida, May 1994,

Bean 7726 (BRI); Beecroft Peninsula, 6.3 km NNW of

Point Perpendicular, S end of Long Beach, Dec 1988,

Winsbury 86 et al. (BRI); Bolaro Mountain Road, 9 km

from Kings Highway, Feb 1988, Richardson 153 et al.

(BRI).

Distribution and habitat : Diospyros

australis occurs in eastern Australia, from

Mt Windsor Tableland, northeast Queensland

to near Batemans Bay, NSW, also inland at

Carnarvon Gorge and vicinity in Queensland

(Map 12), mostly in microphyll to notophyll

vine forest on a variety of soil types.

Phenology : Flowers have been recorded from

August to December and fruit from January to

September.

21. Diospyros pentamera (Woolls & F.Muell.

ex F.Muell.) F.Muell, Austral. Veg. 35 (1867);

Cargillia pentamera Woods & F.Muell. ex

F.Muell., Fragm. 4: 82 (1864). Type: New

South Wales. Clarence River, s.dat., C. Moore

s.n. (lecto [here designated]: MEL 232965).

Cargillia arborea A.Cunn. ex Hiern, Trans.

Cambridge Phil. Soc. 12: 239 (1873), nom.

inval., pro. syn.

Illustration : Cooper & Cooper (2004: 156).

Tree to 35 m. Twigs with appressed long

basidxed pale brown hairs, glabrescent.

Leaves: petiole 2-6 mm long, glabrescent;

lamina elliptic or lanceolate, 3-10 cm long,

1- 4 cm wide; base attenuate; glands mostly

2- 10 on lamina below; margins slightly

recurved near base; apex acuminate; sparsely

appressed pubescent, glabrescent both sides,

minutely papillate below appearing dud pale

green and discolorous; midvein on upper

surface depressed; secondary veins 7-11 pairs,

sometimes indistinct. Male indorescence

axes solitary or few in leaf axils, each axis

4-12 mm long, each with 3-10 dowers,

appressed pubescent. Calyx tube 1.5-1.7 mm

long, lobes (4-)5(-6), 1.2-1.4 mm long, acute;

calyx appressed pubescent outside, glabrous

inside. Corolla tube 1.5-1.7 mm long, lobes

mostly 5, narrowly ovate or oblong, 3-3.5

mm long, appressed pubescent outside but

glabrous on the overlapped margin and inside.

Stamens 15-20, 3.7-4.5 mm long, dlaments

connate at base in pairs or alternipetalous

ones sometimes single and adnate to corolla

tube at base, 1-2 mm long, glabrous towards

base, pilose just below anthers; anthers linear-

rostellate, 2.5-3 mm long, connective pilose;

pistillode c. 1 mm long, pubescent. Female

dowers l(-3) in each axil, each on a pubescent

peduncle 1.5-5 mm long, bracts caducous.

Calyx tube 2.5-3 mm long, lobes (4-)5, 2-2.5

mm long, obtuse; calyx appressed pubescent

outside and inside. Corolla tube 1.5-2.2

mm long, glabrous outside and inside, lobes

(4-)5, oblong or narrowly ovate, 3-3.8 mm

long. Staminodes 10-12, like stamens with

sterile anthers, dlaments single or paired.

Ovary 2-2.5 mm long, sericeous, 3-locular,

ovules 2 per locule, style 1.7-2 mm long,

appressed pubescent, 3-branched for nearly

half its length and glabrous just below the

stigmas. Fruiting calyx cupular, 10-12 mm

diameter, appressed to base of fruit, lobes

broadly triangular, 2-4 mm long; fruit ovoid

or subglobose, 14-18 mm long, 10-15 mm

wide, appressed pubescent, glabrescent,

1-4-seeded; seeds 8-12 mm long. Fig. 7A-F.

Additional selected specimens examined : Queensland.

Cook District: Daintree NP, Adeline Creek headwaters,

Candlenut scrub. May 1999, Forster PIF24569 & Booth

(BRI); SFR 144 Mt Windsor Tableland, Oct 1979,

Moriarty 2685 (BRI); SFR 607, Parish of Cairns, Shoteel

LA, Sep 1986, Gray 4359 (BRI). North Kennedy

District: Scrubby Creek, Herberton Range, Nov 1929,

Kajewski 1359 (BM, BRI); Mt Fox, Nov 1949, Clemens

s.n. (BRI [AQ416986]); Mt Aberdeen NP, west of

Bowen, May 1992, Forster PIF9964 et al. (BRI). South

Kennedy District: SF 679 Crediton, Clarke Range, Apr

2000, Forster PIF25525 (BRI). Port Curtis District:

Resumption LA, SF 391, Bulburin, Dec 1993, Forster

PIF14521 et al. (BRI). Wide Bay District: c. 2 km

from Dundowran Beach on road to Hervey Bay, Nov

1978, Stanley 78125 & Ross (BRI); Mary River Heads,

Pialba, Nov 1987, Forster P1F3458 et al. (BRI); top of

Woowoonga Range, c. 15 km NE of Biggenden, Jun

1979, Young 201 & Randall (BRI). Burnett District:

Dripping Rock, Cania Gorge NP, Mar 1997, Kampf et al.

s.n. (BRI); roadside opposite entrance to Paradise Falls

carpark, Bunya Mountains NP, Dec 2008, Young 2412

(BRI). Moreton District: Brolga Park, Dulong Road, c.

6 km SW of Nambour, Dec 1989, Sharpe 4928 & Bean

(BRI); Wards Scrub, W of Samford near headwaters of

South Pine River, Dec 1983, Jessup 578 & Guymer (BRI);

Beechmont Ridge, Beechmont, Macpherson Range, Oct

188

Austrobaileya 9(2): 155-197(2014)

Fig. 7. A-F: Diospyrospentamera. A. male flower x4. B. dissected corolla showing stamens and pistillode, male flower

x6. C. female flower x4. D. dissected corolla showing staminodes, female flower x6. E. dissected ovary, female flower

x6. F. fruit x2. G-L: Diospyros mabacea. G. male flower bud x4. H. female flower x3,1. dissected corolla showing

stamens, male flower x8. J. dissected ovary, female flower x4. K. dissected corolla showing staminodes, female flower

x4. L. fruit xl.5. A, B from Stanley 78125 & Ross (BRI); C from Jessup 578 & Guymer (BRI); D from Forster PIF3458

et al. (BRI); E from Smith 3618 & Webb (BRI); F from Grimshaw G84 (BRI); G, H from Baeuerlen 1531 (BRI); I-K

from Murray s.n. (BRI [AQ651258]); L. reconstructed from Baeuerlen 1531 (BRI) and unvouchered photo.

189

Jessup, Diospyros in Australia

1969, Schodde 5590 (BRI); O’Reilly’s, Lamington NP,

May 1948, Smith 3618 & Webb (BRI); Qld/NSW border

fence reserve near ‘Wild Mountain’ property. Levers

Plateau, Oct 1993, Grimshaw G84 (BRI). New South

Wales. North Coast: Glenugie Peak, c. 12 miles [20

km] SE of Grafton, Nov 1965, Constable 6408 (BRI);

Dorrigo SF, Oct 1930, White 7512 (BRI). Central

Coast: 3 miles [4.8 km] W of Kiama on Jamberoo Road,

Illawarra District, Mar 1964, Schodde 3474 (BRI).

Distribution and habitat: Diospyros

pentamera is endemic to the eastern coast of

Australia, south of Cape Tribulation, northeast

Queensland to south of Kiama, New South

Wales (Map 13). It occurs in various types

of rainforest from sea level to at least 1100 m

altitude.

Phenology: Flowers have been recorded from

September to May and fruit from February to

July, also October and December.

22. Diospyros mabacea (F.Muell.) F.Muell.,

Syst. Census of Austral. PI. 92 (1883);

Cargillia mabacea F.Muell., Fragm. 5: 162

(1866). Type: New South Wales. Tweed River,

s.dat ., C. Moore s.n. (holo: MEL 92384; iso: K

792758).

Tree to 25 m. Twigs with erect or antrorse

basifixed pale brown hairs, glabrescent.

Leaves: petiole 3-8 mm long, glabrescent;

lamina narrowly elliptic or oblanceolate,

6-16 cm long, 2-5 cm wide; base shortly

attenuate; margins recurved near base; glands

mostly 2-10 scattered on lamina below;

apex acuminate; glabrous above except

main veins, sparsely appressed pubescent,

glabrescent below; midvein on upper surface

depressed, secondary veins 9-13 pairs. Male

inflorescence a fascicle of cymes, each axis

3-4 mm long, each fascicle with up to 6

flowers, pubescent, bracts narrowly ovate,

persistent. Calyx tube 2-2.3 mm long, lobes

4, 0.7-1.2 mm long, acute or obtuse; calyx

sparsely appressed pubescent, glabrescent

outside, glabrous inside. Corolla tube 2-2.5

mm long, lobes 4, narrowly ovate or oblong,

3.5-4 mm long, appressed pubescent outside

but glabrous on the overlapped margin and

inside. Stamens 15 or 16, 4-4.5 mm long,

filaments connate at base in pairs and adnate to

corolla tube at base, 1-1.5 mm long, glabrous;

anthers subulate, 2.4-2.8 mm long, glabrous

or with a few short hairs on the connective;

pistillode c. 1 mm long, pubescent. Female

flowers l(-3) in each axil, each on a pubescent

peduncle 4-6 mm long, bracts oblong,

persistent. Calyx tube 2.5-3 mm long, lobes

4, 4-4.5 mm long, obtuse; calyx sparsely

appressed pubescent outside, glabrous inside.

Corolla tube 2-3 mm long, glabrous outside

and inside, lobes 4, oblong, 6-7 mm long,

apex rounded, recurved. Staminodes 8, like

stamens with sterile anthers, filaments single.

Ovary 3-3.7 mm long, appressed pubescent,

4-locular, ovules 2 per locule, style 3-4 mm

long, appressed pubescent, but glabrous and

scarcely branched below the stigmas. Fruiting

calyx with recurved lobes 5-6 mm long; fruit

ovoid, narrowly ovoid or ellipsoid, 25-45 mm

long, glabrescent, 2-6-seeded; seeds 17-20

mm long. Fig. 7G-L.

Additional selected specimens examined : New South

Wales. North Coast: Murwillumbah, Oct 1896,

Baeuerlen 1531 (BRI, NSW); Oxley River, Tyalgum

Showground, Dec 1983, Floyd AGF2029 (BRI, NSW); 6

km NW of Tyalgum, 300 m NW beyond end of Butlers

Road, Feb 1990, Davies 1553a & Richardson (BRI);

Oxley River Middle Arm Creek, just beyond end of

Butlers Road, NW of Tyalgum, Jul 1981, Guymer 1586

& Jessup (BRI, CANB, NSW); Oxley River tributary,

Eungella, Dec 1983, Floyd AGF2030 (BRI, NSW); Portion

23, Parish of Chill ingham on E bank of Hopping Dicks

Creek, Oct 1996, Murray s.n. (BRI [AQ651258], NSW);

Murwillumbah, Sep 1900, Campbell s.n. (NSW 26402),

Nov 1900, Campbell s.n. (NSW 274846), Mar 1901

Campbell s.n. (NSW 274851); Tweed River District, Oct

1901, Campbell s.n. (BM); Tweed River, North Arm,

Mar 1917, Runners s.n. (NSW); main arm of Brunswick

River, c. 3 km W of Mullumbimby, Feb 1998, 0 ’Donovan

s.n. (NSW).

Distribution and habitat: Diospyros

mabacea is restricted to northeast New South

Wales, occurring in the Tweed and Brunswick

River valleys (Map 11). The species is found in

lowland complex notophyll vine forest.

Phenology: Flowers and fruit have been

recorded in October.

Notes: Diospyros mabacea is listed as

Endangered under the Environment

Protection and Biodiversity Conservation

Act 1999 and the NSW Threatened Species

Conservation Act 1995.

190

Excluded names

Diospyros cor difolia Roxb., PI. Coromandel

1: 38, t. 50 (1795).

Australian specimens referred to this name by

Bentham (1868: 286) are D. rugosula.

Diospyros ellipticifolia (Stokes) Bakh., Gard.

Bull. Straits Settlem. 7: 162 1933); Ferreola

ellipticifolia Stokes, Bot. Mat. Med. 4: 556

(1812), nom. illeg. et nom. superfl. pro Maba

elliptica J.R.Forst. & G.Forst., Char. Gen.

PL, ed. 2. 122, t. 61 (1776); Ebenus elliptica

(J.R.Forst. & G.Forst.) Kuntze, Revis. Gen. Pl.

2: 408 (1891).

Smith (1971) noted that Ferreola ellipticifolia

Stokes is an illegitimate name because it

was published with Maba elliptica J.R.Forst.

& G.Forst. in synonymy. However, he then

used the illegitimate combination D. elliptica

(J.R.Forst. & G.Forst.) RS.Green (a later

homonym of D. elliptica Knowlt. (Dorr

2011). Australian specimens identified as this

species (as the f. australiensis) by Bakhuizen

van den Brink (1941: 65, 429) are referrable to

D. yandina.

Diospyros ferrea (Willd.) Bakh., Gard. Bull.

Straits Settlem. 7: 162 (1933); Ehretia ferrea

Willd., Phytographia 4 (1794).

Bakhuizen van den Brink (1941: 57) stated

“inter tropicos Africae, Asiae et Australiae

dispersa”; however, no Australian specimens

were actually cited. Bakhuizen van den Brink

used species rank taxa to be all encompassing,

with the actual taxa that could be related

to species in a modern sense, usually

circumscribed as varieties or forma.

Diospyros ferrea f. lamponga (Miq.) Bakh.,

Bull. Jard. Bot. Buitenzorg ser. 3,15: 434,440

(1941); Maba lamponga Miq., FI. Ned. Ind.,

Eerste Bijv. 3: 584 (1861).

Bakhuizen van den Brink (1941: 434) stated

‘Habitat in Australia ore tropicali et in

Insulindie’; however, no Australian specimens

were actually cited.

Austrobaileya 9(2): 155-197 (2014)

Diospyros longipes Hiern, J. Bot. 52:

338 (1914). Type citation: “Townsville,

Queensland, 11 Aug. 1913, R.H. Cambage”

Bakhuizen van den Brink (1941: 49)

referred this material to Mimusops elengi L.

(Sapotaceae).

Diospyros major (G.Forst.) Bakh., Bull. Jard.

Bot. Buitenzorg ser. 3, 15: 429 (1941), adnot.

Maba major G.Forst.

This was proposed by Bakhuizen van den

Brink (1941: 429) as an alternative name to D.

ellipticifolia (Stokes) Bakh. (see above). The

name was misapplied by Jacobs & Pickard

(1981: 111) to material from New South Wales;

however, no specimens were cited at the time.

Diospyros montana Roxb., PL Coromandel 1:

37, 48 (1795).

Australian specimens referred to this species

by Bakhuizen van den Brink (1938: 265,

1941: 265-270) are D. rugosida. According

to Kostermans (1981: 22), D. montana is

restricted to southern India and Ceylon (Sri

Lanka).

Diospyros montana f. cordifolia (Roxb.)

Hiern, Trans. Cambridge Philos. Soc. 12: 222

(1873); D. cordifolia Roxb, PL Coromandel 1:

38, t. 50 (1796).

Australian specimens referred to this name

are Diospyros rugosula.

Maba rufa Labill., Serf. Austro-Caledon. 33:

t. 36 (1824).

Australian material identified with this name

by Hiern (1873: 114), Bailey (1900: 963, t. XL)

and Bakhuizen van den Brink (1941: 444) is

referrable to Diospyros cupulosa. Maba rufa

is a New Caledonian endemic now known as

D. labillardierei F.White.

Diospyros whyteana (Hiern) F.White,

Bothalia 7: 488 (1961).

In Australia this species is only known from

cultivated plants (Taplin & Symon 2008: 85).

191

Jessup, Diospyros in Australia

Acknowledgements

This work was partially funded by a grant

from the Australian Biological Resources

Study (ABRS) for the purposes of providing

an account of Ebenaceae for the Flora of

Australia. I would like to thank the directors

curators and staff of BM, BRI, CANB, CNS

(formerly QRS), K, L, MEL, NSW and P

for access to their collections and assistance

in various ways. Mr Andrew Murray

provided information on and specimens

of Diospyros mabacea. Dr Guido Roghi

from the Dipartimento di Geoscienze,

Universita di Padova kindly investigated

Massalongo's publications and specimens in

that institution's Paleontological Museum.

Mr David Fell provided me with useful

advice on the habitat preferences of various

North Queensland species. Mr Frank Zich

(CNS) provided information and photographs

of some specimens. Ms Rebecca Francis

provided technical assistance during early

stages of this project and Mr Will Smith

produced the maps and line drawings.

References

Bailey, F.M. (1900). The Queensland Flora. Part III. J.

Diddams & Co: Brisbane.

- (1913). Comprehensive Catalogue of

Queensland Plants. Government Printer:

Brisbane.

Baker, R.T. (1899). Contributions to a knowledge of the

flora of Australia. Proceedings of the Linnean

Society of New South Wales 24: 437-447.

Bakhuizen van den Brink, R.C. (1936-1955). Revisio

Ebenacearum Malayensium. Bulletin duJardin

Botanique de Buitenzorg, ser. 3, 15: 1-49

(1936); 50-368 (1938); 369-515 (1941); i-xx, pi.

1-92 (1955).

Bentham, G. (1868). Ebenaceae. Flora Australiensis 4:

285-292. L. Reeve: London.

Britton, N.L. & Brown, A. (1913). An Illustrated Flora

of the Northern United States , ed. 2, Vol. 2.

Charles Scribner’s Sons: New York.

Brown, R. (1810). [1960 fascimile], Prodromus florae

Novae Hollandiae et Insulae van-Diemen. J.

Cramer: New York.

Candolle, A. de (1844). Ebenaceae. In A.P de Candolle

(ed.), Prodromus Systematis Naturalis Regni

Vegetabilis 8: 209-243. Masson: Paris.

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

Australian Tropical Rainforest. Nokomis:

Melbourne.

Dorr, L.J. (2011). Diospyros foliosa: the correct name

for D. elliptica , and new combinations in Fijian

and Samoan Diospyros (Ebenaceae). Kew

Bulletin 66: 633-635.

Duangjai S., Wallnofer B., Samuel R., Munzinger J. &

Chase M.W. (2006). Generic delimitation and

relationships in Ebenaceae sensu lato: evidence

from six plastid DNA regions. American

Journal'of Botany 93: 1808-1827.

Duangjai S., Samuel R., Munzinger J., Forest F.,

Wallnofer B., Michael H.J., Barfuss M.H.J.,

Fischer G. & Chase M.W. (2009). A multilocus

plastid phylogenetic analysis of the pantropical

genus Diospyros (Ebenaceae), with an emphasis

on the radiation and biogeographic origins of the

New Caledonian endemic species. Molecular

Phylogenetics and Evolution 52: 602-620.

Endlicher, S. (1839). Ebenaceae. In Genera plantarum

secundum or dines naturales disposita 10: 741-

143.

Frodin, D.G. (2004). History and concepts of big plant

genera. Taxon 53: 753-776.

Hiern W.P (1873). A monograph of Ebenaceae.

Transactions of the Cambridge Philosophical

Society 12: 1-300.

Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray,

B. & Elick, R.W. (2003). Australian Tropical

Rain Forest Plants, Trees, Shrubs and Vines.

CD-ROM. CSIRO Publishing: Melbourne.

Jacobs, S.W. & Pickard, J. (1981). Plants of New South

Wales. D. West, Government Printer: Sydney.

Jessup, L.W. (1994). Ebenaceae. In R.J.F. Henderson

(ed.), Queensland Vascular Plants: Names

and Distribution, p. 101-102. Queensland

Herbarium: Indooroopilly.

- (1997). Ebenaceae. In R.J.F. Henderson (ed.),

Queensland Plants: Names and Distribution , p.

66-67. Queensland Herbarium, Department of

Environment: Indooroopilly.

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

(ed.). Names and Distribution of Queensland

Plants, Algae and Lichens , p.65. Queensland

Herbarium, Environmental Protection Agency:

Brisbane.

192

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

Holland (eds.), Census of the Queensland

Flora 2007 , p. 65-66. Queensland Herbarium,

Environmental Protection Agency: Brisbane.

- (2010). Ebenaceae. In P.D. Bostock & A.E.

Holland (eds ). Census of the Queensland

Flora 2010 , p. 60-61. Queensland Herbarium,

Department of Environment & Resource

Management: Brisbane.

- (2013). Ebenaceae. In P.D. Bostock & A.E.

Holland (eds.). Census of the Queensland

Flora 2013. Queensland Herbarium: Brisbane.

https://data.qld.gov.au/dataset/census-of-the-

queensland-flora-2013

Kostermans, A.J.G.H. (1977). Notes on Asiatic, Pacific

and Australian Diospyros. Blumea 23: 449-474.

- (1981). Ebenaceae. In M.D. Dassanayake &

F.R. Fosberg, (eds.), A Revised Handbook to the

Flora of Ceylon 3: 1-54. Amerind Publishing:

New Delhi.

Massalongo, A.B. (1859). Syllabus Plantarum Fossilium:

Hucusque in Formationibus Tertiariis Agri

Veneti Detectarum. 77. A. Merlo: Veronae.

Mueller, F. (1866). Fragmenta Phytographiae

Australiae 5: 164. Government Printer:

Melbourne.

- (1867). Australian vegetation, indigenous or

introduced, considered especially in its bearings

on the occupation of the territory, and with a

view of unfolding its resources. Intercolonial

Exhibition, 1866-7. Essay 5: 1-38. Blundell &

Austrobaileya 9(2): 155-197 (2014)

Co.: Melbourne.

Singh, V. (2005). Monograph on Indian Diospyros L.

(Persimmon, Ebony) Ebenaceae. Botanical

Survey of India: Kolkata.

Smith, A.C. (1971). Studies of Pacific Island plants,

XXIII. The genus Diospyros (Ebenaceae) in

Fiji, Samoa and Tonga. Journal of the Arnold

Arboretum 52: 369-403.

- (1981). Flora Vitiensis Nova 2: 729-744. Pacific

Tropical Botanical Garden: Lawai, Kauai,

Hawaii.

Stanley, T.D. & Ross, E.M. (1986,2002 reprint). Flora of

South-eastern Queensland Vol. 2. Queensland

Herbarium, Queensland Department of Primary

Industries: Brisbane.

Taplin, R.L. & Symon, D.E. (2008). Remnant

horticultural plants at the site of the former

Newman’s Nursery, 1854-1932. Journal of the

Adelaide Botanic Gardens 22: 85.

Wallnofer, B. (2001). The biology and systematics

of Ebenaceae: a review. Annalen des

Naturhistorischen Museums in Wien , Serie B,

103: 485-512.

- (2004). Ebenaceae. In K. Kubitzki (ed.). The

Families and Genera of Vascular Plants 6: 125—

130. Springer Verlag: Berlin, Heidelberg.

White, F. (1980). Notes on the Ebenaceae. VIII. The

African sections of Diospyros. Bulletin du

Jar din Botanique National de Belgique 50:

445-460.

Map 1. Distribution of Diospyros calycantha in Australia.

Jessup, Diospyros in Australia

193

Map 3. Distribution of Diospyros rugosula • and D. uvida A in Australia.

194 Austrobaileya 9(2): 155-197 (2014)

Map 4. Distribution of Diospyros maritima • and D. fasciculosa A in Australia.

Map 5. Distribution of Diospyros areolifolia A, D. compacta • and D. kaki in Australia.

Jessup, Diospyros in Australia

195

Map 6. Distribution of Diospyros humilis in Australia.

Map 7. Distribution of Diospyros geminata • and D. littorea A in Australia.

196

Austrobaileya 9(2): 155-197 (2014)

Map 8. Distribution of Diospyros laurina in Australia.

Map 9. Distribution of Diospyros peninsularis A, D.

hemicycloides • and D. yandina ■.

Map 11. Distribution of Diospyros granitica • and D.

mabacea A.

Map 10. Distribution of Diospyros pluviatilis and D.

rheophila •.

Jessup, Diospyros in Australia

197

Diploglottis alaticarpa W.E.Cooper (Sapindaceae),

a new species from Queensland’s Wet Tropics

W.E. Cooper

Summary

Cooper, W.E. (2014). Diploglottis alaticarpa W.E.Cooper (Sapindaceae), a new species from

Queensland’s Wet Tropics. Austrobaileya 9(2): 198-202. Diploglottis alaticarpa W.E.Cooper

is described and illustrated. Notes on habitat and distribution are provided, as well as a species

identification key for the genus.

Key Words: Sapindaceae, Diploglottis , Diploglottis alaticarpa , Australia flora, Queensland flora,

new species, taxonomy, rainforest

W.E. Cooper, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor

Road, Smithfield, Queensland 4878, Australia

Introduction

Diploglottis Hook.f. (Sapindaceae) was

noted as a genus of eight endemic species in

Australia by Reynolds (1985:33). A further two

Australian endemic species were described

by Reynolds (1987), viz. D. bernieana

S.T.Reynolds and D. obovata S.T.Reynolds.

Harden (1986) reinstated D. australis (G.Don)

Radik, with D.cunninghamii (Hook.) Hook.f

ex Benth. in synonymy. Leenhouts (1994:

520) synonymised D. cunninghamii (Hook.)

Hook.f. ex Benth. and D. diphyllostegia

(F.Muell.) F.M.Bailey with D. australis

and stated there to be 11 (although this

enumeration was overly incorrect) species

in northeast Australia, with one of those also

occurring in New Guinea and New Caledonia.

Leenhouts (1994) treatment of Diploglottis

diphyllostegia has not been accepted and it

has continued to be recognised as a distinct

entity (Hyland et al. 2003; Cooper & Cooper

2004; Forster & Jessup 2007). Including the

species newly described here, Diploglottis

(Sapindaceae) is now a genus of 11 species,

all occurring in Australia with 10 species

considered endemic. Diploglottis in Australia

is more widespread than indicated by

Leenhouts (1994), occurring in rainforest and

monsoon forests of eastern Australia from

Accepted for publication 4 August 2014

Cape York on northernmost Queensland to

the Illawarra district in southern New South

Wales.

A sterile specimen of an unsual Diploglottis

was first collected by Rigel Jensen before 1994

and determined by the then QRS herbarium

(now CNS) as Sapindaceae (Palmerston

Rigel Jensen s.n) as evidenced in Cooper

(1994: 312); however, the specimen appears

to be lost. Since 1994, subsequent collections

of this plant have been usually included

within D. bracteata Leenh. Additional fertile

specimens have conclusively shown that the

species described below as D. alaticarpa

W.E.Cooper is quite distinct from that species

in leaf and fruiting features. From herbarium

records and field observations, D. alaticarpa

and D. bracteata do not occur together and

are therefore allopatric.

Diploglottis was considered by Acevedo-

Rodriguez et al. (2011) to be falsely

polygamous and Gross (2005) described

D. smithii S.T.Reynolds as monoecious.

Whereas, vouchers from D. alaticarpa with

male flowers ( Cooper 2229 ) and with bisexual

flowers ( Cooper 2230), both collected from

the same tree but on different dates, confirm

the species to be polygamo-dioecious.

Based on Diploglottis species known at

the time, Leenhouts (1994) characterised the

genus as having fruit that are not winged;

Cooper, Diploglottis alaticarpa

however, the new species described herein has

distinctly winged sutures, a newly recognised

trait for Diploglottis.

Materials and methods

The study is based upon the examination of

herbarium material from CNS and BRI, as

well as field observations. All specimens cited

have been seen by the author.

