Editorial Committee

P.I. Forster (editor)

P.D. Bostock (technical advisor)

G.P. Guymer (technical advisor)

Desktop Publishing

Murray Henman

Yvonne Smith

Peter Bostock

Austrobaileya

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

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

This volume (vol. 7, no. 1) published 21 December 2005

Austrobaileya is published once per year.

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

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Email: paul.forster@epa.qld.gov.au

ISSN 0155-4131

Web site: www.epa.qld.gov.au/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.

Contents

Studies in Euphorbiaceae, s.lat. 6. A revision of the genus Poranthera Rudge

( Antidesmeae , Porantherinae ) in Australia

D.A. Halford & R.J.F. Henderson . 1-27

A taxonomic revision of Tarenna Gaertn. and Triflorensia S.T.Reynolds (Rubiaceae:

Ixoroideae. Pavetteae ) in Australia

S. T. Reynolds & PL Forster . 29-55

A taxonomic revision of Actephila Blume (Euphorbiaceae/Phyllanthaceae) in Australia

PI. Forster . 57-98

Studies in Australian Myrsinaceae: Tapeinosperma Hook.f.

B.R. Jackes . 99-110

Notes on the narrow-leaved Ironbarks (Myrtaceae: Eucalyptus subseries Subglaucae )

A.R. Bean . 111-120

Backhousia enata A.J.Ford, Craven & J.Holmes (Myrtaceae), a new species from

north-eastern Queensland

A.J. Ford, LA. Craven & J.J. Brophy . 121-127

Baccania sauropoda D.Meagher (Marchantiophyta: Lepidoziaceae), a new species from

tropical Queensland

D. Meagher . 129-133

Gyrostemon osmus Halford (Gyrostemonaceae), a new species from south-eastern Queensland

D.A. Halford . 135-139

Eucalyptus erosa A.RBean (Myrtaceae), a new stringybark species from central Queensland

A.R. Bean . 141-144

Rediscovery and neotypification of Marsdenia arachnoidea Schltr. (Apocynaceae:

Asclepiadoideae - Marsdenieae), an endangered species from Papua New Guinea

PI. Forster & W. Takeuchi . 145-150

Ludwig Leichhardt’s Australian Plant Collections, 1842-1847

J.L.Dowe . 151-163

Wing structure in seeds of Strangea Meisn. and Stenocarpus R.Br. (Proteaceae)

H. T. Clifford & M.E. Dettmann . 165-170

The rediscovery of Boronia inflexa subsp. grandiflora (Rutaceae)

M.F. Duretto, P. Grimshaw & K. Sparshott . 171-173

New species of Philotheca Rudge (Rutaceae) from Queensland

PI. Forster . 175-181

Papuasian Orchid Studies, 2

P. Ormerod . 183-203

Goodenia elaiosoma I.D.Cowie (Goodeniaceae), a new species from the Top End

of the Northern Territory and a key to the northern species

I. D. Cowie . 205-213

{continued)

Studies in Euphorbiaceae, s.lat. 7. Shonia R.J.F.Hend. & Halford ( Ricinocarpeae,

Ricinocarpinae ), a new Australian endemic genus

D.A. Halford & R.J.F. Henderson .215-228

Cycas scratchleyana F.Muell. (Cycadaceae), a new species record for Queensland

and Australia

P.I. Forster . 229-230

New species of Commersonia J.R.Forst. & G.Forst. (Sterculiaceae) from

Eastern Australia and Vanuatu

G.P. Guymer . 231-250

Orthographic corrections .251

Studies in Euphorbiaceae s.lat. 6. A revision of the genus

Poranthera Rudge ( Antidesmeae , Porantherinae ) in Australia

David A. Halford & Rodney J.E Henderson

Summary

Halford, D A. & Henderson, R.J.F. (2005). Studies in Euphorbiaceae s.lat. 6. A revision of the

genus Poranthera Rudge {Antidesmeae, Porantherinae) in Australia. Austrobaileya 7(1): 1-27.

The genus Poranthera Rudge is revised for Australia. Oreoporanthera Hutch, is accepted as a

taxonomic synonym of Poranthera. Fourteen species of Poranthera are recognised in Australia

and a key is provided for their identification. The following are described here as new: P. dissecta

Halford & R.J.F.Hend., P. florosa Halford & R.J.F.Hend., P. leiosperma Halford & RJ.F.Hend., P.

obovata Halford & R.J.F.Hend. and P. oreophila Halford & R.J.F.Hend. The new combination P.

petalifera (Orchard & J.B.Davies) Halford & R.J.F.Hend., based on Oreoporanthera petalifera

Orchard & J.B.Davies, is made. New species are illustrated, all species are described and distribution

maps provided, and notes on their distribution, habitat and phenology are given. Lectotypes are

chosen for P. microphylla var. intermedia Mull.Arg. and P. arbuscula Sond. Poranthera coerulea

Schwarz is neotypified.

Key Words: Euphorbiaceae, Poranthera , Oreoporanthera , Australian flora, taxonomy,

nomenclature

D A.Halford and R.J.F. Henderson, c/- Queensland Herbarium, Environmental Protection Agency,

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

Introduction

Poranthera Rudge, as accepted here, is a genus

of fifteen species that are annuals or herbaceous

perennials. The genus has representatives in

all Australian states as well as in New Zealand,

with thirteen of the species endemic in Australia

and one endemic in New Zealand. Centres of

diversity are found in south-eastern Australia

and south-western Western Australia. The

species endemic in New Zealand is P. alpina

Cheeseman ex Hook.f. which is not dealt with

here. A description of this species is given in

Cheeseman (1925).

Webster (1994) included Poranthera

Rudge, as well as Oreoporanthera Hutch., in

Euphorbiaceae subfamily Phyllanthoideae

Ascherson, tribe Antidesmeae (Sweet)

Hurusawa, subtribe Porantherinae (Mull.Arg.)

Kohler.

The genus Poranthera was established

by Rudge in 1811 to include a single species,

P. ericifolia Rudge, based on material collected

from Port Jackson, New South Wales (Rudge,

1811). The generic name is derived from Greek

poros, pore, and anthera , anther, in reference

to the species’ anther cells dehiscing by terminal

pores. In his protologue of the genus and

Accepted for publication 30 June 2005

species, Rudge mistakenly described its flowers

as lacking a calyx. He was apparently deceived

by the petaloid nature of their calyx lobes and

was not aware of the presence of small,

inconspicuous petals within the calyx whorl.

Over the subsequent 55 years, another

nine species of Poranthera were described by

various authors. Brongniart (1833) described

P. microphylla and P. corymbosa, Klotzsch

(1848) described P. drummondii, P. ericoides,

P glauca, P. huegelii and P. piceoides and

Sonder (1857) described P. arbuscula while

Baillon (1858) described P. linearoides , the name

of which is an illegitimate synonym of

P. corymbosa.

Later, Baillon (1866) and Muller (1866)

independently accepted Poranthera as

containing only six species and both placed

P. drummondii, P. piceoides and P. arbuscula

in synonymy under P. microphylla, P. ericoides

and P. corymbosa respectively. Muller (Joe. cit.),

however, also recognised seven varieties;

P. corymbosa var. arbuscula (Sond.) Mull.Arg.,

P. corymbosa var. genuina Mull.Arg.

(= P. corymbosa Brongn. var. corymbosa ),

P. microphylla var. diffusa Mull.Arg.,

P. microphylla var. drummondii (Klotzsch)

Miill.Arg., P. microphylla var. genuine/

Miill.Arg. (= P. microphylla Brongn. var.

microphylla ), P. microphylla var. glauca

Miill.Arg. and P. microphylla var. intermedia

Miill.Arg. Seven years later, Bentham (1873), in

his account of Poranthera in Flora

Australiensis, accepted only five species in the

genus, combining P. glauca with P. ericoides.

In the most recent comprehensive work

on Poranthera , Griining (1913) recognised the

five species Bentham enumerated, re-instated

P. drummondii as a species distinct from

P. microphylla , distinguished a number of

varieties in P. microphylla and P. corymbosa

and included P. alpina, a species described by

Hooker (1881) from New Zealand. Griming was

the first to formally divide Poranthera into

subgeneric groups describing Poranthera

subgen. Euporanthera Griining (= Poranthera

subgen. Poranthera ), characterised by having

flowers of both sexes possessing petals and in

terminal racemose or corymbose inflorescences,

and with stipules thin and membraneous, and

Poranthera subgen. Oreoporanthera Griining,

distinguished by its flowers lacking petals,

being solitary and axillary, and with stipules more

or less leathery. Since Griining’s treatment, two

other species have been described, namely

P. triandra (Black 1916) and P. coerulea

(Schwarz 1927).

The genus Oreoporanthera was erected

by Hutchinson in 1969 to accommodate the

species from subalpine communities on the

South Island of New Zealand previously known

as Poranthera alpina Cheeseman ex Hook.f.

(Hutchinson 1969). He distinguished

Oreoporanthera from Poranthera by its

dioecious habit, arrangement of flowers singly

in upper leaf axils, opposite leaves with strongly

recurved margins, and by its flowers which

lacked petals in both sexes and a vestigial ovary

in male flowers. Orchard & Davies (1985) added

a second species, O. petalifera , to this genus.

In their discussion, they noted that their newly

described species, although closely related to

O. alpina , disagreed with Hutchinson’s

description of the genus in having petals in male

and female flowers and a vestigial ovary in male

flowers. The new species described in this paper

as P. oreophila , is similiar to O. alpina and

O. petalifera in its dioecy and strictly opposite

Austrobaileya 7 (1): 1-27 (2005)

leaves with strongly recurved margins.

However, its flowers are borne in short dense

terminal racemes. There is also a high-altitude

form of P. microphylla occurring in the

Australian Alps which resembles P. oreophila

in having opposite leaves with more or less

strongly revolute margins, but it differs from

both those in its monoecy. It is our opinion that

the distinctions used to separate

Oreoporanthera from Poranthera are extremely

tenuous. We therefore concur with Webb et al.

(1988) that the separation of Oreoporanthera

from Poranthera cannot be upheld and that the

species concerned are best placed in a single

genus named Poranthera.

No formal subgeneric classification is

followed in this revision though two informal

groups can be recognised based on habit. These

are either small annuals or small herbaceous

perennials with annual stems dying back to a

more or less woody rootstock (P. microphylla ,

P. dissecta, P. drummondii, P. triandra,

P. coerulea, P. leiosperma, P. petalifera,

P. florosa and P. oreophila ) or comparatively

large annuals or perennials with ± woody stems

(P. ericifolia , P. huegelii, P. corymbosa,

P. obovata and P. ericoides).

Materials and methods

The present study involved examination of

herbarium specimens by both authors, together

with field investigations by the second author

from 1988 to 1992. Altogether, approximately

1100 specimens have been examined and

annotated. These comprise collections from the

following herbaria: AD, BRI, CANB, DNA, HO,

K, LD, MEL, NE, NSW, P and PERTH. The above

acronyms and ones used in the text are those

given by Holmgren et al. (1990). Author

abbreviations follow Brummitt & Powell (1992).

All specimens cited have been examined unless

indicated as unseen n.v. (non visus).

The species treated in the present paper

are listed alphabetically. Descriptions of taxa

were made from dried herbarium specimens,

material preserved in 70% ethanol or dried

material reconstituted by placing in boiling water

for a few minutes. Measurements listed are

based upon the total variation observed in the

herbarium specimens examined. Colour of fresh

Halford & Henderson, Revision of Poranthera

vegetative and floral parts, where given, are

either from herbarium label notes or from

photographs taken by the second author during

field studies. Plant size, habit, flowering and

fruiting times, and habitat data were obtained

from herbarium labels. The morphological data

for this revision were recorded using the DELTA

system (Dallwitz etal. 1993). The distribution

maps were produced with Maplnfo Version 3

and are based on herbarium specimen locality

data.

Taxonomy

Poranthera Rudge, Trans. Linn. Soc. London

10: 301-303 (1811); Poranthera Rudge

subgen. Poranthera , Griming in A. Engler,

Pflanzenr. H.58: 16 (1913); Poranthera

subgen. Euporanthera Griming in A.

Engler, Pflanzenr. Heft 58:16(1913), nom.

inval. Type: P. ericifolia Rudge.

Poranthera subgen. Oreoporanthera Griming

inA. Engler, Pflanzenr. Heft 58:21 (1913).

Type: Poranthera alpina Cheeseman ex

Hook.f.

Oreoporanthera Hutch., Amer. J. Bot. 56(7): 747

(1969). Type: O. alpina (Cheeseman ex

Hook.f.) Hutch., P. alpina Cheeseman ex

Hook.f.

Annuals or herbaceous perennials, monoecious

or rarely dioecious, glabrous. Stems terete, erect,

ascending, procumbent, decumbent or

prostrate, leafy. Leaves stipulate, sessile or

petiolate, alternate or opposite, entire. Stipules

persistent, conspicuous, white, membraneous,

entire or deeply lobed. Flowers pedicellate,

solitary in upper leaf axils or in short dense

terminal racemes that are sometimes umbel-like;

racemes usually with several male and female

flowers, solitary or several in a leafy terminal

corymbose inflorescence, bracteate; bracts leaf-

like. Perianth 2-whorled or sometimes lacking a

corolla in female flowers; calyx with a short tube,

3- or 5(rarely 4)-lobed; lobes imbricate, petaloid,

entire, persistent; petals 3 or 5(rarely 4), free,

always shorter than calyx lobes, persistent.

Glands 4 or 5, antipetalous. Male flowers with

slender pedicels; stamens 3 or 5, antisepalous;

filaments free; anthers 4-celled, dehiscing by

terminal pores; rudimentary ovary present.

Female flowers with mostly stout pedicels;

ovary 3-locular, smooth, glabrous; locules

biovulate; styles 3, free or shortly fused at base,

each spreading, persistent, deeply 2-partite into

linear branches or notched distally. Fruit

capsular, depressed globose, 3- or 6-lobed,

smooth or slightly rugose, glabrous, deshiscing

septicidally into three 2-valved cocci leaving a

persistent columella. Seeds wedge-shaped or

reniform, white or light brown, smooth or

variously sculptured, ecarunculate; endosperm

copious.

A genus of fifteen species in tropical and

temperate Australia and New Zealand. Fourteen

species occur in Australia of which thirteen are

endemic.

Key to Australian species of Poranthera

1. Leaves opposite. 2

Leaves alternate . 7

2. Plants dioecious, perennial with annual stems dying back to rootstock. 3

Plants monoecious, annual or rarely with perennial rootstock. 4

3. Flowers in short dense terminal racemes (N.S.W., Vic.).12. P. oreophila

Flowers solitary in upper leaf axils (Tas.).13. P. petalifera

4. Seeds smooth (W.A., S.A., Vic., N.S.W.).9. P. leiosperma

Seeds tuberculate, granulate or striate. 5

5. Calyces mostly 3-lobed, or if 4- or 5-lobed then with 1 or 2 lobes much

smaller than other 3; seeds granulate (W.A., N.T., S.A.).14. P. triandra

Calyces 5-lobed, rarely 4-lobed; lobes all ± equal in dimensions; seeds

tuberculate or striate. 6

4 Austrobaileya 7 (1): 1-27 (2005)

6. Seeds granulate with translucent granules; stipules deeply dissected (W.A.) . . 3. P. dissecta

Seeds tuberculate with white tubercules; stipules entire or

slightly toothed distally (all States).10. P. microphylla

7. Herbaceous perennials or erect woody annuals to 80(rarely to 150) cm high. 8

Small annuals to 30 cm high. 12

8. Seeds fenestrellate (W.A., S.A.).8. P. huegelii

Seeds reticulate ridged, reticulate-foveate, granulate, verrucate or smooth. 9

9. Leaf laminae >2 cm long (Qld, N.S.W., Vic.).2. P. corymbosa

Leaf laminae < 2 cm long. 10

10. Stipules laciniate (N.S.W.). 5. P. ericifolia

Stipules entire. 11

11. Leaf laminae narrow-obovate, 10-15mmlong, 1.5-3 mm wide(Qld, N.S.W.) . . . ll.P.obovata

Leaf laminae linear, 4-10 mm long, 0.5-1.4 mm wide (W.A.) . 6. P. ericoides

12. Seeds smooth or alveolate . 13

Seeds tuberculate, granulate or striate. 16

13. Seeds smooth (W.A., S.A., Vic., N.S.W.).9. P. leiosperma

Seeds alveolate. 14

14. Flowers in terminal racemes; calyx lobes of male flowers < 1 mm long

(W.A.,N.T.). 1. P. coerulea

Flowers in umbel-like racemes arranged in loose corymbose panicles; calyx

lobes of male flowers 1.1-1.5 mm long. 15

15. Leaf laminae narrow-oblanceolate, cuneate at base (W.A.) .4. P. drummondii

Leaf laminae spathulate, attenuate at base (W.A.).7. P. florosa

16. Stipules deeply lobed (W.A.). 3. P. dissecta

Stipules entire. 17

17. Calyces 3-lobed, rarely 4- or 5-lobed; leaves 2-5.5 mm long; seeds

granulate (W.A., N.T., S.A.).14. P. triandra

Calyces 5-lobed; leaves 6-11 mm long; seeds tuberculate (all States) .... 10. P. microphylla

1. Poranthera coerulea O. Schwarz, Repert.

Spec.Nov.Regni Veg. 24: 87 (1927).

Type: Northern Territory. 8 miles (c. 13 km)

E of Port Darwin, N.T., s.d ., [F.A.K. ] Bleeser

237\ holo: B (destroyed); Thorak’s

Reserve, c. 10 miles ( c. 16 km) SE [of]

Darwin, 21 March 1961, G. Chippendale

7856 (neo [here chosen]: DNA; isoneo:

BRI, NSW).

Illustration : Dunlop etal. (1995: 235, fig. 77),

as Poranthera microphylla.

Monoecious, diffuse to compact annuals to 8

cm high. Stems unbranched or much branched

distally; branchlets erect, smooth, glabrous, 1-

1.5 mm across, with leaf scars obscure. Leaves

subsessile, alternate, widely spaced along

branchlets; stipules white, narrow-triangular,

1.5-2 mm long, entire or erose; laminae narrow-

obovate to obovate, 13-27 mm long, 3-8 mm

wide, flat, smooth and glabrous adaxially and

abaxially, discolorous; midrib impressed

proximally adaxially, prominent abaxially; base

attenuate; apex obtuse to acute. Flowers in short

Halford & Henderson, Revision of Poranthera

dense terminal racemes; racemes solitary or

rarely several in a leafy terminal corymb;

peduncles up to 6 mm long; rachis up to 20 mm

long; bracts narrow-obovate, 2-5 mm long, 0.7-

1 mm wide, acute. Male flowers with pedicels

0.8-1.3 mm long; calyx lobes 5, white or pink,

obovate, 0.6-0.9 mm long, 0.4-0.5 mm wide,

concavo-convex, rounded at apex; petals 5,

white, erect, incurved distally, oblanceolate, 0.3-

0.4 mm long, up to 0.1 mm wide, obtuse, with

margins entire; glands ovate, up to 0.1 mm wide;

stamens 5; filaments 0.7-0.8 mm long, slightly

incurved; anthers c. 0.1 mm long; rudimentary

ovary 3-lobed; lobes clavate, c. 0.4 mm long.

Female flowers with pedicels 0.6-1 mm long,

extending to c. 5 mm long in fruit; calyx lobes 5,

white or pale pink, oblong, 1.1-1.3 mm long,

0.5-0.6 mm wide, ± flat, obtuse at apex; petals 5,

white, linear-oblong, c. 0.3 mm long, c. 0.1 mm

wide, obtuse, with margins entire; ovary

depressed globose, 1.5-1.9 mm across, deeply

6-lobed, emarginate distally, ± smooth; styles

each 0.5-0.7 mm long, deeply 2-partite, slender.

Capsules depressed globose, 1.9-2.1 mm

across, 0.9-1.1 mm long, shallowly 3 lobed,

emarginate and slightly rugose distally. Seeds

reniform, 0.6-0.7 mm long, 0.4-0.5 mm wide, 0.5-

0.6 mm across; testa alveolate, brown.

Additional specimens examined : Western Australia.

Mt Elizabeth track to Munja, Jul 1996, Kenneally

11815 (PERTH). Northern Territory, c. 40 km NW

of Jabiru, Mar 1981, Craven 7691 & Whitbread { CANB,

MEL); Kapalga, Kakadu NP, Mar 1994, Egan 3306

(BRI); Port Darwin, s.d, Holtze 537 (MEL); Melville

Island, Apr 1986, Johnson 4103 (BRI); Yirrkala, Sep

1948, Specht 1020 (BRI); 4 km ESE of Jabiru, E of

Ranger Plant, Apr 1980, Telford 7535 & Wrigiey

(CANB).

Distribution and habitat : Poranthera

coerulea has a disjunct distribution. It occurs

from the Darwin area east to Yirrkala, Northern

Territory, with one isolated population in the

Kimberley region, Western Australia. It is

recorded as growing in open forest or woodland

communities sometimes with a grassy

understorey. The soils are sandy with ironstone

gravel or lateritic red earths, in damp areas on

plains or along damp creek beds. Map 1.

Phenology : Flowers and fruits have been

collected in March, April, July and September.

Affinities : Poranthera coerulea is easily

distinguished from P. microphylla, with which

it has been confused, by having a generally

more robust habit, an alveolate seed coat and

generally longer and broader leaves.

Typification: No type material of the name

Poranthera coerulea has been located by us.

Schwarz (1927) published the name P. coerulea

based on a specimen at B collected in the Darwin

area by F. A.K. Blesser labelled “Port Darwin, 8

miles E (Bleeser no. 237)”. This type is believed

to have been destroyed during the Second

World War (McKee 1963). According to

Orchard (1999), Bleeser also sent specimens

from the Northern Territory to K, MEL and N SW.

No duplicates of the above Bleeser collection

have been located in material of Poranthera on

loan from K, and searches in MEL and NSW

(by the first author) have also been

unsuccessful. The collection Chippendale

7856 (DNA) from approximately 16 km south

east of Darwin is therefore chosen here as

neotype of P. coerulea O.Schwarz. This

collection agrees with the protologue

description except that its leaves are slightly

shorter in length than what Schwarz recorded

for the species.

2. Poranthera corymbosa Brongn., Ann. Sci.

Nat. (Paris) 29: 385 (1833); Poranthera

corymbosa Brongn. var. corymbosa ,

Mull. Arg. in A.DC., Prodr. 15(2):

192(1866); Poranthera corymbosa var.

genuina Mull.Arg. in A.DC., Prodr. 15(2):

192 (1866), nom. inval.; Poranthera

corymbosa var. linarioides Griming in A.

Engler, Pflanzenr. Heft 58:19(1913), nom.

inval;Poranthera linearoides Baill.,

Etude Euphorb. 574, Atlas 45/6, t. 25, figs.

1-9 (1858), nom. illeg. Type: [Australia.]

Port Jackson, s.d.. [C.] Gaudichaud 112

(lecto [here chosen]: P 152752).

Porantheraarbuscula Sond., Linnaea2S. 567

(1856); Poranthera corymbosa var.

arbuscula (Sond.) Mull .Arg. in A. DC.,

Prodr. 15(2): 192(1866). Type: [New South

Wales.] FI. Novae Holl, s.d ., Sieber 116

(lecto [here chosen]: MEL2062902 (ex

herb. Sonder); isolecto: MEL2062901;

2065789).

? Poranthera corymbosa var. sparsifolia

Griming in A. Engler, Pflanzenr. Heft 58:

19 (1913). Type citation: “Neusudwales;

Parramatta (F. Mueller!). Queensland:

Sandy Island (ohne Sammlernamen!).”

(syntypes: n.v.).

Illustrations : Griming (1913:17, fig. 4A-E), as

Poranthera corymbosa var. arbuscula;

James & Harden (1990:394); Jeanes(1999:

73,fig. lie).

Monoecious, erect annuals or herbaceous

perennials to 80 (rarely to 150) cm high. Stems

sparingly branched; branchlets erect, smooth,

glabrous, up to 6 mm across, with leaf scars

prominent. Leaves sessile, alternate, crowded;

stipules white, narrow-triangular, 2.3-3.8 mm

long, entire; laminae linear or narrow-

oblanceolate, (15-)20-90 mm long, 0.3-6 mm

wide, flat with margins recurved to revolute,

smooth and glabrous adaxially and abaxially,

discolorous; midrib impressed proximally

adaxially, prominent abaxially; base attenuate

or cuneate; apex acute to obtuse. Flowers in

short dense terminal umbel-like racemes;

racemes in a lax leafy terminal corymb;

peduncles up to 100 mm long; rachis 3-5 mm

long; bracts narrow-obovate, 1.5-5 mm long,

0.3-1.3 mm wide, obtuse. Male flowers with

pedicels 1-2.7 mm long; calyx lobes 5, white,

obovate, 1.5-2.7 mm long, 0.7-1 mm wide, ± flat,

rounded at apex; petals 5, white, erect, linear to

oblanceolate, 0.7-0.9 mm long, 0.1-0.2 mm wide,

acute, with margins entire; glands 2-lobed, 0.2-

0.4 mm wide; stamens 5; filaments 1.3-1.8 mm

long, slightly incurved; anthers 0.2-0.3 mm

long; rudimentary ovary 3-lobed; lobes clavate,

0.5-0.8 mm long. Female flowers with pedicels

0.8-2.5 mm long, extending to c. 5 mm in fruit;

calyx lobes 5, white, obovate, 1.6-2.2 mm long,

0.8-1.3 mm wide, slightly concavo-convex,

rounded at apex; petals 5, white, oblong, 0.5-

0.8 mm long, 0.1-0.3 mm wide, obtuse, with

margins entire; ovary depressed globose, 1.1-

1.6 mm across, shallowly 6-lobed, slightly

emarginate distally, smooth; styles each 0.9-

1.5 mm long, deeply 2-partite, slender. Capsules

depressed globose, 2.8-3.1 mm across, 1.6-2

mm long, shallowly 3-lobed, emarginate distally,

± smooth. Seeds wedge-shaped, 1.1-1.3 mm

long, 0.8-0.9 mm wide, 0.8-0.9 mm across; testa

reticulate, white.

Selected specimens (from 140 examined ):

Queensland. Darling Downs District: Mt Janet Road,

Passchendaele State Forest, Oct 1997, Bean 12478

Austrobaileya 7 (1): 1-27 (2005)

(BRI); 15.7 km SW of Stanthorpe, portion 87, Stalling

Lane, Nov 1994, Halford Q2221 (BRI). Moreton

District: Mount Ernest, McPherson Range, Sep 1990,

Forster PIF7397 et al. (BRI, CANB, MEL, NSW);

Stags Head, E ridge 2 km E of Mt Clunie, 32 km WSW

of Rathdowney, Sep 1997, Halford Q3546 (BRI). New

South Wales. Carters Creek crossing, c. 16 km NNW

of Nelligen, Feb 1980, Adams 3450 & Gray (CANB,

NSW); 3 km SW of the Old Hut Creek crossing of

Nethercote Road, Nullica State Forest, Oct 1985,

Albrecht 2050 (MEL, NSW); Blackheath Glen, Blue

Mountains, Nov 1969, Burgess s.n. (CANB

[CBG033397]); W side of Cox’s Gap, E of Bylong, Oct

1979, Chapman 1553 (BRI, CANB, MEL, NSW); 2.4

km along the Flagstone Creek track from the Gulf

Road, c. 19 km (direct) just W of N of Emmaville, Oct

1990, Coveny 14619 et al. (BRI, CANB, NSW);

Wombeyan Caves along the road to Mittagong, Oct

1988, Greuter 21387 (NSW); Edinburgh Castle, 8 km

SSE of Woodenbong, Dec 1992, Halford Q1554A (BRI,

NSW); c. 18 km NW of Sandy Hollow heading towards

Merriwa, Sep 1990, Henderson & Turpin H3413 (BRI);

Torrington, New England, Sep 1969, Jones 4068

(CANB); Turpentine Range, c. 10 miles (c. 16 km) W

of Tomerong, Sep 1965, McGillivray 1485 (NSW); c.

13 km W of Tomerong in the Sassafras Range, between

Braidwood and Nowra, Nov 1968, Pullen 4333 (CANB,

NSW); Craters Creek, Currowan State Forest, WNW

of Batemans Bay, Dec 1973, Pullen & Story 8732

(BRI, CANB, NSW); 17 km E of Tianjara Falls along

Turpentine Road, Aug 1984, Stewart 96 et al. (CANB,

MEL). Victoria. Brisbane Ranges NP, Mt Wallace -

Bacchus Marsh Road junction of Aeroplane Road, Oct

1977, Beauglehole ACB56761 & Errey (MEL); Pyrete

Ranges, Oct 1982, Kemp s.n. (MEL 628611); Hard to

Seed Track, 9.5 km SW of Genoa Peak, Oct 1983,

Walsh 1226 (MEL).

Distribution and habitat : Poranthera

corymbosa occurs in south-eastern Australia

from near Stanthorpe and the McPherson

Range, in south-eastern Queensland, through

New South Wales to East Gippsland, Victoria,

with disjunct records from the Grampian, Pyrete

and Brisbane Ranges, in western Victoria. It is

recorded growing in heath, open eucalypt

woodland or forest communities in mostly well-

drained sandy soils overlying a variety of

substrates. Map 2.

Phenology : Flowers have been collected

throughout the year, but mainly from September

to November, fruits from September to January.

Typification: In his protologue of/ 3 , corymbosa,

Brongniart cited two collections, namely “ad

Port-Jackson in Nova-Hollandia ( Gaudichaud

et d’Urville in herb. Mus. Par.)”. These

Gaudichaud and Dumont d’Urville collections

Halford & Henderson, Revision of Poranthera

have been located at P (numbered 152752 and

152751 respectively). The Gaudichaud specimen

is selected here as lectotype of Poranthera

corymbosa Brongn. as it is part of the original

material, agrees with the protologue and is more

than likely the material illustrated in the

protologue.

In his protologue of P. arbuscula, Sonder

cited two collections, namely “Nov. Holland,

austral. (Dr F. Mueller)” and “SieberHerb. Nov.

Holl. no. 116 ex parte”. In MEL, there are three

sheets of Sieber’s collection, numbered 2062901,

2065789 and 2062902, and one specimen labelled

“P. arbuscula, P. cicabricosa, Nov. Holla. Austr.

Ferd. Mueller”, numbered 2062936. They are all

from Sonder’s herbarium. There is also a

specimen in MEL numbered 694305 which is

labelled “Poranthera cicabricosa f mueller (1847)

Encounter Bay [SA]” which matches the

Mueller collection in the Sonder herbarium. We

believe it to be a duplicate of that specimen.

The Mueller and Sieber collections Sonder cited

actually comprise two distinct taxa. The Sieber

collection is conspecific with P. corymbosa ,

while the Mueller collection is undoubtedly

conspecific with P. huegelii as circumscribed

here. The Sieber specimen numbered 2062902

is selected here as lectotype of P. arbuscula

Sond. as it is part of the original material, agrees

with the protologue and is possibly annotated

P. arbuscula by Sonder himself.

Notes: The plant Griming named P. corymbosa

var. sparsifolia is probably conspecific with

P. corymbosa as treated here but we have been

unable to examine the syntypes to verify this.

However, from Gruning’s description it is

reasonable to assume the two taxa are

conspecific and not worthy of formal

recognition. The supposed locality of collection

of the syntype “Queensland, Sandy Island” is

somewhat unusual for P. corymbosa as no other

collections of this species we know of have been

made on any of the coastal islands of

Queensland. However, the locality of collection

given for the other syntype, “Parramatta, F.

Mueller”, is within the presently known

geographical range of P. corymbosa.

Poranthera corymbosa is somewhat

variable in leaf size and plant stature. However,

there seems no justification for recognising

varieties within this species. Some small forms

can be confused with P. ericifolia but can be

distinguished from that by having entire

stipules and generally more open, lax

inflorescences.

3. Poranthera dissecta Halford & R. J.F.Hend.,

sp. nov. quoad habitu P. triandrae

J.M.Black similis autem foliis anguste

ovatis vel oblongis, stipulis profunde

dissectis, floribus calycibus 5-lobis,

pagina seminum minute striata differt.

Olim collectiones P. dissectae ad

P. microphyllam Brongn. relatae fuerunt

sed P. dissecta a P. microphylla stipulis

profunde dissectis, floribus capsulisque

minoribus et pagina seminum striata

differt. Typus: Western Australia. 22 km

ESE of Lake King, 8 Aug 1968, P G. Wilson

6892 (holo: PERTH).

Monoecious, prostrate annuals. Stems much

branched; branchlets smooth, glabrous, up to

0.5 mm across, with leaf scars obscure. Leaves

sessile, opposite or alternate, widely spaced

along branchlets; stipules white, narrow-

triangular, 0.8-1.1 mm long, deeply 2^1-lobed;

laminae narrow-obovate or elliptic, 2-3.6 mm

long, 0.9-1.5 mm wide, flat, smooth and glabrous

adaxially and abaxially, concolorous; midrib

obscure adaxially, slightly raised proximally

abaxially; base attenuate; apex obtuse to

rounded. Flowers in short dense terminal umbel¬

like racemes; racemes solitary; rachis c. 1 mm

long; bracts narrow-obovate, 0.9—1.5 mm long,

0.2-0.4 mm wide, obtuse to rounded. Male

flowers with pedicels 0.5-1.1 mm long; calyx

lobes 5 (or sometimes 4), colour when living

unknown, narrow-obovate or oblong, 0.4-0.5

mm long, 0.1-0.2 mm wide, concave-convex,

acute to obtuse at apex; petals 5, colour when

living unknown, erect, ovate, c. 0.1 mm long, up

to 0.05 mm wide, acute, with margins toothed

distally; glands obscure; stamens 3, filaments

up to 0.2 mm long, straight; anthers up to 0.1

mm long; rudimentary ovary a minute,

hemispherical dome. Female flowers with

pedicels 0.3-0.6 mm long, extending to 1.5 mm

long in fruit; calyx lobes 5, colour when living

unknown, narrow-ovate or oblong, 0.5-0.6 mm

long, 0.1-0.2 mm wide, concavo-convex, acute

at apex; petals obscure or absent; ovary

depressed globose, 0.4-0.6 mm across, deeply

6-lobed, emarginate distally, papillose; styles

each 0.1-0.4 mm long, shortly divided distally,

slender. Capsules depressed globose, 1-1.3 mm

across, 0.6-0.7 mm long, prominently 6-lobed,

emarginate distally, rugose. Seeds wedge-

shaped, c. 0.4 mm long, c. 0.3 mm wide, to c. 0.3

mm across; testa striate with translucent ridges.

Fig. 1

Additional specimens examined : Western Australia.

c. 45 km N of Stokes Inlet, Oct 1968, Eichler 20286

(CANB); Victoria Desert, camp 53, Sep 1891, Helms

s.n. (NSW455348); Karolin [Karlgarin], Sep 1891,

Helms s.n. (MEL2065544); Mt Andrew, c. 120 km SE

of Norseman, Sep 1980, Newbey 7595 (PERTH).

Distribution and habitat: Poranthera dissecta

occurs in scattered populations in southern

Western Australia, in an area more or less

bounded by Norseman, Ravensthorpe and

Karlgarin. It is recorded as growing in well-

drained granitic loamy sand near granite

outcrops. Map 3.

Phenology : Flowers and fruits have been

collected from August to October.

Affinities: Poranthera dissecta resembles

P. triandra in stature but differs from that by

having narrow-ovate or oblong leaves, deeply

dissected stipules, a 5-lobed calyx and a

minutely striate seed surface. Collections of

P. dissecta have been referred in the past to

P. microphylla. Poranthera dissecta differs

from P. microphylla by its deeply dissected

stipules, smaller flowers and capsules and its

striate seed surface.

Etymology: The specific epithet is derived from

Latin, dissectus, dissected, in reference to the

species’ deeply dissected stipules.

4. Poranthera drummondii Klotzsch in J.G.C.

Lehmann, PI. Preiss. 2: 231 (1848);

Poranthera microphylla var. drummondii

(Klotzsch) Miill.Arg. in A.DC., Prodr.

15(2): 193 (1866). Type: [Western

Australia.] In sands of the Swan River, in

1839, J. Drummond s.n. (syn: n.v );

[Western Australia.] Rotenest (Rottnest

Island), 21 Aug 1839, L. Preiss 2048 (syn:

LD 99/018-0884; isosyn: B (ex herb. L.C.

Treviranus), G-DC n.v. (IDC microfiche

800-73 2453: II.4); MEL 2062911,2062926

(ex herb. Sonder)).

Austrobaileya 7 (1): 1-27 (2005)

Monoecious, compact annuals to 15 cm high.

Stems sparingly or much branched; branchlets

erect or ascending, ± smooth, glabrous, up to

1.1 mm across, with leaf scars obscure. Leaves

sessile, alternate, widely spaced along

branchlets; stipules white, narrow-triangular,

1.3-2.5 mm long, entire or erose; laminae narrow-

oblanceolate, 9-31 mm long, 1.4-3.5 mm wide,

flat with margins slightly recurved, smooth and

glabrous adaxially and abaxially, discolorous;

midrib obscure adaxially, prominent or slightly

raised proximally abaxially; base cuneate; apex

rounded to obtuse. Flowers in short dense

terminal umbel-like racemes; racemes solitary

or several in a leafy terminal corymb; peduncles

up to 25 mm long; rachis up to 3 mm long; bracts

narrow-obovate, 2-6 mm long, 0.5-0.9 mm wide,

obtuse to rounded. Male flowers with pedicels

0.8-2.2 mm long; calyx lobes 5, white, narrow-

obovate, 1.4-1.5 mm long, 0.4-0.8 mm wide, ±

flat, rounded at apex; petals 5, white, erect,

narrow-ovate or oblanceolate, 0.1-0.8 mm long,

up to 0.1 mm wide, acute, with margins entire;

glands clavate, c. 0.1 mm wide; stamens 5;

filaments 0.6-1 mm long, incurved; anthers c.

0.1 mm long; rudimentary ovary 3-lobed; lobes

clavate, 0.2-0.6 mm long. Female flowers with

pedicels 0.8-1.5 mm long, extending to 4.5 mm

in fruit; calyx lobes 5, white, narrow-obovate to

oblong, 1-1.5 mm long, 0.4-0.7 mm wide, ±

concavo-convex, rounded at apex; petals 5,

white, narrow-ovate, 0.2-0.6 mm long, 0.1-0.2

mm wide, acute, with margins entire or erose;

ovary depressed globose, 0.9-1 mm across,

shallowly to deeply 6-lobed, emarginate distally,

papillose; styles each 0.5-0.8 mm long, 2-partite,

slender. Capsules depressed globose, 1.7-1.9

mm across, 0.5-0.6 mm long, shallowly 6-lobed,

emarginate distally, somewhat rugose. Seeds

wedge-shaped, 0.6-0.7 mm long, 0.5-0.7 mm

wide, 0.3-0.5 mm across; testa alveolate, grey-

white.

Selected specimens (from 30 examined ): Western

Australia. Guildford, Perth, Nov 1901, Andrews s.n.

(PERTH); Dillon Bay, Oct 1963, Aplin 2749 (PERTH);

Cape Naturaliste, Oct 1978, Aplin 6444 (PERTH);

Cape Leeuwin, S of Augusta, Sep 1965, Beauglehole

ACB12508 (NSW, PERTH); S end of Garden Island,

Oct 1978, Cranfield 28 (PERTH); c. 3 km S of

homestead, Dirk Hartog Island, Sep 1972, George 11348

(PERTH); S outskirts of Kalbarri township, near

Telecom tower, Sep 1988, Henderson H3148A (BRI);

c. 10 km SW of Lake Indoon on Leeman Road, Sep

1977, Hnatiuk 770965 (PERTH); Bremer Bay, Sep

1974, Newbey 4385 (PERTH); Hamelin Bay, Oct 1953,

Halford & Henderson, Revision of Poranthera

Fig. 1 . Poranthera dissecta. A. whole plant xl. B. inflorescence x8. C. male flower x32. D. branchlet with leaf and

stipules xl2. E. fruit from above *32. F. fruit from side x32. A from Wilson 6892 (PERTH); B-F from Eichler

20286 (CANB). Del. W. Smith.

Royce 4644 (PERTH); Nambung NP, Cervantes, Oct

1971, Royce 9780 (PERTH); Yanchep NP, Nov 1965,

Scrymgeour 160 (PERTH); Helena Valley, Sep 1977,

Seabrook 257 (PERTH); Champion Bay, 1889, Sewell

s.n. (MEL2065488); c. 10.7 km from Coorow, along

road to Marchagee, Oct 1983, Short 2202 (MEL);

Rottnest Island, Aug 1956, Storr 172 (PERTH); by

scenic drive just NE of Cape Leeuwin, Nov 1982, Strid

21403 (PERTH).

Distribution and habitat: Poranthera

drummondii is confined to the coastal and

subcoastal districts of south-west Western

Australia, from Dirt Hartog Island southward

to Augusta and eastward to Dillon Bay near

Albany. It is recorded as growing in heathland,

shrubland or open woodland communities in

mostly sandy soils. Map 4.

Phenology: Flowers have been collected from

August to November, fruits from September to

November.

Affinities: Muller (1866) reduced this species

to a variety of P. microphylla , while Bentham

(1873) included it in P. microphylla without

giving it formal recognition. Although

P. drummondii is somewhat similar to the

Western Australian populations of

P. microphylla , it can easily be distinguished

from that by its larger flowers and alveolate seed

surface.

Notes : The collection Henderson H3148B (BRI)

has seeds that are more or less reniform with a

faintly reticulate surface. This gathering was

part of the collection Henderson H3148A (BRI)

which is typical of P. drummondii with wedge-

shaped seeds and an alveolate seed surface.

More collections and further research are

required to determine if this specimen represents

a form requiring formal recognition.

5. Poranthera ericifolia Rudge, Trans. Linn.

Soc. London 10:301-303 (1811). Type: [New

South Wales.] Port Jackson, s.d., [without

collector] (holo: BM n.v. (transparency at

BRI)).

Illustration : James & Harden (1990: 394).

Monoecious, erect, herbaceous perennials 10-

30 cm high. Stems sparingly to much branched

from base; branchlets erect, smooth or papillose,

glabrous, 2-3 mm across, with leaf scars

prominent. Leaves sessile, alternate, crowded;

stipules white, narrow-triangular, 1.4-2 mm long,

laciniate; laminae linear or narrow-oblanceolate,

8-15 mm long, 0.6-2 mm wide, flat to rounded

with margins recurved to revolute, smooth and

glabrous adaxially and abaxially, discolorous;

midrib impressed proximally adaxially, prominent

abaxially; base attenuate or cuneate; apex

obtuse to subacute. Flowers in a short dense

terminal umbel-like racemes; racemes in a

compact leafy terminal corymb; peduncles 10-

20 mm long; rachis 2-5 mm long; bracts narrow-

obovate, 2-4 mm long, 0.5-0.9 mm wide, obtuse.

Male flowers with pedicels 1.5-2 mm long; calyx

lobes 5, pink or white, obovate, 1.7-2.2 mm long,

0.8-1 mm wide, ± concavo-convex, rounded at

apex; petals 5, white, erect, recurved distally,

linear to oblanceolate, 0.9-1.1 mm long, 0.2-0.3

mm wide, acute or obtuse, with margins entire;

glands 2-lobed, 0.2-0.4 mm wide; stamens 5;

filaments 1.8-2 mm long, slightly incurved;

anthers 0.1-0.2 mm long; rudimentary ovary 3-

lobed; lobes clavate, 0.7-1 mm long. Female

flowers with pedicels 0.9-2.2 mm long,

extending to c. 3 mm long in fruit; calyx lobes 5,

white or pink, oblong, 1.5-2 mm long, 0.6-0.8

mm wide, ± concavo-convex, rounded at apex;

petals 5, white, linear-oblong, 0.5-0.9 mm long,

0.1-0.2 mm wide, acute, with margins entire;

ovary depressed globose, 1-1.5 mm across,

deeply 6-lobed, emarginate distally, smooth;

styles each 0.9-1.4 mm long, deeply 2-partite,

Austrobaileya 7 (1): 1-27 (2005)

slender. Capsules depressed globose, 24-2.9

mm across, 1.4-1.6 mm long, shallowly 6-lobed,

emarginate distally, papillose. Seeds wedge-

shaped, 0.9-1.1 mm long, 0.7-0.9 mm wide, 0.8-

0.9 mm across; testa reticulate, white.

Selected specimens {from 60 examined ): New South

Wales. Gospers Mountain, Apr 1983, Benson & Keith

1416 (NSW); near Wyong, Oct 1948, Burbidge 2869

(CANB); Wentworth Falls, King’s Tableland, Nov 1962,

Burgess s.n. (CANB [CBG001710]); Mt Manning

(Bucketty), Sep 1963, Burgess s.n. (CANB

[CBG001619]); Victoria Falls Road, Blue Mountains,

Dec 1947, Constable s.n. (NSW); Belrose, Nov 1981,

Coveny 11055 & Hind (NSW); Kioloa State Forest,

Oct 1966, Evans 2515 (NSW); near Sublime Point,

Leura, Dec 1961, Goode 528 (NSW); on road to Mt

Hay, c. 4 km N of Leura, Blue Mountains NP, Oct

1990, Henderson & Turpin H3419 (BRI); Comleroy

Road near entrance to Wollemi NP, Oct 1987, Hind

5393 et al. (BRI, NSW); 1.8 km along Forest Road

towards Currawong, Sep 1995, Jobson 3785 (MEL);

“Eagle Rock”, N of Mogo Creek Road, 20 km S of

Bucketty towards St Albans, Oct 1996, Jobson 4422

(NSW); Mt Kindarun, 14 miles (c. 22 km) NNW of

Putty, Sep 1959, Johnson & Constable s.n. (NSW);

Walls lookout. Pierces Pass, near Mt Banks, Dec 1970,

Lassak s.n. (NSW); Mt Victoria, Nov-Dee 1889,

Maiden s.n. (NSW); “Big Hill”, Mittagong, Dec 1960,

Mowle 47 (CANB); Tarougra Forest Road, 2 Ion E of

Bodalla along Potato Point Road, Oct 1986, Mullins

702 (CANB); northern Budawang Range, Camping Rock

Creek, S of Sassafras, Oct 1971, Pulley & Telford BR2 70

(CANB); 3 km N of Karuah on Pacific Hwy, Aug 1967,

Telford 211 (CANB); near old sawmill, 10 miles (c. 16

km) S of Sassafras, Oct 1965, Whaite 2916 (NSW).

Distribution and habitat: Poranthera

ericifolia is confined to coastal and subcoastal

areas of New South Wales, from Weston

southwards to Nowra. It is recorded as growing

in heathland, shrubland, and eucalypt woodland

or open forest communities usually in sandy

soils derived from sandstone. Map 5.

Phenology: Flowers have been collected in

April and from July to December, fruits in

February, March and from September to

December.

Notes: Poranthera ericifolia has been

confused with P. corymbosa but is easily

distinguished from that by having laciniate

stipules, smaller leaves and shorter and more

compact habit.

6. Poranthera ericoides Klotzsch in J.G.C.

Lehmann, PI. Preiss. 2: 231 (1848);

Poranthera ericoides Klotzsch var.

ericoides , Domin, Vestn. Krai. Ceske

Halford & Henderson, Revision of Poranthera

Spolecn. Nauk. Tr. Mat.-Prir. {Mem. Soc.

Roy. Sci. Boheme, Prague) 1921/2(2): 58

(1923); Poranthera ericoides var. typica

Domin, Vestn. Krai. Ceske Spolecn. Nauk.

Tr. Mat.-Prir. (Mem. Soc. Roy. Sci.

Boheme, Prague) 1921/2(2): 58 (1923),

nom. inval. Type: [Western Australia.]

Hay, 7 Nov 1840, L. Preiss 2050 (holo: LD

99/018-0885; iso: MEL 2062927 (exherb.

Sonder)).

Poranthera piceoides Klotzsch in J.G.C.

Lehmann, PI. Preiss. 2:232 (1848). Type:

[Western Australia.] In sand in forest near

Bull’s Creek, Perth, Nov 1841, L. Preiss

2044 (syn: LD 99/018-0889; isosyn: G-DC

n.v. (microficheIDC 800-73.2453:1.7); MEL

617990, 2062929 (ex herb. Sonder);

[Western Australia.] In sand in the shade

in forest near Pine-Aple [Pineapple inn,

near Perth, fide Marchant 1990], 14 Sep

1830, L. Preiss 1227 (syn: LD 99/018-0888;

isosyn: G-DC n.v. (microfiche IDC 800-73.

2453:1.7 (top element)); MEL 2062908 (ex

herb. Sonder), 1617991).

Monoecious, compact, herbaceous perennials

to 15 cm high. Stems much branched at base;

branchlets erect or procumbent, smooth or

papillose, glabrous, 0.6-2.5 mm across, with leaf

scars prominent. Leaves ± sessile, alternate,

crowded on branchlets; stipules white, narrow-

triangular, 0.6-1.1 mm long, entire; laminae

linear, 4-10 mm long, 0.5-1.4 mm wide, rounded

with margins recurved to midrib, smooth and

glabrous adaxially and abaxially, concolorous;

midrib obscure adaxially, prominent abaxially;

base cuneate; apex obtuse. Flowers in short

dense terminal umbel-like racemes; racemes

solitary or several in a leafy compact terminal

corymb; peduncles up to 15 mm long; rachis up

to 4 mm long; bracts narrow-obovate, 2-3.5 mm

long, 0.6-0.8 mm wide, acute to obtuse. Male

flowers with pedicels 1-1.7 mm long; calyx lobes

5, white or sometimes with pinkish hue, oblong

or obovate, 1.2-2 mm long, 0.5-0.8 mm wide, ±

flat, rounded at apex; petals 5, white, erect,

oblanceolate, 0.5-0.9 mm long, 0.2-0.4 mm wide,

acute, obtuse to rounded, with margins entire;

glands 2-lobed, 0.1-0.3 mm wide; stamens 5,

filaments 1.3-2 mm long, slightly incurved;

anthers c. 0.2 mm long; rudimentary ovary 3-

lobed; lobes clavate, 0.6-0.8 mm long. Female

flowers with pedicels 0.5-1.5 mm long,

extending to c. 5 mm long in fruit; calyx lobes 5,

white sometimes with a pinkish hue, oblong,

1.3-1.8 mm long, 0.4—0.8 mm wide, ± flat, rounded

at apex; petals 5, white, linear-oblanceolate, 0.7-

0.8 mm long, 0.3-0.4 mm wide, acute to obtuse,

with margins entire; ovary globose, 1-1.7 mm

across, shallowly to deeply 6-lobed, emarginate

distally, smooth or papillose; styles each 1.1-

1.5 mm long, deeply 2-partite, slender. Capsules

depressed globose, 2-2.5 mm across, c. 1.5 mm

long, prominently to shallowly 6-lobed,

emarginate distally, rugose. Seeds wedge-

shaped or rarely reniform, 1-1.2 mm long, 0.6-

0.8 mm wide, 0.6-0.8 mm across; testa smooth

and white or rarely verrucate and pale brown.

Selected specimens (from 35 examined): Western

Australia. Between Kukerin and Lake Grace, Nov

1931, Blackall 1351 (PERTH); near Yanchep, 2 km

from Wanneroo Road towards Gingin, Oct 1981, Craven

6964 (MEL); Wanneroo, Nov 1976, Demarz D6219

(PERTH); Gnangarra, Oct 1945, Gardner s.n.

(PERTH); 18 miles ( c. 29 km) E of Cranbrook, on

Chester Pass Road, Nov 1959, George 398 (PERTH);

3 miles ( c. 5 km) N of Qualeup homestead, Nov 1960,

George 1780 (PERTH); Albany Hwy, N [of] Cranbrook

turnoff, Nov 1961, George 3097 (PERTH); near

junction of Red Gum Pass road with Salt River Road,

near NW boundary of Stirling Range NP, Sep 1988,

Henderson H3197 (BRI); 4.5 km W of Tarin Rock,

Sep 1976, Hnatiuk 761350 (PERTH); between Toll’s

Creek and Solomon’s Well, Oct 1902, Morrison s.n.

(PERTH); 1 mile (c. 1.6 km) E of Tambellup, Oct

1963, Newbey 1155 (PERTH); 7 miles (c. 11 km) S of

Wagin, Oct 1962, Phillips s.n. (CANB [CBG024913],

NSW); near Gingin, and 15 miles ( c. 24 km) from

Muchea, Sep 1968, Phillips 1855 (CANB, NSW); 5

km E of Tincurrin, Oct 1982, Rechinger 59109

(PERTH); Moore River NP, Oct 1971, Royce 9491

(PERTH); Talbot Brook, York, Sep 1921, Sargent s.n.

(PERTH); 3 miles (c. 5 km) S of Gingin, Sep 1966,

Scrymgeour 1308 (PERTH); along No. 2 Rabbit Proof

Fence, c. 40 km SSE of Jerramungup-Ravensthorpe

road and c. 25 km N of Bremer Bay, Oct 1966, Wilson

4353 (PERTH); 20 miles (c. 32 km) from Pingelly

towards Wandering, on north road, Oct 1968, Wrigley

s.n. (CANB [CBG027813]); 11 miles (c. 18 km) from

Kukerin toward Lake Grace, Nov 1968, Wrigley WA/68

5732 (CANB).

Distribution and habitat : Poranthera

ericoides is confined to south-western Western

Australia from Winchester south east to near

Ravensthorpe. It is recorded as growing in

closed heath or mallee communities with a

heath-like understorey in sand or sandy loam

soils sometimes with gravel in the profile.

Map 6.

Phenology : Flowers have been collected from

September to November, fruits in November.

Affinities: Poranthera ericoides has been

confused in the past with P. huegelii.

Poranthera ericoides differs from that in the

texture of its seed surface and in usually having

a more compact habit.

Notes: The inflorescence of P. ericoides is

typically a more or less compact corymb, but

there is a variant which has a more open and lax

inflorescence somewhat resembling those in

some forms of P. huegelii. This variation is

particularly prevalent in material from the

northern end of the range of P. ericoides , e.g.

Craven 6964 (MEL), Phillips 1855 (CANB),

Royce 9491 (PERTH) and Scrymgeour 1308

(PERTH).

There is a collection at PERTH ( Coleby -

Williams 312 ) from near Winchester which is c.

130 km north of the nearest other locality of

collection of P. ericoides (in the Moore River

National Park). This specimen is a plant which

has a low compact habit typical of P. ericoides

but its inflorescence is more corymbose than

what is typical in P. ericoides. The collection

Newbey 4593 at PERTH, from near Chillicup

Pool, is a plant with a more open spreading habit

than that of the typical form of P. ericoides.

More collections, especially of fruiting material,

and further field investigations are required to

determine if any of these forms are worthy of

formal recognition.

7. Poranthera florosa Halford & R.J.F.Hend.,

sp. nov. maxime arete affinis P. drummondii

Klotzsch cum signis multis communiter

autem foliis spathulatis folia angusta

oblanceolata P. drummondii Klotzsch

plerumque latioribus et insuper ab ea

habitu leviter robustiore et inflorescentia

plus florosa differt. Typus: Western

Australia. Thumb Peak Range, 31 Oct

1965, A.S. George 7134 (holo: PERTH; iso:

PERTH).

Poranthera microphylla var. intermedia

Miill.Arg. in A. DC., Prodr. 15(2): 193

(1866). Type: [Western Australia.] In moist

hollows of rocks at eastern side of

mountain, Melville, (Plantagenet), 5 Oct

1840, [ L .] Preiss 2045 (lecto, here chosen:

Austrobaileya 7 (1): 1-27 (2005)

LD99/018-0887 (specimen on left of

sheet); isolecto: MEL 2062930,2062930).

Monoecious, diffuse annuals to 20 cm high.

Stems much branched from base; branchlets

ascending to decumbent, smooth, glabrous, up

to 2.5 mm across, with leaf scars obscure. Leaves

sessile, alternate, widely spaced along

branchlets; stipules white, narrow-triangular,

0.7-1.1 mm long, entire; laminae spathulate, 9-

25 mm long, 2.5-5 mm wide, flat with margins

slightly recurved, smooth and glabrous

adaxially and abaxially, discolorous; midrib

obscure adaxially, prominent abaxially; base

attenuate; apex obtuse to rounded. Flowers in

short dense terminal umbel-like racemes;

racemes several in a leafy terminal corymb;

peduncles up to 25 mm long; rachis up to 3 mm

long; bracts narrow-obovate, 1.8-3 mm long,

0.4-0.8 mm wide, obtuse to rounded. Male

flowers with pedicels 1.5-3.1 mm long; calyx

lobes 5, white, obovate, 1.1-1.5 mm long, 0.5-

0.7 mm wide, ± flat, rounded at apex; petals 5,

white, erect, oblanceolate to narrow-obovate,

0.6-0.7 mm long, 0.1-0.3 mm wide, acute to

obtuse, with margins entire; glands obdeltate,

0.2-0.3 mm wide; stamens 5; filaments 1.1-1.4

mm long, incurved; anthers c. 0.1 mm long;

rudimentary ovary 3-lobed; lobes clavate, 0.5-

0.6 mm long. Female flowers with pedicels 1.2-

2.2 mm long, extending to 5.5 mm long in fruit;

calyx lobes 5, white, oblong-obovate, 1.1-1.3

mm long, 0.4-0.7 mm wide, slightly concavo-

convex, rounded at apex; petals 5, white, oblong,

0.3-0.4 mm long, 0.1-0.2 mm wide, acute, with

margins entire; ovary depressed globose, 0.6-

1 mm across, deeply 6-lobed, emarginate distally,

papillose; styles each 0.5-1.1 mm long, deeply

2-partite, slender. Capsules depressed globose,

1.5-1.7 mm across, 0.9-1.1 mm long, shallowly

6-lobed, emarginate distally, rugose. Seeds

wedge-shaped, 0.6-0.8 mm long, 0.4-0.5 mm

wide, 0.4-0.5 mm across; testa alveolate, grey-

white. Fig. 2.

Selected specimens (from 10 examined ): Western

Australia. Scott River, Dec 1957, Churchill s.n.

(PERTH); Chespeck Road, Northcliffe, Dec 1983,

Cranfield 4880 (PERTH); Fitzgerald River, Sep 1948,

Gardner 9215 (PERTH); base of Talyuberup Peak,

Stirling Range, Oct 1984, Keighery 7278 (CANB,

PERTH); W of Mt Magog, Stirling Range, Nov 1969,

Mann & George 146 (K, PERTH); Plantagenet, Nov

1901, Pritzel 929 (PERTH); Bremer Bay, in 1900,

Wellstead s.n. (PERTH).

Halford & Henderson, Revision of Poranthera

Distribution and habitat : Poranthera florosa

is confined to south-western Western Australia

from Scott River and Boggy Lake eastwards to

the Stirling Range, Thumb Peak Range and Cape

Arid. It is recorded as growing in heath, eucalypt

mallee or woodland communities in stony sandy

clay or clayey loam soils, or on the edge of

granite outcrops. Map 7.

Phenology : Flowers have been collected from

September to December, fruits from October to

December.

Affinities : Poranthera florosa is most closely

related to P. drummondii with which it shares

many attributes. However, it differs from that

by having spathulate leaves which are generally

broader than the narrow-oblanceolate leaves of

P. drummondii. Poranthera florosa also differs

from that by its slightly more robust growth

habit and more floriferous inflorescence.

Typification : In his protologue of

P. microphylla var. intermedia , Muller cited

Preiss no. 2045 at the Berlin Herbarium. The

type material of P. microphylla var. intermedia

has not been located at B, and it is believed to

have been destroyed during World War II.

Duplicates of Preiss no. 2045 have been located

at LD (one sheet) and MEL (two sheets). While

the MEL sheets are all of one species, the sheet

at LD contains material of two species. The

smaller of the two specimens on that sheet, the

fragment on the right, has flowers, fruit and seed

and is referable to P. microphylla as

circumscribed here. The specimen on the left of

the sheet has flowers and is what we consider

to be the species here named P. florosa. The

Preiss duplicates at MEL match the specimen

on the left of the LD sheet. Both specimens on

the LD sheet would fit the brief description in

the protologue of P. micropylla var. intermedia.

To fix the application of the name then, the

specimen on the left of this sheet is chosen

here as lectotype of P. microphylla var.

intermedia.

Etymology: The specific epithet is derived from

Latin, florosus, profusely flowering, and refers

to the more floriferous inflorescences produced

by this species in comparison with those of

P. drummondii.

8. Poranthera huegelii Klotzsch in J.G.C.

Lehmann, PI. Preiss. 2: 231-232 (1848).

Type: [Western Australia.] In subpeaty

sandy soil between dense shrubs on plain

near city of Albany (Plantagenet), Oct

1840. Preiss. hb. 2047( holo: LD 99/018-

0886; iso: B (ex herb. L. C. Treviranus); G-

DC n.v. (microficheIDC 800-73.2453: II. 1);

MEL1617992,2062909 (exherb. Sonder)).

Illustration: Weber (1986: 763, fig. 407A), as

Poranthera ericoides.

Monoecious, erect or rarely spreading,

herbaceous perennials 30-60 cm high. Stems

undivided or sparingly branched; branchlets

erect or sometimes ascending, smooth,

glabrous, up to 4 mm across, with leaf scars

somewhat prominent. Leaves sessile, alternate,

somewhat widely spaced along branchlets;

stipules white, narrow-triangular, 1.1-3 mm long,

entire or laciniate at the base; laminae linear,

narrow-oblanceolate or narrow-elliptic, 9-26 mm

long, 0.5-2.4 mm wide, flat to rounded with

margins recurved, smooth and glabrous adaxially

and abaxially, discolorous; midrib impressed

proximally adaxially, prominent abaxially; base

shortly cuneate; apex acute to obtuse sometimes

terminated with a minute mucro. Flowers in short

dense terminal umbel-like racemes; racemes in

a lax leafy terminal corymb; peduncles up to 30

mm long; rachis up to 3 mm long; bracts narrow-

obovate, 3-6 mm long, 0.5-1 mm wide, subacute

to obtuse. Male flowers with pedicels 0.9-3 mm

long; calyx lobes 5, white, obovate, 1.6-2.4 mm

long, 0.6-0.9 mm wide, ± concavo-convex,

rounded at apex; petals 5, white, erect, linear to

oblanceolate, 0.6-1.6 mm long, 0.2-0.3 mm wide,

acute to rounded, with margins entire; glands

2-lobed, 0.2-0.4 mm wide; stamens 5; filaments

1.7-2.7 mm long; anthers c. 0.2 mm long;

rudimentary ovary 3-lobed; lobes clavate, 0.7-

1.4 mm long. Female flowers with pedicels 0.3-

1.7 mm long, extending to c. 4.5 mm in fruit;

calyx lobes 5, white, oblong, 1.6-2.5 mm long,

0.5-1.1 mm wide, slightly concavo-convex,

rounded at apex; petals 5, white, linear-oblong,

0.5-1.2 mm long, 0.3-0.4 mm wide, acute to

obtuse, with margins entire; ovary subglobose,

1.1-1.6 mm across, shallowly 6-lobed,

emarginate distally, smooth; styles each 1-1.7

mm long, deeply 2-partite, slender. Capsules

depressed globose, c. 3 mm across, c. 2 mm long,

Austrobaileya 7 (1): 1-27 (2005)

Fig. 2. Poranthera flows a. A. whole plant *0.5. B. inflorescence x8. C. male flower x32. D. branchlet with leaf and

stipules x8. E. fruit from above xl6. F. fruit from side xl6. A-D from George 7134 (PERTH); E, F from Churchill

s.n. (PERTH). Del. W. Smith.

shallowly 3-lobed, emarginate distally, ±

smooth. Seeds wedge-shaped, 0.8-1 mm long,

0.6-0.8 mm wide, 0.6-0.7 mm across; testa

fenestrellate, translucent.

Selected specimens (from 90 examined ): Western

Australia. Jarrahdale, Oct 1908, Andrews s.n.

(PERTH); 3 miles ( c. 5 km) from Mt Barker towards

Albany, Oct 1968, Canning s.n. (CANB [CBG038356]);

5 km S of Margaret River township on Caves Road,

Oct 1983, Corrick 8968 (MEL); E of Cape Naturaliste,

near Castle Bay, Oct 1983, Corrick 9005 (MEL); Mt

Dale Road between main picnic area and corner of

Ashendon Road, Darling Range, Oct 1984, Corrick

9369 (CANB, MEL); Scarp Road, S of Serpentine

Pipehead Dam, Nov 1984, Corrick 9396 (MEL);

Thumb Peak Range, Oct 1965, George 7135 (PERTH);

Gooseberry Hill, Oct 1897, Helms s.n. (PERTH); 7.2

km W of Point Malcolm Junction, Sep 1976, Hnatiuk

761083 (PERTH); Washpool Road, 200 m E of Junction

with Knights Road, 6 km S of intersection Woogenilup

and Knights Road, Oct 1985, Hoyle 1117 (CANB);

Norman’s Beach, North Point, 31 km due NE of Albany,

Oct 1985, Hoyle 1481 (CANB); Baby Barnett Hill, 30

km due NNE of Mt Barker township, Oct 1985, Hoyle

1152 (CANB); 31 km S of Manjimup on South-West

Hwy, Oct 1993, Jobson 2513 (MEL); upper slopes of

Mondurup Peak, Oct 1977, Keighery 1195 (PERTH);

3 miles ( c. 5 km) N of Mt Success, Oct 1962, Newbey

594 (PERTH); 35 miles (c. 56 km) S of Perth on

Albany Hwy, near Mt Cooke, Nov 1972, Paust 1070

Halford & Henderson, Revision of Poranthera

(PERTH); summit of Mt Toolbrunup, Mar 1966, Spratt

19 (PERTH). South Australia. Flinders Chase,

Kangaroo Island, near Breakneck Creek, Nov 1973,

Nelson ANU17292 (CANB); Playford Hwy, near Roo

Lagoon, Kangaroo Island, Nov 1989, Overton 1140

(CANB); Ridge road near Yankalilla turn off, Fleurieu

Peninsula, Dec 1967, Symon 5501 (CANB); Flinders

Chase, Kangaroo Island, Oct 1971, Wace ANU13082

(CANB).

Distribution and habitat: Poranthera huegelii

has a disjunct distribution. It occurs in south¬

western Western Australia from New Norcia

southwards to Augusta and eastwards to

Israelite Bay, and in South Australia on

Kangaroo Island and in the South Mount Lofty

Ranges. It is recorded as growing in Eucalyptus

woodland and open forest, and heath-mallee

communities, often in sandy or gravelly lateritic

soils. Map 8.

Phenology : Flowers have been collected in

March and from August to December, fruits in

January, October and December.

Affinities: Poranthera huegelii seems most

closely related to P. corymbosa but is

distinguishable from that and all other species

of Poranthera by its fenestrellate seed coat.

Notes: Poranthera huegelii shows variation in

habit and the form of the inflorescence which

relates somewhat to the geographical

distribution of the species. Specimens from the

western part of its range (from near NewNorica

southward to Augusta) have an erect, more or

less unbranched habit with widely spaced

leaves along the stem and a diffuse corymbose

inflorescence. Specimens from the eastern part

of its range (from Mt Barker eastward to Israelite

Bay, Western Australia, and from Kangaroo

Island and the South Mount Lofty Ranges in

South Australia) tend to have a shorter and

more branched habit, more crowded leaves and

a more compact corymbose inflorescence. This

eastern form has in the past been incorrectly

identified as P. ericoides (e.g. by Weber 1986).

Specimens from the Stirling Range, Western

Australia, (e.g. Keighery 1195 (PERTH) and

Spratt 19 (PERTH)) tend to have an even shorter

stature than those from the eastern part of the

species’ range. More collections and further

investigations would be required to determine

whether or not either of these forms warrants

formal recognition.

9. Poranthera leiosperma Halford &

R.J.F.Hend., sp. nov. maxime arete afifmis

P. microphyllae Brongn. cum signis multis

communiter autem pagina seminum laevi

ad invicem tuberculata, foliis longioribus

(11-15 mm longis vice 6-11 mm longis),

stipulis longioribus (1.3-1.6 mm longis vice

0.8-1.3 mm longis) et insuper ab ea habitu

leviter robustiore differt. Typus: Western

Australia. 47 miles ( c. 76 km) ENE of

Laverton, 28 Sep 1966, A.S. George 8098

(holo: PERTH).

Monoecious, spreading annuals to 10 cm high.

Stems much branched at base; branchlets

procumbent or erect, smooth, glabrous, 0.7-0.9

mm across, with leaf scars obscure. Leaves

sessile, alternate or opposite, widely spaced

along branchlets; stipules white, narrow-

triangular, 1.3-1.6 mm long, entire; laminae

narrow-obovate, 11-15 mm long, 2.1^1.1 mm

wide, flat with margins slightly recurved, smooth

and glabrous adaxially and abaxially,

discolorous; midrib impressed proximally

adaxially, slightly raised proximally abaxially;

base attenuate; apex obtuse. Flowers in short

dense terminal umbel-like racemes; racemes

solitary; peduncles up to 13 mm long; rachis up

to 4 mm long; bracts narrow-obovate, 4-5 mm

long, 0.7-1 mm wide, apically rounded. Male

flowers with pedicels 0.8-1 mm long; calyx lobes

5, white, obovate, 0.7-0.8 mm long, 0.4-0.5 mm

wide, slightly concavo-convex, rounded at apex;

petals 5, white, erect, oblanceolate, 0.2-0.3 mm

long, c. 0.1 mm wide, acute, with margins entire;

glands clavate, c. 0.1 mm wide; stamens 5;

filaments 0.5-0.6 mm long, incurved; anthers c.

0.1 mm long; rudimentary ovary 3-lobed; lobes

clavate, 0.2-0.3 mm long. Female flowers with

pedicels 0.8-1 mm long, extending up to 7 mm

long in fruit; calyx lobes 5, pink, oblong-elliptic,

0.7-0.8 mm long, 0.4-0.5 mm wide, slightly

concavo-convex, rounded at apex; petals 5,

white, linear-oblanceolate or narrow-obtrullate,

c. 0.2 mm long, 0.1-0.2 mm wide, acute, with

margins entire; ovary depressed globose, 0.8-

1 mm across, shallowly 6-lobed, slightly

emarginate distally, smooth; styles each 0.3-

0.5 mm long, deeply 2-partite, slender. Capsules

depressed globose, 1.6-1.8 mm across, 0.9-1

mm long, shallowly 3-lobed, emarginate distally,

± smooth. Seeds wedge-shaped, 0.7-1.2 mm

long, 0.4-0.9 mm wide, 0.5-0.6 mm across; testa

smooth, white. Fig. 3.

Selected specimens (from 25 examined): Western

Australia. Entrance to Cape Le Grand NP, Nov 1979,

Cranfield s.n. (PERTH); 10 km NW of Southern Cross

on Bullfinch Road, Sep 1980, Cranfield 1643 (PERTH);

E of Laverton, Jul 1916, Gardner 172 (PERTH);

Victoria Desert, camp 53, Sep 1891, Helms s.n.

(MEL2065543, NSW273990, NSW455349,

NSW455351); 5.2 km W of Zanthus, Oct 1986,

Keighery & Alford 761 (PERTH); Parker’s Range, in

1890, Merrall s.n. (MEL2065489); Comet Vale, Sep

1953, Royce 4414 (PERTH); 8 miles (c. 13 km) S of

Queen Victoria Spring, Oct 1956, Royce 5531 (PERTH);

24 km E of Depot Springs Homestead, Aug 1970, Wilson

s.n. (PERTH). Northern Territory. Kings Canyon,

George Gill Range, Oct 1966, Beauglehole ACB20404

(DNA, NSW); Reedy Creek Gorge, 4 miles (c. 6 km) E

of Kings Canyon, George Gill Range, Oct 1966,

Beauglehole ACB20473A (NSW). South Australia.

c. 44.5 km direct N of Hypurna Homestead, Danggali

Conservation Park, Sep 1980, Conn 932 (AD); Great

Australian Desert, 148 km N of Cook, Aug 1980, Symon

12238 (CANB); c. 39.5 km direct N of Hypurna

Homestead, Danggali Conservation Park, Sep 1980,

Toelken 6631 & Christensen (AD). New South Wales.

“Top Hut”, NE of Mildura, Oct 1987, Green s.n.

(NSW215901); 40 miles (64.4 km) N of Balranald,

Oct 1970, Mulham 396 (NSW); c. 1 mile (1.6 km) W

of Timor Rock, near Coonabarabran, Jan 1962, Salasoo

2253 (NSW). Victoria. Hattah Lakes NP, Sep 1969,

Anderson s.n. (MEL694357); Kulkyne NP, Oct 1948,

Beauglehole ACB1035 (MEL).

Distribution and habitat: Poranthera

leiosperma is recorded from across southern

Australia with scattered occurrences near

Southern Cross, Western Australia, eastwards

through southern Northern Territory, South

Australia, north-western Victoria to near

Coonabarabran in western New South Wales.

It is recorded as growing in open shrubland

and mallee communities frequently with Triodia

spp. in the understorey in red sandy soils.

Map 9.

Phenology : Flowers and fruits have been

collected in January and from July to November.

Affinities: Poranthera leiosperma is most

closely related to P. microphylla with which it

shares many attributes. However, it differs from

that by having seeds with a smooth rather than

tuberculate surface, longer leaves (11-15 mm

long as compared with 6-11 mm long) and

longer stipules (1.3-1.6 mm long as compared

with 0.8-1.3 mm long). Poranthera leiosperma

also differs from P. microphylla in its slightly

more robust growth habit.

Etymology: The specific epithet is derived from

Greek, leio -, smooth, and spermus , seed, and

Austrobaileya 7 (1): 1-27 (2005)

refers to the smooth seed coat in this species

which contrasts with that in the majority of

Poranthera species which is variously

ornamented.

10. Poranthera microphylla Brongn., Sci.

Nat. (Paris) 29: 385 (1833); Poranthera

microphylla Brongn. var. microphylla ,

Miill.Arg. in A DC., Prodr. 15(2): 193

(1866); Poranthera microphylla var.

genuinaMull.Arg. in A.DC., Prodr. 15(2):

193 (1866), nom. inval. Type: [Australia.]

Port Jackson, s.d., [L.A.] Lesson s.n. (lecto,

here chosen: PI52754 (exherb. Brongn.);

isolecto: P152755).

Poranthera microphylla var. diffusa

Miill.Arg. in A DC., Prodr. 15(2): 193

(1866). Type: [Australia.] New Holland, s.d.

[without collector] (holo: G-DC n.v.

(microfiche IDC 800-73.2453: II. 7)).

Poranthera microphylla var. glauca

Miill.Arg. in A DC., Prodr. 15(2): 193

(1866). Type: [Australia. New South

Wales.] New Holland, in 1816, [without

collector] (syn: G-DC n.v. (microfiche 800-

73.2453. II. 6); syn: B n.v. (? destroyed)).

Poranthera microphylla var. diffusa Ewart,

FI. Victoria 721 (1930 ),nom. illeg. Type:

unknown; no material cited.

Illustrations: Griining (1913:17, fig. 4F-G), as

Poranthera microphylla var. intermedia ;

Cunningham etal. (1982:464); James &.

Harden (1990: 394); Wheeler (1992: 627,

fig. 192A); Kenneally et al. (1996: 106);

Kirkpatrick (1997: fig. 49f); Corrick &

Fuhrer(2000: 82).

Monoecious, diffuse or compact annuals or

rarely perennials, to 30 cm high. Stems much

branched at base then sparingly branched

distally; branchlets ascending to erect, ±

smooth, glabrous, 1.5-2 mm across, with leaf

scars obscure. Leaves sessile or shortly

petiolate, alternate, subopposite or opposite,

widely spaced along branchlets; stipules white,

narrow-ovate, 0.8-1.3 mm long, entire; petioles

where present up to 1.5 mm long; laminae

narrow-oblanceolate to obovate, spathulate or

ovate, 6-11 mm long, 1-4 mm wide, flat with

margins slightly recurved, smooth and glabrous

Halford & Henderson, Revision of Poranthera

Fig. 3. Poranthera leiosperma. A. whole plant *1.5. B. inflorescence *6. C. male flower x32. D. branchlet with

leaves and stipules x6. E. fruit from above xl6. F. fruit from side xl6. A-F from George 8098 (PERTH). Del. W.

Smith.

adaxially and abaxially, discolorous; midrib

obscure adaxially, prominent abaxially; base

attenuate or cuneate; apex obtuse to rounded,

usually shortly apiculate. Flowers in short dense

terminal racemes; racemes solitary; peduncles

up to 20 mm long; rachis up to 5 mm long; bracts

narrow-obovate, 3-4 mm long, 0.5-0.7 mm wide,

obtuse. Male flowers with pedicels 0.5-1.5 mm

long; calyx lobes 5, white or pink, obovate or

oblong, 0.4-1.5 mm long, 0.2-0.8 mm wide, ±

concavo-convex, rounded or obtuse at apex;

petals 5, mostly white, pink or sometimes pale

mauve, erect, oblanceolate or linear, 0.2-0.6 mm

long, 0.1-0.3 mm wide, obtuse, with margins

erose; glands clavate or obdeltate, 0.1-0.4 mm

wide; stamens 5; filaments 0.3-0.9 mm long;

anthers up to 0.1 mm long; rudimentary ovary a

hemispherical dome or 3-lobed; lobes clavate,

0.1-0.4 mm long. Female flowers with pedicels

0.2-1 mm long, extending to 9 mm long in fruit;

calyx lobes 5, white or pale pink with greenish

band along midline, oblong-elliptic, 0.5-1.5 mm

long, 0.2-0.8 mm wide, flat or slightly concavo-

convex distally, obtuse at apex; petals 5, white,

linear-oblanceolate or narrow-obtrullate, 0.1-0.5

mm long, 0.1-0.3 mm wide, acute to obtuse, with

margins minutely erose; ovary depressed

globose, 0.4-1.1 mm across, shallowly to deeply

6-lobed, flat to slightly emarginate distally,

smooth or papillose; styles each 0.2-0.7 mm

long, deeply 2-partite, slender or rarely stout.

Capsules depressed globose, 1.4-2.5 mm

across, 0.6-1.4 mm long, prominently 6-lobed,

slightly emarginate distally, slightly rugose.

Seeds wedge-shaped, 0.4-1.1 mm long, 0.4-0.9

mm wide, 0.3-0.9 mm across; testa tuberculate,

white.

Selected specimens (from 600 examined ): Western

Australia. Hill River crossing on Brand Hwy, c. 6 km

N of Badgingarra, Sep 1988, Henderson H3135 (BRI);

Peak Eleanora, Peak Charles NP, c. 45 km W of Salmon

Gums, Nov 1979, Newbey 6406 (PERTH); Cape Arid

NP, E of Esperance, Dec 1971, Royce 10149 (PERTH).

Northern Territory, c. 15 km N of Borroloola on

Bing Bong Homestead road, Jun 1976, Craven 4243

(CANB); 105 km from Borroloola on Wollogorang

road. May 1987, Craven 8529 et al. (CANB);

Parakeelya swamp, 50 km SW of Nathan River

Homestead, Aug 1985, Latz 10203 (DNA). South

Australia. Hambidge Flora and Fauna Reserve, Oct

1966, Alcock 1138 (BRI, CANB); c. 5 km E along

Casterton Road from the Nangwarry to Penola road,

Oct 1989, Henderson & Turpin H3312 (AD, BRI);

near Comaum, c. 20 km E of Penola, Sep 1962, Hunt

1170 (AD). Queensland. Cook District: Tinaroo Creek

forestry road, 8.2 km past Henry Hannan Drive towards

Mt Haig, Apr 1989, Henderson & Clarkson H3218

(BRI). Darling Downs District: Girraween NP, 4.5 km

ESE of Park Headquarters, Bald Rock Creek, Jan 1993,

Halford Q1593 et al (BRI, MEL). Moreton District:

Old Logan Road, Camira, Apr 1992, Sharpe 5310 &

Bird (BRI, MEL). New South Wales, c. 22 km NE of

Emmaville on Amaroo road, Oct 1993, Coveny 16652

& Whalen (BRI, NSW); Tantawangalo State Forest, 12

km S of Tantawangalo, Apr 1993, Crawford 2254

(CANB, MEL, NSW); on Wollar/Bylong road, c. 30

km SW of Merriwa, Sep 1990, Henderson & Turpin

H3409 (BRI). Australian Capital Territory. Mt

Loch, Feb 1983, Beardsell s.n. (MEL 1562983); Mt

Franklin, Brindabella Range, Mar 1957, Pullen 73

(CANB). Victoria. Mt Samaria Road, c. 65 km NW of

Bridge Creek township, Nov 1996, Clarke 2753 (MEL);

c. 6 km SE of Taggerty, Cathedral Range, Neds Gully

camping area, Nov 1996, Clarke 2726 (MEL);

Gippsland Plains, Briagolong Forest Reserve, 500 m S

of Cemetery Road and 300 m E of Briagolong-Stratford

Road, Oct 1991, Lunt 91/84 (MEL). Tasmania.

Flinders Island, Strzelecki NP, 1.4 km SW of Strzelecki

Peaks, Dec 1988, Crawford 1136 (BRI, CANB);

Wombat Moor, Jan 1948, Curtis s.n. (HO 6359); N of

Austrobaileya 7 (1): 1-27 (2005)

Bessells Road, Sale Rivulet, Dec 1983, Patacsek s.n.

(H072814); Mt Field National Park, Mt Field West,

Feb 1969, Telford EMC2227 (CANB).

Distribution and habitat : Poranthera

microphylla is widespread and is recorded in

all Australian states though it is not found in

the more arid regions of Australia. It also occurs

in New Zealand. It is recorded as growing in a

wide variety of habitats from coastal dunes to

alpine mountain slopes in grassland, heathland,

woodland and open forest communities on a

range of substrates. Map 10.

Phenology : Flowers and fruits have been

collected throughout the year.

Typification: In his protologue of

P. microphylla, Brongniart cited two collections,

namely “in Nova-Hollandia In Montibus

Caeruleis near Port-Jackson (Lesson)” and “ad

fretum Entrecasteaux (Vog. de Baudin, in herb.

Mus. Par.)”. Two sheets of the Lesson collection

and one sheet of the Baudin collection have

been located amongst material on loan to BRI

from P, and are numbered 152754,152755 and

152753 respectively. The Lesson specimen

152754 is selected here as lectotype of

Poranthera microphylla as it is part of the

original material, agrees with the protologue

description and is more than likely the material

illustrated in the protologue.

Notes: Poranthera microphylla displays a large

degree of morphological variation. However,

the variants are not considered sufficiently

distinct enough to warrant formal recognition

here. In general, those plants from populations

in the Kimberley in Western Australia, central

Northern Territory and the Cook District in

northern Queensland have more or less erect

stems, linear leaves and generally comparatively

smaller flowers, fruits and seeds than those of

other forms; plants from south-western Western

Australia and South Australia have a

procumbent to erect habit and leaves narrow-

obovate with a cuneate base; plants in eastern

Australia, from north-eastern Queensland to

Tasmania, generally have a sprawling to weakly

ascending habit and spathulate leaves with an

attenuate base.

There is also a high-altitude form in the

Australian Alps and in the higher altitudes in

Halford & Henderson, Revision of Poranthera

Tasmania which resembles P. oreophila and

P. petalifera in having opposite leaves with

more or less strongly revolute leaf margins and

stout style branches. Some specimens of

P. microphylla seem to possess a perennial

rootstock ( e.g. Pullen 73 (CANB), Beardsell

s.n. (MEL 1562983) and Telford EMC2227

(CANB)). This form of P. microphylla differs

from the other mentioned species by having

monoecious rather than dioecious plants. It also

differs from P oreophila in having tuberculate

rather than smooth seeds and from P. petalifera

in having its flowers in terminal umbel-like

racemes rather than being solitary in leaf axils.

11. Poranthera obovata Halford & R. J.F.Hend.,

sp. nov. ut videtur maxime arete affinis

P. ericifoliae Rudge sed foliis angustis

obovatis vice linearibus usque angustis

oblanceolatis, marginibus foliorum leviter

recurvatis vice valide recurvatis usque

revolutis et stipulis integris vice laciniatis

distinguenda. Typus: Queensland.

Moreton District: Helidon Hills, c. 10 km

N of Helidon township, 7 Sep 1989, R.J.F.

HendersonH3231 (holo: BRI; iso: MEL,

NSW, distribuendi).

Poranthera sp. (Beerwah D.Hassall DH7431),

PI. Forster & D A. Halford (2002, p.73).

Monoecious, spreading to erect, herbaceous

perennials to 20 cm high. Stems unbranched or

sparingly branched; branchlets ascending to

erect or rarely procumbent, smooth or papillose,

glabrous, 1.5-2 mm across, with leaf scars

prominent. Leaves sessile, alternate, crowded

along branchlets; stipules white, narrow-ovate

or narrow-triangular, 1.5-2 mm long, entire;

laminae narrow-obovate, 10-15 mm long, 1.5-3

mm wide, flat with margins slightly recurved,

smooth and glabrous adaxially and abaxially,

discolorous; midrib impressed proximally

adaxially, prominent abaxially; base attenuate;

apex obtuse. Flowers in short dense terminal

racemes; racemes solitary or rarely several in a

leafy terminal corymb; peduncles 10-25 mm

long; rachis 2-30 mm long; bracts narrow-

obovate, 2-7 mm long, 0.5-1.3 mm wide, obtuse

to rounded. Male flowers with pedicels 1.5-2.5

mm long; calyx lobes 5, white to pink or rarely

with pale blue hue, obovate, 1.6-1.7 mm long,

0.5-0.7 mm wide, slightly concavo-convex,

rounded at apex; petals 5, white or pale pink,

erect, recurved distally, terete proximally, c. 0.5

mm long, toe. 0.1 mm across, acute, with margins

entire; glands 2-lobed, 0.2-0.3 mm wide;

stamens 5; filaments 1.4-1.5 mm long, slightly

incurved; anthers 0.2-0.3 mm long; rudimentary

ovary 3-lobed; lobes clavate, 0.5-0.6 mm long.

Female flowers with pedicels 1.5-2 mm long,

extending to c. 6 mm in fruit; calyx lobes 5, white

to pink, oblong, 1.3-2.8 mm long, 0.6-0.7 mm

wide, slightly concavo-convex, rounded at apex;

petals 5 or sometimes absent, white or pink,

subulate, 0.4-0.6 mm long, 0.1-0.2 mm wide,

acute, with margins entire; ovary depressed

globose, 1-1.8 mm across, deeply 6-lobed,

emarginate distally, papillose; styles each c. 1

mm long, deeply 2-partite, slender. Capsules

depressed globose, 2.8-3 mm across, 1.7-2 mm

long; prominently 6-lobed distally, emarginate

distally, papillose. Seeds wedge-shaped, 1.1-

1.2 mm long, c. 2 mm wide, c. 2 mm across; testa

reticulate-foveate with fovea white and ridges

pale brown and translucent. Fig. 4.

Selected specimens (from 30 examined ):

Queensland. Port Curtis District: Kroombit Tops,

State Forest 316, 48 km E of Biloela, Sep 1988, Gibson

TOI430 (BRI). Leichhardt District: Blackdown

Tableland, c. 35 km SE of Blackwater, Sep 1971,

Henderson H967 et at. (BRI, MEL); Mimosa Creek,

Blackdown Tableland, Sep 1974, Williams 74054 (BRI).

Wide Bay District: Oakey Creek road, Blackall Range,

Oct 1990, Sharpe 4977 & Bean (BRI). Moreton

District: State Forest 893 Mt Mee, Aug 1995, Forster

PIF17391 & Leiper (BRI, MEL, NSW); near Forestry

Station, Beerwah, Jun 1974, Hassall DH7431 (BRI);

Mapleton Forestry Scenic Drive, c. 5 km N of Mapleton,

Nov 1987, Sharpe 4751 & Guymer (BRI); Lockyer

State Forest (SF616), Nov 1994, Thompson & Bean

MOR453 (BRI); at foot of Mount Beerburrum, Sep

1959, Thurlow s.n. (BRI [AQ205075]). New South

Wales. 8 km NNW of Glenreagh, Oct 1990, Bean

2453 (BRI); 7 km NNW of Glenreagh, Oct 1990, Bean

2462 (BRI, CANB, K, MEL, NSW); Rocky Creek, 28

km from Grafton towards Coaldale, Feb 1995, Bean

8357 (BRI, NSW).

Distribution and habitat : Poranthera obovata

occurs in eastern Australia from the Blackdown

Tableland near Blackwater in central

Queensland southwards to Glenreagh in

northern New South Wales. It is recorded as

growing in heath, open eucalypt woodland or

forest communities in either shallow sandy soils

usually overlying sandstone or gravelly sandy

clays. Map 11.

Austrobaileya 7 (1): 1-27 (2005)

Fig. 4. Porantliera obovata. A. branchlet with flowers x2. B. male flower xi6. C. female flower xi6. D. branchlet

with leaf and stipules x8. E. fruit from above xl2. F. fruit from side xl2. A, D from Sharpe 4977 & Bean (BRI);

B, C, E, F from Henderson H3588 (BRI). Del. W. Smith.

Phenology : Flowers have been collected in

February, April, June and from August to

December, fruits in February, April, June,

September, October and December.

Affinities : Poranthera obovata seems most

closely related to P. ericifolia but is

distinguished from that by its narrow-obovate

rather than linear to narrow-oblanceolate leaves,

slightly recurved rather than strongly recurved

to revolute leaf margins and entire rather than

laciniate stipules.

Etymology : The specific epithet is derived from

Latin, obovatus , obovate, in reference to the

shape of the species’ leaves.

12. Poranthera oreophila Halford &

R.J.F.Hend., sp. nov. olim P. microphylla

Brongn. confusa sed plantis dioeciis,

seminibus testa laevi et pallide brunnea

distinguenda. P oreophila ut videtur

maxime arete affinis P. petaliferae

(Orchard & J.B.Davies) Halford &

R.J.F.Hend. autem floribus in racemis

terminalibus dispositis et floribus

masculis plerumque minoribus

distinguenda. Typus: New South Wales.

Kosciusko N.R, Gurrangorambla Range,

Mount Morgan, 12 Jan 1994, A.E.

Jenkinson 3 &D.K. Judge (holo: CANB;

iso: BRI, MEL, NSW).

Halford & Henderson, Revision of Poranthera

Dioecious, compact herbaceous perennials to

30 cm high. Stems sparingly branched;

branchlets annual, prostrate or weakly

ascending, ± smooth, glabrous; 0.7-1.1 mm

across, with leaf scars obscure. Leaves sessile

or shortly petiolate, opposite, widely spaced

along branchlets; petioles 0.2-0.5 mm long;

stipules white, narrow-triangular, 0.5-1 mm long,

entire; laminae narrow-ovate, narrow-obovate

or narrow-elliptic, 4.3-6.5 mm long, 1.8-3.8 mm

wide, flat or rounded with margins recurved to

revolute, smooth and glabrous adaxially and

abaxially, discolorous; midrib impressed

proximally adaxially, slightly raised proximally

abaxially; base attenuate; apex acute to obtuse.

Flowers in short dense terminal racemes;

racemes solitary; peduncles 5-10 mm long;

rachis 2-4 mm long; bracts narrow-obovate, 1.1-

2 mm long, 0.5-1.1 mm wide, obtuse. Male

flowers with pedicels 0.5-0.7 mm long; calyx

lobes 5, white, obovate or oblong, 0.9-1.1 mm

long, 0.5-0.9 mm wide, ± flat; rounded at apex;

petals 5, white, erect, recurved distally,

oblanceolate, 0.4—0.5 mm long, 0.1-0.2 mm wide,

acute, with margins ± entire; glands 2-lobed or

subglobose, 0.2-0.4 mm wide; stamens 5;

filaments 0.5-0.9 mm long, ± straight; anthers c.

0.1 mm long; rudimentary ovary 3-lobed, lobes

clavate, 0.2-0.3 mm long. Female flowers with

pedicels up to 1.1 mm long, extending to 7 mm

long in fruit; calyx lobes 5, white sometimes with

mauve tinge, oblong to oblong-obovate, 1.1-

1.2 mm long, 0.8-0.9 mm wide, flat, rounded at

apex; petals 5, white, linear-oblanceolate, 0.2-

0.6 mm long, 0.1-0.2 mm wide, acute, with

margins entire; ovary depressed globose, 1-

1.3 mm across, shallowly 6-lobed, emarginate

distally, smooth or papillose; styles each 0.5-

0.6 mm long, deeply 2-partite, stout. Capsules

depressed globose, 1.7-2.2 mm across, 1.2-1.4

mm long, shallowly 3-lobed, emarginate distally,

± smooth. Seeds wedge-shaped, 0.9-1 mm long,

0.6-0.7 mm wide, 0.8-0.9 mm across; testa

smooth, pale brown or white. Fig. 5.

Selected specimens (from 40 examined ): New South

Wales. Kiandra, 0.5 miles (c. 0.8 km) SW of road

junction, Feb 1969, Briggs 2523 (NSW); Kosciusko

N.P., 2 miles (c. 3.2 km) from Grey Mare Track

junction with Happy Jacks Road, toward Happy Jacks

Pondage, Jan 1966, Carroll 150A (CANB); Jindabyne,

Mar 1907, Hallman s.n. (NSW455328); Grey Mare

Range, c. 2.4 km SW of Grey Mare Hut, Feb 1968,

Rodd 712 (NSW); Rule’s Point N of Kiandra, Jan 1969,

Salasoo 3503 (NSW); Kosciusko NP, Little Peppercorn

Plain, Feb 1981, Taylor 1309 et al. (CANB); Snowy

River between Guthega Dam and Spencers Creek, Jan

1971, Thompson 1008 (NSW); Cooleman Plain, Dec

1962, Walker ANU952 (CANB). Australian Capital

Territory, between Mt Gingera and Blackfellows Gap,

May 1959, Burbidge 6398 & Gray (CANB); Mt

Gingera, Feb 1953, Moore 2323 (CANB); east slope of

Mt Gingera, Brindabella Range, Nov 1966, Schodde

5159 (CANB, K, NSW). Victoria. Bogong High Plains,

Wilkinson Memorial Lodge area, Jan 1966,

Beauglehole ACB15421 (MEL, NSW); near Cope Hut,

Bogong High Plains, Feb 1972, Craven 2184 (CANB);

Bogong High Plains, Nelse Track, Jan 1982, Forbes

774 & van Rees (MEL); Bogong NP, footslopes of Mt

Nelse beside track, Feb 1985, Norris 302 (NSW); Digges’

Holes, Nunniong Plateau, Jan 1949, Wakefield 2665

(MEL); Mt Cobberas, Feb 1974, Willis s.n.

(MEL2062983); Cobberas Mountains, between No. 1

summit and Asses Ears, Feb 1975, Willis s.n.

(MEL2062982).

Distribution and habitat: Poranthera

oreophila occurs in alpine areas of south¬

eastern Australia from the Brindabella Range in

the Australian Capital Territory, through

Kosciusko National Park, New South Wales, to

Nunnjong Tableland and Bogong High Plains,

Victoria. It is recorded as growing mostly in

stony clay loam soils in alpine tussock grassland

and herbfield communities, at altitudes higher

than 1300 metres. Map 12.

Phenology : Flowers and fruits have been

collected from November to May.

Affinities: Poranthera oreophila has been

identified as P. microphylla in the past but can

be distinguished from that species by being

dioecious and having smooth light brown

seeds. Poranthera oreophila seems most

closely related to P. petalifera but can be

distinguished from that by having its flowers in

tenninal racemes and its generally smaller male

flowers. Poranthera oreophila is similar to the

New Zealand endemic P. alpina but can be

distinguished from that by having its flowers in

short dense terminal racemes, both the male and

female flowers possessing petals and male

flowers with a vestigial ovary.

Etymology: The specific epithet is derived from

Greek, oreo-, mountain, and philus , loving, in

reference to the species’ habitat.

Austrobaileya 7 (1): 1-27 (2005)

Fig. 5. Poranthera oreophila. A. branchlet with flowers and fruit *0.8. B. inflorescence of male flowers x8. C. male

flower xi6. D. petal from male flower x32. E. branchlet with leaves and stipules x6. F. inflorescence of female

flowers x8. G. fruit from above xl2. H. fruit from side xf2. A from Moore 2323 (CANB); B-E from Forbes 774

& van Rees (MEL); F-H from Briggs 2523 (NSW). Del. W. Smith.

13. Poranthera petalifera (Orchard &

J.B.Davies) Halford & R. J.EHend., comb,

nov., Oreoporantherapetalifera Orchard

& J.B.Davies, Pap. & Proc. Roy. Soc.

Tasmania 119: 62/63 (1985). Type:

Tasmania. NE ridge of Mt Anne, 31 Dec

1984, J.B. Davies 31 (holo: HO n.v.\ iso:

CHRw.v.,MEL).

Illustrations : Orchard & Davies (1985: fig. 1),

as Oreoporanthera petalifera ; J.

Kirkpatrick (1997: fig. 45c), as

Oreoporanthera petalifera.

Dioecious, compacted herbaceous perennials

to 30 cm high and to 20 cm across. Stems much

branched; branchlets decumbent, smooth,

glabrous, 0.6-0.9 mm across, with leaf scars

obscure. Leaves shortly petiolate, opposite,

crowded or widely spaced along branchlets;

petioles up to 1 mm long; stipules white, narrow-

triangular, 0.8-1.2 mm long, entire; laminae

lanceolate or narrow-obovate, 3.5-7.1 mm long,

0.8-1.7 mm wide, rounded with margins strongly

revolute to midrib, smooth and glabrous

adaxially and abaxially, discolorous; midrib

obscure adaxially, slightly raised proximally

Halford & Henderson, Revision of Poranthera

abaxially; base attenuate; apex acute. Flowers

solitary in upper leaf axils. Male flowers with

pedicels 0.9-2.6 mm long; calyx lobes 5, white,

oblong or obovate, 1.1-2 mm long, 0.6-0.8 mm

wide, ± flat, rounded at apex; petals 5, white,

erect, incurved distally, oblanceolate, 0.6-0.7 mm

long, 0.1-0.2 mm wide, acute, with margins

entire; glands subglobose, 0.3-0.4 mm wide;

stamens 5; filaments 0.8-1.2 mm long, incurved;

anthers 0.3-0.4 mm long; rudimentary ovary 3-

lobed; lobes clavate, 0.2-0.3 mm long. Female

flowers with pedicels up to 1 mm long, extending

to c. 4 mm in fruit; calyx lobes 5, white, oblong

to oblong-obovate, 0.9-2.2 mm long, 0.6-0.9 mm

wide, slightly concavo-convex, rounded at apex;

petals 5, white, linear-oblanceolate or narrow-

ovate, 0.2-0.6 mm long, 0.1-0.3 mm wide, acute

or obtuse, with margins ± entire; ovary globose,

c. 0.6 mm across, shallowly 6-lobed, flat distally,

smooth; styles each c. 0.2 mm long, 2-partite,

stout. Capsules depressed globose, c. 2 mm

across, c. 1.2 mm long, shallowly 3-lobed,

emarginate distally, smooth. Seeds subglobose,

0.8-0.9 mm long, 0.7-0.8 mm across; testa faintly

tuberculate, white or orange brown.

