Editorial Committee

P.I.Forster (editor)

P.D.Bostock (technical advisor)

G.P.Guymer (technical advisor)

Graphic Design

Will Smith

Desktop Publishing

Yvonne Smith

Austrobaileya

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

Vol. 7, No. 3 was published on 30 November 2007

Vol. 7, No. 4 was published on 18 December 2008

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, Environmental Protection Agency.

Austrobaileya 7 (4): 577-737

Contents

A taxonomic revision of Beyeria Miq. (Euphorbiaceae: Ricinocarpeae ,

Ricinocarpinae )

D.A.Halford & R.J.F.Henderson . 577-639

A taxonomic revision of Erythrina L. (Fabaceae: Faboideae ) in Australia

A. R. Be an . 641-658

A remarkable new species of Rimularia Nyl. (lichenized fungi: Trapeliaceae)

from tropical Australia

G.Kantvilas, P.McCarthy & B. Stuckey . 659-663

A new subsection and two new subseries for Boronia Sm. Section Valvatae

(Benth.) Engl. (Rutaceae)

M.F.Duretto . 665-668

Solanum succosum A.R.Bean & Albr. (Solanaceae), a new species allied to

S. chippendalei Symon

A.R.Bean & D.E. Albrecht . 669-675

F.M.Bailey’s taxonomy of A.Meston’s collections from the Bellenden Ker

Expedition of 1904

J.L.Dowe & A.D.Broughton . 677-679

New subspecies for Zieria odorifera J.A.Armstr. (Rutaceae) from northern

New South Wales

M. F.Duretto & P.I.Forster . 681-690

Three new species of Rhodomyrtus (DC.) Rchb. (Myrtaceae) from Papua

New Guinea

N. Snow, JMcFadden & J.P.Atwood . 691-706

Plectranthus batianoffii P.I.Forst. (Famiaceae), a new species from north¬

east Queensland

P. I. Forster . 707-710

Huperzia tetrastichoides A.R.Field & Bostock (Fycopodiaceae) a newly

recognised species of tassel fern from the Wet Tropics of Queensland,

Australia

Ashley R. Field & Peter D.Bostock .711-715

Corsia dispar D.E. Jones &B.Gray (Corsiaceae), a new species from Australia,

and a new combination in Corsia for a New Guinea taxon

D.L. Jones & B.Gray .717-722

Capparis batianoffii Guymer (Capparaceae), a new species from central

coastal Queensland

G.P Guymer . 723-725

Reinstatement of Ammannia triflora Benth. (Fythraceae)

A.R.Bean . 727-728

A new combination in Cissocarpus Rozefelds (Vitaceae), a fossil genus from

Queensland

P.D.Bostock & H.T.Clifford .729

(continued )

Austrobaileya 7 ( 4 ): 577-737

Notes on Palmeria F.Muell. (Monimiaceae) in Australia and the application

of the name Palmeria racemosa (Tul.) A.DC.

G.P.Guymer . 731-733

Two new species of Pandorea Spach (Bignoniaceae) recognised from

Queensland

G.P.Guymer . 735-736

New combinations in Liparis Rich, and Pterostylis R.Br. for two species of

Orchidaceae from Queensland

Peter D.Bostock .737

A taxonomic revision of Beyeria Miq.

(Euphorbiaceae: Ricinocarpeae , Ricinocarpinae )

David A. Halford and Rodney J.F. Henderson

Summary

Halford, D A. & Henderson, R.J.F. (2008). A taxonomic revision of Beyeria Miq. (Euphorbiaceae:

Ricinocarpeae, Ricinocarpinae). Austrobaileya 7(4): 577-639. The genus Beyeria Miq. is endemic

to Australia. Twenty-four species are recognised and a key provided for their identification. The

following species are described here as new: Beyeria apiculata Halford & R. J.F.Hend., B. cockertonii

Halford & R.J.F.Hend., B. constellata Halford & R.J.F.Hend., B. disciformis Halford & R.J.F.Hend.,

B. lanceolata Halford & R.J.F.Hend., B. lapidicola Halford & R.J.F.Hend., B. physaphylla Halford

& R.J.F.Hend., B. rostellata Halford & R.J.F.Hend., B. simplex Halford & R.J.F.Hend., B. sulcata

Halford & R.J.F.Hend. and B. villosa Halford & R.J.F.Hend. A new subspecies and new variety are

described, namely Beyeria cinerea subsp. borealis Halford & R.J.F.Hend. and B. sulcata var. gracilis

Halford & R.J.F.Hend. The new combinations Beyeria sulcata var. brevipes (Airy Shaw) Halford &

R.J.F.Hend., based on B. brevifolia var. brevipes Airy Shaw and B. sulcata var. truncata (Airy Shaw)

Halford & R.J.F.Hend., based on B. brevifolia var. truncata Airy Shaw, are made. Beyeria calycina

var. minor Airy Shaw is raised to specific rank as B. minor (Airy Shaw) Halford & R.J.F.Hend. The

new taxa are illustrated and distinguished from related taxa while all taxa are described and mapped

with notes provided on their habitat, distribution and phenology. Lectotypes are chosen for Beyeria

drummondii Mull.Arg., Beyeria latifolia Baill., Beyeria lechenaultii forma elaeagnoides Baill.,

Beyeria lechenaultii forma pernettioides Baill., Beyeria lechenaultii forma rosmarinoides Baill.,

Beyeria lechenaultii var. latifolia Griming, Beyeria lepidopetala F.Muell., Beyeria opaca F.Muell.,

Beyeria opaca var. linearis Benth., Beyeria viscosa var. amoena Mull.Arg., Beyeria viscosa var.

angustifolia F.Muell. & Tate, Beyeriopsis latifolia Mull.Arg. and Croton viscosus Labill. All known

synonyms are listed here.

Key Words: Euphorbiaceae, Beyeria, Beyeria apiculata, Beyeria cinerea subsp. borealis, Beyeria

cockertonii, Beyeria constellata, Beyeria disciformis, Beyeria lanceolata, Beyeria lapidicola,

Beyeria minor, Beyeria physaphylla, Beyeria rostellata, Beyeria simplex, Beyeria sulcata, Beyeria

sulcata var. brevipes, Beyeria sulcata var. gracilis, Beyeria sulcata var. truncata, Beyeria villosa,

Australia, Australian flora, taxonomy, nomenclature, identification keys

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

Beyeria Miq. is endemic to Australia with 24

species recognised in the present revision.

Species of Beyeria are woody, perennial

shrubs or small trees and the majority grow

in shrubland, heathland or dry sclerophyll

open forest or woodland communities,

in a variety of soils derived from diverse

geologies. The species are largely temperate

in their distribution (Map 1) with the majority

endemic to south-western Australia.

The genus was established in 1844 by

Friedrich Miquel who named a single species,

Beyeria viscosa Miq., based on material

collected by Ludwig Preiss in 1839 at Rottnest

Accepted for publication 1 July 2008

Island, Western Australia (Miquel 1844). The

name commemorates Adriaan de Beyer [de

Beijer] (1773-1843) who Miquel described

as a dedicated worker for his studies of

cryptogramic plants in Batavia (Netherlands).

For more detailed biographical information of

A. de Beyer refer to Bouwer (2006). Miquel

listed the following characters for his new

genus: flowers dioecious and without petals,

calyx 5-partite, anthers extrorse, ovary with

uniovulate locules and stigma pileiform,

entire and sessile. At the time, Miquel

considered Beyeria showed affinities with

the “tribe of Crotons”. However, he noted the

stigma structure distinguished Beyeria from

the genera of the tribe.

In 1845, Johann Klotzsch in Lehmann’s

Plantae Preissianae, published the genus

578

Austrobaileya 7 ( 4 ): 577-639 ( 2008 )

Calyptrostigma Klotzsch to accommodate

Croton viscosus Labill. and in a footnote

described two new species (Calyptrostigma

ledifolium Klotzsch and C. oblongifolium

Klotzsch) based on collections in the Lindley

Herbarium (now at CGE) and Willdenow

Herbarium (now at B-Willd.), respectively

(Klotzsch 1845). Labillardiere based his

Croton viscosus on material he collected from

southern Australia in 1792 while a member of

D’Entrecasteaux’s expedition (Labillardiere

1806). Klotzsch placed Calyptrostigma

within the tribe Crotoneae Blume, but did not

indicate the affinities of his new genus.

In the previous year Miquel (1844) had

stated in the protologue of Beyeria that it

had not been possible to confirm whether

Labillardiere’s Croton viscosus was the

same as his Beyeria viscosa as no specimen

of Croton viscosus was available to him

at the time. However, he considered that

Labillardiere’s species (Croton viscosus)

seemed most likely to belong to his new genus

Beyeria.

Eleven years later, Sonder (1857)

transferred Calyptrostigma ledifolium and

C. oblongifolium to Beyeria and created a

new name, Beyeria preissii Sond., for the

plants Klotzsch described under the name

“C. viscosum Klotzsch” at the same time

specifically excluding Croton viscosus of

Labillardiere. Sonder listed Croton viscosum

Labill. as a synonym of Beyeria viscosa. The

name Beyeria preissii was nomenclaturally

superfluous when published by Sonder, since

he included within this species a duplicate of

the type of B. viscosa Miq.

Halford & Henderson, Revision of Beyeria

In a precursory paper to the publication

of De Candolle’s Prodromus, Jean Muller

(Muller Argovensis) (1865) established the

genus Beyeriopsis and described five species

( B. cygnorumMull.Arg.,B. cinerea Mull.Arg.,

B. latifolia Mull.Arg., B. similis Mull.Arg. and

B. brevifolia Mull.Arg.) based on collections

from south-western Western Australia made

by James Drummond and Augustus Oldfield.

He also transferred Beyeria lepidopetala

F.Muell. to Beyeriopsis. He considered

Beyeriopsis to be closely related to Beyeria

from which it was distinguished on the basis

of its staminal structure. Beyeriopsis was

described as having staminal filaments bifid at

the apex and anthers with 2 discrete separate

locules compared with Beyeria which had

staminal filaments entire and anthers with 2

separate but contiguous locules. Muller also

divided Beyeria into two sections ( B. sect.

Discobeyeria, and B. sect. Eubeyeria ( nom.

inval. = B. sect. Beyeria)) based principally on

the presence or absence of a disc and petals

in the flowers. However, at the time, he did

not arrange the then known species into the

sections, but named two new species ( Beyeria

drummondii Mull.Arg. and B. lasiocarpa

Mull.Arg.) which he indicated belonged to

B. sect. Discobeyeria and B. sect. Beyeria

respectively.

The following year in De Candolle’s

Prodromus , Muller (1866) maintained the

two genera, enumerating seven species in

Beyeriopsis , which included the new species

B. cyanescens Mull.Arg. He divided the genus

into two unnamed ‘sections’, one ‘Flores

dioici’ contained.#. cygnorum,B. lepidopetala,

B. cinerea and B. cyanescens the other ‘Flores

monoici’ contained B. latifolia, B. similis and

B. brevifolia. Muller accepted five species as

belonging to Beyeria , placing B. backhousei

Hook.f. and B. oblongifolia (Klotzsch) Solid,

in synonymy of B. ledifolium (Klotzsch) Sond.

and B. viscosa respectively and recognising

seven varieties; B. viscosa var. genuina

( =B. viscosa var. viscosa), B. viscosa var.

oblongifolia, B. viscosa var. minor, B. viscosa

var. amoena, B. ledifolia var. genuina (=

B. ledifolia var. ledifolia), B. ledifolia var.

backhousei (‘backhousif) and B. ledifolia

var. angustifolia. He maintained the sectional

division in Beyeria with B. sect. Discobeyeria

579

containing a single species B. drummondii,

while if sect. Beyeria contained B. lasiocarpa,

B. viscosa, B. ledifolia and B. opaca F.Muell.

Baillon (1866) considered the differences

in the androecium between Beyeria and

Beyeriopsis were insufficient to warrant

recognition of two genera. However, he didn’t

transfer the Beyeriopsis species of Muller’s

to Beyeria. Baillon transferred the name

Hemistema ( L Hemistemma ’) lechenaultii

DC. to Beyeria, described B. uncinata

Baill. (now in synonymy of Eremophila

sturtii R.Br.), B. lasiocarpa forma denudata

Baill, B. lechenaultii forma genuina

(= B. lechenaultii forma lechenaultii ),

B. lechenaidtii forma pernettioides Baill.,

B. lechenaultii forma myrtoides Baill.,

B. lechenaidtii forma rosmarinoides Baill.,

B. lechenaultii forma salsoloides Baill.,

B. lechenaidtii forma vaccinioides Baill.

and placed Calyptro stigma ledifolium,

C. oblongifolium and B. drummondii in

synonymy of B. lechenaultii (DC.) Baill.,

B. viscosa and B. opaca respectively.

In his account of Beyeria in Flora

Australiensis, Bentham (1873) followed

Baillon in treating Beyeriopsis as congeneric

with Beyeria and transferred Muller’s

Beyeriopsis species to Beyeria. He recognised

13 species, and described B. viscosa var.

linearis Benth. and B. opaca var. linearis

Benth., but did not comment on the

infraspecific taxa previously published by

Muller and Baillon. Bentham arranged the

species into three sections based primarily

on stamen and stigma features. The first

section Beyeria sect. Eubeyeria (nom. inval.

= B. sect. Beyeria) was characterised by

“anthers twice as long as broad, adnate to an

entire or scarcely lobed connective, stigma

entire or scarcely lobed” and included four

species namely B. viscosa, B. opaca (for which

B. lechenaultii,B. ledifoliaSond. non Klotzsch

and B. backhousei were listed as synonyms),

B. lasiocarpa and B. uncinata (=Eremophila

sturtii). The second section Beyeria sect.

Beyeriopsis (nom. superfl.) was characterised

by “anthers short, the locules quite distinct,

either adnate to a deeply 2-lobed connective or

partially free with the connective more entire

and stigmas entire or scarcely lobed” and

580

contained eight species namely B. latifolia,

B. cygnorum (Miill.Arg.) Benth., B. cinerea

(Miill.Arg.) Benth., B. cyanescens (Miill.Arg.)

Benth., B. lepidopetala, B. similis (Miill.Arg.)

Benth., B. brevifolia (Miill.Arg.) Benth. and

B. drummondii (for which Calyptrostigma

ledifolium was listed as a synonym).

The name Beyeria sect. Beyeriopsis was

nomenclaturally superfluous when published

by Bentham, since he included within this

section B. drummondii which was the sole

species Miifler had included in the section he

earlier had named Beyeria sect. Discobeyeria.

The third section Bentham recognised was

Beyeria sect. Oxygyne F.Muell. characterised

by “anthers of Beyeriopsis , stigma deeply

lobed male flowers in a loose raceme” and

contained a single species namely B. tristigma

F.Muell.

In the most recent and complete

taxonomic treatment of Beyeria , Griming

(1913) recognised 12 species and, following

Bentham, grouped them into three sections.

Gruning’s treatment of the genus differed

from Bentham’s in excluding Beyeria uncinata

(= Eremophila sturtii R.Br.), maintaining B.

lechenaultii and B. opaca as separate species

and placing B. drummondii and B. backhousei

in synonymy of B. lechenaultii. Griming

maintained the four varieties of Beyeria viscosa

described by Muller (1866), described B. opaca

var. longifolia Griming, B. lechenaultii var.

latifolia Griming and made new combinations

from B. backhousei, Calyptrostigma ledifolium,

B. lechenaultii forma rosmarinoides and B.

drummondii as the varieties B. lechenaultii var.

backhousei (Hook.) Griming, B. lechenaultii

var. ledifolium (Klotzsch) Griming, Beyeria

lechenaultiivar. rosmarinoides (Bai 11.) Griming

and B. lechenaultii var. drummondii (Mull.

Arg.) Griming.

In the present study the authors have

maintained the sectional division of Beyeria

as presented by Griming (1913) excluding

B. sect. Oxygyne. Halford & Henderson (2005)

established the genus Shonia R.J.F.Hend. &

Halford to accommodate Beyeria tristigma,

the sole species included in B. sect. Oxygyne.

As stated previously (Halford & Henderson

2005, 2007; Webster 1994; Wurdack et

Austrobaileya 7 ( 4 ): 577-639 ( 2008 )

al. 2005) Beyeria is closely related to

Ricinocarpos Desf., Shonia R.J.F.Hend.

& Halford and Bertya Planch, within the

Euphorbiaceae. Beyeria is distinguished

from Ricinocarpos, Shonia and Bertya by the

following combination of features: flowers in

fascicles or cymose clusters or solitary, male

flowers with staminal filaments free, erect

to spreading on a flat or slightly convex to

hemispherical receptacle and stigmas entire,

dilated and forming a cap over the top of

the ovary, discoid or rarely with 2 appressed

limbs or shallowly 3-lobulate. For a key to

the Australian genera of Euphorbiaceae tribe

Ricinocarpeae refer to Halford and Henderson

(2005).

Materials and methods

This revision is based on an assessment

of morphological characters of about 1350

dried herbarium specimens and field studies

undertaken by the second author from 1988 to

1992 and by the first author in 2006. Selected

herbarium collections from herbaria AD, B,

BRI, CANB, HO, K, LD, MEL, NE, NSW,

P and PERTH were studied and annotated.

Acronyms used here and elsewhere to indicate

herbaria holding particular specimens are

those listed by Holmgren et al. (1990). All

specimens cited have been examined by

one or both of the authors, unless indicated

otherwise by ‘ n.v.\

Colour descriptions of vegetative and

floral parts are either from the information on

herbarium labels or from photographs taken

by the second author during field studies.

Measurements listed are based upon the

total variation observed in the herbarium

specimens examined. Information on plant

size, flowering and fruiting times, and habitat

of occurrence were obtained from herbarium

labels. All measurements were made either

on fresh material, dried material, material

preserved in 70% ethanol or dried material

reconstituted by placing in boiling water for

a few minutes. The distribution maps were

produced with Maplnfo Version 3 and are

based on herbarium specimen locality data.

Common abbreviations used in the text are

N.P (National Park), N.R. (Nature Reserve),

N.S.W. (New South Wales), Qld (Queensland),

581

Halford & Henderson, Revision of Beyeria

S.A. (South Australia), S.F./S.F.R. (State Forest/

State Forest Reserve), Tas. (Tasmania), Vic.

(Victoria), W.A. (Western Australia).

Taxonomy

Beyeria Miq., Ann. Sci. Nat. Bot. ser. 3, 1:

350-352, t. 15 (1844). Type: B. viscosa Miq.

Beyeriopsis Mull. Arg., Linnaea 34: 56 (1865).

Type: Beyeriopsis brevifolia Mull.Arg.,

[= Beyeria brevifolia (Miill.Arg.) Benth.]

(lecto: fide Wheeler 1975: 535).

Calyptrostigma Klotzsch in J.G.C.Lehmann,

PI. Preiss. 1(2): 175 (1845). Type: C. viscosum

(Labill.) Klotzsch

Monoecious or rarely dioecious shrubs or

small trees, often resinous on most parts;

branches glabrous or with an indumentum of

stellate or rarely simple hairs. Leaves spirally

alternate, exstipulate, petiolate or rarely

sessile, simple, entire, margins flat, recurved

or revolute, marginal glands absent or present

mostly on blade proximally. Flowers axillary

or terminal on short axillary branchlets,

pedicellate rarely ± sessile, in few-flowered

fascicles or cymose clusters or solitary,

bracteate, gamosepalous; calyx deeply

5(rarely 4 or 6)-lobed, subequal, imbricate

(quincuncial); petals present, 5(rarely 4) or

absent, slightly shorter than or equal in length

to calyx lobes; disc present or absent, of discrete

alternipetalous glands or forming a continuous

glandular ring. Male flowers with receptacle

flat to hemispherical; stamens numerous (>

9); filaments free, ± erect or spreading, mostly

bifid distally; anthers dorsifixed, extrorse, of

two separate, parallel but contiguous cells,

each transverse or lateral on the apex of the

filament, dehiscing by longitudinal slits;

pistillode absent. Female flowers with calyx

lobes persistent, rarely accrescent; petals

sometimes marcescent; ovary 2- or 3(rarely

l)-locular with one pendant ovule in each

locule; style short or ± obsolete; stigma

elobate or shallowly 3-lobulate, calyptriform,

discoid, or rarely bipartite, persistent. Fruits

capsular, ellipsoid, ovoid or subglobose,

mostly trilobate, glabrous or densely hairy,

smooth or papillose, mostly 2- or 3(rarely

l)-seeded, separating septicidally often into

three 2-valved cocci leaving a persistent

columella. Seeds ellipsoid and dorsi-ventrally

compressed, globose or obloid, carunculate;

testa smooth, shiny, often blotched; caruncle

creamy-white, yellowish-white, or light

brown, waxy-fleshy; endosperm copious;

embryo linear, in the middle of the endosperm,

cotyledons longer than the radicle.

Key to sections of Beyeria

Flowers apetalous or if rudimentary petals present then < half the

length of calyx lobes; calyx lobes of male flowers spreading at anthesis

.B. sect. Beyeria

Flowers with petals, slightly shorter than or equal to calyx lobes; calyx lobes

of male flowers ± erect, enclosing or appressed to androecium at anthesis

.B. sect. Beyeriopsis

Beyeria Miq. sect. Beyeria, Miill.Arg.,

Linnaea 34: 58 (1865). Type: B. viscosaMiq.

Beyeria sect. Discobeyeria Mull.Arg.,

Linnaea 34: 58 (1865). Type: B. drummondii

Miill. Arg.

Beyeria sect. Eubeyeria Miill.Arg., Linnaea

34: 58 (1865), nom. inval.

Shrubs or small trees. Flowers apetalous or

petals rarely present as rudimentary lobes.

Male flowers with calyx lobes spreading at

anthesis; receptacle convex to hemispherical;

filaments entire; anthers of two separate,

obloid or linear, parallel but contiguous

cells, each lateral on the apex of the filament.

Female flowers with stigmas calyptriform or

rarely bipartite.

582

Austrobaileya 7(4): 577-639 (2008)

Distribution : The species of Beyeria sect. Australia, Queensland, New South Wales,

Beyeria occur in Western Australia, South Victoria and Tasmania.

Key to species of Beyeria sect. Beyeria

1 Ovaries and fruits densely hairy, non-resinous (N.S.W., Qld, Vic.)

.2. B. lasiocarpa

1. Ovaries and fruits glabrous or with a few scattered hairs, usually resinous .2

2 Stigmas bipartite; lobes ± oblong, c. 1 mm long, appressed to ovary (S.A.)

.5. B. subtecta

2. Stigmas entire, calyptriform.3

3 Abaxial surface of leaf blades densely tomentellous with crispate hairs >

0.1 mm long, never resinous (N.S.W., S.A., Tas., Vic., W.A.).3. B. lechenaultii

3. Abaxial surface of leaf blades glabrous or densely puberulous with hairs

< 0.1 mm long, often obscured by resinous covering.4

4 Leaf blades lanceolate; fruits 10-12 mm long (N.S.W., Vic.).1. B. lanceolata

4. Leaf blades narrow-elliptic to elliptic or narrow-obovate to obovate; fruits

5-9 mm long.5

5 Leaf blades narrow-obovate, 7-20 mm long, < 5 mm wide; ovaries 2-

locular (N.S.W., S.A., Vic.. W.A.).4. B. opaca

5. Leaf blades narrow-elliptic to elliptic or narrow-obovate to obovate, 20-

90 mm long, 5-30 mm wide; ovaries 3-locular (N.S.W., Qld, S.A., Tas.,

Vic., W.A.).6. B. viscosa

1. Beyeria lanceolata Halford & R.J.F.Hend.

species nova B. viscosae Miq. maxime affinis

sed foliorum lamina lanceolata non angusto-

elliptica usque ad obovata et fructibus

longioribus (10-12 mm longis non 6-8 mm

longis) differt. Typus: New South Wales.

Neenah Creek, southern slopes of Nungatta

Mt, c. 48 km SSW of Eden, 16 February 1984,

T.James 519 & M.Taylor (holo: NSW; iso:

MEL).

Illustration : Jeanes (1999: 67, fig. lOj) as

Beyeria viscosa.

Dioecious or occasionally monoecious, tall

slender shrubs or small trees to 6 m high,

resinous on young shoots, buds and adaxial leaf

surface. Main trunk with fine flaky, reddish-

brown bark. Young branchlets of unknown

colour in fresh state, angular becoming terete

with age, glabrous; older branchlets with grey,

± smooth bark. Leaves petiolate; petioles 3-6

mm long, glabrous, 3-6 pairs of sessile glands

on adaxial face; blades lanceolate, 55-120

mm long, 6-16 mm wide, length:width ratio

7-15:1; adaxial surface glabrous and smooth;

abaxial surface papillose and sparsely hairy

with stellate hairs c. 0.7 mm across (obscured

by resinous covering); base cuneate; margins

flat or slightly recurved; apex acute; midvein

impressed adaxially, abaxially prominently

raised, rounded and glabrous; secondary

and tertiary veins obscure; marginal glands

absent. Flowers pedicellate; male flowers

axillary, solitary or in shortly pedunculate

clusters of up to 4 flowers; female flowers

axillary or terminal on a short axillary

branchlet, solitary; peduncles up to 4 mm

long, glabrous; bracts narrow-ovate, up to 3

mm long, rounded at apex, glabrous; pedicels

± angular, glabrous, stouter in female than

in male flowers; calyx lobes 5, of unknown

colour when fresh, glabrous; petals absent;

disc obscure or absent. Male flowers with

pedicels 7-9 mm long; calyx lobes broad-

ovate, 3.2-4.5 mm long, 3-4 mm wide,

concavo-convex, the margins erose, obtuse to

rounded at apex; receptacle c. 2.5 mm across,

glabrous; stamens 40; filaments ± erect,

0.1-0.3 mm long, glabrous, entire; anthers c.

1.3 mm long. Female flowers with pedicels

Halford & Henderson, Revision of Beyeria

6-10 mm long; calyx lobes ± appressed to

and enclosing gynoecium apart from stigma,

triangular to ovate, 2.5-3.5 mm long, 2-3.5

mm wide, the margins entire, rounded to

obtuse at apex; ovary subglobose, ± trigonal,

c. 2.5 mm long, glabrous, thickly resinous, 3-

locular; style c. 0.2 mm long, glabrous; stigma

calyptriform, 1.9-2.5 mm across, umbilicate,

glabrous, with margins entire. Fruits

subglobose, 10-12 mm long, 9-12 mm across,

2- or 3-seeded, glabrous, smooth; persistent

calyx c. one third the length of mature fruit.

Seeds ellipsoid, dorsi-ventrally compressed,

57-6.5 mm long (including caruncle), 4-4.8

mm across, 3-3.1 mm in thickness; testa

dark brown or mottled dark brown and grey;

caruncle c. 1.5 mm long, and 2.5 mm wide,

yellowish-white. Fig. 1.

Additional selected specimens examined: New South

Wales, gully on the E side of Mt Wog Wog, Nalbaugh

N.P., Dec 1985, Albrecht 2343 (MEL, NSW); c. 4.8 km

SSW of Nungatta Peak trig, Apr 1986, Albrecht 2570

(MEL); Rock Flat Creek, Sep 1950, Wakefield 4604

(MEL). Victoria. Mt Dandenong, Aug 1914, Campbell

s.n. (MEL 114246); Dandenong Ranges below Mt

Dandenong, May 1995, Jeanes 176 (MEL); Sportsman

Creek, Sep 1984, Beauglehole ACB77347 (CANB, MEL);

Haunted Stream, Oct 1977, Gullan & Norris 208 (MEL);

Bunga Creek, near Cunningham, Jan 1911, St John s.n.

(MEL 622345); by Castleburn Creek on Dargo road, Sep

1979, Walsh NGW130 (MEL); Mountain Creek, Deddick

track crossing, 4.6 km NW of Mt Joan, Sep 1979, Forbes

167 (MEL, NSW); Wulgulmerang, c. 70 km NNW of

Orbost, Nov 1962, Willis s.n. (CANB, MEL 503120,

NSW); Little River Gorge, c. 75 km N of Orbost, Nov

1968, Willis s.n. (MEL 114337); c. 0.5 km SW of Mt

Merragunegin, Sep 1988, Albrecht 3711 (MEL); Upper

Genoa River, Oct 1948, Wakefield 3186 (MEL); Upper

Genoa River at junction with Yambulla Creek, Oct 1948,

Willis s.n. (MEL 114244).

Distribution and habitat : Beyeria lanceolata

is known from Nungatta and Nalbaugh

National Parks, southern New South Wales

and in the Dandenong Ranges and East

Gippsland, Victoria (Map 2). It grows on

sheltered rocky hillslopes and in gullies

in open eucalypt forest communities with

rainforest understorey shrubs.

Phenology : Flowers have been collected

throughout the year, particularly from August

to January, fruits from September to March.

Affinities : Beyeria lanceolata is

morphologically most similar to B. viscosa but

differs by its lanceolate rather than narrow-

583

elliptic to obovate leaf blades and its larger

subglobose fruit which are 10-12 mm long

compared with 6-8 mm long in B. viscosa.

Etymology : The specific epithet is from the

Latin lanceolatus, lanceolate, and alludes to

the shape of the leaf blades in this species.

2. Beyeria lasiocarpa Mull.Arg., Linnaea 34:

59 (1865); Beyeria lasiocarpa Mull.Arg. forma

lasiocarpa , Baill., Adansonia 6: 307 (1866).

Type: [New South Wales.] Twofold Bay, s.d.,

F.Mueller (holo: G-DC n.v. (microfiche IDC

800-73.2454: II. 7); iso: MEL [4 sheets 114153,

114154, 114156, 114157], NSW 465094).

Illustrations : Costermans (1986: 211); Jeanes

(1999: 67, fig. 10k).

Dioecious or rarely monoecious, spreading,

much-branched shrubs to 3 m high, thinly

resinous on most parts. Young branchlets pale

green and angular becoming grey and terete

with age, glabrous; older branchlets with

grey to black shallowly fissured bark. Leaves

petiolate; petioles 3-7 mm long, glabrous,

with up to 5 pairs of sessile glands on adaxial

face; blades narrow-elliptic, narrow-obovate

or very narrow-ovate, 30-110 mm long, 7-20

mm wide, length:width ratio 5-15:1; adaxial

surface glabrous and ± smooth; abaxial

surface densely hairy with ± sessile, stellate

hairs 0.1-0.2 mm across; base cuneate;

margins slightly recurved; apex obtuse to

acute or rounded, sometimes ultimately

apiculate with extension from midrib;

apiculum slender, < 0.2 mm long; midvein

slightly impressed adaxially, abaxially raised,

glabrous; secondary veins slightly impressed

or obscure adaxially, abaxially slightly raised;

tertiary veins obscure; marginal glands absent.

Flowers pedicellate, males axillary, in shortly

pedunculate cymose clusters of up to 4 flowers

or rarely solitary, females axillary or terminal

on short axillary branchlets, solitary or rarely

in shortly pedunculate cymose clusters of

up to 3 flowers; peduncles up to 5 mm long,

glabrous; bracts ± triangular, 1-3 mm long,

acute at apex, ± glabrous; pedicels glabrous,

stouter and longer on female flowers than

on male flowers; calyx lobes 5, green, white

puberulous abaxially, glabrous adaxially,

the margins erose, rounded at apex; petals

absent; disc ± a continuous ring, glabrous.

584

Austrobaileya 7(4): 577-639 (2008)

Fig. 1. Beyeria lanceolata. A. branchlet with female flowers x 1. B. transverse section of leaf x 9. C. side view of male

flower x 6. D. face view of female flower x 9. E. side view of female flower x 6. F. side view of fruit x 3. G. face view

of fruit x 6. H. abaxial view of seed x 6. I. side view of seed x 60. A-E from Jeanes 176 (MEL), F-I from lames 519

& Taylor (NSW). Del. W.Smith.

Halford & Henderson, Revision of Beyeria

Male flowers with pedicels 4-5 mm long;

calyx lobes broad-ovate, 3-4 mm long, 3-4.5

mm wide; receptacle c. 3 mm across, stellate-

tomentose; stamens 35-50; filaments erect,

1-1.5 mm long, glabrous or with a few simple

hairs proximally; anthers 0.7-1 mm long.

Female flowers with pedicels 7-15 (-20) mm

long; calyx lobes spreading, ovate to broad-

ovate or suborbicular, 3-5 mm long, 1.8-5.5

mm wide; ovary subglobose, ± trigonal, c. 3

mm long, densely hirsute, 3-locular; style up

to 0.7 mm long, hirsute; stigma calyptriform,

3-5 mm across, umbilicate, glabrous, with

margins irregularly toothed. Fruits depressed

globose, 6-10 mm long, 7-10 mm across,

3-seeded, moderately hairy with simple ±

erect hairs <1.5 mm long; persistent calyx

c. one third the length of mature fruit. Seeds

ellipsoid, dorsi-ventrally compressed, 4-5.5

mm long (including caruncle), 2.3-37 mm

across, 2-2.8 mm in thickness; testa black to

dark brown; caruncle 0.6-1.5 mm long, 1.5-

2.3 mm wide, light brown.

Additional selected specimens examined : Queensland.

Wide Bay District: Swain Peak, 7 km NW of Yandina,

Mar 1993, Bean 5808 (BRI); Gheerulla Falls, 5 km W

of Mapleton, Nov 1990, Bean 2686 (BRI). Moreton

District: Neurum Creek, between Mt Delaney & Mt

Archer, Mar 1989, Forster PIF5006 et al. (BRI); 4.3

km past bridge over Little Nerang Creek, Mudgeeraba

to Springbrook road, Feb 1991, Forster PIF7802 &

Leiper (BRI). New South Wales. Upper Guy Fawkes

River, Boundary Creek, Jan 1992, Glimour 7297 (BRI,

MEL, NSW); Doyles River S.F., 88 km SE of Walcha,

on the Oxley Highway, Aug 1985, Foreman 988 (BRI,

MEL, NE, NSW); Deua N.P., Diamond Creek, below

3rd waterfall, Jan 1994, Taws 357 & Scott (NSW);

Carters Creek, Currowan S.F., NW of Batemans Bay,

Dec 1973, Pullen & Story 8747 (BRI, CANB, MEL,

NSW, PERTH); North Brooman S.F., c. 4.8 km NW of

Termed, Nov 1973, Pullen & Story 8625 (BRI, NSW);

Pebbly Beach, 20 km NE of Batemans Bay, Dec 1969,

Briggs 3108 (NSW); Bemboka S.F., 2.6 km at 80° from

Bemboka Peak, Mar 1992, Telford 11564 & Crawford

(BRI, MEL, NSW); summit of Dr Georges Mt, 10 km

NE of Bega on Tanja road, Jul 1995, Jobson 3648 (MEL,

NSW); Mt Dromedary, Dec 1962, Adams 505 (BRI,

MEL, NSW); Egan Peaks N.R., Sep 1984, Albrecht 856

(MEL, NSW); 0.5 km upstream from Newtons Flat along

creek, Nadgee N.R., Oct 1992, Zich 199 (CANB, NSW).

Victoria. Fainting Range track, 1.5 km E of Tambo

River, Jan 1984, Parkes EG147A (MEL); c. 100 m N of

junction between Mottle Range road and Monument

track, 6 km SSE of Mt Tara, Jan 1984, Parkes EG155

(BRI, MEL); between Snowy River at Wood Point and

Long Point track, Aug 1979, Walsh NGW178 (MEL);

beside un-named creek falling N to Genoa River, 3.7 km

NE from township, Oct 1991, Walsh 3205 (BRI, MEL).

585

Distribution and habitat : Beyeria lasiocarpa

occurs in coastal and subcoastal areas of

eastern Australia from Mapleton, south¬

eastern Queensland, southwards through New

South Wales to Bairnsdale, East Gippsland,

Victoria (Map 3). It grows in wet sclerophyll

forest and moist gullies and shaded sites in

dry sclerophyll forest communities and on

rainforest margins.

Phenology : Flowers and fruits have been

collected throughout the year.

Affinities : Beyeria lasiocarpa is similar to

B. viscosa but can be distinguished by its

densely hairy (rather than glabrous) ovary

and fruits and the lack of resin on the abaxial

surface of its leaf blades.

3. Beyeria lechenaultii (DC.) Baill.,

Adansonia 6: 307 (1866), (7 eschenaultif);

Hemistema ('Hemistemmaf lechenaultii DC.,

Syst. Nat. 414 (1817); Beyeria lechenaultii

(DC.) Baill. forma lechenaultii , Baill.,

Adansonia6\ 307 (18 66); Beyeria lechenaultii

(DC.) Baill. var. lechenaultii , Griming in

A.Engler, Pflanzenr. H58: 70 (1913). Type:

[South Australia.] Novo Holl. et ile St.

Francois [St Francis Isles], s.d., Leschenault

s.n. (holo: G-DC n.v., (microfiche IDC 800-

73. 41:1. 1); iso: P 152760 (photo at BRI)).

Beyeria lechenaultii forma elaeagnoides

Baill., Adansonia 6: 308 (1866),

(‘eloeagnoides ’). Type: [Victoria.] near Lake

King, February 1855, F.Mueller s.n. (lecto

[here chosen]: MEL 114170).

Beyeria lechenaultii forma myrtoides Baill.,

Adansonia 6: 308 (1866). Type: [Victoria:]

Cape Otway, s.d., [F.Mueller?] (holo: MEL

114177).

Beyeria lechenaultii forma pernettioides

Baill., Adansonia 6: 307-308 (1866). Type:

[Victoria.] Portland, s.d., W.Allett s.n. (lecto

[here chosen]: MEL 114182).

Beyeria lechenaultii forma rosmarinoides

Baill., Adansonia 6: 308 (1866); Beyeria

lechenaultii var. rosmarinoides (Baill.)

Grlining in A.Engler, Pflanzenr. H58: 70

(1913), nom. superfl. etincor. Type: [Victoria.]

summit of Mount Ligar, s.d., [F.Mueller ?]

(lecto [here chosen]: MEL 114186; isolecto:

MEL 114179).

586

Beyeria lechenaultii forma salsoloides Baill.,

Adansonia 6: 308 (1866). Type: [Victoria.]

N.W. Victoria, s.d., Mr Lockhart Morton s.n.

(holo: MEL 114181).

Beyeria lechenaultii forma vaccinioides

Baill., Adansonia 6: 308 (1866). Type:

[Victoria.] mouth of the Glenelg [River], s.d.,

WAllett s.n. (holo: MEL 114180).

Beyeria lechenaultii var. latifolia Griming in

A.Engler, Pflanzenr. H58: 71 (1913). Type:

[Victoria.] Cape Otway, s.d., [F.Mueller?]

(lecto [here chosen]: MEL 114177).

Beyeria backhousei Hook.f., FI. Tasman. 1:

339 (1857), (‘ backhousii ’); Beyeria ledifolia

var. backhousei (Hook.f.) Miill.Arg. in A.DC.,

Prodr. 15(2): 203 (1866), (‘ backhousii ’);

Beyeria lechenaultii var. backhousei (Hook,

f.) Griming in A.Engler, Pflanzenr. H58: 70

(1913), (‘ backhousii Type: [Tasmania.]

Flinders Island, Bass Straits, s.d., R.C.Gunn

540 (syn: K); [Tasmania.] Flinders Island,

s.d., JBackhouse s.n. (syn: K).

Beyeria drummondii Miill.Arg., Linnaea

34: 58-59 (1865); Beyeria lechenaultii var.

drummondii (Miill.Arg.) Griining in A.Engler,

Pflanzenr. H 58: 70-71 (1913). Type: [Western

Australia.] s.loc., s.d., J.Drummond214 (lecto

[here chosen]: G-DC n.v. [microfiche IDC

800-73. 2454: II. 6]; isolecto: K, PERTH).

Calyptrostigma ledifolium Klotzsch in

J.G.C.Lehmann, PI. Preiss. 1: 176 (1845);

Beyeria ledifolia (Klotzsch) Sond., Linnaea

28: 565 (1857); Beyeria ledifolia (Klotzsch)

Sond. var. ledifolia, Miill.Arg. in A.DC.,

Prodr. 15(2): 203 (1866); Beyeria lechenaultii

var. ledifolia (Klotzsch) Griining in A.Engler,

Pflanzenr. H58: 70 (1913). Type: [Western

Australia.] in New Holland, s.d., J.Drummond

s.n. (holo: n.v.).

Beyeria ledifolia var. angustifolia Miill.Arg.

in A.DC., Prodr. 15(2): 203 (1866). Type:

[Western Australia.] at King George Sound,

1860, Cuming s.n. (holo: G-DC n.v. [microfiche

IDC 800-73. 2455:1. 1],

Beyeria opaca var. linearis Benth., FI.

Austral. 6: 65 (1873), Beyeria lechenaultii var.

rosmarinoides Ewart, FI. Victoria 726 (1930),

nom. illeg. Type: [South Australia.] near

Austrobaileya 7(4): 577-639 (2008)

Adelaide, s.d., Blandowski s.n. (lecto [here

chosen]: K).

Beyeria viscosa var. angustifolia F.Muell. &

Tate, Trans. &Proc. Roy. Soc. South Australia

16: 341 (1896). Type: [Western Australia.]

Fraser Range, 23 October 1891, R. Helms s.n.

(lecto [here chosen]: MEL 1560026; isolecto:

NSW [3 sheets 257570, 257573, 257576,]).

Beyeria lechenaultii var. latifolia Ewart, FI.

Victoria 726 (1931), nom. illeg. non Griining

(1913). Type: not designated.

Beyeria lechenaultii forma genuina Baill.,

Adansonia 6: 307 (1866), nom. inval. [ICBN,

Art. 24.3],

Beyeria ledifolia var. genuina Miill.Arg. in

A.DC., Prodr. 15(2): 203 (1866), nom. inval.

[ICBN, Art. 24.3],

Beyeria lechenaultii var. genuina Griining

in A.Engler, Pflanzenr. H58: 70 (1913), nom.

inval. [ICBN, Art. 24.3],

Illustrations : Cunningham et al. (1982: 453);

Costermans (1986: 211); Weber (1986: 742,

fig. 398a); Jeanes (1999: 67, fig. lOh); Corrick

& Fiihrer (2000: 81).

Dioecious or rarely monoecious, open to

dense, spreading to erect shrubs up to 2 m

high, usually resinous on most parts. Young

branchlets pale green when fresh, angular

becoming terete with age, with resinous

longitudinal ridges, puberulous between

ridges but usually obscured by resinous

covering; older branchlets with grey or pale

brown, ± tessellated bark. Leaves petiolate;

petioles 1-3 mm long, ± glabrous, glands

absent; blades linear to narrow-oblong,

narrow-obovate to obovate, narrow-elliptic

to elliptic or rarely narrow-ovate, 9-40 mm

long, 0.9-15 mm wide, length:width ratio

2-25:1; adaxial surface glabrous and smooth,

usually thickly resinous; abaxial surface

densely woolly with crispate hairs up to 1

mm long; base cuneate, obtuse or rounded;

margins flat or slightly to strongly recurved

sometimes to midrib concealing abaxial leaf

surface; apex acute, rounded to truncate,

emarginate or rarely obcordate, sometimes

ultimately apiculate with extension from

midrib; apiculum stout, up to 0.5 mm long,

Halford & Henderson, Revision of Beyeria

bent upward; midvein obscure or slightly

impressed adaxially, abaxially prominently

raised, rounded and glabrous; secondary

and tertiary veins obscure; marginal glands

occasionally present at base of blade, 1 or

2 per side of midrib, sessile, smooth, c. 0.1

mm across. Flowers pedicellate, males

axillary, solitary or in shortly pedunculate

clusters of up to 3 flowers, females axillary or

terminal on short axillary branchlet, solitary;

peduncles up to 1 mm long, resinous; bracts

narrow-ovate, up to 3 mm long, acute at apex,

± glabrous; pedicels ± angular, puberulous

but usually concealed by resinous covering,

stouter on female than on male flowers;

calyx lobes 5(rarely 4 in male flowers), pale

green; petals absent or rarely present in male

flowers as small rudimentary lobes; disc a

continuous glandular ring, ± fleshy, glabrous.

Male flowers with pedicels 3.5-10 mm long;

calyx lobes broad-ovate or suborbicular, 1-

3.2 mm long, 1-3 mm wide, concavo-convex,

puberulous abaxially medially, glabrous

adaxially, the margins erose, obtuse to

rounded or acute at apex; receptacle 1-1.5 mm

across, densely hairy or with a few scattered

hairs; stamens 10-60; filaments ± erect,

0.1-0.3 mm long, glabrous, entire; anthers

0.3-1 mm long. Female flowers with pedicels

2- 10 mm long; calyx lobes ± appressed to

and enclosing gynoecium apart from stigma,

triangular to ovate, 1-2 mm long, 0.8-1.5

mm wide, puberulous abaxially, glabrous

adaxially, the margins entire, acute, rounded

to obtuse at apex; ovary subglobose and ±

trigonal or laterally compressed, 1.2-2 mm

long, glabrous or sometimes with scattered

stellate hairs, thickly resinous, 3(rarely 2)-

locular; style 0.3-0.7 mm long, glabrous;

stigma calyptriform, 1-1.8 mm across,

shallowly umbilicate, glabrous, with margins

erose. Fruits ellipsoid-subglobose, rarely

3- lobate, 4.5-8 mm long, 3.8-7 mm across,

2- or 3-seeded, glabrous, ± smooth; persistent

calyx c. one third the length of mature fruit.

Seeds ellipsoid, dorsi-ventrally compressed,

3.5-6.8 mm long (including caruncle), 2-4

mm across, 1.5-3.3 mm in thickness; testa

dark brown or mottled dark and light brown;

caruncle 0.7-1.3 mm long, 1.2-2.6 mm wide,

yellowish-white.

587

Distribution and habitat : Beyeria

lechenaultii occurs in southern Australia

south of latitude 30° from near Perth, Western

Australia eastward through South Australia

to the Griffith area, New South Wales, Lake

Entrance, Victoria, Bass Strait Islands and

the north-west coast of Tasmania, with a

disjunct population recorded in the Pilliga

area, New South Wales (Map 4). In inland

areas it mainly grows in shrubland, mallee,

and open woodland communities, in sandy

loams or red earths, on undulating plains or

occasionally in mountain ranges, gorges or

associated with rocky outcrops. In coastal

areas it grows in heathland and shrubland

communities in sandy soils on coastal dunes

or rocky headlands.

Phenology : Flowers have been recorded

throughout the year, particularly from July

to December, fruits in March and April and

from July to January.

Typification: In the protologue of Beyeria

lechenaultii forma elaeagnoides, Baillon

(1866) cited two collections Wilhelmi, Port

Lincoln (herb. EMuell.!)’ and ‘ F.Mueller ,

prope Lake King Island, in virgultis (herb.!)’.

Two collections, which are considered as

syntypes of the name B. lechenaultii forma

elaeagnoides , have been located amongst

material loaned to BRI from MEL [114170 and

114208], The sheet 114170 has been selected

as lectotype because it is part of original

material and has morphology that agrees with

the brief description in Baillon’s protologue of

this forma.

In the protologue of Beyeria lechenaultii

forma pernettioides , Baillon (1866) cited

four collections from F.Mueller’s herbarium

(now MEL). Four collections [114174, 114182,

114195 and 114204] which are considered to

be syntypes and one collection [2062920]

considered to be a possible isosyntype have

been located amongst material on loan to

BRI from MEL. The collection made by

Allett from Portland [114182] is here selected

as the lectotype because it is the most ample

and best preserved of the specimens and

has morphology that agrees with the brief

description given in Baillon’s protologue of

this forma.

588

In the protologue of Beyeria lechenaultii

forma rosmarinoides , Baillon (1866) cited

some 12 collections, two from Preiss’s

herbarium and ten from F. Mueller’s herbarium

(now MEL). The Mueller collection labelled

“summit of Mount Ligar” [MEL 114186] is

here selected as the lectotype of this name

because it is part of the original material and

has morphology that agrees with the brief

description given in Baillon’s protologue of

this forma.

In the protologue of Beyeria lechenaultii

var. latifolia , Griming (1913) cited two

specimens namely ‘without location details,

(F. Mueller !)’ and ‘Victoria. Sorrento,

(Weindorfer !)’. He also made reference

to four names, ‘ Beyeria lechenaidtii var.

/?-£ Baill.’ (= Beyeria lechenaultii forma

pernettioides Baill., Beyeria lechenaultii

forma elaeagnoides Baill., Beyeria

lechenaultii forma myrtoides Baill. and

Beyeria lechenaultii forma vaccinioides

Baill.). We have been unable to locate

either of the two specimens cited. To fix the

application of Griining’s Beyeria lechenaultii

var. latifolia , the type of Beyeria lechenaultii

forma myrtoides Baill., is here selected as

lectotype of Griining’s name because of the

material located it best fits the description

given in Griining’s protologue of this variety.

In the protologue of Beyeria drummondii,

Muller (1865) cited two James Drummond

collections from Western Australia, namely

“In Nova Hollandia austre-occidentali

ad Swan River {Drummond n. 13\ 2141)”.

We have been unable to locate material of

Drummond n. 13 in loans from K or PERTH

and it is not recorded on the microfiche of De

Candolle’s herbarium. However, we have seen

on the microfiche of De Candolle’s herbarium

(G-DC) what we considered to be the syntype

Drummond n. 214. As well we have located

two isosyntypes amongst material loaned

to BRI from PERTH and K. The collection

{Drummond 214 ) that is located at G-DC is

here selected as the lectotype of Muller’s name

because it is part of original material and has

morphology that agrees with the description

in Muller’s protologue of this species.

Austrobaileya 7(4): 577-639 (2008)

In the protologue of Beyeria viscosa var.

angustifolia , Mueller & Tate (1896) cited a

single collection from Fraser Range collected

by R. Helms on the Elder Expedition in the

north-west of South Australia and across the

Great Victoria Desert of Western Australia.

We have located three duplicates (one at

MEL and two at NSW). The MEL specimen

[1560026] is here selected as the lectotype of

this Mueller name because it is part of original

material and has morphology that agrees with

the description in Mueller’s protologue of this

species.

In the protologue of Beyeria opaca var.

linearis , Bentham (1873) cited two collections,

namely ‘Alps on the Macalister, F.Mueller .;

near Adelaide, Blandowski’. A single sheet

containing both specimens was located

amongst material loaned to BRI from K. The

specimen collected by Blandowski (on the left

of the sheet) is here selected as the lectotype

of Bentham’s name because it has morphology

that agrees with the protologue and is the more

ample of these two collections.

Affinities : Beyeria lechenaultii is

morphologically most similar to B. subtecta

but differs in having generally larger leaves

(9-40 mm long x 0.9-15 mm wide compared

with 4-10 mm long x 0.8-1.6 mm wide) and

fruits (4.5-8 mm long x 3.8-7 mm across

compared with c. 3 mm long and 2.5 mm

across), and female flowers with an entire

rather than a bipartite stigma.

Notes: In the past, Beyeria lechenaultii has

been confused with B. sulcata var. sulcata

in the western part of its range but is easily

distinguished from this taxon by having

apetalous flowers or petals reduced to small

rudimentary lobes compared with petalous

flowers in B. sulcata var. sulcata.

This species, as circumscribed here,

varies considerably in the length and width

of its leaves. This variation has led several

authors in the past to formally describe many

infraspecific and specific taxa (Hooker 1857;

Muller 1865; Baillon 1866; Griming 1913).

Much more material has since been collected

and although the extreme forms within this

species differ considerably from each other,

much of the morphological variation appears

589

Halford & Henderson, Revision of Beyeria

to intergrade, making it difficult in assigning

material to one or other of them. For this

reason we have not formally recognised these

variants. However, the more notable variants

are discussed below.

‘Typical variant’ sens. lat.

Leaf blades linear to narrow-oblong or rarely

narrow-obovate, mostly 9-27 mm long, 1.5-

3.5 mm wide, the margins strongly recurved

almost to midrib or sometimes to midrib

concealing abaxial leaf surface, apex mostly

rounded to truncate. Male flowers: calyx

lobes 5, 1.5-3.2 mm long, stamens 20-60.

Female flowers: ovary 3-locular. Fruits 6-8

mm long.

This is the most widespread variant and occurs

from York, Western Australia, eastwards

through South Australia to New South Wales

and north-western Victoria.

Selected specimens examined: Western Australia.

Yallari Reserve [Yallarin], Jun 1925, Franks 43

(PERTH); c. 35 km SE of Coolgardie, on road to

Norseman (Coolgardie - Esperance Highway), Sep 1988,

Henderson H3165 (BRI); 3.5 miles [c. 6 km] E of Eucla,

Oct 1966, George 8511 (NSW, PERTH); McPherson

Rock, 31 km S of Norseman, Aug 1975, Crisp 972

(CANB, PERTH); 27 km S from Balladonia towards

Mt Ragged, Sep 1983, Taylor 1525 & Ollerenshaw

(CANB, MEL); near Point Dover, Jul 1967, Wilson

5946 (PERTH). South Australia, c. 2 km N of New

West Bore, Bibliando Station, Apr 1974, Crisp 733

(AD, CANB); Ceduna, Sep 1968, Canning WA/68 2324

(CANB); 3 Calpatanna Waterhole Conservation Park,

c. 26 km SSE of Streaky Bay, Nov 1989, Davies 1449

& Hadlow (MEL); Bay of Shoals, Kangaroo Island,

Aug 1984, Weston 6 (CANB); by Chauncey’s Line, c.

5 km SE of Harriet Hill and c. 18 km SW of Murray

Bridge, Oct 1958, Schodde 911 (AD). New South Wales.

Pilliga Scrub, Aug 1953, Jordan s.n. (AD 966071380);

c. 2 km N of Goolgowi towards Merriwagga, Oct 1989,

Henderson & Turpin H3323 (BRI, NSW); 4 km W of

Kamarah, Nov 1975, Crisp 1508 (AD, CANB); 2.3 km

WNW of Kamarah on the Ardlethan - Griffith road, Apr

1988, Dolby 88/13 etal. (MEL, NSW). Victoria. Sunset

Country, Millewa South Bore track, c. 0.3 km S of Bore

to first major dune crest, Aug 1986, Lucas 207 (MEL);

Big Desert, 8 km S of Murrayville on Nhill road, Oct

1979, Corrick 6389 (AD, MEL); 30.6 km SE of Walpeup

on road to Patchewollock, Oct 1980, Corrick 6701 et al.

(AD, CANB, MEL); 9 km W of Swan Hill on road to

Sea Lake, Sep 1989, Henderson & Turpin H3270 (BRI);

2.7 km NE of Dimboola, Aug 1995, Jobson 3715 (BRI,

MEL).

‘Broad-leafed variant’

Leaf blades narrow-elliptic to elliptic, narrow-

obovate to obovate or rarely narrow-ovate,

mostly 10-30 mm long, 2.5-13 mm wide,

the margins flat or slightly recurved, apex

obtuse, acute or rounded, rarely emarginate,

sometimes ultimately apiculate with extension

from midrib. Male flowers: calyx lobes 5,2.5—

3 mm long, stamens 20-35. Female flowers:

ovary 3-locular. Fruits 6-7 mm long.

Occurs mostly in coastal areas from Fowlers

Bay, South Australia, east to Lakes Entrance,

Victoria, and onto the Bass Strait Islands

and the north-west corner of Tasmania. Also

occurs on Mt Arapiles and Mitre Rock in

north western Victoria.

The following names placed in synonymy

of Beyeria lechenaultii are applicable to this

variant: B. lechenaultii forma elaeagnoides

Baill., B. lechenaultii forma myrtoides Baill.,

B. lechenaultii forma pernettioides Baill.,

B. lechenaultii forma vaccinioides Baill.,

B. lechenaultii var. latifolia Griming and

B. backhousei Hook.f.

Selected specimens examined: South Australia. Cape

Wiles, Mar 1960, Filson 1608 (MEL); far SW tip of Yorke

Peninsula, Innes N.P, Sep 1989, Smith 89/17 (MEL);

Beachport, Jan 1963, Womersley 10 (MEL, NSW).

Victoria. Mitre Rock, Sep 1996, Ross 3791 (MEL); Mt

Arapiles, Nov 1968, Beauglehole ACB29646 (MEL);

Kalimna Heights, Clara Street, Dec 1995, Stephens 14

(MEL); Glenelg River mouth track to near Ocean Beach

carpark, Dec 1983, Forbes 1925 & Beauglehole (MEL);

Port Campbell N.P, Oct 1966, Finck & Beauglehole

ACB21640 (MEL, NSW); Otways, Parker River, c. 100 m

upstream from mouth, Jan 1991, Albrecht 4696 (MEL);

Cape Schanck coastal park, Nov 1982, Beauglehole

ACB71544 & Elmore (MEL). Tasmania. King Island,

Golf Links, Currie Harbour, Jan 1968, Cameron s.n.

(HO 27189); Erith Island, Kents Group, Bass Strait,

Dec 1987, Whinray 9030 (MEL); Prime Seal Island,

Furneaux Group, Aug 1972, Whinray 1473 (MEL); W

end of Trouser Point, Flinders Island, Oct 1985, Collier

782 (HO); Cape Barren Island, Furneaux Group, Oct

1973, Whinray 606 (MEL); Australian Point, Apr 1984,

Moscal 7874 (HO, MEL); Temma, Feb 1976, Richley s.n.

(HO 27722).

‘Narrow-leafed variant’

Leaf blades linear, mostly 10-30 mm long,

0.9-1.2 mm wide, the margins strongly

recurved to midrib concealing abaxial leaf

surface, apex obtuse, rounded or rarely

obcordate. Male flowers: calyx lobes 4 or

590

5, 1-1.7 mm long, stamens 10-24. Female

flowers: ovary 2- or 3-locular. Fruits 4.5-5

mm long.

Occurs in coastal or near coastal areas

from Beaufort Inlet eastward to near the

Oldfield River in Western Australia.

This variant has generally shorter and

narrower leaves as well as generally smaller

male flowers, occasionally 2-locular ovaries

and generally smaller fruit than the other

variants Newbey 1245 8c 11231 (both PERTH).

Two collections ( Newbey 1245 (PERTH) and

Newbey 11231 (PERTH)), which are relatively

atypical for this variant, have leaves with

obcordate or emarginate apices.

The names Beyeria drummondii Mull.

Arg. and B. lechenaultii var. drummondii

(Milll.Arg.) Griming, placed above in the

synonymy of B. lechenaultii , are applicable to

this variant.

Selected specimens examined: Western Australia.

Beaufort Inlet, Feb 1964, Newbey 1245 (PERTH);

junction of Twertup Creek & Fitzgerald River, Fitzgerald

RiverN.R, Aug m5, Newbey 10946(BRI, PERTH); inlet

S of Fitzgerald, Jul \91\,Aplin 4787 (PERTH); Phillips

River, 19 km from Ravensthorpe on Ravensthorpe

- Albany road, Oct 1966, Muir 4150 (MEL); 30 km SW

of Ravensthorpe, Oct 1986, Newbey 11231 (PERTH);

Quoin Head Campground, Fitzgerald River N.P, W of

Hopetoun, Nov 1996, Lepschi BJL3194 & Laity (BRI,

MEL); Fitzgerald River N.P, c. 50 m NW of Quoin

Head campsite, Apr 2001, Hisiop 2204 (BRI); Fitzgerald

River, Sep 1948, Gardner 9251 (PERTH); Coppermine

Creek, Fitzgerald River N.P, Oct 1970, Royce 9291

(PERTH); Oldfield River at crossing of the Esperance

- Ravensthorpe road, c. 105 km W of Ravensthorpe, Oct

1968, Eichler 20225 , 20226 (PERTH).

4. Beyeria opaca F.Muell., Trans. Philos. Soc.

Victoria 1: 16 (1855); Beyeria opaca F.Muell.

var. opaca , Benth., FI. Austral. 6: 65 (1873).

Type: [Victoria.] Murray, s.d., F.Mueller s.n.

(lecto [here chosen]: MEL 2062921; isolecto:

K).

Beyeria opaca var. latifolia J.M.Black, FI.

5. Austral. , 1 st edn, 357 (1924). Type: [South

Australia.] Ooldea, January 1917, s.coll. (holo:

AD 98046056; iso: AD 966050860).

Beyeria opaca var. typica Griming in

A.Engler, Pflanzenr. H58: 69 (1913), nom.

inval. [ICBN, Art. 24.3],

Austrobaileya 7(4): 577-639 (2008)

Illustrations : Cunningham et al. (1982: 453);

Weber (1986: 742, fig. 398b); Jeanes (1999: 67,

fig. lOi); Corrick 8c Fiihrer (2000: 81).

Dioecious, spreading, much-branched shrubs

to 1 m high, resinous on most parts. Young

branchlets pale green and angular becoming

terete with age, glabrous; older branchlets

with grey to black bark. Leaves petiolate;

petioles 1-E5 mm long, glabrous, glands

absent; blades narrow-obovate, 7-20 mm

long, E2-4 mm wide, length:width ratio 5—

8:1; adaxial surface glabrous and ± smooth,

resinous; abaxial surface papillose and hairy

with ± sessile, stellate hairs c. 0.05 mm

across (obscured by resinous covering); base

cuneate; margins ± flat or slightly recurved;

apex rounded; midvein faintly impressed

adaxially, abaxially faintly raised, glabrous

and resinous; secondary and tertiary veins

obscure; marginal glands occasionally present

at base of blade, 1 per side of midrib, sessile,

smooth, c. 0.1 mm across. Flowers pedicellate,

males axillary, solitary, females axillary or

terminal on short axillary branchlet, solitary;

bracts ± triangular, E3-1.5 mm long, acute at

apex, glabrous; pedicels ± glabrous, stouter

on female flowers than on male flowers; calyx

lobes 5, glabrous, concavo-convex, acute to

rounded at apex; petals absent in male flowers,

absent or rudimentary in female flowers;

disc obscure or absent. Male flowers with

pedicels 4-7 mm long; calyx lobes yellowish

green, reddened in parts, suborbicular or

ovate, 1.5-3.5 mm long, 2-2.8 mm wide, the

margin erose; receptacle 1-E5 mm across,

glabrous; stamens 25-40; filaments erect,

0.8-1 mm long, glabrous; anthers 0.7-0.9 mm

long. Female flowers with pedicels E5-4

mm long; calyx lobes green, ± appressed to

and enclosing gynoecium apart from stigma,

ovate, 0.9-E8 mm long, 0.7-E5 mm wide, the

margin entire; petals when present, oblong, <

0.2 mm long; ovary ellipsoid, 0.8-1 mm long,

glabrous or densely hairy distally, 2-locular;

style ± obsolete; stigma calyptriform, c. 1 mm

across, shallowly umbilicate, glabrous, with

margins ± entire. Fruits ellipsoid, laterally

compressed, 4.5-8.5 mm long, 3.7-7 mm

across, 2.5-4.5 mm in thickness, 1- or 2-

seeded, glabrous, ± smooth; persistent calyx

c. one fifth the length of mature fruit. Seeds

Halford & Henderson, Revision of Beyeria

ellipsoid, dorsi-ventrally compressed, 3.5-5

mm long (including caruncle), 2.2-3 mm

across, 2-2.7 mm in thickness; testa mottled

light to dark brown; caruncle c. 1 mm long

and 1.5 mm wide, creamy-white.

Additional selected specimens examined : Western

Australia, c. 50 m SW of Bushfire Rock road, 46.5 km

E of Hyden, Sep 1991, Mollemans & Mollemans 4624

(BRI); 76.9 km E of Norseman on Eyre Highway, Aug

1995, Cranfield 10049 (BRI, PERTH). South Australia.

1 mile [c. 1.6 km] S of Maralinga, Aug 1956, Forde 466

(MEL); Lake Tallacootra, Oct 1983, Weber 8191 (BRI);

30.5 miles [c. 49 km] from Why alia towards Kimba,

Aug 1968, Canning 2174 (CANB; NSW); 18 km W of

Walker Flat, Sep 1979, Spooner 6530 (AD, NSW); 26 km

NE of Blanchetown on Waikerie road, Oct 1975, Haegi

689 (AD); turnoff to Loxton, just before Alawoona

on Alawoona to Paruna road, Oct 1989, Henderson &

Turpin H3320 (BRI); c. 9 km N of Overland Corner, Sep

1971, Donner 3697 (AD, NSW). Victoria, c. 11.7 km

along road which runs W of the Sunset Tank - Merrinee

road, Sep 1980, Short 1194 & Corrick (AD, CANB,

MEL, NSW); 3 km W of abandoned railway, near Rocket

Lake, Oct 1977, Crisp 3321 (CANB, MEL); 36 miles [c.

58 km] W of turnoff, 14 miles [c. 23 km] N of Birthday

Tank, Sunset Country, Sep 1965, Filson 7449 (AD, MEL,

NSW); Kulkyne S.F., W of the old Calder Highway, 2

miles [ c. 3 km] N of the Lake Hattah to Hattah Station

road, Nov 1958, Aston 152 (MEL); c. 15 km S of Hattah,

heading towards Ouyen on Calder Highway, Sep 1989,

Henderson & Turpin H3268 (BRI); Big Desert, 44 km N

of Broken Bucket bore, Oct 1979, Corrick 6360 (MEL).

New South Wales. 14 miles [c. 22 km] from Mt Hope

to Euabalong, Sep 1969, Dunlop 1554 (CANB); 26 miles

[c. 42 km] from Lake Cargelligo towards Mt Hope, Sep

1966, Phillips s.n. (AD 97031105, CANB [CBG016390]);

c. 22 km E of Rankins Springs, Oct 1972, Jackson 2153

(AD, MEL); Pulletop N.R., c. 20 km SW of Rankins

Springs, Nov 1975, Crisp 1454 (AD, CANB); c. 17 km

S of Weethalle on Weethalle - Barellan road, Sep 1989,

Henderson & Turpin H3258 (BRI).

Distribution and habitat : Beyeria opaca

occurs in southern Australia, from Whyalla,

south-eastern South Australia, eastward to

Weethalle and Lake Cargelligo, New South

Wales and south to Swan Hill in Victoria,

with disjunct populations near Hyden and

Norseman in southern Western Australia and

near Maralinga, Lake Tallacootra and the

Gawler Range, south-western South Australia

(Map 5). It grows in mallee communities on

red sandy soils on sandy flats or dunefields.

Phenology : Flowers have been collected

throughout the year, particularly from August

to November, fruits in March, April and from

June to November.

591

Affinities : Beyeria opaca resembles the

narrow leaf forms of B. viscosa but can be

distinguished by its smaller habit (shrub to 1

m high compared with shrub to small tree to

5 m high) and leaves (7-20 mm long x 1.2-4

mm wide compared with 20-90 mm long x 5-

30 mm wide), 2-locular ovary, and generally

fewer stamens in each male flower.

Typification : In the protologue of Beyeria

opaca , Mueller (1855) did not cite any

particular collection but listed a number of

localities “In the Mallee scrub, between Lake

Lalbert, Lake Tyrrell, and the Murray River”.

Four collections which we consider part of the

original material that Mueller used to draw

up his description of this species have been

located amongst material loaned to BRI from

MEL and K. Two of the MEL sheets [114214

andl 14225] are labelled “Beyeria viscosa

var opaca, ‘Mallee Scrub”. The other MEL

sheet [2062921, ex herb. Sonder] and the K

sheet are labelled “Beyeria opaca, Murray,

Mueller”. The MEL sheet [2062921] is chosen

as the lectotype of this name because it is an

ample specimen and has morphology that

agrees with the description in the protologue.

5. Beyeria subtecta J.M.Black, FI. S.

Austral. 357-358 (1924). Type: [South

Australia.] Cygnet River, Kangaroo Island,

20 October 1908, H.H.D.Griffith s.n. (holo:

AD 9831247).

Illustration : Barker & Dashorst (1984: 139).

Dioecious, much-branched shrubs to 0.6

m high, resinous on most parts. Young

branchlets of unknown colour when fresh,

angular becoming terete with age, puberulous

between resinous longitudinal ridges; hairs

stellate, sessile, c. 0.1 mm across, sometimes

obscure by resin; older branchlets with grey

to black shallowly fissured bark. Leaves

petiolate; petioles 0.5-1.5 mm long, glabrous,

glands absent; blades linear or very narrow-

obovate, 4-10 mm long, 0.8-1.6 mm wide,

length:width ratio 5-7:1; adaxial surface

glabrous and ± smooth, resinous; abaxial

surface densely hairy with ± sessile, stellate

hairs c. 0.1 mm across; base cuneate; margins

recurved to midrib usually concealing

abaxial leaf surface; apex obtuse to rounded,

ultimately apiculate with extension from

592

midrib; apiculum slender, up to 0.4 mm

long; midvein slightly impressed adaxially,

abaxially raised, flattened, glabrous and

resinous on abaxial face; secondary and

tertiary veins obscure; marginal glands

rarely present at base of blade, 1 per side of

midrib, sessile, smooth, c. 0.1 mm across.

Flowers pedicellate, axillary, solitary; bracts

oblong, 0.6-1.2 mm long, acute at apex, ±

glabrous; pedicels stellate pubescent, stouter

on female flowers than on male flowers; calyx

lobes 5(rarely 4 in male flowers), ovate, ±

flat, stellate-puberulous abaxially, glabrous

adaxially, the margins entire; petals absent;

disc a continuous ring, ± fleshy, glabrous.

Male flowers with pedicels 1.5-3 mm long;

calyx lobes light green to yellow or white,

reddened in parts, 1.5-2.2 mm long, 1.2-

1.4 mm wide, acute, obtuse or rounded at

apex; receptacle c. 0.4 mm across, glabrous;

stamens 15-25; filaments erect to spreading,

0.1-0.3 mm long, glabrous; anthers 0.4-0.6

mm long. Female flowers with pedicels up

to 1 mm long; calyx lobes green, ± appressed

to and enclosing gynoecium apart from

stigma, 1.6-2.1 mm long, 0.7-0.8 mm wide,

acute at apex; ovary ellipsoid, dorsi-ventrally

compressed, c. 1 mm long, sparsely to densely

stellate-puberulous (usually obscured by

resinous covering), 2-locular; style c. 0.4 mm

long, sparsely to densely stellate puberulous

(usually obscured by resinous covering);

stigma bipartite; lobes ± oblong, c. 1 mm long,

glabrous, appressed to ovary, with margins

entire. Fruits ellipsoid, c. 3 mm long and 2.5

mm across, 1-seeded, sparsely stellate hairy

distally; persistent calyx c. half the length of

mature fruit. Seed not seen.

Additional specimens examined: South Australia,

along side of road heading ENE towards Point Morrison

near junction with Kingscote to Penneshaw road, Oct

1983, Davies & Bushman 15 (MEL); Beyeria N.P,

Kangaroo Island, Apr 1991, Spencer 1057 & Worboys

(MEL); 700 m S of six way road intersection, 12.5 km SE

of Cygnet River township on Kingscote to Penneshaw

road, Oct 1983, Davies & Bushman 9 (BRI, CANB);

700 m ENE of road intersection NE of Dead Horse

Lagoon, Kangaroo Island, Oct 1983, Davies & Bushman

4 (CANB); road junction 2.3 km NW of intersection N

of Kiowie Station on Kingscote to Penneshaw road, Oct

1983, Davies & Bushman 2 (MEL); 5 miles [ c. 8 km]

from American River, towards Kingscote, Kangaroo

Island, Sep 1965, Phillips SA/65 776 (CANB); proposed

American River Tip, top of Muston Hill, 9.7 km E of

American River, Aug 1984, Weston 92 (CANB).