Measurements of the floral parts and

fruits are based on material preserved in 70%

ethanol as well as fresh material from the

field.

Taxonomy

Diploglottis alaticarpa W.E. Cooper sp.

nov. Similar to D. bracteata Leenh. but

differs in the new leaves (silvery-pink versus

silvery-green), rachis (sericeous becoming

glabrescent versus persistently sericeous),

leaflet upperside (shiny versus dull), primary

vein on leaflet upperside (slightly raised

versus flat or slightly raised within a distinctly

deep and narrow groove), fruit (winged versus

not winged, sparsely hairy versus puberulent,

opening widely and becoming campanulate to

allow seeds to fall versus opening slightly and

not changing shape (never campanulate) with

valves separating and dropping to allow seeds

to fall or whole fruits fall and dehisce on the

ground). Typus: Queensland. Cook District:

Southern side of Palmerston Highway near

Mamu Boardwalk, Wooroonooran National

Park [west of Innisfail], 19 December 2013,

W.Cooper 2232 & R.Jensen (holo: CNS [3

sheets + spirit]; iso: BRI, CANB, K, L, MO

distribuendi).

Sapindaceae (Palmerston Rigel Jensen s.n.);

Cooper & Cooper (1994: 312).

Tree to 22 m, monoecious, dbh to 40 cm; trunk

not fluted, bark lacking distinctive features;

branchlets shallowly ribbed, minutely

lenticellate, glabrous; new growth silvery-

pink and sericeous. Leaves compound,

alternate; rachis + petiole 12-41 cm long,

sericeous but soon glabrescent, flattened

along upperside, ribbed, minutely lenticellate,

pulvinus 7-12 mm long. Leaflets subopposite

to alternate, coriaceous, 9-19; petiolules 7-25

mm long, grooved on upperside, pulvinulus

199

5-12 mm long, glabrescent; lamina oblong,

oblong-obovate or oblong-elliptical, 44-185

mm long, 18-63 mm wide; base cuneate,

sometimes asymmetrical; apex emarginate;

margin entire; upperside glabrous or with

sparse minute hairs on secondary veins;

underside glabrescent with sparse appressed

minute hairs; venation camptodromous;

primary vein slightly raised; secondary

veins 8-15 pairs, slightly raised on upperside

and distinctly raised on underside; tertiary

venation reticulate. Inflorescence an axillary

or pseudo-terminal panicle up to 240 mm

long, sericeous; rachis ribbed, pulvinate;

cymules 4-flowered; bracts caducous, ovate,

base truncate, apex acute, 12-25 mm long,

3-7 mm wide, both sides sericeous. Flowers

actinomorphic, unisexual (staminate) or

apparently bisexual, diameter 2.3-5.3 mm,

buds broadly ovoid; pedicels 1.5-2 mm long,

terete; calyx shortly cupular, 5 lobed, lobes

c. 2 mm long and wide, both sides sericeous;

petals 5, broadly orbicular or obovate, 1.3-2.5

mm long, 1.5-2.5 mm wide, outside sparsely

pilose proximally, pilose inside, margin

ciliate; scale divided to base, crested, pilose;

disk annular, thick, not interrupted, glabrous;

stamens 7 or 8, filaments to 2 mm long, pilose,

hairs proximally denser; anthers oblong,

bilocular, c. 1 mm long, basifixed, dehiscing

laterally, glabrous; pistillode sericeous,

3-celled; ovary sericeous, c. 2 mm long,

3-locular; ovules 1 per locule; style sparsely

hairy. Fruiting pedicel 5-8 mm long; fruit

a 3-(rarely 4-) locular capsule, triangular-

oblate (unless 1 or 2 ovules are aborted then

asymmetricaly oblate or globular), 26-33 mm

long, 23.5-44 mm wide, yellowish-green and

mostly blushed with pink proximally, sparsely

and minutely hairy; sutures winged in basal

half, wings 2.5-4 mm wide; walls leathery,

orange and sericeous inside, opening widely

and becoming campanulate to allow shedding

of seeds; seeds 1-3, lenticular, 14-26 mm

long, 16-22 mm wide, 9-14.5 mm thick,

testa dark brown; aril 2-lobed, orange-red,

completely or almost completely covering

seed, margin thin. Fig. 1.

Additional specimens examined : Queensland. Cook

District: Pullom Road, Palmerston, Sep 2013, Cooper

2228 & Cooper (CNS); loc. cit ., Oct 2013, Cooper 2229

(CNS); loc. cit., Nov 2013, Cooper 2230 (CNS); Pullom

200

Austrobaileya 9(2): 198-202 (2014)

Fig. 1. Diploglottis alaticarpa fruit A. Indehisced fruit viewed from base, showing winged sutures. B. Dehisced fruit

from apex, showing sericeous inner valves and arillate seeds. C. Seed with aril removed. D. Seeds with arils intact

showing pale coloured attachment scar. All natural size. Del. W.T.Cooper

Road, 17 km WNW of South Johnstone, Dec 2003,

Forster PIF29795 & Jensen (BRI); Wooroonooran NP

Palmerston Section, Dec 1998, Jensen 947 (BRI); Near

Bora Creek, Johnstone River, Oct 1995, Jensen 432

(CNS); Crawfords Lookout, Palmerston NP, Dec 2013,

Cooper 2233 & Jensen (CNS); NPR 904, Palmerston,

near Crawfords Lookout, Jan 1997, Jensen 812 (CNS).

Distribution and habitat : Diploglottis

alaticarpa is endemic to the Wet Tropics

bioregion in north-east Queensland, where

it is currently known to occur mostly

within Wooroonooran National Park in the

Palmerston section, between the Johnstone

River and the South Johnstone River west of

Innisfail, altitude 100-362 m (Map 1).

Diploglottis alticarpa grows as an

understory tree in complex mesophyll

vine forest on basalt soil. It commonly co¬

occurs with Aglaia ferruginea C.TWhite &

W.D.Francis, Argyrodendron trifoliolatum

F.Muell., Cardwellia sublimis F.Muell.,

Eupomatia laurina R.Br., Flindersia

brayleyana F.Muell., Myristicaglobosa subsp.

muelleri (Warb.) WJ.de Wilde, Neolitsea

dealbata (R.Br.) Merr., Rhysotoechia

robertsonii (F.Muell.) Radik., Synima

macrophylla S.T.Reynolds and Syzygium

gustavioides (F.M.Bailey) B.Hyland.

Phenology : Flowers have been recorded in

October and November: fruits have been

recorded in December and January.

Affinities : Diploglottis alaticarpa appears to

be most similar to D. bracteata. The leaves,

flowers, arils and seeds are similar but the

two species differ most remarkably by the

Cooper, Diploglottis alaticarpa

colour of new leaf growth (silvery-pink versus

silvery-green), primary vein (virtually flush

with leaf blade versus slightly raised or flush

in a deep groove) and fruit sutures (winged

versus not winged).

201

Etymology : The specific epithet comes from

the Latin alatus (winged), carpus (fruit)

referring to the winged fruit, an unusual trait

for Diploglottis.

Identification key to Diploglottis species

1 Fruit sutures outwardly ribbed or winged.2

1. Fruit sutures smooth or depressed.3

2 Fruit green, sparsely hairy, sutures distinctly winged; leaflets

flat, glabrous or with sparse minute pale hairs. D. alaticarpa

2. Fruit yellow or orange, densely rusty hairy, sutures ribbed not

truly winged; leaflets bullate and rusty hairy. D. bernieana

3 Fruit glabrous or sparsely hairy. 4

3. Fruit velvety or densely hairy.6

4 Aril orange; young growth, peduncles and leaf axes villous with deep red

hairs; leaflets 5-10 pairs. D. pedleyi

4. Aril red (with the exception of a very rare occurrence of orange);

indumentum of pale hairs; leaflets 2-4 pairs.5

5 Fruit valves thick; petiolules 10-35 mm long; inflorescence 10-25

mm long; occurs north of Innisfail NE Qld. D. harpulliodes

5. Fruit valves thin, aril rarely orange; petiolules 2-5 mm long; inflorescence

120-160 mm long; occurs in SE Qld and NE NSW. D. campbellii

6 Leaflets distinctly bullate. 7

6. Leaflets flat, secondary veins may be depressed but not truly bullate.9

7 Indumentum of pale hairs. D. smithii

7. Indumentum of rusty or dark brown hairs.8

8 Sparsely branched or mostly an unbranched tree to 6 m; new

growth pink; occurs north from Cooktown NE Qld. D. macrantha

8. Tree to 35 m; new growth green; occurs south from Mt Walsh

NP SE Qld and NE NSW. D. australis

9 Indumentum silvery; primary vein slightly raised or flush in a deep groove . D. bracteata

9. Indumentum rusty; primary vein not deep with a groove. 10

10 Leaflet apices shortly acuminate; occurs north from Eungella,

central eastern Qld. D. diphyllostegia

10. Leaflet apices obtuse or truncate; occurs from Sarina to Proserpine

area, central eastern Qld. D. obovata

Acknowledgements

I am especially grateful to Rigel Jensen for

field assistance. I also thank Darren Crayn

and Frank Zich for support and access

to CNS herbarium, Bill Cooper for the

illustrations, Andrew Ford and Rigel Jensen

for comments to the manuscript and Steve

Murphy for the distribution map. Permits to

collect in the Wet Tropics were issued by the

Queensland Department of Environment and

Heritage Protection to the Australian Tropical

Herbarium.

202

References

Acevedo-Rodriguez, P., Van Welzen, P.C., Adema, F. &

Van Der Ham, R.W. J.M. (2011). Sapindaceae. In

K. Kubitzki (ed.). Flowering Plants. Eudicots:

Sapindales, Cucurbitales, Myrtaceae. The

Families and Genera of Vascular Plants, pp.

357-407. Springer Verlag: Berlin/Heidelberg.

Cooper, W. & Cooper, W.T. (1994). Fruits of the Rain

Forest. Geo: Sydney.

- (2004). Fruits of the Australian Tropical

Rainforest. Nokomis Editions: Melbourne.

Forster, PI. & Jessup, L. (2007). Sapindaceae. In P.D.

Bostock & A.E. Holland (eds ), Census of the

Queensland Flora 2007 , p. 185. Environmental

Protection Agency: Brisbane.

Gross, C.L. (2005). A comparison of the sexual systems

in the trees from the Australian tropics with

other tropical biomes - more monoecy but why?

American Journal of Botany 92: 907-919.

Austrobaileya 9(2): 198-202 (2014)

Harden, G.J. & Johnson, L.A.S. (1986). A note on

Diploglottis australis (G.Don) Radkl. Telopea

2: 745-748.

Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray,

B. & Elick, R. (2003). Australian Tropical Rain

Forest Plants. Trees, Shrubs and Vines. CD-

ROM. CSIRO Publishing: Melbourne.

Leenhouts, P.W. (1994). Diploglottis. Flora Malesiana

ser.l, 11: 520-522. Rijksherbarium/Hortus

Botanicus: Leiden.

Reynolds, S.T. (1985). Sapindaceae. In A.S. George

(ed.). Flora of A ustralia 25: 4-164. Australian

Biological Resources Study: Canberra.

- (1987). Notes on Sapindaceae, V. Austrobaileya

2: 328-332.

Map 1. Distribution of Diploglottis alaticarpa in northeast Queensland. Shaded area on map indicates Wooroonooran

National Park.

Ptilotus senarius A.R.Bean (Amaranthaceae), a

new species from northern Queensland

A.R. Bean

Summary

Bean, A.R. (2014). Ptilotus senarius A.R.Bean (Amaranthaceae), a new species from northern

Queensland. Austrobaileya 9(2): 203-206. The new species Ptilotus senarius A.R.Bean, recorded

only between Georgetown and Croydon, is described, illustrated, and differentiated from similar

taxa. A conservation status of Presumed Extinct is proposed.

Key Words: Amaranthaceae, Ptilotus, Ptilotus senarius, Australia flora, Queensland flora, new

species, taxonomy, presumed extinct, conservation status

A.R. Bean, Queensland Herbarium, Department of Science, Information Technology, Innovation

and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Email: Tony. Bean@dsitia. qld. gov. au

Introduction

Ptilotus is a genus of about 100 species, of

which 31 occur in the four eastern states of

Australia (Bean 2008). Within the family

Amaranthaceae, it is characterised by the

alternate leaves, the ovoid to cylindrical

inflorescences, single-seeded fruits and the

flowers with five tepals bearing hairs at least

towards the base.

In late 2008, the present author

distinguished the species here described from

herbarium specimens at BRI. They had been

identified by G. Beni as Ptilotus exaltatus var.

semilanatus (now known as P. nobilis subsp.

semilanatus (Lindl.) A.R.Bean).

The new species does appear to have some

affinity to Ptilotus nobilis subsp. semilanatus ,

although it is geographically disjunct from that

taxon. It has similar vegetative characteristics

to P. fusiformis (R.Br.) Poir., but is very

different in floral morphology. The new

species is diagnosed against these two taxa.

Despite a relatively small target search

area, no one has been able to relocate the

species in the field and it is presumed Extinct

at present.

Accepted for publication 4 August 2014

Materials and methods

All herbarium specimens of Ptilotus held

at BRI and MEL have been examined.

Measurements were made from dried

material.

Taxonomy

Ptilotus senarius A.R.Bean sp. nov. P.

nobili subsp. semilanato affinis sed foliis

angustioribus, caulibus pilis stellatis

praeditis, ovario glabro et inflorescentiis

paucifloris differens. Typus: Queensland.

Cook District: Riverview, Gilbert River, 10

May 1967, S.J. Dansie 3873 (holo: BRI; iso:

CANB, distribuendi ; CNS, image seen).

Ptilotus sp. (Gilbert River L.J. Brass 430);

Bostock & Holland (2010: 9).

Slender branched shrub 30-60 cm high, with

a woody base. Stems terete, densely stellate

hairy on new growing tips, but only sparsely

hairy elsewhere. Hairs stellate or pseudo-

stellate (with the radial arms perhaps not

all arising from the same point), 0.15-0.3

mm diameter, broader than high, white, rays

3-12, no central ray discernable. Leaves

alternate, linear, 7-31 mm long, 0.6-1.5 mm

wide, apex acute, base attenuate, petiole

absent, glabrous or with sparse stellate hairs.

Inflorescence terminal, l-6(-8)-flowered,

with flowers disposed at various angles to the

stems and rachis; bracts persistent on rachis,

204

broadly ovate, cymbiform, 27-3.2 mm long,

apex acute, outer surface densely covered by

stellate and verticillate hairs except laterally;

bracteoles caducous, ovate, cymbiform,

3.8-4.6 mm long, apex aristate, outer surface

densely covered by verticillate and stellate

hairs; perianth 15-19 mm long at anthesis;

tepals 5, fresh colour unknown, grey when

dried, linear, straight, apex acute, dorsal surface

densely covered by antrorse verticillate hairs

O. 4-0.6 mm long, and antrorse septate hairs

2.3-6 mm long; distal 1-1.5 mm of tepals

without any hairs attached. Two outer tepals

slightly longer than three inner tepals; outer

tepals glabrous on ventral surface; inner tepals

with dense woolly hairs on proximal one-third

of ventral surface, otherwise glabrous. Fertile

stamens 3, 7-10 mm long; filaments slender,

brown; anthers c. 1.7 mm long, caducous.

Ovary glabrous throughout. Style 8-12 mm

long, mostly straight, but sinuously curved at

about halfway along its length, eccentrically

inserted on ovary; stigma expanded. Figs. 1,2.

Additional specimen examined : Queensland. Cook

District: Gilbert River, Jun 1925, Brass 430 (BRI).

Distribution and habitat : Ptilotus senarius

has been collected only near the Gilbert River,

between Georgetown and Croydon in northern

Queensland. The habitat given for the two

collections is somewhat contradictory: one

label with “in tea-tree forest” i.e. dominated

by Melaleuca spp., most of which favour low-

lying areas; while the other with “on grassy

hills”.

Phenology : Flowers are recorded for May and

June.

Affinities : Ptilotus senarius is perhaps closely

related to P. nobilis subsp. semilanatus, as they

both have narrow leaves, tepals of about the

same length, and similar tepal indumentum.

However, P. senarius is readily distinguished

by the leaves less than 1.5 mm wide and

without sinuate margins (leaves 2-10 mm wide

and with sinuate margins for P. nobilis subsp.

semilanatus), the glabrous ovary (hairy for

P. nobilis subsp. semilanatus), inflorescences

l-6(-8)-flowered (30-100-flowered for

P. nobilis subsp. semilanatus), flowers

projecting at various angles from the rachis

(flowers perpendicular to rachis and stem for

Austrobaileya 9(2): 203-206 (2014)

P. nobilis subsp. semilanatus) and the stems

with stellate hairs (hairs absent or verticillate

for P. nobilis subsp. semilanatus).

Ptilotus senarius is similar to P.fusiformis

in height and form of the plant, and the similarly

linear leaves. The stems and leaves of both

species appear to the naked eye to be glabrous.

However, P. senarius is readily distinguished

by its stems with small stellate hairs

(glabrous for P. fusiformis), inflorescences

l-6(-8)-flowered (12-27-flowered for P.

fusiformis), perianth 15-19 mm long (8-13

mm long for P. fusiformis), and the tepals

with both verticillate and septate hairs on the

dorsal surface (with septate hairs only for P.

fusiformis).

Conservation status: It is considered that

the distribution of the species is relatively

well circumscribed, and nearly 50 years have

elapsed since the most recent known sighting.

A major road passes through the Gilbert

River area, giving ready access to botanists,

and the May-June period (when the previous

collections were made) is a popular time for

travel in this region. Hence one expects that

further serendipitous collections would have

been made since 1967, if the species were

present.

Since the recognition of the species in

late 2008, several active searches have been

made in the Gilbert River area, mainly around

May and June, but also at other times of the

year. During these years, the area has been

subject to a widely varying regime of rainfall

and temperature patterns, including periods

suitable for germination of herb and shrub

species. Much of the area is without major

weed infestation (except in riparian areas),

but cattle grazing has been a continuous land

use for a century or more. Ptilotus species

are known to be sensitive to grazing pressure

(Letnic 2004; Fensham et al. 2010).

Taking into consideration the grazing

threat, the well circumscribed distribution,

the time elapsed since the most recent

collection, and the inability to locate further

plants, a category of Presumed Extinct (X)

is proposed (IUCN 2001).

Bean, Ptilotus senarius

205

Queensland Herbarium (BRI) I

£3. ^3oj\

FftQ00008292

Honor QUEENSLANB

SSS3 S&I&&

„S.J. Dansie

10 May 1967

'i Airtata "‘PE Ig jgla I- l-l- |-> _

Ptilotus exaltatus

semilanatus (Lindlcy ex Mitch.) Maiden & Betehe

det. Q £rNL- VW I

PtiCotuvi Neef

VAr, fWUpudtu (£Wl .) &zA,

M^det. 7/ ^ Herb.

Queensland

herbarium

131712

BRISBANE

Fig. 1. Holotype of Ptilotus senarius (Dansie 3873 ) (BRI).

206

Austrobaileya 9(2): 203-206 (2014)

References

Bean, A.R. (2008). A synopsis of Ptilotus

(Amaranthaceae) in eastern Australia. Telopea

12: 227-250.

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

of the Queensland Flora 2010. Queensland

Herbarium, Department of Environment and

Resource Management: Brisbane.

Fensham, R.J., Fairfax, R.J. & Dwyer, J.M. (2010).

Vegetation responses to the first 20 years of

cattle grazing in an Australian desert. Ecology

91: 681-692.

Iucn (2001). IUCN Red List Categories and Criteria.

Version 3.1. Gland: IUCN - The World

Conservation Union.

Letnic, M. (2004). Cattle grazing in a hummock

grassland regenerating after fire: the short¬

term effects of cattle exclusion on vegetation in

south-western Queensland. Rangeland Journal

26: 34-48.

t

Fig. 2. Inflorescences of Ptilotus senarius (Dansie 3873 )

(BRI).

Note: The isotype at CNS bears the locality

“Riverview, Gilbert River”, whereas the

holotype specimen at BRI merely states

“Gilbert River”.

Etymology: From the Latin senarius meaning

‘consisting of six’. This refers to the few

flowers making up each inflorescence, usually

no more than six.

Acknowledgements

I am grateful to Keith McDonald for his several

dogged attempts to relocate this species in

the field. I thank the Director of MEL for

access to their collections, and Frank Zich

for sending images of the Dansie specimen

at CNS. Peter Bostock kindly translated the

diagnosis into Latin. Will Smith produced the

specimen scans.

Diversity on a tropical sky island: two new species

of Plectranthus L.Her. (Lamiaceae) from the

Hann Tableland, north-east Queensland

Paul I. Forster

Summary

Forster, P.I. (2014). Diversity on a tropical sky island: two new species of Plectranthus L.Her.

(Lamiaceae) from the Hann Tableland, north-east Queensland. Austrobaileya 9(2): 207-215.

A remarkable array of Plectranthus populations occurs on boulder and pavement fields on the

geographically isolated Hann Tableland west of Mareeba in north-east Queensland. Seven species are

present, namely P. bellus P.I.Forst., P. bipartitus P.I.Forst. sp. nov., P. congestus S.T.Blake, P.foetidus

Benth., P. mirus S.T.Blake, P. spectabilis S.T.Blake and P. splendens P.I.Forst. sp. nov. The newly

described species are illustrated and a distribution map is presented to all the species of Plectranthus

on the Hann Tableland.

Key Words: Lamiaceae, Plectranthus , Plectranthus bipartitus , Plectranthus splendens , Australia

flora, Queensland flora, Hann Tableland, new species, taxonomy, sky island, conservation status

P.I.Forster, Queensland Herbarium, Department of Science, Information Technology, Innovation

and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Email: Paul. Forster@dsitia. qld. gov. au

Introduction

The genus Plectranthus L.Her. is diverse in

Australia, with over 40 species described so

far. Most species are boulder or pavement

specialists where they grow in shallow,

skeletal soils. These habitats can be variable

within themselves, with a matrix of ‘micro

environments’ and the plants that occur often

do so in narrow bands or patches determined

by soil depth and moisture. These physical and

edaphic factors, compounded by geographic

isolation and the breeding biology of these

herbs or subshrubs are thought to have driven

speciation in eastern Australia (Forster 2011)

resulting in a mix of widespread species and

narrowly distributed endemics.

The Hann Tableland is a massive,

geographically isolated granite pluton

nearly 30 x 6 km in area and is located

west of Mareeba in north-east Queensland.

The granites of the tableland are part of the

Kennedy Province Granites, specifically the

Whypalla Supersuite and have been dated as

early Permian (275 ±15 Ma) in age (Champion

& Bultitude 2013). The topography of the

Accepted for publication 18 August 2014

Hann Tableland ranges from c. 350 m at its

base with the surrounding foothills to high

points of 950-1000 m. From north to south,

the tableland runs roughly south-southwest

with gentle to very steep slopes. Much of the

tableland is covered in woodland; however,

some gully systems and high points have closed

forests (rainforests). Interspersed throughout

the woodland ecosystems are large areas of

outcropping rock characterised by extensive

areas of pavement and small to massive

boulders or outcrops. These latter habitats

have shallow to no soil cover and usually

have vegetation that is markedly seasonal,

comprising a mixture of grasses, sedges and

other herbs. This ‘ephemeral flush’ vegetation

(Porembski et al. 2000) has relatively few

perennial species; these are generally woody

with either deep or lateral, water seeking

roots, are resurrection plants (e.g. Borya

septentrionalis F.Muell., Cheilanthes spp.), or

in the case of Plectranthus species have fleshy

to succulent rootstocks and foliage.

The Hann Tableland pluton is a ‘sky island’

(Heald 1951; Watling & Donnelly 2006) as it is

surrounded by other geology (metasediments)

at lower altitudes and is geographically

disjunct from other granite massives of the

208

Whypalla Supersuite (e.g. Bakers Blue, Mount

Windsor Tableland/Daintree National Park).

The upland vegetation and flora on the Hann

Tableland is in part refugial in composition

(particularly the rainforest communities, to

a lesser extent the woodland communities);

however, the plant communities on skeletal

soils exhibit a low level of localised endemism

by the presence of species such as Stylidium

elachophyllum A.R.Bean & M.T.Mathieson

and the two species described here. The

presence of these plant species, together with

a number of insects (Bouchard & Brooks

2004) reinforces the hypothesis that the Hann

Tableland comprises an area of biodiversity

endemism. The sharing of some species (e.g.

Plectranthns bellus P.I.Forst., Pterostylis

caligna M.T.Mathieson) from these

communities with similar habitats elsewhere

(e.g. Mount Windsor Tableland/Daintree

National Park) indicates past, relatively recent

dispersal events rather than vicariance. This

hypothesis is based on the reproductive

biology of the plants listed, reiterating my

previous comments (Forster 2011: 387) and

the irregular outcropping petrogenesis of the

granites and their subsequent erosion over the

past 270 Ma (Champion & Bultitude 2013).

This recent dispersal has also undoubtedly

been responsible for the various populations

of Plectranthus known from the Hann

Tableland, especially those known from one

or few populations.

Extensive fieldwork on the Hann Tableland

has revealed a large number of Plectranthus

populations (Map 1). Generally a single or

two species occur at any one site; however,

where two species are present, there is often

subtle edaphic separation of plants based on

soil depth and seasonal water supply. The co¬

occurring species are markedly different in

morphology and appear to be reproductively

isolated as hybrids have not been seen.

This reproductive isolation is apparently

maintained by genetic differences or in some

instances by phenology.

Seven species of Plectranthus have now

been recorded from the Hann Tableland.

Four of these are relatively widespread in

north-east Queensland, viz. P. congestus

Austrobaileya 9(2): 207-215 (2014)

R.Br., P. foetidus Benth., P. mirus S.T.Blake,

P. spectabilis S.T.Blake; one has a more

restricted distribution between the Mount

Windsor Tableland and Daintree National

Park, viz. P. bellus , and two are newly

described in this paper.

Materials and methods

Fieldwork on the Hann Tableland was

undertaken between 2006 and 2013 with

the use of helicopters to explore isolated or

remote areas of suitable habitat in 2010 and

2013; however, due to the rugged vastness of

the area further exploration is still warranted.

Specimens were prepared in the field or from

material cultivated in a garden in Brisbane.

Six or more seedlings were collected from

populations selected for further study

using cultivated plants. Descriptions and

illustrations were prepared from cultivated

plants.

Taxonomy

Plectranthus bipartitus P.I.Forst. sp. nov.

Distinct within Australian Plectranthus by

the bipartite form of the individual cymes

of the verticillasters and the extensive two-

tiered indumentum, particularly glandular

trichomes and ‘micro’ glandular papillae.

Typus: Queensland. Cook District: Hann

Tableland National Park, southern end;

west of Mareeba, 8 April 2013, P.I. Forster

PIF39595 (holo: BRI [2 sheets + spirit]; iso:

CNS, K, MEL distribuendi).