Additional specimens examined : Tasmania. NE ridge

of Mount Anne, Jun 1984, Buchanan & Davies 3725

(HO); loc. cit., Dec 1984, Buchanan 5101, 5102, 5103,

5116, 5117, 5119 (HO); Mt Anne, Feb 1989, Croft

10176 & Richardson (CANB).

Distribution and habitat: Poranthera

petalifera is confined to Mount Anne in south¬

western Tasmania, occuring at altitudes from

800 to 1000 m above sea level. It is recorded as

growing in crevices of exposed dolomite rocks

in dwarf subalpine rainforest. Map 13.

Phenology : Flowers have been collected in

December and February, fruits in June.

Affinities: Poranthera petalifera seems most

closely related to P. oreophila and the New

Zealand endemic P. alpina. It can be

distinguished from the former by having its

flowers borne singly in the axils of the upper

leaves and generally by the larger male flowers.

For comments on the distinguishing features

of P. petalifera from P. alpina , see Orchard &

Davies (1985).

14. Poranthera triandra J.M.Black, Trans. &

Proc. Roy. Soc. South Australia 40: 66-

67, t.7 (1916). Type: [South Australia.]

Yeelanna, Nov 1915, T.G.B. Osborn s.n.

(holo: AD977512A; iso: AD9775128B, K).

Illustration: Weber (1986: 764, fig. 408).

Monoecious, compact annuals to 3 cm high.

Stems much branched; branchlets procumbent

or erect, smooth, glabrous, up to 0.5 mm across,

with leaf scars obscure. Leaves ± sessile,

alternate or opposite, widely spaced along

branchlets; stipules white sometimes with red-

brown striations, narrow-triangular, 0.5-1 mm

long, entire; laminae narrow-obovate to obovate

or narrow-elliptic to elliptic, 2-5.5 mm long, 0.9-

2.2 mm wide, flat, smooth and glabrous adaxially

and abaxially, concolorous; midrib obscure

adaxially, slightly raised proximally abaxially;

base attenuate; apex obtuse to rounded. Flowers

in short dense terminal umbel-like racemes;

racemes solitary or several in a leafy terminal

corymb; peduncles 5-10 mm long; rachis 1-2

mm long; bracts narrow-obovate, 1-1.5 mm long,

0.4-0.5 mm wide, obtuse to rounded. Male

flowers with pedicels 0.3-0.6 mm long; calyx

lobes 3(rarely 4 or 5), pink sometimes with

reddish striations, oblong or broadly ovate, 0.4-

0.5 mm long, 0.1-0.5 mm wide, concavo-convex,

rounded at apex; petals 3, white, erect, ovate,

0.1-0.5 mm long, up to 0.2 mm wide, obtuse to

rounded, with margins entire; glands obscure;

stamens 3; filaments 0.2-0.3 mm long, straight;

anthers up to 0.1 mm long; rudimentary ovary

minute, hemispherical. Female flowers with

pedicels c. 0.3 mm long, extending to c. 3 mm

long in fruit; calyx lobes 3(rarely 4 or 5), white

with red-brown striations, oblong-elliptic or

ovate, 0.5-0.6 mm long, 0.2-0.5 mm wide, slightly

concavo-convex, obtuse or sometimes acute at

apex; petals absent or 3, when present white,

subulate, c. 0.3 mm long, c. 0.05 mm wide, acute,

with margins entire; ovary depressed globose,

0.5-0.9 mm across, deeply 6-lobed, emarginate

distally, smooth or slightly papillose; styles

each 0.1-0.2(-0.4) mm long, shortly divided

distally or rarely 2-partite, stout or rarely slender.

Capsules depressed globose, 0.9-1.3 mm

across, 0.5-0.8 mm long, prominently 6-lobed,

emarginate distally, smooth or papillose. Seeds

wedge-shaped, 0.4-0.6 mm long, 0.3-0.4 mm

wide, 0.3-0.4 mm across; testa granulate, white

or pale brown with white protuberances.

Selected specimens (from 20 examined ): Western

Australia. Eucla, in 1889, Batt s.n. (MEL2065675,

2065674); 4.7 km S of Sunrise Hill road, east boundary

of reserve 33113, Oct 1984, Burgman 4485 (PERTH);

21.7 km due SSE of Peak Eleanora, Sep 1984, Burgman

3599B (PERTH); Parker Range, in 1890, Merrall s.n.

(MEL2065535); Jimberlana Hill, 9 km NE of

Norseman, Aug 1980, Newbey 7194 (PERTH); Middle

Island, Recherche Archipelago, Nov 1973, Weston 8760

& Trudgen (PERTH); Middle Island, Recherche

Archipelago, Nov 1950, Willis s.n. (MEL2063002).

Northern Territory. Reedy Creek Gorge, 4 miles (c.

6.4 km) E of Kings Canyon, George Gill Range, Oct

1966, Beauglehole ACB20473B (NSW); near Reedy

Rockhole, George Gill Range, Jul 1981, Thomson 86

(DNA). South Australia. Well E of Meningie, Oct

1965, Beauglehole ACB14474 & Williams (AD, MEL,

NSW); Monarto South, c. 15 km W of Murray Bridge,

Nov 1945, Cleland s.n. (AD96005033); c. 60 km SE

of Adelaide, Oct 1958, Eichler 15061 (AD); S of Port

Rickaby, Yorke Peninsula, Oct 1980, Heyligers 80162

(CANB); Lucindale, c. 35 km W of Naracoorte, Nov

1934, Ising s.n. (AD97409341); near Murray Bridge,

Sep 1973, Smith 2414 (AD); c. 15 km W of Murray

Bridge on Adelaide-Melbourne railway, Sep 1974,

Spooner 3573 (AD); Kangaroo Island, 1886, Tepper

s.n. (MEL2065666). Victoria. Grampians, in 1892,

Walter s.n. (MEL694310).

Distribution and habitat: P or anthera triandra

occurs in scattered populations from near

Esperance, Lake King and Eucla in Western

Australia, Kangaroo Island, Yorke Peninsula

and Murray Bridge in South Australia, George

Gill Range in the Northern Territory, and the

Grampians, Victoria. It is recorded as growing

in open eucalypt woodland, low shrubland or

open mallee communities in sandy or loamy

sand soils. Map 14.

Phenology: Flowers and fruits have been

collected from July to November

Notes: The Central Australia collections

Thomson 86 (DNA) and Beauglehole

ACB20473B (NSW) are from populations

somewhat isolated from the others of this

species. These specimens differ from others of

the species in having a colliculate rather than

granulate seed surface and the calyx lobes of

its female flowers are narrower than those

observed in other collections. They also lack

the reddish-brown marks that are prominent in

other dried specimens of P. triandra. These

specimens are tentatively included here within

our concept of P. triandra but further collections

and field studies are warranted to establish their

correct identity.

Excluded Names

Poranthera microphylla var. procera Griming

mAEnglQyPflanzenr. Heft 58:18(1913). Type

citation: ‘Western Australia. Swan River

(Drummond without number in Herb.

Berol.)’(holo: B (? destroyed)).

Austrobaileya 7 (1): 1-27 (2005)

Type material appears non existent rendering

the correct placement of this name impossible

for us at this stage. From the protologue

description it is most probably a synonym of

Poranthera microphylla.

Poranthera glauca Klotzsch in J.G.C. Lehmann,

PI. Preiss. 2: 231 (1848); Poranthera ericoides

var. glauca (Klotzsch) Domin, Vestn. Krai.

Ceske Spolecn. Nauk. Tr. Mat.-Prir. (Mem. Soc.

Roy Sci. Boheme, Prague) 1921/2(2): 58 (1923).

Type citation: [Western Australia.] ‘Inarenosis

ad fluvium Cygnorum J. Drummond a. 1839 Herb.

Preiss. sine No. 5 (holo: ?).

We have been unable to locate any type material

rendering the correct placement of this name

impossible for us at this stage. From the

protologue description it is most probably a

synonym of Poranthera huegelii.

Acknowledgements

We would like to thank Dr Gordon Guymer for

making space and facilities available at BRI for

the first author; the directors and curators of

AD, B, CANB, DNA, HO, K, LD, MEL, NE, NSW,

PERTH for loan of their holdings of Poranthera

material for study at BRI; Alex Chapman and

Bob Chinnock for searching for types on our

behalf at E and BM while acting as Australian

Botanical Liaison Officer at K, Les Pedley and

Peter Bostock (BRI) for assistance in translation

of diagnoses into Latin, Will Smith (BRI) for the

excellent illustrations and Peter Bostock for the

maps. Associated fieldwork from 1988 to 1992

by the second author and salary support for

the first author from 1999 and 2000 was funded

by grants from the Australian Biological

Resources Study (ABRS) which are gratefully

acknowledged.

References

Baillon, H.E. (1858). Etude generate du groupe des

Euphorbiacees. Victor Masson: Paris.

Baillon, H.E. (1866). Species Euphorbiacearum

Euphorbiacees Australiensis. Adansonia 6:

282-345.

Bentham, G. (1873). Euphorbiaceae. In Flora

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Black, J.M. (1916). Additions to the Flora of South

Australia no. 9. Transactions and Proceedings

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56-77.

Halford & Henderson, Revision of Poranthera

Brongniart, A.T. (1833). Note sur quelques

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Annales de sciences naturelles. (Paris) 29:

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Brummitt, R.K. & Powell, C.E. (1992). Authors of Plant

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Flora. W A G. Skinner, Government Printer:

Wellington.

Corrick, M.G. & Fuhrer, B.A. (2000). Wildflowers of

Victoria and adjoining areas. Victoria:

Bloomings Books: Hawthorn.

Cunningham, G.M., Mulham, W.E., Milthorpe, P.L. &

Leigh, J.H. (1982). Plants of Western New South

Wales. New South Wales Government Printing

Office: Sydney.

Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1993).

DELTA user’s guide, a general system for

processing taxonomic descriptions , 4 th edn.

CSIRO: East Melbourne.

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of the Darwin Region. Conservation

Commission of the Northern Territory: Darwin.

Forster, PI. & Halford, D.A. (2002). Poranthera. In

R.J.F. Henderson (ed.). Names and

Distribution of Queensland Plants, Algae and

Lichens, p.73. Queensland Herbarium,

Queensland Government Environmental

Protection Agency: Brisbane.

GrUning, G. (1913). Poranthera. IV 147 Euphorbiaceae

- P or anther oideae et Ricinocarpoideae. In

A. Engler (ed.). Das Pflanzenreich, Regni

vegetabilis conspectus H.58: 13-21. Verlag

von Wilhelm Engelmann: Leipzig.

Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (1990).

Index Herbariorum Part 1, 7 th edn. Regnum

Vegetabile 106.

Hooker, J.D. (1881). Poranthera alpina. Hooker’s

leones PIantarum 14: 49-50.

Hutchinson, J. (1969). Tribalism in the family

Euphorbiaceae. American Journal of Botany

56: 738-758.

James, J.A. & Harden, G.J. (1990). Poranthera. In G.J.

Harden (ed.). Flora of New South Wales 1: 394.

New South Wales University Press:

Kensington.

Jeanes, J.A. (1999). Poranthera. In N.G. Walsh & T.J.

Entwisle (eds). Flora of Victoria 4: 71-72.

Inkata Press: Melbourne.

Kenneally, K.F., Edinger, D.C. & Willing, T. (1996).

Broome and Beyond: plants and people of

the Dampier Peninsula, Kimberley, Western

Australia. Department of Conservation and

Land Management: Como, Western Australia.

Kirkpatrick, J. (1997). Alpine Tasmania: an illustrated

guide to the flora and vegetation. Oxford

Press: Melbourne.

Klotzsch, J.F. (1848). Poranthera. In J.G.C. Lehmann

(ed.), Plantae Preissianae 2: 230-232.

Meissner: Hamburg.

Marchant, N.G. (1990). The Western Australian

collecting localities of J.A.L. Preiss.

In PS. Short (ed.). The history of systematic

botany in Australasia , pp. 131-136.

Australian Systematic Botany Society Inc.:

Melbourne.

McKee, H.S. (1963). The Bleeser botanical collection

from northern Australia. Contributions from

the New South Wales National Herbarium 3:

233-234.

Muller, J. (1866). Euphorbiaceae. In A.L.P.P. de

Candolle (ed.) Prodromus Systematis Naturalis

Regni Vegetabilis. 15(2): 189-214. Masson:

Paris.

Orchard, A.E. & Davies, J.A. (1985). Oreoporanthera,

a New Zealand “Endemic” plant genus

discovered in Tasmania. Papers and

Proceedings of the Royal Society of Tasmania

119: 61-63.

Orchard, A.E. (1999). A history of systematic botany

in Australia. Flora of Australia, 2 nd edn, 1:

11-104. CSIRO Publishing: Melbourne.

Rudge, E. (1811). A description of several species of

plants from New Holland. The Transactions

of the Linnean Society of London 10: 283-

303.

Schwarz, O. (1927). Plantae novae vel minus cognitae

Australiae tropicae. Repertorium Specierum

Novarum Regni Vegetabilis 24: 87.

Sonder, O.W. (1857). Plantae Muellerianae -

Euphorbiaceae. Linnaea 28: 562-567.

Webb, C.J., Sykes, W.R. & Garnock-Jones, PJ. (1988).

Flora of New Zealand 4: 623.

Botany Division, D.S.I.R.: Christchurch.

Weber, J.Z. (1986). Euphorbiaceae. In J.P Jessop &

H.R. Toelken (eds). Flora of South Australia,

4 th edn, 2: 735-768. South Australian

Government Printing Division: Adelaide.

Webster, G.L. (1994). Synopsis of the genera and

suprageneric taxa of Euphorbiaceae. Annals

of the Missouri Botanical Garden 81: 33-144.

Wheeler, J.R. (1992). Euphorbiaceae. In J.R. Wheeler

(ed.). Flora of the Kimberley Region, pp. 589-

629. Department of Conservation and Land

Management: Como, Western Australia.

Index to Scientific Names

Names in bold type are accepted names and

those in Roman font are synonyms or invalid

names.

Oreoporanthera Hutch. 3

Oreoporanthera petalifera Orchard & J.B.Davies 23

Poranthera arbuscula Sond. 5

Poranthera coerulea O. Schwarz. 4

Poranthera corymbosa Brongn. 5

Poranthera corymbosa Brongn. var. corymbosa. . 5

Poranthera corymbosa var. arbuscula (Sond.)

Mull.Arg. 5

Poranthera corymbosa var. genuina Mull.Arg. ... 5

Poranthera corymbosa var. linarioides Griming . . 5

Poranthera corymbosa var. sparsifolia Griming. . . 6

Poranthera dissecta Halford & R.J.F.Hend. 7

Poranthera drummondii Klotzsch. 4

Poranthera ericifolia Rudge . 10

Poranthera ericoides Klotzsch. 11

Poranthera ericoides Klotzsch var. ericoides .... 11

Poranthera ericoides var. glauca (Klotzsch)

Domin. 24

Poranthera ericoides var. typica Domin. 11

Austrobaileya 7 (1): 1-27 (2005)

Poranthera florosa Halford & R.J.F.Hend. 12

Poranthera glauca Klotzsch . 24

Poranthera huegelii Klotzsch. 13

Poranthera leiosperma Halford & R.J.F.Hend. . . 15

Poranthera linearoides Baill. 5

Poranthera microphylla Brongn. 16

Poranthera microphylla Brongn. var. microphylla 16

Poranthera microphylla var. diffusa Ewart. 17

Poranthera microphylla var. diffusa Mull.Arg. . . 16

Poranthera microphylla var. drummondii

(Klotzsch) Mull.Arg. 8

Poranthera microphylla var. genuina Mull.Arg. . 16

Poranthera microphylla var. glauca Mull.Arg. . . 17

Poranthera microphylla var. intermedia Mull.Arg. 12

Poranthera microphylla var. procera Griming ... 24

Poranthera obovata Halford & R.J.F.Hend. 19

Poranthera oreophila Halford & R.J.F.Hend. 20

Poranthera petaJifera (Orchard & J.B.Davies)

Halford & R.J.F.Hend. 22

Poranthera piceoides Klotzsch . 11

Poranthera Rudge. 3

Poranthera Rudge subgen. Poranthera . 3

Poranthera subgen. Euporanthera Griming . 3

Poranthera subgen. Oreoporanthera Griming. 3

Poranthera triandra J.M.Black . 23

Halford & Henderson, Revision of Poranthera

Maps 1-14. Distribution of Poranthera species.

A taxonomic revision of Tarenna Gaertn. and Triflorensia S.T.Reynolds

(Rubiaceae: Ixoroideae: Pavetteae) in Australia

S.T. Reynolds & Paul I. Forster

Summary

Reynolds, S.T. & Forster, P.I. (2005). A taxonomic revision of Tarenna Gaertn. and

Triflorensia S.T.Reynolds (Rubiaceae: Ixoroideae'. Pavetteae) in Australia. Austrobaileya 7(1):

29-55. Revisions are presented for the Australian representatives of Pavetteae excluding the

genus Pavetta L„ viz. Tarenna Gaertn. (three species, two subspecies) and Triflorensia S.T.Reynolds

(three species). The following are described as new, Triflorensia S.T.Reynolds genus nov., Tarenna

subgenus Pacifica S.T.Reynolds subg. nov. and Tarenna monticola S.T.Reynolds & RI.Forst. sp.

nov. New combinations are made in Tarenna for T. pentamera (Benth.) S.T.Reynolds and T.

dallachiana subsp. expandens (F.Muell. ex Benth.) S.T.Reynolds & P.I.Forst. and in Triflorensia

for three species previously included in Diplospora or Tarenna, viz. T. australis (Benth.)

S.T.Reynolds, T. cameronii (C.T.White) S.T.Reynolds and T. ixoroides (F.Muell.) S.T.Reynolds.

Lectotypes are selected for several names. All recognised taxa are described with notes on their

affinities, habitat, distribution, conservation status and etymology. Keys to taxa, distribution

maps and line drawings of some of the taxa are also provided.

Key Words: Rubiaceae, Tarenna subgenus Tarenna, Tarenna subgenus Pacifica, Tarenna dallachiana

subsp. expandens, Tarenna monticola, Tarenna pentamera, Triflorensia australis, Triflorensia

cameronii, Triflorensia ixoroides, Australian flora

S.T.Reynolds & RI.Forster*, Queensland Herbarium, Environmental Protection Agency, Brisbane

Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. *author for

correspondence. Email: paul.forster@epa.qld.gov.au

Introduction

The classification of the genera and

suprageneric taxa of Rubiaceae remains in a state

of flux in the light of proposed phylogenies

based on both morphological and molecular

data sets. The formal classification of genera

into subfamilies, tribes and subtribes within

Rubiaceae was last comprehensively reviewed

by Robbrecht (1988) with additions five years

later (Robbrecht 1993), but since then there have

been a number of wide ranging phylogenetic

studies, primarily using molecular data, with

results that have challenged this

morphologically based classification (e.g.

Andreasen et al. 1999; Bremer et al. 1999;

Andreasen & Bremer 2000).

In the current paper a revision is presented

of the Australian species of Tarenna Gaertn.,

together with the description and revision of a

new genus Triflorensia S.T.Reynolds that is

endemic to Australia and proposed to be closely

allied to Tarenna and also to Pavetta L. (see

Reynolds 1993). All three of these genera belong

Accepted for publication 30 June 2005

in the subfamily Ixoroideae , with both Pavetta

and Tarenna being grouped with Dictyandra

Welw. ex Hook.f., Leptactina Hook.f. and

Rutidea DC. in the tribe Pavetteae A.Rich. ex

Dumort. (tribal description given below) on the

basis of both morphological and molecular data

sets (Andreasen & Bremer 2000). In the earlier

classification of Robbrecht (1988), Pavetta and

Tarenna, together with Duperrea Pierre ex

Pitard., Ivor a L., Myonima Comm, ex Juss. and

Versteegia Valeton were all included in a

polyphyletic Pavetteae.

Tarenna has long been treated in a broad

sense by various authors including Bridson

(1979), Bridson & Robbrecht (1985) and Smith

and Darwin (1988). Bridson (1979) recognised

six informal groups under Tarenna s.l. based

mainly on placentation and seed types, for the

African species. The Australian species fall into

three ‘groups’ based more or less on the same

characters used by Bridson (1979), two of these

are retained under Tarenna , whereas one is

transferred to the closely related genus

Triflorensia.

Coffea L., Discospermum Dalzell,

Psilanthus Hook.f. and Tricalysia A.Rich. ex

DC. (Andreasen & Bremer 2000). An account of

the Australian genera and species of Coffeeae

was provided by Forster (2004).

The three species transferred to

Triflorensia have had a chequered

nomenclatural history, starting in Diplospora

DC. (Bentham 1867; Mueller 1875; White 1926)

and most recently being combined in Tarenna

(Ali & Robbrecht 1991). The genus Diplospora

is now restricted to species found in Asia and

Malesia and is included in the tribe Coffeeae

DC. together with Bertiera Aublet., Coffea L.,

Discospermum Dalzel 1, Psilanthus Hook.f. and

Tricalysia A. Rich. ex DC. (Andreasen & Bremer

2000). An account of the Australian genera and

species of Coffeeae was provided by Forster

(2004).

Materials and Methods

The initial draft of this revision was undertaken

by the first author prior to 1998. It has been

updated in 2005 by the second author,

particularly in terms of introductory information

about recent generic phylogenies and new data

pertaining to species distribution, habitat and

conservation status.

The revision is based on morphological

characters derived from herbarium specimens

(dried sheets, spirit material) at BRI, CANB,

DNA, K, NSW and QRS examined by the first

author, with supplementary observations and

measurements by the second author of

specimens accessioned at BRI since 1995.

Measurements encompass the main range of

variation, with aberrant or unusual data indicated

in brackets, e.g. (8-) 12-15.

Common abbreviations in the specimen

citations are L.A. (= Logging Area), N.R (=

National Park), S.F. (= State Forest), S.F.R. (=

State Forest Reserve) and T.R. (= Timber

Reserve).

Species are defined as groups of

populations (1-many) with discontinuities in the

variation of two or more independent character

states of morphology. It is assumed that there

is genetic continuity or at least a shared ancestral

lineage between the different populations of a

Austrobaileya 7 (1): 29-55 (2005)

single species. Although species definitions

remain a matter of opinion and debate, the one

applied here (‘phenetic species concept’) is

widely used and understood (Stebbins 1950;

Cronquist 1988; Stuessy 1990; Levin 1979,2000)

and would also equate to the ‘diagnostic

species concept’ of Judd etal. (2002). Characters

most commonly used in the identification keys

are those of inflorescence morphology, leaf

lamina shape, flower morphology (particularly

shape, size and indumentum cover) and fruit

size and shape.

If only a single, or several minor (e.g.

indumentum colour or density) character state

differences are present and the discontinuity is

geographically based, the rank of subspecies

is used. It is generally considered that

subspecies should differ in only a few minor

characters and that intermediate populations

should exist to demonstrate continuity of

character states (e.g. Stebbins (1950) states

“subspecies....connected....by a series of

intergrading forms” or Stace (1989) states “a

population of several biotypes forming a more

or less distinct regional facies of a species.a

geographical race, ecotype, topodeme or

genoecodeme”). In this revision subspecies are

recognised within Tarenna dallachiana due to

the existence of intermediate forms.

Taxonomy

Tribe PavetteaeA. Rich, ex Dumort., Anal. Fam.

PI. 33 (1829); Bridson & Robbrecht, Bull.

Nat. PlantentuinBelg. 55: 83-115 (1985);

Andreasen & Bremer, Amer. J. Bot. 87:

1743(2000).

Shrubs, small trees or lianas; stipules

interpetiolar, entire; leaves entire, without

raphides; inflorescences terminal on main or

lateral branches, rarely pseudo-axillary,

trichotomously branched. Flowers 4 or 5-

merous; corolla lobes contorted to the left;

corolla tubes long and slender tubes (usually

much longer than its lobes), anthers affixed at

throat and exserted; stylar pollen presentation

present; pollen grains 3-4(-5) colporate; style

exserted or rarely included in the corolla tube,

the stigmas fusiform, clavate, not lobed; ovary

2-locular, axile placentation with 1-many ovules,

the ovules either attached to the central septum,

or embedded on a fleshy placenta which is

Reynolds & Forster, Revision of Tarenna and Triflorensia

attached to the dissepiment usually near the

middle. Fruits fleshy, 1-many seeded; seeds

either 4-many in each cell of the fruit, then very

small, angular and reticulate, or seeds solitary

or 2 in each cell of the fruit, then large ±

hemispherical, rounded dorsally, and with an

adaxial excavation on the ventral side.

Distribution : Mainly Paleotropical. About six

genera, three (one new) in Australia.

Key to the Australian genera of Pavetteae

1. Stipules connate to above its middle into an oblique sheath;

inflorescences terminal on main and lateral branches; bracts at base of

trichotomous inflorescence branches connate, membranous; ovules and

seeds one in each cell of the ovary or fruit; fruits without a thin

crustaceous shiny endocarp.Pavetta

Stipules slightly united at base only; inflorescences terminal on main

branches; bracts at base of trichotomous inflorescence branches free or

connate towards their base; ovules an seeds 1-many in each cell of the

ovary or fruit; fruits with a thin crustaceous shiny

endocarp.2

2. Inflorescence branches terminated by 3-flowered dichasial cymes; bracts

at base of the trichotomous branches of the inflorescence shortly connate;

bracteoles paired, one on each side of the calyx; flowers 4 or 5-merous;

seeds 2 in each locule of the fruit, rounded on top, angular on its sides;

stipules usually obtuse.Triflorensia

Inflorescences branches not terminated by 3-flowered dichasial cymes;

bracts at base of inflorescence branches free; bracteoles 3 or 4, alternate

on the pedicel (one on calyx); flowers 5-merous; seeds either small and 4-

many in each locule of the fruit (if so, then angular, reticulate (like orange

segments)), or large and solitary in each cell of the fruit (if so,

thensubglobose, rounded on top and with a thick rimmed circular

excavation on its ventral side); stipules acuminate.Tarenna

Tarenna Gaertn., Fruct. Sem. 1:139, t. 28 (1788).

Type: T. asiatica (L.)KuntzeexK.Schum.

(based on Rondeletia asiatica L.; syn. T.

zeylanica Gaertn.)

Chomelia L., Opera Varia 210 (1758) nom.

rej.

[Webera auct., non Schreber: Bentham, FI.

Austral. 3: 412 (1867); F.M.Bailey,

Queensland FI. 3: 752(1900)].

[Stylocoryne auct., non Cav.: Domin,

Biblioth. Bot. 617(1929)]

Shrubs or small trees; branchlets terete or 4-

angled towards their tips. Leaves petiolate, often

blackish when dry; stipules persistent, erect,

free or connate towards base, ovate or

triangular-ovate, sometimes articulated towards

base, acuminate or aristate at apex, usually with

a darker central area and paler margins, provided

with a prominently bifid keel (the keel bifid

towards base) and with collectors towards its

base inside. Inflorescences terminal, many-

flowered, corymbiform panicles, trichotomously

branched, the three branches clustered on a

short peduncle or clustered at apex of the

branchlet subtended by the uppermost pair of

leaves; lowermost branches usually patent;

ultimate cymules 2-5-flowered; bracts free or

slightly connate (towards the base), and with

very narrowly lanceolate lateral lobes. Flowers

bisexual, 5 (rarely 4)-merous, fragrant;

pedicellate; bracteoles small, alternate on the

pedicels, subulate or ovate. Calyx tube

subcyathiform or cupular; limb wider and

thinner, 5 (rarely 4)-lobed at apex, with lobes

imbricate, ovate or hemispherical. Corolla white;

tube infundibuliform or cylindrical and usually

dilated towards apex, longer or shorter than the

lobes, usually hairy at throat; corolla lobes 5

(rarely 4), twisted to the left in bud, ± oblong,

patent or reflexed, hairy inside. Stamens 5 (rarely

4), inserted at throat (slightly below the orifice

of corolla tube); anthers long, narrowly ovoid,

with a prolonged ovate connective at apex,

slightly sagittate at base, dorsifixed; filaments

much shorter, glabrous. Disc fleshy, annular.

Ovary 2-locular; ovules 1-many per locule,

ovoid, attached to septum or impressed on the

semicircular placenta which is attached to the

septum; style (with stigma) shortly exceeding

the corolla tube or long exserted; style filiform,

hairy towards middle or glabrous; stigma long,

fusiform or clavate, with lobes coherent for most

of stylar length or stigma bifid at apex. Fruits

shiny, usually blackish when ripe, subglobose,

crowned by persistent calyx, 2-locular; seeds

1-many per locule, if many they are blackish,

angular, rugulose and closely reticulate

resembling orange segments, or if solitary,

subglobose, smooth or finely reticulate on top,

and with a circular excavation (hilar cavity) on

their ventral side; pericarp smooth outside;

endocarp thin, crustaceous, very shiny and

brown.

Austrobaileya 7 (1): 29-55 (2005)

Tarenna has c. 370 species (Wong 1989;

Ridsdale 1998) in tropical Africa, Asia, Pacific

Islands, New Guinea and Australia (three

species).

Etymology: The name Tarenna is derived from

Tarana ” a Sri Lankan plant name.

Diagnostic features: The genus Tarenna (in

Australia) is characterised by its terminal,

trichotomous corymbiform panicles, (4-) 5-

merous flowers, styles with fusiform stigmas,

small, globose, 1-many seeded fruits with a thin,

shiny, crustaceous endocarp, usually blackish

discoloration of its leaves and inflorescences,

4-angular young branchlets, and triangular

acuminate stipules with softer paler margins and

a bifid keel. Tarenna may be distinguished from

other members of the tribe Pavetteae in

Australia, viz. Pavetta and Triflorensia as

indicated above.

Infrageneric Classification

Three subgenera are recognised under Tarenna

here: two of these were previously recognised

as sections by Hooker (1882: 104) and Valeton

(1924: 84). Only the subgenera Psendoixora and

Pacifica occur in Australia.

Key to subgenera

1. Ovules solitary in each cell of the ovary, attached to the septum; seeds

solitary in each cell of the fruit, large, smooth, subglobose, with a thick-

rimmed circular excavation on its ventral side. 3. Pseudoixora

Ovules 4-many in each cell of the ovary, impressed on the suborbicular

placenta which is attached to the septum; seeds 4-many per fruit, small,

angular, densely reticulate, resembling orange segments.2

2. Branchlets usually stout, 4-angular towards their apices; stipules large,

triangular, acuminate or aristate at apex, darker in the centre and with paler

soft margins; inflorescences large, many-flowered; calyx lobes ±

hemispherical or ovate; corolla tube long and slender, usually much longer

than its lobes; style usually much exserted from the corolla tube.2. Pacifica

Branchlets usually slender, terete or ± 4-angular towards their apices; stipules

ovate, cuspidate at apex, coriaceous; inflorescences smaller, loosely

branched and flowered; calyx lobes ovate; corolla tube usually as long as

its lobes; style usually slightly exserted from the corolla tube.1. Tarenna

Reynolds & Forster, Revision of Tarenna and Triflorensia

1. Tarenna subgenus Tarenna

Type: Tarenna asiatica (L.) Kuntze ex K.Schum.

(syn. T. zeylanica Gaertn.).

Weber a sect. Webera Hook f., FI. Brit. India

3: 104(1882). Type: not designated.

Distinguishing characters as indicated in the

key above.

Distribution : Mainland Asia (China, Indian

subcontinent) and south-east Asia (Wong 1989;

Ridsdale 1998).

Notes : Tarenna appressa (King) Corner from

India and T. lanceolata Chun & How ex

W.C.Chen from China also belong to this

subgenus {fide specimens at K, L).

2. Tarenna subgenus Pacifica S.T.Reynolds

subgenus nova quoad placentam et

semina Tarennae subg. Tarennae similis

autem inflorescentiis grandioribus

multifloris, tubis corollae lobis multo

longioribus, lobis calycem

hemisphericalis, stipulis magnis

triangularis marginibus pallentioribus,

mollioribusque praeditis, ramulis 4-

angularibus differt. Typus: Tarenna

sambucina (Forst.f.) Durand ex Drake

(syn. Coffea sambucina Forst.f.).

Tarenna sect. Multiovulatae Valeton in

Lauterbach, Beitr. FI. Papuasien 13, Bot.

Jahrb. 60: 84 (1924). Type: Tarenna

sambucina (Forst.f.) Durand ex Drake

Trees or shrubs with stout, 4-angular

branchlets; large broad triangular stipules

which are acuminate or aristate at apex, and

usually provided with a thicker darker area in

the centre and softer paler edges. Inflorescences

large, usually many-flowered; flowers with long,

slender corolla tubes which are usually much

longer than its lobes; usually ± hemispherical

calyx lobes; long styles which are usually much

exserted from the corolla tube; fusiform stigmas;

ovary with 4-many ovoid ovules which are

impressed on the fleshy placentas; 4-many-

seeded fruits with small, angular reticulate seeds.

Notes : Tarenna subgenus Pacifica has the

ovules, placenta and seeds of Tarenna

subgenus Tarenna , but differs from it by its

stout 4-angular branchlets, the large triangular

stipules, large many-flowered inflorescences,

and long narrow corolla tubes which are usually

much longer than its lobes (see key to

subgenera).

Two Australian species, viz. Tarenna

dallachiana (F.Muell. ex Benth.) S.Moore and

T. monticola S.T.Reynolds & RI.Forst. are

referrable to this subgenus.

Distribution : Australia, New Guinea and islands

of the Pacific and Indian oceans. Two species

occur in Australia {T. dallachiana and

T. monticola), one species in the Pacific Ocean

{T. sambucina), one species in New Guinea

(T. barbellata Valeton) and one species from

the Indian Ocean (T. buruensis Merrill).

Etymology : The name of the subgenus alludes

to the Pacific Region where the species

referrable to this subgenus are commonly found.

Key to the Australian species of Tarenna subgenus Pacifica

Corolla hairy outside, tube 8-22 mm long, lobes erect; seeds 4-16 per fruit,

very rarely more; domatia in angle between lateral veins and midrib absent

below; leaf apex obtuse, subacute or shortly acuminate. 1. T. dallachiana

Corolla glabrous outside (rarely subglabrous), tube 4-7 mm long, lobes

mostly recurved; seeds 14-26 per fruit; tufts of hair in angle between

lateral veins and midrib present below; leaf apex mostly long acuminate . . 2. T. monticola

1. Tarenna dallachiana (F.Muell. ex Benth.)

S.Moore, J. Bot. 64: 94 (1926). Webern

dallachiana F.Muell. ex Benth., FI.

Austral. 3:412(1867); Ixora dallachiana

(F.Muell. ex Benth.) F.Muell., Syst. Census

Australian Plants 1: 75 (1882); Chomelia

dallachiana (F.Muell. ex Benth.) O.Ktze,

Rev. Gen. PI. 1: 278 (1891); Stylocoryna

dallachiana (F.Muell.) Domin, Biblioth.

Bot. 89: 617 (1929). Type: Queensland.

Cook District: Albany Island, date

unknown, W.Hill s.n. (lecto [here

designated]: K; photo BRI).

Small trees 2-8 m high, bark grey or mottled

grey and white, smooth or tessellated; young

branchlets and leaves finely hairy to glabrous;

branchlets and petioles sometimes scurfy.

Leaves petiolate; lamina elliptic, ovate-elliptic,

broadly elliptic or elliptic-oblong, apex abruptly

shortly acuminate, subacute or obtuse, base

obtuse or subacute and attenuate into petiole,

(12-) 14-25 (-27.5) x (3.7-) 5.5-9 (-13) cm,

coriaceous, dries blackish or brownish, greenish

or greenish brown and sometimes slightly

glossy above; upper surface glabrous or hairy

on the main nerves; lower surface usually

pubescent with usually fine hairs all over

(especially young leaves), or hairy only on the

main nerves; lateral nerves 8-14 pairs,

suboblique or patent, mostly arched at their tips;

petioles (1-) 2-3.5 (-5) cm long, sparsely hairy

or glabrous. Stipules 6.5-11 x 3-6 mm, triangular

or ovate, gradually narrowing at apex to a

subulate acumen or cuspidate, usually with a

darker centre and paler softer margins,

puberulent or glabrous, sometimes slightly

viscid, usually with thick, viscid colleters at base

inside. Inflorescences usually densely flowered

(38-152-flowered); the 3 branches subtended

by the uppermost pair of leaves of the

branchlets; peduncles sparsely short hairy;

central branch with 1 or 2 internodes (these 1—

2.2 cm long); lateral ones 2-9 cm long, 12-152-

flowered; ultimate branches 1-1.7 cm long,

terminated by 2 or 3-flowered cymules or 9-24-

flowered cymes; bracts subtending the

inflorescence stipular, ovate, those at junction

of the upper branches subulate; bracteoles 2-

4, alternate on the pedicel, minute (about 1.5

mm long), narrowly ovate or subulate, ciliolate.

Flowers perfumed; pedicels 2-9 mm long. Calyx

1.5-2.5 x 1-2 mm; tube glabrous or puberulous,

dries blackish; limb paler, indistinctly toothed;

calyx lobes small, 0.25-0.5 mm long, ±

hemispherical, obtuse, truncate or retuse at apex,

Austrobaileya 7 (1): 29-55 (2005)

ciliolate. Corolla tube cylindrical, gradually

dilated from base to the mouth or dilated near

mouth only, (8-) 12-22 mm long, 1-3 mm wide

at mouth, 0.7-1.5 mm wide at base, straight or

slightly curved, densely appressed hairy

outside, sparsely appressed hairy inside; corolla

lobes usually erect, elliptic, oblong obovate or

subobovate, 3.5-6.5 x 1.5-3 mm, retuse or

obtuse; disc glabrous. Stamens exserted;

filaments 1-1.8 mm long; anthers 3.5-6.5 mm

long. Styles 2.3-4.5 mm long, exserted (exserted

portion (from corolla tube) 1-1.8 cm long), hairy

towards the middle (from below stigma); ovary

2 locular; ovules (2-) 4-8 in each loculus, ovoid,

impressed on the ± ovoid fleshy placenta which

is attached to the septum. Fruits globose,

topped by remnant calyx rim, dull olive, black

when ripe, 6.5-8 x 6-8 mm; seeds (4—) 8-16 per

fruit, small, c. 1x2 mm, angular, reticulate, black;

fruit stalk 8—12 mm long.

Typification: Two syntypes were originally

cited by Bentham (viz. Rockingham Bay,

Dallachy s.n. (syn: MEL) and Albany Island,

Hill s.n. (syn: K). The Hill collection is the better

of the two and is selected here as lectotype for

the name.

Notes: Tarenna dallachiana is readily

distinguishable by its usually large, glabrous

or finely hairy, elliptic leaves which dry

greenish, greenish-brown or blackish, large

triangular stipules with soft edges and a bifid

keel, 4-angular, darkish, usually scurfy young

branchlets, many-flowered corymbiform

inflorescences, ± hemispherical ciliolate calyx

lobes, long corolla tube (tube dilated near apex

or gradually tapering from mouth to base) which

are appressed hairy outside, short, oblong

corolla lobes, subsessile anthers which are

nearly as long as the corolla lobes, and 4-14

seeded fruits.

The shape and size of leaves, the colour

of dried leaves, the presence or absence of hairs

on young branches and young leaves, and the

length and shape of corolla tube are all very

variable in the specimens available for study of

this species. Two subspecies are recognised

here that merge into each other at the outer parts

of their respective ranges on Cape York

Peninsula.

Etymology : The specific epithet honours the

botanical collector John Dallachy (1808-1871)

who collected many specimens for Ferdinand

Mueller.

Reynolds & Forster, Revision of Tarenna and Triflorensia

Key to subspecies of Tarenna dallachiana

1. Corolla tube 8-12 long, usually gradually dilated towards its mouth (2-3 mm

wide at mouth); pedicels 2-5 mm long; seeds 4-8(-12) in each fruit; leaves

12-25 (-27.5) x 3.7-9.0 (-10.7) cm, dries blackish or dark brown, both

surfaces glabrous or lower surface sparsely hairy on the main nerves;

young branchlets and young leaves sparsely hairy or glabrous

. la. Tarenna dallachiana subsp. dallachiana

Corolla tube 14-22 mm long, cylindrical, usually dilated only towards its

throat (1-2.5 mm wide at mouth); pedicels (2-) 4-9 mm long; seeds (6-

(6-) 8-12 (-16) in each fruit; leaves (10-) 15-22 (-24) x (3-) 6-9.5 (-13)

cm, dries greenish or pale brown; upper surface glabrous or subglabrous,

lower ones covered with soft hairs, or occasionally only the main nerves

hairy; young branchlets and young leaves densely hairy with fine soft

hairs (usually velvety). lb. Tarenna dallachiana subsp. expandens

la. Tarenna dallachiana subsp. dallachiana

Distinguishing characters as indicated in the

key above.

Illustrations : Hyland etal. (1999,2003); Cooper

& Cooper (2004).

Selected specimens examined: Queensland. Cook

District: Upper Parrot Creek, Annan River, 15° 48’S,

145° 15’E, Sep 1948, Brass 20293 (BRI); Mt Amos,

Cooktown, 15° 42’S, 145° 18’E, Nov 1986, Burkitt 61

(BRI); Fly Point, about 2 km SE of Somerset, 10°

45’S, 142° 36’E, Jul 1992, Clarkson 9646 & Neldner

(BRI); Jardine River National Park, 11° 20’S, 142°

45’E, Aug 1987, Fell DGF587 (BRI); Rex Range, Little

Mossman S.F. 141, 16° 32’S, 145° 23’E, Jan 1993,

Forster PIF13071 & Bean (BRI, QRS); Home Rule, 3

km E of Rossville, 15° 45’S, 145° 17’E, Apr 1999,

Forster PIF24247 & Booth (A, BISH, BRI, K, L, MEL,

QRS); Freshwater F.R., Lake Morris road, 16° 56’S,

145° 42’E, May 2005, Forster PIF30957 & Jensen

(BRI, L, MEL, MO, NSW); track to Mt Misery from

Shiptons Flat, 15° 49’S, 145° 13’E, Jun 2005, Forster

PIF31010 & Jensen (BRI, L, MEL, NSW); Claudie

River, 12° 43’S, 143° 17’E, Oct 1981, Hyland 11179

(BRI); Upper Escape River, 11° 10’S, 142° 45’E, Aug

1976, Kanis 2079 (BRI); Endeavour River, 1883, Persieh

25 (MEL); Between Mulgrave and Russell Rivers, 1887,

Sayer (BRI); Lower slopes of Mt Saunders, 15° 25’S,

145° 14’E, Mar 1984, Scarth-Johnson 1410A (BRI);

Bamaga, Oct 1965, Smith 12395 & 12581 (BRI);

Mourilyan, Feb 1923, White (BRI). North Kennedy

District: Clump Point, Feb 1972, Crome 263 (CANB,

QRS); Rockingham Bay, Dallachy s.n. (MEL 1538176,

NSW 19378 - residual syntypes); Strathdickie,

Proserpine, Michael 1264 & 1387 (BRI); Clump Point,

Dec 1969, Winkel s.n. (BRI). South Kennedy District:

Port Mackay, Dietrich 375 (MEL); loc. cit., 1885,

Marguart s.n. (NSW 193797); loc. cit., Oct 1882,

Griffith 93 (BRI). Port Curtis District: By field near

Keppel Bay, Sep 1931, White 8169 (BRI).

Distribution and habitat: Eastern Queensland,

chiefly coastal, from Cape York Peninsula to

Keppel Bay; usually on ridges, gullies, hillsides

and sometimes on river levees, mostly at the

margins of rainforests (Map 1).

Notes: Tarenna dallachiana subsp.

dallachiana is characterised by the darkish

discoloration of its dried leaves and branchlets,

glabrous or sparsely hairy branchlets (usually

with antrorse hairs) and corolla tubes which are

usually dilated from base to apex. The shape

and size of leaves and presence of hairs are

variable in this subspecies and a few forms

appear to be present.

This subspecies usually has scurfy

branchlets, thickly coriaceous, large or middle-

sized, slightly glossy leaves that dry blackish

or brown, and flowers with wide corolla tubes.

Specimens with these characteristics, especially

those from around Proserpine and Mackay,

resemble the syntype from Rockingham Bay.

Specimens from Cape York Peninsula

show considerable variation. Some resemble the

above, whereas others with larger elliptic leaves

that dry brown, and with glabrous branchlets

and leaves resemble the syntype from Albany

Island.

Specimens from Bamaga and Rex Range

differ from the above in their longer, usually

larger leaves, sparsely hairy young branchlets

and young leaves and approach specimens

from around Weipa included under subsp.

expandens.

Austrobaileya 7 (1): 29-55 (2005)

Fig. 1 . Tarenna dallachiana subsp. expandens. A. branch of inflorescence *1. B. flower x2. C. longitudinal section

of flower x2. D. style and stigma x2. E. calyx with bracteole x8. F. longitudinal section of ovary showing ovules

on placenta xl6. G. fruit M. H. seed xl2. All from Booth R241 (BRI)). Del. W. Smith.

Specimens included here that were

collected from the Annan and Claudie Rivers

appear to be quite distinct. They differ from the

others in their smaller usually broad elliptic

leaves, glabrous leaves and branchlets and a

slightly different aspect. All of these specimens

seen are in fruit and their affinities are unclear

at this time.

lb. Tarenna dallachiana subsp. expandens

(F.Muell.) S.T.Reynolds &P.I.Forst. comb,

et stat. nov.; Weber a expandens F. Muell.,

Fragm. 6: 25 (1867); W. dallachiana var.

expandens (F.Muell.) F.Muell., Fragm. 9:

187 (1875); Ixora expandens (F.Muell.)

F.Muell., Syst. Census of Australian

Plants 1:74 (1882); Tarenna expandens

(F.Muell.) S.Moore, J. Bot. 64: 94(1926).

Type: Northern Territory. Adam’s Bay, date

unknown, Hulls s. n. (holo: MEL 1533300

[flowering specimen]).

Tarennafoliosa S.Moore, J. Bot. 64:94 (1926).

Type: Northern Territory. Groote Eylandt,

Feb 1925, S.Moore 116 (holo: BM

[flowering specimen], syn. nov.

Reynolds & Forster, Revision of Tarenna and Triflorensia

Distinguishing characters as given in the key

above. Fig. 1.

Selected specimens examined : Western Australia.

11 km N of King Cascade, 15° 31’S, 125° 19’E, Jun

1987, Kenneally 10556 & Hyland (BRI, PERTH).

Northern Territory. Goromuru River floodplain, 12°

37’S, 136° 14’E, May 1992, Cowie 2843 (BRI, DNA);

Magela Creek upper Catchment, 12° 49’S, 133° OO’E,

Apr 1995, Cowie 5659 & Brennan (DNA); Daly River,

13° 42’S, 130° 33’E, Sep 1975, Dunlop 3595 (DNA);

Garden Point, Melville Island, Sep 1977, Dunlop 4622

(BRI, CANB, DNA); Arnhem Land, Mt Permain, 11°

56’S, 132° 58’E, Oct 1987, Dunlop 7023 (BRI, DNA);

Upper Goomadeer River, 12° 40’S, 133° 34’E, Oct

1987, Dunlop 7227 & Munns (BRI); 5 km SE Lake

Eames, Vanderlin Island, 15° 42’S, 137° 04’E, Aug

1988, Latz 11079 (BRI); Headwater Springs, Green Ant

Creek, 13° 30’S, 131° 14’E, Mar 1989, Leach 2547 &

Dunlop (BRI); Melville Island, Yapilika tributary of

Maxwell Creek, 11° 33’S, 130° 35’E, Jan 1992, Leach

2952 & Cowie (BRI, DNA); Little Angurugu River,

Groote Eylandt, Sep 1973, Levitt 353 (DNA);

Nourlangie Rock area, 12° 57’S, 132° 50’E, Nov 1972,

McKean B774 (DNA); Darwin, Feb 1961, McKee 8263

(BRI, MEL, NSW); Hapgood River, Gapuwuyak, 12°

37’S, 135° 50’E, Sep 1987, Russell-Smith 3432 &

Lucas (BRI, DNA); Bathurst Island, Pimaroo Creek,

11° 23’S, 130° 22’E, Jun 1988, Russell-Smith 5783 &

Lucas (BRI, DNA); 19 km E Jabiru, 12° 37’S, 133°

03’E, Apr 1989, Russell-Smith 8020 (DNA); Mt

Brockman, Kakadu National Park, 12° 50’S, 132°

54’E, Apr 1989, Russell-Smith 8063 & Lucas (BRI,

DNA); Adelaide River crossing, Daly River road, 13°

30’S, 131° 04’E, Dec 1990, Russell-Smith 8372 &

Lucas (BRI, CANB, DNA, K, MEL). Queensland.

Cook District: 30.9 km from the beach at Ussher Point

on the track to the main Peninsula road, 11° 05’S,

142° 35’E, Jul 1992, Clarkson 9689 & Neldner (BRI);

Weipa Concession, N bank of Mission River, Sep 1974,

Dockrill 882 (BRI, QRS); Andoom Bridge area, near

Weipa, Jan 1989, O Reilly 110 (BRI); 11.5 km N of

Weipa Mission, 12° 35’E, 141° 54’S, Dec 1974, Specht

& Salt W113 (BRI).

Distribution and habitat : Northern tropical

Australia from around Darwin to Arnhem Land

in the Northern Territory and near Weipa, on

Cape York Peninsula, Queensland. There is a

single (sterile) collection from Western Australia

in Prince Regent Nature Reserve. Plants usually

grow at the edge of swamps, mangroves and

semi-deciduous vine forests, near permanent

water (viz. spring fed soakage pockets along

creeks and drainage lines and creek line) on

gorges and sandstone ridges, on sandy clay

loam (Map 1).

Notes : Mueller (1867) described Tarenna

expandens (as Webera expandens) as a distinct

species but later reduced it to a variety of Webera

dallachiana (Mueller 1875). Infraspecific rank

for T. expandens is maintained here, because

although the types of these names are quite

different they are connected by intermediate

forms, particularly on northern Cape York

Peninsula. Recent checklists of the Northern

Territory flora have listed this subspecies as

simply T. dallachiana (Dunlop 1987; Liddle et

al. 1994).

Tarenna dallachiana subsp. expandens

is characterised and distinguished from

T. dallachiana subsp. dallachiana by hairy

branchlets and leaves (especially velvety young

leaves), usually greenish or greenish brown

colour of dried leaves (the green colour usually

persisting in the herbarium for a very long time),

flowers with long very slender corolla tubes and

long pedicels. This subspecies varies greatly

in the shape and size of leaves and

inflorescences, hairiness of leaves and

branchlets, colour of dried leaves, and the

number of nerves.

Several forms are distinguishable in the

specimens available for study. The typical form

is represented by specimens from around

Darwin and Melville Island. They resemble the

type specimen of T. expandens in their short,

broadly elliptic or elliptic-oblong leaves which

are usually broad and obtuse at the base, finely

hairy on their lower surface and usually dry

greenish or greenish brown.

Specimens from eastern Amhem Land and

the islands of the Gulf of Carpentaria are variable

in their leaves. Some of these with larger, hairier

leaves resemble the type of T. foliosa (this

species now combined under this subspecies),

whereas those with relatively narrower leaves

are not unlike those from northern Queensland

(viz. Bamaga, Rex Range), except that their

young leaves are velvety, lateral nerves more

oblique, and their petioles longer.

The following collections viz. Dunlop

3595 from Mt Permain, Arnhem Land, Russell-

Smith 8020 from East Jabiru, Dunlop 7227 &

Munns from Upper Goomadeer River, and

McKean B774 from Nourlangie Rock probably

represent a distinct form but specimens are

inadequate to ascertain this at the present. The

specimens differ from the other collections of

this variety in their hairier, usually smaller, elliptic

leaves (10—17 (—19) x 3-4.4 (-7) cm), which are

usually narrow at both ends (mostly acuminate

or acute at apex, acuminate and decurrent into a

long petiole at base); comparatively greater

number of nerves ((9-) 10-14 pairs) on each

leaf, usually very oblique nerves; longer petioles

(1.5-5 cm long), and 8-16-seeded fruits.

Conservation status : Tarenna dallachiana

subsp. expandens is widespread and not

threatened. It is present in Kakadu National Park

in the Northern Territory and Prince Regent

Nature Reserve in Western Australia.

Etymology : The subspecific epithet is derived

from the Latin word expansus (expanding), but

the application is obscure.

2. Tarenna monticola S.T.Reynolds & PI.Forst.

sp. nov. a T. dallachianae (F.Muell. ex

Benth.) S.Moore floribus pedicellisque

brevioribus, corollis glabris, fructibus 14-

26-seminalis, foliis apice acuminatis

domatiis prominentibus pubescentibus

differt. Typus: Queensland. Cook District:

Riflemead Forest Reserve, 2.2 km along

Mt Lewis road from Bushy Creek bridge,

16° 34’S, 145° 18’E, 5 December 2002, A.

FordAF3781 & J. Holmes (holo: BRI [2

sheets + spirit]; iso: QRS [1 sheet + spirit];

K, L, NSW to be distributed).

Tarenna sp. (Mt Lewis B.P.Hyland 3403)

(Reynolds 1997: 184; Forster & Halford

2002:177)

Small slender trees to 16 m; young parts

especially young leaves densely finely hairy;

branchlets with fine appressed hairs at their tips

only. Leaves petiolate; lamina elliptic, abruptly

narrowing at both ends, apex usually long-

acuminate at apex, base acute or acuminate and

decurrent into the petiole, 11.5-19.5 (-22) x 5-

8.5 cm, thinly coriaceous, dries greenish, greyish

or darkish brown with whitish or reddish nerves;

upper surface dullish, glabrous, lower surface

sparsely appressed hairy on the midrib and

usually with prominent domatia in the axils of

the main nerves; midrib sunken above; lateral

nerves 8-11 pairs, suboblique and ascending

at their tips; petioles 1-3 cm long. Stipules

deltoid, long acuminate or aristate at apex and

Austrobaileya 7 (1): 29-55 (2005)

thinner at margins, 8-9 x 4.5-5 mm, appressed

hairy outside, sometimes viscid inside.

Inflorescences trichotomously branched,

densely flowered; peduncles with short patent

hairs; lowermost branch (lateral) 1.8-3 cm long,

29-64-flowered; bracts lanceolate or very

narrowly ovate; bracteoles 2 or 3 per pedicel

(one at base of calyx), narrowly ovate or slightly

subulate, usually ciliolate. Flowers glabrous;

pedicels (1-) 2-5 mm long, puberulous. Calyx

2.5-3.5 xc.2 mm, glabrous or lobes sometimes

subglabrous; calyx lobes prominent, ovate,

obtuse, 0.8-1 x 0.5-0.8 mm. Corolla 8-12.5 mm

long, glabrous or rarely subglabrous outside,

sparsely appressed hairy inside; corolla tube

4-7 mm long, dilated gradually to mouth, 1.5-

2.5 mm wide at mouth, c. 1.5 mm wide at base;

corolla lobes 2.5-4 xc.2 mm, oblong, obtuse,

reflexed. Stamens exserted with filaments to 1.5

mm long and anthers 3.5-5 mm long. Style and

stigma 15-20 mm long, exserted, with exserted

part (from mouth of corolla tube) 6-10 mm long;

style sparsely long hairy in the upper half;

stigma fusiform c. 2 mm long; ovules 7-13 in

each loculus of the ovary, impressed on the

fleshy placenta which is attached to the septum.

Fruits 4-5 x 5-6 mm, subglobose, (14-) 20-26-

seeded; seeds very small, angular, reticulate;

fruit stalk 4-7 mm long. Fig. 2.

Selected specimens examined : Queensland. Cook

District: Copper Lode Falls Dam area Cairns, Dec 1972,

Birch 93 (BRI); T.R. 66, 5.5 km along Mt Lewis road,

16° 35’S, 145° 18’E, Nov 1998, Forster PIF23995 et

at. (BRI); Mt Misery road. East Normanby River, 15°

52% 145° 12’E, Nov 2000, Forster PIF26485 et at.

(A, BRI, MEL, NSW); S.F.R. 143, Windmill L.A., 16°

33’S, 145° 15% Nov 1978, Gray 1067 (BRI, QRS);

[S.JF.R. 1073, 16° 45’S, 145° 30’E, Dec 1961, Hyland

2208 (BRI); Mt Lewis, T.R. 130, Leichhardt L.A., c.

40 miles (64 km) W of Cairns, Dec 1964, Hyland

3403 (BRI); Kuranda, 16° 49’S, 145° 38% Jul 1959,

Jones 1286 (BRI, CANB); Clohesy River (cult. Tolga),

Dec 1991, Sankowsky 1366 (BRI); 3 miles W of

Rumula, Mt Lewis range, Oct 1964, Schodde 4144

(BRI, CANB); S.F.R. 310, Cpt. 21, 17° 15% 145°

40’E, Dec 1961, vonAltena 2203 (BRI). North Kennedy

District: Bluewater State Forest, 55 km NW of

Townsville, 19° 13% 146° 25% Nov 1991, Bean

3798 (BISH, BRI, LAE); Mt Fox, Nov 1949, Clemens

s.n. (BRI [AQ716419]); 18 km W of Bruce Highway,

along Forestry road, Bluewater Creek, S.F., Paluma

Range, 19° 14% 146° 25% Nov 1991, Halford Q681

(BISH, BRI, K, L, MEL, QRS).

Reynolds & Forster, Revision of Tarenna and Triflorensia

Fig. 2. Tarenna monticola. A. flowering branch x0.5. B. branch of inflorescence in detail x2. C. flower x4. D.

longitudinal section of flower x4. E. style and stigma x4. F. anther x8. G. calyx tube with bracteoles x8. H.

longitudinal section of ovary showing ovules on placenta xl6. 1. stipule x4. A from Sankowsky s.n. (BRI[AQ565519]);

B-I from Halford Q681 (BRI). Del. W. Smith.

Distribution and habitat : Northeast

Queensland from Windsor Tableland in the

north to the Paluma Range in the south. Plants

grow in rainforests (complex notophyll

vineforest), usually at high altitudes (480-880

m altitude) on substrates derived from basalt

and metamorphics (Map 2).

Notes: Tarenna monticola is readily

recognisable from the other two species

enumerated here, by the small domatia in the

angle between the main lateral veins and the

midrib on the lower surface of its leaves, coarser

nerves, usually acuminate leaf apex, glabrous

inflorescences, short pedicels, and glabrous

corolla. Juvenile foliage is densely hairy, but

this is not so evident in adult foliage. Tarenna

monticola also differs from T. dallachiana by

its narrower calyx lobes, shorter corolla, and

greater number of seeds (14-26 seeds in each

fruit).

Conservation status: Tarenna monticola is

widespread and not threatened. It is present in

Paluma & Lake Eacham National Parks and a

number of Forest Reserves and State Forests.

Etymology: The specific epithet refers to the

usual habitat of this species, from the Latin

words monti (mountain) and cola (dweller).

Tarenna subgenus Pseudoixora (Hook.f.)

S.T.Reynolds comb, et stat. nov.; Webera

sect. Pseudoixora (‘ pseudixora’’) Hook.f.,

FI. Brit. India 3: 104 (1882). Type: not

designated.

Tarenna sect. Pauciovulatae Valeton, in

Lauterbach, Beitr. FI. von Papuasien 13,

Bot. Jahrb. 60: 85 (1924). Type: not

designated.

Notes: This subgenus differs from typical

Tarenna by its solitary ovule or seed in each

cell of the ovary and fruit, and by its large, ±

smooth, subglobose Ixora -like seed which is

dorsally rounded and provided with a thick-

rimmed circular excavation on its ventral side.

However, its inflorescence, subulate bracteoles

on the pedicels, stipules and branchlets are

similar to those of subg. Tarenna. Bakhuizen

van den Brink Jr. (1975: 47) recognised sect.

Pseudoixora under Tarenna Gaertn. but did not

make a formal combination.

Austrobaileya 7 (1): 29-55 (2005)

Distribution: This subgenus is present in

northern Australia ( T. pentamera ), New Guinea

(e.g. T. guelcheriana Valeton and

T. warburgiana Valeton), south-east Asia (e.g.

T. curtisii F.N. Williams), Mauritius in the Indian

Ocean (T. richardii Drake exB.Verdcourt) and

Africa (Bridson 1979).

Etymology: Hooker (1882) was apparently

referring to the Ixora -like seeds when he named

this subgenus (as sect. Pseudoixora

(‘ Pseudixora’).

3. Tarenna pentamera (Benth.) S.T.Reynolds

comb, nov.; Ixora pentamera Benth., FI.

Austral. 3: 416 (1867). Type: Northern

Territory. Cavern [Chasm] Island, off

Groote Eylandt, 14 January 1803, R. Brown

3445 (lecto [here designated]: BM;

isolecto: K, CANB, MEL).

Tarenna sp. A (Koch 1992: 929).

Shrubs or trees 2-8 m tall; bark light brown or

greyish brown; branchlets greyish, ± 4-gonous

towards apex, densely hairy with short

spreading hairs, subglabrous or glabrous.

Leaves petiolate; lamina elliptic, narrowly elliptic

or elliptic-ovate, apex acute or shortly

acuminate, base subacute and decurrent into

the petiole, sometimes suboblique, (5.5-) 8.5-

15 (—18.5) x (2-) 3-5 (-6.3) cm, coriaceous,

drying greenish to darkish-brown, glossy

above; upper surface glabrous or sparsely hairy,

lower surface densely hairy or only the nerves

finely hairy, usually with small hairy tufts in axil

of the main nerves; midrib sunken above, lateral

nerves 8-12 pairs, slightly oblique or ± arcuate,

impressed above; petioles 1.2-3.1 cm long,

slender, channelled above, finely hairy or

glabrous. Stipules free or slightly connate (near

base), ovate or triangular, 5.5-7 x 2.5-3 mm,

keeled, aristate (especially in young plants),

paler at margins, finely and sparsely appressed

hairy on outside, glabrous inside; colleters

present at the base inside. Inflorescences

trichotomously branched corymbiform panicles

4.5-7 cm long; centre branch with (1-) 2 or 3

internodes; lateral branches 2.5-7.5 cm long,

17-22-flowered; bracts ovate, sometimes very

small and only represented by thick subulate

lateral lobes. Flowers perfumed; pedicels 1-4

mm long; bracteoles 1 or 2 on each pedicel,

Reynolds & Forster, Revision of Tarenna and Trijlorensia

alternate, minute, narrowly ovate or subulate.

Calyx cyathiform, dilated to a broad limb, 2-2.5

mm long, densely adpressed hairy outside; tube

c. 1 mm long and c. 1 mm wide at mouth, with

limb about 1 mm long; lobes small, c. 0.5 x 0.7

mm, hemispherical. Corolla white; tube

infundibuliform, 2-3.5 mm long, dilated near

mouth (1.5-2 mm wide at mouth), glabrous

outside, densely hairy at throat; lobes longer

than tube, 4.5-6 x 1.5-2 mm, obovate, obtuse,

reflexed. Stamens exserted, filaments 0.7-1 mm

long, anthers as long as or longer than corolla

lobes, 3.5-5 mm long, usually with a prolonged

acute apical connective; ovules solitary in each

cell of the ovary, attached to the septum. Style

with stigma 7.5-9 mm long, stigma thick,

elongate, ± clavate. Fruits purple black when

ripe, subglobose, 5-6 x 5.5-6 mm, sometimes

ribbed when dry, 2-seeded; seeds solitary in

each locule, subglobose, rounded on top, flat

below and provided with a circular thickly

rimmed hilar cavity in the centre, smooth or finely

and obscurely reticulate on its dorsal side. Fig.

Selected specimens examined : Western Australia.

Fenelon island, 14° 08’S, 125° 42’E, Jul 1977,

Kenneally 6384 (PERTH); Lone Dingo, 14° 03’S, 125°

43’E, Jan 1982, Kenneally 7839 (BRI, QRS, PERTH);

5.9 km E of Mt Waterloo, 15° 19’S, 125° 07’E, Jun

1987, Kenneally 10480 & Hyland (CANB, PERTH);

Mama Falls, 5 km E of Kalumburu on edge of

escarpment, 14° 18’S, 126° 41’E, Dec 1992, Mitchell

2796B (BRI, PERTH). Sir Graham Moore Island, 13°

56’S, 126° 33’E, Jul 1973, Wilson 11250 (PERTH).

Northern Territory. Popham Bay, Cobourg Peninsula,

11° 16’S, 131°50’E, Jun 1983, Bardsley s.n. (DNA

27284); Bickerton Island, 13° 46’S, 131° 50’E, Jun

1985, Clark 112 & Wightman (DNA); N Coombe

Point, Mountnorris Bay, 11° 26’S, 132° 47’E, Oct

1992, Cowie 3228 (BRI, DNA); Castlereagh Bay,

Dhipirrinjura Outstation, 12° 15’S, 135° 05’E, Aug

1995, Cowie 5978 (BRI, DNA); Crocodile Island, NW

of Milingimbi, 12° 00’S, 134° 50’E, Dec 1998, Cowie

8138 & Harwood (BRI, DNA); Wurrmalmirr Creek,

Elcho Island, 11° 58’S, 135° 37’E, Jul 1975, Dunlop

3851 (BRI, CANB, DNA); Gunn Point, 12° 12’S, 131°

01’E, Nov 1989, Forster P1F5926 & Russell-Smith

(BRI, DNA); 6 km S Brogden Point, Murgenella, 11°

34’S, 133° 04’E, Mar 1987, Russell-Smith 1994 &

Lucas (CANB); 5 km south Snake Bay, Melville Island,

11° 28’S, 130° 41’E, Apr 1987, Russell-Smith 2081 &

Lucas (BRI, DNA); Groote Eylandt, Angurugu, 13° 59’S,

136° 28’E, Jul 1987, Russell-Smith 2873 & Lucas

(DNA); NE of Port Bradshaw, Gove, 12° 27’S, 136°

49’E, Sep 1987, Russell-Smith 3371 & Lucas (DNA);

Melville Island, Rangani Creek, 11° 18’S, 130° 31’E,

Jun 1988, Russell-Smith 5757 & Lucas (BRI, DNA);

Conder Point, Melville Island, 11° 44’S, 131° 17’E,

May 1989, Russell-Smith 8067 & Lucas (BRI, DNA);

Gunn Point, 12° 10’S, 131° 05’E, Dec 1989, Russell-

Smith 8175 & Lucas (BRI, DNA); Yirrkala, 12° 12’S,

136° 47’E, Aug 1948, Specht 938 (BRI, MEL); West

of mouth of West Alligator River, 12° 12’S, 132° 15’E,

May 1978, Webb & Tracey 12257 (BRI, DNA, QRS);

NE Coast of Cape Van Diemen, Melville Island, 11°

10’S, 130° 22’E, May 1978, Webb & Tracey 12478

(CANB).

Distribution and habitat : Northern Australia

with a chiefly coastal distribution from the

Mitchell Plateau, Western Australia, to the Gulf

of Carpentaria, Northern Territory and on

offshore islands; usually in deciduous vine

thickets on coastal dunes (Map 2).

Notes : Tarenna pentamera is characterised by

hairy young branchlets and leaves (especially

lower surface of leaves), acute or acuminate leaf

apex, subulate bracteoles on the pedicels,

flowers with fairly short and wide corolla tubes

which are shorter than its lobes, and by its 2-

seeded fruits with a large solitary subglobose

seed in each locule of the fruit. It resembles

T. asiatica in its inflorescence, the shape of the

corolla and subulate bracteoles, but differs by

its solitary ovule or seed in each cell of the ovary

or fruit, attachment of the ovule, and by its

subglobose seed. It differs from the other

Australian species of Tarenna by its short

corollas and large subglobose solitary seed in

each cell of the fruit. In the other species, the

corolla tube is usually very long and much

longer than its lobes, and the seeds are small

and numerous (more than 4) in each cell of the

fruit, angular, and densely conspicuously

reticulate.

Typification: Several sheets of the type

collection (Brown 3445), are available for study,

the sheet at BM which consists of both

flowering and fruiting specimens is here chosen

as lectotype of this species name, the

isolectotype at CANB has flowers only, whereas

the isolectotypes at K and MEL are fruit only.

Two labels are present on the BM sheet, one

bears the following annotation, presumably

made by Brown, “Cavern Island and island 91 (?),

island 3 (?), No 17 spec.”, whereas the other is

typed “Cavern Island and other islands near

Groote Eylandt, 14 January 1803”. In his

manuscript, Brown cited “Carpentaria, small

island adjoining Cavern Island”, date of

collection 14 January 1803. According to Specht

Fig. 3. Tarenna pentamera. A. flowering branch *1.2. B. flower x5. C. longitudinal section of flower showing a

solitary ovule and anthers x4. D. longitudinal section of ovary showing more detail of solitary ovule. x8. E. anther.

x8. F. style and stigma x6. G. seed x5. All from Cowie 3228 (BRI). Del. W.Smith.

Reynolds & Forster, Revision of Tarenna and Triflorensia

(1958:2) Brown visited Chasm Island, off Groote

Eylandt, on that date. Vallance (1990) stated that

between the 5 th and 17 th of January 1803, Brown

visited Chasm and Finch Island and more

recently Vallance etal. (2001) have categorically

stated that “Brown lands on Chasm Island” on

14 January 1803. Hence it seems reasonable to

assume that Brown’s “Cavern” island is now

Chasm island (13° 40’S, 136°35’E).

Conservation status: Tarenna pentamera is

widespread and not threatened. It has not been

collected from any conservation reserves.

Etymology: The specific epithet is from the

Greek pent (five) and merus (parts) and alludes

to the perianth parts.

Triflorensia S.T.Reynolds genus nov. arte affinis

Tarennae Gaertn. a quo inflorescentia

ramis florentibus quibisque trichotomo

ramosis instructa in cymis dichasiali 3-flora

terminentibus vel floribus (praecipue

lateralibus) tribus ramis quibisque in cyma

3-flora plerumque substitutis, bracteis basi

breviter connatis, bracteolis binatis,

floribus plerumque 4-meris, stipulis

obtusis plerumque tenuibus, ramulorum

ramificatione supra-axillare differt. Typus:

Triflorensia australis (Benth.)

S.T.Reynolds (= Diplospora australis

Benth.)

Shrubs or small trees; branchlets pale brown or

greyish with flaky bark (young ones resinous),

trichotomously branched, the central axis

usually with supra-axillary branching (that is

the branchlet is branched (with three branches)

at some distance above the last node and not at

the node itself, and often provided with a long

internode between the upper trichotomous

branches and last node). Leaves entire,

coriaceous, drying darkish; petiolate. Stipules

connate from middle down or connate only

towards the base with lobes broadly ovate,

obtuse or slightly rounded at apex, thin, slightly

scarious at margins, keeled, occasionally

provided with prominent, thick, rigid lateral awns

and with colleters at base inside. Inflorescences

terminal, trichotomously branched with the

three branches either clustered at the apex of

the branchlet or clustered on a short stalk

subtended by the uppermost pair of leaves,

each branch terminated by a triad (dichasial

cyme); all the flowers of the triad or only the

lateral ones replaced by branches each

terminated by a triad; branches often branched

many times, with middle branch of the main

inflorescence sometimes provided with 1 or 2

internodes, and terminated by 3 branches each

with a triad at its apex; lateral branches of the

inflorescence patent, sometimes 3-branched at

their apex, each branch terminated by a triad;

bracts persistent, those subtending the

inflorescence stipule-like, connate, obtuse at

apex and shortly sheathing at base, those at

the upper junction of the trichotomous

branches shortly connate with depressed ovate,

slightly scarious, erose lobes, and usually

provided with long or short, thick, rigid, lateral

awns; colleters present at base inside. Flowers

bisexual, 4 (or 5) -merous; middle flower of the

triad usually sessile, lateral flowers pedicellate;

bracteoles small, paired, inserted on either side

of the calyx tube, ovate, erose at margins. Calyx

turbinate, with limb wider and thinner, 4 or 5-

lobed; lobes imbricate, thin, hemispherical,

erosed at margins and usually ciliolate. Corolla

tube infundibuliform, slightly shorter than the

lobes; hairy or glabrous inside; corolla lobes 4

(or 5), twisted to the left in bud, longer than the

tube, oblong, obtuse at apex, hairy from about

middle to the base inside, glabrous outside.

Stamens 4 (or 5); anthers introrse, narrowly

ovoid-ellipsoid, with a short apical lobe, thicker

towards the middle (near suture of the lobes),

dorsifixed; filaments very short, usually inserted

below apex of the corolla tube; pollen grains 3-

colporate; disc fleshy, annular. Ovules 2 in each

cell of the ovary, attached to the septum; style

slender, as long as corolla tube; stigma shortly

exceeding the corolla tube, fleshy, lanceolate,

with 2 erect lobes at apex. Fruits globose, green

turning black when ripe, smooth; endocarp

crustaceous, thin, shiny, pale brown; seeds two

(very rarely one) per locule, hemispherical-

angular, smooth and rounded on top, slightly

angular on its sides, flat and with a subcircular

or elliptic excavation in the centre on its ventral

side.

Distribution: This genus is currently endemic

to Australia where it is represented by three

species. It is probably also in New Caledonia as

Tarenna microcarpa (Guillaumin) Jeremie

(Jeremie 1974) from that country resembles

Triflorensia australis in its seeds.

Notes : Triflorensia is characterised by its three

inflorescence branches which are apically

clustered at the branchlet tip or on a short stalk;

each branch is terminated by a three-flowered

dichasium, the middle flower of the triad usually

sessile, the lateral ones stalked, the lateral

flowers or all three flowers usually replaced by

3 branches each, these often branching again

and ultimately terminated by a triad; connate

ovate bracts, which are shortly sheathing at

base, and provided with long or short, rigid,

lateral awns; 4 (or 5) -merous flowers; paired,

small bracteoles (one on each side of the calyx);

thick, lanceolate stigma usually with 2 erect lobes

at apex; small, 4 (rarely 3) -seeded fruits; slightly

angular seeds which are rounded on top,

angular on its side, and with a subcircular

excavation on its flat ventral side; obtuse thin

stipules, and supra-axillary branching (that is

the central branch (of the three branches of the

branchlet) usually branched (with 2 lateral

branches) at some distance above the last node

instead of at the node itself; the distance from

last node to where the upper three branches of

the branchlet diverge is between 5-30 mm long).

Triflorensia differs from Tarenna in the 2

ovules or seeds in each cell of the ovary or

fruit; hemispherical seeds which are rounded

on top and angular on their sides; inflorescence

branches terminated by a triad (the middle flower

Austrobaileya 7 (1): 29-55 (2005)

the oldest and usually sessile, the laterals

pedicellate); connate bracts (connate near base

and often provided with lateral awn-like lobes);

stipules thin, usually obtuse; and the supra-

axillary branching of the primary branch of the

branchlet.

Earlier workers such as J.D.Hooker have

indicated that Diplospora australis Benth. was

wrongly placed in the genus Diplospora (he

had determined the type specimen of this

species at K, ‘certainly not Diplospora ’), and

another worker (hand writing appears to be that

of ? W. Francis from BRI) had determined the

same collection as ‘ Stylocoryne’ (now under

Tarenna ). Diplospora australis and the two

other Australian species remained under the

name Diplospora (e.g. Bailey 1900; White 1926;

Ross 1986) until Ali and Robbrecht (1991)

tranferred them to Tarenna Gaertn. and made

the following combinations viz. Tarenna

australis (Benth.) Ali & Robbr., Tarenna

cameroni (C.T.White) Ali & Robbr., and

Tarenna triflora (Benth.) Ali & Robbr. nom.

illeg. (see under Triflorensia ixoroides).

Etymology : The name of the genus alludes to

the three branches of the inflorescences, each

of which are terminated by a triad, the three

flowers of the triad often replaced by 3 branches

each terminated by a triad.

Key to the species of Triflorensia

1. Inflorescences usually small, the three branches terminated by a 3-flowered

dichasium, or the flowers of each triad usually replaced by three stalked

triads, the flowers of these are occasionally replaced by 3 branches each

terminated by a triad, the whole inflorescence ultimately (3-) 7-13(-25)-

flowered, intemodes rarely present in the central branch; middle flower of

the triad sessile, laterals stalked; peduncles and young branchlets usually

with sparse short hairs; leaves 2^1.5 (-6) x 0.6-1.8 (-2.7) cm. 3. T. ixoroides

Inflorescences usually larger, the three flowers of each branch of the

inflorescences usually replaced by three branches each terminated by a

triad, the branching often repeated many times, the main branch (central

branch) of the inflorescence usually with one or two internodes, the whole

inflorescence ultimately (9-) 15-53-flowered; middle flower (of the 3-

flowered cyme) sessile or shortly stalked, laterals stalked; peduncles and

young branchlets glabrous or sparsely hairy; leaves 5-10 (-12.5) x 1.7-

4.2cm.2

2. Inflorescences 27-53-flowered, peduncles glabrous or sparsely hairy; middle

flower of the 3-flowered cymes sessile; lateral flowers stalked (pedicels 1-

Reynolds & Forster, Revision of Tarenna and Triflorensia 45

1.5 mm long); flowers mostly 4-merous; leaves (5-) 7-10 (-12.5) x 1.7-3.5

(-4.2) cm; petioles 5-12 mm long; fruits globose, 6-10 x 7-10 mm. 1. T. australis

Inflorescences 9-27-flowered, peduncles glabrous; all 3 flowers of the cyme

usually stalked; pedicels 2.5-3 mm long, shorter in the middle flowers;

flowers mostly 5-merous; leaves 5.5-7.2 x (1.8-) 3.2-4.2 cm; petioles 10-

15 mm long; fruits ellipsoid or ellipsoid ovoid, 8-9.5 x 6.5-7 mm. 2. T. cameronii

l.Triflorensia australis (Benth.) S.T.Reynolds

comb.nov.; Diplospora australis Benth.,

FI. Austral. 3: 413 (1867); Tarenna

australis (Benth.) Ali & Robbr., Blumea

35: 302 (1991). Type: Queensland, Cape

York, W. Hill39 (lecto [here designated]:

K).

Illustrations : Hyland etal. (1999,2003).

Small trees or shrubs 2-6 m high; branchlets

pale brown or pale greyish brown; young

branchlets and inflorescences sparsely hairy or

glabrous. Leaves petiolate; lamina elliptic,

subacute or obtuse at apex, subacute and

attenuate into petiole at base, 5-10.2 (-12.5) x

1.7-3.5 (-4.2) cm, coriaceous, thickened at

margins; midrib raised and deeply chanelled

above; lateral nerves 5 or 6 (-8) pairs,

suboblique or subpatent, looping at margins;

distinct both surfaces; reticulate veins not

apparent; domatia absent; petioles 5-12 mm

long; stipules shortly sheathing near its base

and provided with 2 lateral thick rigid awns,

lobes usually thin, broadly ovate, rounded or

obtuse, obscurely keeled. Inflorescences 1.5-

3.5 x 1.7-2.5 cm, peduncles glabrous or

subglabrous, the 3 branches usually clustered

on short stout peduncles (this 3-6 mm long (10

mm long in fruit)), each branch 3-13 mm long,

terminated by a 3-flowered dichasial cyme (the

middle flower of the dichasium sessile, laterals

stalked), all the flowers or only the lateral flowers

of the triad replaced by 3-branches each

terminated by a triad, the three flowers of each

are again replaced by three branches each with

a triad, this is repeated many times, the whole

inflorescence ultimately 27-53-flowered; the

central branch of each inflorescence with 1 or 2

intemodes, these 5-13 mm long; lateral branches

without internodes, with the lowermost branch

7-18-flowered; stalk of ultimate cymes 1-2 mm

long; bract at the base of the three main

branches of the inflorescence, stipular, slightly

sheathing and sometimes provided with lateral

rigid awns, whereas bracts on upper branches

small, and usually without awns. Flowers 4-

merous; central flower sessile; lateral flowers

stalked (pedicels 1-1.5 mm long); bracteoles

ovate, obtuse; calyx 1.5-2 x 1.5-2 mm. Calyx

lobes hemispherical, c. 0.5 x 1 mm, erose at

margins and ciliolate. Corolla infundibuliform,

5-6 mm long, white; tube 2-2.5 x 2.5 mm, hairy

inside, with the hairs denser towards the mouth;

lobes subpatent, 3.5-4.5 mm long, oblong,

obtuse, hairy from about middle down. Anthers

spreading, 3-4 mm long, dorsifixed towards the

base; filaments c. 0.5 mm long; disc fleshy

shorter than calyx limb. Style and stigma 6-7

mm long, style sparsely hairy; stigma thick, as

long as the anthers, slightly exceeding the

corolla tube. Fruits globose, 6—10 x 7-10 mm;

stipe 4—6 mm long. Fig. 5.

Selected specimens examined : Northern Territory

10 miles (16 km) NE Mudginberry Homestead, 12°

36’S, 132° 52’E, Jul 1972, Byrnes 2736 (CANB, DNA);

W side of Gunn Point, Nov 1990, Cowie 1417 &

Dunlop (BRI, DNA, NSW, QRS); East Alligator River,

10 km S of Cannon Hill Ranger Station, 12° 27’S, 132°

58’E, May 1980, Craven 6004 (CANB, DNA); Gunn

Point, 12° 12’S, 131° 00’E, Apr 1984, Dunlop 6676

(DNA); Wessel Islands, 11° 10’S, 136° 44’E, Sep 1974,

Latz 3311 (BRI, DNA); Marchinbar Island, Wessels,

11° 23’S, 136° 35’E, Dec 1987, Russell-Smith 4476 &

Lucas (BRI, DNA, QRS); Kakadu N.P, 4 km W of Hill

301, 12° 31’S, 132° 57’E, Jun 1988, Russell-Smith

5682 & Lucas (BRI, DNA); 19 km E Jabiru, 12° 37’S,

133° 03’E, Apr 1989, Russell-Smith 8053 (BRI, DNA);

12 km E of Mudginberri Homstead, Kakadu N.P., 12°

35’S, 132° 59’E, Jan 1991, Russell-Smith 8401 &

Brock (BRI, DNA); Kakadu National Park, Ngarradj,

12° 29’S, 132° 55’E, Jan 1992, Russell-Smith 8569 &

Lucas (BRI). Queensland. Cook District: Mapoon,

Jul 1901, Bailey s.n. (BRI [AQ124224]); Coast, Mt

Bremer, 26 km NE of Bamaga, 10° 42’S, 142° 31’E,

Feb 1994, Fell DGF3932 et al. (BRI); 102 km NW of

Coen, boundary of Archer Bend N.P. and Merluna

Holding, 13° 20’S, 142° 29’E, Jun 1994, Fell DGF4420

& Buck (BRI, DNA, QRS); 28 km along road to Leo

Creek Mine, Mcllwraith Range, 13° 42’S, 143° 17’E,

Jun 1992, Forster PIF10047 & Tucker (BRI, MEL,

QRS); Mary Valley Scrub, 15° 02’S, 143° 45’E, Jul

1993, Forster PIF13443 et al. (BRI); Bolt Head,

Temple Bay, 12° 15’S, 143° 05’E, Jun 1996, Forster

PIF19392 (BRI); Kroll Creek Holding - Parish of

Kesteven, 13° 43’S, 143° 14’E, Mar 1982, Hyland

11770 (BRI, QRS); T.R. 14, Parish of Kesteven, 13°

43’S, 143° 18’E, Apr 1993, Hyland 14744 (BRI, QRS);

Lankelly Creek road, Mcllwraith Range, 13° 53’S, 143°

15’E, May 1995, Hyland 15341 (BRI, QRS); Dauan

Island, Torres Strait, 9° 25’S, 142° 32’E, Sep 1971,

Lawrie s.n. (BRI [AQ3998]); Cape York, Oct 1849,

Bot 495, McGillivray s.n. (K - residual syntype).

Distribution and habitat: Triflorensia australis

is endemic to tropical, northern Australia where

it has been collected from Arnhem Land in the

Northern Territory and Cape York Peninsula and

Torres Strait in Queensland. Populations are

largely coastal and occur in vinethickets, often

in the lee side of foredunes or coastal cliffs.

Substrates are usually derived from sandstone

or laterite (Map 3).

Typification: A lectotype is selected for the

name based on the better of the two original

syntypes at K.

Notes: Triflorensia australis is readily

distinguishable from the other species by its

larger leaves, and larger, usually branched

inflorescences. It is also markedly allopatric in

its distribution.

There is considerable variation in the

leaves of the specimens available for study.

Generally they are elliptic, and usually medium¬

sized in the majority of specimens seen (e.g.

Dunlop 6676, Cowie 1417 & Dunlop; Hyland

5502 and 10857 resemble the syntypes). A few

specimens have very small or larger leaves but

are probably the result of phenotypic variation,

rather than representing distinct forms, as they

are connected by intermediates throughout

their range. Specimens with small or narrow

leaves (viz. Byrnes 2736 and Tracey 14645 )

resemble T. ixoroides in their leaves, but that

species differs by its very small inflorescences,

whereas specimens with slightly larger leaves

(viz Russell-Smith 5733 & Lucas', Russell-Smith

8569 & Lucas), appear to have a slightly

different aspect in their branching and leaves

(with broader bases) and may be worthy of

formal recognition.

Conservation status: This species is not

considered threatened. It is present in Kakadu

National Park in the Northern Territory and

Archer Bend National Park in Queensland.

Etymology: The specific epithet is from the

Latin word australis (southern) and pertains to

the perceived unusual distribution at the time

of description.

Austrobaileya 7 (1): 29-55 (2005)

2. Triflorensia cameronii (C.T.White)

S.T.Reynolds comb, nov.; Diplospora

cameronii C.T.White,/. Bot. 64:216-217

(1926). Type: Queensland. Moreton

District: Yarraman, October 1924, M.A.

Cameron Y63 (lecto [here designated]:

BRI [AQ717032]).

Tarenna cameronii (C.T.White) Ali & Robbr.,

Blumea 35: 302(1991).

Illustrations: Ali & Robbrecht (1991: 282);

James (1992: 504); Hauser & Blok (1998:

296).

Small trees or shrubs to 4.5 m high; branchlets

pale brown with thin flaky bark; young

branchlets and inflorescence axes slightly

viscid, glabrous. Leaves petiolate; lamina

lanceolate, elliptic or broadly elliptic, obtuse or

abruptly shortly acuminate at apex, obtuse or

subacute and decurrent into the petiole at the

base, 5.5-7.2 (-11.5) x 2-4.2 (-6.5) cm,

coriaceous, drying dark brown, very glossy

above, dull and pale brown below; midrib raised

above, deeply channelled; lateral nerves 6-8 (-

10) pairs, subpatent, looping at margins;

reticulate venation not apparent; petioles 10-

15 mm long. Stipules shortly connate, with lobes

obtuse, thin and colleters present inside.

Inflorescences c. 2.2 x 2.5 cm, with the 3

branches of the inflorescence clustered on a

long or short main stalk (stalk 2-7 mm long), or

clustered at apex of the branchlets, each branch

5-8 (-12) mm long, terminated by a triad; the 3

flowers often replaced by 3 branches each

terminated by a triad, with the whole

inflorescence to 27-flowered; lateral branches

(3-) 7-15-flowered; central branch of the main

inflorescence with 1 or 2 internodes, 5-10 mm

long and 3-branched at apex; bracts shortly

connate and provided with thick lateral awns,

lobes broadly ovate, obtuse, (bracts at the base

of the main 3-branched inflorescence

occasionally with foliaceous lateral lobes,

whereas those subtending the branches have

narrow, scarious lobes). Flowers mostly 5-

merous, middle flower of the cyme sessile or on

short stalks (pedicels to 4 mm long); lateral

flowers stalked (pedicels 2.5-4 mm long);

bracteoles ovate, erose, c. 1 x 1.5 mm. Calyx2-

3.5 x 2.5-5 mm; limb thinner, 5-lobed; lobes

Reynolds & Forster, Revision of Tarenna and Triflorensia

hemispherical, c. 1 x 1.7 mm. Corolla

infundibuliform, 6-7 mm long, white; tube 2-

2.5 x 3.5 mm long, glabrous inside; lobes

oblong, obtuse, c. 4x3 mm, densely erect hairy,

especially towards their base (near mouth of

the tube), glabrous on the margins. Anthers

narrowly ovoid, apiculate, c. 4 mm long;

filaments c. 0.5 mm long. Style with stigma c. 7

mm long; style glabrous; stigma fusiform,

slightly 2-lobed at apex. Fruits ellipsoid or

ellipsoid-ovoid, 8-9.5 x 6.5-7 mm. Fig. 4.

Selected specimens examined : Queensland. Moreton

District: Yarraman, Dec 1924, Cameron Y63 (BRI -

residual syntype); Yarraman L.A., Compartment 7,

Oct 1955, District Forester s.n. [Cameron ] (BRI

[AQ10267]); Yarraman, [S.F.]R. 289, Oct 1955, District

Forester s.n. [Cameron] (BRI [AQ411508]); loc. cit. ,

Jan 1956, District Forester s.n. [Cameron] (BRI

[AQ10266]); Russel’s road, near Pine Mountain, 27°

32’S, 152° 43’E, Feb 1983, Forster PIF1528 et al.

(BRI); 1.5 km SW of Mt Berryman, 27° 44’S, 152°

19’E, Feb 1991, Forster PIF7769 & Sharpe (BRI,

QRS); Anduramba road, Biarra Range between Strange

View and Barnes Hill, 27° ll’S, 152° 06’E, May 1990,

Grimshaw PG2 (BRI); Road to Worlds End Pocket, N

of Ipswich (1.5 km SE of Pine Mountain), Dec 1981,

Guymer 1628 (BRI); Sandy Creek, NW of Joalah N.P,

Tamborine, 27° 48’S, 152° 56’E, Dec 1987, Mertons

s.n. (BRI [AQ408219]). Darling Downs District: Bunya

Mountains N.P., Cherry Plain to Mt Kiangarow, 26°

40’S, 151° 32’E, Jul 2001, Forster PIF27478 & Booth

(BRI, MEL, NSW); Cawdor, near Toowoomba, Jan

1976, Gilbert s.n. (BRI [AQ114754]); Between Gowrie

Junction and Highfields, McKenzie s.n. (BRI

[AQ397008]); near Rogers Park, S.F. 289, Cooyar,

Feb 1991, Rider s.n. (BRI [AQ503777]); Bunya

Mountains, Dec 1883, Scortechinii s.n. (MEL

1583913). New South Wales. Lismore, Oct 1894,

collector unknown (NSW 193805).

Distribution and habitat : Triflorensia

cameronii is endemic to south-eastern

Queensland and northern New South Wales

with a northern limit at Yarraman and a southern

limit near Lismore. Plants occur in dry rainforest

(araucarian microphyll vineforest, semi¬

evergreen vinethicket) on volcanic soils derived

from basalt (Map 3).

Typification : White (1926) clearly indicated that

the type citation covered two specimens with

the same number given for collections made on

different days, viz. “flowering specimens Oct.

1924, fruiting specimens Dec. 1924”. The

flowering specimen is the better of the two

specimens and is selected here as lectotype for

the name.

Notes : Triflorensia cameronii is readily

recognisable from the other two species by its

larger ellipsoid fruits, leaves that are elliptic or

lanceolate and the inflorescences that are much

branched and glabrous.

Conservation status : Triflorensia cameronii is

not considered threatened in Queensland where

it is present in Bunya Mountains and Joalah

National Parks and a number of State Forests.

In New South Wales this species is listed as

Endangered (NSW Scientific Committee 2004).

Etymology : Named for McRae (Mac) Archibald

Cameron (1897-1994), member of the 5 th

Australian Light Horse in World War 1,

subsequently District Forester at Yarraman and

collector of the type.

3. Triflorensia ixoroides (F.Muell.)

S.T.Reynolds comb, nov.; Diplospora

ixoroides F.Muell., Fragm. 9: 182 (1875).

Type: Rockhampton, s. dat ., Dali achy s. n.

(lecto: K; isolecto: MEL) fide McGillivray

(1972) [as Ixora triflora R. Br. ex Benth.

nom illeg. ]

[Ixora triflora auct. non (Forst.) Seem.;

Benth., FI. Austral. 3:416 (1867). pro parte

(quoad specimen “East Coast, Brown 5,1

(pro parte [specimens at K and MEL only]);

Rockhampton, Thozet 2 (MEL);

Rockhampton, Dallachy 1 2 (K, MEL); Broad

Sound, Bowman 3 ( n.v .)]

1 This collection (East Coast, R. Brown ) is a mixed collection of two species. The fruiting specimens (at BM,

CANB) that are labelled Thirsty Sound and annotated (probably by Brown) as ‘ Ixora triflora ’ or ‘ Pavetta triflora ’

(BM specimen) are Ixora queenslandica Fosberg, whereas the flowering specimens (at K, MEL) that are annotated

by Brown as ‘ Canthoides ’ (BM specimen) are Triflorensia ixoroides.

2 The flowering specimens {viz. Rockhampton leg. Dallachy (MEL), Rockhampton leg. Thozet (MEL), and

probably Broad Sound, leg Bowman (see 3 below) match Bentham’s description of the inflorescence and flowers

which are that of Diplospora ixoroides F.Muell.

3 This collection is probably referrable to Triflorensia ixoroides as it has been recorded from this locality.

Fig. 4. Triflorensia cameronii. A. habit (flowering) *0.5. B. detail of inflorescence showing bracts x2. C. detail of

dichasial cyme x3. D. stipule x3. E. calyx with bracteoles x8. F. flower x4. G. petal with anther x6. H. two faces of

anther xl2. I. style with stigma x6. J. fruit x3. K. dorsal and ventral view of seed x6. All from District Forester (BRI

[AQ10266]). Del. W.Smith.

Reynolds & Forster, Revision of Tarenna and Triflorensia

Tarenna triflora (R.Br ex Benth.) Ali & Robbr.,

Blumea 35: 302 (1991) nom. illeg. (based

on Ixora triflora R. Br. ex Benth. nom.

illeg.)

Tarenna sp. (Ka Ka Mundi NP W.J. McDonald

+ 4642) (Reynolds 1997: 184; Forster &

Halford 2002:177)

Illustrations : Jones (1986: 88); Ross (1986:342,

fig. 47M); Ali & Robbrecht (1991:282).

Shrubs or small trees 2.5-6 m high; bark smooth

or slightly rough, pale grey or grey brown;

branchlets pale brown or grey, slightly 4-ridged

towards their apex and sparsely minutely hairy;

nodes dilated; supra-axillary branching frequent

(that is the main axis of the branchlet is

trichotomously branched at some distance

above the last node, with the internode

(between the last node and the upper

trichotomous branch) usually very

conspicuous. Leaves petiolate; lamina elliptic,

subacute at both ends, 2-4.5 (-6) x 0.6-1.8 (-

2.7) cm; margins thickened, coriaceous, dark

green and very shiny on upper surface, pale

green and dull below; midrib raised above and

deeply channelled; lateral nerves 6 or 7 pairs,

subpatent, looping at margins, not apparent on

lower surface; petioles 2^1 mm long. Stipules

connate towards the base, with lobes broadly

ovate, obtuse; colleters present inside,

prominent. Inflorescences resinous when very

young, usually very small, 0.4-1 x c. 1.2 cm,

with 3 branches clustered on a short stalk (this

2-7 mm long), or clustered at apex of the

uppermost pair of leaves; peduncles finely

hairy; each of the three branches of the

inflorescence terminated by a triad; all three

flowers or the lateral ones only are replaced by

three branches each (these branches are 2-7

mm long) and terminated by a triad; flowers of

each triad occasionally replaced by 3 branches

each bearing a sessile flower in the centre and 2

stalked lateral branches; whole inflorescence

ultimately 7-13 (-21 )-flowered; central branch

of the inflorescence rarely with an internode

(this 2-3 mm long); bracts connate towards

base, with lobes very broadly ovate, depressed

on top with fimbriate margins, finely hairy at

base; bracts at base of the 3-branched

inflorescence (or on the top of the main

peduncle) usually with thick and rigid, dark

lateral awns (those subtending the upper

branches are awnless). Flowers (4 or) 5-merous;

middle flower sessile; lateral ones shortly

stalked (pedicels 0.7-1 mm long); bracteoles

ovate, erase; calyx c. 1.7x2 mm, with limb wider;

lobes paler than tube, c. 0.5 x l mm. Corolla

infundibuliform, 5.5-6 mm long, white; tube 2-

2.5 mm long, c. 3 mm wide at mouth, sparsely

hairy inside with long projecting hairs, glabrous

outside; lobes 2.5-3.5 x 2.5-3 mm, oblong with

rounded apex, recurved, densely hairy inside

(from about middle down), glabrous and paler

at margins. Anthers c. 2 mm long, ovoid,

apiculate, darker along the middle (on its dorsal

side); filaments c. 0.5 mm long; disc fleshy,

annular. Style and stigma slightly protruding

from the corolla tube, as long as or shorter than

the anthers, style finely patent hairy; stigma

fleshy, with 2 erect lobes. Fruits subglobose,

5.5-7 x 5.5-9 mm, green turning black; seeds

loose in the mature fruit, subangular, minutely

obscurely pitted or reticulate. Fig. 5.

Selected specimens examined : Queensland. North

Kennedy District: Mt Stuart, near Townsville, 19° 21’S

146° 47’E, Feb 1996, Camming 15819 (BRI). South

Kennedy District: Hazlewood Gorge, 13 km SSW of

Eungella, 21° 15’S, 148° 27°E, Dec 1992, Bean 5267

(AD,BRI). Leichhardt District: Rosewood forest,

Taunton National Park, 23° 31’S, 149° 13’E, Oct 1995,

Brushe 420 et at. (BRI); Connors River, Lake Plattaway,

22° 25’S 148° 55’E, Sep 1993, Fensham 1211 (BRI);

10 km NW of Taroom, 25° 33’S, 149° 38’E, Jan 1996,

Fensham 2864 (BRI); Anvil Peak, Hodgson Range,

22° 41’S, 148° 01’E, Aug 1990, Forster PIF7225 (BRI,

MEL, QRS); Pine Mt, S.F. 79, 21° 44’S, 148° 50’E,

Apr 1991, Forster PIF8014 & McDonald (BRI, QRS);

Expedition N.P., Amphitheatre section, Cannondale

Scrub, 25° 12’S, 148° 59’E, Nov 1998, Forster

PIF23871 & Booth (BRI, MEL, QRS); Palmgrove N.P,

NW of Taroom, 25° 01’S, 149° 15’E, Nov 1998,

Forster PIF23811 & Booth (BRI, MEL, QRS);

Carnarvon N.P., northern boundary of Ka Ka Mundi

section, 24° 41’S, 147° 38’E, Aug 1991, McDonald

4642 & Bean (BRI, CANB, NSW). Port Curtis District:

“Greenfields”, c. 14 km NE of Goovigen, 24° 04’S,

150° 24’E, Mar 1986, Anderson 4113 (BRI); Western

slopes of Mt McGuire, W of Gladstone, 24° 01’S, 151°

02’E, Nov 1996, Aspland 1580 (BRI); Yeppoon Hill,

Oct 1987, Batianoff 9254 & Dillewaard (BRI);

Rockhampton, Jan 1865, Dietrich 2301 (MEL); S.F.

