Austrobaileya

Volume 6 Number 1 2001

AUSTROBAIICVA

A Journal of Plant Systematica

Queensland Herbarium

Queensland Government

Environmental Protection Age ncy

Volume 6

Number 1 2001

A Journal of Plant Systematics

Queensland Herbarium

Queensland Government

Environmental Protection Agency

Editorial Committee

L.W. Jessup (editor)

R.J.F. Henderson (technical advisor)
B.K. Simon (technical advisor)

Desktop Publishing

A.E. Sinclair

Austrobaileya

Vol. 1, No. 1 was published on 1 December 1977
Vol. 5, No. 4 was published on 15 December 2000
Vol. 6, No. 1 was published on 12 December 2001

Austrobaileya is published once per year.

Exchange: This Journal will be distributed on the basis of exchange.

Australian subscribers: Orders for single issues and subscriptions may be placed. The price is
(2001 GST included): A$31.62 per issue for individuals, A$50.60 for institutions, including
postage.

Overseas subscribers: Order for single issues and subscriptions may be placed. The price is
(2001): A$28.75 per issue for individuals, A$46.00 for institutions, including postage.

All correspondence relating to exchange, subscriptions or contributions to this journal should
be addressed to: The Editor, Austrobaileya, Queensland Herbarium, Environmental Protection
Agency (EPA), Brisbane Botanic Gardens, Mt Coot-tha, Mt. Coot-tha Road, Toowong Qld 4066,
Australia.

ISSN0155-4131

© Queensland Herbarium 2001

Austrobaileya is the journal of the Queensland Herbarium and is devoted to publication of
results of sound research and of informed discussion on plant systematics, with special empha¬
sis on Queensland plants.

Opinions expressed by authors are their own and do not necessarily represent the policies or
views of the Queensland Herbarium.

Contents

A taxonomic revision of Convolvulus L. (Convolvulaceae) in Australia

R. W. Johnson. 1

Vanguerieae A.Rich. ex Dum. (Rubiaceae) in Australia, 2. Cyclophyllum Hook.f

S. T. Reynolds & R.J.F. Henderson.41

Revision of the Macrozamia miquelii (F.Muell.) A.DC.

(Zamiaceae section Macrozamia) group

David L. Jones, Paul I. Forster and Ish K. Sharma.67

Oreodendron C. T. White reduced to Phaleria Jack
(Thymelaeaceae, Thymelaeoideae)

B.E. Herber.95

Anew species of Lissanthe R.Br. (Epacridaceae) from Queensland

A.R. Bean.99

Hydnophytum ferrugineum (Rubiaceae: Hydnophytinae), a new species of
ant-plant from Cape York Peninsula, Queensland

Paul I. Forster. 103

Alysicarpus (Leguminosae: Desmodieae) in Australia: a taxonomic revision
Les Pedley.107

Eucalyptus broviniensis (Myrtaceae), a new critically endangered species from
south-eastern Queensland

A.R. Bean. 117

Proiphys infundibularis (Amaryllidaceae), a new species from the Townsville
region of Queensland.

D.L. Jones & J.L. Dowe. 121

Drupe - a term in search of a definition

H. Trevor Clifford and Mary E. Dettmann. 127

A new species of Myriophyllum L. (Haloragaceae) from artesian springs in
Queensland.

D. Halford & R.J. Fensham. 133

Pappus morphology and terminology in Australian and New Zealand thistles
(Asteraceae, tribe Cardueae)

A.R. Bean. 139

Cycas cupida (Cycadaceae), a new species from central Queensland.

Paul I. Forster. 153

New combinations and a new name in Australian Sapotaceae
L.W. Jessup.161

New species of Livistona R. Br. (Arecaceae) from north Queensland and
Papua New Guinea

John L. Dowe and Anders S. Barfod.165

(continued)

Note

Kentrophora S.M. Wilson and Kraft, a new name for an algal genus in tribe
Amansieae (Rhodomelaceae, Rhodophyceae).

R.J.F. Henderson, S.M. Wilson and G.T. Kraft.

175

A taxonomic revision of Convolvulus L.
(Convolvulaceae) in Australia

R.W. Johnson

Summary

Johnson, R.W. A taxonomic revision of Convolvulus L. (Convolvulaceae) in Australia. Austrobaileya
6 (1): 1-39. A revision of the genus Co/ivo/vm/m^L. in Australia is presented. Twelve species are recognised
and described, including four that are new. They are C. graminetinus, C. recurvatus, C. tedmoorei and
C. wimmerensis. In two species, C. angustissimus and C. recurvatus , subspecies have been recognised
and described. C. angustissimus subsp. omnigracilis and C. angustissimus subsp. peninsularum, and
C. recurvatus subsp. nullarborensis are described as new while C. angustissimus subsp. fililobus is a new
combination based on C. erubescens war. fililobus Wawra. A key to identify all species and subspecies is
provided together with distribution maps and illustrations of certain diagnostic characters.

Keywords: Convolvulaceae, Convolvulus, Australia

R.W. Johnson c/- Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens
Mt Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Introduction

The genus Convolvulus was formally
established by Linnaeus (1753) and he
recognised 31 species. From these species, the
name Convolvulus arvensis was selected as the
type of the genus by Hitchcock (1929). In
Genera Plantarum, Linnaeus (1754) attributed
the name to Tourneforte. Linnaeus adopted a
broad concept of the genus including in it
species which are now segregated into other
genera including Calystegia R. Br.
(Convolvulus sepium), Ipomoea L. (C. batatas),
Operculina Silva Manso (C. turpethum),
Merremia Dennst ex Endl. (C. umbellatus),
Evolvulus L. (C. alsinoides) and Xenostegia
D. F. Austin & Staples (C. tridentatus).

The first published record of Convolvulus
from Australia was by Sims (1808) when he
described Convolvulus erubescens from a
specimen collected by Mr Loddiges from New
South Wales. Brown (1810) described three
new species from Australia, namely
C. angustissimus, C. remotus and C. multicaulis.
The latter species is now placed in the genus
Jacquemontia Chois. Later Choisy (1824)

Accepted for publication 16 May 2001

described C. acaulis from Kangaroo Island
while still later Lehmann (1826) described
C. geniculatus though no type was designated.
Subsequently this species was listed as
occurring in Australia. Vriese (1845) described
four new species from Western Australia,
namely C. adscendens, C. huegelii, C. preissii
and C. subpinnatifidus. Mueller (1853)
recognised C. crispifolius, based on a specimen
he collected from Cudnaka in South Australia.
However, in 1864, he concluded that the
extreme variation exhibited by C. erubescens
embraced all previously described species from
Australia including C. crispifolius (Mueller
1864). Bentham (1869) agreed with this
taxomony and decided in his Flora treatment
to recognise only one species, C. erubescens,
to include the segregate species that had already
been described. He also included in
Convolvulus the genera Jacquemontia and
Calystegia. Yet soon after, Bentham & Hooker
(1873) recognised these three genera as being
distinct. Domin (1928) described C. clementii
from northern Western Australia as a new
species occurring across tropical Australia.

Until the late 1980’s, only two species of
Convolvulus were recognised in local and

2

Austrobaileya 6(1): 1-39 (2001)

regional floras, C. arvensis, an early
introduction from Europe, and the highly
variable C. erubescens embracing all previously
described native species. Since then I have
described two species, C. eyreanus and
C. microsepalus from South Australia as new
(Johnson 1987).

According to Mabberley (1997) and
Austin (1998), Convolvulus is a genus of about
100 species mainly of temperate origin, though
Sa’ad (1967) recognised 118 species from the
Mediterranean and Middle East alone while
Austin (1982a), Ooststroom (1953) and
Mabberley (1989) indicated there may be about
250 species in the genus. It is more likely it
comprises about 150 species. Convolvulus is
included in tribe Convolvuleae which also
includes Jacquemontia and Calystegia.

Classification

Ooststroom (1953) recognised three tribes
within the Convolvulaceae. One of these tribes,
Cuscuteae, represented by the genus Cuscuta
L. is regarded as a separate family, Cuscutaceae,
by some authors. The separation of the other
tribes, the Convolvuleae and the Ipomoeeae, is
based on whether their pollen is spinulose
(Ipomoeeae) or smooth (Convolvuleae).
Ooststroom (l.c.) placed Convolvulus in
subtribe Convolvulinae with Calystegia,
Jacquemontia, Merremia, Operculina and
Aniseia Chois. Austin (1973,1975) recognised
9 tribes within Convolvulaceae, two of which
were equivalent to subtribe Convolvulinae of
Ooststroom. His Convolvuleae included
Convolvulus, Calystegia, Polymeria,
Jacquemontia as well as Evolvulus
(Dicranostylinae) while his Merremieae (Austin
1982) included Aniseia, Operculina and
Merremia. This division was based on
cytological data, corolla shape and stylar
characters. However, more recent studies
(Austin 1998) using cladistic analyses suggest
the Merremieae may not form a clade distinct
from the Convolvuleae.

Three sections are recognised within
Convolvulus (Sa’ad 1967), separated on the
presence or absence of spines and whether or
not the stems are twining. All of the Australian
species belong to C. sect. Convolvulus, being
espinose and twining.

Relationships

According to Austin (1973), Convolvulus is
most closely related to Calystegia, Evolvulus,
Polymeria and Jacquemontia. In Australia,
Jacquemontia is clearly distinguished in
possessing triramous hairs while Evolvulus has
biramous hairs. The remaining genera have
simple hairs. The style in Polymeria bears a
much divided stigma with usually 4-8 lobes
unlike that in Convolvulus which has two lobes.
P. distigma, however, has a 2-lobed stigma
though the branches are similar in structure to
the other species of Polymeria and usually have
short lobes at the base. Calystegia is
distinguished by its large bracts which enclose
or closely subtend the calyx. Convolvulus and
Polymeria can also be distinguished from the
other genera in possessing tri-colpate rather
than either pantoporate or pantocolpate pollen
grains.

Materials and Methods

This revision is based principally on herbarium
specimens held at AD, BRI, CANB, HO, MEL,
NE, NSW and PERTH. Only limited field work
in Queensland and the Northern Territory was
possible. In addition, plants were grown from
seed in a glasshouse at BRI. Specimens were
sorted into nominal taxa and, from each,
specimens with flowers and fruit were selected
for comprehensive dissections. Those
specimens which could not be placed within
nominal taxa were also dissected. Attributes
from dissected specimens were then classified
using a variety of numerical taxonomic
programs. While the classifications produced
as a result were not completely satisfactory,
groupings that were produced formed part of
the input into the taxonomic treatment
presented below.

Attributes

Attributes of Australian species of Convolvulus
which are significant diagnostically are
discussed below.

Cotyledons: The cotyledons fall into one of
two morphological groups i.e. those that are:

• bilobed, with the lobes, linear, diverging
and greatly exceeding the base e.g.

C. clementii (Fig. 1A), or

Johnson, Convolvulus L. in Australia

3

B

Fig. 1 . Cotyledons of Convolvulus species. A. C. clementii from S.A. Pastoral Board s.n. (AD98010365); B. C. remotus
from Kenneally 4643 (PERTH).

• oblong to rounded oblong, with two short
rounded apical lobes, always shorter than
the base e.g. C. angustissimus, C. arvensis,
C. crispifolius, C. eyreanus, C. graminetinus,
C. microsepalus, C. recurvatus and
C. remotus (Fig. IB).

Habit : Most species have long trailing and
twining stems. The initial leaves are somewhat
similar in shape in all the Australian species,
being oblong to linear, often hastate, and with
entire to undulate margins. However, the
cauline leaves can vary greatly both between
and within species. In some species such as C.
remotus, C. microsepalus and C. arvensis, leaf
shape is similar from base to the tips of the
stems. In other species such as C. angustissimus
and C. clementii, considerable variation occurs
within a single plant. Lower cauline leaves are
often greatly different in both shape and size
from leaves found on fertile branches. In some
taxa this difference may have taxonomic
significance. This is particularly so within
C. angustissimus. Because of considerable
variation within individuals and within
populations, identification can be difficult from
single specimens taken from one part of a plant.
This problem is compounded by seasonal
conditions. Specimens taken from the same
population can vary greatly when collected in
the spring and late summer, while plants which

are rapidly forced into early flowering and
fruiting can produce inflorescences from the
axils of basal and lower cauline leaves.

Vestiture: Hairs are simple and tubercle-based.
They vary considerably in length and
orientation but there is some consistency in the
vestiture in individual species. Both C. remotus
and C. microsepalus have characteristically
short (< 0.4 mm long), appressed hairs. By
contrast the hairs on C. clementii and
C. angustissimus subsp. angustissimus when
present are commonly irregularly spreading and
often exceed 0.5 mm, and may even approach
1 mm in length. Both C. eyreanus and
C. crispifolius have dense silky semi-appressed
hairs. In many species, the vestiture changes
from that on basal leaves to that on fertile stems
with hairs more likely to be spreading in basal
parts and tightly appressed on terminal shoots.
Terms such as sparsely hairy, moderately hairy
and densely hairy are used in the text.
Moderately hairy is used when the length of
the hairs begins to exceed their distance apart
while densely hairy is used where the density
is such that the vestiture obscures the leaf
surface.

Inflorescence : All species have a simple or
compound dichasial inflorescence which is
axillary. Normally only one inflorescence is
found in a leaf axil but in many species such as

4

C. clementii, C. graminetinus, C. remotus,
C. arvensis, C. tedmoorei, C. recurvatus and
C. erubescens, occasionally two occur. Some
species, such as C. angustissimus and
C. microsepalus, almost always have a simple
inflorescence. In other species, such as
C. clementii , C. erubescens and C. arvensis, it
is common to find compound inflorescences
on most specimens. While bracteoles are paired
on all branches, it is only the outer one that
subtends a bud or flower. In the text, such
inflorescences are referred to as one-sided
dichasia. The bracteoles are usually opposite
in species with solitary flowers but in those with
one-sided dichasia they are often sub-opposite
to alternate, even when a single flower is
present.

Another distinguishing character is the
shape of the pedicel at fruiting. In some species,
such as C. angustissimus, C. wimmerensis and
C. recurvatus, the pedicel becomes strongly
recurved at fruiting while in other species, such
as C. erubescens and C. clementii, the fruiting
pedicel is straight or occasionally is sinuate or
rarely slightly curved. Aside from
C. graminetinus, a straight pedicel at fruiting
appears to be characteristic of northern species
with species restricted to southern areas all
possessing strongly recurved pedicels.

Calyx: The calyx consists of 5 free sepals,
quincuncially arranged. The outer two sepals
are similar or slightly unequal in size but they
are usually somewhat different in shape and /
or size from the inner pair. The intermediate
sepal is a hybrid in shape, the exposed half
resembling that of the outer sepals, the enclosed
half resembling the inner. The shape of the outer
sepals, in particular the apex, is diagnostic. Both
C. arvensis and C. microsepalus have small
outer sepals with a rounded, emarginate apex.
By contrast all other species have an acute to
rounded apex with a ± prominent recurved
apiculum.

Corolla : The corolla is typically funnel-shaped
with a narrow tube which flares distally into a
broad limb. The point at which it flares is
marked by five V-shaped structures where the
midpetaline bands diverge. The length of the
tube to the point where the corolla flares has
use as a possible discriminating attribute. The

Austrobaileya 6(1): 1-39 (2001)

length of the corolla and the diameter of the
limb are difficult to ascertain on herbarium
specimens because the corolla is funnel-shaped
and is flattened during pressing. The length of
the flattened and dried corolla approaches the
petal length and it is for this reason, petal length
has been used in the key to species and in
species descriptions.

Each petal has a distinct mid-petaline
band, hairy on the outside in the upper part and
which tapers towards the apex. The tube is
always glabrous on the outside. The colour of
the corolla varies from white to pale pink, or
occasionally purplish. As in other species of
Convolvulaceae, white-flowered forms have
been recognised e.g. Convolvulus erubescens
var. albus, but such formal recognition appears
trivial.

Stamens : The five stamens are affixed to the
corolla tube at the base and alternate with the
lobes. The lower part of the filament, attached
to the corolla tube, is flared downwards and
usually bears low tubercles on the margin. The
tubercles extend beyond the point of attachment
onto the inner face of the free upper part of the
filaments. The filaments are usually unequal in
length. The pollen grains are spheroidal or
ellipsoidal and 3-colpate. The colpi are linear
in shape and the surface of the pollen grains is
smooth (Fig. 2).

Ovary : In all species the ovary is borne on a
cup-shaped disk. The ovary is 2-celled with 2
basal ovules in each cell and is tipped by a style
with two + cylindrical stigmas. The nature of
the stigmas is a generic character.

Capsule: All species have globular to globular-
ovoid capsules with a distinct beak resulting
from the persistent style base. The capsules
dehisce loculicidally into 2 valves with each
valve often splitting further into 2 parts. The
capsule also dehisces at the base leaving a
persistent dissepiment. The size of the capsule
varies depending on maturity and seasonal
conditions but can be used to discriminate
among species.

Seeds: Under normal circumstances four seeds
are produced in each capsule, two in each cell.
The seeds are therefore ^-globular or globular-

Johnson, Convolvulus L. in Australia

5

Fig. 2. Pollen grain of Convolvulus remotus from Kraehenbuehl 2782 (AD).

ovoid in shape, with two flat faces and the
abaxial face concave. On occasions, fewer
seeds are produced and in these circumstances
the seed shape diverges from the usual pattern.
The size and surface sculpture are diagnostic.
Seeds of all species are illustrated in the text.
The underlying surface is finely and regularly
punctate but is usually covered with low
tubercles or ridges of irregular shape. A wing,
continuous or discontinuous, may or may not
occur. The characteristic patterning on the seed
surface appears to be produced towards
maturity and immature seed may lack any raised
surface structures. However, mature seed of
C. angustissimus subsp. fililobus appears to
have a smooth finely punctate surface without
surface structures. Though no species has hairy
seeds, the tubercles on the surface and wings
on the outer margins in some species, in
particular in C. clementii, appear to be formed
by the fusion of hair-like structures.

Hybridisation: While no study was made of
gene interchange among species, genetic traits
such as recurved pedicels and the one-sided
dichasial inflorescence do appear to flow into
populations of species which normally lack

them. In areas of geographical overlap where
both C. angustissimus subsp. angustissimus and
C. erubescens are found, strongly recurved
pedicels are occasionally found in populations
of C. erubescens.

Biogeography: The genus is primarily
temperate in distribution. The greatest diversity
of species in Australia is found in the south-east
with 8 species being recorded in Victoria, 8 in
South Australia and 10 in New South Wales.
By contrast only 2 species are known from the
Northern Territory, 5 from Western Australia
and 7 from Queensland. Convolvulus is one of
a few genera, including Calystegia and Wilsonia
R.Br., in Convolvulaceae which are concentrated
in temperate regions of Australia. Most genera
and species of this family occurring in Australia
are tropical and sub-tropical in distribution.

Within Convolvulus, three species are
common and widespread with C. angustissimus
occupying moister mainly southern areas of
Australia, C. clementii occupying arid and
semi-arid regions of all mainland states and
C. remotus more common in southern wetter
areas but extending into more arid areas, often

6

on moister sites. The remaining species, aside
from C. graminetinus which is found in
grasslands and grassy forests and woodlands
on mainly clay soils in semi-arid and sub-humid
Queensland and New South Wales, have
relatively restricted distributions.

Taxonomy

Convolvulus L., Sp.Pl. 1:153 (1753). Type:

Convolvulus arvensis L. (lecto: fide

Hitchcock 1929).

Annual or perennial herbs with erect, trailing
or twining stems, or in some extra-Australian
species shrubs or sub-shrubs, somet im es spiny;
stems hairy, more rarely ± glabrous; hairs
simple. Leaves simple, alternate, petiolate or
sub-sessile; blade very variable in shape, entire
to deeply divided, often hastate or sagittate,
with base usually cordate. Flowers axillary in
1- to few-flowered dichasia or in sub-umbellate
heads in some extra-Australian species. Sepals
5, free, with inner and outer pairs subequal and
the intermediate one asymmetric. Corolla
funnel-shaped or more rarely campanulate,
white, pink or mauve with 5 midpetaline bands

Austrobaileya 6(1): 1-39 (2001)

usually hairy outside towards the apex,
otherwise glabrous; limb sub-entire to
shallowly 5-lobed. Stamens 5, included;
filaments epipetalous, alternating with the
petals, terete above, flattened and dilated
downwards, often with low tubercles towards
the base; anthers bilocular, dehiscing
longitudinally; pollen spheroid or ellipsoid,
smooth. Ovary on a cup-shaped disk, bilocular,
each locule with 2 ovules; style simple, filif orm;
stigmas 2, cylindrical, obtuse. Capsule globular
or globular-ovoid, with a persistent style base,
2-4-valved, irregularly circumscissile at the
base but with a persistent dissepiment. Seeds 4
or less by abortion, ^-globular or ^-globular-
ovoid, with a finely punctate surface usually
bearing irregular tubercles or ridges.

Distribution: A genus of c. 150 species found
throughout the temperate and subtropical
regions of both hemispheres, rarely extending
into the tropics.

Etymology: The generic name refers to the
twining habit of most species being based on
the latin convolvere, to roll together or entwine.

Key to native and naturalised (*) species of Convolvulus in Australia

Because of the extreme variability in leaf shape from base to tip of the stem in some species, the leaf
shape used in the key refers to mid-cauline leaves, particularly those subtending inflorescences.

1. Leaves hastate or sagittate, with an entire, oblong or triangular-oblong,

occasionally linear, terminal lobe usually > 2 mm wide, and a pair of entire
or occasionally 2-toothed or 2-lobed basal lobes, ascending lobes absent,
rarely a tooth or a short lobe present on a few leaves at the junction of the

terminal and basal lobes.2

Leaves crenate, serrate to shallowly lobed or if a distinct terminal lobe is
present then basal lobes auriculate and toothed or much divided with
ascending lobes prominent.6

2. Sepals 4 mm or less in length, with a rounded to truncate, emarginate apex,

glabrous to sparsely hairy.3

Sepals 4-7 mm long, rarely shorter, with apex acute to rounded, apiculate.4

3. Petals 15-30 mm long; sepals 3-4 mm long; leaves glabrous or with a few

weakly erect hairs.1. *C. arvensis

Petals 5-10 mm long; sepals usually < 3 mm long; leaves moderately to

sparsely hairy with appressed hairs.2. C. microsepalus

Johnson, Convolvulus L. in Australia

7

4. Sepals and leaves with short appressed or crisped-appressed hairs, or if

hairs spreading then seeds with a fine pattern of low irregular tubercles;
inflorescence a 1-sided dichasium with 1-3 flowers; pedicels at

fruiting straight or sinuate, or if recurved, then petals 7-10 mm long.5

Sepals and leaves with spreading hairs and seeds with sparse low reticulate
ridges; inflorescence of solitary flowers; pedicel at fruiting recurved;
petals 8-25 mm long.9. C. angustissimus

5. Pedicels at fruiting recurved; capsule globular to 5.5 mm long; seeds Va-

globular in shape, to 3.5 mm in length, faces with a fine pattern of low
irregular tubercles; petals to 10 mm long; sepals usually ± glabrous to

sparsely hairy.3. C. graminetinus

Pedicels at fruiting straight or sinuate; capsule globular to globular-ovoid,

5.5-8.5 mm long; seeds ^-ellipsoid in shape, 3-4.8 mm long, faces with
irregular ridges; petals exceeding 10mm in length; sepals moderately to

densely hairy.4. C. remotus

6. Leaves crenate or serrate with basal lobes usually not prominent, sericeous;

hairs ± appressed, commonly > 0.5 mm long.7

Leaves with distinct terminal lobe and much divided basal lobes or basal
lobes auriculate and toothed, not sericeous; hairs usually <0.5 mm long.8

7. Peduncle at flowering < 12 mm long; seeds < 3 mm long; capsule < 4.5

mm in diameter. 5. C. crispifolius

Peduncle at flowering usually > 12 mm long; seeds > 3 mm long; capsule
5-6 mm in diameter.6. C. eyreanus

8. Corolla with petals > 9 mm long; seeds reticulate, tuberculate or smooth.9

Corolla with petals < 9 mm long; seeds tuberculate.12

9. Pedicel at fruiting straight or sinuate; inflorescence a 1-sided dichasium
with 1-3 flowers; peduncle slightly ribbed; margin of terminal lobe usually
undulate to lobed; basal lobes auriculate, dentate; seeds with many laterally

compressed sinuate tubercles.7. C. erubescens

Pedicel at fruiting strongly recurved.10

10. Faces of seeds with a fine pattern of low irregular tubercles; inflorescence

a 1-sided dichasium with 1 or 2 flowers; peduncle and stems often slightly

ribbed; petals to 10 mm long.3. C. graminetinus

Faces of seeds with sparse, low, reticulate ridges; flowers solitary; peduncle
and stems terete; petals 9-25 mm long. 11

11. Leaves densely hairy, silvery coloured, with mainly appressed hairs, with

ascending lobes often at least half the length of the terminal lobe.8. C. wimmerensis

Leaves moderately to sparsely hairy, or if densely hairy, then hairs
ascending or spreading, with ascending lobes rarely half the length of the
terminal lobe.9. C. angustissimus

12. Pedicel at fruiting recurved.13

Pedicel at fruiting ± straight to sinuate.14

13. Sepals ± glabrous to sparsely hairy; pedicel at fruiting 3-12 mm long;

seeds densely tuberculate, wingless.3. C. graminetinus

Sepals moderately to densely hairy, pedicels at fruiting 3-6 mm long;
seeds sparsely tuberculate, with a discontinuous wing of fused hair-like
structures.10. C. recurvatus

Austrobaileya 6(1): 1-39 (2001)

14. Stems slender; seeds 2.5-3.2 mm long, with wing present, often

discontinuous.11. C. clementii

Stems coarse; seeds 3.5-4 mm long with no obvious wing. 12. C. tedmoorei

1. Convolvulus arvensisL., Sp. PI. 1:153 (1753).

Type: Sweden, herb. Linn. 218.1 (lecto:

UNNXfide Meeuse (1957), p. 695.

Perennial with trailing and twining stems arising
from a well developed underground root system;
stems terete, narrowly winged, glabrous or
sparsely to very rarely moderately hairy,
glabrescent, hairs crisped to sinuate and semi-
erect, 0.2-0.5 mm long. Leaves petiolate, of
similar shape from base to tip of the stem (Fig.
3 A) though petiole becoming shorter and blade
smaller towards the tip; petiole 5-25 mm long;
blade ovate to oblong, often triangular-oblong,
hastate or sagittate, 15-60 mm long, 5-40 mm
wide, apex acute to rounded, sometimes
emarginate, mucronulate, base truncate to
cordate, basal lobes triangular, to 10 mm long,
acute to rounded, entire or very rarely with a
tooth on the lower margin, ascending lobes
absent, terminal lobe oblong, 12-55 mm long,
glabrous or with scattered hairs, hairs crisped
and spreading to weakly erect, 0.2-0.5 mm long.
Inflorescence solitary, axillary, bracteolate, a
one-sided dichasium, with 1-4 flowers or rarely
with 2 inflorescences per axil; peduncle terete,
with low ribs, 4-65 mm long, glabrous to sparsely
hairy, rarely moderately hairy, hairs loosely
crisped-appressed to weakly erect, 0.1-0.5 mm
long; bracteoles opposite to subopposite, linear
to narrowly obovate, 1.5-7 mm long, up to 2 mm
wide, apex acute to obtuse, mucronulate, eciliate
or with well developed cilia, sparsely hairy on
the back with hairs mainly on the midrib; pedicel
6-22 mm long, recurved at fruiting, hairs as for
peduncle. Outer sepals oblong, elliptic-oblong
to obovate-oblong, 3-4.2 mm long, rarely
shorter, 1.75-2.8 mm wide, apex rounded or
retuse, glabrous or moderately ciliate and ±
glabrous or sparsely hairy outside, with
occasional hairs along the midrib and near the
tip; inner sepals orbicular to obovate-orbicular,
3-4.5 mm long, 3-4.5 mm wide, apex retuse,
mucronulate, base rounded, ± glabrous. Corolla
funnel-shaped, white to pink, 15-25 mm long,
20-30 mm diameter, flared 4-7 mm above the
base of the tube; petals 17-30 mm long, 13-18

mm wide, with rounded, emarginate to shortly
bilobed lobes, glabrous except for scattered
hairs on the outside of the midpetaline band for
4-8 mm, occasionally up to 15 mm, from the
apex. Stamens 5, unequal; filaments affixed to
the corolla tube for 2.5-3.5 mm from the base,
free for 4.5-10 mm, with short cylindrical
tubercles to 0.2 mm long, mainly on the margins,
from 0.75-1.25 mm above the base of the corolla
and extending for 4-7 mm; anthers oblong to
oblong-elliptic, 2.5-3.2 mm long, 1-1.5 mm wide,
apex truncate, emarginate, base sagittate, basal
lobes 0.7-0.9 mm long. Ovary ovoid, 1.5-2.5 mm
long, on a lobed disk 0.5-0.8 mm high, glabrous
or hairy; style 8-9 mm long; stigmas cylindrical
to narrowly obovate, occasionally falcate,
obtuse, 2.5-4 mm long. Capsule globular to
globular-ovoid, 4—7 mm long, 4-6 mm diameter,
glabrous. Seeds 4, ^-globular-obovoid, 3^f mm
long, c. 3 mm wide, dark-brown to black, surface
finely punctate bearing a fine and ± regular
pattern of small, often laterally compressed,
tubercles, wingless (Fig. 4A).

Selected specimens (113 specimens examined):
Western Australia. Dwarda, Feb 1941, O’Connell
s.n. (PERTH); Boyup Brook, Jan 1949, O’Sullivan
s.n. (PERTH); South Coogee, Feb 1951, Quinlivan s.n.
(PERTH); Fremantle, Mar 1946, Royce s.n. (PERTH);
Bunbury, Dec 1952, Simpson s.n. (PERTH). South
Australia. Yallanda Flat, Feb 1964, Alcock 654A (AD);
Railway Terrace, Edwardstown, Nov 1987, Dashorst
s.n. (BRI); Fulham, c. 8 km W of Adelaide, Nov 1967,
Smith 984 (AD); Adelaide, Jan 1960, Symon 332 (NE).
Queensland. Leichhardt District: Trelinga, 2 miles
W of Wandoan, Mar 1972, Elphinstone s.n. (BRI).
Port Curtis District: Marlborough, Property of
GJ.A.Hack, Jun 1966, Kelly s.n. (BRI). Burnett District:
Monto, Oct 1951, Stubbs s.n. (BRI). Wide Bay District:
2 miles SW of Kilkivan, Feb 1970, Ditchmen s.n. (BRI).
Maranoa District: St George Irrigation, Jan 1969,
Hazard s.n. (BRI). Darling Downs District: Cooper Gully
road, NE of Yangan, Oct 1997, Bean 12426 (BRI).
Moreton District: 6 miles ESE of Gatton, Aug 1968,
Hazard s.n. (BRI). New South Wales. Carrol near
Gunnedah, Jan 1971, Dale s.n. (NSW); Henty, Feb 1949,
McBarron 3078 (NSW); Campbelltown, Feb 1962,
McBarron s.n. (NSW); Parkes, Mar 1925, Swann s.n.
(NSW); Armidale, Nov 1982, Wilson & Lapinpuro
LL11 (NSW). Victoria. Midlands: Avoca, 1894, Martin
s.n. (MEL). Tasmania. Sandy Bay, Hobart, Nov 1941,
Curtis s.n. (HO); New Town Research Laboratories,

Johnson, Convolvulus L. in Australia

9

Fig. 3. Variation in leaf shape of Convolvulus species from base to tip of stems. A. C. arvensis x 1. (1) Failes s.n.
(NE25445a), (2,3) Rodway s.n. (HO 15261); B. C. microsepalus. (1) Pfeiffer & Pfeiffer s.n. (AD96919556) x 2,
(2,3) Orchard 211 (AD) x 1; C. C. graminetinus. (1) McDonald 46 (BRI) x 0.5, (2) McDonald 46 (BRI) x 1, (3)
Fensham 2803 (BRI) x 1, (4) Fensham 1736 (BRI) x 1; D. C. remotus (1-3) Alcock 653G (AD) x 1. Del. W. Smith

Fig. 4. Seeds of Convolvulus species, adaxial (L.H.S.) and abaxial (R.H.S.) surfaces x 10. A. C. arvensis from Curtis
s.n. (H036313); B. C. microsepalus from Copley 192 (AD); C. C. graminetinus from Fensham 3332 (BRI); D. C.
remotus from Lothian 1214 (AD). Del. W. Smith.

10

Austrobaileya 6(1): 1-39 (2001)

Map 1. Distribution of #. Convolvulus arvensis. ★. C. microsepalus. Del. W. Smith.

Feb 1983, Morris 8338 (HO); Granton, Jan 1981,
Orchard 5278 (HO); Royal Park, Hobart, Mar 1961,
Somerville s.n. (HO); Hobart, Jan 1877, Spicer 118
(HO).

Distribution and habitat : C. arvensis is a native
of Eurasia but is now widespread throughout
the temperate regions of the world. In Australia,
it is a weed of cultivation and roadsides, often
on deep fertile clay soils. It has been recorded
from most of the major cultivation regions of
temperate Australia and northwards to the
Darling Downs and central Queensland (Map 1).

Affinities: Though it is not a native of Australia,
C. arvensis most closely resembles
C. microsepalus. Both species have sepals
which are small and similar in shape with a
blunt, somewhat emarginate apex, very
different from those in other Australian species
and both have hastate leaves with entire
margins. However, the latter has much smaller
flowers.

Phenology: Flowering occurs mainly from mid
spring to early autumn (October-March) with
fruiting mainly in early to mid autumn (March-
April).

Notes: The date of introduction of this species
into Australia is unknown. Bentham (1869)

makes no reference to C. arvensis in Australia.
Earliest herbarium records include Woolls s.n.
(MEL) collected in 1869 in New South Wales,
Wilson 12 (MEL) in 1883 in Victoria and Spicer
118 (HO) in 1877 in Tasmania. Woolls (1867)
in his paper on accidental plant introductions
does not mention C. arvensis.

Etymology: The specific epithet is from the
Latin arvensis meaning ‘ pertaining to fields
or cultivated lands’ which refers to the habitat
where the species is commonly found.

2. Convolvulus microsepalus R.W. Johnson,
Austrobaileya 2:410 (1987). Type: South
Australia, Flinders Ranges: c. 51 km N
of Quorn on the Quorn to Hawker road,
8 November 1970, A.E. Orchard 2626
(holo: AD; iso NCU, COLO, n.v.).

Perennial with trailing and twining stems; stems
slender, terete, moderately densely to sparsely
hairy, glabrescent, hairs appressed, 0.15-0.4 mm
long. Leaves similar in shape from base to tip of
the stem (Fig. 3B). Basal leaves petiolate; petiole
short, 2-7 mm long; blade oblong, slightly
hastate, 10-20 mm long, 4-5 mm wide, apex
obtuse to bluntly acute, mucronulate, base
truncate to shallowly cordate. Leaves on fertile
stems petiolate; petiole 3-8 mm long; blade
linear to oblong, sometimes triangular, hastate

Johnson, Convolvulus L. in Australia

or sagittate, 10-45 mm long, 3-20 mm wide,
apex acute to rounded-truncate with a short
recurved mucro, base truncate to cordate, basal
lobes 1-10 mm long, acute to obtuse, entire, 2-
toothed or lobed, ascending lobes absent,
terminal lobe oblong, linear to linear-triangular,
10-40 mm long, 3-8 mm wide, entire to slightly
undulate, moderately dense to sparsely hairy,
hairs silvery, appressed 0.2-0.4 mm long. In
upper parts, leaves similar but smaller.
Inflorescence solitary, axillary, bracteolate, with
solitary flowers; peduncle slender, terete, 8-30
mm long, densely hairy, hairs appressed, 0.3-
0.4 mm long; bracteoles opposite, subulate to
linear, 0.7-1.8 mm long, 0.4-0.5 mm wide,
apex acute, ciliate, moderately to densely hairy
on the back; pedicel 3-12 mm long, recurved
at fruiting, moderately to densely hairy, hairs
appressed 0.2-0.4 mm long. Outer sepals
obovate-elliptic to obovate, 2-3 mm long,
rarely to 4 mm, 2-3 mm wide, apex rounded to
truncate, ± emarginate, glabrous or with some
appressed hairs outside at the base and on the
basal margin; inner sepals orbicular to obovate,
2.5-3.5 mm long, 3-3.5 mm wide, apex
rounded to truncate, emarginate, mucronulate,
base rounded, glabrous. Corolla funnel-shaped,
white or pink, with a creamish-green throat, 5-
8 mm long, 5-13 mm diameter, flared 2-3 mm
above base of the tube; petals 6-9 mm long, 3-
6 mm wide, with rounded apiculate lobes,
glabrous except for a few hairs around the apex.
Stamens 5, slightly unequal; filaments affixed
to the corolla tube for 1-1.5 mm from the base,
free for 1.8-3 mm with low tubercles, mainly
along the margins, from almost the base of the
corolla and extending for up to 3 mm; anthers
oblong to triangular-oblong, 1-1.3 mm long,
0.5-0.75 mm wide, apex rounded, base
sagittate, basal lobes 0.2-0.25 mm long. Ovary
ovoid, 1.5-2 mm long, on a well developed
disk, 0.2-0.4 mm high, glabrous; style 1.5-2.5
mm long, with cylindrical, falcate, obtuse
stigmas, 1.2-2.3 mm long. Capsule globular to
globular-ovoid, 5-7 mm long, 5-7 mm diameter,
glabrous. Seeds 14-globular, 3.5-4 mm long, 3-
3.5 mm wide, dark brown to honey-coloured,
surface finely punctate bearing prominent
raised tubercles of fused hairs, usually laterally
flattened and of irregular shape, to 0.25 mm high,
with interrupted wing comprising fused hair¬
like structures (Fig. 4B).

11

Specimens examined : South Australia. Lake Eyre: 1
km S of Paradise Creek, c. 40 km S of Marree, Jun
1978, Badman 32 (AD). Funders Ranges: Parachilna
Gorge nr Mt Mary, c. 60 km N of Wilpena Pound, Oct
1987, Browne 463 (BRI); c. 16 km N of Hawker, Apr
1966, Copley 192 (AD); 8 miles E of Wilmington,
Mar 1959, Filson 809 (MEL, AD); 5.1 km N of Fred
Hughes (Eukaby) Gold Mine, Parachilna, Sep 1987,
Vonow 584 (BRI). Eastern: c. 6 km NE of Curnamona
(Arkipena Springs road), Apr 1968, Barker 454 (AD);
c. 3 km N of Curnamona Homestead, Apr 1968,
Orchard 211 (AD); Oak Park Homestead, c. 50 km S
of Yunta, Mar 1969, Pfeiffer & Pfeiffer s.n. (AD).
Northern Lofty: reserve between Gladstone and Laura,
Nov 1920, J.M.Black Herb. s.n. (AD). Murray: Upper
Murray Mallee, E of Sutherlands, c. 105 km NE of Adelaide,
Oct 1962, Boehm 376 (AD). New South Wales. 0.5 miles
W of Fowlers Gap Research Station, May 1954, Briggs
s.n. (NE); 42 km E of Broken Hill on Barrier Highway to
Wilcannia, Nov 1989, Palmer 282 (CANB). Victoria.
Wimmera: Wimmera, F. Mueller Herb. [Dallachyl ] (MEL
689646, 689644).

Distribution and habitat : C. microsepalus is
known mainly from eastern South Australia,
where it occurs in the Flinders Ranges and
surrounding areas. Two specimens of it on
sheets bearing F. Mueller labels were collected
from the Wimmera area. It has also been
recorded from south-western NSW (Map 1). It
grows in gravelly clay loam or loamy soils on
open plains carrying chenopod shrublands.

Affinities: C. microsepalus most closely
resembles C. remotus in leaf shape and
indumentum. However the sepals of C. remotus
are about twice as long, are apiculate, not +
emarginate, and are hairy. The corolla of
C. remotus is also much larger. The sepals of
C. microsepalus most closely resemble those
of C. arvensis but the latter has a corolla at least
twice as large.

Phenology: The main flowering period is from
late winter to early autumn but flowers have
occasionally been recorded at other times.
Capsules are produced from spring onwards.

Notes: C. microsepalus was first described as
Convolvulus sp. A in the Flora of South
Australia (Johnson 1986).

Etymology: The specific epithet refers to the
size of the sepals in this species which are by
far the smallest of any species of Convolvulus
recorded from Australia.

12

3. Convolvulus graminetinusR.W. Johnson, sp.
nov. affinis C. remoto R.Br. sed corollis et
sepalis minor, pedicellis ad fructificantem
recurvis et paginis seminum differt. Typus:
Queensland. Leichhardt District: Gregory
Mine Site (23°10’S 148°22’E), 50kmNEof
Emerald, 25 November 1998, R. W. Johnson
5300 (holo: BRI; iso: BRI, CANB, NE,
NSW).

Perennial with trailing and mainly twining
stems; stems terete often ribbed, moderately to
sparsely hairy, glabrescent, hairs crisped-
appressed, mainly 0.2-0.35 mm long, rarely
longer to 0.7 mm and more loosely appressed
to ascending. Leaves may or may not vary in
shape from base to tip of the stem (Fig. 3C).
Basal leaves petiolate; petiole long, often equal
to or longer than the blade; blade oblong-
triangular to linear-triangular, commonly
hastate, 24-40 mm long, 6-16 mm wide, apex
acute to rounded, mucronulate, base cordate,
basal lobes entire or 2-toothed, margin + entire
occasionally undulate, + glabrous to very
sparsely hairy, hairs crisped, appressed to
ascending, 0.25-0.5 mm long. Leaves on fertile
stems petiolate; petiole short, 2-10(-20) mm
long; blade linear, triangular-ovate to narrow
elliptic, hastate or sagittate, 15-60 mm long,
2-15 mm wide, apex acute, mucronulate, base
cordate to truncate, slightly decurrent, basal
lobes linear, 1—10(—15) mm long, entire, two¬
toothed or lobed, or with a short recurved tooth
or lobe on the lower margin, ascending lobes
absent or occasionally becoming prominent in
terminal parts, linear, to 25 mm long and 1-3
mm wide, margin + entire, terminal lobe
narrow-linear or narrow-elliptic to narrow-
oblong, oblong-elliptic or narrow-ovate, 20-55
mm long, 2-8 mm wide, margin + entire to
undulate, occasionally with a few lobes on more
basal leaves, ± glabrous to very sparsely hairy
above, rarely denser, sparse below, hairs
crisped-appressed to ascending, 0.1-0.5 mm
long. In upper parts, petiole and blade shorter
and lobes narrower. Inflorescence solitary,
axillary, bracteolate, a one-sided dichasium
with 1-2 flowers, or occasionally with 2
inflorescences per axil; peduncle terete, wiry,
7-55 mm long, moderately to sparsely hairy,
hairs crisped-appressed, 0.15-0.3 mm long;
bracteoles opposite to subopposite,
occasionally alternate, linear to subulate, 1-2

Austrobaileya 6 (1): 1-39 (2001)

mm long, 0.35-0.5 mm wide, apex acute, with or
without cilia, ± glabrous or very sparsely hairy
on the back; pedicel 3-12 mm long, recurved at
fruiting, hairs as for peduncle, sometimes
denser. Outer sepals obovate to elliptic or
oblong, 3-5(-5.5) mm long, 2.2-3.8 mm wide,
apex acute to obtuse-rounded with a short
recurved apiculum, + ciliate, + glabrous to
sparsely hairy, rarely denser, outside, hairs
crisped, ± appressed to ascending, 0.1-0.4 mm
long; inner sepals obovate, obovate-orbicular
to broadly elliptic, 3.5-5 mm long, 2.5-4 mm
wide, apex rounded, occasionally acuminate,
with a short recurved apiculum, base rounded
to truncate, glabrous or with a few hairs at the
tip. Corolla funnel shaped, pink, occasionally
white, with a greenish-cream throat, 6-10 mm
long, 7.5-15 mm diameter, flared 3-3.8 mm
above the base of the tube; petals 7-11 mm
long, 4-7 mm wide, with rounded-triangular,
apiculate lobes, glabrous except for hairs on
the outside of the midpetaline band for 2-3 mm
from the apex. Stamens 5, unequal; filaments
affixed to the corolla tube for 1.5-2.5 mm from
the base, free for 1.6-3.5 mm, with low
tubercles from 0.5 mm above the base of the
corolla and extending for 2-3.5 mm; anthers
oblong to ovate-oblong, 0.8-1.5 mm long,
0.55-0.85 mm wide, apex obtuse, base
sagittate, basal lobes 0.15-0.4 mm long. Ovary
ovoid, 1-1.3 mm long, on an undulate disk
0.25-0.4 mm high, glabrous; style 2.5-4 mm
long, with cylindrical, slightly flattened, obtuse
stigmas, 1.4-2.2 mm long. Capsule globular,
4-5.5 mm long and diameter, glabrous. Seeds
4, V^-globular to Vi-obovoid-globular, 2.5-3(-
3.5) mm long, 2-3 mm wide, dark brown to
black, surface finely punctate bearing low,
laterally flattened, wavy tubercles forming a
close regular pattern, wingless (Fig. 4C).