Erect herb or subshrub to 100 cm high; foliage

with very strong sweetly aromatic scent when

crushed, somewhat clammy; non-glandular

and glandular trichomes uncoloured, sessile

glands absent. Roots thickened-tuberous

to fibrous. Stems square, erect to rarely

straggling, fleshy-succulent, easily snapped

without obvious stringy fibres, the lower parts

up to 9 mm diameter, pale green, upper parts

with persistent indumentum, non-glandular

trichomes sparse, divaricate to weakly

retrorse, 6-8-celled up to 0.5 mm long,

glandular trichomes dense, variable length

to 0.3 mm. Leaves discolorous, petiolate;

petioles 10-21 x 2-3.5 mm, channelled

on top, non-glandular trichomes sparse,

divaricate, 6-8-celled up to 1 mm long,

Forster, Plectranthus from Hann Tableland

209

glandular trichomes dense, variable length to

0.3 mm; laminae ovate to broadly ovate, firm

fleshy, weakly keeled, 35-110 x 27-105 mm,

crenate with 8-10 teeth up to 12 mm long on

each margin, of similar length along margin,

secondary teeth usually present; tip acute;

base rounded to truncate; upper surface mid¬

green, veins deeply impressed, non-glandular

trichomes sparse, divaricate, 6-8-celled up

to 1 mm long, glandular trichomes sparse,

variable length to 0.3 mm long; lower surface

pale green, somewhat scabrid, veins strongly

raised, non-glandular trichomes sparse

(denser on veins), divaricate, 6-8-celled up

to 1 mm long, glandular trichomes sparse

to moderately dense, variable length to

0.3 mm long. Inflorescence up to 300 mm

long, usually single with no side branches,

pedunculate for 20-30 mm, often with very

small subtending leaves; axis square in cross-

section, non-glandular trichomes sparse,

divaricate, 6-10-celled up to 1.2 mm long and

with numerous ‘micro’ trichomes, glandular

trichomes dense, up to 0.3 mm long and with

dense ‘micro’ papillae; bracts broadly ovate

to obovate, 1.7-2 x 1-1.8 mm, ecomose,

non-glandular trichomes sparse, divaricate,

6-8-celled up to 0.5 mm long, glandular

trichomes dense, up to 0.3 mm long and

with dense ‘micro’ papillae; verticillasters

12-20-flowered, 5-6 mm apart; cymes

very shortly pedunculate for 0.5-1 mm,

bipartite; pedicels 2-2.8 X c. 0.3 mm, non-

glandular trichomes scattered or occasional,

divaricate, 4-8-celled up to 0.6 mm long,

glandular trichomes sparse to dense, up to

0.3 mm long and with dense ‘micro’ papillae.

Flower calyces 2-2.5 mm long, non-glandular

trichomes scattered, divaricate, 6-8-celled

up to 0.8 mm long and with ‘micro’ papillae,

glandular trichomes sparse and with ‘micro’

papillae. Corolla 7-8 mm long, lilac (mauve),

‘micro’ papillae absent; tube 4.8-5 mm long,

curved at 70-90° c. 2 mm from the base,

not curved upwards, ± glabrous or with an

occasional non-glandular trichome 2-4-celled

and up to 0.3 mm long; upper lobes orbicular-

ovate, erect or slightly reflexed, c. 1.5 x

1.5 mm, non-glandular trichomes sparse,

divaricate, 4-6-celled up to 0.5 mm long,

glandular trichomes usually absent; lateral

lobes oblong, c. 1 x 0.7-0.8 mm, glabrous;

lower lobe broadly ovate, 4-4.5 x 3.5-4 mm,

non-glandular trichomes scattered, divaricate,

4-6-celled up to 0.5 mm long, glandular

trichomes scattered; filaments filiform, 7-8 x

c. 0.2 mm, lilac, fused for 2-4 mm from the

base; anthers 0.3-0.4 x 0.25-0.35 mm; style

filiform, 9-10 x c. 0.2 mm, lilac, bifid for c.

O. 1 mm. Fruit calyces 3.5-4 mm long; upper

lobe oblong-ovate, 1.2-1.5 x c. 9 mm; lateral

lobes lanceolate-falcate to lanceolate, 1.5-2

x c. 0.6 mm; lower lobes lanceolate-falcate,

2-2.1 x 0.5-0.7 mm. Nutlets n.v. Fig. 1.

Additional specimens examined : Queensland. Cook

District: Hann Tableland, May 1997, Jago 4344 &

Jensen (BRI); Hann Tableland NP, northern end of

tableland; west of Mareeba, Apr 2013, Forster PIF39690

(BRI, CNS, MEL).

Distribution and habitat. Plectranthus

bipartitus is currently known from two

populations from northern and southern

ends of the Hann Tableland (Map 1). In

both instances, plants occur in skeletal soils

on steep granite slabs at altitudes between

570 and 815 m in association with seasonal

ephemeral flush vegetation, predominantly of

grasses, sedges and annual herbs.

Notes : Plectranthus bipartitus was first

collected by Bob Jago and Rigel Jensen in

1997. It is a large and distinctive plant and

likely to be noticed by botanical collectors due

to its presence on exposed areas of pavement.

Plectranthus bipartitus has a number

of distinct features that are not present in

other Australian species from this genus.

There is a passing superficial similarity to

P. diversus S.T.Blake; however, that species

has a combination of leaf morphology and

indumentum (especially the widespread

occurrence of sessile orange glands), combined

with inflorescence structure that is not present

in P. bipartitus. The bipartite nature of the

cymes in P. bipartitus is unique in Australian

Plectranthus. The widespread occurrence

of two-tiered indumentum layers (Fig IB),

particularly the abundant glandular ‘micro’

papillae on the foliage and inflorescence

of P. bipartitus is also diagnostic. The

occurrence of ‘micro’ papillae on the foliage

of other Australian Plectranthus is relatively

Fig. 1 . Plectranthus bipartitus. A. habit of flowering stem x0.4. B. adaxial leaf surface x0.6. C. abaxial leaf surface

x0.6. D. indumentum on stem indicating two-tiered nature x24. E. bipartite cyme of verticillaster x3. F. floral bract

xl2. G. lateral view of flower x6. H. face view of flower x6.1. lateral view of flower pedicel and calyx xi2. J. lateral

view of fruiting pedicel and calyx xl2. All from Forster PIF39595 (BRI). Del. W.Smith.

Forster, Plectranthus from Hann Tableland

211

widespread; however, usually these vary

more or less continuously in length with

the more obvious cover of fully developed

indumentum.

Both populations of this species contained

many plants in 2013 and no other species

of the genus were present reinforcing the

hypothesis that these plants are not the result

of any recent hybridisation event. The overall

morphology of Plectranthus bipartitus is also

hard to reconcile with a stabilised hybrid

combination from the other species that are

present. The apparent (this could be negated

with further survey) marked disjunction at

the ends of the tableland is repeated to some

extent by that of P. bellus , although this latter

species is known from more populations.

Conservation status : Two populations are

known for Plectranthus bipartitus. Both

occur in a National Park; however, as noted

for P. bellus (Forster 2011), the granite

pavement and boulder fields habitats of the

Hann Tableland are besieged by a range

of alien naturalised species, especially

feral pasture grasses such as Andropogon

gay anus Kunth, Melinis minutiflora P.Beauv,

M. repens (Willd.) Zizka, Sporobolus

pyramidalis P.Beauv.and the daisies Bidens

pilosa L. and Praxelis clematidea R.M.King

& H.Rob. An appropriate conservation status

is Endangered based upon the IUCN (2001)

criterion D.

Etymology : The specific epithet is derived

from the Latin bipartitio (division into two)

and alludes to the structures of the individual

cymes.

Plectranthus splendens P.I.Forst., sp. nov.

differing from P. apreptus S.T.Blake by being

a much more robust erect herb to subshrub

(versus a weakly decumbent herb) with

strongly scented ± firm fleshy foliage with

red sessile glands (versus scentless lax foliage

with yellow sessile glands), the leaves have

fewer teeth (5-8 per margin) that are longer

(up to 8 mm long) (versus leaves with more

teeth [7-15 per margin] that are shorter [up to

3 mm long]) and the inflorescence is initially

weakly comose with much larger (3.2-4 mm

long) bracts (versus an inflorescence that is not

weakly comose and has much smaller (1.2-1.8

mm long) bracts). Typus: Queensland. Cook

District: Hann Tableland National Park,

8 April 2013, P.I. Forster PIF39587 (holo:

BRI [2 sheets + spirit]; iso: CNS, K, MEL

distribuendi).

Erect to rarely decumbent subshrub to 100

cm high; foliage with very strong sickly sweet

scent when crushed, scabrid; non-glandular

and glandular trichomes uncoloured,

sessile glands 8-celled, dark orange. Roots

thickened fibrous. Stems square, erect to

rarely straggling, fie shy-succulent, easily

snapped without obvious stringy fibres, the

lower parts up to 8 mm diameter, pale green

to flushed purple, upper parts with persistent

indumentum, non-glandular trichomes

generally absent or rarely 6-9-celled and up

to 0.8 mm long, glandular trichomes dense,

to 0.8 mm, sessile glands absent. Leaves

discolorous, petiolate; petioles 20-45 x

2-3.5 mm, channelled on top, non-glandular

trichomes absent or scattered, divaricate

and 6-8-celled to 0.8 mm long, glandular

trichomes sparse to dense, to 0.8 mm,

sessile glands absent; laminae broadly ovate

to somewhat subcordate, rarely reniform-

cordate, ± firm fleshy, keeled when young,

flatter with age, 28-70(90) x 28—80(110) mm,

dentate with 5-8 teeth up to 8 mm long on

each margin, of similar length along margin,

secondary teeth usually present; tip acute; base

subcordate to truncate; upper surface glossy

mid-green, scabrid, veins impressed, non-

glandular trichomes sparse to dense, antrorse

to divaricate, 6-8-celled up to 1 mm long,

glandular trichomes sparse to dense, to 0.8 mm

long, sessile glands absent; lower surface pale

green to silver green (due to the indumentum)

and sometimes with a reddish hue (due to

sessile glands), scabrid, veins strongly raised,

non-glandular trichomes sparse to dense,

divaricate, 6-8-celled up to 1 mm long,

glandular trichomes sparse to dense, to 0.8

mm long, sessile glands absent, or scattered

to dense. Inflorescence up to 210 mm long,

single with 1 or 2 side branches, pedunculate

for 15-20 mm; axis square in cross-section,

non-glandular trichomes absent or a few

near vertillasters, divaricate, 6-8-celled up

to 0.8 mm long, glandular trichomes dense,

212

up to 0.8 mm long, sessile glands absent;

bracts broadly ovate to subcordate, 3.2-4

x 3.5-5 mm, loosely comose near top of

inflorescence, but soon shed, non-glandular

trichomes sparse, divaricate, 5-8-celled up

to 0.5 mm long, glandular trichomes sparse

to dense, up to 0.5 mm long, sessile glands

occasional; cymes sessile; verticillasters

10-14-flowered, up to 13 mm apart; pedicels

3.6-6 x c . 0.4 mm, non-glandular trichomes

occasional, divaricate, 4-celled up to 0.1

mm long, glandular trichomes sparse, up to

0.2 mm long, sessile glands absent. Flower

calyces 2.2-2.5 mm long, non-glandular

trichomes scattered, antrorse, 4-celled up

to 0.2 mm long, glandular trichomes sparse,

sessile glands sparse. Corolla 10-11 mm long,

purple and white to deep blue-purple; tube

5.5-6 mm long, curved at c. 90° c. 3 mm

from the base, not curved upwards, glabrous;

upper lobes suborbicular, erect, 2.2-2.5

x 2.2-2.5 mm, non-glandular trichomes

occasional, divaricate, 2-4-celled up to 0.2

mm long, glandular trichomes sparse, sessile

glands occasional; lateral lobes oblong, 2-2.2

x c. 1 mm, glabrous, purple with central

white blotch; lower lobe broadly ovate,

5.5-6 x 4-4.2 mm, non-glandular trichomes

occasional, divaricate, 2-4-celled up to 0.1

mm long, glandular trichomes occasional,

sessile glands occasional or absent; filaments

filiform, 8-9 x c. 0.1 mm, lilac, fused for c. 3

mm from the base; anthers c. 0.4 x 0.3 mm;

style filiform, 8-10 x c. 0.1 mm, lilac, bifid

for c. 0.3 mm. Fruit calyces 3-3.3 mm long;

upper lobe oblong-ovate, 1-1.2 x c. 1.5 mm;

lateral lobes lanceolate-falcate, 1.2-1.3 x

0.8-0.9 mm; lower lobes lanceolate-falcate,

1.2-1.5 x 0.5-0.6 mm. Nutlets ± circular in

outline, flattened to somewhat convex, 0.7-

0.9 x 0.8-0.9 mm, glossy brown, somewhat

tessellate. Fig. 2.

Additional specimens examined: Queensland. Cook

District: Northern end of Hann Tableland, NW of

Mareeba, Apr 2005, Wannan 3955 & Ray (BRI, NSW),

Oct 2005, Wannan 4126 & Jago (BRI); Hann Tableland

NP, May 2010, Forster PIF36941 (BRI, CNS), May

2010, Forster PIF37194 (BRI, CNS, MEL), Mar 2012,

Mathieson MTM1299 (BRI, DNA), Apr 2013, Forster

PIF39592 (BRI, CNS, MEL, NSW), Apr 2013, Forster

PIF39725 (BRI, CNS, MEL, NSW); Hann Tableland,

near radio tower (cult Tolga), May 1995, Sankowsky

1467 & Sankowsky (BRI), Jul 1995, Forster PIF17165

Austrobaileya 9(2): 207-215 (2014)

(BRI, CNS, DNA, L, MEL, NE, PE); Hann Tableland,

near Radar Station, May 2004, McDonald KRM2471

(BRI, DNA, MEL, NSW), May 2006, Forster PIF31718

& McDonald (BRI, MEL, NE, NSW); Hann Tableland,

NW of Mareeba, Oct 1973, Webb & Tracey 11649 (BRI);

Boyle Pocket, Hann Tableland, Mar 2000, Thompson

SLT2613 (BRI).

Distribution and habitat : Plectranthus

splendens is endemic to the Hann Tableland

(Map 1) where it is widespread - occurring

predominantly on areas of pavement, but

is also to be found on large boulders within

areas of woodland or open forest. It is the

most widespread Plectranthus on the Hann

Tableland after P. spectabilis and the two

species are sometimes sympatric.

Notes: Plectranthus splendens might be

related (on the basis of morphology) to the

more coastal occurring P. apreptus S.T.Blake

and some collections from the Hann Tableland

have been identified and distributed under

that name. The two species differ in a number

of characters. P. splendens is a much more

robust erect herb to subshrub with strongly

scented ± firm fleshy foliage with red sessile

glands, the leaves have fewer teeth (5-8 per

margin) that are longer (up to 8 mm long) and

the inflorescence is initially weakly comose

with much larger (3.2-4 mm long) bracts.

By comparison P. apreptus is a weakly

decumbent herb with yellow sessile glands,

scentless thin fleshy foliage, the leaves have

more teeth (7-15 per margin) that are shorter

(up to 3 mm long), the inflorescence is not

weakly comose and has much smaller (1.2-1.8

mm long) bracts.

The populations from the Hann Tableland

included within this species vary in the degree

of indumentum cover and the disposition of

the sessile glands on the foliage. Some plants

(generally throughout a whole population)

have noticeably shaggier foliage (due to

greater density of indumentum) than others.

A couple of populations have very dense

red sessile glands on the leaf undersurface

that impart a reddish colour to the surface;

however, this feature seems to be variable and

can be more or less absent.

This attractive species appears to have been

first collected by Len Webb and Geoff Tracey

in their pioneering botanical explorations of

Forster, Plectranthus from Hann Tableland

213

Fig. 2. Plectranthus splendens. A. habit of flowering stem x0.5. B. adaxial leaf surface xl. C. abaxial leaf surface *1.

D. verticillaster x4. E. floral bract x4. F. lateral view of flower x6. G. face view of flower x6. H. lateral view of flower

pedicel and calyx x8.1. lateral view of fruiting pedicel and calyx x8. All from Forster PIF39587 (BRI). Del. W. Smith.

214

Austrobaileya 9(2): 207-215 (2014)

the Hann Tableland rainforests in 1973. The

species is likely to become widespread in

cultivation as it is easy to grow in frost free

gardens and spreads vigorously by layering.

Most material already in cultivation by

native plant enthusiasts emanates from the

population near the radar station based

on a collection made by Garry and Nada

Sankowsky prior to 1995. This particular

population of plants is notable for the more

or less complete absence of red sessile glands

from the leaf undersurface.

Conservation status: Plectranthus splendens

is widespread on the Hann Tableland with

most of the known populations present in the

National Park. As noted for P. bellus (Forster

2011) and for P. bipartitus (above), the habitats

where this species occurs are besieged by alien

invasive weeds. In 2013, the populations of P.

splendens were little affected by these weed

invasions; however, it is difficult to predict

what may eventuate in the future as the aliens

expand their distribution. An appropriate

conservation coding for P. splendens is

Vulnerable, based on the criterion D2 (IUCN

2001 ).

Etymology: The specific epithet is derived

from the Latin splendens and alludes to the

attractive appearance of this plant.

Acknowledgements

Some of the cited field collections were made

on Bush Blitz (Commonwealth Government)

supported expeditions in 2010 and 2013.

Assistance with field work and plant material

was provided by Garry and Nada Sankowsky

(Tolga), Keith McDonald (Atherton), Mike

Mathieson and Megan Thomas (Queensland

Herbarium); traditional owners for the Hann

Tableland (John & Troy Grainer), National

Parks & Wildlife Service staff (Jonathon

Roth, Robert Miller). Cape York Helicopters

provided excellent logistical support.

References

Bouchard, P. & Brooks, D.R. (2004). Effect of vagility

potential on dispersal and speciation in

rainforest insects. Journal of Evolutionary

Biology 17: 994-1006.

Champion, D.C. & Bultitude, R.J. (2013). The

geochemical and Sr-Nd isotopic characteristics

of Paleozoic fractionated S-types granites of

north Queensland: Implications for S-type

granite petrogenesis. Lithos 162/163: 37-56.

Forster, PI. (2011). Five new species of Plectranthus

F.Her. (Famiaceae) from New South Wales and

Queensland. Austrobaileya 8: 387-404.

Heald, W.F. (1951). Sky islands in Arizona. Natural

History 60: 56-63.

Iucn (2001). Red List Categories and Criteria: Version

3.1. IUCN Species Survival commission. IUCN:

Gland, Switzerland/Cambridge, U.K.

Porembski, S., Becker, U. & Seine, R. (2000). Islands on

islands: habitats on inselbergs. In S. Porembski

& W. Barthlott (eds.), Inselbergs. Ecological

Studies 146: 48-67.

Watling, J.I. & Donnelly, M.A. (2006). Fragments

as islands: a synthesis of faunal responses to

habitat patchiness. Conservation Biology 20:

1016-1025.

Forster, Plectranthus from Hann Tableland

215

Map 1 . Distribution of Plectranthus species on the Hann Tableland, north-east Queensland. Outline indicates the

extent of the National Park.

Four new Queensland species of Solanum L.

allied to S. ellipticum R.Br. (Solanaceae)

A.R. Bean

Summary

Bean, A.R. (2014). Four new Queensland species of Solanum L. allied to S. ellipticum R.Br.

(Solanaceae). Austrobaileya 9(2): 216-228. Four new species of the Solanum ellipticum species group

are described, viz. Solanum adoxum A.R.Bean, S. capitaneum A.R.Bean, S. prolatum A.R.Bean and

the new combination S. chillagoense (Domin) A.R.Bean. The phenetic relationships between these

and other taxa of the S. ellipticum group are clarified by the results of a morphometric analysis, using

18 quantitative characters and 25 operational taxonomic units. The new taxa are illustrated and maps

of their distributions provided as is a map of S. ellipticum. An identification key for all Queensland

species of the S. ellipticum group is provided. Solanum dianthophorum Dunal is formally placed in

synonymy with S. ellipticum R.Br.

Key Words: Solanaceae, Solanum ellipticum, Solanum adoxum, Solanum capitaneum, Solanum

chillagoense, Solanum prolatum, Australia flora, Queensland flora, taxonomy, morphometries, new

species, identification key, conservation status

A.R. Bean, Queensland Herbarium, Department of Science, Information Technology, Innovation

and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia.

Email: Tony. Bean@dsitia.qld.gov. au

Introduction

Solanum ellipticum R.Br. has long been

regarded as a taxonomically challenging

species, encompassing a broad range of

morphological forms. Symon (1981: 193)

referred to “the cluster of species in the S.

ellipticum R.Br. complex”. Bean (2004)

included a key to 16 informal taxonomic

species-groups occurring in Australia, based

on the groupings of Whalen (1984). The

Solanum ellipticum group (Group 27), which

is endemic to Australia, is one of these. In

this key, the Solanum ellipticum group was

distinguished by having green to yellow

non-juicy fruits larger than 12 mm diameter,

monomorphic flowers, calyx prickles 10-

200 per flower, leaves with stellate hairs,

calyx not accrescent, leaves more or less

entire, and finger hairs rarely present. Bean

(2004) included ten species in the Solanum

ellipticum group, including two new species

( S. crebrispinum A.R.Bean, S. senticosum

A.R.Bean). In a more recent paper devoted

to the Solanum ellipticum group (Bean

2011), 13 species were enumerated, after the

description of four new species, reinstatement

Accepted for publication 4 August 2014

of two species, and the exclusion of three

species from the group membership.

The present paper is devoted to the

taxonomy of Solanum ellipticum sens. lat. in

tropical Queensland, the centre of diversity

for the S. ellipticum group. A morphometric

analysis of data for several species in this

group has been carried out. Morphometric

studies have proved useful in helping define

species limits in Solanum (Alvarez et al.

2008; Strickland-Constable et al. 2010). In

the current paper, a morphometric analysis,

illustrated by a dendrogram and ordination

plot, supports the recognition of four new

species ( S. adoxum, S. capitaneum, S.

chillagoense and S. prolatum ) in the S.

ellipticum group.

An interactive key (Bean 2012- onwards),

covering the Solanum species of eastern and

northern Australia, is available on the DELTA

website.

Materials and methods

This revision is based on an examination of

herbarium specimens at BRI. Measurements

of floral parts and fruits were made using

217

Bean, Solanum ellipticum

material preserved in spirit, or reconstituted

in boiling water. Terminology follows Bean

(2004). The distribution maps were compiled

using DIVA-GIS Version 7.5.0, using label

data of specimens at BRI.

Specimens of Solanum ellipticum sens,

lat. were arranged into groups (putative

taxa), based on their overall morphology. A

selection of the most complete specimens

from each group was then used for detailed

measurements, with the individual herbarium

specimens being the Operational Taxonomic

Units (OTUs).

The groups were labelled with an informal

name that refers to the locality of at least some

of the specimens, and the OTUs within each

group were numbered, i.e. Newcastle 1, 2 &

3; Hughenden 1, 2, 3 & 4; Chillagoe 1, 2 &

3; Ravenswood 1 & 2 and Edgbaston 1 &

2. Also included in the analysis were three

samples of S. ellipticum sens, str ., two of S.

callosum A.R.Bean, three of S. crebrispinum

(postulated to be the closest relative of

‘Newcastle’), and three of S. cleistogamum

Symon (a close relative of ‘Edgbaston’ and

‘Ravenswood’).

A morphometric analysis was performed

to help elucidate the phenetic relationships

between the OTUs, and to assist in defining

taxa. The analysis was done using Patn

software, Version 2.30 (Belbin 2004), using

18 quantitative characters (Table 1), and 25

OTUs. The characters chosen were considered

to be potentially useful in separating putative

taxa, and have proved useful in previous

Solanum taxonomic studies e.g. Bean (2004),

Strickland-Constable et al. (2010). The

author aimed to make 10 measurements per

character per OTU, but this was often not

possible, for example when fewer than 10

structures were available for measurement,

or the structure was absent altogether. The

mean value for each set of measurements was

then calculated for each OTU. The values for

Character 1 were recalculated using a Log

(base 10) transformation, and the values for

Character 3 were recalculated using a square

root transformation. This was done so that

these characters did not dominate the analysis,

because of their large values and large range

of values (Belbin 2004). The final matrix was

then imported into Patn.

A hierarchical classification, using

Gower’s metric association measure and

flexible UPGMA was produced (Fig. 1)

and a priori grouping was not invoked. The

ordination analysis used semi-strong hybrid

multidimensional scaling (SSH) to produce

a three-dimensional and a two-dimensional

plot of the phenetic distance between the

OTUs (Fig. 2). Various numbers of groups

were tried in preliminary analyses, but in the

final analysis, eight groups were specified.

crebrispinum3

crebrispinuml

crebrispinum2

Hughenden3

Hughenden4

Hughenden2

Hughendenl

Chillagoel

Chillagoe3

Chillagoe2

Newcastle2

Newcastle 1

NewcastleS

callosum2

callosuml

ellipticum2

ellipticuml

ellipticum3

cieistogamum2

cleistogamum 1

cleistogamum3

Ravenswoob2

Ravenswood 1

Edgbaston 1

Edgbaston2

Fig. 1 . UPGMA dendrogram generated from

agglomerative group fusion using Gower’s metric

association measure.

218

Austrobaileya 9(2): 216-228 (2014)

Table 1. Characters used in the morphometric analysis.

The Kruskal-Wallis (KW) statistic cited for each character gives an indication of the value of each

character, i.e. the higher the value, the greater is the utility of that character in discriminating

between the groups.

Character

KW value

1 .

Leaf length (mm). The largest leaves on each specimen were measured

17.633

2.

Ratio between leaf length and leaf width, for the leaves measured in

Character 1

16.428

3.

Prickles per cm of branchlet. Between two and four sections of branchlet

were chosen for measurement on each specimen

22.341

4.

Number of stellate hairs per 0.25 mm 2 of the upper leaf surface. A hair was

counted as present only if the attachment point was within the rectangular

area measured

14.096

5.

Number of prickles on the upper leaf surface. Leaves were examined under

the microscope as sometimes prickles are as short as 1 mm

19.059

6.

Stellate hair diameter (mm), upper leaf surface. Where multiple layers of

hairs were present, only the uppermost hairs were measured

22.058

7.

Ratio of length of central ray to lateral rays of stellate hair (upper leaf

surface)

15.345

8.

Longest stalk of stellate hair visible in the field of view (upper leaf surface

21.510

9.

Stellate hair diameter (mm), lower leaf surface. Where multiple layers of

hairs were present, only the uppermost hairs were measured

21.267

10.

Ratio of length of central ray and lateral rays of stellate hair (lower leaf

surface).

11.541

11.

Longest stalk of stellate hair visible in the field of view (lower leaf surface)

21.185

12.

Flower number per inflorescence. This measurement includes scars where

flowers have abscised.

20.361

13.

13. Inflorescence rachis length (mm).

19.817

14.

14. Number of prickles on the calyx.

18.625

15.

Calyx lobe length at anthesis (mm). Only flowers at or very close to anthesis

were measured, as calyx lobes lengthen as the bud/flower ages

16.177

16.

Corolla, ratio of apex length: sinus length. Large values indicate a deeply

divided “stellate” corolla, while small values indicate a pentagonal or rotate

corolla

17.585

17.

Fruiting pedicel length (mm)

11.285

18.

Peduncle length of basal fruit (mm). This is the distance from the base of

the inflorescence to the first fruit attachment or scar

17.041

Results

The PATN analysis revealed eight or nine

distinct taxon groupings in the UPGMA

dendrogram (Fig. 1), and these are largely

congruent with the putative taxon sorting

originally made. When the data were

analysed again using untransformed values

for Characters 1 and 3, the same taxon

groupings were obtained. Transforming the

values for Characters 1 and 3 affected only the

Ordination stress value, which was reduced

from 0.0899 to 0.0825.

Bean, Solanum ellipticum

Fig. 2. Two dimensional ordination of the OTUs, using

semi-strong hybrid multidimensional scaling (SSH).

The minimum spanning tree is shown, with eight groups

distinguished by different symbols.

▼ S. adoxum , O S.callosum, □ S. capitaneum,

♦ S. chillagoense, ^ S. crebrispinum,

A S. cleistogamum, • S. ellipticum, I S. prolatum.