583 Wietalaba, 24° 17’S, 151° 13’E, Dec 1995, Forster

PIF18268 et at (BRI). Burnett District: Auburn River

N.P, 25° 43’S, 151° 03’E, Mar 1988, Forster PIF3583

& Ross (BRI, MEL); 4.5 km S of Binjour, 25° 34’S,

151° 27’E, Forster PIF5873 et at (BRI, MEL). Wide

Bay District: 1 km S of Booyal, 25° 13’S, 152° 02’E,

Nov 1987, Forster PIF3283 (BRI); Ashfield road,

Bundaberg, 24° 52’S, 152° 24’E, Jan 1997, Forster

PIF20184 (BRI); 5 km SSW of Bauple, Dec 1982,

Fig. 5. Triflorensia ixoroides. A. flowering branch xl. B. detail of inflorescence *4. C. flower x4. D. longitudinal

section of flower x4. E. part of branchlet showing supra-axillary branching and stipule x2. Triflorensia australis.

F. flowering branch showing stipule and bract x2. G. detail of dichasial cyme x4. H. calyx with bracteoles xi6. A-

D from McDonald 4642 (BRI); E from Bean 5267 (BRI); F-H from Russell-Smith 4516 (BRI). Del. W. Smith.

Reynolds & Forster, Revision of Tarenna and Triflorensia

Guymer 1835 & Dillewaard (BRI); S.F.38, 39 km WSW

of Maryborough, 25° 38’S, 152° 19’E, Oct 1997,

Halford Q3435 & Hucks (BRI). 148° 27’E, Dec 1992,

Bean 5267 (AD, BRI).

Distribution and habitat: Triflorensia

ixoroides has a wide distribution in eastern

Queensland from near Townsville to Childers,

usually on ranges and slopes, in dry rainforests

(araucarian microphyll vineforest, semi¬

evergreen vinethicket), mostly on stony soil

derived from volcanic substrates (basalt,

granite) (Map 3).

Notes: Triflorensia ixoroides is readily

distinguishable from the other two species by

its small leaves and small inflorescences which

are not as branched. The leaves are variable in

the specimens seen in this study, they are

usually small (between 3.5—4 5 x 0.8-2.4 cm) and

narrowly elliptic in the majority of specimens

(typical of this species), whereas a few

specimens have either very small and usually

very narrow leaves 2-3 x 0.6 cm (viz. Fensham

2864 and Beasley 102), or largish leaves, 5-6 x

2-2.7 cm (viz. Guymer 1835 & Dillewaard), or

occasionally both typical small leaves and

largish leaves on the same branchlet (viz.

Johnson 3512 & Batianoff).

Specimens with larger leaves may be

confused with T. australis when sterile, but that

species has a larger inflorescence and is only

recorded from northern Australia.

Typification: The typification and application

of the names Diplospora ixoroides F.Muell. and

Ixora biflora R.Br. ex Benth. is complicated and

confusing due to two species being used as

syntypes for the latter name and to formulate

the species description (Bentham 1867) as

noted by White (1926), Moore (1926) and

McGillivray (1972).

Bentham’s (1867) description of the fruit

and seed is that of an Ixora whereas the

description of the flowers is not that of an Ixora

and is referrable to Diplospora ixoroides

F.Muell. His syntypes except for Brown’s

collection from “East Coast” 1 (which is a mixed

collection) are that of the latter species. Moore

(1926) after a query from C.T. White (BRI) noted

the presence of two species in the collections

labelled as I. triflora R.Br. ex Benth. at K and

that Bentham’s description of the flowers was

different to that of Brown’s manuscript

description of this species. Brown had based

his manuscript description of the flowers of

Ixora triflora from a Solander manuscript that

is based on an Ixora flower in the Banks &

Solander collection 4 . Bentham obviously did

not see this manuscript. Moore (1926)

lectotypified I. triflora R.Br. ex Benth. by

selecting the BM sheet of Brown’s collection

from East Coast ( viz Thirsty Sound) as

lectotype. However, Fosberg (1938) noted the

name I triflora R. Br. ex Benth. was already an

invalid name when published, being predated

by I triflora (Forst.) Seem, from Fiji which was

published in 1866. He provided a new name

Ixora queenslandica Fosberg for this taxon

which is recognised as an extant species in that

genus (Forster & Halford 2002). The name

Tarenna triflora (Benth.) Ali & Robbr. (1991)

based on Ixora triflora R.Br. ex Benth. nom.

illeg. is therefore a superfluous and illegitimate

name.

Diplospora ixoroides F.Muell. was

described in 1875 because Mueller considered

Bentham’s description of I. triflora R.Br. ex

Benth. inappropriate. He combined I. triflora

under D. ixoroides but did not cite any of

Bentham’s syntypes, nor select a lectotype. His

description and the two specimens he cited

under this species name viz. Port Denison,

Fitzalan, and Neerkoll Creek, Bowman, match

the syntypes of I. triflora from Rockhampton,

Thozet (MEL); Rockhampton, Dallachy (MEL),

and the flowering specimens of Brown’s

collection from the East Coast (specimens at K,

MEL), and also probably with Broad Sound,

leg. Bowman (see above).

McGillivray (1972) lectotypified

Diplospora ixoroides F.Muell. (as Ixora triflora

R.Br. ex Benth. nom. illeg.) with one of

Bentham’s syntypes (of I. triflora R. Br. ex

4 Banks and Solander’s collection from the Bay of Islets (BRI) is a flowering specimen of Ixora queenslandica

Fosberg. The illustration provided by Miller, in Britten, lllustr. Cooks Voy. 2: p. 145, tab. 141 (May 1901) based on

this collection is this species and not Diplospora ixoroides as captioned.

Benth.) viz. Rockhampton, Dallachy (lecto K,

isolecto MEL). This specimen, however, is

referrable to D. ixoroides (see above), so he

had therefore actually lectotypified D. ixoroides

F.Muell.

Conservation status : Triflorensia ixoroides is

widespread and not threatened. It is present in

Auburn River, Carnarvon, Expedition,

Palmgrove and Taunton National Parks, as well

as numerous State Forests.

Etymology : The specific epithet undoubtedly

alludes to the superficial appearance of this

species to plants of the unrelated Rubiaceous

genus Ixora.

Acknowledgements

STR is grateful to Les Pedley for the Latin

diagnosis, Rod Henderson for his advice on

nomenclature, Gordon Guymer for checking and

photographing the types during his term as

ABLO at Kew, Diane Bridson for discussions

on Tarenna in Australia, Will Smith for the

illustrations and maps, the Australian Biological

Resources Study for part funding of this

revision, and the curators of the AD, BM,

CANB, DNA, K, L, MEL, NSW, PERTH and QRS

for the loan of specimens including types.

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Austrobaileya 7 (1): 29-55 (2005)

Map 1. Distribution in 1° grids in Australia for Tarenna dallachiana subsp. dallachiana X and

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and Triflorensia cameronii%.

Ataxonomic revision of Actephila Blume

(Euphorbiaceae/Phyllanthaceae) in Australia

Paul I. Forster

Summary

Forster, P.I. (2005). A taxonomic revision of Actephila Blume (Euphorbiaceae/Phyllanthaceae)

in Australia. Austrobaileya 7(1): 57-98. The genus Actephila Blume is revised for Australia.

Fourteen native and endemic species (one with two subspecies) are recognised: A. bella P.I.Forst.

sp. nov., A. championiae P.I.Forst. sp. nov., A. flavescens P.I.Forst. sp. nov., A. foetida Domin,

A. grandifolia (Mull.Arg.) Baill., A. latifolia Benth., A. lindleyi (Steud.) Airy Shaw, A.

longipedicellata P.I.Forst., A. petiolaris Benth subsp. petiolaris, A. petiolaris subsp. jagonis

P.I.Forst. subsp. nov., A. plicata P.I.Forst. sp. nov., A. sessilifolia Benth., A. traceyi P.I.Forst. sp.

nov., A. venusta P.I.Forst. sp. nov., and A. vernicosa P.I.Forst. sp. nov. An identification key to

species is provided. All taxa are described and most species illustrated. Notes are provided on

distribution, habitat, typification, affinities and conservation status for each taxon.

Key Words: Euphorbiaceae, Phyllanthaceae, Australian flora, Actephila bella, Actephila flavescens,

Actephila grandifolia, Actephila longipedicellata, Actephila petiolaris subsp. jagonis, Actephila

plicata , Actephila traceyi, Actephila venusta, Actephila vernicosa

P.I.Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens,

Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: paul.forster@epa.qld.gov.au

Introduction

The genus Actephila was described by Blume

(1825) who did not include any species at the

time. The first species to be included in

Actephila were A. bantamensis Miq. (now a

synonym of A. javanica Miq.) and A. javanica

(Miquel 1859), although other species were

described prior to this date in other genera and

are now included in Actephila (e.g. Dalzell

(1851) with Anomospermum excelsum (now

Actephila excelsa (Dalzell) Mull.Arg.) and

Steudel (1841) with Lithoxylon lindleyi (now

A. lindleyi (Steud.) Airy Shaw). Numerous

species were described between 1865 and 1978

and there are currently around 20 species of

Actephila recognised worldwide from Australia,

China, India, Indonesia, Malaysia, New Guinea,

India, Laos, Sri Lanka, Solomon Islands,

Thailand and Vietnam (Airy Shaw 1980a, 1981;

Philcox 1997; Govaerts etal. 2000; Chayamarit

in press; Li & Hoffmann in press). Most authors

consider that the species level systematics of

the genus is difficult (Govaerts et al. 2000;

Radcliffe-Smith 2001) and it is likely that the

number of species in the genus will increase

once the Malesian species are revised.

Accepted for publication 13 July 2005

Although the traditional taxonomic

placement of Actephila in Euphorbiaceae

subfamily Phyllanthoideae (or family

Phyllanthaceae) has not been challenged

recently, it remains to be rigorously tested using

molecular data and additional information from

the areas of anatomy (particularly seeds),

chemistry and embryology. An initial

classification of Euphorbiaceae s.l. by Muller

(1866) placed the genus together with Amanoa

Aubl., Cluytiandra Mull.Arg., Cyathogyne

Mull.Arg., Discocarpus Klotzsch, Lachnostylis

Turcz., Moacurra Roxb., Payeria Baill.,

Pentabrachion Miill.Arg., Savia Willd. and

Secretania Miill.Arg., in the tribe Phyllantheae

Mull.Arg. subtribe Savieae Mull.Arg. Pax

& Hoffman (1922, 1931) included Actephila

in the subfamily Phyllanthoideae Ascherson,

subtribe Amanoinae Pax & K.Hoffm., together

with Amanoa. Webster (1994) included the

genus in the subfamily Phyllanthoideae , tribe

Wielandieae Baill. ex Hums., together with the

genera Blotia Leandri, Chonocentrum Pierre

ex Pax & K.Hoffm Discocarpus, Gonatogyne

Mull.Arg .,Heywoodia Sim, Lachnostylis, Savia

and Wielandia Baill., but made no definitive

statement on the closeness of any of these

genera to Actephila. A similar disposition

of genera within this tribe was outlined by

Radcliffe-Smith (2001), with the addition of the

genera Chonocentrum Pierre ex Pax & K.Hoffm.

and Petalodiscus (Baill.) Pax. In both of these

recent, morphologically based overviews,

Actephila was keyed out next to Blotia , a genus

from Madagascar. Most recently it has been

proposed that the subfamily Phyllanthoideae

be again recognised at family level as the

Phyllanthaceae (Chase et al. 2001; APG 2003).

Limited molecular analysis of genera placed in

the Phyllanthaceae has revealed that Actephila

is allied to the genera Andrachne L., Leptopus

Decne., Meineckia Baill., P or anther a Rudge

and Zimmermannia Pax, with the closest

relationship to Andrachne and Leptopus

(Wurdack etal. 2004; Samuel etal. 2005).

As with many genera of Euphorbiaceae

s.l., Actephila remains poorly studied in terms

of its anatomy, chemistry or micromorphology.

The available taxonomic classifications of the

genus are based mainly on gross morphology

and it is likely that further molecular analysis

will reveal patterns of relationship to other

genera in Phyllanthaceae that are different again

to those recently proposed. On the basis of

wood anatomy of a single species of Actephila

( A.javanica ), Mennega (1987) considered that

the genus was close to Pentabrachium. Levin

(1986a-c) studied leaf architecture and epidermal

morphology of two species of Actephila and

concluded that they were most similar to those

from the genus Andrachne L. Tokuoka & Tobe

(2001) described the seed coat anatomy of two

species of Actephila from Asia as exotegmic

and the cells of the exotegmen as stellate-

undulate. A similar seed coat anatomy was

found in two species of Savia and led these

authors to speculate that the two genera may

be closely related. They further stated that the

“Wielandieae are morphologically unlikely to

be monophyletic”. The lack of endosperm in

the seeds of Actephila, Amanoa, Spondianthus

and Wielandia was considered primitive within

the group (Tokuoka & Tobe 1995).

Species Delimitation

The morphological or phenetic species concept

used, has been implicit in previous revisions of

Australian Euphorbiaceae s.l. (Forster 1994a-d,

1995a,b, 1996a,b, 1997a,b, 1999a,b, 2003). This

species concept is closely tied to habitat

Austrobaileya 7 (1): 57-98 (2005)

preferences and geographic distribution and is

arrived at from extensive fieldwork in northern

Australia, particularly Queensland where all of

the Australian species are found.

Species are defined as groups of

populations (1 -many) with discontinuities in the

variation of two or more independent character

states of morphology. It is assumed that there

is genetic continuity or at least a shared ancestral

lineage between the different populations of a

single species. Although species definitions

remain a matter of opinion and debate, the one

applied here (‘phenetic species concept’) is

widely used and understood (Stebbins 1950;

van Steenis 1957; Cronquist 1988; Stuessy 1990;

Levin 1979,2000) and would also equate to the

‘diagnostic species concept’ of Judd et al.

( 2002 ).

If only a single, or several minor

(indumentum colour or density) character state

differences are present and the discontinuity is

geographically based, the rank of subspecies

is used. It is generally considered that

subspecies should differ in only a few minor

characters and that intermediate populations

should exist to demonstrate continuity of

character states (e.g. Stebbins (1950) states

“subspecies.... connected .... by a series of

intergrading forms” or Stace (1989) states “a

population of several biotypes forming a more

or less distinct regional facies of a species a

geographical race, ecotype, topodeme or

genoecodeme”).

Australian species of Actephila are the

only euphorbs s.l. occurring in rainforests to

have been examined by electrophoretic

methods, viz. isozyme analysis (Shapcott 1998).

That work examined a small number of

populations from the species A. grandifolia,

A. lindleyi and A. vernicosa and although it

revealed variation, the sampling was

insufficient to draw wider conclusions.

Electrophoretic data is variable in its usefulness

in distinguishing specific taxa in other plant

groups (e.g. cycads: Sharma etal. 1998, 1999,

2004). Morphologically discrete species may

differ little genetically (see references in Sharma

et al. 1998) and the application of isozymes to

taxonomic decisions is limited as speciation can

occur without divergence at enzyme loci or

Forster, Revision of Actephilia

divergence in allozyme loci can occur without

speciation (Crawford 1989). The conserved

nature of allozyme variation responsible for

these apparently low levels of variation (Bossart

& Prowell 1998) may not be reflected in results

from more informative molecular techniques

such as amplified fragment length

polymorphisms (AFLP), microsatellites (ITS) or

inter-simple sequence repeats (ISSR).

The delimitation of species for this genus

in Australia has been difficult, as considerable

variation occurs for each taxon, nearly all of

which are known from a range of disjunct

populations in eastern and northern parts of

the continent (Map 1). Fertile material can be

hard to obtain, and unless care is taken, most

flowers are lost from dried specimens as they

often abscise during the drying process. This

has resulted in a limited sampling of floral

diversity from the available herbarium

collections. Despite this, all of the species

recognised below are distinctive when

encountered in habitat. The habit of the

individual plant and form of the foliage changes

little when the plants are cultivated. Further

characters are undoubtedly to be found in the

morphology of seedlings based on limited

observations of cultivated plants; however, in

the absence of comprehensive information of

this sort it is the intention of this revision to

provide a classification and identification key

that can be used with vegetative adult material.

The first species to be recognised for

Australia was Actephila lindleyi (as the

misapplied name Securinega nitida Lindl.) in

1821 (Airy Shaw 1971). Further species were

added by Mueller (1865) and Baillon (1866a),

although these were considered conspecific

with A. lindleyi by Airy Shaw (1971). The basic

classification framework for Australian species

Map 1. Distribution of Actephila in Australia within 1° grids and species diversity.

was established by Bentham (1873) who named

A. latifolia , A. petiolaris and A. sessilifolia, as

well as recognising A. lindleyi (as A. nitida).

Over 100 years later little had changed with Airy

Shaw (1980b, 1980c, 1981) recognising five

species, A. foetida Domin, A. latifolia,

A. lindleyi, A. petiolaris and A. sessilifolia for

the genus in Australia.

The current revision represents a major

departure from the long-accepted status quo of

species diversity in the Australian species of

Actephila , with the number of species more than

doubled. This has been due to a number of

factors, most notably the considerable

accumulation of herbarium material (together

with supporting spirit collections) during the

1980’s and 1990’s from previously known

populations, and secondly continued botanical

exploration of the rainforests of Australia that

has resulted in the discovery of new

populations and undescribed taxa. The presence

of several of these undescribed species has

been indicated in the allozyme study of Shapcott

(1998) for a small number of species

(A. grandifolia, A. lindleyi, A. vernicosa of this

account), census accounts of the flora of

Queensland (Forster & Henderson 1997; Forster

& Halford 2002), and identification manuals to

rainforest plants (Hyland et al. 1999, 2003;

Cooper & Cooper 2004).

Reproductive Ecology

Ecology of Actephila species has been barely

studied and little more than circumstantial and

anecdotal information is available on their

reproduction, pollination, dispersal and

recruitment. Floyd (1989) stated that A. lindleyi

fruit were “eaten by [the] brown cuckoo-dove”

and that “cuttings strike easily”. All of the

Australian species of Actephila comprise small

trees, shrubs or subshrubs and occur in

rainforest (mature phase) communities in New

South Wales, the Northern Territory and

Queensland. Plants tend to mass flower

following rain between October and December.

Flowering is often associated with the

production of new leaves and the shape and

colour of these unhardened leaves is often quite

different to the mature leaves that are present

by the time of fruiting. This pattern of floral and

Austrobaileya 7 (1): 57-98 (2005)

vegetative phenology is similar to other species

of Euphorbiaceae and Putranjivaceae that occur

in these habitats, e.g. Croton (Forster 2003),

Drypetes (Forster 1997b) or Mallotus (Forster

1999b).

The Australian species of Actephila are

monoecious, a trait commonly associated with

Euphorbiaceae s.l. in this region (Gross 2005).

The flowers are clustered in small axillary

fascicles with staminate flowers usually greatly

outnumbering pistillate ones, but there is no

spatial or temporal separation of the sexes as is

found in species of Croton (Forster 2003) or

Mallotus (Forster 1999b). In Actephila it is not

unusual to observe individuals with

inflorescences where the flowers are all of one

sex (usually male), particularly if the plants are

recovering from a period of stress (usually

drought); however it is uncertain whether this

indicates evolution of androdioecy, “the

presence of male-only and hermaphroditic

plants in a population” (Charlesworth 1984;

Thomson et al. 1989) without quantitative

observations. An alternative explanation is that

Actephila plants may be gender diphasic, i.e.

the plants ‘change sex’ (Lloyd 1982;

Korpelainen 1998; Richardson & Clay 2001), with

many of the ‘male’ plants later having both male

and female flowers.

Pollinators of Actephila are

undocumented, but the shallow, bowl-like

flowers are perhaps most suited to generalist

insect pollinators with short proboscises

(Endress 1994), as has been found in Drypetes

(Forster 1997b) (Euphorbiaceae/Putrajivaceae)

by Williams & Adam (1997) or postulated for

Croton (Forster 2003). The presence of foetid

smelling flowers in species such as Actephila

foetida and A. traceyi may indicate that flies

are the primary pollinators and this group of

insects are sometimes the only pollinators in

dioecious species (Bawa 1980; Charlesworth

1993; Renner & Feil 1993). Although the

possibility of wind pollination (Williams &

Adam 1999) cannot be discounted without

experimentation, it is perhaps less likely due to

the occurrence of Actephila populations in the

rainforest understorey where wind speed is

greatly reduced (Turner 2001).

Forster, Revision of Actephilia

Plants of Actephila when cultivated in

areas well removed from their natural habitats

still produce fruit with fertile seed (R. Jensen, G.

Sankowsky, M.C. Tucker, pers. comm. 2004) and

this has been interpreted as indicating that

generalist pollination by opportunistic insects

is occurring (Tucker & Simmons 2004). The

possibility of apomixis or agamospenny (i.e. “the

development of seeds without preceeding

fertilization”), as found for the euphorbiaceous

Alchornea ilicifolia (Smith 1841; Baillon 1866c;

Endress 1994; Bicknell & Koltunow2004) is more

likely, but needs to be experimentally confirmed.

Agamospermous reproduction may also explain

the apparent morphological uniformity of plants

within any given population of Actephila

resulting from long-term recruitment of

genetically similar individuals.

Pistillate flowers are often persistent on

the plant with both the pedicels and floral parts

increasing in size until the perianth (especially

the petals) withers and the fruit develops. At

what stage these flowers cease to be

functionally receptive to pollination and should

be regarded as developing faiit is uncertain and

a somewhat arbitrary distinction in the

morphological descriptions has been made

based on persistence of the petals. Esser (2003)

considered that the genus Actephila was

characterised by autochorous/diplochorous

fruit dehiscence and dispersal, whereby the

fruits are ballistically explosive (ballistochores,

viz. Turner 2001), followed by myrmechory.

The seeds of Actephila have highly

reduced or no caruncles present, so once they

have been explosively ejected could be

dispersed by ants that move the seeds as a result

of being attracted to a food reward. These

caruncles are significantly smaller (percentage

of the seed volume) than those found in other

Australian Euphorbiaceae s.l. such as Acalypha

L. (Forster 1994b), Austrobwcus Miq. (Forster

1997a), Bertya Planch. (Halford & Henderson

2002a), Croton L. (Forster 2003), Dissiliaria

F.Muell. exBaill. (Forster 1996b), Micrantheum

Desf. (Berg 1975), Monotaxis Brongn. (Halford

& Henderson 2002b), Petalostigma F.Muell.

(Forster & van Welzen 1999), Pseudanthus

Sieber ex Spreng. (Halford & Henderson 2003),

Sankowskya P.EForst. (Forster 1995a),

Stachystemon Planch. (Halford & Henderson

2003) and Wyanbeelia Airy Shaw & B.Hyland

(Forster unpubl.) and their use as a food source

by invertebrates has yet to be observed. Species

with this dispersal pattern are unlikely to have

regular, long-distance dispersal, i.e. greater than

100 m from the parent (Cain et al. 1999) and

most seeds of rainforest trees fall within 10 m of

the parent plant (Turner 2001). This dispersal

pattern usually results in a markedly clumped

disposition of individuals, especially for species

that inhabit the understorey and midstratum of

rainforests, but has only been quantitatively

documented for A. lindleyi at one locality in

south-eastern Queensland (Butler 2004). In

species such as A. bella, A. championiae,

A. latifolia, A. petiolaris, A. plicata,

A. sessilifolia and A. vernicosa , the boundaries

of populations are often sharp, indicating

“dispersal limited distribution” (Primack & Miao

1992). In comparison species such as

A. grandifolia, A. lindleyi and A. venusta do

not always have such a ‘sharp’ boundary to

populations, and although individuals are

clumped, they are generally more widely

scattered within rainforest patches.

Australian Distribution and Diversity

Australia is the major centre of diversity for

Actephila with at least fourteen endemic

species. There are collectively fewer species

combined in the other regions where the genus

has been found. The Australian species are also

the most morphologically diverse with the

observed variation mainly that of leaf

morphology and foliage indumentum. The most

distinctive species in the whole genus is

undoubtedly A. foetida with its massive leaves

and foliage indumentum. Toothed leaves, as

found in A. excelsa from Malesia, have not been

observed in the Australian species.

As yet no infrageneric phylogeny has

been proposed fox Actephila', however, obvious

morphological groupings of the Australian

species are (1 )A. bella, A. foetida, A. petiolaris ,

(2) A. flavescens, A. grandifolia, A. lindleyi,

A. traceyi, A. venusta, A. vernicosa, (3)

A. championiae, A. latifolia, A. plicata, (4)

A. longipedicellata, A. sessilifolia. These

informal groupings are based on morphological

similarities in leaf and fruit morphology. Similar

appearing, yet distinct, species occur in New

Guinea and elsewhere; however my knowledge

of these taxa is insufficient to suggest further

groupings throughout the genus range.

Species of Actephila are restricted to

rainforests in eastern and northern Australia

(Map 1), although they are absent from the

Kimberley of Western Australia and are found

south only to central New South Wales. Species

sympatry is rarely encountered with the only

instances I am familiar with being (1) A. lindleyi

-A. sessilifolia at Bundaberg) and (2)A. plicata

-A. sessilifolia at Mt Dryander near Proserpine.

Hence the current day distribution pattern of

species, especially within rainforests of similar

structure and floristic composition, is

essentially allopatric. This is strictly the case

for the species within each of the groups

outlined above with the geographic

disjunctions ranging from nearby (e.g.

A. championiae - A. plicata ; A. foetida -

A. petiolaris ; A. grandifolia - A. lindleyi ;

A. traceyi-A. venusta ) to far flung (e.g. A. bella

- A. petiolaris ; A. flavescens - A. lindleyi ;

A. longipedicellata - A. sessilifolia).

While there are minor centres of species

diversity in Australian Actephila, i.e. three grid

cells in the ‘Wet Tropics’ and one grid cell near

Miriamvale with three or four species each

(Map 1), there is no overall similarity with the

patterns of species distribution and diversity

found in the Australian taxa of Croton L. with

many more species (Forster 2003) or Mallotus

Lour, that has a similar number of species

(Forster 1999b).

Materials and Methods

This revision is based on herbarium holdings

at BRI, CANB (including CBG), DNA, MEL,

NSW and QRS, photographs or microfiche of

types at G-DC, K, L, P and U, and field

collections and observations of all taxa by the

author. Representative material (including

types) of Actephila species from New Guinea

and Indonesia were examined at BRI, CANB, L

and QRS.

Most species of Actephila remain poorly

collected, and the descriptions for most of the

species in this revision are based on few fertile

collections. Floral descriptions are incomplete

for A. bella, A. flavescens, A. petiolaris subsp.

Austrobaileya 7 (1): 57-98 (2005)

jagonis and A. plicata. Collectors should ensure

that both male and female flowers are obtained

when making specimens of Actephila. It is not

unusual to encounter mass flowering

individuals of Actephila, however, most flowers

are lost from specimens when the material is

handled and dried. If spirit for flower/fruit

pickling is available for use in the field, then

flowers or fruit should be immediately pickled.

Male flowers usually greatly outnumber females

giving the impression that the plants are

dioecious, but careful searching will usually

result in location of material of both sexes.

Floral descriptions were prepared from

material preserved in spirit (FAA or 70% alcohol

and glycerol) or reconstituted by boiling in water

and detergent. Fruit descriptions were prepared

from spirit and dried material. Foliage,

inflorescence and seed descriptions were

prepared from dried material.

The degree of leaf margin recurving is

useful in specific delimitation and distinctions

have been made between ‘flat’ (the leaf lamina

margin is in the same plane as the rest of the

leaf lamina), ‘recurved’ (the leaf lamina margin

is curved in a plane 0-90° to the plane of the leaf

lamina) and ‘revolute’ (the leaf lamina margin is

curved in a plane 90-180° to the plane of the leaf

lamina). Venation terminology largely follows

Hickey (1973) and Ash et al. (1999) with the

recognition of a midrib (1° vein order), lateral

veins (2° vein order) and intercostal veins (3°

and onwards vein orders) within any leaf lamina.

When an intercostal vein comprises a

continuous raised line of cells it is termed

‘distinct’; if it is discontinuous or fades away

into the body of the lamina, it is termed

‘indistinct’. Indumentum cover is described

using the terminology of Hewson (1988), except

that ‘scattered’ is used instead of‘isolated’. The

shapes of leaves, sepals and petals are

described using the terminology of Hickey &

King (2000).

Characters most commonly used in the

identification key are those of indumentum

cover, leaf lamina venation (especially vein order

number and its expression) and leaf lamina

margin form, colour of foliage parts (based on

dried material), flowers (particularly disk form

and size, sepal and petal shape and size). Use

Forster, Revision of Actephilia

of non-vegetative characters in the couplets has

been avoided in most cases but is necessary to

distinguish several species towards the end of

the key.

Common abbreviations in the specimen

citations are N.P. or N.P.R. (National Park or

National Park Reserve), L.A. (Logging Area),

F.R., S.F. or S.F.R (Forest Reserve, State Forest

or State Forest Reserve), T.R. (Timber Reserve)

and V.C.L. (Vacant Crown Land).

NCA is an abbreviation for the

Queensland Nature Conservation Act (1992) and

its associated schedules. Discussions of

conservation rankings are made using the

criteria of the IUCN (2001).

Rainforest structural terminology follows

Webb (1978). The ‘Wet Tropics’ is defined as

that area of north-eastern Queensland which

encompasses the ‘hot, humid vine forests’ from

near Cooktown in the north to Paluma in the

south (Webb & Tracey 1981), and is equivalent

to the bioregion of the same name (Goosem et

al. 1999). For the purposes of the key, ‘Central

Qld’ comprises the pastoral districts of North

Kennedy, South Kennedy and Port Curtis;

‘Southern Qld’ comprises the pastoral districts

of Burnett, Darling Downs, Moreton and Wide

Bay; ‘Cape York Peninsula’ is that part of

Queensland north of the ‘Wet Tropics’.

Taxa are arranged alphabetically.

Suggested affinities between taxa are indicated

in the Notes section for each taxon.

Taxonomy

Actephila Blume, Bijdr. FI. Ned. Ind. 581 (1825).

Savia section Actephila (Blume) Baill., Etude

Euphorb. 571 (1858). Type species:

Actephila javanica Miq.

Lithoxylon Endl., Gen. PI. 2:1122 (1840). Type

species: Securinega nitida Lindley ( non

Willd. 1806).

Anomospermum Dalz., Hook. J. Bot. Kew

Gard. Misc. 3:228 (1851)(now Miers 1851).

Type species: Anomospermum excelsum

Dalz. [= Actephila excelsa (Dalz.)

Muell.Arg.]

Perennial trees, shrubs or subshrubs,

monoecious, evergreen; stems and foliage

without latex. Indumentum of simple trichomes;

scales, glandular trichomes and stinging

trichomes absent. Stipules entire, generally

inconspicuous, deciduous. Leaves alternate,

subverticillate or subopposite, coriaceous or

chartaceous, flat to plicate, ± sessile to petiolate,

simple and elobate, penninerved, entire or

toothed, glands absent. Inflorescences

fasciculate, axillary, extra-axillary or rarely on

short brachyblasts, or solitary, uni- or bisexual,

with flowers in bracteate groups. Male flowers

pedicellate; calyx lobes 4-6, imbricate, ± equal;

petals 2-6, often scale-like or absent; disk extra-

staminal, annular, 4-5 lobed, or entire and

rounded; stamens 3-6, inserted on the disk,

filaments free, filiform; anthers introrse,

bilobate, thecae oblong and longitudinally

dehiscent; pistillode trifid. Female flowers

pedicellate; calyx lobes 4-6, imbricate; petals

2 -6, usually often scale-like or absent; disk

annular or cupular, fleshy; ovary 3-locular,

locules biovulate; styles free or shortly connate

at base, entire or bifid. Fruits capsular, trilobate

to pentalobate, surface rough, glabrous or

pubescent, dehiscing loculicidally and partly

septicidally into bivalved cocci; columella

persistent. Seeds deltoid; testa crustaceous;

albumen absent; caruncles absent or present;

cotyledons thick, sometimes plicate.

Between twenty-five and thirty species

in Asia, Australia, Malesia and Melanesia.

Fourteen species in Australia, all endemic.

Derivation of name : akte (seashore) and

phileo (to love).

Austrobaileya 7 (1): 57-98 (2005)

Key to Australian species of Actephila

1. Leaf lamina chartaceous (thin).2

Leaf lamina coriaceous (thick, leathery) .3

2. Branchlets with dense indumentum when young; leaf petioles 18-78 mm

long, dark olive-green when dry. 4. A. foetida

Branchlets glabrous when young; leaf petioles 3-8 mm long, black-green

when dry.14. A. vernicosa

3. Leaf lamina with up to 4 or 5° vein orders distinct.4

Leaf lamina with up to 3° vein orders distinct (rarely 4° in ,4. lindleyi ).5

4. Leaf petiole length 2-8 mm long; lamina length/petiole length ratio 13.6-42.5

(Cape York Peninsula). 8.A.longipedicellata

Leaf petiole length 9-80 mm long; lamina length/petiole length ratio 2-8.7

(Wet Tropics).9. A. petiolaris

5. Leaflaminawithupto2° vein orders distinct.6

Leaf lamina with up to 3° vein orders distinct (rarely 4° in A. lindleyi ).9

6. Leaf margin flat to slightly recurved.7

Leaf margin revolute.8

7. Leaf petiole cream when dried; lamina base obtuse or rounded (rarely

subcordate) (Central Qld). 1. A. bella

Leaf petiole olive-green when dried; lamina base acute (rarely rounded)

(Cape York Peninsula, Northern Territory). 13. A. venusta

8 . Leaf petiole length 10-44 mm long, green-yellow when dried; lamina base

acute or rounded (rarely weakly cordate). 2. A. championiae

Leaf petiole length 1-7 mm long, grey-fawn when dried; lamina base cordate

to subcordate, often appearing stem-clasping. 11. A. sessilifolia

9. Leaf margin revolute.10

Leaf margin flat to recurved (rarely revolute in A. lindleyi ).11

10. Branchlets cream when dry; leaf petioles cream-yellow when dry; fruit

rounded (Wet Tropics) . 3. A. flavescens

Branchlets brown when dry; leaf petioles olive-green when dry; fruit

pentalobate (Central Qld). 10. A. plicata

11. Midrib of leaf lamina below prominently raised at lamina base (height >

width) then decreasing towards apex (Wet Tropics). 6. A. latifolia

Midrib of leaf lamina below strongly raised at lamina base (height < or ±

equal to width) then decreasing towards apex (Cape York Peninsula, Central

and Southern Qld, NSW).12

12. Male flower sepals 3.5-5 mm long, disk 5-angled; fruit 16-24 mm diameter

(Cape York Peninsula).12.A.traceyi

Male flower sepals 1-3.5 mm long, disk annular or rounded; fruit 11-15 mm

diameter (Central and Southern Qld, NSW).13

13. Abaxial leaf surface pale green and matt-glaucous, often drying brown;

male flower pedicels 10-20 mm long, petals 1.4-1.5 x 0.5-0.7 mm. 5. A. grandifolia

Abaxial leaf surface pale green-yellow and glossy, often drying pale brown;

male flower pedicels 1.5-5 mm long, petals 2-2.5 x 0.8-1.3 mm. 7. A. lindleyi

Forster, Revision of Actephilia

Fig. 1. Actephila bella. A. abaxial leaf surface x]B. side view of male flower x6. C. face view of male flower x6.

D. side view of fruit x4, E. face view of fruit x4. A, D, E from Forster PIF5859 (BRI); B, C from 27 Jan. 2004,

Tucker s.n. (BRI). Del. W. Smith.

1. Actephila bella P.I.Forst., sp. nov. Ab

A. petiolari Benth. petiolo folii cremea

(versus adversum olivaceam), venis

paginae abaxialis folii usque ad tantum

ordine 1-2 (adversum ordines 1-4)

distinguibilibus, floribus masculis

pedicellis longis (13-15 x c. 0.8 mm)

filiformibus (adversum pedicellos breves

validos (3-7 x 0.6-0.8)), petalis oblongis

et longioribus (2.5-3 x 0.4-0.7 mm) (non

petala oblanceolata et breviora (1.8-2.5 x

0.5-1.2 mm) et disco majore (6-7.5 mm

diam. adversum 3.5-6 mm diam.) differt.

Typus: Queensland. Port Curtis District:

Bulburin State Forest (S.F. 67 Bulburin), c. 400

m NW of junction of Granite & Boobook Creeks,

24° 32’S, 151° 30’E, 17 April 1980, W.J.

McDonald 3260, W.J. Fisher & P. Ryan (holo:

BRI [2 sheets])

Actephila sp. (Granite Creek P.I.Forster

PIF5859) (Forster & Henderson 1997: 70;

Forster & Halford 2002:68).

Subshrub or shrub to 3 m high. Indumentum of

simple trichomes, fawn, entire plant is glabrous

unless stated otherwise. Branchlets glossy,

cream-fawn, sparsely lenticellate with age,

glabrous or with scattered to sparse

indumentum when young, glabrescent. Stipules

ovate-triangular, 0.8-1.2 mm long, 0.7-1.2 mm

wide, with scattered cilia. Leaves alternate,

coriaceous, petiolate; petioles 3^17 mm long,

0 .8-1 mm wide, cream, lenticellate, glabrous or

with sparse indumentum; lamina length/petiole

length ratio 2.6-18.7 (n=39); lamina elliptic-

ovate, rarely obovate, ± flat, 56-145 mm long,

23-61 mm wide, length/width ratio 1.9-3.1

(n=39); base obtuse to rounded, rarely

subcordate and unequal; tip acute to acuminate,

rarely rounded; midrib above slightly raised

(height < 5 times the width), pale green, below

strongly raised at lamina base (height ± equal

to width) then decreasing towards apex, pale

olive-green; venation brochidodromous,

comprising 10-14 lateral veins per side of midrib

and intercostal reticulate veins; upper surface

dark green and glossy, lateral venation visible,

intercostal venation visible; lower surface pale

olive-green and matt; lateral and intercostal

venation prominent and slightly raised, 1° and

2° vein orders distinct, 3° and 4° vein orders

indistinct, 5° and onwards vein orders obscure;

margin entire, flat, pale olive-green.

Inflorescence an axillary fascicle 1—1.5 mm

diameter, single sex or with both male and female

flowers. Male flowers: pedicels filiform, 13-15

mm long, c. 0.8 mm diameter; sepals 5, fused for

2.5- 3 mm, lobes oblong-ovate, 2.5-3.5 mm long,

3.5- 5 mm wide, glabrous or sparsely ciliate in

patches towards apices; petals absent or

present as one or two, oblong, 2.5-3 mm long,

0.4-0.7 mm wide; disk conspicuous, fleshy, 1.5-

2 mm high, 6-7.5 mm diameter, 5-angled; stamens

3-5, filaments 2.5-3 mm long, 0.3-0.5 mm

diameter, anthers 0.7-0.8 mm long, 1-1.2 mm

wide. Female flowers not seen; sepals 5, oblong-

ovate, c. 2.5 mm long, 1.8-2 mm wide; petals not

seen; disk conspicuous, fleshy, c. 1.5 mm high,

4.5- 5 mm diameter, 5-angled. Fruit: pedicels c.

18 mm long and 0.5 mm diameter; capsules

depressed-globose to subglobose, rounded, 9-

10 mm long, 12-13 mm diameter, strongly bullate,

green; columella not seen. Seeds not seen. Fig.

1 .

Additional specimens examined : Queensland. Port

Curtis District: SE end of Granite Creek, Bulburin S.F.

391, 24° 38’S, 151° 33’E, Oct 1989, Forster PIF5859

(BRI, L); Bulburin Forest Reserve, Pine Creek, 24°

37’S, 151° 33’E, Jan 2004, Forster PIF29892 & Tucker

(BRI); S.F. 67 Bulburin, Granite Creek, Jul 1978,

McDonald 2374 & Stanton (BRI); Shoalwater Bay

Training Area, c. 4.5 km W of Old Raspberry Vale, 22°

33’S, 150° 20’E, Sep 1993, McDonald 5685 & Meter

(BRI); cult. Kholo, Ipswich (ex S.F. 391 Bulburin, Pine

Creek), Jan 2004, Tucker s.n. (BRI).

Notes: Actephila bella shows some superficial

similarities to A. petiolaris from nearly 5° of

latitude further north, but differs from that

species in the cream leaf petiole (versus olive-

green), the abaxial leaf lamina surface with 1

and 2° vein orders distinct (versus 1-4°

distinct); the male flowers with long (13-15 xc.

0.8 mm), filiform pedicels (versus short (3-7 x

0.6-0.8), stout pedicels), petals that are oblong

and longer (2.5-3 x 0.4-0.7 mm) (versus petals

that are oblanceolate and shorter (1.8-2.5 x 0.5-

1.2 mm)) and the larger disk (6-7.5 mm diameter

versus 3.5-6 mm diameter).

The floral descriptions for this species are

incomplete. I have seen only two male flowers

(from the Pine Creek population) and no female

flowers.

Distribution and habitat: Actephila bella is

known from two 1° grid cells with populations

at Shoalwater Bay and Bulburin Forest Reserve

(several localities) (Map 5). Plants grow in the

understorey of ‘dry’ rainforest (araucarian

notophyll vineforest) on alluvium and

substrates derived from andesite at altitudes

between 160 and 300 m.

Austrobaileya 7 (1): 57-98 (2005)

Conservation status: Actephila bella is an

inconspicuous plant and is likely to be

overlooked by collectors. At Pine Creek (within

the Bulburin Forest Reserve) this species grows

in close association with the critically

endangered Macadamia jansenii C.L.Gross &

P.H.Weston. Individual numbers of the

Actephila at this locality are not high (less than

500 plants). The number of individuals at the

other two localities is not known. Further

survey work is required to determine an area of

occupancy and an estimation of the number of

individuals; however, it is likely that the species

should be listed as Endangered.

Etymology: The specific epithet is formed from

the Latin word bellus (beautiful) and refers to

the flowers of this species.

2. Actephila championiae P.I.Forst., sp. nov. ab

A. latifoliae Benth. differt ramulis cremeis

(adversum atro-brunneos), lamina folii

basi obtusa rotundata rarius infirme

cordata (adversum attenuatam usque

cuneatam), margine revoluto (non piano)

et venis in pagina abaxiali folii usque ad

tantum ordinibus 1-2 (adversum ordines

1-3) distinguibilibus, floribus foemineis

petalis oblanceolatis vel oblongis vel

spathulatis (non obovatis) et majoribus

(3-3.2 x 1-2 mm non 1-2 x 0.5-1 mm) et

discis majoribus (6-7 mm diam. adversum

3-4 mm).

Typus: Queensland. South Kennedy District:

Black Mt area, 21 0 36’ S, 149° 1 O’ E, 20 September

1993, 1.G. Champion870 (holo: BRI; iso: DNA,

MEL).

Actephila sp. (Koumala I.GChampion 870)

(Forster & Henderson 1997: 70; Forster &

Halford 2002:68).

Shrub or small tree to 8 m high. Indumentum of

simple trichomes, uncoloured, entire plant is

glabrous unless stated otherwise. Branchlets

matt to slightly glossy, cream, lenticellate with

age. Stipules triangular, 1.2-2 mm long, 0.8-1.3

mm wide, irregularly ciliate. Leaves alternate,

rarely subverticillate, coriaceous, petiolate;

petioles 10-44 mm long, 1.5-2 mm wide, green-

yellow, lenticellate; lamina length/ petiole length

ratio 3.6-13.2 (n=64); lamina elliptic, obovate,

oblanceolate or oblong, ± flat, 65-220 mm long,

Forster, Revision of Actephilia

Fig. 2. Actephila championiae. A. habit of flowering branch x0.5. B. abaxial leaf surface x0.25. C. side view of

male flower x3. D. face view of male flower x4. E. side view of female flower x3. F. face view of female flower x4.

G. side view of dehiscing fruit xl.5. H. face view of dehiscing fruit xl.5. A, C, D-F from Champion 870 (BRI); B,

G, H from Champion 90OB (BRI). Del. W. Smith.

12-110 mm wide, length/width ratio 1.8-2.8

(n=64); base acute to rounded, rarely weakly

cordate and unequal; tip acute to rounded, rarely

retuse; midrib above ± flat, dark green, below

prominently raised at lamina base (height >

width) then decreasing toward apex, pale

yellow; venation brochidodromous, comprising

10-12 lateral veins per side of midrib and

intercostal reticulate veins; upper surface dark

green and glossy, lateral venation visible,

intercostal venation ± obscure; lower surface

pale green-yellow and matt, lateral and

intercostal venation prominent, only laterals

raised, 1° and 2° vein orders distinct, 3° and 4°

vein orders indistinct, 5° and onwards vein

orders obscure; margin entire, revolute, pale

yellow. Inflorescence an axillary fascicle 3-4 mm

diameter, rarely interpetiolar or on a short

peduncle to 5 mm long, single sex or with both

male and female flowers. Male flowers: pedicels

filiform, 10-15 mm long, 0.8-1.3 mm diameter;

sepals 5, oblong to obovate, fused for 2-2.5

mm at base, free lobes 2.5-4 mm long, 1.7-3 mm

wide, ciliate in patch at apices; petals absent or

one or two present, oblanceolate, oblong or

spathulate, 3.2-4 mm long, 0.8-1.8 mm wide;

disk conspicuous, fleshy, 1.5-2 mm high, 8-9

mm diameter, strongly 5-(6)-angled; stamens 5,

filaments 2.5-4 mm long, c. 0.2 mm diameter,

anthers, 0.4-0.5 mm long, 0.6-0.8 mm wide.

Female flowers: pedicels stout, 10-18 mm long,

0.8-2 mm diameter; sepals 5, obovate, fused at

base for 1-1.5 mm, free lobes 2.8-3 mm long,

1.7-2 mm wide, ciliate in patch at apices; petals

absent or one or two present, oblanceolate,

oblong or spathulate, 3-3.2 mm long, 1-2 mm

wide; disk conspicuous, fleshy, c. 1.5 mm high,

6-7 mm diameter, strongly 5-angled; ovary 3.5-

4 mm long, 4^1.5 mm diameter; styles 3, simple,

shortly connate at base, 1.7-2 mm long, stigmas

subcapitate. Fruit: pedicels 12-25 mm long, 1-

1.5 mm diameter; capsules depressed-globose

to subglobose, rounded to weakly trilobate, 14-

17 mm long, 14-20 mm diameter, weakly bullate,

green; columella c. 9 mm long. Seeds not seen.

Fig. 2.

Additional specimens examined: Queensland. South

Kennedy District: Black Mt., c. 7 km NW of Koumala,

21° 35’S, 149° 10’E, Dec 1993, Champion 900a et al

(AD, BRI, DNA); loc. cit., Dec 1993, Champion 900b

et al. (BRI); loc. cit., Dec 1993, Champion 900c et al.

(BRI, MEL, NSW); loc. cit., Dec 1993, Champion

900d et al. (BRI); Cameron Creek, Black Mt., just N of

join with rail line, c. 6 km NW of Koumala, 21° 55’S,

149° 25’E, Mar 1996, Champion 1348 & Canning

(BRI, L); Cameron Creek, Black Mt., c. 6.3 km NW of

Koumala township, 21° 55’S, 149° 25’E, Mar 1996,

Champion 1349 & Canning (BRI, DNA, NSW); T.R.

179 Kalvin, Black Mt Range, 21° 36’S, 149° H E, Apr

1991, Forster PIF8030 & McDonald (BRI, L, MEL,

QRS).

Notes: Actephila championiae differs from

A. latifolia Benth. in the cream branchlets

(versus dark brown), the leaf lamina with a base

that is obtuse to rounded, rarely weakly cordate

(versus attenuate to cuneate), a margin that is

revolute (versus flat), and abaxially with 1-2°

of vein orders distinct (versus 1-3°); the female

flowers with petals that are oblanceolate, oblong

or spathulate (versus obovate) and larger (3-

3.2 x 1-2 mm versus 1-2 x 0.5-1 mm) and a

larger disk (6-7 mm diameter versus 3-4 mm).

Together with A. latifolia and A. plicata ,

A. championiae may comprise a natural species

group with the species distributed in an

allopatric geographic replacement sequence

from central Queensland to the top of the ‘Wet

Tropics’.

Austrobaileya 7 (1): 57-98 (2005)

Distribution and habitat: Actephila

championiae is narrowly endemic (single 1° grid

cell) in central Queensland where it is restricted

to the Black Mountain Range near Koumala

(Map 5). Plants grow in the understorey of‘dry’

rainforest (semi-deciduous, complex notophyll

vineforest) on substrates derived from acid

volcanic rocks at altitudes between 160 and 200

Conservation status: Actephila championiae

is not present in any conservation reserves.

Further survey work is required in the Black

Mountain Range to determine an area of

occupancy and estimation of the number of

individuals for the species. Actephila

championiae is currently listed as Vulnerable

under the NCA.

Etymology: Actephila championiae is named

to honour Irene Champion of Mackay, in

recognition of her significant contribution to

the documentation and collection of the flora

of central Queensland. Irene has contributed

over 1300 herbarium collections to the

Queensland Herbarium and made repeated visits

to the habitat of this species to ensure the

collection of fertile material after its initial

discovery in a sterile state.

3. Actephila flavescens P.I.Forst. sp. nov. ab

A. lindleyi (Steud.) Airy Shaw stipulis

minoribus (0.5-0.8 x 0.5-0.8 mm) ovatis

(adversum stipulas majores (1-1.7 x 0.8-

1.5 mm) triangulari-ovatas), petiolis flavo-

cremeis (non brunneis usque olivaceis),

lamina folii margine revoluto (adversum

planum usque recurvum), apice acuto

(rarius acuminato) adversum apicem

acutum vel rotundatum (rarius retusum),

costa abaxiali manifeste emersa ad basim

laminae (alto > latitudine) adversum valde

emersam (alto < latitudine), flavescenti

(adversum pallide flavo-viridi) et frutice

majore(14-16 x 14-15 mm non 9-12 x 11 -

13.5 mm) et seminibus majoribus (9-11 x

7-8 mm non 6-7 x 5-6.5 mm) pallide

olivaceis non atro-olivaceis differt.

Typus: Queensland. Cook District: Daintree

National Park, Adeline Creek headwaters,

Candlenut Scrub, 16° 07’S, 145° 03’E, 18 May

1999, P.I. Forster PIF24571 & R. Booth (holo:

BRI; iso: QRS).

Forster, Revision of Actephilia

Shrub or small tree to 5 m high. Indumentum of

simple trichomes, uncoloured, entire plant is

glabrous unless stated otherwise. Branchlets

glossy, cream, lenticellate with age. Stipules

ovate, 0.5-0.8 mm long, 0.5-0.8 mm wide. Leaves

alternate, subverticillate or subopposite,

coriaceous, petiolate; petioles 4-12 mm long,

0 .8-1.2 mm wide, cream-yellow, lenticellate with

age; lamina length/petiole length ratio 6.5-23.8

(n=27); lamina oblanceolate to obovate, rarely

± falcate, ± flat, 57-120 mm long, 23^42 mm wide,

length/width ratio 2-3.1 (n=27); base acute to

attenuate; tip acuminate, rarely acute; midrib

above strongly raised (height < width), dark

green, below prominently raised at lamina base

(height > width) then decreasing towards apex,

pale yellow; venation brochidodromous,

comprising 10 or 11 lateral veins per side of

midrib and intercostal reticulate veins; upper

surface dark green and glossy, lateral and

intercostal venation visible; lower surface pale

green, glossy, lateral venation slightly raised,

intercostal venation visible and not raised, 1-

3° vein orders distinct, 4° and 5° vein orders

indistinct, 6° and onwards vein orders obscure

or absent; margin entire, revolute, pale yellow.

Inflorescence an axillary fascicle 1-3 mm

diameter, rarely interpetiolar, single sex or with

both male and female flowers. Male flowers not

seen. Female flowers not seen at anthesis;

sepals 5, oblong to ovate, 1.5-3 mm long, 0.8-1

mm wide, externally with a few scattered simple

trichomes, apices cucullate, internally glabrous;

petals not seen; disk conspicuous, fleshy, c. 1

mm high and 3 mm diameter, annular; ovary not

seen; styles not seen, stigmas not seen. Fruit:

pedicels 5-35 mm long, c. 1 mm diameter;

capsules depressed-globose, rounded, 14-16

mm long, 14-15 mm diameter, strongly bullate,

green; columella 7-8 mm long. Seeds 9-11 mm

long, 7-8 mm wide, pale olive green; hilum c. 2

mm long, caruncle poorly developed, narrow-

oblong, c. 2 mm long and 0.5 mm wide, cream.

Additional specimens examined : Queensland. Cook

District: Cedar Bay, 15° 47’S, 145° 21’E, Jan 1973,

Dick sub Webb & Tracey 13774 (QRS); Mt Hedley, 3

km ENE of Rossville, T.R. 165, 15° 44’S, 145° 17’E,

Apr 1999, Forster PIF24293 & Booth (BRI, QRS);

Shipton’s Flat on Tin Mine road, 15° 45’S, 145° 10’E,

May 1969, Smith 14358 (BRI).

Notes : This is a poorly collected species and

its morphological description is incomplete. I

have seen only sterile or fruiting material and

collectors should seek flowers when

encountering this plant. Actephila flctvescens

appears to be closely allied to A. lindleyi but is

disjunct by 2° of latitude from populations of

that species and differs in the smaller (0.5-0.8 x

0.5-0.8 mm) ovate stipules (versus larger (1-1.7

x 0 .8-1.5 mm) triangular-ovate stipules); the

cream-yellow petioles (versus brown to olive-

green stipules); the leaf lamina with a revolute

margin (versus flat to recurved), tip that is

acuminate (rarely acute) versus acute or rounded

(rarely retuse), the midrib abaxially prominently

raised at lamina base (height > width) (versus

strongly raised (height < or ± equal to width)

and pale yellow (versus pale green-yellow); and

the larger fruit (14-16 x 14—15 mm versus 9-12

x 11-13.5 mm) and larger seeds (9-11 x 7-8 mm

versus 6-7 x 5-6.5 mm) that are pale olive-green

(versus dark olive green).

Actephila ftavescens appears to form a

natural species group with A. grandifolia,

A. lindleyi, A. traceyi, A. venusta and

A. vernicosa , with all of these species being

geographically allopatric.

Distribution and habitat: Actephila ftavescens

is known from a restricted area (single 1° grid

cell) in the ‘Wet Tropics’ of north-eastern

Queensland between Mt Hedley and Adeline

Creek (Map 2). Plants have been collected from

‘dry’ rainforest (semideciduous, complex

notophyll vineforest) on substrates derived

from granite at altitudes between 480 and 540

Conservation status: Actephila ftavescens is

present in the Daintree National Park and Timber

Reserve 165. At Mt Hedley and Adeline Creek

plants were locally common, but no attempt was

made at the time to determine population sizes.

Further survey is required of suitable habitats

within the known distribution area to determine

an area of occupancy and estimation of the

number of individuals for the species. It is likely

that A. ftavescens is not currently threatened.

Etymology: The specific epithet is derived from

the Latin adjective ftavescens (yellowish, pale

yellow) and refers to the prominent leaf lamina

midrib of this species (when compared to

A. lindleyi ) as viewed from below.

Austrobaileya 7 (1): 57-98 (2005)

Fig. 3. Actephila foetida. A. abaxial leaf surface with detail of indumentum x0.25. B. side view of male flower x6.

C. face view of male flower x7. D. side view of fruit xl.5. E. face view of fruit xl.5. A-C from Forster PIF28216

(BRI); D, E from Jago 6004 (BRI). Del. W. Smith.

4. Actephila foetida Domin, Biblioth. Bot. 89:

315 (1927). Type: Queensland. Cook

District: in hygrodrymio apud rivulum

Harveys Creek, distr. Cairns, December

1909, K. Domin 5839 (lecto [here

designated]: PR528400).

Illustrations : Hyland etal. (1999,2003).

Subshrub to 1 m tall. Indumentum of simple

trichomes, pale fawn. Branchlets glossy, fawn,

densely pubescent when young, glabrescent

and lenticellate with age. Stipules triangular, 1-

2.2 mm long, 1.5-2 mm wide, with scattered to

dense indumentum. Leaves alternate,

chartaceous, petiolate; petioles 18-78 mm long,

1-5 mm wide, dark olive-green, with dense

indumentum when young, glabrescent and

lenticellate; lamina length/petiole length ratio

2.8-12.8 (n=21); lamina broadly elliptic to

obovate, ± flat, 45-530 mm long, 30-213 mm

wide, length/width ratio 1.9-2.7 (n=21); base

cordate to truncate; tip obtuse to long

acuminate; midrib above ± flat, dark olive-green,

below prominently raised at lamina base (height

> width) then decreasing towards apex, dark

olive-green; venation brochidodromous,

comprising 12-15 lateral veins per side of midrib

and intercostal reticulate veins; upper surface

dark olive-green, lateral venation visible,

intercostal venation ± obscure, glabrous or with

an occasional trichome on the midrib; lower

surface pale olive-green, lateral venation

prominent and raised, intercostal venation

prominent but not raised, 1-4° vein orders

distinct, 5° vein order indistinct, 6° and onwards

vein orders obscure, with sparse, hispid

indumentum, apart from on the lateral veins

where it may be dense; margin entire, flat, pale

olive-green. Inflorescence an axillary fascicle 7-

13 mm diameter, with both male and female

flowers. Male flowers: pedicels filiform, 3.5-8

mm long, 0.5-0.7 mm diameter, with sparse, hispid

indumentum; sepals 5, lanceolate-ovate to

obtuse-ovate, 2-2.5 mm long, 1-1.8 mm wide,

ciliate, with sparse indumentum externally,

glabrous internally; petals absent; disk

conspicuous, fleshy, c. 1 mm high, 3.5^1 mm

diameter, irregularly 4 or 5-angled; stamens 4 or

5, filaments 1.3-1.8 mm long, c. 0.2 mm diameter,

glabrous, anthers globose-ovoid, c. 0.4 mm long,

0.3-0.4 mm wide. Female flowers: pedicels

filiform, 7.5-9 mm long, 0.5-0.7 mm diameter, with

sparse, hispid indumentum; sepals 5, obovate,

3^1 mm long, 2-2.8 mm wide, ciliate, with sparse

indumentum externally, glabrous internally;

petals absent; disk conspicuous, fleshy, c. 1

mm high, 4^1.5 mm diameter, irregularly 4 or 5

angled; ovary 1.4-1.8 mm long, 1.8-2 mm

diameter, with dense indumentum; styles 3,

simple, c. 1 mm long, stigmas subcapitate. Fruit:

pedicels 7-35 mm long, 1-2 mm diameter,

Forster, Revision of Actephilia

glabrous; capsules depressed-globose,

rounded to weakly trilobate, 12-13 mm long,

15-19 mm diameter, strongly bullate; columella

3.5-7 mm long. Seeds 8-10 mm long, 7-12 mm

wide, hilum 1.5-2 mm long, caruncle poorly

developed, narrow-oblong, 2.3-3 long, 0.3-0.8

mm wide, cream. Fig. 3.

Additional specimens examined : Queensland. Cook

District: 8 km from Bellenden Ker, 17° 16’S, 145°

55’E, Dec 1992, Bruhl 1022 et al. (BRI, QRS); Harveys

Creek, Jan 1910, Domin 3410 (PR); loc. cit, Dec 1909,

Domin 5838 (PR - residual syntype); Bellenden Ker

cable car Base Station, 17° 16’S, 145° 54’E, Jan 2002,

Forster PIF28216 et al. (BRI); adjacent to Bellenden

Ker Primary School, 17° 16’S, 145° 55’E, Nov 1994,

Gray 5844, 5845 (QRS); loc. cit., Dec 1994, Gray

5881 (QRS); N.P.R. 226 (Harvey Creek), 17° 15’S,

145° 50’E, Jan 1972, Hyland 5788 (BRI, QRS); adjacent

to Bellenden Ker State School on Harvey Creek road,

17° 17’S, 145° 55’E, Mar 1981, Irvine 2092, 2094

(QRS); Harveys Creek, Bellenden Ker, Dec 1991, Jago

C3694 (BRI); Parish of Bellenden Ker, Local Gov.

Res. 806 & part of adjoining road reserve, 17° 16’S,

145° 55’E, Jun 1985, Lyons 2 (BRI); loc. cit., Dec

1985, Lyons 5 (QRS); Hutchinson’s Creek, Cow Bay,

16° 13’S, 145° 24’E, Oct 1996, Small 5 (QRS); Harvey

Creek, Russell River, 1887, Sayer s.n. (MEL); Bellenden

Ker, 17° 17’S, 145° 54’E, Mar 1981, Stocker 1782

(QRS).

Distribution and habitat : Actephila foetida

has a restricted distribution in the Wet Tropics

of north-eastern Queensland with two centres

of distribution in adjacent 1° grid cells: the

vicinity of the township of Bellenden Ker near

and along Harvey Creek and north of the

Daintree River at Hutchinson Creek (Map 4).

Plants occur as understorey shrubs in ‘wet’

lowland rainforest (evergreen, complex

mesophyll vineforest) on alluvium overlying

granite substrates at altitudes from near sea

level to 100 m.

Typification : The protologue for Actephila

foetida states “Nordost-Queensland:

Regenwalder bei Harveys Creek (DOMIN

XII. 1909)”. There are three collections of this

species by Domin at PR, one (PR528398:

fruiting) was collected in January 1910, the other

two (PR528399 & PR528400: both flowering) in

December 1909. The better of these two

(PR528440) has three complete leaves and both

male flowers and fruiting peduncles and is

selected here as lectotype of the name.

Notes : Actephila foetida is poorly known in

terms of the potential variation in its morphology

due to a paucity of fertile collections. It is a

highly distinctive species due to the large leaves

(generally between 20 and 35 cm long) that are

covered in hispid indumentum on the lower

lamina surface and the flowers that lack petals

and that have hispid indumentum on the

pedicels and the calyx. Airy Shaw (1981)

described the fruit of this species as 5-6 mm

long, but these measurements must have been

taken from immature material.

The population at Hutchinson’s Creek

lacks comprehensive fertile collections, but

appear to have sparser indumentum on the

foliage than those from Harvey Creek.

Conservation status : Actephila foetida is

currently listed as Vulnerable under the NCA.

Populations are present in Daintree and

Wooroonooran National Parks. There is no

accurate information about the area of

occupancy for the species or the number of

individuals that exist. The species is locally

common at the base of Bellenden Ker and along

Harvey Creek.

Etymology: The specific epithet is formed from

the Latin word foetidus (stinking, evil smelling)

and refers to the smell of the male flowers.

5. Actephila grandifolia (Mull.Arg.) Baill.,

Adansonia 6: 330 (1866).

Lithoxylon grandifolium Mull.Arg., Linnaea

34: 65 (1865). Type: New South Wales.

(“Nouvelle Hollandia”), Clarence River, F.

Mueller s.n. (holo: G-DC [photo at BRI!]

Illustration: Baillon (1866b: 1.10).

Subshrub or shrub to 3 m high. Indumentum of

simple or pustular-glandular trichomes,

uncoloured to pale ferruginous, entire plant is

glabrous unless stated otherwise. Branchlets

glossy, tan, lenticellate with age, glabrous or

with scattered to sparse pustular-glandular

trichomes, glabrescent. Stipules triangular to

triangular-ovate, 0.8-2 mm long, 0.7-1.5 mm

wide. Leaves alternate, subverticillate or

subopposite, coriaceous, petiolate; petioles 3-

30 mm long, 0.8-2 mm wide, cream, lenticellate

with age; lamina length/petiole length ratio 4.3-

18 (n=184); lamina oblanceolate to obovate,

rarely narrow-elliptic, often weakly falcate, ±

flat, 31-240 mm long, 10-64 mm wide, length/

width ratio 2.3-5.8 (n=184); base acute,

Austrobaileya 7 (1): 57-98 (2005)

Fig. 4. Actephila grandifolia. A. abaxial leaf surface x0.5. B. side view of male flower x4. c. face view of male

flower x4. D. side view of female flower x4. E. face view of male flower x4. F. side view of fruit xl.5. G. face view

of fruit xl.5. A, F, G from Forster PIF25276 (BRI); B-E from Forster PIF25283 (BRI). Del. W. Smith.

acuminate, attenuate or cuneate, often unequal;

tip acute or acuminate, rarely retuse, rounded

or truncate; midrib above slightly raised (height

< 5 times the width), pale yellow, below strongly

raised at lamina base (height < or ± equal to

width) then decreasing towards apex, pale

green-yellow; venation brochidodromous,

comprising 7-14 lateral veins per side of midrib

and intercostal reticulate veins; upper surface

dark green and glossy, lateral and intercostal

venation visible; lower surface pale green, matt-

glaucous, often drying brown, lateral venation

slightly raised, intercostal venation visible and

slightly raised, 1-3° vein orders distinct, 4° and

5° vein orders indistinct, 6° and onwards vein

orders obscure; margin entire to slightly wavy

or undulate, recurved, pale green-yellow.

Inflorescence an axillary fascicle 2-4 mm

diameter, single sex or with both male and female

flowers. Male flowers: pedicels filiform, 10-20

mm long, 0.1-0.4 mm diameter; sepals 5,

obovate, 2-3.5 mm long, 1.8-3 mm wide, apices

with a dense patch of hairs internally, ciliate

and with scattered hairs towards base externally;

petals absent, or one to three present, oblong

to spathulate, 1.4-1.5 mm long, 0.5-0.7 mm wide,

irregularly bidentate; disk conspicuous, fleshy,

0.8-1 mm high, 3-5 mm diameter, annular, white;

stamens 3 or 4, filaments 2.2-4.5 mm long, 0.1-

0.2 mm diameter, anthers globose-ovoid, 0.5-

0.6 mm long, 0.7-0.8 mm wide. Female flowers:

pedicels filiform, 7-28 mm long, 0.4-0.5 mm

diameter, glabrous or with scattered, uncoloured

simple trichomes; sepals 5, obovate, 2.2-3 mm

long, 1-2 mm wide, apices internally densely

ciliate, with scattered to sparse trichomes

towards base externally; petals absent or one

or two present, oblong to spathulate, 1-1.5 mm

long, 0.5-0.8 mm wide; disk conspicuous, fleshy,

0.8-1 mm high, 2.5-5 mm diameter, annular;

ovary 1-3 mm long, 1.2-3 mm diameter; styles

3, simple, shortly connate at base, 0.8-1 mm

long, stigmas subcapitate. Fruit: pedicels 7-28

mm long, 0.5-1 mm diameter; capsules

depressed-globose, rounded, 9-11 mm long,

12-15 mm diameter, strongly bullate, green;

columella 5-8 mm long. Seeds 6-9.5 mm long,

5-8.5 mm wide, dark olive green with irregular

darker green blotches; hilum 2.5-3.5 mm long,

caruncle poorly developed, narrow-oblong, 1-

2.5 mm long, 0.2-0.3 mm wide, cream. Fig. 4.

Selected specimens examined : Queensland. Moreton

District: Lyrebird road, Springbrook, 28° It’S, 153°

15’E, s.dat., Davidson s.n. (BRI [AQ440657]); Camp

Eden, Currumbin Valley, 28° 14’S, 153° 21’E, Dec 1997,

Forster PIF22023 & Leiper (BRI, DNA, NE, QRS);

Springbrook, Lyre Bird Ridge road, 28° ll’S, 153° 15’E,

Jan 2000, Forster PIF25276 & Leiper (BRI, K, L,

MEL, QRS); Springbrook, Repeater Station road to

Forster, Revision of Actephilia

Best of All Lookout, 28° 13’S, 153° 15’E, Jan 2000,

Forster PIF25283 (A, AD, BRI, K, L, MEL, NSW,

QRS); Upper Currumbin Creek, 2 km ESE of Boyd’s

Butte, Aug 1976, McDonald 1497 & Whiteman (BRI,

CANB). New South Wales. Bexhill, 28° 46’S, 153°

2UE, Feb 1891, Baker 120 (NSW); Mullumbimby, 28°

33’S, 153° 30’E, Jun 1894, Baker s.n. (NSW239914,

239915); 6 km NW of Bungalow, 28° 31’S, 153° 28’E,

Jan 1984, Beesley 89 (CANB); 3 km W of Burringbar,

28° 27’S, 153° 26’E, Jan 1984, Beesley 101 (CANB);

Richmond River, Aug 1884, Betche s.n. (NSW239763);

Rocky Creek, Whian Whian S.F., 17.6 km NNE of

Lismore, 28° 39’S, 153° 20’E, Sep 1972, Coveny 4446

& Rodd (BRI, NSW); Big Scrub Reserve on Rocky

Creek, Whian Whian S.F., 4.8 km NNE of Dunoon,

28° 38’S, 153° 20’E, Jan 1980, Coveny 10597 & Hind

(BRI, NSW); Upper Crystal Creek, 28° 16’S, 153° 18’E,

May 1977, Floyd AGF347 (CANB); Hill 1 km E of

Mooball, 28° 27’S, 153° 30’E, Jul 1981, Guymer 81038

& Harden (NSW); Richmond River, Sep 1867,

Henderson 58 (MEL); Red Scrub, Whian Whian, 28°

34’S, 153° 21’E, Feb 1967, Jones 3469 (CANB);

Tintenbar, 28° 48’S, 153° 31’E, Oct 1894, Maiden

s.n. (NSW239894); Eureka, Richmond River, 28° 40’S,

153° 26’E, Jul 1915, Rupp s.n. (MEL47993); Whian

Whian S.F., Big Scrub F.R., 28° 38’S, 153° 20’E, Jun

1986, Shapcott s.n. (BRI [AQ451831]); Marshall Falls,

Alstonville, 28° 50’S, 153° 26’E, Sep 1911, Tanner

s.«.(NSW239853); Red Scrub, Whian Whian S.F., 28°

38’S, 153° 17’E, Aug 1974, Webb & Tracey 11456

(BRI).

Distribution and habitat : Actephila

grandifolia has a restricted distribution in south¬

eastern Queensland and north-eastern New

South Wales where it is known from a single 1 °

grid cell with a northern limit at Springbrook

and a southern limit at Alstonville (Map 5).

Plants grow in the understorey of subtropical

‘wet’ rainforest (complex notophyll vineforest)

or in open forest dominated by eucalypts and

with a scrubby rainforest understorey.

Substrates are soils derived from basalt at

altitudes between 40 and 950 m.

Notes : Actephila grandifolia occurs in the

wetter rainforests (evergreen, complex notophyll

vineforest) of the Springbrook Plateau and

extreme north-east of New South Wales. It is

ecologically allopatric with A. lindleyi, although

the distance between populations of the two

species may not be great. The differences

between Actephila grandifolia and A. lindleyi

(as A. mooreana ) were eloquently outlined by

Baillon (1866b) together with a particularly

detailed plate of the former. Actephila

grandifolia was lumped into the synonymy of

A. lindleyi by Airy Shaw (1971) and as a result

disappeared from botanical view for the last 30

years (e.g. Stanley 1983; James & Harden 1990;

Forster etal. 1991; Forster & Henderson 1997;

Hauser & Blok 1998; Forster & Halford 2002),

despite being recognised in herbaria (BRI,

NSW) prior to this. Airy Shaw must have had

second doubts about this placement, as in 1974

he determined specimens in MEL as A. cf.

grandifolia.

Actephila grandifolia is easily

distinguished from A. lindleyi by the leaves with

cream petioles (versus brown to olive-green),

the foliage being noticeably matt-glaucous

below (versus glossy) and the male flowers with

much longer, filiform pedicels (10-20 x 0.1-0.4

mm versus 1.5-5 x 0.2-0.8 mm), obovate sepals

(versus oblong to ovate) and smaller petals (1.4-

1.5 versus 2-2.5 mm long).

A chromosome count of 2n = 40 was

recorded for material of A. grandifolia collected

near “Lismore” (probably the Whian Whian S.F.

collection cited above) (Shapcott 1998). On the

basis of several morphological and chromosome

characters Shapcott (1998) clearly distinguished

her “Lismore” population from three

populations of A. lindleyi sensu stricta.

However, this was not the case with

electrophoretic data where the “Lismore”

population was indistinguishable from the

A. lindleyi populations at “Mt Glorious” and

“Biggenden”

Conservation status : Actephila grandifolia is

present in Springbrook National Park in

Queensland and Whian Whian State Forest in

New South Wales. Plants are usually locally

common within a population. No information is

available on the area of occupancy for each

population, or the numbers of plants within

individual populations.

This species (as “A. lindleyi ”) is used as

a nest site in the Richmond Range of north¬

eastern New South Wales by the yellow-

throated scrubwren Sericornis citreogularis,

the nests of that species then being used by

the golden-tipped bat Kerivoula papuensis as

a roost site (Schulz 2000). This bird and bat use

a range of other species, so it is likely that the

use of the Actephila is related to abundance

and habit within the site and conservation of

the animals is not dependent on this particular

plant species.

Etymology : The specific epithet is formed from

the Latin words grandis (large) and -folius

(leaved) and alludes to the size of the leaves in

this species when compared to A. lindleyi.

6. Actephila latifolia Benth., FI. Austral. 6: 89

(1873). Type: Queensland. Cape York,

Daemels.n. (lecto: K [photo atBRI \\fide

Airy Shaw (1981: 587)).

Actephila sp. (Little Mossman BH13697)

(Hyland etal. 1999,2003).

Illustrations : Hyland etal. (1999,2003).

Subshrub or shrub to 3 m high. Indumentum of

simple trichomes, uncoloured, entire plant is

glabrous unless stated otherwise. Branchlets

glossy, dark brown, lenticellate with age.

Stipules triangular, 1.5-2 mm long, 0.8-1.2 mm

wide. Leaves alternate to subverticillate,

coriaceous, petiolate; petioles 3-20 mm long,

1-2 mm wide, olive green, occasionally

lenticellate; lamina length/petiole length ratio

10.8-22.5 (n=104); lamina elliptic, narrow-elliptic

or oblanceolate, rarely obovate or orbicular, ±

flat, 40-220 mm long, 16-80 mm wide, length/

width ratio 1.9-4.2 (n=104); base attenuate to

cuneate; tip acute, obtuse or rounded; midrib

above slightly raised (height < 5 times the

width), dark green, below prominently raised at

lamina base (height > width) then decreasing

towards apex, green-yellow; venation

brochidodromous, comprising 10-14 lateral

veins per side of midrib and intercostal reticulate

veins; upper surface dark green and matt, lateral

venation barely visible, intercostal venation

obscure; lower surface olive green and matt,

lateral venation prominent and slightly raised,

intercostal venation visible but not raised, 1-3°

vein orders distinct, 4° vein order indistinct, 5°

and onwards vein orders obscure; margin

entire, flat, green-yellow. Inflorescence an

axillary fascicle 3-5 mm diameter, single sex or

with both male and female flowers. Male flowers:

pedicels filiform, 2.5-7.5 mm long, 0.8-1 mm

diameter; sepals 5, obovate to ovate, 2-3 mm

long, 1.2-1.8 mm wide, apices cucullate and

densely ciliate; petals absent, or rarely present

as a single one, obovate, c. 1.5 mm long and 0.8

mm wide, irregularly bidentate; disk

conspicuous, fleshy, 0.8-1 mm high, 2.5—4 mm

diameter, 5-angled; stamens 3-5, filaments 1.2-

Austrobaileya 7 (1): 57-98 (2005)

2.5 mm long, c. 0.1 mm diameter, anthers globose-

ovoid, 0.3-0.5 mm long, 0.5-0.7 mm wide. Female

flowers: pedicels filiform, 5-7 mm long, 0.8-1

mm diameter; sepals 5, obovate to ovate, 2-2.5

mm long, 1.5-1.8 mm wide, apices pouched and

densely ciliate; petals absent or rarely present

as a single one, obovate, 1-2 mm long, 0.5-1

mm wide; disk conspicuous, fleshy, 0.8-1 mm

high, 3^1 mm diameter, 5 or 6-angled; ovary 2-

2.5 mm long, 2-3 mm diameter; styles 3, simple,

shortly connate at base, 0.8-1 mm long, stigmas

subcapitate. Fruit: pedicels 2-32 mm long, 0.8-

1 mm diameter; capsules depressed-globose,

rounded, 12-14 mm long, 14-17 mm diameter,

strongly bullate, green; columella 6-7 mm long.

Seeds 8-9 mm long, 5.5-6.5 mm wide, olive green

with irregular dark green blotches; hilum 1.4-3

mm long, caruncle poorly developed, narrow-

oblong, 2-3 mm long, 0.8-1 mm wide, cream.

Fig. 5.

Additional specimens examined : Queensland. Cook

District: N.P.R. 142, summit of Mt Cook, 15° 29’S,

145° 16’E, Feb 1992, Fell DGF2416 et al. (QRS); Mt

Perserverence road, near Nissen Creek via Julatten,

16° 33’S, 145° 20’E, Feb 2003, Ford 3838 & Holmes

(BRI, QRS); Bridle L.A., S.F. 607 Dinden, 16° 58’S,

145° 35’E, Jul 1994, Forster PIF15486 et al. (BRI,

QRS); Bridle L.A., S.F. 607 Dinden, 16° 58’S, 145°

36’E, Jul 1994, Forster PIF15491 et al. (BRI, QRS);

Lake Morris, Copperlode Dam, 16° 59’S, 145° 39’E,

Dec 1999, Forster PIF25217 & Booth (BRI, MEL,

QRS); S.F. 607 Dinden, Bridle Creek track, 16° 59’S,

145° 36’E, Jan 2002, Forster PIF28120 et al. (A, AD,

BISH, BRI, MEL, MO, NE, NSW, NY, Z); Currunda

Creek, Redlynch, 7 km W of Cairns, 16° 55’S, 145°

40’E, Dec 2003, Forster PIF29786 & Jensen (A, BRI,

L, MEL, NSW, Z); S.F.R. 141, Little Mossman L.A.,

16° 33’S, 145° 22’E, Jun 1995, Gray 6238, 6239 (QRS);

Special Purposes Reserve, near Lake Placid, 16° 52’S,

145° 40’E, Nov 1991, Jago 737 (QRS); Moores Gully,

16° 48, 145° 40’E, Nov 1987, Sankowsky 756 &

Sankowsky (BRI); cult. Tolga (ex Rex Range), Dec

2004, Sankowsky 2553 & Sankowsky (BRI); Mt

Stuckey area, NW of Starke Station, 14° 56’S, 145°

03’E, Sep 1974, Tracey 14295 (BRI); Macalister Range,

16° 50’S, 145° 40’E, Dec 1999, Wannan 865 & Jago

(NSW); Smithfield - Black Mt road, via Kuranda, 1962,

Webb & Tracey 7269A (BRI); Mt Stuckey, W of Starcke

Station, 14° 56’S, 145° 03’E, Sep 1974, Webb & Tracey

13789 (BRI).

Distribution and habitat : Actephila latifolia

is endemic to north-eastern Queensland over

three 1 ° grid cells in the ‘Wet Tropics’ bioregion

with a northern limit at Mt Stuckey and a

southern limit at Bridle Creek and Lake Morris

(Map 5). Although the type collection is labelled

as ‘Cape York’ it is unlikely that it was made

Forster, Revision of Actephilia

Fig. 5. Actephila latifolia. A. abaxial leaf surface x 0 . 25 . b. side view of male flower x6. C. face view of male flower

x6. D. side view of female flower x6. E. face view of female flower x6. F. side view of fruit xl,5, G. face view of

fruit xl.5. H. ventral view of seed x3. A, F-H from Forster PIF15491 (BRI); B-E from Forster PIF29786 (BRI).

Del. W. Smith.

from the Cape of that name, but rather from

somewhere in the vicinity of Cooktown at the

southern end of Cape York Peninsula. Plants

grow at altitudes between 200 and 500 m, in

‘seasonally dry’ rainforest (complex mesophyll

or complex notophyll vineforests) on alluvial or

rocky substrates derived from basalt, granite,

metamorphics or mudstone.

Notes : Actephila latifolia is a distinctive plant

that appears closely allied to the more southerly

distributed A. championiae and A. plicata.

These species share the characteristics of

leaves that are pale green-yellow to olive green

below, but differ in a number of characters, such

as leaf shape, lamina venation and fruit shape.

As noted previously (Forster 2004),

Actephila latifolia has been mis identified for

some time as an undescribed species (Hyland

etal. 1999,2003).

Conservation status : Actephila latifolia is

present in Mt Cook and Starcke National Parks,

Jumrum Creek Conservation Park and State

Forests 141,607,1229. It is not considered rare

or threatened and none of the known

populations are under direct anthropogenic

threat. There is no information available on the

area of occupancy or the number of individuals

within populations.

Etymology: The specific epithet is derived from

the Latin words latus (broad, wide) and -folius

(leaved) and alludes to the size and shape of

the leaves.

7. Actephila lindleyi (Steud.) Airy Shaw, Kew

Bull. 25:496(1971).

[Securinega nitida auct. Lindl., non Willd.;

Collect. Bot. 19(1821)]

Lithoxylon lindleyi Steud., Nomenclat. Bot.,

ed. 2,2:57 (1841), nom. nov. Type: Cult, in

Hort. Kew, “Nat. of Otaheite” [exact origin

unknown], 1813-1821 (holo: K [photo at

BRI!]).

Lithoxylon nitidum Baill., Et. Gen. Euphorb.

590 (1858), nom. illegit., superfl.

Actephilanitida (Baill.) Benth. & Hook.f. ex

Drake del Castillo, Illustr. insul. Maris

Pacific 177 & 286 (1892), nom. illegit.,

superfl.

Actephila mooreana Baill .,Adansonia 6: 330,

366 (1866). Type: New South Wales. Mount

Lindsay, Richmond River, 1851, C. Moore

211 (holo: P [photo at BRI!]

Illustrations: Floyd (1989: 134); James &

Harden (1990:393); Hauser & Blok (1998:145).

Shrub or small tree to 6 m high. Indumentum of

simple trichomes, uncoloured to pale

ferruginous, entire plant is glabrous unless

stated otherwise. Branchlets glossy, cream to

tan-yellow, lenticellate with age. Stipules

triangular-ovate, 1-1.7 mm long, 0.8-1.5 mm

wide. Leaves alternate, subverticillate or

subopposite, coriaceous, petiolate; petioles 2-

24 mm long, 1-2 mm wide, brown to olive-green,

lenticellate with age; lamina length/petiole

length ratio 5.1-22.7 (n=218); lamina elliptic-

oblong, oblanceolate or obovate, rarely ±

falcate, ± flat, 30-172 mm long, 12-64 mm wide,

length/width ratio 1.6-3.7 (n=218); base acute

to attenuate; tip acute to rounded, rarely retuse;

midrib above slightly raised (height < 5 times

the width), dark green, below strongly raised at

lamina base (height < width) then decreasing

towards apex, pale green-yellow; venation

brochidodromous, comprising 7-12 lateral veins

per side of midrib and intercostal reticulate

veins; upper surface dark green and glossy,

lateral and intercostal venation visible; lower

surface pale green-yellow, glossy, often drying

pale brown, lateral venation slightly raised,

intercostal venation visible and slightly raised,

1-3 (rarely 4)° vein orders distinct, 4° and 5

(rarely 6)° vein orders indistinct, 6 (rarely 7)°

and onwards vein orders obscure; margin entire

to slightly wavy or undulate, flat to recurved

(rarely revolute), pale yellow. Inflorescence an

axillary fascicle 1-4 mm diameter, rarely

Austrobaileya 7 (1): 57-98 (2005)

interpetiolar, single sex or with both male and

female flowers. Male flowers: pedicels filiform,

1.5-5 mm long, 0.2-0.8 mm diameter; sepals 5

(rarely 6), oblong to ovate, 1-3 mm long, 1.2-1.5

mm wide, apices cucullate, ciliate internally;

petals absent, or 1-5, linear-oblong to oblong-

spathulate, 2-2.5 mm long, 0.8-1.3 mm wide,

rarely ciliate; disk conspicuous, fleshy, 0.4-1 mm

high, 1.2-3.5 mm diameter, rounded, yellow;

stamens 4 or 5, filaments 1-3 mm long, 0.1-0.2

mm diameter, anthers globose-ovoid, 0.3-0.6 mm

long, 0.4-0.8 mm wide. Female flowers: pedicels

filiform, 2-14 mm long, 0.4-1 mm diameter,

glabrous or with scattered to sparse,

uncoloured to pale ferruginous simple

trichomes; sepals 5, oblong to ovate, 1-3 mm

long, 0.5-1.5 mm wide, apices cucullate,

internally densely ciliate; petals absent or 1-3,

linear-oblong to oblong-spathulate, 0.8-2.5 mm

long, 0.3-1.5 mm wide; disk conspicuous,

fleshy, 0.5-1 mm high, 2-4 mm diameter, annular

to 5-angled, glabrous, rarely with scattered

trichomes; ovary 0.8-2.5 mm long, 1.2-2.5 mm

diameter; styles 3, simple, shortly connate at

base, 0.5-1 mm long, stigmas subcapitate. Fruit:

pedicels 3-25 mm long, 0.6-1.2 mm diameter;

capsules depressed-globose, ovoid or rounded,

9-12 mm long, 11-13.5 mm diameter, strongly

bullate, olive-green; columella 5-7 mm long.

Seeds 6-7 mm long, 5-6.5 mm wide, dark olive

green; hilum 2-3 mm long, caruncle poorly

developed, ± obscure or narrow-oblong, 1-1.5

mm long, c. 0.2 mm wide, cream. Fig. 6.

Selected specimens examined : Queensland. North

Kennedy District: Hervey Range, off Hervey Range road,

W of Townsville, Sep 1995, Gumming 13728 (BRI).

South Kennedy District: Eastern side of Spencer Gap,

Peak Downs Highway, SW of Eton, 21° 20’S, 148°

56’E, Nov 1989, McDonald 4428 et ai. (BRI, NSW,

QRS). Port Curtis District: Lower reaches of

Koolkoorum Creek, 24° 25’S, 151° 12’E, Dec 1993,

Forster PIF14262 (A, BRI, L, MEL, QRS). Burnett

District: S.F. 95, 20 km WSW of Gayndah, 25° 42’S,

151° 25’E, Mar 1999, Forster PIF24166 & Booth

(AD, BRI, K, L, MEL, NSW, QRS); Kirbys road,

Bundaberg, 24° 49’S, 152° 23’E, Sep 1999, Forster

PIF24918 & Schmitt (BRI); Goodnight Scrub National

Park, 25° 14’S, 151° 54’E, Jan 2004, Forster PIF29902

& Tucker (BRI, MEL, NSW). Wide Bay District: Council

Beauty Spot, 1 km E of Mt Cooroy, 26° 26’S, 152°

58’E, Nov 1990, Forster PIF7587 & Sharpe (BRI, L,

QRS); Utopia, 14 km SSE of Biggenden, 25° 38’S,

152° 06’E, Dec 1991, Forster PIF9215 (BRI, MEL);

Mt Glastonbury, S.F. 242 Glastonbury, 26° 14’S, 152°

27’E, Dec 1991, Forster PIF9290 & Sharpe (BRI, K,

L, MEL, QRS); S.F. 220 Malmaison, Oakview Forestry,

26° 07’S, 152° 20’E, Mar 2000, Forster PIF25469 &

Forster, Revision of Actephilia

Fig. 6. Actephila lindleyi. A. abaxial leaf surface *0.5. B. side view of male flower x6. C. face view of male flower

x6. D. side view of female flower x6. E. face view of female flower x6. A from Forster PIF14262 (BRI); B-E from

Forster PIF24166 (BRI). Del. W. Smith.

Booth (BRI, MEL, QRS). Moreton District: Gold Creek,

2 km WNW of North Arm, 26° 30’S, 152° 55’E, Nov

1990, Forster PIF7581 & Sharpe (BRI, CANB, K, L,

MEL, QRS); 0.5 km SW of McAfees Lookout, S.F.

309, 27° 25’S, 152° 52’E, Feb 1991, Forster PIF7788

& Bird (BRI, QRS); F.R. 809, Mt Glorious, D’Aguilar

Range, NW of Brisbane, 27° 20’S, 152° 45’E, Dec

2004, Forster PIF30403 & Jensen (BRI, L, MEL,

NSW); Mary Cairncross Scenic Reserve, Blackall Range,

3 km SE of Maleny, 26° 49’S, 152° 52’E, Dec 2004,

Forster PIF30426 et ai. (BRI, L, MEL, NSW). New

South Wales, c. 10 miles [16 km] WSW of Dungog

on the road to Gresford, Nov 1970, Blaxell & Coveny

3337 (BRI, CANB, NSW); 1 km E of Toolona Lookout,

headwaters of Hopping Dicks Creek, 28° 16’S, 153°

10’E, Nov 1982, Guymer 1807 & Jessup (BRI);

Wiangaree S.F., c. 10 miles NE of Wiangaree, 28° 33’S,

153° 06’E, Oct 1966, Hayes 2534 et ah (NSW);

Wongawilli, S of Wollongong, 34° 28’S, 150° 45’E,

Aug 1986, Mills s.n. (NSW540765); Border Ranges

N.P., Coff Lyon’s road, near Border Loop, 28° 20’S,

153° 38’E, May 1986, Shapcott s.n. (BRI [AQ451829]);

Yessabah Caves near Kempsey, 31° 06’S, 152° 43’E,

Aug 1967, Telford IRT78 (CANB).

Distribution and habitat : Actephila lindleyi

has the broadest distribution of the Australian

species in this genus (twenty 1° grid cells),

ranging from slightly north of Townsville south

to Wongawilli near Woollongong (Map 2). It is

a rarely encountered tree at its southern limit

(Mills & Jakeman 1995). This species is

invariably found in ‘dry’ rainforests (araucarian

microphyll or semi-deciduous, complex

notophyll vineforests and semi-evergreen

vinethickets) on a variety of substrates derived

from andesites, basalts or metamorphics at

altitudes between 40 and 500 m. At some

localities (e.g. LamingtonNational Park, Bunya

Mountains National Park), this species may be

a dominant part of the rainforest understorey

(Laidlaw et al. 2000; Butler 2003; Kitching et al.

2004).

Notes: Airy Shaw (1971,1981) openly admitted

that A. lindleyi as he circumscribed it, was

“extraordinarily variable... .there is need of much

more material before the limits of its variation

can begin to be understood”. In the present

account considerable material that was

previously identified as A. lindleyi is referred

to the species A. flavescens, A. grandifolia,

A. traceyi, A. venusta and A. vernicosa.

Chromosome counts of 2n = 40 were

recorded for material of A. lindleyi collected at

Mt Glorious and the Border Ranges (Shapcott

1998).

A probable, undescribed taxon allied to

A. lindleyi has been collected at Conway Range

near Airlie Beach (Vouchers: Eastern catchment

of Repulse Creek, Conway N.P, c. 3 km W of

Woodcutter Bay, 20° 22’S, 148° 47’E, Oct 1993,

McDonald 5834 & Champion (BRI); Conway

S.F., c. 9 km SE of Airlie Beach, 20° 20’S, 148°

45’E, Oct 1993, McDonald 5848 & Champion

(BRI); Conway S.F., c. 2 km E of Impulse Creek

crossing, 20° 21’S, 148° 44’E, Feb 1999,

McDonald6623 & Kemp (BRI)) where it occurs

in ‘seasonally dry’ rainforest (semi-deciduous,

complex notophyll vineforest) fringing

watercourses at an altitude of c. 60 m on

substrates derived from granite. This material

differs from A. lindleyi in leaf morphology

(principally venation expression); however,

only fruiting material has been collected so far.

Conservation status : Actephila lindleyi is

widespread and locally abundant in many of

the extant populations. In Queensland this

species is present in Boombana, Fairlies Knob,

Goodnight Scrub, Kondalilla, Lamington, Main

Range, Mapleton Falls, Mt Bauple, Mt

Collosseum, Mt Pinbarren and Nicoll Scrub

National Parks, Forest Reserves 287, 309, 471

and 1344, State Forests 9,28,53,86,95,220,242,

435, 866, 893 and 1294. In New South Wales it

has been recorded from Border Ranges National

Park and Wiangaree State Forest.

Etymology : The species is named for John

Lindley (1799-1865), one time Professor of

Botany at University College, London, and

specialist on Orchidaceae (Stearn 1999).

8. Actephila longipedicellata P.I.Forst. sp. nov.

ab A. sessilifoliae Benth. differt lamina folii

basi acuta usque cuneata (adversum

cordatam usque subcordatam, saepe

aspectu amplexicauli propter petiolum

brevem), numero majore venarum

lateralium (11-15 non 8-11), floribus

masculis pedicellis longioribus (25-34 mm

adversum 8-11 mm), sepalis majoribus

(3.5-5 x 2.5^4 mm adversum 1.8-2.5 x 1.5-

1.8 mm), petalis spathulatis (adversum

petala obovata usque oblonga) majoribus

(2.8-3.5 x 1.3-2.5 mm non 0.8-1.8 x 0.3-1

mm) et disco 5-angulato (adversum

discum annularem) et duplo majore (2.4-5

mm diam. adversum 2-2.8 mm).

Typus: Queensland. Cook District: Garraway

Creekrockpiles, 12°45’S, 143° ll’E, 16April

1988, PL ForsterPIF4227 & D.J. Liddle (holo:

BRI [1 sheet + spirit]; iso: DNA, KEP, SAN,

SAR).

Actephila sp. (Claudie River BH 7803)

(Hyland etal. 1999,2003).

Subshrub or shrub to 4 m high. Indumentum of

simple trichomes, uncoloured, entire plant is

Austrobaileya 7 (1): 57-98 (2005)

glabrous unless stated otherwise. Branchlets

glossy, cream to green, lenticellate with age.

Stipules triangular to triangular-ovate, 0.7-2 mm

long, 0.7-1.2 mm wide, glabrous or irregularly

ciliate. Leaves alternate or subverticillate,

coriaceous, petiolate; petioles 2-8 mm long, 1-

2 mm wide, cream to green, lenticellate; lamina

length/petiole length ratio 13.6-42.5 (n=78);

lamina obovate to oblanceolate, ± flat, 33-195

mm long, 12-68 mm wide, length/width ratio 1.8-

3.7 (n=78); base acute to cuneate; tip acute to

acuminate, rarely rounded; midrib above slightly

raised (height < 5 times the width), pale green,

below prominently raised at lamina base (height

± equal to or > width) then decreasing towards

apex, pale green-yellow; venation

brochidodromous, comprising 11-15 lateral

veins per side of midrib and intercostal reticulate

veins; upper surface dark green and glossy,

lateral and intercostal venation visible; lower

surface pale olive-green and glossy, lateral and

intercostal venation prominent, only laterals

raised, 1-5° vein orders distinct, 6° vein order

indistinct, 7° and onwards vein orders obscure;

margin entire, flat to slightly recurved, pale

yellow. Inflorescence an axillary fascicle 1-4 mm

diameter, rarely interpetiolar, single sex or with

both male and female flowers. Male flowers:

pedicels filiform, 25-34 mm long, 0.5-0.8 mm

diameter; sepals 5, ovate, fused for c. 1 mm at

base, free lobes 3.5-5 mm long, 2.5-4 mm wide;

petals 5, spathulate, 2.8-3.5 mm long, 1.3-2.5

mm wide; disk conspicuous, fleshy, c. 1 mm high,

4.5-5.5 mm diameter, annular to weakly 5-

angled; stamens 5, filaments 3-3.5 mm long, c.

0.5 mm diameter, anthers globose-ovoid, 0.4-

0.5 mm long, 0.7-0.8 mm wide. Female flowers:

pedicels filiform, 20^17 mm long, 0.5-2 mm

diameter; sepals 5, ovate to obovate, fused at

base for c. 1 mm, free lobes 5-6 mm long, 3.5-

4.5 mm wide, glabrous or ciliate along entire

edge; petals 5, oblong to spathulate, 3.5-5.5

mm long, 1-2.5 mm wide; disk conspicuous,

fleshy, c. 1.5 mm high, 5.5-7 mm diameter, annular

to weakly 5-angled, annular; ovary 2-4 mm long,

2-5.5 mm diameter; styles 3-(5), simple, shortly

connate at base, 2-2.5 mm long, stigmas

subcapitate. Fruit: pedicels 30-55 mm long, 0.5-

1 mm diameter; capsules globose, rounded, 20-

25 mm long, 14-17 mm diameter, strongly bullate,

green; columella 7-8 mm long. Seeds 11-15 mm

long, 7-9 mm wide, hilum 3^1 mm long, camncle

Forster, Revision of Actephilia

Fig. 7. Actephila longipedicellata. A. habit of fruiting branch x0.3. B. abaxial leaf surface x0.5. C. side view of

female flower x3. D. face view of female flower x3. E. side view of female flower x4. F. face view of female flower

x4. G. face view of fruit xl. H. side view of fruit xl. A-D, G, H from Forster PIF15462 (BRI); E, F from Fell

DGF2562 (BRI). Del. W. Smith.

poorly developed, narrow-oblong, 4-6 long, c.

0.2 mm wide, cream. Fig. 7.

Additional specimens examined : Queensland.

Cook District: Tozer Gap, Tozer Range, Jul 1948,

Brass 19449 (BRI); Ridges above Claudie River,

4.5 km NE of Mt Tozer, 12° 44’S, 143° 1.5’E, May

1992, Fell DGF2562 (AD, BRI, MEL, QRS);

Kennedy Hill Gorge, 12° 28’S, 143° 16’E, Jun 1989,

Forster PIF5411 & Tucker (BRI); Tozer Range, 12°

44’S, 143° 12’E, Jul 1994, Forster PIF15462 (BRI,

MEL); Claudie River, 12° 45’S, 143° 15’E, Oct

1973, Hyland 6931 (BRI, QRS); loc. cit., Oct 1974,

Hyland 7803 (BRI, QRS); loc. cit., Nov 1998,

Hyland 16147 (BRI, QRS); cult. Tolga (ex Garraway

Creek), 12° 45’S, 143° ll’E, Mar 1996, Sankowsky

1505 (QRS); loc. cit., Oct 1996, Sankowsky 1524

(QRS); loc. cit., Apr 1997, Sankowsky 1557 &

Sankowsky (BRI); loc. cit., Nov 1997, Sankowsky

1614 (QRS); loc. cit.. May 1997, Sankowsky s.n.

(BRI [AQ654612]); Claudie River, 12° 45’S, 143°

15’E, Nov 1977, Stocker 1625 (CANB, QRS); Iron

Range, Jul 1963, Volck AF02647 (QRS).

Distribution and habitat : Actephila

longipedicellata has been collected in three

localities (Garraway Creek and Tozer Gap near

Iron Range, and Kennedy Hill) within a single

1° grid cell on far north Cape York Peninsula,

Queensland (Map 5). Plants grow in the

understorey of ‘seasonally dry’ rainforest

(evergreen complex notophyll vineforest) in

rocky situations (along creeks or on ridges),

with granite boulders in much evidence at

altitudes between 20 and 240 m.

Notes : Actephila longipedicellata was first

collected by Brass in 1948 and collections made

prior to 1999 were usually identified as A.

lindleyi. Actephila longipedicellata is similar

on morphological grounds (short petioles, long

pedicels) to A. sessilifolia , but differs from that

species in the leaf lamina with an acute to

cuneate base (versus cordate to subcordate,

often appearing stem clasping due to short

petiole); the greater number of lateral veins (11-

15 versus 8-11); the male flowers with longer

pedicels (25-34 mm versus 8-11 mm), larger

sepals (3.5-5 x 2.5^1 mm versus 1.8-2.5 x 1.5-

1.8 mm), spathulate petals (versus obovate to

oblong) that are larger (2.8-3.5 x 1.3-2.5 mm

versus 0.8-1.8 x 0.3-1 mm) and a disk (versus

annular) that is twice as large (4.5-5 mm diameter

versus 2-2.8 mm).

status: Actephila

Austrobaileya 7 (1): 57-98 (2005)

longipedicellata has a restricted distribution

with perhaps as many as three populations. The

area of occupancy and the number of individuals

within populations are unknown. One

population is within Iron Range National Park.