Selected specimens (64 specimens examined):
Queensland. North Kennedy District: 72 km NW of
Pentland, Jun 1993, Thompson & Figg HUG443 (BRI).
Mitchell District: Thornleigh Ck, stockroute Malverton
to Gowan Hills, 70 km W of Blackall, Nov 1975, Johnson
3017 (BRI). South Kennedy District: 150 km NW of
Clermont, Aug 1977, Dale 169 (BRI). Leichhardt District:
Peak Downs, Jun 1951, Everist 4378 (BRI); 13 km SE of
Capella, Mar 1995, Fensham 2803 (BRI). Burnett
District: Kingaroy, Oct 1949, Benham s.n. (BRI). Warrego
District: Morven, Apr 1936, Blake 10996 (BRI); Pinnacle
Station, W of Augathella, Mar 1982, Greenfield JT1090
(BRI). Maranoa District: 10 miles SE of Roma, Apr 1961,
Johnson 2075 (BRI). Darling Downs District: between

Johnson, Convolvulus L. in Australia

13

Map 2. Distribution of ★ . Convolvulus graminetinus. %.
C. erubescens.

Clifton and Allora, Nov 1946, Everist & Webb 1251
(BRI); 7 km SW of Toowoomba, Apr 1994, Fensham
1736 (BRI). Moreton District: Cowley Vale, 16 miles E
of Helidon, Aug 1969, Schroder s.n. (BRI). New South
Wales, c. 13 km WSW of Moree on road to
Collarenebri, Sep 1975, Henderson H2352 (BRI);
Tamworth District, Feb 1967, Loveridge s.n. (NSW);
Twenty-one Mile Warrambool between Walgett and
Collarenebri, Nov 1967, McGillivray 2785 (NSW); 5.5
km W of North Star, near entrance to “Mungle”, Sep
1988, Moore 8792 (CANB); 4 km S of Melton Grove
on Darnick road, just N of Willandra, May 1994,
Porteners & Benson 9405019 (NSW); 10 miles NE of
Yetman, Nov 1952, SWQS 1317 (BRI); Iolanthe, c. 25
km SW of Garah, Apr 1976, Wilson 1470 (NSW).

Distribution and habitat : C. graminetinus is
widespread throughout the semi-arid and
subhumid regions of Queensland and New
South Wales though it extends into coastal areas
in south-east Queensland (Map 2). It occurs
mainly on clay soils on alluvial plains with
Eucalyptus coolabah and E. tereticornis and
on rolling downs, often derived from basalt,
dominated by Dichanthium sericeum. In north¬
eastern NSW it is found in Eucalyptus albens
woodlands on clay to clay loam soils and in
Acacia harpophylla - Casuarina cristata
woodlands. In drier areas it occurs on clay soil
downs with chenopods.

Affinities: C. graminetinus appears most
closely related to C. erubescens with its similar
crisped appressed hairs, and its tendency for

the stems and peduncles to be slightly ribbed.
Its seeds have a similar surface pattern and the
inflorescence tends to be a one-sided dichasium
with 1 or 2 flowers. C. erubescens grows in
moister Eucalyptus forests and on the edges of
rainforest whereas C. graminetinus is found in
grasslands and marginal brigalow woodlands
on heavy clay soils in sub-humid to arid areas.
Plants with hastate-sagittate leaves resemble
those of C. remotus. However, C. graminetinus
has smaller and less hairy sepals, smaller
flowers and the seeds are quite different in size,
shape and surface pattern.

Phenology: Flowering and fruiting occurs
throughout the summer to early winter
(N o vember-June).

Notes: Throughout central Queensland,
populations of C. graminetinus tend to have
hastate leaves, + glabrous sepals and crisped
appressed hairs though plants with leaves
bearing prominent ascending lobes are found.
Similar populations are found through the north
western plains of New South Wales with the
presence of ascending lobes on leaves
becoming more common. However, in the north
western slopes region of that state, some
populations, such as Moore 9108 (CANB) have
many spreading hairs on leaves, stems and
sepals. Aside from this character, the leaves are
often deeply divided with well developed
ascending lobes. Two specimens Hoskings
1608 and 1681 (NSW) from the same
population in Oxley Park, Tamworth are
revealing. Hoskings 1608 collected in October
1998 features typical short crisped appressed
hairs and + glabrous sepals in contrast to the
specimen collected in February 1999 which
bears moderately dense spreading hairs on
leaves and sepals similar to those in Moore
9108. A specimen collected by J. Crawford from
near Bingera (CBG12272) contains two separate
branches, each branch representing the two
different forms described above. The specimen,
Portenters & Benson 9405019 , was collected
at a distance from the main area of distribution
of this species. It has longer hairs, longer
bracteoles, much hairier sepals, commonly 2
inflorescences per axil and larger seeds than do
the other specimens. Wilson 1470 also has large
seeds but it otherwise agrees morphologically
with specimens of C. graminetinus from
Queensland.

14

This species has been recorded as a weed
of cultivation on clay soils on the Darling
Downs and Central Highlands of Queensland.

Etymology : The specific epithet is derived from
the latin graminetum, meaning grassland, and
-inum, belonging to. This refers to the most
common habitat where this species occurs.

Conservation Status: This species is
widespread and not endangered at present.

4. Convolvulus remotus R.Br., Prod.: 483
(1810). Type: Australia: South Coast
[Port Lincoln, 4 March 1802], R. Brown
(holo: BM; iso: MEL[MEL689915]).

Convolvulus preissii de Vriese in Lehmann,
Plantae Preissianae 1:346 (1845). Type:
Ad promontorium Cape Riche, 21 Nov
1840, Herb. Preiss no. 1927 (holo: LD).

Convolvulus huegelii de Vriese in Lehmann,
Plantae Preissianae 1: 346 (1845). Type:
In solo limoso haud longe a praedio
rustico Maddington, ad flumiun
Canning River, 2 Nov 1839, Herb.
Preiss no. 1928 (holo: LD).

Perennial with twining, sometimes trailing,
stems; stems terete, sparsely to densely hairy,
hairs short, appressed 0.2-0.4(-0.6) mm long.
Leaves somewhat variable in shape and size
from base to tip of the stem (Lig. 3D). Basal
leaves petiolate; petiole often longer than the
blade, 12-20 mm long; blade triangular,
triangular-oblong to triangular-ovate, bluntly
hastate to sagittate, 10-30 mm long, 3-12 mm
wide, apex acute to rounded, sometimes
emarginate, mucronate, base truncate,
decurrent, terminal lobe entire to slightly
undulate, sparsely hairy above, slightly denser
below, hairs appressed, rarely spreading, 0.1-
0.4(-0.5) mm long. Leaves on fertile stems
petiolate; petiole 2-20 mm long; blade ovate
to triangular or triangular-oblong, bluntly
hastate to auriculate-sagittate, 10-80 mm long,
5-40 mm wide, apex acute to rounded,
sometimes emarginate, mucronate, base
cordate, rarely truncate, basal lobes spreading
or recurved, to 10 mm long, obtuse, entire,
sometimes 2-toothed, occasionally with a
distinct recurved lobe from the lower margin,
very rarely with a short ascending lobe, terminal
lobe linear or oblong, often triangular,

Austrobaileya 6(1): 1-39 (2001)

occasionally narrow elliptic, to 67 mm long,
2-17 mm wide, entire or rarely slightly
undulate, moderately to densely hairy above,
more rarely sparsely hairy, similar to slightly
denser below, hairs appressed, 0.1-0.4(-0.5)
mm long, silvery. In upper parts, petiole shorter
and blades smaller with narrower lobes.
Inflorescence solitary, axillary, bracteolate, a
one-sided dichasium, with 1-2, rarely 3,
flowers, or rarely with 2 inflorescences per axil;
peduncle filiform, 5-42 mm long, moderately
to densely hairy, hairs appressed, 0.2-0.4 mm
long; bracteoles opposite to sub-opposite,
subulate to linear, 1.3-3 mm long, 0.25-0.5 mm
wide, apex acute, ciliate, moderately to sparsely
hairy on the back; pedicel 3-16 mm long, hairs
as for peduncle. Outer sepals broadly elliptic
to almost orbicular, occasionally slightly
obovate, 5-6.5(-7) mm long, 3.5-4.5(-5.5) mm
wide, apex obtuse to rounded, often shortly
apiculate, mucronate, usually ciliate,
moderately to densely, rarely sparsely, hairy
outside, hairs mainly appressed, 0.2-0.4(-0.6)
mm long; inner sepals broadly elliptic to almost
orbicular, often obovate-elliptic, rarely ovate-
elliptic, 4.2-6 mm long, 3-4.5 mm wide, apex
rounded, shortly apiculate, mucronate, base
truncate, face glabrous to sparsely hairy. Corolla
funnel-shaped, pink to pale pink, occasionally
white, mauve or reddish-purple, midpetaline
band brownish-yellow on the outside, throat
creamish-green, 8-12 mm long, 8-20 mm
diameter, flared 3.5-5 mm above the base of
the tube; petals 9-18 mm long, 4-11 mm wide,
with rounded-triangular, emarginate lobes,
often with a short apiculum, glabrous except
for hairs on the outside of the midpetaline band
for up to 5 mm from the apex. Stamens 5,
slightly unequal in length; filaments affixed to
the corolla tube for 1.5-3.5 mm from the base,
free for 2-5 mm, with low tubercles from just
above the base of the corolla and extending for
2.5-5 mm; anthers oblong to triangular-oblong,
1-2 mm long, 0.6-1 mm wide, apex rounded
to emarginate, base sagittate, basal lobes 0.2-
0.5 mm long. Ovary ovoid, 1-2.5 mm long, on
a prominent disk, 0.3-0.5 mm high, glabrous;
style 3.5-6 mm long, with cylindrical to slightly
obovate, obtuse stigmas, 1.3-2.5 mm long.
Capsule globular to globular-ovoid, 5.5-8.5
mm long, 5-7 mm diameter, glabrous. Seeds
4, ^-ellipsoid or Vi-globular-ellipsoid, 3-4.8
mm long, 2-3 mm wide, brown to dark brown

Johnson, Convolvulus L. in Australia

15

often with irregular darker patches and stripes,
surface finely punctate bearing low irregular ±
anastomosing ridges and tubercles and a narrow
+ continuous to broken wing on the outer
margins (Fig. 4D).

Selected specimens (309 specimens examined): Western
Australia. 201 miles E of Kalgoorlie on Trans-Australian
Railway, Jun 1964, Aplin & Trudgen 5762 (PERTH); 1.3
km S on Keating road. Glittering, Dec 1981, Cranfield 1980
(PERTH); 97 km NW of Forrest, Apr 1984, Downing 919
(PERTH); Rocky Pool, Gascoyne River, c. 850 km N of
Perth, Oct 1975, Kenneally 4643 (PERTH). Northern
Territory. Palm Valley, Jul 1965, Beauglehole 10409 (BRI);
Bond Gap, Simpsons Gap National Park, Nov 1980, Latz
8530 (DNA, BRI); Mt Benstead Creek, c. 50 km ENE of
Alice Springs, Jun 1984, Latz 9928 (DNA, BRI). South
Australia. 12 miles N of Bordertown, Nov 1964,
Beauglehole 19821 (AD); c. 5 km NE of McLaren Flat, c.
30 km S of Adelaide, Dec 1976, Bell 75 (AD); Beresford
Hill, Oct 1978, Chorney 991 (AD); 25 km E of Watson,
Aug 1980, Weber 6588 (AD); c. 1 km N of Nudlamutana
Well, c. 20 km N of Balcanoona, Oct 1967, Whibley 2180
(AD).Queensland. “Budgerygar”, 64 km SW of Yaraka,
Nov 1975, Johnson 3045, 3111 (BRI). New South Wales.
Delta road junction, Sturt Highway, E of Wentworth, May
1979, Fox 7905071 (NSW); NE edge of Narran Lake,
Brewarrina, Nov 1967, McGillivray 2856 (NSW); 26 miles
N of Wentworth on road to Broken Hill, Aug 1969, Rodd
s.n. (NSW); Depot Glen, 12kmNofMilparinka, Sep 1990,
Wilson 1646 (NSW). Victoria. Grampians: Mt Arapiles, S
side, upper reaches of N arm of golf course gorge, Nov
1968, Beauglehole 29687 (MEL). Murray Mallee: Lake
Hindmarsh Reserve, Dec 1986, Beauglehole 87629 &
Huebner (MEL); Lake Wallawalla area, c. 4 km NW of
causeway on Lake Wallawalla, Dec 1988, Browne 566
(LTB, BRI); Thurla, South Cardross Lakes, Dec 1964,
Chandler ACB19687 (MEL).

Distribution and habitat : C. remotus occurs
mainly south of the Tropic of Capricorn in all
mainland states. It is also absent from eastern
coastal areas (Map 3). It occurs on a wide
variety of soil types from clays through loams
to sands. In more arid areas, it occurs on
sandhills in Zygochloa grasslands as well as on
alluvial soils along drainage lines. It is
commonly found in chenopod shrublands with
bluebush and saltbush. In semi-arid areas, it has
been recorded from Acacia shrublands,
including mulga, and from open mallee
woodlands and heaths.

Affinities: C. remotus does not appear to be
closely related to the other Australian species.
As noted under C. graminetinus, sterile
specimens can resemble those of that species.

Phenology: Flowering occurs throughout the
year but mainly in spring and early summer
(August-December); fruit are found mainly in
spring to early autumn.

Etymology: Unknown

Conservation Status: This species is
widespread and not endangered at present.

5. Convolvulus crispifolius F. Muell., Linnaea
25:423 (1853), (as “crispifolias”). Type:
South Australia. In montibus nudis petraeis
aliquot milliaria Anglica directione boreali-
orientali a Cudnaka, October 1851, F.
Mueller s.n. (holo: MEL[MEL1544962];
iso: MEL[MEL689518]).

Perennial with short trailing stems, rarely to 1
m long; stems wiry, terete, moderately to
densely hairy, becoming less densely hairy with
age, hairs appressed to loosely ascending, 0.2-
1 mm long. Leaves somewhat variable in shape
and size from base to tip of the stem (Fig. 5 A).
Basal leaves petiolate; petiole long, often longer
than the blade; blade ovate, oblong-ovate to
triangular-ovate, lacking distinct basal lobes,
10-20 mm long, 4-18 mm wide, apex obtuse
to rounded, base truncate to shallowly cordate,
margin crenate, moderately to densely hairy,
hairs appressed, 0.15-0.6 mm long. Leaves on
fertile stems petiolate; petiole short, 2-15 mm
long; blade ovate, oblong-ovate to triangular-
ovate, sometimes slightly hastate, 5-25 mm
long, 4-15 mm wide, apex acute to truncate,
base cordate, + decurrent, margin unevenly
crenate to bluntly serrate, with 7-10 teeth per
side, more deeply indented towards the base or
with short basal lobes, moderately to densely
hairy, hairs appressed, 0.3-0.8 mm long. In
upper parts, basal and ascending lobes
becoming more prominent but rarely exceeding
3 mm in length. Inflorescence solitary, axillary,
bracteolate, a one-sided dichasium with 1-2
flowers; peduncle terete, 2-12 mm long, rarely
extending to 20 mm at fruiting, moderately to
densely sericeous, hairs appressed to loosely
ascending, 0.2-0.5 mm long; bracteoles
opposite, linear-subulate to narrowly ovate,
1.2-2.2 mm long, 0.25-0.3 mm wide, apex
acute, ciliate, densely hairy on the back; pedicel
1.5-4 mm long, recurved at fruiting, hairs as

16

Austrobaileya 6(1): 1-39 (2001)

Fig. 5. Variation in leaf shape of Convolvulus species from base to tip of stems A. C. crispifolius. (1) Copley 571 (AD) x 1,
(2,3) Chinnock 2915 (AD) x 2; B. C. eyreanus. (1) Kuchel 884 (AD) x 1, (2) Hill 163 (AD) x 0.5; (3) Hill 163 (AD) x 1;
C. C. erubescens. (1) Johnson & Pedley 453 (BRI) x 0.5, (2) Simmonds s.n. (BRI-AQ275832) x 0.5, (3) Hind s.n.
(NSW198642) x 0.5, (4) McBarron 12570 (NSW) x 2; D. C. wimmerensis x L (1) Beauglehole ACB83982 (MEL) (2,3)
Beauglehole ACB86599 (MEL). Del. W. Smith.

Johnson, Convolvulus L. in Australia

17

for peduncle. Outer sepals obovate to obovate-
elliptic, 4—4.5 mm long, 3-3.7 mm wide, apex
rounded-obtuse with a distinct recurved
apiculum, moderately to densely ciliate,
sericeous outside, hairs appressed to loosely
ascending, 0.25-0.7 mm long; inner sepals
obovate to obovate-elliptic or obovate-
orbicular, 3.6^1.3 mm long, 3-4 mm wide, apex
rounded or truncate, shortly apiculate, base
obtuse to truncate, glabrous or with hairs on
the midrib. Corolla funnel-shaped, white to
pink, with a whitish-green throat, 5-6 mm long,
6-8 mm diameter, flared 3-4 mm above the
base of the tube; petals 6-8 mm long, 2.5-3.5
mm wide, with irregularly rounded, bluntly
apiculate lobes, glabrous except for sericeous
hairs on the outside of the midpetaline band
for 2-2.5 mm from the apex. Stamens 5,
unequal; filaments affixed to the corolla tube
for 1.5—2 mm from the base, free for 1.5-2.8
mm, with low tubercles, mainly along the
margins, from almost the base of the corolla
and extending for 2-3 mm; anthers oblong to
ovate, 0.65-0.75 mm long, 0.5-0.7 mm wide,
apex truncate or rounded, sometimes
emarginate or apiculate, base sagittate, basal
lobes 0.1-0.15 mm long. Ovary ovoid to ovoid-
elliptic, 1-1.25 mm long, on a distinct disk
0.25-0.3 mm high, glabrous; style furrowed,
2-2.5 mm long, with cylindrical to narrowly
ellipsoid, obtuse stigmas, 1.2-1.5 mm long.
Capsule globular to globular-ovoid, 4-4.5 mm
long, 4-4.5 mm diameter, glabrous. Seeds 4, l A
globular, 2.4-3 mm long, 1.75-2 mm wide,
dark brown to black, surface finely punctate
bearing low, short irregular sinuate ridges and
a narrow, + continuous wing of fused hair
follicles, 0.1-0.15 mm wide (Fig. 6A).

Specimens examined : South Australia. Flinders
Ranges: Wilpena Pound, Sep 1989, Bates 20915 (BRI).
Eyre Peninsula: Hundred of Hawker, Lincoln Highway,
c. 16 km S of Elbow Hill, Jul 1965, Alcock 652 (AD);
Hundred of Hambidge, Flora & Fauna Reserve,
Prominent Hill, NE of Loch, Sep 1965, Alcock 1063
(AD); Lincoln Highway, N of Elbow Hill, Nov 1965,
Alcock, s.n. (AD); Hambidge Flora & Fauna Reserve,
W of Prominent Hill, NE of Lock, Oct 1966, Alcock
1151 (AD); Crown lands, WNW of Kimba, Oct 1981,
Alcock 9001 (AD); 23 km NE of Poochera, N of
Karoultaby, Oct 1975, Chinnock 2915 (AD); Mount
Ive, Gawler Ranges, c. 160 km W of Port Augusta, Sep
1969, Donner 3242 (AD); between Cowell and Arno
Bay, Nov 1961, Kraehenbuehl 526 (AD); junction of
Sections 24 & 14, Hundred of Verran, c. 95 km NNE of
Port Lincoln, Oct 1963, Kuchel 1470 (AD); County

Buxton, Pinkawillinie, c. 36 km WNW of Kimba, Feb
1959, Rohrlach 166 (AD); Price Beach, Oct 1983
Toelken 7741 (BRI); c. 5 km NE of Corrobinnie Hill,
Oct 1981, Weber 6932 (AD). Yorke Peninsula: Hundred
of Wiltunga, NW comer of Sect 168, c. 140 km NNW
of Adelaide, Aug 1966, Copley 571 (AD); Sect 200,
Hundred of Wiltunga, c. 140 km NNW of Adelaide,
Nov 1966, Copley 876 (AD); Ardrossan, c. 80 km NW
of Adelaide, Tate s.n. (AD). Murray: Berri, Jun 1921
J.M.Black Herb s.n.(AD); Berri, Jan 1921, Cleland
s.n.(AD); Mantung District, c. 140 km ENE of Adelaide,
Aug 1924, Cleland s.n. (AD); between Overland comer
and Barmera, Sep 1965, Eichler s.n. (AD). Victoria.
Murray Mallee: Sunset Country, Pheeny’s Track, c.10
km W of S. Bambill Track, Oct 1981, Browne 536
(BRI).

Distribution and habitat : C. crispifolius is
found south of 32°S in the southern part of the
Flinders Ranges and in both the Eyre and Yorke
Peninsulas of South Australia. Its distribution
extends eastward into the Murray-Mallee area
and into far western Victoria (Map 4). It occurs
on sandy and sandy-loam, sometimes rocky,
soils, commonly in mallee scrubs, and on
sandhills.

Affinities: C. crispifolius appears most closely
related to C. eyreanus. Johnson (1987), in
describing C. eyreanus , referred to a form of
that species with small serrate, silvery coloured
leaves which he considered may prove
taxonomically distinct. This form is now
considered referable to C. crispifolius. The latter
species can be distinguished from C. eyreanus
by its small crenate, silvery coloured leaves,
its shorter pedicels and its smaller seeds and
capsules. C. eyreanus has a more vigorous
twining habit; branches from the crown in
C. crispifolius are mainly prostrate and non¬
twining.

Phenology: Flowering has been recorded mainly
from spring to early summer with fruiting
extending into the late summer.

Notes: Though this species was described in
1853, the name has rarely appeared in print.
Strangely, Bentham (1869) did not take account
of the name and appeared unaware of its
publication. Under C. erubescens Sims he
remarked on a remarkable form or variety “.. with
the leaves very densely tomentose and much-
cut and crisped and the peduncles very short
from Cudnaka” which had been collected by
Mueller. This specimen had been selected by
Mueller as the type for the name of his species,

Austrobaileya 6 (1): 1-39 (2001)

Fig. 6. Seeds of Convolvulus species, adaxial (L.H.S.) and abaxial (R.H.S.) surfaces x 10. A. C. crispifolius from Chinnock
2915 (AD); B. C. eyreanus from Donner 3531 (AD); C. C. erubescens from Johnson 1801 (BRI); D. C. wimmerensis from
Beauglehole ACB82670 (MEL). Del. W. Smith.

Map 4. Distribution of •. Convolvulus crispifolius. ★. C. eyreanus. Del. W. Smith.

Johnson, Convolvulus L. in Australia

19

C. crispifolius . Mueller (1864) wrote of the
extreme variation existing within C. erubescens
and reduced C. crispifolias (now spelt as C.
crispifolius) to synonomy under C. erubescens.
I was not aware of its existence until after C.
eyreanus was described.

Etymology : The specific epithet refers to the
characteristically unevenly crenate to bluntly
serrate margins of the leaves.

6. Convolvulus eyreanus R.W. Johnson,
Austrobaileya 2:408 (1987). Type: South
Australia: Frome East, c. 40 km ENE of
Frome Downs Homestead, 23 July 1971,
N.N. Donner 3531 (holo: AD; iso: Altona
Springs, Oshkosh, n.v.).

Perennial with trailing and twining stems; stems
terete, sericeous, moderate to densely hairy on
younger parts becoming less dense with age,
hairs appressed to ascending, 0.2-0.7 mm long.
Leaves variable in shape and size from base to
tip of the stem (Fig. 5B). Basal leaves petiolate;
petiole 3-25 mm long; blade oblong to oblong-
triangular, barely hastate, 8-20 mm long, 3-18
mm wide, apex truncate, mucronulate, base
truncate to barely cordate, margin undulate,
toothed or lobed, with 6-7 lobes/side,
moderately to densely hairy, hairs + appressed,
0.2-0.7 mm long. Leaves on fertile stems
petiolate; petiole 1—10(—12) mm long; blade
ovate to oblong, hastate or sagittate, 7-35 mm
long, 3-20 mm wide, apex acute to rounded,
mucronulate, base cordate, margin serrate to
shallowly lobed, basal lobes often more
prominent, 2-10(-20) mm long, entire to 2 or
3-lobed, ascending lobes linear, 3-16(-22) mm
long, entire to shallowly lobed, terminal lobe
linear to oblong, crenate to shallowly lobed with
5-8 lobes or rounded teeth, densely sericeous
and silvery on both sides, hairs 0.2-0.6 mm
long, + appressed. On terminal branches, basal
lobes becoming more prominent with ascending
lobes to 7-8 mm and basal 2-3 mm long, often
with a secondary recurved 3-toothed lobe.
Inflorescence solitary, axillary, bracteolate, a
one-sided dichasium with 1 or 2, rarely 3
flowers; peduncle slender, terete, 10-30(-60)
mm long, moderately to densely hairy, hairs
appressed to ascending, occasionally erect, 0.2-
0.5 mm long; bracteoles opposite to
subopposite, linear to subulate, 1-3 mm long,

0.3-0.5 mm wide, apex acute, moderately to
densely ciliate, cilia 0.25-0.5 mm long,
moderately to densely hairy on the back, hairs
+ appressed to ascending, 0.2-0.5 mm long;
pedicels terete, 1-10 mm long, hairs as for
peduncle. Outer sepals obovate, obovate-
elliptic to obovate-oblong, 4.2-5.5 mm long,
3-4 mm wide, apex bluntly acute to rounded
with a short recurved apiculum, moderately
ciliate, cilia 0.3-0.7 mm long, sparsest at the
tip, moderately to densely hairy outside, hairs
+ appressed to ascending, 0.2-0.6 mm long;
inner sepals obovate, 3.2-4. 5 mm long, 3-3.7
mm wide, apex rounded, abruptly acuminate,
with a short recurved apiculum, base rounded
to obtuse, glabrous or with occasional hairs
mainly on midrib and at base. Corolla funnel-
shaped, off-white to pink, 6-8 mm long, 6-10
mm diameter, flared 3.5-4 mm above the base
of the tube; petals 8-9 mm long, 3.7-4.5 mm
wide, with well developed rounded-triangular
and obtuse lobes, glabrous except for some
appressed hairs on the outside of the
midpetaline band for 2.5-3 mm from apex.
Stamens 5, unequal; filaments affixed to the
corolla tube for 1.8—2 mm from the base, free
for 2-4.3 mm, with low tubercles to 0.1 mm
long, mainly along the margins, from almost
the base of corolla and extending for 2-3 mm;
anthers oblong to triangular-ovate, 0.75-0.95
mm long, 0.55-0.7 mm wide, apex obtuse to
rounded, apiculate, base sagittate, basal lobes
0.1-0.2 mm long. Ovary ovoid, c. 1 mm long,
on a disk 0.2-0.3 mm high, glabrous or with
an occasional hair in the upper part; style 2.7-
3 mm long, glabrous or hairy, with cylindrical
to very narrowly ellipsoid, occasionally falcate,
stigmas, 1-1.4 mm long. Capsule globular to
globular-ovoid, 6-7 mm long, 5-5.5 mm diameter,
glabrous or with a few hairs at the apex. Seeds
4, ^-globular, 3.2^4.4 mm long, 2.3-3 mm wide,
dark brown, surface finely punctate bearing
irregular shaped tubercles or short wavy ridges,
with a narrow discontinuous to + continuous
wing, 0.1-0.2 mm wide (Fig. 6B).

Selected specimens (43 specimens examined): South
Australia. Lake Eyre Region: Far N Lake Eyre, Central
Hunt Peninsula, just N of Muloorina H/S, Sep 1968,
Cornwall 109 (AD); Mt Gason Bore, Birdsville Track,
c. 250 km NE of Marree, Sep 1960, Filson 3330 (AD,
MEL); Wirragalpina Swamp, c. 46 km WSW of new
Stuart H/S, Mar 1984, Haegi 3353 (BRI); Muloorina
Station between Station and Lake Eyre, Jul 1955, Hill

20

163 (AD); Birdsville Track nr camp at Lake
Palankarinna, c. 30 km N of Dulkaninna H/S, Mar
1972, Jackson 1901 (AD); Mulka Bore Ruins, 3 miles

5 of new H/S, c.155 km NNE of Marree, Aug 1960,
Lothian & Francis 280 (AD); Ooroowilanie, c.165 km
NNE of Marree, Jul 1960, SA Pastoral Board s.n. (AD);
just S of Strangways Railway Siding, 53 km E of William
Creek, Mar 1983, Weber 8851 (BRI). Funders Ranges
Region: Oraparinna National Park, central portion, c.

6 km SSW of headquarters, Jackson 1767 (AD); c. 25
km S of Moolawatana Station, c. 140 km ENE of
Leigh Creek, Aug 1963, Kuchel 884 (AD); c. 8 km W
of Yadlakenna Dam, between Myrtle Springs and
Termination Hill, Nov 1964, Lothian 3385 (AD); Lake
Torrens East, Motpena, c. 24 km WSW of Parachilna,
Aug 1955, SA Pastoral Board s.n. (AD); Ideyaka, Sep
1883, Tate s.n. (AD). Gairdner-Torrens Region: Mulgaria,
Aug 1955, SA Pastoral Board s.n. (AD); Lake Torrens
Basin, c. 15 km W of Yadlakenna Well, c. 30 km NW
of Leigh Creek, Nov 1964, Lothian 3428 (AD). Eastern
Region: Frame Downs Station, Oct 1971, Trezise 348
(AD); 5 km N of North Mulga Outstation on pipeline,
Sep 1987, Vonow 702 (BRI); Lake Frame East, Billeroo
Creek Area, c. 45 km ENE of Frame Downs Homestead,
Jul 1971, Whibley 3455 (AD). Queensland. Gregory
South District: 3 km S of Birdsville via old cement
crossing road heading S, Sep 1995, Edmunds AD 149
(BRI).

Distribution and habitat: C. eyreanus occurs
throughout the north-eastern parts of South
Australia in the basins of Lake Eyre, Lake
Torrens and Lake Lrome. It also grows in the
Simpson Desert extending into Queensland,
south of Birdsville (Map 4). It is found mainly
on sand dunes and associated habitats, often
growing in Acacia shrublands.

Affinities : C. eyreanus is most closely related
to C. crispifolius. It also resembles C. clementii
but can be distinguished from that species by
its more sericeous vestiture, more shallowly
lobed leaves and its seed size and surface
architecture.

Phenology: Llowering and fruiting occurs
mainly from the late winter to early summer.

Etymology: The specific epithet refers to the
name Eyre in Lake Eyre and the Eyre Region
of South Australia, both named in honour of
the Australian explorer, Edwin John Eyre
(1815-1901).

Conservation Status: This species appears to
be widespread in South Australia but is known
from only one collection in Queensland.

Austrobaileya 6(1): 1-39 (2001)

7. Convolvulus erubescens Sims, Curtis’s
Botanical Magazine 27: t. 1067 (1807)
Type: NSW, Loddiges s.n. (holo: not
found).

Convolvulus erubescens var. dilatatus
Choisy in A.DC., Prodr. 9:412 (1845).
Type: 1 ‘Varietas e cultura nota (described
from living plants in the wild - no type)
(v.v.)”.

Perennial with trailing and twining stems; stems
terete, ribbed to narrowly winged, moderately
to sparsely hairy, becoming sparser with age,
hairs mainly crisped-appressed, 0.2-0.35(-0.5)
mm long. Leaves variable in shape and size
from base to tip of the stem (Fig. 5C). Basal
leaves petiolate; petiole long, often equal to or
longer than the blade, 25-35 mm long; blade
triangular-ovate, sagittate, 20-45 mm long, 15-
27 mm wide, apex obtuse, mucronulate, base
cordate, margin undulate to shallowly lobed,
basal lobes not prominent, + glabrous above,
moderately hairy below, hairs crisped, 0.15-
0.4 mm long. Leaves on fertile ste ms petiolate;
petiole 2-25 mm long; blade triangular to
oblong-triangular, occasionally linear,
auriculate to sagittate, 15-60 mm long, 2-40
mm wide, apex acute to obtuse or rounded,
occasionally emarginate, apiculate, base
cordate, margin lobed, basal lobes auriculate,
rounded, slightly prominent with 2-6 teeth or
lobes per side, including an ascending lobe
becoming more prominent in upper parts, to
20 mm long and 6 mm wide, terminal lobe
triangular to oblong triangular, even linear, to
40 mm long and 15 mm wide, margin undulate
to shallowly lobed, + glabrous to moderately
hairy above, moderate to sparse below, hairs
crisped-appressed to loosely ascending,
occasionally semi-erect, 0.2-0.4(-0.5) mm
long. In upper parts, leaves with shorter
petioles, blades narrower triangular-ovate,
sagittate and basal lobe barely 2-toothed with
4 or 5 undulations per side. Inflorescence
solitary, axillary, bracteolate, a one-sided
dichasium, with l-3(-4) flowers, or
occasionally with 2 inflorescences per axil;
peduncle terete to slightly flattened, ribbed, 10-
60 mm long, moderately to sparsely hairy, hairs
crisped appressed 0.15-0.4 mm long; bracteoles
opposite to distinctly alternate, linear to

Johnson, Convolvulus L. in Australia

21

subulate, 1-3 mm long, 0.4-0.7 mm wide, apex
acute, ± ciliate, ± glabrous to sparsely hairy on
the back; pedicel ± ribbed, 5-20 mm long, not or
only very slightly recurved at fruiting, hairs
similar to and often denser than on the peduncle.
Outer sepals obovate to oblong or elliptic, 5-7
mm long, 2.5-5 mm wide, apex acute with a
recurved apiculum, + ciliate, moderately to
sparsely hairy to ± glabrous outside, hairs
crisped, loosely appressed to ascending, 0.15-
0.3(-0.5) mm long; inner sepals obovate to
elliptic, acuminate, 4.6-6.5 mm long, 3.5-5.5
mm wide, apex acute to obtuse with a distinct
curved apiculum, base truncate, ± glabrous,
sometimes sparsely hairy and ciliate. Corolla
funnel-shaped, pink or mauve with a pale
greenish throat, 7-15 mm long, 8-20 mm
diameter, flared 3.5-5 mm above the base of the
tube; petals 10-16 mm long, 7-12 mm wide, with
rounded-triangular, emarginate to apiculate
lobes, glabrous except for hairs on the outside
of the midpetaline band for 4-8 mm from the
apex. Stamens 5; filaments affixed to the corolla
tube for 1.5-3 mm from the base, free for 3-5.2
mm, with low tubercles from 1 mm above the
base of the corolla and extending for 1.75^4 mm;
anthers broadly oblong to triangular-oblong,
1.4—2.2 mm long, 0.6-0.9 mm wide, apex rounded,
emarginate, base sagittate, basal lobes 0.25-0.45
mm long. Ovary globular-ovoid, 1-1.5 mm long,
on a distinct disk 0.25-0.5 mm high, glabrous;
style 3-7 mm long, with cylindrical, obtuse,
stigmas, 1.8-2.35 mm long, suffused with pink.
Capsule iglobular to globular-ovoid, 4.5-6 mm
long, 5.5-6.5 mm diameter, glabrous. Seeds 4,
Vi-globular, 2.8-3.7 mm long, 2.3-2.5 mm wide,
black to dark brown, surface finely punctate
bearing numerous small irregular tubercles and
no distinct wing (Fig. 6C).

Specimens examined : Queensland. Port Curtis District:
Rockhampton, Apr 1867, O’Shanesy 55 (MEL); Neeko[o]l
Creek, s.d., [ Bowman ] s.n. (MEL). Burnett District
Bundaberg, Mar 1980, Stanley 919 (BRI). Darling Downs
District: c. 3 miles S of Mt Mowbullan Guest House on
Bunya Mts - Bell road, May 1958, Johnson & Pedley 453
(BRI); Atkins Lagoon, Pelican, Jan 1980, Lithgow 702
(BRI). Moreton DiSTRict: Tarampa Creek, s.d., Bailey
s.n. (BRI); Mt Mistake, s.d., Simmonds s.n. (BRI).
New South Wales. Clarence River, s.d., Beckler s.n.
(MEL); Mt Annan Botanic Garden, Mt Annan Ridge, 4
km W of Campbelltown, Dec 1985, Hind s.n. (NSW);
St Johns RC Cemerery, Campbelltown, May 1966,
McBarron 12570 (NSW); Darvall Park, Dennistone,
Dec 1978, Coveny 10408 (NSW); RC Cemetery,
Camden, Oct 1965, McBarron 11396 (NSW); 1 mile S

of Cambelltown on Appin Road, Jan 1969, Coveny
779 (NSW); South Creek, Rossmore, 10 miles W by S
of Liverpool, Apr 1968, Johnson 1801 (NSW);
Doonside, Leb 1984, Coveny 11781 (NSW); Maitland
Longbridge on New England Highway, Jan 1981, Medd
160021 (NSW); Singleton, Jun 1912, Breakwell s.n.
(NSW).

Distribution and habitat: C. erubescens is
found in coastal and sub-coastal areas from the
Sydney area in New South Wales to
Rockhampton in Queensland (Map 2). It is
found mainly in wetter eucalypt forests and in
rainforest margins.

Affinities: C. erubescens appears most closely
related to C. graminetinus. However,
C. erubescens has a larger corolla and longer,
straight to sinuate, pedicels at fruiting while in
C. graminetinus the pedicels are shorter and
strongly recurved at fruiting. For more than a
century, C. angustissimus was regarded as
conspecific with C. erubescens but the latter
has a compound dichasial inflorescence and a
distinctly different seed surface pattern which
distinguishes it from that species.

Phenology: Flowering occurs throughout the
late spring to early autumn with fruits recorded
mainly in summer and autumn.

Notes: Though the type specimen was not
located, the illustration, given in Curtis’s
Botanical Magazine and the associated
protologue, relate extremely well to the taxon
described above. Though no type specimen or
description of C. erubescens var. dilatatus
Choisy has been found, based on the specific
and subspecific epithets, I have concluded it
probably falls within my concept of
C. erubescens.

Specimens collected in the Sydney area
indicate there may have been some gene flow
between populations of C. angustissimus and
C. erubescens. Specimens which appear to be
of C. erubescens occasionally have recurved
pedicels but agree with C. erubescens in having
compound inflorescences, hastate leaves with
dentate basal lobes, ribbed peduncles and seeds
with closely patterned tubercles, e.g. Perthville,
Schijf (NSW455842). A specimen labelled as
from Yarra Yarra (MEL 689505) appears to
belong to this species. However the collecting
locality given for it is well outside the normal
range of C. erubescens and the label information

22

is queried.

Etymology : The specific epithet refers to the
flower colour in this species.

Conservation Status : In Queensland,
populations of this species have been
considerably reduced since European
settlement and no collections have been
recorded in the past 20 years. In view of the
uncertainty of the current population it is
probably best to regard it as rare.

8. Convolvulus wimmerensis R.W. Johnson,
sp. nov. affinis C. angustissimo R. Br. sed
pilis densis, appressis et argenteis, et
inflorescentia dichasiali unilaterali differt.
Typus: Victoria. Terrick Terrick State
Park M2, 24 November 1985, A.C.
Beauglehole 82670 (holo: MEL
[MEL689790]).

Perennial with trailing and twining stems; stems
terete, moderately to densely hairy, becoming
sparser with age, hairs crisped-appressed,
silvery, 0.15-0.5 mm long, sometimes with
some ascending and occasionally spreading
hairs to 0.9 mm. Leaves variable in shape and
size from base to tip of the stem (Fig. 5D). Basal
leaves petiolate; petiole long, occasionally
equalling the blade, 12-20 mm long; blade
ovate to oblong, occasionally triangular-oblong,
8-25 mm long, 5-15 mm wide, apex rounded
to truncate, sometimes slightly emarginate, base
cordate to truncate, decurrent, margin crenate
to shallowly lobed, basal lobes soon becoming
more prominent, moderately densely hairy to
+ glabrous, hairs ascending to erect, 0.2-0.7
mm long. Leaves on fertile stems petiolate;
petiole short 2-10 mm long; blade ovate in
outline, 13-30 mm long, 8-20 mm wide, apex
obtuse to rounded, rarely acute, base cordate,
decurrent, deeply divided almost to the base
into 5 lobes, basal pair of lobes linear, 2-12
mm long, toothed or bifid, with a secondary
recurved lobe from the lower margin, ascending
lobes linear, prominent, often more than half
the length of the terminal lobe, up to 20 mm
long, terminal lobe linear to narrowly oblong
or narrowly obovate 10-25 mm long, 1-5 mm
wide, irregularly toothed or lobed particularly
on more basal leaves, undulate to entire above,
moderately to densely hairy, hairs appressed
0.2-0.5 mm long, silvery. In upper parts, lobes

Austrobaileya 6(1): 1-39 (2001)

narrower and entire. Inflorescence solitary,
axillary, bracteolate, a one-sided dichasium with
1-2 flowers; peduncle terete, 10-35 mm long,
moderately to somewhat densely hairy, hairs
short, appressed to slightly ascending 0.15-0.5
mm long; bracteoles opposite to alternate, linear
to subulate, 1-2.5 mm long, 0.4-0.75 mm wide,
apex acute, ciliate, moderately densely hairy on
the back; pedicel 3-6 mm long, up to 10 mm at
fruiting, recurved at fruiting, hairs as for
peduncle. Outer sepals elliptic to obovate, 5-
6.5 mm long, 3-4.5 mm wide, apex barely acute
to rounded with a short recurved apiculum,
ciliate, moderately to densely hairy outside,
hairs appressed to slightly ascending, 0.2-0.5
mm long; inner sepals orbicular-oblong, 5-6 mm
long, 4—4.5 mm wide, apex rounded, with a short
recurved apiculum, base slightly cordate, rare
ciliate hair at the apex, ± glabrous. Corolla funnel-
shaped, pink, c. 10 mm long, 10-12 mm diameter,
flared c. 4-4.5 mm above the base of the tube;
petals 9-12 mm long, 7-8 mm wide, with rounded
apiculate lobes, glabrous except for appressed
hairs on the outside of the midpetaline band
for 3-6 mm from the apex. Stamens 5, unequal;
filaments affixed to the corolla tube for 2-2.5
mm from the base, free for 3-4 mm, with low
scattered tubercles from 1 mm above the base
of the corolla and extending for 2.5-3 mm;
anthers ovate to oblong, 1.25-1.5 mm long,
0.75-0.8 mm wide, apex obtuse, emarginate,
base sagittate, basal lobes to 0.3-0.35 mm long.
Ovary ovoid, 1.25 mm long, on a disk 0.3 mm
high, glabrous; style c. 4 mm long, with
cylindrical, obtuse stigmas, 1.2-1.25 mm long.
Capsule globular, 5.5-6 mm long, c. 5.5 mm
diameter, glabrous. Seeds 4, Vi-globular to V4-
globular-obovoid, 3.2-3.8 mm long, 2.5-3 mm
wide, dark brown, surface finely punctate
bearing scattered low, laterally compressed
tubercles, wingless (Fig. 6D).

Specimens examined: New South Wales. Brooking near
Urana, Crouch s.n. (MEL); Berrigan, Apr 1950, Godden
s.n. (NSW). Victoria. Lowan Mallee: Lowan, Oct 1896,
Reader s.n. (MEL). Midlands: Terrick Terrick State Park
M2, Nov 1985, Beauglehole ACB82670 (MEL). Riverina:
Barmah Regional Park L52, Jan 1986, Beauglehole
ACB83555 (MEL); Waaia via Numurkah, Mar 1959,
Cleaves s.n. (MEL). Wannon: Yarrackigarra Swamp
Wildlife Reserve C38, Nov 1986, Beauglehole ACB86983
(MEL); Nurcoung Lakes Reserve C41, Nov 1986,
Beauglehole ACB86599 (MEL). Wimmera: West Yanac
Wildlife Reserve C3, Sep 1986, Beauglehole ACB83982
(MEL); Lake Marmal Reserve H7, Dec 1985,

Johnson, Convolvulus L. in Australia

23

Map 5. Distribution of #. Convolvulus wimmerensis. ★. C. recurvatus subsp. recurvatus. ▲. C. recurvatus subsp.
nullarborensis. ■ . C. tedmoorei. Del. W. Smith.

Beauglehole ACB82942 (MEL); Mt Jeffcott Flora
Reserve, 17 km ENE of Donald PO, Oct 1979,
Beauglehole & Donald History Group ACB65283
(MEL); Wimmera, Dallachy 101 (MEL); Wimmera,
in 1892, Eckert s.n. (MEL); near Wycheproof, Oct
1917, Watts 778, 785, 786 (MEL); near Dimboola,
s.d., coll. ign. 23 (MEL).

Distribution and habitat : C. wimmerensis has
been most commonly recorded from the
Wimmera region of Victoria, but its distribution
extends along the northern part of the Riverina
District and into New South Wales (Map 5). It
grows on flat to undulating plains in open
Eucalyptus woodlands.

Affinities: C. wimmerensis resembles
C. angustissimus in having strongly recurved
fruiting pedicels and much divided leaves. It
differs in the shape of the lower cauline leaves,
in having dense appressed hairs on cauline
leaves and stems, and in having a one-sided
dichasial inflorescence with commonly 2
flowers. It has a small corolla at the lower end
of the size range for C. angustissimus. It also
resembles C. eyreanus, C. recurvatus,
C. crispifolius and C. clementii but differs from
them in having flowers with longer petals.

Phenology: Flowering and fruiting have been
recorded from November to March.

Etymology : The specific epithet refers to the
Wimmera District of Victoria where the species
occurs.

9. Convolvulus angustissimus R.Br., Prod.

482 (1810); Convolvulus erubescens var.

angustissimus (R.Br.) Choisy in A.DC.,
Prod. 9:412 (1845). Type: Tasmania, Van
Diemens Land near Risdon Cove, in 1804,
R. Brown (holo: BM[Bennett
27 65];iso:MEL[MEL6 8 9920],
MEL[MEL689582]) (see under Notes).

Convolvulus erubescens var. albus
Guilfoyle, Australian Plants: 117 (1911).
Type: “Vic.” (holo: n.v.).