Eight taxon groups are distinguished in

a 2-dimensional ordination, with minimum

spanning tree (Fig. 2). These groups are

accepted here as taxa at species rank.

Species rank is appropriate because each

taxon differs from all its relatives by at least

four characters, and because the phenetic

differences between the putative new species

are as great as or greater than the differences

between established species e.g. S. ellipticum

and S. cleistogamum (Fig. 1).

The ‘Edgbaston’ and ‘Ravenswood’

specimens were originally placed in separate

groups, as the two ‘Ravenswood’ specimens

differ from the two ‘Edgbaston’ in a number

of characters, e.g. the broader leaves,

the abundant glandular stellate hairs, the

somewhat larger-diameter stellate hairs, and

the numerous stellate hairs with a very long

central ray. The differences are reflected by

their strong separation on the dendrogram.

However, I believe that these morphological

differences are largely due to the juvenile

nature of the available ‘Ravenswood’

specimens. The ‘Edgbaston’ specimens are

from older, fully adult plants, lacking any

219

juvenile growth or new shoots - they do

also possess the distinctive indumentum

characters mentioned above, but at very low

frequencies, and their leaves are narrower.

Taxonomy

Solanum adoxum A.R.Bean sp. nov. affinis

S. cleistogamo sed habitu erecto usque

fuso, foliis angustioribus basibus cuneatis

praeditis, corolla profunde lobata et praesentia

in crescentia juvenili pilorum stellatorum

glandibus apicalibus instructorum, differens.

Typus: Queensland. Mitchell District: Near

Measuring Spring, Edgbaston Reserve, NE of

Aramac, 6 April 2012, A.R. Bean 31650 (holo:

BRI; iso: CANB).

Sprawling to erect, rhizomatous perennial

shrub, 0.2-0.8 m high. Branchlets green to

yellow; prickles 0-3 per cm, straight, acicular

or broad-based, 1.5-6 mm long, 5-10 times

longer than wide, with stellate hairs throughout

lower part or glabrous; stellate hairs dense,

0.3-0.5 mm diameter, stalks 0-0.1 mm long;

lateral rays 6-8, porrect; central ray 0.6-2

times as long as laterals, gland-tipped or not

gland-tipped; gland-tipped finger hairs absent

or present, type 2 hairs absent. Adult leaves

elliptical, entire, 1.8-37 cm long, 0.6-1.2

cm wide, 2.9-4.1 times longer than broad;

apex obtuse, base cuneate, oblique part 0-2

mm long, obliqueness index 0-6 percent;

petioles 0.6-1.2 cm long, 23-43% length of

lamina, prickles present or absent. Tipper leaf

surface green to grey; prickles absent; stellate

hairs distributed throughout, protostellae

absent, hair density sparse to dense, 0.1-0.4

mm apart, 0.3-07 mm across, stalks absent

or to 0.1 mm long, lateral rays 7-8, porrect;

central ray 0.5-1.0 times as long as laterals,

not gland-tipped; simple hairs absent; type 2

hairs absent. Lower leaf surface greenish to

white or grey; prickles absent; stellate hairs

sparse to very dense; 0.05-0.4 mm apart,

0.4-0.8 mm diameter, stalks absent or to 0.1

mm long; lateral rays 5-8, porrect; central ray

0.4-0.9 times as long as laterals, not gland-

tipped; simple hairs absent; type 2 hairs

absent. Inflorescence supra-axillary, cymose

(pseudo-racemose); common peduncle 1-26

mm long; rachis 1-45 mm long, prickles

present; 2-5-flowered, with all flowers

220

bisexual, 5-merous; pedicels at anthesis 7-9

mm long, same thickness throughout, prickles

absent or present. Calyx tube at anthesis 1.5-

2.5 mm long; calyx lobes at anthesis deltate,

1.2-2.1 mm long; calyx prickles 8-26 per

flower, 1-3 mm long; calyx stellae dense to

very dense, transparent to white or yellow,

0.25-0.5 mm across, stalks absent or to 0.05

mm long, lateral rays 4-8, central ray 0.7-4

times as long as laterals, gland-tipped or

not gland-tipped, gland-tipped simple hairs

present or absent, type 2 hairs absent. Corolla

purple, 7-9 mm long, deeply lobed, inner

surface glabrous; anthers 3.0-4.7 mm long;

filaments 0.2-0.3 mm long; ovary glabrous or

with type 2 hairs; functional style 5-6.5 mm

long, protruding between anthers, glabrous

or with type 2 hairs. Fruiting calyx lobes less

than or more than half length of mature fruit,

prickles 1-3.5 mm long; mature fruits 1-4

per inflorescence, globose, c. 8 mm diameter,

pedicels 11-21 mm long. Mature fruits not

seen. Juvenile leaves ovate, 3.4-4.4 cm long,

1.4-2.5 cm wide, prickles absent; stellate

hairs 0.45-0.55 mm diameter, some with

eglandular central ray 4-11 times as long as

laterals, others with glandular central ray 0.6-

1 times as long as laterals, lateral rays also

often glandular; gland-tipped simple hairs

frequent, 0.1-0.2 mm long. Fig. 3A-E.

Additional specimens examined: Queensland. Mitchell

District: 93 km E of Muttaburra on stock route through

Sumana Station, Apr 2011, Thompson MUT446 &

Edginton (AD, BRI); Edgbaston, NE of Aramac, Apr

2014, Fensham 6399 (BRI); Ravenswood Station, 20

km ENE of Aramac, May 2006, Thompson MUT312 &

Wilson (BRI); Ravenswood Station, 20 km E of Aramac,

Apr 2012, Bean 31745 (BRI).

Distribution and habitat: Solanum adoxum

is known from three localities - Ravenswood

Station; Edgbaston Reserve; and Sumana,

E of Muttaburra (Map 1). At the latter two

localities, it grows on weathered sand dunes in

association with Triodia longiceps J.M.Black.

At Ravenswood, it has been found in a cleared

area, formerly Gidgee {Acacia cambagei

R.T.Baker) woodland, on sandy soil.

Phenology: Flowers and fruits have been

recorded for April and May.

Affinities: Solanum adoxum is allied to S.

cleistogamum , but differs from it by the

Austrobaileya 9(2): 216-228 (2014)

upright to sprawling habit (prostrate for S.

cleistogamum ), the narrower leaves with

cuneate leaf bases; the deeply lobed corolla

(rotate for S. cleistogamum ); the glandular

stellate hairs and gland-tipped simple hairs

on pedicels and branchlets, at least on young

growth and juvenile plants (no glandular hairs

for S. cleistogamum ), the presence of stellate

hairs with a very long central ray (4-11 times

the laterals), at least on young growth and

juvenile plants (stellate hairs with a very long

central ray absent for S. cleistogamum).

Conservation status: Solanum adoxum is

known from just three subpopulations. The

Ravenswood subpopulation is very small

and extinction at that site is predicted due

to grazing pressure. The Edgbaston site is

within a Bush Heritage reserve, and should be

stable, but fewer than 100 plants are known

there. About six plants were observed in the

Sumana subpopulation (E.J. Thompson, pers.

comm. 2014), and the site is on a stock route.

A conservation status of Endangered is

recommended (IUCN 2001). EN Blab(ii,iv) +

2ab(ii,iv); Cl.

Etymology: From the Greek adoxos, meaning

‘without glory’ or ‘obscure’. This refers

to its rather straggly habit and the lack of

ornamental features.

Solanum capitaneum A.R.Bean sp. nov.

affinis S. crebrispino sed inflorescentiis

6-12-floris, aculeis brevioribus in ramulis,

aculeis 4-22 in calyce (adversum 50-70

in S. crebrispino) et pilis stellatis sparsis

usque moderate densis in pagina superiore

foliorum, differens. Typus: Queensland.

Cook District: 3.6 km by road from Forsayth

towards Georgetown, 3 February 2006, K.R.

McDonald KRM4857 (holo: BRI; iso: BM).

Solanum sp. (Newcastle Range D.E. Symon

4907); Henderson (2002).

Sprawling to erect, rhizomatous perennial

shrub, 0.2-0.3 m high. Branchlets yellow,

rusty or brown; prickles 4-26 per cm, straight,

acicular, 1-4 mm long, 7-11 times longer

than wide, with stellate hairs throughout

lower part; stellate hairs very dense, 1.1-1.4

mm diameter, stalks 0-0.3 mm long; lateral

rays 6-8, porrect; central ray 1-1.5 times

Bean, Solanum ellipticum

as long as laterals, not gland-tipped; type

2 hairs absent. Adult leaves ovate, entire,

8.6- 13.6 cm long, 3.1-5.3 cm wide, 2.1-2.8

times longer than broad; apex obtuse or acute,

base cuneate or obtuse, oblique part 3-8 mm

long, obliqueness index 3-6 percent; petioles

1.6- 3.4 cm long, 18-25% length of lamina,

prickles present. Upper leaf surface green;

prickles absent or present on midvein only,

0-4, straight, acicular, 2-3 mm long; stellate

hairs distributed throughout, protostellae

present, hair density sparse or moderate, 0.5-

1 mm apart, 1-1.7 mm across, stalks absent

or to 0.3 mm long, lateral rays 6-8, porrect;

central ray 0.9-1.3 times as long as laterals,

not gland-tipped; simple hairs absent; type

2 hairs absent. Lower leaf surface greenish-

white, white, or grey; prickles absent; stellate

hair density moderate or dense; 0.25-0.4 mm

apart, 1-1.7 mm diameter, stalks 0.1-0.6 mm

long; lateral rays 6-8, porrect; central ray

0.6-1.1 times as long as laterals, not gland-

tipped; simple hairs absent; type 2 hairs

absent. Inflorescence supra-axillary, cymose

(pseudo-racemose) or 2-branched; common

peduncle 16-38 mm long; rachis 45-110

mm long, prickles present; 6-12-flowered,

with all flowers bisexual, 5-merous; pedicels

at anthesis 9-11 mm long, same thickness

throughout, prickles absent or present. Calyx

tube at anthesis 2-2.5 mm long; calyx lobes

at anthesis attenuate, 1-4 mm long; calyx

prickles 4-22 per flower, 1.5-4 mm long;

calyx stellae very dense, white or transparent

or brown or rusty, 1-1.2 mm across, stalks

0-0.3 mm long, lateral rays 5-7, central ray

0.9-1.5 times as long as laterals, not gland-

tipped, simple hairs absent, type 2 hairs

absent. Corolla purple, 13-15 mm long, rotate

or shallowly lobed, inner surface glabrous;

anthers 6-6.7 mm long; filaments 0.2-0.3 mm

long; ovary with Type 2 hairs only; functional

style 7.5-9 mm long, protruding between

anthers, glabrous. Fruiting calyx lobes less

than half length of mature fruit, prickles 2-3

mm long; mature fruits 3-6 per inflorescence,

globular, c. 17 mm diameter, 2-locular;

placenta stalked, anvil-shaped; interior moist

but not juicy, pericarp 0.8-1.4 mm thick;

pedicels 12—16 mm long. Fig. 3F-J.

221

Additional specimens examined : Queensland. Cook

District: Blackdown, 2.5 km WNW of Nundah Creek

crossing, Nov 2000, Ford AF2489 (BRI); Alma-den,

undated, Bicks.n. (BRI [AQ332188]); 68.4 km by road W

of Mt Surprise, Newcastle Range, Mar 2006, McDonald

KRM4929 (BRI); Agate Creek fossicking area, Apr 2006,

McDonald KRM5207 (BRI); Newcastle Range, near

Georgetown, Jan 2001, McDonaldKRM700 (BRI); c. 32

km E of Georgetown, May 1967, Symon 4904, 4907 (AD,

BRI, CANB, K, MO); Newcastle Range, Routh Gorge,

Jan 1983, Sankowsky 242 & Sankowsky (BRI); 5.6 km

along Agate Creek Road from junction at Forsayth, Apr

2006, McDonald KRM5144 (BRI); 3 km W of Forsayth,

Apr 2003, Wannan 2808 & Graham (BRI); c. 32 km E

of Georgetown, May 1967, Symon 4906 (AD, BRI, MO);

67.3 km by road from Mt Surprise, Newcastle Range,

Apr 2006, McDonald KRM5116 (BRI); Townley, S of

Georgetown, Jul 2001, Fensham 4565 (BRI); Bagstowe,

Stuarts Spring Nature Refuge, c. 95 km S of Georgetown,

May 2011, Mathieson MTM1054 (BRI).

Distribution and habitat: Solanum

capitaneum is found mainly around

Georgetown and Forsayth, with several

records from the Newcastle Range. There is

also an old record from Alma-den (Map 1).

Most (if not all) records are from eucalypt

woodland with sandy soil on granite substrate.

Phenology : Flowers have been recorded from

November to July; mature fruits from January

to July.

Affinities: Solanum capitaneum is

distinguished within the S. ellipticum group

by its large stellate hairs, well-spaced on

the upper leaf surface, and the sometimes

branched inflorescence bearing 6-12 flowers.

It is close to S. crebrispinum , but differs by

the prickles on the branchlets < 6 mm long

(up to 11 mm long for S. crebrispinum ), the

narrower leaves, 2.1-2.8 times longer than

wide (vs. 1.5-2.1 times for S. crebrispinum ),

the upper leaf surface with sparse to

moderately dense stellate hairs (vs. dense for

S. crebrispinum ) on stalks no longer than 0.3

mm (stalks to 0.9 mm for S. crebrispinum );

the absence of prickles on the lower leaf

surface (present for S. crebrispinum ); the

6-12-flowered inflorescences (3-5-flowered

for S. crebrispinum ); flowers on pedicels 9-11

mm long (4-6 mm long for S. crebrispinum );

and the calyx prickles 4-22 per flower (50-70

per flower for S. crebrispinum ).

222

Austrobcdleya 9(2): 216-228 (2014)

Fig. 3A-E: Solanum adoxum A. adult leaf *2. B. stellate hair with short central ray *48. C. stellate hair with long

central ray *48. D. branchlet indumentum with a mixture of stellate and simple glandular hairs *48. E. flower at

anthesis x6. F-J: S. capitaneum. F. adult leaf xl. G. infmctescence xl. H. stellate hair from upper leaf surface x24.1.

corolla x2. J. flower at anthesis with corolla removed x3. A-B from Bean 31650 (BRI); C-D from Thompson MUT312

& Wilson (BRI); E from Bean 31745 (BRI); F-J from McDonaldKRM4857 (BRI). Del. W. Smith.

Bean, Solanum ellipticum

Conservation status : Although the

geographic range of Solanum capitaneum is

not large, it appears to be relatively common

within that range. A status of Least Concern

(IUCN 2001) is recommended.

Etymology : The epithet is from the Latin

capitaneus , head or chief, and refers to the

larger size of stellate hairs in this species

in comparison to most species in the S.

ellipticum group.

Solanum chillagoense (Domin) A.R.Bean

comb, et stat. nov.; S. ellipticum var.

chillagoense Domin, Biblioth. Bot. 89: 588

(1929). Type: Queensland. Cook District:

near Chillagoe, February 1910, K. Domin

s.n. [jI ter Australiense No. 8299 ] (lecto [here

designated]: PR 530913, photo at BRI).

Solanum ellipticum f. albiflora Domin,

Biblioth. Bot. 89: 588 (1929). Type:

Queensland. Cook District: near Chillagoe,

February 1910, K. Domin s.n. (holo: ?PR), n.v.

Sprawling to prostrate, rhizomatous perennial

shrub, 0.2-0.3 m high. Branchlets white,

yellow or brown; prickles 1-9 per cm, straight,

acicular, 1-8 mm long, 8-11 times longer than

wide, with stellate hairs throughout lower part

or sometimes glabrous; stellate hairs very

dense, 0.6-0.8 mm diameter, stalks 0-0.2

mm long; lateral rays 7-8, porrect; central ray

0.4-1.1 times as long as laterals, not gland-

tipped; type 2 hairs absent. Adult leaves ovate,

entire or obscurely lobed, 8.2-13.3 cm long,

3.4-4.6 cm wide, 2.2-2.9 times longer than

broad; apex acuminate or acute, base cuneate,

oblique part 2-7 mm long, obliqueness index

2-7 percent; petioles 2-3.5 cm long, 20-33%

length of lamina, prickles present. Upper

leaf surface green to grey-green; prickles

absent or present on midvein only, 0-5,

straight, acicular, 1-2 mm long; stellate hairs

distributed throughout, protostellae absent,

hair density moderate to dense, 0.25-0.35

mm apart, 0.6-0.8 mm across, stalks absent

or to 0.15 mm long, lateral rays 7-9, porrect;

central ray 0.4-1 times as long as laterals,

not gland-tipped; simple hairs absent; type

2 hairs absent. Lower leaf surface greenish-

white to white; prickles absent; stellate hairs

dense; 0.15-0.3 mm apart, 0.6-0.8 mm

223

diameter, stalks 0-0.3 mm long; lateral rays

7-8, porrect; central ray 0.5-1 times as long

as laterals, not gland-tipped; simple hairs

absent; type 2 hairs absent. Inflorescence

supra-axillary, cymose (pseudo-racemose);

common peduncle 13-34 mm long; rachis 35-

65 mm long, prickles present; 6-9-flowered,

with some flowers bisexual and some male,

5- merous; pedicels at anthesis 7-11 mm long,

same thickness throughout, prickles absent

or present. Calyx tube at anthesis 2.5-3 mm

long; calyx lobes at anthesis attenuate, 2.5-8

mm long; calyx prickles 6-21 per flower, 0.5-

3.5 mm long; calyx stellae very dense, white

or yellow, 0.5-0.7 mm across, stalks 0-0.3

mm long, lateral rays 8, central ray 0.8-1.1

times as long as laterals, not gland-tipped,

simple hairs absent, type 2 hairs absent.

Corolla mauve to purple, 8-10 mm long,

deeply lobed, inner surface glabrous; anthers

3.5- 4.5 mm long; filaments 0.3-0.8 mm long;

ovary with stellate hairs only; functional style

6- 9 mm long, protruding between anthers,

glabrous or with stellate hairs near base.

Fruiting calyx lobes less than half length of

mature fruit, prickles 1-3.5 mm long; mature

fruits 1-3 per inflorescence, oblate, c. 18 mm

diameter, 2-locular; placenta stalked, anvil¬

shaped; interior moist but not juicy, pericarp

2.5- 3 mm thick; pedicels 15-18 mm long.

Seeds pale yellow, 2.3-2.5 mm long. Fig.

4A-E.

Additional specimens examined : Queensland. Cook

District: Chillagoe Caves NP, Royal Archway section,

Mungana, Jan 2002, Forster PIF28141 et at. (BRI);

Chillagoe - Mungana Road, c. 200 m SE of Red Dome

turnoff, Nov 2000, Ford AF2487 (BRI, NSW); 12 km

S of Chillagoe, Burke Developmental Road, Dec 1991,

Gray 5370 (BRI, CNS); Dome Rock, E of Chillagoe,

Mar 2000, McDonald KRM317 (BRI); 11 miles [16.6

km] S of Chillagoe, May 1967, Symon 4874 (BRI); 14

km SSE of Chillagoe beside Burke Developmental Road,

May 2006, Wan nan 4385 & Gray (AD, BRI, NY); SE of

Chillagoe, Mar 2000, McDonald KRM338 (BRI); Metal

Hills [Chillagoe - Mungana Caves] NP, N of Chillagoe,

Feb 2007, Little & Little s.n. (BRI [AQ728603]); Burke

Developmental Road, c. 200 m SE of Red Dome turnoff,

Mungana, May 2001, Ford AF2793 (BRI, NSW);

Royal Arch Cave, Chillagoe NP, Mar 2000, McDonald

KRM337 (BRI).

Distribution and habitat : Solanum

chillagoense is known only from the Chillagoe

area of north Queensland (Map 1). All of the

occurrences (except one) are associated with

224

Austrobaileya 9(2): 216-228 (2014)

Fig. 4A-E: Solanum chillagoense. A. adult leaf *0.6. B. infructescence xl. C. flower at anthesis x3. D. style and

ovary xl2. E. outer surface of fruiting calyx *2. F-I: S. prolatum. F. adult leaf x0.6. G. inflorescence x0.8. H. flower

at anthesis x2. I. stellate hair from upper leaf surface x24. A-B from McDonald KRM338 (BRI); C from Forster

PIF28141 (BRI); D-E from McDonald KRM337 (BRI); F-H from Cumming 23517 (BRI); I from Johnson & Turpin

s.n. (BRI [AQ745940]). Del. W. Smith.

Bean, Solanum ellipticum

limestone karst, and several specimens are

from semi-evergreen vine thicket.

Phenology : Flowers have been recorded from

November to May; mature fruits in May.

Affinities : This species has been identified in

the past as Solanum ellipticum. It differs from

that species by the 6-9-flowered inflorescences

(1-5-flowered for S. ellipticum ), the distinctly

oblate fruits (globose in S. ellipticum ), the

fruiting pericarp 2.5-3 mm thick (0.6-1 mm

thick for S. ellipticum ), the glabrous inner

surface of the corolla (sparsely stellate-hairy

for S. ellipticum ), and the presence of dense

stellate hairs on the ovary (glabrous or with

Type 2 hairs for S. ellipticum). In addition, the

calyx prickles on S. chillagoense occur only

along the midribs.

Conservation status : Although the

geographic range of Solanum chillagoense is

not large, it appears to be relatively common

within that range. A status of Least Concern

(IUCN 2001) is recommended.

Solanum prolatum A.R.Bean sp. nov. affinis

S. elliptico sed aculeis multo magis in ramulis,

aculeis 30-60 in pagina superiore foliorum,

pilis stellatis in pagina superiore foliorum

radio centrali radiis lateralibus 1.5-4 plo

longiore praeditis, inflorescentiis 6-14-floris

flores omnes bisexuales gerentibus et corolla

rotata majore, differens. Typus: Queensland.

Burke District: c. 4 km N of Solway

Downs homestead, c. 90 km directly NW of

Richmond, 14 November 1999, D.C. Johnson

& G.P. Turpin s.n. (holo: BRI [AQ745940]).

Sprawling to prostrate, rhizomatous perennial

shrub, up to 0.4 m high. Branchlets white,

rusty or brown; prickles 24-98 per cm,

straight, acicular, 2-8.5 mm long, 12-16 times

longer than wide, glabrous; stellate hairs very

dense, 0.7-1.1 mm diameter, stalks 0-0.5 mm

long; lateral rays 7-8, porrect; central ray

0.8-1.3 times as long as laterals, not gland-

tipped; type 2 hairs absent. Adult leaves ovate

to broadly ovate, entire or obscurely lobed,

6.3- 11.4 cm long, 4-6.2 cm wide, 1.6-2.4

times longer than broad; apex obtuse or acute,

base obtuse to cordate, oblique part 2.5-7 mm

long, obliqueness index 3-6 percent; petioles

2.4- 4.8 cm long, 30-44% length of lamina,

225

prickles present. Upper leaf surface green

to grey-green; prickles present on midvein

and lateral veins, 30-60, straight, acicular,

2-7 mm long; stellate hairs distributed

throughout, protostellae absent, hairs dense,

0.1-0.4 mm apart, 0.6-1 mm across, stalks

0-0.5 mm long, lateral rays 7-8, porrect or

ascending; central ray 1.5-4 times as long as

laterals, not gland-tipped; simple hairs absent;

type 2 hairs absent. Lower leaf surface grey

to white; prickles present on midvein and

lateral veins, 13-23; stellate hairs dense

to very dense, 0.1-0.25 mm apart, 0.8-1.2

mm diameter, stalks 0-1 mm long; lateral

rays 7-8, porrect or ascending; central ray

1-2 times as long as laterals, not gland-

tipped; simple hairs absent; type 2 hairs

absent. Inflorescence supra-axillary, cymose

(pseudo-racemose); common peduncle 22-43

mm long; rachis 36-82 mm long, prickles

present; 6-14-flowered, all flowers bisexual,

5-merous; pedicels at anthesis 9-21 mm long,

same thickness throughout, prickles present.

Calyx tube at anthesis 1.5-3.5 mm long; calyx

lobes at anthesis attenuate, 3.5-9 mm long;

calyx prickles 36-82 per flower, 1-3.5 mm

long; calyx stellae very dense, transparent or

purplish, 0.7-1 mm across, stalks 0-0.5 mm

long, lateral rays 6-8, central ray 2-3 times

as long as laterals, not gland-tipped, simple

hairs absent, type 2 hairs absent. Corolla

purple, 16-18 mm long, rotate, inner surface

glabrous; anthers 4.6-57 mm long; filaments

0.5-1.2 mm long; ovary glabrous; functional

style 9-9.5 mm long, protruding between

anthers, glabrous. Fruiting calyx lobes with

prickles 2-3.5 mm long; pedicels 11-16 mm

long. Mature fruits not seen. Fig. 4F-I.

Additional specimens examined : Queensland. Burke

District: Sussex Park, Flinders River, Jun 1934, Blake

6258 (BRI); 16.8 km N of Kennedy Development Road

near Hughenden towards Torver Valley, Aug 2005,

Cumming 23517 (AD, BRI). Mitchell District: 35

miles [58.2 km] S of Torrens Creek, Woura Park, Jun

1971, Compton s.n. (BRI [AQ039078]); Woura Park,

56 km S of Torrens Creek, Sep 1972, Bode s.n. (BRI

[AQ005863]).

Distribution and habitat : Solanum prolatum

has a restricted distribution centred on the

town of Hughenden (Map 1). At one location it

occurs on the top of a breakaway with Acacia

cambagei ; at another it grows on a basalt slope

226

with Corymbia dallachiana (Benth.) K.D.Hill

& L.A.S. Johnson and C. terminalis (F.Muell.)

K.D.Hill &L.A.S.Johnson.

Phenology: Flowers have been recorded

between June and September. The fruiting

period is unknown.

Affinities: Solanum prolatum differs from S.

ellipticum by the many more prickles on the

branchlets, the obtuse to cordate leaf bases,

the 30-60 prickles on the upper leaf surface

(0-20 prickles for S. ellipticum ), the stellate

hairs on the upper leaf surface with a central

ray 1.5-4 times the laterals (central ray 0.5-1

times for S. ellipticum ), the 6-14 flowered

inflorescences with all flowers bisexual

(1-5 flowered with some flowers male for

S. ellipticum ), the corolla 16-18 mm long,

rotate, glabrous on the inner surface (corolla

7-12 mm long, deeply to shallowly lobed,

sparsely stellate hairy on inner surface for S.

ellipticum ).

Conservation status: Solanum prolatum

is known from just five specimens, but

from quite a wide area. It is anticipated

that more populations will be found, based

on the availability of habitat. A status of

Data Deficient (IUCN 2001) is therefore

recommended.

Etymology: From the Latin prolatus , meaning

extended or elongated. This is in reference to

the central ray of the stellate hairs, which is

longer in this species than in any other of the

S. ellipticum group.

Solanum ellipticum R.Br., Prodr. 446 (1810).

Type: Queensland. Port Curtis District:

Broadsound, 25 September 1802, R. Brown

Austrobaileya 9(2): 216-228 (2014)

(lecto: BM, fide Symon [1981: 188]; isolecto:

MPU).

Solanum dianthophorum Dunal, Hist. Nat.

Solanum 183 (1813); S. biflorum R.Br., nom.

illeg., non Lour. (1790), syn. nov. Type:

Queensland. Port Curtis District: Port II,

undated [Port Clinton, 21-23 August 1802],

R. Brown s.n. [Bennett number 2668 ] (holo:

BM).

For a description, see Bean (2011).