Further survey work is required to determine

whether this species is deserved of

conservation listing. The population at

Garraway Creek is under localised threat from

destruction of vegetation by tourists and local

inhabitats that use the area for short-term

recreational camping.

Etymology: The specific epithet is derived from

the Latin words longus (long) and pedicellus

(pedicel) and refers to the consistently long

flower pedicels in this species.

9. Actephila petiolaris Benth., FI. Austral. 6:90

(1873). Type: Queensland. “Rockingham

Bay”, Dallachy s.n. (holo: K [photo at

BRI!]).

Subshrub, shrub or small tree to 8 m high.

Indumentum of simple trichomes, fawn or

ferruginous, entire plant is glabrous unless

stated otherwise. Branchlets glossy, fawn-tan

to olive-green, glabrous or with sparse

indumentum when young, lenticellate with age.

Stipules triangular, 1.2-2 mm long, 0.8-1.5 mm

wide. Leaves alternate, coriaceous, petiolate;

petioles 9-80 mm long, 1-1.5 mm wide, olive

green, glabrous or with sparse indumentum,

glabrescent and occasionally lenticellate with

age; lamina length/petiole length ratio 2-8.7

(n=106); lamina elliptic-ovate, ovate or obovate,

± flat, 48-190 mm long, 25-90 mm wide, length/

width ratio 1.5-3.3 (n=106); base cuneate,

obtuse or rounded; tip acute, abruptly acute,

acuminate or caudate; midrib above ± flat, dark

green, below prominently raised at lamina base

(height > width) then decreasing towards apex,

pale olive-green; venation brochidodromous,

comprising 9-12 lateral veins per side of midrib

and intercostal reticulate veins; upper surface

dark green and glossy, lateral venation barely

visible, intercostal venation obscure; lower

surface olive green and glossy, lateral venation

prominent and slightly raised, intercostal

venation visible but not raised, 1^1° vein orders

distinct, 5° vein order indistinct, 6° and onwards

vein orders obscure; margin entire, flat, olive-

Conservation

Forster, Revision of Actephilia

green. Inflorescence an axillary fascicle 2-4 mm

diameter, single sex or with both male and female

flowers. Male flowers: pedicels stout, 3-7 mm

long, 0.6-0.8 mm diameter; sepals 5, oblong to

oblong-ovate, 3-4 mm long, 1.3-2.8 mm wide,

apices glabrous or sparsely ciliate; petals

absent, or rarely one or two present,

oblanceolate, 1.8-2.5 mm long, 0.5-1.2 mm wide,

irregularly lacerate at tip; disk conspicuous,

fleshy, 0.8-1.2 mm high, 3.5-6 mm diameter, 5-

angled; stamens 4 or 5, filaments 2-2.5 mm long,

c. 0.2 mm diameter, anthers globose-ovoid, 0.2-

0.3 mm long, 0.4—0.5 mm wide. Female flowers:

pedicels filiform, 9-35 mm long, 0.5-0.8 mm

diameter; sepals 5, oblong to oblong-ovate,

1.8-3.2 mm long, 1.8-2 mm wide, apices densely

ciliate, occasionally with sparse indumentum

down centre of sepal externally; petals absent

or rarely one or two present, oblanceolate, 1.2-

2 mm long, 0.3-0.5 mm wide; disk conspicuous,

fleshy, 1-2 mm high, 3^1.5 mm diameter, 5-

angled; ovary 2-4 mm long, 2-5 mm diameter;

styles 3, simple, shortly connate at base, 1-1.8

mm long, stigmas subcapitate. Fruit: pedicels

23-30 mm long, c. 1.5 mm diameter; capsules

depressed-globose, rounded, 10-20 mm long,

15-27 mm diameter, strongly bullate, green;

columella 7-9 mm long. Seeds 6-12 mm long, 5-

9 mm wide, olive green; hilum 1-2.5 mm long,

caruncle poorly developed, narrow-oblong, 1-

4 mm long, c. 0.2 mm wide, cream. Fig. 8.

Notes : There is significant variation in the

morphology of the collections referred here to

A. petiolaris , notably the length of the petiole

relevant to the length of the leaf lamina, leaf

shape and indumentum colour and density on

young foliage. This variation is geographically

and ecologically based and is formalised by the

recognition of subspecies.

Etymology : The specific epithet is formed from

the Latin word petiolus (petiole) and alludes to

the long petiole of this species.

Key to subspecies of Actephilapetiolaris

1 Foliage glabrous or with scattered fawn indumentum when young; leaf lamina

length/petiole length ratio 2-7.4. A. petiolaris subsp. petiolaris

1: Foliage with sparse ferruginous-brown indumentum when young; leaf lamina

length/petiole ratio 5-8.7.A. petiolaris subsp. jagonis

9a. Actephila petiolaris subsp. petiolaris

Shrub or small tree to 8 m high. Indumentum

absent, or fawn. Leaf glabrous or with scattered

indumentum when young; leaf lamina length/

petiole length ratio 2-7.4 (n=79); lamina elliptic-

ovate, rarely obovate.

Additional specimens examined: Queensland. Cook

District: Cedar Bay, 15° 47’S, 145° 21’E, Jan 1973,

Dick sub Webb & Tracey 13774 (BRI); Churchill Creek,

Churchill L.A., S.F. 143, 16° 34’S, 145° 19’E, Jul 1995,

Forster PIF17208 & Figg (BRI, QRS); cult. Tolga (ex

Churchill Creek near Julatten), 16° 34’S, 145° 19’E,

Jan 2002, Forster P1F28205, P1F28206 (BRI); S.F.R.

675, Mulgrave L.A., 17° 05’S, 145° 42’E, Mar 1980,

Gray 1665, 1668 (BRI, QRS); S.F.R. 675, Little

Mulgrave Valley, 17° 06’S, 145° 41’E, Jun 1996, Gray

6778 (QRS); S.F.R. 675, 17° 05’S, 145° 40’E, Nov

1976, Hyland 3481RFK (QRS), 3482RFK (QRS),

3483RFK (CANB, QRS); S.F.R. 675 Grafton, Mulgrave

L.A., 17° 05’S, 145° 42’E, Nov 1988, Hyland

25619RFK (QRS); S.F.R. 675, East Mulgrave L.A.,

17° 05’S, 145° 40’E, Dec 1976, Hyland 9249 (QRS);

S.F.R. 675 Grafton, Mulgrave L.A., 17° 05’S, 145°

42’E, Nov 1988, Hyland 13719 (CANB, QRS); Cooper

Creek, east of Turpentine road, 16° 09’S, 145° 24’E,

Jun 2001, Jago 6004 (BRI); Coopers Creek, via

Daintree, Jun 1969, Mazlin AF04347 (QRS); cult.

Tolga (ex Churchill Creek), Nov 1999, Sankowsky

1680, 1681 & Sankowsky (QRS); loc. cit., Dec 2005,

Sankowsky 2554 & Sankowsky (BRI); cult. Tolga (ex

S.F.R. 675 Grafton, Mulgrave L.A., 17° 05’S, 145°

42’E), Dec 2004, Sankowsky 2555 & Sankowsky (BRI);

Coopers Creek, Cape Tribulation, 16° 10’S, 145° 25’E,

Oct 1997, Small 14 (QRS); Gap Creek, c. 8 miles [13.3

km] back from Ayton, May 1969, Smith 14444 (BRI);

Bloomfield, May 1969, VolckAF04311 (QRS).

Distribution and habitat : Actephila petiolaris

subsp. petiolaris is endemic to the ‘Wet Tropics’

Austrobaileya 7 (1): 57-98 (2005)

Fig. 8. Actephila petiolaris subsp. petiolaris. A. habit of flowering branch x0.5. B. abaxial leaf surface x0.5. C.

side view of male flower x6. D. face view of male flower x6. E. side view of female flower x6. E face view of

female flower x6. G. side view of fruit *2. H. face view of fruit x2. A-D from Forster PIF28205 (BRI); E, F from

Forster PIF28206 (BRI); G, H from Forster PIF17208 (BRI). Del. W. Smith.

bioregion of north-eastern Queensland with a

northern limit at Gap Creek and a southern limit

in the Little Mulgrave Valley (Map 3). Despite

the different labelling of collections, there are

only four localities known, viz. Cedar Bay -

Bloomfield - Gap Creek, Churchill Creek, Cooper

Creek and S.F.R. 675. Plants occur at altitudes

between 20-300 m in the understorey of ‘wet’

rainforest (evergreen, complex mesophyll or

notophyll vineforest) on alluvium derived from

metamorphics along watercourses at altitudes

between 100 and 440 m.

Notes : A number of collections are provisionally

placed here, notably those from Cooper Creek

north of the Daintree River. These particular

collections (Mazlin AF04347, Jago 6004 and

Small 14) have particularly large leaves with

caudate leaf apices and show some similarities

to A. foetida. They lack the indumentum and

pattern of interlateral venation that is present

in A. foetida , but share its habit (small shrub)

and leaf size variation. Leaf size appears to be

highly variable in A. petiolaris subsp.

petiolaris , but this is not particularly unusual

in understorey trees and shrubs that are

subjected to variable regimes of light.

Conservation status: Actephila petiolaris

subsp. petiolaris is present in several National

Parks (Cedar Bay, Daintree) and two State

Forests (143, 675). There is no information

available as to the area of occupancy or the

number of individuals within populations. It is

not considered to be under threat.

Forster, Revision of Actephilia

9b. Actephila petiolaris subsp. jagonis

P.I.Forst., subsp. nov. a forma typica

Actephilae petiolaris Benth.

frondescentia juvenile vestita indumento

sparso ferrugineo (adversum glabram vel

indumentum dissitum hinnuleum

juvenile), lamina obovata rarius elliptico-

ovata (adversum ovatam rarius obovatam)

et folii ratione longitudinis laminae ad

longitudine petiolo majore (5-8.7 non 2-

7.4) distinguitur.

Typus: Queensland. Cook District: on track from

Power Station, to Mt Surprise Creek [Barron

Gorge], 16° 51’S, 145°38’E, 23 January 1993,

R.L. Jago 2030 (holo: BRI [1 sheet]).

Subshrub to 2 m tall. Indumentum ferruginous-

brown. Leaf with sparse indumentum when

young; leaf lamina length/ petiole length ratio

5-8.7(n=27); lamina obovate, rarely elliptic-

ovate.

Additional specimens examined : Queensland. Cook

District: Barron Gorge, 16° 50’ S, 145° 38’E, Feb 1994,

Cooper WWC797 & Cooper (QRS); Barron Gorge

National Park, 16° SI’S, 145° 38’E, Feb 1996, Forster

PIF18644 & Jago (BRI, QRS); N.P.R. 880, Parish of

Cairns [Barron Gorge], 16° 5l’S, 145° 38’E, Apr 1994,

Hyland 15085 (QRS).

Distribution and habitat : Actephila petiolaris

subsp. jagonis is known only from the Barron

Gorge National Park near Cairns in the ‘Wet

Tropics’ of north-eastern Queensland (Map 3).

This locality is within the overall geographical

range of A. petiolaris but is disjunct from the

populations of A. petiolaris subsp. petiolaris

and occurs in a different and much ‘drier’

rainforest type. Plants occur on steep slopes in

semi-deciduous microphyll vineforest on

substrates derived from metamorphics at

altitudes between 100 and 330 m.

Notes : This taxon is included as a subspecies

of A. petiolaris with some reservations. Flowers

of this plant have not been seen and it may well

prove to be specifically distinct from A.

petiolaris subsp. petiolaris that is a larger,

generally glabrous plant, with generally longer

leaf petioles and leaf laminas that are nearly

always elliptic-ovate (versus nearly always

obovate). The type collection of A. petiolaris

subsp .jagonis has a single, badly mangled fruit.

This fruit appears smaller than those found on

A. petiolaris subsp. petiolaris , but until better

material is obtained, the uniqueness of the size

disparity cannot be determined.

Conservation status : The only known

population of this taxon is within Barron Gorge

National Park. Survey of the population is

required to determine the area of occupancy,

the number of plants that are present and any

threats that they may face. Due to its occurrence

in a single population this subspecies is

probably deserved of conservation listing.

Etymology: Actephila petiolaris subsp.

jagonis is named to honour R.L. (Bob) Jago of

Cairns, plant collector and amateur botanist

extraordinaire. Bob has contributed over 2000

herbarium collections to the Queensland

Herbarium and was the first to collect this plant.

10. Actephila plicata P.I.Forst., sp. nov. ab A.

latifolia Benth. folii lamina plicata non

plana margine revoluto non piano, floribus

foemineis sepalis longioribus (2.8-3.5 mm

adversum 2-2.5 mm), petalis oblongis

usque oblanceolatis (in ilia obovatis) et

majoribus (2.5-3 x c. 0.8 mm adversum 1-

2 x 0.5-1 mm) et frutice 5-lobato (in ilia

rotundato) differt.

Typus: Queensland. North Kennedy District:

Mt Dryander, left branch Dryander Creek, 15

km N of Proserpine, 20° 16’S, 148° 34’E, 6

February 2004, P.I. Forster PIF29943, R. Jensen

&M.C. Tucker (holo: BRI [1 sheet + spirit]; iso:

MEL, NSW).

Shrub or small tree to 6 m high. Indumentum of

simple trichomes, uncoloured, entire plant is

glabrous unless stated otherwise. Branchlets

glossy, brown, lenticellate with age. Stipules

ovate to triangular-ovate, 1-1.8 mm long, 1-1.8

mm wide, irregularly crenate. Leaves alternate

or subverticillate, coriaceous, petiolate; petioles

3-25 mm long, 1.5-2.5 mm wide, olive-green,

lenticellate; lamina length/petiole length ratio

6-20 (n=l 10); lamina elliptic to obovate, plicate,

37-207 mm long, 20-75 mm wide, length/width

ratio 1.6-3.3 (n=l 10); base acute to cuneate; tip

acute to rounded; midrib above ± flat, dark green,

below prominently raised at lamina base (height

> width) then decreasing towards apex, pale

olive-green; venation brochidodromous,

comprising 10-14 lateral veins per side of midrib

Austrobaileya 7 (1): 57-98 (2005)

Fig. 9. Actephila plicata. A. abaxial leaf surface x0.5. B. side view of fruit *1.5. C. face view of fruit xl.5. D.

ventral view of seed x3. A, D from Cooper WWC1576 (BRI); B, C from Forster PIF29943 (BRI). Del. W. Smith.

and intercostal reticulate veins; upper surface

dark green and glossy, lateral and intercostal

venation visible; lower surface pale olive-green

and matt, lateral and intercostal venation

prominent with laterals raised, 1-3° vein orders

distinct, 4° vein order indistinct, 5° and onwards

vein orders obscure; margin entire, revolute, pale

olive-green. Inflorescence an axillary fascicle 2-

8 mm diameter, rarely interpetiolar, single sex or

with both male and female flowers. Male flowers

not seen. Female flowers: pedicels stout, 8-10

mm long, 1-1.5 mm diameter; sepals 5, obovate,

2.8-3.5 mm long, 1.2-2.2 mm wide, ciliate in patch

at the apices; petals absent or one or two

present, oblong or oblanceolate, 2.5-3 mm long,

c. 0.8 mm wide; disk conspicuous, fleshy, 1—1.5

mm high, 4-5.5 mm diameter, 5-angled; ovary,

styles and stigmas not seen. Fruit: pedicels 6-

10 mm long, 1-1.5 mm diameter; capsules

depressed-globose, strongly pentalobate, 12-

14 mm long, 15-19 mm diameter, strongly bullate,

green; columella 8-12 mm long. Seeds 7-11 mm

long, 5-8.5 mm wide, pale tan and fawn to olive-

green and tan; hilum 2-5 mm long, caruncle

narrow oblong, 2-3 mm long, 0.5-1 mm wide,

cream. Fig. 9.

Additional specimens examined: Queensland. North

Kennedy District: S.F. 387, 2 km SSW of dam wall on

Proserpine River, 20° 23’S, 148° 22’E, May 1991,

Forster PIF8318 & McDonald (BRI, K, L, MEL, QRS);

Mt Dryander S.F., Dryander Creek left branch, 20°

16’S, 148° 34’E, Oct 1999, Forster PIF24997 & Booth

(BRI, MEL, QRS); headwaters of Dryander Creek, Mt

Dryander, 20° 15’S, 148° 35’E, Oct 1969, Webb &

Tracey 10040 (BRI). South Kennedy District: Scawfell

Island, 20° 53’S, 149° 37’E, Nov 1986, Batianoff 6175

& Hegerl (AD, BRI); Scawfell Island National Park,

20° 53’S, 149° 37’E, Nov 1986, Batianoff 6303 (BRI);

Dugong Inlet, Whitsunday Island, 20° 15’S, 148° 57’E,

Sep 1990, Batianoff 900971 & Batianoff ( BRI); Cape

Hillsborough National Park, 20° 15’S, 149° 02’E, Aug

1992, Batianoff 920889 (BRI); Puritan Bay, Conway

National Park, 20° 28’S, 148° 53’E, May 1994,

Batianoff 94053 & Dillewaard (BRI); Keswick Island,

Connie Bay, 20° 54’S, 149° 24’E, Sep 1996, Batianoff

9609111 & Champion (BRI); Cape Hillsborough N.P,

along track to Hidden Valley, 20° 55’S, 149° 02’E, Sep

2001, Cooper WWC1576 & Cooper (BRI); R299

Conway, Repulse Bay, Jun 1981, Dansie AFO5230

(QRS).

Notes: Male flowers have not been examined

for this species. There are morphological

obvious similarities between A. plicata and

A. latifolia that occurs further north; however,

the former differs from A. latifolia in the leaf

lamina being plicate (versus flat) with a revolute

margin (versus flat); the female flowers with

longer sepals (2.8-3.5 mm versus 2-2.5 mm),

petals that are oblong to oblanceolate (versus

obovate) and larger (2-5-3 x c . 0.8 mm versus

1-2 x 0.5-1 mm) and the strongly pentalobate

fruit (versus rounded).

Distribution and habitat: Actephila plicata is

endemic to central Queensland with a northern

limit at Mt Dryander and a southern limit at Cape

Hillsborough National Park where it occurs in

coastal areas of the mainland and on offshore

continental islands (Map 5). Plants occur in ‘dry’

rainforest and vinethicket (littoral microphyll

vineforest, semi-deciduous, complex notophyll

vineforest) on granite substrates from near sea

level to 100 m altitude.

Forster, Revision of Actephilia

Conservation status : Actephila plicata is

present in several National Parks (Cape

Hillsborough, Conway, Dryander, Scawfell

Island) and two State Forests (387,432). There

is no information available as to the area of

occupancy or the number of individuals within

populations. It is not considered under threat.

Etymology : The specific epithet is formed from

the Latin word plicatus (folded) and alludes to

the appearance of the leaf lamina.

11. Actephila sessilifolia Benth., FI. Austral. 6:

90 (1873). Type: Queensland. Port Curtis

District: [“Shrub of 4-6 ft at the Caves

Mountain, 5 miles west of Morinish”],

Thozets.n. (lecto: MEL; isolecto: K [with

no locality data, photo at BRl],Jide Airy

Shaw (1980b: 217).

Illustrations : Calvert etal. (2005:12).

Subshrub or shrub to 6 m high. Indumentum of

simple trichomes, uncoloured or ferruginous,

entire plant is glabrous unless stated otherwise.

Branchlets glossy, fawn to ferruginous brown,

lenticellate with age. Stipules triangular, 1-1.2

mm long, 0.8-1 mm wide, occasionally ciliate.

Leaves alternate or subverticillate, coriaceous,

petiolate, appearing sessile due to shape of

lamina base; petioles 1-7 mm long, 1-2 mm wide,

grey-fawn, lenticellate; lamina length/petiole

length ratio 12.5-125 (n=222); lamina elliptic,

obovate or oblanceolate, rarely orbicular, ± flat,

19-152 mm long, 9-60 mm wide, length/width

ratio 1.5-3.6 (n=222); base cordate to

subcordate, often falsely stem clasping due to

short petiole; tip acute to rounded, rarely

caudate or retuse; midrib above ± flat or slightly

raised (height < 5 times the width) pale green,

below slightly raised at lamina base (height ±

equal to width) then decreasing towards apex,

pale olive-green; venation brochidodromous,

comprising 8-11 lateral veins per side of midrib

and intercostal reticulate veins; upper surface

dark green and glossy, lateral venation visible,

intercostal venation weakly visible to ± obscure;

lower surface pale olive-green and glossy, lateral

and intercostal venation visible, but weakly

raised, 1° and 2° vein orders distinct, 3° vein

order indistinct, 4° and onwards vein orders

obscure; margin entire, weakly revolute, yellow-

green. Inflorescence an axillary fascicle 1-2 mm

diameter, single sex or with both male and female

flowers. Male flowers: pedicels filiform, 6-8 mm

long, 0.3-0.5 mm diameter; sepals 5, obovate to

ovate, 1.8-2 mm long, 1.5-1.8 mm wide, ciliate

on upper margin; petals absent or 1-5, oblong

to obovate, 0.8-1.8 mm long, 0.3-1 mm wide;

disk conspicuous, fleshy, 0.4-0.5 mm high, 2-

2.8 mm diameter, annular; stamens 5, filaments

1-1.2 mm long, c. 0.2 mm diameter, anthers

globose-ovoid, 0.3-04 mm long, 0.4-0.5 mm

wide. Female flowers: pedicels filiform, 20-32

mm long, 0.1-0.2 mm diameter; sepals 5, obtuse

to ovate, 2.1—4 mm long, 1.2-2.8 mm wide,

glabrous or ciliate towards the apices; petals

1-5, oblanceolate, oblong or obovate, 1.2-1.4

mm long, 0.3-1 mm wide; disk conspicuous,

fleshy, 0.8-1 mm high, 2.8-4 mm diameter, 5-

angled; ovary 2-2.5 mm long and 2-4 mm

diameter; styles 3, simple, shortly connate at

base, 1.5-1.8 mm long, stigmas subcapitate.

Fruit: pedicels 22-36 mm long, c. 0.5 mm

diameter; capsules depressed-globose to

subglobose, rounded, 9-11 mm long, 13-16 mm

diameter, weakly to strongly bullate, green to

green-black; columella 4-7 mm long. Seeds 6-9

mm long, 4.5-8 mm wide, olive green or olive

Fig. 10. Actephila sessilifolia. A. abaxial leaf surface

x0.25. B. side view of fruit xl.5. C. face view of

dehiscing fruit xl.5. D. ventral view of seed x3. All

from Forster PIF8113 (BRI). Del. W. Smith

green and dark green in irregular patterns; hilum

2-3.5 mm long, caruncle narrow-oblong, 1.5-5

mm long, 0.5-1 mm wide, cream. Fig. 10.

Selected specimens examined : Queensland. North

Kennedy District: Emmett Creek, Bowling Green Bay

National Park, S of Townsville, Aug 1991, Bean 3605

(BRI); Mt Burrumbush, Bowling Green Bay National

Park, S of Townsville, 19° 28’S, 147° 05’E, Aug 1991,

Bean 3658 (BRI); Lower slopes of Mt Quandong, c. 21

km SW of Proserpine, 20° 29’S, 148° 24’E, May 2002,

Champion 1843 et at (BRI); Emmett Creek, Saddle

Mountain, 19° 26’S, 147° 02’E, Jul 1995, Camming

13611 (BRI); Mt Cataract, W of Townsville, 19° 17’S,

146° 30’E, Oct 1996, Camming 15223 (BRI); Mt

Quandong, Proserpine Valley, May 1982, Dansie

AF05173 (QRS); Seaview Range, Apr 1947, Flecker

N.Q.N.C. 10880 (QRS); N.P.R. 629, Paluma Range,

Rollingstone Creek (West Branch) off Pace road, 19°

04’S, 146° 21’E, Dec 2003, Ford et al. 4222 (QRS);

Dryander Creek left branch, SE slope of Mt Dryander,

20° 16’S, 148° 34’E, Jun 1989, Forster PIF5127 et al.

(BRI); Elliott Toe, Bowling Green Bay National Park,

9 km NNE of Woodstock, 19° 31’S, 146° 52’E, May

1991, Forster PIF8361 & Bean (BRI, QRS); lower

western slopes of Mt Dryander, 20° 15’S, 148° 33’E,

Jul 1974, Moriarty 1866 (QRS); Gregory Creek, 20°

16’S, 148° 36’E, Perry s.n. (BRI [AQ566535]). South

Kennedy District: N of Pine Mt, 8 km NNW of

Marlborough, 22° 45’S, 149° 51’E, Jul 1997, Fensham

3282 (BRI); T.R. 179 Kalvin, 6 km W of Koumala,

21° 36’S, 149° ll’E, Apr 1991, Forster PIF8032 &

McDonald (BRI, MEL, QRS); T.R. 179 Kalvin, 6 km

WSW of Koumala, 21° 37’S, 149° 10’E, Apr 1991,

Forster PIF8045 & McDonald (BRI); Coffee Creek,

NE base of Mt Jukes, 20° 59’S, 148° 57’E, Apr 1991,

Forster PIF8113 & McDonald (BRI); Mt Adder,

southern base of Mt Jukes, 20° 59’S, 148° 55’E, Apr

1991, Forster PIF8118 & McDonald (BRI, MEL, QRS);

S.F. 658 Carawatha, 20° 47’S, 148° 34’E, Apr 1991,

Forster PIF8198 & Tucker (BRI, K, MEL, QRS); S.F.

658 Carawatha, 20° 47’S, 148° 34’E, Jun 1995, Forster

PIF16724 & Tucker (BRI, QRS); Headwaters of Coffee

Creek, eastern side of Mt Jukes (N.P 616 Ossa), 21°

02’S, 148° 55’E, Nov 1989, McDonald 4401 et al.

(BRI); St Helen’s Gap, between Mirani & Calen, 21°

00’S, 148° 42’E, Nov 1989, McDonald 4510 et al.

(BRI, QRS); headwaters of Cameron Creek, E foothills

of Clarke Range, c. 6 km NW of Koumala, 21° 34’S,

149° ll’E, Jun 1994, McDonald 5983 & Champion

(BRI). Leichhardt District: S.F. 82 Connors, headwaters

of East Funnel Creek, 21° 34’S, 149° ll’E, Jun 1994,

McDonald 5976 & Champion (BRI). Port Curtis

District: S.F. 878, W of Rockhampton, 23° 18’S, 150°

09’E, Jul 1993, Fensham 1080 (BRI); Mt Larcom, 5

km NW of Yarwun, 23° 48’S, 151° 05’E, Jan 1994,

Forster PIF14646 (BRI); Curtis Island, 23° 32’S, 151°

06’E, Sep 1994, Thomas s.n. (BRI [AQ582433],

NSW); 25 km NNW of Gladstone, 23° 42’S, 151° 03’E,

Feb 1997, Thompson GLA13 & Turpin (BRI); Rundle

Range State Forest, c. 10 miles [16.7 km] SE of Port

Alma near O’Connor Creek, near mouth of Fitzroy

Austrobaileya 7 (1): 57-98 (2005)

River, 23° 40’S, 150° 59’E, May 1971, Webb & Tracey

10462A (BRI, MEL). Burnett District: McLaughlin’s

road, Yarrol Scrub, 21 km E of Monto, 24° 53’S, 151°

19’E, Dec 2004, Forster PIF30482 & Beard (A, BRI,

L, MEL, MO, NE, NSW). Wide Bay District: Kirbys

Road, Bundaberg, 24° 49’S, 152° 23’E, Sep 1999,

Forster PIF24923 & Schmitt (BRI).

Distribution and habitat: This species is much

more widespread than thought by Airy Shaw

(1980b) who examined material from only three

localities. Actephila sessilifolia has a known

range from the Paluma Range, west of

Townsville, south to Yarrol near Monto (Map

3). Plants are usually locally common in the

understorey of‘dry’ rainforest (semi-deciduous,

complex to simple, mesophyll to notophyll

vineforest) or more rarely vinethickets (semi¬

evergreen vinethicket), on alluvium or rocky

situations, on substrates derived from basalt,

granite or metasediments at altitudes between

30 and 490 m.

Notes: Due to the cordate nature of the leaf

base in Actephila sessilifolia , the shortly

petiolate leaves give the impression that the

leaves are sessile. This character combination

is unique within Actephila. Although this

character state is approached in some

specimens of A. longipedicellata, this latter

species does not have a cordate base to the

leaf lamina

Conservation status: Actephila sessilifolia is

present in Bowling Green Bay, Mt O’Connell,

Pioneer Peaks and Rundle Range National Parks

and State Forests 55, 82,179,207,658 and 878.

Many of the populations are small and

fragmented due to land clearing, fire incursions

and competition from weeds. The populations

in the southern part of the known distribution

range are small and not in conservation

reserves. There is no information available as

to the area of occupancy or the numbers of

individuals within populations. This species is

not considered threatened although it is

currently listed as Rare under the NCA.

Etymology: The specific epithet is fonned from

the Latin words sessili (sessile, stalkless) and-

folius (leaves) and alludes to the superficial

appearance of the foliage.

Forster, Revision of Actephilia

12. Actephila traceyi P.I.Forst., sp. nov. ab

A. lindleyi (Steud.) Airy Shaw folii lamina

numero majore venarum lateralium (11-16

non 7-12), floribus masculis pedicellis

longissimis (5-16 non 1.5-5 mm), sepalis

ovato-spathulatis (adversum sepala

oblonga usque ovata) et longioribus (3.5-

5 non 1-3 mm), disco 5-angulato (in ilia

rotundato), frutice majore (11—15 x 16-24

mm adversum 9-12 x 11-13.5 mm) et

seminibus majoribus (10-14 x 7-12

adversum 6-7 x 5-6.5) pallide olivaceis

usque olivaceis (non atro-olivaceis)

differt.

Typus: Queensland. Cook District: Leo Creek

area, Timber Reserve 14, Mcllwraith Range, 13°

45’S, 143° 23’E, 21 June 1995, P.I.Forster

PIF16894 (holo: BRI [1 sheet + spirit]; iso:

MEL).

Actephila sp. (Rocky River GC.Stocker 1042)

(Forster & Henderson 1997: 70; Forster &

Halford 2002:68).

Subshrub or shrub to 3 m high. Indumentum of

simple trichomes, uncoloured, entire plant is

glabrous unless stated otherwise. Branchlets

glossy, fawn-tan, lenticellate with age. Stipules

narrow-triangular to triangular, 0.5-1.8 mm long,

0.8-1 mm wide. Leaves alternate, subopposite

or subverticillate, coriaceous, petiolate; petioles

3-23 mm long, 1-1.5 mm wide, deeply grooved

on top with distinct rim to 0.5 mm high, yellow-

green, lenticellate with age; lamina length/petiole

length ratio 6.2-18.6 (n=128); lamina obovate

to oblanceolate, rarely elliptic-ovate, ± flat, 55-

170 mm long, 23-66 mm wide, length/width ratio

1.7-3.2 (n=128); base acute to cuneate; tip acute

to acuminate, rarely rounded; midrib above

slightly raised (height < 5 times the width),

yellow-green, below strongly raised at lamina

base (height ± equal to width) then decreasing

towards apex, yellow; venation brochido-

dromous, comprising 11-16 lateral veins per

side of midrib and intercostal reticulate veins;

upper surface dark green and glossy, lateral and

intercostal venation visible; lower surface pale

yellow-green and glossy, lateral and intercostal

venation prominent, slightly raised, 1-3° vein

orders distinct, 4° vein order indistinct, 5° and

onwards vein orders obscure; margin entire, flat,

yellow-green. Inflorescence an axillary fascicle

2-4 mm diameter, single sex or with both male

and female flowers. Male flowers: pedicels

filiform, 5-16 mm long, 0.6-1.2 mm diameter;

sepals 5, ovate-spathulate, 3.5-5 mm long, 2—4

mm wide, ciliate in dense apical patch and with

occasional marginal cilia; petals absent or 1,

narrow-oblong to oblong-spathulate, 1-2 mm

long, 0.2-0.8 mm wide; disk conspicuous,

fleshy, 1-1.8 mm high, 4-6 mm diameter, 5-

angled; stamens 5, filaments 2.5-3.8 mm long,

0.2-0.3 mm diameter, anthers globose-ovoid,

0.4-0.8 mm long, 0.2-0.5 mm wide. Female

flowers: pedicels filiform, 10-22 mm long, 0.7-1

mm diameter; sepals 5, spathulate, 3.5^1 mm

long, 2.2-3 mm wide, glabrous; petals absent;

disk conspicuous, fleshy, 1.5-2 mm high, 5-6

mm diameter, annular; ovary not seen; styles 3,

simple, shortly connate at base, c. 2 mm long,

stigmas subcapitate. Fruit: pedicels 15-35 mm

long, 0.7-1 mm diameter; capsules depressed-

Fig. 11. Actephila traceyi. A. abaxial leaf surface *0.5.

B. side view of male flower *6. C. face view of male

flower x3. D. ventral view of seed x2. A from Forster

PIF10607 (BRI); B, C from Tucker 3006 (BRI); D

from Stocker 1042 (BRI). Del. W. Smith.

globose, rounded, 11-15 mm long, 16-24 mm

diameter, strongly bullate, green; columella 7-

11 mm long. Seeds 10-14 mm long, 7-12 mm

wide, pale olive tan to olive green; hilum 3-5

mm long, caruncle narrow-oblong, 2-4 mm long,

c. 0.5 mm wide, cream. Fig. 11.

Additional specimens examined : Queensland. Cook

District: Between Chester River & Leo Creek, 13° 39’S,

143° 26’E, Jul 1978, Butler 299 (CANB); Gorge of the

Chester River, eastern fall of Mcllwraith Range, 13°

41’S, 143° 24’E, Jul 1978, Clarkson 2349 (BRI); Rocky

River, 13° 49’S, 143° 28’E, Aug 1997, Cooper WWC56

& Jensen (QRS); T.R. 14, Leo Creek Mine area,

Mcllwraith Range, 13° 44’S, 143° 22’E, Jun 1992,

Forster PIF10164 et al. (BRI, QRS); Rocky River Scrub,

Silver Plains Station, 13° 49’S, 143° 27’E, Jun 1992,

Forster PIF10607 & Tucker (BRI, MEL, QRS); Lower

Station Creek Gorge, Mcllwraith Range, 13° 57’S, 143°

19’E, Jun 1997, Forster PIF21281 et al. (BRI, MEL,

NE); Rocky River, 13° 55’S, 143° 30’E, Sep 1971,

Hyland 5477 (BRI, QRS); T.R. 14 Massy, 13° 52’S,

143° 23’E, Nov 1980, Hyland 10899 (BRI, CANB,

QRS); loc. cit., Nov 1980, Hyland 10899 (BRI, QRS);

Mcllwraith Range, near the head of South Massey Creek,

13° 54’S, 143° 19’E, Le Cussan 227 (QRS); cult. Tolga

(ex Leo Creek, track to old mine, 13° 44’S, 143° 22’E),

Dec 2004, Sankowsky 2556 & Sankowsky (BRI); cult.

Tolga (ex Rocky River, 13° 49’S, 143° 27’E), Dec

2004, Sankowsky 2558 & Sankowsky (BRI);

Mcllwraith Range, Oct 1962, Smith 11789 (BRI); T.R.

14, Rocky River area, 13° 40’S, 143° 25’E, Sep 1973,

Stocker 1042 (BRI, QRS); cult. Kholo (ex Leo Creek,

Mcllwraith Range, 13° 44’S, 143° 22’E), Jan 2004,

Tucker 3006 (BRI); cult. Kholo (ex Rocky River, Silver

Plains, 13° 49’S, 143° 27’E), Jan 2004, Tucker 4375

(BRI); T.R. 14, Mcllwraith Range, 13° 45’S, 143° 20’E,

Jul 1977, Unwin 383 (QRS); Goanna Creek, E of

Mcllwraith Range, Nov 1956, Webb 3155 (BRI); Rocky

River on eastern foothills of Mcllwraith Range, 13°

47’S, 143° 25’E, Oct 1969, Webb & Tracey 9341 &

9438 (BRI); headwaters of Lankelly Creek on western

fall of Mcllwraith Range, 13° 52’S, 143° 20’E, Oct

1969, Webb & Tracey 9554 (BRI); North western fall

of Mcllwraith Range at head of Peach Creek, 13° 46’S,

143° 16’E, Oct 1969, Webb & Tracey 9817 (BRI);

Lankelly Creek on western fall of Mcllwraith Range,

13° 54’S, 143° 20’E, Jun 1971, Webb & Tracey 11487

(BRI).

Distribution and habitat : Actephila traceyi is

restricted to the Mcllwraith Range on far

northern Cape York Peninsula, where it has been

found in the catchments of Goanna, Leo,

Massey, Peach and Station Creeks and the

Chester and Rocky Rivers (Map 5). Plants grow

Austrobaileya 7 (1): 57-98 (2005)

at altitudes between 100 and 380 m in ‘ seasonally

dry’ rainforest (semi-deciduous, complex

notophyll to mesophyll vineforest) along

watercourses on alluvium derived from granites

or metamorphics or a combination of the two

substrates.

Notes : Herbarium collections of Actephila

traceyi have been usually identified as

A. lindleyi: ; however, it differs from that species

in the leaf lamina with a greater number of lateral

veins (11-16 versus 7-12); the male flowers with

much longer pedicels (5-16 versus 1.5-5 mm),

sepals that are ovate-spathulate (versus oblong

to ovate) and longer (3.5-5 versus 1-3 mm), a

disk that is 5-angled (versus rounded); larger

fruit (11—15 x 16-24 mm versus 9-12 x 11-13.5

mm); and larger seeds (10-14 x 7-12 mm versus

6-7 x 5-6.5 mm) that are pale olive-green to

olive-green (versus dark olive-green).

A probable, undescribed taxon allied to

A. traceyi has been collected at Bolt Head,

Temple Bay and the Nesbit River (Vouchers: 4.5

km SW of the Nesbit River mouth, 57 km NE of

Coen, Silver Plains, 13° 13’S, 143° 32’E, Aug

1993, FellDGF3448 etal. (BRI, L); Bolt Head,

Temple Bay, 12° 14’S, 143° 04’E, Jun 1996,

Forster PIF19409 etal. (BRI); North of Beach

road, Bolt Head, 12° 15’S, 143° 05’E, Jun 1996,

Gray 6887 (BRI, QRS)) where it occurs in semi-

deciduous notophyll vineforest on stabilised

sand dunes. These sterile collections differ from

A. traceyi in the fewer lateral veins in the leaves.

Conservation status : Actephila traceyi is

commonly encountered throughout its known,

albeit restricted range. There is no information

available on the area of occupancy or the

number of individuals within populations.

Populations are not present in any conservation

reserves, but are not considered under threat.

Etymology : Actephila traceyi is named to

honour J.G (Geoff) Tracey (1930-2004), pioneer

ecologist, explorer and plant collector for CSIRO

in Australian rainforests. Geoff contributed over

10,000 herbarium collections to the Queensland

Herbarium and elsewhere.

Forster, Revision of Actephilia

13. Actephila venusta P.I.Forst., sp. nov. ab

A. lindleyi (Steud.) Airy Shaw venis in

folii lamina abaxiali usque ad tantum

ordinibus 1-2 (adversum ordines 1-3),

floribus pedicellis longioribus (flores

masculi: 5-14 adversum 1.5-5 mm,

foeminei: 13^14 adversum 2-14 mm),

sepalis ad apicibus non cucullatis

(adversum cucullata), disco floris masculi

5-angulato (in ilia rotundato), disco floris

foeminei maximo (1-1.8 x 4-6 mm

adversum 0.5-1 x 2-A mm) differt.

Typus: Queensland. Cook District: Possum

Scrub, road to Stones Crossing from Weipa, 12°

27’S, 142° 09’E, 8 December 1993, P./. Forster

PIF14372 (holo: BRI [3 sheets + spirit]; iso: A,

AD, CANB, DNA, K, L, MEL, NSW).

Actephila sp. (Possum Scrub P.LForster

PIF14372) (Forster & Henderson 1997:70;

Forster & Halford 2002:68).

Actephila sp. (Lockerbie WWC 817) (Hyland

etal. 1999,2003).

Subshrub or shrub to 4 m high. Indumentum of

simple trichomes, ferruginous-tan, entire plant

is glabrous unless stated otherwise. Branchlets

glossy, fawn-tan, lenticellate with age. Stipules

ovate-triangular to triangular, 0.8-2.4 mm long,

O. 7-1.8 mm wide, occasionally irregularly

crenate. Leaves alternate, subverticillate or

subopposite, coriaceous, petiolate; petioles 2-

33 mm long, 0.8-1.2 mm wide, olive-green,

lenticellate; lamina length/petiole length ratio

3.2-17.7 (n=239); lamina oblanceolate, obovate,

rarely elliptic or elliptic-ovate, ± flat, 26-155 mm

long, 13-82 mm wide, length/width ratio 1.2-3.3

(n=239); base acute, rarely rounded, often

unequal; tip acute or rounded, rarely retuse;

midrib above slightly raised (height < 5 times

the width), dark green, below strongly raised at

lamina base (height < or ± equal to width) then

decreasing towards apex, yellow-green;

venation brochidodromous, comprising 8-10

lateral veins per side of midrib and intercostal

reticulate veins; upper surface dark green and

glossy, lateral venation visible, intercostal

venation ± obscure; lower surface pale green

and glossy, lateral and intercostal venation

prominent and slightly raised, 1° and 2° vein

orders distinct, 3° vein order indistinct, 4° and

onwards vein orders obscure; margin entire, flat

to slightly recurved, pale green to yellow-green.

Inflorescence an axillary fascicle 1-4 mm

diameter, single sex or with both male and female

flowers. Male flowers: pedicels filiform, 5-14 mm

long, 0.2-0.4 mm diameter; sepals 5, oblong to

obovate, 2-5 mm long, 1.2-3 mm wide, densely

ciliate in patch at apex; petals absent; disk

conspicuous, fleshy, 1-1.5 mm high, 1.5^1 mm

diameter, 5-angled; stamens 5, filaments 2-3 mm

long, c. 0.1 mm diameter, anthers globose-ovoid,

c. 0.2 mm long and 0.4 mm wide. Female flowers:

pedicels filiform, 13-44 mm long, 0.6-1 mm

diameter; sepals 5, rarely 6, oblong to obovate,

4.5- 6 mm long, 2.5^1.5 mm wide, densely ciliate

in patch at apex; petals absent, of if rarely present

1 or 2, spathulate, c. 2 mm long and 1 mm wide;

disk conspicuous, fleshy, 1-1.8 mm high, 4-6

mm diameter, 5-angled; ovary 2-4.5 mm long,

3.5- 5.5 mm diameter; styles 3, simple, shortly

connate at base, 0.8-2 mm long, stigmas

subcapitate. Fruit: pedicels 13^14 mm long, 0.8-

1.2 mm diameter; capsules depressed-globose,

Fig. 12. Actephila venusta. A. abaxial leaf surface x0.5.

B. side view of male flower x6. C. face view of male

flower x6. All from Forster PIF14372 (BRI). Del. W.

Smith.

rounded, 8-9 mm long, 12-14 mm diameter,

strongly bullate, green to purple-brown;

columella 5-6 mm long. Seeds 5-7 mm long, 4-

6 mm wide, pale olive green; hilum 1.5-2.5 mm

long, caruncle poorly developed, narrow-

oblong, 1-2 mm long, 0.2-0.5 mm wide, cream.

Fig. 12.

Selected specimens examined: Northern Territory.

13 km SW Cape Arnhem, 12° 24’S, 136° 53’E, Feb

1994, Brennan 2587 (DNA); Marchinbar Island, South

Island of Two Island Bay, 11° 04’S, 136° 43’E, Oct

1994, Brennan 2934 (DNA); Elcho Island, 11° 46’S,

135° 55’E, Dec 1987, Dunlop 7589 (BRI, DNA, MEL,

NSW, QRS); Warangaya, Elcho Island, 11° 56’S, 135°

42’E, Sep 1987, Russell-Smith 3286 & Lucas (BRI,

DNA); Elcho Island, 14 km SW Naningburra Point,

11° 53’S, 135° 48’E, Dec 1987, Russell-Smith 4487 &

Lucas (DNA, MEL); NE tip of Elcho Island, 11° 48’S,

135° 56’E, Dec 1987, Russell-Smith 4498 & Lucas

(BRI, DNA); Gove, Dalywoi Bay, 12° 21’S, 136° 55’E,

Feb 1988, Russell-Smith 4669 & Lucas (BRI, DNA,

NSW, QRS); Elcho Island, 11° 55’S, 135° 50’E, Dec

1987, Russell-Smith 4518 & Lucas (BRI, DNA); west

bank near mouth of West Alligator River, 12° 15’S,

132° 15’E, May 1978, Webb & Tracey 12563 (BRI,

CANB, DNA). Queensland. Cook District: Prince of

Wales Island, Torres Strait, 10° 45’S, 142° 15’E, Feb

1975, Cameron 20132, 20133 (QRS); Pajinka, Cape

York, 10° 46’S, 142° 31 ’E, Feb 2001, Cooper

WWC1479 & Jensen (BRI, MEL); Mt Bremer, western

slopes, 26 km NE of Bamaga, Injinoo custodial land,

10° 42’S, 142° 31’E, Feb 1994, Fell DGF3922 et al.

(BRI, QRS); Iron Range N.P, 5.8 km SW of Cape

Weymouth, 12° 39’S, 143° 23’E, Mar 1994, Fell

DGF4114 & Stanton (BRI, MEL, QRS); Lake

Wicheura, 23 km NE of Bamaga, Injinoo custodial

land, 10° 46’S, 142° 33’E, Feb 1994, Fell DGF4074 &

Stanton (BRI); Muddy Bay, Cape York, 10° 43’S, 142°

33’E, Jun 1994, Forster PIF15330 & Tucker (BRI,

DNA); Lockerbie Scrub, 10° 47’S, 142° 29’E, Jun 1994,

Forster PIF15334 & Tucker (BRI, MEL); Mutee Head,

Cape York, 10° 54’S, 142° 14’E, Jun 1994, Forster

PIF15352 (BRI, MEL); Turtle Head Is., 10° 56’S, 142°

40’E, May 1995, Le Cussan 397 (BRI); Yam Island,

Torres Strait, 9° 05’S, 142° 46’E, Nov 1999, Wannan

I486 (BRI).

Distribution and habitat : In Queensland

Actephila venusta is found from islands in the

Torres Strait south to Iron Range. This species

is also found in a handful of localities in the

Northern Territory (Map 2). Plants grow in the

understorey of ‘dry’ rainforests (semi-

deciduous to deciduous, complex microphyll to

notophyll vineforests) on substrates (alluvial

or on ridges) derived from granites, laterised

sandstones, metamorphics or sand ridges at

altitudes between 30 and 430 m.

Austrobaileya 7 (1): 57-98 (2005)

Notes : Actephila venusta is closely allied to

A. flavescens and A. lindleyi and can be

considered a more northerly distributed sister

species. Herbarium collections of A. venusta

have been usually referred to A. lindleyi.

Actephila venusta differs from A. lindleyi in

the leaf lamina abaxially with 1-2° vein orders

distinct (versus 1-3°), the flowers with longer

pedicels (male flowers: 5-14 versus 1.5-5 mm;

female flowers: 13^14 versus 2-14 mm); sepals

that are not cucullate at the apices (versus

cucullate); the male flower disk 5-angled (versus

rounded) and the female flower disk much larger

(1-1.8 x 4-6 mm versus 0.5-1 x 2^1 mm).

Populations of Actephila venusta from the

driest sites (semi-deciduous microphyll

vineforests) in the Northern Territory and

western Cape York Peninsula tend to have

narrower, more oblanceolate leaves than those

from wetter areas (evergreen microphyll to

notophyll vineforest) such as at Bamaga.

A probable, undescribed taxon allied to

A. venusta has been collected in the vicinity of

Bathurst Head (Vouchers: 19.5 km ESE of

Bathurst Head, Kalpowar Pastoral Holding, 14°

20’S, 144° 21’E, Nov 1992, FellDGF2765 &

Stanton (BRI, QRS); 15 km SE of Bathurst Head,

14°21’S, 144° 17’E, Nov 1992, Fell DGF2774 &

Stanton (BRI, MEL, QRS); south of Bathurst

Bay, 14° 20’S, 144° 20’E, Oct 1970, Hyland4841

(BRI, QRS)) where it occurs in ‘seasonally dry’

rainforest (semi-deciduous complex notophyll

vineforest) on granite substrates. These sterile

collections differ in the strongly subverticillate

foliage with small lamina blades that are about

half the length of A. venusta.

Conservation status : Actephila venusta is

widely distributed in areas remote from

development. There is no information available

on the area of occupancy or the numbers of

individuals within populations for this species.

Actephila venusta is present in Iron Range

National Park and is not considered to be

threatened.

Etymology : The specific epithet is derived from

the Latin word venustus (beautiful) and alludes

to the flowers of this species.

Forster, Revision of Actephilia

Fig. 13. Actephila vernicosa. A. habit of flowering branch x0.5. B. abaxial leaf surface *0.5. C. side view of female

flower x6. D. face view of flower x6. E. side view of fruit xl.5. F. face view of fruit xl.5. A, B, E, F from Forster

PIF21249 (BRI); C, D from Cooper WWC1832 (BRI). Del. W. Smith.

14. Actephila vernicosa P.I.Forst., sp. nov. ab

A. lindleyi (Steud.) Airy Shaw differt

stipulis triangularibus usque truncatis

(adversum stipulas triangulari-ovatas)

quae minores sunt (0.7-1 x 0.5-0.8 mm

adversum 1-1.7 x 0.8-1.5 mm), foliis

chartaceis (in ilia coriaceis) petiolis

atroviridibus (non brunneis usque

olivaceis), floribus masculis sepalis

obtusis usque obovatis (adversum sepala

oblonga usque ovata) disco 5-angulato

(adversum rotundato) et frutice trilobato

(non rotundato).

Typus: Queensland. COOK DISTRICT: Millaa

Millaa Falls, 17° 30’S, 145°36’E,23 June 1997,

P.I. ForsterPIF21249, R. Jensen &M.C. Tucker

(holo: BRI [1 sheet + spirit]; iso: DNA, MEL,

NSW).

Actephila sp. (Millaa Millaa RJ 494) (Hyland

etal. 1999,2003).

Actephila sp. (Wooroonooran N.P.,

P.I.Forster PIF17151) (Forster &

Henderson 1997: 70; Forster & Halford

2002 : 68 ).

Subshrub or shrub to 3 m high. Indumentum of

simple trichomes, translucent, entire plant is

glabrous unless stated otherwise. Branchlets

glossy, fawn-tan, lenticellate with age. Stipules

triangular to truncate, 0.7-1 mm long, 0.5-0.8

mm wide. Leaves alternate or subverticillate,

chartaceous, petiolate; petioles 3-8 mm long,

0.8-1 mm wide, black-green, lenticellate; lamina

length/petiole length ratio 7.4-16.8 (n=322);

lamina elliptic, obovate, oblanceolate, rarely

orbicular, ± flat, 20-90 mm long, 11-50 mm wide,

length/width ratio 1.3-3.3 (n=322); base

attenuate to cuneate; tip acute, obtuse or

rounded; midrib above slightly raised (height <

5 times the width), dark green, below strongly

raised at lamina base (height < width) then

decreasing towards apex, pale yellow-green;

venation brochidodromous, comprising 6-9

lateral veins per side of midrib and intercostal

reticulate veins; upper surface dark green and

glossy, lateral venation visible, intercostal

venation ± obscure; lower surface pale green

and glossy, lateral and intercostal venation

prominent and slightly raised, 1-3° vein orders

distinct, 4° and 5° vein orders indistinct, 6° and

onwards vein orders obscure; margin entire, flat,

pale green. Inflorescence an axillary fascicle 0.5-

1.5 mm diameter, single sex or with both male

and female flowers. Male flowers: pedicels

filiform, 4-7 mm long, 0.3-0.5 mm diameter;

sepals 5, obtuse to obovate, 1.8-2 mm long, 1-

1.5 mm wide, ciliate in patches towards apices;

petals absent; disk conspicuous, fleshy, c. 0.5

mm high, 1.5-2.5 mm diameter, 5-angled;

stamens 4 or 5, filaments 1-2 mm long, 0.1-0.2

mm diameter, anthers globose-ovoid, 0.3-0.5 mm

long, 0.2-0.4 mm wide. Female flowers: pedicels

filiform, 5-25 mm long, 0.5-0.7 mm diameter;

sepals 5, obtuse to ovate, 2-2.8 mm long, 1.4-2

mm wide, ciliate in patches towards the apices;

petals absent; disk conspicuous, fleshy, 0.8-1

mm high, 3^1.5 mm diameter, 5-angled; ovary

1.5-3.5 mm long, 1.5-3.5 mm diameter; styles 3,

simple, shortly connate at base, 0.8-1.8 mm

long, stigmas subcapitate. Fruit: pedicels 18-

22 mm long, 0.8-2 mm diameter; capsules

depressed-globose to subglobose, trilobate,

12-17 mm long, 10-13 mm diameter, strongly

bullate, green; columella 5-6 mm long. Seeds

7-8 mm long, 6-7 mm wide, pale tan to olive

green; hilum c. 2 mm long, caruncle absent. Fig.

13.

Additional specimens examined : Queensland. Cook

District: Westcott road. Topaz, 17° 24’S, 145° 41’E,

May 1993, Cooper WWC539, 540 & Cooper (QRS);

he. tit, Nov 1995, Cooper WWC957 & Cooper (QRS);

loc. tit., Dec 1995, Cooper WWC958 & Cooper (QRS);

Ioc. tit., Oct 1996, Cooper WWC1064 & Cooper (BRI,

QRS); loc. tit., Dec 2003, Cooper WWC1832 & Cooper

(BRI); Stockwellia Site, Boonjee, 17° 25’S, 145° 45’E,

Jun 1995, Cooper WWC946 & Cooper (QRS); F.R.185

Robson’s L.A., near Danbulla, Mar 1961, Dansie 2027

(BRI); Danbulla F.R. 185, Aug 1962, Dansie 2397

(BRI); Scenic Reserve, Millaa Millaa Falls, Sep 1982,

Dansie AFOS219 (QRS); Stockwellia track,

Wooroonooran N.P., 17° 22’S, 145° 45’E, Jul 1995,

Forster PIF17151 & Figg (BRI, MEL); Wooroonooran

N.P., Gourka road (old Windin L.A.), 17° 22’S, 145°

42’E, Dec 2003, Forster PIF29778 & Jensen (BRI,

MEL); Stockwellia track, Boonjee, 17° 23’S, 145° 44’E,

Jul 1995, Hunter JH5287 (BRI); East slopes of Mt

Bartle Frere along the Falchetti track to the Mitchell

Bomber Crash Site, 17° 22’S, 145° 51’E, Jul 2001,

Jago 6009 & Gandini (A, BRI, L, MEL, NE, NSW);

Millaa Millaa Falls, 17° 30’S, 145° 36’E, Dec 1995,

Jensen 494 (QRS); S.F.R. 185, Maunder L.A., 17° 10’S,

145° 35’E, May 1972, Nicholson s.n. (QRS28440);

cult. Tolga (ex Boonjee), Apr 1997, Sankowsky 1558

& Sankowsky (BRI); loc. cit., Dec 2004, Sankowsky

2559 & Sankowsky (BRI); Millaa Millaa Falls, 17° 31’S,

145° 37’E, Feb 1986, Shapcott s.n. (BRI [AQ451830]);

VCL, Bartle Frere, E of Glen Allyn Trig., Feb 1962,

Webb & Tracey 5799 (BRI).

Distribution and habitat : Actephila vernicosa

is restricted to a small area bordered by the

Austrobaileya 7 (1): 57-98 (2005)

hamlets or localities of Bartle Frere, Danbulla

forestry, Boonjee and Topaz in the ‘Wet Tropics’

bioregion of north-eastern Queensland (Map 2).

Plants occur as small shrubs or subshrubs in

the understorey of ‘wet’ rainforest (evergreen,

complex notophyll vineforest) on red soil

derived from basalt or occasionally from granite,

at altitudes between 600 and 800 m.

Notes: Actephila vernicosa is an insignificant

plant that has been usually misidentified as

A. lindleyi in herbaria. The population from

Millaa Millaa Falls was one of the five

populations included in the allozyme study of

A. lindleyi s. 1. by Shapcott (1998) where it was

found to be significantly different from the

others (now variously identified as

A. grandifolia and A. lindleyi).

Actephila vernicosa is distinctive among

the Australian species of the genus in the small,

highly glossy, chartaceous leaves and the short

blackish-green petioles. It appears to be allied

to A. lindleyi but differs from that species in

the triangular to truncate stipules (versus

triangular-ovate) that are smaller (0.7-1 x 0.5-

0.8 mm versus 1-1.7 x 0.8-1.5 mm), the

chartaceous leaves (versus coriaceous) with

black-green petioles (versus brown to olive-

green); the male flowers with obtuse to obovate

sepals (versus oblong to ovate) and a 5-angled

disk (versus rounded); and the trilobate fruit

(versus rounded).

Conservation status : Actephila vernicosa has

a restricted distribution, but is locally common

in its known localities. There is no available

information as to the area of occupancy or the

numbers of individuals within populations.

Nearly all populations are present in National

Parks (Wooroonooran) or State Forests (S.F. 185

Danbulla) and are not presently under threat.

Etymology: The specific epithet is derived from

the Latin word vernicosus (varnished) and

alludes to the glossy leaves of this species.

Excluded Names

Actephila mearsii C.T.White, Proc. RoyalSoc.

Queensland 50: 85 (1939).

Base name for Peripentadenia mearsii

(C.T.White) L.S.Sm. (Elaeocarpaceae).

Actephila mearsii was still listed as a valid

species in the genus by Govaerts et al. (2000).

Forster, Revision of Actephilia

Acknowledgements

I wish to thank -

R. Booth (BRI), W. & W.T. Cooper, R. Jensen, G

Leiper, W. McDonald (BRI), W. Takeuchi (LAE),

G. Sankowsky, G. Smyrell and M.C. Tucker for

assistance during fieldwork where collections

of Actephila were made, or for making special

collections of material.

Will Smith (BRI) for the illustrations that were

funded by the Australian Biological Resources

Study (ABRS), along with some funding for

production of a ‘Flora of Australia’ treatment.

Peter Bostock (BRI) for translation of the

diagnoses into Latin.

Alex George, Gordon Guymer (BRI) and Annette

Wilson (ABRS) who whilst Australian Botanical

Liaison Officers at the Royal Botanic Gardens,

Kew, located and photographed various types.

The herbaria CANB, DNA, MEL, NSW, PR and

QRS for the loan of specimens.

Peter van Welzen (L) for assistance with

examination of types at L and U, and for

providing the unpublished account of

Actephila for the Flora of Thailand.

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Airy Shaw, H.K. (1980a). The Euphorbiaceae of New

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Airy Shaw, H.K. (1980b). New or noteworthy Australian

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Studies in Australian Myrsinaceae: TapeinospermaHook.f.

Betsy R. Jackes

Summary

Jackes B.R. (2005). Studies in Australian Myrsinaceae: Tapeinosperma Hook.f. Austrobaileya

7(1): 99-110. In Australia, there are three species recognised in the genus Tapeinosperma. One

new species T. pallidum Jackes is from North Queensland and one new combination and

reinstatement, T. repandulum (F.Muell.) Jackes, is a species that occurs in southern Queensland

and northern New South Wales. Descriptions, an identification key, illustrations and distribution

maps are provided for each species.

Key Words: Myrsinaceae, Tapeinosperma , Tapeinosperma pallidum, Tapeinosperma repandulum,

Australian flora.

B.R. Jackes, School of Tropical Biology, James Cook University, Townsville, Queensland 4811,

Australia.

Introduction

Currently in the genus Tapeinosperma Hook.f.

more than 60 species are recognised. However,

Pipoly and Takeuchi (2004) estimate that there

are about 100 species with many undescribed.

The distribution of described species ranges

from Indonesia east to Fij i and south to Australia

and New Caledonia, with more than half of the

species occurring in the latter area. Although

species have been described for the Philippines

these have all been transferred to Ardisia Sw.

(Stone 1989).

In Australia, two species now placed in

the genus Tapeinosperma were initially

recognised and described as belonging to the

genus Ardisia. In 1902, Mez combined these

two species (A. pseudojambosa F.Muell. and A.

repandulam F.Muell.) and transferred them to

Tapeinosperma as T. pseudojambosa. In

addition, he transferred the species described

as Embelia fliickigeri F.Muell., to

Tapeinosperma. Reynolds (1991) when

undertaking a revision of Embelia considered

that Mez was correct in transferring E.fliickigeri

to Tapeinosperma. However, this species, which

is known only from the type collection, does

not fit the genus as currently circumscribed.

The flowers are unisexual rather than bisexual

and the stamens are free from the corolla or

almost, rather than the upper portion only being

free. As noted by Smith (1973) and Sleumer

(1988) Tapeinosperma is difficult to distinguish

from a number of other genera in the

Accepted for publication 11 July 2005

Myrsinaceae unless the material is flowering.

In New Guinea, vegetative specimens can be

easily confused with Ardisia, Discocalyx Mez,

Fittingia Mez and Loheria Merr. Both Ardisia

and Tapeinosperma have bisexual flowers rather

than the unisexual flowers found in the other

genera; however, flowers in Discocalyx may

appear bisexual at times when the aborted

organs are still reasonably well-developed.

Hence the only reliable character separating all

the species in these three genera, is the nature

of the filament. In Tapeinosperma the apical

portion is ligulate and free from the corolla; in

Discocalyx it is absent or fused to the corolla

(Smith 1973), but in Ardisia the filaments are

free or only fused at the base to form a minute

tube which is adnate to the corolla or the anthers

are sessile or almost so. In addition,

Tapeinosperma can be distinguished from

Ardisia by the ovules being uniseriate rather

than multiseriate as found in most species of

Ardisia ; however, this distinction is not clear

cut for Australian species of Ardisia. In most

non-Australian species of Tapeinosperma the

stigma is obvious, however in the three

Australian species the stigma is punctiform and

similar to Ardisia stigmas. Species belonging

to these two genera in Australia can be readily

distinguished by the nature of the inflorescence

and the shape of the corolla.

In Australia, the species of

Tapeinosperma are distinguished from all other

species of Myrsinaceae by the following

combination of characters: inflorescence

100

paniculate; flowers bisexual; corolla fused at

the base to form a tube; staminal filaments short

but upper portion free and ligulate; ovules

uniseriate, style slender with a punctiform stigma

and fruit 1-seeded (many-seeded in Maesa

which is now in a separate family Maesaceae

(Anderbergera/. 2000)).

Materials and Methods

This study was based on an examination of

morphological characters derived from

herbarium material as well as some fresh

material. Herbarium specimens examined were

from the following Herbaria: BRI, CANB, K, L,

MEL, MO, NSW, QRS. All measurements were

based on dried material although some flowers

were reconstituted in boiling water prior to

examination. Secretory structures of a

schizogenous origin are present in the various

organs and were observed both with a hand

lens and under a microscope. These structures

which are referred to here as glands, vary in

shape and colour between species but appear

to be constant within a species. In thicker leaves

the pellucid glands may appear orange-coloured

or reddish if neighbouring cells are damaged,

as by insects. These cells are not to be confused

with the small globular deep red glands which

only occur in one species ( T. pseudojambosa).

A detailed description of these structures is

given in Otegui et al. (1998) and Jackes (2005).

Areole terminology is as described by Hickey

(1973). Trichomes were examined by light

microscopy and the basic structure confirmed

by SEM. Three types were recorded: uniseriate

2-3 celled; capitate ‘scales’ sunken into the

surrounding epidermis (Fig. lb), (although not

tested, some of these appeared to be secretory

and were similar in appearance to those

recorded by Bliithen and Reifenrath (2003) for

Ardisia pachyrrachis (F.Muell.) F.M.Bailey, as

extrafloral nectaries); stalked glandular papillae

are present on the margins of the anthers and

particularly towards the apex (Fig. la).

Taxonomy

Tapeinosperma Hook.f. in Benth. & Hook.f.,

Gen. PI. 2:647 (1876). Lectotype:

Tapeinosperma vieillardii Hook.f.

(Sleumer 1988).

Derivation of name: From the Greek tapeinos -

low or humble, and sperma - seed, referring to

the small seed.

Austrobaileya 7 (1): 99-110 (2005)

Shrubs or small trees, branches glabrous or

occasionally a few uniseriate hairs present

when young; leaf scars usually obvious, on

older branches lenticels often prominent;

terminal buds may bear flat capitate scales.

Leaves simple, alternate, subopposite to

opposite, often crowded towards the ends of

branches thus appearing pseudowhorled;

shortly petiolate; lamina in Australian species,

chiefly oblanceolate to obovate; glabrous or

with flat capitate scales scattered on the abaxial

surface; adaxial surface often glossy green,

abaxial surface paler, chartaceous to coriaceous;

apex acuminate, base cuneate, margins entire,

undulate, crenulate to irregularly serrulate;

venation obvious, lateral veins often numerous

and curving in from the margin; glands globular,

irregularly rounded to lineate, pellucid, or red

to dark red drying black, some small globular

red glands may be present; for Australian

species, the number of glands per areole lacking

veinlets varies from 1-8, to 25 in areoles with

veinlets. Inflorescence paniculate, (in non-

Australian species the panicle is sometimes

congested) terminal or arising from the axils of

the uppermost leaves; branches usually

reddish, subtended by chartaceous, usually

caducous bracts; pedicel subtended by small

caducous bracts; glands present. Flowers

bisexual, 5-merous, variants occasionally

recorded; glands irregularly rounded and/or

lineate. Calyx deeply divided, cupulifomi, in non-

Australian species may be rotate or occasionally

clavate, lobes thick, margins imbricate in bud,

glands present, pellucid to dark red to black.

Corolla urceolate, in some non-Australian

species campanulate or tubular, lobes may be

shorter than, equal to or longer than the tube,

apex rounded, glands if present globose or

shortly lineate, pellucid, dark red to black.

Stamens: filaments fused at base, adnate to the

corolla-tube at base, free and ligulate distally;

anthers oblong-deltoid, apiculate, dorsifixed,

longitudinally dehiscent, introrse, glandular

papillae present; pollen tricolporate, exine

reticulate. Ovary ovoid or occasionally

obturbinate, flat capitate scales may be present;

style slender elongating after anthesis; stigma

small and discoid or punctiform. Ovules 2-12,

borne in shallow grooves on the outside of the

ovoid placenta or, as in Australian species

embedded. Drupe globular to depressed

globular, reddish, exocarp thin, fleshy when

fresh, numerous glands present, endocarp

Jackes, Revision of Tapeinosperma

crustaceous smooth or angled, glands present;

style usually persistent but often broken. Seed

one, shape various, embryo transverse, thin,

curved, cotyledons elongated slender,

endosperm ruminate or not.

About 60 species are currently recognised

in the genus, chiefly from the New Guinea

region, Australia, New Caledonia and eastwards

to Fiji. The genus has been confused with both

Discocalyx and Ardisia. It may be readily

separated from Discocalyx by the presence of

bisexual flowers; stamens basally adnate to the

corolla tube, but distally free and anthers are

dorsifixed rather than anthers appearing almost

sessile on the corolla tube; and the long slender

style. In Ardisia the staminal filaments are free

or basally connate into a minute tube adnate to

the corolla or in most of the Australian species

the anthers are sessile or almost so. The

inflorescence in Australian species of

Tapeinosperma is paniculate and not umbellate

or congested as in Ardisia and the corolla is

urceolate with the lobes erect or weakly

spreading at maturity as against the corolla

101

being campanulate or rotate with widely

spreading lobes.

The genus consisting of three endemic

Australian species extends from North

Queensland (c. 15°S) along the eastern coastline

to northern New South Wales (c. 28°S). A fourth

species known only from the type collection,

which has been placed in both Embelia and

Tapeinosperma is now considered to be more

closely related to Embelia , because of the

presence of unisexual flowers, stamens free or

scarcely adnate to the corolla and the general

pubescence. Additional specimens are required

before it can be definitely assigned to an

existing genus.

Flowers in the Australian species appear

to be protogynous. The style and stigma

protrude from the urceolate corolla prior to

anther dehiscence and prior to the corolla lobes

becoming erect or even slightly spreading. No

pollen was observed on the style and stigma at

this stage.

Key to species of Tapeinosperma

1. Calyx and corolla with pellucid glands only. Leaf lamina with pellucid

glands only, (4—)6—8 per areole lacking veinlets, to 25 in areoles with

veinlets . 1. T. pallidum

Calyx and corolla with prominent orange to dark red to almost black glands.

Leaf lamina with pellucid glands only, or pellucid and red to dark red

glands, 1-6 per areole or up to 10 in areoles with veinlets.2

2. Corolla with orange and dark-coloured glands, the latter chiefly irregularly

globose, randomly distributed. Leaf lamina with scattered small globular

red to dark red glands, may be sparse; lineate pellucid glands with a

length: width ratio up to 4:1 rarely more, 1—4 per areole. 2. T. pseudojambosa

Corolla with very dark-coloured glands, chiefly lineate, restricted to the

median area. Leaf lamina lacking the small globular red glands;

lineate pellucid glands with a length: width ratio of 2:1 or less, 2-6 per

areole lacking veinlets, up to 10 in areoles with veinlets.3. T. repandulum

1. Tapeinosperma pallidum Jackes sp. nov.

A T. pseudojambosa in lamina folii

absentia glandium rubrarum, a T.

repandulo in lamina folii praesentia

glandium magnarum lineatarum

pellucidarum ratione (longitudo:latitudo)

quam 3:1 majore et aliquot usque 10:1 (in

illo non majus quam 2:1) et ab ambobus

absentia glandium rubrarum atratarumve

in corolla distinguatur. Typus: Queensland.

North Kennedy District: Keough’s scrub,

Portion 53V, Herberton, 17°30’S, 145°28’E,

20 November 1973, B.P. Hyland 7111 (holo:

102

QRS34393; iso: QRS34394, BRI, CANB,

NSW).

Tapeinosperma sp. (Cedar Bay J.G.Tracey

14780) in Henderson (2002).

Shrub or small tree 2-9 m tall, often much-

branched; outer bark smooth, brownish, blaze

pale pink with darker stripes, branchlets pale,

rarely reddish, usually angular with ridges

extending down from the base of the petiole,

leaf scars obvious; scales occasionally present

on terminal bud. Leaves alternate to

pseudowhorled: petiole 0.5-3(-4) mm long;

lamina oblanceolate, obovate, rarely tapering

abruptly to the apex, 4-15.8(-l 8) cm long, 1.2-

4.2(-5.6) cm wide, glabrous, scattered pale-

coloured scales may be present on abaxial

surface, adaxial surface glossy green, abaxial

surface paler, chartaceous; apex acuminate,

base cuneate, margins undulate, entire or

irregularly serrulate to crenulate, midrib

depressed on adaxial surface, prominently

raised on abaxial surface, 18-25 pairs of lateral

veins on either side of the midrib, looping near

the margins. Glands, usually about 6-8 per

areole lacking veinlets, up to c. 25 in an areole

with veinlets; chiefly irregularly globular and

shortly lineate with a length:width ratio of 3:1,

but some lineate with a length: width ratio up to

10:1 also occur, particularly near the midrib,

pellucid. Panicle 4-20 cm long, 4—8(—12) cm

wide, branches reddish, 5-11 primary branches,

the subtending bracts chartaceous, to 25 mm

long, often prominent particularly towards the

base, glands pellucid; panicle usually increases

in size as fruit develop. Pedicels (2-)5-10 mm

long, usually reddish, often curving and

lengthening after anthesis, glands pellucid,

bracts to 2 mm long, capitate scales

occasionally present. Flowers 2.5-3(-4) mm

long, glands pellucid. Calyx cupuliform, green,

tube 0.25-0.5 mm long, lobes triangular 1-1.5

mm long, c. 0.75 mm wide at base, often

spreading, glands pellucid. Corolla urceolate,

white, creamy green to pink, tube 1,5-2(-3) mm

long, lobes 0.5-1 mm long, c. 0.75 mm wide at

base, lobes often slightly spreading, glands

pellucid. Stamens: filaments ligulate, 0.5-1 mm

long, some glandular hairs may be present;

anthers 1.5-2 mm long, deltoid, apiculate, some

pale-coloured glandular papillae towards apex.

Ovary globular to obturbinate 0.7-1 mm long,

capitate scales present; style 2-3 mm long,

elongates after anthesis, stigma punctiform,

Austrobaileya 7 (1): 99-110 (2005)

glands pellucid. Ovules 4 or 5. Fruit globular to

depressed-globular, sometimes ribbed, 5-8 mm

long, 6-8 mm wide, red; style persistent. Fig. 1,

2,5A, 6A.

Selected specimens examined : Queensland. Cook

District: S.F.R.185, Danbulla, 17° 06’S, 145° 36’E, Apr

1992, Cooper & Cooper WWC238 (QRS); S.F.R. 143,

Windmill Creek, 16° 34’S, 145° 15’E, Nov 1997,

Cooper & Cooper WWC1172 (QRS); Mt Demi, 2 miles

[3.2 km] SW of Mossman, Nov 1944, Flecker

NQNC9017 (QRS); Flerberton Range, tributary of

Rocky Creek, 17° 14’S, 145° 25’E, Dec 1996, Ford

1836 (BRI, NSW, QRS); N.P.R. 904, Wooroonooran,

Coolamon Creek, c. 700 m S of Towalla Mine, site 34,

17° 28’S, 145° 44’E, Oct 2001, Ford 3049, Holmes &

Cooper (BRI, NSW, QRS); Davies Creek L.A., S.F. 607,

14.2 km past Davies Creek Falls, 17° 04’S, 145° 36’E,

Mar 1988, Forster PIF3888 (BRI, DNA); Daintree

N.R, Mt Sorrow track before razorback, 2.5 km W of

Cape Tribulation, 16° 04’S, 145° 27’E, Dec 1997,

Forster PIF21975, Booth, Jago & Jensen (BRI, K,

MEL, NSW, QRS); S.F.R. 185, Danbulla, Breach L A.

17° 05’S, 145° 38’E, Dec 1991, Gray 5364 (QRS);

F.R. 194, Atherton District, Mar 1962, Hyland AFO/

2787 (BRI); Mt Lewis road near the CSIRO plot, 16°

31’S, 145° 15’E, Nov 2001, Jago 6060 & Worboys

(BRI); Johnstone River, 1915 Michael 25 (BRI);

Downfall Creek area near northern shore of Tinaroo

Falls Dam, 17° 09’S, 145° 35’E, Sept 1972, Moriarty

1139 (CANB, NSW, QRS); Cedar Bay, 15° 49’S, 145°

20’E, Oct 1972, Tracey 14780 (BRI);. Mt Finnegan, S

of Cooktown 15° 47’S, 145° 17’E, Aug 1972, Webb &

Tracey 10856 (BRI, K); Malanda, Jan 1918, White s.n.

(BRI [AQ91814]). North Kennedy District: Bridge K10,

Douglas Creek, Kirrama, 18° 13’S, 145° 53’E, Nov

1997, Cooper & Cooper WWC1172 (QRS); Murray

River, Rockingham Bay, Nov 1869, Dallachy s.n.

(MEL1612793); Kirrama Range S.F. 344, c. 38 km

NW Kennedy, 18° 01’S, 145° 36’E, Nov 1989, Fell

DGF2000 (BISH, BRI, CANB, MEL, QRS); Mt Fox,

Dec 1954, Volck AFO950 (QRS).

Distribution and habitat : This species is

usually found in the rainforests of northern

Queensland particularly in complex mesophyll

vineforest, although it does occur in drier

situations on a variety of soil types. It extends

from the Cooktown area c. 15° 30’ S south to the

Ingham area c. 18° 30’S (Map 1). Altitudinal

range is from 300 to 1100 m.

Phenology: Flowering occurs in spring and

early summer, with a peak in November to

December. Fruits have been collected from

November through to September.

Notes: This north Queensland species is easily

distinguished when flowering as it is the only

species where there are no red or dark-coloured

glands present in the calyx and corolla. The often

conspicuous bracts subtending inflorescence

branches and the pedicels have pellucid glands

Jackes, Revision of Tapeinosperma

103

Fig. 1. Tapeinosperma pallidum. A. glandular papilla on anther. B. capitate scale on ovary. All from Cooper &

Cooper WWC1172 (QRS). Scale bar = 10pm

only, unlike T. repandulum where the bracts are

much smaller and have dark lineate glands, the

bracts in T. pseudojambosa are very early

caducous. In vegetative specimens, the

absence of red globular glands in the leaves

separates it from T. pseudojambosa. The number

of glands per areole, (4-)6-8 in areoles lacking

veinlets and up to 25 where veinlets are present

in the areoles, will usually readily separate this

species from both T. pseudojambosa and

T. repandidum where there are fewer glands per

areole, the latter species only occurs in southern

Queensland and northern New South Wales.

However, the latter is readily separated on the

length: width ratio of the lineate pellucid glands

which rarely exceed 2:1 compared to T. pallidum

where many exceed 3:1 and usually some much

longer ones occur with a ratio of up to 10:1.

Although usually an erect, much

branched shrub or small tree, there are a number

of records of the main stem lying horizontally

with branches arising vertically.

Etymology : From the Latin pallidus - pale,

referring to the pale-coloured flowers where

only pellucid glands are present in the calyx

and corolla.

2. Tapeinosperma pseudojambosa (F.Muell.)

Mez in Engler, Pflanzenr. 1(IV: 236): 170

(1902).

Ardisia pseudojambosa F. Muell., Fragm .4:81

(1864). Type: Queensland. North Kennedy

District: Mt Elliot, Fitzalan (lecto [here

chosen]: MEL1612779; isolecto:

MEL1612768).

Shrubs or small trees 1-12 m tall, trunk slender,

bark smooth, nondescript to mottled greyish-

brown, lenticellate, blaze cream to pinkish-red;

branchlets reddish with small ridges extending

down from base of the petiole. Leaves alternate

to subopposite, petiole reddish, 1-6 mm long,

often thickened, small flange may be present.

Lamina broadly obovate, usually abruptly

tapering to acuminate apex, 7-17.3 cm long, 2.2-

6.5 cm wide, glabrous or with rufous-coloured,

capitate scales on abaxial surface, chartaceous;

margins undulate, entire or crenulate, midrib

depressed on adaxial surface, prominently

raised on the abaxial surface; >23 lateral veins

on either side of the midrib; glands 1-4 per

areole rarely more; globular to shortly lineate

pellucid with a length: width ratio up to 4:1, rarely

pellucid glands absent, scattered small globular

red glands present, distribution and density

variable. Panicle 3-10 cm long, 3-11 cm wide,

enlarging as fruit develops, 4-10 primary

branches, reddish, the subtending bracts very

soon caducous, rarely present at maturity;

pedicels 4-7 mm long at anthesis, often curving

so flowers appear pendulous, glands obvious

dark red, bracts caduous c. 1mm long. Flowers

3—4 mm long, 5-merous, occasional 6-merous,

dark-coloured glands prominent on calyx,

corolla and style, pellucid glands sometimes

present. Calyx cupuliform, tube 0.25-0.5 mm

long, lobes triangular 1-1.5 mm long, c. 1 mm

wide at base, glands prominent, orange to dark

red. Corolla urceolate, creamy-green, white,

cream to pale pink, tube 1.5-3 mm long, lobes

triangular 0.5-1.5 mm long, c. 1mm wide at base;

orange to dark red glands prominent on lobes,

appearing verrucose, some pellucid glands may

104

Austrobaileya 7 (1): 99-110 (2005)

Fig. 2. Tapeinosperma pallidum. Scan of Cooper & Cooper WWC1172 (QRS).

Jackes, Revision of Tapeinosperma

be present on tube. Stamens: filaments thick,

ligulate, 0.5-1 mm long; anthers 2-2.5 mm long,

deltoid, apiculate, glandular rufous-coloured

papillae present. Ovary conical to depressed-

globular, 1-1.5 mm wide, c. 1 mm high, scattered

rufous-coloured capitate scales present, style

2-3(-5) mm long elongating after anthesis,

glands red, stigma punctiform. Ovules 6 or 7.

Fruit depressed-globular 6-7 mm long, 7-8 mm

diam., dark pink to red, glands black when dry,

style persistent. Fig. 3, 5B, 6B.

Selected specimens examined : Queensland. North

Kennedy District: Dugong Inlet, Whitsunday Is, 20° 05’ S,

148° 57’E, Sept 1990, Batianoff 900991A (BISH, BRI,

LAE); near South Pinnacle, 25 km SW of Townsville,

19° 24’S, 146° 38’E, Sept 1992, Bean 5053 (BRI);

Bluewater Range, WNW of Townsville, 19° 13’S, 146°

24’E, Nov 1996, Gumming 15306 (BRI); S.F.R. 344,

Kirrama road c. 1 km NNW of Mt Collins, site 62, 18°

10’S, 42° 18’E, Jun 2002, Ford 3494 & Holmes (BRI,

QRS); Mt Dryander S.F. Dryander Creek, left branch,

20° 16’S, 148° 34’E, Oct 1999, Forster P1F25000 &

Booth (BRI, DNA, K, L, MEL, QRS); Strathdickie,

near Proserpine, 20° 25’S, 148° 35’E, Michael 1116

(BRI); Dryander Creek, c. 2 km N of Gregory and c. 20

km N of Proserpine, 20° 24’S, 148° 35’E, Nov 1985,

Sharpe 4173 (BRI, CANB, K). South Kennedy District:

T.R. 179, Kalvin, 6 km W of Koumala, 21° 36’S, 149°

ll’E, Apr 1991, Forster PIF8033 & McDonald (BRI,

K, MEL, QRS); Mt Beatrice N.P northern tributary of

Catherine Creek, 21° 52’S, 148° 40’E, Jul 1993, Forster

PIF13393 & Tucker (BRI, QRS); Eungella N.P. Broken

River Circuit Track, 20° 55’S, 148° 35’E, Mar 1990,

Pearson 213 (BRI); Massey Creek, Eungella Range,

Oct 1951, Smith 4780 (BRI); Cathu S.F. 20° 50’S,

148° 35’E, Nov 1981, Young 438 (BRI). Leichhardt

District: On ranges south west from Sarina, May 1927,

Francis s.n. (BRI [AQ91823]). Port Curtis District:

Fitzroy River, Jun 1865, Bowman s.n. (MEL1612762);

Rockhampton, Jan 1862, Dallachy 111 (MEL);

Rockhampton, Oct 1865, Dietrich 1501 (MEL);

Colosseum Creek, T.R. 115, Many Peaks Range, 24°

23’S, 151° 27’E, Jan 1994, Forster PIF14635 (BRI,

MEL, QRS); Granite Creek, Bulburin S.F. 24° 33’S,

151° 31’E, Dec 1982, Guymer 1819 & Dillewaard

(BRI, CANB); Shoalwater Bay Training Area, site

SW07, Polygon range, E of Mt Mulgrave, 20° 37’S,

150° 17’E, Sep 1994, McDonald 5748 & Tweedie (BRI);

Rockhampton, Thozet 14 (MEL); Jeffrey’s property

“Shirley”, Agnes Waters road, turn left at Bindaree

then left at Jeffrey’s road, 24° 18’S, 151° 36’E, Oct

1998, Worthington 1937 (BRI).

Distribution and habitat: This species is found

chiefly in rainforest on drier sites or along creeks

growing on a variety of substrates. It is

frequently found as an understorey shrub

growing in association with Argyrodendron

spp. as canopy emergents and species of Gossia.

105

It extends from west of Cardwell on the drier

side of the Cardwell Range (c. 18°S) south to

Bulburin State Forest area (24° 33’ S) (Map 2).

Altitudinal range is from 70-600 m.

Phenology: Flowers from August to February

with a peak from September to early November.

Fruits have been collected from November to

May. The period from fertilisation to maturation

is approximately two months, but fruit may

remain attached for longer periods.

Notes: This species is easily distinguished from

the other two species by the presence of small

globular red to dark red glands scattered

throughout the leaf lamina, often at least one

per areole; however, in some specimens they

are sparse and in thick leaves they may be

difficult to find. Most areoles are simple and

lack veinlets, and there are l^l(-6) glands per

areole. The number of glands per areole

combined with the small globular red glands

can be used to identify sterile specimens of this

species. The obovate leaves abruptly tapering

to the apex are more common in this species

than in the others. It is the only species where

the orange to dark-coloured glands are

randomly distributed throughout the petals.

Etymology: From the Latin pseudo - false, and

the genus Jambosa Adans., a synonym of

Syzygium indicating that it is similar in

appearance to certain species of Syzygium.

3. Tapeinosperma repandulum (F.Muell.) Jackes

comb, nov.; Ardisia repandula F.Muell.

Fragm. 4: 82 (1864). Type: New South

Wales. North Coast District: Richmond

River, C. Moore, (lecto [here chosen]:

MEL654527).

Tapeinosperma sp. (Woombye A.R. Bean 994)

in Henderson (2002).

Shrubs or small trees 1.5-5 m tall, crown often

sparse and open; branchlets angular, may be

winged, reddish, leaf scars present, terminal

buds with small capitate scales. Leaves alternate,

subopposite to opposite, often in

pseudowhorls; petiole reddish, 2-4 mm long,

margins often with a small wing that continues

down the stem; lamina lanceolate to

oblanceolate rarely obovate, tapering to the apex

7.8-19.3 cm long, 2.3-5.6 cm wide, glabrous or

pale capitate scales sparse, adaxial surface

106

Austrobaileya 7 (1): 99-110 (2005)

QUEENSLAND HERBARIUM (BRI)

Flora of Queensland North Kennedy

Tapernosperma pseudoJambosa (F.Muell.) Mez

Coll. A.R.Bean 5053 27 SEP 1992

19°24'S 146°38 , E Alt. 600 m.

Depth m.

Near South Pinnacle, 25km SW of Townsville,

In rainforest gully with Raloghia, Archontophenix,

Coveniella.

Shrub 3 metres high.

Flowers white.

Det- Myrsinaceae

Dups,

*May be cited as computerised collection number AQ 549144

(Archival Paper)

Queensland herbarium (bri)

Brisbane Australia

aq 549144

Fig. 3. Tapeinosperma pseudojambosa. Scan of Bean 5053 (BRI).

Jackes, Revision of Tapeinosperma

glossy, abaxial surface paler, chartaceous; apex

acuminate, base cuneate, margins undulate,

irregularly serrulate to crenulate; midrib

depressed on adaxial surface, prominent on

abaxial surface, 15-25 lateral veins on either side

of the midrib, looping towards the margin.

Glands c. 2-6 per areole in areoles lacking

veinlets, up to 10 in an areole with veinlets;

globular to shortly lineate, length:width ratio <

2:1, pellucid. Panicle, 2-8 cm long, 2-3 cm wide,

3-5 primary branches, the subtending bracts

small, caducous. Pedicels 3-9 mm long, with

dark-coloured lineate glands, elongating and

curving as fruit matures; subtending bracts

chartaceous c. 1 mm long with dark-coloured

lineate glands. Flowers c. 3 mm long. Calyx

cupuliform, tube c. 0.75 mm long, lobes triangular

c. 0.75 mm long, 0.5 mm wide at base, dark-

coloured glands present. Corolla urceolate,

cream margins with dark to black centre, tube

1.5 mm long, lobes c. 1.5 mm long, glands

pellucid towards the margins with dark-coloured

chiefly lineate glands concentrated into the

median area of the lobes, drying black. Stamens:

filaments ligulate, 1 mm long, anthers 2 mm long

deltoid, apiculate, dark-coloured glands usually

present in apex, glandular papillae pale. Ovary

globose, c. 1 mm diam., pale capitate scales

present, style at anthesis 2.5-3 mm long

elongating as fruit develops, dark glands

present, stigma punctiform. Ovules 4 or 5. Fruit

depressed-globular, 6-7 mm long, 8 mm diam.,

dark pink to red. Fig. 4,5C, 6C.

Additional specimens examined'. Queensland. Wide

Bay District: Kin Kin, Mar 1916, Francis & White s.n.

(BRI [AQ91810], NSW26760); Kin Kin, Dec 1919,

Francis s.n. (BRI [AQ91812]); Between compartments

1 and 2, Mothar L.A., S.F. 393 Woodum, c. 15 km SE

of Gympie, 16° 15’S, 152° 49’E, Jan 1989, McDonald

4262 (BISH, BRI, MO, NSW); East Cedar Creek c. 7

km N of Mapleton, 26° 33’S, 152° 52’E, Jan 1990,

Sharpe 4934 & Thomas (BRI); Near Gympie-Imbil,

26° 20’S, 152° 40’E, Oct 1982, Williams s.n.

(NSW259576, UNE). Moreton District: Brolga Park,

beside Dulong road, W of Woombye, 26° 39’S, 152°

54’E, Jan 1989, Bean 994 (BRI); Brolga Park, 6 km

W of Woombye, 26° 39’S, 152° 54’S, Dec 1989, Forster

PIF6148, Bean & Tucker (BRI); c. 0.6 km SE of Mt

Wagawn, 28° 15’S, 153° 13’E, Feb 1986, Guymer 2109

(BRI); Landsborough, Shirley s.n. (BRI [AQ91822]);

Yandina, Mar 1891, Simmonds s.n. (BRI [AQ912821]);

Eudlo, Nov 1891, Simmonds s.n. (BRI [AQ91582]);

Mooloolah Scrubs, Dec 1890, Unknown (BRI

[AQ91818]); Maroochie, Mar 1891, Unknown (BRI

[AQ91829]); Eumundi, May 1892, Unknown (BRI

[AQ91813]); Eudlo Scrubs, Nov 1896, Unknown

(BRI[AQ91825]). New South Wales. North Coast

District: Couchy Creek, below Sphinx Lookout,

107

Springbrook, May 1977, Floyd 351 (BRI, CANB,

NSW); Richmond River, Hodge s.n. (MEL1612772);

Oxley River, c. 12 km NW of Tyalgum, 28° 15’S, 153°

10’E, Jun 1986, Williams & Bird 86005 (BRI).

Distribution and habitat: This species is found

along streams and in subtropical rainforest

areas, often in very moist habitats within these

forests. It has been collected from the Gympie

area (c. 26°S) south to the northern rivers area

ofNew South Wales (c. 28°S) (Map 1). Substrate

varies from rhyolite to volcanic derived soils

and it has been found at altitudes ranging from

80 to 700 m.

Phenology: Flowers have been collected in

summer and early autumn, while fruits have been

collected throughout the year but chiefly from

May to July.

Notes: Tapeinosperma repandulum is a typical

understorey species readily distinguished from

the other species by the distribution of the dark-

coloured glands in the corolla that are

concentrated in the centre of the corolla lobes,

with pellucid glands towards the margins.

Vegetatively it is the only species with lineate

pellucid glands rarely exceeding a length: width

ratio equal to or less than 2:1.

Etymology: From the Latin repandus - margin

is wavy or somewhat uneven, referring to the

margin of the leaf lamina.

Excluded species

Tapeinosperma fliickigeri (F.Muell.) Mez in

Engler \Pjlanzenr. 1(IV:326): 171 (1902)=Embelia

fliickigeri F.Muell. Viet. Nat. 8:200(1892).

Type: Queensland. Cook District: Russell River,

S. Johnson s.n. (holo: MEL1612758; iso: K).

Known only from the type collection and

presumed extinct. All existing specimens seen

were of poor quality. Additional material is

required to confirm placement.

Acknowledgements

I am grateful to the Directors of the following

herbaria - BRI, CANB, K, L, MEL, MO, NSW

and QRS for the loan of specimens or

permission to examine specimens and use their

facilities. My thanks to Peter Bostock for the

108

Austrobaileya 7 (1): 99-110 (2005)

0*1 12 .

FLORA OF AUSTRALIA PROJECT

QUEENSLAND HERBARIUM,

TTQr>£. r*/OG

BOTANIC GARDENS, BRISBANE.

f) tU-er j'C'i i-y - -Q v X,

/Cv~* , (jQ~ jcy^fx,

^ylq.

Accepted Name

7<5yoe/>-70 spe.i'rr^a, ps&u-cLojamUoscL-

Date /3 . //. Herb. BRI

Det -

Date ■ / x

Fig. 4. Tapeinosperma repandulum. Scan of Francis [AQ91812] (BRI).

Jackes, Revision of Tapeinosperma 109

Fig 5. A. portion of leaves (mid section) showing the pattern of distribution of glands within the areoles, stippled

glands are pellucid or orange coloured, solid glands are small globular red to dark red glands. A. Tapeinosperma

pallidum (from Dallachy s.n. (MEL612793); B. T. pseudojambosa (from Clemens s.n. (BRI [AQ 91827]); C. T.

repandulum (from Moore 212 (MEL)). Scale bar = 1 mm.

Fig. 6. Tapeinosperma pallidum. A. flower. T. pseudojambosa B. flower. T. repandulum C. flower. Lightly

stippled glands are pellucid. Scale bar = 1mm. A from Cooper and Cooper 1172 (QRS); B from McDonald 5748 &

Tweedie (BRI); C from Francis s.n. (BRI [AQ 91812]).

110

Latin diagnosis and to Adella Edwards for

preparing the maps and scanning specimens.

References

Anderberg, A.A., Stahl, B. & Kallersjo, M. (2000).

Maesaceae , a new primuloid family in the order

Ericales s.l. Taxon 49: 183-187.

Bluthen, N., & Reifenrath, K. (2003). Extrafloral

nectaries in an Australian rainforest: structure

and distribution. Australian Journal of Botany

51: 515-527.

Henderson, R.F.J. (ed.) (2002). Names and Distribution

of Queensland Plants, Algae and Lichens , p.

119. Environmental Protection Agency:

Brisbane.

Hickey, L.J. (1973). Classification of the architecture

of dicotyledonous leaves. American Journal

of Botany 60: 17-33.

Jackes, B.R. (2005). Revision of Myrsine (Myrsinaceae)

in Australia. Australian Systematic Botany 18:

399-438.

Mez, C.C. (1902). Myrsinaceae. In A. Engler (ed.) Das

Pflanzenreich 9(IV.236): 1-437. Cramer:

Weinheim/Bergstrafe.

Otegui, M.S., Gaspar, M.L., Maldonado, S., Varetti, E.L.

Austrobaileya 7 (1): 99-110 (2005)

& Pollero, R. (1998). Studies on tissues

associated with hydroxybenzoquinone

secretion in Myrsine laetevirens (Myrsinaceae).

Nordic Journal of Botany 19: 71-85.

Pipoly, J.J., & Takeuchi, W. (2004). New species of

Tapeinosperma and Discocalyx (Myrsinaceae)

from Morobe Province, Papua New Guinea.

Harvard Papers in Botany 8: 153-159.

Reynolds, S.T. (1991). The genus Embelia N.Burman

(Myrsinaceae) in Australia. Austrobaileya 3:

361-367.

Sleumer, H. (1988). The genera Discocalyx Mez,

Fittingia Mez, Loheria Merr., and

Tapeinosperma Hook.f. (Myrsinaceae) in New

Guinea. Blumea 33: 81-107.

Smith, A.C. (1973). Studies of Pacific Island Plants,

XXV. The Myrsinaceae of the Fijian region;

Tapeinosperma. Journal of the Arnold

Arboretum. 54: 228-263.

Stone, B.C. (1989). New and noteworthy Malesian

Myrsinaceae, III. On the genus Ardisia Sw. in

Borneo. Proceedings of the Academy of

Natural Sciences of Philadelphia. 141: 263-

306.

Map 1. Distribution of Tapeinosperma pallidum • and

T. repandulum A.

0 150 300 _600

• ••

Map 2. Distribution of Tapeinosperma pseudojambosa •.

Notes on the narrow-leaved Ironbarks (Myrtaceae: Eucalyptus

subseries Subglaucae )

A.R. Bean

Summary

Bean, A.R. (2005). Notes on the narrow-leaved ironbarks (Myrtaceae: Eucalyptus subseries

Subglaucae). Austrobaileya 7(1): 111-120. The taxonomic history of the narrow-leaved ironbarks

is discussed, as are the problems associated with the taxonomic delimitation of its member

species, particularly E. crebra sens. lat. and E. drepanophylla sens. lot. Eucalyptus elegans sp.

nov. is described and illustrated. The identity of the poorly known E. bowmanii Benth. is

discussed, and the circumscription of E. tholiformis is expanded.

Key Words: Myrtaceae, ironbarks, eucalypt taxonomy. Eucalyptus crebra, Eucalyptus

drepanophylla, Eucalyptus elegans, Eucalyptus tholiformis, Queensland flora. New South Wales

flora

A.R. Bean, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens,

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

Introduction

Ironbark is the common name given to a group

of Eucalyptus species with very hard, thick,

fissured grey to black bark. This common name

was first applied by the early colonists of New

South Wales and Queensland and it must have

been in common parlance by the 1840’s, as the

term was frequently used by Leichhardt (1847)

to describe eucalypt trees he saw with these

features.

Ironbark is perhaps the most distinctive

of all bark types in Eucalyptus sens, lat., and

more than any other bark type, it reflects

taxonomic affinity. In other words, all ironbark

species are related to each other, whereas

species with ‘gum’, ‘box’ or ‘stringy’ bark may

belong to more than one taxonomic section or

even subgenus.

Pryor & Johnson (1971) placed most

ironbarks into two informal series, viz. E. series

‘ Pruinosae’ and E. series ‘ Paniculatae\

Another series, E. series ‘ Melliodorae\

contained the two unusual ironbarks

E. sideroxylon and E. tricarpa.

Brooker (1985) showed that the informal

name ‘ Pruinosae’ is inappropriate, as

E. pruinosa Schauer is box-barked, whereas all

Accepted for publication 30 June 2005

other species in the series are ironbarks. He

proposed the series be renamed as informal

E. series ‘ Crebrae\ More recently, the informal

classification of Pryor & Johnson (1971) has

been replaced by formal names (i.e. using the

rules of the ICBN) and the ironbark series under

discussion became E. series Siderophloiae

Blakely. The eucalypt volume of the Flora of

Australia (Chippendale 1988) listed 12 species

belonging to E. series Siderophloiae. Since

then, 12 additional ironbark species belonging

to that series have been described by Brooker

& Bean (1987), Johnson & Hill (1990), Hill &

Johnson (1991), Bean & Brooker (1994), Hill

(1997a) andHill (1997b).

Brooker (2000) nominated four areas of

“poor resolution” within Eucalyptus , meaning

species groups poorly understood

taxonomically. The E. ser. Siderophloiae

ironbark group was one of these. Brooker ( loc.

cit .) maintained that E. ser. Siderophloiae could

be divided into two subseries; E. subser.

Subglaucae (the subject of this paper) and

E. subser. Jugatae Blakely. The latter is a small

but complex group of taxa characterised by

sessile opposite leaves retained in the crown of

mature trees. Brooker listed 20 species for the

E. subser. Subglaucae, excluding

E. drepanophylla (as it was listed as a synonym

of E. crebra in Brooker & Kleinig (1994)) and

E.farinosa (perhaps merely by omission).