Convolvulus geniculatus Lehm., Semina in
horto botanico Hamburgense 1826
collecta quae pro mutua commutatione
offeruntur (1826). Type: not cited.

Perennial with trailing and twining stems; stems
terete, densely hairy to ± glabrous in younger
parts, becoming sparser with age; hairs short,
± appressed, 0.2-0.5 mm long, in older parts
becoming erect and spreading and up to 0.8 mm
long, the relative abundance of erect and
appressed hairs varying among subspecies.
Leaves variable in shape and size from base to
tip of the stem. Basal leaves petiolate; petiole
long, often longer than the blade; blade ovate
to oblong, sometimes triangular, or linear,
occasionally slightly hastate, 5-35 mm long,
2-15 (-20) mm wide, apex acute to rounded-
truncate, occasionally emarginate, base tapering
to cordate, decurrent, margin entire, undulate,
irregularly crenate to shallowly lobed, basal and
ascending lobes barely more prominent,
moderately hairy to ± glabrous, hairs short,
appressed to longer, erect, depending on
subspecies. Leaf size and shape of lower cauline
leaves very variable depending on subspecies.

24

Leaves on fertile stems petiolate; petiole 2-20
mm long; blade narrowly ovate to ovate in
outline, 10-65 mm long, apex acute to obtuse,
mucronulate, occasionally rounded-emarginate,
more acute in upper parts, base tapering to
cordate, decurrent, hastate to deeply 3-5-lobed
from the base, basal lobes linear to obovate,
spreading to recurved, to 17 mm long, apex
obtuse or 2-toothed, often with a 2-toothed,
recurved secondary lobe on the lower margin,
ascending lobes linear, up to 30 mm long, 1-8
mm wide, terminal lobe narrow-linear to
narrow-oblong, occasionally triangular, 10-60
mm long, 1-8 mm wide, margin entire, rarely
undulate to slightly lobed, moderately hairy to
± glabrous above, sometimes slightly denser
below, hairs appressed to crisped-appressed,
0.15-0.35 mm long on some subspecies, mainly
ascending to semi-erect, 0.25-0.6 mm long on
others. In upper parts, petiole shorter, blade
shorter and with narrow acute lobes, often with
short, sometimes bifid, basal lobes and a very
narrow ascending lobe. Inflorescence solitary,
axillary, bracteolate with solitary flowers, very
rarely a one-sided dichasium with 2 flowers or
with 2 inflorescences per axil; peduncle terete,
4-50 mm long, moderately to sparsely hairy,
hairs mainly appressed, occasionally spreading;
bracteoles opposite, rarely sub-opposite, linear,
subulate to narrowly ovate, 1-4 mm long, 0.2-
0.5 mm wide, apex acute, ciliate, moderately
hairy to glabrous on the back; pedicel often
darker than the peduncle 3-23 mm long,
recurved at fruiting, hairs similar to and often
denser than on the peduncle. Outer sepals
obovate, obovate-oblong, to elliptic, (3.5-)4-
6.5 mm long, 2-5 mm wide, apex acute to
rounded with a short recurved apiculum, ciliate,
more rarely + eciliate (in subsp. fililobus),
moderately hairy to glabrous outside, hairs
appressed to spreading, depending on
subspecies; inner sepals obovate to obovate-
orbicular, more rarely elliptic or oblong, 3.5-6
mm long, 2.5—4.5 mm wide, apex rounded, with
a short recurved apiculum, base rounded to
truncate, + glabrous or with some hairs around
the apex and upper spine. Corolla funnel-
shaped, pink, with a paler throat, rarely white,
7-20 mm long, 7-20 mm diameter, flared 2.5-
6 mm above base of the tube; petals 8-25 mm
long, 3-12 mm wide, with rounded to rounded-
triangular, erose, often emarginate, barely
apiculate lobes, glabrous except for hairs on

Austrobaileya 6 (1): 1-39 (2001)

the outside of the midpetaline band for 1-9 mm
from the apex. Stamens 5, slightly unequal;
filaments affixed to the corolla tube for 1.5-3
mm from the base, free for 1.75-5 mm, with
low tubercles from 0.75 mm above the base of
the corolla and extending for 1-4 mm; anthers
oblong to ovate, 0.8-2.75 mm long, 0.5-1 mm
wide, apex rounded, emarginate, base sagittate,
basal lobes 0.2-0.7 mm long. Ovary ovoid 1-
1.8 mm long, on a well developed disk, 0.25-
0.6 mm high, glabrous; style 3-10 mm long,
with cylindrical to narrowly ovoid, obtuse
stigmas, 1-2.5 mm long. Capsule globular to
globular-ovoid, 4-8 mm long, 4-7.5 mm
diameter, glabrous. Seeds 4, 14-globular to
slightly 14-globular-obovoid, 2.9-4 mm long,
2.2-3.5 mm wide, dark brown to black, surface
finely punctate usually bearing low reticulate
± continuous ridges, smooth in subsp. fililobus,
wing not prominent, discontinuous to absent.

Affinities: C. angustissimus most closely
resembles C. wimmerensis. C. recurvatus has
similar recurved pedicels subtending mature
capsules but its pedicels are much shorter than
those in C. angustissimus, its flowers smaller
and its inflorescences often compound.

Notes: Brown (1810) gave no indication of the
type locality in Tasmania. He collected
specimens of C. angustissimus in Tasmania in
1804 and these collections are now represented
in the Natural History Museum, London on two
herbarium sheets (David Moore, BM, pers.
com.)

1. Bennett 2767 - One of three labels lists a
collecting locality as “Port Dalrymple prope
littoraJan: 1804”.

2. Bennett 2765 - Two labels are present on
the sheet, one states “In campis prope Baie
du Nord quandum Frederick Harvey
(Henry) Bay diet Feb: 1804”, the other “Van
Diemens Land near Risdon Cove” [Type of
Convolvulus angustissimus R.Br. according
to the sheet].

Based on this I have accepted the
designation of the latter collection as the
holotype. A further two specimens of this taxon
collected by Brown and held at MEL list only
“Van Diemens Land” as the collecting locality
and I have assumed these are isotypes.

Johnson, Convolvulus L. in Australia

25

Although the description of
C. geniculatus Lehm. is a very general one, and
no type specimen has been cited, on the basis
of its solitary flowers and geniculate pedicels
at fruiting, it is highly likely it is conspecific
with C. angustissimus.

This species shows considerable
variation. While the simple inflorescence,
relatively large flowers and recurved fruiting
pedicels characterize this taxon, other attributes
are very variable, particularly its leaf shape and
vestiture. Further field work is probably needed
for a better understanding of the complex. The
specimen collected by Brown from Risdon
Cove consists of the plant crown and upper
leaves which have much divided narrow basal
lobes, narrow ascending lobes and a narrow-
linear entire terminal lobe. Leaves in the lower
part of the plant are absent. A good match for
the type specimen is one collected at Ross
{Burns 7). All other specimens borrowed from
HO have ovate, oblong or triangular, hastate,
lower cauline leaves, becoming much divided
up the stem with leaves on terminal branches
of older plants with very narrow lobes
resembling those on the holotype. Parham
( pers. comm.) did not find any specimens in
HO with narrowly lobed, much divided leaves
to the base of the stem. In addition, in visits to
both Risdon Cove and Ross, Parham did not
find any plants with narrow, deeply divided,

lower cauline leaves.

However in Victorian specimens seen,
many have a few linear hastate basal leaves
which are abruptly replaced distally by lobed
leaves with very narrow linear lobes up to 2 mm
in width. These leaves resemble those on the
type specimen. The basal leaves absent from
the type specimen may have been similar to the
broad divided leaves found on other Tasmanian
specimens or the very narrow leaves found on
the type may have occurred along the stem to
the base as occurs in many Victorian specimens.
For this revision I have accepted the former
possibility. On this basis, all Tasmanian material
is of C. angustissimus subsp. angustissimus.

Both C. angustissimus subsp.
omnigracilis and C. angustissimus subsp.
fililobus are reasonably distinctive but much
variation is present within C. angustissimus
subsp. angustissimus and C. angustissimus
subsp. peninsularum. Because of the great
variation, resulting from ontogenetic
development and geographic distribution, and
the influence of time of germination and length
of the growing season on morphology, it is
difficult to classify the existing variation on the
basis of herbarium specimens. In addition,
hybridisation further confounds attempts at
classification. More field work will be necessary
to clarify the variation which exists within these
taxa.

Four subspecies are recognised and can be distinguished as follows.

1. Lower cauline leaves broad and much divided with the terminal lobe more

than 1.5 mm wide; lobes gradually becoming narrower towards the tip

.9a. C. angustissimus subsp. angustissimus

Cauline leaves with narrow lobes almost from the base of the plant; lobes
<1.5 mm wide, or if wider then basal lobes spreading and clavate

and terminal lobe distinctly obovate-clavate. 2

2. Flowers 14-25 mm long; flowering pedicels 8-18 mm long; outer sepals

moderately to sparsely hairy, usually ciliate at the tip; seeds with sparse

low reticulate ridges.9b. C. angustissimus subsp. omnigracilis

Flowers 9-14 mm long; flowering pedicels 4-8 mm long.3

3. Seeds smooth; sepals + glabrous to sparsely hairy, with hairs mainly

appressed, leaves with ascending lobes often > 1/3 the length of the terminal

lobe and lacking distinctive basal lobes.9c. C. angustissimus subsp. fililobus

Seeds with sparse low reticulate ridges; sepals moderately to sparsely hairy
with ascending and spreading hairs common; leaves with ascending lobes
mostly < 1/3 the length of the terminal lobe; lower cauline leaves with
distinct, spreading, clavate basal lobes.9d. C. angustissimus subsp.peninsularum

26

9a. Convolvulus angustissimus R. Br. subsp.

angustissimus

Convolvulus adscendens de Vriese in
Lehmann, Plantae Preissianae 1: 346
(1845). Type: In arenosis apertis distr.
York, 16 March 1839, Herb. Preiss No.
1924 (holo: LD; iso: MEL[MEL689918,
MEL689919]).

Convolvulus subpinnatifidus de Vreise in
Lehmann, Plantae Preissianae 1: 347
(1845). Type: In solo sublimoso fertili
prope Beljarup, Hay, 4 November 1840,
Herb. Preiss No. 1925 (holo: LD; iso:
MEL[MEL689916, MEL689917]).

Convolvulus acaulis Choisy in A.DC. Prodr.
9:406 (1 Jan 1845). Type: NovaHollandia
(ins. Kanguroos) (PARIS), (holo: P).

Stems densely to moderately hairy in younger
parts, becoming sparser with age, hairs +
appressed, 0.2-0.4 mm long, on older parts
becoming erect and spreading and up to 0.8 mm
long. Leaves variable in shape and size from
base to tip of the stem (Lig. 7 A). Blade of basal
leaves ovate to oblong, sometimes triangular
or linear, occasionally slightly hastate, 5-25
mm long, 4-15 mm wide, apex acute to
rounded-truncate, occasionally emarginate,
base truncate to cordate, decurrent, margin
irregularly crenate to serrate to shallowly lobed,
basal and ascending lobes barely more
prominent, moderately hairy to ± glabrous, hairs
semi-erect, 0.25-0.8 mm long. Leaves on fertile
stems petiolate; petiole 2-20 mm long; blade
ovate to triangular-ovate in outline, 10-65 mm
long, 2-40 mm wide, apex acute, rarely obtuse
to rounded-emarginate, more acute in upper
parts, base cordate, decurrent, often 3-5 lobed
from the base, basal lobes linear to narrowly
oblong to 17 mm long, entire or 2-toothed, often
with a 2-toothed recurved secondary lobe on
the lower margin, ascending lobes linear, up to
30 mm long, 1-8 mm wide, terminal lobe linear to
narrow-oblong, often triangular, 10-60 mm long,
1-8 mm wide, entire, rarely undulate to slightly
lobed, moderately hairy to ± glabrous above,
slightly denser below, hairs mainly ascending
to semi-erect, 0.25-0.6 mm long. In upper parts,
petiole shorter, lobes becoming narrower, more
acute, sometimes with very short basal and
ascending lobes and a long narrow-linear

Austrobaileya 6 (1): 1-39 (2001)

terminal lobe. Peduncle 5-50 mm long; pedicel
3-23 mm long. Outer sepals obovate to obovate-
oblong, more rarely elliptic, 4-6 mm long, 2.5-5
mm wide, ciliate, moderately to sparsely hairy
outside, hairs loosely ascending to spreading,
some appressed, or more rarely almost all
appressed, 0.15-0.5 mm long; inner sepals
obovate to obovate-orbicular, rarely elliptic,
4.5-5.7 mm long, 3-4.5 mm wide, + glabrous
to sparsely hairy outside. Petals 9-21 mm long,
5-12 mm wide. Capsule globular to globular-
ovoid, 4-8 mm long, 4-7 mm diameter. Seeds
2.9-4 mm long, 2.2-3.5 mm wide bearing low
reticulate ± continuous ridges (Pig. 8A).

Selected specimens (230 specimens examined): Western
Australia. King Georges Sound, s.d., Muir s.n.
(MEL689547); Harvey, Nov 1916, Stoward s.n. (PERTH).
South Australia. Big Heath NP, S portion, c. 7 km SE of
Nine Mile Well, Nov 1969, Jackson 1605 (AD); Mt Lofty
Ridge Wildlife Reserve, Feb 1969, Sexton s.n. (AD);
Lenswood Agricultural Research Centre, north east Nov
1978, Spooner 6205 (AD); near su mmi t of Mt Barker, c.
30 km SE of Adelaide, Dec 1964, Whibley 1526 (AD); c.
20 km SE of Mt Gambier, Wilson 799 (AD). Queensland.
Burnett District: Narayen, Mar 1973, coll. ign. N1334
(BRI). Maranoa District: Stanhope Downs, 44 km by
road NW of Roma, Nov 1996, Thomas s.n. (BRI).
Darling Downs District: Kildonan, Feb 1936, Blake
10528 (BRI). New South Wales. Molong, Nov 1906,
Boorman s.n. (NSW); NSW-ACT border adjoining
Queanbeyan rubbish tip, Jan 1983, Coveny & Hind
11502 (BRI, NSW); 12 miles NE of Albury on Hume
Highway, Oct 1967, Muir 4605 (MEL); Sinclair
Lookout, 14.4 km W of Glen Innis, Mar 1987, Plat,
Coveny & Dunn 7 (BRI); SE of Nimmitabel, Nov 1960,
Salasoo 1980 (NSW); Armidale, UNE hill, Nov 1959,
Winterhalder s.n. (NE). Victoria. Midlands: 7 miles
SW of Thoona, Nov 1960, Muir 1756 (MEL). Riverina:
Farran’s Lookout on Murray Valley Highway, 3 miles
N of Towong, Oct 1961, Muir 2395 (MEL). Victorian
Volcanic Plain: Warrnock road, c. 14 km SSW of
Chetwynd, Nov 1982, Corrick 8472 (BRI, MEL).
Wannon: Portland, Gorae West, 1946, Beauglehole
38437 (MEL). Tasmania. Poatina, Nov 1986,
Buchanan 8854 (HO); Domain, Hobart, Oct 1942,
Curtis s.n. (HO); Cape Portland, Oct 1983, Moscal s.n.
(HO); Township Lagoon, Nov 1983, Moscal 3894 (HO,
MEL).

Distribution and habitat : C. angustissimus
subsp. angustissimus has been recorded from
all States but not from the Northern Territory
(Map 6). It occurs in Tasmania mainly in the
Hobart-Launceston area with no records from
the western part of the State. It is widespread
in Victoria but is absent from the north-west
and east coast. It extends into the south-east of
South Australia with other populations in the

Johnson, Convolvulus L. in Australia

27

Adelaide area and on the Yorke and Eyre
Peninsulas. In Western Australia, it is restricted
to the south-western corner. In eastern
Australia, it extends northwards through the
tableland areas of New South Wales into
southern Queensland, where it is found mainly
west of the Great Dividing Range as far north
as Springsure. It is found on level to hilly terrain
in mainly loamy and clay soils, less commonly
in sandy and rocky soils. It grows in grassy
eucalypt woodlands and forests and in
grasslands which develop following the
clearing of the woodlands.

Phenology: Flowering occurs mainly in early
spring to mid autumn (September-April) while
fruiting occurs from late spring to late autumn
(November-May).

Notes: As circumscribed here, considerable
variation exists within this subspecies. The
typical form with large flowers and long
pedicels is found in Tasmania and southern
Victoria and extends into south-eastern South
Australia and north into New South Wales.
Similar forms are found at Narayen,
Queensland (AQ 637866), at Armidale in
northern New South Wales and at the
Wombeyan Cave ( Constable NSW56058). In
the Kosciusko region of New South Wales and
nearby areas of Victoria, specimens (e.g. Muir
2395, Makinson 975, Walsh 279, Forbes 576)
have denser erect hairs. However similar
vestiture is also found on occasional specimens
throughout the range of this subspecies. In drier
parts of Victoria on through South Australia to
Western Australia there is a gradual reduction
in the size of the corolla with petals on most
specimens rarely exceeding 14 mm in length.

In Queensland, a variant with moderately
dense ascending to erect hairs up to 0.75 mm
long is found in the Charleville district
{Clements AQ275818, Bailey AQ275821). This
variant has black seeds, 4.5-5 mm long, which
are larger than those from other populations.
In addition the surface ridges on the seeds are
more distinctly raised and the discontinuous
wing more prominent. Both are formed by fused
hair-like structures. No collections have been
made of this population since 1945 and the
study of further material may warrant its
recognition as a distinct taxon.

9b. Convolvulus angustissimus subsp.
omnigracilis R.W. Johnson, subsp. nov.
affinis C. angustissimo R. Br. subsp.
angustissimo sed foliis valde anguste
lobatis fere basi differt. Typus: Victoria.
Victorian Volcanic Plain: Nerrin-Nerrin
- Woorndoo road, 4 km WSW of Mt
Hamilton, near Pagel’s Lane (37°48’S
142°56 , 30”E), 27 November 1983, S.J.
Forbes & N. Scarlett 1867 (holo:MEL
[MEL67409]; iso: BRI [AQ420954]).

Stems moderately densely to sparsely hairy,
becoming sparser with age, hairs + appressed,
occasionally spreading, 0.15-0.4 mm long.
Leaves are somewhat variable in shape and size
from base to tip of the stem though lobes are
narrowly linear throughout (Fig. 7B). Blade of
basal leaves linear, linear-elliptic to narrow
oblong, hastate, 5-35 mm long, 2-10 mm wide,
apex obtuse to rounded, rarely emarginate, base
tapering to truncate, margin entire to slightly
undulate but soon with narrow bifid basal lobes,
well developed ascending and terminal lobes,
sparsely hairy to + glabrous, rarely moderately
hairy, hairs ± appressed, 0.15-0.4 mm long.
Leaves on fertile stems petiolate; petiole short,
3.5-15 mm long; blade linear to ovate in
outline, 12-60 mm long, apex acute to almost
obtuse, base tapering to cordate, hastate to
deeply lobed, basal lobes linear, often short,
0.5-5 mm long, occasionally to 12 mm long,
sometimes with a recurved secondary lobe from
the lower margin, ascending lobes linear to 25
mm long, < 1 mm wide, terminal lobe linear or
obovate-linear 12-60 mm long, < 2 mm wide, +
glabrous to sparsely hairy, hairs appressed,
0.15-0.35 mm long, margin entire. In upper parts,
leaves smaller but of similar shape. Peduncle 5-
35 mm long; pedicel 5-18 mm long. Outer sepals
obovate to elliptic, 4.5-6 mm long, 2.8-3.7 mm
wide, apex acute to rounded with a shortly
recurved apiculum, ciliate, moderately to
sparsely hairy outside, hairs appressed, 0.1-
0.3 mm long; inner sepals obovate, 4.8-6 mm
long, 3-4 mm wide, ± glabrous to sparsely hairy
outside. Petals 14-25 mm long, 8-12 mm wide.
Capsule globular, 5-8 mm long, 4.5-7 mm
diameter. Seeds 3.5—4 mm long, 2.7-3.3 mm wide
bearing low reticulate ± continuous ridges (Fig.
8B).

28

Austrobaileya 6 (1): 1-39 (2001)

Fig. 7. Variation in leaf shape of Convolvulus species from base to tip of stems. A. C. angustissimus subsp.
angustissimus. (1) Moscal 3894 (HO) x 1, (2) coll. ign. s.n. (H036311) X 1, (3) Rodway s.n. (H015256) x 1, (4) Tate s.n.
(AD97605515) x 2; B. C. angustissimus subsp. omnigracilis. (1) Stones s.n. (MEL503463) x 2, {2) Baker s.n. (MEL536246)
x 1, (3) Forbes 1867 & Scarlett (MEL) x 2; C. C. angustissimus subsp. fililobus x 2. (1) Beauglehole ACB87768 (MEL),
(2) Aston 2367 (MEL), (3) Beauglehole ACB66176 (MEL); D. C. angustissimus subsp. peninsularum. (1) Eichler 14063
(AD) x 2, (2) Eichler 14063 (AD) x 1, (3) Alcock 654E (AD) x 2, (4) Alcock 654B (AD) x 1. Del. W. Smith.

Selected specimens (36 specimens examined):
Victoria. East Gippsland: Amboyne Creek Area, 11 km
NW of Tubbut PO, Jan 1980, Beauglehole ACB67710
(MEL). Eastern Highlands: Whittlesea, Mar 1904, Baker
s.n. (MEL 536248). Midlands: 15 km W of
Maryborough PO, N of abandoned goldmine J7, Nov
1979, Beauglehole & Maryborough FNC ACB66592
(MEL); 8.2 km SE of Ararat, Jan 1995, Zich & Young
258 (CANB). Otway Plains: Limeburners Lagoon Flora
& Fauna Reserve P3, Mar 1982, Beauglehole & Errey
ACB70341 (MEL). Victorian Volcanic Plain: Corio,
Geelong Area, Feb 1964, Anderson s.n. (MEL 503341);
Laverton, 13 miles WSW of Melbourne, 0.5 miles NE
of Laverton, Nov 1962, Aston 847 (MEL);
Broadmeadows & Glenroy, Oct 1903, Baker s.n. (MEL
536247); Campbellfield, Broadmeadows, Nov 1900,
Baker s.n. (MEL 536250); Preston, Nov 1899, Baker
s.n. (MEL 536249); Deer Park, Sep 1900, Baker s.n.
(MEL 536246); Werribee, Dec 1899, Baker s.n. (MEL
536255); Keilor, n.d., Cowle s.n. (MEL 579995; MEL
689658); Nerrin-Nerrin Woorndoo road, 4 km WSW
of Mt Hamilton, Nov 1983, Forbes & Scarlett 1867
(MEL); Williamstown Butts, Dec 1953, Hansen s.n.
(MEL 689793); Camperdown-Foxhow road, 25 km

WSW of Cressy PO, Oct 1977, Hirth s.n. (MEL
1513254); Campbellfield, Melbourne, Oct 1978, Muir
6277 (MEL); Lake Corangamite, SW of Cundare, Oct
1982, Scarlett s.n. (BRI [AQ377773]); Rifle Range,
Williamstown, Dec 1943, Smith 43/104 (MEL); Altona,
Oct 1975, Stones s.n. (MEL 503463); St Albans
Railway, Oct 1975, Stones s.n.(MEL 503472).

Distribution and habitat: C. angustissimus
subsp. omnigracilis is restricted to Victoria
being found around Melbourne and extending
to the west and north-west to near Ararat and
St Arnaud (Map 7). It grows mainly in grassy
communities on plains on grey to yellow clay
loam or clay soils. These soils are commonly
derived from basalt or Ordovician shales.
Populations also occur around Lake
Corangamite, sometimes on Coxiella shell
deposits.

Phenology : Flowering occurs mainly in mid

Johnson, Convolvulus L. in Australia

29

Fig. 8. Seeds of Convolvulus augustissimus, adaxial (L.H.S.) and abaxial (R.H.S.) surfaces x 10. A. C. angustissimus
subsp. angustissimus from Hosking 661 (NSW); B. C. angustissimus subsp. omnigracilis from Smith 43/104 (MEL); C. C.
angustissimus subsp .fililobus from Aston 2367 (MEL); NB. Seeds of C. angustissimus subsp. peninsularum not available.
Del. W. Smith.

Map 7. Distribution of •. Convolvulus angustissimus subsp. omnigracilis. ★. C. angustissimus subsp. fililobus.
▲ . C. angustissimus subsp. peninsularum. Del. W. Smith.

30

spring to early summer (October-December)
with some flowering throughout late summer
and autumn. Fruits are produced throughout
the spring to autumn period.

Etymology : The subspecific epithet refers to
the lobes of the much divided leaves of this
subspecies which are slender in all leaves on
the plant.

9c. Convolvulus angustissimus subsp.
fililobus (Wawra) R.W. Johnson, comb,
nov. Convolvulus erubescens var.
fililobus Wawra, Itinera Principum S.
Coburgi 1: 102 (1883). Type: Victoria.
Prarien des Murraygebietes (holo: W).

Stems mainly trailing, moderately to sparsely
hairy, becoming sparser with age, hairs short
appressed, 0.15-0.3(-0.5) mm long. Blade of
basal leaves linear, linear-elliptic to narrowly
oblong, hastate, 5-20 mm long, 5-8 mm wide,
apex obtuse to rounded, mucronulate, base
tapering to truncate, margin undulate, often
toothed or lobed towards the base, sparsely
hairy to ± glabrous, hairs appressed, 0.1-0.4
mm long, but soon becoming deeply and
narrowly 5-lobed. Leaves on fertile stems
petiolate; petiole short 2.5-8 mm long; blade
ovate in outline, 15-40 mm long, apex acute to
almost obtuse, base tapering to shallowly
cordate, deeply 5-lobed, basal lobes linear, 1-
7(-10) mm long, somet im es with a recurved
secondary lobe from the basal margin,
ascending lobes narrowly linear to 20 mm long,
< 1 mm wide, terminal lobe narrowly linear,
15-40 mm long, <1.5 mm, entire, moderately
to very sparsely hairy, hairs appressed, 0.1-0.3
mm long. In upper parts, leaves smaller, often
with prominent terminal lobe and short but
distinct narrowly linear basal and ascending
lobes (Fig. 7C). Peduncle 4-13(-20) mm long;
pedicel 4-8 mm long, occasionally extending
to 12 mm at fruiting. Outer sepals obovate to
elliptic, 3.5-5(-5.5) mm long, 2.4-3.5 mm
wide, eciliate or with a few scattered cilia,
sparsely hairy to + glabrous, very rarely
moderately hairy outside, hairs mainly
appressed, 0.15-0.25 mm long. Inner sepals
obovate to oblong, 3.5-5(-5.5) mm long, 2.7-
3.8 mm wide, glabrous. Petals 9-14 mm long, 5-
9 mm wide. Capsule globular, 6-6.5 mm long, c.
5.5 mm diameter. Seeds 3-4 mm long, 2.2-2.6
mm wide, golden brown with darker mottling,

Austrobaileya 6 (1): 1-39 (2001)

surface finely punctate without any
protuberances, scurfy with a distinct ridge on
the outer margins, occasionally with a very
narrow discontinuous wing of fused hair-like
structures (Fig. 8C).

Specimens examined: New South Wales. 13 km E of
Urana, on Urana to Lockhart road, Nov 1982, Aston 2367
(MEL.BRI); Falkiner Memorial Field Station, Deniliquin,
Nov 1975, Crisp 1813 (CBG); Falkiner Memorial Field
Station, Deniliquin, Dec 1945, Willoughby 77 (CANB).
Victoria. Grampians: Mt Arapiles SW side, c. 0.75 miles
W of Natimuk Golf Course, Nov 1968, Beauglehole
ACB29738 (MEL); Mitre Rock, 10 km W of Natimuk PO,
Nov 1979, Beauglehole ACB66176 (MEL); Grampians, 6.4
km from Stawell on road to Halls Gap, Nov 1959, Symon
86 (NE). Riverina: Gaynor Swamp Wildlife Reserve,
Apr 1981, Beauglehole ACB68838 (MEL); Spence
Bridge Education Area L38, Sep 1985, Beauglehole
ACB80211 (MEL); Tocumwal Regional Park, Sep
1985, Beauglehole ACB81273 (MEL); Barmah State
Park L53, Nov 1985, Beauglehole ACB82215 (MEL);
Barmah State Forest L52, Nov 1985, Beauglehole
ACB82460 (MEL); Murray River Reserve G36, Dec
1985, Beauglehole ACB83322 (MEL); Hunter, 11 km
NW of Elmore, Nov 1989, Davies & Hadlow 1334
(CBG); Tatura, Nov 1945, Gauba s.n.(CBG 12623);13
miles S Shepparton on Goulburn Valley Highway, Oct
1967, Muir 4632 (MEL); Rushworth-Murchison Road,

6 km ESE of Rushworth M34, Dec 1981, Muir 7000
(MEL); c. 1 km S of Wunghnu between Goulburn Valley
Highway and railway line, Nov 1981, Stebbing s.n.
(MEL 642833). Wannon: Mooree Historic Reserve Cl9,
Nov 1986, Beauglehole ACB87426 (MEL); Kialla
Roadside Reserve, Dec 1986, Beauglehole ACB87649
(MEL); Tarranjurk Roadside Reserve C17, Dec 1986,
Beauglehole ACB87768 (MEL). Wimmera: Jeffcott
Roadside Reserve, Dec 1986, Beauglehole ACB87967
(MEL); 9 miles from Quambatook towards Dumosa,
Oct 1963, Phillips 17 (CBG).

Distribution and habitat: C. angustissimus
subsp. fililobus is found in an arc from central
western Victoria, west of Horsham, through the
Riverina District of Victoria and extending into
New South Wales in the southern part of the
south western plains in the Urana-Deniliquin
area (Map 7). It grows on clay and clay loam
soils on plains.

Phenology : Flowering occurs mainly in early
spring to early summer (September -
December) while fruit are produced throughout
spring to early autumn.

Etymology: The subspecific epithet refers to
the finely divided leaves characteristic of plants
of this subspecies.

Johnson, Convolvulus L. in Australia

31

9d. Convolvulus angustissimus subsp.
peninsularum R.W. Johnson, subsp. nov.

affinis C. angustissimo R. Br. subsp.
angustissimo sed foliis anguste lobatis e
basi et foliis humilibus lobis basalibus
distincte effusis obovatis differt. Typus:
South Australia. Yorke Penninsula
Region: Innis National Park, 3 km S of
northern boundary (35°15’S 136°55’E),
9 October 1974, C.R. Alcock 4733 (holo:
AD[AD97523049]; iso: SYD, n.v.).

Stems moderately hairy to + glabrous,
glabrescent, hairs appressed to crisped-
appressed, 0.2-0.4 mm long though towards
the base, hairs more spreading and longer.
Leaves variable in shape and size from base to
tip of the stem (Fig. 7D). Blade of basal leaves
oblong hastate, 5-10 mm long, 3-8 mm wide,
apex obtuse to rounded, base cordate, margin
undulate, sparsely to moderately hairy, hairs
spreading, 0.25-0.5 mm long, blades soon
becoming longer and narrower with distinct,
spreading, entire, 2-toothed or bi-lobed basal
lobes and a narrow-linear to linear obovate
terminal lobe up to 25 mm long, rarely more
than 4 mm wide with an undulate to distinctly
lobed margin, hairs becoming more appressed.
Leaves on fertile stems petiolate; petiole 3-12
mm long; blade narrow-linear or narrowly ovate
in outline, 15-35 mm long, apex obtuse,
mucronulate, base cordate, sometimes truncate,
basal pair of lobes linear to obovate, lateral to
recurved, up to 7 mm long, apex obtuse, toothed,
often with a recurved secondary lobe from the
lower margin, ascending lobes linear, up to 10
mm long, terminal lobe linear to linear-obovate,
15-35 mm long, up to 5 mm wide, margin entire
or slightly undulate, moderately hairy to +
glabrous, hairs spreading to appressed, 0.2-0.5
mm long. In upper parts, leaves linear, hastate,
often with short basal lobes, sometimes bifid,
sometimes with ascending lobes, and a very
narrow terminal lobe. Peduncle 4-20(-35) mm
long; pedicel 3-8 mm long. Outer sepals oblong,
obovate-oblong, rarely elliptic, 4-5.5 mm long,
2-4 mm wide, apex rounded apiculate, ciliate,
moderately dense to sparsely hairy outside,
hairs appressed, ascending to spreading, 0.2-
0.5 mm long; inner sepals obovate-oblong to
elliptic, 4-5.5 mm long, 2.5-3.5 mm wide,
moderately hairy to glabrous. Petals 8-13 mm
long, 3-10 mm wide. Capsule globular 5-6 mm

long, c. 5 mm diameter. Seeds 3.2-3.3 mm long,
c. 2.5 mm wide, surface finely punctate bearing
low reticulate ridges.

Selected specimens (28 specimens examined): South
Australia, nr Yallunda Flat, midway between Tumby Bay
and Cummins, Oct 1964, Alcock 654A(AD); [Port] Lincoln,
Northshore, Nov 1964, Alcock 654B (AD); Coontra Creek,
Lincoln Highway, Nov 1964, Alcock 654C (AD); near
Lipson, Lincoln Highway, 10 km N of Tumby Bay, Dec
1964, Alcock 654E (AD); near Boston House, 3 km N of
Port Lincoln, Jan 1965, Alcock 654L (AD); Lincoln
Highway, 3 km N of Port Lincoln, Oct 1964, Alcock s.n.
(AD); Hundred of Blessing, E End Camp, S of Bascombe
Well, Oct 1967, Alcock 1481 (AD); Hincks NP, N-S track.
Hundred of Nicholls, Oct 1968, Alcock 2365 (AD); Innis
NP, 3 km S of N boundary, Oct 1974, Alcock 4733 (AD);
Coontra Creek, near Tumby Bay, Jan 1965, Alcock 654
(AD); Pondalowie Bay, c. 10 km NW of Stenhouse Bay,
Oct 1965, Blaylock 43 (AD); Port Lincoln, in 1875, Browne
s.n. (MEL); sea coast, c. 4 km S of Corny Point, Sep 1957,
Eichler 14063 (AD); Hallett Cove, c. 20 km SSW of
Adelaide, Mar 1937, Ising s.n. (AD); Sellicks Beach scrub,
Nov 1968, Kraehenbuehl 2657, 2670 (AD); Hallett Cove
Conservation Park, Dec 1978, [ Launer ] 9 (AD); Hundred
of Noarlunga Sec. 190, Nov 1966, Parsons 200 (AD); Hog
Bay E end, Kangaroo Island, Nov 1883, Tate s.n. (AD);
Venus Bay, s.d., Warburton. s.n. (MEL); Hundred of
Blessing, c. 10 km SW of Bascombes Well, Oct 1967,
Wheeler 662 (AD).

Distribution and habitat: C. angustissimus
subsp. peninsularum is found on the Eyre and
Yorke peninsulas in South Australia and on
Kangaroo Island (Map 7). It is found on
seashores and coastal cliffs but also extends
inland, occuring mainly in heathlands and
grasslands.

Phenology: Flowers are found in late spring
and throughout summer with fruits being
produced soon after flowering.

Notes: This subspecies includes very distinctive
variants which occur in coastal areas mainly in
the southern parts of the Yorke and Eyre
Peninsulas. It is characterised by spreading
basal lobes on the lower leaves. It becomes
difficult from incomplete herbarium specimens
to distinguish this subspecies from smaller
flowered forms of C. angustissimus subsp.
angustissimus which occur to the north of it.
Further field work will be needed to understand
better the relative distributions of these
subspecies. Few seeds of C. angustissimus
subsp. peninsularum were available for study
and further information on seed size and
sculpture is needed to assess their relevance.

32

Etymology : The specific epithet refers to the
known distribution of this subspecies which is
found mainly on the Yorke and Eyre peninsulas
in South Australia.

10. Convolvulus recurvatus R.W. Johnson, sp.
nov. affinis C. clementio Domin sed
pedicellis ad fructificans brevioribus et
recurvis differt. Typus: South Australia.
Bute District, c. 140 km NNW of
Adelaide, 28 October 1966, B. Copley
827 (holo: AD; iso: K, n.v.).

Perennial with trailing and twining stems; stems
terete, moderately hairy, becoming sparser with
age, hairs appressed to ascending and
spreading, 0.2-0.8 mm long. Leaves variable
in shape and size from base to tip of the stem.
Basal leaves petiolate; petiole long, often longer
than the blade, 5-20 mm long; blade linear to
oblong, sometimes triangular, 6-20 mm long,
4-15 mm wide, apex obtuse to rounded,
mucronulate, sometimes emarginate, base
truncate to cordate, sometimes hastate, margin
+ entire, undulate to irregularly crenate or
lobed, soon becoming 3-5-lobed with lobes
having lobed to entire margins, + glabrous, soon
becoming moderately hairy, hairs loosely
appressed to spreading 0.15-0.6 mm long.
Leaves subtending inflorescences petiolate;
petiole 2-20 mm long; blade ovate in outline,
usually divided almost to the base into 3-5
lobes, 7-35 mm long, 5-20 mm wide, apex
bluntly obtuse to truncate, mucronulate,
sometimes emarginate, base cordate, decurrent,
basal pair of lobes linear to narrowly oblong,
1-12 mm long, often 2-3-toothed with a
secondary recurved lobe from the lower margin,
ascending lobes linear to narrowly obovate
contracted towards the base up to 20 mm long,
1-4 mm wide, often more than half the length
of the terminal lobe, terminal lobe linear to
narrowly obovate or oblong, contracted towards
the base, 5-25 mm long,1.5-6(-10) mm wide,
margin undulate to irregularly crenate, more
rarely entire, sometimes shallowly lobed,
sparsely to moderately hairy, hairs appressed,
crisped-appressed to spreading, 0.1-0.5 mm
long. In upper parts, leaves with shorter
petioles, lobes becoming narrower and, except
for the basal lobes, always entire. Inflorescence,
solitary, axillary, bracteolate, a one-sided
dichasium with lor 2 flowers, or occasionally

Austrobaileya 6 (1): 1-39 (2001)

with 2 inflorescences per axil; peduncle terete,
5-25 mm long, moderately to sparsely hairy,
hairs appressed to spreading, 0.1-0.6 mm long;
bracteoles opposite, occasionally alternate,
linear, 0.8-2.5 mm long, 0.25-0.6 mm wide,
apex acute, ciliate, sparsely to moderately hairy
on the back; pedicel 2-6(-8) mm long, recurved
at fruiting, hairs denser and more appressed
than on the peduncle. Outer sepals obovate,
obovate-oblong to broadly elliptic, 3-5 mm
long, 2.5-3.5 mm wide, apex acute to rounded
with a short recurved apiculum, ciliate,
moderately to densely hairy outside, hairs short
appressed, ascending or erect, 0.1-0.6 mm
long; inner sepals orbicular to obovate-
orbicular or obovate-elliptic, 3-6 mm long, 2-
4.5 mm wide, apex obtuse to rounded or slightly
emarginate, with a short recurved apiculum,
face glabrous or with a few hairs around the
apex and down the spine. Corolla funnel-
shaped, white or pink, 5-8 mm long, 6-8 mm
diameter, flared 2-4 mm above the base of the
tube, petals 5-9 mm long, 2.5-6 mm wide, with
rounded-triangular to rounded-oblong,
emarginate to apiculate lobes, glabrous except
for hairs on the midpetaline band for 1.5-3.5
mm from the apex. Stamens 5, unequal,
filaments affixed to the corolla tube for 1-2.3
mm from the base, free for 1.2-3 mm, with low
scattered tubercles from 0.5 mm above the base
of the corolla and extending for 1-1.5 mm;
anthers ovate-oblong to oblong-elliptic, 0.5-1
mm long, 0.35-0.7 mm wide, apex rounded,
emarginate, often apiculate, base sagittate, basal
lobes 0.15-0.25 mm long. Ovary ovoid, 1-1.5
mm long, on a disk 0.25 mm high, glabrous;
style 1.5-2.0 mm long, with cylindrical to
narrowly ovoid, obtuse stigmas, 0.6-2 mm long.
Capsule globular, 4-5 mm long, 4-5.5 mm
diameter, glabrous. Seeds 4,14-globular, 2.5-3.5
mm long, 2-2.5 mm wide, dark brown, surface
finely punctate bearing numerous laterally
compressed, wavy tubercles and an irregular,
narrow discontinuous wing of fused hair-like
structures (Lig. 10A&B).

Affinities: C. recurvatus most closely resembles
C. clementii in the small size of its corolla, its
much divided leaves and compound
inflorescence. However, it is readily
distinguished from that species by its short and
strongly recurved pedicels subtending the
mature capsules. The cauline leaves of

Johnson, Convolvulus L. in Australia

33

C. recurvatus also tend to be smaller and more Etymology: The specific epithet was chosen
finely divided. to highlight the characteristic recurved pedicels

which support mature capsules.

Two subspecies are recognised and can be distinguished as follows
Hairs on stems and leaves mainly ascending or spreading;

petals 7-9 mm long; outer sepals 4-5 mm long. 10a. C. recurvatus subsp. recurvatus

Hairs on stems and leaves mainly appressed;

petals 5-7 mm long; outer to 4 mm long. 10b. C. recurvatus subsp. nullarborensis

10a. Convolvulus recurvatus R.W. Johnson
subsp. recurvatus

Perennial with trailing and twining stems, with
± appressed hairs especially towards the tip,
but also with many spreading, 0.2-0.8 mm long.
Leaves subtending inflorescences petiolate;
petiole 2-20 mm long; blade 10-35 mm long, 5-
20 mm wide (Fig. 9A). Inflorescence with 1-2
flowers. Pedicel 2-6 (-8) mm long. Outer sepals
obovate, 4-5 mm long, 2.5-3.5 mm wide,
moderately to densely hairy outside, hairs
mainly erect or ascending, 0.25-0.6 mm long;
inner sepals 3.5-5 mm long, 3.0-4.5 mm wide.
Corolla 6-8 mm long; petals 7-9 mm long, 3-4
mm wide. Stamens affixed to the corolla tube for
1.5-2.3 mm from the base, free for 2-3 mm;
anthers 0.8-1.0 mm long, 0.65-0.7 mm wide,
basal lobes 0.2-0.25 mm long. Style 2.5-3.0 mm
long; stigmatic lobes 1.0-2.0 mm long.

Specimens examined : South Australia. Siam Station,
c. 100 km W of Pt Augusta, Apr 1921, Black s.n. (AD
97524062); between Pt Wakefield & Kilpara, c. 12
km NW of Port Wakefield, Sep 1967, Blaycock 606
(AD); Wangianna, 40 km W Marree on railway, Apr
1941, Cleland s.n. (AD 97218292); Mt Lyndhurst
Station, c. 45 km N of Leigh Creek, May 1924, Cleland
s.n. (AD 97217274); Curnamona, c. 110 km N of
Yunta, Dec 1930, Cleland s.n. (AD 97218289); Bute
District, c. 140 km NNW of Adelaide, Oct 1966,
Copley 827 (AD); gate, 3 km E of Ediacara, c. 50 km
W of Leigh Creek, Sep 1963, Lothian 2422 (AD).
New South Wales. Near Fowlers Gap, Sep 1952, Anon
(NE 21249); Fowlers Gap Station, 110 km N from
Broken Hill, Dec 1988, Browne 557 (LTB); Bogan
Gate, Apr 1924, Ising 2156 (NSW); 16 miles S of
Cobar on Nymagee road, Sep 1966, Moore 4488
(CANB); Nucha Lake, 30 km E of Fowlers Gap, Jun
1979, Paijmans 3331 (CANB); Trangie, May 1965,
Robards TR2 (NSW). Victoria. MURRAY MALLEE:
Red Cliffs, S of Mildura, Sep 1964, Beauglehole
ACB38486 (MEL); Wyperfeld NP, c. 0.5 miles SW of
Wonga Hut, Sep 1968, Beauglehole ACB28484 (MEL);
between Lakes Moumpoull and Kondarin, 9.7 km NE
of Hattah, Aug 1977, Cameron 8683 (MEL). LOWAN
MALLEE: Wyperfeld NP, Callitris Plain N boundary,
Oct 1968, Beauglehole & Finck ACB29216 (MEL).

Distribution and habitat : C. recurvatus subsp.
recurvatus is found south of about 30°S from
north of Leigh Creek in South Australia in the
west to Trangie in New South Wales in the east.
It extends south into far north-western Victoria
and southern South Australia to the west of
Port Augusta (Map 5). Its distribution partly
overlaps the south-eastern part of the range of
C. clementii. It grows on red brown earths and
sandy and loamy soils generally in low-lying
areas such as flood plains and dry lake bottoms.
It has been recorded from Eucalyptus
woodlands and mallee communities.

Phenology: Flowering occurs throughout the
year but mainly in late winter to early spring
(August-October); fruit have been recorded
mainly from early autumn to late spring (March-
November).

Notes: The specimens collected in the
Wyperfeld National Park area in Victoria cited
above are immature and further collections from
the area will be necessary to confirm their
identity.

10b. Convolvulus recurvatus subsp.
nullarborensis R.W. Johnson, subsp. nov.

affinis C. recurvato R.W. Johnson subsp.
recurvato sed corollis et sepalis
parvioribus et pilis plerumque adpressis
et brevioribus differt. Typus: South
Australia. Nullabor Region: c. 1 km NW
of Cook, 16 September 1960, P. Wilson
1692 (holo: AD, iso: BRI, MEL).

