Editorial Committee

P.I.Forster (editor)

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Graphic Design

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Austrobaileya

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

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

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Austrobaileya 8 ( 4 ): 441-723 ( 2012 )

Contents

A taxonomic revision of Euphorbia section Anisophyllum Roeper

(Euphorbiaceae) in Australia

D. A.Halford & W.K.Harris . 441-600

Utricularia corneliana R.W. Jobson (Lentibulariaceae), a new species from

the North Kennedy district of Queensland

R.W. Jobson . 601-607

Stylidium elachophyllum A.R.Bean & M.T.Mathieson (Stylidiaceae), a new

species from northern Queensland

A.R.Bean & M.T.Mathieson . 608-612

Poikilogyne cor difolia (Cogn.) Mansf., a newly recorded genus and species of

Melastomataceae for Australia

A. Field .613-615

Croton lucens RI.Forst. (Euphorbiaceae), a new species from south-east

Queensland

P.I.Forster .616-623

Capsule dehiscence in Viola betonicifolia Sm. (Violaceae)

R.J. Little & G.Leiper . 624-633

A revision of Calyptochloa C.E.Hubb. (Poaceae), with two new species and

a new subspecies

E. J.Thompson & B.K.Simon . 634-652

The Australian species of Blainvillea Cass. (Asteraceae: Ecliptinae)

A. E.Orchard . 653-669

A taxonomic revision of Hollandaea F.Muell. (Proteaceae)

A.J.Ford& P.H.Weston . 670-687

Nomenclature and synonymy of several Goodenia R.Br. (Goodeniaceae)

species from northern Australia

A. E. Holland . 688-695

Types of enigmatic north-Queensland Orchids from the Dockrill herbarium

A.R.Field & F.A.Zich . 696-698

New species and subspecies of Ipomoea L. (Convolvulaceae) from northern

Australia and a key to the Australian species

R.W. Johnson . 699-723

NOT AS PUBLISHED - plates 27 & 28

edited to match published captions

A taxonomic revision of Euphorbia section Anisophyllum

Roeper (Euphorbiaceae) in Australia

David A. Halford & Wayne K. Harris

Summary

Halford, D A. & Harris, W.K. (2012). A taxonomic revision of Euphorbia section Anisophyllum

Roeper (Euphorbiaceae) in Australia. Auslrobaileya 8(4): 441-600. A systematic study of Euphorbia

section Anisophyllum Roeper in Australia is presented. A total of 58 species are recognised of which

51 are native (48 endemic) and seven are naturalised. Twenty one species are described here as new: E.

accedens Halford & W.K.Harris, E. albrechtii Halford & W.K.Harris, E. crassimarginata Halford &

W.K.Harris, E.fitzroyensis Halford & W.K.Harris, E. gregoriensis Halford & W.K.Harris, E. hassallii

Halford & W.K.Harris, E. laciniloba Halford & W.K.Harris, E. litticola Halford & W.K.Harris,

E. macdonaldii Halford & W.K.Harris, E. multifaria Halford & W.K.Harris, E. occulta Halford &

W.K.Harris, E. papillata Halford & W.K.Harris, E. papillifolia Halford & W.K.Harris, E. philochalix

Halford & W.K.Harris, E. porcata Halford & W.K.Harris, E. psilosperma Halford & W.K.Harris, E.

thelephora Halford & W.K.Harris, E. trigonosperma Halford & W.K.Harris, E. verrucitesta Halford &

W.K.Harris, E. vicina Halford & W.K.Harris and E. victoriensis Halford & W.K.Harris. New species

are illustrated while all species are described and their distributional range mapped, and notes on their

distribution, habitat and phenology are given. Eleven new varieties are described as new: E. australis

var. glabra Halford & W.K.Harris, E. australis var. hispidula Halford & W.K.Harris, E. ferdinandi

var. appendiculata Halford & W.K.Harris, E. ferdinandi var. saxosiplaniticola Halford & W.K.Harris,

E. inappendiculata var. robustior Halford & W.K.Harris, E. mitchelJiana var. longiloba Halford &

W.K.Harris, E. papillata var. Jaevicaulis Halford & W.K.Harris, E. papillifolia var. polyandra Halford

& W.K.Harris, E. thelephora var. australis Halford & W.K.Harris, E. thelephora var. rugosa Halford

& W.K.Harris, and E. vaccaria var. erucoides Halford & W.K.Harris. The new combinations E.

macdonaldii var. potentillina (Baill.) Halford & W.K.Harris, based on E. australis var. potentillina

Baill., E. mitchelJiana var. filipes (Benth.) Halford & W.K.Harris, based on E. filipes Benth., and

E. schultzii var. comans (W.Fitzg.) Halford & W.K.Harris based on E. comans W.Fitzg., are made.

Lectotypes are chosen for E. alsiniftora Baill., E. armstrongiana Boiss., E. australis var. semiglabra

Domin, E. comans W.Fitzg., E. distans W.Fitzg., E. erythrantha F.Muell., E. inappendiculata Domin,

E. filipes Benth., E. macgillivrayi Boiss., E. macgillivrayi var. yarrabensis Domin, E. macgillivrayi var.

pseudoserndata Domin, E. macgillivrayi f. glabrata Domin, E. mitchelliana Boiss., E. mitchelJiana

var. cairnsiana Domin, E. mitchelJiana var. dietrichiae Domin, E. mitchelJiana var. glauca Benth., E.

mitchelJiana var. stenophylla Benth., E. petala Ewart & L.R.Kerr, E. schizolepis F.Muell. ex Boiss. and

E. schultzii Benth. Keys to identify the species and varieties are provided.

Key Words: Euphorbiaceae, Euphorbia section Anisophyllum Roeper, Chamaesyce , Australia flora,

taxonomy, nomenclature, identification keys, E. accendens Halford & W.K.Harris, E. albrechtii

Halford & W.K.Harris, E. australis var. glabra Halford & W.K.Harris, E. australis var. hispidula

Halford & W.K.Harris, E. crassimarginata Halford & W.K.Harris, E. ferdinandi var. appendiculata

Halford & W.K.Harris, E. ferdinandi var. saxosiplantiticola Halford & W.K.Harris, E. fitzroyensis

Halford & W.K.Harris, E. gregoriensis Halford & W.K.Harris, E. hassallii Halford & W.K.Harris,

E. inappendiculata var. robustior Halford & W.K.Harris, E. laciniloba Halford & W.K.Harris, E.

litticola Halford & W.K.Harris, E. macdonaldii Halford & W.K.Harris, E. mitchelJiana var. longiloba

Halford & W.K.Harris, E. multifaria Halford & W.K.Harris, E. occulta Halford & W.K.Harris, E.

papillata Halford & W.K.Harris, E. papillata var. laevicaulis Halford & W.K.Harris, E. papillifolia

Halford & W.K.Harris, E. papillifolia var. polyandra Halford & W.K.Harris, E. philochalix Halford &

W.K.Harris, E. porcata Halford & W.K.Harris, E. psilosperma Halford & W.K.Harris, E. thelephora

Halford & W.K.Harris, E. thelephora var. australis Halford & W.K.Harris, E. thelephora var. rugosa

Halford &W.K.Harris, E. trigonosperma Halford & W.K.Harris, E. vaccaria var. erucoides Halford &

W.K.Harris, E. verrucitesta Halford & W.K.Harris, E. vicina Halford & W.K.Harris, E. victoriensis

Halford & W.K.Harris

D.A. Halford & W.K. Harris, cl- Queensland Herbarium, Department of Science, Information

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

Queensland 4066, Australia

Accepted for publication 28 September 2012

442

Introduction

In its broadest sense Euphorbia L. is one of

the largest genera of angiosperms in the world

comprising at least 2100 species and has an

almost cosmopolitan distribution (Riina &

Berry 2012). The morphological element that

unifies this diverse genus is the cyathium

which consists of a central pistillate flower

surrounded by five clusters of staminate

flowers within a cup-like involucre with

a ring of lobes and glands on the rim. This

unique reproductive structure resembles a

bisexual flower and is generally interpreted

as a complex pseudanthium. Recent studies

present evidence that the cyathium could

involve the expression of floral as well as

inflorescence developmental pathways

(Prenner & Rudall 2007; Prenner et al. 2011).

Euphorbia s.l. includes an astonishing

array of vegetative forms from small leafy

annuals to large cactus-like trees (Horn et

al. 2012). Given the size and morphological

diversity of the genus it is not surprising

that the taxonomy of this genus is complex

and often considered controversial. This

is reflected in a long history of attempts to

dismember the genus into numerous more

homogeneous groups (e.g. Chamaesyce Gray,

Monadenium Pax, Pedilanthus Neck, ex Poit.,

Poinsettia Graham and Synadenium Boiss.).

The genus Chamaesyce is one of the more

easily recognizable segregates that is generally

considered to be a ‘natural’ taxon and more or

less well defined by a series of morphological,

physiological and developmental characters

(Dressier 1961; Webster et al. 1975; Hassall

1976; Evans & Kinghorn 1977; Koutnik 1987;

Hayden 1988; Benedi & Orell 1992; Yang &

Berry 2011). It is characterised by having a

sympodial growth pattern, predominance of

C 4 photosynthesis with the associated Kranz

leaf anatomy, leaves opposite (predominantly

distichous) and mostly asymmetrical at the

base with interpetiolar stipules and seeds

ecarunculate. The genus is widely distributed

in subtropical and tropical regions throughout

the world, with a particular centre of diversity

in North America and is estimated to contain

c. 350 species (Hassall 1977; Yang & Berry

2011). As in other regions of the world the

Austrobaileya 8(4): 441-600 (2012)

recognition of Chamaesyce in Australia has

had limited acceptance (James & Harden

1990; Forster & Henderson 1995).

The current trends are tending to favour

the adoption of a broad concept of the genus

Euphorbia as recent phylogenetic analyses

involving morphological characters (Park &

Elisens 2000) and molecular sequence data

(Steinmann & Porter 2002; Bruyns et al.

2006; Park & Jansen 2007; Zimmermann

et al. 2010; Bruyns et al. 2011; Horn et al.

2012) indicate that the previously recognised

segregated genera are nested well within

Euphorbia. The six molecular phylogenies

resolved Euphorbia s.l. into four major, well

supported clades (A-D) which are formally

recognised as subgenera (A = E. subg.

Rhizanthium (Boiss.) Wheeler; B = E. subg.

Esula Pers.; C = E. subg. Euphorbia ; D = E.

subg. Chamaesyce Raf.) (Bruyns et al. 2006,

2011; Horn et al. 2012). Yang et al. (2012) have

recircumscribed E. subg. Chamaesyce and

presented a formal classification, recognizing

15 sections. The previous segregated genus

Chamaesyce is deeply imbedded within E.

subg. Chamaesyce and formally recognised

at the sectional level namely E. sect.

Anisophyllum Roeper (Yang et al. 2012).

Euphorbia section Anisophyllum in

Australia

In De Candolle’s Prodromus , Boissier (1862)

enumerated 15 species of Euphorbia sect.

Anisophyllum that are relevant to the Australian

flora. In his account of Euphorbia in Flora

Australiensis , Bentham (1873) recognised

17 species within E. sect. Anisophyllum , one

species E. hirta L. (as E. pilulifera L.) appears

to be an early naturalisation. Since Bentham’s

work numerous species (18) and infraspecific

(13) taxa based on Australian material have

been described by various botanists (e.g.

Fitzgerald 1918; Domin 1927; Thomson 1992).

The taxonomy of the Australian species of E.

sect. Anisophyllum has not been reviewed on

a national level since 1977. Hassall (1977)

as part of his doctoral studies on the tribe

Euphorbieae in Australia, recognised 22

native species as well as 14 infraspecific taxa.

He supported the recognition of Chamaesyce

(E. sect. Anisophyllum ) as distinct from

Euphorbia and transferred 15 names to

443

Halford & Harris, Euphorbia section Anisophyllum in Australia

Chamaesyce (Hassall 1976). He proposed

a number of new taxa (species, subspecies

and varieties) and lectotypifications but these

were never validly published and the majority

of his taxonomy on this group has remained

unpublished.

The first author found that while preparing

an account of Euphorbia for the Flora of

Australia that it was apparent there were

considerable taxonomic problems in this

section of the genus. Many more collections

and information relevant to the taxa concerned

have become available since Hassalfs work. A

number of the species were previously poorly

known and there was confusion over the

application of some names. We have found,

as Bentham and Hassall have before us, that

the specific and infraspecific boundaries in

the Australian E. sect. Anisophyllum are not

easily defined. Although some morphological

features such as seeds are reasonably

diagnostic for individual species, variation

across most characters is extensive. However,

it is felt that separation of these entities at

the specific rank is more desirable than

having a few ‘mega-species’ with numerous

infraspecific units. While the present study

resolves many taxonomic issues in the

Australian species of E. sect. Anisophyllum ,

still other problems remain to be examined in

more detail.

We have recognised 58 species of which

51 are native (48 endemic) and seven are

naturalised. Three native species {Euphorbia

bifida Hook. & Arn., E. obliqua Endl. and E.

pallens Dillwyn) extend to South East Asia

and the south west Pacific. The species of

Euphorbia sect. Anisophyllum are widespread

in Australia and are present in all States

although only a handful of species occur in

Victoria (Vic) and Tasmania has none. Most

species occur in the northern and central part

of the continent where there are six main

centres of species’ richness: Hamersley -

Pilbara (Western Australia [WA]), Kimberley

- Victoria River (WA & Northern Territory

[NT]), north western Highlands (Queensland

[Qld]), Einasleigh - Desert Uplands (Qld),

MacDonnell Ranges - Finke (NT & South

Australia [SA]) and northern Flinders Range

(SA).

Cytology

The cytology findings reported by Hassall

(1976, 1977) have been re-interpreted in light

of the present study. The voucher collections

(lodged at BRI) from that study have been re¬

determined in line with the taxa recognised

here (Appendix 1). Hassall recorded two base

chromosome numbers for the Australian taxa,

n = 8 & 11 as well as an apparent tetraploid {n

= 22) race of E. dallachyana Baill.

Groupings of species within E. sect.

Anisophyllum in Australia

Boissier (1862) arranged the then known

species in E. sect. Anisophyllum into eight

subsections. The Australian taxa that he

dealt with were placed into four subsections:

E. subsect. Chamaesyce Boiss. (E.

armstrongiana Boiss, E. australis Boiss., E.

drummondii Boiss., E. erythrantha F.Muell.

[= Euphorbia australis var. erythrantha

(F.Muell.) Benth., in this revision], E.

minutifolia Boiss.[= E. dallachyana , in this

revision]); E. subsect. Elegantes Boiss. (E.

schhizolepis); E. subsect. Hypericifoliae

Boiss. (E. baueri , unplaced in this revision,

E. bifida , E. macgillivrayi (= E. bifida , in this

revision), E. micradenia (= E. bifida , in this

revision), E. mitchelliana Boiss., E. muellerii

Boiss.); E. subsect. Sclerophyllae Boiss.

{E. atoto (= E. pallens , in this revision), E.

myrtoides Boiss., E. obliqua). Bentham (1873)

did not recognise any subdivisions within E.

sect. Anisophyllum and made no comment

regarding Boissier’s groupings.

Yang & Berry (2011) found Boissier’s

classification to be of little value in

designating monophyletic groups within

Euphorbia sect. Anisophyllum. Their data

supported three main lineages within the

group namely the “Acuta”, “Peplis” and

“Hypericifolia” clades. All the Australian

species sampled (E. australis , E coghlanii

F. M.Bailey, E. dallachyana , E. carissoides

F.M.Bailey, E. schultzii Benth., E. schizolepis

F.Muell. ex Boiss., E. sp. nov. ‘Australia’

= E. occulta Halford & W.K.Harris) were

found to group within the large cosmopolitan

distributed Hypericifolia clade (= E. subsect.

Hypericifoliae Boiss., Yang et al. (2012).

444

Hassall (1977) evaluated the relationships

within the Australian species of Chamaesyce

(= E. sect. Anisophyllum ) using numerical

techniques involving morphological and

cytological data. He noted a good correlation

in the data and organised the Australian

species into three groups that essentially

corresponded to karyotype variation which

parallelled, to some degree, patterns in

morphology in the Australian species. Our

observations generally support Hassalfs

delimitation of the Australian species into

three groups. These groups are:

Group 1 (E. sect. Chamaesyce [Hassall

1977]): This, represented by 35 species,

is the largest and most widespread of the

three groups. The species are found mostly

in inland habitats. The plants are generally

of small stature (mostly <40 cm) and often

with prostrate or decumbent stems; leaves

are small (2-25 x 1-9 mm); cyathia solitary

at the nodes although often clustered on

shortened leafy lateral branchlets; seeds small

(0.7-2.2 mm long), generally tetraquetrous in

cross-section, with surfaces variously finely

sculptured or smooth, mostly mucilaginous

when moistened. Fourteen of the species have

been recorded having chromosome number n

=11 and one species with a tetraploid race n =

22 (Appendix 1).

Within this group there are a number

of subgroupings that we perceive to be

morphologically similar species. These are:

subgroup australis: E. accedens

Halford & W.K.Harris, E. australis , E.

careyi F.Muell., E. centralis B.G.Thomson,

E. macdonaldii Halford & W.K.Harris, E.

occulta , E. schutlzii, E. thelephora Halford &

W.K.Harris, E. vaccaria Baill.

subgroup drummondii: E. drummondii,

E. dallachyana , E. fitzroyensis Halford

& W.K.Harris, E. flindersica Halford

& W.K.Harris, E. gregoriensis Halford

& W.K.Harris, E. hassallii Halford &

W.K.Harris, E. philochalix Halford &

W.K.Harris, E. porcata Halford & W.K.Harris,

E. verrucitesta Halford & W.K.Harris.

subgroup ferdinandi: E. crassimarginata

Halford & W.K.Harris, E. ferdinandi

Austrobaileya 8(4): 441-600 (2012)

F.Muell., E. papillata Halford & W.K.Harris,

E. multifaria Halford & W.K.Harris.

subgroup ophiolitica: E. laciniloba

Halford & W.K.Harris, E. maconochieana

B.G.Thomson, E. ophiolitica (P.I.Forst.)

Y.Yang, E. papillifolia Halford & W.K.Harris.

subgroup petala: E. alhrechtii Halford

& W.K.Harris, E. cinerea W.Fitzg., E. petala

Ewart & L.R.Kerr.

subgroup wheerleri: E. myrtoides, E.

sharkoensis Baill., E. victoriensis Halford &

W.K.Harris, E. wheeleri Baill.

Unplaced species: E. inappendiculata Domin,

E. armstrongiana.

Group 2 (E. sect. Sclerophyllae [Hassall

1977]): This group is represented by 13

species. The species are in coastal and inland

habitats in mostly northern Australia. The

plants are generally of larger stature (up to 80

cm) and often with decumbent or erect stems;

leaves are generally larger (5-80 x 1-22

mm); cyathia solitary at the nodes or in lax to

congested, leafy to bracteose dichasia; seeds

generally larger (1-2.2 mm long), tetragonous,

trigonous, biconvex or suborbicular in cross-

section, with surfaces variously finely

sculptured or smooth, non-mucilaginous

or mucilaginous when moistened. Five

of the species have been recorded having

chromosome number n =8 (Appendix 1).

The following groupings are perceived to be

morphologically similar species.

subgroup coghlanii: E. biconvexa

Domin, E. clementii Domin, E. coghlanii ,

E. psilosperma Halford & W.K.Harris, E.

trigonosperma Halford & W.K.Harris.

subgroup pallens: E. litticola Halford

& W.K.Harris, E. muelleri , E. obliqua, E.

pallens , E. psammogeton P.S.Green.

subgroup bifida: E. bifida,E. mitchelliana ,

E. vicina Halford & W.K.Harris.

Group 3 (E. sect. ‘Elegantes’ [Hassall 1977]):

This group is represented by 3 species. The

species occur in inland habitats of northern

Australia. The plants are generally of larger

stature (up to 120 cm) and mostly decumbent to

erect stems; leaves are ± large (5-36 x 3-19 mm);

445

Halford & Harris, Euphorbia section Anisophyllum in Australia

cyathia solitary at the nodes; seeds large (1.9-

3.2 mm long), tetragonous or tetraquetrous in

cross-section, with surfaces variously coarsely

sculptured or smooth, and mucilaginous when

moistened. The chromosome number for these

species is unknown.

Included species: E. carissoides, E.

kimberleyensis B.G.Thomson, E. schizolepis.

The groups and subgroups have not been

given formal names for it is more appropriate

that this be done within the context of a

broader investigation. In the following

‘Taxonomy’ section we have arranged the

species alphabetically.

Materials and methods

This study has been primarily herbarium

based, together with field work undertaken

by the authors. Herbarium collections on

loan to BRI from herbaria AD, DNA, HO,

MEL, NSW, NT and PERTH were studied

and annotated, and selected material from G,

K, P and W was also seen. The Herbarium

acronyms follow Holmgren et al. (1990). All

specimens cited in this revision have been

examined by one or both of the authors, unless

indicated otherwise by ‘n.v\ Those specimens

seen only as a digital image either online

(http://www.jstor.org) or images sent from

other herbaria in lieu of sending the specimen

are indicated by ‘image seen’ while those seen

only as a microfiche image are indicated by the

citation of the relevant IDC microfiche. Unless

otherwise stated the species in this treatment

are endemic to Australia. Species considered

naturalised in Australia are indicated by an

asterisk ‘*’ preceding the species name in the

‘key to species’ and in the main species entry

in the ‘Taxonomy’ section. The descriptions

of the species’ naturalised in Australia are

based on the variation observed in specimens

collected from Australia and therefore may

not reflect the full variation observed in the

species over its native range.

The revision is based on an assessment of

morphological characters of about 5300 dried

herbarium collections. We used the alpha

taxonomic techniques of studying characters

of the specimens and repeatedly sorting the

specimens into groups until the most efficient

characters for sorting had been determined.

Measurements in tenths of a millimetre were

made with a dissecting microscope at 10 to 40

times magnification and eyepiece graticule.

Descriptions of colour of vegetative and

floral parts are either from the information on

herbarium labels or information recorded by

the authors during field studies. The ventral

surface of the seed is the side facing towards

the centre of the capsule before dehiscence;

the dorsal surface is the side facing away

from the centre of the capsule. Seed shape is

the outline of the seed when the dorsal surface

is viewed radially. 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. 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 was

obtained from herbarium labels. Locality data

on herbarium specimens were used to generate

the distribution maps using DIVA-GIS version

7.5.0.0.

Seed photographs were taken with a Nikon

DS-Fil microscope camera attached to a

stereomicroscope (LeicaMZ6). Single images

were combined with Helicon Focus version

5.2 (Helicon Soft, http://HeliconFocus.com)

to produce composite images to increase the

depth of field. A small number of mature seeds

of each taxa where tested for the production of

a mucilaginous layer by using the method of

Jordan & Hayden (1992). Seeds were hydrated

in water for 5 min, then examined under a

dissecting microscope for the presence of

mucilage.

Although the taxa studied flower and

fruit early in their development there are

many herbarium collections that we have

been unable to identify to species because

the collections were of insufficient quality or

lacking sufficient floral or fruiting parts (e.g.

seeds).

Common abbreviations used in the

specimen citations include N.R (National

Park), N.R. (Nature Reserve) and S.F. (State

Forest).

446

Taxonomy

Euphorbia sect. Anisophyllum Roeper in

J.E.Duby, Bot. Gall. 2 nd edn, 1: 412 (1828).

Type: Euphorbia peplis L. (lecto: fide

Wheeler [1941: 111]).

Chamaesyce Gray, Nat. Arr. Brit. PI. 2: 260

(1821). Type: Chamaesyce maritima Gray,

nom. illeg., = C. peplis (L.) Prokh. (lecto: fide

Millspaugh [1909: 300] n.v., fromYang et al.

[ 2012 ]).

Monoecious (rarely dioecious) annuals or

herbaceous perennials, few to many stems

arising from crown of taproot, milky latex

in all parts, growth sympodial, the primary

axis exhibits no further development above

the epicotyl, growth continues through

development of lateral or secondary axes.

Stipules present, persistent, connate forming

Austrobaileya 8(4): 441-600 (2012)

interpetiolar sheath, entire or deeply bipartite.

Leaves opposite, distichous, petiolate,

usually with Kranz anatomy, the blades

entire to variously toothed, base is usually

markedly asymmetrical. Cyathia terminal

but appearing axillary, solitary at the nodes,

or in lax to congested, leafy to bracteose

dichasia. Involucre with 4 (rarely 5 or 6)

glands; glands with or without petaloid

appendages. Staminate flowers few to many,

pedicellate, naked and monandrous. Pistillate

flowers pedicellate, naked, hypogynous disc

entire or fimbriate; ovary 3-locular with

one pendant ovule in each locule; styles 3,

free or basally connate, bifid or entire. Fruit

capsular, 3-seeded, separating septicidally

into three 2-valved cocci leaving a persistent

columella. Seeds ecarunculate, smooth or

variously sculptured, mucilaginous or non-

mucilaginous when moistened.

Key to species of Euphorbia section Anisophyllum in Australia (distribution in states

indicated by acronyms)

1 .

2 .

6 .

Capsules with hairs.2

Capsules glabrous.28

Indumentum on stems consisting of two hairs types; white short ±

appressed crispate hairs to 0.5 mm long and yellow spreading segmented

hairs to 1.5 mm long; cyathia in dense capitate, terminal or axillary

cymose clusters.

Indumentum on stems consisting of one hair type or absent; cyathia

solitary, clustered on congested lateral branchlets, or in lax to congested

leafy terminal and axillary dichasial cymes.

Stems prostrate or rarely decumbent; cyathia in terminal cymose clusters

(NSW, Qld) 37. *E. ophthalmica

Stems ascending to erect; cyathia in terminal andaxillary cymose clusters

(NSW, NT, Qld, WA).21. *E. hirta

Capsules with hairs unevenly distributed over surface either restricted to

keels or towards the base of the capsule.5

Capsules with hairs evenly distributed over surface.10

Seeds >1.9 mm long (WA).24. E. kimberleyensis

Seeds <1.9 mm long.6

Capsules with hairs mostly towards the base; hairs >0.8 mm long (NT,

WA).54. E. vaccaria

Capsules with hairs restricted to keels; hairs mostly <0.8 mm long .7

Halford & Harris, Euphorbia section Anisophyllum in Australia 447

7 Capsules <1.5 mm across; involucral gland appendages <0.2 mm

long; staminate flowers per cyathium <10; stems prostrate or weakly

ascending, glabrous on lower surface (NSW, NT, Qld, WA).44. *E. prostrata

7. Capsules >1.5 mm across or if less then involucral gland appendages

>0.2 mm long, staminate flowers per cyathium >10 and stems erect

or if prostrate or ascending then evenly hairy.8

8 Capsules distinctly wider than long, 2-3 mm across, deeply 3-lobulate

(NT, Qld, WA).48. E. schultzii

8. Capsules ± as long as wide, 1.5-2.3 mm across, shallowly 3-lobulate.9

9 Leaf blades 7-70 mm long, glabrous or with a few scattered hairs, with

margins entire or finely serrulate; seeds tetragonous or trigonous in

transverse section; seed surfaces with 3-7 prominent narrow rounded

transverse ridges (NT, Qld,WA).30. E. mitchelliana

9. Leaf blades 3-7 mm long, sparsely to densely hairy, with margins often

coarsely serrate; seeds tetraquetrous in transverse section; seed surfaces

with low faint to distinct irregular ridges (NT, Qld, SA, WA) .9. E. centralis

10 Seeds >1.9 mm long.11

10. Seeds <1.9 mm long.12

11 Stems decumbent to erect, moderately to densely hairy with hairs to

2 mm; involucral gland appendages dentate to deeply laciniate, hairy

abaxially or if glabrous then styles entire (NT, WA).47. E. schizolepis

11. Stems usually prostrate, sparsely to densely hairy with hairs to 0.3 mm

long; involucral gland appendages entire or shallowly lobed, glabrous;

styles bifid (NT).31. E. muelleri

12 Gland appendages erect concealing involucral glands; involucral lobes

obovate, 0.8-1 mm long (Qld).35. E. occulta

12. Gland appendages spreading radially not concealing involucral gland or

absent; involucral lobes triangular to broad-triangular or subulate, 0.2-0.8 mm long .... 13

13 Styles entire or scarcely notched.14

13. Styles bifid for at least 1/3 of their length.18

14 Seeds <1 mm long (NT, Qld).27. E. macdonaldii

14. Seeds >1 mm long.15

15 Involucral glands patelliform, either ± planar or shallowly concave; gland

appendages deeply laciniate with narrow attenuate teeth, >0.4 mm long

(NT, Qld, SA, WA).9. E. centralis

15. Involucral glands cupuliform, deeply sunken in centre with distinct

thickened rim; gland appendages entire or shallowly irregularly lobed

or bluntly toothed, up to 0.6 mm long.16

16 Capsules distinctly longer than wide (NSW, NT, Qld, SA).51. E. thelephora

16. Capsules ± as long as wide or distinctly wider than long.17

17 Capsules distinctly wider than long; leaf margins serrate; dorsal faces of

seeds with 3-5 prominent ± transverse or irregular, broad ridges (NT,

Qld, WA).48. E. schultzii

17. Capsules ± as long as wide; leaf margins entire or minutely toothed

distally; dorsal faces of seeds with 5-7 prominent ± transverse, narrow

ridges (NT, Qld).1. E. accedens

448 Austrobaileya 8(4): 441-600 (2012)

18 Pedicel of mature capsules <1 mm long; capsules not completely exserted

from involucre at maturity causing the involucre to split (NT, Qld,

WA) 52. *E. thymifolia

18. Pedicel of mature capsules >1 mm long; capsules completely exserted

from involucre at maturity.19

19 Cyathia in lax terminal dichasia or monochasia or in congested axillary

bracteose dichasia.20

19. Cyathia solitary at nodes or in congested short lateral leafy branches

(leaves on lateral branches usually smaller in size than main branches

but not reduced to bracts).22

20 Capsules minutely papillose; gland appendages absent or if present then

<0.1 mm long (WA).56. E. vicina

20. Capsules smooth; gland appendages present, >0.1 mm long.21

21 Cyathia in axillary or terminal congested dichasial cymes (NSW, NT,

Qld, WA).6. E. bifida

21. Cyathia in terminal lax dichasial or monochasial cymes (NT, Qld

WA).30. E. mitchelliana

22 Hairs on capsules appressed; seeds <1 mm long; annual herbs to 10 cm

high with prostrate or weakly ascending stems; leaf blades green often

with elongate purple spot centrally; stipules 1-2.3 mm long; hairs on

stem up to 0.8 mm long (all mainland States).29. *E. maculata

22. Hairs on capsules spreading or if appressed then either seeds >1 mm

long or herbaceous perennials with ascending to erect stems or leaf

blades yellowish green or stipules <1 mm long or hairs on

stems to 0.2 mm long.23

23 Capsules depressed ovate or transversely elliptic in lateral view,

distinctly wider than long, acutely keeled; leaf blades ovate, oblong or

obovate, >1.5 times longer than wide (NT, Qld, WA).48. E. schultzii

23. Capsules broad-ovate to very broad-ovate or broad-elliptic in lateral view,

as long as wide or if depressed ovate then only slightly wider than long,

obtusely keeled and leaf blades elliptic to broad elliptic and <1.5 times

longer than wide.24

24 Gland appendages deeply laciniate with narrow attenuate teeth (NT,

Qld, SA, WA).9. E. centralis

24. Gland appendages entire, bluntly lobed or acutely toothed.25

25 Gland appendages white rarely pale pink, entire; stems prostrate; capsules

conspicuously papillose and only sparsely hairy with hairs up to 0.1

mm long; leaf blades strongly asymmetric at base (NT, WA) ... 28. E. maconochieana

25. Gland appendages red, yellow or if white or pink then irregularly

lobed or toothed; stems prostrate, ascending to erect; capsules

smooth or if papillose then only minutely so and sparsely to densely

hairy with hairs 0.1-0.5 mm long; leaf blades weakly to strongly

asymmetric at base.26

26 Stems longitudinal ribbed (Qld).36. E. ophiolitica

26. Stems not longitudinal ribbed.27

Halford & Harris, Euphorbia section Anisophyllum in Australia 449

27 Cyathiamostly unisexual; hairs on stems <0.2 mm long; involucral glands

± flat, 0.3-0.5 mm long by 0.4-0.8 mm wide (WA).7. E. careyi

27. Cyathia bisexual; hairs on stems >0.2 mm long or if equal to

or shorter then involucral glands concave or with shallow central

pit, <0.3 mm long and <0.4 mm wide (all mainland States except Vic) . . . 4. E. australis

28 Dorsal faces of seeds smooth, without ornamentation or surface

irregularities; seeds tetragonous, trigonal, biconvex or suborbicular in

transverse section.29

28. Dorsal faces of seeds sculptured or variously ornamented sometimes only

obscurely so or if smooth then seed tetraquetrous in transverse section.37

29 Involucral glands, sessile, pressed flat against the outer surface of the

involucre; gland appendages absent; involucral lobes oblong to obovate,

deeply laciniate distally, >1 mm long (WA).11. E. clementii

29. Involucral glands stipitate, spreading; gland appendages usually present

sometimes inconspicuous; involucral lobes triangular to subulate,

mostly <1 mm long.30

30 Capsules >3.5 long; seeds >1.9 mm long; styles >1.5 mm long;

herbaceous woody perennials to 120 cm high (Qld).8. E. carissoides

30. Capsules <3.5 mm long; seeds <1.9 mm long; styles mostly <1.5 mm

long; herbaceous perennials mostly less than 80 cm high.31

31 Seeds biconvex in transverse section (NT, Qld, SA, WA).5. E. biconvexa

31. Seeds trigonous or suborbicular or tetragonous in transverse section.32

32 Capsules >3 mm long; stipules >2 mm long (NT, Qld, WA).26. E. litticola

32. Capsules <3 mm long; stipules <2 mm long.33

33 Leaf blades <5 mm wide; styles entire or scarcely divided distally (NT,

WA).46. E. psilosperma

33. Leaf blades >5 mm wide or if less then styles bifid for one third or more

of their length.34

34 Stipules triangular to broad-triangular, pubescent on adaxial surface;

seeds not becoming mucilaginous when moistened; growing on sandy

coastal foreshores.35

34. Stipules subulate to narrow-triangular, glabrous on adaxial surface; seeds

mucilaginous when moistened; growing in inland areas on various soils

and coastal sands.36

35 Leaf blades <20 mm long and <10 mm wide; cyathia solitary in upper

axils (Qld).34. E. obliqua

35. Leaf blades >20 mm long and >10 mm wide; cyathia in congested dichasial

cymes (Qld).38. E. pallens

36 Seeds suborbicular in transverse section; plants most commonly growing

on clay soils (NT, Qld, WA).12. E. coghlanii

36. Seeds trigonous in transverse section; plants most commonly growing

on sandy soils (NT, Qld, WA).53. E. trigonosperma

37 Dorsal and ventral surfaces of seeds with 4-6 transverse grooves or

foveate to reticulate-foveate.38

37. Dorsal and ventral surfaces of seeds smooth, or faintly or distinctly

transversely or irregularly ridged.39

450

38.

39.

40.

41.

42.

43.

44.

45.

46.

47.

48.

Austrobaileya 8(4): 441-600 (2012)

Dorsal and ventral surfaces of seeds with 4-6 transverse grooves (Fig.

25B) (NT, WA).3. E. armstrongiana

Dorsal and ventral surfaces of seeds foveate to reticulate-foveate

(Figs 28L & M) (all mainland States except Vic).58. E. wheeleri

Seeds >1.9 mm long, >1.5 mm wide.40

Seeds <1.9 mm long or if >1.9 mm then <1.5 mm wide.41

Capsules papillose; surface of seeds with low rounded irregular ridges

with exotesta distinctly thicker on the ridges; gland appendages dentate

to laciniate, (0.5) 1.3-2.5 mm long (WA).24. E. kimberleyensis

Capsules smooth; surface of seeds with faint narrow irregular ridges

with exotesta of even thickness over surface; gland appendages entire

or shallowly lobed, 0.6-1.5 mm long (NT).31. E. muelleri

Cyathia in lax to congested leafy terminal dichasia or monochasia or in

congested axillary bracteose dichasia.42

Cyathia solitary at the nodes, sometimes clustered on short leafy lateral

branches.46

Seeds orbicular or very broad-ovate in outline, >1.5 mm long, >1.2 mm

wide, ± suborbicular or slightly trigonal in transverse section, surfaces

faintly irregularly ridged; capsules >2.5 mm long and >3 mm wide

(NSW, Qld).45. E. psammogeton

Seeds ovate to elliptic or broad-ovate in outline, <1.5 mm long or if 1.5 mm

long then <1.2 mm wide, tetragonous in transverse section or if

trigonal in transverse section then surface distinctly ridged; capsules

<2.5 mm long and <3 mm wide.43

Capsules minutely papillose; gland appendages absent or if present <0.1

mm long (WA).56. E. vicina

Capsules smooth; gland appendages present, >0.1 mm long.44

Stipules broad-triangular, 0.3-0.6 mm long, with margins lacerate, teeth

gland-tipped (NSW, NT, Qld, WA).22. *E. hyssopifolia

Stipules deeply bipartite, lobes subulate to narrow-triangular, 0.6-2 mm

long, with margins entire or laciniate, sometimes with teeth gland-

tipped .45

Cyathia in terminal lax dichasial or monochasial cymes, usually above

subtending leaves (NT, Qld, WA).30. E. mitchelliana

Cyathia in congested dichasial cymes, mostly axillary on short

lateral branchlets, usually shorter than subtending leaf (NSW, NT, Qld,

WA) 6. E. bifida

Stems hairy, sometimes only proximally.47

Stems glabrous.55

Styles hairy, bifid; capsules smooth, slightly wider than long (Qld). . .40. E. papillifolia

Styles glabrous or if with a few scattered hairs then style entire,

and capsules papillose and distinctly longer than wide.48

Styles entire, 0.3-1.1 mm long; capsules widest at the middle (NSW, NT,

Qld, SA).51. E. thelephora

Styles bifid or if entire then 0.2-0.3 mm long and capsules usually wider

below the middle

Halford & Harris, Euphorbia section Anisophyllum in Australia 451

49 Involucral glands cupuliform, deeply sunken in centre with distinct rim.50

49. Involucral glands patelliform, either ± planar or convave with tangential

trough but not deeply sunken in centre with distinct rim.52

50 Capsules distinctly wider than long, 1.5-1.8 x 2-2.7 mm, deeply

3-lobulate (NT, Qld, WA).48. E. schultzii

50. Capsules only slightly wider than long, 1.4-2.1 * 1.5-2 mm, shallowly

3-lobulate.51

51 Exotesta ± of even thickness; gland appendages <0.1 mm long or if longer

then usually deeply lobed; leaves usually >3 times as long as wide (NT,

Qld, WA).20. E. hassallii

51. Exotesta of uneven thickness, always thicker on ridges; gland

appendages >0.1 mm long, entire; leaves <3 times as long as wide

(NT).19. E. gregoriensis

52 Gland appendages conspicuous, >0.3 mm long; staminate flowers

per cyathia mostly 10 or more; stems sparsely to densely hairy with

appressed to spreading hairs <0.5 mm long.

52. Gland appendages inconspicuous and <0.3 mm long or absent; staminate

flowers per cyathia <10; stems sparsely hairy with spreading hairs

>0.5 mm long.

53 Gland appendages reniform, entire; capsules papillose (visible at 10x

mag.); styles 0.7-1 mm long (NT, WA).28. E. maconochieana

53. Gland appendages obdeltoid to oblong with toothed margin; capsules

smooth; styles 0.4-0.6 mm long (Qld).25. E. laciniloba

54 Styles bifid; involucral glands 0.05-0.15 x 0.1-0.2 mm; gland appendages

<0.1 mm long (NSW, NT, Qld, SA, WA).23. E. inappendiculata

54. Styles entire; involucral glands 0.2-0.3 x 0.4-0.5 mm; gland appendages

>0.1 mm long (NT, Qld).7. E. macdonaldii

55 Seeds faces ± smooth.56

55. Seeds faces sculptured or variously ornamented sometimes only obscurely so .... 65

56 Capsules widest below the middle; seeds <1 mm long .57

56. Capsules widest at the middle or if widest below middle then seeds >1 mm long. ... 58

57 Involucral glands cupuliform; leaf blades oblong or obovate, >1.5 times as

long as wide, margins minutely toothed distally (NT, WA).10. E. cinerea

57. Involucral glands patelliform; leaf blades oblong, elliptic to broad

elliptic, <1.5 times as long as wide, margins serrulate (NT, Qld) ... 27. E. macdonaldii

58 Involucral glands cupuliform, surface deeply sunken, often with thickened rim .... 59

58. Involucral glands patelliform, surface flat or concave.63

59 Capsules distinctly longer than wide.60

59. Capsules more or less as long as wide.61

60 Gland appendages >0.1 mm long; seeds >0.7 mm wide (radially) (NSW,

NT, Qld, SA) 40. E. papillata

60. Gland appendages absent or if present then <0.1 mm long or if

0.1 mm long then seeds <0.7 mm wide (radially) (NSW, NT, Qld, SA,

WA).16. E. ferdinandi

452

Austrobaileya 8(4): 441-600 (2012)

61 Stems and capsules conspicuously papillose (visible at 10x mag.) (NSW,

NT, Qld, SA) 40. E. papillata

61. Stems smooth; capsules smooth or inconspicuously papillose (visible at

40x mag.).62

62 Leaf blades <2 times as long as wide, oblong, oblong-obovate to obovate,

2.8-7 x 1.6-3.8 mm; stems prostrate, rarely ascending to erect; stipules

0.4-1 mm long (NSW, Qld, SA, Vic, WA).32. E. multifaria

62. Leaf blades >2 times longer than wide, narrow oblong, 6-11 x 2-3.6 mm;

stems ascending to erect, rarely prostrate; stipules 0.8-1.5 mm long (NT,

Qld).39. E. papillata

63 Styles entire (NSW, NT, Qld, SA).51. E. thelephora

63. Styles bifid 1/3—1/2 of their length.64

64 Plants monoecious; gland appendages deeply dentate or divided to base,

0.3-0.4 mm long; capsules minutely papillose (visible at 40x mag.);

growing in sandy clay soils among rocky outcrops and on gravelly hill

slopes (SA).18. E. flindersica

64. Plants dioecious rarely monoecious; gland appendages entire, erose or

shallowly irregularly lobed, mostly 0.4-0.9 mm long rarely shorter;

capsules smooth or rarely minutely papillose; growing in red clay

loams to heavy black clays, rarely sandy soils on plains or undulating

to hilly terrain (NSW, Qld).40. E. papillifolia

65 Capsules >1.2 times wide as long, transversely broad-elliptic in lateral

view, deeply 3-lobulate; involucral glands cupuliform; gland appendages

entire (NT, Qld, WA).48. E. schultzii

65. Capsules <1.2 times wide as long, or if >1.2 times as long as wide then

either capsules broad-ovate to depressed ovate in lateral view and

shallowly 3-lobulate or involucral glands patelliform or gland

appendages toothed or deeply irregularly lobed.66

66 Seeds <1 mm long.67

66. Seeds >1 mm long.71

67 Styles entire.68

67. Styles bifid for 1/3—1/2 of their length.69

68 Involucral glands cupuliform; leaf blades >1.5 times as long as wide,

margins minutely toothed distally (NT, WA).10. E. cinerea

68. Involucral glands patelliform; leaf blades <1.5 times as long as

wide, margins serrulate (NT, Qld).27. E. macdonaldii

69 Stems rooting at nodes; stipules 0.5-0.6 mm long, broadly triangular, not

bilobed (NSW, Qld, SA).49. *E. serpens

69. Stems not rooting at nodes; stipules 0.9-1.2 mm long, bilobed, lobes triangular . ... 70

70 Dorsal faces of seed concave, with prominent irregular rounded ridges;

gland appendages entire, crenulate or shallowly irregularly lobed (NT,

WA) 42. E. philochalix

70. Dorsal faces of seed planar, with faint rounded, irregularly ridges; gland

appendages toothed or deeply lobed (WA).50. E. sharkoensis

Halford & Harris, Euphorbia section Anisophyllum in Australia 453

71 Involucral glands patelliform, flat or concave with tangential trough, rim

of gland not thickened.72

71. Involucral glands cupuliform, deeply sunken in the centre, usually with

thickened rim.88

72.

73.

74.

75.

76.

77.

78.

79.

80.

81.

82 .

Styles entire or scarcely bifid; capsules broader towards the base; gland

appendages >0.3 mm long (NT, Qld).41. E. petala

Styles bifid for 1/3—1/2 of their length or if entire then capsules

broader towards the middle or gland appendages absent or <0.3 mm

long.73

Stipules on at least one side of stem triangular to broadly triangular, with

toothed margins but not deeply bipartite.74

Stipules deeply bipartite, with lobes subulate to triangular.76

Gland appendages <0.4 mm long and <0.8 mm wide; staminate flowers

per cyathium <10; hypogynous disc often laciniate (all mainland

States).14. E. dallachyana

Gland appendages >0.4 mm long and >0.8 mm wide; staminate flowers

per cyathium mostly >10; hypogynous disc entire.75

Leaf blades oblong to oblong-elliptic, >1.5 times as long as wide; seeds

<1.5 mm long (Qld).25. E. laciniloba

Leaf blades elliptic to broad-elliptic, <1.5 times as long as wide; seeds

>1.5 mm long (Qld).36. E. ophiolitica

Gland appendages <0.2 mm long or absent.77

Gland appendages >0.2 mm long.83

Capsules >2 mmlong; stems ascendingto erect, rarely prostrate; staminate

flowers per cyathium 10-30; styles 0.5-0.9 mm long (WA).33. E. myrtoides

Capsules <2 mm long; stems prostrate rarely erect; staminate flowers

per cyathium 5-15; styles 0.2-0.6 mm long.78

Styles entire, 0.2-0.3 mm long (NSW, Qld, SA, Vic, WA).32. E. multifaria

Styles bifid for 1/3—1/2 of their length, 0.3-0.6 mm long.79

Leaves minutely papillose (visible at 40x mag.).80

Leaves ± smooth.81

Involucral glands well developed, 0.1-0.3 mm long and 0.3-0.5 mm wide;

gland appendages >0.1 mm long (NSW, Qld).40. E. papillifolia

Involucral glands poorly formed up to 0.1 mm long and 0.1-0.2 mm

wide; gland appendages absent or <0.1 mm long (NSW, NT, Qld, SA,

WA).23. E. inappendiculata

Stipules >1 mm long; involucral glands up to 0.1 mm long (NSW, NT, Qld,

SA, WA).23. E. inappendiculata

Stipules <1 mm long; involucral glands >0.1 mm long.82

Dorsal faces of seeds undulate or obscurely ridged; seeds 1-1.5 mm long,

growing in sandy soils on rocky limestone foreshores and coastal sand

dunes (north-western WA).50. E. sharkoensis

Dorsal faces of seeds distinctly ridged; seeds 1.5-1.7 mm long; growing

on inland sand dunes in the Great Victoria Desert (NT, SA,WA) . . . .57. E. victoriensis

454

Austrobaileya 8(4): 441-600 (2012)

83 Capsulesminutelypapillose(visibleat40x mag.); stems notlongitudinally

ridged (SA).18. E. flindersica

83. Capsules smooth, or if minutely papillose then stems longitudinally

ridged when young.84

84 Staminate flowers <10 per cyathia or if 10 then capsules <2 mm long;

seeds <1.5 mm long.85

84. Staminate flowers >10 per cyathia or if 10 then capsules >2 mm long;

seeds >1.5 mm long.86

85 Leaves ± smooth; growing in sandy soils on rocky limestone foreshores

and coastal sand dunes (north-western WA).50. E. sharkoensis

85. Leaves minutely papillose (visible at 40x mag.); growing in clay

soils (mostly derived from basalt) in central Queensland (NSW,

Qld).40. E. papillifolia

86 Leaf blades elliptic to broad elliptic; 1-1.5 times as long as wide (Qld). 36. E. ophiolitica

86. Leaf blades oblong, ovate, obovate or oblong-elliptic, 1.5-2.2 times as

long as wide.87

87 Capsules >2.2 mm long; involucral glands 0.3-0.6 mm long; plants to 40

cm tall, monoecious; stems spreading to erect rarely prostrate; leaves

bright green to yellow-green (WA).33. E. myrtoides

87. Capsules <2.2 mmlong; involucral glands 0.2-0.4 mm long; plants to 10 cm

tall, mostly dioecious; stems prostrate to spreading; leaves blue-green

(NSW, Qld) 40. E. papillifolia

88 Stipules on at least one side of stem broadly triangular, not deeply

bipartite; hypogynous disc mostly laciniate; stems prostrate, rarely

erect or ascending (all mainland States).14. E. dallachyana

88. Stipules subulate to triangular, deeply bipartite; hypogynous disc mostly

entire; stems ascending to erect or prostrate.89

89 Gland appendages >0.1 mm long.90

89. Gland appendages <0.1 mm long or absent.100

90 Capsules broadest towards the base.91

90. Capsules broadest towards the equator.94

91 Dorsal faces of seeds with medial irregular longitudinal ridge (NSW, NT,

Qld, SA, WA).43. E. porcata

91. Dorsal faces of seeds irregularly ridged but without medial longitudinal

ridge.92

92 Styles entire or scarcely bifid at the tip (NT, Qld).41. E. petala

92. Styles bifid for 1/3—1/2 of their length.93

93 Dorsal faces of seed with low faint irregular ridges; plants usually robust

herbaceous perennials to 30 cm tall; stems erect rarely prostrate; gland

appendages 0.2-0.4 mm long (NT, WA).2. E. albrechtii

93. Dorsal faces of seeds with prominent irregular ridges; plants

slender herbaceous perennials to 15 cm tall; stems prostrate

rarely ascending; gland appendages 0.1-0.3 mm long (NT,

WA) 42. E. philochalix

455

Halford & Harris, Euphorbia section Anisophyllum in Australia

94 Leaf blades >3 times as long as wide; seeds 0.8-1.3 mm long, distinctly

or obscurely irregularly ridged; styles scarcely bifid to divided c. 1/3 of

their length; involucral glands 0.3-0.5 mm wide (NT, Qld, WA).20. E. hassallii

94. Leaf blades <3 times as long as wide or if up to 3.3 times

as long as wide then either seeds >1.3 mm long and obscurely

irregularly ridged, or styles entire, or involucral glands <0.2 mm

wide.95

95 Seeds <1.2 mm long or if >1.2 then dorsal faces of seeds with a medial

longitudinal ridge .96

95. Seeds >1.2 mm long; dorsal faces of seeds smooth or with irregular

ridges.98

96 Styles entire; capsules minutely papillose; stems longitudinally ridged

(NT, Qld) 39. E. papillata

96. Styles divided for 1/3—1/2 of their length; capsules smooth; stems not

longitudinally ridged.97

97 Dorsal faces of seeds irregularly ridged (NSW, NT, Qld, SA,

WA). 23. E. inappendiculata

97. Dorsal faces of seeds with a medial longitudinal ridge (NSW, NT, Qld, SA,

WA).43. E. porcata

98 Styles entire, 0.1-0.3 mm long; capsules papillose (NT, Qld).39. E. papillata

98. Styles divided for 1/3—1/2 of their length, 0.3-0.5 mm long; capsules

smooth or papillose.99

99 Erect annual to 40 cm tall; exotesta of uneven thickness, always thicker

on ridges; involucral lobes >0.4 mm long (NT).19. E. gregoriensis

99. Prostrate to procumbent perennial; exotesta of even thickness;

involucral lobes <0.4 mm long (Qld).40. E. papillifolia

100 Gland appendages absent; involucral glands well developed 0.2-0.3

mm long and 0.4-0.5 mm wide with conspicuous thickened rim; leaf

blades >3 times as long as wide and >7 mm long (NT, Qld). . . . 13. E. crassimarginata

100. Gland appendages usually present but inconspicuous or if absent then

either leaf blades <3 times as long as wide and mostly <7 mm long

or involucral glands poorly formed up to 0.1 mm long and 0.1-0.2 mm

wide; involucral glands with or without thickened rim. 101

101 Dorsal surface of seeds with 6-9 transverse ridges; exotesta of uneven

thickness, always thicker on ridges (WA).17. E. fitzroyensis

101. Dorsalfacesofseedswithirregularridges,ortuberculateoriftransversely

ridged then with 3-6 ridges on dorsal faces of seed; exotesta ± of even

thickness over seed surface. 102

102 Styles entire or scarcely notched at the apex, 0.2-0.3 mm long; dorsal

surface of seeds distinctly transversely or irregularly ridged; capsules

broadest towards the middle.103

102. Styles bifid for 1/3-2/3 of their length or if entire or scarcely notched

at the apex then either dorsal faces of seeds obscurely undulate or

capsules broadest towards the base; styles 0.2-0.6 mm long

105

456

Austrobaileya 8(4): 441-600 (2012)

103 Leaves <6 mm long; stems not longitudinally ribbed; dorsal face of

seeds irregularly ridged; exotesta of uneven thickness over seed surface

(thicker on ridges) (NSW, Qld, SA, Vic, WA).32. E. multifaria

103. Leaves >6 mm long or if <6 mm long then stems longitudinally ribbed,

dorsal faces of seeds with 3-6 transverse ridges and exotesta of ± even

thickness over surface.104

104 Leaves rounded at apex; dorsal faces of seeds with 3-6 transverse

ridges (NSW, NT, Qld, WA).15. E. drummondii

104. Leaves acute to obtuse at apex, or if rounded then dorsal faces

of seeds with irregular ridges (NT, Qld, WA).20. E. hassallii

105 Stipules subulate, 1-1.7 mm long; involucral glands poorly formed up to

0.1 mm long and 0.1-0.2 mm wide (NSW, NT, Qld, SA, WA). . . 23. E. inappendiculata

105. Stipules narrow-triangular to triangular or if subulate then <1.1 mm

long; involucral glands well formed, 0.1-0.2 mm long by 0.1-0.7 mm

wide.106

106 Largest leaves >8 mm long, mostly >3 times as long as wide (NT, Qld,

WA) 20. E. hassallii

106. Largest leaves <8 mm long, <3 times as long as wide.107

107 Dorsal faces of seeds with an irregularly medial longitudinal ridge

(NSW, NT, Qld, SA, WA).43. E. porcata

107. Dorsal faces of seeds undulate, faintly irregularly ridged or with

irregular wart-like protuberances.108

108 Dorsal faces of seeds with irregular wart-like protuberances (NSW,

SA, WA).55. E. verrucitesta

108. Dorsal faces of seeds undulate or only faintly irregularly ridged

(NSW, Qld, SA, Vic, WA).32. E. multifaria

1. Euphorbia accedensHalford&W.K.Harris,

species nova fieri potest, ut E. accedens

permisceatur cum E. australi var. hispidula

Halford & W.K.Harris et E. thelephora var.

rugosa Halford & W.K.Harris. Ab E. australi

var. hispidula seminibus longioribus 1.3-1.4

mm longis (in vicem 0.9-1.2 mm longis), 5-7

porcis ± transversis rotundatus superficierum

dorsalium seminis (in vicem undulatis vel

porcatis leviter irregulatim), stylis integris

vel vix bifidis (in vicem bifidis 1/3-2/3

longitudinis) necnon ab E. thelephora var.

rugosa seminibus brevis 1.3-1.4 mm longis

(in vicem 1.5-1.7 mm longis), capsulis in

latere visis lato-ellipticus ± longioribus quam

latioribus 1.6-2 mm longis 1.9-2.2 mm latis

(in vicem in latere visis 1.8-2.2 mm longis

1.4-1.8 mm latis distincte longioribus quam

latioribus) differt. Euphorbia accedens formis

E. schultzii Benth. similis sed capsulis in

latere visis ± longioribus quam latioribus 1.6-

2 mm longis 1.9-2.2 mm latis leviter 3-lobatis

(in vicem in latere visis depresse ovatis vel

transverse ellipticis distincte brevioribus

quam latioribus valde 3-lobatis), seminum

superficiebus porcis transversis rotundatis vel

porcatis irregulariter (in vicem 3-5 porcis)

marginibus folium integris distaliter dentatis

minute (in vicem marginibus folium pro

parte maxima dentatis grosse) differt. Typus:

Queensland. Burke District: Undilla Station,

NW of Mt Isa, 18 July 2008, R.Booth &

D.T.Kelman CAM36-7 (holo: BRI).

Monoecious, herbaceous perennial to 5

cm high, with few to many stems arising

from thickened woody taproot. Stems

erect or prostrate, sparingly to much

branched, smooth, with a moderately dense

indumentum; hairs spreading, ± straight, 0.2-

0.8 mm long, white. Interpetiolar stipules

subulate, 0.3-0.5 mm long, deeply bipartite,

with indumentum as for stems; margin entire

457

Halford & Harris, Euphorbia section Anisophyllum in Australia

or laciniate. Leaves: petiole 1-1.5 mm long,

smooth, with indumentum as for stems; blade

oblong or oblong-obovate, 5-12 mm long,

3-6 mm wide, 1.5-2.5 times longer than

wide; adaxial and abaxial surfaces pale blue-

green sometimes suffused with reddish tinge

or reddish tinge along margin, papillose, with

a moderately dense indumentum consisting

of spreading, ± straight hairs 0.2-0.8

mm long; base asymmetric with one side

cordate, the other obtuse to cuneate; margin

entire or minutely toothed distally; apex

obtuse to rounded sometimes with apiculate

tip. Cyathia solitary at the nodes, often

clustered on short leafy lateral branchlets

with subtending leaves slightly smaller than

primary stem leaves; peduncles 0.1-0.4 mm

long. Involucres cupuliform, 0.8-1.1 mm

long, 1-1.2 mm across; lobes 5, triangular,

0.4-0.5 mm long, margin fimbriate; glands 4,

stipitate, cupuliform, with distinct tangential

trough and thickened rim, transverse-oblong

in outline, 0.2-0.3 mm long, 0.3-0.5 mm

wide, yellow or pink; gland appendages

conspicuous, spreading radially, very broad-

obovate or transverse-oblong, 0.2-0.5 mm

long, 0.3-0.8 mm wide, white to pink,

glabrous, margin entire, erose or shallowly

lobed; bracteoles 0.8-0.9 mm long, adnate for

c. 1/3 of their length to involucre, free portion

divided into numerous subulate glabrous or

hirsute segments. Staminate flowers 4-7

per cyathium; pedicels 0.6-0.8 mm long;

staminal filaments c. 0.1 mm long. Pistillate

flowers: styles 0.4-0.5 mm long, spreading,

minutely papillose, sparsely hairy, entire or

scarcely bifid, with terete apices. Capsules

exserted from involucre on pedicel to 1.5 mm

long, broad-elliptic in lateral view, 1.6-2 mm

long, 1.9-2.2 mm across, shallowly 3-lobate

with keels acute, papillose, with a moderately

dense indumentum consisting of spreading

hairs 0.1-0.2 mm long; hypogynous disc

entire. Seeds ovate in outline, 1.3-1.4 mm

long, 0.7-0.8 mm tangentially, 0.7-0.8 mm

radially, tetraquetrous in cross section;

dorsal faces planar; ventral faces planar to

concave; all faces with 5-7 prominent ±

transverse, rounded ridges; exotesta thin,

of uneven thickness, distinctly thicker on

ridges, grey-white, microreticulate, becoming

mucilaginous when moistened; endotesta

dark brown. Fig. 1.

Additional selected specimens examined: Northern

Territory. Cattle Creek Stud, Wave Hill, Apr 1963,

Napier s.n. (DNA 10288); c. 75 km NW of Lake

Surprise, Aug 1991, Latz 12167 (NT); c. 66 km N [of]

Tennant Creek, upper Attack Creek catchment. May

1996, Albrecht 7628 & Latz (NT); near Alroy Downs

Station, May 1947, Blake 17889 (BRI, DNA); 24 km NW

of Wauchope Roadhouse, gas pipeline, Jun 2007, Latz

22770 (NT). Queensland. Burke District: Flora Downs

Station, c. 120 km NW of Mt Isa, May 2001, Bailey &

Kelman s.n. (BRI [AQ697493]); Lake Julius Road, 24

km to Mt Isa Copper stack at 58deg., Apr 1998, Fell

5346 (BRI); Fort Constantine Station, N of Cloncurry,

Jun 2003, Booth 3401 & Kelman (BRI); 90 km SSW of

Cloncurry on the Duchess Road, Feb 2001, Wannan 2071

& Jago (BRI). Gregory North District: 10 miles [c. 16

km] N of Duchess, May 1963, Gittins 731 (BRI, NSW).

Distribution and habitat : Euphorbia

accedens occurs from the Tanami Desert,

NT to Cloncurry, in north-western Qld (Map

1). It grows in eucalypt woodland or Triodia

grassland communities on loam or clay soils

on low limestone rises or hills, or sometimes

on alluvial plains.

Phenology: Flowers and fruits have been

collected in February, April to June, August

and November.

Notes: Euphorbia accedens may be confused

with E. australis var. hispidula Halford &

W.K.Harris and E. thelephora var. rugosa

Halford & W.K.Harris. It differs from E.

australis var. hispidula in having longer seeds

(1.3-1.4 mm long versus 0.9-1.2 mm long for

E. australis var. hispidula ), 5-7 prominent ±

transverse, rounded ridges on the dorsal seed

faces (versus undulate or faintly irregular

ridged for E. australis var. hispidula) and

entire or scarcely bifid styles (versus bifid for

1/3-2/3 of their length for E. australis var.

hispidula. It differs from E. thelephora var.

rugosa in having shorter seeds (1.3-1.4 mm

long versus 1.5-1.7 mm long for E. thelephora

var. rugosa ) and capsules broad-elliptic in

lateral view that are more or less as long as

wide, 1.6-2 mm long, 1.9-2.2 mm across

(versus elliptic in lateral view, 1.8-2.2 mm

long, 1.4-1.8 mm across and distinctly longer

than wide for E. thelephora var. rugosa).

458

Austrobaileya 8(4): 441-600 (2012)

Fig. 1. Euphorbia accedens. A. branchlet with cyathia xl2. B. leaf *4. C. stipules *16. D. cyathia with female flower

at anthesis x32. E. capsule with cyathia xl6. F. cyathial gland, adaxial view *32. G. capsule, top view xl6. H. capsule,

lateral view xl6. A, D from Gittins 731 (BRI); B, C, E-H from Booth & Kelman CAM36-7 (BRI). Del. W.Smith.

Euphorbia accedens is similar to some

forms of E. schultzii but differs in having

capsules broad-elliptic in lateral view that are

more or less as long as wide, 1.6-2 mm long,

1.9-2.2 mm across and shallowly 3-lobate

(versus depressed ovate or transversely

elliptic in lateral view and distinctly shorter

than wide, 1.5-2.2 mm long, 2-3 mm across,

deeply 3-lobate for E. schultzii), seed surface

with 5-7 prominent ± transverse, rounded

ridges or irregularly ridged (versus 3-5 ridges

for E. schultzii ) and leaf margins entire or

minutely toothed distally (versus leaf margins

mostly coarsely toothed for E. schultzii).

Etymology: The specific epithet is from Latin

accedens, an indeclinable participle meaning

approaching, coming near to, or resembling.

This alludes to this species’ similarity to

the taxa E. australis var. hispidula and E.

thelephora var. rugosa.

459

Halford & Harris, Euphorbia section Anisophyllum in Australia

2. Euphorbia albrechtii Halford & r

W.K.Harris, species nova arete similis E. 1

cinereae W.Fitzg. et E. petalae Ewart & t

L.R.Kerr glandis involucralibus capsulis C

seminibus sed a duobus habitu robustiore 1

et stylis 1/3—1/2 longitudinis dividis differt. i

Insuper ab E. cinerea glandulae appendicibus S

longioribus capsulis seminibusque grandioribus \

et ab E. petala glandulae appendicibus (

brevioribus floribus staminalibus paucioribus (

in quoque cyathio differt. Nomen E. a

drummondii saepe misapplicatum pro earn. ]

Euphorbia albrechtii autem ab ilia glandulae (

appendicibus grandioribus 0.2-0.4 mm t

longis (in vicem <0.1 mm longis) capsulis 1

latissimis versus basim acute carinatis (ad a

vicem latissimis ad aequatorem carinatis s

obtuse) habitu robustiore seminibus porcatis S

irregulariter (in vicem superficies praeditis C

porcis distinctis transversis) differt. t

Typus: Northern Territory. 8 km NE [of] f

Bloods Range Outstation, 1 October 2001, a

D.E.Albrecht 10075 (holo: NT). r

Monoecious, herbaceous perennial to 30 cm

high, many stems arising from crown of thick |

cylindrical taproot, the whole plant glabrous.

Stems erect to ascending (rarely prostrate), z

sparingly to much branched, longitudinally {

ridged when young. Interpetiolar stipules j

subulate or narrow-triangular, 0.5-1.1 mm c

long, deeply bipartite to base, glabrous; (

margin laciniate. Leaves: petiole 0.5-0.6 mm k

long, smooth; blade narrow-oblong, oblong J

or obovate, 6-9 mm long, 2.3-4 mm wide, g

1.8-3 times longer than wide, smooth, green i

sometimes with reddish tinge on margin; l

base asymmetric with one side rounded to

cordate, the other rounded; margin sparingly r

minutely toothed distally; apex rounded. i

Cyathia solitary at the nodes, often appearing c

clustered on short leafy lateral branchlets 1

with subtending leaves usually slightly .

smaller than primary stem leaves; peduncles ^

c. 0.3 mm long. Involucres campanulate or i

turbinate, 0.8-1 (-1.4) mm long, 0.7-1.1 mm J

across; lobes 5, triangular, 0.2-0.4 mm long, J

margin entire or fimbriate; glands 4, shortly }

stipitate, cupuliform, with distinct central (

pit, transverse-oblong in outline, 0.1-0.4 mm l

long, 0.4-0.7 mm wide, cream or red; gland *

appendages conspicuous, spreading radially,

reniform or depressed obovate, 0.2-0.4 mm

long, 0.6-1 mm wide, pink to red or cream,

glabrous, entire or shallowly lobed; bracteoles

0.4-0.6 mm long, adnate for c. 1/4 of their

length to involucre, free portion divided

into numerous subulate glabrous segments.

Staminate flowers 2-5 per cyathium;

pedicels 0.8-1.1 mm long; staminal filaments

0.1-0.3 mm long. Pistillate flowers: styles

0.3-0.5 mm long, spreading with recurved

apices, smooth, glabrous, each bifid for

1/3—1/2 of their length, the apices terete.

Capsules exserted from involucre on pedicel

to 2 mm long, broad to very broad-ovate in

lateral view, 1.5-1.8 mm long, 1.7-2.2 mm

across, shallowly 3-lobate with keels acute,

smooth, glabrous; hypogynous disc entire.

Seeds ovate in outline, 1-1.3 mm long, 0.5-

0.8 mm tangentially, 0.5-0.7 mm radially,

tetraquetrous in cross section; dorsal faces

planar to concave; ventral faces concave;

all faces with low faint irregular smooth

ridges; exotesta thin, of even thickness over

surface, white, microreticulate, becoming

mucilaginous when moistened; endotesta red-

brown Figs 2, 25A.

Additional selected specimens examined: Western

Australia. Kimberley, May 1984, Fatchen 965 (AD);

justNW of Wolf Creek Crater, Apr 1979, George 15290A

(BRI, PERTH); Great Sandy Desert, Tanami Track,

c. 2 km W of NT - WA border, c. 225 km SE of Halls

Creek, May 1976, Beauglehole ACB51018 (PERTH); 82

km ESE of Telegraph Line on Ankatell Ridge Road, Aug

1977, George 14822 (BRI, PERTH); Great Sandy Desert,

May 1979, George 15712 (BRI, PERTH); edge of Great

Sandy Desert, c. 9 km by road ESE of edge of Gregory

Range, at 106 km post WNW from Telfer Mine, Aug

1977, Jackson 2984 (AD, HO, MEL); Lake Henern, May

1981, Cane 46 (DNA); 9 miles [c. 14 km] W of Mt Webb,

Jul 1967, George 9037 (PERTH); Patience Well, Gibson

Desert, Feb 2001, Campbell 395 (PERTH); 10 miles [c.

16 km] N of Giles, Aug 1962, Kuchel 107 (AD); Serpents

Glen, Carnarvon Range, Little Sandy Desert, Aug 2001,

Kenneally K12165 & Edinger E2629 (PERTH); Everard

Junction, Gibson Desert N.R., May 1988, Pearson

395 (PERTH); 81 miles [c. 130 km] SW of Warburton

Mission, Aug 1961, George 2900 (PERTH). Northern

Territory. Gibson Creek, c. 56 km N of Tennant Creek,

July 1968, Orchard 861 (AD, DNA); 28 miles [c. 45 km]

W [of] Frewena R/H, Nov 1971, Latz 1846 (DNA, NT);

Lake Surprise, Mala Paddock near camp, Sep 1993,

Parsons 626 (DNA); Stuart Highway, between Tennant

Creek and Barrow Creek, c. 11 km N of Wauchope, Jul

1964, Eichler 17921 (AD); 6 km S [of] Camel Bore,

Newhaven Reserve, May 2002, Latz 18657 (NT); c. 6

km SW [of] Midway Bore, c. 38 km SE [of] Yuendumu,

460

Austrobaileya 8(4): 441-600 (2012)

Fig. 2. Euphorbia albrechtii. A. habit *0.5. B. branchlet with cyathia *8. C. leaf *8. D. stipules xl6. E. cyathia with

female flower x24. F. cyathial gland with appendage, adaxial view x32. G. capsule, top view xl6. H. capsule, lateral

view xl6. A-C, E, F from Albrecht 10075 (NT); D from Albrecht 6304 (DNA); G, H from Fatchen 965 (AD). Del.

W. Smith.

461

Halford & Harris, Euphorbia section Anisophyllum in Australia

Mar 1995, Albrecht 6304 (DNA, MEL); 62 miles [c. 100 J

km] NNW of Alice Springs, Jun 1958, Winkworth 1458 c

(DNA, NT); Hamilton Downs, No. 2 desert bore, Apr <

1956, Chippendale 2019 (DNA, MEL). 1

Distribution and habitat : Euphorbia ^

albrechtii extends from the Little and Great \

Sandy Deserts, WA, eastward through the 1

southern NT to the Qld border (Map 2). It c

grows in Triodia grassland communities on r

red sandy soils on sand plains, inland sand f

dunes or dune swales. It has often been noted

on specimen labels to be growing in areas c

“recently burnt”. 1

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from May to August.

Notes : Euphorbia albrechtii is most similar to

E. cinerea and E. petala which have similar

involucral glands, capsules and seeds. It

differs from both species by having a more

robust habit and styles divided for 1/3—1/2 of

their length. It also differs from E. cinerea in

having longer gland appendages and larger

capsules and seeds. It differs from E. petala

in having shorter gland appendages and

fewer staminate flowers per cyathia. These

differences are summarized in Table 1.

Table 1. Morphological comparison between Euphorbia albrechtiU E. cinerea and E. petala

Character

E. albrechtii

E. cinerea

E. petala

habit

robust erect

herbaceous

perennial rarely

prostrate

slender prostrate

herbaceous

perennial

slender prostrate

herbaceous

perennial

involucre length (mm)

0.8-1

1.1-1.5

gland appendage length

(mm)

0.2-0.4

0.1-0.3

0.3-1.3

staminate flowers per

cyathium

2-5

1-5

7-10

style division

bifid for 1/3—1/2 of

their length

entire

entire or scarcely

bifid at the tip

capsule size (mm)

1.5-1.8 x 1.7-2.2

1.1-1.3 x 1.3-1.5

1.7-1.8 x 1.7-2.2

seed length (mm)

1-1.3

0.7-0.8

1-1.2

The name Euphorbia drummondii has

often been applied to this species in the past.

E. albrechtii differs in having larger gland

appendages 0.2-0.4 mm long (versus <0.1

mm long for E. drummondii ), capsules that

are broadest towards the base and acutely

keeled (versus broadest towards the equator

and obtusely keeled for E. drummondii ),

more robust habit, and seeds with surfaces

irregularly ridged (versus seed surfaces

with 3-6 distinct transverse ridges for E.

drummondii ).

Etymology : The species is named in honour

of botanist David E. Albrecht, Alice Springs

Herbarium (1993-2012), who has made

significant contributions to our knowledge of

the flora of Central Australia.

3. Euphorbia armstrongiana Boiss., in

A.DC., Prodr. 15(2): 47-48 (1862). Type:

[Northern Territory.] Port Essington, s.d.,

[J. W] Armstrong 530 (lecto [here designated]:

K 186465 [ex Herb. Hook.]; isolecto: K 186464

[ex Herb. Benth.]).

Monoecious, annual with lax habit to 60

cm high, few to several stems arising from

slender taproot. Stems decumbent to erect

(rarely prostrate, Lazarides & Adams 146

[DNA]), sparingly to much branched,

smooth, glabrous or sparsely to moderately

462

pilose with spreading white hairs 07-2.5

mm long. Interpetiolar stipules triangular,

0.3-0.6 mm long, entire or deeply bipartite

to base, glabrous adaxially, hairy abaxially

with straight stiff white hairs to 0.3 mm

long; margin entire or laciniate. Leaves:

petiole 1-2.3 mm long, smooth, glabrous

or with indumentum as for stems; blade

oblong, elliptic or occasionally broad-elliptic

to rotund, 7-20 mm long, 5-10 mm wide,

1-2.2 times longer than wide; both surfaces

smooth; adaxial surface green occasionally

with reddish tinge, glabrous; abaxial surface

pale green, glabrous or sparsely pilose with

spreading, ± straight hairs 0.9-1.1 mm long;

base asymmetric with one side cordate,

the other obtuse to rounded; margin entire

or sparingly minutely toothed, sometimes

only proximally; apex rounded to obtuse,

sometimes minutely apiculate. Cyathia

solitary at the upper nodes; peduncles 1-3.5

mm long. Involucres cupuliform, 0.9-1.4 mm

long, 1.3-1.8 mm across; lobes 5, triangular,

0.4-0.5 mm long, margin fimbriate; glands

4, stipitate, patelliform, planar or shallowly

concave, transverse-oblong in outline,

0.4-0.6 mm long, 0.6-1.1 mm wide, pale

green; gland appendages conspicuous to

inconspicuous, spreading radially, transverse-

linear or lunate, 0.1-0.25 mm long, 0.9-1.3

mm wide, white or occasionally pink,

glabrous, margin ± entire; bracteoles 0.8—1

mm long, adnate for c. 1/2 of their length to

involucre, free portion divided into numerous

± linear hairy segments. Staminate flowers

18-22 per cyathium; pedicels 1.3-1.5 mm

long; staminal filaments 0.4-0.5 mm long.

Pistillate flowers: styles 0.7-0.9 mm long,

connate at the base into a column for c. 1/4 of

their length, erect, recurved distally, smooth,

glabrous, each bifid for c. 1/3 of their length,

the apices terete. Capsules exserted from

involucre on pedicel to 5 mm long, very

Austrobaileya 8(4): 441-600 (2012)

broad-ovate in lateral view, 2.5-3 mm long,

2.8-3.8 mm across, shallowly 3-lobate with

keels obtuse, smooth, glabrous; hypogynous

disc entire. Seeds broad-ovate in outline, 1.6-

1.9 mm long, 1.2-1.4 mm tangentially, 1.1-1.4

mm radially, tetragonous in cross section;

dorsal and ventral faces ± planar, with 4-6

distinct transverse grooves; exotesta thin, ±

of even thickness over surface, pale brown

or pale grey, micropapillate, not becoming

mucilaginous when moistened; endotesta red-

brown.

Distribution: Euphorbia armstrongiana

extends from the Kimberley region, WA to

north-western NT.

Typification: In his protologue of E.

armstrongiana, Boissier (1862) states “Ad Port.

Essington Australiae tropicae (Armstrong

n. 530!)..(v.s. in h. Kew.)” Two sheets of the

Armstrong collection have been located

amongst material on loan to BRI from K,

and are numbered (186464 ex Herbarium

Benthamianum 1854) and (186465 ex

Herbarium Hookerianum 1867) respectively.

The specimen numbered 186465 is selected

here as lectotype of E. armstrongiana as it is

part of the original material, morphologically

agrees with the protologue description and is

the more ample of the two specimens.

Notes: Euphorbia armstrongiana is easily

distinguished from all other species of E.

sect. Anisophyllum in Australia by its very

distinctive seed sculpturing and generally

slender stems with elongated internodes.

The seed surface is sulcate with 4-6 distinct

transverse grooves (Fig. 25B).

As circumscribed here, E. armstrongiana

exhibits discontinuous variation in branchlet

indumentum associated with a geographical

disjunction. These taxa are here recognised as

varieties which can be distinguished using the

following key.

Key to varieties of Euphorbia armstrongiana

Stems hairy, sometimes only proximally

Stems glabrous.

3a. E. armstrongiana var. armstrongiana

.3b. E. armstrongiana var. distans

463

Halford & Harris, Euphorbia section Anisophyllum in Australia

3a. Euphorbia armstrongiana var.

armstrongiana

Illustration: Dunlop etal. (1995: 218, fig. 72).

Stems with a sparse to moderately dense

indumentum of spreading, ± straight, white

hairs 0.7-2.5 mm long.

Additional selected specimens examined : Northern

Territory. Cobourg Peninsula, Smith Point, Mar 2002,

Rider & Firth 1773 (DNA); c. 25 km NE of Jabiru,

Mar 1981, Craven 6608 (AD, BRI, DNA); 10 km S of

Oenpelli, Arnhem Land, May 1988, Munir 5831 (AD);

Litchfield N.P, Mar 1995, Cowie 5323 & Taylor (BRI);

Mary River, May 1989, Clarke 1776 (BRI); c. 6 miles [c.

10 km] N of Pine Creek township. Mar 1965, Lazarides

& Adams 146 (DNA, NSW); UDP [Gunlom] Falls, 108

km NE of Pine Creek (Waterfall Creek N.R.), May 1983,

Barker 506 (AD, NSW); Kakadu N.P, 19.1 km S of

old Darwin road, along road to Pine Creek, Apr 1992,

Halford Q1124 (BRI); Kakadu Stage 3, Kambolgie

Creek, Apr 1993, Egan 2129 (BRI, DNA); Edith River

area, Apr 1999, Michell & Risler 2339 (DNA).

Distribution and habitat: Euphorbia

armstrongiana var. armstrongiana occurs

in north-western NT, in an area more or

less bounded by the Cobourg Peninsula,

Oenpelli, Katherine and Litchfield N.P. (Map

3a). It grows in eucalypt woodland/forest

communities sometimes in Allosyncarpia

forest communities on sandy soils mostly

derived from sandstone. It is commonly found

amongst boulders on sandstone plateaux,

scree slopes, rocky hillsides or rocky crevices.

Phenology: Flowers and fruits have been

collected from March to May.

3b. Euphorbia armstrongiana var. distans

(W.Fitzg.) Halford & W.K.Harris combinatio

et status nova; Euphorbia distans W.Fitzg.,

J. & Proc. Roy. Soc. Western Australia 3:

160 (1916). Type: Western Australia. Mount

Broome, May 1905, W.VFitzgerald818 (lecto

[here designated]: PERTH 2847248; isolecto:

NSW 98807).

Illustration: Wheeler (1992: figs 183C, 184F,

185F, 186F), as Euphorbia distans.

Stems glabrous.

Additional selected specimens examined: Western

Australia. West Governor Island, Napier Broome Bay,

May 1984, Willis s.n. (MEL 296755); West Montalivet

Island, Bonoparte Archipelago, Jun 1992, Kenneally

7727<5(PERTH); ‘Shelly Beach Island’ in Prince Frederick

Harbour, May 1987, Kenneally 10008 (PERTH); fire plot

2, 15 km W of airstrip on Mitchell Falls track, Mitchell

Plateau, Oct 1981, Farrell 953 (PERTH); creek entering

inlet of Talbot Bay, 23 km SE of Cockatoo Island, May

1983, Fryxell & Craven 3889 (PERTH); Long Island,

Buccaneer Archipelago, Jun 1982, Hopkins BA136

(PERTH); Sunday Island, Buccaneer Archipelago, Jun

1982, Hopkins BA7 (PERTH); Barker River Gorge, 4 km

N of Mt Hart Station homestead. King Leopold Ranges,

Jun 1988, Edinger 611 (PERTH); Yellow Man Creek, 10

km NE of Mount Hart homestead. May 1998, Kenneally

12145 (PERTH); Fern Creek, King Leopold Range, 1.3

km W of Mt Bell, May 1988, Cranfield 6723 (PERTH).

Distribution and habitat: Euphorbia

armstrongiana var. distans occurs in the

Kimberley, WA, from Bonaparte Archipelago

south to King Leopold Range (Map 3b).

It grows in woodland communities along

creeks, on hillslopes or on flat terrain on clay

to sandy soils derived from granite, sandstone

or basalt. It is also found growing in rock

crevices in vine thicket communities and

amongst coastal rocks on beach conglomerate

or beach sand.

Phenology: Flowers and fruits have been

collected from April to July with one

collection in October.

Typification: Fitzgerald (1918) based his

description of Euphorbia distans on material

he collected from near Mt Broome in the

Kimberley, WA. Two sheets of a collection

made by Fitzgerald from Mt Broome have

been located amongst material on loan to BRI

from PERTH [2847248] and NSW [98807],

The sheet numbered 2847248 from PERTH

is here chosen as lectotype because it is the

best preserved and more ample of the two

specimens and morphologically agrees with

the protologue description of this species.

4. Euphorbia australis Boiss., Cent.

Euphorb. 15 (1860); Chamaesyce australis

(Boiss.) D.C.Hassall, Aust. J. Bot. 24: 640

(1976). Type: [Western Australia.] baie des

chiens marins [bay of the sea dogs (Shark

Bay)], [September 1818], C.Gaudichaud 1400

(holo: G-DC [G 149371] n.v. (image seen)

(IDC microfiche 800-73. 2421: III. 1); iso: P

313100).

Monoecious, herbaceous perennial to 30 cm

high, fewto many stems arising from thickened

taproot. Stems prostrate, ascending to erect,

much branched, smooth, with a sparse to dense

464

indumentum; hairs spreading, ± straight, 0.1-

0.9 mm long, white. Interpetiolar stipules

narrow-triangular, 0.5-1 mm long, deeply

bipartite, with indumentum as for stems;

margin laciniate. Leaves: petiole c. 1 mm

long, smooth, indumentum as for stems; blade

oblong, obovate or oblong-elliptic, 4-16 mm

long, 2-9 mm wide, 1.5-2.4 times longer than

wide; adaxial surface green or red, smooth or

minutely papillose, glabrous or with a sparse

to dense indumentum consisting of ascending

to spreading, ± straight hairs, 0.1-0.8 mm

long; abaxial surface green or red, minutely

papillose, with a sparse to dense indumentum

as for abaxial surface; base asymmetric

with one side cordate the other rounded;

margin entire or toothed distally; apex

rounded. Cyathia solitary at the nodes, often

clustered on short leafy lateral branchlets

with subtending leaves smaller than the

primary stem leaves; peduncles 0.6-0.7 mm

long. Involucres turbinate, 0.9-1.2 mm long,

0.9-1 mm across; lobes 5, triangular, 0.3-0.6

mm long, margin entire or ciliate; glands 4,

stipitate, patelliform or cupuliform, concave

or with shallow central pit, transverse-oblong

or transverse-elliptic in outline, 0.1-0.3 mm

long, 0.3-0.6 mm wide, red or yellow; gland

appendages conspicuous to inconspicuous,

spreading radially, transverse-oblong

to obdeltoid, 0.1-0.4 mm long, 0.3-0.9

mm wide, red or yellow or creamy white,

glabrous; margin dentate or irregularly lobed;

bracteoles 0.5-1 mm long, adnate for c. 1/5 of

their length to involucre, free portion entire or

divided into numerous ± subulate glabrous or

hairy segments. Staminate flowers 5-10 per

cyathium; pedicels 1-1.2 mm long; staminal

filaments 0.1-0.2 mm long. Pistillate flowers:

styles 0.3-0.6 mm long, spreading and

Austrobaileya 8(4): 441-600 (2012)

recurved distally, smooth, glabrous, bifid for

1/3-2/3 of their length, with apices terete.

Capsules exserted from involucre on pedicel

to 2 mm long, ± broad-elliptic in lateral

view, 1.4-2 mm long, 1.5-2.1 mm across,

shallowly 3-lobate with keels obtuse, smooth

or minutely papillose, with a moderately

dense to dense indumentum; hairs spreading,

0.1-0.7 mm long; hypogynous disc entire.

Seeds broad-ovate in outline, 0.8-1.6 mm

long, 0.6-0.9 mm tangentially, 0.6-0.9 mm

radially, tetraquetrous in cross section; dorsal

faces planar or convex; ventral faces planar

or concave; all faces faintly to distinctly

irregularly ridged or undulate; exotesta thin,

of even thickness over surface, grey-white

or brown, microreticulate, mostly becoming

mucilaginous when moistened; endotesta red-

brown.

Distribution and habitat : Euphorbia

australis is widespread across the arid zone

of Australia occurring in all mainland States

excluding Vic.

Notes: Euphorbia australis is consistently

hairy on most parts. It is most similar to E.

careyi, E. centralis and E. vaccaria. For

features distinguishing E. australis from

those species, refer to the ‘Notes’ section

under the species concerned.

As circumscribed here, this species is

a morphologically variable complex that

exhibits a wide range of variation in habit and

length of indumentum, as well as variations

in size of seeds, degree of serration of the

leaf margin and degree of lobing on gland

appendages. Four varieties are here formally

recognised which can be distinguished using

the following key.

Key to varieties of Euphorbia australis

1 Upper surface of leaf blades glabrous or rarely with a few

scattered hairs.4c. E. australis var. glabra

1. Upper surface of leaf blades sparsely to densely hairy.2

2 Leaf margins entire; plants moderately to densely villose; seeds <1.3 mm

long.4e. E. australis var. subtomentosa

2. Leaf margins toothed sometimes only distally; plants hispid to

hispidulous, pilose or sparsely to densely villose; seeds 1-1.6 mm

long

Halford & Harris, Euphorbia section Anisophyllum in Australia 465

3 Plants hispid to hispidulous; hairs on capsules mostly <0.3 mm long;

dorsal faces of seed faintly irregularly ridged or undulate. 4d. E. australis var. hispidula

3. Plants villose or pilose; hairs on capsules >0.3 mm long; dorsal faces of

seed distinctly irregularly ridged.4

4 Plants pilose to villose; leaf blades 9-16 mm long by 3-9 mm wide;

cyathia mostly green; glands yellow to red; gland appendages often

white to creamy yellow, shallowly lobed or toothed; grows in coastal

and subcoastal areas.4a. E. australis var. australis

4. Plants villose; leaf blades 3.2-8 mm long by 2-5 mm wide; cyathia and

glands red; gland appendages red, usually acutely toothed;

grows in arid inland areas.4b. E. australis var. erythrantha

4a. Euphorbia australis var. australis

Herb to 10 cm high; stems usually prostrate,

rarely ascending. Indumentum pilose to

villose; hairs (0.2) 0.6-0.9 mm long. Leaves

oblong to oblong-elliptic, 9-16 mm long, 3-9

mm wide, margin toothed distally, adaxial and

abaxial surfaces pilose to villose. Involucres

mostly green; glands 0.1-0.3 mm long, 0.3-

0.5 mm wide, yellow to red; appendages

transverse-oblong to obdeltoid, 0.1-0.2(0.4)

mm long, 0.2-0.5(0.9) mm wide, shallowly to

deeply lobed, mostly white to creamy yellow.

Capsules 1.5-1.8 mm long, 1.8-2 mm wide;

hairs 0.4-0.5 mm long. Seeds 1.1-1.6 mm

long, 0.6-0.9 mm tangentially, 0.6-0.9 mm

radially, with distinct irregular ridges on

dorsal faces.

Additional selected specimens examined : Western

Australia. Varanus Island, Lowendale Group, May

1991, Thomson 3500 (DNA); Barrow Island, Oct 1980,

Buckley 6731 (PERTH); Sandy Island, Flying Foam

Passage, Dampier Archipelago, May 1960, Royce 6386

(PERTH); Burrup Peninsula, May 1991, Thomson 3508

(PERTH); between Port Hedland & Mundabullangana

Station, Feb 1962, George 3345 (PERTH); Thevenard

Island, W of Onslow, May 1960, Royce 6367 (PERTH);

North West Cape, Aug 1960, George 1385 (PERTH);

near Norcape Lodge, Exmouth, Jul 1977, McFarland

2 (BRI, PERTH); Beagle Island, 50 miles [c. 80 km]

NE of Onslow, May 1960, Royce 6375 (PERTH); Ward

Reef road due north of C platform, Thevenard Island,

May 1990, White MRW040 (PERTH); Thevenard Island,

Jun 1988, Long VL260 (DNA); Bay of Rest, William

Preston Point, Exmouth Gulf, Aug 1980, Kenneally

7377 (PERTH); 21 miles [c. 34 km] S of Learmonth, Jun

1961, George 2447 (PERTH); 23 miles \c. 37 km] S of

Learmonth, Aug 1960, George 1264 (PERTH); 63 miles

[c. 101 km] S of Learmonth, Aug 1960, George 1426

(PERTH); 12 km E of Cape Cuvier, N of Carnarvon,

Aug 1995, Keighery & Gibson 883 (PERTH); Carnarvon

- Quobba Road, c. 33 km N of Carnarvon, Jul 1969,

Wilson 8376 (PERTH); Carnarvon on Gascoyne River

bank. May 1995, Cranfield 9682 (PERTH); Shark Bay,

Peron Peninsula, S of Carnarvon, Sep 1940, Blackall

4644 (PERTH); Bundegee Wash, Sep 1998, Mitchell-

Smith 41 (PERTH).

Distribution and habitat : Euphorbia

australis var. australis occurs on the islands

and coastal and subcoastal parts of north¬

western WA from near Port Hedland south

to Peron Peninsula, Shark Bay (Map 4a). It

grows on sandy soils on foreshores, coastal

sandhills, creek banks or alluvial plains.

Phenology: Flowers and fruits have been

collected sporadically throughout the year

mostly from May to August.

4b. Euphorbia australis var. erythrantha

(F.Muell.) Benth., FI. Austral. 6: 48 (1873);

Euphorbia erythrantha F.Muell., Fragm. 2:

152 (1861). Type: [New South Wales.] Barrier

Range, [1860] Viet. Exped. [ H.Beckler .$.«.]

(lecto [here designated]: MEL 1560386;

isolecto: MEL 1560385).

Herb to 10 cm high; stems usually prostrate,

rarely ascending to erect. Indumentum villose;

hairs (0.2) 0.6-0.9 mm long. Leaves oblong,

suborbicular to obovate, 3.2-8 mm long, 2-5

mm wide, margin toothed distally, adaxial and

abaxial surfaces villose. Involucres mostly

red; glands 0.1-0.2 mm long, 0.3-0.6 mm

wide, red; appendages obdeltoid, 04-0.2(0.4)

mm long, 0.2-0.5(0.9) mm wide, shallowly

to deeply toothed, red. Capsules c. 1.8 mm

long and 2 mm wide; hairs 0.4-0.6 mm long.

Seeds with faint to distinct irregular ridges

on dorsal faces, 1.2-1.6 mm long, 0.7-0.8 mm

tangentially, 0.7-0.9 mm radially. n= 11.

466

Additional selected specimens examined: Western

Australia, near Lake Hopkins, Jul 1967, George 8895

(PERTH); Elder Creek, 2 miles [c. 3 km] W of Warburton,

Aug 1962, George 3830 (PERTH); Mt Eveline, E of

Warburton, Aug 1962, George 3886 (PERTH); just

to N of Blackstone Range, Jun 1977, Burbidge 20/77

& Fuller (PERTH). Northern Territory. Thomas

Reservoir, Apr 1987, Thomson 1715 (DNA). South

Australia. Evelyn Creek, c. 11 km E of Mt Willoughby

homestead, Sep 1966, Shaw 513 (AD); Evelyn Downs,

Sep 1955, Ising (AD 966150252); 55 km S of Mt

Willoughby homestead. May 1984, Badman 1095 (AD);

Balcanoona, near Nudlamutana Well, Oct 1967, Eichler

19623 (AD); Arkaroola Station, Sep 1971, Kuchel 2936

(AD); Mt Chambers, Oct 1975, Whibley 5623 (AD);

Paralana Hot Springs, Nov 1993, Bates 35069 (AD);

Mount Harper Island, Apr 1993, Bates 31973 (AD); c.

3 km E of Aroona Dam, Sep 1972, Lothian 5229 (AD).

New South Wales. 9 miles [c. 14 km] N of Silverton,

May 1973, Hassall 7314 (BRI); Broken Hill, Sep 1919,

Morris 7 (BRI); McDougalls Well, N of Broken Hill,

Mar 1972, Milthorpe 708 (NSW); Mundi Mundi Creek,

May 1979, Fox 7905006 (NSW); Conservation paddock,

Fowler’s Gap Station, c. 16 km SSW of Lake Bancannia,

Aug 1973, Cunningham 1345 & Milthorpe (NSW); back

paddock, Gnalta Station, 95 miles [ c . 153 km] NE of

Broken Hill, Sep 1973, Cunningham 1197 & Milthorpe

(NSW).

Distribution andhabitat: Euphorbia australis

var. erythrantha occurs in the southern arid

zone where it extends from Warburton, WA,

through SA to White Cliffs, in western NSW

(Map 4b). It grows mostly in bare open areas

or in open shrubland communities on rocky

or stony slopes of sandstone, limestone or

granite, stony plains, or in rock crevices on

breakaway cliffs. The soils are mostly skeletal

to shallow red brown clay loams, clays or

occasionally sands.

Phenology : Flowers and fruits have been

collected throughout the year.

Typification: In the protologue of Euphorbia

erythrantha , Mueller (Joe. cit .) cited “In

deserto circum montes Barrier Range. Dr

Beckler”. Two sheets considered to be part

of the collections used by Mueller have been

located in material on loan to BRI from MEL,

a: “near Barrier Range, Beckler” [MEL

1560385]; b: “Barrier Range, Viet. Exped.”

[MEL 1560386], Both sheets have a blue

label attached with the name E. erythrantha

written in Mueller’s hand. The material on

these sheets agrees with the description in

the protologue and it is considered to be type

material. Sheet MEL 1560386 is chosen as the

Austrobaileya 8(4): 441-600 (2012)

lectotype for E. erythrantha because it is the

more ample specimen.

Dr Hermann Beckler was the medical

officer and botanist to the original Burke and

Wills, Victorian Exploration Expedition in

1860. Willis (1962) noted that Beckler never

reached the Barrier Range and that Mueller

and Bentham incorrectly assign many of

Beckler’s collections from the Scrope Range

( c . 90 km E of the Barrier Range) to this

locality.

Notes: Euphorbia australis var. erythrantha

is similar to E. australis var. subtomentosa

with its villose indumentum and red cyathia,

glands and gland appendages. It differs in

having toothed leaf margins and mostly

acutely toothed gland appendages.

It can be difficult to distinguish between E.

australis var. erythrantha and some forms of

E. centralis with which it is sympatric in parts

of eastern WA and northern SA. For features

distinguishing E. australis var. erythrantha

from E. centralis , refer to the ‘Notes’ section

under that species.

4c. Euphorbia australis var. glabra Halford

& W.K.Harris, varietas nova arete similis

E. australi var. subtomentosae Domin

quoad indumentum et formam foliorum.

Ab omnibus aliis varietatis E. australis

superficie foliorum supra glabris differt.

Typus: Western Australia. Hamersley Range

National Park, 4.2 km along Mt Bruce

homestead (abandoned) track from Marandoo

East Road, 14 May 1980, M.Trudgen 2579

(holo: PERTH; there are three unmounted

duplicates (isotypes) with the holotype sheet).

Herb; stems prostrate. Indumentum pilose;

hairs 0.4-1.3 mm long. Leaf blades elliptic

to broadly elliptic, 3-8 mm long, 2-6 mm

wide, margin entire; adaxial surface glabrous

or rarely with a few scattered pilose hairs;

abaxial surface pilose. Involucres mostly

green; glands 0.1-0.2 mm long, 0.3-0.6

mm wide, pale green or red; appendages

transverse-oblong or obdeltoid, 0.2-0.3 mm

long, 0.5-0.6 mm wide, entire, mostly pink.

Capsules 1.7-2 mm long, 1.6-2.1 mm wide;

hairs 0.5-0.7 mm long. Seeds with low faint

irregular ridges on dorsal faces, 1.4-1.5 mm

467

Halford & Harris, Euphorbia section Anisophyllum in Australia

long, 07-0.8 mm tangentially, 0.8-0.9 mm

radially.

Additional specimens examined: Western Australia.

Erallinga Pool, Hamersley Ranges, Apr 1997, Trudgen

MET18971 (PERTH); Caves Creek, Mt Brockman Road,

50 km W of Hamersley Station homestead, Sep 2006,

Halford Q9256A (BRI); Karijini N.P., Mt Bruce Flats,

13 km E of Marandoo Hill, 7.3 km SE of Mt Bruce, 9.3

km S of Mt Oxer, 5.7 km W of Mt Howieson, Hamersley

Range, Mt Bruce S, Sep 1998, van Leeuwen 3861 (BRI).

Distribution and habitat : Euphorbia australis

var. glabra is restricted to the Hamersley

Range, Pilbara, WA where it is known only

from a few scattered localities (Map 4c). It

grows on banks of semi-permanent pools or

creeklines, or alluvial flats, in Eucalyptus

camaldulensis Dehnh. open forest or E.

victrix L.A.S.Johnson & K.D.Hill low forest

on sandy to clayey-loam alluvium.

Phenology : Flowers and fruits have been

collected from April to June and September.

Notes : Euphorbia australis var. glabra is

most similar to E. australis var. subtomentosa

in indumentum and leaf shape. It differs from

all varieties of E. australis by its glabrous

upper leaf surface.

Etymology: The varietal epithet is from Latin

glaber , glabrous, in reference to the lack of

indumentum on the adaxial surface of the

leaves of this variety.

4d. Euphorbia australis var. hispidula

Halford & W.K.Harris, varietas nova ab aliis

varietatis E. australis differt a combinatiore

characterum sequente: indumentum caulium

foliorum capsularum undique hispidulosum

usque hispidum, margines foliorum distale

dentatae, pili in capsulis usque 3 mm longis,

superficies dorsales seminum porcatae leviter

irregulariter vel undulatae. Typus: Western

Australia. Mt Nameless, near Tom Price, 22

September 2006, D.Halford Q9240 (holo:

BRI; iso: MEL, PERTH, distribuendi ).

Herb to 30 cm high; stems prostrate, rarely

ascending to erect. Indumentum on stem and

leaves hispidulous to hispid, moderately dense

with hairs 0.1-0.6 mm long. Leaves oblong,

oblong-elliptic or oblong-obovate, 4-10 mm

long, 2-6 m wide; margin toothed distally.

Involucres mostly red; glands c. 0.1 mm

long, 0.2-0.3 mm wide; gland appendages

transverse-oblong, shallowly to deeply lobed,

0.1-0.2 (0.4) mm long, 0.2-0.5(0.9) mm wide.

Capsules 1.4-1.9 mm long, 1.5-2.1 mm

wide; hairs 0.1-0.3 mm long. Seeds faintly

irregularly ridged or undulate, 0.9-1.2 mm

long, 0.6-0.8 mm tangentially, 0.6-0.8 mm

radially.

Additional selected specimens examined : Western

Australia. Djaluwon Creek, near S end of Lake Gregory,

Apr 1979, George 15376 (PERTH); 27 miles [c. 43 km]

N of Lookout Rocks, May 1947, Royce 1855 (PERTH);

10.9 km S of the Mt Bruce turnoff on the Wittenoon to

Nanutarra Road, Sep 1991, Trudgen MET10610 & Maley

(PERTH); MtNameless, near Tom Price, Jul 1980 , Atkins

838 (PERTH); c. 35 km W along Mt Brockman Road from

Hamersley Station homestead, Sep 2006, Halford Q9259

(BRI); 6 km W of Mujina Claypan, and 20 km ENE of

Mt Windell, Feb 1987, Mollemans 2306 (AD, PERTH);

Hamersley Range N.P., 16.4 km from Milli Milli Springs

on the track to Coppin Pools, May 1980, Trudgen 2443/a

(PERTH); 3.25 km WSW of Packsaddle Hill, Hamersley

Ranges, Jul 1997, Trudgen 16711 (PERTH); 21 km N

of Juna Downs homestead, Aug 1973, Trudgen 380 &

Merton (PERTH); Hamersley Range N.P, above Dales

Gorge, Aug 1974, Carr 4823 & Beauglehole 48601

(NSW); c. 45 km NE of Tom Price along Karijini Drive,

W of Marrandoo, Sep 2006, Halford Q9207 (BRI). c.

30 km W along Karijini Drive from Ranger Station,

Karijini N.P., Sep 2006, Halford Q9201 (BRI); 4.5 km

SSE of West Angela Hill, Hamersley Ranges, Jul 1997,

Trudgen 16723 (PERTH); Little Sandy Desert, Apr 1979,

Mitchell 579 (DNA, PERTH); Little Sandy Desert, Apr

1979, Mitchell 663 (PERTH). Northern Territory. 7

km NW [of] Tanami Mine, Sep 1990 Latz 11821 (MEL,

NT); Warrego, May 1993, Egan 2214 (DNA); Tennant

Creek, Mary Ann Dam, May 1993, Egan 2267 (DNA);

35.5 miles [ c . 57 km] NNW of Wauchope township, Aug

1956, Lazarides 5850 (MEL, NSW); Devil’s Marbles,

Jun 1955, Chippendale 1912 (DNA, NSW).

Distribution and habitat: Euphorbia

australis var. hispidula occurs from near Tom

Price, Pilbara, WA to Tennant Creek, NT

(Map 4d). It grows in open Triodia grassland

or Acacia! vtmWQQ shrubland with Triodia sp. in

the understorey on skeletal stony or gravelly

clay loams on ironstone hills or ridges, or on

sandy to loam soils on sandplains.

Phenology: Flowers and fruits have been

collected from April to October.

Notes: Euphorbia australis var. hispidula

differs from the other varieties of E. australis

by the following combination of characters:

the indumentum is hispidulous to hispid on

stems, leaves and capsules, leaf margins

468

toothed distally, hairs on capsules up to 0.3

mm long and dorsal faces of seeds faintly

irregularly ridged or undulate.

Etymology : The varietal epithet is from Latin

hispidus , hispid (covered with erect, rigid

hairs), in reference to the indumentum on

most surfaces of the variety.

4e. Euphorbia australis var. subtomentosa

Domin, Biblioth. Bot. 89(4): 310 (1927 ‘1926’).

Type: Western Australia. Inter Ashburton et

De Gray River s.d., E.Clement s.n. (holo: PR

528303; iso: K 186467, 186468).

Herb to 20 cm high; stems usually prostrate,

rarely ascending to erect. Indumentum

sparsely to densely villose; hairs 0.1-0.9 mm

long. Leaves oblong, suborbicular to obovate,

2.5-7 mm long, 2-4 mm wide, margin

entire, adaxial and abaxial surfaces villose.

Involucres mostly red; glands 0-0.25 mm

long, 0.1-0.4 mm wide, red or yellow; gland

appendages transverse-oblong, 0.1-0.3 mm

long, 0.3-0.4 mm wide, shallowly to deeply

toothed, red, yellow or white. Capsules 1.4-

1.8 mm long, 1.4-1.8 mm wide; hairs 0.5-0.8

mm long. Seeds with faint to distinct irregular

ridges on dorsal faces, 0.8-1.3 mm long, 0.5-

0.6 mm tangentially, 0.5-0.6 mm radially, n =

11. Fig. 25C.

Additional selected specimens examined : Western

Australia, just W of Wolf Creek Crater, Apr 1979,

George 15312 (PERTH); Meentheena Station

Conservation Reserve, 12.7 km ESE of Bullgarina Hill,

7.3 km ESE of King Rock Hole, 30 km E of Mt Edgar,

25.8 km N of Baroona Hill, May 2000, van Leeuwen

4497 (BRI, PERTH); 4 km SW of Two Sisters, c. 145

km SE of Shay Gap, Jul 1984, Newbey 10481 (PERTH);

Hamersley Range N.P., just S of Marandoo Ridge near

entrance to ‘Grimace Gulch’, Sep 1978, Trudgen 2253

(PERTH); Hamersley Range N.P, 0.3 km from Mindi

Springs on track to Juna Downs, Mar[?] 1980, Trudgen

2616 (PERTH); Rudall River, Aug 1971, Wilson 10308

(PERTH); Rudall River, May 1971, George 10641

(PERTH); Turee Creek, Turee Station, c. 40 km E of

Paraburdoo, Sep 2006, Halford Q9233A (BRI); Jacobs

Creek Gully, May 1995, Cranfield 9701 (PERTH); 1 km

NE of Thompsons Mill, Mt Gould Station, May 1986,

Cranfield 5434 (PERTH); c. 5 km NE of Wellington

Range on road from Wiluna to Carnegie homestead,

Sep 1984, Wilson 11975 (PERTH). Northern Territory.

Bunda Station, Jun 1994, Egan 4215 (DNA); Wave Hill

Station, Mar 1997, Michell 634 & Mangion (DNA);

Supplejack Station, 38 km W of homestead, Sep 1970,

Henshall 2329 (DNA); Kings Canyon, 1.5 km W [of]

George Gill Range, Jul 1981, Thomson 74 (DNA).

Austrobaileya 8(4): 441-600 (2012)

Queensland. Burke District: 251.7 km S ofNormanton,

May 1976, Hassall 7638 (BRI); 63.5 km by road S of

Burke & Wills Roadhouse on Burke Development

Road, Mar 2005, McDonald KRM4131 (BRI); 12 km

NE of Mt Isa, May 1983, Schmid 673 (BRI); Cloncurry

-Normanton road, 8.1 km N of Quamby, Jun 1999, Bean

15172 (BRI); 20 km W of Cloncurry, on road to Mount

Isa, Jun 1991, Halford Q443 (BRI, MEL).

Distribution and habitat : Euphorbia

australis var. subtomentosa is widespread in

inland WA extending across the NT to western

Qld (Map 4e). It grows in Triodia grassland,

Acac/tf/eucalypt woodland communities on

skeletal to shallow gravelly sand to clay soils

on plains, alluvial flats or rocky hills.

Phenology : Flowers and fruits have been

collected from January to October.

Notes: Euphorbia australis var. subtomentosa

is similar to E. australis var. erythrantha.

For features distinguishing of E. australis

var. subtomentosa from E. australis var.

erythrantha , refer to the ‘Notes’ section under

that variety.

As circumscribed here E. australis var.

subtomentosa has two distinct forms. Though

the extremes of these variants are distinctive

they do merge into one another with numerous

intermediates in the Pilbara region where

their distributions overlap. The two forms are

as follows:

Typical form: Stems prostrate, usually red in

colour, hairs on stems >0.4 mm long; leaves

pale green often with red blush over surfaces

or red tinge along margins; cyathia usually

red rarely grey-green, seeds 1-1.3 mm long.

This is common throughout the western part

of the range for this variety (e.g. Cranfield

5434, 9701, Trudgen 2253).

Eastern form: Stems ascending to erect or

prostrate, grey-green rarely red in colour,

hairs on stems mostly <0.4 mm long; leaves

mostly grey-green in colour; cyathia mostly

grey or grey-green rarely red, seeds 0.8-1 mm

long. This is common in north-western Qld

but extends to the Kimberley and the Pilbara

region, WA (e.g. Egan 4215, George 15312,

Hassall 7638).

469

Halford & Harris, Euphorbia section Anisophyllum in Australia

5. Euphorbia biconvexa Domin, Biblioth.

Bot. 89(4): 308 (1927 ‘1926’); Chamaesyce

biconvexa (Domin) D.C.Hassall, Aust. J.

Bot. 24: 640 (1976). Type: Queensland.

[Cook District:] apud opp. Mungana, in colle

calcareo, February 1910, K.Domin s.n. (holo:

PR 528286).

Illustration : Weber (1986: 749, fig. 401B), as

E. coghlanii.

Monoecious, annual or herbaceous perennial

with short-lived stems produced from thick

cylindrical taproot, to 80 (100) cm high, few

to many stems arising from rootstock. Stems

ascending to erect (rarely prostrate), sparingly

to much branched, smooth, glabrous or rarely

sparsely pilose with ± spreading white hairs

0.2—1 mm long. Interpetiolar stipules

narrow-triangular, 0.6-1.5 mm long, bifid or

deeply bipartite, glabrous; margin laciniate

(rarely entire). Leaves: petiole 0.7-1 mm long,

smooth, glabrous; blade oblong, narrow-

ovate or ovate, sometimes falcate, 8-36 mm

long, 2.5-9 mm wide, 2-6 times longer than

wide; adaxial surface green to dark green or

glaucous, sometimes with reddish tinge on

margin, smooth, glabrous; abaxial surface

paler than adaxial surface, smooth, glabrous or

rarely sparsely pilose with spreading, crispate

hairs to 1 mm long; base asymmetric with

one side cordate, the other rounded to cordate;

margin entire or serrulate distally; apex acute

to obtuse or rounded. Cyathia solitary at the

upper nodes, sometimes clustered on short

leafy lateral branchlets with subtending

leaves smaller than the primary stem leaves;

peduncles 1-2 mm long. Involucres turbinate,

0.8-1 mm long, 0.9-1.1 mm across; lobes

5, triangular or subulate, 0.3-0.5 mm long,

margin entire or laciniate; glands 4, stipitate,

patelliform, planar or concave, transverse-

oblong or orbicular in outline, 0.1-0.2 mm

long, 0.1-0.4 mm wide, yellowish green;

gland appendages conspicuous, spreading

radially, broad-obovate or reniform, 0.2-0.4

mm long, 0.3-0.8 mm wide, white (rarely

red), glabrous, margin entire; bracteoles 0.8-1

mm long, adnate for c. 1/3 of their length to

involucre, free portion divided into numerous

subulate glabrous segments. Staminate

flowers 1-15 per cyathium; pedicels c. 0.9

mm long; staminal filaments 0.3-0.4 mm

long. Pistillate flowers: styles 0.4-0.5 mm

long, spreading to erect, recurved distally,

smooth, glabrous, each bifid for c. 1/2 of their

length, the apices terete. Capsules exserted

from involucre on pedicel to 3 mm long,

transversely broad-elliptic in lateral view,

1.6-2 mm long; 2-2.5 mm across, deeply

3-lobate with keels acute, smooth, glabrous;

hypogynous disc entire. Seeds narrow-ovate

or elliptic in outline, 1—1.5 mm long, 0.5-

0.9 mm tangentially, 0.9-1.1 mm radially,

laterally compressed, biconvex in transverse

section; faces convex, smooth; exotesta thin,

of even thickness over surface, pale grey,

microreticulate, becoming mucilaginous

when moistened; endotesta brown, n = 8 Fig.

25D.

Additional selected specimens examined: Western

Australia. Djaluwon Creek, near S end of Lake

Gregory, Apr 1979, George 15384 (PERTH); Mt

Windell road corridor, 8 km NW of Mt Winded, 3.6

km ESE of Karijini N.P. headquarters. Mar 1992, van

Leeuwen 1105 (PERTH); Hamersley Range N.P, c. 0.5

km S from Dales Gorge turnoff, on Yampire Gorge -

Juna Downs homestead road, Aug 1977, Jackson 2938

(AD, MEL); 25 miles [c.40 km] S of Sir Frederick

Range, Aug 1962, Symon 2289 (AD); Wooramel River

crossing, Innouendy Station, May 1995, Cranfield 9733

(PERTH); 30 km S [of] Karratha, May 1991, Thomson

3501 (DNA). Northern Territory. Cutta Cutta Caves

Reserve, Feb 1989, Thomson 3271 (DNA); 2 km S of

Daly Waters on the Stuart Highway, Feb 1988, Thomson

2235 (BRI); 22 km N [of] Ucharonidge homestead, Feb

1989, Thomson 3247 (BRI); 5 km NW of Wycliffe Wed,

May 1983, Halford 83536 (DNA); Palm Valley, Sep 1958,

Chippendale 4912 (AD, DNA, MEL); Uluru (Ayres

Rock - Mt Olga) N.P., Kata Tjuta (the Olgas), Kata Tjuta

Lookout walk, on the Docker River Road, 45 km WNW

of Ranger Station, May 1988, Lazar ides & Palmer 63

(AD, DNA). Queensland. Cook District: near Telecom

Tower, Chillagoe, Mar 1990, Forster PIF6529 (BRI).

Burke District: Hughenden - Mt Isa Road, c. 104

km W of Julia Creek, Apr 1975, Halliday 424 (BRI);

Lookout and carpark area. Porcupine Gorge, 65 km N

of Hughenden, Apr 1997, Archer 340 (BRI); 27 km E

of Cloncurry; beside the Flinders Highway, Feb 2001,

Wannan 2114 & Jago (BRI). Mitchell District: Lochern

N.P, N of Stonehenge, Mar 1998, Forster PIF22371 &

Booth (BRI, MEL). Gregory North District: Mayne

River crossing on Winton to Jundah Road, Apr 1986,

Neldner 2525 & Stanley (BRI). South Australia. 5 km

SW end of Callyamurra Waterhole, Apr 1987, Reid 685

(AD); minor branch of Red Mulga Creek, May 1987,

Symon 14447 (AD, HO, MEL).

470

Distribution and habitat: Euphorbia

biconvexa is endemic to Australia and

widespread across the northern part of the

continent from Carnarvon, WA, through

the northern and southern central areas of

the NT and far northern SA to Clermont, in

central Qld (Map 5). It grows in a wide range

of grassland, or eucalypti cacia woodland

communities on sandy to clay soils on plains,

rocky hillsides or along drainage lines.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from February to September.

Notes: Euphorbia biconvexa is similar to

E. coghlanii and E. trigonosperma in habit,

general appearance and seed surface texture

but differs in having seeds that are lenticulate

in transverse section compared with

suborbicular for E. coghlanii and trigonous

for E. trigonosperma.

6. Euphorbia bifida Hook. & Arn., Bot.

Beechey Voy. 213 (1837); Chamaesyce bifida

(Hook. & Arn.) T.Kuros., Acta Phytotax.

Geobot. 51(2): 212 (2001 ‘2000’). Type:

China, “collected about the neighbourhood

of Macao and the Islands adjacent”, 20 July -

30 August 1830, Rev. G.H.Vachell 240 (holo:

E-GL n.v.; iso: K n.v. (cibachrome seen).

Euphorbia vachellii Hook. & Arn., Bot.

Beechey Voy. 213 (1837); Chamaesyce

vachellii (Hook. & Arn.) H.Hara, Enum.

Spermat. Jap. 3: 44 (1954). Type: [China.]

Macao, 17 July 1830, G.H.Vachell 241 (holo:

E-GL n.v. (image seen); iso: K n.v).

Euphorbia micradenia Boiss., in A.DC.,

Prodr. 15(2.2): 27 (1862); E. macgillivrayi

var. micradenia (Boiss.) Domin, Biblioth.

Bot. 89(4): 311 (1927 ‘1926’); Chamaesyce

micradenia (Boiss.) D.C.Hassall, Aust. J.

Bot. 24: 640 (1976). Type: [Queensland.

Cook District:] Albany Island, August 1855,

F. Mueller s.n. (holo: K 186475; iso: G-DC n.v.

(microfiche IDC 800-73. 2419: II. 4), MEL

530408, P 698542 n.v. (image seen).

Euphorbia macgillivrayi Boiss., in A.DC.,

Prodr. 15(2.2): 26 (1862); E. macgillivrayi

Boiss. var. macgillivrayi , Domin, Biblioth.

Bot. 89(4): 865 (1927 T926’); Chamaesyce

Austrobaileya 8(4): 441-600 (2012)

macgillivrayi (Boiss.) D.C.Hassall, Aust.

J. Bot. 24: 640 (1976). Type: [Queensland.

South Kennedy District:] Port Molle,

December 1847, JMcG [J.MacGillivray\ s.n.

(lecto [here designated]: K 186474).

Euphorbia mitchelliana var. stenophylla

Benth., FI. Austral. 6: 47 (1873). Type:

[Northern Territory.] Port Darwin, 7

December 1871, [M] Schultz 854; (lecto [here

designated]: K 186483).

Euphorbia macgillivrayi var. yarrabensis

Domin, Biblioth. Bot. 89(4): 311 (1927

T926’). Type: Queensland. [Cook District:]

In xerodrymio apud opp. Yarraba, January

1910, K.Domin s.n. (lecto [here designated]:

PR 528309).

Euphorbia macgillivrayi var. pseudoserrulata

Domin, Biblioth. Bot. 89(4): 311 (1927 T926’).

Type: Queensland. [Cook District:] apud

opp. Chillagoe, February 1910, K.Domin s.n.

(lecto [here designated]: PR 528314).

Euphorbia macgillivrayi f. glabrata Domin,

Biblioth. Bot. 89(4): 311 (1927 T926’). Type:

Queensland. [Moreton District:] prope

Brisbane River, 1863-1865, A.Dietrich 406

(lecto [here designated]: PR 528316).

Euphorbia serrulata Reinw. ex Blume, Bijdr.

FI. Neerl. Ind. 635 (1826 T825’), non Thuill.

Type: Timor, s.d., [C.G.C.] Reinwardt s.n.

(Herb. Lugd. Bat. No. 903.159-328) (holo: L

n.v. [photo at BRI]).

Illustrations : Brock (1988:183), as Euphorbia

vachellii; Wheeler (1992: figs 184P, 185P,

186P (1992), as E. vachellii.

Monoecious, annual or herbaceous perennial

with short-lived stems produced from thick

cylindrical taproot, to 80 cm high, one to few

stems arising from rootstock. Stems erect

to ascending, sparingly branched, smooth,

glabrous or densely puberulous on young

stems becoming sparser with age; hairs weakly

appressed to spreading, crispate, 0.1-0.5 mm

long, white. Interpetiolar stipules subulate,

0.8-2 mm long, deeply bipartite, glabrous;

margin laciniate. Leaves: petiole 1-3 mm

long, smooth, glabrous or with indumentum

as for stems; blade linear to oblong, lanceolate

to narrow-ovate or narrow-elliptic, 20-80

471

Halford & Harris, Euphorbia section Anisophyllum in Australia

mm long, 2-12 mm wide, 2.6-20 times longer

than wide; adaxial surface green sometimes

with reddish tinge on margin, smooth,

glabrous or sparsely pubescent with weakly

appressed to ascending, crispate hairs 0.3-0.6

mm long; abaxial surface pale green, smooth,

glabrous or with indumentum as for adaxial

surface; base mostly asymmetric with one

side cordate or obtuse, the other cuneate to

rounded or slightly cordate; margin serrate

sometimes only distally (rarely entire); apex

acute to obtuse. Cyathia grouped together

in congested 2-7 branched dichasial cymes

together with a solitary cyathium at the distal

nodes; peduncles 3-15 mm long; bracts

subulate to narrowly triangular or leaf-like

but much smaller than the primary stem

leaves; cyathial peduncles 1-4(6) mm long.

Involucres turbinate, 0.7-1.4 mm long, 1-1.3

mm across; lobes 5, triangular, 0.4-0.7 mm

long, margin entire or laciniate; glands 4,

stipitate, patelliform or cupuliform, planar

or with distinct central pit, transverse-

oblong to transverse-elliptic or orbicular in

outline, 0.1-0.3 mm long, 0.2-0.4 mm wide,

red or yellowish green; gland appendages

conspicuous (rarely inconspicuous), spreading

radially, broad-obovate to obdeltoid or

oblong, 0.3-2.2 mm long, 0.3-2 mm wide,

white, glabrous, margin entire; bracteoles

0.8-1.7 mm long, adnate for 1/3—1/2 of their

length to involucre, free portion divided into

few to numerous subulate glabrous or hirsute

segments. Staminate flowers 3-30 per

cyathium; pedicels 1.2-1.6 mm long; staminal

filaments 0.2-0.3 mm long. Pistillate flowers:

styles 0.3-0.8 mm long, sometimes connate

at the base into a column for c. 1/7 of their

length, spreading, smooth, glabrous, each

bifid for 1/3-2/3 of their length, the apices

stout terete. Capsules exserted from involucre

on pedicel to 3 mm long, very broad-ovate

or broad-elliptic in lateral view, 1.7-2.1 mm

long, 1.9-2.5 mm across, shallowly 3-lobate

with keels obtuse, smooth, glabrous or with a

sparse indumentum consisting of ± appressed

hairs 0.2-0.3 mm long; hypogynous disc

entire. Seeds broad-ovate in outline, 1.1-1.5

mm long, 0.8-1.1 mm tangentially, 0.8-1

mm radially, tetragonous or trigonal in cross

section; dorsal and ventral faces convex,

with 3-6 distinct narrow rounded transverse

ridges; exotesta thin, of even thickness over

surface, white, microreticulate, becoming

mucilaginous when moistened; endotesta red

brown to dark brown, n = 8. Fig. 25E.

Additional selected specimens examined : Western

Australia. ‘Duncan’s Swamp’, WSW of Amax Campsite,

Mitchell Plateau, May 1978, Kenneaily 6775 (PERTH);

‘Rocky Creek’, 27.3 km NW of Doongan Station on

Gibb River - Kalumburu Road, Junl987, Edinger 291

(PERTH); adjacent to Rocky Creek, 27.3 km NW of

Doongan Station, Jun 1987, Koch 544 (PERTH); Calder

River crossing, 80 km NNW of Beverley Springs, Jan

1993, Barrett MDB139 (PERTH). Northern Territory.

Finniss River Station, Aug 1997, Cowie 7644 & Mangion

(DNA); East Alligator floodplain, Jun 1992, Cowie 3031

(DNA); mouth of Daly River, near Palmerston Island,

Feb 1994, Leach 4006 (DNA); 41 miles [c. 66 km] E

[of] Pine Creek, Apr 1962, Nelson 301 (AD, BRI, DNA,

MEL); Kakadu N.P., Gwalmek, Jan 1995, Russell-

Smith 9182 (DNA); South Bay, Bickerton Island, Jun

1948, Specht 533 (AD, BRI, MEL); west side of Cape

Shield, May 1993, Cowie 4069 & Leach (DNA); Maria

Island, Jul 1972, Dunlop 2837 (DNA). Queensland.

Cook District: 1 km E of junction of Piccaninny and

Scrubby Creek, Archer Bend N.P, Jun 1993, Neldner

4102 (BRI). North Kennedy District: 4.8 km S of

Inkerman, Bruce Highway, Sep 1976, Williams 76079

(BRI); 6 km N of Elliot River, 67 km SE of Home Hill,

Apr 1975, McDonald 1371 & Batianoff (BRI), Nelly Bay,

near Dingo Beach, north of Proserpine, Apr 2002, Bean

18676 (BRI). South Kennedy District: Hay Point, Nov

1978, Stanley 78304 (BRI). Leichhardt District: Sutton

Development Road, ‘Wedelia’, c. 1 km W of Kemmis

Creek turnoff, Feb 1994, Champion 1019 & Dixon

(BRI). Burnett District: Branch Creek road, 9.5 km

NNW of Binjour, Feb 2005, Bean 23475 (BRI). Moreton

District: Opossum Creek, Springfield, Ipswich, Jan

1996, Bird s.n. (BRI [AQ487777]). New South Wales.

Roseberry, Apr 1947, Jones 4 (BRI [AQ202910]).

Distribution and habitat : Euphorbia bifida

occurs from South East Asia to Australia.

In Australia it is widespread across coastal

and subcoastal parts of northern Australia

from Derby, WA, to Cairns, Qld and south

along the east coast to Rosebery (near

Kyogle) in northern NSW (Map 6). It grows

in a wide variety of habitats; grassland,

Melaleuca/eucalypt woodland or open forest

communities; on rocky seashores, coastal

dunes, swamps, alluvial plains or rocky

hillsides. The soils vary from sands to

cracking clays.

Phenology : Flowers and fruits have been

collected throughout the year.

All

Typification: Boissier (1862) cited two

collections in his protologue of E.

macgillivrayi namely “Ad Port Molle

Australiae (M’Gillivray!), Gould Island

(Id. forma major!) ... (v.s. in herb. Kew!)”

Two collections, which are considered as

syntypes of the name E. macgillivrayi ,

have been located amongst material of

Euphorbia on loan to BRI from K [a: Voyage

of Rattlesnake, Bot. 192, Port Molle, Dec

?3?/47, JMcG. (K) [K 186474]; b: Voyage

of Rattlesnake, Bot 258, Gould Island, May

22nd 1848, John MacGillivray [K 186473]].

The collection from Port Molle [K 186474] is

here selected as lectotype because it is part

of the original material, is the more ample

and better preserved and has morphology that

matches the description in the protologue of

this species. Both syntypes are referable to E.

bifida as applied here.

Bentham (1873) cited four collections

made by M. Schultz from Port Darwin in his

protologue of E. mitchelliana var. stenophylla

namely “Port Darwin, Schultz, n. 38, 505, 549

and 854”. The four collections a: Port Darwin,

North Australia, s.d., [Schultz] 38 , from R.

Schomburgk Oct 1869 (K 186481); b: Port

Darwin, N. Australia, 3/70, F. Schultz 549

comm. R. Schomburgk (K 186482); c: Port

Darwin, 7/12/71, Schultz 854 (K 186483); d:

Port Darwin, N. Australia, 3 1870, Schultz

505 comm. R. Schomburgk (K186484), have

been located amongst material of Euphorbia

on loan to BRI from K. B.G. Thomson

annotated the collection Schultz 854 (K

186483) as lectotype in 1989. However, his

choice has not been published. We agree with

his choice and the collection Schultz 854 (K

186483) is here selected as lectotype because

it is part of the original material, is the more

ample and complete of the four collections

and has morphology that best matches the

description in the protologue of this variety.

The syntypes Schultz 38 and 549 are referable

to E. bifida , while Schultz 505 is referable to E.

mitchelliana var. mitchelliana as applied here.

Domin (1827) cited four of his collections

from northern Queensland in his protologue

of E. macgillivrayi var. pseudoserrulata

namely “Queensland: bei Chillagoe sehr

Austrobaileya 8(4): 441-600 (2012)

typisch (DOMIN II. 1910); Savannenwalder

bei Pentland (DOMIN III. 1910); ...Picnic

Hill in der Nahe der Russell-Miindung

(DOMIN I. 1910); ...Beech Mts. (DOMIN

III. 1910)”. Five sheets [PR 528311, 528312,

528313, 528314, 528315] which are considered

as syntypes of the name E. macgillivrayi var.

pseudoserrulata , have been located amongst

material of Euphorbia on loan to BRI from PR.