Distribution and habitat: Solanum

ellipticum is widespread in the eastern half

of Queensland, excluding the far north and

the high-rainfall areas of the south-east (Map

2). It grows on a diverse range of habitats and

soils, where the drainage is good and high

levels of sunlight are available.

Notes: Bean (2004) accepted Solanum

dianthophorum as a possibly distinct

species, but restricted its use to the type

specimen that was collected in what is now

the Shoalwater Bay Training Area, north of

Rockhampton. Recent biodiversity surveys

there have resulted in the collection of several

Solanum ellipticum specimens from a range

of habitats. At least one of them was collected

from littoral rainforest on sand near a beach.

This is the presumed habitat of Brown’s

collection of S. dianthophorum , described by

him as “arenosus prope littus”. The absence

of a distinct S. dianthophorum-likQ plant in

that habitat (or any other habitat), combined

with the presence of typical S. ellipticum

there, has convinced me that the type of S.

dianthophorum is merely an anomaly and that

it should be reduced to synonymy with the

widespread S. ellipticum.

Key to the Queensland species of the Solanum ellipticum group

1 Style eccentric, strongly bent near its base.2

1. Style erect, sometimes curled near apex.3

2 Branchlet hairs not floccose (stalks 0-0.2 mm); calyx prickles 0.5-2 mm

long; corolla 9-13 mm long.S. emmottii

2. Branchlet hairs floccose (stalks 0-2 mm); calyx prickles 2-4 mm long;

corolla 14-20 mm long

3 Inflorescences 1-5-flowered

3. Inflorescences 6-12-flowered

S. lithophilum

. 4

10

Bean, Solarium ellipticum 227

4 Calyx prickles more than 30.5

4. Calyx prickles 0-30.7

5 Branchlet prickles 0-10/cm; corolla inner surface sparsely hairy.S. ellipticum

5. Branchlet prickles 18-42/cm; corolla inner surface glabrous.6

6 Leaves rusty-coloured, base cuneate; calyx stellate hairs 0.4-0.6 mm

across; style and ovary glabrous; branchlet stellate hairs 0.6-0.9 mm

diameter.S. senticosum

6. Leaves white to yellowish, base obtuse to cordate; calyx stellate hairs

0.7-0.9 mm across; style and ovary with short glandular hairs; branchlet

stellate hairs 0.9-1.2 mm diameter.S. crebrispinum

7 Stellate hairs of upper leaf surface with a central ray 1.1-2 times longer

than lateral rays; corolla rotate, white to mauve.S. cleistogamum

7. Stellate hairs of upper leaf surface with a central ray 0.5-1 times longer

than lateral rays; corolla shallowly to deeply lobed, purple.8

8 Adult leaves 0.7-1.2 cm wide, 2.9-4.1 times longer than broad, lower

surface green; branchlets green.S. adoxum

8. Adult leaves 1.4-5.2 cm wide, 1.6-2.9 times longer than broad, lower

surface greenish-white to grey; branchlets grey to brown.9

9 Leaves 2.2-3.5 cm long, prickles absent from upper leaf surface; petioles

58-77% length of lamina; calyx prickles 0-4.S. unispinum

9. Leaves 3.5-14 cm long, prickles 2-20 on upper leaf surface; petioles

20-45% length of lamina; calyx prickles (2-)5-30.S. ellipticum

10 Stellate hairs of upper leaf surface with central ray 1.5-4 times longer

than laterals; corolla 16-18 mm long, rotate.S. prolatum

10 . Stellate hairs of upper leaf surface with central ray 0.4-1.3 times longer

than laterals; corolla 8-15 mm long, deeply lobed or shallowly lobed. 11

11 Stellate hairs of upper leaf surface 0.05-0.2 mm apart, centre to centre

(dense to very dense). 12

11 . Stellate hairs of upper leaf surface 0.25-1.0 mm apart, centre to centre

(sparse to dense). 13

12 Stellate hairs consistently present on inner surface of corolla; rachis

prickles present; branchlets terete; mature fruits 18-20 mm diameter . . . . S. callosum

12 . Stellate hairs consistently absent from inner surface of corolla; rachis

prickles absent; branchlets usually ridged; mature fruits 14-17 mm

diameter.S. quadriloculatum

13 Stellate hairs of upper leaf surface 1-1.7 mm across, and 0.5-1 mm apart,

centre to centre; all flowers bisexual; calyx prickles scattered; corolla

13-15 mm long, rotate or shallowly lobed; ovary with short glandular

hairs only.S. capitaneum

13 . Stellate hairs of upper leaf surface 0.6-0.8 mm across, and 0.25-0.35 mm

apart, centre to centre; some flowers bisexual and some male; calyx

prickles confined to midveins; corolla 8-10 mm long, deeply lobed;

ovary with stellate hairs.S. chillagoense

228

Acknowledgements

I thank Keith McDonald for his numerous

specimen collections, Will Smith for the

illustrations, and Peter Bostock for the Latin

diagnoses. Ailsa Holland, Yumiko Baba and

Melinda Laidlaw assisted with the use of

Patn. Ailsa Holland made helpful suggestions

on an earlier draft of this paper.

References

Alvarez, N., Peralta, I., Salas, A. & Spooner, D.M.

(2008). A morphometric study of species

boundaries of the wild potato Solarium

brevicaule complex: replicated field trials in

Argentina and Peru. Plant Systematics and

Evolution 274: 37-45.

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: 639-816.

- (2011). New and reinstated species of the

Solanum ellipticum R.Br. (Solanaceae) species

group. Austrobaileya 8: 412-430.

Austrobaileya 9 ( 2 ): 216-228 ( 2014 )

- (2012- onwards). Solanum species of eastern

and northern Australia. Version 29 th June 2013.

http://delta-intkey.com Accessed 3 April 2014.

Belbin, L. (2004). Patn for Windows, Version 2.30.

http://www. patn. com. au/default. htm Accessed

13 April 2014.

Henderson, R. J.F. (ed.) (2002). Names and Distribution

of Queensland Plants, Algae and Lichens.

Environmental Protection Agency: Brisbane.

Iucn (2001). IUCN Red List of Categories and

Criteria: Version 3.1. IUCN Species Survival

Commission: Gland (Switzerland)/Cambridge

(United Kingdom).

Strickland-Con stable, R., Schneider, H., Ansell,

S.W., Russell, S.J. & Knapp, S. (2010).

Species identity in the Solanum bahamense

species group (Solanaceae, Solanum subgenus

Leptostemonum). Taxon 59: 209-226.

Symon, D.E. (1981). A revision of the genus Solanum

in Australia. Journal of the Adelaide Botanic

Gardens 4: 1-367.

Whalen, M.D. (1984). Conspectus of species groups

in Solanum subgenus Leptostemonum. Gentes

Herbarum 12: 179-282.

Map 1. Distribution of Solanum adoxum ▲,

S. capitaneum ■, S. chillagoense A and

S. prolatum • based on BRI specimens.

Map 2. Distribution of Solanum ellipticum based on BRI

specimens.

Systematics of Tephrosia Pers. (Fabaceae: Millettiae)

in Queensland: 1. A summary of the classification of

the genus, with the recognition of two new species

allied to T varians (F.M.Bailey) C.T.White

Les Pedley

Summary

Pedley, L. (2014). Systematics of Tephrosia Pers. (Fabaceae: Millettiae) in Queensland: 1. A

summary of the classification of the genus, with the recognition of two new species allied to T.

varians (F.M.Bailey) C.T.White. Austrobaileya 9(2): 229-243. The classification and nomenclature

of Tephrosia is reviewed and the names Brissonia Desv., Tephrosia section Brissonia DC. and

Tephrosia section Recueria Benth. are lectotypified. An account of the species group surrounding T.

varians is provided with description of the new species T. delicatula Pedley and T. turpinii Pedley.

Key Words: Leguminosae, Fabaceae, Brissonia, Tephrosia, Tephrosia section Brissonia, Tephrosia

section Recueria, Tephrosia delicatula, Tephrosia turpinii, Tephrosia varians, Australia flora,

Queensland flora, taxonomy, new species, identification key, distribution maps

Les Pedley, cl- Queensland Herbarium, Department of Science, Information Technology, Innovation

and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Email: Les.Pedley@dsitia. qld.gov. au

Introduction

Tephrosia Pers. is a ‘taxonomically difficult’

genus of an estimated 300 - 400 species

(Lewis et al. 2005) distributed mainly in

the tropics and subtropics of both the Old

and New World. This figure could well be

an underestimate; however, as studies of the

genus in eastern Australia suggest that there

are up to three times as many undescribed

species as there are already described ones.

This is the first of a series of papers that

review the systematics of the genus over the

eastern part of its range in Australia.

Classification & Nomenclature

Wood (1949) gave a clear account of the

convoluted nomenclatural history of

Tephrosia. In brief, the name is conserved with

the previously published Erebinthus Mitch.

(1769), Needhamia Scop. (1777) and Reineria

Moench (1802) as nomina rejecienda. Cracca

L. (1753) is also the same as Tephrosia but the

name has been rejected in favour of Cracca

Benth. (1853). This in turn, is also conserved

but is now considered a synonym of Coursetia

Accepted for publication 3 September 2014

DC. (for example, by Estrada & Martinez

2003). Though Tephrosia was conserved at the

International Botanical Congress of 1905 {fide

Wood op. cit .), adherents of the American

Code’ continued to recognise Cracca L. until

the 1930s. Conservation merely endorsed the

decision of Candolle (1826) to adopt the name

because Linnaeus applied the name Cracca

in Flora Zeylandica (1747) differently from

his use of it in Species Plantarum (1753).

However, Weatherby (1935) opined that

“neither Tephrosia nor Cracca L. ought ever

to have been proposed”. Today conservation

of Tephrosia would certainly be accepted but

probably not that of Cracca Benth.

Persoon (1807) accepted 39 species (two

described for the first time) in Tephrosia

but did not subdivide the genus in any way.

Candolle (1826) had little doubt that the genus

would have to be divided into several genera,

but that in the state of knowledge at the time

he believed it more prudent to erect simple

sections 1 . Candolle (1825) had recognised

1 ‘ J’ai peu doute qu’il devra un jour etre divise en plu-

sieurs genres; mais dans l’etat actuel des connaissances,

j’ai era plus prudent d’en former de simple sections’.

230

74 species (18 of them with some doubt)

distributed among four sections, namely:

Brissonia DC., Craccoides DC., Reineria

Moench and Mundulea DC. The type of the

genus, T. purpurea (L.) Pers. (see Jarvis 2007),

was referred by Candolle to section Reineria ,

which therefore becomes section Tephrosia.

Brissonia was published by Necker in

Elementa Botanica (1790) as a “ species

naturalis These were given unitary

designations which resemble generic names,

but are not to be treated as such. Necker’s

publication is listed in Appendix IV of the

ICBN among Opera unitqua appressa and

the name Brissonia is therefore not validly

published. It was later validated as a generic

name by Desvaux (1814). Though his name

is illegitimate as he cited two generic names

as synonyms, it does not prevent its use as

a sectional name. Candolle used Brissonia

as a sectional name, attributing it to Necker,

but did not include in it any of the species of

Brissonia Desv. He treated these among the

‘Species non satis notae’. The sectional name

Brissonia must therefore be attributed to him

alone, not to (Desv.) DC. Lectotypification

of the names Brissonia Desv. and Tephrosia

section Brissonia DC. is essential if

classifications of the genus after Candolle

are discussed. Three of the species of sect.

Craccoides are now referred to Coursetia

DC., and the other to Lotus L. (fide Ottley

1944: 101).

Brissonia Desv., J. Bot. (Desvaux) 3: 78

(1814). Lectotype (chosen here): Tephrosia

coronillifolia Desv., loc. cit..

Tephrosia section Brissonia DC., Prodr.

2: 249 (1825). Lectotype (chosen here):

Tephrosia virginiana (L.) Pers.

Meyer (1836) made a significant, perhaps

undervalued, contribution to the classification

of the genus. He described Apodynomene

E.Mey., differing from Tephrosia in having

ovate stipules similar to its spathaceous

caducous bracts, and transverse seeds. In

it he recognised two sections: Epibrissonia

E.Mey. (three species) with barbate styles

and Epireineria E.Mey. (one species) with

Austrobaileya 9 ( 2 ): 229-243 ( 2014 )

glabrous styles. Harvey (1862) reduced the

genus to sectional rank under Tephrosia.

Bentham (1859) placed South American

species in two sections: Brissonia (three

species) and Recueria sect. nov. (four species).

They differed mainly in attributes of pods

and calyces. Candolle (1825) had previously

put two species of section Recueria in

section Reineria (= section Tephrosia ), and to

endorse such placement section Recueria is

lectotypified here.

Tephrosia section Recueria Benth., FI. Bras.

(Martius) 15(1): 47 (1859). Lectotype (chosen

here): Tephrosia leptostachya DC.

T. leptostachya is sometimes considered to

be conspecific with T. purpurea , the type of

Tephrosia.

Later Bentham (1863) considered the genus

as a whole. Since his classification greatly

influenced subsequent workers, it deserves

detailed examination. Probably following the

principles set out by him and Hooker for the

publication of Genera Plantarum (see Stevens

1996) he adopted a wide circumscription of the

genus as he did for other large legume genera

such as Desmodium Desv. and Rhynchosia

Lour. He reviewed the classification, adding

generic synonyms, and recognised three

sections: (!) Brissonia , (ii) Tephrosia (as

Reineria) and (iii) Requienia (described by

Candolle as a distinct genus). He did not

mention either section Recueria or section

Craccoides and excluded section Mundulea ,

which he had raised, rather perfunctorily,

possibly acting on a suggestion of Candolle

(1826), to generic rank (Bentham 1852). The

genera Apodynomene , Macronyx Dalziel and

Pogonostigma Boiss. were discussed briefly

but included in section Tephrosia , though he

noted the first of these might be recognised

as a subsection. As part of a large regional

work, Bentham (1864) described 24 species

all except one as new. The other, T. purpurea ,

was accepted with the addition of five

varieties all confined to Australia. He referred

all species “with the exception perhaps of T.

flammea and T. crocea to section Reineria ,

with terminal or leaf-opposed racemes or

axillary clustered pedicels, and to the large

231

Pedley, Tephrosia in Queensland: 1

subsection with subulate or small stipules,

except T. venulosa [sic, perhaps a lapsus

calami for T. reticulata Benth.] in which they

are broad and striate but not so much so as in

the S. African Apodynomenes”.

J.G. Baker (1878) treated Brissonia as a

subgenus, with T. Candida DC. as sole species,

as well as subgenus Macronyx (Dalziel) Baker

(one species) and subgenus Reineria (DC.)

Baker. In his diagnosis of section Brissonia

Candolle (1825) had “Stylus lateriter barbatus”

but referred T. Candida to section Mundulea

though noting in the description of the section

that it was exceptional in having “stylus ...

barbatus”. He (Candolle 1826) discussed its

position in the genus, although it is difficult

to understand on what grounds his decision

was made. Bentham had included Macronyx

in section Reineria. E.G. Baker (1926) divided

the genus into three: (i) Eutephrosia [= section

Tephrosia ], which included section Brissonia

and section Reineria , (ii) Pogonostigma Boiss.

(included by Bentham in section Reinera ), and

(iii) Requienia. The latter two contained two

and one species respectively. The remaining

142 species were distributed among 42 series,

none formally named.

Since Bentham and the Bakers many of

the critical studies of the genus have been

concentrated at herb. Kew with emphasis

on the flora of the former British colonies

in eastern and southern Africa. Notable

exceptions are Domin (1926), Forbes (1948)

and Wood (1949). Domin’s contribution to

the taxonomy of the Australian species is

significant, though not a monograph as it was

termed by Wood (1949). He recognised 22

species, one ( T. baueri Benth.) included by

Bentham under T. purpurea , two previously

described by him (Domin 1912) and nine

described as new, one of them ( T. affinis) with

an illegitimate name. He did not list 15 of the 23

species accepted by Bentham (1864). Forbes

revised South African species of the genus.

She described 67 species, more than a third

of them for the first time, distributed among

four unnamed sections. These sections seem

aids to identification rather than groupings of

taxonomically related species; for example,

in its protologue T. inandensis H.M.L.Forbes

was compared with T. grandiflora, which was

placed in a different section. Wood (1949)

published a comprehensive account of the

“barbistyled” species of Tephrosia in north

America. He noted that in his studies that

the New World species fell “into two rather

natural groups: those with glabrous styles and

those with bearded or barbate styles”.

In preliminary studies for the Flora of

Tropical East Africa , Gillett (1958) also

recognised that the genus included some

species with glabrous styles and others

with styles with indumentum of some sort

(“barbistyled”). This is more or less in accord

with distinctions between Candolle’s sections

Reineria and Brissonia. He also pointed out

that T. aurantiaca Harms and a few allied

species constituted a distinct group which

might be considered outside the glabri-/

barbistlyed division of the genus. The basis for

this suggestion was inter alia the reticulation

of the tertiary veins of the leaflets of the

species. Wood (1949) had already emphasised

the significance of both the indumentum of the

style and the tertiary venation of the leaflets.

Brummitt (1980) took up Gillett’s and

presumably Wood’s work and formally

recognised Tephrosia subgenus Barbistyla,

without referring either to section or subgenus

Brissonia. He did not pursue Gillett’s notes

on T. aurantiaca , failing to mention the

detailed discussion of the occurrence of

pubescent styles among African species

(Gillett 1959). Instead he concluded that “it

seems best to refer the entire T. aurantiaca

group, including T. hockingii subsp.

hirsutostyla (Dewit) Gillett to the typical

subgenus” and seems to have considered

the venation of leaflets of little significance.

He discussed the status of satellite genera,

three of which had been described after the

Bakers’ treatments. Of these he maintained

Requienia and Ptycolobium Harms as distinct

but believed that Caulocarpus E.G.Baker and

Linophyllum Hutch, were not different from

Tephrosia. Bosman & de Haas (1983) rejected

Brummitt’s subgenera on the grounds of

a lack of correlation between penicillate

stigmas and indumentum of styles, and

“furthermore it appeared to be impossible

232

to correlate the pubescence of the style

with any other character, except that that

the barbistyled species have larger flowers

...” Of the 20 species treated by them only

the four introduced species are barbistyled.

They foreshadowed the treatment of Geesink

(1984) who admitted to only a limited

knowledge of Tephrosia. He recognised all

four genera discussed by Brummitt. Schrire

(1987) supported Brummitt’s division of the

genus and added significantly to definitions of

the subgenera. Brummitt (2007) presented a

key to 70 species; species 1-30 were referred

to subgenus Tephrosia , species 32-70 to

subgenus Barhistyla. The affinities of species

31, T. miranda Brummitt (known from only

two specimens), were considered obscure. It

has unusual pods but he suggested it might be

related to T. zoutpansbergensis Bremek. and

T. villosa (L.) Pers. despite its pubescent style.

Both Brummitt (2007) and Schrire (1987)

placed T. Candida in T. subgenus Barbistlya.

Unless the subgenus is defined more narrowly

to exclude T. Candida , it should be referred to

T. subgenus Brissonia (DC.) Baker.

The position of Mundulea (DC.) Benth.

vis-a-vis Tephrosia and Millettia Wight &

Arn. has been disputed. Geesink (1984) noted

it to be scarcely distinct from Millettia. Schrire

(1990) when discussing Tephrosiapondoensis

(Codd) Schrire noted the differences between

Tephrosia and Mundulea. He more or less

dismissed the diagnostic value of attributes

of leaf venation and dehiscence of pods,

but stated that floral characters have proved

more consistent in separating the two. Later

some species of Mundulea were referred to

the newly described Pyranthus Du Puy &

Labat and Sylvichadsia Labat & Du Puy by

Du Puy & Labat (1995) and Labat & Du Puy

(1998) respectively. Both genera appear to be

more closely related to another Madagascan

genus Chadsia Bojer than to either Tephrosia

or Millettia. Du Puy & Labat (2002) treated

Mundulea as a distinct genus, but combined

the key to its species with the key to those

of Tephrosia ; mainly because, contrary to

the opinion of Schrire, attributes of their

pods provided the most reliable differential

characters and pods were often not available.

Verdcourt (2007) iterated Bentham (1863) in

Austrobaileya 9(2): 229-243 (2014)

noting that Mundulea “links Tephrosia with

Millettia , but to combine it with either would

lead to problems of definition. Certainly

the three cannot be united, but Mundulea

could perhaps be considered a subgenus

of Tephrosia ”. I have examined only a few

specimens of M. sericea (Willd.) A.Chev.

the most widespread species of the genus;

its calyx lobes are unlike those of any other

species, African, Asian, north American or

Australian I have seen but its general facies

would place it in Tephrosia. Its dilated

staminal filaments seem a minor attribute in

distinguishing genera.

Differing opinions on the circumscription

and status of Tephrosia subgenus Barbistyla

seem to have inhibited the evaluation of

characters other than stylar indumentum that

might be significant for the classification of

Tephrosia as a whole. Many workers have

commented on the venation (or nervature) of

the leaflets that they considered characteristic

of the genus. For example, Forbes (1948):

“There is one constant and conspicuous

character by which the genus maybe readily

recognised, the close distinct penninerved

venation of the leaflets”; Bosman & de Haas

(1983): “The intersecondary nerves are

relatively numerous, parallel, rather straight

(never S-shaped) and curved slightly upwards

to the margin in which they usually end”;

Geesink (1984): “the nerves are straight,

forming a sharp angle (usually 10°-30°)

with the midrib”. Wood (1949: 210) and

Gillett (1958) recognised minor variants of

this pattern; Wood: “More or less parallel

lateral veins are given off obliquely from

the midrib of the leaflet in all species. The

areoles formed by the veinlets between these

veins are,... either elongate or isodiametric”;

Gillett: “T aurantiaca s. lat. is undoubtedly

a very natural group, distinguished by ...

rather large leaflets... with rather wide spaces

between the chief lateral nerves and a very

prominent network of venules connecting

them”. Rudd (1991), in a key to genera

distinguished Tephrosia from Mundulea and

Millettia by its “numerous straight, closely

parallel lateral veins mostly extending to the

margin”.

233

Pedley, Tephrosia in Queensland: 1

Cowie (2004) drew attention to the

differences and diversity of Australian

species. In defining four groups of Australian

species differing in their venation patterns,

he upset established views. His first group,

with T. phaeosperma F.Muell. ex Benth. as an

example, corresponds to the accepted ‘typical’

pattern ; the second and third are difficult to

place; the fourth is distinctive. It consists of

“species with well spaced secondary veins

curving or dichotomising before the margin,

the intersecondary venation often closely

reticulate, prominent or not”. Bentham (1864)

seems to have recognised this group indirectly

where, in his key to species, the first five have

“veins anastomosing or reticulate within the

margin”. It possibly requires the recognition

of a genus distinct from Tephrosia and studies

of Queensland species have strengthened my

opinion that this is warranted.

Since Tephrosia reticulata Benth. and

related species in the Northern Territory

require further study, which I cannot

undertake, I have retained the three

reticulately veined species treated below in

Tephrosia. In the protologue of T. oxalidea

R.Butcher & P.J.H.Hurter, Butcher & Hurter

(2012) provided detailed description of the

venation of a reticulately veined species.

Their somewhat diagrammatic illustrations

of leaflets of T. coriacea Benth. (Fig. 1A) and

T. rosea F.Muell. ex Benth. (Fig. ID) contrast

the reticulate venation of the former with the

‘typical’ venation of the latter.

In the relaxed atmosphere of the final

session of the 16th Botanical Congress in St

Louis in 1999, I suggested to some friends

from herb. Kew that about 12 Australian

species from northern Australia characterised

by leaves with reticulate veins might be

segregated as a separate genus (quoted in

Lewis et al. 2005).

Except for indumentum of the style,

attributes of inflorescences and flowers

have been used in only a rather indefinite

way to delimit infrageneric taxa in

Tephrosia. Limited study of living and

herbarium specimens; however, suggests

that Apodynomene might be recognised as a

distinct genus. In practice it has been treated

as a coherent group of species by a succession

of authors. Forbes (1948) placed most of the

species in her ‘section 4’, a notable exception

being T. inandensis in ‘section 3’, though,

in its protologue, she compared it with T.

grandiflora (L’Her. ex Ait.) Pers. (section

4), a species that is naturalised in the West

Indies. Wood (1949: 374) noted that it is easily

recognised: “The large, promptly deciduous

bracts and broad stipules were largely the

basis of the segregate genus Apodynomene

E.Mey., of which this is the type-species.

Although stipules and bracts of this type

characterize a group of South African species,

the absence of secondary bracts is indeed

anomalous, there seem to be no real reasons

for separating this group as a distinct genus”.

He considered Harvey’s (1862) treatment of

the group as a section might prove to be a more

reasonable disposition. It should be noted that

Wood effectively selected T. grandiflora as

generitype of Apodynomene. Schrire (1987)

in his notes on T. subgenus Barbistyla ,

commented: “within the subgenus Barbistyla ,

inflorescence characters (bracts) and fruit

characters (pod size and seed position)

separate the T. grandiflora allies from those

of T. longipes [the type of the name T. subg.

Barbistyla ]. Inflorescence specialisation has

been particularly marked in the T. grandiflora

alliance”. He keyed 11 species of this alliance

more or less together (species No. 11-21)

though he did not use absence of secondary

bracts as a key character.

It might be expected that molecular

analysis could suggest a plausible scheme

for generic and infrageneric classification of

Tephrosia and its allies. LPWG (2013) gave

a comprehensive account of the advances

and deficiencies in the ‘building a high-

resolution molecular phylogeny of legumes’.

For an antipodean taxonomist it was not at

all encouraging. Among the desiderata was

“we must prioritise species-level phylogenetic

investigations of known non-phylogenetic

genera”. How is one to know whether Tephrosia

is phylogenetic or not? In the phylogram of

Kajita et al. (2001) T. heckmanniana Harms

and T. grandiflora formed a clade sister to one

consisting of Mundulea sericea and Chadsia

versicolor Bojer. da Silva et al. (2012), in a

234

study concentrated on Lonchocarpus Bojer,

confirmed the monophyly of Tephrosia ,

presumably using Candolle’s (1825) or

Bentham’s (1863) classification. Seven species

were used in the study (not all shown in the

published phylogram), all with ‘typical’

venation. A detailed molecular analysis of

the genus, including species with ‘atypical’

venation, and purportedly related genera is

necessary.

I have no doubt that Candolle (1826)

was correct in his assessment that one day

Tephrosia will need to be divided into several

genera, but also agree with Cowie (2004) that

“there is a need for further investigation of

generic limits and infrageneric subdivisions”.

Materials and methods

The species treated here are characterised by

the reticulate venation of their leaflets. They

fall into the fourth group of species defined

by Cowie (2004) though the secondary

and tertiary veins of their leaflets are more

prominent than those of the species described

by him. Only one of them ( T. gyropoda

Cowie) occurs in Queensland, near the

Northern Territory border, some 600 km west

of the nearest collecting locality of any of the

other three species covered here. To some

extent it resembles T. delicatula , but differs in

the often cuneate bases of its leaflets which

are sparsely pubescent on both surfaces and,

above all, by its arillate seeds.

In this and future papers in the series,

only the primary bract (the one subtending

Austrobaileya 9(2): 229-243 (2014)

a fascicle of flowers in inflorescences) is

described. Secondary bracts occur at the

base of each flower of fascicles of all species

examined, though sometimes they are early

deciduous. Inflorescences of the type found in

Apodynomene seem not to occur.

In all descriptions some modifications

of customary terms have been adopted.