Perennial with mainly trailing, more rarely
twining stems, with appressed hairs 0.2-0.4 mm
long with an occasional longer spreading hair.
Leaves subtending inflorescences petiolate;
petiole 2-10 mm long; blade 7-25 mm long, 6-
20 mm wide (Fig. 9B). Inflorescence commonly
1-flowered, more rarely with 2 flowers. Pedicel

34

Austrobaileya 6 (1): 1-39 (2001)

Fig. 9. Variation in leaf shape of Convolvulus species from base to tip of stems. A. C. recurvatus subsp. recurvatus. (1)
Copley 823 (AD) x 1, (2) Robards TR2 (NSW) x 2, (3) Copley 823 (AD) x 2; B. C. recurvatus subsp. nullarborensis x 2.
(1) lsing s.n. (AD966061026), (2,3) Wilson 1710 (AD); C. C. clementii x 1. (1) Lay 254 (AD), (2,3) Law 3 (BRI); D. C.
tedmoorei. (1) Moore 5863 (CANB) x 1 (2) Moore 5863 (CANB) x 0.5, (3) McKean s.n. (CANB301718) x 1.
Del. W. Smith.

Fig. 10. Seeds of Convolvulus species, adaxial (L.H.S.) and abaxial (R.H.S.) surfaces x 10. A C. recurvatus subsp. recurvatus
from lsing 2156 (NSW); B. C. recurvatus subsp. nullarborensis from George 8482 (PERTH); C. C. clementii from George
3938 (PERTH); D. C. tedmoorei from McKean 5169 (CANB). Del. W. Smith.

Johnson, Convolvulus L. in Australia

35

2-5 mm long. Outer sepals obovate, obovate-
oblong to broadly elliptic, 3-4 mm long, 2.4-3
mm wide, moderately hairy outside, hairs short
appressed to ascending, 0.1-0.4 mm long; inner
sepals 3^4 mm long, 2-3 mm wide. Corolla 5-6
mm long, petals 5-7 mm long, 2.5-3 mm wide.
Stamens affixed to the corolla tube for 1-1.4 mm
from the base, free for 1.2-2.75 mm; anthers 0.5-
0.7 mm long, 0.35-0.6 mm wide, basal lobes up
to 0.15 mm long. Style 1.5-1.9 mm long; stigmatic
lobes, 0.6-1.4 mm long.

Specimens examined : Western Australia. 2 miles S
of Reid, Nullarbor Plain, Sep 1962, Aplin 1671
(PERTH); Nullarbor Plain, c. 115 miles NW of Reid,
Oct 1966, George 8482 (PERTH). South Australia.
Nullarbor Region: Nullarbor Plain N part, c. 22 km N
of Cook, Aug 1980, Donner 7216 (AD); Hughes, Sep
1920, Ising 1528 (AD, BRI, MEL); c. 1 km NW of
Cook, Sep 1960, Wilson 1692 (AD, BRI, MEL); 6 km
E of Watson, c. 240 km E of WA border on railway
line, Sep 1960, Wilson 1710 (AD, BRI). Flinders Ranges
Region: Leigh Creek on railway to Alice Springs, Oct
1953, Lothian s.n.(AD); Leigh Creek, opp. Scout Hut,
c. 230 km NNE of Port Augusta, Oct 1969, Lothian
5053 (AD). Eastern Region: Koonamore Station, c. 60
km N of Yunta, Dec 1926, Black J.M.Herb s.n. (AD);
Koonamore Vegetation Reserve, c. 400 km NNE of
Adelaide, May 1971, Crisp 181(AD); Koonamore
Vegetation Reserve, 60 km N of Yunta, Mar 1962,
Lange s.n. (AD); Koonamore, c. 60 km N of Yunta,
Nov 1927, Paltridge s.n. (AD). Eyre Penninsula Region:
Gawler Range, 2 km W of Wartaka H/S, Oct 1983,
Weber 7921 (BRI).

Distribution and habitat : C. recurvatus subsp.
nullarborensis occurs in Western Australia and
South Australia on the Nullarbor Plain (Map
5) where it grows in chenopod shrublands on
clay loam soils derived from limestone. Other
populations recorded from central South
Australia have been assigned to this subspecies.
Three specimens from the Koonamore area
appear to belong to this taxon. In particular
Crisp 181 clearly resembles Ising 1528 which
was collected from Hughes on the Nullarbor
Plain. Two specimens from Leigh Creek
0 Lothian s.n. (AD96212336) and Lothian 5053
possess the short recurved fruiting pedicels and
short appressed hairs of C. recurvatus subsp.
nullarborensis and though the leaves appear
much less divided I have included them under
this subspecies. The latter populations grow in
arid shrub steppes.

Phenology: Flowering and fruiting have been
recorded from late winter to mid spring, from

August to October.

Etymology: The subspecific epithet refers to
the Nullarbor Plain from which the most typical
specimens of this subspecies have been
collected.

11. Convolvulus clementii Domin, Biblioth.
Bot. 89: 539 (1928) Type: Nordwest-
Australien: zwischen Ashburton und De
Grey River, E. Clement (holo: PR).

Convolvulus clementii var. biflorus Domin,
Biblioth. Bot. 89: 539 (1928). Type:
Queensland. Sandsteinhugel der
Dividing Range bei Jericho, Domin III
1910 (holo: PR).

Perennial with trailing and twining stems; stems
terete, occasionally with low ribs, moderately
densely to sparsely hairy, becoming sparser
with age, hairs ± appressed especially towards
the tip of the stem but in lower parts ascending
to spreading hairs are common, 0.2-0.8 mm
long. Leaves variable in shape and size from
base to tip of the stem (Fig. 9C). Basal leaves
petiolate; petiole often equal to or longer than
the blade, 6-30 mm long; blade oblong to ovate,
sometimes triangular, sometimes hastate or
auriculate, 7-30 mm long, 5-15 mm wide, apex
obtuse to rounded, occasionally emarginate,
mucronulate, base truncate to cordate, margins
± entire to undulate, soon becoming lobed with
a prominent pair of basal lobes, a pair of
ascending lobes and a terminal lobe, all
undulate to irregularly lobed, + glabrous but
soon becoming moderately hairy, hairs mainly
spreading, 0.3-0.5 mm long. Leaves subtending
inflorescences petiolate; petiole 2-30 mm long;
blade ovate in outline, 10-60 mm long, apex
obtuse to rounded, emarginate, mucronulate,
base cordate with a broad sinus, deeply lobed,
basal pair of lobes linear or narrowly-oblong,
2-15 mm long, with a secondary recurved lobe
from the lower margin, often 2-3 toothed,
ascending lobes linear to narrowly oblong,
occasionally slightly obovate, to 30 mm long,
1-3 mm wide, terminal lobe linear to linear-
obovate, up to 55 mm long, 2-10 mm wide,
occasionally triangular-oblong and up to 30 mm
wide, margin entire, undulate, toothed to
shallowly lobed, moderately to sparsely hairy
becoming glabrous, usually slightly denser

36

below, hairs crisped-appressed to spreading,
0.2-0.6 mm long. In upper parts, leaves deeply
lobed, lobes becoming narrower and aside from
the basal lobes, with entire margins.
Inflorescence solitary, axillary, bracteolate, a
one-sided dichasium with lor 2, rarely 3 flowers,
or occasionally with 2 inflorescences per axil;
peduncle terete, 5-50 mm long, moderately
densely to sparsely hairy, hairs crisped
appressed, ascending and spreading, 0.2-0.6
mm long; bracteoles opposite to alternate, linear
to subulate or rarely narrowly elliptic, 1-4.5 mm
long, 0.25-0.5 mm wide, apex acute, ciliate,
moderately hairy to ± glabrous on the back;
pedicel 3-15 mm long, rarely longer at fruiting,
straight to sinuate or slightly curved at fruiting,
hairs often denser and more appressed than for
peduncle. Outer sepals obovate, orbicular to
broadly elliptic, 4-5.5(-6) mm long, 2.5^4-.5 mm
wide, apex barely acute to rounded with a
recurved apiculum, ciliate, moderately to
sparsely, occasionally densely, hairy outside,
hairs crisped-appressed, ascending and
spreading, 0.2-0.7 mm long; inner sepals
orbicular, obovate to obovate-elliptic, 3.5-5.5
mm long, 2.5-4 mm wide, apex obtuse to
rounded-truncate with a recurved apiculum,
base rounded to truncate, eciliate or with a few
hairs on the upper margin, face + glabrous with
some hairs around the apiculum and midrib.
Corolla funnel-shaped, white to pink, rarely
purplish, 5-8 mm long, 4-10 mm diameter, flared
2.5-4 mm above the base of the tube; petals 6-
9 mm long, 1.5-4 mm wide, with rounded-
triangular, broadly emarginate lobes, with a short
apiculum, glabrous except for hairs on the
outside of the midpetaline band for 1-3 mm from
the apex. Stamens 5, slightly unequal; filaments
affixed to the corolla tube for 1.5-2.5 mm from
the base, free for 1.5-3 mm, with scattered low
tubercles, mainly along the margins, from 0.5
mm above the base of the corolla and extending
for 1-3 mm; anthers oblong, oblong-elliptic to
triangular-ovate, 0.5-0.9 mm long, 0.3-0.5 mm
wide, apex obtuse to rounded emarginate, base
sagittate, basal lobes 0.1-0.3 mm long. Ovary
ovoid to globular ovoid, 1-1.5 mm long, on a
disk 0.2-0.25 mm high, glabrous; style 2-3 mm
long, with cylindrical to narrowly ovoid,
sometimes falcate, obtuse stigmas, 1.3-1.9 mm
long, often suffused with purple. Capsule
globular to globular-ovoid, 4-7 mm long, 4-6

Austrobaileya 6 (1): 1-39 (2001)

mm diameter, glabrous. Seeds 4,14-globular, 2.5-
3.2 mm long, 2-2.7 mm wide, dark brown to black,
surface finely punctate bearing irregular short
raised ridges and tubercles, made of fused hairs
and a serrated to + continuous wing of fused
hairs to 0.25 mm wide (Fig. 10C).

Selected specimens (170 specimens examined):
Western Australia. Between Menzies and Comet Vale,
Sep 1939, Blackall 4193 (PERTH); 4.4 miles N of
Meekatharra, Oct 1973, Demarz 4701 (PERTH); 4
km S of Mt Magnet, Sep 1984, Demerz 10196
(PERTH); 1 mile E of Yanrey Homestead, Aug 1960,
George 1154 (PERTH); Wingkilina, Hinckley Ranges,
Jun 1981, Kalotas 868 (BRI). Northern Territory.
Argadargada H/S Bore, Sep 1954, Chippendale 308
(DNA); 1 mile S of Elkedra H/S, Oct 1956, Chippendale
3097 (DNA); Mt Ebenezar H/S, 57 km W of Stuart
Highway on road to Ayers Rock, Sep 1979, Johnson
3364 (BRI); 10 miles SSW of Alice Springs, Oct 1956,
Lazarides 6101 (DNA); Uluru NP - Docker River road, 51
km WNW of Ranger Station, Aug 1988, Lazarides &
Palmer 542 (BRI). South Australia. Far East N of
Curnamona Gate & boundary fence, c. 130 km N of Yunta,
Nov 1962, Lothian 1216 (AD); NW Region Mt Davies,
Tomlinson Range, Sep 1955, SA Pastoral Board 107 (AD);
Lake Eyre Basin NE, Cadelga Waterhole, 10 km S & 60
km E[W] of Queensland border, Aug 1973, SA Pastoral
Board s.n. (AD); Everard Range at foot of Mt Areteinna,
Sep 1968, Spooner 136 (AD); Lake Frame NE, c. lOkmW
of Quinyambie H/S, Jul 1971, Whibley 3535 (AD).
Queensland. Burke District: 15 km NW of Richmond,
May 1974, Byrnes 3047 (BRI). Gregory North District:
13 km S of Boulia, Oct 1984, Neldner 1631 (BRI).
Mitchell District: Winton, Jul 1934, Blake 6551 (BRI).
Leichhardt District: Wandoan, Nov 1930, Hubbard
5021 (BRI). Gregory South District: 35 miles ENE of
Nappamerry, May 1971, Silcock S363 (BRI). Warrego
District: 6 km E of Charleville along Morven road,
Mar 1976, Purdie & Boyland 200/23 (BRI). Maranoa
District: 8 miles E of Weengallen, Nov 1961, Pedley
914 (BRI). Da rti ng Downs District: c. 12 miles W of
Meandarra, Mar 1959, Johnson 749 (BRI). New South
Wales. Donalds Plain, 75 km W of Cobar, Oct 1963,
Constable 4655 (NSW); Bulloo River, in 1887,
Lockhart Morton s.n. (MEL); NE edge of Narran Lake,
near Brewarrina, Nov 1967, McGillivray 2856A (NSW);
Paika, Balranald, Mar 1910, McPherson s.n. (MEL);

11 km E of Warratta Bore, 32 km E of Milparinka,
Oct 1976, Wilson 1643 (NSW). Victoria. Murray
Mallee: Red Cliffs, Nov 1987, Browne 474 (LTB, BRI).

Distribution and habitat : C. clementii is found
south of about 20° S, from around Dampier in
Western Australia across the Northern Territory
to Richmond in Queensland. It extends
southwards to the southern parts of Western
Australia and across southern Australia to far
north-western areas of Victoria. It is common
throughout semi-arid New South Wales and
Queensland and is absent within this region in

Johnson, Convolvulus L. in Australia

37

Map 8. Distribution of Convolvulus clementii. Del. W. Smith. 0 &

V

only the most arid areas of the continent (Map
8). It is found mainly on plains, often in flooded
and swampy situations, but also on clay pans
between sand dunes. It grows mainly in grassy
woodlands, commonly with Eucalyptus
populnea, E. intertexta and Acacia aneura,
where it occurs in red-brown sandy and loamy
duplex soils and in sands. It is also found in
grassy woodlands with E. coolabah and
E. largiflorens along floodplains where soils are
usually brown to grey clays, sandy clays to clay
loams. It is occasionally found in Astrebla
grasslands.

Affinities: It appears most closely related to
C. recurvatus which can be distinguished from
this species by its short pedicels which are
strongly recurved at fruiting.

Phenology: Flowering and fruiting have been
recorded throughout the year though flowering
may be more common in late winter to mid
spring (August-October) and fruit are found
mainly in the early autumn to late spring
(March-No vember).

Etymology: This species was named in honour
of Dr E. Clement, a botanical collector who lived
in Western Australia and who sent the type
specimen to Dr K. Domin in Prague.

Conservation Status: This species is
widespread.

12. Convolvulus tedmoorei R.W. Johnson, sp.
nov. affinis C. clementio Domin sed
caulibus et foliis multis grossis et
seminibus grandioribus et alis destitutis
differt. Typus: New South Wales. 5 miles
[8 km] NW of Louth, 7 September 1971,
C.W.E. Moore 5863 (holo: CANB; iso:
BRI, NSW).

Perennial with trailing stems; stems coarse,
terete, moderately to sparsely hairy, hairs
weakly ascending, 0.25-0.4 mm long. Leaves
somewhat similar in shape from base to tip of
the stem (Fig. 9D). Basal leaves petiolate;
petiole long, often as long as the blade; blade
ovate, apex rounded to emarginate, base
cordate, shallowly lobed particularly at the
base, sparsely hairy, hairs as for the stem.
Leaves on fertile stems petiolate; petiole 10-
40 mm long; blade broadly ovate, oblong or
oblong-triangular, 20-50 mm long, 15-40 mm
wide, apex rounded to emarginate, base cordate,
decurrent, basal lobes broad, 5-10 mm long,
with 2 or 3 rounded emarginate lobes,
ascending lobes oblong, 5-12 mm long, 2-8 mm
wide, terminal lobe oblong 10-25 mm long, 3-
20 mm wide, margin undulate to shallowly lobed,
moderately to sparsely hairy, hairs crisped and

38

loosely appressed to weakly ascending, 0.25-
0.4 mm long. In upper parts, leaves similar in
shape but shorter and with narrower lobes.
Inflorescence solitary, axillary, bracteolate, a
one-sided dichasium with 1 or 2 flowers, or
commonly with 2 inflorescences in each axil;
peduncle terete, 7-30 mm long, moderately hairy,
hairs crisped-appressed to weakly ascending,
0.25-0.5 mm long; bracteoles opposite to sub¬
opposite, narrowly triangular, 1.5-2.5 mm long,
acute, ciliate, moderately to sparsely hairy on
the back; pedicel thicker than the peduncle,
dilated upwards, 6-13 mm long, more densely
hairy than the peduncle. Outer sepals elliptic to
obovate, becoming orbicular, 5-6 mm long, 4.5-
5.5 mm wide, obtuse to barely acute with a short
recurved apiculum, becoming rounded
acuminate, mucronate, ciliate particularly in the
upper half, sparsely to moderately hairy outside,
hairs ascending 0.15-0.3 mm long; inner sepals
obovate, 5-6 mm long, 4.8-6 mm wide, apex
rounded, shortly apiculate, base truncate,
glabrous except for short cilia around the apex.
Corolla funnel-shaped, 5-lobed, white to pink?,
c. 7 mm long and c. 8 mm diameter, flared c. 4.5
mm above the base of the tube; petals 8-9 mm
long, 2.5-3 mm wide, with well developed
oblong, rounded, emarginate lobes, glabrous
except for dense hairs on the outside of the
midpetaline band for c. 1.5 mm from the tip.
Stamens 5, unequal; filaments affixed to the
corolla tube for c. 2 mm from the base, free for
2-3 mm with low tubercles, mainly along the
margins, from almost the base of the corolla
tube and extending for c. 2.5 mm; anthers ovate
to triangular-ovate, 0.75-0.85 mm long, 0.6-
0.65 mm wide, apex rounded-emarginate, base
sagittate, basal lobes 0.1-0.2 mm long. Ovary
ovoid, 1.25-1.3 mm long on a disk, 0.3 mm
high, glabrous; style stout, c. 3.5 mm long, with
stout cylindrical stigmas, 1-1.1 mm long.
Capsule globular, 6-7 mm long and wide,
glabrous, 2-valved. Seeds 4, !4-globular to X A-
globular-obovoid, 3.5-3.8 mm long, 2.8-3.2
mm wide, very dark brown to black, surface
finely punctate bearing numerous laterally
compressed and wavy tubercles, with no
obvious wing (Fig. 10D).

Specimens examined: New South Wales. Toganmain
Station, Darlington Point, Jun 1969, McKean 5169
(CANB); 5 miles NW of Louth, Sep 1971, Moore 5863
(CANB, BRI, NSW).

Austrobaileya 6 (1): 1-39 (2001)

Distribution and habitat : C. tedmoorei is
known from only two areas on the flood plains
of the Darling and Murrumbidgee Rivers in
central western New South Wales (Map 5). It
grows in self-mulching grey clay soils.

Affinities: This species most closely resembles
C. clementii but it can be distinguished by its
more prostrate and fleshy habit, its coarse stems
and its larger capsules and seeds. The seed
surface structure is also a distinguishing
character as can be seen in Fig. 10, as is the
lack of a wing on the seed.

Notes: Only one flower of this species was
available for dissection so information on its
floral attributes is limited.

Etymology: This species is named in honour
of C.W.E.(Ted) Moore, a distinguished CSIRO
plant ecologist, who brought this taxon to my
notice and collected one of the two known
collections of it.

Conservation Status : This species is known
from only two localities and both collections
were made about thirty years ago. It is certainly
a rare species and should possibly be accorded
a higher conservation status than rare.

Acknowledgements

I am deeply grateful for the assistance provided
by Jeny Calway and Hans Dillewaard in
growing and collecting data from plants in the
BRI glasshouse at Indooroopilly in Brisbane.
In particular, I would like to thank Hans for his
photographic assistance. The generous help
given by John Parham in collecting material
from key areas in Tasmania is also appreciated.
Many thanks are also due to Will Smith for the
illustrations. I would also like to thank the
referees for their careful reading of the
manuscript and helpful co mm ents.

The curators of AD, HO, MEL, NE, NSW
and PERTH are thanked for providing loans of
specimens for this study. My special thanks
are given to Dr G.P. Guymer, Director of the
Queensland Herbarium, for allowing me to use
the herbarium and providing space and facilities
to enable me to continue my research.

Johnson, Convolvulus L. in Australia

39

References

Austin, D.F. (1973). The American Erycibeae
(Convolvulaceae): Maripa, Dicranostyles and
Lysiostyles 1. Systematics. Annals of the Missouri
Botanic Garden 60: 306-412.

-(1975). Convolvulaceae. In R.E. Woodson Jr &

R.W. Schery (eds), Flora of Panama. Annals of
the Missouri Botanic Garden 62: 157-224.

-(1982). Convolvulaceae. In Z.L. de Febres &

J.A. Steyermark (eds), Flora of Venezuela 8(3):
15-239. Caracas: Instituto Nacional de
Parques.

-(1982a). Convolvulaceae. In G. Harling & B.

Sparre (eds), Flora of Ecuador No. 15. Stockholm:
Swedish Research Councils.

-(1998). Convolvulaceae. In J.A. Steyermark, PE.

Berry& K. Holst (eds), Flora of Venezuelan
Guiana 4: 377^-24. St Louis: Missouri Botanic
Gardens Press.

Bentham, G (1869). Convolvulaceae. In Flora Australiensis
4: 410-442. London: L. Reeve & Co.

Bentham, G. & J.D. Hooker (1873). Genera Plantarum
2: 874-875. London: L. Reeve & Co.

Brown, R. (1810). Prodromus Florae Novae
Hollandiae et Insulae van Diemen. London:
Johnson & Co.

Choisy, J.D. (1845). Convolvulaceae. In A. de Candolle,
Prodromus systematis naturalis regni vegetabilis
9: 323^162. Paris: Fortin, Masson.

Domin, K. (1928). Convolvulus. Beitrage zur Flora und
Pflanzengeorgraphie Australiens. Bibliotheca
Botanica 89(6): 1093-1094.

Hitchock, A.S. (1929).Nomenclature; proposals by
British botanists. In International Botanical
Congress 5 th ed., 1930. Cambridge, England.

Johnson R.W. (1986). Convolvulaceae. In J.P Jessop &
H.R. Toelkin (eds), Flora of South Australia
Part III: 1133-1145. Adelaide: South Australia
Government Printing Division.

-(1987). Two new species of Convolvulus L.

(Convolvulaceae) from South Australia.
Austrobaileya 2 (4): 408-411.

Lehmann, J.G.C. (1826). Semina in horto botanico
Hambergensi 1826 collecta quae pro mutua
commutatione offeruntur. Hamburg: Meissner.

Linnaeus, C. (1753). Species Plantarum. London: Ray
Society.

Mabberley, D.J. (1989). The Plant-Book. Cambridge:
Cambridge University Press.

-(1997). The Plant-Book. Second Edition.

Cambridge: Cambridge University Press.

Meeuse, A.D.J. (1957). The South African Convolvulaceae.
Bothalia 6: 641-792.

Mueller, F. (1853). Diagnosis et descriptions plantarum
novarum quas in Nova Hollandia australi
praecipue in regionibus interioribus. Linnaea 25:
423^424.

-(1864). The Vegetation of the Chatham-Islands.

Melbourne: Government Printer.

Ooststroom, S.J. van (1953). Convolvulaceae. In C.G.G.J.
Steenis (ed.). Flora Malesiana, Ser.1.4: 388-513.
Djakarta: Noordhoff-Kolff.

Sa’ad, F. (1967). The Convolvulus species of the Canary
Isles, the Mediterranean region, and the Near and
Middle East. Rotterdam.

Sims, J. (1808). Curtis’s Botanical Magazine 27:1.1067.

Vriese, W.H.de (1845). Convolvulaceae. In J.GC. Lehmann
(ed.) Plantae Preissianae 1: 345-347. Hamburg:
Meissner.

Woolls W. (1867). Plants Introduced Accidentally (1866).
In. A Contribution to the Flora of Australia: 136—
152. Sydney: F. White.

(1754). Genera Plantarum. Stockholm:
Laurentii Salvii.

Vanguerieae A.Rich. ex Dum. (Rubiaceae) in Australia, 2.
Cyclophyllum Hook.f.

S.T. Reynolds & R.J.F. Henderson

Summary

Reynolds, S.T. & Henderson, R.J.F. (2001). Vanguerieae A.Rich. ex Dum. in Australia, 2. Cyclophyllum
Hook.f. Austrobaileya 6 (1): 41-66. As a result of critical studies of Australian species that have, in the
past, been included in Canthium Lam., the genus Cyclophyllum Hook.f. (Rubiaceae, Vanguerieae) is
now accepted as occurring in Australia. A revision of this genus in this continent is presented here. Nine
of its species occur in Australia, five of which are described here as new, namely C. longipetalum
S.T.Reynolds & R.J.F.Hend., C. maritimum S.T.Reynolds & R.J.F.Hend., C. multiflorum S.T.Reynolds
& R.J.F.Hend., C. protractum S.T.Reynolds & R.J.F.Hend. and C. rostellatum S.T.Reynolds &

R. J.F.Hend. The new combinations C. brevipes (Merr. & L.M.Perry) S.T.Reynolds & R.J.F.Hend.,
C. coprosmoides (F.Muell.) S.T.Reynolds & R.J.F.Hend., C. costatum (C.T.White) S.T.Reynolds
& R.J.F.Hend. and C. schultzii (O.Schwarz) S.T.Reynolds & R.J.F.Hend. are provided for the
remaining four. The new combination C. coprosmoides var. spathulatum (O.Schwarz) S.T.Reynolds
& R.J.F.Hend. is provided for a distinctive variety of C. coprosmoides, and C. schultzii
forma angustifolium S.T.Reynolds & R.J.F.Hend. for a distinctive form of C. schultzii. All recognised
taxa are described, and keys to identify them as well as maps showing their known distribution are
provided. Line drawings representing some of the recognised taxa are also provided.

Keywords: Rubiaceae, Vanguerieae, Cyclophyllum, Australia

S. T. Reynolds & RJ.F. Henderson cl- Queensland Herbarium, Environmental Protection Agency, Brisbane
Botanic Gardens Mt Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Introduction

Psydrax Gaertn. and Cyclophyllum Hook.f.
have of recent times been combined under
Canthium Lam., but based mainly on the work
of Bridson (1985, 1987, 1992) and studies of
much Australian material, these genera plus
Everistia, the only other genus with taxa
previously included in Canthium in Australia,
are now considered worthy of recognition in
Australia. As a result, the genus Canthium in
the strict sense is found not to occur in this
country. A key to distinguish Cyclophyllum
from Psydrax and Everistia has been provided
by Reynolds and Henderson (1999).

Bridson (1987) accepted that there existed
a group of species allied to Pyrostria
Commerson ex Juss. from Africa, and centred
on the New Caledonian Cyclophyllum
deplanchei Hook.f., which could be recognised
as either a distinct genus with the name
Cyclophyllum Hook.f., an infra-generic group
within Pyrostria or even as an infra-generic
group within Canthium. At that time, she left
the question of what rank to recognise this

Accepted for publication 16 March 2001

group at as a matter still “to be settled”. She
diagnosed the group and listed Canthium barbatum
(GForst.) Seem, and C. sessilifolium A.Gray, from
Fiji, C. brevipes Merr. & L.M.Perry, C. caudatum
(Valeton) S.Moore, C. longiflorum (Valeton) Merr.
& L.M.Perry and C. valetonianum S.Moore from
New Guinea, and C. coprosmoides F.Muell. and
C. costatum C.T.White from Australia as species
she believed “should be considered for transfer to
Cyclophyllum ,, .

Smith & Darwin (1988, p.232), in dealing
with the group in Fiji, recognised Cyclophyllum
as a distinct genus and included three species
from that country in it. Guilluamin had earlier
(Guilluamin, 1948) recognised 14 species from
New Caledonia as belonging to this genus. As
stated above, our studies have led us to accept
that Cyclophyllum should be recognised as a
distinct genus.

Although the species treated here as
Cyclophyllum costatum (from north Queensland)
and C. schultzii (from the Northern Territory) are
very distinctive, herbarium material that was
previously included under Canthium

42

coprosmoides (from Queensland) was found to
be very variable and, in fact, represented a
number of undescribed taxa in addition to
Cyclophyllum coprosmoides in the strict sense.
These new taxa have been formally described
in this account.

As stated previously (Reynolds &
Henderson, l.c.), this study was based mostly
on herbarium material, but measurements given
for leaves, inflorescences, flowers and fruits are
based on dried, fresh or spirit material. In the
list of specimens cited, only the herbaria from
which specimens have been seen are recorded.
State subdivisions (pastoral districts) are
provided for Queensland collections only. The
taxonomic concepts accepted here are those of
the first author and result from her many years
of detailed herbarium studies.

Taxonomy

Cyclophyllum Hook.f. in Benth. & Hook, f., Gen. PI.
2: 535 (April 1873). Canthium sect.
Cyclophyllum (Hook,f.) Baillon, Adansonia 12:
184 (1879); from Greek cyclo- (circular) and -
phyllus (-leaved), in reference to the leaf blades
of the type.Type: Cyclophyllum deplanchei
Hookf.

Trees or shrubs, usually with horizontal
branches; branchlets without spines. Leaves
stipulate, petiolate, usually coriaceous; stipules
interpetiolar, ovate to deltoid, acuminate or with
a subulate tip. Inflorescences axillary, of 1 to
few-flowered, umbelliform, sessile or shortly
pedunculate cymes, or sometimes with flowers
along rudimentary inflorescence branchlets.
Flowers in bud obtuse or acuminate at apex, 5-
merous (sometimes 4-merous in C. costatum ),
pedicellate, each subtended by a single bract;
calyx tube cupuliform except in C. costatum
where long attenuated into pedicel; corolla tube
slender, longer than lobes, hypocrateriform,
coriaceous, hairy adaxially with dense, white,
moniliform hairs projecting from its mouth and
obscuring anthers; stamens (4 or) 5, with
filaments very short, and anthers ovoid,
dorsifixed, introrse, the locules attached by their
dorsal surface to a brown coloured connective
with pallid margins; ovary 2-locular; ovules

Austrobaileya 6 (1): 41-66 (2001)

solitary in each locule, pendulous; style as long
as the corolla tube; stigma attached to the style
at its rounded base, fleshy, capitate, usually
obscurely 2-lobed. Fruit transversely ellipsoid,
broadly obovoid to obcordiform, smooth, entire
(or ribbed in C. costatum)-, pyrenes exceedingly
woody, usually broadly hemispherical and
depressed distally, smooth or slightly granular
or rugose when dry.

Distribution: About 30 species, occurring in
New Caledonia, Fiji, the Moluccas, New
Guinea and Australia; nine species in Australia.

Notes: The genus Cyclophyllum is characterised
by its axillary, fasciculate inflorescences, and
by its fleshy flowers with a long
hypocrateriform corolla tube with moniliform
hairs protruding from the throat, introrse
shortly stalked anthers with a dorsal brown-
coloured connective, style which shortly
exceeds the corolla tube, and capitate stigma
which is attached to the style at its convex base.

In the Australian species, the flowers are
clustered on very reduced, thickened, broad,
knob-like peduncles, each flower being
subtended by a bract, or the flowers and bracts
are clustered distally on a short, more or less
slender peduncle.

Affinities: Cyclophyllum Hook.f. is closely
related to Pyrostria Commerson ex Juss. from
Africa, and Bridson (1987) included it as one
of five informal groups she recognised within
that genus. However, Guilluamin (1948) and
Smith & Darwin (1988) recognised it as a
distinct genus. Cyclophyllum is also related to
Everistia and Psydrax and resembles those
genera in the placement of its cotyledons, but
differs from them by its long fleshy
hypocrateriform corolla tube which is always
longer than the corolla lobes, anthers with a
distal appendage, and its capitate stigma. It
resembles Canthium, with which it was
previously combined, in its habit, inflorescence
attributes, anthers with a dark dorsal
membranous connective, and in style and
stigma attributes, but the latter genus has
cotyledons placed perpendicular to the ventral
face of the seed and a consistently shorter
corolla tube.

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

43

Key to species of Cyclophyllum in Australia

1. Calyx tube elongate, attenuate at base into a slightly winged pedicel, together

with pedicel 11-19 mm long; fruits strongly ridged; leaves thin, slightly

membranous when dry.1. C. costatum

Calyx tube short, cupular, attached to a slender, rounded pedicel, together
with pedicel less than 10 mm long; fruits evenly rounded or slightly
ridged; leaf blades ± coriaceous when dry.2

2. Domatia present on leaf blades, usually conspicuous.3

Domatia absent on leaf blades or if present, small and inconspicuous. 5

3. Cymes (2-)5-12-flowered, sessile; branchlets densely covered with

exceedingly conspicuous lenticels; leaf blades (6.7-)9-13 cm long.7. C. multiflorum

Cymes (lor)2-5-flowered, shortly stalked or sessile; branchlets with few,
usually inconspicuous lenticels or lenticels absent; leaf blades usually

5.5- 9.5 cm long.4

4. Corolla 8.5-9 mm long; cymes 2-5-flowered, sessile; leaf blades 5.6-7.7

(-8.3) x 2.7-3.2(-4) cm, usually abruptly acuminate at apex; petioles

3-8 mm long.5. C. protractum

Corolla 13-14 mm long; cymes (1 or)2-4-flowered, on short peduncles
or sessile; leaf blades 5.5-7(-9.5) x (2.2-)3.3-5.2 cm, acuminate or
subacute at apex; petioles 5-16 mm long.4. C. longipetalum

5. Inflorescences distinctly pedunculate; peduncles (1 or) 2-6 mm long. 6

Inflorescences sessile or with reduced peduncles; peduncles (where present)

0.5-1.5 mm long.7

6. Peduncle branched or simple; cymes (2-)7-ll(-16)-flowered; leaf blades

8.5- 16.2 x 1.8-7.2(-8.2) cm; fruits 8-12 x 16-23 mm, usually

obcordiform, broadly 2-lobed distally; corolla 6-9 mm long.8. C. schultzii

Peduncle simple; cymes 2-7-flowered; leaf blades 3.8-8.2 x 2.1-4.6(-5.2)
cm; fruits 8.5-11.5 x c.8.5 mm, transversely ellipsoid, usually depressed
distally; corolla 9-10 mm long.6. C. maritimum

7. Cymes (4-)7-10-flowered; bracts glabrous; leaf blades usually acuminate

or subcaudate at apex, 6.2-14.7 x 2.5-5.5 cm, drying blackish coloured

or dark brown on both surfaces.9. C. brevipes

Cymes (1 or)2-4-flowered; bracts pubescent; leaf blades rounded, obtuse,
retuse, subacute or subacuminate at apex, 3.6—11(—13.5) x 1.6-5.2(-6.7)
cm, drying dark or greyish brown adaxially, pale grey or yellow green
abaxially.8

8. Petioles 8-10 mm long; cymes (l-)3-6-flowered; corolla 10-12 mm

long, cream to orange or brownish coloured; apex of flower buds and
corolla lobes acuminate or rarely obtuse; bracts usually with rusty brown

hairs.3. C. rostellatum

Petioles 4-7 mm long; cymes (1 or )2-4-flowered; corolla 6.5—8(—11) mm
long, cream or deep yellow coloured; apex of flower buds and corolla
lobes obtuse; bracts with white or pale brown coloured hairs or

glabrous.2. C. coprosmoides

44

Austrobaileya 6 (1): 41-66 (2001)

Conspectus of some diagnostic attributes in Cyclophyllum

Leaf blades membranous when dry; fruits usually strongly ribbed

C. costatum

Leaf blades with prominent domatia

C. longipetalum,
C.multiflorum,

C. protractum

Leaf blades usually with small, obscure domatia

C. brevipes,

C. coprosmoides,

C. maritimum,

C. schultzii

Inflorescences usually 5- or more-flowered

C. maritimum,

C. multiflorum,

C. schultzii,

C. brevipes

Inflorescences usually 4- or less-flowered

C. coprosmoides,

C. costatum,

C. longipetalum,

C. protractum

1. Cyclophyllum costatum (C.T.White)
S.T.Reynolds & R .J.F.Hend., comb, nov.;
Canthium costatum C.T.White, Contr. Am.
Arb. 4: 99 (1933). Type: Queensland.
Cook District: Daintree River, 11 March
1932, L.J. Brass 2261 (holo: ?An.v.; iso:
BRI).

Trees 4-8 m high; bark brown or whitish
coloured, slightly fissured; branchlets slightly
angular distally, reddish brown coloured but
usually with a white bloom, the older ones with
small, whitish coloured lenticels. Leaves with
stipules deltoid, keeled, attenuated into a narrow
subulate apex 6-7 mm long; petioles (1-) 6-7 mm
long; blades elliptic to ovate-elliptic, (4.2-)7-
7.7 x (1.9-)3.3-4.2 cm, with apex acute or
abruptly obtuse and subacuminate, and base
abruptly obtuse or subacute and attenuate into
petiole, glabrous, thick and fleshy but
membranous when dry, pale green in colour,
glossy adaxially and matt abaxially; midrib
slender, slightly sunken adaxially but
prominently raised abaxially; lateral nerves in
4-7 pairs, extremely slender, arcuate and looping
at margins; reticulate venation obscure; domatia
absent. Cymes 1-3-flowered, with flowers
fasciculate on a short, thick, knob-like peduncle.
Flowers in bud acute at apex, 4 or 5-merous,
strongly perfumed; pedicels 10-17 mm long,
slender; calyx elongate with a long, flattened,
narrowly turbinate, longitudinally ribbed tube
attenuating into the pedicel, together with

pedicel 11-19 mm long, with a short, 4- or 5-
denticulate limb; lobes subequal, narrowly
ovate to ovate, 1-1.5 mm long, unicostate;
corolla pale yellow, 13-15 mm long, with tube
10-11 mm long, sparsely hairy at mouth; lobes
ovate, cuculate at apex, abruptly long-
acuminate or cuspidate with a long acumen,
tricostate, 3-3.5 x 1.5-1.75 mm, scurfy
adaxially, densely papillose abaxially on and
near margins with usually 2 rows of papillae;
disc shorter than calyx limb, glabrous; stamens
with anthers ellipsoid or + oblongoid,
apiculate, c.lmm long. Fruits on pendulous
pedicels 6-11mm long, reddish coloured when
ripe, laterally compressed and 2-lobed, broadly
obovate to + obcordate in broad side view, 1.3-
1.4 x c.1.4 cm, strongly angled with 8 or 9
prominent longitudinal ridges, or fruits
ellipsoidal and c.10 mm x 6 mm; pyrenes
slightly rugose. (Fig. 1,F-I).

Other specimens examined: Queensland. Cook District:
Daintree River, Dec 1929, Kajewski 1450 (BRI); ditto,
Aug 1985, Williams 85120 (BRI); Mossman, Aug 1957,
Volck 1388 (BRI); Little Falls Creek, Aug 1957, Smith
10050A (BRI); on road up to Mt Windsor Tableland,
16°17’S, 145°05’E, Jul 1978, Webb & Tracey 11409 (BRI);
Chowchilla Logging Area, State Forest Reserve 144, Mt
Windsor, 16°17’S, 145°05’E, Jul 1978, Unwin 608 (QRS);
State Forest Reserve 144, Oct 1975, Irvine 1632 (BRI).

Distribution and habitat: North Queensland;
in rainforest at altitudes of 960-1095 m
(Map 1).

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

45

Fig. 1 . Cyclophyllum coprosmoides var. coprosmoides . A. flowering branch x 0.6. B. flower x 5. C. LS of flower x 5.
D. fruit x 2. E. pyrenes x 3. A, Jessup s.n. (from Lebler 1978, p.529); B & C, Anderson 4100 (BRI); D & E, White 7287
(BRI). Cyclophyllum costatum. F. flower x 5. G. & H. fruits x 2. I. pyrenes x 3. F, Kajewski 1450 (BRI); G-I, Webb &
Tracey 11409 (BRI).

46

Austrobaileya 6 (1): 41-66 (2001)

Map 1 . Distribution of Cyclophyllum costatum ★and Cyclophyllum coprosmoides var. coprosmoides #.

Notes: Cyclophyllum costatum is readily
distinguishable by its long-attenuate somewhat
laterally compressed calyx tube, complanate
winged pedicels, usually strongly ribbed fruits,
and leaves + membranous when dry.

Conservation status: This species, as Canthium
costatum, is listed as vulnerable under the
Regulations of Queensland Nature
Conservation Act (as amended 2000) and the
Environment Protection and Biodiversity
Conservation Act 1999.

2. Cyclophyllum coprosmoides (F.Muell.)
S.T.Reynolds & R.J.F.Hend., comb, nov.;
Canthium coprosmoides F.Muell., Philos.
Inst. Vic. Trans. 3: 47 (1858). Type:
Dawson River, date unknown, Mueller s.n.
(syn: MEF [MEF1538527]); ditto,
December 1856, Mueller s.n. (syn: MEF
[MEF1538570, p.p.]); Mackenzie River,
date unknown, Mueller s.n. (syn: MEF
[MEF1538570, p.p.]); Brisbane River,
August 1855, Mueller s.n. (syn: MEF
[MEF1538569]).

[Plectronia barbata sensu F.Muell., Fragm.

9: 186 (1875), non (GForst.) F.Muell.]

Shrubs or trees 0.9-10.5 m high, with usually
opposite branching; bark grey mottled with pale
grey or white; branchlets usually slightly 4-
angular distally, pale greyish coloured or very
pale brown or brownish-grey, usually slightly
scurfy distally and with minute spreading white
hairs, usually with conspicuous minute red resin
granules and small, round, + pustular whitish
coloured lenticels. Leaves with stipules ovate
or triangular, keeled and tapered into a short or
long, folded apex, glabrous or sparsely white
hairy towards base adaxially; petioles 3-7mm
long, narrowly winged or ridged distally; blades
elliptic, elliptic-oblong, obovate or spathulate,
2.5-8(-12.5) x 1.6-4.6(-6.7) cm, with apex
obtuse, + rounded, slightly retuse or truncate,
subacute or rarely abruptly shortly acuminate
with obtuse acumen, and base + cuneate or
narrowly cuneate and decurrent into the petiole,
thin or thick coriaceous, with both upper and
lower surfaces usually with a slight sheen, the
upper one dark green or slightly yellowish-
green but drying dark brown to blackish and
usually dull above, and the lower one pale green

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

but drying greenish-yellow, grey or pale brown
with darker brown or white nerves and
occasionally blotched, concave along midrib on
adaxial surface; lateral nerves in 3-6 pairs,
slender, slightly oblique or arcuate, looping and
anatomising near margins; margins flat or
slightly recurved; domatia, if present, small.
Cymes (lor)2-4-flowered, sessile; bracts or
hairy or glabrous. Flowers in bud obtuse or
subacute at apex; pedicels (1.5—)3—5 mm long,
minute spreading hairy or glabrous; calyx 2-
2.5 x 1.75-1.5 mm, sparsely hairy or glabrous
except for ciliolate lobes; limb short, with lobes
minute, ovate; corolla white or cream,
becoming yellow with age, 6.5—8(—11) mm
long, with tube greenish yellow, 3-8 mm long,
c.2 mm wide at mouth, sparsely hairy adaxially
but densely hairy at throat; lobes ovate-elliptic,
± patent or slightly recurved, abruptly acute and
cucullate at apex, 3-5 x 1.5-2 mm, glabrous,
usually very scurfy adaxially, streaked with
whitish or reddish brown streaks, usually
densely papillose towards the apex adaxially
on the margins, sparsely papillose abaxially;
disc fleshy, as long as or shorter than calyx limb;
stamens with filaments c.0.5 mm long and
anthers ellipsoid, apiculate at apex, tailed at
base, 1.5-1.75 mm long; style with stigma 7-8
mm long; stigma c.1.5 mm x 1.75-2 mm long;
ovary walls (especially of mature ovaries)
usually covered with numerous reddish
resinous cells. Fruits on spreading to
pendulous pedicels 6-9 mm long, orange or
orange-yellow when ripe, ellipsoid to obovoid,
slightly obcordiform or transversely ellipsoid,
slightly lobed at apex, when dry shallowly
grooved between the lobes, 6-10 x 8-12 mm;
pyrenes broadly hemispherical, smooth or
slightly rugose.

Distribution and habitat: Eastern Queensland,
from Fitzroy Island, east of Cairns, to Brisbane
River; along creeks and river banks, on ridges
along rivers and on rocky headlands.

Notes: Cyclophyllum coprosmoides, as based
on specimens filed under the name Canthium
coprosmoides in various herbaria, was found
to be a very variable species within which seven
distinct taxa were distinguishable. Why this

47

very different material had been included in one
taxon in the past was probably because, with
the exception of the distinctive Canthium
costatum (now Cyclophyllum costatum) from
north Queensland and C. schultzii (now
Cyclophyllum schultzii ) from the Northern
Territory, Canthium coprosmoides was the only
name available for specimens of the Canthium
alliance with clustered umbelliform
inflorescences, and flowers with a corolla with
a long hypocrateriform tube and short lobes.
However, critical examination of the above
specimens led to the recognition of five new
species, namely Cyclophyllum longipetalum,
C. maritimum, C. multiflorum, C. protractum
and C. rostellatum, in addition to C.
coprosmoides which is now considered to
contain two varieties.

Some very early collections of this
alliance from northern Queensland were
previously identified as of Canthium barbatum
(or Plectronia barbata ), a Pacific species. That
species, however, differs from the above
Australian material by its very acuminate,
usually membranous leaves. Cyclophyllum
coprosmoides is distinguishable by its shortly
stalked, elliptic, elliptic-oblong or obovate
coriaceous leaf blades with obtuse, rounded or
subacute apex, usually hairy young branchlets,
sessile l-3(or 4)-flowered cymes, obtuse or
subacute flower buds, corollas 6.5—8(—11) mm
long, and obtuse or abruptly subacute corolla
lobes which are scurfy adaxially.