112

Problems of recognising taxa

By the end of the 20 th century, nearly all of

the distinct ironbark taxa had been formally

described, leaving the morass of Eucalyptus

crebra and E. drepanophylla , both very

widespread and very variable in their

circumscription.

It was intended (by A.R. Bean & M.W.

McDonald) to elucidate the taxonomy of these

species using a combination of herbarium

studies, field studies throughout their range,

seedling trials and allozyme analysis. Initial

herbarium sorting of taxa was hampered by the

incomplete nature of most specimens which

lacked either mature buds or fruits or both. Very

few specimens included juvenile leaves. This

made it difficult to compare and classify

specimens.

Field studies did reveal a number of

seemingly distinct taxa from various parts of

Queensland and New South Wales, by allowing

access to a more complete range of fertile

material and juvenile leaf morphology. On that

basis, tentative taxa were erected and given

phrase names e.g. Eucalyptus sp. (Stannary Hills

G.W. Althofer 402), and specimens in the

Queensland Herbarium were thus annotated.

However, curation of the herbarium material

remained very difficult, as there were often no

characters that could consistently separate taxa.

Bean and McDonald collected seed of nearly

all of the tentative taxa identified from the field studies

for later allozyme analysis. For each provenance, the

seed collection was derived from five parent trees in

close proximity. For the tentative taxa perceived to

be widespread, up to four provenances were

collected, and for E. crebra and E. drepanophylla ,

seed collections from their type localities were

included. The allozyme study was undertaken (see

Achour 2003) and the results showed that

“Siderophoiae taxa were characterised by high levels

of allozyme variation and relatively similar allelic

frequencies. This is reflected in the lack of bootstrap

support for the clustering of most taxa...”

Seedling morphology has proved to be

taxonomically very useful in many groups of

eucalypts. Hence a comprehensive ironbark seedling

trial was undertaken by the present author. Seedlings

of all of the tentative taxa, as well as most of the

Austrobaileya 7 (1): 111-120 (2005)

already described species of the subseries were

raised to a height of30-40 cm, and scored for a

variety of characters. While the seedling

characteristics of the established species

(sometimes quite diagnostic) were confirmed,

there was little differentiation between the

tentative taxa, and the variation within taxa was

considerable. None of the tentative taxa (except

E. elegans, see below) could be distinguished

on the basis of seedling morphology.

Further field examinations, away from

“core areas” of the tentative taxa showed that

most intergraded extensively with the

geographically adjacent taxon, to such an extent

that additional taxa are impossible to define and

diagnose.

A similarly difficult taxonomic situation

exists in the Red Gum group {Eucalyptus subser.

Erythroxyla Blakely). Brooker & Slee (2000)

documented intergrades between several pair

combinations of long established species, and

they highlighted the difficulties of identifying

and classifying these intergrading forms.

Holman (2002) has studied the apparent

intergradation between the two ironbark

species, Eucalyptus whitei (of E. subser.

Subglaucae) mdE. melanophloia (of E. subser.

Jugatae). He has shown that there is almost no

genetic difference between them. These taxa

intergrade over a vast area of central

Queensland, forming a cline that “represents a

single cohesion species and a single

evolutionary lineage”. This research refutes

having these two species in different subseries.

Because the end points of the whitei-

melanophloia cline are so morphologically

dissimilar, the intergrades are very obvious. In

the case of the tentative taxa considered in this

paper, there are probably similar patterns of

clinal variation, but as the endpoints of the clines

are morphologically quite similar, the intergrades

are more cryptic.

A combination of these factors has forced

me to abandon nearly all of these tentative new

taxa, and revert to a broadly circumscribed E.

crebra and E. drepanophylla. While most of

the intergrading forms are referable to these two

species, there are several other species

involved. Intergrades have been observed

Bean, Narrow-leaved Ironbarks

113

between the following species pairs e.g.

E. drepanophylla and E. xanthoclada,

E. whitei and E. crebra, E. quadricostata and

E. xanthoclada, E. cullenii and E. crebra,

E. drepanophylla and E. granitica, E. atrata

and E. crebra.

Practicality and utility requires that the

existing species continue to be recognised, but

we need to be aware that some populations

cannot be identified with certainty.

One new species (E. elegans ) is described

here. Uniquely, it often grows in a mosaic pattern

with E. crebra without any intergradation or

hybridisation, and is considered sufficiently

distinct by virtue of its exceptionally small buds

and fruits, linear juvenile leaves, and linear to

narrowly lanceolate adult leaves. The poorly

known E. bowmanii Benth. is discussed, and

its identity is reaffirmed. The circumscription of

E. tholiformis has been expanded.

A key to the taxa comprising E. subseries

Subglaucae is presented. Of necessity, it uses

some characters observable only in the field. It

must be pointed out that for the narrow-leaved

ironbarks, imperfect herbarium specimens often

cannot be identified.

Taxonomy

Eucalyptus subser. Subglaucae Blakely, Key

Eucalypts 59, 250 (1934). Type: E.

siderophloia Benth.

Lignotuberous trees, rarely reduced to mallee

form. Bark on trunk rough, grey to black,

longitudinally furrowed, usually very hard and

unyielding, impregnated with kino, comprising

many very small to quite large lamellae;

commonly referred to as ‘ ironbark’. Rough bark

persistent on trunk and at least on the major

branches, and often persistent throughout. All

parts glabrous. Cotyledons reniform. Seedling

and juvenile stems more or less terete; seedling

leaves petiolate, opposite for 4-9 pairs; juvenile

leaves petiolate to subsessile, alternate. Adult

stems more or less terete, pith glands absent;

adult leaves glabrous, petiolate, alternate,

concolorous, margins entire; lateral veins at 45-

60 degrees to midrib; venation very densely

reticulate; intramarginal vein close to margin or

sometimes confluent with it; oil glands

numerous to occasional, circular, island or

intersectional, sometimes absent; petioles

terete. Conflorescences compound, pseudo¬

terminal and in upper leaf axils, unit

inflorescences umbellate, 3-7-flowered,

peduncles erect, pedicels present. Buds

smooth. Outer operculum shed early in bud

development. Stamens annular, in several

whorls, white, all fertile, irregularly flexed,

anthers adnate, basifixed, globoid, opening by

lateral non-confluent slits. Stigma blunt. Ovary

3-5 locular. Ovules in four longitudinal rows on

placenta. Fruits woody, with narrow

staminophore, disc descending or rarely

ascending, valve tips not connate; peduncles

erect, pedicels present. Seeds ellipsoidal,

somewhat flattened, grey-black, shallowly

reticulate, not winged, hilum ventral. Fertile

seeds much larger and darker than chaff.

Eucalyptus elegans A.R.Bean sp. nov. affinis

E. crebra autem foliis juvenibus linearibus

2.5-6 mm latis, foliis adultis pro parte

maxima linearibus, alabastris maturis 3-4

mm longis, fructibus 2.6-3.9 mm longis

differt. Typus: Queensland. Darling

Downs District: north end of Bendidee

National Park, NE of Goondiwindi, 27

November 1999, A.R. Bean 15864 (holo:

BRI; iso: CANB,NSW).

E. sp. (Chinchilla L. Pedley 4022) in Henderson

( 2002 )

E. sp. (Inglewood P. Grimshaw+ PG846) in

Henderson (2002)

E. sp. (GilgandraD.J. Carr+ 352) in Henderson

( 2002 )

Tree 8-28 m high, rough bark persistent

throughout. Seedling leaves opposite for 3^1

pairs, narrowly lanceolate, 55—61 x 8-11 mm,

discolourous, not glaucous, apex acute or

obtuse, margins entire, petioles 3^4 mm long.

Juvenile leaves linear, 45-83 x 2.5-6 mm, 12-22

times longer than wide, discolorous, apex acute,

base cuneate; petioles 2-4.5 mm long. Adult

leaves with petioles 9-14 mm long; lamina

narrowly lanceolate to linear, or falcate, 75-165

x 7-11 mm, 7-20 times longer than wide, apex

acute to attenuate, base cuneate, green to

bluish-grey, dull. Vein network dense, oil glands

appearing as isolated islands within the areoles.

114

Intramarginal vein present, single, close to

margin. Inflorescences largely terminal,

compound, the lower umbels axillary. Unit

umbels 3-7-flowered; peduncles terete, 2.5-7

mm long, 0.6-0.8 mm wide; pedicels 2-3.5 mm

long; buds obovoid to ellipsoid, at maturity 3-

4 mm long, 2.2-2.8 mm wide; hypanthium

wrinkled when dry, but not ribbed. Operculum

conical to hemispherical-umbonate, 1.1-2 mm

long, 1.8-2 mm wide, not ribbed, shorter than

hypanthium. Stamens white, outer whorls c. 2.5

mm long and inner whorls c. 1.3 mm long. Style

1-1.5 mm long at anthesis, stigma blunt or

dilated. Fruiting peduncles 1.5-6 mm long, 0.7-

0.9 mm wide; pedicels 1-3 mm long; fruits

cupular, circular in cross-section, not or very

faintly ribbed, 2.5-4 mm long, 2.7^1 mm wide,

staminophore 0.2-0.3 mm wide, disc obliquely

to vertically descending; valves 3-4, valve tips

exserted or at rim level. Seeds grey to black, c.

0.8 mm long, flattened, surface +/- reticulate.

Fig. 1.

Selected specimens examined : Queensland. Burnett

District: 2 km SW of ‘Rockybar’, Mar 1990, Bean

1433 (BRI); 30 miles [50 km] SW of Mundubbera, Sep

1969, Pedley 2892 (BRI). Darling Downs District:

Bracker State Forest, S of Inglewood, Dec 1990, Bean

2736 (BRI, CANB); S of Jackson, Sep 1948, Blake

18224 (BRI); Barakula S.F., Apr 1975, Brooker B4782

(BRI, CANB); 1.5 km E on road 4.9 km N of Barakula

Forest Office, Sep 1974, Chippendale GC1079 &

Brennan (BRI, CANB); Winfield Road, Weiambilla, N

of Tara, Sep 1998, Foley s.n. (BRI); Bracker S.F. 81, 2

km N of Brush Creek, 20.9 km S of Inglewood, Jul

1994, Grimshaw PG846 & Taylor (BRI); 5.8 km N of

Miles, Jan 2000, McDonald KRM245 (BRI); 23 miles

[38.3 km] N of Chinchilla, Nov 1968, Pedley 2782

(BRI); near Cecil Plains, Dec 1969, Pedley 3073 (BRI);

15 km WNW of Chinchilla, Dec 1972, Pedley 4022

(BRI); 23.2 km along Burncluith road, Barakula S.F.,

Oct 1993, Slee 3441 (BRI); c. 13.1 miles [21.1 km] W

of Condamine River on Moonie Highway, May 1961,

Smith 11321 (BRI); Bybera, via Inglewood, May 1934,

White 10059 (BRI). New South Wales. 28 miles [45

km] S of Gilgandra, Oct 1976, Carr & Carr 352 (AD,

BRI, CANB, MEL, NSW); Pilliga Scrub, Oct 1976,

Carr & Carr s.n. (BRI, CANB, MEL, NSW); Pilliga

Scrub, 25 km E of Baradine, Dec 1973, Streimann 703

(BRI, CANB).

Distribution and habitat: E. elegans extends

from Eidsvold to Inglewood in Queensland, and

again in the Narrabri - Gilgandra area of New

South Wales (Map 1). It grows on flat or gently

undulating ground with sandy soil near the

surface. Common associates are Callitris

glaucophylla and Casuarina luehmannii.

Austrobaileya 7 (1): 111-120 (2005)

Phenology: Flowers are recorded from June to

December; fruits may be found all year round.

Notes: Eucalyptus elegans is distinguished by

its tall habit, straight trunk, linear juvenile leaves,

mostly linear adult leaves, and very small buds

and fruits. In the Chinchilla and Inglewood

areas, it often grows in close proximity to

E. crebra, but without intergradation, with

E. crebra favouring the more hilly terrain.

Conservation status: The species is common

and widespread. No conservation coding is

required.

Etymology: From the Latin elegans meaning

elegant, attractive. This is a reference to the

fonn of the tree and the appearance of the crown.

Eucalyptus bowmanii Benth.

Bentham (1867) described E. bowmanii on the

basis of a single specimen collected by Edward

Bowman. This specimen, although in flower, is

deficient in many ways. All of the opercula have

been shed from the flowers, and none remains

with the specimen. There are just two

inflorescences, no fruits, and only five leaves.

There is no collecting locality, other than

‘Queensland’, and there is no indication of bark

type.

Because of the incomplete nature of the

specimen and lack of location and other

information, the identity of E. bowmanii has

long been in doubt. Maiden (1909) considered

it to be a doubtful species, either conspecific

with E. hemiphloia (=E. moluccana ), or closely

related to it. Blakely (1934) accepted it as a

distinct species, placing it alongside

E. drepanophylla. Pryor and Johnson (1971)

regarded it as a synonym of E. fibrosa subsp.

fibrosa. G.M. Chippendale examined the type

specimen in 1973 and determined it as E. fibrosa

subsp. fibrosa.

The present author has also recently

examined the type of E. bowmanii , and concurs

that E. bowmanii is conspecific with E. fibrosa

subsp. fibrosa. A brief description, gleaned from

the type specimen, is given here:

Adult leaves lanceolate, 23-34 mm wide, lateral

veins at around 45 degrees to the midrib,

Bean, Narrow-leaved Ironbarks

115

Fig. 1. Eucalyptus elegans. A. juvenile leaves x0.8. B. branchlet bearing axillary bud clusters and fruits x0.8. C.

umbel of mature buds (some buds have been shed) x4. D. hypanthium (at anthesis, stamens removed) showing

staminophore and style x8. G. mature fruits x8. A from Bean 16658 (BRI); B-E from Bean 15864 (BRI). Del. W.

Smith.

116

intramarginal vein c. 1.5 mm from margin, petioles

18-23 mm long. Inflorescences axillary, 7-

flowered; peduncles c. 14 mm long, c. 3 mm wide

at apex, angular; pedicels indistinct, thick,

angular; floral hypanthia (including pedicel)

narrowly obconic, 7-8 mm long, 4-4.6 mm wide

at distal end; staminophore c. 0.5 mm wide;

stamens white; anthers adnate, basifixed.

The basifixed adnate anthers confirm its

placement in E. sect. Adnataria. The two

axillary inflorescences present on the specimen

would suggest otherwise, as species in this

section are noted for their terminal

inflorescences, but they can and do have some

axillary umbels as well. The type of E. bowmanii

appears to coincide with the form of E. fibrosa

subsp .fibrosa that grows around Glen Geddes

and Marlborough, north of Rockhampton. The

recent collections of Batianoff9812232 et al,

Anderson s.n., and Batianoff 91081 & Robins ,

(all held at BRI), match the type almost perfectly.

Eucalyptus tholiformis A.R.Bean & Brooker,

Austrobaileya 4(2): 187 (1994). Type:

Queensland. Leichhardt District: Salvator

Rosa National Park, on ridge east of the

Sentinel, 18 May 1986, A.R. Bean 444

(holo: BRI; iso: BRI, MEL).

E. sp. (Rewan H.A.Kerswell 1) in Henderson

( 2002 )

Selected specimens examined : Queensland.

Leichhardt District: 19.8 km from Goonyella turnoff

towards Clermont, Aug 1986, Bean 504 (BRI); Nathan

Gorge, Oct 1989, Bean 1138 (BRI); Lookout track,

S.F. 236, SW of Blackwater, Nov 2002, Bean 19571

Austrobaileya 7 (1): 111-120 (2005)

(BRI); ‘Alice Wells’ on northern boundary of Mt

Playfair Station, Aug 1977, Blaxell 1448 & Armstrong

(BRI, NSW); near Yoothapinna Holding NW of Injune,

Apr 1975, Brooker 4856 (BRI, CANB); 12.5 km from

Mantuan Downs turn-off towards Springsure, Oct 1987,

Brooker 9781 (BRI); 83.7 km W of Springsure on

Tambo road, Aug 1984, Hill 1196 et al. (BRI, CANB,

DNA, MEL, NSW, PERTH); SE Boundary of Rewan

Holding, Portion 3 Wyseby, Apr 1987, Kerswell 1 (BRI);

Expedition Range, 27 km ESE of Rolleston township,

Aug 1961, Lazarides & Story 17 (BRI, CANB). South

Kennedy District: 69 km from Alpha towards Tambo,

Dec 1988, Hill 3607 & Stanberg (BRI, CANB, NSW).

Maranoa District: road to West Branch Camp, Mt

Moffatt N.P., Dec 1997, Bean 12845 (BRI, CANB).

Notes: The circumscription of this species has

been expanded to include populations with a

lesser amount of smooth bark on the upper

branches than at the type locality, and fruits

with a level or descending disc. These

populations were previously included under

E. sp. ‘Rewan’. The distribution of

E. tholiformis is hence extended considerably

to the east (Map 1). All of these forms are united

by the juvenile leaves that are consistently

broadly ovate, and 55-95 x 25-50 mm.

Eucalyptus crebra F.Muell. and Eucalyptus

drepanophylla Benth.

These two species are at the heart of the

taxonomic problems in the ironbarks, and

specimens from some locations e.g. Nebo,

Marlborough, are difficult to assign. However,

in other areas (e.g. Pentland, Clairview) they

co-occur without intergradation. The main

distinguishing characters are given in the table

below.

Table 1. Morphological comparison of Eucalyptus crebra & E. drepanophylla

Character

E. crebra

E. drepanophylla

junction of operculum

and hypanthium

(immature buds)

no constriction

distinct constriction, sometimes

referred to as egg-in-eggcup

rim of fruit (comprising

staminophore and part of

disc)

0.4-0.8 mm wide

0.7-1.2 mm wide

juvenile leaves

linear to narrowly lanceolate,

grey to blue-grey, not glossy

lanceolate, often pale lime-green,

sometimes +/- glossy

Bean, Narrow-leaved Ironbarks

117

118

Austrobaileya 7 (1): 111-120 (2005)

The traditional distinctions made between although E. drepanophylla , on average, has

these species, i.e. larger adult leaves, buds and broader adult leaves than E. crebra.

fruits for E. drepanophylla , do not hold,

Key to the narrow-leaved ironbarks (Eucalyptus subser. Subglaucae )

1 Buds and fruits with 4 obvious longitudinal ribs.2

Buds and fruits with 1-2 faint ribs, or ribs absent.3

2 Adult leaves ovate to orbicular, 3-6 cm wide; branchlets pruinose; pedicels 3-5 mm long

.E. farinosa

Adult leaves lanceolate, 1.5-2.2 cm wide; branchlets not pruinose; pedicels 5-7 mm long

.E. quadricostata

3 Apex of some or all adult leaves obtuse or retuse.4

Apex of all adult leaves acute to acuminate .5

4 Leaves lemon scented; fruits 5-6.5 mm long, pedicels 4-6 mm long. E. staigeriana

Leaves not lemon scented; fruits 3.5-5.5 mm long, pedicels 0-2mm long.E. jensenii

5 Inflorescences predominantly simple, axillary.E. rhombica

Inflorescences predominantly compound, terminal.6

6 Juvenile leaves broadly-ovate to orbicular (<2.5 times longer than wide).7

Juvenile leaves more than 2.5 times longer than wide.10

7 Juvenile leaves 2.5-5 cm wide.8

Juvenile leaves 5-12 cm wide.9

8 Outer branches smooth; adult leaves 15-30 mm wide. E. tholiformis

Bark rough throughout; adult leaves 10-20 mm wide.E. fracta

9 Buds and branchlets pruinose; fruits 6-8 mm long; umbels 7-11 flowered.

. E. fibrosa subsp. nubila

Buds and branchlets not pruinose; fruits 7-12 mm long; umbels 5-9 flowered.

.E. fibrosa subsp. fibrosa

10 Outer branches (up to 6 cm diameter) conspicuously smooth barked .11

Smooth bark absent or confined to branches <2.5 cm diameter.13

11 Branches 6-12 cm diameter smooth-barked . E. decorticans

Branches 6-12 cm diameter rough-barked.12

12 Juvenile leaves green, lanceolate, 1.5-2.5 cm wide. E. taurina

Juvenile leaves grey-green, linear, 0.8-1.3 cm wide. E. beaniana

13 Immature buds egg-in-eggcup .14

Immature buds without any constriction.16

14 Juvenile leaves glaucous.E. paedoglauca

Juvenile leaves green to grey-green, not glaucous.15

Bean, Narrow-leaved Ironbarks

119

15 Fruits large, 5-8 x 5-8 mm; mature buds egg-in-eggcup. E. xanthoclada

Fruits small, 4-6 x 4-5 mm; mature buds without constriction. E. drepanophylla

16 Juvenile leaves linear to narrowly lanceolate (7-20 times longer than wide).17

Juvenile leaves lanceolate to ovate (2.5-7 times longer than wide) .20

17 Fruiting disc broad and convex. E. cullenii

Fruiting disc narrow, descending .18

18 Adult leaves lacking intramarginal vein. E. exilipes

Adult leaves with intramarginal vein.19

19 Juvenile leaves 2.5-6 mm wide; fruits 2.5^1 mm long. E. elegans

Juvenile leaves 6-14 mm wide; fruits 4-7 mm long. E. crebra

20 Adult leaves bright green, quite glossy; juvenile leaves subsessile. E. granitica

Adult leaves blue-grey to green but not glossy; juvenile leaves distinctly petiolate.21

21 Operculum hemispherical to ellipsoid-truncate, apex obtuse .22

Operculum conical, apex acute.23

22 Buds and branchlets usually pruinose; operculum hemispherical.E. atrata

Buds and branchlets never pruinose; operculum ellipsoid-truncate. E. ophitica

23 Fruits typically ellipsoid-truncate; adult leaves 10-18 mm wide.E.whitei

Fruits cupular to obconical; adult leaves 18-30 mm wide. E. siderophloia

Acknowledgements

I am very grateful to Maurice McDonald for his

encouragement and discussions, his logistical

support during the field component of the work,

and for organising and supervising the allozyme

analysis for the ironbarks. Ian Brooker has

provided support and encouragement over

many years, and first suggested a revision of

the ironbarks to me two decades ago. My

numerous taxonomic discussions with him have

been invaluable. I also thank Will Smith for the

illustration and map, and Les Pedley for the

Latin diagnosis.

References

Achour, P. (2003). Phylogeny in the Red Ironbarks

(Eucalyptus series Siderophloiae). http://

www. anbg. gov. au/cpbr/summer-scholarship/

2003-projects/achour-ironbark.html

Bean, A.R. & Brooker, M.I.H. (1994). Four new species

of ironbark (Eucalyptus L’Herit., Myrtaceae)

from southern Queensland. Austrobaileya 4:

187-94.

Bentham, G. (1867). Eucalyptus. In Flora Australiensis

3: 185-261. L.Reeve & Co.: London.

Blakely, W.F. (1934). A Key to the Eucalypts. The

Worker Trustees: Sydney.

Brooker, M.I.H. (1985). The Ironbarks allied to

Eucalyptus crebra F.Muell. and the description

of a new species, E. quadricostata , in the group.

Austrobaileya 2: 148-152.

Brooker, M.I.H. (2000). A new classification of the

genus Eucalyptus L’Her. (Myrtaceae).

Australian Systematic Botany 13: 79-148.

Brooker, M.I.H. & Bean, A.R. (1987). Two new

ironbarks and a new bloodwood ( Eucalyptus ,

Myrtaceae) from Queensland. Brunonia 10:

189-200.

Brooker, M.I.H. & Kleinig, D.A. (1994). Field Guide

to Eucalypts , Volume 3. Inkata Press: Sydney.

120

Brooker, M.I.H. & Slee, A. V (2000). Studies in the Red

Gums of South-eastern Australia with particular

emphasis on Eucalyptus subser. Erythroxyla.

Australian Forestry 63: 86-106.

Chippendale, G.M. (1988). Flora of Australia Volume

19, Myrtaceae, Eucalyptus, Angophora.

Australian Government Publishing Service:

Canberra.

Henderson, R. J.F. (ed.) (2002). Names and Distribution

of Queensland Plants, Algae and Lichens.

Environmental Protection Agency: Brisbane.

Hill, K.D. (1997a). New species in Angophora and

Eucalyptus (Myrtaceae) from New South

Wales. Telopea 7: 97-109.

Hill, K.D. (1997b). New taxa in Eucalyptus

(Myrtaceae) from New South Wales and

Queensland. Telopea 7: 187-98.

Hill, K.D. & Johnson, L.A.S. (1991). Systematic studies

in the eucalypts - 4; New taxa in Eucalyptus

(Myrtaceae). Telopea 4: 321-49.

Austrobaileya 7 (1): 111-120 (2005)

Holman, J. (2002). Clines, species and eucalypts: an

evolutionary perspective. Unpublished Ph.D.

thesis. Griffith University: Nathan,

http: // www4. gu.edu.au: 8080/adt-root/uploads/

appro ved/adt-QGU20030527.124144/public/

02Whole.pdf

Johnson, L.A.S. & Hill, K.D. (1990). New taxa and

combinations in Eucalyptus and Angophora

(Myrtaceae). Telopea 4: 37-108.

Leichhardt, L. (1847). Journal of an Overland

Expedition in Australia. T. & W. Boone:

London, facsimile edition.

Maiden, J.H. (1909). Critical Revision of Eucalyptus,

Part X. Government Printer: Sydney.

Pryor, L.D. & Johnson, L.A.S. (1971). A Classification

of the Eucalypts. Australian National

University: Canberra.

Backhousia enata A.J.Ford, Craven & J.Holmes (Myrtaceae),

a new species from north-eastern Queensland

A. J. Ford 1 , L.A. Craven 2 & J. J. Brophy 3

Summary

Ford, A.J., Craven, L.A. & Brophy, J.J. (2005). Backhousia enata A.J.Ford, Craven & J.Holmes

(Myrtaceae), a new species from north-eastern Queensland. Austrobaileya 7(1): 121-127.

Backhousia enata A.J.Ford, Craven & J.Holmes is described, illustrated and compared with a

putatively related species. Notes on habitat, distribution, conservation status and an analysis of

essential oils are provided. The oil profile of the new species is most similar to that of B.

sciadophora F.Muell. whereas morphologically B. enata is very similar to B. myrtifolia Hook. &

Harvey, to which we believe it is most closely related. A revised key to the species of Backhousia

is presented.

Key Words: Myrtaceae, taxonomy, identification key, Australia, Queensland flora, Backhousia

enata, essential oils.

'A.J. Ford, CSIRO, Sustainable Ecosystems and Rainforest-CRC, Tropical Forest Research Centre,

PO Box 780, Atherton, Queensland 4883, Australia

2 L.A. Craven, Australian National Herbarium, CPBR, CSIRO Plant Industry, GPO Box 1600,

Canberra, ACT 2601, Australia

3 J.J. Brophy, School of Chemistry, University of New South Wales, UNSW, Sydney, New South

Wales 2052, Australia

Introduction

Backhousia is a genus of about nine species of

trees and shrubs and is endemic to Australia

(Guymer 1988; Bean 2003). The relationships of

the genus are with another Australian endemic

genus, Choricarpia , with which it shares a

distinctive embryo type (Wilson et ah 2001).

One of the species, Backhousia citriodora

F.Muell, is cultivated for its essential oils that

are used as flavouring in the food industry

(Doran etal. 2001).

During routine and targetted botanical

surveys for the Rainforest Co-operative

Research Centre (Rainforest-CRC), an unusual

member of the Myrtaceae was observed and

collected along the Tully River, north-eastern

Queensland. Subsequent visits yielded both

flowering and fruiting specimens, examination

of which indicated to us that it was a new

species of Backhousia. Accordingly, B. enata

is described below.

To complement a recent survey of the

essential oils of Backhousia (Brophy et ah

1995), samples of B. enata foliage were obtained

and the essential oils analysed.

Accepted for publication 30 June 2005

Taxonomy

Backhousia ewafa A.J.Ford, Craven & J.Holmes,

sp. nov. A B. myrtifolia Hook. & Harvey

ramulis 4-alatis, bracteis brevioribus (2-

2.6 mm longis), lobis calycis brevioribus

(1.9-2.2 mm longis), petalis complanatis

usque cucullatis, stylo breviore (2.2-3.3

mm longo) differt. Typus: Queensland.

North Kennedy District: Alcock Forest

Reserve, rafting access point No. 9, 5.2

km from Tully River Camping Area [NW

of Tully], 14 January 2003, A. Ford3792,

G. Sankowsky & N. Sankowsky (holo:

BRI; iso: CANB, K, L, MEL, MO, NE, NSW,

QRS, SYD).

Backhousia sp. (Tully River) in Cooper &

Cooper (2004:342)

Single to multistemmed large shrubs or trees,

5-15 m high; trunk diameters to 20 cm;

buttresses absent; trees usually with numerous

coppice shoots and smaller juvenile plants with

coppice and vegetative self-layering shoots;

bark of main trunk rough, minutely fissured, more

or less flaky, grey-brown; upper branches

mostly smooth, developing a flaky character

with increasing age; outer blaze (and wood) tan,

very fibrous. Spreading to antrorse, colourless

122

Austrobaileya 7 (1): 121-127 (2005)

to whitish uniseriate hairs to 0.7mm long present

on branchlets, leaves, petioles, inflorescence

axes, stipe, hypanthia and calyx lobes.

Branchlets conspicuously winged, the wing

extending from the base of each side of the

petiole to the node below; puberulent, hairs to

0.7 mm long; gland dotted; the branchlets

becoming glabrous and terete with age. Stipules

two or three, moderately persistent,

ferrugineous, setose, filiform, to 1.1 mm long.

Juvenile and adult leaves similar. Cataphylls to

13x6 mm, usually inserted proximally on each

vegetative growth unit, occasional vestigial

cataphylls (lacking any lamina but inserted at a

node) present amongst optimally developed

leaves. Leaves opposite, petiolate, decussate,

discolorous, dull on adaxial surface and shiny

on abaxial surface; lamina ovate to elliptic-

obovate, 24-38 x 11-18 mm, base cuneate to

roundly cuneate, apex acute to acute-acuminate;

margin flat or slightly recurved; both surfaces

with hairs to 0.4 mm long, densest along the

midvein, more or less glabrescent except for the

midvein; midvein raised on each surface, more

prominent on adaxial surface; oil glands

moderately dense and conspicuous on each

surface; venation brochidodromous,

inconspicuous on adaxial surface, primary

venation conspicuous on abaxial surface,

secondary venation discernible, tertiary

venation not discernible, 8-12 primary lateral

veins on each side of midvein; intramarginal

vein complete, 0.4—0.8 mm from margin, indented

to slightly looping at junction with lateral veins.

Petioles 1.5^1 mm long, channelled on adaxial

surface, puberulent, glandular. Inflorescences

in the upper leaf axils, paniculate, composed of

three to (usually) six flowers, the central flower

position of the inflorescence aborted and often

represented by a caducous, hairy, linear and

bract-like appendage c.l mm long; primary

inflorescence axis 6-10 mm long, weakly 4-

angled (not terete) and with shallow

longitudinal grooves on the abaxial and adaxial

surfaces, hairs to 0.7 mm long; secondary

inflorescence axis 1.9-2.1 mm long, hairs to 0.7

mm long. Bracts elliptic-ovate, 2.0-2.6 x 1.0-1.2

mm, caducous, apex obtuse to bluntly acute,

abaxial surface with hairs mostly in proximal half

and on the keeled midvein, adaxial surface

glabrous, margin hairy, oil glands conspicuous.

Bracteoles 1.5-2.0 mm long, narrowly elliptic,

abaxial surface with hairs mostly on the keeled

midvein, adaxial surface glabrous, oil glands

conspicuous, red-brown to orange-brown

finger-like colleters c. 0.15 mm long inserted

between bracteoles and at the base of the bracts.

Flowers white, perigynous. Hypanthium

stipitate, campanulate, 27-3.3 mm x 2.2-2.4 mm

including stipe, the outer surface with hairs,

glabrous inside except for scattered hairs on

the swollen disc adjacent to the calyx lobes and

the filaments, stipe 1.0-1.3 x c. 0.7 mm. Calyx

lobes 5, persistent, accrescent, flat, spreading,

greenish, sub-equal (3 outer and 2 inner), ovate

to triangular, 1.9-2.2 x 1.6-2.2 mm, apices obtuse

(outer) or bluntly acute (inner); on abaxial

surface moderately hairy more so in proximal

half, glabrous to minutely puberulent on adaxial

surface, oil glands conspicuous. Petals

caducous, 5, white, flat to cucullate, 1.8-2.0 x

1.6-1.9 mm, shortly clawed, ovate to nearly

orbicular, glabrous, obscurely veined, oil glands

sparse, alternating with calyx lobes. Stamens

56-70, in two whorls; filaments free, of variable

length in the same flower, 1-3.6 mm long, very

slender and becoming thread-like towards the

apex, glabrous; anthers dorsifixed, versatile, c.

0.15 x 0.2 mm, dehiscing laterally through

longitudinal slits. Style 2.2-3.3 mm long,

minutely hairy to ± glabrous, with sparse oil

glands, straight, tapering slightly towards the

apex, inserted in a shallow depression on the

ovary summit, stigma punctiform; ovary c. 1 mm

wide, summit glabrous or with scattered hairs,

flat to slightly convex, with a row of hairs c. 0.2

mm long on the periphery of the ovary wall which

extend above the ovary, adnate to the

hypanthium for about half the ovary length, 2-

locular, 6 or 7 ovules per locule, placentas axile.

Fruit dry, indehiscent, 2-2.2 x 2-2.3 mm

excluding calyx lobes; calyx lobes spreading,

to 5 x 4 mm; style elongating to c. 5 mm following

anthesis, puberulous. Seeds 5-7 per locule, c. 1

mm long, 3-faced with the abaxial face convex.

Fig. 1.

Additional specimens examined : Queensland.

NORTH KENNEDY DISTRICT. Alcock Forest

Reserve, rafting access point No. 9, 5.2 km from Tully

River Camping Area, Feb 2002, Ford 3275 & Holmes

(BRI, QRS); loc. cit ., Dec 2002, Ford 3780 & Holmes

(BRI, CANB, L, MEL, NSW, QRS); loc. cit., Mar 2003,

Ford 3844, Holmes & Cameron (BRI, CANB); Tully

River, Apr 2002, Cooper & Cooper WWC1727 (QRS);

near Carter Creek Falls above Carter Creek, Walter

Hill Range, c. 20 km S of Millaa Millaa, Jul 2003,

Graham s.n. (BRI, QRS).

Ford et al., Backhousia enata

123

Fig. 1. Backhousia enata. A. flowering branchlet x4. B. half flower x6. C. branchlet habit, showing wings x6. All

from Ford 3792 (QRS). Del. W.T. Cooper.

Distribution and habitat : Backhousia enata

is endemic to the Wet Tropics bioregion in

north-eastern Queensland, where it is currently

known to occur only in the Tully River

catchment, north-west of Tully (Map 1). It

inhabits notophyll vine-forest/rainforest on

soils derived from rhyolite and basalt. At the

type locality, which is riparian and where the

substrate is rhyolite, canopy species include

Xanthostemon chrysanthus, Buckinghamia

celsissima, Pseudoweinmannia lachnocarpa,

Carnarvonia araliifolia, Syzygium

tierneyanum and Carallia brachiata. Small

trees and shrubs at this locality include

Bursaria tenuifolia, Atractocarpus fitzalanii,

Chionanthus ramiflorus, Croton triacros,

Codiaeum variegatum var. moluccanum,

Mallotus polyadenos and Schefflera

actinophylla. However, in the Carter Creek

area, which is on a razor ridge with a basalt

substrate, Agathis robusta is the dominant

canopy tree. Here the understorey is very

sparse and consists of Hedraianthera

porphyropetala, Alyxia ilicifolia, Hoya

australis and Peperomia blanda var.

floribunda. Altitudinal range, from existing

specimens, is 80-310 m.

Phenology : Flowers have been recorded in

January; fruits have been recorded in March.

124

Austrobaileya 7 (1): 121-127 (2005)

Notes: On morphological features, B. enata

appears to be most closely related to B.

myrtifolia , and will key to that species in Guymer

(1988). It differs in critical features highlighted

in Table 1. In addition, B. myrtifolia and B. enata

are geographically separated, using current

data, by about 1,000 km.

The winged condition of adult branchlets

in B. enata is unusual within the genus. Some

species of Backhousia, e.g. B. bancroftii , B.

myrtifolia and B. hughesii, exhibit winged stems

but only when either at the seedling or small

sapling stage, or on coppice shoots, not on

adult specimens.

Special mention needs to be made of some

aberrant and widely disjunct fertile specimens

of B. myrtifolia , viz. Camira (Qld), Bird s.n.

(BRI[AQ563613]) (BRI); Border Ranges

National Park (NSW), ForsterPIF15109 (BRI);

Miriam Vale (Qld), Brushe TOI208 (BRI) and

Albion Park (NSW), Pullen 4239 (BRI). These

collections have very slight ridges on the adult

branchlets. Furthermore, the dimensions at the

lower extremes of measurements listed in Table

1 within B. myrtifolia are mostly attributable to

the Bird s.n. specimen. The Bird s.n. collection

vouchered an essential oil analysis reported

upon by Brophy et al. (1995); significantly, the

oils were consistent with those in other sampled

populations of B. myrtifolia. As the

aforementioned specimens lack the distinctive

features of B. enata listed in Table 1, they

should be regarded as merely morphologically

atypical forms of B. myrtifolia.

The leaves of B. enata are recorded as

being aromatic with a menthol-like smell. New

growth is red-purple. Its flowers lack any

distinctive odour.

The three outer calyx lobes are unusual in

that they have a thickened brown-red-purple

margin, whereas the two inner calyx lobes

usually lack this combined feature, presumably

this is a function of these tissues being exposed

in the bud stage. Of all the B. myrtifolia

specimens examined, only Brushe TOI208 has

calyx lobes with thickened margins. Although

not mentioned on the label, it is possible that

these thickened margins were coloured when

fresh and therefore similar to the calyx lobe

condition of B. enata. In addition to the stipules

that occur on each side of the petiole, several

short, colleter-like structures are inserted

between the axillary bud and the adaxial surface

of the petiole. These structures appear to be

homologous with similar structures discussed

by Weberling (1966) for Kania Schltr., by

Weberling (2000) for Tepualia Griseb. and by

Craven (1987) with respect to their occurrence

in Calytrix Labill.

The leaf oil of B. enata is dominated by

monoterpenes and these compounds

accounted for approximately 90% or more of

the oil, the remainder being sesquiterpenes. Our

analysis (voucher, Ford 3792) showed that the

principal components were a-pinene (14-17%)

and (3-pinene (36-42%), with terpinen-4-ol (5-

8%) and p-cymene (2-5%) being the next most

abundant components. The major sesquiterpene

was spathulenol (3-5%), with no other

sesquiterpene being greater than 1.5%. No

aromatic components were detected in the oil

of this species.

The composition of the oil from B. enata

bears no similarity to that obtained from B.

myrtifolia , the major components of which,

either singly or together, are the aromatic ethers:

methyl eugenol, E-methyl isoeugenol and

elemicin (Brophy etal. 1995). The oil of B. enata

is most similar to that obtained from B.

sciadophora F.Muell. which contained

significant amounts of a-pinene(44-55%), |3-

pinene (2-8%) and limonene (6-13%). In this

species too, monoterpenes accounted for

greater than 90% (Brophy et al. 1995). No

samples were analysed from the Carter Creek

populations.

Conservation Status: All existing collections

have been made within the World Heritage Area

of the Wet Tropics bioregion. As this species

has only recently been discovered it is

premature to discuss its conservation status.

Currently there are four known discrete

populations, with the total number of individuals

being less than 200. A preliminary search of the

Tully River in adjacent, and suitable, riparian

habitats failed to detect any further individuals.

However, as there are an estimated 100 plants

in the Carter Creek area (Graham, pers comm.

2003) it is not unexpected that several more

populations may exist within the poorly known

and rugged expanse of the Koolmoon, Cochable

Ford et al., Backhousia enata

125

Table 1. Differences between B. enata and B. myrtifolia . Dimensions of floral parts refer to flowers at

anthesis only, as calyx lobes and style elongate during fruit maturation.

Character

B. enata

B. myrtifolia

Branchlet shape (on adult and fertile

branchlets only)

conspicuously winged

terete (very rarely with slight ridges)

Bract shape

elliptic-ovate

narrow ovate-narrow elliptic

Bract dimensions

2.0-2.6 x 1.0-1.2 mm

3.5-6.1 x 1.1-2.0 mm

Primary inflorescence axis length

6-10 mm

14—40 mm

Calyx lobe dimensions

1.9-2.2 x 1.6-2.2 mm

3.1-11.0 x 1.6-5.0 mm

Outer calyx lobe apex

obtuse

acute

Petal dimensions

1.8-2.0 x 1.6-1.9 mm

2.1-3.8 x 1.7-3.0 mm

Style length

2.2-3.3 mm

5.0-8.1 mm

and Cannabullen Creek catchments. Therefore

it is appropriate to give B. enata a Data Deficient

(DD) status using the guidelines of the IUCN

( 2001 ).

Etymology : The specific epithet is from the

Greek, enatos , ninth, and refers to the locality,

Access point 9, on the Tully River. It was at this

locality that the type collection was made.

The following key is based upon that given in

Guymer (1988) with the addition of Backhousia

oligantha A.R.Bean and the present species.

B. anisata Vickery was removed by Wilson et

al. (2000) to a new genus allied to Syzygium

Gaertn., Anetholea Peter G. Wilson. However,

the distinction of Anetholea from Syzygium is

slight and the species has been transferred to

the latter genus (Craven & Biffin 2005).

Key to the species of Backhousia

1. Hypanthium stipe filiform (c. 0.25 mm diameter), (6—)8—l 8 mm long.2

Hypanthium stipe slender ( c . 0.5 mm diameter), 0.5-8 mm long.6

2. Leaves lemon-scented, narrow ovate, apex acuminate; branchlets and

inflorescences pubescent. SE Qld to NE Qld. B. citriodora F.Muell.

Leaves not lemon-scented, orbicular, ovate, obovate, or narrow obovate,

apex obtuse; branchlets and inflorescences puberulent or glabrous.3

3. Inflorescences of 3-11 dichasial clusters, 5-7 cm long; inner calyx lobes

4—5 mm long. NE Qld. B. hughesii C.T.White

Inflorescences of lor 2 (or 3) dichasial clusters, 3-5 cm long; inner calyx

lobes 2-2.5 mm long. 4

4. Inflorescence axes and hypanthia glabrous; leaves orbicular to ovate,

(4—)5-7.8 x 2.5-3.5 cm. SE Qld to NE NSW.B. sciadophora F.Muell.

Inflorescence axes and hypanthia puberulent.5

5. Trunk bark shed in strips; leaves ovate, obovate or narrow obovate;

inflorescences 8-20-flowered. CE Qld to SE Qld. B. kingii Guymer

126

Austrobaileya 7 (1): 121-127 (2005)

Trunk bark deciduous throughout; leaves elliptic or narrow ovate;

inflorescences (l-)3-flowered. CE Qld to SE Qld.B. oligantha A.R.Bean

6. Trunk bark deciduous throughout. CE Qld to SE Qld.B. oligantha A.R.Bean

Trunk bark rough, persistent throughout.7

7. Inflorescences terminal and axillary. NE Qld.

. B. bancroftii F.M.Bailey & F.Muell. ex F.M.Bailey

Inflorescences axillary.8

8. Adult branchlets conspicuously winged. NE Qld .... B. enata A. J.Ford, Craven & J.Holmes

Adult branchlets unwinged (rarely 4-ridged).9

9. Leaves ovate to narrow ovate, 2.2-6.5 x 0.9-4 cm; calyx lobes +/- equal,

5-13 mm long in fruit. SE Qld to SE NSW.B. myrtifolia Hook. & Harvey

Leaves narrow ovate to linear-oblong, 1.7-3.4 x 0.4-1 cm; inner calyx lobes

larger than outer, 2-3 mm long in fruit. NE NSW to NE Qld.B. angustifolia F.Muell.

Acknowledgements

The authors wish to thank Bill Cooper for the

illustrations. Peter Bostock provided the

distribution map. Field assistance was provided

to the first author by Wendy Cooper, Gary and

Nada Sankowsky and Bob Hewett. Maurie

Tucker, Kris Kupsch and Nan Nicholson

supplied additional useful information. Bob

Goldsack is thanked for help with the oil

distillations. Permits to collect in Alcock Forest

Reserve were issued by the Queensland Parks

and Wildlife Service. Andrew Graham (ex-

CSIRO) provided the information and specimen

for the Carter Creek populations. Peter Wilson

gave useful suggestions to the first author

during a visit to NSW. The curators and staff at

BRI, CANB, NSW and QRS are thanked for

allowing access to specimens and the use of

their facilities. An anonymous referee is thanked

for their useful comments.

References

Bean, A.R. (2003). Backhousia oligantha (Myrtaceae),

a new species from Queensland. Austrobaileya

6: 533-536.

Brophy, J.J., Goldsack, R.J., Folkes, C.J.R. & Forster,

P I. (1995). Leaf Oils of the Genus Backhousia

(Myrtaceae). Journal of Essential Oil Research

7: 237-254.

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

Australian Tropical Rainforest. Nokomis

Editions: Melbourne.

Craven, L.A. (1987). A taxonomic revision of Calytrix

Labill. (Myrtaceae). Brunonia 10: 1-138.

Craven, L.A. & Biffin, E. (2005). Anetholea anisata

transferred to, and two new Australian taxa of,

Syzygium (Myrtaceae). Blumea 50: 157-162.

Doran, J.C., Brophy, J.J., Lassak, E.V. & House, A.P.N.

(2001). Backhousia citriodora F.Muell. -

Rediscovery and chemical characterization of

the 1-citronellal form and aspects of its breeding

system. Flavour & Fragrance Journal 16:

325-328.

Guymer, G.P. (1988). A new species of Backhousia

Hook. & Harvey (Myrtaceae) from

Queensland and a reappraisal of Backhousia

floribunda A.J. Scott. Austrobaileya 2: 567-

569.

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rediistcatsenglish.pdf).

Weberling, F. (1966). Additional notes on the

Myrtaceous affinity of Kania eugenioides

Schltr. Kew Bulletin 20: 517-520.

Weberling, F. (2000). Tepualia stipularis (Myrtaceae),

una planta con appendices estipuliformes y

estipulas rudimentarias. Kurtziana 28: 313-

315.

Wilson, P.G., O’Brien, M.M. & Quinn, C.J. (2000).

Anetholea (Myrtaceae), a new genus for

Backhousia anisata : a cryptic member of the

Acmena alliance. Australian Systematic Botany

13: 429-435.

Ford et al., Backhousia enata

127

Wilson, P.G., O’Brien, M.M., Gadek, P.A. & Quinn, C.J.

(2001). Myrtaceae revisited: a reassessment

of infrafamilial groups. American Journal of

Botany 88: 2013-2025.

Map 1. Distribution of Backhousia enata ▲ in the ‘Wet Tropics’ of Queensland

Bazzania sauropoda D.Meagher (Marchantiophyta: Lepidoziaceae),

a new species from tropical Queensland

David Meagher

Summary

Meagher, D. (2005). Bazzania sauropoda D.Meagher (Marchantiophyta: Lepidoziaceae), a new

species from tropical Queensland. Austrobaileya 7(1): 129-133. Bazzania sauropoda sp. nov.

(Marchantiophyta: Lepidoziaceae), a liverwort from tropical rainforest in north-eastern Queensland,

is described, and its known distribution described.

Key Words: Lepidoziaceae, Bazzania sauropoda , bryophyte, liverwort, Marchantiophyta,

Queensland flora

D. Meagher, School of Botany, The University of Melbourne, Victoria 3010, Australia

Introduction

The genus Bazzania S.F. Gray (Lepidoziaceae)

comprises leafy liverworts with two rows of

lateral leaves inserted incubously on the stem,

a row of underleaves on the ventral side of the

stem, and minutely leafy ventral flagella arising

from the axils of underleaves. In almost all species

the branches grow as strongly as the stem from

which they arise, so that the branching is

distinctly Y-shaped and resembles dichotomous

branching. For this reason, such branching is

called ‘ pseudodichotomous \ The branches are

always of the Frullania type; that is, the branch

replaces the ventral half of a lateral leaf, leaving

the other half of the leaf in the branch junction

on the dorsal side.

Numerous specimens of Bazzania have

been collected in Queensland, but the names

given to them have been largely incorrect. Many

have been identified as Bazzania involuta

(Mont.) Trev., but that species is confined to

New Zealand and Tasmania (Meagher in press).

In Australia, the diversity of Bazzania

species generally increases with decreasing

latitude, culminating in a maximum diversity in

the Wet Tropics World Heritage Area, where at

least twenty species occur (Meagher 2003,

2005). Most of the species in northern

Queensland are already known from New

Guinea, Borneo, Java or other islands to the

north, but some are undescribed, including the

species now described here.

Accepted for publication 12 July 2005

Taxonomy

Bazzania sauropoda D.Meagher sp. nov.

Bazzania foliis suboppositis, non vittatis,

asymmetrice ovato-lingulatis, apicibus

asymmetrice 3-dentatis sed aliter integris;

amphigastriis incrassatis, praecipue

profunde 3-lobatis sed plerumque cum

lobis minoribus in lateribus; cellulis folii

valde incrassatis et nodulosis, omnibus

magnitudine similaribus; cellulis

amphigastriorum similaribus sed

minoribus; perianthio ubique

quadricarinato.

Typus: Australia, Queensland. Cook District:

summit of Mount Lewis, Sept 1986, G.A.M. Scott

s.n. (holo: MELU261; iso: BRI, CANB, F).

Plants robust, dioecious, strongly

anisophyllous, pendent. Branching

pseudodichotomous, the branches of

Frullania-typQ, spreading widely (usually

> 90°); branch dorsal half-leaf ± symmetric,

narrowly ovate, tapering to an acute, undivided

(rarely bifid) apex; first branch underleaf 1-fid

or nearly 2-fid, not connate with the adjacent

stem underleaf although closely situated to it.

Leaves subopposite, spreading widely when

moist but folded strongly to the ventral side of

the stem when dry, not vittate, asymmetrically

ovate-lingulate, mostly 1.2-1.6 mm x 0.65-0.8

mm, with an arched dorsal margin and a straight

to slightly incurved ventral margin; apex much

narrower than the rest of the leaf, mostly 3-fid

but sometimes obscurely so, rarely with an extra

130

Austrobaileya 7 (1): 129-133 (2005)

lobe or two; apical teeth short and blunt, sinuses

between lobes ranging from widely lunate to

narrowly triangular. Cells in mid-leaf mostly 20-

30(^10) x 20-25(-30) pm, slightly smaller near

the margins, in more-or-less regular longitudinal

rows, very thick-walled, strongly nodulose, with

massive, confluent and bulging trigones. Oil

bodies transparent, 2-5 per cell, very variable

in shape, from globular to ellipsoid, spindle-

shaped or almost oblong, undivided, slightly

lumpy. Underleaves mostly 0.45-0.60 mmx 0.35-

0.50 mm, connate with leaves on one side only,

variable in shape but mostly deeply 3-fid, lobes

often ending in a small apiculus; additional

smaller lobes sometimes present. Cells of the

underleaves as for the leaves but slightly

smaller. Perianth about 2.5 mm long, almost

sessile on a very short ventral branch, unevenly

4-keeled throughout its length, narrowing to a

very shortly ciliate mouth; outermost bracts

(bracteoles) with broad lobes, intermediate

bracts deeply and narrowly lobed, innermost

bracts ± laciniate; cells of intermediate and

innermost bracts ± rectangular, thick-walled,

most (except those in laciniae) with a strong

central papilla. Flagella numerous, long and thin,

sparsely to moderately leafy. Fig. 1.

Additional specimens examined : Queensland. Cook

District: Mount Lewis, Aug 1995, Brown 95/358 (NSW);

Platypus Creek, Jan 1936, Flecker s.n. (CANB); Lamb

Range, Apr 2005, Meagher 2101 (MELU); Mount

Lewis, Feb 1983, Streimann 29787, 29791 (CANB);

Lamb Range, Jun 1984, Streimann 29828 (CANB);

Bellenden Ker, South Peak, Mar 1983, Streimann

27348 (CANB); Mount Finnigan, Cedar Bay N.P., Oct

1995, Streimann 57207, 57208, 57209, 57226, 57135

(CANB); Windsor Tableland, Jun 1984, Streimann

29586 (CANB). North Kennedy District: Paluma Range,

Cloudy Creek, Jun 2005, Dalton s.n. (MELU); Cardwell

Range, Jun 1984, Streimann 28625 (CANB).

Distribution and habitat: Bazzania sauropoda

is known only from montane tropical rainforest

at seven locations in the Wet Tropics World

Heritage Area (Map 1). It is usually an epiphyte

on small trees, but one specimen was found

growing on rock.

Notes: Most species of Bazzania (and its partial

synonym Mastigobryum Gott., Lindenb. &

Nees) reported from the Australian tropics are

well-known species that occur also in South

East Asia and Melanesia (Meagher 2002,2005;

McCarthy 2003). Stephani(1908,1924) described

many species from the region and illustrated

them all in his unpublished ‘leones’. Significant

studies and reviews of the genus Bazzania have

since been undertaken for South East Asia and

Melanesia by Evans (1933), Grolle (1968,1972),

Herzog (1931,1949,1953), Kitigawa(1972,1973,

1979,1980), Meijer (1960) and Tixier (1985), and

for other nearby and distant regions of the

world by Amell (1963), Engel & Merrill (1994),

Engel & Schuster (1988), Fulford (1963), Grolle

& Schultze-Motel (1973), Hattori & Mizutani

(1958), Hodgson (1954), Jones (1975), Long &

Grolle (1990), Magill & Schelpe (1979), Mizutani

(1967), Poes (1969,1994), Scott(1985), Smith

(1990) and Wiggington & Grolle (1996). Among

the hundreds of species reported and described

by these authors, none resembles the present

species.

In Bazzania sauropoda the leaves are

much the same shape as some other species,

but the shape of the underleaves is very

distinctive and does not resemble that of any

other species. Furthermore, certain microscopic

characters such as nodulose cells of more-or-

less uniform size throughout the leaves and

underleaves, and massive bulging trigones, are

very uncommon in Bazzania.

Etymology: The specific epithet refers to the

shape of the underleaves, which is reminiscent

of the footprint of a large lizard.

Acknowledgements

Thanks to Dr John Engel (Field Museum,

Chicago, USA) for correcting the description

and for valuable comments on the manuscript;

Andi Cairns (James Cook University) for

organising permits to collect new material; the

Queensland Environmental Protection Agency

for permission to collect material on public land;

Dr Peter Wilson (National Herbarium of New

South Wales) for kindly correcting the Latin

diagnosis; Dr Elizabeth Brown (National

Herbarium of New South Wales) for important

corrections and suggestions on the manuscript;

and to the curators at the Australian National

Botanic Gardens Herbarium in Canberra (CANB,

CBG), the School of Botany Herbarium at The

University of Melbourne (MELU, MUCV) and

the National Herbarium of New South Wales

(NSW) for loans of specimens.

Meagher, Bazzania sauropoda

131

Fig. 1 . Bazzania sauropoda. A. dorsal view of portion of moist plant with two young gynoecia. B. lateral leaves.

C. underleaves. D. first branch underleaf and adjacent stem underleaf. E. underleaf. F. cells in centre of leaf. G.

female bracts (left to right) outermost, intermediate, innermost. H. perianth (dry). I. cells of innermost bract.

(Scale bars: A 5 mm, B-E, G-H 0.5 mm, F, I 0.05 mm, H 1 mm.).

132

Austrobaileya 7 (1): 129-133 (2005)

Map 1. Distribution of Bazzania sauropoda. 1: Mount

Finnigan. 2: Windsor Tableland. 3: Mount Lewis. 4:

Lamb Range. 5: Bellenden Ker. 6: Cardwell Range. 7:

Paluma Range.

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Enegl, J.J. & Merrill, G.L.S. (1994). Studies of New

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Society of New Zealand 82(1): 7-24.

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Kitagawa, N. (1972). Notes on little-known species of

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known species of Hepaticae, 26-50. Journal

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Kitagawa, N. (1980). New Guinean species of the genus

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68: 381-440.

McCarthy, P. (2003). Catalogue of Australian

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of southern Africa. Memoirs of the Botanical

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Meagher, D. (2002). New bryophyte records for

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367.

Mizutani, M. (1967). Studies of the Himalayan species

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North Viet-Nam. Journal of the Hattori

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Gyrostemon osmus Halford (Gyrostemonaceae), a new species

from south-eastern Queensland

D.A. Halford

Summary

Halford, D A. (2005). Gyrostemon osmus Halford (Gyrostemonaceae), a new species from south¬

eastern Queensland. Austrobaileya 7(1): 135-139. Gyrostemon osmus Halford is described,

illustrated and diagnosed against related species. Notes on habitat and distribution are provided. A

new section, G. sect. Monoecia Halford is proposed to accommodate the new species.

Key Words: Gyrostemonaceae, Gyrostemon , Gyrostemon osmus, Gyrostemon section Monoecia,

Queensland, Australian flora

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

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

Introduction

Gyrostemon Desf. is a genus of dioecious

shrubs and small trees distributed throughout

southern and arid Australia. In George’s (1982)

Flora of Australia treatment of the genus, he

recognised a total of twelve species, eight of

which are endemic in Western Australia. Until

now two species G. ramulosus Desf. and

G. tepperi(FMue\\. exHWalter)A.S.Georgewere

recorded for Queensland, from the far west of

the State.

The species described here as G osmus

was first brought to my attention in May 2004

when a specimen was sent to the Queensland

Herbarium for identification. The specimen was

initially misidentified as Codonocarpus

attenuatus (Hook.) H.Walter. The habit and

broad flat leaf lamina of G osmus gives this

species a superficial resemblance to

Codonocarpus attenuatus. However, G osmus

has fruiting carpels that dehisce along the

midline of the dorsal surface, a character that

distinguishes the genus Gyrostemon from

Codonocarpus which has fruiting carpels that

dehisce along the ventral surface.

Taxonomy

Gyrostemon osmus Halford, sp. nov. ut videtur

maxime arete affinis G racimigero autem

planta monoecia foliis grandioribus 6-20

x 0.7-3 cm (vice 1-5 x c. 0.2 cm) praedita,

Accepted for publication 18 July 2005

floribus maribus 13-15 staminibus in

verticillum unum dispositis (vice 45-50

staminibus in 4-5 series dispositis),

floribus femineis in pedicellis 2-3 mm

longis (comparitis ad flores femineos in

pedicellis 2-3 mm longis), stigmatibus 2.5

mm longis, ± linearibus (vice 1-1.5 mm

longis aliquantum petaloideis) differt.

Typus: Queensland. Moreton District:

Pages Pinnacle, c. 10 km WSW of

Mudgeeraba, 11 January 2005, D. Halford

Q8348 (holo: BRI; iso: AD, DNA, MEL,

MO, NSW, PERTH distribuendi).

Monoecious, short-lived perennial subshrub to

2.5 m high, single-stemmed or much branched

near ground. Branches hollow, stout, ± terete,

stiffly erect with few lateral branchlets, glabrous,

sparsely papillose when young becoming

smooth with age, green to yellowish orange

becoming orange to reddish orange in colour

with age, no cork development. Stipules

triangular, 0.2-0.7 mm long, acute. Leaves ±

sessile, spirally alternate, spreading,

discolorous; lamina flat, thin, lanceolate or very

narrowly elliptic-ovate, 6-20 cm long, 0.7-3 cm

wide, glabrous and smooth except for scattered

protuberances on margins and along mid and

lateral veins; base cuneate; apex attenuate;

midvein slightly raised on adaxial surface,

prominent on abaxial surface; lateral veins

widely spaced, c. 70° to midvein. Inflorescences

racemose, axillary, with 1-20 female flowers

basal on the axis and 1-60 males flowers distal

136

Austrobaileya 7 (1): 135-139 (2005)

to them; axis 0.8 to 10 cm long, papillose; bracts

linear-lanceolate 1.5-3.5 mm long, pale brown.

Male flowers on pedicels 1-2.7 mm long; calyx

cupular, c. 2 mm across, green, divided to c. 1/3

into 7 or 8 lobes; lobes broadly triangular, acute

at apex; stamens 12-15 in 1 whorl; filaments to

0.2 mm long, stout; anthers ± obloid-clavate,

1.8-2.2 mm long, rounded to retuse at apex, 2-

locular, dehiscing by lateral longitudinal slits;

rudimentary carpels present around central disc;

disc c. 0.8 mm diameter, flat or slightly convex, ±

smooth. Female flowers sessile or shortly

pedicellate; pedicels to 0.6 mm long; calyx

persistent, cupular, c. 1.5 mm across, green,

divided V 3 to into 5-8 lobes; lobes ±

triangular, obtuse to acute at apex; androecium

obsolete; carpels 10-12, free, attached around

central column; ovule 1 per carpel, attached near

centre of column; stigmas as many as carpels,

spreading at anthesis, marcescent, shortly

connate at base, ± linear, 2.5^1 mm long, ±

tetragonous in transverse section, glabrous,

longitudinally sulcate on adaxial surface,

tuberculate on abaxial surface; central column

expanded at top into disc; disc 0.4-0.5 mm

across, tuberculate. Fruits depressed globose,

3-5 mm long, 6-9 mm across; fruiting carpels

laterally compressed, reniform, scarious, ±

inflated, ± smooth and rounded on dorsal

surface, dehiscing along longitudinal midline

of dorsal surface into 2 segments which readily

separate from the persistent central column.

Seeds ± comma-shaped in lateral view, 1.7-2

mm long, dark brown, transversely rugose; aril

white, enclosing base of seed. Fig. 1 & 2.

Additional specimens examined: Queensland.

Moreton District: Pages Pinnacle, May 2004, Halford

Q8232 (BRI); Pages Pinnacle, c. 100 m SE of summit,

9.5 km WSW of Mudgeeraba, Apr 2004, O’Donnell 1

(BRI).

Distribution and habitat : Gyrostemon osmus

is endemic in south-eastern Queensland, where

it is currently known from the type locality on

Pages Pinnacle west of Mudgeeraba. It grows

in a tall open heath on narrow ledges and steep

slopes of a razorback ridge on shallow well-

drained greyish loam soils derived from rhyolite.

Associated plants include Allocasuarina

littoralis, Leptospermum microcarpum,

Lepidosperma later ale, Xanthorrhoea latifolia

subsp. latifolia, Dendrobium kingianum and

Platysace lanceolata.

Phenology : Flowers have been recorded from

January, April and May.

Affinities : In George’s (1982) keys to the species

of Gyrostemon, G. osmus will key to G. tepperi

in the key based on male plants and to G.

racemiger in the key based on female plants. A

comparison of diagnostic differences between

G. osmus, G. tepperi and G racemiger is provided

in Table 1.

Gyrostemon osmus is unique within

Gyrostemon and is easily distinguished from

all other species by the following combination

of characters: plants monoecious; leaf lamina

flat and thin, 0.7-3 cm wide; flowers in axillary

racemes; 12-15 stamens in single whorl; female

flowers sessile or with pedicel to 0.6 mm long;

10-12 carpels per flower; fruits depressed

globose with fruiting carpels not thickened

along midline of dorsal surface; stems without

corky bark.

Some may consider these morphological

differences sufficient to warrant the recognition

of a new genus. However, I feel it would be

premature to do so in the absence of more

complete knowledge of the variation within the

genus. George (2003) indicates that there are

several inadequately known taxa of Gyrostemon

in Western Australia.

Keighery (1985) recognised two sections

within Gyrostemon namely, G sect. Gyrostemon

and G. sect. Didymotheca (Hook.f) Keighery.

These sections were based on the number of

staminal whorls and number of carpels.

Gyrostemon osmus does not fit into either of

the sections as circumscribed by Keighery. The

attributes of a single whorl of stamens and 10-

12 carpels, together with plants monoecious,

and leaf laminae thin and flat, warrant the

segregation of G. osmus to a new section within

Gyrostemon. I here describe Gyrostemon sect

Monoecia to accommodate G. osmus. The

differences between the three sections of

Gyrostemon are set out in Table 2.

Gyrostemon sect. Monoecia Halford sectio nova.

A Gyrostemone sect. Gyrostemone et G. sect.

Didymotheca differt plantis monoeciis, foliis

teneris plus minusve planis (in illis plantae

dioeciae, foliis crassis pro parte maxima

filiformibus usque teretibus) etiam a sect.

Gyrostemone verticillo unico staminum (in illo

Halford, Gyrostemon osmus

137

Fig. 1. Gyrostemon osmus. A. inflorescence with male and female flowers *6. B. male flower viewed from above

x8. C. male flower viewed from below ><8. D. female flower viewed from side x 8. E. female flower viewed from

above x8. F. fruit viewed from above x6. G. lateral view of fruit x6. H. lateral cross section of fruit showing seed

attachment xlO. I. lateral view of seed xl6. J. ventral view of seed xl6. All from Halford Q8232 (BRI). Del. W.

Smith.

138

Austrobaileya 7 (1): 135-139 (2005)

Table 1. Morphological comparison of Gyrostemon osmus , G. tepperi and G. racemiger.

Character

G. osmus

G. tepperi

G racemiger

breeding system

monoecious

dioecious

leaf lamina shape

flat, lanceolate or very

narrowly elliptic ovate

linear-terete

leaf lamina dimensions

6-20 x 0.7-3 cm

0.5-3 x c. 0.1 cm

1-5 x c. 0.2 cm

male flowers per raceme

up to 50

1-3

up to 14

no. of staminal whorls

4-5

stamen number

12-15

7-10

45-60

female flowers per

raceme

up to 20

solitary or up to 3

up to 8

female flower pedicel

length

obsolete or up to 0.6 mm

1-2 mm

2-3 mm

carpel number

10-12

1 or 2

10-18 or more

stigma length

2.5-4 mm

c. 1 mm

1-1.5 mm

stigma shape

± linear

petaloid

± petaloid

aril

enclosing base of seed

almost enclosing whole

seed

enclosing up to !4 of the

seed

duobus usque aliquot) et a sect. Didymotheca

carpellis 10-12 (in illo 1-4) differ! Typus:

G. osmus Halford.

Conservation status : Gyrostemon osmus is

currently known only from the type locality.

Pages Pinnacle is not within a conservation

reserve; however, it is in an area set aside as

part of the Water Catchment area for the Hinze

Dam. The population has been observed to

fluctuate dramatically with as few as 5

individuals in August 2004 to approximately

1000 plants in January 2005. The increase in

population size was in response to a fire that

burnt through the habitat in October 2004.

Preliminary field observations indicate G osmus

has a soil-stored seedbank, the germination of

which appears to be induced by fire.

This species fulfils the criteria of “Critically

Endangered” under the IUCN (2001) categories

(CR Blac(iv); B2ac(iv)). Further searches of

likely habitat are required in south-eastern

Queensland and northern New South Wales.

Etymology : The specific epithet is derived from

the Greek osme meaning ‘odor, smell’; in

reference to the faint pungent odor which

emanates from the plant when crushed.

Acknowledgements

I am grateful to Will Smith for the excellent

illustration, Peter Bostock for the photograph

used in Fig. 2 and the translation of the diagnosis

of G. sect. Monoecia into Latin, Les Pedley for

the translation of the diagnosis of G osmus into

Latin and Gordon Guymer, Director of BRI, for

Halford, Gyrostemon osmus

139

Table 2. Morphological comparison of Gyrostemon sect. Gyrostemon , G sect. Didymotheca

and G sect. Monoecia

Character

G. sect. Gyrostemon

G. sect. Didymotheca

G. sect. Monoecia

breeding system

dioecious

monoecious

leaf texture

thick

thin

leaf shape

filiform to linear-terete

narrowly lanceolate,

narrowly obovate, linear to

linear-terete

lanceolate or very

narrowly elliptic-ovate

Number of staminal

whorls

2-numerous

Number of stamens

11-100

7-14

12-15

Number of carpels

2-30

1-4

10-12

assistance in the field and for making available

working space and facilities at BRI. An

anonymous referee is thanked for useful

comments.

References

George, A.S. (1982). Gyrostemonaceae. Flora of

Australia 8: 362-378. Australian Biological

Resources Study/CSIRO Publishing: Melbourne.

George, A.S. (2003). Gyrostemonaceae. In Kubitzki

(ed.). The Families and Genera of Vascular

Plants 5: 213-217. Springer-Verlag: Berlin,

Heidelberg.

Iucn (2001). IUCN Red List Categories: version 3.L

IUCN Species Survival Commission. IUCN:

Gland, Switzerland.

Keighery, G.J. (1985). Walteranthus , a new genus of

Gyrostemonaceae from Western Australia.

Botanische Jahrbucher fur Systematik

Pflanzengeschichte und Pflanzengeographie

106: 107-113.

Fig. 2. Gyrostemon osmus. Branchlet with flowers and

fruit. Photo: P.D. Bostock.

Eucalyptus erosa A.R.Bean (Myrtaceae), a new stringybark

species from central Queensland

A.R. Bean

Summary

Bean, A.R. (2005). Eucalyptus erosa A.R.Bean (Myrtaceae), a new Stringybark species from

central Queensland. Austrobaileya 7(1): 141-144. Eucalyptus erosa , a new species of the

stringybark group (E. ser. Pachyphloius Blakely) is described, illustrated and diagnosed against

closely related species.

Key Words: Myrtaceae, stringybark, eucalypt taxonomy. Eucalyptus erosa, Queensland flora.

A.R. Bean, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens

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

Introduction

Eucalyptus series Pachyphloius Blakely

comprises taxa commonly known as the ‘true

stringybarks’ (Ladiges and Humphries 1986).

They are characterised by the thick

longitudinally fibrous bark persistent

throughout the trunk and most or all branches,

and the seedlings with emergent oil glands

bearing numerous radiating hairs, in appearance

like stellate hairs. Frequently they exhibit sessile

buds and fruits, and oblique adult leaf bases.

Chippendale (1988) enumerated 21 species for

the group while Brooker (2000) listed 28 species.

The majority of these species occur in eastern

New South Wales and eastern Victoria, but

several species extend to Queensland, mostly

in higher altitude areas along the southern parts

of the Great Dividing Range in the south-east.

On the high ranges and plateaux between

Injune and Springsure, including the Consuelo

Tableland, two species of E. series

Pachyphloius are indigenous. One is

E. laevopinea R.T.Baker, which also grows

widely on the New England Tableland much

further south. On the Consuelo Tableland, it

forms a superb tall forest, “the best natural

stringybark forest in existence” (I. Brooker pers.

comm.). The second species (described here) is

most closely related to E. mackieana. It grows

on deep sandy soils derived from the

weathering of quartzose sandstone rock, and is

also a tree of impressive proportions.

Accepted for publication 30 June 2005

Taxonomy

Eucalyptus erosa A.R.Bean sp. nov. affinis E.

mackieanae sed plantulae foliis latioribus

densius hirsutis, pedicellis longioribus,

fructus disco piano et cotyledonibus

latioribus differens. Typus: Queensland.

Maranoa District : 13 km N of Mt Moffatt

Homestead, on road to Consuelo

Tableland, 27 April 1981, D.F. Blaxell 1883

& L.A.S. Johnson (holo: BRI; iso: CANB,

MEL, NSW).

Eucalyptus sp. (Carnarvon Range) in Ladiges

& Humphries (1986), Neish et al. (1995)

Eucalyptus sp. (Mt Moffatt HS D.F. BlaxelH

1883) in Bean et al. (2002)

Well-formed tree to 30 metres high,

lignotuberous, trunk straight. Bark persistent

except on small branches, coarsely fibrous to

stringy, grey on outside, brown underneath.

Juvenile leaves sub-opposite to alternate,

lanceolate to narrowly-ovate, 5-7 x 1.5-2 cm,

strongly discolorous, apex acute, base cuneate

to obtuse, margins minutely denticulate, petioles

0.2-0.3 cm long; stems, petioles and leaves

covered with clusters of simple radiating hairs,

in appearance like stellate hairs, each cluster

0.3-0.5 mm diameter. Intermediate leaves

broadly lanceolate, 7.5-9.5 x 2.5-3.5 cm,

glabrous, slightly discolorous, apex apiculate,

base oblique. Adult leaves falcate or lanceolate,

7-14 x 1.1-2.3 cm, alternate, concolorous,

penninerved, lateral veins at 30-45° to the

142

Austrobaileya 7 (1): 141-144 (2005)

midrib; reticulation very sparse, with numerous

large island oil glands, several per areole;

intramarginal vein present, 0.5-1.3 mm from the

margin; apex acute, mucronate or uncinate, base

cuneate to oblique, margins erose due to

marginal glands (4-9 on each side of lamina);

petioles 10-16 mm long. Inflorescences axillary,

unbranched, 9-11 (-13) flowered umbellasters;

peduncles angular to flattened, 7-14 mm long;

pedicels 2-4.5 mm long; immature buds

ellipsoidal; mature buds obovoid, 5.5-7 mm long,

3.5-4 mm in diameter, smooth and without ridges.

Hypanthium obconical, 3.5-4 mm long;

operculum conical to apiculate, 2^1 mm long,

scar absent; stamens white, irregularly flexed,

all fertile. Anthers reniform to cordate, versatile,

anther slits confluent. Style terete, stigma blunt

or tapered. Ovary 3-4-1 ocular, ovules in 2 vertical

rows. Fruits cupular to hemispherical, 4—6 mm

long, 6-7.5 mm in diameter, disc flat, 0.8-1.4 mm

wide, valves 3-4, at rim level, pedicels 2.5-5 mm

long. Seeds pyramidal to D-shaped, 1.6-2.0 mm

long, not toothed, dark brown, surface faintly

reticulate, hilum terminal. Chaff similar in size

and shape, but paler. Fig. 1.

Additional specimens examined : Queensland.

Maranoa District: 146 km NW of Injune on road to

Consuelo Tableland, Sep 1977, Blaxell 1529 &

Armstrong (BRI, NSW); 155 km NW of Injune on

road to Consuelo Tableland, Sep 1977, Blaxell 1524 &

Armstrong (BRI, NSW); between Warrong and Mt

Moffatt, NW of Injune, Apr 1975, Brooker B4867,

B4868 (BRI, CANB); Great Dividing Range, c. 80 km

SW of Rolleston, Kenniff Caves, Jun 1977, Crisp 3068

(BRI, CANB, NSW); 6 km W of Mt Moffatt HS, May

1982, Neldner & Thomas 749 (BRI). Warrego District:

c. 0.5 km N of Pumphole Spring, Dooloogarah Creek,

Feb 1977, Martensz 1148, 1149 & Johnston (BRI,

CANB).

Distribution and habitat : Eucalyptus erosa is

confined to the Mt Moffatt section of Carnarvon

National Park, north-west of Injune, and the

adjacent grazing property “Dooloogarah”. It

grows on coarse sandy colluvials derived from

outcropping quartzose sandstone, at altitudes

of700-900 metres. It occurs in association with

Angophora leiocarpa (G J.Leach) K.R.Thiele &

Ladiges, Eucalyptus grisea L.A.S.Johnson &

K.D.Hill and Callitris glaucophylla Joy

Thomps. & L.A.S.Johnson.

Phenology : Flowers have been recorded for

May. Fruits may be found throughout the year.

Notes : The most remarkable feature of E. erosa

is the consistent and conspicuous presence of

marginal glands on the adult leaves. On the

newly emerging leaves (new growth), they are

readily observed as pustular glands crowded

towards the distal end of the leaf (Fig. Id). On

fully grown leaves, they are more or less evenly

distributed along the margins (Fig. lc). By this

stage, the glands have usually erupted from the

surface along the margin, leaving a slightly

sunken, elliptical structure, that is then more

appropriately called a leaf-margin lenticel (Neish

et al. 1995). In this species and a few others

(e.g. E. denticidata ), the leaf-margin lenticels

are associated with irregularities in the leaf

margin that are conspicuous to the naked eye.

In other species, the leaf margin irregularities

are much more subtle.

“Irregular leaf margins” was one of the

characters used by Ladiges & Humphries (1986)

in their cladistic analysis of the stringybark

group. They noted its presence in E. erosa (as

E. sp. Carnarvon Range), but not for any other

member of the Stringybark group. Neish et al.

(1995) documented the occurrence of this

character in five species of Eucalyptus ,

including one additional species of stringybark,

E. laevopinea. However, this character is much

more widespread in the genus than has yet been

reported (Bean, unpubl. obs.).

E. erosa differs from the closely related

E. mackieana by its juvenile leaves 1.5-2 cm

wide (0.5-1 cm wide for E. mackieana ), the

seedling leaves with more than 30 hair clusters

per square centimetre (vs. <12 for

E. mackieana ), the pedicels 2.5-5 mm long in

faiit (1-2.5 mm long in E. mackieana ), and fruits

with a flat disc (convex disc in E. mackieana).

According to Ladiges & Humphreys (1986),

E. erosa has cotyledons 10-13 mm wide,

compared to 5-7 mm wide for E. mackieana.

E. mackieana has long been spelt

E. mckieana, but Recommendation 60C.4 of the

IUCN (Greuter et al. 2000), stated that “The

Scottish patronymic prefix Mac, Me or M’,

meaning “son of’, should be spelled ‘mac’ and

united with the rest of the name”.

Conservation status : There are no appreciable

threats to this species. Under the IUCN criteria

(IUCN 2001), E. erosa classifies as ‘Least

Bean, Eucalyptus erosa

143

Fig. 1. Eucalyptus erosa. A. juvenile leaves *0.5. B. branchlet bearing axillary bud clusters x0.5. C. adult leaf

showing the marginal glands *1. D. newly emerging adult leaves with pustular marginal glands x4. E. umbellaster of

mature buds (some buds have been shed) x3. F. umbellaster of mature fruits (some fruits have been shed) x3. G.

mature fruit, viewed from above x3. A,C from Maxell 1883 & Johnson ; B,D from Crisp 3068 ; E from Neldner &

Thomas 749 ; F,G from Martensz 1149 & Johnston (all BRI). Del. W. Smith.

144

Austrobaileya 7 (1): 141-144 (2005)

Concern’. Hence no conservation status is

recommended.

Etymology : The specific epithet is derived from

the Latin erosus , referring to the erose

(corroded, irregularly toothed, or apparently

gnawed) margins of the adult leaves in this

species.

Acknowledgements

I am grateful to Will Smith for the illustrations

and Peter Bostock for the Latin diagnosis.

References

Bean, A.R., Guymer, G.P. & Jessup, L.W. (2002).

Myrtaceae. In R.J.F. Henderson (ed.), Names

and Distribution of Queensland Plants, Algae

and Lichens, pp. 119-32. Environmental

Protection Agency: Brisbane.

Brooker, M.I.H. (2000). A new classification of the

genus Eucalyptus L’Her. (Myrtaceae).

Australian Systematic Botany 13: 79-148.

Chippendale, G.M. (1988). Eucalyptus, Angophora

(Myrtaceae), Flora of Australia Vol. 19: 142—

55. Australian Government Publishing Service:

Canberra.

Greuter, W., McNeill, J., Barrie, F.R., Burdet, H.M.,

Demoulin, V., Filgueiras, T.S., Nicholson, D.H.,

Silva, PC., Skog, J.E., Trehane, P, Turland, N.J.

& Hawksworth, P.L. (2000). International

Code of Botanical Nomenclature (Saint Louis

Code). Regnum Vegetabile , Volume 138.

International Association for Plant Taxonomy

(Europe).

Iucn (2001). IUCN Red List Categories and Criteria:

Version 3.1. IUCN Species survival

Commission. IUCN: Gland, Switzerland/

Cambridge, UK.

Ladiges, PY. & Humphries, C.J. (1986). Relationships

in the stringybarks. Eucalyptus L’Herit.

informal subgenus Monocalyptus series

Capitellatae and Olsenianae. phylogenetic

hypotheses, biogeography and classification.

Australian Journal of Botany 34: 603-632.

Neish, P.G., Drinnan, A.N. & Ladiges, PY. (1995).

Anatomy of leaf-margin lenticels in Eucalyptus

denticulata and three other eucalypts.

Australian Journal of Botany 43: 211-221.

Rediscovery and neotypification of Marsdenia arachnoidea Schltr.

(Apocynaceae: Asclepiadoideae - Marsdenieae), an

endangered species from Papua New Guinea

Paul I. Forster 1 & Wayne Takeuchi 2

Summary

Forster, P.I. & Takeuchi, W. (2005). Rediscovery and neotypification of Marsdenia arachnoidea

Schltr. (Apocynaceae: Asclepiadoideae - Marsdenieae), an endangered species from Papua New

Guinea. Austrobaileya 7(1): 145-150. The rediscovery of the long-lost Marsdenia arachnoidea

Schltr. is reported. A neotype is selected for the name M. arachnoidea and the species is described

and illustrated. It is endemic to Morobe Province of north-eastern Papua New Guinea, where the

only known populations are under threat from habitat alteration.

Key Words: Asclepiadaceae, Apocynaceae, Marsdenia arachnoidea, taxonomy, rediscovery,

conservation. New Guinea flora

‘PI. Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens,

Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: paul.forster@epa.qld.gov.au

2 W. Takeuchi, Herbaria and Arnold Arboretum of Harvard University, c/o National Forest Authority

Planning Division and the PNG Forest Research Institute, PO. Box 314, Lae, Morobe Province

411, Papua New Guinea.

Introduction

The genus Marsdenia R.Br. was recently

revised by Forster (1995) for Australia and

Papuasia (i.e. New Guinea, the Bismarck

Archipelago, and the Solomon Islands).

Including the single species described two

years later (Forster 1997) there are currently 79

species recognized for the region.

Several species from New Guinea were not

fully treated in Forster (1995) due to the

destruction of type specimens at the Berlin-

Dahlem Herbarium (B) in World War II, and the

absence of conspecific collections referable to

the lost types. Among these incompletely

treated species was the distinctive M.

arachnoidea Schltr. from north-eastern Papua

New Guinea (PNG).

Marsdenia arachnoidea is distinguished

by its long, filiform calyces, and by the unusual

corona lobes with a retuse apex and basally

involuted margins. Although the original

collections were lost long ago, M. arachnoidea

was fortuitously among the few species

illustrated by Schlechter (1913). In the absence

of authentically named specimens, Schlechter’s

illustration was previously the most effective

Accepted for publication 11 July 2005

means for identifying the species. With the

recent rediscovery of this plant, a more

comprehensive description of M. arachnoidea

can now be presented. In the following account

the new gatherings are illustrated and the

conservation status of extant populations is

also summarized.

Taxonomy

Marsdenia arachnoidea Schltr., Bot. Jahrb.

Syst. 50:151(1913).

Type Citation: “Nordostl. Neu-Guinea: kleinere

liane in den waldern oberhalb Dschischungari,

im Wariagebiet, ca. 1000 m ii M. (Schlechter n.

19845. - bliihend im Juni 1909)”

Type: Papua New Guinea. Morobe Province:

Lae, hills near Taraka, 6° 37’S, 146° 55’E, 25

October 2003, W. Takeuchi & D. Ama 16667 (neo

[here designated]: BRI; isoneo: A, LAE).

Illustration : Schlechter (1913:151, fig. 10).

Wiry vine, prostrate or scandent, herbaceous,

latex clear. Roots not seen. Indumentum of pale

yellow-brown trichomes on most parts. Stems

twining, cylindrical, up to 2 mm diameter, laxly

provided with trichomes when young, usually

146

Austrobaileya 7 (1): 145-150 (2005)

corky with age; internodes up to 14 cm long.

Leaves petiolate; lamina lanceolate or oblong-

lanceolate, up to 16 cm long and 5 cm wide,

chartaceous, discolorous, upper surface dark

green, glabrous except for scattered trichomes

on midribs, lower surface pale green, trichomes

short and sparse, denser on veins; colleters 8-

10 at lamina base, conical, pale brown; venation

brochidodromous, areolate in l(-2) series near

the margin, laterals 9-10 pairs, closing by

supramedial loops, bifacially prominent,

interlateral nervation reticulate, obscure above,

raised below; petiole 20-40 mm long, c. 0.8 mm

diameter, adaxially grooved, purple, trichomes

dense. Cymes racemiform, up to 10 mm long;

peduncles up to 8 mm long and c. 2 mm diameter,

trichomes scattered; bracts filiform-lanceolate,

up to 5 mm long, purple, with dense trichomes

to 0.7 mm long. Flowers 13-15 mm long, 50-55

mm diameter; pedicels c. 7 mm long and 1 mm

diameter, reddish-purple, with sparse, erect

trichomes to 0.5 mm long. Sepals filiform-

lanceolate, 18-19 mm long, c. 3.5 mm wide at

base and c. 0.5 mm wide at tip, reddish-purple,

with dense, erect trichomes to 0.5 mm long;

colleters two at each sinus base. Corolla

subrotate, reddish-purple and partly off-white

in the tube, glossy; tube flattened, c. 7 mm long

and 13 mm diameter, glabrous, with a raised

mound under each corona lobe, not pouched at

top; lobes filiform-lanceolate, fleshy, 24—25 mm

long, c. 9 mm wide at base and 0.5 mm wide at

tip, ciliate, otherwise glabrous; coralline corona

absent. Staminal corona c. 7 mm long, 13-14

mm diameter, cream, lobes not in close proximity

to one another, not overtopping style-head;

each lobe c. 7 mm long, somewhat involuted or

pouched at sides near the base, basal portion

roughly rectangular, c. 3 mm wide at base, upper

portion spathulate with a slightly retuse apex,

c. 1 mm wide at tip. Staminal column 5-5.5 mm

long, c. 4.5 mm diameter; anther appendages

pandurate, c. 1 mm long and 1.5 mm wide; alar

fissure c. 3 mm long. Style-head depressed-

globose, c. 3 mm long and wide, apiculate tip c.

1 mm long and 0.2 mm wide. Ovaries not

dissected. Pollinariac. 0.8 mm long, 1.6-1.7 mm

wide; pollinia ellipsoid, c. 0.8 mm long and 0.5

mm wide, yellow; corpusculum ovate, c. 0.7 mm

long and 0.4 mm wide, tan-brown; caudicles c.

0.3 mm long and 0.1 mm wide. Follicles and seeds

not seen. Fig. 1.

Additional specimen examined : Papua New Guinea.

Morobe Province: Lae, hills near Taraka, 6° 39’S, 146°

56’E, Aug 2004, Takeuchi, Ama & Morris 17029 (A,

BRI, CANB, LAE).

Distribution and habitat : Many of the

circumstances surrounding the original

discovery of M. arachnoidea are still unknown.

Although the principal details from Schlechter’s

expeditions have been compiled in summary

form (itineraries, place names, and trip routes;

see appendices in Schlechter 1911-14), the type

locality ‘above Dschischungarf is nowadays

impossible to place precisely as there is no

longer any record of such a village (e.g.,

Department of District Administration 1968;

Anonymous 1970). By reconstructing

Schlechter’s itinerary, it is otherwise clear that

the species was first discovered somewhere in

the south-easternmost part of the Bowutu

Range (Dischore Range fide Schlechter), in

proximity to what is today known as Mt Hahl

(Map 1). A recent trip to this general vicinity

(by one of the authors) was aborted by land

ownership disputes. Investigation of the lower

drainage by orchidologists has previously

resulted in the rediscovery of several Schlechter

species (Clements & Ziesing 1990). Although

the Bowutu Range is the most extensive series

of ultrabasic environments in New Guinea, the

presumed type locality for M. arachnoidea is

apparently outside the serpentine zone (see

Dow & Davies 1964). While this is not entirely

certain because of the geographic imprecision

of the type, the new record at Taraka is clearly

from a non-ultrabasic habitat.

At the Taraka locality, M. arachnoidea

occurs as an umbrophile in remnant patches of

lowland rainforest between 200-300 m elevation.

On the current system of forest resource

classification (Hammermaster & Saunders

1995a) the new site is mapped as a medium-

crowned low elevation forest (structural code

Hm). The entire subdistrict is experiencing

severe impacts from anthropogenic

development, and the latest vegetation overlays

are already outdated (Hammermaster &

Saunders 1995b). The regrowth mosaic around

Lae currently extends to a front about 16 km

outside the city.

The rediscovery of M. arachnoidea is

extremely ironic, in view of the plant’s prolonged

Forster & Takeuchi, Rediscovery of Marsdenia arachnoidea

147

Fig. 1 . Marsdenia arachnoidea. A. stem with flowering inflorescence x0.5. B. base of leaf lamina with colleters *6.

C. adaxial leaf surface showing venation x0.5. D. bud *1.5. E. face view of flower *4.5. F. back view of flower

shosing sepals xl. G. face view of staminal column and corona x3. H. side view of staminal column and corona x3.

I. face view of pollinarium x20. All from Takeuchi 16667 (BRI). Del. W. Smith.

148

Austrobaileya 7 (1): 145-150 (2005)

absence from the scientific record and the

belated realization that it has been present all

this time on the outskirts of PNG’s second

largest city. The new occurrence is just 5 km

from the University of Technology, the principal

training center for forestry science in the

country, and is only 7-8 km from the National

Herbarium. Tapeinosperma magnified Pipoly &

Takeuchi was recently described from the same

habitat as the new populations of

M. arachnoidea and the two species occur

together in the Taraka understory.

Typification: For various reasons (logistics,

land access, habitat alteration), it is unlikely that

M. arachnoidea will be recollected at the

location where it was found by Schlechter nearly

100 years ago. In view of the desirability of

typifying such an unusual species, a neotype

has been designated from the recent collections

obtained near Lae.

Notes: Although the Taraka collections are

referable to Marsdenia arachnoidea , there are

some minor differences between these

specimens and the one illustrated by Schlechter

(1913). In the type collection, the sepals were

noticeably longer than the corolla lobes,

whereas in the new collections they are slightly

shorter, and sometimes as small as one-half the

length of the corolla. Irrespective of this

variation, the filiform sepals are highly

distinctive and quite unlike those in any other

species of Marsdenia. The single leaf depicted

in Schlechter’s figure is also much longer and

with more lateral veins (possibly up to 15,

versus 9 or 10) than in the specimens at hand.

These distinctions may warrant future

recognition at the infraspecific level, but such

determination will require examination of

specimens from the original locality.

Marsdenia arachnoidea has morphological

affinities to M. hemiptera H.Rchb. and

M. jensenii PI.Forst. from Australia, and to

M. brunnea PI.Forst., M grandis PI.Forst. and

M. lorea S.Moore from New Guinea. The

diagnostic differences between these species

are discussed in Forster (1995,1997).

Conservation status: Based on existing

information, the conservation outlook for

M. arachnoidea is precarious. Morobe

Province is probably the most comprehensively

collected area inNew Guinea (Conn 1994; Johns

1995) and this rather spectacular species having

been collected only thrice in nearly 100 years

indicates that it is genuinely rare, rather than

merely overlooked. Shortly after the neotype

was obtained, the collection site for Takeuchi

& Ama 16667 was destroyed by forest-clearing

in a process recalling the discovery of

Madangia inflata PI.Forst., D.J.Liddle &

I.M.Liddle from a similarly impacted area (Forster

et al. 1997). Forest removal by subsistence

agriculture and industrial logging is a serious

and ongoing process in many parts of PNG.

A disproportionate number of asclepiads

from New Guinea are under collected and

consequently poorly known. This is exemplified

by the startling fact that twenty-one of the fifty

species of Papuasian Marsdenia that are

currently recognised (Forster 1996) are

represented by single collections (Forster 1995).

As is often the case with relatively

inconspicuous taxa, it is likely that additional

populations will be eventually discovered for

the single-record plants (Forster 2000).

Although the conservation status of M.

arachnoidea is undoubtedly ‘Data Deficient’

(IUCN 2001), the recently documented

occurrences can be provisionally considered

as ‘Critically Endangered’ under the Criterion

B2 as the area of occupancy is under 10 km 2

with one extant location (a) and there is a

continuing decline in the area, extent and/or

quality of habitat (b(iii)) and of the number of

locations or subpopulations (b(iv)).

Etymology: The specific epithet is derived from

the Latin arachnoideus (cobwebby), a probable

allusion by Schlechter to the appearance of the

flowers and their superficial similarity to species

of Brachystelma R.Br. ex Sims from South Africa

and Papua New Guinea with which he was

familiar (Schlechter 1895,1913; Nicholas 1992;

Forster 1994). A number of Brachystelma

species (e.g. B. dinteri Schltr., B. elongatum

(Schltr.) N.E.Br., B. glabriflorum (F.Muell.)

Schltr., B. longifolium (Schltr.) N.E.Br. and B.

meyerianum Schltr.) have a dense indument of

trichomes on adaxial corolline surfaces. This

indument can impart a ‘fuzzy’ or ‘cobwebby’

appearance to the flowers.

Forster & Takeuchi, Rediscovery of Marsdenia arachnoidea

149

Acknowledgements

Thanks to Will Smith (BRI) for preparing the

illustrations and to Demas Ama and Ephraim

Morris for assistance with the fieldwork.

References

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Papua and New Guinea. The Library,

University of Papua and New Guinea: Waigani.

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results of the Schlechter-Lauterbach

Commemorative Expedition 1989-90: studies

in the phylogeny of the Australasian

Orchidaceae. Australian National Botanic

Gardens, Occasional Publication 11.

Australian National Botanic Gardens: Canberra.

Conn, B.J. (1994). Documentation of the flora of New

Guinea. In C.-I. Peng & C.H. Chou (eds.).

Biodiversity and Terrestrial Ecosystems, pp.

122-156. Academia Sinica Monograph Series

No. 14. Institute of Botany: Taipei.

Department of District Administration. (1968). Village

Directory. Territory of Papua and New Guinea.

G.W. Reid, Acting Govt. Printer: Port Moresby.

Dow, D.B. & Davies, H.L. (1964). The geology of the

Bowutu Mountains, New Guinea. Australia

Bureau Mineral Resources, Geology and

Geophysics Report 75: 1-31.

Forster, PI. (1994). Type collections of African

Asclepiadaceae in the National Herbarium of

Victoria (MEL). Muelleria 8: 141-149.

Forster, P.I. (1995). Circumscription of Marsdenia

(Asclepiadaceae: Marsdenieae), with a revision

of the genus in Australia and Papuasia.

Australian Systematic Botany 8: 709-933.

Forster, P.I. (1996). A checklist of the Asclepiadaceae

of Papuasia. Science in New Guinea 22: 15-

22 .

Forster, P.I. (1997). Marsdenia jensenii

(Asclepiadaceae: Marsdenieae), a new species

from north-eastern Queensland. Austrobaileya

5: 59-61.

Forster, P.I. (2000). Rediscovery of Dischidia

torricellensis (Schltr.) PI.Forst., an unusual

epiphytic asclepiad from New Guinea.

Austrobaileya 5: 725-728.

Forster, P.I., Liddle, D.J. & Liddle, I.M. (1997).

Madangia inflata (Asclepiadaceae:

Marsdenieae), a new genus and species from

Papua New Guinea. Austrobaileya 5: 53-57.

Hammermaster, E.T. & Saunders, J.C. (1995a). Forest

Resources and Vegetation Mapping of Papua

New Guinea. PNGRIS Publ. no. 4. CSIRO and

AIDAB: Canberra.

Hammermaster, E.T. & Saunders, J.C. (1995b). Forest

Resources and Vegetation Mapping of Papua

New Guinea. 1:250,000 vegetation map

overlays separately issued as working copies

to PNGRIS Publ. no. 4. CSIRO and AIDAB:

CSIRO.

Iucn (2001). IUCN Red List Categories and Criteria

Version 3.1. IUCN - The World Conservation

Union: Gland (Switzerland).

Johns, R.J. (1995). Malesia - an introduction. Curtis’s

Botanical Magazine n.s. 12: 52-62.

Nicholas, A. (1992). The Asclepiadaceous works of

Rudolf F. Schlechter (1872-1925).

Will denow ia 22: 215-264.

Schlechter, R. (1895). Beitrage zur Kenntnis

sudafrikanischer Asclepiadaceen. Botanische

Jahrbiicher Systematische 20, Beiblatt zu den

Botanischen Jahrbiichen 51: 1-56.

Schlechter, R. (1911-14). Die Orchidaceen von

Deutsch-Neu-Guinea. Fedde Repertorium

Specierum Novarum Regni Vegetabilis, Beiheft

1: 1-1079. English translation 1982, D.F.

Blaxell (ed). The Australian Orchid Foundation:

Melbourne.

Schlechter, R. (1913). Die Asclepiadaceen von Deutsch-

Neu-Guinea. Botanische Jahrbiicher fur

Systematik Pflanzengeschichte und

Pflanzengeographie 50: 81-164.

150

Austrobaileya 7 (1): 145-150 (2005)

Map 1 . Island of New Guinea. Localities for M. arachnoidea. A. The type locality (above Dschischungari) as

inferred from Schlechter’s itinerary, Morobe Province. B. Taraka, Lae subdistrict, Morobe Province.

Ludwig Leichhardt’s Australian plant collections, 1842-1847

John Leslie Dowe

Summary

Dowe, J.L. (2005). Ludwig Leichhardt’s Australian plant collections, 1842-1847. Austrobaileya

7(1): 151-163. The contribution made by Ludwig Leichhardt to Australian botany is examined

with reference to his collections and their use as types. About 2800 specimens collected by

Leichhardt were located, of which at least 78 specimens have been designated as types. Leichhardt’s

records of species distribution, and manuscript names, are also examined. Forty-nine eponyms,

named for Leichhardt, are listed.

Key Words: Australian botany, Ludwig Leichhardt, Overland Expedition, botanical exploration

J.L. Dowe, Australian Centre for Tropical Freshwater Research, James Cook University, Douglas,

Queensland 4811, Australia.

Introduction

The plant collections and botanical

observations that were made by Australia’s

colonial explorers contributed to the major

taxonomic and systematic treatments produced

at the time. Works such as Flora Australiensis

(Bentham 1863-1878) and Fragmenta

Phytographiae Australiae (Mueller 1858-1882),

were predominantly based on collections by

explorers such as Mitchell, Sturt, Gregory, Giles,

and Leichhardt, among others, as well as

botanists, such as Banks & Solander, Brown,

Caley, Dietrich, C. Moore and Cunningham

(Maiden 1908; Blake 1955; Hall 1978; Orchard

1999).

Ludwig Leichhardt

The exploration activities of [Friedrich Wilhelm]

Ludwig Leichhardt (b. Prussia, 23 Oct. 1813, d.

central Australia? 1848) have made him one of

Australia’s most memorable and controversial

colonial figures (Jack 1921; Neumayer 1944;

Chisholm 1955; Allingham 1977; Desmond 1977;

Dalton 1986; Roderick 1988; Barker & Barker

1990; Priessnitz 1991; Levitus 1995; Peam2001).

His contribution to botany, though well

recognised by some (Maiden 1908; Moore 1920;

Blake 1955; Hall 1978; Webster 1980; Filson

1992; Orchard 1999; Short 2003), has been oddly

disparaged by others who have chosen to

emphasise his enigmatic and controversial

status (Chisholm 1955; Webb 2003). This paper

aims to provide an overview of his collecting

Accepted for publication 12 July 2005

activities, the use of his specimens as types,

and his contribution to botany in Australia.