Austrobaileya 7(4): 577-639 (2008)

Distribution and habitat : Beyeria subtecta is

endemic to Kangaroo Island, South Australia

(Map 6). It grows in Eucalyptus cneorifolia

and Melaleuca uncinata heathland and

mallee on clay on a laterite substrate and in

Casuarina shrubland and mallee on white

sand with ironstone over clay intruded with

limestone.

Phenology : Flowers have been collected from

August to October, fruits in October.

Affinities: Beyeria subtectaismorphologically

most similar to B. lechenaultii but differs in

having generally smaller leaves (4-10 mm

long x 0.8-1.6 mm wide compared with 9-

40 mm long x 0.9-15 mm wide) and fruits

(c. 3 mm long and 2.5 mm across compared

with 4.5-8 mm long x 3.8-7 mm across), and

female flowers with a bipartite rather than an

entire stigma.

Notes : Beyeria subtecta is listed as

‘Vulnerable’ in the South Australian

threatened species list under the National

Parks and Wildlife Act (South Australia)

1972.

6. Beyeria viscosa Miq., Ann. Sci. Nat. Bot.

ser. 3,1: 350-352, 1 .15 (1844); Beyeria viscosa

Miq. var. viscosa , Mull.Arg. in A.DC., Prodr.

5(2): 202 (1866). Type: [Western Australia] in

colliculis arenosis insulae Rottenest, N.Holl.,

19 August 1839, L.Preiss 2387 (holo: U 19718;

iso: G-DC n.v. [microfiche IDC 800-73. 2454:

III. 1], LD 99/018-0879, MEL [3 sheets

2062937 ex Sonder Herb., 2062918 ex Sonder

Herb., 114267 ex Steetz Herb.], P 275782).

Beyeria viscosa var. amoena Mull.Arg.

in A.DC., Prodr. 15(2): 202 (1866). Type:

southern Australia, 1863, F.Mueller s.n. (lecto

[here chosen]: G-DC n.v. [microfiche IDC

800-73. 2454: III. 7]; iso?: K).

Beyeria viscosa var. latifolia Benth.,

FI. Austral. 6: 65 (1873). Type: Western

Australia. Swan River to Cape Richer [Riche],

s.d., J.Drummond 5th coll. n. 217 (holo: K [ex

herb. Benth.]; iso: K [ex herb. Hook.]).

Beyeria viscosa var. minor Mull.Arg. in

A.DC., Prodr. 5(2): 202 (1866). Type: [New

South Wales.] interior west from Wellington

Valley, 1825, A.C.Cunningham s.n. (holo: G-

DC n.v. [microfiche IDC 800-73. 2454: III.

6 ]).

593

Halford & Henderson, Revision of Beyeria

Calyptrostigma oblongifolium Klotzsch in

J.G.C.Lehmann, PL Preiss. 1(2): 176 (1845);

Beyeria oblongifolia (Klotzsch) Sond.,

Linnaea 28: 564 (1857); Beyeria viscosa

var. oblongifolia (Klotzsch) Miill.Arg. in

A.DC., Prodr. 15(2): 202 (1866). Syntypes:

[Australia.] Nova Hollandia, s.d., s.coll. (B-

Willd. 17848 fol. 1 n.v. [microfiche IDC 7440.

1291: III. 4]); s.loc., s.d., Labillardiere s.n. (B-

Willd. 17849 fol. 1 and 2. n.v. [microfiche IDC

7440. 1291: III. 6,7]).

Croton viscosus Labill., Nov. Holl. PI. 2: 72-

73, t. 222 (1806), ( £ viscosum ’); Calyptro stigma

viscosum (Labill.)Klotzschin J.G.C.Lehmann,

PL Preiss. 1(2): 176 (1845). Type: [Australia.]

Nova Hollandia, ora austro-occidentalis, s.d.,

Labillardiere s.n. (lecto [here chosen]: FI

165908 n.v. [transparency at BRI]).

Beyeria viscosa var. obovata C.T.White, Proc.

Roy. Soc. Queensland 50: 86 (1939). Type:

Queensland. Mitchell District. Torrens

Creek, 19 March 1933, C.T.White 873L (holo:

BRI; iso: A n.v., BRI, K n.v.).

Beyeriapreissii Sond., Linnaea 28: 564 (1857)

nom. superfl. Type: [Western Australia.]

s.loc. [s.d. in sandy hills on Rottnest Island,

19 August 1839], [J.A.L.] Preiss 2387 (lecto

(here chosen): MEL 2062918 ex Sonder Herb.;

isolecto: LD 99/018-0879, MEL [2 sheets

2062937 ex Sonder Herb, 114267 ex Steetz

Herb.], G-DC n.v. [microfiche IDC 800-73.

2454: III. 1], P 275782, U 19718).

Beyeria viscosa var. genuina Miill.Arg. in

A.DC., Prodr. 15(2): 202 (1866), nom. inval.

[ICBN, Art. 24.3],

Illustrations : Cunningham et al. (1982: 453);

Rippey & Rowland (1995: 107); Jeanes (1999:

67, fig. 10j).

Dioecious or rarely monoecious, dense

rounded or slender shrubs or rarely small

trees to 5 m high, usually resinous on young

shoots, buds and adaxial surface of leaf. Main

trunk with flaky, grey bark. Young branchlets

of unknown colour in fresh state, angular

becoming ± terete with age, densely hairy with

minute appressed hairs (obscured by resinous

covering); older branches with grey, ± smooth

or shallowly fissured bark. Leaves petiolate;

petioles 3-6 mm long, glabrous, 6 or 7 pairs of

sessile glands on adaxial face; blades narrow-

elliptic to elliptic, oblong elliptic, narrow-

obovate to obovate, 20-90 mm long, 5-30 mm

wide, length:width ratio 3-7:1; adaxial surface

glabrous and smooth; abaxial surface hairy

with stellate hairs c. 0.1 mm across, usually

covered with resin; base obtuse to cuneate or

attenuate; margins flat or slightly recurved;

apex rounded, obtuse, acute or rarely retuse

to emarginate; midvein impressed adaxially,

abaxially prominently raised, rounded and

glabrous; secondary and tertiary veins

obscure or slightly impressed adaxially,

obscure or visible abaxially; marginal glands

absent. Flowers pedicellate, males axillary,

solitary or in shortly pedunculate racemose

clusters of up to 5 flowers, females axillary

or terminal on short axillary branchlet,

solitary or rarely in shortly pedunculate lax

clusters of up to 3 flowers; peduncles up to

2-4 mm long, glabrous or if puberulous,

then indumentum usually concealed by

resinous covering; bracts oblong, 1-3 mm

long, rounded at apex, glabrous; pedicels

angular, puberulous, indumentum usually

concealed by resinous covering, stouter on

female flowers than on male flowers; calyx

lobes 5(rarely 4 or 6), pale green, glabrous or

puberulous and usually concealed by resinous

covering; petals absent; disc obscure, absent

or ± a continuous ring. Male flowers with

pedicels 3-10 mm long; calyx lobes oblong,

ovate, broad-ovate or suborbicular, 2.8-5.2

mm long, 2.6-4 mm wide, concavo-convex,

the margins erose, obtuse to rounded at apex;

receptacle c. 2 mm across, glabrous or hairy;

stamens 30-70; filaments ± erect, 0.5-1.8 mm

long, glabrous or rarely with scattered hairs

proximally, entire; anthers 0.9-2 mm long.

Female flowers with pedicels 5-20 mm long;

calyx lobes ± appressed to and enclosing

gynoecium apart from stigma, oblong or

suborbicular, 2.2-3 mm long, 1.5-2.3 mm

wide, plano-convex, the margins entire,

rounded to obtuse at apex; ovary subglobose,

± trigonal, 1-2.6 mm long, glabrous, thickly

resinous, 3-locular; style obsolete, < 0.1 mm

long; stigma calyptriform, 1.8-2 mm across,

glabrous, with margins erose. Fruits globose,

weakly to strongly trilobate, green or purple,

6-8 mm long, 6-8.5 (-10) mm across, 3-

seeded, glabrous or rarely with scattered

594

stellate hairs, resinous, smooth; persistent

calyx c. one third the length of mature fruit.

Seeds ellipsoid, dorsi-ventrally compressed,

4.5-72 mm long (including caruncle), 2.6-4

mm across, 1.9-3.5 mm in thickness; testa

light brown, brown to reddish brown, usually

mottled; caruncle 1-2 mm long, 1.5-2 mm

wide, yellowish-white.

Additional selected specimens examined : Western

Australia, near S end of Useless Harbour in Sharks Bay,

1863, Brown s.n. (MEL 114268); East Wallabi Island

[Houtman Abrolhos], Jul 1970, Ashby 3263 (PERTH);

Garden Island, Sep 1978, Marchant s.n. (PERTH

06845983); near upper carpark. Bunker Bay, Cape

Naturaliste, WNW of Busselton, Sep 1988, Henderson

H3200 (BRI); SE peninsula. Middle Island, Recherche

Archipelago, Nov 1973, Weston 8791 & Trudgen

(CANB, PERTH). South Australia. Boston Point, Eyre

Peninsula, [in 1851?,] Wilhelm s.n. (MEL 2065872).

Queensland. North Kennedy District. Sally’s Mesa, 34

km from Greenvale on Charters Towers road, Feb 1994,

Forster PIF14975 & Bean (BRI). Mitchell District. 41

km NW of Torrens Creek, Apr 1993, Thompson HUG410

et al. (BRI). Leichhardt District. Little St Peter, 10

miles [c. 16 km] N of Springsure, Sep 1985, O’Keeffe

781 (BRI, CANB). Darling Downs District. Dunmore

S.F., 49.3 km WSW of Dalby, Apr 1995, Halford Q2488

(BRI). New South Wales. MacIntyre Falls, 35.2 km NW

of Ashford on the Wallangarra road. Mar 1987, Coveny

12517 et al. (BRI, NSW); c. 6 km N of Cobar on road

to Bourke, Oct 1989, Henderson & Turpin H3367 (BRI,

NSW); Mt Wombelong, Warrumbungle Ranges, Dec

1973, Streimann HS550 (AD, BRI, CANB); Bungonia

Gorge, S of Marulan, Nov 1966, Pullen 4166 (BRI, MEL,

NSW); The Rock NR, 30 km SW of Wagga Wagga, Nov

1975, Crisp 1835 (AD, BRI, NSW); Erskine Creek,

Nepean River, Sep 1888, Maiden s.n. (NSW 463865).

Victoria. Sorrento, Nov 1903, Weindorfer s.n. (MEL

114270). Tasmania. 10 miles [c. 16 km] from Launceston,

Jan 1949, Burbidge 2941 (CANB, HO); St Pauls River

E of Cutoff Hill, May 1985, Collier 519 (HO); Longley,

beside North West Bay River, Jan 1989, Davies 782 &

Ollerenshaw (BRI, MEL); above old quarry on Mystery

Caves track, Jan 1983, Forbes 1334 (CANB, HO, MEL).

Distribution and habitat : Beyeria viscosa

has a disjunct distribution occurring in

western and eastern Australia (Map 7). In

south-western Western Australia it occurs in

scattered populations in coastal areas from

Shark Bay south to Yallingup and on the south

coast in the Archipelago of the Recherche

near Esperance. It grows in heathland and

shrubland communities in sandy soils over

limestone or rarely on sandy clay soils over

granite. There is a single record (George 14178

(PERTH)) from a slope near a creek in Jarrah/

Marri forest on sandy loam over laterite.

Austrobaileya 7(4): 577-639 (2008)

In eastern Australia it occurs in subcoastal

and inland areas from Ingham, north-eastern

Queensland south to the southern coast of

New South Wales, Flinders Island, eastern

Tasmania, and single collections from Boston

Point, Eyre Peninsula, South Australia, and

Sorrento, Victoria. In inland areas it grows

mostly in mallee or Eucalyptus woodland

communities, in skeletal sandy loams or red

earths, in gullies, on ridges, rocky slopes and

hilltops. In subcoastal areas it grows in mostly

Eucalyptus dominated open forests in gullies,

along river banks and on gentle slopes.

Phenology : Flowers have been recorded

throughout the year, particularly from June to

January, fruits in August to November.

Typification : In the protologue of Croton

viscosus , Labillardiere (1806) did not cite any

particular collection but made the statement

“Habitat in terra Van-Leuwin”. The location

cited in the protologue refers to the area around

the present town of Esperance and Esperance

Bay, Western Australia (Nelson 1974, 1975).

Labillardiere made botanical collections in

this area in 1792 as well as in Tasmania while

a member of D’Entrecasteaux’s expedition.

Labillardiere’s Herbarium now resides in

Florence (FI-WEBB). On our behalf Alex

Chapman, while Australian Botanical Liaison

Officer at K, visited FI and searched for relevant

type material. We have seen photographs

of two specimens from Labillardiere’s

Herbarium, namely sheets stamped with

Herbarium Webbianum numbers 165908 and

165910. The sheet numbered 165908 has pinned

to it a printed label “Herbarium Webbianum,

ex Herb. Labillardiere” with the hand written

annotation ‘Nova Hollandia, ora austro-

occidentalis’, two pages with a handwritten

draft of the description that appeared in the

protologue of Croton viscosus and three small

slips of paper with handwritten notes mostly of

which are indecipherable. The four specimens

that are attached to the sheet all appear to be

part of a single collection, although three bear

female flowers or fruit and the other bears

male flowers. The sheet numbered 165910

has a printed label “Herbarium Webbianum,

ex Herb. Labillardiere” with the annotation

‘Nova Hollandia et terra Dieman’ and three

small slips of paper with handwritten notes that

Halford & Henderson, Revision of Beyeria

are mostly indecipherable. The four specimens

that are attached to the sheet appear to be from

two separate collections. The sheet numbered

165908 is here selected a lectotype for the

name Croton viscosus as it is part of the

original material and has morphology that

agrees with the description in the protologue.

Muller (1866) cited two collections in his

protologue of Beyeria viscosa var. amoena

namely “In Nova Hollandia in monte Flinders

(hb. Hook!)” and “in Australia Felice (Ferd.

Muell.! in hb. DC.)”. The collection in the

Hooker herbarium has not been located in

material on loan from Kew of this genus.

The collection in G-DC is selected here as

lectotype of this name.

Affinities : Beyeria viscosa is morphologically

similar to B. lanceolata and B. lasiocarpa. It

differs from B. lanceolata in having narrow-

elliptic to obovate rather than lanceolate leaf

blades and smaller globose fruit (6-8 mm

long compared with 10-12 mm long). Beyeria

viscosa differs from B. lasiocarpa by having

glabrous (rather than densely hairy) ovaries

and fruits and a resinous abaxial surface of

its leaf blades.

Notes : The combination Beyeria oblongifolia

was also made by J.D.Hooker in December

1857, four months after Sonder’s.

In his protologue of Beyeria viscosa ,

Miquel listed “ Croton viscosum Labillard..?”.

This doubtful inclusion of Labillardiere’s name

has led many authors to presume that Miquel

had made a new combination and therefore

ascribe the authorship of the name B. viscosa

as (Labill.) Miq. However, Miquel made the

statement in the protologue of B. viscosa that

no specimen of Croton viscosus Labill. was

available to him and it had not been possible

to confirm that Croton viscosus Labill.

was the same as his species. We contend

that this clearly indicates that Miquel was

not transferring Labillardiere’s name but

publishing a new name for which Miquel

should be attributed the sole authorship.

This species as circumscribed here varies

considerably in the shape, length and width of

its leaf blades. This variation has led several

authors in the past to describe a number of

infraspecific and specific taxa (Muller 1866;

595

Bentham 1873; White 1939). Much more

material has since been collected and although

the extreme forms within this species differ

considerably, much of the morphological

variation appears to intergrade especially

with the variants in eastern Australia, making

it difficult to assign material unequivocally

to one or other of them. For this reason we

have not formally recognised any variants

here. However, the more notable variants are

discussed below.

In Western Australia, specimens typically

have large (30-70 mm long, 10-30 m wide)

leaves that are narrow-elliptic to elliptic or

rarely narrow-obovate and 2.2 to 3.5 times

as long as wide, obtuse to cuneate leaf bases,

rounded to obtuse leaf apices, secondary

and tertiary veins visible on abaxial surface

of leaves, and a glabrous receptacle in male

flowers. This variant includes the type of the

name Beyeria viscosa Miq.

In Tasmania and Victoria, specimens have

leaves that are generally narrower and more

attenuate (40-90 mm long, 8-18 mm wide)

and 3-5 times as long as wide, cuneate leaf

bases, obtuse to acute leaf apices, secondary

and tertiary veins obscure or faintly visible on

abaxial surface of leaves, and a glabrous or

sparsely hairy receptacle in male flowers.

In southern Queensland and New South

Wales, specimens have typically smaller (20-

50 mm long, 4-10 mm wide) leaves that are

narrow-obovate and 3 to 6 times as long as

wide, cuneate leaf bases, rounded to obtuse

or rarely retuse leaf apices, secondary and

tertiary veins obscure on abaxial leaf surface

of leaves and densely hairy receptacle in male

flowers. In the more mesic habitats in Central

and South Coast, and Southern Tablelands of

New South Wales, the leaves of this variant

are generally broader and longer (50-70 mm

long, 15-20 mm wide) than populations in

drier habitats.

In northern Queensland, specimens have

typically smaller (20-55 mm long, 10-30

mm wide) narrow-obovate to obovate leaves

that 1.8 to 3 times as long as wide, cuneate

to attenuate leaf bases, rounded to retuse or

emarginate leaf apices, secondary and tertiary

veins obscure on abaxial surface of leaves and

a densely hairy receptacle in male flowers.

596

Beyeria sect. Beyeriopsis (Miill.Arg.) Benth.,

FL Austral. 6: 63, 66 (1873); Beyeriopsis Mull.

Arg., Linnaea 34: 56 (1865). Type: B. brevifolia

(Miill.Arg.) Benth.

Shrubs. Flowers petalous, slightly shorter

than or equal to calyx lobes. Male flowers

calyx lobes ± erect enclosing or appressed

to androecium at anthesis; receptacle flat to

convex; filaments entire or bifid distally;

anthers of two separate, obloid, parallel but

Austrobaileya 7(4): 577-639 (2008)

contiguous cells, each transverse on the apex

of the filament; connective extending beyond

anther cells. Female flowers with stigmas

calyptriform, discoid or shallowly 3-lobulate.

Distribution : The species of Beyeria sect.

Beyeriopsis are confined to south-western

Western Australia.

Key to species of Beyeria sect. Beyeriopsis

1 Young branchlets hairy.2

1. Young branchlets glabrous.9

2 Young branchlets densely hairy between glabrous longitudinal ridges;

calyx lobes prominently keeled abaxially.12. B. constellata

2. Young branchlets evenly covered with a sparse to dense indumentum;

calyx lobes rounded abaxially.3

3 Leaf blades > 5 mm wide.16. B. latifolia

3. Leaf blades < 5 mm wide.4

4 Young branchlets with a sparse to moderately dense indumentum of

stellate hairs; rays of hairs spreading to ascending.5

4. Young branchlets with a sparse to dense indumentum of simple or bifid

hairs; hairs ascending to erect.6

5 Pedicels < 7 mm long.10. B. cinerea

5. Pedicels > 7 mm long.17. B. lepidopetala

6 Pedicels > 5 mm long.8. B. brevifolia

6. Pedicels < 5 mm long.7

7 Young branchlets resinous, sparsely hairy; hairs <0.3 mm long; ovaries

3-locular.11. B. cockertonii

7. Young branchlets non-resinous, moderately to densely hairy; hairs > 0.3;

ovaries 2-locular.8

8 Flowers pedicellate; pedicels 2-4 mm long; ovaries glabrous.22. B. simplex

8. Flowers ± sessile; ovaries densely hairy.24. B. villosa

9 Young branchlets longitudinally grooved.10

9. Young branchlets not longitudinally grooved.15

10 Fruiting calyces 4-7 mm long.9. B. calycina

10. Fruiting calyces < 4 mm long .11

11 Leaf apices rounded or obtuse, without excurrent midrib.12

11. Leaf apices rounded, obtuse, acute to attenuate or truncate, ultimately

terminated with excurrent midrib.13

12 Leaf blades < 6 mm long.19. B. physaphylla

12. Leaf blades > 6 mm long.20. B. rostellata

Halford & Henderson, Revision of Beyeria 597

13 Pedicels hairy proximally with minute erect glandular hairs c. 0.05 mm

long. 13. B. disciformis

13. Pedicels glabrous.14

14 Calyx lobes of female flowers suborbicular to very broad-ovate, c. 2.5

mm long; fruiting calyces c. half of fruit length. 18. B. minor

14. Calyx lobes of female flowers broad-oblong or broad-obovate, 0.8-1.3

mm long; fruiting calyces < one third of fruit length. 23. B. sulcata

15 Leaf apices acute to obtuse, ultimately apiculate with extension from

midrib .16

15. Leaf apices obtuse to rounded, without excurrent midrib.18

16 Branchlets and leaves non-resinous; leaf apices with slender apiculum,

0.3-0.6 mm long. 7. B. apiculata

16. Branchlets and leaves mostly thickly resinous; leaf apices with stout

apiculum, up to 0.2 mm long.17

17 Leaf blades 1.4-2 mm wide, length: width ratio < 10; leaf apices rounded

.15. B. lapidicola

17. Leaf blades 0.9-1.5 mm wide, length: width ratio 10-20; leaf apices acute to

attenuate. 23. B. sulcata

18 Adaxial surface of leaf blades smooth, non-resinous. 8. B. brevifolia

18. Adaxial surface of leaf blades papillose, tuberculate, usually resinous.19

19 Male flowers with pedicels 7-11 mm long; ovaries without horn-like

appendages. 15. B. lapidicola

19. Male flowers with pedicels 0.5-4 mm long; ovaries crowned by 4-6 horn¬

like appendages;.20

20 Leaf blades 5-16 mm long; pedicels of male flowers 1-2 mm long; pedicels

of female flowers 1-3 mm long; subapical appendage on fruit up to 1

mm long. 14. B. gardneri

20. Leaf blades 20-50 mm long; pedicels of male flowers 2-3 mm long;

pedicels of female flowers 5-10 mm long; subapical appendage on fruit

up to 2 mm long. 21. B. similis

7. Beyeria apiculata Halford & R.J.F.Hend.

spcies nova B. brevifolia (Miill.Arg.) Benth.

maxime affinis sed foliis sessilibus vel

breviter petiolatis petiolis usque ad 0.5 mm

longis non petiolis 1-2 mm longis, folii

lamina plerumque longiore, 10-40 mm longa

non 5-15 mm longa, et apiculato ad apicem,

flore mare staminibus 9—11 non c. 30 et flore

femineo ovario 3-loculato non 2-loculato

distinguenda est. Typus: Western Australia.

O’Briens Lookout, c. 12 km NW of Wongan

Hills, 14 September 1988, R.J.F.Henderson

H3157 (holo: BRI; iso: MEL, PERTH, MO,

distribuendi).

Monoecious, spreading shrubs to 1.5 m high,

not resinous. Young branchlets pale green

and angular becoming reddish brown and

terete with age, glabrous; older branchlets

with grey to black shallowly fissured bark.

Leaves sessile or shortly petiolate; petioles

up to 0.5 mm long, glabrous; blades linear,

10-40 mm long, 1-1.3 mm wide, length:

width ratio 10-30:1; adaxial surface glabrous

and ± smooth; abaxial surface hairy with

± sessile, stellate hairs c. 0.1mm across,

base cuneate; margins revolute to midrib

concealing abaxial leaf surface; apex obtuse

to acute, ultimately apiculate with extension

from midrib; apiculum slender, 0.3-0.6 mm

long, recurved; midvein obscure adaxially,

abaxially raised, flattened and glabrous on

abaxial face; secondary and tertiary veins

obscure; marginal glands rarely present on

598

blade, up to 1 mm from blade base, 1 per

side of midrib, sessile, smooth, c. 0.1 mm

across. Flowers pedicellate, axillary, solitary

or rarely 2 per axil; bracts ± triangular or

narrow-oblong, 0.4-1.5 mm long, acute at

apex, ± glabrous; pedicels ± glabrous except

for scattered minute erect hairs, stouter and

longer on female flowers than on male flowers;

calyx lobes 5, green, suborbicular, glabrous,

concave adaxially, gibbose abaxially, the

margins erose, rounded at apex; petals slightly

shorter than or equal to calyx lobes, erect,

suborbicular or depressed obovate, glabrous

abaxially, sparsely to moderately villose

adaxially, the margins erose distally; disc

obscure or absent. Male flowers with pedicels

2- 4 mm long; calyx lobes 1-1.3 mm long,

1-1.1 mm wide, surrounding androecium at

anthesis; petals 0.5-0.6 mm long, 0.5-0.6 mm

wide; receptacle 0.8-1 mm across, minutely

hairy; stamens 9-11; filaments erect, 0.1-0.3

mm long, glabrous, bifid distally; anthers 0.2-

0.3 mm long. Female flowers with pedicels

5-12 mm long; calyx lobes ± appressed to

and enclosing gynoecium apart from stigma,

1-1.3 mm long, 0.8-1.1 mm wide; petals

marcescent, 1-1.3 mm long, 0.8-1 mm wide;

ovary ellipsoid, trilobate, 0.6-1 mm long,

glabrous, 3-locular; style ± obsolete; stigma

± discoid, 0.4-0.5 mm across, glabrous,

with margins entire. Fruits subellipsoid,

3- 4 mm long, 3.4-4 mm across, 1-3-seeded,

glabrous, smooth; persistent calyx c. one third

the length of mature fruit. Seeds ellipsoid,

dorsi-ventrally compressed, 3.5-37 mm long

(including caruncle), 1.9-2.1 mm across, 1.4-

1.6 mm in thickness; testa light to dark brown;

caruncle 0.6-0.8 mm long, 0.8-1.3 mm wide,

light brown. Fig. 2.

Additional specimens examined : Western Australia. 6

miles [c. 10 km] from Three Springs, towards Arrino,

Sep 1968, Phillips WA/68 955 (CANB); TV Translator

Hill, 13.5 km NE of township of Wongan Hills on

Piawaning road, Oct 1984, KenneaUy 9344 (CANB,

PERTH); E edge of the Wongan Hills, c. 15 km N of the

township of Wongan Hills, Aug 1980, KenneaUy 7409

(PERTH).

Distribution and habitat : Beyeria apiculata

occurs in south-western Western Australia

where it is known from near Wongan Hills

and Three Springs (Map 8). It is recorded as

growing in a Casuarina/Eucalyptus woodland

Austrobaileya 7 ( 4 ): 577-639 ( 2008 )

community on shallow stony red loamy soil

on a lateritic rise.

Phenology : Flowers have been collected

from August to October, fruits in August and

September.

Affinities : B. apiculata is morphologically

most similar to B. brevifolia (Mull.Arg.)

Benth. but can be distinguished by its more or

less sessile to shortly petiolate leaves (petioles

up to 0.5 mm long compared with petioles 1-2

mm long), generally longer leaf blades (10-

40 mm long compared with 5-15 mm long),

its apiculate leaf apex (rather than obtuse to

rounded in B. brevifolia ), fewer number of

stamens per male flower (9-11 compared with

c. 30) and its female flowers with a 3-locular

rather than a 2-locular ovary.

Etymology : The specific epithet is from Latin

apiculatus , ending abruptly in a short point,

and refers to the apiculate apex of the leaf

blades of this species.

8. Beyeria brevifolia (Mull.Arg.) Benth.,

FI. Austral. 6: 67-68 (1873); Beyeriopsis

brevifolia Mull.Arg., Linnaea 34: 58 (1865);

Beyeria brevifolia (Mull.Arg.) Benth. var.

brevifolia , Airy Shaw, Kew Bull. 26: 69 (1971).

Type: [Western Australia.] Swan River, s.d .,

[J.] Drummond ser. 4. n. 215 [holo: G-DC n.v.

(microfiche IDC 800-73. 2454: II. 5]; iso: K,

PERTH).

Monoecious, erect, virgate shrubs to 1.8 m

high, not resinous. Young branchlets reddish

brown, ± angular, glabrous or rarely sparsely

to moderately hairy and glabrescent; hairs

simple, erect to ascending, c. 0.1 mm long;

older branchlets terete, with grey shallowly

fissured bark. Leaves petiolate; petioles 1-2

mm long, glabrous; blades narrow-oblong

to linear, 5-15 mm long, 0.8-1.6 mm wide,

length:width ratio 6-12:1; adaxial surface

glabrous and ± smooth; abaxial surface

hairy with ± sessile, stellate hairs <0.1 mm

across; base cuneate; margins recurved or

revolute to midrib concealing abaxial leaf

surface; apex obtuse to rounded, straight or

slightly recurved; midvein obscure adaxially,

abaxially raised, flattened and glabrous on

abaxial face; secondary and tertiary veins

obscure; marginal glands absent. Flowers

pedicellate, axillary or terminal on short

Halford & Henderson, Revision of Beyeria

599

Fig. 2. Beyeria apiculata. A. branchlet with male and female flowers x 2. B. side view of leaf apex x 12. C. abaxial

view of leaf apex x 12. D. transverse section of leaf x 24. E. side view of male flower x 15. F. face view of male flower

x 15. G. side view of female flower x 12. H. face view of female flower x 12.1. adaxial view of calyx lobe from female

flower x 24. J. side view of calyx lobe from female flower x 24. K. transverse section of stigma and distal half of ovary

x 24. L. face view of stigma x 24. M. side view of fruit x 8. N. face view of fruit x 8. O. abaxial view of seed x 10. P.

side view of seed x 10. A-P from Henderson H3157 (BRI). Del. W. Smith.

600

axillary branchlets, solitary; bracts ±

triangular, up to 0.4 mm long, acute at apex, ±

glabrous; pedicels ± glabrous, slightly stouter

on female flowers than on male flowers; calyx

lobes 5, yellowish green, suborbicular to very

broad-ovate, glabrous, concavo-convex, the

margins erose, rounded to broad-obtuse at

apex; petals slightly shorter than calyx lobes,

erect, depressed obovate, glabrous abaxially,

sparsely to moderately villose adaxially

distally, the margins erose distally; disc of

5 discrete glands; glands fleshy, glabrous.

Male flowers with pedicels 5-15 mm long;

calyx lobes 0.8-1 mm long, 0.9-1.1 mm wide,

surrounding androecium at anthesis; petals

0.6-0.9 mm long, 0.5-1.4 mm wide; disc

glands c. 0.3 mm long, ±truncate or lobed

distally, canaliculate adaxially; receptacle

1-1.3 mm across, glabrous; stamens 30;

filaments erect, 0.1-0.3 mm long, glabrous,

bifid distally; anthers c. 0.3 mm long. Female

flowers with pedicels 7-13 mm long; calyx

lobes ± appressed to and enclosing gynoecium

apart from stigma, 0.9-1.3 mm long, 0.8-1

mm wide; petals marcescent, 0.6-1 mm long,

0.6-1 mm wide; disc glands 0.4-0.6 mm

long, concavo-convex, rounded, truncate or

shallowly lobed at apex; ovary ellipsoid, 0.9-

1.3 mm long, glabrous, 2-locular; style c. 0.1

mm long, glabrous; stigma calyptriform, 0.9-

1.4 mm across, shallowly umbilicate, glabrous,

with margins entire. Fruits broad-ovoid or

obliquely ellipsoid, laterally compressed,

5-6.7 mm long, 3.4-6 mm across, 1- or 2-

seeded, glabrous, smooth; persistent calyx

c. one fifth the length of mature fruit. Seeds

ellipsoid, dorsi-ventrally compressed, c. 5.2

mm long (including caruncle), 3.2 mm across

and 2.7 mm in thickness; testa light to dark

brown, shiny; caruncle c. 1.7 mm long and 2

mm wide, yellowish-brown.

Additional selected specimens examined: Western

Australia. 5 km N of Nyabing, Sep 1979, Crisp 6145

et al. (BRI); NW slopes of Mt Short, c. 14 km NW of

Ravensthorpe, Sep 1988, Henderson H3188 (BRI); near

NW base of Mt Short, c. 14 km NW of Ravensthorpe,

Sep 1988, Henderson H3189 (BRI); Mt Short, Sep 1968,

Wilson 7980 (PERTH); Ravensthorpe Shire, Hamersley

Inlet Reserve, track to mouth of Hamersley Inlet, c. 500

m S of turnoff to CALM campsite, Aug 2001, Hislop

2286 (BRI, PERTH); Mt Desmond, Aug 1964, Gardner

14074 (PERTH); E side of Mt Desmond, Apr 1962,

George 3661 (PERTH); SW side of Mt Desmond, c. 10

km ESE of Ravensthorpe, Sep 1988, Henderson H3191

Austrobaileya 7(4): 577-639 (2008)

(BRI); 11 km ESE of Ravensthorpe, Oct 1966, Muir 4217

(MEL); 8 km S of Ravensthorpe on road to Hopetoun,

Aug 1968, Wilson 7075 (PERTH); Mt Short, 10 miles

[c. 16 km] N of Ravensthorpe, Nov 1968, Wrigley s.n.

(CANB [CBG036514], PERTH); 77.5 miles [c. 124.7 km]

N of Albany on Jerramungup road, Jan 1975, Keighery

357 (PERTH); 2 miles [c. 3.2 km] from coast, E of

Fitzgerald Inlet, Sep 1970, Aplin 3706 (BRI, PERTH);

Fitzgerald River, May 1964, Gardner 14765 (PERTH);

Mt Maxwell (Fitzgerald River N.P.), Sep 1985, Newbey

10966 (BRI, PERTH).

Distribution and habitat: Beyeria brevifolia

occurs in south-western Western Australia

where it is known from near Jerramungup,

north-eastwards to Mt Desmond near

Ravensthorpe, with disjunct populations near

Harrismith and Nyabing (Map 9). It grows in

mallee communities with heath understorey,

on gravelly, sandy loam or stony clay soils on

hillsides and undulating sandy plains.

Phenology : Flowers have been collected

in January, April, May and from August to

November, fruits from August to October.

Affinities: Beyeria brevifolia is

morphologically most similar to B. apiculata.

For features distinguishing B. brevifolia from

B. apiculata , refer to the Affinities’ section

under that species.

Notes: The collections from the disjunct

populations at Harrismith and Nyabing ( Frizell

& Morrison (PERTH) from Harrismith and

Crisp 6145 et al. (BRI) from Nyabing) differ

from the other collections of this species

examined by having a sparse to moderately

dense indumentum of simple hairs on the

young branchlets. Further collections and

field studies are warranted to establish the

significance of this difference.

9. Beyeria calycina Airy Shaw, Kew Bull. 26:

70 (1971); Beyeria calycina Airy Shaw var.

calycina , Airy Shaw, Kew Bull. 26: 70 (1971).

Type: Western Australia. Coolgardie District:

17 miles [ c. 27 km] W of Lake King turnoff

from Coolgardie - Esperance road, near

granite outcrop, October 1963, P.R.Jefferies

631018 (holo: PERTH; iso: PERTH, K).

Monoecious, spreading, much-branched

shrubs to 1 (-1.5) m high, resinous on most

parts. Young branchlets yellowish green, ±

angular, longitudinally grooved, glabrous,

thickly resinous; older branchlets terete, with

Halford & Henderson, Revision of Beyeria

grey to black shallowly fissured bark. Leaves

petiolate; petioles 0.8-2 mm long, glabrous,

minutely papillose (usually obscured by

resinous covering); blades linear, 15-35 mm

long, 2-3 mm wide, length:width ratio 7-12:1;

adaxial surface ± glabrous, minutely papillose

(obscured by thick resinous covering);

abaxial surface densely hairy with ± sessile,

stellate hairs c. 1 mm across; base cuneate

or attenuate; margins recurved to midrib

concealing abaxial leaf surface; apex obtuse

to acute, ultimately apiculate with extension

from midrib; apiculum stout, c. 0.3 mm long,

bent upward; midvein slightly impressed

adaxially, abaxially raised, flattened, glabrous

and resinous on abaxial face; secondary

and tertiary veins obscure; marginal glands

present on blade, ± regularly spaced from base

to apex of blade, up to 12 per side of midrib,

sessile, smooth, c. 0.1 mm across. Flowers

pedicellate, axillary, solitary; bracts ± ovate,

3-4 mm long, obtuse at apex, glabrous

abaxially, stellate hairy adaxially; pedicels

± glabrous except for scattered minute erect

hairs proximally, stouter on female flowers

than on male flowers; calyx lobes 5, green,

glabrous, rounded to obtuse at apex; petals ±

equal in length to calyx lobes, erect, obovate,

sparsely to moderately villose on both surfaces

proximally, the margins erose distally; disc of

5 discrete glands; glands fleshy, c. 0.3 mm

long, glabrous, dorsi-ventrally compressed.

Male flowers with pedicels 3-4 mm long;

calyx lobes suborbicular or very broad-

ovate, 1.2-1.5 mm long, 1.5-2.5 mm wide,

surrounding androecium at anthesis; petals c.

1.5 mm long and 1 mm wide; receptacle c. 1

mm across, minutely hairy; stamens 15-20;

filaments erect, 0.2-0.6 mm long, glabrous,

entire; anthers c. 0.5 mm long. Female

flowers with pedicels c. 5 mm long; calyx

lobes accrescent, appressed to and enclosing

gynoecium apart from stigma, broad-obovate,

2-3.5 mm long (up to 7 mm long in fruit),

2.5-3 mm wide; petals marcescent, 2-2.5

mm long, 1.5-2 mm wide; ovary ellipsoid,

trilobate, c. 1.5 mm long, glabrous, minutely

papillose, 3-locular; style 0.6-1 mm long,

glabrous; stigma discoid, c. 0.3 mm across, the

same width as style, glabrous, with margins

entire. Fruits broad-ellipsoid, c. 4.5 mm

long, 4-4.5 mm across, 1-3-seeded, glabrous,

601

minutely papillose; persistent calyx longer

than length of mature fruit. Seeds ellipsoid,

dorsi-ventrally compressed, 3.5-4.2 mm

long (including caruncle), 2-2.5 mm across,

1.5-1.8 mm in thickness; testa light to dark

brown; caruncle 0.5-0.9 mm long, 1-1.4 mm

wide, creamy-white.

Additional selected specimens examined: Western

Australia, c. 27 km SW of Daniell, between Norseman

and Salmon Gums, on road to Lake King, Sep 1988,

Henderson H3177 (BRI); 21 km W of Norseman

- Esperance road, along Frank Hann road, Nov 1994,

Cranfield 9389 (BRI, PERTH); 14 miles [c. 22 km] W of

junction of Norseman - Esperance and Lake King roads.

May 1964, Jefferies 640511 (PERTH); [without locality,]

Oct 1961, D.E. White s.n. (PERTH).

Distribution and habitat : Beyeria calycina

occurs in south-western Western Australia

where it is known from between Norseman

and Salmon Gums (Map 10). It grows in

Casuarina shrubland with scattered mallee

eucalypts and a shrubby layer on red brown

sandy loam on undulating plains.

Phenology : Flowers have been collected in

September and October, fruits in October.

Affinities : Beyeria calycina , B. disciformis

Halford & R.J.F.Hend. and B. minor (Airy

Shaw) Halford & R.J.F.Hend. are the only

Beyeria species that have accrescent calyces.

For features distinguishing B. calycina from

those species, refer to the Affinities’ section

under the species concerned.

10. Beyeria cinerea (Mull.Arg.) Benth., FI.

Austral. 6: 66 (1873); Beyeriopsis cinerea

Miill.Arg., Linnaea 34: 57 (1865). Type:

[Western Australia.] in western New Holland

at Swan River, s.d., [J.\ Drummond 724 (holo:

G-DC n.v. [microfiche IDC 800-73. 2454: II.

2]; iso: W n.v. [photo at BRI], K).

Dioecious or sometimes monoecious, open,

spreading, erect or prostrate shrubs to 0.7

m high, not resinous. Young branchlets of

unknown colour when fresh, terete, sparsely

to moderately hairy, glabrescent with

age; hairs stellate, stipitate, up to 0.8 mm

across; older branchlets with greyish white,

shallowly fissured bark. Leaves petiolate;

petioles 0.5-2.7 mm long, moderately hairy

with indumentum as for branchlets; blades

narrow-ovate to broad-ovate, narrow-elliptic

or triangular, 3-17 mm long, 1.2-5 mm wide,

602

length:width ratio 2-4:1; adaxial surface

sparsely hairy with stipitate stellate hairs c.

0.7 mm across, becoming glabrous with age

though remaining sparsely tuberculate by

persistent hair bases; abaxial surface densely

hairy with sessile and stipitate stellate hairs

up to 0.4 mm across; base cordate, cuneate

to truncate; margin recurved or revolute

sometimes to midrib concealing abaxial leaf

surface; apex obtuse or acute; midvein obscure

or faintly impressed adaxially, abaxially

raised, moderately hairy with stellate hairs;

secondary and tertiary venation obscure;

marginal glands absent. Flowers pedicellate,

axillary, males fasciculate in 2- or 3-flowered

fascicles or solitary, females solitary; bracts

ovate or oblong, 0.4-0.6 mm long, obtuse

at tip, glabrous or with a few stellate hairs;

pedicels slender, 1-3.5 (-7) mm long,

glabrous or with a few stellate hairs; calyx

lobes 5, yellow green to light green, concave

and glabrous adaxially, gibbose with a few

stipitate stellate hairs abaxially, the margins

erose, obtuse to rounded at apex; petals slightly

shorter than or equal to calyx lobes, erect,

the margins erose; disc of 5 discrete glands;

glands fleshy, 0.2-0.3 mm long, laterally

compressed, glabrous, truncate to rounded

or irregularly lobed. Male flowers with

calyx lobes suborbicular to depressed ovate,

1-1.3 mm long, 1-1.6 mm wide, surrounding

androecium at anthesis; petals suborbicular or

depressed obovate, 0.5-0.8 mm long, 0.7-1

mm wide, glabrous abaxially, with scattered

stellate hairs adaxially proximally; receptacle

c. 1 mm across, stellate-tomentose; stamens

20-40; filaments erect, 0.2-0.5 mm long,

glabrous, bifid distally; anthers 0.2-0.3 mm

long. Female flowers with calyx lobes ±

appressed to and enclosing gynoecium apart

from stigma, suborbicular or broad-ovate,

0.7-1.1 mm long, 0.7-0.8 mm wide; petals

suborbicular ortransverse-elliptic,0.3-0.6mm

long, 0.3-0.9 mm wide, glabrous abaxially,

stellate-tomentose adaxially distally; ovary

subglobose, 0.5-0.7 mm long, stellate-

Leaf blades with truncate to cuneate bases

Leaf blades with cordate bases.

Austrobaileya 7(4): 577-639 (2008)

pubescent or glabrous, mostly 1-locular by

abortion, rarely 2-locular; style c. 0.2 mm

long, stellate-pubescent; stigma ± discoid,

0.4-0.5 mm wide, glabrous, sulcate adaxially,

with margins entire, slightly recurved. Fruits

obliquely or transverse-pyriform, 4-6 mm

long, 2.5-3.8 mm across, 1- or rarely 2-

seeded, glabrous or with scattered stellate

hairs; persistent calyx c. one fifth the length

of mature fruit. Seeds globose or broad-

ellipsoid, slightly dorsi-ventrally compressed,

2.9-4.5 mm long (including caruncle), 2-2.7

mm wide, 2-2.5 mm in thickness; testa

mottled cream and dark brown; caruncle

0.7-1.7 mm long, 0.8-1.5 mm across, creamy-

white to yellowish-white.

Affinities : Beyeria cinerea is morphologically

most similar to B. lepidopetala F.Muell.

but differs from that in having generally

smaller leaves (3-17 mm long x 1.2-5 mm

wide compared with 15-20 mm long x 1.5-3

mm wide) that are narrow-ovate to broad-

ovate, narrow-elliptic or triangular rather

than linear in outline and shorter pedicels

(1-7 mm long compared with 8-30 mm long).

Beyeria cinerea is also similar to B. latifolia

Baill. but differs from it in having generally

smaller leaves (3-17 mm long x 1.2-5 mm

wide compared with 10-30 mm long x 6-12

mm wide) and smaller fruits (4-6 mm long x

2.5-3.8 mm across compared with 6-6.5 mm

long x 4-5.5 mm across) that are more or less

pyriform and 1-seeded rather than ovoid and

1-3-seeded.

Beyeria cinerea , as circumscribed here,

occurs on the islands and in coastal areas from

Exmouth to Mandurah, Western Australia.

The species exhibits some discontinuous

variation in leaf blade shape associated with

a geographical disjunction. These differences

are considered to be sufficient to warrant

formal recognition of taxa at subspecific rank.

Two subspecies are here formally described

and they can be distinguished using the

following key.

10a. B. cinerea subsp. cinerea

10b. B. cinerea subsp. borealis

Halford & Henderson, Revision of Beyeria

10a. Beyeria cinerea (Miill.Arg.) Benth.

subsp. cinerea

Beyeriopsis cygnorum Miill.Arg., Linnaea

34: 56-57 (1865); Beyeria cygnorum (Mull.

Arg.) Benth., FI. Austral. 6: 66 (1873). Type:

[Western Australia.] Swan River, s.d., [J.\

Drummond 85 (holo: G-DC n.v. (microfiche

IDC 800-73. 2454: I. 8); iso: MEL [2 sheets

114150, 114151], PERTH, K n.v. (photo at

BRI)).

Leaves with petioles 1-2 mm long; leaf blades

narrow-ovate to ovate or narrow-elliptic, 3.5-

17 mm long, 1.2-5 mm wide; base cuneate to

truncate; margins recurved or revolute; apex

obtuse to acute.

Additional selected specimens examined : Western

Australia. Horrocks Beach, Aug 1984, Bates 3919

(AD); 15.8 km N of Leeman - Eneabba road, turnoff

on Dlawong road, Jul 1992, Cranfield & Spencer 8260

(PERTH); Beehive Gully, Leeman, 1982, Foulds 33

(PERTH); 3 miles [c. 5 km] E of Jurien Bay, Sep 1966,

George 7810 (PERTH); Parrot Ridge, Yanchep N.P,

50 km N of Perth, Sep 1989, Keighery 11165 (CANB);

Reabold Hill, Floreat Park, 8 km W [of] Perth, Nov

1987, Keighery 9262 (PERTH); Buckland Hill, Mosman

Park, Sept 1989, Kenneally 11007 (PERTH); The Plains,

Mandurah road, Oct 1967, George 9202 (PERTH);

Madora, Perth to Mandurah, Sep 1983, Keighery s.n.

(PERTH).

Distribution and habitat : Beyeria cinerea

subsp. cinerea occurs in south-western

Western Australia where it is known from

Horrocks Beach, south of Dongara, southward

to near Mandurah (Map 11). It grows in

coastal heath and shrubland communities on

sandy soils over limestone.

Phenology : Flowers have been collected in

July and from September to November, fruits

from September to November.

Notes : The name Beyeria cygnorum which is

here placed in synonymy is listed as Priority

Three under DEC Conservation Codes for

Western Australian Flora (Florabase, http://

florabase.dec.wa.gov.au [accessed June

2008]).

10b. Beyeria cinerea subsp. borealis Halford

& R.J.EHend. subspecies nova a Beyeria

cinerea (Miill.Arg.) Benth. subsp. cinerea

folii lamina generatim minore, 3-5.2 mm

longa x 1.2-4.3 mm lata non 3.5-17 mm longa

x 1.2-5 mm lata, basi laminae folii cordata

603

non truncata ad cuneata et marginibus folii

laminae folii revolutioribus differt. Typus:

Western Australia. N end of Passage Paddock,

Dirk Hartog Island, 2 September 1972,

A.S.George 11384 (holo: PERTH; iso: BRI,

CANB).

Beyeriopsis cyanescens Miill.Arg. in A.DC.,

Prodr. 15(2): 200 (1866); Beyeria cyanescens

(Miill.Arg.) Benth., FI. Austral. 6: 66-67

(1873). Type citation: “In Nova Hollandia, in

Iles-Steriles (hb. Kunth! in hb. berol. ex Mus.

Paris).” n.v.

Illustration : Griining (1913: 74, fig. 12) as

Beyeria cyanescens.

Leaves with petioles 0.5-2.5 mm long; blades

triangular or narrow- to broad-ovate, 3-5.2

mm long, 1.2-4.3 mm wide; base cordate;

margins revolute to midrib usually concealing

abaxial surface of leaf blade; apex obtuse.

Additional selected specimens examined : Western

Australia. E of Pitgrammunne Well on Yardie Creek

Station, Cape Range, Sep 1964, Chadwick 1434 (PERTH);

± 5 miles [c. 8 km] N of Yardie Creek, May 1965, George

s.n. (PERTH); Learmonth road, 44 miles [c. 71 km] S

of Bullara turnoff, Feb 1962, George 3284 (PERTH); 5

miles [c. 8 km] N of Cardabia Station turnoff, Minilya

- Exmouth road, Sep 1970, George 10349 (PERTH);

Bernier Island, Shark Bay, Jul 1988, Morat 8104

(PERTH); Blow Hole, Carnarvon, Aug 1976, Wittwer

W1797 (PERTH); Dorre Island, Jul 1959, Royce 5897 ;

5896 (PERTH); ibid, Aug 1977, Weston 10566 (PERTH);

Quoin Bluff area. Shark Bay, Jun 1974, Kenneally 1345

(PERTH); Red Bluff, c. 3 km S of Kalbarri, May 1968,

Wilson 6507 (PERTH).

Distribution and habitat : Beyeria cinerea

subsp. borealis is confined to coastal areas

from Exmouth southwards to Kalbarri in

Western Australia (Map 12). It grows in low

scrub communities on red sandy soils on

limestone rises.

Phenology : Flowers have been collected in

February and from June to September, fruits

in May, July and September.

Affinities: Beyeria cinerea subsp. borealis

differs from B. cinerea subsp. cinerea by its

generally smaller leaf blades (3-5.2 mm long

x 1.2-4.3 mm wide compared with 3.5-17

mm long x 1.2-5 mm wide) which are cordate

rather than truncate to cuneate at the base, and

its more strongly revolute leaf blade margins.

604

Etymology : The subspecific epithet is from

Latin, borealis , northern, in reference to this

subspecies’ distribution in relation to that of

the other subspecies.

11. Beyeria cockertonii Halford &

R.J.F.Hend., species nova B. brevifoliae

(Miill.Arg.) Benth. maxime affinis sed strato

viscido tenui in partibus maximis, habitu

breviore (usque ad 0.25 m alto non usque ad

1.8 m alto), pedicellis brevioribus (1-3 mm

longis non 5-15 mm longis) et flore femineo

stigmate 3-lobulato non calyptriformi et

ovario 3-loculato non 2-loculato distinguenda

est. Typus: Western Australia. SE of

Ravensthorpe [precise locality withheld for

conservation purposes], 5 September 2006,

D.Halford Q9139 & G.Cockerton (holo:

PERTH; iso: BRI).

Beyeria sp. Bandalup Hill (G.Cockerton

7553), in Florabase, http://florabase.dec.wa.

gov.au [accessed June 2008],

Monoecious, erect shrubs to 0.25 m high,

resinous on most parts. Young branchlets

yellow, ± angular, sparsely hairy; hairs simple,

erect, c. 0.1 mm long; older branchlets terete,

with grey irregularly tessellated bark. Leaves

petiolate; petioles 0.5-1 mm long, glabrous;

blades narrow-oblong to linear, 5-8 mm long,

0.8-1.2 mm wide, length:width ratio 6-9:1;

adaxial surface glabrous and ± smooth;

abaxial surface hairy with stellate hairs up

to 0.5 mm across; base cuneate; margins

recurved to midrib concealing abaxial leaf

surface; apex rounded to truncate, slightly

recurved; midvein obscure adaxially, abaxially

raised, flattened and glabrous on abaxial

face; secondary and tertiary veins obscure;

marginal glands absent. Flowers pedicellate,

axillary, solitary; bracts ± triangular, up

to 0.4 mm long, acute at apex, glabrous;

pedicels ± glabrous or with a few minute hairs

proximally, slightly stouter on female flowers

than on male flowers; calyx lobes 5, yellow,

glabrous, concavo-convex, the margins erose;

petals, erect, the margins erose distally; disc

of 5 discrete glands; glands fleshy, glabrous.

Male flowers with pedicels 1-3 mm long;

calyx lobes suborbicular to very broad-ovate,

c. 1.3 mm long and 1.2 mm wide, surrounding

androecium at anthesis, rounded to broad-

Austrobaileya 7(4): 577-639 (2008)

obtuse at apex; petals slightly shorter than

or equal to calyx lobes, depressed obovate,

c. 0.8 mm long and 0.9 mm wide, glabrous

abaxially, sparsely to moderately villose

adaxially proximally; disc glands c. 0.2 mm

long, ±truncate or lobed distally, canaliculate

adaxially; receptacle 0.8-1.1 mm across,

stellate hairy; stamens 10-16; filaments

erect, c. 0.1 mm long, glabrous, entire or

bifid distally; anthers c. 0.4 mm long. Female

flowers with pedicels 1—1.5 mm long; calyx

lobes ± appressed to and enclosing gynoecium

apart from stigma, ovate to broad-ovate, c.

1.3 mm long and 0.8 mm wide, acute at apex;

petals less than half the length of the calyx

lobes, marcescent(?), broad-obovate, c. 0.4

mm long and 0.4 mm wide; disc glands c. 0.1

mm long, rounded to truncate at apex; ovary

ellipsoid, c. 0.9 mm long, glabrous, 3-locular;

style 0.1-0.3 mm long, glabrous; stigma 3-

lobulate, glabrous; lobes erect, triangular, c.

0.2 mm long, with margins entire. Fruits ±

ellipsoid, 3-4 mm long, 2-3 mm across, 1-3-

seeded, glabrous, smooth; persistent calyx c.

one third the length of mature fruit. Seeds

ellipsoid, dorsi-ventrally compressed, c. 2.5

mm long (including caruncle), 1.5 mm across

and c. 1 mm in thickness; testa dark brown;

caruncle c. 0.5 mm long and 0.7 mm wide,

pale brown. Fig. 3.

Additional specimens examined: Western Australia,

[localities withheld] Feb 2003, Landcare Services

GC7553A (PERTH); Jun 2002, Cockerton GC7553 (BRI,

PERTH).

Distribution and habitat: Beyeria cockertonii

occurs in south-western Western Australia

where it is known only from a single site near

Ravensthorpe (Map 13). It grows in mallee-

heath on shallow rocky clay soils on hill

slopes and hilltops.

Phenology: Flowers have been collected in

June and September.

Affinities: Beyeria cockertonii is

morphologically most similar to B. brevifolia

but can be distinguished by the thin, viscid

covering over most parts, shorter habit (up to

0.25 m high compared with up to 1.8 m high),

shorter pedicels (1-3 mm long compared with

5-15 mm long), and female flowers with a 3-

lobulate rather than calyptriform stigma and a

3-locular rather than 2-locular ovary.

Halford & Henderson, Revision of Beyeria

605

Fig. 3. Beyeria cockertonii. A. branchlet with male flowers x 4. B. branchlet with immature fruit x 4. C. abaxial view

of leaf apex x 12. D. side view of leaf apex x 12. E. transverse section of leaf x 24. F. side view of male flower x 12. G.

face view of male flower x 12. H. side view of female flower x 24.1. face view of female flower x 22. J. adaxial view of

calyx lobe from female flower x 24. K. side view of fruit x 12. L. face view of fruit x 12. M. abaxial view of seed x 16.

N. side view of seed x 16. A-N from Halford Q9139 & Cockerton (BRI). Del. W. Smith.

606

Notes : Beyeria cockertonii is listed as

Declared Rare Flora under the Western

Australian Wildlife Conservation Act 1950,

under the name Beyeria sp. Bandalup Hill

(G.Cockerton 7553) (Florabase, http://

florabase.dec.wa.gov.au [accessed June

2008]).

Etymology : The specific epithet honours

Geoff Cockerton (Managing Director,

Landcare Services Pty Ltd) who discovered

and collected this species and subsequently

brought it to our attention.

12. Beyeria constellata Halford & R. J.F.Hend.

species nova floribus maribus et femineis

calyce lobis manifeste carinatis a speciebus

omnibus ceteris Beyeriae clare distincta.

Typus: Western Australia, c. 4 km E of

Cadoux, 14 September 1988 ,R.J.F.Henderson

H3158 (holo: BRI; iso: K, MEL, PERTH,

distribuendi).

Monoecious, erect much-branched shrubs

to 0.6 m high, resinous on most parts. Young

branchlets white to pale green, ± angular

becoming terete with age, longitudinally

ridged, densely hairy between resinous

ridges; hairs stellate, sessile, 0.2-0.3 mm

across; older branchlets with grey to black

shallowly furrowed bark. Leaves petiolate;

petioles 0.7-1.3 mm long, hairy adaxially,

glabrous abaxially; blades oblong or narrow-

elliptic, 10-20 mm long, 2-5 mm wide, length:

width ratio 3-7:1; adaxial surface glabrous,

minutely papillose (obscured by thick

resinous covering); abaxial surface densely

hairy with ± sessile, stellate hairs up to 0.8

mm across; base abruptly cuneate; margins

recurved to revolute occasionally to midrib

concealing abaxial leaf surface; apex rounded

or sometimes retuse; midvein impressed

adaxially, abaxially prominently raised and

flattened, glabrous and resinous on abaxial

face; secondary and tertiary veins obscure;

marginal glands absent. Flowers pedicellate,

axillary or terminal on short axillary

branchlets, solitary or rarely 2 per axil; bracts

± foliose, oblong, up to 1 mm long, rounded

at apex, ± glabrous adaxially, stellate hairy

abaxially; pedicels slender or stout, sparsely

hairy proximally with minute erect glandular

hairs c. 0.1 mm long (mostly obscured by

Austrobaileya 7(4): 577-639 (2008)

resinous covering); calyx lobes 5, light green

to yellow-green with flushes of red especially

along the margins, suborbicular, concave

adaxially, prominently keeled abaxially,

glabrous, the margins usually erose, obtuse

to rounded at apex; petals slightly shorter

than or equal to calyx lobes, ± erect, glabrous

abaxially, densely villose abaxially; disc of

5 discrete glands; glands fleshy, 0.3-0.4 mm

long, glabrous, rounded to truncate. Male

flowers with pedicels 13-16 mm long; calyx

lobes 1.6-1.8 mm long, 1.7-2.3 mm wide,

surrounding androecium at anthesis; petals

transverse-elliptic, 0.5-0.6 mm long, 0.5-0.6

mm wide, with erose margins; receptacle

2- 2.5 mm across, stellate hairy; stamens

30-60; filaments erect, 1-1.3 mm long,

glabrous, entire or bifid distally; anthers 0.3-

0.4 mm long. Female flowers with pedicels

8-18 mm long; calyx lobes ± appressed

to and enclosing gynoecium apart from

stigma, 1.5-1.8 mm long, 1.6-1.7 mm wide;

petals persistent, broad-ovate, 1.4-1.6 mm

long, 1.6-1.8 mm wide, with entire margins;

ovary subglobose, trigonal, c. 1.2 mm long,

densely hairy proximally, 3-locular; style

0.5-0.7 mm long, glabrous; stigma ± discoid,

3- lobulate, glabrous, with margins entire,

recurved. Fruits subglobose, trilobate, 4-6

mm long, 5-6 mm across, mostly 3-seeded,

hairy proximally, glabrous, papillose distally

(obscured by resinous covering); persistent

calyx c. half the length of mature fruit. Seeds

subglobose, dorsi-ventrally compressed, c. 3.6

mm long (including caruncle), 3.2 mm across

and 2.4 mm in thickness; testa of unknown

colour when fresh; caruncle c. 1 mm long and

1.5 mm wide, creamy-white. Fig. 4.

Additional specimens examined : Western Australia. 5

km W [E] of Cadoux at Johnson road turnoff, Sep 1983,

Purdie 5293 (CANB); Cadoux Pump road, c. 7 km E

of Cadoux on road to Koorda, near Rabbit Proof Fence

road, Sep 2006, Halford Q9145 & Cockerton (BRI,

MEL, PERTH); no.2 Rabbit Fence, E of Manmanning,

Sep 1982, Smith 129 (CANB, MEL, PERTH).

Distribution and habitat : Beyeria constellata

is confined to the Cadoux and Manmanning

districts in south-western Western Australia

(Map 14). It grows in scrub with Eucalyptus

oldfieldii on sand over laterite and in

Eucalyptus woodland on grey sandy soil.

Halford & Henderson, Revision of Beyeria

607

Fig. 4. Beyeria constellata. A. branchlet with female flowers x 2. B. branchlet showing stellate indumentum x 6. C.

side view of leaf apex x 6. D. abaxial view of leaf apex x 6. E. transverse section of leaf x 12. F. side view of male flower

x 6. G. face view of male flower x 6. H. side view of female flower x 6.1. face view of female flower x 6. J. side view of

calyx lobe from female flower x 12. K. side view of female flower with calyx and petals removed, showing disc glands,

ovary and style x 6. L. side view of fruit x 6. M. face view of fruit x 6. N. side view of seed x 10. O. abaxial view of

seed x 10. A-0 from Henderson H3158 (BRI). Del. W.Smith.

608

Phenology : Flowers and fruits have been

collected in September.

Affinities : Beyeria constellata when in flower

or fruit is not easily confused with any other

species of Beyeria. The prominent keel on

the calyx lobes of male and female flowers

clearly distinguishes it from all other Beyeria

species.

Etymology : The specific epithet is from Latin,

constellatus , studded with stars, in reference

to the conspicuous star-like appearance of the

flowers of this species.

13. Beyeria disciformis Halford &

R.J.F.Hend. species nova B. minori (Airy

Shaw) Halford & R.J.F.Hend. maxime affinis

nee non B. calycinae Airy Shaw affinis. Ab

ilia foliis laminis longioribus et proportione

angustioribus, 10-16 mm longis x 1.3-2.3

mm latis non 5-10 mm longis x 1.6-2.5 mm

latis, angusti-oblongis usque ad linearis non

angusti-oblongis usque ad angusti-ellipticis,

floribus femineis stigmate minore, 0.1-0.3

mm lato non 0.7-0.9 mm lato, pedicellis

glandulosis pubescentibus non glabris

et bracteis ovatis et dense pubescentibus

adaxialiter non angusti-oblongis et plene

glabris differt. Ab hac foliorum laminis et

petiolis et bracteis brevioribus et floribus

et fructibus plerumque parvioribus differt.

Typus: Western Australia, c. 2 km from

Buntine along road to Wubin, 13 September

1988, R.J.F.Henderson H3155 (holo: BRI; iso:

CANB, K, MEL, PERTH, distribuendi).

Monoecious, spreading, much-branched

shrubs to 1 m high, resinous on most parts.

Young branchlets pale green, ± angular

becoming terete with age, longitudinally

grooved, glabrous; older branchlets with

grey to black shallowly fissured bark. Leaves

petiolate; petioles 0.5-1 mm long, glabrous,

minutely papillose (usually obscured by

resinous covering); blades narrow-oblong

to linear, 10-16 mm long, 1.3-2.3 mm

wide, length:width ratio 6-8:1, recurved

distally; adaxial surface glabrous, minutely

papillose (usually obscured by resinous

covering); abaxial surface densely hairy with

± sessile, stellate hairs up to 0.6 mm across;

base cuneate; margins recurved to midrib

concealing abaxial leaf surface; apex rounded

Austrobaileya 7(4): 577-639 (2008)

or obtuse, ultimately apiculate with extension

from midrib; apiculum stout, 0.3-0.6 mm

long, recurved; midvein slightly raised or

impressed adaxially, abaxially prominently

raised and flattened, glabrous and resinous

on abaxial face; secondary and tertiary veins

obscure; marginal glands occasionally present

on blade, up to 2 mm from base, 1 per side of

midrib, sessile, smooth, up to 0.2 mm across.

Flowers pedicellate, axillary, solitary; bracts

ovate, 0.5-1.3 mm long, cymbiform, acute at

apex, glabrous abaxially, densely hairy with

weakly ascending, simple hairs up to 0.3 mm

long adaxially; pedicels slender, sparsely

hairy proximally with minute erect glandular

hairs c. 0.05 mm long; calyx lobes 5, pale

green, suborbicular to very broad-ovate,

glabrous, concavo-convex, the margins entire

or erose, obtuse to rounded at apex; petals

slightly shorter than or equal to calyx lobes,

concavo-convex, the margins erose; disc

obscure or absent. Male flowers with pedicels

2-4 mm long; calyx lobes 0.8-1.3 mm long,

1.4- 1.7 mm wide, surrounding androecium at

anthesis; petals suborbicular, 0.7-1.1 mm long,

0.8-1.4 mm wide, glabrous abaxially, densely

villose adaxially proximally; receptacle 0.9-

1.1 mm across, minutely hairy; stamens 15;

filaments erect, 0.2-0.4 mm long, glabrous,

entire or bifid distally; anthers 0.2-0.3 mm

long. Female flowers with pedicels 2-6 mm

long; calyx lobes accrescent, ± appressed to

and enclosing gynoecium apart from stigma,

1.4- 2 mm long (2-3 mm long in fruit), 1.7-2.4

mm wide; petals marcescent, suborbicular

to broad-obovate, 0.8-1.5 mm long, 1-1.5

mm wide, glabrous abaxially, densely villose

adaxially; ovary subglobose, trigonal, 0.9-1.1

mm long, glabrous, 3-locular; style c. 0.1 mm

long, glabrous; stigma discoid, 0.1-0.3 mm

across, glabrous, with margins entire. Fruits

subglobose, 3.5-4 mm long, 3-4 mm across,

2- or 3-seeded, glabrous, smooth; persistent

calyx one half to two thirds the length of

mature fruit. Seed not seen. Fig. 5.

Additional specimens examined: Western Australia.

2 miles [c. 3 km] N of Wubin, Sep 1966, Knox 660902

(PERTH); Wubin, Oct 1965, Knox 651043 (PERTH).

Distribution and habitat: Beyeria disciformis

is known only from the vicinity of Wubin in

south-western Western Australia (Map 15).

Halford & Henderson, Revision of Beyeria

609

Fig. 5. Beyeria disciformis. A. branchlet with male and female flowers x 2. B. branchlet showing longitudinal grooves

x 6. C. adaxial view of leaf x 4. D. side view of leaf apex x 12. E. abaxial view of leaf apex x 12. F. transverse section of

leaf x 24. G. side view of male flower with bracts x 12. H. face view of male flower x 12.1. side view of female flower

with bracts x 12. J. face view of female flower x 12. K. side view of calyx lobe from female flower x 12. L. side view

of fruit x 6. M. face view of fruit x 6. A-M from Henderson H3155 (BRI). Del. W. Smith.

610

Austrobaileya 7(4): 577-639 (2008)

It grows on hard red loamy soil in remnant

Acacia/Hakea/Eucalyptus shrubland.

Phenology : Flowers have been collected in

September and October, fruits in October.

Affinities : Beyeria disciformis is similar to

B. minor (Airy Shaw) Halford & R.J.F.Hend.

and B. calycina. It differs from the former in

having longer and proportionally narrower

leaf blades, a different leaf blade shape,

Table 1. Comparison of morphological characters for Beyeria disciformis , B. minor and

B. calycina

Character

B. disciformis

B. minor

B. calycina

leaf shape

narrow-oblong to linear

narrow-oblong to

narrow-elliptic

linear

leaf dimensions (mm)

10-16 x l.3-2.3

5-10 x 1.6-2.5

15-35 x 2-3

petiole length (mm)

0.5-1

0.6-1.2

0.8-2

stigma width (mm)

0.1-0.3

0.7-0.9

c. 0.3

pedicel

glandular hairy

proximally

glabrous

with scattered

hairs proximally

bract shape

ovate

narrow-oblong

ovate

bract dimensions (mm)

0.5-1

0.6-1.1

3-4

bract adaxial surface

hairy

glabrous

hairy

calyx lobe length (mm)

in male flower

0.8-1.3

c. 1.5

1.2-1.5

in female flower

1.4-2

c. 2.5

2-3.5

petal length (mm)

in male flower

0.7-1.1

c. 1

c. 1.5

in female flower

0.8-1.5

0.9-1

2-2.5

fruit dimensions (mm)

3.5-4 x 3-4

4.5-5.5 x 4-5

c. 4.5 x 4-4.5

smaller stigmas, glandular hairy pedicels

and ovate bracts which are densely hairy

adaxially compared with narrow-oblong and

wholly glabrous bracts in B. minor. Beyeria

disciformis differs from B. calycina in having

generally shorter leaf blades, petioles and

bracts and generally smaller flowers and

fruits. These differences are summarized in

Table 1

Etymology : The specific epithet is from Latin

disciformis , disk-like, and refers to the shape

of the stigma in this species.

14. Beyeria gardneri Airy Shaw, Kew Bull.

26: 68-69 (1971). Type: Western Australia.

Irwin District: Murchison River, 30 August

1931, C.A.Gardner 2588 (holo: PERTH; iso:

PERTH).

Monoecious, slender, open shrubs to 0.7 m

high, usually resinous on most parts. Young

branchlets of unknown colour when fresh,

terete, glabrous, tuberculate; older branchlets

with greyish white shallowly fissured bark.

Leaves petiolate; petioles 0.8-2 mm long,

glabrous, minutely papillose (usually obscured

by resinous covering); blades narrow-oblong

to linear, 5-16 mm long, 0.8-1.4 mm wide,

length:width ratio 6-12:1; adaxial surface

glabrous, sparsely tuberculate; abaxial

surface densely hairy with hairs < 0.05 mm

long; base cuneate; margins recurved to

midrib concealing abaxial leaf surface; apex

obtuse to rounded, recurved; midvein obscure

adaxially, abaxially raised and rounded,

glabrous; secondary and tertiary veins

obscure; marginal glands absent. Flowers

pedicellate, axillary, solitary or fasciculate

Halford & Henderson, Revision of Beyeria

in 2- or 3-flowered fascicles; bracts ovate,

up to 0.8 mm long, acute at apex, ± glabrous;

pedicels ± glabrous, stouter on female

flowers than on male flowers; calyx lobes 5,

yellowish green, suborbicular to very broad-

ovate, glabrous, concave adaxially, gibbose

abaxially, the margins entire, rounded to

obtuse at apex; petals slightly shorter than

or equal to calyx lobes, erect, suborbicular

or very broad-ovate, glabrous abaxially,

sparsely to moderately villose adaxially, the

margins ± entire; disc of 5 discrete glands;

glands thin, ± oblong, 0.3-0.6, dorsi-ventrally

compressed, glabrous, acute to rounded. Male

flowers with pedicels 1-2 mm long; calyx

lobes 0.5-0.9 mm long, 0.9-1.3 mm wide,

surrounding androecium at anthesis; petals

0.5-0.9 mm long, 0.9-1 mm wide; receptacle

c. 0.8 mm across, minutely hairy; stamens

10-12; filaments erect, 0.1-0.4 mm long,

glabrous, bifid distally; anthers 0.2-0.3 mm

long. Female flowers with pedicels 1-3 mm

long; calyx lobes ± appressed to and enclosing

gynoecium apart from stigma, 0.9-1.3 mm

long, 1.3-1.4 mm wide; petals marcescent,

c. 1.3 mm long and 1.1 mm wide; ovary

ellipsoid, ± bilobate, c. 1 mm long, glabrous,

2-locular, with 4 short subapical appendages;

style ± obsolete; stigma discoid, 0.4-0.5

mm across, glabrous, with margins entire.