Affinities: Cyclophyllum coprosmoides is most
closely related to C. rostellatum in its more or
less si mil ar leaves, sessile cymes and number
of flowers in each cyme, but that species differs
from it by its longer petioles (8.0-10 mm long),
usually narrower leaf blades, rostrate flower
buds and acuminate corolla lobes. However,
specimens which appear to be intermediate
between these species are occasionally found
(see under C. rostellatum ).

Variability: The leaves of this species are very
variable but the material can be divided into
two subordinate taxa which are accepted here
as varieties following O. Schwarz (1927).

48

Austrobaileya 6 (1): 41-66 (2001)

Key to varieties of Cyclophyllum coprosmoides

1. Leaf blades elliptic, elliptic-oblong or subobovate, 5-9.3 x 2.7-5.2 cm,
rarely less, with apex obtuse, subacute or shortly obtusely acuminate, and
base obtuse or subacute; lateral nerves in 3-6 pairs; petioles 4-7 mm

long.2a. C. coprosmoides var. coprosmoides

Leaf blades obovate to spathulate, 2.5-4.8 x 1.6-2.3 cm, rarely more, with
apex obtuse or rounded, and base cuneate and decurrent into petiole; lateral
nerves in 3 or 4 pairs; petioles 3-4 mm long.2b. C. coprosmoides var. spathulatum

2a. C. coprosmoides var. coprosmoides Leaf
blades elliptic, elliptic-oblong or
subobovate, dull green or slightly
yellowish green, drying yellowish green
to pale greyish coloured on the abaxial
surface. (Fig. 1A).

Representative specimens: Queensland. North Kennedy
District: Proserpine River, 7 km ESE of Proserpine, Nov
1985, Sharpe 4202 (BRI). South Kennedy District: Cape
Hillsborough National Park, Hidden Valley road, 1.5 km S
of picnic area, Jan 1990, Thompson 163 (BRI). Leichhardt
District: Dawson Range, Blackdown Tableland,
23°44’S,149 o 07’E, Jun 1977, Telford 5778 (CANB); Isla
Gorge, about 28 km SW of Theodore, Aug 1973, Sharpe
620 & Hockings (BRI). Port Curtis District: Deepwater
National Park, 40 km E of Miriamvale, Jul 1989, Gibson
1609 (BRI); Raspberry Vale, Apr 1945, Blake 15547 &
Webb (BRI); Callide, Oct 1947, Smith 3585 (BRI);
Greenfields, about 14 km NE of Goovigen, Mar 1986,
Anderson 4115 (BRI). Wide Bay District: Black Gin Creek,
Timber Reserve 580,25°29’S,151° 55’E, Apr 1990, Forster
PIF6594 (BRI); Mt Walsh, 6.5 km S of Biggenden,
25°34’S,152°02’E, May 1977, Telford 5335 (CANB).
Moreton District: Base of Mt Coolum, Apr 1945, Clemens
s.n. (BRI); Brisbane, Gold Creek road, Brookfield, Jan
1984, Williams 84011 (BRI).

Distribution and habitat: Eastern Queensland,
from near Proserpine to Brisbane; on sandstone
ranges, ridges, stony hillsides and gullies; in
remnant scrubs on sandy stony soil, at altitudes
up to 450 m. (Map 1).

Variability: Leaf blades in this variety are very
variable in shape; specimens from the vicinity
of Brisbane usually have small, elliptic leaf
blades which are obtuse, rounded or
subacuminate at apex, and which dry olive-
green on the abaxial surface, whereas
specimens from ridges further north, near the
Dawson and Mackenzie Rivers as well as from
near Proserpine and Cape Hillsborough, usually
have larger, broadly elliptic or elliptic-oblong
to subobovate leaf blades which usually dry a
very pale grey on the abaxial surface. These
latter specimens also have flowers that are
slightly larger than those on specimens from

south-eastern Queensland.

Note: Specimens with more or less obovate leaf
blades can be confused with those of
C. coprosmoides var. spathulatum, but the latter
variety differs from the former in its smaller
leaf blades or by having a fewer number of nerves
in those blades.

2b. C. coprosmoides var. spathulatum

(O.Schwarz) S.T.Reynolds & R.J.F.Hend.,
comb.nov.; Plectronia coprosmoides var.
spathulata O.Schwarz, Feddes Repert. 24:
102 (1927). Type: East Coast, Brown s.n.
(lecto [here designated]: BRI; isolecto: CANB,
MEF, NSW).

Leaf blades obovate or spathulate, rounded,
obtuse or occasionally retuse at the apex and
cuneate at the base, of varying shades of green
to grey but paler on the abaxial surface than on
the adaxial surface.

Specimens examined: Queensland. Cook District: Fitzroy
Island, 16°55S, 146°03E, Jan 1997, Jago 4233 (BRI). North
Kennedy District: Harold Island, 20 o 14’S,149°09’E, Nov
1983, Bat ianoff3404 & Dillewaard (BRI). South Kennedy
District: Port Mackay, date unknown, Dietrich 1042
(MEL); Bailey Islet, May 1969, Firth s.n. (BRI). Port Curtis
District: Gladstone, Dec 1864, Dietrich 1229 (MEL); ditto,
near Keppel Bay, Jan 1865, Dietrich 2388 (MEL); Sloping
Island, S of N Keppel Island, Oct 1987, Batianojf 9197 &
Dillewaard (BRI). Wide Bay District: Fraser Island, May
1967, Baxter 797 (BRI); ditto, Sep 1979, Coutts 3 (BRI).

Distribution and habitat: Eastern coastal
Queensland, commonly on offshore islands;
usually along beaches, on rocky headlands and
on frontal dunes. (Map 2).

Notes: Leaf blades in specimens from offshore
islands, especially in those collected from plants
growing in deep sand on high dunes (e.g.
Batianoff 9197 & Dillewaard ) are
comparatively very small, and are clustered on
usually stunted branchlets with the nodes very
close together. Leaf blades in those from plants

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

49

Map 2. Distribution of Cyclophyllum rostellatum ★, Cyclophyllum protractum A and Cyclophyllum coprosmoides var.
spathulatum #.

growing on the mainland, e.g. from near
Gladstone, are usually larger and resemble
those in some specimens of the typical variety.
However, that variety has leaf blades with a
greater number of lateral nerves than are present
in the leaf blades of this variety.

Typification: O. Schwarz (1927) cited six
collections when describing Plectronia
coprosmoides var. spathulata, namely [1] East
Co [a] st, R.Br[own], [2] Gladstone, Dietrich
no. 1229, [3] [locality unknown], Banks et
Solander s.n., [4] Rockingham Bay, [collector
unknown], [5] Queensland [without precise
locality], F. Mueller [s.n.] and [6] Richmond
River, [collector unknown]. These syntypes
were all apparently housed in herbarium B in
Berlin and are probably now no longer in
existence. As to isosyntype material, only
Brown s.n. (BRI, CANB, MEL, NSW) and
Dietrich 1229 (MEL [MEL1538514]) has been
available for study here. The former collection
agrees with the protologue and matches
specimens included in this variety here, whereas
Dietrich 1229 has larger leaf blades which
approach in size those on specimens included

in Cyclophyllum coprosmoides var.
coprosmoides here. Therefore, based on the
material that was available for this study,
Brown’s duplicate specimen at BRI, stated only
to have been collected on the “East Coast” of
Australia, is chosen here as lectotype of
Schwarz’s varietal name. Brown’s “East Coast”
locality is almost certainly in coastal eastern
Queensland.

The locality of collection of the Banks
and Solander specimen cited by Schwarz was
probably the Endeavour River in north
Queensland. Banks and Solander specimens
in BRI and CANB of the related C. rostellatum
are definitely stated to have been collected from
the Endeavour River and are possibly/probably
conspecific with the specimen seen by Schwarz.

3. Cyclophyllum rostellatum S.T.Reynolds
& R.J.F.Hend. sp. nov.; C. coprosmoidi
(F.Muell.) S.T.Reynolds & R.J.F.Hend.
similis sed foliis petiolis longioribus,
floribus in statu alabastro subrostratis ad
apicem, corollae lobis plerumque
acuminatis vel caudatis et fructibus

50

obcordiformibus differt. Typus:
Queensland. Cook District: 9 km N of
Batavia Downs on the Peninsula
Development Road, 12 o 35’S,142°10’E,
altitude 100 m, 22 April 1990,
J.R. Clarkson 8505 & V.J. Neldner{ holo:
BRI). Canthium sp. (Mt Rose A.R.Bean
1978), S.T. Reynolds (1997, p.180).

Small trees 2-6 m high; trunks 12-15 cm in
diameter; bark grey mottled with white,
tessellated; branchlets quadrangular distally,
whitish or yellowish-white coloured, densely
lenticellate, glabrous. Leaves with stipules
triangular, keeled and attenuated into a long
folded apex, thin and often paleaceous and ±
fimbriate at margins, and with long rust-
coloured hairs on the margins and at base
adaxially; petioles 8-10 mm long; blades
elliptic-oblong to subobovate, (3.7—)6.5—11(—
13.5) x (1.6-)2.7-3.4(-5.5) cm, with apex
obtuse or + rounded, and base cuneate and
attenuate into the petiole, glabrous, thick
coriaceous, the adaxial surfaces drying brown,
the abaxial ones drying olive green, greyish-
brown, pale grey or whitish coloured (in young
leaves) and often with whitish coloured flecks
and blotches; midrib slightly channelled
adaxially; lateral nerves in 4 or 5 pairs, slender,
± arcuate and looping near margins; reticulate
venation obscure or absent; domatia very rarely
present when inconspicuous. Cymes (l-)3-
6-flowered, with flowers fasciculate on a short,
thick, knob-like peduncle; bracts rust-coloured
hairy. Flowers in bud acuminate and somewhat
rostrate at apex, rarely obtuse or subacute,
strongly perfumed; pedicels 2-3.5 mm long;
calyx glabrous, c.2 x 1.5 mm, with short,
denticulate limb with ciliolate lobes; corolla
creamy yellow to orange or brown, 10-12 mm
long (only 5 mm long in one specimen), with
tube (2-) 4.5-7 mm long, 1-1.5 mm wide at
mouth, sparsely hairy adaxially but densely
hairy at mouth; lobes elliptic, abruptly long
acuminate at apex with acumen usually about
half the length of the lamina of corolla lobe, or
subacute at apex, 2.5-4.5 x 1-2 mm, glabrous,
scurfy adaxially, densely papillose along
margins and at attenuated slightly folded apex;
stamens with filaments c.0.5 mm long and
anthers c.1.5 mm x 1 mm; style with stigma
7.5-8.5 mm long; stigma c.0.75 x 0.75 mm.
Fruits on erect, spreading to pendulous

Austrobaileya 6 (1): 41-66 (2001)

pedicels (3-)6-8 mm long, orange-yellow,
orange-brown or reddish coloured when ripe,
obcordiform, slightly lobed at apex,
prominently longitudinally 2-lobed, when dry
with deep and narrow grooves between the
lobes and with short ribs towards the base, 6-
11 x 5-11 mm, or rarely 1-lobed; pyrenes
depressed ovoid. (Fig. 2).

Selected specimens: Queensland. Cook District: Edith
Falls, Jardine River, 11°09’S,142°30’E, Oct 1989, O’Reilly
541 (BRI); Brown’s Creek, Pascoe River, Jul 1948, Brass
19583 (BRI); north bank of Pascoe River near Youngman’s
Crossing, 12°33’S,143°14’E, Nov 1977, Tracey 14597
(BRI); MtTozer, Iron Range, 12°45’S,143 0 13’E, Oct 1968,
Webb & Tracey 8719 (BRI); Tozer [Iron] Range, 0.5 mile
[0.8 km] E of Mt Tozer, Jul 1948, Brass 19401 (BRI,
CANB); Mcllwraith Range, Leo Creek Road, Timber
Reserve 14, 13°43’S,143°20’E, Sep 1975, Hyland 3330
(BRI, CANB, NSW, QRS); ditto, Sep 1975, Sanderson 748
(QRS); Mcllwraith Range, Lankelly Creek, Timber Reserve
9, Jun 1992, Forster PIF10382 & Tucker (BRI); Mt Rose
area, NW of Cooktown, 15°20’S,145°02’E, Jul 1990, Bean
1978 (BRI); Ayton Road, Jun 1962, Gittins 566 (BRI,
NSW).

Distribution and habitat: Far northern
Queensland from Torres Strait islands to near
Cooktown; on ranges, rocky slopes, gorges and
gullies, usually near permanent streams; in
rainforests, mostly on soils derived from granite
at altitudes of 250-600 m (Map 2).

Notes: The inclusion of Torres Strait Islands in
the distributional range of C. rostellatum is
tentative because only scanty, sterile material
presumably of this species from this locality
has been seen.

The colour of its fresh leaves is recorded
as bluish green by F.S. Smith {Smith 11834,
BRI).

C. rostellatum is characterised by its
subrostrate flower buds, acuminate or subacute
corolla lobes, whitish coloured glabrous
branchlets, usually narrow elliptic leaf blades
on long petioles and its rusty hairy stipules and
bracts. It is closely related to C. coprosmoides
which it resembles at first sight in its leaves
and few-flowered cymes but that species differs
from C. rostellatum by its obtuse flower buds
and corolla lobes, usually broader leaves, short
petioles and hairy and scurfy young branchlets.
These species, however, are connected by
intergrades (see below) but the extremes are
quite distinct.

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

51

Fig. 2. Cyclophyllum rostellatum. A. flowering branch x 0.8. B. detail of inflorescence x 2. C. flower x 5. D. LS of
flower x 5. E. stamen (adaxial view) x 10. F. stamen (abaxial veiw) x 10. G. fruit x 3. H. pyrene x 3. A-H, Clarkson 8505
& Neldner (BRI). Del. W. Smith.

52

Variability: The shape and size of leaf blades
and the shape of the flower buds in this species
are quite variable. Specimens with narrowly
elliptic or subobovate leaf blades which dry
greyish or greyish-brown on the abaxial surface
and have dark coloured nerves and paler
coloured blotches, subrostrate flower buds and
acuminate corolla lobes are typical of this
species. Whereas specimens with very small
thick leaves (e.g. Hyland 3330 and Sanderson
748 from Mcllwraith Range, and Brass 19401A
from Mt Tozer Range above) probably
represent a distinct form or variant, they lack
flowers. Examination of these, would be
necessary to be certain of this. Moreover,
specimens with broad leaves with acuminate
flower buds (e.g. Bean 1978 and Gittins 566
above), and others with narrow leaves but with
obtuse to subacute flower buds are also
included here. These probably represent
intergrades between this species and
C. coprosmoides, but examination/study of
more such specimens would be necessary to
be certain of this. The broad-leaved form, as
represented by Bean 1978 and Gittins 566
above, resembles the latter species but differs
from that in the nature of its flower buds, overall
aspect, longer petioles and glabrous branchlets.

Etymology: The specific epithet, from Latin
rostellatus, beak-like, refers to the beak-like
apex of the flower buds.

4. Cyclophyllum longipetalum S.T.Reynolds
& R.J.F.Hend. sp. nov. differt a
C. coprosmoide (F.Muell.) S.T.Reynolds
& R.J.F.Hend. foliis conspicue foveolatis
acuminatis subacutisve et petiolis
corollisque longioribus. Typus: [New
South Wales.] Port Jackson, February
1805, R. Brown s.n. (holo: MEL
[MEL1538081]; iso: ?BM n.v., BRI
[AQ123681] CANB,NSW [NSW193831]).

Canthium sp. (Cooroy S.T.Blake+ 15507),
S.T. Reynolds (1997, p.180).

Trees 2.5-20 m high; trunks to 38 cm in
diameter; branchlets glabrous, quadrangular
distally, very pale brown often mottled with
white, dotted with minute resin glands and small
whitish coloured lenticels. Leaves with stipules
triangular, keeled, attenuated into a long folded
apex; petioles 5-16 mm long; blades elliptic,

Austrobaileya 6 (1): 41-66 (2001)

(5.5-)7-9.5 (rarely -11.5) x (2.2-)3.3-4.3(-5.2,
rarely -5.7) cm, usually abruptly narrowing and
acuminate at both ends, or with apex abruptly
acuminate or subacute, and base acute and
decurrent into the petiole, glabrous, thin,
coriaceous, with adaxial surface pale green and
dull or slightly glossy but drying dull green,
pale greenish-yellow or dark brown and usually
blotched, and abaxial surface usually paler than
adaxial one; lateral nerves in 4 or 5 pairs,
slightly arcuate and looping near margins,
obscure adaxially; domatia present, prominent,
few, on each side of midrib. Cymes 1-4-
flowered, sessile or subsessile with peduncles
1-2 mm long; bracts hairy. Flowers in bud
obtuse at apex, strongly scented; pedicels 4-6
mm long, usually stout; calyx 2.5-3 mm long,
with tube + cupular but attenuate into the
pedicel and with a short lobed limb; lobes
minute, ovate, usually ciliolate; corolla 12-15
mm long, with tube white becoming brown with
age, 8-12 mm long and c.2 mm wide, sparsely
hairy adaxially and densely hairy at mouth;
lobes white becoming greenish orange with age,
ovate-elliptic, 3.5-5 x 1.5-2 mm, folded at apex
where + abruptly shortly acuminate or
cuspidate, scurfy adaxially, glabrous abaxially
and densely papillose on the margins; disc
slightly shorter than the calyx limb, fleshy;
anthers ovoid-ellipsoid, c.2 mm long, apiculate
distally, tailed proximally, on filaments c.0.5
mm long; style with stigma 10-13 mm long;
stigma 2-2.5 x 2-2.5 mm. Fruits on erect,
spreading to pendulous pedicels 7.5-12 mm
long, orange to red when ripe, obovoid,
depressed or shallowly lobed at apex, deeply
grooved between lobes when dry, 1-1.25 x 1.2-
1.5 cm; pyrenes depressed ovoid. (Fig. 3).

Selected specimens: Queensland. Wide Bay District:
Fraser Island, Nov 1915, collector unknown [? Morrison
s.n.] (BRI); ditto. May 1967, Baxter 845 (BRI); Cooroy,
Apr 1945, Blake 15507 & Webb (BRI). Moreton District:
Rocky Creek, State Forest 249, SE of Yandina,
26 0 35’S,152 0 59’E, Apr 1990, Bean 1503 (BRI); Mt
Glorious, Jan 1945, Clemens s.n. (BRI); ditto, Apr 1956,
Hoogland 5237 (CANB); ditto, Apr 1999, Phillips 199
(BRI); Lyrebird Ridge road, northern end of Springbrook
Plateau, 28 0 irS,153°15’E, Dec 1993, Grimshaw 269
(BRI). New South Wales. Lennox Head, Ballina, Apr 1892,
Bauerlen 799 (MEL, NSW); Lismore, Oct 1891, Bauerlen
s.n. (MEL); Whian Whain State Forest, Gibberagunyah
Mountain, 28°35 , S,153°19’E, May 1968, Jones 3825
(CANB); Port Macquarie, Feb 1895, Brown s.n. (NSW
[NSW193746]); Hastings River, date unknown, Beckler
s.n.(MEL [MEL1538257]; NSW [NSW193747]);

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

53

Fig. 3. Cyclophyllum longipetalum. A. flowering branch x 0.6. B. detail of inflorescence x 2. C. flower x 5. D. fruit x
2. E. LS of fruit showing embryo x 2. F. pyrene x 2. A-C, Brown s.n. [MEL1538081] (MEL); D-F, Peberdy s.n.
[AQ468261] (BRI). Del. W. Smith.

Kurrajong, Blue Mountains, date unknown, Woolls s.n.
(MEL [MEL1538498]); Kalandra Sanctuary, Mona Vale,
33°44’S,151°17’E, Oct 1977, Coveny 9697 (NSW);Foley’s
Bush, Foxground, 34°43’S,150°46’E, Nov 1906, Hay s.n.
(NSW).

Distribution and habitat: South-eastern
Queensland to Illawarra District, New South
Wales; usually on steep ranges, ridges and
hillsides, in shallow stony soil; in rainforests at
altitudes of 135-600 m (Map 3).

Notes: C. longipetalum is characterised by its
few-flowered cymes, flowers with long corolla
tubes and foveolate, usually subacuminate
leaves with long petioles. It resembles
C. coprosmoides at first sight in its leaves and
inflorescences, but differs from that species as
follows:-

C. longipetalum has glabrous branchlets,
subacuminate or acute leaves with petioles 5-
16 mm long and domatia present and
prominent, sessile or subsessile cymes with
peduncles 1-2 mm long, and flowers with a
corolla 12-15 mm long. C. coprosmoides has
branchlets usually scurfy and hairy distally,
obtuse or slightly round-tipped leaves with
petioles 3-7 mm long and domatia, if present,
small, sessile cymes and flowers with a corolla
6.5—8(—11) mm long.

Etymology: The specific epithet, from Latin
longus, long, and petalum, petal, refers to the
comparatively long corollas in flowers of this
species.

54

Austrobaileya 6 (1): 41-66 (2001)

115 120 125 130 135 140 145 150 155

Map 3. Distribution of Cyclophyllum longipetalum A,Cyclophyllum multiflorumn and Cyclophyllum schultzii forma
schultzii #-

5. Cyclophyllum protractum S.T.Reynolds &
R.J.F.Hend. sp. nov.; C. coprosmoidi
(F.Muell.)S.T.Reynolds & R.J.F.Hend.
similis sed foliis foveolatis apice
acuminato protractove et inflorescentiis
2-5-floribis differt. Typus: Queensland.
Cook District: State Forest Reserve 144
Whypalla, Chowchilla Logging Area,
16°18’S,145°05’E, altitude 1000 m, 2
February 1988, B.P.M. Hyland 13500
(holo: BRI; iso: ?QRS n.v.).

Canthium sp. (Copper-Lode Falls C.H.Gittins
2211), S.T. Reynolds (1997, p.180).

Trees 8-15 m high; trunks 10-20 cm in
diameter; branchlets glabrous, greyish coloured
mottled with white and dotted with whitish or
dark coloured, + pustulate lenticels. Leaves
with stipules broadly ovate, keeled, attenuated
into a short lobe distally; petioles 3-8 mm long;
blades elliptic-ovate, 5.6-7.7(-8.3) x 2.7-3.2(-
4) cm, with apex usually abruptly long
acuminate or ± caudate, and base subacute,

glabrous, thin, coriaceous, drying dark brown
adaxially, pale brown abaxially and sometimes
irregularly blotched; lateral nerves in 3-5 pairs,
± arcuate and looping near margins; secondary
veins loosely reticulate, usually obscure;
domatia present, usually conspicuous. Cymes
2-5-flowered, on peduncles 0.5-1.5 mm long;
bracts glabrous, smooth or ciliolate. Flowers
in bud obtuse at apex, strongly perfumed;
pedicels 2-3 mm long; calyx 1.5-2 x c.1.75
mm, cupular, glabrous, with a short, lobed limb;
lobes minute, ovate, glabrous or rarely sparsely
ciliolate; corolla 8.5-9 mm long, cream, yellow
or orange with age, with tube 4.5-6 mm long
and c.1.5 mm wide at mouth, sparsely hairy
adaxially but densely hairy at mouth; lobes
elliptic-ovate, 3.5—4.5 x c. 1.5 mm, glabrous but
densely papillose at the slightly cucullate apex;
stamens with filaments c.0.5 mm long and
anthers c. 1.5 mm long; style with stigma 6-
7.5 mm long; stigma c. 1.25 x 1.25 mm. Fruits
on erect to spreading or decurving pedicels 8-
10 mm long, red when ripe, transversely
ellipsoid or ± obcordiform, slightly truncate to

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

lobed at apex, deeply grooved between lobes
when dry, 7.5-9 x 9-11 mm; pyrenes smooth.

Selected specimens: Queensland. Cook District: State
Forest Reserve 144, Agapetes Logging Area,
16°17’S,145°05’E, Dec 1979, Hyland 10178 (BRI, QRS);
State Forest Reserve 143, Little Mossman Logging Area,
16°32’S,145 0 23’E, nearMossman, Oct 1978, Moriaty 2470
(QRS); State Forest Reserve 607, Shoteel Logging Area,
Mickies Pocket, 16°55’S,145 0 36’E, [NE of Mareeba], Dec
1981, Gray 2318 (BRI, QRS); Copper-lode Falls Dam site,
c.6 miles [c.9.5 km] S of Cairns, 16°56 , S,145°46’E, Sep
1970, Gittins 2211 (BRI).

Distribution and habitat: North Queensland,
from Daintree River to near Cairns and
Mareeba; on ranges in rainforest at altitudes of
400-1000 m (Map 2).

Notes: C. protractum is characterised by its
usually abruptly acuminate, prominently
foveolate leaves, 2-5-flowered shortly
pedunculate cymes and transversely ellipsoid
or subobcordiform fruits. It resembles
C. maritimum and C. coprosmoides in the shape
and size of its leaves and few-flowered cymes,
but the former species differs from
C. protractum by its efoveolate leaves,
peduncles (l-)2.5-6 mm long, (2-)4-7-
flowered cymes, and transversely ellipsoid
fruits which are slightly depressed at the apex
and only shallowly lobed when dry.
C. coprosmoides differs from C. protractum by
having leaves with domatia, when present,
comparatively small, and the abaxial surface
of the blade is greenish yellow, greyish or pale
brown coloured when dry, and its sessile, 1-4-
flowered cymes.

Etymology: The specific epithet, from Latin
protractus, drawn out, refers to the usually
protracted apex of this species’ leaf blades.

6. Cyclophyllum maritimum S.T.Reynolds
& R.J.F.Hend. sp. nov. primo aspectu
C. protracto S.T.Reynolds & R.J.F.Hend.
persimilis, sed foliis efoveolatis apice
obtuso et inflorescentiis distincte
pedunculatis differt. Typus: Queensland.
Cook District: Base of Mt Cook, near
Cooktown, January 1982, V.Scarth-
Johnson 1218A (holo: BRI).

Canthium sp. (Lizard Island R.L.Specht+
LI181), S.T. Reynolds (1997, p.180).

55

Shrubs or small trees 3-10 m high; bark mottled
grey, slightly rough; branchlets glabrous, very
pale brown or pale grey, with dense, whitish or
dark coloured lenticels. Leaves with stipules
triangular, keeled, attenuated into a short lobe
distally; petioles 5-8 mm long, channelled
adaxially; blades elliptic, broadly elliptic or
elliptic-ovate, (3.8-) 5.5-8.2 x (2.1-)3^L6(-5.2)
cm, with apex obtuse or abruptly shortly
acuminate, and base abruptly obtuse or
subacute and decurrent into petiole, with
margins slightly recurved, thin, coriaceous, the
adaxial surface slightly glossy, drying brown,
blackish or dark greyish-green and with midrib
occasionally sunken proximally, the abaxial
surface usually paler brown with nerves darker
coloured; lateral nerves slender, in 4-6 pairs,
suboblique or arcuate and looping near margins;
secondary venation usually obscure; domatia
absent or very rarely present but obscure.
Cymes (2-)4-7-flowered, on peduncles (1—)2.5—
6 mm long; bracts glabrous. Flowers in bud
subacute or obtuse at apex, strongly perfumed;
calyx c.2 x 1.5 mm, glabrous, with a short,
denticulate limb; corolla 9-10 mm long, yellow,
with tube 5-6 mm long and 1.5-2 mm wide at
mouth, sparsely hairy adaxially but densely
hairy at mouth; lobes sublanceolate, 3.5-5.5 x
1.5-1.75 mm, scurfy adaxially, glabrous abaxially,
densely papillose distally on the acute and
slightly reflexed apex; stamens with filaments
c.0.5 mm long and anthers c.1.5 x 1 mm; style
with stigma 7-8 mm long; stigma c.1.25 x 1.25
mm. Fruits on erect, spreading or decurving
pedicels 6-10 mm long, greenish orange to red
when ripe but drying black, transversely
ellipsoid, obovoid or obliquely ellipsoid,
slightly depressed at apex, with abroad shallow
channel between the lobes when dry, 8.5-11.5
x 8.5-14 mm; pyrenes smooth. (Fig. 4).

Selected specimens: Queensland. Cook District: Torres
Strait, Duan Island, Sep 1971, Lawrie s.n. (BRI); ditto,
Yorke Island, 9°45’S,143 0 25’E, Jan 1971, Oct 1911, Lawrie
s.n. (BRI); ditto, Jun 1995, Waterhouse BMW3640 (BRI);
Bamaga District, Jacky Jacky Creek, May 1962, Webb &
Tracey 5997 (BRI); 9.5 km S of Captain Billy Landing,
H 0 38’S,142°5rE, Mar 1992, Clarkson 9257 & Neldner
(BRI); Shelburne Holdings, track between Round Point and
Conical Hill, Nov 1985, Gunness AG1929 (BRI); Lizard
Island, Jun 1973, Specht LI112 (BRI); Finch Bay near
Cooktown, Mar 1966, Smith 13103 (BRI), ditto, Apr 1973,
Henderson\\\6()A (BRI); Cooktown, Jul 1943, Blake 15066
(BRI); Endeavour River, in 1882, Persieh 761 (BRI,

56

Austrobaileya 6 (1): 41-66 (2001)

Fig. 4. Cyclophyllum maritimum. A. flowering branch x 0.8. B. detail of inflorescence x 2. C. flowers x 5. D. LS of
flower x 5. E. fruit x 2. F. pyrene x 3. A & B, Gunness AG1929 (BRI); C & D, Clarkson 8636 & Neldner (BRI); E & F,
Clarkson 7699 & Neldner (BRI). Del. W. Smith.

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

57

Map 4. Distribution of Cyclophyllum maritimum.

MEL); Walker Bay, 15 0 31’S,144 0 16’E, Dec 1988,
Clarkson 7699 & Neldner (BRI).

Distribution and habitat: North-eastern
Queensland, from Torres Strait Islands to the
Daintree River, common around Cooktown; in
coastal scrubs on headlands, frontal sand dunes,
in gullies, along tidal creeks and rivers; in sandy
and sandy rocky soil (Map 4).

Affinities: Cyclophyllum maritimum is
characterised by its thick, slightly glossy,
efoveolate leaf blades, its pedunculate, (2-) 4-
7-flowered, umbelliform cymes, and its large,
broadly ellipsoid fruits which are shallowly
channelled between the lobes and depressed at
the apex. It resembles C. protractum at first
sight in its leaves, but that species differs from
C. maritimum in having leaf blades with
prominent domatia and mostly abruptly long-
acuminate at the apex, subsessile or sessile
cymes, and fruits deeply grooved when dry.

Variability: The leaves of C. maritimum are
variable. Specimens from around Cooktown
which have elliptic leaf blades which are obtuse
at both ends are typical of this species but

specimens from Lizard Island usually have
thicker, narrower, often comparatively smaller
leaf blades which are much narrower at their
base. These latter specimens possibly represent
a distinct form or variant but the material
presently available is insufficient to determine
if this is correct.

Notes: Collections from near Captain Billy
Landing in Cape York Peninsula which have
comparatively larger and broader leaf blades
are only tentatively included here. Their leaves
approach some of those in specimens included
under C. coprosmoides here but in most other
characters they appear to be of this species.
More material for examination is needed to be
certain of their affinities.

Collector Margaret Lawrie recorded that
on Yorke Island in the Torres Strait ( Lawrie
[AQ3960, AQ123642], BRI), this species is
known by the native name “Uru”, and that the
fruits are inedible [for humans].

Etymology: The specific epithet, from Latin
maritimus, growing by the sea, refers to the
usual habitat of this species.

58

7. Cyclophyllum multiflorum S.T.Reynolds
& R.J.F.Hend. sp. nov. quoad
inflorescentiam C. schultzii (O.Schwarz)
S.T.Reynolds & R.J.F.Hend. accedens sed
inflorescentiis sessilibus, corollis
longioribus fructibus obovoideis differt.
Typus: Queensland. North Kennedy
District: Edmund Kennedy National
Park near Cardwell, 18°13’S, 146°00’E,
16 December 1991, A.R. Bean 3869
(holo: BRI).

Canthium sp. (Kuranda G.Sankowsky+
680), S.T. Reynolds (1997, p.180).

Small trees 4-10 m high; bark light grey,
smooth; branchlets whitish coloured or very
pale brown, usually resin-gland dotted and very
resinous distally, conspicuously densely
lenticellate with usually small white lenticels,
puberulous throughout or glabrous distally.
Leaves with stipules triangular, keeled,
cuspidate with a long folded apex (that on
stipules distal on branchlets even longer),
sparsely hairy adaxially at base; petioles (2.5-
) 4-8 mm long, channelled adaxially; blades
elliptic, elliptic-oblong or + oblanceolate, (6.7)
9-12 (-13) x (2.9-)4.3-5.5(-6) cm, usually
abruptly obtuse and attenuate at both ends, or
with apex acuminate or obtuse and base acute
or subacute and attenuate into the petiole,
glabrous, thin or thick, coriaceous, drying dark
or reddish brown adaxially, olive green, yellow-
green or pale greyish coloured (especially in
young leaves) and usually speckled and
blotched with white abaxially; midrib slightly
channelled towards the base adaxially; lateral
nerves in 5 or 6 pairs, slightly arcuate or
obliquely arched, looping near margins, drying
reddish-brown in young leaves; secondary
venation very openly reticulate; domatia
present, few on each side of the midrib,
conspicuous. Cymes (2-)6-9(-14)-flowered,
sessile or subsessile; bracts hairy. Flowers in
bud obtuse at apex; pedicels (l-)2.5-6 mm
long, puberulous or glabrous; calyx c.2.5 x 2
mm, glabrous; limb short, denticulate; corolla
cream coloured or pale yellow, orange or brown
with age, (9—) 11.5—13.5 mm long, with tube
(7-)8-10 mm long, 1-1.5 mm wide at mouth,
sparsely hairy adaxially but densely hairy at
mouth; lobes ovate, abruptly acute and ±
cucullate at apex, (2-) 3-4.5 x 1.5-2 mm,

Austrobaileya 6(1): 41-66 (2001)

glabrous, scurfy adaxially, papillose on margins
and more densely at apex; disc shorter than the
calyx limb; stamens with filaments c.0.5 mm
long and anthers c.1.5 mm long; style with
stigma 10-11 mm long; stigma c.0.75 x 0.75
mm. Fruits on erect to spreading, slender
pedicels (6-) 8-15 mm long, orange-red when
ripe, obovoid or subellipsoid when slightly
wider above the middle, laterally compressed,
truncate at apex, when dried slightly
rhomboidal and deeply grooved, or with
shallow broad channels between the two lobes,
8-12 x 6-11 mm; pyrenes smooth or slightly
rugose. (Fig. 5).

Representative specimens: Queensland. Cook District:
Daintree River, Dec 1929, Kajewski 1465 (MEL); ditto,
Oct 1968, Webb & Tracey 11371 (BRI); Mowbray River,
Jan 1932, Brass 1955 (BRI); Noah Creek,
16 o 08’S,145°27’E, Dec 1986, Sankowsky 571 &Sankowsky
(BRI); State Forest Reserve 1073, Buchan Logging Area
near Kuranda, 16°46’S,145 0 37’E, Jan 1979, Gray 1256
(BRI); Bridle Creek about 12 miles [19.2 km] SE of
Mareeba, Nov 1973, Hartley 14123 & Hyland (BRI,
CANB); Russell River, in 1892, Johnson s.n. (MEL);
Bellenden Ker Range, Oct 1977, Jago 19 (QRS); Mt Bartle
Frere, in 1882, Johnson [MEL1538185] (MEL); ditto, Oct
1935, Blake 9803 (BRI); State Forest Reserve 194, East
Barron, 17 0 21’S,143°27"E, Nov 1981, Gray 2241 (BRI);
Fenby’s Gap, 17°52 , S,146°05 , E, Oct 1951, Smith 4901
(BRI). North Kennedy District: Murray River, Oct 1867,
Dallachy [MEL1538502] (MEL); Murray Upper,
18°05 , S,145°42 , E, Feb 1991, Cooke 464 (BRI); Wigham
Creek Crossing, 37 km NW of Ingham on road to
Broadwater Creek State Forest Park, 18 0 27’S,145°59’E,
Nov 1992, Halford 689 (BRI); Fa mil y Islands, Sep 1864,
Dallachy [MEL1538260] (MEL); Edmund Kennedy
National Park near Cardwell, 18 o 13’S,146 o 06’E,Dec 1991,
Bean 3869 (BRI); approx. 6.5 kmNNW of Cardwell, Oct
1978, Thorsbome & Thorsbome 300 (BRI); Mt Fox, Apr
1949, Clemens s.n. (BRI); ditto, Dec 1949, Clemens s.n.
(BRI).

Distribution and habitat: North-eastern
Queensland, from the Daintree River to near
Cardwell; usually along creeks, near swampy
lowland forests, on banks of brackish lagoons,
beach ridges and coastal sands; in vine thickets
on sandy soil (Map 3).

Affinities: Cyclophyllum multiflorum is readily
distinguishable by its 5-9-flowered, sessile
cymes, usually hairy branchlets and pedicels,
usually densely lenticellate branchlets,
prominently foveolate leaf blades with arcuate
lateral nerves and greenish-yellow or olive
coloured discoloration of the abaxial surfaces
of dried leaf blades. It resembles C. schultzii in

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

59

Fig. 5. Cyclophyllum multiflorum. A. flowering branch x 0.8. B. detail of inflorescence x 0.8. C. flower x 5. D. LS.
of flower x 5. E. detail of anthers and stigma (viewed from above) x 10. F. fruit x 3. G. pyrene x 3. A-E, Bean 3869
(BRI); F & Q Sankowsky 684 (BRI). Del. W. Smith.

60

its many-flowered inflorescences, but that
differs from this species by its distinct, single
or occasionally branched pedunculate
inflorescences, its flowers with shorter corollas
with tubes less than twice as long as the lobes,
and its usually subcordiform fruits which are
usually deeply and widely 2-lobed apically. In
addition, C. multiflorum resembles Canthium
longiflorum (Valeton) Merr. & L.M.Perry from
New Guinea, a species also belonging in
Cyclophyllum, in its sessile, many-flowered
inflorescences, but that species differs from the
former in its longer and larger leaf blades, and
longer pedicels.

Variability: The shape, size and texture of leaf
blades and the presence or absence of hairs on
branchlets and inflorescence axes are very
variable in this species. The majority of
specimens examined have hairy branchlets and
peduncles whereas two collections (Jago 19 and
Gray 2241 above) differ in having these parts
glabrous. Specimens from near Cardwell, the
Atherton Tableland and the Daintree River, with
elliptic, sublanceolate or elliptic-oblong, thick
leaf blades with a subacute or acuminate apex
are typical of this species. Collections from near
the Russell River, Mt Bartle Frere and Mt
Bellenden Ker usually have comparatively
smaller, thinner leaf blades with subacute, ±
obtuse or shortly acuminate apex. They
probably represent a distinct form of this
species, but more specimens especially ones in
flower are necessary to be certain of this.

Etymology: The specific epithet, from Latin
multiflorus, abounding in flowers, refers to the
many-flowered inflorescences in this species.

8. Cyclophyllum schultzii

(O.Schwarz)S.T.Reynolds & R J.F.Hend.
comb, nov.; Plectronia schultzii
O.Schwarz, Feddes Repert. 24: 101
(1927); Type: [Northern Territory.] Port
Darwin, 16 mil es E [of], banks of Howard
Creek, Bleeser 81 (?Bf, n.v.). Canthium
schultzii (O.Schwarz) Chippendale, Proc.
Linn. Soc. NSW 96(4): 208 (1972).

Shrubs or trees 1.5-10 m high; bark mottled
with grey or brownish-grey colouration, smooth
or finely fissured; branchlets very pale to dark
brown, glabrous, usually dotted with minute
pale brown lenticels. Leaves with stipules

Austrobaileya 6(1): 41-66 (2001)

triangular, keeled, attenuated into a long or short,
narrow folded apex; petioles 5-15 mm long;
blades broadly or narrowly elliptic, elliptic-
ovate, elliptic-oblong to ± oblanceolate, 8.5-
14(-16.2) x 1.8-7.2(-8.2) cm, with apex abruptly
and shortly, bluntly acuminate, subacute or
obtuse, and base abruptly obtuse or subacute,
and usually attenuate into the petiole, thin or
thick, coriaceous, glabrous, with adaxial
surfaces glossy or dull green and drying
brownish or blackish coloured and sometimes
with white blotches, and abaxial surfaces pale
green or slightly glaucous and drying pale
brown to pale greyish brown, sometimes with
midrib and nerves darker coloured; lateral
nerves in (4—)6—11 pairs, obliquely arched or
arcuate and looping near margins, prominent;
secondary venation very openly reticulate;
margins flat or sometimes slightly recurved;
domatia usually present but usually
inconspicuous. Cymes 2-11 (-16) flowered,
with a simple or once-branched glabrous
peduncle 1-5 mm long; branches, where
present, each terminated by a cymule of 4-8
flowers; bracts minute, glabrous. Flowers in
bud obtuse at apex; pedicels (1.5—)3—5 mm
long; calyx c.2 x 2 mm, cupuliform with a very
short limb and minute ovate lobes, glabrous or
with a few hairs on the lobes; corolla white,
becoming yellowish with age, 6-9 mm long,
with tube 3-5 mm long, 2-2.5 mm wide at
mouth, densely hairy adaxially, the hairs at the
mouth much denser and longer; lobes elliptic,
2.5-5.5 x 1.5-2 mm, obtuse, slightly recurved
and cucullate at apex, glabrous or sparsely hairy
towards the base adaxially, sparsely papillose
abaxially; disc shorter than calyx limb, fleshy,
glabrous; stamens erect, with filaments broad,
c.0.5 mm long, and anthers ovoid, apiculate,
2-2.5 mm long; style with stigma 6-7 mm long,
slightly protruding from mouth of corolla tube
but shorter than anthers; stigma broad, +
orbicular but 2-lobed. Fruits on erect to
ascending pedicels 5-7 mm long, pale reddish
pink when ripe, fleshy, transversely ellipsoid
or obcordate, depressed and lobed at apex, 8-
12 x 16-23 mm, or fruits occasionally 1-lobed
when slighty ellipsoid or obliquely obovoid, 8-
10 x 5-8 mm; pyrenes rugose.

Affinities: Cyclophyllum schultzii is readily
distinguishable by its usually 6-11-flowered,
pedunculate inflorescences, short corollas with

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

61

dense hairs at the mouth of the corolla tube,
and broadly obcordiform (sometimes
ellipsoidal) fruits. This species is closely related
to C. brevipes (Merr. & L.M.Perry)
S.T.Reynolds & R.J.F.Hend. in its more or less
similar leaves, flowers and fruits, but that
species differs from C. schultzii in its sessile
inflorescences with fewer flowers, and corollas
which are glabrous at the mouth of the tube.
(See under C. brevipes below.)

Variability: Attributes of the leaves of this
species are very variable. Though two forms
are recognised here based on these attributes,
specimens are sometimes difficult to place in
either taxon as they appear intermediate
between these forms. Never-the-less, the
extremes of these taxa are very distinctive.

DNA); Catchment of Habgood River, Gapuwiyak,
12°39’S,135°52'E, Dec 1987, Russell-Smith 4362 & Lucas
(BRI); Warangaya, Elcho Island, ll o 56’S,135 0 42’E, Sep
1987, Russell-Smith 3299 & Lucas (BRI).

Distribution and habitat: Arnhem Land,
Northern Territory, and offshore islands; chiefly
in coastal areas, at the edge of wet monsoon
forest. (Map 3).

Variability: The leaf blades in this form are
quite variable. Collections from near Howard
River, Howard Springs Creek (type locality),
Darwin River, Meckitt Creek, Black Jungle and
Channel Point have comparatively large,
broadly elliptic to elliptic-ovate leaf blades
which are usually abruptly obtuse at apex and
base, or are abruptly shortly and bluntly
acuminate at the apex, whereas specimens from

Key to forms of Cyclophyllum schultzii

1. Leaf blades elliptic or elliptic-ovate, (4.2-)6-7.2(-8.2 cm) wide, less than
twice as long as broad, usually obtuse at apex and base, thick, coriaceous,
on petioles 7-15 mm long; lateral nerves in 6-11 pairs

.8a. C. schultzii forma schultzii

Leaf blades narrowly elliptic or lanceolate, 1.8-3.3(-3.9) cm wide, 3 to 4
times as long as broad, usually subacute at both apex and base, thin,
coriaceous, on petioles 5-6 mm long; lateral nerves in 4-7 pairs

.8b. C. schultzii forma angustifolium

8a. C. schultzii forma schultzii

Leaf blades broadly elliptic or elliptic-oblong,
thick, coriaceous, green on adaxial surface,
pale green abaxially, usually drying brownish
coloured with nerves paler coloured.

Selected specimens: Northern Territory. Melville Island,
Garden Point, ll o 24’S,130°25E,Nov 1986, Dunlop 6845
(DNA); ditto, Apr 1987, Russell-Smith 2134 & Lucas
(DNA); Leader Creek, Gunn Point, 12°12’S,130°06E, Mar
1983, Wightman 225 (DNA); Black Jungle,
12°32’S,131°13’E, Oct 1985, Clark 27 & Wightman (BRI,
CANB, DNA); Darwin River Quarry area,
12°49’S,130°59’E, Nov 1978, Rankin 1602 (DNA);
Channel Point, 13°08’S,130°15’E, Nov 1985, Clark 76
(BRI, CANB, DNA); ditto, Jan 1986, Wightman 2509
(DNA); Howard River headwaters, 12 o 32’S,131 o 07’E, Feb
1990, Taylor 10 (DNA); Howard Springs Creek,
12°27’S,131°04E, Oct 1974, Dunlop 3686 (BRI); Adelaide
River, 13°14’S,131°05’E, Dec 1971, McKean 178 (CANB,

other areas usually have smaller or narrower
leaves. Some of those from near Adelaide River
have comparatively narrow leaves and are not
too different in appearance from some of the
specimens included under C. schultzii forma
angustifolium here. However, the leaves in that
form are usually much narrower and usually
thinner in texture than those included here under
C. schultzii forma schultzii. A few collections
from north-eastern Arnhem Land resemble ones
from far northern Queensland included under
C. brevipes here in their thin, usually elliptic or
elliptic-oblong leaf blades which possess
prominent reddish coloured dots and flecks.
However, in that species the inflorescences are
sessile, have unbranched peduncles and fewer
flowers, and the leaves are comparatively
shortly petiolate (see also under C. brevipes ).