The collection from Chillagoe [PR 528314] is

here selected as lectotype because it is part of

the original material, is the more ample and,

as Domin comments in the protologue, this

collection is very typical of this variety. In

1989, B.G.Thomson annotated the collection

from Pentland, Domin [PR 528312] as

lectotype but this was not published. All of

the syntypes [PR 528311, 528312, 528313,

528314, 528315] are referable to E. bifida as

applied here.

Domin (1827) in his protologue of

E. macgillivrayi var. yarrabensis states

“Nordost-Queensland: Savannenwalder in der

Ebene bei Yarraba (DOMIN I. 1910)”. Three

collections (PR 528308, 528309, 528310)

which are considered by us as syntypes of the

name E. macgillivrayi var. yarrabensis , have

been located amongst material of Euphorbia

on loan to BRI from PR. The collection (PR

528309) is here selected as lectotype because

it is part of the original material, is the more

ample and complete of the three collections

and has morphology that best matches the

description in the protologue of this variety. In

1989, B.G.Thomson annotated the collection

Domin [PR 528310] as lectotype but this

was not published. All of the syntypes [PR

528308, 528309, 528310] are referable to E.

bifida as applied here.

Domin (1927) cited two collections in his

protologue of E. macgillivrayi forma glabrata

namely “sind z. B. die von A. Dietrch sub

no. 406 und 1031”. Two collections, which

are considered as syntypes of the name E.

macgillivrayi forma glabrata , have been

located amongst material of Euphorbia on

loan to BRI from PR [a: Queensland, “prope

Brisbane River, 1863-1865, A. Dietrich 406”

(PR 528316); b: Queensland, “prope Brisbane

River, 1863-1865, A. Dietrich 1031” (PR

473

Halford & Harris, Euphorbia section Anisophyllum in Australia

528317). The collection Dietrich 406 (PR r

528316) is here selected as lectotype because s

it is part of the original material, the more \

ample and better preserved of the syntypes of 3

this forma. Both syntypes are referable to E.

bifida as applied here. ^

Notes : Euphorbia bifida is similar to E. I

mitchelliana in seed shape and seed surface t

ornamentation but differs from that in having

its cyathia grouped together in congested 2-7

branched dichasial cymes together with a ^

solitary cyathium at the distal nodes. ^

As circumscribed here, Euphorbia ^

bifida is morphologically variable. There is

considerable variation in the general habit,

indumentum, and leaf dimensions and

shape. Although the extreme forms within

this species differ considerably from each f

other, much of the morphological variation

appears to intergrade, making it difficulty ^

in assigning some material to one or other

form. For this reason we have not formally

recognised these variants. A more detailed

analysis of Australian populations might lead ,

to the recognition of infraspecific taxa, but ^

a study of the species across its entire range

would be desirable to place this variation in

context. The more notable variants observed 4

in Australia are:

Annual form: fibrous rootstock, stems £

erect and little branched, leaves linear to c

lanceolate 5-23 times as along as wide, J

margins conspicuously serrate, (e.g. Leach t

4006, Kenneally 6775, Cowie 1235). This is \

widespread and the most commonly collected i

form and is recorded from Kimberley, WA £

across the NT to Qld. t

Perennial form: perennial rootstock with C

erect annual stems, leaves narrow-ovate to

narrow-oblong, 3-8 times as long as wide, ( -

margins not conspicuously serrate {Dunlop

5593, Nelson 301, Short 5025). This occurs in J

the NT and Qld.

Coastal form often on frontal dunes: perennial c

rootstock, stems mostly prostrate, leaves C

narrow-oblong to oblong or narrow-elliptic £

to elliptic 18-25 mm long, 2-3 times as long £

as wide. (e.g. Bean 18676, McDonald 1371 C

& Batianoff, Stanley 78304). It is commonly (

recorded along the east coast from Cape York

south to Fraser Island. The following names

placed in synonymy are applicable to this

variant: E. macgillivrayi and E. micradenia.

7. Euphorbia careyi F.Muell., Fragm. 11: 64

(1881). Type: Western Australia. Fortescue

River, in 1878, H.S.Carey s.n. (holo: MEL

61116).

Euphorbia bouleyi S.Moore, J. Linn. Soc., Bot.

45: 212 (1920). Type: [Western Australia.]

NW Coast, s.d., de Bouley s.n. (holo: BM

812174).

Dioecious (rarely monoecious), herbaceous

perennial, 15-80 cm high, often with

compact fan-like habit, single or few stems

arising from roostock. Stems ascending

or erect (rarely prostrate, Royce 7141, 7077

[PERTH]), much branched, smooth, with a

sparse to moderately dense indumentum;

hairs appressed to ascending, curved to

crispate or sometimes straight, to 0.1 (0.2)

mm long, white. Interpetiolar stipules

narrow-triangular, 0.2-0.5 mm long, deeply

bipartite, pubescent; margin laciniate.

Leaves: petiole 0.2-0.6 mm long, smooth,

with indumentum as for stems; blade oblong,

obovate or oblong-elliptic, 2-9 mm long, 1.5-

5 mm wide, 1.2-1.8 times longer than wide;

adaxial surface yellow-green or light to dark

green, smooth, with a sparse to moderately

dense indumentum (rarely glabrous, Harris

WKH2224 [BRI]); hairs spreading, curved

to crispate, to 0.1 mm long; abaxial surface

paler than adaxial surface, smooth, with

indumentum as for adaxial surface; base

asymmetric with one side rounded to cordate,

the other cuneate; margin serrulate to serrate

or entire; apex rounded to obtuse. Cyathia

solitary at the upper nodes; peduncles 0.4-1

mm long. Involucres turbinate to cupuliform,

0.5-1.1 mm long, 0.8-1.3 mm across; lobes 5,

oblong or triangular, 0.4-0.5 mm long, margin

laciniate; glands 4, subsessile, patelliform,

planar or shallowly concave, transverse-

oblong to transverse-elliptic in outline,

0.3-0.5 mm long, 0.4-0.8 mm wide, yellow;

appendages conspicuous to inconspicuous or

absent, spreading radially; transverse-oblong,

0.2-0.3 mm long, 0.5-0.9 mm wide, yellow

(rarely red), glabrous, margin dentate or

474

shallowly lobed; bracteoles 0.5-1.3 mm long,

adnate for 1/2-2/3 of their length to involucre,

free portion divided into numerous subulate

glabrous or hirsute segments. Staminate

flowers 6-25 per cyathium; pedicels 1-1.4

mm long; staminal filaments 0.3-0.4 mm

long. Pistillate flowers: styles 0.5-0.8 mm

long, connate at the base into a column

for 1/5—1/7 of their length, spreading with

recurved apices, smooth, glabrous or with

a few minute erect hairs proximally, each

bifid for 1/2-2/3 of their length, the apices

terete. Capsules exserted from involucre on

pedicel to 3 mm long, broad-elliptic or broad

to very broad-ovate in lateral view, 1.4-1.6

mm long, 1.6-1.9 mm across, shallowly

3-lobate with keels acute, papillose, densely

hairy; hairs evenly distributed over capsule,

spreading or appressed-ascending, to 0.1 mm

long; hypogynous disc entire or laciniate.

Seeds broad-ovate in outline, 0.9-1.2 mm

long, 07-0.8 mm tangentially, 0.7-0.8 mm

radially, tetraquetrous in cross section; dorsal

faces ± planar; ventral faces planar or slightly

concave; all faces with low faint irregular

ridges; exotesta thin, of even thickness, white

to pale brown, microreticulate, becoming

mucilaginous when moistened; endotesta red

brown or pale brown. Fig. 25F.

Additional selected specimens examined: Western

Australia. Karratha, Jul 1981, Craig 240 (PERTH);

Enderby Island, Jun 1980, Onus 72 (PERTH); Depuch

Island, May 1962, Royce 7077 (PERTH); floodplain

of Robe River, c. 8 km SE of Pannawonica, Sep 2003,

Maier & McCreery 511 (BRI); c. 50 km S of Dampier

on Hamersley Iron Railway line road, Apr 1995,

Mitchell PRP225 (PERTH); c. 3 km E of Deepdale

Outcamp where the Robe River crosses the Deepdale to

Millstream Road, Nov 1996, Mitchell PRP1492 (BRI);

Hamersley Ranges, valley of Bungaroo Creek, 15.4 km

SE of Yathala (Old Yalleen) Well, Apr 1995, Trudgen

MET12311 et al. (PERTH); Tambrey Station, Sep 1969,

Brooker 2122 (PERTH); Tombourah Creek, Marble Bar,

Aug 1989, De Jong s.n. (PERTH 4026241); Meentheena

Conservation Reserve, 6.5 km SSE of Meetheena Station

homestead. May 2001, van Leeuwen 4804 (PERTH);

Carawine Gorge, Sep 1991, Wilson & Rowe 913 (NSW);

2.5 km SW of Silver Grass Peak, 30 km NNE of Mt

Farquhar, Jul 1999, Backhouse BEM22 et al. (PERTH);

32.2 miles [c. 51.8 km] from Roebourne turnoff on Tom

Price to Wittenoom Road, Aug 1971, Ashby 4182 (AD,

PERTH); Chichester Range, c. 200 km (215 km by road)

S of Port Hedland, along the road to the Hamersley

Range, Apr 1977, Eichler 22546 (PERTH); Barlee

Range N.R., 3.2 km S of Wongida Well, 12.5 km SW of

Mt Florry, 11.6 km E of Minnie Spring, Barlee Range,

Austrobaileya 8(4): 441-600 (2012)

Aug 1993, van Leeuwen 1502 (PERTH); Barlee Range,

Henry River, Aug 1961, Royce 6535 (PERTH); 5 km W

of Ashburton Downs homestead, Jul 1977, Mitchell 424

(PERTH); Newman, Jul 1981, Deighton 110 (PERTH).

Distribution and habitat : Euphorbia careyi

is confined to the Pilbara, north-western WA,

occurring from the Dampier Archipelago,

south to Barlee Range and east to Carawine

Gorge on the Oakover River (Map 7). It

grows in hummock grassland, shrubland or

low woodland communities on shallow stony,

sandy to clay loam soils, on mostly rocky

scree slopes or stony hillsides.

Phenology : Flowers and fruits have been

collected from February to November

(with most collections made from June to

September).

Notes : Euphorbia careyi can be confused

with E. australis but is distinguished by its

more or less fan-like habit, generally shorter

indumentum, larger involucral glands,

generally green or yellow-green leaves and

dioecy.

The typical form of this species has an

indumentum of short, appressed to ascending

crispate to curved hairs. A less common

variant occurring in the southern half of the

species range has an indumentum of short,

spreading, ± straight hairs (e.g. Ashby 4182 ,

Deighton 110).

The following collections are tentatively

placed here. Although their general leaf

shape, indumentum and involucral glands are

consistent with Euphorbia careyi , they differ

in having shorter styles, darker green leaves

with a reddish tinge along margins and coarser

toothed leaf margins (Robe River, between

Onslow and Roebourne, Aug 1966, Butler 37

(PERTH); Burrup Peninsula, near Karratha,

Aug 2004, Harris WKH2206 (BRI); on track

up Table Top Hill, Karratha, Aug 2004,

Harris WKH2203 (BRI); ‘Grimace Gulch’,

through Maraddoo Ridge, Hamersley Range

N.P, Sep 1978, Trudgen 2296 (PERTH);

Dolphin Island, Dampier Archipelago, May

2000, Trudgen 23472 (BRI); ESE of Whim

Creek, 7.3 km and 1.3 km S of North West

Coastal highway, Jun 2000, Trudgen 23473

(BRI); Burrup Peninsula, 1.4 km almost due

S of Holden Point between Dampier Port and

475

Halford & Harris, Euphorbia section Anisophyllum in Australia

North West Shelf Gas plant, Jun 2000, Long

MET &A026 (BRI).

8. Euphorbia carissoides F.M.Bailey,

Queensland Agric. J. 16: 449 (1906);

Chamaesyce carissoides (F.M.Bailey) P.I.Forst.

& R.J.F.Hend., Novon 5: 323 (1995). Type:

Queensland. [Cook District:] Herberton,

[March 1906,] R.C.Ringrose s.n. (holo: BRI

[AQ342536]; iso: K).

Euphorbia schizolepis var. glabra Benth.,

FI. Austral. 6: 47 (1873). Type: [Queensland.

Cook District:] Gulf of Carpentaria, s.d., DM

[F.Mueller] s.n. (holo: K 186491; iso: G-DC

n.v. [IDC microfiche 800-73. 2416:1. 4], MEL

6122).

Illustration: Forster (1993: 271, fig. 1).

Monoecious, herbaceous woody perennial

with few stems arising from woody taproot,

25-120 cm high, the whole plant glabrous.

Stems ascending to erect; sparingly to much

branched, smooth, often with grey-white

bloom. Interpetiolar stipules subulate,

0.1-2 mm long, entire or deeply bipartite,

glabrous; margin entire or laciniate. Leaves:

petiole 1-2.5 mm long, smooth; blade ovate,

5-36 mm long, 3.5-19 mm wide, 1.4-2

times longer than wide; adaxial and abaxial

surfaces grey-green or blue green often with

red to pink tinge and whitish bloom, smooth;

base symmetric, cordate; margin serrate;

apex acute or obtuse, sometimes apiculate.

Cyathia solitary at the nodes; peduncles

1.5-6.5 mm long. Involucres turbinate,

2-2.5 mm long, 1.5-2.5 mm across; lobes

5, triangular, 0.5-1 mm long, margin entire;

glands 4, shortly stipitate, patelliform, planar

or shallowly concave, transverse-oblong to

reniform in outline, 0.7-0.8 mm long, 1,3—

1.8 mm wide, mostly red; gland appendages

conspicuous, spreading radially, transversely

oblong, 0.7-1.4 mm long, 2-2.5 mm wide,

white turning pink to red with age, glabrous,

margin irregularly dentate; bracteoles 2-2.5

mm long, divided into 2-5 ± linear plumose

segments. Staminate flowers 15-20 per

cyathium; pedicels 1.5-2.5 mm long; staminal

filaments 0.7-0.8 mm long. Pistillate flowers:

styles 1.5-2.3 mm long, connate at the base

into a column for c. 1/3 of their length,

erect, spreading distally, smooth, glabrous,

entire, the apices terete. Capsules exserted

from involucre on pedicel to 5 mm long,

depressed ovate or transversely broad-elliptic

in lateral view, 3.5-4 mm long, 4-5 mm

across, shallowly 3-lobate with keels obtuse,

smooth, glabrous; hypogynous disc entire.

Seeds ovate to broad-ovate in outline, 1.9-2.4

mm long, 1.5-1.9 mm tangentially, 1.6-1.9

mm radially, tetragonous in cross-section;

dorsal faces planar or slightly convex; ventral

faces planar or slightly concave; all faces ±

smooth; exotesta of even thickness, c. 0.1 mm

thick, grey-white, microreticulate, becoming

mucilaginous when moistened; endotesta

brown or red-brown. Fig. 25G.

Additional selected specimens examined: Queensland.

Cook District: Bridge Creek Pastoral Holding, Nov

2000, Stanton 305 (BRI); Blue Hills, Mt Surprise

Gemfields, Jul 1987, Champion 253 (BRI); Blue Hills,

49 km from Mt Surprise township. Mar 1988, Champion

342 (BRI); O’Briens Creek fossicking area, c. 33 km

NW (in straight line) of township of Mt Surprise, Sep

2000, Champion 1673 (BRI); Crystal Creek, Jul 1996,

Ford 1752 (BRI); Pannikin Springs area, 29 km W of

Mungana, Jan 1993, Forster PIF12998 & Bean (BRI);

Pannikin Springs area, Blackdown Station, May 1999,

Forster PIF24369 & Booth (BRI, MEL); loc. cit ., May

2000, ForsterPIF24405 & Booth (BRI, MEL); Aroonbeta

Holding, Walsh River Gorge, near mouth of St Helena

Creek, Jan 1989, Godwin C3159 (BRI); 13.5 km S along

Lappa to Mt Garnet Road, Apr 2005, Forster PIF30791

& McDonald (BRI); 28.8 km S along Lappa to Mt

Garnet Road, Apr 2005, Forster PIF30805 & McDonald

(BRI); Stannary Hills, Aug 1908, Bancroft 207 (BRI);

Newcastle Range, 24 km E of Georgetown, Aug 1997,

Bean 12235 (BRI, MEL); 40 km N of Georgetown, May

1977, Rice 2443 (BRI); O’Briens Creek gemfields via Mt

Surprise, Jul 1994, Coveny 16760 et al. (BRI).

Distribution and habitat: Euphorbia

carissoides is restricted to north-eastern Qld,

occurring from near Georgetown and east to

Stannary Hills, with a disjunct occurrence

near Hopevale (Map 8). It grows on clifflines,

rocky outcrops or hillsides in shrubland or

eucalypt low open woodland communities

on generally shallow soils derived from

sandstone, granite or rhyolite substrates.

Phenology: Flowers and fruits have been

collected in January, March to May, July to

September and November.

Notes: Euphorbia carissoides does not appear

to have any close relatives in Australia and is

easily distinguished from other Australian

476

species of Euphorbia by the following

combination of characteristics: glabrous,

upright woody subshrub; smooth and glabrous

capsules; styles entire; seeds tetraquetrous in

transverse section with a smooth seed coat.

The species is reported to probably be

fire intolerant and only persists in areas not

regularly burnt (Forster 1993).

9. Euphorbia centralis B.G.Thomson,

Nuytsia 8: 353, fig. 2 (1992); Chamaesyce

centralis (B.G.Thomson) P.I.Forst. &

R.J.F.Hend., Novon 5: 323 (1995). Type:

Northern Territory. 3 km SW of Alice Springs,

18 January 1990, B.G.Thomson 3408 (holo:

DNA w.v.; iso: AD n.v., BRI, CANB n.v.,fide

Thomson [1992: 353]).

Illustration : Thomson (1992: 355, fig. 2).

Monoecious (rarely dioecious), annual or

herbaceous perennial with short-lived stems

produced from slender woody taproot, to

30 cm high, few to many stems arising

from rootstock. Stems erect or occasionally

decumbent, much branched, smooth,

moderately dense to densely hispidulous,

pubescent or pilose; hairs spreading or weakly

ascending, straight or crispate, 0.1-1.1 mm

long, white. Interpetiolar stipules narrow-

triangular, 0.2-0.5 mm long, deeply bipartite,

indumentum as for stems; margin lacerate.

Leaves: petiole 0.5-1.3 mm long, smooth,

with indumentum as for stems; blade elliptic

to broad-elliptic, rotund or obovate, 3-7

mm long, 2-5 mm wide, 1.1-2 times longer

than wide; adaxial and abaxial surfaces

dark green to grey green (often with reddish

tinge especially on margin), smooth, with

a sparse to moderately dense indumentum;

hairs spreading, ± straight, 0.4-0.6 mm

long; base asymmetric, one side rounded to

cordate, the other cuneate to obtuse; margin

coarsely serrate or sparsely minutely toothed;

apex rounded. Cyathia solitary at the nodes;

peduncles 0.2-1 mm long. Involucres

turbinate, 0.8-1 mm long, 1-1.1 mm across;

lobes 5, triangular, 0.3-0.5 mm long,

margin entire or laciniate; glands 4, stipitate,

patelliform, shallowly concave, transverse-

oblong to transverse-elliptic in outline, 0.1-

0.3 mm long, 0.4-0.6 mm wide, cream; gland

Austrobaileya 8(4): 441-600 (2012)

appendages conspicuous, spreading radially,

broad-obovate, 0.4-1.3 mm long, 0.7-1.3

mm wide, pink to red, pale green or cream,

glabrous, margin deeply laciniate; bracteoles

0.9-1 mm long, adnate for 1/3 of their length to

involucre, free portion divided into numerous

subulate glabrous segments. Staminate

flowers 15-20 per cyathium; pedicels c. 1.1

mm long; staminal filaments 0.1-0.2 mm long.

Pistillate flowers: styles 0.4-0.6 mm long,

connate at the base into a column for c. 1/10

of their length, ascending, recurved distally,

smooth, glabrous, each scarcely bifid or

divided c. 1/2 of their length, the apices terete.

Capsules exserted from involucre on pedicel

to 3.5 mm long, elliptic or rarely broad to very

broad-ovate in lateral view, 1.5-2 mm long,

1.5-2 mm across, shallowly 3-lobate with

keels acute, smooth or minutely papillose,

sparsely to densely hairy; hairs mostly evenly

distributed over capsule rarely confined to

keels, spreading, 0.1-1 mm long; hypogynous

disc entire. Seeds ovate in outline, 1.3-1.6 mm

long, 0.7-0.8 mm tangentially, 0.7-0.9 mm

radially, tetraquetrous in cross section; dorsal

faces flat to concave; ventral faces concave;

all faces with low faint to distinct irregular

ridges; exotesta thin, of even thickness, white

or pale brown, microreticulate, becoming

mucilaginous when moistened; endotesta red-

brown. n= 11. Fig. 25H.

Additional selected specimens examined : Western

Australia. Pass of the Abencerrages, Rawlinson

Range, Jul 1963, George 4888 (PERTH); near N base

of Mt Aloysius, Jul 1963, George 5229 (PERTH); near

Mt Squires, Barrow Range, Aug 1891, Helms s.n. (AD

96832153, MEL 2178635). Northern Territory. 66.3

km E of Stuart Highway towards ‘Epenarra’, May 2005,

Bean 23798 (BRI); 22 miles [c. 35 km] SSW of Georgina

Downs Station, Mar 1953, Perry 3471 (BRI, DNA); 20

miles [c. 32 km] NNE [of] Huckitta homestead, Jul 1970,

Latz 670 (AD, DNA, MEL); 1 km SW Muranji Rockhole,

Mt Winter, Oct 1986, Thomson 1552 (AD, DNA);

Chewings Range, N face of range across from Giles Yard

Spring, Feb 1990, Thomson 3434 (DNA); Harts Range,

E-end, Plot 1216, May 1988, Thomson 2413 (DNA,

MEL); Penny Springs, Kings Canyon, Jul 1985, Leach

680 (DNA); 12.5 km N [of] Tempe Downs homestead,

Aug 1988, Barritt 324 (DNA, MEL); Atherrita Bore,

52 miles [c. 84 km] SE [of] Todd River homestead,

Aug 1964, Nelson 1303 (AD, BRI, DNA, NSW); Mann

Range, 30 miles [c. 48 km] NE [of] Mt Davies Camp,

Oct 1970, Latz 908 (AD, DNA); 35 km E [of] Horseshoe

Bend homestead, Nov 1993, Albrecht 5571 (AD, DNA,

MEL). Queensland. Gregory North District: 21 km

477

Halford & Harris, Euphorbia section Anisophyllum in Australia

WNW of‘Marion Downs’, Sep 1978, Purdie 1336 (BRI);

Glenormiston, Toko Range, c. 1.8 km S of S-bend Gorge,

Mulligan River on track to Cravens Peak homestead, Jun

2010, Halford Q9867 & Forster (BRI). South Australia,

between Deering Hills and Mann Ranges, c. 18 km NE

of Mt Cooperinna, Sep 1978, Barker 3437 (AD, NSW); at

foot of Mt Harriet, Sep 1963, Whibley 947 (AD); 7 km E

of Marys Well, De Rose Hill Station, Aug 1992, Badman

6108 (AD); Beresford Hill, Oct 1978, Chorney 992 (AD,

NSW); 2 km SW of Bulgunnia Station homestead, Nov

1992, Badman 6572 (AD).

Distribution and habitat: Euphorbia centralis

occurs in central Australia from Tennant

Creek, NT, south to Bulgunnia Station (S of

Coober Pedy), SA and from Cavanagh and

Barrow Ranges, WA east to Boulia, western

Qld with a disjunct occurrence near Quilpie,

south-western Qld (Map 9). It grows mostly

in hummock grassland or Acacia shrubland

communities on rocky scree slopes or stony

hillsides. The soils are sandy, and often

shallow and rocky, derived from granite,

sandstone or limestone substrates.

Phenology: Flowers and fruits have been

collected throughout the year, particularly

from March to October.

Notes: Euphorbia centralis is similar to E.

australis and can be difficult to distinguish

from E. australis var. erythrantha in northern

SA where these taxa distributions overlap.

Mature plants of Euphorbia centralis exhibit

a rounded shrub-like habit and have gland

appendages 0.4-1.3 mm long that are deeply

laciniate (versus 04-0.2(0.4) mm long and

shallowly to deeply toothed for E. australis

var. erythrantha).

10. Euphorbia cinerea W.Fitzg., J. & Proc.

Roy. Soc. Western Australia 3: 161 (1918).

Type: Western Australia. Devil’s Pass

[Windjana Gorge], Napier Range, May 1905,

W.VFitzgerald 591 (holo: PERTH).

Monoecious, herbaceous perennial with many

short-lived stems produced from slender

taproot, the whole plant glabrous. Stems

prostrate, sparingly branched, longitudinally

ridged. Inter petiolar stipules subulate,

0.6-0.8 mm long, deeply bipartite, glabrous;

margin laciniate. Leaves: petiole 0.2-0.7

mm long, smooth; blade oblong or obovate,

4-8 mm long, 2.6-5 mm wide, 1.6-2 times

longer than wide, smooth, pale green; base

asymmetric with one side cordate, the other

obtuse; margin sparsely minutely toothed

distally; apex rounded to retuse. Cyathia

solitary at nodes, often clustered on short

leafy lateral branchlets; peduncles 0.1-0.2 mm

long. Involucres campanulate or turbinate,

0.8-1 mm long, 0.6-0.8 mm across; lobes 5,

triangular, 0.2-0.5 mm long, margin entire

or serrulate; glands 4, stipitate, cupuliform,

with distinct central pit, transverse-oblong or

transverse-elliptic in outline, c. 0.1 mm long,

0.2-0.4 mm wide, pink; gland appendages

conspicuous, spreading radially, reniform,

0.1-0.3 mm long, 0.4-0.8 mm wide, pink to

white, glabrous, margin entire or shallowly

lobed; bracteoles 0.6-0.8 mm long, entire or

divided into few subulate glabrous segments.

Staminate flowers 1-5 per cyathium; pedicels

0.9-1.1 mm long; staminal filaments 0.1-0.3

mm long. Pistillate flowers: styles 0.3-0.4

mm long, spreading, smooth, glabrous, entire,

the apices terete. Capsules exserted from

involucre on pedicel to 1.5 mm long, broad

to very broad-ovate in lateral view, 1.1-1.3

mm long, 1.3-1.5 mm across, shallowly

3-lobate with keels acute, smooth, glabrous;

hypogynous disc entire. Seeds ovate in outline,

0.7-0.8 mm long, 0.4-0.5 mm tangentially,

0.4-0.5 mm radially, tetraquetrous in cross

section; dorsal faces slightly concave; ventral

faces concave; all surfaces smooth or with

1 or 2 low faint ridges; exotesta thin, of

even thickness, grey-white, microreticulate,

becoming mucilaginous when moistened;

endotesta pale brown or red-brown. Fig. 251.

Additional selected specimens examined : Western

Australia, upper reaches of Barker River, 2 km N of Mt

Hart homestead, Jun 1987, Edinger 421 (PERTH); Black

Stone Mine, Leopold Range, May 1988, Cranfield 6728

(PERTH); 3 km S of Karriwell Bore; 100 km E of Derby,

Jun 1988, Wilson 12788 (PERTH); Windjana Gorge N.P,

Aug 1982, Conrick 1083 (AD); Camballin Road, 14.7 km

5 of junction with Derby - Fitzroy Crossing Road, Apr

1985, Aplin 128 etal. (PERTH); Windjana Gorge, Napier

Range, Jul 1974, Carr 3865 & Beauglehole ACB47643

(NSW, PERTH); 1 mile [c. 1.6 km] from Calwynyardah

turnoff, Fitzroy road. Mar 1967, Power 311 (PERTH);

c. 6 km SE of Gibb River Road along road to Mt

House Station, May 1985, Aplin 991 et al. (PERTH);

Geikie Gorge N.P., Jul 1974, Carr 3773 & Beauglehole

ACB47551 (NSW, PERTH); Gogo [Station], Apr 1951,

Gardner 10016 (PERTH); E of junction of Kununurra

- Wyndham - Halls Creek road, Jul 1974, Carr 3201

6 Beauglehole ACB46979 (NSW, PERTH); Ord River,

NNW of Goosehole Yard, Bungle Bungle N.P., Jun 1989,

478

Menkhorst 933 (DNA). Northern Territory. Birrindudu

Station, Jun 1994, Egan 3826 (DNA).

Distribution and habitat: Euphorbia cinerea

occurs in the southern Kimberley, WA from

near Derby and eastward to adjacent parts of

the NT (Map 10). It grows on red sand to loam

soils on plains or floodplains in grassy open

shrubland or woodland communities, or in

stony red soils on hills in hummock grassland

communities.

Phenology: Flowers and fruits have been

collected from March to August.

Notes: Euphorbia cinerea resembles E.

albrechtii and E. petala. For features

distinguishing E. cinerea from E. albrechtii

and E. petala , refer to the ‘Notes’ section

under E. albrechtii.

This species has been misidentified as

E. drummondii (e.g. Wheeler et al. 1992). It

is easily distinguished from E. drummondii

by its generally smaller habit, capsules and

seeds, capsule shape and seed sculpturing.

The collection site on the type sheet

(PERTH) is recorded as “Devil’s Pass” rather

than Wingrah Pass, as recorded in Fitzgerald’s

protologue for this species. Royce (in Wilson

1974) confirmed that the names refer to the

same locality. Devil’s Pass is now known as

Windjana Gorge.

11. Euphorbia clementii Domin, Biblioth.

Bot. 89(4): 308-309 (1927 ‘1926’). Type:

[Western Australia.] inter Ashburton et

DeGrey River, in 1900, E.Clement s.n. (holo:

PR 528292).

Monoecious, annual (?) to 60 cm high, few

to many stems arising from base, the whole

plant glabrous. Stems ascending to erect

(rarely semiprostrate, Mitchell PRP288

[PERTH]), sparingly to much branched,

smooth. Interpetiolar stipules subulate,

0.5-1.5 mm long, deeply bipartite, glabrous;

margin entire or laciniate. Leaves: petiole 1-2

mm long, smooth; blade oblong or oblong-

obovate, 12-18 mm long, 5-9 mm wide, 2.1-

2.3 times longer than wide; adaxial surface

green, smooth; abaxial surface pale green,

smooth; base symmetric, rounded; margin

entire; apex rounded. Cyathia solitary at

Austrobaileya 8(4): 441-600 (2012)

the nodes, mostly on distal portion of stems.

Involucres cupuliform, 1.5-2 mm long, 1.8-

3.5 mm across; lobes 5, oblong or obovate,

1-1.3 mm long, margin deeply laciniate

distally; glands 4, sessile, appressed to abaxial

surface of involucre, patelliform, planar,

transverse-elliptic to orbicular in outline,

0.8-1.1 mm long, 0.7-1.4 mm wide, yellowish

green; gland appendages absent; bracteoles

1.3-1.6 mm long, divided into numerous ±

linear hirsute segments. Staminate flowers

60-75 per cyathium; pedicels 1.5-2 mm

long; staminal filaments 0.4-0.8 mm long.

Pistillate flowers: styles 0.7-1 mm long,

ascending, smooth, glabrous, each bifid

for c. 1/2 of their length, the apices clavate.

Capsules exserted from involucre on pedicel

to 6 mm long, broad-elliptic or transversely

broad-elliptic in lateral view, 2.7-3 mm long,

3.2-3.5 mm across, deeply 3-lobate with

keels obtuse, smooth, glabrous; hypogynous

disc entire. Seeds oblong or elliptic to broad-

elliptic in outline, 1.9-2 mm long, 1.3-1.5 mm

tangentially, 1.3-1.5 mm radially, trigonal

to suborbicular in cross section; dorsal and

ventral faces convex, smooth; exotesta thin, of

even thickness, grey-white, microreticulate,

becoming mucilaginous when moistened;

endotesta red-brown to brown. Fig. 25J.

Additional selected specimens examined: Western

Australia. 20 km E [of] De Grey River crossing on

Port Hedland/Broome Road, May 1991, Thomson 3498

(AD, BRI, NT); HGM site 4, Y9 Crustal Deposit, Yarrie

Iron Ore Mine site, c. 190 km E of Port Hedland, Apr

1998, Maier s.n. (PERTH 5086590); Gorge Creek,

on Port Hedland - Marble Bar Road, Apr 2006, Bean

25126 (BRI); Taiga Gap, S of Coongan Siding, Jun 1941,

Burbidge 1053 (PERTH); 15.8 km N of Taiga, Sep 1986,

Chinnock 6987 (AD); c. 37 km SE of Yandeyarra on

southern access track from Great Northern Highway,

Apr 1995, Mitchell PRP288 (PERTH).

Distribution and habitat: Euphorbia

clementii is confined to the Pilbara, north¬

western WA, occurring in an area more or

less bounded by Port Hedland, Marble Bar

and the Mungaroona Range (Map 11). It

grows in Triodia grassland communities, on

gravelly clay loam soils on slopes or stony

rises or on sandy soils on plains. It is often

noted on specimen labels to be growing in

areas “recently burnt”.

479

Halford & Harris, Euphorbia section Anisophyllum in Australia

Phenology : Flowers and fruits have been t

collected from April to June. £

Notes: Euphorbia clementii is a distinctive

species but appears most closely related to E. ,

biconvexa, E. coghlanii and E. trigonosperma. d

It is easily distinguished from all three species j.

by its large, sessile, disk-like involucral

glands that are appressed to the outside of

the involucral cup (versus shortly stipitate c

involucral glands spreading radially from g

the involucral rim and perpendicular to the j

outside of the involucral cup for E. biconvexa ,

E. coghlanii and E. trigonosperma ), involucral

lobes oblong or obovate, 1-1.3 mm long ^

(versus triangular or subulate and 0.3-1.1 mm g

long for E. biconvexa , E. coghlanii and E.

trigonosperma) and more numerous staminate

flowers per cyathia (60-75 per cyathia versus

up to 35 per cyathia for E. biconvexa , E.

coghlanii and E. trigonosperma). ^

12. Euphorbia coghlanii F.M.Bailey, 2

Queensland Dept. Agric. Stock Bot. Bull. 13: s

12 (1896); Chamaesyce coghlanii (F.M.Bailey) c

D.C.Hassall ex P.I.Forst. & R.J.F.Hend., \

Nov on 5: 323 (1995). Type: Queensland. I

Gregory North District: Roxborough, c

Georgina River, December 1895, F.M.Bailey c

s.n. (holo: BRI [AQ342538]); Epitype (here £

chosen): Queensland. Burke District: 20 km f

(by road) E of Musselbrook Mining Camp C

on road to Ridgepole Waterhole, 175 km N C

of Camooweal - Lawn Hill National Park, c

27 April 1995, R.W.Johnson MRS379 & :

M.B. Thomas (BRI). e

Monoecious, annual or herbaceous perennial ^

with short-lived stems produced from slender

woody taproot, to 80 cm high, few to many

stems arising from rootstock. Stems erect,

sparingly to much branched, smooth, ( -

glabrous or with a sparse to moderately dense

indumentum (pubescent or pilose); hairs ±

appressed or spreading, crispate, 0.1-0.4 mm

long, white. Interpetiolar stipules narrow-

triangular or subulate, 0.6-1.5 mm long, £

entire or bipartite, glabrous; margin entire |

or laciniate. Leaves: petiole 0.7-2 mm long,

smooth, glabrous or with indumentum as /

for stems; blade oblong, lanceolate, narrow- {

ovate to broad-ovate or oblong-elliptic, 14-36 g

mm long, 2-10 mm wide, 2.2-9 times longer ^

than wide; adaxial surface subglaucous, dull

green or pale green (often with reddish tinge),

smooth, glabrous; abaxial surface glaucous

or pale green, smooth, glabrous or sparsely

hairy with appressed-ascending to ascending,

± straight or crispate hairs 0.1-1 mm long;

base asymmetric with one side cordate, the

other rounded; margin entire or serrulate;

apex acute to rounded. Cyathia solitary at

the upper nodes, sometimes clustered on

short leafy lateral branchlets with subtending

leaves slightly smaller than the primary stem

leaves; peduncles 1-9 mm long. Involucres

campanulate or turbinate, 0.8-1.2 mm long,

1.1-1.3 mm across; lobes 5, triangular to

subulate, 0.4-0.5 mm long, margin entire

or laciniate; glands 4, stipitate, patelliform,

planar to concave (rarely cupuliform with

distinct central pit), transverse-elliptic to ±

orbicular in outline, 0.1-0.2 mm long, 0.1-

0.6 mm wide, red or yellowish green; gland

appendages conspicuous to inconspicuous,

spreading radially, very broad-obovate to

obdeltoid, 0.1-0.5 mm long, 0.5-0.9 mm

wide, red or white, glabrous, margin entire;

bracteoles 0.7-0.8 mm long, adnate for c. 1/4

of their length to involucre, free portion entire

or divided into subulate glabrous segments.

Staminate flowers 5-25 per cyathium;

pedicels 1.1-1.4 mm long; staminal filaments

0.2-0.3 mm long. Pistillate flowers: styles

0.6-0.8 mm long, erect to spreading, recurved

distally, smooth, glabrous, each bifid for 1/3—

2/3 of their length, the apices terete. Capsules

exserted from involucre on pedicel to 4 mm

long, transversely broad-elliptic in lateral

view, 1.8-2 mm long, 1.8-2.4 mm across,

deeply 3-lobate with keels obtuse, smooth,

glabrous; hypogynous disc entire. Seeds

oblong to ovate in outline, 1.2-1.4 mm long,

0.8-1 mm tangentially, 0.8-1 mm radially,

suborbicular in cross section; dorsal and

ventral faces convex, smooth; exotesta thin, of

even thickness, grey-white, microreticulate,

becoming mucilaginous when moistened;

endotesta red-brown to dark brown, n = 8.

Fig. 25K.

Additional selected specimens examined: Western

Australia. 10 km from Kununurra on road to Ivanhoe

Springs, Apr 1985, Aplin 568 et al. (PERTH); 15 km

S of Victoria Highway, 40 km E of Kununurra along

Victoria Highway, Apr 1989, Halford H51 (BRI);

480

Barrow Island, Oct 1980, Buckley 6901 (PERTH); 2.4

km WSW of Coondewanna Hill, Hamersley Ranges, Jun

1997, Trudgen 16673 (PERTH); c. 1 km S of Newman

on Great Northern Highway, Mar 1994, Mitchell PRP120

(PERTH); 1 km SSW of Erallinga Pool, Hamersley

Ranges, Apr 1997, Trudgen 15267 (PERTH); 4.75 km

ESE of West Angela Hill, Hamersley Ranges, Jun 1997,

Trudgen 16729 (PERTH). Northern Territory. 2 km

NW of Timber Creek, Dec 1994, Barritt 1625 (DNA,

MEL); Longreach Waterhole, Elliott, Feb 1988, Thomson

2187 (AD, BRI); Muckaty, near homestead. Mar 1955,

Chippendale 1071 (BRI, DNA); Brunette Downs Station

on stockroute to racecourse, Aug 1986, Henshall 4127

(AD, DNA); 5 miles [c. 8 km] W of Frewena Roadhouse,

Jul 1971, Henry 194 (BRI); Soudan Station, No. 103 Bore,

Nov 1986, Henshall 4132 (DNA). Queensland. Burke

District: Beames Brook on Burketown to Camooweal

Road, May 2000, Jago 5699 & Wannan (BRI); 67 km

WNW of Mt Isa, 6 km N of Mingera Creek, Apr 1989,

Harris 335 (BRI); Roxmere Station, S of Cloncurry,

Apr 2003, Booth 3179 & Kelman (BRI); 2 km by road

from Hughenden towards Winton, Apr 2006, Halford

Q9015 & Batianoff (BRI). Mitchell District: 30 miles

[c. 48 km] W of Blackall, May 1975, Hassall 7530 (BRI).

Gregory North District: BladensburgN.P., S of Winton

and 2 km NE of Bladensburg homestead. Mar 1998,

Forster PIF22278 & Booth (BRI, MEL). Leichhardt

District: 15 km SE of Capella, Mar 1995, Fensham 2572

(BRI); 16 miles [c. 26 km] NW of Rolleston, May 1975,

Hassall 7536 (BRI).

Distribution and habitat : Euphorbia

coghlanii occurs across northern Australia

from the Pilbara and Kimberley, WA,

extending through the northern central region

of the NT, and eastward to central Qld (Map

12). It grows in grassland or open woodland

communities, on mostly dark cracking clay

soils rarely on sands, on plains, alluvial flats

or along drainage lines. It has been rarely

recorded in white coastal sands {Buckley 6901

[PERTH]).

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from January to August.

Notes : Euphorbia coghlanii is similar to E.

biconvexa and E. trigonosperma but differs in

having seeds that are suborbicular in transverse

section versus lenticulate for E. biconvexa and

trigonous for E. trigonosperma. It also differs

from E. trigonosperma in habitat preference

as well. Euphorbia coghlanii inhabits mostly

dark cracking clay soils on plains, alluvial

flats compared to sandy soils on coastal or

inland dunes, in well-drained sandy soils on

stony slopes of sandstone and limestone hills,

Austrobaileya 8(4): 441-600 (2012)

and in sandy alluvium along drainage lines.

The holotype of Bailey’s Euphorbia

coghlanii is in poor condition. It currently

consists of a few almost leafless stems as

well as a fragment packet containing a few

fragmented leaves and stems, and a single

cyathium. The Bailey collection (BRI

[AQ342538]) as well as the illustration with

the protologue of E. coghlanii lack sufficient

details to fix the application of the name. As

there is no other original material available

to clarify the application of E. coghlanii , an

epitype is here nominated. The collection

Johnson MRS379 & M.B. Thomas (BRI [AQ

587084]) is chosen here as epitype. This

collection has morphology that is similar

to the remaining fragments of the original

material and matches the description in the

protologue.

13. Euphorbia crassimarginata Halford &

W.K.Harris, species nova similis generatim

E. drummondii Boiss. sed ab ea glandulis

plerumque majoribus (0.2-0.3 x 0.4-0.5 mm)

ore manifestioribus incrassatis appendicem

petaloidam carens (ad vicem 0.15-0.2 x 0.2-

0.4 mm glandulae appendicibus transverso-

linearis usque 1 mm longis) stylis 0.4-0.5

longis bifidis apicibus teretibus (ad vicem

0.2-0.3 mm longis integris vel vix bifidis)

differt. Typus: Queensland. Burke District:

31 km by road from junction of Burke and Gulf

Development Roads towards Flinders River,

16 January 2005, K.R.McDonald KRM3414

(holo: BRI, iso: DNA, distribuendi ).

Monoecious, annual to 40 cm high, with

few stems arising from fibrous rootstock,

the whole plant glabrous. Stems erect

(rarely prostrate, Pullen 8875 [BRI]),

sparingly branched, longitudinally ridged.

Interpetiolar stipules narrow-triangular,

0.7-1 mm long, deeply bipartite, glabrous;

margin laciniate. Leaves: petiole 0.5-0.8 mm

long, smooth; blade narrow-oblong or narrow-

ovate, sometimes falcate, 7.5-17 mm long,

1.7-3.4 mm wide, 3.6-5.9 times longer than

wide; adaxial and abaxial surfaces green to

pale green often with reddish tinge especially

along margins, smooth or minutely papillose;

base asymmetric with one side obtuse, the

other rounded; margin sparsely minutely

481

Halford & Harris, Euphorbia section Anisophyllum in Australia

toothed distally; apex obtuse or acute.

Cyathia solitary at nodes, often clustered on

short leafy lateral branchlets; peduncles 0.4-1

mm long. Involucres turbinate, 0.7-0.8 mm

long, 0.8-0.9 mm across; lobes 5, triangular,

0.4-0.5 mm long, margin fimbriate; glands

4, stipitate, cupuliform, with distinct central

pit and thicken rim, transverse-elliptic or

orbicular in outline, 0.2-0.3 mm long, 0.4-0.5

mm wide, red to purple; gland appendages

absent; bracteoles subulate, 0.5-0.6 mm long,

adnate for c. 1/3 of their length to involucre,

glabrous. Staminate flowers 5 per cyathium;

pedicels 0.7-0.8 mm long; staminal filaments

c. 0.1 mm long. Pistillate flowers: styles 0.4-

0.5 mm long, spreading, smooth, glabrous,

each bifid for 1/3-2/3 of their length, the

apices terete. Capsules exserted from

involucre on pedicel to 1.8 mm long, broad-

elliptic in lateral view, 1.7-1.9 mm long, 1.6-

1.8 mm across, shallowly 3-lobate with keels

obtuse, smooth, glabrous; hypogynous disc

entire. Seeds oblong-ovate in outline, 1.2-1.5

mm long, 0.7-0.8 mm tangentially, 0.7-0.8

mm radially, tetraquetrous in cross section;

dorsal faces planar; ventral faces concave;

all faces with 3-6 faint to distinct transverse

ridges; exotesta of uneven thickness,

distinctly thicker on transverse ridges, white

to pale brown, microreticulate, becoming

mucilaginous when moistened; endotesta red-

brown. n = 11. Figs 3, 25L.

Additional selected specimens examined : Northern

Territory. 27 km SW [of] Daly Waters, Feb 1989,

Thomson 3147 (DNA); 22 km N [of] Ucharonidge

homestead, Feb 1989, Thomson 3246 (AD); Tanumbirini

Station, Carpentaria Highway, Feb 1988, Thomson 2203

(DNA). Queensland. Burke District: c. 20 miles [ c. 32

km] W of Burketown, Jul 1974, Hassall 7472 (BRI); c.

15 km SW of Normanton, along the road to Mogoura

Station, Apr 1974, Pullen 8875 (BRI); 28.4 km by road

towards Flinders River from junction of Burke and Gulf

Development Roads, Jan 2005, McDonald KRM3449

(BRI); 5 km S of Normanton, May 1976, Hassall 7633

(BRI); 15 km S of Normanton on Cloncurry road, 200

m W of highway, Apr 2004, Thompson BUR157 &

Newton (BRI); near Eight Mile Creek, c. 40 km NNE of

Normanton, on track c. 2.5 km off Burke Developmental

Road, Jun 2001, Turpin GPT873 & Thompson (BRI); c.

70 miles [c. 113 km] S of Burketown, Jul 1974, Hassall

s.n. (BRI [AQ475167]); 17.5 km N of Donors Hill

turnoff, on Cloncurry - Normanton Road, May 1976,

Farrell TF414 (BRI); Bang Bang Jumpup on Cloncurry

-Normanton Road, 105 km S of Normanton, Mar 1977,

Farrell TF793 (BRI); 90 km NNW of Mt Isa on remote

station track. May 2006, Booth CAM08-10 & Kelman

(BRI); 54.7 km by road S of Flinders River road bridge.

Mar 2005, McDonaldKRM4047 (BRI); 125.7 km by road

S of Flinders River road bridge. Mar 2005, McDonald

KRM4059 (BRI). Mitchell District: Ventcher Station,

82 km SW of Torrens Creek, Apr 2005, Thompson

TAN208 & Booth (BRI); 83 km SSE of Torrens Creek on

Jednock Station, Apr 2005, Thompson TAN236 & Booth

(BRI).

Distribution and habitat : Euphorbia

crassimarginata extends from Daly Waters,

NT, eastward to near Hughenden, Qld (Map

13). It grows in eucalypt or Melaleuca open

woodland, or Acacia woodland communities,

on plains or low lateritic hills, and in Mitchell

grassland on plains. The soils are variable in

texture: sand, clay loam to cracking clays.

Phenology : Flowers and fruits have been

collected from January to July.

Notes : Euphorbia crassimarginata bears a

general resemblance to E. drummondii but

differs from it by having generally larger

glands (0.2-0.3 x 0.4-0.5 mm) with more

pronounced thickened rim and lacking a

petaloid appendage (versus 0.15-0.2 x 0.2-0.4

mm, gland appendages present, transverse-

linear, to 0.1 mm long for E. drummondii ), and

styles 0.4-0.5 mm long, each bifid for 1/3-2/3

of their length, the apices terete (versus 0.2-

0.3 mm long, each entire or scarcely bifid, the

apices clavate for E. drummondii ).

Etymology: The specific epithet is from Latin

crassus , thick, and - marginatus , margined,

in reference to the thickened margin of the

involucral glands of the species.

14. Euphorbia dallachyana Baill .,Adansonia

6: 285-286 (30 July 1866); Chamaesyce

dallachyana (Baill.) D.C.Hassall, Aust. J.

Bot. 24: 640 (1976). Type: Queensland.

[Port Curtis District:] Rockhampton, s.d.,

[J.Dallachy s.n.] (holo: P 313101; iso: MEL

68199).

Euphorbia minutifolia Boiss., in A.DC.,

Prodr. 15(2): 1263 (late August 1866).

Type: [Queensland. Cook District:] Gulf

of Carpentaria, s.d., [W.Landsborough s.n]

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

2421: II. 8]; iso: MEL 68205, 68204).

Illustration: Auld & Medd (1987: 160).

482

Austrobaileya 8(4): 441-600 (2012)

Fig. 3. Euphorbia crassimarginata. A. habit *0.6. B. branchlet with cyathia *4. C. leaf *4. D. stipules *12. E. cyathia

with female flower x32. F. capsule with cyathia *16. G. cyathial gland, adaxial view x32. H. capsule, top view xl6.1.

capsule, lateral view xl6. A from McDonaldKRM3414 (BRI); B-I from McDonaldKRM3449 (BRI). Del. W.Smith.

483

Halford & Harris, Euphorbia section Anisophyllum in Australia

Monoecious, annual or herbaceous perennial

with short-lived stems produced from slender

woody taproot, to 5(20) cm high, many

stems arising from rootstock, the whole plant

glabrous. Stems prostrate (rarely ascending to

erect), much branched, smooth. Interpetiolar

stipules triangular to broad-triangular, 0.2-

1.1 mm long, entire or bipartite, glabrous;

margin lacerate. Leaves: petiole 0.5-1.5 mm

long, smooth; blade oblong, ovate, obovate

to broad-obovate, elliptic to broad-elliptic or

subrotund, 2-9 mm long, 1.3-7 mm wide, 1-2

times longer than wide; adaxial and abaxial

surfaces green or blue-green (sometimes

tinged red especially along margins),

smooth (rarely minutely papillose); base

asymmetric with one side rounded to cordate,

the other obtuse to rounded; margin entire

or minutely toothed distally; apex obtuse,

mucronate or retuse. Cyathia solitary at the

nodes; peduncles 0.5-5 mm long, smooth.

Involucres turbinate, 0.6-0.9 mm long, 0.5-1

mm across; lobes 5, triangular to subulate,

0.2-0.4 mm long, margin fimbriate; glands 4,

stipitate, cupuliform, with a shallow central

pit, transverse-oblong, or transverse-elliptic

in outline, 0.1-0.2 mm long, 0.2-0.4 mm

wide, pink or pale green; gland appendages

conspicuous to inconspicuous, spreading

radially, transverse-linear, 0.05-0.3 mm long,

0.2-0.6 mm wide, pink or white, glabrous,

margin entire or shallowly lobed; bracteoles

0.5-0.8 mm long, adnate for 1/3 of their

length to involucre, free portion divided into

few to numerous subulate glabrous or hirsute

segments. Staminate flowers 5-10 per

cyathium; pedicels 0.6-1 mm long; staminal

filaments 0.1-0.2 mm long. Pistillate flowers:

styles 0.1-0.4 mm long, spreading to erect,

smooth, glabrous, bifid for up to 1/2 of their

length, the apices clavate or terete. Capsules

exserted from involucre on pedicel to 2.2 mm

long, transversely broad-elliptic in lateral

view, 1.5-2.2 mm long, 1.5-2.4 mm across,

shallowly 3-lobate with keels obtuse, smooth;

hypogynous disc laciniate or entire. Seeds

ovate or broad-ovate in outline, 1.1-1.5 mm

long, 0.8-0.95 tangentially, 0.7-0.9 mm

radially, tetraquetrous or tetragonous in

cross section; dorsal and ventral faces planar

to convex, with 2-4 faint transverse ridges;

exotesta thin, of even thickness, grey-white,

microreticulate, becoming mucilaginous

when moistened; endotesta brown or red-

brown. n - 11, 22. Fig. 25M.

Additional selected specimens examined : Western

Australia. Fitzroy River, Great Northern Highway,

Broome - Derby Road, Jul 1974, Carr 4305 &

Beauglehole 48083 (NSW; PERTH); corner of Kargotich

& Mundijong Roads, Mundijong, May 2009, Hislop

3880 (BRI, MICH). Northern Territory. Dashwood

Crossing, Jul 1995, Booth 1031 (DNA); Kalkaringi,

Mar 1990, Thomson 3492 (DNA); Lake Nash Station,

Beantree Bore, Jul 2000, Risler 472 (DNA); between

Top Springs & Timber Creek, Jul 1974, Carr 2739 &

Beauglehole 46518 (MEL, NSW). Queensland. Burke

District: Gregory River crossing, Riversleigh Station,

May 1990, Latz 11639 (DNA). South Kennedy Disctrict:

5 km from Moray Downs homestead on Clermont Road,

May 1995, Forster P1F16701 & Figg (BRI). Port Curtis

District: GlenGeddes, c. 45 kmNNW of Rockhampton,

Nov 2003, Halford Q8076 & Forster (BRI, MEL, NSW).

Moreton District: Blenheim, W of Laidley, Dec 2004,

Forster PIF30528 (BRI); White Swamp Road, S of

Boonah, Mar 2004, Bean 21752 (BRI). South Australia.

c. 6 km S of Blanchetown, Feb 1973, Weber 3521 (AD);

c. 1 km NW of Big Bend S of Swan Reach, Apr 1973,

Donner 4123 (AD); Adelaide Botanic Gardens, Car

Park, Jan 1992, Smith 2958 (AD). New South Wales. 0.4

km S of Gurley Creek, on Moree - Narrabri Road, Jan

1996, Bean 9509 (BRI, MEL, NSW); alongside Ironbark

Creek, W of Woodsreef Mine, Mar 1994, Hosking 958

(MEL, NSW); Buchanan Lamington Colliery, 4.8 km

from Warkworth, Jun 1975, Coveny 6554 & Powell

(NSW); 10 km W of Balranald, Dec 1987, Thomson

2300 (NSW [mixed collection with E. serpens ], NT);

20.9 km SE along Aratula Road towards Tocumwal off

the Deniliquin - Finley Road, Apr 1988, Coveny 12903

etal. (MEL, NSW). Victoria. 1.5 km NE [of] Colignan,

Apr 1983, Forbes 1416 (MEL); Johnson’s Bend, Murray

River near Mildura, May 1969, Craven 1575 (MEL);

Yarrawonga, Jan 1982, Aston 2180 (MEL).

Distribution and habitat: Euphorbia

dallachyana is widespread in eastern

Australia with scattered localities in northern

and south-western WA, and north-western

NT (Map 14). It grows in grassland, eucalypt

woodland/open forest communities on plains,

alluvial flats or river and creek banks, rarely

on rocky hillsides. The soils vary from sand

to cracking clays. It is often recorded in

disturbed areas such as roadsides, in urban

parklands, along railways, pastures and

cultivated land.

Phenology: Flowers and fruits have been

collected throughout the year, particularly

from November to May.

484

Notes : Euphorbia dallachyana is

characterised by the lack of an indumentum;

stems not longitudinally ridged; stipules

triangular to broad-triangular; glands

cupuliform with a shallow central pit; gland

appendages always present although small

(to 0.3 mm long) and transverse-linear

with margin entire or shallowly lobed; seed

surfaces with 2-4 faint transverse ridges;

capsules transversely broad-elliptic in lateral

view.

Bentham (1873) placed Euphorbia

dallachyana in synonymy under E.

drummodii. Hassall (1976) reinstated it to

specific rank (as Chamaesyce dallachyana ).

Euphorbia dallachyana is similar to E.

drummondii in its involucral glands, gland

appendages, and capsule shape and size. It

differs from that species in its leaf form and

stipule form, and seed surface sculpturing.

Euphorbia dallachyana as circumscribed

here is variable in style and hypogynous disc

morphology. The typical variant occurs widely

throughout the species distributional range

although it is more common in the northern

part of the range. It has styles bifid for c. 1/2

of their length with the apices ± terete and the

hypogynous disc with a laciniate fringe. The

hypogynous fringe is not always present at the

base of all capsules on the one plant and can

vary from just a few slender subulate lobes to

a much divided broad lobe (e.g. Hislop 3880 ,

Hosking 958 , Smith 2958 , Halford Q8076 &

Forster ). Hassall (1977) records this typical

variant as a tetraploid with a base chromosome

number n = 22.

A second variant occurring in the southern

part of the species range (through NSW, SA,

Vic and WA) has styles that are bifid for up to

1/3 of their length with ± erect, stout clavate

apices and the hypogynous disc undivided

(e.g. Donner 4123 , Craven 1575 , Coveny

12903 et al ., Aston 2180). This variant was

referred to E. drummondii (as Chamaesyce

drummondii) by Hassall (1977) who recorded

a chromosome number n — 11. We believe

that this variant is better placed under E.

dallachyana as it is morphologically similar

in seed sculpturing, and stipule and leaf

form. Further collections and field studies are

Austrobaileya 8(4): 441-600 (2012)

warranted to establish the significance of this

variation.

15. Euphorbia drummondii Boiss., Cent.

Euphorb. 14 (1860); E. drummondii Boiss.

var. drummondii , Baillon, Adansonia 6: 285

(1866); Chamaesyce drummondii (Boiss.)

Sojak, Cas. Ndr. Muz., Odd. Prlr. 140: 169

(1972). Type: [Western Australia.] Swan

River, s.d., [J] Drummond 670 (holo: G

191672 n.v. [image seen]; iso: G 191671 n.v.

[image seen], K 186497, P 313102).

Monoecious, annual or herbaceous perennial

with short-lived stems produced from slender

woody taproot, to 15 cm high, few to many

stems arising from rootstock, the whole

plant glabrous. Stems prostrate or erect,

sparingly branched, longitudinally ridged.

Interpetiolar stipules subulate, 0.5-0.9

mm long, deeply bipartite, glabrous; margin

laciniate. Leaves: petiole 0.5-1.1 mm long,

smooth; blade narrow-oblong to oblong, broad-

ovate or oblanceolate , sometimes slightly

falcate, 4.5-16 mm long, 2.3-5 mm wide, 1.7-

5 times longer than wide; adaxial and abaxial

surfaces blue-green or red, smooth (rarely

minutely papillose); base asymmetric with

one side cordate, the other obtuse to rounded;

margin serrulate; apex rounded. Cyathia

solitary at the upper nodes, often clustered

on short leafy lateral branchlets; peduncles

c. 0.7 mm long. Involucres campanulate,

0.6-0.7 mm long, 0.8-1 mm across; lobes 5,

triangular, 0.4-0.5 mm long, margin entire

or fimbriate; glands 4, stipitate, cupuliform,

with distinct central pit and thickened rim;

transverse-oblong or orbicular in outline,

0.15-0.2 mm long, 0.2-0.4 mm wide, yellow;

gland appendages inconspicuous, spreading

radially, transverse-linear, to 0.1 mm long,

0.4-0.5 mm wide, white, glabrous, margin

entire; bracteoles 0.5-0.6 mm long, adnate for

c. 1/5 of their length to involucre, free portion

divided into few to numerous, subulate

hirsute segments. Staminate flowers 5-8 per

cyathium; pedicels 0.7-1.1 mm long; staminal

filaments c. 0.1 mm long. Pistillate flowers:

styles 0.2-0.3 mm long, ascending, smooth,

glabrous, entire or scarcely bifid, the apices

clavate. Capsules exserted from involucre

on pedicel to 2.5 mm long, broad-elliptic in

485

Halford & Harris, Euphorbia section Anisophyllum in Australia

lateral view, 1.6-1.8 mm long, 1.7-1.9 mm

across, shallowly 3-lobate with keels obtuse,

smooth, glabrous; hypogynous disc entire.

Seeds oblong-ovate in outline, 1.1-1.6 mm

long, 0.8-0.9 mm tangentially, 0.7-0.9 mm

radially, tetraquetrous in cross section; dorsal

faces planar; ventral faces concave; all faces

with 3-6 distinct transverse ridges; exotesta

thin, of ± even thickness over surface, grey-

white, microreticulate or minutely granulate,

becoming mucilaginous when moistened;

endotesta red-brown, n- 11. Figs 4, 25N.

Additional selected specimens examined : Western

Australia. Karijini N.P., 9.2 km E of Mt Bruce, 8.5 km

5 of Mt Oxer, 8.7 km WNW of Mt Howieson, Mt Bruce

Flats, May 1992, van Leeuwen 1185 (PERTH); 9 km N of

West Angela Hill, Hamersley Ranges, Jul 1997, Trudgen

16679 (PERTH); 33 miles [c. 53 km] NE of Cosmo

Newbery on road to Warburton, Aug 1962, George 3755

(PERTH); Kurrajong Rockhole, N end of Hunt Range,

May 1978, Keighery 1747 (PERTH); near Cunderdin,

Feb 1963, Aplin 2205 (MEL, PERTH); Lapsley’s

property, near Dulyalbin Rock, S of Moorine Rock, Jan

1984, Dodd 91 (PERTH). Northern Territory. Simpson

Gap, Feb 2009, Albrecht 12733 (BRI); Heavitree Range,

base of road to Telecom West Gap Tower road, Nov

1986, Thomson s.n. (DNA A80432); SE [of] White

Range, Arltunga Historical Reserve, May 1980, Kaiotas

482 (DNA); Kings Canyon, ‘Garden of Eden’, Jul 1981,

Thomson 45 (DNA). Queensland. Mitchell District:

c. 9.5 km E of Warang homestead, site on track to

the Sandstone Wall. White Mountains N.P, 57 km by

road N from Torrens Creek, Apr 2000, Thomas 1483

6 Thompson (BRI); Leander Station, Longreach, Apr

1977, Greenfield JT893 (BRI); Milo Station, NNW

of Adavale, Aug 2009, Forster PIF35305 & Thomas

(BRI). Leichhardt District: near junction of Old Rig

road and Mapala - Glenhaughton Road, c. 75 km NW

of Taroom, Mar 2004, Halford Q8214 & Edginton

(BRI); 60 km WNW of Taroom, Glenhaughton Road,

Mar 2004, Halford Q8191 & Edginton (BRI). Warrego

District: 5 km SE of Charleville, Mulga Master site,

Charleville experimental Reserve, Apr 1972, Everist

9842 (BRI). Maranoa District: 11 km SE of Gradule,

W of Goondiwindi, Sep 2001, Bean 17825 (BRI). New

South Wales. Mulwarrina Creek, Mulgowen Station,

35 miles [c. 56 km] S of Bourke, Oct 1963, Constable

4568B (NSW); N side of Mt Gunderbooka, Nov 1987,

Wilson 163 & Wilson (NSW); 5.6 km W of Girilambone,

Mitchell Highway between Nyngan and Coolabah, Aug

1987, Wiecek 35 et al. (NSW); 4 miles [c. 6 km] NW of

Girilambone, on Mitchell Highway, Sep 1968, Thompson

s.n. (NSW 520554); Forbes District, Apr 1899, Cox s.n.

(NSW 541459).

Distribution and habitat : Euphorbia

drummondii occurs widely and disjunctly

across mainland Australia from the Pilbara

region and south-western WA, through

southern NT to central and southern Qld, and

northern NSW (Map 15). It grows in open

eucalypt /Acacia dominated communities on

well drained sandy to clay-loam soils, usually

on rocky hills, plains and alluvial flats.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from March to August.

Typification: The isotype {Drummond 670 ) at

Kew consist of two sheets (186497, 186498).

The specimen on sheet 186497 matches

the holotype and isotypes at G and P, while

specimen on sheet 186498 is referable to E.

dallachyana as applied here.

Notes : Euphorbia drummondii is

characterised by the following features: the

whole plant glabrous, stems longitudinally

ridged, glands cupuliform with distinct

central pit and thickened rim, gland

appendages although small (to 0.1 mm long)

always present, transverse-linear with entire

margins; seed surfaces with 3-6 distinct

transverse ridges, capsules broad-elliptic in

lateral view, and styles short (0.2-0.3 mm

long), entire or scarcely bifid with somewhat

thick clavate apices.

The name Euphorbia drummondii has

been widely applied in the past. This wide

application of the name starts with Bentham

(1873) whose concept of E. drummondii

included three previously recognised

species (E. ferdinandi , E. dallachyana , E.

sharkoensis ). The need to recognise a wider

range of taxa has become obvious from the

specimens collected since Benthanfs work.

Fitzgerald (1918), Ewart & Kerr (1926), Domin

(1927) and Hassall (1977) started to unravel

this unwieldy complex when they reinstated

previously named species or described new

ones. Hassall (1977) proposed a number of

subspecies and varieties for the species, but

these were never validly published. We have

reinstated E. ferdinandi and E. sharkoensis

as distinct species and recognised eleven

new species that have not previously been

recognised as distinct from E. drummondii.

These are: E. albrechtii, E. flindersica , E.

gregoriensis, E. hassallii , E. multifaria , E.

papillata , E. papillifolia , E. philochalix , E.

porcata, E. verrucitesta and E. victoriensis.

486

Austrobaileya 8(4): 441-600 (2012)

Fig. 4. Euphorbia drummondii. A. habit x0.8. B. branchlet with cyathia x4. C. leaf x 4. D. stipules *12, E. cyathia with

female flower x24. F. capsule with cyathia xl6. G. cyathial gland with appendage, adaxial view x32. H. capsule, top

view xl6.1. capsule, lateral view xl6. A & C from Halford Q8191 & Edginton (BRI); B, D-I from Forster PIF35305

& Thomas (BRI). Del.W. Smith.

16. Euphorbia ferdinandi Baill., Adansonia

6: 284-285 (1866). Type: [New South Wales.]

Mt Goningberri [Koonenberry Mountain], 31

December 1860, NE Exped. [H.Beckler s.n .]

(holo: P 313103; iso: MEL 2179535).

Monoecious, herbaceous perennial, to 10

cm high, few to many annual stems arising

from woody taproot, the whole plant

glabrous. Stems prostrate or decumbent to

ascending, sparingly to much branched,

487

Halford & Harris, Euphorbia section Anisophyllum in Australia

smooth or sometimes longitudinally ridged.

Interpetiolar stipules subulate, narrow-

triangular or triangular, 0.3-1.2 mm long,

bifid or deeply bipartite, glabrous; margin

shallowly to deeply laciniate. Leaves: petiole

0.2-0.8 mm long, smooth; blade obovate,

oblong-obovate, oblong or elliptic, 3-8 mm

long, 1.4-4.2 mm wide, 1.4-2.5 times longer

than wide; adaxial and abaxial surfaces green

or blue-green (sometimes with reddish tinge

especially on margins), smooth or papillose;

base asymmetric with one side cordate

to rounded, the other cuneate to obtuse;

margin sparingly minutely toothed distally

or entire; apex rounded. Cyathia solitary at

the nodes, sometimes clustered on short leafy

lateral branchlets; peduncles 0.2-0.5 mm

long. Involucres turbinate to campanulate,

0.5-0.9 mm long, 0.6-1 mm across; lobes 5,

triangular, 0.15-0.5 mm long, margin entire

or fimbriate; glands 4, stipitate, cupuliform,

with distinct central pit and thickened rim,

transverse-elliptic or orbicular in outline, 0.1-

0.3 mm long, 0.1-0.4 mm wide, pink, orange

to red or yellowish green; gland appendages

mostly absent, rarely present, spreading

radially, transverse-linear, to 0.1 mm long,

0.3-0.4 mm wide, red or white, glabrous,

margin entire; bracteoles 0.5-0.8 mm long,

adnate for 1/3 of their length to involucre,

free portion either an entire subulate glabrous

lobe or divided into few to numerous subulate

glabrous or hirsute segments. Staminate

flowers 3-5 per cyathium; pedicels 0.7-1

mm long; staminal filaments c. 0.1 mm long.

Pistillate flowers: styles 0.2-0.4 mm long,

spreading or ascending to erect, smooth,

glabrous, entire (rarely scarcely bifid,

Halford Q9212 [BRI]), the apices terete or

clavate. Capsules exserted from involucre on

pedicel to 17(2.5) mm long, elliptic in lateral

view, 1.4-2.1 mm long, 1.2-1.7 mm across,

shallowly 3-lobate with keels obtuse to acute,

smooth or papillose, glabrous; hypogynous

disc entire. Seeds narrow-ovate in outline,

1.1-1.6 mm long, 0.5-0.7 mm tangentially,

0.4-0.6 mm radially, tetraquetrous in cross

section; dorsal faces planar or concave;

ventral faces concave; all faces smooth;

exotesta thin, of even thickness, pale brown or

white, microreticulate or minutely granulate,

becoming mucilaginous when moistened;

endotesta brown or red-brown.

Distribution : Euphorbia ferdinandi is

widespread in arid Australia from near

Wiluna and Laverton, WA, eastward through

the NT and SA to western Qld and north¬

western NSW, with disjunct populations near

Tom Price, Pilbara, WA.

Notes : Bentham (1873) placed Euphorbia

ferdinandi in synonymy under E. drummondii.

Subsequently the name E. drummondii has

been widely applied to this species. Euphorbia

ferdinandi differs from E. drummondii , in

having smooth seeds (versus seed surfaces

with 3-6 distinct transverse ridges for E.

drummondii ) and capsules distinctly longer

than wide (capsules elliptic in lateral view,

1.4-2.1 x 1.2-1.7 mm versus broad-elliptic

in lateral view, 1.6-1.8 x 1.7-1.9 mm for E.

drummondii ).

As recognised here, E. ferdinandi is

variable in degree of gland appendage

development and size of capsules and seeds.

This has led us to recognise three varieties

which can be distinguished by the following

key.

Key to varieties of Euphorbia ferdinandi

1 Gland appendages present. 16b. E. ferdinandi var. appendiculata

1. Gland appendages absent. 2

2 Capsules 1.4-1.9 x 1.3-1.7 mm, <1.3 times long as wide, distinctly

minutely papillose (visible at 40x mag.); seeds 1.1-1.3 mm

long. 16a E. ferdinandi var. ferdinandi

2. Capsules 1.8-2.1 x 1.2-1.5 mm, >1.3 times as long as wide,

faintly minutely papillose (visible at 40x mag.); seeds 1.4-1.6 mm

long. 6c. E. ferdinandi var. saxosiplaniticola

488

16a. Euphorbia ferdinandi var. ferdinandi

Stems prostrate or decumbent, sometimes

longitudinally ridged. Leaves: petiole 0.2-0.8

mm long; blade obovate or oblong-obovate,

3.4-8 mm long, 1.4-4.2 mm wide, 1.9-2.5

times longer than wide; adaxial and abaxial

surfaces papillose; margin sparingly minutely

toothed distally or entire. Involucres

turbinate to campanulate, 0.7-0.8 mm long,

0.8-1 mm across; gland appendages absent.

Pistillate flowers: styles 0.2-0.4 mm long,

entire or rarely scarcely bifid, with terete

apices. Capsules 1.4-1.9 mm long, 1.3-1.7

mm across, papillose (sometimes only

minutely so and may appear smooth without

40x mag.). Seeds 1.1-1.3 mm long, 0.5-0.7

mm tangentially, 0.5-0.6 mm radially, n = 11.

Figs 5, 250.

Additional selected specimens examined : Western

Australia. Warburton, Oct 1963, de Graff s.n. (PERTH

2437104); Mt Eveline, E of Warburton, Aug 1962,

George 3869 (PERTH); 28 miles [c. 45 km] N of

Warburton, Jul 1963, George 5315 (PERTH); 35 km

W of NT/WA border on Lasseter Highway, Apr 1992,

Zich 46 (NSW). Northern Territory. Barkly Highway,

May 1993, Egan 2232 (DNA); Stirling/Ti Tree boundary.

May 1979, Kalotas 354 (DNA); Hermannsburg Road, 52

km E of Hermannsburg, Aug 1988, Barritt 454 (DNA);

eastern Golfcourse suburb, 4 km E Alice Springs,

Mar 1988, Barritt 22 (DNA, MEL); Hamilton Downs

Road, 14 miles [c. 22 km] NW [of] Alice Springs, Jan

1964, Nelson 843 (DNA, NSW); Finke Road, 6 km E

of Wellmullinna Creek, Aug 1988, Barritt 602 (DNA,

MEL). Queensland. Burke District: 12.5 km SE of

Malbon, May 1985, Neldner 1687 & Stanley (BRI).

Gregory North District: Moonah Creek, 7 km E of Oban

homestead. May 1985, Neldner 2250 & Stanley (BRI).

Warrego District: Currawinya N.P, Thargomindah

turnoff on Eulo to Hunger ford Road, Mar 1997, Forster

PIF20367 & Watson (BRI). Gregory South District:

117 miles [c. 188 km] NW of Quilpie on Windorah Road,

May 1975, Hassall 7514 (BRI). South Australia, c. 6.5

km by road NNE of turnoff from Pipalyatjara road, on

road to Waltjitjata, Tomkinson Ranges, Sep 1978, Stove

487 (AD); Patchawara Bore, Innamincka Station, Nov

1987, Conrick 2220 (AD, MEL); 13.5 km by road SSE of

Duff Creek crossing and 52 km NNW of William Creek

Hotel on Oodnadatta track. Mar 1984, Donner 9924

(AD); 3 km N of Carl Dour Tank, Mobella Station, Aug

1995, Badman 7920 (AD); 23 km SE of Lake Bring, Oct

1987, Symon NPWS1443 (AD). New South Wales. Depot

Glen, 12 km N of Milparinka, Oct 1976, Pickard 3142

(NSW); Stud Creek, Gorge Loop Road, Sturt N.P, Aug

1991, Duncan s.n. (NSW 249230).

Distribution and habitat : Euphorbia

ferdinandi var. ferdinandi is widespread

Austrobaileya 8(4): 441-600 (2012)

in central Australia from near Wiluna and

Laverton, WA, eastward through NT and SA

to western Qld and north-western NSW, with

disjunct populations near Tom Price, Pilbara,

WA (Map 16a). It grows on sandy to sandy

loam soils in Acacia shrubland/woodland or

hummock grassland communities, mostly

on plains or along watercourses, rarely on

sandstone escarpments.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from May to October.

16b. Euphorbia ferdinandi var.

appendiculata Halford & W.K.Harris,

varietas nova ab aliis varietatibus E.

ferdinandi Baill. appendicibus glandulis

praesentibus et etiam habitu foliisque

minoribus differt. Typus: Queensland.

Gregory North District: 42 km S along

Old Cork Road from Winton - Boulia Road,

24 September 2005, D.Halford Q8604 &

M.B.Thomas (holo: BRI; iso: AD, MEL).

Stems prostrate or rarely decumbent to

ascending, longitudinally ridged. Leaves:

petiole 0.3-0.7 mm long; blade oblong or

elliptic, 3-6 mm long, 1.6-3.6 mm wide,

1.4-2 times longer than wide; adaxial surface

smooth (rarely faintly papillose); abaxial

surface smooth; margin entire. Involucres

campanulate, 0.5-0.8 mm long, 0.6-0.8 mm

across; gland appendages inconspicuous,

transverse-linear, to 0.1 mm long, 0.3-0.4 mm

wide. Pistillate flowers: styles 0.2-0.3 mm

long, entire, with clavate apices. Capsules

1.7-2 mm long, 1.3-1.6 mm across, smooth

or rarely faintly papillose. Seeds 1.3-1.4 mm

long, 0.5-0.6 mm tangentially, 0.5-0.6 mm

radially, n— 11. Fig. 25P.

Additional selected specimens examined : Northern

Territory. Hamilton Downs Station, Jan 1987, Thomson

1736 (DNA); Burt Plain, 60 km N of Alice Springs, Sep

1986, Thomson 1468 (DNA). Queensland. Mitchell

District: Idalia N.P., SW of Blackall, Jun 1990, Rogers

s.n. (BRI [AQ624140]). Gregory North District:

Diamantina N.P., track to Scotts Tank, Apr 1997, Forster

PIF20678 & Holland (BRI); 153 km S of Boulia, May

1976, Hassall 7649 (BRI). Warrego District: c. 40

miles [ c. 64 km] S of Charleville on Cunnamulla Road,

Aug 1973, Hassall 7384 (BRI); Currawinya N.P., Paroo

River floodplain south of Caiwarra ruins. Mar 1997,

Forster PIF20507 & Watson (BRI); Currawinya N.P.,

north of Hungerford, Apr 1994, Swain 604 (BRI); 33.5

Halford & Harris, Euphorbia section Anisophyllum in Australia

489

Fig. 5. Euphorbia ferdinandi \ar.ferdinandi. A. habit *0.6. B. branchlet with cyathia *6. C. leaf *8. D. papillae on

lower leaf surface x32. E. stipules xl6. F. cyathia with female flower x32. G. capsule with cyathia xl6. H. cyathial

gland, adaxial view x32. I. capsule, top view xl6. J. capsule, lateral view xl6. A from Barritt 454 (DNA); B-J from

Kalotas 354 (DNA). Del. W. Smith.

490

km E of Eulo, towards Cunnamulla, Aug 2006, Bean

25669 (BRI). Gregory South District: c. 9 km due W of

Tickalara homestead, c. 170 km SW of Thargomindah,

1995, Kemp 895 & Fairfax (BRI). South Australia. 25.7

km from Birdsville road, E of Macumba homestead.

Browns Creek, Sep 1986, Ballingall 2238 (BRI); c. 65

km N of Innamincka, Aug 1975, Weber 4680 (AD);

Marree, Jun 1930, George s.n. (AD 98597032); Myall

Well, Andamooka Station, Oct 1989, Badman 3862

(AD). New South Wales, walking track c. 2.25 km from

Mt Wood summit, Sturt N.R, Sep 1989, Wiecek 313 et

al. (NSW); Rossmore Station, 20 miles [c. 32 km] NE of

Bourke, Oct 1963, Constable 4481 (NSW).

Distribution and habitat : Euphorbia

ferdinandi var. appendiculata occurs from

near Alice Springs, NT, north east to Julia

Creek, Qld, south to north-eastern SA and

east to Bourke, western NSW (Map 16b). It

commonly grows in grassland communities

on cracking clay soils, or chenopod shrubland

communities on stony clay soils on plains or

alluvial flats.

Phenology : Flowers and fruits have been

collected throughout the year.

Notes : Euphorbia ferdinandi var.

appendiculata differs from the other varieties

by having gland appendages and it is also

generally smaller in habit and leaves.

Etymology : The varietal epithet is Latin

appendiculatus, with small appendages, in

reference to the presences of gland appendages

in this variety.

16c. Euphorbia ferdinandi var.

saxosiplaniticola Halford & W.K.Harris,

varietas nova ab E. ferdinandi Baill.

var. ferdinandi capsulis angustioribus et

plerumque longioribus 1.8-2.1 x 1.2-1.5 mm

>1.3 plo longioribus quam latioribus tantum

minute papillosis (in vicem 1.4-1.9 x 1.3-1.7

mm <1.3 plo longioribus quam latioribus

distincte minute papillosis), seminibus

longioribus 1.4-1.6 mm longis (in vicem

1.1-1.3 mm longis) necnon E. ferdinandi

var. appendiculata Halford & W.K.Harris

appendices glandulae carens differt. Typus:

South Australia. Poonia Waterhole, 5

September 1986, J.S.Weber 9140 (holo: AD;

iso: three duplicates to be distributed).

Stems prostrate or decumbent. Leaves:

petiole 0.3-0.5 mm long; blade oblong,

obovate or elliptic, 4.2-7 mm long, 2.1-3

Austrobaileya 8(4): 441-600 (2012)

mm wide, 2.3-2.5 times longer than wide;

adaxial and abaxial surfaces smooth; margin

entire or sparingly minutely toothed distally.

Involucres turbinate, 0.8-0.9 mm long, 0.6-

0.7 mm across; gland appendages absent.

Pistillate flowers: styles 0.2-0.3 mm long,

entire, with terete apices. Capsules 1.8-2.1

mm long, 1.2-1.5 mm across, smooth or

faintly papillose. Seeds 1.4-1.6 mm long,

0.5-0.7 mm tangentially, 0.4-0.5 mm radially.

Fig. 25Q.

Additional selected specimens examined : Northern

Territory. Allitra Tableland, Simpson Desert, Aug

1992, Latz 12450 (DNA); Mac Clarke Reserve, Andado

Station, Aug 1992, Coulson 61 & Latz (DNA); Mt

Wilyunpa, Aug 1992, Coidson 35 (DNA); Alice Springs,

Andado Station, Aug 1997, Michell 391 & Calliss

(DNA). South Australia. Pedirka, Aug 1932, Ising (AD

966031149); Minnie Downs, Apr 1924, Reese 130 (AD);

8 km SSW of homestead, Dulkaninna Station, Dec 1995,

Badman 8796 (AD).

Distribution and habitat : Euphorbia

ferdinandi var. saxosiplaniticola occurs in

central Australia from Allitra Tableland, NT

southward to near Maree, SA (Map 16c).

It grows on clay soils on gibber plains or

tablelands.

Phenology : Flowers and fruits have been

collected in April, August, September and

December.

Notes : Euphorbia ferdinandi var.

saxosiplaniticola differs from E. ferdinandi

var. ferdinandi in having narrower and

generally longer capsules, 1.8-2.1 mm long

by 1.2-1.5 mm wide, >1.3 times as long as

wide, that are only faintly minutely papillose

(versus capsules 1.4-1.9 mm long by 1.3—1.7

mm wide, <1.3 times long as wide, that are

distinctly minutely papillose for E. ferdinandi

var. ferdinandi) and longer seeds 1.4-1.6

mm long (versus seeds 1.1-1.3 mm long for

E. ferdinandi var. ferdinandi). Euphorbia

ferdinandi var. saxosiplaniticola differs from

Euphorbia ferdinandi var. appendiculata in

lacking glandular appendages.

Etymology : The varietal epithet is from

Latin saxosa , rocky or stony places, planities,

flat surface, plain, and -cola, dweller or

inhabitant, in reference to the gibber plains of

Central Australia where this variety has been

recorded growing.

491

Halford & Harris, Euphorbia section Anisophyllum in Australia

17. Euphorbia fitzroyensis Halford &

W.K.Harris, species nova similis E.

drummondii Boiss. et E. hassallii Halford

& W.K.Harris a duabus autem superficiebus

seminum porcis 6-9 angustis rotundatis

transversis (ad vicem porcis 3-6 transversis

in E. drummondii et porcis 3-5 prominentibus

rotundatis transversus irregularisque in E.

hassallii ) necnon ab E. drummondii stylis

1/3—1/2 longitudinis bifidis (ad vicem stylis

integris vel vix bifidis) et exotesta manifeste

crassa in porcis transversis (ad vicem exotesta

aeque crassa in superficie seminum) differt.

Typus: Western Australia. Liveringa Station,

near Fitzroy River, 20 April 1985, T.E.H.Aplin

172 et al. (holo: PERTH).

Monoecious, annual to 30 cm high, few

to many stems arising from rootstock, the

whole plant glabrous. Stems erect, sparingly

to much branched, longitudinally ridged.

Interpetiolar stipules subulate or narrow-

triangular, 0.4-0.8 mm long, deeply bipartite,

glabrous; margin entire or laciniate. Leaves:

petiole 0.3-0.5 mm long, smooth; blade

narrow-oblong, 5-9 mm long, 1.5-3.1 mm

wide, 2.5-4.2 times longer than wide; adaxial

and abaxial surfaces colour unknown, smooth

or minutely papillose (visible at 40 x mag.);

base asymmetric with one side cordate, the

other obtuse; margin entire or sparingly

minutely toothed distally; apex rounded or

obtuse. Cyathia solitary at the nodes, often

clustered on short leafy lateral branchlets

with subtending leaves slightly smaller than

the primary stem leaves; peduncles 0.2-1.4

mm long. Involucres turbinate, 0.5-0.8 mm

long, 0.4-0.6 mm across; lobes 5, triangular,

0.3-0.4 mm long, margin entire; glands 4,

stipitate, cupuliform, with distinct central pit,

transverse-oblong in outline, 0.1-0.15 mm

long, 0.2-0.4 mm wide, colour unknown;

gland appendages inconspicuous, spreading

radially, transverse-linear; c. 0.05 mm long

and 0.35 mm wide, glabrous, colour unknown,

margin entire; bracteoles 0.6-0.8 mm long,

adnate for 1/4 of their length to involucre, free

portion entire subulate glabrous. Staminate

flowers 2-4 per cyathium; pedicels 0.8-1

mm long; staminal filaments c. 0.1 mm long.

Pistillate flowers: styles 0.2-0.3 mm long,

spreading or ascending, smooth, glabrous,

each bifid for 1/3—1/2 of their length, the apices

clavate or terete. Capsules exserted from

involucre on pedicels to 2 mm long; broad-

elliptic in lateral view, 1.6-1.8 mm long, 1.6-

1.7 mm across, shallowly 3-lobate with keels

obtuse, smooth, glabrous; hypogynous disc

entire. Seeds narrow-ovate in outline, 1.2-1.4

mm long, 0.7-0.8 mm tangentially, 0.6-0.7

mm radially, tetraquetrous in cross section;

dorsal faces planar; ventral faces planar or

concave; all faces with 6-9 prominent narrow

rounded transverse ridges; exotesta of uneven

thickness, distinctly thicker on transverse

ridges, white or pale brown, microreticulate;

endotesta brown or red-brown. Figs 6, 25R.

Additional specimens examined : Western Australia.

Fitzroy River crossing, S of Derby, May 1991, Thomson

3475 (NT); Fitzroy River, Aug 1906, Fitzgeralds.n. 1567

(PERTH); Fitzroy River, Aug 1906, Fitzgerald (NSW

521748); Liveringa, Jan 1953, Broadbent 590 (BM);

Geikie Gorge N.P., Jul 1974, Carr 3803 & Beauglehole

47581 (NSW, PERTH); Gogo Station, Fitzroy Crossing,

May 1962, Royce 7005 (PERTH).

Distribution and habitat : Euphorbia

fitzroyensis is confined to the vicinity of

the Fitzroy River, WA (Map 17). It grows

on alluvial plains in eucalypt woodland

communities on heavy loam to silty clay

soils, and in Triodia grassland communities

on sandy soils.

Phenology : Flowers and fruits have been

collected from April to August.

Notes : Euphorbia fitzroyensis is similar to E.

drummondii and E. hassallii. It differs from

both species by having seed surfaces with 6-9

narrow rounded transverse ridges (versus 3-6

transverse ridges for E. drummondii , and 3-5

prominent rounded transverse or irregular

ridges for E. hassallii). It also differs from E.

drummondii in having styles bifidly divided

for 1/3—1/2 of their length (versus styles entire

or scarcely bifid for E. drummondii ) and

exotesta is distinctly thicker on transverse

ridges (versus exotesta is of even thickness

over seed surface for E. drummondii).

The collection (Royce 7005) is tentatively

placed here. Although its general form,

leaf shape, involucral glands and seeds are

consistent with E. fitzroyensis , it differs in

having stems lacking longitudinal ridges.

492

Austrobaileya 8(4): 441-600 (2012)

Fig. 6. Euphorbiafitzroyensis. A. habit *0.4. B. branchlet with cyathia x8. C. leaf x6. D. stipules xl6. E. cyathia with

female flower x32. F. capsule with cyathia xl6. G. cyathial gland with appendage, adaxial view x32. H. capsule, top

view xl6.1. capsule, lateral view xl6. A-I from Aplin 172 (PERTH). Del. W. Smith.

493

Halford & Harris, Euphorbia section Anisophyllum in Australia

Etymology : The specific epithet refers to

the Fitzroy River area in the Kimberley, WA

where the species occurs.

18. Euphorbia flindersica Halford &

W.K.Harris, J. Adelaide Botanic Gardens 24:

43-45 (2010). Type: South Australia. MtGee,

15 September 1973, R.H.Kuchel 3169 (holo:

AD).

Illustration : Halford & Harris (2010: 44, fig.

1 ).

Monoecious, herbaceous perennial to 10 cm

high, with short-lived stems produced from

thick somewhat woody rootstock, the whole

plant glabrous. Stems prostrate to erect,

much branched, smooth or faintly papillose.

Interpetiolar stipules triangular 0.4-0.6 mm

long, deeply bipartite, glabrous; margin entire

or laciniate. Leaves: petiole 0.2-1.2 mm long,

smooth; blade oblong or obovate, 1.3-6.7 mm

long, 1-3.8 mm wide, 1.5-1.8 times longer

than wide, both surfaces minutely papillose

(visible at 40x mag.); adaxial surface mostly

green sometimes with reddish tinge on

margin; abaxial surface grey; base strongly

asymmetric with one side cordate to rounded,

the other cuneate to rounded; margins entire

or sparsely minutely toothed distally; apex

rounded. Cyathia solitary at the nodes;

peduncles 0.3-0.7 mm long. Involucres

campanulate or cupuliform, 0.8-1 mm long,

0.6-1.2 mm across; lobes 5, triangular, 0.3-

0.4 mm long, margins ciliate; glands 4, shortly

stipitate, patelliform, planar or shallowly

concave, transverse-oblong to transverse-

elliptic, 0.1-0.3 mm long, 0.3-0.5 mm wide,

red or yellowish green; gland appendages

conspicuous, spreading radially, obdeltoid,

0.3-0.4 mm long, 0.6-0.8 mm wide, pink or

red, glabrous, dentate or irregularly lobed;

bracteoles 0.5-0.9 mm long, adnate for 1/3 of

their length to involucre, free portion divided

into few to numerous subulate glabrous

segments. Staminate flowers 10-15 per

cyathium; pedicels 0.7-1 mm long; staminal

filaments c. 0.1 mm long. Pistillate flowers:

styles c. 0.5 mm long, spreading, smooth,

glabrous, each bifid to 1/4—1/3 of their length,

apices slender terete. Capsules exserted from

involucre on pedicel to 2.7 mm long, ovate

to broad-ovate in lateral view, 1.5-1.8 mm

long, 1.7-2.2 mm across, shallowly 3-lobate

with keels acute, papillose; hypogynous disc

entire. Seeds ovate in outline, 1.1-1.3 mm

long, 0.7-0.8 mm tangentially, 0.6-0.8 mm

radially, tetraquetrous in cross section; dorsal

faces planar; ventral faces planar or shallowly

convex; all faces smooth or with faint irregular

ridges; exotesta thin, of even thickness, cream

or pale brown, microreticulate, becoming

mucilaginous when moistened; endotesta

reddish brown, n = 11. Fig. 26A.

Additional selected specimens examined: South

Australia. Gorge Creek of Myrtle Springs, c. 24 km

NW of Leigh Creek, Sep 1962, Lothian 1077 (AD);

Paralana Springs, 125 km NE of Blinman, Aug 1968,

Carrick 2059 (AD); south branch of Paralana Hot

Springs Creek, Aug 1964, Saddler s.n. (AD 96532103);

Nepouie Springs, Apr 1994, Bates 37341 (AD); Grindell

Hut, Balcanoona Station, July 1980, Williams 11205

(AD); Italowie Creek, Aug 1979, Conrick AD100 (AD);

Chambers Gorge, near Mt Chambers, c. 60 km ENE

of Blinman, Sep 1956, Eichler 12559 (AD); Chambers

Gorge, c. 80 km ENE of Parachilna, Sep 1973, Whibley

3922 (AD); Parachilna Gorge, Aug 1963, Sharrad 1404

(AD); upper Bunyeroo Gorge, c. 50 km NNE of Hawker,

Oct 1958, Kraehenbuehl 14 (AD); The Bunkers, Bunkers

Range, Apr 1989, James 16 (AD); Arkaroola Sanctuary,

Ridge Top road, Oct 1971, Kuchel 3039 (AD); western

portion [of] Oraparinna N.P, Sep 1971, Weber 2710

(AD); Brachina Gorge, Sep 1961, Symon 1400 (AD);

Moralana Station, road & rail crossing Bunyeroo Creek,

Jul 1987, Symon 14628 (AD); Arkaba Station, Oct 1925,

Beck s.n. (AD 96938301).

Distribution and habitat : Euphorbia

flindersica is restricted to the northern

Flinders Ranges, SA, occurring from near

Leigh Creek to Hawker (Map 18). It grows on

sandy clay soils on rocky outcrops or gravelly

hill slopes.

Phenology : Flowers and fruits have been

collected from April to October.

Notes : Euphorbia flindersica is similar to E.

ferdinandi and E. papillata in having papillose

capsules and leaves but differs from both in

having involucral glands patelliform, planar

or shallowly concave (versus cupuliform, with

distinct central pit and thickened rim for E.

ferdinandi and E. papillata).

Euphorbia multifaria and E. verrucitesta

overlap the distributional range of E.

flindersica and all three species were

previously included in E. drummondii s.l.

in SA (Weber 1986). Euphorbia flindersica

494

differs from these species by its planar or

shallowly concave involucral glands, larger

and conspicuous gland appendages, 0.3-0.4

mm long, which are dentate or irregularly

lobed.

Hassall (1977) proposed the name E.

drummondii subsp. blackii for this taxon but

this name was never validly published.

19. Euphorbia gregoriensis Halford

& W.K.Harris, species nova similis E.

schultzii var. comanti (W.Fitzg.) Halford

& W.K.Harris in habitu et indumento et

forma foliorum et in amplitudine differt

autem capsulis angustioribus (1.9-2 mm)

longitudine latitudinem aequi leviter 3-lobatis

(ad vicem 2-3 mm latis manifeste latioribus

quam latioribus profunde 3-lobatis). Similis

E. hassallii Halford & W.K.Harris in

forma capsularum differt autem exotesta

inaequaliter crassa semper crassiore in porcis

(ad vicem appendicibus <0.1 mm longis aut

si longioribus nunc lobatis profunde) paginis

foliorum falcato-oblong is 9.8-12 x 4-6

mm <3 plo longioribus quam latiorbus (ad

vicem paginis foliorum anguste oblongis vel

anguste ovatis 8.2-25 x 2-4.8 mm >3 plo

longioribus quam latis). Typus: Northern

Territory. Gregory National Park, 14 km NE

of Bullita O/S [outstation], 8 February 1986,

B.G.Thomson 1114 (holo: DNA).

Monoecious, annual to 40 cm high, one

to few stems arising from base, the whole

plant glabrous except for a few scattered

hairs on stems proximally. Stems erect,

sparingly branched, smooth, glabrous or

sparsely hairy proximally; hairs spreading,

curled or straight, 0.5-1.0 mm long, white.

Interpetiolar stipules subulate, 0.5-1.3

mm long, bipartite, glabrous; margin entire.

Leaves: petiole 0.5-0.8 mm long, smooth;

blade falcate-oblong, 9.8-12 mm long, 4-6

mm wide, 1.8-2.7 times longer than wide;

adaxial and abaxial surfaces green, minutely

papillose to glabrous; base asymmetric,

cordate; margin serrate at the apex and along

one side or sparingly toothed distally; apex

rounded or obtuse. Cyathia solitary at the

nodes, sometimes clustered on short leafy

lateral branchlets with subtending leaves

slightly smaller than the primary stem leaves;

Austrobaileya 8(4): 441-600 (2012)

peduncles 0.5-0.7 mm long. Involucres

campanulate or turbinate, 0.8-1 mm long,

0.7-0.8 mm across; lobes 5, triangular,

0.5-0.6 mm long, margin fimbriate; glands

4, stipitate ( c. 0.075 mm long), cupuliform,

with distinct central pit, transverse-oblong

in outline, c. 0.2 mm long, 0.3-0.4 mm

wide, pale green or pink; gland appendages

conspicuous, spreading radially, very broad-

obovate, 0.1-0.4 mm long, 0.6-0.8 mm

wide, pink or white, glabrous, margin entire;

bracteoles c. 0.8 mm long, adnate for 1/3—

1/2 of their length to involucre, free portion

entire, subulate or divided into numerous

subulate segments, glabrous. Staminate

flowers 7-10 per cyathium; pedicels 1-1.1

mm long; staminal filaments 0.1-0.2 mm

long. Pistillate flowers: styles 0.4-0.5 mm

long, ascending, smooth, glabrous, each bifid

for c. 1/3 of their length, the apices terete.

Capsules exserted from involucre on pedicel

to 2.5 mm long, broad-elliptic in lateral view,

1.9-2 mm long, 1.9-2 mm across, shallowly

3-lobate with keels acute, smooth or minutely

papillose, glabrous; hypogynous disc entire.

Seeds ovate in outline, 1.3-1.5 mm long; 0.8-

0.9 mm tangentially, 0.8-0.9 mm radially,

tetraquetrous in cross section; dorsal and

ventral faces planar, with prominent broad

rounded irregular ridges; exotesta of uneven

thickness, distinctly thicker on ridges, white

or pale brown, microtuberculate, becoming

mucilaginous when moistened; endotesta

brown or red-brown. Figs 7, 26B.

Additional selected specimens examined: Northern

Territory. 50 km [ c . 80 km] S [of] Bullita O/S [outstation],

Gregory N.P., Mar 1991, Thomson 3453 (DNA, NT); 20

miles [c. 32 km] S [of] Timber Creek Police Station,

May 1959, Chippendale 6050 (DNA, MEL, NSW);

Gregory N.P, 8 km NW of Bullita O/S [outstation], Feb

1986, Thomson 1082 (DNA); Gregory N.P., 3 km S of

Bullita O/S [outstation], Feb 1986, Thomson 963 (DNA);

Bullita Station, Gregory N.P, Feb 1986, Wightman 2591

& Clark (DNA); Gregory N.P, Feb 1986, Thomson 1166

(DNA); Gregory N.P., Bullock Paddock Creek valley,

Apr 1996, O’Neill 24 (DNA); Gregory N.P, Station Hill,

Feb 1992, Cowie 2502 & Brocklehurst (DNA); Wickham

River, Mar 1992, Brocklehurst 616 (DNA).

Distribution and habitat : Euphorbia

gregoriensis is restricted to the hills of the

Gregory N.P. and Victoria River Station,

NT (Map 19). It grows in eucalypt open

woodland communities on shallow soils on

Halford & Harris, Euphorbia section Anisophyllum in Australia

495

Fig. 7 . Euphorbia gregoriensis. A. habit *0.4. B. leaf x4. C. stipules *12. D. cyathia with female flower x24. E. capsule

with cyathia xl2. F. cyathial gland with appendage, adaxial view x32. G. capsule, top view xl2. H. capsule, lateral

view xl2. A & B from Thomson 1114 (DNA); C & D from Thomson 1082 (DNA); E-H from Brocklehurst 616 (DNA).

Del.W. Smith.

496

rocky sandstone mesa tops, scree slopes or

hill sides.

Phenology : Flowers and fruits have been

collected from February to May.

Notes: Euphorbia gregoriensis is

morphologically similar to E. schultzii var.

comans (W.Fitzg.) Halford & W.K.Harris in

habit, indumentum, and leaf shape and size,

but differs by having narrower capsules (1.9-

2 mm) that are as long as wide and shallowly

3-lobate (versus 2-3 mm across, distinctly

broader than long and deeply 3-lobate

for E. schultzii var. comans). Euphorbia

gregoriensis is similar to E. hassallii in

capsule shape but differs by having exotesta

of uneven thickness, always thicker on ridges

(versus exotesta ± of even thickness for E.

hassallii), gland appendages >0.1 mm long

and entire (versus gland appendages <0.1

mm long or if longer then appendages deeply

lobed for E. hassallii) and leaf blades falcate-

oblong, 9.8-12 x 4-6 mm, <3 times as long

as wide (versus leaf blades narrow-oblong or

narrow-ovate, 8.2-25 x 2-4.8 mm, >3 times

as long as wide for E. hassallii).

The collection Chippendale 6111 (3.3

miles [ c. 5.3 km] E [of] Willeroo, May 1959

[DNA, NSW]) is tentatively referred to here as

it most resembles E. gregoriensis. However,

it is atypical in having a longitudinal band

of short crispate hairs along its stems and its

seeds lack the thick endotesta that is typical

of this species.

Etymology : The specific epithet refers to the

Gregory N.P., in the NT where the species

occurs.

20. Euphorbia hassallii Halford &

W.K.Harris, species nova similis E.

drummondii Boiss. sed plerumque stylis

longioribus 0.3-0.4 mm longis bifidis usque

1/3 longitudinis apicibus teretibus (ad vicem

0.2-0.3 mm longis integris vel vix bifidis

apicibus clavatus) necnon appendicibus

glandulae interdum aut carentibus aut

leviter usque profunde lobatis (ad vicem

appendicibus glandulae semper qui sunt

<0.1 mm longis) differt. Typus: Northern

Territory. Victoria Highway, 30 km W of

Katherine, 23 December 1994, M.J.A.Barritt

Austrobaileya 8(4): 441-600 (2012)

1700 (holo: BRI, iso: DNA, MEL).

Monoecious, annual or herbaceous perennial

with slender taproot, to 30 cm high, few

stems arising from rootstock. Stems erect or

ascending (rarely prostrate), sparingly to much

branched, smooth, sometimes longitudinally

ridged, glabrous or with a sparse indumentum;

hairs spreading, ± straight, 0.2-0.4 mm

long, white. Interpetiolar stipules subulate,

0.5-1.1 mm long, deeply bipartite, glabrous;

margin entire. Leaves: petiole 0.4-0.6 mm

long, smooth, glabrous; blade narrow-oblong

or narrow-ovate, 8.2-25 mm long, 2-4.8 mm

wide, 3.7-8.8(32) times longer than wide;

adaxial surface dark green sometimes with

reddish tinge, minutely papillose, glabrous;

abaxial surface paler than adaxial surface,

minutely papillose, glabrous or with a sparse

indumentum; hairs appressed-ascending,

straight, 0.2-0.3 mm long; base asymmetric

with one side cordate, the other obtuse to

cordate; margin entire or sparingly minutely

toothed distally; apex rounded, obtuse or

acute. Cyathia solitary at the upper nodes,

often clustered on short leafy lateral branchlets

with subtending leaves slightly smaller than

the primary stem leaves, peduncles 0.3-0.5

mm long, smooth, glabrous. Involucres

campanulate or turbinate, 0.6-0.9 mm long,

0.8-1 mm across; lobes 5, triangular, 0.3-0.5

mm long, margin entire or fimbriate; glands

4, stipitate, cupuliform, with distinct central

pit and thickened rim, transverse-oblong

in outline, 0.1-0.2 mm long, 0.3-0.5 mm

wide, red; gland appendages conspicuous or

absent, spreading radially, transverse-linear

or obdeltoid, to 0.4 mm long, 0.3-0.4 mm

wide, pink or red, glabrous, margin shallowly

to deeply lobed; bracteoles 0.5-0.9 mm long,

free or adnate for up to 1/3 of their length to

involucre, free portion entire and subulate or

divided into few subulate segments, glabrous.

Staminate flowers 3-10 per cyathium;

pedicels c. 0.8 mm long; staminal filaments

c. 0.1 mm long. Pistillate flowers: styles 0.3-

0.4 mm long, spreading, smooth, glabrous,

each scarcely bifid or bifid for c. 1/3 of their

length, the apices terete. Capsules exerted

from involucre on pedicel to 2 mm long,

broad-elliptic in lateral view, 1.5-1.8 mm

long, 1.6-1.9 mm across, shallowly 3-lobate

497

Halford & Harris, Euphorbia section Anisophyllum in Australia

with keels obtuse, smooth or minutely 1

papillose, glabrous; hypogynous disc entire. c

Seeds ovate in outline, 1.1-1.3 mm long, 1

07-0.8 mm tangentially, 0.7-0.8 mm radially, f

tetraquetrous or tetragonous in cross section; (

dorsal faces planar or convex; ventral faces 1

planar; all faces with 3-5 prominent rounded I

transverse or irregular ridges; exotesta thin, of a

even thickness, grey-white, microreticulate, t

becoming mucilaginous when moistened; 2

endotesta reddish brown, w = 11. Figs 8, 26C. 1

Additional selected specimens examined : Western

Australia. 1 km E [of] Durack River crossing. Home s

Valley - Gibb River Road, Mar 1991, Thomson 3461

(NT); 19 km SE of East Wyndham on Kununurra

Road, Jul 1974, Carr 3224 & Beauglehole 47002 3

(NSW, PERTH); Bobby Creek, 20.3 km N of turn off a

to Beagle Bay on Cape Leveque - Broome Road, Apr ^

1988, Kenneally 10638 (PERTH); One Arm Point, NE of

airstrip. Mar 1996, Carter BJC708 (BRI, PERTH); NE of C

airstrip One Arm Point, Apr 1993, Carter 629 (PERTH); 1

W of Dragon Tree Soak, Great Sandy Desert, Aug 1977, C

George 14803 (BRI, PERTH); 80 Mile Beach, near g

Caravan Park, May 1991, Thomson 3662 (DNA, NT). +

Northern Territory. Twin Falls, Mar 1982, Dunlop 6219

6 Taylor (BRI); Mataranka, Elsey N.P, Feb 1994, Egan "

3199 (DNA); Keep River N.P, Apr 1991, Evans 3760 I

(BRI); Station Hill, Gregory N.P, Feb 1986, Wightman J

2746 & Clark (DNA); Gregory N.P, Jasper Creek, Apr |<

1996, Booth 1561 & Woodward (DNA); Limmen Bight

River upper reaches, Jan 1989, Thomson 2859 (DNA); l

7 Mile Spring, Gallipoli Station, Jan 1989, Latz 11226 z

(DNA). Queensland. Cook District: 60.5 km by road /

E of Croydon, Gregory River area, Jan 2005, McDonald

KRM3489 (BRI). Burke District: Westmoreland ^

Station, just W of Hells Gate, 15 km SE of homestead, c

May 2005, Booth 4292 & Thompson (BRI); 28.5 km SSW 8

of Hells Gate Roadhouse on turnoff Lagoons Station, ,

Apr 2006, Thompson WES274 & Edginton (BRI); 54

km NW of Burketown on Escott Station, Apr 2007, 2

Thompson WES1392 & Wilson (BRI); 9.1 km by road 1

W of Gilbert River Road crossing towards Croydon, Jan

2005, McDonald KRM2402 (BRI); Calton Hills Station, 2

N of Mt Isa, May 2004, Booth 3488 & Kelman (BRI). (

Distribution andhabitat : Euphorbia hassallii

occurs across northern Australia from the j

Kimberley, WA, through the northern regions j

of the NT and extending to near Emerald in

central Qld (Map 20). It grows in eucalypt 1

open forest/woodland, Acacia shrubland or 1

tussock grassland communities on sandy to (

clay soils on plateaux, rocky slopes or plains. c

Phenology : Flowers and fruits have been ^

collected from December to July. 1

Notes : Euphorbia hassallii is similar to E.

drummondii but differs in having generally

longer styles (0.3-0.4 mm long) that are bifid

for up to a 1/3 of their length, with terete apices

(versus 0.2-0.3 mm long, entire or scarcely

bifid, with clavate apices for E. drummondii).

It also differs in generally having longer gland

appendages (when present) that are shallowly

to deeply lobed (versus gland appendages

always present, entire and less than 0.1 mm

long for E. drummondii) and stems often with

a sparse indumentum proximally (versus

stems always glabrous for E. drummondii).

Euphorbia hassallii is a morphologically

variable complex and with further collections

and study may be subdivided into a number of

taxa. One of the more recognisable variants

occurs in the Kimberley, WA. This variant

tends to have a prostrate habit, smaller leaves,

capsules and seeds, and deeply lobed gland

appendages. Representative specimens of

this variant are: 15 km N [of] Kalumburu

Mission, Apr 1991, Thomson 3468 (DNA, NT,

PERTH); Sir Graham Moore Island, Jul 1973,

Wilson 11252 (PERTH); Durack Range, 90

km WSW of Kununurra, Feb 1993, Keighery

140 & Gibson (PERTH); Old Pago Mission,

22 kmNNE of Kalumburu, Jun 1990, Edinger

732 (PERTH).

Etymology : The species is named in honour

of Dr David Hassall, botanist and landscape

architect, whose studies on the Australian

Euphorbieae (Hassall 1977) have been of

assistance in understanding the complexity of

Euphorbia in Australia.

21. *Euphorbia hirta L., Sp. PI. 454. 1753;

Chamaesyce hirta (L.) Millsp., Publ. Field

Columb. Mus., Bot. Ser. 2: 303 (1909). Type:

“Habitat in India” (lecto: LINN [Herb. Linn.

No. 630. 7] n.v. (image seen) (IDC microfiche

177. 320: II. 5), fide Wheeler (1939: 72)).

Euphorbia pilulifera f. humifusa Domin,

Biblioth. Bot. 89(4): 312 (1927 ‘1926’). Type:

Queensland. [North Kennedy District:] apud

opp. Pentland, February 1910, K.Domin s.n.

(holo: PR 528327).

Euphorbia pilulifera f. rubromaculata

Domin, Biblioth. Bot. 89(4): 312 (1927 ‘1926’).

Type: not designated.

498

Austrobaileya 8(4): 441-600 (2012)

Fig. 8. Euphorbia hassallii. A. habit *0.3. B. branchlet with cyathia x4. C. leaf x4. D. stipules xl6. E. cyathia with

female flower x24. F. capsule with cyathia xl2. G. cyathial gland with appendage, adaxial view x32. H. capsule, top

view xl2.1. capsule, lateral view xl2. A-I from Barritt 1700 (BRI). Del. W.Smith.

499

Halford & Harris, Euphorbia section Anisophyllum in Australia

Euphorbia pilulifera f. viridis Domin, C

Biblioth. Bot. 89(4): 312 (1927 ‘1926’). Type: s

not designated. c

Euphorbia chrysochaeta W.Fitzg., J. & Proc.

Roy. Soc. Western Australia 2: 162 (1918).

Type: Western Australia. May River, nr.

Emmanuel’s Yards, May 1905, W.V Fitzgerald

436 (lecto [here designated]: PERTH n.v. J

(photo at BRI); isolecto: (NSW 612208). j

Illustrations: James & Harden (1990: 429), i

as Chamaesyce hirta, Lin et al. (1991: 229, f

fig. 8), as Chamaesyce hirta ; Wheeler (1992: r

figs 183E, 1841, 1851, 1861); Aboriginal 1

Communities of the Northern Territory (1993: 1

285-286); Dunlop et al. (1995: 218, fig. 72); f

Kleinschmidt et al. (1996: 31); Harris (2001: (

32, fig. 1), as Chamaesyce hirta. t

Monoecious, annual to 30 cm high, with one c

to a few stems arising from the base. Stems J

ascending to erect, sparingly branched,

smooth, sparsely to densely hairy; indumentum ,

consisting of white weakly appressed crispate

hairs to 0.5 mm long interspersed with yellow

spreading ± straight segmented hairs to 1.5

mm long. Interpetiolar stipules narrow-

triangular, 1-1.8 mm long, bifid or deeply

bipartite, sparsely hairy abaxially with white

ascending hairs 0.2-0.3 mm long; margin ^

laciniate. Leaves: petiole 0.5-3 mm long,

smooth, with indumentum as for stems; blade /

ovate or elliptic, 10-50 mm long, 5-21 mm

wide, 1.6-2 times longer than wide; adaxial (

surfaces red or dark green with reddish tinge, s

smooth, with a sparse indumentum consisting (

of white ± appressed curved hairs 0.2-0.4 j

mm long; abaxial surface pale green often ^

with reddish tinge, smooth, with a sparse to i

moderately dense indumentum consisting of l

white ± appressed crispate or curved hairs 1

up to 1.1 mm long and scattered yellow ± 1

straight segmented hairs up to 1.5 mm long; (

base asymmetric with on side rounded, the s

other cuneate; margin serrulate; apex acute. (

Cyathia in dense (7-160 cyathia) terminal j.

and axillary capitate, cymose clusters to ^

15 mm in diameter on peduncles 2-17 mm i

long; bracts subulate to narrow-triangular, I

to 0.5 mm long; cyathial peduncles 0.2-1.5

mm long. Involucres turbinate, 0.5-0.9 mm l

long, 0.5-0.7 mm across; lobes 5, triangular, c

0.2-0.3 mm long, margin laciniate; glands 4,

stipitate, cupuliform, concave or with shallow

central pit, orbicular in outlline, 0.1-0.2 mm

long, 0.1-0.2 mm wide, red or purple; gland

appendages present and inconspicuous or

absent, spreading radially, transverse-oblong

or lunate, to 0.15 mm long, 0.1-0.3 mm

wide, pink to white, glabrous, margin entire;

bracteoles 0.4-0.5 mm long, adnate for c. 2/3 of

their length to involucre, free portion divided

into ± linear hairy segments. Staminate

flowers 4-8 per cyathium; pedicels 0.3-0.8

mm long; staminal filaments 0.1-0.3 mm

long. Pistillate flowers: styles 0.2-0.3 mm

long, spreading, smooth, glabrous, each bifid

for 1/2-2/3 of their length, the apices clavate.

Capsules exserted from involucre on pedicel

to 1.2 mm long, depressed ovate in lateral view,

1- 1.4 mm long, 0.9-1.4 mm across, shallowly

3-lobate with keels acute, smooth, pubescent;

hairs appressed, c. 0.1 mm long; hypogynous

disc entire. Seeds ovate in outline, 0.7—1 mm

long, 0.4-0.5 mm tangentially, 0.4-0.5 mm

radially, tetraquetrous in cross section; dorsal

and ventral faces planar to concave, with

2- 4 faint acute transverse ridges; exotesta

very thin, of even thickness, grey-white,

microreticulate or micropapillate, becoming

mucilaginous when moistened; endotesta pale

brown or red-brown. n = 9. Fig. 26D.

Additional selected specimens examined : Western

Australia. Kalumburu Mission, Apr 1991, Willing

244 (PERTH); Kununurra, Mar 1978, Aplin 6247

(PERTH); Chinatown, Broome, Aug 1985, Kenneally

9390 (PERTH); Onslow, Feb 1985, Dodd 173 & Madin

(PERTH); Carnarvon, Apr 1969, Haw son s.n. (PERTH

2846187). Northern Territory. Cobourg Peninsula;

Black Point, Apr 1993, Cowie 1535 (DNA, MEL);

Nabarlek, Oct 1987, Dunlop 7147 (DNA, MEL);

Bullita Station, Gregory N.P, Feb 1992, Cowie 2388 &

Brocklehurst (DNA); Borroloola townsite. Mar 1979,

Kalotas 239 (DNA); Docker River town. Mar 1980,

Henshall 2894 (DNA). Queensland. Cook District:

Unigan N.R., Weipa, Mar 1990, Forster PIF6507 &

O’Reilly (BRI). Burke District: Warang homestead

site. White Mountains N.P, 57 km by road N of Torrens

Creek, Flinders River catchment, Apr 2000, Thomas

1902 & Thompson (BRI). Leichhardt District:

Springsure Creek, Springsure, Aug 2004, Halford Q8318

(BRI). Wide Bay District: 1.5 km NNE of Didcot, Jan

1996, Forster PIF18301 (BRI, MEL). Moreton District:

Emu Creek, Benarkin S.F., c. 11 km SE of Blackbutt,

Feb 2003, Halford Q7478 (BRI). New South Wales.

Billinudgel, Middle Pocket Road, Lacks Creek, Apr

2003, Forster PIF29321 (BRI); c. 2 miles [c. 3 km] NW

of Coaldale, Jul 1969, Clark 1848 et al. (NSW); Grafton,

500

Feb 1946, Flintoff s.n. (NSW 931); Whip Mountain, near

Macksville, Mar 1981; Ennis s.n. (NSW 373094); 30 km

from Singleton alongside the Putty Road, Mar 2001,

Hosting 2012 (MEL, NSW).

Distribution and habitat : A native of the New

World, now a common pantropical weed. In

Australia, the species has become widespread

across northern Australia from Carnarvon,

WA through the NT to Qld south to Sydney,

NSW (Map 21). It is a common weed of

drainage lines, roadsides, lawns, garden beds

and cultivated land.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from February to June.

Notes: Euphorbia hirta is easily distinguished

from the native Australian Euphorbia species

by its indumentum of long yellowish coloured

hairs on stems and petioles. Euphorbia hirta

is very similar to E. ophthalmica Pers., but

E. hirta has a more robust, erect habit with

generally larger leaves and cyathia in axillary

as well as terminal cymose glomerules.

22. *Euphorbia hyssopifolia L., Syst.

Nat. 10 th edn, 1048 (1759); Chamaesyce

hyssopifolia (L.) Small, Bull. New York Bot.

Gard. 3: 429 (1905). Type: Jamaica, s.d.,

P.Browne s.n. (lecto: LINN [Herb. Linn. No.

630.9] n.v. (image seen) (IDC microfiche 177.

320: II. l\fide Fawcett & Rendle [1920: 339]).

Illustrations: Lin etal. (1991: 232, fig. 10), as

Chamaesyce hyssopifolia ; James & Harden

(1990: 429), as Chamaesyce nutans ; Carolin

& Clarke (1991: 58), as Euphorbia nutans ;

Wilson et al. (1995: 109-110).

Monoecious, annual or sometimes

herbaceous perennial with slender taproot, to

60 cm high, one or a few stems arising from

rootstock. Stems ascending, erect (rarely

prostrate), sparingly to much branched,

smooth, glabrous or sparsely pubescent with

hairs in two longitudinal bands; hairs to 0.4

mm long, weakly spreading, crispate, white.

Interpetiolar stipules broad-triangular,

0.3-0.6 mm long, glabrous; margin lacerate

with the teeth often gland-tipped. Leaves:

petiole 0.5-2 mm long, smooth, glabrous or

with indumentum as for stems; blade narrow-

ovate or narrow-elliptic (rarely slightly

Austrobaileya 8(4): 441-600 (2012)

falcate), 10-30 mm long, 4-11 mm wide,

2.5-5 times longer than wide; adaxial surface

green sometimes with irregularly shaped

reddish blotches or suffused with reddish

tinge, smooth, glabrous or sparsely hairy

with spreading, ± straight hairs to 1 mm long;

abaxial surface pale green, smooth; glabrous

or sparsely hairy with ascending, ± straight

hairs to 1 mm long; base asymmetric with

one side rounded to cordate, the other obtuse;

margin serrulate; apex obtuse to rounded.

Cyathia in lax 2-4 branched dichasial

cymes together with a solitary cyathium at

the distal nodes; peduncles 10-25 mm long;

bracts leaf-like but smaller than primary

stem leaves; cyathial peduncles 1-1.5 mm

long. Involucres turbinate, 0.4-0.9 mm long,

0.8-1.2 mm across; lobes 5, triangular to

subulate, 0.1-0.3 mm long, margin laciniate;

glands 4, shortly stipitate, patelliform, planar

or shallowly concave, transverse-oblong in

outline, 0.1-0.4 mm long, 0.3-0.6 mm wide,

pale green or pink; gland appendages present

and conspicuous to inconspicuous, spreading

radially, transverse-oblong, transverse-

linear or reniform, 0.1-0.4 mm long, 0.3-

0.6 mm wide, white or pink, margin entire;

bracteoles 0.4-0.6 mm long, adnate for c.

2/3 of their length to involucre, free portion

deeply divided into ± linear glabrous or

hairy segments. Staminate flowers 5-12 per

cyathium; pedicels c. 1 mm long; staminal

filaments 0.1-0.4 mm long. Pistillate flowers:

styles 0.2-0.4 mm long, erect to ascending,

smooth, glabrous, each bifid for c. 1/2 of

their length, the apices terete. Capsules

exserted from involucre on pedicel to 2 mm

long, depressed ovate or transversely broad-

elliptic in lateral view, 1.5-2 mm long, 1.5-2

mm across, shallowly to deeply 3-lobate with

keels acute, smooth, glabrous; hypogynous

disc entire. Seeds broad-ovate in outline, 0.9-

1.2 mm long, 0.5-0.9 mm tangentially, 0.7-

0.8 mm radially, tetragonous in cross section;

dorsal faces convex; ventral faces planar

to convex; all faces with 2-4 prominent ±

transverse acute ridges; exotesta thin, of

even thickness over surface, grey-white,

micropapillate, becoming mucilaginous when

moistened; endotesta pale brown to dark

brown. Fig. 26E.

501

Halford & Harris, Euphorbia section Anisophyllum in Australia

Additional selected specimens examined : Western

Australia. Howatharra, SE of Northampton, 2001,

Anon. (PERTH 6096956); Kitchener Street, Victoria

Park, Perth, Apr 1997, Lepschi 3410 (BRI, PERTH);

Quarantine Office, Kewdale, Perth, Apr 2002, Buckley

s.n. (BRI [AQ558138]). Northern Territory. Gove Golf

Club, Booth 2158 (DNA); Tennant Creek township, Nov

1996, Latz 15020 (DNA); Undoolya Road, Alice Springs,

Dec 1991, Thomson 3573 (DNA). Queensland. Burke

District: Normanton, Mar 1999, Waterhouse BMW5136

(BRI). North Kennedy District: Townsville Field

Training Area (Dotswood sector). Camp McAliney (350

man camp), Apr 2001, Waterhouse BMW6195 (BRI); 2

McPhenox Street, Charters Towers, Jan 1986, Bolton 546

(BRI). Leichhardt District: Charles Street, Springsure,

Aug 2004, Halford Q8314 (BRI); 57 km NW of

Wandoan, Jan 2003, Hodgkinson s.n. (BRI [AQ732941]).

Port Curtis District: Heron Island, Oct 1998, Batianoff

981087 (BRI). Darling Downs District: 2 km W of

Macalister on road to Chinchilla, Feb 2004, Halford

Q8140 & Harris (BRI). Moreton District: Chapel

Hill Reservoir, Fleming Road, c. 9 km W of Brisbane,

Oct 1997, Bean 12761 (BRI); Brocks Road, Currumbin

Valley, Feb 2004, Halford 08160 & Edginton (BRI). New

South Wales. Clarrie Hall Dam, SW of Murwillumbah,

Feb 2000, Bean 16009 (BRI); Billinudgel, Middle

Pocket Road, Lacks Creek, Apr 2003, Forster PIF29322

(BRI); Byron Bay, Nov 1987, Coveny 12780 et al. (BRI);

30 km from Singleton alongside the Putty Road, Mar

2001, Hosking 2013 (MEL, NSW); Wentworth Avenue,

East Lakes, Jan 1984, Coveny 11772 & Wilson (NSW).

Australian Capital Territory. Majura Avenue at

junction with Officer Crescent, Dickson, Canberra, Feb

2006, Lepschi 5432 & Mallinson (BRI).

Distribution and habitat : Euphorbia

hyssopifolia is native to the tropic and

subtropic regions of the Americas, now

naturalised in tropical Africa, Asia and

Australia. In Australia, the species is

recorded in all mainland Australian States

except Vic and SA (Map 22). The species

frequently colonises sunny open positions, on

sandy beaches, road verges, garden beds or

disturbed ground.

Phenology : Flowers and fruits have been

collected throughout the year.

Notes : Euphorbia hyssopifolia is recognised

by having usually ascending to erect stems,

broad-triangular interpetiolar stipules 0.3-0.6

mm long with lacerate margins and the teeth

often gland-tipped, glabrous capsules and

seeds surfaces with 2-4 prominent acute ±

transverse ridges.

Collections from NSW here identified as

E. hyssopifolia have been previously named

Chamaesyce nutans (Lag.) Small (James &

Harden 1990). Euphorbia hyssopifolia and

E. nutans Lag. are native to North America

and considered closely related. The main

differences between the two species appear

to be in seed sculpturing (seeds surfaces

transversely ridged for E. hyssopifolia versus

wrinkled for E. nutans ), capsule size (1.5-2.1

mm long for E. hyssopifolia versus 1.9-2.3

mm long for E. nutans) (Burch 1966; Elmore

& McDaniel 1986).

23. Euphorbia inappendiculata Domin,

Biblioth. Bot. 89(4): 309 (1927 ‘1926’);

Chamaesyce inappendiculata (Domin)

D. C.Hassall, Aust. J. Bot. 24: 640 (1976).

Type: [Western Australia.] between the

Ashurton and the De Gray Rivers, s.d .,

E. Clement s.n. (lecto [here designated]: PR

528295; isolecto: PR 528293).

Monoecious, annual or herbaceous perennial

to 30 cm high, few to many stems arising

from slender or thickened woody taproot.

Stems prostrate, decumbent or ascending to

erect, sparingly to much branched, glabrous

or sparsely pilose with spreading, straight or

curved white hairs 0.5-0.9 mm long, smooth.

Interpetiolar stipules subulate, 1—1.8 mm

long, bipartite, glabrous; margin entire,

laciniate or irregularly toothed distally.

Leaves: petiole 0.3-8 mm long, smooth,

glabrous; blade narrow-oblong to oblong,

oblanceolate, obovate, oblong-obovate or

elliptic, 3-14 mm long, 1.5-6.5 mm wide,

1.5-4(47) times longer than wide; both

surfaces green sometimes suffused with red

pigmentation, smooth or minutely papillose;

adaxial surface glabrous; abaxial surface

glabrous or sparsely hairy with spreading,

straight to curved hairs to 0.8 mm long;

base asymmetric with one side cordate or

rounded, the other cuneate or obtuse; margin

entire or sparingly minutely toothed distally;

apex obtuse to rounded. Cyathia solitary

at the nodes, often clustered on short leafy

lateral branchlets with subtending leaves

slightly smaller than the primary stem

leaves; peduncles 0.1-0.8 mm long, smooth,

glabrous. Involucres turbinate, campanulate

or cupuliform, 0.4-0.8 mm long, 0.5-07 mm

across; lobes 5, triangular or subulate, 0.1-0.5

502

mm long, margin entire, laciniate or sparsely

ciliate; glands 4, stipitate, patelliform,

concave with tangential trough or cupuliform

with shallow central pit and sometimes with

thickened margin, transverse-oblong or ±

orbicular in outline, 0.05-0.15 mm long,

0.1-0.2 mm wide, pink or red; appendages

conspicuous or absent, spreading radially,

transverse-linear, 0.07-0.3 mm long, 0.2-0.4

mm wide, white, glabrous, margin entire or

shallowly lobed; bracteoles 0.4-0.6 mm long,

adnate for c. 1/6 of their length to involucre,

free portion entire or divided into a few

subulate segments, glabrous. Staminate

flowers 3-5 per cyathium; pedicels 0.6-1 mm

long; staminal filaments 0.1-0.2 mm long.