Describing the architecture of leaves I have

followed the terminology of Butcher & Hurter

(2012, see particularly their fig. 1) with the

addition of the term ‘leaf axis’, which denotes

the length of petiole + rachis. Length of the

leaf axis in descriptions removes uncertainty

as to whether or not ‘leaf-length’ includes the

length of the terminal leaflet. It does not. The

calyx is 5-lobed, the lobes usually unequal,

the upper pair (vexillary) united to some

extent, the lowermost (carinal) often longer

than the others. In most Australian species the

calyx is therefore distinctly bilabiate. I have

considered the two vexillary lobes as one,

divided to some degree. The rather indefinite

terms ‘notched’, ‘split’ and ‘divided’ are

used to note the type of division. I have

introduced the term ‘knob’ to describe the

protuberances that usually occur on the basal

part of the staminal sheath and adjacent part

of the basally free anther-filament. These are

sometimes called calluses or callosities but I

have restricted the term callus {pi. calli) to the

thickening of the basal part of the standard.

The calli and knobs probably determine the

attitude of the flower at anthesis and have

roles in its pollination.

Taxonomy

Key to species allied to Tephrosia varians from eastern Queensland

1 Leaves with (9—)11—15 leaflets, each (12-)16-24 mm x 8—12(—14) mm,

(1-)1.5-2.2 times longer than wide; pedicels 10-16 mm long; flowers

8-10 mm long, apricot-orange; keel petals glabrous.T. delicatula

1. Leaves with (3-)5-7(-9) leaflets, each 25-60 mm x 6-25 mm, 1.3-4(-7)

times longer than wide; pedicels 3-15 mm long; flowers 8-20 mm long,

orange or yellow; keel petals pubescent or glabrous (or a few hairs on

lower margin).2

235

Pedley, Tephrosia in Queensland: 1

2 Flowers yellow, 8-11 mm long; keel petals with loosely appressed hairs

on lower margins or in proximal or distal half of lamina; pedicels 3-6

mm long; pods 4-6 mm wide.T. varians

2. Flowers apricot or orange, 15-20 mm long; keel petals glabrous or with

a few appressed hairs on lower margins; pedicels 5-15 mm long; pods

6-10 mm wide.T. turpinii

Tephrosia varians (F.M.Bailey) C.T.White,

Proc. Roy. Soc. Queensland 53: 214 (1942);

Galactia varians F.M.Bailey, Bot. Bull. Dept.

Agric. Queensland 10: 22 (1895); Queensl. FI.

2: 430, t. 14 (1900). Type: Queensland. Cook

District: Coolgarra [17°34'S 145°12'E], April

1895, M.Butler s.n. (holo: BRI [AQ22860]).

Tephrosia lutea F.Muell., Fragm. 5: 9 (1865)

nom. invalidum & 9: 64 (1875) pro syn.

Sprawling (sub)shrub to c. 40 cm tall: annual

stems from perennial carrot-like taproot.

Branchlets at base of plant terete often with

scattered ± spreading hairs to 0.5 mm long,

becoming fluted, glabrous in upper part;

stipules linear to narrow-triangular, indurated,

3-6 mm long, 1-8 veins depending on width;

young growth bronzed. Leaves with 5-7(-9)

leaflets; axis 70-90(-140) mm long, petiole

3-7.5(-9) cm long, interjugal rachis 15—30(—

38) mm long, ultrajugal rachis (5-)10-25(-28)

mm long; leaflets somewhat discolorous,

oblong, ovate, occasionally obovate, (25-

)30-60(-70) mm long, 6-25(-40) mm, rarely

more, wide, 1.3-4 times longer than wide,

rounded, sometimes truncate or emarginate

at tip, cordate or occasionally rounded at

base, glabrous except rarely a few hairs on

margins or along midrib beneath; petiolules

glabrous or with a few hairs, 1-2 mm long.

Inflorescence terminal or leaf-opposed, 30-50

cm or more long, sparsely flowered in distal

half; fascicles of 2 or 3 flowers, usually with

an additional bud; subtending bract, to 2 mm

long deciduous when only about as long as

developing fascicles; pedicels 3-6 mm long,

glabrous or sparsely appressed pubescent.

Flowers 8-11 mm long, yellow, occasionally

with red suffusion at base of standard; calyx

glabrous to moderately dense pubescent, hairs

short, appressed; tube 2-2.5 mm long, upper

lobe wide-triangular or rarely rounded, 0.8-

1.5 mm long, notched 0.1-0.3 mm, lateral lobes

triangular l-2(-2.4) mm long, sometimes

curved when long, lower usually longer than

laterals l-2.3(-3.2) mm long, sometimes

acuminate. Corolla: standard concave,

hemispherical or reniform, emarginate,

5-8 mm long, 7-12 mm wide, usually with

small thick prominent calli, claw (l-)2-2.5

mm long; wing petals longer than keel, often

markedly so, 5.5-8.5 mm long, (2.5-)3-5

mm wide, auriculate, claw 2.2-3(-3.5) mm

long; keel 4-5(-6) mm long, 2-2.7(-3) mm

wide, pubescent with loosely appressed hairs

along lower margin or in proximal half, claw

2-2.5(-3) mm long. Staminal sheath glabrous,

knobs absent or poorly developed, anthers

0.4-0.6(-0.8) mm long; ovary pubescent with

usually sparse appressed hairs, style flat and

glabrous, geniculate at tip (secus Lee 1948),

stigmatic surface inside. Pods oblong, flat,

straight or slightly decurved, 45-60 mm long,

4-6 mm wide; valves cartilaginous, glabrous

or sparsely appressed pubescent, up to 10-12

seeds, c. 4 mm between their centres, spongy

tissue between them. Seeds (few seen mature)

spherical, olive-brown, 3.6-3.8 mm diam.;

small rim aril. Fig. 1.

Additional selected specimens (all BRI): Queensland.

Cook District: Cairns, Dec 1941, Blake 14508 ; c. 5 km

SSW of Beagle North Camp, c. 3.7 km SSW of Weipa, Dec

1981, Clarkson 4177 ; 2 km S of the Big Coleman River

on the Coen to Musgrave Road, May 1987, Clarkson

7118 & Simon. ; 30 km N of Wenlock River crossing on

Peninsula Development Road, Aug 1987, Clarkson 7326\

10 km S of Wenlock River on Peninsula Development

Road, Apr 1990, Clarkson 8503 & Neldner ; 7 miles [11

km] N of Moreton Telegraph Station, Jul 1968, Gittins

1826\ Whitewater Station, near boundary of Undara

NP, Dec 2004, McDonald KRM3230 ; Undara NP, E of

Mount Surprise, Jan 2005, McDonaldKRM3314\ Undara

NP, E of Barkers Knob, Dec 2005, McDonaldKRM4684-

junction of Herberton - Petford - Irvinebank Roads, Feb

2006, McDonald KRM4485 ; 17.2 km W of Irvinebank,

Mar 2007, McDonald KRM6176 ; near Emu Creek

Station, 4.1 km E of road bridge, Jan 2008, McDonald

KRM7118; SE of Mareeba, Jan 1982, Pedley 4828 (+K,

MEL); c. 28 km SSE of Laura, Jun 2006, Wannan 4684

& Ray ; E of Musgrave on road to Marina Plains Station,

Mar 2007, Wannan 4684 & Beasley. North Kennedy

District: c. 0.5 km E of Button Rock, Mount Zero

236

Austrobaileya 9(2): 229-243 (2014)

Fig. 1. Tephrosia varians. A. twig with inflorescence x0.4. B. trifoliolate leaf (leaflets unusually large) ><0.4. C. leaflet,

showing venation xl. D. calyx, outer surface x6. E. standard x4. F. keel petal x6. G. wing petal x6. H. pod (slightly

immature) x2. I. seed x8. A from McDonald KRM4685 (BRI); B from McDonald KRM10308 (BRI); C-H from

McDonaldKRM8682 (BRI); I from Wannan 4449 (BRI). Del. W. Smith.

237

Pedley, Tephrosia in Queensland: 1

property, c. 80 km W of Townsville, Jan 2007, Camming

24593 ; Stuarts River [near Townsville], in 1891, Johnson

s.n. (+MEL 582909 & 582910); Herberton, Jan 1912,

Kenny s.n. (AQ238666); Stuart, near Townsville, Nov

1940, Watts s.n. (AQ238661).

Distribution and habitat : The species

ranges from the northern part of Cape York

Peninsula to about Townsville and inland

to about Kidston (19°S, 144°E) (Map 1). It

occurs in eucalypt woodland on well drained

soils, mainly granitic sands in the south and

red earths in the north.

Notes: One specimen ( Clarkson 8503 &

Neldner ) from south of the Wenlock River on

the Peninsula Development Road and about

20 km from the collecting locality of Gittins

1826 (cited above) is unusual. It is unifoliolate

with longer petiolules and has shorter pods

with more crowded seeds.

Mueller (1865) distinguished a species,

evidently close to Tephrosia reticulata Benth,

from the shores of Rockingham Bay. He

stated that, if it differed, except for its ‘perfect

glabrosity’ [my translation of the Latin], was

to be called T. lutea. The name is not validly

published as it is clearly a provisional one.

Later (1875) he referred the species to T.

reticulata.

Tephrosia turpinii Pedley, sp. nov. affinis

T. varianti a qua floribus grandioribus

aurantiis vel armeniacis non luteis petalis

carinae plerumque glabris vel interdum pilis

paucis secus marginem inferum praeditis et

leguminibus latioribus paucioriseminalibus

differt. Typus: Queensland. Cook District:

near Emu Creek Station, 4.1 km by road E

of Emu Creek bridge, 11 January 2008, K.R.

McDonald KRM7119 & A. Ford (holo: BRI;

iso: DNA).

Tephrosia sp. (Petford J.R. Clarkson 2774A);

Holland & Pedley in Bostock & Holland

( 2010 ).

Similar in size and habit to T. varians.

Young stems slightly angular becoming

terete, indumentum of sparse appressed

hairs 0.1-0.3 mm long, glabrescent; stipules

linear to triangular, 3-6 mm long with

up to 5 veins when wide. Leaves of (3-

)5-7 leaflets, rarely extremely minutely

stipellate; axis (5.5-)7-12.5(-14) cm long,

petiole 3-7(-8) cm long, interjugal rachis

15-35(-45) mm long, ultrajugal rachis 4-20

mm long, rarely terminal leaflets digitate;

leaflets slightly discolorous, oblong, elliptic

or ovate, 25-55(-65) mm long, 8-22(-25)

mm wide, (1.3-)2-3.5(-4) times longer than

wide, rounded or somewhat truncate at tip,

minutely mucronulate, rounded, often slightly

emarginate or subcordate at base, glabrous

above, glabrous or with extremely sparse

short hairs beneath, petiolules 1-2 mm long.

Inflorescence terminal or leaf opposed,

10-25(-35) cm long, open; fascicles of 2 or

3 flowers, subtending bract ovate, deciduous

when c. 1 mm long, pedicel 5-15 mm long,

sparse to dense appressed hairs. Flower 15-20

mm long, apricot or orange; calyx sparse to

moderately dense appressed hairs; tube (2.2-)

2.5 mm long, upper lobe wide triangular

(1.5—)2—3 mm long, usually notched 0.3-0.7

mm, occasionally sinus between lobes wide,

lateral lobes (1.5—)2—3 mm long, lower usually

longer, (1.5-)2-4.5 mm long, all tending to

be acuminate when long. Corolla: standard

broadly transversely obovate or ± circular, (9-

)11—16 mm long, (11—)12.5—16(—20) mm wide,

thickened at base or with small thick calli,

claw 2-3(-4) mm long; wing petals as long

as or slightly longer than keel, (8—)12—15 mm

long, 4.5-7.5 mm wide, not or only slightly

auriculate, claw 2-3.5 mm wide; keel petals

8-12 mm long, 4-5(-6) mm wide, glabrous or

occasionally sparse appressed hairs on lower

margin; staminal sheath glabrous, knobs

absent or poorly developed, anthers 0.8—1(—

1.2) mm long. Ovary with dense appressed

hairs; style glabrous, geniculate at tip,

stigmatic surface inside angle occasionally

with a few short hairs. Pods flat, straight,

oblong, (40-)55-60 mm long, 6-10 mm

wide, with up to 6 or 7 seeds, (4-)5.5-7 mm

between their centres with spongy tissue

between them; valves coriaceous, glabrous

or with a few scattered short appressed hairs.

Seeds flat, oblong in outline, 4.5-5 x 4-4.5

mm, small rim aril. Fig. 2.

Additional selected specimens (all BRI): Queensland.

Cook District: 16 km W of North Kennedy River

on road from Fairview to Kimba Station, Dec 1981,

Clarkson 4205 ; Burke Development Road, c. 11 km W

of Petford, Feb 1981, Clarkson 2774A ; Morgan’s Folly,

238

Austrobcdleya 9(2): 229-243 (2014)

Fig. 2. Teplirosia turpinii. A. twig with inflorescence x0.8. B. leaflet, showing venation xl. C. calyx, outer surface

x4. D. standard x3. E. keel petal x4. F. wing petal x4. G. pod (slightly immature) x2. H. seed x8. A from Wannan 1945

(BRI); B from McDonaldKRM7119 (BRI); C-F from FordAF2498 (BRI); G from McDonaldKRM3257 (BRI); H from

Wannan 5259 (BRI). Del. W. Smith.

239

Pedley, Tephrosia in Queensland: 1

38 km along road to Blackdown Station, off Chillagoe

to Wrotham Park road, Feb 1994, Forster PIF14754

(+DNA); Chillagoe, Jan 1931, Hubbard & Winders 6759

(+K; cited by White (1942) as T. various ); Emu Creek

Station between Emu Creek and Walsh River, Nov 2005,

McDonald KRM4614 et al. (+DNA, NSW); 3.7 km by

road W of Almaden, Dec 2005, McDonald KRM4649

(+MEL); 64.4 kmN of Laura near Weiss Creek, Nov 2010,

McDonald KRM10086 (+DNA, MEL); Lappa, Janl982,

Pedley 4838A\ DPI experimental area, Kalinga Station,

Jan 1976, Staples IBS2241 (distributed as T. various );

Tate River, Feb 1938, Straughan s.n. [AQ238667]; cited

by White (1942) as T. varians).

Distribution and habitat : Tephrosia turpinii

ranges from the northern part of Cape York

Peninsula to 18°S latitude a little west of

the Great Dividing Range (Map 1). Unlike

T. varians it has not been collected in the

Townsville area. It occurs in similar habitats

to those of T. varians ; that is: in eucalypt

woodland on well drained soils mainly on

granitic sands in the south and red earths in

the north. The two species occasionally grow

together.

Etymology : The species is named for my

friend and colleague Mr G.P. (‘Gerry’) Turpin,

an elder of the Mbabarum people whose land

includes that part of the distribution range

where the species commonly occurs.

Tephrosia delicatula Pedley, sp. nov. affinis

T. varianti a qua foliis foliolis pluribus

minoribus petiolulis longioribus praeditis

pedicellis longioribus floribus grandioribus

aurantiacis non luteis carinae petalis ±glabris

leguminibus longioribus angustoribusque

differt. Typus: Queensland. Cook District:

6.8 km along Battle Camp Road from West

Mclvor Road junction, 8 January 2008, K.R.

McDonald KRM7106 (holo: BRI; iso: CNS,

DNA, K, MEL distribuendi ).

Twiggy multistemmed shrub to 50 cm

tall, annual stems from perennial parsnip¬

like taproot; young stems reddish, terete,

glabrous; young growth bronzed; stipules

linear subulate to c. 6 mm long. Leaves with

(9—)11—15 leaflets; axis 90-125 mm long,

petiole 15-30(-35) mm long, interjugal rachis

(8—)12—15(—18) mm long, ultrajugal rachis

(2-)5-10 mm long; leaflets oblong elliptic or

somewhat obovate, the terminal tending to

be smaller than the laterals, (12-)16-24 mm

long, 8—12(—14) mm wide, (l-)1.5-2.2(-2.4)

times longer than wide, rounded, minutely

mucronulate at tip, obtuse at base, slightly

discolorous, glabrous or with a few scattered

appressed hairs beneath, veins raised on both

surfaces; secondary veins 8-15 on each side of

midrib; petiolules 2-3 mm long. Inflorescence

terminal, to c. 30 cm long, fascicles sparsely

arranged in distal half, occasionally a fascicle

in axil of subtending leaf; fascicle 3-flowered,

subtending bract deciduous well before

anthesis, 0.5-2 mm long, pedicel 10-16 mm

long, glabrous or with scattered appressed

hairs. Flowers 8-10 mm long, apricot-orange;

calyx with sparse short appressed hairs,

tube 2-2.5 mm long, upper lobe obtuse or

occasionally acuminate, 1-1.6 mm long,

notched for c. 0.2 mm, lateral lobes and lower

lobe triangular, occasionally acuminate,

laterals 1-1.6 mm long, lower 1.5-3 mm.

Corolla: standard ± square, emarginate, 7-10

mm long, 10-11 mm wide, only thickened at

base or calli well defined, claw wide, 1-1.3 mm

long; wing petals longer and wider than keel,

7.5-9 mm long, 3.5-4.5 mm wide, shortly

auriculate, claw 2-3 mm long; keel petals

5-6 mm long, 2.8-3.5 mm wide, glabrous,

claw 2-3 mm long. Staminal sheath glabrous,

knobs moderately developed, anthers 0.5-0.6

mm long. Ovary with dense appressed hairs;

style flat and glabrous, geniculate at tip with

stigmatic surface in angle. Pods (dehisced)

straight, sparse appressed hairs, c. 50 mm

long, 3-3.5 mm wide. Seeds spherical or

almost so (only 3 seen mature), pale khaki in

colour, 3.3-3.4 x 2.5-3.3 mm. Fig. 3.

Additional specimens examined (all BRI, duplicates

to be distributed as indicated): Queensland. Cook

District: Battle Camp Road, Jan 2008, McDonald

KRM7112 (CANB, DNA, K, MEL, NSW), Apr 2008,

McDonald KRM7219 & Covacevich.

Distribution and habitat : Tephrosia

delicatula is quite common in the vicinity

of the type locality some 35 km NW of

Cooktown (Map 1), where it is found in

eucalypt woodland dominated by Eucalyptus

phoenicea F.Muell.or E. tetrodonta F.Muell.

and Corymbia nesophila (Blakely) K.D.Hill

& L.A.S.Johnson on shallow soil on low

sandstone hills.

240

Austrobaileya 9(2): 229-243 (2014)

Fig. 3. Tephrosia delicatula. A. twig with inflorescence x0.5. B. leaf *0.8. C. leaflet, showing venation x2. D. calyx,

outer surface *6. E. standard x4. F. keel petal x6. G. wing petal x6. H. pod x2. I. seed x8. A, D-G from McDonald

KRM7106 (BRI); B & C from McDonaldKRM7112 (BRI); H & I from McDonald KRM7219 (BRI). Del. W. Smith.

241

Pedley, Tephrosia in Queensland: 1

Notes: A specimen ( Bean 5564 & Forster

[BRI]) collected some 10 km west of the type

locality and another (, S.L.Thompson ST13056

[BRI]), c. 240 km west of it, possibly represent

hybrids with Tephrosia varians or T. turpinii.

They have lanceolate leaflets longer than

those of T. delicatula though their venation

patterns are similar.

Etymology : The specific epithet is from the

Latin delicata and -ula and alludes to the

smaller leaflets with finer venation when

compared to those of both T. varians and T.

turpinii.

Acknowledgements

I wish to thank W. Smith for preparing the

figures and map. Over a long period of time

Tve been able to access herbarium collections

both in Australia and overseas, particularly at

Kew; and I wish to thank the relevant staff

for this.

References

Baker, E.G. (1926). The Leguminosae of Tropical

Africa. Erasmus Press: Gent.

Baker, J.G. (1876). Leguminosae. In J.D. Hooker, Flora

of British India 2: 56-306. L. Reeve & Co.:

London.

Bentham, G. (1852). Leguminosae. In F.A.W. Miquel,

Plantae Junghuhnianae 205-257. H R. de

Breuk/J.B. Bailliere: Lugduni-Batavorum/

Parisiis.

- (1859). Tephrosia. In C.F.P. von Martius,

Flora Brasiliensis 15(1): 45-49. R. Oldenburg:

Lipsiae.

- (1863). Leguminosae. In G. Bentham & J.D.

Hooker, Genera Plant arum 2: 434-600. L.

Reeve & Co.: London.

- (1864). Tephrosia. In Flora Australiensis 2:

202-211. L. Reeve & Co.: London.

Bosman, M.T.M. & De Haas, A.J.P (1983). A revision

of Tephrosia (Leguminosae-Papilionoideae) in

Malesia. Blumea 28: 421-487.

Brummitt, R.K. (1980). Reconsideration of the genera

Ptycholobium , Caulinocarpus , Lupinophyllum

and Requinea in relation to Tephrosia

(Leguminosae-Papilionoideae). Kew Bulletin

35: 459-473.

- (2007). Tephrosia. In Flora Zambesiaca

3(3): 119-204. Flora Zambesiaca Managing

Committe: London.

Butcher, R. & Hurter, P. J.H. (2012). Tephrosia oxalidea

(Fabaceae: Millettieae), a new species from the

Pilbara and Gasgoyne bioregions of Western

Australia. Nuytsia 22: 341-349.

Candolle, A.P. de (1825). Leguminosae. In Prodromus

systematis naturalis regni vegetabilis 2: 93-

524. Treuttel et Wtirtz: Paris.

- (1826). § Du genera Galega et du Tephrosia.

Memoires sur la famille des Legumineuses

263-268. A. Belin: Paris.

Cowie I.D. (2004). New species and lectotypifications

of some reticulate nerved Tephrosia (Fabaceae)

from north-west Australia and the genus

Paratephrosia is re-evaluated. Nuytsia 15:

163-185.

Desvaux, A.N. (1814). Journal de Botanique, Redigepar

une Societe di Botanistes 3: 78.

Domin, K. (1912). Fifth Contribution to the Flora of

Australia. Repertorium Specierum Novarum

Regni Vegetabilis 11: 261-264.

- (1926). Tephrosia. In Beitrage zur Flora

Australiens. Bibliotheca Botanica 89: 192-203.

Du Puy, D.J & Labat, J.-N. (1995). Pyranthus , a new

genus of Leguminosae. Kew Bulletin 50: 73-80.

- (2002). Tephrosia and Mundulea. In D.J. Du

Puy (ed.). The Legminosae of Madagascar 395-

424. Royal Botanic Gardens: Kew.

Estrada, A.E. & Martinez, A. (2003). Los Generos de

Leguminosas del Norte Mexico. Sida, Botanical

Miscellany No. 25.

Forbes, H.M.L. (1948). A revision of the South African

species of Tephrosia Pers. Bothalia 4: 951-1001.

Geesink, R. (1984). Scala Millettiarum. Leiden Botanical

Series 8: 1—131.

Gillett, J. (1958). Notes on Tephrosia in tropical Africa.

Kew Bulletin 13: 111-132.

- (1959). The indumentum of the style in the

taxonomy of Tephrosia in Africa, with notes on

T. virgata H.M.Forbes and T. euchroa Verdoorn.

Kew Bulletin 13: 414-419.

Harvey, W.H. (1862). Leguminosae. In W.H. Harvey &

O.W. Sonder, Flora Capensis 2: 1-285. Hodges,

Smith & Co.: Dublin.

Holland, A.E. & Pedley, L. (2010). Fabaceae. In P.D.

Bostock & A.E. Holland (eds ). Census of the

Queensland Flora 2010\ pp. 67-80. Queensland

Herbarium, Department of Environment and

Resource Management: Brisbane.

Jarvis, C. (2007). Order out of Chaos. Linnaean plant

names and their types. Linnean Society &

Natural History Museum: London.

242

Kajita,T., Ohashi, H., Bailey, C.D. & Doyle, J.J. (2001).

rbcL and legume phylogeny with particular

reference to Phaseoleae, Millettieae and allies.

Systematic Botany 26: 515-536.

Labat, J-N. & Du Puy, D.J. (1998). Sylvichadsia, a

new genus of Leguminosae -Papilionoideae -

Millettiae endemic to Madagascar. Adansonia ,

ser. 3. 20: 163-171.

Lewis, G., Schrire, B., Mackinder, B. & Lock, M. (eds.).

(2005). Legumes of the World. Royal Botanic

Gardens: Kew.

Linnaeus, C. (1747). Flora Zeylanica. Laurentii Salvii:

Stockholm.

- (1753). Species Plantarum. Laurentii Salvii:

Stockholm.

LPWG (2013). Legume phylogeny and classification

in the 21st century: progress, prospects and

lessons for other species-rich clades. Taxon 62:

217-248.

Meyer. E.H.F. (1836). Commentariorum de Plantis

Africae Australioris 111. A.P. Voss: Lipsiae.

Mueller, F. (1865). FragmentaPhytographiaeAustraliae

5: 9. Government Printer: Melbourne.

- (1875). Fragmenta Phytographiae Australiae 9:

64. Government Printer: Melbourne.

Necker, N. J. de (1790). Elementa Botanica. Societatum

typographicam: Neuwied.

Ottley, A.M. (1944). The American Loti with special

consideration of a proposed new section

Seripeteria. Brittonia 5: 81-123.

Persoon, C.H. (1807). Tephrosia. In Synopsis Plantarum

2(2): 328-330. C.F. Cramerum: Paris.

Rudd V.E. (1991). Fabaceae (Leguminosae). In M.D.

Dassanayake & F.R. Fosberg (eds.), A Revised

Hand Book to the Flora of Ceylon 7: 144-145.

Amerind Publishing Co. Pvt. Ltd.: New Dehli.

Schrire, B.D. (1987). A synopsis of Tephrosia subg.

Barbistyla (Fabaceae) in southern Africa.

Bothalia 17: 7-15.

- (1990). Tephrosia pondoensis. The Flowering

Plants of Africa 51: t. 2036.

Silva, M. J. Da, Quieroz, L.P. De, Tozzi, A.M.G. A, Lewis,

G.P., Sousa A.P. De (2012). Phylogeny and

biogeography of Lonchocarpus sensu lato and

its allies in the tribe Millettieae (Leguminosae,

Papilionoideae). Taxon 61: 93-108.

Austrobaileya 9(2): 229-243 (2014)

Stafleu, F.A. & Cowan, R.S. (1976). Taxonomic

Literature 2 nd ed. Bohn, Scheltema & Holkema:

Utrecht.

Stevens, PF. (1996). Why conventional classifications

form no basis for comparative studies - a

cognative perspective on Bentham and Hooker’s

“Genera Plantarum”. American Journal of

Botany 83(6). Abstract 556.

Verdcourt, B. (2007). Mundulea. In Flora Zambesiaca

3(3): 117-119. Flora Zambesica Managing

Committee: London.

Weatherby, C.A. (1935). Cracca In A. Rehder et al.

Conservation of later generic homonyms.

Bulletin of Miscellaneous Information, Royal

Gardens, Kew 1935: 341-544.

Wood, C.E. (1949). The American barbistyled species

of Tephrosia. Rhodora 51: 193-231, 233-302,

305-364, 369-384.

Pedley, Tephrosia in Queensland: 1

243

Map 1. Distribution of Tephrosia delicatula T. turpinii A and T. varians •. N.B. specimens discussed under the

Notes Section for T. delicatula are not mapped.

C.T. White’s botanical survey and

collections from Papua in 1918

A.R. Bean

Summary

Bean, A.R. (2014). C.T. White’s botanical survey and collections from Papua in 1918. Austrobaileya

9(2): 244-262. In July and August 1918, Cyril Tenison White spent four and a half weeks in Papua

New Guinea, during which time he went on two extended excursions and collected 829 botanical

specimens, of which at least 722 are known to be extant, representing 562 species. 32 new species

and two new varieties were named from specimens collected by White during his survey, including

15 new taxa authored or co-authored by White. This paper provides an itinerary for his time in

Papua, accurate geocodes for his collections, and descriptive and photographic background for his

excursions. The significance and legacy of White’s Papuan survey are discussed.