62

These latter specimens of C. schultzii forma
schultzii probably have only young leaves on
them but study of more specimens would be
necessary to establish if this is true.

8b. C. schultzii forma angustifolium

S.T.Reynolds & R.J.F.Hend. forma nova
a C. schultzii forma schultzii foliis lamina
angustiora, textura plusminusve
membranaceiore, nervis lateralibus
paucioribus praeditis differt. Typus:
Northern Territory. Deaf Adder Creek
Gorge, 18 November 1972, P. Martensz
AE324 (holo: BRI; iso: CANB, DNA).

Leaf blades narrowly elliptic, thin, coriaceous,
reportedly discolorous and ± glaucous on their
abaxial surface when fresh, usually drying
blackish coloured or brown, with 4-7 pairs of
slender and usually obliquely arched lateral
nerves. (Fig. 6).

Selected specimens: Northern Territory. Finniss River,
Jan 1973, Byrnes 2398 (BRI); approximately 19 miles
[c.30.4 km] NNW of Oenpelli Mission, 12 o 04’S,133°—
’E, Feb 1973, Lazarides 7708 (BRI, CANB, DNA); Cooper
Creek, first billabong below Nabarlek Camp,
12°20’S,133 o 20’E, Sep 1978, Rice 2990 (CANB); Upper
East Alligator River, 12°49’S,133 0 22’E, Oct \9%l,Russell-
Smith 3869 & Lucas (DNA); Waterfall Creek, UDP Falls
area, 13° 26 , S,132°25’E, Jul 1978, Rankin 1467 (DNA,
PERTH); Edith Falls Reserve, 14 0 12 , S,132°ll , E,Oct 1977,
Must 1658 (DNA); Katherine River, Katherine, Oct 1958,
Chippendale 5042 (BRI, DNA, PERTH); ditto, Dec 1963,
Adams 802 & 803 (BRI, CANB); Katherine Gorge National
Park, Oct 1968, Byrnes 536 (BRI, DNA, PERTH).

Distribution and habitat: Arnhem Land,
Northern Territory; usually along rivers, creeks
and lagoons, or on sandstone slopes above river;
in monsoon forests, usually on sandy soil (Map

5).

Notes: Cyclophyllum schultzii forma
angustifolium is characterised by its
comparatively thin, narrowly elliptic or
lanceolate, thin leaf blades with 4-7 pairs of
slender, suboblique looping lateral nerves.

Although O. Schwarz (1927, p.101),
when describing Cyclophyllum schultzii (as
Plectronia schultzii ), made note of a narrow¬
leaved form of this species and cited collections
as representing such, namely Adelaide River,
Schultz 440, 447, 503, and Finniso River
District, Bleeser A7, he did not formally name
this form. Only duplicates of Schultz’s

Austrobaileya 6(1): 41-66 (2001)

collections from Adelaide River have been
available for this study. They ( Schultz 440
[MEL1538523], 447 [MEL1538524] and 503
[MEL 1538493] (all MEL)) all fall within the
range of variability here accepted for C. schultzii
forma schultzii. On the other hand, Bleeser A7
from the ‘Finniso’ [Finniss] River District is/
was (?) probably the only part of Schwarz’s
material referrable to C. schultzii forma
angustifolium for specimens of this form have
since been collected (in 1973) in the Finniss
River area by Norm Byrnes (see Byrnes 2398
in the list of specimens above).

Etymology: The epithet, from Latin angustus,
narrow, and folium, leaf, refers to the narrow
leaf blades in this taxon.

9. Cyclophyllum brevipes (Merr. &
L.M.Perry) S.T.Reynolds & R.J.F.Hend.
comb, nov.; Canthium brevipes Merr. &
L.M.Perry, J. Arn. Arb. 26: 231-232
(1945). Type: Papua New Guinea.
Western Division: Penzara, between
Morehead and Wassi Kussa Rivers,
December 1936, L.J. Brass 8443 (iso:
BRI).

Shrubs or small trees 2-6 m high, usually
with spreading branches; bark mottled grey;
branchlets slightly quadrangular distally, grey
or reddish brown, glabrous, usually dotted with
white lenticels. Leaves with stipules ovate,
keeled, attenuated into a usually short folded
apex, glabrous; petioles 3-9 mm long; blades
elliptic, (6.2-)10-13.5 (-14.7) x (2.5-)3.7-5(-
5.5) cm, with apex shortly acuminate or
subcaudate, and base subobtuse, abruptly
obtuse or ± rounded, thin (more so when dry),
coriaceous, glabrous, drying blackish coloured
or brown, sometimes blotched and marked with
reddish coloured resin dots, deeply channelled
over midrib proximally on the adaxial surface;
lateral nerves very slender, obliquely arched, ±
obscure adaxially; secondary venation
reticulate, obscure; domatia usually present,
usually small and inconspicuous. Cymes
(4 -) 6-8(-10)-flowered, sessile; bracts glabrous.
Flowers in bud obtuse at apex, perfumed;
pedicels 2-3 mm long; calyx c.2 x.2.5 mm,
glabrous; limb short, denticulate with lobes
ovate, minute, glabrous or with scattered hairs
distally; corolla white or yellow, 5.5-7.5 mm long,

Vanguerieae in Australia, 2. Cyclophyllum Hook.f.

63

Fig. 6. Cyclophyllum schultzii forma angustifolium. A. flowering branch x 0.8. B. pedunculate inflorescence with
branchesx2. C.flowerx5. D. LS of flower x 5. E.fruitx3. F.pyrenex3. A-D, Adams 803 (BRI); E & F, Byrnes 1187
(DNA). Del. W. Smith.

64

Austrobaileya 6 (1): 41-66 (2001)

with tube (2.5-)3.5—4.5 x c. 1.5 mm, sparsely hairy
adaxially but with hairs more dense at mouth;
lobes subelliptic, 3-4 x 1-1.5 mm, obtuse or
subacute and ± cucullate at apex, glabrous,
densely papillate at apex and on margins; disc
fleshy, as long as the calyx limb; stamens with
filaments c.0.5 mm long and anthers 1.5-1.75
mm long; style with stigma 7-7.5 mm long;
stigma c.l mm long. Fruits on ascending to
spreading pedicels 2.5-6.5 mm long, bright red
when ripe, broadly ellipsoid to obcordate,
slightly retuse or deeply and widely 2-lobed on
top, 8-10 x 10-18 mm; pyrenes with slightly
granular surface when dry.

Selected specimens: Papua New Guinea. Lake
Daviumbu, Middle Fly River, Aug 1936, Brass 7470
(BRI); Western District, Weam, Bensbach subdistrict,
8°38’S,141°07 , E, Jul 1967, Ridsdale [NGF33541]
(BRI, NSW); Pangoa Airstrip, Lake Murray, Morehead
subdistrict, 8°05’S,141 o 15’E, Mar 1968, Millar
[NGF35421] (BRI). Queensland. Cook District: Cape
York, 11°—’S,141°—’E, Nov 1955, White 1157 (BRI);
Bamaga, U°1-’S, 142°3-’E, Oct 1965, Smith 12381
(BRI); ditto, Oct 1965, Jones 3826 (CANB); Cockatoo
Creek, at Telegraph Crossing Road, 11°39’S,142 0 27’E,
Mar 1992, Johnson 5098 (BRI); Vyse Crossing, 19 km
N of Lorim Point, Weipa, 12°30’S,141 o 53’E, Jan 1981,
Morton AM1030 (BRI, MEL); Rocky Creek,
12°06’S,142°33’E, Jul 1984, Puttock & King

[UNSW16940] (BRI, UNSW); Wenlock River, southern
bank at Moreton Telegraph Station, 12°27’S,142°38’E,
Oct 1989, Neldner 2804 & Clarkson (BRI); Wabum
Creek, about 18 km from Aurukun on road to Merluna,
13°16’S,141°50 , E, Dec 1981, Clarkson 4131A (BRI,
QRS); Coconut Creek, downstream from Beagle North
Camp, about 40 km SSW of Weipa, 13 o 01’S,141°47’E,
Dec 1981, Clarkson 4163 (BRI); Lakefield National
Park, Pocket Waterhole, “Bizant”, 14°4-’S,144°0-’E,
Oct 1985, Williams 85264 (BRI); Hann River,
15°12’S,143°52 , E, Nov 1971, Stocker 842 (BRI);
Magnificent Creek, Kowanyama, 15 0 2-’S,141°4-’E,
Mar 1990, Birchley 4 (BRI).

Distribution and habitat: Southern Papua New
Guinea to Cape York Peninsula, Australia;
usually in riparian rainforests on sandy soil.
(Map 5).

Affinities: Cyclophyllum brevipes is
characterised by its shortly petiolate elliptic
leaves which usually dry blackish coloured or
dark brown, its 4-10-flowered sessile cymes,
and its obcordate or ellipsoid fruits. It is closely
related to C. schultzii with which it shares more
or less similar leaves, inflorescences, flowers
and fruits, but that species differs from
C. brevipes mainly by its pedunculate
inflorescences.

Vanguerieae in Australia, 2. Cyclophyllum Hook.f. °

These two species may be distinguished using the following key.

1. Inflorescences pedunculate, with (2—)4—11(—16) flowers; peduncle
sometimes branched; corolla tube 3-5 mm long, 2-2.5 mm wide at
mouth where very densely hairy; fruit usually obcordiform, deeply

2-lobed at apex, 8-12 x 16-23 mm; petiole 5-15 mm long. C. schultzii

Inflorescences sessile, (4-)7-10-flowered; corolla tube 2.5-4.5 mm long, to
1.5 mm wide at mouth where sparsely hairy; fruit ellipsoid or obcordiform,
retuse or deeply 2-lobed at apex, 8-10 x 10-18mm; petiole 3-9 mm long.C. brevipes

Notes: The Queensland specimens match
closely those from Papua New Guinea especially
the isotype of C. brevipes at BRI, and the one
collected by Len Brass from Lake Daviumbu on
the Middle Fly River, also at BRI. Queensland
plants are recorded as growing in habitats similar
to those for plants in Papua New Guinea.

The fruits of this species are reported as
edible [for humans].

Variability: This species varies greatly in the
shape and size of its leaves and fruits, both in
Australia and in Papua New Guinea.
Specimens with shortly petiolate, blackish
brown or brown, elliptic leaf blades,
obcordiform fruits and glabrous flowers are
typical of this species. However, specimens
with narrowly elliptic leaf blades which dry pale
brown and have typical obcordiform fruits, or
ones with typical elliptic leaf blades but with
small obovoid fruits, are also present. These
probably represent distinct forms of the species,
but more specimens for study would be
necessary to be certain if this is correct.
Moreover, a few specimens, e.g. Jones 3826
and Johnson 5098 cited above, resemble
specimens from eastern Arnhem Land included
here in C. schultzii in their thin, elliptic leaf
blades which are prominently marked with
numerous, small, reddish coloured dots or
flecks but that species differs from C. brevipes
by its pedunculate inflorescences which have
a greater number of flowers in each.

Acknowledgements

We thank Les Pedley for help with the Latin
diagnoses, colleagues at BRI for collecting many
of the ‘ Canthium' species and for their
comments about the habitat of the species
concerned, Clyde Dunlop for his comments on
the manuscript, and Laurie Jessup, Gordon
Guymer and Clyde Dunlop for their assistance

in locating specimens and photographing types
and literature during their term as Australian
Botanical Liaison Officer at The Herbarium,
Royal Botanic Gardens, Kew, England, Diane
Bridson for her comments on Psydrax and
Canthium, and assistance with specimens
during visits to Kew (S.T.R. and, later, R.J.F.H),
Will Smith for the illustrations and maps, David
Halford and Kym Sparshott for assistance with
measurements, maps and illustrations, the
Directors/Keepers of the herbaria AD, BM,
CANB, CGE, DNA, K, L, MEL, NSW, P,
PERTH, PVNH, QRS and UNSW for allowing
me (S.T.R.) full access to specimens in their
institutions and for the loan of herbarium
material, and the Australian Biological
Resource Study, ABRS, Environment Australia
f or a grant to (S.T.R.) to undertake research in
the genus Canthium in Australia.

References

Bridson, D.M. (1985). The reinstatement of Psydrax
(Rubiaceae subfamily Cinchonoideae tribe
Vanguierieae ) and a revision of the African species.
Kew Bulletin 40(4):687-725.

-(1987). Studies in African Rubiaceae-

Vanguerieae: a new circumscription of
Pyrostria and a new subgenus, Canthium
subgenus Bullockia. Kew Bulletin 42(3):611-
639.

-(1992). The genus Canthium (Rubiaceae-

Vanguerieae) in Tropical Africa. Kew Bulletin
47(3):353-401.

Guilluamin, A. (1948). Cyclophyllum. Flore analytique
et Synoptique de la Nouvelle-Caledonie
Phanerogames 332-333. Office de la
Recherche Scientifique Coloniale, 22 Rue
Oudinot Paris.

Lebler, B.A. (1978). The Canthiurms of South-eastern
Queensland. Queensland Agricultural Journal
104 (6): 527-532. Brisbane: Government
Printer.

Austrobaileya 6 (1): 41-66 (2001)

66

Reynolds, S.T. (1997). Canthium, in R.J.F. Henderson
(ed), Queensland Plants Names and
Distribution 180-181. Brisbane: Queensland
Herbarium, Department of Environment.

Reynolds, S.T. & Henderson, R.J.F. (1999). Vanguerieae
A.Rich, ex Dum. (Rubiaceae) in Australia, 1.
Everistia S.T.Reynolds & R.J.F.Hend.
Austrobaileya 5(2): 353-361.

Schwarz, O. (1927). Plantae novae vel minus cognitae
Australiae tropicae. Feddes Repert. 24: 80-
109.

Smith A.C. & Darwin, S.P. (1988). Psydrax Gaertner,
Cyclophyllum Hook, f., in Flora Vitiensis Nova
(4): 229 - 231 .

Revision of the Macrozamia miquelii (F.Muell.)
A.DC. (Zamiaceae section Macrozamia) group

David L. Jones*, Paul I. Forster** and Ish K. Sharma*

Summary

Jones, David L., Forster, Paul I., Sharma Ish K. (2001). Revision of the Macrozamia miquelii (F.
Muell) A.DC ( Zamiaceae section Macrozamia ) group. Austrobaileya 6 (1): 67-94. The
Macrozamia miquelii group consists of seven species, M. cardiacensis P.I.Forst. & D.L.Jones, M.
douglasii W.Hill ex F.M.Bailey, M. longispina P.I.Forst. & D.L.Jones, M. macleayi Miq., M.
miquelii (F.Muell.) A.DC., M. mountperriensis F.M.Bailey and M. serpentina D.L.Jones & P.I.Forst.
sp.nov., all occurring in eastern Queensland. The complicated typification of Macrozamia
douglasii W.Hill ex F.M.Bailey and Encephalartos douglasii F.Muell. is resolved with lectotypes
selected for both names. A key to the species in the group is provided and all species are illustrated.
A partial electrophetic analysis of the complex is also presented.

Keywords: Zamiaceae, Key, taxonomy, electrophoresis, Macrozamia miquelii, Macrozamia
serpentina

* Centre for Plant Biodiversity Research, Australian National Herbarium, G.P.O. Box 1600, Canberra,
Australian Capital Territory, 2601, Australia.

** Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens Mt Coot-tha,
Mt Coot-tha Road,Toowong, Queensland, 4066, Australia

Introduction

The genus Macrozamia Miq. consists of thirty-
eight named species and a few others yet to be
recognised formally. The genus is endemic to
mainland Australia with the vast majority of
species (c. 36) occurring in the eastern states
of Queensland and New South Wales. Another
six (including unnamed species) are known to
occur in Western Australia and a single, isolated
relict species is restricted to Central Australia.
While some species of Macrozamia, such as
M. riedlei (Fisch. ex Gaudich.) C.A.Gardner
and M. communis L.A.S Johnson, have a wide
distribution, others, such as M. cranei
D.L.Jones & P.I.Forst. and M. viridis D.L.Jones
& P.I.Forst., are narrow endemics restricted to
relatively small areas and specialised habitats.

This paper examines the systematics,
morphology and relationships of the group of
species centred around Macrozamia miquelii
(F.Muell.) A.DC. in Macrozamia section
Macrozamia. This group is defined by the

Accepted for publication 30 March 2001

intermediate size of the plants, untwisted or
slightly twisted leaves that have spinescent
petioles and hypostomatic leaflets with the
lower leaflets grading into pinnacanths or the
latter absent in some species. These features
are shared by a group of three species allied to
M. communis L.A.S.Johnson, but the
M. miquelii group can be distinguished from
these by narrow-based spines on the female
sporophylls (c. 2-5 mm wide cf. 5-12 mm wide
in the M. communis group). Species of the
M. miquelii group form small to large cycads
that are generally much larger than species in
Macrozamia section Parazamia and smaller
than the largest species of M. section
Macrozamia such as M. macdonnellii (F.Muell.
ex Miq.) A.DC., M. moorei F.Muell. and
M. johnsonii D.L.Jones & K.D.Hill, all of
which have amphistomatic leaflets. All of these
species in Macrozamia section Macrozamia
appear to be insect pollinated, either by thrips,
beetles or a combination of the two (Forster
et al. 1994; Forster, unpubl.; Mound & Terry
2000 ).

The earliest described species of the
group, Macrozamia miquelii, was first named
in Encephalartos by Mueller (1862), but was

68

soon transferred to Macrozamia by De Candolle
(1868). Additional species were added by
Miquel (1868a,b) and Bailey (1883, 1886).
Johnson (1959) took an extremely conservative
view of the complex reducing all published
names to synonymy of M. miquelii and also
lectotypified this name using a type from
Rockhampton (Forster 1999a). Further new
species were recently added by Forster & Jones
(1998) as part of the Flora of Australia account
compiled by Hill (1998).

Although concise descriptions of five
species from this group have been provided in
the Flora of Australia by Hill (1998), there are
numerous discrepancies in measurements
between our observations and his accounts of
Macrozamia douglasii, miquelii and
M. mountperriensis, necessitating redescription
of these species. Additionally the
circumscription of M. miquelii is altered in the
current paper with M. macleayi being
resurrected from synonymy. Hence we provide
detailed descriptions for all seven species in the
group.

Materials and Methods

All species dealt with in this paper were
examined in the field. Measurements cited here
were made mainly from living material of adult
plants, with some supplementation from
herbarium collections, particularly those at BRI,
CANB and MEL. Juvenile material of all
species will generally have quite dissimilar
leaves in terms of dimensions and component
parts. The key is designed to be used with fertile
material and adult leaves. When referring to
“leaflets” in the descriptions, it indicates the

Austrobaileya 6(1): 67-94 (2001)

total number of leaflets in the leaf. Leaflet
dimensions are based on mature leaves and the
leaflets from the central portion of the leaf. More
accurate identifications will be arrived at if
collectors either use the key on plants in situ or
collect a range of leaves and cones, as there is
often considerable variation even on a single
individual. All types have been seen unless
indicated n.v. Specimens of known sex are cited
as A (female) or B (male), or C (seedling)
following the collector’s number. Some locality
details in the citation of specimens are
abbreviated or omitted for conservation
purposes.

Limited allozyme analysis has been
attempted for the complex but should not be
regarded as all-encompassing. One species
(M serpentina ) was not included in the analysis
due to the loss of collected samples. The
methodology used for the starch gel
electrophoresis is the same as previously
described (Sharma et al. 1999). Table 1 shows
localities of species, vouchers and the number
of samples used for analysis. Nei’s genetic
distance coefficient (Nei 1978) was used to
measure the level of genetic differentiation
among populations and species. Based on these
values, a dendrogram (Fig. 1) was constructed
(see Sharma et al. 1999 for details), which
revealed that all the populations clustered into
groups corresponding with the respective
species.

Population codes, voucher numbers and
sample size of the six species of Macrozamia
examined in this study are listed below.
Voucher details are given in the specimen
citation for individual taxa (Table 1).

Table 1. Population codes, voucher details and sample size of
collections used in isoenzyme analysis.

M. mountperriensis

BRO

Forster PIF9343 & Machin

26

SNY

Forster PIF13321 & Machin

26

GOO

Forster PIF2771

14

SEA

Forster PIF13372 & Machin

26

MON

Forster PIF13984 & Machin

14

SCH

Jones DLJ 6340 & Jones

26

Jones et al. Macrozamia miquelii group

69

M. longispina

WOO Forster PIF13374 & Machin 26

WID Forster PIF12137A-C& Machin 32

M. miquelii

BYE Jones DLJ 9393 32

ARC Forster PIF12253A-B & Machin 32

KPL Forster PIF12268 & Machin 32

MOR Forster PIF12251A-C & Machin 32

M. macleayi

BLK Forster PIF12277A-B & Machin 32

BUL Forster PIF12289A-B & Machin 28

BUR Forster PIF12299A-B & Machin 28

BAN Forster PIF13983 & Machin 28

KAL Forster PIF13980A-B & Machin 28

KOO Forster PIF12285A-B & Machin 28

IMB Forster PIF13146 & Machin 28

M. douglasii

DOG Forster PIF9346 & Machin 26

COO Machin PM17 21

M. cardiacensis

CAR Forster PIF13109A-C & Machin 32

Taxonomy

Key to species of the Macrozamia miquelii group

1. Lower 8 or more leaflets reduced to pinnacanths.2

Lower 4 or less leaflets reduced to pinnacanths, or pinnacanths absent.5

2. Female cones narrowly ovoid; male cones 35-40 cm long.1. M. cardiacensis

Female cones cylindrical to barrel-shaped; male cones 12-28 cm long.3

3. Male cones 2.5-3.5 cm diameter; microsporophylls 8-12 x 4-8 mm.7. M. serpentina

Male cones 3.5-6.5 cm diameter; microsporophylls 12-25 x 8-15 mm.4

4. Leaves glossy above, thin-textured; female cones 7-10 cm diameter.4. M. macleayi

Leaves dull to slightly glossy above, thick-textured; female cones 10-15

cm diameter.5. M. miquelii

5. Lower 1-4 leaflets reduced to pinnacanths.2. M. douglasii

Pinnacanths absent.6

70

Austrobaileya 6(1): 67-94 (2001)

1

2

3

4

5

i 6

J-1-1-1_I_I_I_I_I_I_I_I_l

.40 .33 .27 .20 .13 .07 .00

Fig. 1. Allozyme Based Dendrogram of Macrozamia species and populations based on Nei’s unbiased Genetic Distance.
1 =M.mountperriensis\ 2=M.longispina; 3=M.miquelii; 4=M.macleayr, 5=M.douglasii\ 6=M.cardiacensis.

Jones et al. Macrozamia miquelii group

71

Fig. 2. Macrozamia cardiacensis. Adult in habitat with P.Machin for scale. Forster PIF12151 & Machin. Photo: P.I.Forster.

6. Leaflets 50-110,6-9 mm wide; male cones 12-25 x 3-4 cm; distal

megasporophylls with spines 2.5-4 cm long. 6. M. mountperriensis

Leaflets 100-140,3-6 mm wide; male cones 8-15 x 2.5^1 cm; distal

megasporophylls with spines 4.5-7 cm long. 3. M. longispina

1. Macrozamia cardiacensis P.I.Forst. &
D.L.Jones in P.McCarthy (ed.), FI.
Australia 48: 717 (1998). Type:
Queensland. Wide Bay District: Cardiac
Hill, Mt Walsh National Park, 26
February 1993, P.I.Forster 13109A &
P.Machin (holo: BRI).

Caudex usually subterranean, occasionally
emergent, erect, columnar or barrel-shaped, to
40 cm long, 20-40 cm diam., unbranched.
Young leaves light green. Mature leaves
elliptic-lanceolate in outline, 1-2 m long,
obliquely erect to spreading, dark green, glossy,
flat in cross-section, arching in profile, 10-20 in
a moderately dense crown; expanded leaf base
9-12 cm x 3-4 cm, covered with grey-brown,
soft woolly hairs; petiole (including the
expanded base) 30-40 cm long, 1.5-2 cm wide
at the first leaflet, flat adaxially, convex and
angular abaxially; rhachis straight, not twisted,
flat adaxially, the cross-section similar to that

of the petiole; leaflets 100-140 per leaf, inserted
at about 40° to the rhachis, crowded, distal
leaflets densely packed, proximal leaflets
progressively more widely spaced, linear, 20-
31 cm x 0.8-1.2 cm, flat, moderately thin-
textured, hypostomatic, dark green & glossy
adaxially, paler beneath, contracted proximally
to a pale yellow callous base, margins flat,
entire, tapered to a pungent apex; lower 4-7
pairs of leaflets gradually reduced to rigid
pinnacanths. Male cones 1-5, cylindrical, 35-
40 cm x 6.5-8 cm, straight or curved with age,
green; peduncle 15-30 cm x 1.5-2.5 cm,
elliptical to round in cross-section;
microsporophylls cuneate, 2.2-3.2 cm x 1-1.5
cm, with an erect, apical spine 0.2-2.5 cm long.
Female cones 1 or 2, narrowly ovoid, 32-36
cm x 11-14 cm, green; peduncle 20-25 x 1.6-
2.5 cm, elliptical to rounded in cross-section;
megasporophylls 4-4.5 cm x 3.5-4.7 cm,
broadly cuneate, with an erect apical spine 1-5
cm long, the distal sporophylls with spines 4.5-

Fig. 3. Macrozamia cardiacensis. Female plant with cones. Forster PIF13109A & Machin (type). Photo: P.I.Forster.

Austrobaileya 6(1): 67-94 (2001)

Fig. 4. Macrozamia cardiacensis. Detail of female cones. Forster PIF13109A & Machin (type). Photo: P.I.Forster.

Jones et al. Macrozamia miquelii group

73

Map 1. Distribution of Macrozamia cardiacensisX, M. douglasii +, M. longispinaW, M. macleayi A , M. miquelii ♦,
M. mountperriensis ★, M. serpentina # based on herbarium records in BRI, CANB and MEL.

5 cm long. Seeds oblong to ovoid, 2.2-3.5 cm x
1.2-2 cm, sarcotesta orange to dark red. (Fig.
2-4).

Selected specimens : Queensland. Wide Bay District: Mt
Walsh N.R, Coast Range, Oct 1992, Forster PIF12151 &
Machin (BRI); ditto, Feb 1993, Forster PIF13109B &
Machin (BRI); ditto. May 1994, Forster PIF15232 & Bean
(BRI); ditto, Mar 1995, Forster PIF16367 (BRI); Coast
Range, Oct 1995, Grimshaw PG2212 (BRI); ditto,
Grimshaw PG2401 (BRI); Mt Walsh N.P., Coast Range,
Aug 1992, Machin PM16-19 (BRI); Mt Walsh, 6.5 km
S of Biggenden, May 1977, Telford 5322 (CANB).

Distribution and habitat : M. cardiacensis (Map
1) is restricted to the Mount Walsh National
Park near Biggenden in the Wide Bay District
of Queensland. It grows in skeletal soils on
steep to precipitous slopes on substrates derived
from rhyolites or andesites at altitudes between

500 and 640 m in open eucalypt forest.
Associated canopy species are Eucalyptus
andrewsii, E. decolor , E. acmenoides and
Corymbia citriodora.

Phenology : Cones shed pollen in November
and December. Both male and female cones are
attended by the thrips Cycadothrips chadwickii
and a species of Tranes (P.I.Forster, pers. obs.
Dec. 2000). Macrozamia cardiacensis is the
only species in the M. miquelii group that has
been observed to have both of these insects
present. Ripe seeds are dispersed from March
to May.

Notes : Plants usually have a subterranean trunk
but on rocky sites or in shallow soils an
emergent trunk is often formed.

74

Austrobaileya 6(1): 67-94 (2001)

Macrozamia cardiacensis is similar to
both M. miquelii and M. douglasii. It differs
from M. miquelii by its larger cones with longer
apical spines on the distal megasporophylls and
from M. douglasii by its more numerous
pinnacanths, less prominent callous base and
longer apical spines on the distal
megasporophylls. It is perhaps most closely
allied to M. douglasii on morphological
characters, and the limited allozyme analysis
based on few populations of the two species
tends to support this (Fig. 1).

Conservation status : Although of restricted
distribution M. cardiacensis is locally abundant
in two localities, growing in dense colonies with
strong seedling recruitment. This species is
listed as Rare on the schedules of the
Queensland Nature Conservation Act 1992
although it is conserved in Mt Walsh National
Park.

Etymology: Named for the type locality of
“Cardiac Hill” and also alluding to the difficulty
experienced by the co-collectors of the type in
transporting the heavy female cones.

2. Macrozamia douglasii W.Hill ex
F.M.Bailey, Syn. Queensland FI. 500
(1883).Type: Queensland. Wide Bay
District: [comprises a pressed leaf]
Fraser Island, [forwarded 24 April 1882
to K by J.Pink], Sheridan s.n. (lecto (here
designated): K, photo!).

Encephalartos douglasii F.Muell., Chem.
Drugg. Australas. 5: 80-81 (14 February
1883); Macrozamia tridentata var.
douglasii (F.Muell.) J.Schust. in
A.Engler, Pflanzenr. 99 (IV, I): 90
(1932). Type: Queensland. Wide Bay
District: [comprises 5 bags of fruit & a
photograph of an intact cone] Fraser
Island, [collected prior to 17 Jan 1883
when forwarded to Mueller],
A.McDowall s.n. (lecto [here
designated]: MEL269564; isolecto: K [2
sheets of three portions of the one
pressed frond, plus a copy of the photo
of the intact cone] photo!).

Illustrations : Ballard (1936: t. 3310,3311);
Jones (1993: 237).

Caudex subterranean or emergent, to 1 m tall
and 70 cm diam., unbranched. Young leaves
light green. Mature leaves elliptic-lanceolate in
outline, 2-3.5 m long, erect to obliquely erect
or widely spreading, bright green to dark green,
flat in cross-section, arching in profile, 30-90
in a dense crown; expanded leaf base 10-15
cm x 3-4 cm, covered with greyish-white, soft
woolly hairs; petiole (including the expanded
base) 40-60 cm long, 12-20 mm wide at the
first leaflet, green, flat or grooved adaxially,
convex and angular abaxially; rhachis straight,
not twisted or slightly twisted, pale green, the
cross-section similar to that of the petiole;
leaflets 120-200, inserted at about 40° to the
rhachis, widely spreading, moderately crowded,
distal leaflets densely packed, proximal leaflets
becoming more widely spaced, linear,flat,
moderately thin-textured, hypostomatic, 25-35
cm x 8-12 mm, dark green and glossy adaxially,
paler beneath, contracted proximally to a
prominent white callous base, margins entire,
tapered to a sharply pointed apex; lower leaflets
gradually reduced to 1-4 pairs of rigid,
moderately long, yellowish pinnacanths. Male
cones 1-3, cylindrical, 20-35 cm x 5-7 cm,
straight or curved with age, green; peduncle 20-
35 cm x 2-3 cm, elliptical in cross-section;
microsporophylls cuneate, 2-3.5 cm x 1.5-2.5
cm, with an erect, apical spine 0.5-4 cm long.
Female cones 1-3, cylindrical to barrel-shaped,
35-45 cm x 10-18 cm, green; peduncle 25-45
cmx 2-3 cm; megasporophylls broadly cuneate,
2.5-3.5 cm x 4-6 cm, with an erect, apical spine
1-4 cm long, the distal sporophylls with spines
3-4 cm long. Seeds oblong, 2.8-4 cm x 1.8-
2.5 cm, sarcotesta orange to red. (Figs. 5-8).

Selected specimens : Queensland. Wide Bay District:
Cowra Logging Area, Tuan State Forest, Jan 1992, Forster
PIF9346 A & B & Machin (BRI, CANB); ditto, Nov 1992,
Forster PIF12330 Sc Machin (BRI); Great Sandy National
Park - Cooloola section, Aug 2000, Forster PIF25968 &
Kokubugata (BRI); Fraser Island, Nov 1930, Hubbard 4552
(BRI; K n.v.); Fraser Island, c. 0.1 km from Dilli towards
Ungowa, Apr 1992, Jones 9414 & Jones (CANB); Fraser
Island, c. 2 km W of Eumong, May 1992, Jones 9429 &
Jones (BRI, CANB); Shark Creek, opposite southern end
of Fraser Island, Jan 1928 ,Kajewski 1 (BRI); Fraser Island,
May 1992, Machin (CANB); Tuan Ck, near Tin Can Bay,
May 1992, Machin (CANB); Cooloola area, S.F. 451
Womalah, Jun 1993, Machin PM7 (BRI); Cooloola, Stutz
Hut road, S.F. 915, Jul 1992, Machin PM21-22 (BRI);
Cooloola, off Tinnanbar road, S.F. 451 Womalah, Jul 1992,
Machin PM23-25 (BRI); Cooloola State Forest, Sep
1970, Moriarty 434 (CANB); Fraser Island, Mar 1951,

Jones et al. Macrozamia miquelii group

75

Fig. 5. Macrozamia douglasii. Habit. Fraser Island. Photo: D.L.Jones.

Webb SW4761 (CANB); Fraser Island, Oct 1921, White of TraneS weevils (Forster et al 1994). Ripe

AQ142034 (BRI). seeds are dispersed from March to April.

Distribution and habitat: Macrozamia
douglasii (Map 1) occurs on Fraser Island,
where it is extremely abundant, and on adjacent
coastal districts near Cooloola north of Noosa,
both regions in the Wide Bay District. It grows
at altitudes between 10-80 m in tall open forests
developed on old coastal dunes, along streams
and on the fringes of rainforest, with some
specimens occurring within the rainforest
canopy. The soils are deep grey to white sands
with a permanent water table.

Phenology: Pollen shedding occurs in
November with the cones attended by hordes

Typification: The typification of this twice
published species is very complex. The species
was first named Encephalartos douglasii by
Mueller (1883) and soon after, and apparently
independently, as Macrozamia douglasii by
Bailey (1883). Both names were validations of
the nomen nudum Macrozamia douglasii
W.Hill (1879), but when used in Macrozamia ,
Bailey’s name takes precedence. Bailey did not
explicitly cite a type specimen (which was not
unusual at the time) and it appears that a
progression of material was sent to various
herbaria and botanic gardens by a range of
collectors around this time with the resultant

76

Austrobaileya 6(1): 67-94 (2001)

Fig. 6. Macrozamia douglasii. Female cone. Fraser Island. Photo: D.L.Jones.

descriptions being compiled from all (or at least
most) of them. Bailey named this species “After
the Hon. John Douglas” thus validating the
unpublished name of Walter Hill’s. Further on
he states “( Fraser’s Island.)”. This has been
incorrectly interpreted as “Fraser Island, Qld,
Douglas; holo: BRI” by Hill (1998); however,
no such specimen exists at BRI. In the Synopsis
Bailey did not explicitly cite who the collectors
of any new taxa were. Hence it does not
necessarily follow that Douglas collected
material of this species at all, and it is probable
that the material used by Bailey in his
description was collected by W. Hill or one of
his contacts. W. Hill first coined the specific
epithet and stated “Frazer’s Island. During my

visit to the above island, in April last, I took the
opportunity of making a collection of the
various seeds and plants flourishing there, and
I am glad to say that I was enabled to secure a
number of very interesting specimens, including
Macrozamia douglasii, W. H. ( Goulbine ) (the
nuts of this tree are largely used as an article of
food by the aborigines)”(Hill 1879). Ballard
(1936) cites collections by W. Hill in 1881, J.
Pink in 1882 and A. McDowall in 1883, but does
not state where these collections are deposited.

We have been able to locate four separate
collections of this species from the early 1880’s
deposited in K and MEL, all of which must be
critically examined for the typification of these

Jones et al. Macrozamia miquelii group

77

Fig. 7. Macrozamia douglasii. Male cones. Forster PIF12330 & Machin. Photo: PI.Forster.

Fig. 8. Macrozamia douglasii. Detail of apical portions of male cones. Forster PIF12330 & Machin. Photo: PI.Forster.

78

Austrobaileya 6(1): 67-94 (2001)

two names. No potential type material was
located at BRI, despite extensive searching.
None of these collections are by John Douglas.
There are only seven collections by Douglas at
BRI, all of which are ferns collected in June
1893 from Thursday Island where he was
Government Resident and Police Magistrate
(Joyce 1972).

Collections in question are: Specimen #1:
MEL237356 (labelled as syntype). This
comprises a pressed, partial and somewhat
immature leaf and is labelled as ” Encephalartos
Douglasii F.vM. Macroz. Douglasii Ex horto.
bot. Brisbane 1881”.

Specimen #2: MEL269564 (labelled as
syntype). This comprises four bags of seed and
cone remnants and is accompanied by a
photographic print of an intact cone in a wired
(or perhaps string) cage and a long letter to
Mueller from A.McDowall of Maryborough
dated 17 January 1883. According to the letter
four ripe cones were originally sent from Fraser
Island. It is reasonable to assume that the
collector of this material was A.McDowall. A
copy of this photographic print is also at K.

Specimen #3: K. This comprises a pressed
partial leaf and is accompanied by a letter by
James Pink of the Brisbane Botanic Gardens
to Thiselton-Dyer at K that is dated 24 April
1882. In the letter Pink quite clearly states that
the material of Macrozamia was collected by
“Mr Sheridan Collector of Customs & Police
Magistrate Maryborough”.

Specimen #4: K. This comprises a whole
pressed leaf that is divided into three pieces and
mounted on two sheets. The material is
accompanied by a letter from Mueller to
Thiselton-Dyer dated 17 February 1883 where
it is stated that the collection is by A.McDowall.
This material appears to be the leaf material that
originally accompanied the four bags of seed in
MEF (see specimen #2). Mueller states in the
letter “It was my intention, dear Mr Dyer, to
have sent you a female amentum of
Encephalartos Douglasii by this mail, but it is
not yet dry, nor is the photograph likely to get
ready in time. So I shall keep back also leaves
and male cone til next post”.

Mueller (1883) in the description of

E. douglasii quite clearly states that “through
the kindness of Mr. A.McDowall, the districts’
lands office and surveyor of Maryborough,
Wide Bay, to place this cycadeous plant on
diagnostic record”. The name Encephalartos
douglasii F.Muell. is adequately typified by the
A.McDowall collection (specimen # 2), as this
material is annotated by Mueller, collected prior
to the publication of the name and referred to
in the protologue. The collection has been split
between MEF and K, with the former
designated as lectotype and the latter
isolectotype for this name.

This then leaves the problem of
typification for Macrozamia douglasii W.Hill
ex Bailey. As no type was explicitly cited, we
are left with a number of options, either
selection of a lectotype from the available
collections or selection of a neotype. We cannot
prove that the McDowall collection was seen
by Bailey and it would appear that McDowall
sent the material directly to Mueller, hence it is
not appropriate to consider the type of both
names to be the same.

Bailey would undoubtedly have seen the
live material acquired by Hill or Sheridan. It is
quite possible that Mueller sent a copy of his
description of Encephalartos douglasii to
Bailey and so the name Macrozamia douglasii
is merely a new name in a different genus for
the entity based on the McDowall specimen.
The collection by Sheridan at K (specimen #
3), although only of a pressed leaf, is here
designated as the lectotype for the name
Macrozamia douglasii as it comprises a mature
leaf undoubtedly prepared from a live plant in
the Brisbane Botanic Gardens, and is likely to
have been seen by Bailey.

Notes: Macrozamia douglasii is similar to
M. miquelii but can be distinguished by the
lustrous dark green leaves, extremely
conspicuous, intensely white callous bases and
only the lowest 1-4 pairs of leaflets reduced to
spine-like pinnacanths. In comparison to
M. douglasii, M. cardiacensis has fewer leaves
in the crown, pale yellow callous bases and
smaller cones with longer spine-like
appendages on the distal megasporophylls. The
allozyme analysis indicates a closer relationship
between M. douglasii and M. cardiacensis than

Jones et al. Macrozamia miquelii group

79

between M. douglasii and the other species
(Fig. 1).

The male cones of M. douglasii are
destroyed rapidly by the larvae of a host-
specific Curculionid weevil (Tranes sp.) after
shedding pollen (Forster et al. 1994). This
activity results in a complete lack of male cones
on plants once pollen shedding has ceased and
is one reason for a paucity of preserved material
in herbaria. Hill (1998) states that “pollen cones
not seen”, yet these are described by both
Mueller (1883), Bailey (1883) and Ballard
(1936) and well illustrated in the latter.

Conservation status: This species is locally
abundant and grows in scattered, sometimes
dense colonies with strong seedling
recruitment. It is not rare or threatened and is
very well conserved in Cooloola and Fraser
Island National Parks.

Etymology: Named for John Douglas (1828-
1904), a politician and administrator in
Queensland and British New Guinea (Joyce
1972). Douglas, a short-term Premier of
Queensland, was considered a “leader in
Brisbane’s intellectual, literary and religious
circles” and became involved in a number of
societies (e.g. Royal Society of Queensland,
Acclimatization Society of Queensland) along
with Hill and Bailey. The naming can be
interpreted as being one of political patronage.
Port Douglas in north Queensland is also named
after him.

3. Macrozamia longispina P.I.Forst. &
D.L.Jones in P.McCarthy (ed), Flora of
Australia 48: 111 (1998). Type:
Queensland. Wide Bay District: State
Forest 639, Wrattens, 25 October 1992,
P.I.Forster PIF12137B & P.Machin
(holo: BRI).

Illustrations : Forster & Osborne (2001).

Caudex subterranean or less commonly an erect
emergent trunk to 30 cm long and 30 cm diam.,
unbranched. Young leaves light green. Mature
leaves elliptic-lanceolate in outline, 1-1.5 m
long, obliquely erect to spreading, dark green,
glossy, flat in cross-section, arching in profile,
6-20 in a moderately dense crown; expanded
leaf base 25-35 x 1.2-1.5 cm, covered with

grey-brown, soft woolly hairs; petiole
(including the expanded base) 40-50 cm long,
0.4-0.5 cm wide at lowest pair of leaflets, flat
adaxially, convex and angular abaxially; rhachis
straight, not twisted, green, the cross-section
similar to that of the petiole; leaflets 100-140,
inserted at about 40° to the rhachis, widely
spreading, crowded, distal leaflets densely
packed, proximal leaflets more widely spaced,
narrowly linear-lanceolate, 25-32 cm x 3-6
mm, flat, thin-textured, hypostomatic, dark
green and glossy adaxially, paler beneath,
contracted proximally to a white callous base,
margins entire, tapered to a pungent apex; lower
leaflets reduced in size but not spine-like.
Male cones 1-5, cylindrical to fusiform, 8-15
cm x 2.5-4 cm, straight or curved with age,
green; peduncle 15-28 cm x 0.5-1.5 cm;
microsporophylls cuneate, 1.2-1.6 cm x 0.8-
1.3 cm, with an erect, apical spine 0.5-2 cm
long. Female cones 1-3, narrowly ovoid, 13-
19 cm x 6-8 cm, green; peduncle 20-26 x 1-

1.5 cm; megasporophylls broadly cuneate, 2.8-

3.5 x 3.0-3.7 cm, with an erect, apical spine
2.5-7 cm long, the distal sporophylls with
spines 4.5-7 cm long. Seeds oblong to obovoid,
2-2.5 cm x 1.5-2 cm, sarcotesta orange to red.
(Figs. 9-10).

Selected specimens: Queensland. Wide Bay District:
Myravale, Dec 1964, Clifford (AQ142047 & AQ321067)
(BRI); ibid, Apr 1965, Clifford (AQ142047) (BRI); S.F.639
Wrattens, Oct 1992, Forster PIF12137A-C (BRI); ditto,
Feb 1993, Forster PIF13140 (BRI); Widgee Mt, Oct 1992,
Forster PIF12120A & B (BRI); ditto, Feb 1993, Forster
PIF13143 & Machin (BRI); ditto, Apr 1996, Forster
PIF19138 & Leiper (BRI); S.F. 57 St Mary, Jun 1993,
Forster PIF 13374 & Machin (BRI).

Distribution and habitat : Macrozamia
longispina (Map 1) occurs in south-eastern
Queensland where it is mainly restricted to the
vicinity of Glastonbury and Widgee west of
Gympie, with an outlying population to the
north at State Forest 57. It grows at altitudes
between 420 and 680 m on slopes and ridges in
tall moist eucalypt forest in shallow skeletal soils
derived from serpentinite, or more rarely on deep
sand. Common canopy species include
Eucalyptus biturbinata, E. acmenoides,
E. tereticornis and Corymbia intermedia.

Phenology: Cones shed pollen in October and
November and are attended by the thrips

80

Austrobaileya 6(1): 67-94 (2001)

Fig. 9. Macrozamia longispina. Adult male plant in habitat. Forster PIF12137B & Machin. Photo: PI.Forster.

Cycadothrips chadwickii (Forster et al. 1994).
Ripe seed are dispersed in March and April.