Pistillate flowers: styles 0.2-0.5 mm long,

spreading to ascending, smooth, glabrous,

each bifid for 1/3-2/3 of their length, the apices

terete. Capsules exserted from involucre on

pedicel to 1.5 mm long, elliptic to transversely

broad-elliptic (rarely very broad-ovate) in

lateral view, 1.3-1.6 mm long, 1.2-1.8 mm

across, shallowly 3-lobate with keels acute or

obtuse, smooth or minutely papillose (visible

at 40x mag.), glabrous; hypogynous disc

entire. Seeds ovate in outline, 0.9-1.3 mm

long, 0.5-0.8 mm tangentially, 0.5-0.8 mm

radially, tetraquetrous in cross section; dorsal

faces planar, concave or convex; ventral faces

planar or concave; all surfaces smooth or with

faint narrow rounded irregular or transverse

ridges; exotesta thin, of even thickness over

Austrobaileya 8(4): 441-600 (2012)

surface, white or grey-white, microreticulate,

non mucilaginous or becoming mucilaginous

when moistened; endotesta pale brown to

brown.

Distribution and habitat : Euphorbia

inappendiculata is widespread in arid

Australia from the Pilbara, WA through

central NT to north-eastern SA and into

central Qld and western NSW.

Notes : Euphorbia inappendiculata is similar

to E. drummondii but differs in having longer

subulate stipules (1-1.8 mm long versus 0.5-

0.9 mm long for E. drummondii ), generally

smaller glands, 0.1-0.15 x 0.1-0.2 mm (versus

0.15-0.2 x 0.2-0.4 mm for E. drummondii )

that are patelliform, and planar to shallowly

concave (versus glands cupuliform, with a

distinct central pit and thickened rim for E.

drummondii ), and seeds surfaces smooth

or with faint narrow rounded irregular or

transverse ridges (versus seeds surfaces

with 3-6 distinct transverse ridges for E.

drummondii ).

Euphorbia inappendiculata exhibits

some discontinous variation in indumentum

and gland appendage characters with some

geographical discontinuity. This variation

is considered sufficient to warrant formal

recognition of three varieties within this

species which can be distinguished using the

following key.

Key to varieties of Euphorbia inappendiculata

1 Stems pilose; surface of seeds ± smooth or faintly undulate; exotesta not

becoming mucilaginous when moistened 23a. E. inappendiculata var. inappendiculata

1. Stems glabrous; surface of seeds with faint narrow rounded irregular or

transverse ridges; exotesta becoming mucilaginous when moistened.2

2 Gland appendages <0.1 mm long or absent; seeds 1.1-1.3 mm long;

leaf blades 7-9 x 2.2-4 mm.23b. E. inappendiculata var. queenslandica

2. Gland appendages 0.1-0.3 mm long; seeds 1-1.1 mm long; leaf blades

6-13 x 3.2-6.2 mm.23c. E. inappendiculata var. robustior

503

Halford & Harris, Euphorbia section Anisophyllum in Australia

23a. Euphorbia inappendiculata var.

inappendiculata

Annuals or herbaceous perennials. Stems

prostrate, sparsely pilose with spreading,

straight or curved white hairs 0.5-0.9 mm

long. Leaves: petiole 0.3-0.7 mm long; blade

oblong, elliptic or oblong-obovate, 3-9 mm

long, 1.5-5 mm wide, 1.5-2 times longer

than wide; adaxial surface glabrous; abaxial

surface glabrous, or with sparse spreading

trichomes that are straight to curved, to 0.8 mm

long, Involucres campanulate or cupuliform,

0.7-0.8 mm long, 0.5-0.6 mm across; glands,

patelliform, concave with tangential trough,

transverse-oblong in outline, c. 0.15 mm long,

0.1-0.2 mm wide; appendages transverse-

linear, <0.1 mm long or absent. Seeds 0.9-1.1

mm long, 0.5-0.6 mm tangentially, 0.5-0.6

mm radially, tetraquetrous in cross section;

facets smooth or faintly undulate. Fig. 26F.

Additional selected specimens examined : Western

Australia. 10 km W [of] Fortesque River bridge,

Panawonica Railway, May 1991, Thomson 3503 (NT,

PERTH); 5 km N [of] Fortesque River crossing,

Panawonica/Millstream Road, May 1991, Thomson 3505

(NT); Two Mile Creek, Warralong Station, May 1941,

Burbidge 762 (PERTH); Barlee Range N.R., 10.8 km W

of Mt Palgrave, 7.8 km NE of Mt Maitland, 11.3 km ESE

of Mt Padbury, Barlee Range, Aug 1993, van Leeuwen

1437 (PERTH).

Distribution and habitat : Euphorbia

inappendiculata var. inappendiculata is

restricted to the Pilbara, WA, occurring from

Barlee Range to Warralong Station (Map

23a). It is recorded as growing on heavy clay

soils on open plains or gentle slopes.

Phenology : Flowers and fruits have been

collected in May and August.

Typification: Domin (1927) based his

description of E. inappendiculata on material

collected by Clement from “Nordwest-

Australien: zwischen Ashburton - und De

Gray River”. Two sheets of a collection made

by Clement from between the Ashurton and

the De Gray Rivers, WA have been located

amongst material on loan to BRI from PR

and are numbered PR 528295 and PR 528293

respectively. The sheet PR 528295 is here

selected as lectotype of E. inappendiculata

because it is part of the original material, has

morphology that matches the description in

the protologue, and is the best preserved and

more ample of the two specimens.

23b. Euphorbia inappendiculata var.

queenslandica Domin, Biblioth. Bot. 89(4):

309 (1927 ‘1926’). Type: Queensland. [Burke

District:] apud opp. Hughenden et Cloncurry,

February 1910, K.Domin s.n. (holo: PR

528294).

Chamaesyce sp. B.; James & Harden (1990:

428).

Euphorbia “Marree” (F.J. Badman 776);

Barker (1993: 50).

Chamaesyce sp. Marree (F.J.Badman 776);

Barker (2005: 84).

Annuals to 15 cm high, whole plant glabrous.

Stems prostrate, decumbent or ascending to

erect. Leaves: petiole 0.3-0.6 mm long; blade

narrow-oblong, oblanceolate or obovate,

7-9(14) mm long, 2.2-4 mm wide, 2.2-4(47)

times longer than wide; adaxial and abaxial

surfaces glabrous. Involucres turbinate,

0.5-0.6 mm long, 0.5-0.6 mm across; glands

cupuliform, often poorly formed with shallow

central pit and sometimes with thickened

margin, ± orbicular in outline, to 0.1 mm long,

0.1-0.2 mm wide; appendages transverse-

linear, <0.1 mm long or absent. Seeds 1.1-1.3

mm long, 0.5-07 mm tangentially, 0.5-07

mm radially, tetraquetrous in cross section;

facets with faint narrow rounded irregular

ridges, n — 11. Figs 9, 26G.

Additional selected specimens examined: Western

Australia. Calico Creek, 25 km W of Nicholson

Station, May \913,Aplin 5326 (PERTH); 1.1 km S of Mt

Brockman, 22 km W of Hamersley Station homestead,

Sep 2006, Halford Q9262 (BRI). Northern Territory.

Victoria River Crossing, between Top Springs & Timber

Creek, Jul 1974, Carr 2751 & Beauglehole 46530 (MEL,

NSW); Boree Creek, 6 m[iles] [c. 10 km] N [of] No. 48

bore. Brunette Downs, Apr 1970, Latz 576 (AD, DNA);

24 km SW [of] Delmore Downs homestead, Jul 2000,

Albrecht 9255 (NT); 5 km SW [of] Alcoota Station

homestead, Jul 2000, Albrecht 9209 & Latz (NT); Alice

Springs Shooting complex claypan (Conlans Lagoon),

Feb 2007, Albrecht 12101 & Duguid (BRI); 15 km N [of]

Mt Dare homestead, Andado Station, Apr 1997, Latz

15192 (DNA, NT). Queensland. Burke District: ‘Lydia

Downs’ c. 45 miles [c. 72 km] NW of Maxwelton, Jan

1966, Pedley 1958 (BRI). Mitchell District: 36.9 km

SE of Winton on Longreach Road, Sep 1984, Chinnock

6109 (AD); c. 55 miles [c. 88 km] NW of Longreach,

Jul 1974, Hassall 7438 (BRI). Gregory North District:

504

Austrobaileya 8(4): 441-600 (2012)

Fig. 9. Euphorbia inappendiculata var. queenslandica. A. habit x0.5. B. branchlet with cyathia x2. C. stipules x16. D.

cyathia with female flower x32. E. capsule with cyathia xl6. F. cyathial gland, adaxial view x48. G. capsule, oblique

view x24. All from Halford 8128 (BRI). Del. W. Smith.

505

Halford & Harris, Euphorbia section Anisophyllum in Australia

Long Hole, Winton Water Supply, Mar 1998, Forster

PIF22313 & Booth (BRI). Warrego District: 14 km E

of Cunnamulla on road to St George, Feb 2004, Halford

Q8128 & Harris (BRI). South Australia. Site 4W,

Coongie Lakes Survey, Feb 1988, Gillen 1211 (AD);

Peake Creek, on the William Creek - Oodnadatta Road,

Feb 1983, Weber 8947 (AD); Frome Downs Station, Apr

1968, Barker 444 (AD). New South Wales. Walkdens

Plain, Bourke, Mar 1974, Milthorpe & 2179 Cunningham

(NSW); ‘Glen Hope’, Ingleby paddock, c. 10 km N of

White Cliffs, Apr 1976, Lawrie 1893 (NSW); 16 km S

of Bourke on edge of Cobar Road, Mar 1973, Keane 6

(NSW); Fowlers Gap Research Station, Broken Hill, Mar

1968, Young MCB15023 (NSW); Pooncarie site Poo031,

4 km S of ‘Melton Grove’ on the Darnick Road (just N

of Willandra Creek), May 1994, Porteners 9405015 &

Benson (NSW).

Distribution and habitat : Euphorbia

inappendiculata var. queenslandica is

widespread in arid and semi-arid eastern

Australia, extending from central NT and

north-eastern SA into central Qld and

western NSW, with disjunct occurrences in

the Hamersley Range and near the Nicholson

River, WA, and Victoria River, north-western

NT (Map 23b). It grows in Astrebla spp.

grassland, Acacia cambagei low woodland, or

grassland/chenopod shrubland communities

on cracking clay soils on plains or gently

undulating terrain.

Phenology : Flowers and fruits have been

collected throughout the year.

Notes : Euphorbia inappendiculata var.

queenslandica differs from E. inappendiculata

var. inappendiculata in having glabrous

stems, seed surface with faint narrow rounded

irregular ridges, and exotesta that becomes

mucilaginous when moistened. For features

distinguishing E. inappendiculata var.

queenslandica from E. inappendicidata var.

robustior , refer to the ‘Notes’ section under

that variety.

23c. Euphorbia inappendiculata var.

robustior Halford & W.K.Harris, varietas

nova a varietatis aliis E. inappendiculatae

Domin plerumque caulibus robustioribus

appendicibus glandulae majoribus 0.1-

0.3 mm longis (ad vicem <0.1 mm in E.

inappendiculata var. inappendiculata et

var. queenslandica Domin) necnon ab

E. inappendiculata var. inappendiculata

caulibus glabris necnon E. inappendiculata

var. queenslandica seminibus brevioribus

1-1.1 mm longis (ad vicem 1.1-1.3 mm longis)

differt. Typus: Northern Territory. 6 miles [c.

10 km] east [of] No. 7 bore near Elliott, 19

February 1969, P.K.Latz 435 (holo: DNA, iso:

BRI; according to label information on the

holotype sheet there are duplicates (isotypes)

lodged at K n.v., MO «.v.).

Annuals or herbaceous perennials to 30 cm

high, whole plant glabrous. Stems mostly

prostrate or rarely ascending. Leaves: petiole

0.8-2 mm long; blade oblong, obovate, or

oblong-obovate, 6-13.2 mm long, 3.2-6.2

mm wide; 1.8-3 times longer than wide;

adaxial and abaxial surfaces glabrous.

Involucres turbinate, 0.4-0.7 mm long,

0.5-0.7 mm across; glands cupuliform,

often poorly formed, concave, orbicular in

outline, to 0.1 mm long, 0.1-0.2 mm wide;

appendages oblong, 0.1-0.3 mm long, 0.2-

0.4 mm wide. Seeds 1-1.1 mm long, 0.6-

0.8 mm tangentially, 0.6-0.8 mm radially,

tetraquetrous or tetragonous in cross section;

facets with faint narrow rounded irregular or

transverse ridges. Fig. 26H.

Additional selected specimens examined : Northern

Territory. Stuart Highway at Newcastle Creek, Mar

1979, Kalotas 225 (DNA); 7 km E of No. 7 Bore,

Newcastle Waters Station, Mar 1979, Kalotas 208, 210

(DNA); 36 km SW [of] Ucharonidge homestead, Feb

1989, Thomson 3235 (DNA); 5 km SW [of] No. 9 Bore,

Brunchilly Station, Jun 1984, Low 127 (DNA); 30 miles

[c. 48 km] NW of Rockhampton Downs Station, Jul

1948, Perry 1588 (DNA); Alexandria Station, 15 km NW

of homestead. Mar 1981, Henshall 3519 (DNA); 10 km

NNE of Connells Bore (Pictorella Swamp), May 1982,

Latz 9139 (DNA); Connells Lagoon, Sep 1986, Piercey

s.n. (DNA [A0088677], NT 88677). Queensland. Burke

District: c. 40 km NW of Burke and Wills Roadhouse

along Wills Developmental Road (160 km SSE of

Burketown), 1995, Kemp 890 & Fairfax (BRI); 25 km E

of Richmond, May 1974, Byrnes 3017 (BRI).

Distribution and habitat : Euphorbia

inappendiculata var. robustior occurs from

Newcastle Waters, NT, east to Richmond,

north-western Qld (Map 23c). It grows in

mostly Astrebla spp. grassland or chenopod

shrubland communities on clay loam to

cracking clay soils on plains or floodouts.

Phenology : Flowers and fruits have been

collected from February to September.

506

Notes : Euphorbia inappendiculata var.

robustior differs from the other varieties

of E. inappendiculata by its generally more

robust stems and larger gland appendages

(0.1-0.3 mm long versus <0.1 mm long for

E. inappendiculata var. inappendiculata

and E. inappendiculata var. queenslandica).

It also differs from E. inappendiculata var.

inappendiculata in having glabrous stems).

It differs from E. inappendiculata var.

queenslandica in having shorter seeds (1-

1.1 mm long versus 1.1-1.3 mm long for E.

inappendiculata var. queenslandica).

Etymology : The varietal epithet is from Latin

robustus , robust, and the comparative suffix;

-ior more so, to a greater degree, in reference

to the more robust habit of this variety when

compared with the other varieties of this

species.

24. Euphorbia kimberleyensis

B.G.Thomson, Nuytsia 8: 358, fig. 4 (1992).

Type: Western Australia. Palm Woodland,

Mitchell Plateau, West Kimberley, 15 June

1976, K.F.Kenneally 4921 (holo: PERTH; iso:

CANB n.v.,fide Thomson [1992: 358]).

Illustrations : Wheeler (1992: figs 184Q,

185Q, 186Q), as Euphorbia sp. A; Thomson

(1992: 359, fig. 4).

Monoecious, annual to 20 cm high, few to

many stems arising from fibrous roots. Stems

mostly prostrate or occasionally decumbent

to ascending, much branched, moderately

densely hairy on one side, smooth; hairs

ascending-spreading, curved (retrorse) or

occasionally straight, to 1 mm long, white.

Interpetiolar stipules subulate to triangular,

0.8-1.9 mm long, entire or bipartite, glabrous;

margin ± entire or laciniate. Leaves: petiole

0.6-2 mm long, smooth, glabrous; blade

ovate to broad-ovate, occasionally slightly

falcate, 9-27 mm long, 6-16 mm wide, 1.2-

1.8 times longer than wide; adaxial surface

green developing purplish tinge with age;

abaxial surface pale green; both surfaces

smooth, glabrous; base asymmetric with one

side cordate, the other obtuse to rounded;

margin entire or sparingly minutely toothed;

apex obtuse to rounded sometimes with short

acuminate or apiculate tip. Cyathia solitary at

the nodes, sometimes clustered on short leafy

Austrobaileya 8(4): 441-600 (2012)

lateral branchlets with subtending leaves

slightly smaller than the primary stem leaves;

peduncles 1-4 mm long, smooth, glabrous.

Involucres turbinate, 1.5-2 mm long, 1.5-

2.5 mm across; lobes 5, triangular, 0.6-1.1

mm long, margin entire; glands 4, stipitate,

patelliform, planar or shallowly concave,

transverse-oblong in outline, 0.5-0.7 mm

long, 0.8-1.5 mm wide, red; gland appendages

conspicuous, spreading radially, ± obdeltoid,

(0.5)13-2.5 mm long, 1.2-2.5 mm wide, pink

to white, glabrous, margin dentate to laciniate;

bracteoles 1.4-1.9 mm long, adnate for c.

1/2 of their length to involucre, free portion

divided into numerous subulate plumose

segments. Staminate flowers 19-32 per

cyathium; pedicels 1.9-2.1 mm long; staminal

filaments 0.4-0.8 mm long. Pistillate flowers:

styles 0.7-1.5 mm long, erect with spreading

apices, smooth, glabrous, entire, the apices

terete. Capsules exserted from involucre on

pedicel to 5 mm long, broad to very broad-

ovate or broad-elliptic in lateral view, 3.5-4

mm long, 3.8-4 mm across, shallowly

3-lobate with keels acute, papillose, glabrous

or rarely sparsely hairy with hairs confined to

keels, spreading, to 1.5 mm long; hypogynous

disc entire. Seeds ovate in outline, 2-2.8 mm

long, 1.5-1.8 mm tangentially, 1.5-1.8 mm

radially, tetraquetrous in cross section; dorsal

faces ± planar; ventral faces planar or slightly

concave; all surfaces with low rounded

irregular ridges; exotesta of uneven thickness

over surface, distinctly thicker on the ridges,

pale brown, microreticulate, becoming

mucilaginous when moistened; endotesta

dark brown. Fig. 261.

Additional selected specimens examined : Western

Australia. Port Warrender area, Apr 1988, Dunlop

7881 (DNA); area of Carson Volcanics towards Port

Warrender off the later ite plateau. May 1978, Kenneally

6704 (PERTH); eastern margin of Mitchell Plateau, Apr

1991, Willing 326 (PERTH); near Lone Dingo VT, 9 km

SW of Warrender Hill, Jun 1987, Alford 551 (PERTH); 5

km N [of] Theda homestead, Apr 1991, Thomson 3466

(AD); Kalumburu road, 108.9 km by road N of junction

with Gibb River and Ellenbrae Road, Apr 1985, A pi in

721 etal. (PERTH); Silver Gull Creek at spring, c. 14 km

SE of Cockatoo Island, Apr 1983, Fryxell & Craven 3868

(PERTH); Gibbings Island, Buccaneer Archipelago, Jun

1982, Kenneally 8443 (PERTH); Warren Arms fishing

camp, Coppermine Creek, Buccaneer Archipeligo,

Apr 1997, Brockway CB136 (PERTH); Koolan Island,

Jun 1985, Fryxell 4582 et al. (PERTH); garden behind

house, Warren Arms Camp, Apr 1992, Mitchell 2252

507

Halford & Harris, Euphorbia section Anisophyllum in Australia

(PERTH); Crocodile Creek, 5 km E of W end of Koolan

Island, May 1986, Kenneally 9719 (PERTH); Milliwindi

track opposite Bold Bluff, Apr 1988, Cranfield 6382

(PERTH); 5 km N [of] King Leopold Ranges, Apr 1991,

Thomson 3469 (AD, BRI).

Distribution and habitat : Euphorbia

kimberleyensis is restricted to the Kimberley,

WA, occurring from the Buccaneer

Archipelago, east to the King Leopold

Range and north east to the Port Warrender

and near Kalumburu (Map 24). It grows in

eucalypt open woodland or Triodia grassland

communities in rocky sites along creeklines

or on hillsides. The soils are recorded as loam

or red clay. The geological substrate may be

sandstone, laterite or basalt.

Phenology : Flowers and fruits have been

collected from April to June.

Notes : Euphorbia kimberleyensis seems

most closely related to E. schizolepis. It

differs by its glabrous leaves, cyathia and

gland appendages, stems glabrous or with

longitudinal bands of hairs, capsules glabrous

or rarely with a few hairs along the keels

(versus leaves, cyathia, gland appendages,

capsules and stems densely hairy for E.

schizolepis) and fewer staminate flowers per

cyathium (15-25 per cyathium versus 30-40

per cyathium for E. schizoplepis).

25. Euphorbia laciniloba Halford &

W.K.Harris, species nova similis E.

petalae Ewart & L.R.Kerr et E. ophioliticae

(P.I.Forst.) Y.Yang sed ab E. petala stylis

1/3—1/2 longitudinis bifidis (ad vicem

stylis integris vel vix bifidis), appendicibus

glandulae profunde laciniatis lobis acutis

usque attenuatis (ad vicem dentatis vel

breviter lobatis rotundatis obtusisve),

crescentibus plerumque in solo argillaceo (ad

vicem crescentibus in solo arenario necnon

seminibus minoribus 1.1-1.6 x 0.7-1 x 0.7-1

mm (ad vicem seminibus 1.7-1.8 x 1-1.1 x

1-1.1 mm), foliis oblongis usque oblongo-

ellipticis vel ovatis 1.6-2.8 plo longioribus

quam latioribus (ad vicem ellipticis usque

lato-ellipticis 1-1.5 plo longioribus quam

latioribus) differt. Typus: Queensland.

Port Curtis District: Marlborough Creek

crossing, 3.4 km W of Marlborough on road to

Sarina, 28 November 2004, D.Halford Q8783

& G.Batianoff (holo: BRI, iso: MICH).

Monoecious, herbaceous perennial to 10

cm high, many annual stems arising from

woody taproot. Stems prostrate or weakly

ascending, much branched, longitudinally

ridged, glabrous or rarely sparsely hairy;

hairs spreading, straight to 0.15 mm long,

white. Interpetiolar stipules narrowly

to broadly triangular, 0.2-0.9 mm long,

entire or deeply bipartite, glabrous; margin

laciniate. Leaves: petiole 0.4-0.9 mm long,

smooth, glabrous; blade oblong or oblong-

elliptic, 4.3-10.5 mm long, 2.2-5 mm wide,

1.6-2.2 times longer than wide; both surfaces

minutely papillose (visible at 40x mag.),

glabrous; adaxial surface mainly blue-green

with red along the margins; abaxial surface

similar to but paler than adaxial surface; base

asymmetric with one side shallowly cordate,

the other cuneate or obtuse; margin serrulate;

apex rounded. Cyathia solitary at the nodes;

peduncles 1.3-2 mm long, smooth, glabrous,

or with a few isolated hairs to 0.1 mm long.

Involucres campanulate, 0.8-1.1 mm long,

0.8-1.5 mm across; lobes 5, triangular,

0.4-0.5 mm long, margin fimbriate; glands

4, stipitate, patelliform, planar or shallowly

concave, transverse-oblong in outline, 0.2-

0.4 mm long, 0.5-1 mm wide, yellowish

green or red; gland appendages conspicuous,

spreading radially, obdeltoid or oblong, 0.4-1

mm long, 0.8-1.5 mm wide, white or pink,

glabrous, margin laciniate; bracteoles 0.7—1

mm long, adnate for 1/3—1/2 of their length to

involucre, free portion divided into numerous

subulate hirsute segments. Staminate

flowers (5)10-17 per cyathium; pedicels

1-1.2 mm long; staminal filaments c. 0.1

mm long. Pistillate flowers: styles 0.4-0.6

mm long, spreading, smooth, glabrous, each

bifid forl/3-1/2 of their length, the apices

terete. Capsules exserted from involucre

on pedicel to 3 mm long, depressed ovate

in lateral view, 1.6-1.7 mm long, 2-2.1 mm

across, shallowly 3-lobate with keels obtuse,

smooth, glabrous; hypogynous disc entire.

Seeds ovate in outline, (0.9)1.1-1.4 mm long,

(07)0.8-0.9 mm tangentially, 0.7-0.9 mm

radially, tetraquetrous or tetragonous in cross

section; dorsal faces planar or convex; ventral

faces planar or concave; all surfaces with

faint narrow rounded transverse or irregular

ridges; exotesta thin, of even thickness

508

over surface, grey-white, microreticulate,

becoming mucilaginous when moistened;

endotesta brown. Figs 10, 26J.

Additional selected specimens examined : Queensland.

Leichhardt District: 35 km NE of Capella, Mar 1995,

Fensham 2784 (BRI); Warren State Farm, Mar 1920,

Francis s.n. (BRI [AQ202944]); 35 km SE of Springsure,

Jan 1996, Fensham 2387 (BRI); Palmgrove N.P., NW of

Taroom, Bigge Range, Nov 1998, Forster PIF23707 &

Booth (BRI). Port Curtis District: Marlborough area.

Gap Creek Road, Spring Creek, Oct 2001, Batianoff

01102GNB et al. (BRI); near Koolkoorum Creek, S.F.

121, Dawes Range, Feb 1994, Thompson CAL150 et al.

(BRI). Burnett District: Kingaroy, Apr 1947, Smith

3051 (BRI). Maranoa District: N of Mt Rugged, Mt

Moffatt N.P., Dec 1997, Bean 12876 (BRI).

Distribution and habitat : Euphorbia

laciniloba is restricted to central-east and

south-east Qld, from Marlbrough south to the

Bunya Mountains and west to the Carnarvon

Range (Map 25). It grows in eucalypt

woodland or Poa grassland communities on

alluvial flats, hills or plains. The soils are

loam to clays, rarely sand, mostly derived

from basalt substrates, rarely from sandstone

or mudstone.

Phenology : Flowers and fruits have been

collected from October to July.

Notes : Euphorbia laciniloba is similar to E.

petala and E. ophiolitica (RI.Forst.) Y.Yang.

It differs from E. petala by having styles

bifid 1/3 to 1/2 of their length (versus styles

entire or scarcely bifid for E. petala ), gland

appendages deeply laciniate with acute to

attenuate lobes (versus toothed or shallowly

lobed with lobes rounded or obtuse at tip

for E. petala ), capsules only slightly broader

at the base (versus distinctly broader at the

base for E. petala) and growing on mostly

clay soils (versus growing on sandy soils for

E. petala). It differs from E. ophiolitica by

having smaller seeds, 1.1-1.6 x 0.7-1 x 0.7-1

mm (versus 1.7-1.8 x 1—1.1 x 1-1.1 mm for E.

ophiolitica) and leaf blades oblong to oblong-

elliptic or ovate, 1.6-2.8 times as long as wide

(versus elliptic to broad elliptic, 1-1.5 times

long as wide for E. ophiolitica).

Etymology : The specific epithet is from Latin

laciniatus , slashed into narrow divisions with

tapered-pointed incisions, and lobus , lobe, in

reference to the divided gland appendages of

this species.

Austrobaileya 8(4): 441-600 (2012)

26. Euphorbia litticola Halford &

W.K.Harris, species nova quoad habitationem

litoralem foliorum crassitudinem,

cyathiorum dispositionem, seminum

formam, amplitudinem sculturamque similis

E. pallenti Dillwyn sed habitu robustiore

caulibus crassioribus usque 5 mm diam. in

internodiis (ad vicem caulibus usque 3.5

mm) stipulis chartaceis lato-ovatis 1.5-3

mm longis glabris superficiebus ambabus (ad

vicem stipulis ± coriaceis triangularibus 1-1.5

mm longis pubescentibus in superficiebus

adaxialibus) basibus foliorum symmetricis

cordatisque usque auriculatis (ad vicem

basibus asymmetricis hinc cordatis illinc

obtusis usque leviter cordatis) differt. Typus:

Northern Territory. Melville Island, 7 km NW

of Point Elly, 28 November 1989, P.I.Forster

PIF6113 & R.Petherick (holo: BRI; iso: MEL,

according to label information on the holotype

sheet there are duplicates (isotypes) lodged at

CNS [formerly QRS] n.v. & DNA n.v.).

Euphorbia levis var. imbricata Boiss.,

in A.DC., Prodr. 15(2): 13 (1862). Type:

[Northern Territory.] Port Essington, s.d.,

[J.Wi] Armstrongs.n. (holo: K).

Illustrations : Wheeler (1992: figs 183A, 184A,

185A, 186A) as Euphorbia atoto ; Dunlop et

al. (1995: 218, fig. 72), as Euphorbia atoto.

Monoecious, herbaceous perennial to 70(150)

cm high, many stems arising from woody

rootstock, the whole plant glabrous. Stems

decumbent to erect (rarely rhizomatous,

Dunlop 9793 & Wightman [DNA]), much

branched, smooth. Interpetiolar stipules

broad-triangular, 2-3 mm long, chartaceous,

reddish brown, entire or bifid at apex,

glabrous; margin lacerate. Leaves: petiole

1-2 mm long, smooth; blade oblong, ovate

or oblong-elliptic, 18-32 mm long, 8-16

mm wide, 1.8-2.5 times longer than wide;

adaxial surface blue-green sometimes with

reddish tinge, smooth or minutely papillose;

abaxial surface similar to but paler than

adaxial surface; base ± symmetric, auriculate

to cordate; margin entire or serrulate; apex

obtuse to rounded with short apiculate

tip. Cyathia in congested 2 or 3 branched

dichasial cymes together with a solitary

cyathium at the distal nodes; peduncles 5-10

Halford & Harris, Euphorbia section Anisophyllum in Australia

509

Fig. 10. Euphorbia laciniloba. A. habit *0.6. B. branchlet with cyathia *4. C. leaf *6. D. stipules xl2. E. cyathia with

female flower x24. F. capsule with cyathia xl6. G. cyathial gland with appendage, adaxial view x24. H. capsule, top

view xl6.1. capsule, lateral view xl6. All from Halford Q8793 & Batianoff (BRI). Del. W.Smith.

510

mm long; bracts leaf-like but smaller than the

primary stem leaves; cyathial peduncles 1-9

mm long. Involucres turbinate or cupuliform,

1.5-1.8 mm long, 1.6-1.8 mm across; lobes 5,

triangular, 0.6-0.8 mm long, margin laciniate;

glands 4, stipitate, patelliform, planar or

shallowly concave, transverse-elliptic in

outline, 0.4-0.5 mm long, 0.6-1 mm wide,

green or yellowish green; gland appendages

inconspicuous or absent, spreading radially,

transverse-linear or lunate, 0.1-0.3 mm long,

0.9-1 mm wide, white, glabrous, margin

entire or dentate; bracteoles 1.5-1.9 mm

long, adnate for 1/2-2/3 of their length to

involucre, free portion divided into numerous

subulate glabrous segments. Staminate

flowers c. 25 per cyathium; pedicels 1.5-2.2

mm long; staminal filaments 0.6-0.7 mm

long. Pistillate flowers: styles 0.8-1 mm

long, ascending, smooth, glabrous, each bifid

for c. 1/2 of their length, the apices terete.

Capsules exserted from involucre on pedicel

to 6 mm long, very broad-ovate in lateral

view, 3-3.2 mm long, 37-4.2 mm across,

shallowly 3-lobate with keels obtuse, smooth,

glabrous; hypogynous disc entire. Seeds very

broad-elliptic in outline, 1.7-1.8 mm long,

1.4-1.6 mm tangentially, 1.5-1.7 mm radially,

suborbicular in cross section, dorsal and

ventral faces convex, smooth; exotesta thin,

of even thickness over surface, chalky-white,

microreticulate, not becoming mucilaginous

when moistened; endotesta brown. Figs 11,

26K.

Additional selected specimens examined : Western

Australia. Cape Anjo, Jul 1973, Wilson 11303 (PERTH);

Wollaston Island, Jun 1972, Marchant 72/316 (PERTH);

Krait Bay, Cape Voltaire, Bonaparte Archipelago, May

1998, Mitchell 5422 (BRI, PERTH); Vansittart Bay at E

end of airstrip on Anjo Peninsula, Jun 1992, Kenneally

11232 (DNA, PERTH). Northern Territory. New Year

Island, Apr 1995, Booth 518 (DNA); Melville Island,

Penelli Beach, Jun 1992, Cowie 2187 & Cowie (DNA);

Cobourg Peninsula, Trepang Bay, Apr 1993, Cowie 3605

(DNA); Mountnorris Bay, foot of Cobourg Peninsula,

c. 35 km W of Murgenella, Jun 1988, Munir 6125 (AD,

BRI); Black Point, Cobourg Peninsula, Oct 1968, Byrnes

NB1094 & Maconochie (AD, DNA); Annesley Point,

Malay Bay, Jun 1988, Munir 6097 (AD, BRI); Grant

Island, May 1992, Dunlop 8983 (DNA); Murgenella,

Malay Bay, Jul 1985, Wightman 1971 (DNA); Arnhem

Land, S side Anuru Bay, Oct 1992, Cowie 3123 (DNA,

MEL); Cotton Island, May 1992, Cowie 2941 (DNA);

English Company Islands, Truant Island, Jul 1992, Leach

3038 (DNA); Wigram Island, Outstation, Aug 1995,

Austrobaileya 8(4): 441-600 (2012)

Cowie 6062 (DNA); Port Bradshaw, Sep 1993, Dunlop

9793 & Wightman (DNA). Queensland. Cook District:

Aurukun Reserve, 5 km S of Kirke River mouth, Oct

1978, Smyth s.n. (BRI [AQ412998]). Burke District: 10

km W of Massacre Inlet, Wentworth Station, Dec 1984,

Halford 841217 (DNA); Charlie Bush Bay, Mornington

Island, Sep 1981, Fosberg 62011 (BRI).

Distribution and habitat : Euphorbia litticola

occurs in coastal areas from Bonaparte

Archipelago, Kimberley, WA, across the NT

to Aurukun on the west coast of Cape York

Peninsula, Qld (Map 26). It most likely

extends into the Indonesian Archipelago. It

grows on sandy coastal foreshores usually

just above the strand line.

Phenology : Flowers and fruits have been

collected throughout the year, more frequently

from April to July.

Notes : Euphorbia litticola is similar to E.

pallens Dillwyn in its coastal habitat, leaf

thickness, cyathia arrangement and seed

shape, size and sculpturing. It differs from E.

pallens by its more robust habit with thicker

stems (up to 5 mm diameter at internodes as

compared with up to 3.5 mm diameter for E.

pallens ), stipules chartaceous, broad-ovate,

1.5-3 mm long and glabrous on both surfaces

(versus stipules ± leathery, triangular, 1-1.5

mm long and pubescent on the adaxial surface

for E. pallens) and leaf bases symmetrical

cordate to auriculate (versus asymmetrical,

with one side cordate, the other obtuse to

shallowly cordate for E. pallens).

Etymology : The specific epithet is from Latin

littus , sea-shore, beach, and - icola , dweller

or inhabitant, in reference to the where this

species has been recorded growing.

27. Euphorbia macdonaldii Halford &

W.K.Harris, species nova similis E. australi

Boiss. sed stylis integris plerumque

brevioribus 0.2-0.3 mm longis (ad vicem 0.3-

0.6 mm longis bifidis 1/3-2/3 longitudinis)

capsulis lato-usque per lato-ovatis aspectu

laterali manifeste latioribus basim versus (ad

vicem lato ellipticis aspectu laterali manifeste

latioribus aequatorem versus) plerumque

seminibus minoribus 0.7-0.9 x 0.5-0.6 x

0.5-0.7 mm (ad vicem seminibus 0.8-1.6

x 0.6-0.9 x 0.6-0.9 mm) differt. Typus:

Queensland. Cook District: 1 km S of Lappa

511

Halford & Harris, Euphorbia section Anisophyllum in Australia

on Mt Garnet road, 12 April 2005, P.I.Forster e

PIF30732 & K.R. McDonald (holo: BRI, iso: f

MEL, NSW, distribuendi ). 1

Monoecious (rarely dioecious), herbaceous \

perennial to 15 cm high, few to many

stems arising from woody taproot. Stems

prostrate or ascending, much branched,

smooth or sometimes longitudinally ridged,

with a sparse to dense indumentum (rarely r

glabrous); hairs spreading, ± straight, 0.5-

1.0 mm long, white. Interpetiolar stipules

narrow-triangular to triangular, 0.2-07 mm ^

long, bipartite, indumentum as for stems or

glabrous; margin laciniate. Leaves: petiole

0.4-1.2 mm long, smooth, glabrous or with j;

indumentum as for stems; blade oblong,

oblong-elliptic, elliptic to broad-elliptic or

oblong-obovate, 4-9.5 mm long, 2.5-7 mm 1

wide, 1-1.9 times longer than wide; adaxial t

and abaxial surfaces red to pink or green f

with reddish tinge over surface or along \

margin, smooth (rarely papillose, Henshall I

379 [BRI]), glabrous or with a moderately j

dense indumentum; hairs spreading, straight, ^

0.1-0.3 mm long; base asymmetric with one

side cordate to rounded, the other cuneate

to rounded; margin serrulate to serrate

or only toothed distally; apex rounded to

retuse. Cyathia solitary at the nodes, often

clustered on short leafy lateral branchlets .

with subtending leaves slightly smaller than '

the primary stem leaves; peduncles 0.4-0.6

mm long, smooth, glabrous. Involucres ,

turbinate, 0.8-1.2 mm long, 0.7-1.3 mm ^

across; lobes 5, triangular, 0.4-0.5 mm long,

margin entire; glands 4, stipitate, patelliform,

planar or shallowly concave, transverse- e

oblong to transverse-elliptic in outline, 0.2- a

0.4 mm long, 0.3-0.6 mm wide, pink, cream a

to yellow; gland appendages inconspicuous, c

spreading radially, transverse-oblong to

transverse-linear or lunate, 0.1-0.4 mm long,

0.5-0.8 mm wide, pink, glabrous, margin

dentate, irregularly shallowly lobed or entire;

bracteoles 0.1-0.8 mm long, divided into few ,

to numerous subulate glabrous segments. ,

Staminate flowers 5-15 per cyathium;

pedicels 0.8-1.1 mm long; staminal filaments

0.1-0.2 mm long. Pistillate flowers: styles

0.2-0.3 mm long, spreading to ascending,

smooth, glabrous or pubescent abaxially,

entire, the apices terete. Capsules exserted

from involucre on pedicel to 2.5 mm long,

broad to very broad-ovate in lateral view, 1.1-

1.4 mm long, 1.3-1.6 mm across, shallowly

3-lobate with keels obtuse or acute, ± smooth

or minutely papillose, glabrous or sparsely to

densely hairy; hairs white spreading, 0.1-0.3

mm long; hypogynous disc entire. Seeds

ovate in outline, 0.7-0.9 mm long, 0.5-0.6

mm tangentially, 0.5-0.6 mm radially,

tetraquetrous in cross section; dorsal and

ventral faces planar or concave, smooth or

with faint transverse or irregular ridges;

exotesta thin, of even thickness over surface,

grey-white or pale brown, microreticulate,

becoming mucilaginous when moistened;

endotesta red-brown.

Distribution and habitat : Euphorbia

macdonaldii occurs in north-eastern Qld,

from near Chillagoe south to near Mt Coolon

with a disjunct population in the Nicholson

River area, NT.

Notes : Euphorbia macdonaldii is similar to

E. australis but differs by having entire styles

which are generally shorter (0.2-0.3 mm long

versus styles 0.3-0.6 mm long and bifid for

1/3-2/3 of the length for E. australis), broad

to very broad-ovate capsules in lateral view

which are distinctly broader towards the base

(versus broad-elliptic in lateral view and

distinctly broader towards the equator for E.

australis ) and generally smaller seeds (0.7-

0.9 x 0.5-0.6 x 0.5-0.7 mm versus 0.8—1.6 x

0.6-0.9 x 0.6-0.9 mm for E. australis).

As circumscribed here, this species

exhibits discontinuty in density, distribution

and length of the indumentum. Two varieties

are here formally recognised which can be

distinguished using the following key.

Etymology : The specific epithet honours

Keith R. Mcdonald, formerly of the

Threatened Species Unit, Department of

Environment & Heritage Protection, whose

botanical collections have added much to the

knowledge of the flora of Queensland.

512

Austrobaileya 8(4): 441-600 (2012)

Fig. 11. Euphorbia litticola. A. habit *0.4. B. branchlet with cyathia *3. C. leaf *3. D. stipules x6. E. cyathia with

female flower xl2. F. capsule with cyathia x6. G. cyathial gland with appendage, adaxial view x24. H. capsule, top view

x6. I. capsule, lateral view ><6. A-D, F-I from Forster PIF6113 & Petherick (BRI); E from Cowie et al. 2187 (DNA).

Del. W.Smith.

513

Halford & Harris, Euphorbia section Anisophyllum in Australia

Key to varieties of Euphorbia macdonaldii

Stems, leaves and capsules with a moderately dense to dense indumentum;

hairs 0.1-0.5 mm long. 27a. E. macdonaldii var. macdonaldii

Stems, leaves and capsules glabrous or with a sparse indumenutm; hairs 0.5-

1 mm long. 27b. E. macdonaldii var. potentillina

27a. Euphorbia macdonaldii var.

macdonaldii

Euphorbia australis var. canescens Domin,

Biblioth. Bot. 89(4): 310 (1927 ‘1926’). Type:

Queensland. [North Kennedy District:] Mt

Remarkable apud opp. Pentland, Feb 1910,

K.Domin s.n. (holo: PR 528297).

Herbaceous perennials to 10 cm high, many

stems arising from woody taproot. Stems

prostrate or weakly ascending, moderately

dense to dense hairy (hispid); hairs 0.1-0.5

mm long. Leaves: petiole 0.5-1 mm long;

blade oblong-elliptic or oblong-obovate,

4-9.5 mm long, 2.5-5 mm wide, 1.4-1.9

times longer than wide; adaxial and abaxial

surfaces smooth (rarely papillose), with a

moderately dense indumentum consisting

of hairs 0.1-0.3 mm long; margin toothed

distally; apex rounded. Involucres 0.8-1.2

mm long, 0.7-1.3 mm across; glands 0.2-0.4

mm long, 0.3-0.6 mm wide, pink or cream to

yellow; gland appendages transverse-oblong

to transverse-linear, to 0.4 mm long, 0.5-0.8

mm wide, pink, margin dentate or irregularly

shallowly lobed. Staminate flowers 5-15 per

cyathium. Capsules shallowly 3-lobate with

keels obtuse, ± smooth, sparsely to densely

hairy; hairs spreading, 0.1-0.3 mm long. Figs

12, 26L.

Additional selected specimens examined : Northern

Territory. Nicholson River area, Jun 1974, Henshall 379

(BRI, DNA). Queensland. Cook District: 16 kmNW of

Mt Garnet on Lappa Road, Jun 2000, Forster PIF25802

(BRI); 13.5 km S along Lappa to Mt Garnet Road,

Apr 2005, Forster PIF30797 & McDonald (BRI); 11.4

km along Sundown Road from junction with Kennedy

Highway, Apr 2006, McDonald KRM5072 (BRI);

Pinchgut Creek, base of Pinchgut Hill, NE of Chillagoe,

Mar 2005, McDonald KRM3913 & Little (BRI); 36 [km]

SW of Mt Garnet beside the road to ‘Sundown’, Mar

2005, Wannan 3888 (BRI); 11.5 km along Sundown

Road to Almaden/Mt Surprise off Kennedy Highway,

May 2006, Forster PIF31477 & McDonald (BRI); 28.8

km S along Lappa to Mt Garnet Road, Apr 2005, Forster

PIF30802 & McDonald { BRI); 16 km NW of Mt Garnet,

on road to Lappa, Jan 1993, Bean 5462 & Forster (BRI);

Stannary Hills, 11 km S of Mutchilba, Portion 603, May

2006, Forster PIF31590 & McDonald (BRI); Stannary

Hills, 9.5 km S of Mutchilba, Portion 603, May 2006,

Forster PIF31469 & McDonald (BRI); Copperfield

River, Kidston Goldmine Water Supply Dam, Gilbert

Range, Feb 1994, Forster PIF14891 & Bean (BRI). North

Kennedy District: 22.9 km S of turnoff to Einasleigh

on Kennedy Developmental Road, Jul 2000, Cumming

19867 (BRI); Humpybong track. High Range, c. 40 km

SW of Townsville, Feb 1999, Cumming 18617 (BRI); on

track towards lottery turnoff, just W of Warrigal Creek,

7.2 km ESE of Flinders Highway, SW of Pentland, Aug

1988, Cumming 8282 (BRI); Pentland, Jun 1934, Blake

6061 (BRI); 100 km S of Charters Towers on road

to Lornesleigh Station, May 2006, Halford Q9079 &

Batianoff [\3K\)\ Charters Towers, s.d., Plant 190 (BRI);

6 km E of Mt Cooper homestead, Jun 1992, Thompson

CHA133 & Sharpe (BRI). South Kennedy District:

Mt Coolon - Collinsville Road, 0.9 km E of Deception

Creek - W side of road site 96/3, Jan 1996, Champion

1296 & Pollock (BRI).

Distribution and habitat : Euphorbia

macdonaldii var. macdonaldii is confined to

north-eastern Qld, from near Chillagoe south

to near Mt Coolon with a distjunct population

in the Nicholson River area, NT (Map 27a). It

grows in eucalypt woodland communities on

sandy soils most often on rocky granitic hills,

but also on rhyolitic or sandstone hills.

Phenology : Flowers and fruits have been

collected from January to August.

27b. Euphorbia macdonaldii var.

potentillina (Baill.) Halford & W.K.Harris

combinatio nova; Euphorbia australis var.

potentillina Baill., Adansonia 6: 283-284

(1866). Type: Queensland, s.loc., s.d., [E.M]

Bowman s.n. [202/62] (holo: P 698528,

element top left hand corner, n.v. (image

seen); iso: MEL 1560384).

Euphorbia australis var. semiglabra Domin,

Biblioth. Bot. 89(4): 310 (1927 ‘1926’). Type:

Queensland. [Cook District:] apud opp.

Chillagoe, February 1910, K.Domin s.n.

(lecto, [here designated]: PR 528301).

514

Austrobaileya 8(4): 441-600 (2012)

Fig. 12. Euphorbia macdonaldii var. macdonaldii. A. habit ><0.4. B. branchlet with cyathia x8. C. leaf x8. D.

indumentum on lower leaf surface xl6. E. stipules xi6. F. cyathia with female flower x24. G. cyathial gland with

appendage, adaxial view x32. H. capsule with cyathia x24. I. capsule, top view xl6. J. capsule, lateral view xl6. A

from Forster PIF31477 & McDonald (BRI); B-D, F-J from Forster PIF30732 & McDonald (BRI); E from McDonald

KRM5072 (BRI). Del. W.Smith.

515

Halford & Harris, Euphorbia section Anisophyllum in Australia

Herbaceous perennials to 15 cm high, with

few to many annual stems arising from

slightly thickened rootstock. Stems prostrate

or ascending, sparsely hairy (pilose) (rarely

glabrous); hairs 0.5-1.0 mm long. Leaves:

petiole 0.4-1.2 mm long; blade oblong, elliptic

to broad-elliptic, 5.5-9 mm long, 4.5-7 mm

wide, 1-1.5 times longer than wide; adaxial

and abaxial surfaces smooth, glabrous or

with scattered hairs 0.5-1 mm long; margin

serrulate to serrate; apex rounded to retuse.

Involucres 0.9-1 mm long, 1-1.1 mm across;

glands 0.2-0.3 mm long, 0.4-0.5 mm wide,

pink; gland appendages transverse-oblong

or lunate, 0.1—0.2 mm long, 0.7-0.8 mm

wide, pink, margin entire or shallowly lobed.

Staminate flowers 5 per cyathium. Capsules

shallowly 3-lobate with keels acute, smooth

or minutely papillose, glabrous, n = 11. Fig.

26M.

Additional selected specimens examined : Queensland.

Cook District: 6.7 km along Pormpuraaw Road from

Gulf Development Road junction near Musgrave, May

2010, McDonald KRM9190 (BRI); on a hillside to the

E of the road to the OK Mine, on Nychum Station and

15 km WSW of the homestead, Aug 2003, Fox IDF2428

(BRI); 7.2 km by road towards Ootann from junction

with Burke Development Road near Almaden, Jan 2005,

McDonald KRM3512 (BRI); 24 km W of Dimbulah,

May 1976, Hassall 7618 (BRI); 45 km by road S of Mt

Garnet, Feb 2006, McDonald KRM4795 (BRI); Mt Zero

property, off Ewan - Laroona Road, c. 100 km W of

Townsville, Feb 2005, Camming 23180 (BRI).

Distribution and habitat : Euphorbia

macdonaldii var. potentillina occurs from

near Musgrave, Cape York Peninsula, south

to the Paluma Range, Qld (Map 27b). It is

recorded as growing in eucalypt woodland

communities on sandy soils on small hills,

most often on granite but also on rhyolite.

Phenology : Flowers and fruits have been

collected in January, February and August.

Typification: Domin (1927) cited two of his

collections in the protologue of Euphorbia

australis var. semiglabra namely “Chillagoe

(DOMIN II. 1910) ... Mungana (DOMIN II.

1910)”. Two collections, which are considered

as syntypes of the name E. australis var.

semiglabra , have been located amongst

material of Euphorbia on loan to BRI from

PR [a: Euphorbia australis var. typica,

Queensland: in xerodrymio apud opp.

Mungana, Jan 1910, Domin (Iter Australiense

nr 5741), (PR 528300); b: Euphorbia australis

var. typica , Queensland: apud opp. Chillagoe,

Feb 1910, Domin (Iter Australiense nr 5742)

(PR 528301). The collection from near

Chillagoe [PR 528301] is here selected as

lectotype because it is part of the original

material and has morphology that matches

the description in the protologue of this

variety. The syntype collected near Mungana

(PR 528300) is referable to E. schultzii var.

comans.

Notes : Euphorbia macdonaldii var.

potentillina differs from the typical variety by

the characters set out in the key above.

28. Euphorbia maconochieana

B.G.Thomson, Nuytsia 8: 354, 356, fig. 3

(1992). Type: Northern Territory. Cahill’s

Crossing, Victoria River crossing on the

Top Springs to Victoria River Downs Road,

26 March 1990, B.G.Thomson 3486 (holo:

DNA n.v.; iso: AD n.v., BRI, PERTH n.v., fide

Thomson [1992: 354]).

Illustrations : Wheeler (1992: figs 184S, 185S,

186S), as Euphorbia sp. C; Thomson (1992:

355, fig. 3).

Monoecious, annual, few to many stems

arising from the base. Stems prostrate,

sparingly to much branched, ± smooth, with a

sparse to moderately dense indumentum; hairs

± spreading or appressed, straight, crispate or

curved, to 0.2 mm long, white. Interpetiolar

stipules narrow-triangular, 0.4-1 mm long,

bipartite, with indumentum as for stems;

margin entire. Leaves: petiole c. 0.5 mm

long, smooth, with indumentum as for stems;

blade narrow-oblong to oblong, narrow-ovate

to ovate or obovate, 7-16 mm long, 2-7 mm

wide, 1.8-2.7 times longer than wide; adaxial

surface colour unknown, smooth or papillose,

glabrous or with a sparse to moderately dense

indumentum consisting of spreading, straight

or curved hairs to 0.1 mm long; abaxial surface

colour unknown, smooth or papillose, with

a sparse to moderately dense indumentum

consisting of spreading, curved hairs to 0.3 mm

long; base asymmetric with one side cordate,

the other cuneate; margin sparingly minutely

to coarsely toothed or entire; apex rounded,

516

obtuse or acute. Cyathia solitary at the nodes,

often clustered on short leafy lateral branchlets

with subtending leaves slightly smaller than

the primary stem leaves; peduncles 1-2.5

mm long, smooth, with indumenutm as for

stems. Involucres cupuliform, 1.3-1.5 mm

long, 1.5-1.8 mm across; lobes 5, triangular,

0.5-0.8 mm long, margin fimbriate; glands

4, stipitate, patelliform, planar or shallowly

concave, transverse-oblong or transverse-

elliptic in outline, 0.3-0.4 mm long, 0.4-0.5

mm wide, pink to dark red; gland appendages

conspicuous, spreading radially, reniform,

(0.1)0.3-1.5 mm long, 0.5-17 mm wide, white

or rarely pale pink, glabrous, margin entire;

bracteoles 0.9-1.6 mm long, adnate for c.

1/2 of their length to involucre, free portion

divided into numerous subulate hirsute

segments. Staminate flowers 15-32 per

cyathium; pedicels 1-1.4 mm long; staminal

filaments 0.4-0.5 mm long. Pistillate

flowers: styles 0.7-1 mm long, connate at

the base into a column for c. 1/8 of their

length, erect with spreading apices, papillose

proximally, glabrous, each bifid for 1/3-2/3

of their length, the apices terete. Capsules

exserted from involucre on pedicel to 3.5

mm long, very broad-ovate or broad-elliptic

in lateral view, 1.7-1.8 mm long, 1.7-2.2 mm

across, shallowly 3-lobate with keels obtuse,

papillose, glabrous or sparsely hairy; hairs

spreading, c. 0.1 mm long; hypogynous disc

entire. Seeds ovate in outline, 1-1.1 mm long,

0.6-0.8 mm tangentially, 0.6-0.7 mm radially,

tetraquetrous to tetragonous in cross section;

dorsal faces convex; ventral faces planar or

slightly concave; all faces with faint narrow

rounded irregular ridges; exotesta thin, of

even thickness over surface, pale brown,

microreticulate, becoming mucilaginous when

moistened; endotesta brown. Fig. 26N.

Additional selected specimens examined : Western

Australia, drain 7, Packsaddle Creek, Aug 1973,

Kenneally 1941 (PERTH); Smoke Creek, SW of Lake

Argyle, May 1980, Weston 12185 (PERTH); Nicholson

Road, off Duncan Highway, Apr 1993, Done s. n. (PERTH

2984318); 47 km S of Forrest River crossing on Duncan

Highway, Apr 1977, Eichler 22400 (MEL); 3 km SE of

Brooking Gorge, Apr 1988, Cranfield 6438 (PERTH);

Bow River Diamond Mine, 200 km S of Kununurra,

Apr 1991, Barrett BR14 (PERTH); Kununurra, Mar

1978, Aplin 6286b (PERTH). Northern Territory. 10

km NNE of Twins Mount on Palm Creek, Mar 1989,

Austrobaileya 8(4): 441-600 (2012)

Leach 2376 & Dunlop (DNA); 46 miles [c. 74 kn] SW

of Birrimbah Outstation, Jun 1949, Perry 2078 (DNA);

Gregory Creek, Mar 1992, Brocklehurst 615 (DNA);

Auvergne Station, plot 923, Mar 1998, Harwood 437 &

Brocklehurst (DNA); Newry Station, 37 km E of WA/

NT border, along Victoria Highway, Apr 1989, Halford

H61 (BRI, DNA); 5 km W [of] Top Springs Roadhouse,

Mar 1990, Thomson 3489 (BRI); 51 miles [c. 82 km] E

[of] Victoria River Downs, May 1959, Chippendale 6095

(DNA, MEL); 4 miles [c. 6 km] S of Willeroo Outstation,

Jun 1949, Perry 2025 & Lazarides (BRI, DNA, MEL); 3

km W [of] Top Springs Roadhouse, Mar 1990, Thomson

3488 (DNA); Victoria River District, site 30, Apr 1990,

Manning 569 (DNA); Kalkaringi, Mar 1990, Thomson

3491 (DNA); Kelly Station, Mayl994, Egan 3937 (BRI,

DNA); 10 km S [of] Victoria River Crossing, between

Top Springs and Timber Creek, Jul 1974, Carr 2730 &

Beauglehole 46509 (MEL, NSW).

Distribution and habitat : Euphorbia

maconochieana extends from Brooking

Gorge near Fitzroy Crossing, Kimberley, WA,

north east to the Fitzmaurice River and east

to Wavehill Station, in the north-western part

of the NT (Map 28). It grows in grassland or

occasionally in open woodland communities,

on dark heavy clays on plains or low rolling

hills.

Phenology : Flowers and fruits have been

collected from February to August with one

collection in October.

Notes : Euphorbia maconochieana is similar

to E. papillifolia but differs from that species

in having longer styles, distinctly papillose

capsules and smaller seeds.

29. *Euphorbia maculata L., Sp. PI. 455

(1753); Chamaesyce maculata (L.) Small, FI.

S.E. U.S. [Small] 713 (1903). Type: “habitat

in America sepientrionali” (lecto: LINN

lHerb. Linn. No. 630.11 ] n.v. (image seen)

(IDC microfiche 177. 320: III. 3), fide Croizat

[1962: 191]).

Euphorbia supina Rafi, Amer. Monthly Mag.

2: 119 (1817); Chamaesyce supina (Raf.)

Moldenke, Annot. Classified List Moldenke

Collect. Numbers 135 (1939). Type: North

America, not designated.

Illustrations : Lin et al. (1991: 240, fig. 15)

as Chamaesyce maculata ; Weber (1986: 749,

fig. 401K); James & Harden (1990: 429), as

Chamaesyce supina ; Jeanes (1999: 61, fig. 9J).

517

Halford & Harris, Euphorbia section Anisophyllum in Australia

Monoecious, annual to 10 cm high, many

stems arising from slender taproot. Stems

prostrate to weakly ascending, much branched,

with a moderately dense indumentum on one

side, smooth; hairs spreading or ascending-

spreading, curved, to 0.8 mm long, white.

Interpetiolar stipules subulate, 1-2.3 mm

long, deeply bipartite; glabrous or with a few

hairs on margin; margin entire or laciniate

proximally. Leaves: petiole 1-1.8 mm long,

smooth, with indumentum as for stems; blade

narrow-oblong to oblong, 5-13 mm long,

2-4.5 mm wide, 1.7-4 times longer than wide;

adaxial surface green to blue green, usually

with elongated red spot centrally and reddish

tinge along margins, smooth, glabrous; abaxial

surface pale green, smooth, with a sparse to

moderately dense indumentum consisting of

appressed-ascending to spreading, curved

hairs 0.3-0.5 mm long; base asymmetric

with one side rounded to auriculate, the

other cuneate to obtuse; margin serrulate;

apex acute to obtuse or rounded. Cyathia

solitary at the nodes, often clustered on short

leafy lateral branchlets with subtending

leaves slightly smaller than the primary stem

leaves; peduncles 0.4-1 mm long, smooth,

with a few scattered hairs or glabrous.

Involucres turbinate, 0.4-0.8 mm long,

0.5-0.8 mm across; lobes (4)5, triangular,

0.2-0.4 mm long, margin fimbriate; glands

4, stipitate, cupuliform, with shallow central

pit, transverse-oblong in outline, 0.1-0.15

mm long, 0.2-0.3 mm wide, pink, cream

or yellowish green; gland appendages

conspicuous, spreading radially, transverse-

oblong, 0.1-0.3 mm long, 0.3-0.7 mm wide,

white or pink, glabrous, margin entire or

irregularly shallowly lobed; bracteoles

0.5-0.7 mm long, adnate for c. 1/5 of their

length to involucre, free portion divided into

few subulate hirsute segments. Staminate

flowers 4-5 per cyathium; pedicels 0.6-1 mm

long; staminal filaments 0.1-0.15 mm long.

Pistillate flowers: styles 0.3-0.5 mm long,

spreading, smooth, glabrous, each bifid for c.

1/3 of their length, apices clavate. Capsules

exserted from involucre on pedicel to 2.3 mm

long, very broad-ovate to depressed ovate in

lateral view, 1.5-1.6 mm long, 1.5-1.8 mm

across, shallowly 3-lobate with keels acute,

smooth or minutely papillose, with a sparse

to moderately dense indumentum; hairs

appressed, 0.1-0.2 mm long; hypogynous disc

entire. Seeds ovate or broad-ovate in outline,

0.8-1 mm long, 0.6-0.7 mm tangentially,

0.6-0.7 mm radially, tetraquetrous in cross

section; dorsal and ventral faces ± planar or

concave, with 2-5 rounded transverse ridges;

exotesta thin, of even thickness over surface,

pale brown or grey-white, microreticulate,

becoming mucilaginous when moistened;

endotesta red-brown. Fig. 260.

Additional selected specimens examined : Western

Australia. Broome, Sep 1992, Kenneally KFK11346

(PERTH); South Perth, Apr 1982, Perry 1294 (PERTH);

Narrogin, Mar 2002, Warren 654 & Rose (PERTH).

Queensland. Maranoa District: Cavanough Park,

St George, Feb 2004, Halford Q8108 & Harris (BRI).

Darling Downs District: 200 m S of Mackenzie Road

on Stanthorpe bypass, Apr 2002, Halford Q7512 &

Batianoff (BRI). Moreton District: Brisbane Botanic

Gardens, Mt Coot-tha, Jan 1999, Halford Z1753 (BRI).

South Australia. Lofty South, 25 Canopus Avenue,

Hope Valley, Jan 1989, Alcock 11028 (AD, BRI); 59

Thomas Street, Unley, Adelaide, Feb 1984, Symon 13784

(AD, BRI); 21 Para Road, Evanston, Mar 1976, Alcock

5361 (AD). New South Wales. Tamworth City Council

Nursery, Oxley Park, Tamworth, Apr 1999, Hosking 1707

& Bayliss (MEL); Botanic Gardens, Sydney, Feb 1976,

Rodd 3021 (BRI, NSW); Buronga, Silver City Highway,

near the bottle shop at Stanley Wineries, Dec 2000,

Browne 1126 (NSW). Australian Capital Territory.

Australian National Botanic Gardens, Feb 1990, Telford

10899 (AD, MEL). Victoria. Cobram township, Apr

2001, McManus s.n. (MEL 2096409); St James Parade,

Elsternwick, Melbourne, Apr 1985, LeBreton 25 (MEL,

NSW); South Yarra, Royal Botanic Gardens, Mar 1998,

Clarke 2865 (MEL).

Distribution and habitat : Euphorbia

maculata is most likely native to North

America, but has become widely naturalised

around the world. In Australia it is naturalised

in all mainland States (Map 29). It grows in

mostly urban areas on roadsides, in lawns,

garden beds and pathways.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from December to May.

Notes : In Australia, the name Euphorbia

thymifolia L. has often been misapplied

to this species (Anderson 1939; Jacobs

& Pickard 1981; Stanley & Ross 1983).

Euphorbia maculata can be distinguished

from E. thymifolia by having capsules at

518

maturity fully exserted from the involucre

versus the capsules at maturity half included

within the involucre causing the involucre to

split in E. thymifolia. Another morphological

difference between these two species is that

the cyathial gland appendages of E. thymifolia

are generally unequal in size versus mostly

equal forfi. maculata (P. Berry, pers. comm).

We have not observed this difference in the

Australian collections of the two species.

Euphorbia maculata may be confused

with another naturalised weedy species in

Australia, E. prostrata Aiton. It differs from

E. prostrata in having ± appressed hairs

evenly spread over the capsules, whereas

the hairs on the capsules of E. prostrata are

spreading and confined to the keels.

30. Euphorbia mitchelliana Boiss., in

A.DC., Prodr. 15(2): 25 (1862); Chamaesyce

mitchelliana (Boiss.) D.C.Hassall, Aust. J.

Bot. 24: 640 (1976). Type: [Queensland. Port

Curtis District:] Port Curtis [Gladstone area],

November 1847, JMacGillivray 83 (lecto

[here designated]: K 186478, element on left;

isolecto: K 186479).

Monoecious, annual or herbaceous perennial

with woody taproot, 80 cm high, few to many

stems arising from rootstock. Stems prostrate,

decumbent or erect (rarely rhizomatous),

sparsely to densely hairy or glabrous, smooth;

hairs ascending to spreading, crispate, 0.1-

0.8 mm long, white. Interpetiolar stipules

subulate, 0.6-1 mm long, entire or bipartite,

glabrous or pubescent; margin entire or

laciniate. Leaves: petiole 0.8-2 mm long,

smooth, glabrous or with indumentum

as for stems; blade linear to lanceolate,

narrow-oblong to oblong, elliptic to broad-

elliptic or ovate, 7-70 mm long, 1-12 mm

wide, 1.5-30 times longer than wide; both

surfaces smooth; adaxial surface dark green

or subglaucous, glabrous or sparsely pilose

with spreading, straight to crispate hairs

0.4-1.3 mm long; abaxial surface pale green

or glaucous, glabrous or sparsely to densely

pilose with ascending to spreading, crispate

hairs 0.1-1.3 mm long; base symmetric or

asymmetric, cordate or obtuse; margin entire

or minutely serrulate distally with gland-

tipped teeth; apex rounded, obtuse or acute.

Austrobaileya 8(4): 441-600 (2012)

Cyathia in lax terminal dichasial cymes

(sometimes becoming monochasial distally);

bracts leaf-like but much smaller than the

primary stem leaves, subulate to narrowly

triangular; cyathial peduncles 1-6 mm

long. Involucres turbinate or campanulate,

0.6-1.3 mm long, 0.7-1.8 mm across; lobes

5, triangular to subulate, 0.3-0.8 mm long,

margin ciliate or laciniate; glands 4, stipitate,

patelliform, planar or shallowly concave,

transverse-oblong to transverse-elliptic

in outline, 0.1-1.2 mm long, 0.3-1.6 mm

wide, yellowish green; gland appendages

inconspicuous to conspicuous, spreading

radially, broad-obovate, obdeltoid or oblong,

0.1-1.5 mm long, 0.5-2.2 mm wide, pink or

white, glabrous, margin entire or shallowly to

deeply irregularly lobed; bracteoles 0.5-0.6

mm long, adnate for 1/2-2/3 of their length to

involucre, free portion divided into a few to

many subulate glabrous or hirsute segments.

Staminate flowers 5-50 per cyathium;

pedicels 0.8-2.2 mm long; staminal filaments

0.2-0.6 mm long. Pistillate flowers: styles

0.4-1 mm long, spreading, smooth, glabrous,

each bifid for 1/4-2/3 of their length, the apices

terete. Capsules exserted from involucre on

pedicel to 2.7 mm long, transversely broad-

elliptic in lateral view, 1.5-2 mm long, 1.6-

2.3 mm across, shallowly 3-lobate with keels

obtuse, smooth, glabrous or with a sparse to

moderately dense indumentum consisting

of spreading or appressed hairs 0.1-0.4 mm

long; hairs mostly evenly distributed over

capsule rarely confined to keels; hypogynous

disc entire. Seeds ovate to elliptic in outline,

1.2-1.4 mm long, 0.8-1 mm tangentially,

0.8-1 mm radially, tetragonous or trigonal

in cross section; dorsal faces convex; ventral

faces convex or ± planar; all faces with 3-7

prominent narrow rounded transverse ridges;

exotesta thin, of even thickness over surface,

grey-white or pale brown, microreticulate,

becoming mucilaginous when moistened;

endotesta red-brown.

Distribution and habitat : Euphorbia

mitchelliana is widespread across northern

Australia from One Arm Point, WA,

eastward through northern parts of the NT to

Maryborough, in south eastern Qld.

519

Halford & Harris, Euphorbia section Anisophyllum in Australia

Typification: Boissier (1862) in his protologue

of Euphorbia mitcheUiana states “Ad

Port Curtis Australiae (Mitchell No. 231!,

MacGillivray n. 83 et 63 in herb. Kew!)” Four

sheets (five collections) which are considered

as syntypes of the name E. mitcheUiana , have

been located amongst material of Euphorbia

on loan to BRI from K [a: “[Queensland.

Belyando River] bed of large sandy river,

sub-tropical New Holland, 11 Aug 1846,

Lieut.-Col. Sir T.L. Mitchell 231”. There

are two specimens on this sheet: element on

left (K 186533) and an element on right (K

186532). The element K 186533 has attached

to its stem a small label of white paper with

“Euphorbia” and the letters “E.C.T.” written

in pencil. The writing on the label appears to

be in Alan Cunningham’s hand. Although it is

certainly one of the elements used by Boissier

in describing E. mitcheUiana it is clearly not

part of the Mitchell 231 collection and not

from the Belyando River area; b: “Voyage

of Rattlesnake B. 63, Port Curtis, sandy

beaches, Nov./47. J MG [J. McGillivray]” [K

186480]; c: “Vogage of Rattlesnake B. 83,

Port Curtis, Nov./47, JMG[J. McGillivray]”

[K 186478], Also on this sheet is a R.Brown

collection from the Northumberland Isles (K

186477). This is not considered to be part of

the syntype material; d: “Vog of Rattlesnake

B.83, Port Curtis, Dr McGillivray [ex herb

Hook.]” [K 186479], The collection (186478)

is here selected as lectotype because it is part

of the original material, is the more ample

and complete of the five collections and has

morphology that best matches the description

in the protologue of this species. In 1989, B.G.

Thomson annotated the K (186533) collection

as lectotype but this was not published. All

the syntypes are referable to E. mitcheUiana

var. mitcheUiana as applied here.

Notes : Euphorbia mitcheUiana is similar

to E. bifida in seed shape and seed surface

ornamentation but differs from that in having

cyathia in lax terminal dichasial cymes

sometimes becoming monochasial distally.

As circumscribed here, E. mitcheUiana

is morphologically variable. There is

considerable variation in the general habit of

plants, vestiture, leaf shape and size, involucre

shape, and gland appendages shape, size and

degree of lobing. This variation is considered

sufficient to warrant formal recognition of

three varieties within this species which can

be distinguished using the following key.

Key to varieties of Euphorbia mitcheUiana

1 Involucres turbinate <1 mm across; staminate flowers <20 per cyathium;

gland appendages small, to 0.6 mm long, usually narrower than or

equal to gland width, toothed or deeply lobed . . 30a. E. mitcheUiana var. mitcheUiana

1. Involucres campanulate, >1 mm across; staminate flowers usually 20-50

per cyathium; gland appendages large, 0.5-1.5 mm long, usually wider

than gland width, entire or shallowly lobed.2

2 Stems prostrate or decumbent; leaf blades ovate, elliptic to oblong-elliptic,

10-40 mm long, 5-12 mm wide, 1.5-4.5(6) times as long as

wide. 30b. E. mitcheUiana var. filipes

2. Stems erect to ascending, rarely prostrate; leaf blades linear to

narrow-oblong, or narrow-lanceolate, 14-45 mm long, 2-7 mm

wide, 5.5-20 times as long as wide.30c. E. mitcheUiana var. longiloba

30a. Euphorbia mitcheUiana var.

mitcheUiana

Euphorbia mitcheUiana var. hirta Boiss., in

A.DC., Prodr. 15(2): 25 (1862); Chamaesyce

mitcheUiana var. hirta (Boiss.) D.C.Hassall,

Aust. J. Bot. 24: 640 (1976). Type:

[Queensland.] bed of large sandy river

[Belyando River], 11 August 184 6, Lieut.-Col.

Sir T.L.Mitchell 231 (holo: K 186532, element

on the right).

520

Euphorbia mitchelliana var. oblongifolia

Boiss., in ADC., Prodr. 15(2): 25 (1862).

Type: [Australia.] Nova Holl. , s.d., F.Bauer

s.n. (holo: W).

Euphorbia mitchelliana var. cairnsiana

Domin, Biblioth. Bot. 89(4): 307 (1927

T926’). Type: Queensland. [Cook District:]

in silvis mixtis apud opp. Cairns solo arenoso,

December 1909, K.Domin s.n. (lecto [here

designated]: PR 528276).

Euphorbia mitchelliana var. dietrichiae

Domin, Biblioth. Bot. 89(4): 307(1927 ‘1926’).

Type: [Queensland. Moreton District:] prope

Brisbane River, 1863-1865, A.Dietrich 1882

(lecto [here designated]: PR 528274; isolecto:

HBG 516202 n.v. (image seen), LD 1045446

n.v. (image seen)).

Euphorbia mitchelliana var. filifolia Domin,

Biblioth. Bot. 89(4): 307 (1927 ‘1926’). Type:

Queensland. [Cook District:] in xerodrymio

ad pedem montis Metal Mts apud opp.

Chillagoe, February 1910 , K.Domin s.n. (holo:

PR 528279).

Slender annual or herbaceous perennial

with few to many stems arising from woody

rootstock, to 60 cm high. Stems decumbent

to erect (rarely prostrate), sparsely hairy

or glabrous. Leaf blades linear to narrow-

oblong or narrow-lanceolate (rarely elliptic),

15-60 mm long, 1-10 mm wide, (2.5)7-30

times as long as wide, glabrous or sparsely

hairy on both surfaces; apex rounded, obtuse

or acute. Involucres turbinate, 0.6-1 mm

long, 0.7-1 mm across; glands 0.1-0.3 mm

long, 0.3-0.6 mm wide; gland appendages

inconspicuous to conspicuous, broad-

obovate or oblong, 0.1-0.6 mm long, 0.5-0.6

mm wide, pink or white, margin toothed or

deeply lobed. Staminate flowers 5-15 per

cyathium. Capsules glabrous or with a sparse

indumentum consisting of appressed hairs, n

= 8 Fig. 26P.

Additional selected specimens examined : Western

Australia. 95 km W along Gibb River Road from the

Great Northern Highway, Apr 1989, Halford H27 (BRI);

4 km SW of One Arm Point, Apr 1992, Carter 509

(PERTH); 2 km NW One Arm Point, Jan 1989, Carter

346 (DNA, PERTH). Northern Territory. Grant Island,

Apr 1995, Booth 638 (DNA); Douglas Daly Research

Farm, Jan 1998, Michell 503 (DNA); old BHP flying

Austrobaileya 8(4): 441-600 (2012)

strip, Arnhem Land, Jun 1972, Symon 7729 (AD);

Bickerton Island, South Bay, Apr 1993, Cowie 3881 &

Leach (DNA); c. 29 miles [c. 46 km] NE [of] Maranboy

Police Station, Mar 1965, Lazarides 19 & Adams (DNA,

MEL); Rose River, Gulf of Carpentaria, Jul 1972, Dunlop

2701 (DNA); Limmen Bight River upper reaches, Jan

1989, Thomson 2860 (DNA); 80 km SW [of] Elliott,

Feb 1989, Thomson 3210 (DNA). Queensland. Cook

District: 5 km E of Irvinebank near Jumna Mine, Feb

2004, McDonald KRM1750 (BRI); 5.2 km E of Davies

Creek Road from Kennedy Highway, Apr 1992, Neldner

3843 (BRI). Burke District: 75.5 km by road from

junction of Gulf and Burke Development Roads, towards

Croydon, Jan 2005, McDonald KRM3470 (BRI); c. 6 km

SW of Normanton along the road to ‘Mogoura’ Station,

Apr 1974, Pullen 8842 (BRI, NSW). North Kennedy

District: Castle Hill, Townsville, Feb 1992, Bean 4032

(BRI); near Mt Woodhouse, SW of Ayr, Oct 1950, Blake

18657 (BRI). South Kennedy District: just E of Great

Dividing Range, c. 36 km NNW of Yarrowmere Station

homestead, Oct 1983, Henderson H2872 et al. (BRI).

Mitchell District: 37 km W of Jericho on the Jericho

- Barcaldine Road, Jul 1975, Beeston 1249C (BRI).

Gregory North District: Hamilton River, 5 km NW of

‘Toolebuc’, Jun 1978, Purdie 1209 (BRI). Port Curtis

District: near Castle Tower N.P., c. 19.5 km NW of

Bororen, May 1995, Thompson CAL215 & Turpin (BRI).

Distribution and habitat : Euphorbia

mitchelliana var. mitchelliana is widespread

across northern Australia from One Arm

Point, WA, eastward through northern NT

to Maryborough, south-eastern Qld (Map

30a). It grows in a wide variety of Melaleuca/

eucalypt woodland/open forest communities

on rocky hills, plains or coastal dunes. The

soils are mostly sandy in texture and derived

from a variety of substrates.

Phenology : Flowers and fruits have been

collected throughout the year.

Typification: Domin (1827) in his protologue

of E. mitchelliana var. cairnsiana states

“Nordost-Queensland: Mischwalder liei

Cairns, auf Sand (Domin XII. 1909)”. Two

sheets (PR 528273, PR 528276), which are

considered by us as syntypes of the name

E. mitchelliana var. cairnsiana , have been

located amongst material of Euphorbia on

loan to BRI from PR. The collection (PR

528276) is here selected as lectotype because

it is part of the original material, is the more

ample and complete of the two sheets and has

morphology matching the description in the

protologue of this variety. Both syntypes are

referable to E. mitchelliana var. mitchelliana

as applied here.

521

Halford & Harris, Euphorbia section Anisophyllum in Australia

Domin (1827) in his protologue of

Euphorbia mitchelliana var. dietrichiae states

“Queensland: s.L, A. Dietrich No. 640, 873,

925, 988, 1882; Savannenwalder bei Mareeba

und zwischen Chillagoe und dem Walsh River

(Domin II. 1910)”. A total of seven sheets

\A.Dietrich 640 (PR 528284); A.Dietrich 873

(PR 528275); A.Dietrich 925 (PR 528282);

A.Dietrich 988 (PR 528285); A.Dietrich 1882

(PR 528274); Feb 1910, K.Domin s.n. (PR

528271); Feb 1910, K.Domin s.n. (PR 528272),

which are considered by us as syntypes of the

name E. mitchelliana var. cairnsiana, have

been located amongst material of Euphorbia

on loan to BRI from PR. The collection

A.Dietrich 1882 (PR 528274) is here selected

as lectotype because it is part of the original

material, the more ample and complete of the

syntypes seen and has morphology that best

matches the description in the protologue of

this variety. All the syntypes are referable to

E. mitchelliana var. mitchelliana as applied

here.

Notes: Euphorbia mitchelliana var.

mitchelliana has leaf blades that are typically

linear to narrow-oblong, or narrow-lanceolate.

Leaf blades of collections from the central

Qld coast and growing on coastal sands are

generally elliptic in outline and resemble

those of E. mitchelliana var. filipes. Some

representative specimens of this variant are:

Port Curtis District: Statue Bay, c. 5 km SSE

of Yeppoon, Aug 1976, Henderson H2407

(BRI); Curtis Island, N of Southend towards

Connor Bluff, Mar 1966, Blake 22558 (BRI,

MEL), Port Curtis [Gladstone area], Nov

1847, MacGillivray 63 (K).

30b. Euphorbia mitchelliana var. filipes

(Benth.) Halford & W.K.Harris combinatio

et status nova; Euphorbia filipes Benth., FI.

Austral. 6: 51 (1873); Euphorbia macgillivrayi

var. filipes (Benth.) Domin, Biblioth. Bot.

89(4): 311 (1927 ‘1926’); Chamaesyce filipes

(Benth.) D.C.Hassall, Aust. J. Bot. 24: 640

(1976). Type: [Queensland. Burke District:]

Sweers Island, s.d ., D.Henne s.n. (lecto [here

designated]: K 186472; isolecto: K 186469,

MEL 2189021).

Euphorbia alsiniflora Baill., Adansonia 6:

288 (30 July 1866); Chamaesyce alsiniflora

(Baill.) D.C.Hassall, Aust. J. Bot. 24: 640

(1976). Type: [Queensland. Burke District:]

Bentink [Bentinck] Island, s.d., s.coll. (lecto

[here designated]: P 313099; isolecto: G-DC

n.v. (microfiche IDC 800-73. 2419: II. 2)).

Herbaceous perennial to 50 cm high, with

woody taproot, few to many stems arising

from rootstock. Stems prostrate, decumbent

(rarely erect), densely hairy or glabrous.

Leaf blades ovate, elliptic to oblong-elliptic,

10-40 mm long, 3—12 mm wide, 1.5-4.5(6)

times as long as wide; glabrous or sparsely

to densely hairy on both surfaces, margin

entire; apex rounded, obtuse. Involucres

campanulate, 0.8-1.2 mm long, 1.2-1.5 mm

across; glands 0.4-0.5 mm long, 0.4-0.7 mm

wide; gland appendages conspicuous, broad-

obovate, obdeltoid, 0.5-1.5 mm long, 1.1-2.2

mm wide, white, margin entire or shallowly

lobed. Staminate flowers stamens 20-50 per

cyathium. Capsules glabrous or with a sparse

to moderately dense indumentum consisting

of spreading hairs, n = 8.

Additional selected specimens examined : Northern

Territory. Greenhill Island, Apr 1994, Taylor 168

(DNA); North Goulburn Island, May 1992, Dunlop 9052

(DNA); Little Mooroongga Island, Jun 1996, Booth 1981

(DNA); Mouth of Glyde River, Dhipirrinjura, Jun 1996,

Booth 1840 (DNA); Bremer Island, Jul 1992, Leach 2995

(DNA); Nyanantu Creek near mouth, Jan 1989, Thomson

2886 (BRI, MEL, NSW); West Island, Sir Edward Pellew

Group, Aug 1988, Thomson 2694 (BRI); Vanderlin

Island, Sir Edward Pellew Group, Jul 1988, Thomson

2544 (DNA); 13 miles [c. 21 km] SSW [of] Borroloola,

Jun 1971, Dunlop 2202 (DNA); SE of Calvert River

mouth, Jan 1989, Brock 446 (DNA). Queensland. Cook

District: Mapoon Beach, S ofPortMusgrave, Dec 1980,

Morton AM1011 (BRI, MEL); Weipa, Nanam Beach, Jul

1980, Morton AM767 (BRI); Lake Patricia, Weipa, Dec

1993, Forster PIF14412 (BRI); Edward River Mission,

Jun 1968, Pedley 2690 (BRI). Burke District: 77 km

N of Escott homestead by track, towards Point Parker.

Jul 1987, Dalliston HC251 (BRI); Sweers Island, South

Wellesley Islands, southern Guif of Carpentaria, Nov

2002, Thomas SW13 & Pedley (BRI); North Bountiful

Island, South Wellesley Group, Gulf of Carpentaria,

Nov 2002, Thomas B0152 & Pedley (BRI); 10 km W

[of] Massacre Inlet, Wentworth Station, Dec 1984,

Thompson 855 (BRI).

Distribution and habitat: Euphorbia

mitchelliana var. filipes occurs along the

northern coast of Australia from Coburg

Peninsula, NT, eastward to Weipa, Qld (Map

30b). It grows mostly in grassland, open

woodland or on the edge of monsoon vine

522

thicket on sandy soils on coastal floodplain,

dunes or beach ridges.

Phenology : Flowers and fruits have been

collected throughout the year.

Typification: Bentham (1873) in his

protologue of E. filipes states “N. Australia,

Islands of the Gulf of Carpentaria, R. Brown,

Henne; Fitzmaurice river, F. Mueller”. Four

collections (K 186469, 186470, 186471,

186472) which are considered by us as

syntypes of the name E. filipes , have been

located amongst material of Euphorbia on

loan to BRI from K. The collection (K 186472)

is here selected as lectotype because it is part

of the original material, has morphology that

best matches the description in the protologue

of this variety. All of the syntypes [K 186469,

186470, 186471, 186472] are referable to E.

mitchelliana wax. filipes as applied here.

Baillon (1866) cited two collections in his

protologue of Euphorbia alsiniflora namely

“N.? Bentink island (herb. Mus., ex herb. F.

Muell.!). - Martin”, “Mount King, Glensly

river (herb. F. Muell.!)”. Two collections (MEL

68196, P 313099) which are considered by us

as syntypes of the name E. alsiniflora , have

been located amongst material of Euphorbia

on loan to BRI from MEL and P [a: Mount

King, Glensly [Glenelg] River, [Dr. J.]Martin

s.n. (MEL 68196); b: Bentink [Bentinck]

Island, [without date or collector] (P 313099).

The collection from Bentinck Island (P

313099) is here selected as lectotype because

it is the more ample and best preserved of

the syntypes and has morphology that best

matches the description in the protologue

of this species. In 1989, B.G. Thomson

annotated the collection from Mount King,

Martin s.n. [MEL 68196] as lectotype but this

was not published. Due to the poor quality of

the material and lack of seed we are unable

to confidently identify the Martin s.n. [MEL

68196] collection. It is most likely referable to

either E. trigonosperma or E. biconvexa.

Notes: Euphorbia mitchelliana var. filipes

differs from the typical form by having larger

involucres (0.8-1.2 x 1.2-1.5 mm versus

0.6-1 x 0.7-1 mm for E. mitchelliana var.

mitchelliana ), more staminate flowers per

Austrobaileya 8(4): 441-600 (2012)

cyathia (20-50 per cyathia versus 5-15 per

cyathia for E. mitchelliana var. mitchelliana)

and larger gland appendages (0.5-1.5 x 1.1-

2.2 mm versus 0.1-0.6 x 0.5-0.6 mm for E.

mitchelliana var. mitchelliana). For features

distinguishing Euphorbia mitchelliana var.

filipes from E. mitchelliana var. longiloba see

the ‘Notes’ section under that variety.

30c. Euphorbia mitchelliana var. longiloba

Halford & W.K.Harris, varietas nova ab

E. mitchelliana Boiss. var. mitchelliana

involucro majore 1-1.3 x 1.2-1.8 mm (ad

vicem involucro 0.6-1 x 0.7-1 mm longo)

floribus staminatis pluribus 35-50 in quoque

cyathio) ad vicem floribus 5-15 in quoque

cyathio) appendicibus glandulae plerumque

majioribus 0.5-1.5 x 0.5-1.7 mm (ad vicem

appendicibus 0.1-0.6 x 0.5-0.6 mm) necnon

ab E. mitchelliana var. filipedi (Benth.)

Halford & W.K.Harris foliis linearibus usque

angusto-oblongis vel angusto lanceolatis

5.5-20 plo longiorbus quam latioribus (ad

vicem foliis ovatis ellipticis usque oblongo-

ellipticis 1.5-4.5(6) plo longioribus quam

latioribus) differt. Typus: Northern Territory.

40 km W [of] Seven Emus, 22 January 1989,

B.G.Thomson 2948 (holo: BRI, according to

label information on the holotype sheet there

are duplicates (isotypes) lodged at DNA n.v .,

NT n.v.).

Euphorbiapubicaulis S.Moore, J. Bot. 64: 97

(1926). Type: [Northern Territory.] Groote

Eylandt, March 1925, G.H.Wilkins 175 (holo:

BM 812173).

Herbaceous perennial to 80 cm high, with

many stems arising from woody rootstock.

Stems erect to ascending (rarely prostrate

or rhizomatous), glabrous (rarely sparsely

hairy). Leaf blade linear to narrow-oblong,

or narrow-lanceolate, 14-45 mm long, 2-7

mm wide, 5.5-20 times longer than wide;

both surfaces glabrous (rarely sparsely

hairy); margin entire or minutely serrulate

distally with gland-tipped teeth; apex acute.

Involucres campanulate, 1-1.3 mm long, 1.2-

1.8 mm across; glands 0.1-0.5 mm long, 0.3-

0.8 mm wide; gland appendages conspicuous,

broad-obovate, obdeltoid, 0.5-1.5 mm long,

0.5-1.7 mm wide, pink or white, margin entire

or shallowly lobed. Staminate flowers 35-50

523

Halford & Harris, Euphorbia section Anisophyllum in Australia

per cyathium. Capsules glabrous, n = 8.

Additional selected specimens examined : Western

Australia. Dillon’s Spring, Oct 1906, Fitzgerald s.n.

(NSW 612843). Northern Territory. North Goulburn

Island, Apr 1995, Booth 808 (DNA); Tin Camp Creek,

Nov 1991, Brennan 1635 (DNA); 24 miles [c. 38 km]

E [of] OT Downs homestead. Mar 1959, Chippendale

5527 (BR1, DNA, MEL, NSW); Mooroongga Island,

Garmalatjirrna Outstation, Aug 1995, Cowie 5967

(DNA); 10 km S of Roper River mouth, Nov 1987, Dunlop

7364 (DNA); Allia Creek, Feb 1989, Dunlop 7967 & Leach

(BRI, MEL); 25 km E [of] Bulman, Arnhem Land, Jun

1990, Dunlop 8657 & White (AD, BRI, MEL); Banyella,

Blue Mud Bay, May 1993, Dunlop 9419 & Leach

(DNA); 15 km S of Elliott, Jan 1993, Egan 1349 (DNA);

N of Nhulunbuy, Oct 1993, Egan 2698 (BRI, DNA);

Nhulunbuy, mine rehabilitation site, Feb 1982, Him s.n.

(BRI [AQ512374], DNA [D0051128], MEL 1583977);

c. 40 km SSW of Nathan River homestead, Aug 1985,

Latz 10104 (AD, DNA); 7 miles [c. 11 km] NE of Legune

Station, Jul 1949, Perry 2585 & Lazarides (BRI, DNA,

MEL); Sir Edward Pellew Islands, West Island, Jan 1989,

Russell-Smith 6742 & Lucas (MEL); Boomerang Creek,

Merlin Mining Lease, Apr 1996, Smith 3810 (DNA);

40 km NNW [of] Wollogorang homestead, Jan 1989,

Thomson 3021 (DNA). Queensland. Cook District:

Wenlock, Batavia River, Jul 1948, Brass 19689 (BRI);

13.6 km W of the track from Rutland Plains to Inkerman

on the track to White Waterhole, May 1992, Clarkson

9538 & Neldner (BRI). Burke District: Westmoreland,

off road past Hells Gate, May 1997, Forster PIF21071

& Booth (BRI); 169 km S of Normanton, May 1976,

Hassall 7636 (BRI).

Distribution and habitat : Euphorbia

mitchelliana var. longiloba occurs from near

Wyndham WA, through northern NT to

the Qld border, with scattered occurrences

on Cape York Peninsula, Qld (Map 30c). It

grows in eucalypt woodland or open forest

communities on hills or plains on mostly

sandy soils, also recorded growing on coastal

sand dunes.

Phenology : Flowers and fruits have been

collected throughout the year.

Notes : Euphorbia mitchelliana var. longiloba

resembles the typical form in leaf size and

shape but differs from the that by having

larger involucres (1-1.3 x 1.2-1.8 mm versus

0.6-1 x 0.7-1 mm for E. mitchelliana var.

mitchelliana ), more staminate flowers per

cyathia (35-50 per cyathia versus 5-15 per

cyathia for E. mitchelliana var. mitchelliana)

and generally larger gland appendages (0.5-

1.5 x 0.5-1.7 mm versus 0.1-0.6 x 0.5-0.6

mm for E. mitchelliana var. mitchelliana).

Euphorbia mitchelliana var. longiloba differs

from E. mitchelliana var. filipes by having

leaves linear to narrow-oblong, or narrow-

lanceolate, that are 5.5-20 times as long as

wide (versus leaves ovate, ellipitc to oblong-

elliptic, 1.5-4.5(6) times as long as wide for

E. mitchelliana \wc.filipes.

Etymology: The varietal epithet is from Latin

longus , long, and lobus, lobe, in reference to

the longer gland appendages of this variety

compare to the gland appendages of the type

variety.

31. Euphorbia muelleri Boiss., in A DC.,

Prodr. 15(2): 27 (1862). Type: [Northern

Territory.] tropical Australia, s.dF.Mueller

s.n. (holo: K 186485; iso: P 698540 n.v. (image

seen).

Illustration: Dunlop etal. (1995: 218, fig. 72).

Monoecious (rarely dioecious), herbaceous

perennial, few to many annual stems arising

from thickened woody rootstock. Stems

prostrate (rarely erect, Leach 3258 [DNA]),

smooth, glabrous or with a sparse to dense

indumentum; hairs spreading, straight, 0.1-

0.3 mm long, white. Interpetiolar stipules

narrow-triangular to broad-triangular, 0.4-

1.5 mm long, entire, bifid to deeply bipartite,

glabrous or hairy abaxially with hairs as for

stems; margin entire or lacerate with gland-

tipped teeth. Leaves: petiole 0.4-2 mm long,

smooth, glabrous or with indumentum as for

stems; blade ovate to broad-ovate or elliptic

to rotund, (4) 9-20 (27) mm long, (3) 7-16

(18) mm wide, 1-1.4 times longer than wide;

adaxial surface green occasionally with

reddish tinge, smooth, glabrous (rarely with

indumentum as for abaxial surface); abaxial

surface pale green or red, smooth, mostly with

a sparse to moderately dense indumentum

consisting of spreading, straight hairs to 0.2

mm long (rarely glabrous); base asymmetric

with one side rounded to shallowly cordate,

the other obtuse or rounded; margin entire;

apex rounded to obtuse or retuse. Cyathia

solitary at the nodes; peduncles 3-11 mm long,

smooth, glabrous or with indumentum as for

stems. Involucres cupuliform to turbinate,

1.5-2.5 mm long, 1.7-2.8 mm across; lobes 5,

triangular, 0.7-1.4 mm long, margin entire or

524

ciliate; glands 4, stipitate, patelliform, planar

or shallowly concave, transverse-oblong

or reniform in outline, 0.3-0.6 mm long,

0.8-1.5 mm wide, colour unknown; gland

appendages conspicuous, spreading radially,

elliptic, broad-obovate to very broad-obovate

or transverse-oblong, 0.6-1.5 mm long,

1.3-3.5 mm wide, white, glabrous, margin

entire or shallowly lobed; bracteoles 1.7-2

mm long, adnate for c. 1/2 of their length

to involucre, free portion deeply divided

into numerous subulate hirsute segments.

Staminate flowers 50-60 per cyathium;

pedicels 2.1-2.5 mm long; staminal filaments

0.4-0.6 mm long. Pistillate flowers: styles

0.8-1 mm long, ascending, spreading distally,

smooth, glabrous, each bifid for c. 1/2 of their

length, the apices clavate. Capsules exserted

from involucre on pedicel to 10 mm long,

transversely broad-elliptic in lateral view, 2.8-

3.5 mm long, 3.5-3.8 mm across, shallowly

3-lobate with keels obtuse, smooth, glabrous

or sparsely hairy; hairs spreading, to 0.2 mm

long; hypogynous disc entire. Seeds ovate

or broad-ovate in outline, 1.9-2.7 mm long,

1.6-2 mm tangentially, 1.5-1.9 mm radially,

tetragonous in cross section; dorsal faces

convex; ventral faces ± planar; all faces with

faint narrow irregular ridges; exotesta thin,

of even thickness over surface, grey-white,

microreticulate, becoming mucilagionous

when moistened; endotesta brown. Fig. 26Q.

Additional selected specimens examined : Northern

Territory. Finniss River Road to Litchfield N.P., Sep

1992, Leach 3258 (DNA); Lowther Road, Virginia, Nov

1984, Wightman 1769 (DNA); Howard River floodplain,

E side of Gunn Point Road, Mar 2001, Cowie 9059

(BRI); 8 miles [c. 13 km] NE [of] Humpty Doo, Jun

1972, McKean B550 (DNA); Shoal Bay Road, Aug 1973,

Parker 144 (DNA); Koongarra area, Apr 1979, Rankin

2026 (DNA); Muirella Park, Kakadu N.P, Dec 1980,

Dunlop 5623 (DNA); 2 km N of Nabarlek airstrip, Apr

1979, Rankin 2211 (DNA); Nabarlek, Nov 1988, Hinz 56

(DNA); c. 7 km S of Daly River Road from Blackfellow

Creek crossing, Tipperary Station, Aug 1986, Strong

925 (DNA); 1.5 km W [of] Wangi Falls, Oct 1984,

Sivertsen 972 (DNA); Daly River Mission area. Mar

1993, Wightman 6024 (DNA); Litchfield N.P, Lost City

Road, Mar 1994, Egan 3358 (DNA); Litchfield Park,

Jun 1998, Michell & Risler 1588 (DNA); 11.2 miles [c.

18 km] S [of] Batchelor, Mar 1961, Chippendale 7728

(DNA, MEL, NSW); 2 km N of Hayes Creek, Nov

1987, Smith 890 (DNA); Kakadu N.P., 6 km E of South

Alligator River, Old Darwin road, Nov 1986, Cowie 432

(DNA); 35 km SSW of Cooinda on Pine Creek Road,

Austrobaileya 8(4): 441-600 (2012)

May 1980, Craven 5602 (DNA); Mann River; c. 5 km S

[of] headwaters, Jun 1992, Wilson 1451 (DNA); Kakadu

Highway, 600 m from Pine Creek Road junction, Nov

1991, Brennan 1618 (DNA).

Distribution and habitat : Euphorbia

muelleri is restricted to the northern part

of the NT, from the Finniss River area, east

to Nabarlek (Map 31). It grows in eucalypt

woodland/open forest or Melaleuca woodland

communities on mostly sandy soils on rocky

ridges, hillslopes or plateaux.

Phenology : Flowers and fruits have been

collected in March, April, June and from

August to December.

Notes: Euphorbia muelleri is a distinctive

species that bears a general resemblance

to E. pallens but differs from it by having

larger seeds, capsules and cyathia, longer

gland appendages, more staminate flowers

per cyathia, seed surface with faint narrow

irregular ridges and cyathia solitary at the

nodes. It also grows in a different habitat.

32. Euphorbia multifaria Halford &

W.K.Harris, species nova habitu generali

glandulis involucri et glandulae appendicibus

arete similis E. verrucitestae Halford &

W.K.Harris sed capsulis majoribus lato-

ellipticis 1.5-1.7 x 1.5-2.1 mm (ad vicem

capsulis perlato-ovatis 1.3-1.5 x 1.5-1.7 mm)

seminum superficiebus laevibus vel leviter

irregulariter porcatis (ad vicem superficiebus

verrucis irregularibus in seminis superficiebus

ornatis) differt. Aliquae formae similis E.

ferdinandi Baill. sed ab ea distinguendae

seminibus latioribus 1.1-1.4 x 0.7-0.8 mm

(ad vicem seminibus 1.1-1.6 x 0.5-0.7 mm)

capsulis lato-ellipticis latis quam latioribus

1.5-1.9 x 1.5-2.1 mm (ad vicem ellipticis

latis quam manifeste longioribus). Euphorbia

multifaria quondam eadem atque E.

drummondii Boiss. considerata est sed habitu

plerumque minori seminum superficiebus

laevibus vel porcatis leviter irregulariter (ad

vicem superficiebus 3-6 porcis distinctis

transversis) differt. Typus: South Australia.

c. 27 kmN ofKingoonya, 7 September 1966,

N.N.Donner 1684 (holo: AD).

Monoecious, annual or herbaceous perennial

with thickened rootstock, to 10 cm high,

many stems arising from base, the whole plant

525

Halford & Harris, Euphorbia section Anisophyllum in Australia

glabrous. Stems prostrate (rarely ascending r

to erect), smooth or often longitudinally c

ridged. Interpetiolar stipules triangular, r

0.3-1 mm long, deeply bipartite, glabrous; 1

margin laciniate. Leaves: petiole 0.2-0.6 /

mm long, smooth; blade oblong, obovate, /

oblong-obovate or elliptic, 2.8-7 mm long, f

1.6-3.8 mm wide, 1.8-1.9 times longer than 1

wide; adaxial surface dull light blue-green *

often with reddish patches along the midline, 6

smooth (rarely minutely papillose); abaxial (

surface green, smooth; base asymmetric £

with one side rounded to shallowly cordate, ^

the other obtuse to cuneate; margin sparingly ,

minutely toothed; apex rounded. Cyathia 4

solitary at the nodes, sometimes clustered on 1

short leafy lateral branchlets with subtending k

leaves slightly smaller than the primary stem ^

leaves; peduncles 0.3-0.4 mm long, smooth. f

Involucres campanulate or turbinate, 0.5- *

0.6 mm long, 0.6-0.8 mm across; lobes I

5, triangular, 0.3-0.5 mm long, margin *

fimbriate; glands 4, stipitate, cupuliform, c

with distinct central pit and thickened rim, (

transverse-oblong or orbicular in outline, (

0.1-0.3 mm long, 0.2-0.4 mm wide, red; c

gland appendages inconspicuous, spreading £

radially, transverse-linear, <0.1 mm long, c. ^

0.1 mm wide, pink, glabrous, margin entire; c

bracteoles 0.5-0.8 mm long, adnate for c. t

1/5 of their length to involucre, free portion 1

divided into few subulate glabrous or hirsute 1

segments. Staminate flowers 4-10 per p

cyathium; pedicels 0.7-1 mm long; staminal 7

filaments c. 0.2 mm long. Pistillate flowers: ^

styles 0.2-0.3 mm long, erect to ascending,

smooth, glabrous, each entire to scarcely bifid ^

or bifid for 1/3—1/2 of their length, the apices

stout. Capsules exserted from involucre A

on pedicel to 2.7 mm long, broad-elliptic

in lateral view, 1.5-1.9 mm long, 1.5-2.1 ^

mm across, shallowly 3-lobate with keels (

obtuse, smooth (rarely minutely papillose), '

glabrous; hypogynous disc entire (rarely

laciniate). Seeds ovate in outline, 1.1-1.4

mm long, 0.7-0.8 mm tangentially, 0.6-0.7

mm radially, tetraquetrous in cross section;

dorsal faces planar or concave; ventral faces

concave; all faces smooth or with faint narrow 1

irregular ridges; exotesta of even thickness c

over surface or sometimes thicker on ridges,

microreticulate or microgranulate, grey-white

or pale brown, becoming mucilaginous when

moistened; endotesta pale brown to red-

brown. n = 11. Figs 13, 26R, 27A.

Additional selected specimens examined : Western

Australia. 16.2 km W along road to Weano Gorge

from Yampire Gorge to Juna Downs Road, Hamersley

Range N.P., Jun 1989, Trudgen 7000 (PERTH); Lake

Mason Station, Jul 1941, Bennett 3 (PERTH); Queen

Victoria Spring, Mar 1992, Pearson DJP1778 (PERTH);

6 km S of Mundrabilla Hotel, Sep 1984, Downing 987

(PERTH); c. 16 km S of Norseman, Apr 1999, Davis

8807 (PERTH); 44 km NE of Balladoma, Mar 1984,

Keighery 7328 (PERTH); 20.9 km S of Caiguna via

Baxter’s Memorial track, Aug 1983, Fitzgerald s.n.

(PERTH 02846934). Queensland. Warrego District:

46 km SE [of] Charleville along Boatman Road, Mar

1976, Purdie 87 & Boyland (BRI); c. 10 miles [c. 16

km] N of Hungerford, May 1973, Hassall 7331 (BRI).

South Australia. 12 km S of Nefertiti Gate, Mobella

Station, Sep 1995, Badman 8146 (AD); 6 km W of Roxby

Downs, Apr 1989, Badman 2073 (AD); Knowles Cave,

Nullarbor Plain, Feb 1967, Symon 4648 (AD, NSW);

Eyre Peninsula, Yudnapinna, Jul 1966, Rogers 460 (AD);

Koonamore Vegetation Reserve, 60 km N of Yunta, Jun

1968, Carrick 1688 (AD, HO, MEL); c. 5 km ex Yunta

on Waukaringa Road, Mar 1968, Barker 252 (AD);

Quondong, KiKi - Lilydale fence, Apr 1967, Barker 24

(AD). New South Wales. Fowlers Gap, N of Broken Hill,

Oct 1975, Jacobs 2102 (NSW); 10.6 km NE of ‘Glenoca’

homestead, on road to ‘Boorungie’ homestead, 26

km NE of Little Topan Hotel, Oct 1971, DeNardi 881

(NSW); 29 miles [c. 47 km] by roadNNW of Wilcannia,

on road to White Cliffs, May 1969, Briggs 2709 (NSW);

API Gypsum Mine, ‘Marlow’, 23 km NNW [of] Conoble

Railway Station, Mar 1973, Pickard 1944 (NSW); near

Darling River, 16 km NNE of Wentworth and 2 km S of

Tapio Station, Aug 1978, Muir 5837 (MEL). Victoria.

Meridian Road, 13 km S of Benetook, Sep 1981, Corrick

7409 (AD, MEL).

Distribution and habitat : Euphorbia

multifaria is widespread in southern Australia

from near Narrogin and Mount Magnet, WA,

east through southern SA, north-western

Vic and western NSW to near Mitchell, in

southern Qld, with an outlier population in

the Hamersley Range, north-western WA

(Map 32). It commonly grows in shrubland

or woodland communities on sandy soils on

undulating plains or on aeolian dunes. It is

also recorded on shallow soils on rocky slopes

or outcrops, and in chenopod shrubland

communities on clay soils on flats.

Phenology : Flowers and fruits have been

collected throughout the year.

526

Austrobaileya 8(4): 441-600 (2012)

Fig. 13. Euphorbia multifaria. A. habit xl. B. branchlet with cyathia x8. C. leaf x8. D. stipules xl6. E. cyathia with

female flower x24. F. cyathia with capsule xl6. G. cyathial gland with appendage, adaxial view x24. H. capsule, top

view xl6.1. capsule, lateral view xl6. A from Barker 24 (AD); B-I from Dormer 1684 (AD). Del. W.Smith.

527

Halford & Harris, Euphorbia section Anisophyllum in Australia

Notes : Euphorbia multifaria is very similar to

E. verrucitesta in its general habit, involucral

glands and gland appendages. It differs from

E. verrucitesta by its larger broad-elliptic

capsules, 1.5-1.9 x 1.5-2.1 mm (versus

capsules very broad-ovate, 1.3-1.5 x 1.5-1.7

mm for E. verrucitesta) and smooth or faintly

irregularly ridged seed surfaces (versus

irregular wart-like protuberances on the seed

surface for E. verrucitesta).

Some forms of E. multifaria resemble

E. ferdinandi but can be distinguished from

that species by its broader seeds (1.1-1.4 x

0.7-0.8 mm versus 1.1-1.6 x 0.5-0.7 mm for

E. ferdinandi) and broad-elliptic capsules that

are as long as wide, 1.5-1.9 x 1.5-2.1 mm

(versus capsules elliptic, 1.4-2.1 x 1.2-1.7

mm, that are distinctly longer than wide for

E. ferdinandi).

Euphorbia multifaria has been previously

identified as E. drummondii but differs from

that species by having smooth or faintly

irregular ridged seed surfaces (versus seed

surfaces with 3-6 distinct transverse ridges

for E. drummondii) and generally smaller

habit.

As recognised here Euphorbia multifaria

is a morphologically variable complex and

with further collections and study may be

subdivided into a number of taxa. One of

the more recognisable variants has the seed

surface faintly irregularly ridged with a

chalky grey-white endotesta (Fig. 27A). This

form has been recorded from Norseman, WA

east across the Nullarbor Plain to Immarna

Siding, SA (e.g. Davis 8807 , Keighery 7328,

Symon 4648). The more typical observed seed

surface is smooth or faintly undulate (Fig.

26R) which is found throughout the species

range.

Etymology : The specific epithet is from

Latin, multifarius, having great variety, and

is in reference to the variation of this species

seed surface texture, and style architecture.

33. Euphorbia myrtoides Boiss. in A.DC.,

Prodr. 15(2): 15 (1862); Chamaesyce

myrtoides (Boiss.) D.C.Hassall, Aust. J. Bot.

24: 640 (1976). Type: [Western Australia.]

Despard [Depuch?] Island, s.d., [5.] Bynoe s.n.

(holo: K 186487; iso: BM n.v. [image seen]).

Monoecious, herbaceous perennial 10-40 cm

high, few to many stems arising from woody

rootstock, the whole plant glabrous. Stems

ascending to erect (rarely prostrate), sparingly

to much branched, smooth. Interpetiolar

stipules subulate or triangular, 0.3-1 mm

long, bipartite, glabrous; margin laciniate or

lacerate. Leaves: petiole 0.5-2.5 mm long,

smooth; blade ovate, obovate, oblong or

oblong-elliptic, 5-17 mm long, 3-9 mm wide,

1.5-2 times longer than wide; adaxial surface

bright green or yellow-green, smooth; abaxial

surface pale green or grey-green, smooth; base

asymmetric with one side cordate, the other

obtuse to rounded; margin entire or serrulate

to serrate; apex rounded. Cyathia solitary at

the distal nodes, sometimes clustered on short

leafy lateral branchlets with subtending leaves

slightly smaller than the primary stem leaves;

peduncles 0.5-2.5 mm long. Involucres

cupuliform, 0.8-1.1 mm long, 0.8-1.5 mm

across; lobes 5, narrow-triangular to broad-

triangular, 0.3-0.5 mm long, margin entire

or fimbriate; glands 4, stipitate, patelliform,

planar or concave, transverse-elliptic to

reniform in outline, 0.3-0.6 mm long, 0.4-1.2

mm wide, yellow, yellow-green or red; gland

appendages absent or conspicuous, spreading

radially, transverse-oblong, 0.1-0.6 mm long,

1-1.3 mm wide, white, pink or red, glabrous,

margin entire or dentate to shallowly lobed;

bracteoles 1-1.2 mm long, adnate for 1/3—

1/2 of their length to involucre, free portion

divided into few to numerous subulate

glabrous or hirsute segments. Staminate

flowers 12-30 per cyathium; pedicels 1.1-1.5

mm long; staminal filaments 0.1-0.4 mm long.

Pistillate flowers: styles 0.5-0.9 mm long,

connate at the base into a column for c. 1/5

of their length, spreading, smooth, glabrous,

each bifid for 1/4—1/2 of their length, the apices

terete. Capsules exserted from involucre on

pedicel to 4.5 mm long, elliptic to broad-

elliptic in lateral view, 2.2-2.8 mm long, 2.2-

2.6 mm across, shallowly 3-lobate with keels

obtuse, smooth, glabrous; hypogynous disc

entire. Seeds ovate in outline, (1.3)1.5-2 mm

long, 0.8-1.1 mm tangentially, 0.8-1.1 mm

radially, tetraquetrous in cross section; dorsal

faces planar or convex; ventral faces planar

to concave; all faces with faint to prominent

528

narrow irregular ridges; exotesta thin, of even

thickness over surface, white or pale brown,

microreticulate, becoming mucilaginous

when moistened; endotesta brown. Fig. 27B.

Additional selected specimens examined : Western

Australia. Pender Bay, Dampierland, 4.5 km NE [of]

Cape Borda, Mar 1989, Keighery 10573 (PERTH);

1 km from Port Smith camping area. Port Smith, Apr

1992, Zich 154 (MEL, NSW); 1 km W of Mandora

Station homestead, Aug 1997, Mitchell PRP1779 (BRI,

PERTH); 21 km NE of Sandfire Roadhouse, Great

Northern Highway, Sep 1978, Beauglehole ACB59327 &

Errey (PERTH); N of Dragon Tree Soak, Great Sandy

Desert, Aug 1977, George 14770 (PERTH); Back Beach,

Karratha, July 1981, Craig 265 (PERTH); Onslow,

Feb 1985, Dodd 171 & Madin (PERTH); Onslow, Sep

2006, Halford Q9294 (BRI); near Thels Well, Mardie

Station, Aug 2002, Thompson JPS164 (PERTH); near

Lakes Percival and Wooloomba, Aug 1962, Johnson s.n.

(DNA D9762); Tobin Lake, Great Sandy Desert, May

1979, George 15641 (PERTH); near Norcape Lodge,

Exmouth, Jul 1977, McFarland s.n. (BRI [AQ234287]);

near S boundary of Cane River Station, 80 km SE of

Onslow, May 1999, Edinger 1495 (PERTH); Lake Auld,

July 1967, George 9144 (PERTH); Kennedy Range

N.P, 30 km NE of Binthalya, Aug 1994, Keighery &

Gibson 1518 (PERTH); Muggon Station, N boundary

of Spinifex Paddock, 3.7 km W of Spinifex Well, Sep

1999, Patrick 3202 et al. (PERTH); Cape Boileau, 35 km

N [of] Broome, May 1991, Thomson 3497 (DNA); Port

Hedland, May 1991, Thomson 3665 (DNA); Anketell

Ridge, Great Sandy Desert, May 1979, Mitchell 1160

(DNA).

Distribution and habitat : Euphorbia

myrtoides occurs in north-western WA from

Carnarvon to Derby and inland to the Great

Sandy Desert (Map 33). It grows in open

shrubland, or Spinifex/Triodia grassland

communities on crests or upper slopes of

coastal beach dunes and inland sand dunes.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from April to September.

Notes : The holotype at Kew has a label with

hand written annotation in ink “Euphorbia,

Despard Island, NW. Aust., Bynoe” This

locality appears to be erroneous. There is no

record of an island of this name along the WA

coast. Benjamin Bynoe was the surgeon on

the voyage of the Beagle (1937-1843). Hall

(1978) noted that Bynoe chiefly collected at

Depuch Island and in the Abolhos in north¬

western Australia. Depuch Island is the most

likely collection site for the type specimen.

The isotype in the BM has a printed label

Austrobaileya 8(4): 441-600 (2012)

“VOYAGE OF H.M.S. BEAGLE, 1839-40

PRESENTED 1842” with a hand annotation

“Depuch Island, Bynoe Coll., Capts. Wickham

& Stokes”.

There is variation in the prominence and

degree of seed surface sculpturing and habit,

but the variation grades from one form to

another form.

Euphorbia myrtoides is morphologically

most similar to E. wheeleri but can be

distinguished from that by its faint to

prominent narrow irregular ridges on the seed

surface (versus foveate to reticulate-foveate

seed surface for E. wheeleri).

Some of the southern populations

of Euphorbia myrtoides approach E.

sharkoensis in general facies; however, it

can be distinguished from that species by its

generally larger capsules and seeds.

34. Euphorbia obliqua Endl., Prodr. FI.

Norfolk. 85 (1833); Chamaesyce obliqua

(Endl.) J.Florence, Bull. Mus. Natl. Hist. Nat.,

B, Adansonia , 18(3-4): 241 (1996). Type:

Norfolk Island, s.d., F.L.Bauer s.n. (holo: W

n.v.).

Illustrations : Green (1993: 315, fig. 2, Bl-3;

1994: 236, fig. 47, E-G).

Monoecious, herbaceous perennial, many

stems arising from thickened woody

rootstock. Stems prostrate, sparingly to much

branched, smooth, glabrous. Interpetiolar

stipules triangular to broad-triangular, 0.6-1

mm long, entire or bifid, glabrous abaxially,

hairy adaxially with appressed hairs c. 0.1 mm

long; margin dentate. Leaves: petiole 1-2 mm

long, smooth; blade elliptic to broad-elliptic,

9-20 mm long, 6-10 mm wide, 1.2-1.8 times

longer than wide; adaxial surface green to

blue-green, smooth, glabrous, abaxial surface

pale blue-green, smooth, glabrous, base

asymmetric with one side cordate, the other

obtuse to rounded or cordate; margin entire;

apex obtuse to rounded. Cyathia solitary at

the distal nodes; peduncles 2-3 mm long.

Involucres turbinate, 1.3-2 mm long, 1-1.5

mm across; lobes 5, triangular, 0.4-0.5 mm

long, margin entire or laciniate; glands 4,

stipitate, patelliform, ± planar, transverse-

oblong in outline, 0.3-0.4 mm long, 0.6-0.7

529

Halford & Harris, Euphorbia section Anisophyllum in Australia

mm wide, yellowish green; gland appendages 1

conspicuous, spreading radially, transverse- I

linear or lunate, c. 0.1 mm long, 0.6-0.9 I

mm wide, white, glabrous, margin entire; 1

bracteoles 1.3-1.5 mm long, adnate for 1/3- c

1/2 of their length to involucre, free portion c

divided into numerous subulate hirsute I

segments. Staminate flowers 15-25 per §

cyathium; pedicels 1.1-1.6 mm long; staminal

filaments 0.5-0.6 mm long. Pistillate ^

flowers: styles 0.5-0.6 mm long, connate c

at the base into a column for c. 1/8 of their j

length, spreading, smooth, glabrous, each ^

bifid for 1/2-2/3 of their length, the apices ^

terete. Capsules exserted from involucre on t

pedicel to 4 mm long, very broad-ovate or 8

transversely broad-elliptic in lateral view,

2.2-2.3 mm long, 2.5-3 mm across, shallowly

3-lobate with keels obtuse, smooth, glabrous; 1

hypogynous disc entire. Seeds very broad- s

elliptic in outline; 1.4-1.5 mm long; 1.2-1.4 r

mm tangentially, 1.2-1.3 mm radially, t

suborbicular in cross section; dorsal and f

ventral faces convex, smooth; exotesta thin, (

of even thickness over surface, chalky-white, /

microreticulate, not becoming mucilaginous

when moistened; endotesta pale brown. Fig. ,

27C. f

Additional selected specimens examined : Queensland. j

North Kennedy District: Holbourne Island, Great j.

Barrier Reef, 39 km from Bowen, Mar 1971, Heatwoie

s.n. (BRI [AQ7736]); Hayman Island, Jun 1934, While c

10181 (BRI); Deloraine Island, Nov 1985, Batianoff3479 C

& Dalliston (BRI). South Kennedy District: Penrith £

Island, 70 km E of Mackay, Nov 1986, Batianoff 6040 ^

(BRI). Port Curtis District: North West Island, Aug

1968, Baxter 977 (BRI); Masthead Island, May 1998, i

Batianoff980527 (BRI); Masthead Island, Great Barrier ^

Reef, Capricorn Group, c. 56 km NE of Gladstone, Dec

1970, Heatwoie s.n. (BRI [AQ6541]); Tryon Island, Great ^

Barrier Reef, Capricorn Group, c. 106 km from Yeppoon, <-

Aug 1971, Heatwoie s.n. (BRI [AQ7777]); Capricorn

Islands, Great Barrier Reef, Jan 1932, Macgillivray r

s.n. (BRI [AQ202444]); North West Island N.P, Great

Barrier Reef Marine Park, May 2000, Batianoff205031 C

(BRI); North West Island, Aug 1979, Nugent s.n. (BRI ^

[AQ454415]); Wilson Island, Great Barrier Reef, 'N

Capricorn Group, c. 91 km NE of Gladstone, Oct 1969, q

Cameron s.n. (BRI [AQ7790]); Wreck Island N.P, ^

Great Barrier Reef Marine Park, May 2000, Batianoff

205071 (BRI, NSW); Heron Island, Capricorn Group, (

Jun 1973, Fosberg 55079 (BRI); Heron Island, Dec 1983, I

Smith 834 & Heatwoie (BRI); Tryon Island, May 2000, \

Batianoff205042 (BRI); West Hoskyn Island N.P., May /

2000, Batianoff205137 (BRI, NSW). V

Distribution and habitat : In the Western

Pacific Euphorbia obliqua occurs on Norfolk

Island, Vanuatu and New Caledonia (Green

1994). In Australia it occurs along the east

coast of Qld from Bowen to Gladstone on

continental islands and coral cays of the Great

Barrier Reef (Map 34). It grows on coastal

sands on frontal beach dunes.

Phenology : Flowers and fruits have been

collected throughout the year.

Notes: Euphorbia obliqua , E. litticola, E.

pallens and E. psammogeton all grow in

very similar sandy foreshore habitats along

the north and east coast of Australia but are

allopatric in their distributions.

Euphorbia obliqua can be confused with

E. psammogeton. It differs from that by its

smooth seeds (versus faint irregular narrow

rounded ridges), smaller gland appendages,

to 0.1 mm long (versus 0.2-0.4 mm long

for E. psammogeton ), and solitary cyathia

(versus cyathia in dichasial cymes for E.

psammogeton).

In Qld, in the past, Euphorbia obliqua has

been confused with E. pallens (previously

referred to by the name E. atoto G.Forst.).

It differs from that by its more slender

habit, cyathia solitary in upper axils (versus

cyathia arranged in terminal or axillary lax to

congested dichasial cymes for E. pallens) and

generally smaller leaves (9-20 x 6-10 mm

versus 24-35 x 11-22 mm for E. pallens).

35. Euphorbia occulta Halford &

W.K.Harris, species nova similis aliquantum

E. careyi F.Muell. a qua glandularum

appendicibus erectis cuculliformibus (ad

vicem appendicibus aut absentibus aut

radial iter extendentibus) glandis involucribus

cupuliformibus lacuna centrali instructis (ad

vicem appendicibus patelliformibus planis

vel leviter concavis) bracteis involucribus

obovatis (ad vicem bracteis oblongis

usque triangularibus distinguenda. Typus:

Queensland. Burke District: 12.3 km along

Lake Moondara Road from Barkly Highway,

N of Mt Isa, 1 July 2011, D.Halford QM524A

(holo: BRI, iso: DNA, K, MEL, MICH, NT,

distribuendi).

530

Dioecious, herbaceous perennial to 20 cm

high, with many stems arising from woody

rootstock. Stems decumbent to erect, much

branched, smooth, with moderately dense

to dense indumentum; hairs spreading,

straight, to 0.1 mm long, white. Interpetiolar

stipules triangular, c. 0.1 mm long, bifid or

deeply bipartite, with indumentum as for

stems; margin ± entire. Leaves: petiole 0.5-1

mm long, smooth, with indumentum as for

stems; blade elliptic to broad-elliptic or ±

rotund, 4-5 mm long, 2.8-5 mm wide, 1.2-

1.5 times longer than wide; adaxial surface

yellow-green, smooth, with moderately

dense indumentum consisting of spreading,

straight, hairs to 0.1 mm long; abaxial

surface yellow-green, minutely papillose,

with indumentum as for adaxial surface; base

asymmetric with one side cordate to obtuse,

the other rounded; margin sparingly serrulate

distally; apex rounded to retuse. Cyathia

solitary at the nodes; peduncles 0.2-0.5 mm

long. Involucres turbinate, 0.7-1 mm long,

c. 1 mm across; lobes 5, obovate, 0.8-1 mm

long, margin laciniate distally; glands 4,

sessile, cupuliform, with distinct central pit,

transverse-oblong in outline, c. 0.1 mm long,

0.4-0.5 mm wide, yellow; gland appendages

conspicuous, erect, transverse-oblong,

0.3-0.5 mm long, 0.7-0.9 mm wide, yellow,

glabrous, margin irregularly lobed distally;

bracteoles 1.3-1.5 mm long, adnateforc. 1/3 of

their length to involucre, free portion divided

into numerous subulate glabrous segments.

Staminate flowers c. 20 per cyathium;

pedicels c. 0.8 mm long; staminal filaments c.

0.3 mm long. Pistillate flowers: styles 0.7-1.2

mm long, connate at the base into a column

for c. 1/4 of their length, erect to spreading

and slightly recurved distally, smooth,

sparsely hairy, each bifid for 1/3—1/2 of their

length, the apices terete. Capsules exserted

from involucre on pedicel to 1.5 mm long,

broad-elliptic or transversely broad-elliptic

in lateral view, 2-2.5 mm long, 2-2.6 mm

across, shallowly 3-lobate with obtuse keels,

smooth, with moderately dense indumentum;

hairs spreading, c. 0.1 mm long; hypogynous

disc entire. Seeds ovate in outline, 1.3-1.5

mm long, 0.7-0.8 mm tangentially, 0.8-0.9

mm radially, tetragonous in cross section;

Austrobaileya 8(4): 441-600 (2012)

dorsal faces ± planar; ventral facet concave;

all faces with distinct rounded irregular

ridges; exotesta thin, of even thickness over

surface, white, microreticulate, becoming

mucilaginous when moistened; endotesta pale

brown. Figs 14, 27D.

Additional specimens examined : Queensland. Burke

District: 12.3 km along Lake Moondara Road from

Barkly Highway, N of Mt Isa, Jul 2011, Halford OM524B

(BRI, DNA, MEL, MICH); Rifle Creek Station, S of Mt

Isa, May 2005, Booth 3536 & Kelman (BRI); Rifle Creek,

5 of Mt Isa, May 2005, Booth 3536A & Kelman (BRI);

10.6 km ENE of Rifle Creek Dam, Aug 2001, Kelman

6 Kelman s.n. (BRI [AQ641966]); Rifle Creek Station,

8 km ENE of River Creek Dam, Apr 2002, Kelman s.n.

(BRI [AQ729647]).

Distribution and habitat : Euphorbia occulta

is known only from the rocky hills in the

vicinity of Mtlsa, north-western Qld (Map 35).

It grows in low open woodland of Eucalyptus

leucophloia Brooker and Corymbia

aparrerinja K.D.Hill & L.A.S.Johnson with

a ground layer usually dominated by tussock

grasses or Triodia pungens on stony skeletal

loam soils.

Phenology : Flowers and fruits have been

collected in May, July and August.

Notes : Euphorbia occulta somewhat

resembles E. careyi from which it can be

distinguished by its erect hood-like gland

appendages (versus gland appendages

spreading radially or absent for E. careyi ),

cupuliform involucral glands with distinct

central pit, (versus patelliform with planar

or shallowly concave for E. careyi ), and

obovate involucral bracts (versus ± oblong to

triangular for E. careyi ).

Etymology: The specific epithet is from

Latin, occultus , secret, hidden, in reference to

the concealment of the involucral glands by

the erect hood-like gland appendages of this

species.

36. Euphorbia ophiolitica (P.I.Forst.) Y.Yang,

Taxon 61: 783 (2012); Chamaesyce ophiolitica

P.I.Forst., Austrobaileya 5: 711-712 (2000).

Type: Queensland. Port Curtis District:

First Sugarloaf, 9.5 km W of Canoona,

1 March 1994, P.I.Forster PIF15042 &

A.R.Bean (holo: BRI; iso: AD n.v., DNA n.v.,

MEL n.v., fide Forster [2000: 711]).

Halford & Harris, Euphorbia section Anisophyllum in Australia

531

Fig. 14. Euphorbia occulta. A. branchlet with male cyathia x8. B. branchlet with female cyathia x8. C. leaf x8. D.

stipules xl6. E. cyathia with male flowers xl6. F. cyathia with female flower xl6. G. involucral bract x32. H. cyathial

gland with appendage, adaxial view xl6.1. cyathial gland with appendage, lateral view xl6. J. capsule with cyathia xl2.

K. capsule, top view xl2. A, E & G from Booth 3536A & Kelman (BRI); B-D, F & H-K from Booth 3536 & Kelman

(BRI). Del. W.Smith

532

Illustration : Forster (2000: 713, fig. 1).

Monoecious or dioecious, annual or

herbaceous perennial with slender woody

taproot, few to many stems arising from

rootstock. Stems prostrate, sparingly to

much branched, longitudinally ridged, with

a moderately dense indumentum or glabrous;

hairs spreading, ± straight, 0.1-0.4 mm

long, white. Interpetiolar stipules subulate

to triangular, 0.2-0.6 mm long, entire or

bipartite, glabrous or pubescent; margin

laciniate. Leaves: petiole 0.5-1 mm long,

smooth, with indumentum as for stems or

glabrous; blade elliptic or broad-elliptic,

5-10 mm long, 4-8.5 mm wide, 1—1.5 times

longer than wide; adaxial surface glaucous,

blue-green sometimes red along margin,

minutely papillose (visible at 40 x mag.), with

moderately dense indumentum consisting of

spreading, straight hairs 0.1-0.3 mm long or

glabrous; abaxial surface pale blue-green,

with surface texture and indumentum as for

adaxial surface; base asymmetric with one

side cordate, the other obtuse to rounded;

margin serrulate or sparingly minutely

toothed distally; apex rounded. Cyathia

solitary at the nodes; cyathial peduncles

0.2-2.5 mm long. Involucres campanulate,

0.8-1.4 mm long, 0.8-1.5 mm across; lobes

(rarely 4)5, broad-triangular, 0.3-0.4 mm

long, margin fimbriate; glands 4(rarely 5),

stipitate, patelliform, planar or shallowly

concave, transverse-oblong or reniform in

outline, 0.4-0.5 mm long, 0.6-1 mm wide,

pale green; gland appendages conspicuous,

spreading radially, oblong, 0.4-0.5 mm long,

0.8-1.3 mm wide, white, glabrous, margin ±

entire or shallowly lobed; bracteoles 0.6-0.8

mm long, adnate for c. 1/3 of their length to

involucre, free portion divided into subulate

hirsute segments. Staminate flowers 20-25

per cyathium; pedicels 0.5-1.3 mm long;

staminal filaments 0.1-0.2 mm long. Pistillate

flowers: styles 0.4-0.6 mm long, spreading,

smooth, glabrous or minutely pubescent, each

bifid for 1/3—1/2 of their length, the apices

terete. Capsules exserted from the involucre

on pedicel to 2.5 mm long, very broad-ovate

or depressed ovate in lateral view, 1.6-2.6 mm

long, 2.1-2.8 mm across, shallowly 3-lobate

with keels obtuse, smooth, glabrous or with

Austrobaileya 8(4): 441-600 (2012)

moderately dense indumentum; hairs evenly

distributed over capsule, spreading, 0.1-0.2

mm long; hypogynous disc entire. Seeds

ovate in outline, 1.7-1.8 mm long, 1-1.1 mm

tangentially, 1-1.1 mm radially, tetragonous

in cross section; dorsal faces convex; ventral

faces planar or slightly concave; all faces with

narrow rounded irregular or transverse ridges;

exotesta thin, of even thickness over surface,

grey-white, microreticulate, becoming

mucilaginous when moistened; endotesta pale

brown or red-brown. Fig. 27E.