Fruits subellipsoid, ± laterally compressed

when 2-seeded, 4-5.5 mm long, 4.5-5.5 mm

across, with 2 or rarely 4 horn-like subapical

appendages, 1- or rarely 2-seeded, glabrous,

smooth; persistent calyx c. one fifth the

length of mature fruit; subapical appendages

up to 1 mm long. Seeds subglobose, slightly

dorsi-ventrally compressed, c. 4 mm long

611

(including caruncle), 2.8-3 mm across, 2.7-3

mm in thickness; testa light to dark brown;

caruncle c. 1.2 mm long and 0.7 mm wide,

creamy-white.

Additional specimens examined: Western Australia,

[localities withheld] Aug 1961, Gardner 13318 (PERTH);

Aug 1931, Gardner2588 (PERTH); Aug 1931, Gardner &

Blackall 598 (PERTH); Sep 1983, Purdie 5220 (CANB);

Oct 1963, Chadwick 1672 (PERTH); Sep 1970, Chapman

1292 (PERTH); Sep 1991, Cranfield & Spencer 8068

(CANB); Sep 1977, Hnatiuk 771198 (PERTH); Sep 1988,

Griffin 5229 (PERTH); Oct 1971, Royce 9615 (PERTH).

Distribution and habitat : Beyeria gardneri

is confined to subcoastal areas from Kalbarri

southwards to Badgingarra and Watheroo in

south-western Western Australia (Map 16).

It grows in open heathland and shrubland

communities on sandy soils mostly on

undulating sand plains.

Phenology : Flowers and fruits have been

collected from August to October.

Affinities : Beyeria gardneri is

morphologically most similar to B. similis

(Miill.Arg.) Benth. and both these species

have fruits with subapical appendages which

distinguishes them from other species of

Beyeria. Beyeria gardneri differs from

B. similis in having generally smaller leaf

blades, shorter pedicels and shorter subapical

appendages on its fruits. These differences

are summarized in Table 2.

Notes : Beyeria gardneri is listed as Priority

One under DEC Conservation Codes for

Western Australian Flora (Florabase, http://

florabase.dec.wa.gov.au [accessed June 2008]).

Table 2. Comparison of morphological characters for Beyeria gardneri and B. similis

Character

B. gardneri

B. similis

leaf dimensions (mm)

5-16x0.8-1.4

20-50 x 1.6-2.5

pedicel length (mm)

of male flowers

1-2

2-3

of female flowers

1-3

5-10

fruit: subapical appendage length (mm)

up to 1

up to 2

612

15. Beyeria lapidicola Halford & R. J.F.Hend.

species nova B. disciformi Halford &

R. J.F.Hend. et B. minori (Airy Shaw) Halford

& R. J.F.Hend. maxime affinis. Ab ilia floribus

pedicello 7-13 mm longo non 2-6 mm longo

et bracteis oblongis, plusminusve foliosis et

adaxialiter glabris non ovatis cymbiformibus

et adaxialiter dense pubescentibus differt. Ab

hac calycis lobis ovatis usque lato-ovatis non

suborbicularibus vel lato-ovatissimis, floribus

femineis stigmate 0.4-0.6 mm lato non 0.7-

0.9 mm lato, folii apice obtuso non obtuso-

apiculato et floribus femineis fructiferis

calycis lobis non auctis differt. Typus:

Western Australia, near Wiluna [precise

locality withheld for conservation purposes],

18 August 2006, A.Markey & S.Dillon 4114

(holo: PERTH).

Beyeria sp. Murchison (B. Jeanes s.n.

7/7/2005), in Florabase, http://florabase.dec.

wa.gov.au [accessed June 2008],

Monoecious, spreading, much-branched

shrubs to 1 m high, resinous on most parts.

Young branchlets pale yellowish green, ±

angular becoming terete with age, sometimes

shallowly longitudinally grooved, glabrous;

older branchlets with grey to black shallowly

fissured bark. Leaves petiolate; petioles

1-1.6 mm long, glabrous, minutely papillose

(usually obscured by resinous covering);

blades narrow-oblong, 8-16 mm long,

1.4-2 mm wide; adaxial surface glabrous,

minutely papillose (usually obscured by

resinous covering); abaxial surface densely

hairy with ± sessile, stellate hairs up to 0.3

mm across; base cuneate; margins recurved

to midrib usually concealing abaxial leaf

surface; apex rounded, ultimately apiculate

(usually obscured by resinous covering) with

extension from midrib; apiculum stout, up

to 0.1 mm long; midvein slightly impressed

adaxially, abaxially prominently raised

and flattened, glabrous and resinous on

abaxial face; secondary and tertiary veins

obscure; marginal glands absent. Flowers

pedicellate, axillary or terminal on short

axillary branchlets, solitary; bracts oblong, ±

foliose, up to 2.5 mm long, rounded at apex,

glabrous adaxially, stellate hairy abaxially;

pedicels slender, papillose (mostly obscured

by resinous covering), stouter on female

Austrobaileya 7(4): 577-639 (2008)

flowers than on male flowers; calyx lobes 5,

pale green, ovate to broad-ovate, glabrous,

concavo-convex, the margins entire or erose,

obtuse to rounded at apex; petals slightly

shorter than calyx lobes, ± flat, the margins

entire; disc of 5 discrete glands; glands fleshy,

0.2-0.3 mm long, laterally compressed,

glabrous, truncate. Male flowers with pedicels

7-11 mm long; calyx lobes 1.2-1.4 mm long,

1.4-1.7 mm wide, surrounding androecium at

anthesis; petals orbicular, c. 1 mm long and 1

mm wide, glabrous abaxially, sparsely villose

adaxially; receptacle c. 1 mm across, minutely

hairy; stamens 20; filaments erect, 0.2-0.4

mm long, glabrous, entire or bifid distally;

anthers 0.2-0.3 mm long. Female flowers

with pedicels 7-13 mm long; calyx lobes ±

appressed to and enclosing gynoecium apart

from stigma, 1.1-2 mm long, 0.9-1.5 mm

wide; petals marcescent, suborbicular, 0.7-1

mm long, c. 0.7 mm wide, glabrous abaxially,

densely villose adaxially; ovary subglobose,

trigonal, c. 1.5 mm long, glabrous, 3-locular;

style ± obsolete; stigma calyptriform, 0.4-0.6

mm across, glabrous, with margins shallowly

lobed. Fruits ± ellipsoid, c. 4.5 mm long

and 4 mm across, 2- or 3-seeded, glabrous,

smooth; persistent calyx c. one fifth the length

of mature fruit. Seed not seen. Fig. 6.

Additional specimens examined: Western Australia,

[precise localities withheld] Jul 2006, Capobianco

AC702-14 (BRI); Sep 2006, Meissner 736 & Bayliss

(PERTH); Sep 2006, Meissner 737 & Bayliss (BRI); Jul

2005, Jeanes s.n. (BRI).

Distribution and habitat : Beyeria lapidicola

occurs on banded ironstone ranges near

Meekatharra, Wiluna and Menzies in south¬

western Western Australia (Map 17). It

grows in shrubland communities of Callitris

glaucophylla and Acacia spp. or Acacia

aneura on sandy loam soils, mostly on banded

ironstone hills.

Phenology : Flowers and fruits have been

collected from July to September.

Affinities: Beyeria lapidicola is most

similar to B. disciformis and B. minor (Airy

Shaw) Halford & R. J.F.Hend. It differs from

B. disciformis in having fruit with longer

pedicels which are 7-13 mm long compared

with 2-6 mm long, bracts which are oblong,

± foliose and glabrous adaxially compared

Halford & Henderson, Revision of Beyeria

613

Fig. 6. Beyeria lapidicola. A. branchlet with male flowers x 6. B. branchlet showing longitudinal grooves * 4. C.

abaxial view of leaf apex x 9. D. side view of leaf apex x 9. E. transverse section of leaf x 12. F. side view of male flower

x 9. G. face view of male flower x 12. H. side view of female flower x 9.1. face view of female flower x 12. J. side view

of calyx lobe from female flower x 24. K. side view of fruit x 6. L. face view of fruit x 6. A, C-G from Meissner 737 &

Bayliss (BRI); B, H-J from Jeanes s.n ., Jul 2005 (BRI); K&L from Markey 4114 & Dillon (PERTH). Del. W. Smith.

614

with ovate, cymbiform bracts that are densely

hairy adaxially and calyx lobes that do not

enlarge as the fruit matures. It differs from

B. minor in having the calyx lobes shaped

differently (ovate to broad-ovate compared

with suborbicular or very broad-ovate),

female flowers with a smaller stigma which

is 0.4-0.6 mm across compared with 0.7-0.9

mm across, an evenly rounded rather than an

obtuse and apiculate leaf apex and calyx lobes

that do not enlarge as the fruit matures.

Notes : Beyeria lapidicola is listed as Priority

Two under DEC Conservation Codes for

Western Australian Flora, under the name

Beyeria sp. Murchison (B. Jeanes s.n.

7/7/2005) (Florabase, http://florabase.dec.

wa.gov.au [accessed June 2008]).

Etymology : The specific epithet is derived

from Latin lapis , stone, and - cola , dweller

or inhabitant, in reference to the banded

ironstone hills where this species has been

recorded as growing.

16. Beyeria latifolia Baill., Adansonia 6:

304 (1866). Type: [Western Australia.] stony

gully west from Mt Bland, s.d., s.coll. (lecto

[here chosen]: MEL 114159).

Beyeriopsis latifolia Mull.Arg., Linnaea 34:

57 (1865). Type: [Western Australia.] Point

Henry, s.d., \A.F !] Oldfield s.n. (lecto [here

chosen]: G-DC n.v. [microfiche IDC 800-73

2454 II. 3, top element]; isolecto: K, MEL

114160).

Monoecious sometimes dioecious, spreading,

much-branched shrubs to 0.7 m high,

sometimes resinous on young shoots and

flower buds. Young branchlets pale green, ±

terete, densely hairy; hairs stellate, stipitate

or sessile, c. 0.4 mm across; older branchlets

with grey shallowly fissured bark. Leaves

petiolate; petioles 4-7 mm long, sparsely

to densely hairy with stellate hairs as for

branchlets; blades ovate to broad-ovate, 10-

30 mm long, 6-12 mm wide, length:width

ratio 1.5-3:1; adaxial surface glabrous or with

scattered stipitate stellate hairs c. 0.4 mm

across, becoming glabrous with age, smooth

or with scattered tubercles from persistent

hair bases; abaxial surface densely hairy

with ± sessile, stellate hairs c. 1 mm across;

base obtuse to truncate; margins recurved to

Austrobaileya 7(4): 577-639 (2008)

revolute; apex obtuse to rounded; midvein

slightly impressed adaxially, abaxially raised,

sparsely to densely hairy; secondary veins

obscure adaxially, abaxially raised; tertiary

veins obscure; marginal glands rarely present

on blade, up to 1 mm from blade base, 1 per

side of midrib, ± sessile, smooth, c. 0.1 mm

across. Flowers pedicellate, axillary, solitary;

bracts ± ovate, c. 0.9 mm long, acute at apex,

stellate hairy; pedicels sparsely to densely

hairy with hairs as for branchlets, slightly

stouter on female flowers than on male flowers;

calyx lobes 5, yellowish green, suborbicular

or ovate, concave and glabrous adaxially,

gibbose with a few stellate hairs abaxially, the

margins erose, rounded to obtuse to acute at

apex; petals slightly shorter than or equal to

calyx lobes, erect, suborbicular or depressed

elliptic, glabrous abaxially, sparsely villose

adaxially; disc of 5 discrete glands; glands

fleshy, dorsi-ventrally compressed, c. 0.3 mm

long, glabrous, truncate or irregularly lobed.

Male flowers with pedicels 5-10 mm long;

calyx lobes 1.5-1.7 mm long, 0.9-1.5 mm

wide, surrounding androecium at anthesis;

petals 0.5-1.5 mm long, 1.2-1.8 mm wide,

the margins erose distally; receptacle c. 1

mm across, minutely hairy; stamens 9-11;

filaments erect, 0.1-0.3 mm long, glabrous,

bifid distally; anthers c. 0.4 mm long. Female

flowers with pedicels 3-11 mm long; calyx

lobes ± appressed to and enclosing gynoecium

apart from stigma, 0.5-1 mm long, 0.7-1.4 mm

wide; petals marcescent, 0.7-0.9 mm long,

0.7-0.9 mm wide, the margins entire; ovary

± ellipsoid, 0.9-1.3 mm long, densely stellate

hairy, 2- or 3-locular; style ± obsolete; stigma

calyptriform, 0.8-1 mm across, shallowly

umbilicate, glabrous, with margins entire.

Fruits ± ovoid, 6-6.5 mm long, 4-5.5 mm

across, 1-3-seeded, sparsely hairy; persistent

calyx c. one tenth the length of mature fruit.

Seeds ellipsoid, dorsi-ventrally compressed,

c. 6 mm long (including caruncle), 3 mm

across and 2.5 mm in thickness; testa light to

dark brown; caruncle c. 2 mm long and 1 mm

wide, light brown.

Additional specimens examined: Western Australia.

Gairdner River, Qualup [?], Oct 1928, Gardner 2225

(PERTH); 20 miles [c. 32 km] W of Salmon Gums,

Jan 1977, Wittwer W2000 (PERTH); SE of Middle Mt

Barren, Fitzgerald River Reserve, Dec 1970, George

Halford & Henderson, Revision of Beyeria

10590 (CANB, PERTH); 2 km S of Thumb Peak,

Fitzgerald River N.P, Oct 1976, Newbey 4845 (CANB);

West Mt Barren, Oct 1963, Aplin 2769 (PERTH); ibid ,

Nov 1960, George 1786 (PERTH); ibid , Oct 1965,

George s.n. (PERTH); Mt Bland, Res. 24048, Jul 1970,

George 10055 (PERTH).

Distribution and habitat : Beyeria latifolia

is confined to the south coast of Western

Australia where it occurs east of Albany

from Point Henry north-east to Thumb Peak

in the Fitzgerald River National Park, with a

disjunct population near Salmon Gums (Map

18). It grows in coastal scrub and mallee

scrub communities on mostly sandy soils in

sheltered sites.

615

Phenology : Flowers have been collected in

January, July and from October to December,

fruits in October and November.

Affinities : Beyeria latifolia is morphologically

most similar to B. cinerea and B. lepidopetala

F.Muell. Beyeria latifolia differs from the

former by its leaf blade size, and fruit shape

and size. Beyeria latifolia can be distinguished

from B. lepidopetala by its leaf blade shape

and size and shorter pedicels on female

flowers. These differences are summarized in

Table 3

Table 3. Comparison of morphological characters for Beyeria latifolia , B. cinerea and

B. lepidopetala

Character

B. latifolia

B. cinerea

B. lepidopetala

leaf dimensions (mm)

10-30 x 6-12

3-17 x 1.2-5

15-20 x 1.5-3

leaf shape

ovate to broad-ovate

narrow-ovate

to broad-ovate,

narrow-elliptic,

triangular

linear

pedicel length (mm)

of female flowers

3-11

1-7

10-30

fruit dimensions (mm)

fruit shape

6-6.5 x 4-5.5

ovoid

4-6 x 2.5-3.8

pyriform

fruit not seen

Typification : In the protologue of Beyeria

latifolia , Baillon (1866) cited three syntypes

“Oldfield , n. 831, in vallibus umbrosis ad

Portum Henry, Austral, austr.-occid.; Lare?

in lapidosis orient. Montis Bland (herb.

F.Muell .).; Drummond , Swan-River (ser. 4, n.

216?)”. Two sheets of what is considered to be

type material have been located in material

loaned to BRI from MEL. These sheets

are: Point Henry, Oldfield 831 [114160] and

Mt Bland, [114159]. In the absence of further

suitable material, the collection from Mt

Bland [114159] is here selected as lectotype of

this name.

In the protologue of Beyeriopsis latifolia ,

Muller (1865) cited two collections “Point

Henry (Oldfield, comm, Dr, F. Muell. in herb.

DC!)” and “Swan River (Drummond ser. 4.

n. 216!)”. These collections are mounted on a

single sheet in G-DC. The Oldfield collection

is selected as lectotype of this name. The

Oldfield collection in G-DC has the date 1863

on the lower edge of the Botanical Museum of

Melbourne label. This we believed is the year

in which the material was received by Muller

Argoviensis from F. Mueller and is not the

date of its collection.

17. Beyeria lepidopetala F.Muell., Fragm.

1: 230 (1859); Beyeriopsis lepidopetala

(F.Muell.) Mull.Arg., Linnaea 34: 57 (1865).

Type: [Western Australia.] Murchison, s.d .,

A.[Fl\ Oldfield s.n. (lecto[here chosen]: MEL

98605; isolecto: K, MEL [2 sheets 114318,

2062919]).

Illustration : Hopper et al. (1990: 83).

Monoecious or dioecious, erect, open

shrubs to 4 m high. Young branchlets of

unknown colour when fresh, terete, densely

hairy, becoming glabrous with age though

616

remaining tuberculate by persistent hair

bases; hairs stellate, shortly stipitate, up

to 0.5 mm across; older branchlets with

grey shallowly furrowed bark. Leaves

petiolate; petioles 1-2 mm long, hairy with

indumentum as for branchlets; blades linear,

15-20 mm long, 1.5-3 mm wide, length:

width ratio 5-10:1; adaxial surface sparsely

hairywith stipitate stellate hairs c. 0.4 mm

across, becoming glabrous with age though

remaining sparsely tuberculate by persistent

hair bases; abaxial surface densely hairy

with ± sessile, stellate hairs c. 0.5 mm across;

base cuneate or obtuse; margin revolute or

recurved usually to midrib concealing abaxial

leaf surface; apex obtuse to acute; midvein

slightly impressed adaxially, abaxially raised,

densely stellate hairy; secondary and tertiary

veins obscure; marginal glands occasionally

present at base of blade, 1 per side of midrib,

sessile, smooth, c. 0.1 mm across. Flowers

pedicellate, axillary, solitary; bracts foliose,

up to 5 mm long; pedicels sparsely stellate

hairy, slightly stouter and longer on female

flowers than on male flowers; calyx lobes 5, of

unknown colour when fresh, suborbicular or

broad-ovate, glabrous, concavo-convex, the

margins erose, rounded at apex; petals slightly

shorter than calyx lobes, erect, suborbicular,

glabrous abaxially, sparsely villose adaxially,

the margins erose distally; disc of 5 discrete

glands; glands fleshy, 0.4-0.5 mm long,

glabrous, rounded. Male flowers with pedicels

8-10 mm long; calyx lobes c. 1.5 mm long

Austrobaileya 7(4): 577-639 (2008)

and 1.5 mm wide, surrounding androecium at

anthesis; petals c. 1.1 mm long and 1.4 mm

wide; receptacle 0.8-1 mm across, minutely

hairy; stamens 50; filaments erect, c. 0.2

mm long, glabrous, bifid distally; anthers c.

0.2 mm long. Female flowers with pedicels

10-30 mm long; calyx lobes ± appressed to

and enclosing gynoecium apart from stigma,

c. 1.5 mm long and 1.5 mm wide; petals

marcescent, c. 1 mm long and 1.2 mm wide;

ovary ellipsoid, c. 1 mm long, densely stellate

hairy, 3-locular; style ± obsolete; stigma

discoid, c. 0.8 mm across, glabrous, deeply

tri-sulcate with margins entire. Mature fruit

and seed not seen.

Additional specimens examined: Western Australia,

[localities withheld] Aug 2003, Wildflower Society

Bnshland Survey GDNI/56 (BRI); Aug 1974, Cranfield

9326 (PERTH).

Distribution and habitat : Beyeria

lepidopetala is known only from a small area

near Geraldton, Western Australia (Map 19).

It grows on yellow sandy clay.

Phenology : Flowers have been collected in

August.

Affinities: Beyeria lepidopetala is

morphologically most similar to B. cinerea

and B. latifolia. It can be separated from both

species by its leaf shape and size and pedicel

length of the female flowers. These differences

are summarized in Table 4.

Table 4. Comparison of morphological characters for Beyeria lepidopetala , B. latifolia and

B. cinerea

Character

B. lepidopetala

B. cinerea

B. latifolia

leaf dimensions (mm)

15-20 x 1.5-3

3-17 x 1.2-5

10-30 x 6-12

leaf shape

linear

narrow-ovate to broad-

ovate narrow-elliptic,

triangular

ovate to broad-

ovate

pedicel length (mm)

of female flowers

10-30

1-7

3-11

Typification : In the protologue of Beyeria

lepidopetala , Mueller (1865) cited a single

collection “In locis rupestribus ad flumen

Murchison. A. Oldfield Four sheets of what

is considered to be original material have

been located (three at MEL [98605, 114318

and 2062919] and one at K). The MEL sheet

[98605] is selected here as lectotype of this

Halford & Henderson, Revision of Beyeria

name because it is an ample specimen and has

morphology that agrees with the description

in the protologue.

Notes : Listed as Declared Rare Flora under

the Western Australian Wildlife Conservation

Act 1950 (Florabase, http://florabase.dec.

wa.gov.au [accessed June 2008]).

18. Beyeria minor (Airy Shaw) Halford &

R. J.F.Hend. combinatio et status nova

Basionym: Beyeria calycina var. minor Airy

Shaw, Kew Bull. 26: 70-71 (1971). Type:

Western Australia. 85 miles [ c. 136 km] E of

Merredin, October 1964, P.R. Jefferies 641006

(holo: PERTH; iso: PERTH).

Monoecious, erect shrubs to 0.8 m high,

resinous on most parts. Young branchlets

of unknown colour when fresh, ± angular,

longitudinally grooved, glabrous, thickly

resinous; older branchlets terete, with grey

to black shallowly fissured bark. Leaves

petiolate; petioles 0.6-1.2 mm long, glabrous,

minutely papillose (usually obscured by

resinous covering); blades narrow-oblong

to narrow-elliptic, 5-10 mm long, 1.6-2.5

mm wide, length:width ratio 3-4:1; adaxial

surface glabrous, minutely papillose (usually

obscured by resinous covering); abaxial

surface densely hairy with ± sessile, stellate

hairs up to 0.9 mm across; base obtuse;

margins recurved to midrib concealing

abaxial leaf surface; apex rounded or obtuse,

ultimately apiculate with extension from

midrib; apiculum stout, up to 0.5 mm long,

bent upward; midvein obscure or slightly

raised adaxially, abaxially prominently raised

and flattened, glabrous and resinous on

abaxial face; secondary and tertiary veins

obscure; marginal glands occasionally present

on blade, up to 1 mm from blade base, 1 per

side of midrib, sessile, smooth, up to 0.2 mm

across. Flowers pedicellate, axillary, solitary;

bracts narrow-oblong, 0.6-1.1 mm long, acute

at apex, glabrous; pedicels glabrous, papillose

usually obscured by resinous covering, stouter

and longer on female flowers than on male

flowers; calyx lobes 5, of unknown colour

when fresh, suborbicular to very broad-ovate,

glabrous, concavo-convex, obtuse to rounded

at apex; petals slightly shorter than or equal

to calyx lobes, suborbicular to oblate, erect,

617

glabrous abaxially, densely villose adaxially

proximally, the margins erose; disc obscure

or absent in male flowers, of 5 discrete

glands in female flowers. Male flowers with

pedicels 2-6 mm long; calyx lobes c. 1.5

mm long and 2 mm wide, the margins erose,

surrounding androecium at anthesis; petals c.

1 mm long, and 1.3 mm wide; receptacle c.

1.5 mm across, minutely hairy; stamens 20;

filaments erect, c. 0.3 mm long, glabrous,

entire or bifid distally; anthers c. 0.4 mm

long. Female flowers with pedicels 5-9 mm

long; calyx lobes somewhat accrescent, ±

appressed to and enclosing gynoecium apart

from stigma, c. 2.5 mm long (c. 3 mm long in

fruit) and 2.6 mm wide, the margins entire;

petals marcescent, 0.9-1 mm long, 1.3-1.5

mm wide; disc glands fleshy, ovate, c. 0.4 mm

long, dorsi-ventrally compressed, glabrous,

rounded; ovary subglobose, c. 1.2 mm long,

glabrous, 3-locular; style c. 0.1 mm long,

glabrous; stigma discoid to calyptriform, 0.7-

0.9 mm across, glabrous, with margins entire,

± incurved. Fruits subglobose, 4.5-5.5 mm

long, 4-5 mm across, mostly 2- or 3-seeded,

glabrous, smooth; persistent calyx c. half

the length of mature fruit. Seeds ellipsoid,

dorsi-ventrally compressed, 3.5-4.2 mm long

(including caruncle), 2.6-2.9 mm across,

2-2.2 mm in thickness; testa light to dark

brown; caruncle c. 0.7 mm long and 1 mm

wide, creamy-white.

Additional selected specimens examined: Western

Australia. 6 km N of Hickey Ricken Soak, c. 60 km N

of Bullfinch, Sep 1970, Wilson 8768 (PERTH); North

Bungulla, Sep 1926, Gardner s.n. (PERTH); 800 m

NNE of summit of Mt Holland, 87.5 km ENE of Hyden,

Aug 1990, Mollemans 3338 (BRI, PERTH).

Distribution and habitat : Beyeria minor

is known from scattered localities near

Southern Cross, Kellerberrin and Hyden in

south-western Western Australia (Map 20).

It grows in Acacia shrubland communities on

red brown earths or deep sandy soils.

Phenology : Flowers have been collected from

August to October, fruits in October.

Affinities: Beyeria minor is morphologically

most similar to B. calycina and B. disciformis.

It differs from the former by its smaller leaf

blades (5-10 mm long x 1.6-2.5 mm wide

compared with 15-35 mm long x 2-3 mm

618

wide) that are narrow-oblong to narrow-elliptic

rather than linear in outline, and its smaller

accrescent calyx lobes in female flowers (that

are up to 3 mm long compared with up to 7 mm

long). For features distinguishing B. minor

from B. disciformis , refer to the ‘Affinities’

section under that species.

19. Beyeria physaphylla Halford &

R.J.F.Hend. species nova strato viscido

tenui in partibus maxime et ramulis glabris

et longitudinaliter sulcatis B. calycinae Airy

Shaw et B. minori (Airy Shaw) Halford &

R.J.F.Hend. ut videtur maxime affinis sed ab

utroque forma et amplitudine foliorum et apice

laminae folii rotundato non apiculato et calycis

lobis non accrescentibus post florescentiam

distinguenda est. Typus: Western Australia.

c. 5.5 km ESE of Scaddan, on Scaddan road

to Bostock Swamp, 19 September 1988,

R.J.F.Henderson H3180 (holo: BRI; iso: K,

MEL, PERTH, distribuendi).

Beyeria sp. Scaddan (P. van der Moezel

PGV161), in Florabase, http://florabase.dec.

wa.gov.au [accessed June 2008],

Monoecious or dioecious, erect, much-

branched shrubs to 0.5 m high, thinly resinous

on most parts. Young branchlets light brown,

± terete, longitudinally sulcate, glabrous;

older branchlets with grey tessellated bark.

Leaves mostly crowded on short lateral

branchlets, petiolate; petioles 0.5-1 mm long,

glabrous; blades obovate, 2.8-4.5 mm long,

1.5-2.2 mm wide, length:width ratio 1.5-

2:1; adaxial surface glabrous and ± smooth,

resinous; abaxial surface densely hairy

with ± sessile, stellate hairs up to 0.6 mm

across; base cuneate; margins recurved to

midrib concealing abaxial leaf surface; apex

rounded; midvein obscure adaxially, abaxially

raised, ± flattened, glabrous and resinous on

abaxial face; secondary and tertiary veins

obscure; marginal glands absent. Flowers

pedicellate, axillary, solitary; bracts oblong or

obovate, 0.5-1.3 mm long, rounded at apex,

glabrous; pedicels glabrous, stouter on female

flowers than on male flowers; calyx lobes 5,

suborbicular, glabrous, concave adaxially,

gibbose abaxially, the margins erose, obtuse

to rounded at apex; petals slightly shorter

than or equal to calyx lobes, erect, glabrous

Austrobaileya 7(4): 577-639 (2008)

abaxially, sparsely to moderately villose

adaxially, the margins erose distally; disc

obscure or absent. Male flowers with pedicels

2-4 mm long; calyx lobes 1-1.3 mm long,

1-1.1 mm wide, surrounding androecium at

anthesis; petals suborbicular or depressed

obovate, 0.5-0.6 mm long, 0.5-0.6 mm wide;

receptacle 0.8-1 mm across, minutely hairy;

stamens 9-11; filaments erect, 0.1-0.3 mm

long, glabrous, bifid distally; anthers 0.2-0.3

mm long. Female flowers with pedicels 3-4

mm long; calyx lobes ± appressed to and

enclosing gynoecium apart from stigma,

0.9-1 mm long, 0.8-1.1 mm wide; petals

marcescent, broad-oblong, 0.5-0.8 mm long,

0.5-0.8 mm wide; ovary subglobose, c. 0.5

mm long, glabrous, 3-locular; style c. 0.1 mm

long, glabrous; stigma calyptriform, c. 0.9 mm

across, glabrous except for minute scattered

hairs on margin, with margins entire. Fruits

± ellipsoid, sometimes laterally compressed,

4-4.7 mm long, 3.2-47 mm across, 1- or 2-

seeded, glabrous, ± smooth; persistent calyx

c. one fifth the length of mature fruit. Seeds

ellipsoid, dorsi-ventrally compressed, c. 3.5

mm long (including caruncle), 2.5 mm across

and 2.1 mm in thickness; testa mottled, light

and dark brown; caruncle c. 0.8 mm long and

1.6 mm wide, light brown. Fig. 7.

Additional specimens examined : Western Australia.

c. 5.5 km WSW of Scaddan, on Scaddan road to Bostock

Swamp, 19 Sep 1988, Henderson H3179 (BRI); 10 km E

of Scaddan on Scaddan road, Aug 1982, van der Moezel

PGV161 (PERTH).

Distribution and habitat : Beyeria

physaphylla is known only from near

Scaddan, south-western Western Australia

(Map 21). It grows in mallee eucalypt

woodland with a shrubby layer of Melaleuca

spp., Hakea spp. and Leptospermum sp., on

grey sandy soil on the edge of a salt lake.

Phenology : Flowers and fruits have been

collected in September.

Affinities : Beyeria physaphylla is most

similar to B. calycina and B. minor in

having a thin viscid covering over most

parts and branchlets that are glabrous and

longitudinally sulcate. Beyeria physaphylla

can be distinguished from both these species

by its leaf blade shape and size, its lack of an

apiculum at the leaf apex, and in its calyx

Halford & Henderson, Revision of Beyeria

619

Fig. 7. Beyeria physaphylla. A. branchlet with male flowers x 4. B. side view of leaf x 6. C. adaxial view of leaf x 6.

D. transverse section of leaf x 12. E. side view of male flower x 12. F. face view of male flower x 12. G. side view of

female flower x 12. H. face view of female flower x 12.1, side view of ovary and style on receptacle x 15. J. transverse

section of stigma and ovary x 15. K. side view of calyx lobe from female flower x 15. L. abaxial view of calyx lobe

from female flower x 15. M. side view of fruit x 6. N. face view of fruit x 6. O. abaxial view of seed x 10. R side view

of seed x 10. A-P from Henderson H3180 (BRI). Del. W. Smith.

620 Austrobaileya 7(4): 577-639 (2008)

Table 5. Comparison of morphological characters for Beyeria calycina , B. minor and

B. physaphylla

Character

B. calycina

B. minor

B. physaphylla

leaf shape

linear

narrow-oblong to

narrow-elliptic

obovate

leaf dimensions (mm)

15-35 x 2-3

5-10 x 1.6-2.5

3-4.5 x 1.6-2.2

leaf apex

apiculate

rounded

calyx lobes after flowering

accrescent

not enlarging

lobes which do not enlarge as the fruit

matures. These differences are summarized

in Table 5.

Etymology : The specific epithet is from

Greek, physao , puff up, distend, inflate, and

-phyllus , -leaved, and refers to the inflated

appearance of the leaf blades of this species.

20. Beyeria rostellata Halford & R. J.F.Hend.

species nova B. disciformi Halford &

R. J.F.Hend. et B. minori (Airy Shaw) Halford

& R.J.F.Hend. ut videtur maxime affinis sed

ab utroque sed foliorum petiolo longiore

(1.5-2 mm longo non 0.5-1 mm longo ut in

B. disciformi vel 0.6-1.2 mm longo ut in B.

minore ), stylo in floribus femineis longiore

(c. 0.6 mm longo non c. 0.1 mm longo ut in

B. disciformi et B. minore ), pilis stellatis

in pagina abaxiali laminae folii minoribus

(0.05 mm diam. non 0.6 mm diam. ut in

B. disciformi et B. minore), calycis lobis

lato-oblongis, oblongis vel angusto-ovatis

non suborbicularis vel latissime ovatis et

apice laminae folii rotundato non obtuso et

apiculato valido differt. A B. minore lamina

folii angusto-oblonga usque ad lineari non

oblonga usque ad oblongo-elliptica et calycis

lobis non accrescentibus nee non differt.

Typus: Western Australia, [precise locality

withheld] Jackson Range, 7 September 2006,

D.Halford Q9143 & G.Cockerton (holo:

PERTH; iso: BRI, MEL, MO).

Beyeria sp. Jackson Range (R. Cranfield & P.

Spencer 7751), in Florabase, http://florabase.

dec.wa.gov.au [accessed June 2008],

Monoecious, erect, shrubs to 1.8 m high,

resinous on most parts, thickly so on young

shoots and adaxial leaf surface. Young

branchlets of unknown colour when fresh

state, ± terete, longitudinally sulcate,

glabrous; older branchlets with grey to black

shallowly fissured bark. Leaves petiolate;

petioles 1.5-2 mm long, glabrous, minutely

papillose (usually obscured by resinous

covering); blades narrow-oblong to linear,

13-20 mm long, 1.2-2.3 mm wide, length:

width ratio 5-9:1; adaxial surface glabrous

and minutely papillose (usually obscured by

resinous covering); abaxial surface densely

hairy with sessile, stellate hairs up to 0.5

mm across; base cuneate or obtuse; margins

recurved usually to midrib concealing

abaxial leaf surface; apex rounded, ultimately

terminated by sessile gland; midvein

impressed adaxially, abaxially prominently

raised, ± flattened and glabrous on abaxial

face; secondary and tertiary veins obscure;

marginal glands absent. Flowers pedicellate,

axillary, solitary; bracts oblong, c. 1.1 mm

long, rounded at apex, glabrous; pedicels ±

glabrous except for minute erect glandular

hairs proximally, slightly stouter and longer

on female flowers than on male flowers; calyx

lobes 5, green, glabrous, slightly concavo-

convex, the margins ± entire, rounded at

apex; petals slightly shorter than or equal to

calyx lobes, erect, glabrous abaxially, densely

villose adaxially proximally, the margins

erose; disc of 5 discrete glands; glands fleshy,

c. 0.2 mm long, laterally compressed, glabrous.

Male flowers with pedicels 3-5 mm long;

calyx lobes broad-oblong, 1.3-1.5 mm long,

0.8-1 mm wide, surrounding androecium at

anthesis; petals depressed obovate, c. 1.4 mm

long and 1.6 mm wide; disc glands truncate

or irregularly lobed distally; receptacle 0.9-

1.1 mm across, minutely hairy; stamens 25;

Halford & Henderson, Revision of Beyeria

621

filaments erect, 0.3-0.5 mm long, glabrous,

entire; anthers 0.4-0.5 mm long. Female

flowers with pedicels 5-7 mm long; calyx

lobes ± appressed and enclosing gynoecium

apart from stigma, oblong or narrow-ovate,

c. 2.3 mm long and 1.3 mm wide; petals

caducous, obovate, c. 1.4 mm long and 0.9

mm wide; disc glands entire, rounded; ovary

subglobose, 0.9-1.1 mm long, glabrous,

3-locular; style c. 0.6 mm long, glabrous;

stigma discoid, c. 0.7 mm across, glabrous,

with margins ± entire. Fruits ellipsoid,

5-5.5 mm long, 3.5-4 mm across, 2- or 3-

seeded, glabrous, ± smooth; persistent calyx

c. one third the length of mature fruit. Seeds

ellipsoid, dorsi-ventrally compressed, 4.5-6.0

mm long (including caruncle), 2.5-2.6 mm

across, 1.8-2.1 mm in thickness; testa dark

brown; caruncle c. 0.7 mm long and 1.2 mm

wide, light brown. Fig. 8.

Additional selected specimens examined : Western

Australia, [precise localities withheld for conservation

purposes] May 1978, Keighery 1640 (PERTH); Jul 1990,

Mollemans & Mollemans 3101 (BRI, CANB); Nov

1996, Sweedman 4365 (PERTH); Sep 1989, Cranfield

& Spencer 7751 (PERTH); Nov 2000, Carlino 174.TC20

(PERTH); Oct 2003, Cockerton 9106 (BRI); Nov 2000,

Mattiske 175-2/249 (PERTH); Nov 2000, Mattiske 193-

2/586 (PERTH); Aug 2002, Bull s.n. (PERTH); Oct

2000, Mattiske J39-149 (PERTH); Sep 1981, Newbey

9024 (PERTH).

Distribution and habitat : Beyeria rostellata

is known only from the slopes and summit

of Mt Jackson Range, south-western Western

Australia (Map 22). It grows in Acacia

and Eucalyptus tall shrubland or low open

woodland on skeletal red sandy to clay soils

over banded ironstone substrates.

Phenology : Flowers have been collected

in May, July and September, fruits in July,

September and December.

Affinities : Beyeria rostellata is most similar

to B. disciformis and B. minor. The differences

are summarized in Table 6.

Table 6. Comparison of morphological characters for Beyeria rostellata , B. disciformis and

B. minor

Character

B. rostellata

B. disciformis

B. minor

petiole length (mm)

1.5-2

0.5-1

0.6-1.2

leaf shape

narrow-oblong to

linear

oblong to oblong-

elliptic

narrow-oblong to

narrow-elliptic

leaf length (mm)

13-20

10-16

5-10

leaf apex

rounded, without

stout apiculum

rounded or obtuse, with

stout apiculum

rounded or obtuse,

with stout apiculum

calyx lobes shape

broad-oblong,

oblong, narrow

ovate

suborbicular to very

broad-ovate

suborbicular to very

broad-ovate

calyx lobes after

flowering

not enlarging

accrescent

style length (mm)

c. 0.6

c. 0.1

Notes: Beyeria rostellata is listed as Priority

One under DEC Conservation Codes for

Western Australian Flora, under the name

Beyeria sp. Jackson Range (R. Cranfield & P.

Spencer 7751) (Florabase, http://florabase.dec.

wa.gov.au [accessed June 2008]).

Etymology: The specific epithet is from Latin

rostellatus , provided with a short beak, and

refers to the prominent persistent style on the

fruit of this species.

622

Austrobaileya 7(4): 577-639 (2008)

Fig. 8. Beyeria rostellata. A. branchlet with fruit x 2. B. branchlet showing longitudinal grooves x 8. C. abaxial view

of leaf apex x 6. D. side view of leaf apex x 6. E. transverse section of leaf x 12. F. side view of male flower x 12. G.

face view of male flower x 12. H. side view of female flower x 6.1. face view of female flower x 12. J. side view of calyx

lobe from female flower x 12. K. side view of fruit x 4. L. face view of fruit x 4. M. abaxial view of seed x 8. N. side

view of seed x 8. A-N from Halford & Cockerton Q9143 (BRI). Del. W. Smith.

Halford & Henderson, Revision of Beyeria

21. Beyeria similis (Miill.Arg.) Benth., FI.

Austral. 6: 67 (1873); Beyeriopsis similis Mull.

Arg., Linnaea 34: 58 (1865). Type: [Western

Australia.] Swan River, [1852,] J.Drummond

ser. 6 no. 86 (holo: G-DC n.v. [microfiche

IDC 800-73, 2454: IT, 4]; iso: MEL [3 sheets

114226, 114227, 114228], PERTH, K).

Monoecious, erect, much-branched shrubs

to 0.3 m high, resinous on most parts. Young

branchlets of unknown colour, angular

becoming terete with age, glabrous; older

branchlets with grey to black shallowly

fissured bark. Leaves ± sessile; blades linear,

20-35 (-50) mm long, 0.8-1.5 mm wide,

length:width ratio 25-34:1; adaxial surface

glabrous, tuberculate with yellow resinous

dots or rarely smooth; abaxial surface

hairy with ± sessile, stellate hairs c. 0.05

mm across; base cuneate; margins revolute

usually to midrib concealing abaxial leaf

surface; apex obtuse to rounded, recurved;

midvein obscure adaxially, abaxially raised,

± flattened, glabrous and resinous; secondary

and tertiary veins obscure; marginal glands

absent. Flowers pedicellate, axillary, solitary

or fasciculate in 2-4-flowered fascicles;

bracts ± oblong, 0.5-0.7 mm long, acute at

apex, glabrous; pedicels ± glabrous except for

scattered minute erect simple hairs proximally,

stouter and longer on female flowers than

on male flowers; calyx lobes 5, of unknown

colour when fresh, suborbicular, glabrous,

concave adaxially, gibbose abaxially, the

margins ± entire, rounded at apex; petals

slightly shorter than or equal to calyx lobes,

erect, suborbicular, glabrous abaxially, the

margins entire; disc of 5 discrete glands in

male flowers, obscure or absent in female

flowers. Male flowers with pedicels 2-3

mm long; calyx lobes 0.9-1.4 mm long, 0.8-

1.8 mm wide, surrounding androecium at

anthesis; petals 0.7-0.9 mm long, 0.8-1.2 mm

wide; sparsely hairy adaxially proximally;

disc glands thin, triangular, < 0.2 mm long,

dorsi-ventrally compressed, glabrous, acute;

receptacle c. 0.9 mm across, glabrous;

stamens 15; filaments erect, 0.4-0.6 mm

long, glabrous, bifid distally; anthers c. 0.2

mm long. Female flowers with pedicels

5-10 mm long; calyx lobes ± appressed to

and enclosing gynoecium apart from stigma,

623

0.7-1 mm long, 1.2-1.7 mm wide; petals

marcescent, 0.4-0.7 mm long, 0.4-0.6 mm

wide, glabrous abaxially; ovary ellipsoid, c.

1 mm long, glabrous, 2- or 3-locular, with

4 or 6 short subapical appendages; style ±

obsolete; stigma calyptriform, c. 0.4 mm

across, glabrous, with margins entire. Fruits

subellipsoid, 3-6 mm long (excluding horns),

4-5.5 mm across, with 2 small horn-like

subapical appendages, 1-seeded, glabrous, ±

smooth; persistent calyx c. one fifth the length

of mature fruit; subapical appendages up to 2

mm long. Seeds ellipsoid, 4.3-47 mm long

(including caruncle), 3-3.5 mm across, 3-3.5

mm in thickness; testa light to dark brown;

caruncle 0.5-0.8 mm long, 1-1.8 mm wide,

creamy-white.

Additional specimens examined : Western Australia,

[precise localities withheld for conservation purposes],

Oct 1974, George 12908 (BRI); Mt Peron, Aug 1949,

Gardner 9410 (PERTH); Aug 1979, Griffin 2003

(PERTH); Sep 1976, Johnson 3299 (BRI); Sep 1976,

Briggs 6371 (NSW).

Distribution and habitat : Beyeria similis is

confined to the Mt Lesueur area north-east of

Jurien Bay, Western Australia (Map 23). It

grows in shrubland communities on sandstone

ridges, and in heath communities with low

eucalypts on sandy clay soils in valleys and

on shallow grey sand among massive lateritic

duricrust boulders.

Phenology : Flowers and fruits have been

collected in August and September.

Affinities : Beyeria similis is morphologically

most similar to B. gardneri. For distinguishing

characters refer to the Affinities’ section

under B. gardneri.

Notes : Beyeria similis is listed as Priority

Three under DEC Conservation Codes for

Western Australian Flora (Florabase, http://

florabase.dec.wa.gov.au [accessed June

2008]).

624

22. Beyeria simplex Halford & R.J.F.Hend.

species nova B. latifoliae (Mtill.Arg.) Baill.

et B. lepidopetalae Mtill.Arg. ut videtur

maxime affinis sed ab utroque ramulis et

petiolis indumento pilorum simplicium non

stellatorum et ovariis glabris distinguenda est.

A B. latifolia lamina folii angusto-oblonga

vel angustissimo-ovata non ovata ad late

ovata et pedicellis et pagina adaxiali loborum

calycis glabra non pilifera nee non differt.

A B. lepidopetala pedicellis brevioribus

(2-4 mm longis non 8-30 mm longis) nee

non differt. Typus: Western Australia. Cape

Arid National Park, at foot of Mt Ragged,

5 December 1971, R.D.Royce Will (holo:

PERTH; iso: CANB).

Beyeria sp. B Mount Ragged (A.S.George

7422), in Florabase, http://florabase.dec.

wa.gov.au [accessed June 2008],

Monoecious shrubs to c. 0.3 m high,

not resinous. Young branchlets ± terete,

moderately to densely hairy, glabrescent with

age; hairs simple, ascending to spreading,

0.4-0.9 mm long; older branchlets with grey

shallowly fissured bark. Leaves petiolate;

petioles 1-2 mm long, moderately hairy with

hairs as for young branchlets; blades narrow-

oblong or very narrow-ovate, 7-17 mm

long, 1.7-3.5 mm wide, length:width ratio

4-5:1; adaxial surface glabrous and ± smooth;

abaxial surface densely hairy with ± sessile,

stellate hairs c. 0.4 mm across; base obtuse;

margins revolute usually to midrib concealing

abaxial leaf surface; apex rounded; midvein

slightly impressed adaxially, abaxially raised,

moderately hairy with simple hairs up to

0.4 mm long; secondary and tertiary veins

obscure; marginal glands usually present on

blade, up to 1 mm from base, 1 per side of

midrib, ± sessile, smooth, c. 0.1 mm across.

Flowers pedicellate, axillary, solitary; bracts

oblong, 0.1-0.8 mm long, obtuse or rounded

at apex, ± glabrous; pedicels 2-4 mm long,

glabrous, stouter on female flowers than on

male flowers; calyx lobes 5, of unknown

colour when fresh, suborbicular, glabrous,

concave adaxially, gibbose abaxially, the

margins erose, obtuse to rounded at apex;

petals slightly shorter than calyx lobes, erect,

transverse-elliptic, glabrous abaxially, densely

villose adaxially, the margins erose distally;

Austrobaileya 7(4): 577-639 (2008)

disc of 5 discrete glands; glands fleshy, 0.4-

0.5 mm long, laterally compressed, glabrous,

truncate. Male flowers with calyx lobes c.

1.4 mm long and 1.5 mm wide, surrounding

androecium at anthesis; petals 0.9-1 mm long,

1.6-1.8 mm wide; receptacle c. 1.8 mm across,

glabrous; stamens 20-25; filaments erect,

0.3-0.6 mm long, glabrous, bifid distally;

anthers 0.3-0.5 mm long. Female flowers

with calyx lobes ± appressed to and enclosing

gynoecium apart from stigma, c. 1.4 mm

long, and 1.3 mm wide; petals marcescent, c.

0.8 mm long, 0.9-1 mm wide; ovary broad-

ovoid, laterally compressed, c. 1.2 mm long,

glabrous, 2-locular; style 0.1-0.2 mm long,

glabrous; stigma calyptriform, c. 0.3 mm

across, shallowly umbilicate, glabrous, with

margins ± entire. Fruits ± ovoid, laterally

compressed, c. 6 mm long, 5.3 mm across

and 3.6 mm in thickness, 1- or 2-seeded,

glabrous, smooth; persistent calyx c. one fifth

the length of mature fruit. Seeds ellipsoid,

dorsi-ventrally compressed, c. 5.2 mm long

(including caruncle), 3.4 mm across and 2.8

mm in thickness; testa light brown; caruncle

c. 0.6 mm long and 1 mm wide, light brown.

Fig. 9.

Additional specimens examined : Western Australia.

Mt Ragged, Jan 1966, George 7422 (BRI, PERTH); ibid ,

Apr 1996, Barrett 670 (PERTH); lower western slopes of

Mt Ragged, Dec 1999, Hislop 1956B (PERTH); western

slopes of Mt Ragged, Cape Arid, Mar 1987, Keighery &

Alford 1503 (PERTH).

Distribution and habitat : Beyeria simplex

occurs on the south coast of Western Australia

where it is known from Mt Ragged in Cape

Arid National Park (Map 24). It grows in

mallee heath on mountain slopes in stony

sandy soil and in gullies on rocky slopes.

Phenology : Flowers and fruits have been

collected in September, October, December

and March.

Affinities: Beyeria simplex is most similar

to B. latifolia and B. lepidopetala but differs

from both of these species in having an

indumentum of simple rather than stellate

hairs on the branchlets and petioles, and

a glabrous ovary. Beyeria simplex further

differs from B. latifolia in having narrow-

oblong or very narrow-ovate rather than

ovate to broad-ovate leaves, glabrous rather

than hairy pedicels, and calyx lobes that are

Halford & Henderson, Revision of Beyeria

625

Fig. 9. Beyeria simplex. A. branchlet with flowers x 2. B. branchlet showing indumentum x 8. C. adaxial view of leaf

x 4. D. transverse section of leaf x 12. E. side view of male flower x 12. F. face view of male flower x 12. G. side view

of female flower x 12. H. face view of female flower x 12.1. side view of calyx lobe from female flower x 12. J. Stigma

x 24. K. transverse section of stigma and distal half of ovary x 24. L. side view of fruit x 6. M. face view of fruit x 6.

N. abaxial view of seed x 8. O. side view of seed x 8. A from George 7422 (BRI), B-O from Royce Will (PERTH).

Del. W.Smith.

626

glabrous on the adaxial surface rather than

hairy. Beyeria simplex also differs from B.

lepidopetala in having shorter pedicels which

are 2-4 mm long compared with 8-30 mm

long as in that species.

Etymology : The specific epithet is from Latin

simplex , simple, undivided or unbranched,

and refers to the simple hairs on the branchlets

of this species.

23. Beyeria sulcata Halford & R.J.F.Hend.

species nova B. brevifoliae (Miill.Arg.)

Benth. affinis sed foliis costa excurrenti ad

apicem, pedicellis robustioribus et generatim

brevioribus (1-9 mm longis non 5-15 mm

longis), ramulis plerumque sulcatis, foliis

sessilibus vel petiolis usque ad 0.5 mm

longis non petiolis semper 1-2 mm longis et

surculis immaturis viscidis differt. Beyeria

sulcata cum B. lechenaultii (DC.) Baill. antea

confusa sed floribus ambo maribus et femineis

petalis, calycis lobis florum marium plus

minusve erectis et androecium sub anthesi

cingentibus facile distinguenda. Typus:

Western Australia, c. 13 km NE of Bruce

Rock on road to Merredin, 16 September

1988, R.J.F.Henderson H3161 (holo: BRI; iso:

K, MEL, PERTH, distribuendi).

Monoecious, erect to spreading shrubs to 2 m

high, usually resinous on most parts. Young

branchlets of unknown colour when fresh,

± angular, mostly longitudinally grooved,

glabrous; older branchlets with grey to black

flaking or shallowly fissured bark. Leaves

sessile or shortly petiolate; petioles up to 1.7

mm long, glabrous, minutely papillose (usually

obscured by resinous covering); blades narrow-

obovate or linear, 7-35 mm long, 0.8-2.2

mm wide, length:width ratio 8-35:1; adaxial

surface glabrous, minutely papillose (usually

obscured by resinous covering); abaxial

surface glabrous and papillose or densely

hairy with sessile, stellate hairs up to 0.3 mm

across; base cuneate to attenuate; margins

recurved usually to midrib concealing abaxial

leaf surface; apex rounded, obtuse, acute to

attenuate or truncate, ultimately apiculate with

extension from midrib; apiculum stout, up to

0.3 mm long, bent upward; midvein slightly

raised adaxially, abaxially prominently

raised, ± flattened and glabrous; secondary

and tertiary veins obscure; marginal glands

Austrobaileya 7(4): 577-639 (2008)

absent or present on blade, up to 10 mm from

base, 1 or 2 per side, sessile, smooth, up to

0.2 mm across. Flowers pedicellate, axillary,

solitary or fasciculate in 2- or 3-flowered

fascicles; bracts ovate, 0.3-0.7 mm long,

acute to obtuse at apex, glabrous; pedicels

glabrous, slightly stouter on female flowers

than on male flowers; calyx 5, of unknown

colour when fresh, concavo-convex, glabrous,

the margins entire or erose, marginal glands

usually present, rounded to truncate at apex;

petals slightly shorter than or equal to calyx,

suborbicular, erect, the margins erose; disc

in male flowers of 5 discrete glands, female

flowers absent or obscure. Male flowers with

pedicels 0.8-3.5 mm long; calyx lobes broad-

obovate or broad-oblong, 0.8-1 mm long,

0.6-0.9 mm wide, surrounding androecium

at anthesis; petals 0.6-1 mm long, 0.5-1

mm wide, glabrous abaxially, sparsely

villose adaxially proximally; disc glands

fleshy, filiform, up to 0.2 mm long, glabrous;

receptacle c. 0.9 mm across, sparsely stellate

hairy; stamens 7-11; filaments erect, 0.1-0.3

mm long, glabrous, shortly bifid distally;

anthers 0.3-0.4 mm long. Female flowers

with pedicels 0.7-7.5 mm long; calyx lobes

± appressed and enclosing gynoecium apart

from stigma, broad-oblong or broad-obovate,

0.8-1.3 mm long, 0.4-0.9 mm wide; petals

marcescent, 0.5-0.8 mm long, 0.4-0.7 mm

wide, glabrous; ovary ovoid and laterally

compressed or subglobose, 0.8-1.3 mm long,

glabrous, 2- or 3-locular; style ± obsolete;

stigma calyptriform, 0.5-0.7 mm across,

sulcate adaxially, glabrous, with margins

entire. Fruits obloid or broad-ovoid, 2.7-47

mm long, 2.4-3.8 mm across, 2- or 3-seeded,

glabrous, ± smooth; persistent calyx up to one

third the length of mature fruit. Seeds obloid

or ellipsoid, dorsi-ventrally compressed, 2.7-

4.2 mm long (including caruncle), 1.6-2.2

mm across, 1.5-2 mm in thickness; testa dark

brown; caruncle 0.9-1 mm long, c. 0.8 mm

wide, light brown.

Distribution : Beyeria sulcata is endemic to

south-western Western Australia.

Affinities : Beyeria sulcata is similar to

B. brevifolia but differs in having leaves with

the midrib excurrent at the apex, pedicels

which are more robust and generally shorter

(1-9 mm long compared with 5-15 mm long),

Halford & Henderson, Revision of Beyeria

mostly sulcate branchlets, leaves which are

sessile or with a petiole up to 0.5 mm long

(compared with leaves with petioles 1-2 mm

long), and its generally viscid young shoots

compared with non-viscid shoots.

Beyeria sulcata has in the past been

confused with B. lechenaultii (DC.) Baill. but

is easily distinguished in having petals in both

male and female flowers and with the calyx

lobes of male flowers more or less erect and

surrounding the androecium at anthesis.

627

Etymology : The specific epithet is from Latin

sulcatus , furrowed or grooved, and refers to

the longitudinal grooves on the branchlets of

this species.

Notes : Beyeria sulcata exhibits some

discontinuous variation in minor leaf and

stem characters with little geographical

discontinuity. This variation is considered

sufficient to warrant formal recognition of

four varieties within this species which can

be distinguished using the following key.

1 Leaf apices with a truncate apiculum.23d. B. sulcata var. truncata

1. Leaf apices with an acute apiculum.2

2 Young branchlets not distinctly grooved; pedicels 2-9 mm long; leaf

apices acute to attenuate.23c. B. sulcata var. gracilis

2. Young branchlets distinctly longitudinally grooved; pedicels 1-4 mm

long; leaf apices rounded to obtuse.3

3 Leaf blades linear, 0.8-1.1 mm wide.23b. B. sulcata var. brevipes

3. Leaf blades narrow-obovate, 1.3-2.5 mm wide.23a. B. sulcata var. sulcata

23a. Beyeria sulcata Halford & R.J.F.Hend.

var. sulcata

Beyeria brevifolia var. robustior Airy Shaw,

Kew Bull. 26: 69 (1971). Type: Western

Australia. Merredin, 28 November 1923,

M.Koch R2996 (holo: K [2 sheets]; ?iso:

PERTH; MEL [3 sheets]; NSW).

Bark on lower stems ± smooth, shiny,

bronze-coloured, with peeling, reddish

coloured papery strips. Young branchlets

distinctly longitudinally grooved. Leaves

sessile or with petioles up to 0.7 mm long;

blades narrow-obovate, 7-16 (-23) mm long,

1.3-2.5 mm wide, apex rounded or obtuse,

ultimately apiculate with extension from

midrib; apiculum stout, up to 0.2 mm long.

Male flowers with pedicels 1.2-1.7 mm long.

Female flowers with pedicels 0.7-1.5 (-3) mm

long; ovary 2- or 3-locular. Fig. 10 & lla-d.

Additional selected specimens examined : Western

Australia. Bronti [Brontie], Aug 1953, Gardner 12186

(PERTH); along State Vermin Fence No. 7, 60 km

NNW of Southern Cross, Nov 1985, Dodd 222 (CANB,

PERTH); Coolgardie, Oct 1934, Gardner s.n. (PERTH);

Calooli, 9 miles [c. 14 km] SW of Coolgardie, Jul 1925,

Franks s.n. (PERTH); Queen Victoria Springs N.R.

study area, Jun 1987, Pearson 229 (PERTH); Queen

Victoria Spring, N of Zanthus, Oct 1956, Royce 5523

(PERTH); between Cundeelee and Queen Victoria

Spring, Oct 1956, Royce 5516 (PERTH); 12 miles [c.

19 km] S of Queen Victoria Spring, Jan 1956, Royce

5327 (PERTH); near Campion, Aug 1945, Gardner s.n.

(PERTH); Campion, Sep 1945, Gardner 7635 (PERTH);

Booraan [Booran], Aug 1950, Gardner 9540 (MEL,

PERTH); Southern Cross, Aug 1974, Wittwer 1291

(PERTH); 13 km E of Southern Cross, Mar 1966, Wilson

4038 (PERTH); 11 miles [c. 18 km] E of Southern Cross,

Oct 1963, Jefferies 631006 (PERTH); Karalee, Mar 1957,

Jefferies 573074 (PERTH); c. 27 km ESE of Southern

Cross, on road to Coolgardie, Sep 1988, Henderson

H3162 (BRI); 13 km NE of Mt Hampton, c. 50 km SSW

of Southern Cross, Sep 1979, Newbey 5849 (PERTH); 28

miles [ c. 45 km] S of Coolgardie, on Norseman road, Sep

1965, Beauglehole ACB13305 (NSW, PERTH); 11 miles

[c. 18 km] from Lake Grace on Ravensthorpe road, Oct

1965, Knox 65X022 (PERTH); No. 1 Rabbit Proof Fence,

c. 32 km E of Lake King Township, Aug 1968, Wilson

6969 (PERTH).

Distribution and habitat : Beyeria sulcata

var. sulcata occurs in southern Western

Australia in an area more or less bounded

by Kularin in the west, eastward to Queen

Victoria Spring on the southern edge of the

Great Victoria Desert and southwards to

Frank Hann National Park and Lake Grace

(Map 25). It grows in open woodland or

hummock grassland communities on mostly

sandy soils.

628

Austrobaileya 7(4): 577-639 (2008)

Fig. 10. Beyeria sulcata var. sulcata. A. branchlet with flowers x 3. B. branchlet showing longitudinal grooves x 6. C.

abaxial view of leaf apex x 12. D. side view of leaf apex x 12. E. transverse section of leaf x 12. F. side view of male

flower x 24. G. face view of male flower x 24. H. side view of female flower x 24.1. face view of female flower x 24. J.

side view of calyx lobe from female flower x 24. K. side view of ovary and style x 24. L. transverse section of stigma

and ovary x 24. M. side view of fruit x 6. N. face view of fruit x 6. O. abaxial view of seed x 12. P. side view of seed x

12. A-P from Henderson H3161 (BRI). Del. W.Smith.

Halford & Henderson, Revision of Beyeria

Phenology : Flowers have been collected

throughout the year, particularly from August

to November, fruits in February, March and

from June to November.

23b. Beyeria sulcata var. brevipes (Airy

Shaw) Halford & R.J.F.Hend. combinatio

nova

Basionym: Beyeria brevifolia var. brevipes

Airy Shaw, Kew Bull. 26: 69 (1971). Type:

Western Australia. 370 mile peg, Coolgardie

- Norseman road, May 1964, P.R.Jefferies

640515 (holo: K; iso: PERTH).

Bark on lower stems unknown. Young

branchlets longitudinally grooved. Leaves

sessile or with petioles up to 0.7 mm long;

blades linear, 15-22 mm long, 0.8-1.1 mm

wide, apex rounded or obtuse, ultimately

apiculate with extension from midrib;

apiculum stout, up to 0.3 mm long. Male

flowers with pedicels 1-3 mm long. Female

flowers with pedicels 1.5-4 mm long; ovary

2- or 3-locular. Fig. lle-h.

Additional selected specimens examined : Western

Australia, c. 44 km SE of Coolgardie, on road to

Norseman, Sep 1988, Henderson H3171 (BRI); 27

miles [c. 43 km] southward from Coolgardie, Mar 1953,

Gardner 11141 (PERTH); N of Norseman, White s.n.

(PERTH); ibid ; s.dat, White 611008 (PERTH); just north

of Norseman, Mar 1957, Jefferies 573059 (PERTH);

34 km E of Sinclair Soak, c. 93 km NE of Norseman,

Aug 1980, Newbey 7083 (PERTH); North Ironcap,

May 1978, Keighery 1676 (PERTH); near south end of

Lake Cowan, c. 5 km N of Norseman, Jul 1967, Wilson

6052 (PERTH); c. 30 km SSW of Norseman, on road to

Salmon Gums, Sep 1988, Henderson H3174 (BRI); c. 40

km SSE of Norseman, on road to Salmon Gums, Aug

1988, Henderson H3176 (BRI).

Distribution and habitat: Beyeria sulcata var.

brevipes is known from south-western Western

Australia in an area more or less bounded

by Southern Cross, Coolgardie and Salmon

Gums (Map 26). It is recorded as growing

on well-drained sandy soils sometimes with

gravel in mallee communities.

Phenology: Flowers have been collected

in March, May and from July to November,

fruits in May, July and from September to

November.

Affinities: Beyeria sulcata var. brevipes

differs from B. sulcata var. sulcata by its

generally narrower and linear leaf blades.

629

23c. Beyeria sulcata var. gracilis Halford &

R.J.F.Hend. varietas nova foliorum lamina

acuta usque attenuata apiculo usque ad 0.2

mm longum, petiolis et pedicellis plerumque

longioribus et ramulis juvenibus non distincte

longitudinalitersulcatis a varietatibus omnibus

ceteris Beyeriae sulcatae differt. Typus:

Western Australia. 74 km W of Kumarl, c.

122 km N of Esperance, 10 October 1966,

P.G.Wilson 5715 (holo: PERTH).

Beyeria sp. C South West (A.S.George 9878),

in Florabase, http://florabase.dec.wa.gov.au

[accessed June 2008],

Bark on lower stems shallowly fissured,

persistent, dull, grey, rough. Young branchlets

not distinctly longitudinally grooved. Leaves

with petioles 1-1.7 mm long; blades ± linear,

15-35 mm long, 0.9-1.5 mm wide, with the

apex acute to attenuate, ultimately terminated

with an apiculate extension from midrib;

apiculum stout, up to 0.2 mm long. Male

flowers with pedicels 2-3.5 mm long. Female

flowers with pedicels 2-9 mm long; ovary 2-

or 3-locular. Fig. Hi—1.

Additional selected specimens examined: Western

Australia. Hi Vallee Property, Warradarge, towards NW

corner of Butte, Jul 2001, Hislop & Hort MH2253 (BRI);

SW of Norseman, Oct 1961, White 611015 (PERTH);

3 km ENE of Salmon Gums, Mar 1980, Newbey 6671

(PERTH); Newdegate Research Station, 14 km W of

Newdegate, Aug 1968, Wilson 7031 (PERTH); near 260

mile post between Lake Grace and Lake King, Oct 1963,

Jefferies 631020 (PERTH); 25 km W of Lake Grace,

Sep 1967, Wilson 6212 (PERTH); Dunn Rock N.R.,

30 km SW of Lake King, Apr 1984, Backshall DJB87

(PERTH); c. 10 km SW [of] Lake Lockhart near access

road, Oct 1982, Coates AC25 (PERTH); near junction of

West and Phillips Rivers, S of Ravensthorpe, Nov 1965,

George 7179 (PERTH); 56 miles [c. 90 km] W of the

junction of Norseman - Esperance and Lake King roads.

May 1964, Jefferies 640512 (PERTH); 31 km due ENE

of Muckinwobert Rock, 1 km NE of Melaleuca road on

West Point road, Sep 1984, Burgman 3995 (PERTH); 14

km E of Grass Patch on Steddy’s road. May 1982, van

der Moezel PGV70 (PERTH); Esperance, May 1974,

Edmiston E717 (PERTH); 25.5 km NE of Mt Ridley, May

1990, Archer 12059014 (MEL); 24 km NW of Clyde Hill,

May 1983, Burgman & McNee 1210 (PERTH).

Distribution and habitat: Beyeria sulcata var.

gracilis is confined to south-western Western

Australia where it occurs from near Kondinin

eastwards to Clyde Hill, north north east of

Esperance, with a disjunct population near

Eneabba (Map 27). It grows in open shrub

mallee or heath communities on well-drained

sandy soils sometimes with gravel.

630

Austrobaileya 7(4): 577-639 (2008)

Fig. 11. Leaf shape of Beyeria sulcata varieties. B. sulcata var. sulcata. A. abaxial view of leaf x 3. B. side view of

leaf apex x 6. C. abaxial view of leaf apex x 6. D. transverse section of leaf x 12. B. sulcata var. brevipes. E. abaxial

view of leaf x 6. F. side view of leaf apex x 12. G. abaxial view of leaf apex x 12. H. transverse section of leaf x 24.

B. sulcata var. gracilis. I. abaxial view of leaf x 3. J. side view of leaf apex x 12. K. abaxial view of leaf apex x 12. L.

transverse section of leaf x 24. B. sulcata var. truncata. M. abaxial view of leaf x 6. N. side view of leaf apex x 12. O.

abaxial view of leaf apex x 12. P. transverse section of leaf x 24. A-D from Pearson 229 (PERTH); E-H from Jefferies

640515 (PERTH); I-L from Wilson 5715 (PERTH); M-P from White 611017 (PERTH). Del. W. Smith.

Phenology : Flowers have been collected from

March to November, fruits in May, June and

from August to October.

Affinities: Beyeria sulcata var. gracilis

differs from the other varieties of Beyeria

sulcata in having an acute to attenuate leaf

blade apex with an apiculum up to 0.2 mm

long, generally longer petioles and pedicels,

and its young branchlets not being distinctly

longitudinally grooved.

Etymology : The specific epithet is from

Latin gracilis , thin or slender, and refers to

the generally narrower leaves of this variety

compared with those of B. sulcata var.

sulcata.

631

Halford & Henderson, Revision of Beyeria

23d. Beyeria sulcata var. truncata (Airy

Shaw) Halford & R.J.F.Hend. combinatio

nova

Basionym: Beyeria brevifolia var. truncata

Airy Shaw, Kew Bull. 26: 69 (1971). Type:

Western Australia. 92 miles [ c. 147 km]

from Lake King turnoff from Coolgardie -

Esperance road, October 1963, P.R. Jefferies

631019 (holo: PERTH; iso: PERTH).

Bark on lower stems unknown. Young

branchlets longitudinally grooved. Leaves

sessile or with petioles up to 0.7 mm long;

blades linear, 10—13 mm long, 0.8-1.2 mm

wide, apex rounded to truncate, ultimately

apiculate with truncate extension from midrib.

Male flowers with pedicels 0.8-1.3 mm long.

Female flowers with pedicels c. 2.4 mm long;

ovary 2-locular. Fig. llm-p.

Additional specimens examined: Western Australia.

92 miles [c. 148 km] W of Norseman - Esperance road

along Lake King road. May 1964, Jefferies 640514

(PERTH); 34 miles [c. 55 km] N of Ravensthorpe, Oct

1965, Knox 65X034M (PERTH); SW of Norseman, s.d..

White 611017 (PERTH).

Distribution and habitat: Beyeria sulcata var.

truncata is known from a small area east of

Lake King, south-western Western Australia

(Map 28). It grows on gravelly sand.

Phenology: Flowers have been collected in

May and October, fruits in October.

Affinities: Beyeria sulcata var. truncata

differs from other varieties of B. sulcata by

its much more prominent midrib on the lower

surface of leaf blades and the truncate leaf

apex rather than an apiculate excurrent midrib.

It also differs from B. sulcata var. sulcata and

B. sulcata var. gracilis by its narrower leaf

blades.

24. Beyeria villosa Halford & R.J.F.Hend.

species nova a speciebus omnibus ceteris

Beyeriae floribus plus minusve sessilibus

et comparate parvis et ramulis juvenibus

indumento villoso facile distinguibilis est.

Typus: Western Australia. Ravensthorpe

Range [precise locality withheld for

conservation purposes], 25 October 1987,

K.Newbey 11802 (holo: PERTH; iso: BRI).