Methods

Citations and other references to collections

made by Leichhardt were located in Flora

Australiensis (Bentham 1863-1878), and

Fragmenta Phytographiae Australiae

(Mueller 1858-1882), as well as other relevant

works. The determination and updating of

nomenclature were primarily achieved through

access to Henderson (2002), APNI (2004), IPNI

(2004), and published volumes in the Flora of

Australia series. Unpublished Queensland

Herbarium data, based on Henderson (2002)

were designated as the standard in the event of

conflicting nomenclature. Literature pertaining

to Leichhardt’s Overland Expedition was

examined and parts relevant to vegetation or

botany were appraised and where appropriate

were used as references within the text. Searches

for Leichhardt’s plant specimens were done in

B, BM, BRI, E, HO, K, MEL, NSW and P

Herbarium acronyms are according to Holmgren

etal. (1990).

Results

Arriving in Australia in February 1842 from

Germany where he received an education in the

natural sciences at Gottingen and Berlin

Universities, Leichhardt undertook three major

expeditions. The first was the Overland

Expedition from the Darling Downs to Port

152

Austrobaileya 7 (1): 151-163 (2005)

Essington, Oct. 1844-Dec. 1845 (Leichhardt

1847b). The second was an unsuccessful

attempt to cross Australia from east to west,

which commenced in Brisbane in December

1846, but was compelled to return because of

illness and the loss of stock after six months

(Bunce 1859; Sprod 1989). The third, also an

attempt to cross Australia from east to west,

commenced in March 1848, and resulted in the

disappearance of Leichhardt and his entire

party, of which no remains have ever been

found (Connell 1980).

Of the specimens collected by

Leichhardt during his five years of activity in

Australia, the number would have been

considerably greater had Leichhardt not been

forced to abandon most of his plant and

geology specimens during the Overland

Expedition because of the drowning of the pack-

horses, and other unfortunate incidents. In

correspondence to his brother-in-law C.

Schmalfuss [24 January 1846], Leichhardt

reported that, during the Expedition, he had to

“... .burn about 3000 driedplants. ...” whilst in

correspondence to G. Durando of the Paris

herbarium [20 May 1846] he wrote “....As my

collection increased, I surrounded the different

packages with green hide, which when dry,

formed a fine box round them, and protected

them from hard usage to which they were

exposed. ...the time came when I had to open

all my fine green hide boxes, to make a poor

choice of the dried plants, and to throw the

greatest number of them away unable to carry

them any farther, as four of my pack horses

drowned, and the means of carrying my

collections of plants and geological specimens

were consequently destroyed. I fully lost 4-5000

specimens. There are however still some very

interesting remnants...” (Politzer 1944;

Aurousseau 1968). Furthermore, additional

losses were incurred when a bullock carrying

his remaining specimens “plunged into a deep

pond, where he was quietly swimming about

and enjoying himself, whilst I was almost crying

with vexation at seeing all my plants

thoroughly soaked' (Leichhardt 1847b, p. 469).

The majority of Leichhardt’s extant specimens

were collected in locations that are now

southeast Queensland, central and northern

New South Wales, and the Port Essington area

in the Northern Territory (Webster 1980).

Leichhardt published his diary of the Overland

Expedition in the form of a journal that was edited

by geographer and surveyor P.P. King

(Leichhardt 1847b). Hooker (1860, p. cxxi) noted

that the journal, at the time, was “....by far the

fullest published detailed account of the

tropical vegetation of the interior of Australia

that we possess ”.

At the time that Leichhardt was engaged

in his exploration activities, systematic studies

of the Australian flora were relatively

rudimentary. Leichhardt’s primary references

were those of Brown (1810,1830) and volumes

1-7 of Candolle and Candolle (1823-1839)

(Aurousseau 1968). Those works dealt primarily

with the plants of the coastal and near-coastal

regions, of settled areas and where exploration

had previously occurred. The floras of central

and northern Australia were yet to be fully

appreciated and studied, and it is in this context

that Leichhardt framed his scientific attention.

Apart from Leichhardt’s specimens

subsequently being used in the formulation of

taxonomic descriptions, Leichhardt’s botanical

observations in the journal of the Overland

Expedition were often quoted. For example,

Bentham (1864, p. 297) noted in his description

of Erythrophleum chlorostachys (F.Muell.)

Baill., as “also in Leichhardt’s collection, and

said to be his ‘leguminous Iron-bark tree ”’

and Leichhardt’s original: “a leguminous tree,

with the darkfissured bark of the Ironbark ....”

(Leichhardt, 1847b, p. 242). Mention was made

by Bentham (1866, p. 408) with regards to

Gardenia edulis F.Muell., [= G vilhelmii Domin]

of the Gilbert and Lynd Rivers, “....the

“Breadfruit-tree” of Leichhardt ....”, though

correctly the ‘ Bread tree ’ in the Journal

(Leichhardt 1847b, p. 273). Mueller (1866a), in

his taxonomic account of Xylomelum

scottianum (F.Muell.) F.Muell., suggested that

this was probably the species noted by

Leichhardt in his Journal. The journal entry of

May 22,1845, in the vicinity of Separation Creek

was: “.... a Xylomelum, with smooth and smaller

seed-vessels than those of X. pyriforme”

(Leichhardt 1847b, p. 263). Mueller, in naming

Bauhinia leichhardtii F.Muell., noted that

Leichhardt “referred to it repeatedly in his

joumaT (Mueller, 1859, p. 51), and with reference

to Loranthus grandibracteus, Mueller (1860,

Dowe, Leichhardt’s Australian plant collections

p. 150) noted that this species was “ cursorily

mentioned in the diary of that lamented

traveller ”.

It was beyond the scope of this paper to

both confirm the existence and whereabouts of

all Leichhardt’s collections, and to determine

the type status of those specimens that have

been designated as types. However, herbarium

searches yielded about 2800 specimens, which

most likely form the bulk of Leichhardt’s

collections. Of these, at least 78 specimens have

been designated as types (Appendix).

Leichhardt published no taxonomic

work, although he provided accounts that were

published within the botanical domain,

including an overview of his botanical activities

of 1842-1844 (Leichhardt 1845), letters sent to G.

Durando of Paris, (Leichhardt 1846), lecture

transcripts (Leichhardt 1847a) that contained

proposed species names (Table 1), and his

Journal of an Overland Expedition (Leichhardt

1847b). These publications outlined the extent

of Leichhardt’s travels and his plant collecting

activities. As well, many minor articles were

published in newspapers and magazines in

Australia, England and Germany (Aurousseau

1968; Hoare 1981).

Heward (1847) provided an account of the

botany of Leichhardt’s Overland Expedition

153

which drew primarily from Leichhardt’s paper

published in the Tasmanian Journal of Natural

Science (Leichhardt 1847a), and it is reasonable

to assume, based on Heward making no

reference to specimens, that he did not examine

any Leichhardt specimens collected during the

Overland Expedition. More recently, Jackes

(1990) provided an assessment of Leichhardt’s

botany for the Lynd, Mitchell and Nassau Rivers

area, based on Leichhardt’s descriptions and

names. That account identified species as noted

by Leichhardt in his Journal.

With further regard to taxonomy,

Leichhardt was cited as author of Kallstroemia

minuta (Leichh. exBenth.)Engl. (Engler 1897;

Barker 1998), with Engler’s transfer of

Tribulopsis minutus Leichh. ex Benth. to

Kallstroemia. The type for Tribulus minutus

Leichh. ex Benth. is “ Queensland (?).

Leichhardt’s Expedition ” and is extant in K.

Engler’s work is an illegal combination and he

should have cited this taxon as Tribulus minutus

Leichh. ex Benth., not as a Tribulopsis , as

Tribulus is the genus in which Bentham (1863)

formerly established the taxon. The reason that

Bentham originally cited Leichhardt as the

author is not documented, and it could be

surmised that Leichhardt may have provided

some notes about the species. Bentham’s (1863)

author citation of ‘ Leichh. in Herb. F. Muell. ’

Table 1. Plant names used by Leichhardt (1845, 1847), and which have not been

taxonomically accepted and are designated nomina nuda.

Taxon

Reference and citation

Acacia

equisetifolia

Joum. Overt Exped. 356 (1847) "In a patch of rusty-gum forest we found Acacia

equisetifolia.... ”

Tas. Joum. Nat. Sc. 3: 97 (1847) "Grevillea ceratophylla, and Acacia equisetifolia, were

first met with in latitude 19° 19'."

Casuarina

villosa

Joum. Overt Exped. 49 (1847) "a new species of forest oak, which deserves the name

Casuarina villosa, for its bark looks quite villous"

Grevillea

lanceolata

Tas. Joum. Nat. Sc. 3: 97 (1847) "Grevillea lanceolata, a new species with broad

lanceolate leaf, thus named by me, showed itself first at the Suttor, where it was growing

on a light sandy soil with Pandanus spiralis."

Zamia australis

London J. Bot. 4: 279 (1845) "The grass tree (Xanthorrhoea) gives a peculiar character

to many spots, and Zamia australis is no less striking."

154

Austrobaileya 7 (1): 151-163 (2005)

suggests this. To my knowledge, the only

published reference to Tribulus made by

Leichhardt is that in which he noted the

presence of “a species of Tribulus ” at Comet

Creek (Leichhardt 1847b, p. 87). Barker (1998, p.

32) stated that the type for T. minutus was

“[Queensland, Leichhardt District], before Canal

[pastoral run] and afterwards, dat ., L.

Leichhardts.n. ( syn. MEL s.n.,p.p., excluding

lower LHS specimen); without locality,

Leichhardt’s Expedition, s.dat., Herb. Mueller

{syn. K-Herb. Hooker, type seen photographs

in the Eichler manuscripts)”. Furthermore,

Barker (1998) indicated that there was a

specimen in MEL {Leichhardt 26, MEL) that

was annotated by Leichhardt as “Tribulus

parviflorus mi hr from Bokkara Flats, 3 January

1847. Barker placed this specimen under

Tribulus micrococcus Domin.

Leichhardt provided material for an

illustration of the fungus Aseroe rubra Labill.

(Phallaceae) that was published in an account

of the fungi in the Hooker collection (Berkeley

1844). Labillardiere (1806) had earlier described

the fungus. Leichhardt collected the specimen

in Sydney in 1842. With regards to examination

of specialised collections, Mueller (1866b)

provided a summary of the ferns collected by

Leichhardt, and Filson (1992) noted that Mueller

sent to European researchers a small number of

lichen specimens that were collected by

Leichhardt.

Conclusion

The contribution made by Leichhardt toward

the development of botany in Australia is

acknowledged in a number of ways. Mueller,

Bentham and others noted over 500 Leichhardt

collections in works, about 2800 specimens have

been located in herbaria, and at least 78

specimens have been designated as types. In

relative terms this is a high proportion of types

to numbers collected. Leichhardt’s name is

commemorated in at least 49 taxa (Table 2), of

which about half are currently in use. In addition,

Leichhardt’s journal provided valuable

distribution information that was subsequently

used by taxonomists in various treatments.

Leichhardt’s descriptions of the potential

pastoral value of areas that he passed through

were used in determining patterns of settlement

and development of pastoral and mining

industries.

Acknowledgments

I would like to thank the following people who

contributed in many ways to this paper: Tony

Bean (BRI), Helen Cohen (MEL), Barry Conn

(NSW), Roberta Cowan (ABLO-Kew, 2003),

Marco Duretto (HO), Rod Fensham (BRI), Paul

Forster (BRI), Catherine Gallagher (MEL), Alex

George (ABLO-Kew, 2005), Ken Hill (NSW),

Ailsa Holland (BRI), Betsy Jackes (JCT), Bob

Makinson (NSW), Les Pedley (BRI), Chris

Quinn, Annette Wilson (ABLO-Kew, 2004) and

Peter Wilson (NSW).

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Leichhardt. Cambridge University Press:

London.

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7. L.Reeve & Co.: London.

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Dowe, Leichhardt’s Australian plant collections

155

Table 2. Taxa named for Ludwig Leichhardt. Currently accepted taxa honouring Leichhardt

indicated in bold

Name honouring Leichhardt

Current Accepted Name for taxon

Acacia decurrens var. leichhardtii Benth

Acacia oshanesii F.Muell. & Maiden

Acacia leichhardtii Benth

no change

Acacia mollissima var. leichhardtii F.MuelL ex

Maiden

Acaciapubescens (Vent.) RBr.

Alsophila leichhardtiana F.Muell.

Cyathea leichhardtiana (F.Muell) Copel.

Amanoa leichhardti Bail!

Bridelia leichhardtii Bail! ex MulLArg.

Anthocercis leichhardtii F.Muell.

Duboisia leichhardtii (F.Muell.) F.Muell.

Aristida leichhardtiana Domin

No change

Bauhinia leichhardtii F.Muell.

Lysiphyllum cunninghamii (Benth) de Wit

Carex inversa RBr. var. leichhardtii Boeck.

Carex inversa RBr.

Chorizema leichhardtii F.Muell.

Isotropis filicaulis Benth

Clematis aristata var. leichhardtiana Kuntze

No change

Commersonia leichhardtii Benth

No change

Datura leichhardtii F.Muell. ex Benth.

No change

Dendmhyprmm leichardtii A. Jaeger

Camptochaete leichhardtii (A.Jaeger) Broth

Dicranum leichhardtii Hampe

Dicranoloma leichhardtii (Hampe) Watts & White!

Digitaria macractinia subsp. leichhardtiana Henrard

No change

Eriostemon leichhardtii F.Muell.

Halfordia kendack (Montrouz.) Guillaumin

Eriostemon myoporoides var. leichhardtii Benth.

Philotheca glasshousiensis (Domin) P.I.Forst.

Eriostemon trachyphyllus var. leichhardtii Benth

Philotheca glasshousiensis (Domin) P.I.Forst.

Eucalyptus leichhardtii F.M.Bailey

Corymbia leichhardtii (F.M.Bailey) K.D.Hill &

L. A. S. Johnson

Euphoria leichhardtii Benth

Dimocarpus longan Lour.

Flindersia leichardtii C.D.C.

Flindersia bennettiana Benth

Euphoria leichhardtii var. hebepetala Benth

Arytera foveolata F.Muell

Grevillea leichardtii S.Moore

Grevilleapungens RBr.

Halfordia leichhardtii F.Muell. ex Guillaumin

Halfordia kendack (Montrouz.) Guillaumin

Harpullia leichhardtii F.Muell. ex Benth.

No change

Hypnum leichardtii Hampe

Camptochaete leichhardtii (Hampe) Broth.

Kunzea opposita var. leichhardtii Byrnes

No change

Leichhardtia F.Muell.

Phyllanthus L.

Leichardtia R Br.

Marsdenia R Br.

Leichhardtia T. Shepard

Callitris Vent.

Leichardtia australis RBr.

Marsdenia australis (RBr.) Druce

156

Austrobaileya 7 (1): 151-163 (2005)

Table 2 (continued) . Taxa named for Ludwig Leichhardt. Currently accepted taxa honouring

Leichhardt indicated in bold

Name honouring Leichhardt

Current Accepted Name for taxon

Lobelia leichhardii E.Wimm.

No change

Livistona leichhardtii F.Muell.

Livistona humilis R.Br.

Macropteranthes leichhardtii F.Muell ex Benth.

No change

Marsdenia leichhardtiana F.Muell.

Marsdenia australis (R.Br.) Druce

Morinda leichhardtii F.Muell.

No change

Najas leichhardtii Magnus

No change

Neckera leichhardtii Hampe

Neckera pennata Hedw.

Parsonsia leichhardtii F.Muell.

No change

Prostanthera leichhardtii Benth.

Prostanthera ringens Benth.

Psoralea leichhardtii F.Muell

Indigofera glandulosa Willd.

Rotala occultiflora var. leichhardtii Koehne

Rotala occultiflora Koehne var. occultiflora

Rubus moorei var. leichardtianus Domin

Rubus moorei F.Muell. var. moorei

Sarcocephalus leichhardtii F.Muell.

Nauclea orientalis (L.) L.

Unona leichardtii F.Muell.

Melodorum leichhardtii F.Muell.

Urera leichhardtiana Wedd.

Dendrocnide photinophylla (Kuntil) Chew

Urostigma leichhardtii Miq.

Ficus platypoda var. leichhardtii (Miq.) R.J.F.Hend.

Vitex leichhardtii F.Muell

Gmelina leichhardtii F.Muell.

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People and plants in Australia , pp. 183-219

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Marshall, Hamilton, Kent & Co. Ltd.: London.

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Novae Hollandiae Plantarum Specimen 2:

124.

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Holland, by Dr. Ludwig Leickhardt, 1842-44;

extracted from his letters to M. G. Durando,

of Paris. London Journal of Botany 4: 278-

291.

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Extracted from a letter from Dr. Ludwig

Leickhardt of New Holland to M. Durando, of

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660.

Leichhardt, L. (1847a). Lectures on the geology, botany,

natural history, and capabilities of the country

between Moreton Bay and Port Essington.

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81-113.

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expedition in Australia, from Moreton Bay to

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Boone: London.

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Proceedings of the Royal Society of New South

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plants. Transactions of the Philosophical

Institute of Victoria 3: 40-63.

Mueller, F. (1860). Essay on the plants collected by

Mr. Eugene Fitzalan, during Lieut. Smith’s

expedition to the estuary of the Burdekin.

Government Printer: Melbourne.

Mueller, F. (1866a). Xylomelum scottianum.

Fragmenta phytographiae Australiae 5: 174.

Government Printer: Melbourne.

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continentalem in habitantium imprimis de

earum distributione agens breviarium.

Fragmenta phytographiae Australiae 5: 111-

142. Government Printer: Melbourne.

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Leichhardt’s letters from Australia during the

years March 23, 1842, to April 3, 1848, pp.

71-95. Pan Publishers: Melbourne.

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103.

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medici in botanicis: Australian Flora and the

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Politzer, L.L. (1944). Dr. Ludwig Leichhardt’s letters

from Australia during the years March 23,

1842, to April 3, 1848. Pan Publishers:

Melbourne.

Priessnitz, H. (1991). The ‘Vossification’ of Ludwig

Leichhardt. In D. Walker & J. Tampke (eds.).

From Berlin to the Burdekin: the German

contribution to the development of Australian

science, exploration and the arts, pp. 196-

217. New South Wales University Press:

Kensington.

Roderick, C. (1988). Leichhardt the dauntless explorer.

Angus and Robertson: North Ryde.

Short, P. (2003). In pursuit of plants. University of

Western Australia Press: Crawley (WA).

Sprod, D. (1989). Proud intrepid heart: Leichhardt’s

first attempt to the Swan River 1846-1847.

Blubber Head Press: Hobart.

Webb, J.B. (2003). The botanical endeavour: journey

towards a flora of Australia. Surrey Beatty &

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Webster, E.M. (1980). Whirlwinds in the plain: Ludwig

Leichhardt - friends, foes and history.

Melbourne University Press: Melbourne.

158

Austrobaileya 7 (1): 151-163 (2005)

Appendix: Specimens, collected by Ludwig Leichhardt in Australia, that have been designated as

types, with updated nomenclature, original literature citations and herbaria where specimens are kept. If

specimens have been selected as lectotypes or have become lectoparatypes due to lectotypification this

has not been indicated.

Akaniaceae

Cupania lucens F.MuelL, Fragm. 3: 44 (1862) =

Akania bidwillii (Hogg.) Mabb., Plant Book 707

(1990)

Ad sinum Moreton Bay;

Leichhardt.

cited but not

located [not at K or

BM]

Anacardiaceae

Rhus viticifolia F.Muell. ex Benth., FI Austral. 1: 489

(1863) = Rhus tomentosa L., Sp.Pl. 266 (1753)

Queensland (?) Leichhardt

Apocynaceae

Parsonsia leichhardtii F.MuelL, Fragm. 6: 128 (1868)

Ad sinum Wide Bay;

Leichhardt

MEL

Asteraceae

Cassinia theodori F.MuelL, Fragm. 5: 148 (1866)

In parietibus phonolithicis ad

originem fluvii Gwydir; Dr.

Ludw. Leichhardt

Asteraceae

Helichrysum diotophyllum F.MuelL, Fragm. 5: 150

(1866) = Ozothamnus diotophyllus (F.Muell.) Anderb.,

Op. Bot. 104: 89 (1991)

In A ustralia orientali

subtropica ad flumen

Dogwood Creek; Leichhardt

et Bunce

MEL, K

Boraginaceae

Cordia ixiocarpa F.Muell, Fragm. 1: 59 (1858) =

Cordia myxa var. ixiophylla (F.MuelL) Domin, Biblioth.

Bot. 89(4): 1097 (1930)

In collibus humilibus ad

flumen Gilbert. Leichhardt,

Mueller.

cited but not

located[not at K or

BM]

Caesalpiniaceae

Cassia concinna Benth., FI. Austral. 2:291 (1864) =

Chamaecrista concinna (Benth) Pedley, FI. Aust. 12:

143, 196 (1998)

N.S. Wales. Mount Flinders,

Leichhardt

MEL

Campanulaceae

Lobelia leichhardii E.Wimm, Das Pflanzenreich 107:

584 (1953)

Australien: Morton Bay, 1845

(Leichhard! - Hb. Paris)

Casuarinaceae

Casuarina equisetifolia var. micmcarpa F.Muell.,

Fragm. 6: 17 (1867) - Casuarina cunninghamiana

Miq. subsp. cunninghamiana , Revisio critica

Casuarinarum 56 (1848)

ad Glendon (Leichh.).

MEL

Chenopodiaceae

Atriplex stipitata Benth, FI. Austral. 5: 168 (1870)

N.S. Wales, also in Leichhardt's

collection

MEL

Chrysobalanaceae

Parinari nonda F.Muell ex Benth, FI. Austral. 2: 426

(1864)

N.Australia. From the Upper

Lind to Van Diem an's river,

Gulf of Carpentaria,

Leichhardt

MEL

Combretaceae

Macropteranthes leichhardtii F.Muell. ex Benth., FI.

Austral. 2: 505 (1864)

Queensland. Ruined Castle

Creek Leichhardt.

MEL, K

Combretaceae

Terminaliaplatyptera F.Muell., Fragm. 2: 151 (1861)

In plantiebus terrae Arnhem's

Land et circum sinum

Carpentariae.

MEL

Cyathaceae

Alsophila leichhardtiana F.Muell., Fragm. 5: 53 (1865)

= Cyathea leichhardtiana (F.Muell.) Copel., Philipp. J.

Sc., C 6: 360 (1911)

Moreton Bay, Fern-tree Creek,

Bunya Mountain, L.

Leichhardt.

MEL

Dowe, Leichhardt’s Australian plant collections

159

Appendix (continued)

Cyperaceae

Carex inversa var. leichardtii Boeck., Linnea 39: 70

(1875) = Carex inversa R.Br., Prodr. 242 (1810)

Nova Holland., Paramatto

(Leichardt)

MEL

Cyperaceae

Cladium scleroides F.Muell., Fragm. 9: 12 (1875)

=Exocarya scleroides (F.Muell.) Benth., leones

Plantarum 13: t.1206 (1877)

In silvis Araucariae Bidwilli

prope fluvios Burnett's et

Dawson 's River, Leichhardt

MEL

Epacridaceae

Styphelia exolasia F.Muell., Fragm. 6: 34 (1867) =

Leucopogon exolasius (F.MuelL) Benth., FI. Austral. 4:

217 (1868)

In vicinia pagi Camden Dre

Ludiv. Leichhardt anno 1843

detecta

Epacridaceae

Stypheliapleiosperma F.MuelL, Fragm. 6: 41 (1867) =

Leucopogonpleiospermus (F.Muell) Benth., FI.

Austral. 4: 207 (1868)

In Australia orientali (loco

speciali non indicato),

Leichhardt

Euphorbiaceae

Bridelia leichhardtii Bail! ex MiilLArg. in Candolle, A.P.

de & Candolle, A.L.P.P. de (ed.), Prodromus 15(2): 499

(1886)

In Nova Hollandia ad

Moreton Bay (Leichhardt in

hb. Mus. Paris).

Euphorbiaceae

Claoxylon australe Baill. ex MiilLArg., Etude Generate

du Groupe des Euphorbiacees : 493 (1858)

herb. Mus. Nouvelle-

Hollande. Coll. Leichhard,

1845

Euphorbiaceae

Croton stigmatosus F.Muell., Fragm. 4: 140 (1864)

ad sinum Moreton Bay, Dr

Leichhardt

G-DC, MEL, P

Euphorbiaceae

Hemecyclia lasiogyna F.Muell., Fragm. 4: 119 (1864)

= Drypetes deplanchei (Brongn. & Griseb.) Merr., J.

Arnold Arbor. 32: 199 (1951)

In locis fontanis ad portum

Essingtoni; Leichhardt

MEL

Euphorbiaceae

Micrantheum ericoides var. juniperinum Grun., Das

Pflanzenreich 58: 25 (1913)

Queensland (Leichhardt!)

MEL

Fabaceae

Bossiaea brownii Benth., FI. Austral. 2: 163 (1864)

Queensland, also in

Leichhardt's Collection

MEL

Fabaceae

Chorizema leichhardtii F.Muell, Fragm. 4: 20 (1863)

= Isotropis filicaulis Benth., FI. Austral. 2: 40 (1864)

In clivis arenoso- rupestribus

ad sinum marinum Wide Bay;

Dr. Ludw. Leichhardt

MEL

Fabaceae

Flemingia parviflora Benth., FI. Austral. 2: 269 (1864)

Queensland. Lynedoch valley,

Leichhardt

MEL

Fabaceae

Hovea heterophylla A.Cunn ex Hook.f f decipiens

Domin, Biblioth. Bot. 89: 129 (1928) = Hovea linearis

(Sm) R Br., Hortus Kewensis Edn. 2, 4: 275 (1812)

sandstone hills towards

Brisbane, Leichhardt

MEL

160

Austrobaileya 7 (1): 151-163 (2005)

Appendix (continued)

Fabaceae

Pultenaea paleacea Wild. var. obtusata Benth., FI.

Austral. 2: 116 (1864) = Pultenaea paleacea Wild. var.

paleacea, Trans. Linn. Soc. London 9: 246 (1808)

between Suggerah and Lake

Macquoy, Leichhardt

MEL

Fabaceae

Sophora fraseri Benth., FI. Austral. 2: 274 (1864)

Queensland. Mur rum - Mur rum

Creek, Leichhardt

MEL

Fabaceae

Swainsona brachycarpa Benth, FI. Austral. 2: 217

(1864)

Queensland. Condamine river

and Darling Downs,

Leichhardt

MEL, NSW

Fabaceae

Swainsona oroboides F.Muell. ex Benth., FI. Austral. 2:

222(1864)

N.S. Wales, head of Gwydir,

Leichhardt

MEL

Fabaceae

Templetonia muelleri Benth, FI. Austral. 2: 169 (1864)

= Templetonia stenophylla (F.MuelL) J.M. Black, Flora

of South Australia 4: 304 (1929)

Queensland. Wide Bay,

Bidwill, Leichhardt

MEL

Fabaceae

Tephrosia filipes Benth., FI. Austral. 2: 208 (1864)

Queensland. Erythrina Creek,

Leichhardt

MEL

Fabaceae

Tephrosia juncea Benth, FI. Austral. 2: 208 (1864)

Queensland, also in

Leichhardt's collection

MEL

Fabaceae

Tephrosia purpurea Pers. var. rufescens Benth., FI.

Austral. 2: 210 (1864) — Tephrosia rufula Pedley,

Austrobaileya 1: 38 (1977)

Archer's Hill, Leichhardt

MEL, NSW

Mimosaceae

Acaciapolybotrya var foliolosa Benth., FI. Austral. 2:

414 (1864) = Acacia glaucocarpa Maiden & Blakely,

Proc. Roy. Soc. Queensland 38: 120 (1927)

Queensland, limestone hills,

Leichhardt

MEL

Mimosaceae

Acacia undulifolm var. humilis Benth., FI. Austral. 2:

356 (1864) =Acacia hubbardiana Pedley, Contr.

Queensland Herb. 4: 2 (1969)

also the Brisbane specimens

from F. Mueller and

Leichhardt

MEL

Mimosaceae

Acacia leichhardtii Benth., FI. Austral. 2: 372 (1864)

Queensland. Expedition

range, Leichhardt

K, NSW

Musci

Neckera leichhardtii Hampe, Linnaea 36: 520 (1870) =

Neckera pennata Hedw., Sp. Muse. Frond. 200 (1801)

Hab. New South Wales leg.

Leichhard

Myrsinaceae

Myrsine subsessilis F.Muell, Fragm. 4: 81 (1864) =

Rapanea subsessilis (F.Muell) Mez., Das Pflanzenreich

9: 354 (1902)

In silvis ad sinum Moreton

Bay, Dr. Leichhardt

NSW

Myrtaceae

Eucalyptus albem Miq. ex Benth, FI. Austral. 3: 219

(1867)

N.S. Wales, between Alford's

and the Range, "Box,"

Leichhardt

MEL

Myrtaceae

Eucalyptus fibrosa F.MuelL, J. Linn. Soc., Botany 3: 87

(1859)

In montibus nemorosis ad

flumen Brisbane. Anth.

aestate.

MEL

Dowe, Leichhardt’s Australian plant collections

161

Appendix (continued)

Myrtaceae

Thryptomene polyandra F.Muell, Fragm. 4: 77 (1864)

= Homalocalyx polyandrus (F.Muell) F.Muell. ex

Berth., FI. Austral. 3: 56 (1867).

In quadam parte Australiae

baud in dicat a detexit Dr.

Ludw. Leichhardt.

MEL

Najadaceae

Najas leichhardtii Magnus, Beitrage zur Kenntniss der

Gattung Najas L. \ 46, 50,52, t. 8, figs 1-8 (1870).

bei einer von Leichhardt in

Australien gesammelten

Pflanze

Phallaceae

Aseroe actinobola Corda, leones Fung, cogn .6: 23

(1854) = Aseroe rubra LabilL, Novae Hollandiae

Plantarum Specimen 2: 124 (1806).

Mr. Leichardt theilte die

Zeichnung und Beschreibung

des frischen und so

merkwurdigen Pilzes folgends

mit

Poaceae

Aristida leichhardtiana Domin, Regni Veg. 9: 551

(1911)

Queensland: Dry-beef Creek,

coll. Leichhardt.

BRI

Poaceae

Aristida vagans var. gracillima Berth., FI. Austral. 7:

563 (1878) = Aristida gracilipes (Domin) Henrad, Med.

Rijks-Herb 54: 209 (1926)

Cameroons Brush, Leichhardt

BRI

Poaceae

Festuca latispicea F.Muell, Fragm. 8: 127 (1874) =

Glyceria latispicea (F.Muell.) Berth., FI. Austral. 7:

658 (1878)

Ad Gwydir et Myall-Creek,

Leichhardt

MEL

Poaceae

Festuca loliiformis F.Muell., Fragm. 8: 128 (1874) =

Tripogon loliiformis (F.MuelL) C.E. Hubb., Bull. Misc.

Inform. 10: 448 (1934)

Ad Moreton's Bay et

Charley's Creek, Leichhardt

cited but not

located [not at K or

BM]

Poaceae

Panicum buncei F.Muell ex Berth., FI. Austral. 7: 487

(1878)

Queensland. Bokhara Flats,

Leichhardt

K, BRI

Poaceae

Sporobolus caroli Mez, Repert. Spec. Regi. Veg. 299

(1921)

Australien, Liverpool-plains

(Herb. hort. Sydney)

MEL

Pontederiaceae

Limnostachys cyanea F.Muell., Fragm. 1: 24 (1858) =

Monochoria cyanea (F.Muell.) F.Muell, Fragm. 8: 44

(1873)

In terra Arnhem's Land.

Leichhardt

K, MEL

Proteaceae

Grevillea leichardtii S.Moore, J. Linn. Soc. Bot. 45:

211 (1920) = Grevillea pungens R.Br., Trans. Linn.

Soc. London 10: 175 (1810)

N. Australia, "Sandy

scrubland west side of gulf'

[Carpentaria]; Leichardt

BM, K, MEL

Proteaceae

Grevillea leiophylla F.Muell. ex Berth., FI. Austral. 5:

471 (1870).

Queensland. Glasshouse

ranges, Moreton Bay,

F. Mueller, and probably the

same neighbourhood,

Leichhardt

MEL, NSW

Proteaceae

Grevillea rubicunda S.Moore, J. Linn. Soc. Bot. 45:

210(1920)

N. Australia, "Westward of

the Gulf [Carpentaria],

table-land of the South

Alligator"; Leichardt

162

Austrobaileya 7 (1): 151-163 (2005)

Appendix (continued)

Proteaceae

Grevillea singuliflora F.MuelL, Fragm. 6: 92 (1867)

Ad rivum Dogwood-Creek

Dogwood Creek, Leichhardt

BM, MEL

Proteaceae

Lomatia silaifolia var. induta F.Muell. ex Benth., FI.

Austral. 5: 537 (1870) = Lomatia silaifolia var.

silaifolia (Sm) RBr., Trans. Linn. Soc. London 10: 199

(1810)

Brisbane river, Moreton Bay,

Leichhardt, F. Mueller

BM, K

Ranunculaceae

Clematis aristata var. leichhardtiana Kuntze, Verb. Bot.

Vereins Prov. Brandenburg 26: 156 (1885)

Australia (11842 Leichhardt,

mus. bot. beml.); Norfolk

Island; Lord Howe's Island;

Nova Caledonia (IGillivray)

cited but not

located [not at K or

BM]

Rhamnaceae

Emmenosperma alphitonioides F.Muell., Fragm. 3: 63

(1862)

ad amnem Piri Creek; Dr

Ludw. Leichhardt

cited but not

located[not at K or

BM]

RosaceaeRubus moorei F.Muell. var. leichhardtianus

Domin, Regni. Veg. 1133 (1913) = Rubus moorei

F.Muell. var. moorei

New South Wales: From the

Creek Brush to Archers

Station, leg. Leichhardt

MEL

RubiaceaeAsperula geminifolia F.Muell., Fragm. 5: 147

(1899)

In pratis ripariis inter

flumina Dawson's River et

Brisbane's River; Leichhardt,

F.M.; nee non prope urbem

Newcastle, Leichhardt

cited but not

located[not at K or

BM]

RutaceaeEriostemon leiebardtii F.Muell Fragm 5:5

(1866) = Halfordia kendack (Montrouz.) Guillaumin,

Notulae Systematicae (Paris) 98 (1911)

Secus rivos silvaticos in

vicinia sinus marini Moreton

Bay. Dr. Ludw. Leichhardt

MEL

Rutaceae

Eriostemon trachyphyllus var. leichhardtii Benth., FI.

Austral. 1: 333 (1863) =Philotheca glasshousiensis

(Domin.) P.I.Forst., Austrobaileya 7: 178 (2005)

"From Brroa " (N.S. Wales?)

Leichhardt

MEL

Rutaceae

Flindersia leiebardtii C.DC., Monographic

Phanerogamarum 1: 731 (1878) = Flindersia

bennettiana Benth., FI. Austral. 1: 389 (1863)

In Moreton Bay (Leichardt in

herb. Mus. Par.)

Santalaceae

Choretrum candollei F.Muell. ex Benth, FI. Austral. 6:

219 (1873)

N.S. Wales. Liverpool plains,

Leichhardt

MEL

Sapindaceae

Euphoria leichhardtii Benth var. hebepetala Benth, FI.

Austral. 1: 468 (1863) = Arytera foveolata F.Muell,

Trans. & Proc. Philos. Instit. Vic. 3: 24 (1859)

"Nurrum Nurrum ",

Leichhardt (Herb. F.Muell.)

MEL

Sapindaceae

Euphoria leichhardtii Benth FI. Austral. 1: 468 (1863)

= Dimocarpus longan Lour., FI. Cochin. 233 (1790)

Queensland (?), Leichhardt

(Herb. F. Muell).

MEL

Sapindaceae

Harpullia leichhardtii F.Muell. ex Benth, FI. Austral. 1:

470 (1863)

N.Australia. Port Essington,

Leichhardt

MEL

Dowe, Leichhardt’s Australian plant collections 163

Appendix (continued)

Simaroubaceae

Ailanthnspunctata F.Muei, Fragm. 3:42 (1863] =

Pentaceras australis (F.Muell.) Benth., [Rutaceae] FI.

Austral. 1:365 (1863)

in silvis Australiae orient alis

subtropicae; e.g.

MacConnell's Brush, L.

Leichhardt

cited but not

located [not at K or

BM]

Solanaceae

Anthocercis leichhardtii F.Muell., Fragm. 6: 142 (1868)

= Duboisia leichhardtii (F.Muei) F.Muei, Syst.

Census Austral. PI. 97 (1883)

In Australia orientali

extratropica; Leichhardt

K, MEL

Sterculiaceae

Commersonia leichhardtii Benth., FI. Austral. 1: 242

(1863)

Queensland. Head of Boyd

river, Leichhardt, in Herb. F.

Muell.

MEL

Stereaceae

Thelephora leichhardtiana Lev., Ann. Sci. Nat. ser.3, 5:

148 (1846) = Stereum ostrea Blume & Nees, Epicr.

Syst. Mycol. 547 (1838)

Queensland.

Urticaceae

Hyrtanandra lythroides F.Muell., Fragm. 5: 194 (1866)

= Pouzolzia hirta (Blume) Hassk., Cat. Hort. Bogor

800 (1844)

Ad flumen Lynd's River ejus

originem versus, nec non ad

fluvium South Alligator-River

in terra tabulari; Dr.

Leichhardt.

cited but not

located[not at K or

BM]

Urticaceae

Elatostema reticulatum Wedd., Ann. Sci. Nat. Bot. ser.

4, 1: 188 (1854)

Nov. Holland. (Leichhard, pi.

exs., n.?).

NSW

Urticaceae

Urera leichardiana Wedd., Ann. Sci. Nat. ser 4, 1: 178

(1854) = Dendrocnide photinophylla (Kunth) Chew,

Gard. Bull. Singapore 21: 205 (1965)

Nov. Holland. (Leichard, pi.

exs. n. 26).

Urticaceae

Urera excelsa Wedd., Ann. Sci. Nat. ser 4, 1: 178

(1854) = Dendrocnide excelsa (Wedd.) Chew, Gard.

Bull. Singapore 21: 205 (1965)

Nova Hollandia (Leichhard,

pi. exs. n. 17).

Verbenaceae

Vitex leichhardtii F.Muei, Fragm. 3: 58 (1862) =

Gmelina leichhardth (F.Muei) Benth., FI. Austral. 5:

66 (1870).

In silvis ad amnem Myall

Creek Australiae orient alis

subtropicae; Dr. Ludwig

Leichhardt.

MEL

Zygophyllaceae

Tribulus minutus Leichh ex Benth., FI. Austral. 1: 291

(1863)

Queensland (?), Leichhardt's

Expedition.

K, MEL, NSW

Wing structure in seeds of Strangea Meisn. and

Stenocarpus R.Br. (Proteaceae)

H. Trevor Clifford 1 and Mary E. Dettmann 2

Summary

Clifford, H.T. & Dettman, M.E. (2005). Wing structure in seeds of Strangea Meisn. and

Stenocarpus R.Br. (Proteaceae). Austrobaileya 7(1): 165-170. In Stenocarpus the inner wing of

the seed derives from the mesophyll of the outer integument whereas with Strangea this wing

develops from the inner epidermis of that integument. The outer wing in both genera derives

from the outer epidermis of the outer integument. Contrary to the present circumscription of the

Subtribe Stenocarpinae in neither genus is the inner integument involved in wing production. An

emended diagnosis for the Subtribe Stenocarpinae is provided.

Key Words: Proteaceae, Stenocarpus, Strangea , seed morphology, subtribe Stenocarpinae

‘H.T.Clifford, c/o Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic

Gardens, Mt Coot-tha Rd, Toowong, Queensland 4067, Australia

2 M.E. Dettman, School of Integrative Biology, University of Queensland, Queensland 4072, Australia

Introduction

Strangea Meisn. and Stenocarpus R.Br.

(Proteaceae) constitute the Subtribe

Stenocarpinae L.A.S.Johnson & B.G.Briggs,

within the Tribe Embothrieae Rchb. as defined

by Johnson and Briggs (1975). The Subtribe is

distinguished from the others of the

Embothrieae in possessing, amongst other

attributes, seeds with “2 adjacent wings; outer

developed from the proximal part of the funicle

and folded around the inner wing which is

developed from the inner integument”. Douglas

(1995) stated that the Stenocarpinae “possess

a fasciculate wheel-like or 1-flowered

inflorescence and a distinctive interseminal

layer”. As shown below in neither genus is the

inner wing derived from the inner integument.

The seed coats of these and other genera

of Proteaceae are formed from the outer

integument and the basal tissues of the ovule,

so the seed is extotestal (Netolitzky 1926, Comer

1976, Manning & Brits 1993). The ‘crystal layer’

identified in the seeds of many genera develops

from the inner epidermis of the outer integument

(Corner 1976, Kausik 1940, Manning & Brits

1993, Netolitzky 1926, Strohschen 1986a,b,c) and

is one cell thick. In the majority of these cells a

single crystal and/or sand of calcium oxalate is

present (Jordaan 1946a,b, Strohschen

Accepted for publication 30 June 2005

1986a,b,c). Due to the breakdown of the inner

integument the embryo sheath is formed largely

from two cuticles. The outer forms between the

two integuments and the inner between the

inner epidermis of the inner integument and the

nucellus (Strohschen 1986a,b,c). The embryo

sheath is sac-like and envelops all tissues that

he within the outer cuticle and is usually stained

with tannin especially about the hypostase. On

account of its shape it is unfortunate the term

embryo sac is not available for this structure.

In the current paper we describe the seed

morphology of Strangea and Stenocarpus and

provide an emended diagnosis for the tribe

Stenocarpinae.

Materials and methods

The seeds and fruits of Strangea linearis

(Voucher: Dettmann s.n. [AQ751259]) were

collected from Banksia aemula-Eucalyptus

signata woodland on North Stradbroke Island

and those of Stenocarpus sinuatus (Louden)

Endl. (Voucher: Clifford s.n. [AQ751257])and

Stenocarpus salignus (Voucher: Clifford s.n.

[AQ751258]) from trees cultivated in Brisbane.

Voucher specimens have been deposited in the

Queensland Herbarium.

The structure of the seed wings was

determined from a study of ovules and seeds

166

Austrobaileya 7 (1): 165-170 (2005)

made optically transparent by soaking in a dilute

solution of commercial bleach and after rinsing

thoroughly with water were stained with 1%

safranin. They were mounted in glycerine jelly

before examination in transmitted and reflected

light at magnifications up to x200.

For purposes of comparison, the

developing seed wings of several proteaceous

genera, other than Strangea and Stenocarpus,

were also investigated. These included Banksia

L.f., Buckinghamia F.Muell., Cardwellia

F.Muell., Grevillea R.Br. ex Knight, Hakea

Schrad., Lomatia R.Br., Orites R.Br. and

Xylomelum Sm.

Results

The seeds of both Strangea and Stenocarpus

derive from ovules that are initially hemitropous

but due to their differential growth and that of

the follicle, the seeds, at maturity, may appear

to be anatropous. This apparent change in the

orientation of the ovule with age may result from

the extension of the raphe relative to the distal

portion of the ovule or from the anterior end of

the sclereid strand being confused with the

hilum (Fig. 1). The vascular trace in both genera

is embedded in a strand of sclereids on the hilar

side of the wing, but towards the base of the

seed the strand reverses direction and, after

passing through the tissue surrounded by the

base of the outer integument, branches radially

to form a radiating cap over the hypostase. On

account of its change of direction the vascular

trace assumes the form of a loop or sharp hook

to which the name “hooked raphe” was given

by Johnson & Briggs (1963).

Whereas the embryo is close to the

chalazal region in the seeds of Stenocarpus, in

Strangea it is quite distant therefrom (Fig. 1).

This distinction between the genera was

commented upon by Johnson & Briggs (1963)

who noted that, “The most important difference

lies in the downward projection of the vascular

tissue in the chalazal region.

The seeds of all species of Strangea are

rectangular in outline and of those reported for

10mm 10mm 10 mm

Fig.l. Longitudinal sections of mature seeds of Strangea and Stenocarpus species in the plane which includes the

hilum, chalaza and micropyle , (diagrammatic and micropyle directed towards the base of the follicle). A. Strangea

linearis; B. Stenocarpus sinuatus; C. Stenocarpus salignus.

Abbreviations: c, chalaza; cl, crystal layer; e, embryo; es, embryo sheath; h, hilum; hy, hypostase; me, mesophyll;

mi, micropyle; oe, outer epidermis of outer integument; r, raphe; s, sclereid strand; v, vascular trace.

167

Clifford & Dettman, Seeds of Strangea and Stenocarpus

2mm

Fig. 2. Transverse sections of mature seed wings of

Strangea and Stenocarpus species, (diagrammatic). A.

Strangea linearis ; B. Stenocarpus sinuatus ; C.

Stenocarpus salignus.

Stenocarpus they are either rectangular or sub-

circular (Foreman 1995). All species of both

genera are flattened in the plane that includes

the micropyle, chalaza and hilum. In each genus

the outer integument loosely invests the

remainder of the seed in a membranous sheath.

At maturity this splits longitudinally along the

side opposite the hilum in all species of Strangea

but in Stenocarpus splits across the apical end

and the whole length of either the side opposite

the hilum or along both sides of the sheath

(Fig. 2).

With the seeds of Strangea the split

penetrates the outer integument resulting in the

formation of two membranous cloak-like open

sheaths, commonly referred to as wings. The

outer wing, which derives from the outer

epidermis of the outer integument remains

attached to the sclereid strand. Although

separate from the inner wing it is folded along

the line of the sclereid strand and is attached

only at the chalaza. The inner wing is the

crystalliferous inner epidermis of the same

integument. The embryo and its enclosing

sheath are suspended from the vascular trace

that extends from behind the base of the outer

integument to the hypostase (Fig. 1A, 3A)

thereby resembling a seed on a long funicle and

was so interpreted by Bailey (1913), Drummond

(1853) and Gardner (1933,1942).

With the seeds of Stenocarpus the

situation is more complex. Here the outer

enveloping wing, as with Strangea , is formed

from the outer epidermis of the outer integument

but the inner develops from its mesophyll, a

tissue not formed in Strangea seeds. The outer

wing may split, as in S. sinuatus, from the base

to the apex of the seed, along the side opposite

the hilum, thereby forming a laterally

compressed cowl-like sheath over the

mesophyll as in S. sinuatus. In contrast, the

outer epidermis of a S. salignus seed splits

across its chalazal end and along both the hilum

and non-hilum sides to produce two wing-like

membranes (Maiden 1904, Schwartzbarth 1905),

which are attached to a flap of tissue formed

from the apex of the integument. On the hilum

side of the seed a split develops on both sides

of the sclereid strand. In both species the crystal

layer (inner epidermis of the outer integument)

immediately surrounds the embryo sheath to

which it is tightly appressed. The mesophyll

and outer epidermis (inner and outer wings)

surround the crystal layer and are extended

beyond the micropyle to form the apical half of

the seed (Fig. 3).

Discussion

The origin of the two hinge-like wings

surrounding the embryo sheath of Strangea

seeds were misinterpreted by Gardner (1933,

1942) when he proposed Diploptera

C.A.Gardner as a new genus to accommodate

Hakea stenocarpoides Benth. Following Bailey

(1913) and Drummond (1853) he regarded the

wings as part of the fruit and the embryo sheath

with its enclosed embryo as the seed. Later he

appreciated that Diploptera and Strangea were

congeneric (Gardner 1942). Initially Strangea

was based on a single collection of S. linearis

168

Meisn., for which only one seed, “bordered on

one side with a slightly thickened nerviform

margin” (Meisner 1855, p.67), was available. As

there was no mention of a wing the seed may

have been immature. In the same paper Meisner

described Grevillea cynanchicarpa Meisn.

from Western Australia and although he was

inclined to include it in Strangea did not do so

because the specimen lacked seeds, the

morphology of which he regarded as diagnostic

of the genus.

However, the seeds had been described

by Drummond (1853), the collector of the

specimen, and Meisner quoted a portion of his

description, after translating it into Latin. It read

as follows “1-spermo, semine apice alato, samara

Fraxini imitante”. Unfortunately he left out the

remainder of the sentence which read as

follows, “each seed is enclosed by three

membranous wings; the outer and inner are

smooth and brown, not unlike the wings of the

seed of Hakea and Banksia in substance; the

middle membrane is of light brown colour and

very brittle” (Drummond 1853, p. 182). From one

who possessed only a hand lens this

description of the seed is amazingly accurate.

Furthermore, in reporting that the middle wing,

now known to be the crystal layer, is brittle,

Drummond reveals he was exceptionally

Austrobaileya 7 (1): 165-170 (2005)

observant. Some years later Mueller (1871)

transferred the species to Strangea and in an

expanded description of the genus noted the

seeds possessed two wings that were derived

from the seed coat.

In the above account stress has been

placed on the crystal layer for determining the

structure of winged Proteaceae seeds, but

Gardner (1933), when he described Diploptera,

did not appear to have been aware of its

taxonomic importance. At that date the presence

of a crystal layer in some of these seed coats

had been reported by Netolitzky (1926) who

correctly placed the crystals in the inner

epidermis of the outer integument. In the non-

winged seeds of M acadamia F.Muell. a crystal

layer had also been reported by Francis (1928)

who incorrectly interpreted it as being the inner

integument. Unaware that in some taxa of

Proteoideae the outer integument and pericarp

are fused, Filla (1926) regarded the crystal layer

of Semina Adans. emend. Salisb. seeds as part

of the inner fruit wall. Although Brough (1933)

described in detail both the anatomical and the

morphological development of Grevillea

robusta R.Br. seeds he did not report the

presence of crystals in the inner epidermis of

the outer integument. Not until the studies of

Jordaan (1946a,b) was the taxonomic

5mm

Fig. 3. Longitudinal section of mature seeds of Strangea and Stenocarpus species in the plane at right angles to

that in Fig.l (diagrammatic). A. Strangea linearis; B. Stenocarpus sinuatus; C. Stenocarpus saligna. Abbreviations

as listed in Legend to Fig. 1.

All sections taken at level x-

-x of Fig. 1. Abbreviations as listed in Legend to Fig. 1

Clifford & Dettman, Seeds of Strangea and Stenocarpus

169

significance of the crystal layer appreciated.

Although the structure of the seed-wings of

Strangea and Stenocarpus are not identical, in

neither genus is the inner integument involved

in their formation as claimed by Johnson and

Briggs (1975) and Flora of Australia 16: 360

(1995), so the following emended diagnosis of

the Subtribe Stenocarpinae is proposed:

Proteaceae, Subtribe Stenocarpinae L A S.

Johnson & B.G. Briggs emend. H.T.Clifford &

M.E.Dettmann

Conflorescentia fasciculiformis vel rotata vel

floribus paucis aliquando unico; flores ±

zygomorphi; ovula 1-2 vel 2 usque ad

numerosa; semina 1 vel multa duo et alata.

Chromosomata n— 11. Genus typicum:

Stenocarpus R.Br.

Conflorescence of fascicles or wheel-like or

flowers few, sometimes solitary; flowers ±

zygomorphic; ovules 1-2 or 2-many; seeds 1-

several, two-winged.

Acknowledgements

The assistance of Natalie Camilleri in preparing

the diagrams is much appreciated. The authors

are also indebted to the Curator, Ross

McKinnon AM, for permission to collect living

material from the Brisbane Botanic Garden, Mt

Coot-tha, Dr Paula Rudall of the Royal Botanic

Gardens, Kew for providing literature and an

unnamed referee for helpful comments on the

original manuscript.

References

Bailey, F.M. (1913). Comprehensive Catalogue of

Queensland Plants. Government Printer:

Brisbane.

Brough, P. (1933). The life-history of Grevillea robusta

[Cunn.]. The Proceedings of the Linnean

Society of New South Wales 58: 33-73.

Corner, E.J.H. (1976). The seeds of Dicotyledons.

Cambridge University Press: Cambridge.

Douglas, A.W. (1995). Affinities. In P. McCarthy (ed.).

Flora of Australia 16, Elaeagnaceae, Proteaceae

1: 6-14. CSIRO Australia: Melbourne.

Drummond, J. (1853). On the botany of the north western

District of Western Australia. Hooker’s Journal

of Botany and Kew Garden Miscellany 5: 177—

183.

Filla, F. (1926). Das Perikarp der Proteaceae. Flora

120: 99-142.

Forman, D.B. (1995). Stenocarpus. In P.McCarthy (ed.).

Flora of Australia 16, Elaegnaceae,

Proteaceae 1:363-369. CSIRO Australia:

Melbourne.

Francis, W.D. (1928). The anatomy of the Australian

bush nut ( Macadam ia ternifolia). Proceedings

of the Royal Society of Queensland 39: 43-

52, + plate 1.

Gardner, G.A. (1933). Contributiones florae australiae

occidentalis No. VIII. Journal of the Royal

Society of Western Australia 19: 79-93.

Gardner, G.A. (1942). Contributiones florae australiae

occidentalis No.XI. Journal of the Royal

Society of Western Australia 27: 165-210.

Johnson, L A S. & Briggs, B.G. (1963). Evolution in

the Proteaceae. Australian Journal of Botany

11: 21-61.

Johnson, L.A.S. & Briggs, B.G. (1975). On the

Proteaceae - the evolution and classification

of a southern family. Botanical Journal of the

Linnean Society 70: 83-182.

Johnson, L.A.S. & Briggs, B.G. (1995). Flora of Australia

16, Elaeagnaceae, Proteaceae 1. CSIRO

Australia: Melbourne.

Jordaan, P.G. (1946a). Die Saadknop en Embriologie

van Leucadendron (with Summary in English).

Annale van die Universiteit van Stellenbosch.

Jaargang XXIII, Reeks A, No.7 (1945): 1-38.

-(1946b). Die Saadknop en Embriologie van

Leucospermum conocarpum R.Br. (with

Summary in English). Annale van die

Universitiet van Stellenbosch. Jahrgang XXIII,

Reeks A, No.7 (1945): 39-53.

Kausik, S.B. (1940). Studies in the Proteaceae. IV.

Structure and Development of the Ovule of

Hakea saligna Knight. Annals of Botany (n.s.)

4: 73-80.

Maiden, J.H. (1904). The Forest Flora of New South

Wales. Part VI. No. 22. Government Printer:

Sydney.

Manning, J.C. & Brits, G.J. (1993). Seed coat

development in Leucospermum cordifolium

(Knight) Fourcade (Proteaceae) and seed

covering structures in Proteaceae. Botanical

Journal of the Linnean Society 112: 139-148.

Meisner, C.F. (1855). New Proteaceae of Australia.

Hooker’s Journal of Botany and Kew

Miscellany 7: 65-78.

Mueller, F. (1871). Fragmenta Phytographie

australiae 7: 131-133. Government Printer:

Melbourne.

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Netolitzky, F. (1926). Anatomie der Angiospermen-

Samen. In K.Linsbauer (ed.), Handbuch der

Pflanzenanatomie 10: 364-67. Gebruder

Borntraeger: Berlin.

Schwartzbarth, J. (1905). Anatomische Untersuchungen

von Proteaceen-Fruchten und Samen. Beihefte

zum Botanischen Centralblatt 18: 27-78.

Strohschen, B. (1986a). Contributions to the biology

of useful plants. 4. Anatomical studies of fruit

development of the Macadamia Nut

(Macadamia integrifolia Maiden and Betche).

Angewandte Botanik 60: 239-247.

Strohschen, B. (1986b). Contributions to the biology

of useful plants. 5. Anatomical studies of fruit

development and fruit classification of the

Monkey Nut ( Hicksbeachia pinnatifolia

F.Muell.). Angewandte botanik 60: 249-256.

Strohschen, B. (1986c). Contributions to the biology

of useful plants. 6. Anatomical studies of fruit

development and fruit classification of

Persoonia pinifolia R.Br. Angewandte Botanik

60: 251-265.

The rediscovery of Boronia inflexa subsp. grandiflora (Rutaceae)

Marco F. Duretto 1 , Paul Grimshaw 2 & Kym Sparshott 3

Summary

Duretto, M.F., Grimshaw, P. & Sparshott, K. (2005). The rediscovery of Boronia inflexa subsp.

grandiflora (Rutaceae). Austrobaileya 7(1): 171-173. Details of the first collection of Boronia

inflexa Duretto subsp. grandiflora Duretto with accurate site and habitat information are presented.

The descriptions for the species and subspecies are revised and an amended key to the subspecies

of B. inflexa is provided. The habitat of the B. inflexa subsp. grandiflora is discussed for the first

time and it is recommended that the conservation status of the subspecies be revised to ‘Critically

Endangered’.

Key Words: Rutaceae, Boronia inflexa, Boronia inflexa subsp. grandiflora, Queensland flora

'M.F. Duretto, Tasmanian Herbarium, Tasmanian Museum and Art Gallery, Private Bag 4, Hobart,

Tasmania 7001, Australia. Email: marco.duretto@tmag.tas.gov.au.

2 P. Grimshaw, Biodiversity Assessment and Management Pty Ltd, 38 Middle St, Cleveland,

Queensland 4163, Australia. Email: paul@biodiversity.tv.

3 K. Sparshott, Queensland Parks and Wildlife Service, Park Services, Southern Region, Toowoomba

District Office, PO Box 731, Toowoomba, Queensland 4350, Australia. Email:

kym.sparshott@epa.qld.gov.au.

Introduction

Boronia inflexa Duretto subsp. grandiflora

Duretto was described in a revision of Boronia

section Cyanothamnus for eastern Australia

(Duretto 2003). This taxon was first identified

by Ross (1983) as a large-flowered form of

B. bipinnata from the Granite Belt of south¬

eastern Queensland. It may have been part of

what was referred to by McDonald et al. (1995)

as B. sp. aff. bipinnata in their Flora of

Girraween and Bald Rock National Parks.

Boronia inflexa contains four subspecies, two

of which have large flowers: B. inflexa subsp.

grandiflora (Qld) and B. inflexa subsp.

torringtonensis Duretto (N.S.W.). The other two

subspecies, B. inflexa subsp. inflexa and B.

inflexa subsp. montiazura Duretto, have smaller

flowers and are both found in the ‘granite belt’

of south-eastern Queensland. The former is also

known from an isolated population in Gibraltar

Range (N.S.W.).

In the discussion under B. inflexa subsp.

grandiflora, Duretto (2003) noted that the taxon

was known from two collections with vague

collection details. As these collections were from

the western edge of the known distribution of

B. inflexa subsp. inflexa there was the

Accepted for publication 30 June 2005

possibility that these two taxa were not

taxonomically distinct but represented extremes

in flower size. Given that B. inflexa subsp.

grandiflora is a large flowered and showy taxon

and thus would tend to be noticed, the lack of

collections since 1973 indicated that it was

possibly extinct.

During surveys of the north-west portion

of Girraween National Park in November 2000,

two of the authors (Grimshaw & Sparshott)

collected material of a large flowered taxon

which had affinities to B. bipinnata. This

material was made available to Duretto in late

2003 who detennined that the material could be

assigned to B. inflexa subsp. grandiflora. The

type locality of this taxon is ‘property of W.

McDonagh, Lyra’ which is close to where the

recent collection was made in Girraween

National Park. It would be reasonable to

suppose that these two collections have come

from the same population.

Revised descriptions for B. inflexa and

B. inflexa subsp. grandiflora are given below,

together with the first account of the habit and

habitat of the latter. The revised descriptions

have necessitated minor amendments of the key

to the subspecies of B. inflexa as published by

Duretto (2003).

172

Austrobaileya 7 (1): 171-173 (2005)

Taxonomy

Boronia inflexa Duretto, Muelleria 17:40, figs

2I-K (2003). Type: Queensland. Darling

Downs District: Mt Norman, Girraween

National Park, 28° 52’S 151° 58’E, 27

September 1977, J. Armstrong 1149 &

J.M. Powell ( holo: BRI [AQ383403]; iso:

CANB 8305848, NSW 385918).

Erect, woody shrub to 2 m tall and 3 m wide.

Branchlets not or slightly glandular tuberculate,

hispidulous or pilose; hairs concentrated

between leaf decurrencies, becoming glabrous

with age, hairs to 0.5(-0.75) mm long. Leaves

imparipinnate or rarely bipinnate (subsp.

inflexa ), 3-5(-7)-foliolate, entire leaf in outline

6-25 mm long, 6-35 mm wide, not obviously

glandular, glabrous to minutely pilose; petiole

3-10 mm long; rachis segments 3-9 mm long;

terminal leaflets 1-16 mm long, 0.5-2.5 mm wide,

linear, flat, concolorous, dorsiventral, margin

entire, tip acute; lateral leaflets similar to terminal

leaflets or longer, 1.5-16 mm long. Inflorescence

l-3(-7)-flowered, not obviously glandular,

glabrous or glabrescent, smaller to slightly

longer than leaves; peduncles 0.5-9 mm long,

secondary inflorescence units 0.5-1 mm long;

prophylls 0.5-3 mm long, glabrous or

glabrescent with a few hairs towards apex or

minutely ciliate; metaxyphylls 0.5-1.5 mm long;

anthopodia 1-3 mm long. Sepals debate to

narrowly deltate, 1.5-3 mm long, 0.75-1.5 mm

wide, not obviously glandular, flat, glabrous to

minutely ciliate, tip acute or acuminate due to

involute margins. Petals white to pink, 2.5-7.5

mm long, not obviously glandular, glabrous to

minutely ciliate, persistent. Staminal filaments

pilose, slightly glandular tuberculate towards

apex; anther loculi glabrous, apiculum minute,

glabrous to pilose. Ovary glabrous; style

glabrescent to pilose; stigma entire, minute,

scarcely wider than the style. Cocci 3-3.5 mm

long, 1-2 mm wide, glabrous or with few hairs

along suture. Seed dull, grey, 2.5-3 mm long, 1-

1.5 mm wide, irregularly rugulose, without wax

platelets between tubercula.

Conservation status: Three of the four

subspecies of B. inflexa, namely B. inflexa

subsp. grandiflora , B. inflexa subsp.

montiazura and B. inflexa subsp.

torringtonensis , can be considered Critically

Endangered (IUCN 2001) though the last two

lack satisfactory field data. B. inflexa subsp.

inflexa is found over a wider area but accurate

population sizes are unknown. Populations

seen in GibraltarNational Park (see Duretto 2003)

and Girraween National Park (Grimshaw pers

obs.) contained few plants.

Given that populations appear to be

small and occur over very limited areas, the

species as a whole, could be considered to be

Critically Endangered. Only B. inflexa subsp.

inflexa and B. inflexa subsp. grandiflora are

conserved in National Parks.

Key to the subspecies of Boronia inflexa

1. Leaflets minutely pilose.

Leaflets glabrous or glabrescent

Boronia inflexa subsp. grandiflora Duretto,

Muelleria 17:43, figs 2N-0 (2003). Type:

Queensland. Darling Downs District: On

property of W.McDonagh, Lyra, Qld, 22

October 1962 ,K.N. Shea SI 24 (holo: BRI

[AQ151006], transparency MEL 2068529).

Shrub to c. 1.5 m tall. Branchlets not glandular

. subsp. inflexa

. . . subsp. grandiflora

. . . subsp. montiazura

subsp. torringtonensis

tuberculate, pilose, mainly between leaf

decurrencies, hairs to 0.5(-0.75) mm long.

Leaves imparipinnate, 3(-7)-foliolate, entire leaf

in outline (9-) 15-25 mm long, (8-) 17-35 mm

wide, sparsely and minutely pilose; petiole 4-

10 mm long; rachis segments 3-9 mm long;

leaflets 4-15 mm long, 0.75-1.25 mm wide.

Inflorescence axillary, 1-3-flowered, smaller or

2. Petals 2-3.5 mm long; anther apiculum glabrous.

Petals 6-7.5 mm long; anther apiculum with a few hairs . . .

3. Terminal leaflets 3-7 mm long, 0.5-0.75 mm wide.

Terminal leaflets (4—)10—16 mm long, (0.75—)1—1.25 mm wide

Duretto et al., Rediscovery of Boronia inflexa

c. as long as leaves; peduncles 2-7 mm long;

prophylls 1-3 mm long, ciliate towards apex;

anthopodia 1.5-3 mm long. Sepals deltate, 2-3

mm long, 1.25-1.5 mm wide, glabrescent along

margin and/or towards apex, tip acuminate due

to involute margins. Petals pale pink and tinged

with white, 6-7.5 mm long, minutely ciliate.

Anther apicula with a few simple hairs. Style

pilose. Cocci and seed not seen.

Specimens examined : Queensland. DARLING

DOWNS DISTRICT: Near Amiens, SE Qld, Sep 1973,

Harslett s.n. (NE2638 [transparencies BRI, MEL

2068530]); NW section of Girraween NP near Lyra,

Dec 2000, Grimshaw PG2857 & Sparshott (BRI,

NSW).

Distribution and ecology : The only known

extant population of B. inflexa subsp.

grandiflora is found in Girraween National Park

on the Granite Belt of southeast Queensland.

The two older known collections give imprecise

locality information of near Amiens and Lyra,

also in the Granite Belt. The population in

Girraween National Park was found in

Eucalyptus prava, E. andrewsii and E. caleyi

woodland with a dense, shrubby and fairly

diverse understorey. Scattered throughout the

area are granitic pavements and outcrops.

Flowering material has been collected from

September to December.

Conservation status : Duretto (2003) indicated

that a conservation code of 2K (following the

format of Briggs & Leigh 1996) was appropriate

for B. inflexa subsp. grandiflora as only two

collections with vague locality information were

known. The third collection of this taxon

0 Grimshaw PG2857 & Sparshott ) has precise

locality and habitat information and was

recently made within Girraween National Park.

The taxon was seen in only one valley during a

survey of the area (Grimshaw & Sparshott, pers.

obs.) and approximately a dozen mature plants

were seen over an area of 1-2 hectares. The site

was burnt in October 2002 (Sparshott, pers.

obs.) although it is not known how the

population was affected. One of the authors

(Grimshaw) has travelled extensively in the park

over many years (in part as a park ranger

between 1973 and 1982) and has not observed

it elsewhere. Given this additional information,

conservation codes of 2ECt (following Briggs

& Leigh 1996), or CR (Critically Endangered;

IUCN 2001) are appropriate. The taxon satisfies

173

Critically Endangered criterion D [Population

size estimated to number fewer than 50 mature

individuals] of the IUCN Red List. Potential

threats to the taxon’s survival would include

wildfires (especially if these occurred too

frequently), any development [including roads,

fire trails and walking tracks] in the area, and

weed incursions. Further surveys are urgently

required to determine if there are any other

populations of this taxon, as is monitoring of

the only known population.

Acknowledgements

The authors would like to thank Gintaras

Kantvilas (HO) and an anonymous reviewer

who made useful comments on an earlier draft

of this manuscript.

References

Iucn (2001). IUCN Red List Categories and Criteria:

version 3.1. IUCN Species Survival

Commission. IUCN: Gland, Switzerland/

Cambridge, UK.

Briggs, J.D. & Leigh, J.H. (1996). Rare or Threatened

Australian Plants , revised edition. CSIRO

Australia: Collingwood.

Duretto, M.F. (2003). Notes on Boronia (Rutaceae) in

eastern and northern Australia. Muelleria 17:

19-135.

McDonald, B., Gravatt, C., Grimshaw, P. & Williams, J.

(1995). The Flora of Girraween and Bald

Rock National Parks. Queensland Herbarium,

Queensland Department of Environment and

Heritage: Brisbane.

Ross, E.M. (1983). Rutaceae. In T.D. Stanley & E.M.

Ross (eds). Flora of South-eastern Queensland

1: 440-470. Queensland Department of

Primary Industries miscellaneous publication

81020: Brisbane.

New species of Philotheca Rudge (Rutaceae) from Queensland

Paul I. Forster

Summary

Forster, P.I. (2005). New species of Philotheca Rudge (Rutaceae) from Queensland. Austrobaileya

7(1): 175-181. Taxa previously allocated subspecies rank under Philotheca myoporoides are

elevated to species rank as P. conduplicata (Paul G. Wilson) P.I.Forst., P. epilosa (Paul G. Wilson)

P.I.Forst., P. glasshousiensis (Domin) P.I.Forst., P. obovatifolia (Bayly) P.I.Forst. and P.

queenslandica (C.T.White) P.I.Forst. Diagnostic descriptions and a key are provided to the

Queensland species of Philotheca section Erionema (F.Muell.) Paul G. Wilson.

Key Words: Rutaceae, taxonomy, Philotheca conduplicata, Philotheca epilosa, Philotheca

glasshousiensis, Philotheca myoporoides, Philotheca obovatifolia, Philotheca queenslandica,

Queensland flora

PI. Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens,

Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: paul.forster@epa.qld.gov.au

Introduction

The genus Philotheca Rudge with c. 50 species

is endemic to Australia and was last revised by

Bayly (1998) and Wilson (1998a). Six species of

Philotheca in the two sections P. section

Philotheca and P. section Erionema (F.Muell.)

Paul G. Wilson have been recognised for

Queensland (Wilson 1998a). Philotheca

acrolopha Paul G. Wilson, P. ciliata Hook.,

P. cuticularis Paul G. Wilson, P. difformis

(A.Cunn. ex Endl.) Paul G. Wilson and

P. sporadica (Bayly) Paul G. Wilson belong to

P. section Philotheca and P. myoporoides s.lat.

to P. section Erionema.

Philotheca myoporoides (DC.) Bayly

s.lat. was revised by Bayly (1998) who

recognised one species with nine subspecies

distributed in eastern Australia. Four of these

subspecies are found in south-eastern

Queensland. The rank at which taxa in

P. myoporoides s.lat. have been recognised is

based on the pioneering work of Wilson (1970)

when he first revised the group (then part of

Eriostemon). Wilson (1970) adopted a broad

species concept of complex species with

numerous phenetically defined subspecies in

the genera Eriostemon mdPhebalium. In recent

times, recognition at species rank of some of

these subspecific taxa, or other new taxa that

might also have been named as subspecies

when using the concepts of Wilson (1970,

Accepted for publication 5 August 2005

1998a,b), has occurred in the genera Leionema,

Phebalium and Philotheca (e.g. Rozefelds

2001; Forster 2003; Horton et al. 2004; Walsh

2004; Weston & Turton 2004). Bayly (1998)

stated “for the most part, taxa in the Philotheca

myoporoides complex are both morphologically

and geographically distinct.most taxa overlap

on a number of these features, and each is largely

defined by some unique combination of

attributes”.

As noted elsewhere (Forster 2005), it is

considered that plant species can be defined as

groups of populations (1-many) with

discontinuities in the variation of two or more

independent character states of morphology. It

is assumed that there is genetic continuity or at

least a shared ancestral lineage between the

different populations of a single species, albeit

in most cases in the absence of non-

morphological data. Specific taxa defined in this

manner are in the sense of the ‘phenetic species

concept’ (Stebbins 1950; van Steenis 1957;

Levin 1979,2000; Cronquist 1988; Stuessy 1990).

If only a single, or several minor (e.g.

indumentum colour or density) character state

differences are present and the discontinuity is

geographically based, the rank of subspecies

is often used. Subspecies should differ in only

a few minor characters and intermediate

populations should exist to demonstrate

continuity of character states (e.g. Stebbins

(1950) states “subspecies.... connected .... by a

series of intergrading forms” or Stace (1989)

176

Austrobaileya 7 (1): 175-181 (2005)

states “a population of several biotypes

forming a more or less distinct regional facies

of a species.a geographical race, ecotype,

topodeme or genoecodeme”).

Neither Wilson (1970) nor Bayly (1998)

have demonstrated that intermediate

populations exist for the four subspecies of P.

myoporoides s.lat. that occur in Queensland,

either between the four subspecies, or between

them and other subspecies in the superspecies.

Weston & Harden’s (2002) statement that

“intergrades [are] known between taxa where

they grow together” appears to not be based

on any published data and may derive from

Wilson’s (1970) observation that there were

“distinct morphological groups which occupy

separate geographical areas (except at one point

where two of them may overlap)”. There are

clear differences between the four subspecies

based on habit, stem, leaf and staminal filament

characters. Recognition of these four

Queensland taxa at species rank is expedited

here to enable use of the names in a forthcoming

census of the Queensland flora. Two of the taxa

have been previously recognised at species

level in the genus Eriostemon (Domin 1926;

White 1942). Revision of the remaining

subspecies of P. myoporoides s.lat. in the states

of New South Wales and Victoria is left in

abeyance for workers familiar with the relevant

taxa.

Materials and methods

The results in this paper are based on the

examination of herbarium material at BRI, MEL

and NSW. All taxa have been studied in the

field. Although descriptions of the individual

taxa can be sourced in Wilson (1970) and Bayly

(1998); new, concise descriptions are provided

below to assist in identification of the

Queensland taxa.

Taxonomy

Key to the Queensland species of Philotheca section Erionema (F.Muell.) Paul G Wilson

1. Stems sparsely to densely warty-verrucose.2

Stems more or less smooth, or with only scattered, slightly raised glands.4

2. Leaves strongly concave when dry; staminal filaments eciliate or with only

scattered long hairs towards the apex.

Leaves ± flat or weakly concave when dry; staminal filaments ciliate for

entire length and with dense long hairs towards the apex.

3. Stems densely glandular-verrucose; leaves oblanceolate to oblong, 20-52

mm long x 5-11 mm wide, length/width ratio 3-5.7. P. glasshousiensis

Stems sparsely glandular-verrucose; leaves obovate, 28-60 mm long x

(11-)15-28 mm wide, length/width ratio 2.2-3.3. P. obovatifolia

4. Leaves elliptic, falcate and strongly conduplicate when dry, 30-65 mm long;

petals 7-11 mm long; capsules 7-9 mm long. P. conduplicata

Leaves narrow-elliptic to narrow-obovate, flat when dry, 7-30 mm long;

petals 5-8 mm long; capsules 5-7 mm long. P. queenslandica

P. epilosa

New Combinations

Philotheca conduplicata (Paul G Wilson)

P.I.Forst. comb. & stat. no Eriostemon

myoporoides subsp. conduplicata Paul

G Wilson, Nnytsia 1:41 (1970); Philotheca

myoporoides subsp. conduplicata (Paul

G Wilson) Bayly, Muelleria 11:124 (1998).

Type: New South Wales. Howell, August

1905, J.H. Maiden & J.L. Boorman s.n.

(holo: NSW68742).

Illustrations : Williams (1984: 127), as

Eriostemon myoporoides subsp.

Forster, Philotheca

177

conduplicatus; Bayly(1998:114, fig. ID)

& Weston & Harden (2002: 297), as

Philotheca myoporoides subsp.

conduplicata.

Woody shrub to 2 m high; rootstock not

rhizomatous. Branchlets ± smooth or with only

scattered glands. Leaves laxly clustered near

stem apices; leaf laminae narrow-elliptic,

conduplicate and falcate when dry, 40-75 mm

long, 6-14 mm wide, length/width ratio 4-10.4;

apex obtuse to acute, mucronate; base attenuate.

Inflorescences pedunculate to 4 mm, with 1-4

flowers; pedicels 2-4.5mm long, sparsely to

densely glandular-verrucose. Petals elliptic-

oblong to oblanceolate, 6-11 mm long, 2.5-3.5

mm wide, adaxially papillose, abaxially glabrous

apart from scattered to dense, verrucose glands

in the middle towards the apex, cream,

sometimes with a pink tinge. Staminal filaments

sparsely ciliate for entire length, apex sparsely

pilose with hairs to 0.5 mm long. Cocci erect,

prominently rostrate, 7-9 mm long.

Selected specimens examined'. Queensland. Darling

Downs District: GlenAplin, Jul 1949 , Althofer s.n. (BRI

[AQ151760]); Wyberba, Dec 1962, Blake 22003 (BRI);

near Wallangarra, Nov 1944, Clemens s.n. (BRI

[AQ151759]); near Ballandean, Oct 1944, Clemens

s.n. (BRI [AQ151763]); 3.3 km SE of Glen Aplin, Sep

1974, Gittins 2793 (BRI); Daphne Peak, Braemar, Glen

Aplin, Sep 1963, Gordon 6404 (BRI); ridge S of Bald

Mt, extreme W section of Girraween N.P., 28° 52’S,

151° 54’E, Sep 1994, Grimshaw PG970 & Turpin (BRI,

NSW); 18 km SW of Stanthorpe, Portion 89,

Nundubbermere Road, 28° 43’S, 151° 45 1 E, Oct 1994,

Halford Q2291 (BRI); Red Rock Gorge, Sundown N.P,

Aug 1983, Haselgrove SD57 (BRI); Fletcher, Mt

Pleasant Fauna Sanctuary, Aug 1964, Philp s.n. (BRI

[AQ151758]); Ballandean, Oct 1933, White 9401 (BRI).

New South Wales. Northern slopes of Bald Knob, 10

km Wof Woodenburg, 28° 22’S, 152° 32’E, Sep 1998,

Bean 13795 (BRI, NSW); WNW of Glen Innes via

Wellingrove, Kings Plains N.P, 29° 34’S, 151° 22’E,

Coveny 16604 & Whalen (BRI, NSW); Jonquil Knob,

Torrington State Recreation area, off Duck Creek trail,

29° ll’S, 151° 32’E, Jul 2001, Forster PIF27435 (BRI,

MEL, NSW); Torrington, 29° 19’S, 151° 42’E, Oct

1969, Jones 4115 (BRI).

Distribution and habitat : Philotheca

conduplicata is endemic to the Granite Belt of

south-east Queensland (Girraween N.P.,

Sundown N.P., Ballandean, Glen Aplin) and

adjacent north-east New South Wales (Bald

Knob, Kings Plains N.P, Torrington). Plants

grow in rocky areas on granite substrates at

altitudes between 700 and 900 m.

Notes : Philotheca conduplicata is distinctive

in a combination of characters, viz. the leaves

are laxly clustered at the stem apices and the

laminae are narrow-elliptic, falcate and strongly

conduplicate when dry. It is also the tallest of

this group of Queensland species, with

individuals up to 2 m observed.

Philotheca epilosa (Paul G. Wilson) P.I.Forst.

comb. & stat. nov.; Eriostemon

myoporoides subsp. epilosus Paul G.

Wilson, Nuytsia 1: 41 (1970); Philotheca

myoporoides subsp. epilosa (Paul G.

Wilson) Bayly, Muelleria 11: 120 (1998).

Type: Queensland. Darling Downs

District: Wallangarra, November 1906,

S.L. Boorman s.n. (holo: NSW69255; iso:

NSW68741).

Illustrations : Ross (1983: 466, fig. 71J) &

Williams (1987: 119), as Eriostemon

myoporoides subsp. epilosus ; Bayly

(1998: 114, fig. IB) & Weston & Harden

(2002: 296) as Philotheca myoporoides

subsp. epilosa.

Woody subshrub to 1 m high; rootstock not

rhizomatous. Branchlets densely glandular-

verrucose. Leaves densely clustered near stem

apices; leaf laminae obovate to oblanceolate,

strongly concave when dry, 4-25 mm long, 2.2-

8 mm wide, length/width 1.6^4.2; apex obtuse

to rounded, strongly mucronate; base cuneate

to truncate. Inflorescences pedunculate to 3 mm,

with a solitary flower; pedicels 2-5 mm long,

densely glandular-verrucose. Petals elliptic-

oblong to obovate, 5-8 mm long, 2-3.5 mm wide,

adaxially papillose, abaxially glabrous apart from

scattered, verrucose glands in the middle

towards the apex, cream, often tinged pink.

Staminal filaments eciliate or only with scattered

cilia for entire length, apex epilose or with only

a few scattered hairs to 0.2 mm long. Cocci erect,

prominently rostrate, 5-7 mm long.

Selected specimens examined : Queensland. Darling

Downs District: Wallangarra, Nov 1973, Abell s.n. (BRI

[AQ13990]); Upper reaches of Bald Rock Creek,

Girraween N.P., Sep 1993, Bean 6369 & Forster (BRI);

Portion 90, Wyberba, near Girraween N.P., 28° 50’S,

151° 54’E, Sep 1993, Bean 6383 & Forster (BRI);

Girraween N.P, Nov 1971, Blake 23658 (BRI); Amiens,

Nov 1956, Bowen s.n. (BRI [AQ151773]); Summit of

Mt Norman, Wallangarra, Nov 1944, Clemens 44562

(BRI); Signboard Mt, Girraween N.P., 28° 49’S, 151°

178

Austrobaileya 7 (1): 175-181 (2005)

58’E, Aug 1995, Forster PIF17582 & Figg (BRI);

Wyberba, 1961, Hockings s.n. (BRI [AQ151769]);

Portion 125 Broadwater, N side of Girraween N.P,

Aug 1974, McDonald 264 (BRI); Stanthorpe, s.dat.,

Macpherson s.n. (BRI [AQ151772]); Amiens, at foot

of “Sow & Pigs”, Oct 1963, Pedley 1485 (BRI);

Property of W. McDonagh, Lyra, Oct 1962, Shea S118

(BRI); Stanthorpe, Aug 1930, Westcott 12 (BRI). New

South Wales. Boonoo Boonoo N.P., NE of

Tenterfield, near Boonoo Boonoo River, 28° 51’S, 152°

08’E, Sep 2001, Copeland 3182 (BRI, MEL, NSW);

29 km N of Tenterfield off the Mt Lindesay Highway,

Bungoona track to summit. Bald Rock N.P, 28° 51’S,

152° 02’E, Oct 1993, Coveny 16563 & Whalen (BRI,

NSW); Bald Rock, Bald Rock N.P, 28° 51’S, 152° 02’E,

May 1995, Hunter 3202 (BRI, NE); 4 miles [6.4 km]

NNE of Boonoo Boonoo, 28° 53’S, 152° 08’E, Dec

1971, Thurtell 3896 & Coveny (BRI, NSW); Basket

Swamp, 25 km by road NNE of Tenterfield, Boonoo

S.F., 28° 54’S 152° 09’E, Oct 1992, Wilson 8507 (BRI,

NSW).

Distribution and habitat : Philotheca epilosa

is endemic to the Granite Belt of south-east

Queensland (Girraween N.P., Amiens,

Stanthorpe) and adjacent north-east New South

Wales (Boonoo Boonoo, Bald Rock N.P. Plants

grow in open areas of rock slabs and pavements

on granite substrates at altitudes between 800

and 1100 m.

Notes : Philotheca epilosa is almost sympatric

with P. conduplicata in Girraween National Park

near Wyberba; however, I know of no instances

where both taxa have been collected from the

same locality. Despite this near geographical

proximity of the two taxa, there are no

specimens that indicate introgression or hybrid

individuals. The absence of intermediates was

first noted by Wilson (1970), viz. “no herbarium

specimens showing any suggestion of

integradation between the two”. Philotheca

epilosa is distinctive in the small obovate to

oblanceolate leaf laminae that dry strongly

concave and the staminal filaments that are

eciliate or with only scattered cilia towards the

apex.

Philotheca glasshousiensis (Domin) P.I.Forst.

comb, nov.; Eriostemon glasshousiensis

Domin, Biblioth. Bot. 89:286(1926). Type:

Queensland. Moreton District:

Glasshouse Mountains, slopes of Mt

Coonowrin, September 1909, C.T. White

s.n. (holo: PR?, n.v.; iso: BRI [AQ314418]).

Eriostemon trachyphyllus var. leichhardtii

Benth., FI. Austral. 1: 333 (1863); E.

myoporoides subsp. leichhardtii (Benth.)

Paul G. Wilson, Nuytsia 1: 41 (1970);

Philotheca myoporoides subsp.

leichhardtii (Benth.) Bayly, Muelleria 11:

124 (1998). Type: Brroa [= Mt Beerwah,

Glasshouse Mountains], L. Leichhardt

(holo: K n.v. fide Wilson (1970: 41); iso:

MEL4536).

Illustration : Bayly (1998: 114, fig. lc), as

Philotheca myoporoides subsp.

leichhardtii.

Woody subshrub to 1 m high; rootstock not

rhizomatous. Branchlets densely glandular-

verrucose. Leaves densely clustered near stem

apices; leaf laminae oblanceolate to oblong-

obcuneate, ± flat or somewhat incurved when

dry, 20-52 mm long, 5-11 mm wide, length/width

ratio of 3-5.7; apex rounded to tmncate, abruptly

apiculate; base cuneate. Inflorescences

pedunculate to 5 mm; with l^f flowers; pedicels

4-12 mm long, scattered glandular-verrucose.

Petals elliptic-oblong, 6-8.5 mm long, 2.5-3 mm

wide, adaxially papillose, abaxially glabrous or

with scattered, verrucose glands in the middle

towards the apex, cream. Staminal filaments

woolly ciliate for entire length, apex densely

long-pilose with hairs to 1.5 mm long. Cocci

erect, prominently rostrate, 7-11 mm long.

Selected specimens examined : Queensland. Port

Curtis District: S.F. 316, Kroombit Tops, 24° 21’S,

150° 56’E, Feb 1995, Forster PIF16249 (BRI, K, NSW);

Kroombit Tops Forest Reserve, 24° 21’S, 150° 56’E,

Nov 2003, Forster PIF29772 (BRI, HO, MEL); S.F.

316, Kroombit Tops, 51 km SSW of Calliope, 24°

25’S, 150° 56’E, May 1988, Gibson TOH147 (BRI);

ridge to N of Kroombit Creek, c. 6 km SSW of Forestry

Barracks, Kroombit Tops S.F., 24° 25’S, 150° 57’E,

Dec 1983, McDonald 3865 (BRI, NSW). Burnett

District: Cania Gorge, May 1977, Olsen 3538 & Byrnes

(BRI); Cania Gorge, c. 26 km NW of Monto, Aug

1980, Sharpe 2620 (BRI). Wide Bay District: Mt

Cooroora, S.F. 963, 2 km W of Pomona, 26° 22’S,

152° 50’E, Sep 1996, Bean 10746 (BRI, MEL); Mt

Cooroora, Pomona, 26° 22’S, 152° 50’E, Feb 1995,

Forster PIF16121 (BRI, MEL); loc. cit, Oct 1999,

Forster PIF24972 & Booth (BRI, MEL). Moreton

District: top of Glasshouses, Jul 1879, Bailey s.n. (BRI

[AQ151787]); Mt Beerwah N.P, 26° 54’S, 152° 53’E,

Aug 1990, Bean 2150 (BRI); Mt Tibrogargan N.P,

26° 56’S, 152° 57’E, Apr 1993, Bean 5911 (BRI); 1

km NNE of Mt Tibrogargan, N of Beerburrum, 26°

55’S, 152° 57’E, Feb 1997, Bean 11666 (BRI); near

Forster, Philotheca

179

top of Mt Ngungun, Jun 1951, Blake 18765 (BRI); Mt

Beerwah, Aug 1966, Blake 22761 (BRI); Mt

Coonowrin, 26° 54’S, 152° 55’E, Jun 1968, Smith

13967 (BRI); Coochin Hills, east peak, 26° 52’S, 152°

56’E, Aug 1968, Smith 14035 (BRI); Coochin Hills,

near summit of W peak on N side, 26° 52’S, 152° 56’E,

Aug 1968, Smith 14046 (BRI); Glasshouses, Coonoorin

(Crookneck) Peak, Aug 1968, Webster 14990 & Hildreth

(BRI); (BRI); Top of Mt Ngungun, Glasshouse

Mountains, Mar 1931, White 7648 (BRI).

Distribution and habitat : Philotheca

glasshousiensis is endemic to south-eastern

Queensland where it occurs in disjunct

populations with a northern limit at Kroombit

Tops and a southern limit in the Glasshouse

Mountains. Plants occur in small, scattered

populations on rocky outcrops and clifflines

on substrates derived from metamorphosed

sandstones, rhyolite and trachyte at altitudes

between 100 and 900m.

Notes : This species occurs in close

geographical proximity to P. queenslandica but

is not “broadly sympatric” with that taxon as

stated by Bayly (1998). As noted by Bayly

(1998) [there are] “no specimens that could not

be readily assigned to one taxon or the other”.

Philotheca queenslandica and

P. glasshousiensis do not show any evidence

of introgression; hence, the observed variation

does not fulfil the criteria necessary for them to

be allocated subspecies rank.

Philotheca glasshousiensis is perhaps

more closely related to P. obovatifolia and both

share the character of staminal filaments that

are ciliate for the entire length. However, it

differs from that species by having smaller (20-

52 x 5-11 mm) leaves that are oblanceolate to

oblong-obcuneate with a length/width ratio of

3-5.7, and the staminal filaments have long-

pilose hairs towards the apex. In comparison

P. obovatifolia has broadly obovate leaves that

are noticeably larger (28-60 x (11-) 15-28 mm)

with a length/width ratio of 2.2-3.3, and staminal

filaments with only a few scattered hairs

towards the apex.

Philotheca obovatifolia (Bayly) PI.Forst. stat.

nov.; Philotheca myoporoides subsp.

obovatifolia Bayly, Muelleria 11: 123

(1998). Type: Queensland. Moreton

District: Mt Ernest, 10 November 1992,

PI.Forster PIF12364 & G. Leiper (holo:

BRI; iso: MEL).

Illustrations : Bayly (1998: 114, fig. 4), Logan

River Branch S.G.A.P (Qld Region Inc.)

(2002: 323) & Weston & Harden (2002:

296), as Philotheca myoporoides subsp.

obovatifolia.

Woody subshrub to 1 m high; rootstock not

rhizomatous. Branchlets sparsely glandular-

verrucose. Leaves densely clustered near stem

apices; leaf laminae broadly obovate, ± flat, 28-

60 mm long, (11—) 15-28 mm wide, length/width

ratio of 2.2-3.3; apex obtuse to slightly retuse,

shortly mucronate; base cuneate to truncate.

Inflorescences pedunculate to 9 mm, with 1-5

flowers; pedicels 5-9 mm long, sparsely

glandular-verrucose. Petals elliptic-oblong,

8-9 mm long, 3.5^4 mm wide, adaxially papillose,

abaxially glabrous, apart from scattered

verrucose glands in the middle towards the

apex, cream, tinged with pink. Staminal filaments

ciliate for entire length, apex with scattered,

longer hairs to 1 mm long. Cocci not seen.

Additional specimens examined: Queensland.

Moreton District: Mt Barney, near saddle, 28° 17’S,

152° 42’E, Jun 1972, Bell 538 (BRI); Mt Barney, Oct

1935, Everist 1390 (BRI); loc. cit., Sep 1949, Everist

4137 (BRI); Mt Barney, upper slopes SE ridge of East

Peak, 28° 17’S, 152° 41’E, Sep 1994, Forster PIF15720

(BRI, CANB, K, NSW); Mt Barney, summit area East

Peak, 28° 17’S, 152° 41’E, Sep 1994, Forster PIF15722

(AD, BRI); Mt Barney, Dec 1974, McDonald s.n. (BRI

[AQ151747]); Mt Lindesay, Jul 1934, Michael 2067

(BRI); loc. cit., Jul 1935, Michael 2218 (BRI); Mt

Barney, Aug 1931, White 7829 (BRI). New South

Wales. Werrikimbe N.P, Mt Werrikimbe summit, 31°

13, 152° 13’E, Aug 1998, Copeland 1042 (BRI, NE).

Distribution and habitat : Philotheca

obovatifolia is endemic to the extreme south¬

east of Queensland (Mt Barney, Mt Ernest, Mt

Lindesay) and Mt Werrikimbe in north-east New

South Wales. Plants occur in montane (800-

1300 m) heathland, shrubland or woodland on

rocky substrates derived from granite (Mt

Werrikimbe) or rhyolite (Mt Barney, Mt Ernest).

The species appears to be extinct on Mt

Lindesay as it has not been sighted or collected

there in the last 20 years despite extensive

searches.

Notes: Philotheca obovatifolia is a very

distinctive, large leaved taxon in this species

complex. The single collection from Mt

Werrikimbe has stems that are only weakly

glandular-verrucose.

180

Austrobaileya 7 (1): 175-181 (2005)

Philotheca queenslandica (C.T.White) P.I.Forst.

comb, nov.; Eriostemon queenslandicus

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

53: 207 (1942); Eriostemon myoporoides

subsp. queenslandicus (C.T.White) Paul

G Wilson, Nuytsia 1:41 (1970); Philotheca

myoporoides subsp. queenslandica

(C.T.White) Bayly, Muelleria 11: 125

(1998). Type: Queensland. Moreton

District: Caloundra, 23 August 1933, S.L.

Everist454 (holo: BRI).

Illustrations : Williams (1979: 108), as

Eriostemon myoporoides subsp.

queenslandicus, Bayly (1998: 114, fig. If)

& Haslam (2004: 90), as Philotheca

myoporoides subsp. queenslandicus.

Wiry stemmed subshrub to 1 m high; rootstock

often rhizomatous. Branchlets sparsely

glandular verrucose. Leaves densely clustered

near stem apices; leaf laminae narrow-elliptic to

narrow-obovate, flat to concave, 9-30 mm long,

1.5-7 mm wide, length/width ratio of 2.7-10;

apex obtuse to acute, shortly mucronate; base

attenuate to cuneate. Inflorescences

pedunculate to 3 mm, with a solitary flower;

pedicels 3-6 mm long, sparsely glandular-

verrucose. Petals elliptic-oblong, 4-10 mm long,

1.4-3.8 mm wide, adaxially papillose, abaxially

glabrous, apart from scattered verrucose glands

in the middle towards the apex, cream, tinged

with pink. Staminal filaments densely ciliate for

entire length, apex with dense, longer hairs to

1.5 mm long. Cocci erect, bluntly apiculate to

shortly rostrate, 5-8 mm long.

Selected specimens examined : Queensland. Wide Bay

District: Sandy Creek, c. 10 miles W of Tin Can Bay,

Sep 1967, Brass 33696 (BRI); near Boonooroo, Sep

1948, Clemens s.n. (BRI [AQ151812]); Noosa Shire

land, 2.5 km W of Lake Weyba, 26° 25’S, 153° 02’E,

Jan 2003, Forster PIF29234 & Thomas (BRI); Old

Hollett road, 2 km W of Lake Weyba, 26° 25’S, 153°

02’E, Mar 2003, Forster PIF29278 & Thomas (BRI,

MEL, NSW); Peregian, opposite to Lake Weyba N.P.,

26° 29’S, 153° 06’E, Sep 1992, Franks AJG9209018

& Suosaari (BRI); Cooloola Way, Cooloola N.P, 26°

04’S, 152° 59’E, Aug 1994, Grimshaw G896 & Turpin

(BRI); Cooloola, just inside northern boundary of

R1093, east of Noosa River, Jul 1971, Harrold 84

(BRI); Noosa Heads, May 1953, Hunt s.n. (BRI

[AQ151800]); Lake Cootharaba, s.dat.. Keys & Wedd

s.n. (BRI [AQ151803]); near Poverty Point, Tin Can

Bay, 25° 57’S, 153° 03’E, Jun 1978, Williams 78061

(BRI). Moreton District: Maroochie, Jul 1879, Bailey

s.n. (BRI [AQ151796]); Caloundra, Aug 1932, Blake

4133 (BRI); 0.8 km S of Mt Coolum turnoff (Suncoast

Beach Drive) on David Low Way, 26° 34’S, 153° 05’E,

Jun 1992, Doust 357 & Brown (BRI, NSW); Roys Road,

Beerwah, 26° 51’S, 153° 00’E, Oct 1999, Forster

PIF24936 (AD, BRI, MEL, NSW); 1.6 km NE of

Landsborough, 26° 47’S, 152° 58’E, Aug 1988, Ross

3181 (BRI, MEL); 3 km N of Coolum Beach, Jan

1977, Sharpe 2140 (BRI); Bribie Island - Caboolture

road, about halfway. Mar 1964, van Royen 9339 (BRI);

North of Currumundi Lakes, Caloundra, 26° 46’S, 153°

08’E, Mar 1971, Williams s.n. (BRI [AQ151794])..

Distribution and habitat : Philotheca

queenslandica is endemic to south-eastern

Queensland with a northern limit at Boonooroo

and a southern limit at Bribie Island. Plants

occur in lowland, wallum heathlands in boggy

or sandy soils that are seasonally moist to

inundated at altitudes below 100 m.

Notes: Philotheca queenslandica is the most

herbaceous of all the species enumerated here

(the others are all noticeably woody subshrubs

or shrubs) and has been observed to possess a

rhizomatous rootstock (not apparent in the other

taxa). It perhaps most closely resembles

P. conduplicata; however, it differs from that

species in the smaller leaves (9-30 mm long,

versus 30-65 mm long), the leaf shape (narrow-

elliptic to narrow- obovate, flat to concave when

dry, versus elliptic, falcate and strongly

conduplicate when dry) and the smaller flowers

(petals 4-10 mm long versus 7-11 mm long) and

capsules (5-8 mm long versus 7-9 mm long).

Acknowledgements

Thanks to G Guymer (BRI) for comments on the

manuscript and the Directors of MEL and NSW

for access to collections.

References

Bayly, M.J. (1998). Notes on the Eriostemon

myoporoides (Rutaceae) species complex,

including new names and a new generic

placement in Philotheca. Muelleria 11: 113—

126.

Cronquist, A. (1988). The Evolution and Classification

of Flowering Plants , 2 nd edition. New York

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Domin, K. (1926). Eriostemon. In ‘Beitrage zur Flora

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Forster, PI. (2003). Phebalium distans P.I.Forst.

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Forster, Philotheca

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Ross (eds.). Flora of South-eastern

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Philotheca (Rutaceae). Muelleria 15: 19-26.

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Biosystematics. 2 nd edition. Edward Arnold:

London, Melbourne, Auckland.

Stebbins, G.L. (1950). Variation and Evolution in

Plants. Columbia University Press: New York.

Stuessy, T.F. (1990). Plant Taxonomy. Columbia

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delimitation. Flora Malesiana , ser. I, 5:

C LX VII-C CXXIX.

Walsh, N.G. (2004). A new species of Leionema

(Rutaceae) from south-eastern New South

Wales. Telopea 10: 805-810.

Weston, P.H. & Harden, G.J. (2002). Philotheca. In

GJ. Harden (ed.). Flora of New South Wales 2 nd

edition, 2: 292-299. University of New South

Wales Press: Kensington.

Weston, P.H. & Turton, M. (2004). Phebalium bifidum

(Rutaceae), a new species from the Capertee

Valley, New South Wales. Telopea 10: 787-

792.

White, C.T. (1942). Contributions to the Queensland

Flora, No. 7. Proceedings of the Royal Society

of Queensland 53: 201-228.

Williams, K.A.W. (1979). Native Plants of Queensland.

Vol. 1. K.A.W. Williams: North Ipswich.

Williams, K.A.W. (1984). Native Plants of Queensland.

Vol. 2. K.A.W. Williams: North Ipswich.

Williams, K.A.W. (1987). Native Plants of Queensland.

Vol. 3. K.A.W. Williams: North Ipswich.

Wilson, P.G. (1970). A taxonomic revision of the genera

Crowea, Eriostemon and Phebalium

(Rutaceae). Nuytsia 1: 1-155.

Wilson, P.G. (1998a). A taxonomic review of the genera

Eriostemon and Philotheca. Nuytsia 12: 239-

265.

Wilson, P.G. (1998b). New species and nomenclatural

changes in Phebalium and related genera

(Rutaceae). Nuytsia 12: 267-288.