Additional selected specimens examined : Queensland.

Port Curtis District: South Percy Island, 50 km NE

of Arthur Point, Shoalwater Bay, Oct 1989, Batianoff

11422 et al. (BRI); Mt Wheeler, Rockhampton, Jan 1989,

Specht 3 & Reeves (BRI, NSW); Gumigil Mining Lease,

18 km SSW of Marlborough, Jul 2000, Champion 540 &

Whereat (BRI); on Rockhampton - Marlborough road,

50 km from Rockhampton, May 1960, Johnson 1720

(BRI); 9.5 km W of Canoona, First Sugarloaf, Jan 1988,

Forster PIF3393 (BRI).

Distribution and habitat : Euphorbia

ophiolitica is restricted to central Qld,

from Percy Isles (east of Mackay) to the

Marlborough district (north of Rockhampton)

(Map 36). It grows on stony hillsides or

ridges, in open woodland or rarely grassland

communities, on shallow stony soils derived

from serpentinite rocks.

Phenology : Flowers and fruits have been

collected in January, May, July and October.

Notes : Euphorbia ophiolitica is similar to E.

laciniloba and E. papillifolia. For features

distinguishing E. ophiolitica from those

species, refer to the ‘Notes’ section under the

species concerned.

37. *Euphorbia ophthalmica Pers., Syn. PI.

2: 13 (1807); Chamaesyce ophthalmica (Pers.)

D.G.Burch, Ann. Missouri Bot. Gard. 53: 98

(1966). Type: [Brazil.] Rio de Janeiro, [July

1767,] [P] Commerson [238] (holo: P-JU n.v.

[IDC microfiche 6206. 1187, II. 2]).

Illustrations : Burger & Huft (1995: 19, fig. 6);

Harris (2001: 32, fig. 2).

Monoecious, annual to 8 cm high, few to many

stems arising from base. Stems prostrate,

decumbent or weakly ascending, sparingly

to much branched, smooth, moderately dense

to densely hairy; indumentum consisting

533

Halford & Harris, Euphorbia section Anisophyllum in Australia

of white weakly appressed crispate hairs

to 0.5 mm long interspersed with yellow

spreading ± straight segmented hairs to 1.5

mm long. Interpetiolar stipules subulate to

narrow-triangular, 0.5-2 mm long, bifid to

deeply bipartite, sparsely hairy with white

ascending hairs to 0.4 mm long; margin

laciniate. Leaves: petiole 0.5-2.5 mm long,

smooth, with indumentum as for stems; blade

oblong or narrow-ovate to ovate, 6-15 mm

long, 3-8 mm wide 1.5-2 times longer than

wide; adaxial surface green occasionally with

reddish tinge on margin, smooth, glabrous

or sparsely hairy with white, spreading to

ascending, ± straight hairs 0.6-0.9 mm long;

abaxial surface pale green, smooth, with a

moderately dense to dense indumentum of

white, spreading to ascending, ± straight hairs

to 1.2 mm long; base asymmetric with one side

rounded, the other cuneate; margin serrulate;

apex acute to obtuse. Cyathia in dense (10-30

cyathia) capitate, terminal cymose clusters to

10 mm in diameter on peduncles 1-7.5 mm

long; bracts subulate to narrowly triangular,

to 0.7 mm long; cyathial peduncles 0.5-1

mm long. Involucres turbinate, 0.6-0.8 mm

long, c. 0.5 mm across; lobes 5, subulate,

0.1-0.2 mm long, margin laciniate; glands

4, stipitate, cupuliform, with shallow central

pit and sometimes thickened rim, transverse-

elliptic to orbicular in outline, c. 0.1 mm long

c. 0.1 mm wide, pink or pale green; gland

appendages present and conspicuous or

absent, spreading radially, transverse-oblong,

to 0.3 mm long and 0.3 mm wide, white,

glabrous, margin irregularly lobed; bracteoles

filamentous, 0.5-0.6 mm long, glabrous,

entire or laciniate distally. Staminate flowers

2-5 per cyathium; pedicels 0.3-0.5 mm

long; staminal filaments c. 0.1 mm long.

Pistillate flowers: styles 0.3-0.5 mm long,

erect to spreading, smooth, glabrous, each

bifid for 1/2-2/3 of their length, the apices

clavate. Capsules exserted from involucre

on pedicel to 1.7 mm long, depressed ovate

in lateral view, 0.8-1.5 mm long, 1.1-1.3 mm

across, shallowly 3-lobate with keels acute,

smooth, with moderately dense indumentum;

hairs appressed, 0.1-0.2 mm long, evenly

distributed over capsule; hypogynous disc

entire. Seeds ovate in outline, 0.7-1 mm

long, 0.5-0.6 mm tangentially, 0.4-0.5 mm

radially, tetraquetrous in cross section; dorsal

and ventral faces planar to concave; all faces

with 5-7 faint narrow acute transverse ridges;

exotesta thin, of even thickness, pale brown,

microgranulate especially along ridges,

becoming mucilaginous when moistened;

endotesta pale red-brown. Fig. 27F.

Additional selected specimens examined: Queensland.

Burnett District: Walter Street, Kingaroy, Mar 2004,

Halford Q8170 & Edginton (BRI). Moreton District:

Lookout, Dulong road, c. 4.5 km W of Nambour,

Apr 2003, Halford Q7510 (BRI); Brisbane Botanic

Gardens, Mt Coot-tha, Mar 2000, Harris 123 (BRI);

between Woolley and Woodstock Streets, Taringa, 6

km from Brisbane GPO, Apr 2000, Bean 16179 (BRI,

MEL); Brisbane, Salisbury, Feb 2001, Batianoff210201

(BRI); 15 Whittaker Street, North Ipswich, Apr 1992,

Williams 92002 (BRI); 0.8 km along Sugarloaf Road,

Warrill View, SSW of Ipswich, Nov 2000, Bean 17023

(BRI); Newmarket end, Ashgrove Avenue, Newmarket,

Brisbane, Mar 2000, Forster PIF25475 (AD, BRI);

Ormeau Railway Station, between Beenleigh & Nerang,

May 2003, Bean 20384 (BRI); Guanaba, Coomera

River, 5 km SW of Oxenford, property I. Cairns, Apr

2003, Forster PIF29300 (BRI); 8 km S of Canungra

on road to O’Reillys, near winery. Mar 2003, Fechner

s.n. & Holland (BRI [AQ733643]). New South Wales.

Far North Coast Council depot, Wyrallah Road, East

Lismore, Nov 2000, Hosking 1965 & Scott (BRI).

Distribution and habitat : Euphorbia

ophthalmica is a native of southern Florida,

the West Indies and Argentina and is

naturalised in Australia. In Australia it occurs

from Gladstone, Qld south to Lismore, NSW

(Map 37). It grows along roadsides, in garden

beds or lawns.

Phenology : Flowers and fruits have been

collected from November to May.

Notes: Euphorbia ophthlamica is easily

distinguished from the native Australian

Euphorbia species by its indumentum of

long yellowish coloured hairs on stems and

petioles. Euphorbia ophthalmica is closely

related to E. hirta , but may be distinguished by

its slender, prostrate habit, generally shorter

leaf blade (blade to 1.5 cm long compared

with blade to 4.5 cm long for E. hirta), and

terminal cymose glomerules compare with

axillary and terminal for E. hirta.

534

38. Euphorbia pallens Dillwyn, Rev. Hortus

Malab. 54 (1839); Chamaesyce pallens

(Dillwyn) V.S.Raju, J. Econ. Taxon. Bot.

28: 92 (2004). Type: [India], illustration of

Ben-Pala, Rheede, Hort. Malab. 10: 115, t. 58

(1690).

Illustrations : Lin et al. (1991: 224, fig 4), as

Chamaesyce atoto ; Forster (1994: 9, fig 3-5)

as Euphorbia atoto.

Monoecious, herbaceous perennial to 50(80)

cm high, few to many stems arising from

thickened woody rootstock. Stems ascending

to erect or prostrate, much branched,

smooth, often with whitish bloom glabrous.

Interpetiolar stipules triangular, 1-1.5 mm

long, abaxial surface glabrous or hairy with

white ascending hairs c. 0.1 mm long, adaxial

surface hairy with white ascending hairs c.

0.1 mm long; margin lacerate or fimbriate.

Leaves: petiole 1.5-2.7 mm long, smooth;

blade oblong, ovate or elliptic, 24-35 mm

long, 11-22 mm wide, 1.4-2.7 times longer

than wide; adaxial surface blue-green or pale

green with whitish bloom, smooth or minutely

papillose, glabrous abaxial surface pale blue-

green sometimes with reddish tinge, smooth

glabrous, base asymmetric with one side

cordate, the other obtuse or shallowly cordate;

margin entire; apex obtuse to rounded or

retuse, sometimes with apiculate tip. Cyathia

in congested 2-5 branched dichasial cymes

together with a solitary cyathium at the distal

nodes; peduncles 5-20 mm long; bracts leaf¬

like but smaller than the primary stem leaves;

cyathial peduncles 2-5 mm long. Involucres

turbinate or cupuliform, 1.4-1.5 mm long,

1.5- 1.8 mm across; lobes 5, narrow-triangular

to broad-triangular, 0.5-0.6 mm long,

margin entire, fimbriate or laciniate; glands

4, stipitate, patelliform, planar or shallowly

concave, transverse-elliptic or orbicular in

outline, 0.3-0.4 mm long, 0.5-0.6 mm wide,

pale green; gland appendages present but

inconspicuous, spreading radially, transverse-

linear, to 0.1 mm long, 0.5-0.6 mm wide, pink

or white, glabrous, margin entire; bracteoles

1.5- 1.9 mm long, adnate for 1/3—1/2 of their

length to involucre, free portion divided into

numerous linear glabrous or hirsute segments.

Staminate flowers 5-20 per cyathium;

pedicels 1-2 mm long; staminal filaments

Austrobaileya 8(4): 441-600 (2012)

0.5-0.7 mm long. Pistillate flowers: styles

0.4-0.6 mm long, ascending, spreading

distally, smooth, glabrous, each bifid for 1/3—

1/2 of their length, the apices dorsiventrally

flattened. Capsules exserted from involucre

on pedicel to 6 mm long, very broad-ovate or

transversely broad-elliptic in lateral view, 2.3-

2.8 mm long, 2.8-3.8 mm across, shallowly

or deeply 3-lobate with keels obtuse, smooth,

glabrous; hypogynous disc entire. Seeds very

broad-elliptic or very broad-ovate in outline,

1.5-1.7 mm long, 1.2-1.5 mm tangentially,

1.3-1.4 mm radially, trigonal or suborbicular

in cross section; dorsal and ventral faces

convex, smooth; exotesta thin, of even

thickness over surface, white or pale brown,

microreticulate; not becoming mucilaginous

when moistened; endotesta brown. Fig. 27G.

Additional selected specimens examined : Queensland.

Cook District: Kerr Island, Torres Strait, Jan 2007,

Waterhouse BMW7498 (BRI); Deliverance Island, Apr

1996, Waterhouse BMW3797 (BRI); Ida Point, c. 4 km

SE of Cape York, Oct 1965, Smith 12524 (BRI); Muttee

Head, Cape York, Mar 1990, Forster PIF6433 (BRI);

Muddy Bay, Cape York, Jun 1994, Forster PIF15316

& Tucker (BRI); Saunders Island, Nov 1973, Stoddart

5073 (BRI); Bathurst Bay, Aug 1974, Hyland 7403

(BRI); Howick Island, Oct 1973, Stoddart 4857 (BRI);

Coquet Island, Howick Group, Mar 1984, Godwin s.n.

(BRI [AQ440547]); Sinclair Island, Aug 1973, Stoddart

4190 (BRI, MEL); Eagle Island, Great Barrier Reef, Jan

1973, Heatwole s.n. (BRI [AQ202465]); Ingram Island,

Jul 1973, Stoddart 4045 (BRI, NSW); Lizard Island,

Sep 1967, Heatwole 68 (BRI); Eagle Island, Oct 1973,

Stoddart 4821 (BRI); Three Isles, Sep 1973, Stoddart

4470 (BRI); 2 km N of settlement at Bramston Beach,

12 km E of Babinda, Apr 1975, McDonald 1490 &

BatianoffiBRl); Mission Beach, Dec 1949, Clemens s.n.

(BRI [AQ202454]). North Kennedy District: Edmund

Kennedy N.P., near Cardwell, Dec 1991, Bean 3874

(BRI); Hinchinbrook Island, W of Kirkville Hills, Aug

1970, Everist 9657 (BRI); George Point, Hinchinbrook

Island, Apr 1994, Camming 12773 (BRI).

Distribution and habitat : Euphorbia

pallens ranges from Sri Lanka through

Malaysia, southern China to Australia and

the southwestern Pacific Islands. In Australia,

it occurs in coastal areas from Torres Strait

south along the east coast of Cape York

Peninsula to Hinchinbrook Island (Map 38).

It grows on coastal sands on frontal beach

dunes and coral cays.

Phenology : Flowers and fruits have been

collected throughout the year.

535

Halford & Harris, Euphorbia section Anisophyllum in Australia

Notes : The name Euphorbia atoto G.Forst.

(Chamaesyce atoto (G.Forst.) Croizat) has

been previously applied to this species in

Australia (Forster 1994; Forster & Halford

2010), as well as in south east Asia and the

Pacific (Hurusawa 1954; Smith 1981; Lin et

al. 1991). Smith (1981) selected a lectotype

for the name E. atoto from material at Kew.

Florence (1996) set forth an opposing view on

the lectotypification of E. atoto on the grounds

that the specimen selected was in serious

conflict with the protologue and that there

was another element available that was part

of the original material and morphologically

agreed with the description in the protologue

(ICBN Art 9.17). Esser & Chayamarit (2001)

have put forward the argument that “it is not

obvious whether the discrepancy discussed

by Florence in the inflorescence architecture

justifies this decision”. We have examined

microfiche, photographs and digital images

of the relevant material collected by the

Forsters which resides in a number of herbaria

(P-Forst, BM, G, GEOT, UPS-THUNB) and

we concur with Florence’s conclusions. As

circumscribed by Florence (1996) E. atoto is

endemic to Tahiti. Therefore, another name is

required for the widespread seashore species.

Euphorbia pallens Dillwyn seems to be the

next available name.

Euphorbia pallens is smiliar to E.

litticola, E. obliqua and E. psammogeton.

For features distinguishing E. pallens from

E. litticola , refer to the ‘Notes’ section under

that relevant species. Euphorbia pallens can

be distinguished from E. obliqua by having

its cyathia arranged in terminal or axillary lax

to congested dichasial cymes (compared with

cyathia solitary in upper axils for E. obliqua)

and generally larger leaves (24-35 x 11-22

mm as compared with 9-20 x 6-10 mm for

E. obliqua). It differs from E. psammogeton

by its smooth rather than rugulose seeds, and

smaller gland appendages (up to 0.1 mm long

as compared with 0.2-0.4 mm long for E.

obliqua).

Hassall (1977) recorded a chromosome

number n = 8 for two voucher collections that

were identified as E. atoto at the time, (RLS

[R.L.Specht] 7478, Lizard Island; and GLW

[G.L.Webster] 73118, Currimine [Kurrimine]

Beach, both from north Queensland).

Unfortunately, we have been unable to locate

the vouchers that were apparently lodged

in BRI and cannot confirm that they are E.

pallens.

39. Euphorbia papillata Halford &

W.K.Harris, species nova arete similis

E. ferdinandi Baill. sed habitu robustiore

seminibus latioribus 0.7-1 mm latis (ad vicem

0.5-0.7 mm latis), glandulae appendicibus

majoribus 0.1-0.2 mm long is (ad vicem

appendicibus aut absentibus aut usque 0.1

mm longis), floribus staminatis 5-10 in

quoque cyathio (in vicem floribus paucioribus

3- 5 in quoque cyathio) differt. Euphorbia

papillata quondam saepe putabatur eadem

E. drummondii Boiss. esse sed seminum

superficiebus ± laevibus (ad vicem

superficiebus 3-6 porcis transversis distinctis

praeditis),foliorumfructuumque superficiebus

papillosis (ad vicem superficiebus laevibus),

glandulae appendicibus 0.1-0.2 mm longis

(ad vicem appendicibus usque 0.1 mm longis)

distinguenda. Typus: Northern Territory.

Georgina Station, 8 km NW of No. 14 Bore,

22 September 1992, P.K.Latz 12754 (holo:

BRI; iso: MEL, NT).

Monoecious, herbaceous perennial to 25 cm

high, many stems arising from woody taproot

crown, the whole plant glabrous. Stems

prostrate or ascending to erect, much branched,

papillose or smooth, often longitudinally

ridged. Interpetiolar stipules subulate or

narrow-triangular, 0.8-1.6 mm long, bipartite,

glabrous; margin laciniate. Leaves: petiole

0.3-0.7 mm long, smooth or papillose; blade

obovate, oblong-obovate or narrow-oblong,

4- 11 mm long, 2-5.5 mm wide, 1.6-3.1 times

longer than wide; adaxial surface dark green

or blue-green, papillose; abaxial surface

blue-green but paler than adaxial surface,

papillose; base asymmetric with one side

cordate, the other cuneate or obtuse; margin

serrulate distally; apex rounded or mucronate.

Cyathia solitary at the nodes, often gathered

together on short leafy lateral branchlets with

subtending leaves usually slightly smaller

than primary stem leaves; peduncles 0.3-0.6

mm long, papillose. Involucres campanulate

or turbinate, 0.7-1.1 mm long, 0.8-1.4 mm

536

across; lobes 5, triangular, 0.3-0.5 mm long,

margin entire or fimbriate; glands 4, shortly

stipitate, cupuliform, with distinct central

pit and thickened rim, transverse-oblong

or transverse-elliptic in outline, 0.15-0.3

mm long, 0.25-0.4 mm wide, pink to red or

pale green; gland appendages conspicuous,

spreading radially, transverse-oblong,

transverse-linear or reniform, 0.1-0.2 mm

long, 0.5-07 mm wide, pink or white,

glabrous, margin entire or shallowly lobed;

bracteoles obovate, c. 1 mm long, divided

into numerous subulate hirsute segments.

Staminte flowers 5-10 per cyathium;

pedicels 0.9-1.1 mm long; staminal filaments

c. 0.1 mm long. Pistillate flowers: styles 0.1-

0.4 mm long, spreading, ascending to erect,

smooth or papillose, glabrous, each entire or

bifid for c. 1/2 of their length, the apices stout

terete. Capsules exserted from involucre

on pedicel to 2 mm long; broad-elliptic in

lateral view, 1.9-2.3 mm long, 1.6-2.4 mm

across, shallowly 3-lobate with keels obtuse

or acute, papillose, glabrous; hypogynous

disc entire. Seeds ovate in outline, 1.3-1.6

mm long, 0.7-1 mm tangentially, 0.6-1 mm

radially, tetraquetrous in cross section; dorsal

faces ± planar, smooth or with faint narrow

medial longitudinal ridge; ventral faces

planar or concave, smooth; exotesta thin,

of even thickness over surface, grey-white,

microreticulate or microgranulate, becoming

mucilaginous when moistened; endotesta pale

brown to red brown.

Austrobaileya 8(4): 441-600 (2012)

Distribution and habitat : Euphorbia

papillata is widespread in southern NT and

western Qld, extending into northern SA

and north-western NSW, but absent from the

Simpson, Sturt Stony and Strzelecki Deserts.

Etymology : The specific epithet is from Latin

papillatus , having papillae, in reference to

the papillate surface of the capsules, leaves

and sometimes the young branchlets of this

species.

Notes : Euphorbia papillata closely resembles

E.ferdinandi but differs from that by its more

robust habit, broader seeds (0.7-1 mm versus

0.5-0.7 mm for E. ferdinandi ), larger gland

appendages (0.1-0.2 mm long versus gland

appendages absent or up to 0.1 mm long for E.

ferdinandi ), and more staminate flowers per

cyathium (5-10 versus 3-5 for E. ferdinandi).

Euphorbia papillata was previously

often identified as E. drummondii but it

is distinguished from that species by its ±

smooth seed surfaces (versus seed surfaces

with 3-6 distinct transverse ridges for E.

drummondii ), papillose surfaces on leaves

and fruits (versus smooth for E. drummondii ),

and larger gland appendages (0.1-0.2 mm

long versus gland appendages up to 0.1 mm

long for E. drummondii ).

As accepted here, E. papillata is very

variable in leaf size and stem texture. Two

varieties are recognised which can be

distinguished by the following key.

Key to varieties of Euphorbia papillata

Stems papillose.39a. E. papillata var. papillata

Stems ± smooth, lacking papillae.39b. E. papillata var. laevicaulis

39a. Euphorbia papillata var. papillata

Chamaesyce sp. (Pathungra A.Gunness

AG2118); Forster & Halford (2010: 65).

Stems papillose. Leaves: petiole 0.3-0.7 mm

long, papillose; blade obovate or oblong-

obovate, 4-10 mm long, 2-5.5 mm wide,

1.6-2.3 times longer than wide. Involucres

campanulate or turbinate, 0.9-1.1 mm long,

1—1.4 mm across; lobes 0.4-0.5 mm long,

margin fimbriate, n— 41. Figs 15, 27H.

Additional selected specimens examined: Northern

Territory. Phillip Creek Station, May 1993, Egan 2287

(BRI); Alexandria Station, 5 km NE [of] Buchanan

Creek, Nov 1986, Henshall 4121 (DNA, NT); 35.5 miles

[c. 57 km] NNW [of] Wauchope township, Aug 1956,

Lazarides 5849 (BRI, DNA); Wakaya Desert, May

1993, Latz 13086 (DNA, MEL); 40 km N [of] Plenty

Highway on Sandover Highway, Apr 1988, Kimbel 68

(DNA, NT); 12 miles [c. 19 km] W of Stuart Highway,

along Yuendumu Road, Apr 1967, Maconochie 62 (AD,

DNA, MEL); Illamurta Springs area; 7.5 km west, Aug

1988, Barritt 323 (DNA); Uluru (Ayers Rock - Mt

Olga) N.P.; on the Docker River road, 53.6 km WNW

Halford & Harris, Euphorbia section Anisophyllum in Australia

537

Fig. 15. Euphorbia papillata var. papillata. A. habit *0.6. B. branchlet with cyathia *4. C. leaf *8. D. indumentum

on upper leaf surface x32. E. stipules x24. F. cyathia with female flower x24. G. capsule with cyathia xl6. H. cyathial

gland with appendage, adaxial view x32.1. capsule, top view *16. J. capsule, lateral view xl6. A, C & D from Latz

12754 (NT); B, E-J from Halford Q8597 & Thomas (BRI). Del. W. Smith.

538

of the Ranger Station, Aug 1988, Lazarides & Palmer

539 (MEL). Queensland. Burke District: 1 km W of

Torrens Creek (township) along Flinders Highway,

towards Hughenden, Apr 2006, Halford Q9020 &

Batianoff (BRI). Gregory North District: 31 km from

Winton, Bladensburg N.P., Sep 2005, Halford 08597 &

Thomas (BRI); 60 km S of Winton on road to Opalton,

Sep 2005, Halford Q8600 & Thomas (BRI); Jundah -

Winton road, 38 km S of Mayne River crossing. May

2004, Bean 22495 (BRI). Warrego District: c. 13 km

E of Quilpie, near junction of Adavale Road and Quilpie

- Cheepie Road, Jun 2002, Pollock ABP1301 & Walsh

(BRI). Gregory South District: 81 km by road NW of

Thylungra and 197 km NW of Quilpie on the Windorah

Road, Mar 2001, Thomas 1948 & Fechner (BRI). South

Australia. Everard Ranges, Wildcat Bore, Sep 1963,

Eichler 17467 (AD); near Marla, Oct 1998, Bates 51375

(AD); c. 25 km W of Tallaringa Well, May 1967, Lothian

3841 (AD). New South Wales. 4 km W of Weebah Gate,

NSW - Qld border, Nov 1976, Pickard 3305 (NSW).

Distribution and habitat: Euphorbiapapillata

var. papillata is widespread in southern NT

and western Qld, extending into northern SA

and north-western NSW, but absent from the

Simpson, Sturt Stony and Strzelecki Deserts

(Map 39a). It grows in mulga woodland,

eucalypt open woodland, open shrubland

or tussock grassland communities on red

sandy to loam soils on plains, low lateritic

or limestone rises; also rarely recorded on

black clay soils in Mitchell grassland or open

woodland communities.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from March to June.

39b. Euphorbia papillata var. laevicaulis

Halford & W.K.Harris, varietas nova

ab E. papillata var. papillata caulibus ±

laevibus papillis carentibus differt. Typus:

Queensland. Mitchell District: Moorrinya

National Park, 7 May 2006, E.J.Thompson &

G.W. Wilson TAN551 (holo: BRI).

Stems smooth. Leaves: petiole 0.5-0.7 mm

long, smooth; blade narrow-oblong, 6-11 mm

long, 2-3.6 mm wide, 2.2-3.1 times longer

than wide. Involucres turbinate, 0.7-0.8 mm

long, c. 0.8 mm across; lobes 0.3-0.4 mm

long, margin entire or fimbriate. Fig. 271.

Additional selected specimens examined: Northern

Territory. 29 km S [of] Renner Springs, Jun 1977,

Parker 942 (BRI). Queensland. Burke District: 26 km

N of Burke and Wills Roadhouse, Mar 2003, McDonald

KRM1332 (BRI); Glengalla, 63 miles [c. 101 km] N of

Maxwelton, Jun 1947, Everist 3035 (BRI); 1 km W of

Austrobaileya 8(4): 441-600 (2012)

Torrens Creek (township) along Flinders Highway,

towards Hughenden, Apr 2006, Halford Q9021 &

Batianoff (BRI). North Kennedy District: Flinders

Highway, 36 km E of Torrens Creek, Jul 2000, Bean

16764 (BRI); 20 km from Charters Towers, towards

Clermont, Apr 2002, Bean 18961 (BRI, MEL). South

Kennedy District: 6 km N of Ulcanban homestead. Mar

2002, Turpin GPT740 & Thompson (BRI). Mitchell

District: 72 km S of Prairie on stock route, 1 km E of

Holmleigh homestead. Mar 2004, Thompson TAN153 &

Camming (BRI); 32 km SSE of Prairie on Stock Route

503 bordering Ashton Station, Mar 2004, Thompson

TAN132 & Camming (BRI); Corinda Station, 10 km NE

of Corinda homestead. May 2006, Thompson TAN264 &

Wilson (BRI); Corinda Station, 16.1 km N of Corinda

homestead. May 2006, Thompson TAN333 & Wilson

(BRI); MoorinyaN.P., c. 100 km S of Torrens Creek, Apr

2005, Booth 3707 & Thompson (BRI); Corinda, 100 km

N of Aramac, Mar 2004, Camming 22475 & Thompson

(BRI); 50 km from Longreach, towards Winton, May

2004, Bean 22594 (BRI).

Distribution and habitat: Euphorbia

papillata var. laevicaulis extends from

Cloncurry eastward to Charters Towers and

south to near Longreach, Qld; with a disjunct

occurrence near Renner Springs, NT (Map

39b). It is recorded growing in a variety

of habitats: tussock/hummock grassland,

Eremophila mitchellii Benth. low open

woodland, mulga woodland and eucalypt

woodland/open forest, on mostly undulating

plains. The soils vary from sands to clays.

Phenology: Flowers and fruits have been

collected from January, March to July and

October.

Notes: Euphorbia papillata var. laevicaulis

differs from E. papillata var. papillata by its

± smooth stems that lack papillae.

Etymology: The varietal epithet is from Latin

laevis , smooth, and caulis , stem, in reference

to the more or less smooth surface of the

stems of this variety.

40. Euphorbia papillifolia Halford &

W.K.Harris, species nova quoad formam

sculturamque seminum superficierum

similis E. ophioliticae (P.I.Forst.) Y.Yang

differt autem foliis oblongis ovatisve 1.5-2.8

plo longioribus quam latioribus (ad vicem

foliis ellipticis usque late ellipticus 1-1.5

plo longioribus quam latioribus) glandis

minoribus 0.1-0.4 x 0.3-0.8 mm (ad vicem

glandulis 0.4-0.5 x 0.6-1 mm) seminibus

minoribus 1.3-1.6 x 0.7-1 x 0.8-1 mm (ad

539

Halford & Harris, Euphorbia section Anisophyllum in Australia

vicem seminibus 17-1.8 x 1-1.1 H 1—1.1 mm)

plerumque stipulis longioribus 0.5-1.6 mm

longis (ad vicem stipulis 0.2-0.6 mm longis).

Euphorbia papillifolia crescit in solis argillis

repantis saepe a basalto orituris quoniam E.

ophiolitica in solis a saxis serpentinis orituris.

Typus: Queensland. Port Curtis District:

South Kariboe Creek crossing, Burnett

Highway, between Biloela and Monto, 19

March 2010, D.HalfordQ9771 & G.P.Guymer

(holo: BRI, iso: MEL, NSW, distribuendi ).

Monoecious or dioecious, herbaceous

perennial to 10 cm high, few to many stems

arising from woody rootstock. Stems

prostrate or decumbent, sparingly to much

branched, often longitudinally ridged,

glabrous (rarely with a moderately dense

indumentum, Allison TH5034 [BRI]); hairs

spreading, straight, 0.1-0.4 mm long, white.

Interpetiolar stipules subulate or narrow-

triangular, 0.5-1.6 mm long, bifid or deeply

bipartite, glabrous; margin entire, laciniate or

sometimes fimbriate. Leaves: petiole 0.6-1.1

mm long, smooth, glabrous; blade oblong or

ovate, 7-11 mm long, 2.5-9 mm wide, 1.8-

2.8 times longer than wide; adaxial surface

blue-green sometimes with reddish tinge

along margin; minutely papillose (visible at

40 x mag.), glabrous (rarely with a moderately

dense indumentum consisting of spreading,

straight hairs to 0.4 mm long, abaxial surface

pale grey to white; minutely papillose (visible

at 40 x mag.), glabrous or with indumentum

as of adaxial surface; base asymmetric

with one side cordate, the other cuneate

or obtuse to rounded; margin serrulate or

sparingly minutely toothed sometimes only

distally (rarely entire); apex acute to obtuse

or rounded. Cyathia solitary at the nodes,

often gathered together on short leafy lateral

branches; peduncles 0.6-1.3(2.3) mm long,

smooth, glabrous. Involucres turbinate or

cupuliform, 0.6-1.5 mm long, 0.6-1.8 mm

across; lobes 5 or 6, triangular, 0.2-0.4 mm

long, margin laciniate or fimbriate; glands

4-6, stipitate, patelliform, planar or concave,

transverse-oblong or reniform in outline,

0.1-0.4 mm long, 0.3-0.8 mm wide, pink,

red or yellowish green; gland appendages

conspicuous, spreading radially, transverse-

oblong, broad-obovate to very broad-obovate,

obdeltoid or reniform, 0.1-0.9 mm long, 0.9-

1.4 mm wide, pink or white, glabrous, margin

entire, erose or shallowly lobed; bracteoles

0.5-0.6 mm long, adnate for c. 1/2 of their

length to involucre, free portion divided

into few linear hirsute segments. Staminate

flowers 1-30 per cyathium; pedicels 0.8-1.3

mm long; staminal filaments 0.1-0.3 mm long.

Pistillate flowers: styles 0.3-0.8 mm long,

connate at the base into a column for c. 1/6

of their length, spreading, minutely papillose

or smooth, pubescent or glabrous, each bifid

for 1/3—1/2 of their length, the apices terete.

Capsules exserted from involucre on pedicel

to 3.5 mm long, broad-elliptic or transversely

broad-elliptic in lateral view, 1.5-2.1 mm

long, 1.7-2.2 mm across, shallowly 3-lobate

with keels obtuse, smooth, glabrous;

hypogynous disc entire. Seeds ovate to broad-

ovate in outline, 1.3-1.6 mm long, 0.7-1 mm

tangentially, 0.8-1 mm radially, tetraquetrous

or tetragonous in cross section; dorsal faces

convex; ventral faces planar to concave; all

faces smooth, faintly undulate or with faint

to distinct narrow rounded irregular ridges;

exotesta thin, of even thickness over surface,

grey-white, microreticulate, becoming

mucilaginous when moistened; endotesta pale

brown.

Distribution and habitat : Euphorbia

papillifolia is widespread in eastern Australia

from near Mount Surprise, north-eastern Qld,

south to the Bathurst district, NSW.

Etymology : The specific epithet is from

Latin papilliatus , having papillae, and - folius ,

leaved, in reference to the papillose leaf

surface of this species.

Notes : Euphorbia papillifolia is similar to E.

ophiolitica in seed shape and seed surface

texture. It differs in having oblong or ovate

leaves 1.5-2.8 times longer than wide (versus

elliptic to broadly elliptic, 1-1.5 times longer

than wide for E. ophiolitica), smaller glands

(0.1-0.4 x 0.3-0.8 mm versus 0.4-0.5 x

0.6-1 mm), smaller seed dimensions (1.3-1.6

x 0.7-1 x 0.8-1 mm versus 1.7-1.8 x 1-1.1

x 1 - 1.1 mm) and generally longer stipules

(0.5-1.6 mm long versus 0.2-0.6 mm long).

They also differ in habitat with E. papillifolia

occurring on cracking clay soils often derived

540

Austrobaileya 8(4): 441-600 (2012)

from basalt substrate while E. ophiolitica is

restricted to soils derived from serpentinite

rocks.

As accepted here, E. papillifolia varies

in breeding system, the number of gland per

Key to varieties of Euphorbia papillifolia

Plants monoecious; gland appendages 0.1-0.4 mm long; glands 4 per

cyathia; staminate flowers per cyathia <15; styles 0.3-0.5 mm

long.40a. E. papillifolia var. papillifolia

Plants dioecious (rarely monoecious); gland appendages 0.4-0.9 mm long;

glands 4-6 per cyathia; staminate flowers per cyathia >15, styles

0.5-0.8 mm long.40b. E. papillifolia var. polyandra

cyathia and length of gland appendages. This

variation is considered sufficient to warrant

formal recognition of two varieties within this

species which can be distinguished using the

following key.

40a. Euphorbia papillifolia var. papillifolia

Monoecious, herbaceous perennials.

Stems prostrate, glabrous or rarely with a

moderately dense indumentum. Involucres

turbinate, 0.6-1 mm long, 0.6-1.1 mm across;

lobes 5, triangular, 0.2-0.3 mm long; glands

4, 0.1-0.3 mm long, 0.3-0.5 mm wide; gland

appendages 0.1-0.4 mm long, 0.3-0.9 mm

wide. Staminate flowers 1-10 per cyathium.

Female flowers with styles 0.3-0.5 mm long.

Figs 16, 27J.

Additional selected specimens examined : Queensland.

Cook District: Undara N.P., NW of Tobacco Spring,

Dec 2006, McDonaldKRM6036 (BRI). Burke District:

c. 92 km N of Hughenden, Apr 1998, Thompson

HUG531 et al. (BRI). North Kennedy District: Mt

Fox, Dec 1949, Clemens s.n. (BRI [AQ202423]); several

kms SE of turn off to Ingham, Jun 1999, Addicott 113

(BRI); Burdekin River, 40 km N of Charters Towers, c.

1 km S of Keelbottom Mountain, Jul 1981, Sharpe 2894

(BRI); N of Bowen, May 1997, Calvert & Lockyers 1

(BRI); Millview, 15 km N of Charters Towers, Jul

2006, Hooker NH675 (BRI). Leichhardt District: 29

km S of Springsure, Mar 1995, Fensham 2607 (BRI);

12 km SE of Rolleston, Jan 1996, Fensham 2422 (BRI).

Port Curtis District: Surfus Hill, E of Marlborough

Creek, Nov 1997, McCabe C2 (BRI); Glen Geddes,

near Rockhampton, Jun 1965, Beauglehole ACB3567

(MEL, NSW); South Kariboe Creek, SE of Biloela,

Jan 1996, Bean 9637 (BRI). Burnett District: c. 8 km

NW of ‘Rawbelle’, W of Monto, Jun 1996, Bean 10379

(BRI). Warrego District: junction of Landsborough &

Mitchell Highways, 5.5 km S of Augathella, Oct 2011,

Halford QM625 (BRI). Maranoa District: The Lamen,

ESE of Roma, Jan 1998, Bean 12989 (BRI).

Distribution and habitat : Euphorbia

papillifolia var. papillifolia is widespread in

central east Qld from Undara N.R, east to Mt

Fox (west of Ingham) and south to Roma and

Monto (Map 40a). It inhabits Dichanthium

spp. grassland or eucalypt/Acacza woodland

on plains or gently undulating country. The

soils are dark brown to black cracking clays

often derived from basalt substrates.

Phenology : Flowers and fruits have been

collected throughout the year.

40b. Euphorbia papillifolia var. polyandra

Halford & W.K.Harris, varietas nova

a varietate typica plerumque glandulae

appendicibus majoribus 0.4-0.9 * 0.9-1.4

mm (ad vicem appendicibus 0.1-0.4 x 0.3-0.9

mm), staminibus pluribus in quoque cyathio

(15-30 non 1-7), stylis longioribus 0.5-0.7 mm

longis (ad vicem stylis 0.3-0.5 mm longis),

interdum seminibus majoribus 1.3-1.6 x 0.9-

1 x 0.9-1 mm (ad vicem seminibus 1.3-1.4 x

0.7-0.9 x 0.8-0.9 mm) differt. Typus: New

South Wales, eastern bank of Lake Inverell,

c. 4 km SE of Inverell, 11 December 2002,

L.M.Copeland 3483 (holo: BRI, according to

label information on the holotype sheet there

are duplicates (isotypes) lodged at CANB, L,

MEL, NE, NSW-all n.v).

Chamaesyce sp. A.; James & Harden (1990:

428).

Dioecious (rarely monoecious), herbaceous

perennials. Stems prostrate or decumbent,

glabrous. Involucres turbinate or cupuliform,

0.9-1.5 mm long, 0.8-1.8 mm across; lobes

5-6, triangular, 0.3-0.4 mm long; glands

4-6, 0.2-0.3 mm long, 0.5-0.8 mm wide;

Halford & Harris, Euphorbia section Anisophyllum in Australia

541

Fig. 16. Euphorbia papillifolia var. papillifolia. A. habit x0.5. B. branchlet with cyathia *6. C. leaf x6. D. papillae

on lower leaf surface x32. E. stipules x24. F. cyathia with female flower x24. G. cyathia with capsule xl2. H. cyathial

gland with appendage, adaxial view x32.1. capsule, top view xl2. J. capsule, lateral view xl2. All from Halford Q9771

& Guymer (BRI). Del. W.Smith.

542

gland appendages (0.1)0.4-0.9 mm long,

0.9-1.4 mm wide. Staminate flowers 15-30

per cyathium. Pistillate flowers with styles

0.5-0.7 mm long. n= 11.

Additional selected specimens examined : Queensland.

Leichhardt District: Warren State Farm, Mar 1920,

Francis s.n. (BRI [AQ202944]); Barfield, Jan 1945,

Scarlett S228 (BRI); 1.2 km along Lyndley Lane, N

of Jimbour, Nov 1997, Bean 12531 (BRI). New South

Wales. Mt Russell, near Inverell, Dec 1915, Breakwell

s.n. (NSW 614327); 9.3 km NE of Inverell onNullamanna

Road, May 1985, Wilson 6178 (NSW); Inverell, Apr

1913, Boorman s.n. (NSW 612144); Sinclair Lookout,

Waterloo Range, 14.4 km W of Glen Innes, Mar 1987,

Coveny 12489 et al. (NSW); 40 km ENE of Narrabri,

Sep 1976, Hassall 7669 (BRI); entrance to Mt Kaputar

N.P. on Dawsons Spring Road, 28 km ENE of Narrabri,

Nov 1976, Coveny 8893 & Roy (NSW); Binnaway

Road, 5 miles [ c . 8 km] SSE of Coonabarabran, Jan

1962, Salasoo 2313 (NSW); east facing slope to north of

Dulegal Arboretum, Chaffey Dam, Feb 1993, Hosking

688 (MEL); Kelso, Mar 1933, McKie s.n. (NSW 614324).

Distribution and habitat: Euphorbia

papillifolia var. polyandra occurs from

Springsure, Qld, south to the Bathurst district,

NSW (Map 40b). Itgrows in grassy Eucalyptus

woodland or ThemedalDichanthium grassland

on plains or undulating to hilly terrain. The

soils are mostly red clay loams to black clays,

rarely sandy. It is also recorded as a weed of

cultivation.

Phenology : Flowers and fruits have been

collected throughout the year.

Notes: Euphorbia papillifolia var. polyandra

differs from the type variety in having

generally larger gland appendages (0.4-0.9

x 0.9-1.4 mm versus 0.1-0.4 x 0.3-0.9 mm),

more stamens per cyathia (15-30 versus 1-7),

longer styles (0.5-0.7 mm long versus 0.3-0.5

mm long) and generally larger seeds (1.3-1.6

x 0.9-1 x 0.9-1 mm versus 1.3-1.4 x 0.7-0.9

x 0.8-0.9 mm).

Bentham (1873) cited a Dr Beckler

collection from Warwick, Qld (MEL 68198)

under E. alsiniflora with the comment that it

appears to be the same species. The Beckler

collection is here identified as E. papillifolia

var. polyandra.

Hassall (1977) proposed the name E.

drummondii subsp. bleckeri for this taxon but

this name was never validly published.

Austrobaileya 8(4): 441-600 (2012)

Etymology: The varietal epithet is from Greek

polys , many, and andrus , man (stamen), in

reference to the many more stamens present

per cyathia in this variety compared with the

type variety.

41. Euphorbia petala Ewart & L.R.Kerr,

Proc. Roy. Soc. Victoria 39: 1, fig. 1 (1926);

Chamaesyce petala (Ewart & L.R.Kerr)

P.I.Forst. & R.J.F.Hend., Novon 5: 323 (1995).

Type: [Northern Territory.] Wycliffe Well,

June 1924, [A.J. Ewart s.n] (lecto [here

designated]: MEL 1560387; isolecto: MEL

503409).

Illustration: Ewart & Kerr (1926: 2, fig 1).

Monoecious, herbaceous perennial, few to

many stems arising from woody taproot, the

whole plant glabrous. Stems prostrate, much

branched, smooth. Interpetiolar stipules

narrow-triangular to subulate, 0.6-1.5 mm

long, bipartite, glabrous; margin laciniate.

Leaves: petiole 0.5-0.8 mm long, smooth;

blade oblong, obovate or elliptic, 6-11 mm long,

3-7 mm wide, 1.5-2.8 times longer than wide;

adaxial surface pale blue-green or dull light

green, smooth; abaxial surface similar to but

paler than adaxial surface; base asymmetric

with one side auriculate to cordate, the

other shallowly cordate to rounded; margin

serrulate or sparingly minutely toothed; apex

rounded to shallowly retuse. Cyathia solitary

at the nodes, often gathered together on short

leafy lateral branchlets with subtending

leaves usually slightly smaller than primary

stem leaves; peduncles 0.2-0.8(1.2) mm long,

smooth. Involucres campanulate or turbinate,

1.1-1.5 mm long, 1-1.2 mm across; lobes 5,

triangular, 0.3-0.5 mm long, margin entire

or fimbriate; glands 4, stipitate, patelliform,

concave, or cupuliform with central tangential

pit, transverse-oblong in outline, 0.1-0.5 mm

long, 0.4-0.5 mm wide, pink or red; gland

appendages conspicuous, spreading radially,

obdeltoid, 0.3-1.3 mm long, 1.1-2 mm wide,

pink or white, glabrous, margin irregularly

toothed or shallowly lobed distally; bracteoles

0.7-0.8 mm long, adnate for c. 1/3 of their

length to involucre, free portion divided into

few to numerous, subulate glabrous segments.

Staminate flowers 7-10 per cyathium;

pedicels 0.9-1.3 mm long; staminal filaments

543

Halford & Harris, Euphorbia section Anisophyllum in Australia

c. 0.1 mm long. Pistillate flowers: styles 0.4-

0.5 mm long, spreading, smooth, glabrous,

each entire or scarcely bifid, the apices terete.

Capsules exserted from involucre on pedicel

to 2.5 mm long, broad to very broad-ovate

in lateral view, 1.7-1.8 mm long, 1.7-2 mm

across, shallowly 3-lobate with keels acute,

smooth, glabrous; hypogynous disc entire.

Seeds ovate in outline, 1-1.2 mm long, 0.6-

0.7 mm tangentially, 0.6-0.8 mm radially,

tetraquetrous in cross section; dorsal faces

planar or convex; ventral faces concave; all

faces with faint narrow irregular ridges;

exotesta thin, of even thickness over surface,

grey-white, microreticulate, becoming

mucilaginous when moistened; endotesta pale

brown or red-brown. Fig. 27K.

Additional selected specimens examined : Northern

Territory. Panu Panu Soakage, near Lake Surprise, Aug

1991, Latz 12135 (DNA, MEL); 50 km W of Newcastle

Waters homestead, Feb 1989, Thomson 3187 (DNA);

60 km NE [of] Bob Well, Lander River, Jul 1989, Latz

11556 (MEL); 11.4 km N of Elliot on Stuart Highway,

Apr 1983, Barker 194 (AD, NSW); 32 km S [of] Elliott

on Stuart Highway, Mar 1991, Thomson 3450 (DNA); 8

km N [of] Renner Springs, Stuart Highway, Feb 1988,

Thomson 2240 (DNA); 40 km ESE [of] Barrow Creek,

May 1996, Albrecht 7563 & Latz (DNA); Elkedra

Station, 7 km NW of homestead on road to Hatches

Creek via jump-up, Aug 1979, Morton 210 (DNA,

MEL); Little Lagoon, Groote Eylandt, Apr 1948, Specht

220 (AD, BRI, MEL, NSW); Calvert River mouth, Jun

1987, Thomson 1894 (DNA); Old Highland Plains, Jul

1971, Latz 1741 (DNA). Queensland. Burke District:

Nicholson River crossing, 3 km E of Doomadgee, Jun

1991, Halford 0607 (BRI); Musselbrook Creek Gorge,

27.6 km (by road) NE of Musselbrook Mining Camp,

175 km N of Camooweal - Lawn Hill N.P., Apr 1995,

Johnson MRS598 & Thomas (BRI); 88.5 km along

Richmond Road, 0.7 km S of Prospect Station turnoff;

Esmeralda Station, Apr 2007, McDonald KRM6433

(BRI); Paradise Swamp, c. 10 km NW of Burke & Wills

Roadhouse and N of the Burketown Development Road,

Feb 2006, Fox IDF3904 & Wilson (BRI); Richmond

- Croydon Road, 109.0 km from Richmond, Jul 1998,

Bean 13409 (BRI); Bunda Bunda, NE of Julia Creek, Apr

1999, Fensham 3759 (BRI). North Kennedy District:

29 km S of Yarrowmere homestead on track to Bowie

homestead. Whistling Duck Creek, Apr 2006, Halford

Q9035a & Batianoff (BRI). South Kennedy District: c.

18 km of Moonoomoo Station on road to Yarrowmere

Station from Bowie Station, Oct 1983, Henderson H2804

etal. (BRI).

Distribution and habitat : Euphorbia petala

occurs from Groote Eylandt, Newcastle

Waters and Lake Surprise in NT, eastward

to Charter Towers and near Aramac, north

Qld (Map 41). It grows in Triodia grassland

or open eucalypt/melaleuca woodland/open

forest on sand plains, sandstone rises, inland

sand dunes or sandy river terraces. It has also

been recorded in low monsoon forest. The

soils are sandy to sandy loam.

Phenology : Flowers and fruits have been

collected from February to October.

Typification: There are two sheets [1560387

& 503409] at MEL collected by Ewart at

Wycliffe Well in June 1924 and annotated with

the name E. petala that qualify as syntypes

of E. petala. The sheet MEL 1560387 is the

more ample specimen and is selected here as

lectotype of the name.

Notes : Euphorbia petala is similar to E.

albrechtii, E. cinerea and E. ophiolitica. For

features distinguishing E. petala from E.

albrechtii and E. cinerea , refer to the ‘Notes’

section under E. albrechtii.

42. Euphorbia philochalix Halford &

W.K.Harris, species nova quoad habitum

generali et formam seminis capsulaeque

similis E. verrucitestae Halford & W.K.Harris

sed ab ea glandulae appendicibus longioribus

0.1-0.3 mm longis (ad vicem appendicibus c.

0.5 mm longis), seminibus 0.9-1.1 mm longis

interdum brevibus (ad vicem seminibus 1.1-

1.2(1.3) mm longis), seminum superficiebus

porcis prominentibus angustis praeditis

irregularis (ad vicem superficiebus verrucosis)

distinguenda. Euphorbia philochalix

quondam putabatur eadem E. drummondii

Boiss. esse sed seminum superficiebus

irregulariter porcatis (ad vicem superficiebus

porcis 3-6 distinctis transversis), capsulis

latis usque perlatis 1.3-1.5(1.8) x 1.5-1.8(2.1)

mm plerumque minoribus (ad vicem capsulis

1.6—1.8 x 1.7-1.9 mm), seminibus late obovatis

(ad vicem seminibus oblongo-ovatis), stylis

bifidis 1/3—1/2 longitudinis (ad vicem stylis

integris vel vix bifidis) differt. Typus:

Western Australia. 37 km S of Denham, 24

May 1991, B.G.Thomson 3515 (holo: DNA;

iso: NT, according to label information on the

holotype sheet there are duplicates (isotypes)

lodged at PERTH n.v., K n.v).

544

Monoecious, annual to 15 cm high, many

stems arising from slender taproot, the

whole plant glabrous. Stems prostrate or

ascending, much branched, smooth or

often longitudinally ridged. Interpetiolar

stipules narrow-triangular, 0.9-1.2 mm

long, bipartite, glabrous; margin laciniate.

Leaves: petiole 0.1-1 mm long, smooth;

blade obovate or elliptic, 4.3-67 mm long,

2.3-4.2 mm wide, 1.2-1.9 times longer than

wide; adaxial and abaxial surfaces green

sometimes with reddish tinge especially along

margins, smooth or minutely papillose; base

symmetric, obtuse to cuneate or asymmetrical

with one side cordate, the other obtuse or

rounded; margin entire or sparingly minutely

toothed distally; apex retuse or rounded.

Cyathia solitary at the nodes, peduncles

0.3-0.5 mm long. Involucres campanulate

or turbinate, 0.8-1.1 mm long, 0.9-1.2 mm

across; lobes 5, triangular, 0.4-0.6 mm

long, margin entire or fimbriate; glands 4,

stipitate, cupuliform, with shallow central

pit and somewhat thickened rim, transverse-

oblong in outline, 0.1-0.3 mm long, 0.2-0.5

mm wide, pink or red; gland appendages

conspicuous, spreading radially, obdeltoid or

oblong, 0.1-0.3 mm long, 0.5-0.7 mm wide,

pink, glabrous, margin entire, crenulate or

shallowly irregularly lobed; bracteoles 0.8-

1.1 mm long, adnate for c. 1/3 of their length

to involucre, free portion divided into few

subulate hirsute segments. Staminate flowers

5-10 per cyathium; pedicels 1-1.5 mm long;

staminal filaments c. 0.1 mm long. Pistillate

flowers: styles 0.3-0.5 mm long, spreading

or ascending, smooth, glabrous, each bifid

for 1/3—1/2 of their length, the apices stout

terete. Capsules exserted from involucre on

pedicel to 2 mm long, broad to very broad-

ovate in lateral view, 1.3-1.5 mm long, 1.6-

1.8 mm across, shallowly 3-lobate with keels

acute, smooth, glabrous; hypogynous disc

entire. Seeds broad-ovate in outline, 0.9-1.1

mm long, 0.6-0.8 mm tangentially, 0.6-0.8

mm radially, tetraquetrous in cross section;

dorsal and ventral faces concave, with

prominent irregular rounded ridges; exotesta

thin, of even thickness over surface, white

or pale brown, microreticulate, becoming

mucilaginous when moistened; endotesta red-

brown. n= 11. Figs 17, 27L.

Austrobaileya 8(4): 441-600 (2012)

Additional selected specimens examined : Western

Australia, c. 8.5 km S of Useless Loop township

and Trig Station, Shark Bay, Oct 1997, Markey 1563

(PERTH); Salutation Island, Freycinet Estuary, Shark

Bay, Sep 1989, Alford 1316 (PERTH); 1 mile [c. 1.6 km]

5 of Useless Loop, Aug 1970, Aplin 3386 (PERTH);

Three Bay Island, Shark Bay, Sep 1989, Alford 1285

(PERTH); Baudin Island, Freycinet Estuary, Shark Bay,

Sep 1989, Alford s.n. (PERTH 4591437); 15 miles [c.

24 km] W of Wiluna, Aug 1973, Hassall 73101 (BRI);

Kalbarri N.P, Aug 1994, Cranfield 9272 (PERTH);

[Houtman] Abrolhos Islands, Nov 1897, Helms s.n.

(PERTH 2436752, 2436817); Little Rat Island, Easter

Group, Abrolhos Islands, Nov 1999, Harvey s.n.

(PERTH 6228739); Sachse Property, Snake Soak, Sep

2002, Davis WW76-35 (PERTH); c. 15 km SE of Bonnie

Rock, Sep 2001, Sage WW43-33 etal. (PERTH); c. 18.2

km NNW of Mt Dimer, Hunt Range, Jaurdi Station, Jul

1995, Gibson 3708 & Lyons (PERTH); 50 miles [c. 80

km] N of Kalgoorlie, Aug 1973, Hassall 73100 (BRI);

17 miles [c. 27 km] N of Kalgoorlie, Aug 1973, Hassall

7398 (BRI)[mixed collection with E. multifaria ]; near

Broad Arrow, 38 kmN of Kalgoorlie, Sep 1927, Gardner

6 Blackall s.n. (PERTH 2437201). Northern Territory.

Old Curtain Springs homestead, SW Mt Connor, Sep

1986, Thomson 1442 (DNA) [mixed collection with E.

porcata].

Distribution and habitat : Euphorbia

philochalix occurs from Shark Bay and the

Houtman Abrolhos Islands, east to near

Kalgoorlie, WA, with a disjunct population

near Mt Connor in southern NT (Map 42). It

grows in low shrubland or low eucalypt open

woodland communities on sand dunes, gentle

undulating terrain or limestone rises. The soils

are mostly brown to red sands or sandy clays,

often over limestone.

Phenology : Flowers and fruits have been

collected from May to November.

Notes : Euphorbia philochalix is similar to E.

verrucitesta in its general habit, and seed and

capsule shape. It may be distinguished from

that species by its longer gland appendages

(0.1-0.3 mm long versus c. 0.5 mm long for E.

verrucitesta ), generally shorter seeds (0.9-1.1

mm long versus 1.1-1.2(1.3) mm long for E.

verrucitesta ) and prominent narrow irregular

ridged seed surfaces (versus irregular wart¬

like protuberances on the seed surface for E.

verrucitesta).

Euphorbia philochalix has previously been

identified as E. drummondii. but differs from

that species in having prominent irregularly

ridged seed surfaces (versus seed surfaces

Halford & Harris, Euphorbia section Anisophyllum in Australia

545

Fig. 17. Euphorbiaphilochalix. A. habit *0.6. B. branchlet with cyathia x6. C. leaf *8. D. stipules xl6. E. cyathiawith

female flower x24. F. cyathia with fruit *16. G. cyathial gland with appendage, adaxial view *32. H. capsule, top view

xl6.1. capsule, lateral view xl6. A from Thomson 3515 (DNA); B-C, & F-I from Alford 1285 (PERTH); E from Hassall

73100 (BRI). Del. W.Smith

546

with 3-6 distinct transverse ridges for E.

drummondii ), capsules broad to very broad

ovate in outline which are generally smaller

in dimensions (1.3-1.5(1.8) x 1.5-1.8(2.1) mm

versus broadly elliptic in outline, 1.6-1.8 x

1.7-1.9 mm for E. drummondii ), seeds broad-

obovate in outline (versus oblong-obovate for

E. drummondii ) and styles that are bifid for

1/3—1/2 of their length (versus style entire or

scarcely bifid for E. drummondii ).

Etymology : The specific epithet is a noun in

apposition and is from Greek philo -, loving,

fond of, and chalix, calcium, in reference to

the usual habitat of this species, namely on

soils associated with limestone.

43. Euphorbia porcata Halford &

W.K.Harris, species nova quondam

putabatur eadem E. drummondii Boiss.

esse sed superficie dorsalis seminis porca

irregulari longitudinali in medio praedita

superficie ventrali seminis ± laevis (ad vicem

superficiebus seminis porcis 3-6 distinctis

transversis), glandulis involucralibus 0.1-0.15

x 0.1-0.2 mm sine ore incrassata (ad vicem

glandulis 0.15-2 x 0.2-0.4 mm) differt. Saepe

dicitur E. porcata cum E. ferdinandi Baill.

var. ferdinandi crescat sed stylis 1/3—1/2

longitudinis bifidis (ad vicem stylis integris

vel vix bifidis), glandulis involucralibus sine

ore incrassato seminis superficie dorsali

porca longitudinali irregulari ad medio

praedita (ad vicem superficie laevi), capsulis

longitudine latitudinem aequa perlato-ovatis

vel lato-ellipticis a latere visus 1.4-2 x 1 4-2

mm (ad vicem capsulis 1.4-1.9 x 1.3-1.7 mm)

distinguenda. Typus: Queensland. Gregory

North District: Ethabuka Station, c. 3 km

S of homestead, 112 kmNW ofBedourie, 26

June 2010, D.Halford QM84 & P.I.Forster

(holo: BRI, iso: AD, DNA, NSW, PERTH,

distribuendi ).

Monoecious, annual or herbaceous perennial

with thickened woody taproot, to 30 cm high,

many stems arising from rootstock, the whole

plant glabrous. Stems mostly prostrate or

ascending, sometimes erect, much branched,

smooth. Interpetiolar stipules narrow-

triangular, 0.5-1.1 mm long, bipartite,

glabrous; margin serrulate or laciniate.

Leaves: petiole c. 0.2 mm long, smooth; blade

Austrobaileya 8(4): 441-600 (2012)

oblong or oblanceolate, 4-8 mm long, 2-5 mm

wide, 1.6-2.4 times longer than wide; adaxial

surface pale green or reddish green, smooth

(rarely minutely papillose); abaxial surface

similar to but paler than adaxial surface; base

asymmetric with one side cordate or truncate,

the other cuneate or obtuse; margin serrulate

or sparingly minutely toothed; apex retuse

or rounded. Cyathia solitary at the nodes,

sometimes gathered together on short leafy

lateral branchlets with subtending leaves

slightly smaller than primary stem leaves;

peduncles c. 0.2 mm long, smooth. Involucres

turbinate, 0.5-0.8 mm long, 0.6-0.8 mm

across; lobes 5, triangular, 0.2-0.3 mm long,

margin entire; glands 4, stipitate, cupuliform,

with distinct central pit, transverse-oblong

to orbicular in outline, 0.1-0.15 mm long,

0.1-0.2 mm wide, red; gland appendages

inconspicuous or absent, spreading radially,

transverse-linear, 0.05-0.15 mm long, c. 0.2

mm wide, pink, glabrous, margin entire;

bracteoles 0.5-0.7 mm long, adnate for 1/3—

1/2 of their length to involucre, free portion

entire or divided into few to numerous

subulate glabrous segments. Staminate

flowers 3-10 per cyathium; pedicels 0.6-

1.1 mm long; staminal filaments 0.15 mm

long. Pistillate flowers: styles 0.2-0.4 mm

long, spreading, recurved distally, smooth,

glabrous, each bifid for 1/3—1/2 of their length,

the apices slender and terete or stout and

clavate. Capsules exserted from involucre on

pedicel to 1.3 mm long, very broad-ovate or

broad-elliptic in lateral view, 1.4-2 mm long;

1.4-2 mm across, shallowly 3-lobate with

keels obtuse, smooth, glabrous; hypogynous

disc entire. Seeds ovate in outline, 1-1.3

mm long, 0.5-0.7 mm tangentially, 0.6-0.7

mm radially, tetraquetrous in cross section;

dorsal faces convex or concave, with ± medial

longitudinal irregular ridge; ventral faces

concave, usually smooth or faintly irregularly

ridged; exotesta thin, of even thickness over

surface, white or pale brown, microreticulate,

becoming mucilaginous when moistened;

endotesta pale brown to red-brown, n— 11.

Figs 18, 27M.

Halford & Harris, Euphorbia section Anisophyllum in Australia

547

Fig. 18. Euphorbia porcata. A. habit *0.4. B. branchlet with cyathia *8. C. leaf x8. D. stipules xl2. E. cyathia with

female flower x24. F. cyathia with fruit xl6. G. cyathial gland with appendage, adaxial view x32. H. capsule, top view

xl6. I. capsule, lateral view xl6. A from Halford QM9 & Forster (BRI); B-I from Halford QM30 & Forster (BRI).

Del.W. Smith.

548

Additional selected specimens examined : Western

Australia. 18 km N of Overlander Roadhouse on North

West Coastal Highway, c. 33 km S of Gladstone, Sep

1985, Wilson 12201 (BRI, PERTH); 42 km from Agnew

along road to Wiluna, Sep 1982, Strid 20156 (PERTH);

Damboring, c. 46 km N of Wongan Hills, Mar 1968,

Wilson 6488 (PERTH); Rundall River N.P., Little

Sandy Desert, Apr 1979, Mitchell 873 (DNA). Northern

Territory. 46 km N [of] Three-Ways; Stuart Highway,

Feb 1988, Thomson 2241 (DNA); Stirling Bore, 20 miles

[c. 32 km] S of Barrow Creek township, Sep 1955, Perry

5345 (BRI, DNA, MEL, NSW); George Gill Range,

Stokes Creek, Oct 1986, Thomson 1534 (DNA); Maryvale

Station, junction of Blackhill Bore and Maryvale Road,

Aug 1988 Barritt 745 (DNA); Armata Road, 27 km S

[of] t/off from old road to Uluru, Apr 1988, Thomson

2298 (DNA). Queensland. Gregory North District:

Cravens Peak, 194 km by road SW of Boulia, Painted

Gorge, 12 km by road S of 12 Mile Bore, Apr 2007,

Thomas 3505 & Turpin (BRI); Cravens Peak, 13.6

km S along shotline track from Ocean Bore - Painted

Gorge Track, 135 km SW of Boulia, Jun 2010, Halford

QM9 & Forster (BRI); c. 33 km N of Bedourie on road

(Diamantina Development Road) to Boulia, Jun 2010,

Halford QM30 & Forster (BRI). Warrego District: 75

km NW of Charleville on road to Adavale, Sep 2005,

Halford Q8594 & Thomas (BRI). Gregory South

District: 25 miles [c. 40 km] E of Birdsville, May 1975,

Hassall 7521 (BRI); 3 miles [c. 4.8 km] N of Santos, May

1973, Hassall 7343 (BRI). South Australia. Dulkaninna

Station, Apr 1997, Smyth 48 (AD, MEL); Dulangari,

May 1986, Conrick 2057 (AD); Arabana Hill, 2 km N of

Lake Arthur, Mar 1987, Badman 1960 (AD); 0.5 miles

[c. 800 m] SW of Hawker Gate, on Quinyambie Station,

Jun 1955, Johnson 970 & Constable (NSW). New South

Wales. Cameron Corner, Sturt N.P, Sep 1989, Wiecek

250 et al. (BRI, NSW); 1.5 km NE of Joulnie homestead.

May 1973, Hassall 7318 (BRI); W bank of the salt lake,

45 km SSE of Milparinka, Sep 1971, DeNardi 822 (AD,

NSW); S shore of Menindee Lake, Kinchega N.P., Jun

1984, Dal by 84/06 (NSW).

Distribution and habitat: Euphorbiaporcata

is widespread across the arid zone in Australia

where it extends from WA, through SA and

the southern part of the NT to south-western

Qld and western NSW (Map 43).

It is recorded growing on sand dunes,

floodouts or in drainage lines or dune

swales, in hummock grassland, Acacia spp.

shrubland/woodland or Eucalyptus spp.

woodland communities, on mostly sandy to

sandy loam soils.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from April to October.

Notes: Euphorbia porcata has previously been

identified as E. drummondii, but differs from

Austrobaileya 8(4): 441-600 (2012)

that species in its dorsal seed surface having

a medial longitudinal irregular ridge and the

ventral seed surface ± smooth (versus seed

surfaces with 3-6 distinct transverse ridges

for E. drummondii ), and smaller involucral

glands that lack a thickened rim (0.1-0.15 x

0.1-0.2 mm versus 0.15-0.2 x 0.2-0.4 mm for

E. drummondii).

Euphorbia porcata has often been recorded

growing with E. ferdinandi var. ferdinandi.

It is distinguished from that species by its

divided styles (styles bifid for 1/3—1/2 of their

length versus entire or scarcely bifid for E.

ferdinandi var. ferdinandi ), involucral glands

lacking a thickened rim, dorsal seed surface

having a medial longitudinal irregular ridge

(versus seed surface smooth for E. ferdinandi

var. ferdinandi) and capsules as wide as long

(very broad-ovate or broad-elliptic in lateral

view, 1.4-2 mm x 1.4-2 mm versus elliptic

in lateral view, 1.4-1.9 x 1.3-17 mm for E.

ferdinandi var. ferdinandi).

The seeds are more or less uniform over

the eastern part of the species range. The

collections seen of this species from WA have

slightly shorter seeds with a more well defined

medial longitudinal ridge. Further collections

and field studies are warranted to establish the

significance of this variation.

Etymology: The specific epithet is from Latin

porcatus, ridged, in reference to the irregular

longitudinal medial ridge along the dorsal

faces of the seeds of this species.

44. *Euphorbia prostrata Aiton, Hort.

Kew. 2: 139 (1789); Chamaesyce prostrata

(Aiton) Small, FI. S.E. U.S. [Small] 713, 1333

(1903). Type: cultivated at Kew, origin in the

West Indies, in 1758, P.Miller s.n. (holo: BM

510671, n.v).

Illustrations: Auld & Medd (1987: 161);

Carter (1988: 422, fig. 78, 2); James & Harden

(1990: 428) as Chamaesyce prostrata; Lin

et al. (1991: 244, fig. 18), as Chamaesyce

prostrata.

Monoecious, annual to 5 cm high, many

stems arising from slender taproot. Stems

prostrate to weakly ascending, much branched,

smooth, with moderately dense to dense

549

Halford & Harris, Euphorbia section Anisophyllum in Australia

indumentum in a longitudinal band; hairs

spreading, straight to recurved, to 0.3 mm

long. Interpetiolar stipules narrow-triangular

to triangular, 0.3-1 mm long, bipartite or entire,

glabrous or occasionally with scattered hairs;

margin laciniate or obscurely toothed. Leaves:

petiole 0.4-1 mm long, smooth, glabrous or

nearly so; blade oblong, elliptic or rotund,

2.2-8.5 mm long, 1.2-4.5 mm wide, 1.5-2

times longer than wide; adaxial surface green

to blue-green occasionally with reddish tinge

along margin, smooth, glabrous; abaxial

surface grey-green occasionally with reddish

tinge along the midline, smooth, glabrous or

with sparse to moderately dense indumentum

consisting of spreading hairs 0.1-0.3 mm

long; base asymmetric with one side shallowly

cordate, the other rounded; margin sparingly

minutely toothed; apex acute to rounded.

Cyathia solitary at the nodes, often gathered

together on short leafy lateral branchlets with

subtending leaves usually slightly smaller

than primary stem leaves; peduncles 0.5-

2.5 mm long, smooth, glabrous. Involucres

turbinate, 0.5-07 mm long, 0.4-0.6 mm

across; lobes 5, triangular, 0.1-0.2 mm

long, margin fimbriate; glands 4, stipitate,

patelliform or cupuliform, with a tangential

trough or distinct central pit with a somewhat

thickened rim, transverse-oblong to orbicular

in outline, 0.05-0.1 mm long, 0.1-0.3 mm

wide, red or reddish-purple; gland appendages

conspicuous, spreading radially, transverse-

oblong, 0.05-0.1 mm long, 0.1-0.2 mm wide,

pink, red or white, glabrous, margin entire or

undulate; bracteoles filamentous, 0.4-0.5 mm

long, glabrous. Staminate flowers 3-5 per

cyathium; pedicels 0.5-0.7 mm long; staminal

filaments c. 0.1 mm long. Pistillate flowers:

styles 0.2-0.3 mm long, erect to spreading,

recurved distally, smooth, glabrous, each

bifid for c. 1/2 of their length, the apices

clavate. Capsules exserted from involucre on

pedicel to 2 mm long, broad to very broad-

ovate in lateral view, 1-1.5 mm long, 1.1-1.5

mm across, shallowly 3-lobate with keels

acute, smooth, with a sparse to moderately

dense indumentum mostly confined to the

keels; hairs spreading, 04-0.2(0.5) mm long;

hypogynous disc entire. Seeds ovate in outline,

0.8-1 mm long, 0.5-0.6 mm tangentially,

0.4-0.6 mm radially, tetraquetrous in cross

section; dorsal and ventral faces ± planar,

with 4-7 distinct transverse acute ridges;

exotesta thin, of even thickness over surface,

pale brown or grey-white, microreticulate or

microgranulate especially on ridge crests,

becoming mucilaginous (very thin layer)

when moistened; endotesta pale brown. Fig.

27N.

Additional selected specimens examined : Western

Australia, near work shed, Karijini Ranger Station,

Karijini N.P., Sep 2006, Halford Q9213 (BRI, PERTH);

Naval Base, Fremantle to Rockingham, Jan 1984,

Keighery 6533 (PERTH); roadside (W) on causeway N

end, W of T27 electricity pole. Garden Island, Jul 2003,

Dodd 847 (PERTH); 2.5 km W of Mundijong, junction

Mundijong and Kargotich Roads, Mar 1999, Davis 8796

(BRI, PERTH); corner of Kargotich & Mundijong Roads,

Mundijong, May 2009, Hislop 3878 (BRI, MICH).

Northern Territory. Camfield Station homestead, Jul

1983, Kernot s.n. (DNA [D0024788]); Brunette Downs

Station, Apr 1988, Kimbel 34 (DNA); Tangentyere

Nursery, Alice Springs, Nov 1995, Albrecht 7079 (DNA).

Queensland. Cook District: Boigu Island, c. 8 km S of

PNG mainland, Oct 1992, Waterhouse BMW1444 (BRI).

Burke District: Palm Grove, Lawn Hill N.P, May

1990, Latz 11635 (DNA). Leichhardt District: Charles

Street, Springsure, Aug 2004, Halford Q8320 (BRI);

Leichhardt Hotel, Taroom, Mar 2004, Halford Q8189

& Edginton (BRI). Port Curtis District: Raglan Creek

crossing, 21 km W of Mt Larcom, Nov 2003, Halford

Q8066 & Forster (BRI). Burnett District: Lions Park,

Kingaroy Street, Kingaroy, Mar 2004, Halford Q8171

& Edginton (BRI). Moreton District: Peregian Beach,

Avocet Parade, Jan 2000, Harris 112 (BRI); Guanaba,

Coomera River, 5 km SW of Oxenford, property of I.

Cairns, Apr 2003, Forster PIF29296 (BRI). New South

Wales. Kyogle District, Dec 1957, Vane s.n. (NSW

541519); Lismore, Mar 1957, Flynn s.n. (NSW 612664);

Narrabri Golf Club, 1996, s.coll. (NSW 398987); Bective

Travelling Stock Route, alongside Peel River, Jan 1995,

Hosking 1074 & Sullivan (NSW, MEL); Tamworth

Regional Botanic Gardens, Oxley Park, Tamworth, Mar

1999, Hosking 1692 & Bayliss (MEL, NSW); Strathfield,

Mar 1954, Ford s.n. (NSW 612601).

Distribution and habitat : Euphorbia

prostrata is a native to tropical and subtropical

America now naturalised throughout much of

the warmer regions of the world, including

Australia. In Australia it is common in urban

centres in coastal and subcoastal parts of Qld

and NSW, with scattered occurrences through

north-western Qld, NT and in the Pilbara

and Perth regions, WA (Map 44). It grows in

sunny situations on disturbed open ground,

road sides, river banks, lawns, garden beds

and in cracks of footpaths.

550

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from January to March.

Notes : In Australia, Euphorbia prostrata

has been confused with E. maculata and E.

australis. E. prostrata is easily distinguished

by having capsules sparsely to moderately

densely hairy along the keels and more or less

glabrous between the keels, and seed surface

with 4-7 distinct transverse acute ridges.

There are two forms of this naturalised

species in Australia. The most common and

widespread form has a slender habit with hairs

c. 0.2 mm long on the stems and the leaves

are ± glabrous except for a few scattered hairs

on the abaxial surface and along the margins.

The second form has a more robust habit with

hairs up to 0.3 mm long on the stems and

the leaves have a sparse to moderately dense

indumentum consisting of spreading, hairs

0.1-0.3 mm long. The longer indumentum

gives the plants a shaggy appearance. This

second form is recorded in the Perth region,

WA (e.g. Davis 8796, Keighery 6533, Hislop

3878 ) and at a single locality near Wandoan,

Qld (90 km NW of Wandoan, Jan 2005, Ward

s.n. [BRI (AQ723922)]).

45. Euphorbia psammogeton PS.Green,

Kew Bull. 48: 314 (1993); Chamaesyce

psammogeton (PS.Green) P.I.Forst. &

R. J.F.Hend., Novon 5: 323 (1995). Type: [New

South Wales.] Lord Howe Island, Blinky

Beach, 13 November 1963, P.S.Green 1625

(holo: K 186489).

Euphorbia atoto var. imbricata Boiss.,

in A.DC., Prodr. 15(2): 13 (1862). Type:

[Queensland. Port Curtis District:] Port

Curtis, November 1847, \J.\ Macgillivray B65

(holo: K).

Illustrations : Stanley & Ross (1983: 414, fig.

65B), as E. sparrmannii, James & Harden

(1990: 429), as Chamaesyce sparrmannii.

Green (1993: 315, fig. 2, Al-5; 1994: 236, fig.

47, A-D).

Monoecious, herbaceous perennial, to 35 cm

high, few to many stems arising from woody

taproot. Stems prostrate or ascending to

erect, sparingly to much branched, smooth,

Austrobaileya 8(4): 441-600 (2012)

glabrous. Interpetiolar stipules triangular to

broad-triangular, 1-2 mm long, entire or bifid

to bipartite, glabrous abaxially, hairy adaxially

with conspicuous ascending white hairs to

0.15 mm long; margin irregularly serrulate.

Leaves: petiole 2-3 mm long, smooth; blade

obovate, elliptic or oblong-elliptic, 12-28 mm

long, 5-17 mm wide, 1.3-2.4 times longer than

wide; adaxial surface green, blue-green or

grey-green, smooth, glabrous; abaxial surface

similar to but paler than adaxial surface;

base asymmetric with one side cordate, the

other rounded; margin sparingly minutely

toothed or entire; apex obtuse to rounded.

Cyathia in congested 4-6 branched dichasial

cymes together with a solitary cyathium at

the distal nodes; peduncles 4-15 mm long;

bracts leaf-like but smaller than the primary

stem leaves; cyathial peduncles 1-3(6) mm

long. Involucres turbinate, 1.5-2 mm long,

1.5-2 mm across; lobes 5, triangular, 0.4-0.8

mm long, margin entire or laciniate; glands

4, stipitate, patelliform, planar or shallowly

concave, transverse-oblong in outline, 0.4-0.5

mm long, 0.7-1 mm wide, pale green; gland

appendages conspicuous, spreading radially,

transverse-linear or lunate, 0.2-0.4 mm long,

0.6-1.3 mm wide, white, glabrous, margin

entire; bracteoles 0.9-1.2 mm long, adnate

for 1/3—1/2 of their length to involucre, free

portion divided into numerous ± linear hirsute

segments. Staminate flowers (2)5-10(20) per

cyathium; pedicels 0.9-1.6 mm long; staminal

filaments 0.3-0.5 mm long. Pistillate flowers:

styles 0.7-0.8 mm long, spreading, recurved

distally, smooth and glabrous, each bifid for c.

1/2 of their length, the apices terete. Capsules

exserted from involucre to 3.5 mm long,

very broad-ovate in lateral view, 2.8-3 mm

long, 3.5-3.9 mm across, shallowly or deeply

3-lobate with keels obtuse, smooth, glabrous;

hypogynous disc entire. Seeds orbicular or

very broad-ovate in outline, 1.5-1.8 mm long,

1.4-1.5 mm tangentially, 1.4-1.6 mm radially,

± suborbicular or somewhat trigonal in cross

section; ventral and dorsal faces convex, with

irregular narrow rounded ridges; exotesta

thin, of even thickness over surface, white,

microreticulate, not becoming mucilaginous

when moistened; endotesta red-brown, n = 8.

Fig. 270.

551

Halford & Harris, Euphorbia section Anisophyllum in Australia

Additional selected specimens examined: Queensland.

South Kennedy District: Eimeo Beach, c. 10 km N

of Mackay, Aug 1976, Henderson H2400 (BRI). Port

Curtis District: Freshwater Creek, Shoalwater Bay

Training Area, Nov 2001, Denley Samp55 (BRI); beach

between Emu Point and Rocky Point at Emu Park, Jul

1977, Batianoff & McDonald 109 (BRI); Deepwater

N. P, 40 km E of Miriam Yale, Oct 1989, Gibson TOI868

(BRI). Wide Bay District: Coonarr Beach, SE of

Bundaberg, Nov 1996, Bean 11243 (BRI); Woodgate,

Sep 2004, Halford Q8340a (BRI, MEL, NSW). Moreton

District: Bishop Island, E of Brisbane, Apr 1932, Blake

3336 & Everist (BRI); Hercules Bank, S of Brisbane

River, Jul 1930, Hubbard 3487 (BRI); unnamed island

between Tiger Mullet Channel and Whalleys Gutter, c.

6.5 km ENE of Jacobs Well, Sep 2003, Halford Q7915

& Batianoff (BRI). New South Wales. Fingal Point

[Head], c. 2.5 miles [c. 4 km] SE of Tweed Heads, Jun

1962, Constable 3050 (NSW); Angourie - Headland S of

blue pools small path, Yurigu N.P, Sep 1993, Clemesha

s.n. (NSW 279169); S of Woolgoolga - Headland, Jan

1971, Whaite & Whaite 3471 (NSW); Lord Howe Island,

North Beach, Mar 1971, Rodd 1759 (NSW); Lord Howe

Island, Blinky Beach, Mar 2001, Le Cussan 1123 (BRI);

Big Gibber, Myall Lakes, E of Bombah Point, Feb 1969,

Blaxell 207 (NSW); Wamberal Beach, c. 10 km E of

Gosford, Aug 1985, Bishop 803 etal. (NSW); Narrabeen,

Nov 1912, Hamilton s.n. (NSW 612475); Currarong

- Beecroft Head, Apr 1936, Rodway 2182 (NSW);

Currarong, Dec 1987, Clarke s.n. (NSW 612457).

Distribution and habitat : Euphorbia

psammogeton occurs along the east coast

from Mackay, Qld, south to Currarong, on the

south coast of NSW and Lord Howe Island

(Map 45). It grows on beach sands often on

exposed frontal dunes just above the high tide

mark.

Phenology : Flowers and fruits have been

collected throughout the year.

Notes : Euphorbia psammogeton is similar

to E. obliqua and E. pallens. It differs from

both species by having the seed surface with

faint irregular narrow rounded ridges (versus

seed surface smooth for E. obliqua and E.

pallens ), and larger gland appendages (0.2-

O. 4 mm long versus up to 0.1 mm long for E.

obliqua and E. pallens). It also differs from

E. obliqua by having its cyathia in dichasial

cymes (versus cyathia solitary at the nodes for

E. obliqua).

46. Euphorbia psilosperma Halford &

W.K.Harris, species nova ut videtur maxime

arete cum E. biconvexa Domin, E. coghlanii

F. M.Bailey, E. trigonosperma Halford &

W.K.Harris cognata sed ab eis capsulis

majoribus, seminibus comparate latioribus et

stylis ± integris. Hae differentiae in tabula 2

breviter repetant. Typus: Western Australia.

227 km SW [of] Broome on Great Northern

Highway, 11 May 1991, B.G.Thomson 3630

(holo: NT; iso: DNA).

Monoecious, herbaceous perennial to 20 cm

high, many stems arising from woody taproot.

Stems ascending to erect, much branched,

smooth, glabrous (rarely with a few scattered

white, spreading hairs c. 0.1 mm long).

Interpetiolar stipules narrow-triangular,

0.5-0.6 mm long, bipartite, glabrous; margin

entire. Leaves: petiole 1-2 mm long, smooth,

glabrous; blade narrow-oblong or narrow-

ovate, 7-20 mm long, 1-5 mm wide, 3.6-10

times longer than wide; adaxial surface green,

smooth, mostly glabrous; abaxial surface pale

green, smooth, glabrous; base asymmetric

with one side cordate, the other obtuse; margin

entire; apex acute to obtuse. Cyathia solitary

at the nodes, occasionally clustered on short

leafy lateral branchlets with subtending leaves

slightly smaller than primary stem leaves;

peduncles c. 1 mm long, smooth, glabrous.

Involucres turbinate, 0.7-0.8 mm long,

0.8-0.9 mm across; lobes 5, triangular, c. 0.5

mm long, margin entire or ciliate; glands 4,

stipitate, cupuliform, with shallow central

pit or longitudinal trough, transverse-elliptic

in outline, 0.2-0.3 mm long, 0.4-0.5 mm

wide, pale green or pink; gland appendages

conspicuous, spreading radially, transverse-

elliptic, 0.1-0.4 mm long, 0.4-1.2 mm wide,

white, glabrous, margin entire; bracteoles

0.5-0.8 mm long, adnate for c. 1/2 of their

length to involucre, free portion divided into

few subulate glabrous segments. Staminate

flowers 5-10 per cyathium; pedicels 0.7-1.2

mm long; staminal filaments 0.2-0.3 mm

long. Pistillate flowers: styles 0.4-0.6 mm

long, spreading, smooth, glabrous, each entire

or scarcely bifid distally, the apices terete.

Capsules exserted from involucre on pedicel

to 2.5 mm long, transversely broad-elliptic in

lateral view, 2-2.2 mm long, 2.8-3 mm across,

deeply 3-lobate with keels obtuse, smooth,

glabrous; hypogynous disc entire. Seeds

very broad-ovate in outline, 1.2-1.4 mm long,

0.9-1.2 mm tangentially, 0.9-1.3 mm radially,

tetragonous, trigonal or suborbicular in

552

cross section; dorsal and ventral faces planar

or convex, smooth; exotesta thin, of even

thickness over surface, grey-white, ± smooth,

becoming mucilaginous when moistened;

endotesta red-brown. Figs 19, 27P.

Additional selected specimens examined: Western

Australia. Derby to Broome road, 20.3 km S (by road)

of Derby, Apr 1985, Aplin 77 et al. (PERTH); 63 miles

\e. 101 km] E of Derby on road to Fitzroy Crossing,

Mar 1967, Power 190 (PERTH); just NW of Wolf Creek

Crater, Apr 1979, George 15290 (PERTH); between

Port Hedland and Mundabullangana Station, Feb 1962,

George 3339 (PERTH). Northern Territory. 2 km S

of Daly Waters on Stuart Highway, Feb 1988, Thomson

2237 (DNA); 28 km S [of] Lajamanu-Tanami, Feb 1988,

Wilson 1003 (DNA); Barkly Tableland, Attack Creek,

Stuart Highway, Jul 1974, Carr 2567 & Beaugiehoie

ACB46346 (MEL, NSW).

Distribution and habitat : Euphorbia

psilosperma occurs in scattered localities

from near Port Hedland, WA, eastward to

Daly Waters, NT (Map 46). It grows on red

sands on aeolian dunes or plains in Acacia

open shrubland or mixed low woodland

communities. It has been noted on specimen

labels, George 3339 & 15290 (PERTH), to be

growing in areas “recently burnt”.

Phenology : Flowers and fruits have been

collected in February and April.

Notes : Euphorbia psilosperma seems most

closely related to E. biconvexa, E. coghlanii

and E. trigonosperma. It differs from these

species by having larger capsules, relatively

broader seeds and more or less entire styles.

These differences are summarized in Table 2.

Etymology : The specific epithet is from Greek

psilo-, bare, bald, smooth, and -spermits,

seeded, in reference to the smooth seeds of

this species.

47. Euphorbia schizolepis F.Muell. ex

Boiss., in A.DC., Prodr. 15(2): 20 (1862);

Euphorbia schizolepis F.Muell. ex Boiss. var.

schizolepis , Benth., FI. Austral. 6: 47 (1873).

Type: [Northern Territory.] Hooker’s Creek,

s.d., F.Mueller s.n. (lecto [here designated]: K

186490; isolecto: MEL 61123).

Illustrations : Wheeler (1992: figs 184L,

185L, 186L); Dunlop et al. (1995: 218, fig. 72).

Austrobaileya 8(4): 441-600 (2012)

Monoecious, herbaceous perennial to 30

cm high, few to many annual stems arising

from crown of thick woody taproot. Stems

decumbent or erect, sparingly to much

branched, smooth, with a moderately dense

to dense indumentum; hairs white, weakly

appressed to ascending or spreading,

crispate, to (0.4)2 mm long. Interpetiolar

stipules subulate, 1-2.1 mm long, entire or

bipartite, hairy as for stems; margin entire

or laciniate or parted into few filiform

segments. Leaves: petiole 1-3 mm long,

smooth, with indumentum as for stems;

blade ovate to broad-ovate, sometimes

somewhat falcate, 11-22 mm long, 7-18 mm

wide, 1.2-3 times longer than wide; adaxial

surface green or glaucous, papillose, with a

sparse to moderately dense indumentum;

hairs spreading to ascending, crispate, to

0.8 mm long; abaxial surface similar to but

paler than adaxial surface; base asymmetric

with one side cordate, the other rounded;

margin serrulate sometimes only distally;

apex obtuse or acute. Cyathia solitary at the

nodes; peduncles 2-5.5 mm long, smooth,

indumentum as for stems. Involucres

turbinate, 1.7-2.5 mm long, 2.5-3 mm across;

lobes 5, triangular, 0.5-1 mm long, margin

entire; glands 4, stipitate, patelliform, planar

or shallowly concave, transverse-oblong or

transverse-elliptic in outline, 0.5-1.1 mm

long, 1-2.2 mm wide, pale green; gland

appendages conspicuous, spreading radially,

obdeltoid or oblong, 0.6-2 mm long, 2-3 mm

wide, pink or white, glabrous or moderately

densely hairy on abaxial surface and on

adaxial surface distally, margin toothed or

deeply laciniate; bracteoles 1.6-3 mm long,

adnate for 1/3—1/2 of their length to involucre,

free portion divided into few ± linear villose

or plumose segments. Staminate flowers

30-40 per cyathium; pedicels 2.2-3.8 mm

long; staminal filaments 07-1(1.5) mm long.

Pistillate flowers: styles 1.2-27 mm long,

connate at the base into a column for c. 1/4

of their length, erect to spreading, recurved

distally, smooth or papillose abaxially,

glabrous or pubescent abaxially, each bifid

for 1/3—1/4 of their length or entire, the apices

terete. Capsules exserted from involucre on

pedicel to 7 mm long, transversely broad-

Halford & Harris, Euphorbia section Anisophyllum in Australia

553

Fig. 19. Euphorbia psilosperma. A. habit *0.8. B. branchlet with cyathia x8. C. leaf *4. D. stipule xl6. E. cyathium

with female flower x24. F. capsule with cyathium xl2. G. cyathial gland with appendage, adaxial view x32. H. capsule,

top view xl2.1. capsule, lateral view xl2. A from Thomson 3630 (NT); B from Power 190 (PERTH); C, D, F-I from

Thomson 2237 (DNA); E from Wilson 1003 (DNA). Del. W.Smith.

554

elliptic in lateral view, 3.2-5 mm long; 3.6-5

mm across, shallowly to deeply 3-lobate

with keels obtuse, papillose, with a dense

indumentum; hairs spreading or appressed-

ascending, to 2 mm long; hypogynous

disc entire. Seeds oblong-ovate in outline,

2.5- 3.2 mm long, 1.8-2.3 mm tangentially,

1.6- 1.9 mm radially, ± tetragonous in cross

section; dorsal faces convex; ventral faces ±

planar; all faces with 4-6 prominent broad,

flat-topped, transverse ridges; exotesta of

uneven thickness over surface, distinctly

thicker on ridges, pale brown or grey-white,

microreticulate, becoming mucilaginous

when moistened; endotesta dark brown. Fig.

27Q.

Additional selected specimens examined : Western

Australia, old irrigated Research Station, which is 2

km SW of Kimberley Research Station, Kununurra, Mar

1997, Mitchell 4528 (BRI, PERTH); Smoke Creek, SW

of Lake Argyle, May 1980, Weston 12186 (PERTH); 2

km N of Lennard Gorge, King Leopold Range, Jun 1992,

Halford Q1448 (BRI, PERTH); Napier Range, Tunnel

Creek, Apr 1988, Dunlop 7753 & Simon (BRI); Kelly

Bore, c. 25 km E of Ord River homestead, Apr 1977,

Eichler 22385 (PERTH); 6 miles [c.10 km] NE of Gordon

Downs Station, Jul 1949, Perry 2441 (AD, BRI, DNA,

NSW). Northern Territory. Annaburroo Station, c. 1

km E of homestead T/O [turnoff]. May 2002, Risler 1788

& Cusack (DNA); Old Jim Jim Road, Kakadu N.P, Mar

1987, Clark 872 (DNA); Arnhem Land, Donydji, Jun

1989, Dunlop 8514 & White (DNA); 7 miles [c. 11 km]

W [of] Caledon Bay turnoff, Jun 1972, Lot: 2842 (DNA,

NSW); Kakadu Stage 3, road to Gunlom, Apr 1993,

Egan 1937 (DNA); Nitmiluk N.P, NE corner of Park,

Apr 2002, Michell 4056 (DNA); Arnhem Land, near

Emu Springs, Sep 1999, Cowie 8368 & Harwood (DNA);

3 miles [ c . 5 km] N of Katherine, Mar 1964, Lazarides

7057 (AD, BRI, DNA); Mataranka, Elsey N.P, Feb 1994,

Egan 3238 & Cowie (BRI); Blackfella Creek, Newry

Station, Mar 1989, Russell-Smith 7617 (DNA); Balbarini

Creek, 20 miles [c. 32 km] E [of] O.T. Downs, Mar 1959,

Chippendale 5524 (AD, BRI, DNA, NSW); Kalkaringi,

Mar 1990, Thomson 3485 (DNA); near Kalkarindji,

Mar 1997, Michell & Calliss 398 (DNA); 10 km S of

Mt Stanford, Jun 1974, Latz 5361 (BRI); Cattle Creek

Station, Aug 1994, Egan 4282 (DNA).

Distribution and habitat : Euphorbia

schizolepis is confined to northern Australia

from Wyndham, WA, east through

north-western NT to Gove; with disjunct

occurrences in the Napier and King Leopold

Ranges, WA and near Cape Crawford, NT

(Map 47). It grows in a variety of habitats,

including tussock grassland, shrubland or

eucalypt woodland communities on dark clay

Austrobaileya 8(4): 441-600 (2012)

soils on plains, or in eucalypt forest/woodland

or vine thicket communities on sandy to loam

soils on ridges, rocky hills and slopes, or

alluvial flats.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from January to July.

Typification: Boissier (1862) cited two

collections in his protologue of Euphorbia

schizolepis namely “Ad Hooker’s Creek

et in sinu Carpentarie Novae Hollandiae

sept. (F. Muller!).(v.s. in h. Kew)”. Two

collections, which are considered syntypes

of the name E. schizolepis , have been located

amongst material of Euphorbia on loan to

BRI from K [a: Euphorbia schizolepis F.von

Mueller, Hookers Creek, DM (K 186490); b:

Euphorbia schizolepis F.von Mueller, Gulf of

Carpentaria, DM (K 186491)].

The collection (a) from Hookers Creek (K

186490) is here selected as lectotype because

it is part of the original material and has

morphology that best matches the description

in the protologue of this species. There is

a collection at MEL [61123] “ Euphorbia

schizolepis F. Mueller, Upper Victoria River”

that is a very good match to the material

from Hookers Creek (K 186490) and is most

likely part of the same collection and is cited

here as an isolectotype. The collection (b)

from the Gulf of Carpentaria (K 186491) is

the holotype of Bentham’s E. schizolepis

var. glabra. Duplicates of this collection

have been located at G-DC (IDC microfiche

800-73. 2416: I. 4) and MEL [61122], All are

referable to E. carissoides.

Notes : Euphorbia schizolepis is a distinctive

species that seems most closely related to E.

kimberleyensis. For features distinguishing

E. schizolepis from E. kimberleyensis , refer

to the ‘Notes’ section under that species.

The hairs on the stems, leaves and capsules

of E. schizolepis are typically long (up to 2

mm) and weakly ascending to spreading. The

specimens Forster PIF6007 (Gove Peninsula,

11 km along Dalywoi Bay road [BRI]) and

Wannan 2640 & Anderson (Woodcutters

Mine, S of Darwin [BRI]) are atypical in

having short ( c . 0.2 mm long) appressed hairs.

Halford & Harris, Euphorbia section Anisophyllum in Australia 555

Table 2. Comparison of some morphological characters between Euphorbia psilosperma ,

E. biconvexa, E. coghlanii and E. trigonosperma.

Character

E. psilosperma

E. biconvexa

E. coghlanii

E. trigonosperma

seed shape in

outline

very broad-

obovate

narrow-obovate

or elliptic

oblong to

obovate

obovate to very

broad-obovate

seed shape

in transverse

section

tetrogonous,

trigonal,

suborbicular

biconvex

suborbicular

trigonal

seed size

(mm)

L2-l.4x0.9-

1.2 x 0.9-1.3

1-1.5 xO.5-0.9

x 0.9-1.1

1.2-1.4 x

0.8-1 x 0.8-1

1.2-1.6X 0.6-1 x

0.8-1

capsule size

(mm)

2-2.2 x 2.8-3

1.6-2 x 2-2.5

oo oo

1.5-2.1 x 1.9-2.6

style division

entire or

scarcely bifid at

the tip

bifid for 1/2 of

their length

bifid for

1/3-2/3 of

their length

bifid for 1/2-2/3

of their length

The species as circumscribed here varies

in the degree of style division. The typical

form has styles divided for 1/3—1/4 of their

length (e.g. Lazarides 7057 ' Mitchell 4528 ) and

is commonly observed in collections from the

Kimberley and Victoria River regions in the

southern part of the species range. A second

form with entire styles (e.g. Chippendale

5524 , Perry 2441 ) is more commonly recorded

in the northern part of the species range from

Darwin to Arnhem Land.

48. Euphorbia schultzii Benth., FI. Austral.

6: 47-48 (1873); Chamaesyce schultzii

(Benth.) D.C.Hassall, Aust. J. Bot. 24: 640

(1976). Type: [Northern Territory.] Port

Darwin, s.d., [M. Schultz] 844 (comm, by

Schomburgk, 7 December 1871) (lecto [here

designated]: K 186494).

Monoecious, annual or herbaceous perennial

with thickened woody taproot, 10-40 cm

high, with few to many stems arising from

base. Stems decumbent to erect or prostrate,

sparingly to much branched, smooth, with

a moderately dense indumentum (rarely

glabrous); hairs appressed to spreading,

curved or straight, 0.1—1 mm long, white.

Interpetiolar stipules subulate to narrow-

triangular, 0.4-1.2 mm long, deeply bipartite,

glabrous or with indumentum as for stems;

margin entire or laciniate with teeth often

gland-tipped. Leaves: petiole 0.4-1 mm long,

smooth, with indumentum as for stems; blade

narrow-oblong to oblong, narrow-ovate to

ovate or oblong-obovate to obovate, (3)4-16

mm long, 1.5-9 mm wide, 1.2-2.9 times

longer than wide; adaxial surface green

often with reddish tinge, smooth or papillose,

glabrous or with a sparse to moderately dense

indumentum consisting of ascending to

spreading, ± straight hairs 0.2-0.8 mm long;

abaxial surface pale green, pink, smooth or

papillose; glabrous or sparse to moderately

dense indumentum consisting of appressed-

ascending to spreading, ± straight hairs

0.1-0.8 mm long; base asymmetric with one

side cordate to auriculate, the other rounded

to cordate; margin serrate to serrulate, often

only along one side and distally (rarely entire);

apex obtuse to acute or rounded. Cyathia

solitary at the nodes, often clustered on short

leafy lateral branchlets with subtending leaves

slightly smaller than primary stem leaves;

peduncles 0.1-1.5 mm long. Involucres

turbinate to cupuliform, 0.8-0.9 mm long,

0.7-1.4 mm across; lobes 5, triangular, c. 0.4

mm long, margin fimbriate or entire; glands 4,

stipitate, cupuliform, with distinct tangential

trough and thickened rim, transverse-oblong

in outline, 0.1-0.3 mm long, 0.4-0.6 mm

wide, pink; gland appendages conspicuous,

spreading radially, very broad-obovate or

transverse-oblong, 0.1-0.6 mm long, 0.4-0.8

mm wide, white to pink, glabrous, margin

entire, erose or shallowly lobed; bracteoles

0.5-0.8 mm long, adnate for c. 1/5 of their

556

length to involucre, free portion divided

into numerous subulate glabrous segments.

Staminate flowers 4-20(27) per cyathium;

pedicels 0.7-1 mm long; staminal filaments

0.1-0.3 mm long. Pistillate flowers: styles

0.2-0.6 mm long, spreading, recurved distally,

smooth, glabrous or sparsely hairy, entire

or bifid for 1/3-2/3 the length, with terete

apices. Capsules exserted from involucre

on pedicel to 3 mm long, depressed ovate or

transversely elliptic to broad-elliptic in lateral

view, broadest at or below the equator, 1.5-

2.2 mm long, 2-3 mm across, deeply 3-lobate

with keels acute, papillose (rarely smooth),

glabrous or with a sparse to moderately

dense indumentum consisting of appressed to

spreading hairs to 0.2 mm long; hairs evenly

distributed over surface (rarely confined to

keels); hypogynous disc laciniate or entire.

Seeds ovate or broad-ovate in outline, 1—1.5

mm long, 0.6-0.8 mm tangentially, 0.6-0.8

mm radially, tetraquetrous in cross section;

dorsal faces convex to planar; ventral

faces planar to concave; all faces with 3-5

prominent ± transverse or irregular, rounded

ridges; exotesta thin, of even thickness over

surface or of uneven thickness and distinctly

thicker on ridges, grey-white, microreticulate,

becoming mucilaginous when moistened;

endotesta pale to dark brown.

Distribution and habitat: Euphorbia schultzii

is widespread across northern Australia

from the Kimberley, WA through the NT to

northern Qld.

Austrobaileya 8(4): 441-600 (2012)

Typification : Bentham (1873) in his

protologue of Euphorbia schultzii cited four

collections by M. Schultz from Port Darwin

settlement, NT (“N. Australia. Port Darwin,

Schultz, n. 15, 237, 844 and 879”). All of

these have been located amongst material of

Euphorbia on loan to BRI from K: Schultz 15

(K 186495); Schultz 844 (K 186494); Schultz

879 (K 186493); Schultz 237 (K 186492).

The collection Schultz 844 (K 186494) is

here chosen as the lectotype of Euphorbia

schidtzii because it has morphology that

agrees with the description in the protologue,

is the best preserved and most ample of the

original material, and has mature capsules

and seed attached. The syntypes Schultz 15

(K 186495); Schultz 844 (K 186494); Schultz

879 (K 186493); Schultz 237 (K 186492) are

all referable to E. schultzii var. schultzii as

applied here.

Notes : Euphorbia schultzii has been confused

with E. careyi and E. australis in the past. It

can be distinguished from both species by

the capsules that are distinctly broader than

long (1.5-2.2 mm long x 2-3 mm across)

and deeply 3-lobate (versus capsules that are

more or less as long as wide (1.4-2 x 1.5-2.1

mm for E. australis ; 1.4-1.6 x 1.6-1.9 mm

for E. careyi ). It also differs from E. careyi

in having cupuliform involucral glands with

a distinct tangential trough and thickened

rim rather than patelliform with planar or

shallowly concave gland surface.

Two varieties are recognised which can be

distinguished using the following key.

Key to varieties of Euphorbia schultzii

Capsule hairy.48a. E. schultzii var. schultzii

Capsule glabrous.48b. E. schultzii var. comans

48a. Euphorbia schultzii var. schultzii

Euphorbia australis var. glaucescens Boiss.,

in A.DC., Prodr. 15(2): 36 (1862). Type:

[Queensland. Cook District:] sandy banks of

the Gilbert River, s.d., F.Mueller s.n. (holo: K

186466; iso: MEL 503406, P 698528 element

on the right hand side of sheet).

Illustrations: Wheeler (1992: figs 184M,

185M, 186M); Dunlop et al. (1995: 218, fig.

72).

Stems erect or prostrate, with a moderately

dense indumentum; hairs appressed or

spreading, curved or straight, 0.1-1 mm long.

Leaf blades: narrow-oblong to oblong, ovate

557

Halford & Harris, Euphorbia section Anisophyllum in Australia

or oblong-obovate to obovate, (3)4-16 mm

long, 1.5-9 mm wide, 1.7-2.9 times longer than

wide; adaxial surface glabrous or with a sparse

to moderately dense indumentum consisting

of ascending to spreading, ± straight hairs

0.2-0.8 mm long; abaxial surface glabrous

or with a sparse indumentum consisting of

hairs similar to those on adaxial surface; base

asymmetric with one side cordate, the other

cuneate to rounded. Involucres turbinate

to cupuliform, 0.8-0.9 mm long, 0.7-1.4

mm across; gland appendages conspicuous,

spreading radially, transverse-oblong, 0.1-0.6

mm long, 0.7-0.8 mm wide, margin entire

or irregularly toothed. Staminate flowers

10-20 per cyathium. Pistillate flowers: styles

0.2-0.6 mm long, glabrous or sparsely hairy,

entire or bifid for 1/3-2/3 of their length.

Capsules depressed ovate or transversely

elliptic in lateral view, 1.5-2.2 mm long,

2-3 mm across, with a sparse to moderately

dense indumentum consisting of appressed to

spreading hairs to 0.2 mm long; hairs evenly

distributed over surface (rarely confined to

keels); hypogynous disc laciniate or entire, n

= 11 Fig. 27R.

Additional selected specimens examined: Western

Australia, along the Ord River at Ivanhoe Crossing, Apr

1977, Eichler 22177 (PERTH); base of Mt Nyulasy, 198

km N of Halls Creek on road to Kununurra, c. 35 km N

of Turkey Creek, Apr 1985, Aplin 1347 et al. (PERTH);

Windjana Gorge N.P, outside drip line immediately in

front of Carpenters Gap rockshelter, July 1997, Wallis

LW97A/37 (PERTH); along Millie Windie Road (from

Lennard River crossing), Jun 1988, Wilson 12952

(PERTH); Ord River between Springvale and Bedford,

Apr 1972, Aplin 4851 (PERTH); 39 km S of Forrest

River on Duncan Highway, Apr 1977, George 14452

(PERTH); Geike Range, near Geike Gorge, Apr 1988,

Simon 3986 & Cranfield (BRI). Northern Territory.

Meckitt Creek, 2 km S [of] boat ramp, Dec 1986,

Wightman 3301 (DNA); Cape Hotham, Escape Cliff,

Mar 1993, Cowie 3315 (DNA); 11.2 miles [c. 18 km] S

[of] Batchelor, Mar 1961, Chippendale 7739 (BRI, DNA,

MEL, NSW); Fitzmaurice River, Feb 1994, Leach 4071

(BRI, MEL); Binjari Settlement, 15 km W of Katherine,

Mar 1996, Barritt 2094 (DNA); Gregory N.P, past Matt

Wilson Lookout, Feb 1992, Cowie 2239 & Brocklehurst

(DNA, MEL). Queensland. Cook District: Royal Arch

Cave section, Chillagoe - Mungana Caves N.P, Jan

2005, McDonald KRM3542 & Little (BRI); Newcastle

Range between Georgetown and Mt Surprise, Jan 2005,

McDonald KRM3499 (BRI); Mt Eliza, 8 km NW of

Mount Surprise, Jan 1993, Bean 5498 & Forster (BRI).

Burke District: E of the Burke Development Road

at the Bang Bang Jumpup, Feb 2006, Fox IDF3829 &

Willson (BRI). North Kennedy District: ‘Maidavale’, E

of Mingela, Apr 1991, Bean 2952 (BRI); c. 67 km S of

Charters Towers near Mt Malakoff, Jun 1998, Thompson

CHA451 & Turpin (BRI).

Distribution and habitat : Euphorbia

schultzii var. schultzii occurs across northern

Australia from the Kimberley region, WA,

through the northern area of the NT to near

Townsville, north Qld (Map 48a). It grows in

eucalypt/Acacz'tf woodland or eucalypt open

forest communities on rocky hills, plateaux

or riverine flats, rarely in Spinifex grassland

on coastal dunes or rocky foreshores. The

soils are mostly sandy to loam derived from a

range of substrates.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from January to June.

Notes : As accepted here, there is some

variation in the indumentum form and

distribution, and style division of Euphorbia

schultzii var. schultzii. Generally, specimens

fall into those with curved, appressed to

ascending hairs 0.1-0.3 mm long and those

with ± straight, spreading hairs 0.2-1 mm

long. Both forms occur throughout the

variety’s distributional range.

Euphorbia schultzii var. schultzii typically

has styles divided for 1/3-2/3 of their length.

A number of specimens with entire styles (e.g.

Bean 2952, McDonald KRM3499) have been

collected from the eastern end (Mt Isa to the

Townsville region, Qld) of the varieties range.

The indumentum on the capsules of this

variety is typically evenly distributed over

the capsule surface. Specimens Eichler 22177

(from WA) and Fox IDF3829 (from Qld)

are atypical in having the hairs on capsules

restricted to their keels.

Collections from the Kimberley, WA

generally have narrower ridges on the seed

surface and generally shorter (<0.2 mm

long) spreading hairs on the stems leaves and

capsules (e.g. Wilson 12952, Aplin 4851, Aplin

1347 et al). Further investigations of these

forms are warranted.

558

48b. Euphorbia schultzii var. comans

(W.Fitzg.) Halford & W.K.Harris combinatio

et status nova; Euphorbia comans W.Fitzg.,

J & Proc. Roy. Soc. Western Australia 3:

161 (1918). Type: Western Australia. Derby,

April 1905, W.VFitzgerald 193 (lecto [here

designated]: PERTH 2847221; isolecto: NSW

98808).

Illustration : Wheeler (1992: figs 184D, 185D,

186D), as Euphorbia comans.

Stems decumbent to erect (rarely prostrate),

with a moderately dense indumentum

(rarely glabrous); hairs spreading, ± straight,

0.2-0.9 mm long. Leaf blades: narrow-

oblong to oblong, narrow-ovate to ovate,

6-15 mm long, 2-9 mm wide, 1.2-2.5 times

longer than wide; adaxial surface glabrous

or with a sparse indumentum consisting of

spreading, ± straight hairs 0.5-0.6 mm long;

abaxial surface glabrous or with a sparse to

moderately dense indumentum consisting

of spreading, ± straight hairs 0.5-0.8 mm

long; base asymmetric with one side cordate

to auriculate, the other rounded to cordate.

Involucres turbinate, c. 0.7 mm long, 0.7-1.4

mm across; gland appendages conspicuous,

spreading radially, very broad-obovate or

transverse-oblong, 0.1-0.4 mm long, 0.4-0.6

mm wide, margin entire, erose or shallowly

lobed. Staminate flowers 4-20(27) per

cyathium. Pistillate flowers: styles 0.2-

0.4 mm long, glabrous, bifid for 1/3-2/3

the length. Capsules depressed ovate or

transversely elliptic to broad-elliptic in lateral

view, 1.5-1.8 mm long, 2-2.7 mm across,

glabrous; hypogynous disc entire. Fig. 28A.

Additional selected specimens examined : Western

Australia. 11 miles [c. 18 km] S of Derby, Jan 1971, Allan

565 (PERTH); Manguel Creek Station S of the Broome

to Derby Road, Apr 1968, Payne D (PERTH). Northern

Territory. Victoria Highway, 57 km W of Katherine,

Dec 1994, Barritt 1693 (DNA, MEL); Nitmiluk N.P,

Mar 2001, Michell 3211 (DNA); Cutta Cutta Reserve,

Jan 1993, Egan 831 (DNA); 43.3 miles [c. 70 km] SE

[of] Top Springs Store, Apr 1959, Chippendale 5795

(DNA, MEL); 6 km N of Dunmarra, Jan 1994, Egan

3025 (DNA); Foelsche River, Jan 1989, Thomson 2956

(DNA); 30 miles [c. 48 km] SSW of Wavehill Station,

Jun 1949, Perry 2220 (BRI, DNA); Birrindudu Range,

NW Tanami Desert, May 2004, Latz 19970 & Brennan

(NT); 50 km W [of] Newcastle Waters homestead, Feb

1989, Thomson 3188 (DNA); 20 km NE [of] Elliott, Feb

1989, Thomson 3218 (DNA); 3 miles [c.4.8 km] NE [of]

Austrobaileya 8(4): 441-600 (2012)

Newcastle Waters, Mar 1959, Chippendale 5431 (BRI,

DNA, MEL); 50 km N of Renner Springs on the Stuart

Highway, Feb 1988, Thomson 2238 (DNA); Mittiebah

Station, Mitchiebo Waterhole, Mar 1981, Henshall 3481

(AD, BRI); 46 km N of Three Ways on Stuart Highway,

Feb 1988, Thomson 2242 (DNA). Queensland. Burke

District: 28.2 km S by road from Musselbrook Mining

Camp on road to Camooweal, Apr 1995, Thomas &

Johnson MRS435 (BRI); 1 km N of Dugald River

crossing on Julia Creek - Normanton Road, Mar 1977,

Farrell TF782 (BRI); 28.4 km by road from junction of

Burke and Gulf Development Roads, towards Flinders

River, Jan 2005, McDonaldKRM3447 (BRI).

Distribution and habitat : Euphorbia

schultzii var. comans occurs from Derby,

WA, across the north of the NT to Cloncurry,

north-western Qld (Map 48b). It grows in

eucalypt open woodland or Acacia/Grevillea

shrubland communities on sandy soils, on

alluvial flats, sand plains, sandstone plateaux

or rocky rhyolite, granite or laterite hills. Also

found in vine thicket on basalt soils.

Phenology : Flowers and fruits have been

collected from December to June (mostly

from January to April).

Typification: Fitzgerald (1918) based his

description of Euphorbia comans on material

collected by him from near Derby, Denham

and King Rivers, Kimberley, WA. All of

these have been located amongst material of

Euphorbia on loan to BRI from PERTH and

NSW: Derby, Apr 1905, W.VFitzgerald 193

(PERTH 2847221, NSW 98808); Denham

River, Oct 1906, W.VFitzgerald s.n. (NSW

612560); King River, Oct 1906, W.VFitzgerald

s.n. (NSW 612552). The collection Fitzgerald

193 (PERTH 2847221) is here chosen as

the lectotype of E. comans because it has

morphology that agrees with the description

in the protologue, and is the best preserved

and most ample of the original material. All

the syntypes seen are referable to E. schultzii

var. comans as applied here.

Notes : We have reduced Euphorbia comans

to a synonym of E. schultzii and recognised

the taxon at the varietal level. It differs from

E. schultzii var. schultzii in having glabrous

capsules.

Most of the specimens of this variety seen

by us have typically spreading hairs on the

stems. A rare form with glabrous stems is

559

Halford & Harris, Euphorbia section Anisophyllum in Australia

represented by Strong 975 , 976 (Sailor Creek,

Rosewood Station, May 1987 [DNA]) and

Russell-Smith 7614 & Lucas (Blackfellow

creek, Newry Station, Mar 1989 [BRI, NT]).

49. *Euphorbia serpens Kunth, in Humb.,

Bonpl. & Kunth, Nov. Gen. Sp. 2: 52 (1817);

Chamaesyce serpens (Kunth) Small, FI. S.E.

U.S. [Small] 709 (1903). Type: Venezuela.

“Cumana prope Bordones et Punta Araya”,

F.W.H.A. von Humboldt & A.Bonpland 407

(holo: P-Bonpl. n.v. [IDC microfiche 6209-1.

32: II. 4]).

Illustration : Lin et al. (1991: 246, fig. 9), as

Chamaesyce serpens.

Monoecious, annual or herbaceous perennial

with slender taproot, many stems arising

from rootstock, the whole plant glabrous.

Stems prostrate, much branched, smooth,

rooting at nodes. Interpetiolar stipules

broad-triangular, 0.5-0.6 mm long, glabrous;

margin lacerate distally. Leaves: petiole 0.3-

0.6 mm long, smooth; blade broad-oblong,

broad-elliptic or rotund, 2.5-4 mm long, 2.5—

3.7 mm wide, 1-1.2 times longer than wide;

adaxial and abaxial surfaces green, smooth;

base ± symmetric, cordate; margin entire;

apex retuse. Cyathia solitary at the nodes;

peduncles 0.5-2 mm long, smooth, glabrous.

Involucres turbinate, 0.5-0.6 mm long, 0.7-

0.8 mm across; lobes 5, triangular, 0.3-0.4

mm long, margin fimbriate; glands 4, stipitate,

patelliform, planar or shallowly concave,

transverse-oblong in outline, c. 0.15 mm long,

0.15-0.3 mm wide, purple; gland appendages

conspicuous, spreading radially, transverse-

oblong or obdeltoid, 0.1-0.2 mm long, 0.3-0.6

mm wide, white, glabrous, margin entire or

shallowly lobed; bracteoles 0.8-1 mm long,

adnate for c. 1/5 of their length to involucre,

free portion divided into few ± linear hirsute

segments. Staminate flowers 5-8 per

cyathium; pedicels c. 1 mm long; staminal

filaments c. 0.1 mm long. Pistillate flowers:

styles c. 0.4 mm long, spreading, smooth,

glabrous, each bifid for c. 1/4—1/2 of their

length, the apices terete. Capsules exserted

from involucre on pedicel to 1.5 mm long,

broad-elliptic in lateral view, 1.2-1.4 mm long,

1.4-1.5 mm across, shallowly 3-lobate with

keels obtuse, smooth, glabrous; hypogynous

disc laciniate. Seeds broad-ovate in outline,

0.8-0.9 mm long, 0.6-0.7 mm tangentially,

0.6-0.7 mm radially, tetraquetrous in cross

section; dorsal faces planar; ventral faces

planar or concave; all faces with 2-4 faint

narrow rounded, transverse ridges; exotesta

thin, of even thickness over surface, white,

microreticulate, becoming mucilaginous

when moistened; endotesta pale brown. Fig.

28B.

Additional selected specimens examined: South

Australia. Hundred of Wiltunga, section 302, May

1967, Coley 1316 (AD). Queensland. Burke District:

Resolution Street, Hughenden, Sep 2009, Halford

Q9699 (BRI). North Kennedy District: Dotterel Close,

Douglas, Townsville, Sep 2006, Hooker NH693 (BRI);

Racecourse Road, Ross River South Bank, Townsville,

Jan 2005, Camming RJC23142 (BRI). Maranoa

District: McDowall Street, Roma, Feb 2004, Halford

Q8135 & Harris (BRI). Darling Downs District:

Jimbour, railway line 1 km NW of township, Jan 2001,

Forster PIF26562 & Harris (BRI, MEL). Moreton

District: Tinchi Tamba Wetland Reserve, Brisbane, Dec

2009, Halford Q99745 (BRI). New South Wales. 10 km

W of Balranald, Dec 1987, Thomson 2300 (NSW [mixed

collection with E. dallachyana], NT).

Distribution and habitat : Euphorbia serpens

is native of the Americas and is naturalised

in Asia, Africa and Australia. In Australia, at

present it occurs in scattered localities; in Qld

from Townsville, Hughenden, Roma, Jimbour

and the Brisbane area, in western NSW near

Balranald; and on northern Yorke Peninsula,

SA (Map 49). It grows in disturbed sites,

including roadsides, railway easements,

footpaths, urban parklands, garden beds and

lawns.

Phenology : Flowers and fruits have been

collected in January, February, May, August

and December.

Notes : Euphorbia serpens is somewhat

similar to and easily confused with the native

E. dallachyana. It differs by having stems

rooting at the nodes and generally smaller

leaves, capsules and seeds.

The collection, Copley 1316 (AD) (from a

home garden, Yorke Peninsula, SA) is placed

here; however, it is atypical when compared

with the rest of the material of E. serpens in

Australia in having somewhat longer stipules

that are bipartite.

560

50. Euphorbia sharkoensis Baill., Adansonia

6: 287 (1866). Type: [Western Australia.]

Sharks Bay, Useless Harbour, in 1863, Maitl.

Brown s.n. (holo: P 698526 n.v. [image seen];

iso: MEL 503410).

Euphorbia drummondii var. rubescens

Benth., FI. Austral. 6: 49 (1873). Type:

[Western Australia.] Dirk Hartog’s Island,

January 1822, A.Cunningham 223 (holo: K

186499).

Monoecious, annual or herbaceous perennial

to 30 cm high, few to many stems arising from

slender taproot, the whole plant glabrous.

Stems prostrate or erect, sparingly to much

branched, smooth, sometimes longitudinally

ridged. Interpetiolar stipules subulate,

0.1-0.9 mm long, bipartite, glabrous; margin

entire or laciniate. Leaves: petiole 0.5-1.5

mm long, smooth; blade oblong, elliptic or

obovate to oblong-obovate, 3.1-14 mm long,

2.2-7 mm wide, 1.4-2.8 times longer than

wide; adaxial and abaxial surface blue-green

or green sometimes with reddish tinge along

margin, smooth, glabrous; base asymmetric

with one side cordate, the other cuneate or

rounded; margin entire or serrulate to serrate;

apex rounded to obtuse. Cyathia solitary

at the nodes; peduncles 0.3—1 mm long,

smooth, glabrous. Involucres campanulate

or turbinate, 0.5-0.9 mm long, 0.8-1.2 mm

across; lobes 5, triangular, 0.2-0.5 mm

long, margin fimbriate or laciniate; glands

4, stipitate, patelliform, planar or shallowly

concave with tangential trough, transverse-

oblong in outline, 0.2-0.3 mm long, 0.4-0.6

mm wide, red; gland appendages conspicuous,

spreading radially, transverse-oblong, 0.1-0.3

mm long, 0.4-0.7 mm wide, red or white,

glabrous, margin toothed or deeply lobed;

bracteoles 0.4-1 mm long, adnate for c. 1/3 of

their length to involucre, free portion divided

into few subulate glabrous or hirsute segments.

Staminate flowers 3-8 per cyathium;

pedicels 0.4-1.2 mm long; staminal filaments

c. 0.1 mm long. Pistillate flowers: styles 0.4-

0.6 mm long, spreading or ascending, smooth

and glabrous, each bifid for 1/3—1/4 of their

length, the apices terete. Capsules exserted

from involucre on pedicel to 3 mm long, very

broad-ovate or broad-elliptic in lateral view,

Austrobaileya 8(4): 441-600 (2012)

1.5-2 mm long, 1.5-2.3 mm across, shallowly

3-lobate with keels acute or obtuse, smooth,

glabrous; hypogynous disc entire (rarely

laciniate). Seeds ovate in outline, (0.9)1—1.5

mm long, 0.7-0.8 mm tangentially, 0.6-0.8

mm radially, tetraquetrous in cross section;

dorsal faces planar; ventral faces concave; all

faces with faint rounded, irregularly ridges;

exotesta thin, of even thickness over surface,

or sometimes slightly thicker on ridges, white,

microreticulate, becoming mucilaginous

when moistened; endotesta red brown, n = 11.

Additional selected specimens examined : Western

Australia. Airlie Island, July 1987, Long VL158

(PERTH); Burgundy Bay (Bomb site area). Alpha

Island, Montebello Islands, Oct 2000, Kenneally 11610

(PERTH); Varanus Island, Lowendale Group, May 1991,

Thomson 3496 (DNA, PERTH); Barrow Island, Oct

1980, Buckley 6956 (PERTH); Barrow Island, Oct 1980,

Buckley 7159 (PERTH); Legendre Island, May 1991,

Thomson 3506 (PERTH); Legendre Island, Dampier

Archipelago, Jun 1962, Royce 7322 (PERTH); 3.5 miles

[c. 5.6 km] S of Exmouth township. May 1965, George

6591 (PERTH); Ward Reef track on Thevenard Island,

May 1990, White MRW041 (PERTH); 10.5 km S of

Exmouth P.O., on track to beach, 0.5 km E of highway,

Jul 1977, McFarland & McFarland 4 (BRI, PERTH);

c. 30 km S of Exmouth, along W side of highway, Jul

1977, McFarland & McFarland s.n. (PERTH 3995879);

near Norcape Lodge, Exmouth, Jul 1977, McFarland &

McFarland s.n. (BRI [AQ234287]); top of Beagle Hill,

Pt [Point] Quobba, c. 50 km NNW of Carnarvon, Sep

1987, Wilson 12623 (PERTH); 25 miles [c. 40 km] N of

Carnarvon on Quobba Road, Sep 1970, George 10139

(BRI, PERTH); Dorre Island, Aug 1977, Weston 10547

(PERTH); S of Eagle Bluff, Peron Peninsula, Shark

Bay, c. 3.5 km SSE of intersection of Eagle Bluff Road

and Denham - Hamelin Road, Oct 1997, Markey 1625

(PERTH); Taillefer Isthmus, Shark Bay, c. 6 km NNW of

repeater station site near Goulet Bluff, Oct 1997, Markey

1797 (PERTH); c. 10 km N of Bibby Giddy Outcamp

on Heirisson Prong, 30 m SSE of Clough’s Bar track at

c. 2.4 km along track from junction with Useless Loop

Road, Sep 1997, Markey 1733 (PERTH); 10 miles [c. 16

km] S of Overlander (near Hamelin Pool), Aug 1973,

Hassall 73103 (BRI); Port Gregory rubbish tip, 2.5 km

SE of town, July 1997, Davis 3612 (PERTH).

Distribution and habitat : Euphorbia

sharkoensis is confined to the islands and

coastal areas of north western WA, from

Roeburne to Shark Bay, with a disjunct

occurrence near Port Gregory (Map 50).

It grows in open shrubland or heathland

communities on sandy soils on rocky

limestone foreshores or coastal sand dunes.

561

Halford & Harris, Euphorbia section Anisophyllum in Australia

Phenology : Flowers and fruits have been

collected from May to October.

Notes: Euphorbia sharkoensis seems

most closely related to E. myrtoides. It can

be distinguished from that species by its

generally smaller habit, smaller involucral

glands, seeds and capsules, longitudinally

ribbed stems, faintly irregularly ridged seed

surface and fewer staminate flowers per

cyathium.

Euphorbia sharkoensis has often been

previously identified as E. drummondii. It

differs from that species by having involucral

glands with a planar or shallowly concave

gland surface, gland appendages 0.1-0.3 mm

long, with a toothed or deeply lobed margin

and seeds with a faintly irregularly ridged

surface.

Plants from Barrow Island, Montebello

Islands and from islands of the Dampier

Archipelago (e.g. Thomson 3496, Buckley

7159, Kenneally 11610 ) differ from the typical

form by having a more woody habit, stems

that are more prominently longitudinally

ridged, and leaves that are generally thicker

and smaller.

The two collections (4-5 miles [6-8 km]

N of Yardie Creek, May 1965, George 6652

[PERTH]; Onslow, Sep 2006, Halford Q9293

[BRI]) are tentatively placed here. They

both differ from the typical form in having

gland appendages that are not as toothed or

lobed, and more prominent seed sculpturing.

These variants warrant further collecting and

investigation.

51. Euphorbia thelephora Halford &

W.K.Harris, species nova saepe cum E.

inappendiculata Domin confusa sed stylis

integris (ad vicem stylis 1/2-2/3 longitudinis

plerumque capsulis longioribus), 1.5-2 mm

longis (ad vicem capsulis 1.3-1.6 mm longis),

distincte papillosis (ad vicem capsulis tantum

aliquando minute papillosis), seminibus

plerumque longioribus 1.3-1.7 mm longis (ad

vicem seminibus 0.9-1.3 mm longis) differt.

Typus: Queensland. Burke District: 83 km S

of Normanton on road to Cloncurry, 1 August

2011, D.Halford QM515 (holo: BRI; iso: AD,

DNA, MEL, MICH, NSW, P, distribuendi ).

Monoecious or dioecious, herbaceous

perennial to 30 cm high, few to many annual

stems arising from woody rootstock. Stems

prostrate, decumbent or erect, sparingly to

much branched, smooth or papillose, with a

sparse to dense indumentum (rarely glabrous);

hairs spreading, ± straight, 0.1-1.5 mm long,

white. Interpetiolar stipules subulate, 0.3-

1.3 mm long, bipartite, papillose, glabrous

or hairy as for stems; margin entire or

laciniate. Leaves: petiole 0.3-1 mm long,

papillose (rarely smooth), indumentum as

for stems; blade oblong, obovate, elliptic or

oblong-elliptic to rotund, 2-8 mm long, 1-6

mm wide, 1-2.4 times longer than wide;

adaxial surface green, grey-green, sometimes

with reddish tinge along margin, smooth

or papillose, glabrous or with a sparse to

dense indumentum consisting of spreading,

straight to curved hairs 0.1-0.8 mm long;

abaxial surface green or pale, papillose;

glabrous or with a indumentum as for adaxial

surface; base asymmetric with one side

cordate, the other rounded to obtuse; margin

entire or obscurely toothed distally; apex

rounded. Cyathia solitary at the nodes, often

clustered on short leafy lateral branchlets

with subtending leaves slightly smaller than

primary stem leaves; peduncles 0.2-0.5 mm

long, papillose, indumentum as for stems.