Beyeria sp. A Ravensthorpe (A.S.George

9474), in Florabase, http://florabase.dec.

wa.gov.au [accessed June 2008],

Monoecious, erect and spreading, much-

branched shrubs to 1.5 m high, resinous on

calyx lobes and adaxial leaf surface. Young

branchlets of unknown colour when fresh, ±

angular soon becoming terete, densely hairy

with simple and bifid hairs up to 0.7 mm long,

glabrescent with age; older branchlets with

grey to black shallowly fissured bark. Leaves

petiolate; petioles 0.9-1.8 mm long, densely

hairy with hairs as for young branchlets;

blades narrow-oblong to linear, 7-14 mm

long, 1.4-2.1 mm wide, length:width ratio 5-

7:1, slightly recurved distally; adaxial surface

almost glabrous except for scattered simple

hairs, ± smooth, thickly resinous; abaxial

surface densely hairy with ± sessile, stellate

hairs c. 0.5 mm across; base abruptly cuneate;

margins revolute to midrib concealing

abaxial leaf surface; apex rounded to

slightly truncate; midvein slightly impressed

adaxially, abaxially raised, densely hairy with

hairs as for abaxial leaf surface; secondary

and tertiary veins obscure; marginal glands

absent. Flowers ± sessile, axillary, solitary or

fasciculate in 2- or 3-flowered fascicles; bracts

narrow-ovate, 0.8-1.1 mm long, acute at apex,

densely hairy with simple appressed hairs up

to 0.6 mm long; calyx lobes 5, of unknown

colour when fresh, hairy adaxially proximally,

glabrous adaxially, concavo-convex, the

margins entire or erose, rounded at apex;

petals slightly shorter than or equal to calyx

lobes, erect, glabrous abaxially and adaxially,

the margins entire or erose distally; disc of 5

discrete glands; glands fleshy, ± terete, 0.2-

0.3 mm long, glabrous. Male flowers with

calyx lobes very broad-ovate, c. 0.6 mm long

and 0.5 mm wide, surrounding androecium

at anthesis; petals broad-elliptic, c. 0.6 mm

long and 0.5 mm wide; receptacle c. 0.6 mm

across, glabrous; stamens 10; filaments erect,

0.3-0.4 mm long, glabrous, bifid distally;

anthers 0.2-0.3 mm long. Female flowers

with calyx lobes ± appressed to and enclosing

gynoecium apart from stigma, oblong or

broad-ovate, c. 0.5 mm long and 0.3 mm wide;

petals marcescent, obovate, c. 0.6 mm long

and 0.5 mm wide; ovary subglobose, laterally

compressed, c. 1 mm long, densely hairy, 2-

locular; style 0.2-0.4 mm long, pubescent

proximally; stigma discoid, c. 0.4 mm long,

shallowly umbilicate, glabrous, with margins

632

entire, recurved. Fruits subglobose or

obpyriform, 3-5 mm long, 2.3-4 mm across,

1-seeded, sparsely to moderately hairy;

persistent calyx c. one quarter the length of

mature fruit. Seeds ellipsoid, 3-4 mm long

(including caruncle), 2-3 mm across; testa

brown to black; caruncle 1-1.3 mm long, c. 1

mm wide, light brown. Fig. 12.

Additional specimens examined : Western Australia,

[precise localities withheld for conservation purposes]

Feb 2005, Craig 6248 (PERTH); Apr 2004, Craig

6084 (PERTH); Feb 1998, Craig 3626 (PERTH); May

2004, Craig 6093 (PERTH); Dec 2004, Mappin 10535

(PERTH); Aug 1969, George 9474 (BRI, PERTH).

Distribution and habitat : Beyeria villosa

occurs near Ravensthorpe in south-western

Western Australia (Map 29). It grows in

mallee or tall shrubland communities often

with Eucalyptus indurata and Melaleuca

pauperiflora on well-drained, calcareous grey

to brown clayey soils.

Phenology : Flowers have been collected in

August and October, fruits in October.

Affinities: Beyeria villosa is not easily

confused with any other species of Beyeria.

It can be distinguished by its comparatively

small, ± sessile flowers and the villose

indumentum on its young branchlets.

Notes: Beyeria villosa is listed as Priority

Four under DEC Conservation Codes for

Western Australian Flora, under the name

Beyeria sp. A Ravensthorpe (A.S.George

9474) (Florabase, http://florabase.dec.wa.gov.

au [accessed June 2008]).

Etymology: The specific epithet is from

Latin villosus , shaggy hairy, and refers to the

indumentum on the young branchlets of this

species.

Excluded and doubtful names

Beyeria bickertonensis Specht, Rec. American-

Australian Sci. Exped. Arnhem Land 3:

249, fig. 8 (1958) = Shonia bickertonensis

(Specht) Halford & R. J.F.Hend.,^ Halford &

Henderson (2005).

Beyeria lasiocarpa forma denudata Baill.,

Adansonia 6: 307 (1866), (‘ denudatum ’).

Type: [Australia.] New England, [s. dat.,]

C.Stuart s.n. (herb. F.Muell.) (holo: ?MEL,

Austrobaileya 7(4): 577-639 (2008)

n.v .) [= possibly Beyeria lasiocarpa Mull.

Arg.].

Beyeria loranthoides Baill., Etude Euphorb.

403 (1858). Type: [Australia. Tasmania or Port

Essington,] 5. dat., Leguille [Guillou ] (holo:

?P n.v.) [= Tasmannia sp.? (Winteraceae)].

Baillon (1858), in the protologue of

B. loranthoides , indicated some taxonomic

doubt as to whether or not this species belonged

in the genus Beyeria. Muller (1866) noted that

Baillon had personally communicated to him

that this species was probably better placed in

Drimys (Winteraceae).

Beyeria opaca var. longifolia Griming in

A.Engler, Pflanzenr. H58: 69 (1913). Type:

[Queensland.] Brisbane, s. dat., anon, per

F.M.Bailey s.n. (holo: B?) [= possibly Beyeria

lasiocarpa Mull.Arg.].

Beyeria tristigma F.Muell., Fragm. 6: 181

(1868) [= Shonia tristigma (F.Muell.) Halford

& R.J.F.Hend, fide Halford & Henderson

(2005)].

Beyeria uncinataVMxxQW. ex Baill., Adansonia

6: 306 (1866) [= Eremophila sturtiiR.Br.,fide

Davies (1987)].

Beyeria virgata Ewart, Proc. Roy. Soc.

Victoria n.s. 33: 226-227 (1921) [= Bertya

virgata (Ewart) Halford & R.J.F.Hend., fide

Halford & Henderson (2002)].

Acknowledgements

We would like to thank Gordon Guymer,

Director of BRI for making working space and

facilities available at BRI for the first author,

and the directors and curators of AD, CANB,

DNA, HO, K, LD, MEL, NE, NSW, P and

PERTH for the loan of their relevant holdings for

study at BRI. The following persons provided

assistance and they are thanked sincerely for

their efforts; Alex Chapman, Roberta Cowan,

Alex George and Peter Bostock for searching

for types on our behalf while Australian

Botanical Liaison Officers at K, Wifi Smith

(BRI) for the excellent illustrations, Gerry

Turpin, Paul Robins and Andrew Franks (all

BRI) for fieldwork assistance undertaken by

the second author and Peter Bostock (BRI) for

preparing the maps. We are indebted to Geoff

Cockerton (Managing Director, Landcare

Halford & Henderson, Revision of Beyeria

633

Fig. 12. Beyeria villosa. A. branchlet with male and female flowers x 4. B. abaxial view of leaf apex x 12. C. side view

of leaf apex x 12. D. transverse section of leaf x 12. E. side view of male flower x 24. F. face view of male flower x 24.

G. side view of female flower x 16. H. face view of female flower x 16.1, abaxial view of calyx lobe from female flower

x 24. J. adaxial view of calyx lobe from female flower x 24. K. transverse section of stigma and ovary x 12. L. side

view of fruit x 4. M. face view of fruit x 4. N. abaxial view of seed x 12. O. side view of seed x 12. A-0 from Newby

11802 (BRI). Del. W. Smith.

634

Holdings Pty Ltd) who generously gave his

time, financial and technical support in 2006

that allowed the first author the opportunity to

study a number of the species in the field. BHP

Billiton Ravensthorpe Nickel Operation is

thanked for permission to access sites on mining

leases in the Ravensthorpe area. Associated

fieldwork from 1988 to 1992 by the second

author and salary support in 1999 and 2000

for the first author was funded by grants from

the Australian Biological Resources Study

(ABRS), Environment Australia, which are

gratefully acknowledged.

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636

Austrobaileya 7(4): 577-639 (2008)

Index to Scientific Names

Names in bold type are accepted names

and those in light type are synonyms etc.

The numbers refer to the pages of the above

taxonomic treatment.

Bertya virgata (Ewart) Halford & R. J.F.Hend

.632

BeyeriaMiq.581

Beyeria apiculata Halford & R. J.F.Hend.

.597

Beyeria backhousei Hook.f..586

Beyeria bickertonensis Specht.632

Beyeria brevifolia (Mull.Arg.) Benth.598

Beyeria brevifolia var. brevifolia .598

Beyeria brevifolia var. brevipes Airy Shaw

.629

Beyeria brevifolia var. robustior Airy Shaw

.627

Beyeria brevifolia var. truncata Airy Shaw

.631

Beyeria calycina Airy Shaw.600

Beyeria calycina var. calycina .600

Beyeria calycina var. minor Airy Shaw . . . 617

Beyeria cinerea (Mull.Arg.) Benth.601

Beyeria cinerea subsp. borealis Halford &

R. J.F.Hend.603

Beyeria cinerea subsp. cinerea .603

Beyeria cockertonii Halford & R. J.F.Hend.

.604

Beyeria constellata Halford & R. J.F.Hend.

.606

Beyeria cyanescens (Mull.Arg.) Benth.603

Beyeria cygnorum (Mull.Arg.) Benth.603

Beyeria disciformis Halford & R. J.F.Hend.

.608

Beyeria drummondii Mull.Arg.586

Beyeria gardneri Airy Shaw.610

Beyeria lanceolata Halford & R. J.F.Hend.

.582

Beyeria lapidicola Halford & R. J.F.Hend.

.612

Beyeria lasiocarpa Mull.Arg.583

Beyeria lasiocarpa forma denudata Baill.

.632

Beyeria lasiocarpa forma lasiocarpa .583

Beyeria latifolia Baill.614

Beyeria lechenaultii (DC.) Baill.585

Beyeria lechenaultii forma elaeagnoides

Baill.585

Beyeria lechenaultii forma genuina Baill. . . 586

Beyeria lechenaultii forma lechenaultii . . . 585

Beyeria lechenaultii forma myrtoides Baill.

.585

Beyeria lechenaultii forma pernettioides

Baill.585

Beyeria lechenaultii forma rosmarinoides

Baill.585

Beyeria lechenaultii forma salsoloides Baill.

.586

Beyeria lechenaultii forma vaccinioides

Baill.586

Beyeria lechenaultii var. backhousei (Hook.

f.) Griming.586

Beyeria lechenaultii var. drummondii (Mull.

Arg.) Griming.586

Beyeria lechenaultii var. genuina Griming

.586

Beyeria lechenaultii var. latifolia Ewart

.586

Beyeria lechenaultii var. latifolia Griming

.586

Beyeria lechenaultii var. lechenaultii .585

Beyeria lechenaultii var. ledifolia (Klotzsch)

Griming.586

Beyeria lechenaultii var. rosmarinoides

(Baill.) Griming.585

Beyeria lechenaultii var. rosmarinoides

Ewart.585

Beyeria ledifolia (Klotzsch) Sond.586

Beyeria ledifolia var. angustifolia Mull.Arg.

.586

Beyeria ledifolia var. backhousei (Hook.f.)

Mull.Arg.586

Beyeria ledifolia var. genuina Mull.Arg.

.586

Beyeria ledifolia var. ledifolia .586

Beyeria lepidopetala F.Muell.615

Beyeria loranthoides Baill.632

Beyeria minor (Airy Shaw) Halford &

R. J.F.Hend.617

Beyeria oblongifolia (Klotzsch) Sond.593

Beyeria opaca F.Muell.590

Beyeria opaca var. latifolia J.M.Black .... 590

Halford & Henderson, Revision of Beyeria

Beyeria opaca var. linearis Benth.586

Beyeria opaca var. longifolia Griming .... 632

Beyeria opaca var. opaca .590

Beyeria opaca var. typica Griming.590

Beyeria physaphylla Halford & R. J.F.Hend.

.618

Beyeria preissii Sond.593

Beyeria rostellata Halford & R. J.F.Hend.

.620

Beyeria Miq. sect. Beyeria.581

Beyeria sect. Beyeriopsis (Mull.Arg.)

Benth.596

Beyeria sect. Discobeyeria Mull.Arg.581

Beyeria sect. Eubeyeria Mull.Arg.581

Beyeria similis (Mull.Arg.) Benth.623

Beyeria simplex Halford & R. J.F.Hend. . . . 624

Beyeria sp. A Ravensthorpe (A.S. George

9474). 631

Beyeria sp. B Mount Ragged (A.S. George

7422). 624

Beyeria sp. Bandalup Hill (G. Cockerton

7553). 604

Beyeria sp. C Southwest (A.S. George 9878

.629

Beyeria sp. Jackson Range (R. Cranfield &

R Spencer 7751). 620

Beyeria sp. Murchison (B. Jeanes s.n.

7/7/2005). 612

Beyeria sp. Scaddan (R van der Moezel

PGV161).618

Beyeria subtecta J.M.Black.591

Beyeria sulcata Halford & R. J.F.Hend.

.626

Beyeria sulcata var. brevipes (Airy Shaw)

Halford & R. J.F.Hend.629

Beyeria sulcata var. gracilis Halford &

R. J.F.Hend.629

Beyeria sulcata var. sulcata .626

Beyeria sulcata var. truncata (Airy Shaw)

Halford &R. J.F.Hend.631

Beyeria tristigma F.Muell.632

Beyeria uncinata F.Muell. ex Baill.632

Beyeria villosa Halford & R. J.F.Hend.631

Beyeria virgata Ewart.632

Beyeria viscosa Miq.592

Beyeria viscosa var. amoena Mull.Arg. . . . 592

Beyeria viscosa var. angustifolia F.Muell. &

Tate.586

637

Beyeria viscosa var. genuina Mull.Arg. . . . 593

Beyeria viscosa var. latifolia Benth.592

Beyeria viscosa var. minor Mull.Arg.592

Beyeria viscosa var. oblongifolia (Klotzsch)

Mull.Arg.593

Beyeria viscosa var. obovata C.T.White

.593

Beyeria viscosa var. viscosa .592

Beyeriopsis Mull.Arg.581, 596

Beyeriopsis brevifolia Mull.Arg.598

Beyeriopsis cinerea Mull.Arg.601

Beyeriopsis cyanescens Mull.Arg.603

Beyeriopsis cygnorum Mull.Arg.603

Beyeriopsis latifolia Mull.Arg.614

Beyeriopsis lepidopetala (F.Muell.) Mull.

Arg.615

Beyeriopsis similis Mull.Arg.623

Calyptrostigma Klotzsch.581

Calyptrostigma ledifolium Klotzsch.586

Calyptrostigma oblongifolium Klotzsch . . . 593

Calyptrostigma viscosum (Labill.) Klotzsch . 593

Croton viscosus Labill.593

Eremophila sturtii R.Br.632

Hemistema lechenaultii DC.585

Shonia bickertonensis (Specht) Halford &

R. J.F.Hend.632

Shonia tristigma (F.Muell.) Halford &

R. J.F.Hend.632

Austrobaileya 7(4): 577-639 (2008)

Maps 2-16. Distribution of Beyeria species. 2. B. lanceolata 3. B. lasiocarpa 4. B. lechenaultii 5. B. opaca

6. B. subtecta 7. B. viscosa 8. B. apiculata 9. B. brevifolia 10. B. calycina 11. B. cinerea subsp. cinerea 12. B.

cinerea subsp. borealis 13. B. cockertonii 14. B. constellata 15. B. disciformis 16. B. gardneri

639

Halford & Henderson, Revision of Beyeria

Maps 17-29. Distribution of Beyeria species. 17. B. lapidicola 18. B. latifolia 19. B. lepidopetala 20. B. minor

21. B. physaphylla 22. B. rostellata 23. B. similis 24. B. simplex 25. B. sulcata var. sulcata 26. B. sulcata var.

brevipes 27. B. sulcata var. gracilis 28. B. sulcata var. truncata 29. B. villosa

A taxonomic revision of Erythrina L.

(Fabaceae: Faboideae) in Australia

A.R. Bean

Summary

Bean, A.R. (2008). A taxonomic revision of Erythrina L. (Fabaceae: Faboideae ) in Australia.

Austrobaileya 7(4): 641-658. Erythrina L. is represented in Australia by the following native or

naturalised taxa; *E. caffra Thunb., *E. crista-galli L., E. fusca Lour., E. insularis F.M.Bailey,

E. numerosa A.R.Bean, *E. speciosa Andrews, *E. x sykesii , E. variegata L., E. vespertilio subsp.

biloba (F.Muell.) A.R.Bean and E. vespertilio Benth. subsp. vespertilio. Erythrina fusca is newly

recorded for Australia. Distribution maps and illustrations are provided for all taxa. A dichotomous

identification key catering for both flowering and fruiting material is included.

Key Words: Erythrina, Fabaceae, Australia, taxonomy, identification key, new species

A.R.Bean, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens,

Mt Coot-tha road, Toowong, Queensland 4066, Australia. Email: tony.bean@epa.qld.gov.au

Introduction

Erythrina L. is a genus comprising around

120 species, occurring throughout the tropics

and subtropics of the world, but with two-

thirds of the species in the Americas (Krukoff

& Barneby 1974).

The last comprehensive account of

Erythrina in Australia (Bentham 1864)

recorded just two species. Krukoff &

Barneby (1974), in their conspectus of the

genus, also recorded just two indigenous

Erythrina species (E. vespertilio and

E. insularis ). Krukoff (1982) again recorded

two indigenous species for Australia. He

maintained^, vespertilio , added E. variegata ,

but excluded E. insularis , which he then

regarded as a synonym of E. vespertilio. The

present account includes five native and four

naturalised taxa. This increase is a reflection

of field-based knowledge of the native taxa

and the recent naturalisation of some long-

cultivated taxa.

Erythrina species are frequently cultivated

as ornamental plants because of their showy

(usually red) flowers. In developing countries,

they are also used for shade or as a convenient

trellis for vines such as pepper, betle leaves,

vanilla, and yam (Whistler & Elevitch 2006).

Accepted for publication 1 August 2008

Some Erythrina are used as a “living fence”,

particularly cultivars chosen for their upright

slender growth habit. Stem cuttings, up to a

metre high and 10 cm diameter, will sprout

roots in moist ground, and when planted

close together, quickly develop into a prickly

impenetrable fence. The ease by which some

Erythrina taxa propagate themselves by

cuttings or even by fallen branches is a factor

in the spread of some alien taxa in Australia,

particularly E. x sykesii.

Materials and methods

The data and descriptions presented here

are based largely on a morphological study

of herbarium specimens at BRI (including

spirit material), NSW and MEL, and images

of type specimens at K. Most taxa have

been examined in the field by the author.

Leaf shapes follow Hickey & King (2000).

Measurements of floral parts are based on

fresh material, or material preserved in 70%

alcohol, or reconstituted. Measurements of

leaves, pods and seeds are based on dried

herbarium material. All measurements are

based on material collected from Australia,

except for Erythrina fusca , where additional

measurements were obtained from New

Guinea specimens. Curved structures (e.g.

standard petal, pod) were measured along the

curvature, and not directly from end to end.

Pod length includes the stipe.

642

Good quality herbarium specimens of

Erythrina are difficult to collect and preserve.

For example, it is rarely possible to collect

flowers and mature fruits at the same time,

and many species are leafless when in flower.

Furthermore, pressed specimens are very

prone to “falling apart” during the drying

process. The leaves and flowers readily abscise

from the branchlets, so that one is often left

with a pile of separate leaflets, flowers, twigs

etc. With these difficulties in mind, I have

attempted to create an identification key that

will cater for imperfect herbarium specimens,

which, it must be said, form the greater part.

Erythrina speciosa is included in the key

to species but not treated in the main text.

A description of it may be found in Green

(1994).

The following abbreviations are used

throughout the paper: L.A. = Logging Area,

N.P. = National Park; N.S.W. = New South

Wales, N.T. = Northern Territory, Qld =

Queensland, S.F. = State Forest, T.R. = Timber

Reserve, W.A. = Western Australia.

Taxonomy

Erythrina L., Sp. PI. 2: 706 (1753), Gen. PI.

ed. 5, 316 (1754). Type: E. corallodendron L.,

[lecto: fide Britton (1924: 427)].

Etymology : from the Greek erythros meaning

red, in reference to the bright red flowers

possessed by most species in the genus.

Austrobaileya 7(4): 641-658 (2008)

Trees, shrubs, or rarely herbaceous resprouters.

Large conical prickles usually present on trunk

and branches, occasionally on the petioles.

Leaves spirally arranged, pinnatelytrifoliolate,

in most species deciduous; stipules present but

often caducous, stipels fleshy and glandular.

Leaflets pinnately-veined though often 3-

veined at the base, intramarginal veins absent.

Inflorescences terminal or axillary, widely

spreading from distal parts of branchlets,

pseudoracemose, indeterminate, oldest

flowers at the base, flowers borne in fascicles,

bracts and bracteoles deciduous. Calyx with

a tubular or obconical hypanthium, often

split down one side at anthesis; calyx lobes

or teeth 1-5, usually obscure and sometimes

lacking. Corolla showy, the keel and wing

petals usually much smaller than the standard.

Stamens 10, with 9 filaments connate above

the middle, alternately longer and shorter,

the upper filament free or basally connate

with the staminal tube; anthers bithecate

with longitudinal dehiscence, dorsifixed.

Style terminal, incurved, not bearded. Fruits

dehiscent, linear or linear-oblong, often

sickle-shaped, more or less moniliform. Seeds

estrophiolate, ovoid, ellipsoid or sub-globose,

with a very hard testa.

Chromosome number: 2 n = 42 (Hennessy

1991)

Key to native and naturalised taxa of Erythrina in Australia

1 Leaflets elliptical (broadest near the middle), 1.5-2.3 times longer than

wide.2

1. Leaflets not elliptical (deltoid, obdeltoid, rhomboid, or variously lobed),

length 0.19-1.5 times that of width.3

2 Leaflets with minute waxy reticulations below * 1 2 ; one or two prickles

usually present on petioles; pedicels 13-25 mm at anthesis; standard

petal and keel petals pink to red; seeds black with brown streaks, 2-6

per pod.7. *E. crista-galli

2. Leaflets without waxy reticulations below; petioles without prickles;

pedicels 5-13 mm at anthesis; standard petal and keel petals mainly

yellow-orange; seeds uniformly dark brown, 6-12 per pod.8. E. fusca

high magnification needed; not obvious on every leaflet

Bean, Revision of Erythrina 643

3 Indumentum persistent on abaxial veins of fully expanded leaves; standard

petal straight, 7-11 mm wide. *E. speciosa Andr. 2

3. Indumentum absent from fully expanded leaves; standard petal at least

slightly curved, 12-46 mm wide.4

4 Indumentum comprising tiny stellate or dendritic hairs; keel petals free

(indigenous taxa).5

4. Indumentum comprising simple hairs; keel petals connate (naturalised

taxa).9

5 Leaflet margins not sinuate or lobed, strongly 3-veined at base with veins

converging at point of petiole attachment (littoral habitats).6

5. Leaflet margins sinuate or with lateral lobes, either not 3-veined at base

or veins converging inside lamina (not in littoral areas).7

6 Terminal branchlets with dark conical prickles; corolla orange-red at

anthesis; pods indehiscent, only slightly constricted between seeds,

with raised anastomosing veins; seeds brown to purplish-brown. 1. E. variegata

6. Terminal branchlets without prickles; corolla greenish-yellow at anthesis;

pods dehiscent, highly constricted between seeds, smooth or faintly

striate; seeds bright red. 2. E. insu laris

7 Leaflets broadly V-shaped, central lobe absent or up to 12 mm long,

lateral lobes parallel-sided, 32-66 mm long and 5-17 mm wide

.3b. E. vespertilio subsp. biloba

7. Leaflets ± rhomboid or obdeltoid, central lobe well developed (8-40 mm

long), lateral lobes tapering, 20-50 mm wide at their base.8

8 Pods 7-14 cm long, containing 1—4(—5) seeds; standard 32-40 mm

long,anthers 2.1—2.8(—3.1) mm long; fascicles 8—15(—25) mm apart

.3a. E. vespertilio subsp. vespertilio

8. Pods 13-24 cm long, containing 5-11 seeds; standard 39-54 mm long,

anthers 2.8-4.3 mm long; fascicles 1.5-8 mm apart. 4. E. numerosa

9 Standard slightly recurved; terminal leaflet 95-180 mm long; fruits never

develop. 6. *E. x sykesii

9. Standard strongly recurved; terminal leaflet 55-105 mm long; fruits

consistently develop. 5. *E. caffra

2 this species is included in the key on the basis of reported naturalisation on Norfolk Island (Green 1994). No naturalised

material has been seen by the present author.

1. Erythrina variegata L., Herb. Amboin. 10

(1754). Type: illustration of “Gelala Alba” in

Rumphius, Herb. Amboin. 2: 234, t. 77 (holo:

the illustration),^^ Merrill (1917: 33, 276).

Erythrina corailodendrum var. orientalis L.,

Sp. PI. 706 (1753); E. variegata var. orientalis

(L.) Merr., Interpr. Herb. Amboin. 276 (1917).

Type: India. Malabar. “Mouricou” in Rheede,

Hort. Malab. 6: 13, t. 7 (lecto: the illustration),

fide Krukoff & Barneby (1974: 431).

Erythrina indica Lam., Encycl. 2(1): 391

(1786). Types: t. 7 of Rheede, Hort. Malab. 6:

13 (syn: n.vfi t. 76 ‘Gelala litorea’ of Rumph.,

Herb. Amboin., 2: 234 (syn: n.v).

Erythrina phlebocarpa F.M.Bailey,

Queensland Agric. J. 1: 368 (1897);

E. indica var. phlebocarpa (F.M.Bailey)

Domin, Biblioth. Bot. 89: 222 (1926). Type:

Queensland. Cook District: Newcastle Bay,

Cape York Peninsula, s.dat., F.L. Jar dine

s.n. (lecto: BRI [AQ22856], the sheet

bearing the fruit ),fide Krukoff (1972: 135).

644

Illustrations : Cooper & Cooper (2004: 210);

Verdcourt (1979: 428); Whistler & Elevitch

(2006: 331, 333); Williams (1979: 110).

Tree 5-25 m high, deciduous. Bark mostly

smooth, with shallow longitudinal furrows,

grey to grey-brown. Trunk and branches

bearing large scattered conical prickles,

terminal branchlets with small black

prickles. Indumentum comprising minute

stellate or dendritic hairs, confined to apical

section of branchlets, developing leaves,

ovaries and young fruits. Leaves uniformly

green or sometimes strikingly variegated

green and yellow, not waxy below. Petioles

unarmed, terete, 40-140 mm long; petiolule

of basal leaflets 6-11 mm long; terminal

leaflet broadly ovate, deltoid or rhomboid,

70-190 mm long, 80-210 mm wide, even

larger on young plants, length/breadth ratio

0.70-1.05, apex acute to acuminate, base

obtuse or truncate; lateral lobes absent; basal

leaflets similar but conspicuously smaller.

Leaflet venation penninerved, basal pair of

lateral veins converging at point of petiole

attachment. Inflorescences borne on leafless

branches, >100-flowered, 20-40 cm long,

rachis and peduncles about same length,

peduncles 6-9 mm diameter; fascicles 3-

flowered, often two or three fascicles at same

position along rachis, each fascicle group 4-6

mm apart on rachis, pedicels 5-9 mm long

at anthesis; calyx tubular in bud, 19-35 mm

long, a ventral longitudinal split developing

as corolla expands, extending almost to base

of calyx, apex truncate or with 2-3 filiform

lobes each 2-6 mm long; all petals red to

scarlet, prominently veined; standard 52-68

x 16-23 mm, apex obtuse, recurved and

exposing staminal column; wings and keel

similar, wings 16-20 mm long; keel petals

16-20 mm long, free, obtuse; stamens 57-70

mm long, anthers 2.9-4.3 mm long, brown;

ovary hairy, ovules 11-16. Pods straight, 8-

25 cm long, 1.7-2.8 cm wide at widest point,

scarcely narrowed between the seeds, outer

surface with raised anastomosing veins, ±

indehiscent and fragmenting when mature,

glabrous at maturity, seeds usually not easily

visible. Seeds 2-10 per pod, ellipsoidal,

13-18 mm long, uniform in colour, brown to

purplish-brown, with a dark hilum. Indian

coral tree. Fig. IF

Austrobaileya 7(4): 641-658 (2008)

Additional specimens examined: Northern Territory.

Cato River, 4 km S of mouth, on headland, Apr 1996,

Cowie 6705 (MEL); Inglis Island, Dec 1987, Dunlop 7437

(BRI, DNA). Queensland. Cook District: Dauan Island,

Torres Strait, Sep 1971, Lawrie s.n. (BRI [AQ003901]);

Warraber Island, Torres Strait, Jul 1996, Waterhouse

3903 (BRI); Murray Island, Aug 1970, Lawrie 105

(BRI); Palfrey Island, near Lizard Island, Jul 1990,

Batianoff 12135 (BRI); Quarantine Bay, Cooktown, Aug

1977, Scarth-Johnson 574A (BRI); Low Isles, 12 km NE

of Port Douglas, Jun 1973, Everist 9964 (BRI); Green

Island, Aug 1973, Stoddart 4247 (BRI); Clump Point,

ESE of El Arish, Sep 1974, Moriarty 1574 (BRI, CANB);

Clump Point, near Mission Beach, Jan 2002, Cooper

WWC1668 & Cooper (BRI). North Kennedy District:

Goold Island, NE Side, Aug 1970, Everist 9696 (BRI);

Cardwell Gap, 13 kmNNE of Ingham, Oct 1976, Everist

s.n. (BRI [AQ 198159]); Port Denison, s.dat ., Fitzalan

s.n. (MEL); Daydream Island, Apr 1990, Batianoff

900461 (BRI); Conway N.P, Airlie Beach, Funnel Bay,

May 1994, Batianoff 9405221 (BRI); Wilson’s Beach,

Conway, SE of Proserpine at head of Repulse Bay,

Apr 1985, Rodd 4439 & Hardie (BRI, NSW); Hayman

Island, Jun 1934, White 10123 (BRI). South Kennedy

District: Lindeman Island, Nov 1985, Batianoff 3332

& Dalliston (BRI); Cape Hillsborough, R60, Ossa, May

1975, Hyland 8220 (BRI); Dolphin Heads, Mackay, Sep

1994, Batianoff 940912 & Saltman (BRI). Port Curtis

District: Eurimbula N.P, 6 kmNW of Agnes Water, Oct

1990, Gibson 1360 (BRI, MEL).

Distribution and habitat : Erythrina

variegata is widely distributed in coastal areas

from Africa to the eastern Pacific Ocean,

including Tanzania, Madagascar, India,

Burma, Vietnam, southern China, Malesia,

New Caledonia and Tahiti (Krukoff 1972).

It is not native to North or South America

but was introduced there over 200 years ago.

In Australia, it is indigenous on the north¬

eastern coast of N.T., in Qld from Torres Strait

and along the east coast as far south as Agnes

Waters (Map 6). It also occurs on some island

territories of Australia, viz. Christmas Island

and North Keeling Island (Du Puy & Telford

1993). It inhabits littoral rainforest or open

forest behind beaches and headlands, and

along the tidal sections of rivers.

Phenology : Flowers from August to October;

fruits are recorded for nearly every month of

the year.

Notes : Krukoff (1972) insisted that the

leaves of the type collection of Erythrina

phlebocarpa belong to the African species

E. lysistemon , while the fruits are of

E. variegata. He did not give any reasons for

identifying the leaves as E. lysistemon , and I

cannot discern any distinctive feature of the

Bean, Revision of Erythrina

645

Fig. 1. Terminal leaflets of Erythrina spp. A. E. vespertilio subsp. vespertilio. B. E. vespertilio subsp. biloba

C. E. caffra. D. E. crista-galli. E. E. numerosa. F. E. variegata. A from Bean 8492 ; B from McRae s.n. (AQ501724);

C from Stephens GBS12 ; D from Hall & Bird s.n. (AQ659717); E from Grimshaw G8 et al. ; F from Moriarty 1574. All

BRI. Del. B. Connell.

foliage of E. lysistemon that would support

the claim. The foliage is entirely consistent

with E. variegata , and it seems clear that all

parts of the type collection are referable to

that species. Krukoff’s lectotypification of

E. phlebocarpa on the fruiting sheet at BRI

must however be followed.

This species is widely utilised by mankind

throughout its range, as a fodder tree, a trellis

plant, or as a living fence (Whistler & Elevitch

2006 ).

Etymology : From the Latin variegatus

meaning variegated. A variant of the

species has leaves displaying variegation

646

(conspicuously white veins on an otherwise

green leaf). Apparently this variant was the

one described by Linnaeus.

2. Erythrina insularisF.M.Bailey,(9wee«,s7a«<i

Agric. J. 1:228 (1897). Type: Queensland. Cook

District: Turtle Island, June 1897, F.M. Bailey

29 (holo: BRI; iso: MEL).

Illustration: Cooper & Cooper (2004: 210).

Tree 4-12 m high, deciduous. Bark mostly

smooth, with pale longitudinal fissures, grey

to brown. Trunk and branches with or without

prickles, terminal branchlets without prickles.

Indumentum comprising minute stellate or

dendritic hairs, confined to apical section of

branchlets, developing leaves, ovaries and

young fruits. Leaves uniformly green, not

waxy below. Petioles unarmed, terete, 45-135

mm long; petiolule of basal leaflets 4-10 mm

long; terminal leaflet deltoid to rhomboid,

60-100 mm long, 55-120 mm wide, length/

breadth ratio 0.78-1.11(—1.32), apex obtuse

or acute; base obtuse to broadly cuneate;

margins sometimes slightly sinuate. Leaflet

venation penninerved, basal pair of lateral

veins converging at point of petiole attachment

(i.e. 3-veined at base). Inflorescences borne on

leafless branches, 15-40-flowered, 7-19 cm

long, rachis and peduncles about same length,

peduncles 3.5-6.3 mm diameter; fascicles

3-flowered, often two or three fascicles at

same position along rachis, each fascicle

group 2-7 mm apart on rachis, pedicels 7-11

mm long at anthesis; calyx tubular in bud,

15-19 mm long, truncate, a longitudinal

ventral split developing as corolla expands,

extending almost to base of calyx; petals

initially pale yellow, senescing to orange or

scarlet, prominently veined; standard 25-41

x 12-15 mm, apex obtuse, strongly recurved

and exposing staminal column, wings and

keel very similar, wings 12-16 mm long; keel

petals 14-15 mm long, free, obtuse; stamens

35-40 mm long, anthers 3.2-4.2 mm long,

brown; ovary hairy, ovules 8-11. Pods often

curved at maturity, 7-17 cm long, 1.3-1.8 cm

wide at widest point, conspicuously narrowed

between the seeds, glabrous, dehiscent, outer

surface with faint raised parallel veins. Seeds

2-7, ellipsoidal, 9.5-12 mm long, uniform

in colour, scarlet to red, with a pale or dark

Austrobaileya 7(4): 641-658 (2008)

hilum, often remaining attached to pod and

easily visible. Fig. 2B.

Additional specimens examined : Queensland.

Cook District: Gabba Island, Torres Strait, Jul 1996,

Waterhouse 3891 (BRI); Dalrymple Island, Jan 1998,

Waterhouse 4766 (BRI); Pumpkin Island, Jun 2003,

Wannan 3101 & Buosi (BRI); Coconut Island, central

Torres Strait, Sep 1971, Lawrie s.n. (BRI [AQ008846]);

Coconut Island, Aug 1994, Wannan 64 (BRI); western

end of Warraber Island, Torres Strait, May 2003,

Hacks LAH164 (BRI, CANB); Warraber (formerly

Sue) Island, Torres Strait, Jun 1995, Waterhouse 3668

(BRI); Milman Island, Cape York, Jan 1995, Card MC24

(BRI); On coast, near Cowal Creek and False Orford

Ness, Sep 1987, Gitay HG101 (BRI); Hannibal Island,

near Shelburne Bay about 16 km W of Helby Hill, Jul

1969, Done s.n. (BRI [AQ007594]); Bird Island, Nov

1973, Stoddart 5090 (BRI); Farmer Island N.P., Great

Barrier Reef Marine Park, Dec 2004, Batianoff 31229

(BRI); Farmer Island, Feb 1991, Card FAR1 (BRI); ex

Haggerstone Island, cultivated Tolga, Oct 2001, Ford

AF3028 (BRI).

Distribution and habitat: Erythrina insularis

is endemic to Qld. It is confined to the Torres

Strait and north-eastern Cape York Peninsula

(Map 4). It inhabits semi-evergreen notophyll

vine forest, often as an emergent. Nearly all

records are from coral islands or cays, but

there is a single known occurrence from a

continental island (Gabba Island), and one

record from the mainland (Cowal Creek).

Phenology: Flowers from July to November;

fruits from December to July.

Notes: Adema (1996) included Erythrina

merrilliana Krukoff as a synonym of

E. insularis. The former was described from

Papua New Guinea (Central Province, 700

metres altitude). New Guinea specimens

may be consistently distinguished by the 5-6

mm long calyx lobes and the orange to red

corolla (calyx lobes lacking, corolla initially

pale-yellow, senescent flowers orange for

E. insularis). These characters, in addition to

marked differences in habitat, indicate that

they are distinct species.

Erythrina insularis is sympatric with

E. variegata. For example, both species

have been collected from Warraber island

{Waterhouse 3903\ Hucks 164). Sterile

specimens of E. insularis and E. variegata

are of very similar appearance, but can be

distinguished by the presence (E. variegata)

or absence (E. insularis) of prickles on the

Bean, Revision of Erythrina

647

Fig. 2. Terminal leaflets of Erythrina spp. A. E. vespertilio subsp. vespertilio. B. E. insularis. C. E. fusca.

D. E. x sykesii. A from Hinz 533 ; B from Hucks 164 ; C from Morris s.n. (AQ497997); D from Batianoff0302118. All

BRI. Del. B. Connell.

terminal branchlets. Fertile plants may be

readily identified. The flowers of E. insularis

are pale greenish-yellow on the day of opening,

changing to orange-red when senescent,

standard 25-41 mm long ( E. variegata

flowers orange-red at all stages, standard

52-68 mm long); the pods of E. insularis are

smooth or faintly striate, moniliform and the

seeds bright red (pods reticulately veined, not

constricted, seeds brown or purplish-brown

for E. variegata).

Bailey noted in the protologue that some

trees were completely without prickles, and

some recent collectors have noted the same.

Card (2001) recorded that (for Milman Island)

prickles or spines are absent from the stems

of the species.

Card (2001) discovered that the seeds of

Erythrina insularis , in contrast to E. variegata

and E. fusca, do not float in seawater, but rather

sink immediately to the bottom. However,

she said that seeds could still be ocean

dispersed when attached to pods or caught

up in rafts of drift material. This dispersal

difficulty perhaps explains why E. insularis

has such a restricted distribution compared to

E. variegata and E. fusca.

Etymology: From the Latin insularis, meaning

an island, because of the island location of the

first collection.

648

3. Erythrina vespertilio Benth. in T.L.

Mitchell, J. Exped. Trop. Australia 218

(1848); Corallodendron vespertilio (Benth.)

Kuntze, Revis. Gen. PI. 1: 173 (1891). Type:

Queensland. Warrego District. Sub¬

tropical New Holland [near Mt Faraday], 29

June 1846, T.L. Mitchell 189 (lecto: K , fide

Krukoff & Barneby (1974: 435)).

Tree 3-20 m high, deciduous. Bark corky,

fissured or furrowed, creamy, grey, grey-

brown or yellowish cream-grey. Trunk and

branches bearing prickles, terminal branchlets

with or without prickles. Indumentum

comprising minute stellate or dendritic hairs,

confined to apical section of branchlets,

developing leaves, ovaries and young fruits.

Leaves uniformly green, not waxy below.

Leaflet venation penninerved, either not 3-

veined at base or veins converging above base

of leaflet. Inflorescences borne on leafless

branches, 13-35-flowered, 11-33 cm long,

rachis and peduncle about the same length,

peduncle 1.5-4.2 mm diameter; fascicles 1-3-

flowered, often two fascicles at same position

along rachis, each fascicle group 8—15(—25)

mm apart on rachis, pedicels 12-24 mm

long at anthesis; calyx tubular in bud, 15-21

mm long, truncate or with minute lobes, a

longitudinal lateral split developing as corolla

expands, extending almost to base of calyx;

Austrobaileya 7(4): 641-658 (2008)

all petals orange-red to scarlet, prominently

veined; standard 32-40 x 16-20 mm, apex

obtuse, strongly recurved and exposing

staminal column; wing and keel petals about

equal in size and shape; wings 8-16 x 3-5.5

mm long; keel petals 10-13 x 3-5.5 mm

long, free, obtuse; stamens 35-45 mm long,

anthers 2.1—2.8(—3.1) mm long, pale brown to

brown; ovary densely stellate-hairy, ovules

6-8. Pods slightly curved when mature, 7-14

cm long, 1.1-1.5 cm wide at widest point,

conspicuously narrowed between the seeds,

glabrous at maturity, dehiscent and with seeds

easily visible, outer surface smooth or with

faint raised parallel veins. Seeds 1-4, rarely

5, ellipsoidal, uniform in colour, orange to

scarlet, with a pale hilum. Bat’s wing coral

tree.

Distribution : This species is endemic to

Australia and comprises two subspecies.

Typification: There are two sheets at K

collected by T. Mitchell, both available to

Bentham when drawing up the protologue.

Krukoff & Barneby (1974) cited “ Mitchell

189 (1846) (K-holotype)”, interpreted here as

lectotypification of the name.

Etymology : From the Latin vespertilio , a bat.

This refers to the shape of the leaflets, which

resemble the profile of a bat in flight.

The subspecies can be distinguished as follows:

Leaflets broadly V-shaped; central lobe absent or up to 12 mm long, lateral

lobes parallel-sided, 32-66 mm long and 5-17 mm wide; seeds 11.5—

16.5 mm long.subsp. biloba

Leaflets rhomboid to obdeltoid, central lobe well developed (8-40 mm

long), lateral lobes tapering, 20-50 mm wide at their base; seeds 10-11.5

mm long.subsp. vespertilio

3a. Erythrina vespertilio Benth. subsp.

vespertilio

Erythrina vespertilio var. vespertilio , Domin,

Biblioth. Bot. 89: 221 (1926).

Erythrina vespertilio var. typica Domin,

Biblioth. Bot. 89: 221 (1926), nom. illeg.

Illustrations: Brock (1988: 149), as

E. variegata var. orientally Williams (1979:

111); Cooper & Cooper (2004: 211).

Petioles unarmed, terete, 40-82 mm long;

petiolule of basal leaflets 5-6 mm long;

terminal leaflet 28-85 mm long, 33-112 mm

wide, length/breadth ratio mostly 0.6-0.84, to

1.06 on Cape York peninsula; base cuneate;

terminal lobe 8-40 mm long, apex acuminate,

acute or obtuse; margins sinuate, forming

obtuse, tapering, often prominent lateral lobes

14-38 mm long, 20-50 mm wide at their base.

Seeds 10-11.5 mm long. Figs 1A, 2A, 3.

Bean, Revision of Erythrina

649

Fig. 3. Erythrina vespertilio subsp. vespertilio. A. leafy branchlet * 0.5. B. single inflorescence bearing buds and

flowers x 0.5. C, E. the two halves of an atypically long fruit containing seeds x 0.5. D. typical fruit x 0.5. F, G. seed x

3. A & B from Forster PIF7686 (BRI); C-G from Forster PIF7730 (BRI). Del. W. Smith.

650

Additional selected specimens examined: Western

Australia, near Poompangala Hill, 9 km NW of

Kalumburu, May 1993, Cowie 4277 (MEL); on road to

Warrender, past Lone Dingo turn off, Mitchell Plateau,

Sep 1978, Clayton-Greene s.n. (MEL2117860A); near

Ord River, 1884, Johnston s.n. (MEL72472A); on

ridge between Calder River and billabong, c. 5 km N

of Walcott Inlet, Manja Camp, Aug 2001, Shepherd 62

(MEL). Northern Territory. Groote Eylandt, Hemple

Bay, May 1948, Spechts.n. (BRI [AQ230649]); Nabarlek,

Jun 1989, Hinz 533 (BRI); Oenpelli, Oct 1948, Specht

1309 (BRI); about SW of Brock’s Creek, Jul 1946, Blake

16478 (BRI); Rockhole Creek, Nitmiluk, Jul 1990,

Evans 3285 (BRI); near mouth of Foelsche River, Jul

1948, Perry 1831 (BRI); Guy Cave Area, 16 miles Cave

Reserve, south of Katherine, May 1978, Webb 13028

(BRI). Queensland. Cook District: Thursday Island,

Jul 1975, Stocker 1238 (BRI); Archer River, Wenlock

- Coen road, Jul 1948, Brass 19760 (BRI); Larrod’s

Hill, Sep 1976, Hyland 8955 (BRI). Burke District:

Karumba, Aug 1943, Blake 15132 (BRI, MEL). North

Kennedy District: Lansdown, c. 25 miles [40 km] S of

Townsville, Mar 1971, Robertson T203 (BRI); Mt Abbot,

50 km W of Bowen, Oct 1992, Bean 5202 (BRI). South

Kennedy District: 2 km SW of Dudgeon Point, Mackay,

Feb 1993, Batianoff930252 (BRI); 1 km S of Cape River,

on Gregory Developmental Road, Nov 1991, Thompson

128 (BRI). Leichhardt District: c. 3.2 km S of Umolo

turnoff, Nov 1973, Williams 367 (BRI). Port Curtis

District: Pine Mt, Marlborough, May 1998, Batianoff

9805213 (BRI); Mt Castletower N.P, eastern slopes.

Many Peaks Range, Feb 1995, Forster PIF16319 (BRI).

Burnett District: Bania S.F., north of Mt Perry, Mar

1995, Bean 8492 (BRI, MEL); 6 km NNE of Coalstoun

Lakes, Apr 1991, Forster PIF7860 (BRI). Wide Bay

District: Delley Road, N of Biggenden, Nov 2001, Bean

18057 (BRI, MEL); 2 km NNE of Didcot, Nov 1997,

Forster PIF21858 (BRI). Moreton District: Redbank

Plains, Ipswich, Jan 1992, Bird s.n. (BRI [AQ520215]).

Distribution and habitat : This subspecies is

widespread across northern and north-eastern

Australia (Kimberley region of W.A., Top End

of N.T. and the eastern half of Qld). In Qld it

extends as far north as Thursday Island, and

south to around Ipswich (Map 8).

Phenology : For northern Australia (the

Top End of N.T., Gulf of Carpentaria, far

north Qld), flowering occurs from June to

September; in central and southern Qld,

flowering occurs from October to December;

fruits may be found from November to April.

Notes: While the leaflets of Erythrina

vespertilio subsp. vespertilio are strongly 3-

veined at the base, the veins almost always

converge at a point inside the lamina surface,

i.e. removed from the point of insertion of the

petiole. This provides a very useful way to

Austrobaileya 7(4): 641-658 (2008)

distinguish sterile specimens of E. vespertilio

from most other Australian species.

3b. Erythrina vespertilio subsp. biloba

(F.Muell.) A.R.Bean, combinatio et status

nova

Basionym: Erythrina biloba F.Muell.,

Hooker’s J. Bot. Kew Gard. Misc. 9: 21

(1857); E. vespertilio var. biloba (F.Muell.)

Domin, Biblioth. Bot. 89: 221 (1926). Type:

Northern Territory or Western Australia, at

the foot of a little granite range at the head

of Hooker’s Creek and Sturt’s Creek, January

1856, F Mueller s.n. (holo: MET 72396; iso:

K).

Illustration : Milson (2000: 82-83).

Petioles unarmed, terete, 48-110 mm long;

petiolules 3-5 mm long; leaflets broadly Y-

shaped, terminal leaflet 14-36 mm long, 33-

144 mm wide, length/breadth ratio 0.19-0.32

(-0.51); base cuneate to attenuate; terminal

lobe often absent, sometimes present, 0-12

mm long, apex acute or obtuse; lateral lobes

32-66 mm long, 5-17 mm wide, straight or

somewhat recurved, length/breadth ratio 3.3-

9.1, obtuse, parallel-sided. Seeds 11.5-16.5

mm long. Fig. IB.

Additional selected specimens examined : Western

Australia. Nickol Bay, 1876, Crouch s.n. (MEL);

Blackfellows Creek crossing on Winnama Yards track,

3 km S of Texas Downs road, Jul 1984, Scarlett 84-387

(MEL); Fortescue River, 1878, Forrest s.n. (MEL).

Northern Territory. 6 miles [10 km] N of Aileron

Homestead, Mar 1955, Winkworth 868 (BRI); Barrow

Creek, 1880, Chandler s.n. (MEL); Yarraji, 63 km NE

Bob Well, Lander River, Jul 1989, Latz 11553 (MEL);

Singleton, 240 miles [386 km] N of Alice Springs, Jan

1950, Everist 4241 (BRI); 13 miles [21 km] NW of

Napperby Homestead, May 1956, Chippendale 2059

(BRI, NSW); c. 4 miles [c. 6 km] NNW of Mt Ziel, Jul

1968, Beauglehole 27180 (MEL); N side of Mt Liebig,

Jul 1988, Corrick 10336 (MEL); Hatches Creek, Nov

1968, Byrnes NB1132 (BRI); Redbank Gorge, Heavitree

Range, Sep 1969, Gittins 2038 (BRI, NSW). South

Australia, north of Oodnadata, 1915, Freeman s.n.

(NSW). Queensland. Burke District: Lake Moondara

road, Mt Isa, Nov 1996, Dean SB72 (BRI); 58 miles

[93 km] SSE of Camooweal, May 1948, Perry 729

(BRI). Mitchell District; Grey Range, about 3 km W

of Mount Grey, on Blackall - Yaraka road, Nov 1990,

McRae s.n. (BRI [AQ501724]); Parkgate, c. 55 km SE

of Barcaldine, Jan 1995, Smyrell GS105 (BRI). Gregory

North District: Bladensburg N.P., S of Winton, Opalton

road. Mar 1998, Forster PIF22195 & Booth (BRI, MEL);

Ray more - Kyabra road, 60 km NNW of Ero manga, Jan

651

Bean, Revision of Erythrina

1998, Pegler LKP152 (BRI). Warrego District: Idalia

N.P., Emmet Pocket, Nov 1990, Morgan CM35 (BRI);

Adavale, Mar 1922, Graw s.n. (BRI [AQ230644]).

Distribution and habitat : Widely distributed

in central Australia, north of about 28° S

latitude, including W.A., N.T., Qld and the far

north of S.A. (Map 7). It is not known from

N.S W It grows on a wide range of soil types,

often in locations that receive extra water e.g.

base of rocky hills, creek banks etc.

Phenology : Flowers are recorded from

October to March; fruits from January to

May.

Notes : This subspecies differs from

subspecies vespertilio mainly in leaf shape.

The difference is however, so marked, and so

well correlated to geographical distribution,

that a rank of subspecies is considered to be

appropriate. The two subspecies do show some

intergradation in leaf form, as evidenced by

three specimens at BRI ( Hyland 6149 , Morgan

CM35, and Clemens s.n. [AQ199198]), but the

great majority of collections may be readily

assigned to one or the other. The seed size

also appears to be diagnostic, as for each of

the eight seed-bearing specimens at BRI and

MEL, the seed length of Erythrina vespertilio

subsp. biloba exceeds that for E. vespertilio

subsp. vespertilio.

Etymology: from the Latin bi meaning two

and lobus meaning lobes. This is a reference

to the distinctive leaflet morphology.

4. Erythrina numerosa A.R.Bean species

nova Species nova affinis E. vespertilio

sed leguminibus longioribus semina 5-11

continentibus, ovulorum numeris majoribus,

caudice laevi unsulcato, floribus in

inflorescentia magis dense aggregata, antheris

longioribus et corolla pallidiore (aurantiaca

usque vitellina) differt. Typus: Queensland.

Moreton District: Splityard Creek,

Wivenhoe Dam, 10 February 1991, L.H.Bird

& P.R.Sharpe s.n. (holo: BRI [AQ547555]; iso:

A, CANB, K, MEL, NSW, US, distribuendi).

Erythrina sp. (Croftby P.I.Forster+ PIF6209)

in Bostock & Holland (2007).

Illustrations: Nicholson & Nicholson (1991:

26), as E. vespertilio ; Williams (1979: 110), as

Erythrina sp.

Tree 5-20 m high, deciduous. Bark smooth,

not furrowed, somewhat shiny, creamy, grey,

brownish or greenish, with conical corky

outgrowths surmounted by prickles. Trunk,

branches and terminal branchlets all bearing

prickles. Indumentum comprising minute

stellate hairs, confined to apical section of

branchlets, developing leaves, ovaries and

young fruits. Leaves uniformly green, not

waxy below. Petioles unarmed, terete, 45-120

mm long; petiolule of basal leaflets 3.5-7 mm

long; terminal leaflet 60-95 mm long, 78-113

mm wide, length/breadth ratio 0.63-1.09,

apex obtuse; base obtuse to cuneate; margins

sinuate, forming obtuse, tapering, often

prominent lateral lobes. Leaflet venation

penninerved, either not 3-veined at base

or veins converging above base of leaflet.

Inflorescences borne on leafless branches,

15-60-flowered, 14-46 cm long, rachis

shorter than peduncle, peduncles 3.5-9 mm

diameter; fascicles 3-flowered, often two or

three fascicles at same position along rachis,

each fascicle group 1.5-8 mm apart on rachis,

pedicels 9-17 mm long at anthesis; calyx

tubular in bud, 17-25 mm long, truncate or

with minute lobes, a longitudinal lateral split

developing as corolla expands, extending

almost to base of calyx; all petals salmon-

coloured to orange, prominently veined;

standard 39-54 x 21-28 mm, apex obtuse,

strongly recurved and exposing staminal

column, wings slightly longer than keel,

wings 15-18 mm long; keel petals 13-16 mm

long, free, obtuse; stamens 35-48 mm long,

anthers 2.8-4.3 mm long, yellow to brown;

ovary densely stellate hairy, ovules 12-16.

Pods strongly curved at maturity, 13-24

cm long, 1.1-1.6 cm wide at widest point,

conspicuously narrowed between the seeds,

dehiscent when mature with seeds easily

visible, glabrous at maturity, surface smooth

or striate. Seeds 5-11, ellipsoidal, 9-13 mm

long, uniform in colour, orange to red, with

a pale hilum. Pine Mountain coral tree , Pine

Mountain corkwood. Figs IE, 3.

Additional selected specimens examined : Queensland.

Burnett District: Meandu Mine, Tarong Coal, Feb

1994, Forster PIF14838 & Smyrell (BRI). Wide Bay

District: Oakview S.F. 220, 2.8 km SSE past barracks,

Jul 1990, Forster PIF6939 (BRI, MEF); 14.1 km from

Kenilworth, along Booloumba Creek road, Nov 1990,

652

Austrobaileya 7(4): 641-658 (2008)

Fig. 4. Erythrina numerosa. A. leafy branchlet x 0.4. B. single inflorescence bearing buds and flowers x 0.4. C. mature

fruit containing seeds x 0.5. D, E. seed x 3. A, B from Forster PIF7618 (BRI); C-E from Forster PIF7777 (BRI). Del.

W. Smith.

Bean, Revision of Erythrina

Bean 2647 (BRI); Wide Bay, Aug 1843, Leichhardt s.n.

(MEL). Darling Downs District: 2 miles [3 km] S of

Maclagan, Nov 1970, Rawlings s.n. (BRI [AQ199200]).

Moreton District: Commissioners View, c. 10km E of

Blackbutt, Oct 2001, Bean 17998 (BRI); World’s End

Pocket at end of Pine Mountain Road, N of Ipswich, Nov

1981, Bird s.n. (BRI [AQ347049]); World’s End Pocket,

Pine Mountain, Nov 1978, Williams 78249 (BRI); Petrie,

18 miles [29 km] N of Brisbane, Dec 1931, Blake 3071

(BRI); S.F. 1355 Kipper Creek, Nov 1994, Forster

PIF15888 (BRI); Splityard Creek, Wivenhoe Dam, Nov

1990, Forster PIF7618 et al. (BRI); Mt Crosby cliffs,

Brisbane River, Feb 1991, Bird & Collins s.n. (BRI

[AQ547556]); Barnes Hill Pipeline Reserve, Mt Crosby

road, Mt Crosby, Oct 1993, Grimshaw G8 et al. (BRI);

Moggill Scrub, undated, Stuart 238 (MEL); Brisbane

River, Jul 1855, Mueller s.n. (MEL); c. 2 km NE of

Perrys Knob, Mt Marrow area, 4.2 km SSE of Marburg,

Oct 1999, Pollock ABP708 & Leach (BRI); Neumann’s

Lookout, 2 km W of Mount Berryman, Feb 1991, Forster

PIF7777 (BRI, MEL); Pimpama, Mar 1889, Simmonds

115 (BRI); property of A. Schumacher, Milford, SE

of Boonah on Rathdowney road, Nov 1981, Bird s.n.

(BRI [AQ347246]); Boonah - Croftby road, 4.8 km S of

Carney’s Creek crossing, Oct 2000, Bean 16908 (BRI).

New South Wales. North Coast: Acacia Creek, via

Killarney, Dec 1905, Dunn s.n. (NSW); Mt Warning, Jan

1936, Fraser s.n. (NSW); Tyalgum Ridge, Limpinwood

Nature Reserve, c. 25 km WNW of Murwillumbah, Dec

1977, Coveny 9932 & Haegi (BRI, NSW); Bog Onion

road, Mebbin S.F., c. 7 miles [c. 11 km] S of Tyalgum,

Nov 1966, Hayes 2590 et al. (NSW); Mt Mullengen, 4

miles [6 km] E of Ramornie, Jul 1922, Blakely & Shiress

s.n. (NSW).

Distribution and habitat: Erythrina

numerosa is distributed from Nymboida in

New South Wales (Harden et al. 2006) to

Kilkivan in Queensland (Map 5). It grows

in araucarian notophyll vine forest or on its

margins.

Phenology : Flowers are recorded from

October to December; fruits from February

to June.

Notes: Erythrina numerosa differs from

E. vespertilio by the peduncles 3.5-8.5 mm

diameter (vs. 1.5-4.2 for E. vespertilio),

fascicles 1.5-8 mm apart (vs. 8-15), standard

petal 39-54 mm long (vs. 32-40 mm), keel

petals 13.5-15.5 mm long (vs. 10-13), anthers

2.8-4.3 mm long (vs. 2.1—2.8(—3.1) mm), ovary

with 12-16 ovules (vs. 6-8), pods 13-24 cm

long and containing 5-11 seeds (vs. 7-14 cm

long and containing 1—4(—5) seeds). In the

field, it is conspicuously different by virtue

of the smooth, unfurrowed trunk, sometimes

rather shiny; in contrast E. vespertilio has

653

a dull corky, ridged trunk. The orange to

salmon-coloured corolla of E. numerosa is

markedly paler than that of E. vespertilio,

in which the corolla is varies from scarlet to

orange-red.

Etymology: The epithet refers to the numerous

seeds in each pod, in comparison with

Erythrina vespertilio and other Australian

species.

5. *Erythrina caffra Thunb., FI. Cap. Ed.

2, 559 (1823); Duchassaingia caffra (Thunb.)

Walp.,Z/««aea23:741 (1850); Corallodendron

caffrum (Thunb.) Kuntze, Revis. Gen. PI. 1:

172 (1891). Type: South Africa, “e. cap. b.

spei Thunberg”, C.P. Thunberg s.n. (holo:

UPS-Thunb. 16302 [microfiche!]).

Tree 6-10 m high, briefly deciduous. Bark

mostly smooth, with shallow longitudinal

furrows, grey to grey-brown. Trunk and

branches bearing large scattered conical

prickles, terminal branchlets with prickles.

Indumentum comprising minute simple

hairs, confined to apical section of branchlets,

developing leaves, ovaries and young fruits.

Leaves uniformly green, not waxy below.

Petioles unarmed, terete, 50-100 mm long;

petiolule of basal leaflets 4-7 mm long;

terminal leaflet deltoid, rhomboid or broadly

trullate, 55-105 mm long, 50-100 mm wide,

length/breadth ratio 1.04-1.31, apex acute to

acuminate; base obtuse or truncate; lateral

lobes absent; basal leaflets similar but smaller

than the terminal one. Leaflet venation

penninerved, basal pair of lateral veins

converging at point of petiole attachment.

Inflorescences borne on leafless branches, 50-

100-flowered, 10-18 cm long, rachis usually

shorter than peduncle, peduncles 4-5 mm

diameter; fascicles 3-flowered, often two or

three fascicles at same position along rachis,

each fascicle group 3-7 mm apart on rachis,

pedicels 5-8mmlongatanthesis; calyx tubular

in bud, 9-12 mm long, truncate or with minute

lobes, splitting apically as corolla expands, at

anthesis shortly bilabiate; all petals orange to

scarlet, prominently veined; standard 35-55

x 14-28 mm, apex obtuse, strongly recurved

and exposing staminal column, wings and

keel similar, wings 13-24 mm long; keel

petals 12-21 mm long, fused, obtuse; stamens

654

38-50 mm long, anthers 2.3-4.3 mm long,

yellow to brown; ovary hairy, ovules 8-10.

Pods not strongly curved when mature, 6-10

cm long, 0.9-1.2 cm wide at widest point,

conspicuously narrowed between the seeds,

outer surface smooth, dehiscent, seeds easily

visible, glabrous at maturity. Seeds 2-4,

ellipsoidal, 6-8 mm long, uniform in colour,

red, with a dark hilum. Kaffir coral tree. Fig.

Additional specimens examined : Queensland.

Moreton District: SW side of Mt Coot-tha, at Chapel

Hill Environmental Education Hut, off Fleming road,

Sep 1998, Stephens GBS12 (BRI); Forestry Road, just

past lookout, Mt Coot-tha, Sep 1999, Boyle TPB50 &

Lahey (BRI).

Distribution and habitat : Erythrina caffra is

sparingly naturalised around Brisbane (Map

1), where it has been found on hillsides in

woodland dominated by Eucalyptus spp. It is

also recorded as naturalised on Norfolk Island

(Green 1994).

Phenology : Flowers recorded for May and

September; fruits in November.

Notes : Erythrina caffra originated in South

Africa, in coastal forests of the Eastern Cape

and KwaZulu-Natal (Hennessey 1991). It has

been cultivated in Brisbane since at least 1933

(White 9026 , BRI).

Etymology : The species name caffra refers

to Kaffraria, an old name for a part of the

Eastern Cape region of Africa.

6. *Erythrina x sykesii Barneby & Krukoff,

Lloydia 37: 447 (1974). Type: Queensland,

cultivated at Brisbane Botanic Gardens, 12

June 1933, C.T.White 9073 (holo: NY; iso:

BRI).

Tree 2.5-10 m high, deciduous. Bark mostly

smooth, with pale longitudinal fissures,

grey to brown. Trunk and branches with

prickles, terminal branchlets with or without

prickles. Indumentum comprising minute

simple hairs, confined to apical section of

branchlets, developing leaves and ovaries.

Leaves uniformly green, not waxy below.

Petioles unarmed, terete, 80-170 mm long;

petiolule of basal leaflets 6-10 mm long;

terminal leaflet deltoid to rhomboid, 95-180

mm long, 80-230 mm wide, length/breadth

ratio 0.78-1.16, apex acuminate or acute; base

Austrobaileya 7(4): 641-658 (2008)

obtuse to broadly cuneate. Leaflet venation

penninerved, basal pair of lateral veins

converging at point of petiole attachment (i.e.

3-veined at base). Inflorescences borne on

leafless branches, 30-100-flowered, 17-30 cm

long, rachis and peduncles about same length,

peduncles 4-6 mm diameter; fascicles 3-

flowered, often two or three fascicles at same

position along rachis, each fascicle group 1-8

mm apart on rachis, pedicels 4-6 mm long

at anthesis; calyx tubular in bud, 9-12 mm

long, 2-lipped and 14-18 mm long at anthesis;

petals red, prominently veined; standard 48-

64 x 18-24 mm, apex acute, slightly recurved,

staminal column exposed, wings and keel

similar, wings 20-28 mm long; keel petals

18-27 mm long, fused to each other for most

of their length, obtuse; stamens 55-60 mm

long, anthers 2.8-4.5 mm long, yellow; ovary

hairy, ovules 11-13. Fruits do not develop.

Thorny coral tree. Fig. 2D.

Additional specimens examined: Queensland. Cook

District: Bromfield Crater, Atherton Tableland, Jul

1966, Brass 33486 (BRI); Egan Creek L.A., SW corner,

T.R. 756, Parish of Jordan, Nov 1958, Smith 10478 (BRI).

North Kennedy District: Herberton, north Queensland

Tablelands, Feb 2003, Batianoff 0302118 & Batianoff

(BRI); North Cedar Creek, Ravenshoe, Feb 2003,

Batianoff0302119 & Batianoff ( BRI). Moreton District:

Nambour, Apr 1970, O’Mara 104 (BRI); Golden Beach

Caloundra, Sunshine Coast, Aug 1998, Batianoff98083

& Batianoff (BRI); North Stradbroke Island, Amity

Point, Kindaria Street, Aug 2003, Batianoff 030899

& Boyle (BRI); Curtis road, Mt Tamborine, Sep 2000,

Bean 16823 (BRI). New South Wales. North Coast:

Fingal Head, Jul 1998, Batianoff 980712 (BRI, MET);

Byron Bay, Cape Byron main beach, Jul 1998, Batianoff

980721 (BRI); Richmond River, Casino, Jul 2004, Bean

22676 (BRI); Valla, 20 km N of Nambucca Heads and

50 km S of Coffs Harbour, Dec 2004, Hall s.n. (BRI

[AQ612871]); Woolooware Bay, Cronulla, on Kurnell

road, Jul 1976, Coveny 7753 & Hind (NSW); ruins of

Ellamatta Homestead, Reddells road, c. 5 km W of

Kembla Grange, Jul 1996, Jobson 4316 (NSW).

Distribution and habitat : Erythrina x sykesii

is naturalised in New Zealand (Webb

et al. 1988) and Australia. In Australia,

naturalisations are known from some coastal

parts of Qld and N.S.W. (including Lord Howe

Island), and around Perth (Paczkowska &

Chapman 2000) in W.A. (Map 9). It inhabits

cleared or disturbed places where the soil is

frequently moist, particularly near creeks.

Bean, Revision of Erythrina

Phenology : Flowers are recorded from July

to November.

Notes : The parentage of Erythrina x sykesii

is unknown. Krukoff & Barneby (1974)

suggested that the parents could be E.

coralloides DC. and E. lysistemon Hutch.

This suggestion was repeated by Spencer

(2002). Hennessy (1991) gave E. speciosa

Andr. and E. caffra as the probable parents,

and cited morphological features allying it to

those species. The origin of E. x sykesii is also

unknown but it has been cultivated for several

decades in both Australia and New Zealand

(Spencer 2002). It does not produce fruits,

but vegetative pieces, even sections of trunk,

will sprout roots readily (Esler & Edgar 1997;

pers. obs.).

Etymology: Named for W.R. (Bill) Sykes, a

New Zealand botanist.

7. *Erythrina crista-galli L., Mant. PI. 99

(1767); Micropteryx crista-galli (L.) Walp.,

Linnaea 23: 740 (1840); Corallodendron

crista-galli (L.) Kuntze, Rev is. Gen. PI. 172

(1891). Type: Brazil, undated, D. Vandelli

s.n. (lecto: LINN, Cat. no. 888.4 fide Howard

(1988)).

Illustration: Whistler & Elevitch (2006:

332).

Evergreen tree 4-8 m high, trunk and branches

without prickles, terminal branchlets without

prickles. Bark corky, deeply furrowed, grey

to brown. Indumentum comprising minute

simple hairs, confined to apical section of

branchlets, developing leaves, ovaries and

young fruits. Leaves uniformly green, minutely

reticulately waxy below. Petioles frequently

bearing 1 or 2 recurved prickles, terete, 60-

140 mm long; petiolule of basal leaflets 8-13

mm long; terminal leaflet 58-130 mm long,

25-77 mm wide, length/breadth ratio 1.7-2.3,

apex acute, base cuneate, lateral lobes absent;

basal leaflets identical to, or slightly smaller

than the terminal one. Leaflet venation

penninerved, without a basal pair of lateral

veins. Inflorescences in leaf axils or at end of

leafy branchlets, 12-80-flowered, 10-40 cm

long, rachis and peduncle not distinguishable;

fascicles 2-3- flowered, one or sometimes two

fascicles at any position along branchlet, each

fascicle (group) 2-32 mm apart on branchlet,

655

pedicels 13-25 mm long at anthesis; calyx

broadly campanulate in bud, 9-13 mm long,

truncate with two small deciduous teeth, not

splitting longitudinally as corolla expands;

petals prominently veined; standard pink, red

or crimson on inner surface, pink on outer

surface, broadly-elliptic, 40-55 x 34-46 mm,

apex obtuse or emarginate, strongly recurved

and exposing staminal column, wings and

keel very dissimilar, wings white, 9-19 mm

long, wholly or largely obscured by calyx,

obtuse; keel petals pink, red or crimson, 30-

45 mm long, fused, incurved, enclosing much

of staminal column, acute; stamens 33-55

mm long; anthers 1.9-2.7 mm long, yellow;

ovary hairy, ovules 16-21. Pods glabrous at

maturity, 14-27 cm long, 1.3-1.6 cm wide at

widest point, somewhat narrowed between the

seeds, outer surface smooth, pods dehiscent

and slightly curved when mature with seeds

easily visible. Seeds 2-6, ellipsoidal, 12-16

mm long, not uniform in colour, black with

brown streaks, with a pale hilum. Cockspur

coral tree, Cock’s comb coral tree. Fig. ID

Additional specimens examined : Queensland. Wide

Bay District: Tozer Gully, Cootharaba road, Gympie,

Nov 2000, Bean 17044 (BRI). Moreton District:

intersection of Rode and Webster roads, Stafford

Heights, Nov 1999, Wrench s.n. (BRI [AQ665547]);

Cannon Hill College, Cannon Hill, 7 km E of Brisbane

GPO, Oct 2000, Bean 16979 (BRI, NSW); Brisbane

River, Moggill Ferry Reserve, Sep 2000, Batianoff

200816 (BRI); Paradise Road, Willawong, c. 15 km S of

Brisbane CBD, May 2002, Bean 19028 (BRI); tributary

of Blunder Creek, Richlands, Brisbane, Oct 1997, Hall

& Bird s.n. (BRI [AQ659717]); Greenbank, Brisbane,

Nov 1991, Robins s.n. (BRI [AQ628572]). New South

Wales. North Coast: 5 km SW of Lismore, Mar 1984,

Swarbrick 7404 (BRI); Wilson’s River floodplain, beside

Bruxner Highway, c. 3 km SW of Lismore, Oct 1990,

Smith JMBS935 (NSW); Casino, Jan 1991, Clarke s.n.

(NSW).

Distribution and habitat: Erythrina crista-

galli is native to Argentina, Brazil, Bolivia,

Paraguay and Uruguay (Krukoff & Barneby

1974). It has been widely cultivated in

tropical and sub-tropical parts of the world,

including South Africa, California (U.S.A.)

and Australia. In Australia it has become

naturalised in coastal Qld and N.S.W. (Map

2) (listed as a noxious weed in the latter

state), where it inhabits sunny swampy areas,

including areas where the groundwater is

brackish.

656

Phenology : Flowers have been recorded for

nearly every month of the year; fruits (rarely

collected) recorded in May, November and

December.

Notes : This species has been cultivated in

Australia since at least 1924 ( White 2400,

BRI). It is the national flower of Argentina

(Krukoff & Barneby 1974).