Papuasian Orchid Studies, 2

Paul Ormerod

Summary

P. Ormerod (2005). Papuasian Orchid Studies, 2. Austrobaileya 7(1): 183-203. The new species

Dendrobium eymanum, D. flebiliflorum, D. paragnomus, D. spenceanum, D. spinuliferum, D.

stipiticola, Glomera pseudomonanthos, Pseuderia takeuchii, Robiquetia brassii and Tainia

serratiloba are described from Papuasia (Indonesian Papua, Papua New Guinea, Solomon Islands).

All species are illustrated and notes are provided on their affinities, distribution and habitats.

Key Words: Orchidaceae, Dendrobium eymanum, Dendrobium flebiliflorum, Dendrobium

paragnomus, Dendrobium spenceanum, Dendrobium spinuliferum, Dendrobium stipiticola,

Glomera pseudomonanthos, Pseuderia takeuchii, Robiquetia brassii, Tainia serratiloba, Papua

New Guinea flora, Indonesian Papua flora, new species

P. Ormerod, PO. Box 8210, Cairns, Queensland 4870, Australia.

Introduction

This paper is the second in a series devoted to

the study of Papuasian Orchids and deals with

fourteen species belonging to five genera. The

first part (Ormerod 2003) dealt with five species

and two subspecies belonging to the genera

Appendicula Blume, Dendrobium Sw. and

Eulophia R.Br. ex Lindl. The orchid flora of

Papuasia (Indonesian Papua, Papua New

Guinea, Solomon Islands) comprises

approximately 2800 species distributed in about

137 genera. Despite this richness there is

comparatively little basic taxonomic work in

progress. This paper is thus an attempt to

redress the latter situation and draws attention

to a number of infrequently collected and

previously undescribed taxa.

Materials and methods

This study is mostly based on examination of

rehydrated herbarium materials from Papuasia.

Living and spirit-preserved material was

examined only for the description of

Dendrobium paragnomus. Herbarium

collections were studied from AMES, BRI, K,

LAE and NSW. All specimens have been seen

unless otherwise indicated.

Taxonomy

Dendrobium Sw.

In the broad sense of this genus the island of

New Guinea has about 420 species (Schuiteman

Accepted for publication 8 August 2005

& de Vogel 2001). The notes below mostly

concern taxa in the sections Latouria and

Pedilonum ; however, a new species is also

named from the section Herpetophytum.

Dendrobium sect. Herpetophytum Schltr.

This section contains sixteen accepted species

all endemic to New Guinea. The known taxa are

D. appendicula Schltr., D. decumbens Schltr.,

D. disoides Schltr., D. fusciflorum Ormd.,

D.glossorhynchoides Schltr., D herpetho-

phytum Schltr., D. hippocrepiferum Schltr.,

D.millarae A.D.Hawkes, D. minimiflorum Gilli,

D. nigricans Schltr., D. oxychilum Schltr., D.

podocarpifolium Schltr., D. scopula Schltr.

(syn.: D. lucidum Schltr.), D. vagabundum

A.D.Hawkes & Heller, D. vestigiiferum J.J.Sm.

and D. vonroemeri J.J.Sm. The plants are eas¬

ily mistaken in herbaria for specimens of

Podochilus Blume due to a superficially simi¬

lar habit of wiry, branching stems with small

leaves. Apart from the distinctive habit, plants

of section Herpetophytum are characterized by

having single-flowered inflorescences,

nonresupinate flowers lasting several days and

connate lateral sepals which form a frontally

closed oblongoid mentum.

Recently I added Poaephyllum fuscum

Ridl. to this section as Dendrobium fusciflorum

with the comment that it has the longest leaves

(to 43 mm) in the section (Ormerod 2002). This

remark is incorrect; both D. podocarpifolium

Schltr. and D. vagabundum A.D.Hawkes &

Heller have longer leaves (to 60mm long).

184

Austrobaileya 7 (1): 183-203 (2005)

Dendrobium stipiticola Ormerod sp. nov.

Species nova in sectione Herpetophyto

singularis, foliis carnosis brevisque,

labellis subbilobatis et epichilio

obsolescenti statim dignoscenda. Typus:

Papua New Guinea. Western Highlands

Province: Mt. Hagen Subdistrict, Tomba

to Mt. Hagen track, 26 May 1972, PF.

Stevens & J.F. Veldkamp LAE 54919

(holo: AMES).

Epiphytic herb. Roots terete, 0.5-0.8 cm thick.

Primary stem at least 30 cm long and 0.09-0.11

cm thick; with stemlets to 7.5 cm long and 0.1-

0.15 cm thick arising every 1.5-4 cm, these root

at the base and become bare in the lower half as

the leaves dehisce. Leaves fleshy, subfalcate-

oblong and obtuse in lateral view, mid dull green,

to 6 mm long, one side c. 1.75 mm wide; leaf

sheaths smooth. Inflorescences single, axillary,

one-flowered, c. 1 mm long. Flowers

nonresupinate; pedicel plus ovary clavate, c. 4

mm long; sepals yellow in bud but dirty white

when open, lip edged purple. Dorsal sepal

oblong-elliptic, obtuse, margins involute in

upper half, c. 6.5 mm long and 2.5 mm wide;

lateral sepals broadly ovate-elliptic, with an

oblong-elliptic basal dilation c. 2.5 mm long, that

forms with the columnfoot a c. 3 mm long

oblongoid and retrorse mentum, obtuse,

margins involute in apical third, c. 5 mm long at

midline and c. 3 mm wide halfway. Petals oblong-

elliptic, subacute, c. 6 mm long and 2 mm wide;

labellum with claw c. 3 mm long and 2 mm wide

thence expanding into a transversely elliptic-

renifonn blade c. 4 mm long and 6mm wide with

broadly elliptic-subquadrate broadly rounded

sidelobes and a minute recessed emarginate

midlobe. Column semiterete, c. 2.5 mm long with

columnfoot about 3 mm long. Fig. 1.

Distribution and habitat : Papua New Guinea,

Western Highlands Province. Lower montane

forest, c. 2980 m. The type was collected from a

fallen log.

Notes : This species is unique in the section

due to its combination of short fleshy leaves

and its flowers with a sub-bilobed lip with a

tiny recessed midlobe. The collectors of the type

also believed they saw the same species lower

down at 2560 m.

Etymology : The specific epithet is derived from

the classical Latin stipitis (stalk or stem; in

classical times applied to a log, stump or trunk

of a tree) and the compound cola (a dweller) in

reference to where the type specimen was found

growing.

Dendrobium sect. Latouria (Blume) Miq.

There are about 53 currently accepted species

in this section. Another four are added here

along with some notes on synonymy and a

previously overlooked combination. The

previous revision of sect. Latouria by Cribb

(1983) dealt with 48 taxa and is still a quite useful

account for orchid hobbyists since all the later

described entities are rarely encountered in

horticulture and then usually in specialist

collections.

Dendrobium eymanum Ormerod sp. nov.

Species nova affinis D. acutisepalo J.J.Sm. sed

carinis labelli glabris et apicis oblique

elevates differt. Typus: Indonesia. Papua

Province: Wissel Lakes area, Enarotali to

Koegapa, Egogitoagapa to Enarotali, 29

March 1939, P.J. Eyma 4814 (holo: AMES

[right hand specimen]; iso: K, L n.v .).

Dendrobium acutisepalum auct. non J.J.Sm.:

van Royen( 1979:314,/?./?.); Cribb(1983:

305 ,p.p).

Roots fleshy, terete, 0.2-0.3 cm thick. Stem 5-6

leaved in upper half, terete, to 24 cm long and

0.4-0.5 cm thick; internodes 0.85-3.5 cm long.

Leaves stiffly coriaceous, erect, elliptic, obtuse

to weakly bilobed, 3-5 cm long and 1.7-2.4 cm

wide. Inflorescences subterminal, laxly 7-10

flowered, to 22 cm long; peduncles 12-14.5 cm

long; sheathing bracts 2-3, scattered, tubular,

clasping, 1.2-1.4 cm long; rachis to 7.5 cm long;

floral bracts lanceolate-cymbiform, acute, 0.6-

0.9 cm long and 0.2-0.3 cm wide. Flowers

resupinate, colour unknown; pedicel plus ovary

terete, becoming weakly obconical and with low

(irregular or wavy) ribs 20-28 mm long. Dorsal

sepal oblong, acute, c. 13 mm long and 4.1-5

mm wide; lateral sepals ovate-elliptic, acute,

medially carinate, c. 13 mm long and 10.5 mm

wide basally, at middle c. 5 mm wide, forming

with the columnfoot a conical 7.5-8 mm long

Ormerod, Papuasian Orchid Studies

185

5mm

Fig. 1. Dendrobium stipiticola. A. apex of stemlet with old inflorescence remnant. B. longitudinal section of

pedicel plus ovary and column with lateral sepal. C. adaxial view of dorsal sepal. D. adaxial view of petal. E. adaxial

view of lateral sepal. F. adaxial view of labellum. All from Stevens & Veldkamp LAE 54919 (AMES). Scales as

indicated.

mentum. Petals obovate-elliptic, c. 13.5 mm long

and 5 mm wide; labellum trilobed, c. 17 mm long;

hypochile c. 16 mm long to tip of sidelobes

(including the c. 5 mm long ‘claw’) and 12 mm

wide; sidelobes obliquely obcuneate, obtuse;

callus of three thickly laminate carinae most

prominent between the sidelobes, the midkeel

only evident apically, apices rounded and

raising forward, c. 2 mm high at termination at

base of epichile; epichile concave, transversely

elliptic-trapeziform, subacute, c. 5.5 mm long and

4 mm wide. Column short, stout, c. 2 mm long;

columnfoot 7.5-8 mm long. Fig. 2.

Distribution and habitat : Indonesia, Papua

Province. Heath vegetation at c. 3000 m.

Notes : Florally this species strongly resembles

Dendrobium acutisepalum J.J.Sm. from which

it may be distinguished by the slightly longer

mentum (7.5-8 mm vs. 6 mm), an oblong (versus

elliptic) dorsal sepal, a labellum callus with

glabrous, thickly laminate carinae that raise up

and forward at termination and the transversely

elliptic-trapeziform (or subtrilobate) midlobe. In

D. acutisepalum the lip has a callus with thicker

keels that are verrucose above in the upper half

which do not raise forward obliquely at

termination and the midlobe varies from rhombic

to oblong-lanceolate but is never subtrilobate.

On the type sheet of D. eymanum I have

chosen the specimen on the right hand side of

the sheet to be the holotype. The material on

this sheet is not a mixture it is just that the

specimen on the left hand side has slightly

younger flowers.

Dendrobium eymanum probably occurs

at higher altitudes (3000 m+) since it was

collected in an area of heath vegetation which

is typically an element of the alpine flora.

Etymology : The specific epithet honours PJ.

Eyma, collector of the type specimen.

186

Austrobaileya 7 (1): 183-203 (2005)

Fig. 2. Dendrobium eymanum. A. lateral view of flower. B. adaxial view of dorsal sepal. C. adaxial view of petal.

D. lateral view of labellum. E. labellum minus midlobe with sidelobes spread. F. column. All from Eyma 4814

(AMES). A-E and F to respective scales.

Ormerod, Papuasian Orchid Studies

Dendrobium flebiliflorum Ormerod sp. nov.

Species nova inter species generis Dendrobii

mento floris semilunatis, labello trilobatis

et epichilio transverse ellipticis distincta.

Typus: Papua New Guinea. Southern

Highlands Province: Ialibu area?,

received 22 January 2003, cult. G.C.

Stockers.n. (holo: BRI [AQ751261]).

Stems narrowly clavate, shallowly sulcate, erect,

bifoliate, c. 18 cm long and 0.25-0.3 cm wide

basally, c. 0.4 cm wide at middle. Leaves broadly

oblong, acute, 3-6 cm long and 1.5-1.9 cm wide.

Inflorescences subterminal, erect, 2-3 flowered,

up to 4.5 cm long; peduncles up to 3.2 cm long;

rachis weakly fractiflex (zig zag), up to 1.3 cm

long; floral bracts ovate, acute, c. 0.4 cm long

and 0.2 cm wide. Flowers resupinate; pedicel

plus ovary clavate, 1.6-1.7 cm long; sepals

greenish tinted brownish, whitish at base.

Dorsal sepal broadly ovate, acute, c. 7 mm long

and 5 mm wide; lateral sepals broadly ovate,

acute, forming with the columnfoot an 8-9 mm

long incurved-semilunate obtuse mentum, c. 8

mm long and 9.5 mm wide. Petals broadly

oblong-subcuneate, acute, c. 8 mm long and 3

mm wide; labellum trilobed, c.10 mm long;

hypochile with obliquely oblong sidelobes, half

of apex minutely denticulate, other half

rounded-entire, sidelobes c. 6 mm long and 4

mm wide basally then slowly narrowing to c. 3

mm wide; callus of three laminate keels highest

at their termination at the base of the midlobe;

epichile transversely elliptic-reniform, margins

minutely erose-denticulate, c. 3 mm long and 6

mm wide. Column short, c. 3.5 mm long and c. 3

mm wide at base; anther cap c. 1 mm long;

columnfoot c. 8 mm long. Fig. 3.

Distribution : Papua New Guinea. The precise

distribution of this species is unknown.

Notes : This plant is a singular, small-flowered

species of which I know no close relatives. It is

distinctive in having flowers with a blunt,

semilunate mentum combined with a trilobed

lip with a low, trilaminate callus and transversely

elliptic midlobe.

Etymology : The specific epithet is derived from

the Latin flebilis (mournful or sad) and florus

(flowered) in reference to the dull colour of the

flowers.

187

Dendrobium montis-yulei Kraenzl., in A.Engler,

Pflanzenr. IV 50. II. B. 21,45:150 (1910).

Type: Papua New Guinea. Central Province:

nearMt. Yule, W. McGregor s.n. (holo: Bt;

iso: HBG (photograph and sketch seen)).

Dendrobium terrestre J.J.Sm., Bull. Jard. Bot.

Buitenz. 2: 10 (1911), syn. nov. Type:

Indonesia. Papua Province: Mt. Goliath,

March 1911, A.C. de Kock 119 (holo: BO

n.v.).

Additional specimen examined : Indonesia. Papua

Province: Wissel Lakes, Enarotali, May 1960, Vink &

Schram 8599 (AMES).

Distribution : Indonesia (Papua Province);

Papua New Guinea (Central Province).

Notes : Cribb (1983) correctly suspected that

D. montis-yulei was the earlier name for D.

terrestre but due to the unavailability of type

material he did not combine the two. Since then

the type specimen (or part of it) has been located

in Hamburg. Dr. Dariusz Szlachetko kindly sent

an analytical drawing and photograph of the

latter, interpretation of which confirms the need

for the above reduction.

Cribb (1983) also noted two albinistic

variants from Papua New Guinea. A similar form

also seems to occur in Indonesian Papua

Province, since according to the collectors of

the above number from Wissel Lakes, it has

white flowers, a green lip and a column coloured

green with red.

Dendrobium spenceanum Ormerod sp. nov.

Species nova affinis D. rarifloro J.J.Sm. sed

petalis obovatis et epichilio labelli

profunde emarginatis differt. Typus:

Indonesia. Papua Province: 4 km SW of

Bernhard Camp, Idenburg River, March

1939, L.J. Brass 13324 (holo: AMES).

Epiphytic herb. Rhizome abbreviated. Roots

terete, relatively fleshy, to 1.8 mm thick. Stems

unifoliate, when ovoid (then c. 13 mm long and

6 mm thick), fusiform when older, c. 25 mm long

and 5.5 mm thick, lower part c. 2 mm thick. Leaves

stiffly coriaceous, oblong-elliptic, acute, c. 35

mm long and 13 mm wide. Inflorescences

pseudoterminal, one flowered, c. 27 mm long;

Austrobaileya 7 (1): 183-203 (2005)

5mm

Fig. 3. Dendrobium flebiliflorum. A. lateral view of flower. B. adaxial view of dorsal sepal. C. adaxial view of petal.

D. adaxial view of lateral sepal; E. lateral view of labellum. F. adaxial view of labellum. All from Stocker s.n. (BRI

[AQ751261]). A and B-F to respective scales.

sheathing bracts c. 2.5 mm long; floral bracts c.

2 mm long. Flowers c. 11 mm long; pedicel plus

ovary weakly ribbed, narrowly clavate,

glabrous; sepals and petals dirty white, lip

brown. Dorsal sepal elliptic, obtuse, c. 13 mm

long and 6.5 mm wide; lateral sepals broadly

elliptic, obtuse, c. 12 mm long and 11 mm wide,

forming with the columnfoot a c. 5 mm long

bluntly conical mentum. Petals obovate,

minutely apiculate c. 12 mm long and 7 mm wide;

labellum trilobed, flabellate-obdeltate, c. 16 mm

long and 17 mm wide; callus c. 9 mm long, thickly

tricarinate in apical third, its lateral carinae

weakly evident basally, the midkeel most

Ormerod, Papuasian Orchid Studies

189

evident apically; sidelobes broadly obovate-

subquadrate, rounded, c. 6.5 mm wide (measured

obliquely in free part); epichile transversely

oblong-bilobulate (each side obliquely

subquadrate), deeply emarginate, c. 3 mm long

and 7 mm wide. Column semiterete-conical, c.

2.5 mm long; columnfoot c. 5 mm long. Fig. 4.

Distribution and habitat : Indonesia, Papua

Province. Lower montane mossy forest at c. 900

Notes : The plant habit and general lip shape

(including the callus) of this species is

reminiscent of D. rariflorum J.J.Sm. It differs

from that species in having obovate petals and

a lip with a transversely oblong-bilobulate

midlobe. In D. rariflorum the petals are rhombic

and the lip has an elliptic-subrhombic midlobe

longer than broad.

Etymology : The species is named after Mr. Phil

Spence who has long maintained a specialist

interest in the taxa of section Latouria with

regards to both cultivation and taxonomy.

Dendrobium spiimlifemm Ormerod sp. nov.

Ab omnibus aliis speciebus sectionis Latouriae

inflorescentiis sub-unifloris, floribus

pallide viridibus, labello flabellatis humili-

carinatis et nervis pro medio molliter

spinuliferis longis distinguitur. Typus:

Indonesia. Papua Province: 15 km SW of

Bernhard Camp, Idenburg River, January

1939, L.J. Brass 12051 (holo: AMES).

Epiphytic herb. Rhizome abbreviated. Roots

several, terete, c. 1 mm thick. Stems bifoliate,

narrowly clavate, to 65 mm long and 3 mm thick.

Leaves coriaceous, lanceolate, acute, c. 37 mm

long and 7 mm wide. Inflorescences subterminal,

one (two?) flowered, c. 5 mm long; floral bracts

broadly ovate, cupular, acute, c. 2 mm long.

Flowers resupinate; pedicel plus ovary terete,

obconically dilated apically, c. 20 mm long,

glabrous; sepals and petals pale green, lip

pencilled with purple. Dorsal sepal ovate-

elliptic, acute, c. 14 mm long and 7 mm wide;

lateral sepals ovate-deltate, acute, c. 15 mm long

and 12 mm wide, forming with the columnfoot a

c. 8 mm long bluntly conical mentum. Petals

oblong-lanceolate, acute, c. 13.5 mm long and 4

mm wide; labellum trilobed, broadly clawed

thence widely flabellate-obdeltate; hypochile c.

17 mm long and 20 mm wide with the c. 8 mm

wide sidelobes irregularly erose-truncate

apically; keels three, rather low, separated, each

broadly grooved above, the lateral ones each

tenninating in a line of softly fleshy spinuliform

projections and also flanked alongside the outer

edges with a line of the same projections which

also occur occasionally on the veins of the

sidelobes; epichile stiffly flexible, transversely

elliptic-reniform when spread, margin irregular,

apex decurved, acute, c. 5 mm long and 10 mm

wide. Column short, subconical, c. 3 mm long;

columnfoot 7.5-8 mm long. Fig. 5.

Distribution and habitat : Indonesia, Papua

Province. Lower montane mossy forest at c.

1800 m.

Notes : This species seems to stand alone in

section Latouria due to its combination of rare

characters such as the very short inflorescence

of one relatively large flower, the large sidelobes

irregularly toothed at the front and the low

separated keels terminating in and flanked by

soft spinuliform projections.

The type specimen has only one leaf but

an abscission scar on that leaf-bearing stem

indicates that two leaves were originally present.

Etymology : The specific epithet is derived from

the Latin spinula (a small thorn or prickle) and

the compound -fer (carrying or bearing), in

reference to the labellum with spinuliform

processes.

Dendrobium stenopterum (Rchb.f.) Chadim,

Orchadian4:28 (1972).

Dendrobium macrophyllum A.Rich. var.

stenopterum Rchb.f., Gard. Chron. 3 rd

series3: 393 (1888).

Type: Sine loc. [? New Guinea], May 1888,

cult. W. Bull s.n. (holo: W n.v., icon

W49207, microfiche seen).

Distribution: Not known.

Notes : The above combination has been

overlooked probably because Chadim (1972)

made it among the bibliography at the end of

his article. He definitely accepted the name

D. stenopterum for a species different from

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Fig. 4. Dendrobium spenceanum. A. plant. B. lateral view of flower. C. lateral profile of callus. D. adaxial view of

labellum. E. adaxial view of petal. F. adaxial view of dorsal sepal. All from Brass 13324 (AMES). A and B-F to

respective scales.

Ormerod, Papuasian Orchid Studies

191

Fig. 5. Dendrobium spinuliferum. A. plant. B. adaxial view of labellum. C. adaxial view of dorsal sepal. D. adaxial

view of petal. E. column. All from Brass 12051 (AMES). A and B-E to respective scales.

192

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D. macrophyllum A.Rich, even though the

transfer is found only in the bibliography.

Dendrobium stenopterum is a member of

the D. polysema Schltr. complex which occurs

in the Philippines, Sulawesi to New Guinea and

down to Vanuatu. It has been very difficult for

me to distinguish species in this group as all

the forms seem to run into each other. However

Phil Spence (pers. comm.) has found

discriminatory characters for taxa in this complex

and he intends to elaborate on these in a

proposed book on section Latouria.

Dendrobium sect. Pedilonum Blume

Dauncey (2003) published a revision of this

section, accepting 47 species. To this section I

would add D. limpidum Schuit. & de Vogel,

D. scabrifolium Ridl. (? = D. crenatifolium

J.J.Sm.), D. usitae Yukawa (a natural hybrid

between D. bullenianum Rchb.f. and

D. goldschmidtianum Kraenzl.) and

D. paragnomus which is described below. Also

I restate my position on D. obtusum Schltr. and

extend the distribution of one species to

Indonesian Papua.

Dendrobium nothofagicola T.M.Reeve,

Orchadian 7: 134 (1982). Type: Papua

New Guinea. Enga Province: Wabag

Distr., Yulimandaka via Sopas, (fl. in cult.

June 1981), P. Ken sub T. Reeve 551 (holo:

LAE, n. v.; iso: E, K, NSW n.v.).

Additional specimen examined : Indonesia. Papua

Province: 18 km NE of Lake Habbema, Bele River,

Nov 1938, Brass 11099 (AMES).

Distribution : Indonesia (Papua Province);

Papua New Guinea.

Notes : This species has not been previously

recorded for the Indonesian province of Papua.

The flowers of this collection were recorded as

being orange-yellow and are typical for the

species.

Dendrobium obtusum Schltr., inK.Schum. &

Lauterb., Fl. Schutzgeb. Sudsee, Nachtr.

2:177(1905). Type: Papua New Guinea.

West Sepik Province: Torricelli Ranges,

April 1902, R. Schlechter 14451 (holo:

Distribution : Indonesia (Papua Province);

Papua New Guinea (including Bougainville).

Notes : Dauncey (2003) has accepted D.

concavissimum J.J.Sm. as the correct name for

this species whilst accurately noting that the

name D. lauterbachianum A.D.Hawkes is a

superfluous and illegitimate replacement for D.

obtusum Schltr.

I have previously outlined my arguments

why D. obtusum was a valid name and not a

homonym of “D. obtusa” Rchb.f. 1861 (Ormerod

1995). The latter name occurs in the index of

Walper’s Annales (Reichenbach 1861: 1150)

under Dendrobium. It is quite obvious that the

epithet and reference “D. obtusa Bl. (s.) 500.

no. 24” refers to Dendrocolla obtusa Blume

which is listed by Reichenbach as a synonym

of Sarcochilusobtusus (Blume) Rchb.f.

It cannot be argued that Dendrobium

obtusa of 1861 is a valid name or transfer

because it was NOT accepted by the supposed

author who correctly indicates by placing an

“(s.)” with it in the index that it is a synonym.

Thus the name “D. obtusa” is invalidly

published because it is not accepted (Greuter

et al. 2000, ) and it thus has no status under the

ICBN {Art. 12). Therefore the later D. obtusum

of Schlechter must be accepted as a valid name

and not a homonym.

Dendrobium paragnomus Ormerod sp. nov.

Species nova affinis Dendrobio gnomo Ames

sed foliis latiusculis, inflorescentiis

sessilis, floribus glabris (non scabris) et

marginibus labelli integris (non minute

erosis) differens. Typus: Papua New

Guinea? cult. G.C. Stocker sub P. Ormerod

23 (holo: BRI).

Dendrobium sp. aff. D. gnomus: Lavarack et

al. (2000:241).

Roots filamentous, white. Rhizome short. Stems

clustered, 6-10 leaved in apical half, to 35 cm

long and 0.6 cm thick; intemodes cylindrical,

0.5-1.5 cm long. Leaves oblong-lanceolate,

subacute, 2.7-3.5 cm long, 1.2-0.9 cm wide;

sheaths purple-red with slightly raised nerves,

ashy white when old, ± truncate. Inflorescences

sessile, c. 3 mm long, four-flowered, on leafless

Ormerod, Papuasian Orchid Studies

193

stems or below leaves on leafy stems; floral

bracts deltate-cymbiform, subacute, 5-6 mm

long and 4 mm wide. Flowers resupinate; pedicel

plus ovary narrowly clavate, ribbed, 7-8 mm

long, glabrous, unscented, long lasting; sepals,

petals and lip dark vivid mauve. Dorsal sepal

ovate-elliptic, subacute, c. 8 mm long and 3.7

mm wide; lateral sepals obliquely ovate-elliptic,

subacute, c. 13 mm long and 5 mm wide, forming

with the column-foot a mentum c. 5 mm long.

Petals oblong, acute, 7.5-8 mm long and 2.5 mm

wide; labellum obovate-trulliform, subacute,

apical 1/3 obtusely deflexed, c. 12 mm long and

5 mm wide, the basal 1/4 concave with fluid

inside, c. 3 mm long and 2.5 mm wide, separated

from the lamina by a transverse septum,

constriction at septum c. 2 mm wide, lamina c. 9

mm long and 5 mm wide. Column c. 2.5 mm long;

columnfoot appressedto ovary, c. 4.5 mm long.

Fig. 6.

Distribution : Papua New Guinea?

Notes: This species is closely related to

D. gnomus Ames from the Solomon Islands but

has slightly broader leaves, sessile

inflorescences, glabrous (not scabrous) flowers

with the labellum margin entire (not minutely

erose). Dauncey (pers. comm.) kindly confirmed

that the present taxon was undescribed.

Dr. Geoff Stocker obtained this plant from

Papua New Guinea in the early 1970’s but was

unaware of where it was originally collected. It

is an attractive species now fairly common in

Australian orchid collections thanks to Dr.

Stocker’s efforts in propagating it.

Etymology: The specific epithet is derived from

the Greek compound para (near or similar to)

and gnomus (a dwarf), in reference to the

similarity of the species to D. gnomus Ames.

Glomera Blume

There are 41 species in New Guinea so far

recognised for this genus that extends from

Thailand (Seidenfaden & Pedersen 2003) to

Samoa. Several species remain to be described,

some of them quite large plants. The flowers

are quite similar in most species but diagnostic

characters are often readily detected in the

vegetative parts of the plants.

Glomera pseudomonanthos Ormerod sp. nov.

Glomerae subpetiolatae Schltr. arete affinis sed

caulibus 4-7 mm (non 2-3 mm) diam., foliis

oblongis 9-13 mm latis (non ligulatis-

lanceolatis 4.5-7 mm latis), et floribus

virgineis praeter labellum viride apice

atroviridi usque purpureo-nigro

(adversum flores virgineos tantum apice

labelli atroviride) distinguenda. Typus:

Papua New Guinea. Morobe Province:

Dawon, 12 June 1964, A. Millar

NGF23408 (holo: LAE; iso: AMES, BRI,

K; BO, CANB, L n.v.).

Epiphytic or terrestrial herb. Rhizome short. Stem

terete at base, becoming complanate, 29-36.5

cm long and 0.2-0.5 cm wide, laxly 4-8-leaved

or more. Leaves ±oblong, sub-equally and

obtusely bilobed, subpetiolate, 3-6.5 cm long

and 0.9-1.3 cm wide, light to dark glossy green;

sheaths obcuneate, apex truncate, smooth,

lightly striate, 1.1-2.3 cm long and 0.4-0.7 cm

wide. Inflorescences terminal, capitate,

subglobose, c. 1.5 cm wide; subtending sheath

ovate-elliptic, acute, c. 2 cm long and 1.2 cm

wide. Flowers resupinate; pedicel plus ovary

six-ribbed, 5-7 mm long, glabrous; sepals and

petals white to greenish-white, lip green to dark

green with apex dark green to purple-black.

Dorsal sepal broadly elliptic, shortly acuminate,

5-nerved, 4-4.5 mm long and 3-3.5 mm wide;

lateral sepals elliptic-quadrate, very shortly

acuminate, 5-nerved, connate to the back of the

spur, c. 5 mm long and 3.5 mm wide. Petals

obovate-elliptic, very shortly and obtusely

apiculate, c. 4.5 mm long and 3 mm wide; labellum

lamina fleshy, obovate, obtuse, c. 2 mm long

and wide. Spur subglobose, c. 2 mm long.

Column c. 1.6 mm long. Fig. 7.

Additional specimens examined : Indonesia. Papua

Province: Okwalimkam, river headwaters, June 1967,

Ridsdale & Galore NGF33197 (LAE). Papua New

Guinea. Enga Province: Laiagam, Mar 1983, Reeve

1051 (K). Southern Highlands Province: Tari Distr., Komo

Subd., Eanda, Jul 1980, Reeve 2587 (K, NSW).

Distribution and habitat: Indonesia (Papua

Province); Papua New Guinea. Lower montane

forest, 1220-2200 m.

Notes: Glomera pseudomonanthos is a very

distinctive species easily recognised by its

194

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5mm

Fig. 6. Dendrobium paragnomus. A. lateral view of flower. B. lateral view of flower minus sepals and petals. C.

adaxial view of labellum. D. adaxial view of lateral sepal. E. adaxial view of dorsal sepal. F. adaxial view of petal.

All from Stocker sub Ormerod 23 (BRI). A-C and D-F to respective scales.

Ormerod, Papuasian Orchid Studies

195

3mm

5cm

Fig. 7. Glomera pseudomonanthos. A. plant with apical inflorescence. B. adaxial view of dorsal sepal. C. adaxial

view of petal. D. adaxial view of lateral sepal. E. lateral view of flower minus sepals and petals. F. adaxial view of

labellum and spur. A from Millar NGF23408 (LAE); B-F from Ridsdale & Galore NGF33197 (LAE). A and B-F to

respective scales.

196

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flattened stems and leaf sheaths, oblong leaves

and white flowers with a green lip and darker

apex. Only G. subpetiolata Schltr. appears to be

closely related, this has more slender stems,

narrower leaves and white flowers with only

the apex of the lip dark green. One collection of

G subpetiolata was available for comparison,

it was collected near Bupu Village in Morobe

Province, Papua New Guinea [ Millar

NGF22613 (LAE)].

Etymology: The specific epithet is derived from

the Greek compound pseudo (false) and

monanthos, from the compound mono (one),

and anthos (flower). This is in reference to the

vegetative habit that resembles that of certain

Dendrobium species in sect. Biloba J.J.Sm.

(formerly sect. Monanthos Schltr.).

Pseuderia Schltr.

In New Guinea fourteen described species may

be recognized in this genus. Yukawa et al. (1993,

1996) have argued for the removal of this genus

from the Dendrobiinae based on chloroplast

DNA restriction site variation and rbcL

sequences. I agree with Yukawa and his

colleagues since there is also good

morphological evidence for such a proposal.

Pseuderia differs from all Dendrobiinae in

possessing convolute leaves, an elongated

column lacking a columnfoot and the

incompletely divided pollinia. Yukawa et al.

(1993, 1996) suggest placing Pseuderia in the

Podochileae, but admit (1996: 173) that there

is a lack of evidence to support such a move.

The following species is easily recognised

among the known taxa in New Guinea by its

narrow lanceolate leaves.

Pseuderia takeuchii Ormerod sp. nov.

Species nova P. brevifolio J.J.Sm. affinis sed

foliis 3-5plo angustioribus, floribus rubro-

brunneis et labello prope apicem area

densa pubescenti praedito differt. Typus:

Papua New Guinea. East Sepik Province:

Sitipa River headwaters, near Gahom

Village, 5 September 1990, W. Takeuchi

6632 (holo: BRI; iso: AMES).

Epiphytic herb. Roots and lower part of plant

not seen. Stem terete, branching, laxly foliose,

part preserved 32 cm long, in lower part to 3.5

mm thick, in upper part 1-1.75 mm thick;

internodes 4-16 mm long, averaging 8 mm long

on branchlets. Leaves lanceolate, acute, thin,

48.5-60.5 mm long and 3-5.5 mm wide; leaf

sheaths striate, minutely black-brown hairy.

Inflorescences to 5 mm long, arising

suboppositely to leaf lamina, probably

successively one-flowered. Flowers resupinate;

pedicel plus ovary terete, c. 6.5 mm long, red-

brown. Dorsal sepal linear-ligulate, obtuse,

triveined, c. 12 mm long and 2 mm wide; lateral

sepals falcate, ligulate, apiculate, triveined, c.

10 mm long and 2.5 mm wide. Petals slightly

falcate, linear-ligulate, obtuse, triveined, c. 9 mm

long and 1.6-1.75 mm wide; labellum obovate-

oblanceolate, obtuse, apical third laxly

pubescent becoming densely pubescent, lower

half with a keel canaliculate (appearing

bicarinate) for c. 3 mm before it becomes

unicarinate, in total lip c. 8 mm long and 3.5 mm

wide, at very base c. 0.5 mm wide where attached.

Column slender, gently arcuate, c. 5.5 mm long.

Fig. 8.

Distribution and habitat : Papua New Guinea

(East Sepik Province). Alluvial forests with a tall

canopy in the flood zone of the river, c. 100 m.

Notes: This species is similar to Pseuderia

brevifolia J.J.Sm. (also from the New Guinea

north coast) in having short leaves; however, it

differs from that plant in having the leaves 3-5

times narrower, red-brown flowers and a lip with

a dense pubescent patch near its apex. In

P. brevifolia the leaves are wider (1.3-1.9 cm

wide), the flowers are yellowish with red to violet

dots and the lip is loosely papillose to

pubescent.

Etymology: The specific epithet honours W.

Takeuchi, collector of the type specimen.

Robiquetia Gaud.

Five species of this genus are recognised as

occurring in New Guinea, these are:

R. ascendens Gaud. (Syn.: Saccolabium

mooreanum Rolfe), R. camptocentrum (Schltr.)

J.J.Sm., R. flexa (Rchb.f.) Garay (Syn.:

Ornithochilus moretonii F.M.Bailey),

R. gracilistipes (Schltr.) J.J.Sm. and R. hamata

Schltr. Besides the one described below there

appears to be at least one more new taxon that

occurs on New Ireland; however, material is

insufficient to describe it at this time.

Ormerod, Papuasian Orchid Studies

197

5mm

Fig. 8. Pseuderia takeuchii. A. lateral view of flower. B. adaxial view of dorsal sepal. C. adaxial view of petal. D.

adaxial view of lateral sepal. E. adaxial view of labellum. F. column and labellum, the latter unequally folded

longitudinally due to pressing. All from Takeuchi 6632 (BRI). A, B-E and F to respective scales.

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Fig. 9. Robiquetia brassii. A. plant with flowering inflorescence. B. lateral view of flower. C. lateral view of column

and labellum. D. adaxial view of lateral sepal; E. abaxial view of dorsal sepal. F. adaxial view of petal. G. adaxial view

of labellum. H. adaxial view of column (r=rostellum, stp=stipes, v=viscidium). A from Brass 11716 (BRI); B-H

from Kalkman & Nicolas 4159 (AMES). A, B-F, G and H to respective scales.

Ormerod, Papuasian Orchid Studies

Robiquetia brassii Ormerod sp. nov.

Species nova affinis R. camptocentrum (Schltr.)

J.J.Sm. sed floribus labello epichilo

adaxialiter ecalloso non bicalloso differt.

Typus: Indonesia. Papua Province: Balim

River, December 1938, L.J. Brass 11716

(holo: BRI; iso: AMES).

Epiphytic herb. Stem elongate, leafy, at least

15-24 cm long and 0.5-0.85 cm thick but 1.3-2

cm wide across top of sheaths. Roots terete,

elongate, at least 27 cm long and 0.3-0.4 cm

thick. Leaves 3-7 or more, oblong to ligulate,

thickly coriaceous, emarginate to obliquely

truncate, lobes obtuse to truncate, from c. 6 cm

long and 2.1 cm wide to c. 23 cm long and 3.6 cm

wide; sheaths obliquely truncate opposite leaf

lamina, 1.5^1.4 cm long. Inflorescences 13.5-30

cm long, to 0.4 cm thick; peduncle 11-19 cm

long; peduncular sheaths three, scattered,

tubular, clasping, acute, 0.7-1.35 cm long; rachis

2.5-11 cm long, densely 15 to many-flowered;

floral bracts ovate-lanceolate, acute, 0.5-1 cm

long and 0.2-0.3 cm wide. Flowers resupinate;

pedicel plus ovary cylindric to weakly clavate,

laxly covered in short scales, 5.5-6 mm long;

sepals and petals fleshy, red to dirty purple.

Dorsal sepal obovate-elliptic, obtuse, concave,

trinerved, 4.5-6 mm long and 2.5-3.5 mm wide,

sparsely covered in short scales outside; lateral

sepals broadly oblique-elliptic, obtuse, 4-6 mm

long and 2.8-4 mm wide, sparsely covered in

short scales outside. Petals obovate-elliptic,

obtuse, 1-2-nerved, 4-5.5 mm long and 3-4 mm

wide; labellum trilobed, spurred, connate to

ventral sides of the column for c. 1 mm, 4-5 mm

long and 5-6.5 mm wide; hypochile with broadly

rounded sidelobes 2-3.5 mm long and 2-2.5 mm

wide, each inside at front with a fleshy callosity

c. 1 mm long, between sidelobes a low short

median keel terminating at the epichile base;

epichile fleshy, ovate, subacute, 1.5-2 mm long;

spur conical-cylindric, acute to obtuse, straight

to gently geniculate, 5-10 mm long. Column

short, stout, hamate, c. 2 mm long and 1.5-2.5

mm wide laterally. Fig. 9 & 10.

Additional specimen examined: Indonesia. Papua

Province: Star Mts., Sibil Valley, May 1959, Kalkman

& Nicolas 4159 (AMES).

199

Distribution and habitat : Indonesia (Papua

Province). Lower montane forest, epiphyte on

riverbank trees (both records), 1200-1600 m.

Notes : This species is closely related to

Robiquetia camptocentrum (Schltr.) J.J.Sm.

R. brassii is distinguished by the fleshy, entire

labellum epichile which is not split medially

above with each side thickened into an ovoid

to globose callosity. Furthermore, in

R. camptocentrum a distinct sinus is present

between the lateral lobes and the epichile whilst

when these are viewed laterally in R. brassii

this sinus is absent.

Robiquetia hamata is also very similar to

R. brassii that also occurs in the Indonesian

province of Papua [ Eyma 4718 (AMES, BRI,

K) from WisselLakes], However; in R. hamata

the labellum epichile is split medially above with

diverging margins and it is hollowed out like

the bow of a boat whilst in R. brassii the epichile

is solid.

The holotype of R. brassii is a small-leaved

portion of the plant whilst the isotype in AMES

and the other known collection have leaves 3-4

times as long. In rehydrated material the flowers

appear to have a thickened median line on the

exterior of the sepals but this feature is probably

an artifact of drying associated with the fleshy

flowers.

Etymology : The specific epithet honours

L. J.Brass, collector of the type specimen.

TainiaBlume

If Tainia is interpreted in the strict sense then

only one species has been recognised as

occurring in New Guinea until now. The first

taxon reported from New Guinea was Tainia

trinervis (Blume) Rchb.f. which was at first

described by Schlechter under the name

T. parviflora. It is now known to occur in the

Moluccas, New Guinea, Bougainville and

northeast Australia.

The second species that is described

below, was first recognised by Mr. Walter

Kittredge (HUH) but due to an oversight, the

single collection was not loaned when the genus

was monographed by Turner (1992).

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Fig. 10. Robiquetia brassii. A. lateral view of flower. B. lateral view of labellum and column (minus anther and

pollinarium). C. adaxial view of labellum. D. lateral view of column minus anther with pollinia on the stigma. E.

adaxial view of dorsal sepal. F. adaxial view of petal. G. adaxial view of lateral sepal. All from Brass 11716 (AMES).

A, B and C-G to respective scales.

Ormerod, Papuasian Orchid Studies

201

Fig. 11. Tainia serratiloba. A. plant with flowering inflorescence. B. adaxial view of labellum. C. adaxial view of

spread epichile. D. adaxial view of dorsal sepal. E. adaxial view of petal. F. adaxial view of lateral sepal. G. lateral

view of column. H. lateral view of column minus anther and with wings spread out. All from Brass 12308 (AMES).

A and D-G to respective scales. B, C, H not to scale.

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Austrobaileya 7 (1): 183-203 (2005)

Tainia serratiloba Ormerod sp. nov.

Planta pro genere pusilla, inflorescentiis

pauciflora, floribus labello trilobatis

tricarinatis et lobis lateralibus eroso-

laceratis. Typus: Indonesia. Papua

Province: 15 km SW of Bernhard Camp,

Idenburg River, January 1939, L.J. Brass

12308 (holo: AMES).

Epiphytic herb. Roots few, elongate. Rhizome

terete, internodes 8-11 mm long and 1-3 mm

thick; rhizome sheaths tubular, clasping,

obliquely truncate, c. 1 mm long. Pseudobulbs

30-40 mm apart, narrowly conical-cylindric,

unifoliate, fused(?) to rhizome for up to 7.5 mm,

up to 12.5 mm long and 2 mm thick. Leaves ovate-

lanceolate to oblong-lanceolate, acute, 13-

nerved, 48-66 mm long and 11-14 mm wide.

Inflorescences 62-130 mm long; peduncles

resembling pseudobulbs basally, 59-75 mm long

and to 2 mm thick; sheathing bracts three, laxly

spaced, tubular, obliquely truncate, acute, 8-15

mm long; rachis laxly 1-4-flowered, 21-55 mm

long; floral bracts tubular, obliquely truncate,

acute, 6.5-9.5 mm long. Flowers resupinate;

pedicel plus ovary narrowly clavate, 9-12.5 mm

long; sepals brown with yellow tips, lip white.

Dorsal sepal ligulate-oblanceolate, acute, c. 15.5

mm long and 3.5 mm wide; lateral sepals falcate,

oblong-ligulate, acute, median nerve weakly

carinate externally, c. 10 mm long and 2.8 mm

wide. Petals oblong-ligulate, falcate, acute, c.

13 mm long and 3.25 mm wide; labellum trilobed,

c. 7.5 mm long and 6 mm wide, with three

subcrispate-undulate keels each grooved on the

upper edge; hypochile c. 4.5 mm long, with

obliquely subquadrate, erose-lacerate sidelobes

c. 1 mm long and 2 mm wide; epichile

subquadrate, minutely apiculate, c. 3 mm long

and 4.5 mm wide. Column broadly winged,

without anther c. 6.5 mm long, c. 3 mm wide

laterally at apex, c. 1.2 mm wide laterally near

base, lobe of columnwings c. 0.9 mm wide;

pollinia eight. Capsule c. 26 mm long and 6.5

mm thick with pedicel 10 mm long. Fig. 11.

Distribution and habitat : Indonesia (Papua

Province). Lower montane forest, rainforest

gully, c. 1750 m.

Notes : This plant superficially resembles

Collabium pumilum (J.J.Sm.) Seidenf. from

Sulawesi in habit, presumably because it has

also adapted to a montane mossy habitat. Tainia

serratiloba does not appear to have any close

relatives in Tainia and stands alone in the

genus.

Etymology : The specific epithet is derived from

the Latin serratus (serrated or notched), and

the Latin compound lobus derived from the

Greek lobos (a lobe or pod), in reference to the

serrated lateral lobes of the labellum.

Acknowledgements

I wish to thank W. Kittredge (HUH), G. Romero

(AMES), G. Stocker, D.L. Szlachetko and T.

Yukawa (TNS) for their help with various

matters. The Directors of the cited herbaria

allowed access to collections in their care. P.

Bostock (BRI) contributed the majority of the

Latin diagnoses whilst R.J.F. Henderson (ex

BRI) contributed the Latin diagnoses of

Glomera pseudomonanthos and Robiquetia

brassii.

References

Chadim, V.A. (1972). Dendrobium macrophyllum

A.Rich. Orchadian 4: 25-31.

Cribb, RJ. (1983). A revision of Dendrobium sect.

Latouria. Kew Bulletin 38: 229-306.

Dauncey, E.A. (2003). A taxonomic revision of

Dendrobium section Pedilonum

(Orchidaceae). Harvard Papers in Botany 7:

151-320.

Greuter, W., McNeill, J., Barrie, F.R., Burdet, H.M.,

Demoulin, V., Filgueiras, T.S., Nicolson, D.H.,

Silva, P.C., Skog, J.E., Trehane, R, Turland, N.J.

& Hawksworth, D.L. (2000). International

Code of Botanical Nomenclature (Saint Louis

Code). Regnum Vegetabile 138: 1-474.

Kraenzlin, F. (1910). Orchidacea e-Monandreae-

Dendrobiinae Part I. In A.Engler (ed ). Das

Pflanzenreich IV. 50. II. B. 21, 45: 1-382.

Lavarack, P.S., Harris, W. & Stocker, G. (2000).

Dendrobium and its Relatives. Kangaroo Press:

Roseville East.

Ormerod, P. (1995). Complementary studies of

Solomon Archipelago orchids. Australian

Orchid Review 60: 35-39.

Ormerod, P. (2002). Notes on the orchids of New

Guinea and the Pacific. Part 7. Oasis (Dora

Creek) 2(3): 8-9, 12.

Ormerod, P. (2003). Papuasian Orchid Studies. Oasis

(Dora Creek) 3(4): 3-6.

Ormerod, Papuasian Orchid Studies

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Reichenbach, H.G. (1861). Index. In W.G. Walpers (ed ),

Annales Botanices Systematicae 6: 1150.

Schlechter, R. (1912). Dendrobium Sw. Repertorium

specierum novarum regni vegetabilis, Beihefte

1; 440-643.

Seidenfaden, G & Pedersen, H.AE. (2003). Contributions

to the orchid flora of Thailand XIV. Nordic

Journal of Botany 22: 525-534.

Schuiteman, A. & De Vogel, E.F. (2001). Orchids of New

Guinea. Vol. I. Illustrated Checklist and

Genera. CD-ROM. ETI/National Herbarium

of the Netherlands: Leiden.

Turner, H. (1992). A revision of the orchid genera

Ania, Hancockia, Mischobulbum and Tainia.

Orchid Monographs 6: 43-100 + figures.

Van Royen, P. (1979). Orchidaceae. In The Alpine

Flora of New Guinea. 2: 51-811. J.Cramer:

Vaduz.

Yukawa, T., Kurita, S., Nishida, M. & Hasebe, M. (1993).

Phylogenetic implications of chloroplast DNA

restriction site variation in Subtribe

Dendrobiinae (Orchidaceae). Lindleyana 8:

211 - 221 .

Yukawa, T., Ohba, H., Cameron, K.M. & Chase, M.W.

(1996). Chloroplast DNA phylogeny of

Subtribe Dendrobiinae (Orchidaceae): Insights

from a combined analysis based on rbcL

sequences and restriction site variation.

Journal of Plant Research 109: 169-176.

Goodenia elaiosoma I.D.Cowie (Goodeniaceae), a new species from the

Top End of the Northern Territory and a key to the northern species

IanD.Cowie

Summary

Cowie, I D. (2005). Goodenia elaiosoma I.D.Cowie (Goodeniaceae), a new species from the Top

End of the Northern Territory and a key to the northern species. Austrobaileya 7(1): 205-213.

The new species Goodenia elaiosoma I.D. Cowie ( Goodenia section Goodenia , subsection

Borealis ) is described from the western Top End of the Northern Territory. The species is

compared with the superficially similar, tropical species G. armstrongiana and G. debilis. A key to

the 43 Goodenia species found in the northern Northern Territory is provided.

Key Words: Goodeniaceae, Goodenia elaiosoma, Goodenia armstrongiana, Goodenia debilis,

new species, key to species. Northern Territory, tropical Australia.

I.D.Cowie, Northern Territory Herbarium, Department of Natural Resources, Environment and

the Arts, P.O. Box 496, Palmerston Northern Territory 0831, Australia.

Introduction

The seasonally waterlogged, infertile, sandy

soils of the monsoonal northern Northern

Territory (NT) support a diverse assemblage of

specialist plant species. Indeed, the area has

been identified as a world centre of diversity

for the genus Utricularia L. (Lentibulariaceae)

(Taylor 1989) with 36 taxa known from the area.

This diversity is also reflected in other genera

associated with such soils including Eriocaulon

L. with 21 species in the northern NT (Leach

2000) and Lindernia All. (under revision but

more than 25 taxa in the northern NT (W.R.

Barker unpublished data). To a greater or lesser

degree, speciation has also occurred in genera

such as Byblis Salisb. Calandrinia Kunth,

Centrolepis Labill., Drosera L., Fimbristylis

Vahl, Mitrasacme Labill., Oldenlandia L.,

Stylidium Willd., Trithuria Hook.f., and

Typhonium Schott occurring on seasonally

water logged substrates and extends to the

wetter end of the drainage continuum with local

speciation in shallow water, aquatic genera such

as Hygrochloa Lazarides and Nymphoides Hill.

Many of the taxa involved are endemic to the

NT, with relatively restricted distributions on

the lowlands primarily associated with the

Koolpinyah Surface of Williams (1969) or the

Western Arnhem Land Plateau (Woinarski et

al. in prep.). The species composition of this

Accepted for publication 10 August 2005

flora appears to vary in response to subtle

changes in the texture, drainage and period of

inundation of these colluvial and alluvial sandy

soils found in drainage depressions, on

upstream floodplains, along minor drainage

lines and in seepage areas.

In the Darwin area, these soils are

extensively exploited as a source of fine sand

for building and construction (Doyle 2001) and

the long-term conservation status of the flora

they support is of some concern. During

surveys of this flora in the Darwin-Kakadu area

to clarify the distribution, abundance and

conservation status of Utricularia species in

particular, collections were made of an

undescribed Goodenia Sm. This taxon,

although allied to G. armstrongiana de Vriese

and G. debilis A.E.Holland & T.P.Boyle, has

characters which clearly set it aside from those

species and is described here.

Goodenia Sm. is a genus of c. 190 species

almost endemic to Australia and occurring in all

States and Territories. Seventy-two species are

known from the NT. The genus has recently

been treated for Australia by Carolin (1990;

1992), although no treatment exists specifically

for the NT. A key to the 43 Goodenia species

found in the NT north of 17° S latitude is

provided here to assist with their identification.

206

Austrobaileya 7 (1): 205-213 (2005)

Materials and methods

Morphological characters were examined using

fresh and dried material at DNA. Floral

characters were primarily measured from

rehydrated and spirit-preserved material with

supplementary measurements of dried flowers.

The terminology used for Goodenia

follows that generally accepted for the genus

(Carolin 1990,1992; Holland & Boyle 2002). In

particular, the more or less reduced cauline

leaves or scales which subtend the flowers are

termed bracts. Appendages (bracteoles) may

also be present on the flower stalk, which if

ebracteolate is termed a pedicel. Where

bracteoles are present, that part below the

bracteole is termed a peduncle and the part

above a pedicel. The term calyx lobes is used

here to describe the free parts of the calyx,

otherwise known as sepals in the literature. The

adaxial corolla lobes are those either side of the

split in the side of the corolla tube while the

abaxial corolla lobes are the three more connate

lobes between them. Species in the key are

numbered according to the Flora of Australia

treatment (Carolin 1992), to allow easy reference

to descriptions there and users of the key are

referred to that treatment for authors of plant

names.

Taxonomy

Goodenia elaiosoma ID. Cowie, sp.nov. Arete

affinis G. debili A.E. Holland & T.P. Boyle

a corolla purpureo-brunnea et praecipue

a semine oblongo cum hilo basali et

elaiosomate, sed sine labro a qua differt.

Typus: Northern Territory. E of Hope Inlet,

12°25’S, 131° 06’E, 3 June 2000, ID. Cowie

8891 (holo: DNA; iso: AD, B, BRI, CANB,

K, L, MEL, MO, NSW, NY, PERTH).

Decumbent annual herb, stems to 70 cm long,

base single stemmed, softly spongy.

Indumentum of white to stramineous, straight,

antrorse, closely appressed hairs 0.3-0.7 mm

long, sparse on stems, pedicels, leaf

undersurfaces and capsules, more dense on new

growth, ovary, calyx and exterior of floral tube;

inner surface of corolla in lower half with fine,

weak, hyaline hairs. Leaves alternate, antrorse,

all cauline, sessile, linear, 6-35 mm long, 1-2

mm wide, gradually diminishing in length

towards the stem apex, glabrous above, margins

entire, slightly recurved. Inflorescence a

bracteate raceme. Flowers solitary; bracts

foliaceous; pedicels filiform, patent, 10-18 mm

long, bracteoles absent. Sepals adnate to ovary

almost to apex; lobes linear, 1.0-1.7 mm long.

Corolla 5-7 mm long, purple brown, end of

lobes and wings cream to dark maroon, enations

absent, pouch absent; tube 1.2-1.9 mm long;

abaxial lobes narrowly oblong, 3.3^1 mm long,

c. 0.8 mm wide, free for 1.7-2.5 mm, wings equal,

1.3- 1.7 mm long, 0.8-1.3 mm wide, minutely

serrate to erose; adaxial lobes free, narrowly

oblong, auricle 1-1.3 mm long, c. 0.7 mm wide,

wings strongly unequal, wing above auricle

usually rudimentary for most of its length, 0.5-

1.7 mm long, 0.1-0.5 mm wide, opposite wing

well developed, 1.2-1.5 mm long, 0.7-1 mm wide,

minutely serrate to erose. Stamens 2.1-2.3 mm

long; filaments linear, 1.4-1.8 mm long; anthers

narrowly oblong, 0.4-0.5 mm long, apiculate.

Style 22-2.1 mm long, indusium broadly

obovate, 0.6-1 mm long, 0.8-1 mm wide, basal

tuft of hairs absent, sometimes with a few

subapical hairs to 0.5 mm long, otherwise

glabrous, apex concave, bristles 0.1-0.5 mm

long, to 0.5 mm long on upper (abaxial) lip. Ovary

inferior; ovules 6-7. Capsule ellipsoidal, 3.7-

4.5 mm long, dehiscing deeply. Seed

stramineous, oblong, biconvex in cross-section,

1.4- 2.0 mm long, 0.9-1.1 mm wide, smooth,

glossy, obscurely areolate, rimless, wing absent,

base asymmetric with a hyaline elaiosome. Fig.

1 .

Additional specimens examined : Northern Territory.

Wangi Road, near Finniss River crossing, 12° 58’S,

130° 45’E, May 2004, Cowie 9993 & Jacka (BRI,

CANB, DNA, L, NT, MEL, MO); Shoal Bay area, E of

Hope Inlet, NT, 12° 25’S, 131° 06’E, May 2000, Cowie

8885 & Kerrigan (BRI, CANB, DNA, MEL, MO,

NSW); Howard River Floodplain, 12° 28’S, 131° 06’E,

May 2001, Kerrigan 341 & Cowie (DNA, MEL); 19

km NNW of Twin Falls, 13° 08’S, 132° 45’E, Jun 1980,

Craven 6293 (DNA).

Distribution and habitat : Endemic to the

northern NT and known from Finniss River

southwest of Darwin to Kakadu National Park

(Map 1). Grows in Eriachne burkittii grassland/

sedgeland or Pandanus low open woodland,

on the margins of non-estuarine floodplains or

in seasonally inundated drainage depressions,

on sandy substrates.

Cowie, Goodenia elaiosoma

207

Fig. 1. Goodenia elaiosoma. A. habit. B. indumentum of stem. C. flower. D. capsule. E. seed, all from Cowie 8891

(DNA). Scale bars: A. 1 cm; B. 0.5 mm; C, D, E.l mm. Del. M. Osterkamp Marsden.

208

Austrobaileya 7 (1): 205-213 (2005)

NT.

Phenology : Flowering and fruiting are recorded

from April to June.

Notes : Goodenia elaiosoma is most closely

related to G debilis but differs in having purple-

brown flowers and particularly in the seed which

is oblong, lacks a rim and has a basal hilum and

elaiosome (Fig. 2). G. debilis has cream or yellow

flowers with brownish markings and the seed is

elliptic to elliptic-lanceolate, has a narrow rim,

with the hilum obliquely lateral at base (Fig. 2).

Although specimens of both G debilis and

G armstrongiana have a narrow membrane

around the hilum, this is apparently air-filled

and not filled with oily material as in

G. elaiosoma. G. elaiosoma shares with

G debilis narrow entire leaves, a shorter corolla,

few ovules and an appressed indumentum. Both

species lack the prominent basal tuft of hairs

on the indusium.

Goodenia elaiosoma differs from

G armstrongiana especially by the reduced

wing on the adaxial corolla lobes, purple-brown

corolla, and the smaller, essentially smooth,

oblong, biconvex, rimless, glossy seed with a

basal elaiosome. Other differences include the

stem which is single and softly spongy at the

base; linear leaves; straight, closely appressed

indumentum; indusium lacking an often

prominent basal tuft of hairs; usually smaller

corolla, and fewer ovules. Although variable in

leaf dimensions and flower colour,

G armstrongiana is characterised interalia by

antrorse leaves which are more or less dentate,

often with basal auricles, a sparse indumentum

of curved, appressed to ascending strigose

hairs, long filiform peduncles, white to yellow

corolla, an indusium with a prominent basal tuft

of hairs to 1.5 mm long and seeds which are

obovate-oblong to oblanceolate, flattened,

verrucate, with a well developed rim, the hilum

obliquely lateral at the base (Fig. 2). This species

is multistemmed at the base with wiry stems.

G elaiosoma is sometimes sympatric with

G armstrongiana (e.g. at Finniss River), where

both species maintain their identities without

forming hybrids. Populations of G elaiosoma

are geographically disjunct from those of the

north Queensland G debilis.

Like Goodenia armstrongiana and

G debilis , G elaiosoma belongs in G section

Goodenia , subsection Borealis Carolin (Carolin

1992).

Conservation status : The species is known

from five locations, with a range extending some

250 km. Two populations fall within Shoal Bay

Conservation Reserve east of Darwin while a

third lies in Kakadu National Park. A fourth

population is known from an area previously

disturbed by mining. Although plants at each

population were locally common, with plants at

Finniss River abundant and estimated at

approximately 1 plant m 2 over an area of several

hectares, little is known of the true size and

extent of populations. Outside of the two

reserves, the poorly drained sands inhabited

by this species in the Darwin area have been

extensively taken up for mining and are heavily

exploited. Despite extensive survey work in

suitable habitat in the Darwin area, no other

populations were found. However, large areas

Cowie, Goodenia elaiosoma

209

Fig. 2. Seeds of Goodenia armstrongiana, G. debilis and G. elaiosoma. From left to right A. G. armstrongiana

(Orr 222). B. G. armstrongiana (Cowie 8576). C. G. debilis (Forster 22614). D. G. elaiosoma (Cowie 8891).

of its range to the east of Darwin and in Kakadu Etymology: From the Greek, elaion - oil and

National Park have not been intensively soma - a body, a reference to the elaiosome

surveyed at an appropriate scale or time of year. found on the seed.

Using IUCN criteria, a conservation code of

Data Deficient is recommended (IUCN 2001).

Key to Goodenia species in the Northern Territory north of 17°S latitude

Species in the key are numbered according to the Flora of Australia treatment (Carolin 1992), to

allow easy reference to descriptions there and users of the key are referred to that treatment for

authors of plant names. It is recognized that a number of complexes are in need of further study (e.g.

G. janamba Carolin - G. coronopifolia R.Br., G. byrnesii Carolin - G. malvina Carolin - G. campestris

Carolin and G. leiosperma Carolin - G sp. Melville Island).

1. Style 2-4 fid. GROUP 1

Style unbranched.2

2. Herbs with stoloniferous stems or plants compact and cushion-like; flowers

2 mm long or less, dark red to red-purple, lobes equal, wings obsolete or

absent. GROUP 2

Decumbent to erect herbs, stems not stoloniferous; flowers almost always

more than 2 mm long, blue, purple-brown to yellow or white; wings usually

well developed, rarely absent.3

3. Leaves either mostly clustered on a short rosette-like basal stock or scattered

along a short stem; inflorescence distinct with few leaf-like bracts (i.e.

scapose or scapiform). GROUP 3

Stems foliose throughout or with leaf-like bracts predominant in

inflorescence (stems sometimes reduced or sometimes a few basal leaves

present). GROUP 4

210

Austrobaileya 7 (1): 205-213 (2005)

GROUP 1

1. Style 4-fid; corolla yellow-brown. 176. G quadrifida

Style 3-fid; corolla yellow, purple or brownish-purple.2

2. Corolla yellow, at least on wings.3

Corolla purple or brownish-purple .6

3. Simple hairs predominating on young leaves, bracts and pedicels, rarely

almost glabrous.160. G pilosa

Glandular hairs predominating on young leaves, bracts and pedicels.4

4. Seeds smooth; corolla with simple strigose hairs outside, a few glandular

hairs sometimes present.170. G neglecta

Seeds granulate to verrucate; corolla glandular-pubescent outside.5

5. Cauline leaves usually with 1 or 2 acute teeth at the base, thus auriculate;

central indusium ovate to broad ovate; corolla (5.5-) 14-20 mm long .... 172. G holtzeana

Cauline leaves petiolate or sessile, never with two broad basal teeth; central

indusium depressed-ovate; corolla 10-12 mm long. 173. G heppleana

6. Seeds granulate to verrucate; corolla glandular pubescent outside, simple

strigose hairs absent. 175. G purpurea

Seeds smooth; corolla with simple strigose hairs outside, glandular hairs

also present.7

7. Glandular hairs numerous and conspicuous on the flower and pedicel; leaves

oblanceolate to narrowly oblanceolate; corolla 8-15 mm long . 174. G symonii

Glandular hairs few and inconspicuous on the flower and pedicel; leaves

very narrowly lanceolate to linear; corolla 7-8 mm long. 168. G porphyrea

GROUP 2

1. Leaves to 80 mm long; inflorescence a compact head at ground level 165. G chthonocephala

Leaves to 25 mm long; flowers solitary or in condensed terminal racemes.2

2. Hairs on leaves simple. 178. G kakadu

Hairs on leaves stellate. 177. G pumilio

GROUP 3

1. Leaves scattered along a short stem.2

Leaves mostly on a short rosette-like basal stock.3

2. Leaves linear, less than 5 mm wide; capsule cylindrical; seeds reticulate-

foveolate. 13. G gloeophylla

Leaves oblanceolate, more than 10 mm wide; capsule ovoid; seeds colliculate-

punctate. 27. G scaevolina

3. Seeds much less than 1 mm diam., numerous in capsule; bracteoles present.4

Seeds more than 2 mm diam., often few in capsule; bracteoles absent.10

4. Corolla lobes white to purple or blue.5

Corolla lobes yellow or yellow and purple .8

5. Corolla 2-3(-5) mm long mostly white with pale purple markings; corolla

wings almost obsolete; seeds smooth. 19. G minutiflora

Corolla 4 mm or more long, wings 1-2 mm wide .6

Cowie, Goodenia elaiosoma

211

6. Leaves resinous-viscid with glandular and simple hairs; corolla dark purple-

blue; capsule 1.5-3 mm long; seeds smooth to reticulate. 20. G viscidula

Leaves not viscid, glabrous or with simple hairs; corolla almost white to

purple; capsule 2-5 mm long; seeds smooth.7

7. Corolla purple to white-mauve, 8-13 mm long, wings 1-2 mm wide .... 18. G purpurascens

Corolla bluish-purple, 4-5 mm long; wings c. 1 mm wide. 21. G paludicola

8. Plant annual; scapes with glandular and patent simple hairs; abaxial corolla

lobes yellow, adaxial corolla lobes purplish; leaves usually obovate; seeds

reticulate-foveolate. 16. G bicolor

Plant usually perennial; scapes glabrous or only with simple hairs; corolla

lobes yellow throughout; leaves mostly linear-lanceolate to linear-

oblanceolate; seeds smooth.9

9. Leaf veins obscure; indusium notched and folded longitudinally. 11. G. gracilis

Leaf veins 3, prominent; indusium concave. 15. G lamprosperma

10. Calyx lobes markedly unequal, adaxial calyx lobe much longer than others;

seeds reticulate, wing 0.2-0.4 mm wide; dwarf plant. 136. G redacta

Calyx lobes all equal, adaxial calyx lobe equal to others; seeds smooth or

reticulate, wing to 1 mm wide; erect plant.11

11. Leaves linear to oblong-elliptic, dentate to pinnatisect, lobes acute; indusium

obdeltoid; seeds reticulate, wing c. 0.5 mm wide. 106. G coronopifolia

Leaves narrowly oblong to oblanceolate with a few blunt teeth; indusium

broadly oblong; seeds smooth, wing c. 1 mm wide. 130. G janamba

GROUP 4

1. Cauline leaves and bracts appressed to stem, antrorse (pointing upwards)

. 166. G armstrongiana

Cauline leaves and bracts spreading to patent, antrorse or not.2

2. Ovary with mostly glandular hairs or calyx lobes attached in lower half of

capsule or both.3

Ovary with mostly simple hairs (often tubercular based) or

glabrous; calyx lobes attached in upper half of capsule.8

3. Cauline leaves terete or linear, less than 2.5 mm wide.4

Cauline leaves linear to obovate, more than 3 mm wide .6

4. Upper stems strongly flexuose (zig-zag); ovules 4-6; seeds aculeate (covered

in prickles), wing 0.1-0.2 mm wide.138. G cirrifica

Upper stems more or less straight; ovules more than 20; seeds colliculate

(covered with small rounded projections), wing 1 mm or more wide.5

5. Plant single stemmed at base; leaves mostly more than 40 mm long, not

viscid and varnished; abaxial corolla lobes often wingless or with reduced

wings. 141. G triodiophila

Plant many-stemmed at base; leaves mostly less than 40 mm long, often

viscid and varnished; abaxial corolla lobes with well developed wings . 139. G armitiana

212

Austrobaileya 7 (1): 205-213 (2005)

6. Annual herb; leaves and stems with simple hairs only; corolla 11-17 mm

long; northern. 162. G potamica

Perennial herb or subshrub; leaves and stems with glandular hairs; corolla

20-25 mm long; southern.7

7. Leaves ovate to orbicular; corolla yellow; seeds verrucose (covered with

small wart-like projections), wingless; southern Gulf. 46. G grandiflora

Leaves oblanceolate to obovate; corolla blue to mauve; seeds colliculate-

punctate, wing c. 0.1 mm wide; Victoria River area. 27. G scaevolina

8. Calyx lobes markedly unequal, adaxial calyx lobe much longer than others;

seeds reticulate, wing 0.2-0.4 mm wide. 136. G redacta

Calyx lobes all equal, adaxial calyx lobe equal to others; seeds smooth to

verrucose or aculeate, wing usually absent.9

9. Flowers sessile; corolla to 5 mm long; seeds verrucose, granulose . . . 164. G subauriculata

Flowers pedicellate; corolla more than 5 mm long; seeds various.10

10. Indusium folded longitudinally, with furrow on upper surface; corolla 5-7

mm long; seeds aculeate (covered in prickles), reticulate, wing 0.1-0.2 mm

wide. 137. G odonnellii

Indusium concave, not folded (rarely grooved); corolla 7 mm or more long.11

11. Calyx lobes 7-12 mm long and 1.5-2 wide.12

Calyx lobes less than 7 mm long or less than 1 mm wide or both (rarely more

than 7 mm long but then 1 mm wide or less) .13

12. Seeds verrucose to aculeate. 154. G sepalosa

Seeds smooth. 158. G durackiana

13. Seeds smooth to minutely colliculate-alveolate or reticulate.14

Seeds verrucose, granulose .18

14. Seeds biconvex, lacking a rim or wing, basal elaiosome present; corolla

purple-brown; calyx lobes less than 2 mm long. G elaiosoma

Seeds flattened or concavo-convex, rimmed or winged, lacking an elaiosome;

corolla more or less yellow; calyx lobes usually more than 2 mm long .15

15. Cauline leaves entire, petiolate, base long attenuate; seeds reticulate,

prominently winged.108. G strangfordii

Cauline leaves and bracts usually dentate, sessile or with a short petiole,

base rounded to cordate; seeds smooth or minutely colliculate-alveolate,

rimmed but almost wingless.16

16. Inner surface of corolla densely pubescent; calyx lobes 5-11 mm long; corolla

15-23 mm long; abaxial corolla lobes 5-8 mm long; capsule 5-8 mm diam. 157. G leiosperma

Inner surface of corolla with few hairs; calyx lobes 2-5 mm long; corolla less

than 14 mm long; abaxial corolla lobes 2-5 mm long; capsule less than 5 mm diam.17

17. Stems glabrous, ovary glabrous or with a few strigose hairs. 167. G argillacea

Stems with scattered hairs, ovary densely hairy . . G sp. Melville Island (N.B.Byrnes 666)

18. Cauline leaves and bracts shortly petiolate, auricles absent or inconspicuous;

calyx lobes 1.2-2 mm wide. 156. G brachypoda

Cauline upper leaves and bracts sessile and often conspicuously auriculate;

calyx lobes 1 mm or less wide.19

Cowie, Goodenia elaiosoma

213

19. Corolla 7-14 mm long; stems glabrous or with a few coarse hairs .20

Corolla 15-25 mm long; stems sparsely hirsute to hispid, rarely glabrous.21

20. Calyx lobes 1-2 mm long; corolla yellow, 7-8 mm long, abaxial corolla lobes

shorter than their connate part . 160. G campestris

Calyx lobes 2-A mm long; corolla mauve, sometimes more or less yellow

towards top, 9-14 mm long; abaxial corolla lobes at least as long as their

connate part. 161. G malvina

21. Capsule ellipsoidal; calyx lobes 0.4-0.6(-0.8) mm wide, hispid; leaves (7-)

10-20 mm wide.163. G hispida

Capsule more or less globular; calyx lobes 0.8-1 mm wide, with a few marginal

bristles; leaves 5-10 mm wide.159. G byrnesii

Acknowledgements

I wish to thank Clyde Dunlop for valuable

discussions in early stages of preparation of

this paper. Raelee Kerrigan and Bob Harwood

first brought the species to my attention and

assisted with fieldwork. Sally Jacka kindly

showed me the Finniss River population. The

key to species is derived, in part, from an as yet

unpublished key to 14 species found in the

Darwin Region prepared by Clyde Dunlop, Greg

Leach and myself. Annette Wilson offered

guidance during my short visit to Kew, whilst

Laurie Jessup and Alex Chapman during their

respective terms as ABLO searched at Kew for

the type of G armstrongiana and literature on

my behalf. Dale Dixon and Raelee Kerrigan read

and commented on the manuscript. Monica

Osterkamp Madsen prepared the illustration.

References

Carolin, R.C. (1990). Nomenclatural notes and new

taxa in the genus Goodenia (Goodeniaceae).

Telopea 3: 517-570.

Carolin, R.C. (1992). Goodenia. Flora of Australia

35: 147-281. CSIRO Publishing: Melbourne.

Doyle, N. (2001). Extractive minerals within the outer

Darwin area. Report 14. Northern Territory

Geological Survey: Darwin.

Holland, A.E. & Boyle, T.P. (2002). Four new species

of Goodenia Smith (Goodeniaceae) from

Queensland. Austrobaileya 6: 253-266.

Iucn (2001). IUCN Red List Categories and Criteria:

Version 3.1. IUCN Species Survival

Commission. IUCN: Gland and Cambridge.

Leach, GJ. (2000). Notes and new species of Eriocaulon

(Eriocaulaceae) from Australia. Australian

Systematic Botany 13: 755-772.

Taylor, P. (1989). The Genus Utricularia - A taxonomic

monograph. Her Majesty’s Stationary Office:

London.

Williams, M.A.J. (1969). Geomorphology of the

Adelaide-Alligator area. In R. Story, M.A.J.

Williams, A.D.L. Hooper, R E. O’Ferrall &

J.R. McAlpine (eds.). Lands of the Adelaide-

Alligator Area, Northern Territory , pp. 71-94.

Land Research Series No. 25. CSIRO:

Melbourne.

Woinarski, J.C.Z., Hempel, C., Cowie, I., Brennan, K.,

Kerrigan, R., Leach, G. & Russell-Smith, J. (in

prep.) Distributional patterning of restricted

range plant species in the Northern Territory,

Australia.

Studies in Euphorbiaceae s.lat. 7

Shonia R.J.F.Hend. & Halford ( Ricinocarpeae , Ricin o car pin ae),

a new Australian endemic genus

David A. Halford & Rodney J.E Henderson

Summary

Halford, D.A. & Henderson, R.J.F. (2005). Studies in Euphorbiaceae s.lat. 7. Shonia R.J.F.Hend.

& Halford {Ricinocarpeae, Ricinocarpinae ), a new Australian endemic genus. Austrobaileya 7(1):

215-228. The new, Australian endemic genus Shonia R.J.F.Hend. & Halford, is described and

compared with Beyeria Miq. and Ricinocarpos Desf., to which it is closely allied, and Bertya

Planch. Four species are recognised and a key is provided for their identification. Two species,

Shonia carinata Halford & RJ.F.Hend. and S. territorialis Halford & R.J.F.Hend., are described as

new, and the new combinations S. tristigma (F.Muell.) Halford & R J.F.Hend. and S. bickertonensis

(Specht) Halford & R.J.F.Hend., based on Beyeria tristigma F.Muell. and Beyeria bickertonensis

Specht respectively, are made for the other two. Two new subspecies are recognised in S. tristigma ,

namely S. tristigma subsp. parvifolia Halford & R.J.F.Hend. and S. tristigma subsp. borealis

Halford & RJ.F.Hend. All species are described, maps provided for their distribution, and information

on their phenology and the habitat of their occurrence is given. The new species are illustrated

and a lectotype is chosen for Beyeria tristigma F.Muell.

Key Words: Euphorbiaceae, Ricinocarpeae , Ricinocarpinae, Shonia , Shonia bickertonensis, Shonia

carinata, Shonia territoralis, Shonia tristigma, Shonia tristigma subsp. borealis, Shonia tristigma

subsp. parvifolia, Beyeria sect. Oxygyne, Australian flora, taxonomy

D.A. Halford & R.J.F. Henderson, cl- Queensland Herbarium, Environmental Protection Agency,

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

Introduction

Following a review of the attributes of taxa

included in the Australian endemic genus

Beyeria Miq., it became clear that the taxon

Ferdinand Mueller called B. tristigma in 1868,

and included as the only species in his Beyeria

sect. Oxygyne , differs significantly in attributes

of its inflorescence and female flowers from

those characteristic of B. viscosa Miq., which

includes the type of the genus name Beyeria

Miq. Beyeria tristigma agrees more with

Ricinocarpos Desf in respect to its female floral

attributes than to Beyeria while still retaining

attributes in common with the latter genus.

Segregation of this taxon as a distinct genus

seems warranted because of these fundamental

differences, and the existence of other taxa

possessing floral attributes similar to those of

B. tristigma. This idea of segregation is not

entirely new. Bentham (1873: 68) was clearly

perplexed as to where to place B. tristigma for

he noted “the racemose male flowers, the petals

Accepted for publication 16 August 2005

much longer than the calyx, and the divided

stigma or style bring this species near to

Ricinocarpos , but the stamens are entirely those

of Beyeria , and the stigmatic lobes are closely

recurved as in that genus, to which on the whole

it appears to be the nearest related”. However;

Bentham, like Mueller (1868) before him,

considered that it was sufficiently distinct from

the other Beyeria species then recognised to

accord it sole membership of a distinct section

of this genus that Mueller had named Beyeria

sect. Oxygyne. Bentham characterised this

section as having “Anthers of Beyeriopsis”,

which he had previously accepted {op. cit. : 63,

66) as Beyeria sect. Beyeriopsis, the “stigma

deeply divided into 3 narrow lobes”, which he

later described as flat and recurved, and “male

flowers in a loose raceme”.

In his account of Beyeria , Griming (1913)

recognised the uniqueness of B. tristigma and

accepted it as a “transition form” from

Ricinocarpos to Beyeria but persisted in

including it in Beyeria.

216

Austrobaileya 7 (1): 215-228 (2005)

From their overall morphology as well as

their porate pollen grains with a “crotonoid”

pattern of polygonally arranged sexinous

processes, Beyeria tristigma and its allies are

clearly related to Beyeria and to Ricinocarpos

and thus belong with them in Mueller

Argoviensis’s tribe Ricinocarpeae , and within

that to subtribe Ricinocarpinae Webster

(Webster 1994). We have come to the

conclusion that these species would be as out

of place in Ricinocarpos , as they are in Beyeria ,

and that they appear to occupy a morphological

position between those two genera. We have,

therefore, established the genus Shonia to

accommodate them. The structure of their

inflorescences and flowers provide the main

characters to distinguish them from Beyeria and

Ricinocarpos.

Shonia differs from Beyeria by the

following combination of characters:

inflorescences racemose with usually one or

two female flowers basal on the axis with few to

several male flowers distal to them, the female

flowers with styles with three horizontally

spreading and entire stigmatic lobes sometimes

becoming decurved or recurved with age.

Shonia differs from Ricinocarpos by the

following combination of characters: male

flowers with anthers terminal and transverse on

the filaments, an androecium on a flat or slightly

convex receptacle, and female flowers with

entire stigmatic lobes.

Recent molecular studies of the

uniovulate Euphorbiaceae (Wurdack etal. 2005)

suggest a close relationship between Beyeria ,

Ricinocarpos and Bertya Planch, (tribe

Ricinocarpeae , subtribe Bertyinae Mtill.Arg.),

another endemic Australian Euphorbiaceae

genus. For a complete revision of Bertya see

Halford and Henderson (2002). A comparison

of important diagnostic differences between

Bertya , Beyeria , Ricinocarpos and Shonia is

provided in Table 1.

Key to Australian genera of Euphorbiaceae tribe Ricinocarpeae based on female flowers

1. Stigma entire, dilated and forming a cap over the top of the ovary, discoid or

rarely with 2 appressed entire limbs or shallowly 3-lobulate. Beyeria

Stigma with 3 horizontally spreading and divergent limbs sometimes

becoming decurved or recurved with age; limbs entire or 2- to 5-lobed .2

2. Stigmatic limbs entire.Shonia

Stigmatic limbs divided.3

3. Disc present.Ricinocarpos

Disc absent. Bertya

Key to Australian genera of Euphorbiaceae tribe Ricinocarpeae based on male flowers

1. Flowers with staminal filaments erect to spreading from a central cylindrical

column formed by fusion of the bases of filaments.2

Flowers with staminal filaments erect to spreading on a flat or slightly convex to

hemispherical receptacle, with filaments free.3

2. Petals conspicuous, or if absent then calyx spreading at anthesis.Ricinocarpos

Petals absent or rudimentary, calyx strongly recurved at anthesis. Bertya

3. Flowers in racemes.Shonia

Flowers in few flowered fascicles or cymose clusters, or solitary. Beyeria

Halford & Henderson, Revision of Shonia

217

Table 1. Morphological comparison oiBertya, Beyeria, Ricinocarpos and Shonia

Character

Bertya

Beyeria

Ricinocarpos

Shonia

arrangement of

flowers

paired or in

umbelliform clusters,

or solitary

in fascicles or cymose

clusters, or solitary

in umbelliform

clusters, or racemes,

or solitary

in racemes

male flowers:

petals

absent or rudimentary

present or absent

present

disc

absent

present or absent

present

stamens

spreading from central

column formed by

fusion of bases of

filaments

erect on flat to

hemispherical

receptacle

erect to spreading

from central column

formed by fusion of

bases of filaments

erect on flat or

slightly convex

receptacle

position of anthers

in relation to

filament

distal and longitudinal

terminal and transverse

or distal and

longitudinal

distal and longitudinal

terminal and

transverse

female flowers:

petals

absent or rudimentary

present or absent

present or rarely

absent

present

disc

absent

present or absent

present

stigmatie limbs:

degree of division

deeply 2- to several-

lobed

elobate and

calyptriform or discoid,

or rarely bipartite or

shallowly 3-lobulate

deeply 2(rarely 3 to

5)-lobed

entire

stigmatie limbs:

orientation

spreading to

ascending rarely

recurved distally

appressed

spreading and

divergent

spreading

sometimes

becoming

decurved or

recurved with

age

Materials and methods

The present study involved examination of

herbarium specimens by the authors together

with field investigations by the second author

from 1988 to 1992. Altogether, approximately 80

specimens have been examined and annotated,

comprising collections from the following

herbaria: BRI, CANB, DNA, K, MEL. Acronyms

used in the text to indicate herbaria holding

particular specimens follow Holmgren et al.

(1990). All specimens cited have been seen

unless indicated as n.v. (non visus).

Descriptions of taxa were made from dried

herbarium specimens, material preserved in 70%

ethanol and glycerol, or dried material

reconstituted by placing in boiling water for a

few minutes. Colour of fresh vegetative and

floral parts are either from herbarium label notes

or from photographs taken by the second author

during field studies. Plant size, habit, flowering

and fruiting times and habitat data were

obtained from herbarium labels. The distribution

maps were produced with Maplnfo Version 3

and were based on herbarium specimen locality

data.

218

Austrobaileya 7 (1): 215-228 (2005)

Taxonomy

Shonia R.J.F.Hend. & Halford, gen. nov.

affmis Beyeriae Miq. QtRicinocarpi Desf.

Ab ilia floribus in racemis apicalibus vel

apparenter axillaribus dispositis floribus

plerumque ambobus maribus femineisque

in quoque racemo (non floribus solitariis

vel fascicularibus vel cymosis in foliorum

axillis), petalis bene evolutis plerumque

quam sepala multo longioribus raro

aequantibus (non petalis carentibus vel

petalis sepala brevioribus), stylorum ramis

stigmaticis 3 e columna brevi orientibus,

linearibus integris dorsi-ventraliter

complanatis (non stylis peltatis et inverse

cupulatis vel inverse patelliformibus

marginibus integris vel laceratis) differt.

Ab hoc stylorum ramis stigmaticis integris

dorsi-ventraliter complanatis (non

distaliter bifidis vel raro ad 5-lobatis lobis

teretibus vel dorsi-ventraliter complanatis

sed abaxialiter sulcatis), staminalibus

filamentis e basi liberis (non proximaliter

diverse connatis) differt. Typus: Shonia

tristigma (F.Muell.) Halford & R.J.F.Hend.

(= Beyeria tristigma F.Muell.)

Beyeria sect. Oxygyne F.Muell., Fragm. 6:1812

(1868), nom. inval., F.Muell. ex Benth., FI.

Australiensis 6: 68 (1873). Type: Beyeria

tristigma F.Muell. (= Shonia tristigma

(F.Muell.) Halford & RJ.F.Hend.).

Monoecious or rarely dioecious shrubs or rarely

small trees; stems erect or ascending, branching,

the branches leafy throughout. Leaves spirally

alternate, exstipulate, shortly petiolate,

persistent, with laminae glabrous and smooth

adaxially, stellate pubescent abaxially except for

the glabrous surface of the raised midvein;

secondary veins obscure; basi-laminar glands

absent. Inflorescences racemose or rarely

paniculate, terminal on stems or on axillary

branches sometimes reduced so inflorescences

Key to species of Shonia

appear axillary, with one or two female flowers

basal on the axis and with few to several male

flowers distal to them, or rarely with all flowers

male, bracteate. Flowers pedicellate with

pedicels stouter in female flowers than those of

male flowers; calyx deeply 5- or rarely 4-lobed,

with margins of lobes entire; petals present; disc

present. Male flowers with a disc of discrete

alternipetalous glands or a continuous

glandular ring; stamens 10-30, free and erect

on a flat or slightly convex receptacle; anthers

dorsifixed, extrorse, of two separate obloid,

parallel but contiguous locules each transverse

on a slightly swollen connective at the apex of

filament, dehiscing by longitudinal slits;

pistillodes absent. Female flowers with calyx

lobes persistent and appressed to ovary, with a

disc of a continuous glandular ring; ovary 3-

locular with one pendant ovule in each locule;

style short or ± obsolete; stigma with 3

horizontally spreading and divergent limbs,

sometimes becoming decurved or recurved with

age; limbs entire, ± dorsi-ventrally flattened, red,

glabrous, persistent. Fruits capsular, trilobate,

smooth, glabrous, separating septicidally into

three 2-valved cocci leaving a persistent

columella. Seeds obloid, smooth, carunculate;

caruncle waxy-fleshy; endosperm copious;

cotyledons narrower than the radicle.

The genus is endemic in Australia and

contains four species distributed in tropical

north eastern Northern Territory and central and

north eastern Queensland.

Etymology : We have named this genus Shonia

in memory of the second author’s late wife, Enid

Lynette ‘Shoni’ Henderson, accidentally killed

when their car crashed on a holiday/fieldtrip in

the Northern Territory in 1991 looking for

‘Stenolobeae’ taxa. The second author wishes

to acknowledge with gratitude her support for

him and his botanical work over more than 30

years. The name should be pronounced

“SHOWN-ee-a”.

1. Calyx lobes prominently keeled abaxially.4. S. carinata

Calyx lobes evenly rounded abaxially, not prominently keeled.2

2. Leaf laminae narrow-elliptic or narrow-oblanceolate to narrow-obovate, with

a length/width ratio less than 6:1.1. S. tristigma

Leaf laminae narrow-lanceolate or linear, with a length/width ratio of 6:1 or

greater.

Halford & Henderson, Revision of Shonia

219

3. Leaf laminae 3-10 mm wide, with a length/width ratio of 15:1 or less. 2. S. territorialis

Leaf laminae less than 3 mm wide, with a length/width ratio greater than 15:1

. 3. S. bickertonensis

Note: A specimen apparently of this genus

collected from Blackdown, WNW of Chillagoe

in north Queensland, in 1997 (Ford 1883 (BRI))

is from a shrub to 3 m tall which has leaves with

laminae linear and less than 2 mm wide. Also,

these leaves are obtuse and ultimately rounded

at the apex. As the material is incomplete, lacking

mature flowers and fruit, it is not possible to

place it in any of the above species

unequivocally. Further investigation may find

it to represent yet another species of Shonia or

perhaps another subspecies of S. tristigma.

1. Shonia tristigma (F.Muell.) Halford &

R. J.F.Hend., comb, nov.; Beyeria tristigma

F.Muell., Fragm. 6:181 (1868). Type:

Queensland. North Kennedy District:

Hinchinbrook Island, 8 November 1867,

[J. Dallachy s.n .] (lecto [here chosen]:

MEL 114230; isolecto:K).

Monoecious, erect to spreading shrubs or rarely

small trees to 4 m high, thinly viscid on most

parts; bark shallowly fissured, reddish brown.

Branchlets somewhat angular, becoming ± terete

with age, glabrous. Leaves with petioles 1-2

mm long, flattened adaxially, rounded abaxially,

glabrous and smooth; laminae narrow-elliptic,

narrow-oblanceolate to narrow-obovate, 10-60

mm long, 2-15 mm wide, with a length/width

ratio of 2-8:1, cuneate to attenuate at the base,

rounded to obtuse or acute at the apex and

recurved along the margins; adaxial surface

green, with midvein impressed; abaxial surface

white, with stellate hairs soft and up to c. 0.2

mm across, and with the raised midvein rounded

in cross section. Inflorescences racemose,

usually of one or two female flowers basal on

the axis with 5-12 male flowers distal to them;

primary axis 13^40 mm long, glabrous, smooth;

bracts triangular, c. 0.5 mm long, glabrous,

caducous. Flowers with calyx 5-lobed, of

unknown colour when fresh, glabrous; tube c.

0.1 mm long; lobes ± spreading; petals 5, of

unknown colour when fresh, glabrous except

for a tuft of hairs near the base on the adaxial

surface. Male flowers with pedicels 5-25 mm

long, angular, glabrous; calyx lobes oblong, 0.8-

1 mm long, 0.6-0.7 mm wide, obtuse to rounded

at the apex; petals broadly ovate or suborbicular,

1.4—1.6 mm long, 1.3—1.5 mm wide, rounded to

obtuse at the apex, recurved to revolute distally;

disc a ± continuous glandular ring, fleshy, lobed,

glabrous; receptacle convex, 0.8-1 mm across,

minutely hairy; stamens 20-25; filaments 0.3-

0.5 mm long, ± glabrous; anthers 0.3-0.4 mm

long. Female flowers with pedicels 10-30 mm

long, slender and circular in transverse section

proximally, becoming stouter and 5-quetrous

distally, glabrous or densely pubescent distally;

calyx lobes oblong, 1-1.2 mm long, 0.7-0.8 mm

wide, obtuse to rounded at the apex; petals

suborbicular, c. 1.2 mm long and 1.2 mm wide,

rounded at the apex, recurved distally; disc

membranous, entire; ovary subglobose,

trilobate, 1.3-1.5 mm across, ± smooth,

glabrous; style c. 0.2 mm long; stigmatic limbs

1-2 mm long. Fruits broadly ellipsoid or

subglobose, 5.5-7 mm long, 5-7 mm across.

Seeds 4.2-5.5 mm long (including caruncle), 3-

3.2 mm wide, 2.5-2.7 mm across; testa smooth,

dark brown; caruncle ± hemispherical, c. 0.5 mm

long and 1.2 mm across, ± smooth, creamy-

white.

Distribution and habitat: Shonia tristigma is

confined to the east coast of north-eastern

Queensland where it occurs in disjunct

populations from Cape York southwards to

Hinchinbrook Island, a little north of Ingham.

Typification: One collection is cited in the

protologue of Beyeria tristigma, namely “In

insula Hinchinbrook-Island prope

Rockingham’s Bay, Dallachy”. Two apparently

relevant sheets from this locality have been

located in material on loan to BRI from MEL

and K. The sheet in MEL [114230] is without

information on the collector but is dated as

collected on 8 November 1867, while the sheet

in K has information on the collector but is

without the date of collection. We considered

both to be part of the original material and hence

the MEL sheet is here selected as lectotype of

Mueller’s name.

220

Austrobaileya 7 (1): 215-228 (2005)

Notes: As circumscribed here, S. tristigma formally recognised by the establishment of

exhibits some variation in leaf shape and size as three subspecies within this species,

well as habitat differences. This variation is here

Key to the subspecies of Shonia tristigma

1. Leaf laminae narrow-elliptic, 30-60 mm long, 10-15 mm wide.

. la. S. tristigma subsp. tristigma

Leaf laminae narrow-oblanceolate to narrow-obovate, 10^40 mm long, 2-10 mm wide. 2

2. Shrubs to 0.5 m high; leaf laminae less than 20 mm long, 2-5 mm wide

. lc. S. tristigma subsp. parvifolia

Shrubs (0.2-) 0.5-4 m high; leaf laminae 20 mm or more long, mostly more

than 5 mm wide . lb. S. tristigma subsp. borealis

la. Shonia tristigma (F.Muell.) Halford &

R.J.F.Hend. subsp. tristigma

Slender shrubs to 2.5 m high. Leaf laminae

narrow-elliptic, 30-60 mm long, 10-15 mm wide.

Additional specimens examined: Queensland. North

Kennedy District: Zoe Bay, Hinchinbrook Island, Aug

1951, Blake 18859 (BRI); Mt Bowen, Hinchinbrook

Island, Jun 1991, Camming 11222 (BRI); Mt

Diamantina, Hinchinbrook Island, Jul 1991, Cumming

11279 (BRI); Little Ramsay Bay, Hinchinbrook Island,

Aug 1975, Sharpe 1593 (BRI); loc. cit. , Aug 1975,

Sharpe 1666 (BRI); northern end of Little Ramsay

Bay, Aug 1975, Sharpe 1682 (BRI); Zoe Bay,

Hinchinbrook Island, Oct 1982, Tracey 15491 (BRI,

CANB).

Distribution and habitat: Shonia tristigma

subsp. tristigma is endemic to Hinchinbrook

Island. It is recorded as growing on steep south

or south-east facing rocky slopes in heathland

or Banksia plagiocarpa and Allocasuarina

littoralis closed shrubland communities, on

steep north facing rocky slopes in rainforest

and on plains of alluvial soils in Corymbia

tessellaris woodland. Map 1.

Phenology: Flowers have been collected in July,

August and October, fruits in July and August.

lb. Shonia tristigma subsp. borealis Halford

& RJ.F.Hend., subsp. nov. folii lamina

angusta-oblanceolata usque angusta-

obovata, 20-40 mm longa et (3-) 5-10 mm

lata ab subspeciebus aliis S. tristigmatis

differt. Typus: Queensland. Cook District:

Tozers Gap, 30 June 1994, P.7. Forster

PIF15396 (holo: BRI; iso: K,MEL,NSW,

QRS distribuendi ).

Erect to spreading shrubs (0.2—) 0.5-4 m high

or rarely a small tree. Leaf laminae narrow-

oblanceolate to narrow-obovate, 20-40 mm long

and (3-) 5-10 mm wide. Fig. 1.

Selected specimens (from 36 examined):

Queensland. Cook District: slopes of Mt Fraser, Apr

1932, Brass 2430 (BRI); Newcastle Bay, 2.5 miles [c.

4 km] S of Somerset, May 1948, Brass 18772 (BRI,

CANB, K); Brown’s Creek, Pascoe River, Jun 1948,

Brass 19170 (BRI, CANB, K); Tozer Range, Jun 1948,

Brass 19351 (BRI, CANB, K); beach dune S of Cape

Bedford, Jul 1980, Clarkson 3292 (BRI, CANB);

between Mclvor River and Cape Flattery, Nov 1972,

Dockrill 616 (BRI); c. 1 km upstream from Brown

Creek crossing. Iron Range road, Apr 1988, Forster

PIF4161 & Liddle (BRI); 16 km E of Maloneys

Springs, Bromley station, Jul 1991, Forster PIF8842

(BRI); Mt Tozer, Iron Range N.P., Jul 1994, Forster

PIF15431 (BRI); Silver Plains, 4 km WNW of Colmer

Point, Jun 1999, Forster P1F24645 & Booth (BRI);

Tozers Gap, Aug 1965, Gittins 1082 (BRI); Olive River,

Sep 1974, Hyland 7444 (BRI, CANB); near Cape

Bedford, Jun 1968, Pedley 2617 (BRI); near Mclvor

River, Sep 1960, Smith 11144 (BRI); Iron Range, along

road between Browns Creek and Tozers Gap, Jul 1988,

Thomas 290 (BRI); Olive River, Sep 1974, Webb &

Tracey 13801 (BRI); between Starcke station

homestead and Cape Flattery, Jul 1976, Webb & Tracey

13833 (BRI).

Distribution and habitat: Shonia tristigma

subsp. borealis occurs in coastal and

subcoastal populations from Cape York

southwards to Mt Fraser near Mossman. It is

recorded as growing on coastal or near-coastal

sand dunes, in sandy soils on rocky slopes and

rocky headlands, in heathland, shrubland,

woodland and open forest with heath

understorey, and in vine thicket communities.

Map 2.

Halford & Henderson, Revision of Shonia

221

Fig. 1. Shonia tristigma subsp. borealis. A. Flowering branchlet. x3. B. face view of male flower. xi2. C. side view

of female flower. xi2. D. lateral view of stamen, x 40. E. face view of anther. x40. F. side view of female flower.

xl2. G. face view of fruit. x6. H. ventral view of seed. x6. I. lateral view of seed. x6. J. transverse section of leaf.

xl2. A-J from Forster PIF15396 (BRI). Del. W.Smith.

222

Austrobaileya 7 (1): 215-228 (2005)

Phenology : Flowers have been collected from

February to October, fruits from May to

September.

Notes : Shonia tristigma subsp. borealis is

variable in its stature which to some degree

appears to be related to habitat. Plants on

exposed coastal sites or in heathland

communities tend to be low shrubs 0.2-1 m high

whereas plants in more sheltered sites such as

woodland or forest communities tend to be

larger shrubs or small trees 1—4 m high.

The collection Brass 2430 from the slopes

of Mt Fraser (BRI) differs from all other

collections of this subspecies by having an

indumentum of stellate hairs on the branchlets

as compared with glabrous branchlets in other

collections seen. This collection is also markedly

disjunct from the other collections cited for this

subspecies.

Etymology : The subspecific epithet is from

Latin borealis , north or northern, and refers to

the northerly distribution of this subspecies

compared with the remainder of this species.

lc. Shonia tristigma subsp. parvifolia Halford

& RJ.F.Hend., subsp. nov. folii lamina

angusta-oblanceolata, 2-5 mm lata et 10-

17 mm longa ab subspeciebus aliis S.

tristigmatis differ! Typus: Queensland.

Cook District: 7 km W of Jowalbinna, 5

July 1990, A.R. Bean 1717 (holo: BRI).

Erect shrubs to 0.5 m high. Leaf laminae narrow-

oblanceolate, 10-17 mm long, 2-5 mm wide.

Additional specimens examined : Queensland. Cook

District: 6 km S of Jowalbinna turnoff on Maytown

track, Nov 1983, Clarkson 5053 (BRI, CANB); Sandy

Creek, 6 km NE of Jowalbinna and 25 km SSE of

Laura, Jul 1998, Wannan 908 et cd. (BRI).

Distribution and habitat : Shonia tristigma

subsp. parvifolia is known only from near

Jowalbinna which is approximately 30 km SSW

of Laura, on Cape York Peninsula. This taxon is

recorded as growing in a small ravine in

Eucalyptus/Corymbia woodland around large

sandstone boulders, on sandy soil in

Corymbia!Eucalyptus woodland and on

skeletal soils on sandstone slopes with

Eucalyptusphoenicea. Map 3.

Phenology : Flowers have been collected in July

and November, fruits in July.

Etymology : The subspecific epithet is derived

from Latin parvus , small, and - folius , -leaved,

and refers to the relatively small leaves of this

subspecies compared with those in other

subspecies of this species.