Involucres cupuliform or turbinate, 0.7-

1.4 mm long, 0.7-1.4 mm across; lobes 5,

triangular, 0.3-0.6 mm long, margin entire

or ciliate; glands 4, stipitate, patelliform,

planar to shallowly concave, or cupuliform

with distinct tangential trough and thickened

rim, transverse-oblong, orbicular or reniform

in outline, 0.1-0.3 mm long; 0.2-0.5(0.9)

mm wide, red or yellow; gland appendages

conspicuous to inconspicuous or absent,

spreading radially, transverse-oblong or

triangular, 0.05-0.4 mm long, 0.05-1.1 mm

wide, white or red, glabrous, margin entire

or lobed; bracteoles 0.6-1 mm long, adnate

for c. 1/5 of their length to involucre, free

portion divided into few to many subulate,

glabrous or hirsute segments. Staminate

562

flowers 1-13 per cyathium; pedicels 0.6-1.2

mm long; staminal filaments 0.1-0.2 mm

long. Pistillate flowers: styles (0.2)0.3—1.1

mm long, spreading, recurved distally,

smooth or papillose abaxially, glabrous or

sparsely hairy proximally, entire, the apices

terete. Capsules exserted from involucre on

pedicel to 2.3 mm long, elliptic to broad-

elliptic in lateral view, 1.5-2.2 mm long,

1.2-1.8 mm across, shallowly 3-lobate

with keels acute to obtuse, papillose rarely

smooth, glabrous or with a sparse to dense

indumentum consisting of spreading hairs to

0.5 mm long; hypogynous disc entire. Seeds

narrow-ovate or ovate, 1.2-1.7 mm long, 0.5-

0.8 mm tangentially, 0.5-0.7 mm radially,

tetraquetrous in cross section; dorsal faces ±

planar, smooth or with numerous, prominent,

narrow rounded ± transverse ridges or with a

single medial longitudinal ridge; ventral faces

planar to concave, smooth or with numerous,

prominent, narrow rounded ± transverse

ridges; exotesta thin, of even thickness over

surface or of uneven thickness and distinctly

thicker on ridges, grey-white, pale brown,

microreticulate, becoming mucilaginous

when moistened; endotesta brown to red-

brown.

Austrobaileya 8(4): 441-600 (2012)

Distribution and habitat : Euphorbia

thelephora occurs from the Barkly Tableland,

NT, eastward to Normanton, Qld and south to

Coober Pedy, SA and east to Tibooburra in far

north-western NSW.

Notes: Euphorbia thelephora has been

confused with E. inappendiculata. It differs

from that species by having styles entire

(versus styles bifid for 1/2—2/3 of their

length), generally longer capsules (1.5-2

mm long versus capsules 1.3-1.6 mm long

for E. inappendiculata) which are distinctly

papillose (versus capsules only occasionally

minutely papillose for E. inappendiculata ),

and generally longer seeds (1.3-1.7 mm

long versus 0.9-1.3 mm long for E.

inappendiculata).

Etymology : The specific epithet is from Greek

thelephorus , bearing nipple-like projections,

in reference to the papillate surface on the

capsules and leaves in the typical form of this

species.

Notes : Euphorbia thelephora varies in

attributes of its indumentum and seeds.

On this basis, three varieties are formally

recognised here and can be distinguished

using the following key.

Key to varieties of Euphorbia thelephora

1 Seeds with transverse ridges. 51c. E. thelephora var. rugosa

1. Seeds smooth or finely granulate. 2

2 Leaves and capsules glabrous or with a few scattered hairs 51a. E. thelephora var. thelephora

2. Leaves and capsules sparsely to densely hairy. 51b. E. thelephora var. australis

51a. Euphorbia thelephora var. thelephora

Euphorbia sp. Beddome Range (D.E.Albrecht

5656); Short etal. (2011: 31).

Stem glabrous or with a sparse indumentum

of spreading hairs to 0.5(1) mm long. Leaves

with adaxial and abaxial surfaces glabrous

or rarely with a few scattered spreading

hairs to 0.3 mm long. Involucres turbinate,

0.7-0.8 mm long, 0.7-0.9 mm across; glands

patelliform, planar or shallowly concave,

transverse-oblong or orbicular in outline,

0.1-0.3 mm long, 0.2-0.5 mm wide, yellow;

gland appendages inconspicuous or absent,

to 0.2 mm long, 0.05-0.2 mm wide, white.

Staminate flowers 1-4 per cyathium.

Capsules elliptic in lateral view, papillose,

glabrous or rarely with a few scattered

spreading hairs to 0.3 mm long. Seeds narrow-

ovate, dorsal and ventral faces smooth, n = 11.

Figs 20, 28C.

Additional selected specimens examined: Northern

Territory. 2 miles [c. 3.2 km] W [of] Avon Downs, Jun

1960, Chippendale 7262 (AD, BRI, DNA, MEL); 10 km

N of Brunette Downs turnoff on Borroloola Road, Sep

1978, Farrell TF901 (BRI); c. 12 miles [c. 19 km] W of

Camooweal, Jul 1974, Hassall 7448 (BRI); James River

Halford & Harris, Euphorbia section Anisophyllum in Australia

563

Fig. 20. Euphorbia thelephora var. thelephora. A. habit *0.4. B. branchlet with cyathia *8. C. leaf *8. D. papillae on

lower leaf surface x32. E. stipules xl6. F. cyathia with female flower x24. G. capsule with cyathia xl6. H. cyathial gland

with appendage, adaxial view x32.1. capsule, top view xl6. J. capsule, lateral view xl6. A, C-E from Chippendale 7262

(BRI); B, F-J from Halford Q8602 & Thomas (BRI). Del. W. Smith.

564

at crossing, Barkley Highway, Avon Downs, Jun 1968,

Nicholls 860 (NT); Beddome Range, New Crown Station,

Nov 1993, Albrecht 5656 (BRI, DNA, NT); Beddome

Range, May 1974, Latz 5232 (BRI). Queensland. Burke

District: 8 km S of Normanton, May 1976, Hassall 7634

(BRI); 16 km SE of Leichhardt Falls on the Burketown

- Donors Hill Road, Apr 1974, Pullen 8939 (BRI); c. 20

km W [of] Undilla, Burketown-Camooweal Road, May

1990, Latz 11640 (DNA); Hughenden - Mt Isa Road, c.

99 km W of Julia Creek, Apr 1975, Halliday 421 (BRI);

‘Somerville’, 50 miles [c. 80 km] NE of Maxwelton,

May 1963, Entwistle 3 (BRI). Gregory North District:

33 km N of Boulia, May 1976, Hassall 7646 (BRI); 130

km W of Winton on road to Boulia, Sep 2005, Halford

Q8602 & Thomas (BRI); 5 km WNW of ‘Red Hill’,

Jun 1978, Purdie 1228 (BRI); 27 km WNW of ‘Marion

Downs’, Sep 1978, Purdie 1436 (BRI); 124 km ESE of

Bedourie, May 1976, Hassall 7656 (BRI); Diamantina

N.P., N boundary of park, Apr 1997, Forster PIF20755 &

Holland (BRI). Gregory South District: 18 miles [c. 29

km] NE of Betoota, May 1975, Hassall 7519 (AD, BRI);

70 km NW of Monkira homestead. May 1985, Neldner

1870 & Stanley (BRI). South Australia. Pandie Pandie

Station, gibber plain 3.5 km S of Lake Moorayepe, June

1983, Alexander 2277 (AD); Pedirka, Aug 1932, Ising

3012 (AD); 5 km S of Copper Hill Station homestead,

Aug 1992, Badman 5972 (AD).

Distribution and habitat : Euphorbia

thelephora var. thelephora occurs from

the Barkly Tableland, NT, eastward to

Normanton, Qld, and south to Coober Pedy,

SA and into north-western NSW (Map 51a).

It commonly grows in Dichanthium/Astrebla

grassland, Atalaya and Grevillea woodland or

Acacia cambagei R.T.Baker tall shrubland, on

stony clay or cracking clay soils on undulating

plains. The variety also inhabits Eucalyptus

camaldulensis Dehnh. low open woodland

along drainage lines, open grassland and

woodland on the slopes of small mesas or low

stony hills.

51b. Euphorbia thelephora var. australis

Halford & W.K.Harris, varietas nova ab

E. thelephora var. thelephora indumento

caulium foliorumque pilis sparsis usque

densis (ad vicem glabris vel pilis paucis

dispersis), floribus staminibus 5-13 in quoque

cyathio (ad vicem floribus staminibus 1-4

in quoque cyathio) necnon ab E. thelephora

var. rugosa seminis superficie laevi (ad

vicem superficie porcis ± transversis multis

prominentibus angustis rotundatis praedito)

differt. Typus: Queensland. Gregory North

District: c. 20 km E along Donahue Highway

from Qld/NT border, W of Boulia, 15 October

Austrobaileya 8(4): 441-600 (2012)

2011, D.Halford QM604 (holo: BRI, iso: AD,

DNA, MICH, NSW, distribuendi).

Stem with a sparse to dense indumentum

of spreading hairs to 0.8 mm long. Leaves:

adaxial and abaxial surfaces with a sparse to

dense indumentum of spreading hairs to 0.8

mm long. Involucres turbinate, cupuliform,

0.8-1.4 mm long, 1-1.4 mm across; glands

patelliform, planar to shallowly concave or

rarely cupuliform with distinct tangential

trough, transverse-oblong or reniform in

outline, 0.1-0.3 mm long; 0.3-0.5(0.9) mm

wide, red or yellow; gland appendages

conspicuous or absent, transverse-oblong,

0.1-0.4 mm long, 0.4-0.5(1.1) mm wide, white

or red. Staminate flowers 5-13 per cyathium.

Capsules elliptic to broad-elliptic in lateral

view, papillose (rarely smooth), densely hairy

with weakly spreading hairs 0.3-0.5 mm

long. Seeds narrow-ovate to ovate, dorsal and

ventral faces smooth. Fig. 28D.

Additional selected specimens examined: Northern

Territory. Horseshoe Bend Road, 5 km N of Lilia Creek,

Aug 1988, Barritt 776 (DNA [mixed collection with E.

thelephora var. rugosa]). Queensland. Burke District:

Silver Hills, 15 km N of Richmond, Apr 1977, Farrell

TF807 (BRI). Gregory North District: Roxborough

Downs, Jan 1896, Bailey s.n. (BRI [AQ202490]) [mixed

collection with E. thelephora var. rugosa ]. South

Australia. 14 km E of Roxby Downs homestead, Apr

1989, Badman 2032 (AD); 8 km SE of Netting Dam,

Arm of Lake Torrens, Andamooka Station, Jul 1989,

Badman 3247 (AD); 1 km S of homestead, Muloorina

Station, Apr 1996, Badman 8903 (AD); c. 8 km NE of

Koonamore, Curnamona Road, Apr 1968, Barker 351

(AD); Borefield Road, Apr 1997, Bates 46870 (AD);

Waukaringa Gold Mine, 30 km N of Yunta, Oct 1968,

Carrick 1724 (AD); Beresford Hill, Oct 1978, Chorney

992 (AD [mixed collection with E. centralis], NSW);

Bongadillina Creek, Warrina, 130 km N of William

Creek, Jun 1985, Conrick 1801 (AD); c. 10 km SSW of

Mt Gunson Copper Mines W of Pernatty Lagoon, Oct

1966, Eichler 18833 (AD); picnic ground c. 6 miles (c.

9 km) W of Marree, Jul 1955, Hill 100 (AD); Evelyn

Downs, Oct 1955, Ising s.n. (AD 966150240); c. 22 km of

Stuart Creek crossing (near Blower Waterhole) on track

to Coward Springs from Stuart Creek homestead. Mar

1984, Jackson 5147 (AD); Mt Lyndhurst, Oct 1898, Koch

245 (AD); Carriewerloo Station, 3 km E of homestead,

Jun 1992, Michael 353 (AD); Carriewerloo Station, 1

km NW of SE corner of Horseshoe paddock. Mar 1993,

Michael 536 (AD); c. 2 km NE of Gordon, c. 16 km SW

of Hawker, Oct 1958, Schodde 965 (AD); Dalhousie, Jun

1987, Symon 14569b New South Wales. 18.6 km N of

Tibooburra on Silver City Highway in Sturt N.P., Sep

1989, Coveny 13612 et al. (NSW).

565

Halford & Harris, Euphorbia section Anisophyllum in Australia

Distribution and habitat : Euphorbia

thelephora var. australis extends from near

Finke, NT, south east to near Yunta, SA, with

disjunct populations in western NSW and

north-western Qld (Map 51b). It grows in

herbland communities on clayey loam or clay

soils on gibber plains, in low open shrubland

communities on skeletal clay soils on rocky

calcareous/gypseous hills, or in loam or sandy

clay soils on alluvial flats or creek banks.

Phenology : Flowers and fruits have been

collected in January, March to October.

Notes : Euphorbia thelephora var. australis

differs from E. thelephora var. thelephora

by having a sparse to dense indumentum on

its stems and leaves (versus glabrous or with

a few scattered hairs for E. thelephora var.

thelephora ) and 5-13 staminate flowers per

cyathium (versus 1-4 staminate flowers for

E. thelephora var. thelephora). Euphorbia

thelephora var. australis differs from E.

thelephora var. rugosa by having a smooth

seed surface (versus seed surface with

numerous prominent narrow rounded ±

transverse ridges).

Etymology : The varietal epithet is Latin

australis , south, southern, in reference to this

variety’s distribution in relation to that of the

other varieties of this species.

51c. Euphorbia thelephora var. rugosa

Halford & W.K.Harris, varietas nova ab aliis

varietatibus E. thelephorae seminis superficie

porcis ± transversis multis prominentibus

angustis rotundatis praedita (ad vicem

superficie laevi) necnon ab E. thelephora var.

thelephora indumento caulium foliorumque

sparso usque denso (ad vicem glabro vel pilis

paucis dispersis) et floribus staminatis 7-10 in

quoque cyathio (ad vicem floribus staminatis

1-4 in quoque cyathio) differt. Typus:

Northern Territory. 5 km W along Hatt Road

from Stuart Highway, S of Alice Springs, 18

October 2011, D.Halford QM614 (holo: BRI;

iso: AD, DNA, MEL, MICH, distribuendi ).

Stem with a dense indumentum of spreading

hairs to 0.8 mm long. Leaves: adaxial and

abaxial surfaces with a dense indumentum of

spreading hairs to 0.8 mm long. Involucres

cupuliform, 0.8-1.4 mm long, 0.9-1.4 mm

across; glands cupuliform, with distinct

tangential trough and thickened rim,

transverse-oblong in outline, 0.2-0.3 mm

long; 0.3-0.5 mm wide, red; gland appendages

conspicuous, transverse-oblong, 0.2-0.3 mm

long, 0.6-1 mm wide, pink to red. Staminate

flowers 7-10 per cyathium. Capsules elliptic

in lateral view, papillose (rarely smooth),

densely hairy with weakly spreading hairs

0.3-0.5 mm long. Seeds narrow-ovate to

ovate, dorsal and ventral faces with numerous

prominent narrow rounded ± transverse

ridges, n — 11. Fig. 28E.

Additional selected specimens examined: Northern

Territory. Manners Creek homestead, Sep 1954,

Chippendale 390 (NSW); 5 km W of Stuart Highway,

road to Pine Gap, Dec 1993, Albrecht 5728 (NT); c. 4

km NW Mt Ebenezer homestead, Oct 1992, Nelson 2860

(DNA); Horseshoe Bend Road, 5 km N of Lilia Creek,

Aug 1988, Barritt 776 (DNA) [mixed collection with E.

thelephora var. australis ]. Queensland. Gregory North

District: c. 73 km S of Cloncurry, May 1976, Etas sail

7643 (BRI); Roxborough Downs, Jan 1896, Bailey s.n.

(BRI [AQ202490]) [mixed collection with E. thelephora

var. australis ]; Glenormiston, 10 miles [c. 16 km] W of

homestead, Aug 1949, Colliver s.n. (BRI [AQ202489]);

23 km NE of Old Cork homestead, near Diamantina

River, Nov 1986, Neldner 2599 & Nicolson (BRI);

Currawilla, c. 100 miles [ c. 161 km] W of Windorah,

Jun 1949, Everist 3913 (BRI); Georgina River, Sep 1910,

Bick 101 (BRI). South Australia. Dalhousie, Jun 1987,

Symon 14568 (AD); Macumba, ,s\ d. , Tate 126 (AD).

Distribution and habitat : Euphorbia

thelephora var. rugosa occurs in central

Australia from Mt Ebenezer, in southern

NT and Macumba, in northern SA, north

east to the Diamantina River, in western Qld

(Map 51c). It grows in Acacia shrubland or

woodland communities on sandy loam, clay

loam or stony clay soils on plains or low

calcareous hills.

Phenology : Flowers and fruits have been

recorded in January, May, June, and from

August to November.

Notes : Euphorbia thelephora var. rugosa

differs from the other varieties of E. thelephora

by having a seed surface with numerous

prominent narrow rounded ± transverse

ridges (versus smooth seed surface for the

other E. thelephora varieties). Euphorbia

thelephora var. rugosa also differs from E.

thelephora var. thelephora by having a sparse

to dense indumentum on its stems and leaves

566

(versus glabrous or with a few scattered

hairs for E. thelephora var. thelephora) and

7-10 staminate flowers per cyathium (versus

1-4 staminate flowers for E. thelephora var.

thelephora).

Etymology: The varietal epithet is from Latin

rugosus, wrinkled, in reference to the seed

surface texture of this variety.

52. *Euphorbia thymifolia L., Sp. PI. 1: 454

(1753); Chamaesyce thymifolia (L.) Millsp.,

Publ. Field Mus., Bot. Ser. 2(11): 412 (1916).

Type: “habitat in India” (lecto: LINN [Herb.

Linn. No. 630.10] n.v. (IDC microfiche 177.

320: III. 3), fide Wheeler [1941: 253]).

Illustration : Lin et al. (1991: 248, fig. 20), as

Chamaesyce thymifolia.

Monoecious, herbaceous perennial, with

many annual stems arising from crown

of slender taproot. Stems prostrate, much

branched, smooth, with a moderately

dense indumentum on upper surface; hairs

appressed to ascending, crispate, to 0.4 mm

long, white. Interpetiolar stipules subulate,

0.5-1.3 mm long, bipartite, indumentum as

for stems; margin laciniate. Leaves: petiole

0.8-1 mm long, smooth, glabrous or with a

few scattered hairs; blade oblong to ovate,

elliptic or oblong-elliptic, 5-8 mm long,

2.5-5 mm wide, 1.6-1.8 times longer than

wide; adaxial surface green to blue green,

occasionally red, smooth, glabrous; abaxial

surface pale green, smooth, with a sparse to

moderately dense indumentum consisting of

appressed-ascending, crispate hairs 0.3-0.4

mm long; base asymmetric with one side

rounded to shallowly cordate, the other obtuse

to rounded; margin serrulate, sometimes only

distally; apex rounded. Cyathia solitary at

the nodes, but often clustered on short leafy

axillary branchlets with subtending leaves

usually much smaller than primary stem

leaves; peduncles 0.2-0.5 mm long, smooth,

glabrous. Involucres turbinate, 0.5-0.8 mm

long, c. 0.5 mm across; lobes 5, triangular,

c. 0.2 mm long, margin fimbriate; glands

4, stipitate, patelliform, shallowly concave,

transverse-oblong to transverse-elliptic in

outline, c. 0.1 mm long, 0.2-0.3 mm wide, red;

gland appendages inconspicuous, spreading

Austrobaileya 8(4): 441-600 (2012)

radially, transverse-oblong or obdeltoid,

c. 0.1 mm long and 0.2 mm wide, white to

pink, glabrous, margin entire or shallowly

lobed; bracteoles 0.5-0.6 mm long, adnate

for c. 1/5 of their length to involucre, free

portion divided into ± linear hairy segments.

Staminate flowers 3-5 per cyathium; pedicels

0.7-0.8 mm long; staminal filaments c. 0.1

mm long. Pistillate flowers: styles 0.4-0.7

mm long, spreading, smooth, glabrous, each

bifid for 1/2—2/3 of their length, the apices

terete. Capsules at maturity on pedicel 0.5-

0.6 mm long, not completely exserted from

involucre, rupturing side of involucre, broad

to very broad-ovate in lateral view, c. 1 mm

long, 1-1.1 mm across, shallowly 3-lobate

with keels acute, smooth, with a moderately

dense indumentum; hairs appressed, 0.1-0.2

mm long; hypogynous disc entire. Seeds

ovate in outline, 0.6-0.8 mm long, 0.4-0.5

mm tangentially, 0.4-0.5 mm radially,

tetraquetrous in cross section; dorsal and

ventral surfaces concave with 3 or 4 distinct,

narrow, rounded, transverse ridges; exotesta

thin, of even thickness over surface, white,

microreticulate, becoming mucilaginous

when moistened; endotesta pale brown. Fig.

28F.

Additional selected specimens examined: Western

Australia. Broome, May 1991, Thomson 3481 (NT).

Northern Territory. Palmerston, Driver High School,

Jan 1995, Cowie 5179 (BRI, MEL); Howard Springs

Nature Park, Jun 1984, Rankin 2961 (DNA). Queensland.

Cook District: Injinoo, Mar 1993, Waterhouse

BMW2813 (BRI); Heathlands, Mar 1992, Johnson 5199

& Sharpe (BRI); Beames Street, Mareeba, Apr 1983,

Clarkson 4597 (BRI); Cowley Beach, Feb 1978, Hopkins

1511 & Graham (BRI). Burke District: Gregory River

crossing, Riversleigh Station, May 1990, Latz 11638

(DNA). North Kennedy District: Townsville, Feb

1980, Stanley 8051 (BRI); Bowen, Delta Horticultural

Research Station, Jul 1980, Swarbrick WNA28 (BRI).

South Kennedy District: edge of Eungella Dam, W of

Eungella, Feb 2003, Bean 20068 & Champion (BRI).

Port Curtis District: Mt Morgan Railway Station, Nov

2004, Batianoff 0411545 & Halford (BRI). Moreton

District: 2 Orion Court, Rothwell, northern suburb of

Brisbane, Mar 2007, Austin s.n. (BRI [AQ790414]).

Distribution and habitat: Euphorbia

thymifolia is most likely native to the New

World, but has become widely naturalised

in the Old World tropics. In Australia it has

become naturalised in the coastal tropical

and subtropical regions of Qld with isolated

567

Halford & Harris, Euphorbia section Anisophyllum in Australia

occurrences in Broome, WA, the Darwin

region, NT and Riversleigh Station, north¬

western Qld (Map 52). It grows mostly in

sunny situations on disturbed ground, lawns,

garden beds, and is frequently recorded

growing along pathways.

Phenology : Flowers and fruits have been

recorded from November to August.

Notes : Euphorbia thymifolia may be confused

with another two introduced weedy species, E.

maculata and E. prostrata. It is distinguished

from both of these species in having generally

smaller capsules and seeds, longer and more

divided style limbs, and the capsules on short

pedicels at maturity causing the involucre to

split.

53. Euphorbia trigonosperma Halford

& W.K.Harris, species nova similis E.

biconvexae Domin et E. coghlanii F.M.Bailey

sed seminibus trigonis transversal iter

(ad vicem seminibus lenticularibus in E.

biconvexa et suborbicularibus in E. coghlanii )

differt. Typus: Queensland. Cook District:

eastern flank of Newcastle Range, W of Mount

Surprise, 27 December 2006, K.R.McDonald

KRM6016 (holo: BRI).

Euphorbia mitchelliana var. glauca Benth.,

FI. Austral. 6: 47 (1873). Type: [Western

Australia.] Nickol Bay, [in 1861,] Gregory

Expedition [P. Walcott s.n .] (lecto [here

designated]: MEL 1560383).

Euphorbia sp. Hale River (B.G. Thomson

3395); Short etal. (2011:31).

Monoecious, herbaceous perennial to 50

(100) cm high, few stems arising from

slender woody taproot. Stems mostly erect

or sometimes decumbent to ascending,

sparingly to much branched, smooth, with a

sparse to moderately dense indumentum or

glabrous; hairs spreading, ± straight, 0.2-

0.5 mm long, white. Interpetiolar stipules

narrow-triangular, 0.8-1.4 mm long, entire,

bifid or bipartite, glabrous; margin entire.

Leaves: petiole 0.8-1.2 mm long, smooth,

glabrous; blade narrow-oblong or ovate,

sometimes falcate, 12-30 mm long, 4-13

mm wide, 1.6-4.2 times longer than wide;

adaxial surface light to dark green or grey

green, smooth (rarely minutely papillose),

glabrous; abaxial surface paler than adaxial

surface, smooth (rarely minutely papillose),

glabrous or (rarely with scattered spreading

hairs c. 1 mm long); base asymmetric

or symmetric with one side cordate, the

other rounded or cordate; margin entire or

sparingly minutely or coarsely toothed; apex

rounded, obtuse or acute. Cyathia solitary at

the nodes, but mostly clustered in congested

monochasial or dichasial cymes on shortened

axillary branchlets with subtending leaves

much smaller primary stem leaves; cyathial

peduncles 1-10 mm long. Involucres

campanulate or turbinate, 1-1.8 mm long, 0.9-

2 mm across; lobes 5, triangular to subulate,

0.4-1.1 mm long, margin laciniate; glands

4, stipitate, patelliform, planar, transverse-

oblong or orbicular in outline, 0.2-0.4 mm

long, 0.2-1 mm wide, green or yellowish

green; gland appendages conspicuous (rarely

absent), spreading radially, transverse-elliptic

or very broad-obovate, 0.1-1.6 mm long, 0.4-

2.3 mm wide, white, pink or yellowish-white,

glabrous, margin entire; bracteoles 1-1.5 mm

long, adnate for 1/2-2/3 of their length to

involucre, free portion divided into numerous

subulate glabrous segments. Staminate

flowers 2-32 per cyathium; pedicels 0.9-2.2

mm long; staminal filaments 0.3-0.6 mm

long. Pistillate flowers: styles 0.8-1.5 mm

long, connate at the base into a column for up

to 1/5 of their length, spreading, sometimes

recurved distally, smooth, glabrous, each

bifid for 1/2-2/3 of their length, the apices

terete. Capsules exserted from involucre on

pedicel to 4 mm long, elliptic in lateral view,

1.5-2.1 mm long, 1.9-2.6 mm across, deeply

3-lobate with keels obtuse, smooth, glabrous;

hypogynous disc entire. Seeds ovate or

elliptic in outline, (1)1.2-1.6 mm long, 0.6-1

mm tangentially, 0.8-1 mm radially, trigonal

in cross section; dorsal faces convex, smooth;

ventral faces planar or convex, smooth;

exotesta thin, of even thickness over surface,

pale brown, microreticulate, becoming

mucilaginous when moistened; endotesta

dark brown. Figs 21, 28G.

568

Additional selected specimens examined : Western

Australia, block 16 along track next to main channel,

Kununurra, Mar 1978, Aplin 6315 (BRI, PERTH); 6 km

SW of Mt Chalmers and 26 km from Dog Chain Creek

on road to Derby, Jun 1988, Wilson 12931 (PERTH);

86 km NE of Sandfire Roadhouse, Great Northern

Highway, Sep 1978, Beauglehole ACB59231 & Errey

(PERTH); Munda Creek, Mundabullangana Station,

W of Port Hedland, Feb 1962, George 3369 (PERTH);

Gregory Gorge, W Midstream, May 1976, Keighery

738 (PERTH); Hamersley Range N.P, first hill on east

side of road to Hancock Gorge from Mt Bruce - Mt

Tom Price Road, May 1980, Trudgen 2688 (PERTH);

33 miles [c. 53 km] N of Carnarvon on road to Quobba,

Feb 1962, George 3255 (PERTH). Northern Territory.

Gregory N.P, Feb 1992, Cowie 2212 & Brocklehurst

(MEL); Gold Creek, Wollogorang, Jan 1989, Russell-

Smith 6890 & Lucas (DNA); Upper Robinson River, Jan

1989, Thomson 3060 (BRI, DNA); Birrindudu, Jun 1994,

Egan 3776 (DNA); 26 km NW Barkly homestead, Apr

1993, Parsons 388 (DNA); 13 km SSE [of] Sangsters

Bore, Tanami Desert, Jun 1991, Latz 11958 (MEL);

Amarata Waterhole, Hale River, Nov 1989, Thomson

3395 (DNA). Queensland. Burke District: 34.6 km SW

of Hells Gate Roadhouse, turn off Lagoons, Apr 2006,

Thompson WES292 & Edginton (BRI); Lawn Hill N.P,

Musselbrook section, Murray Springs, Apr 2003, Booth

3205 & Kelman (BRI); E of Jump-up, 33.6 km (by road)

E of Musselbrook Mining Camp, May 1995, Johnson

MRS1028 & Thomas (BRI); 30 miles [c. 48 km] SE of

Riversleigh Station, Jun 1948, Perry 1428 (DNA); 11 km

N and 2 km W of Mt Isa, Jun 1983, Schmid 654 (BRI);

c. 7 km NW of Croydon, just W of Welcome Creek,

Dec 1998, Wannan 980 (BRI); 14.9 km by road from

Georgetown towards Forsyth, Feb 2006, McDonald

KRM4841 (BRI). South Australia. Christmas Creek

near Crispe Bore, May 1987, Symon 14390 (AD).

Distribution and habitat : Euphorbia

trigonosperma occurs across northern

Australia from the Pilbara and Kimberley,

WA, extending through the northern central

area of NT, and eastward to north-western

Qld, with outlying occurrences in southern

NT and far northern SA (Map 53). It

grows in grassland, shrubland or woodland

communities, in sandy soils on coastal or

inland dunes, in well-drained sandy soils on

stony slopes of sandstone or limestone hills,

and in sandy alluvium along drainage lines.

Phenology : Flowers and fruits have been

collected throughout the year, particularly

from January to September.

Notes: Euphorbia trigonosperma seems

most closely related to E. biconvexa and

E. coghlanii. It differs from both species in

having seeds that are trigonous in transverse

section (versus lenticulate for E. biconvexa

Austrobaileya 8(4): 441-600 (2012)

and suborbicular for E. coghlanii ), larger

involucral glands (0.2-0.4 x 0.2-1 mm versus

O. 1-0.2 x 0.1-0.6 mm for E. biconvexa and E.

coghlanii) and longer styles (0.8-1.5 mm long

versus 0.4-0.8 mm for E. biconvexa and E.

coghlanii).

This species as circumscribed here varies

considerably in the shape and thickness of

its leaf blades, number of staminate flowers

per cyathia and size of its gland appendages.

Further collections and field studies are

warranted to establish the significance of this

variation.

54. Euphorbia vaccaria Baill, Adansonia

6: 286 (1866). Type: [Western Australia.]

Hierson Island, Nickol [Nichol] Bay, s.d.,

P. Walcott s.n. (lecto: MEL 1551017, fide

Thomson [1992: 354]; isolecto: P 313104).

Monoecious, herbaceous perennial to 2 cm

high, with many annual stems arising from

a thickened woody taproot. Stems prostrate,

sparingly to much branched, smooth, with

a sparse to moderately dense indumentum;

hairs spreading, ± straight, 0.8-1.8 mm

long, white. Interpetiolar stipules narrow-

triangular, 0.7-0.8 mm long, deeply bipartite,

with indumentum as for stems; margin

laciniate. Leaves: petiole 1-1.5 mm long,

smooth, with indumentum as for stems; blade

oblong, oblong-elliptic, ovate or obovate,

4-9 mm long, 1.5-4 mm wide, 1.2-2.5

times longer than wide; adaxial and abaixal

surfaces pale green to green, smooth, with

moderately dense to dense indumentum

consisting of ascending to spreading, straight

hairs 0.3-1 mm long; base asymmetric with

one side truncate, cordate to obtuse, the other

cuneate to obtuse or rounded; margin entire

or minutely toothed distally; apex obtuse to

rounded. Cyathia solitary at the nodes, often

clustered on short leafy lateral branchlets

with subtending leaves slightly smaller than

primary stem leaves; peduncles 0.2-1.2 mm

long, smooth, hairy. Involucres campanulate,

1.3-2 mm long, 0.8-1.4 mm across; lobes 5,

triangular, 0.4-0.5 mm long, margin entire;

glands 4, stipitate, cupuliform, with tangential

trough, transverse-oblong in outline, 0.1-0.2

mm long, 0.2-0.4 mm wide, dark red; gland

appendages conspicuous, spreading radially,

Halford & Harris, Euphorbia section Anisophyllum in Australia

569

Fig. 21. Euphorbia trigonosperma. A. habit *0.4. B. branchlet with cyathia x4. C. leaf *2. D. stipules x8. E. cyathia

with female flower xi2. F. cyathia with immature capsule x8. G. cyathial gland with appendage, adaxial view xl2. H.

capsule, top view xl2.1. capsule, lateral view xl2. A from McDonaldKRM6016 (BRI); B-I from Thompson WES292

& Edginton (BRI). Del. W. Smith.

570

obdeltoid, 0.3-0.5 mm long, 0.6-1 mm wide,

pink, red or white, glabrous, margin toothed or

irregularly lobed; bracteoles 0.6-0.8 mm long,

adnate for c. 1/3 of their length to involucre,

free portion divided into ± linear hairy or

glabrous segments. Staminate flowers 3-5

per cyathium; pedicels 1.2-1.6 mm long;

staminal filaments 0.1-0.2 mm long. Pistillate

flowers: styles 0.3-0.5 mm long, ascending

and spreading distally, minutely papillose,

glabrous, each bifid for 1/3—1/2 of their length,

the apices terete. Capsules exserted from

involucre on pedicel to 3 mm long, broad to

very broad-ovate in lateral view, 1.3-1.7 mm

long, 1.2-1.9 mm across, shallowly 3-lobate

with keels obtuse, smooth or minutely

papillose, with a dense indumentum mostly

confined to the proximal half of the capsule;

hairs spreading, to 1.5 mm long; hypogynous

disc entire. Seeds ovate in outline, 0.8-1.2

mm long, 0.5-0.9 mm tangentially, 0.5-0.8

mm radially, tetraquetrous in cross section;

dorsal faces ± planar or slightly concave, with

faint narrow rounded irregular ridges; ventral

faces concave, ± smooth; exotesta thin, of

even thickness over surface, grey-white

Austrobaileya 8(4): 441-600 (2012)

or pale brown, microreticulate, becoming

mucilaginous when moistened; endotesta pale

brown or reddish brown.

Distribution and habitat : Euphorbia

vaccaria occurs from near Onslow, WA east

to Victoria River Crossing and the Tanami

Desert, NT.

Notes: Euphorbia vaccaria is similar to

and often grows with E. australis var.

subtomentosa where their distributions

overlap. Euphorbia vaccaria differs from E.

australis by the indumentum of the capsules

being confined to the proximal half of the

capsule and consisting of hairs 0.8-1.5 mm

long (versus hairs evenly disturbed over the

capsule surface and consisting of hairs up to

0.7 mm long for E. australis).

Euphorbia vaccaria exhibits some

discontinuous variation in indumentum

and leaf characters with little geographical

discontinuity. This variation is considered

sufficient to warrant formal recognition of

two varieties within this species which can be

distinguished using the following key.

Key to varieties of Euphorbia vaccaria

Leaves subtending cyathia on short lateral branchlets entire or minutely

toothed; indumentum 0.8-1 mm long.54a.E. vaccaria var. vaccaria

Leaves subtending cyathia on short lateral branchlets distinctly toothed

distally; indumentum 1-1.8 mm long.54b. E. vaccaria var. erucoides

54a. Euphorbia vaccaria var. vaccaria

Stems prostrate, with a moderately dense

indumentum; hairs 0.8-1 mm long. Leaf

blades oblong, ovate or obovate, 4-8 mm

long, 2-4 mm wide, 1.6-2.5 times longer

than wide; adaxial and abaxial surfaces with

a moderately dense indumentum of hairs

0.3-1 mm long; base asymmetric with one

side truncate to shallowly cordate, the other

rounded; margin entire or minutely toothed

distally; apex obtuse to rounded. Involucral

gland appendages 0.1-0.4 mm long, c. 0.5

mm wide, pink, red or white, margin toothed

or irregularly lobed. Pistillate flowers with

styles 0.3-0.4 mm long. Capsules exserted

from involucre on pedicel to 2.5 mm long,

1.3-1.7 mm long, 1.2-1.7 mm across, hairs to

0.8 mm long. Fig. 28H.

Additional selected specimens examined : Western

Australia. Cape Boileau, 35 km N of Broome, May

1991, Thomson 3484 (DNA); 436.3 km NE of Port

Hedland towards Broome, Sep 1995, Lolly TRL711

(BRI); 258 km SW of La Grange Mission turnoff. Great

Northern Highway, Aug 1974, Beauglehole ACB48293

& Carr (PERTH); Nalgi Station, Jul 1941, Burbidge

1341 (PERTH); Anna Plains, Jul 1941, Burbidge 1477

(PERTH); Great Sandy Desert, May 1984, Fatchen

849 (AD); 44 km (by road) along North West Coastal

Highway from Roebourne towards Whim Creek, Sep

2006, Halford Q9150 (BRI); Chichester Range, c. 13 km

N of Roebourne - Wittenoom Road, N of Mt Florence,

Sep 2006, Halford Q9269 (BRI); 49.4 km N (by road) of

Auski Roadhouse along Great Northern Highway, Sep

571

Halford & Harris, Euphorbia section Anisophyllum in Australia

2006, Halford Q9181a (BRI); SW corner of Cane River

Station, 80 km SE of Onslow, May 1999, Edinger 1455

(PERTH); 2 km NE of North West Coastal Highway,

in southern section of Cane River Station, May 1999,

Edinger 1665 (PERTH); 0.7 km SSW of Packsaddle Hill,

Hamersley Ranges, Sep 1997, Trudgen 16607 (PERTH);

31.3 km S of Nullagine on Nullagine - Newman Road,

Aug 2004, Harris WKH2235 (BRI); Patience Well,

Gibson Desert, Jun 2001, Campbell 2489 (PERTH);

Bungle Bungle N.P, Ord River at Blue Holes, Jul 1989,

Menkhorst 695 (DNA). Northern Territory. 28.9 km

E of Victoria River Crossing, Gregory N.P, Jun 2000,

Kerrigan 213 & Risler (DNA); 5 miles [c. 8 km] SSE [of]

Mongrel Downs homestead, Aug 1970, Latz 750 (AD,

DNA); 45 miles [c. 72 km] SW [of] Mongrel Downs

homestead, Aug 1970, Parker 283 (DNA, MEL, NSW);

46 miles \c. 74 km] E [of] Mongrel Downs homestead,

Apr 1971, Dunlop 2108 (DNA); Lake Mackay, Oct 1992,

Latz 12845 (MEL).

Distribution and habitat : Euphorbia

vaccaria var. vaccaria is found from near

Onslow, WA east to Victoria River Crossing

and the Tanami Desert, NT (Map 54a). It

grows in hummock grassland, Eucalyptus!

Acacia shrubland/woodland communities

on sandy loam to sandy clay soils on alluvial

flats, or stony plains or hills. It has also been

recorded growing on the margins of saltpans.

Phenology : Flowers and fruits have been

collected mostly from April to September.

54b. Euphorbia vaccaria var. erucoides

Halford & W.K.Harris, varietas nova ab E.

vaccaria var. vaccaria foliis subtendentibus

distaliter dentatis (ad vicem foliis integris

vel minute dentatis) 1-1.8 mm longi (ad

vicem pilis 0.8-1 mm longis), glandulae

appendicibus plerumque longioribus 0.3-0.5

mm longis (ad vicem appendicibus 0.1-0.4

mm longis) differt. Typus: Western Australia.

Fortescue River floodplain, c. 15 km N

of Auski Roadhouse, old Great Northern

Highway, 15 September 2006, D.Halford

Q9188 (holo: BRI; iso: MEL, PERTH).

Stems prostrate, with a sparse to moderately

dense indumentum; hairs 1-1.8 mm long.

Leaf blades obovate or oblong-elliptic,

4-9 mm long, 1.5-4 mm wide, 1.2-2.5

times longer than wide; adaxial and abaxial

surfaces with sparse or moderately dense

indumentum of hairs 0.5-1 mm long; base

asymmetric with one side cordate to obtuse,

the other cuneate to obtuse; margin toothed

distally; apex rounded. Involucral gland

appendages 0.3-0.5 mm long, 0.6-1 mm

wide, red, margin shallowly irregularly

lobed. Pistillate flowers with styles 0.4-0.5

mm long. Capsules exserted from involucre

on pedicel to 3 mm long, 1.4-1.5 mm long,

1.6-1.9 mm across, hairs 1-1.5 mm long.

Additional selected specimens examined : Western

Australia. Gregory Gorge, c. 20 km downstream

from Midstream, May 1976, Keighery 779 (PERTH);

Midstream, Sep 1969, Brooker 2057 (PERTH); Oakover

River, 19 km S of Two Sisters, c. 160 km SE of Shay

Gap, July 1984, Newbey 10424 (PERTH); near northern

shore of Gud Lake, Canning Stock Route, Aug 1989,

Barker 191 (PERTH); Cave Creek, 15 km NW of Mt

Brockman Station (abandoned) on homestead block,

Jul 1999, Backhouse BEM14 et al. (PERTH); Caves

Creek, Hamersley Station, c. 75 km along Mt Brockman

road W of Hamersley Station homestead, Sep 2006,

Halford Q9521 (BRI); Caves Creek, Mt Brockman road

50 km W of Hamersley Station homestead, Sep 2006,

Halford Q9256 (BRI); Hamersley Range N.P, 0.3 km

from Mindi Spring on track to Coppin Pools, May

1980, Trudgen 2615 (PERTH); Turee Creek, 1.5 km W

of Mindi Springs, Hamersley Range N.P, Jun 1975,

Trudgen 1351 (PERTH); Hamersley Range N.P, 2.3 km

from Mindi Springs on track to Mild Midi Springs, May

1980, Trudgen 2385 (PERTH); 25 km E of Karijini Drive

along service road of Hamersley Iron railway, Sep 2006,

Halford Q9208 (BRI); Roy Hid to Munjina Road, 7.2

km W of intersection with Nullagine - Newman Road,

Sep 2004, Harris WKH2238 (BRI); Turee Creek, Turee

Station, c. 40 km E of Paraburdoo, Sep 2006, Halford

Q9233B (BRI); 14.5 km W of Rhodes Ridge on Weed

Wold Creek Road, Aug 1973, Trudgen 420 (PERTH);

Mundiwindi, Great Northern Highway, Jun 1970, Briggs

3603 (NSW, PERTH); Little Sandy Desert, 26.6 km

ESE of Burranbar Pool on Savory Creek, 37.9 km NE

of Cooma Wed, 61.8 km N of Terminal Lake, 38.9 km

NNW of the Dean Hills, Apr 1997, van Leeuwen 3043

(BRI).

Distribution and habitat: Euphorbiavaccaria

var. erucoides occurs in the Chichester and

Hamersley Ranges, and extending eastward

to Guli Lake, in the Great Sandy Desert

(Map 54b). It grows in Eucalyptus!Melaleuca

riparian forest or woodland communities on

sandy loam to loam or gravelly alluvium in

dry river beds, on creek and river banks or

alluvial flats, and in hummock grassland,

mallee or mulga woodland communities on

sandy loam to sandy clay soils often with

calcrete on plains.

Phenology : Flowers and fruits have been

collected from mostly from May to September.

572

Notes : Euphorbia vaccaria var. erucoides

differs from E. vaccaria var. vaccaria in

having subtending leaves distinctly toothed

distally (versus entire or minutely toothed

for E. vaccaria var. vaccaria ), indumentum

1-1.8 mm long (versus 0.8-1 mm long for

E. vaccaria var. vaccaria) and generally

longer gland appendages, 0.3-0.5 mm long

(versus 0.1-0.4 mm long for E. vaccaria var.

vaccaria).

Etymology : The varietal epithet is Latin eruca ,

caterpillar, and - oides , like, resembling, in

reference to the colloquial name ‘caterpillar

plant’ used by ecological investigators in the

Pilbara region for this taxon. The cyathia are

clustered on short leafy lateral branchlets,

the clusters somewhat resemble a hairy

caterpillar.

55. Euphorbia verrucitesta Halford &

W.K.Harris, species nova ut videtur arete

affinis E. philochalicis Halford & W.K.Harris

sed glandulae appendicibus brevioribus

c. 0.5 mm longis (ad vicem appendicibus

0.1-0.3 mm longis), seminibus plerumque

longioribus 1.1-1.2(1.3) mm longis (ad vicem

seminibus 0.9-1.1 mm longis), seminum

superficiebus verrucis irregularibus ornatis

(ad vicem superficiebus porcis prominentibus

irregularibus ornatis) differt. Euphorbia

verrucitesta habitu generali, glandibus

involucralibus, glandulae appendicibus E.

multifariae Halford & W.K.Harris persimilis

sed differt autem capsulis minoribus 1.3-1.5 x

1.5-1.7 mm perlato-ovatis in ambitu (ad vicem

capsulis 1.5-1.9 x 1.5-2.1 mm lato-ellipticis

in ambitu), seminum superficiebus verrucis

irregularibus ornatis (ad vicem superficiebus

laevibus vel leviter irregulariter porcatis).

Ante hac eandem cum E. drummondii Boiss.

putata est sed seminum superficiebus verrucis

parvis irregularibus (ad vicem superficiebus

3-6 porcis transversis), capsulis minoribus

1.3-1.5 x 1.5-1.7 mm (ad vicem capsulis 1.6-

1.8 x 1.7-1.9 mm), seminibus lato-obovatis in

ambitu (ad vicem seminibus oblongo-ovatis in

ambitu), stylis bifidis 1/3—1/2 longitudinis (ad

vicem stylis integris vel vix bifidis) differt.

Typus: South Australia. 85 km S of Oldea, 25

September 1960, P. Wilson 1845 (holo: AD).

Austrobaileya 8(4): 441-600 (2012)

Monoecious, annual to 5 cm high, many

stems arising from slender taproot, the

whole plant glabrous. Stems prostrate, much

branched, smooth. Interpetiolar stipules

subulate or narrow-triangular, 0.4-1 mm

long, bipartite, glabrous; margin laciniate.

Leaves: petiole 0.3-0.7 mm long, smooth;

blade obovate, oblong or oblong-elliptic,

3.5- 7 mm long, 2-4.5 mm wide, 1.4-2.1

times longer than wide; adaxial and abaxial

surfaces green sometimes with reddish tinge

especially along margins, smooth or minutely

papillose; base symmetric, rounded to cuneate

or asymmetrical with one side cordate

or rounded, the other cuneate, obtuse or

rounded; margin entire or sparingly minutely

toothed distally; apex retuse or rounded.

Cyathia solitary at the nodes, peduncles

0.3-0.6 mm long. Involucres turbinate,

0.8-1.1 mm long, 0.9-1 mm across; lobes 5,

triangular, 0.3-0.5 mm long, margin entire,

fimbriate or laciniate; glands 4, stipitate,

cupuliform, with shallow central pit and

somewhat thickened rim, transverse-oblong

in outline, 0.1-0.2 mm long, 0.2-0.4 mm

wide, red; gland appendages inconspicuous

or absent, spreading radially, transverse-

linear, obdeltoid or oblong, c. 0.05 mm long,

0.1-0.2 mm wide, pink, glabrous, margin

entire; bracteoles 0.5-0.7 mm long, adnate for

c. 1/3 of their length to involucre, free portion

divided into few subulate glabrous or hirsute

segments. Staminate flowers 4 or 5 (10) per

cyathium; pedicels 0.9-1.1 mm long; staminal

filaments c. 0.1 mm long. Pistillate flowers:

styles 0.2-0.4 mm long, spreading, smooth,

glabrous, each bifid for 1/3—1/2 of their length,

the apices stout and terete. Capsules exserted

from involucre on pedicel to 2 mm long, very

broad-ovate in lateral view, 1.4-1.5 mm long,

1.5- 1.7 mm across, shallowly 3-lobate with

keels obtuse, smooth, glabrous; hypogynous

disc entire. Seeds broad-ovate in outline,

1-1.2(1.3) mm long, 0.7-0.8 mm tangentially,

0.6-0.7 mm radially, tetraquetrous in cross

section; dorsal and ventral faces planar or

shallowly concave, with prominent rounded

irregular wart-like protuberances; exotesta

thin, of even thickness over surface, white

or pale brown, microreticulate, becoming

mucilaginous when moistened; endotesta red-

brown. Figs 22, 281.

Halford & Harris, Euphorbia section Anisophyllum in Australia

573

Fig. 22. Euphorbia verrucitesta. A. leaf xl2. B. stipules xl6. C. cyathia with female flower x32. D. cyathia with fruit

xl6. E. cyathial gland with appendage, adaxial view x32. G. capsule, lateral view xl6. F. capsule, top view xl6. All

from Wilson 1845 (AD). Del. W.Smith.

Additional selected specimens examined : Western

Australia. 16 km E of Cocklebiddy, Oct 1984, Keighery

7543 (PERTH); 104 km S of Neale Junction, Great

Victoria Desert, Jul 1974, George 11932 (PERTH); 52

km W of Naretha, Jun 1966, Goodall 2762 (PERTH).

South Australia. S of Uno Range near Lake Gilles,

Jul 1992, Bates 28625 (AD); Koonamore, near ‘Bindyi’

Research Hut, Aug 1966, Orchard 55 (AD); near Shell

Hill, Dec 1983, Spooner 9115 (AD); R.Lencer Reserve,

Punthari, Mar 1997, Spooner 16298 (AD). New South

Wales. 2 km W along N boundary track from main road,

them c. 100 m S in Bluff Paddock, Nulla Nulla Station,

Nov 2000, Jobson 6586 et al. (NSW).

Distribution and habitat: Euphorbia

verrucitesta occurs from near Southern

Cross and Esperance, WA, eastward across

the Nullarbor through southern SA to far

south western NSW (Map 55). It has been

recorded growing in Acacia papyrocarpa

Benth. woodland and chenopod shrubland

on limestone plains and in Austrostipa open

grassland on sand plain and interdune swales.

It is also recorded on rocky mesas and granite

hills. The soils are sands or loams often

associated with limestone or gypsum.

Phenology: Flowers and fruits have been

collected throughout the year, but mostly in

October.

Notes: Euphorbia verrucitesta seems most

closely related to E. philochalix. It may be

distinguished from that species by its shorter

gland appendages (c. 0.5 mm long versus

0.1-0.3 mm long for E. philochalix ), generally

longer seeds (1.1-1.2(1.3) mm long versus 0.9-

1.1 mm long for E. philochalix) and irregular

wart-like protuberances on the seed surface

(versus prominent irregular rounded ridges

for E. philochalix).

Euphorbia verrucitesta is very similar

to E. multifaria in general habit, involucral

glands and gland appendages. It differs

from E. verrucitesta by its smaller capsules

1.3-1.5 x 1.5-17 mm that are very broad-

ovate in outline (versus 1.5-1.9 x 1.5-2.1 mm,

broad-elliptic in outline for E. multifaria) and

irregular wart-like protuberances on the seed

surface (versus smooth or faintly irregularly

ridged seed surface for E. multifaria).

574

Euphorbia verrucitesta has previously

been identified as E. drummondii , but differs

from that species in having small irregular

wart-like protuberances on the seed surface

(versus seed surfaces with 3-6 distinct

transverse ridges for E. drummondii ), smaller

capsules (1.3-1.5 x 1.5-17 mm versus 1.6-1.8

x 17-1.9 mm for E. drummondii ), seeds broad-

obovate in outline (versus oblong-obovate for

E. drummondii ) and styles that are bifid for

1/3—1/2 of their length (versus styles entire or

scarcely bifid for E. drummondii).

Etymology : The specific epithet is from

Latin verrucus , warts, and testa , seed coat, in

reference to the warty appearance of the seed

surface of this species.

56. Euphorbia vicina Halford & W.K.Harris,

species nova ante hac eandem falso cum

E. mitchelliana Boiss. vel E. schultzii

Benth. putata est. Cum generaliter similem

E. mitchellianae tamen cyathiis in cymis

dichasialibus congestis (in vicem cymis

laxis terminalis dichasialibus vel aliquando

monochasialibus distal iter nascentibus)

differt. Ab E. schultzii in cymis congestis

dispositis (ad vicem solitariis vel in ramulis

abbreviatis lateralibus fasciculatis), capsulis

1.4-1.6 x 1.9-2 mm lato-ellipticis a latere

visis leviter 3-lobatis carinis obtusis praeditis

(ad vicem capsulis 1.5-2.2 x 2-3 mm

depresse ovatis vel transverse ellipticis usque

lato-ellipticis a latere visis valde 3-lobatis

carinis acutis praeditis), seminibus tetragonis

in sectione transversali (ad vicem seminibus

tetraquetris) differt. Typus: Western

Australia. 5 km N [of] Theda homestead, 2

April 1991, B.G.Thomson 3467 (holo: PERTH;

iso: NT).

Monoecious, annual to 30 cm high, with

few stems arising from slender taproot.

Stems ascending to erect, sparingly to much

branched, smooth, with a sparse to moderately

dense indumentum; hairs ascending, curved,

0.2-07 mm long, white. Interpetiolar

stipules narrow-triangular, 0.6-1 mm

long, deeply bipartite, glabrous or with

indumentum as for stems; margin laciniate

with the teeth often gland-tipped. Leaves:

petiole 1-1.5 mm long, smooth, glabrous or

with indumentum as for stems; blade narrow-

Austrobaileya 8(4): 441-600 (2012)

ovate to ovate, 12-21 mm long, 2-9 mm wide,

2- 5.5 times longer than wide; adaxial and

abaxial surfaces green, smooth or minutely

papillose, with a sparse to moderately dense

indumentum consisting of ascending, curved

hairs 0.4-1.2 mm long (rarely glabrous); base

asymmetric with one side cordate, the other

rounded to obtuse; margin serrulate (rarely

entire); apex acute. Cyathia clustered in

congested dichasial cymes on short axillary

branchlets with bracts leaf-like but much

smaller than primary stem leaves; cyathial

peduncles c. 0.5(2) mm long. Involucres

turbinate to cupuliform, 0.7-1 mm long, 0.8-

1.3 mm across; lobes 5, triangular, 0.5-0.6

mm long, margin entire or laciniate; glands

4, stipitate, cupuliform, with shallow central

pit, transverse-oblong to orbicular in outline,

0.2-0.3 mm long, 0.2-0.3 mm wide, red;

gland appendages inconspicuous or absent,

spreading radially, obdeltoid, c. 0.1 mm long

and 0.1 mm wide, pink, glabrous, margin

entire; bracteoles 0.4-0.6 mm long, adnate

for c. 1/4 of their length to involucre, free

portion divided into ± linear hairy segments.

Staminate flowers 5-10 per cyathium;

pedicels 0.6-0.8 mm long; staminal filaments

0.1-0.2 mm long. Pistillate flowers: styles

0.3-0.4 (0.9) mm long, connate at the base

into a column for up to 1/5 of their length,

spreading, ± smooth, glabrous, each bifid

for 1/2-2/3 of their length, the apices terete.

Capsules exserted from involucre on pedicel

to 2 mm long, broad-elliptic in lateral view,

1.4-1.6 mm long, 1.9-2 mm across, shallowly

3- lobate with keels obtuse, papillose, with a

sparse indumentum consisting of spreading

white hairs 0.1-0.3 mm long (rarely glabrous);

hypogynous disc entire. Seeds broad-ovate

in outline, 1.1-1.3 mm long, 07-0.8 mm

tangentially, 07-0.8 mm radially, tetragonous

in cross section; dorsal faces ± convex;

ventral faces concave or planar; all faces with

faint regularly ridges or with 3 or 4 faint or

prominent, acute narrow transverse ridges;

exotesta thin, of even thickness over surface,

grey-white, microreticulate, becoming

mucilaginous when moistened; endotesta

brown. Figs 23, 28J.

Halford & Harris, Euphorbia section Anisophyllum in Australia

575

Fig. 23. Euphorbia vicina. A. habit x0.4. B. leaf x4. C. stipules x24. D. cyathia with female flower x24. E. cyathia with

capsule xl6. F. cyathial gland with appendage, adaxial view x32. G. capsule, top view xl6. H. capsule, lateral view xl6.

A from Keighery 4928 (PERTH); B-H from Fraser s.n. (PERTH 3079074). Del. W.Smith.

576

Additional specimens examined: Western Australia.

Mitchell Plateau, Apr 1982, Keighery 4928 (PERTH);

‘Black Mud Swamp’, 24 km from Amax Campsite on

road to Mitchell River Station, Jun 1976, Kenneally

5303/A (PERTH); summit of Poompangala Hill, c. 8 km

W of Kalumburu, Apr 1991, Willing 249 (PERTH); 2

km E of junction of Charnley and Calder River, Eastern

Walcott Inlet, May 1983, Milewski 207 (PERTH); Mt

Hart Station, Feb 1951, Fraser s.n. (PERTH 3079066,

3079074).

Distribution and habitat : Euphorbia vicina

is restricted to the Kimberley, WA, occurring

from the Mitchell Plateau and Kalumburu

areas south to Mt Hart Station (Map 56). It

grows in moist areas in herbland, grassland

or woodland communities on shallow sandy

soils, on lateritic mesas or sandstone plateaux.

Phenology : Flowers and fruits have been

collected from February to June.

Notes : Euphorbia vicina has been in the past

erroneously referred to E. mitchelliana or

E. schultzii. Although E. vicina does bear

a general resemblance to E. mitchelliana

it differs from that in having cyathia in

congested dichasial cymes (versus lax

terminal dichasial or sometimes becoming

monochasial distally for E. mitchelliana)

and capsules with a papillose surface (versus

smooth for E. mitchelliana).

Euphorbia vicina differs from E. schultzii

in having cyathia in congested cymes (versus

solitary or clustered on shortened leafy lateral

branchlets for E. schultzii ), capsules 1.4-1.6

x 1.9-2 mm, broad-elliptic in lateral view

and shallowly 3-lobate with keels obtuse

(versus 1.5-2.2 x 2-3 mm, depressed ovate or

transverse-elliptic to broad-elliptic in lateral

view and deeply 3-lobate with keels acute for

E. schultzii) and seeds tetragonous in cross

section (versus tetraquetrous in cross section

for E. schultzii).

We have seen very few ample collections

of this species and consequently the

morphological variation within this species

is poorly understood. The specimen Milewski

207 is tentatively placed here. However,

it has somewhat larger capsules and more

prominent transversely ridged seeds than

what are considered typical for this species.

Austrobaileya 8(4): 441-600 (2012)

Etymology : The specific epithet is from Latin

vicinus, near, neighbouring, in reference to

the species’ morphological similarity to E.

mitchelliana.

57. Euphorbia victoriensis Halford &

W.K.Harris, species nova arete similis

morphologice E. wheeleri Baill. in simili

sede crescens sed in tractu geographice

disjuncto. Habitu plerumque minori seminum

superficiebus porcis 4-6 prominentibus

angustis rotundatis (ad vicem superficiebus

foveatis vel reticulo-foveatis), capsulis

minoribus 1.8-1.9 mm longis (ad vicem

capsulis 2-2.8 mm longis) differt. Ante hac

eandem cum E. drummondii Boiss. putata

est sed seminum superficiebus porcis 4-6

prominentibus angustis rotundatis (ad

vicem porcis 3-6 transversis distinctis),

stipulis brevioribus 0.3-0.4 mm longis (ad

vicem stipulis 0.5-0.9 mm longis), glandis

involucralibus patelliformibus superficiebus

planis concavisve (ad vicem glandulis

cupuliformibus lacuna centrali distincta et

ore incrassata praeditis) stylis 0.4-0.6 mm

longis 1/3—1/2 longitudinis bifidis apicibus

teretibus ornatis (ad vicem stylis 0.2-0.3 mm

longis integris vel vix bifidis apicibus clavatis

ornatis) differt. Typus: Western Australia, c.

43 km W of Serpentine Lakes, 18 July 1972,

N. N.Donner 3943 (holo: AD).

Monoecious, annual or herbaceous perennial

with slender woody taproot, few to many

stems arising from rootstock, the whole

plant glabrous. Stems prostrate or rarely

erect, sparingly to much branched, smooth.

Interpetiolar stipules narrow-triangular,

O. 3-0.4 mm long, deeply bipartite; margin

laciniate. Leaves: petiole 0.6-1 mm long,

smooth; blade oblong or oblong-obovate,

3.6-7.2(11.5) mm long, 2.5-34(6.4) mm wide,

1.4-2.4 times longer than wide; adaxial

and abaxial surfaces green sometimes with

reddish tinge; smooth; base asymmetric with

one side cordate or obtuse, the other cuneate;

margin serrulate, sometimes only distally

(rarely entire); apex rounded. Cyathia solitary

at the distal nodes; peduncles c. 0.5 mm long,

smooth. Involucres turbinate, 0.7-0.9 mm

long, 0.9-1.1 mm across; lobes 5, triangular,

0.3-0.4 mm long, margin fimbriate; glands

577

Halford & Harris, Euphorbia section Anisophyllum in Australia

4, stipitate, patelliform, planar to shallowly

concave, transverse-oblong or reniform in

outline, 0.2-0.3 mm long, 0.4-0.7 mm wide,

yellowish green sometimes with reddish

tinge; gland appendages inconspicuous or

absent, spreading radially, transverse-linear,

to 0.1 mm long, 0.6-0.7 mm wide, white often

with reddish tinge, glabrous, margin erose;

bracteoles 0.8-1 mm long, adnate for c. 1/4 of

their length to involucre, free portion divided

into ± linear hairy segments. Staminate

flowers 8-15 per cyathium; pedicels 0.8-1

mm long; staminal filaments c. 0.1 mm

long. Pistillate flowers: styles 0.4-0.6 mm

long, spreading, recurved distally, smooth,

glabrous, each bifid for 1/3—1/2 of their length,

the apices terete. Capsules exserted from

involucre on pedicel to 2 mm long, broad-

elliptic in lateral view, 1.8-1.9 mm long,

1.9-2.1 mm across, shallowly 3-lobate with

keels obtuse, smooth; hypogynous disc entire.

Seeds ovate in outline, 1.5-1.7 mm long,

0.9-1 mm tangentially, 0.9-1 mm radially,

tetraquetrous in cross section; dorsal faces

convex; ventral faces planar; all faces with

4-6 prominent, narrow, rounded, irregular

ridges; exotesta thin, of even thickness

over surface, grey-white, microreticulate,

becoming mucilaginous when moistened;

endotesta red-brown or dark brown. Figs 24,

28K.

Additional selected specimens examined : Western

Australia. 11 miles [ c. 18 km] E of Notabilis Hill,

Gunbarrel Highway, Gibson Desert, Jul 1963, George

5364 (PERTH); 33 miles [c. 53 km] SE of Windulda

(Warburton Road), Aug 1962, George 4009 (PERTH).

Northern Territory, near Mt Connor, Jul 1958, Cleland

s.n. (AD 966060055). South Australia, near WA border

and Serpentine Lakes on road W of Emu, Jul 1979,

Williams 10671 (AD); 155 km W of Vokes Hill Junction,

Aug 1980, Alcock 8272 (AD); Anne Beaded Highway,

unnamed Conservation Park, Great Victoria Desert,

Aug 2001, Friebe SI 12 (AD); 35 km W of Vokes Hill

road junction, Jul 1979, Williams 10535 (AD); c. 52 km

W of Vokes Hill, corner on Connie Sue Highway, Aug

1980, Mowling s.n. (AD 98597043); 132 km N of Cook,

Aug 1980, Alcock 7980 (AD); 52 km N [of] Muckera

Rockhole, Oct 1987, Canty 2383 (AD); 98 km N of Cook

on the road to Vokes Corner, Aug 1980, Donner 7513

(AD).

Distribution and habitat: Euphorbia

victoriensis occurs in the Great Victoria Desert

in WA and SA. It has also been recorded from

the southern corner of the Gibson Desert, WA

and the south western corner of the NT (Map

57). It has been recorded growing in open

woodland communities on sand dunes.

Phenology : Flowers and fruits have been

collected from July to October.

Notes: Euphorbia victoriensis is

morphologically most similar to E. wheeleri

and grows in a similar habitat although

occurring in separate geographical areas.

Euphorbia victoriensis differs in having a

generally smaller habit, seeds surfaces with

4-6 prominent, narrow, rounded, irregular

ridges (versus seed surfaces foveate or

reticulate-foveate for E. wheeleri) and smaller

capsules (1.8-1.9 mm long versus 2-2.8 mm

long for E. wheeleri).

Euphorbia victoriensis has been previously

identified as E. drummondii. It differs from

that species in having seed surfaces with

4-6 prominent, narrow, rounded, irregular

ridges (versus 3-6 distinct transverse ridges

for E. drummondii ), shorter stipules (0.3-0.4

mm long versus 0.5-0.9 for E. drummondii ),

involucral glands patelliform with a planar or

concave surface (versus glands cupuliform,

with distinct central pit and thickened rim

for E. drummondii) and styles 0.4-0.6 mm

long and bifid for 1/3—1/2 of their length, with

terete apices (versus styles 0.2-0.3 mm long,

entire or scarcely bifid, with clavate apices for

E. drummondii).

Etymology: The species epithet refers to the

Great Victoria Desert, in WA and SA, where

this species occurs.

58. Euphorbia wheeleri Baill., Adansonia

6: 286-287 (1866); Chamaesyce wheeleri

(Baill.) D.C.Hassall, Aust. J. Bot. 24: 640

(1976). Type: [Queensland. Gregory South

District:] between Stokes Range and Coopers

Creek, s.d., Dr Wheeler s.n. (holo: P 313105;

iso: MEL 1520270).

Illustration: Weber (1986: 755, fig. 403C).

Monoecious (rarely dioecious), herbaceous

perennial to 50 cm high, with many stems

arising from crown of thickened woody

taproot, the whole plant glabrous. Stems

ascending to erect or occasionally prostrate to

decumbent, smooth. Interpetiolar stipules

578

Austrobaileya 8(4): 441-600 (2012)

Fig. 24. Euphorbia victoriensis. A. habit *0.6. B. branchlet with cyathia *8. C. leaf *8. D. stipules *16. E. cyathia with

female flower x32. F. cyathia with fruit xl6. G. cyathial gland with appendage, adaxial view x32. H. capsule, top view

xl6.1. capsule, lateral view xl6. A from Canty 2383 (AD); B-I from Donner 3943 (AD). Del. W.Smith.

579

Halford & Harris, Euphorbia section Anisophyllum in Australia

triangular or subulate, 0.6-1.6 mm long,

deeply bipartite; margin laciniate. Leaves:

petiole 0.3-1.4 mm long, smooth; blade

oblong-elliptic, elliptic, oblong or sometimes

obovate, 4-18 mm long, 2-8 mm wide, 1.7-

2.4 times longer than wide; adaxial surface

bright or dark green, smooth; abaxial surface

pale green, smooth; base asymmetric with one

side cordate, the other side cuneate; margin

serrulate sometimes only distally or entire;

apex rounded or retuse. Cyathia solitary at

the nodes; peduncles 0.5-2 mm long, smooth.

Involucres turbinate or cupuliform, 1-1.4 mm

long, 0.9-1.9 mm across; lobes 5, triangular,

0.4-0.5 mm long, margin fimbriate; glands

4, stipitate, patelliform, planar or shallowly

concave, transverse-oblong to ± reniform

in outline, 0.2-0.4 mm long, 0.4-0.8 mm

wide, yellow, pink or red; gland appendages

conspicuous to inconspicuous, spreading

radially, lunate, 0.1-0.4 mm long, 0.8-1 mm

wide, pink or white, glabrous, margin entire

or shallowly irregularly lobed; bracteoles 0.8-

1 mm long, adnate for c. 1/3 of their length to

involucre, free portion divided into ± linear

hairy segments. Staminate flowers 20-25 per

cyathium; pedicels 1-1.2 mm long; staminal

filaments 0.3-0.4 mm long. Pistillate

flowers: styles 0.4-0.8 mm long, ascending

to erect, spreading distally, smooth, each

bifid for 1/2—2/3 of their length, the apices

terete. Capsules exserted from involucre on

pedicel to 3 mm long, elliptic to broad-elliptic

in lateral view, 2-2.8 mm long, 1.9-2.4 mm

across, shallowly 3-lobate with keels obtuse,

smooth; hypogynous disc entire. Seeds ovate

in outline, 1.5-2.2 mm long, 0.8-1.2 mm

tangentially, 0.7-1.1 mm radially, tetragonous

in cross section; dorsal faces convex; ventral

faces ± planar; all faces foveate or reticulate -

foveate; exotesta thin, of even thickness over

surface, white, microreticulate, becoming

mucilaginous when moistened; endotesta pale

brown, n = 11. Figs 28L, M.

Additional selected specimens examined : Western

Australia, c. 10 km SW of rockhole, Wilson Cliffs,

Great Sandy Desert, May 1977, deGraaf 36 (PERTH);

60 miles [c. 96 km] NW of Giles, Aug 1962, Kuchel 280

(AD). Northern Territory. Lake Surprise area. Mar

1973, Henry 621 (DNA); False Mt Russell, 78 miles [c.

125 km] WSW [of] The Granites, Aug 1970, Latz 716

(AD, MEL); Finke River Railway Xing [crossing], Nov

1993, Albrecht 5531 (DNA, NT); Maryvale Station,

junction of Blackhill Bore and Maryvale Road, Aug

1988, Barritt 746 (DNA); 1.5 km SSW [of] Indracowra

homestead, Aug 2000, Albrecht 9376 (NT); Amerada

Petroleum Corp. No. 1 Hale River, Nov 1966, Symon

4376 (AD). Queensland. Gregory North District:

Monkira Station towards Cluny Station, Sep 1969,

Gittins 1952 (BRI, NSW); 164 km from Windorah on

Bedourie Road, Apr 1997, Forster PIF20607 & Holland

(BRI, MEL). Gregroy South District: 10 miles [c. 16

km] E of Birdsville, May 1975, Hassall 7523 (BRI); 40

miles [ c. 64 km] W of Windorah, May 1975, Hassall

7527 (BRI); Mt Howitt Station, 128 km W of Eromanga,

Jul 1936, Blake 11921 (AD, BRI); 83 km by road NW

of Quilpie on the road to Windorah, Mar 2001, Thomas

1936 & Fechner (BRI). South Australia. Simpson

Desert, Aug 1991, Jessop 124 (AD, BRI); Birdsville

Track, Goyder Lagoon, Aug 1975, Jackson 2733 (AD);

Oodnadatta, Dec 1977, Knight 212 (AD); Mungeranie

homestead, Sep 1956, Cleland s.n. (AD 96806425); 5

km SSW of Olympic Dam Mine, May 1989, Badman

2588 (AD). New South Wales. 2.7 km ESE of Cameron

Corner, Oct 1986, Rodd 5773 et al. (AD, NSW).

Distribution and habitat: Euphorbia wheeleri

is widespread in central Australia from the

Great Sandy and Gibson Deserts, in WA,

through southern NT and northern-east SA to

south-western Qld and far north-western NSW

(Map 58). It grows primarily in open shrubland

or Zygochloa grassland communities on deep

red sands or sandy loams on desert dunes, also

recorded on interdune flats, sandy river banks,

and in Triodia grassland on deep red sand on

open sandplains.

Phenology: Flowers and fruits have been

collected throughout the year, particularly

from May to October.

Notes: Euphorbia wheeleri is morphologically

most similar to E. myrtoides and E. victoriensis.

In addition to its geographic separation for

those species, it is easily distinguished from

both species by its foveate to reticulate-foveate

seed surface.

As accepted here, Euphorbia wheeleri

shows some variation in the seed shape and

seed surface texture. The typical form occurs

across the species range, and has seeds that

are generally slender (1.6-2.2 x 0.8-1 x 0.7-

0.9 mm) with a foveate surface (Fig. 26M).

A second less common form is found only

in the region from near Finke, NT, south to

Arcoona, SA. The seeds are generally shorter

and broader (1.5-1.9 x 1 . 1-1.2 x 1 — 1.1 mm) and

have a reticulate-foveate surface (Fig. 26N).

Representative specimens of this second

580

form are: Northern Territory. Bundooma,

Jun 1935, Cleland s.n. (AD 97405252); Uluru

N.P., 34 km WNW of Ranger Station, May

1988, Lazarides 153 & Palmer (AD, DNA);

Erldunda Station, Jun 1935, Cleland s.n.

(AD 97405248). South Australia, near Marla,

Apr 1997, Bates 47351 (AD); Pedirka, Aug

1932, Ising s.n. (AD 966031152); c. 8 km E

of Macumba homestead, Sep 1931, Ising

s.n. (AD 966030967); Moonlands Bore,

Todmorden Station, Aug 1992, Badman

5869 (AD 99326003); Macumba Station,

Nov 1950, Ising s.n. (AD 966150249); off

Borefield road, near Canegrass Swamp, Apr

1997, Bates 46879 (AD); Arcoona, Sep 1927,

Murray 170 (AD). This variant may, with

further collections and research, be found to

represent a distinct taxon.

Excluded names

Euphorbia atoto G.Forst., FI. Ins. Austr. 36

(1786); Chamaesyce atoto (G.Forst.) Croizat,

in O. Degener, FI. Hawaiiensis Fam. 190.

(1936). Type: Society Islands, Tahiti, s.d.,

Herbier Forster 110 (lecto: P-Forst n.v. [image

seen]; isolecto: BM 1014903 n.v. [image seen],

G n.v., GEOT n.v. [image seen], UPS-THUNB

n.v. (IDC microfiche 1036-16. 479, II. 1 ),fide

Florence [1996: 242-243]).

As circumscribed by Florence (1996),

Euphorbia atoto is endemic to Tahiti. The

Australian specimens previously assigned to

this species are referrable to E. litticola, E.

obliqua or E. pallens.

Euphorbia chamaesyce F., Sp. PI. 1: 455

(1753). Type: “Habitat in Europa australi,

Sibiria” unknown locality, s.d., Lofling 373

(lecto: FINN [Herb. Linn. No. 630.15] n.v.

(image seen) (IDC microfiche 177. 320: I. 1),

fide Khan [1964: 152]).

In Australia, this name has been

misapplied to Euphorbia drummondii, E.

maculata, E. prostrata and E. thymifolia

(Hooker 1855-1859; Dunlop et al. 1995;

Paczkowska & Chapman 2000).

Euphorbia drummondii var. dallachyana

Baill., Adansonia 6: 285 (30 July 1866). Type

citation: “Dallachy, “Pine Plains”, cum typo

(Herb. F. Muell.!) - Murray, Cooper’s creek

(herb. F. Muell.!)”.

Austrobaileya 8(4): 441-600 (2012)

We have been unable to locate the type

material of this variety. There is a collection

in MEF (2179478) with the information

“Euphorbia drummondii, Baill. Cooper’s

Creek” hand written on a blue Phytologic

Museum of Melbourne label. There is no

indication that it was seen by Baillon or that it

is E. drummondii var. dallachyana of Baillon.

This collection is referrable to E. dallachyana.

Euphorbia drummondii var. erythropeplis

Baill., Adansonia 6: 285 (30 July 1866). Type:

Western Australia. Murchison R., [without

date,] Oldf. [Oldfield] 1082 (holo: MEF

2179193).

There is insufficient material to place this

name with any certainty. The seeds and fruits

of this taxon are unknown. The involucral

glands and gland appendages are similar to

Euphorbia drummondii, however, the leaves

are smaller and thicker, and more or less

entire.

Euphorbia bracteolaris Boiss., Cent.

Euphorb. 8 (1860). Euphorbia hypericifolia

var. bracteolaris (Boiss.) Ewart, Proc. Roy.

Soc. Victoria 19: 41 (1907). Type citation: “In

montibus Nilagiricis Indiae (Perrottet sub n.

1832 ex parte).”

This name is not applicable to any

Australian taxon. The Australian specimen

(Elder Expl. Exp. 1892 lat 27 deg. 5 mS., long.

119 deg. 15 m. E [MEF 2179930]) assigned by

Ewart ( loc. cit.) to this variety is referable to

Euphorbia coghlanii.

Euphorbia pilulifera L., Sp. PL. 454 (1753),

nom. rej., fide Esser & Cafferty (2001);

(2006). McNeill et al. (2006). Type: “Habitat

in India.” (lecto: LINN [Herb. Linn. No.

630.8] (image seen ),fide Brown et al. (1911:

497-498)).

In Australia, the name Euphorbia

pilulifera has been misapplied to the species

E. hirta (Bentham 1873; Domin 1927). The

correct name for the type of the name E.

piluliferia is E. parviflora L. This name is not

applicable to any Australian species.

581

Halford & Harris, Euphorbia section Anisophyllum in Australia

Acknowledgements

We wish to thank the Directors/Curators

of AD, BM, DNA, K, MEL, NSW, NT,

P, PERTH, PR and W for the loan of their

holdings requested by us for study at BRI, and

G and P for providing digital images of type

material. We are deeply grateful to Les Pedley

for translating the diagnoses into Latin, Will

Smith for the excellent photographs, drawings,

and maps, Paul Berry (MICH) and Ya Yang

(MICH) for their assistance in the identification/

confirmation of a number of the naturalised

species in Australia, Mike Hislop (PERTH),

Keith McDonald (formerly of Department of

Environment & Heritage Protection), David

Fell (3D Environmental) and colleagues at BRI

for collecting additional material of Euphorbia.

We thank the Australian Botanical Liaison

Officers (Peter Bostock (2001-2002), Roberta

Cowan and Alex George (2004-2005), Jeremy

Bruhl (2007-2008) and Tony Orchard (2008-

2009)) at Kew for searching and photography

of types, Stephen van Leeuwen (Department

of Environment and Conservation, WA) for

the opportunity to undertake field work in the

Pilbara region of Western Australia in 2004

and 2006. We would like to thank Dr Gordon

Guymer for making facilities and space

available at BRI for the authors. The study was

partially funded by grants from the Australian

Biological Resources Study (ABRS),

Department of Sustainability, Environment,

Water, Population and Communities, which is

gratefully acknowledged.

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Fig. 25.

Halford & Harris, Euphorbia section Anisophyllum in Australia

583

Fig. 25. Dorsal and lateral views of Euphorbia seeds: A. E. albrechtii [Chippendale 2019 (DNA)]. B. E.

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584

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Fig. 26.

Halford & Harris, Euphorbia section Anisophyllum in Australia

585

Fig. 26. Dorsal and lateral views of Euphorbia seeds: A. E.flindersica [Kuchnel 3169 (AD)]. B. E. gregoriensis

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6125 (BRI)]. L. E. macdonaldii var. macdonaldii [McDonald KRM3913 & Little (BRI)]. M. E. macdonaldii var.

potentillina [McDonald KRM3512 (BRI)]. N. E. maconochieana [Perry & Lazarides 2025 (BRI)]. O. E. maculata

[Halford Q7512 & Batianoff (BRI)]. P. E. mitchelliana var. mitchelliana [Neldner 3843 (BRI)]. Q. E. muelleri [Leach

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Fig. 27.

587

Halford & Harris, Euphorbia section Anisophyllum in Australia

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588

Austrobaileya 8(4): 441-600 (2012)

Fig. 28. Dorsal and lateral views of Euphorbia seeds: A. E. schultzii var. comans [Henshall 3481 (BRI)]. B. E. serpens

[Halford 08135 & Harris (BRI)]. C. E. thelephora var. thelephora [Neldner & Stanley 1870 (BRI)]. D. E. thelephora

var. australis [Jackson 5147 (AD)]. E. E. thelephora var. rugosa [Nelson 2860 (DNA)]. F. E. thymifolia [Clarkson 4597

(BRI)]. G. E. trigonosperma [McDonald KRM4841 (BRI)]. H. E. vaccaria var. vaccaria [Halford Q9181A (BRI)].

I. E. verrucitesta [Spooner 16298 (AD)]. J. E. vicina [Willing 249 (PERTH)]. K. E. victoriensis [Cleland s.n. (AD

966060055)]. L. E. wheeleri (typical form) [Hassall 7527 (BRI)]. M. E. wheeleri [Bates 46879 (AD)]. Scale bars = 1

mm.

589

Halford & Harris, Euphorbia section Anisophyllum in Australia

Maps 1-10.

590

Austrobaileya 8(4): 441-600 (2012)

E. ferdinandi

var. saxosiplaniticola

16C

Maps 11-23A.

591

Halford & Harris, Euphorbia section Anisophyllum in Australia

Maps 23B-33.

592

Austrobaileya 8(4): 441-600 (2012)

Maps 34-46.

593

Halford & Harris, Euphorbia section Anisophyllum in Australia

48B

E. thelephora

var. thelephora

E. schultzii

var. comans

51A

51B

E. trigonosperma

E. thelephora

var. australis

E. verrucitesta

51C

E. vaccaria

var. vaccaria

E. thelephora

var. rugosa

54A

Maps 47-57.

594

Austrobaileya 8(4): 441-600 (2012)

Map 58.

Appendix 1. List of chromosome numbers for Australian taxa of Euphorbia sect.

Anisophyllum (data from Hassall 1977 and BRI database).

Taxon

Queensland Herbarium AQ number of

voucher specimens

E. australis var. erythrantha

475107

E. australis var. subtomentosa

475110, 475112, 475113, 478151

E. biconvexa

475032, 475033, 475037, 475041, 475042,

475043, 475044, 475045, 475046

E. bifida (coastal form)

475047,475048, 475050, 475088

E. bifida (narrow leaf form)

475072, 475074, 475075

E. coghlanii

475034, 475035, 475036, 475038, 475057

E. crassimarginata

475165, 475167, 475168

E. dallachyana

475134, 475135, 475136, 475137, 475138,

475139, 475140, 475141, 475142, 475143,

475144, 475145, 475146, 475147, 475148,

475150,475151, 475152, 475153, 475154,

475155, 475156, 475157, 475170

E. dallachyana

447303, 447304, 447305, 447306, 447311,

447317, 447318, 447319, 447320,447321,

447322, 447323, 447327, 447278, 447279,

475115, 475116, 475117, 475118, 475121

E. drummondii

447280, 447300, 447301, 447307, 447308,

447309, 447310, 447312, 447325, 447326,

447328, 447330, 447331, 447332, 447333,

447334, 447335, 447337, 447339, 447345,

447346, 447347, 447348, 475085, 475087,

475163, 475164

E. ferdinandi var. ferdinandi

475097, 475098, 475099, 475100, 475101,

475103

E. ferdinandi var. appendiculata

447284, 447289, 447290, 447292

Halford & Harris, Euphorbia section Anisophyllum in Australia

595

Taxon

Queensland Herbarium AQ number of

voucher specimens

E. flindersica

475128

E. hassallii

475125, 475127

E. inappendiculata var.

447283, 447298, 447343, 475122, 475123,

queenslandica

475161

E. macdonaldii var. potentillina

475106

E. mitchelliana var. longiloba

475055, 475058

E. mitchelliana var. mitchelliana

475054, 475059, 475060, 475062, 475061,

475063, 475064,475065, 475066

E. multifaria

447287, 447288, 447295, 447296, 447313,

447314, 447315, 447336

E. papillata var. papillata

475093, 475096, 475104

E. papillifolia var. polyandra

475129

E. philochalix

475130, 475131,475132

E. porcata

447299, 447294, 475120, 447324, 447285,

447316, 447286, 447282, 447297

E. psammogeton

475049

E. schultzii

475105

E. thelephora var. rugosa

475111

E. thelephora var. thelephora

475069, 475070, 475071, 475095

E. wheeleri

475080, 475082,475083, 475084, 475077,

475078, 475079, 475240

Index to Scientific Names

Names in bold type are accepted names and those in light type are synonyms, etc. The numbers

refer to the taxa enumerated above and excl. refers to ‘Excluded names’ section.

Chamaesyce alsiniflora (Baill.) D.C.Hassall.30b

Chamaesyce atoto (G.Forst.) Croizat.excl.

Chamaesyce australis (Boiss.) D.C.Hassall.4

Chamaesyce biconvexa (Domin) D.C.Hassall.5

Chamaesyce bifida (Hook. & Arn.) T.Kuros.6

Chamaesyce carissoides (F.M.Bailey) P.I.Forst. & R. J.F.Hend.8

Chamaesyce centralis (B.G.Thomson) P.I.Forst. & R. J.F.Hend.9

Chamaesyce coghlanii (F.M.Bailey) D.C.Hassall ex P.I.Forst. & R. J.F.Hend.12

Chamaesyce dallachyana (Baill.) D.C.Hassall.14

Chamaesyce drummondii (Boiss.) Sojak.15

Chamaesyce filipes (Benth.) D.C.Hassall.30b

Chamaesyce hirta (L.) Millsp.21

Chamaesyce hyssopifolia (L.) Small.22

Chamaesyce inappendiculata (Domin) D.C.Hassall.23

596 Austrobaileya 8(4): 441-600 (2012)

Chamaesyce macgillivrayi (Boiss.) D.C.Hassall.6

Chamaesyce maculata (L.) Small.29

Chamaesyce micradenia (Boiss.) D.C.Hassall.6

Chamaesyce mitchelliana (Boiss.) D.C.Hassall.30

Chamaesyce mitchelliana var. hirta (Boiss.) D.C.Hassall.30a

Chamaesyce myrtoides (Boiss.) D.C.Hassall.33

Chamaesyce obliqua (Endl.) J.Florence.34

Chamaesyce ophiolitica P.I.Forst.36

Chamaesyce ophthalmica (Pers.) D.G.Burch.37

Chamaesycepallens (Dillwyn) V.S.Raju.38

Chamaesyce petala (Ewart & F.R.Kerr) P.I.Forst. & R. J.F.Hend.41

Chamaesyce prostrata (Aiton) Small.44

Chamaesycepsammogeton (P.S.Green) P.I.Forst. & R.J.F.Hend.45

Chamaesyce schultzii (Benth.) D.C.Hassall.48

Chamaesyce serpens (Kunth) Small.49

Chamaesyce sp. (Pathungra A.Gunness AG2118).39a

Chamaesyce sp. A.40b

Chamaesyce sp. B.23b

Chamaesyce sp. Marree (F.J.Badman 776).23b

Chamaesyce supina (Raf.) Moldenke.29

Chamaesyce thymifolia (F.) Millsp.52

Chamaesyce vachellii (Hook. & Arn.) H.Hara.6

Chamaesyce wheeleri (Baill.) D.C.Hassall.58.

Euphorbia “Marree” (F.J. Badman 776).23b

Euphorbia accedens Halford & W.K.Harris.1

Euphorbia albrechtii Halford & W.K.Harris.2

Euphorbia alsiniflora Baill.30b

Euphorbia armstrongiana Boiss.3

Euphorbia armstrongiana Boiss. var. armstrongiana .3a

Euphorbia armstrongiana var. distans (W.Fitzg.) Halford & W.K.Harris .3b

Euphorbia atoto G.Forst.excl.

Euphorbia atoto var. imbricata Boiss.45

Euphorbia australis Boiss.4

Euphorbia australis Boiss. var. australis .4a

Euphorbia australis var. canescens Domin.27a

Euphorbia australis var. erythrantha (F.Muell.) Benth.4b

Euphorbia australis var. glabra Halford & W.K.Harris.4c

Halford & Harris, Euphorbia section Anisophyllum in Australia 597

Euphorbia australis var. glaucescens Boiss.48a

Euphorbia australis var. hispidula Halford & W.K.Harris.4d

Euphorbia australis var. potentillina Baill.27b

Euphorbia australis var. semiglabra Domin.27b

Euphorbia australis var. subtomentosa Domin.4e

Euphorbia baueri Engelm. ex Boiss.6

Euphorbia biconvexa Domin.5

Euphorbia bifida Hook. & Arn.6

Euphorbia bouleyi S.Moore.7

Euphorbia bracteolaris Boiss.excl.

Euphorbia careyi F.Muell.7

Euphorbia carissoides F.M.Bailey.8

Euphorbia centralis B.G.Thomson.9

Euphorbia chamaesyce F.excl.

Euphorbia chrysochaeta W.Fitzg.21

Euph orbia cin erea W. Fitzg.10

Euphorbia clementii Domin.11

Euphorbia coghlanii F.M.Bailey.12

Euphorbia comans W.Fitzg.48b

Euphorbia crassimarginata Halford & W.K.Harris.13

Euphorbia dallachyana Baill.14

Euphorbia distans W.Fitzg.3b

Euphorbia drummondii Boiss.15

Euphorbia drummondii var. dallachyana Baill.excl.

Euphorbia drummondii Boiss. var. drummondii .15

Euphorbia drummondii var. erythropeplis Baill.excl.

Euphorbia drummondii var. rubescens Benth.50

Euphorbia erythrantha F.Muell.4b

Euphorbia ferdinandi Baill.16

Euphorbia ferdinandi var. appendiculata Halford & W.K.Harris.16b

Euphorbia ferdinandi Baill. var. ferdinandi .16a

Euphorbia ferdinandi var. saxosiplaniticola Halford & W.K.Harris.16c

Euphorbiafilipes Benth.30b

Euphorbia fitzroyensis Halford & W.K.Harris.17

Euphorbia flindersica Halford & W.K.Harris.18

Euphorbia gregoriensis Halford & W.K.Harris.19

Euphorbia hassallii Halford & W.K.Harris .20

598 Austrobaileya 8(4): 441-600 (2012)

Euphorbia hirta L.21

Euphorbia hypericifolia var. bracteolaris (Boiss.) Ewart.excl.

Euphorbia hyssopifolia L.22

Euphorbia inappendiculata Domin.23

Euphorbia inappendiculata Domin var. inappendiculata .23a

Euphorbia inappendiculata var. queenslandica Domin.23b

Euphorbia inappendiculata var. robustior Halford & W.K.Harris.23c

Euphorbia kimberleyensis B.G.Thomson.24

Euphorbia laciniloba Halford & W.K.Harris.25

Euphorbia levis var. imbricata Boiss.26

Euphorbia litticola Halford & W.K.Harris.26

Euphorbia macdonaldii Halford & W.K.Harris.27

Euphorbia macdonaldii Halford & W.K.Harris var. macdonaldii .27a

Euphorbia macdonaldii var. potentillina (Baill.) Halford & W.K.Harris.27b

Euphorbia macgillivrayi Boiss.6

Euphorbia macgillivrayi f. glabrata Domin.6

Euphorbia macgillivrayi vm.filipes (Benth.) Domin.30b

Euphorbia macgillivrayi Boiss. var. macgillivrayi .6

Euphorbia macgillivrayi var. micradenia (Boiss.) Domin.6

Euphorbia macgillivrayi var. pseudoserrulata Domin.6

Euphorbia macgillivrayi var. yarrabensis Domin.6

Euphorbia maconochieana B.G.Thomson.28

Euphorbia maculata L.29

Euphorbia micradenia Boiss.6

Euphorbia minutifolia Boiss.14

Euphorbia mitchelliana Boiss.30

Euphorbia mitchelliana var. cairnsiana Domin.30a

Euphorbia mitchelliana var. dietrichiae Domin.30a

Euphorbia mitchelliana var .filifolia Domin .30a

Euphorbia mitchelliana var. filipes (Benth.) Halford & W.K.Harris.30b

Euphorbia mitchelliana var. glauca Benth.53

Euphorbia mitchelliana var. hirta Boiss.30a

Euphorbia mitchelliana var. longiloba Halford & W.K.Harris.30c

Euphorbia mitchelliana Boiss. var. mitchelliana. .30a

Euphorbia mitchelliana var. oblongifolia Boiss.30a

Euphorbia mitchelliana var. stenophylla Benth.6

Euphorbia muelleri Boiss.31

Halford & Harris, Euphorbia section Anisophyllum in Australia 599

Euphorbia multifaria Halford & W.K.Harris.32

Euphorbia myrtoides Boiss.33

Euphorbia obliqua Endl.34

Euphorbia occulta Halford & W.K.Harris.35

Euphorbia ophiolitica (PLForst.) Y.Yang.36

Euph orbia oph th almica Pers.37

Euphorbia paliens Dillwyn.38

Euphorbiapapillata Halford & W.K.Harris.39

Euphorbia papillata var. laevicaulis Halford & W.K.Harris.39b

Euphorbia papillata Halford & W.K.Harris var. papillata .39a

Euphorbiapapillifolia Halford & W.K.Harris.40

Euphorbia papillifolia Halford & W.K.Harris var. papillifolia .40a

Euphorbia papillifolia var. polyandra Halford & W.K.Harris.40b

Euphorbiapetala Ewart & L.R.Kerr.41

Euphorbiaphilochalix Halford & W.K.Harris.42

Euphorbia pilulifera L.excl.

Euphorbia pilulifera f. humifusa Domin.21

Euphorbia pilulifera f. rubromaculata Domin.21

Euphorbia pilulifera f. viridis Domin.21

Euphorbiaporcata Halford & W.K.Harris.43

Euphorbia prostrata Aiton.44

Euph orbia psammogeton P. S Green.45

Euphorbiapsilosperma Halford & W.K.Harris .46

Euphorbiapubicaulis S.Moore.30c

Euphorbia schizolepis F.Muell. ex Boiss.47

Euphorbia schizolepis var. glabra Benth.8

Euphorbia schizolepis F.Muell. ex Boiss. var. schizolepis .47

Euphorbia schultzii Benth.48

Euphorbia schultzii var. comans (W.Fitzg.) Halford & W.K.Harris.48b

Euphorbia schultzii Benth. var. schultzii .48a

Euphorbia serpens Kunth.49

Euphorbia sharkoensis Baill.50

Euphorbia sp. Beddome Range (D.E.Albrecht 5656). 51a

Euphorbia sp. Hale River (B.G. Thomson 3395). 53

Euphorbia supinaRaf. .29

Euphorbia thelephora Halford & W.K.Harris.51

Euphorbia thelephora var. australis Halford & W.K.Harris.51b

600

Austrobaileya 8(4): 441-600 (2012)

Euphorbia thelephora var. rugosa Halford & W.K.Harris.51c

Euphorbia thelephora Halford & W.K.Harris var. thelephora .51a

Euph orbia thymifolia L.52

Euphorbia trigonosperma Halford & W.K. Harris.53

Euphorbia vaccaria Baill.54

Euphorbia vaccaria var. erucoides Halford & W.K.Harris.54b

Euphorbia vaccaria Baill. var. vaccaria .54a

Euphorbia vachellii Hook. & Arn.6

Euphorbia verrucitesta Halford & W.K.Harris .55

Euphorbia vicina Halford & W.K.Harris.56

Euphorbia victoriensis Halford & W.K.Harris.57

Euphorbia wheeled Baill.58

Utricularia corneliana R.W.Jobson (Lentibulariaceae), a new

species from the North Kennedy district of Queensland

Richard W. Jobson

Summary

Jobson, R.W. (2012). Utricularia corneliana R.W.Jobson, (Lentibulariaceae), a new species from

the North Kennedy district of Queensland. Austrobaileya 8(4): 601-607. Utricularia corneliana

R.W.Jobson, possibly endemic to the Minnamoolka area of northern Queensland, is described,

illustrated, and differentiated from the local, and closely related African and South American species.

Notes are provided on habitat and ecology, and conservation status. A key to Australian and related

suspended aquatic species of Utricularia is provided.

Key Words: Lentibulariaceae, Utricularia , Utricularia corneliana , Australia flora, Queensland flora,

new species, taxonomy, bladderwort, aquatic

R.W.Jobson, National Herbarium of New South Wales, Royal Botanic Garden and Domain Trust, Mrs

Macquaries Road, Sydney, NSW 2000, Australia. Email: r ichard.iobson@,rbgsvd.nsw.gov.au

Introduction

Utricularia L. (Lentibulariceae) is a

monophyletic genus of carnivorous

angiosperms containing at least 219

recognised species worldwide (Taylor 1989;

Gassin 1993; Lowrie 1998, 2002; Lowrie et

al. 2008; Jobson 2012), mostly distributed

in subtropical and tropical regions (Taylor

1989). In his monograph of Utricularia ,

Taylor (1989) delimited the genus into the

following two subgenera: Polypompholyx

(Lehm.) P.Taylor (three species), including

two sections ( Tridentaria P.Taylor and

Polypompholyx ), and Utricularia (211

species), including 35 sectional groupings.

In line with the results of Jobson et al. (2003)

the genus has since been divided into three

subgenera, viz. Polypompholyx , Bivalvaria

S.Kurz and Utricularia (Reut & Jobson 2010).

Australia has c. 62 species (47 endemic), from

the subgenera Polypompholyx (c. 40 species),

Bivalvaria (13 species), and Utricularia (nine

species).

Based on its suspended aquatic habit, and

morphological characters such as bladder-trap

form, the absence of bracteoles, the presence

of basifixed bracts, and dehiscence of seed

via a circumscissile suture of the capsule,

the new species described here ( Utricularia

corneliana, Figs. 1, 2A) is considered a

member of subgenus Utricularia section

Utricularia. This section consists mainly

of species with a fully suspended aquatic

habit (Taylor 1989; Jobson et al. 2003). In

the current paper, the distribution, habitat

and morphological differences between

U. corneliana and the other Australian

suspended aquatic species, U. aurea Lour.,

U. australis R.Br., U. gibba L., U. muelleri

Kamienski, and U. stellar is L.f., are discussed.

Also provided is a comparative discussion of

the tropical African U. reflexa Oliver and U.

raynalii P.Taylor, and the South American

U. warmingii Kamienski, three species that

have several characters in common with U.

corneliana.

Methods and materials

This study is based on a single collection from

a single site. The specimen was divided into

two spirit preserved (70% ethanol) accessions

that are deposited at NSW and BRI.

The author examined suspended aquatic

species of Utricularia (U. aurea , U. australis ,

U. gibba , U. muelleri and U. stellaris) that

are deposited at BRI and NSW, finding

no indication that wrongly identified U.

Accepted for publication 24 July 2012

602

corneliana had previously been collected.

Taxonomy

Utricularia corneliana R.W.Jobson, species

nova U. reflexae similis sed limbo inferiore

quam superiore majore differt. Typus:

Australia: Queensland. North Kennedy

District: S of Mt Garnet, 9 June 2011,

R.W.Jobson 1281 (holo: NSW; iso: BRI).

Small perennial, suspended aquatic herb.

Rhizoids not present. Stolons filiform 5-15

cm long, 0.3-0.5 mm thick, unbranched,

terete, sparsely hairy, internodes 6-8 mm

long. Leaves numerous, circular in outline, ±

amplexicaul, 3-5 mm long, slightly flattened,

divided at the base into 2 primary segments,

with 3 further dichotomously divided

segments, the ultimate segments apically

and laterally setulose. Traps 1 (2) per leaf,

inserted in the angle between the first and

second division segments, occasionally also

in the third, stalked, ovoid 2-2.6 mm long,

mouth lateral with two dorsal, setiform, often

recurved appendages 1-2 mm long, sometime

2 or 3 simple lateral setae. Internal glands

4-armed, narrowly cylindrical up to 90 p long,

~5 p in diameter (Fig. 2B). Inflorescence

weakly erect, emergent, 2-3 cm long, arising

along the stolon from nodes at intervals of

c. 3.5 cm. Peduncle filiform 0.5-0.6 mm

thick, terete, glandular, sparsely hairy on

lower portion, mostly glabrous above first

bract. Scales and bracteoles absent. Bracts

basifixed, amplexicaule, c. 1.3 mm long and

0.9 mm in diameter, apex rounded or truncate.

Flowers 1-3 on an elongated raceme axis;

pedicels filiform, erect at anthesis, deflexed in

fruit 3-5.5 mm long. Lowest flower probably

cleistogamous. Calyx lobes subequal, upper

lobe slightly longer, ovate 3-3.5 mm long,

2-2.2 mm in diameter. Corolla 4.5-9.3 mm

long, yellow, with few brown nerves on the

basal portion of the upper lip, densely covered

with fine multicellular hairs on dorsal surfaces;

upper lip broadly ovate with apex rounded

4-6.5 mm long, 3.8-5.5 mm in diameter, the

lower half of dorsal surface covered in hairs;

lower lip limb smaller, bilobed, with a single

prominent, slightly emarginate swelling at

the base; spur cylindrical at base, curved,

Austrobaileya 8(4): 601-607 (2012)

slightly flattened and tapering mid-way with

apex rounded, almost as long as lower lip

(when lip is flattened). Filaments curved c.

1.6 mm long. Ovary globose. Capsule 3.2-37

mm long, 2-3 mm in diameter, walls fleshy,

circumscissile dehiscence. Seeds thinly

lenticular 0.8-1 mm in diameter, with a broad,

softly angled and translucent, mildly dentate

edged, marginal wing of irregular testa cells

with raised anticlinal walls (Fig. 2C). Pollen

17-18 colporate, 30 x 30 p, Jobson 1281

(NSW). Fig. 1.

Distribution and habitat : Utricularia

corneliana is thus far, only known from a

single Swamp, south of Mt Garnet in the

Minnamoolka area. This ephemeral swamp

with a circumference of c. 4 km is fringed by

Eucalyptus platyphylla F.Muell., E. sp., and

Melaleuca nervosa (Lindl.) Cheel woodland

(Fig. 3). Plants of the bladderwort were

infrequent in a single corner of the swamp

(c. 5 x 5 m), in water to c. 20 cm deep, with

Aldrovanda vesiculosa L., aquatic grasses,

Eleocharis sp., Marsilea mutica Mett.,

Myriophyllum simulans Orchard, Nympoides

indica (L.) Kuntze, Utricularia aurea , U.

gibba and U. stellaris. This black soil swamp

is based upon a basalt and sand substrate at an

elevation of c. 700 m.

Phenology : Flowers and fruits recorded

in June. Further research is required to

determine extent of flowering season.

Notes: Utricularia corneliana is

geographically isolated and grows

sympatrically with three other suspended

aquatic Utricularia species; however,

morphologically it shares most characters

in common with U. reflexa , a variable

species endemic to tropical Africa and

Madagascar (Taylor 1989: fig. 194, p. 640).

Molecular phylogenetic data also support this

relationship (Jobson et al., in prep.) and negate

the possibility of a localised hybridisation

event. These two species share a bright yellow

corolla, similarly shaped bracts, bilobed

lower corolla lip, and have traps invariably

positioned at the angle between leaf segments

(Fig. 1)

Jobson, Utricularia comeliana

603

There are two, perhaps closely allied

species, that also have traps positioned in

the angle between leaf segments; Utricularia

raynalii (tropical Africa) (Taylor 1989:

fig. 195, p. 642), and U. warmingii (South

America) (Taylor 1989: fig. 196, p. 644).

These two species differ from U. reflexa and

U. corneliana by both possessing prismatic

shaped seeds; a rose pink corolla and spongy

lower leaf segments in the former and a light

yellow corolla and inflated peduncles in the

latter (Taylor 1989).

There are several characters that

differentiate Utricularia corneliana from

U. reflexa , namely a lack of rhizoids in the

former; an upper corolla lip that is longer

than the lower, with an upper lip rear surface

sparsely hairy only on the lower half (Fig. 1),

versus an entirely hairy surface in U. reflexa

(Taylor 1989: fig. 194, p. 640); internal trap

quadrifid gland arms that are 14 versus 30

times as long as they are wide (Fig 2B, versus

Taylor 1989: fig. C, p. 17); flat lenticular

shaped seeds ( c. 1 mm in diameter) (Fig. 1,

2C), versus disc shaped seeds (0.4-0.8 mm in

diameter) that are 2-3 times wider than thick

(Taylor 1989: fig. 194, p. 640).

Conservation status : After a search along

the circumference of the type locality swamp

(Fig. 3), plants were not observed anywhere

else. Two nearby swamps ( c. 10 and 15 km

away respectively) were also examined with

no other sightings.

Considering the limited geographic

distribution of Utricularia corneliana and its

low frequency at the collection site, it is likely

that this plant is extremely rare. The collection

site is on leasehold land and is therefore not

protected.

If Utricularia corneliana is more

widespread than appears, the question

remains as to why it had not been collected

before this study? One possible answer is that

the flowers of U. corneliana resemble those of

other local Utricularia species (U. aurea , U.

gibba , and U. stellaris) in the general shape

and colour (yellow), blending in with these

more common species.

It could also be the case that habitat

destruction, erosion, weed infestation, and

associated eutrophication of swamps and

lagoons, early on in the agricultural history

of the region, has reduced the population size

of Utricularia corneliana. An example of a

local disappearance of a fellow suspended

aquatic species is that of U tubulata F.Muell.,

the type specimen of which ( Armit 222

[MEL1513562]) was collected in 1875 on

‘Cashmere’ (now ‘Glen Ruth’ and ‘Goshen’

stations), about 15 km E of the U. corneliana

site. Armit recorded the plant as “floating in

swamps and lagoons” on “Cashmere”, but

it has not since been collected anywhere in

Queensland, except for a single site in the far

north-west corner of the state {Jacobs 1465

[NSW]).

A more intensive survey of this area

of Queensland is warranted to determine

presence and extent of both the above

species; although it is likely that Utricularia

corneliana has mostly suffered the same early

fate as that of the local U. tubulata. At present

the conservation status of U. corneliana

should be regarded as Data Deficient.

Etymology: The specific epithet is in honour

of Cornelia M. Jobson, the author’s wife and

field assistant.

Acknowledgements

I thank Roderick Fensham and Peter

Bostock (BRI) for providing information on

specimens. I also thank Catherine Wardrop for

providing the detailed illustrations presented

in this paper, Peter Wilson for preparing the

Latin diagnosis, Marco Duretto for providing

helpful comments on the manuscript (all

NSW), Lubomir Adamec (Czech Republic)

for supplying plant material, and Cornelia

Jobson for help with fieldwork. Scientific

Purposes permits were obtained through the

Queensland Department of Environment and

Resource Management (WITK08454010,

WISP08454110). This work was partly

supported by ABRS grant RFL212-45.

604

Austrobaileya 8(4): 601-607 (2012)

Key to Australian and related suspended aquatic species of Utricularia (modified from

Taylor 1989)

Abbreviations: NSW (New South Wales), Qld (Queensland), NT (Northern Territory), SA

(South Australia), Tas (Tasmania), WA (Western Australia)

1 Leaves verticillate; peduncle inflated; corolla very pale pink with a very

slender spur 1.5-2 cm long.U. tubulata (Qld, NT, WA)

1. Leaves not verticillate (some semiverticillate); peduncle not inflated;

corolla yellow.2

2 Peduncle with whorl of usually inflated leaf-like structures at or above

base; primary segments of leaves 3-6.3

2. Peduncle without a whorl of inflated leaf-like organs; primary segments of

leaves 2.5

3 Inflated leaf-like organs fusiform, arising from base, or near base

of peduncle.U. aurea (NSW, Qld, NT, WA)

3. Inflated leaf-like organs ellipsoid, arising some distance above base

of peduncle.4

4 Inflated leaf-like organs sessile with capillary segments arising from

distal half only; seeds disk shaped, angular (not winged); calyx about

equal in length to capsule.U. stellaris (NSW, Qld, NT, WA)

4. Inflated leaf-like organs stipitate with capillary segments arising from

distal half and from base; seeds lenticular, narrowly winged; calyx

much shorter than capsule.U. muelleri (Qld, NT, WA)

5 Corolla externally pubescent; traps always inserted at angle between leaf segments .... 6

5. Corolla externally glabrous; traps lateral on leaf segments.7

6 Corolla upper lip longer than lower; seed flat, lenticular.U. corneliana (Qld)

6. Corolla upper lip equal to or shorter than lower; seed thick, disk¬

shaped .U. reflexa (tropical Africa, Madagascar)

7 Leaves with ultimate segments few (2-8); upper corolla lip larger than

lower.U. gibba (All states except SA, Tas)

7. Leaves with ultimate segments numerous (20-80); upper corolla lip

smaller than lower.U. australis (All states)

Jobson, Utricularia comeliana

605

Fig. 1 . Utricularia comeliana. A. habit x5.5. B. leaf segments with traps x7. C. bladder-trap in lateral view xlO. D.

bract with pedicel base in situ xlO. E. sepals with exposed ovary x7. F. flower in frontal view x5.5. G. flower in rear

view x5.5. H. flower in lateral view x5.5. I. fruiting capsule with calyx x7. J. stamen x20. K. seed x40. A-K from

Jobson 1281 (NSW).

606

Austrobaileya 8(4): 601-607 (2012)

Fig. 2. Utricularia corneliana. A. habit. B. Internal quadrifid gland of bladder trap. C. flat lenticular seed. A-C from

Jobson 1281 (NSW).

Jobson, Utricularia comeliana

607

Fig. 3. Shallow swamp habitat holding the observed population of Utricularia comeliana. Insert is a topographic map

of northern Queensland showing vicinity of collection site (red box).

References

Gassin R.J. (1993). Utricularia beaugleholei

(Lentibulariaceae: subgenus Utricularia :

section Pleiochasia ), a new species from south¬

eastern Australia. Muelleria 8: 37-42.

Jobson, R.W. (2012). A new species of Utricularia

(Lentibulariaceae) from northern Queensland,

Australia. Telopea 14: 49—57.

Jobson, R.W., Playford, J., Cameron, K.M. &

Albert, V.A. (2003). Molecular phylogeny

of Lentibulariaceae inferred from rpsl6 and

trnl-f chloroplast gene regions: implications

for character evolution and biogeography.

Systematic Botany 28: 157-171.

Lowrie, A. (1998). A new species of Utricularia

(Lentibulariaceae) from the south-west of

Western Australia. Nuytsia 12: 37-41.

- (2002). Utricularia petertaylorii

(Lentibulariaceae), a new species from the

south-west of Western Australia. Nuytsia 14:

405-410.

Lowrie, A., Cowie, I.D. & Conran, J.G. (2008). A

new species and section of Utricularia

(Lentibulariaceae) from northern Australia.

Telopea 12: 31-46.

Reut, M. & Jobson, R.W. (2010). A phylogenetic study

of subgenus Polypompholyx. a parallel radiation

of Utricularia (Lentibulariaceae) throughout

Australasia. Australian Systematic Botany 23:

152-161.

Taylor, P. (1989). The genus Utricularia. Kew Bulletin

Additional Series XIV. HMSO: London.

Stylidium elachophyllum A.R.Bean & M.T. Mathieson

(Stylidiaceae), a new species from northern Queensland

A.R. Bean & M.T. Mathieson

Summary

Bean, A.R. & Mathieson, M.T. (2012). Stylidium elachophyllum A.R.Bean & M.T.Mathieson

(Stylidiaceae), a new species from northern Queensland. Austrobaileya 8(4): 608-612. Stylidium

elachophyllum is described as new. It is an ephemeral herb known only from the Hann Tableland in

northern Queensland, with a morphological affinity to S. fissilobum F.Muell., S. prophyllum Lowrie

& Kenneally and S. oviflorum A.R.Bean. The species is illustrated and diagnosed against related

taxa. Its conservation status is assessed and a status of Endangered is recommended.

Key Words: Stylidiaceae, Stylidium , Stylidium elachophyllum , Australia flora, Queensland flora, new

species, taxonomy, conservation status

A.R.Bean & M.T.Mathieson, Queensland Herbarium, Department of Science, Information

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

Queensland 4066, Australia. Email: Tony.Bean@science.dsitia.qld.gov.au. Email: Michael.

Mathie son@science. dsitiat. qld. gov. au

Introduction

The tropical triggerplants of Australia are

diverse and difficult to study as they are

nearly all ephemeral and are fertile just after

the wet season when field access to them is

difficult. A number of new Stylidium species

from tropical Australia have been described

in recent years (Lowrie & Kenneally 1997,

1999; Bean 1997, 2000, 2010).

During a visit to the Hann Tableland, west

of Mareeba in north Queensland in May 2010,

the second author discovered a population of

Stylidium Sw. ex Willd. that did not match

any named species. No material could be

found at BRI or CNS that corresponded with

this species. Further material was collected

in March 2012, enabling its unnamed status

to be confirmed and its formal description

herein.

Materials and methods

This research is based on a study of herbarium

specimens and associated spirit collection at

BRI and CNS, and field studies at the type

locality. All measurements have been made

from material preserved in spirit.

RE refers to Regional Ecosystem,

descriptions of which can be viewed at (in

Accepted for publication 13 July 2012

this case): http://www.derm.qld.gov.au/

wildlife-ecosvstems/biodiversitv/regional

ecosvstems/search.php?&page=31

Taxonomy

Stylidium elachophyllum A.R.Bean &

M.T.Mathieson species nova; affinis Stylidio

fissilobo sed absentia glandium paracorollae,

apicibus integris loborum posteriorum

anteriorumque (in S. fissilobo utrique bilobi),

labello e basi sinus anterioris affixo (in S.

fissilobo parti externae tubi corollae affixo)

et capsulis 6-9 mm longis (non 12-21 mm

longis) differens. Typus: Queensland. Cook

District: Hann Tableland National Park, NW

of Mareeba, 27 March 2012, M.T. Mathieson

MTM1292 (holo: BRI [1 sheet + spirit]; iso:

CNS, distribuendi).

Annual herb, 3-10 cm high. Glandular hairs

0.05-0.1 mm long, glands dark red to brown,

globose. Rootstock not thickened. Stems

present, glabrous. Leaves 2-6, scattered

along stems, green, deltate, 0.5-1.7 mm long,

0.3-0.6 mm wide, sessile, glabrous, apex

obtuse or acute, base truncate. Scapes absent.

Inflorescences 1.5-3 cm long, determinate;

branches monochasially cymose. Bracts

deltate to lanceolate, 0.9-1.5 mm long,

glabrous, apex obtuse or acute. Pedicels

absent. Hypanthium linear, glandular-hairy at

Bean & Mathieson, Stylidium elachophyllum

distal end only. Sepals elliptical, 3 free and 2

mostly fused, 1.2-1.5 mm long, 0.3-0.4 mm

wide, sparsely glandular-hairy, apex obtuse.

Corolla pink, tinged yellow at the junction of

anterior and posterior petals, glandular-hairy

on tube only. Corolla tube 17-2.1 mm long,

with sinus on anterior side only. Paracorolla

(throat appendages) discontinuous, glabrous,

decurrent with corolla, free part c. 0.1 mm

high; lobes 4-6, all similar, 2 opposite the

anterior petals, 2-4 opposite the posterior

petals; glands absent. Labellum attached at

base of anterior sinus of corolla tube, ovate

or lanceolate, 0.5-0.8 mm long, glabrous;

terminal appendage absent or present, to 0.3

mm long; basal appendages absent. Corolla

lobes divided to the throat, with a central

longitudinal vein; anterior lobes obovate, 0.7-

1.1 mm long, 0.4-0.7 mm wide, entire, obtuse;

posterior lobes obovate, 1.5-1.8 mm long,

0.8-1.2 mm wide, entire, obtuse. Column

3-4 mm long, of uniform width throughout,

glabrous, lateral lobes absent, spur absent.

Stigma sessile. Capsule linear, without raised

longitudinal ribs, 6-9 mm long excluding

sepals, 0.6-0.7 mm wide, halves coherent

distally. Seeds ellipsoidal, 0.2-0.25 mm long,

brown, colliculate. Fig. 1-3

Additional specimen examined : Queensland. Cook

District: Hann Tableland N.P., NW of Mareeba, May

2010, Mathieson MTM814 & Forster (BRI).

Distribution and habitat: Stylidium

elachophyllum is endemic to northern

Queensland and has been found only in the

Hann Tableland N.P., about 20 km north¬

west of Mareeba. It grows in ephemeral

vegetation in shallow soil-filled depressions

or fringing mats of Borya septentrionalis

F.Muell. on granite pavements (RE7.12.37)

at approximately 800 m altitude. Vascular

flora associated with this habitat at this

site includes Xyris pauciflora Willd.,

Polycarpaea spirostylis F.Muell., Eriocaulon

pusillum R.Br., Drosera indica L., Byblis

liniflora Salisb., Utricularia chrysantha

R.Br., Stylidium oviflorum A.R.Bean, S.

capillare R.Br., annual grasses including

Schizachyrium pachyarthron C.A.Gardner,

Dimeria ornithopoda Trin. and Sacciolopis

indica (L.) Chase, and annual sedges including

Fimbristylis furva R.Br. and Fuirena ciliaris

609

(L.) Roxb.. These ephemeral species grow

during the wet season when there is a largely

continuous supply of water, either as rain or

run-off, but completely desiccate during the

dry season.

Vegetation surrounding the granite

pavements is comprised of sclerophyll

woodland dominated by Eucalyptus

portuensis K.D.Hill, E. atrata L.A.S. Johnson

& K.D.Hill, E. lockyeri Blaxell & K.D.Hill,

Corymbia intermedia (R.T.Baker) K.D.Hill

& L.A.S. Johnson and C. leichhardtii

(F.M. Bailey) K.D.Hill & L.A.S. Johnson,

generally with scattered Melaleuca viridiflora

Sol. ex Gaertn. and Xanthorrhoea johnsonii

A.T.Lee in the understorey. The area is in

general quite fire-prone.

Phenology: Flowers and fruits are recorded

for March, when plants were observed to be

abundant, and May, when very few plants

were observed as the habitat dried out.

Affinities: Stylidium elachophyllum is

morphologically allied to S. fissilobum

F.Muell., S. prophyllum Lowrie & Kenneally

and S. oviflorum of S. subgenus Andersonia

(R.Br. ex G.Don) Mildbr. (see Bean 2000).

It shares with these species the small scale¬

like leaves that are scattered along the stem of

the plant, and the corolla lobes divided to the

throat or almost so.

Stylidum elachophyllum differs from S.

fissilobum by having entire corolla lobe apices

(bilobed in S. fissilobum ), a labellum which

is attached at the base of the anterior sinus

(attached to the outside of the corolla tube

in S. fissilobum ), and capsules which are 6-9

mm long (12-21 mm long in S. fissilobum). S.

elachophyllum differs from S. oviflorum in

possessing a pink corolla (white and yellow

in S. oviflorum ) with smaller anterior and

posterior lobes, a labellum which is attached

at the base of the anterior sinus (attached to

the outside of the corolla tube in S. oviflorum ),

a shorter column that lacks lateral lobes,

capsules which are 6-9 mm long (18-25

mm long for S. oviflorum ), and a colliculate

seed surface (smooth in S. oviflorum ). S.

elachophyllum differs from S. prophyllum in

having glandular-hairy sepals (glabrous in

610 Austrobaileya 8(4): 608-612 (2012)

Fig. 1. Stylidium elacliopliyllum. A. whole plant *1.5. B. flower x8. C. calyx lobes xl6. D. corolla and column xi6. E.

undehisced capsule x8. All from Mathieson MTM1292 (BRI). Del. W. Smith

S. prophyllum), a smaller and more obscure

paracorolla, much smaller anterior and

posterior corolla lobes, a column which is

3-4 mm long (5-6 mm in S. prophyllum ), and

capsules which are 6-9 mm long (11-17 mm

in S. prophyllum).

Conservation status : This species is only

known from a single location on the Hann

Tableland within Hann Tableland National

Park. It was not encountered elsewhere on the

Hann Tableland despite extensive searches in

similar habitats during two expeditions.

The population at the type locality

is estimated to be between 200 and 500

plants occupying an area of less than five

hectares. With encroachment of introduced,

hyperinvasive grasses (viz. giant rat’s tail

grass Sporoboluspyramidalis (Lam.) Hitchc.,

grader grass Themeda quadrivalvis (L.)

Kuntze and gamba grass Andropogon gayanus

Kunth) and the rampant herbaceous weed

Praxelis clematidea R.M.King & H.Rob.

into the ephemeral flush wetland habitats on

granite outcrops immediately adjacent to the

Bean & Mathieson, Stylidium elachophyllum

Fig. 3. Stylidium elachophyllum , front view of flower, photographed in habitat (Mathieson MTM1292 )

612

Austrobaileya 8 ( 4 ): 608-612 ( 2012 )

type locality, this population is considered

greatly threatened. Applying criteria of

the IUCN (IUCN 2001), the recommended

conservation status is Endangered (B2b(iii);

C2a(ii)).

Etymology : From the Greek elacho- small,

and phyllon - leaf, in reference to the very

small scale-like leaves of this species.

Acknowledgements

Collections of this species were made on a

Bush Blitz nature discovery expedition in

2010 and, in 2012, using a Bush Blitz Tactical

Taxonomy Grant awarded to MTM; both

funded by the Commonwealth Government in

part. We thank Will Smith for the illustrations,

and Peter Bostock for the Latin diagnosis.

The following people and organisations are

thanked for their assistance that made the

survey possible: Keith McDonald (Dept, of

Environment & Heritage Protection (DEHP)-

Atherton) for logistical support and co¬

ordination; Traditional Owners for the Hann

Tableland, John and Troy Grainer; Cairns/

Mareeba National Parks & Wildlife Service

staff (Jonathon Roth, Robert Miller) for

logistical and field support; Cape York Tenure

Unit (DEHP - Cairns) (Georgianna Fien,

Eric Wason); Southedge Research Station

for accommodation and use of facilities;

Cape York Helicopters for safe and efficient

helicopter provision.

References

Bean, A.R. (1999). A revision of Stylidium sect. Debilia

Mildbr., S. sect. Floodia Mildbr. and 5. sect.

Lanata A.R.Bean (Stylidiaceae). Austrobaileya

5: 427-455.

- (2000). A revision of Stylidium subg. Andersonia

(R.Br. ex G.Don) Mildbr. (Stylidiaceae).

Austrobaileya 5: 589-649.

- (2010). Four new species of Stylidium Sw.

(Stylidiaceae) from northern Australia.

Austrobaileya 8: 107-223.

Iucn (2001). IUCN Red List of Categories and

Criteria: Version 3.1. IUCN Species Survival

Commission: Gland (Switzerland)/Cambridge

(United Kingdom).

Lowrie, A. & Kenneally, K.F. (1997). Eight new species

of triggerplant {Stylidium. Stylidiaceae) from

northern Australia. Nuytsia 11: 213-215.

- (1999). Stylidium candelabrum (Stylidiaceae),

a new species from the Northern Territory,

Australia. Nuytsia 13: 251-254.

Austrobaileya 8(4): 613-615 (2012)

SHORT COMMUNICATION

Poikilogyne cordifolia (Cogn.) Mansf., a newly recorded

genus and species of Melastomataceae for Australia

Ashley R. Field

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

Brisbane Botanic Gardens, Mt Coot-tha, Toowong, Queensland 4066, Australia & Australian Tropical

Herbarium, James Cook University Cairns Campus, Smithfield, Queensland 4878, Australia. Email:

Ashlev.Field@science.dsitia.qld.gov.au

Melastomataceae is a diverse family of plants

in the order Myrtales (A.P.G. 2009). The

family comprises more than 4500 species

and 150 genera (Renner 1993) and exhibits

its highest diversity in the Neotropics, with

secondary centres of diversity occurring in

the Malesian and African regions (Renner

1993; Cellinese 2007). Even though Cape

York Peninsula, Queensland is relatively

close to the Malesian region it harbours only

five indigenous Melastomataceae species

from four genera (Bostock & Holland

2010). Approximately 60% of the Australian

Melastomataceae flora has been introduced by

humans via the horticultural trade, including

the significant environmental weeds Clidemia

hirta (L.) D.Don, Mikania micrantha Kunth

and Miconia calvescens DC. (Bostock &

Holland 2010). Therefore, upon receiving an

attractive and unidentified Melastomataceae

collected from Cape York Peninsula it was

initially expected that it belonged to a weedy

horticultural species. Closer examination of

the buds, leaf type, fruit type and anthers

indicated that it belonged to the widespread

New Guinea species Poikilogyne cordifolia

Mansf and that it is likely to be indigenous.

It is the first record of the genus Poikilogyne

Baker f. outside Malesia.

Poikilogyne is a genus of more than 20

species with a centre of diversity in New

Guinea where there are a number of narrow

range endemics (Cellinese 2007). Most

Poikilogyne are terrestrial shrubs or vines

with ovate to orbicular leaves with major

veins radiating from the cordate leaf base.

Accepted for publication 30 October 2012

These veins are interconnected by cross veins

which give the leaves a laddered appearance

typical of many Melastomataceae. The leaf

margins of many Poikilogyne , particularly

the montane species, are toothed in some

way and the indument of the leaves, buds and

stems is variable with many species being

scurfy or villose. The inflorescences are

usually terminal and multi-branched and bear

5-merous flowers (also 4-6 merous) with 10

actinomorphic stamens and a campanulate

five ribbed hypanthium (also 4-6 ribbed).

The anthers are described by Cellinese

(2007) as having a very small terminal pore,

a prominently thickened connective and an

obtuse basal spur. The fruits are dehiscent,

longitudinally splitting capsules borne in

a distinct 10 ribbed cup (also 8-12 ribbed)

formed by the hypanthium. The fruit of

Poikilogyne are dry and capsular and the

seeds are small. This is consistent with the

wind-dispersal mode for Melastomataceae

rather than bird dispersal found in species

with berry type fruits (Styles & Rosselli

1993).

Maxwell (1983) reduced Poikilogyne

ledermannii Mansf. to varietal rank within P.

cordifolia but did not provide any characters

to distinguish the two varieties. Based on

Mansfield’s (1926) circumscription of the two

taxa, and on material held in BRI that have

been determined by J.F.Maxwell, I consider

that the Australian specimen belongs to

Poikilogyne cordifolia var. cordifolia.

Poikilogyne cordifolia (Cogn.) Mansf., Bot.

Jahrb. Syst. 60: 111 (1926); Allomorphia

cordifolia Cogn., FI. Kais. Wilh. Land [K.M.

Schumann & M.U. Hollrung] 87 (1889). Type:

614

Papua New Guinea, s.loc., in 1886, [U.M.]

Hollrung 135 (holo: Bf; iso: BR 5214778,

scan !).

Poikilogyne cordifolia var. cordifolia;

J.F.Maxwell, Gard. Bull. Singapore 35: 223

(1983).

Description of Australian specimen :

Branching woody shrub to 1 m tall with

tetrangular stems bearing a distinct keel

along the corners of young and mature stems.