Etymology : The epithet is derived from the

Latin, and means ‘cock’s comb’, almost

certainly in reference to the linear fan-shaped

arrangement of the stamens.

8. Erythrina fusca Lour., FI. Cochinch.

427 (1790); Corallodendron fuscum (Lour.)

Kuntze, Revis. Gen. PI. 1: 173 (1891).

Types: Cochinchina, Louriero s.n. (syn: not

found); illustration of ‘ Gelala aquatictf in

Rumphius, Herb. Amboin., 2: 235, t. 78 (syn:

the illustration).

Evergreen tree to 23 m high. Bark ± smooth,

though cracked and with large pustular

lenticels, olive-brown. Trunk and branches

with or without prickles, terminal branchlets

with prickles. Indumentum comprising

minute simple hairs, confined to apical section

of branchlets, developing leaves, ovaries and

young fruits. Leaves uniformly green, not

waxy below. Petioles unarmed, terete, 50-100

mm long; petiolule of basal leaflets 7-10 mm

long; terminal leaflet 70-120 mm long, 43-76

mm wide, length/breadth ratio 1.58-1.81, apex

acute; base obtuse to cuneate; lateral lobes

absent; basal leaflets similar but smaller than

the terminal one. Leafletvenation penninerved,

basal pair of lateral veins converging at point

of petiole attachment. Inflorescences borne

on leafless branches, 18-30-flowered, 10-26

cm long, rachis and peduncles about same

length, peduncles 2.5-4.5 mm diameter;

fascicles 3-flowered, each fascicle group 6-25

mm apart on rachis, pedicels 5-13 mm long

at anthesis; calyx tubular in bud, 13-16 mm

long, truncate or apiculate, a longitudinal split

developing as corolla expands, campanulate

and asymmetrical at anthesis; all petals

prominently veined; standard 38-52 x 25-30

mm, yellow to orange, apex obtuse, strongly

recurved and exposing staminal column;

wings and keel similar; wings 18-22 mm

long, creamy-white proximally, maroon

Austrobaileya 7(4): 641-658 (2008)

distally; keel petals 20-27 mm long, creamy-

white, fused, obtuse; stamens 33-40 mm

long, anthers 2.2-37 mm long, yellow; ovary

hairy, ovule number unknown. Pods glabrous

at maturity, 11-20 cm long, 1.2-1.5 cm wide

at widest point, slightly narrowed between the

seeds, outer surface smooth, pods dehiscent

and straight to slightly curved when mature.

Seeds 6-12, ellipsoidal, 12-18 mm long,

uniform in colour, dark brown. Coral tree.

Fig. 2C.

Additional specimens examined : Queensland. Cook

District: Daintree Mission, Sep 1948, Smith 4036

(BRI); Daintree River, below the township of Daintree,

Oct 2000, Morris s.n. (BRI [AQ497997]).

Distribution and habitat : Erythrina fusca

is regarded as the most widespread species

of the genus, indigenous to coastal areas of

the tropics where annual rainfall is high,

including eastern Africa, India, Malesia, many

Pacific islands and western South America. In

Australia, this species is apparently confined

to the Daintree area of north Qld (Map 3),

where the annual rainfall exceeds 2500 mm

and the dominant vegetation type is lowland

rainforest.

Phenology : Flowers recorded in September;

fruits in October.

Notes : Erythrina fusca has not previously

been reported as occurring in Australia,

despite Smith’s collection from the Daintree

area in 1948. Apparently it was previously

considered that the collection was made from

a cultivated specimen. The second collection

(cited above) has confirmed the non-cultivated

status of E. fusca in Australia. However its

origin status in Australia (indigenous or alien)

remains unclear. Only one tree is currently

known to local naturalists.

Drift seeds of Erythrina fusca (but not

plants) have been recorded from islands of

the Capricorn - Bunker group off the coast

of Gladstone. These are thought to originate

from islands of the Pacific (Smith 1992).

Etymology : From the Latin fuscus meaning

‘dark, dusky’. This perhaps alludes to the very

dark-coloured seeds of this species.

657

Bean, Revision of Erythrina

Excluded names

Erythrina acanthocarpa E.Mey.

This species was recorded as perhaps

naturalised in South Australia (Jessop &

Toelken 1986), but according to R. Barker

(pers. comm. 2007), a recent re-examination

of the relevant specimen revealed some

follow-up documentation to indicate that it

was not established.

Erythrina lysistemon Hutch.

Krukoff (1972) identified the leaves of

the type collection of E. phlehocarpa as

belonging to E. lysistemon. This is refuted

here (see discussion under E. variegata). As

this was the only non-cultivated record of

E. lysistemon for Australia, it is excluded

from this account.

Acknowledgements

I am grateful to Paul Forster and Ailsa Holland

for advice and documentation regarding

Erythrina numerosa, and Peter Bostock for

the Latin diagnosis and distribution maps.

Ellen Terrell and Dan Irby sent information

about the occurrence of E. fusca at Daintree.

Will Smith and Brian Connell provided the

illustrations.

References

Adema, F. (1996). Notes on Malesian Fabaceae

(Legummosae-Papilionoideae), 1. The genus

Erythrina L. Blumea 41: 463-468.

Bentham, G. (1864). Erythrina. In Flora Australiensis 2:

252-254. L. Reeve & Co.: London.

Bostock, RD. & Holland, A.E. (eds.) (2007). Census

of the Queensland Flora 2007. Queensland

Herbarium, Environmental Protection Agency:

Brisbane.

Britton, N.L. (1924). Scientific Survey of Porto Rico

& the Virgin Islands , Volume 5. New York

Academy of Sciences: New York.

Brock, J. (1988). Top End Native Plants. J. Brock:

Darwin.

Card, M.A. (2001). Vegetation dynamics of Milman

Island, northern Great Barrier Reef

Australia. PhD thesis, James Cook University:

Townsville.

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

Australian Tropical Rainforest. Nokomis

Editions Pty Ltd: Melbourne.

Du Puy, D.J. & Telford, I.R.H. (1993). Fabaceae. In

H.J. Hewson & H.S. Thompson (eds). Flora

of Australia , Oceanic islands 2, 50: 204-36.

Australian Government Publishing Service:

Canberra.

Esler, A. & Edgar, E. (1997). Erythrina x sykesii.

Auckland Botanical Society Journal 52: 39-42.

Green, PS. (1994). Fabaceae. In A.G.S. Wilson (ed.).

Flora of Australia , Oceanic Islands 1, 49:

172-197. Australian Government Publishing

Service: Canberra.

Harden, G. J., Mcdonald, W.J.F. & Williams, J.B. (2006).

Rainforest Trees and Shrubs - A field guide to

their identification. Gwen Harden Publishing:

Nambucca Heads, New South Wales.

Hennessy, E.F.F. (1991). Erythrineae in southern Africa.

Bothalia 21: 1-25.

Hickey, M. & King, C. (2000). The Cambridge illustrated

glossary of Botanical terms. Cambridge

University Press: Cambridge.

Howard, R.A. (1988). Flora of the Lesser Antilles,

Leeward and Windward Islands, Volume 4.

Arnold Arboretum: Cambridge.

Jessop, J.P & Toelken, H.R. (1986). Flora of South

Australia Part II Leguminosae-Rubiaceae.

South Australian Government Printing

Division: Adelaide.

Krukoff, B.A. (1972). Notes on Asiatic-Polynesian-

Australian species of Erythrina , II. Journal of

the Arnold Arboretum 53: 128-139.

- (1982). Notes on the species of Erythrina. XVIII.

Allertonia 3: 121-138.

Krukoff, B.A. & Barneby, R.C. (1974). Conspectus of

species of the genus Erythrina. Lloydia 37:

332-459.

Merrill, E.D. (1917). An Interpretation of Rumphius’s

Herbarium Amboinense. Manila Bureau of

Printing: Manila.

Milson, J. (2000). Trees and shrubs of north-west

Queensland. Queensland Department of

Primary Industries: Brisbane.

Nicholson, N. & Nicholson, H. (1991). Australian

Rainforest Plants III. Terania Rainforest

Nursery: The Channon, New South Wales.

Paczkowska, G. & Chapman, A.R. (2000). The Western

Australian Flora, a descriptive catalogue. The

Wildflower Society of Western Australia (Inc.),

the Western Australian Herbarium, CALM

and the Botanic Gardens and Parks Authority:

Perth.

Smith, J.M.B. (1992). Patterns of disseminule dispersal

by drift in the southern Coral Sea. New Zealand

Journal of Botany 30: 57-67.

658

Spencer, R. (2002). Erythrina L. In R.D. Spencer (ed.),

Horticultural Flora of South-eastern Australia

3: 198-201. University of New South Wales

Press: Sydney.

Verdcourt, B. (1979). A Manual of New Guinea Legumes.

Office of Forests, Division of Botany: Lae.

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

Flora of New Zealand , Volume 4, Naturalised

pteridophytes, gymnosperms, dicotyledons.

Botany Division DSIR: Christchurch.

Austrobaileya 7(4): 641-658 (2008)

Whistler, W. & Elevitch, C.R. (2006). Erythrina

variegata (coral tree). In C.R. Elevitch (ed).

Traditional trees of Pacific Islands: their

culture, environment and use, pp. 329-343.

Permanent Agriculture Resources: Hawaii.

Williams, K.A.W. (1979). Native Plants of Queensland.

Volume 1. K.A.W. Williams: North Ipswich,

Australia.

Distribution of Erythrina spp. Maps 1-9.

Map 1. E. caffra. Map 2. E. crista-galli. Map 3. E.fusca. Map 4. E. insularis. Map 5. E. numerosa. Map 6. E. variegata.

Map 7. E. vespertilio subsp. biloba. Map 8. E. vespertilio subsp. vespertilio. Map 9. E. x sykesii.

A remarkable new species of Rimularia Nyl. (lichenized

fungi: Trapeliaceae) from tropical Australia

Gintaras Kantvilas 1 , Patrick McCarthy 2 & Benjamin Stuckey 3

Summary

Kantvilas, G., McCarthy, P.M. & Stuckey, B. (2008). A remarkable new species of Rimularia Nyl.

(lichenized fungi: Trapeliaceae) from tropical Australia. Austrobaileya 7(4): 659-663. The new

species Rimularia cerebriformis Kantvilas is described, based on a collection and field observations

from dry eucalypt woodland in the Northern Territory, Australia. It represents an unusual occurrence

of a genus that occurs mostly in moist environments in cool to cold, temperate latitudes of both

hemispheres. Relationships with other species of the genus are discussed briefly.

Key Words: lichens, Rimularia cerebriformis , Trapeliaceae, Northern Territory, Australia

'G.Kantvilas, Tasmanian Herbarium, Private Bag <

2 P.M.McCarthy, Australian Biological Resources

Australia

3 B.Stuckey, Northern Territory Herbarium, PO.

Australia

Introduction

Rimularia Nyl. is a genus of crustose lichens,

widespread in cool to cold temperate areas

of both hemispheres. General accounts of

the genus are provided by Hertel & Rambold

(1990) and Lumbsch (1997) and, although

many additional species have been described

since that time, the delimitation of the genus

has remained generally unchanged. It is

characterised by the presence of a Chlorella-

type photobiont, lecideine apothecia with

a well-developed, carbonised, cupulate

excipulum, branched and anastomosing

paraphyses, simple, non-halonate ascospores

and, in particular, eight-spored asci of the

Rimularia-typQ in the sense of Hertel &

Rambold (1990) and Hafellner (1984); that

is, with a well-developed tholus with amyloid

flanks and an amyloid cap, and generally

weakly amyloid internal structures. Species

of Rimularia occur on rocks, bark, soil,

bryophytes and other lichens.

Ten species of Rimularia have been

recorded for Australia (McCarthy 2007), seven

of which are found in Tasmania (Coppins

& Kantvilas 2001; Kantvilas & Elix 2007),

highlighting the predeliction of the genus

for cool, moist environments. Indeed, of the

approximately 25 known species in the world,

Accepted for publication 10 January 2008

t, Hobart, Tasmania 7001, Australia

Study, G.P.O. Box 787, Canberra, A C T. 2601,

Box 496, Palmerston, Northern Territory 0831,

only two, R. globulispora from New Guinea

(Aptroot & Sipman 1991) and R. gyromuscosa

from Taiwan (Aptroot & Sparrius 2003) occur

in low latitudes. However, both of these are

restricted to high elevations; the former is also

known from Tasmania (Coppins & Kantvilas

2001). Hence the discovery of a species in

tropical Australia, growing on exposed rocks

in a lowland, very harsh environment, can be

regarded as very unusual. This species is new

to science and is described below.

Materials and methods

The study is based on collections and

observations made by the authors, and on

comparative material in herbaria, chiefly in

the Tasmanian Herbarium (HO). Anatomical

and morphological observations were

undertaken using light microscopy, with thin

hand-cut sections mounted in water, 10%

KOH, Lactophenol Cotton Blue, and Lugol’s

Iodine after pretreatment with KOH (the last

indicated as K/I). Ascospore measurements

are presented in the format: smallest

measurement-meaw-largest measurement,

and are based on 60 observations. Ascus

measurements are presented in the same

way but, owing to the paucity of material,

are based on 20 observations only. Chemical

composition was determined by thin-layer

chromatography using standard methods

(Orange et al. 2001).

660

Taxonomy

Rimularia cerebriformis Kantvilas species

nova apotheciis intense gyrosis et igitur

Rimulariae gyrizanti R. gyromuscosaeque

optime congruens sed thallo inconspicuo,

acidum gyrophoricum continent^ ascosporis

10-19 pm longis, 5-10 pm latis differens.

Typus: Northern Territory. Edith Falls,

14°12'S, 132°13'E, on exposed sandstone

outcrops in river gorge, 22 September 2007,

G.Kantvilas 291/07 (holo: HO; iso: DNA).

Thallus inapparent, very thin and

discontinuous. Photobiont cells very sparse

and limited to the base of the apothecia,

Chlorella-Mke , with individual cells globose

to rhomboid, mostly 6-8 pm wide, occurring

singly or in pairs, surrounded by a gelatinous

sheath; colonies of unidentified cyanobacteria

also present. Apothecia numerous and

scattered, 0.5-0.8 mm wide, jet-black, strongly

convex to subglobose, broadly adnate at the

base, with disc intensely gyrose-contorted and

margin indistinct and mostly camouflaged by

the gyrae of the disc; in section comprising

a mass of amorphous, opaque dark brown,

sterile excipular tissue to c. 500 pm high,

imbedded in the upper part with ‘strands’

or Tocules’ of chiefly colourless hymenial

tissue, 60-80 pm high and 50-80 pm wide.

Excipulum in section K+ olive brownish at

the edges, comprising irregularly roundish

cells to 10 pm wide. Hymenium colourless

to pale brownish in the uppermost part, K/I+

pale blue; asci eight-spored, broadly clavate,

40-47. <5-55(-62) x (13-)15-77.S-22 pm, of

the typical Rimularia- type (sensn Hafellner

1984); paraphyses rather dense, frequently

branched and anastomosing, robust, 1-3 pm

thick, frequently rather moniliform and of

variable thickness; apices neither capitate

nor pigmented. Ascospores ovate, broadly

ellipsoid to oblong, hyaline to occasionally

slightly pale brownish, 10-73.6-19 x (5-)6-

7.3-9(-10) pm. Pyncidia not observed. Fig. 1

& 2 .

Chemistry : gyrophoric acid (major) and

lecanoric acid (minor) detected by thin-layer

chromatography of the apothecia.

Distribution and habitat : The new species

is known only from the type locality in the

Austrobaileya 7(4): 659-663 (2008)

upper reaches of a gorge cut by the Edith

River. It grows on coarse, quartz sandstone

associated with the Kombolgie Sub Group,

formed between 1730 and 1822 mya (Kruse

et al. 1994). The site is very exposed and,

although protected from seasonal burning

and floodwaters, may be subject to the

microclimatic effects of the waterfall nearby

during months of heavy rainfall. Vegetation

in the surrounding area included a sparse mix

of riparian and lithophytic species including

Xanthostemon eucalyptoides F.Muell., Ficus

atricha D.J.Dixon, Syzigium eucalyptoides

(F.Muell.) B.Hyland subsp. eucalyptoides,

Cymbopogonprocerus R.Br. and species from

the broader, low open woodland communities

dominated by Corymbia K.D.Hill &

L.A.S.Johnson, Eucalyptus L’Herit., Acacia

Mill, and Triodia R.Br. The occurrence of

lichens at the locality is extremely sporadic.

The surfaces of rock exposed to direct

sunlight are almost bare, and even Rimularia

cerebriformis itself appears to be very

localised. More sheltered, shaded surfaces,

often within clefts or beneath underhangs,

are also very poorly colonised, although

an unidentified species of Buellia was not

uncommon.

Leliyn (Edith Falls) is situated in the

wet-dry tropics of northern Australia and

experiences year round high temperatures.

The average maximum and minimum

temperatures for Katherine, c. 50 km to

the south, are 34°C and 20°C respectively;

the type locality could expect even higher

temperatures. The area receives a majority of

its annual rainfall (1282 mm) during heavy

monsoonal rains between December and

March.

Notes : This species is detected in the

field solely by the presence of its jet-black

apothecia (Fig. 1), and even under low-

power magnification, no thallus is visible.

Scattered dull olive-brown to olive-green,

irregular granules, nestling amongst the

coarse grains of the substratum in the vicinity

of the apothecia, belong to unidentified

cyanobacteria. The degree of penetration of

the lichen hyphae into the rock substratum,

as determined by discoloration of the latter,

appears to be limited to about the outermost

0.5 mm.

Kantvilas et al., Rimularia cerebriformis

661

Fig. 1. Rimularia cerebriformis habit. Scale = 0.5 mm.

The gyrose apothecia of the new species

align it to Rimularia gyrizans (Nyl.) Hertel

& Rambold, which is widespread in cool

temperate areas of Europe and North America,

to R. mullensis (Stirt.) Coppins, endemic to

Great Britain, and to R. gyromuscosa Aptroot,

endemic to subalpine Taiwan. These species

differ from R. cerebriformis mainly by having

a well-developed thallus. There are also

chemical differences: the first contains stictic

(major) and norstictic (minor) acids, the second

contains norstictic acid whereas the last lacks

lichen substances. In addition, the ascospores

of all of these species are markedly smaller

than those of R. cerebriformis'. 8-11 x 4.7-8

pm in R. gyrizans and (6-) 8-11 x 4-7 pm in

R. mullensis (Hawksworth & Coppins 1992),

and 10-12.5 x 5-7 pm in R. gyromuscosa

(Aptroot & Sparrius 2003).

Amongst the Australian species of

Rimularia , Rimularia cerebriformis is easily

distinguished by its gyrose apothecia.

There is one taxon of as yet undetermined

identity in alpine Tasmania that has similar

apothecia, but this has a well-developed

thallus containing norstictic acid. Gyrophoric

acid is a relatively common substance in

Rimularia , and amongst the Australian

species, is also known from R. australis

Hertel & Rambold, R. exigua Hertel &

Rambold, R. insularis (Nyl.) Rambold &

Hertel and R. ramboldiana Kantvilas &

Elix; none of these species could in any way

be confused with R. cerebriformis. Indeed

the most superficially similar lichen known

to us is Stephanocyclos henssenianus Hertel,

a predominantly subantarctic, saxicolous

lichen also known from Tasmania. That

species lacks a conspicuous thallus and has

prominent, subglobose, gyrose apothecia.

However, it also has unequivocally Porpidia-

type asci and belongs in the Porpidiaceae.

662

Austrobaileya 7(4): 659-663 (2008)

Fig. 2. Rimularia cerebriformis anatomy. A. asci, paraphyses and ascospores, with amyloid tissues stippled. Scale

=20 pm. B. vertical section through apothecium (schematic), showing carbonised, massive excipulum and ‘locules’ of

hymenial tissue. Scale = 100 pm.

Acknowledgements

We thank Prof. Jack Elix for confirming the

chemical composition of the new species and

Dr Jean Jarman for preparing Figure 2 for

publication. The companionship of colleagues

from the CHAH-2007 field trip in the Northern

Territory is cheerfully acknowledged.

References

Aptroot, A. & Sipman, H.J.M. (1991). New lichens and

lichen records from New Guinea. Willdenowia

20: 221-256.

Aptroot, A. & Sparrius, L.B. (2003). New microlichens

from Taiwan. Fungal Diversity 14: 1-50.

Coppins, B. J. & Kantvilas, G. (2001). Four new species

of Rimularia Nyl. (Agyriaceae). Bibliotheca

Lichenologica 78: 35-48.

Hafellner, J. (1984). Studien in Richtung einer

naturlicheren Gliederung der Sammelfamilien

Lecanoraceae und Lecideaceae. Beiheft zur

Nova Hedwigia 79: 241-371.

Hawksworth, D.L. & Coppins, B.J. (1992). Lecidea

Ach. (1803). In O.W.Purvis, B.J.Coppins,

D.L.Hawksworth, RW.James & D.M.Moore

(eds ). The Lichen Flora of Great Britain and

Ireland, pp. 318-336. British Lichen Society:

London.

Hertel, H. & Rambold, G. (1990). Zur Kenntnis

der Familie Rimulariaceae (Lecanorales).

Bibliotheca Lichenologica 38: 145-189.

Kantvilas, G. & Elix, J.A. (2007). Additions to the

lichen family Agyriaceae Corda from Tasmania.

Bibliotheca Lichenologica 95: 317-333.

Kruse, P.D., Sweet, I.P, Stuart-Smith, P.G., Wygralak,

A.S., Pieters, PE. & Crick, I.H. (1994).

Katherine SD/53-9 1:250 000, Geological Map

Series. Explanatory Notes Northern Territory

Geological Survey.

Lumbsch, H.T. (1997). Systematic studies in the sub¬

order Agyriineae (Lecanorales). Journal of the

Hattori Botanical Laboratory 83: 1-73.

663

Kantvilas el al ., Rimularia cerebriformis

McCarthy, P.M. (2007). Checklist of the Lichens of

Australia and its Island Territories. Australian

Biological Resources Study, Canberra. Version

13 November 2007. http:/www.anbg.gov.au/

abrs/lichenlist/introduction.html

Orange, A., James, P.W. & White, F.J. (2001).

Microchemical Methods for the Identification

of Lichens. British Lichen Society: London.

A new subsection and two new subseries for Boronia

Sm. section Valvatae (Benth.) Engl. (Rutaceae)

Marco E Duretto

Summary

Duretto, M.F. (2008). A new subsection and two new subseries for Boronia Sm. section Valvatae

(Benth.) Engl. (Rutaceae). Austrobaileya 7(4): 665-668. Three infrageneric taxa are newly described

for Boronia Sm. section Valvatae (Benth.) Engl.: these being B. subsection Anomalae Duretto, and

B. subseries Lanceolatae Duretto and B. subseries Rosmarinifoliae Duretto of B. series Valvatae.

Descriptions and notes on the distribution and species composition of each new taxon and B. subseries

Valvatae are provided.

Key Words: Rutaceae, Boronia, Boronia section Valvatae, Boronia subsection Anomalae, Boronia

series Valvatae, Boronia subseries Valvatae, Boronia subseries Lanceolatae, Boronia subseries

Rosmarinifoliae

M.F.Duretto, Tasmanian Herbarium, Tasmanian Museum and Art Gallery, Private Bag 4, Hobart,

Tasmania 7001, Australia. Email: marco.duretto@tmag.tas.gov.au

Introduction

Boronia Sm. section Valvatae (Benth.)

Engl, sensu lato was the subject of cladistic

analyses using morphological and anatomical

characters that resulted in the publication of

the first infrageneric classification for the

section (Duretto & Ladiges 1999; Duretto

1999). In those studies a number of taxonomic

issues were identified and some taxa were

treated as incertae sedis and a number of

species and species-groups were informally

identified. In anticipation of the Flora of

Australia account of the genus these informal

taxa are being formally described, where

appropriate, or placed in synonomy (see also

Duretto accepted), and the position of one of

the taxa treated as incertae sedis is resolved.

Boronia anomala Duretto was treated

as incertae sedis in B. section Valvatae as

its taxonomic relationships with the four

formally recognised subsections were unclear

(Duretto 1999). A new subsection is described

here to accommodate it. The main reason for

describing a new subsection for this species

is to have the formal taxonomy reflect the

current understanding of the relationships

of species in the section. Boronia anomala

is an unusual species and though it appears

to be more closely related to B. subsection

Accepted for publication 2 June 2008

Grandisepalae Duretto (NW Australia) this

is not certain (Duretto 1999) and requires

further investigation. The current formal

classification of B. section Valvatae will now

reflect current understanding of the group and

taxonomically isolated taxa should be easily

identifiable by being members of higher

taxonomic taxa, in this case a subsection.

Species that are the sole representatives

of higher taxonomic groups have a high

conservation value (see Humphries et al.

1995) and this information is more easily

extracted from a formal classification than it

is from informal and/or uncertain ones. The

formal description of an infrageneric taxon

to accommodate B. anomala will hopefully

prompt further research on this species.

One of the larger clades identified by

Duretto & Ladiges (1999) and Duretto (1999)

was Boronia series Valvatae which contains

24 species. Support for the nodes within

B. series Valvatae was weak and the entire

clade was susceptible to collapse if taxa and/

or characters were deleted and/or added to the

cladistic dataset (Duretto & Ladiges 1999;

Duretto 1999; Duretto et al. 2004). With this

in mind, a cautious approach was adopted

and the four main clades in B. series Valvatae

were treated as informal species-groups: viz.

the B. alulata species-group, the B. foetida

species-group, the B. lanceolata species-

666

group and the B. rosmarinifolia species-group

(Duretto & Ladiges 1999; Duretto 1999).

These groups remained stable in later analyses

with the addition of new taxa (Duretto 2003;

Duretto et al. 2004). The B. foetida and the

B. lanceolata species-groups, with additional

collections and a re-evaluation of their

morphological descriptions, have proved to be

indistinguishable and are thus combined.

These three informal species-groups

are here formally described as subseries.

This is being done firstly, because they are

higher taxa worthy of formal recognition and

in hindsight the approach adopted in 1999

was overly cautious, and secondly, a formal

taxonomy is more stable and of greater value

compared to an informal classification which

can become problematic in the long term and

invariably does not become adopted. The

Boronia alulata species-group contains the

type species of B. section Valvatae , B. alulata

Sol. ex Benth., and becomes B. subseries

Valvatae.

Three species in Boronia series Valvatae ,

B. chartacea P.H.Weston, B. hapalophylla

Duretto, F.J.Edwards & P.G.Edwards, and

B. ledifolia (Vent.) DC., fall outside these

new subseries and are treated as incertae

sedis (Duretto & Ladiges 1999; Duretto

1999; Duretto et al. 2004). Further research

using additional data (especially molecular)

is required to elucidate relationships of these

taxa.

Taxonomy

Boronia section Valvatae (Benth.)Engl., Nat.

Pflanzen. 3(4): 135 (1896); Boronia series

Valvatae Benth., FI. Austral. 1: 308, 311

(1863).

Type species: Boronia alulata Benth., fide.

Duretto (1999: 56).

The section is classified into five subsections:

subsection Anomalae Duretto (described

below, northern Western Australia (N WA)),

subsection Bowmaniorum Duretto (Cape

York, northern Queensland (Qld)), subsection

Grandisepalae Duretto (N WA, Northern

Territory (NT), north-west Qld), subsection

Ternatae Duretto (south-western WA) and

Austrobaileya 7(4): 665-668 (2008)

subsection Valvatae (NT, Qld, New South

Wales (NSW), Victoria) (see Duretto 1999 for

full descriptions).

Boronia subsection Anomalae Duretto,

subsectio nova a subsectione typica pilis non

stellatis, petalis caducis, filamentis staminum

clavatis, antheris antipetalis multo longioribus

quam antheris antisepal is differt.

Type species: Boronia anomala Duretto

Glabrous except for adaxial surface of petals

and staminal filaments; stellate hairs absent.

Branches with decurrent leaf bases. Leaves

imparipinnate; rhachis segments linear;

lamina dorsiventral, epicuticular wax platelets

absent; midrib impressed on the adaxial

surface, not raised on the abaxial surface,

with tightly packed tissue between midvein

and abaxial epidermis. Inflorescence 1-3-

flowered, axillary. Sepals much smaller than

petals [aestivation unknown], persistent with

fruit. Petals without a raised midrib abaxially,

valvate in bud, caducous with fruit. Stamens:

filaments clavate, suddenly narrowing at apex

so as to appear truncated before connecting

to anther, pilose along the margins below the

glandular tip; anthers attached to the apex of

the filament; antipetalous anthers much larger

than antisepalous anthers; anther-apiculum

minute. Seed (mature not seen) black, shiny,

possibly slightly rugulose, adaxial side

flattened and without ridge.

Notes: Boronia subsection Anomalae is

distinguished from the four other subsections

of B. section Valvatae by the lack of stellate

hairs and the deciduous petals. The presence

of stellate hairs and persistent petals are

apomorphies for B. section Valvatae (less

B. anomala) (Duretto & Ladiges 1999;

Duretto 1999). Other apomorphies for the

section are the axillary inflorescences and

the sepals and petals being valvate in bud.

Boronia anomala has these features apart

from the sepal aestivation character which is

unknown for the species. Sepal aestivation

and mature seed morphology are key

characters in the classification of Boronia

and both are unknown for B. anomala. The

seed possibly being slightly rugulose is

interesting as in Boronia this feature only

Duretto, New infrageneric taxa in Boronia section Valvatae

occurs elsewhere in B. sections Imbricatae

Engler and Cyanothamnus (Lindl.) F.Muell.

Together with B. subsection Grandisepalae,

which is also confined to north-western

Australia, B. anomala shares the pinched

staminal filaments and relatively large

antepetalous anthers but lacks the large

sepals characteristic of B. subsection

Grandisepalae. In the analysis described by

Duretto (1999), where the sepal aestivation

was scored as ‘unknown’, B. anomala was

sister to B. subsection Grandisepalae in 80%

of the found trees and to either B. subsection

Valvatae or B. subsection Bowmaniorum in

the remaining trees. In these cases it would

appear that the lack of stellate hairs and

deciduous petals are secondary losses for the

species.

It is worth noting that geographically

restricted and taxonomically isolated

subgeneric taxa in Boronia with large

numbers of autapomorphies are not unusual

in north-western Australia: see also B. series

Quadrilatae Duretto and B. series Rupicola

Duretto (Duretto & Ladiges 1999; Duretto

1999, accepted).

The subsection is monotypic and confined

to the Kimberley Region of northern Western

Australia (see Duretto 1999 for additional

information on Boronia anomala).

Boronia series Valvatae

A series of four subseries (see below) that

contain 21 species, plus an additional three

species, Boronia chartacea, B. hapalophylla

and B. ledifolia , that are treated as incertae

sedis (see above). Full descriptions, keys

to taxa and notes on all but two species are

given by Duretto (1999). Boronia beeronensis

Duretto and B. hapalophylla are discussed

in full by Duretto (2003) and Duretto et al.

(2004) respectively.

Boronia subseries Valvatae

Boronia alulata species-group, fide Duretto,

(1999:63)

Leaves petiolate, imparipinnate, the younger

distal leaves not unifoliolate; leaflets linear to

broadly elliptic to spathulate, obtuse to acute,

margins slightly recurved to revolute, midribs

667

impressed on the adaxial surface, raised on

the abaxial surface, the cells between midvein

and abaxial epidermis with secondary

thickening. Inflorescence 1-many flowered.

Sepals narrowly ovate-deltate to narrowly

deflate (except B. umbellata ), acute; abaxial

surface with a moderately dense indumentum,

often dark in colour.

Notes: Boronia subseries Valvatae is

characterised by the following combination

of characters: imparipinnate leaves that have

raised midribs with secondary thickening on

the abaxial surface, and the narrowly deflate

sepals. Boronia umbellata RH.Weston does

not have narrowly deflate sepals but is the

sister species of B. mollis A.Cunn. ex Lindl.

(Duretto & Ladiges 1999; Duretto 1999) and

the lack of the narrowly deflate sepals appears

to be a secondary loss.

Boronia subseries Valvatae contains eight

species (B. alulata, B. amabilis S.T.Blake,

B. angustisepala Duretto, B. hoipolloi Duretto,

B. mollis, B. obovata C.T.White, B. quinkanesis

Duretto, B. umbellata) found from Cape York

(N Qld) to the Sydney region (NSW) and in

north-western Queensand (Fig. 10 in Duretto

1999).

Boronia subseries Lanceolatae Duretto,

subseries nova a subserie typica sepalis

late deltatis abaxialiter dense tomentosis, ad

apicem acuminatis usque acutis differt.

Type species: Boronia lanceolata F.Muell.

Boronia lanceolata species-group, fide

Duretto, Muelleria (1999: 74)

Boronia foetida species-group, fide Duretto,

Muelleria (1999: 84)

Leaves petiolate, imparipinnate or simple,

leaves often becoming simple distally; leaflets

or simple leaves narrowly to broadly elliptic,

acute, the margins plane to slightly recurved

(sometimes revolute on drying); midrib

impressed on the adaxial surface, raised

prominently on the abaxial surface, the cells

between midvein and abaxial epidermis with

secondary thickening. Inflorescence 1-many-

flowered. Sepals broadly deflate, acuminate

to acute, abaxial surface with a dense, fawn

stellate indumentum.

668

Notes : Boronia subseries Lanceolatae is

characterised by the presence of ovate-

deltate sepals, and petiolate leaves that have

prominently raised midribs with secondary

thickening. All species have large simple

leaves except B. duiganiae Duretto which

usually has pinnate leaves, and B. odorata

Duretto which sometimes has juvenile leaves

that are trifoliolate.

Boronia subseries Lanceolatae contains

seven species (B. bella Duretto, B. duiganiae ,

B. excelsa Duretto, B. foetida Duretto,

B. jensziae Duretto, B. lanceolata, B. odorata )

and is found in the ‘Top End’ of the Northern

Territory and eastern Queensland (Figs 11 &

13 in Duretto 1999).

Boronia subseries Rosmarinifoliae Duretto,

subseries nova a subserie typica foliis

simplicibus sessilibus, cellulis costae abaxialis

sine densatione secundaria differt.

Type species: Boronia rosmarinifolia A.Cunn.

ex Endl.

Boronia rosmarinifolia species-group, fide

Duretto (1999: 78)

Leaves sessile, simple, linear to elliptic to

obovate, obtuse, margin plane to revolute;

midvein impressed slightly on the adaxial

surface, raised slightly on the abaxial surface,

the cells between the midvein and the abaxial

epidermis without secondary thickening.

Inflorescence l(-3)-flowered. Sepals ovate-

deltate, acute to acuminate, abaxial surface

with a dense fawn indumentum.

Notes : Boronia subseries Rosmarinifoliae is

characterised by small, sessile, simple leaves

without secondary thickening in the cells of

the abaxial midrib.

Boronia subseries Rosmarinifoliae

contains six species ( B. beeronensis, B.forsteri

Duretto, B. glabra (Maiden & Betche) Cheel,

B. palasepala Duretto, B. rosmarinifolia ,

B. splendida Duretto) and is found in south¬

eastern Queensland and north-eastern New

South Wales (Fig. 12 in Duretto 1999; Fig. 15

in Duretto 2003).

Austrobaileya 7(4): 665-668 (2008)

Acknowledgement

I’d like to thank Neville Walsh for completing

the Latin diagnoses.

References

Duretto, M.F. (1999). Systematics of Boronia section

Valvatae sensu lato (Rutaceae). Muelleria 12:

1-131.

- (2003). Notes on Boronia (Rutaceae) in eastern and

northern Australia. Muelleria 17: 19-135.

- (accepted). A reassessment of Boronia (Rutaceae)

in the Northern Territory with a key to species,

the decription of one new species and the

reduction of another. The Beagle

Duretto, M.F., Edwards, J. & Edwards, P. (2004).

Boronia hapalophylla (Rutaceae), a new and

restricted species from north-eastern New

South Wales. Telopea 10: 705-710.

Duretto, M.F. & Ladiges, P.Y. (1999). A cladistic

analysis of Boronia section Valvatae (Rutaceae).

Australian Systematic Botany 11: 636-665.

Humphries, C.J., Williams, PH. &Vane-Wright, R.I.

(1995). Measuring biodiversity value for

conservation. Annual Review of Ecology and

Systematics 26: 93-111.

Solanum succosum A.R.Bean & Albr. (Solanaceae),

a new species allied to S. chippendalei Symon

A.R.Bean 1 & D.E.Albrecht 2

Summary

Bean, A.R. & Albrecht, D.E. (2008). Solanum succosum A.R.Bean & Albr. (Solanaceae), a new species

allied to S. chippendalei Symon. Austrobaileya 7(4): 669-675. The new species, Solanum succosum

from Queensland and the Northern Territory, is described and illustrated. It is closely allied to

S. chippendalei for which a comparative description is provided. Distribution maps are presented for

both species and S. succosum is illustrated.

Key Words: Solanum succosum, Solanum chippendalei , Solanaceae, taxonomy, Australian flora,

identification

'A.R.Bean, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens, Mt

Coot-tha road, Toowong 4066, Queensland, Australia. Email: tony.bean@epa.qld.gov.au

2 D.E. Albrecht, Alice Springs Herbarium, Department of Natural Resources, Environment and the Arts,

PO Box 1120, Alice Springs, Northern Territory, 0871, Australia. Email: dave.albrecht@nt.gov.au

Introduction

In his revision of Australian Solanum ,

Symon (1981) identified a group of closely

related species ( S. melanospermum F.Muell,

S. chippendalei Symon, S. phlomoides

A.Cunn. ex Benth. and S. beaugleholei

Symon), which he termed a “quartet of species

with common ancestry”. He did not outline

the distinguishing features of the group, but

these species may be characterised by the

strongly andromonoecious inflorescences

(one bisexual flower at the base and numerous

male flowers above), the calyx not accrescent

(although the calyx lobes may be almost as

long as the mature fruit), the black seeds, and

the fruits 21-40 mm diameter (among the

largest for Solanum spp. in Australia).

Solanum chippendalei sensu Symon,

extends from the eastern Pilbara region in

Western Australia to Winton in northwest

Queensland, in semi-arid hummock grassland

and shrubland to woodland habitats. Field and

herbarium observations by the authors indicate

Symon’s concept of S. chippendalei comprises

two species: S. chippendalei and S. succosum

A.R.Bean & Albr., here described as new.

Accepted for publication 6 June 2008

Materials and methods

The present study is based on herbarium

specimens at BRI (including material in spirit),

CANB, DNA, NT, MEL and PERTH as well

as field studies by the authors in Queensland

(Qld), southern Northern Territory (NT)

and the Pilbara region of Western Australia

(WA).

The species descriptions and terminology

presented in this paper follow those of Bean

(2004) and all data gathered during this

study have been added to an existing DELTA

(Dallwitz et al. 1993) dataset. An interactive

key covering Solanum spp. of eastern

Australia is available at the DELTA website

(http://delta-intkey.com/).

Prickle width is measured just above the

base of the prickle, where the prickle surface

is at 45° to the branch. The prickle length is

the distance from this position to the apex of

the prickle. Adult leaves are those adjacent to,

or distal from, the inflorescences.

Taxonomy

Solanum succosum A.R.Bean & Albr.

species nova affine S. chippendalei sed a quo

foliis inferioribus profunde lobatis, filamentis

multo longioribus dorsifixis, seminibus

minoribus, et fructibus persistentibus cavitate

interna liquido impleta et pericarpio tenuiore

praeditis, differens. Typus: Queensland.

Burke District: c. 8 km along road to Lady

670

Loretta Mine, turnoff c. 65 km from Mt Isa,

24 August 1993, T.R.Lally 101 (holo: BRI; iso:

CANB, DNA).

Solarium sp. Juicy Fruit (P.K.Latz 18938) in

Albrecht et al. (2007).

Illustration : Symon (1981: 273), as Solanum

chippendalei.

Erect, rhizomatous perennial shrub 0.4-1.5 m

high. Adult branchlets terete or ridged, white,

grey, yellow, rusty or brown; prickles 3.5-40

per cm, straight, acicular, 1-10 (-13) mm

long, 7-16 times longer than wide, with dense

stellate hairs on lower part; stellate hairs very

dense, 0.7-1.3 (-1.6) mm diameter, stalks

0-1.6 (-2) mm long, lateral rays 7-8, porrect

or ascending; central ray 1-1.5 times as long

as laterals, not gland tipped. Adult leaves

lanceolate, elliptical or ovate, shallowly or

deeply lobed throughout, lobes 2-4 on each

side, obtuse, lobing index 1.2-5, at least the

lower leaves always deeply lobed; lamina

3-12 cm long, 1.4-5.3 (-8) cm wide, 1.6-3.5

times longer than broad; apex obtuse or acute;

base cuneate or obtuse; oblique part 0-11 mm

long, obliqueness index 0-10 percent; petioles

0.8-2.1 cm long, 20-30% length of lamina,

prickles present. Upper leaf surface grey-

green or grey, prickles 0-8, straight, acicular

or broad-based, 3-10 mm long; stellate hairs

distributed throughout, stellate hairs dense

to very dense, 0.15-0.3 mm apart, 0.5-1 mm

across, stalks 0-0.5 mm long; lateral rays 6-9,

porrect or ascending; central ray 1-2.5 times

as long as laterals, not gland-tipped; simple

hairs absent; Type 2 hairs absent. Lower leaf

surface grey, white, or yellowish, prickles 0-5,

straight, acicular or broad-based; stellate hairs

dense to very dense, 0.1-0.2 mm apart, 0.7-1.5

mm diameter, stalks 0-0.8 mm long; lateral

rays 6-8, porrect; central ray 1-1.8 times as

long as laterals, not gland tipped; simple hairs

absent; Type 2 hairs absent. Inflorescence

supra-axillary, cymose (pseudo-racemose),

common peduncle 0-3 mm long, rhachis

prickles present, 7-11-flowered, with one

usually markedly larger bisexual flower, the

rest male. Pedicels at anthesis 5-23 mm long,

prickles absent or present. Calyx prickles on

basal bisexual flower 35-125, each 2-6 mm

long; calyx prickles on distal male flowers 0-

Austrobaileya 7(4): 669-675 (2008)

15, each 1-5 mm long; calyx tube at anthesis

3-5 mm long, lobes attenuate, 6.5-11 mm

long. Calyx stellate hairs very dense, white,

0.7-0.9 mm across, stalks 0-1.5 mm long,

lateral rays 7-8; central ray 1-1.5 times as

long as laterals, not gland tipped; simple hairs

absent; Type 2 hairs absent. Corolla purple,

10-33 mm long, rotate, inner surface glabrous

or rarely with a few stellate hairs at tip of lobes.

Anthers 5.8-8 mm long, dorsifixed towards

base. Filaments (1.6-) 2-3.3 mm long. Ovary

glabrous. Functional style glabrous, 10-12.5

mm long, erect near base, then bent, displacing

one anther. Fruiting calyx lobes less than or

more than half length of mature fruit, lobes

not recurved or strongly recurved; prickles

2-6 mm long. Fruits 1 per inflorescence,

broadly ellipsoidal or less commonly globular,

21-30 mm diameter at maturity, yellow or

yellowish-green, eventually turning very dark

brown and prune-like, 2-locular, placenta

stalked, anvil-shaped, extending the length of

fruit; pericarp initially 1.5-4 mm thick, the

innermost surface gelatinous and gradually

deliquescing, finally 1-1.5 mm thick; internal

cavity of fruit fluid-filled when fruits green

or yellow. Pedicels at fruiting stage 20-55

mm long, 1.3-2.5 mm thick at midpoint.

Seeds black, flattened, 3-4 mm long, 27-3.2

mm wide. Fig. 1.

Additional selected specimens examined : Queensland.

Gregory North District: 47.8 km from Dajarra, towards

Mt Isa, May 2005, Bean 23697 (BRI, UT); Trough

Tank, Placer Pacific Osborne exploration lease, 31km

N of Pathungra, May 1993, Gunness AG2187 (BRI); 70

miles [113 km] W of Winton on Boulia road. May 1966,

Pedley 2000 (BRI); 15 km NE of ‘Burnham’, Jun 1979,

Purdie 1551 (BRI). Burke District: Mt Isa, Nov 1930,

MacCallum s.n. (BRI); 3.9 km by road along Gunpowder

road from junction with highway, Jun 2004, McDonald

KRM2861 (BRI); Camooweal Caves N.P., Apr 2001,

McDonald KRM849 (BRI); Corella River, 88 km W

of Cloncurry on road to Mt Isa, Jul 2003, McKenzie

RAM03/77 (BRI). Northern Territory. Burt Plain: 15

miles [24 km] N of Alice Springs, May 1968, Nelson 1698

(MEL, NT). Darwin Coastal: 2 miles [3 km] E Howard

Creek on Koolpinya road, Jan 1964, Robinson 236 (DNA).

Davenport Murchison Range: 3.5 km N of Whistleduck

Creek camping area, Davenport Range N.P., May 2005,

Bean 23802 (BRI, UT); near Dixon Creek, N of Devils

Marbles, May 2005, Bean 23832 (BRI, UT); 13.2km S of

Renner Springs, Jun 2005, Bean 24167 (BRI, NY, UT).

Mitchell Grass Downs: 18 miles [29 km] S of Elliot, Feb

1969, Latz 426 (NT). Ord-Victoria Plains: Mt Sanford,

Aug 2000, Brock 124 (B, NT); Humbert River Station,

Jun 1974, Latz 5324 (AD, MO, NT). Sturt Plateau: 3.5

Bean & Albrecht, Solarium succosum

671

Fig. 1. Solanum succosum. A. flowering branchlet * 1. B. adult leaf x 1. C. calyx and fruit, close to maturity x 1. D.

transverse section of fruit x 1.5. E. seed x 8. A from McDonald KRM862 (BRI); B from Bean 23802 (BRI); C-E from

McDonaldKRM856 { BRI). Del. W. Smith.

km S of Three Ways, towards Tennant Creek, Apr 2001,

McDonald KRM856 (BRI); N of Renner Springs, Apr

2001, McDonald KRM862 (BRI); 16 km S of Larrimah,

Jun 1972, Symon 7639 (AD, B, NSW, NT); 1 km N of

Larrimah, Jun 1975, Symon 10353 (AD, CANB, MO,

NT). Tanami: 7 miles S of Tennant Creek town. Mar

1955, Chippendale 956 (CANB, DNA, MEL, NT, S); 27

km S of Tennant Creek, July 1977, Latz 7419 (AD, NT);

Newcastle Waters, 1887, Giles s.n. (MEL).

Distribution and habitat: Solanum succosum

is found in summer rainfall dominated semi-

arid and monsoonal parts of northwest Qld

(from Lawn Hill National Park to Winton)

and north central to northwest NT (from about

Humbert River Station in the northwest to as

far south as about Wauchope). Two specimens

(just north of Alice Springs on the verge of the

Stuart Highway, and east of Howard Creek in

the Darwin Coast bioregion) are distant from

the main range of the species (Map 2). The

former occurrence is undoubtedly due to

human influence but the status of the latter

is equivocal. Most records of the species are

from Triodia dominated habitats on gravelly

hillslopes, and gravelly or sandy plains and

creeklines. A few records have been made

from along tracks through Mitchell grass

672

country on clay plains. Germination of seeds

and clonal regrowth are both evident after

fire.

Phenology : This species may flower and fruit

at any time of year, generally in response to a

fire or rainfall event.

Notes: Solanum succosum is closely allied

to S. chippendalei but differs in a number of

respects. In S. succosum , the lower leaves (and

often all leaves) are deeply lobed (not entire

or shallowly lobed), the anthers are dorsifixed

(not basifixed), the filaments are (1.6-) 2.0-

3.3 mm long (compared to 5-7 mm long). The

mature fruits are mostly longer than broad

(although sometimes globular), have liquid-

filled internal cavity, the pericarp is initially

1.5-3 mm thick, becoming thinner with age,

and the placenta is longitudinally continuous.

The fruits are very firmly attached to the

calyx, and detached only with difficulty, then

leaving a conical plug on the calyx. Fruits

detached in this manner invariably exhibit

two pores near the end of the fruit through

which juice and seeds may be squirted. The

seeds are 3-4 mm long and 27-3.2 mm wide,

the lower leaf surface has stellate hairs 07-1.6

mm diameter, and stellate hairs on branchlets

are 0.8-1.3 mm diameter.

By contrast, in Solanum chippendalei the

leaves have shallow lobes throughout, or are

(frequently) entire. The filaments are 0.5-1.5

mm long and basally attached; the mature

fruits are globular, lack liquid in the internal

cavity, the pericarp is 4-75 mm thick, not

becoming thinner with age, and the placenta

does not extend to the distal end of the fruit.

The fruits are not firmly attached to the calyx,

and mature fruits may often be seen lying on

the ground after falling from the plant. The

scar on the fruiting calyx is flat and circular.

Detached fruits never exhibit pores. Seeds are

(3.2-) 3.7-5 (-5.3) mm long and (2.5-) 2.8-4

(-4.6) mm wide, the lower leaf surface has

stellate hairs 0.5-1 mm diameter, and stellate

hairs on branchlets are 0.4-1 mm diameter.

Ethnobotany: Kaytetye and Anmatyerr

peoples recognise Solanum succosum as

distinct from S.chippendalei. Both language

groups use the name tyatyarlkwerre for

S. succosum and regard the fruit as inedible,

Austrobaileya 7 ( 4 ): 669-675 ( 2008 )

one woman saying they are poisonous (Myf

Turpin, pers. comm. 2007). Anmatyerr

people use the names anakety or antyewal

for S. chippendalei and regard the fruit highly

(Latz 1995).

Etymology: From the Latin succosus meaning

‘juicy’. This refers to the liquid-filled internal

cavity of mature fruits of this species, which

contrasts strongly with the dry internal cavity

of the related Solanum chippendalei.

Solanum chippendalei Symon, J. Adelaide

Bot. Gard. 4: 272 (1981). Type: Western

Australia, base of the Sir Frederick Range, 1

August 1962, D.E.Symon 2272 (holo: AD; iso:

AD, CANB, PERTH).

Illustrations: Symon (1981: 267, fig. 119);

Latz (1995: 271-272).

Erect, rhizomatous perennial shrub, 0.25-

1.5 m high. Adult branchlets white, grey,

yellow, rusty or brown; prickles 4-26 per cm,

straight, acicular, 2.5-8 mm long, 8-17 times

longer than wide, with scattered stellate hairs

on lower part. Adult branchlets with stellate

hairs very dense, 0.4-1 mm diameter, stalks

up to 0.8 mm long, lateral rays 7-9, porrect

or ascending; central ray 07-1.6 times as

long as laterals, not gland tipped. Adult

leaves lanceolate, elliptical or ovate, entire

or shallowly lobed throughout, lobes when

present 3-5 on each side, obtuse, lobing index

1-2; lamina 3-13.2 cm long, 1.6-5.1 cm wide,

1.5-3.3 times longer than broad; apex obtuse

or acute; base cuneate or obtuse; oblique

part 2-12 mm long, obliqueness index 2-11

percent; petioles 1.2-4.3 cm long, 16-38%

length of lamina, prickles present. Upper

leaf surface grey-green or grey, prickles 0-4,

straight, acicular, 2-4 (-8) mm long; stellate

hairs distributed throughout, stellate hairs

dense to very dense, 0.05-0.25 (-0.3) mm

apart, 0.5-0.9 (-1.2) mm across, stalks 0.1-

O. 5 (-0.7) mm long; lateral rays 7-8, porrect;

central ray 0.8-1.5 times as long as laterals, not

gland tipped; simple hairs absent; Type 2 hairs

absent. Lower leaf surface white, to yellowish,

prickles 0-5 (-10), straight, acicular; stellate

hairs very dense, 0.05-0.2 (-0.25) mm apart,

0.5-1 mm diameter, stalks 0.1-07 (-1) mm

long; lateral rays 7-8, porrect; central ray 0.7-

1.5 times as long as laterals, not gland tipped;

Bean & Albrecht, Solanum succosum

673

simple hairs absent; Type 2 hairs absent.

Inflorescence leaf-opposed or supra-axillary,

cymose, common peduncle up to 33 mm

long, rhachis prickles absent or present, 8-30-

flowered, with one bisexual flower, the rest

male. Flowers 5-merous, markedly dimorphic,

with a larger pricklier basal flower. Pedicels

at anthesis 5-27 mm long, prickles absent

or present. Calyx prickles on basal bisexual

flower 70-130, each 2-9 mm long; calyx

prickles on distal male flowers 0-12, each

2-4.5 mm long; calyx tube at anthesis 2-6

mm long, lobes deflate to attenuate, 8-21 mm

long. Calyx stellae very dense, white, 0.5-0.8

mm across, stalks 0-0.8 mm long, lateral

rays 7-9; central ray 1-1.5 times as long as

laterals, not gland tipped; simple hairs absent;

Type 2 hairs absent. Corolla purple, 7-28 mm

long, rotate or shallowly lobed, inner surface

glabrous or sparsely stellate-hairy. Anthers

5-7 mm long, basifixed. Filaments 0.5-1.5

mm long. Ovary glabrous or with stellate

hairs. Functional style glabrous, 12-13 mm

long, erect near base, then bent, displacing

one anther. Fruiting calyx lobes more than

half length of mature fruit, lobes not recurved

or rarely strongly recurved; prickles 1-7 mm

long. Fruits 1 per inflorescence, globular

to obloid, 23-40 mm diameter at maturity,

yellow or yellowish-green, 2-locular, placenta

not apparent at distal end, pericarp 4-7.5 mm

thick, not deliquescing; internal cavity of fruit

not fluid-filled. Pedicels at fruiting stage 15-

50 mm long, 1.5-2.7 mm thick at midpoint.

Seeds black, flattened, (3.2-) 3.7-5 (-5.3) mm

long, (2.5-) 2.8-4 (-4.6) mm wide.

Additional selected specimens examined: Western

Australia, along No. 1 Rabbit Proof fence, 7 miles

[11 km] N of Sandy Creek, May 1947, Royce 1638

(PERTH); Dovers Hills, northern Gibson Desert, Jul

1967, George 8977 (PERTH); 50 miles [80 km] W of

Giles, Jun 1968, Howard s.n. (PERTH); Canning Stock

Route, Jul 1976, Palmer 27 (PERTH); Bloodwood Bore,

Balweena Reserve, Aug 1969, Nelson 1925 (PERTH);

S side of Petermann Ranges, W of N.T. - W.A. border,

Sep 1978, Beauglehole 60602 & Errey (PERTH); 14 km

S of Kumarina, N of Meekatharra, May 1975, Symon

10000 (PERTH); 18 km S of Two Sisters, c. 160 km SE

of Shay Gap, Jul 1984, Newbey 10403 (PERTH); Balgo

Hills, via Derby, 15 June 1967, McNamara lc (PERTH).

Northern Territory. Burt Plain: 21 miles [34 km] SW

of Napperby Station, Sep 1956, Lazarides 6001 (CANB,

NT, PERTH); Stirling Bore, 5 miles [8 km] S of Barrow

Creek township, Sep 1955, Perry 5349 (K, L, MEL, NSW,

NT, US). Davenport Murchison Range: Elkedra Station,

May 1977, Latz 6962 (ADW, BRI, NSW, NT); 98 km E

of Three Ways on Barkly Highway, Jun 2004, McKenzie

RAM04/63 (BRI). Great Sandy Desert: Wartupunya

Rockhole, Jan 1972, Latz 2137 (ADW, NT); Pulca

Currinya, Mt Wedge Station, Sep 1976, Latz 6601 (ADW,

CANB, NT). Macdonnell Ranges: Mt Liebig area, Apr

1972, Latz 2272 (ADW, NT); Haasts Bluff airstrip, Nov

1976, Latz 6667 (ADW, CANB, NT). Ord Victoria

Plain: road to Lajamanu from Buntine Highway, May

2004, Brennan 6299 (AD, DNA, NT, PERTH). Tanami:

Stuart Highway near Ali Curung turnoff, Jun 2006,

Albrecht 12049 (BRI, NT); The Granites Tenements,

Dec 1984, Kalotas 1797 (ADW, NT, PERTH).

Distribution and habitat : Solanum

chippendalei is found in arid and semi-arid

parts of north central to southwest NT (from

Wave Hill Station in the northwest to Georgina

Downs in the northeast, and to Docker River

in the far southwest), extending westward

into WA as far west as the Pilbara (Map 1).

A specimen from Alice Springs {Latz 22296 )

is certainly a recent introduction from seed

discarded by aborigines. It occurs in Triodia

dominated communities on sandy or gravelly

plains and hillslopes. Germination and clonal

regrowth is strongly encouraged by fire or

soil disturbance and it is commonly found on

roadsides.

Phenology: Flowers and fruits can be found

at any time of the year depending on timing

of rains and/or fire.

Notes: Solanum chippendalei is closely

allied to S. succosum (see notes under that

species). It is also very close to S. phlomoides

A.Cunn. ex Benth. Both S. chippendalei and

S. phlomoides , as currently understood, are

quite variable, and distinguishing between

them is difficult.

Ethnobotany: The fruit of Solanum

chippendalei is an outstandingly important

plant food for central Australian aborigines

throughout its distribution range, and it

has various language names. The seeds

are removed and the thick pericarp eaten,

which has a rather bland taste reminiscent of

cantaloupe (rockmelon). Although relatively

high in vitamin C, it has quite low protein

and fat values (Latz 1995). The ripe fruit has

excellent keeping qualities. Those not eaten

fresh are sometimes threaded onto skewers

or made into pasteballs, dried and stored

for considerable periods. In recent times

674

Alyawarr people (NT) have deliberately

established colonies of S. chippendalei south

of its natural range by scattering seeds near

their camp sites (Latz 1995). This practice

has also been reported by Walsh (in prep.) for

the Martu people of the western Great Sandy

Desert area in the vicinity of Rudall River

and the central Canning Stock Route. Martu

elders assert that they have always scattered

the seeds of S. chippendalei and believe that

it is their role to disperse productive plants

(Walsh, in prep.).

Several language groups, e.g. Warlpiri,

Pitjantjatjara, Martu and probably Pintubi

recognise two forms of Solarium chippendalei

(Latz 1995; Walsh, in prep.). The form found

on hillsides is called ngayaki (Warlpiri), pura

(Pitjantjatjara) and piljiwin (Martu), and the

sandplain form is called wanakiji (Warlpiri)

and pura (Martu) (Latz 1995; Walsh, in

prep.). In our investigations we have not found

any reliable morphological characters that

differentiate the habitat forms recognised by

some aboriginal groups.

Etymology : Named for George Chippendale

(1921-extant), a botanist who worked for

many years in the arid parts of Northern

Territory, based at Alice Springs.

Acknowledgements

We thank the Directors of CANB and PERTH

for the loan of specimens, the Directors of

MEL and DNA for access to those Herbaria,

Peter Latz for alerting us to the distinctiveness

of Solanum succosum , Tim Collins for

growing plants for study, Myf Turpin, Glenn

Wightman and Fiona Walsh for ethnobotanical

information, Will Smith (BRI) for the

illustrations, and Peter Bostock for the Latin

diagnosis and for producing the distribution

maps.

References

Albrecht, D.E., Duguid, A.W., Latz, P.K., Harris, M.G.

& Coulson, H. (2007). Vascular Plant Checklist

for the Southern Bioregions of the Northern

Territory. 2nd edition. Department of Natural

Resources, Environment and the Arts: Alice

Springs.

Austrobaileya 7(4): 669-675 (2008)

Bean, A.R. (2004). The taxonomy and ecology of

Solanum subg. Leptostemonum (Dunal) Bitter

(Solanaceae) in Queensland and north-eastern

New South Wales. Austrobaileya 6: 639-816.

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

onwards). User’s guide to the DELTA system:

a general system for processing taxonomic

descriptions, 4 th edition. http://delta-intkey.

com/

Latz, P.K. (1995). Bushfires andBushtucker. Aboriginal

plant use in Central Australia. IAD Press: Alice

Springs.

Symon, D.E. (1981). A revision of the genus Solanum

in Australia. Journal of the Adelaide Botanic

Gardens A. 1-367.

Walsh, F. (in prep.). Land use and management systems

of Martu Aborigines: implications for land

management in the Sandy Deserts, Australia.

Doctoral thesis. University of Western

Australia, Departments of Anthropology and

Plant Biology.

Bean & Albrecht, Solarium succosum

675

.Map 2. Distribution of Solarium succosum.

F.M.Bailey’s taxonomy of A.Meston’s collections

from the Bellenden Ker Expedition of 1904

John Leslie Dowe 1 & Alan D. Broughton 2

Summary

Dowe, J.L. & Broughton, A.D. (2008). F.M.Bailey’s taxonomy of A.Meston’s collections from the

Bellenden Ker Expedition of 1904. Austrobaileya 7(4): 677-679. The four new taxa described by

F.M.Bailey for collections made by A.Meston during his 1904 ascent of Bellenden Ker Range, were

first published in a Parliamentary Paper tabled 1st June 1904. This preceded his publication in the

Queensland Agricultural Journal, on 1st July 1904. A note is also provided on place names associated

with Bellenden Ker Range and their recommended spelling.

Key Words: F.M.Bailey, A.Meston, Bellenden Ker Range, Queensland botany

‘John Leslie Dowe, Australian Centre for Tropical Freshwater Research, James Cook University,

Townsville, Queensland 4811, Australia. E-mail: John.Dowe@jcu.edu.au

2 Alan D. Broughton, Cairns Historical Society, Cnr Lake & Shields St., Cairns, Queensland 4870,

Australia.

Introduction

F.M.Bailey’s contribution to the botany of

Bellenden Ker Range is mostly contained

in two reports, and based on collections

made during expeditions in 1889 and 1904.

Both expeditions were organized and led

by Archibald Meston (Meston 1889, 1904).

Bailey’s only ascent of the range was during

the 1889 expedition, during which some

600 specimens were collected. An account

of the 1889 expedition itinerary and route,

and a re-assessment of the taxonomic and

nomenclatural results were provided by Dowe

& Broughton (2007). The expedition of 1904

was conducted under the auspices of the

Queensland Government Lands Department,

and was not so much a scientific expedition

but one which “confined itself almost entirely

to the economic plants ... or promise to be

of practical value to mankind” and to have

“obtained as many specimens as possible for

Mr Bailey’s herbarium” (Meston 1904, p. 3

[p. 1009]; Table 1). In the introductory section

of the report, Meston wrote short accounts on

Timbers, Gums, Wood for Veneer, and on the

Aborigines and their names for various plant

species.

Table. 1. Meston’s itinerary for the 1904 ascent of Bellenden Ker Range

Activity

Date

Departed Brisbane for Cairns

12 January

Arrived Cairns

17 January

Departed Cairns for Bellenden Ker Range

19 January

Summit camp

24-29 January

Returned to Cairns

8 March

Departed Cairns for Brisbane

9 March

Arrived Brisbane

14 March

Accepted for publication 1 July 2008

678

In the botanical section of the report,

Bailey listed 200 taxa that were collected

by Meston or his indigenous collectors (see

figure in Clarke 2008: 145). The specimens

were seen and examined by Bailey upon

their arrival in Brisbane. A search of herbaria

[BRI, BM, K, MEL, NSW] revealed 109

specimens are extant and otherwise attributed

to Meston from the 1904 expedition. It must

be assumed that some of Meston’s collections

may not have been conserved as herbarium

specimens, or otherwise have not survived,

hence the disparity between the number of

species listed by Bailey in the report and the

number of specimens that have been located.

F.M.Bailey’s botanical report of the 1904

expedition

Four new species were described by Bailey

from Meston’s collections from the Bellenden

Austrobaileya 7(4): 677-679 (2008)

Ker Range expedition of 1904 (Table 2).

Citations from the taxonomic literature give

the place of publication of the new species

as the Queensland Agricultural Journal of

July 1904 (Bailey 1904b). However, there is a

prior publication which should be cited as the

place of valid publication of the four species,

a Parliamentary Paper that was tabled 1 June

1904 (Bailey 1904a) (pers. comm., D. Embury,

Queensland Parliamentary Service). Tabling

of a Parliamentary Paper qualifies as ‘effective

and valid publication’ under the International

Code of Botanical Nomenclature (McNeill

et al., 2006). The protologues were identical

apart from some very minor grammatical and

punctuation changes in each publication, and

each new taxon was appended as being a ‘ sp.

novl in both publications.

Table 2. New taxa described by F.M.Bailey from Meston’s 1904 Bellenden Ker Range

expedition collections

Taxon/Species

Bailey,

1904a

Bailey,

1904b

Type citation

Location of

types

Pentapanax bellendenkerensis

F.M.Bailey [= Polyscias

bellendenkerensis (F.M.Bailey)

Philipson] [Araliaceae]

p. 12

p. 491

Summit of

Bellenden-

Ker, Meston’s

expedition, 1904,

A.Meston 68

holo: BRI

Psychotria coelospermum

F.M.Bailey [Rubiaceae]

p. 13

p. 492

Barron R, 1877,

Bailey; Meston’s

Expedition to

Bellenden-Ker,

1904

Bailey syntype

specimen at

BRI; Meston

syntype

specimen not

located* **

Lucuma obpyriformis

F.M.Bailey [= Pouteria

obpyriformis (F.M.Bailey)

Baehni = Sersalisia sericea

(Aiton) R.Br.] [Sapotaceae]

p. 13

p. 493

Meston’s

Bellenden-Ker

Expedition, 1904

[Mt Toressa,

1904, Meston 29]

holo: BRI

Bulbophyllum lageniforme

F.M.Bailey [Orchidaceae]

p. 16

p. 494

Bellenden-Ker

Expedition, 1904,

A.Meston s.n.

**lecto: BRI;

isolecto: K

*PI.Forster, pers. comm. April 2008.

**selected by Clements (1989).

Do we & Broughton, Bellenden Ker Expedition of 1904

Places names associated with Bellenden

Ker Range

According to the Queensland Place Names

Gazetteer, the official spelling of the name

Bellenden Ker does not use a hyphen. In Dowe

& Broughton (2007) we followed Meston’s

(1889) account of his report to Parliament,

‘Report by A. Meston on the Government

Scientific Expedition to the Bellenden-Ker

Range (Wooroonooran) North Queensland'’

where he used the hyphenated spelling and

continued to use this form in all his subsequent

publications. The use of a hyphen appears to

have ceased in the 1920s, and it has more or

less been consistently unhyphenated since

then, although Clarke (2008) has recently

maintained the older usage.

The highest peak on the northern section

of the Bellenden Ker Range is listed in the

Gazetteer as ‘Centre Peak’, rather than ‘Mt

Bellenden Ker’ or ‘Mt Bellenden Ker Centre

Peak’ ( cf. Dowe & Broughton 2007).

Acknowledgements

We would like to thank the Directors of BRI,

MEL and NSW for supplying information

about their collections; Jenny Tonkin for

information about Meston’s collections in

K and BM; and David Embury, Queensland

Parliamentary Service, for answering queries

concerning the date of tabling of Bailey’s

Report.

References

Bailey, F.M. (1904a). Report by the Government Botanist

on botanical specimens collected by Mr. A.

Meston on or about the Bellenden-Ker Range.

Queensland: Parliamentary Papers printed

during the Third Session of the Fourteenth

Parliament, pp. 9-17 [pp. 1015-1023 in

repaginated and overprinted bound copy], G.A.

Vaughan, Government Printer: Brisbane.

- (1904b). Contributions to the flora of Queensland.

Queensland Agricultural Journal 15: 491-495.

Clarke, P.A. (2008). Aboriginal Plant Collectors:

botanists and Australian aboriginal people in

the nineteenth century. Rosenburg Publishing

Pty Ltd.: Dural.

Clements, M.A., (1989). Catalogue of Australian

Orchidaceae. Australian Orchid Research 1:

16.

679

Dowe, J.L. & Broughton, A.D. (2007). F.M.Bailey’s

ascent of Mt Bellenden-Ker in 1889, and notes

on the publication priority of new vascular plant

species from the Expedition. Austrobaileya 7:

555-566.

Meston, A. (1889). Report by A. Meston on the

government scientific expedition to the

Bellenden-Ker Range (Wooroonooran),

north Queensland. Queensland: Votes and

Proceedings of the Legislative Assembly

4: 1-13 [pp. 1205-1217 in repaginated and

overprinted bound copy], James C. Beal,

Government Printer: Brisbane.

- (1904). Report by Mr. A. Meston on Expedition

to the Bellenden-Ker Range. Queensland:

Parliamentary Papers printed during the Third

Session of the Fourteenth Parliament, pp. 1-17

[pp. 1007-1023 in repaginated and overprinted

bound copy], G.A.Vaughan, Government

Printer: Brisbane.

McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin,

V., Hawksworth, D.L., Marhold, K.,

Nicolson, D.H., Prado, J., Silva, PC., Skog,

J.E., Wiersema, J.H. & Turland, N.J. (2006)

International Code of Botanical Nomenclature

(VIENNA CODE), adopted by the Seventeenth

International Botanical Congress, Vienna,

Austria, July 2005. Regnum Vegetabile 146.

A.R.G. Gantner: Verlag KG.

New subspecies for Zieria odorifera J.A.Armstr.

(Rutaceae) from northern New South Wales

Marco F. Duretto 1 & Paul I. Forster 2

Summary

Duretto, M.F. & Forster, P.I. (2008). New subspecies for Zieria odorifera J.A.Armstr. (Rutaceae) from

northern New South Wales. Austrobaileya 7(4): 681-690. Three new subspecies of Zieria odorifera

endemic to northern New South Wales are described and illustrated, these being: subsp. copelandii

Duretto & P.I.Forst,, subsp. warrabahensis Duretto & P.I.Forst. and subsp. williamsii Duretto &

P.I.Forst. A key to the four subspecies of Z. odorifera is provided.

Key Words: Rutaceae, Zieria, Zieria odorifera, Zieria odorifera subsp. copelandii, Zieria odorifera

subsp. odorifera, Zieria odorifera subsp. warrabahensis, Zieria odorifera subsp. williamsii

'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. I. Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens

Mt Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: paul.forster@epa.

qld.gov.au

Introduction

Zieria Sm. (Rutaceae) contains c. 60 species

restricted to eastern Australia (Queensland,

New South Wales, Victoria, South Australia

and Tasmania) and one species, Z. chevalieri

Virot, which is endemic to New Caledonia (see

Armstrong 2002; Duretto & Forster 2007).

Two species were identified in the

Flora of New South Wales under the phrase

names Ziera sp. ‘Cathedral Rock’ (Williams

95303) and Z. sp. ‘Oxley Wild Rivers N.P.’

(Copeland 2174) (Armstrong & Harden 2002).

In anticipation of the forthcoming account

of the genus for the Flora of Australia these

informal taxonomic concepts were evaluated.

Both concepts were found to be conspecific

with Z. odorifera J.A.Armstr. and represent

variants of a new subspecies which is

described below, along with two other new

subspecies. These four subspecies form a

geographic replacement series throughout

the overall species range and can be viewed

as representing distinct regional elements in

the process of speciation via isolation and

local genetic drift. The subspecies are distinct

in character combinations predominantly

composed of indumentum type and cover and

the size of some foliar and floral characters.

Materials and methods

This paper is based on herbarium collections

housed in BRI, CANB, MEL, NE and NSW.