2. Shonia territorialis Halford & RJ.F.Hend.

sp. nov. S. tristigmati et S. bickertonensi

ut videtur artissime affinis. Ab S.

tristigmate folii lamina (40-) 60-80 mm non

10-60 mm longa et angusta-lanceolata vel

lineari non angusta-elliptica vel angusta -

oblanceolata ad angusta-ovata, floris

maris pedicello 2-10 mm non 5-25 mm

longo et flore femineo stylo 0.2-0.6 mm

non circa 0.2 mm longo et limbis

stigmaticis 1.3-2.5 mm non 1-2 mm longis

differt. Ab S. bickertonensi folii lamina 3-

10 mm non 0.7-2.5 mm lata et ratione

longitudinis latitudinibus 15:1 vel minus

non ultra 15:1 et inflorescentiis axe

primario 20-60 mm non 10-30 mm longo

differt. Typus: Northern Territory. Magela

Creek upper catchment, 10 April 1995, ID.

Cowie 5737 & K. Brennan (holo: DNA).

Monoecious or rarely dioecious shrubs to 1 m

high, thinly viscid on most parts; bark shallowly

fissured, grey. Branchlets somewhat angular

becoming ± terete with age, glabrous. Leaves

with petioles 1-3.5 mm long, flattened or slightly

grooved adaxially, rounded abaxially, glabrous

and smooth; laminae narrow-lanceolate or linear,

(40-) 60-80 mm long, (3-) 5-10 mm wide, with a

length/width ratio of 8-15:1, cuneate to

attenuate at the base, acute at the apex and

recurved along the margins; adaxial surface

green, with midvein impressed; abaxial surface

white, with stellate hairs soft and up to 0.2 mm

across, and with the raised midvein angular in

cross section. Inflorescences racemose or rarely

paniculate, usually of one or two female flowers

basal on the axis with 7-10 male flowers distal

to them or rarely of all male flowers; primary

axis 20-60 mm long, glabrous, smooth; bracts

triangular, c. 0.5 mm long, glabrous, caducous.

Flowers with calyx 5-lobed, of unknown colour

when fresh, glabrous; tube c. 0.1 mm long; lobes

± spreading; petals 5, of unknown colour when

fresh, ± glabrous. Male flowers with pedicels

Halford & Henderson, Revision of Shonia

223

2-10 (-15) mm long, ± terete, glabrous; calyx

lobes triangular, 0.8-1.5 mm long, 0.6-0.7 mm

wide, acute at the apex; petals broadly elliptic

or suborbicular, 1.8-2 mm long, 1.4-1.6 mm wide,

rounded to obtuse at the apex; disc a ±

continuous glandular ring, fleshy, lobed,

glabrous; receptacle slightly convex, c. 0.8 mm

across, tomentose; stamens c. 20; filaments 0.3-

0.5 mm long, ± glabrous; anthers 0.3-0.4 mm

long. Female flowers with pedicels 25-50 mm

long, slender proximally becoming stouter

distally, ± terete, glabrous; calyx lobes

triangular, 0.8-1.5 mm long, 0.5-0.8 mm wide,

acute at the apex; petals broadly elliptic or

suborbicular, c. 1.2 mm long and 1 mm wide,

rounded at the apex, recurved distally; disc a ±

continuous glandular ring, fleshy, lobed,

glabrous; ovary subglobose, trilobate, 1-1.5 mm

across, ± smooth, glabrous; style 0.2-0.6 mm

long; stigmatic limbs 1.3-2.5 mm long. Fruits

broadly ellipsoid, 6-7 mm long, 5-6 mm across.

Seeds c. 6.5 mm long (including caruncle), 3.5

mm wide and c. 3 mm across; testa dark brown;

caruncle ± hemispherical, c. 0.8 mm long and 1.7

mm across, smooth, creamy-white. Fig. 2.

Additional specimens examined : Northern Territory

South Magela Gorge, 8 km SE of Mt Brockman, Jan

1991, Brennan Bre892 (DNA); 12 miles [ c. 19 km] S

of Mt Brockman, Jul 1972, Byrnes 2712 (BRI, CANB,

NSW); 27 km SE of Jabiru, Feb 1973, Craven 2418

(BRI, CANB, DNA); c. 10 miles (16 km) ESE ofNoranda

Mining Camp, Jun 1972, Schodde AE40 (CANB,

DNA).

Distribution and habitat : Shonia territorialis

is confined to Mt Brockman and the nearby

Arnhem Land Plateau, Northern Territory. It is

recorded as growing in monsoon forest

communities in sandy soils in gullies or gorges.

Map 4.

Phenology : Flowers have been collected in

January, February, April, June and July, fruits in

January.

Affinities: Shonia territorialis seems most

closely related to S. tristigma and

S. bickertonensis. It can be distinguished from

S. tristigma by having generally longer leaf

laminae (which are (40-) 60-80 mm long

compared with 10-60 mm long in S. tristigma)

that are narrow-lanceolate or linear rather than

narrow-elliptic or narrow-oblanceolate to

narrow-obovate, male flowers with generally

shorter pedicels (which are 2-10 mm long

compared with 5-25 mm long in S. tristigma)

and female flowers with longer styles (which

are 0.2-0.6 mm long compared with up to 0.2

mm long in S. tristigma). Shonia territorialis

differs from S. bickertonensis by having leaf

laminas wider (being 3-10 mm wide compared

with 0.7-2.5 mm wide in S. bickertonensis ) and

proportionally shorter (having a length/width

ratio equal to or less than 15:1 compared with

more than 15:1 in S. bickertonensis ) and

generally longer inflorescences (with the

primary axis being 20-60 mm long compared with

10-30 mm long in S. bickertonensis).

Etymology: The specific epithet is from Latin

territorium , territory, and -alis, belonging to,

and refers to the distribution of this species

being wholly within Australia’s Northern

Territory.

3. Shonia bickertonensis (Specht) Halford &

R.J.F.Hend., comb, nov.; Beyeria

bickertonensis Specht, Rec. American-

Australian Sci. Expedition Arnhem Land

3: 249, fig. 8 (1958). Type: Northern

Territory. South Bay, Bickerton Island, in

gorge in sandstone hills, 21 June 1948,

R.L. Specht 641 (holo: BRI; iso: CANB,

K,MEL).

Monoecious, multi-stemmed shrubs to 3 m high,

thinly viscid on most parts; bark smooth or

shallowly fissured, grey or brown. Branchlets

somewhat angular becoming ± terete with age,

glabrous. Leaves with petioles 1-2 mm long,

flattened adaxially, rounded abaxially, glabrous;

laminae linear, 30-50 (-70) mm long, 0.7-2.5 mm

wide, with a length/width ratio of 18-55:1,

cuneate at the base, ± acute at the apex and

with margins recurved to the midrib; adaxial

surface green, with midrib impressed; abaxial

surface white, with stellate hairs soft and up to

c. 0.2 mm across, and with the raised midvein

angular in cross section. Inflorescences

racemose, usually of one or two female flowers

basal on the axis with 5-15 male flowers distal

to them; primary axis 10-30 mm long, glabrous,

smooth; bracts triangular, c. 0.5 mm long,

glabrous, caducous. Flowers with calyx 5-lobed,

of unknown colour when fresh, glabrous; tube

c. 0.1 mm long; lobes ± spreading; petals 5, of

unknown colour when fresh, glabrous. Male

flowers with pedicels 1.5-10 mm long, terete,

224

Austrobaileya 7 (1): 215-228 (2005)

Fig. 2. Shonia territorialis. A flowering branchlet. x2. B. male flower viewed from above. xl2. C. male flower,

oblique view. xl2. D. female flowers, lateral view. xl2. E. transverse section of leaf. xl6. A from Craven 2418

(CANB); B & C from Cowie 5737 & Brennan (DNA); D & E from Brennan Bre892 (DNA). Del. W.Smith.

Halford & Henderson, Revision of Shonia

225

glabrous; calyx lobes oblong, 1.5-1.8 mm long,

0.4-0.6 mm wide, rounded to obtuse at the apex;

petals broadly ovate, 2.3-2.5 mm long, 1.2-1.4

mm wide, acute to obtuse at the apex; disc a ±

continuous glandular ring, fleshy, lobed,

glabrous; receptacle convex, c. 0.8 mm across,

densely hairy; stamens 20-30; filaments 0.4-

0.8 mm long, ± glabrous; anthers 0.3-0.4 mm

long. Female flowers with pedicels 15^10 mm

long, ± terete, slender proximally becoming

stouter distally, glabrous; calyx lobes ovate, 0.5-

1 mm long, 0.4-0.6 mm wide, acute at the apex;

petals ovate or elliptic, 1.7-1.8 mm long, 1-1.2

mm wide, rounded at the apex; disc a ±

continuous glandular ring, fleshy, lobed,

glabrous; ovary subglobose, trilobate, c. 1.2 mm

across, ± smooth, glabrous; style 0.1-0.5 mm

long; stigmatic limbs 1.7-3 mm long. Fruits

broadly ellipsoid, 5.5-7 mm long, 4.5-5 mm

across. Seeds 4.8-5.2 mm long (including

caruncle), 2.4-3 mm wide, 2-2.6 mm across; testa

dark brown; camncle ± hemispherical, 0.8-1 mm

long, 1.2-2 mm across, ± smooth, creamy-white.

Additional specimens examined : Northern Territory.

28 km NW of Oenpelli, Feb 1973, Craven 2260 (CANB,

DNA); Mount Permain, Arnhem Land, Oct 1987,

Dunlop 7034 (BRI, MEL); Bickerton Island, Apr 1993,

Dunlop 9323 & Leach (BRI); Bickerton Island, North

Bay, May 1993, Egan 2368 (DNA); Umbakumba Road,

Groote Eylandt, 6 May 1972, Levitt s.n. (CANB, DNA);

Umbakumba Road, Groote Eylandt, 9 May 1972, Levitt

s.n. (DNA); Douglas Springs, Eva Valley Station, Jun

1988, Lucas 73 (BRI); Kakadu N.P., Douglas Springs,

Jul 1991, Menkhorst 1038 (DNA, MEL); Mt

Borradaile, Amhemland, Jul 1995, Metcalfe s.n. (DNA);

Ruined City, 50 km NNE of Ngukurr, Nov 1987, Russell-

Smith 4194 & Lucas (DNA); Upper East Alligator

River, Arnhem Land, Apr 1988, Russell-Smith 5229 &

Lucas (DNA); Eva Valley Station, 5 km E of Ranford

Hill, Jun 1988, Russell-Smith 5656 & Lucas (DNA);

4.5 km SE of Mt Borradaile, Jun 1992, Taylor 89

(DNA).

Distribution and habitat : Shonia

bickertonensis is confined to Arnhem Land and

eastern off-shore islands of the Northern

Territory. It is recorded as growing in sandy

soils on sandstone hills and in gorges. Map 5.

Phenology : Flowers have been collected in

February, April to July and September to

November, fruits in April, May, July and

October.

4. Shonia carinata Halford & R. J.F.Hend. sp.

nov. affinis S. bickertonensi Specht foliis

linearis sed calycis lobis manifeste

carinatis ab alter et speciebus omnibus

aliis Shoniae differt. Typus: Queensland.

Maranoa District: Summit of Junction

Ridge, N of Marlong Arch, Mt Moffatt

NP, 27 October 1998, A.R. Bean 14305

(holo: BRI; iso: CANB, MEL, NSW

distribuendi ),

Beyeria sp. (Bull Creek Gorge B.O’Keeffe

573), in Forster & Halford (2002:69).

Monoecious, erect to spreading multi-stemmed

shrubs to 2 m high, viscid on most parts; bark

rough and scaly, ± dark grey. Branchlets

somewhat angular, becoming ± terete with age,

glabrous. Leaves with petioles 0.5-1.2 mm long,

± terete, glabrous, smooth; laminae linear, 12-

40 mm long, 0.8-1.4 mm wide, with a length/

width ratio of 20-40:1, cuneate at the base,

obtuse to rounded or acute at the apex and with

the margins recurved to midrib; adaxial surface

yellow-green, with midvein ± obscure; abaxial

surface white, with stellate hairs soft and up to

c. 0.1 mm across, and with the raised midrib

angular in cross section. Inflorescences

racemose, usually of one or two female flowers

basal on the axis with 3-5 male flowers distal to

them; primary axis 3-10 mm long, glabrous;

bracts ovate, up to 0.4 mm long, glabrous,

caducous. Flowers with calyx 5 (rarely 4)-lobed,

green, glabrous; tube c. 0.1 mm long; lobes ±

erect; petals 5 or rarely 4, of unknown colour

when fresh, glabrous except for a tuft of hairs

near the base on adaxial surface. Male flowers

with pedicels 6-9 mm long, ± terete, glabrous;

calyx lobes oblong, 0.8-1.1 mm long, 0.5-0.8

mm wide, obtuse to rounded at the apex,

prominently keeled abaxially; petals broadly

obovate, 1.4-1.6 mm long, 0.6-1.2 mm wide,

rounded at the apex, recurved to revolute

distally; disc of 5 discrete glands; glands dorsi-

ventrally compressed, truncate to obtuse,

glabrous; receptacle ± flat or slightly convex,

0.8-1 mm across, minutely hairy; stamens c. 10;

filaments 0.7-0.8 mm long, glabrous; anthers

0.3-0.4 mm long. Female flowers with pedicels

12-17 mm long, slender and circular in

transverse section proximally, becoming stouter

and 5-quetrous distally, glabrous; calyx lobes

226

Austrobaileya 7 (1): 215-228 (2005)

Fig. 3. Shonia carinata. A.flowering branchlet. x3. B. side view of male flower. *12. C. longitudinal section of

male flower. xl5. D. lateral view of stamen. x30. E. face view of anther. x30. F. side view of female flower. *12.

G. calyx lobe. xl5. H. face view of fruit. x6. I. ventral view of seed. x9. J. lateral view of seed. x9. K. transverse

section of leaf. x30. A-H & K from Henderson & Turpin H3562 (BRI); I & J from Bean 15369 & McDonald

(BRI). Del. W.Smith.

Halford & Henderson, Revision of Shonia

227

oblong, c. 1.2 mm long, 0.4-0.5 mm wide, obtuse

to rounded at the apex, prominently keeled

abaxially; petals suborbicular, c. 1.2 mm long

and 1.2 mm wide, rounded at the apex, tumid

abaxially, recurved distally; disc membranous,

entire; ovary ellipsoid, trilobate, 1-1.3 mm

across, smooth, glabrous; style c. 0.1 mm long;

stigmatic limbs 0.3-0.4 mm long. Fruits broadly

ellipsoid, 5-5.2 mm long, 5.2-5.4 mm across.

Seeds c. 5 mm long (including caruncle), c. 2.5

mm wide and 2 mm across; testa dark brown;

caruncle ± hemispherical, c. 1 mm long and 1.2

mm across, ± smooth, creamy-white. Fig. 3.

Additional specimens examined : Queensland.

Leichhardt District: Salvator Rosa N.R, S side of Cories

Bluff, Sep 1990, Ballingall 2682 (BRI); Bull Creek

Gorge S of Springsure - Tambo Road, Sep 1999, Bean

15365 & McDonald (BRI); loc. cit., Sep 1999, Bean

15369 & McDonald (BRI); Bull Creek Gorge, c. 60 km

NE of Tambo, Aug 1991, Henderson & Turpin H3558

(BRI); loc. cit., Aug 1991, Henderson & Turpin H3562

(BRI); Bull Creek Gorge area, Sep 1984, O’Keeffe 573

(BRI); loc. cit., Sep 1984, O’Keeffe 660 (BRI); near Mt

Salvator, Sep 1992, O’Keeffe 991 (BRI); Salvator Rosa

N.R, 170 km SW of Springsure, Sep 1987, Thomas

255 (BRI). Maranoa District: Mt Mobil Holding, 15-

20 km W of Umberilla homestead, Nov 1990,

Grimshaw CHR18 (BRI).

Distribution and habitat : Shonia carinata is

confined to central Queensland in an area more

or less bounded by Tambo, Springsure and

Mitchell. It is recorded as growing in shallow

sandy soils associated with rocky sandstone

slopes and cliff lines in woodlands dominated

by Eucalyptus cloeziana F.Muell. and/or

E. decorticans (F.M.Bailey) Maiden and with a

shrubby understorey. Map 6.

Phenology : Flowers have been collected from

August to November, fruits in August and

September.

Affinities : Shonia carinata is similar to

S. bickertonensis in having linear leaves but

can be distinguished from that and all other

species of Shonia by its abaxially prominently

keeled calyx lobes.

Etymology : The specific epithet is from the

Latin carinatus , keeled, and refers to the

prominent keeled calyx lobes in the flowers of

this species.

Acknowledgments

We would like to thank Gordon Guymer for

making space and facilities available at BRI for

the first author; the directors and curators of

CANB, DNA, K and MEL for the loan of their

relevant holdings for study at BRI. The following

persons provided assistance in this study and

they are thanked sincerely for their efforts; Will

Smith (BRI) for the excellent illustrations, Gerry

Turpin (BRI) for assistance with fieldwork

undertaken by the second author and Peter

Bostock (BRI) for preparing the maps and

allowing use of his Botanical Latin (‘Translat’)

computer program to help construct the Latin

description and diagnoses. Associated

fieldwork between 1988 and 1992 by the second

author and salary support in 1999 and 2000 for

the first author were funded by grants from the

Australian Biological Resources Study (ABRS)

which are gratefully acknowledged.

References

Bentham, G. (1873). Beyeria. In Flora Australiensis 6:

63-68. L. Reeve & Co.: London.

Forster, RI. & Halford, D.A. (2002). Euphorbiaceae.

In R.J.F. Henderson (ed.), Names and

Distribution of Queensland Plants, Algae and

Lichens, pp. 68-74. Environmental Protection

Agency: Brisbane.

Halford, D.A. & Henderson, R.J.F. (2002). Studies in

Euphorbiaceae A.L.Juss. sens. lat. 3. A revision

of Bertya Planch. (Ricinocarpeae Mtill.Arg.,

Bertyinae Mtill.Arg.). Austrobaileya 6(2): 187—

245.

Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (1990).

Index Herbariorum. Parti. The Herbaria of

the World , 8 th edn. New York Botanic Gardens:

New York.

Gruning, G. (1913). IV. 147 Euphorbiaceae -

Porantheroideae et Ricinocarpoideae. In A.

Engler (ed.). Das Pflanzenreich, Regni

vegetabilis conspectus H.58: 1-97. (1968

facsimile-H.R. Engelmann (J. Cramer):

Weinheim/Bergstrasse).

Mueller, F. (1868). Fragmenta Phytographiae

Australiae 6: 181/182. Victorian Government:

Melbourne.

Webster, G.L. (1994). Synopsis of the genera and

suprageneric taxa of Euphorbiaceae. Annals

of the Missouri Botanical Garden 81: 33-144.

Wurdack, K.J., Hoffmann, P. & Chase, M.W. (2005).

Molecular phylogenetic analysis of uniovulate

Euphorbiaceae (Euphorbiaceae sensu stricto )

using plastid RBCL and TRNL-F DNA

sequences. American Journal of Botany 92:

1397-1420.

228

Austrobaileya 7 (1): 215-228 (2005)

Maps 1-6. Distribution of Shonia taxa.

Austrobaileya 7 (1): 229-230 (2005)

229

Short Communication

Cycas scratchleyana F.Muell. (Cycadaceae),

a new species record for Queensland and Australia

Paul I. Forster

P I. Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens,

Mt Coot-tha Road, Toowong, Queensland 4066. Email: paul.forster@epa.qld.gov.au

The genus Cycas has a wide distribution in

Australia, the Western Pacific, Malesia, Asia

and Madagascar. Thirty-eight species have

been recently recognised as occurring in

Australia (including its offshore islands and

territories) and six species in adjacent Papuasia

(Indonesia - Papua Province, Papua New

Guinea, Solomon Islands) (Hill 1996,1998,2004;

Hill etal. 2004; Claussen2005). Queensland has

17 species and is a major centre of diversity for

Cycas (Forster 2004).

In 2003, Bruce Wannan and Ing Toh

collected material of a Cycas on Mer (Murray)

Island in Torres Strait. Only a single plant was

observed (B. Wannan, pers. comm. 2004). This

collection is conspecific with Cycas

scratchleyana F.Muell. known previously only

from Papua New Guinea (Hill 1994; Hill et ail.

2004); hence their collection is a new species

record for both Queensland and Australia.

Although the collection is from ± natural

vegetation (Regional Ecosystem RE 3.2.5., but

floristically closer to RE 3.2.13), the collectors

note that it was made in sandy soil in a village.

Although Mer Island is politically part of

Queensland, it is immediately adjacent to the

mainland of Papua New Guinea and has at least

one other plant species ( Alectryon

repandodentatus Radik.) for which it is the only

locality in Australia. One other species (Apluda

mutica L.) is found only there and on two other

islands in Torres Strait with regard to its

Australian occurrence.

Mer Island is quite small in area and hosts

a variably sized population of people with many

links to the mainland of Papua New Guinea.

While it is feasible that this cycad may have

been transported from the mainland of Papua

New Guinea at some time and perhaps planted

at the locality it is unlikely to have happened in

recent times due to the large size of the plant (c.

15-16 m tall). Alternatively the seed may have

been transported as a food source in the past,

as species of Cycas are utilised in this manner

by indigenous people in Australia (Beck 1992)

and Papua New Guinea (see Hill 1994 for a list

of vernacular names for this species in Papua

New Guinea). This cycad colony on Mer Island

is geographically quite close to other recorded

populations of C. scratchleyana in Papua New

Guinea, hence it is also quite possible that the

population is natural and its occurrence on the

island is without anthropogenic influence.

Cycas scratchleyana F.Muell., Viet. Nat. 2(2):

18(1885). Type: Papua New Guinea. Mt

Bedford, Jala-River, Dedouri-Country,

1883, W.Armits.n. (holo: MEL 68059; iso:

Kh.v.).

Specimens examined (all at BRI): Papua New

Guinea. CENTRAL PROVINCE: Kubuna, Nov

1938, Brass 5643\ Kuriva Sawmill, Hiritano

Highway road leading into company dry dock, 9°

10’S, 147° 00’E, Larivita & Maru LAE70598;

Mekeo district, Jul/Aug 1918, White 794. GULF

PROVINCE: Ihu, Vailalu River, Feb 1926, Brass

994 ; above Purari River, 63 km at 65° from

Baimuru, 7° 15’S, 145° 20’E, Mar 1974, Croft et

al. LAE61174 ; Mina River, upstream from limit of

ship navigability. Mar 1974, Womersley NGF46475.

MILNE BAY PROVINCE: track between Tutubea

& Lake Lavu, Fergusson Island, 9° 33’S, 150° 45’E,

Nov 1976, Croft et al. LAE68785A ; c. 2 km inland

of Medino, N Coast of Cape Vogel Peninsula, Sep

1954, Hoogland 4750. WESTERN PROVINCE: Fly

Accepted for publication 16 August 2005

230

Austrobaileya 7 (1): 229-230 (2005)

River, 528 mile Camp, May 1936, Brass 6752.

Australia, Queensland. Cook District: Mer

(Murray Island) in Torres Strait, 9° 55’S, 144° 02’E,

May 2003, Wannan 2886 & Toh.

References

BECK, W. (1992). Aboriginal preparation of Cycas

seeds in Australia. Economic Botany 46: 133—

147.

CLAUSSEN, J. (2005). Native Plants of Christmas

Island. Flora of Australia Supplementary Series

No. 22. Australian Biological Resources Study:

Canberra.

FORSTER, RI. (2004). The cycads of Queensland -

diversity and conservation. Palms & Cycads

No. 82: 4-28.

HILL, K.D. (1994). The Cycas rumphii complex

(Cycadaceae) in New Guinea and the Western

Pacific. Australian Systematic Botany 7: 543-

567.

HILL, K.D. (1996). A taxonomic revision of the genus

Cycas (Cycadaceae) in Australia. Telopea 7:

1-64.

HILL, K.D. (1998). Cycadophyta. In P. McCarthy

(ed.). Flora of Australia 48: 597-661. CSIRO

Publications: Melbourne.

HILL, K.D. (2004). Cycas Candida (Cycadaceae), a

new species from Queensland together with an

extension of range and amended description

of Cycas yorkiana. Telopea 10: 607-611.

HILL, K.D., STEVENSON, D.W. & OSBORNE, R.

(2004). The World List of Cycads. In T. Walters

& R. Osborne (eds.), Cycad Classification:

Concepts and Recommendations , pp. 219-235.

CAB International Publishing: Wallingford.

New species of Commersonia J.R.Forst. & G.Forst. (Sterculiaceae) from

eastern Australia and Vanuatu

GP. Guymer

Summary

Guymer, G.P. (2005). New species of Commersonia J.R.Forst. & G.Forst. (Sterculiaceae) from

eastern Australia and Vanuatu. Austrobaileya 7(1): 231-250. Ten new species of Commersonia

are described: Commersonia argentea, C. beeronensis, C. johnsonii, C. leiperi, C. pearnii, C.

pedleyi, C. reticulata, C. rossii and C. viscidula from eastern Australia, and C. obliqua from

Vanuatu. The new combination C. novoguinensis (Gilli) Guymer is made based on Disaster

novoguinensis Gilli. The new subgenus Commersonia subgenus Verticillata Guymer is described for

the species with deeply 3-lobed petaloid staminodes: C.fraseri, C. rossii, C. argentea, C. beeronensis

and C. viscidula. An epitype is selected for C. bartramia. All new taxa are described and most

illustrated, with notes on distribution, habitat and conservation status. An identification key to

the Queensland species of Commersonia is provided.

Key Words: Sterculiaceae, Commersonia subgenus Verticillata, Commersonia argentea,

Commersonia beeronensis , Commersonia johnsonii , Commersonia leiperi, Commersonia

novoguinensis, Commersonia obliqua, Commersonia pearnii, Commersonia pedleyi,

Commersonia reticulata, Commersonia rossii, Commersonia viscidula, new species, new

combination, taxonomy, Australia, Vanuatu, Papua New Guinea, Queensland

G.P. Guymer, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens,

Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: Gordon.Guymer@epa.qld.gov.au

Introduction

The genus Commersonia was described by J.R.

Forster and G.A. Forster in Characteres

Genernm (Forster & Forster 1775) and was

based on C. echinata , a species from Tahiti

(Society Islands). However, George Forster in a

letter to Christian Voss in 1792 indicated he was

the author of the plant descriptions “He [J.R.

Forster] had no part in the plant descriptions

[for Characteres Generum ]; I completed them

all together with Sparrman,...” (Hoare 1975:170).

Commersonia is a genus of about 30

species occurring in south east Asia and the

south west Pacific with the majority of its

species occurring in Australia. The extra-

Australian species include Commersonia

echinata J.R.Forst. & GForst., endemic to Tahiti,

C. obliqua described here from Vanuatu, C.

novoguinensis (a new combination in this

paper) from montane New Guinea and the

widespread C. bartramia (L.) Merr. that ranges

from China, throughout SE Asia, New Guinea,

New Caledonia, Solomon Islands and Australia.

The Australian species occur in Western

Accepted for publication 28 November 2005

Australia, southern Northern Territory, South

Australia, Victoria, New South Wales and

Queensland.

The majority of new species described

here have been previously recognised or newly

discovered and are required for the forthcoming

census of Queensland plants. The new species

C. pedleyi described in this paper has a single

staminode or a central and two lateral

staminodes between each pair of stamens and

its other characters are consistent with the

species being positioned in Commersonia. The

genus Rulingia RBr. was described by Brown

(1820) and distinguished from Commersonia by

having a single staminode between each pair of

stamens. The characters of the staminodes of

C. pedleyi indicate that the genus Rulingia

requires reassessment with respect to its

relationship to Commersonia.

Materials and methods

This study is based upon the examination of

herbarium material at BM, BRI, CANB, G, K, L,

MEL, NSW and P, and for some species field

collections and observations by the author. The

232

Austrobaileya 7 (1): 231-250 (2005)

herbarium acronyms follow Holmgren et al.

(1990). All specimens cited have been seen by

the author. Bibliographic data are provided for

a number of previously described species that

some of the new taxa have been confused with

in the past, viz. C. bartramia, C. fraseri, C.

leichhardtii. The latter species is fully

described and typified to enable application of

the name. One species is resurrected from

synonymy under another genus (viz. C.

novoguinensis).

Measurements are sometimes abbreviated

with x indicating length x width. Descriptions

of the flowers were prepared from material

preserved in spirit or reconstituted material.

Common abbreviations in the specimen

citations are L.A. (Logging Area), N.P. (National

Park), S.F. (State Forest). The abbreviation NCA

is for the Queensland Nature Conservation Act

1992 and its associated regulations. The

conservation status of species is based on the

criteria of the IUCN (2001).

Vegetation terminology for Queensland

follows that of the Vegetation Management Act

1999 and its associated regulations. The

abbreviation RE refers to regional ecosystem

and the regional ecosystem description

database atwww.epa.qld.gov.au/REDD.

Taxonomy

Commersonia J.R.Forst. & G.Forst., Char. Gen.

43 (1775). Type: C. echinata J.R.Forst. &

GForst.

Disaster Ann. Naturhist. Mus. Wien 83:454-

455 (1980). Type: D. novoguinensis Gilli

Distribution: About 30 species in Australia, SE

Asia, Malesia and Melanesia.

Key to Queensland species of Commersonia

1. Flowers with deeply 3-lobed petaloid staminodes; capsules with 5 chartaceous

wings 2-6 mm long (subg. Verticil lata) . 2

Flowers with filiform lateral staminodes less than half as long as central

staminode lobe or rarely only with one staminode between each pair of

the stamens; capsules without chartaceous wings or with 5 short papery

upper wings 1.5-2 mm long (C. pedleyi ) (subg. Commersonia ). 5

2. Leaves and inflorescences with mixture of red glandular hairs and stellate

hairs. SE Qld & NE NSW. 1. C. viscidula

Leaves and inflorescences with predominantly stellate hairs. 3

3. Leaves glaucous above. Central Qld. 2. C. argentea

Leaves green above . 4

4. Branchlets, leaves and inflorescences with golden brown indumentum;

petals 4.5-4.8 mm long; ovules 6 per loculus. SE Qld. 3. C. beeronensis

Branchlets, leaves and inflorescences with cream or pale brown indumentum;

petals 5-5.5 mm long; ovules 8 per loculus. SE Qld, NSW .4. C. fraseri

5. Trees or rarely a montane shrub 2 m high; leaves 9-24 x 6-13 cm,

inflorescences 4-12 cm long, 50-100-flowered. E Qld, NE NSW . . 6. C. bartramia sens. lat.

Shrubs, prostrate or erect to 1 m high; leaves 1.4-11.5 x 0.4-5 cm;

inflorescences 1-2.3 cm long, 1-10-flowered. 6

6. Flowers red; leaves glaucous, ovate to ovate-lanceolate, 3.5-7 x 1.5-5 cm.

NEQld. 9. C. reticulata

Flowers white, yellow, pink or pale green; leaves 1.4-11.5 x 0.4-3 cm. 7

7. Flowers 14—17 mm diameter. Central Qld.10. C. johnsonii

Flowers 4.5-7 mm diameter. 8

Guymer, New species of Commersonia

233

8. Leaves glaucous and velutinous above, margins markedly undulate;

peduncles 0.9-1.2 mm long. Central subcoastal Qld. 11. C. pearnii

Leaves green and pubescent above, margins not markedly undulate;

peduncles 3-8 mm long. 9

9. Flowers pink; leaves lobed. S inland Qld. 14. C. pedleyi

Flowers white or yellow; leaves serrate or crenate. 10

10. Flowers white, 6-7 mm diameter; leaf blades lanceolate, 1.5-11.5 x 0.4-1.6 cm.

SEQld. 12. C. leiperi

Flowers yellow, 12-15 mm diameter; leaf blades ovate to lanceolate,

3.5-8.5 x 1-3 cm. Central eastern Qld.13. C. leichhardtii

Commersonia subgenus Verticillata Guymer,

subg. nov. affinis subgeneri Commersoniae

sed staminodiis profunde trilobis

petaloideis, pedicellis articulatis et capsulis

5-alatis differt. Typus: Commersonia

viscidula Guymer.

Notes : The subgenus differs from subgenus

Commersonia by its deeply 3-lobed petaloid

staminodes, the articulate pedicels and 5-

winged capsules. It contains 5 species and is

restricted in distribution to eastern subtropical

Australia.

Etymology: The subgenus name Verticillata

refers to the upper half of the whorled central

staminodes being radiate while the lower half

enclose the ovary.

1. Commersonia viscidula Guymer, sp. nov.

affinis C. fraseri sed ab ea foliis ramulis

inflorescentiis calycibus viscidis differt.

Typus: Queensland. Wide Bay District: Mt

Tinbeerwah, between Cooroy and Tewantin, 26°

24’S, 152° 59’E, 4 October 1981, G.P. Guymer

1614 (holo: BRI; iso BRI, CANB, K, MEL, NSW,

PERTH).

Commersonia sip. 1 (Stanley & Ross 1986).

Commersonia sp. (Mt Tinbeerwah GP. Guymer

1786) (Guymer 2002).

Spreading shrub 1-3 m, multistemmed, bark

grey with viscid branchlets, leaves,

inflorescences and calyces (glandular hairs

usually red, 0.5-2.8 mm long), suckering from

rhizomes; branchlets pubescent (hairs stellate

0.3-0.6 mm diameter, red-glandular 1-2 mm long),

glabrescent. Leaves pale green above, glaucous

below; blades ovate or ovate-lanceolate,

occasionally lower leaves 3-lobed, 5-16 x 2.5-

8.5 (-13) cm; margins undulate, crenate to

serrate, teeth 20-34 pairs, up to 2 mm long; bases

truncate or slightly cordate or oblique; apices

acute; pubescent above (hairs mid-dense,

stellate 0.2-2.4 mm diameter, red-glandular 0.1-

0.7 mm long) and velutinous below (hairs

stellate 0.4-1.4 mm diameter, red-glandular 0.2-

0.8 mm long on midvein and lateral veins); 5-

veined at base, venation slightly impressed

above, raised below, lateral veins 7-9 pairs;

petioles 6-15 mm long, pubescent (hairs stellate

& red-glandular). Stipules linear-triangular, 3-

10x1-2 mm, pubescent (hairs stellate & red-

glandular). Inflorescences 10-30-flowered, 3.5-

8 cm long; peduncles 6-25 mm long; bracts

lanceolate, 5-7 x 0.7-1.2 mm. Flowers white,

sepals sometimes with a pale pink flush, 7-10

mm diameter; pedicels 4-9 mm long, articulate

below the middle. Calyces 3.8—4.2 mm long,

pubescent outside (hairs stellate & red-

glandular), puberulous inside (hairs simple &

red-glandular); lobes ovate-acute, 2.2-32 x 2.4-

3.6 mm. Petals 3.5-5 mm long, central lobe

spathulate or obovate, margins undulate, apex

shallowly 2-4-lobed, 2.9-3.8 mm long, 1.7-2 mm

wide at apex, 0.7-0.8 mm wide at base; lateral

lobes rounded, 0.8-1.1 x 0.9-1.2 mm. Staminal

tube 0.4-0.6 mm long; central staminodes

narrowly spathulate, 4—6 mm long, apex 0.6-0.7

mm wide, middle c. 0.2 mm wide, 0.8-1 mm wide

at base, lower half forming a cone over the

ovary, upper half radiate; lateral staminodes

linear-spathulate, 2-3 mm long, 0.12-0.16 mm

wide at base, 0.2-0.4 mm wide near apex. Ovary

ovoid, 1.1-1.3 mm diameter, glabrous or with

rudimentary bristles and stellate hairs, 5-lobed

and 5-winged, margins of sutures pink or red,

234

ovules 3 or 4 per loculus; styles coherent at

apex, 0.4-0.5 mm long; stigmas globular, 0.1-

0.12 mm diameter. Capsules dark brown 18-28

mm diam., 5-winged, wings chartaceous, 4-6 mm

long; bristles 2.5-6 mm long, with stellate and

Austrobaileya 7 (1): 231-250 (2005)

simple hairs. Seeds 1-3 per loculus, black,

obovoid, angular, 2.5-2.8 x 1.8-2 mm, smooth

with fine longitudinal striations; strophiole fawn

or pale yellow, 1-1.3 mm long. Fig. 1.

Fig. 1. Commersonia viscidula. A. flowering branchlet. x 0.8. B. open flower, x 8. C. young flower before

staminodes open, x 8. All from Guymer 1853 (BRI).

Guymer, New species of Commersonia

235

Additional specimens examined (total of 28 seen):

Queensland. Burnett District: Track to Coongarra

Rock, 11.5 km ENE of Coalstoun Lakes, 25° 34’S,

151° 59’E, Sep 2002, Forster PIF28849 (BRI);

Summit of Coongarra Rock, S.F. 1344, 25° 39’S,

151° 58’E, Mar 1994, Thomas COOl (BRI). Wide

Bay District: Woowonga Creek, foothills of

Woowonga Range, 25° 29’S, 152° 08’E, Oct 1995,

Young 2040 (BRI); Summit of Mt Walsh, Mt Walsh

N.P., 25° 34’S, 152° 03’E, Aug 1996, Forster

PIF19549 (BRI); Towards the top of Biggenden

Bluff, Oct 1930, White 732 7 (BRI); 1.5 km SSW of

Biggenden Bluff, Mt Walsh N.P., 25° 35’S, 152°

01’E, Sep 2002, Bean 19232 (BRI); Summit of Mt

Tinbeerwah, Tewantin S.F. 959, 26° 23’S, 152° 58’E,

Aug 1994, Grimshaw G902 (BRI); Mt Tinbeerwah,

Apr 1962, Everist 7166 (BRI); loc. cit., Aug 1982,

Guymer 1780 (BRI); loc. cit., Guymer 1786 (AD,

BRI, CANB, NSW); loc. cit., Jan 1970, Henderson

H533 (BRI). New South Wales. Northern

Tablelands: Severn River Gorge, W of Appletree

Flat, 29° 08’S, 150° 59’E, Dec 1990, Bean 2761

(BRI); 8.5 km along Upper Rocky River road, 29°

10’S, 152° 14’E, c. 40 km SE of Tenterfield, Dec

2003, Bean 21455 (BRI). North Coast: Macleay

River, Nov 1906, Boorman s.n. (BRI [AQ81584],

G, NSW); Gloucester Buckets, Sep 1897, Maiden

s.n. (NSW, W); Darkey Creek, 35 km SW of

Singleton, Dec 1976, Guymer 841 (BRI); loc. cit.,

Oct 1970, Fisher 262 (BRI); Nymboida N.P., 29°

40’S, 152° 31’E, W of Grafton, s.dat., s.coll. (BRI);

Old Darkey Creek, c. 35 km S of Singleton on

Windsor road, Feb 1980, Williams s.n. (BRI

[AQ780689]); Parsons Creek, between Howes

Valley & Bulga, Feb 1963, Pullen 3804 (BRI).

Distribution and habitat : Commersonia

viscidula occurs from Woowonga Range,

south-east Queensland to Bulga, central

coast of New South Wales. It grows in open

shrublands, heathlands dominated by

Leptospermum microcarpum , woodlands

and eucalypt forests. The species is found

on hills, upper slopes and along creeks on

soils varying from rock pavements, to sandy

or alluvial.

Phenology : Flowers have been collected from

August to April and fruits in August, October,

December, February to April.

Affinities : Commersonia viscidula is related to

C. fraseri but differs from that species in its

viscid leaves, branchlets, inflorescences and

calyces.

Conservation status : The species has a wide

distribution and is represented in conservation

areas across its distributional range. It is not

considered to be at risk.

Etymology : The epithet is derived from the

Latin word viscidulus , referring to the sticky

leaves, branchlets and inflorescences that

characterise this species.

2. Commersonia argentea Guymer, sp. nov.

affinis C. fraseri sed ab ea ramulis foliis

inflorescentiisque argenteis velutinis

differt.

Typus: Queensland. Burnett District: 0.3 km E

of Cadarga intersection on road to Allies

Creek, 16 November 1981, G.P. Guymer

1642 (holo: BRI; iso: BRI, K, MEL, NSW).

Commersonia sp. 2 (Stanley & Ross 1986).

Commersonia sp. (Cadarga GRGuymer 1642)

(Guymer 2002).

Shrub 1.5-4 m tall, suckering from rhizomes;

branchlets silvery velutinous (stellate hairs 0.2-

1.2 mm diameter, and occasional glandular hair

to 0.2 mm long). Leaves silvery or glaucous,

discolorous; blades ovate to ovate-acute,

(2.5-) 3.5-13 x (1-) 2-7.5 (-8) cm; margins

crenulate, teeth 34-39 pairs, up to 1 mm long;

bases cordate or truncate and oblique; apices

obtuse; white stellate-velutinous above (hairs

0.2-0.7 mm diameter) and below (hairs 0.15-0.4

mm diam.); 5-veined at the base, midvein and

lateral veins slightly impressed above, veins

raised below, lateral veins 6-7 pairs; petioles

4-8 mm long, stellate-velutinous. Stipules

triangular, 4-6 x 0.7-1 mm, early deciduous,

stellate-pubescent. Inflorescences 10-30-

flowered, 2-5 cm long; peduncles 10-20 mm

long; axes white stellate-velutinous (hairs 0.15-

0.7 mm diameter & glandular hairs to 0.4 mm

long). Flowers white or cream, 9-10 mm

diameter; pedicels 9-11 mm long, articulate 3.5-

3.8 mm from flower base. Calyx lobes 2.7-3 x

1.9-2.2 mm, white stellate-velutinous outside,

sparsely puberulous inside. Petals 4.5-5.1 mm

long, central lobe spathulate, shallowly 3 or 4-

lobed at apex, margins undulate, 3.5-3.8 mm

long, 0.9-1.0 mm wide at base, 1.2-1.3 mm wide

at apex, lateral lobes rounded, 1.1-1.3 x 0.7-0.8

mm. Staminal tube 0.5-0.6 mm long. Staminodes:

central spathulate, 4.6-5.3 mm long, 0.2-0.3 mm

wide in middle, 0.7-0.8 mm wide at base, apex

0.5-0.6 mm wide and shallowly 3-lobed, lobes

rounded; laterals linear-spathulate, 3.8-4.3 mm

long, c. 0.1 mm wide in lower half and 0.3 mm

236

Austrobaileya 7 (1): 231-250 (2005)

wide near apex. Ovary 5-lobed and 5-winged,

0.9-1 mm diameter, with rudimentary bristles and

stellate hairs; ovules 6 per loculus; styles c. 0.5

mm long, free; stigmas clavate, free, c. 0.08 mm

diameter. Capsules and seeds not seen.

Additional specimens examined : Queensland.

Mitchell District: Bull Creek Gorge, Sep 1984,

O’Keeffe s.n. (BRI [AQ418119]). Leichhardt District:

40 km N of Arcadia Valley, Mar 1990, Ambrose s.n.

(BRI [AQ470721 ]); Nogoa River Gorge, Mt

Playfair Holding, Nov 1985, Don s.n. (BRI

[AQ398750]); Injune - Rolleston road, 86 km N of

Injune, Mar 1994, Halford Q2161 (BRI); Castlevale

Station, W of Springsure, Feb 1984, Jackes s.n.

(BRI [AQ544572]); 70 km E of Tambo, road to

Salvator Rosa N.P., Nov 1989, McRobert s.n. (BRI

[AQ459483]). Burnett District: 0.3 km E of Cadarga

intersection on road to Allies Creek, Nov 1981,

Guymer 1642 (BRI, K, MEL, NSW); Kragra district,

Feb 1980, Hando 137 (BRI); Kragra, near

Chinchilla, Jan 1980, Lithgow 687 (BRI); Cadarga,

Dec 1981, Lithgow 1009 (BRI).

Distribution and habitat : Commersonia

argentea is confined to central and south east

Queensland from the Carnarvon Range to the

Chinchilla area where it is a component of

eucalypt open forest on ridges, or undulating

terrain or along watercourses and occurs in

sandy soils. It is often found in stands that can

cover up to an hectare due to its suckering from

rhizomes.

Phenology : Flowers have been recorded from

November to January, and in March.

Affinities : Commersonia argentea is related to

C. fraseri but differs from that species by is

silvery velutinous branchlets, leaves and

inflorescences.

Conservation status : This species has a

sporadic distribution but has numerous plants

at most locations due to its suckering habit. It

occurs near the National Park estate but has

not yet been conserved. It is not considered to

be at risk.

Etymology: The specific epithet argentea

(Latin for silver) refers to the silver coloured

leaves of this species.

3. Commersonia beeronensis Guymer, sp. nov.

affinis C. fraseri sed ab ea foliis ramulis

inflorescentiisque aureis pubescentibus

differt.

Typus: Queensland. Burnett District: Beeron

Holding, Pretty Face Mountain, 11 September

1996, P.I. Forster PIF19610 (holo: BRI; iso: BRI,

K, MEL, NSW).

Commersonia sp. (Beeron PI.Forster PIF4658)

(Guymer 2002).

Shrub 1-2.5 m tall, suckering from rhizomes;

branchlets golden pubescent (stellate hairs 0.3-

1 mm diameter), glabrescent. Leaves green, paler

below; blades entire or lower leaves sometimes

3-lobed, ovate to lanceolate 5-12 x l^l cm;

margins denticulate, teeth 24-30 pairs, up to 1

mm long; bases truncate or sometimes cordate;

apices acuminate; pubescent above (hairs mid-

dense, stellate 0.4-0.8 mm diameter, glandular

<0.1 mm long), golden brown stellate pubescent

below (stellate hairs 0.5-1.2 mm diameter); 5-

veined at the base, midvein and lateral veins

slightly impressed above, veins raised below,

lateral veins 5-8 pairs; petioles stellate-

pubescent, 4-7 mm long. Stipules triangular or

ovate, 1.5-6 x 0.7-1 mm, early deciduous,

stellate-pubescent. Inflorescences 9-24-

flowered, 2-5 cm long; peduncles 6-10 mm long,

stellate-pubescent. Flowers cream, 6-7 mm

diameter; pedicels 2.5-5 mm long, articulate 1-

1.5 mm from flower base. Calyx lobes triangular,

2-2.2 x 2-2.5 mm, stellate-pubescent outside,

sparsely puberulous inside. Petals 4.5-4.8 mm

long; central lobe obovate, entire, 3.5-3.8 mm

long, 1.2-1.5 mm wide at apex, 0.7-0.8 mm wide

at base; lateral lobes rounded, 0.8-0.9 x 0.9-1

mm. Staminal tube 0.3-0.4 mm long; central

staminodes narrowly spathulate, 5-5.3 mm long,

0.2-0.3 mm wide in middle, 1.1-1.3 mm wide at

base, apex 0.5-0.6 mm wide; lateral staminodes

linear-spathulate, 3.8-4 mm long, c. 0.1 mm wide

in lower half and 0.3 mm wide near apex. Ovary

5-lobed and 5-winged, 0.8-0.9 mm diameter,

with rudimentary bristles and stellate hairs;

ovules 6 per loculus; styles twisted, 0.5-0.6

mm long, coherent at apex; stigmas clavate,

free, c. 0.06 mm diameter. Capsules 5-winged,

18-22 mm diameter, dark brown, wings

chartaceous, 4.5-6 mm long; bristles 1.4-2.1

mm long, with stellate hairs. Seeds 1-3 per

loculus, ovoid and angular, tuberculate, 2.5-

2.9 x 1.9-2.1 mm, dark brown to black;

strophiole 0.8-1.1 mm long, fawn or tan. Fig.

2 .

Guymer, New species of Commersonia

237

Fig. 2. Commersonia beeronensis. A. flowering branchlet. x 1. B. flower. * 8. A from Halford Q1810 (BRI); B

from Forster PIF4658 (BRI).

Additional specimens examined : Queensland.

Burnett District: 3 km WNW of Manar Homestead,

Aug 1988, Forster PIF4658 (BRI); Beeron Holding, 5

km WSW of Toondahra Homestead, Sep 1989, Forster

PIF5760 (BRI); Beeron Holding, 7 km W of Toondahra

Homestead, Aug 1990, Forster PIF7125 (BRI); Beeron

Holding, 45 km S of Mundubbera, Nov 2001, Franks

AJF011014 (BRI); Beeron Holding, ‘Rocky Paddock’,

47.8 km SSW of Gayndah, Aug 1993, Halford Q1810

(BRI).

Distribution and habitat: Commersonia

beeronensis is confined to National Park

(previously Beeron Holding), in SE Queensland.

It occurs on granite outcrops, on slopes and at

base of granite domes in open forests and

woodlands dominated by Eucalyptus dura, E.

petalophylla, E. acmenoides and Lophostemon

confertus (RE 11.12.20).

Phenology : Flowers and fruits have been

collected in August, September and November.

Affinities: Commersonia beeronensis is related

to C.fraseri but differs from that species by the

golden pubescent branchlets, leaves and

inflorescences.

Conservation status: Commersonia

beeronensis is known only from Beeron

Holding that is now a National Park. It is an

uncommon plant where it occurs in the Park

and is currently listed as Vulnerable under the

NCA. Severe wildfires kill the above ground

parts of this plant with regeneration occurring

from subterranean suckers and seed.

238

Austrobaileya 7 (1): 231-250 (2005)

Etymology : The specific epithet refers to

Beeron Holding, the only known location for

the species.

4. Commersonia fraseri J.Gay, Mem. Mus. Par.

10: 215-216, t.4 (1824); Commersonia

fraseri J.Gay var, fraseri, Mem. Mus. Par.

10:216 (1824); C. fraseri var. macrophylla

J.Gay, Mem. Mus. Par. 10: 216-218, t. 4.1

(1824). Type: “Habitat in Novae Hollandiae

NovaWalesiameridionali ... .Cl. Fraser

[553]” (holo: P; iso: K).

Commersoniafraseri var. microphylla J.Gay,

Mem.Mus.Par. 10:218,14.2(1824). Type:

“Nullum huic specimini numerum apposuit

Cl. Fraser” (holo: P).

Notes : Gay described Commersonia fraseri

based on two specimens collected by Charles

Fraser and given to Gaudichaud on his visit to

Australia on Freycinef s voyage 1817-1820. Gay

recognised two varieties of C. fraseri , var.

macrophylla and var. microphylla but not a

type variety. The detailed description of C.

fraseri var. macrophylla indicates this to be the

type variety. Gay indicated his C. fraseri var.

microphylla was possibly a proper species “an

species propria?”.

5. Commersonia rossii Guymer, sp. nov. afifmis

C. fraseri sed ab ea foliis majoribus et

trichomatibus stellatus majoribus (0.5-1.8

mm diametro) in ramulis, inflorescentiis et

floribus differ!

Typus: New South Wales. South Coast: On the

southern side of the Lochiel - South Pambula

road, c. 2 km E of Lochiel, 4 October 1985, D.E.

Albrecht2047 (holo: MEL; iso: BRI).

Commersonia sp. 1 (Short 1996).

Shrub or small tree, 4-10 m tall, suckering from

rhizomes; branchlets rusty brown stellate-

tomentose (hairs 1.2-1.8 mm diameter). Leaves

green, paler below; blades ovate to ovate-

lanceolate, 6.5-25 (-31) x 2.8-15 (-26) cm;

margins irregularly crenate, teeth 15-30 pairs,

up to 3 mm long; bases truncate or cordate;

apices acuminate; pubescent above (hairs

sparse to mid-dense, stellate 0.1-1.8 mm

diameter as well as simple), stellate-tomentose

below (hairs 0.2-1.3 mm diameter); 5- or 7-veined

at the base, midvein and lateral veins slightly

impressed above, veins raised below, lateral

veins 7-9 pairs; petioles 9-20 (-32) mm long,

stellate-tomentose. Stipules triangular, 3-10 x

1-4 mm long, early deciduous, stellate-

pubescent. Inflorescences 18-60-flowered, 3.5-

9 cm long; peduncles (3-) 10-22 mm long; bracts

ovate-lanceolate, acute, 2.8-6 x 0.9-1.7 mm; axes

rusty brown stellate-tomentose (hairs 0.5-0.8

mm diameter). Flowers white or cream, sepals

occasionally pale pink, 9-11 mm diameter;

pedicels 2.5-5.5 mm long, articulate 1.2-3 mm

from flower base. Calyces stellate-pubescent

outside, puberulous inside, 4.5—4.8 mm long;

lobes ovate-acute, 3.5—4.5 x 2.4-2.8 mm wide.

Petals obovate, entire or 2-lobed at apex, 4.3-

4.5 mm long; central lobe spathulate, 2.8-3.1

mm long, 0.4-0.6 mm wide in middle, 1.6-1.7 mm

wide at apex; lateral lobes rounded, 1-1.2 x 0.9-

1 mm long. Staminal tube 0.3-0.4 mm long; central

staminodes spathulate, margins undulate, apex

2 or 3-lobed (lobes to c. 0.5 mm long), 5-6 mm

long, 0.3-0.35 mm wide in middle, 0.7-0.9 mm

wide at base, apex 0.7-1.3 mm wide; laterals

linear-spathulate, entire or 2-lobed (lobes to c.

0.5 mm long), 3-4 mm long, 0.1-0.15 mm wide in

lower half, 0.4—0.7 mm wide near apex. Ovary 5-

lobed and winged, 1-1.3 mm diameter, with

rudimentary bristles and stellate hairs; ovules 8

per loculus; styles 0.5-0.6 mm long, coherent at

the top; stigmas clavate, free, c. 0.8 mm diameter.

Capsules 5-winged, 15-25 mm diameter.

Additional specimens examined : New

South Wales. South Coast: Near Egans Peak Nature

Reserve, Sep 1984, Albrecht 912 (BRI, MEL); Lochiel

- South Pambula road, c. 2 km E of Lochiel, Oct 1985,

Albrecht2047{ BRI, MEL); 8.2 km W along Broadwater

road from Princess Highway, Oct 1991, Albrecht 4769

(BRI, MEL); Mt Dromedary road, SW of Narooma,

Oct 1963, Adams 687 (BRI, CANB, E, L); 6.4 km E of

Bega, Oct 1984, Stewart 154 & Whigham (BRI,

CANB); Deep Creek, Eden-Bombala road, Jun 1959,

Webb & Tracey 3642 (BRI). Victoria. Gippsland:

Combienbar road, 2 miles [3.3 km] NE of junction of

Errinundraroad, Sep 1970, Beauglehole ACB34205

(BRI, MEL).

Distribution and habitat : Commersonia rossii

is confined to the South Coast, New South

Wales to East Gippsland, Victoria. It occurs in

wet sclerophyll forest, open forest, riparian

forests dominated by Tristaniopsis laurina and

the margins of warm-temperate rainforest.

Phenology : Flowers have been collected in

August and September.

Guymer, New species of Commersonia

239

Affinities : Commersonia rossii is related to C.

fraseri but differs from that species by its larger

leaves (6.5-25 x 2.8-15 cm), and larger stellate

indumentum (hairs 0.5-1.8 mm diameter) on the

branchlets, leaves and inflorescences.

Etymology : Named in honour of Dr Jim Ross,

Chief Botanist, National Herbarium of Victoria.

Commersonia subgenus Commersonia

Type: Commersonia echinata J.R.Forst. &

GForst.

6. Commersonia obliqua Guymer, sp. nov. affinis

C. echinatae sed ab ea foliis velutinis

lanceolatis usque triangulari-lanceolatis,

apicibus anguste acuminatis et foliorum

basibus valde manifestis obliquis differt.

Typus: Vanuatu. Efate Island: Undine Bay, 28

April 1928, S.F. Kajewski 238 (holo: BRI; iso:

K, P).

Shrubs or trees 3-15 m tall; branchlets stellate-

pubescent (hairs 0.15-0.5 mm diameter),

glabrescent. Leaves green, glaucous below;

blades lanceolate or triangular-lanceolate,

glabrous above except for a few scattered hairs

along primary veins, white or pale grey

velutinous below (stellate hairs 0.1-0.2 mm

diameter), (3-) 5-17.7 x 1.8-6.2 cm; apices

narrowly acuminate; bases cordate and

markedly oblique; margins irregularly serrate,

teeth to 2 mm long; midvein and lateral veins

slightly sunken above, raised below; petioles

4-11.5 mm long, pubescent. Stipules narrowly

triangular, 2.5-6 x 0.5-1.2 mm, early deciduous,

stellate-pubescent. Inflorescences 2.5-5.7 cm

long, stellate-pubescent. Flowers white, 5-6

mm diameter. Sepals stellate-pubescent,

glabrous at base inside; lobes broadly-ovate,

acute, 1.3-2.0 mm long, 1.7-2.5 mm wide; petals

2.5-3.3 mm long, tube 0.9-1.1 mm long, ligule

1.7-2 mm long, 0.4-0.5 mm wide, glabrous

outside, stellate-puberulent inside. Staminal

tube 0.4-0.5 mm long; central staminodes

lanceolate, acute, 1.6-2.1 x 0.5-0.7 mm; lateral

staminodes 0.5-0.75 mm long, stellate-

puberulent. Ovary 5-lobed, c. 0.8 mm diameter;

ovules (2-) 4 per loculus; styles connate above,

free below, 0.45-0.55 mm long; stigmas capitate.

Capsules 27-32 mm diameter, bristles 7-12 mm

long, stellate-pubescent. Seeds 2 per loculus,

botuliform, 2.5-2.7 x 1.5-17 mm, dark brown to

black; strophiole pale yellow, filamentous (5-

8), 2-2.5 mm long. Fig. 3A.

Additional specimens examined : Vanuatu. Efate

Island: Undine Bay, Apr 1928, Kajewski 238 (BRI,

K, P); loc. cit., Aug 1896, Morrison s.n. (K).

Erromango Island: Umbornale, Jul 1930,

Cheeseman 5 (K); Nouankao River & vicinity, Aug

1971, Chew Wee-Lek RSNH 139 (K); between

Nouankao Camp & Ipota, c. 10 km E of Ipota, Aug

1971, Green RSNH 1312 (K, P); Camp 5, Nevel,

Jul 1966, Johnson 2 (K, P); Dillous Bay, Jul 1896,

Morrison s.n. (K); tableland, July 1896, Morrison

s.n. (K). Tanna Is: May 1968, Bernardi 13124 (K,

P, W). Aneityum Island: 3 miles [5 km] NE of

Anelgauhat, in 1955, Cheeseman s.n. (BM, P); Oct

1858, McGillivray 5 (BM, P); Anelgauhat, 5 & 13

June 1896, Morrison s.n. (K); Anumy Valley, W

side, Jun 1896, Morrison s.n. (K); S of Ithurum, Jun

1896, Morrison s.n. (K); S of Eili, Jul 1971, Raynal

RSNH 16135 (K); Oct 1858, Saunders 5 (K);

Anelgauhat Bay, Sep 1929, Wilson in Kajewski 958

(K, P).

Distribution and habitat : Commersonia

obliqua is known from Vanuatu in tropical

lowland rainforest.

Phenology : Flowers have been collected in

April to September and fruits from April to

October.

Affinities : Commersonia obliqua is allied to C.

echinata from Tahiti but differs from that

species by its lanceolate to triangular-lanceolate

velutinous leaves, narrowly acuminate leaf

apices and markedly pronounced oblique leaf

bases.

Etymology: The epithet is from Latin obliquus

referring to the markedly oblique leaf bases.

7. Commersonia bartramia (L.) Merr., Interpr.

Rumph. Herb. Amboin. 362 (1917);

Muntingia bartramia L., Amoen. Acad.

4: 124 (1759). Type: Illustration of

Restiaria alba by Rumphius, Herb.

Amboin. 3: t. 119 (1743). Epitype [here

chosen]: Indonesia. Ambon: Kolukken, 1

April 1918, Kornassi 1071 (BRI; isoepi:

K,P).

Distribution : This species is widespread

throughout Asia from China, throughout SE

Asia, New Guinea, Solomon Islands, New

Caledonia, Fiji and Australia.

240

Austrobaileya 7 (1): 231-250 (2005)

Notes : The illustration of Restiaria alba in The degree of variation shown in leaf and

Rumphius ( loc. cit .) is stylised and lacks detail. floral characters of specimens currently referred

An epitype ( Kornassi 1071 ) is selected to to Commersonia bartramia indicate that

supplement the illustration and fix the further taxa may be included in this complex,

application of the name Commersonia

bartramia.

Fig. 3. Commersonia novoguinensis. A. flowering branchlet. x 0.75. Commersonia obliqua. B. fruiting branchlet.

x 0.75. A from Craven & Schodde 1332 (BRI); B from Kajewski 958 (BRI).

Guymer, New species of Commersonia

241

8. Commersonia novoguinensis (Gilli) Guymer,

comb. nov.

Disaster novoguinensis Gilli, Ann. Naturhist.

Mus. Wien 83:454-455 (1980). Type: Papua

New Guinea. Bachufer bei Kompiam,

1650m, February 1974,4. Gilli G557 (holo:

W).

Notes : This species was described by Gilli under

a new genus and subsequently placed in the

synonymy of Commersonia bartramia by van

Steenis (1982). The species is allied to C.

bartramia but differs from that species in its

ferruginous branchlets, leaves, inflorescences,

flowers and fruits, and by its strongly reticulate

veins and leaves being glabrescent above. Fig.

3B.

Additional specimens examined : Papua New

Guinea. Western Highlands Province: 1 mile [1.6

km] S of Birip on Kompiam-Wabag Road, May

1965, Flenley ANU2815 (K, L); Near Yogos village,

lower Tale valley, Jun 1960, Hoogland & Schodde

6660 (BRI); Kopiago, 5° 22’S, 142° 33’E, Nov

1968, Vandenberg & Galore NGF42051 (BRI,

CANB, L, K); Sirunki, pathway to swamp, below

Kaiamanda corner, Aug 1962, Walker ANU594 (K,

L); Wurup area, 10 miles [16 km] SE of Mt Hagen,

Jul 1966, Wheeler ANU6219 (BRI, CANB, K, L);

Kompiam, Jul 1959, Womersley NGF11250 (BRI);

Between Kompiam & Porget, 5° 30’S, 143° 40’E,

Jul 1959, Womersley NGF11251 (BRI, K). Eastern

Highlands Province: Mt Wilhelm, July 1959, Brass

30767 (K, L); Keglsugl, 6° 00’S, 145° 00’E, Feb

1964, Millar NGF23108 (BRI, CANB, L); Pengagl

Creek, 5° 45’S, 145° 05’E, Aug 1964, Sayers &

Millar NGF19927 (BRI, CANB, L); Above Keglsugl,

5° 50’S, 145° 05’E, May 1966, Streimann, Kairo

& Maniak NGF27751 (BRI, CANB, K, L). Morobe

Province: near Haumga, Apr 1966, Craven &

Schodde 1107 (BRI, CANB, K, L); Headwaters of

Langimar River, Apr 1966, Craven & Schodde 1332

(BRI, K, L); Edie Creek road, c. 4 miles [6.7 km]

SW of Wau, Oct 1963, Hartley 12304 (BRI); Near

Wagau airstrip, Dec 1963, Sayers sub Hartley 12585

(BRI); Wagau, 6° 45’S, 146° 50’E, Nov 1963,

Womersley NGF17821 (BRI, CANB, L). Southern

Highlands Province: near Kiburu, Mendi Valley, Jul

1961, Schodde 1429 (K, L). Central Province: Between

Efogi & Launumu, 9° 09’S, 147° 40’E, Oct 1973, Croft

& Lelean LAE60557 (BRI, CANB, K, L); Mt Pigini,

SE of Efogi, 9° 10’S, 147° 35’E, Sep 1970, Stevens

LAE50445 (BRI, CANB, K, L).

Distribution and habitat : Commersonia

novoguinensis is restricted to Papua New

Guinea at altitudes of between 1200-3000 m from

Kopiago in the Western Highlands Province to

Efogi in Central Province and occurs in montane

rainforest, on rainforest margins and in eucalypt

forest.

Phenology : Flowers have been collected in

April to July and September to November, with

fruits collected in February, May, July, August

and December.

9. Commersonia reticulata Guymer, sp. nov.

affinis C. leiperi et C. pearnii sed ab iis

floribus rubris, staminodiis erectis

lateralibus et foliis ovatis usque ovato-

lanceolatis differt.

Typus: Queensland. North Kennedy District:

Princess Hills section, Fumholtz National Park,

18° 19’S, 145°25’E, 10April2005, P.I. Forster

PIF30680 & K.R. McDonald (holo: BRI; iso:

AD, BRI, CANB, DNA, K, F, MO, MEF, NE,

NSW, PERTH).

Shrubs or subshrubs 0.3-1 mtall, suckering from

rhizomes; branchlets pubescent (hairs stellate,

O. 4-1.2 mm diameter), glabrescent. Leaves

glaucous or silvery, paler below; blades broadly

ovate to ovate-lanceolate, rarely 3-lobed, (3-) 3.5-

7 x 1.5-5 cm; margins irregularly crenate, teeth 7-

20 pairs, to 2 mm long; bases cordate; apices

obtuse or rounded; tomentose above (stellate

hairs 0.5-1.25 mm diameter) and below (stellate

hairs 0.75-1.5 mm diam.); 5-veined at the base,

veins raised below, impressed above, lateral

veins 5-6 pairs; petioles 4-7 mm long. Stipules

narrowly triangular, 5-8 mm long, stellate-

pubescent. Inflorescences 2-5-flowered, 10-15

mm long; peduncles 3-3.5 mm long; axes

stellate-pubescent (hairs 0.6-1.2 mm diameter);

bracts linear-triangular, 4-6 x 0.3-0.5 mm wide,

stellate-pubescent. Flowers red or bright red,

9-11 mm diameter, pedicels 0.2-0.3 mm long.

Calyces 5-5.5 mm long, stellate-pubescent

outside, pubescent inside (hairs stellate &

glandular); lobes ovate-acute, 2.9-3.2 x 3.8^1

mm. Petals obovate, 4-4.5 mm long; central

lobes rounded, 1.4-1.5 x 2-2.4 mm; lateral lobes

rounded, 1.1-1.2x1.1-1.2 mm long. Staminal

tube 1.8-2 mm long; central staminodes ovate-

acute, 3.5-3.8 x 1-1.2 mm; lateral staminodes

filifonn, smooth or minutely papillate, erect, 0.7-

0.8 x 0.15-0.16 mm. Ovary slightly 5-lobed, 1-

1.5 mm diameter, with rudimentary bristles;

ovules 6 per loculus; styles 1-1.2 mm long;

stigmas capitate, 0.15-0.2 mm diameter.

Capsules and seeds not seen. Fig. 4.

242

Additional specimens examined: Queensland.

North Kennedy District: Cashmere [Creek],

Rockingham Bay, s.dat., Armit s.n. (MEL

2126043); Princess Hills access road, Lumholtz

N.P., Jul 1998, Bean 13723 1 (BRI, MEL); High

Range, W of Townsville, Jan 1997, Camming

15495 2 (BRI); 43 km along Gunnawarra to

Minnamoolka road from junction with Kennedy

Development road, just N of Wombinoo entrance,

18° 03’ S, 145° 12’E, Apr 2005, Forster

PIF3065 7 & McDonald 3 (BRI, DNA, NSW,

MEL); Between Minnamoolka and Gunnawarra,

Jan 1976, Hyland 8606 3 (BRI, QRS); c. 40 miles

[66.7 km] S of Mt Garnet, Oct 1967, Morain

199 3 (BRI); Princess Hills, Lumholtz N.P., May

1996, Williams 574 1 (BRI).

Distribution and habitat : Commersonia

reticulata is known from south of Mt Garnet

to Townsville in north-east Queensland. It

occurs in mixed open forest to woodland

commonly including Corymbia clarksoniana,

Eucalyptus portuensis, E. crebra and

Corymbia citriodora on red kandosols on

Tertiary surfaces (RE 9.5.5) or mixed open forest

including Eucalyptus portuensis, E. crebra,

Corymbia clarksoniana and C. citriodora on

Austrobaileya 7 (1): 231-250 (2005)

shallow soils on metamorphic hills and ranges

(RE 9.11.4).

Phenology : Flowers have been recorded for

January, April, May, July and October.

Affinities : Commersonia reticulata is related

to C. leiperi and C. pearnii but differs from

those species by its red flowers, erect lateral

staminodes and ovate to ovate-lanceolate

leaves.

Conservation status : Commersonia reticulata

is conserved in Lumholtz N.R and is locally

abundant due to suckering in its area of

occurrence (up to 5 ha: Forster PIF30680 ; up

to 4 ha: Forster PIF30657). The species is not

considered to be at risk although it remains

poorly surveyed with only three extant

populations known ( 1_3 in specimens cited

above).

Etymology : The epithet is from Latin reticula

referring to the prominently reticulate leaf

venation that characterise this species.

Fig. 4. Commersonia reticulata. A. flowering branchlet. x 1.5. B. flower, x 6. All from Forster PIF30680 (BRI).

Guymer, New species of Commersonia

243

10. Commersonia johnsonii Guymer sp. nov.

affinis C. leiperi et C. pearnii sed ab iis

floribus majoribus (14—17 mm diam.),

petalis longioribus (5.5-6.5 mm) et ovulis

10 per loculum differt.

Typus: Queensland. Leichhardt District: Zamia

Range, 3 kmNNW of Springsure, 1 March 1960,

R.W. Johnson 1398 (holo: BRI; iso: BRI, AD,

CANB, DNA, K, MEL, NSW).

Commersonia sp. (Zamia Range R.W. Johnson

1398) (Guymer 2002).

Low spreading shrub 0.3-1 m tall, suckering

from rhizomes; branchlets yellowish-brown

stellate pubescent (hairs 0.1-1.2 mm diameter).

Leaves green or slightly glaucous; blades

lanceolate or lower leaves occasionally ovate

and 3-lobed, (1.5-) 2-10 x 0.5-2 (-2.2) cm;

margins crenate, teeth 6-14 pairs, obtuse or

rounded, to 1 mm long; bases truncate; apices

obtuse; yellowish-brown stellate-pubescent

above and below (hairs 1-1.75 mm diameter); 5-

veined at the base, venation impressed above,

raised below, lateral veins 4-7 pairs; petioles

pubescent, 3-12 mm long. Stipules narrowly

triangular, 5-9 x 0.5-1 mm, stellate-pubescent.

Inflorescences 3-flowered, 12-20 mm long;

peduncles 3-4 mm long; axes stellate-

pubescent; bracts linear or narrowly triangular,

4-10 mm long. Flowers white to pale pink, 14-

17 mm diameter, pedicels 1-4 mm long. Calyces

pubescent inside with simple, glandular (0.05-

0.1 mm long) and some stellate hairs, stellate-

pubescent outside, 7-8.5 mm long; lobes

triangular, 4-5.5 x 4.5-6.5 mm. Petals 4.8-6.5 mm

long, central lobe ovate, 2-2.5 x l-l .4 mm, lateral

lobes 0.8-1.2 mm long. Staminal tube 1-1.2 mm

long; central staminodes triangular, caudate,

2.3- 2.5 x 1.1-1.3 mm; lateral staminodes filiform,

comiculate, minutely papillate, erect, protruding

slightly above anther filament, 0.5-0.8 x 0.12-

0.15 mm long. Ovary 1—1.1 mm diameter, with

rudimentary bristles; ovules 10 per loculus;

styles 0.5-0.6 mm long; stigmas clavate, c. 0.15

mm diameter. Capsules 12-18 mm diameter,

bristles crinkled, 3-4 mm long and with stellate

hairs; seeds 1 or 2 per loculus, ovoid, 2-2.5 x

1.4- 1.6 mm, tuberculate with longitudinal ridges,

fawn to black; strophiole clear or white, 0.5-0.7

mm long. Fig. 5.

Additional specimens examined: Queensland.

Leichhardt District: On road to Fred’s Gorge,

Minerva Hills N.P., Dec 1996, Johnstone 709

(BRI); Zamia Range, 3 km NNW of Springsure,

Mar 1960, Johnson 1398 (BRI); 1.7 km W of Fred’s

Gorge lookout, Minerva Hills N.P., Sep 2005,

Thomas 3015 & Halford (BRI, MEL, NSW, CANB,

DNA, PERTH); Mt Zamia, near Springsure, Jan

1961, McLaughlin 63 (BRI); loc. cit., Sep 1962,

Story 264 (BRI); loc. cit., Jun 1985, Young 1050

(BRI); Minerva Hills N.P., W of Gorge Lookout,

Mar 1999, Stephens SP7 (BRI); Mt Zamia, 4 km N

of Springsure, Apr 1985, Rodd 4565 (BRI); Mt

Zamia E.P. [now N.P.], near Springsure, Aug 1990,

Bean 2077 (BRI); loc. cit.. Mar 1988, Donnelly

s.n. (BRI [AQ437350]); near Springsure, Jul 1980,

O’Keeffe 14 (BRI); From the cliffs, Springsure, in

1871, Roth 208 (MEL).

Distribution and habitat: Commersonia

johnsonii is restricted to Zamia Range in

Minerva N.P., near Springsure, central

Queensland. It is found in Corymbia citriodora

woodland (RE 11,8.4a) or shrubland of Acacia

curvinervia (RE 11.8.7) on Minerva Hills

volcanics (plugs, domes and dykes) of alkaline

quartz, trachyte and rhyolite on hillsides in

shallow stony soils. Other species present can

include Eucalyptus exserta, Acacia curvinervia

and Alphitonia excelsa with a grass layer of

Cymbopogon and Triodia species. Plants of this

species are common along a 500 m stretch of

road, 1.7 km W of Fred’s Gorge lookout

(M.Thomas pers. comm).

Phenology: Flowers are recorded for March,

June to September, December and January.

Fruits are recorded for March, April, August,

September, December and January.

Affinities: Commersonia johnsonii is related

to C. leiperi and C. pearnii but differs from

these species by its larger flowers (14-17 mm

diameter), longer petals (5.5-6.5) and 10 ovules

per loculus.

Conservation status: The species has a

restricted distribution on Mt Zamia and its total

known populations are contained within

Minerva N.P Further survey work is required

to determine its area of occupancy and

population numbers.

Etymology: The species is named in honour of

Robert William Johnson, botanist and

previously Director of the Queensland

Herbarium. He collected the type material of this

species in 1960 and continues to make

significant botanical contributions to our

244

Austrobaileya 7 (1): 231-250 (2005)

Fig. 5. Commersonia johnsonii. A. flowering branchlet. x 2. B. flower x 4. C. fruit, x 2. All from Thomas 3015 &

Halford (BRI).

understanding of the flora and vegetation of

Queensland.

11. Commersonia pearnii Guymer, sp. nov.

affinis C. leiperi sed ab ea foliis glaucis

velutinis marginibus valde undulatis et

lobis 5-7 rotundatis obtusisve, petiolis

longioribus et pedunculis brevioribus

(0.9-12 mm longis), differt.

Typus: Queensland. Leichhardt District:

Blackdown Tableland National Park, Charlevue

Lookout, 23° 44’ S, 149° 02’E, 20 March 2005, PI.

Forster PIF30624 (holo: BRI, iso: BRI, AD,

CANB, DNA, Q K, L, MEL, MO, NE, NSW, P,

PERTH).

Shrub 0.3-1 m tall, suckering from rhizomes;

branchlets stellate-pubescent (hairs pale yellow,

0.3-1.2 mm diam.), glabrescent. Leaves

glaucous, discolorous; blades oblong-ovate,

1.4-5 x 0.8-1.6 cm; margins with 5 to 7 rounded

or obtuse lobes 1-10 mm long, markedly

undulate; bases cordate; apices rounded;

stellate-velutinous above (hairs 0.4-0.6 mm

diameter) and below (stellate hairs 0.6-1 mm

diameter); 3-5-veined at the base, midvein and

lateral veins slightly impressed above, raised

below, lateral veins 5-6 pairs; petioles

pubescent, 4-10 mm long. Stipules narrowly to

linear triangular, 3-6 x 0.2-0.5 mm.

Inflorescences 3-8-flowered, 3-5 mm long;

peduncles 0.9-1.2 mm long; bracts narrowly

triangular, pale yellow stellate-pubescent,

deciduous, 2-3.5 x 0.2-0.3 mm; axes stellate-

Guymer, New species of Commersonia

245

Fig. 6. Commersonia pearnii. A. flowering branchlet. x 1.5. B. flower, x 16. C. opening bud x 16. All from Forster

PIF30624 (BRI).

pubescent. Flowers white, cream to pale green,

4-5 mm diameter; pedicels 0.5-0.8 mm long.

Calyx lobes triangular-ovate, 1.4-1.5 x 1.4-1.5

mm. Petals 1.3-1.5 x 1.1-1.2 mm wide; central

lobes Ungulate, 1.2-1.4 x 1.1-1.5 mm; lateral

lobes rounded, 0.7-0.8 x 0.5-0.6 mm. Staminal

tube 0.5-0.6 mm long; central staminodes

triangular, glabrous inside, papillate outside, 1-

1.1 x 0.5-0.7 mm wide; lateral staminodes

filiform, corniculate, curved across central

staminode, papillate, 0.4-0.5 x 0.1-0.13 mm.

Ovary slightly 5-lobed, 0.7-0.8 mm diameter,

glabrous or with rudimentary bristles; ovules

3-5 per loculus; styles coherent, c. 0.25 mm

long; stigmas free, globular, 0.07-0.1 mm

diameter. Capsules and seeds not seen. Fig. 6.

Additional specimens examined : Queensland.

Leichhardt District: Blackdown Tableland, Sep 1981,

Pearson 384 (BRI); Charlevue Lookout, on Loop

Road, Blackdown Tableland N.P., 23° 44’S, 149° 02’E,

Dec 1996, Johnstone 701 & Davies (BRI, NSW).

Distribution and habitat: Commersonia

pearnii is known only from Blackdown

Tableland and occurs in open-forest (to

woodland) with a range of canopy species

including Eucalyptus cloeziana, E.

melanoleuca, E. sphaerocarpa, E. suffulgens ,

E. baileyana , Corymbia bunites, C.

246

Austrobaileya 7 (1): 231-250 (2005)

hendersonii, C. trachyphloia, Corymbia

leichhardtii and C. citriodora (RE 11.10.13). It

occurs on sandstone scarps and tablelands with

shallow soils formed from Cainozoic to

Proterozoic consolidated, medium to coarse¬

grained sediments. The species is locally

abundant over about one hectare at Charlevue

Lookout (P. Forster, pers. comm).

Phenology : Flowers have been collected in

March, September and October.

Affinities: Commersonia pearnii is related to

C. leiperi but differs from this species by its

glaucous velutinous leaves with strongly

undulated margins and 5-7 rounded or obtuse

lobes, longer petioles (4-10 mm long) and its

shorter peduncles (0.9-F2 mm long).

Conservation status: Commersonia pearnii has

a very restricted distribution being confined to

Charlevue Lookout, Blackdown Tableland and

the species is conserved in Blackdown

Tableland N.P Survey work is required to

determine the species area of occupancy and

population numbers.

Etymology: Named in honour of Professor

John Pearn AM, RFD, KStJ, Professor of

Paediatrics and Child Health at the University

of Queensland, doctor, educator, humanitarian

and author of medical and botanical works.

12. Commersonia leiperi Guymer, sp. nov.

affinis C. pearnii sed ab ea foliis aureo-

fuscis stellato-pubescentia marginibus

serratis paribus 6-7 dentium praeditis,

petiolis brevioribus (1-2.5 mm longis) et

pedunculis longioribus (3-8 mm longis)

differt.

Typus: Queensland. Wide Bay District: State

Forest 1294,20 km SE of Childers, 1 February

1996, K.M. Sparshott KMS744 & R. Price

(holo: BRI; iso: BRI).

Erect or prostrate shrub 0.2-1 m high, suckering

from rhizomes; bark brown, lenticellate;

branchlets golden stellate-pubescent (hairs 0.2-

F2 mm diameter). Leaves green, paler below;

blades lanceolate, 1.5-11.5 x 0.4-1.6 cm; margins

serrate, teeth 4-7 pairs, 0.5^1 mm long; bases

truncate; apices acute or obtuse; golden brown

pubescent above (hairs mid-dense, simple,

biramous & stellate 0.4-1 mm diameter), stellate-

pubescent below (hairs 0.4-1 mm diam.); 3-

veined at base, veins slightly impressed above,

laterals and midrib raised below, lateral veins 4-

5 pairs; petioles pubescent, 1-2.5 mm long.

Stipules linear, 1.4-1.7 x 0.1-0.15 mm.

Inflorescences cymose, 3-12-flowered;

peduncles 3-8 mm long; bracts linear, 1-2 mm

long; axes stellate-pubescent. Flowers white,

pink in bud, 4.5-5.5 mm diameter; pedicels 1-4

mm long, pubescent (hairs stellate and

glandular). Calyx pubescent outside (stellate

hairs 0.2-0.4 mm diameter & red glandular hairs

to 0.4 mm long), puberulous inside (hairs stellate

6 glandular); lobes ovate-acute, 1.5-1.6 x 1.3—

1.4 mm. Petals 1.7-1.9 x 1.5-1.7 mm; central lobe

ovate, 0.8-1.3 x 0.7-0.9 mm; lateral lobes

rounded, 1.6-1.8 x 1.5-1.7 mm. Staminaltube

0.8-1 mm long; central staminodes ovate-acute,

0.8-0.9 x 0.5-0.6 mm, papillate outside and on

margins, glabrous inside; lateral staminodes

filiform, comiculate, curved in front of central

staminode, 0.3-0.4 x 0.2-0.25 mm, papillate.

Ovary slightly 5-lobed, 0.75-0.85 mm diameter,

with rudimentary bristles; ovules 6 per loculus;

styles coherent, 0.4-0.5 mm long; stigmas

capitate, coherent, 0.12-0.15 mm diameter.

Capsules and seeds not seen. Fig. 7.

Additional specimens examined : Queensland. Wide

Bay District: S.F. 57, St Mary, 25° 40’S, 152° 25’E, Apr

1996, Forster PIF19113 (BRI); Joe. cit. , 25° 40’S, 152°

24’E, Aug 1996, Forster PIF19501 (BRI, MEL); cult.

Beenleigh (ex S.F. 57, St Mary), Mar 1997, Leiper s.n.

(BRI [AQ654851]); Coonarr Creek, near Elliott River,

c. 20 km SSE of Bundaberg, s.dat, s.coll. (BRI); Wongi

Forestry, Childers, Feb 1988, Randall 502 (BRI);

Distribution and habitat: Commersonia leiperi

is restricted to a single degree square from just

south of Bundaberg to State Forest 57 St Mary,

southwest of Maryborough, in south east

Queensland. It has been recorded from

woodlands and open forests of Angophora

leiocarpa. Eucalyptus cloeziana and E.

acmenoides in sandy soil (RE 12.9-10.21).

Phenology: Flowers have been collected in

February and April.

Affinities: Commersonia leiperi is related to

C. pearnii but differs from that species by its

golden brown stellate-pubescent leaves, serrate

leaf margins with 6-7 teeth, shorter petioles (1-

2.5 mm long) and its longer peduncles (3-8 mm

long).

Guymer, New species of Commersonia

247

Fig. 7. Commersonia leiperi. A. flowering branchlet. x 2. B. flower from above, x 10. C. flower from the side, x

10. All from Leiper s.n. (BRI [AQ654851]).

Conservation status'. Commersonia leiperi has

a restricted distribution of less than 100 km but

has significant populations conserved in State

Forests. The species requires detailed survey

to determine the area of occupancy and number

of individuals.

Etymology: This species is named in honour of

Glenn Leiper, educator, amateur botanist,

photographer and author. Glenn originally

discovered this species in State Forest 57 St

Mary and has cultivated and flowered it at

Beenleigh enabling illustration of the flowers.

13. Commersonia leichhardtii Benth., FI.

Austral. 1:242-243 (1863). Type: Queensland:

Head of the Boyd river, 29 January 1847,

Leichhardt 574 (lecto [here chosen]: MEL

236401).

Shrubs 0.2-0.5 m tall, suckering from rhizomes;

branchlets golden-brown stellate-pubescent

(hairs 0.4-1.2 mm diameter), glabrescent. Leaves

green or slightly silvery, discolorous; blades

ovate to lanceolate, occasionally lower leaves

shallowly 3-lobed, 3.5-8.5 x 1-3 cm; margins

crenate, teeth 7-22 pairs, to 2 mm long; bases

248

Austrobaileya 7 (1): 231-250 (2005)

truncate or slightly cordate; apices obtuse;

stellate pubescent above (hairs 0.4-1 mm

diameter) and below (hairs 0.6-1.8 mm diameter);

5-veined at the base, midvein and lateral veins

impressed above, raised below, lateral veins 5-

7 pairs; petioles 4-12 (-14) mm long, stellate-

pubescent. Stipules narrowly triangular, 3-12.8

x 0.8-1 mm, deciduous, stellate-pubescent.

Inflorescences 3-5-flowered, 1.5-2 mm long;

peduncles 5-6 mm long; axes stellate-

pubescent (hairs 0.0.6-1.4 mm diameter); bracts

narrowly triangular, 5-8 x 0.2-0.3 mm long,

stellate-pubescent. Flowers yellow, 12-15 mm

diameter; pedicels 1-1.8 mm long. Calyces 9-10

mm long, stellate-pubescent outside (hairs 0.4-

1.8 mm diameter); lobes ovate-triangular, 5-6.5

x 4-6 mm, pubescent inside (hairs stellate &

simple). Petals 4.5-5 mm long; central lobe ovate,

2.3-2.5 x 1.2-1.4 mm; lateral lobes rounded, 1-

1.2 x 1-1.8 mm. Staminal tube 0.8-1 mm long;

central staminodes triangular, caudate, 2.5-4 x

1-1.5 mm; laterals erect, corniculate, smooth,

0.5-0.9 x 0.2-0.25 mm. Ovary slightly 5-lobed,

1—1.3 mm diameter, and with rudimentary

bristles; ovules 10-12 per loculus; styles 0.7-1

mm long, free; stigmas clavate, free, 0.2-0.3 mm

diameter. Capsules and seeds not seen.

Additional specimens examined : Queensland.

Leichhardt District: Head of Boyd River [Creek],

Jan 1847, Leichhardt s.n. (K). Burnett District:

Gorge, Oaky L.A., Coominglah S.F., NW of Monto,

Mar 1996, Bean 10147 (BRI, MEL); loc. cit., Jun

1996, Bean 10414 (BRI). Darling Downs District:

Turkey Mt, Barakula S.F. 302, Nov 1987, Forster

PIF15173 (BRI); Barakula S.F. 302, Sep 1996,

Forster PIF19627 & Ryan (BRI); Turkey Mt track,

Barakula S.F., Nov 1984, R. Hando 370 (BRI);

Turkey Mt fire lookout, Barakula S.F., V. Hando

379 (BRI); loc. cit.. May 1985, V Hando 380 (BRI).

Distribution and habitat : Commersonia

leichhardtii is confined to central Queensland

from Boyd Creek, west of Taroom to Barakula

State Forest, near Chinchilla. It occurs in

woodlands or open forests of Corymbia

citriodora. Eucalyptus crebra and E.

acmenoides on sandstone ridges or stony

hillsides (RE 11.7.6).

Phenology : Flowers have been collected in

March, June and November.

Typijication: The type citation for this name

states “Head of Boyd river, Leichhardt, in Herb.

F. Muell. ”. Although an unnumbered collection

of Leichhardt’s was retained at Kew, the

numbered specimen in MEL is superior in quality

and is used to typify the name.

14. Commersonia pedleyi Guymer sp. nov.

affinis C. leiperi et C. pearnii sed ab iis

floribus roseis 6-7 mm diam., ovulis 10

per loculum, foliorum marginibus loborum

partibus 5-16 praeditis, differt.

Typus: Queensland. Darling Downs District:

13 miles [21.7 km] W ofWestmar, 19 October

1959, L. Pedley 518 (holo: BRI; iso: BRI, AD, K,

MEL, NSW).

Rulingia sp. 1 (Stanley & Ross 1986).

Rulingia sp. (Westmar L.Pedley 518) (Guymer

2002 ).

Low spreading or prostrate shrub to 60 cm tall,

20-150 cm across, suckering from rhizomes;

branchlets golden pubescent, glabrescent.

Leaves dull green, paler below; blades usually

linear-lanceolate to lanceolate, 3-lobed,

occasionally ovate and palmate, 2-7 x 0.4-4 cm;

margins lobed, lobes 5-16 pairs, 1-4 mm long;

bases truncate, cordate, occasionally oblique;

apices obtuse or rounded; sparsely pubescent

above (hairs simple 0.1-0.3, glandular and

stellate 0.3-0.5 mm diameter), stellate pubescent

below (hairs mid-dense, 0.2-0.8 mm diameter);

3-veined at base, midvein and lateral veins

impressed above, veins raised below, lateral

veins 4-9 pairs; petioles 2-5 mm long, stellate-

pubescent. Stipules narrowly triangular, 2-5 mm

long, stellate-pubescent. Inflorescences 7-10-

flowered, 10-23 mm long; peduncles 4-5 mm

long; bracts linear, 2-9 mm long; axes stellate-

pubescent. Flowers at first white, later becoming

pink, 6-7 mm diameter; pedicels 1-3 (-9) mm

long. Calyces 4-6.5 mm long, stellate pubescent

outside, pubescent inside (hairs stellate, simple

and glandular); lobes ovate-acute, 3-3.5 x 2.5-

3.2 mm. Petals 4-5.5 mm long; central lobes

linear-lanceolate, 2.7-4.5 mm long, apex

rounded, 0.3-0.45 mm wide, c. 0.5 mm wide at

base; basal lobes rounded, 0.7-1 mm long.

Staminal tube 1.2-1.4 mm long; central

staminodes narrowly triangular, caudate,

reflexed, 4.3^1.6 x 0.8-1 mm; lateral staminodes

present or absent, filiform, erect, 0.35-0.5 mm

long, papillate. Ovary 5-lobed, 1.2-1.3 mm

diameter, with rudimentary bristles and stellate

Guymer, New species of Commersonia

249

hairs; ovules 9-10 per loculus; styles free, 0.5-

0.6 mm long; stigmas 0.1-0.16 mm diameter.

Capsules 9-12 mm diameter, with 5 papery

protruding upper wings (along sutures) 1.5-2

mm long, bristles slightly crinkled, 1-2.3 mm long

with simple and stellate hairs; seeds 4 or 5 per

loculus, ovoid, 2-2.4 x 1-1.5 mm, tuberculate,

black; strophiole 1-1.8 mm long, fawn. Fig. 8.

Additional specimens examined : Queensland.

Maranoa District: 80 km E of St George, Apr 1984,

Ballingall 1370 (BRI); 44 miles [73.3 km] NE of St

George, Aug 1956, Everist 5832 (BRI); Megine to

Brucedale road, c. 40 km S of Roma, Sep 1995,

Grimshaw PG2171 (BRI); St George - Bollon road,

Sep 1983, Jahnke s.n. (BRI [AQ398475]); 56 km NNW

of Talwood, Feb 1968, McDonald 375 (BRI); 3 km E

of Yuleba, Oct 1973, s.coll. (BRI [AQ11346]). Darling

Downs District: 1.6 km S of Miles, Mar 1994, Bean

7582 (BRI); 18.2 km W of Westmar, Moonie Highway,

Oct 1984, Forster PIF1918 (BRI); Miles, Mar 1950,

Gordon DMG155 (BRI); 19.9 km W of Westmar, Sep

2005, Halford Q8584 & Thomas (BRI); c. 24 km W

of Westmar, Sep 2005, Halford Q8586 & Thomas

(BRI); Gurulmundi, near Miles, Sep 1977, Hondo s.n.

(BRI [AQ264195]); Main Range, near Gurulmundi, Nov

1930, Hubbard 5111 (BRI, K); Moonie, Dec 1970,

Pedersen CO4 (BRI); Main Range, 3 miles [5 km] W

of Gurulmundi, Sep 1961, Pedley 879 (BRI); 5 km W

of Gurulmundi, Jan 1984, Pedley 5 029 (BRI); 10 km

from Gurulmundi on Woleebee road, Aug 1977, Ross

s.n. (BRI [AQ228855]); Glenmorgan, Oct 1969, Smith

s.n. (BRI [AQ377200, 377339]); 17 km W of Westmar,

Sep 1988, Thomas s.n. (BRI [AQ477366]); 16 km W

of Gurulmundi on Woleebee road, Oct 1973, Williams

75043 (BRI).

Distribution and habitat : Commersonia

pedleyi has a distributional range of 160 km from

Gurulmundi, north of Miles in the western

Darling Downs District, to west of St George, in

the eastern Maranoa District, southern

Queensland. It occurs in heathlands,

woodlands, open forests and along roadsides

and is found on sandy soils, red lateritic earths,

sand ridges and brown loams.

Phenology: Flowers have been collected in

August to January, March and April, and fruits

in October, November and March.

Affinities: Commersonia pedleyi is allied to C.

leiperi and C. pearnii but differs from those

species by its pink flowers 6-7 mm diameter, its

10 ovules per loculus and its leaf margins with

5-16 pairs of lobes.

Conservation status: The species has a wide

range of occurrence (160 km) and is conserved

in the National Park estate. It is not considered

to be risk.

Etymology: Named in honour of Leslie Pedley,

botanist and formerly Assistant Director at the

Queensland Herbarium, who made extensive

collections of the species (including the type)

and who first recognised this species was

unique and new to science.

Fig. 8. Commersonia pedleyi. A. flowering branchlet. x 2. B. flower, x 8. All from Forster PIF1918 (BRI).

250

Austrobaileya 7(1): 231-250 (2005)

Acknowledgements

I would like to thank the directors and staff of

BM, CANB, E, Q K, MEL, NSW, P and W for

access to their collections; Will Smith for the

illustrations; Peter Bostock for the translations

of the diagnoses into Latin; and Megan

Thomas, David Halford, Glenn Leiper, Keith

McDonald and Paul Forster for their collections

of excellent specimens.

References

Brown, R. (1820). Rulingia pannosa. In J. Sims (ed.),

Curtis’s Botanical Magazine 48: t. 2191.

Forster, J.R. & Forster, J.G. (1775). Characteres

Generum Plantarum. White et al . London.

Guymer, G.R (2002). Sterculiaceae. In R.J.F. Henderson

(ed.). Names and Distribution of Queensland

Plants, Algae and Lichens. Environmental

Protection Agency: Brisbane.

Hoare, M.E. (1975). The tactless philosopher: Johann

Reinhold Forster (1729-98). Melbourne: The

Hawthorn Press.

Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (1990).

Index Herbariorum. Part 1. The Herbaria of

the World. 8 th edn. New York Botanic Garden:

New York.

Iucn (2001). IUCN Red List Categories and Criteria.

Version 3.1. IUCN-The World Conservation

Union: Gland/Cambridge.

Short, PS. (1996). Sterculiaceae. In N.G. Walsh & T.J.

Entwisle (eds.). Flora of Victoria, Dicotyledons.

3: 324-331. Inkata Press: Melbourne.

Stanley, T.D. & Ross, E.M. (1986). Flora of south¬

eastern Queensland. 2: 87. Queensland

Department of Primary Industries: Brisbane.

Van Steenis, C.G.G.J. (1982). Sterculiaceae. In C.G.G.J.

van Steenis (ed.). Critical notes on New Guinea

plants described by A. Gilli. Blumea 28: 169.

Austrobaileya 7(1): 251 (2005)

Orthographic Corrections

Austrobaileya 6(4): 2004

251

p. 616, species 18.

The correct spelling of the species name,

according to Article 23.5 of the current

International Code of Botanical Nomenclature,

is Cor chorus subargenteus since the specific

epithet, according to the author, is derived from

Latin sub-, somewhat and argentea, silvery, and

is in reference to the appearance of the foliage

of this species. The adjective argenteus (W.T.

Stearn, Botanical Latin Fourth Edition 1992,

p.371) is a Group A adjective which is declined

as the stem argente- with terminations -us

(nominative singular masculine case, as in this

situation), -a (nominative singular feminine

case) or -um (nominative singular neuter case).

p. 843, forma 6d(ii)

The correct spelling of the forma name

according to Article 24.2 of the current

International Code of Botanical Nomenclature

is Psydrax odorata forma parviflorifera since

the forma epithet, according to the authors, is

based on “Latin parvus , ‘small’, and florifer ,

‘flowerbearing’, [which] refers to the small

inflorescence in this form”. The adjective

florifer is declined with terminations -er (for

nominative singular masculine case), -era (for

nominative singular feminine, as in this

situation) or -erum (for nominative singular

neuter case) (W.T. Stearn, Botanical Latin

Fourth Edition 1992, p.91).

Austrobaileya 7 (1): 1-252 (2005)

Studies in Euphorbiaceae, s.lat. 7. Shonia RJ.F.Hend. & Halford ( Ricinocarpeae,

Ricinocarpinae), a new Australian endemic genus

D.A. Halford & R.J.F. Henderson . 215-228

Cycas scratchleyana F.Muell. (Cycadaceae), a new species record for Queensland and Australia

P.I. Forster . 229-230

New species of Commersonia J.R.Forst. & GForst. (Sterculiaceae) from Eastern Australia and Vanuatu

G.P. Guymer . 231-250

Orthographic corrections.251

Austrobaileya 7(1): 1-252 (2005)

Contents

Studies in Euphorbiaceae, s.lat. 6. A revision of the genus Poranthera Rudge

( Antidesmeae , Porantherinae ) in Australia

D.A. Halford & R.J.F. Henderson . 1-27

A taxonomic revision of Tarenna Gaertn. and Triflorensia S.T.Reynolds (Rubiaceae:

Ixoroideae: Pavetteae) in Australia

S. T. Reynolds & PI. Forster . 29-55

A taxonomic revision of Actephila Blume (Euphorbiaceae/Phyllanthaceae) in Australia

PI. Forster . 57-97

Studies in Australian Myrsinaceae: Tapeinosperma Hook.f.

B.R. Jackes . 99-110

Notes on the narrow-leaved Ironbarks (Myrtaceae: Eucalyptus subseries Subglaucae)

A.RBean . 111-120

Backhousia enata A.J.Ford, Craven & J.Holmes (Myrtaceae), a new species from

north-eastern Queensland

A.J. Ford, L.A. Craven & J.J. Brophy . 121-127

Bazzania sauropoda D.Meagher (Marchantiophyta: Lepidoziaceae), a new species from

tropical Queensland

D. Meagher . 129-133

Gyrostemon osmus Halford (Gyrostemonaceae), a new species from south-eastern Queensland

D.A. Halford . 135-139

Eucalyptus erosa A.R.Bean (Myrtaceae), a new stringybark species from central Queensland

A.RBean . 141-144

Rediscovery and neotypification of Marsdenia arachnoidea Schltr. (Apocynaceae:

Asclepiadoideae - Marsdenieae), an endangered species from Papua New Guinea

PI. Forster & W. Takeuchi . 145-150

Ludwig Leichhardt’s Australian Plant Collections, 1842-1847

J.L.Dowe . 151-163

Wing structure in seeds of Strangea Meisn. and Stenocarpus R.Br. (Proteaceae)

H. T. Clifford & M.E. Dettmann . 165-170

The rediscovery of Boronia inflexa subsp. grandiflora (Rutaceae)

M.F. Duretto, P. Grimshaw & K. Sparshott . 171-173

New species of Philotheca Rudge (Rutaceae) from Queensland

PI. Forster . 175-181

Papuasian Orchid Studies, 2

P. Ormerod . 183-203

Goodenia elaiosoma LD.Cowie (Goodeniaceae), a new species from the Top End

of the Northern Territory and a key to the northern species

I. D. Cowie . 205-213

{continued inside back cover)