Immature shoots with a reddish-pink blush,

mature bark pale yellow-brown with a few

raised corky lenticels. All parts of stems,

leaves, flowers and fruits glabrous. Leaves

and branchlets opposite with a distinct

interpetiolar scar. Petioles fleshy, 16-110 mm

long, reddish pink. Leaves orbicular-cordate,

60-160 mm long by 60-120 mm wide,

with a distinctly cordate base and a weakly

acuminate apex; midvein and 2-4 pairs of

major lateral veins all radiating from the leaf

base petiole junction and interconnected by

secondary cross veins; all venation prominent

on both surfaces and distinctly raised on

the abaxial surface. Inflorescences terminal

thyrses, 50-150 mm long and 50-150 mm

wide, reddish-pink, with flowers opening

sequentially. Inflorescence bracts minute,

reddish-pink, sometimes absent. Flowers

5-merous, occasionally 4 or 6-merous within

the same inflorescence. Pedicels 7-9 mm long

and 1 mm in diameter. Hypanthium 9-14 mm

long and 4-9 mm in diameter, pink, with 5

apically ribbed obtuse calyx lobes. Petals

free (imbricate in bud), rounded-oval with

a small keel apically, 6-9 mm long and 4-9

mm wide, bicolored, pink and white. Anthers

actinomorphic, 4-6 mm long with a terminal

pore and a small basal spur. Filaments 5-6

mm long at anthesis. Anthers and filaments

folded inwards when flowers first open.

Ovary half-inferior, truncate, style less than

5 mm long when flowers open elongating

to 13 mm long and bearing a simple white

to reddish-pink stigma. Inflorescences and

fruit becoming dry and woody at maturity

and remaining attached to the plant in

branch sinuses. Fruits dry, dehiscent,

loculidically splitting, capsules with sepals

and hypanthium reducing to a thin partially

Austrobaileya 8 ( 4 ): 613-615 ( 2012 )

decayed dry membrane bearing 10 persistent

woody longitudinal ribs, corresponding with

the mid-vein and marginal veins of the sepals,

which grip the ovary. Seeds numerous per

locule, less than 1 mm long, brown. Fig 1.

Selected specimens examined : Indonesia. Papua.

Bernhard Camp, Idenburg River, Mar 1939, Brass

13069 (BRI). Papua New Guinea. Owen Stanley Range

between Mts Brown & Clarence, May 1926, Brass

1488 (BRI). Australia. Queensland. Cook District:

Munburra, Starcke, Sep 2012, Thompson 691 (BRI,

CNS).

Distribution and habitat : This species is

presently known in Australia from a single

record at Munburra, Starcke on eastern Cape

York Peninsula where it was growing on steep

metamorphic rocky slopes on an ecotone

between Eucalyptus and Lophostemon

dominated forest and rainforest. This

species is widespread and abundant in New

Guinea and it is expected that it will be more

widespread in other similar, poorly accessed,

habitats on eastern Cape York Peninsula.

Notes : The site where Poikilogyne cordifolia

was collected is remote, undisturbed and

did not contain any introduced plant species

(S.Thompson pers. comm.). This species is

unknown in cultivation in Australia and it is

considered unlikely that it has been introduced

via the horticultural trade. Combined, these

factors indicate that Poikilogyne cordifolia

is indigenous to Australia and that it has

naturally dispersed to Australia from Papua

New Guinea. Poikilogyne cordifolia is readily

distinguished from all other Australian

Melastomataceae by its large orbicular-heart

shaped cordate leaves and by its capsular

loculidically splitting fruit which are clasped

by the ribs of the sepals and hypanthium.

Conservation status : Insufficient data are

available to accurately assess the conservation

status of Poikilogyne cordifolia in Australia.

Poikilogyne cordifolia is abundant in New

Guinea and could be considered secure there.

Acknowledgments

I would like to thank Simon Thompson for

collecting the voucher material and also

Nicoletta Cellinese and Trevor Whiffin for

confirming the identification.

Field, Poiklogyne cordifolia

References

A.P.G. [=Angiosperm Phylogeny Group.] Ill (2009). An

update of the Angiosperm Phylogeny Group

classification for the orders and families of

flowering plants: APG III. Botanical Journal of

the Linnaean Society 161: 105-121.

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

of the Queensland Flora 2010. Queensland

Herbarium, Department of Environment and

Resource Management: Brisbane.

Cellinese, N. (2007). Two new species in the genus

Poikilogyne (Melastomataceae) from Papua

New Guinea. Novon 17: 20-23.

615

Mansfield, R. (1926). Die Melastomataceen von

Papuasian. Botanische Jahrbiicher fur

Systematik, Pflanzengeschichte und

Pflanzengeographie 60: 105-143.

Maxwell, J.F. (1983). Taxonomic and nomenclatural

notes on Oxyspora,Anereincleistus, Poikilogyne

and All'omorphia (Melastomataceae). Gardens’

Bulletin Singapore 35: 209-226.

Renner, S.S. (1993). Phylogeny and classification of the

Melastomataceae and Memecylaceae. Nordic

Journal of Botany 13: 519-540.

Styles, F.G. & Rosselli, L. (1993). Consumption of fruits

of the Melastomataceae by birds: how diffuse is

coevolution? Vegetatio 107/108: 57-73.

Fig. 1. Poikilogyne cordifolia. A. flowering and fruiting branch. B. lateral view of flower buds. C. lateral and apical

view of mature dehiscent capsules. All from fresh material of Thompson 691 (BRI). Photographs by Andrea Lim.

Croton lucens P.I.Forst. (Euphorbiaceae), a new

species from south-east Queensland

Paul I. Forster

Summary

Forster, PI. (2012). Croton lucens P.I.Forst. (Euphorbiaceae), a new species from south-east

Queensland. Austrobaileya 8(4): 616-623. A shrubby, perennial species of Croton L. from the

Gympie area in south-east Queensland is newly described as Croton lucens P.I.Forst. This new species

is known from only three small populations (less than 100 plants) and is considered Endangered. An

identification key is provided for Croton species from south-east Queensland that have foliage that is

penninerved and fawn-silver to silver-white below.

Key Words: Euphorbiaceae, Croton , Croton lucens , Australia flora, Queensland flora, identification

key, taxonomy, new species, endangered conservation status

P.I.Forster, Queensland Herbarium, Department of Science, Information Technology, Innovation

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

Email: Paul.Forster@science.dsitia.qld.gov.au

Introduction

A revision of Australian Croton L. was

published ten years ago with twenty-seven

native species recognised (Forster 2003). An

additional native species (C. dichromifolius

P.I.Forst.) from Cape York Peninsula was

added by Forster (2010).

Fieldwork near Gympie, south-east

Queensland in July 2009 resulted in the

discovery of several populations of what

appeared to be a previously unknown and

undescribed species of Croton ; however, this

material was sterile and no collections were

made at the time. Subsequent revisits to the

three known localities have resulted in several

collections of fertile material and it is now

possible to describe these in the current paper

as the new species Croton lucens P.I.Forst.

Under the schema of Forster (2003),

this species can be placed in Group 5,

along with Croton capitis-york Airy Shaw,

C. dichromifolius , C. insularis Baill., C.

mamillatus P.I.Forst., C. phebalioides Muell.

Arg., C. simulans P.I.Forst. and C. stigmatosus

F.Muell. This group of species comprises

all shrubs or small trees with penninerved

foliage that is ± silver-white below due to

dense adpressed indumentum of trichomes

Accepted for publication 10 July 2012

and/or scales. Unpublished molecular analysis

of Australian Croton indicates that these

species (with C. stockeri Airy Shaw added)

may be a relatively natural grouping (P. Berry

& B. van Ee pers. comm. Feb 2010), although

C. lucens remains to be sequenced and

compared to these species. An identification

key is provided for the south-east Queensland

representatives of this group of Crotons.

Materials and methods

Data presented and discussed in this paper are

based on field collections and observations

made between 2009 and 2012 with specimens

deposited in the Queensland Herbarium

and duplicate distribution as indicated.

The morphological description (especially

indumentum types) is modelled on those of

Forster (2003). Venation terminology largely

follows Hickey (1973) and Ash et al. (1999)

with the recognition of a midrib (1° vein order),

lateral veins (2° vein order) and intercostal

veins (3° and onwards vein orders) within

any leaf lamina. When an intercostal vein

comprises a continuous raised line of cells it

is termed ‘distinct’; if it is discontinuous or

fades away into the body of the lamina, it is

termed ‘indistinct’. Indumentum cover is

described using the terminology of Hewson

(1988), except that ‘scattered’ is used instead

of‘isolated’. The shapes of leaves, sepals and

Forster, Croton lucens

617

petals are described using the terminology

of Hickey & King (2000). Length and

width dimensions are indicated as length

measurement x width measurement followed

by the measurement unit.

Abbreviations used in text: NCA (Nature

Conservation Act 1992 ) http://www.

legislation, qld. gov, au/legisltn/current/n/

naturecona92.pdf

Taxonomy

Croton lucens RI.Forst., species nova; C.

insulari Baill. simillimus sed ab eo nectariis

extrafloral ibus absentibus vel tubulosis

(non sessilibus circularibusque), lamina

folii idioblastis densis reflectivis in pagina

superiore praedita (adversum apparenter

nullam) et floribus masculis minoribus

staminibus paucioribus (10 vel 11 non 14-

18) differt. Typus: Queensland. Wide Bay

District: Marys Creek State Forest (S.F.124),

Groundwater Logging Area, Spring Gully;

18 km SW of Gympie, 10 February 2012,

P.I. Forster PIF38604 (holo: BRI [2 sheets +

spirit]; iso: MEL, MICH).

Perennial shrub or small tree to 5 m tall,

evergreen, monoecious; indumentum

primarily of sessile peltate scales, uncoloured

to pale silver to silver-yellow or silver-brown

(with age) (appearing fawn-silver to silver en

masse), peltate scales silver-yellow to silver-

brown in central column and uncoloured to

pale silver-brown in the rays, other trichomes

uncoloured. Bark flaky in irregular patches,

cream-fawn. Branchlets ± rounded, with

dense interlocking peltate scales when young

and on most leafy stems, glabrescent; stipules

subulate, entire, 6-8 x 0.4-0.5 mm, with

dense peltate scales coloured brown-fawn.

Leaves alternate, thinly coriaceous, markedly

discolorous, petiolate; petioles 3.5-5.5 x

1-1.2 mm, deeply channelled on top, with

dense interlocking peltate scales; lamina

elliptic-ovate to obovate, 13-75 x 4-26 mm,

penninerved with 5-7 lateral 2° veins per side

of 1° midrib; upper surface glossy, dark green,

1° and 2° veins barely visible, 3° veins obscure,

sparse and often deeply embedded peltate

scales over entire surface, dense reflective

idioblasts; lower surface fawn-silver, 1° veins

raised, 2° veins scarcely visible and indistinct,

3° veins obscure, surface completely obscured

by dense, interlocking peltate scales, neither

scabrid nor velutinous; margins slightly

undulate, not toothed; foliar glands absent; tip

acute to acuminate, base cuneate; extrafloral

nectaries apparently absent on most leaves,

occasionally present and tubular to c. 0.8 x 0.2

mm, c. 3 mm from lamina base. Inflorescence

short, up to 14 mm long, unbranched, usually

bisexual and androgynous, mixed glomerules

not observed, pedunculate for 1-1.5 mm, axis

with dense, interlocking peltate scales; bracts

lanceolate-ovate, much reduced, 0.2-0.3 x

0.1-0.2 mm with dense peltate scales (no

trichomes). Male flowers c. 5 mm long and

4 mm diameter (includes pedicels), singly or

two open per inflorescence, generally not in

glomerules and spirally spaced 0.2-0.3 mm

apart; pedicels c. 1.5 x 0.2 mm, with dense

peltate scales; sepals valvate, lanceolate-

ovate, 1.4-1.5 x 1-1.2 mm, with dense

peltate scales coloured silver-brown; petals

lanceolate to weakly oblanceolate, 1.2-1.4

x 0.7-0.8 mm, densely marginally ciliate

and with a dense tuft of interlocking peltate

scales in apical 1/3; stamens 10 or 11, with

dense simple trichomes at base, filaments

filiform, 3-3.5 x 0.1-0.2 mm, glabrous,

anthers oblong-ovoid, 0.7-0.8 x 0.5-0.6 mm,

cream, glabrous. Female flowers held singly

and spaced up to 1 mm apart on inflorescence

axis; pedicels c. 6 x 1 mm (up to 9 mm long in

fruit), with dense, interlocking peltate scales;

sepals valvate, lanceolate-ovate, 1.3-1.5 x c.

1 mm, with dense marginal cilia and dense

peltate scales externally; petals absent; styles

3, linear-flabellate to 2 mm long, multifid and

once divided for 1.1-1.8 mm, barely connate

at base, with dense peltate scales to above the

point of division, occasional minute simple

glandular trichomes on tips; ovary 3-locular,

c. 3x3 mm, with dense interlocking peltate

scales. Fruits trilobate, globose-ovoid, 7-8

x 7-8 mm, with dense interlocking peltate

scales. Seeds ovoid, 3.5-6 mm long, 2.8-4

mm wide, 1.5-1.7 mm thick, pale brown with

occasionally dark brown blotches, ventral

surface bifacial, dorsal surface rounded,

618

Austrobaileya 8(4): 616-623 (2012)

Fig. 1. Croton lucens. A. habit of fertile branchlet x0.6. B. leaf abaxial surface showing 1° midrib and 2° lateral venation

xl.C. node with single inflorescence and $ flowers x4. D. dense interlocking peltate scales present on most vegetative parts

x24. E. node with stipule x4. F. lateral view of S flower xl2. G. apical view of $ flower xl2. H. lateral view of $ flower x8.

I. apical view of fruit x4. J. lateral view of fruit x4. K. ventral view of seed x6. L. lateral view of seed x6. A-F from Forster

PIF38604 (BRI); G & H from Leiper s.n., Jun 2012 (BRI); I-L from Leiper s.n., Jan 2012 (BRI). Del. W.Smith.

Forster, Croton lucens

619

micropylar ridge 2.8-4.8 mm long, caruncle

broadly depressed-ovate, 0.6-1 mm long,

0.8-1.2 mm wide, cream. Figs. 1-4.

Additional specimens examined. Queensland. Wide

Bay District: Marys Creek State Forest (S.F. 124), near

Marys Creek Bridge; 13 km WSW of Gympie, Jan 2012,

Leiper s.n. (BRI, MEL); ditto loc., Feb 2012, Leiper &

Smyrell s.n. (BRI, NSW); ditto loc ., Jun 2012, Leiper s.n.

(BRI).

Distribution and habitat : Croton lucens

has so far been found only in Marys Creek

State Forest to the southwest of Gympie

in south-east Queensland. Plants grow in

the understorey of hoop-pine vineforest

(araucarian microphyll vineforest) on shallow,

stony, red soils that are derived from a complex

mixture of metasediments and volcanic

intrusions. The geology of this area is highly

complex and comprises substrates that have

been intermixed and altered by subsequent

volcanic activity via vents and limited scale

flows (Ostwald 1992; Sivell & McCullock

2001; Sivell & Waterhouse 1998). The red

colour of the soils at the localities of the three

known populations may be due to weathering

of andesite (or less likely basalt or granite)

intrusions; however, limited observation of

outcropping rocks indicated a predominance

of more ancient Permian metasediments.

Notes: Croton lucens has markedly

discolorous foliage, although this in itself is

a common character in the genus. The dense,

interlocking peltate scales impart a fawn-silver

appearance to the lower surface of the leaf

lamina, although on close examination with

magnification the scales appear silver-yellow

to silver-brown in the central column and

uncoloured to pale silver-brown in the rays.

While many Crotons have this sort of foliage,

C. lucens is notable for the ‘shimmering’

appearance of the foliage, which seems to be

a result of light reflection from the numerous

embedded idioblasts on the abaxial surface of

the leaf lamina (Fig. 2).

Fig. 2. Croton lucens. Foliage showing glossy, reflective adaxial surface and silver-white abaxial surface (from Leiper

& Smyrell s.n. [BRI]). Photo: G.Leiper.

620

Austrobaileya 8(4): 616-623 (2012)

Fig. 3. Croton lucens. Fruit and male flower (from Leiper & Smyrell s.n. [BRI]). Photo: G.Leiper.

In some respects, the general habit of

Croton lucens is most similar to C.mamillatus;

however, the newly described species differs

from the latter in numerous character states of

foliage, flowers and fruit (Table 1). In terms

of the foliage indumentum, C. lucens is most

similar to C. insularis\ however it differs from

this species in the extrafloral nectaries (absent

or tubular versus sessile and circular), the leaf

lamina with dense reflective idioblasts above

(versus apparently none) and the smaller male

flowers with fewer stamens (10 or 11 versus

14-18). The inflorescences of C. lucens (Figs.

3 & 4) are consistently much shorter (up to 14

mm long) than those of C. insularis (up to 120

mm long [Forster 2003]) and as a result have

far fewer flowers per individual unit. Under

some drought conditions the latter species

may also have relatively short inflorescences

(less than 50 mm long); however, these are

usually intermixed on the same individual

with longer, more typical ones. Despite the

sympatry between these two superficially

similar species, no intermediates have been

observed.

Very little fertile material was available for

description of this species and the degree of

variation in floral part numbers and sizes may

be more than documented here. Collectors

should pay particular attention to locating

female flowers as these have been rarely

encountered and are more inconspicuous

when compared to the males.

Conservation status : Croton lucens is

infrequent at Marys Creek State Forest

with only three small populations (less than

100 plants observed) known; all are highly

localised and two are in close proximity (less

than 1 km apart). It is likely that the three

populations are merely subpopulations of a

single metapopulation.

This State Forest has been largely altered

by conversion of the natural forest to hoop-

pine ( Araucaria cunninghamii Aiton ex

A.Cunn.) plantations; most now on a second

Forster, Croton lucens

621

Table 1. Comparison of morphological character states for some south-east Queensland

species of Croton

Character

state

C. insular is

C. lucens

C. mamillatus

C. phebaliodes

C. stigmatosus

Branchlet

indumentum

dense,

interlocking

peltate scales

dense,

interlocking

peltate scales

dense peltate

trichomes

dense stellate

trichomes

dense peltate

scales and

scattered stellate

trichomes

Extrafloral

nectaries

on leaf

usually 2 at

top of petiole,

sessile, circular

general

absent, or

tubular and

near lamina

base

generally

absent, or

sessile circular

usually 2 at

lamina base,

sessile, ellipsoid

usually 2 at

lamina base,

stipitate and

circular

Leaf lamina

margin

entire or

somewhat

sinuate

somewhat

sinuate

± entire

or weakly

denticulate

entire, sinuate

or denticulate

with 11-18 teeth

denticulate with

14-24 teeth

Leaf lamina 2°

lateral veins

5-8, ±

obscure due to

indumentum

5-7

12-14

7-14

12-15

Leaf lamina

adaxial

surface

indumentum

cover

dense,

interlocking

peltate scales

dense,

interlocking

peltate scales

dense peltate

trichomes and

peltate scales

dense,

interlocking

peltate trichomes

dense stellate

trichomes and

dense peltate

scales

Male flower

pedicel length

2.3-5 mm

c. 1.5 mm

2.5-3 mm

1.5-3.5 mm

2-4 mm

Male flower

sepal shape

and size

lanceolate-

triangular,

1.5-2.5 x

1.2-1.6 mm

lanceolate-

ovate, 1.4-1.5

x 1-1.2 mm

lanceolate-ovate

to ovate, 1.8-2.2

x 1.4-1.5 mm

lanceolate-ovate

to obovate, 1.2-

2.7 x 1.3-1.8 mm

lanceolate-

ovate, ovate or

obovate, 1.5-2.2

x 0.4-1.4 mm

Male flower

petal shape

and size

oblanceolate,

1.7-2.6 x

0.6-1 mm

lanceolate

to weakly

oblanceolate,

1.2-1.4 x

0.7-0.8 mm

obovate, 1.5-2

x 0.6-0.7 mm

obovate, 1.3-3

x 0.5-0.7 mm

obovate, 1.5-2.2

x 0.3-8 mm

Style division

once

twice

Stamen

number

14-18

10 or 11

9 or 10

10-12

Fruit surface

indumentum

and

protuberances

dense

interlocking

peltate scales,

no mamillate

protuberances

dense

interlocking

peltate scales,

no mamillate

protuberances

dense stellate

trichomes on

fleshy mamillate

protuberances

dense, sessile and

stalked stellate

trichomes

dense, stalked

trichomes on

fleshy mamillate

protuberances

622

Austrobaileya 8(4): 616-623 (2012)

Fig. 4. Croton lucens. Female flower (from Leiper s.n ., Jun 2012 [BRI]). Photo: G.Leiper.

rotation. The vineforest community now

exists mainly as small patches or narrow

strips (forest breaks) alongside watercourses

or on very steep slopes. The degree of habitat

fragmentation due to this historical forest

conversion is very high and connectivity

between many of the fragments is likely

to be minimal for plant species that are not

particularly vagile via vertebrate or wind

dispersal.

Compounding the habitat fragmentation

are severe and extensive infestations of

numerous alien environmental weeds,

including Aristolochia elegans Mast,

Asparagus africanus Lam., Dolichandra

unguis-cati (L.) L.Lohmann, Lantana

camara L. and Solarium mauritianum Scop.

These weeds are present not only in the areas

of forestry plantation, but are also widespread

within the canopy of the vineforest remnants

where no attempts at control are presently

being made.

Populations of several threatened

plant species listed under the NCA such as

Cupaniopsis shirleyana (F.M.Bailey) Radik.,

Floydia praealta (F.M.Bailey) LA.S.Johnson

& B.G.Briggs, Fontainea venosa Jessup &

Guymer and Macadamia integrifolia Maiden

& Betch co-exist with Croton lucens ; hence

in an ideal world the known localities for this

new species would be co-managed with the

others.

Under the IUCN (2001) criteria, this

species can be assessed as Endangered on

the criterion D.

Etymology : The specific epithet is derived

from the Latin lucens (shining, polished,

glistening) and alludes to the strikingly

discolorous foliage that appears to shimmer

in a light breeze.

Forster, Croton lucens 623

Key to Croton species with peiminerved and markedly discolorous foliage (+ silver-white

or fawn-silver below) in south-east Queensland

1 Branchlet indumentum and leaf lamina adaxial surface indumentum

comprising only dense interlocking peltate scales.2

1. Branchlet indumentum and leaf lamina adaxial surface indumentum of

dense peltate or stellate trichomes, or a mixture of peltate scales and

peltate or stellate trichomes.3

2 Extrafloral nectaries 2 at top of petiole, sessile and circular; male flower

pedicel length 2.3-5 mm, stamens 14-18.C. insularis

2. Extrafloral nectaries absent, or 2 near lamina base and tubular; male

flower pedicel length <2 mm, stamens 10 or 11.C. lucens

3 Branchlet indumentum of dense peltate trichomes; fruit surface with

dense stellate trichomes on fleshy mamillate protuberances.C. mamillatus

3. Branchlet indumentum of dense stellate trichomes, or dense peltate scales

and scattered stellate trichomes; fruit surface with dense, sessile and

stalked stellate trichomes or with dense, stalked trichomes on fleshy

mamillate protuberances.4

4 Branchlet indumentum of dense stellate trichomes only; extrafloral

nectaries usually 2 at lamina base, sessile, ellipsoid; leaf lamina adaxial

surface indumentum of dense, overlapping peltate trichomes.C. phebalioides

4. Branchlet indumentum of dense peltate scales and scattered stellate

trichomes; extrafloral nectaries usually 2 at lamina base, stipitate and

circular; leaf lamina adaxial surface indumentum of dense stellate

trichomes and dense peltate scales.C. stigmatosus

Acknowledgements

Repeated visits in pursuit of fertile material

to two of the localities for this species were

undertaken by Glenn Leiper and Greg

Smyrell. Will Smith drew the illustrations and

Peter Bostock provided the Latin diagnosis.

References

Ash, A., Ellis, B., Hickey, L.J., Johnson, K., Wilf, P. &

Wing, S. (1999). Manual of Leaf Architecture.

Smithsonian Institution: Washington.

Berry, P.E., Hipp, A.L., Wurdack, K.J., Van Ee, B. &

Riina, R. (2005). Molecular phylogenetics of

the giant genus Croton and tribe Crotoneae

(Euphorbiaceae sensu stricto) using ITS and

TRNL-TRNF DNA sequence data. American

Journal of Botany 92: 1520-1534.

Forster, PI. (2003). A taxonomic revision of Croton L.

(Euphorbiaceae) in Australia. Austrobaileya 6:

349-436.

- (2010). Croton dichromifolius P.I.Forst.

(Euphorbiaceae), a new species from Cape York

Peninsula, Queensland. Austrobaileya 8: 143—

149.

Hewson, H. (1988). Plant Indumentum. A Handbook of

Terminology. Australian Flora & Fauna Series

No. 9. Australian Government Publishing

Service: Canberra.

Hickey, L. J. (1973). Classification of the architecture of

dicotyledonous leaves. American Journal of

Botany 60: 17-33.

Hickey, M. & King, C. (2000). The Cambridge

Illustrated Glossary of Botanical Terms.

Cambridge University Press: Cambridge.

Iucn (2001). IUCN Red List Categories and Criteria.

Version 3.1. Gland: IUCN - The World

Conservation Union.

Ostwald, J. (1992). Mineralogy, paragenesis and

genesis of the braunite deposits of the Mary

Valley manganese belt, Queensland, Australia.

Mineralium Deposita 27: 326-335.

Sivell, W.J. & Mcculloch, M.T. (2001). Geochemical

and Nd-isotopic systematics of the Permo-

Triassic Gympie Group, southeast Queensland.

Australian Journal of Earth Sciences 48: 377-

393.

Sivell, W.J. & Waterhouse, J.B. (1988). Petrogenesis

of Gympie Group volcanics: evidence for

remnants of an early Permian volcanic arc in

eastern Australia. Lithos 21: 81-95.

Capsule Dehiscence in Viola betonicifolia Sm. (Violaceae)

R. John Little 1 & Glenn Leiper 2

Summary

Little, J. & Leiper, G. (2012). Capsule dehiscence in Viola betonicifolia Sm. (Violaceae).

Austrobaileya 8(4): 624-633. Seed dispersal syndromes in Viola are reviewed and the sequence of

events culminating in the dispersal of seeds from capsules of Viola betonicifolia is documented. Seed

parameters (length x width) and measurements of distances travelled after being ballistically ejected

from a capsule valve were determined. Preliminary observations were recorded of the approximate

length of time for a mature capsule to open and the approximate time for an open capsule to eject all

seeds from its three valves.

Key Words: Violaceae, Viola, Viola betonicifolia , capsule dehiscence, diplochory, myrmecochory

'R John Little, MagnaFlora, 16 Pebble River Circle, Sacramento, California 95831, U.S.A. Email:

bugsandbunnies4u@sbcglobal.net

2 Glenn Leiper, Tweedvale Street, Beenleigh, Queensland 4207, Australia. Email: leiper30@,bigpond.

com

Introduction

The effective dispersal of seeds to sites

where seedlings can successfully establish is

critical to most vascular plants that reproduce

sexually (Forster 2007). Understanding

the methods by which seeds are dispersed

provides valuable information about a species’

potential for distribution and colonisation,

and potential taxonomic relationships. Two

distinct seed dispersal syndromes have been

recognised in Viola. 1. myrmecochory (seed

dispersal by ants following passive release

from capsules) and 2. diplochory (explosive

ejection of seeds followed by attraction and

dispersal by ants) (Beattie & Lyons 1975).

Characters attributable to the myrmecochory

syndrome include a prostrate peduncle during

dehiscence with the capsule on or near the

ground; a calyx often swollen during fruiting;

a capsule with walls not greatly thickened;

seeds never under pressure during dehiscence;

seeds with a large and conspicuous elaiosome;

and seeds that are dispersed passively beneath

the parent plant. Characters attributable to the

diplochory syndrome include a tall peduncle

that is erect during dehiscence and carries the

capsule above the leaves; a calyx that does not

enlarge during fruit development; a capsule

Accepted for publication 9 November 2012

with walls thickened with sclereids that

produce pressure on seeds; seeds with a small

elaiosome; and seeds that are ballistically

dispersed 1-5 m from the parent plant. A

third type, called ballistic or autochory

(explosive ejection of seeds away from the

parent plant not related to ant dispersal) has

been suggested for Viola , but was not detected

as a distinct syndrome in the species studied

by Beattie & Lyons (1975).

Perhaps the first documented observation

of capsule dehiscence in Viola was that of

Leavitt (1902) for Viola rotundifolia Michx., a

perennial species from North America. Based

on the distance that seedlings germinated in

his garden from a single ‘mother’ plant, the

species appeared able to ballistically eject its

seeds 1.5-2.7 m (Leavitt 1902). Harrington

(1903) stated, “after dehiscence [the capsule

valves] fold lengthwise and eject the seeds

with some force,” and Bare (1979) said, “As

each segment dries, it slowly folds together

lengthwise, and the resultant pressure on the

seeds throws them several feet through the

air.”

The method by which the capsules of

diplochorous Viola species dehisce has

been described variously as “sometimes

dehiscing explosively” (Lawrence 1951),

Little & Lei per, Capsule dehiscence in Viola betonicifolia

“the matured capsule opens and ejects the

seeds in a moment” (Ohkawara & Higashi

1994), “seeds are explosively ejected from the

capsules” (Douglas & Ryan 1998), “opening

± explosively on maturity” (Thiele & Prober

2003), and “at maturity the capsule springs

open and the seeds are forcefully ejected”

(Karlsson etal. 2012). These descriptions give

the impression capsules ‘explode’ to release

their seeds, in a manner similar to some genera

in the Euphorbieae (Forster 2007). These

descriptions obscure the actual mechanism

by which capsules of many Viola species open

and how the seeds are subsequently dispersed.

Other workers describe how seeds are

released from the capsule, e.g., “explosively

ejected”, “ballistically dispersed”, or

“ballistically ejected”, which more accurately

describes how seeds are dispersed (Turnbull

& Culver 1983; Bulow-Olsen 1984; Brooks &

McGregor 1986; Douglas & Ryan 1998; Little

& McKinney in prep). However, there appear

to be no detailed studies for diplochorous

species documenting the actual process by

which Viola capsules open and how the seeds

are dispersed.

This note focuses on Viola betonicifolia

Sm. subsp. betonicifolia , a species native to

Australia. This species possesses characters

attributable to the diplochorous seed dispersal

syndrome, i.e., a tall peduncle with the capsule

above the leaves at the time of dehiscence, a

calyx that does not enlarge when the capsule

develops, seeds with a small elaiosome,

and seeds that are ballistically ejected some

distance from the parent plant.

During a visit by the first author to

Queensland and New South Wales, Australia,

from November 2011 through January 2012, an

opportunity arose to observe mature capsules

of Viola betonicifolia and V. he derace a Labill.

splitting open and to observe how seeds were

ejected from capsules of V betonicifolia. The

second author made numerous additional

observations in 2012 of how capsules open

in V betonicifolia and documented distances

that seeds were ejected.

625

Questions we attempted to answer in this

study for Viola betonicifolia included:

• What is the sequence of events for the

peduncle and capsule leading up to seed

dispersal?

• Does the inverted peduncle of

cleistogamous capsules remain pendant

or become erect?

• How long does it take for a mature

capsule to open?

• How long before seeds begin to disperse

from an open capsule?

• Are seeds dispersed randomly from

a valve, or is there a pattern (e.g., do

seeds disperse first from the center, the

proximal, or the distal ends of a valve)?

• What are minimum and maximum

dispersal distances?

For comparison with other Viola species

we documented the number of seeds produced

by cleistogamous flowers (abbreviated CL,

which are self-pollinated flowers, with minute

or no petals); measurements of seed lengths;

and morphological details of chasmogamous

flowers (abbreviated CH, which are flowers

with showy petals).

Materials and methods

There were two rounds of observations.

The first author observed and photographed

Viola betonicifolia plants near Mt Lindesay

in south-eastern Queensland, in December

2011 and 20 January 2012 during visits with

the second author. Initial observations of

capsules opening and seeds being ballistically

ejected were made by the first author on 22

January 2012 from a plant collected as a

voucher specimen on 20 January 2012, but

not yet pressed. In the field, the collected

plant was placed in a cup of water. The plant

had two capsules which for convenience are

discussed herein as A and B. Capsule A was

photographed at different intervals after it

had opened until all seeds were ejected. The

number of seeds in each valve was determined

for Capsule A, as was the approximate length

of time from when it began to open until

all seeds were ejected. While attention was

focused on Capsule A, Capsule B had opened

and ejected an unknown number of seeds.

626

Observation of Capsule B began when it

had a total of 10 seeds remaining among the

three valves. For both capsules, the time of

day and number of seeds remaining in each

valve were recorded at periodic intervals and

documented with photographs.

Although valuable data were obtained

during this first period of observation by the

first author, dispersal distances could only

be roughly approximated due the fact that it

was difficult to observe where the dehisced

seeds had landed. Dr Paul Forster encouraged

the authors to gather better data on dispersal

distances. The second author volunteered

to continue a second round of observations

and to measure dispersal distances and

seed parameters. Eventually, container-

grown plants were obtained and maintained

outdoors by the second author in Beenleigh,

Queensland. During this study, no CH

flowers were present. Plants with CL capsules

that appeared ready to open were brought

indoors and dispersal distances were able to

be measured under controlled circumstances.

To determine dispersal distances, white

flannelette sheets with roughened surfaces to

minimize bounce of seeds were placed around

the containers and the distances where seeds

landed were measured by two observers.

Photographs were taken of maturing

capsules to document

a) the movement of the peduncle;

b) the position of capsules prior to splitting

and opening;

c) whether capsules open slowly or

explosively;

d) the number of seeds per capsule; and

e) if seeds are randomly dispersed from a

valve or if there is a pattern by which

they are dispersed.

Seed measurements (length x width) were

made by the second author with a micrometer.

Statistics were calculated with Microsoft®

Excel.

Results

Viola betonicifolia subsp. betonicifolia is a

perennial, acaulescent species native to the

east coast of Australia from South Australia

Austrobaileya 8(4): 624-633 (2012)

and Tasmania to Queensland; it is also found

in New Guinea and the Philippines. It occurs

in a wide range of habitats from coastal dunes

and sclerophyllous forest to alpine herbfields

(Adams 1982). The species produces both

CH and CL flowers. The species is available

for horticulture commercially and is sold in

Australia mainly through specialist native

plant nurseries.

Floral Morphology

Fig. 1 is a typical CH flower of Viola

betonicifolia. The lateral petals are bearded

(Adams 1982; James 1990a,b). We found that

CH flowers on some plants at Mt Lindesay

also have hairs on the two upper petals (Fig.

1), which has not previously been reported.

The CL flowers of V betonicifolia have sepals

but no petals (Fig. 2).

Fig. 1. Typical CH Viola betonicifolia flower; Mt

Lindesay, Queensland. Photo: J. Little

Fig. 2. A CL flower developing from the base of the

plant. Sepals are visible enclosing the ovary. Petals are

absent. Photo: G. Leiper

Little & Leiper, Capsule dehiscence in Viola betonicifolia

Capsules

After the ovules are fertilized, the capsules

of CL flowers begin maturing in an inverted

position, typical of CL flowers of many

Viola species. As the capsule matures the

peduncle elongates, lifting the capsule to

an elevated position above many of the

leaves. It transitions from an inverted to

an upright position in about 24 hours (Fig.

3). The capsule also becomes noticeably

paler, possibly because the xylem ceases

functioning. The capsule becomes dehydrated

to facilitate splitting open and eventual seed

dispersal. The number of fertile seeds can be

counted when the capsule is upright. In about

another 24 hours, the capsule begins to split

open (Fig. 4), the three valves begin to spread

back and become completely separated (Fig.

5). Eventually the valves become more or less

parallel to the ground (Fig. 6). Occasionally,

a few seeds are ejected before the capsule is

fully open.

Fig. 3. Capsule from CL flower on peduncle transitioning

from inverted to upright position. Photo: G. Leiper

As the capsule valves begin to dry, the

edges move toward each other. The pressure

of the constricting valves squeezes the

seeds which eventually causes them to be

ballistically ejected. We observed that seeds at

the distal end of valves were ejected first (Fig.

6) and those at the proximal end were ejected

last (Fig. 7). However, for other capsules

627

Fig. 4. Capsule in fully upright position beginning to

split open. Note shrivelled sepals. Photo: G. Leiper

Fig. 5. Capsule completely split open with valves

beginning to spread apart. Each valve is ca. 13 mm long.

Photo: J. Little

628

Austrobaileya 8(4): 624-633 (2012)

Fig. 6. Seeds in a valve about midway through dehiscence.

Note the distal end of these valves contracted before the

proximal end. Photo: J. Little

seeds at the proximal end were ejected first.

Most capsules observed in this study (n=7)

ejected all their seeds (Fig. 8).

Observations of Seed Dehiscence

Capsule A (recorded by first author): Capsule

A was first observed open and in an upright

position at 1121 with a total of 35 seeds (Table

1) . The length of time it took to transition

from an inverted to upright position was not

observed nor was the time noted when the

first seed was ejected. All seeds were ejected

from the capsule in about 2.3 hr (138 minutes).

Valve 1, which initially had two more seeds

than the other valves, retained its seeds for a

longer period of time. Valve 3 was the first to

eject all its seeds.

Capsule B (recorded by first author): This

capsule was first noticed when it was open

and after it had ejected an unknown number

of seeds. Observations of Capsule B began at

1603 when it had 10 seeds remaining (Table

2) . Nine seeds were dispersed in the next 23

minutes. The time when the last seed was

ejected from Valve 3 was not observed, but no

seeds were present at 1715.

Fig. 7. One valve completely dehisced; 2 valves with one

seed remaining at proximal end of valve. Photo: J. Little

Fig. 8. Valves fully dehisced and completely clasped shut.

Photo: J. Little

The second author timed a capsule that

started opening at around 1400, was almost

fully open at 1445, and was fully open at

1530. During the 45 minute interval between

1445 and 1530, some seeds were observed

being ejected from the capsule. This capsule

took about 1.5 hr to fully open and all seeds

had been ejected by 1610. Thus, after it fully

Little & Lei per, Capsule dehiscence in Viola betonicifolia 629

Table 1. Seed dispersal times of Viola betonicifolia (Capsule A)

No. of Seeds in Valve

Time

Valve 1

Valve 2

Valve 3

Total

1121

1300

1303

1304

1311

1312

1338

1339

Table 2. Seed dispersal times of Viola betonicifolia (Capsule B)

No. of Seeds in Valve

Time

Valve 1

Valve 2

Valve 3

Total

1603

1604

1610

1626

1715

opened, it took 40 minutes until all seeds

were ejected (some seeds dispersed before it

was fully open).

Seeds

In the ovary, the ovoid-shaped seeds are

attached to the placenta at the narrow end

of the seed at the location of the elaiosome.

Typical of many Viola species, the seeds are

smooth and shiny when fully mature and fresh

(Fig. 9). Mature seeds appear black or brown.

Under magnification, they appear mottled

black and brown (Fig. 9). The elaiosome is

white or whitish when fresh (Fig. 9). The

mean number of seeds per CL capsule was

19 (n=7 capsules, 133 seeds; range 8-35; SD

8.42; Table 3). The mean number of seeds per

capsule is considered preliminary; additional

counts need to be made utilizing a larger data

set. Raw data for seed lengths and widths are

available from the authors.

Mean seed length was 1.44 mm (n=75

seeds, five capsules); range 1.10-1.71 mm;

SD 0.160. The length of five seeds was not

determined because they were inadvertently

Fig. 9. Viola betonicifolia seeds emphasising elaiosomes.

Lines are mm. Photo: J. Little

crushed when measured. The mean seed

width was 1.11 mm (n=74 seeds, five capsules);

range 0.98-1.30 mm; SD 0.075. The width of

six seeds was not included because the colour

of five was markedly paler than other seeds

in the capsule and thus were assumed to not

be fully mature and/or possibly not viable;

another seed was inadvertently crushed when

measured.

630

Dispersal Distances

The distances that seeds travelled after being

ballistically ejected from capsule valves are

summarized in Table 4. The mean distance

for all seeds (n=85) was 148 cm; range 13-321

cm; SD 77.76.

The majority of seeds were dispersed a

distance of 101 to 200 cm (Fig. 10). Two seeds

exceeded 300 cm (306 and 321 cm).

Austrobaileya 8(4): 624-633 (2012)

Table 3. Numbers of seeds per capsule in

Viola betonicifolia

Capsule No.

Number of seeds

35 a

a Determined by first author; all others by second author.

Table 4. Seed dispersal distances in Viola betonicifolia

Dispersal

Date

Temp

(°C)

Total number

of seeds per

capsule

Mean distance/

Range/ Standard

Deviation (SD)

Distances (cm)

19 February

2012

n/a

78.04 cm J

17-154.5 cm; SD

41.36

17, 33.5, 37.5, 50, 72, 76.5, 95, 98.5,

106, 118, 154.5

22 February

2012

28.5

140 cm/13-256

cm; SD 66.36

13, 13.5, 56, 85, 90.5, 105, 132.5,

134.5, 139.5, 157, 159, 168, 174.5,

175, 175.5, 188.5,210, 226.5, 256

1 March

2012

31.5

158.02 cm/ 27-

253 cm; SD 67.85

27, 52.5, 55.5, 62, 79, 93.5, 104, 128,

133.5, 146.5, 161, 180, 188, 191.5,

198.200.5, 204,211.5,213,213,

224.5, 226, 247, 253

2 March

2012

181.16 cm/22.5-

321 cm; SD 78.62

22.5, 83, 103, 141.5, 144, 150, 166.5,

177, 216, 217.5, 218, 226, 236.5, 295,

321

8 March

2012

161.06 cm J 20-

306 cm; SD 97.45

20, 33, 49.5, 76, 96.5, 106.5, 121.5,

141, 152.5, 177.5, 208.5,215, 284,

294, 295.5, 306

Discussion

Cleistogamous flowers (CL)

We documented the presence of CL flowers in

Viola betonicifolia. The presence or absence of

CL flowers in this species is not mentioned in

most Australian floras or guidebooks (Adams

1982; Stanley & Ross 1983; James 1990a,b;

Robinson 1994; Entwisle 1996; Fairley &

Moore 2000; Anon 2007; Duretto 2009;

Elliot & Jones 2010). While not specifically

mentioning cleistogamous flowers in V

betonicifolia , Williams (1979) referenced the

presence of “self-pollinating flowers”. Of the

73 Viola species in North American, 60 are

known to produce CL flowers, nine species do

not, and the condition in four species remains

unknown (Little & McKinney in prep). The

number of Viola species in the Australian

flora that produce CL flowers has not been

determined.

The fact that all CL capsules examined

in this study contained mostly fertile seeds

suggests that the pollination mechanism in

CL flowers of Viola betonicifolia is highly

efficient. Mayers & Lord (1983a,b) reported an

interesting situation for V. odorata L. where

the pollen grains in CL flowers germinate

while still in the undehisced anther sacs; the

Little & Lei per, Capsule dehiscence in Viola betonicifolia

631

-D

1-25 26-100 101-200 201-300 300+

Distances (cm)

Fig. 10. Dispersal distance frequency histogram (n=84) for Viola betonicifolia

pollen tubes then penetrate the sac and grow

out towards the nearby stigma. Karlsson et al.

(2012) stated that preliminary observations

in other Viola species suggest that pollen is

simply released in close proximity to the

stigma on the recurved style. The mechanism

by which CL flowers of V betonicifolia achieve

self-pollination is unknown and needs study.

Capsules

Observations of mature capsules of Viola

betonicifolia and other Australian species by

the authors, and unpublished observations

by the first author on several Californian

species, show that the capsules of putatively

diplochorous Viola species do not open

‘explosively,’ but instead open rather slowly.

Based on this study and observations of other

Viola species, the capsule itself does not

‘explode.’

Capsules of Viola species that disperse

seeds ballistically are usually on erect

peduncles, whereas capsules that passively

release their seeds (e.g. V odorata ) usually

point downward (Beattie & Lyons 1975).

Capsules of V betonicifolia are on erect

peduncles prior to seed dispersal, typical

of diplochorous species that eject seeds

ballistically. Upon drying, contraction of

each of the three capsule valves squeezes the

seeds, which are then ballistically ejected.

We observed that V betonicifolia seeds were

sometimes ejected before the valves were

completely open. This occurrence needs to be

taken into account when determining the total

number of seeds per valve and per capsule.

The morphology of mature capsules of

CH and CL flowers are indistinguishable.

Only CL capsules were available for this

study. Dispersal in CH capsules has not been

studied. However, based on observations

of CH capsules of other Viola species in

Australia and California, we suspect that

CH capsules of V betonicifolia dehisce in a

manner similar to CL capsules. Although

empirical data are not available, it seems

plausible that temperature, humidity, and soil

moisture are important factors in the length of

time it takes for a capsule to change from an

inverted to an upright position, to split open,

and to begin dehiscing seeds.

How long does it take a mature capsule to

open? Based on one capsule that was timed, it

took 1.5 hr to fully open. Although only one

632

capsule was timed, it was observed that all

capsules opened slowly. None ‘exploded’ in a

spontaneous release of seeds. After a capsule

is fully open, how long before seeds begin to

disperse? Data from two capsules showed that

one took 2.3 hr while another took 40 minutes

to disperse all their seeds. Are seeds randomly

dispersed from a valve or is there a pattern?

We observed that seeds were dispersed first

from the centre and then randomly from

either the proximal or distal ends of the valve.

Beattie & Lyons (1975) observed that seeds

of Viola species they studied were usually

dispersed first from the centre of the valve.

Seeds and Dispersal Distances

The position of the individual valves relative

to ground level prior to dehiscence affects the

trajectory and dispersal distance for a given

seed. The seed dispersal distances that we

report are measures of the horizontal distance

from the plant and may not reflect the actual

distance travelled. For example, a seed could

be ejected 200 cm vertically, but land only 20

cm from the plant. The dispersal distance in

this case would be measured as 20 cm. We

did in fact observe that some seeds were

expelled vertically while others were expelled

± horizontally. Further research is needed to

determine if position in the valve correlates

with the distance a seed is ejected, other

factors being equal.

The mean dispersal distance for all Viola

betonicifolia seeds was 149 cm (n=84), range

13-321 cm. The frequency histogram of

dispersal distances approximates a bell curve

(Fig. 10). Dispersal distances of seeds from

V betonicifolia CH flowers remain to be

determined.

Beattie & Lyons (1975) reported distances

of ballistically dispersed seeds for CH

capsules of seven perennial Viola species

native to eastern North America. They also

reported data for CL capsules for three of

the seven species they studied (V striata

Aiton, V blanda Willd. and V papilionacea

= V. sororia Willd. var. sororia). Because

dispersal distances of seeds from CH capsules

were longer for each species compared to CL

capsules, a reasonable comparison of Beattie

Austrobaileya 8(4): 624-633 (2012)

& Lyons’ (1975) data with V betonicifolia was

made by considering only the results from CL

capsules: V striata , mean dispersal distance

was 110 cm (n=50), range 20-220 cm; V

blanda , mean dispersal distance was 80 cm

(n=18), range 30-220 cm; and Vpapilionacea ,

mean dispersal distance was 100 cm (n=527),

range 2-210 cm. The mean dispersal distance

of V betonicifolia seeds from CL capsules

(149 cm) exceeded the three species reported

by Beattie & Lyons (1975) by 110, 80, and

100 cm. In addition, the maximum dispersal

distance of V betonicifolia (321 cm) exceeded

the three species reported by Beattie & Lyons

(1975) of 220, 220, and 210 cm.

Forster (2007) documented that

ballistically ejected seeds of Euphorbia

obesa Hook. (Euphorbiaceae) were viscid

resulting in their sticking to soil, pebbles

and vegetation. We found Viola betonicifolia

seeds were buoyant in water, but not viscid

and thus do not stick to surfaces. In coastal

areas of Queensland, V betonicifolia grows

with Melaleuca quinquenervia (Cav.)

S.T.Blake in swampy areas that become

seasonally or periodically inundated from

rainwater. In addition to ants, seed dispersal

may be facilitated by being buoyant and

transported in water during storm events. The

potential that dispersal of Viola seeds can be

facilitated by water has not been investigated.

Anecdotal observations of V betonicifolia

plants growing in Tines’ in coastal wetlands

in Queensland where water has obviously

carried them and receded (pers. comm., F.

Jordan), suggests that water dispersal may

be an important, heretofore overlooked

mechanism for dispersal of this species in

Australia.

Acknowledgements

The authors appreciated the comments of K.R.

Thiele on an earlier draft and encouragement

of P.I. Forster to conduct additional research.

Little & Lei per, Capsule dehiscence in Viola betonicifolia

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440-444. Lothian Books: Melbourne.

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Sydney District. An Identification Guide

(Revised Edition), p. 84-85. Kangaroo Press:

Kenthurst.

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(Euphorbiaceae). Euphorbia World 3: 26-30.

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Plants of Colorado. Reprinted for Grove Press

by University Microfilms International: Ann

Arbor, Michigan.

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-(1990b). Violaceae. In G.J.Harden (ed.). Flora of

New South Wales 1: 435-441. Royal Botanic

Gardens: Sydney.

Karlsson, T., Marcussen, T., Wind, P. & Jonsell, B.

(2012). Violaceae. In Nordic Flora , version

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A revision of Calyptochloa C.E.Hubb. (Poaceae),

with two new species and a new subspecies

E.J. Thompson & B.K. Simon

Summary

Thompson, E.J. & Simon, B.K. (2012). A revision of Calyptochloa C.E.Hubb. (Poaceae), with two

new species and a new subspecies. Austrobaileya 8(4): 634-652. Two new species of Calyptochloa

C.E.Hubb. ( Calyptochloa cylindrosperma E. J.Thomps. & B.K.Simon and C. johnsoniana E. J.Thomps.

& B.K.Simon) endemic to central Queensland, and a new subspecies of Calyptochloa gracillima

C.E.Hubb. (C. gracillima subsp. ipsviciensis E.J.Thomps. & B.K.Simon) endemic to southeast

Queensland are described and illustrated.

Key Words: Poaceae, Paniceae, panicoid, cleistogamous, Calyptochloa, Calyptochloa

cylindrosperma, Calyptochloa gracillima subsp. gracillima, Calyptochloa gracillima subsp.

ipsviciensis, Calyptochloa johnsoniana, Cleistochloa, Queensland flora, taxonomy, new species, new

subspecies, identification key

E.J. Thompson, c/o Queensland Herbarium, Department of Science, Information Technology,

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

Australia.

B .K. Simon, c/o Queensland Herbarium, Department of Science, Information Technology, Innovation

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

Introduction

Calyptochloa C.E.Hubb., an endemic

Australian genus, is placed in the subfamily

Panicoideae Link, tribe Paniceae R.Br. This

tribe is characterised by the spikelets having

a pair of dimorphic florets with the lower

often incomplete, male or sterile, and falling

entire, the upper fertile, and by the relative

induration of the glumes and lemmas (Clayton

& Renvoize 1986; Kellogg & Campbell 1987).

Calyptochloa is amphigamous by having two

types of inflorescences, viz. in terminal and

axillary positions. The terminal inflorescence

(Connor 1979), is a spike-like raceme with

chasmogamous (CH) pedicillate spikelets

that open at maturity and thereby potentially

cross-fertilise. Conversely, the axillary

inflorescence usually consists of a single

sessile cleistogamous (enclosed self-fertilising

flowers) (CL) spikelet which is hidden within

semi-woody to woody leaf sheaths at each of

several nodes along the culm. In the summer

wet season, the axillary spikelets are produced

at nodes with the terminal inflorescence

above. At other times of the year, these chains

Accepted for publication 20 August 2012

of axillary spikelets may be produced in the

absence of terminal inflorescences. Webster

(1987) stated that the CL spikelets occur

singly or in pairs but we have not observed

paired spikelets in any specimens at BRI,

including those cited by Webster (1987), until

we examined the type specimen for one of the

new species described herein (Calyptochloa

johnsoniana E.J.Thomps. & B.K.Simon). In

Calyptochloa , the CL spikelets are obligately

self-fertilised and never open. Plants of

Calyptochloa retain the CL spikelets for a few

months enclosed in the leaf sheaths before

disarticulation at the culm nodes or at the leaf

sheath bases which then fall at maturity with

subsequent dispersal of the caryopses.

Calyptochloa has remained a monotypic

genus since description with only C. gracillima

C.E.Hubb. recognised until now (Hubbard

1933b; Tothill & Hacker 1983). The genus is

characterised by the perennial mat-forming

growth habit and the fertile leaf sheaths

which enclose the CL spikelets. Clifford &

Ludlow (1972) differentiated Calyptochloa

from other Queensland grass genera in their

key using “stems disarticulating at the nodes

at maturity” and “prostrate to creeping” habit.

Thompson & Simon, Calyptochloa

The genus is both clonal (stoloniferous) and

cleistogamous, a rare combination in grasses

(Campbell et al. 1983).

Cleistochloa C.E.Hubb. (Hubbard

1933a), another perennial panicoid genus

from Australia and New Guinea, was

listed by Connor (1981) with Calyptochloa

amongst 13 genera world wide that possess

clandestine axillary CL spikelets and belong

in four different subfamilies of the Poaceae.

Seven of these genera have amphigamous

inflorescences and dimorphic spikelets of

which Calyptochloa and Cleistochloa are

the only panicoid genera. Dimorphochloa

S.T.Blake (Blake 1941; Simon et al. 2010),

which is also an Australian CL panicoid

genus, was correctly omitted from Connor’s

(1981) list taking into account that this genus

had not been synonymised with Cleistochloa

(Clayton & Renvoize 1986; Webster 1987)

at the time. Although Dimorphochloa has

amphigamous inflorescences it differs from

these other genera in terms of the CL spikelets

as follows: similar to the CH spikelets,

located apically on branchlets below the

terminal inflorescences, and not hidden in

the leaf sheaths at anthesis. Amphicarpum

Kunth, another CL panicoid genus from

eastern North America, was also omitted

from Connor’s (1981) list. Amphicarpum has

amphigamous inflorescences and dimorphic

spikelets but differs by the subterranean CL

spikelets (rhizanthogenes) which are borne at

the tips of rhizomes.

Connor (1981) reported that the

clandestine spikelets are a secondary source

of seed with most of the seed produced in the

terminal inflorescences. For the Australian

genera the reverse is true with most or all of

the caryopses produced in the CL spikelets.

Of about 30 specimens of Calyptochloa

gracillima possessing terminal inflorescences

inspected at BRI, only one had CH caryopses.

No specimens of Cleistochloa at BRI were

observed to have CH caryopses, thereby

confirming this same observation made by

Hubbard (1933a). The Australian genera

with clandestine spikelets share features,

indicative of obligate or habitual cleistogamy

(Connor 1979), which when compared to the

635

CH spikelets (Campbell et al. 1983) include

the following:

a) reduced CL inflorescence size, usually one

spikelet compared to a raceme or reduced

panicle

b) CL lodicules absent

c) reduced size of CL anthers usually

enravelled in reduced styles

d) upper floret with lemma and palea

convolute towards the apex tightly

enclosing the anthers and styles at anthesis

compared to gaping, and

e) the CL caryopses a little larger than the

CH caryopses when present.

Campbell et al. (1983) provided a detailed

classification of CL species comprising

four types based on factors that relate to

prevention of the spikelets from opening

including leaf sheath, spikelet parts or the soil

conditions. Campbell et al. (1983) classified

Calyptochloa and Cleistochloa as type II

where fertilisation occurs in spikelets hidden

in the lowermost sheaths and this type is

usually associated with major inflorescence

and spikelet differentiation. Chase (1908)

referred to these clandestine CL spikelets at

or near the ground as cleistogenes. However,

Calyptochloa and Cleistochloa have CL

spikelets enclosed in the sheaths in upper

axils at fertilisation and the upper floret has

modifications including revolute lemma and

palea, and lodicules are absent, which prevent

the floret from opening. These characteristics

match type 1 of Campbell et al. (1983), where

fertilisation takes place within the leaf sheaths

of the middle to upper part of the stem but the

spikelet may be exserted at maturity.

Hubbard (1933a) stated that as for the

American CL grasses, the Australian species

are found in arid regions or dry places within

humid regions. Calyptochloa is distributed

from tropical central Queensland with hot

humid summers and monsoonal wet season

to warm temperate south-eastern Queensland

with warm humid summers (Map 1).

Calyptochloa spp. are found in mostly well

shaded habitats in a variety of vegetation

communities frequently dominated by Acacia

spp. on gently undulating to steeply sloping

636

Austrobaileya 8 ( 4 ): 634-652 ( 2012 )

terrain with shallow to skeletal soils derived

from a variety of geology but often on

landscapes with lateritic profiles.

In the current paper we provide a taxonomic

account of Calyptochloa , trebling the number

of species. Some of these additional species

have been recognised for some time; however,

their description is now possible following

collection of material critical for character

delimitation. Other taxa currently listed under

Calyptochloa include C. sp. (Charters Towers

E. J.Thompson+ CHA554) (Simon et al. 2010)

and C. sp. (Duaringa K.D.Addison 42) (in BRI

HERBRECS database accessed July 2012);

both have same similar features to the species

described in this paper, but may ultimately

be described in other genera. These taxa are

new members of the group of Australian

panicoid grasses with axillary CL spikelets

and are the subject of further study. They have

overlapping distribution and habitat to other

members of the group and often occur with

Thyridolepis xerophila (Domin) S.T.Blake

which is also a CL panicoid grass but the CL

spikelets are in the terminal inflorescences

and this species lacks axillary spikelets. On

a number of occasions up to three to four of

these CL species have been observed growing

together.

Materials and methods

Morphological data were obtained from

dried herbarium material at BRI, and from

cultivated plants transplanted from the field.

Numerous terminal spikelets and leaf sheaths

were dissected to examine the contents and

describe the characteristics of the spikelets.

Caryopsis germination trials were conducted

during one summer over a two month

period using sealable containers in outdoor

conditions with periods of direct sunlight and

no artificial lighting, shade or heating.

Habitat descriptions provided include

Regional Ecosystems (REs) which are defined

by DERM (2011). Botanical terminology

follows Beentje (2010). Common abbreviations

used in specimen citations include N.R

(National Park), S.F. (State Forest).

Taxonomy

Calyptochloa C.E.Hubb., Hook. Icon. PI. 33:

t. 3210 (1933). Type species: C. gracillima

C.E.Hubb.

Decumbent mat forming perennials; rhizomes

absent. Stolons wiry, c. 1 mm thick; mid-culm

internodes hollow. Culms differentiated,

sterile and fertile, ascending from stolons.

Fertile culms preceded by a portion of sterile

culm; disarticulating at nodes or retained.

Leaves ultimately disarticulating; margin

undulate on one side, thickened, scabrid,

white, with scattered tubercle-based hairs to

4 mm long at least at base; adaxial surface

usually with scattered to moderately dense,

erect simple hairs; abaxial surface with

moderately dense, erect simple hairs. Mature

fertile leaf sheaths disarticulating or retained,

semi-woody to woody, enclosing from c. half

to most of the length of the internode with

scattered appressed to ascending tubercle-

based bristles between ribs, with or without

simple hairs; outer margin with dense, simple

appressed to ascending simple hairs. Fertile

culm internodes retained within leaf sheaths

or bowing and protruding, scabrid along ribs

with occasional simple hairs to 0.5 mm long

between ribs. Sterile leaf sheaths retained;

usually two types of hairs, with scattered

appressed to ascending stiff tubercle-based

hairs and sometimes ascending simple hairs.

Sterile culm internodes with moderately

dense to dense appressed to ascending,

normal to flagelliform simple hairs to 2 mm

long between ribs. Ligule a fringe of hairs,

c. 0.3 mm long. Inflorescences of two kinds,

chasmogamous terminal and cleistogamous

axillary. Terminal inflorescences spike-like.

Spikelets appressed to rachis, pedicillate,

adaxial, elliptic, dorsiventrally compressed.

Lower glume flat, chartaceous, glabrous

except at base, apex acute; frequently absent,

if present then restricted to apical spikelets.

Upper glume as long as spikelet, ovate, flat,

chartaceous, 5-nerved, dense simple hairs at

base and usually moderately dense simple

hairs to 2 mm over lower 30 to 60% and most

of margin, upper portion glabrous; apex acute

to truncate. Rachilla inconspicuous between

florets. Lower floret sterile; lemma ovate,

flat, chartaceous, densely hairy with simple

Thompson & Simon, Calyptochloa

hairs at base and moderately hairy over lower

60 to 80%, upper portion glabrous, margin

moderately hairy with hairs to 2 mm long;

apex acute to obtuse. Palea absent. Upper

floret fertile, shorter than the lower and

slightly indurated; lemma ovate in dorsiventral

view, convolute, chartaceous, glabrous,

3-nerved, apex acute with minutely scabrid

awn; palea ovate, convolute, chartaceous,

glabrous, 2-nerved; apex acute. Lodicules, 2.

Anthers, 3. Caryopsis rarely present. Axillary

inflorescences usually a single cleistogamous

spikelet at 5-10 contiguous culm internodes

often from immediately below terminal

inflorescence; spikelets enclosed within

leaf sheaths which are scarsely enlarged to

conspicuously swollen towards the base where

the walls are thicker, semi-woody to woody.

Spikelets sessile, adaxial, narrow elliptic in

dorsiventral view, slightly indurated. Lower

glume absent. Upper glume lanceolate, flat,

shorter than spikelet, chartaceous, glabrous

except for base with scattered short simple

hairs, 3-nerved; apex acute to truncate.

Rachilla inconspicuous between florets.

Lower floret sterile; lemma elliptic, boat¬

shaped, two-keeled, chartaceous, glabrous

except for base, 5-nerved; apex obtuse.

Palea absent. Upper floret fertile, more than

c. 80% of length of first; lemma lanceolate,

convolute, chartaceous, glabrous, apex acute

with minutely scabrid awn; palea lanceolate,

convolute, chartaceous, glabrous, obscurely

5-nerved; apex acute to shortly awned.

Lodicules absent. Stamens 3. Caryopsis tan

637

to light brown, shallowly grooved at least on

lower half, on adaxial face; hylum broadly

elliptic, c. 40% of caryopsis length.

Notes: Calyptochloa differs from the other

Australian cleistogamous panicoid genera

by having terminal spikelets dorsi-ventrally

compressed compared to spikelets elliptic in

cross-section; the upper floret of the terminal

spikelets about 60 to 70% of the spikelet length

compared to equal to the spikelet length;

the axillary spikelets retained within semi-

woody to woody leaf sheaths compared to the

spikelets exposed, partially hidden or hidden

within cartilaginous leaf sheaths; axillary

spikelets lacking spongy tissue at the base of

the lower lemma; axillary caryopsis grooved

on the adaxial face compared to face convex;

and differential indumentum type on the

sterile and fertile culm internodes compared

to little or no difference in the indumentum

types.

Preliminary results from caryopsis

germination and seedling trials for most of the

Calyptochloa spp. recognised here, indicate

some variation in dormancy, cotyledon

characters and seedling survival. Germination

for the trial was sporadic but frequently

temporally clustered giving an impression

that dormancy may be broken by a period of

several hot days. Seedling survival was poor

for most taxa suggesting that survival may be

affected by nutrient status and/or acidity of the

potting medium and is potentially dependent

on mycorrhiza. Investigations are continuing

into these aspects.

Key to Calyptochloa species

1 Fertile culm internode bowed and protruding from leaf sheath with

chartaceous margins; axillary spikelet with upper glume >4.8 mm

long; upper glume of terminal spikelets scabrid in mid third portion . .3. C. johnsoniana

1. Fertile culm internode retained within leaf sheath with margins semi-

woody to woody; axillary spikelet with upper glume <4.5 mm long;

upper glume of terminal spikelets sparsely hairy to pilose with simple

hairs to 1 mm long in mid third portion.2

2 Lower portion of fertile leaf sheath conspicuously swollen to 2.7 mm wide,

wall 0.3-0.5 mm thick; axillary spikelets 3.5-5.5 mm long (excluding

awn); terminal spikelets 3-4.6 mm long (excluding awn) 1. C. gracillima

2. Lower portion of fertile leaf sheath slightly swollen to 1.4 mm wide, wall

0.2-0.3 mm thick; axillary spikelets 6-7.5 mm long (excluding awn);

terminal spikelets 5-6 mm long (excluding awn).2. C. cylindrosperma

638

1. Calyptochloa gracillima C.E.Hubb.,

Hook. Icon. PI. 33: t. 3210, 1-6 (1933). Type:

Queensland. Burnett District: Munduberra,

April 1931, H.S.Bloxsome 9 (holo: BRI; iso:

BRI, K [photo BRI]).

Decumbent stoloniferous perennial.

Ascending branches to 40 cm tall, copiously

branched with 7-30 nodes. Stolons to c. 2 m

long. Mid-culm leaf blades 12-40 mm long,

2.5-6 mm wide; adaxial surface with sparse

hairs 0.5-2 mm long; abaxial surface with

moderately dense simple hairs 0.5-1 mm

long. Mature fertile leaf sheaths retained,

convolute, woody. Fertile culm internodes

14-40 mm long. Sterile leaf sheaths with or

without tubercle-based bristles 0.3-0.8 mm

long and occasionally simple hairs 1.5-3 mm

long; outer margin hairs dense, 0.4-1 mm

long. Terminal inflorescences on axes 1.5-3

cm long, 5-8-flowered. Spikelets 2.3-5 mm

long (without awn), 1-1.8 mm wide; lateral

Austrobaileya 8(4): 634-652 (2012)

pedicels 0.3-1.6 mm long; ultimate pedicel

2.5-5.5 mm long. Lower glume triangular

to lanceolate, 0.2-1.8 mm long; apex acute.

Upper glume 3-5 mm long. Lower lemma

2.3-5 mm long; apex acute. Upper lemma

2.2-3.5 mm long, awn 0.5-3 mm long;

lodicules c. 0.2 mm long; palea 2-3 mm

long, rarely awned. Anther 1.5-2 mm long.

Caryopsis (1.6-1.8) 2.2-2.5 mm, rarely

present. Axillary inflorescences present at

3-10 internodes. Spikelets 3.5-5.5 mm long

(without awn), 0.8-1.1 mm wide. Upper

glume 0.5-3.5 mm long, apex acute. Lower

lemma 3-5.5 mm long. Upper floret subequal

to lower. Upper lemma body 3.5-5.5 mm

long, awn 0.5-2.6 mm long; palea 3-3.8 mm

long. Anthers 0.3-0.7 mm long. Caryopsis

approximately plano-convex, 2-3.5 mm long,

0.7-0.8 mm wide. Measurements in bold

type are from Hubbard (1933b) and were not

repeatable from the specimens examined.

Key to subspecies of Calyptochloa gracillima

la Axillary spikelets 4-5.5 mm long (excluding awn) x 1-1.1 mm

wide, anthers 0.3-0.4 mm long; terminal spikelets with lower

glume when present c. 0.2 mm long and upper glume

apex obtuse to truncate.C. gracillima subsp. gracillima

lb Axillary spikelets 3.5-4.2 mm long (excluding awn) x 0.8-0.9 mm wide,

anthers 0.4-0.7 mm long; terminal spikelets with lower glume when

present 0.8-1.8 mm long and upper glume apex acute . . C. gracillima subsp. ipsviciensis

la. C. gracillima subsp. gracillima

Decumbent stoloniferous perennial. Ascending

branches to 25 cm tall, copiously branched

with 10-30 nodes. Stolons to c. 1.5 m long.

Mid-culm leaf blades 25-40 mm long, 2.5-5

mm wide; adaxial surface with sparse to

moderately dense simple hairs 0.3-1.6 mm

long and usually some tubercle based hairs

to 3 mm long on margin at base; abaxial

surface with moderately dense simple hairs

0.2-0.8 mm long. Mature fertile leaf sheaths

10-17 mm long, 1.5-3 mm wide near base

with wall 0.3-0.4 mm thick. Sterile leaf

sheaths with tubercle-based bristles c. 0.3

mm long and simple hairs c. 1.3 mm long.

Terminal inflorescences on axes 1-3 cm

long, 5-8-flowered. Spikelets 3-5 mm long

(without awn), 1.3-1.8 mm wide; lateral

pedicels 0.4-2 mm long, apical pedicel

2- 4.5 mm long. Lower glume triangular to

lanceolate, 0.2-1.3 mm long. Upper glume

3- 5 mm long; apex truncate. Lower lemma

3-5 mm long. Upper lemma 3-3.5 mm long,

awn 2-3 mm long; lodicules 0.2-0.4 mm

long; palea 2.5-3 mm long, rarely awned,

awn to 2 mm long. Anthers (0.5-1) 1.6—2 mm

long. Caryopsis (1.6-1.8) c. 2.3, rarely seen.

Axillary inflorescences usually present at 5

(3-10) internodes. Spikelets 4-5.5 mm long

(without awn), 1-1.1 mm wide. Upper glume

0.5-1.5 mm long. Lower lemma 4-5.5 mm

long. Upper lemma body 4-5.5 mm long, awn

2-2.6 mm long; palea 3-3.8 mm long. Anthers

0.3-0.4 mm long. Caryopsis 2-3.5 mm long,

0.7-0.8 mm wide. Fig. 1 & 2.

Thompson & Simon, Calyptochloa

639

Fig. 1 . Axillary spikelet of Calyptochloa gracillima subsp. gracillima. A. leaf sheath enclosing axillary spikelet x8.

B. upper glume facing *12. C. side view xl2. D. lower lemma facing xl2. E. upper glume xl2. F. lower lemma xl2. G.

cross-sectional view of lower lemma xl2. H. upper lemma xl2.1. upper palea xl2. J. caryopsis xl6. K. cross-sectional

view of caryopsis xl6. A-K from Blake 19976 (BRI). Del. W. Smith.

640

Austrobaileya 8(4): 634-652 (2012)

Fig. 2. Terminal spikelet of Calyptochloa gracillima subsp. gracillima. A. upper glume facing xi6. B. side view xl6.

C. lower glume *24. D upper glume xl6. E. lower lemma facing xl6. F. upper lemma xl6. G. upper palea xl6. H.

caryopsis xl6. I. cross-sectional view of caryopsis xl6. A-F from Blake 19976 (BRI); H-I from Bean 20216 (BRI).

Del. W.Smith.

Measurements in bold type are from Hubbard

(1933b) which were not repeatable from the

specimens examined.

Additional selected specimens examined : Queensland.

North Kennedy District: On edge of road 70 km SSE of

Charters Towers, May 2012, Thompson & Simon CHA795

(BRI). South Kennedy District: Edge of highway, 53

km NW of Clermont, May 2012, Thompson & Simon

EJT875 (BRI); 4 km (direct) NW of haul road overpass,

near Newlands coal mine, WNW of Glendon, Jun 2009,

Bean 29028 , (BRI). Leichhardt District: Edge of road,

34 km SW of Springsure, Apr 2012, Thompson & Simon

EJT830 (BRI); site of Brigalow Research Station, 20

miles [32 km] NW of Theodore, Apr 1963, Johnson 2642

(BRI); 17 km W of Baralaba, on road to Woorabinda,

Mar 2005, Bean 23519 (BRI); Near Bun Bun Kundoo

Spring, Ka Ka Mundi N.R, via Springsure, May 1999,

Bean 14846 (BRI); 16.6 km along Roche Creek Road, E

of Wandoan, Mar 2010, Bean 29485 (BRI). Port Curtis

District: Gogango, May 1956, Blake 19976 , (BRI);

Marmor, Mar 1943, Blake 14819 (BRI); Hibbs Road,

N of Jambin, Apr 2003, Bean 20216 (BRI). Maranoa

District: 20 miles [32 km] W of Mitchell, Mar 1936,

Blake 10951 (BRI). Darling Downs District: Edge of

track, Barakula S.F., 32 km NW of Chinchilla, Apr 2012,

Thompson & Simon EJT786 (BRI).

Distribution and habitat : Calyptochloa

gracillima subsp. gracillima is endemic to

central Queensland (Map 1). At its most

southern limits, it occurs on a range of soil

types e.g. clay under brigalow (Acacia

Thompson & Simon, Calyptochloa

harpophylla F.Muell. ex Benth.) (RE 11.3.1),

sandy duplex soils to skeletal soils on laterite

and shallow sandy soils on sandstone in

ironbark woodland (commonly Eucalyptus

fibrosa subsp. mibila (Maiden & Blakely)

L.A.S.Johnson) (RE 10.7.7). Other REs

represented include 11.5.3 and 11.5.4. Further

north it occurs on mostly lateritic landscapes

overlapping with the distribution area of C.

cylindrosperma but the two species are rarely

seen together. REs represented include 11.7.2

and 11.7.6. C. gracillima subsp. gracillima

has a much broader habitat range than C.

cylindrosperma , C. johnsoniana and C.

gracillima subsp. ipsviciensis, which is also

reflected in its broader overall distribution.

Phenology : Calyptochloa gracillima subsp.

gracillima flowers from December to March

during the wet season. The cleistogamous

spikelets are produced over a broader seasonal

period.

Notes : Caryopsis germination trials indicate

differences between the subspecies of

Calyptochloa gracillima. Initial trials have

revealed more rapid germination of C.

gracillima subsp. ipsviciensis and better

seedling survival than for the nominative

subspecies.

Conservation status : This subspecies is

widely distributed over a large area and is

usually common in the habitats where it

occurs suggesting this subspecies is Least

Concern (IUCN 2001).

lb. Calyptochloa gracillima subsp.

ipsviciensis E.J.Thomps. & B.K. Simon,

subspecies nova similar to C. gracillima

C.E.Hubb. subsp. gracillima differing by

the axillary spikelets mostly shorter (3.5-4.2

mm versus 4-5.5 mm) and narrower (0.8-

0.9 mm versus 1-1.1 mm); longer anthers

(0.4-0.7 versus 0.3-0.4); and by the terminal

spikelets with an acute apex of upper glume

(versus obtuse to truncate), and longer lower

glumes when present (0.8-1.8 mm versus

<0.2 mm). Typus: Queensland, Moreton

District: Council reserve, cnr Reservoir

Lane and Kholo Road, Ipswich, 4 April

2012, E.J.Thompson MOR711 (holo: BRI; iso:

CANB, K, L, MO, NSW, SI, US).

641

Decumbent stoloniferous perennial.

Ascending branches to 40 cm tall, copiously

branched with 10-30 nodes. Stolons to c. 3 m

long. Mid-culm leaf blades 20-36 mm long,

2.5-5 mm wide; adaxial surface with sparse

hairs 0.5-2 mm long; abaxial surface with

moderately dense simple hairs 0.5-1 mm long.

Mature fertile leaf sheaths 10-15 mm long,

1.2- 2.5 mm wide near base with wall 0.3-0.4

mm thick. Sterile leaf sheaths with tubercle-

based bristles 0.3-0.7 mm long and simple

hairs 1.5-3 mm long. Terminal inflorescences

on axes 1.5-3 cm long, 5-8-flowered.

Spikelets 3-4.6 mm long (without awn), 1-1.6

mm wide; lateral pedicels 1-1.6 mm long,

apical pedicel 2.5-4 mm long. Lower glume

lanceolate, 0.7-1.8 mm long. Upper glume

2.3- 4.6 mm long; apex acute. Lower lemma

2.3-4.6 mm long. Upper floret lemma 2.2-3.2

mm long, awn 0.5-2.4 mm long; lodicules 0.2

mm long; palea 2-2.7 mm long, apex acute.

Anther, 1.5-2 mm long. Caryopsis not seen.

Axillary inflorescences usually present at 4

(3-5) internodes. Spikelets 3.5-4.2 mm long

(without awn), 0.8-1 mm wide. Upper glume

0.7-3.5 mm long. Lower lemma 3.5-4.2 mm

long. Upper lemma body 3-4.2 mm long, awn

0.5-2.5 mm long; palea 27-3.5 mm long.

Anthers 0.4-0.5 mm long. Caryopsis 2.3-37

mm long, 0.5-0.8 mm wide. Fig. 3 & 4.

Additional specimens examined : Queensland.

Moreton District: Edge of powerline easement off

South Deebing Creek Road, Deebing Heights, Feb 2012,

Thompson MOR689 & Simon (BRI, CANB, K, SI); Edge

of Kerners Road, Yamanto near Ipswich, Aug 2011,

Thompson EJT497 (BRI, CANB, MO); Edge of Kerners

Road, Yamanto near Ipswich, Feb 2012, Thompson

MOR688 & Simon (BRI); Council reserve, corner

Reservior Lane and Kholo Road, Ipswich, May 2002,

Thompson MOR739 & Simon (BRI): Ipswich Council

reserve, end of Powers Road, off Kholo Road, c. 1 km S

of Brisbane River crossing, c. 6 km N of Ipswich; Mar

2012, Thompson MOR709 (BRI); Edge of Kholo Road,

c. 1 km SE of Brisbane River crossing near corner of

Blackwall Road, c. 6 km N of Ipswich, Mar 2012,

Thompson MOR693 (BRI, CANB, NSW, RSA).

Distribution and habitat : Calyptochloa

gracillima subsp. ipsviciensis is endemic

to southeast Queensland in the vicinity of

Ipswich (Map 1) where it is known from

a few small areas. It is an uncommon to

dominant species in woodlands dominated by

Eucalyptus spp. including E. crebra F.Muell.

642

Austrobaileya 8(4): 634-652 (2012)

Fig. 3. Axillary spikelet of Calyptochloa gracillima subsp. ipsviciensis. A. habit *0.6. B. leaf sheath enclosing axillary

spikelet x8. C. upper glume facing xl6. D. side view xl6. E. lower lemma facing xl6. F. lower lemma xl6. G. cross-

sectional view of lower lemma xl6. H. upper glume xl6.1. upper lemma xl6. J. upper palea xl6. K. caryopsis xl6. L.

cross-sectional view of caryopsis xl6. A-L from Thompson MOR689 & Simon (BRI). Del. W. Smith

Thompson & Simon, Calyptochloa

643

Fig. 4. Terminal spikelet of Calyptochloa gracillima subsp. ipsviciensis. A. upper glume facing xl2. B. side view xl2.

C. lower glume xl6. D. upper glume xl6. E. lower lemma xl6. F. upper lemma xl6. G. upper palea xl6. H. stamens and

stigmas x!6. A-H from Thompson MOR689 & Simon (BRI). Del. W. Smith.

and E. moluccana Roxb. and/or Corymbia

citriodora subsp. variegata (F.Muell.)

A.R.Bean & M.W.McDonald on loam to clay

loam duplex soils derived from shale on gently

undulating to hilly terrain. REs represented

include 12.9-10.2, 12.9-10.3 and 12.9-10.19.

Associated ground layer species include

Aristida caput-medusae Domin, Cleistochloa

subjuncea C.E.Hubb. and Theme da triandra

Forssk. The habitat is typically moderately

shaded.

Phenology : Calyptochloa gracillima subsp.

ipsviciensis flowers from December to March

during the wet season. The cleistogamous

spikelets are produced over a broader seasonal

period.

Notes: Until 2011 there were no specimen

records of Calyptochloa gracillima at BRI

from the Moreton Pastoral District near

Ipswich. These new records represent a

disjunction of over 200 km from the previous

known southern limit of the species.

Calyptochloa gracillima subsp. ipsviciensis

644

is similar to C. gracillima subsp. gracillima

in growth habit but on average it is taller,

the mats cover a greater area and the leaves

are more yellowish green. C. gracillima

subsp. ipsviciensis also differs by the mostly

thinner walled fertile leaf sheaths, and often

the proportionally shorter fertile leaf sheath

in relation to the internode length. Generally

the fertile leaf sheaths cover about half the

length of the culm internodes whereas for C.

gracillima subsp. gracillima the leaf sheath

usually covers most of the length of the culm

internode. Only spikelets towards the apex

of racemes of terminal inflorescences have a

lower glume present, but it is often absent.

The distribution of this subspecies

overlaps with Ottochloagracillima C.E.Hubb.

and Entolasia marginata (R.Br.) Hughes,

both of which it could easily be confused

with in the field in terms of growth habit

and leaf colour and size although to date

these species have not been seen growing

with C. gracillima. Ottochloa gracillima and

Entolasia marginata are distinguishable in

the field by the branched inflorescences, the

smaller glabrous spikelets and the abaxial leaf

surface which is glabrous to sparsely hairy.

Etymology : The subspecies epithet is derived

in reference to the name of the nearby city of

Ipswich where it has been found.

Conservation status: Calyptochloa gracillima

subsp. ispviciensis is only known from a few

locations near the urban centre of Ipswich, two

of which are Ipswich City Council reserves. At

two locations only one or two plants or mats

have been observed. The very restricted range

and the few small populations suggest this

subspecies should be considered Critically

Endangered (criterion Bla,b [IUCN 2001]).

Current threats include invasion from weeds

such as Megathyrsus maxima var. pubiglumis

(K.Schum.) B.K.Simon & S.W.L.Jacobs

and Lantana montevidensis (Spreng.) Briq.,

inappropriate burning regimes, urbanisation

and road construction.

2. Calyptochloa cylindrospermaE.J.Thomps.

& B.K.Simon, species nova similar to C.

gracillima C.E.Hubb. differing by the degree

of swelling of the mature fertile leaf sheaths

Austrobaileya 8(4): 634-652 (2012)

(slightly versus conspicuous) with thinner

walls (0.2-0.3 mm versus 0.3-0.5 mm); the

longer axillary spikelets (6-7.5 mm versus

3.5-5.5 mm) with longer caryopses (3.8-4

mm versus 2-3.7 mm) and shape (cylindrical

versus plano-convex); the longer terminal

spikelets (5-6 mm versus 3-5 mm) with

longer anthers (2.5-2.6 mm versus 1.6-2

mm) and longer upper glume (5-6 mm versus

2.3-5 mm). Typus: Queensland. North

Kennedy District: 16 km SW of Charters

Towers on edge of road, 7.5 km W of Black

Jack, 30 March 2011, E.J.Thompson CHA769,

B.K.Simon & M.Edginton (holo: BRI; iso:

CANB, K, L, MO, NSW, SI, US).

Calyptochloa sp. (Blackjack E.J.Thompson+

CHA769) (in BRI HERBRECS database

accessed July 2012).

Decumbent stoloniferous perennial. Ascending

branches to 40 cm tall, copiously branched

with 10-30 nodes. Stolons to c. 0.5 m long.

Mid-culm leaf blades 15-30 mm long, 2-4

mm wide; adaxial surface with scattered to

moderately dense simple hairs to 0.5-1.6 mm

long; abaxial surface with moderately dense

simple hairs 1-2 mm long. Mature fertile

leaf sheaths retained, semi-woody, 15-20

mm long, 1.2-1.7 mm wide near base with

wall 0.2-0.3 mm thick; tubercle-based hairs

0.7-1.4 mm long between nerves, simple hairs

absent; outer margin hairs to 1 mm long.

Fertile culm internodes 20-45 mm long.

Sterile leaf sheaths with scattered tubercle-

based hairs 0.4-1 mm long and some simple

hairs 0.5-2 mm long. Terminal inflorescences

on axes 2-5 cm long, 5-10-flowered. Spikelets

5-6 mm long (without awn), 1.5-2 mm wide;

lateral pedicels 0.5-1.5 mm long; ultimate

pedicels 3-8 mm long; ultimate spikelets

frequently longer than basal spikelets. Lower

glume triangular, 0.1-0.5 mm long. Upper

glume 5-6 mm long; apex acute. Lower

lemma 5-6 mm long; apex acute. Upper

lemma body 3-4 mm long, awn to 2.5-4 mm

long. Lodicules c. 0.3 mm long. Upper palea

3-4 mm long. Anthers 2.5-2.6 mm long.

Caryopsis not seen. Axillary inflorescences

enclosed in leaf sheaths within scarcely

enlarged basal portion usually present at 3-5

internodes. Spikelets 6-7.5 mm long (without

Thompson & Simon, Calyptochloa

awn), 07-0.9 mm wide. Upper glume 07-4.5

mm long, apex acute. Lower lemma 6-75 mm

long. Upper floret subequal to lower. Upper

lemma body 6-7.5 mm long, awn 1.5-2.5 mm

long. Palea 5.5-6 mm long; acute to shortly

awned. Anthers c. 0.6 mm long. Caryopsis

cylindrical, 3.8-4 mm long, 0.5-0.6 mm

wide. Fig. 5 & 6.

Additional specimens examined (c. 55 collections

examined): Queensland. North Kennedy District:

near Charters Towers, Apr 1943, Blake 14904 (BRI);

50 km NW of Charters Towers, Dec 2011, Thompson

CHA779 (BRI); 16 km SW of Charters Towers on edge

of road. Mar 2011, Thompson CHA767 et at. (BRI);

16 km W of Charters Towers on edge of Capricorn

Highway, Mar 2011, Thompson CHA773 et al. (BRI);

16 km SW of Charters Towers on road to Jesmond, Mar

2002, Thompson CHA556 & Turpin (BRI, CANB, RSA);

20 km SW of Charters Towers, May 2012, Thompson

CHA786 & Simon (BRI, CANB, K); 15 km NE of Mt

Cooper Homestead, Jun 1992, Thompson CHA332 &

Sharpe (BRI); 88 km SE of Charters Towers, May 2012,

Thompson CHA801 & Simon (BRI, CANB, K). South

Kennedy District: (site plot 53) 8.5 km SW of Mt Hope

Homestead, Apr 1992, Thompson BUC508 & Simon

(BRI); Blackwood N.P., 160 km S of Charters Towers,

Mar 1998, Gumming 16888 (BRI); ditto loc ., Dec 2011,

Thompson CHA776 (BRI).

Distribution and habitat: Calyptochloa

cylindrosperma is known from central

Queensland near Charters Towers (Map 1). It

usually grows as the dominant ground cover,

commonly in woodland of Acacia shirleyi

Maiden, with or without A. catenulata

C.T.White, on lateritic landscapes on mostly

Tertiary plateaux with gently undulating red

soil, occasionally jump-ups with shallow

soils, or sometimes on shallow soils in

sheltered gullies on quartzose sandstone.

Associated ground layer species include

Cleistochloa subjuncea, Thyridolepis

xerophila and Aristida caput-medusae.

Regional Ecosystems represented include

10.7.3a and b, and 11.7.2. The habitat is

typically well shaded.

Phenology: Calyptochloa cylindrosperma

flowers from December to March during the

wet season. Axillary spikelets are produced

over a broader seasonal period.

Notes: Calyptochloa cylindrosperma is

similar to C. gracillima in growth habit (Table

1) and they have been found growing together

at the transition of habitat from skeletal soil to

645

deep red soil with the latter habitat occupied

by C. cylindrosperma.

Only spikelets towards the apex of racemes

of terminal inflorescences have a lower glume

present; however, it is often absent.

Calyptochloa cylindrosperma also has an

overlapping distribution and shares habitat

with Calyptochloa sp. (Charters Towers

E.J.Thompson+ CHA554). This latter species

differs by the terminal spikelets having the

upper glume and lower lemma convex in

cross-section and the upper floret as long as

the spikelet, and the axillary inflorescences

consisting of two types, one with paired

spikelets, sessile and pedicillate, the other a

single sessile spikelet.

Etymology : The specific epithet is from the

Greek cylindro- (cylindrical) and -sperma

(seed) in reference to the shape of the

cleistogamous caryopses.

Conservation status: Calyptochloa

cylindrosperma is common at several

locations in northern central Queensland but

has a very restricted range with narrow habitat

diversity. The small populations suggest this

species should be considered as Critically

Endangered (criterion Bla,b) (IUCN 2001).

3. Calyptochloa johnsoniana E.J.Thomps.

& B.K.Simon, species nova similar to C.

gracillima differing by the longer axillary

spikelets (6-6.1 mm versus 3.5-5.5 mm) with

a longer upper glume (>4.8 mm versus <4.5

mm); the fertile leaf sheaths (woody abaxially

and chartaceous adaxially versus semi-woody

to woody for the whole circumference); the

fertile culm internodes (bowed adjacent

to the spikelets and exserted from the leaf

sheath versus culm retained within the leaf

sheath); upper glume of the terminal spikelet

(scabrid versus pilose). Typus: Queensland.

Leichhardt District: Duaringa, December

1976, R. W. Strickland s.n. (holo: BRI

[AQ670557]).

Decumbent stoloniferous perennial.

Ascending branches to 90 cm tall, copiously

branched with 10-20 nodes. Stolons to c. 0.5

m long. Mid-culm leaf blades 20-40 mm long,

3-4 mm wide; adaxial surface with scattered

646

Austrobaileya 8(4): 634-652 (2012)

Fig. 5. Axillary spikelet of Calyptochloa cylindrosperma. A. habit x0.7. B. leaf sheath enclosing axillary spikelet x6.

C. spikelet with upper glume facing x8. D. upper glume xl6. E. lower lemma xl6. F. x-sectional view of lower lemma

xl6. G. upper lemma xl6. H. upper palea xl6.1. grain face view xl6. J. grain side view xl6. K. grain cross-sectional

view xl6. A-K from Thompson CHA767 et al. (BRI). Del. W.Smith.

Thompson & Simon, Calyptochloa

647

Fig. 6. Terminal spikelet of Calyptochloa cylindrosperma. A. side view x8. B. lower glume facing x8. C. upper glume

facing x8. D. lower glume xl6. E. upper glume xl2. F. lower lemma xl2. G. upper lemma xl2. H. upper palea xl2.1.

gynoecium and stamens x!6. A-I from Thompson CHA767 etal. (BRI). Del. W. Smith.

648

Austrobaileya 8(4): 634-652 (2012)

Table 1. Comparison of morphological characters for Calyptochloa taxa*

Character state

C. cylindrosperma

C. gracillima

subsp.

gracillima

C. gracillima

subsp.

ipsviciensis

C. johnsoniana

Terminal

spikelets

Spikelet length x

width (mm)

5-6 x 1.5-2

3-5 x 1.3-1.8

3-4.6 x 1.1-1.6

4.5- 5.1 x

1.5- 1.7

Lower glume length

(mm), apex shape

0.1-0.5, acute

<0.2, obtuse-

truncate

0.8-1.8, acute

not observed

Upper glume length

(mm), apex shape

5-6, acute

3-5, obtuse to

truncate

2.3-4.6, acute

4.5-5.1, truncate

Lower lemma length

(mm)

5-6

3-4.8

2.3-4.6

4.5-5.1

Upper lemma length

(mm)

3-4

3-3.5

2.2-3.2

3.1-3.5

Upper lemma awn

length (mm)

2.5-4

2-3

0.5-2.4

1-2

Upper palea length

(mm)

3-4

2.5-3

2-2.7

3-3.3

Caryopsis length

(mm)

not observed

2.3 (1.6-1.8*)

not observed

Anther length (mm)

2.5-2.6

1.6-2 (0.5-1*)

c.1.5

c.1.6

Axillary

spikelets

Spikelet length x

width (mm)

6-7.5 x 0.7-0.9

4-5.5 x 1-1.1

3.5-4.2 x

0.8-0.9

6-6.1

Upper glume length

(mm)

0.7-4.5

0.5-1.5

0.7-3.5

4.8-5.1

Upper lemma &

lower lemma length

(mm)

6-7.5

4-5.5

3.5-4.2

4-5

Upper lemma awn

length (mm)

1.5-3

2-2.6

0.5-2.5

c.4.5

Upper palea length

(mm)

5.5-6

3-3.8

2.7-3.5

4-4.5

Caryopsis length x

widthe (mm) shape

3.8-4 x 0.5-0.6,

cylindrical

2-3.5 x 0.7-0.8,

c. plano-convex

2.3-3.7 x 0.5-

0.8, c. plano¬

convex

c.4xl,

cylindrical

Anther length (mm)

c. 0.6

0.3-0.4

0.4-0.7

c. 0.3

Width of mature

fertile leaf sheath

(mm)

1.2-1.4

1.5-2.7, mostly

c. 2

1.2-2.5, mostly

c. 1.5

Thickness of mature

fertile leaf sheath at

abaxial wall (mm)

0.2-0.3

0.3-0.5

0.3-0.4

c. 0.3

data from Hubbard (1933b)

Thompson & Simon, Calyptochloa

simple and tubercle-based hairs 0.5-2 mm

long; abaxial surface with scattered simple

and tubercle-based hairs 0.5-2 mm long.

Mature fertile leaf sheaths disarticulating,

10-20 mm long, c. 1.5 mm wide near base

with wall to 0.3 mm thick on abaxial side,

tapering to chartaceous margins; tubercle-

based trichomes c. 0.5 mm long between

nerves, simple hairs absent; outer margin

hairs to 1 mm long. Fertile culm internodes,

13-20 mm long, protruding from leaf sheath

and bowing around caryopsis. Sterile leaf

sheaths with scattered tubercle-based hairs

c. 0.3 mm long. Sterile culm internodes with

medium density simple hairs to 2 mm long.

Terminal inflorescences on axes 2-5 cm

long, 5-6-flowered. Spikelets 4.5-5.5 mm

long (without awn), 1.5-1.7 mm wide; lateral

pedicels 0.3-1 mm long; ultimate pedicels 4-5

mm long. Lower glume not observed. Upper

glume 4.5-5.1 mm long, body with simple

hairs to 1.5 mm long at base and scabrid

for 60% of length; margins with simple

hairs to 2 mm long for 75% of length; apex

truncate. Lower lemma 4.5-5.1 mm long,

dense tubercle-based hairs to 2.5 mm long

for 75% of length; apex obtuse. Upper lemma

body 3.1-3.5 mm long, awn to 1-2 mm long.

Lodicules c. 0.2 mm long. Upper palea 3-3.3

mm long. Anthers c. 1.6 mm long. Caryopsis

not seen. Axillary inflorescences, spikelets

single or rarely paired, one sessile and the

other pedicellate, pedicel c. 6.5 mm long,

enclosed within leaf sheaths with scarsely

enlarged basal portion, usually present at 3-5

internodes. Spikelets 6-6.1 mm long (without

awn), 1.4-1.5 mm wide. Upper glume 4.8-5.1

mm long, apex obtuse to truncate. Lower

lemma 6-6.1 mm long. Upper floret c. 80% of

length of lower. Upper lemma body 4-5 mm

long, apex with two lateral lobes 0.3-0.5 mm

long and awn 3-4.5 mm long. Palea 4-4.5

mm long. Anthers c. 0.3 mm long. Caryopsis

cylindrical, c. 4 mm long and 1 mm wide. Fig.

7 & 8.

Distribution and habitat: The species is

known only from the type specimen collected

from a red soil plateau near Duaringa (Map

1). The notes on the specimen label do not

provide details about the habitat; however,

from our existing knowledge it is very likely

649

to be woodland dominated by Acacia shirleyi

(RE 11.7.2).

Phenology: Flowers in December and

probably through to March during the wet

season.

Notes: Calyptochloa johnsoniana has an

overlapping distribution and habitat with C.

gracillima subsp. gracillima (Table 1) and

C. sp. (Duaringa K.D. Addison 42). Because

of the similar growth habit and leaves, C.

johnsoniana could easily be confused in the

field with C. sp. (Duaringa K.D. Addison

42) which differs by characters including the

following: the scabrid ellipsoid cleistogamous

spikelets, with woody upper glume and lower

lemma, in axillary racemes; the terminal

inflorescences being a reduced panicle; the

terminal spikelets with upper glume and

lower lemma having elliptical cross-section

and upper floret equal to the spikelet length.

Conservation status: This species is only

known from a single specimen from the type

locality near Duaringa. Pending the discovery

of additional populations that may extend

the geographical range, we recommend that

this species should be considered Critically

Endangered (criterion Bla-b [IUCN 2001]).

Etymology: The specific epithet is in honour

of Dr Robert W. Johnson (1930-2012), former

Director at the Queensland Herbarium from

1976-1990.

Achnowledgements

We are very grateful to Dr. G.P. Guymer and

A. Holland for their critical review of a draft

of the manuscript, and Will Smith for the

botanical illustrations and map. Many thanks

to Steven Priday who first brought attention

to the location of plants of the new subspecies

at Ipswich, and David Moore for additional

locations.

650

Austrobaileya 8(4): 634-652 (2012)

Fig. 7. Axillary spikelet of Calyptochloa johnsoniana. A. leaf sheath enclosing spikelet x3. B. side view x8. C. upper

glume xl2. D. lower lemma xl2. E. lower lemma xl2. F. upper lemma xl2. G. upper palea xl2. H. immature caryopsis

xl2.1. cross-sectional view of immature caryopsis xl2. A-I from Strickland s.n. (BRI [AQ670557]). Del. W.Smith.

Thompson & Simon, Calyptochloa

651

Fig. 8. Terminal spikelet of Calyptochloa johnsoniana. A. portion of culm with terminal inflorescence *1; B. side

view xl2; C. upper glume xl2; D. lower lemma xl2; E. upper floret x. A-E from Strickland s.n. (BRI [AQ670557]).

Del. W.Smith.

References

Beentje, H. (2010). The Kew Plant Glossary an

illustrated dictionary of plant terms. Kew

Publishing: Kew.

Blake, S.T. (1941). New genera of Australian grasses.

University of Queensland Papers Department

of Biology 1(19): 1-12 + plates.

Campbell, C.S., Quinn, J.A., Cheplick, G.P., Bell, T.J.

(1983). Cleistogamy in grasses. Annual Review

of Ecology and Systematics 14: 411-441.

Chase, A. (1908). Axillary cleistogenes in some

American grasses. American Journal of Botany

5: 254-258.

Clayton, W.D. & Renvoize, S.A. (1986). Miscellaneous

notes on Panicoid grasses. Genera Graminum

Grasses of the World. Kew Bulletin Additional

Series 13: 1-389.

Clifford, H.T. & Ludlow, G. (1972). Keys to the Families

and Genera of Queensland Flowering Plants

(Magnoliophyta). University of Queensland

Press: St Lucia.

Connor, H.E. (1979). Breeding systems in the grasses:

a survey. New Zealand Journal of Botany 17:

547-574.

- (1981). Evolution of reproductive systems in

the Gramineae. Annals of Missouri Botanical

Garden 68: 48-74.

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652

Derm (2011). Regional Ecosystem Description Database

(REDD). Version 6.0b Updated January 2011,

(Department of Environment and Resource

Management: Brisbane), http: //w w w. derm.

qld.gov.au/wildlifeecosvstems/biodiversitv/

regionalecosvstems/

Hubbard, C.E. (1933a). Cleistochloa subjuncea

C.E.Hubbard. Hooker’s leones Plantarum 33

(3209): 1-6.

- (1933b). Calyptochloa gracillima C. E.Hubbard.

Hooker’s leones Plantarum 33 (3210): 1-3.

Iucn (2001). IUCN Red list Categories and Criteria.

Version 3.1. IUCN Species Survival

Commission: Gland/Cambridge.

Kellogg, E.A. & Campbell, C.S. (1987). Phylogenetic

analysis of the Gramineae. In T.R.Soderstrom

et al. (eds.). Grass Systematics and Evolution ,

pp. 310-322. Smithsonian Institution Press:

Washington.

Simon, B.K., Sharp, D. & Thompson, E.J. (2010).

Poaceae. In P.D.Bostock & A.E.Holland (eds ),

Census of the Queensland Flora 2010 , p. 142—

160. Department of Environment and Resource

Management: Brisbane.

Tothill, J.C. & Hacker, J.B. (1983). The Grasses

of Southern Queensland. University of

Queensland Press: St Lucia.

Map 1. Distribution of Calyptochloa cylindrosperma

(★), C. gracillima subsp. gracillima (•), C. gracillima

subsp. ipsviciensis (A) and C. johnsoniana (V)

Webster, R.D. (1987). The Australian Paniceae

(Poaceae). J.Cramer: Berlin.

The Australian species of Blainvillea ass. (Asteraceae: Ecliptinae)

A.E. Orchard

Summary

Orchard, A.E. (2012). The Australian species of Blainvillea Cass. (Asteraceae: Ecliptinae).

Austrobaileya 8(4): 653-669. Characters separating the Australian species of Blainvillea from the

closely related Wedelia are discussed. Four species are recognised in Blainvillea ; these are keyed,

described and illustrated ( Blainvillea acmella (L.) Philipson, B. calcicola Orchard, sp. nov., B.

cunninghamii (DC.) Orchard, comb. nov. and B. gayana Cass.). Of these, the first three are considered

native, while B. gayana is a naturalised alien. Examination of extra-Australian specimens reveals

that these four species account for the genus worldwide and a partial global synonymy is presented.

Key Words: Asteraceae, Ecliptinae, Blainvillea, Blainvillea acmella, Blainvillea calcicola, Blainvillea

cunninghamii, Blainvillea gayana, Australia flora, taxonomy, new species, identification key

A.E.Orchard, c/o Australian Biological Resources Study, GPO Box 787, Canberra ACT 2001,

Australia. Email: tony.orchard@environment.gov.au

Introduction

The pantropical genus Blainvillea Cass,

comprises four species, although at various

times as many as ten have been recognised (e.g.

Bentham & Hooker 1873). It was described

by Cassini (1823), with a single species, B.

rhomboidea Cass. Other species were later

described in Blainvillea , or transferred from

genera such as Verbesina L., Spilanthes Jacq.,

Eclipta L. and Wedelia Jacq. For example,

de Candolle (1836a) recognised five species,

B. rhomboidea , B. latifolia (L.f.) DC. (based

on Eclipta latifolia L.f.), B. gayana Cass.,

B. prieuriana DC. and B. biaristata DC. He

also accepted (de Candolle 1836b) 34 Wedelia

species, one of which (W cunninghamii DC.)

is here transferred to Blainvillea. Bentham

(1867) recognised no Blainvillea species for

Australia, although his “ Wedelia urticifolia ”

was a misidentification of what is here

accepted as B. cunninghamii.

The type species of Blainvillea was

established when Koster & Philipson (1950)

identified Cassini’s B. rhomboidea as the

species described by Linnaeus as Verbesina

acmella , and by de Candolle as Blainvillea

latifolia , and treated them all under the

new combination Blainvillea acmella (L.)

Philipson.

Accepted for publication 22 August 2012

Specht (1958) described a new species of

Blainvillea (B. dubia Specht) from Arnhem

Land, the only one to that period with an

Australian type, noting the difficulty in

assigning it to Blainvillea versus Wedelia.

More recently, the species B. gayana Cass.,

native to Africa, has been noted as locally

naturalised near Mackay and Proserpine,

Queensland. Other Queensland material has

been tentatively identified as “ Moonia sp. QP\

and “ Wedelia sp. (Marrett River J.Elsol 680 &

T. Stanley)”. In the Northern Territory, some

specimens have been assigned the temporary

informal name Wedelia sp. Limestone

(J.Russell-Smith 7865) N.T. Herbarium.

Generic circumscription of Blainvillea

Even a cursory examination of the synonymy

of currently recognised taxa will reveal that

taxonomists over the last nearly 200 years

have struggled with generic delimitation in

the Wedelia group of the subtribe Ecliptinae

Less., with taxa moving between Wedelia ,

Blainvillea , Eclipta , Wollastonia DC. ex

Decne., Pentalepis F.Muell., Moonia Arn.

and Pascalia Ortega. Pentalepis is one of

the more complex examples: Mueller (1863)

described the genus Pentalepis with two

species. Bentham (1867) treated them as

Moonia in Flora Australiensis. They were

subsequently transferred to Chrysogonum L.

by Mueller (1882). Stuessy then reduced both

654

Moonia (Stuessy 1975) and Chrysogonum

(Stuessy 1977) to monospecific genera from

India / Ceylon and the Americas respectively,

referring the Australian species to “aff.

Blainvillea”. Lawrence (1992) recognised

three Australian taxa in Chrysogonum , while

Karis etal. (1993) resurrected Pentalepis, with

three species). At various times Wollastonia,

Pentalepis and Pascalia have been included

in Wedelia , but are here and elsewhere (e.g.

Panero 2007) considered distinct.

In preparing a treatment of Blainvillea for

Flora of Australia it thus became necessary to

first define the genus, particularly in respect

to Wedelia. It should be noted that here and

elsewhere in this paper the name Wedelia is

used as currently accepted by many authors

to include W. spilanthoides and about six

related northern Australian taxa. The correct

generic placement of these and related

Malesian taxa will be the subject of a separate

paper (in prep.). Unfortunately, there is no

recent monograph covering either Wedelia

or Blainvillea. Regional treatments such as

Grierson (1980) use characters to separate the

genera that work within their limited taxon

range (in that case, floret colour), but have

limited utility elsewhere. Within Australia,

the dearth of northern Floras means that few

attempts have been made to define the genera.

Bailey (1900) used “Ray-florets with small

ligules. Achenes not winged.” ( =Blainvillea )

versus “Ray-florets with large yellow ligules.

Achenes thick.” ( =Wedelia ) as key characters,

but he recognised only one species of

Blainvillea and five of ‘ Wedelia ” (one of

which is now Wollastonia). This limited

coverage, plus the vague qualitative nature

of his characters, makes them unsuitable

for current purposes, where there are four

species of Blainvillea and about six species of

“ Wedelia ”, plus Wollastonia. Ewart & Davies

(1917) did not recognise Blainvillea , but one of

their “ Wedelia ” species (W urticifolia , treated

here as B. cunninghamii ) was distinguished

from the rest by “Five or six of the outer

involucral bracts more leaf-like and longer

than the others” versus “Outer involucral

bracts not longer than the inner ones” (=

three species of “ Wedelia ” of which one is

now Wollastonia). The only recent attempt

Austrobaileya 8(4): 653-669 (2012)

to key these genera within Australia was that

of Toelken (1983), who used the characters

“Fruits curved; scales on receptacle folded”

(=Blainvillea , one sp.) versus “Fruits straight;

scales on receptacle chaffy” (=” Wedelia ”, six

spp., including Wollastonia). In a world-wide

summary, Panero (2007) used the character

“Cypselae with elaiosomes” ( =Wedelia ) versus

“Cypselae without elaiosomes” (= Blainvillea )

to key the two genera. Elaiosomes are present

in many if not all core American Wedelia

species, but not in Australian “ Wedelia :”.

Examination of the Australian taxa;

however, reveals a more useful set of

characters, based on the paleae (receptacle

bracts). In Blainvillea these are more or less

oblong, truncate, laciniate and sometimes

ciliate at the apex, scarious throughout

(sometimes the central tooth greenish), and

folded strongly around the florets (becoming

flatter but still semi-clasping in fruit). In

“Wedelia ” the paleae are more or less

lanceolate, tapering gradually to an acute

apex, not laciniate, and green and fleshy in

the upper part (scarious below). In Australian

taxa of Blainvillea the number of ray florets

is usually 2-5 (rarely 8) and disc florets 2-8,

with about 4-6 (-16) achenes maturing per

capitulum. In “ Wedelia ” the number of florets

is usually much higher, resulting in broader

more hemispherical to depressed globular

capitula, appearing spiny from the rigidly

erect palea tips.

Adoption of this set of characters means

that Wedelia cunninghamii (mainly in

the Northern Territory, but extending to

Queensland and Western Australia) must

be transferred to Blainvillea , where it is

synonymous with (and the epithet pre-dates) B.

dubia. The name Wedelia urticifolia has been

misapplied to these plants (and sometimes

to B. acmella) since Bentham (1867). Much

material in herbaria of B. cunninghamii has in

the past also been misidentified as B. acmella

(syn. B. latifolia). Blainvillea acmella is

present in Australia, but only in Torres Strait

and a few near-coastal localities on Cape York

Peninsula. Wedelia urticifolia does not occur

in Australia. Blainvillea gayana is naturalised

in a few localities in Queensland. Finally, a

Orchard, Australian Blainvillea

655

new species of Blainvillea ( B. calcicola) is

described from two small areas in the Northern

Territory, making a total of four species (one

naturalised, three native) of Blainvillea from

Australia. These are described and illustrated

below, and as they comprise the whole

genus world-wide, a partial extra-Australian

synonymy is provided.

The genus Eisenmannia Sch.Bip. (a generic

synonym listed below) with a single species,

E. clandestina Sch.Bip. was foreshadowed by

Hochstetter (1841), but as a name only. The

following year the species was relisted as a

synonym of Blainvillea gayana (Schnizlein

1842: 135). A duplicate of the collection on

which this invalid name was based is held

in K (in agris Sorghi graminosis ad montem

Cordofanum Arasch-Cool, 16 October 1839,

Kotschyi iter Nubicum, K410220!, ex herb.

Bentham). It is B. acmella.

Materials and methods

This paper arises from work undertaken to

prepare a treatment of subtribe Ecliptinae (and

other groups of the “Heliantheae alliance”)

for the Flora of Australia. It is based upon

examination of all available material in the

major Australian herbaria (AD, BRI, CANB,

DNA, HO, MEL, NE, NSW, PERTH),

plus some in BM, G, G-DC, K, L, and P.

All specimens cited have been seen unless

indicated n.v.

Common abbreviations used in the text

include: N.P (National Park), NT (Northern

Territory), Qld (Queensland) and WA

(Western Australia).

Taxonomy

Blainvillea Cass., Diet. Sci. Nat. 29:

493 (1823), after Henri M. Ducrotay de

Blainville (1777-1850), professor of zoology,

comparative anatomy and physiology, Paris.

Type: Blainvillea rhomboidea Cass. [=B.

acmella (L.) Philipson]

Eisenmannia Hochst., Flora 24 (1,

Intelligenzblf. 42 (1841), nom. inval., nom.

nud.

Annual or perennial herbs with taproot; stems

erect. Leaves opposite (sometimes becoming

alternate near the inflorescence), simple,

3-veined, petiolate; base cuneate, apex acute;

margins crenate to serrate. Capitula in leafy

dichotomous cymes or openly paniculate,

radiate or almost disciform; involucral

bracts in 2 series; outer bracts usually green;

inner bracts sometimes scarious; receptacle

paleaceous; paleae scarious, oblong, truncate

and laciniate (sometimes also ciliate) at apex,

conduplicate, enclosing florets; ray florets

female and fertile, with ligule minutely 2-(or

3-) lobed, yellow or white (mauve-white in

B. gayana ); disc florets bisexual and fertile,

5-merous, with corolla yellow or white;

pappus of scales (often minute) and/or 1-

several awns. Ray achenes 2 or 3 angled,

usually dorsiventrally compressed, smooth

to weakly rugose; disc achenes obpyramidal

to obovoid, 2-4-angled, usually laterally

compressed. Pappus of 0-5 awns, and/or with

a few (usually very short), unequal, shortly

connate scales.

A pantropical genus of four species. In

Australia three native (one endemic) and one

introduced. All occur in northern Australia.

The achenes of most species are very

minutely wrinkled transversely under 10*

magnification. These fine transverse wrinkles

underlie any other rugosities.

656

Austrobaileya 8(4): 653-669 (2012)

Key to the species of Blainvillea

1 Achenes all ±subcylindrical (angled but not noticeably compressed), c. 5x

as long as wide, awns 2-8, stiff, 1.5-3.5 mm long.2

1. Achenes compressed, cuneate to obovoid, 2-2.5x as long as wide; awns

0-2, weak, 0.5-2.0 mm long.3

2 Awns 2 (or 3), terete.1. B. gayana

2. Awns 5-8, flattened (i.e. anatomically, lengthened slender scales).2. B. calcicola

3 Achenes convex at apex, 3-4 (-4.5) mm long, smooth, awns c. 0.5 mm

long (if present).3. B. cuiminghamii

3. Achenes truncate or sunken at apex, usually with raised corners at

summit of angles, 4-5 mm long, weakly rugose; awns 1-1.3 mm long

(if present).4. B. acmella

1. Blainvillea gayana Cass., Diet. Sci. Nat.,

edn. 2,47: 90 (1827). Type citation: “Senegal,

M.Gay Type: Jardin de Luxembourg, la

4 Septembre 1826. Secher par Hardy en

mon absence, des grains m’ont ete envoyees

de Richardtol (Senegal) sous le nom d’

Ageratum? Bidens? Semer en Avril 1825

(holo: K 410221); iso: Blainvillea, cultive au

Luxembourg en 1826, les graines venant du

Senegal, P 69616 (photo!); (see typification

note below).

Blainvillea prieureana DC., Prodr. 5: 492

(1836). Type citation: “In Senegalia superiore

ad montes Bakel legit cl. Leprieur (v.s. comm,

a cl. inv.)” Type: Senegambie, s.dat., Leprieur

s.n. (holo: G 23503, photo!).

Oligogyne burchellii Hook., Icon. PI. 1: t.

101 (1837), non Aspilia burchellii Baker,

nec Wedelia burchellii (Baker) B.Turner;

Calyptocarpus burchellii (Hook.) Sch.Bip.,

Bot. Zeitung (Berlin) 24: 165 (1866). Type

citation: “Rio Janeiro. Wm. J. Burchell, Esq.

(n. 12)”. Type: K, n.v.

Illustrations : Hooker (1837: t. 101),

as Oligogyne burchellii. Pacific Island

Ecosystems at Risk (2011).

Erect annual herb to 2 m tall; stems softly

pilose. Leaves opposite below, alternate above,

broadly ovate to deltoid, becoming lanceolate

in inflorescence, 30-120 mm long, 15-85

mm wide, shallowly crenate-serrate, pilose

adaxially, more densely pilose abaxially,

especially on veins; petioles 6-30 mm long.

Capitula elongated-hemispherical, 10-12

mm diameter; involucral bracts lanceolate,

subacute, green, pilose near apex, glabrous

and striate below. Paleae oblong, stramineous,

scarious, with laciniate apex, striate, glabrous

except midrib. Ray florets 4-8; corolla mauve

to white, c. 3 mm long; ligule 2-lobed. Disc

florets 6-8; corolla white, c. 3.5 mm long.

Achenes 6-16, narrowly cylindrical, brown

to black, very finely transversely wrinkled,

finely and sparsely pilose or subglabrous. Ray

achenes subcylindrical, triquetrous, 4.5-5

mm long, slightly curved; pappus of 3 erect

awns. Disc achenes subcylindrical, 2 or 3

angled, 5.5-6.5 mm long, ± straight; pappus

of 2 (or 3) erect awns. Awns 1.5-2.5 mm long,

antrorsely pilose, on a rostrum c. 0.5 mm

long. Fig. 1.

Additional selected specimens examined : Africa

(selection only): Cape Verde Islands: s. loc., 1895,

Cardosas.n. (K). Ethiopia: Abaye (BlueNile) Gorge, Sep

1973, Gilbert & Getachew 3089 (K). Somalia: 14 km on

road between Luuq and Beled Xaawo, Jun 1989, Thulin &

Mohamed 6960 (K). Djibouti: between Djibouti and Arta,

Apr 1993, Collenette 8699 (K); Cameroun: Colline de

Boboyo, Sep 1964, Bounouge 97 (K). Nigeria: Panshanu

Pass, Aug 1962, Lawler & Hall 188 (K). Kenya: Marsabit

N.P, Feb 2005, Muasya NMK487 et al. (K); Tanzania:

Mkomazi Game Reserve, Apr 1995, Abdallah & Vollesen

95/12 (K). Senegal: 20-23 km from Dendoudi, Oct 1988,

Lawesson 5307 (K). The Gambia: Yundum, Nov 1979,

Terry 3231 (K). Ivory Coast: Pronoi, Nov 1971, Audrun

798 (K). Botswana: Ngamiland, Sennonore, Mann, Apr

1994, Smith 5696 (CANB). South America: Bolivia:

57 km al S de Las Petas, May 2008, Wood 24869 et al.

(K). Brazil: Bresil Meridional, 1838, Guillemin 177 (K).

Australia: Queensland. South Kennedy District: 35

km N of Mackay, Kuttabul, Brangus Court, Apr 2002,

Warren s.n. (BRI); 6.7 km along Dingo Beach Road, N

of Proserpine, Apr 2000, Bean 16327 (BRI, K, NSW);

Royston Park, property of Williamsons, W of Bruce

Highway, c. 6 km N of township of Kuttabul, Apr 2002,

Warren 2 (BRI).

Orchard, Australian Blainvillea

657

Fig. 1. Blainvillea gayana : A. lower leaf x0.6. B. upper leaf x0.6. C. young fruiting capitulum x6. D, E. involucral

bracts x6. F, G. paleae x6. H. ray floret xl2.1. disc floret xl2. J. ray achene x6. K. diagramatic transverse section of the

ray achene shown as J. L. detail of summit of ray achene xl2. M. disc achene x6. N. diagramatic transverse section of

the disc achene shown as M. O. detail of summit of disc achene xl2. A-C, & J-0 from Bean 16327 (BRI), D-I from

Warren s.n. (BRI [AQ555637]). Del. A.E.Orchard.

658

Distribution and habitat : The species is

a native of northern and tropical eastern,

western and southern Africa (as far south as

Botswana) where it is common in open areas.

It is also known from the northern part of

South America (e.g. Bolivia, Brazil) where

it is possibly an introduction via the slave

trade. In Australia (Map 1), several small

infestations have been recorded in cleared land

(remnant Corymbia clarksoniana (D.J.Carr

& S.G.M.Carr) K.D.Hill, C. tessellaris

(F.Muell.) K.D.Hill & L.A.S.Johnson and E.

platyphylla F.Muell. woodland, on alluvial

flats, stony gravels on hills, grassy cleared

pastures) and adjacent to roadsides near

Mackay and Proserpine, in North and South

Kennedy districts of northern Queensland.

While limited in distribution it can be locally

abundant.

Phenology : Flowers and fruits recorded in

April.

Typification : There are two specimens, one

in Paris (P), the other in Kew (K), which

seem to be type material for Blainvillea

gayana. The one in Kew has attached to it

a copy of the original printed description,

with an annotation “Diet, des Sc. nat. tom.

XLVII, publie le 23 e Mai 1827” (apparently

not in Cassini’s hand), an annotation in

French of subsequent discoveries of the plant

(apparently in Cassini’s hand), and a copy of

a letter, almost certainly to Gay, signed by

Cassini, dated “Paris, ce 24 novembre 1826”,

in which he says “J’ai soigneusement examine

votre pi ante du Senegal: ille apparient

bien a mon genre Blainvillea : mais c’est

indubitablement une nouvelle espece, que ja

vous demande la permission de nominer BL

gayana, ...” It seems likely from this that the

Kew specimen is the original material, that in

Paris being a secondary duplicate (although

labelled “Holotype”), and the K specimen is

thus regarded here as the holotype, and that

in P an isotype.

Although I have seen no authentic material

of Oligogyne burchellii , from Hooker’s plate

there is no doubt that his plant is conspecific

with B. gayana. This has been confirmed by

N.Hind (K ,pers. comm).

Austrobaileya 8(4): 653-669 (2012)

Notes : Easily distinguished from other

Australian species of Blainvillea by its

subcylindrical achenes with 2 or 3 long stiff

awns. It is most likely to be confused with

B. acmella, but differs in its capitula being

narrower, particularly in flower (longer than

broad), while those of B. acmella are usually

broader than long. The awns on the achene

of B. gayana are always straight and usually

2 mm or more long. Those of B. acmella are

usually less than 2 mm and often curved

and weaker. Its relationships, however, are

probably with B. calcicola , with which it

shares its long slender achenes.

2. Blainvillea calcicola Orchard species

nova; resembling B. cimninghamii, but

differing in having achenes which are

cylindrical or barely compressed, c. 5.5 mm

long, with 5 to 8 of the pappus scales greatly

elongated (2-3.5 mm long) and serving as

pseudo-awns. Typus: Northern Territory.

Mathison Creek, Willeroo, 15 March 1989,

J.Russell-Smith 7865 & D.Lucas (holo: DNA

42377; iso: BRI [AQ481744]).

Wedelia sp. Limestone (J. Russell-Smith

7865) N.T.Herbarium; Short etal. (2011: 19).

Erect annual herb (15-) 80-100 cm tall; stems

softly pilose. Leaves all opposite, ovate,

45-70 mm long, 20-40 mm wide, shallowly

crenate, moderate to densely, softly pilose

adaxially, more densely pilose abaxially,

especially on veins, with sessile golden

glands between veins; petioles 15-20 mm

long. Capitula obconical, 5-6 mm diameter;

involucral bracts lanceolate, acute, green,

densely pilose throughout. Paleae oblong,

stramineous, scarious, with laciniate and

ciliate apex, striate, with short appressed hairs

and sessile golden glands dorsally. Ray florets

2 or 3; corolla yellow, 3-4 mm long; ligule 2

or 3-lobed. Disc florets 2 or 3; corolla yellow,

4- 5 mm long. Achenes 4-6, grey, very finely

transversely wrinkled, shortly pilose at apex

and on angles. Ray achenes ±cylindrical

(not noticeably compressed), c. 5.5 mm long,

±straight, minutely tuberculate; pappus of

5- 8 erect to subpatent, flattened pilose scales

2-3.5 mm long. Disc achenes similar, but

smooth, usually lacking tubercles. Fig. 2.

Orchard, Australian Blainvillea

659

Fig. 2. Blainvillea calcicola : A. leaf x0.6. B. leaf margin, abaxial view, showing hairs and glands (simplified) xl.5.

C. fruiting capitulum x6. D, E. involucral bracts x6. F. palea, dorsal view x6. G. palea, lateral view x6. H. disc floret

x6.1. corolla of ray floret x6. J. corolla of young disc floret x6. K. ray achene x6. L. apical detail of ray achene xl2. M.

disc achene x6. All from Russell-Smith 7865 & Lucas (A-G, I, J from BRI; H, K-M from DNA). Del. A.E.Orchard.

660

Additional specimens examined : Northern Territory.

Mathison Creek, Willeroo, Mar 1989, Dunlop 8318 &

Leach (DNA); Timber Creek, Mar 1998, Michel 1246

(DNA).

Distribution and habitat : Blainvillea

calcicola is endemic to the Northern Territory,

where it is found in vine thickets on karst,

with Celtis philippinensis Blanco and Trema

tomentosa (Roxb.) Hara, in a limited area near

Willeroo and Timber Creek (Map 2).

Phenology : Flowers and fruits present in

March.

Notes : In Blainvillea calcicola the scales

at the apex of the achene are larger and

perform the function of awns. It appears to

be restricted to karst formations, but note that

the superficially similar B. cunninghamii can

also be found on this substrate. Blainvillea

cunninghamii is distinguished, inter alia , by

flattened achenes with pappus scales less than

0.5 mm long. The bodies of the achenes of B.

calcicola are similar to those of B. gayana ,

but the latter has a pappus of 2 or 3 long stiff

awns, not c. 7 long flattened scales.

3. Blainvillea cunninghamii (DC.) Orchard

comb, nov.; Wedelia cunninghamii DC.,

Prodr. 5: 540 (1836). Type citation: “In

rupestribus insularum Goulburn ad oram

borealum Australiae, mart. flor. legit cl.

A.Cunningham.” Type: Australia: Northern

Territory. Verbesina sp., grassy rocky spots,

Goulburn Island, North Coast, March 1818,

[A.Cunningham] 59 (holo: G); Goulburn

Island (1st Voyage of Mermaid), 1818,

A.Cunningham 184 (iso: BM 820301);

Verbesina sp., grassy rocky spots, Goulburn

Island, 28 March 1818, A. Cunningham s.n.

(iso: K); (see typification note below).

Blainvillea dubia Specht, Rec. Amer.-Austral.

Sci. Exped. Arnhem Land, 3, Bot. PI. Ecol.

3: 314 (1958), syn. nov. Type citation: [N.T.]

“South Bay, Bickerton Island {Eucalyptus

alba-E. polycarpa woodland): 524. N.

eg” Type: Australia: Northern Territory.

South Bay, Bickerton Island, in the Gulf of

C arpentar ia (13° 45’ S, 136° 6’E), 10 June 1948,

R.L.Specht 524 (holo: BRI; iso: AD 96149100,

CANB 63749, K, L 1861, MEL 59399, NSW).

Austrobaileya 8(4): 653-669 (2012)

[Wedelia urticifolia auct. non DC.: Bentham

(1867: 538); Bailey (1900: 861); Ewart &

Davies (1917: 280)]

Illustration : Specht (1958: 315, fig. 26), as

Blainvillea dubia.

Erect aromatic annual herb (0.4-) 1-1.2 (-2)

m tall; stems sparsely, softly pilose. Leaves

all opposite, ovate (becoming narrower

in inflorescence) 80-100 (-135) mm long,

35-50 (-90) mm wide, coarsely crenate,

sparsely, softly pilose adaxially, sparsely

pilose on veins abaxially with sparse sessile

golden glands between veins; petioles 15-20

mm long. Capitula hemispherical, 3-5 mm

diameter; involucral bracts lanceolate, acute,

green, densely pilose throughout or in upper

half, with inner bracts broader. Paleae oblong,

stramineous, scarious, striate, with truncate

laciniate and ciliate apex, glabrous dorsally or

with sparse sessile golden glands. Ray florets

2 or 3; corolla yellow, c. 3.5 mm long; ligule

2-lobed. Disc florets 4-6; corolla yellow, c. 2.5

mm long. Achenes 5 or 6, grey to black, very

finely transversely wrinkled, shortly pilose

apically. Ray achenes obovoid-trigonous,

compressed, 3-4 (-4.5) mm long, with apex

convex, without raised corners at summit of

angles, smooth (lacking tubercles), ±straight;

pappus of minute (0.1-0.2 mm long) scales,

sometimes with 1 or 2 extended as short soft

“awns” c. 0.5 mm long. Disc achenes similar

but 2-angled. Fig. 3.

Additional selected specimens examined : Western

Australia. Kalumburu Mission, May 1998, Mitchell

5472 (DNA); 2.5 km N of Face Point, Carson Escarpment,

Mar 1989, Keighery 10666 (CANB, PERTH); Steep

Head Island, Admiralty Gulf, Apr 2006, Mitchell 8570

(CANB). Northern Territory. Stuart Highway, c. 11

miles [c. 18 km] SE of Katherine, Apr 1964, Adams

932 (BRI, CANB, K, L, NSW, NT); headwaters of the

Liverpool River, Apr 1984, Craven & Wightman 8353

(CANB, DNA, MEL); Rocky Bay, Yirrkala, Mar 1988,

Russell-Smith 5170 & Lucas (BRI, DNA); Waterfall

Creek, Apr 1984, Wightman 1281 & Dunlop (BRI,

CANB, DNA). Queensland. Cook District: Lakefield

N.P, 1.6 km S of mouth of North Kennedy River, Apr

1992, Neldner 3775 & Clarkson (DNA); Stanley Island,

May 1995, Le Cussan 539 (BRI).

Orchard, Australian Blainvillea

661

Fig. 3. Blainvillea cunninghamii : A. midstem leaf x0.6. B. upper leaf *0.6. C. capitulum x6. D, E. outer involucral

bracts x6. F. inner involucral bract x6. G. palea, dorsal view *6. H. palea, lateral view *6.1. ray floret *12. J. disc floret

xl2. K. adaxial view of ray achene x6. L. diagramatic transverse section of K. M. abaxial (dorsal) view of ray achene

x6. N. apical detail of ray achene xl2. O. disc achene x6. P. diagramatic transverse section of disc achene. Q. apical

detail of disc achene xl2. A-H, K-Q, from Wightman 1281 & Dunlop (BRI); figures I, J from Craven & Wightman

8353 (MEL). Del. A.E.Orchard.