Our descriptions follow the format used by

Duretto & Forster (2007).

Taxonomy

The key to the species of Zieria found in New

South Wales (Armstrong & Harden 2002)

requires amendment to accommodate all

taxa found in this state. For example, with

Z. odorifera the key (p. 278: Group 3, couplet

4), and later the description of the species,

indicate that the fruit is pubescent or hirsute,

though many of the specimens Armstrong

(2002) cites in the protologue have glabrous

fruit. In addition, some specimens have

stellate hairs dominating the indumentum

of the stems and so belong to Group 4 of

Armstrong & Harden (2002) and thus will not

key out to Z. odorifera. To key out Z. odorifera

in the Flora of New South Wales the following

couplets need to be inserted.

Accepted for publication 9 July 2008

682

Austrobaileya 7(4): 681-690 (2008)

Zieria Group 3 (Armstrong & Harden (2002, p. 278)

3* Petals white or rarely white with pink tips; inflorescence mostly 1-3-

flowered, shorter than or longer than leaves.

4 Abaxial surface of sepals glabrous.11. Z. adenophora

4* Abaxial surface of sepals variously hairy.

5 Leaflets cuneate-obovate to obcordate, margins flat to slightly recurved;

petals 2-2.5 mm long.12. Z. obcordata

5* Leaflets leaflets narrow elliptic to narrow oblanceolate to oblanceolate,

margins slightly recurved to revolute; petals 2-4.5 mm long.8. Z. odorifera

Zieria Group 4 (Armstrong & Harden (2002, p. 279)

2* Lower surface of leaflets glabrous or hirsute, but never velvety

10a Leaflets usually <3.5 mm wide; petioles < 6 mm long.

10b Petioles 1-1.5 mm long; leaflets 2.5-5 mm long.8. Z. odorifera

10b* Petioles 4-6 mm long; leaflets 17-25 mm long.23. Z. floydii

10a* Leaflets mostly 4-7 mm wide; petioles 8-20 mm long.24. Z. smithii

Zieria odorifera J.A.Armstr., Austral. Syst.

Bot. 15: 412 (2002). Type: New South Wales.

North Western Slopes: Warrumbungle

Range, Burrumbuckle Rock, 8 November

1977, M.D.Crisp 3609 (holo: CANB

[CBG7707801]; iso: NSW 455488).

Erect to prostrate, sometimes spindly, shrub,

to 1 m tall. Branches not or weakly to strongly

glandular verrucose, sparsely to densely

pilose with mainly simple and bifid hairs, or

stellate hairy, hair distribution even, rarely

concentrated between decurrent leaf bases

which are poorly developed or absent. Leaves

palmately trifoliolate, strongly odoriferous,

glabrescent to pilose; petioles 1-2 mm

long, not or weakly to strongly glandular

verrucose; terminal leaflets narrow elliptic to

narrow oblanceolate to oblanceolate, 2.5-10 x

1-5 mm, tips obtuse to acute, margins slightly

recurved to revolute, smooth to glandular

dentate, midrib raised abaxially and not to

strongly glandular verrucose, adaxial surface

not or strongly glandular verrucose; lateral

leaflets similar to terminal leaflets though

usually smaller. Inflorescence axillary,

shorter to longer than the subtending leaves,

l-3(-12)-flowered; peduncle 1-7 mm long,

not or weakly glandular verrucose, pilose

with simple, bifid and trifid hairs or sparsely

stellate hairy, often hairs concentrated

between decurrent bract bases; bracts narrow-

elliptic to narrow-lanceolate, persistent,

1-4 mm long, leaflet-like; pedicels 1-5 mm

long, with a sparse to moderately dense

indumentum of stellate and/or bifid and/or

simple hairs. Sepals ovate-deltate to deflate,

1-2 x 0.7-1.5 mm, slightly imbricate in bud,

not or weakly glandular verrucose, tips acute

or rarely slightly acuminate, adaxial surface

glabrous or sparsely pilose, abaxial surface

glabrous or variously hairy. Petals elliptic, 2-

4.5 x 1.2-2.5 mm, white to pale pink, slightly

imbricate in bud, not obviously glandular or

with a few obvious glands near tip, adaxial

surface glabrous to sparsely pilose, abaxial

surface with a dense indumentum of mainly

minute stellate hairs or glabrous. Staminal

filaments 0.75-1.2 mm long, glabrous, smooth;

anthers 0.5-0.8 mm long, apiculum absent

or minute. Gynoecium glabrous or hirsute.

Cocci 3-3.5 mm x 1.75-2.5 mm, smooth to

slightly glandular verrucose, glabrous or with

a sparse to dense indumentum of simple, bifid

and/or stellate hairs, base of style sometimes

persistent forming a minute apical apiculum.

Seeds 2-2.5 x 1-1.5 mm, black, striate, with

white material between striations.

Duretto & Forster, New subspecies for Zieria odorifera from New South Wales

Distribution and habitat : Zieria odorifera

is endemic to New South Wales and occurs

in rocky areas from Kaputar N.P. and the

Inverell area east to Cathedral Range N.R and

south to Warrumbungle N.R with isolated

683

occurrences further south near Mendooran

and Molong. The species is usually found in

rocky situations in heathland, shrubland and

woodland.

Key to subspecies

1 Petals glabrous or glabrescent on abaxial surface.subsp. copelandii

1. Petals densely hirsute on abaxial surface.2

2 Branches with an indumentum of mainly stellate hairs; sepals c. 1 mm

long, glabrous or with a sparse indumentum of stellate hairs on the

abaxial surface.subsp. warrabahensis

2. Branches pilose, with no or few stellate hairs; sepals 1-2 mm long,

variously pilose on the abaxial surface though a few stellate hairs may

also be present.3

3 Petals 4-4.5 mm long [though smaller petals c. 3.5 mm long may rarely

be also present]; leaflets usually oblanceolate with smooth or slightly

glandular dentate margins.subsp. odorifera

3. Petals 2-2.5(-3.5) mm long; leaflets usually narrowly elliptic or narrowly

oblanceolate with margins slightly to prominently glandular dentate . . subsp. williamsii

Zieria odorifera J.A.Armstr. subsp.

odorifera

Illustrations : Armstrong (2002: 412,

413); Armstrong & Harden (2002: 282) [as

Z. odorifera ].

Erect or spreading shrub, to 1 m tall. Branches

moderately densely to densely pilose, with

mainly simple and bifid hairs. Leaves pilose;

petioles 1-2 mm long, not or weakly glandular

verrucose; terminal leaflets narrow elliptic to

narrow oblanceolate to oblanceolate, 3.5-10 x

2-5 mm, margins smooth to slightly glandular

dentate, adaxial surface not or slightly

glandular verrucose. Inflorescence usually

longer than the subtending leaves, 1—3(—7)-

flowered; peduncle 1.5-7 mm long, pilose with

simple, bifid and trifid hairs; pedicels 1-5 mm

long, with a moderately dense indumentum of

mainly stellate and bifid hairs. Sepals 1.5-2

x 1.2-1.5 mm, abaxial surfaces minutely

pilose, sometimes with bifid and stellate hairs

as well. Petals (3.5-) 4-4.5 x (1.7-) 2-2.5

mm, adaxial surface sparsely pilose, abaxial

surface with a dense indumentum of mainly

minute stellate hairs. Ovary hirsute; style

hirsute at base. Cocci smooth, with a sparse

to dense indumentum of simple, bifid and/or

stellate hairs or rarely (Mt Bulga) glabrous.

Additional specimens examined: New South Wales.

North Western Slopes: Summit of Mt Bulga, May

2003, Lewer & Chaffey s.n. (NSW); Walking track to

Belougery Split Rock, Warrumbungle N.R, Oct 1993,

Johnstone 414 & Burrell (MEL); Warrumbungle N.P.,

Oct 1964, Vickery & Fraser s.n. (NSW); Split Rock,

24 km NW of Coonabarabran, Dec 1973, Streimann

s.n. (CANB, NSW); Burbie Spring to Ailinn Plateau,

Warrumbungle N.P, Oct 1974, Harden s.n. (NE);

Belougery Split Rock, Warrumbungle N.P., Oct 1974,

Harden s.n. (NE); Warrumbungles Range, Sep 1966,

Frazer s.n. (NE); Warrumbungle Mts, May 1946,

Althofer 89 (NSW); 2.3 km E of Burrumbuckle Rock,

Warrumbungle N.P, Oct 1978, Crisp 4345 (CANB); Mt

Wombelong, Warrumbungle N.P., Dec 1973, Streimann

s.n. (CANB). Central-West Slopes: near Mendooran,

Sep 1945, Althofer & Dripstone 40 (MEL); c. 9 km

(direct) of Molong, Bocoble Gap, Oct 1992, Makinson

1196 (BRI, CANB, MEL, NE, NSW).

Distribution and habitat : Zieria odorifera

subsp. odorifera appears to be widespread in

Warrumbungle N.P. In addition, three isolated

collections have been made outside the park

area to the north at Mount Bulga, and south

near Mendooran and Molong. The subspecies

is found in heath, shrubland and low eucalypt

woodland on sandstones and trachyte, often

in rocky situations.

684

Phenology : Flowers have been collected

from September to December, and fruit from

October to December.

Notes : The material from Mount Bulga

differs from the remaining material of this

subspecies in having glabrous fruit and leaves

that are slightly more glandular verrucose.

Hair density on the fruits is quite variable

in material from elsewhere with the type

having sparse indumentum on the fruit,

while some other specimens have a dense

indumentum. The fruit of Zieria odorifera

subsp. williamsii may also be glabrous to

densely hirsute so the taxonomic value of this

character is questionable with such a small

sample size. Only one plant was recorded on

Mt Bulga at the time of the collection and so

further field research is required to determine

if the variation noted warrants taxonomic

recognition.

Conservation status : Few collections have

notes on population size though a few collectors

(e.g. Crisp 3609 , Johnstone 414 & Burrell ,

Makinson 1196 ) noted that the taxon was rare.

Zieria odorifera subsp. odorifera has been

mainly found in Warrumbungle N.P. though

its abundance there is unknown at this stage.

Surveys are required to ascertain the size and

extent of the known populations and the area

of occupancy, prior to making a conservation

assessment; however, a provisional assessment

would be Least Concern (IUCN 2001).

Zieria odorifera subsp. copelandii Duretto &

P.I.Forst., subspecies nova a subspecie typica

petalis sepalisque glabris differt. Typus: New

South Wales. North Western Slopes: Mount

Kaputar National Park, 5 October 2002,

L.M.Copeland 3432 (holo: NE 80023A; iso:

BRI, CANB; NSW n.v).

Erect shrub to 20 cm tall. Branches with

a sparse indumentum of simple, bifid and

stellate hairs. Leaves with a few simple hairs

on midvein and/or margin; petioles 1-1.5 mm

long, weakly glandular verrucose; terminal

leaflets narrow elliptic to narrow oblanceolate

to oblanceolate, 5-8.5 x 1.5-3.5 mm, margins

entire to slightly glandular dentate, adaxial

surface not glandular verrucose. Inflorescence

usually c. equal to subtending leaves, 1—3(—7)-

flowered; peduncle 3-7 mm long, pilose with

Austrobaileya 7(4): 681-690 (2008)

simple, bifid and trifid hairs concentrated

between decurrent bract bases; pedicels

1.5-3.5 mm long, with a sparse to moderately

dense stellate indumentum. Sepals 1-1.5 x 1-

1.3 mm, abaxial surfaces glabrous or with few

stellate hairs towards base. Petals c. 3.5 x 1.7

mm, adaxial surface glabrous or glabrescent,

abaxial surface glabrous or with a few hairs

near margins. Gynoecium glabrous. Cocci

smooth, glabrous. Fig. 1.

Additional specimens examined : New South Wales.

North Western Slopes: 33 km from Narrabri to Mt

Kaputar, Aug 1973, Muffet M3/95 (CANB); Mt Kaputar

N.P., c. 1.5 km from park entrance on road to Mt Kaputar,

Oct 1989, Jones 5182A (CANB).

Distribution and habitat : Zieria odorifera

subsp. copelandii is known only from Mount

Kaputar N.P. It has been collected from

rocky outcrops in heath with Prostanthera

cruciflora , Calytrix tetragona and Kunzea

sp.

Phenology : Flowers and fruits have been

collected in October.

Conservation status : Zieria odorifera subsp.

copelandii appears to be restricted to Mount

Kaputar N.P. with collectors ( Copeland

3432 , Jones 5182A ) noting that it was rare.

Surveys are required to ascertain the size

and extent of the known populations and

the area of occupancy prior to ascertaining

its conservation status. The restricted

distribution would indicate that a status of at

least Vulnerable based on the criterion D2 is

warranted (IUCN 2001).

Etymology : The name honours Dr Lachlan

Copeland, a botanist based in north-eastern

New South Wales, whose collections have

added much to the knowledge of the flora of

New South Wales.

Zieria odorifera subsp. warrabahensis

Duretto & P.I.Forst., subspecies nova a

subspecie typica indumento in caulibus

sepalisque ex pilis praecipue stellatis constato

(in illis expilis praecipue simplicibus vel bifidis

constato) differt. Typus: New South Wales.

North Western Slopes: Warrabah National

Park, 22 October 2006, L.M.Copeland 4113 &

D.M.Raets (holo: NE 88972; iso: BRI, CANB,

HO, NSW).

Duretto & Forster, New subspecies for Zieria odorifera from New South Wales

685

Fig. 1. Zieria odorifera subsp. copelandii. A. branch with single flowering inflorescence x 5. B. abaxial view of leaf x

8. C. inflorescence with flower x 12. D. adaxial view of petal x 30. All from Copeland 3432 (BRI). Del. B. Connell.

Erect shrub to 40 cm tall. Branches with a

moderately dense indumentum of mainly

stellate and some bifid hairs. Leaves sparsely

pilose sometimes with hairs concentrated on

midribs; petioles 1-1.5 mm long, glandular

verrucose; terminal leaflets narrowly elliptic

to oblanceolate, 2.5-5 x 1-3 mm, margins

slightly to obviously glandular dentate, adaxial

surface not glandular verrucose. Inflorescence

usually longer than the subtending leaves, 1-

3-flowered; peduncle 3-5 mm long, sparsely

stellate hairy; pedicels 1-1.5 mm long, with

686

a sparse stellate indumentum. Sepals c. lx

1 mm, abaxial surfaces glabrous or with a

sparse indumentum of minute stellate hairs.

Petals c. 2.5 x 1.2 mm, adaxial surface

sparsely pilose, abaxial surface with a dense

stellate indumentum. Gynoecium glabrous.

Cocci and seed not seen. Fig. 2.

Additional specimen examined: New South Wales.

North Western Slopes: Warrabah, W of Kingstown,

Sep 1995, Hunter 3590 (CANB, NE, NSW).

Distribution and habitat: Zieria odorifera

subsp. warrabahensis is restricted to

Warrabah N.P. where it is found growing in

skeletal sandy soils over granite in herbland,

heath, and woodland of Eucalyptus prava and

Callitris endlicheri.

Phenology: Flowers have been collected in

September and October.

Conservation status: Collectors have noted

{Copeland 4113 ) that Zieria odorifera subsp.

warrabahensis is occasional and localised

in its distribution. Surveys are required to

ascertain the size and extent of the known

populations and the area of occupancy

before providing an accurate conservation

assessment. The highly restricted distribution

would suggest that a status of Endangered

may be warranted based on the criterion C2ii

(IUCN 2001).

Etymology: This subspecies is named after

the range it is apparently restricted to.

Zieria odorifera subsp. williamsii Duretto &

P.I.Forst., subspecies nova a subspecie typica

petalis brevioribus plerumque 2-2.5 mm

longis (in illis plerumque 4-4.5 mm longis)

differt. Typus: New South Wales. Northern

Tablelands: Oxley Wild Rivers National

Park, 12 October 2002, L.M.Copeland 3437

& D.A.Carter (holo: NE 80033A; iso: BRI,

CANB, MEL, K n.v ., NSW).

Zieria sp. ‘Cathedral Rock’ (J.B.Williams

95303); Armstrong & Harden (2002: 282)

Zieria sp. ‘Oxley Wild Rivers N.P’ (Copeland

2174); Armstrong & Harden (2002: 283)

Illustrations: Armstrong & Harden

(2002: 282, as Zieria sp. ‘Cathedral Rock’

(J.B.Williams 95303); 283, as Zieria sp. ‘Oxley

Wild Rivers N.P’ (Copeland 2174)).

Austrobaileya 7(4): 681-690 (2008)

Erect or rarely prostrate or spreading shrub

to 0.5 m tall. Branches with a moderately

dense to dense indumentum of mainly simple

and bifid (occasionally trifid and stellate)

hairs. Leaves sparsely to moderately densely

pilose, sometimes hairs only on midribs;

petioles 1-2 mm long, weakly glandular

verrucose; terminal leaflets narrow elliptic to

narrow oblanceolate, (3-) 5-10 x 0.7-2 mm,

slightly or prominently glandular dentate,

adaxial surface slightly to strongly glandular

verrucose. Inflorescence shorter to slightly

longer or rarely much longer (Copeton Dam

area) than the subtending leaves, 1-3 (-12)-

flowered; peduncle 1-4 (-6) mm long, with

a moderately dense indumentum of mainly

simple and bifid hairs; pedicels 1-3 mm

long, with a moderately dense indumentum

of simple, bifid hairs or stellate hairs.

Sepals 1-1.5 x 0.7-1.3 mm, abaxial surfaces

with a moderately dense simple and bifid

indumentum sometimes with few stellate

hairs. Petals 2-2.5 (-3.5, Copeton Dam area)

x 1.2-1.5 mm long, adaxial surface sparsely

pilose, abaxial surface with a moderately

dense to dense indumentum of mainly stellate

hairs. Gynoecium glabrous to hairy on outer

margin. Cocci slightly glandular verrucose,

glabrous to densely pilose. Fig. 3, 4.

Additional specimens examined: New South Wales.

Northern Tablelands: Sydenham near Barraba, Oct

1914, Rupp s.n. (NSW); Inverell, Aug 1910, Boorman

s.n. (NSW); Goonoowigall Bushland Reserve, near

The Peaks, c. 6 km S of Inverell, Feb 1993, Sipple 44

& Beckers (NE); Murchison S.F., W side of Copeton

Dam, Feb 2001, Copeland 2836 & Westaway (CANB,

NE, NSW); 0.5 km N of Howell, Sep 1985, Southwell

H85-042 (CANB); 2.4 km from Howell on track to

Copeton Lake recreation area, Sep 1985, Southwell

H85-039 (CANB); Copes Creek, S of Gilgai, Nov 1989,

Williams s.n. (NE, NSW); Howell district, 70 km N of

road to Copeton Dam, 8.4 km by road W of Inverell

- Bundarra road, Oct 2001, Copeland 3222 (CANB, NE,

NSW); Howell, Aug 1905, Boorman s.n. (HO, CANB,

MEL, NE, NSW); 2 km E of Howell, Sep 1966, Williams

s.n. (NE50498A); 3 km E of Howell, Sep 1966, Williams

s.n. (NSW651474, NE50532A); ibid , Dec 1971, Williams

s.n. (H0544742, NE28958); 3.2 km E of Howell, Nov

1973, Armstrong 683 (NSW); ibid , Dec 1971, Williams

s.n. (HO, NE); 1 km S of Howell, Oct 1989, Griffith &

Williams s.n. (NE); 2 km S of Howell, 1967, Williams

s.n. (BRI [AQ510733], NSW651470); Single N.P, Mar

2000, Copeland 2487 (NE, NSW); The Basin Nature

Reserve, c. 20 km E of Bundarra, 2.8 km S of ‘Lutana’,

Nov 2001, Copeland 3260 & Croft (C ANB, NE); Parlour

Mountains, ‘Hardacres’ property, c. 38 km NW of

Duretto & Forster, New subspecies for Zieria odorifera from New South Wales

687

Fig. 2. Zieria odorifera subsp. warrabahensis. A. branch with several flowering inflorescences * 8. B. abaxial view

of leaf x 10. C. lateral view of flower x 20. D. indumentum on leaf bearing branchlet x 20. All from Copeland 4113

(BRI). Del. B. Connell.

Armidale, Aug 1991, Hardaker s.n. (NE); Snowy Range,

Biffen’s Block, 6.5 km WNW of Round Mt, Nov 1995,

Metcalf & Williams s.n. (NE); The Pinnacle, 20 km W

of Armidale, Jun 2007, Telford 13165 & Bruhl (HO, NE,

NSW); The Pinnacle, 24 km W of Armidale, Oct 1975,

Wissman s.n. (NSW); Biffen’s property. Maiden Creek

- adjoins western side of Cathedral Rock N.P, c. 15 km

E of Wollomombi, Nov 1995, Williams 95303 & Metcalf

(NSW); Oxley Wild Rivers N.P, Nov 1999, Copeland

2174 et al. , Nov 1999 (CANB, NE, NSW).

Distribution and habitat : Zieria odorifera

subsp. williamsii is known from scattered

populations between Inverell, Cathedral

Rock N.R and Oxley Wild Rivers N.R It

is found on various rock types including

granite, porphyry and other volcanics in

heath, shrubland (dominated by, e.g. Acacia

neriifolia ) and woodland (dominated by, e.g.

Eucalyptus prava, E. laevopinea, E. dealbata

or Callitris endlicheri).

Phenology: Flowers have been collected from

August to November, and fruit from August

to December (March, June).

688

Notes : Zieria sp. ‘Cathedral Rock’ (Williams

95303) and Z. sp. ‘Oxley Wild Rivers N.P.’

(fide Armstrong & Harden 2002) were

separated from other species of Zieria on

a few characters that included plant habit,

inflorescence size relative to leaves, number of

flowers per inflorescence and the pubescence

of the cocci. These characters were found to be

unreliable when a large number of collections

were examined from the area between Howell

and Cathedral Rock. Across the geographic

range of Z. odorifera subsp. williamsii plants

can have glabrous or hirsute fruit, and be

erect, prostrate, ascending, etc.; thus these

proposed diagnostic characters do not have

much taxonomic relevance, they are simply

variation. In addition, the collections from

Howell that were included with Z. odorifera

by Armstrong (2002) and Armstrong &

Harden (2002) do not have petals c. 4.5 mm

in length but c. 2.5 mm in length. Zieria

sp. ‘Cathedral Rock’ (Williams 95303) and

Austrobaileya 7(4): 681-690 (2008)

Z. sp. ‘Oxley Wild Rivers N.P.’ are based on

a few specimens that are forms of the taxon

recognised here which is variable in plant

habit and in the amount of pubescence on the

leaves and fruit (Fig. 3 & 4). The pubescence

of the abaxial surface of the petals, and the

composition of the indumentum is uniform

and thus of taxonomic value for Z. odorifera

(see key above).

Specimens from the Copeton Dam (e.g.

Copeland 3222 ) and Inverell (e.g. Sipple 44

& Beckers ) areas are almost intermediate

between Zieria odorifera subsp. odorifera

and the more ‘typical’ forms of Z. odorifera

subsp. williamsii in that they have longer

peduncles (to 6 mm long), inflorescences

that are much longer than the leaves, larger

petals (to 3.5 mm long) and leaves that are not

always glandular dentate. These somewhat

intermediate specimens are allocated with

a little difficulty to the latter subspecies and

Fig. 3 .Zieria odorifera subsp. williamsii (less hairy variant). A. branch with several flowering inflorescences x 4. B.

abaxial view of leaf x 8. C. detail of abaxial leaf surface showing indumentum cover x 20. D. inflorescence with flower

and buds x 8. E. tip of sepal x 20. F-G. views of stamens x 20. All from Copeland 3437 (BRI). Del. W. Smith.

Duretto & Forster, New subspecies for Zieria odorifera from New South Wales

689

Fig. 4. Zieria odorifera subsp. williamsii (more hairy variant). A. branch with several flowering inflorescences *

2. B. abaxial view of leaf x 6. C. detail of abaxial leaf surface showing indumentum cover x 20. D. inflorescence with

flower and buds x 8. E. tip of sepal x 20. F-G. views of stamens x 20. All from Williams s.n. (BRI [AQ510733]). Del.

W. Smith.

are one of the reasons why these taxa are

being recognised at that rank. Field research

with detailed population studies are required

to determine if the variation seen on these

few specimens is a true reflection of what is

occurring in situ. Nevertheless outside these

areas the smaller petals (2-2.5 mm long) and

the narrow leaflets with glandular dentate

margins easily separate Z. odorifera subsp.

williamsii from the typical variety.

A putative hybrid between Zieria

odorifera subsp. williamsii and Z. cytisoides

{Copeland 3439 & Carter ) has been collected

in the Oxley Wild Rivers N.P. The single plant

was isolated and plants of both parent species

were present nearby (collector’s notes).

Conservation status : In some areas collectors

(e.g. Copeland 3260, Williams 95303 ) have

indicated that this taxon is rare while elsewhere

(e.g. Armstrong 683, Telford 13165 ) it was

noted as being common though localised. It is

known from Oxley Wild Rivers and Cathedral

Rock N.R’s. Surveys are required to ascertain

the size and extent of the known populations

and the area of occupancy prior to ascertaining

an appropriate conservation status, though it

can be provisionally assessed as Endangered

on the basis of criterion B2a,b(i-v) (IUCN

2001 ).

Etymology : This subspecies is named in

honour of John B. Williams (1932-2005) who

did much to advance Australian botany (see

Bruhl et al. 2005).

Acknowledgements

We thank Brian Connell and Will Smith for

the illustrations; Peter Bostock for translation

of the diagnoses into Latin and the distribution

map; and the Directors of CANB, MEL, NE

and NSW for making available material for

research.

Austrobaileya 7(4): 681-690 (2008)

690

References

Armstrong, J.A. (2002). Zieria (Rutaceae): a systematic

and evolutionary study. Australian Systematic

Botany 15: 277-463.

Armstrong, J.A. & Harden, G.J. (2002). Zieria. In G.

Harden (ed.) Flora of New South Wales, 2 nd

edition, 2: 277-288. The University of New

South Wales Press: Sydney.

Bruhl, J.J., Bale, C., Boyd, R., Boyd, K., Clark,

D., Harden, G., Telford, 1, Whalley, W. &

White, G. (2005). Vale John B. Williams 1932-

2005. Australian Systematic Botany Society

Newsletter 124: 9-14.

Duretto, M.F. & Forster, PI. (2007). A taxonomic

revision of the genus Zieria Sm. (Rutaceae) in

Queensland. Austrobaileya 7: 473-544.

Iucn (2001). IUCN Red List Categories and Criteria.

Version 3.1. IUCN Species Survival

Commission. IUCN: Gland, Switzerland and

Cambridge, UK.

Three new species of Rhodomyrtus (DC.) Rchb.

(Myrtaceae) from Papua New Guinea 5

Neil Snow 14 , Jessie McFadden 2 & Joshua P. Atwood 1 ’ 3

Summary

Snow, N., McFadden, J., & Atwood, J.P. (2008). Three new species of Rhodomyrtus (DC.) Rchb.

(Myrtaceae) from Papua New Guinea. Austrobaileya 7(4): 691-706. Three new species of

Rhodomyrtus (DC.) Rchb. are described from Papua New Guinea: Rhodomyrtus longisepala N. Snow

& J.McFadden, Rhodomyrtus guymeriana N.Snow & J.P.Atwood, and Rhodomyrtus misimana

N.Snow. Detailed descriptions, information on distributions and habitats, and conservation

recommendations are provided for each new species. A tabular summary of some key vegetative and

reproductive features is presented for species of Rhodomyrtus sensu lato. A membranous testa for

three species of Rhodomyrtus is discussed for the first time, as is the occurrence of densely-packed,

hair-like fibres comprising much of the outer testa in some species. Large, dark-maroon coloured

cells in the fruit of some species impart visible pits on the outer surface of the seed coats in the dried

fruits. Gelatinous endosperm is reported for the first time in the genus Rhodomyrtus , being present

in Rhodomyrtus guymeriana. Transitions between the typological distinctions of eucamptodromous

and brochidodromous leaf venation types are discussed for Rhodomyrtus surigaoensis.

Key Words: Rhodomyrtus longisepala, Rhodomyrtus guymeriana, Rhodomyrtus misimana,

Rhodomyrtus surigaoensis, Myrtaceae, conservation, Papua New Guinea, Indonesia, seed

morphology, systematics, biogeography, Owen Stanley Terrane, Kutu Terrane, Finisterre Terrane

'N.Snow, Herbarium Pacificum, Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817,

U.S.A. Email: neil.snow@bishopmuseum.org

2 J.McFadden, Conifer High School, Conifer, Colorado 80433, U S A.

3 J P.Atwood, Department of Natural Resources Science, University of Rhode Island, Kingston,

Rhode Island 02881, U.S.A.

4 author for correspondence

5 Pacific Biological Survey contribution no. 2008-032 from the Bishop Museum

Introduction

The genus Rhodomyrtus (DC.) Rchb. comprises

22 currently recognized species (Table 1), which

includes several species described (Guymer 1991;

Snow 2006; herein) since the revision by Scott

(1978). The native range of Rhodomyrtus extends

from India and Sri Lanka to the Philippines

south through Malesia to northeastern Australia

and New Caledonia and Fiji (Scott 1978; Snow

2000, 2006). One species, Rhodomyrtus

tomentosa (authorities in Table 1), the type

species of the genus, is a highly aggressive weed

in some tropical and semi-tropical areas, such

as Hawaii and Florida (Wagner et al. 1999). The

purpose of this paper is to describe three new

species from Papua New Guinea and evaluate

their conservation status, discuss and provide

Accepted for publication 14 July 2008

a tabular summary of some key vegetative and

reproductive characters of Rhodomyrtus.

Materials and methods

Characters from herbarium specimens

were measured following a descriptive

format similar to Snow (2007). Descriptive

terminology generally follows the Systematics

Association Committee (1962). Leaf venation

follows Hickey (1973; see also Simpson

(2006)), whereas herbarium acronyms follow

Holmgren & Holmgren (1998). Flowering

material was unavailable for Rhodomyrtus

guymeriana and/?, misimana , but descriptions

are as complete as possible given limited

material. Two of the new species are known

only from their type gatherings, whereas

R. misimana is known from two gatherings.

Fruits and seeds were dissected from several

specimens housed at BISH (and one from K,

with permission) by rehydrating for several

minutes in boiling water.

692

Taxonomy

Rhodomyrtus guymeriana N.Snow et

J.P.Atwood, species nova R. eleganti affinis

sed fructu costato differt. Typus: New

Guinea. Central Province; Mori River, c.

15 km northeast of Cape Rodney, 10°04'S,

148°32'E, 30 August 1969, R. Pullen 8167

(holo: A; iso: BRI, CANB [2 sheets], K).

Trees to 6 m tall. Bark of main stem unknown.

Branchlets laterally compressed, wingless;

epidermis villous (hairs whitish to reddish),

oil glands absent. Stipules (“enations” of

some authors) of several dark, relatively

long setose hair-like structures (“type

A” stipules sensu Snow et al. 2003: 6-7).

Petioles 6-18 mm, flat above, eglandular,

villous to tomentose becoming glabrous.

Leaves coriaceous, discolorous, surfaces

matt, venation brochidodromous. Leaf

blades 9-16 x 3.3-7.5 cm, obovate to elliptic;

base cuneate; apex mostly acute and often

mucronate (infrequently broadly obtuse);

margin flat. Adaxial leaf surface glabrescent

to glabrate; midvein relatively broad, flush or

slightly raised proximally; secondary veins

distinct (tertiary veins less so); oil glands

not visible. Abaxial leaf surface sparsely to

moderately villous, especially on midvein and

secondary veins; secondary and tertiary veins

raised; intramarginal veins at midpoint of

leaf prominent, 1-2.5 mm from leaf margin;

margin flat; oil glands common but prominent

only with magnification. Inflorescence lateral,

evidently a solitary monad (more material

needed). Flowering material unknown. Fruit

(submature) baccate, broadly fusiform, 35-

42 x 20-27 mm, 9-ribbed (ribs up to 5 mm

deep), shortly villous throughout, pinkish

when fresh; calyx lobes erect. Locules (from

fruit) evidently 2 (or if 3 then one crushed);

placentation axile. Seeds at least 5 (only half

of one dried fruit available for analysis),

embedded in pulpy flesh that adheres tightly

to the thickly membranous testa. Embryo

tightly circinate, c. 7 mm long and 3.2 mm

thick at widest point, rounded, densely but

minutely glandular; endosperm gelatinous

but scant. Fig. 1, 2.

Distribution and habitat : Attempts herein

to reconcile the coordinates indicated on

Austrobaileya 7(4): 691-706 (2008)

the collection label with modern satellite

imagery (Google Earth®, 2008) suggest that

the collection site from the Mori River was

approximately 160 km southeast of Port

Moresby and c. 19 km northeast (not 15 km

as per label) of Port Rodney (Map 1). This

interpretation seems feasible since a road

cutting through forest meets the Mori River

at c. 10°04'24.85"S, 148°30'55.45"E, which

corresponds closely to the stated elevation

of the type gathering of c. 30 m. If this

interpretation is correct, it suggests the species

occurs on the Kutu Terrane (Pigram & Davies

1987). The same recent satellite imagery

indicates moderate levels of deforestation in

an arc from the vicinity of Saint Stephens

Mission, approximately 8 km to the northeast

of the presumed collection site, more or less

continuously to the south-southeast, roughly

3 km from the presumed collection site. The

amount of potential alluvial habitat nearby

that appears to be mostly forested includes an

area of approximately 12.5 linear km to the

southeast towards the town of Domara. To

the northwest of the apparent collection site

much higher amounts of deforestation have

occurred west of the Mori River.

Phenology : Known only from immature fruit

in late August; likely flowering at least 2-4

weeks earlier.

Notes: Rhodomyrtus guymeriana most

closely resembles Rhodomyrtus elegans The

species are similar by virtue of their relatively

long and narrow petioles, leaf shapes and

texture, and mucronate apices on some (but

not all) leaves. The species differ by the

brochidodromous venation (Hickey 1973)

and barely perceptible (if at all) oil glands on

the adaxial leaf surface of R. guymeriana ,

versus the eucamptodromous venation and

dense, small, but easily seen oil glands on the

adaxial leaf surface of R. elegans. In addition,

the short pubescence on the secondary and

tertiary veins on the abaxial leaf surface is

denser on R. guymeriana than R. elegans. It

should be noted that some specimens included

by Scott (1978) in R. elegans may represent

another undescribed species.

Rhodomyrtus guymeriana and R. elegans

differ most prominently in their fruiting

Snow et al ., New species of Rhodomyrtus

693

Fie** O' new fiuim*

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Fig. 1. Rhodomyrtus guymeriana. Holotype, Pullen 8167 (A)

694

Austrobaileya 7(4): 691-706 (2008)

Table 1. Some key vegetative and reproductive characters of Rhodomyrtus s.l.

Venation patterns (Hickey 1973) can be brochidodromous (br), eucamptodromous (eu) or

acrodromous (ac). The hypanthium in the flower can be smooth (sm), longitudinally ribbed (lr),

or winged (wi). Placentation is axile (ax) or parietal (pa). Fruits are pulpy (pu) or not pulpy

(np). Seed arrangement in the locules (“seeds”) is irregular (ir) or stacked (st). Thin horizontal

membranous partitions between seeds (“partitions”) are present (+), absent (-), or consisting of

large, dark maroon cells embedded into the testa (dmc). Outer testal layers are sclerotic (sc) or

membranous (mb). Embryos are C-shaped (c) or circinate (ci). Data taken from specimens at

BISH; character states not yet known or uncertain designated by “?” Excluding polymorphisms

within a species and uncertainties, at least nine permutations of these variables occur among the

species (see Discussion).

Species

venation

hypanthium

placentation

fruit

seeds

partitions

testa

type

embryo

shape

R. canescens

C.T.White & Francis

sm, lr

R. effusa Guymer

R. elegans (Blume)

A. J. Scott

R. guymeriana

N.Snow & J.P. Atwood

R. kaweaensis N.Snow

R. Ianata Guymer

dmc

R. locellata

(Guillaumin) Burret

dmc

R. longisepala N.Snow

& J.McFadden

R. macrocarpa Benth.

R. mengenensis

N.Snow

R. misimana N.Snow

R. montana Guymer

R. novoguineensis

Diels

R. obovata C.T.White

+, -

R. pervagata Guymer

R. pinnatinervis

C.T.White

dmc

R. psidioides (G.Don)

Benth.

+ 5 -

R. salomonensis

(C.T.White) A. J. Scott

ac, eu

dmc

R. sericea Burret

sm, lr

R. surigaoensis Elmer

dmc

R. tomentosa (Aiton)

Hassk.

R. trineura (F.Muell.)

F.Muell. ex Benth.

+ 5 -

Snow et al. , New species of Rhodomyrtus

695

Fig. 2. Rhodomyrtus guymeriana. Close up of fruit on holotype showing prominent longitudinal wings.

morphology. Scott (1978) indicated that

fruits of R. elegans could be smooth or

have approximately 10 ribs. All of the type

specimens for R. guymeriana , which Scott

(1978) included in R. elegans , have nine

deep longitudinal ridges, and the furrows

between each ridge are flat. The flat furrow,

as opposed to having a U- or V-shaped profile,

along with the thin but evenly thickened ribs

and their height of up to 5 mm above the

fruit surface, indicates the ribs may appear

more wing-like when fresh, although their

prominence may decrease as the fruits

increasingly mature and, presumably, become

more plump. Longitudinal ribbing in fruits is

a homoplastic character state that recurs in

many baccate genera of Myrtaceae (e.g. Snow

et al. 2003, Snow 2007; Salywon & Landrum

2007). Although the ribbing can be weak or

only moderate in some species of Myrtaceae

(e.g., Rhodamnia angustifolia N.Snow &

Guymer and R. dumicola Guymer & Jessup

(Snow 2007)), we are unaware of species in

which variation in the cross-sectional shape

of fruits ranges from completely lacking in

ribs to strongly ribbed. We have re-examined

some fruiting specimens cited by Scott (1978)

as R. elegans (e.g., Ridsdale NGF31720,

Streimann & KairoNGF17479, both at BISH),

which with their globose overall and terete

cross-sectional fruit shapes are quite unlike

those of R. guymeriana , with its fusiform

overall shape and nine longitudinal ridges. A

further but subtle difference exists between

the indumentum of the fruits. Mature fruits of

R. elegans bear individual and appressed hairs

that are less than 0.3 mm long; on older fruits

(e.g. Ridsdale NGF31720 , BISH) the fruit

sheds some of its indumentum with increasing

maturity. In contrast, individual hairs on

the sub-mature fruits of R. guymeriana are

approximately 0.5 mm long and somewhat

696

more erect. Finally, the seeds of R. elegans

(e.g. Ridsdale NGF31720, BISH) appear to be

arranged haphazardly between the irregularly

oriented but relatively thick partitions. This

arrangement contrasts with the regular

vertical rows (typically 2 per locule in each of

3 locules) separated by thin partitions, typical

for the acrodromous species of Rhodomyrtus.

Conservation status : In the absence of

focused surveys IUCN (2001) guidelines

suggest a conservation assessment of “Data

Deficient” (DD). Callmander et al. (2005),

citing the urgent global need for more rapid

and effective conservation of plants, presented

compelling reasons for avoiding designations

of “data deficient”, particularly as they relate

to tropical taxa, and suggested instead the use

of either “threatened/potentially threatened”

or “not threatened”. One of the more poignant

reasons in their view for the latter strategy is

that targeted, species-specific field surveys

are unrealistic across much of the tropics and

have lead to significant underestimates of

the percentage of plants worldwide that are

threatened. Underestimations of threatened

taxa interfere with planning efforts that require

knowledge of threatened taxa and an accurate

assessment of overall rarity (Callmander et

al. 2005). The potentially available habitat

nearby for Rhodomyrtus guymeriana suggests

a designation of “not threatened” (IUCN

2001), but because the species has not been

collected elsewhere we feel the designation

of “Potentially Threatened” following the

recommendations of Callmander et al. (2005)

is more appropriate.

Etymology : The specific epithet honours

Dr. Gordon P. Guymer (b. 1953), Director of

the Queensland Herbarium (BRI) and fellow

student of Australasian Myrtaceae (Guymer

& Jessup 1986; Guymer 1988a, b; Snow &

Guymer 1999a,b; Snow & Guymer 2001a,b;

Snow et al. 2003).

Rhodomyrtus longisepala N.Snow et

J.McFadden, species nova a speciebus ceteris

omnibus generis lobis calycis acuminatis,

venatione acrodrma distincta, venis

trichomatibus moderate usque dense tectis

abaxialiter, et a R. novoguineensi Diels foliis

ad apicem acuminatioribus, lobis calcyis

Austrobaileya 7(4): 691-706 (2008)

longioribus differt. Typus: Papua New

Guinea. Morobe Province: Labu Swamp,

lowland rainforest on margin of swamp,

woodline on landslip, 6°42'S, 146°53'E, 11

July 1989, W. Takeuchi 4550 (holo: BISH [2

sheets]; iso: BISH [plus spirit], BRI, CANB,

NY [2 sheets]).

Shrubs to 3.5 m tall; branches numerous

and arching. Bark of main stem unknown.

Branchlets rounded, wings lacking; epidermis

brownish, becoming fissured, tomentose-

villous and reddish-brown (increasingly

whitish and tomentose with age), oil glands

not visible. Stipules hair-like (“type A” sensu

Snow et al. 2003: 6-7). Petioles 5-6.5 mm,

densely tomentose-villous (hairs reddish-

brown becoming whitish); adaxial surface

flattened. Leaves coriaceous, discolorous,

surfaces matt, venation perfect (sometimes

imperfect) basal or suprabasal acrodromous.

Leaf blades 9.3-14.1 x 2.8-4.5 cm, narrowly

elliptic; base cuneate to rounded; apex

acuminate and usually somewhat falcate;

margin flat to slightly revolute. Adaxial leaf

surface (at maturity) with scattered appressed

hairs on midvein but otherwise glabrous;

midvein impressed, becoming flush distally

at apex, secondary veins distinct but tertiary

veins faint; oil glands not evident on dried

material. Abaxial leaf surface villous, hairs

reddish-brown and arising primarily from

veins; secondary and tertiary veins raised;

intramarginal veins at midpoint of leaf c. 0.5

mm from leaf margin; margin flat; oil glands

small and sparse but visible when backlit.

Inflorescence terminal or mostly axillary,

solitary or rarely two per axil; flowers mostly

solitary or less frequently in simple cymes

or few-flowered racemes; leaf bracts present

near apex of peduncle, 16-29 mm long,

moderately to densely villous, sometimes

acrodromous; peduncles 1-11 mm, densely

tomentose-villous (hairs reddish-brown).

Bracteoles 2, 4.5-9 mm long, 0.5-1 mm wide

(at base), narrowly elliptic, apex attenuate

and exceeding base of calyx lobes in flower,

densely villous, typically soon deciduous.

Hypanthium obconic, 7-10 mm, densely

villous (hairs reddish-brown), cross-section

round. Calyx lobes 5, lobes free above, distinct

in bud, 5-8 mm, narrowly triangular, apex

Snow et al ., New species of Rhodomyrtus

697

Fig. 3. Rhodomyrtus longisepala. A. Branchlet; abaxial portion of leaf in detail showing indumentum (middle

of far right). B. Abaxial (upper figure) and adaxial (lower figure) views of flower with detail of stamens and style.

C. Longitudinal section (left) of developing fruit illustrating vertical placement of ovules/young seeds; young fruit

illustrating indumentum, bracteoles, and elongate calyx lobes. All from Takeuchi 4550 (BISH). Del. B. Angell.

698

acuminate, densely villous above and below

(hairs reddish-brown), ascending to erect in

fruit. Petals white, 5-7.5 x 4-5.2 mm, ovate

to broadly obovate or elliptic, apex rounded

to acute, sometimes asymmetric at base;

adaxial surface glabrous, sparsely glandular;

abaxial surface densely villous throughout.

Stamens multiseriate, at least 100 (more

accurate counts not possible with existing

material); filaments 2-4 mm; staminal disk

and ovary disk densely short hairy (hairs

whitish); anther sacs subcylindrical, 0.6-0.8

mm, basifixed to sub-basifixed, dehiscing

via longitudinal slits, eglandular. Ovary 3-

locular, placentation axile, ovules numerous

in 2 distinct vertical rows per placenta. Styles

c. 6 mm, glabrous; stigma somewhat capitate.

Fruit baccate, globular to subcylindrical at

maturity, rounded at base, 18-19 x 14-16

mm, densely tomentose-villous but becoming

nearly glabrous at maturity, reportedly

(collection label) white when ripe. Seeds

(from point of attachment) broadly obovate to

oblate, c. 2 mm long; horizontal membranes

between seeds lacking (Snow 1999, 2000);

testa hard, light yellowish-brown, individual

cells increasingly elongated and enlarged

distally from point of attachment. Embryos

c-shaped. Fig. 3.

Distribution and habitat: Rhodomyrtus

longisepala is known only from the type

gathering from the western part of Labu

Swamp, from lowland rainforest on the

swamp margin west of Lae (Map 1). This

low-lying area is composed primarily of

sediments largely derived from the adjacent

Owen Stanley (to the south) and Finisterre (to

the north) terranes.

Phenology : Known only at the late-flowering,

early-fruiting stage in mid-July. Flowering

likely commences at least 2-3 weeks earlier,

and mature fruits could be expected to last

into early September, if not longer.

Notes: When fresh the leaves are said to

be coriaceous, medium green above, and

yellowish-green below with a reddish-brown

tomentum. According to the collection label

the specimen has numerous long and arching

branchlets.

Austrobaileya 7 ( 4 ): 691-706 ( 2008 )

On New Guinea, Rhodomyrtus

longisepala is a close morphological match

in many respects for the relatively common

R. novoguineensis , but the latter has shorter,

rounded calyx lobes. Amongst the Australian

species, R. longisepala somewhat resembles

R. pervagata, which occurs in northeast

Queensland at elevations of 400-1250 metres

(Guymer 1991). The latter also generally

has rounded (rather than attenuate) calyx

lobes, although in Ford 3295 (BISH) they

are somewhat more acute. Likewise, the

Australian taxon R. trineura may have acute

calyx lobes, although they are considerably

shorter than those of R. longisepala.

This new species was collected as part of

anticancer and anti-AIDS screening for the

National Cancer Institute (sample numbers

Q6607319-R, Q6607320-S, Q6607321-T,

Q6607322-U, and Q6607323-V).

Conservation status: Recent satellite imagery

suggests the area immediately adjacentto Labu

Swamp is little if at all deforested. However,

since the species Rhodomyrtus longisepala

is only known from one gathering the

appropriate designation for it is “Potentially

Threatened”, based on the rationale provided

above for R. guymeriana.

Etymology: The specific epithet is in reference

to the attenuate calyx lobes, which distinguish

it from its congeners.

Rhodomyrtus misimana N.Snow, species

nova R. montanae et R. lanatae affinis

sed foliis subtus non nisi sparsim villosis

differt. Typus: Papua New Guinea. Milne

Bay Province: Misima Island, Mt Oeatau,

26 March 1979, K. Damas LAE74597 (holo:

BISH; iso [n.v.]: A, BRI, CANB, E, K, L,

LAE).

Shrubs to trees, 2-14 m tall. Outer bark light

brown, thin and papery; inner bark light green,

thinly fibrous. Youngest branchlets and leaves

densely villous with reddish-brown hairs, the

adaxial leaf surfaces becoming increasingly

glabrate during ontogeny. Branchlets rounded,

wings lacking; densely villous (to somewhat

tomentose) with reddish-brown hairs.

Stipules of several dark, relatively long setose

hair-like structures (“type A” sensu Snow et

al. 2003: 6-7). Petioles 3.5-5 mm, densely

Snow et al ., New species of Rhodomyrtus

699

HOLOTYPE 44li62

cilia aliiim-liu X. £nov

ilueifuballcya 7C4): . 2008.

Ty-no coni' If ec J bv S, Si

Lat ■ r i Long .: ■

Habitat : njntona ro.rfttt te>o3*y),

I'i-u unpo.atu, larieuoute, r.ldrib

brasn proniriant bale*, surd

■. r ios-uru axiel. auJatiify, bfiiry

.. ,-l _-L,r. ItMlfl. fell till

iK-luicai Collect Iona of tfce Xatloaei

Kertarius, DPI, OiTlee of Furttti , Lae.

Hi. Mo,,log. Kiaie-a laUmi.

u. „u Hlnino* Pro vines Kilns Boy,

Fig. 4. Rhodomyrtus misimana. Holotype specimen (Damas LAE74597 [BISH]).

700

villous with reddish-brown hairs, adaxial

surface channeled (but this obscured by dense

indumentum). Leaves coriaceous, strongly

discolorous (drying dark brown adaxially),

surfaces matt, venation perfect, suprabasal

acrodromous. Leaf blades (1.7-) 5-9.2 x (l-

)1.5—2.5 cm, narrowly ovate, base cuneate to

rounded; apex acute to acuminate; margin flat.

Adaxial surface indumentum (at maturity)

comprised of appressed to somewhat villous

hairs along midvein and lateral primary

veins (whitish to reddish-brown), becoming

less dense proximally; midvein impressed

proximally, becoming nearly flush distally,

secondary veins faint; oil glands not evident

on dried material. Abaxial leaf surface villous,

hairs reddish-brown and mostly arising from

veins; secondary and tertiary veins prominent

and slightly raised; intramarginal veins at

midpoint of leaf <0.5 mm from margin; oil

glands small but dense. Inflorescence axillary

or terminal, flowers 1 or mostly 2 per axil;

pedicels 3.5-5.5 mm long, densely villous.

Bracteoles 2, very broadly ovate, 2.5-3 mm

long, leaf-like, sparsely hairy adaxially,

densely hairy abaxially. Hypanthium obconic

3-4.5 mm long, densely villous, cross-section

round. Calyx lobes 5, free and distinct in

bud, broadly rounded apically, densely

villous, c. 2.5 mm long. Petals sparsely to

moderately villous adaxially, densely villous

abaxially (material otherwise limited and

only fragmentary). Stamens unknown. Style

c. 6.5 mm, somewhat hairy proximally. Ovary

(from fruit) 3-locular, placentation axile, each

locule bearing 2 distinct vertical rows of seeds

and thin septal walls. Fruit baccate, 5.5-7

x 4-5 mm (at submaturity), subcylindrical,

rounded at base, densely villous becoming

less so, reportedly (from collection label)

greenish when young, crowned by persistent

calyx lobes. Horizontal, thin membranes

present between individual seeds (evidently

complete or nearly so) (Snow 1999, 2000).

Testa sclerotic. Embryos not seen but probably

c-shaped based on seed shape. Fig. 4 & 5.

Additional specimen examined : Papua New Guinea.

Milne Bay Province: Louisiade Archipelago, Misima

Island, Mt Oiatau, Mar 1979, Harrison-Gagne 2155

(BISH).

Distribution and habitat : The new species

has been collected at c. 800 and 980 metres

Austrobaileya 7(4): 691-706 (2008)

in montane, mossy forests on Misima Island

(Map 1), which has a narrow east-west axis

approximately 40 km long and 10 km wide

at its greatest dimensions. According to

Pigram & Davies (1987), Misima Island and

the larger southeasterly islands of Tagula, and

Rossel (Yela), which are located in the Milne

Bay Province, are geological outliers of the

Owen Stanley Terrane, which has a much

greater expanse as a prominent component

of the mountainous areas (some exceeding

3800 metres) along the interface of the Gulf,

Morobe, Central, and Northern provinces.

These outlying islands are also known as

the Louisiade Archipelago, a predominantly

volcanic series of islands with associated

coral reefs. The collection label of the

holotype indicates coordinates of 11°22'S,

154°11'E. However, these coordinates are

probably in error, as they fall on Yela (Rossel)

Island, which is located some 145 km to the

southeast of Misima Island. The holotype

of Rhodomyrtus misimana was reported

previously as a range extension of R. montana

(Csizmadi 2006: 54).

Phenology : Fruiting (immature) known only

in late March. The flowering period likely

commences in early February and fruiting

may extend through much of April or later.

Notes : The two collections of Rhodomyrtus

misimana are virtually indistinguishable. The

paratype specimen is also from Mt. Oeatau

and was collected on the same day (likely as

part of the same collecting party).

Rhodomyrtus misimana closely resembles

R. montana. However, the abaxial indumentum

on the leaves of the new species arises from

the venation only, and the oil glands are dense

and easily visible; in R. montana it is densely

tomentose-villous throughout the abaxial

surface (Fig. 6). In addition, R. misimana has

a matt adaxial leaf surface and predominantly

villous indumentum on the branchlets,

whereas R. montana has a glossy adaxial

surface and indumentum on the branchlets

that is more tomentose but includes longer,

villous hairs. Rhodomyrtus misimana occurs

from 800-940 metres elevation on Misima

Island on Mt Oeatau (as “Oiatau” in some

spellings), which is approximately 235 km

Snow et al ., New species of Rhodomyrtus

east of the southeastern-most part of New

Guinea (Papua Peninsula; Map 1). In contrast,

R. montana is restricted to 1900-2300 metres

in the Arfak (Mt. Kobreimot) and Nettoti

ranges in the Vogelkop Peninsula of Papua

Province, Indonesia, approximately 2400 km

northeast of Misima Island.

Rhodomyrtus misimana also resembles

R. lanata. However, the latter species has

much larger leaves (6-13.3 cm long [Guymer

1991]) and shares with R. montana the densely

tomentose-villous abaxial leaf surface.

Rhodomyrtus lanata occurs mostly in the

Morobe and Central Provinces of Papua New

Guinea at elevations from 1100-2300 metres,

although one collection is known from

Sandaun (= West Sepik) Province (Sayers

NGF19502 ) (Guymer 1991).

Conservation status: The collection label of

the holotype indicates the plant was common

when collected in 1979. The species appears

to be restricted to an elevational range of

approximately 180 metres on one island, and

we thus feel the most appropriate designation is

“Potentially Threatened” (following rationale

cited above for Rhodomyrtus guymeriana).

Etymology: The specific epithet reflects the

evident endemism of the species to Misima

Island.

Discussion

The description of the three new species

of Rhodomyrtus provides the opportunity

to summarise and discuss some of the

characters that differentiate species of the

genus as it is presently circumscribed (Table

1), and to report several character states for

the first time. Ignoring polymorphisms and

uncertainties contained therein, Table 1

reveals at least nine permutations of character

states among those presented, even though it

only partially summarizes character variation

in Rhodomyrtus. For example, indumentum

on petals and stigma shapes have been shown

to be of taxonomic utility in Rhodomyrtus and

related genera (Csizmadi 2006; Snow 2008)

but are not reported, since they need additional

study. The most common character syndrome

shown in Table 1 occurs in Rhodomyrtus

effusa and five other species (R. longisepala ,

701

R. montana , R. novoguineensis, R. pervagata,

and R. trineura), all of which have acrodromous

venation (sensu Hickey 1973).

The gelatinous endosperm found in

Rhodomyrtus guymeriana is the first report

for the genus. It is also known for some

species of Gossia (Snow et al. 2003) and has

been reported for Eugenia (van Wyk & Botha

1984), but is otherwise rare among baccate

genera in the family. Other species, especially

those with circinate embryos and fibres

surrounding the testa (see below), should

be re-examined with fresh material for the

presence of endosperm.

The circinate embryos of Rhodomyrtus

guymeriana are shared by R. elegans

(Hartley TGH10646 [A]) and R. macrocarpa

(Brass 33718 [QRS], Smith 112392 [BISH]).

The embryos of these three species are also

densely but minutely glandular. Among

the baccate genera of Myrtaceae, circinate

embryos are known also in Gossia (Snow et

al. 2003), Campomanesia Ruiz & Pav., and

Blepharocalyx salicifolius (H.B.K.) O.Berg

(Landrum & Kawasaki 1997). Species of

Rhodomyrtus possessing circinate embryos

also have a membranous testa (details

below) and either eucamptodromous or

brochidodromous venation; the distinction

between venation types was not recognized

previously for the genus (e.g. Csizmadi 2006;

Snow 2006). Re-analysis of the leaf venation

pattern of R. surigaoensis indicates a type

somewhat intermediate between acrodromous

and brochidodromous. Snow (2007) similarly

reported limitations of the use of leaf venation

typologies for some species of Rhodamnia.

At least two of the three newly described

species have membranous outer testal layers

(Rhodomyrtus elegans and R. guymeriana)

that also exhibit a type of testal morphology

previously unreported for the genus, and which

may represent a first report for Myrtaceae.

This form possesses several layers of

parallel, tightly packed, relatively long fibres

that are tightly appressed to the outer testa

(R. guymeriana [isotype, K] and R. elegans

[Ridsdale NGF31720 , BISH]). The fibres

are oriented radially across the more or less

circular seed and collectively impart a thickly

702

Austrobaileya 7(4): 691-706 (2008)

Fig. 5. Close-up of abaxial leaf surfaces highlighting differences in density of indumentum. Upper half: Rhodomyrtus

misimana, holotype (Damas LAE74597 [BISH]); Lower half: Rhodomyrtus montana, Sleumer & Vink BW14152

(BISH).

Snow et al ., New species of Rhodomyrtus

membranous texture to the seed. The fibrous

testa surrounds a much thinner membranous

layer that lies adjacent to the embryo. These

non- or only weakly sclerified fibres may

represent a transitional evolutionary stage

between a membranous testa and sclerotic

testa, the latter of which is common in many

genera of the Myrteae (Wilson et al. 2005;

which also has been known widely as subtribe

Myrtinae [e.g., Landrum & Kawasaki 1997;

Landrum & Sharp 1989]).

The presence of a membranous testa, as

seen in three species of Rhodomyrtus (Table 1)

is of interest because few genera in the Myrteae

(sensu Wilson et al. 2005) from Australasia

or those from the neotropics (Landrum &

Kawasaki 1997) share this character state. The

mostly Australian genus Pilidiostigma has a

leathery testa (Snow 2004), and the recently

erected monotypic Brazilian genus Curitiba

Salywon & Landrum has a coriaceous testa

(Salywon & Landrum 2007). Among species

of Rhodomyrtus those with a membranous

testa have either brochidodromous or

eucamptodromous leaf venation. Preliminary

DNA sequence analyses using sequences

from ITS-1 and ITS-2 that sampled broadly

in Rhodomyrtus and putatively related genera

have placed Pilidiostigma near (and in one case

sister to) a clade that included many (but not

exclusively) species of Rhodomyrtus having

brochidodromous venation and a membranous

testa (Csizmadi 2006: 31; Salywon et al. in

prep.). The results of Wilson et al. (2005;

Fig. 1 therein) using DNA sequences from

matK also suggested a close phylogenetic

relationship between the eucamptodromous-

veined species R. macrocarpa Benth. and

Pilidiostigma , although their sampling regime

was limited within genera.

Another character reported here for the first

time among the baccate genera of Myrtaceae

(to the best of our knowledge) is large, dark

maroon-coloured cells (as observed on dried

fruits that have been cut in trans-section),

which are tightly adherent to the outer wall of

the seed coats (e.g. in Rhodomyrtus locellata,

R. pinnatinervis, R. salomonensis, and

R. surigaoensis). The large cells are easily

visible without magnification and leave visible

pits in the surface of the testa, suggesting the

703

large cells assume their shape prior to the

sclerotization of the outer testal cells. Species

of Rhodomyrtus with the enlarged dark maroon

cells all have eucamptodromous venation,

apart from R. surigaoensis , whose venation is

intermediate between eucamptodromous and

acrodromous.

Species of Rhodomyrtus with axile

placentation (all but R. macrocarpa) typically

have one of two types of seed arrangement.

In most species the seeds are stacked like

poker chips in two regular rows per ovule,

with little if any pulpy tissue between the

seeds (e.g., as per Psidium guajava L.);

this group includes the type of the genus,

R. tomentosa (e.g. Fosberg 37709, BISH). In

contrast, species such as R. guymeriana and

R. elegans have seeds irregularly embedded

in a juicy, pulpy matrix in which the seeds are

separated by somewhat irregularly oriented

partitions. Rhodomyrtus macrocarpa differs

from all other species in the genus by its

parietal placentation and unilocular carpels,

although its eucamptodromous venation and

membranous testa bearing radial fibres on the

outer-most layers suggest a close relationship

with R. psidioides and R. elegans. The

character variation (Table 1) for fruiting

material was derived mostly from dried

material, and additional studies using fresh

material with larger sample sizes are not

only needed, but may require some of the

information herein to be amended.

With the description of these three new

species, the number of species now known

for Rhodomyrtus s. lat. is 22, a figure

approximately double that of earlier reports

(Scott, 1978; Snow 2000). Rhodomyrtus

trine ura (Guymer 1991) and R. tomentosa

(Scott 1978) have each been recognized as

having infraspecific taxa, raising the total

number of taxa in the genus to as high as 24.

Additional collections at BISH and elsewhere

suggest that one or more additional new species

of Rhodomyrtus may await description from

New Guinea, but better reproductive material

must be seen first.

With 13 species, the island of New Guinea

has the highest diversity of Rhodomyrtus,

compared to seven species in Australia.

704

Rhodomyrtus macrocarpa occurs in both areas

(New Guinea comprises Papua New Guinea

and Papua Province of Indonesia) (Snow

2006) . (Note: “ Rhodomyrtus longipetiolatcT

[Snow 2006: 338] should have read Gossia

longipetiolata N.Snow.) Of the five species of

Rhodomyrtus described recently (Snow 2006

and herein), four are known only from the

type gathering. Given that many regions of

New Guinea harbor localized endemic species

of vascular plants (van Welzen 1995) and that

much of the island remains barely if at all

surveyed (Stevens 1989; Conn 1994; Takeuchi

2007) , we predict that further exploration will

reveal additional novelties in Rhodomyrtus.

As reported previously, Rhodomyrtus

is demonstrably polyphyletic based on

morphology (Snow 1999,2000) and ITS-1 and

ITS-2 sequence data (Csizmadi 2006; Snow

2006). The genus almost certainly will be split

in the future pending additional data. Given the

acrodromous leaf venation of R. tomentosa (the

type species of the genus) and R. longisepala

and R. misimana , and aspects of their fruit

and seed morphology, it seems unlikely the

latter two taxa will require name changes. In

contrast, the brochidodromous venation in

R. guymeriana , coupled with some aspects of

its fruit morphology, suggest it may require a

new generic combination in the future along

with several other species in the genus. Future

studies of Rhodomyrtus should sequence

additional molecular markers and carry

out detailed analyses of floral and fruiting

morphology using fresh material. However,

before Rhodomyrtus is split into (presumably)

two or more monophyletic genera, additional

sequencing and morphological studies also

are needed for the putatively related genera

Archirhodomyrtus Burret, Octamyrtus Diels,

and Kanakomyrtus N.Snow (Snow 2008).

Acknowledgments

We thank curators at A, BRI, CANB, K, L,

NY, QRS and US for loaning material or access

to specimens. For R. longisepala we extend

our appreciation to Lyn Craven (CANB) for

the Latin diagnosis and Bobbi Angell for the

beautiful illustration. Laura Jennings and Eve

Lucas (K) provided assistance with specimens

and permission to dissect a fruit. Peter Wilson

Austrobaileya 7(4): 691-706 (2008)

(NSW) graciously read the manuscript

before its submission and suggested several

improvements. Peter Bostock (BRI) provided

the blank map for PNG for Map 1. Mahalo

to Arnold Hori (BISH) for providing some

of the digital images herein. The description

of R. longisepala represents a portion of a

M.Sc. thesis (Csizmadi 2006) submitted to

the University of Northern Colorado by J.M.,

who graciously acknowledges support at

UNC from the Albert M. Winchester, Robert

Sund, Richard and Chris Monfort, Cross

Family, and Gerald D. Schmidt awards. The

internship of J.P A. at the Bishop Museum was

supported by a NSF grant (IGERT 0504103)

to Peter August, University of Rhode Island

Coastal Institute. N.S. thanks William and

Christiane Anderson (MICH) for assistance

interpreting the correct generic authorship of

Rhodomyrtus.

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Guymer, G.P. (1988a). Rhodamnia pauciovulata , a

new species of Myrtaceae from Queensland.

Austrobaileya 2: 515-516.

- (1986b). A new species of Backhousia Hook. &

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reappraisal of Backhousia floribunda A. J. Scott.

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Austrobaileya 2: 228-234.

Hickey, L.J. (1973). Classification of the architecture of

dicotyledonous leaves. American Journal of

Botany 60: 17-33.

705

Snow et al. , New species of Rhodomyrtus

Holmgren, P.K. & Holmgren, N. H. (1998 onwards;

continuously updated). Index Herbariorum.

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of some American Myrtinae (Myrtaceae):

Psidium and related genera. Systematic Botany

14: 370-376.

Landrum, L.R. & Kawasaki, M.L. (1997). The genera

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706

Austrobaileya 7(4): 691-706 (2008)

Map 1 . Distribution of three new species of Rhodomyrtus from Papua New Guinea. R. guymeriana • :

R. longisepala ♦ .and R. misimana («—, lower right).

Plectranthus batianoffii P.I.Forst. (Lamiaceae), a

new species from north-east Queensland

Paul I. Forster

Summary

Forster, P.I. (2008). Plectranthus batianoffii P.I.Forst. (Lamiaceae), a new species from north-east

Queensland. Austrobaileya 7(4): 707-710. A new species ( Plectranthus batianoffii P.I.Forst.) known

from several continental islands in north-east Queensland is described and illustrated. A conservation

status of Vulnerable is recommended.

Key Words: Lamiaceae, Plectranthus batianoffii , new species, Australian flora, Queensland flora

P.I.Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens

Mt Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: paul.forster@epa.

qld.gov.au

Introduction

The genus Plectranthus is highly diverse in

Australia with at least thirty-five described

species (Blake 1971; Forster 1992,1994,1996,

1997, 1999). As noted previously (Duretto &

Forster 2007), speciation in the genus has

occurred where populations occur in isolated

areas such as mountain peaks or ranges

and where suitable habitat exists on areas

of exposed rock pavements and outcrops.

This suite of habitat factors has resulted in

the “islands on islands” effect (Porembski

et al. 2000) and is widely recognized as

being a driving force in genetic diversity

and speciation (Carlquist 1974; Grant 1981;

Hopper 2000; Seine et al. 2000). The rapid

maturation of Plectranthus individuals from

seed (one growing season) with the potential

for repeated generation turnover is thought

to enable relatively rapid speciation within a

given locality once new genetic changes arise

in a population.

Accumulation of a number of field

collections, together with a critical

examination of previously described taxa and

herbarium collections, now enable a further

new species to be named. The new species

has a restricted distribution on several

offshore continental islands in north-east

Queensland.

Accepted for publication 15 September 2008

Materials and methods

This paper is based on collections in

Australian herbaria, particularly BRI and

QRS. Live material of Plectranthus batianoffii

was cultivated for nearly 20 years in Brisbane

enabling observation of variation under

different ecological conditions.

Taxonomy

Plectranthus batianoffii P.I.Forst., species

nova P. foetido Benth. affinis, sed habitu

suffruticis decumbentis usque ad 30 cm

altitudine non fruticiformi usque ad 2 m

altitudine, foliis vix odoratis non valde

aromaticis, pagina inferiore folii laminae

trichomatibus antrorsis non-glandularibus

(adversum divaricatis vel retrorsis) praedita,

verticillastris flores plures (12-18 non 6-11)

ferentibus differens. Typus: Queensland.

Cook District: Palfrey Island, 14°42'S,

145°28'E, 23 July 1990, G.N.Batianoff 12118

(holo: BRI [1 sheet]).

Prostrate to semi-erect herb to subshrub up

to 30 cm high; foliage with ± no scent when

crushed, slightly clammy; non-glandular and

glandular trichomes clear to purplish, sessile

glands 8-celled and orange. Roots fibrous.

Stems ± square, erect to straggling, succulent,

the lower parts up to 10 mm diameter,

grey-green, non-glandular trichomes sparse,

antrorse, 6-10-celled up to 2 mm long,

glandular trichomes scattered, sessile glands

scattered. Leaves discolorous, petiolate;

petioles 5-18 mm long, 3-4 mm wide, weakly

channelled above, non-glandular trichomes

708

dense, antrorse, 6-12-celledupto 3.2 mm long,

glandular trichomes absent, sessile glands

scattered; lamina broadly ovate to triangular,

± succulent, 30-60 mm long, 30-50 mm wide,

crenate with 15-19 teeth up to 2 mm long on

each margin, widest above middle, secondary

teeth usually present; tip acute; base truncate;

upper surface grey-green, veins impressed and

velutinous, non-glandular trichomes dense,

divaricate to antrorse, 6-12-celled up to 1.5

mm long, glandular trichomes absent, sessile

glands absent; lower surface silver-green,

veins raised and velutinous, non-glandular

trichomes dense, antrorse, 6-12-celled up

to 2 mm long, glandular trichomes absent,

sessile glands dense. Inflorescence up to

200 mm long, comprising 1-3 pedunculate

branches from near the base; verticillasters

12-18-flowered, up to 8 mm apart; pedicels

1.6-1.8 mm long, c. 0.3 mm diameter,

non-glandular trichomes sparse, antrorse,

6-8-celled up to 1 mm long, glandular

trichomes scattered, sessile glands absent;

cymes sessile; axis square in cross-section,

non-glandular trichomes sparse, antrorse,

6-12-celled up to 2 mm long, glandular

trichomes absent, sessile glands scattered;

bracts broadly-ovate, 1.6-1.8 mm long, 2-2.2

mm wide, not forming a coma, non-glandular

trichomes sparse, antrorse, 6-10-celled up

to 1 mm long, glandular trichomes absent,

sessile glands sparse. Flowering calyx 3.2-

3.5 mm long, non-glandular trichomes dense,

antrorse, 6-8-celled up to 1.2 mm long,

glandular trichomes scattered, sessile glands

sparse. Fruiting calyx 3.8-4.5 mm long; upper

lobe obovate to ovate, 2-2.5 mm long, 1.8-2.2

mm wide; lateral lobes lanceolate, 1.9-2.2

mm long, 0.6-0.8 mm wide; lower lobes

lanceolate-falcate, 2-2.3 mm long, 0.5-07

mm wide. Corolla 8.5-10 mm long, pale lilac;

tube 4.5-5.5 mm long, weakly curved at 110—

120° 2-2.5 mm from base, slightly inflated

upwards, glabrous or with non-glandular

trichomes scattered, divaricate, 2-4-celled

to 0.3 mm long; upper lobes suborbicular,

recurved, 1.7-1.9 mm long, 1.6-2 mm wide,

non-glandular trichomes sparse, divaricate,

2-4-celled up to 0.4 mm long, glandular

trichomes absent, sessile glands sparse;

lateral lobes oblong, 1-1.2 mm long, 0.5-0.7

Austrobaileya 7(4): 707-710 (2008)

mm wide, non-glandular trichomes absent,

glandular trichomes absent, sessile glands

scattered; lower lobe broadly ovate, 3.2-4.5

mm long, 3.5-4 mm long, non-glandular

trichomes sparse, divaricate, 2-6-celled up

to 0.5 mm long, glandular trichomes absent,

sessile glands scattered. Filaments filiform,

7-8 mm long, c. 0.3 mm diameter, lilac, fused

for 4-5 mm from base; anthers c. 0.4 mm

long and 0.3 mm wide. Style filiform, 7-8

mm long, lilac, bifid for c. 0.3 mm. Nutlets

± circular in outline, ± flattened, c. 0.9 mm

long, 0.8 mm wide and 0.5 mm thick, smooth,

glossy brown. Fig. 1.