Notes: Macrozamia longispina is similar to
M. macleayi, M. miquelii and M. mountperriensis
but differs from all these species by possessing
leaves with fewer, narrower leaflets and
markedly longer spines on the distal
sporophylls of both the male and female cones.
Macrozamia longispina is undoubtedly most
closely allied to M. mountperriensis and differs
in the leaves with fewer leaflets (usually 52-
58 versus 70-86), narrower median leaflets
(4.5-6 mm versus 6.5-9 mm) and in the apical
sporophylls of the male cones having much
longer spines (15-20 mm versus 3-13 mm).
The allozyme analysis also supports the close
relationship of these two species with the
outlying and sand-dwelling population of
M. longispina at State Forest 57 clustering more
closely with the serpentine populations of that
species, rather than the geographically nearby
populations of M. mountperriensis (Fig. 1).

Conservation status: Although of restricted
distribution the species is extremely abundant

where it occurs with large numbers present in
Wrattens State Forest and at one restricted
locality in St Mary State Forest. Populations
near Glastonbury appear to have been largely
eradicated by local landowners because of their
toxicity to domestic stock. The species is listed
as Rare on the schedules of the Queenslnad
Nature Conservation Act 1992.

Etymology: The specific epithet is derived from
the Latin words longi (longer) and spina (a
spine), alluding to the long apical spines on the
distal micro sporophylls.

4. Macrozamia macleayi Miq., Arch. Neerl.
Sci. Exact. Nat. 3(5): 250 (post June
1868). Type: Queensland. Moreton
District: Moreton Bay, s.coll., s.dat. (holo
U028258),/z<7c Forster (1999b).

Macrozamia cylindrica C.Moore, J. & Proc.
Roy. Soc. New South Wales 17: 119
(1884); M. spiralis var. cylindrica
(C.Moore) Maiden & Betche, Census
New South Wales PI. 9 (1916), nom. illeg.
non Regel (1876); M. tridentata subsp.

Jones el al. Macrozamia miquelii group

Fig. 10. Macrozamia longispina. Fruiting cone. Forster PIF19138 & Leiper. Photo: P.I.Forster.

cylindrica (C.Moore) J.Schust. in
A.Engler, Pflanzenr. 99(IV, I): 91
(1932). Type: New South Wales,
between the Upper Richmond and
Clarence River, 1861, CMoore (holo:
NSW).

Illustrations : Williams (1984:187); Stanley
& Ross (1989:451) [all asM. miquelii ].

Caudex usually subterranean, occasionally
emergent, erect, columnar or barrel-shaped, to
35 cm tall and 30 cm diam., unbranched. Young
leaves light green. Mature leaves elliptic-
lanceolate in outline, 0.5-2 m long, obliquely
erect to spreading, dark green, glossy, flat in

cross-section, arching in profile, 6-50 in a
moderately dense crown; expanded leaf base
8-12 cm x 2.5-3.5 cm, covered with grey-
brown, soft woolly hairs; petiole (including the
expanded base) 10-30 cm long, 0.8-1.2 cm
wide at the first leaflet, greenish, flat adaxially,
convex and angular abaxially; rhachis straight,
not twisted or slightly twisted, pale green, the
cross-section similar to that of the petiole;
leaflets 80-160, inserted at about 40° to the
rhachis, widely spreading, moderately crowded,
distal leaflets densely packed, proximal leaflets
becoming more widely spaced, narrowly linear-
lanceolate, 15-50 cm x 6-9 mm, flat, thin-
textured, hypostomatic, dark green and glossy

82

Austrobaileya 6(1): 67-94 (2001)

Fig. 11. Macrozamia macleayi. Adult male plant in habitat. Forster PIF12277B & Machin. Photo: P.I.Forster.

adaxially, slightly paler beneath, contracted
proximally to a whitish callous base, margins
entire, tapered to a pungent apex; lower leaflets
gradually reduced to 8-28 pairs of rigid
pinnacanths. Male cones 1-5, cylindrical, 15-
20 cm x 3.8-6.5 cm, straight or curved with
age, green; peduncle 15-29 cm x 1.5-2 cm,
elliptical to round in cross-section;
microsporophylls broadly cuneate, 1.5-2.5 cm
x 1-1.5 cm, with an erect, apical spine 0.5-1.5
cm long. Female cones 1 or 2, cylindrical to
barrel-shaped, 19-30 cm x 7-10 cm, green;
peduncle 15-41 cm x 2-3 cm, elliptical in
cross-section, furrowed; megasporophylls
broadly wedge-shaped, 1.5-2.5 cm x 3-3.5 cm,
with an erect apical spine 0.5-4.5 cm long, the
distal sporophylls with spines 2.5-4.5 cm long.
Seeds oblong to obovoid, 2.5-3.5 cm x 1.5-
2.5 cm, sarcotesta orange to red. Figs. 11-13.

Selected specimens examined : Queensland. Port
Curtis District: Mt Colosseum, Nov 1992, Forster
PIF12248 & Machin (BRI); Blackmans Gap, Nov 1992,
Forster PIF12277A & B & Machin (BRI); Koolkoorum
Creek Scientific Area 54, S.F. 121, Nov 1992, Forster
PIF 12285A & B & Machin (BRI); 8.5 km along road
to Bulburin forestry camp site, Nov 1992, Forster
PIF12289A & B & Machin (BRI); hill c. 2 km S of Mt
Colosseum, S of Miriam Vale, May 1992, Jones 9394,

Jones & Forster (BRI, CANB). Burnett District: Mt
Takilberan, Wanbar S.F., Aug 1995, Crane 1268 (BRI);
Burnett Range, S.F.54, Nov 1992, Forster PIF12299A
& B & Machin (BRI); Kalpowar to Gin Gin road, Sep
1993, Forster PIF13980A & B & Machin (BRI). Wide
Bay District: Burnett Range, 1.5 km NW of Mt Bania,
Sep 1993, Forster PIF13983 & Machin (BRI); Mt
Gaeta, c. 36 km N of Mt Perry township, Oct 1993,
Forster PIF14174 (BRI); S.F.256 Imbil, Mitchell L.A.,
Feb 1993, Forster PIF13146A & B & Machin (BRI).
Moreton District: Flaggy Creek tributary, 6 km E of Lake
Manchester, May 1992 ,Bostock 1348 (BRI); Scientific Area
2, S.F. 309, May 1992, Forster PIF9885B 8c Machin (BRI);
Brisbane Forest Park, Jan 1992, Forster PIF9355 & Machin
(BRI); ditto, May 1992, Forster PIF9886A & Machin
(BRI); 2 km NW of Mt Beerburrum, Jun 1994, Forster
PIF15239 & Machin (BRI); Jolly’s Lookout track to South
Boundary road, Brisbane Lorest Park, Nov 1991, Machin
AQ517082 (BRI); Brisbane Lorest Park, May 1993, Machin
(Jones 11536) (CANB); S.L. 309, Enoggera, Apr 1970,
Moriarty 120 (BRI, CANB); Brookfield, Dec 1888,
Simmonds AQ142052 (BRI); MtNebo, Dec 1960, Trapnell
AQ142053 (BRI).

Distribution and habitat : Macrozamia
macleayi (Map 1) is disjunctly distributed from
Mt Colosseum near Miriam Yale in the Port
Curtis District, south to the Brisbane region in
the Moreton District, with a reported disjunct
occurrence in northeastern New South Wales
based on the 1861 type collection of
M. cylindrica “between the Upper Richmond

Jones el al. Macrozamia miquelii group

83

Fig. 12. Macrozamia macleayi. Female cone. Imbil. Photo: D.L.Jones.

River and Clarence River”(Harden 1990; Hill
1998). There are no recent collections from
New South Wales and this record should be
regarded as extremely dubious. Plants grow at
altitudes from 100 to 500 m in the understorey
of araucarian microphyll vineforest or in open
forest that is often dominated by Eucalyptus
acmenoides, E. crebra, E. major, E. microcorys
and Lophostemon confertus.

Phenology: Cones shed pollen October and
November and are attended by the thrips
Cycadothrips chadwickii (Forster et al. 1994).
Ripe seeds are dispersed March and April.

Typification: See discussion in Forster (1999b).

Notes: This species is most closely related to,
and has previously been included under,
M. miquelii (cf. Johnson 1959; Hill 1998), but it
has a more southerly distribution, and can be
distinguished by its glossier, thinner-textured
leaflets and smaller female cones. The allozyme
analysis indicates that the northern populations
near Miriam Vale are more closely related to one
another than the disjunct population at Imbil
(Fig. 1). Isozyme analysis incorporating the
populations near Brisbane would be useful in
further elucidation of these relationships.

Conservation status: This species is abundant
and well conserved in Brisbane Forest Park

84

Austrobaileya 6(1): 67-94 (2001)

Fig. 13. Macrozamia macleayi. Male and female cones. Forster PIF12277A & B & Machin. Photo: P.I.Forster.

where it is present in both National Parks and
State Forests. The northern populations are well
represented in State Forests.

Etymology: Probably named for William John
Macleay (1820-1891), scientist and pastoralist
and a stalwart of the Linnean Society of New
South Wales (Australian Encylopaedia 1996).
Miquel (1868a,b) states “Nascur in Nova
Hollandia orientali, in regione fl. Moreton, ubi
probabititer detexit MacLeay”.

5. Macrozamia miquelii (F.Muell.) A.DC., Prodr.
16(2): 535 (1868).

Encephalartos miquelii F.Muell., Fragm. 3:
38 (1862); M. tridentata var. miquelii
(F.Muell.) J.Schust., in A.Engler,
Pflanzenr. 99(IV, I): 90 (1932);
M. tridentata var. oblongifolia Regel,
Trudy Imp. S.-Peterburgsk. Bot. Sada 4:
320 (1876), M. miquelii cited as
basionym; M. tridentata f. oblongifolia
(Regel) J.Schust., in A.Engler,
Pflanzenr. 99(IY, I): 93 (1932). type:
Queensland. Rockhampton, A.Thozet
(lecto MEL; isolecto: K, n.v.) designated

by Johnson (1959).

Macrozamia tridentata f. milkaui J.Schust.,
in A.Engler, Pflanzenr. 99(IV, I): 90
(1932). type: Queensland, around
Rockhampton, 11 May 1902, L.Diels
8249 (holo: B, destroyed).

Illustrations: Jones (1993: 248, upper
plate).

Caudex usually subterranean, occasionally
emergent, erect, columnar or barrel-shaped, to
50 cm long and 45 cm diam., unbranched.
Young leaves light green. Mature leaves
elliptic-lanceolate in outline, 0.5-2.3 m long,
obliquely erect to spreading, dark green, dull
to slightly glossy, flat in cross-section, arching
in profile, 20-80 in a dense crown; expanded
leaf base 8-15 cm x 3-4 cm, covered with grey-
brown, soft woolly hairs; petiole (including the
expanded base) 20-40 cm long, 8-20 mm wide
at the first leaflet, greenish, flat adaxially,
convex and angular abaxially; rhachis straight,
not twisted or slightly twisted, pale green, the
cross-section similar to that of the petiole;
leaflets 80-180, inserted at about 40° to the
rhachis, widely spreading, moderately crowded,

Jones et al. Macrozamia miquelii group

85

Fig. 14. Macrozamia miquelii. Adult male plant in habitat. Forster PIF12262B & Machin. Photo: P.I.Forster.

distal leaflets densely packed, proximal leaflets
becoming more widely spaced, linear, 15-50
cm x 6-12 mm, flat, thick-textured,
hypostomatic, dark green and dull to slightly
glossy adaxially, slightly paler beneath,
contracted proximally to a whitish or rarely
reddish callous base, margins entire, tapered to
a pungent apex; lower leaflets gradually
reduced to 6-16 pairs of rigid pinnacanths.
Male cones 1-5, cylindrical to fusiform, 12-
28 cm x 3.8-6.5 cm, straight or curved with
age, green; peduncle 11-30 cm x 1.5-2.5 cm,
elliptical to round in cross-section;
microsporophylls broadly cuneate, 1.2-2.5 cm
x 0.8-1.5 cm, with an erect, apical spine 0.2-2

cm long. Female cones 1-3, cylindrical to barrel¬
shaped, 25-40 cm x 10-15 cm, green; peduncle
10-30 cm x 2-3 cm, elliptical in cross-section,
furrowed; megasporophylls broadly wedge-
shaped, 1.5-3 cm x 2.5-4 cm, with an erect,
apical spine 0-4.5 cm long, the distal
sporophylls with spines 2-4.5 cm long. Seeds
oblong to ovoid, 2-3 cm x 1.5-2 cm, sarcotesta
yellowish, light orange or red. (Figs. 14-16).

Selected specimens : Queensland. Port Curtis District:
Western slopes of Mt Beserker, Beserker Range, Apr 1985,
ForsterV\¥\99\ (BRI); Mt Morgan, top of Razorback, Nov
1992, Forster PIF12251A-C (BRI); Mt Archer summit,
Nov 1992, Forster PIF12252A & B & Machin (BRI);
Mt Archer, 2 km from summit, Nov 1992, Forster
PIF12253A & B & Machin (BRI); Waterpark Creek,

86

Austrobaileya 6(1): 67-94 (2001)

Fig. 15. Macrozamia miquelii. Female cone. Byfield. Photo: D.L.Jones.

SE of Byfield township, Nov 1992, Forster
PIF12259A-C & Machin (BRI); 4 km NW of Byfield
township, Nov 1992, Forster PIF12262A & B &
Machin (BRI); northern end of Windmill Plains, Nov
1992, Forster PIF12263A & B & Machin (BRI); Ross
Range, 6.5 km along Keppel Sands road off Emu Park
road, Nov 1992, Forster PIF12268 & Machin (BRI);
Byfield N.P., The Peaks area, 8 km NE of Byfield, Sep
2000, Forster PIF26232 & Booth (BRI); c.7.4 km N
of Farnborough State School, beside road to Byfield,
May 1992, Jones 9393 & Jones (CANB).

Distribution and habitat: Macrozamia miquelii
(Map 1) is distributed in the Port Curtis District
from near Mt Larcom to north of Byfield. Plants
grow at altitudes between 10 and 540 m in open
forest or woodland dominated by eucalypts,
often on stony soil (e.g. trachyte at The Peaks),

but at Byfield they grow on deep sand. At some
localities the dominant canopy species is
Corymbia citriodora and Eucalyptus crebra,
whereas at others it is Corymbia intermedia
and Eucalyptus umbra.

Phenology: Cones shed pollen from October
to November and are attended by the thrips
Cycadothrips chadwickii (Forster et al. 1994).
Ripe seeds are dispersed in March and April.

Typification: When originally named, this
species had three collections from different
localities cited. Johnson (1959) lectotypified the
name using the Thozet collection from
Rockhampton, although Mueller had himself

Jones et al. Macrozamia miquelii group

87

Fig. 16. Macrozamia miquelii. Male and female cones. Forster PIF12263A & B & Machin. Photo: PI.Forster.

already indicated a preferred application (Forster
1999a). A proposal to conserve the name with
Johnson’s lectotype (Forster 1999a) has been
rejected by the IAPT committee for
nomenclature (Brummitt 2001) as it was
considered that Mueller’s later attempt at
restricting application of the name did not
equate to lectotypification.

Notes : Macrozamia miquelii is very closely
allied to M. macleayi (which has a more
southerly distribution), but can be distinguished
from that species by its duller, thicker-textured
leaflets and larger female cones. The close
relationship of these two species is reinforced
by the allozyme analysis (Fig. 1). The female
cones of M. miquelii are also very similar to
those of M. douglasii, but M. miquelii has 6-
16 pairs of prominent pinnacanths towards the
base of the leaf as opposed to 1-4 pairs in
M. douglasii. M. serpentina is also allied to
M. miquelii but is smaller growing with fewer,
shorter leaves, smaller leaflets, smaller male and
female cones and smaller seeds. Macrozamia
cardiacensis, M. longispina and M. mountperriensis
are all superficially similar to M. miquelii. See
under those species for differences.

Conservation status: Widespread and
abundant with a number of populations in
conservation reserves (e.g. Byfield National
Park, Mt Archer National Park).

Etymology: Named for F.A.W.Miquel (1811-
1871), Dutch botanist and early pioneer of
cycad systematic s.

6. Macrozamia mountperriensis F.M.Bailey,
Syn. Queensland FI., Suppl. 1: 50 (1886).
Macrozamia tridentata subsp.
mountperriensis (F.M.Bailey) J.Schust., in
A.Engler, Pflanzenr. 99(IV,I): 89
(1932).Type: Queensland. Adjacent to
Schuh Lookout, W of Mt Perry, 29 Aug.
1990, D.L. Jones 6340 & B.E. Jones (neo:
CANB; isoneo: BRI, NSW), fide Forster
& Jones (1992).

Illustrations : Bailey (1913: 516, 517);
Jones (1993: 250).

Caudex usually subterranean, occasionally
emergent, cylindrical to columnar, to 30 cm long
and 40 cm diam. unbranched. Young leaves light
green to yellowish green. Mature leaves elliptic-
lanceolate in outline, 0.6-1.5 m long, obliquely

Austrobaileya 6(1): 67-94 (2001)

Fig. 17. Macrozamia mountperriensis. Adult female plant in habitat. Forster PIF9343. Photo: P.I.Forster.

erect to spreading, light green to yellowish
green, flat in cross-section, arching in profile,
10-80 in a dense crown; expanded leaf base 6-
12 cm x 2-3.5 cm, covered with light grey, soft
woolly hairs; petiole (including the expanded
base) 30-65 cm long, 7-10 mm wide at the first
leaflet, pale green, ridged above, convex and
angular beneath; rhachis straight, not twisted
or with a slight twist, greenish white, the cross-
section similar to that of the petiole; leaflets
50-110, inserted at about 40° to the rhachis,
moderately crowded, evenly spaced throughout
except those towards the base which are more
widely spaced, linear, 20-35 cm x 6-9 mm, flat,
hypostomatic, thin-textured, light green to
yellowish, contracted proximally to a whitish
callous base, margins entire, tapered to a sharp
apex; lower leaflets slightly smaller but not
spine-like. Male cones 1-4, cylindrical, 12-25
cm x 2.5-4 cm, usually curved, green; peduncle
12-22 cm x 1-1.5 cm, elliptical in cross-section;
microsporophylls cuneate, 1-1.5 cm x 0.6-1.4
cm, with an erect, apical spine 0.2-1.2 cm long.
Female cones 1 or 2, cylindrical to barrel-shaped,
17-30 cm x 6-10 cm, green; peduncle 23^-0 cm
x 1.5-2.5 cm; megasporophylls broadly
cuneate, 1.5-3 cm x 2.5-3.5 cm, with an erect,

apical, spine 0.5-4 cm long, the distal
sporophylls with spines 2.5-4 cm long. Seeds
ovoid to oblong, 2-2.5 cmx 1.5-2 cm, sarcotesta
orange to red or yellow. (Figs. 17-18).

Selected specimens’. Queensland. Wide Bay District:
Stony Creek, 4 km E of Didcot, Dec 1984, Forster PIF1966
(BRI); Farrels Scrub, Deep Creek road, Oct 1991, Forster
PIF9128 (BRI, CANB, MEL); S.F.1294 Brooweena, Jan
1992, Forster PIF9343 & Machin (BRI, MEL); Fairlies
Knob N.P., Seaview Range, Dec 1992, Forster
PIF12571 (BRI); Stoney Range, S.F.38, Jun 1993,
Forster PIF13321 (BRI); Seaview Range, Doongul L.A.,
S.F. 1294, Jun 1993, Forster PIF13372 (BRI); S.F. 57
St Mary, Jun 1993, Forster PIF13374 & Machin (BRI);
7 km from Mt Perry on Monto road, Sep 1993, Forster
PIF13984 (BRI). Burnett District: 2 km SW of
Boolbunda Rock, May 1986, Forster PIF2423 (BRI);
Goodnight Scrub, Kalliwa Creek area, Dec 1986, Forster
PIF2771 (BRI); Goodnight Scrub, May 1991, Jones
9388, Jones & Forster (CANB).

Distribution and habitat : Macrozamia
mountperriensis (Map 1) is distributed from the
Mount Perry region to the west of Bundaberg,
south to Brooweena and east to Aramara. This
species grows at altitudes between 200 and
450m on sheltered slopes, ridges and gullies
under sparse tall dry sclerophyll open forest or
woodland in gravelly loams derived from granite

Jones et al. Macrozamia miquelii group

89

Fig. 18. Macrozamia mountperriensis. Female cone. Mt Perry neotype locality. Photo: D.L.Jones.

or granodiorite (rarely sandstone), and also in
Araucarian microphyll vine forests on red-brown
volcanic loams. Associated canopy species in
open forest and woodland include Angophora
leiocarpa, Corymbia citriodora,
C. trachyphloia, Eucalyptus acmenoides,
E. crebra and E. siderophloia.

Phenology: Pollen shedding occurs in October
and November. The cones are attended by the
thrips Cycadothrips chadwickii (Forster et al.
1994). Ripe seed is shed in March and April.

Notes: Macrozamia mountperriensis can be
distinguished from M. miquelii by its shorter
paler green leaves with proportionately longer

petioles, the absence of pinnacanths, much
smaller cones and smaller seeds. M. longispina
has more, narrower leaflets, smaller male cones
and the distal megasporophylls have longer
spines (see discussion under that species).

Conservation status: M. mountperriensis is
locally abundant in extensive colonies with
strong seedling recruitment. The species is
widespread and well represented in a number
of National Parks and several State Forests. It
is not rare or threatened at present.

Etymology: Named for the township of Mount
Perry where the species was first discovered.

90

Austrobaileya 6(1): 67-94 (2001)

Fig. 19. Macrozamia serpentina. Adult female plant in habitat. Forster PIF9408A & Machin. Photo: P.I.Forster.

7. Macrozamia serpentina D.L.Jones and
P.I.Forst. sp. nov. affinis M. miquelii
(F.Muell.) A.DC. sed statura minor foliis
paucioribus (5-12 adversum 20-80),
foliolis paucioribus (70-100 adversum
80-180), strobilis masculinis parvioribus
tenuioribus (2.5-3.5 cm diam. adversum
3.8-6.5 cm) et strobilis femineis
parvioribus tenuioribus (14-22 x 6.5-8
cm adversum 25-40 x 10-15 cm) spinis
distalibus brevioribus (1.5-2 cm
adversum 2-4.5 cm) differens. Typus:
Queensland. Port Curtis District: 1 km
W of Glen Geddes, 4 November 1992,
P.I.Forster PIF12266 & P.Machin (holo:

BRI; iso: CANB).

Macrozamia sp. (Marlborough P.I.Forster
+PIF12269A) (Forster 1997).

Caudex subterranean, ovoid, to 30 cm long, 15-
25 cm diam., unbranched. Young leaves light
green. Mature leaves broadly elliptic in outline,
0.2-1 m long, obliquely erect to spreading, dark
green, semi-glossy, flat in cross-section, arching
in profile, 5-12 in a sparse crown; expanded
leaf base 6-10 cm x 2-3 cm, covered with grey,
soft woolly hairs; petiole (including the
expanded base) 2-15 cm long, 6-12 mm wide at
the first leaflet, pale green, flat to shallowly
convex adaxially, angular abaxially; rhachis

Jones et al. Macrozamia miquelii group

Fig. 20. Macrozamia serpentina. Female cone. Glen Geddes. Photo: D.L Jones.

straight, not twisted or slightly twisted, pale
green, the cross-section similar to that of the
petiole; leaflets 70-100, inserted at 40-60° to
the rhachis, widely spreading, uncrowded, distal
leaflets densely packed, proximal
leaflets becoming more widely spaced, linear,
15-45 cm x 4-8 mm, flat, thick-textured,
hypostomatic, dark green and glossy adaxially,
slightly paler beneath, contracted proximally to
a white callous base, margins entire, tapered to
a sharp apex; lower leaflets gradually reduced
to 5-8 pairs of rigid pinnacanths. Male cones
1-3, cylindrical, 12-20 cm x2.5-3.5 cm, straight,
curved or slightly twisted with age, green;
peduncle 10-18 cm x 7-9 mm, round in cross-
section, furrowed; microsporophylls cuneate,

8-12 mm x 4-8 mm, with an erect, apical spine
0-7 mm long, the distal sporophylls with the
longest spines. Female cones 1 or 2, cylindrical
to barrel-shaped, 14-22 cm x 6.5- 8 cm, green;
peduncle 14-20 cm x 12-14 mm, round in cross-
section, furrowed; megasporophylls broadly
wedge-shaped, 1.5-2.5 cm x 3-3.5 cm, with an
erect, apical, spine 0.5-2 cm long, the distal
sporophylls with spines 1-2 cm long. Seeds
oblong to ovoid, 1.8-2.5 cm x 1.5-2 cm,
sarcotesta light orange to red. (Figs. 19-21).

Selected specimens examined: Queensland. Leichhardt
District: Marlborough - Glenprairie road, Jul 1998,
Thompson 1267 & Fox (BRI). Port Curtis District
Coorumburra Station, SSW of Marlborough, May 1998,
Batianoff 980533W (BRI); Mt Fairview, W of

92

Austrobaileya 6(1): 67-94 (2001)

Fig. 21. Macrozamia serpentina. Male cones. Forster PIF12266 & Machin. Photo: P.I.Forster.

Ridgelands, S.F.114, May 1998, Batianoff 980534W
(BRI); Glen Geddes, Jul 1987, Champion 295 (BRI);
Between Canoona and Glen Geddes, Dec 1966, Everist
7945 (BRI); Glen Geddes, Jan 1992, Forster PIF9408
(BRI, CANB); NNE of Marlborough, Nov 1992,
Forster PIF12269A & B & Machin (BRI); southern
slopes of Mt Slopeaway, Nov 1992, Forster PIF12273A
& B & Machin (BRI); E of Glenavon Homestead, Five
Mile Creek Headwaters, Mar 1994, Forster PIF15041
& Bean (BRI); Glen Geddes, 6 May 1992, Jones 9389
B & Jones (CANB); Glen Geddes, Oct 1991, Machin
AQ517058 (BRI); Ramilles Block, Jan 1989, Specht
146 & Reeves (BRI); Glen Geddes, Jan 1989, Specht
314 & Reeves (BRI).

Distribution and habitat: Macrozamia
serpentina (Map 1) occurs in the Leichhardt
and Port Curtis Districts between Marlborough
and Yaamba, north of Rockhampton. It grows
at altitudes between 80 and 160 m in low
woodland with a mixed grassy and shrubby
understorey in red clay loams over
serpentinites. Associated canopy species
include Corymbia xanthope and Eucalyptus
fibrosa.

Phenology. Cones shed pollen from October to
November and are attended by the thrips
Cycadothrips chadwickii (Forster et al. 1994).
Ripe seed are shed from March to May.

Notes: Macrozamia serpentina was not
mentioned in the account of Hill (1998),
although it had been listed as an undescribed
species by Forster (1994 & 1997). Macrozamia
serpentina is closely allied to M. miquelii but
smaller growing with fewer (5-12 versus 20-
80), shorter leaves, smaller and fewer (70-100
versus 80-180), more widely spaced leaflets,
smaller and thinner (2.5-3.5 cm diameter versus

3.8-6.5 cm) male cones, and smaller and thinner
(14-22 x 6.5-8 cm versus 25-40 x 10-15 cm)
female cones that have shorter apical spines on
the megasporophylls (1.5-2 cm versus 2-4.5
cm). Incomplete (not included in Fig. 1) allozyme
analysis of this species indicates that it is most
closely related to M. miquelii.

Conservation status: This species is listed as
Endangered on the schedules of the Queensland
Nature Conservation Act 1992. It is poorly
conserved with no populations in conservation
reserves. This species co-exists with a range of
restricted endemics on serpentinite (Batianoff
et al. 1991, 2000), in communities which are
themselves threatened by clearing, mining and
agriculture.

Etymology: The epithet is derived from the
Latin serpentinus, growing on serpentine rock.

Names of Uncertain Application

Macrozamia mackenzii Hort. Ex Mast., Gard.
Chron., n.s. 7: 665 (1877); M. tridentata
var. mackenzii (Hort. ex Mast.) J.Schust.,
inA.Engler, Pflanzenr. 99(IV,I): 90 (1932).
Type: ‘growing in the botanic garden at
Brisbane, under the charge of Mr W. Hill’.
No specimen is known to exist but an
illustration in the Gardeners Chronicle of
a cultivated plant places the species as a
member of the M. miquelii group. This
illustration is not diagnostic and is
inadequate for identification of the taxon
concerned.

Jones et al. Macrozamia miquelii group

93

Encephalartos spiralis var. major Miq.,
Verslagen Meded. Afd. Natuurk. Kon.
Akad. Wetensch. 15: 368 (1863). Type:
“Broad-Sound & Moreton Bays, F.Mueller;
Jervis Bay, N.S.W., F.Mueller. ; Moreton Bay,
C. Stuart.

We have not seen any of these syntypes. Given
the broad geographic origin of the different
collections it is certain that more than one
species is involved.

Acknowledgements

Thanks to Peter Machin for his enthusiastic and
prolonged interest and field assistance with this
project; Peter Bostock (BRI) for the map and
translation of the diagnosis into Latin; Corinna
Broers, Marion Garratt, Karina FitzGerald and
Barbara Jones for technical assistance at
CANB; Bob Chinnock (AD) and Laurie Jessup
(BRI) in their capacity as Australian Botanical
Liaison Officers for examining material at Royal
Botanic Gardens, Kew; Marco Duretto (MEL)
for rechecking specimens in MEL; Greg Smyrell
for determination of the northern limit of M.
miquelii’, Halina Winters (BRI) for library
research on John Douglas and Will Smith (BRI)
for scanning the illustrations.

References

The Australian Encyclopaedia. (1996). Yol. 5. 6 th ed.
Terry Hills: Australian Geographic Pty Ltd.

Bailey, F.M. (1883). Synopsis of the Queensland Flora.
Brisbane: Government Printer.

-(1886). Synopsis of the Queensland Flora,

Supplement 1. Brisbane: Government Printer.

-(1913). Comprehensive Catalogue of the

Queensland Flora. Brisbane: Government
Printer.

Ballard, F. (1936). Macrozamia douglasii W.Hill ex
F.M.Bailey. Hooker’s leones Plantarum 34:
3310, 3311.

Batianoff, GN., Reeves, R.D. & Specht, R.L.(1991). The
serpentinite flora of the humid subtropics of
eastern Australia. Proceedings of the Royal Society
of Queensland 101: 137-157.

Batianoff, GN., Neldner, Y.J. & Singh, S.(2000). Vascular
plant census and floristic analysis of serpentine
landscapes in central Queensland. Proceedings of
the Royal Society of Queensland 109: 1-30.

Brummitt, R.K. (2001). Report of the Committee for
Spermatophyta: 51. Taxon 50: 559-568

De Candolle, A. (1868). Prodromus Systematis
Universalis Regni Vegetabilis 16(2): 535. Paris:
V. Mas son.

Forster, PI. (1994). Gymnosperms. In R.J.F.Henderson
(ed.), Queensland Vascular Plants: Names &
Distribution, pp. 344-345. Indooroopilly:
Queensland Government.

-(1997). Spermatophyta - Gymnospermae. In

R.J.F.Henderson (ed.), Queensland Plants: Names
& Distribution, pp. 211-212. Indooroopilly:
Department of Environment.

-(1999a). Proposal to conserve the name

Encephalartos miquelii (Zamiaceae) with a
conserved type. Taxon 48: 569-570.

-(1999b). Typification and application of the

name Macrozamia macleayi Miq. (Zamiaceae).
Austrobaileya 5: 577.

Forster PI. & Jones D.L. (1992). Neotypification of
Macrozamia mountperriensis (Zamiaceae) with
notes on its distribution. Telopea 5: 289-290.

-(1998). Macrozamia cardiacensis sp.nov., M.

longispina sp. nov. In P.McCarthy (ed.), Flora of
Australia 48: 717. Melbourne: CSIRO
Publishing.

Forster PI. & Osborne R. (2001). Focus on Macrozamia
longispina P.I.Forst. & D.L.Jones (Zamiaceae,
section Parazamia). Encephalartos 66: 8-12,
15.

Forster, P.I., Machin, P, Mound, L. & Wilson G. (1994).
Insects associated with the reproductive
structures of cycads in Queensland and north¬
east New South Wales, Australia. Biotropica
26: 217-222.

Harden, G.J. (1990). Zamiaceae. In G.J.Harden (ed.).
Flora of New South Wales 1: 74-78.
Kensington: University of New South Wales
Press.

Hill, K.D. (1998). Cycadophyta. In P.McCarthy (ed.).
Flora of Australia 48: 597-661. Melbourne:
CSIRO Publishing.

Hill, W. (1879). Report on the Brisbane Botanic Garden.
Brisbane: Government Printer.

Johnson, L.A.S. (1959). The f a milies of cycads and the
Zamiaceae of Australia. Proceedings of the
Linnean Society of New South Wales 84: 64-
117.

Jones, D.L. (1993). Cycads of the World. Chatswood:
Reed Books.

Joyce, R.B. (1972). Douglas, John. [1828-1904], In
B.Nairn et al. (eds.), Australian Dictionary of
Biography. Vol. 4: 1851-1890, pp. 89-92.
Melbourne: Melbourne University Press.

Austrobaileya 6(1): 67-94 (2001)

94

Miquel, F.A.W. (1868a June). Nouveaux materiaux pour
servira la connaissance des Cycadees. Archives
Neerlandaises des Sciences Exactes et
Naturelles 3(5): 193-254.

-(1868b October). Nouveaux materiaux pour

servira la connaissance des Cycadees.
Adansonia 9: 29-73.

Mound, L.A. & Terry, I. (2000). Thrips pollination of
the central Australian cycad, Macrozamia
macdonnellii (Cycadales). International Journal
of Plant Science 162: 147-154.

Mueller, F. (1862). Fragmenta Phytographie
Australiae 3: 38. Melbourne: Government
Printer.

-(1883). Remarks on an undescribed

Encephalartos from Queensland. The Chemist
& Druggist (Australasia) Supplement 5: 80-
81.

Nei, M. (1978). Estimation of average heterozygosity
and genetic distance from a small number of
individuals. Genetics 89: 583-590.

Sharma, I.K., Jones, D.L., Forster, P.I. & Young, A.G.
(1999). Low isozymic differentiation among
five species of the Macrozamia heteromera
group (Zamiaceae). Biochemical Systematics
and Ecology 27: 67-77.

Stanley, T.D. & Ross, E.M.(1989). Flora of south¬
eastern Queensland. Volume 3. Brisbane:
Queensland Department of Primary Industries.

Williams, K.A.W.(1984). Native Plants of Queensland.
2: 187. Ipswich: K.A.W.Williams.

Oreodendron C. T. White reduced to Phaleria Jack
(Thymelaeaceae, Thymelaeoideae)

B.E.Herber

Summary

Herber, B.E. (2001). Oreodendron C.T. White reduced to Phaleria Jack (Thymelaeaceae,
Thymelaeoideae). Austrobaileya 6 (1): 95-97. The monospecific genus Oreodendron , endemic in
Queensland, has been distinguished from the related genus Phaleria by the absence of involucral bracts,
the presence of a short pedicel and the arrangement and exposition of stamens. A study of these characters
in Oreodendron shows that involucral bracts are present before anthesis. Furthermore, the presence of a
short pedicel as well as the arrangement and exposition of stamens, said to be characteristic of Oreodendron ,
also occur in Phaleria. Therefore, Oreodendron is reduced here to synonymy of Phaleria and the new
combination Phaleria biflora (C. T. White) Herber is made for White’s species. Akey to identify the four
Australian species of Phaleria is provided.

Key words: Thymelaeaceae, Oreodendron, Phaleria

B.E. Herber, Institut fur Allgemeine Botanik Ohnhorststrasse 18, 22609 Hamburg, Germany;
email: Bjoemherber@web.de

Introduction

The genus Oreodendron C. T. White with one
species, O. biflorum C. T. White was established
in 1933 based on a collection of S. F. Kajewski
from Thornton Peak (“Mount Alexander”),
Queensland, Australia. White (1933) stated that
his new genus is “closely allied” to Phaleria
Jack, a genus of about 30 species distributed
from Malesia to Sri Lanka, Micronesia, the
Samoan Islands, Tonga and Australia. Till now
the Australian species recognised are
P. clerodendron (F. Muell.) Benth., P. octandra
(L.) Baill. and P. chermsideana (Bailey) C. T.
White. Phaleria is mainly characterized by its
bilocular ovary. In subfamily Thymelaeoideae,
the gynoecium is typically unilocular, only in
tribe Phalerieae, comprising Oreodendron,
Phaleria and the African genus Peddiea Harv.,
are there bilocular ovaries. In addition to
Oreodendron and Phaleria sharing this
character, they also share a non-articulated
floral tube, a diplostemonous androecium,
terminal insertion of the style and the presence
of a floral disk. White (1933: 74-75) claimed
that Oreodendron differs from Phaleria in
“...the absence of involucral bracts at the top
of the peduncle, in the flowers being pedicellate

not sessile and in the anthers being arranged in
two very distinct series, the lower series being
included”. The genus Oreodendron is widely
recognized (e.g. Hutchinson 1967, Takhtajan
1997). On the other hand, Domke (1934), who
was not able to examine any material of
Oreodendron, accepted the genus only with
reservation. In her contribution for the Flora of
Australia, Rye & Heads (1990: 130) remarked
“[t]his genus is very closely related to Phaleria
and needs further study to determine whether
or not it should be retained as a separate genus”
and of O. biflorum that it “[s]omewhat
resembles Phaleria chermsideana ”.

Results and Discussion

Absence of involucral bracts.- The capitate
inflorescences of Phaleria are provided with
two to more bracts in the distal part of the
peduncle, which are often deciduous before or
at anthesis. In Oreodendron biflorum, two
involucral bracts are developed (Fig.lA,B).
These two bracts are minute and deciduous
before anthesis. On peduncles of open flowers,
it is almost impossible to detect their scars and
they are easily overlooked.

Accepted for publication 2 March 2001

96

Austrobaileya 6(1): 95-97 (2001)

Presence of a pedicel - The pedicel of
Oreodendron biflorum is about 0.6 mm in
length (Fig. 1 A,B). Short pedicels are common
in Phaleria too. In all Australian species
pedicels up to 0.3 mm length sometimes occur
(see Rye & Heads 1990). Neither Ding Hou
(1960) nor Smith (1981) mentioned the
presence of a pedicel in Phaleria, which may
be due to its length of less than 1 mm. In such
cases a distinction between sessile, sub-sessile
or pedicellate flowers is questionable.

Arrangement of stamens - The stamens
in Oreodendron are arranged in two clearly
separate series as described by White (1933).
The same basic arrangement is found in
Phaleria, where the stamens are usually
inserted in two distinct series on the upper half
of the floral tube as well.

Exposition of stamens.- In Oreodendron
the anthers of the upper whorl are slightly
exserted, whereas those of the lower whorl are
included in the floral tube (Fig. 1C,D). The
position of anthers in Phaleria is sometimes
as in Oreodendron (e. g. as in Phaleria
disperma (Forst. f.) Baill.), or all anthers are
either exserted or rarely included. Heterostyly
is common in Phaleria and occurs in two of
the Australian species, P. clerodendron and
P chermsideana. It is unlikely but cannot be
excluded that Oreodendron is heterostylic too.
Nevertheless, neither the occurrence of
heterostyly nor the arrangement or exposition
of stamens provides characters to discriminate
between Oreodendron and Phaleria.

Not a single character that would justify
the generic distinction of Oreodendron has been
confirmed. The arguments used by White
(1933) for treating Oreodendron as a genus
distinct from Phaleria Jack have proved to be
based on incomplete observations or are
irrelevant for the generic delimitation in
question. Therefore, Oreodendron is reduced
to a synonym under Phaleria and the following
combination is made:

Phaleria Jack, Malayan Misc. 2: 59 (1822).
Type: P. capitata Jack

Oreodendron C. T. White, Contrib. Arnold
Arbor. 4: 74 (1933), synon. nov. Type:
O.biflorum C. T White

Phaleria biflora (C. T. White) Herber, comb,
nov.

Oreodendron biflorum C. T. White, Contrib.
Arnold Arbor. 4: 74-75 (1933). Type:
Queensland. Cook District: Thornton
Peak, 18 Dec 1929, S. F. Kajewski 1499
(iso: BRI, B) Queensland

Selected specimens: Queensland. Cook District:
Daintree NP, Black Mtn area, Daintree River headwaters,
May 1998, Forster el al. PIF22957 (BRI); Thornton Peak,
Dec 1946, Flecker 7093 (BRI); North Mary L.A., SF 143,
Jul 1994, Forster et al. PIF 15627 (BRI).

Phaleria now includes four species in Australia:
P. clerodendron (F. Muell.) Benth., P. octandra
(L.) Baill., P. chermsideana (Bailey) C. T. White
and P biflora (C. T. White) Herber. Most similar
to P. biflora is P. chermsideana, which can be
distinguished as outlined in the key below.

Key to Australian species of Phaleria

1. Leaf blade (l-)4-7(-9) cm long; involucral bracts 2.2

Leaf blade (8-)13-17(-24) cm long; involucral bracts 4.3

2. Leaf blade ovate; inflorescence 2 (rarely 3)-flowered; septum of fruit thinner

than seed-coat. P. biflora

Leaf blade elliptic. Inflorescence 4-10-flowered. Septum of fruit thicker
than seed-coat. P. chermsideana

3. Inflorescence 5-7-flowered; floral tube at anthesis 25 mm long or more;

fruit when mature >23 mm long. P. clerodendron

Inflorescence 8-25-flowered; floral tube at anthesis up to 17 mm long; fruit
when mature < 20 mm long. P. octandra

Herber, Oreodendron reduced to Phaleria

97

Fig. 1 . Phaleria biflora. A, inflorescence at early stage of
anthesis; B, inflorescence before anthesis; C, D longitudinal
sections of a young flower. A-D, Kajewski 1499 (B). Scale
bar: A-D = 1 mm.

Specimens of other species examined:

Phaleria chermsideana : Queensland. Wide bay District:
Norval Park c. 13km N of Yandaran, NW of Bundaberg,
Oct 1996, Bean 11149 (BRI); Norville Park, N of
Bundaberg, [no date], Randall 629 (BRI); 20miles (32km)
NW of Bundaberg, Dec 1982, Sarnadsky s.n. [AQ348777]
(BRI). Darling Downs District: Moss Gardens, border
fence, Jan 1990, Forster et al. PIF6217 (BRI); The Head
near Killamey, Feb 1968, Jones 3659 (BRI); Norville Park,
32km NW of Bundaberg, Jul 1983, Sarnadsky s.n.
[AQ339630](BRI). Moreton District: O’Reilly’s Guest
House, Nov 1969, Hockings 2 [AQ363220] (BRI); Mt
Mistake, Oct 1969, Smith s.n. [AQ410839] (BRI); Mt
Mistake, May 1948, Smith & Webb 3692 (BRI). New South
Wales: Near Kyogle, Oct 1943, Jones s.n. [AQ
85756] (BRI).

Phaleria clerodendron: Queensland Cook District:
Henrietta Creek on Palmerston Highway, Nov, 1979,
Clarkson 2737 (BRI); Cedar Creek between Bingil Bay
and ElArish, Dec 1984, Jessup 740 (BRI); Russell River
near bridge on Bruce highway, Jul 1963, Jones s.n.
[AQ85766] (BRI).

Phaleria disperma : A. Whistler 6600 (HBG), 4453 (B).
Phaleria octandra : Northern Territory: Darwin and Gulf
District: Water Quarry rainforest, Melville Island, Dec
1984, Jones 1701 (BRI); 2km NW of Yirrkala, Feb 1988,
Russell-SmithAlQn (BRI), North central Arnhem land 19km
N of Mirmgatja, Nov 1987, Russell-Smith 3980 (BRI);
Yirrkala, Aug 1948, Specht 806 (BRI); Banjo jungle, Snake
Bay, Melville Island, near airstrip, May 1978, Webb &
Tracey 12486 (BRI). Queensland: Cook District: Yam
Is, forest adjacent to airfield, Oct 1981, Clarkson 4025
(BRI); Shiptons Flat, Nov 1989, Jessup et al. GJD2830
(BRI); Cape York, Bamaga, Sep 1963, Jones 2554 (BRI);
SFR 191 Wongabel, near Atherton, Jan 1963, Rudder
AF02569 (BRI). South Kennedy District: Netherdale,

Mirani Shire, Apr 1980, McConnell 2013 (BRI); Mt
Mandurana NP 602, c 20km WNW of Mackay, May 1990,
McDonald 4581 (BRI); Mueller s.n. (P).

Acknowledgements

I extend my thanks to K. Kubitzki for helpful
co mm ents and to C. Bayer for the instructive
discussion and for reviewing the manuscript.
Furthermore, I wish to thank the curators of
BRI, B and P for the loan of material in their
care, and acknowledge the support of Deutsche
Forschungsgemeinschaft (Ku 174/15-1).

References

Ding Hou (1960). Thymelaeaceae. In Flora Malesiana I,
6:1-48. Djakarta: Noordhoff - Kolff N.V.

Domke, W. (1934). Untersuchungen liber die geographische
und systematische Gliederung der Thymelaeaceae.
Biblioth. Bot. Ill: 1-151.

Hutchinson, J. (1967). The genera of flowering plants
(Angiospermae) 2. Oxford: Clarendon Press.

Rye, B. L. & Heads, M. J. (1990). Thymelaeaceae. In A. S.
George (ed.), Flora of Australia 18: 122-215.
Canberra: Australian Government Publishing
Service.

Smith, A. C. (1981). Thymelaeaceae. In Flora Vitiensis Nova
2: 580-592. Honolulu, Hawaii: SB Printers, Inc.

Takhtajan, A. (1997). Diversity and classification of
flowering plants. New York: Columbia
University Press.