Key Words: historical botany, history, C.T. White, Brisbane, Papua, Papua New Guinea, herbarium

specimens

A.R. Bean, Queensland Herbarium, Department of Science, Information Technology, Innovation

and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia.

E-mail: Tony.Bean@dsitia.qld.gov.au

Introduction

Cyril Tenison White (1890-1950) was the

grandson of the very well known Queensland

Colonial Botanist, F.M. Bailey, and it was

from him that he learnt his botanical skills.

Upon the resignation of his uncle, J.F. Bailey

from the position of Government Botanist,

the position was given to C.T. White, initially

in an acting capacity, but formalised in

November 1917 1 . He was just 27 years old.

He had apparently been offered ‘an important

scientific post in the Pacific’ just prior to

his permanent appointment as Government

Botanist 2 . Judging by his correspondence and

other archived material, he was an enthusiastic

and hard-working man, and even in the

early stages of his career, an accomplished

taxonomist and collector (Fig. 1).

This paper provides a detailed examination

of C.T. White’s pioneering botanical survey

and collections in southern Papua New Guinea

(then known as the Territory of Papua) in July

and August 1918. The aims are to determine

accurate dates and geocodes for the places

visited by White; to provide descriptive

background for his excursions, and to analyse

the plant collections he made.

Accepted for publication 5 August 2014

Fig. 1 . C.T. White, c. 1915 3

Materials and methods

The Queensland Herbarium specimen

database (Herbrecs) was used to generate a

spreadsheet of White’s 1918 collections in

245

Bean, C.T. White in Papua

Papua. The author has located all of those

specimens, now contained in the collection

at BRI, to assess their quality, to check for

the presence of a field label, and to uncover

any information not recorded on the database.

Details of specimens not at BRI were extracted

from White (1922) and other publications of

the time, and from communication with other

herbaria, namely A, FI and NSW. The ‘Trove’

website (National Library of Australia

2009-onwards) has been extensively used, to

elucidate travel dates from Brisbane to Papua

and back, and for descriptive articles published

in newspapers of the time. A few letters from

the Queensland Herbarium archives have shed

light on some aspects of the trip, and visits to

the National Archives (Brisbane office) and

the University of Queensland (Fryer Library)

provided further details.

Some notes supplied by White (1922)

on floristics and vegetation are quoted here,

and these are supplemented with text and

photographs by other people who travelled in

Papua around the same period.

The latitudes and longitudes for the

various locations cited by White have been

determined using both contemporary maps

(e.g. O’Malley & Stanley 1916) and modern

sources (e.g. GeoNames website, Google

Earth).

Herbarium acronyms (e.g. A, BRI) follow

Thiers (continuously updated).

The journey from Australia

In June 1918, the Lieutenant Governor of

Papua, Horace “Judge” Murray, invited White

to visit Papua as his guest (White 1922).

White must have jumped at the opportunity,

as he arranged a visit for the following month.

Murray and White left Brisbane together

on the steamship ‘Marsina’ 4 on the 16 th July

1918 5 . The ‘Marsina’ (Fig. 2) had left Sydney

three days earlier 6 and was bound for Rabaul

via Port Moresby 7 . Steamer passage from

Brisbane to Port Moresby at that time took

about seven days, and White and Murray

probably arrived in Port Moresby during the

night of the 22 nd /23 rd July.

White collected plants around Port

Moresby for the first two days of his visit.

He then went on two overland excursions,

both times in company with Evan R. Stanley

(1885-1924), the Government Geologist for

Papua, who had been living and working in

Papua since January 1911 (Davies 1987). No

doubt Stanley organised the excursions and

chose the routes.

The residents of Port Moresby were clearly

underwhelmed by the visit of C.T. White. The

local newspaper does not mention his name

at any time during the visit - not even on the

passenger list 8 for the ship that carried him to

Port Moresby! Local residents returning from

time in Australia were written up, mining

engineers and businessmen were discussed at

length, but a visiting botanist, it seems, was a

non-event.

Port Moresby in 1918

World War I was still in progress, and every

issue of ‘The Papuan Courier’ was full of war

news from Europe, but life in Port Moresby

was seemingly normal. Steam ships cruised

around the western Pacific unimpeded and

trade between Australia and Papua was

vigorous and regular. There was a steady

export of goods such as copra, rubber, coffee

and sisal hemp 9 , and economic development

in Papua was booming.

Port Moresby had an estimated 1000

European inhabitants in 1914, and probably

more in 1918. It boasted two banks, a school,

two hotels, a telephone service (Stuart 1970),

and a 500,000 gallon cement reservoir, from

which water was reticulated throughout the

town 10 . An advertisement for one of the hotels

boasted “Lighted throughout by Electricity”

and “Dining Room furnished with electric

fans”, while the other hotel’s advertisement

meekly proclaimed, “Premises lighted by

Acetylene gas” 11 .

Port Moresbians had access to refrigerated

goods, as a “cold storage company”

commenced operations there in 1913, with

facilities for storing “mutton, meat, butter,

fruit and vegetables” 12 . Ice was available for

purchase at one penny per pound. 13 Motor

vehicles had arrived in Port Moresby by 1918.

246

Austrobaileya 9(2): 244-262 (2014)

Fig. 2. The steamship ‘Marsina’, on which White travelled to Port Moresby

At the news of an armistice in the war in

November 1918, it was reported that:

Motor cars screeched their owners’ gladness, and

choruses by the parading residents added to the

unwonted noise ... some of the weary ones then made

their way homeward, while others still on celebration

bent, motored to Sapphire Creek 14 .

At the time of White’s visit, Port

Moresbians were bemoaning the cold weather:

The extremely cold weather experienced during

the past month has caused a great quantity of sickness,

and an epidemic of colds has taken place. Old residents

of Papua say they cannot remember such climatic

conditions 15 .

The lowest overnight temperature at Port

Moresby for that year was 68°F (20°C) on the

31 st July 16 . A few months later, Papuans and

Australians alike were on high alert 17 for the

arrival of the influenza pandemic that killed

millions of people around the world.

Specimen numbering

Although White (1922) stated that the

invitation to visit Papua was “for the purpose

of studying its vegetation”, it was a plant

inventory and specimen collecting survey for

White, and only some general notes on the

vegetation were recorded.

For the first part of his career, C.T. White

used specimen numbers only sporadically,

and the great majority of his specimens

are unnumbered. It wasn’t until the mid-

1920s that he sequentially numbered his

collections on a more regular basis. His trip

to Papua; however, provides an exception,

as nearly all of the specimens have a unique

number, starting at 1 and finishing with 827,

so on the face of it, 827 specimens were

collected. However, in two instances, White

inadvertently applied the same number to

two unrelated specimens, so that the actual

number of specimens was 829. All numbers

247

Bean, C.T. White in Papua

up to 134 were for specimens labelled ‘Port

Moresby’, his initial destination, and the

ensuing numbers relate to his first excursion

eastward to Sogeri Plateau and Javararie.

This gives one hope that the numbering was

strictly chronological, but closer examination

reveals that the sequence of White’s collecting

numbers is only approximately chronological,

and the numbering sequence often fails to

correspond to the sequence of localities

along the route. For the second excursion, the

numbering is even more jumbled with respect

to the locations. It is clear therefore that White

did not allocate numbers to his specimens at

the time of collection, and this is corroborated

by the absence of a collection number on any

of the surviving field slips.

The last few numbers (819-827) were

allocated to specimens collected during the

first excursion. This reveals that numbering

was not done until after the second excursion

at the earliest. Because the numbers appear

only on the final stamped herbarium label, in

ink, I believe that White did not number his

specimens until he returned to Brisbane.

Specimen labels

Almost all of White’s specimens from his

1918 trip have a rectangular thin paper

Queensland Herbarium label with the

following information stamped upon it ‘Hab.

Papuan collections C.T. White No. July/Aug.

1918’, where ‘Hab.’ refers to the locality of

the specimen. Upon these labels, White has

handwritten (in ink) the species name, locality

and the specimen number (Fig. 3).

In addition, 40% of the BRI specimens have a

small slip of paper (either lying loose or glued

to the sheet) that bears White’s handwriting

in coloured pencil (Fig. 4). Such slips often

give just a locality name, but sometimes also

indicate the exact date of collection, or give a

brief description of the plant and even a rough

drawing of a fruit etc. Some of the slips are

quite dilapidated, and show signs of having

been wet, so they were clearly inserted with

the specimens at the time of collection. White

included such slips (or field labels) with all of

his specimens while in the field, to remind him

of the location and morphological features,

QUEENSLAND HERBARIUM.

BOTANIC GARDENS, BRISBANE,

77

Fig. 3. Example of the herbarium label for specimens

from White’s Papuan trip. The number showing through

from the reverse side is a herbarium database acquisition

number.

Fig 4. Example of one of White’s field slips. This one is

more detailed than most.

but many were not ultimately retained with

the herbarium specimen.

For his specimen No. 825, Phrynium

kaniense Loes. & G.M.Schulze, the only

locality given is ‘Papua, Central Division’.

Upon the herbarium label, White has

written “Unfortunately, I have lost my field

label with this, so cannot give more precise

habitat record”. This notation makes it clear

that he relied entirely upon the field slips for

the locality and other data that accompany

his specimens, and appear in his report

(White 1922). For example, the field label

for Castanopsis acnminatissima (Blume)

Rehder (labelled by White as Castanopsis

Schlenckerae) says “Large tree, dense foliage,

Common, Mafulu, 13/8/18”, while the notes

248

given by White (1922) for that species are

“Mafulu. Large tree, dense foliage, common”.

For about 14% of the specimens at BRI, there

is a field label that specifies the actual date

of collection. From these it has been possible

to piece together White’s daily movements

during the two excursions.

The specimens

White collected 829 specimens in an

estimated 21 collecting days and is a tribute

to his endurance and enthusiasm. He confined

his collections to the vascular plants, but

collected across many families.

Nearly all of White’s specimens are well

pressed and without mould (Fig. 5). However,

the quality is often poor; on many specimens

the leaves have detached from the stems,

and for a proportion of those, the leaves are

fragmented and have been placed inside a

packet; some specimens consist of fruits or

leaves only. The specimen quality varies with

both plant group and locality; for example the

grasses, sedges and ferns contain a higher

proportion of good fertile specimens than

average, while very few specimens from the

Mafulu and Deva Deva areas are of good

quality. About 26% of his specimens at BRI

are good intact fertile specimens. Because

knowledge of the New Guinea flora was

so rudimentary, White obviously opted for

quantity rather than quality. He does not

enlighten us as to the vicissitudes attending

his botanical collecting, except to say that the

trip “was of a very hurried nature” (Rogers &

White 1920). One major difficulty would have

been the rain - in many of the areas he visited,

such as Mafulu, “every afternoon is wet, and

most of the nights as well” (Grimshaw 1913c).

The poor quality of many specimens can likely

be attributed to his inexperience in collecting

under those conditions. It is also possible that

White did not have access to his plant presses

during the week-long trip back to Brisbane on

the steam ship, in which case any still-moist

specimens would have deteriorated.

There are many gaps in the number

sequence for specimens at BRI, White’s

‘home’ herbarium. There are three possible

reasons for a missing number at BRI.

Austrobaileya 9(2): 244-262 (2014)

1. For 18 specimens the herbarium label has

been lost, and with it the collection number,

thus causing a gap in the number sequence. In

this category are all of the ‘carpological only’

collections.

2. White or others may have discarded some

specimens after they had been numbered

- this is thought to account for most of the

missing specimens (see below), but it is not

possible to determine how many have been

lost in this way.

3. White sent all material of that specimen to

another herbarium. Some specimens in this

category have been traced:

Material of Rubiaceae and Acanthaceae

was sent to Spencer Moore at the British

Museum (BM), who subsequently sent White

a letter listing identifications for those 18 , and

numerous specimens of Zingiberaceae and

Marantaceae were passed onto H.N. Ridley

at Singapore. D. Prain, the then Director of

Kew Gardens, identified his specimens of

Acalypha, Macaranga and Mallotus (White

1922). He also sent three bamboo specimens

to Kew (numbers 238, 327 & 365) for the

examination of Otto Stapf 19 , but presumably

he had no response, as White (1922) does

not acknowledge a contribution by Stapf.

Only one of the three bamboo specimens

(number 327) is present at BRI and it remains

unidentified as Bambusa sp. He sent an

unknown number of palm specimens to

Odoardo Beccari at Florence (FI), but these

remained unidentified as Beccari died in 1920.

He sent a few pandan specimens to Ugolino

Martelli, also at FI. There are two isotype

specimens ( Psychotria mafuluensis S.Moore,

no. 416 & Psychotria whitei S.Moore, no. 702)

at the Arnold Arboretum in Boston, U.S.A.,

but no other 1918 White specimens are

currently known at A. Numerous specimens

were sent to the New South Wales herbarium

(NSW); Eucalyptus for the examination of

J.H. Maiden, Loranthaceae for W. Blakely,

and ferns for T. Whitelegge; all of these were

apparently returned to BRI, although some

duplicates remain at NSW. White evidently

sent some specimens to Friedrich Markgraf

at Friedrich Wilhelm University in Berlin,

Bean, C.T. White in Papua

249

QUEENSLAND HERBARIUM,

BOTANIC GARDENS, BRISBANE.

- cy a ./

Piper inacroplner Pennant (1800).

vide: Mew. In J. Arn, Arbor,

29: 191 (me)

Studied by: CHEW, Wee-lek. AA$ - $7° ■

078183

Fig. 5. One of White’s Papuan specimens (Piper macropiper Pennant, no. 255)

250

because on the labels of a few specimens at

BRI, he has written ‘seen by Dr Markgraf,

Berlin’.

Statistics

At BRI, there are 665 numbered specimens,

and 15 unnumbered (most of the latter were

previously numbered, but have lost their

herbarium label), making a total of 680

specimens. To this can be added 15 specimens

not at BRI but known to be present at another

herbarium due to being sent by White.

By comparing the holdings at BRI with

the annotated species list presented in White

(1922), it is apparent that there are 27 species

(all apparently collected by White, hence

27 specimens) collected by White, not now

present at BRI, and not listed in any letter or

publication associated with any of the authors

mentioned above. These include Heteropogon

contortus (L.) P.Beauv. ex Roem. & Schult.,

Elatostema sessile J.R.Forst. & G.Forst. and

Opilia amentacea Roxb. These are perhaps

present in another herbarium. Adding in these

27 specimens brings the total to 722.

However, that still leaves 107 specimens

for which nothing is known, except White’s

allocated specimen number. From these

we can determine that 24%of the missing

specimens are from the number series of

the first excursion (Sogeri, Sapphire Ck etc.;

135-414 & 819-827), 13% of the missing

specimens are from the Yule Island/Mekeo

District number series (705-818), 61% are

from the mountainous regions of the second

excursion, Mafulu, Deva Deva and Dilava, i.e.

the number series 415-704; and just 2% are

from the Port Moresby number series (1-134).

Taking into account the poor quality of many

of White’s extant specimens from the wet

mountainous areas of the second excursion, it

is difficult to avoid the conclusion that there

were further poor quality specimens that

White numbered but later discarded.

Of the 722 collections that are known, the

number of species represented is estimated at

562. This is many fewer than the number of

specimens, for the simple reason that White

collected some species twice or more times

(maximum was five times - for Acalypha

Austrobaileya 9(2): 244-262 (2014)

hellwigii Warb., Scaevola oppositifolia Roxb.

and Microlepia pseudohirta Rosenst.). In

most cases the duplicated species collections

were made from the different excursions or

at least from widely separated localities, but

surprisingly often they were from the same

location. For at least some of those in the

latter category, the morphology of the two

collections is very similar. This leads one to

suspect that the two specimens were actually

part of the same gathering that became

physically separated from each other during

drying activities, and then inadvertently

numbered separately by White upon his

return.

Port Moresby collection

White ranged widely around Port Moresby

in his quest for plant specimens; they include

some mangrove species, other maritime

plants, grassland and woodland species

and a couple from rainforest gullies. White

(1922) described the vegetation around Port

Moresby:

It consists for the most part of grass-covered hills with

scattered white-barked eucalypts {E. papuana and E.

alba ) of rather stunted growth dotted about. Other very

common trees on the hills are Alstonia scholaris (Milky

Pine), and Albizzia procera. A cycad (Cycas media ) is

also very abundant... Round about the rocky sea-coast,

near the town, Cochlospermum gillivraei (a small tree)

is conspicuous on account of its numerous large, yellow,

buttercup-like flowers.

It was the height of the dry season, and

most herbaceous plants would have already

died, with many shrubby species senescent or

torpid. However, this did not prevent White

making a very extensive collection. The

specimen quality is often mediocre, reflecting

the seasonal conditions. 137 Port Moresby

specimens are known: 125 from the number

series 1-134, plus no. 513, no. 820 and 10

unnumbered (Queensland Herbarium 2013),

all collected within walking distance of the

town centre. These could all be geocoded

with 1600 metre precision centred on 9° 28’

30”S, 147° 09’ 30”E. About 20 Port Moresby

specimens have field labels specifying the

date as either the 23 rd or the 24 th July, and it

is likely that the great majority were collected

on these two dates.

251

Bean, C.T. White in Papua

White rested in Port Moresby for a few

days following the first excursion, although

he undoubtedly spent considerable time

attending to the drying of his specimens.

There are four dated specimens from this

time, all collected on 6 th August.

First excursion

On the 25 th July, Stanley and White started

on an excursion to the Sogeri Plateau and

Javararie (Fig. 6), Table 1 . They travelled on

horseback 20 , and 15 native carriers followed

on foot (White 1922). In and near Port

Moresby, the carriers were paid with money

(one shilling a day), but away from the town

they were paid in salt or matches 21 . White

was interested to learn that salt was highly

prized and that for a tablespoon of salt, the

natives would trade a sugar-bag full of sweet

potatoes 22 . They headed along the only major

road out of the town, to the plantations of the

Laloki River (15.5 miles [25 km] from Port

Moresby), and onto Sapphire Creek (18 miles

[29 km] from Port Moresby), where they

camped for the night. 23

After Sapphire Creek, the track took

them onto the base of the Astrolabe Range.

White made numerous collections labelled

‘Astrolabe Range’ (26-27 July), presumably

on or near this steep section. This area is the

Varirata or Wariarata plateau. Judging by

the collection dates, White and Stanley did

not stop there until the return journey. They

reached Koitaki and Sogeri on the 28 th July.

Beaver (1905) said of Sogeri:

Sigoro [Sogeri] is one of the prettiest places I have come

across in New Guinea. Some 50 acres of sloping ground

have been cleared of jungle... the rainfall is rather

better than at Warirata and the soil is even richer. The

butterflies are of gorgeous colour, and range in size from

an eighth of an inch across to over 8 inches. The jungle

birds are equally brilliant in their plumage.

The Sogeri plateau was the site of a large

rubber plantation 24 established about 1910,

and in full production by the time of White’s

visit. White (1922) described the vegetation of

Sogeri plateau:

... the vegetation is very rich and tropical, the plateau

being mostly covered with heavy rain-forest in which

the usual Malayan orders and genera predominate.

Zingiberaceae and Marantaceae are particularly

abundant. Mucuna kroetkei is a forest climber with

long pendulous racemes of brilliant scarlet flowers

and is known locally as the “D’Albertis Creeper” 25 ...

Stanley and White camped on the range

between Sogeri and Javararie (now Iawarere)

on the evening of the 28 th July, perhaps leaving

the porters at Sogeri. On the 29 th July, Stanley

and White rode to Javararie, their furthest

point on the excursion. This was the site of

another rubber plantation, one that produced

some of the finest rubber in the Territory

(White 1922). They probably returned to

Sogeri late on the 29 th .

After making further collections around

Sogeri, White and Stanley made a detour to

Bisiatabu and Hombrom Bluff, to the north

of the main track they had been following.

They probably arrived in Bisiatabu late on

30 th July, as all collections from Bisiatabu and

Hombrom Bluff are dated 31 st July. Beaver

(1905) described the terrain near Hombrom

Bluff:

After quitting Tabouri, we left the Warirata track, and

struck off through deep gorges and across steep spurs.

The track was very narrow and rough — 800 feet below,

the river seemed a mere silver thread. Presently the

eye catches the scarp of Hombrom Bluff, rising almost

perpendicularly from the river bank.

Stanley and White then returned to

Varirata plateau by the evening of 1 st August,

where White made a dozen or so collections

with the locality given as “Mt Warirata”.

White (1922) described the vegetation of

these areas:

On the Astrolabe Range (about 3,000 feet), Hombrom

Bluff, Mt Warirata etc., the vegetation for the most part

is of an open character, the principal forest trees being

eucalypts (principally E. tereticornis) with patches of

E. alba and E. clavigera , Casuarina nodiflora, Banksia

dentata, Melaleuca sp. (a Paper-barked Tea-tree),

Diplanthera tetraphylla, Grevillea pinnatifida and

Timonius Rumphii. At Bisiatabu, I was interested to find

Nepenthes moorei to be a common plant in the poorer

open, dry, forest country.

They camped on the lower reaches of the

Laloki River on the afternoon of 2 nd August,

arriving early enough to allow White to make

about 12 specimen collections. They would

have arrived back in Port Moresby in the early

afternoon of the 3 rd August.

252

Austrobaileya 9 ( 2 ): 244-262 ( 2014 )

Table 1. Summary of localities visited during White’s first excursion to Papua

Locality used by White

Collecting date(s)

(year is 1918)

Latitude

Longitude

Altitude 1

(metres)

Astrolabe Range

26 July, 27 July, 30

July, 1 August

9° 25’S

147° 22’E

Between Koitaki

and Sogeri

28 July

9° 24’S

147° 29’E

490

Bisiatabu

31 July

9° 24’S

147° 25’E

510

Hombrom Bluff

31 July

9° 23’ S

147° 20’E

653

Javararie (now

Iawarere/ Iawarare)

29 July

9° 27’S

147° 37’E

548

Koitaki (now Koitaki

Plantation)

28 July

9° 24’S

147° 28’E

479

Laloki River

2 August

9° 24’S

147° 18’E

200

Lawsons Lookout

31 July

9° 24’S

147° 25’E

510

Mt Warirata (now

Varirata / Wariarata

Plateau)

1 August

9° 25’S

147° 22’E

573

Ranges between

Sogeri and Javararie

28 July, 29 July

9° 26’S

147° 33’E

Rona Falls (now

Rouna Falls)

27 July

9° 25’S

147° 23’E

Sapphire Creek

25 July

9° 24’S

147° 19’E

220

Sogeri 2

28 July, 29 July

9° 24’S

147° 29’E

500

Footnotes

'Geocodes and altitudes mainly from Geonames website.

2 The modern location of Sogeri differs considerably from that used by O’Malley & Stanley (1916), and hence by White.

Second excursion

On their second excursion, Stanley and White

travelled into the mountainous region of

Mafulu, via Yule Island and Bioto (Fig. 6).

Stanley attended the first annual dinner

of the Public Service Association at Port

Moresby on the evening of the 6 th August 26 , and

it is likely that White was present as a guest.

Stanley evidently wanted to delay the start of

the second excursion so that he could attend

this dinner. No details have been discovered

about their departure or for the timing of the

early part of the journey, but to reach Mafulu

by the 11 th August (see below), they must have

left Port Moresby straight after the dinner, or

in the early hours of the following day. It is

not known which vessel they boarded to reach

Yule Island, but a journey there by steamer

from Port Moresby took about six hours 27 .

At Yule Island, where there was a

Government mission, Stanley employed 25

native porters for “carrying the camping outfit,

collecting gear, specimens etc” (White 1922).

They travelled from the island (probably

late on the morning of 7 th August) by native

canoes to the nearby mainland, following

the Ethel River (Bioto Creek, Fig. 7) to the

small village of Bioto. White commented

that “time did not permit of much collecting

along the banks of the Ethel River” but also

that he “collected specimens of the apparently

little-known Sonneratia lanceolata ” (White

1922). This Sonneratia specimen is the only

one at BRI that bears the locality ‘Ethel

River’, although one additional species, Luffa

aegyptiaca Mill., is listed in White (1922).

Stanley and White did not stay the night at

Bioto, nor spend much time there. Murray

(1912) said of Bioto:

Bean, C.T. White in Papua

253

Fig. 6. Routes taken by White during his two excursions in Papua.

254

The mosquitos swarm there even in the dry season in

the blazing midday sun, and the imagination staggers at

the thought of what they might be at night. I for one have

never passed a night there, and trust that I never shall.

In contrast to the first excursion, the second

was entirely on foot. White (1922) stated that

from Bioto it was a “five days’ march to the

mountains” and that the following places

were “stopped at en route. Kubunah, Fofofofo,

Dilava, Deva Deva, Mafulu and Bella Vista”.

A few of White’s ‘Mafulu’ collections are

dated ‘11/8/1918’. The fact that they reached

Mafulu by the 11 th August means that they did

not dally, and made only overnight stops at

each village or mission station on their way

inland. It seems very likely that White did

little or no collecting until he reached Mafulu.

In the early stages of the journey, the porters

would have been heavily loaded with the food

required to sustain them for the two-week

Austrobaileya 9(2): 244-262 (2014)

trip; only in the latter stages of the trip (when

food stores were reduced) would they have

had the capacity to carry the extra weight

and bulk of White’s botanical specimens. A

quick inward trip, with all collecting done

on the return journey was the stated strategy

employed by the Archbold Expedition many

years later (Archbold & Rand 1935).

White did not describe the route they

took or the places they camped, but a series

of newspaper articles by an unknown author

(Anon. 1921a, 1921b, 1921c) detailed a trip

following the same route a few years later.

This party also started their expedition from

Yule Island, landed at Bioto, spent their first

night at Kubuna, and travelled onto Mafulu.

In addition, Beatrice Grimshaw had written

a very interesting account of a trip to Mafulu

a few years earlier (Grimshaw 1913a, 1913b,

1913c).

Fig. 7. Bioto Creek, featuring Nypa palms (Nypa fruticans Wurmb) 28 .

255

Bean, C.T. White in Papua

Stanley, White and the porters walked to

Kubuna for the first night’s camp. They were

no doubt anxious to reach the slightly cooler

foothills where the mosquitos were not so

frequent. Anon. (1921a) described the track

from Bioto to Kubuna thus:

We ... wound our way through random grass fires in the

sunblasted country beyond it, along an uphill stony track,

which burned the foot through the sole of the boot. A few

miles out, we stepped out of the eucalyptus belt into the

ficus scrub, out of the dry heat into the moist, ... we had

plunged into tropical scrub, which became steadily denser

and ran ahead of us for many miles ... A three hours’

steady tramp brought us to the first of the hill creeks,

where we halted to boil tea. The carriers came up, running

with sweat, and sat in the water among the rocks.

The next overnight stop for Stanley and

White was at Fofo Fofo as described by Anon.

(1921b).

Fofo-Fofo ... is a clearing of a few acres in the surrounding

jungle at a height of something between 2000 and 2500

feet - a clearing made for the rest-hut... Fofo-fofo is what

the natives call the bell-bird, which leads the morning

chorus here.

The next night they stopped at Dilava (Fig. 8)

as described by Grimshaw (1913a).

After an almost endless pull up, round a comer we

came upon the tiny station of Dilava, perched like a

swallow’s nest on the top of a sharp-edged peak ... on

top of the tiny peak were crowded together some small

stick buildings - Sister’s house, kitchen, school. Father’s

house, and a church.