Additional specimens examined : Queensland. Cook

District: Stanley Island, Jun 1995, Le Cussan 606

(BRI); [all subsequent collections from Lizard Island]

Oct 1967, Heatwole 75 (BRI); Jul 1969, Heatwole s.n.

(BRI [AQ007940]); Dec 1974, Specht LI218 & Specht

(BRI); Dec 1974, Specht LI416 & Specht (BRI); Sep

1988, Batianoff 10216 (BRI); Jul 1990, Batianoff12087

(BRI, MEL). Cultivated: Indooroopilly (ex Lizard

Island), Nov 1994, Forster 15894 (BRI).

Distribution and habitat: Plectranthus

batianoffii has been found only on Lizard,

Palfrey and Stanley Islands on the north-east

Queensland coast. Plants occur on granite

outcrops and pavements or occasionally in

boulder strewn grassland.

Notes: Plectranthus batianoffii was first

collected in 1967 by Hal Heatwole; however,

the taxonomic identity of the plants was not

questioned until the late 1980s when George

Batianoff first collected material.

On the basis of gross morphology, the

most obvious allied species to Plectranthus

batianoffii appears to be P. foetidus Benth.

which is distributed to the south of Lizard and

Palfrey Islands. The new species differs from

P. foetidus by the decumbent subshrub habit

to 30 cm high (versus a shrub up to 2 m tall),

the barely scented foliage (versus strongly

aromatic), the lower surface of the leaf lamina

with antrorse non-glandular trichomes (versus

divaricate to retrorse) and the vertillasters

with more flowers (12-18 versus 6-11).

Etymology: The specific epithet honours

George N. Batianoff, botanist at the

Queensland Herbarium and specialist on

Queensland’s island floras.

Forster, Plectranthus batianoffii

709

Fig. 1. Plectranthus batianoffii. A. flowering stem x 0.5. B. abaxial view of leaf x 1.5. C. adaxial view of leaf x 1.5.

D. inflorescence bract x 6. E. lateral view of flower x 6. F. face view of flower x 6. All from Forster PIF15894 (BRI).

Del. W. Smith & B. Connell.

Conservation status : Plectranthus batianoffii can be assessed as Vulnerable on the criterion

is known from three populations, all situated D2 (IUCN 2001).

on rather small offshore islands. The species

710

Acknowledgements

Thanks to George Batianoff for bringing

this species to the attention of the author and

collecting live material; Will Smith and Brian

Connell for the illustrations and Peter Bostock

for the Latin diagnosis.

References

Blake, S.T. (1971). A revision of Plectranthus (Labiatae)

in Australasia. Contributions from the

Queensland Herbarium No. 9: 1-120.

Carlquist, S. (1974). Island Biology. Columbia

University Press: New York.

Duretto, M.F. & Forster, PI. (2007). A taxonomic

revision of the genus Zieria Sm. in Queensland.

Austrobaileya 7: 473-544.

Forster, PI. (1992). Five new species of Plectranthus

(Lamiaceae) from Queensland. Austrobaileya

3: 729-740.

- (1994). Ten new species of Plectranthus L’Her.

(Lamiaceae) from Queensland. Austrobaileya

4: 159-186.

Austrobaileya 7(4): 707-710 (2008)

- (1999). Plectranthus fasciculatus (Lamiaceae), a

new species from north-eastern Queensland,

Australia. Haseltonia 6: 14-16.

Grant, V. (1981). Plant Speciation, 2 nd edition. Colombia

University Press: New York.

Hopper, S.D. (2000). Floristics of Australian granitoid

inselberg vegetation. In S.Porembski &

W.Barthlott (eds.), Inselbergs. Ecological

Studies 146: 391-407.

Iucn (2001). IUCN Red List Categories and Criteria.

Version 3.1. IUCN Species Survival

Commission. IUCN: Gland, Switzerland and

Cambridge, UK.

Porembski, S., Becker, U. & Seine, R. (2000). Islands on

islands: habitats on inselbergs. In S. Porembski

& W.Barthlott (eds.), Inselbergs. Ecological

Studies 146: 48-67.

Seine, R., Porembski, S. & Becker, U. (2000).

Phytogeography. In S. Porembski & W. Barthlott

(eds.), Inselbergs. Ecological Studies 146: 435-

449.

_ (1996). Plectranthus alioplectus , P. gratus and P.

spectabilis (Lamiaceae), a trio of rock-outcrop

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Haseltonia 4: 47-56.

- (1997). Plectranthus amoenus and

P. thallassoscopicus (Lamiaceae), new species

from north-eastern Queensland, Australia.

Austrobaileya 4: 653-660.

Huperzia tetrastichoides A.R.Field & Bostock

(Lycopodiaceae) a newly recognised species of tassel

fern from the Wet Tropics of Queensland, Australia

Ashley RField 1 & Peter D.Bostock 2

Summary

Field, A.R. & Bostock, P.D. (2008). Huperzia tetrastichoides A.R.Field & Bostock (Lycopodiaceae), a

newly recognised species of tassel fern from the Wet Tropics of Queensland, Australia. Austrobaileya

7(4): 711-715. An Australian endemic species of tassel fern Huperzia tetrastichoides A.R.Field and

Bostock is described as new. This species was previously confused with the non-Australian species

Huperzia prolifera (Blume) Trevis. The new species is restricted to upland Queensland rainforests

from the Windsor Tableland south to the Clarke Range west of Mackay. It is listed as Vulnerable under

the Queensland Nature Conservation Act 1992. A dichotomous identification key to the Australian

species of Huperzia is provided.

’A.R.Field, School of Marine and Tropical Biology, James Cook University, Douglas Townsville

Queensland 4811 Australia. Email: ashley.field@jcu.edu.au\

2 P.D.Bostock, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens,

Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: peter.bostock@epa.qld.gov.au

Key Words: Huperzia , Lycopodium , Huperzia prolifera , Huperzia tetrastichoides, tassel-fern,

Lycopodiaceae

Introduction

Huperzia Bernh. is the largest genus of the

Lycopodiaceae, an ancient and cosmopolitan

plant family (Ollgaard 1987). Twelve species

of Huperzia have been recorded in Australia

(Bostock & Holland 2007). Ten of these

species are epiphytic or epilithic in rainforests

in the Queensland tropics (Chinnock 1998).

Following examination of living plants and

herbarium specimens from Asia and Australia

we have concluded that the entity hitherto

recognised as Huperzia prolifera (Blume)

Trevis. ( sensu Andrews 1990; Chinnock

1998; Goodger et al. 2008) in Australia is

not conspecific with that Malesian species

(Blume 1828) (photos of lectotype sheets at

Leiden L0057380 and L0057381 seen). It is

described in this paper as a new species and

considered to be endemic to Australia.

Materials and methods

This study is based on field observations and

collections in north Queensland and Malaysia,

together with examination of herbarium

collections at BRI, CANB and CNS (formerly

QRS).

Accepted for publication 23 September 2008

Taxonomy

Huperzia tetrastichoides AR.Field &

Bostock, species nova antea H. prolifera

confusa, a qua microphyllis triangulari-

ovatis, carinatis et orthostichis in seriebus

quatuor (non lineari-lanceolatis, ut minimum

lycophyllis planis et microphyllis omnibus

heterostichis) differt; ad H. tetrasticham

arctissime accedit, a qua lycophyllis ad

angulum 20-50° patentibus non aequaliter

adpressis et imbricatis differt. Typus:

Queensland. Cook District: Millaa Millaa

Falls, Atherton Tableland, 14 June 2004,

A.R.Field and O.Rawlins 1139 (holo: BRI; iso:

CNS, distribuendi).

Sporophyte: Epiphytic herbaceous plant with

indeterminate isodichotomous pendulous

shoots arising from a tufted root system.

Lycophylls and sporophylls subopposite,

decussate, orthostichous in four rows.

Lycophylls triangular-ovate, carinate, thin

but firm, 5-12 mm long, 2-6 mm wide, in

basal sterile divisions diverging 20°-50° from

stem, gradually transforming to adpressed,

imbricate sporophylls in sharply quadrangular

terminal fertile divisions. Sporophylls

triangular, carinate, thin but firm, 4-6 mm

712

Austrobaileya 7(4): 711-715 (2008)

Fig. 1. Huperziaprolifera. A. infertile shoot section and B. fertile shoot section; Huperzia tetrastichoides. C. infertile

shoot section and D. fertile shoot section; Huperzia tetrasticha. E. infertile shoot section and F. fertile shoot section ;

A-F scale bar = 2 cm; Huperzia tetrastichoides. G. habit showing isodichotomous pendant branching, carinate sterile

lycophylls and sporophylls, lycophylls arranged in four rows and quadrangular strobili; G scale bar = 5 cm. A & B from

scan of L57380 and L57381; C & D, G from Field ARF1139 (BRI); E & F from Field ARF0815 (BRI). Del. A. Field.

713

Field & Bostock, Huperzia tetrastichoides

long, c. 3 mm wide at base. Sporangia c. 2 mm

in diameter, each completely concealed by

its subtending sporophyll. Stems pale green,

lycophylls and sporophylls usually dull dark

green. Gametophyte: unknown.

Additional specimens examined [precise localities

withheld]: Queensland. Cook District: Mossman, May

2004, Field ARF723, Lloyd & Toh (BRI); NW of Julatten,

May 1989, Jones 4266 & Clements (BRI, CANB);

Rumula, Oct 2004, Field & Field ARF826 (BRI); Black

Mt, Jul 1999, Jago 5307, Wannan & Worboys (BRI).

North Kennedy District: Koombooloomba, Feb 2003,

Gray 8439 (CANB); Paluma, Apr 2003, FieldARF613 &

Cairns (BRI). South Kennedy District: Eungella, Jan

1994, Pollock 127 & Pearson (BRI); Eungella, Feb 2005,

Field ARF888 & Field (BRI); Crediton, Jun 2001, Ford

AF2871 (BRI, QRS).

Distribution and habitat : Huperzia

tetrastichoides is an uncommon canopy

epiphyte of upland notophyll vineforest from

Mt Finnigan at c. 15°50’S, south to the Clarke

Range west of Mackay at c. 21 °S (Map 1). It

is most prevalent on the Evelyn, Atherton and

Mt Carbine Tablelands and descends to lower

altitudes in Mossman Gorge. No specimens

have been recorded from outside Australia

thus it is considered endemic to the Wet

Tropics of Queensland.

Notes : Huperzia tetrastichoides has

carinate, triangular to ovate microphylls

that are orthostichous in four rows, whereas

H. prolifer a has mostly flat, linear to lanceolate,

heterostichous microphylls (Fig. 1A-B, G).

Huperzia tetrastichoides resembles the

Malesian species Huperzia tetrasticha

(Fig. 1E-F) but differs from it by having

divergent rather than uniformly adpressed,

imbricate lycophylls (Fig. 1C-D). Huperzia

tetrastichoides resembles the endemic

Australian species Huperzia marsupiiformis

(D.L.Jones & B.Gray)Holub and Huperzia

lockyeri (D.L.Jones & B.Gray)Holub. It

differs from Huperzia marsupiiformis by

having carinate, triangular-ovate, acutely

pointed lycophylls rather than flat, oval, blunt

lycophylls and from H lockyeri by having

carinate not flat lycophylls and by strobili that

are quadrangular throughout rather than terete

in some parts.

Conservation status : Currently listed as

vulnerable under the Queensland Nature

Conservation Act 1992.

Etymology : The specific epithet reflects

the similarity of this species to Huperzia

tetrasticha ; it is formed from the epithet

tetrasticha based on the Latin tetra, four of,

and stichos, a row or line of things, and the

suffix -oides, like or resembling.

Common name : This species is commonly

known as the ‘bootlace tassel fern’ or the

‘Queensland square tassel fern’ in the nursery

trade.

Key to Australian Huperzia

1 Plants with erect fertile shoots.2

1. Plants with pendant or nodding fertile shoots.3

2 Lycophylls entire; bulbils present in lycophyll axils.H. australiana

2. Lycophylls serrate; bulbils lacking.H. serrata

3 Shoots homophyllous; sporophylls spreading widely.4

3. Shoots heterophyllous; sporophylls adpressed.5

4 Shoots glaucous blue grey; lycophylls lanceolate.H. dalhousieana

4. Shoots glossy green; lycophylls linear-lanceolate.H. squarrosa

5 Strobili 1-2 mm thick.6

5. Strobili 3-5 mm thick.7

6 Lycophylls sessile, decurrent in four distinct rows . . .

6. Lycophylls petiolate, not decurrent in four distinct rows

H. phlegmarioides

. . H. phlegmaria

714

Austrobaileya 7(4): 711-715 (2008)

.H. filiformis

7 Lycophylls thin and linear, margins minutely serrate

7. Lycophylls lanceolate to ovate, margins entire.8

8 Lycophylls carinate or cupped in cross section.9

8. Lycophylls flat in cross section.10

9 Basal lycophylls triangular-ovate, arranged in four rows; lycophylls and

sporophylls thin and coriaceous.H. tetrastichoides

9. Basal lycophylls lanceolate, arranged in more than four rows; lycophylls

and sporophylls thick and succulent.H. carinata

10 Lycophylls with obtuse apices.H. marsupiiformis

10. Lycophylls with acute apices.11

11 Sporophyll apices attenuate and spreading; stems fleshy.H. lockyeri

11. Sporophyll apices obtuse to acute and recurved; stems woody.H. varia

Acknowledgments

This project was supported by a grant from the

Queensland Government ‘Growing the Smart

State Program’. A.Field wishes to thank Holly

Field, Harry and Rita Kupke, and Owen and

Coral Rawlins for access to additional live

Huperzia.

References

Andrews, S.B. (1990). Ferns of Queensland. Queensland

Department of Primary Industries: Brisbane.

Blume, K.L. (1828). Enumeratio plantarum Javae 2:

261-273. J.W. van Leeuwen: Leiden.

Bostock, P.D. & Holland, A.E. (2007). Census of

the Queensland Flora 2007. Queensland

Herbarium, Environmental Protection Agency:

Brisbane.

Chinnock, R.J. (1998). Lycopodiaceae. In P.M.McCarthy

(ed.). Flora of Australia 48: 66-85. ABRS/

CSIRO Publishing: Canberra/Melbourne.

Goodger, J.Q.D., Whincup, A.L., Field, A.R., Holtum,

J A M. & Woodrow, I.E. (2008). Variation in

huperzine A and B in Australasian Huperzia

species. Biochemical Systematics and Ecology

36: 612-618

0llgaard, B. (1987). A revised classification of the

Lycopodiaceae s. I at. Opera Botanica 92: 253-

278.

Field & Bostock, Huperzia tetrastichoides

715

Corsia dispar D.L.Jones & B.Gray (Corsiaceae), a

new species from Australia, and a new combination

in Corsia for a New Guinea taxon

David L. Jones 1 & Bruce Gray 2

Summary

Jones, D.L. & Gray, B. (2008). Corsia dispar D.L.Jones & B.Gray (Corsiaceae), a new species from

Australia, and a new combination in Corsia for a New Guinea taxon. Austrobaileya 7(4): 717-722.

Corsia Becc., a highly specialised monocotyledonous genus, has been recorded from Australia for

40 years; however, the single known species has not been previously studied in detail, nor formally

described. The new species C. dispar D.L.Jones & B.Gray is named and illustrated and notes on its

taxonomy and ecology are provided. Corsia wiakabui (Takeuchi & Pipoly) D.L.Jones & B.Gray is

newly recognised at specific rank, based on C. purpurata L.O.Williams var. wiakabui Takeuchi &

Pipoly.

Key Words: Corsiaceae, Corsia, Corsia dispar, Corsia purpurata var. wiakabui, Corsia wiakabui,

new species, new combination, Queensland flora, Australian flora. New Guinea flora

‘D.L. Jones, cl Centre for Plant Biodiversity Research, Australian National Herbarium, GPO Box

1600, Canberra, A.C.T. 2601, Australia.

2 B.Gray, c/ Australian Tropical Herbarium, PO. Box 6811, Cairns, Queensland 4870, Australia.

Introduction

Corsia Becc. is a genus of about 25 species

developed primarily in New Guinea where

there are c. 23 species; two others occur in

the Solomon Islands and a solitary species is

recorded for northern Australia (Van Royen

1972). Somewhat orchid-like, and often

mistaken for a member of that group of plants,

they can be immediately distinguished by the

presence of a ring of stamens surrounding

a more or less central style. The stamens

are connate basally and with the style form

a partially united structure. They are not

fully fused, unlike the gynostemium in

Orchidaceae. Additionally their pollen grains

are free and not coherent to form pollinia

(Rudall & Eastman 2002).

Species of Corsia are seasonally deciduous,

mycoheterotrophic terrestrial herbs which are

usually discovered by accident rather than

design. They grow in high rainfall montane

forests where there is an accumulation of

surface litter. Most species grow in areas

of difficult access, have a short or limited

flowering period, are only above ground when

flowering or fruiting, and their life cycle is

Accepted for publication 15 September 2008

intimately associated with periods of heavy

rainfall. The majority of species appear to

be narrow endemics with a restricted range.

In fact Van Royen (1972) notes that in New

Guinea “almost every mountain range seems

to have its own species”. All species are

generally so elusive and difficult to see in the

gloomy surroundings where they grow, that

they are undoubtedly under collected, although

occasionally they have been observed to be

locally abundant (Cribb 1985).

The Australian species of Corsia was

first collected by Bruce Gray in 1968 and

identified to generic level by Jim Willis at

MEL. Correspondence by the senior author

with Pieter Van Royen established that it

was an undescribed species with affinities to

C. unguiculata Schltr. from New Guinea, but

with a number of distinctive features which

readily separated it from all New Guinea

taxa. Van Royen (1972) placed all the then

known taxa in two sections, both of which he

described.

The family Corsiaceae has traditionally

been grouped with Burmanniaceae and

Thismiaceae in the Burmanniales (Dahlgen et

al. 1985); however, a phylogeny inferred from

718

large-subunit (26S) ribosomal DNA sequences

suggests that the order Burmanniales is

polyphyletic (Neyland 2002). A second

similar phylogenetic study also suggests that

the Corsiaceae is polyphyletic and provides

compelling evidence for the inclusion of

Corsia within the Liliales (Neyland &

Hennigan 2003).

Generic features of Corsia

Since Corsia has some interesting specialised

characters and is of very limited distribution

in Australia, some notes on its generic features

may be pertinent. Species of Corsia are highly

specialised, achlorophyllous, cryptic herbs

with a mycoheterotrophic life style. The plants

are leafless, with fine filamentous roots which

ramify for surprisingly long distances through

leaf litter. The rhizome is much reduced and

grows in increments which terminate in

a flower stem. The thin, wiry scape bears

sheathing bracts and a single terminal flower

which, in the New Guinea species, is erect

in bud. The perianth segments consist of

three petals and two lateral sepals which are

similar or subsimilar in shape and size. The

third sepal is greatly expanded and modified

to form a labellum (note that in Orchidaceae

the labellum is a modified petal). All species

of Corsia from New Guinea have the labellum

held erect or more or less held horizontally like

an umbrella above the stamens and style, and

all have six stamens. The staminal filaments

are connate basally and also fused with

the base of the style to form a column-like

structure. The filaments are erect in bud and

also in the early stages of anthesis and then

the anthers fall off and the filaments recurve,

leaving the style to elongate and the stigmatic

lobes to expand and become receptive. The

fruit is an elongated three-valved capsule with

the valves spreading outwards at maturity,

the placentas detaching from the outer walls

and the seeds hanging from long thread-like

funicles.

Infrageneric Classification of Corsia

Two sections, Sessilis and Unguiculatis,

were recognised within Corsia by Van

Royen (1972), but as section Sessilis contains

C. ornata Becc., the designated type of the

genus, it must be recognised as section Corsia

Austrobaileya 7(4): 717-722 (2008)

(ICBN, autonym rule, article 22). Additionally

the adjectival epithet unguiculatis should be

Unguiculatae (ICBN, article 21.2). In both

section Corsia and section Unguiculatae , the

ovary is straight or slightly curved, the buds

are erect, the labellum is held erect or above

the sepals and petals like an umbrella and there

are six functional stamens and no staminodes.

These sections can be distinguished thus:

Section Corsia - Petals and sepals linear,

rarely ovate-linear, usually hanging down

beneath the labellum; basal callus attached

to the ovary with a broad base. Type species:

C. ornata Becc.

Section Unguiculatae Van Royen - Petals

and lateral sepals ovate, much shorter than the

labellum, curved inwards to form a cup-like

structure which holds the stamens and style;

basal callus a narrow linear lamella. Type

species: C. unguiculata Schltr.

Unique features of the Australian species

The Australian species of Corsia is quite

divergent from all New Guinea taxa and differs

in at least four significant morphological

features.

1. The ovary is strongly arcuate to uncinate

and as a consequence the buds are cernuous

(ovary straight to slightly arcuate in New

Guinea taxa, with erect buds).

2. The labellum is held below the perianth

segments (held erect or above the perianth

segments like an umbrella in the New Guinea

taxa).

3. The lateral sepals and petals recurve

away from the stamens and style (in section

Unguiculatae these organs hang down beneath

the labellum, whereas in section Corsia they

curve inwards to form a cup-like structure

which holds the stamens and style).

4. The flowers have five stamens and a

staminode (six stamens and no staminodia in

the New Guinea species).

The Australian species does not align into

either of the two sections as circumscribed

above. A new infrageneric taxon may be

required; however, this should wait until a

detailed phylogenetic analysis is carried out.

719

Jones & Gray, Corsia dispar

Taxonomy

Corsia dispar D.L.Jones et B.Gray, species

nova ab C. unguiculata Schltr. ovario

recurvissimo; floribus ferrugineis, labello

ovato ad basim cuneato et limbo multo

crassior et callo brevi atque calcariformi,

differt. Typus: Queensland. Cook District:

Longlands Gap, Herberton Range, 17°28'S,

145°38'E, 6 April 1999, B.Gray 7516 (holo:

QRS*; iso: BRI, CANB, MEL, MO, NSW).

Corsia sp. (Herberton Range B.Gray 3994)

(Bostock 2007).

Brownish pink terrestrial herb growing in

loose colonies. Roots filamentous, branched,

arising from nodes on the rhizome. Rhizome

horizontal, c. 4 mm thick, produced in annual

increments. Leaves reduced to sheathing

bracts. Scape erect, 8-18 cm long, 1.5-2 mm

thick, terete, brownish pink to pale mauve.

Bracts closely sheathing, 4-7, ovate to ovate-

lanceolate, 10-25 mm long, 4-6 mm wide,

pallid, acuminate. Pedicel 4-6 cm long. Ovary

arcuate to uncinate, 15-20 mm long, c. 1.5 mm

wide, brownish pink with reddish ribs. Flower

solitary, 2-2.6 cm long, 1.3-1.6 cm wide,

mostly brownish pink to light reddish brown.

Lateral sepals and petals recurved away from

the stamens and style, asymmetrical, ovate,

7-8 mm long, 2.5-3 mm wide, translucent

white, irregularly mottled with purplish-red,

the apex long-acuminate, darker, papillate,

contracting and becoming filiform with age.

Labellum, including limb, 20-30 mm long,

projecting below the perianth segments;

limb projecting downwards, c. 5 mm long

and 2 mm wide, whitish, winged; lamina at

right angles to the limb, ovate, 15-25 mm

long, 10-14 mm wide, light reddish brown to

brownish pink with a cream basal edge, with

16-24 darker longitudinal nerves, acuminate

to long-acuminate. Callus a small projecting

reddish-black spur at the junction of the

labellum limb and lamina. Stamens 5, and a

staminode on the medial side, held beneath

the lateral sepals and petals; filaments c. 1.5

mm long, white, connate basally, initially

erect then recurved; anthers oblong-elliptic,

c. 1.7 mm long, yellow, abscising with age;

*all specimens from QRS are now housed in CNS

staminode anterior to the style, similar in

size and shape to a filament, remaining erect.

Style columnar, c. 1.7 mm long; stigma three-

lobed, yellow. Capsule erect, linear-oblong,

20-35 mm long, 2-2.5 mm wide, brownish.

Seeds red-brown to blackish, with translucent

funicles. Fig. 1

Additional specimens examined: Queensland. Cook

District: Windsor Tableland, 23 km past Spencer

Creek, S.F. 144, 16°14'S, 145°00'E, Apr 2002, Booth

3081, Jensen & Cooper (BRI); W of Karnak via

Mossman, 16°23'S, 145°16'E, Jan 1995, Cooper &

Cooper WWC 869 (QRS); Mt Lewis, 16°35'S, 145°15'E,

Jan 1994, Cooper & Cooper WWC720 (QRS); S.F.R.

194, Western, Compartment 52, 17°17'S, 145°27'E,

Feb 1967, Dansie AFO4010 (BRI, QRS); S.F.R. 194,

Western, Compartment 53,17°18'S, 145°25'E, Feb 1976,

Hyland 8632 (QRS); Massey Creek, Ravenshoe - Millaa

Millaa road, 17°35'S, 145°35'E, Feb 1977, Gray 274

(QRS); Massey Creek, Ravenshoe - Millaa Millaa road,

17°36'S, 145°33'E, May 2002, Holmes 204 (BRI); Massey

Creek, Portion 297, Parish of Dirran, 17°37'S, 145°35'E,

Apr 1985, Gray 3994 (BRI, QRS); Massey Creek near

Ravenshoe, 17°37'S, 145°35'E, Mar 2004, Cooper &

Jensen WWC 1837 (CANB); e.lkmN of Chamillan

Creek crossing, Ravenshoe - Koombooloomba road,

17°42'S, 145°31'E, Mar 1968, Gray s.n. (CANB); road

to Koombooloomba Dam, 1 km N of Charmillan Creek,

17°42'S, 145°31'E, Mar 2004, Gray 8901 (QRS).

Distribution and ecology: Corsia dispar

is widespread on the Atherton Tableland,

Mt Lewis and Windsor Tableland areas in

north-eastern Queensland, but is seldom

collected. It grows on slopes close to streams

in tall rainforest on both basalt and granite

formations. Plants are found in small, loose

groups among leaf litter in gravelly loam.

Recorded altitude ranges from about 900 to

1100 m.

Notes: Corsia dispar would seem to be most

closely related to C. unguiculata as the two

species share an extended basal limb on

the labellum and a narrow callus. The new

species can however be readily distinguished

by its strongly recurved ovary (straight

in C. unguiculata ), light reddish brown

flowers (purplish-red in C. unguiculata),

ovate labellum with a cuneate base (broadly

ovate with a cordate or truncate base in

C. unguiculata), much thicker basal limb and,

with a short, erect, spur-like callus (narrow

upright crested ridge in C. unguiculata).

The roots of Corsia dispar definitely

branch although Van Royen (1972) records

720

Austrobaileya 7(4): 717-722 (2008)

Fig. 1. Corsia dispar. A. flowering plant. B. flower, side view and front view. C. top of flower showing tepals, stamens,

staminode, labellum limb, base of labellum lamina and callus. D. bud. E. capsule. A & B from Gray 7516 (QRS); C

from Gray 8901 (QRS); D & E from Cooper WWC1837 (QRS). Del. B. Gray.

721

Jones & Gray, Corsia dispar

roots in the New Guinea species as

unbranched. This appears to be an oversight

as several collections from New Guinea at

CANB also have branched roots. The flowers

of C. dispar are protandrous as previously

noted in other taxa (Smith 1907; Gibbs

1917; Van Royen 1972). Van Royen (1972)

also noted in his genus description that the

anthers are introrse, however the anthers are

extrorse at anthesis in C. dispar. They are also

noted in other publications as being extrorse

(Dahlgren et al. 1985). Similarly in the family

description Van Royen noted that the pedicels

usually elongated after flowering but, whereas

in C. dispar a thickening of the pedicel after

fertilisation has been observed, no increase in

length was measured on fruiting specimens.

Phenology : Plants flower mainly in January-

March (rarely as late as May) and fruit in

March-April (once in June). In seasons

of regular or continuous rainfall, new

inflorescences are produced from the rhizomes

and extend the flowering and fruiting period.

Etymology : The specific epithet is from the

Latin dispar , disparis , unlike, dissimilar,

different, in reference to the distinctive

characters of the Australian species.

New combination for a New Guinea taxon

The rank chosen for Corsia purpurata var.

wiakabui (Takeuchi & Pipoly 1998) seems

inappropriate in such a highly specialised

genus as Corsia. Any perceived relationship

by the authors between Corsia purpurata

and its variety must be considered tenuous,

as species of Corsia are generally considered

to be narrow endemics (Van Royen 1972). By

contrast Corsia purpurata , which occurs in

the vicinity of Lake Habbema in Irian Jaya,

and C. purpurata var. wiakabui , described

from specimens collected in New Ireland, are

separated by a geographical distance of more

than 2000 km and occur on different tectonic

plates. Morphologically Corsia purpurata

var. wiakabui can be distinguished from C.

purpurata by its much larger labellum (16-19

mm x 20-23 mm) which is tranversely ovate

to flabellate with a nearly truncate base. By

contrast the labellum of Corsia purpurata

is 10-13 mm x 9-10 mm, elliptic-rhomboid

in shape and with a cuneate base. From

these biogeographical and morphological

considerations it seems best to consider

these taxa specifically distinct and a new

combination is here made at that rank.

Corsia wiakabui (Takeuchi & Pipoly)

D.L. Jones & B.Gray, combinatio et status

nova

Basionym: Corsia purpurata L.O.Williams

var. wiakabui Takeuchi & Pipoly, Sida 18:

164 (1998). Type: Papua New Guinea. New

Ireland: Hans Meyer Range, pond next to

“Lake Camp”, 28 January 1994, W.Takeuchi

& J. Wiakabu (holo: LAE, n.v).

Acknowledgements

We thank the late Jim Willis and the late

Pieter Van Royen for their contributions very

early in this study, Wendy and Bill Cooper

and Rigel Jensen for reigniting our interest

following the bounteous wet season of 2004,

Laurie Adams for the Latin diagnosis and

discussions about the botanical code, Mark

Clements for discussions on the geography of

New Guinea, Peter Bostock for information on

the Corsia collections at BRI and Rebel Elick

for assistance at QRS. Karina Richards, Mark

Clements and Wendy Cooper commented on

the manuscript.

References

Bostock, P.D. (2007). Corsiaceae. In P.D. Bostock &

A.E. Holland (eds.). Census of the Queensland

Flora 2007 , p. 53. Environmental Protection

Agency: Brisbane.

Cribb, PJ. (1985). The saprophytic genus Corsia in the

Solomon Islands. Kew Magazine 2: 320-323.

Dahlgren, R.M.T., Clifford, H.T. & Yeo, P.F. (1985).

The Families of the Monocotyledons'. Structure,

Evolution and Taxonomy. Springer-Verlag:

Berlin, Heidelberg, New York, Tokyo.

Gibbs, L.S. (1917). Phytogeography and Flora of the

Arfak Mountains , p. 105. Taylor & Francis:

London.

Neyland, R. (2002). A phylogeny inferred from the

large-subunit (26S) ribosomal DNA sequences

suggests that Burmanniales are polyphyletic.

Australian Systematic Botany 15: 19-28.

Neyland, R. & Hennigan, M. (2003). A phylogenetic

analysis of large-subunit (26S) ribosome

DNA sequences suggests that Corsiaceae are

polyphyletic. New Zealand Journal of Botany

41: 1-11.

Austrobaileya 7(4): 717-722 (2008)

722

Rudall, RJ. & Eastman, A. (2002). The questionable

affinities of Corsia (Corsiaceae): evidence

from floral anatomy and pollen morphology.

Botanical Journal of the Linnaean Society 138:

315-324.

Smith, J.J. (1907). Corsiaceae. H.A.Lorentz (ed.),

Resultats de L’Expedition Scientifique

Neerlandaise a la Nouvelle-Guinee. Nova

Guinea 8(1): 197.

Takeuchi, W. & Pipoly, J.J. (1998). New flowering plants

from southern New Ireland, Papua New Guinea.

Sida 18: 161-168.

Van Royen, P. (1972). Sertulum Papuanum 17. Corsiaceae

of New Guinea and surrounding areas. Webbia

27: 223-255.

Capparis batianoffii Guymer (Capparaceae), a new

species from central coastal Queensland

G.P. Guymer

Summary

Guymer, G.P. (2008). Capparis batianoffii Guymer (Capparaceae), a new species from central coastal

Queensland. Austrobaileya 7(4): 723-725 (2008). The new species Capparis batianoffii (i Capparis

section Monostichiocalyx Radik.) is described and illustrated, together with information on its

distribution, habitat and conservation status.

Key Words: Capparis batianoffii , Capparaceae, Queensland flora, Australian flora, new species

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

An undescribed species of Capparis L.

belonging to C. section Monstichiocalyx

Radik, has been recognised to occur

on Gloucester Island, central coastal

Queensland, since it was first discovered by

George Batianoff in 1992. Further collections

of this species were made by Batianoff in

1994. It has been referred to as Capparis sp.

(Gloucester Island G.N. Batianoff 920912) at

the Queensland Herbarium (Jessup 2007).

Materials and methods

The paper is based on specimens held at

the Queensland Herbarium (BRI). The

terminology for Capparis follows that used

by Jacobs (1965). Measurements of the flowers

are based on dried or reconstituted material.

Taxonomy

Capparis batianoffii Guymer, species nova

affinis C. sarmentoso Cunn. ex Benth. sed ab

ea staminibus 8 (adversum 19-22) et foliis,

sepalis, petalis et gynophoris pubescentibus

(non glabellis vel glabrescentibus) differt.

Typus: Queensland. North Kennedy

District: Gloucester Island, East coast, 5 km

N of Chinaman’s Rock, 1 September 1992,

G.N.Batianoff 920912 (holo: BRI; iso: BRI,

K, MEL distribuendi).

Capparis sp. (Gloucester Island G.N. Batianoff

920912) (Jessup 2007)

Accepted for publication 24 September 2008

Climbing scrambling shrubs to 6 m, branches

pendulous. Stipular spines present, acicular,

2-3 mm long, recurved, cream or brown,

pubescent with upper half glabrous. Branchlets

pubescent with sandy or pale brown contorted

simple hairs. Leaves distichous; lamina ovate,

obovate, ovate-lanceolate or lanceolate,

10-20 x 3.5-7 mm; apices obtuse, rounded

or retuse, with a small mucro to 1 mm long;

bases cuneate; pubescent above (mid-dense)

and below with sandy or pale brown simple

contorted hairs (0.1-) 0.2-0.75 mm long;

margins slightly recurved; lateral veins 5-7

pairs; venation slightly raised above and below,

midrib sunken above, raised below; petioles

pubescent, 1-1.5 mm long. Inflorescence

axillary, solitary or a collateral pair; pedicels

9-10 mm long, with sandy or pale brown

simple hairs. Flowers white, aromatic, buds

globose. Sepals 4; outer two cymbiform,

5.6-7 x 4-4.5 mm, inner two ovate, 4.8-5.5

x 3.5-4 mm, densely pubescent outside with

sandy or pale brown simple hairs 0.3-0.7 mm

long, glabrous inside. Petals 4; obovate, 10-11

x 3.5-4 mm, pubescent outside with sandy

or pale brown simple hairs 0.3-0.7 mm long,

glabrous inside. Stamens 8; anthers 2.8-3.3

x 1.2-1.4 mm; filaments filiform, 18-20 mm

long, glabrous. Gynophore filiform, 17-18

x 0.2-0.3 mm diameter, glabrous above the

middle, pubescent below with sandy or pale

brown simple hairs 0.3-0.6 mm long. Ovary

ovoid, 2.8-3.3 x 1.2-1.3 mm, smooth, glabrous;

style 1—1.2 mm long; stigma globular, c. 0.5

mm diameter; placentas 4, ovules 16-26, in 2

rows. Berry not seen. Fig. 1.

Fig. 1. Capparis batianoffii. A. flowering branchlet x 2. B. leaves x 4. C. adaxial leaf surface x 12. D. flower x 3. F.

gynophore and ovary x 4. All from Batianoff920912 (BRI). Del. W. Smith.

Additional specimens examined : Queensland. North

Kennedy District [all Gloucester Island]: E coast, site

28, April 1994, Batianoff 940401Z & Figg (BRI); E

side bay, site 16, April 1994, Batianoff 940446 & Figg

(BRI).

Distribution and habitat : Capparis

batianoffii is known only from Gloucester

Island, off the central coast of Queensland

from three locations and six individual plants

(Batianoff etal. 1997). It occurs in Araucarian

vine thickets (Regional ecosystem 8.12.11) on

slopes amongst granitic boulders from near

sea-level to 400 m.

Notes : Capparis batianoffii appears to be

allied to C. sarmentosa but differs from this

725

Guymer, Capparis batianoffii

species by its fewer stamens (8 cf 19-22)

and its pubescent leaves, sepals, petals and

gynophores.

Capparis batianoffii will key to Capparis

quiniflora DC. in Flora of Australia (Hewson

1984) but it is readily distinguished from this

species by its much smaller leaf blades (10-20

mm cf 55-120 mm) and petioles (1-1.5 mm

cf 5-17 mm), and inflorescences (single or

collateral pair cf 2-10-flowered racemes).

Conservation status : Capparis batianoffii is

known to occur only on Gloucester Island.

The island of 2,960 hectares is part of the

Gloucester Islands National Park and lies

within the Great Barrier Reef World Heritage

Area. The species is known from three

locations on the Island with a known area of

occupancy of less than 20 km 2 . Batianoff et

al. (1997) reported six individual plants from

these three locations. The species is threatened

from chance stochastic events (e.g. drought,

fire) and its conservation status is assessed as

vulnerable based on criterion D (it has an area

of occupancy of less than 20km and fewer

than 1000 mature individuals) (IUCN 2001).

Etymology: Named for George Nicholas

Batianoff, Principal Botanist, Queensland

Herbarium, who discovered and made the

first collections of this species and who has

published extensively on the vegetation of

coastal Queensland.

Acknowledgements

I thank Wifi Smith (BRI) for the illustrations

and Peter Bostock for assistance with the

Latin diagnosis.

References

Batianoff, G.N., Dillewaard, H.A. & Franks, A.J.

(1997). Mackay/Whitsunday Coast Vegetation

and Floristics. Part 1: Description, Inventory

and Conservation. Queensland Herbarium,

Department of Environment: Indooroopilly.

Hewson, H.J. (1984). Capparaceae. In A.S. George

(ed.). Flora of Australia 8: 207-231. Australian

Biological Resources Study: Canberra.

IUCN (2001). IUCN Red List Categories and

Criteria. Version 3.1. IUCN Species Survival

Commission, IUCN: Gland, Switzerland/

Cambridge, United Kingdom.

Jacobs, M. (1965). The genus Capparis (Capparaceae)

from the Indus to the Pacific. Blumea 12: 385-

541.

Jessup, L.W. (2007). Capparaceae. In P.D. Bostock &

A.E. Holland (eds.). Census of the Queensland

Flora 2007, p. 42. Queensland Herbarium,

Environmentai Protection Agency: Brisbane.

Austrobaileya 7(4): 727-728 (2008) 727

SHORT COMMUNICATION

Reinstatement of Ammannia triflora Benth. (Lythraceae)

A.R. Bean

Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens, Mt Coot-tha

Road, Toowong, Queensland 4066, Australia. Email: tony.bean@epa.qld.gov.au

Bentham (1867) named Ammannia trijiora

based on a Robert Brown collection from

the islands of the Gulf of Carpentaria. The

name was in use for many years, until the

publication of the Lythraceae treatment in the

Flora of Australia series.

Therein, Hewson (1990) declared A. trijiora

Benth. to be an illegitimate name, with the

statement “ nom. illeg. non Wallich (1828)”.

Hewson renamed the species A. pubijiora

(Koehne) Hewson.

Wallich’s publication, referred to by

Hewson, is his “Numerical List of Dried

Specimens of plant, in the East India

Company’s Museum (1828-1849)”, often cited

as “Wallich’s Catalogue”. Ammannia triflora

Wall, which appeared in this publication, is

a nomen nudum , and hence not a legitimately

publishednew species. Nor can it be considered

a new combination based on Lythrum triflorum

L.f. or its homotypic synonym Nesaea trijiora

(L.f.) Kunth (published in 1825), as Wallich

did not give any indication that his name

might be based on either of these.

Therefore, Ammannia triflora Benth. is a

legitimate name, and is the oldest available

name for the species that Hewson called A.

pubiflora. Ironically, A. pubiflora (Koehne)

Hewson is an illegitimate name, a later

homonym of A. pubiflora (Koehne) Sosn.,

validly published in 1915.

Another matter in need of clarification is

the collection date of the type. Chapman et al.

(2001) listed the place and date of collection

of Ammannia pubiflora (=A. triflora Benth.)

as “Carpentaria island a Novr 27 1802”. This

was a transcription of the label of Brown’s

specimen at BM. Carpentaria island ‘a’ is

Sweers Island (Chapman et al. 2001).

Flinders arrived at Sweers Island on

the 16 th November 1802, and did not leave

until the 30 th November. Brown is known

to have collected on Sweers Island on

several occasions, with his final visit on 27 th

November (Vallance et al. 2001). However,

Hewson (1990) gave Robert Brown’s date of

collection as 27 th August 1802. This is clearly

an error, as on this date the expedition was

still on the east coast of Queensland, around

Shoalwater Bay.

Interestingly, the Queensland Herbarium

does not hold any subsequent specimens of

A. triflora from Sweers Island. This is despite

visits there by several botanists, including

J.F.Bailey, J.Shirley, D.Henne, L.Pedley and

M.Thomas. In fact, only Robert Brown has

collected the species from (what is now)

Queensland.

The revised synonymy for Ammania trijiora

is as follows:

Ammannia triflora Benth., FI. Austral. 3:

297 (1867); Nesaea lanceolata var. pubijiora

Koehne, Bot. Jahrb. Syst. 3: 326 (1882);

Ammannia pubiflora (Koehne) Hewson,

FI. Australia 18: 321 (1990), nom. illeg. non

(Koehne) Sosn. (1915). Type: [Queensland]

Sweers Island, 27 November 1802, R.Brown

(lecto: BM [here designated]; isolecto: BRI;

NSW).

[Ammannia triflora Wall., Numer. List n.

6323 (1832), nom. nud.\

Accepted for publication 2 June 2008

Austrobaileya 7(4): 727-728 (2008)

728

References

Bentham, G. (1867). Flora Australiensis. Volume 3.

L.Reeve & Co.: London.

Chapman, A.R., Moore, D.T., Rees, R.G. & Groves, E.W.

(2001). Robert Brown’s Australian Botanical

specimens, 1801-1805 at the BM. Available at:

http: //florabase. calm, wa. gov. au/bro wn

Hewson, H.J. (1990). Ammannia. In A S.George (ed),

Flora of Australia 18: 97-100, 321.

Vallance, T.G., Moore, D.T. & Groves, E.W. (2001).

Nature’s Investigator: The diary of Robert

Brown in Australia, 1801-1805. Australian

Biological Resources Study: Canberra.

Austrrobaileya 7(4): 729 (2008)

SHORT COMMUNICATION

729

A new combination in Cissocarpus Rozefelds

(Vitaceae), a fossil genus from Queensland

Peter D. Rostock 1 & H. Trevor Clifford 2

‘Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens, Mt Coot-tha

Road, Toowong, Queensland 4066, Australia. Email: peter.bostock@epa.qld.gov.au

2 Honorary Associate, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic

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

Recent molecular, phylogenetic analyses of

Cissus L. (Rossetto et al 2002, Rossetto et

al. 2007; Soejima & Wen 2006), together

with the transfer of Cissus opaca F.Muell.

to Clematicissus Planch. (Jackes & Rossetto

2006) prompted the present authors to re¬

examine names in this genus. This study

led to a misplaced combination in Cissus ,

namely Cissus jackesiae Rozefelds, for a

species described only from fossil seeds.

As this species was cited as the type of the

current form genus Cissocarpus Rozefelds (as

‘Ciccocarpus jackesiae) an orthographically

correctable error for Cissocarpus ), we felt

that Cissus jackesiae should be transferred to

that genus.

The reference to “aff. Cissocarpus jackesii

Rozefelds” in Carpenter et al. (2004: 706), is

construed here as a nomen nudum , since it

fails Art. 33.4 of the International Code of

Botanical Nomenclature (ICBN) (McNeill

et al. 2006) viz. ‘that a new combination is

not validly published unless its basionym

(name-bringing or epithet-bringing synonym)

... is clearly indicated and a full and direct

reference given to its author and place of valid

publication, with page or plate reference and

date’. It is also orthographically incorrect

in that the name Cissus jackesiae Rozefelds

commemorates Betsy R. Jackes and so under

Art. 60.11 should have the feminine ending

-iae as recognised by Rozefelds (1990).

Accepted for publication 25 August 2008

Cissocarpus jackesiae (Rozefelds) Bostock

& Clifford, combinatio nova

Basionym: Cissus jackesiae Rozefelds in

J.G. Douglas & D.C. Christophel (eds.),

Proceedings of the 3rd IOP Conference,

Melbourne, 1988 (1990: 124). A-Z Printers:

Melbourne.

References

Carpenter, R. J., Hill, R. S., Greenwood, D. R., Partridge,

A.D. & Banks, M.A. (2004). No snow in the

mountains: early Eocene plant fossils from

Hotham Heights, Victoria, Australia. Australian

Journal of Botany 52: 684-718.

Jackes, B.R. & Rossetto, M. (2006). A new combination

in Clematicissus Planch. (Vitaceae). Telopea 11:

390-391.

McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V,

Hawksworth, D.L., Marhold, K., Nicolson,

D.H., Prado, J., Silva, PC., Skog, J.E.,

Wiersema, J.H. & Turland, N.J. (eds.) (2006).

International Code of Botanical Nomenclature

(Vienna Code). (Regnum Vegetabile, 146).

A.R.G. Gantner Verlag KG: Vienna.

Rossetto, M., Jackes, B.R., Scott, K.D. & Henry,

R.J. (2002). Is the genus Cissus (Vitaceae)

monophyletic: evidence from plastid and

nuclear ribosomal DNA. Systematic Botany 27:

522-533.

Rossetto, M., Crayn, D.M., Jackes, B.R. & Porter, C.

(2007). Molecular evolution of the Australian

Vitaceae. Canadian Journal of Botany 85:

722-730.

Soejima, A. & Wen, J. (2006). Phylogenetic analysis

of the grape family (Vitaceae) based on three

chloroplast markers. American Journal of

Botany 93: 278-287.

Rozefelds, A.C. (1990). A mid Tertiary rainforest

flora from Capella, Central Queensland.

In J.G. Douglas & D C. Christophel (eds ),

Proceedings of the 3rd IOP Conference,

Melbourne, 1988, pp. 123-136. A-Z Printers:

Melbourne.

Austrobaileya 7(4): 731-733 (2008)

731

SHORT COMMUNICATION

Notes on Palmeria F.Muell. (Monimiaceae) in Australia and

the application of the name Palmeria racemosa (Tul.) A.DC.

G.P. Giiymer

Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens, Mt Coot-tha

Road, Toowong, Queensland 4066, Australia. Email: gordon.guymer@epa.qld.gov.au

Whiffin & Foreman (2007) revised the genus

Palmeria F.Muell. (Monimiaceae) forth q Flora

of Australia and recognised three species,

viz. P. foremanii Whiffin, P. hypotephra

(F.Muell.)Domin and P. scandens F.Muell.

They considered Palmeria scandens to include

P. coriacea C.T.White and P. racemosa (Tul.)

A.DC. However, the legitimate name for this

species when these species are considered to

be synonymous is P. racemosa as its basionym

pre-dates P. scandens by nine years.

Based on examination of herbarium

material and literature at the Queensland

Herbarium five species of Palmeria (viz.

P. coriacea , P. foremanii , P. hypotephra ,

P. racemosa and P. scandens) are recognised

for Australia. This is due to reinstatement of

P. coriacea from north-east Queensland and

recognition of P. racemosa for south-east

Queensland and eastern New South Wales

populations previously included erroneously

in P. scandens.

Key to the Australian Palmeria species

1 Leaves velutinous below and densely covered with white or cream

stellate hairs (0.2-0.5 (-0.7) mm diameter). Central coast and

NE Qld.3. P. hypotephra

1. Leaves glabrous or pubescent below, with pale brown or golden

stellate hairs (0.3-1.5 mm diameter) or stellate and simple

hairs, not covering the leaf surface.2

2 Leaves glabrous below or with an occasional stellate hair (0.1-

0.4 mm diameter) along the midrib. NE Qld (Thornton Peak,

Mt Spurgeon & Tinaroo Range above 900 m).1. P. coriacea

2. Leaves pubescent below with either predominantly stellate

hairs (0.3-1.3 mm diameter) or a mixture of simple and stellate

hairs.3

3 Leaves pubescent below with simple hairs (0.3-1.1 mm long) and

stellate hairs (0.3-1.3 mm diameter) (occasionally with only

simple hairs). Central coast to Cape York, Qld.5. P. scandens

3. Leaves pubescent below with stellate hairs (0.3-1.5 mm diameter) and an

occasional simple hair.4

4 Leaf pubescence below of stellate hairs (0.3-1.5 mm diameter) touching

to overlapping. SE Qld, NE NSW.2. P. foremanii

4. Leaf pubescence below of stellate hairs (0.3-1.1 mm diameter)

restricted to midrib, or midrib and lateral veins. SE Qld, E NSW

.4. P. racemosa

Accepted for publication 8 September 2008

732

Synopsis of Australian species

1. Palmeria coriacea C.T.White, Proc.

Roy. Soc. Queensland 47: 75 (1936). Type:

Queensland. Cook District: Thornton Peak,

14 March 1932, L.J. Brass 2282 (holo: BRI).

Additional selected specimens examined : Queensland.

Cook District: Mt Misery, NE of Mt Carbine, Nov

1988, Jessup GJM857, Guymer & McDonald (BRI);

Thornton Peak, Sep 1937, Tryons.n. (BRI [AQ415815]);

Mt Spurgeon, Sep 1936, White 11050 (BRI); Timber

Reserve 142, Zarda Logging Area, Jun 1973, Irvine

651 (BRI); Mt Lewis road, Aug 1995, Brown 95/294

f Wiecek & Radford (BRI); Tinaroo Range, Jun 1949,

Flecker NQNC12894 (BRI).

Distribution and habitat: The species is

endemic to north-east Queensland from Mt

Misery to Tinaroo Range in rainforest at

altitudes above 900 m.

2. Palmeria foremanii Whiffin, FI. Aust.

2: 454 (2007). Type: Queensland. Darling

Downs District: The Head, near source of

Teviot Creek, 4 May 1978, K.A. W.Williams

78049 (holo: BRI; iso: NSW, CANB n.v., K

n.v).

Palmeria scandens var. hirsuta Domin,

Biblioth. Bot. 89: 120 (1926). Type:

Queensland: Moreton District: “Regenwalder

der Tambourine Mts”, January 1910, K.Domin

s.n. (holo: PR, n.v.).

Additional selected specimens examined : Queensland.

Moreton District: Mt Glorious, Apr 1961, Cribb s.n.

(BRI [AQ170425]); Tamborine Mt, Jan 1924, Simmonds

s.n. (BRI [AQ63652]); Numinbah F.R., Springbrook

Plateau, Jul 2006, Halford Q9114 (BRI); Lyrebird Ridge

road, Springbrook, May 1984, Guymer 1892 (BRI); ditto

loc., Jan 2000, Forster PIF2527 & Leiper (BRI). Darling

Downs District: Near Moss Gardens, The Head to

Killarney road, Jan 2000, Forster PIF25273 (BRI). New

South Wales. North Coast; Acacia Plateau, Mar 1944,

White 12567 (BRI); N Yabbra road, Yabbra S.F., E of

Urbenville, May 1998, Bean 13233 (BRI, NSW); Track

to Minyon Falls, Apr 1984, Guymer 1881 (BRI); Whian

Whian S.F., Jun 1945, White 12832 (BRI).

Distribution and habitat : The species

is endemic to eastern Australia from Mt

Glorious, Queensland to Whian Whian State

Forest, New South Wales at altitudes between

100 and 1100 m. It occurs in rainforests on

soils derived mainly from basalt.

Austrobaileya 7(4): 731-733 (2008)

3. Palmeria hypotephra (F.Muell.) Domin,

Repert. Spec. Nov. Regni Veg. 12: 390 (1913);

Morinda hypotephra F.Muell., Vic. Nat. 6: 55

(1889). Type: Queensland. Cook District: Mt

Bellenden Ker, in 1887, W.A.Sayer s.n. (lecto:

MEL, n.v); fide Whiffin & Foreman (2007).

Additional selected specimens examined : Queensland.

Cook District: Top of Mt Hartley, Jul 1995, Forster

PIF17320 & Figg (BRI); Whyanbeel, Dec 1978, Gray

1185 (BRI); Mt Bellenden Ker summit, Dec 2001,

Forster PIF27935 & PIF27942 (BRI); The Boulders, W

of Babinda, Jul 1999, Forster PIF24665 (BRI); Westcott

road. Topaz, Nov 1989, Forster PIF25154, Booth &

Cooper (BRI); Tully Falls Weir, May 2003, Forster

PIF29392 (BRI). North Kennedy District: Millet Farm,

Ravenshoe, Apr 1940, Samundsett 3 (BRI); Alma Gap

S.F., 22 km NW of Cardwell, Oct 1988, Jessup GJM2404,

Guymer & McDonald (BRI); Paluma, Jun 1988, Jackes 1

(BRI). South Kennedy District: Dalrymple road to Mt

William, Eungella N.P, s.d., Pearson SP462 (BRI).

Distribution and habitat: The species is

endemic to north-east Queensland from

Mt Hartley to Eungella, west of Mackay. It

occurs in simple microphyll fern thicket, low

microphyll vineforest and complex notophyll

and mesophyll vineforest from near sea-level

to 1570 m.

4. Palmeria racemosa (Tul.) A.DC., Prodr.

16(2): 657 (1868); Hedycarya racemosa Tul.,

Ann. Sci., Nat. Bot., ser. 4, 3: 45 (1855). Type:

Nova Hollandia, in 1834, Hiigel s.n. (syn: P

n.v/, W n.v)\ Nova Hollandia, s.d., Baume s.n.

(syn: P n.v.; W n.v).

Additional selected specimens examined : Queensland.

Wide Bay District: Kin Kin, Mar 1916, Francis s.n.

(BRI [AQ63661]). Moreton District: Sunday Creek

road. Conondales, Jan 2002, Forster PIF28094 & Leiper

(BRI). New South Wales. Northern Tablelands: Slaty

Fire trail, Girard S.F., ENE of Tenterfield, Apr 2004,

Bean 21931 (BRI). North Coast: Dorngo S.F., Oct 1930,

White 7503 (BRI); Below Cameron’s Camp on Hasting

Forest Highway, Mar 1978, Hind 2234 (BRI); Upper

Williams River, near Salisbury, Mar 1938, White 11603

(BRI); Macquarie Pass, Jan 1962, Burgess 15243 (BRI).

Distribution and habitat: The species is

endemic to eastern Australia from Kin Kin,

south-east Queensland to Bateman’s Bay,

south coast of New South Wales. It occurs in

rainforests and on rainforest margins from

near sea level to 1000 m altitude.

733

Guymer, Notes on Palmeria

Notes : Hiigefs specimen of Palmeria

racemosa would have been collected between

April and October 1834 when he visited areas

from the Hunter River to Illawarra, New

South Wales (Clark 1994).

5. Palmeria scandens F.Muell., Fragm. 4:

152 (1864); Palmeria scandens var. scandens ,

Domin, Biblioth. Bot. 89: 120 (1926). Type:

Queensland. North Kennedy District:

Rockingham Bay, in 1863, J.Dallachy s.n.

(lecto: MEL 2050674 n.v.); fide Whiffin &

Foreman (2007).

Additional selected specimens examined : Queensland.

Cook District: Mt Finnegan summit area, Oct 1999,

Forster PIF25037 & Booth (BRI); Noah Creek, Dec

2001, Forster PIF27896, Young & Booth (BRI); S.F.

194, Mt Baldy, Dec 2001, Forster PIF27998 (BRI);

Jordan Logging Area, 16.5 km SE of Millaa Millaa, Oct

1988, Jessup GJM2070, Guymer & McDonald (BRI);

Yarrabah, Aug 1918, Michael s.n. (BRI [AQ63664]).

North Kennedy District: Bishop Peak, Hinchinbrook

Channel N.P., N of Ingham, May 1991, Bean 3244 (BRI);

Headwaters of Dryander Creek, Mt Dryander, Oct 1969,

Webb & Tracey 10034 (BRI). South Kennedy District:

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

PIF25509 & Booth (BRI).

Distribution and habitat : The species is

endemic to Queensland from the Mcllwraith

Range, Cape York to Eungella National Park,

west of Mackay, in rainforests or on rainforest

margins from near sea-level to 1200 m

altitude.

Discussion

Brophy et al. (2004) analysed the essential

oils from the leaves of Australian species

of Palmeria (Monimiaceae) excluding

P. coriacea. The results from their samples

using the classification above shows that

P.foremanii ( Forster PIF25277 ) has eight

unique compounds, P. hypotephra ( Forster

PIF27935 ) has four unique compounds,

P. racemosa (Forster PIF28094 ) has five

unique compounds and P. scandens ( Forster

PIF25509, PIF25037 & PIF27596 ) has no

unique compounds with the exception of

Forster PIF27998 from Mt Baldy which

has five unique compounds. This collection

differs from the other P. scandens in having

only simple hairs on the branchlets and leaves

and may represent a distinct taxon.

References

Brophy, J.J., Goldsack, R.J. & Forster, PI. (2004).

Essential oils from the leaves of the Australian

species of Palmeria (Monimiaceae). Journal of

Essential Oils Research 16: 312-317.

Clark, D. (1994, translated and edited). Baron von

Hugel’s New Holland journal: November 1833

- October 1834. Melbourne University Press

at the Miegunyah Press in association with the

State Library of New South Wales: Melbourne.

Whiffin, T. & Foreman, D.B. (2007). Palmeria,

Monimiaceae. In A.J.G. Wilson (ed.). Flora

of Australia 2: 88-91. Australian Biological

Resources Study/CSIRO Publishing: Canberra/

Melbourne.

Austrobaileya 7(4): 735-736 (2008)

735

SHORT COMMUNICATION

Two new species of Pandorea Spach (Bignoniaceae)

recognised from 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

The following new combinations in Pandorea

Spach are required to formally recognise two

Queensland species that are distinct from

Pandorea pandorana (J.Kenn. & Andrews)

Steenis, the Norfolk Island trumpet flower.

Bignonia pandorana J.Kenn. & Andrews

(the basionym of Pandorea pandorana) was

described from plants grown from seeds sent

by Colonel William Paterson, then stationed

at Norfolk Island, to James Lee and John

Kennedy, nurserymen at Hammersmith,

London who first raised it to flowering stage

in 1793. Flowering material was used by

John Kennedy in Andrews (1800) for the

written description and Andrews based his

illustration on plants “in the collection of J.

Vere, Esq. Kensington Gore”. Most databases

and publications cite the authorship as merely

Andrews’; however, the description of this

species was most likely to have been by John

Kennedy (Stafleu & Cowan 1976) who was

Andrew’s father in-law.

Green (1990, 1994) has cast doubt on a

Norfolk Island origin for this species and

believed a mistake was made in its provenance,

stating that it came from Sydney or from

cultivated material imported from New South

Wales. There is no evidence to suggest this

occurred as Colonel William Paterson was

stationed on Norfolk Island from November

1791 to March 1793 and collected seeds and

specimens that he sent to Banks (see Paterson

correspondence to Banks, State Library of

NSW/80239-47.jpg) and in 1794 discussed

with Banks the publication of his memoranda

on the natural history of Norfolk Island

(Paterson’s manuscript with illustrations by

Accepted for publication 8 September 2008

John Doody that Paterson sent to Banks is

now held in the State Library of New South

Wales, Sydney). The species was reported by

Kennedy in Andrews ( loc. cit) to be attacked

severely on Norfolk Island by a “white downy

insect of the genus Aphis” that covered the

whole plant and prevented vegetables being

grown on the Island. Interestingly, Paterson

in a letter to Banks dated 23 May 1793 (State

Library of NSW CY3008/244) refers to a box

he is sending with “some of the destructive

fly of Norfolk Island”. Banks has written in

the letter’s margin ‘a small cicada’, perhaps

referring to the Norfolk Island cicada (Kikihia

convicta (Distant)); however, this is thought

unlikely as the actual insect causing the

damage.

Andrews did not have a herbarium

(Stafleu & Cowan 1976) but based on his

illustration of Bignonia pandorana it has

5-11 lanceolate leaflets with serrate-dentate

margins, white flowers with red inside the

tube and with a red upper style and stigma.

The illustration appears to be more closely

allied to Green’s (1994) Pandorea pandorana

subsp. austrocaledonica (Bureau) PS.Green

from Lord Howe Island than to any mainland

Australian Pandorea. The first settlement

of Norfolk Island (1788 to 1814) resulted in

the clearing of 25% of the Island’s native

vegetation and may have caused the extinction

of Pandorea pandorana. Green (1990)

supports his argument that Pandorea did not

naturally occur on the island by the fact that

Bauer did not collect or illustrate this species

during his stay there from August 1804 to

February 1805 as it was not mentioned by

Endlicher (1833). If Kennedy’s claim that

the species was susceptible to insect attack

is indeed true, together with the fact that 619

hectares had been cleared by 1796, then this

736

species may well have disappeared before

Bauer’s arrival on Norfolk Island.

Pandorea floribunda (A.Cunn. ex DC.)

Guymer, combinatio nova

Basionym: Tecoma floribunda A.Cunn. ex

DC., Prodr. 9: 225 (1845). Type: “In Novae

Hollandiae rupestribus ad Moreton-bay”,

thickets and margins of creeks, Moreton Bay,

in 1828, A. Cunningham s.n. (holo: G-DC,

microfiche BRI!).

Pandorea sp. (K.A.Williams 86020) (Guymer

2007)

Additional selected specimens examined : Queensland.

Port Curtis District: S.F. 67, Sep 1985, Gibson 778

(BRI, NSW). Burnett District: Bunya Mountains, Oct

1917, Swain s.n. (BRI [AQ219029]). Wide Bay District:

Fraser Island, near Lake Wabby, Aug 1941, Blake 14385

(BRI). Moreton District: Dulong road, Dulong, Oct

1989, Sharpe 4898 (BRI); Yarraman, Aug 1944, Clemens

s.n. (BRI [AQ219049]); 5 km N of Dayboro, Mt Mee

road, Sep 1989, Henderson H3230 & Guymer (BRI);

Skyline Drive, Kholo, 10 km NW of Ipswich, Sep 1990,

Bird s.n. (BRI, BISH, MO, NSW); Near Ankida N.R.,

Springbrook, Sep 2005, Thompson MOR587 (BRI); Mt

Lindesay, base of mountain, Oct 1932, White 1552 (BRI).

Darling Downs District: Spicers Peak, Main Range

N.P, Sep 1995, Forster PIF17648 (BRI); 0.7 km W of

Moss Gardens, E of Killarney, Sep 2002, Bean 19370

(BRI). New South Wales. North Coast: “Moore Park”,

Old Grevillea, Sep 1972, Coveny 4567 & Rodd (BRI);

Lismore, Aug 1891, Baeuerlen 480 (BRI, NSW).

Distribution and habitat : The species occurs

from Gladstone, south-east Queensland to

Lismore, north-east New South Wales in

rainforest, on the margins of rainforests and

in adjoining eucalypt forests and woodlands,

from near sea-level to 1200 m.

Notes : This species is distinguished from

Pandorea pandorana by its pale yellow or

cream flowers and 3-5 broadly ovate to ovate,

entire leaflets (3-8 x 1.5-5 cm).

Pandorea linearis (F.M.Bailey) Guymer,

combinatio et status nova

Basionym: Tecoma australis var. linearis

F.M.Bailey, Queensl. FI. 4: 1134, pi. xlv

(1901); Pandorea australis subsp. linearis

(F.M.Bailey) Steenis, Rec. Trav. Bot. Neerl.

24: 863 (1927). Type: Herberton, s.d.,

J.F.Bailey s.n. (lecto[here designated]: BRI

[AQ218976]).

Austrobaileya 7(4): 735-736 (2008)

Additional selected specimens examined : Queensland.

Cook District: Stannary Hills, 15 km S of Mutchilba,

May 2006, Forster PIF31678 & McDonald (BRI,

DNA, NSW); 6 km E of Irvinebank, May 1974, Staples

260574/2 (BRI); On Silver Valley road, 3.4 km from Mt

Molloy, Apr 1988, Forster PIF3972 (BRI); 12 km along

Deadman’sroad, off Silver Valley road, Feb 1996, Forster

PIF18414 & Ryan (BRI). North Kennedy District:

Between St Pauls and Steward Head, E of Herberton, Jun

2005, McDonald KRM4275 (BRI); Ewan road, Mt Spec

area, 17.6 km from Paluma, Sep 1974, Williams 58 (BRI);

17.4 km W of Ivy Teahouse (Paluma), Aug 1994, Kemp

57OH & Kutt (BRI); 5.6 km SE of Hidden Valley, along

Paluma Road, Feb 2000, Pollock ABP817 & Edginton

(BRI); Between Bluewater Gorge and Pattersons Gorge,

Paluma Range, Cumming 20659 (BRI).

Distribution and habitat : The species occurs

from Stannary Hills to Paluma Range, north¬

east Queensland in eucalypt woodlands, open

forests and shrublands.

Notes : Pandorea linearis is distinguished

from P. pandorana by its 9-13 linear to

linear-lanceolate entire leaflets, the lateral

leaflets 1.1-4.2 x 0.15-0.6 cm.

The type collection of Tecoma australis var.

linearis comprises 3 sheets at BRI collected

by J.F.Bailey from Herberton. The BRI

collection AQ218976 is here selected lectotype

as it agrees with Bailey’s description and has

leaves and flowers.

References

Andrews, H.C. (1800). The Botanist’s Repository, for new

and rare plants. Vol. 2: Plate 86. H.C . Andrews:

London.

Endlicher, S. (1833). Prodromus Florae Norfolkicae

sive Catalogus stirpium quae in Insula Norfolk

annis a Ferdinando Bauer collecta depictae.

Fridericum Beck Universitatis Bibliopolam:

Vindobonae.

Green, PS. (1990). Notes relating to the floras ofNorfolk

and Lord Howe Islands III. Kew Bulletin 45:

235-245.

-(1994). Bignoniaceae. In A.J.G. Wilson (ed). Flora

of Australia 49: 343-344. Australian Biological

Resources Study: Canberra.

Guymer, G.P (2007). Bignoniaceae. In P.D. Bostock &

A .E. Holland, Census of the Queensland Flora

2007. Queensland Herbarium, Environmental

Protection Agency: Brisbane.

Stafleu, F.A. & Cowan, R.S. (1976, 2 nd edition).

Taxonomic literature. Volume 1: A-G. Bohn,

Scheltema & Holkema: Utrecht.

Austrobaileya 7(4): 737 (2008)

SHORT COMMUNICATION

737

New combinations in Liparis Rich, and Pterostylis R.Br. for two species of

Orchidaceae from Queensland

Peter D. Bostock

Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens, Mt Coot-tha

Road, Toowong, Queensland 4066, Australia. Email: peter.bostock@,epa.qld.gov.au

A number of segregate genera in Orchidaceae

have been recently published (e.g. Bunochilus

D.L.Jones & M.A.Clem.—segregated from

Pterostylis R.Br.) or reinstated (e.g. Diteilis

Raf.—segregated from Liparis Rich.) that are

not currently recognised by the Queensland

Herbarium. Two endemic Queensland species

have been named in Bunochilus (Jones

2006a) and Diteilis (Jones 2006b) that require

combinations in Pterostylis and Liparis

respectively.

Pterostylis vitrea (D.L.Jones) Bostock,

combinatio nova

Basionym: Bunochilus vitreus D.L.Jones,

Austral. Orchid Res. 5: 116 (2006).

Liparis petricola (D.L.Jones & B.Gray)

Bostock, combinatio nova

Basionym: Diteilis petricola D.L.Jones &

B.Gray, Austral. Orchid. Res. 5: 74 (2006).

References

Jones, D.L. (2006a). Towards a revision of Bunochilus

D.L.Jones & M.A.Clem. Australian Orchid

Research 5: 112-142.

Jones, D.L. (2006b). Miscellaneous new species of

Australian Orchidaceae. Australian Orchid

Research 5: 45-111.

Accepted for publication 30 September 2008

Referees consulted for Volume 7 of Austrobaileya

Acceptance of papers has depended on the outcome of review by referees. Those consulted

for the current volume are listed below. Several were consulted on more than one occasion.

Sincere thanks are extended to all these people whose expertise has helped to maintain journal

standards.

D.Albrecht

B.Lepschi

E. Lucas

M.J.Bayly

A.R.Bean

A.E.Orchard

P.D.Bostock

E. A. Brown

L.Pedley

P.Brownsey

J.J.Bruhl

S .Razafimdimison

R.W.Rogers

R.J.Chinnock

J.R.Clarkson

B.Schrire

B.Coppins

PS. Short

A.Slee

A.Davis

G.W. Staples

M.F.Duretto

I.R.Telford

A.S.George

K.Thiele

D.Goyder

G.P.Guymer

P.van Welzen

W.K.Harris

N.G.Walsh

A.E.Holland

J.Wege

PHoffmann

P.H.Weston

K.L.Wilson

J.Jeanes

Paul G.Wilson

L.W. Jessup

Peter G.Wilson

A ustrobaileya 7( 1 -4): 1-737

Index to Austrobaileya - volume 7

New names are in bold

Acacia 347, 381

Acacia argentea 354

Acacia argentina 348, 349

Acacia burrana 348, 350

Acacia calligera 381

Acacia conferta 347

Acacia cyclops 381

Acacia decurrens var. leichhardtii 155

Acacia eglandulosa 381

Acacia equisetifolia 153

Acacia glaucocarpa 160

Acacia hubbardiana 160

Acacia juncifolia subsp. serpentinicola 381

Acacia leichhardtii 155, 160

Acacia leptostachya 354

Acacia ligulata var. minor 381

Acacia lumholtzii 351

Acacia mariae 348

Acacia microcephala 352

Acacia microcybe 352

Acacia mirbelii 381

Acacia mollissima v ar. leichhardtii 155

Acacia oshanesii 155

Acacia polybotrya var. foliolosa 160

Acacia pubescens 155

Acacia rubricaulis 354, 355

Acacia salicina var. minor 381

Acacia serpentinicola 381

Acacia sp. (Burra Range R Jobson 467) 349

Acacia sp. (Gwambagwine F.Carter 2) 348

Acacia sp. (Harmer Creek J.R.Clarkson+ 9133)

354

Acacia sp. (Mt Leach Range D.G.Fell+

DF1704) 351

Acacia tindaleae 347, 348

Acacia undulifolia var. humilis 160

Acacia webbii 352, 353

Acronychia chooreechillum 562

Actephila 57, 58, 59, 60, 61, 62, 64, 66

Actephila bella 64, 65, 99

Actephila championiae 64, 66, 67, 68, 99

Actephila excelsa 57, 63

Actephila flavescens 64, 68, 69

Actephila foetida 60, 64, 70, 71, 99

Actephila grandifolia 64, 71, 72, 73, 99

Actephila latifolia 64, 74, 75, 99

Actephila lindleyi 59, 64, 75, 77, 78, 97

Actephila longipedicellata 64, 78, 79, 99

Actephila mearsii 92

Actephila mooreana 76

Actephila nitida 76

Actephila petiolaris 64, 80, 81

Actephila petiolaris subsp .jagonis 83

Actephila petiolaris subsp. petiolaris 81, 82

Actephilaplicata 64, 83, 84, 99

Actephila sessilifolia 64, 85, 86, 97

Actephila sp. (Claudie River BH 7803) 78

Actephila sp. (Granite Creek P. I. Forster

PIF5859) 65

Actephila sp. (Koumala I.G.Champion 870)

Actephila sp. (Little Mossman BH13697) 74

Actephila sp. (Lockerbie WWC 817) 89

Actephila sp. (Millaa Millaa RJ 494) 91

Actephila sp. (Possum Scrub P.I.Forster

PIF14372) 89

Actephila sp. (Rocky River G.C.Stocker 1042)

Actephila sp. (Wooroonooran N.P, P.I.Forster

PIF17151) 91

Actephila traceyi 64, 87, 88, 99

Actephila venusta 64, 89, 90

Actephila vernicosa 64, 91, 92

Adiantaceae 341

Aidia racemosa 273

Ailanthus punctata 162

Akania bidwillii 158

Albrecht, D.E. with Bean, A.R. (2008).