662

Distribution and habitat: Blainvillea

cunninghamii is native to northern Australia,

from near Kalumburu Mission in WA, across

the NT north of about Katherine, to mainly

inland Cape York Peninsula in Qld, on sandy

and loamy soils, in grassland, woodland

understorey and the margins of vine thickets

and rainforest, from sea level to less than 200

m in NT, but usually at 350-550 m in Qld

(Map 3)

Phenology: Flowers present (Aug.-) Jan-

May (-June), fruits (Jan.-) Mar-June (-July).

Typification: The holotype of Wedelia

cunninghamii in G is numbered 59, while

an apparent duplicate in BM is numbered

184, and another in K is unnumbered. The

numbers applied to his specimens by Allan

Cunningham are consignment list numbers,

rather than unique collection numbers as

currently understood. The number “184”

was the consignment list number when the

specimen was sent from Cunningham to

Banks and Aiton from Sydney in 1818, while

“59” was the consignment list number when

he sent replicate material to Candolle from

London in 1834. It seems that when he sent

duplicates of his Australian collections to de

Candolle in 1834, Cunningham renumbered

them in a new list. Unfortunately this

consignment list has not been located.

However, Cunningham only collected this

taxon once from Goulburn Island, so despite

the difference in numbers, the G, K and BM

specimens should be treated as replicates. A

detailed account of Cunningham’s numbering

system and the disposal of his collections

will be published elsewhere (Orchard, several

papers in prep.).

Notes: This is the plant collected as

“Buphthalmum acuminatum” by R.Brown at

Morgan’s Island, Blue Mud Bay, on 20 Jan.

1803 (BM, NSW), and later by A.Cunningham

at South Goulburn Island in March 1818 (BM,

G, K). Its presence in Australia thus almost

certainly predates European settlement, and

it is here considered native. Bentham (1867)

treated the Brown and Cunningham specimens

as Wedelia urticifolia DC., an Asian/Malesian

species, and in this he was followed by Bailey

(1900), Ewart & Davies (1917) and some later

Austrobaileya 8(4): 653-669 (2012)

authors. Wedelia urticifolia has paleae which

are rigid, lanceolate, with an acuminate, very

acute or acicular apex (Backer & Bakhuizen

van den Brink 1965), not truncate and laciniate

as in Blainvillea. True Wedelia urticifolia has

not been collected in Australia, although it is

present in southern Indonesia.

Blainvillea cunninghamii is most similar

to, and likely most closely related to, the

predominantly Indian / Sri Lankan distributed

B. acmella. Evolution of the former from

the latter by genetic drift, from an isolated

introduction to Australia at a remote time is

not inconceivable. The current presence of

B. acmella in Australia is almost certainly a

recent event.

Plants growing in very damp shaded

positions have larger leaves and capitula, and

achenes to 4.5 mm long.

4. Blainvillea acmella (L.) Philipson, Blumea

6: 350 (1950); Verbesina acmella L., Sp. PI.

2: 901 (1753); Spilanthes acmella (L.) Murray,

Syst. Veg. 731 (1784); Ceratocephalus

acmella (L.) Kuntze, Revis. Gen. PI. 1: 326

(1891). Type: Habitat in Zeylona (lecto: Herb.

Hermann 2: 10, No. 309, BM 594573, photo!),

fide Koster & Philipson (1950: 349).

? Verbesina dichotoma Murray, Comment.

Soc. Regiae Sci. Gott. 2: 15, PI. 4 (1780),

nom. rej. prop:, Blainvillea dichotoma

(Murray) Hemsl., Biol. Cent.-Amer., Bot. 4:

112 (1887). Type citation: “[Hortus Regiae

Goettingen] Floruit medio et exeunte m.

Augusto in vaporario...” Type: n.v ., probably

not preserved. See note below.

Eclipta latifolia L.f., Siipp. PI. 378 (1782);

Blainvillea latifolia (L.f.) DC. in R.Wight,

Contr. Bot. India 17 (1834). Type citation:

’’Habitat in India orientali.” Type: n.v.

Verbesina lavenia Roxb., Hort. Bengal. 62

(1814), & Roxb., FI. Ind. 3: 442 (1832), nom.

illeg., non V. lavenia L., Sp. PI. 2: 902 (1753)

[=Adenostemma lavenia (L.) Kuntze],

^Blainvillea rhomboidea Cass., Diet. Sci.

Nat. 2nd edn, 29: 493 (1823), nom. rej. prop.

Type citation: ’’Cultives au Jardin du Roi,

ou ils fleurissent vers le milieu du mois de

septembre.” Type: Blainvillea rhomboidea

Orchard, Australian Blainvillea

663

H.Cass. [scripsit Cassini] H.P.Coucher

[Hortus Parisiense Grown], 15 October 1817,

Herb. J.Gay (holo: n.v. ?P; iso: K 487624

[Presented by Dr Hooker, February 1868]).

See note below.

Verbesina dichotoma Wall., Numer. List [Cat]

3204A (in part), B, C, D, E (1831), nom. nud.,

& nom. illeg, non V. dichotoma Murray.

Based on [India], 1849, Wallich herb. no. H.9.

3204 H, K, ex herb. Bentham.

Blainvillea alba Edgew., Trans. Linn. Soc. 20:

70 (1846). Type citation: [N.W. India] ’’Pinjar

Dhun, in arvis, Prov. Sirhind, Indiae Bor-

Occ. Sept” Type: India. Pinjar Dhun [sic] in

arvis, 1844, M.P.Edgeworth s.n. (holo: K, ex

herb. Bentham).

Blainvillea hispida Edgew., Trans. Linn. Soc.

20: 70 (1846). Type citation: [N.W.India]

’’Himala, in arvis, alt. ped. 4000-5000,

Junio.” Type: India. Himalaya in arvis, altit.

4-5000 ped., 1844, M.P.Edgeworth s.n. (holo:

K, ex herb. Bentham).

Wedelia sp. (Marrett River J.A.ElsoH 680);

Holland (1997: 34).

Illustrations : Bailey (1906: 86); Koster &

Philipson (1950: 350, fig. 1); Matthew (1982:

PI. 356); Hajra et al. (1993: 378, fig. 101).

Erect annual aromatic herb (0.6-) 1.2-E5 m

tall; stems hispid with sparse hairs swollen

at base. Leaves opposite below, alternate

above, ovate to lanceolate, 80-100 mm long,

(35-) 40-50 mm wide, coarsely crenate,

adaxially sparsely scabrid, abaxially densely

scabrid especially on veins, with coarse

white hairs swollen at base, and sessile

golden glands between veins; petioles 15-

20 mm long. Capitula hemispherical, 6-10

mm diameter; involucral bracts lanceolate

to linear-lanceolate, acute, green, densely

hispid throughout; inner bracts ovate, acute to

acuminate, green above, stramineous below.

Paleae oblong, stramineous, scarious, with

laciniate and sparsely ciliate apex, striate,

shortly pilose dorsally with numerous sessile

golden glands. Ray florets 4-6; corolla

yellow, c. 3 mm long; ligule 2-lobed. Discs

florets 5-7; corolla yellow, c. 2 mm long.

Achenes 6-10, grey to black, shortly pilose

in upper half. Ray achenes cylindrical-

trigonous, somewhat compressed, 4-5 mm

long, often slightly curved, apically truncate

or sunken, the angles crowned by short peaks,

weakly rugose; pappus of very short scales

(sometimes absent), usually with 1 or 2 short

awns 1-1.3 mm long. Disc achenes obovoid,

2-angled, compressed, 5-6 mm long, smooth

apart from fine transverse wrinkles; pappus

as ray achenes. Fig. 4.

Additional selected specimens examined: Africa:

South Sudan: Tokar Delta, Apr 1949, Bally 6978 (K).

Sudan: (Nubia) Mt Cordafanum Arash-Cool, Oct 1839,

Kotschy s.n. (K, ex herb. Bentham). India: Madras,

Hatiguda, Feb 1884, Gamble 13801 (K); Andhara

Pradesh, Khammam district, Oct 1988, Seethapathi

Rao s.n. (K); Bangalore, Nandi Hills road, Jan 1973,

Burtt RHT18356 et al. (K); Salem, Servarayans, Nov

1978, Venugopal & Jayaseelan RHT18781 (K); Jashpur,

Dumarkona, Sep 1941, Mooney 1860 (K). Sri Lanka:

Ceylon, 1848, Gardner 602 (K, ex herb. Bentham);

Polonnaruwa, near Recumbent Buddah statue, Feb 1969,

Grierson 1026 (CANB). Australia: Western Australia,

eastern margin of Mitchell Plateau, Apr 1991, Willing

335 (PERTH). Queensland. Cook District: Mabuiag,

Torres Strait, 13 Apr. 2000, Waterhouse 5847 (BRI,

CANB); Thursday Island, May 1893, Cowley s.n. (BRI);

Albany Island, May 1995, Le Cussan 435 (BRI); Cape

York, s.d. [c. 1866], Daemal s.n. (MEL); Roko Island,

Feb 2002, Waterhouse 6351 (BRI, CANB, US); Marrett

River, Princess Charlotte Bay, May 1979, Esol 680 &

Stanley (BRI); Pickford Road, Biboohra, Mar 2000,

Clayton s.n. (BRI, K).

Distribution and habitat: The species is native

to India and Sri Lanka with occasional records

from Africa (Sudan, possibly elsewhere),

probably representing introductions. It has

been reported from China (Chowdhery

1995) but no specimens from there were seen

during this study. Literature records for the

Americas are mainly misidentifications of B.

gayana although those from the Galapagos

Islands are B. acmella. In Australia (Map

4), it is widespread in the islands of Torres

Strait, and occasional in coastal localities

mainly on northern Cape York Peninsula,

with occasional occurrences further south.

A single collection from Mitchell Plateau,

WA, may represent an introduction. Plants

are usually found in near-coastal localities

near sea level, often on alkaline soils (coral

cays, beach sand, shell banks), sometimes

extending into the understorey of woodland

and vine thickets.

664

Austrobaileya 8(4): 653-669 (2012)

Fig. 4. Blainvillea acmella: A. leaf x0.6. B. fruiting capitulum x6. C, D. involucral bracts x6. E. palea, dorsal view

x6. F. palea lateral view x6. G. ray floret, corolla removed xl2. H. ray floret corolla xl2.1. disc floret, corolla removed

xl2. J. disc floret corolla xl2. K, Q. ray achene, adaxial view x6. L. diagramatic transverse section of K. M. ray achene,

abaxial (dorsal) view x6. N. disc achene x6. O. diagramatic transverse section of N. P. apical detail of ray achene xl2.

R. apical detail of ray achene Q xl2. A-P from Clayton s.n. (BRI [AQ490047]); Q, R from Le Cussan 435 (BRI). Del.

A.E.Orchard.

Orchard, Australian Blainvillea

665

Phenology: Flowers recorded in February to

July, fruits in March to July.

Typification: The name Verbesina dichotoma

Murray was based on material grown in the

Royal Garden at Goettingen and no surviving

authentic specimen has been located.

Murray’s description and plate are somewhat

ambiguous, but on balance, the illustration

of rather broad capitula, and the description

and illustration of the achenes as relatively

broad with rather short awns, suggests that

Murray’s plant was B. acmella rather than B.

gayana. However, because this name would

have priority over B. gayana if proven to be

conspecific, it is being proposed for rejection

against that name in a separate paper.

The combination Blainvillea dichotoma

has been made at least twice. Hemsley (1887)

first made the combination, for Galapagos

Island plants, with the cryptic listing of

“Blainvillea dichotoma, Cass.” This is an

indirect reference to Cassini (1829) in which

the combination is implied but not actually

made, with the observation “Blainvillea. =

An? Verbesina dichotoma, Moench (1794).”

Moench (1794) lists Verbesina dichotoma ,

attributing it to Murray. The correct author

attribution is therefore Blainvillea dichotoma

(Murray) Hemsl. Stewart (1911) listed

Hemsley’s combination as “B[lainvillea]

dichotoma (Murr.) Cass. ace. to Hemsl. Biolog.

Cent.-Am.Bot. IV. 112 (1886-1888)”, citing

Verbesina dichotoma Murray, Comment.

Soc. Regiae Sci. Gott. 2: 15, PI. 4 (1780),

as basionym, and B. rhomboidea Cass, as a

synonym. Later authors (e.g. Wiggins & Porter

1971; McMullen 1999; Jorgensen & Leon-

Yanez 1999) have attributed the combination

as Blainvillea dichotoma (Murray) Stewart,

which is incorrect. From the descriptions and

illustrations in these works, the Galapagos

plants are B. acmella rather than B. gayana.

IPNI also lists “ Blainvillea dichotoma

Luetzelb., Estud. Bot. Nordeste i. (Publ. Insp.

Fed. Obras Contr. Secc., Rio de Janeiro,No.

57, Ser. I.A.) 41 (1922-3), nomen”. It is not

clear whether this nomen nudum was in fact

an attempt at a new combination or merely

usage of Hemsley’s combination.

Blainvillea rhomboidea was described

from plants cultivated in the Royal Botanic

Garden, Paris, from seed of unstated origin.

I have seen no specimens from the Paris

herbarium which could be considered to be

type material, but in Kew there is a sheet with

a printed label “Herb. J.Gay. Presented by Dr.

Hooker, February 1868.” and annotated by

Cassini “Blainvillea rhomboidea, H. Cass.”

and “H[ortusj. P[arisiense], Coucher. 15 e

Octobre 1817.” This is here described as an

isotype, but may in fact be the holotype. It has

capitula with somewhat immature achenes.

The achenes are somewhat intermediate

between what is here accepted as B. acmella

and B. gayana , but the few more mature

ones are rugose and proportionately broad,

and these, plus the broad hemispherical

capitula suggest placement with B. acmella ,

as suggested by Koster & Philipson (1950).

However, because this name would, if found

to be conspecific with B. gayana , threaten that

well-established name, it is being proposed

for rejection against B. gayana in a separate

paper.

The name Verbesina dichotoma Wall.,

invalid because of lack of a description, was

not one of those validated later by G.Don

(see Sprague [1925] for discussion). It is in

any case an illegitimate later homonym of V

dichotoma Murray.

Notes: This species has been confused with

Blainvillea cunninghamii, but differs in its

stiffer, coarser hairs (with distinctly swollen

bases), usually more numerous florets per

capitulum, larger achenes of which at least

the ray achenes are weakly rugose (smooth in

B. cunninghamii ) and the apex of the achenes

is truncate or slightly sunken, with the

angles continued upwards into short peaks.

In Queensland, where both species occur, B.

cunninghamii is usually found inland in damp

forested situations at altitudes to 350-550

m, while B. acmella is mostly a strand plant

of the Torres Strait islands and coastal Cape

York Peninsula.

The embryology of this taxon was

described in detail (under the name B.

rhomboidea Cass.) by Sundara Rajan (1972).

666

Species excludendae et incertae sedis

Blainvillea brasiliensis (Nees & Mart.)

S.F.Blake, Proc. Biol. Soc. Washington

38: 85 (1925); Galophthalmum brasiliense

Nees & Mart., Nov. Act. Acad. Leopold.-

Carol. Nat. Cur. 12: 8, pi. 2 (1824);

Calyptocarpus brasiliensis (Nees & Mart.)

B. Turner, Phytologia 64: 214 (1988). Type

citation: [Brazil], “Ad viam Felisbertiam.”

[Maximilian],

Now generally treated as Calyptocarpus

brasiliensis.

Blainvillea biaristata DC., Prodr. 5: 492

(1836); Calyptocarpus biaristatus (DC.)

H.Rob., Phytologia 41: 34 (1978). Type

citation: “In Brasiliae prov. Rio-Grande (v.s.

in h. Mus. reg. Par. a Mus. imp. Bras, sub n.

873 miss.)” Type: Bresil, Province de Rio

Grande, 1833, C.Gaudichaud 086c, Herbier

Imperial du Bresil No. 873 (holo: P 710003

photo!).

Robinson (1978) advocated transfer of

all of Blainvillea subgenus Oligogyne from

Blainvillea to Calyptocarpus. In the case of B.

biaristatus, this has recently been accepted in

e.g., Zuloaga et al. (2008).

Oligogyne bahiensis DC., Prodr. 5: 629

(1836); Blainvillea bahiensis (DC.) Baker, FI.

Bras. (Mart.) 6: 177 (1884); Calyptocarpus

bahiensis (DC.) Sch.Bip., Bot. Zeitung (Berlin)

24: 165 (1866). Type citation: “Circa Bahiam

Brasiliae legit cl. Blanchet (pi. exs. n. 1706!)”.

Type: n.v. See above under B. biaristata.

Oligogyne megapotamica DC., Prodr. 5:

629 (1836); Calyptocarpus megapotamica

(DC.) Sch.Bip., Bot. Zeitung (Berlin) 24: 165

(1866). Type citation: “In Brasiliae provincia

Rio-Grande (h. Mus. imp. Bras. n. 874)”.

Type: Bresil, Province de Rio Grande, 1833,

C. Gaudichauds.n., Herbier Imperial du Bresil

No. 874, (holo: P 710004, photo!).

Oligogyne tampicana DC. Prodr. 5: 629

(1836); Blainvillea tampicana (DC.) Benth. &

Hook.f., Gen. PI. 2: 370 (1873). Type citation:

“circa Tampico de Tamaulipas Mexicanorum

legit cl. Berlanier (pi. exs. n. 61!).” Type:

Mexico: Tampico de Tamaulipas, [10

February] 1827, Berlandier 61 (holo: G-DC?

Austrobaileya 8(4): 653-669 (2012)

n.v., iso: P 709867, 709868 & 709869 photo!).

A synonym of Calyptocarpus vialis Less.

(McVaugh & Smith 1967).

Oligogyne synedrelloides Hook.f. & Arn., J.

Bot. (Hooker) 3: 316 (1841) [Note that IPNI

also lists the name “ Blainvillea synedrelloides

Benth. & Hook.f., Gen. PI. 2: 370 (1873)”, but

this combination was only implied, not made,

in that publication]. Type citation: “Rio

Grande, Tweedie.”

A synonym of Calyptocarpus biaristatus

(DC.) H.Rob. (Robinson 1978; Zuloaga et al.

2008).

Blainvillea polycephala Gardner, London J.

Bot. 7: 89 (1848). Type citation: [Brazil] “In

dry bushy places near the city of Maranham,

May 1841 [G.Gardner]”

Blainvillea racemosa Gardner, London J.

Bot. 7: 89 (1848); Blainvillea rhomboidea

var. racemosa (Gardner) Baker, FI. Bras.

(Martius) 6: 176 (1884). Type citation: “In

dry, sandy, shady places near Villa de Ico,

Province of Ceara, Aug. 1838”, [G.Gardner].

Type: Brasilia tropica, Prov. Ceara, 1839 [sic],

G.Gardner 1740 (holo: K 54385, photo!).

“Blainvillea amazonica Benth. & Hook.f.,

Gen. PI. 2: 370 (1873)”. This combination

is listed in IPNI, but was only implied, not

validly made, in the place cited. Bentham &

Hooker actually used the name Lipochaeta

amazonica Poepp. & Endl. (Nov. Gen. Sp. PI.

3: 49, t. 256 [1843]). Type citation: “Crescit

in insulis arenosis fluminis Amazonum inter

Ega et Rio negra”.

From the description and plate, it appears that

this is probably a Wedelia species.

‘Blainvillea tenuicaulis Benth. & Hook.f.

Gen. PI. 2: 370 (1873)”. This combination is

listed in IPNI, but was only implied, not validly

made, in the place cited. Bentham & Hooker

actually used the name Wedelia tenuicaulis

Hook.f., which is of doubtful application,

(see Global Compositae Checklist: http://

compositae.landcareresearch.co.nz/Default.

aspx). - Wedelia tenuicaulis Hook.f., Trans.

Linn. Soc. London 20: 213 (1847). Type

citation: [Galapagos], “Albemarle Island, Mr

Orchard, Australian Blainvillea

667

Macrae” Type: Galapagos, Albemarle Island,

s.dat., McRae s.n. (holo: K 487623 photo!; iso:

Galapagos, s.dat., Macrae s.n. (US 385744

photo!).

Hooker, in describing Wedelia tenuicaulis ,

considered that it was close to W. discoidea,

and described it as having discoid capitula

with all florets tubular and hermaphrodite.

The type in K has attached to it a series of

drawings and notes by Hooker. These show a

palea which is of the Blainvillea type (broad,

truncate and laciniate apically), and an achene

which is fusiform with 2 erect awns. Although

the leaves are rather narrow for this species, it

seems likely that this is a rather depauperate

specimen of Blainvillea acmella {cf. notes

under B. acmella regarding B. dichotoma ),

and that the ray florets were either missing or

overlooked by Hooker. It is not Trigonopterum

(Macraea) laricifolia (Hook.f.) W.L.Wagner

& H.Rob.

Blainvillea lanceolata J.G.Baker, FI. Bras.

(Mart.) 6: 176 (1884). Type citation: “Habitat

in prov. Alto Amazonas in silvis Japurensibus:

Martius.”

Blainvillea dallo-vedovae Terrac., Annuar.

Reale 1st. Bot. Roma 5: 107 (1894). Type

citation: [Somalia], Reference not traced.

Probably B. gayana.

Blainvillea dichotoma Luetzelb., Estud. Bot.

Nordeste i. (Publ. Insp. Fed. Obras Contr.

Secc., Rio de Janeiro , No. 57, Ser. I.A.) 41

(1922-3).

Listed in Index Kewensis as a nom. nud., i.e.

nom. inval. Not to be confused with the (also

invalid) Verbesina dichotoma Wall. [= B.

acmella] or Verbesina dichotoma Murray (=

Blainvillea dichotoma (Murray) Hemsl.).

Acknowledgements

I thank the directors of herbaria who kindly

loaned or made available material for this

study (AD, BM, BRI, CANB, DNA, G,

HO, K, L, MEL, NE, NSW, PERTH). The

Australian Biological Resources Study

(ABRS) and Australian National Herbarium

(CANB) both provided office and research

facilities, and the work was carried out as part

of a research consultancy from ABRS. Three

overseas herbaria (BM, G, K) kindly provided

access to their collections during my visit in

May 2012, and their staff are thanked for

their hospitality. Nicholas Hind (K) provided

useful robust discussion on generic and

specific concepts in the Ecliptinae. Annette

Wilson (ABRS) kindly assisted with scanning

and manipulation of images, and suggested

improvements to a first draft. I also thank

ABRS for permission to use in this paper

figures 1-4, for which it holds the copyright.

These figures are destined for a future Flora

of Australia volume.

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A taxonomic revision of Hollandaea F.Muell. (Proteaceae)

A. J. Ford 1 & Peter H. Weston 2

Summary

Ford, A.J. & Weston, P.H. (2012). A taxonomic revision of Hollandaea F.Muell. (Proteaceae).

Austrobaileya 8(4): 670-687. The genus Hollandaea (Proteaceae) is revised, redescribed and

distinguished from its closest relative, the genus Helicia. Hollandaea porphyrocarpa A.J.Ford &

P.H.Weston and H. diabolica A.J.Ford & P.H.Weston are newly described, illustrated and diagnosed

from related species. Notes on habitat, distribution, and conservation status for all four species of

Hollandaea are provided. A key to the species of Hollandaea is presented.

Key Words: Proteaceae, Hollandaea, Hollandaea diabolica , Hollandaea porphyrocarpa , Hollandaea

riparia , Hollandaea sayeriana , Australia flora, Queensland flora, taxonomy, identification key, new

species

’A.J. Ford, CSIRO, Ecosystem Sciences, Tropical Forest Research Centre, PO Box 780, Atherton,

Queensland 4883, Australia.

2 Peter H. Weston, National Herbarium of New South Wales, Royal Botanic Gardens and Domain

Trust, Department of Environment & Conservation (NSW), Royal Botanic Gardens, Mrs Macquaries

Road, Sydney, NSW 2000, Australia.

Introduction

Hollandaea F.Muell. is a genus of four species

that is endemic to Australia (Cooper & Cooper

2004) with all species being confined to the

Wet Tropics area of north eastern Queensland.

The name Hollandaea first appeared in the

literature in April 1887 (Anon. 1887) prior to

its formal publication in June 1887 (Mueller

1887), being noted in a report of a meeting

at which Mueller exhibited specimens and

indicated his intention to publish.

Hollandaea sayeriana (F.Muell.)

L.S.Sm. was originally described as Helicia

sayeriana F.Muell. by Mueller (1886). The

following year, as outlined above, Mueller

erected Hollandaea and transferred Helicia

sayeriana to it, as H. sayeri F.Muell. (Mueller

1887). Unfortunately, this species name was

illegitimate because the specific epithet was

nomenclaturally superfluous when published.

Bailey (1899) described Hollandaea

lamingtoniana F.M.Bailey which appeared

as such in The Queensland Flora (Bailey

1901), but was rightly transferred to Helicia

Lour, by Smith (1952), who attributed the

new combination to C.T.White. Engler (1888)

overlooked or was unaware of these taxa in

Accepted for publication 30 May 2012

his treatment of the Proteaceae and Smith

(1956) made the legitimate combination H.

sayeriana. Hollandaea became accepted as a

distinct monotypic genus (see e.g. Johnson &

Briggs 1963; Venkata Rao 1971; Johnson &

Briggs 1975), differing from all other genera

of Proteaceae in producing a follicular fruit

containing numerous wingless seeds.

Johnson & Briggs (1963: 42) placed

Hollandaea in the subfamily Grevilleoideae

Engl, as a genus incertae sedis on the grounds

that it “... shows little advancement from

the primitive Grevilleoid condition, except

perhaps in its undivided leaves, thick wingless

seeds with obliquely arranged cotyledons,

fully adnate filaments, and bright pink flowers

(crimson in bud), which do not, however,

show any particular structural adaptation

to ornithophily” By “primitive Grevilleoid

condition” they meant a hypothetical

ancestral suite of states including follicular

fruits containing numerous, winged seeds, the

production of pseudoracemose inflorescences

bearing lateral pairs of actinomorphic flowers

and a haploid chromosome number of n = 14.

Venkata Rao (1971) agreed with Johnson &

Briggs in placing Hollandaea in subfamily

Grevilleoideae and placed it in his tribe

Telopeeae Venk.Rao on the basis of its multiple

Ford & Weston, Revision of Hollandaea

ovules and follicular fruit. Moreover, he placed

Hollandaea , together with Knightia R.Br. and

Darlingia F.Muell, in the (nomenclaturally

invalid) new subtribe Hollandinae Venk.Rao.

Venkata Rao’s Telopeeae was considered by

Johnson & Briggs (1975) to be polyphyletic,

as it included parts of their tribes Embothrieae

L.A.S.Johnson & B.G.Briggs, Knightieae

L.A.S.Johnson & B.G.Briggs and Helicieae

L.A.S.Johnson & B.G.Briggs. They argued

(Johnson & Briggs 1975: 108) that Hollandaea

had been shown by their studies to be “more

probably a member of the Heliciean branch

of the subfamily” and that it shared “only

primitive characters with Knightieae or

Embothrieae”. Johnson & Briggs’ tribe

Helicieae included Hollandaea in its own

subtribe, Hollandaeinae L.A.S.Johnson &

B.G.Briggs, as well as Triunia L.A.S.Johnson

& B.G.Briggs in the monotypic subtribe

Triuniinae L.A.S.Johnson & B.G.Briggs

and two other genera, Xylomelum Sm.

and Helicia, in the subtribe Heliciinae

L.A.S.Johnson & B.G.Briggs. These four

genera were grouped together on the basis of

the putatively synapomorphic loss of divided

pre-adult leaves, a character state that is found

in numerous other genera of Grevilleoideae.

These other genera were placed by Johnson

& Briggs in other tribes on the basis of other

putative synapomorphies.

Hollandaea remained a monotypic genus

until Hyland (1995) added a second species,

describing the highly restricted Hollandaea

riparia B.Hyland from the Roaring Meg

Creek area, south of Cooktown. This species

was sampled, along with species of 45 other

genera, in the first published molecular

phylogenetic analysis of the Proteaceae (Hoot

& Douglas 1998). Hoot and Douglas’ analysis

included genera that had been proposed by

Venkata Rao (1971 - Knightia) and Johnson &

Briggs (1975 - Helicia, Xylomelum , Triunia) to

be closely related to Hollandaea and showed

very strongly (100% bootstrap support) that

the sister group of Hollandaea is Helicia ,

contrary to all previous classifications. The

relationships of this clade were unresolved,

forming part of a polytomy of 11 lineages

that together constituted the Grevilleoideae

in their best estimate of relationships (Hoot

671

& Douglas 1998: 309: figure 2). The grouping

of Hollandaea and Helicia as sister taxa was

also found by Weston & Barker (2006) in their

supertree analysis of the results of published

and unpublished molecular phylogenies of the

Proteaceae. They recognised this clade as the

subtribe Heliciinae , one of four subtribes plus

four ungrouped genera in the tribe Roupaleae

Meisn. in their new classification of the

Proteaceae. Monophyly of the Roupaleae

is poorly supported as is resolution of its

internal relationships. Conflicting evidence

from different molecular data sets has

prevented identification of the sister group

of the Heliciinae with confidence: rbcL

cpDNA sequences strongly group Helicia

and Hollandaea with Knightia (Barker et

al. 2007) but this putative relationship is not

corroborated by analyses of other chloroplast

loci (Hoot & Douglas 1998) nor by analysis

of ITS nuclear ribosomal DNA sequences

(Weston & Barker 2006). A supermatrix

analysis of the Proteaceae conducted by

Sauquet et al. (2009), using all available

DNA sequences (1 nuclear and 7 chloroplast

regions) and phylogenetic analyses using

both Bayesian probability and maximum

parsimony criteria, again grouped Helicia

with Hollandaea as a clade, with a posterior

probability of 1.0 and 100% parsimony

bootstrap support. In this analysis, the sister

group of the Heliciinae was resolved as the

monotypic genus Megahertzia A.S.George &

B.Hyland, with a posterior probability of 0.99

but parsimony bootstrap support of only 59%.

Weston & Barker’s (2006) subtribe

Heliciinae is characterised morphologically

by the possession of anatropous ovules that

develop, on fertilization, into thick, unwinged

seeds. These states are found elsewhere in the

tribe Roupaleae. Xylomelum has anatropous

ovules and Triunia has fleshy and unwinged

seeds (Weston 2006). Detailed analysis of

the distribution of these character states in

the Proteaceae is needed to determine which,

if either, of them is synapomorphic for the

Heliciinae.

Hollandaea and Helicia are distinguished

morphologically from one another by their

ovary, fruit and seed morphology (Foreman

672

1995; Hyland 1995). The ovary of Hollandaea

contains ovules borne in an oblique

orientation from lateral placentae. Until

now, Hollandaea has been characterised as

consistently having four or more ovules per

ovary but this now needs to be amended, as

the here-described Hollandaea diabolica

has 2-4 ovules. On fertilization, these

develop into angular seeds that are released

at maturity from a follicular fruit. The ovary

of Helicia contains two erect ovules borne

from basal placentae. On fertilization, one (or

rarely both) of these develops into a globose

to ovoid or ellipsoid seed (hemispherical

to hemielliptical if both ovules develop),

enclosed within an indehiscent (drupaceous

or dry) fruit. Outgroup comparisons with

other members of the tribe Roupaleae and

with its sister group (tribe Banksieae Dumort.

plus Sphalmium B.G.Briggs, B.Hyland &

L.A.S. Johnson according to Weston & Barker

[2006]) suggest that indehiscent fruit may

be a synapomorphy for Helicia. Johnson

& Briggs (1975) assumed that numerous

ovules per carpel is plesiomorphous for the

Proteaceae as a whole and for all subgroups

that include multiovulate taxa. According to

this interpretation, reduction to two ovules

would also be a synapomorphy for Helicia.

However, outgroup comparison suggests

the possibility that multiple ovules might

be more parsimoniously interpreted as a

synapomorphy for Hollandaea.

In the early 1990s two new proteaceous

taxa were observed and collected from the

vicinity of a walking track on the Main Coast

Range near Mossman, north-east Queensland.

Significantly, one of these taxa (described

below as Hollandaea diabolica A.J.Ford &

P.H.Weston) was investigated by Carpenter

(1994) for leaf cuticle morphology. In his

opinion, “cuticular morphology.... indicates

that this taxon cannot be aligned with any

described genus, although it is most likely

to be allied to Helicieae”. Since that time,

additional specimens have been collected

and new locations of each species have

been documented. The currently accepted

morphological circumscription of Hollandaea

includes one of these taxa and needs to be

Austrobaileya 8(4): 670-687 (2012)

modified only minimally, by expanding the

range of ovule numbers to 2-20, to encompass

the other. Both taxa are described below as

new species of Hollandaea.

Materials and methods

The study is based upon examination of

herbarium material from BRI, CNS and NSW

with field observations by the first author. All

specimens cited have been seen by one or

both authors.

Measurements of the floral parts and fruits

of Hollandaea are based on material preserved

in 70% ethanol. Common abbreviations in

the specimen citations are: dbh (diameter

at breast height), E.P (Experimental Plot),

L.A. (Logging Area), N.P.R. (National Park

Reserve), S.F.R. (State Forest Reserve) and

T.R. (Timber Reserve).

Estimates of extent of occurrence sensu

IUCN (2001) were derived from validation

of original collection localities. These data

points were loaded into ESRI ArcView 3.2 and

the draw polygon feature used to calculate the

area between the points. Area of occupancy

estimates were derived from a digital Regional

Ecosystem map together with the first author’s

knowledge of vegetation types and habitats

within the Wet Tropics bioregion (hereafter

referred to as the Wet Tropics) (Environment

Australia 2005). These estimates are not

strictly RE driven, therefore they possibly

represent a relatively more accurate picture of

occurrence and occupancy.

Species names in the distribution and

habitat notes are those that are currently

accepted by the Queensland Herbarium

(BRI).

The abbreviation RE in the distribution and

habitat notes refers to Regional Ecosystem,

descriptions of which can be viewed at:

www.derm.qld.gov.au/wildlife-ecosystem/

biodiversity/regional_ecosystems/index.php

NCA is an abbreviation for the Queensland

Nature Conservation Act (1992) and its

associated schedules. Discussions of

conservation status are made in reference to

the criteria of the IUCN (2001).

Ford & Weston, Revision of Hollandaea

673

Species are arranged alphabetically.

Suggested affinities between species are

indicated in the Affinities section for each

taxon.

Character phylogenies of qualitative

morphological characters were reconstructed

by mapping them onto a phylogenetic tree of

proteaceous genera (Sauquet et al. 2009) using

the parsimony option in Mesquite version

2.75. The following characters (and their

states) were analysed in this way: trichome

morphology (basifixed or medifixed),

ovule form (anatropous, hemitropous or

orthotropous); ovule number (two, more

than two), ovule placentation (basal, lateral,

apical), micropyle orientation (micropyle

pointing towards base of locule or pointing

obliquely across locule), cotyledon thickness

(laminar or fleshy), seed wing (absent or

present).

Character phylogenies

The four character states that diagnose the

subtribe Heliciinae from other members of

the tribe Roupaleae - possession of medifixed

trichomes, anatropous ovules, fleshy

cotyledons, and unwinged seeds were all

resolved unequivocally as synapomorphies

for the Heliciinae on the tree of Sauquet et al.

(2009). Of the character states distinguishing

Hollandaea from Helicia , multiple ovules was

resolved as synapomorphic for Hollandaea

and indehiscent fruit as synapomorphic for

Helicia.

Taxonomy

Hollandaea F.Muell., Australas. Chem.

Druggist 2(6): 173 (June 1887). Type: H.

sayeriana (F.Muell.) L.S.Sm.

Derivation of name : Named after Sir Henry

Thurston Holland, first Viscount Knutsford

(1825-1914), Secretary of State for the

Colonies between 1887 and 1892.

Single or multistemmed trees or large shrubs.

Trichomes medifixed. Bark compact and

close, usually inconspicuously lenticellate,

lacking any significant features. Leaves

alternate, spirally inserted, simple; seedlings

lacking cataphylls; juvenile leaves unlobed,

prominently toothed or rarely entire; adult

leaves entire or inconspicuously toothed,

venation brochidodromous, primary venation

conspicuous on both surfaces, secondary

venation discernible, tertiary venation

discernible. Petioles swollen, pulvinate.

Conflorescence an elongated unbranched

raceme of flower pairs, lateral, cauliflorous,

ramiflorous or borne just proximal to the

lowest/oldest leaves, usually pendulous,

composed of 54-224 flowers; each flower

pair subtended by a scale-like bract (common

bract); common peduncle present. Flowers

bisexual, pedicellate on common peduncle,

hypogynous. Floral bract conspicuous,

inserted variously along pedicel. Perianth

actinomorphic, differentiated into a narrow

basal claw and clavate apical limb in mature

bud; tepals 4, not connate, glabrous adaxially,

glabrous or hairy abaxially, splitting with

individual tepals coiling and reflexed at

anthesis, caducous following anthesis.

Stamens 4, equal; filaments adnate to tepals;

anthers 2-locular, with a distinct mucro,

dehiscing introrsely through longitudinal

slits. Hypogynous glands 4, free or connate,

fleshy. Gynoecium glabrous; ovary sessile,

ovules anatropous, obliquely oriented, 2-20;

placentae lateral; style straight; tip clavate,

functioning as a pollen presenter; stigma

terminal. Fruit dehiscent, follicular, usually

asymmetrical; style persistent. Seeds fleshy,

unwinged, usually angular, 1-20 per fruit,

testa thin. Germination usually hypogeal.

Cotyledons fleshy. 2 n = 28

Distribution : Endemic to the Wet Tropics,

Queensland, Australia; four species.

674

Austrobaileya 8(4): 670-687 (2012)

Key to the species of Hollandaea

The following key allows for both flowering and fruiting specimens. (See Cooper & Cooper

2004: 414, 415 for exquisite fruit illustrations).

1 Flowers present.2

1. Fruit present.5

2 Tepal abaxial surface clothed in medifixed hairs.3

2. Tepal abaxial surface glabrous.4

3 Leaf base cuneate, conflorescence axis > 150 mm long.4. H. sayeriana

3. Leaf base attenuate, conflorescence axis < 110 mm long.2. H. porphyrocarpa

4 Leaf margin recurved, leaf apex obtuse to retuse, conflorescence axis

hairy, tepals creamish, yellowish or with green hue.1. H. diabolica

4. Leaf margin flat, leaf apex obtuse to acute, conflorescence axis glabrous,

tepals purplish or purple with blue-green hue.3. H. riparia

5 Fruit surface sculptured.6

5. Fruit surface smooth.7

6 Fruit smoothly (and shallowly) wrinkled to the touch, seeds 14-20 mm

long, leaf apex acute-obtuse.3. H. riparia

6. Fruit roughly (and deeply) wrinkled to the touch, seeds 22-25 mm long,

leaf apex retuse-obtuse.1. H. diabolica

7 Fruit green when ripe, > 60 mm long.4. H. sayeriana

7. Fruit purple when ripe, < 50 mm long.2. H. porphyrocarpa

1. Hollandaea diabolica A.J.Ford &

P.H.Weston, species nova. Distinguished from

H. riparia by fruit shape (± equidimensional

versus longer than wide), seed size (22-25 mm

versus 14-20 mm), leaf apex (obtuse-retuse

versus obtuse-acute), leaf length:width ratio

(<3.6:1 versus >4:1), tepal colour (creamish or

yellowish versus purplish) and ovule number

(2-4 versus 6-8). Typus: Queensland. Cook

District: Pinnacle Rock Track [Daintree

National Park, NW of Mossman], 1 February

1996, B. Hyland 25914RFK (holo: BRI; iso:

CNS, NSW).

Proteaceae sp. ‘Devils Thumb’; Carpenter

(1994: 291, 292).

Orites sp. (Pinnacle Rock Track WWC 867);

Hyland et al. (2003).

Hollandaea sp. (Devils Thumb); Cooper &

Cooper (2004: 414).

Hollandaea sp. (Devils Thumb P.I.Forster+

PIF10720); Forster & Edginton (2007: 171;

2010: 165).

Illustrations : Cooper & Cooper (2004: 414)

as Hollandaea sp. (Devils Thumb); Hyland et

al. (2003) as Orites sp. (Pinnacle Rock Track

WWC 867).

Single stemmed canopy or subcanopy trees to

25 m high, with trunk diameters to 50 cm dbh

recorded; buttresses absent. Bark nondescript.

Terminal and axillary buds clothed in dark

brown, minute, medifixed hairs. Branchlets

initially somewhat angled and sparsely clothed

in mostly pale-coloured minute medifixed

hairs, becoming terete and glabrous.

Seedlings: first leaves sub-opposite, toothed,

stem hairy. Juvenile leaves simple, unlobed,

prominently toothed. Leaves alternate,

petiolate, discolorous, dull or slightly shiny

on adaxial surface and very pale (± pruinose)

on abaxial surface; lamina elliptic to elliptic -

obovate, 69-155 x 20-55 mm, base attenuate,

apex retuse to obtuse with acumen absent;

margin usually recurved; both surfaces

glabrous although rare pale-coloured minute

medifixed hairs may be present on midvein

on adaxial surface; midvein raised on each

Ford & Weston, Revision of Hollandaea

675

surface, more prominent on abaxial surface;

venation slightly bullate on adaxial surface,

equally conspicuous on both surfaces, primary

venation conspicuous on both surfaces with

5-8 primary lateral veins on each side of

midvein. Petioles slightly swollen, 3-4 mm

long, slightly convex on adaxial surface;

glabrous or with rare, minute, dark-coloured

medifixed hairs, appearing to be winged and

much longer due to the attenuate nature of

the leaf base. Conflorescences ramiflorous or

just below the lowest/oldest leaves, composed

of 54-140 flowers; conflorescence axis 48-

120 mm long, weakly angled (not terete),

longitudinally striated (when dry), moderately

clothed in brown medifixed hairs. Common

bracts c. 0.7 mm long, ovate, with mostly

marginal hairs. Floral bracts c. 0.5 mm long,

ovate, glabrous except for minute pale brown

marginal hairs, inserted usually about halfway

along pedicel. Pedicels terete, 2.1-3 mm long,

sparsely clothed in brown medifixed hairs,

paired on a peduncle c. 1 mm long. Tepals

27-33 mm long and clavate at apex in mature

bud, splitting distally at first with individual

tepals coiling and reflexed at anthesis, apex

acute, cream, yellow or greenish, glabrous on

abaxial and adaxial surface, Stamens with

free filament tips 0.4-0.8 mm long, inserted

c. 5 mm from tepal apex; anthers 4-5 mm

long (including a blunt appendage/mucro of

c. 0.3 mm long). Hypogynous glands free,

fleshy. Style 20-24 mm long; pollen presenter

3-4 mm long; ovary 1.9-2.6 mm long,

ovules 2-4, placenta marginal. Fruit 40-50

mm diameter, asymmetrical but appearing

globose to spherical, deeply wrinkled (with a

honeycomb texture), orange-green, leathery

on outer surface, woody on inner surface,

style persistent. Seeds 1-4 per fruit, 22-25

mm long on the longest axis, usually smooth,

testa thin, radicle c. 2mm long. Germination

hypogeal. Fig. 1.

Additional specimens examined : Queensland. Cook

District: Pinnacle Rock Track, 4 km W of Karnak, Jun

1992, Forster PIF10720 et al. (BRI, CNS); Pinnacle Rock

Track, Feb 1996, Hyland 25913RFK (CNS); loc. ciX., Feb

1996, Hyland 25909RFK (CNS); West of Karnak, via

Mossman, Jan 1995, Cooper 867 & Cooper (BRI, CNS);

loc. cit ., Jan 1995, Cooper 868 & Cooper (BRI, CNS);

Daintree N.P, Pinnacle Rock Track, NW of Mossman,

just before the Gleichenia area, Oct 2005, Ford 4747 et

al. (BRI, NSW, CNS); loc. cit. , Oct 2005, Ford 4750 et

al. (BRI, NSW, CNS); Pinnacle Rock, Whyanbeel, Sep

1991, Sankowsky 1205 (CNS); East Mulgrave River,

Bellenden Ker, Nov 1995, Jensen 525 (CNS).

Distribution and habitat : Hollandaea

diabolica is endemic to the Wet Tropics

bioregion in north-eastern Queensland,

where it is currently known to occur on the

eastern fall of the Main Coast Range west of

Mossman (see above) and an area on the East

Mulgrave River to the south of Mt Bellenden

Ker (Map 1). On the Main Coast Range it

inhabits very wet mountainous notophyll

vine-forests/rainforests on soils derived

from granite. Here the common canopy

species include: Acmena hemilampra subsp.

orophila B.Hyland, Balanops australiana

F.Muell., Elaeocarpus sp. (Mossman Bluff

D.G.Fell 1666), Halfordia kendack (Montrouz.)

Guillaumin, Planchonella euphlebia (F.Muell.)

W.D.Francis, Syzygium spp. and Sphalmium

racemosum (C.T.White) B.G.Briggs, B.Hyland

& L.A.S.Johnson. Common small trees and

shrubs include: Chionanthus axillaris R.Br.,

Haplostichanthus submontanus Jessup subsp.

submontanus, Linospadix minor (W.Hill)

F. Muell., Oraniopsis appendiculata (F. M. Bailey)

J.Dransf., A.K.Irvine & N.W.Uhl, Pittosporum

rubiginosum A.Cunn., Schistocarpaeajohnsonii

F.Muell., Steganthera cooperorum Whiffin,

S. macooraia (F.M.Bailey) P.K.Endress and

Symplocos graniticola Jessup. No floristic

assemblage information is available for the

East Mulgrave River population. Altitudinal

range, from existing specimens, is 450-1000

Hollandaea diabolica has been collected

in RE7.12.16a (East Mulgrave and Main Coast

Range populations) and 7.12.20 (Main Coast

Range population), the former regarded as a

very common and widespread community

within the Wet Tropics. However, it is very

likely that the two populations have different

floristic associations as they are on either

side of the Black Mountain Corridor (BMC)

and several of the species listed above do not

occur south of the BMC. Although both H.

diabolica and H. porphyrocarpa occur in RE

7.12.20 they have not yet been recorded as co¬

occurring.

Fig. 1. Hollandaea diabolica. A. branchlet with inflorescences x0.4. B. paired flowers at anthesis showing floral bracts

x3. C. close up of ovary showing hypogynous glands and floral bract x3. D. lateral view of fruit with persistent style

xl. A-C from Hyland 25914RFK (CNS); D from Cooper 868 & Cooper (CNS). Del. W.Smith

Ford & Weston, Revision of Hollandaea

677

Phenology: Flowers have been recorded

in January and February; fruits have been

recorded in January.

Notes: The woody remains of the fruit are

usually conspicuous under mature trees.

These remains, and fallen yellowish leaves,

are often the only clues that this species is in

the area. The perianth is recorded as being

creamish to yellow or greenish; the style is

pink to mauve and cream at the apex; the

ovary is pink and hypogynous glands are

yellowish.

Juvenile leaves have an acute-acuminate

apex and lack the conspicuous retuse-

obtuse apex of adult leaves. With such an

apex, seedlings (and saplings) could easily

be confused with local glabrous species of

Helicia.

The East Mulgrave River collections are

sterile and no fruit remnants were recorded.

Until fertile collections of this population

are made to suggest otherwise we have

incorporated that population’s data into

the above descriptions and the following

assessment.

Affinities: Hollandaea diabolica appears

to be most closely related to H. riparia, in

that they share the features of ± pruinose

abaxial leaf surface, wrinkled/verrucose

fruit surface, glabrous abaxial tepal surface

and a relatively low ovule number. H.

diabolica can be distinguished from H.

riparia on the following features: fruit shape

(± equidimensional versus longer than wide,

respectively), larger seeds (22-25 mm versus

14-20 mm), leaf apex (obtuse-retuse versus

obtuse-acute), leaf length:width ratio (<3.6:1

versus >4:1), usually shorter conflorescence

axis (48-120 mm versus 110-155 mm) and

fewer ovules (2-4 versus 6-8). Comparisons

between all species of Hollandaea are

provided in Table 1.

Conservation Status. All existing collections

have been made within the World Heritage

Area of the Wet Tropics bioregion. Hollandaea

diabolica only occurs in the Daintree and

Wooroonooran National Parks. As it has a

very disjunct and narrow geographical range,

with an extent of occurrence estimated to be

Map 1. Distribution of Hollandaea species in north-east

Queensland (H. diabolica ▼, H. porphyrocarpa •, H.

riparia ▲, H. sayeriana ■)

less than 50 km 2 and an area of occupancy

estimated to be less than 5 km 2 , it is considered

at risk at this time. An approximate estimate

of the population sizes is not known, but

an optimistic guess of less than 100 mature

individuals on the Main Coast Range, and

perhaps a similar number at the East Mulgrave

River is neither conservative nor extravagant.

A thorough search of nearby areas is required

to determine the population size and structure.

Nonetheless, due to the extremely disjunct

nature, limited distribution and estimated

population sizes we recommend H. diabolica

678

Austrobaileya 8(4): 670-687 (2012)

being listed as “Vulnerable” under the IUCN

(2001) criteria as it fulfills the criteria under

categories VU, D1 and D2.

Etymology : The specific epithet has been

formed arbitrarily and comes from the Latin

diabolus, (devil) in reference to the locality,

Devils Thumb.

2. Hollandaea porphyrocarpa A.J.Ford &

P.H.Weston, species nova. Distinguished

from H. sayeriana by the leaf base (attenuate

versus cuneate), fruit size (35-45 mm versus

60-150 mm), conflorescence axis length

(52-103 mm versus 150-380 mm) and

anther length (4.5-6 mm versus 2-3.1 mm).

Typus: Queensland. Cook District: Daintree

National Park, Pinnacle Rock Track, NW

of Mossman, beyond the Gleichenia area,

13 October 2005, A. Ford 4742, W.Cooper &

R.Russell (holo: BRI [2 sheets + spirit]; iso:

CNS, K, L, MEL, MO, NSW).

Hollandaea sp. (Pinnacle Rock Track

PI.Forster+ PIF10714); Forster & Edginton

(2007: 171; 2010: 165).

Hollandaea sp. (Pinnacle Rock Track

PIF10714); Hyland etal. (2003).

Hollandaea sp. (Pinnacle Rock Track);

Cooper & Cooper (2004: 415).

Illustrations : Cooper & Cooper (2004: 415)

as Hollandaea sp. (Pinnacle Rock Track);

Hyland et al. (2003) as Hollandaea sp.

(Pinnacle Rock Track PIF10714).

Single stemmed canopy or subcanopy trees

to 15 m high, with trunk diameters to 20 cm

dbh recorded; buttresses absent. Terminal

and axillary buds clothed in minute, dark

brown hairs. Branchlets terete, glabrous to

glabrescent, smooth. Seedlings: cotyledons

obovate, 15-17 x 13-17 mm, base sagittate;

hypocotyl glabrous, first leaves sub¬

opposite, toothed, stem hairy. Juvenile leaves

prominently toothed with 4-8 teeth on each

side. Adult leaves petiolate, discolorous, dull

on adaxial and abaxial surfaces; lamina elliptic

to elliptic-obovate, 90-157 x 32-60 mm, base

attenuate, apex acuminate to acute-acuminate

with acumen 3-12 mm long; margin flat

or slightly recurved, entire or sparsely

denticulate, both surfaces glabrous; midvein

raised on each surface, more prominently

so on abaxial surface; faintly 3-veined at

base, primary venation conspicuous on both

surfaces with 6-10 primary lateral veins on

each side of the midvein, equally conspicuous

on both surfaces. Petioles swollen, pulvinate,

4-6 mm long, ± terete, glabrous, appearing to

be winged and much longer than they are due

to the very attenuate shape of the leaf base.

Conflorescences cauliflorous, ramiflorous

or just below the lowest/oldest leaves,

composed of 58-106 flowers; conflorescence

axis 52-103 mm long, weakly angled (not

terete) when dry, densely clothed in pale to

mid-brown hairs. Common bracts c. 0.9 mm

long, lanceolate, with mostly marginal hairs.

Common peduncle of flower pair c. 2 mm

long. Floral bracts triangular to ovate, c. 0.7

mm long, glabrous except for minute, pale

brown marginal hairs, inserted usually about

halfway along pedicel. Pedicels terete, 3-4.5

mm long, clothed in pale brown hairs. Tepals

26-29 mm long in mature bud, splitting

proximally at first with individual tepals

coiling and reflexed at anthesis, apex acute,

pink, moderately clothed on abaxial surface

in pale brown hairs, glabrous adaxially.

Stamens with free filament tips c. 0.4 mm

long, inserted c. 7 mm from tepal apex;

anthers 4.5-6 mm long (including a mucro of

c. 0.7 mm long). Hypogynous glands connate,

fleshy. Style 16-20 mm long; pollen presenter

c. 5 mm long; ovary 2.5-3 mm long, ovules

12-16, placenta marginal. Fruit leathery

(neither fleshy nor woody), 35-45 mm long, c.

30 mm diameter, asymmetrical and somewhat

hemispherical in shape with one side nearly

flat, smooth, purplish, style persistent. Seeds

angular, many-faced with the abaxial face

convex, 6-9 per fruit, 13-16 mm long on

the longest axis, testa thin, radicle c. 3mm

long, adaxial surface of cotyledons flat.

Germination epigeal to hypogeal (cotyledons

usually splitting the testa and forming a

distinctive hypocotyl). Fig. 2.

Additional specimens examined : Queensland. Cook

District: Pinnacle Rock Track, 4.5 km W of Karnak,

Mossman, Jun 1992, ForsterPIF10714 etal. (BRI, CNS);

Pinnacle Rock Track, Feb 1996, Hyland 25910RFK

(CNS); Ridges above the Mossman River in the Mossman

Gorge, May 1991, Russell s.n. (CNS); Pinnacle Rock,

Whyanbeel, Sep 1991, Sankowsky 1206 (CNS); West of

Ford & Weston, Revision of Hollandaea

679

Fig. 2. Hollandaeaporphyrocarpa. A. branchlet with inflorescences *0.4. B. paired flowers at anthesis x3. C. close

up of ovary showing hypogynous glands and floral bract x3. D. lateral view of fruit with persistent style xf A-C from

Ford 4742 et al. (BRI); D from Cooper 865 & Cooper (CNS). Del. W.Smith

680

Karnak, via Mossman, Dec 1994, Cooper 865 & Cooper

(BRI, CNS); Daintree N.P., Pinnacle Rock Track, NW of

Mossman, beyond the Gleichenia area, Oct 2005, Ford

4745, Cooper & Russell (BRI, CNS, NSW).

Distribution and habitat : Hollandaea

porphyrocarpa is endemic to the Wet Tropics

bioregion in north-eastern Queensland,

where it is currently known to occur on

the eastern fall of the Main Coast Range

west of Mossman (Map 1). This area is

locally known as either “Devils Thumb”

or “Pinnacle Rock”. It inhabits very wet

mountainous notophyll to microphyll vine-

forests or vine-fern thickets/rainforests on

soils derived from granite. Common canopy

species include: Cryptocarya corrugata

C.T.White & W.D.Francis, Elaeocarpus elliffii

B. Hyland & Coode, Halfordia kendack,

Myrsine oreophila Jackes, Niemeyera sp.

(Mt Lewis A.K.Irvine 1402) and Sphalmium

racemosum. Common small trees and shrubs

include: Ardisia pachyrrhachis (F.Muell.)

F.M.Bailey, Austromuellera valida B.Hyland,

Baloghia parviflora C.T.White, Catalepidia

heyana (F.M.Bailey) P.H.Weston, Chionanthus

axillaris , Pittosporum rubiginosum, Psychotria

spp., Steganthera cooperorum , Triunia montana

(C.T.White) Foreman and Wendlandia connata

C. T.White. Conspicuous understory species

include: Cyathea rebeccae (F.Muell.) Domin,

Linospadix apetiolata Dowe & A.K.Irvine

and Morinda podistra Halford & A.J.Ford.

Altitudinal range, from existing specimens, is

1000-1090 m.

Hollandaea porphyrocarpa has only been

collected in RE7.12.20.

Phenology : Flowers have been recorded in

May, June, September and October; fruits

have been recorded in December.

Notes : The commonly recorded cauliflorous

inflorescences make this species unmistakable

in the field. This is one of only three

proteaceous species in the tropical rainforests

of northern Queensland to have this feature.

The perianth is rosy pink in bud and at

anthesis; the style is violet at the base and

the apex, whilst the mid sections are white;

hypogynous glands are creamish yellow and

the ovary is purple. New leafy growth is

Austrobaileya 8(4): 670-687 (2012)

recorded as being green.

The flowers have a slight sweet scent and

Bridled Honeyeaters {Lichenostomusfrenatus

(Ramsay)) have been observed visiting them.

Affinities : Hollandaea porphyrocarpa

appears to be most closely related to H.

sayeriana , in that they share a smooth

fruit surface, abaxial tepal surface clothed

in medifixed hairs and a relatively high

ovule number. H. porphyrocarpa can be

distinguished from H. sayeriana on the

following features: leaf base (attenuate versus

cuneate, respectively), smaller fruit (35-45

mm versus 60-150 mm), smaller seeds

(13-16 mm versus [12-] 20-30 mm), shorter

conflorescence axis (52-103 mm versus 150—

380 mm) and longer anthers (4.5-6 mm versus

2-3.1 mm). Comparisons between all species

of Hollandaea are provided in Table 1.

Conservation status : All existing collections

have been made within the World Heritage

Area of the Wet Tropics bioregion. Hollandaea

porphyrocarpa has only been collected

in one mountainous area, mostly along a

walking track, within the Daintree National

Park, west of Mossman. It has a very narrow

geographical range, from the Pinnacle Rock

Track area south towards the Mossman River

(R. Russell, personal communication, 2006),

with an extent of occurrence estimated to be

less than 30 km 2 and an area of occupancy

estimated to be less than 15 km 2 and is

considered at risk at this time. An estimate

of the size of the single population is not

known, but an optimistic guess of less than

300 mature individuals is not extravagant. A

thorough search to cover areas away from the

walking track is necessary to gain a better

understanding of the population size and

structure. Nonetheless, due to the extremely

narrow distribution and estimated population

size we would recommend H porphyrocarpa

being listed as “Vulnerable” under the IUCN

(2001) as it fulfills the criteria under categories

VU, D1 and D2.

Etymology : The specific epithet is derived

from the Greek porphyreos (purple, dark red)

and carpos (fruit) and alludes to the unique

purple-coloured fruit of this species.

Ford & Weston, Revision of Hollandaea

681

3. Hollandaea riparia B.Hyland, FI.

Australia 16: 499 (1995). Type: Queensland.

Cook District: Timber Reserve 165, Baird

Logging Area, Qld, 22 September 1980,

B.Hyland 10626 (holo: CNS; iso: BRI; CANB

n.v.).

Illustrations : Cooper & Cooper (2004:

414); Hyland (1995: Fig. 139); Nicholson &

Nicholson (2000: 38); Hyland et al. (2003).

Single or usually multistemmed large shrubs

or small trees 4-6 m high, with trunk

diameters to 15 cm dbh recorded; buttresses

absent. Bark nondescript. Terminal and

axillary buds clothed in dark brown, minute

medifixed hairs. Branchlets initially slightly

angled to terete and glabrous, the angled

branchlets becoming terete. Seedlings:

cotyledons with peltate base; first leaves

alternate, toothed; stem hairy. Juvenile leaves

simple, unlobed, prominently to sparsely

toothed (or rarely entire). Leaves alternate,

petiolate, discolorous, dull or slightly shiny

on adaxial surface and very pale to pruinose

on abaxial surface; lamina narrow-elliptic

or oblong to oblanceolate-obovate, 71-205

x 12-39 mm, base attenuate, apex obtuse-

acute with acumen absent; margin flat when

fresh and slightly recurved when dry; both

surfaces glabrous with occasional dark-

coloured medifixed hairs present on midvein

on abaxial surface; midvein raised on each

surface, more prominent on abaxial surface;

venation equally conspicuous on both

surfaces, primary venation conspicuous on

both surfaces with 6-9 primary lateral veins

on each side of midvein. Petioles slightly

swollen, 0.5-3 mm long, ± flat on adaxial

surface, glabrous or with very rare minute,

dark coloured medifixed hairs, appearing

to be winged and much longer due to the

extremely attenuate nature of the leaf base.

Conflorescences ramiflorous or just below the

lowest/oldest leaves, composed of 130-198

flowers; conflorescence axis 110-155 mm

long, weakly angled (not terete), longitudinally

striated (when dry), glabrous. Common bracts

narrow-ovate, c. 0.8 mm long, glabrous except

for one or two long hairs at apex. Floral bracts

ovate-lanceolate, c. 0.7 mm long, glabrous

except for one or two minute hairs at the apex,

inserted at base or apex of pedicel. Pedicels

terete, 1.9-2.2 mm long, glabrous, paired on a

peduncle c. 1 mm long. Tepals 28-32 mm long

and clavate at apex in mature bud, splitting

proximally at first with individual tepals

coiling and reflexed at anthesis, apex acute,

purplish to purple-green-bluish, glabrous

on abaxial and adaxial surface. Stamens

with filaments 0.1-0.3 mm long, inserted c.

6 mm from tepal apex; anthers 5.5-6 mm

long (including a blunt appendage/mucro of

c. 0.6 mm long). Hypogynous glands free,

the apex with a fringe of finger-like papillae,

fleshy. Style 21-25 mm long; pollen presenter

3.2-4.3 mm long; ovary 2.2-2.6 mm long,

ovules 6-8, placenta marginal. Fruit leathery

on both surfaces, 25-50 mm long x 22-26

mm diameter, asymmetrical, semi-discoid

(appearing “half-moon” shaped in side view),

deeply wrinkled (verrucose), green and

pruinose, style persistent. Seeds smooth or

angular, 2-8 per fruit, 14-20 mm long on the

longest axis, testa thin, radicle c. 1.2 mm long.

Germination hypogeal. Fig. 3.

Additional specimens examined : Queensland. Cook

District: Roaring Meg Creek, Oct 1984, Sankowsky s.n.

(CNS); T.R. 165 Roaring Meg Creek, Apr 1997, Ford

1887 (BRI, CNS); loc. cit ., Nov 1996, Ford 1808 (BRI,

CNS); T.R. 165 Alexandra L A, Jun 1977, Hyland 9390

(CNS); T.R. 165 Noah Alexandra L.A, Roaring Meg

Creek, Oct 1997, Ford 2002 (CNS); T.R. 106 Parish of

Noah Baird L.A, Roaring Meg Creek, Jul 1997, Hyland

260I9RFK (CNS); loc. cit., Jul 1997, Hyland 26020RFK

(CNS).

Distribution and habitat : Hollandaea riparia

is endemic to the Wet Tropics bioregion

in north-eastern Queensland, where it is

currently only known to occur as a rheophyte

in the Roaring Meg Creek catchment (south

of Cooktown) (Map 1). It inhabits the

riparian zone of creek sides in notophyll-

mesophyll vine-forests/rainforests on alluvial

soils derived from granite. The dominant

canopy species is Xanthostemon chrysanthus

(F.Muell.) F.Muell. ex Benth. Other canopy

species include Acmena hemilampra (F.Muell.

ex F.M.Bailey) Merr. & L.M.Perry subsp.

hemilampra , Blepharocarya involucrigera

F.Muell., Buckinghamia ferruginiflora

Foreman & B.Hyland, Gymnostoma

australiana L.A. S. Johnson, Ormosia

ormondii (F.Muell.) Merr. and Tristaniopsis

682

Austrobaileya 8(4): 670-687 (2012)

Fig. 3. Hollandaea riparia. A. branchlet with inflorescences *0.4. B. paired flowers at anthesis showing floral bracts

x3. C. close up of ovary showing hypogynous glands and floral bract x3. D. lateral view of fruit with persistent style

xl. A-C from Ford 1887 (CNS); D from Ford2002 (CNS). Del. W. Smith

Ford & Weston, Revision of Hollandaea

683

exiliflora (F.Muell.) Peter G.Wilson &

J.T.Waterh. Common small trees and shrubs

include Acronychia acronychioides (F.Muell.)

T.G.Hartley, Chionanthus ramiflorus,

Choriceras majus Airy Shaw, Diospyros

sp. (Baird LA B.P.Hyland 9374), Dinghoua

globularis (Ding Hou) R.H.Archer and

Phyllanthus brassii C.T.White. Altitudinal

range, from existing specimens, is 250-350

Hollandaea riparia has been collected in

RE7.3.49a (usually) and 7.3.10a (rarely).

Phenology : Flowers have been recorded in

April and June; fruits have been recorded

from September to November.

Notes : The perianth is recorded as purplish

to purple-green-bluish; the style is pale

pink to pink-purple; the ovary is pink and

hypogynous glands are yellowish. Juvenile

leaves and adult leaves are similar.

Affinities : Hollandaea riparia appears to

be most closely related to H. diabolica in

that they share the features of ± pruinose

abaxial leaf surface, wrinkled/verrucose fruit

surface, glabrous abaxial tepal surface and a

relatively low ovule number. H. riparia can

be distinguished from H. diabolica on the

following features: fruit shape (longer than

wide versus ± equidimensional, respectively),

smaller seeds (14-20 mm versus 22-25 mm),

leaf apex (obtuse-acute versus obtuse-retuse),

leaf length:width ratio (>4:1 versus <3.6:1),

usually longer conflorescence axis (110-155

mm versus 48-120 mm) and more ovules (6-8

versus 2-4). Comparisons between all species

of Hollandaea are provided in Table 1.

Conservation Status : All existing collections

have been made within the World Heritage

Area of the Wet Tropics bioregion within

Timber Reserve 165. Currently, Hollandaea

riparia is listed under the NCA as Vulnerable.

It has a very narrow geographical range,

with an extent of occurrence estimated to be

less than 20 km 2 and an area of occupancy

estimated to be less than 5 km 2 , and is

considered at risk at this time. An approximate

estimate of the population sizes is not known,

but an optimistic guess of less than 300

mature individuals is neither conservative

nor extravagant. Nonetheless, due to the

extremely limited distribution and estimated

population sizes we agree that H. riparia be

listed as “Vulnerable” under the IUCN (2001)

as it fulfills the criteria under categories VU

D1 and D2.

Etymology : The specific epithet is derived

from the Latin word riparius (the bank of

a stream) and alludes to the habitat of this

species.

4. Hollandaea sayeriana (F.Muell.) L.S.Sm.,

Proc. Roy. Soc. Queensland 67: 39 (1956);

Helicia sayeriana F.Muell., Viet. Naturalist

3: 93 (1886); Hollandaea sayeri F.Muell.,

Australas. Chem. Druggist 2: 173 (1887),

nom. illeg. Type: Queensland. Cook District:

Mt Bellenden Ker, s.dat., W.Sayer s.n. (holo:

MEL).

Illustrations : Williams (1984: 159); Wrigley

& Fagg (1991: 404); Nicholson & Nicholson

(1994: 41); Hyland (1995: 392); Hyland et al.

(2003); Cooper & Cooper (2004: 415).

Single-stemmed subcanopy trees to 17 m

high, with trunk diameters to 30 cm dbh

recorded (but usually much less); buttresses

absent. Bark nondescript. Terminal and

axillary buds clothed in minute, dark

brown medifixed hairs. Branchlets initially

slightly angled and clothed in minute red-

brown medifixed hairs, becoming terete and

glabrous, smooth. Seedlings: first leaves

alternate; toothed, stem hairy. Juvenile

leaves simple, unlobed, prominently toothed.

Leaves alternate, petiolate, discolorous, dull

on adaxial and abaxial surfaces, the abaxial

surface pale lime-green; lamina broadly

elliptic to obovate, 90-260 x (52-)90-171

mm, base cuneate, apex acute-obtuse to

acuminate with acumen to 8 mm long; margin

flat or slightly recurved, denticulate or rarely

entire; abaxial surface glabrescent; midvein

raised on each surface, more prominent

on abaxial surface, on adaxial surface the

midvein is raised proximally and depressed

distally; venation equally conspicuous on

both surfaces, primary venation conspicuous

on both surfaces with 7-10 primary lateral

veins on each side of the midvein. Petioles

swollen, pulvinate, 2-7 mm long, ± terete,

684

glabrescent. Conflorescences ramiflorous or

just below the lowest/oldest leaves, composed

of 92-224 flowers; conflorescence axis 150—

380 mm long, angled (not terete) when dry,

densely clothed in dark brown, medifixed

hairs. Common bracts lanceolate, 0.8 mm

long, with mostly marginal hairs. Floral bracts

lanceolate to ovate, c. 0.7 mm long, glabrous

except for minute, dark brown marginal hairs,

inserted in lower half of pedicel (and usually

near base of pedicel). Pedicel terete, 2.9-3.5

mm long, clothed in dark brown medifixed

hairs, paired on a peduncle c. 1 mm long.

Tepals 20-26 mm long and clavate at apex

in mature bud, splitting proximally at first

with individual tepals coiling and reflexed at

anthesis, apex acute, pink, moderately clothed

on abaxial surface with hairs as for pedicels,

glabrous adaxially. Stamens with free

filament tips up to c. 0.1 mm long, inserted

c. 5 mm from tepal apex; anthers 3.1-3.9 mm

long (including a mucro of c. 0.4 mm long).

Hypogynous glands connate, the apex with

a fringe of globose papillae, fleshy. Style

17-21 mm long; pollen presenter c. 2 mm

long; ovary 2.6-3.1 mm long, ovules 12-20,

placenta marginal. Fruit leathery (neither

fleshy nor woody), 60-150 mm long, 21-43

mm diameter, asymmetrical and somewhat

ellipsoidal in shape with one side nearly flat,

smooth, green, style persistent. Seeds angular

to hemispherical or shortly cylindrical, often

many-faced with the abaxial face convex,

3-20 per fruit, 12-30 mm long on the longest

axis, testa thin, radicle 1-2 mm long, adaxial

surface of cotyledons flat. Germination

hypogeal. Sayers’s silky oak.

Additional selected specimens examined : Queensland.

Cook District: Cucania, Apr 1948, Stephens 12344

(CNS); Bellenden Ker bottom cable station. May 1996,

Gray 6725 (CNS); The Boulders, on walk to lookout,

7 km (by road) west of Babinda, Jun 1992, Conn 3635

(BRI, CNS, MEL, NSW); The Boulders, Babinda, Mar

1980, Jago (R.L.) 415 (CNS); Weinert’s Creek Babinda,

May 1978, Jago (R.L.) 89 (CNS); N.P.R. 226, Bellenden

Ker, junction of Windin Falls road and Bartle Frere road,

Apr 1995, Jensen 218 (CNS); Old Boonjie road. May

1981, Foreman 103 (CNS); S.F.R. 310, Windin L.A.,

Nov 1987, Hyland 25216RFK (CNS); loc. cit. , May 1982,

Gray 2551 (CNS); Topaz road, Apr 1974, Stocker 1158

(BRI, CNS); Towalla road, Towalla, May 1993, Cooper

534 & Cooper (CNS); S.F.R. 755 Boonjee L.A., Apr

1972, Hyland 5935 (BRI, CNS); S.F.R. 755, Gosschalk

L A, E.P./34, Nov 1976, Unwin 125 (CNS); S.F.R. 755,

Austrobaileya 8(4): 670-687 (2012)

Badgery L.A., Nov 1981, Hyland 11272 (CNS); Henrietta

Creek, Palmerston Highway, Jan 1993, Cooper 479 &

Cooper (CNS); T.R. 1244 Palmerston, Mar 1979, Gray

1316 (CNS); S.F.R. 756, Maalan L.A., Jan 1979, Dansie

20138 (CNS).

Distribution and habitat : Hollandaea

sayeriana is endemic to the Wet Tropics

bioregion in north-eastern Queensland,

where it is currently known to occur between

Mt Bellenden Ker (south of Cairns) and

the Innisfail area, including the eastern

edge of the Atherton Tableland (Map 1).

It inhabits very wet mesophyll to (rarely)

notophyll vine-forests/rainforests on soils

derived from basalt, granite and fine grained

metasediments (mudstones). Common canopy

species throughout its range include Alstonia

scholaris (L.) R.Br., Backhousia bancroftii

F.M.Bailey & F.Muell. ex F.M.Bailey,

Beilschmiedia bancroftii (F.M.Bailey)

C.T.White, Cardwellia sublimis F.Muell.,

Cryptocarya oblata F.M.Bailey, Elaeocarpus

ruminatus F.Muell., Endiandra bessaphila

B. Hyland, Ficus pleurocarpa F.Muell.,

Ficus variegata Blume, Franciscodendron

laurifolium (F.Muell.) B.Hyland & Steenis,

Myristica globosa subsp. muelleri (Warb.)

W.J.de Wilde and Syzygium gustavioides

(F.M.Bailey) B.Hyland. Common small trees

and shrubs throughout its range include

Apodytes brachystylis F.Muell., Ardisia

bre\ipedata F.Muell., Atractocarpus hirtus

(F.Muell.) Puttock, Brombya platynema

F.Muell., Gossia dallachiana (F.Muell. ex

Benth.) N.Snow & Guymer, Hypsophila

dielsiana Loes., Irvingbaileya australis

(C.T.White) R.A.Howard, Psychotria sp.

(Utchee Creek H. Flecker NQNC5313),

Rockinghamia angustifolia (Benth.) Airy

Shaw and Symplocos hayesii C.T.White &

W.D.Francis. Altitudinal range, from existing

specimens, is from near sea-level to 800 m.

Hollandaea sayeriana has been collected

or reliably reported in the following REs:

7.8.1a (occasionally), 7.8.2a (commonly),

7.8.12 (rarely), 7.11.1a (rarely), 7.11.12a

(occasionally), 7.11.28 (rarely), 7.11.29a

(rarely), 7.11.29b (rarely) and 7.12.1a

(commonly).

Ford & Weston, Revision of Hollandaea

685

Table 1. Morphological comparison of Hollandaea species (fruit and seed features mostly

taken from Cooper & Cooper [2004: 413-415])

Characters

H. diabolica

H. riparia

H. porphyrocarpa

H. sayeriana

Fruit surface

sculptured

smooth

Fruit colour

orange-green

green

purple

green

Fruit length

40-50 mm

25-50 mm

35-45 mm

60-150 mm

Seed colour

brown

cream/yellow/

green

cream/green

whitish

Seed length

22-25 mm

14-20 mm

13-16 mm

12-30 mm

Seeds per fruit

1-4

2-8

6-9

3-20

Leaf apex

retuse-

obtuse

obtuse-acute

acute-acuminate

acute-obtuse

to acuminate

Leaf base

attenuate

cuneate

Leaf length:width

2.6-3.6

4-8

2.5-2.8

1.2-2.2

Pedicel length

2.1-3 mm

1.9-2.2 mm

3-4.5 mm

2.9-3.5 mm

Conflorescence

axis length

48-120 mm

110-155 mm

52-103 mm

150-380 mm

Conflorescence

axis pubescence

moderately

hairy

glabrous

densely hairy

Tepal length

27-33 mm

28-32 mm

26-29 mm

20-26 mm

Tepal condition

(abaxial surface)

glabrous

clothed in

medifixed hairs

clothed in

medifixed hairs

Style length

20-24 mm

21-25 mm

16-20 mm

17-21 mm

Ovule number

2-4

6-8

12-16

12-20

Anther length

4-5 mm

4-6 mm

4.5-6 mm

2-3.9 mm

Filament length

0.4-0.8 mm

0.1-0.3 mm

c. 0.4 mm

0-0.1 mm

Hypogynous glands

free

connate

Phenology : Flowers have been recorded in

April, May, June and October; fruits have been

recorded in January, March and November.

Notes : The perianth is pink to purple-red

in bud and pink, crimson or purple-pink

at anthesis. New leafy growth is recorded

as being blue-black or maroon. The flower

aroma has been variously described as being:

‘sweet like honey’, ‘sweetly fragrant’ and also

as ‘strongly unpleasant’.

Affinities : Hollandaea sayeriana appears to

be most closely related to H. porphyrocarpa in

that they share a smooth fruit surface, abaxial

tepal surface clothed in medifixed hairs and

a relatively high ovule number. H. sayeriana

can be distinguished from H. porphyrocarpa

on the following features: leaf base (cuneate

versus attenuate, respectively), larger fruit

(60-150 mm versus 35-45 mm), larger seeds

([12—] 20-30 mm versus 13—16 mm), longer

conflorescence axis (150-380 mm versus

52-103 mm) and shorter anthers (2-3.1 mm

versus 4.5-6 mm). Comparisons between all

species of Hollandaea are provided in Table

1 .

Conservation status : Most existing

collections have been made in Wooroonooran

National Park which is within the World

Heritage Area of the Wet Tropics bioregion.

Currently, Hollandaea sayeriana is listed

under the NCA as Near Threatened. It is

estimated to have an extent of occurrence of

1100 km 2 and an area of occupancy of620 km 2 .

We recommend H. sayeriana being retained

as Near Threatened as it fulfills the criteria

under categories Near Threatened A, E and

Austrobaileya 8(4): 670-687 (2012)

possibly C. We feel it would be premature to

suggest a “Near Threatened” status under the

IUCN guidelines, even though it may well be,

as it is unlikely that it would qualify for any

higher status in the future given that there is

no evidence to support either the necessary

population sizes or number of populations or

any decline under any of the other necessary

criteria.

Etymology : The species is named for W.A.

Sayer, a 19th century Australian naturalist,

botanical collector and collector of the type

specimen.

Excluded names

Hollandaea lamingtoniana F.M. Bailey,

Queensland Agric. J. 5:390 (1899) = Helicia

lamingtoniana (F.M.Bailey) C.T.White ex

L.S.Sm.

Acknowledgements

The authors wish to thank Will Smith for

the illustrations. Peter Bostock provided the

distribution maps. Wendy Cooper and Rupert

Russell are warmly thanked for their patience

on the arduous walk to “The Pinnacle” to

show the first author the newly described

plants for the first time. Rupert Russell also

provided additional useful information on

the distribution of both new species. Garry

Sankowsky kindly escorted both authors on

different occasions to Roaring Meg Creek

to see and collect Hollandaea riparia. Lyn

Craven provided the necessary nomenclatural

rules to allow the authors to refer to the locality

“Devils Thumb” in a legitimate and sensible

manner. Brendan Lepschi and Kirsten

Cowley brought nomenclatural and literature

changes in APNI (Australian Plant Name

Index) for Hollandaea to our attention. Kevin

Thiele and Bob Makinson made numerous

improvements to an earlier draft. Permits

to collect in Daintree National Park were

issued by the Department of Environment

and Resource Management. Staff at CNS,

especially Frank Zich, are warmly thanked

for allowing access to specimens and the use

of their facilities.

References

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Victorian Naturalist 3: 169-170.

Australian Plant Name Index (2011). [accessed 23

February 2011 and 1 April 2012], http://www.

anbg.gov.au/cgi-bin/apni

Bailey, F.M. (1899). Botany: Contributions to the flora of

Queensland. Queensland Agricultural Journal

5: 390.

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H.J.Diddams & Co.: Brisbane.

Barker N.P, Weston, P.H., Rutschmann, F. & Sauquet,

H. (2007). Molecular dating of the ‘Gondwanan’

plant family Proteaceae is only partially

congruent with the timing of the break-up of

Gondwana. Journal of Biogeography 34: 2012-

2027.

Carpenter, R.J. (1994). Cuticular morphology and

aspects of the ecology and fossil history of north

Queensland rainforest Proteaceae. Botanical

Journal of the Linnean Society 116: 249-303.

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

Australian Tropical Rainforest. Nokomis

Editions: Melbourne.

Engler, A. (1888). Die Natiirlichen Pflanzenfamilien,

Teil III, vol. 2, fascicle 29. Wilhelm Engelmann:

Leipzig

Environment Australia (2005). Revision of the Interim

Biogeographic Regionalisation for Australia

(IBRA) and Development of Version 5.1 -

Summary report (2000). Updated, IBRA

Version 6.1 (Digital Data, metadata), [accessed

24 June 2009], http://www.environment.gov.au/

parks/nrs/science/bioregion-framework/ibra/.

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Australia: Melbourne.

Forster, PI. & Edginton, M. (2007). Proteaceae. In

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

of the Queensland Flora 2007 , pp. 169-172.

Queensland Herbarium, Environmental

Protection Agency: Brisbane.

- (2010). Proteaceae. In P.D.Bostock &

A.E.Holland (eds.). Census of the Queensland

Flora 2010, pp. 163-166. Queensland

Herbarium, Department of Environment &

Resource Management: Brisbane.

Hoot, S.B. & Douglas, A.W. (1998). Phylogeny of the

Proteaceae based on atpB and atpB-rbcL

intergenic spacer region sequences. Australian

Systematic Botany 11: 301-320.

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(ed.). Flora of Australian 16: 391-393. CSIRO

Australia: Melbourne.

Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray,

B. & Elick, R.W. (2003). Australian Tropical

Rain Forest Plants. Trees, Shrubs and Vines.

CD-ROM. CSIRO Publishing: Melbourne.

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IUCN Species Survival Commission. IUCN:

Gland, Switzerland.

Johnson, L.A.S. & Briggs, B.G. (1963). Evolution in the

Proteaceae. Australian Journal of Botany 11:

21-61.

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classification of a southern family. Botanical

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Mueller, F. (1886). Descriptions of new Australian

plants. Victorian Naturalist 3: 92-93.

- (1887). Notes on Australian plants. Australasian

Chemist and Druggist 2: 173.

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Rainforest Plants IV. Terania Rainforest

Publishing: The Channon.

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Rainforest Publishing: The Channon.

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Barker, N.P, Cantrill, D.J., Mast, A.R. &

Savolainen, V. (2009). Contrasted patterns of

hyperdiversification in Mediterranean hotspots.

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Sciences of the U.S.A. 106: 221-225.

Smith, L.S. (1952). Opisthiolepis , a new genus of

Proteaceae from Queensland. Proceedings of

the Royal Society of Queensland 62: 79-81.

- (1956). New species of and notes on Queensland

plants. Proceedings of the Royal Society of

Queensland 67: 29-40.

Venkata Rao, C. (1971). Proteaceae. Council of

Scientific & Industrial Research: New Delhi.

Weston, PH. (2006). Proteaceae. In K.Kubitzki (ed.),

Families and Genera of Vascular Plants 9:

364-404. Springer Verlag: Berlin.

Weston, PH. & Barker, N.P. (2006). A new

suprageneric classification of the Proteaceae,

with an annotated checklist of genera. Telopea

11: 314-344.

Williams, K.A. (1984). Native Plants of Queensland ,

Vol. 1. K.A.W.Williams: Ipswich.

Wrigley, J.W. & Fagg , M. (1991). Banksias, Waratahs &

Grevilleas, and all other plants in the Australian

Proteaceae family. Collins Angus & Robertson

Publishers Pty Limited: North Ryde.

Nomenclature and synonymy of several Goodenia R.Br.

(Goodeniaceae) species from northern Australia

Ailsa E. Holland

Summary

Holland, A.E. (2012). Nomenclature and synonymy of several Goodenia R.Br. (Goodeniaceae)

species from northern Australia. Austrobaileya 8 ( 4 ): 688 - 695 . The nomenclature, characters and

morphological variation for several species of Goodenia from the Northern Territory and Queensland

are investigated, and the names G. paludicola Carolin and G. heteroptera (F.Muell.) B.D. Jacks, are

newly recorded as synonyms of G. minutiflora F.Muell. and G. pilosa (R.Br.) Carolin respectively.

New amended descriptions are provided, along with notes on identification and conservation.

Key Words: Goodeniaceae, Goodenia, Goodenia heteroptera, Goodenia minutiflora, Goodenia

paludicola, Goodenia pilosa, threatened species, Queensland flora, new synonym

Ailsa E. Holland, Queensland Herbarium, Department of Science, Information Technology,

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

Australia. E-mail: Ailsa.Holland@science.dsitia.qld.gov.au

Introduction

Goodenia paludicola Carolin and G.

heteroptera (F.Muell.) B.D.Jacks. are

currently listed as Near Threatened under

Queensland’s Nature Conservation Act 1992.

These two species have been difficult to

distinguish from G. minutiflora F.Muell. and

G. pilosa (R.Br.) Carolin respectively, based

on the available literature. These species all

occur in the seasonally wet areas of northern

Queensland (Qld), and in the northern part

of the Northern Territory (NT). Ephemerals

in these areas plus adjacent parts of Western

Australia (WA), have previously been poorly

collected due to inaccessibility during the

wet season. The taxonomic and conservation

status of these species is here re-examined

in light of the many additional herbarium

collections now available.

Materials and methods

Diagnostic characters from the literature

were examined (Carolin 1990, 1992) and

are discussed in light of observations made

on the type material, additional specimen

collections, and field observations. Type

specimens from SYD and MEL, along with

the full set of specimens now available at

BRI were examined: 22 specimens of G.

Accepted for publication 31 August 2012

minutiflora/G. paludicola (including one from

the NT); and more than 150 specimens of G.

pilosa/G. heteroptera (including 12 from the

NT). Measurements were made by steel ruler

or micrometer. Field observations were made

in several areas of north Qld.

1. Goodenia paludicola Carolin and G.

minutiflora F.Muell.

The type specimen of Goodenia paludicola

was designated by Carolin (1990) as

“holotype: Queensland: 14 miles (22.4 km)

NW of Corinda on road to Westmoreland,

R.C.Carolin 9147, 7 May 1974 (NSW).

Isotype: SYD”. This collection was not

found in NSW (see acknowledgements). The

isotype in SYD is not designated as such,

but named “ Goodenia udicola Carolin” on

the label (7.5.1974) and annotated “ Goodenia

paludicola correct name Jan 1989”. As the

only type material found, this specimen is

here designated as lectotype for the name

G. paludicola. Another cited specimen

(<Carolin 8766 [SYD]), was also annotated

as “ Goodenia udicola Carolin” (10.12.1974)

and further annotated as “isotype” possibly

at a later time, but as this name was never

published it can be ignored.

The remaining three specimens cited

by Carolin in the protologue of Goodenia

paludicola (Carolin 1990) are all from the

Holland, Goodenia nomenclature and synonymy

NT. These have been re-determined as G.

purpurascens R.Br. based on the much longer

corollas, and are therefore excluded from this

study (see acknowledgements).

The lectotype of Goodenia minntiflora,

designated by Carolin (1990), was also

examined “Lectotype: Queensland: Between

the Norman and Gilbert rivers, T. Gulliver

68 (MEL 23997). Isolectotypes: MEL 23998,

22243*” the latter an additional collection by

Gulliver from the Norman River. Bentham

(1868) fails to mention G. minutiflora,

although he describes “var. ? minutaF MuqW”

under G. purpurascens , a later synonym (see

taxonomy below).

Discussion of characters

Mueller (1874) described Goodenia

minutiflora as a small herb [8-15 mm] with

cauline leaves rarely opposite and then little

different to bracteoles, the calyx lobes semi-

linear, short, corolla minute [ c. 3.2 mm], the

lobes equal and not winged, seeds many,

minute. Carolin (1990, 1992) distinguished

his new species G. paludicola from G.

minutiflora by the larger corolla that is deeper

in colour, and the auricles on the superior

lobes enclosing the indusium.

Peduncles. Carolin (1990,1992) described the

peduncles of G. paludicola as “to 3 mm long”.

The peduncle is normally described as “the

part of the stalk below the bracteoles” Carolin

(1992: 10). However, in this case it appears

that the distance measured is that between the

two uppermost bracts (bracteoles) which are

subopposite on the stem. The inflorescences

(both species) are complex thyrse-like

panicles with indeterminate growth, the

bracts subtending each new branch initially

opposite and then increasingly smaller and

offset by up to 3 mm towards the plant apex.

The peduncles in this case are correctly

interpreted as the distance between the

uppermost bracts and the point of branching

of the inflorescence directly below. This

measurement was observed to be 4-20 mm

on the type material of G. paludicola.

The peduncles of the type material of

Goodenia minutiflora and the other specimens

examined of this species, were also observed

689

to be 4-20 mm (described as 4-10 mm in

Carolin 1992).

Pedicels. The pedicel length for Goodenia

paludicola was described as 7-9 mm by

Carolin (1990, 1992). This does not appear to

be correct even for the type material which

was observed to have pedicels 3-15 mm long.

The pedicels of the type material of

Goodenia minutiflora were observed to be

5-16 mm long (described as 5-7 mm by

Carolin 1992). The other specimens examined

were observed to have pedicel lengths 3-29

mm.

Sepal length. The sepal length for Goodenia

paludicola is described as 1.5-1.8 mm

(Carolin 1990, 1992) but the sepal length

observed for the type material was less than 1

mm. The length of sepals for the type material

of G. minutiflora , along with the majority of

specimens examined, was also less than 1

mm, although a number of specimens had

slightly longer sepals, up to 1.8 mm long.

Corolla length. Corolla length was used by

Carolin (1990, 1992) to distinguish Goodenia

paludicola (4-5 mm) from G. minutiflora

(2-3 mm). However, the type material of G.

paludicola was observed to have corollas 2-3

mm long. The type material of G. minutiflora

was also observed to have corollas 2-3 mm

long, and the corollas of the other specimens

examined were observed to be 2-6 mm long.

Therefore the species cannot be distinguished

by corolla length.

Corolla colour. The deeper corolla colour

was one of the characters used to distinguish

Goodenia paludicola from G. minutiflora.

Carolin (1990, 1992) describes the corolla

colour of G. paludicola as bluish-purple,

and on the label of the type material it is

described as pale purple. Similar specimens

collected from the same area as the type of

G. paludicola had corolla colour described on

the labels as purple-red (Forster PIF20926 &

Booth), white to pale mauve (Pullen 9123), or

purple (Thompson WES724 & Hogan).

The colour of the corolla of G. minutiflora

was described as purple by Mueller (1874) and

“white suffused purple or purplish outside”

690

by Carolin (1992), but corolla colour is not

mentioned on the labels of this type material.

The corolla colour of additional specimens

examined were variously described as white,

mauve or red-purple. It is apparent therefore

that corolla colour is not a reliable character

for distinguishing these two proposed species.

Corolla lobe auricles. Goodenia paludicola

was distinguished from G. minutiflora by

“the auricles on the superior lobes enclose the

indusium” (Carolin 1990, 1992). The indusia

of Goodenia species are often enclosed or

“sheltered” below the auricles in the early

stages of flowering (personal observation) and

this is variable on the specimens examined.

The “enclosing” of indusia is possibly

dependant on the age of the flower or pressing

process.

Conclusion

Based on the above observations, the type

material of Goodenia paludicola appears

to represent a small part of the variation

apparent across the range of G. minutiflora

and is therefore here treated as a synonym of

that species. The description of G. minutiflora

is here amended from Carolin (1979, 1990,

1992) to accommodate the additional

observed variation.

Taxonomy

Goodenia minutiflora F.Muell., Fragm.

8: 244 (1874) Type: [Queensland. Cook

District:] Between the Norman and Gilbert

Rivers, 1874, TGulliver 68 (lecto: MEL,

isolecto: MEL,fide Carolin [1990]).

Goodeniapurpurascens var. minima F.Muell.

ex Benth., FI. Austral. 4: 78 (1868); G. minima

(Benth.) Domin, Biblioth. Bot. 22: 643 (1929).

Type: Northern Territory. Upper Victoria R.

[River], s.dat., F.Mueller s.n. (holo: K n.v:,

iso: MEL).

Goodenia paludicola Carolin, Telopea 3:

536 (1990), syn. nov. Type: Queensland.

Burke District: 14 miles [c. 22 km] NW of

Corinda on road to Westmoreland, 7 May

1974, R.C.Carolin 9147 (lecto: SYD, here

designated).

Slender annual herb ascending to 25 cm tall.

Austrobaileya 8(4): 668-695 (2012)

Stems and pedicels with sparse patent hairs

to 0.7 mm long, mixed with shorter glandular

hairs. Leaves linear to oblanceolate, entire

or with a few teeth, 1-11 cm long, 1-10 mm

wide, acute at apex, tapered at base, glabrous

or nearly so. Inflorescences comprising

compound, leafy thyrse-like panicles, each

branch subtended by a reduced leaf-like bract,

the bracts sub-opposite below, becoming

smaller and more distant, separated by up to

3 mm towards the apex. Bracts mostly linear,

1-35 mm long; peduncles 4-20 mm long;

pedicels 3-28 mm long, articulate c. 1 mm

below the ovary. Sepals partly adnate to the

ovary, lanceolate to ovate, acute, 0.5-1.8 mm

long, pubescent with simple and glandular

hairs. Corolla 2-6 mm long, white, mauve,

purple or red-purple; outer surface with simple

patent hairs and shorter glandular hairs; inner

surface glabrous, without enations; anterior

pouch obscure. Corolla lobes with wings c.

0.5 mm long and wide; abaxial lobes 1-1.5

mm long; adaxial lobes 1.5-2 mm, the auricle

+ wing c. 1 mm long. Stamen filaments 1-1.5

mm long; anthers c. 0.3 mm long. Ovary

glandular pubescent; septum nearly as long

as loculus; ovules numerous; style to 3 mm

long, with scattered patent hairs in the upper

part; indusium somewhat oblong, widening

at the top, 0.3-0.5 mm long, glabrous below,

orifice bristles longer on upper lip, very

short on lower lip. Fruit ovoid, 2-4 mm long,

covered with patent simple hairs, sometimes

mixed with shorter glandular hairs; 2-valved,

the valves entire. Seeds slightly biconvex,

elliptic, 0.3-0.4 mm long, smooth, glossy,

with a narrow rim to 0.1 mm.

Additional selected specimens examined : Northern

Territory. 17.9 km from Borroloola T-junction, towards

Wollogorang, Jun 1999, Bean 15116 (BRI, DNA,

MEL). Queensland. Burke District: Upper Cliffdale

Creek area, c. 23 km NW of old “Corinda” Outstation

on the Doomadgee - “Westmoreland” Road, May

1974, Pullen 9123 (AD, BRI, CANB, HO, MEL); 49

km from Doomadgee on Hells Gate Road, May 1997,

Forster P1F20926 & Booth (BRI); Bowthorn Station,

31.5 km NNE of Bowthorn homestead, Jun 2006,

Thompson WES724 & Hogan (BRI); 4 miles [6.4 km] N

of Maggieville on Myravale Road, May 1974, Carolin

8766 (SYD); Melville Creek, on road N of Normanton,

May 2004, McDonald KRM2306 & Covacevich (BRI);

10 km along the Normanton to Mogoura Road, SW

of Normanton, Apr 1974, Pullen 8851 (BRI, CANB);

31.1 km by road W of Croydon, Apr 2009, McDonald

Holland, Goodenia nomenclature and synonymy

KRM8388 (BRI); 27.1 km along Claraville Station Road

from Croydon, Apr 2007, McDonald KRM6490 (BRI).

Cook District: 140.7 km along Chillagoe Road from

junction with Normanton - Karumba Road, May 2004,

McDonaldKRM2325 & Covacevich (BRI); 13 km along

Forsayth Road from Georgetown, Mar 2008, McDonald

KRM7471 (BRI, NSW).

Distribution and habitat : Goodenia

minutiflora is distributed across northwestern

Qld from Croydon to the Borroloola area in

the NT. Plants occurs on the edges of lagoons,

creeks, and wet depressions, drainage lines

or soaks and on clay or sand plains, often

associated with Melaleuca or Eucalyptus and

Corymbia dominated open woodlands.

Affinities : Goodenia species that are

morphologically similar and with pink,

purple or white flowers are: G. purpurascens

R.Br. (NT, WA & Qld), G. viscidula Carolin

(NT & Qld) and G. gloeophylla Carolin (NT

& WA). However, G. gloeophylla and G.

viscidula are both viscid glandular plants and

G. purpurascens has longer corollas 8-12 mm

long (Carolin 1992).

Phenology : The species is an ephemeral herb

appearing between March and June after rain.

Conservation status : A widespread species

(currently Least Concern), potentially

threatened in some areas by weed/pasture

invasion of habitat.

2. Goodenia heteroptera (F.Muell.)

B.D.Jacks. and Goodenia pilosa (R.Br.)

Carolin

Goodenia heteroptera has been known only

from the holotype collected in the Newcastle

Range in northern Qld ( W.E.Armit 377 [MEL

68192]) (Carolin 1979; 1992). The more

widespread and variable G. pilosa , originally

described from an island in the Gulf of

Carpentaria, has been considered to occur

widely across northern Qld and the northern

part of the NT, as well as New Guinea,

Indonesia, China and the Philippines (Carolin

1979, 1992; AVH 2012).

Discussion of characters

Mueller (1876) distinguished Goodenia

heteroptera by the pedicels several times

shorter than the calyx, and the very small

yellow corolla with short superior lobes with

691

unequal wings, a trifid style and wingless

seeds (Mueller did not mention G. pilosa).

Brown (1810) described G. pilosa as a

herbaceous pilose annual, the leaves dentate

or incised, with basal auriculate flowers on

axillary, ebracteate one flowered peduncles,

the fruit reflexed. Bentham (1868) also

described G. pilosa but failed to mention G.

heteroptera.

Carolin (1979) distinguished Calogyne

heteroptera from C. pilosa by the presence

of glandular hairs. He also described three

races of C. pilosa (Table 1). Carolin (1992)

further distinguished Goodenia heteroptera

by the length of the corolla ( c. 6 mm long), the

presence of glandular hairs on the corolla, the

sinus between the auricle and the wing on the

adaxial corolla lobes, and the wing above the

auricle narrower than the opposite one. Both

descriptions of this species are based entirely

on the holotype.

Habit. The holotype of Goodenia heteroptera

consists of a single small rosette. This

material represents an early rosette stage

of development only. Goodenia species

occurring in ephemeral wetlands in northern

Australia commonly complete their life cycle

quickly by flowering at an early stage, and then

if suitable conditions persist, further growth

and inflorescence development occurs.

The type material of Goodenia pilosa

includes several plants with considerable stem

development; however, all stages from rosette

to trailing stems have been observed among

the specimens examined. Carolin (1979,1992)

also described a variety of habits for G. pilosa

in his races (Table 1).

Hair type. The presence of glandular hairs

was one of the characters distinguishing

G. heteroptera (Carolin (1979, 1992) and

these are present on the holotype. However,

plants with or without glandular hairs were

observed to be otherwise indistinguishable

on the specimens examined. It is probable

that these glandular hairs are more common

on specimens collected at moist sites (e.g.

McDonald KRM321 , Clarkson 8979) and may

even be lost as the environment dries out.

Carolin (1992) also described this situation for

692

Austrobaileya 8 ( 4 ): 668-695 ( 2012 )

Table 1. Forms of Goodenia pilosa as described by Carolin (1990,1992)

Race A = form i

Race B = form ii

Race C = form iii

distribution

Arnhem Land, NT

north Qld

habit

sprawling

compact

sprawling

leaf

indumentum

conspicuously hirsute

almost glabrous

few hairs

pedicels

well developed

short, to 4 mm

usually more than 4 mm

corolla hair

outside

hirsute-pubescent

few simple hairs

hirsute-pubescent

seed surface

glossy, smooth

prominently

verrucose

verrucose to almost

smooth in centre

G. purpurascens, where the plants growing in

creek beds are more glandular hairy.

Hair density. The holotype of Goodenia

heteroptera has leaves with sparse hairs,

described by Carolin (1979) as “scattered’

while the type material (MEL 68195) of G.

pilosa is quite hairy. However, hair density

was observed to be quite variable across the

range on the specimens examined, with those

from the NT usually more densely hairy,

(along with the type material of G. pilosa)

although Carolin’s race “B” from Arnhem

Land was described as “almost glabrous”.

Leaf length and width. The holotype of

Goodenia heteroptera has only small rosette

leaves, commonly found on immature

material across the range. Most mature

specimens from north-west Qld and the NT

were observed to have longer, more narrow

leaves similar to the type material of G. pilosa ;

however, leaf shape varied continuously from

north-east Qld to the NT.

Pedicel length. The very short pedicel length

was one of the characters used by Mueller

(1876) to distinguish Goodenia heteroptera

and the holotype was observed to have short

pedicels c. 1 mm long. Carolin (1979, 1992)

described the pedicels of G. heteroptera as

“scarcely 5 mm long” and those of G. pilosa as

“2-2.5 mm” (although race “C” is described

with pedicels more than 4 mm long). The

pedicel length observed on the specimens

examined varied continuously from 1-22 mm

long, suggesting that the measurement given

by Carolin is an error and should read “2-25

mm”.

Corolla length. Corolla length was one of

the characters used by Carolin (1979, 1992)

to distinguish G. heteroptera ( c . 6 mm long)

from G. pilosa (8-15 mm long). The holotype

of G. heteroptera has one small flower in

the packet, the corolla measuring 6.5 mm

long. Corolla length was observed to vary

from 6 to 13 mm for the other specimens

examined, with corollas tending to be longer

on specimens from the NT (10-13 mm).

Wing above the auricle. The unequal wings

on the superior (adaxial) lobes was mentioned

by Mueller (1974), Bentham (1868) and

Carolin (1979, 1992) as a distinguishing

character for Goodenia heteroptera. Carolin

also described the presence of a sinus

between wing and auricle, as an additional

distinguishing character. On all specimens

examined, the wing above the auricle is

shorter, and the wing-auricle boundary is

not clear cut. It was observed to be either

seamless, folded or broken on various

specimens. The appearance of a sinus on

the holotype of G. heteroptera is therefore

probably a misinterpretation of folding in the

transition zone where it joins to the more rigid

auricle. In the field, the auricles of G. pilosa

together form an “umbrella” over the indusia

during the early stages of flowering, and a fold

or sinus probably develops as the flower ages,

or during pressing.

Holland, Goodenia nomenclature and synonymy

Seed surface. Carolin (1990, 1992) describes

Goodenia heteroptera as having a verrucose-

granular seed surface and G. pilosa as

having both smooth (race A) or verrucose

surfaces (races B and C). The holotype of G.

heteroptera , along with most of the eastern

Qld material has verrucose seeds, sometimes

very faintly so, or verrucose with an almost

smooth centre, representing transitional states

(e.g. Wannan 3534, Clarkson 8979, Bean

16419). The type material of G. pilosa has

smooth seeds.

Conclusion

I conclude that the holotype of Goodenia

heteroptera, though flowering and fruiting,

represents an early rosette stage of

development, and represents just one part of

the variation of the variable and widespread G.

pilosa. The type specimen corresponds most

closely with Carolin’s Race B from Arnhem

Land (Carolin 1979), but the variability across

the range does not support formal recognition

of these races. Specimens examined from

north-west Qld and the NT usually have longer,

more narrow leaves, denser indumentum and

longer corollas (Race A), but these characters

vary across the range of specimens examined.

The name G. heteroptera is therefore treated

as a synonym of G. pilosa and the description

of G. pilosa is amended from Carolin (1979,

1992) to accommodate the corrections and

additional observed variation.

Taxonomy

Goodenia pilosa (R.Br.) Carolin, Telopea

3: 365 (1990); Calogyne pilosa R.Br., Prodr.

579 (1810); Goodenia dubia Sprengel, Syst.

Veg. 1: 271 (1824), nom. illeg. Type: Northern

Territory. Carpentaria, island no. 12 [North

Is., Sir Edward Pellew Group,], 16 December

1802 ,R.Browns.n. (lecto: BMn.v., isolecto: K

[photo!], MEL,fide Carolin [1979: 5]).

Balingayum decumbens Blanco, FI. Filip. 187

(1837). Type: not designated.

Calogyne chinensis Benth., J. Linn. Soc.,

Bot. 5: 78 (1861). Type: China, near Amoy

[Xiamen], s,dat., H.F.Hance 1422 (holo: K,

photo!, fide Carolin [1992: 275]).

693

Calogyne heteroptera F.Muell., Fragm. 10:

43 (1876); Goodenia heteroptera (F.Muell.)

B.D.Jacks., Index Kew. 1: 1056 (1895), syn.

nov.; Carolin, Telopea 3: 565 (1990), nom.

superfl. Type: Queensland. Cook District:

Newcastle Ra. [Range], 5. dat., W.E.Armit 377

(holo: MEL).

Prostrate annual, initially a rosette to 5

cm high, later seasonal growth spreading

with prostrate stems to 50 cm long. Plants

sparsely to moderately pilose with translucent

hairs to 1.5 mm long, spreading or oblique,

sometimes mixed with smaller glandular hairs

particularly on lower leaves, stems and sepals.

Leaves sessile, variable in shape, obovate to

lanceolate or linear, acute, broadly acute or

rounded at apex; margins entire or shallowly

and distantly dentate, sparsely hirsute mainly

on the margins and midvein. Basal leaves

obovate, tapered at base, 1—4(—8) cm long,

0.3-1 cm wide; cauline leaves alternating

on stem, oblanceolate to nearly linear, to 15

cm long, and up to 1 cm wide, base tapered

or auriculate with several spreading lobes.

Racemes to 35 cm long, the flowers subtended

by leaf-like bracts, and often clustered near

the apex; pedicels 1-22 mm long, sparsely

pilose, not articulate, bracteoles absent.

Sepals attached just below the top of the

ovary, lanceolate to narrowly oblong, 2.2-5

mm long, ciliate. Corolla 7-15 mm long, pale

yellow with brown or red stripes; outer surface

sparsely pilose; inner surface hairy in throat,

without enations or pouch; abaxial corolla

lobes 3-5 mm long, wings 1-1.5 mm wide,

extending 1/2 to 2/3 of the lobe; adaxial lobes

3-7 mm long, the auricle well developed, c.

1.5 mm long and wide, and joined to wing on

that side, sometimes with a fold or shallow

sinus between; wings ± equal, 1-1.5 mm long

and wide. Ovary pilose, sometimes shorter

glandular hairs also present; septum short;

ovules 10-15; style 3-fid, the middle indusium

shorter, semicircular, c. 0.5 mm long. Fruit

subglobular, 2.5-5 mm diameter, valves

persistent, gaping. Seed pale brown, ovate to

elliptic with a prominent rim, 2-4.5 mm long,

smooth to verrucose, wing absent.

694

Additional selected specimens examined : Northern

Territory. English Company Islands, Cotton Is., Jul

1992, Leach 3102 (BRI, DNA, K); E side Mt Norris

Bay, Oct 1992, Cowie 3246 (BRI, DNA, MEL); Kakadu

N.P., Jaribu East near Aerodrome, May 1988, Weber

9680 (AD, BRI); Yirrkala, Aug 1948, Specht 913 (BRI,

DNA); Echo Is., Ritjirriur Swamp, Jul 1975, Late 6124

(BRI, CANB, DNA, L, NSW). Queensland. Burke

District: Eight Mile Yard W of Doomadgee Aboriginal

Mission Station near Nicholson River, May 1974,

Pullen 9071 (BRI, CANB); 9.1 km (by road) ENE of

Musselbrook mining camp on Musselbrook Creek, 175

km N of Camooweal, Lawn Hill N.P., Apr 1995, Johnson

MRS526 & Thomas (BRI); Clara River, 40.3 km along

Richmond Road from Prospect Station turnoff. Mar

2008, McDonald KRM7584A (BRI). Cook District:

Thursday Island near satellite receiving dishes on

Milman Hill, Apr 1986, Clarkson 6456 (BRI, NSW);

58.7 km N of the Archer River crossing on the Coen to

Weipa Road, Apr 1991, Clarkson 8979 & Neldner (BRI,

NSW); 3 km E of junction of Kennedy River & Lakes

Creek, Lakefield N.P., May 1992, Neldner 4012 (BRI);

Isabella Falls 23.5 km E of Normanby River on Laura to

Cooktown Road, Jun 1985, Clarkson 5968 (BRI, CANB,

DNA, L, PERTH); Hann Tableland N.P., headwaters of

Gap Creek, central part of Tableland, W of Mareeba,

May 2010, Forster PIF37250 & Thomas (BRI, MEL);

150 metres from Rookwood Creek, Rookwood Station,

Mar 2000, McDonald KRM321 (BRI); 51.3 km along

O’Briens Creek Road from Mount Surprise, Mar 2007,

McDonald KRM6348 (BRI); Fog Creek Station, c. 1.2

km W of Fog Creek, about 21 km N of the homestead,

180 km N of Richmond, Apr 2004, Kahler TH7910

& Appelman (BRI). North Kennedy District: S of

Cardwell, May 2004, Wannan 3534 (BRI); Doug Haigh

Drive, NE of Ravenswood, Apr 2010, Holland 2068 &

Lovatt (BRI, DNA); Lannercost S.F., Oak Hills Road,

W of Ingham, May 2003, Ford AF3954 (BRI, MEL,

NSW); Bruce Highway, 13 km W of Bowen, May 2000,

Bean 16477 (BRI, MEL); 2.9 km along Airport Road,

S of Proserpine, May 2000, Bean 16419 (BRI). South

Kennedy District: 24.3 km from Proserpine towards

Mackay, Apr 1991, May 2000, Bean 2902 (BRI); Ilbilbie

- Notch Point Road, c. 4 km due W of coast, Apr 1994,

Champion 1056 & Pollock (BRI).

Distribution and habitat: Goodenia pilosa

is distributed across northern Australia, from

Mackay to Cape York in Qld, and across the

top end of the NT (AVH 2012); also in New

Guinea, Indonesia, China and the Philippines.

It occurs in seasonally wet areas, on the edges

of lagoons, creeks, and wet depressions,

drainage lines or soaks, on clay or sand

plains, often associated with Melaleuca or

Eucalyptus and Corymbia open woodland.

Affinities: Goodenia species that are similar

to G. pilosa and that have yellow flowers and

branched styles are G. holtzeana (Specht)

Austrobaileya 8 ( 4 ): 668-695 ( 2012 )

Carolin (NT & WA), G. heppleana (W.Fitzg.)

Carolin (NT & WA) and G. neglecta (Carolin)

Carolin (NT). However, G. holtzeana is an

erect, viscid herb with longer corollas (14-20

mm long); G. heppleana is also glandular

but is distinguished by the fruit which do not

gape open at maturity. Goodenia neglecta

is the most similar to G. pilosa , but can be

distinguished by the glandular hairy bracts

and pedicels (Carolin 1992).

Phenology: Goodenia pilosa is an ephemeral

herb appearing between March and June after

rain.

Conservation status: A widespread and

common species (currently Least Concern),

potentially threatened in some areas by weed/

pasture invasion.

Acknowledgements

Many thanks go to Keith McDonald (formerly

of the Department of Environment & Heritage

Protection) whose untiring collection efforts

have made this investigation possible. I

also wish to thank Murray Henwood (SYD)

and Dale Dixon (NSW) for searching for

type material of G. paludicola, ; Pina Milne

(MEL) for the loan of type material of G.

heteroptera , G. pilosa and G. minutiflora; and

Ian Cowie (DNA) and Dave Albrecht (NT)

for finding and measuring the three Northern

Territory specimens cited by Carolin (now re¬

determined as G. purpurascens).

References

AVH (2012). Australia’s Virtual Herbarium. Council

of Heads of Australasian Herbaria. http://www.

chah,gov.au/avh . Accessed 30 May 2012.

Bentham, G. (1868). Goodenia and Calogyne. In Flora

Australiensis 4: 50-81. L.Reeve & Co.: London.

Brown, R. (1810). Calogyne pilosa. Prodromus Florae

Novae Hollandiae et Insulae Van-Diemen 579.

J. Johnson: London.

Carolin, R.C. (1979). The genus Calogyne R.Br. in

Australia. Brunonia 2: 1-17.

- (1990). Nomenclatural notes and new taxa in

the genus Goodenia (Goodeniaceae). Telopea 3:

517-570.

- (1992). Goodenia. In A.S.George (ed.). Flora of

Australia 35: 147-281. Australian Government

Publishing Service: Canberra.

695

Holland, Goodenia nomenclature and synonymy

Mueller, F. (1874). Goodenia minutiflora. Fragmenta

Phytographiae Australiae 8: 244. Government

Printer: Melbourne.

- (1876). Calogyne heteroptera. Fragmenta

Phytographiae Australiae 10: 43. Government

Printer: Melbourne.

Austrobaileya 8(4): 696-698 (2012)

SHORT COMMUNICATION

Types of enigmatic north-Queensland

Orchids from the Dockrill herbarium

Ashley R. Field 1,2 & Frank A. Zich 1

'Australian Tropical Herbarium, Sir Robert Norman Building, James Cook University Cairns

Campus, PO Box 6811, Cairns, Queensland 4870 Australia.

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

Brisbane Botanic Gardens, Toowong, Queensland 4066 Australia. E-mail: Ashley.Field@science.

dsitia.qld.gov.au

Alick William Dockrill (1915-2011), author

of the classic work Australian Indigenous

Orchids (Dockrill 1969, 1992) donated his

personal orchid herbarium of approximately

1600 specimens to the Australian National

Herbarium - Atherton (QRS) which is now

integrated with the Australian Tropical

Herbarium - Cairns (CNS). A number of

duplicates of types have been recovered

from this material including those for five

poorly known taxa: Dendrobium masonii

Rupp, Dendrobium baseyanum St.Cloud,

Dendrobium x faederatum St.Cloud, Eria

intermedia Dockrill and Oberonia attenuata

Dockrill. These five taxa are enigmatic

because they have not been re-recorded in the

wild at or near their type localities since they

were described and also because two of them

are considered to be extinct in the wild under

State and Commonwealth legislation.

We review the status of these five type

specimens and provide a current interpretation

as to their correct taxonomic placement. All

specimens have been seen unless indicated

as n.v. Accepted names are given in bold

type. Type specimens originally accessioned

at QRS are cited with their QRS numbers,

whereas the recent duplicates are accessioned

with CNS numbers.

1. Dendrobium masonii Rupp, Austral.

Orchid Rev. 18: 18 (1953); Diplocaulobium

masonii (Rupp) Dockrill, Orchadian 1(11)

(1965). Type citation: “Cape Tribulation,

North Queensland, 1950, W.W. Mason,

Accepted for publication 20 August 2012

Junr.; flowering in Sydney, N.S.W., in early

November, 1952”. Type: Queensland. Cook

District: Cape Tribulation, 1950, W.W.Mason

s.n. (holo: NSW 22393, fide M.A. Clements

8 May 1986; iso: CNS 137337 [“conveyed by

St Cloud” hand written on Dockrilfs original

folder]).

Dendrobium masonii Rupp has not been re¬

recorded in the wild since it was described,

despite subsequent exploration of the type

locality (Lavarack & Gray 1985). Dockrill

(1992) reported that the type locality had

been cleared for grazing whereas B. Gray

(pers. comm.) reported that the host tree for

the plant from which the type specimen was

prepared had fallen into Bailey Creek.

Upon accessioning the Dockrill herbarium

a duplicate of the W.W. Mason Jnr. collection

was located (CNS 137337).This specimen

appears to be the basis of the habit illustration

in Dockrill (1965), whereas the flowers in the

illustration appear to have been redrawn from

Rupp’s original 1952 figure (Dockrill 1992).

Dockrilfs specimen is annotated as having

been ‘conveyed by S.F. St Cloud’ who was one

of Rupp’s correspondents, known to Rupp as

his ‘Cairns huntsman’ (Gilbert 1992).

Rupp recorded the type as having been

sent to him by W.W. Mason Jnr. from Cape

Tribulation in 1950 as a sterile living plant

which he presumed was a Bulbophyllum

(Rupp 1953). Rupp lodged the living material

at the Sydney Botanic Gardens glasshouse in

1951; it was sent on to Mr & Mrs Loader of

Castlecrag in 1952 and then returned to Rupp

when it flowered (Rupp 1953). The holotype

697

Field & Zich, Dockrill orchid types

is therefore a fifth-hand collection that passed

through several glass-house collections of

cultivated plants. This raises the possibility

that a mix-up occurred in cultivation and that

this taxon is a spurious record for Australia.

Dendrobium masonii was transferred to

the genus Diplocaulobium by Dockrill (1965)

and is currently considered to be a synonym

of Diplocaulobium stelliferum (J.J.Sm.)

A.D.Hawkes from Malesia (Lavarack et. al.

2000). The species is considered to be extinct

in Australia.

It is possible that W.W. Mason Jnr. supplied

Rupp with a specimen of a similar orchid such

as Abaxianthus convexus (Blume) M.A.Clem.

& D.L. Jones, a species that is abundant at the

type locality but was relatively unknown in

Australia at the timeD. masonii was described.

Specimens collected by W.W. Mason Jnr and

A.W. Dockrill when they returned to the type

locality in 1962 clearly belong to A. convexus

(CNS 137455 & CNS 137456).

2. Dockrillia baseyana (St.Cloud) Rauschert,

Feddes Repert. 94: 446 (1983); Dendrobium

baseyanum St.Cloud, N. Queensland

Naturalist 23(110): 1 (1955). Type citation:

“Type in North Queensland Herbarium,

Cairns. Habitat, Kings Plains [south west of

Cooktown], N.Q. Coll. F.L. Basey s.n Type:

(holo: QRS ex CAIRNS, not found; iso: CNS

ex Dockrill herbarium, destroyed; lecto:

‘Kings Plains, North Queensland’, F.L. Basey

s.n. original illustration,^^ Clements (1989:

46).

The F.L. Basey holotype of Dendrobium

baseyanum St. Cloud said to be at CAIRNS

(integrated into QRS in 1971, now integrated

into CNS) was reported missing by Clements

(1989) who nominated the original illustration

as lectotype. Flowers from the F.L. Basey

‘Kings Plains’ specimen were located in the

Dockrill herbarium; however, this duplicate

is an insect-destroyed fragment and no

material remained that was suitable for a type

specimen. Dockrill annotated this specimen

that one of the two flowers was inconsistent

with the description.

This taxon is currently recognised as

a synonym of Dockrillia calamiformis

(G.Lodd.) M.A.Clem. & D.L.Jones (Clements

& Jones 1996).

3. Dockrillia * foederata (St. Cloud)

Rauschert, Feddes Repert. 94: 446 (1983);

Dendrobium foederatum St.Cloud, N.

Queensland Naturalist 23(111): 2 (1955).

Type citation: “Type in North Queensland

Herbarium, Cairns. Growing on Heritiera

littoralis in mangrove swamp, Aeroglen, near

Cairns, leg. J. Dyson-Holland, September,

1954, flowering in cultivation, October 1954

to January 1955.” Type: Queensland. Cook

District: Aeroglen, Cairns, 28 December

1954 , J. Dyson-Holland s.n. (holo: QRS 44141;

iso: CNS 137336).

The holotype of Dendrobium foederatum

St.Cloud lacks flower parts and roots. A

duplicate of this collection was located in the

Dockrill herbarium (CNS 137336). Although

this isotype specimen is insect-damaged it

includes a number of flowers and roots and

thus remains useful for research. A third more

complete specimen (CNS 44140) collected by

L.J.Brass from cultivation may be descended

from the type (i.e. a clonotype).

This taxon is thought to be a natural hybrid

between Dockrillia rigida (R.Br.) Rauschert

and D. calamiformis (Lodd.) M.A.Clem.

& D.L. Jones (the latter originally as

Dendrobium teretifolium R.Br. [= Dockrillia

teretifolia (R.Br.) Brieger] in St Cloud [1955]).

The describing author was of the opinion that

“evidence is against a natural hybrid” (St

Cloud 1955); however, subsequent authors

have all listed it as a naturally occurring

hybrid species (Rauschert 1983; Clements

1989).

4. Eria intermedia Dockrill, Austral. PI. 120

(1964); Bryobium intermedium (Dockrill)

D.L.Jones & M.A.Clem., Orchadian 15(2):

88 (2005). Type citation: “Whitfield

Range, North Queensland (A.W. Dockrill

26/12/1961 - Herb. BRI).” Type: Queensland.

Cook District: Whitfield Rge [Range], 26

December 1961, A.W.Dockrill s.n. (holo: BRI

[AQ279580]; iso: QRS 129124).

A duplicate of the holotype (QRS 129124)

and a cultivated specimen of Eria intermedia

(CNS 132279) were located in the Dockrill

698

herbarium. These collections all originate

from the Dockrill type collection from the

Whitfield Range, Cairns on 26 Dec 1961,

which remains the only authentic herbarium

record of this taxon in Australia.

This taxon is currently recognised as

a synonym of Eria dischorensis Schltr.

(Dockrill 1969), a species that has a

distribution primarily in Papua New Guinea,

although Jones & Clements (2005) recognised

it as Bryobium intermedium and endemic to

Australia.

5. Oberonia attenuata Dockrill, N.

Queensland Naturalist 29(126): 4, figs A-I

(1960). Type citation: “North Queensland,

Mossman River 12-6-1960, Leg. A.W.

Dockrill”. Type: Queensland. Cook District:

Mossman River, 12 June 1960, A.W.Dockrill

s.n. (holo: BRI [AQ279632]; iso: CNS 137338).

Oberonia attenuata has not been recollected

in Australia since it was described in 1960

and is listed as Extinct under State and

Commonwealth legislation. Duplicates of

the holotype and two previous unrecorded

collections were found in the Dockrill

herbarium, greatly adding to the known

material of this species. O. attenuata is

unique among Australian Oberonia in having

long (up to 160 mm) and narrow (4-8 mm)

pendulous falcate-subulate leaves and flowers

with a labellum bearing deeply bifid or trifid

side and front lobes. The cause of the apparent

extinction of this species at the type locality is

poorly understood. Further searches for this

species are needed over a wider range than

previously surveyed.

Additional specimens examined : Queensland. Cook

District: Mossman River, Jan 1960, Archer et al. s.n.

(CNS 137339); Babinda, May 1956, Wilkie s.n. (CNS

137340). Additional replicates of these specimens will

be distributed to BRI and CANB.

References

Clements, M.A. (1989). Catalogue of Australian

Orchidaceae. Australian Orchid Research 1.

Australian Orchid Foundation: Essendon.

Clements, M.A. & Jones, D.L. (1996). New species of

Dendrobiinae (Orchidaceae) from Papua New

Guinea. Lasianthera 1: 8-25.

Dockrill, A.W. (1960). A new species of Oberonia

(Orchidaceae) from north Queensland. The

Austrobaileya 8(4): 696-698 (2012)

North Queensland Naturalist 29( 126): 4.

- (1965). The genera Diplocaulobium and

Ephemerantha in Australia. The Orchadian

1(11): 132-134.

- (1969). Australian Indigenous Orchids. Society

for Growing Australia Plants: Sydney.

- (1992). Australian Indigenous Orchids 1: 310-

311. Surrey Beatty & Sons: Chipping Norton.

Gilbert, L.A. (1992). The orchid man: the life, work and

memoirs of the Rev. H.M.R. Rupp, 1872-1956.

Kangaroo Press: Kenthurst.

Jones, D.L. & Clements, M.A. (2005). Miscellaneous

nomenclature notes and changes in Australian,

New Guinea and New Zealand Orchidaceae.

The Orchadian 15: 33-42.

Lavarack, PS. & Gray, B. (1985). Australian Tropical

Orchids. Nelson: Melbourne.

Lavarack, PS., Harris, W.E. & Stocker, G. (2000).

Dendrobium and its relatives. Kangaroo Press;

East Roseville.

Rauschert, S. (1983). Beitrag zur Nomenklatur der

Orchidaceae. Feddes Repertorium. Zeitschrift

fur Botanische Taxonomie und Geobotanik 94:

433-471.

Rupp, H.M.R. (1953). A new Australian Dendrobium.

Australian Orchid Review 18: 18.

St. Cloud, S.F. (1955). New species of Dendrobium

from north Queensland. The North Queensland

Naturalist 23(111): 2-4.

New species and subspecies of Ipomoea L. (Convolvulaceae)

from northern Australia and a key to the Australian species

R.W. Johnson

Summary

Johnson, R.W. (2012). New species and subspecies of Ipomoea L. (Convolvulaceae) from northern

Australia and a key to the Australian species. Austrobaileya 8(4): 699-723. Ipomoea bracteolata

R.W. Johnson, I. densivestita R.W. Johnson, I. dunlopii R.W. Johnson, I. funicularis R.W. Johnson, 1.

kalumburu R.W.Johnson, 1. limosa R.W.Johnson, I. tolmerana R.W.Johnson, I. tolmerana subsp.

occidentalis R.W.Johnson and I. versipellis R.W.Johnson are described as new species or subspecies.

All newtaxa are illustrated and notes are given on their distribution, habitat, phenology, affinities and

conservation status. A key to all Australian species of Ipomoea is provided.

Key Words: Convolvulaceae, Ipomoea, Ipomoea bracteolata, Ipomoea densivestita, Ipomoea

dunlopii, Ipomoea kalumburu, Ipomoea funicularis, Ipomoea limosa, Ipomoea tolmerana, Ipomoea

tolmerana subsp. occidentalis, Ipomoea versipellis, Australian flora, identification key, new species,

taxonomy

R.W. Johnsonf, c/o Queensland Herbarium, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong,

Queensland 4066, Australia.

Introduction

Ipomoea L. is a genus of more than 600 species

(Austin & Hauman 1996) found mainly in

tropical and subtropical regions, particularly

in the Americas. Bentham (1869) listed 38

species in his Flora Australiensis treatment,

but many have since been transferred to

segregate genera such as Davenportia

R.W.Johnson, Jacquemontia Choisy,

Lepistemon Blume, Merremia Dennst. ex

Endl. and Xenostegia D.F.Austin & Staples.

In Australia, there are more than 50 species,

including 16 which have become naturalised.

Most of the native species are endemic.

This paper has been prepared as a

precursor to an account of Convolvulaceae

for the Flora of Australia. While it covers

most of the currently unresolved problems in

the genus, both Ipomoea abrupta R.Br. and I

gracilis R.Br. embrace considerable variation

and need more careful study.

Materials and methods

This paper is based mainly on specimens held

at BRI and borrowed from DNA and PERTH.

Common abbreviations used in the citation

of specimens include N.P. (National Park).

Accepted for publication 9 July 2012

f Deceased

Abbreviations for Australian states are NSW

(New South Wales), NT (Northern Territory),

Qld (Queensland), WA (Western Australia),

Vic (Victoria).

Taxonomy

Ipomoea bracteolata R.W.Johnson species

nova; affinis I yardiensi A.S.George, sed

caulibus serpentibus et volubilibus (non

fruticosis), sepalis multo longioribus, >15

mm longis (non 8-11 mm longis) et bracteolis

persistentibus et multo longioribus, 10-15

mm longis (non 2-3 mm longis) differens.

Planta perennis, caules serpentes et volubiles.

Folia simplicia, lamina ovata usque ad

late ovata, integra. Inflorescentia cymosa,

1-flora. Sepala exteriora ovata, acuta, ±

laevia. Corolla infundibuliformis taeniis

mesopetalinis pilosis. Semina puberula,

caespite pilorum longiorum ad hilum. Typus:

Northern Territory. Darwin And Gulf: near

quarry road, Batchelor, (13°02'S 13P05'E),

15 February 1991, 1. Cowie 1384 & P. Munns

(holo: BRI; iso: DNA n.v., MEL n.v).