The terrain was now very rugged indeed

(Grimshaw 1913b).

The track to Deva-Deva ... wet by daily rains, is so

slippery that no-one could walk on it without nailed

boots. No white person, as a matter of fact, ever attempts

to do so ... on leaving Dilava the track winds down and

down from the 3000 feet height on which the mission

stands to 1000 feet or so above sea level. After crossing

the river, it takes breath and shoots up to 4200 feet in

height. The tall tree-like bamboo of the lower range gives

way to the exquisite bamboo liane ... begonias, shell-

pink and scarlet, dangling their full racemes along the

track; balsams, pale coral colour; crimson honeysuckle;

white, white graceful nameless flowers, appearing in the

dim greenness of the bush.

Fig 8. Dilava Mission in 1921 29 .

256

Their next night was spent at Deva Deva (Fig.

9) as described by Anon. (1921c).

Deva-Deva, like Dilava, is sited on a tiny promontory of

hill-spur which juts out over a deep valley, filled with a

sea [of] cloud morning and evening.

White reported that the 25 porters they

employed at Yule Island deserted them at Deva

Deva. Other natives were soon requisitioned,

including four women:

Austrobaileya 9(2): 244-262 (2014)

One of the females not only carried a pack weighing 35

lbs, but a young pig which she commandeered on the

wayside, and upon which she bestowed much care 31 .

The last leg was from Deva Deva to

Mafulu. The distance was 13 miles, a distance

that the “practiced walkers of the mission

could traverse in four hours” (Grimshaw

1913b).

Fig 9. Deva Deva in 1921 30 .

It was very, very lovely ... there were streams all the

way, dropping down from unseen heights among white

and pink orchids, and tree-ferns like umbrellas of green

lace ... there were little flowers like wild roses; there were

raspberries, sweet and good; the dim places of the forest

were lit with scarlet flames of balsam; and the pines 32 , by

now, began to thicken.

On the 11 th August, Stanley and White

arrived at Mafulu. Up to this point, they had

been travelling through dense jungle, but at

Mafulu (Anon. 1921c):

The valley opens into reaches of open grass land, rich,

sunny, well-watered, well-drained soil. Here the villages

grow sweet com in abundance, yams in places cover the

hillsides for miles, bananas are common....

Archbold & Rand (1935) hypothesised that

the mountain-valley grasslands are formed by

deliberate burning, but that “conditions which

are dry enough for extensive burning probably

occur only once in several years”. They

further commented that the grasslands were

maintained by the continual clearing of the

secondary growth for gardens by the natives.

White (1922) did not mention grassland at all,

but instead stated:

257

Bean, C.T. White in Papua

In the ranges about Mafulu (about 4000 feet) the vegetation

is extremely rich and varied, consisting almost entirely

of heavy rain-forest. Among trees the ordinary Malayan

types predominate; ferns, lycopods, begonias, palms,

bamboos and other typical tropical forms are abundant.

The occurrence of Grevillea is a connecting link with

the flora of Australia, while Quercus, Castanopsis and

Begonia are Asiatic types not as yet found in Australia.

Davies (1987) recorded that during White

& Stanley’s trip, Stanley made numerous

observations on the geology of the Mafulu

area, and discovered a considerable range in

age of the volcanic rocks there, from Miocene

to Pleistocene. It is clear that Mafulu was

Stanley’s main objective. Stanley and White

spent less than three days at Mafulu (11—13 th

August), leaving on the afternoon of the 13 th .

There are 100 extant White collections from

there, the largest number for any locality

except Port Moresby.

There is nothing in Papua more beautiful than the little

mission-house in Mafulu. The house itself, church and

kitchen are built of sawn boards—almost a necessity in

this damp and chilly climate. The site is magnificent—a

little peak four thousand feet high, set in the midst of a

ring of splendid mountains ... From the verandah, one

can see the giant pandanus of the upper ranges dotting

the green slopes of the neighbouring peaks (Grimshaw

1913c).

The only collections that White made

beyond Mafulu were from Bella Vista station,

where there was a garden tended by a priest

from Alsace province of France. It was located

along the track a few kilometres past Mafulu.

White made only eight collections from Bella

Vista, so he didn’t spend much time there. He

must have visited Bella Vista on one of the

days that they were encamped at Mafulu.

Anon. (1921c) said of Bella Vista:

The oranges in the Alsatian garden of Bella Vista, beyond

Mafulu, are the equal of any that grow in the valley of

the Murray or in California. We received them by the

armful: we could consume no more when we had eaten

half a dozen each. Wild balsam flowers of varied colours

... adorn the track ... right up the Mafulu valley.

As noted before, both the dates on the

extant field labels, and the sequence of

numbering suggest very strongly that White’s

collections were made firstly at Mafulu and

then during the return journey to Yule Island.

From the relatively few dated field labels,

it has been possible to roughly determine

White’s itinerary, and to infer that the total

duration of the journey (from Yule Island

back to Yule Island (second day)) was 13

days, that is, from the 7 th August to the 19 th

August (Table 2). The numbers of specimens

collected at each location allows an

estimation of how long the party spent there.

A consideration of the number of specimens

collected, the dates and distances gives rise

to the inevitable conclusion that White must

have been snatching specimens from daylight

till dusk for much of the return journey, with

the pressing and note-taking done after dark

in the huts.

The following itinerary is hypothesized:

White collects at Mafulu (100 known

specimens) and Bella Vista (8 specimens) on

the 11 th , 12 th and 13 th August, and walks back

to Deva Deva for the night of the 13 th . He

collects at Deva Deva (50 known specimens)

on the 14 th , and travels onto Dilava for the

evening of the 14 th . Further collecting at

Dilava (25 known specimens) on the morning

of the 15 th and on to Fofofofo (16 specimens)

for the night. On the 16 th they travel from

Fofofofo to Diene (3 specimens) and onto

Kubunah (3 specimens), a distance of about

20 km as the crow flies. On the 17 th , they travel

from Kubunah through the Melceo District (31

specimens) to Bioto (7 specimens), and then

by canoe via Ethel River (2 specimens) back

to Yule Island, possibly arriving after dark.

On the 18 th and 19 th , collecting is carried out

at Yule Island (70 specimens).

The Mekeo District was a name used

for a large amorphous tract of lowland

country adjacent to Yule Island. It seems that

White applied the name ‘Mekeo District’ to

specimens he collected between Kubunah and

Bioto, as there are no specimens with the latter

designation, but there are several specimens

labelled ‘between Kubunah and Fofofofo’ or

‘between Fofofofo and Dilava’.

White left Papua on the steamship

‘Morinda’, along with 29 other passengers 33 ,

on the 23 rd August 34 , and he arrived back in

Brisbane early on the 30 th August 1918 35 . The

total time away from Brisbane was just six

and a half weeks, with four and a half weeks

spent in Papua.

258

Austrobaileya 9(2): 244-262 (2014)

Table 2. Summary of localities visited during White’s second excursion in Papua

Locality used by

White

Collecting date(s)

(year is 1918)

Latitude 1

Longitude 2

Altitude 3

(metres)

Bella Vista (now

Belavista)

12 August or 13

August

8° 33’S

147°03’E

1450

Bioto

17 August

8° 44’S

146° 37’E

15

Deva Deva

14-15 August

8°33’S

146° 58’E

770

Diene

16 August

8° 38’S

146° 51’E

480

Dilava

14-15 August

8° 35’S

146° 56’E

870

Ethel River (now

Bioto Creek)

17 August

8° 46’S

146° 36’E

5

Fofofofo (now Fofo

Fofo)

15 August

8° 37’S

146° 53’E

Kubunah (now

Kubuna Mission)

16 August

8° 42’S

146° 45’E

100

Mafulu

11-13 August

8°33’S

147° 00’E

1250

Mekeo district

17 August

8° 42’S

146° 40’E

50

Yule Island 4

18-19 August

8° 49’S

146° 32’E

10

Footnotes

'seconds of arc have been truncated

Seconds of arc have been truncated

3 Altitude figures have mostly been gleaned from Archbold & Rand (1935)

4 Yule Island geocode is for Kairuku Government station (see Murray 1912). One of White’s specimens (n. 772) actually

specifies ‘Kairuku’ as the locality

Results of the survey

Despite the brevity of his Papuan visit,

and the climatic challenges for specimen

collecting, as evidenced by the poor quality of

many specimens, the survey was undoubtedly

significant in terms of new taxa discovered

(Table 3). Several botanists had preceded

White on the route of his first excursion to

Sogeri and Javararie, but he was the first

botanist to collect specimens from the

non-coastal areas of his second excursion,

i.e. between Kubuna and Mafulu. As a

consequence, numerous species collected

from his second excursion were undescribed.

34 new taxa were named from specimens

collected by White during his trip, including

15 new taxa authored or co-authored by White

(Table 3)

The taxonomic scope of his collection

White made numerous collections of some

plant families or groups, while other families

or groups were apparently ignored.

He made an impressive collection of 90

ferns, including 14 Dennstaedtiaceae and

9 Pteridaceae, perhaps with the knowledge

that he could later ask for identifications from

Thomas Whitelegge at NSW He collected 70

grasses (Poaceae), 50 pea-flowers (Fabaceae),

and 43 Rubiaceae. Other well collected

families include the Gingers (Zingiberaceae),

Marantaceae, Asteraceae, Acanthaceae, and

Euphorbiaceae.

On the other hand, White collected no

filmy ferns or tree-ferns, no water-plants,

and nothing from the families Araliaceae,

Celastraceae, Cunoniaceae, Flacourtiaceae

and Icacinaceae, all of which are highly

speciose inNew Guinea. Other omissions were

the families Chrysobalanaceae, Clusiaceae,

Loganiaceae, Monimiaceae, Musaceae,

Symplocaceae and Thymelaeaceae. The

genus Begonia , with scores of species in New

Guinea, is represented by two collections; he

collected no gymnosperms apart from two

259

Bean, C.T. White in Papua

Cycas collections, and just three collections

each of Gesneriaceae and Annonaceae.

In many cases, the 1918 specimen collected

by C.T. White was the first New Guinea

record for that species to be incorporated in

the Queensland Herbarium. With the many

ensuing collectors and collections, nearly all

species have been collected numerous times

since, but there are still some species not

represented at BRI by any specimen other

than that of White ( Amaracarpus cimeifolius

Valeton, Brachyachne ambigua Ohwi,

Dendrocnide cordata (Warb. ex H. J.RWinkl.)

Chew, Echinochloa picta (K.D.Koenig)

P.W.Michael, Eriolopha ovalifolia Ridl.,

Macaranga coggygria Airy Shaw, Pandanus

yuleensis H.St.John, Riedelia whitei Ridl.,

Timonius cryptophlebus S.Moore and

Waltheria indica L.).

Naturalised species

White’s collection of 50 naturalised

species is an important subset of his Papuan

collection and represents the earliest

substantial vouchering of naturalised taxa

in New Guinea. Some are perennial shrubs

e.g. Jatropha gossypiifolia L., but most

are herbaceous annuals e.g. Bidens pilosa

L., and several vines are also included e.g.

Cardiospermum halicacabum L. The major

represented families are Malvaceae (9),

Poaceae (9), Asteraceae (7) and Fabaceae (6).

Most of the naturalised species were collected

at Port Moresby, but three were collected

from Yule Island, three ( Bixa orellana L„

Dichrocephala integrifolia (L.f.) Kuntze and

Urena lobata L.) were gathered from Mafulu,

three from Sogeri, two from Sapphire Creek,

two from Laloki River, and one each from

Bella Vista, Bisiatabu, Astrolabe Range

and Javararie. There is a marked positive

correlation between the number of naturalised

species collected and the permanence or size

of the human settlement. His collection also

included at least one cultivated species {Ficus

callophylla Blume, from Port Moresby).

Despite the fact that White had been in

Papua for just four and a half weeks, he was,

in the 1920s, the only Australian botanist in

the Government employ who knew anything

about the flora of Papua, so he was quickly

considered an expert. He promoted exploration

and collecting by others, and in 1925 he was

instrumental in having a young Toowoomba

man, Leonard Brass, appointed as botanical

collector for the Arnold Arboretum in Boston,

U.S.A. (Frodin 1990: 200). Brass was to

become a very significant plant collector in

New Guinea. White himself returned to New

Guinea in 1944 and 1945.

Acknowledgements

My research has been facilitated by the

marvellous resource ‘Trove’, a website

provided by the National Library of Australia.

All of the newspaper articles cited herein were

located using this website. I acknowledge

the State Library of Victoria for the use of

the image of the ‘Marsina’, and the National

Library of Australia for the use of three

images taken by Frank Hurley. I thank the

librarians at the Fryer Library, University of

Queensland, for their willing assistance, and

the staff of the National Archives facility at

Cannon Hill, Brisbane. I am grateful to Chiara

Nepi (FI) for information on Dr Beccari and

his correspondence with C.T. White. Anthony

Brach, Liz Francis, Lisa Pearson and Emily

Wood of Harvard University helpfully

responded to my queries. Will Smith (BRI)

produced the excursion maps, and Peter

Bostock checked the identity of some ferns.

References

Anonymous (1921a). A Walk in Papua. Out of the dry

belt. The lower hills. The Sydney Morning

Herald 5 Oct 1921, p. 10. http://trove.nla.gov.au.

Accessed 8 June 2013.

-(1921b). A Walk in Papua. Birds of Fofo-Fofo. The

Dilava Villagers. The Sydney Morning Herald 6

Oct 1921, p. 8. http://trove.nia.gov.au. Accessed

8 June 2013.

-(1921c). A Walk in Papua. Roadmakers’ Camp. The

Lovely Mafulu Valley. The Sydney Morning

Herald 8 Oct 1921, p. 12. http://trove.nla.gov.au.

Accessed 8 June 2013.

Archbold, R. & Rand, A.L. (1935). Results of the

Archbold Expeditions. No. 7. Summary of the

1933-1934 Papuan Expedition. Bulletin of the

American Museum of Natural History 68: 527-

597.

260

Austrobaileya 9(2): 244—262 (2014)

Table 3. New taxa named from White’s 1918 Papuan specimens

Taxon name

Family

Buchananiapapuana C.T.White

Anacardiaceae

Cupaniopsis insularis Radik.

Sapindaceae

Cupaniopsis papuana Radik.

Sapindaceae

Dendrochilum murrayi Rogers & C.T.White

Orchidaceae

Desmodiumpapuanum C.T.White

Fabaceae

Diospyros ferrea var. villosa Bakh.

Ebenaceae

Eriolopha ovalifolia Ridl.

Zingiberaceae

Goody era hispidula Rogers & C.T.White

Orchidaceae

Grevillea densiflora C.T.White

Proteaceae

Grevillea subargentea C.T.White

Proteaceae

Habenaria ramosa Rogers & C.T.White

Orchidaceae

Helicia latifolia C.T.White

Proteaceae

Helicia validinervis C.T.White

Proteaceae

Hemigraphis ciliata S.Moore

Acanthaceae

Hemigraphis whitei S.Moore

Acanthaceae

Hidemacanthus S.Moore

Acanthaceae

Hulemacanthus whitei S.Moore

Acanthaceae

Ixora whitei S.Moore

Rubiaceae

Justiciaplatyphylla S.Moore

Acanthaceae

Loranthus barbellatus Blakely

Loranthaceae

Mucuna Stanleyi C.T.White

Fabaceae

Mussaenda whitei S.Moore

Rubiaceae

Phacelophrynium whitei Ridl.

Marantaceae

Physurus bicalcaratus Rogers & C.T.White

Orchidaceae

Psychotria decorifolia S.Moore

Rubiaceae

Psychotria mafuluensis S.Moore

Rubiaceae

Psychotria whitei S.Moore

Rubiaceae

Riedelia lanatiligulata Ridl.

Zingiberaceae

Riedelia whitei Ridl.

Zingiberaceae

Ripogonum papuanum C.T.White

Ripogonaceae

Semecarpus undulata C.T.White

Anacardiaceae

Spathoglottispapuana var. puberiflora Rogers & C.T.White

Orchidaceae

Terminalia okari C.T.White

Combretaceae

Timonius cryptophlebus S.Moore

Rubiaceae

261

Bean, C.T. White in Papua

Beaver, W. (1905). Over New Guinea Mountains. The

Argus, 14 October 1905, page 5. http://trove.nla.

gov.au. Accessed 20 May 2013.

Davies, H.L. (1987). Evan Richard Stanley, pioneer

geologist in Papua New Guinea. BMR Journal

of Australian Geology and Geophysics 10:

153-177.

Frodin, D.G. (1990). Explorers, institutions and outside

influences: botany north of Thursday. In PS.

Short (ed.). History of Systematic Botany in

Australasia, pp. 193-215. Australian Systematic

Botany Society Inc.: Burwood, Victoria.

Grimshaw, B.E. (1913a). Among the Pygmies of Papua

and elsewhere VII. The Sydney Morning

Herald, 25 January 1913, p. 5. http://trove.nla.

gov.au. Accessed 22 June 2013.

-(1913b). Among the Pygmies of Papua and

elsewhere VIII. The Sydney Morning Herald,

1 February 1913, p. 5. http://trove.nla.gov.au.

Accessed 22 June 2013.

-(1913c). Among the Pygmies of Papua and

elsewhere IX. The Sydney Morning Herald,

8 February 1913, p. 5. http://trove.nla.gov.au.

Accessed 8 June 2013.

Murray, J.H.P. (1912). Papua or British New Guinea.

T. Fisher Unwin: London. A Project Gutenberg

Australia ebook. http://gutenberg.net.au/

ebooksl2/1202531h.html. Accessed 17 June

2013.

National Library Of Australia (2009-onwards). Trove.

http://trove.nla.gov.au. Accessed 13 December

2013.

O’malley, J.T. & Stanley, E.R. (1916). Map of the

Central Division, Papua. Mitchell Library,

Public Library of New South Wales.

Queensland Herbarium (2013). Specimen label

information (HERBRECS), viewed 3 June

2013.

Rogers, R.S. & White, C.T. (1920). A contribution to

the orchidaceous flora of Papua (British New

Guinea). Transactions and Proceedings of the

Royal Society of South Australia 44: 110-119.

Stuart, I. (1970). Port Moresby: Yesterday and Today.

Pacific Publications: Sydney.

Thiers, B. (continuously updated). Index Herbariorum:

A global directory of public herbaria and

associated staff. New York Botanical Garden’s

Virtual Herbarium, http://trove.nla.gov.au.

Accessed 5 October 2013.

White, C.T. (1922). A contribution to our knowledge

of the flora of Papua (British New Guinea).

Proceedings of the Royal Society of Queensland

34: 5-65.

262

Austrobaileya 9(2): 244—262 (2014)

Endnotes

1 Newspaper notification; The Brisbane Courier, 10 November 1917, p. 11.

2 Newspaper article; The Capricornian, 17 November 1917, p. 27.

3 photograph in Queensland Herbarium archives. The object in White’s hands is not known.

4 Letter from C.T. White to W. Bradtke, dated 13 July 1918. (Queensland Herbarium archives).

5 Newspaper article: The Brisbane Courier 17 July 1918, p. 11.

6 Newspaper notification; The Sydney Morning Herald, 15 July 1917, p. 8.

7 Newspaper advertisement; The Brisbane Courier, 1 October 1918, p. 2.

8 Newspaper notification; The Papuan Courier, 26 July 1918, p. 4.

9 Newspaper article. Chronicle, 7 December 1918, p. 30.

10 Newspaper article. The Register, 7 September 1916, p. 7.

11 Newspaper advertisement. The Papuan Courier, 9 August 1918, p. 8. (Fryer Library

microfilm).

12 Newspaper article, Townsville Daily Bulletin, 4 January 1913, p. 12.

13 Newspaper article. The Brisbane Courier, 26 February 1914, p. 7.

14 Newspaper article, Cairns Post, 20 November 1918, p. 7.

15 Newspaper article. The Papuan Courier, 9 August 1918, p. 4.

16 Newspaper article. The Papuan Courier, 9 August 1918, p. 4.

17 Newspaper article, Cairns Post, 13 December 1918, p. 5.

18 Lists compiled by S. Le Moore, undated. (Queensland Herbarium archives).

19 Letter from C.T. White to D. Prain, Director, Kew Gardens, dated 28th February 1919. (Queensland Herbarium

archives).

20 Newspaper article - The Queenslander, 14 September 1918, p. 11.

21 Newspaper article - The Queenslander, 14 September 1918, p. 11.

22 The Border Morning Mail and Riverina Times, 17 September 1918, page 4

23 Some of White’s Sapphire Creek specimens are dated 25 th July.

24 Townsville Daily Bulletin, 4 January 1913, p. 12.

25 Named for the well known Italian collector Luigi D’Albertis (1841-1901).

26 Newspaper article. The Papuan Courier, 9 August 1918, p. 5.

27 Newspaper article. The Sydney Morning Herald, 14 April 1915, p. 5.

28 Frank Hurley, 1921, National Library of Australia nla.pic-vn3314284.

29 Frank Hurley, 1921, National Library of Australia nla.pic-vn3314344.

30 Frank Hurley, 1921, National Library of Australia nla.pic-vn3316087.

31 Newspaper article. The Border Morning Mail and Riverina Times, 17 September 1918, p. 4.

32 Presumably Hoop Pine ( Araucaria cunninghamii).

33 Newspaper notification; The Papuan Courier, 30 August 1918, p. 4.

34 Based on seven day’s travel time from Port Moresby to Brisbane.

35 The arrival date of 30 th August in Brisbane has been confirmed by a passenger list for the Morinda (National

Archives of Australia, Brisbane office). An “a.m.” arrival in Brisbane is based on travel time of 2.5 days from the

reported arrival in Sydney on the evening of 1 September (The Sydney Morning Herald, 2 September 1918, p. 8).

The botanical collections of Ebenezer Cowley

John Leslie Dowe

Summary

Dowe, J.L. (2014). The botanical collections of Ebenezer Cowley. Austrobaileya 9(2): 263-278.

Ebenezer Cowley (1849-1899) was active as a botanical collector in north Queensland and British

New Guinea (now Papua New Guinea) during 1889-1898. His official role was Overseer of the

Kamerunga State Nursery. Cowley sent all his specimens to F.M.Bailey, Government Botanist in

Brisbane. About 370 Cowley specimens have been located, and most of these, c. 330, are extant

at BRI. A small number (c. 40) of duplicates and unicates are housed at BM, CNS, DNA, K, L,

M, MEL and NSW. The provenance of Cowley’s north Queensland collections included mostly the

Kamerunga and Barron River areas. 47 Cowley specimens are significant in typification, mainly

through the taxonomic work of F.M.Bailey: of these, 19 are associated with species names in current

use. The taxa that involved Cowley specimens in typification are listed with annotations. Cowley’s

1 st daughter, Clara Moxon (nee Cowley) (1875-1905) also sent plant specimens to, and corresponded

with, F.M.Bailey.

Key Words: Ebenezer Cowley, Clara Cowley, Queensland plant collectors, Kamerunga State Nursery,

typification of north Queensland plants, F.M.Bailey

John Leslie Dowe, Australian Tropical Herbarium, James Cook University, Cairns, Qld, Australia.

Email: john.dowe@jcu.edu.au

Introduction

Ebenezer Cowley (born 18 Apr. 1849, died 9

Feb. 1899) (Fig. 1) was active as a botanical

collector in north Queensland and Papua New

Guinea (then known as British New Guinea)

during 1890-1898. His birth was registered

in Cirencester District, England, and his first

years were spent in Fairford, Gloucestershire

(Barrett 2013a). Details of his early schooling

are unknown, but he migrated to South

Africa prior to 1871 and trained at the Durban

Botanic Gardens as a horticulturist (Gibbney

& Smith 1987). In South Africa, he studied

the flora of Natal whilst engaged in the sugar

milling industry (Stephens 1981). Leaving

South Africa in 1873, he travelled to Australia

and later to Fiji (1874?) where he was involved

with sugar milling. Moving to Queensland in

1881, he managed the CSR Victoria Plantation,

near Ingham, until 1885 (Griggs 2003), after

which he joined his cousin Alfred Sandlings

Cowley (later Sir Alfred) in pastoral and

sugar-cropping activities on properties in the

Herbert River area and Mourilyan (Stephens

1981; Saunders 2013).

Accepted for publication 18 August 2014

Fig. 1. Photograph of Ebenezer Cowley, 1890.

Photographer unknown, courtesy of the Cairns

Historical Society.

264

In 1889, Cowley was appointed the first

Overseer of the Kamerunga State Nursery near

Cairns, a position he held until his death from

liver disease in 1899 (Stephens 1981; George

2009). In Cowley’s first year at Kamerunga,

Government Botanist F.M.Bailey noted that

Cowley had “during the past year collected

specimens of several interesting plants”

(Bailey 1891). Cowley contributed to matters

regarding tropical horticulture and was

regularly quoted with respect to his knowledge

on the subject (Shelton 1892; Anon. 1896;

Sim 2001, 2012). Griggs (2003) noted the

apparent lack of formal training with regard

to Cowley’s sugar varietal research activities,

and the subsequent relocation of sugar cane

research to Mackay in central Queensland in

the late 1890s (Griggs 2011).

The Kamerunga State Nursery (from

1940 known as Kamerunga Research Station)

was established in 1889, as an initiative of

the newly formed Queensland Department

of Agriculture. The Nursery’s primary

emphases were trialling and experimenting

with tropical crops of commercial interest

(Thomatis 1895; Benson 1914; Stephens 1981;

Barker & Byford 1988). Because of wartime

austerity, the Nursery closed in 1916, and the

land was put up for lease. In 1940, part of the

grounds were reacquired by the Department

of Agriculture and Stock, and used as a

research station until the late 1990s (Cairns

Plan 2005). The grounds are now occupied

by Cairns Water, private housing, part of the

Kamerunga Conservation Reserve, and vacant

allotments. Some of the former plantings still

exist as remnants within the area.

During the period when Cowley was

Overseer at the Nursery, some significant

introductions were trialled (Bailey

1910), including Carthamus tinctorius L.

(safflower), Castilla elastica Sesse (Panama

rubber), Cocos nucifera L. (coconut palm),

Coffea arabica L. (coffee). Cola acuminata

(PBeauv.) Schott & Endl. (cola nut), Elaeis

guineensis Jacq. (oil palm), Ficus elastica

Roxb. ex Hornem. (India rubber), Garcinia

mangostana L. (mangosteen), Hevea

brasiliensis (Willd. ex A.Juss.) Muell.Arg.

(para rubber), Maranta arundinacea L.

Austrobaileya 9(2): 263-278 (2014)

(white arrowroot), Musa acuminata Colla

(banana), Musa textilis Nee (Manilla hemp),

Nicotiana tabacum L. (tobacco), Spondias

cytherea Sonn. (vi-apple), Thea chinensis

Sims (tea), Theobroma cacao L. (cocoa) and

Vanilla planifolia Andrews (vanilla). Possibly

the most significant introductions included

sugar varieties (Griggs 2003, 2011), some

of which were collected by Cowley during

visits to Papua New Guinea in 1892-1893

(van Steenis-Krusemen 1950), and by Henry

Tryon, then Qld Government Entomologist,

in 1895 (Stephens 1981; Griggs 2003, 2011).

A breeding program at the Nursery involving

forms of Caricapapaya L. (pawpaw) resulted

in the development of the ‘Cowleyii’ and

the ‘New Guinea’ varieties - the former a

dioecious type and the latter an hermaphrodite

(Stephens 1981). Not so fortuitous was the

introduction of a current weed of national

significance Prosopis pallida (Humb. &