Solanum succosum A.R.Bean & Albr.

(Solanaceae), a new species allied to S.

chippendalei Symon 669

Alectryon repandodentatus 229

Alphitonia 377

Alphitonia obtusifolia 377

Alphitonia pomaderroides 377

Alsophila leichhardtiana 155, 158

Alsophila rebeccae var. commutata 561

Amanoa leichhardti 155

Ammannia triflora 725

Anomospermum 63

Anomospermum excelsum 63

Anthocercis leichhardtii 155, 162

Apluda mutica 229

Apocynaceae 145

Ardisia 99, 101

Ardisia pseudojambosa 103

Ardisia repandula 105

Aristida gracilipes 161

Aristida leichhardtiana 155, 161

Aristida vagans var. gracillima 161

Arthrochilus lavarackiana 385

Arthropteris palisotii 561

Arytera foveolata 155, 162

Aseroe actinobola 161

Aseroe rubra 154, 161

Asperula geminifolia 162

Austrobaileya 7(1 -4): 1-737

Aspidium acuminatum var. villosum 561

Aspidium ramosum var. lineare 561

Asteraceae 567

Atriplex stipitata 158

Atwood, J.P. with Snow, N. & McFadden, J.

(2008). Three new species of Rhodomyrtus

(DC.) Rchb. (Myrtaceae) from Papua New

Guinea 689

Backhousia angustifolia 126

Backhousia bancroftii 126

Backhousia citriodora 121,125

Backhousia enata 121, 123, 126, 127

Backhousia hughesii 125

Backhousia kingii 126

Backhousia myrtifolia 126

Backhousia oligantha 125, 126

Backhousia sciadophora 126

Baculariapalmeriana 561

Bailey, F.M. 555, 559, 677

Bambusa moreheadiana 561

Bauhinia leichhardtii 152, 155

Bazzania 129

Bazzania involuta 129

Bazzania sauropoda 129, 130, 131, 132

Bean, A.R. (2005). Eucalyptus erosa

(Myrtaceae), a new stringybark species

from central Queensland 141

Bean, A.R. (2005). Notes on the narrow¬

leaved Ironbarks (Myrtaceae: Eucalyptus

subseries Subglaucae) 111

Bean, A.R. (2006). A new combination in

Alphitonia Endl. (Rhamnaceae) 377

Bean, A.R. (2006). Eucalyptus megasepala

A.R.Bean (Myrtaceae), a new species from

Queensland allied to E. tetrodontaV. Muell.

(Myrtaceae) 305

Bean, A.R. (2007). A taxonomic revision of

Callitriche L. (Callitrichaceae) in Australia

545

Bean, A.R. (2008). A taxonomic revision of

Erythrina L. (Fabaceae: Faboideae ) in

Australia 641

Bean, A.R. (2008). Reinstatement ofAmmannia

triflora Benth. (Lythraceae) 725

Bean, A.R. & Albrecht, D.E. (2008). Solanum

succosum A.R.Bean & Albr. (Solanaceae),

a new species allied to S. chippendalei

Symon 669

Bellenden Ker 555, 677

Bertya 216, 217

Bertya gummifera var. psiloclada 424

Bertya mitchellii 442

Bertya mitchellii var. mitchellii 442

Bertya psiloclada 424

Bertya virgata 632

Beyeria 215, 216, 217, 577 578

Beyeria 581

Beyeria apiculata 597, 599

Beyeria backhousei 586

Beyeria bickertonensis 223, 632

Beyeria brevifolia 598

Beyeria brevifolia var. brevifolia 598

Beyeria brevifolia var. brevipes 629

Beyeria brevifolia var. robustior 627

Beyeria brevifolia var. truncata 631

Beyeria calycina 600

Beyeria calycina var. calycina 600

Beyeria calycina var. minor 617

Beyeria cinerea 601

Beyeria cinerea subsp. borealis 603

Beyeria cinerea subsp. cinerea 603

Beyeria cockertonii 604, 605

Beyeria constellata 606, 607

Beyeria cyanescens 603

Beyeria cygnorum 603

Beyeria disciformis 608, 609

Beyeria drummondii 586

Beyeria gardneri 610

Beyeria lanceolata 582, 584

Beyeria lapidicola 612, 613

Beyeria lasiocarpa 583

Beyeria lasiocarpa f. denudata 632

Beyeria lasiocarpa f. lasiocarpa 583

Beyeria latifolia 614

Beyeria lechenaultii 585

Beyeria lechenaultii f. elaeagnoides 585

Beyeria lechenaultii f. genuina 586

Beyeria lechenaultii f. lechenaultii 585

Beyeria lechenaultii f. myrtoides 585

Beyeria lechenaultii f. pernettioides 585

Beyeria lechenaultii f. rosmarinoides 585

Beyeria lechenaultii f. salsoloides 586

Beyeria lechenaultii f. vaccinioides 586

Beyeria lechenaultii var. backhousei 586

Beyeria lechenaultii var. drummondii 586

Beyeria lechenaultii var. genuina 586

Beyeria lechenaultii var. latifolia 586

Beyeria lechenaultii var. lechenaultii 585

Beyeria lechenaultii var. ledifolia 586

Beyeria lechenaultii var. rosmarinoides 585

Beyeria ledifolia 586

Beyeria ledifolia var. angustifolia 586

Beyeria ledifolia var. backhousei 586

Beyeria ledifolia var. genuina 586

Beyeria ledifolia var. ledifolia 586

Beyeria lepidopetala 615

Beyeria loranthoides 632

Beyeria minor 617

Beyeria oblongifolia 593

Beyeria opaca 590

Beyeria opaca var. latifolia 590

Beyeria opaca var. linearis 586

Beyeria opaca var. longifolia 632

Auslrobai/eya 7(1-4): 1-737

Beyeria opaca var. opaca 590

Beyeria opaca var. typica 590

Beyeriaphysaphylla 618, 619

Beyeria preissii 593

Beyeria rostellata 620, 622

Beyeria sect. Beyeria 581, 582

Beyeria sect. Beyeriopsis 596

Beyeria sect. Discobeyeria 581

Beyeria sect. Eubeyeria 581

Beyeria sect. Oxygyne 215

Beyeria similis 623

Beyeria simplex 624, 625

Beyeria sp. (Bull Creek Gorge B.O’Keeffe

573) 227

Beyeria sp. A Ravensthorpe (A.S. George

9474) 631

Beyeria sp. B Mount Ragged (A.S. George

7422) 624

Beyeria sp. Bandalup Hill (G. Cockerton 7553)

604

Beyeria sp. C South West (A.S. George 9878)

629

Beyeria sp. Jackson Range (R. Cranfield & R

Spencer 7751) 620

Beyeria sp. Murchison (B. Jeanes s.n. 7/7/2005)

612

Beyeria sp. Scaddan (P. van der Moezel

PGV161) 618

Beyeria subtecta 591

Beyeria sulcata 626

Beyeria sulcata var. brevipes 629

Beyeria sulcata var. gracilis 629

Beyeria sulcata var. sulcata 626

Beyeria sulcata var. truncata 631

Beyeria tristigma 215, 216, 219, 632

Beyeria uncinata 632

Beyeria villosa 631

Beyeria virgata 632

Beyeria viscosa 592

Beyeria viscosa var. amoena 592

Beyeria viscosa var. angustifolia 586

Beyeria viscosa var. genuina 593

Beyeria viscosa var. latifolia 592

Beyeria viscosa var. minor 592

Beyeria viscosa var. oblongifolia 593

Beyeria viscosa var. obovata 593

Beyeria viscosa var. viscosa 592

Beyeriopsis 581, 596

Beyeriopsis brevifolia 598

Beyeriopsis cinerea 601

Beyeriopsis cyanescens 603

Beyeriopsis cygnorum 603

Beyeriopsis latifolia 614

Beyeriopsis lepidopetala 615

Beyeriopsis similis 623

Bignoniaceae 735

Blechnum whelanii 561

Booth, R. & Moore, D. (2006). Reduction of

Carex rhytidocarpa Nelmes (Cyperaceae)

to a synonym of C. inversa R.Br. 373

Boronia 665

Boronia inflexa 172

Boronia inflexa subsp. grandiflora 171, 172

Boronia inflexa subsp. inflexa 172

Boronia inflexa subsp. montiazura 172

Boronia inflexa subsp. torringtonensis 172

Boronia minutiflora 520

Boronia sect. Cyanothamnus 171

Boronia sect. Valvatae 665, 666

Boronia ser. Valvatae 667

Boronia subsect. Anomalae 666

Boronia subser. Lanceolatae 667

Boronia subser. Rosmarinifoliae 668

Boronia subser. Valvatae 667

Bossiaea brownii 159

Bostock, P.D. (2008). New combinations in

Liparis Rich, and Pterostylis R.Br. for two

species of Orchidaceae from Queensland

737

Bostock, P.D. & Clifford, H.T. (2008). A new

combination in Cissocarpus Rozefelds

(Vitaceae), a fossil genus from Queensland

729

Bostock, P.D. with Field, A.R. (2008). Huperzia

tetrastichoides A.R.Field & Bostock

(Lycopodiaceae), a newly recognised

species of tassel fern from the Wet Tropics

of Queensland, Australia 711

Bostock, P.D. with Forster, PI. & Kokubugata,

G. (2006). Chromosome records for four

species of Pellaea sect. Platyloma (J.Sm.)

Hook. & Baker (Adiantaceae) from

Australia 341

Bridelia leichhardtii 155, 159

Brophy, J.J. with Ford, A.J. & Craven, L.A.

(2005). Backhousia enata (Myrtaceae), a

new species from north-eastern Queensland

121

Broughton, A.D. with Dowe, J.L. (2007).

F.M.Bailey’s ascent of Mt Bellenden-Ker in

1889, and notes on the publication priority

of new vascular plant species from the

Expedition 555

Broughton, A.D. with Dowe, J.L. (2008).

F.M.Bailey’s taxonomy of A.Meston’s

collections from the Bellenden Ker

Expedition of 1904 677

Bryophyllum x houghtonii 383

Bryophyllum daigremontianum 383

Bryophyllum delagoense 383

Bulbophyllum lageniforme 678

Bulbophyllum toressae 561

Bunochilus vitreus 737

Austrobaileya 7(1 -4): 1-737

Butler, D.W. with Holland, A.E. (2007).

Trioncinia patens A.E.Holland &

D.W.Butler (Asteraceae: Coreopsideae :

Chrysanthellinae), a new and endangered

species from central Queensland 567

Callitrichaceae 545

Callitriche 545

Callitriche antarctica 552

Callitriche brachycarpa 547

Callitriche brutia 549

Callitriche brutia var. brutia 550

Callitriche capricorni 548

Callitriche cyclocarpa 550

Callitriche deflexa 549

Callitriche hamulata 549

Callitriche macropteryx 547

Callitriche marginata 549

Callitriche muelleri 547

Callitriche palustris 551

Callitriche palustris var. palustris 551

Callitriche platycarpa 552

Callitriche sonderi 548

Callitriche stagnalis 551

Callitriche umbonata 550

Callitriche verna 551

Calyptrostigma 581

Calyptrostsigma ledifolium 586

Calyptrostigma oblongifolium 593

Calyptrostigma viscosum 593

Camptochaete leichhardtii 155

Capparaceae 723

Capparis batianoffii 723, 724

Capparis sp. (Gloucester Island G.N. Batianoff

920912) 723

Carex inversa 373, 375

Carex inversa f. major 374

Carex inversa f. parvula 374

Carex inversa var. leichhardtii 374

Carex inversa var. major 374

Carex inversa var. minor 374

Carex lophocarpa 374, 375

Carex rhytidocarpa 373, 374

Cassia concinna 158

Cassinia theodori 158

Casuarina cunninghamiana subsp.

cunninghamiana 158

Casuarina equisetifolia var. microcarpa 158

Casuarina villosa 153

Chamaecrista concinna 158

Chomelia 31

Chomelia dallachiana 34

Choretrum can dollei 162

Chorizema leichhardtii 155,159

Cissocarpus 729

Cissocarpus jackesiae 729

Citrus inodora 561

Cladium scleroides 159

Claoxylon 451

Claoxylon angustifolium 455, 456

Claoxylon australe 159, 457, 459

Claoxylon australe var. australe 457

Claoxylon australe var. dentata 457

Claoxylon australe var. latifolia 464

Claoxylon australe var. laxijlora 457

Claoxylon delicatum 460

Claoxylon hillii 460, 462

Claoxylon indicum 454, 455

Claoxylon indicum var. novoguineense 460

Claoxylon tenerifolium 453, 454, 463, 464

Claoxylon tenerifolium subsp. boreale 466,

467

Claoxylon tenerifolium subsp. tenerifolium

464, 465

Clarke, C. & Kruger, R. (2006). Nepenthes

tenax C.Clarke & R.Kruger (Nepenthaceae),

a new species from Cape York Peninsula,

Queensland 319

Clematis aristata var. leichhardtiana 155,

162

Clifford, H.T. & Dettmann, M E. (2005). Wing

structure in seeds of Strangea Meisn. and

Stenocarpus R.Br. (Proteaceae) 165

Clifford, H.T. with Bostock, P.D. (2008). A

new combination in Cissocarpus Rozefelds

(Vitaceae), a fossil genus from Queensland

729

Collabium pumilum 202

Commersonia 231, 232, 365

Commersonia argentea 235

Commersonia bartramia 239, 371

Commersonia beeronensis 236

Commersonia echinata 239

Commersoniafraseri 238

Commersonia fraseri var. fraseri 238

Commersonia fraseri var. macrophylla 238

Commersonia fraseri var. microphylla 238

Commersonia inglewoodensis 369, 370, 372

Commersonia johnsonii 243, 371

Commersonia leichhardtii 155, 163,247,371

Commersonia leiperi 246, 247, 372

Commersonia macrostipulata 367, 371

Commersonia novoguinensis 241

Commersonia obliqua 239

Commersonia pearnii 244, 245, 371

Commersonia pedleyi 248, 249, 372

Commersonia perkinsiana 365, 366, 372

Commersonia procumbens 371

Commersonia reticulata 241, 242, 371

Commersonia rossii 238

Commersonia sp. (Beeron P.I.Forster PIF4658)

236, 237

Commersonia sp. (Cadarga G.P.Guymer 1642)

235

Auslrobai/eya 7(1-4): 1-737

Commersonia sp. (Kuranda K.Williams 211)

367

Commersonia sp. (Mt Tinbeerwah G.P.

Guymer 1786) 233

Commersonia sp. (Zamia Range R.W. Johnson

1398) 243

Commersonia sp. 1 233, 238

Commersonia sp. 2 235

Commersonia subg. Commersonia 371

Commersonia subg. Verticillata 233

Commersonia viscidula 233, 234

Convolvulaceae 311

Cordia ixiocarpa 158

Cordia myxa var. ixiophylla 158

Corsia dispar 111, 719, 720

Corsiapurpurata var. wiakabui 721

Corsia sect. Corsia 718

Corsia sect. Unguiculatae 718

Corsia sp. (Herberton Range B.Gray 3994)

719

Corsia wiakabui 721

Corsiaceae 717

Corymbia leichhardtii 155

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 205

Crassulaceae 383

Craven, L.A. with Brophy, J.J. & Ford, A.J.

(2005). Backhousia enata (Myrtaceae), a

new species from north-eastern Queensland

121

Crepidomanes barnardianum 563

Croton stigmatosus 159

Croton viscosus 593

Cryptandra D019989 Jabiru 300

Cryptandra debilis 302

Cryptandrafiliformis 302

Cryptandra gemmata 302

Cryptandra intratropica 302

Cryptandra monticola 302

Cryptandra pogonoloba subsp. pogonoloba

302

Cryptandra pogonoloba subsp. septentrionalis

302

Cryptandra sp. ‘Jabiru East’, Kellermann et al.

(2005) 300

Cryptandra sp. 5 (Jabiru ; L.A.Craven 6484)

300

Cryptandra sp. Jabiru (L.A.Craven 6484)

300

Cryptandra triplex 299, 300, 301, 302, 303

Cupania lucens 158

Cupaniopsis anacardioides 297

Cupaniopsis baileyana 297

Cupaniopsis cooperorum 296

Cupaniopsis dallachyi 293, 296, 297, 298

Cupaniopsis diploglottoides 297

Cupaniopsisflagelliformis 297

Cupaniopsisfleckeri 297

Cupaniopsis foveolata 297

Cupaniopsis newmanii 297

Cupaniopsispapillosa 293, 295, 296, 297

Cupaniopsis parvifolia 297

Cupaniopsis serrata 297

Cupaniopsis shirleyana 296

Cupaniopsis simulatus 297

Cupaniopsis sp. (Tully Falls) 294

Cupaniopsis tomentella 297

Cupaniopsis wadsworthii 296

Cyanocarpus 561

Cyanocarpus nortoniana 561

Cyathea baileyana 561

Cyathealeichhardtiana 155,158

Cycas scratchleyana 229

Cyrtandra baileyi 561

Datura leichhardtii 155

Dendrobium 183

Dendrobium acutisepalum 185

Dendrobium eymanum 184, 185, 186

Dendrobium flebiliflorum 187, 188

Dendrobium fusciflorum 183

Dendrobium macrophyllum var. stenopterum

189

Dendrobium montis-yulei 187

Dendrobium nothofagicola 192

Dendrobium obtusa 192

Dendrobium obtusum 192

Dendrobium paragnomus 183, 192, 194

Dendrobium sect. Herpetophytum 183

Dendrobium sect. Latouria 184

Dendrobium sect. Pedilonum 192

Dendrobium sp. aff. D. gnomus 192

Dendrobium spenceanum 187, 190

Dendrobium spinuliferum 189, 191

Dendrobium stenopterum 189, 192

Dendrobium stipiticola 184, 185

Dendrobium terrestre 187

Dendrobium toressae 561

Dendrocnide excelsa 163

Dendrocnide photinophylla 156, 163

Dendrocolla obtusa 192

Dendrohypnum leichardtii 155

Denhamia viridissima 561

Derris koolgibberah 561

Dettmann, M.E. with Clifford, H.T. (2005).

Wing structure in seeds of Strangea Meisn.

and Stenocarpus R.Br. (Proteaceae) 165

Dicranoloma leichhardtii 155

Dicranum leichhardtii 155

Digitaria macractinia subsp. leichhardtiana

155

Dimeria glabriuscula 561

Austrobaileya 7(1 -4): 1-737

Dimeria ornithopoda 561

Dimocarpus longan subsp. longan 155, 162

Diploptera 167, 168

Diplospora australis 43, 44, 45

Diplospora cameronii 46

Diplospora ixoroides 47, 51, 52

Disaster 232

Disaster novoguinensis 232, 241

Disco calyx 99, 101

Discocarpus 57

Diteilis petricola 737

Dowe, J.L. (2005). Ludwig Leichhardt’s

Australian plant collections, 1842-1847

151

Dowe, J.L. & Broughton, A.D. (2007).

F.M.Bailey’s ascent of Mt Bellenden-Ker in

1889, and notes on the publication priority

of new vascular plant species from the

Expedition 555

Dowe, J.L. & Broughton, A.D. (2008).

F.M.Bailey’s taxonomy of A.Meston’s

collections from the Bellenden Ker

Expedition of 1904 677

Drosera 205

Drypetes deplanchei 159

Duboisia leichhardtii 155, 162

Duretto, M.F., Grimshaw, P. & Sparshott, K.

(2005). The rediscovery of Boronia inflexa

subsp. grandifiora (Rutaceae) 171

Duretto, M.F. & Forster, PI. (2007). A

taxonomic revision of the genus Zieria Sm.

(Rutaceae) in Queensland 473

Duretto, M.F. (2008). A new subsection and

two new subseries for Boronia Sm. sect.

Valvatae (Benth.) Engl. (Rutaceae) 665

Duretto, M.F. & Forster, PL (2008). New

subspecies for Zieria odorifera J. A. Armstr.

(Rutaceae) from northern New South Wales

681

Elatostema reticulatum 163

Embeliafliickigeri 99, 107

Emmenosperma alphitonioides 162

Eremophila sturtii 632

Eriostemon glasshousiensis 178

Eriostemon leichhardtii 155, 162

Eriostemon myoporoides subsp. conduplicatus

176

Eriostemon myoporoides subsp. epilosus 177

Eriostemon myoporoides subsp. leichhardtii

178

Eriostemon myoporoides subsp.

queenslandicus 180

Eriostemon myoporoides var. leichhardtii

155, 178

Eriostemon queenslandicus 180

Eriostemon trachyphyllus var. leichhardtii

155, 162, 178

Erythrina 641, 642

Erythrina x sykesii 654

Erythrina acanthocarpa 656

Erythrina caffra 643, 653

Erythrina crista-galli 642, 655

Erythrina fusca 642, 656

Erythrina insularis 643, 646

Erythrina lysistemon 657

Erythrina numerosa 643,651,652

Erythrina sp. (Croftby PI.Forster+ PIF6209)

651

Erythrina speciosa 643

Erythrina variegata 643

Erythrina vespertilio 648

Erythrina vespertilio subsp. biloba 643, 650

Erythrina vespertilio subsp. vespertilio 643,

648, 649

Erythrina vespertilio var. typica 648

Erythrina vespertilio var. vespertilio 648

Erythrophleum chlorostachys 152

Eucalyptus albens 160

Eucalyptus atrata 119

Eucalyptus beaniana 118

Eucalyptus bowmanii 114

Eucalyptus cloeziana 227

Eucalyptus crebra 112, 116, 119

Eucalyptus cullenii 119

Eucalyptus dec or tic ans 118

Eucalyptus drepanophylla 116,119

Eucalyptus elegans 113, 114, 115, 117, 119

Eucalyptus erosa 141,142, 143

Eucalyptus exilipes 119

Eucalyptus farinosa 118

Eucalyptus fibrosa subsp. fibrosa 118, 160

Eucalyptus fibrosa subsp. nubila 118

Eucalyptus fracta 118

Eucalyptus granitica 119

Eucalyptus jensenii 118

Eucalyptus leichhardtii 155

Eucalyptus mackieana 141, 142

Eucalyptus megasepala 305, 307, 308, 309,

310

Eucalyptus ophitica 119

Eucalyptus paedoglauca 118

Eucalyptus phoenicea 222

Eucalyptus prava 173

Eucalyptus quadricostata 118

Eucalyptus rhombica 118

Eucalyptus siderophloia 119

Eucalyptus sp. (Carnarvon Range) 141

Eucalyptus sp. (Chinchilla L. Pedley 4022)

113

Eucalyptus sp. (Gilgandra D.J. Carr+ 352)

113

Eucalyptus sp. (Inglewood P. Grimshaw+

PG846) 113

Auslrobai/eya 7(1-4): 1-737

Eucalyptus sp. (Mt Moffatt HS D.F. Blaxell+

1883) 141

Eucalyptus staigeriana 118

Eucalyptus subser. Subglaucae 111, 113, 118

Eucalyptus taurina 118

Eucalyptus tetrodonta 305, 306, 309, 310

Eucalyptus tholiformis 116, 117, 118

Eucalyptus whitei 112,119

Eucalyptus xanthoclada 119

Euphorbiaceae 215, 387, 451, 573, 577

Euphoria leichhardtii 155, 162

Euphoria leichhardtii var. hebepetala 155,

162

Exocarya scleroides 159,563

Fell, D.G., The distribution, habitat and

conservation status of Suregada glomerulata

(Blume) Baill. (Euphorbiaceae) from north¬

eastern Cape York Peninsula, Queensland

573

Festuca latispicea 161

Festuca loliiformis 161

Ficus crassipes 562

Ficus platypoda var. leichhardtii 156

Field, A.R. & Bostock, P.D. (2008). Huperzia

tetrastichoides A.R.Field & Bostock

(Lycopodiaceae), a newly recognised

species of tassel fern from the Wet Tropics

of Queensland, Australia 711

Flemingia parviflora 159

Ford, A.J. & Halford, DA. (2006).

Gynochthodes oresbia Halford & A.J.Ford

(Rubiaceae), a new and cryptic species

from north-east Queensland 357

Ford, A.J., Craven, L.A. & Brophy, J.J. (2005).

Backhousia enata (Myrtaceae), a new

species from north-eastern Queensland

121

Forster, PI. (2005). A taxonomic revision

of Actephila Blume (Euphorbiaceae/

Phyllanthaceae) in Australia 57

Forster, PI. (2005). Cycas scratchleyana

F.Muell. (Cycadaceae), a new species record

for Queensland and Australia 229

Forster, P.I. (2005). New species of Philotheca

Rudge (Rutaceae) from Queensland 175

Forster, P.I. & Takeuchi, W. (2005). Rediscovery

and neotypification of Marsdenia

arachnoidea Schltr. (Apocynaceae:

Asclepiadoideae - Marsdenieae), an

endangered species from Papua New

Guinea 145

Forster, P.I. (2006). Bryophyllum x houghtonii

(D.B.Ward) P.I.Forst., a new combination

in Crassulaceae for the hybrid Mother of

Millions 383

Forster, P.I. (2006). Cupaniopsis papillosa

P.I.Forst. (Sapindaceae), a new species from

the ‘Wet Tropics’ of north-east Queensland

293

Forster, P.I. (2006). Jagera madida P.I.Forst.

(Sapindaceae), a new name and change

of rank for J. javanica subsp. australiana

Leenh. 379

Forster, P.I. (2006). Mischarytera megaphylla

P.I.Forst. (Sapindaceae), a new species from

the ‘Wet Tropics’ of north-east Queensland

279

Forster, P.I. (2006). Synima reynoldsiae

P.I.Forst. (Sapindaceae), a new species from

the ‘Wet Tropics’ of north-east Queensland.

285

Forster, P.I. (2007). A taxonomic revision

of Claoxylon A.Juss. (Euphorbiaceae) in

Australia 451

Forster, P.I. (2008). Plectranthus batianoffii

P.I.Forst. (Lamiaceae), a new species from

north-east Queensland 707

Forster, P.I. with Duretto, M.F. (2007). A

taxonomic revision of the genus Zieria Sm.

(Rutaceae) in Queensland 473

Forster, P.I. with Duretto, M.F. (2008). New

subspecies for Zieria odorifera J.A.Armstr.

(Rutaceae) from northern New South Wales

681

Forster, P.I. with Kokubugata, G. and Bostock,

P.D. (2006). Chromosome records for

four species of Pellaea sect. Platyloma

(J.Sm.) Hook. & Baker (Adiantaceae) from

Australia 341

Forster, P.I. with Reynolds, S.T. (2005). A

taxonomic revision of Tarenna Gaertn.

and Triflorensia S.T.Reynolds (Rubiaceae:

Ixoroideae : Pavetteae ) in Australia 29

Forster, P.I. with Reynolds, S.T. (2006). A

taxonomic revision of Ixora L. (Rubiaceae:

Ixoroideae ) in Australia 253

Garcinia mestonii 562

Gardenia edulis 152

Glomera 193

Glomera pseudomonanthos 193, 195

Glyceria latispicea 161

Gmelina leichhardtii 156,163

Goodenia 205

Goodenia argillacea 212

Goodenia armitiana 211

Goodenia armstrongiana 209, 211

Goodenia bicolor 211

Goodenia brachypoda 212

Goodenia byrnesii 213

Goodenia campestris 213

Goodenia chthonocephala 210

Austrobaileya 7(1 -4): 1-737

Goodenia cirrifica 211

Goodenia coronopifolia 211

Goodenia durackiana 212

Goodenia elaiosoma 205, 206, 207, 208, 212

Goodenia gloeophylla 210

Goodenia gracilis 211

Goodenia grandiflora 212

Goodenia heppleana 210

Goodenia hispida 213

Goodenia holtzeana 210

Goodenia janamba 211

Goodenia kakadu 210

Goodenia lamprosperma 211

Goodenia leiosperma 212

Goodenia malvina 213

Goodenia minutiflora 210

Goodenia neglecta 210

Goodenia odonnellii 212

Goodenia paludicola 211

Goodenia pilosa 210

Goodenia porphyrea 210

Goodenia potamica 212

Goodenia pumilio 210

Goodenia purpurascens 211

Goodenia purpurea 210

Goodenia quadrifida 210

Goodenia redacta 211,212

Goodenia scaevolina 210, 212

Goodenia sepalosa 212

Goodenia sp. Melville Island (N.B.Byrnes

666) 212

Goodenia strangfordii 212

Goodenia subauriculata 212

Goodenia symonii 210

Goodenia triodiophila 211

Goodenia viscidula 211

Goodeniaceae 205

Gossia 325

Gossia aneityensis 331,338

Gossia eugenioides 331, 338

Gossiafloribunda 331, 338

Gossia longipetiolata 328, 329, 330, 331

Gossia randiana 331,338

Gossia salomonensis 331, 338

Gossia scottiana 326, 327, 328, 331, 338

Gossia versteeghii 331,338

Grammitis albosetosa 563

Gray, B. with Jones, D.L. (2008). Corsia

dispar D.L.Jones & B.Gray (Corsiaceae),

a new species from Australia, and a new

combination in Corsia for a New Guinea

taxon 717

Grevillea ceratophylla 153

Grevillea lanceolata 153

Grevillea leichhardtii 155, 161

Grevillea leiophylla 161

Grevillea pungens 155, 161

Grevillea rubicunda 161

Grevillea singuliflora 162

Grimshaw, P. with Sparshott, K. & Duretto,

M.F. (2005). The rediscovery of Boronia

inflexa subsp. grandiflora (Rutaceae) 171

Guymer, G.P. (2005). New species of

Commersonia J.G.Forst. & G.Forst.

(Sterculiaceae) from eastern Australia and

Vanuatu 231

Guymer, G.P. (2006). New species of

Commersonia J.R.Forst. & G.Forst.

(Sterculiaceae) from Queensland 365

Guymer, G.P. (2008). Capparis batianoffii

Guymer (Capparaceae), a new species from

central coastal Queensland 723

Guymer, G.P. (2008). Notes on Palmeria

F.Muell. (Monimiaceae) in Australia and

the application of the name Palmeria

racemosa (Tul.) A.DC. 731

Guymer, G.P. (2008). Two new species of

Pandorea Spach (Bignoniaceae) recognised

from Queensland 735

Gynochthodes australiensis 362, 363

Gvnochthodes oresbia 357, 358,359,361,362,

363

Gynochthodes sessilis 362, 363

Gynochtodes sp. (Lamb Range) 358

Gyrostemon osmus 135, 136, 137, 138

Gyrostemon racemiger 138

Gyrostemon sect. Didymotheca 139

Gyrostemon sect. Gyrostemon 139

Gyrostemon sect. Monoecia 136,139

Gyrostemon tepperi 138

Gyrostemonaceae 135

Halford, D.A. (2005). Gyrostemon osmus

(Gyrostemonaceae), a new species from

south-eastern Queensland 135

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

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 215

Halford, D.A. & Henderson, R.J.F. (2007).

A taxonomic revision of Ricinocarpos

Desf. (Euphorbiaceae: Ricinocarpeae ,

Ricinocarpinae ) 387

Halford, D.A. & Henderson, R.J.F. (2008).

A taxonomic revision of Beyeria

Miq. (Euphorbiaceae: Ricinocarpeae ,

Ricinocarpinae ) 577

Auslrobai/eya 7 ( 1 - 4 ): 1-737

Halford, D.A. with Ford, A.J. (2006).

Gynochthodes oresbia Halford & A.J.Ford

(Rubiaceae), a new and cryptic species

from north-east Queensland 357

Halfordia kendack 155, 162

Halfordia leichhardtii 155

Harpullia frutescens 562

Harpullia leichhardtii 155, 162

Helichrysum diotophyllum 158

Helicia 561

Helicia nortoniana 561

Helicia whelanii 562

Hemecyclia lasiogyna 159

Hemistema lechenaultii 585

Henderson, R.J.F. with Halford, D.A. (2005).

Studies in Euphorbiaceae, s.lat. 7. Shonia

R.J.F.Hend. & Halford ( Ricinocarpeae,

Ricinocarpinae ), a new Australian endemic

genus 215

Henderson, R.J.F. with Halford, D.A. (2005).

Studies in Euphorbiaceae, s.lat. 6. A

revision of the genus Poranthera Rudge

( Antidesmeae, Porantherinae ) in Australia

Henderson, R.J.F. with Halford, D.A. (2007).

A taxonomic revision of Ricinocarpos

Desf. (Euphorbiaceae: Ricinocarpeae,

Ricinocarpinae ) 387

Henderson, R.J.F. with Halford, D.A.

(2008). A taxonomic revision of Beyeria

Miq. (Euphorbiaceae: Ricinocarpeae,

Ricinocarpinae ) 577

Holland, A.E. & Butler, D.W. (2007).

Trioncinia patens A.E.Holland &

D.W.Butler (Asteraceae: Coreopsideae:

Chrysanthellinae), a new and endangered

species from central Queensland 567

Homalocalyx polyandrus 161

Hovea heterophylla 159

Hovea linearis 159

Huperzia australiana 713

Hnperzia carinata 714

Huperzia dalhousieana 713

Huperzia filifor mis 714

Huperzia lockyeri 714

Huperzia marsupiiformis 714

Huperzia phlegmaria 713

Huperzia phlegmarioides 713

Huperziaprolifer a 712

Huperzia serrata 713

Huperzia squarrosa 713

Huperzia tetrasticha 712

Huperzia tetrastichoides 711, 712, 714

Huperzia varia 714

Hymenophyllum baileyanum 562

Hymenophyllum subdimidiatum 562

Hymenophyllum trichomanoides 562

Hymenophyllum tunbridgense var. exsertum

562

Hypnum leichardtii 155

Hyptiandra bidwillii var. grandiuscula 562

Hyrtanandra lythroides 163

Ichnanthus pallens var. major 562

Indigofera glandulosa 156

Ipomoeapolpha 311, 312, 313

Ipomoea polpha subsp. latzii 313

Ipomoea polpha subsp. polpha 313

Ipomoea polpha subsp. weirana 313, 314

Isotropisfilicaulis 155, 159

Ixora 29, 253, 255

Ixora baileyana 257, 259, 277

Ixora beckleri 258, 261, 277

Ixora bijlora 52, 260, 263, 277

Ixora biflora var. biflora 260

Ixora biflora var. fleckeri 260

Ixora biflora var. typica 260

Ixora chinensis 272

Ixora coccinea 256, 257, 272, 277

Ixora dallachiana 34, 273

Ixora expandens 36, 273

Ixorafinlaysoniana 272, 278

Ixora foetida 21A

Ixora indica 273

Ixora klanderiana 268

Ixora kochii 268

Ixora oreogena 262, 265, 278

Ixora orophila 254, 273

Ixora parviflora 272

Ixora pavetta 273

Ixora pentamera 40, 273

Ixora queenslandica 47, 52, 264, 267, 277

Ixora sect. Ixora 256

Ixora sect. Pavettopsis 256

Ixora sect. Phylleilema 256

Ixora sect. Pogonanthus 256

Ixora sect. Vitixora 256

Ixora sp. (N. Mary L.A. B.Hyland 8618) 262

Ixora sp. (North Mary LA B.P.Hyland 8618)

262

Ixora thozetiana 273

Ixora timorensis 268, 271, 278

Ixora tomentosa 274

Ixora triflora 47, 49, 51, 52, 264, 274

Ixora vinosa 274

Jackes, B.R. (2005). Studies in Australian

Myrsinaceae: Tapeinosperma Hook.f. 99

Jagerajavanica 379

Jagera javanica subsp. australiana 379

Jagera madida 379

Jagera serrata 379

Johnson, R.W. (2006). The enigmatic Ipomoea

polpha R.W. Johnson (Convolvulaceae) 311

Austrobaileya 7(1 - 4 ): 1-737

Jones, D.L. & Gray, B. (2008). Corsia dispar

D.L.Jones & B.Gray (Corsiaceae), a

new species from Australia, and a new

combination in Corsia for a New Guinea

taxon 717

Kalanchoe x houghtonii 383

Kallstroemia minuta 153

Kantvilas, G., McCarthy, P.M. & Stuckey,

B. (2008). A remarkable new species

of Rimularia Nyl. (lichenized fungi:

Trapeliaceae) from tropical Australia 659

Kellermann, J. (2006). Cryptandra triplex

K.R.Thiele ex Kellermann, a new species of

Rhamnaceae ( Pomaderreae ) from Arnhem

Land, Northern Territory 299

Kokubugata, G., Bostock, P.D. & Forster,

PI. (2006). Chromosome records for

four species of Pellaea sect. Platyloma

(J.Sm.) Hook. & Baker (Adiantaceae) from

Australia 341

Kruger, R. with Clarke, C. (2006). Nepenthes

tenax C.Clarke & R. Kruger (Nepenthaceae),

a new species from Cape York Peninsula,

Queensland 319

Kunzea opposita var. leichhardtii 155

Lamiaceae 707

Lasianthus graciliflorus 257

Lastreopsis microsora 561

Lavarack, PS. (2006). Arthrochilus

Javarackiana (D.L.Jones) Lavarack, a new

combination in Orchidaceae 385

Leichardtia 155

Leichardtia australis 155

Leichhardtia 155

Leptospermum wooroonooran 562

Leucopogon exolasius 159

Leucopogon pleiospermus 159

Limnostachys cyanea 161

Linospadix palmerianus 561

Liparis 737

Liparis petricola 737

Lithoxylon 63

Lithoxylon grandifolium 71

Lithoxylon lindleyi 57, 76

Lithoxylon nitidum 76

Livistona humilis 156

Livistona leichhardtii 156

Loheria 99

Lomatia silaifolia var. induta 162

Lomatia silaifolia var. silaifolia 162

Loranthus grandibracteus 152

Lucuma obpyriformis 678

Lycopodiaceae 711

Lysiphyllum cunninghamii 155

Lythraceae 727

Macadamia whelanii 562

Macropteranthes leichhardtii 156, 158

Madangia inflata 148

Maesa 100

Marsdenia 155

Marsdenia arachnoidea 145, 147, 148

Marsdenia australis 155, 156

Marsdenia leichhardtiana 156

Mastigobryum 130

McCarthy, P.M. with Stuckey, B. & Kantvilas,

G., (2008). A remarkable new species

of Rimularia Nyl. (lichenized fungi:

Trapeliaceae) from tropical Australia 659

McFadden, J., with Atwood, J.P & Snow, N.

(2008). Three new species of Rhodomyrtus

(DC.) Rchb. (Myrtaceae) from Papua New

Guinea 691

Meagher, D. (2005). Bazzania sauropoda

(Marchantiophyta: Lepidoziaceae), a new

species from tropical Queensland 129

Melicope chooreechillum 562

Melodorum leichhardtii 156

Mercurialis angustifolia 455

Mercurialis australis 457

Mercurialis tenerifolia 463

Meston, A. 675

Micrantheum ericoides var. juniper inum 159

Mischarytera lautereriana 281

Mischarytera macrobotrys 281

Mischarytera megaphylla 279, 280, 281, 282

Mischarytera sp. (Oliver Creek L.J.Webb+

10903) 280

Monimiaceae 731

Monochoria cyanea 161

Moore, D. with Booth, R. (2006). Reduction of

Carex rhytidocarpa Nelmes (Cyperaceae)

to a synonym of C. inversa R.Br. 373

Morinda leichhardtii 156

Morinda sp. (Black Leaves BG 1677) 358

Morinda sp. (Mt Finnigan L.J.Webb+ 12114)

358

Mullerochloa moreheadiana 561

Muntingia bartramia 239

Myrsinaceae 99

Myrsine subsessilis 160

Myrtaceae 111, 121, 141, 305, 325, 691

Myrtus metrosideros 562

Najas leichhardtii 156, 161

Nauclea oriental!s 156

Neckera leichhardtii 156,160

Neckera pennata 156, 160

Nepenthaceae 319

Nepenthes mirabilis 319, 323

Nepenthes rowanae 319, 323

Nepenthes tenax 319, 321, 323

Austrobaileya7(\-4)'. 1-737

Oberonia pusilla 562

Octarrhena pusilla 562

Omphalea queenslandiae 562

Orchidaceae 183, 385, 737

Oreoporanthera 1, 3

Oreoporantherapetalifera 22, 23

Orites fragrans 562

Ormerod, P. (2005). Papuasian Orchid Studies,

2 183

Ornithochilus moretonii 196

Ozothamnus diotophyllus 158

Palmeria 731

Palmeria coriacea 731, 732

Palmeria foremanii 731, 732

Palmeria hypotephra 731, 732

Palmeria racemosa 731,732

Palmeria scandens 731,733

Pandanus spiralis 153

Pan dore a 735

Pandorea australis subsp. linearis 736

Pandoreafloribunda 736

Pandorea linearis 736

Pandorea sp. (K.A.Williams 86020) 736

Panicum buncei 161

Panicum incomtum 562

Panicum prenticeanum 562

Panicum vicinum 562

Parinari nonda 158

Parsonsia leichhardtii 156, 158

Pavetta 29, 31

Pavetta australiensis 273, 274

Pavetta brownii 274

Pavetta timorensis 268

Pavetta triflora 47

Pavetteae 30

Pedley, L. (2006). Nomenclatural notes on

Acacia Mill. (Leguminosae - Mimosaceae),

consequential to the conservation of its

name 381

Pedley, L. (2006). Notes on Acacia Mill.

(Leguminosae: Mimosoideae ), chiefly from

Queensland, 5 347

Pellaea calidirupium 342, 343, 344

Pellaea falcata 342, 343, 344

Pellaea nana 342, 343, 344

Pellaea paradoxa 342, 343, 344

Pellaea sect. Platyloma 341

Pentaceras australis 162

Pentapanax bellendenkerensis 678

Peripentadenia mearsii 92

Philotheca 175

Philotheca acrolopha 175

Philotheca conduplicata 176, 177

Philotheca epilosa 176, 177, 178

Philotheca glasshousiensis 176, 178, 179

Philotheca myoporoides 175

Philotheca myoporoides subsp. conduplicata

176, 177

Philotheca myoporoides subsp. epilosa 177

Philotheca myoporoides subsp. leichhardtii

155, 162, 178

Philotheca myoporoides subsp. obovatifolia

179

Philotheca myoporoides subsp. queenslandica

180

Philotheca obovatifolia 176, 179

Philotheca queenslandica 176, 179, 180

Philotheca sect. Erionema 176

Phoringopsis lavarackiana 385

Phyllanthaceae 57

Phyllanthus 155

Piper mestonii 562

Plagianthus monoicus 437

Plectranthus batianoffti 707, 709

Poaephyllum fuscum 183

Polypodium albosetosum 563

Polyscias bellendenkerensis 678

Pomaderreae 299

P or anther a 1, 3, 27, 58

P or anther a alpina 2, 3

Poranthera arbuscula 5

Poranthera cicabricosa 7

Poranthera coerulea 4, 5

Poranthera corymbosa 4, 5, 6, 7

Poranthera corymbosa var. arbuscula 5, 6

Poranthera corymbosa var. corymbosa 5

Poranthera corymbosa var. genuina 5

Poranthera corymbosa var. linarioides 5

Poranthera corymbosa var. sparsifolia 5

Poranthera dissecta 4, 7, 8

Poranthera drummondii 4, 8, 9, 27

Poranthera ericifolia 4, 10, 27

Poranthera ericoides 4, 10, 12, 27

Poranthera ericoides var. ericoides 10

Poranthera ericoides var. glauca 24

Poranthera ericoides var. typica 11

Poranthera florosa 4, 12, 13, 27

Poranthera glauca 24

Poranthera huegelii 4, 13, 15, 24, 27

Poranthera leiosperma 3, 4, 15, 16, 27

Poranthera linearoides 5

Poranthera microphylla 4, 16, 18, 24, 27

Poranthera microphylla var. diffusa 16

Poranthera microphylla var. drummondii 8

Poranthera microphylla var. genuina 16

Poranthera microphylla var. glauca 16

Poranthera microphylla var. intermedia 12,

Poranthera microphylla var. microphylla 16

Poranthera microphylla var. procera 24

Poranthera obovata 4, 19, 27

Poranthera oreophila 3, 20, 21, 22, 27

Porantherapetalifera 3, 22, 23, 27

Poranthera piceoides 11

Poranthera sp. (Beerwah D.Hassall DH7431)

Poranthera subgen. Euporanthera 2, 3

Poranthera subgen. Oreoporanthera 2, 3

Poranthera subgen. Poranthera 2, 3

Poranthera triandra 3, 4, 23, 24, 27

Porantherinae 1

Pouteria obpyriformis 678

Pouzolzia hirta 163

Prostanthera leichhardtii 156

Prostanthera ringens 156

Proteaceae subtribe Stenocarpinae 169

Pseuderia 193

Pseuderia brevifolia 196

Pseuderia takeuchii 196, 197

Pseudoixora 32, 40

Psilanthus 30

Psoralea leichhardtii 15 6

Psychotria coelospermum 678

Psydrax montigena

Pterostylis 737

Pterostylis vitrea 737

Pultenaea paleacea var. obtusata 160

Pultenaea paleacea var. paleacea 160

Quassia baileyana 562

Racosperma calligerum 381

Racosperma leptostachyum 354

Racosperma ligulatum var. minus 381

Racosperma microcephalum 352

Racosperma serpenticola 381

Racosperma tindaleae 347

Rapanea subsessilis 160

Reynolds, S.T. & Forster, PI. (2005). A

taxonomic revision of Tarenna Gaertn.

and Triflorensia S.T.Reynolds (Rubiaceae:

Ixoroideae : Pavetteae ) in Australia 29

Reynolds, S.T. & Forster, PI. (2006). A

taxonomic revision of Ixora L. (Rubiaceae:

Ixoroideae ) in Australia. 253

Rhamnaceae 299

Rhodomyrtus 691

Rhodomyrtus canescens 694

Rhodomyrtus effusa 694

Rhodomyrtus e/egans 337, 338, 694

Rhodomyrtus guymeriana 692, 693, 694,

695

Rhodomyrtus kaweaensis 331, 332, 334, 335,

338, 694

Rhodomyrtus lanata 335, 338, 694

Rhodomyrtus locellata 337, 338, 694

Rhodomyrtus longipetiolata 338

Rhodomyrtus longisepala 694, 696, 697

Rhodomyrtus macrocarpa 337, 338, 694

Austroba.ileya 7(1 - 4 ): 1-737

Rhodomyrtus mengenensis 333, 335, 336,

338, 339, 694

Rhodomyrtus misimana 694, 698, 699, 702

Rhodomyrtus montana 335, 338, 694, 702

Rhodomyrtus novoguineensis 335, 338, 694

Rhodomyrtus obovata 337, 338, 694

Rhodomyrtus pervagata 694

Rhodomyrtuspinnatinervis 335, 338, 694

Rhodomyrtus psidioides 694

Rhodomyrtus salomonensis 335, 338, 694

Rhodomyrtus sericea 694

Rhodomyrtus surigaoensis 335, 338, 694

Rhodomyrtus tomentosa 335, 338, 694

Rhodomyrtus trineura 694

Rhus tomentosa 158

Rhus viticifolia 158

Ricinocarpos 215, 216, 217, 387, 392, 397

Ricinocarpos bowmanii 389, 394, 396, 397

Ricinocarpos bowmanii var. albus 397

Ricinocarpos bowmanii var. bowmanii 397

Ricinocarpos bowmanii var. genuinus 397

Ricinocarpos bowmanii var. planus 397

Ricinocarpos brevis 389, 394, 395, 399, 400

Ricinocarpos caniana 389, 394, 396, 401,

403

Ricinocarpos crispatus 389, 394, 396, 402,

405

Ricinocarpos cyanescens 389, 393, 396, 406

Ricinocarpos glaucus 389, 393, 395, 407

Ricinocarpos glaucus var. cyanescens 406

Ricinocarpos glaucus var. genuinus 407

Ricinocarpos glaucus var. glaucus 407

Ricinocarpos glaucus var. jasminoides 407

Ricinocarpos glaucus var. undulatus 435

Ricinocarpos gloria-medii 389, 397, 408

Ricinocarpos graniticus 389, 393, 395, 410,

411

Ricinocarpos ledifolius 389, 394, 396, 412

Ricinocarpos linearifolius 389, 394, 395, 396,

413, 415

Ricinocarpos major 443

Ricinocarpos marginatus 389, 394, 397, 416

Ricinocarpos megalanthus 422

Ricinocarpos megalocarpus 389, 393, 396,

417, 419

Ricinocarpos mitchellii 443

Ricinocarpos muricatus 389, 393, 395, 440

Ricinocarpos neocaledonicus 443

Ricinocarpos oliganthus 389, 393, 395, 418

Ricinocarpospilifer 389, 393, 395, 420, 421

Ricinocarpospinifolius 389, 393, 396, 422

Ricinocarposproximus 422

Ricinocarpospsilocladus 389, 393, 395, 424

Ricinocarpos puberulus 397

Ricinocarpos rosmarinifolius 389, 394, 397,

425

Auslrobai/eya 7 ( 1 - 4 ): 1-737

Ricinocarpos ruminatus 389, 393, 396, 426,

427

Ricinocarpos sect. Amonodiscus 443

Ricinocarpos sect. Apetalidion 440

Ricinocarpos sect. Bertya 397

Ricinocarpos sect. Euricinocarpus 397

Ricinocarpos sect. Polystaphylos 440

Ricinocarpos sect. Ricinocarpos 397

Ricinocarpos sect. Scissostylus 438

Ricinocarpos sidifolius 422

Ricinocarpos sp. (Blackdown Tableland

R.J.Henderson H610) 413

Ricinocarpos sp. (Cania Gorge K.A.Williams

80221) 401

Ricinocarpos sp. (Mt Alto C.T.White 10633)

441

Ricinocarpos sp. (Mt C alder Station

M.E.Ballingall MEB1588) 404

Ricinocarpos sp. (Planet Downs A.R.Bean

14223) 426

Ricinocarpos sp. (Westmar T.J.McDonald

263) 429

Ricinocarpos sp. A. 433

Ricinocarpos sp. A. Kimberley Flora

(A.S.George 14118) 433

Ricinocarpos speciosus 389, 394, 396, 428

Ricinocarpos stylosus 389, 393, 395, 438

Ricinocarpos tasmanicus 443

Ricinocarpos trachyphyllus 389, 394, 396,

429, 431

Ricinocarpos trichophorus 389, 394, 395, 430

Ricinocarpos trichophyllus 389, 394, 397,

432, 434

Ricinocarpos tuberculatus 389, 393, 396, 434

Ricinocarpos undulatus 389, 393, 395, 435

Ricinocarpos velutinus 389, 394, 395, 437

Ricinocarpos verrucosus 389, 393, 396, 441,

442

Rimularia 659

Rimularia cerebriformis 660, 661, 662

Robiquetia 196

Robiquetia brassii 198, 199, 200

Robiquetia camptocentrum 199

Robiquetia hamata 199

Roeperia bowmanii 397

Roeperia cyanescens 406

Roeperia glauca 407

Roeperia ledifolia 412

Roeperia major 442

Roeperia marginata 416

Roeperia muricata 440

Roeperia pinifolia 422

Roeperia psiloclada 424

Roeperia rosmarinifolia 425

Roeperia speciosa 428

Roeperia trichophora 430

Roeperia tuberculata 434

Roeperia velutina 437

Rondeletia asiatica 31

Rotala occultiflora var. leichhardtii 156

Rotala occultiflora var. occultiflora 156

Rubiaceae 29, 253, 357

Rubiaceae tribe Pavetteae 30

Rubus moorei v ar. leichardtianus 156, 162

Rubus moorei var. moorei 156, 162

Rulingia procumbens 371

Rutaceae 171, 175, 473, 665, 681

Saccolabium mooreanum 196

Sapindaceae 279, 285, 293, 379

Sapindaceae sp. (Noah Creek BG 6026) 279

Sarcocephalus leichhardtii 156

Sarcochilus obtusus 192

Sarcotoechia sp. (Mountain Sarcotoechia

WWC100) 286

Sarcotoechia sp. (Mt Carbine L.W.Jessup +

GJM995) 286

Savia sect. Actephila 63

Scaevola enantophylla 563

Scaevola scandens 563

Scleria ustulata 563

Scleroglossum wooroonooran 563

Securinega nitida 63, 75

Serruria 168

Sersalisia sericea 678

Shonia 215, 216, 217, 218, 228

Shonia bickertonensis 219, 223, 225, 632

Shonia carinata 218, 225, 226, 228

Shonia territorialis 219, 222, 223, 224

Shonia tristigma 218, 219, 220, 632

Shonia tristigma subsp. borealis 220, 221

Shonia tristigma subsp. parvifolia 220, 222

Shonia tristigma subsp. tristigma 220, 222

Snow, N. (2006). New species of Gossia

N.Snow & Guymer and Rhodomyrtus

(DC.) Hassk. (Myrtaceae) from Papua New

Guinea 325

Snow, N., McFadden, J. & Atwood, J.P (2008).

Three new species of Rhodomyrtus (DC.)

Rchb. (Myrtaceae) from Papua New Guinea

691

Solanaceae 669

Solanum chippendalei 669, 672

Solanum sp. Juicy Fruit (P.K.Fatz 18938) 670

Solanum succosum 669,671

Sophora fraseri 160

Sorghum laxiflorum 563

Sparshott, K. with Duretto, M.F. & Grimshaw,

P. (2005). The rediscovery of Boronia

inflexa subsp. grandiflora (Rutaceae) 171

Sporobolus caroli 161

Stenocarpinae 169

Stenocarpus 165, 166, 167, 169

Stenocarpus salignus 165

Stenocarpus sinuatus 165

Sterculiaceae 231, 365

Stereum ostrea 163

Strangea 165, 166, 167, 168, 169

Strangea linearis 165

Strychnos bancroftiana 563

Strychnos minor 563

Stuckey, B. with Kantvilas, G. & McCarthy,

P.M. (2008). A remarkable new species

of Rimularia Nyl. (lichenized fungi:

Trapeliaceae) from tropical Australia 659

Stylocoryna dallachiana 34

Stylocoryne 31, 44

Styphelia exolasia 159

Styphelia pleiosperma 159

Suregada glomerulata 573

Swainsona brachycarpa 160

Swainsona oroboides 160

Symplocos paucistaminea 563

Synima cordierorum 289

Synima macrophylla 289

Synima reynoldsiae 285, 286, 288, 289, 290

Tainia 199

Tainia serratiloba 199,201,202

Tainia trinervis 199

Takeuchi, W., with Forster, PI. (2005).

Rediscovery and neotypification

of Marsdenia arachnoidea Schltr.

(Apocynaceae: Asclepiadoideae

Marsdenieae ), an endangered species from

Papua New Guinea 145

Tapeinosperma 99, 100, 101

Tapeinosperma flueckigeri 107

Tapeinosperma pallidum 101, 104, 109, 110

Tapeinosperma pseudojambosa 101, 103, 106,

110

Tapeinosperma repandulum 101, 105, 107,

108, 110

Tapeinosperma sp. (Cedar Bay J.G.Tracey

14780) 102

Tapeinosperma sp. (Woombye A.R. Bean 994)

105

Tapeinosperma vieillardii 100

Tarenna 29, 30, 31 32, 44

Tarenna appressa 33

Tarenna asiatica 33

Tarenna australis 45

Tarenna cameronii 46

Tarenna dallachiana 30, 33, 34, 35

Tarenna dallachiana subsp. dallachiana 33,

35, 54

Tarenna dallachiana subsp. expandens 35,

36, 37, 38, 273

Tarenna expandens 36, 37

Tarenna foliosa 36

Tarenna monticola 33, 38, 39, 40, 54

Austrobaileya 7 ( 1 - 4 ): 1-737

Tarenna pentamera 40, 41, 42, 43

Tarenna sambucina 33

Tarenna sect. Multiovulatae 33

Tarenna sect. Pauciovulatae 40

Tarenna sp. (Ka Ka Mundi NP W. J. McDonald

+ 4642) 49

Tarenna sp. (Mt Lewis B.P.Hyland 3403) 38

Tarenna sp. A 40

Tarenna subg. Pacifica 32, 33

Tarenna subg. Pseudoixora 32, 40

Tarenna subg. Tarenna 32, 33

Tarenna triflora 44, 49, 52

Tecoma australis var. linearis 736

Tecomafloribunda 736

Templetonia muelleri 160

Templetonia stenophylla 160

Tephrosiafilipes 160

Tephrosiajuncea 160

Tephrosiapurpurea var. rufescens 160

Tephrosia rufula 160

Terminalia platyptera 158

Thelephora leichhardtiana 163

Thryptomene polyandra 161

Trapeliaceae 659

Tribulopsis minutus 153

Tribulus micrococcus 154

Tribulus minutus 153, 163

Tribulus parviflorus 154

Tricalysia 30

Trichomanes barnardianum 563

Trifiorensia 29, 31, 43, 44

Trifiorensia australis 43, 45, 46, 55

Trifiorensia cameronii 45, 46, 47, 48

Trifiorensia ixoroides 44, 47, 50, 51, 52, 274

Trionciniapatens 567, 568, 569

Trioncinia retroflexa 568

Tripogon loliiformis 161

Unona leichhardtii 156

Urera excelsa 163

Urera leichardiana 163

Urera leichhardtiana 156

Uromyrtus metrosideros 562

Urostigma le ichhardtii 156

Vacoparis laxiflorum 563

Vallisneria gracilis 563

Vallisneria nana 563

Vignea inversa 373

Vitex leichhardtii 156, 163

Vittaria ( Taeniopsis ) wooroonooran 563

Webera 31

Webera dallachiana 34, 37

Webera dallachiana var. expandens 36

Webera expandens 36

Webera sect. Pseudoixora (‘pseudixora) 40

Auslrobai/eya 7 ( 1 - 4 ): 1-737

Webera sect. Webera 33

Xylomelum scottianum 152

Zamia australis 153

Ziera sp. ‘Cathedral Rock’ (Williams 95303)

681

Zieria 473

Zieria actites 480, 481

Zieria adenodonta 476, 482

Zieria adenophora 682

Zieria alata 480, 482, 484

Zieria arborescens 477, 479, 485

Zieria arborescens subsp. arborescens 485

Zieria arborescens subsp. glabrifolia 485

Zieria aspalathoides 478, 479, 486

Zieria aspalathoides subsp. aspalathoides

487

Zieria aspalathoides subsp. brachyphylla 487,

488

Zieria aspalathoides var. (Springsure

L.Cockburn AQ195493) 487

Zieria aspalathoides var. intermedia 536

Zieria aspalathoides var. obovatum 524

Zieria bifida 479, 488, 490

Zieria boolbunda 476, 480, 491, 492

Zieria cephalophila 477, 492, 493

Zieria collina 477, 494

Zieria compacta All, 478, 495

Zieria compacta var. glabrata 506

Zieria compacta var. robusta 506

Zieria cytisoides All, 497

Zieria distans All, 497, 499

Zieria eungellaensis 500, 501

Zieria exsul All, 502, 504

Zieriafloydii 682

Zieria fraseri 478, 503, 506

Zieria fraseri subsp. compacta 495

Zieria fraseri subsp. fraseri 506

Zieria fraseri subsp. robusta 505, 506

Zieriafurfuracea 476, 508

Zieria furfuracea subsp. (Belmont Scrub

Anon. AQ152898) 509

Zieriafurfuraceasubsp. (KinKinV.K.Moriarty

134) 508

Zieria furfuracea subsp. euthadenia 508

Zieria furfuracea subsp. gymnocarpa 508,

509

Zieria graniticola 476, 479, 509, 511

Zieria granulata var. adenodonta 482

Zieria hydroscopica A19, 512, 513

Zieria inexpectata 476,514, 515

Zieria insularis 479, 516, 517

Zieria laevigata 478, 518

Zieria laevigata subsp. (Wyberba T.L.Ryan

37) 518

Zieria laevigata var. fraseri 503

Zieria laevigata var. laxiflora 518

Zieria laxiflora 478,518

Zieria madida 480, 519, 521

Zieria minutifiora 478, 479, 520

Zieria minutifiora subsp. (Danbulla L.J.Webb

5732) 522

Zieria minutifiora subsp. minutifiora 522

Zieria minutifiora subsp. trichocarpa 522

Zieria montana 480, 523

Zieria obcordata 682

Zieria obovata 478, 479, 524

Zieria odorifera 681, 682

Zieria odorifera subsp. copelandii 684, 685

Zieria odorifera subsp. warrabahensis 684,

687

Zieria odorifera subsp. williamsii 686, 688,

689

Zieria rimulosa 479, 525

Zieria robertsiorum 479, 480, 525

Zieria scopulus 480, 526, 528

Zieria smithii 476, 479, 528, 682

Zieria smithii subsp. (SF144 B.Gray 428) 516

Zieria smithii subsp. tomentosa 528

Zieria southwellii 479, 530

Zieria sp. (Amiens L.Pedley 1518) 509

Zieria sp. (Binjour PI.Forster PIF14134) 533

Zieria sp. (Brolga Park A.R.Bean 1002) 489

Zieria sp. (Coominglah A.R.Bean 8959) 512

Zieria sp. (Flinders Peak S.L.Everist 1169)

527

Zieria sp. (Herberton J.A.Armstrong 1025)

524

Zieria sp. (Lamington G.Leiper AQ502702)

530

Zieria sp. (Monogorilby PI. Forster PIF1004)

535

Zieria sp. (Mooloolaba G.Leiper AQ636552)

502

Zieria sp. (Mt Ballow, G.Leiper AQ473220)

529

Zieria sp. (Mt Larcom N.Gibson TOI8) 480

Zieria sp. (Pieter Botte M.Godwin C2471)

519

Zieria sp. (Ravenshoe E.W.Bick 116) 536

Zieria sp. (Russell River S.Johnson in 1892)

522

Zieria sp. (Sydney Heads A.R.Bean 2562)

492

Zieria sp. (Thornton Peak J.R.Clarkson 5556)

519

Zieria sp. (White Mt D.G.FelH DGF1257)

531

Zieria sp. ‘Cathedral Rock’ (J.B.Williams

95303) 686

Zieria sp. ‘Oxley Wild Rivers N.P’ (Copeland

2174) 686

Zieria sp. 1 535

A ustrobaileya 7 ( 1 - 4 ): 1-737

Zieria sp. 12 (sp. “L”, Russell River, S. Johnson

s.n. 1892 [189]) 522

Zieria sp. nov. (Mt William) 500

Zieria sp. Oxley Wild Rivers N.R (Copeland

2174) 681

Zieria tenuis 477,531,532

Zieria vagans All , 478, 533, 534

Zieria verrucosa 476, 535

Zieria whitei All , 478, 536, 537

Ziziphus pomaderroides 377, 378

Austrobaileya 7(1-4): 1-737

Contents

Number 1 2005

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 & P.I. Forster .29-55

A taxonomic revision of Actephila Blume (Euphorbiaceae/Phyllanthaceae)

in Australia

P.I. 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, 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.R. Bean .141-144

Rediscovery and neotypification of Marsdenia arachnoidea Schltr.

(Apocynaceae: Asclepiadoideae - Marsdenieae ), an endangered species

from Papua New Guinea

P.I. 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

P.I. Forster .175-181

Papuasian Orchid Studies, 2

P. Ormerod . 183-203

Austrobaileya 7(1-4): 1-737

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

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

Number 2 2006

A taxonomic revision of Ixora R.Br. (Rubiaceae: Ixoroideae ) in Australia

S.T. Reynolds & P.I. Forster . 253-278

Mischarytera megaphylla RI.Forst. (Sapindaceae), a new species from the

‘Wet Tropics’ of north-east Queensland

P.I. Forster . 279-283

Synima reynoldsiae RI.Forst. (Sapindaceae), a new species from the ‘Wet

Tropics’ of north-east Queensland

P.I Forster . 285-291

Cupaniopsispapillosa RI.Forst. (Sapindaceae), a new species from the ‘Wet

Tropics’ of north-east Queensland

P.I. Forster . 293-298

Cryptandra triplex K.R.Thiele ex Kellermann, a new species of Rhamnaceae

( Pomaderreae ) from Arnhem Land, Northern Territory

J. Kellermann . 299-303

Eucalyptus megasepala A.R.Bean (Myrtaceae), a new species from

Queensland allied to E. tetrodonta F.Muell.

A.R. Bean . 305-310

The enigmatic Ipomoea polpha R.W. Johnson (Convolvulaceae)

R.W. Johnson .311-317

Nepenthes tenax C.Clarke & R.Kruger (Nepenthaceae), a new species from

Cape York Peninsula, Queensland

C. Clarke & R. Kruger .319-324

New species of Gossia N.Snow & Guymer and Rhodomyrtus (DC.) Hassk.

(Myrtaceae) from Papua New Guinea

N. Snow . 325-340

Chromosome records for four species of Pellaea section Platyloma (J.Sm.)

Hook. & Baker (Adiantaceae) from Australia

G. Kokubugata, P.D. Bostock & P.I. Forster . 341-345

Notes on Acacia Mill. (Leguminosae: Mimosoideae ), chiefly from

Auslrobai/eya 7(1-4): 1-737

Queensland, 5

L. Pedley . 347-356

Gynochthodes oresbia Halford & A.J.Ford (Rubiaceae), a new and cryptic

species from north-east Queensland

A.J. Ford &D.A. Halford . 357-364

Three new species of Commersonia J.R.Forst. & G.Forst. (Sterculiaceae)

from Queensland

G.P. Guymer . 365-372

Reduction of Carex rhytidocarpa Nelmes (Cyperaceae) to a synonym of C.

inversa R.Br.

R. Booth & D. Moore . 373-375

A new combination in Alphitonia Endl. (Rhamnaceae)

A.R. Bean . 376-378

Jagera madida P.I.Forst. (Sapindaceae), a new name and change of rank for

J. javanica subsp. australiana Leenh.

P. I. Forster .379

Nomenclatural notes on Acacia Mill. (Leguminosae - Mimosoideae ),

consequential to the conservation of its name

L. Pedley . 381-382

Bryophyllum x houghtonii (D.B.Ward) P.I.Forst., a new combination in

Crassulaceae for the hybrid Mother of Millions

P. I. Forster .383

Arthrochilus lavarackiana (D.L.Jones) Lavarack, a new combination in

Orchidaceae

P.S. Lavarack .385

Number 3 2007

A taxonomic revision of Ricinocarpos Desf. (Euphorbiaceae: Ricinocarpeae,

Ricinocarpinae )

D. A. Halford & R.J.F.Render son . 387-449

A taxonomic revision of Claoxylon A. Juss. (Euphorbiaceae) in Australia

P.I.Forster . 451-472

A taxonomic revision of the genus Zieria Sm. (Rutaceae) in Queensland

M.F.Duretto & P.I.Forster . 473-544

A taxonomic revision of Callitriche L. (Callitrichaceae) in Australia

A. R. Be an . 545-554

F.M.Bailey’s ascent of Mt Bellenden-Ker in 1889, and notes on the publication

priority of new vascular plant species from the Expedition

J.L.Dowe & A.D.Broughton . 555-566

Trioncinia patens A.E.Holland & D.W.Butler (Asteraceae), a new species

from Queensland

A.E.Holland & D.W.Butler . 567-571

The distribution, habitat and conservation status of Suregada glomerulata

(Blume) Baill. (Euphorbiaceae) from north-eastern Cape York Peninsula,

Auslrobai/eya 7(1-4): 1-737

Queensland

D.G.Fell . 573-575

Number 4 2008

A taxonomic revision of Beyeria Miq. (Euphorbiaceae: Ricinocarpeae ,

Ricinocarpinae )

D.A.Halford & R.J.F.Henderson . 577-639

A taxonomic revision of Erythrina L. (Fabaceae: Faboideae ) in Australia

A. R. Be an . 641-658

A remarkable new species of Rimularia Nyl. (lichenized fungi: Trapeliaceae)

from tropical Australia

G.Kantvilas, P.McCarthy & B. Stuckey . 659-663

A new subsection and two new subseries for Boronia Sm. Section Valvatae

(Benth.) Engl. (Rutaceae)

M.F.Duretto . 665-668

Solanum succosum A.R.Bean & Albr. (Solanaceae), a new species allied to

S. chippendalei Symon

A.R.Bean & D.E.Albrecht . 669-675

F.M.Bailey’s taxonomy of A.Meston’s collections from the Bellenden Ker

Expedition of 1904

J.L.Dowe & A.D.Broughton . 677-679

New subspecies for Zieria odorifera J.A.Armstr. (Rutaceae) from northern

New South Wales

M. F.Duretto & P.I.Forster . 681-690

Three new species of Rhodomyrtus (DC.) Rchb. (Myrtaceae) from Papua

New Guinea

N. Snow, JMcFadden & J.P.Atwood . 691-706

Plectranthus batianoffii P.I.Forst. (Lamiaceae), a new species from north¬

east Queensland

P.I.Forster . 707-710

Huperzia tetrastichoides A.R.Field & Bostock (Fycopodiaceae) a newly

recognised species of tassel fern from the Wet Tropics of Queensland,

Australia

Ashley R. Field & Peter D.Bostock .711-715

Corsia dispar D.E. Jones &B.Gray (Corsiaceae), a new species from Australia,

and a new combination in Corsia for a New Guinea taxon

D.L. Jones & B.Gray .717-722

Capparis batianoffii Guymer (Capparaceae), a new species from central

coastal Queensland

G.P Guymer . 723-725

Reinstatement of Ammannia triflora Benth. (Fythraceae)

A.R.Bean . 727-728

A new combination in Cissocarpus Rozefelds (Vitaceae), a fossil genus from

Queensland