Ipomoea sp. Cobourg (G.M.Wightman 380);

Short et al. (2011).

Ipomoea sp. Mt Fyfe (R.L.Barrett &

M.D.Barrett RLB 1526); Western Australian

Herbarium (2011).

700

Perennial with trailing and twining stems;

stems terete, herbaceous, moderately densely

to densely hairy, hairs crisped-appressed

to ascending or spreading, 0.7-2 mm long.

Leaves simple, petiolate; petiole 7-20 mm

long, 0.13-0.4 times as long as the blade,

vestiture as for stem; blade ovate to broadly

ovate or ovate-oblong, 27-65 mm long, 13-55

mm wide with a length:width ratio of 1.1-2.2,

entire, apex obtuse to rounded, rarely sub¬

acute, with a short mucro, c. 0.6 mm long,

base rounded to sub-cordate, densely to

moderately densely hairy on both sides with

semi-appressed to ascending hairs 0.5-2 mm

long, midrib with 5-7 pairs of secondary

veins, raised below. Inflorescence axillary,

cymose, 1-flowered; peduncle terete, 13-60

mm long, vestiture as for the stem; bracteoles

opposite, herbaceous, narrowly ovate-

lanceolate, sometimes acuminate, 10-20 mm

long, 3-6 mm wide, vestiture as for leaves,

persistent at flowering and early fruiting,

apex acute, mucronulate, attenuate to rounded

at base; pedicels terete, 5-10 mm in flower,

15-23 mm in fruit, vestiture as for the stem.

Outer sepals narrowly ovate, ovate-elliptic

or lanceolate, 20-25 mm long, 6-10 mm

wide in flower, with a length:width ratio of

2.5-3.5, apex acute, mucronulate, attenuate to

rounded at base, smooth, moderately densely

to densely hairy with hairs up to 2 mm long;

inner sepals narrowly ovate-lanceolate,

20-25 mm long, 3.5-4 mm wide, apex acute,

mucronulate, attenuate to rounded at base,

with a hyaline basal margin, moderately

densely hairy in the upper part and along the

spine. Corolla funnel-shaped with a slender

tube, pale pink to pale mauve with a darker

throat, 50-80 mm long, 35-55 mm diameter;

petals 65-95 mm long, 25-35 mm wide, lobes

rounded, midpetaline band moderately hairy

to within 20-40 mm of base of corolla, hairs

0.3-0.8 mm long, mainly appressed, antrorse.

Stamens included, attached to the base of the

corolla for c. 20 mm; filaments free for 10-15

mm, with multicellular hairs around the point

of attachment; anthers oblong, sagittate, c.

2.8 mm long with basal lobes 0.6-0.7 mm

long; pollen globular, spinulose. Ovary

2-locular, glabrous; style 30-40 mm long;

stigma biglobular. Capsule ovoid, c. 10 mm

Austrobaileya 8(4): 699-723 (2012

long, valvate-dehiscent, glabrous; seeds 4, c. 7

mm long, dark brown, sparsely to moderately

puberulent, with slightly longer hairs, 0.2-0.4

mm long, on the margins and a tuft of long

whitish hairs to 1 mm around the hilum. Fig.

1 .

Additional specimens examined : Western Australia.

Mitchell Plateau, river plain on Camp Creek below

crusher, Feb 1979, Beard 8323 (BRI); 2 km SW of

Mt Fyfe, Drysdale River Station, Jan 2001, Barrett

& Barrett RLB1526 (PERTH). Northern Territory.

Cobourg Peninsula, Smith Point, May 1983, Wightman

380 (DNA); Cobourg Peninsula, SE of Port Bremer

pearl farm, Feb 2005, Cowie & Brennan 10440 (DNA);

Cobourg Peninsula, track to Observation Cliff, Feb

1994, Egan 3130 (BRI); Kakadu N.P. on track to

Nangalor Gallery, Feb 1991, Brennan 964 (DNA); near

T/O [turnoff] to Batchelor quarry. Mar 1991, Cowie 1581

& Munns (BRI, DNA).

Distribution and habitat : Ipomoea

bracteolata occurs in the Drysdale River

- Mitchell Plateau area of the Kimberley,

Western Australia, in the north western area

of the Northern Territory on the Cobourg

Peninsula and near Batchelor and Jabiru

(Map 1). It grows in eucalypt woodland

communities, often with Eucalyptus miniata

A.Cunn. ex Schauer and Corymbia, on sandy

soils on rocky hills.

Phenology : Flowers have been recorded from

January to May; immature fruits have been

recorded in February.

Affinities: Ipomoea bracteolata is a rather

distinctive species. It resembles I. yardiensis

in having a corolla with a hairy mid-petaline

band but it is a vine with trailing and twining

stems, not a shrub. Its bracts are persistent and

large, unlike the small bracts of I. yardiensis

which abscise pre-flowering.

Conservation status: It is regarded as a

priority 1 taxon in Western Australia (Western

Australian Herbarium 2011). It is not listed

among the threatened species in the Northern

Territory (Short et al. 2011).

Etymology: The specific epithet is derived

from the Latin bracteola, meaning a bracteole.

This refers to the pair of distinctive bracteoles

at the base of the pedicel.

Johnson, New species of Ipomoea

701

Fig. 1. Ipomoea bracteolata. A. flowering branch x l.B. outer sepal at flowering x4. C. inner sepal at flowering *4. A

from Cowie & Brennan 10440 (DNA); B-C from Barrett RLB1526 & Barrett (PERTH). Del. W.Smith

Ipomoea densivestita R.W. Johnson species

nova; affinis I. abruptae R.Br., sed seminibus

puberulis, pilis longis in marginibus externis

(non lanatis) et sepal is pilosis (non glabris)

differens. Planta perennis, caules repentes

et volubiles. Folia simplicia, lamina late

ovata, integra. Inflorescentia cymosa,

1-4-flora. Sepala exteriora ± laevia. Corolla

infundibuliformis, glabra. Semina sparsim

puberula, pilis longis in marginibus externis.

Typus: Queensland. Cook District: entrance

to Maidenhair Grotto, Chillagoe, 25 January

1984, M.Godwin C2541 (holo: BRI).

Ipomoea lasiophylla Domin, Biblioth. Bot.

89(6): 536 (1928), nom. illeg., non Hallier f.

(1893). Type: Queensland. Cook District:

in collibus caraticis apud opp. Mungana,

February 1910, K.Domin s.n. (holo: PR

530547, photo BRI).

Ipomoea sp. (Mungana L.J.Webb+ 10184);

Johnson (2007, 2010).

Perennial vine with trailing and twining

stems; stems terete, woody at the base,

moderately to densely hairy, hairs tubercle-

based, simple or bifid, 0.1-0.5 mm long.

Leaves simple, petiolate; petiole 20-70 mm

long, 0.35-0.7 times as long as the blade; blade

broadly ovate to ovate-orbicular, 40-150 mm

long, 30-120 mm wide with a length:width

ratio of 1.1-1.6, apex obtuse to rounded,

sometimes slightly emarginate, mucronulate,

base cordate with a narrow sinus to 20% of

702

blade length, densely pubescent on both sides,

midrib with a pair of small slit-shaped glands

at the base and 9-12 pairs of secondary veins.

Inflorescence axillary, compound cymose,

1-5-flowered; peduncle stout, terete, 5-75

mm long, vestiture as for stem; bracteoles

opposite, herbaceous, oblong to obovate-

oblong, rounded at apex, abscissing prior to

flowering; pedicels terete, dilated upwards,

10-30 mm long, vestiture slightly less dense

than peduncle, with 5 small slit-shaped glands

below the calyx. Outer sepals concave,

broadly ovate, broadly oblong to orbicular,

8-10 mm long, 7-9 mm wide in flower,

apex rounded, base rounded, thickened

with a narrow hyaline margin, surface

smooth, densely to moderately densely hairy,

extending to 14 mm x 14 mm at fruiting,

becoming bullate and sparsely hairy; inner

sepals of similar shape and size to inner, hairs

restricted to spine. Corolla funnel-shaped,

with a cylindrical tube, pink to pale purple,

midpetaline band and throat darker, 50-60

mm long, 50-60 mm diameter; petals to

70 mm long, lobes rounded, emarginate,

midpetaline band glabrous. Stamens 5,

included, attached to the base of the corolla for

c. 14 mm; filaments free for 13-16 mm, with

multicellular hairs from just below the point

of attachment upwards for c. 4 mm; anthers

narrow-lanceolate, sagittate, c. 5 mm long and

1 mm wide; pollen globular, spinulose. Ovary

hemispherical, 2-locular, glabrous; style c.

27 mm long; stigma biglobular. Capsule

globular, 8-11 mm long, with a persistent style

base, 7-11 mm diameter, valvate-dehiscent,

glabrous; seeds 4, 5-7 mm long, brown,

moderately to sparsely puberulent, with long

hairs to 2 mm on the outer margins. Fig. 2.

Additional specimens examined : Queensland. Cook

District: Palmerville Holding, Oct 1990, Godwin C3649

(BRI); Belgravia, 3 km W of Mungana, May 1999,

Gray 7553 (BRI); Mungana N.R, The Archways, NW of

Chillagoe, Feb 1996, Forster PIF18598 & Ryan (BRI);

4 miles [c. 6 km] N of Mungana, May 1970, Webb &

Tracey 10184 (BRI); Belgravia, 2.5 km E of Mungana,

Feb 1999, Gray 7445 (BRI); Royal Arch Tower, c. 5 km

SW of Chillagoe, Mar 1987, Clarkson 6829 & McDonald

(BRI).

Distribution and habitat: Ipomoeadensivestita

is endemic to a small area in north-eastern

Queensland (Map 2). Collections have been

Austrobaileya 8(4): 699-723 (2012

made from near Chillagoe and Palmerville. It

grows in deciduous vine thickets on limestone

outcrops.

Phenology: Flowers have been recorded from

January to May; fruit recorded in May and

October.

Affinities: Ipomoea densevestita is related

to I. abrupta, which is extremely variable

and contains a complex of taxa including

I. velutina R.Br. and the Western Australian

taxa, /. sp. (Carson Escarpment A.S.George

13732) and /. sp. (Cascade Creek R.L &

M.D.Barrett RLB3738). In a narrow sense

I. abrupta is a glabrous liana while I. velutina

has pubescent foliage though the calyx is ±

glabrous. The seeds of I. abrupta sens. I at. are

covered in woolly hairs, unlike I. densivestita

which has puberulent seeds with longer hairs

on the outer margins. Ipomoea abrupta sens,

lat. grows in sandstone areas, mainly in forests

whereas I. densivestita appears restricted to

vine thicket vegetation growing on limestone.

More studies of specimens in the I. abrupta

complex are required.

Conservation status: Ipomoea densivestita

occurs in a small geographical area near

Chillagoe and Palmerville in north-eastern

Queensland. It is rare in that area and requires

conservation assessment.

Etymology: The specific epithet is derived

from the Latin densus, meaning dense and

vestitus , meaning vestiture. This refers to the

dense indumentum on stems and leaves and

reflects the intent of the epithet, lasiophylla

used by Domin.

Ipomoea dunlopii R.W. Johnson species

nova; affinis I. abruptae R.Br., sed sepalis

breviora, laevibus (non bullatis), praesertim

fructiferis longitudinis 1/3 ostendens (non

± perfecte capsulam maturam includentibus)

differens. Planta perennis, caules serpentes

et volubiles. Folia simplicia, late ovata,

interdum hastata. Inflorescentia cymosa,

l-3(-5)-flora. Sepala exteriora late ovata

usque ad orbicularia, ± laevia. Corolla

infundibuliformis, glabra. Semina lanata.

Typus: Northern Territory. Darwin & Gulf:

Mt Brockman Outlier, 15 km SE of Jabiru, 20

April 1989, R.W.Johnson 4732 (holo: BRI; iso:

DNA, PERTH, distribuendi).

Johnson, New species of Ipomoea

Glabrous liana with trailing and twining

stems; stems terete, woody at the base.

Leaves simple, petiolate; petiole 25-75 mm

long, 0.35-0.7 times as long as the blade;

blade narrowly to broadly ovate or triangular,

sometimes with rounded hastate lobes at the

base, 50-150 mm long, 15-120 mm wide,

with a length:width ratio of 1-4, apex acute,

mucronate, base truncate to cordate with

a broad sinus, midrib with a pair of slit¬

shaped glands at the base and 7-12 pairs

703

of secondary veins and coarsely reticulate

venation. Inflorescence axillary, compound

cymose, 1-6(1 l)-flowered; peduncle stout,

terete, 2-30 mm long; bracteoles opposite,

herbaceous, abscissing prior to flowering;

pedicels terete, dilated upwards, 12-35 mm

long, with 5 small slit-shaped glands below the

calyx. Outer sepals concave, broadly ovate

to broadly oblong 5.5-8.5 mm long, 6-7 mm

wide in flower, becoming ovate-orbicular in

fruit and extending to 10 mm x 9.5 mm, apex

Fig. 2. Ipomoea densivestita. A. flowering branch x0.8. B. outer sepal at flowering M. C. inner sepal at flowering *4.

A-C from Clarkson 6829 & McDonald (BRI). Del. W.Smith.

704

obtuse to rounded, base obtuse to rounded,

thickened with a narrow hyaline margin,

surface smooth, rarely slightly wrinkled;

inner sepals broadly oblong, broadly ovate or

orbicular, 7.5-9 mm long, 8-9.5 mm wide,

becoming broader in fruit and extending to

10 mm x n.5 mm, apex rounded, slightly

emarginate, base rounded to subcordate.

Corolla funnel-shaped, 55-70 mm long,

mauve to purple; petals 65-85 mm long,

lobes rounded, emarginate, midpetaline band

glabrous. Stamens 5, included, attached to the

base of the corolla for 10-11 mm; filaments

free for 15-24 mm, with multicellular hairs

from just below the point of attachment for

6-7 mm; anthers linear to oblong, sagittate,

6-7 mm long, c. 1 mm wide with basal lobes

0.75-1 mm long; pollen globular, spinulose.

Ovary 2-locular, glabrous; style 32-40 mm

long; stigma biglobular. Capsule globular to

depressed-globular, 10-13 mm long, 10-14

mm diameter, valvate-dehiscent, glabrous;

seeds 4, 6-7 mm long, black, covered in very

long dense, sinuous, white to pale brown hairs

to 4 mm long, longest on back and margins.

Fig. 3.

Additional specimens examined: Northern Territory.

10 km SW of Oenpelli Aboriginal Settlement, May 1988,

Weber 9860 (AD, BRI, DNA); Cannon Hill Airstrip,

Jan 1973, Martensz AE612 (BRI, DNA); Kakadu N.P.,

Ngarradj, Jan 1992, Russell-Smith 8562 & Lucas (BRI);

East Alligator River, Feb 1973, Dunlop 3204 (BRI); 2

km SW Ngarradj Creek crossing, Jabiru - Oenpelli

Road, Jan 1984, Russell-Smith 1175 (DNA); SE of Mt

Howship, Feb 1984, Dunlop 6638 (DNA); Arnhem Land,

19 km E of Jabiru, Apr 1989, Johnson 4520, 4527, 4600

(BRI); Upper East Alligator River, Arnhem Land, Feb

1991, Brock 768 & Russell-Smith (BRI, DNA); Kakadu

N.R, Koongarra, Feb 1992, Russell-Smith 8709 (DNA);

Mt Brockman, s.dat., Smyth s.n. (BRI); Mt Brockman

Outlier, 15 km SE of Jabiru, Apr 1989, Johnson 4634,

4786, 4823 (BRI); Kakadu N.R, South Magela Gorge,

8 km SW of Mt Brockman, Jabiru Town, Jan 1991,

Brennan 901 (DNA); Kakadu N.R, Baroalba Creek,

upstream from springs, Jabiru Town, Dec 1990, Brennan

781 (DNA); near Buffalo Springs, Mt Brockman, 4 km

NNE of Koongarra, May 1980, Craven 5764 (BRI,

CANB, DNA); Little Nourlangie Rock, Mar 1978,

Dunlop 4703 (BRI, DNA); Deaf Adder Gorge, Feb 1977,

Fox 2507 (DNA); 6 km S of Mt Gilruth, Arnhem Land,

Mar 1984, Jones 1537 (BRI, DNA); top of Jim Jim Falls,

Jan 1981, Dunlop 5694 (BRI, DNA); Kakadu, graveside

gorge, Dec 1986, Brock 192 (DNA); 1 km upstream from

Twin Falls, Mar 1988, Fensham 777 (DNA).

Austrobaileya 8(4): 699-723 (2012

Distribution and habitat: Ipomoea dunlopii

is found in a small area of Arnhem Land S

of Oenpelli (Map 2). It occurs on the top,

and in flats and gullies, of broken sandstone

plateaux. It grows in vine thickets, sandstone

heaths and eucalypt forests and woodlands on

sandy soils.

Phenology: Flowers have been recorded from

January to April; fruit have been recorded

from March to May.

Affinities: Ipomoea dunlopii is related to

I. abrupta which occurs mainly in coastal and

subcoastal areas. It differs from I. abrupta in

having sepals, at fruiting, which are smooth,

shorter and enclose only the lower two-thirds

of the capsule. In I. abrupta the sepals become

bullate and enclose the mature capsule.

Juvenile foliage in I. dunlopii can become

narrow and distinctly hastate which I have not

noticed in I. abrupta.

Conservation status: It is found in a relatively

restricted but protected area in Arnhem Land

and is not known to be threatened.

Etymology: This species is named in honour

of a friend and colleague, Clyde Dunlop,

who made possible my trip to Arnhem Land,

where the type of this species was collected.

Ipomoea funicularis R.W. Johnson species

nova; affinis I. gracili R.Br., sed caulibus

funicularibus (non herbaceis), foliis semper

hastato-sagittatis (raro hastatis), sepal is

exterioribus sub anthesi longioribus, 10-17

mm longis (non 8-13 mm longis) et petalis

longioribus, 60-75 mm longis (non 45-60

mm longis) differens. Planta perennis,

caules repentes et volubiles, lignescentes.

Folia simplicia, lamina hastata usque ad

sagittata. Inflorescentia cymosa, 1-2-flora.

Sepala exteriora ovata, acuta, nervis 3-5,

longitudinalibus, elevatis, tuberculatis

et rugosis in dimidio inferiora. Corolla

infundibuliformis, glabra. Semina puberula,

caespite longiorum pilorum ad hilum. Typus:

Queensland. Cook District: Undara National

Park, NE of Barkers Knob, 14 February 2005,

K.R.McDonaldKRM3610 & C.O’Keefe (holo:

BRI, iso: DNA, PERTH, distribuendi).

Johnson, New species of Ipomoea

Ipomoea gracilis var. sagittata F.Muell.,

Fragm. 6: 99 (1868). Type: Queensland.

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

(lecto: MEL 2260143, here designated;

isolecto: MEL 2260141).

Perennial with a tuberous root and trailing

stems, twining at the tips; stems terete, tough

and fibrous, glabrous, sometimes sparsely

hairy. Leaves simple, petiolate; petiole

14-50 mm long, 0.25-0.65 times as long as

the blade, ± glabrous, occasionally with a

few, weak, sinuate hairs and low tubercles;

blade hastate to hastate-sagittate, 55-100 mm

705

long, 30-60 mm wide, with a narrow-linear

terminal lobe, 40-90 mm long, 2-18 mm

wide, and a pair of spreading linear to linear-

triangular basal lobes, apex narrowly obtuse

to rounded, sometimes retuse, mucronate,

base subcordate to cordate with a wide sinus

up to 22% of blade, glabrous, midrib with a

pair of slit-shaped basal glands and 6-8 pairs

of secondary veins. Inflorescence axillary,

cymose, 1-2-flowered, rarely compound;

peduncle terete, stout, 2-20 mm long,

±glabrous, occasionally with vestiture as for

petiole; bracteoles opposite, herbaceous,

Fig. 3. Ipomoea dunlopii. A. fruiting branch x0.8. B. outer sepal at flowering x4. C. inner sepal at flowering x4. A

from Johnson 4786 (BRI); B-C from Johnson 4732 (BRI). Del. W.Smith

706

narrowly triangular, 3-8 mm long, apex acute,

persistent; pedicels terete, dilated upwards,

7-16 mm long, bearing 5 slit-shaped glands

below the calyx. Outer sepals ovate, narrowly

ovate or ovate-elliptic to ovate-lanceolate,

10- 15 mm long, 3.5-5.5 mm wide in flower,

extending to 8 mm wide in fruit, apex acute,

often inrolled, base rounded, surface with

3- 5 prominent longitudinal veins, with

some tubercles and transverse ridges; inner

sepals narrowly ovate to ovate-elliptic,

11- 16 mm long, 5-6 mm wide, extending

to 9 mm wide in fruit, apex inrolled, acute,

base truncate to sub-cordate, with 3 slightly

raised longitudinal veins and a narrow

hyaline margin, broader at the base. Corolla

funnel-shaped, 50-60 mm long; petals 60-75

mm long, pink, often with a darker centre,

lobes rounded, apiculate, midpetaline band

glabrous. Stamens 5, included, attached to

the base of the corolla for 8-10 mm; filaments

free for 10-22 mm, with dense, sinuate,

multicellular hairs from just below the point

of attachment upwards for ± 6 mm; anthers

linear to narrowly ovate-lanceolate, sagittate,

3.5-4 mm long, with basal lobes 0.5-0.75

mm long; pollen globular, spinulose. Ovary

2-locular, ovoid with a prominent disk; style

27-40 mm long; stigma biglobular. Capsule

globular to depressed globular, 8-10 mm

long, valvate-dehiscent, glabrous; seeds 4,

4- 5 mm long, dark brown, puberulent with a

tuft of long silvery brown hairs to c. 1.5 mm

around the hilum. Fig. 4.

Additional selected specimens (from 29 examined ):

Queensland. Burke District: 38.6 km SW of Hells Gate

Roadhouse on Bowthorn Station, Apr 2006, Thompson

& Edginton WES331 (BRI). Cook District: c. 28 km

SSE of Laura, just S of Hell’s Gate, Jun 2006, Wannan

& Caldwell 4457 (BRI); Mareeba, 0.5 miles [c. 1 km]

S on Atherton Road, Jan 1962, Webb 5528 (BRI); 5

km by road from junction with Burke Development

Road towards Ootann, Jan 2005, McDonald KRM3516

(BRI); Almaden - Mt Surprise Road, near Almaden,

Jun 2008, McDonald KRM5327 & Sellars (BRI); 35

km from Almaden on road to Mt Surprise, Jan 1992,

Forster PIF9620 (BRI); Blue Hills, Mt Surprise, 49 km

from Mt Surprise Township, Mar 1988, Champion 363

(BRI); 3 km E of Mt Surprise, Feb 2004, McDonald

KRM1791 (BRI); Undara N.P., NE of Barkers Knob, Dec

2006, McDonald KRM4681 (BRI); 62.2 km by road W

of Mt Surprise on E approach to Newcastle Range, Mar

2006, McDonald KRM4928 (BRI); Newcastle Range,

64 km from Mt Surprise on Georgetown Road, Mar

1988, Forster PIF3811 (BRI); Black Rock near Lynd

Austrobaileya 8(4): 699-723 (2012

on Hughenden Road, Feb 1988, Horsup 6 (BRI); 19.7

km by road towards Forsayth from Einasleigh, Mar

2005, McDonald KRM3787 (BRI); Newcastle Range,

Agate Creek catchment, Simpson’s Gully, Apr 2006,

McDonald KRM5245 (BRI). North Kennedy District:

Herberton, Jan 1912, Kenny s.n. (BRI); Castle Hill,

Townsville, Mar 1992, Bean 4099 (BRI); Mt Norman,

55 km S of Townsville, Jan 1999, Camming 18473

(BRI); Princess Dam Road, Lumholtz N.P., Jan 2003,

McDonald KRM1226 (BRI). South Kennedy District:

Broken River Range, Apr 1978, Byrnes & Clarkson 3672

(BRI). Mitchell District: c. 32 km SE of Hughenden,

Apr 1998, Thompson & Turpin HUG582 (BRI).

Distribution and habitat : Ipomoea

funicularis occurs in north-eastern

Queensland from south of Laura, south to

the Broken River Range, west of Mackay

and inland to the Gulf of Carpentaria, near

Burketown, SSE of Georgetown and to the

south-east of Hughenden (Map 1). It grows

in eucalypt woodland communities mainly

with ironbarks and bloodwoods on sandy,

sometimes gravelly soils, on rocky slopes and

low hills, derived mainly from granite.

Phenology : Flowers have been recorded from

December to June; fruits have been recorded

from December to April.

Affinities: Ipomoea funicularis is closely

related to I. gracilis. It has a much more

robust habit with wiry, semi-woody stems

much tougher than the herbaceous stems of I.

gracilis. Its leaves are always hastate-sagittate

at the base, though on rare occasions I.

gracilis can have hastate lobes. The calyx and

corolla of I. funicularis are larger than that of

I. gracilis. Its foliage can also resemble that of

I. brassii C.T.White but the species are clearly

distinguished by seed vestiture. Ipomoea

funicularis has puberulent seeds with a tuft

of longer hairs at the hilum whereas the seeds

of I. brassii are villose. The peduncle is also

much longer than the pedicel in I. brassii and

the corolla lacks the darker centre which is

found in I funicularis.

Conservation status : Ipomoea funicularis

is a widespread species and not known to be

threatened.

Etymology : The specific epithet is derived

from the Latin funicularis, meaning rope¬

like. This refers to tough, fibrous texture of

the stems, which distinguishes this species

from I. gracilis which has herbaceous stems.

Johnson, New species of Ipomoea

Ipomoea kalumburu R.W. Johnson species

nova; affinis/. muelleri Benth., sed seminibus

puberulis, caespite pilorum longiorum ad

hilum (non villosis) et bracteis, corolla et

sepalis longioribus differens. Planta perennis,

caules serpentes. Folia simplicia, lamina

late ovata usque ad triangulata, integera.

Inflorescentia cymosa, l-(2-3)-flora. Sepala

exteriora ovata, acuta, ± laevia. Corolla

infundibuliformis, glabra. Semina puberula,

caespite pilorum longiorum ad hilum.

Typus: Western Australia. Cattle paddock,

eastern bank of King Edward River, 1.06 km

from Kalumburu Mission, 31 March 2001,

A.A.Mitchell 6685 (holo: PERTH; iso: BRI).

707

Ipomoea sp. Kalumburu (A.A.Mitchell 3869B);

Short et al. (2011); Western Australian Herbarium

( 2011 ).

Glabrous perennial with trailing, scrambling

and climbing stems; stems terete, herbaceous,

sometimes rooting at the basal nodes. Leaves

simple, petiolate; petiole 15-110 mm long,

0.45-0.8(-l.3) times as long as the blade;

blade broadly ovate-triangular to deltoid,

entire, 25-55(-100) mm long, 25-45(-70)

mm wide with a length:width ratio of 1.1-1.6,

apex obtuse to narrowly rounded, emarginate,

mucronate, base truncate to shallowly cordate

with wide sinus up to 17% of blade, midrib

with a pair of slit-shaped glands at the base

Fig. 4. Ipomoea funicularis. A. flowering branch xl.5. B. outer sepal at flowering ><4. C. inner sepal at flowering *4.

A-C from McDonaldKRM4928 (BRI). Del. W.Smith.

708

and 3-6 pair of secondary veins often with

a purplish or reddish tinge. Inflorescence

axillary, cymose, l-(2-3)-flowered; peduncle

terete, slender, 2-120 mm long; bracteoles

opposite to sub-opposite, herbaceous,

narrowly triangular-ovate, 3-7 mm long,

apex acute, persistent; pedicels terete, dilated

upwards, 6-16 mm long, bearing 5 linear

slit-shaped glands below the calyx. Outer

sepals ovate, narrowly ovate or ovate-oblong

to narrowly ovate-lanceolate, 8-16 mm long,

4-5.5 mm wide in flower, becoming broader

in fruit, apex acute, often inrolled, apiculate,

base rounded, surface with 3 longitudinal

veins, often slightly raised, smooth or with

an occasional tubercle; inner sepals narrowly

ovate-triangular to narrowly deltoid, 10-14

mm long, 5.5-7 mm wide, apex acute, rarely

obtuse, often inrolled, base subcordate with a

narrow hyaline margin, broader at the base.

Corolla funnel-shaped, 40-55 mm long,

40-50 mm diameter, mauve to purple; petals

45-65 mm long, lobes rounded, apiculate,

midpetaline band glabrous. Stamens 5,

included, attached to the base of the corolla

for 6-8 mm; filaments free for 9-16 mm,

with dense, sinuate, multicellular hairs, from

just below the point of attachment upwards

for 5-7 mm; anthers linear to narrowly ovate-

lanceolate, sagittate, 2.3-2.8 mm long, with

basal lobes 0.5-0.6 mm long; pollen globular,

spinulose. Ovary 2-locular, hemispherical

with a prominent disk; style 22-25 mm

long; stigma biglobular. Capsule ovoid to

globular, 10-12 mm long, valvate-dehiscent,

glabrous; seeds 4, 5-6 mm long, dark brown,

puberulent with a tuft of long whitish hairs

around the hilum. Fig. 5.

Additional specimens examined : Western Australia.

S of Loure Creek, Kalumburu Mission, Oct 2002,

Mitchell 7356 (PERTH); cattle paddock, Kalumburu

Mission, N Kimberley, May 1996, Brockway CB80

(BRI, PERTH); c. 0.5 km S of Kalumburu Mission, S

bank King Edward River, Mar 1995, Mitchell 3869

(BRI, PERTH); Kalumburu Mission cattle paddock,

levee King Edward River, Jun 1996, Mitchell 4428 (BRI,

PERTH); south bank of the King Edward River, about 1

km S of Kalumburu Mission, cattle paddock, Feb 1966,

Mitchell 4256 (PERTH).

Austrobaileya 8(4): 699-723 (2012

Distribution and habitat: Ipomoea kalumburu

occurs at Kalumburu, north Kimberley,

Western Australia (Map 1). It is found mainly

in grasslands with Sorghum, Themeda and

Heteropogon on sandy alluvial soils.

Phenology: Flowers have been recorded

between January and October; fruit between

January and June.

Affinities: Ipomoea kalumburu resembles I.

muelleri , but can be distinguished by its longer

calyx and corolla. Its seeds are puberulous,

with a tuft of longer hairs around the hilum,

whereas in I. muelleri the seeds are villous. It

also resembles Ipomoea limosa R.W. Johnson,

but it has a simple cymose inflorescence with

one, rarely 2 or 3 flowers and ± smooth calyx

lobes, unlike I. limosa which has simple and

compound cymes, often many flowered, and

sepals with distinct longitudinal veins which

are tuberculate and rugose, particularly in the

lower half.

Conservation status: It is regarded as a

priority 1 taxon in Western Australia (Western

Australian Herbarium 2011).

Etymology: The specific epithet, kalumburu,

refers to the name of the aboriginal settlement

in northern Western Australia where it is

found. Kalumburu is an aboriginal word

meaning “Path by the river or river crossing”.

Ipomoea limosa R.W. Johnson species nova;

affinis I. argillicola R.W. Johnson, sed laminis

ovato-lanceolatis, sagittatis vel hastatis, <

1.6-plo longioribus quam latioribus (non late

ovatis, cordatis, > 2-plo longioribus quam

latioribus) differens. Planta perennis, caules

serpentes. Folia simplicia, lamina ovato-

lanceolata, sagittata vel hastata. Inflorescentia

cymosa. l-3(-5)-flora. Sepala exteriora ovata,

acuta, nervis 3-5, elevatis, longitudinalibus,

tuberculatis et rugosis in dimidio inferiore.

Corolla infundibuliformis, glabra. Semina

puberula, caespite pilorum longiorum ad

hilum. Typus: Western Australia, c. 1 km E

of Dumas Lookout, Ord Irrigation Project,

23 April 1983, K.L.Wilson 4810 (holo: BRI;

iso: NSW n.v., PERTH).

Johnson, New species of Ipomoea

709

Fig. 5. Ipomoea kalumburu. A. flowering branch x0.8. B. outer sepal at flowering *4. C. inner sepal at flowering x4.

All from Mitchell 3869 (PERTH). Del. W.Smith.

710

Ipomoea sp. (Kununurra T.E.Aplin 6307);

Johnson (2007, 2010).

Perennial with trailing, occasionally climbing

stems; stems herbaceous, stout, terete,

becoming hollow, glabrous to very sparsely

hairy, young stem tips resinous. Leaves

simple, petiolate; petiole 30-140 mm long,

0.4-1.4 times as long as the blade, glabrous,

sometimes with scattered low tubercles;

blade ovate to ovate-lanceolate or oblong-

lanceolate, sagittate, 50-160 mm long, 25-65

mm wide, length:width ratiol.2-2.6, at times

becoming hastate-sagittate with a narrow

triangular terminal lobe 30-80 mm long,

15-45 mm wide extending to 105 mm wide

at the bluntly hastate base with length:width

ratio decreasing to 1, apex narrowly obtuse to

rounded, often emarginate, mucronate, with a

broadly rounded sinus, 5-25% of the length

of the blade, glabrous, midrib with prominent

slit-shaped glands at the base and 6-8 pairs

of secondary veins. Inflorescence axillary,

cymose, 1-5-flowered; peduncle terete,

4—110(—180) mm long, glabrous, sometimes

sparsely tuberculate; bracteoles opposite

to sub-opposite, herbaceous, narrowly

ovate-lanceolate, 3—10(—18) mm long, acute,

mucronulate, attenuate to rounded at base

occasionally becoming foliate, clavate and

rounded at the apex, glabrous, persistent at

flowering and early fruiting; pedicels terete,

dilated upwards, 5-25 mm long, glabrous,

with 5 small slit-shaped glands under the

calyx. Outer sepals ovate-lanceolate, oblong-

lanceolate or narrowly oblong, 13-20 mm

long, 4.5-6.5 mm wide in flower, extending

to 8.5 mm wide in fruit, apex acute, inrolled,

mucronulate, base attenuate to rounded, thick

with a thin hyaline margin, bearing 3-5

raised longitudinal veins tuberculate in the

lower two-thirds, rugose towards the base,

glabrous; inner sepals ovate-lanceolate to

deltoid-lanceolate, 16-19 mm long, 7-8 mm

wide, apex acute, inrolled, mucronulate, base

truncate to subcordate, with 3-5 slightly

raised longitudinal veins and a hyaline basal

margin, glabrous. Corolla funnel-shaped,

50-60 mm long, pale pink to mauve; petals

55-75 mm long, lobes rounded, apiculate,

midpetaline band glabrous. Stamens 5

included, attached to the base of the corolla

Austrobaileya 8(4): 699-723 (2012

for 10-15 mm; filaments free for 10-16 mm,

with multicellular hairs from just below the

point of attachment upwards for 6-8 mm;

anthers oblong-lanceolate, sagittate, 3.2-3.6

mm long with basal lobes 0.6-0.8 mm long;

pollen globular, spinulose. Ovary ovoid with

a prominent disk, 2-locular, glabrous; style

28-35 mm long; stigma biglobular. Capsule

globular to depressed globular, 9-12 mm

long, 9-12 mm diameter, valvate-dehiscent,

glabrous; seeds 4, 5.5-6.5 mm long, brown

to dark brown, puberulent, with a tuft of long

hairs to 2 mm around the hilum. Fig. 6.

Additional specimens examined : Western Australia.

Kimberley Research Station, Ord River, Apr 1958,

Burbidge 5774 (PERTH); 3 km N of Kununurra, Sep

1974, Must 1270 (BRI, PERTH); Block 48, along track on

edge of block near channel, Kununurra, Mar 1978, Aplin

6307 (BRI, PERTH); Kununurra, Mar 1967, Hardy s.n.

(PERTH3639940); Kununurra, May 1967, Scrymgeour

1696 (PERTH); Charnley River Crossing, 57 km N of

Beverley Springs Station Homestead, Dec 1994, Barrett

RLB998 (PERTH); 2 km SW of Beverley Springs Station

Homestead, Feb 1995, Barrett RLB983 (PERTH);

Ord River, 1948, Langfield 16 (PERTH). Northern

Territory. Bradshaw Field Training Area, headwaters of

Little Fitzmaurice River, 65 km from Timber Creek, Apr

2007, Cowie 11520 & Stuckey (DNA); Limmen N.P, c.

25 km N of Limmen Bight River, Apr 2008, Lewis 753

(DNA); 40.5 miles [65 km] S of Willeroo Homestead,

May 1960, Chippendale 6895 (DNA); 25 miles [40 km]

N of Daly Waters, Mar 1955, Winkworth 1112 (DNA);

Tanumbirini Station, Carpentaria Highway, Feb 1988,

Thomson 2247 (DNA); 4 miles [6.5 km] S of Dunmarra,

Mar 1955, Chippendale 1060 (BRI, DNA); 8.5 km S of

Dunmarra, May 1994, Latz 13708 (BRI, DNA); Cattle

Creek Station, 5.6 km N of Mt Gordon, May 2004,

Cowie 10339 & Coleman (DNA). Queensland. Burke

District: 13.5 km SW of Normanton, Apr 1996, Milson

JM1049 (BRI); Normanton - Burketown Road, Jan

2001, McDonald KRM692 (BRI); 6.1 km by road along

Burke Development Road from junction with Gulf

Development Road, Jan 2005, McDonald KRM3444

(BRI); 6.4 km along Burke Development Road from

junction with Gulf Development Road, Apr 2007,

McDonald KRM6532 (BRI); 6.1 km from Burke and

Gulf Development Roads junction, Apr 2009, McDonald

KRM8411 (BRI-spirit only).

Distribution and habitat : Ipomoea limosa

occurs from east of Derby to Kununurra in

the Kimberley region of Western Australia,

eastwards to south of the Gulf of Carpentaria,

near Normanton in Queensland (Map 2). It

grows in grasslands with Sorghum , Themeda

and Dichanthium ; open woodlands with

emergent eucalypts such as coolibah and box,

occasionally with Melaleuca, on alluvial clay

soil plains, commonly in moister areas.

Johnson, New species of Ipomoea

711

Fig. 6. Ipomoea limosa. A. flowering branch x0.8. B. outer sepal at flowering x4. C. inner sepal at flowering x4. A

from McDonald KRM3444 (BRI) and photos of McDonald KRM6532, 8411 (BRI); B-C from McDonald KRM8411

(BRI). Del. W. Smith.

Phenology : Flowers have been recorded from

January to May and also in September; fruits

have been recorded in April and May.

Affinities : Ipomoea limosa resembles I.

argillicola but can be distinguished by its leaf

shape which is ovate-lanceolate, >2 times as

long as broad and sagittate or hastate at the

base. In I. argillicola the leaves are broadly

ovate, <1.6 times as long as wide and cordate

to truncate at the base. I. limosa also has much

shorter peduncles and pedicels. I. limosa is

found in moist depressions on clay soil plains.

I. argillicola occurs on cracking clay soils in

grasslands and grassy eucalypt woodlands.

Conservation status : Ipomoea limosa is

widely spread throughout northern Australia

and is not known to be threatened.

Etymology : The specific epithet is derived

from the Latin limosa, meaning muddy. This

refers to the habitat of this species, which

commonly grows in muddy places.

712

Ipomoea tolmerana R.W. Johnson species

nova; affinis I. gramineae R.Br., sed laminis

hastatis (non integris), sepalis externis rugosis

(non laevibus) et corollis infundibuliformibus

(non hypocrateriformibus) differens. Planta

perennis, caules serpentes et volubiles. Folia

simplicia, lamina peranguste linearis, vix

usque ad manifeste hastata. Inflorescentia

cymosa, l-(2-3)-flora. Sepala exteriora ovata,

acuta, nervis 3-5, longitudinalibus, elevatis,

tuberculatis et rugosis in dimidio inferiore.

Corolla infundibuliformis, glabra. Semina

puberula, caespite pilorum longiorum ad

hilum. Typus: Northern Territory. Litchfield

National Park, Walker Creek Ranger Station,

14 March 1995, I.D.Cowie 5366 & S.Taylor

(holo: BRI [AQ597032]; iso: DNA).

Perennial with a tuberous root and trailing

and twining stems, rooting at the lower nodes;

stems terete, herbaceous, glabrous or with a

few hairs at the leaf nodes, occasionally very

sparsely to moderately hairy with tubercle-

based hairs to 0.5 mm. Leaves simple,

petiolate; petiole 7-45 mm long, 0.1-0.3

times as long as the blade, vestiture as for the

stem; blade narrowly linear to very narrowly

ovate, 60-150 mm long, with a slightly to

distinctly hastate base, terminal lobe 1-6

mm wide, basal lobes 4-50 mm long, 1-2

mm wide, often recurved, apex acute to

narrowly obtuse, apiculate, base truncate to

subcordate, glabrous, occasionally sparsely

hairy mainly below, with hairs confined

to major veins, erect to retrorse, tubercle-

based, midrib with a pair of slit-like glands

at the base. Inflorescence axillary, cymose,

1, rarely 2-3-flowered; peduncle terete, 2-30

mm long, vestiture as for the stem; bracteoles

opposite to subopposite, herbaceous, narrowly

triangular, 2-6 mm long, acute, glabrous to

sparsely hairy, persistent; pedicels terete,

Austrobaileya 8(4): 699-723 (2012

slightly dilated upwards, 3-12 mm long in

flower, extending to 23 mm in fruit, glabrous

to moderately hairy, with glands below the

calyx. Outer sepals narrowly lanceolate to

narrowly ovate-elliptic, 8-16 mm long, 3.5-5.5

mm wide in flower, becoming wider in fruit,

apex acute, apiculate, attenuate to rounded

at base, with 3-5 prominent longitudinal

tuberculate nerves, reticulate in lower part,

glabrous; inner sepals ovate, ovate-lanceolate

or oblong-lanceolate, 10-16 mm long, 4.5-6

mm wide, apex acute, with a rolled apiculate

tip, base truncate with a broad hyaline

margin, glabrous. Corolla funnel-shaped

with a slender tube, 35-60 mm long, 30-50

mm diameter, white, lilac to mauve; petals

40-70 mm long, midpetaline band glabrous.

Stamens 5, included, attached to the base

of the corolla for 8-9 mm; filaments free for

13-17 mm, with multicellular hairs from just

below the point of attachment for 7-11 mm;

anthers oblong, sagittate, 2.8-3.5 mm long

with basal lobes 0.5-0.6 mm; pollen globular,

spinulose. Ovary 2-locular, glabrous; style

26-28 mm long; stigma biglobular. Capsule

globular, c. 10 mm long, 8-10 mm diameter,

valvate-dehiscent, glabrous; seeds 4, 4.5-5.5

mm long, dark brown, sparsely to moderately

densely puberulent, with a tuft of long whitish

hairs to 2 mm around the hilum.

Affinities: Ipomoea tolmerana resembles

I. graminea in the vegetative state, but its

leaves are always barely to distinctly hastate.

Its relationships; however, lie more closely

with the Australian representatives of the

I. gracilis group (Austin et al. 1993) with

which it shares a funnel-shaped corolla and

puberulent seeds with a tuft of longer hairs

around the hilum. From all of these related

species it can be distinguished by its long

narrow-linear, barely to distinctly hastate

leaves.

Key to the subspecies of I. tolmerana

1 Corolla white, petals 50-70 mm long; outer sepals 9-16 x 5-5.5 mm and inner

sepals 15-17 x 5-6 mm, at flowering.subsp. tolmerana

1. Corolla lilac to mauve, petals 40-50 mm long; outer sepals 8-12 x 5-5.5 mm and

inner sepals 11-13 x 6.5-7.5 mm, at flowering.subsp. occidentalis

Johnson, New species of Ipomoea

Etymology : Named after Mt Tolmer where

the initial collections were made. Since then,

the taxon has been commonly referred to

as “Ipomoea Tolmer” and “I. tolmeri” The

mountain was named in honour of Alexander

Tolmer (1815-1890), Commissioner of Police

in South Australia.

Ipomoea tolmerana R.W. Johnson subsp.

tolmerana

Ipomoea sp. Tolmer (C.R.Dunlop 6787);

Short et al. (2011).

Leaf blades 60-150 mm long, basal lobes to

50 mm long; inflorescence cymose with 1 or

2 flowers, rarely 3-flowered; peduncles 2-30

mm long; pedicels 3-12 mm long, glabrous

to moderately hairy; outer sepals 9-16 mm

long, 5-5.5 mm wide and inner sepals 15-17

mm long, 5-6 mm wide, at flowering; corolla

white, petals 50-70 mm long; seeds 5-5.5 mm

long.

Additional specimens examined: Northern Territory.

Narbarlek, Arnhem Land, Mar 1989, Hinz 447 (BRI,

DNA); Western Arnhem Land, c. 91 km ESE of Jabiru,

Mar 2000, Cowie 8678 (DNA); Walker Creek Ranger

Station, Mar 1995, Cowie 5341 & Taylor (DNA);

Litchfield N.R, road from Batchelor, Nov 1992, Leach

3356 (DNA); Tabletop Range, near Mt Tolmer, May 1985,

Dunlop 6787 (DNA); Litchfield N P., near Lost City,

Apr 1999, Cowie 8293 & Brennan (BRI, DNA); Round

Jungle, Kakadu, Feb 1989, Russell-Smith & Lucas 8005

(DNA); Litchfield N.P. south, track to Tableland Creek,

Feb 1996, Cowie 6193 & Booth (BRI, DNA); Marrawal

Plateau, near Bloomfield Springs, Feb 1996, Brennan

3208 (DNA); Umbawarra Gorge, Feb 1978, Gano s.n.

(DNA); Nitmiluk N.P., Site 566, Marrawal Plateau, Apr

2001, Brennan 5630 (DNA); Nitmiluk N.P., Apr 2001,

Michell & Boyce 3360 (DNA); Nitmiluk N.P, Site 324,

Mar 2001, Risler & Waetke 1429 (DNA); c. 29 miles

\c. 47 km] NE of Maranboy Police Station, Mar 1965,

Lazarides & Adams 18 (DNA).

Distribution and habitat: Ipomoea tolmerana

subsp. tolmerana occurs in northern Northern

Territory, from Litchfield N.R, eastward to

Narbalek and Maranboy (Map 3). It grows

on sandstone plateaux, on sandy soils, often

deep, in eucalypt open forest and woodland,

often with Eucalyptus tetrodonta F.Muell.

and E. miniata and a grassy under storey of

Sorghum , Themeda and Aristida and some

shrubs.

713

Phenology: Flowers have been recorded from

February to May, also in November; fruit

have been recorded from March to April, also

in November.

Conservation status: It is a widespread

species and not known to be at risk. It is not

listed among the threatened species in the

Northern Territory (Short et al. 2011).

Ipomoea tolmerana subsp. occidentalis

R.W.Johnson subspecies nova; a subspecie

typica corolla rosa usque ad malvina (in

subspecie typica alba) et petalis brevioribus,

40-50 mm longis (non 50-70 mm longis),

sepalis exterioribus sub anthesi brevioribus,

8-12 mm longis (non 9-16 mm longis) et

sepalis interioribus sub anthesi brevioribus et

latioribus, 11-13 x 6.5-7.5 mm (non 15-17 x

5-6 mm) differens. Typus: Western Australia.

1 km S of the roadhouse and 50 km N of

Broome on the Beagle Bay Track, Dampier

Peninsula, 27 March 2001, A.A.Mitchell 6669

(holo: BRI; iso: DNA, PERTH).

Ipomoea sp. A; Wheeler & Marchant (1992).

Ipomoea sp. A Kimberley Flora (L.J. Penn

84); Western Australian Herbarium (2011).

Leaf blades 60-120 mm long, basal lobes to

25 mm long; inflorescence cymose, 1-, rarely

2-flowered; peduncles 3-15(-25) mm long;

pedicels 8-14(-22) mm long, glabrous to

sparsely hairy; outer sepals 8-12 mm long,

5-5.5 mm wide and inner sepals 11-13 mm

long, 6.5-7.5 mm wide, at flowering; corolla

lilac to mauve, petals 40-50 mm long; seeds

4.5-5.5 mm long. Fig. 7.

Additional specimens examined: Western Australia.

Caffarelli Island, c. 300 km N of Derby, Apr 2000,

Mitchell 6123 (BRI, DNA, PERTH); King Hall Island,

Buccaneer Archipelago, Jun 1982, Hopkins BA0542

(PERTH); 81 km N from Willie Creek turn off on Beagle

Bay Road, Mar 1986, Foulkes 412 (BRI, DNA, PERTH).

Distribution and habitat: Ipomoea tolmerana

subsp. occidentalis occurs in Dampierland,

north-western Western Australia, north to the

Buccaneer Archipelago (Map 3). It grows on

pindan plains, in open savannah woodland on

sandy loam soils.

714 Austrobaileya 8 ( 4 ): 699-723 (2012

Fig. 7. Ipomoea tolmerana subsp. occidentalis. A. flowering branch *1. B-C. leaf variation xl. D. outer sepal at

flowering x4. E. inner sepal at flowering x4. A-E from Mitchell 6669 (PERTH - holotype). Del. W.Smith.

715

Johnson, New species of Ipomoea

Phenology : Flowers and fruits have been

recorded in March and April.

Notes: Ipomoea tolmerana subsp. occidentalis

differs from I. tolmerana subsp. tolmerana in

the colour of the corolla. Populations of blue

flowered species of Ipomoea occasionally

sport white variants. However, in this case,

flower colour in specimens of both the eastern

and western subspecies appears constant. It

also appears to have a smaller corolla and its

outer sepals at flowering are shorter and inner

sepals both shorter and wider than in the type

subspecies.

Conservation status: It is regarded as a

priority 1 taxon in Western Australia (Western

Australian Herbarium 2011).

Etymology: The specific epithet is derived

from the Latin occidentalis, meaning western.

This refers to the western distribution of this

subspecies.

Ipomoea versipellis R.W. Johnson species

nova; affinis I antonschmidii R.W. Johnson,

sed laminis ± glabris usque ad dense

piliferis (in ea semper dense piliferis),

vix usque ad valde hastatis (non semper

integris) et sepalis exterioribus sub anthesi

longioribus, 10—16 mm longis (non 16-21

mm longis). Planta perennis, caules serpentes

or volubiles. Folia simplicia, lamina ovata,

hastata usque ad sagittata. Inflorescentia

cymosa, 1-3-flora. Sepala exteriora laevia.

Corolla infundibuliformis glabra. Semina

pubescentes, caespite pilorum longiorum ad

hilum. Typus: Northern Territory. Darwin &

Gulf: Keep River National Park, Spirit Hills,

c. 17 km NNW of Bullo River Homestead,

22 March 2009, 1.D.Cowie 12288 (holo: DNA

[D191134]; iso: BRI, CANB n.v., MEL n.v.,

MO n.v., PERTH n.v).

Ipomoea sp. B; Wheeler & Marchant (1992).

Ipomoea sp. OT Station (S.T.Blake 17676);

Short etal. (2011).

Perennial with trailing and scrambling stems;

stems terete, stout, herbaceous, becoming

somewhat woody, very sparsely to densely

hairy, hairs stout, tubercle-based, straight,

curved to sinuate, ± erect, 0.3-1.2 mm long.

Leaves simple, petiolate; petiole 10-50

mm long, 0.3-1 times as long as the blade,

vestiture as for the stem; blade broadly ovate

or ovate-oblong, barely to distinctly hastate to

sagittate, 20-70 mm long, 10-50 mm wide,

apex obtuse to rounded, rarely barely acute,

mucronate, base subcordate to cordate with

sinus up to 40% of the blade, ± glabrous to

densely hairy above, sparsely (hairs mainly

on major veins) to densely hairy below,

midrib with glands at the base and 4-7 pairs

of secondary veins. Inflorescence axillary,

cymose, l-5(-10)-flowered; peduncle terete,

10-60 mm long, vestiture as for the stem;

bracteoles opposite, herbaceous, narrow-

linear to linear-subulate, 4-10(-20) mm long,

0.5-1 mm wide, acute, vestiture as for leaves,

persistent; pedicels terete, 3—15(—23) mm

long, vestiture as for the stem, with distinct

slit-like glands 0.5-1 mm long under the calyx.

Outer sepals narrowly ovate, ovate to ovate-

elliptic, apiculate, 9-16 mm long, 3.5-5.5

mm wide in flower, with a length: width ratio

of 2.5-3.5, extending to 8 mm wide in fruit,

apex acute, with 5 raised ribs, moderately to

densely hairy; inner sepals ovate to broadly

ovate, acuminate, 10-15 mm long, 5.5-6.5

mm wide in flower, broader in fruit, apex

rolled, acute, base truncate, with a broad

hyaline margin, especially at the base, with

3-5 veins, glabrous or with hairs on the veins.

Corolla funnel-shaped, 30-40 mm long, 30-

40 mm diameter, deep pink to purple; petals

40-65 mm long, midpetaline band glabrous.

Stamens 5, included, attached to the base of

the corolla for 7-9 mm; filaments free for

9-17 mm, with multicellular hairs from just

below the point of attachment upwards for

6-9 mm; anthers oblong, sagittate, 3-4 mm

long, c. 1 mm wide, with basal lobes 0.5-0.75

mm long; pollen globular, spinulose. Ovary

2-locular on a prominent disk, glabrous; style

20-24 mm long; stigma biglobular. Capsule

globular, c. 11 mm long, with a persistent

style base, valvate-dehiscent, glabrous; seeds

4, 4.5-5.5 mm long, dark brown to black,

densely puberulent with a tuft of long whitish

brown hairs to 2 mm around the hilum. Fig. 8.

Additional specimens examined: Western Australia.

Durack River, c. 80 km SSW of Wyndham, May 1976,

Beauglehole 51440 (BRI, PERTH); Canyon Creek,

near meatworks on Packsaddle Plain, Kununurra, Mar

1993, Mitchell 2824 (BRI, PERTH); Lake Kununurra

716

Austrobaileya 8(4): 699-723 (2012

Fig. 8. Ipomoea versipellis. A. cauline leaf xl. B. cauline leaf *0.5. C. cauline leaf xl. D. outer sepal at flowering x4.

E. inner sepal at flowering x4. A-B from Cowie 12288 (DNA - holotype); C from Brennan 4404 (DNA); D-E from

Mitchell 2824 (BRI). Del. W. Smith.

(Ord River), upper reaches of Noogoora Burr Creek,

Nov 2001, Handasyde TH01 301, 301b (BRI, PERTH).

Northern Territory. Upper Fitzmaurice River, Mar

1989, Leach & Dunlop 2458 (BRI, DNA); Yamburran

Range, Mar 1989, Leach & Dunlop 2470 (BRI, DNA);

Yamburran Range, Mar 1989, Russell-Smith & Lucas

7682 (DNA); Bradshaw Station, near fire plot 3, Feb 1999,

Michell 2181 (DNA); Spirit Hills Conservation Reserve,

Mar 2006, Dixon 1598 (BRI, DNA); Spirit Hills, c. 18

km N of Bullo River Homestead, Apr 2007, Kerrigan

1197 (DNA); Bullo River Station, towards western

boundary, Bullo 29, Mar 2006, Lewis 499 (DNA); 45

km SSW of Legune Station, Mar 1989, Russell-Smith &

Brock 7565 (BRI, DNA); Keep River N.P., Spirit Hills

area, c. 35 km SW of Bullo River Homestead, Mar 2009,

Cowie 12309 (BRI, DNA); Gregory N.P, headwater

of tributary of East Baines River, c. 46 km due SW of

Timber Creek, Apr 1996, Walsh 4357 (DNA, MEL);

Gregory N.P, Jasper Creek, Apr 1996, Woodward 12 &

Booth (DNA); Jasper Gorge area, 11 km SW of gorge,

Apr 2000, Brennan 4404 (DNA); Gregory N.P, Snake

Creek, Apr 1996, Booth & Jones 1673 (BRI, DNA);

Gregory N.P, near top of falls, upstream on creek, c. 45

km SSW of Bullita Outstation, Apr 1996, Duretto 1119

Johnson, New species of Ipomoea

& Davies (DNA); O.T. Station, May 1947, Blake 17676

(BRI, DNA); Gregory N.P., Depot Creek, c. 27 km ENE

of Limbunya, Apr 1996, Cowie & Jones 6242 (BRI,

DNA, PERTH).

Distribution and habitat: Ipomoea

versipellis occurs in the Drysdale River

area of the Kimberley, Western Australia,

extending into the Northern Territory, as

far north as Litchfield N.R, as far south

as Gregory N.R and eastward to the O.T.

Station, SW of Borroloola (Map 3). It grows

in a variety of vegetation communities from

eucalypt woodlands to riverine forests and

vine thickets on sandy soils on rocky slopes

in dissected sandstone country.

Phenology : Flowers have been recorded from

February to May and also in November; fruits

have been recorded from February to May,

717

also in August and November.

Affinities: Ipomoea versipellis is closely

related to I. antonschmidii which occurs to the

east in north-western Queensland. However,

the latter species is always densely hairy and

its leaves are never hastate. It also has longer

sepals.

Conservation status: It is a widespread

species and not known to be at risk.

Etymology: The specific epithet is from the

Latin versipellis, meaning shape-changing,

capable of altering its appearance. This refers

to the variation that occurs in leaf shape

from ± ovate-triangular to deeply hastate

and indumentum from densely hairy to very

sparsely hairy.

Key to species of native and naturalised* Ipomoea found in Australia; endemic Australian

species indicated®

1 Leaves pinnately cut into numerous linear segments; corolla scarlet,

salver-shaped (NSW, NT, Qld, WA).*1. quamoclit

1. Leaves not as above, or if so, then corolla not scarlet and salver-shaped.2

2 Outer sepals with awn >2 mm long, at or below the apex; corolla

salver-shaped.3

2. Outer sepals without a long awn; corolla funnel-shaped or campanulate,

rarely salver-shaped.4

3 Corolla small, scarlet, <5 cm long, limb <3 cm across (NSW, NT, Qld,

WA) *1. hederifolia

3. Corolla large, white, >5 cm long, limb >6 cm across (NSW, Qld).*1. alba

4 Corolla small, <1.5 cm long, rarely longer; sepals fimbriate; peduncles <2

cm long or absent.5

4. Corolla >1.5 cm long or if less, then sepals not fimbriate; peduncles

usually > 2mm long.9

5 Herbs with erect or decumbent stems; corolla pink to purple, never white,

mid-petaline bands glabrous (NSW, NT, Qld, SA, WA).I. polymorpha

5. Annual or herbaceous perennial vines with stems twining, climbing or

trailing; corolla pink, purple or white, mid-petaline bands hairy.6

6 Ovary and capsule hairy; corolla pink to purple (NT, Qld, WA).I. eriocarpa

6. Ovary and capsule glabrous; corolla white.7

7 Leaf base attenuate, rounded to truncate, never cordate (NSW, NT, Qld,

SA, WA) I. lonchophylla®

7. Leaf base cordate.8

8 Leaves glabrous or very sparsely hairy on upper surface, moderately

pilose on lower surface (arid NT, Qld, SA, WA).I. racemigera®

718 Austrobaileya 8(4): 699-723 (2012

8. Leaves evenly pilose on both surfaces (NSW, NT, Qld, WA).I. plebeia

9 Ovary and capsule 3-locular.10

9. Ovary and capsule 2 or 4-locular.15

10 Leaves digitate with 3-5 lobes, lobes coarsely and irregularly toothed to pinnatifid ... 11

10. Leaves entire or 3-5 lobed, not digitate, lobes entire.12

11 Corolla <1.5 cm long; sepals 2.8-5 mm long in flower (NT, Qld, WA).I. coptica

11. Corolla >4 cm long; sepals 6-14 mm long in flower (NT, Qld).I. diversifolia

12 Sepals <15 mm long, apex acute (NSW, NT, Qld).*1. purpurea

12. Sepals mostly >15 mm long with apex long attenuate to acuminate .13

13 Sepals with ±appressed hairs <1 mm long; corolla limb >5 cm diameter;

flowers in cymose clusters on long peduncles (NSW, Qld, SA, Vic, WA).... *1. indica

13. Sepals with ascending to spreading hairs >1 mm long; corolla limb <5 cm

diameter; flowers usually solitary or in 3’s.14

14 Sepals abruptly narrowed with the long subacute tips strongly spreading

or curved, hairs spreading, 3-4 mm long (Qld).*1. hederacea

14. Sepals gradually narrowed with the long acute tips suberect, straight,

scarcely spreading, hairs ascending to spreading < 3 mm long (NT, Qld, WA)... *1. nil

15 Leaves palmately divided to the base or almost so.16

15. Leaves entire or bearing only basal lobes, or if palmately lobed, then

lobes cut less than two thirds of distance to base of blade.17

16 Leaves and stems patently hairy (coastal NT, WA).*1. pes-tigridis

16. Leaves and stems glabrous (NSW, Qld, SA, WA).*1. cairica

17 Leaves narrow-linear, occasionally distinctly hastate, with a length:width

ratio of 6-45; corolla white or lilac to mauve.18

17. Leaves not linear, with a length:width ratio <5, if narrow-linear and

hastate then corolla pink to purple.19

18 Leaves entire; outer sepals much shorter than inner, surface smooth;

corolla salver-shaped (NT, Qld, WA).I. graminea E

18. Leaves barely to distinctly hastate; sepals equal in length, longitudinal

veins rugose; corolla funnel-shaped (NT, WA).I. tolmerana E

19 Corolla salver-shaped, white, rarely pale pink, >8 cm long; sepals mainly

>20 mm long in fruit.20

19. Corolla funnel-shaped to campanulate, pink or purple, rarely white or

yellow, mainly <8 cm long; sepals mainly <20 mm long in fruit.23

20 Stamens inserted near mouth of corolla tube (Qld).I. aculeata

20. Stamens inserted near the base of the corolla tube.21

21 Leaves, sepals and mid-petaline bands of corolla hairy (Qld).I. samtronanensis E

21. Leaves, sepals and mid-petaline bands of corolla glabrous.22

22 Leaves mostly 3-5 lobed; stamens exserted; outer sepals much shorter

than inner sepals (WA).I. trichosperma

22. Leaves mostly entire; stamens included; outer sepals equal or slightly

shorter than inner sepals (NT, Qld, WA).I. violacea (syn. I. macrantha)

23 Marsh or aquatic plants; stems thick, hollow, rooting at nodes; capsules

Johnson, New species of Ipomoea 719

indehiscent or dehiscing irregularly.24

23. Rarely in marshy situations; stems herbaceous or woody; capsules valvate-dehiscent . . 25

24 Corolla creamy-white, 1.5-2.5 cm long (NSW, NT, Qld, SA, WA) ... I. diamantmensis E

24. Corolla pink or pale lilac, often with a purple centre, rarely white, 3-6 cm

long (NT, Qld, WA).I. aquatica

25 Leaves strongly emarginate to bilobed; stems trailing, rooting at the

nodes; plants fleshy; growing on coastal dunes.26

25. Leaves not strongly emarginate to bilobed.27

26 Corolla pink or purple; peduncles 3-16 cm long (NSW, NT, Qld, WA) .... I. pes-caprae

26. Corolla white or yellowish with a purple centre; peduncles <2 cm long

(NT, Qld).I. imperati

27 Sepals distinctly fimbriate; ovary usually hairy.28

27. Sepals glabrous, occasionally hairy, not distinctly fimbriate; ovary mainly glabrous ... 29

28 Corolla 3-4 cm long (Qld, WA, also cultivated).*1. batatas

28. Corolla 1.5-2 cm long (NT, Qld, WA).*1. triloba

29 Leaves palmatifid with 5-7 lobes (Qld).I. mauritiana

29. Leaves entire or bearing only basal lobes.30

30 Corolla yellow, if whitish then corolla <5 cm long with a dark purple centre.31

30. Corolla pink or purple, rarely whitish, if whitish then corolla >5 cm long.34

31 Leaves fleshy, leaves ovate to oblong, apex obtuse to emarginated; outer

sepals >8 mm long (NT, Qld).I. imperati

31. Leaves herbaceous, ovate to broadly ovate, acuminate, acute; outer sepals

3-6 mm long.32

32 Corolla <1.5 cm long, yellow; sepals with some stout hairs 0.5-1.5 mm

long, rarely glabrous (Qld).I. brownii E

32. Corolla >2 cm long, rarely less, yellow to whitish, with a dark purple

centre; sepals glabrous or sparsely pubescent with hairs <0.5 mm long.33

33 Corolla 2-2.5 cm long, pale yellow to whitish (Qld).*1. obscura

33. Corolla 27-5.5 cm long, yellow (Qld).*1. ochracea

34 Leaves glabrous or sparsely hairy.35

34. Leaves moderately to densely hairy.54

35 Sepals ovate, length:width ratio >2, apex acute to narrowly obtuse.36

35. Sepals broadly ovate to orbicular, length: width ratio <2, apex broadly obtuse to rounded 46

36 Seeds puberulent usually with a tuft of hairs around the hilum.37

36. Seeds with longer hairs on the margins, sometimes glabrous.44

37 Peduncle equal or shorter than the pedicels, mainly <1.5 cm long,.38

37. Peduncle longer than the pedicels, mainly >1.5 cm long.40

38 Leaves strongly hastate with narrow-linear terminal lobe and recurved

linear to linear-triangular basal lobes; stems fibrous, becoming woody

(Qld).I. fimicularis E

38. Leaves broadly ovate-triangular to deltoid, ovate to broad-ovate or

ovate-oblong, truncate, sagittate to deeply cordate, rarely hastate at

base; stems herbaceous.39

720 Austrobaileya 8(4): 699-723 (2012

39 Outer sepals bearing 3-5 raised, tuberculate, longitudinal veins (NT,

Qld, WA).I. gracilis*

39. Outer sepals with a ± smooth surface, if veins raised then ± smooth

(WA).I. kalumburu E

40 Corolla sky blue with a paler throat; sepals <7 mm long; seeds lacking

tuft of hairs around hilum (Qld).*1. tricolor

40. Corolla pink to purple, occasionally white, usually with a darker centre;

sepals >7 mm long; seeds with a tuft of long hairs around the hilum.41

41 Outer sepals with a ±smooth surface, if veins raised then ±smooth (WA). . I. kalumburu E

41. Outer sepals bearing 3-5 raised, tuberculate, longitudinal veins.42

42 Leaves broad-ovate to ±reniform, cordate at the base (NT, Qld, WA).I. argillicola E

42. Leaves narrow-ovate to oblong-lanceolate, hastate to sagittate at the base.43

43 Stems, petioles and sepals at flowering with scattered stout hairs (NT, WA) . I. versipellis E

43. Stems, petioles and sepals at flowering glabrous or rarely with occasional

weak, not stout hairs and never on sepals (NT, Qld, WA).I. limosa E

44 Seeds glabrous or with hairs on the margins (Qld).I. tiliacea

44. Seeds villous.45

45 Corolla <4 cm long; sepals at flowering <11 mm long; leaves entire (NT,

Qld, SA, WA).I. muelleri*

45. Corolla >4 cm long; sepals at flowering >11 mm long; leaves usually with

two rounded hastate lobes (NT, Qld).I. brassii E

46 Plant herbaceous with trailing stems, often rooting at the nodes, sometimes

twining at the tips.47

46. Shrub or liana.49

47 Peduncle <2 cm long; corolla 3-4.5 cm long; sepals 6-9 mm long; seeds

glabrous (Qld).I. littoralis

47. Peduncle >2 cm long; corolla 5-9 cm long; sepals 7.5-25 mm long; seeds

glabrous to villous.48

48 Corolla pink to purple; bracts 2-7 mm long, persistent; seeds glabrous to

sparsely pubescent (NT, Qld).I. polpha E

48. Corolla white with a red-purple centre; bracts >10 mm long, caducous;

seeds villous with longer hairs on margins and around the hilum (SA) . . . *1. pandurata

49 Peduncles >3 cm long, at least 3 times as long as pedicel.50

49. Peduncles mainly <3 cm long, never 3 times as long as pedicel.51

50 Erect shrubs; corolla with pubescent mid-petaline band (NT, Qld, SA,

WA).*1. carnea

50. Liana; corolla glabrous (Qld).I. tiliacea

51 Sepals equal to sub-equal, mainly <13 mm long at flowering (NT, Qld,

WA).52

51. Inner sepals longer than outer, 12-22 mm long at flowering.53

52 Fruiting calyx equalling and enclosing the capsule, 11-55 mm long,

strongly wrinkled, capsule with prominent style base.I. abrupta E

52. Fruiting calyx not enclosing the capsule and not or barely wrinkled,

capsule without prominent style base.I. dunlopii E

Johnson, New species of Ipomoea 721

53 Leaves broad-ovate, orbicular or reniform, base cordate (NT, Qld, WA).I. costata E

53. Leaves lanceolate, usually rounded-hastate or sagittate, occasionally

deeply lobed at the base (Qld, WA).I. calobra E

54 Corolla with tomentose mid-petalline bands.55

54. Corolla glabrous.56

55 Shrub, stems not twining; sepals 8-11 mm long; bracts 2-3 mm long

(WA).I. yardiensis E

55. Stems twining; sepals >15 mm long; bracts 10-15 mm long (NT, WA) ... I. bracteolata E

56 Sepals at flowering broadly ovate, oblong or orbicular, obtuse, outer surface smooth . . 57

56. Sepals at flowering ovate, acuminate to lanceolate, acute, surface with 5

raised, longitudinal veins.58

57 Sepals at flowering to 11 mm long, if longer then sepals sparsely hairy;

leaves acute to narrowly obtuse (NT, Qld, WA).I. abrupta E

57. Sepals at flowering 8-10 mm long, moderately to densely hairy; leaves

obtuse to rounded (Qld).I. densivestita E

58 Leaves not lobed at the base; sepals 18-21 mm long (NT, Qld).I. antonschmidii E

58. Leaves usually hastate to sagittate at the base; sepals <16 mm long (NT,

WA).I. versipellis E

Acknowledgements

I appreciate the continuing support provided

by Gordon Guymer, Director of BRI, which

enables me to continue my taxonomic research.

I thank Will Smith (BRI) for the illustrations,

Hans Dillewaard and Jenny Calway for their

technical assistance and David Halford for his

contributions while preparing the account for

the Flora of Australia.

References

Austin, D.F. & Hauman, Z. (1996). A synopsis of

Ipomoea (Convolvulaceae) in the Americas.

Taxon 45: 3-38.

Austin, D.F. Jarrett, R.L. & Johnson, R.W. (1993).

Ipomoea gracilis R.Brown (Convolvulaceae)

and its allies. Bulletin of the Torrey Botanical

Club 120: 49-59.

Bentham, G. (1869). Ipomoea. In Flora Australiensis 4:

412-428. L.Reeve & Co.: London.

Johnson, R.W. (2007). Convolvulaceae. In RD. Bostock

& A.E. Holland (eds.), Census of the Queensland

Flora 2007, pp. 51—53. Queensland Herbarium,

Environmental Protection Agency: Brisbane.

- (2010). Convolvulaceae. In P.D. Bostock &

A.E. Holland (eds.). Census of the Queensland

Flora 2010, pp. 46-48. Queensland Herbarium,

Department of Environment & Resource

Management: Brisbane.

Short, PS., Albrecht, D.E., Cowie, I.D., Lewis, D.L.

& Stuckey, B.M. (eds.) (2011). Checklist of

the Vascular Plants of the Northern Territory.

Northern Territory Herbarium, Department of

Natural Resources, Environment, The Arts and

Sport: Darwin.

Western Australian Herbarium (2011). FloraBase —

The Western Australian Flora. Department

of Environment and Conservation, http://

florabase.dec.wa.gov.au/

Wheeler, J.R. & Marchant, N.G. (1992).

Convolvulaceae. In J.R.Wheeler et al. (ed).

Flora of the Kimberley Region, pp. 737-759.

Western Australian Herbarium: Perth.

722

Austrobaileya 8(4): 699-723 (2012

Map 1. Distribution of Ipomoea bracteolata O, Z funicularis □, I. kalumburu A.

Map 2. Distribution of Ipomoea densivestita O, Z dunlopii □, Z limosa A.

Johnson, New species of Ipomoea

723

Map 3. Distribution of Ipomoea tolmerana subsp. occidentalis O, I. tolmerana subsp. tolmerana □, I. versipellis A.

Referees consulted for Volume 8 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

D.F.Austin

R. Barrett

A.R.Bean

P.Berry

J.J.Bruhl

I.D.Cowie

D.L.Jones

M.F.Duretto

A.J.Ford

G.RGuymer

M.T.Mathieson

D.Nicolle

S .Razafimandimbison

R.Riina

K. Shepherd

P.S. Short

G.W. Staples

K.Thiele

I. R.Thompson

J. Veldkamp

N.G.Walsh

B.Wannan

J.Wege

Peter G.Wilson

F.Zich

Index to Austrobaileya - volume 8

New names are in Bold

Adenostemma lavenia 662

Amaracarpus 155

Amaracarpus corymbosus 156

Amaracarpus heteropus 156

Amaracarpus lanceolatus 156

Amaracarpus longifolius 156

Amaracarpus nematopodus 156, 157

Amaracarpus urophyllus 156

Amaranthaceae 267

Ardisia 2

Ardisia bakeri 3, 5, 23

Ardisia bifaria 19

Ardisia brevipedata 3, 4, 7, 23

Ardisia brevipedata var. depauperata 11, 223

Ardisia brevipedata var. brevipedata 4

Ardisia colorata 18

Ardisia crenata 3, 6, 8, 9, 23

Ardisia crispa 8

Ardisia depauperata 3, 10, 11, 23, 223

Ardisia eliiptica 3, 12, 13, 14, 23

Ardisia fasciculata 3, 14, 15

Ardisia humilis 14

Ardisia hylandii 223

Ardisia pachyrrhachis 3, 16, 17, 23

Ardisia pseudojambosa 21

Ardisia racemosa 3

Ardisia repandula 21

Ardisia sanguinolenta 3, 18

Ardisia solanacea 14

Ardisia subgenus Tetrardisia 18

Arthragrostis aristispicula 190

Arthragrostis brassiana 188

Arthragrostis brassiana var. brassiana

188, 189, 190

Arthragrostis brassiana var. minutiflora

188, 189, 190

Arthragrostis clarksoniana 190

Arthragrostis deschampsioides 189, 190

Asteraceae 97, 103, 280, 340, 653

Austromyrtus alaternoides 180

Austromyrtus aneityensis 181

Austromyrtus aphthosa 180

Austromyrtus cataractarum 181

Austromyrtus clusioides 180

Austromyrtus conspicua 180

Austromyrtus kanalaensis 179

Austromyrtus nigripes 181

Austromyrtus pancheri 181

Austromyrtus prolixa 181

Austromyrtus vieillardii 181

autochory 624

Balingayum decumbens 693

Bean, A.R. (2009). Homoranthus tricolor

(Myrtaceae), a new species from south¬

eastern Queensland 77

Bean, A.R. (2009). Reinstatement of Enydra

woollsii F.Muell. (Asteraceae:

Heliantheae) 103

Bean, A.R. (2009). Taxonomic and

nomenclatural notes on the Eastern

grey boxes (Eucalyptus ser. Moluccanae

Chippendale, Myrtaceae) and the

reinstatement of Eucalyptus woollsiana

R.T. Baker 25

Bean, A.R. (2009). The identity of Centaurea

riparia DC. (Asteraceae) 97

Bean, A.R. (2010). Anew subspecies of

Eucalyptus sideroxylon A.Cunn. ex Woolls

(Myrtaceae) from Queensland 139

Bean, A.R. (2010). Crudia abbreviata

A.R.Bean (Caesalpiniaceae) a new

species from Cape York Peninsula

Queensland 151

Bean, A.R. (2010). Four new species of

Stylidium Sw. (Stylidiaceae) from

northern Australia 107

Bean, A.R. (2010). Two new species of

Solanum L. (Solanaceae) from central

Queensland 165

Bean, A.R. (2011). Two new species of

Pluchea Cass. (Asteraceae: Plucheinae)

from Queensland, Australia 340

Bean, A.R. (2011). A taxonomic revision of

Pterocaulon section Monenteles (Labill.)

Kuntze (Asteraceae: Inuleae-Plucheinae)

280

Bean, A.R. (2011). Hydrocharis dubia

(Blume) Backer (Hydrocharitaceae) is

an alien species in Australia 435

Bean, A.R. (2011). New and reinstated

species of the Solanum ellipticum R.Br.

(Solanaceae) species group 412

Bean, A.R. & Mathieson, M.T. (2012).

Stylidium elachophyllum A.R.Bean &

M.T.Mathieson (Stylidiaceae), a new

species from northern Queensland 608

Bean, A.R. & Miller, C.H. (2011).

A taxonomic revision of Nyssanthes

R.Br. (Amaranthaceae) 267

Bladhia 2

Bladhia brevipedata 4

Blainvillea 653, 655

Blainvillea acmella 656, 662, 664, 669

Blainvillea alba 663

Blainvillea amazonica 666

Blainvillea biaristata 666

Blainvillea brasiliensis 666

Blainvillea calcicola 656, 658, 669

Blainvillea cunninghammi 656, 660,

661,669

Blainvillea dallo-vedovae 667

Blainvillea dichotoma 662, 667

Blainvillea dubia 660

Blainvillea gayana 656, 657, 669

Blainvillea hispida 663

Blainvillea lanceolata 667

Blainvillea latifolia 662

Blainvillea polycephala 666

Blainvillea prieureana 656

Blainvillea racemosa 666

Blainvillea rhomboidea 662

Blainvillea rhomboidea var. racemosa 666

Blainvillea synedrelloides 666

Blainvillea tampicana 666

Blainvillea tenuicaulis 666

Booth, R., Moore, D J. & Hodgon, J. (2009).

Four new species of Cyperus L.

(Cyperaceae) from northern Queensland

Boraginaceae 335

Brachiaria notochthona 216

Brachiaria occidentalis 215

Brachiaria occidentalis var. ciliata 216

Bryobium intermedium 697

Bryonia cucumeroides 371

Cadetia clausa 123

Cadetia collinsii 123

Cadetia maideniana 123

Cadetia taylori 123, 124

Cadetia uniflos 119, 120, 122, 123

Cadetia wariana 123

Caesalpiniaceae 151

Calogyne chinensis 693

Calogyne heteroptera 693

Calyptocarpus bahiensis 666

Calyptocarpus biaristatus 666

Calyptocarpus brasiliensis 666

Calyptocarpus megapotamica 666

Calyptochloa 636

Calyptochloa cylindrosperma 637, 644,

646, 647, 648, 652

Calyptochloa gracillima 637, 638

Calyptochloa gracillima subsp. gracillima

638, 639, 640

Calyptochloa gracillima subsp. ipsviciensis

641,642, 643

Calyptochloa johnsoniana 645, 648, 650,

651,652

Calyptochloa sp. (Blackjack E.J.Thompson+

CHA769) 644

Cenchrus advena 191

Cenchrus americanus 191

Cenchrus basedowii 191

Cenchrus brevisetosus 190, 192

Cenchrus ciliaris 191

Cenchrus clandestinus 191

Cenchrus elymoides 190, 191

Cenchrus elymoides var. brevisetosus 192

Cenchrus longisetus 191

Cenchrus macrourus 191

Cenchrus pedicellatus 191

Cenchrus pedicellatus subsp. unispiculus 191

Cenchrus pennisetiformis 191

Cenchrus polystachios 191

Cenchrus purpurascens 191

Cenchrus purpureus 191

Cenchrus setaceus 191

Cenchrus setigerus 191

Cenchrus spinifex 192

Cenchrus thunbergii 191

Centratherum australianum 97

Centratherum punctatum subsp.

australianum 97

Centratherum riparium 97

Ceratocephalus acmella 662

Chamaeraphis minuta 214

Chamaesyce 446

Chamaesyce alsiniflora 521

Chamaesyce atoto 580

Chamaesyce australis 463

Chamaesyce biconvexa 469

Chamaesyce bifida 470

Chamaesyce carissoides 475

Chamaesyce centralis 476

Chamaesyce coghlanii 479

Chamaesyce dallachyana 481

Chamaesyce drummondii 484

Chamaesyce filipes 521

Chamaesyce hirta 497, 585

Chamaesyce hyssopifolia 500

Chamaesyce inappendicidata 501

Chamaesyce maculata 516

Chamaesyce micradenia 470

Chamaesyce mitchelliana 518

Chamaesyce mitchelliana var. hirta 519

Chamaesyce myrtoides 527

Chamaesyce obliqua 528

Chamaesyce ophiolitica 530

Chamaesyce ophthalmica 532

Chamaesyce pallens 534

Chamaesyce petala 542

Chamaesyce prostrata 548, 587, 592

Chamaesyce psammogeton 550

Chamaesyce schultzii 555

Chamaesyce serpens 559

Chamaesyce sp. (Pathungra A.Gunness

AG2118) 536

Chamaesyce sp. A 540

Chamaesyce sp. B 503

Chamaesyce sp. Marree (F.J.Badman 776)

503

Chamaesyce snpina 516

Chamaesyce thymifolia 566

Chamaesyce vachellii 470

Chamaesyce w heeler i 577

Citrus australasica 134

Citrus garrawayi 134

Citrus wakonai 134, 135

Coelospermum dasylobum 74

Coelospermum paniculatum 74

Coelospermum purpureum 70, 72, 74

Convolvulaceae 47, 55, 171, 699

Convolvulus brownii 6 1

Convolvulus caespitosus 57

Convolvulus cinerascens 57

Convolvulus cymosus 62

Convolvulus dissectus 57

Convolvulus gemellus 57

Convolvulus hirtus 57

Convolvulus incisa 57

Convolvulus peltatus 61

Convolvulus pentaphyllus 57

Convolvulus quinatus 6 1

Convolvulus umbellatus 62

Conyza redolens 306

Cooper, W.E. & de Boer, H.J. (2011). A

taxonomic revision of Trichosanthes L.

(Cucurbitaceae) in Australia including

one new species from the Northern

Territory. 364

Cooper, W.E. & Ford, A. J. (2010).

Trichosanthes odontosperma W.E.Cooper

& A.J.Ford (Cucurbitaceae) a new species

from Queensland’s Wet Tropics 125

Crotalaria inaequalis 65, 67, 68

Croton capitis-york 147, 148

Croton dichromifolius 144, 145, 147, 148

Croton insularis 621, 623

Croton lucens 617, 618, 619, 620, 621,

622, 623

Croton mamillatus 621, 623

Croton phebalioides 621, 623

Croton simulans 146, 147, 148

Croton stigmatosus 621, 623

Croton stockeri 147, 148

Crudia abbreviata 151, 152, 153, 154

Crudia papuana 152

Crudia sp. (Archer River BH 3078RFK) 151

Cucumeroides 366, 371

Cucurbitaceae 125, 364

Cunoniaceae 252

Cycadaceae 356

Cycas media 361

Cycas ophiolitica 361

Cycas terry ana 358, 359, 360, 361, 362, 363

Cyperaceae 35

Cyperus alaticaulis 37, 40, 41, 46

Cyperus eboracensis 37, 40, 41, 46

Cyperus montis-sellae 38

Cyperus multispiceus 37, 42, 44, 46

Cyperuspedunculosus 35, 37, 38

Cyperus peduncidosus var. alatus 38

Cyperus peduncidosus var. atrocastaneus38

Cyperus pedunculosus var. floribundus 38

Cyperus pedunculosus var. longebracteatus

Cyperus pedunculosus var. pedunculosus 38

Cyperus sharpei 37, 43, 44, 46

Cyperus sp. (Cape York J.R.Clarkson +8126)

Cyperus sp. (Chester River J.R.Clarkson

2392)40

Cyperus sp. (Herberton PR. Sharpe 1449) 43

Cyperus sp. (The Boulders J.A.Elsol 818) 42

Cyrtococcum accrescens 193

Cyrtococcum brevisetosus 190

Cyrtococcum capitis-york\93

Cyrtococcum oxyphyllum 193

Cyrtococcum patens 193

Davenportia 171

Davenportia davenportii 172, 173, 174, 176

de Boer, H.J. with Cooper, W.E. (2011). A

taxonomic revision of Trichosanthes L.

(Cucurbitaceae) in Australia including one

new species from the Northern Territory

364

Dendrobium baseyanum 697

Dendrobium foederatum 697

Dendrobium masonii 696

Dendrobium uniflos 120

Digitaria basaltica 194, 195, 198

Digitaria breviglumis 199, 200, 201

Digitaria cowiei 194, 195, 198

Digitaria diminuta 199, 200, 202

Digitaria dolleryi 195, 196, 198

Digitaria fumida 199, 204, 205

Digitaria orbata 199, 202, 203

Digitaria sharpeana 196, 197, 198

Digitaria sp. (Amungee Mungee Stn

I.D.Cowie+ 1752) 194

Digitaria sp. (Chesterton NP C.Dollery 354)

196

Digitaria sp. (Great Basalt Wall R. J.Fensham

2183) 194

Digitaria sp. (Mt Mulligan J.R.Clarkson

5821)200

Digitaria sp. (Sunnybank S.T.Blake 5300)

196

Digitaria sp. (Surat S.T.Blake 21286) 198

Digitaria veldkampiana 197, 198

Diplocaulobium masonii 696

Diplochory 624

Dockrill 696

Dockrillia x foederata 697

Dockrillia baseyana 697

Dockrillia calamiformis 697

Duperreya commixta 48, 50, 54

Duperreya 47

Duperreya halfordii 48, 50, 51, 52, 54

Duperreya sericea 48, 49, 50, 54

Duretto, M.F. & Janes, J.K. (2010).

Three new combinations in Pterostylis

(Orchidaceae) 221

Eclipta latifolia 662

Ecliptinae 653

Eisenmannia 655

Embothrieae 671

Entolasia minutifolia 204, 206

Entolasia sp. (Miles S.T.Blake 7709) 204

Entolasia sp. A 204

Enydra fluctuans 104

Enydra woollsii 103, 104

Episyzygium oahuense 178

Eria dischorensis 698

Eria intermedia 697

Eucalyptus albens 27, 30

Eucalyptus hemiphloia 31

Eucalyptus hemiphloia var. albens 30

Eucalyptus hemiphloia var. microcarpa 32

Eucalyptus melanophloia 347, 348

Eucalyptus melanophloia subsp.

melanophloia 349, 352

Eucalyptus melanophloia subsp. nana x E.

normantonensis hybrid 354, 355

Eucalyptus melanophloia subsp. nana 350,

351,352, 353, 354

Eucalyptus microcarpa 27, 32, 33

Eucalyptus moluccana 27, 31, 32

Eucalyptus odorata var. woollsiana 27

Eucalyptus pilligaensis 27

Eucalyptus section Adnataria 347

Eucalyptus series Moluccanae 26

Eucalyptus series Siderophloiae 347

Eucalyptus sideroxylon 139

Eucalyptus sideroxylon subsp. (Waaje

N.B.Byrnes 3955) 139

Eucalyptus sideroxylon subsp. improcera

139, 140

Eucalyptus subgenus Symphyomyrtus 347

Eucalyptus subsection Apicales 347

Eucalyptus subseries Jugatae 347

Eucalyptus woollsiana 27, 28

Eugenia aphthosa 180

Eugenia bifaria 183

Eugenia cataractarum 181

Eugenia clusioides 180

Eugenia colnettiana 180

Eugenia diversifolia 180

Eugenia heckelii 181

Eugenia kanalaensis 179

Eugenia melastomatifolia 183

Eugenia pancheri 181

Eugenia pseudovenosa 178

Eugenia thompsonii 182

Eugenia trukensis 182

Eugenia vieillardii 181

Euphorbia accedens 447, 456, 458, 589

Euphorbia albrechtii 454, 459, 460, 461,

583, 589

Euphorbia alsiniflora 521

Euphorbia armstrongiana 450, 461

Euphorbia armstrongiana var.

armstrongiana 462, 463, 589

Euphorbia armstrongiana var. distans 462,

463, 583, 589

Euphorbia atoto 580

Euphorbia atoto var. imbricata 550

Euphorbia australis 449, 463

Euphorbia australis var. australis 465, 589

Euphorbia australis var. canescens 513

Euphorbia australis var. erythrantha 465,

589

Euphorbia australis var. glabra 464, 466,

589

Euphorbia australis var. glaucescens 556

Euphorbia australis var. hispidula 465,

467, 589

Euphorbia australis var. potentillina 513

Euphorbia australis var. semiglabra 513

Euphorbia australis var. subtomentosa 464,

468, 583, 589

Euphorbia biconvexa 449, 469, 555, 583, 589

Euphorbia bifida 448, 450, 470, 583

Euphorbia bouleyi 473

Euphorbia bracteolaris 580

Euphorbia careyi 449, 473, 583, 589

Euphorbia carissoides 449, 475, 583, 589

Euphorbia centralis 447, 448, 476, 583, 589

Euphorbia chamaesyce 580

Euphorbia chrysochaeta 499

Euphorbia cinerea 451, 452, 461, 477, 583,

589

Euphorbia clementii 449, 478, 583, 590

Euphorbia coghlcmii 449, 479, 555, 583, 590

Euphorbia comans 558

Euphorbia crassimarginata 455, 480, 482,

583, 590

Euphorbia dallachyana 453, 454, 481, 583,

590

Euphorbia drummondii 456, 484, 486, 583,

590

Euphorbia drummondii var. dallachyana 580

Euphorbia drummondii var. drummondii 484

Euphorbia drummondii var. erythropeplis 580

Euphorbia drummondii var. rubescens560

Euphorbia ferdinandi 451, 486

Euphorbia ferdinandi var. appendiculata

487, 488, 583, 590

Euphorbia ferdinandi var. ferdinandi 487,

488, 489, 583, 590

Euphorbia ferdinandi var. saxosiplaniticola

487, 490, 583, 590

Euphorbia filipes 521

Euphorbia fitzroyensis 455, 491, 492, 583,

590

Euphorbia flindersica 452, 454, 493, 585,

590

Euphorbia gregoriensis 451, 455, 494, 495,

585, 590

Euphorbia hassallii 451, 455, 456, 496,

498, 585, 590

Euphorbia hirta 446, 497, 590

Euphorbia hypericifolia var. bracteolaris 580

Euphorbia hyssopifolia 450, 500, 585, 590

Euphorbia inappendiculata 451, 453, 455,

456, 501,507

Euphorbia inappendiculata var.

inappendiculata 502, 503, 585, 590

Euphorbia inappendiculata var.

queenslandica 502, 503, 504, 585, 591

Euphorbia inappendiculata var. robustior

502, 505, 585, 591

Euphorbia kimberleyensis 446, 450, 506,

585, 591

Euphorbia laciniloba 451, 585, 591

Euphorbia levis var. imbricata 508

Euphorbia litticola 449, 508, 509, 511, 585,

591

Euphorbia macdonaldii 447, 451, 452, 510

Euphorbia macdonaldii var. macdonaldii

513.514.585, 591

Euphorbia macdonaldii var. potentillina

513.585, 591

Euphorbia macgillivrayi 470

Euphorbia macgillivrayi f. glabrata 470

Euphorbia macgillivrayi vm. filipes 521

Euphorbia macgillivrayi var. macgillivrayi

470

Euphorbia macgillivrayi var. micradenia 470

Euphorbia macgillivrayi var. pseudoserrulata

470

Euphorbia macgillivrayi var. yarrabensis 470

Euphorbia maconochieana 448, 451, 515,

585, 591

Euphorbia maculata 448, 516, 585, 591

Euphorbia “Marree” (F.J. Badman 776) 503

Euphorbia micradenia 470

Euphorbia minutifolia 481

Euphorbia mitchelliana 447, 448, 450, 518

Euphorbia mitchelliana var. cairnsiana 520

Euphorbia mitchelliana var. dietrichiae 520

Euphorbia mitchelliana vdj.filifolia 520

Euphorbia mitchelliana var. filipes 519,

521.591

Euphorbia mitchelliana var. glauca 567

Euphorbia mitchelliana var. hirta 519

Euphorbia mitchelliana var. longiloba 519,

522, 591

Euphorbia mitchelliana var. mitchelliana

519,585, 591

Euphorbia mitchelliana var. oblongifolia 520

Euphorbia mitchelliana var. stenophylla 470

Euphorbia muelleri 447, 450, 523, 585, 591

Euphorbia multifaria 452, 453 456, 524,

526, 585, 591

Euphorbia myrtoides 453, 454, 527, 587, 591

Euphorbia obliqua 449, 528, 587, 592

Euphorbia occulta 447, 529, 531, 587, 592

Euphorbia ophiolitica 448, 453, 454, 530,

587, 592

Euphorbia ophthalmica 446, 532, 587, 592

Euphorbia pallens 449, 534, 587, 592

Euphorbiapapillata 451, 452, 455, 535

Euphorbia papillata var. laevicaulis 536,

538, 592

Euphorbia papillata var. papillata 536, 537,

587, 592

Euphorbiapapillifolia 450, 452, 453, 454,

455, 538, 587

Euphorbia papillifolia var. papillifolia 540,

541.592

Euphorbia papillifolia var. polyandra 540,

592

Euphorbiapetala 453, 454, 461, 542, 587,

592

Euphorbiaphilochalix 452, 454, 543, 545,

587, 592

Euphorbia pilulifera 580

Euphorbia pilulifera f. humifusa 497

Euphorbia pilulifera f. rubromaculata 497

Euphorbia pilulifera f. viridis 499

Euphorbiaporcata 454, 455, 456, 546, 547,

587, 592

Euphorbia prostrata 447, 548

Euphorbiapsammogeton 450, 550, 587, 592

Euphorbiapsilosperma 449, 551, 553, 555,

587, 592

Euphorbia pubicaulis 522

Euphorbia schizolepis 447, 450, 552, 587,

593

Euphorbia schizolepis var. glabra 475

Euphorbia schultzii 447, 448, 451, 452, 555,

587

Euphorbia schultzii var. comans 556, 558,

588, 593

Euphorbia schultzii var. schultzii 556, 593

Euphorbia section Anisophyllum 446

Euphorbia serpens 452, 559, 588, 593

Euphorbia serrulata 470

Euphorbia sharkoensis 452, 453, 454, 560,

593

Euphorbia sp. Beddome Range (D.E.Albrecht

5656)562

Euphorbia sp. Hale River (B.G. Thomson

3395)567

Euphorbia supina 516

Euphorbia thelephora 447, 450, 452, 561

Euphorbia thelephora var. australis 562,

564, 588, 593

Euphorbia thelephora var. rugosa 562, 565,

588, 593

Euphorbia thelephora var. thelephora 562,

563, 588, 593

Euphorbia thymifolia 448, 566, 588, 593

Euphorbia trigonosperma 449, 555, 567,

569, 588, 593

Euphorbia vaccaria 446, 568

Euphorbia vaccaria var. erucoides 570, 571,

593

Euphorbia vaccaria var. vaccaria 570, 588,

593

Euphorbia vachellii 470

Euphorbia verrucitesta 456, 572, 573, 588,

593

Euphorbia vicina 448, 450, 574, 575, 588,

593

Euphorbia victoriensis 453, 576, 578, 588,

593

Euphorbia wheeleri 450, 577, 588, 593

Euphorbiaceae 143, 616

Evolvulus hederaceus 57

Fabaceae 65, 159, 438

Fell, D.G. & Stanton, D.J. (2011). Luvunga

monophylla (DC.) Mabb. (Rutaceae): a

new species for Queensland. 431

Field, A. (2012). Poikilogyne cordifolia

(Cogn.) Mansf., a newly recorded genus

and species of Melastomataceae for

Australia 613

Field, A. & Zich, F.A. (2012). Types of

enigmatic north-Queensland Orchids from

the Dockrill herbarium 696

Ford, A.J. & Hasenpusch, J.W. (2009).

Rediscovery of Uncaria cordata (Lour.)

Merr. var. cordata (Rubiaceae: Naucleeae)

in Australia 99

Ford, A.J. & Weston, P.H. (2012). A

taxonomic revision of Hollandaea

F.Muell. (Proteaceae) 670

Ford, A.J. & Zich, F. (2011). Wilkiea

kaarruana Zich & A. J.Ford

(Monimiaceae) a new species from

north-east Queensland 405

Ford, A.J. with Cooper, W.E. (2010).

Trichosanthes odontosperma W.E.Cooper

& A. J.Ford (Cucurbitaceae) a new species

from Queensland’s Wet Tropics 125

Ford, A.J. with Halford, D A. (2009).

Coelospermum purpureum Halford &

A. J.Ford (Rubiaceae), a new species from

north-east Queensland 69

Ford, A.J. with Halford, D.A. (2009). Two

new species of Morinda L. (Rubiaceae)

from north-east Queensland 81

Forster, PI. (2010). Book Review ^Australian

Palms - biogeography, ecology and

systematics 132

Forster, PI. (2010). Book Review, Rainforest

Restoration Manual for South-Eastern

Australia 118

Forster, PI. (2010). Book Review, The

Flowering of Australia’s Rainforests - a

plant and pollination miscellany 149

Forster, PI. (2010). Croton dichromifolius

PI.Forst. (Euphorbiaceae) a new species

from Cape York Peninsula Queensland

143

Forster, PI. (2010). The genus Amaracarpus

Blume (Rubiaceae) in mainland Australia

155

Forster, PI. (2011). Cycas terryana PI.Forst.

(Cycadaceae) a new species from central

Queensland 356

Forster, PI. (2011). Five new species of

Plectranthus L.Her. (Lamiaceae) from

New South Wales and Queensland 387

Forster, PI. (2012). Croton lucens PI.Forst.

(Euphorbiaceae), a new species from

south-east Queensland 616

Forster, PI. & Smith, M.W. (2010). Citrus

wakonai PI.Forst. & M.W.Sm. (Rutaceae)

a new species from Goodenough Island,

Papua New Guinea 133

Galophthalmum brasiliense 666

Geissoieae 256

Geissois lachnocarpa 263

Gen.(Aq 124851) sp. (Boonjie L.J.Webb+

6837A) 85

Gnaphalium cylindrostachyum 306

Gnaphalium redolens 327

Goodenia heteroptera 691, 693

Goodenia minima 689

Goodenia minutiflora 688

Goodeniapaludicola 688, 689

Goodenia pilosa 691, 692, 693

Goodenia purpurascens var. minima 689

Goodeniaceae 688

Gossia alaternoides 180

Gossia alaternoides var. conspicua 180

Gossia alaternoides var. pulchrifolius 180

Gossia aneityensis 181

Gossia aphthosa 180

Gossia clusioides 180

Gossia colnettiana 180

Gossia diversifolia 180

Gossia kuakuense 181

Gossia nigripes 181

Gossia pancheri 181

Gossia vieillardii 181

Gossia virotii 182

Grevilleoideae 670

Guajava cattleiana 178

Guajava guineensis 179

Guymer, G.P (2009). Mischocarpus ailae

Guymer (Sapindaceae), a new species

from the Mount Warning caldera,

Australia 91

Halford, D.A. & Ford, A.J. (2009).

Coelospermum purpureum Halford &

A.J.Ford (Rubiaceae), a new species from

north-east Queensland 69

Halford, D.A. & Ford, A.J. (2009). Two new

species of Morinda L. (Rubiaceae) from

north-east Queensland 81

Halford, D.A. & Harris, W.K. (2012). A

taxonomic revision of Euphorbia section

Anisophyllum Roeper (Euphorbiaceae) in

Australia 441

Harris, W.K. with Halford, DA. (2012). A

taxonomic revision of Euphorbia section

Anisophyllum Roeper (Euphorbiaceae) in

Australia 441

Hasenpusch, J.W. with Ford, A.J. (2009).

Rediscovery of Uncaria cordata (Lour.)

Merr. var. cordata (Rubiaceae: Naucleeae)

in Australia 99

Helicia sayeriana 683

Helicieae 671

Heliciinae 671

Heliotropium cunninghamii 338, 339

Heliotropium microspermum 335, 336, 337,

338

Heliotropium rhadinostachyum 338, 339

Hodgon, J. with Booth, R. & Moore, D. J.

(2009). Four new species of Cyperus L.

(Cyperaceae) from northern Queensland

Holland, A.E. (2009). Crotalaria inaequalis

A.E.Holland (Fabaceae), a new species

from the Gulf Plains, Queensland 65

Holland, A.E. (2010). Notes on Zornia

J.F.Geml. 1: a new species from north

Queensland and a new synonym 159

Holland, A.E. (2011). Reinstatement of Vigna

suberecta Benth. (Fabaceae: Phaseoleae)

438

Holland, A.E. (2012). Nomenclature and

synonymy of several Goodenia R.Br.

(Goodeniaceae) species from northern

Australia 688

Hollandaea 670, 673

Hollandaea diabolica 674, 676, 677, 685

Hollandaea porphyrocarpa 674, 677, 678,

679, 685

Hollandaea riparia 674, 677, 681, 682, 685

Hollandaea sayeri 683

Hollandaea sayeriana 67 A, 677, 683, 685

Hollandaea sp. (Devils Thumb P.I.Forster+

PIF10720) 674

Hollandaea sp. (Devils Thumb) 674

Hollandaea sp. (Pinnacle Rock Track

PI. Forster PIF10714)678

Hollandaea sp. (Pinnacle Rock Track

PI.Forster+ PIF 10714) 678

Hollandaea sp. (Pinnacle Rock Track) 678

Hollandaeinae 671

Hollandinae 671

Homoranthus tricolor 77,78

Hydrocharis dubia 435, 436

Hydrocharis morsus-ranae 435

Hydrocharitaceae435

Ipomoea 699

Ipomoea abrupta 720, 721

Ipomoea aculeata 718

Ipomoea aegyptia 57

Ipomoea alba 717

Ipomoea antonschmidii 721

Ipomoea aquatica 719

Ipomoea argillicola 720

Ipomoea batatas 719

Ipomoea bracteolata 699, 701, 721, 722

Ipomoea brassii 720

Ipomoea brownii 7 19

Ipomoea cairica 718

Ipomoea calobra 721

Ipomoea carnea 720

Ipomoea chryseides 57

Ipomoea cinerascens 57

Ipomoea coptica 718

Ipomoea costata 721

Ipomoea cymosa 62

Ipomoea davenportii 57, 173

Ipomoea densivestita 701, 702, 721, 722

Ipomoea diamantinensis 719

Ipomoea diversifolia 718

Ipomoea dunlopii 702, 705, 720, 722

Ipomoea eriocarpa 717

Ipomoea flava 57

Ipomoea funicularis 704, 705, 719, 722

Ipomoea gracilis 720

Ipomoea gracilis var. sagittata 705

Ipomoea graminea 718

Ipomoea hederacea 718

Ipomoea hederifolia 717

Ipomoea hirsuta 61

Ipomoea imperati 719

Ipomoea incisa 57, 718

Ipomoea kalumburu 707, 709, 720, 722

Ipomoea lasiophylla 701

Ipomoea limosa 708, 711, 720, 722

Ipomoea linifolia 57

Ipomoea littoralis 720

Ipomoea lonchophylla 717

Ipomoea mauritiana 719

Ipomoea modesta 49

Ipomoea muelleri 720

Ipomoea nil 718

Ipomoea obscura 719

Ipomoea ochracea 719

Ipomoea pandurata 720

Ipomoea peltata 61

Ipomoea pentaphylla 57

Ipomoea pes-caprae 719

Ipomoea pes-tigridis 718

Ipomoea plebeia 718

Ipomoea polpha 720

Ipomoeapolymorpha 111

Ipomoea purpurea 718

Ipomoea quamoclit 111

Ipomoea quinata 61

Ipomoea quinquefolia 62

Ipomoea racemigera 111

Ipomoea saintronanensis 718

Ipomoea sinuata 57

Ipomoea sp. (Kununurra T.E.Aplin 6307) 710

Ipomoea sp. (Mungana L. J.Webb+ 10184)

701

Ipomoea sp. A Kimberley Flora (L J. Penn

84)713

Ipomoea sp. A 713

Ipomoea sp. B 715

Ipomoea sp. Cobourg (G.M.Wightman 380)

699

Ipomoea sp. Kalumburu (A.A.Mitchell

3869B) 707

Ipomoea sp. Mt Fyfe (R.L.Barrett &

M.D.Barrett RLB 1526) 699

Ipomoea sp. OT Station (S.T.Blake 17676)

715

Ipomoea sp. Tolmer (C.R.Dunlop 6787) 713

Ipomoea tiliacea 720

Ipomoea tolmer anal 12

Ipomoea tolmerana subsp. tolmerana 713,

723

Ipomoea tolmerana subsp. occidentalis 713,

718, 723

Ipomoea trichosperma 718

Ipomoea tricolor 720

Ipomoea triloba 719

Ipomoea tuberosa 62

Ipomoea versipellis 715, 716, 720, 721, 723

Ipomoea violacea 718

Ipomoea yardiensis 721

Isachne miliacea var. minutula 205

Isachne minutula 205, 206

Isachne pulchella 205

Isachne sharpii 207, 208

Isachne sp. (Cape Tribulation R.L. Jago 4560)

207

Isachne sp. A 207

Jackes, B.R. (2009). Taxonomic revision of

Australian Myrsinaceae: Ardisia Sw. and

Tetrardisia Mqz. 1

Jackes, B.R. (2010). Ardisia hylandii Jackes:

a new name for Ardisia depauperata

(Domin) Jackes 223

Jambosa bifaria 183

Jambosa melastomatifolia 183

James, J.K. with Duretto, M.F. (2010).

Three new combinations in Pterostylis

( Orchidaceae ) 221

Jessup, L.W. (2011). A taxonomic revision of

Symplocos Jacq. (Symplocaceae) in

Australia 225

Jobson, R.W. (2012). Utricularia corneliana

R.W. Jobson (Lentibulariaceae), a new

species from the North Kennedy district of

Queensland 601

Johnson, R.W. (2009). A conspectus of

Merremia Dennst. ex Endl.

(Convolvulaceae) in Australia with the

addition of two species 55

Johnson, R.W. (2009). Duperreya Gaudich.

(Convolvulaceae) revisited 47

Johnson, R.W. (2010). Davenportia

R.W.Johnson, a new genus of

Convolvulaceae ( Merremieae ) from

central Australia 171

Johnson, R.W. (2012). New species and

subspecies of Ipomoea L.

(Convolvulaceae) from northern Australia

and a key to the Australian species 699

Kleinig D. withNicolle, D. (2011).

Eucalyptus melanophloia subsp. nana

D.Nicolle & Kleinig a new mallee

ironbark (E. series Siderophloiae Blakely;

Myrtaceae) from central Australia and

north western Queensland. 347

Knightieae 671

Lamiaceae 387

Leiper, G. with Little, R.J. (2012). Capsule

dehiscence in Viola betonicifolia Sm.

(Violaceae) 624

Lentibulariaceae 601

Limnobium dubium 435

Lipochaeta amazonica 666

Little, R.J. & Leiper, G. (2012). Capsule

dehiscence in Viola betonicifolia Sm.

(Violaceae) 624

Luvunga 431

Luvunga monophylla 431

Mathieson, M.T. (2010). Cadetia uniflos

(F.M.Bailey) M.T.Mathieson a new

combination in Orchidaceae 119

Mathieson, M.T. with Bean, A.R. (2012).

Stylidium elachophyllum A.R.Bean &

M.T.Mathieson (Stylidiaceae), a new

species from northern Queensland 608

Melastomataceae 613

Merremia 55

Merremia aegyptia 56, 57

Merremia caespitosa 57

Merremia chryseides 57

Merremia da\enportii 56, 57, 173

Merremia dissecta 55,57

Merremia gemella 56, 57

Merremia gemella var. gemella 57

Merremia hederacea 56, 57

Merremia hirta 56, 57

Merremia hirta var. hirta 57

Merremia incisa 56, 57, 59, 62

Merremia kimberleyensis 56, 60, 61, 62

Merremia nymphaeifolia 61

Merremia peltata 56, 61

Merremia pentaphylla 57

Merremia quinata 56, 61

Merremia quinquefolia 56, 62

Merremia sp. B 58

Merremia tuberosa 55,62

Merremia umbellata 56, 62

Merremia umbellata subsp. orientalis 62

Merremia umbellata var. orientalis 62

Miller, C.H. with Bean, A.R. (2011). A

taxonomic revision of Nyssanthes R.Br.

(Amaranthaceae) 267

Mischocarpus ailae 91, 93, 94

Mischocarpus albescens 94

Mischocarpus anodontus 94

Mischocarpus australis 95

Mischocarpus exangulatus 94

Mischocarpus grandissimus 95

Mischocarpus lachnocarpus 91, 94

Mischocarpus macrocarpus 95

Mischocarpus montanus 94

Mischocarpus pyriformis 95

Mischocarpus stipitatus 95

Monenteles forsteri 306

Monenteles glandulosus 308

Monenteles glandulosum var. velutinum 311

Monenteles globiferus 298

Monenteles intermedius 298

Monenteles redolens 306

Monenteles serrulatus 308

Monenteles sphacelatus 314

Monenteles sphaeranthoides 317

Monenteles spicatus 306

Monenteles tomentosus 327

Monenteles verbascifolius 323

Monimiaceae 405

Moore, D.J. with Booth, R. & Hodgon, J.

(2009). Four new species of Cyperus L.

(Cyperaceae) from northern Queensland

Morinda constipata 82, 84, 90

Morinda hypotephra 85

Morinda retropila 85, 87, 90

Morinda sp. (Bellenden Ker W.Cooper+

1526)82

Morinda sp. (Bellenden Ker) 82

Morinda sp. (Boonjee) 85

Morinda sp. 1 85

Morinda sp. 2 82

Morindeae 69

Mosiera guineensis 179

myrmechory 624

Myrsinaceae 1, 223

Myrtaceae 25, 77, 139, 177, 347

Myrtus alaternoides 180

Myrtus aneityensis 181

Myrtus conspicua 180

Myrtus diversifolia 180

Myrtus flavida 180

Myrtus guineensis 179

Myrtus nigripes 181

Myrtus prolixa 181

Myrtus pulchrifolius 180

Myrtus virotii 182

Nicolle, D. & Kleinig, D. (2011). Eucalyptus

melanophloia subsp. nana D.Nicolle &

Kleinig a new mallee ironbark {E. series

Siderophloiae Blakely; Myrtaceae) from

central Australia and north western

Queensland 347

Nyssanthes 267, 268

Nyssanthes diffusa 269, 271, 273, 279

Nyssanthes erecta 269, 270, 278

Nyssanthes impervia 269, 274, 275, 279

Nyssanthes longistyla 269, 276, 277, 279

Nyssanthes media 271

Oberonia attenuata 698

Oligochaetochilus 221

Oligochaetochilus mystacinus 221

Oligogyne bahiensis 666

Oligogyne burchellii 656

Oligogyne megapotamica 666

Oligogyne synedrelloides 666

Oligogyne tampicana 666

Operculina peltata 61

Oplismenus mollis 209, 210

Oplismenus undulatifolius var. mollis 209

Orchard, A.E. (2012). The Australian species

of Blainvillea Cass. (Asteraceae: Ecliptinae)

653

Orchidaceae 119, 221

Orites sp. (Pinnacle Rock Track WWC 867)

674

Paniceae 187, 634

Panicoideae 634

Panicum breviglume 200

Panicum heteroneuron 216

Panicum minutulum 205

Panicum notochthonum 216

Panicum pallide-fuscum 215

Panicum parviflorum var. pilosum 204

Paractaenum 210

Paspalidium johnsonii 206, 211, 212

Paspalidium sp. (Musselbrook R.W.Johnson+

MRS792) 212

Pellow, B. with Rozefelds, A.C. (2011). A

taxonomic revision of Pseudoweinmannia

Engl. (Cunoniaceae: Geissoieae) 252

Pennisetum advena 191

Pennisetum alopecuroides 191

Pennisetum basedowii 191

Pennisetum ciliare 191

Pennisetum clandestinum 191

Pennisetum elymoides 191

Pennisetum glaucum 191

Pennisetum macrourum 191

Pennisetum pedicellatum 191

Pennisetum pedicellatum subsp. unispiculum

191

Pennisetum pennisetiforme 191

Pennisetum polystachion 191

Pennisetum purpureum 191

Pennisetum setaceum 191

Pennisetum setigerum 191

Pennisetum thunbergii 191

Pennisetum villosum 191

Petrorchis 221

Plagiosetum 210

Plectranthus bellus 388, 389, 390, 391, 392

Plectranthus caldericola 392, 393, 395

Plectranthus fragrantissimus 394, 395, 396

Plectranthus gratus 402

Plectranthus insularis 395, 398, 399, 400

Plectranthus intraterraneus 399

Plectranthus parviflorus 399

Plectranthus sp. (Restoration Island J. Le

Cussan 285) 401

Plectranthus spectabilis 391

Plectranthus venustus 401, 402, 403

P/wcto 340, 345

Pluchea baccharoides 345

Pluchea dentex 345

Pluchea dioscoridis 340

Pluchea dunlopii 345

Pluchea ferdinandi-muelleri 345

Pluchea indica 345

Pluchea punctata 343, 344, 345, 346

Pluchea rubelliflora 345

Pluchea xanthina 340, 342, 345, 346

Poaceae 187, 634

Poikilogyne 613

Poikilogyne cordifolia 613

Poikilogyne cordifolia var. cordifolia 614,

615

Porana commixta 48

Porana sericea 49

Proteaceae 670

Proteaceae sp. ‘Devils Thumb’ 674

Pseudoraphis minuta 214

Pseudoraphis jagonis 212, 213, 214

Pseudoraphis minuta var. laevis 214

Pseudoraphis minuta var. minuta 214

Pseudoraphis sp. (Port Douglas R.L. Jago

6610)212

Pseudoweinmannia 252, 260

Pseudoweinmannia apetala 254, 257, 259,

261,262, 266

Pseudoweinmannia lachnocarpa 255, 257,

259, 261,262, 263, 266

Psidium cattleianum 177, 178

Psidium cattleianum var. purpureum 178

Psidium cattleianum var. pyriformis 178

Psidium floribundum 181

Psidium guineense 179

Psidium kuakuense 181

Psychotria nematopoda 156

Pterocarpus sp. (Archer River B.RHyland

3078)151

Pterocaulon 280

Pterocaulon billardierei 306

Pterocaulon brachyanthum 286, 288, 289,

290, 330

Pterocaulon ciliosum 285, 288, 290, 291,

292, 330

Pterocaulon ciliosum x P. serrulatum var.

serrulatum 327

Pterocaulon cylindrostachyum 306

Pterocaulon discolor 288, 290, 293, 294, 331

Pterocaulon glandulosum 308

Pterocaulon glandulosum var. glandulosum

308

Pterocaulon globuliflorum 288, 295, 296,

297,331

Pterocaulon intermedium 288, 297, 298,

299, 331

Pterocaulon intermedium x p paradoxum

327

Pterocaulon niveum 287, 300, 301, 302, 330

Pterocaulon paradoxum 288, 302, 303, 304,

332

Pterocaulon redolens 288, 302, 306, 307, 332

Pterocaulon serratum 311

Pterocaulon serrulatum 288, 308

Pterocaulon serrulatum var. serrulatum 284,

310,311,313, 333

Pterocaulon serrulatum var. velutinum x p

sphacelation 327

Pterocaulon serrulatum var. velutinum 284,

311,312, 333

Pterocaulon sp. (Yarrowmere Station E.J.

Thompson+ BUC340) 298

Pterocaulon sp. A Kimberley Flora

(B.J. Carter 599) 298

Pterocaulon sp. A 298

Pterocaulon sphacelatum 285, 288, 313, 314,

316, 333

Pterocaulon sphaeranthoides 288, 313, 317,

318, 331

Pterocaulon spicatum 306

Pterocaulon tomentosus 327

Pterocaulon tricholobum 287, 319, 320, 322,

334

Pterocaulon verbascifolium 288, 322, 323,

324, 334

Pterocaulon xenicum 288, 322, 325, 326, 331

Pterostylis mystacina 221

Pterostylis section Oligochaetochilus 221

Pterostylis section Parviflorae 221

Pterostylis section Speculantha 221

Pterostylis series Parviflorae 221

Randia sp. (Boonjee BG5345) 70

Randia sp. (Boonjee) 70

Randia sp. (Boonjie L.W.Jessup+ GJM264)

Restiaria cordata 99

Rhodomyrtus kaweaensis 183

Rhodomyrtus misimana 184

Rhodomyrtus montana 184

Roupaleae 671, 673

Rozefelds, A.C. & Pellow, B. (2011). A

taxonomic revision of Pseudoweinmannia

Engl. (Cunoniaceae: Geissoieae ) 252

Rubiaceae gen. nov. sp. (Boonjee) 85

Rubiaceae 69, 81, 99, 155

Rutaceae 133, 431

Sapindaceae 91

Setaria glauca subsp. subtesselata 215

Setaria glauca var. pallide-fusca 215

Setaria pallide-fusca 215

Setariapumila subsp. pallide-fusca 215

Setaria pumila subsp. subtesselata 215

Simon, B.K. (2010). New taxa nomenclatural

changes and notes on Australian grasses in

the tribe Paniceae (Poaceae: Panicoideae)

187

Simon, B.K. with Thompson, E.J. (2012).

A revision of Calyptochloa C.E.Hubb.

(Poaceae), with two new species and a

new subspecies 634

Smith, M.W. with Forster, PI. (2010). Citrus

wakonai P.I.Forst. & M.W.Sm. (Rutaceae)

a new species from Goodenough Island,

Papua New Guinea 133

Snow, N. & Veldkamp, J.F. (2010).

Miscellaneous taxonomic and

nomenclatural notes for Myrtaceael77

Solanaceae 165, 412

Solanum aridicola 414, 415, 417, 429

Solanum callosum 414, 416, 419, 430

Solanum cleistogamum 414, 419

Solanum crebrispinum 414

Solanum dianthophorum 414

Solanum ellipticum 412, 413, 414, 421

Solanum ellipticum form “b” 415

Solanum ellipticum var. ellipticum 421

Solanum ellipticum var. Foot hills (G.J.Leach

1145) 424

Solanum ellipticum var. mollibaccalis 424

Solanum ellipticum var. Ranges

(D.E.Albrecht 5807) 415

Solanum ellipticum var. typicum 421

Solanum emmottii 414, 422, 423, 430

Solanum horridum 414

Solanum lithophilum 414, 415, 424, 426, 430

Solanum orgadophilum 166, 169, 170

Solanum petrophilum var. pedicellatum 419

Solanum pisinnum 165, 167, 170

Solanum quadriloculatum 414

Solanum senticosum 414

Solanum unispinum 414, 425, 428, 429

Speculantha 221

Speculantha section Elongatae 221

Sphaeranthus elongatus 306

Spilanthes acmella 662

Stanton, D.J. with Fell, D.G. (2011). Luvunga

monophylla (DC.) Mabb. (Rutaceae):

a new species for Queensland 431

Stylidiaceae 107, 608

Stylidium centrolepoides 115, 116

Stylidium elachophyllum 608, 610, 611

Stylidium exiguum 107, 108, 109, 117

Stylidium nominatum 107, 110

Stylidium notabile 112, 113, 114

Stylidium osculum 108, 110, 111, 117

Symplocaceae 225

Symplocos 225, 226

Symplocos ampulliformis 227, 228, 229, 230,

231,232

Symplocos baeuerlenii 228, 231, 232

Symplocos boonjee 227, 229, 230, 232, 233

Symplocos bullata 227, 229, 230, 234

Symplocos candelabrum 228, 230, 235

Symplocos cochinchinensis subsp. thwaitesii

248

Symplocos cochinchinensis var. candelabrum

235

Symplocos cochinchinensis var. gittonsii 238

Symplocos cochinchinensis var. glaberrima

239

Symplocos cochinchinensis var. montana 249

Symplocos cochinchinensis var. pilosiuscula

246

Symplocos cochinchinensis var. stawellii 247

Symplocos cochinchinensis var. thwaitesii

248

Symplocos crassiramifera 228, 230, 235, 236

Symplocos cyanocarpa 230, 236

Symplocos cyanocarpa var. cyanocarpa 229,

236, 237, 238

Symplocos cyanocarpa var. pilosa 228, 237,

238

Symplocos gittinsii 229, 230, 238, 239

Symplocos glabra 228, 229, 230, 237, 239,

240

Symplocos graniticola 229, 230, 240, 241

Symplocos harroldii 228, 231, 241, 242

Symplocos hayesii 229, 231, 237, 242, 243

Symplocos hylandii 229, 230, 243, 244, 245

Symplocos oresbia 228, 229, 230, 244, 245

Symplocospaucistaminea 229, 230, 245, 246

Symplocospuberula 228, 230, 246, 247

Symplocos sp. (Boonjie B.RHyland 2753)

233

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

244

Symplocos sp. (North Mary B.Gray 2543)

234

Symplocos sp. ‘Mt Lewis’ 240

Symplocos sp. 1. 241

Symplocos spicata var. australis 247

Symplocos stawellii 229, 231, 247, 248

Symplocos stawellii var. montana 249

Symplocos stawellii var. stawellii 247

Symplocos thwaitesii 228, 230, 231, 248, 249

Symplocos wooroonooran 228, 230, 245,

249, 250

Syzygium bifarium 183

Syzygium melastomatifolium 183

Syzygium thompsonii 182

Tapeinosperma pseudojambosa 21

Tapeinosperma repandulum 21

Tessaria redolens 306

Tetrardisia 18

Tetrardisia bifaria 19, 23

Tetrardisia disticha 19

Thompson, E.J. & Simon, B.K. (2012).

A revision of Calyptochloa C.E.Hubb.

(Poaceae), with two new species and a

new subspecies 634

Thompson, E.J. (2011). Heliotropium

microspermum E.J.Thomps.

(Boraginaceae) a new species from

Queensland 335

Trichosanthes 125, 364, 366

Trichosanthes ambrozii 368

Trichosanthes ascendens 373

Trichosanthes baviensis 373

Trichosanthes boninensis 373

Trichosanthes brevibracteata 368

Trichosanthes cavaleriei 373

Trichosanthes chinensis 371

Trichosanthes chingiana 373

Trichosanthes cucumerina 368

Trichosanthes cucumerina var. cucumerina

368, 369, 385

Trichosanthes cucumeroides 371

Trichosanthes cucumeroides var.

dicoelosperma 372

Trichosanthes cucumeroides var. hainanensis

373

Trichosanthes cucumeroides var. stenocarpa

373

Trichosanthes dicoelospermum 372

Trichosanthes formosana 373

Trichosanthes hainanensis 373

Trichosanthes hearnii 371

Trichosanthes himalensis 371

Trichosanthes holtzei 373

Trichosanthes horsfieldii 371

Trichosanthes mafuluensis 373

Trichosanthes matsudai 373

Trichosanthes morrisii 368, 381, 382, 386

Trichosanthes odontosperma 126, 128, 129,

131,368, 375

Trichosanthes okamotoi 373

Trichosanthes ovigera 371

Trichosanthes pachyrrhachis 368

Trichosanthes pedatifolia 368

Trichosanthespentaphylla 365, 368, 379,

380, 386

Trichosanthes pierrei 373

Trichosanthespilosa 368, 371, 372, 385

Trichosanthes reniformis 368

Trichosanthes rostrata 373

Trichosanthes section Cucumeroides 367

Trichosanthes section Cucumeroides 371

Trichosanthes section Edulis 367, 375

Trichosanthes section Foliobracteola 361,

376

Trichosanthes section Involucraria 367, 378

Trichosanthes section Trichosanthes 361, 368

Trichosanthes sp. (D55403) 381

Trichosanthes sp. (Mt Lewis B.Gray 167) 126

Trichosanthes sp. (Mt Lewis BG 167) 126

Trichosanthes sp. (Mt Lewis) 126

Trichosanthes sp. (Williams 1987: 306 126

Trichosanthes sp. A (Telford 1982: 196) 126

Trichosanthes sp. Fine Leaf (L.A.Craven

7930)369

Trichosanthes sp. Kakadu (C.R. Dunlop

6639)381

Trichosanthes sp. Nitmiluk 368

Trichosanthes subvelutina 365, 368, 376,

377, 385

Trichosanthes trichocarpa 313

Trichosanthes vanoverberghii 313

Triuniinae 61 1

Uncaria callophylla 100, 101

Uncaria cordata 99, 101

Uncaria cordata var. cordata 99, 100

Uncaria lanosa var. appendiculata 100, 101

Uragoga nematopoda 156

Urochloa gilesii subsp. occidentals 215

Urochloa gilesii var. gilesii 216

Urochloa gilesii var. nothochthona 216, 217

Urochloa notochthona 216

Urochloa occidentalis 215

Urochloa occidentalis var. ciliata 215, 216

Urochloa occidentalis var. occidentalis 215

Utricularia aurea 606

Utricularia corneliana 602, 604, 605, 606

Utricularia gibba 606

Utricularia muelleri 606

Utricularia reflexa 606

Utricularia stellaris 606

Utricularia tubulata 606

Veldkamp, J.F. with Snow, N. (2010).

Miscellaneous taxonomic and

nomenclatural notes for Myrtaceae. 177

Verbesina acmella 662

Verbesina dichotoma 662, 663

Verbesina lavenia 662

Vigna lanceolata 440

Vigna sp. (Jimbour A.R.Bean 12534) 438

Vigna suberecta 438, 440

Viola betonicifolia 624, 626, 627, 628, 629

Violaceae 624

Wedelia 653

Wedelia cunninghamii 660

Wedelia sp. (Marrett River J.A.Elsol+ 680)

663

Wedelia sp. Limestone (J. Russell-Smith

7865) N.T. Herbarium 658

Wedelia tenuicaulis 666

Wedelia urticifolia 660

Weinmannia apetala 261

Weinmannia lachnocarpa 263

Weinmannia lachnocarpa var. parvifolia 263

Weston, P.H. with Ford, A.J. (2012). A

taxonomic revision of Hollandaea

F.Muell. (Proteaceae) 670

Wilkiea austroqueenslandica 408

Wilkiea hylandii 408

Wilkiea kaarruana 406, 408, 410, 411

Wilkiea smithii 408

Wilkiea sp. A 406

Wilkiea sp. Palmerston (B.P.Hyland 80) 406

Wilkiea sp. Palmerston (BH 80RFK) 406

Wilkiea sp. Palmerston 406

Windmannia lachnocarpa 263

Xanthomyrtus hienghenensis 179, 180

Xanthomyrtus hienghenensis var.

hienghenensis 180

Xanthomyrtus hienghenensis var. latifolia 180

Xanthomyrtus kanalaensis 179

Zich, F. withFord, A.J. (2011). Wilkiea

kaarruana Zich & A. J.Ford

(Monimiaceae) a new species from north¬

east Queensland 405

Zich, F.A. with Field, A. (2012). Types of

enigmatic north-Queensland Orchids

from the Dockrill herbarium 696

Zornia chaetophora 162

Zornia dyctiocarpa DC. var. dyctiocarpa 160

Zornia floribunda 162

Zornia macdonaldii 159, 161, 162, 163

Zornia pedunculata 160

Zornia stirlingii 162