White, C. T. (1933). Ligneous plants of north Queensland.
Contr. Arnold. Arbor. 4: 74-75.

A new species of Lissanthe R.Br.
(Epacridaceae) from Queensland

A.R. Bean

Summary

Bean, A.R. (2001). A new species of Lissanthe R.Br. (Epacridaceae) from Queensland. Austrobaileya
6(1): 99-102. Lissanthe brevistyla, a new species from the serpentinite deposits near Rockhampton, is
described and illustrated. It is compared to related taxa, and notes on the conservation status are provided.

Key words: Lissanthe brevistyla, Lissanthe, Epacridaceae, sepentinite, Queensland flora, taxonomy.

A.R. Bean, Queensland Herbarium, Environmenal Protection Agency, Brisbane Botanic Gardens Mt
Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Introduction

The serpentinite geological deposits near
Marlborough, north of Rockhampton contain
a high proportion of rare or endemic plant
species, some of which are very restricted in
distribution (Batianoff etal. 2000). The species
under consideration here came to notice only
recently, and only after a mining lease had been
established over the area of its occurrence.

A revision of the whole of Epacridaceae
is being undertaken at the National Herbarium
of New South Wales by Elizabeth Brown and
colleagues. Since the results of this revision are
some years away, the opportunity is taken here
to describe this rare Queensland epacrid.

The new species shares with Lissanthe
R.Br. the following features: spicate
inflorescence on terminal growth (not on old
wood), with flowers subtended by a solitary
persistent bract and two bracteoles; the
bracteoles somewhat removed from the sepals;
the inner surface of the corolla tube glabrous
in the lower half, but hairy towards the throat;
the corolla lobes valvate in bud; and the 5-7-
locular drupaceous fruits.

The generic placement is not certain, as
this taxon belongs to a group of closely related
genera that include Lissanthe, Cyathodes s. lat.,
Acrotriche and some Leucopogon spp. (Brown
and Crayn, pers. comm.). The pedicel lacks the
degree of elongation usually associated with
species of the genus Lissanthe, and the filaments

Accepted for publication 20 April 2001

are unusually wide and thick (Fig. lc, Id).

The only other Queensland taxon
currently ascribed to Lissanthe is L. strigosa
subsp. subulata (R.Br.) J.M.Powell (Powell &
Wiecek 1994). However, Powell (1992)
foreshadowed the transfer of some species of
Leucopogon R.Br. to Lissanthe, namely
Leucopogon pedicellatus and L. pleiospermus.
Determinavit slips at BRI annotated by Powell
indicate a similar position for Leucopogon
pluriloculatus. While these species have the
densely hairy corolla lobes, traditionally
indicative of Leucopogon, they otherwise
conform to the characteristics of Lissanthe.

Taxonomy

Lissanthe brevistyla A.R.Bean sp. nov. affinis
L. strigosae autem inflorescentia
terminali spicata, pedicellis brevioribus,
stylo ovarioque glabro, fructibus
grandioribus purpureocyaneis differt.
Typus: Queensland. Port Curtis
District: Gumigil Mining Lease, 16.5 km
and 189° from Marlborough Motel on
Bruce Highway, 17 November 1999, 1.G.
Champion 1569 & B. Tangey (holo: BRI;
iso: NSW).

Spreading shrub to 1.5 m high and 2.5 m across.
Branchlets puberulous. Leaves lanceolate to
narrowly lanceolate, 10-25 x 1.8-3 mm,
glabrous, dark green and flat to convex above,
whitish below, usually 7-veined, tapering into
a pungent point 0.8-1.3 mm long, margins

100

Austrobaileya 6(1): 99-102 (2001)

Fig.l. Lissanthe brevistyla. A. flowering branchlet x 0.8. B. side view of flower showing the persistent bract (each subtending
a flower), one (of two) bracteoles, sepals and corolla tube x 16. C. internal view of flower x 16. D. lateral view of stamen
x 32. E. leaf, abaxial surface x 4. F. mature fruit x 4. G. transverse section of endocarp of fruit x 8. A, Champion 1570 &
Tangey, B-E, Champion 1554 & Tangey, F, G, Champion 1565 & Tangey (all BRI). Del. W. Smith.

Bean, Lissanthe

101

entire, 7-9 vascular bundles per leaf; petiole
0.8-1.5 mm long. Inflorescence a dense
auxotelic spike, up to 25 mm long, comprising
18-26 bisexual flowers, in the axils of the
uppermost leaves or apparently terminal.
Rachis puberulous. Pedicels obscure.
Subtending bract orbicular, c. 1 x 1 mm,
sparsely puberulous to glabrous, persistent.
Bracteoles 2, persistent, 0.7-1.1 mm long,
keeled, obtuse, ciliolate, placed just below but
not or scarcely overlapping the sepals. Sepals
5, imbricate, broadly ovate, 0.5-1 mm long,
0.5-0.8 mm wide, apex obtuse, margins
ciliolate. Corolla white or pink; tube urceolate,
1.8-2.6 mm long, far exceeding sepals,
glabrous externally, with long, flexuose, very
sparse to moderately dense hairs internally
towards distal end; lobes 5, valvate in bud,
triangular, 0.7-0.9 mm long, acute, spreading
or recurved, glabrous externally, with long,
flexuose, sparse hairs on inner surface. Stamens
5, alternating with corolla lobes. Filaments 0.4—
0.5 mm long, 0.25-0.3 mm wide and thick,
hairy at base. Anthers free, brown, 0.4-0.6 mm
long, slightly exserted from corolla tube,
dorsifixed, attached above their middle onto the
filaments. Ovary glabrous, 6 or7-locular, each
locule containing 1 ovule. Style terminal on the
ovary summit, terete, 0.5-0.7 mm long, much
shorter than corolla tube; stigma small. Mature
fruits globose to depressed-globose, 4-6 mm
long, 4-6.5 mm diameter, smooth (when fresh),
style persistent, pedicel 1-1.5 mm long;
pericarp glabrous, purple-blue in colour;
mesocarp fleshy; endocarp woody, ribbed, not
separating into pyrenes. Fig. 1.

Specimens examined: Queensland. Port Curtis District:
SF 114, site 4, Mt Fairview, 65 km NW of Rockhampton,
Dec 1998, Batianoff 981222 et al. (BRI); Poachers Gully,
Gumigil Mining Lease, 16.2 km and 185° from
Marlborough Motel on Bruce Highway, Nov 1999,
Champion 1544 & Tangey (BRI); ‘Magpie’, 17.6 km and
171° from Marlborough Motel on Bruce Highway, Nov
1999, Champion 1554 & Tangey (BRI, MEL); ‘Currawong’
on Gumigil’s Ramilies Lease, 19.4 km and 167° from
Marlborough Motel on Bruce Highway, Nov 1999,
Champion 1557 & Tangey (BRI, CANB); Gumigil Mining
Lease, 16.5 km and 189° from Marlborough Motel on Bruce
Highway, Nov 1999, Champion 1570 & Tangey (BRI);
Marlborough Ck, 25 km SW of Marlborough, Nov 1997,
McCabe s.n. (BRI).

Distribution and habitat: Endemic to the
Marlborough area of central coastal
Queensland. It is confined to steep hillsides in
woodland dominated by Eucalyptus fibrosa
F.Muell. and E. xanthope Brooker & A.R.Bean,
and may be associated with other shrub species
such as Hakea trineura F.Muell., Pimelea
leptospermoides F.Muell. and Leucopogon
cuspidatus R.Br. It grows in red gravelly soil
or on stony scree, derived from ultramafic
rocks.

Phenology: Flowers and fruits are recorded for
November.

Affinities: L. brevistyla differs fromL. strigosa
ssp. subulata by its almost sessile flowers,
shorter style (relative to corolla tube length),
the larger purple-blue fruits (small and white
for subulata ), and glabrous style and ovary
(densely hairy for subulata ).

Of the species currently under
Leucopogon, L. brevistyla most resembles
Leucopogon pedicellatus, as they both have a
glabrous style and ovary, and a style shorter
than the corolla tube. However, L. pedicellatus
has longer pedicels, and much longer and more
pilose corolla lobes.

Conservation status: Applying the criteria of
the IUCN (Anon. 1994), a category of
‘vulnerable’ is proposed (Criteria B1, B2 (b, c)).

Notes: This species is undoubtedly
lignotuberous, as it grows in a very fire-prone
area. It may spread vegetatively (by rhizomes),
as dense clumps covering more than 10 m 2 have
been observed (I. Champion pers. comm.).

Etymology: From the Latin brevi- short and
stylus- style, referring to the style which is
shorter than in related species.

Acknowledgements

Thanks are due to Irene Champion for making
special collections of this species, to Will Smith
for the illustrations, and Les Pedley for the Latin
diagnosis.

102

Austrobaileya 6(1): 99-102 (2001)

References

Anonymous (1994). IUCN Red List Categories.
International Union for Conservation of Nature
and Natural Resources: Switzerland.

Batianoff, G.N., Neldner, VJ. & Singh, S. (2000). Vascular
Plant Census and Floristic Analysis of Serpentine
Landscapes in Central Queensland. Proceedings
of the Royal Society of Queensland 109: 1-30.

Powell, J.M. (1992). Lissanthe, in GW.Harden (ed.), Flora
of New South Wales 3: 417-9. Sydney: New
South Wales University Press.

Powell, J.M. & Wiecek, B.M. (1994). Morphological
variation in Lissanthe strigosa (Epacridaceae) in
New South Wales. Telopea 5(4): 735—41.

Hydnophytum ferrugineum (Rubiaceae: Hydnophytinae),
a new species of ant-plant
from Cape York Peninsula, Queensland

Paul I. Forster

Summary

Forster, P.I. (2001). Hydnophytum ferrugineum (Rubiaceae: Hydnophytinae), a new species of
ant-plant from Cape York Peninsula, Queensland. Austrobaileya 6 (1): 103-106. A new species
of ant-plant Hydnophytum ferrugineum P.I.Forst. is described and illustrated. Notes on the
distribution, habitat and conservation status are provided for the new species which is restricted to
upland rainforests on granite in the Mcllwraith Range, Cape York Peninsula where it is known
from a small area. A conservation status of Vulnerable is recommended. A key to the Australian
species of Hydnophytinae is provided.

Keywords: Hydnophytum - Australia; Hydnophytum ferrugineum', Hydnophytum moseleyanum',
Myrmecodia', ant-plants.

P.I. Forster, Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens Mt
Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Introduction

The tribe Hydnophytinae Huxley & Jebb of the
Rubiaceae comprises the genera Anthorrhiza
Huxley & Jebb, Hydnophytum Jack,
Myrmecodia Jack, Myrmephytum Becc. and
Squamellaria Becc. Apart from Hydnophytum,
the other genera have been revised in a series
of papers by C.R.Huxley and M.H.P.Jebb
(Huxley & Jebb 1991, 1993; Jebb 1991). The
genus Hydnophytum Jack comprises c. 50
species and although a monograph has been
promised (e.g. Huxley 1993) it has yet to
materialise. A single species of Hydnophytum
has been long recognised to occur in Australia
and was initially identified as H. formicarium
Becc. by Bailey (1900). Huxley (1982)
referred this species to H. papuanum Becc., but
later annotated herbarium material as H.
moseleyanum Becc. the name by which it is
now known in Queensland (Reynolds &
Halford 1997).

The presence of a second species of
Hydnophytum in Australia was alluded to firstly
by Huxley (1982) as “ Hydnophytum sp. 1”
based on a collection by the redoubtable
L.J.Brass (#19885, but incorrectly as 1885) at
Leo Creek in the Mcllwraith Range on Cape
York Peninsula and secondly by D.L.Jones in
Elliot & Jones (1990) as “ Hydnophytum

Accepted for publication 27 June 2001

species. An apparently undescribed species
which occurs in the higher ranges of Cape York
Peninsula”. The Brass collection as held at
BRI is sterile and this may have precluded
Huxley from describing it. It has now been
possible to collect fertile material of this entity
enabling a description to be made and a name
formally published.

Taxonomy

Hydnophytum ferrugineum P.I.Forst., sp.
nov. Ab Hydnophyto moseleyano bracteis
inflorescentiae trichomatibus ferrugineis
densis usque ad 2 mm longis obsitis, foliis
valde discoloribus superne atroviridibus
nitidis (adversum folia plus minus
concoloria superne pallide viridia
impolita), venis lateralibus 5 vel 6 in
costae latere utroque (adversum venas
laterales 3-4 in costae latere utroque)
differt. Typus: Queensland. Cook
District: Timber Reserve 14, Leo Creek,
9 July 1997, P.I.Forster PIF21406 et al.
(holo: BRI [3 sheets + spirit]; iso: K,
MEL).

Epiphytic subshrub with succulent tuber and
stems. Tuber horizontal to pendent, globose-
spherical to variously misshapen, somewhat
mounded, unridged, up to 30 cm diameter,

104

Austrobaileya 6(1): 103-106 (2001)

spineless, glabrous, surface often with entrance
holes for ants over the entire surface, cream-
silver; chambers complex and forming a
honeycomb of chambers. Stems solitary, rarely
2 or 3, up to 60 cm long, much branched, up to
10 mm diameter, spineless, glabrous, clypeoli
absent. Interpetiolar stipules triangular-
lanceolate, 3-3.5 mm long, 2-3.5 mm wide,
caducous, glabrous. Leaves mesomorphic;
petiole 2-6 mm long, 1.5-2 mm wide, generally
as broad as long, channelled above, glabrous;
lamina elliptic to elliptic-ovate, 10-60 mm long,
7-35 mm wide, strongly discolorous, upper
surface dark glossy green and venation obscure,
lower surface pale matt-green and with 5 or 6
indistinct lateral veins; tip obtuse to rounded;
base cuneate. Inflorescence sessile, laterally
displaced to the leaf axil, surrounded by dense
bracts, each bract with dense ferruginous
trichomes to 2 mm long. Flowers 4-merous.
Corolla 4-5 mm long, c. 4 mm diameter, white;
tube 2-3.5 mm long, 1.8-2 mm diameter,
externally glabrous, internally with dense
trichomes in top of throat forming a connivent
mass around anthers; lobes lanceolate-ovate,
fleshy and cucullate, upwardly apiculate, c. 2
mm long, 1.2-1.5 mm wide; anthers inserted,
1-1.1 mm long and c. 0.5 mm wide; stigma 2-
lobed, c. 2.5 mm long. Fruit a fleshy oblong
drupe, c. 6 mm long, white. Fig. 1.

Additional specimen examined: Queensland. Cook
D isTRict: Leo Creek, Upper Nesbit River, Aug 1948,
Brass 19885 (BRI).

Distribution & habitat: Hydnophytum
ferrugineum is thus far known only from the
headwaters of Leo Creek in the Mcllwraith
Range on Cape York Peninsula. Plants grow
as epiphytes in the canopy of complex
notophyll vineforest on granite substrates at
altitudes above 500 m.

Notes: This species is clearly distinct because
of the strongly discolorous, glossy foliage and
the ferruginous trichomes that are copiously
distributed in the inflorescences. When
compared to H. moseleyanum it differs in the
inflorescence bracts beset with dense
ferruginous trichomes up to 2 mm long (versus
sparse uncoloured trichomes less than 0.5 mm
long) and the leaves being strongly discolorous
and dark glossy-green above and with 5 or 6
lateral veins per side of midrib (versus more or
less concolorous and pale matt-green above and
with 3 or 4 lateral veins per side of midrib).
None of the other New Guinea species
represented in the BRI collections appears
closely similar.

Although the genera Hydnophytum and
Myrmecodia are readily distinguishable, it is
felt opportune to provide a key to all of the
Australian species of Hydnophytinae. There
is now increased horticultural interest in these
plants worldwide (e.g. Plummer 2000) and aids
to their accurate identification are required.

Key to the Australian species of Hydnophytinae

1. Tuber and stem with spines, especially around ant exit-holes and

inflorescences; stems thick, branching sparse or absent; inflorescences

sunken.2

Tuber and stems spineless; stems slender and freely branching;
inflorescences pedunculate to sessile. 4

2. Stems several from tuber; leaf lamina fleshy-succulent; fruit

white. Myrmecodia beccarii Hook.f.

Stems generally single from tuber, rarely with side branches; leaf

lamina coriaceous to leathery; fruit orange-red to pink.3

3. Stems with spiny, shield-like outgrowths (clypeoli) at the base of each leaf

.Myrmecodia platytyrea Becc.

Stems without clypeoli, but with elongated depressions filled with bracts

and hairs at the base of each leaf.Myrmecodia tuberosa Jack

Forester, Hydnophytum ferrugineum

105

Fig. 1. Hydnophytum ferrugineum. A. whole plant, x 0.3. B. undersurface of leaf showing venation pattern, x 2. C. close-
up of tuber surface showing ant exit-holes, x 1. D. node with axillary inflorescence and emerging flower, x 3. E. side view
of flower, x 8. F. dissection of corolla showing anthers and hairs in throat, x 8. G. seedling, x 2. All from Forster 21406
(BRI). Del. W. Smith.

106

Austrobaileya 6(1): 103-106 (2001)

4. Leaf lamina with 5 or 6 lateral nerves per side of midrib, strongly

discolorous, dark green and glossy above; inflorescence with copious

ferruginous indumentum.Hydnophytum ferrugineum P.I.Forst.

Leaf lamina with 3 or 4 lateral nerves per side of midrib, more or less
concolorous, pale green and matt above; inflorescences with scattered
uncoloured indumentum.Hydnophytum moseleyanum Becc.

Conservation status: H. ferrugineum is locally
common but appears to be very restricted in its
distribution. Apart from damage by occasional
cyclones there are no apparent natural threats.
This species is however potentially under threat
by hobbyist collectors of ant-plants. The Leo
Creek locality is well-known for a number of
endemic or rarely encountered Orchidaceae
taxa and has been exploited by collectors over
the last 30 years. Orchid collectors often also
collect ant-plants and so the formal description
of this species will undoubtedly arouse further
interest in the locality. Large, mature ant-plants
are very susceptible to damage and their
removal is rarely followed by successful re¬
establishment. It is possible to grow ant-plants
from seed and this is the best way to establish
an ex-situ population of this species.
Suggestions for the culture and propagation of
H. ferrugineum may be found in D.L. Jones in
Elliot & Jones (1990) or adapted from Forster
( 2000 ).

Applying IUCN guidelines (Anon.
1994), a category of ‘vulnerable’ is proposed
(Criteria C2, Dl, D2).

Etymology: The specific epithet is based on
the Latin ferrugineus and alludes to the colour
of the dense hairs present on the inflorescences
of this species.

Acknowledgements

W.Smith (BRI) provided the figure.
RD.Bostock (BRI) translated the diagnosis into
Latin and commented on the manuscript.
Assistance with the difficult fieldwork

undertaken to collect this plant was provided
by R.Jensen, GKenning and M.C.Tucker.

References

Anonymous, (1994). IUCN Red List Categories.
International Union for Conservation of Nature
and Natural Resources: Switzerland.

Bailey, F.M. (1900). Hydnophytum Jack. Queensland Flora
3: 773-774. Brisbane: H.J.Diddams & Co.

Forster, P.I. (2000). The ant, the butterfly and the
ant-plant: notes on Myrmecodia

beccarii( Rubiaceae), a vulnerable Queensland
endemic. Haseltonia 7: 2-7.

Huxley, C.R.(1982). Ant-epiphytes of Australia. In
R.C.Buckley (ed.), Ant-plant Interactions in
Australia, pp. 63-73. The Hague: Dr.W.Junk
Publishers.

-(1993). The tuberous epiphytes of the Rubiaceae

6: A taxono mi c history of the Hydnophytinae.
Blumea 37: 335-340.

Huxley, C.R. & Jebb, M.H.P. (1991). The tuberous
epiphytes of the Rubiaceae 3: A revision of
Myrmephytum to include Myrmedoma. Blumea
36: 43-52.

-(1993). The tuberous epiphytes of the Rubiaceae

5: A revision of Myrmecodia. Blumea 37:
271-334.

Jebb, M.H.P. (1991). The tuberous epiphytes of the
Rubiaceae 4: A revision of Squamellaria. Blumea
36: 53-61.

Plummer, N.W. (2000). Cultivation of the epiphytic
ant-plants, Hydnophytum and Myrmecodia.
Cactus & Succulent Journal (Los Angeles) 72:
142-147.

Reynolds, S.T. & Halford, D.A. (1997). Rubiaceae. In
R.J.F.Henderson (ed.), Queensland Plants Names
& Distribution, pp. 180-184. Brisbane: Dept,
of Environment.

Alysicarpus (Leguminosae: Desmodieae)
in Australia: a taxonomic revision

Les Pedley

Summary

Pedley, Les (2001). Alysicarpus (Leguminosae: Desmodieae) in Australia. Austrobaileya 6 (1):
107-116. Nine species of Alysicarpus occur in northern Australia, three of them naturalised. All
species are described and a key to their identification and notes on their geographical ranges and
habitats are provided. Alysicarpus aurantiacus, A. major and A. suffruticosus are described as new.

Key words: Australia: Alysicarpus; Alysicarpus aurantiacus; Alysicarpus major; Alysicarpus suffruticosus.

Les Pedley, c/- Queensland Herbarium, Environmental Protection Agency, Mt Coot-tha Botanic Gardens,
Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Introduction

Alysicarpus is a genus of some 30 species
restricted to the tropics and subtropics of the
Old World. It is characterised by its scarious
calyx with rather complex venation (Fig. 1) and
the turgid articles of its pod, indehiscent, and
round or oval in cross-section. Species of the
genus are usually described as having leaves
unifoliolate or ‘very rarely’ (Hutchinson 1964,
Verdcourt 1971) or ‘less often’ (Ali 1977)
trifoliolate. The leaves of all specimens from
Australia examined are unifoliolate It seems
that species with predominantly trifoliolate
leaves should be referred to Desmodiastrum
Prain (Pramanik & Thothathri 1986).

No critical review of the Australia species
has been published. Bentham (1864) recognised
three species: A. vaginalis (L.) DC., A. longifolius
Wight & Arn. and A. rugosus (Willd.) DC., all
of them occurring also in India. From his notes
to A. rugosus it is evident that he took a wide
view of species. Schindler recognised that some
specimens seen by Bentham represented
species different from the Indian ones and
described A. brownii, A. muelleri and A.
schomburgkii. Probably because of its inflated
pod he also transferred Desmodium
campylocaulon F. Muell. ex Benth. to
Alysicarpus , describing Alysicarpus sect.
Desmodiopsis to accommodate it. The transfer

did nothing to sharpen the circumscription of
Desmodium but a lot to obscure the limits of
Alysicarpus. Desmodium campylocaulon is
currently referred to Desmodium , albeit in a
distinct section (Pedley 1999). Backer (1911)
raised A. rugosus var. ludens Baker to specific
rank and referred to it plants grown in Java
from material received from Thursday Is.,
Queensland. Domin (1926) followed Bentham
but described A. rugosus var. longe-exsertus
and identified two specimens as A. rugosus var.
ludens Baker. Presumably he was unaware of
Backer’s treatment of the latter as he remarked
that he had the impression that the taxon would
make a good species. White & Francis (1920)
recorded A. bupleurifolius as naturalised in
Queensland.

Both A. vaginalis and A. rugosus occur
in Africa, and critical work associated with the
Flora Congo-Beige, Ruanda-Urundi and the
Flora of Tropical East Africa has influenced the
perception of species in Australia. Leonard
(1954) distinguished A. ovalifolius from
A. vaginalis ; both appear to be recent
introductions into Australia. The occurrence of
the last has obscured the presence of two
endemic species: A. major, a poorly collected
species from the Kimberley region of Western
Australia and A. aurantiacus, an eastern
Australian species that extends to New Guinea.

Accepted for publication 29 June 2001

108

Austrobaileya 6 (1): 107-116 (2001)

Key to native and naturalised (*) species of Alysicarpus

1. Compact shrub to 1 m tall; stems and both surfaces of leaflets villose with

dense spreading hairs 0.8-1mm long; petioles 2-3 mm long; pod unknown

.8. A. suffruticosus

Annuals or perennials, prostrate or erect, usually sparingly branched; stems
and leaflets only sparsely hairy, not villose.2

2. Pod not, or only slightly, constricted between the articles; articles smooth

or obscurely reticulately veined when dry; annual or perennial plants

.3

Pod markedly constricted between the articles; articles predominantly
reticulately veined; plants annual.8

3. Articles of pod glabrous, not at all rugose or reticulately veined; pod slightly

constricted between the articles; calyx lobes overlapping at base of fruit

.3. *A. bupleurifolius

Articles of pod glabrous or pubescent, pod not constricted between them;
calyx lobes not overlapping at base of fruit.4

4. Leaflet lanceolate, 7-9 cm long, prominently reticulately veined on both

surfaces; pod flattened.2. A. brownii

Leaflet orbicular, broadly oblong, ovate or obovate, or lanceolate (sometimes
all on a single plant), rarely more than 7cm long (only in A. schomburgkii)
obscurely reticulately veined when dry; articles scarcely flattened.5

5. Flowers in short, rather open, racemes, terminal and in upper axils forming

an open leafy panicle; leaflet 10-20(-24) mm wide; articles 2-3 mm long;

plants annual. 6. *A. ovalifolius

Flowers in rather dense terminal racemes, seldom in upper axils; leaflets
3-12 mm wide; articles to 4 mm long; plants perennial.6

6. Racemes dense, flowers overlapping at anthesis; pod articles somewhat

flattened, pubescent, 3.5-4 mm long; seeds not seen, probably smaller

than those of A. aurantiaceus and A. vaginalis .4. A. major

Racemes open or dense; pod articles scarcely flattened, glabrous, to 3.5
mm long; seeds c. 2 x 1.5 mm.7

7. Leaflets (2-) 3-15 times longer than wide, widest below the middle when

elongate, acute, subacute, or rarely, in the lower part of the plant, rounded
at the apex, drying greyish; inflorescence rather open at anthesis; calyx

5.5-7.7mm long; standard 6.5-8 x 4-7 mm; petals orange or yellow. 1. A. aurantiacus

Leaflets 1.3-2.5 (-4) t im es longer than wide, widest at or above the middle,
rounded at the apex, drying brown or greenish; inflorescences dense at
anthesis; calyx 4.5-5 mm long; standard c. 6 x 3.4mm; petals at least
tinged pink, violet or purple.9. *A. vaginalis

8. Calyx 7-9 mm long; pod articles 2.5-4 mm long, 3-3.3 (-4) mm wide,

coarsely reticulate, pale straw-coloured when mature.5. A. muelleri

Calyx 4-6 mm long; pod articles 1.5-2.5 mm long, 1.5-2 mm wide, finely
transversely reticulate, black when mature. 7. A. schomburgkii

Pedley, Alysicarpus in Australia

109

Fig. Alysicarpus aurantiacus. A. Flower (x8), just before opening. B. Calyx (x8), flattened out, lower lobe at right. C.
Pod (x4), slightly immature. A, B, L.A. Craven 324; C, A.R. Bean 2901.

1. Alysicarpus aurantiacus Pedley, sp. nov.
affinis et saepe confusus A. vaginali (L.)
DC. a quo foliolis elongatioribus, 3-
15plo, rarius 2plo, (non 1.3-2.5 plo)
longioribus quam latis, infra medium
latissimis ubi elongatis, cinereis in sicco,
inflorescentia floribus sub anthesi non
confertis, calycibus longioribus, 5.5-7.7
mm non 4.5-5 mm, petalis aurantiacis
flavisve non roseis purpureis violaceisve
tinctis differt. Typus: Queenland. North
Kennedy District: 45 km S of Townsville,
19°32’S, 146°45’E, 13 May 1979, L.
Pedley 4588 (holo: BRI; iso: MEL).

Alysicarpus vaginalis auct. non (L.) DC:
Bentham, FI. Austral. 2: 239 (1864);
Verdcourt, Man. New Guinea Leg.: 420
t. 98 (1979).

Prostrate or ascending perennial herb. Stem
glabrous or with scattered appressed or
ascending hairs and/or sparse weak uncinate
hairs; stipules 6-15 (-18) mm long. Leaflet
(10—) 15-45 (-50) mm long, 3-11 mm wide,
(2-) 3-11 (-15) times longer than wide,
narrowly lanceolate to narrowly ovate or
oblong, subcordate to rounded at the base,
subacute or rarely, on lower part or plant, obtuse
at tip, mucronulate, glabrous or with scattered
long hairs above, sparse or moderately dense
appressed hairs beneath or hairs only on veins;
petiolules 0.5-1 mm long; stipels somewhat
longer than petiolules; petioles (3-) 5-8 (-10)
mm long. Inflorescences rather open, terminal,

to c. 50 mm long; rachis with uncinate hairs;
bracts ovate c. 4 mm long; pedicels paired, 1-
2 mm long. Flowers orange or yellowish: calyx

5.5- 7.7 mm long, the tube 1.7-2.2 mm, the
lobes narrowly acute, not overlapping at base,
the upper to 3.5 mm long, slightly shorter than
the others, bifid at apex; corolla: standard
obovate, 6.5-8 mm long, 4-7 mm wide; wings

5.5- 6.5 mm long, clawed, lamina oblong; keel
petals about as long as wings; ovary uncinately
pubescent. Pods with up to 8 articles, straight,
not constricted between seeds, to c. 20 mm long;
articles obscurely reticulate, uncinately
pubescent, (2-) 2.5-3.5 mm long, 2-2.5mm
wide; seeds 1.4-2 mm long, 1.2-1.5 mm wide.
Fig. 1.

Selected specimens: Papua New Guinea. Central
District: Moitaka, 9°30’S 147°10’E, Dec 1964, Gillison
NGF 22038 (BRI); Milne Bay District: near Modino, north
east of Cape Vogel Pen., Sep 1954, Hoogland 4676 (BRI).
Queensland. Cook District: Lockerbie, 10 miles [16 km]
WSW of Somerset, Apr 1948, Brass 18463 (BRI);
Endeavour River, in 1882, Persietz 261 (MEL). North
Kennedy District: Saltwater Creek [Rockingham Bay],
Nov 1865, Dallachy (MEL); Townsville, Feb 1918, White
s.n. (BRI). South Kennedy District: Kelsey Creek, Michael
814 (BRI); 24.3 km from Proserpine towards Mackay,
20°35’S 148°37’E, Apr 1991, Bean 2901 (BRI). Port
Curtis District: Rockhampton, Feb 1868, 0’Shanesy No. 15
ser. 9 (MEL); Bororen, Oct 1935, Kohler (BRI).

Distribution and habitat : Alysicarpus
aurantiacus extends from southern New
Guinea through coastal districts of Queensland
from Cape York to about Maryborough. It
grows among grasses (often Heteropogon

110

Austrobaileya 6 (1): 107-116 (2001)

contortus or Themeda triandra ) usually in
eucalypt communities.

Affinities: The species is closely related to,
and often confused with, A. vaginalis. It differs
in its more open inflorescences, its more
elongate, usually subacute leaflets and orange
or yellowish flowers. Its leaflets dry greyish
whereas those of A. vaginalis are brown or
retain a greenish tinge.

Etymology: The specific epithet is Latin
meaning ‘orange’, a reference to the colour of
the flowers of the species.

2. Alysicarpus brownii Schindler, Trans. &
Proc. Bot. Soc. Edinb. 26:286 (1915).
Type: Northern Australia, [Melville Bay
(Arnhem Bay North), 16 February 1803,
see below], R. Brown, ‘Iter Australiense
4176’ (holo: E; iso: BM, K).

A. longifolius auct non (Spreng.) Wight &
Arn.: Benth., FI. Austral. 2:239 (1864).

Perennial with erect annual stems to 2.5 m tall
(teste Maconochie 2000) from persistent
rootstock. Stem terete, faintly longitudinally
striate, uncinately pubescent, often with a few
appressed hyaline hairs c. 0.5 mm long,
glabrescent; stipules scarious, long pointed 15-
25(-30) mm long. Leaflet narrowly lanceolate
or linear lanceolate, sharply acute, slightly
cordate or rounded at base, the margin slightly
inrolled, (65-) 85-130 (-160) mm long, 3.5-
10 (-15) mm wide, (10-) 12-22 (-27) times
longer than wide, rather coriaceous,
conspicuously reticulately veined on both
surfaces, glabrous except for some uncinate
hairs on veins and long appressed ones on
margins and midribs beneath; petiolule 0.5-1.5
mm long; stipels cartilaginous, about as long
as petiolules; petioles (3-)7-12 mm long.
Inflorescence terminal, open in fruit, to c. 30
cm long; rachis densely uncinately pubescent;
primary bracts ovate acuminate, 4-6 mm long,
secondary bracts linear about half as long, all
early deciduous; pedicels paired, densely
pubescent, 0.5-3 mm long at anthesis, slightly
longer in fruit. Flowers yellow or orange; calyx
4-5.5 mm long, densely uncinately pubescent,
with stiff hyaline hairs on lobes, tube 1.6-2 mm
long, lobes 2.2-3.5, the upper one slightly

shorter than the others, bifid for c. 1 mm; corolla:
standard broadly obovate, 6.5-8 m long, 5-6.5
mm wide; wings 6-7.3 mm long (including claw
1-2 mm long), 1.6-2 mm wide; keel petals about
as long as keel, not clawed; ovary uncinately
pubescent, occasionally with some hyaline hairs
at apex extending to base of style. Pods with up
to five somewhat flattened articles each 3.5-4
mm long, 2-2.5 mm wide (somewhat smaller than
described by Schindler), obscurely reticulately
veined, uncinately pubescent, slightly
thickened at the septum when dry; seeds not
seen.

Selected specimens: Northern Territory. Berry Springs,
12°42’S 131°00E, Dec 1974, Must 1337 (DNA); Katherine
Experimental Station, 14°27’S 132°15’E, Jan 1978,
Maconochie 2310 (DNA, K); 9.5 km S. of Fish River
homestead turn-off, Jun 1974, Maconochie 2000 (BRI,
DNA, K); no definite locality (possibly Nabarlek, 12°22’S
133°22’E), Apr 1982, Hinz s.n. (DNA).

Distribution and habitat: The species is
confined to the north of the Northern Territory
where it is reported to occur in eucalypt
communities, probably in seasonally
waterlogged sites. It is poorly collected and is
evidently rare.

Affinities: In its protologue, Schindler noted
Alysicarpus brownii to be nearly related to
A. vaginalis. It differs in its erect habit, longer
leaves, larger flattened pod-articles and colour
of the flowers. It is probably closer to the
yellow-flowered endemic A. major but differs
in its habit and size and shape of leaflets.

Notes: Type material at E and K has no details
of a collecting locality, but the isotype at BM is
well labelled. The article of the pods on the
specimens examined are somewhat smaller than
as described by Schindler.

3. Alysicarpus bupleurifolius (L.) DC., Prodr.
2: 352 (1825); White & Francis, Bot. Bull.
(QldDepart. Ag. & Stock) 22: 15 (1920);
Meeuwen, Reinwardtia 6: 88 (1961);
Verdcourt, Man New Guinea Leg. 420. t.
98G (fruit) (1979); Nguyen, FI. Camb.
Laos Vietnam 23:130 t. 22, 1-5 (1987);
Pedley, Rev. Handb. FI. Ceylon 10: 153
(1996). Hedysarum bupleurifolium L.,
Sp. PI. 745 (1753). Type: herb. Linnaeus
921.3 (LINN).

Pedley, Alysicarpus in Australia

111

Erect or ascending annual herb. Branches with
covering of appressed hairs, becoming
glabrous; stipules acute, 3-10 mm long. Leaflet
narrowly ovate on young plants, lanceolate to
linear on older ones, rounded at the base acute
at the apex, usually 30-70 mm long, 3-4.5 mm
wide, 10-20 times longer than wide,
occasionally down to 5 mm long, up to 9 mm
wide and only twice as long as wide, glabrous
on upper surface, sparse appressed hairs on
lower; prominently reticulately veined on both
surfaces; petiolules 0.5-1 mm long; stipels
minute or absent; petioles 2-4 mm long.
Inflorescences terminal, dense, to 20 mm long
in flower, open, to 50 mm or more in fruit;
rachis glabrous; pairs of flowers subtended by
acute or acuminate bract, deciduous at anthesis;
pedicels 0.8-1.5 mm long, with ascending
brown hairs, becoming glabrous. Flowers:
calyx rather stiff, angled towards the base, 6-7
mm long, the lobes equal in length, strongly
overlapping at the base, 4-5 mm long, the upper
entire or shortly bifid, glabrous; corolla orange
and purple; standard oblong 4-5 mm long, 1.5-
2.5 m wide, the wings and keel petals about
equal in length, shorter, about 4 mm long; ovary
glabrous. Pods to 15 mm long, slightly to
markedly exserted from the calyx, not
constricted between the articles; articles 3-8,
cylindrical or slightly flattened, 1.6-2 mm long,
1.3-2 mm wide, smooth or very obscurely
reticulately veined, glabrous; seeds almost
cubic, about 1 mm long and 0.8 mm wide.

Selected specimens (all BRI ): Queensland. Cook District:
Mareeba, Apr 1962, McKee 9069. North Kennedy District:
Cromarty, near Townsville, Mar 1933, White 8842.
Leichhardt District: Camp Gully, 45 miles [72 km] SSW
of St Lawrence, May 1962, Johnson 2293. Port Curtis
District: Rodds Bay, near Gladstone, Mar 1955, Shaw s.n.
Wide Bay District: Bundaberg, near city centre. Mar 1980,
Stanley 962. Moreton District: The Gap, Brisbane, Feb
1963, Blake 22024 (CANB, K, MEXU).

Distribution and habitat : The species ranges
from India, Sri Lanka and south-east Asia
through Malesia to Australia. In Australia it
occurs in coastal and subcoastal districts
between Mareeba and Brisbane, on roadsides
and in eucalypt communities often on drainage
lines. Some collectors have noted its being
relished by cattle. It is probably a recent
introduction to Australia first collected near
Townsville in 1918.

Affinities : Alysicarpus bupleurifolius is a well
defined species showing little variation in
Australia and is not often confused with any
other species. In habit it resembles
A. schomburgkii which, however, has pods
strongly reticulately veined, constricted
between the articles.

4. Alysicarpus major Pedley, sp. nov. similis
A. vaginali (L.) DC. a quo floribus flavidis
(non subroseis purpurascentibusve)
vexillo majore c. 8 mm longo, leguminum
articulis aliquantum complanatis
pubescentibus longioribus 3.5-4 mm
(non 2-3 mm) longis differt. Typus:
Western Australia. Donkeys Creek, 29 km
S of ‘Drysdale River’ Station, c. 15°27’S
126°20’E, 25 May 1975, D.E. Symon
10165 (holo: AD; iso: PERTH & (n.v.)
NSW).

Prostrate or ascending perennial. Stems from
stout tap-root with scattered short weak
uncinate hairs, glabrescent; stipules (8-) 12-
15 mm long. Leaflet somewhat dimorphic, short
towards base of plant, 15-25 (-30) mm long,
7-14 (-16) mm wide, (1.4—) 1.8-2.4 (-3) times
longer than wide, oblong (rarely ovate), obtuse
mucronulate at the apex, cordate at the base,
glabrous above with stiff appressed hairs
beneath, strongly reticulately veined on both
surfaces; petiolule c. 1 mm long; stipels about
as long as the petiolules; petiole 4-7 mm long.
Inflorescence terminal, rather dense at anthesis,
pedicels paired, 1.5-2 mm long, bracts not seen.
Flowers: calyx 5-5.5 mm long with minute
uncinate hairs on tube and longer apressed hairs
on lobes, tube 1.8-2 mm long, upper lobe 3-
3.5 mm long, bifid for 1-1.5 mm, lateral lobes
3.2-3.5 mm long, lower lobe 3-3.2; corolla:
standard (orange) c. 8 mm long, 3.5 mm wide,
wings 5.3-5.6 mm long, the claw 1.2-1.3 m
long, c. 1.6 mm wide, keel petals (reddish) c. 6
mm long, not clawed; ovary with a few long
hairs at the apex. Pod to c. 20 mm long with up
to 5 articles, not contracted, and sometimes with
no perceptible junction, between them; articles
slightly flattened, uncinately hairy, 3.5-4 mm
long, c. 2.5 mm wide, the most proximal and
distal larger than others; seeds not seen.

112

Austrobaileya 6 (1): 107-116 (2001)

Selected specimens : Western Australia. Sir Graham
Moore Is., S side, 13°56’S 120°33’E, Jul 1973, Paul G.
Wilson 11310 (PERTH); ... Secure Bay, 16°27’S
124°13’E, May 1993, Mitchell 3130 (K, PERTH);King
Leopold Ranges, close to road crossing of Fern
Creek,(17°09’ S 125° 16’E) c. 310 m, Apr 1988, Sands
4211 (BRI, K, PERTH).

Distribution and habitat: Alysicarpus major
is known only from the Kimberley region of
Western Australia where it is reported to grow
on laterite.

Affinities: The species is related to A. vaginalis
from which it differs in having orange and rust-
red petals and larger, somewhat flattened hairy
pod-articles. It also has some affinity with the
endemic A. aurantiacus from eastern Australia
which also has yellow or orange petals and
pubescent pods but the articles of
A. aurantiacus are smaller and its leaflet
narrower.

Etymology: The epithet, Latin meaning ‘larger’
or ‘greater’, alludes to the plant’s having
somewhat larger flowers and fruits than its
relative A. vaginalis.

5. Alysicarpus muelleri Schindler, Rep. spec,
nov. reg. veg. 22: 269 (1926). Type:
Victoria River and Depot Creek, F.
Mueller (holo or iso: K; iso (?): MEL).

A. muelleri var. clementii Schindler, op. cit .:
270 (1926). Syntypes: Western
Australia: between Ashburton and Yule
Rivers, Clement s.n. (iso: K); between
Ashburton and De Grey Rivers, Clement
s.n. (iso: K).

A. rugosus var. longe-exsertus Domin,
Biblioth. Bot. 89: 217 (1926). Type: in
fl. Flinders River ad Hughenden, Feb
1910, K. Domin Iter australiense ‘4706’
(lecto, chosen here: PR, herb, no
527404).

A. rugosus auct non (Willd.) DC.; Benth,
Fl. Austral. 2: 239 (1864).

Erect annual to 1 m. tall. Stems with
indumentum of short uncinate hairs and
scattered long (0.5-1.5 mm) straight spreading
or ascending hairs; stipules somewhat
membranous, 3-10 (-20) mm long. Feaflet
linear, oblong ovate-lanceolate or narrowly

ovate, obtuse or subacute, mucronulate at the
apex, rounded or truncate at base, 25-75 mm
long, 5-22 mm wide, 2.3-5 (-7) times longer than
wide, glabrous or with scattered long hairs
above, scattered long hairs and usually
moderately dense microscopic uncinate hairs
beneath; petiolules 1-1.5 (-2) mm long, stipels
shorter than petiolules 0.3-1 (-1.5) mm long;
petioles 2.5-6 (-7.5) mm long. Inflorescence
terminal, dense; rachis with indumentum
similar to that of stems, though occasionally
with multicellular glandular hairs towards the
tip; bracts concave, ovate, acuminate 5-6 (-8)
mm long, 2-2.5 mm wide, indumentum similar
to that of calyx, subtending two flowers; pedicel
1.5-4 mm long. Flowers with yellow standard
and orange or red wings; calyx 7-9 mm long,
the tube c. 1 mm long, the lobes distinctly
overlapping at the base at anthesis, the upper
united almost to the tip, the lower slightly longer
than the others; corolla: standard obovate,
slightly retuse, 5-6.5 mm long, 3-4 mm wide;
wing 4-4.5 mm long, including claw 0.8-1.5
mm; keel petals strongly tapered to base but
not clawed, about as long as the wings, ovary
densely puberulent. Pod of usually 4 articles
(occasionally 3 or 5) exserted from calyx;
articles 2.5-4 mm long, 3-3.3 (-4) mm wide,
coarsely transversely reticulate with scattered
minute uncinate hairs, pale straw coloured
when mature; seeds subglobular 2-2.4 mm long
slightly less than 2 mm wide.

Selected specimens : Western Australia. Road Paddock
c. 3km S of Kalumburu Mission 14°18’05”S 126°39’08”E,
Jun 1966, Mitchell 4437 (BRI, PERTH); Ord River, W of
‘Ord River’ Station 17°26’S 128°49’E, Apr 1977, Pullen
10754 (BRI, CANB). Northern Territory. Katherine
Gorge Road, 5 miles [8 km] NE of Katherine, Mar
1964, Adams 903 (BRI, CANB, K); McArthur River,
16°46’S 135°45’E, May 1947, Blake 17812 (BRI, K);
25Vi miles [41 km] W of 'Rockhampton Downs’, Jun
1960, Chippendale NT 7125 (BRI, CANB).
Queensland. Burke District: Kilty Plain, 18 km NW
of Mt Isa, 20°37’S 139°21’E, Mar 1987, Harris 152
(BRI); ‘Lydia Downs’ c. 45 miles [72 km] NW of
Maxwellton, Jan 1966, Pedley 1960 (BRI). Leichhardt
District: ‘Lenton Downs’ 65 miles [104 km] WNW of
Nebo, May 1962, Johnson 2308 (BRI).

Distribution and habitat: Alysicarpus muelleri
occurs across tropical Australia. It is a summer¬
growing annual virtually restricted to dark grey
and brown clay soil in grassland dominated by
Mitchell grass (Astrebla spp.) and Flinders
grasses (Iseilema spp.). In the eastern part of

Pedley, Alysicarpus in Australia

113

its range, however, it has been collected from
sandy soils that have been tilled.

Affinities : The plant has been confused with
A. rugosus sensu stricto, a plant not found in
Australia, which has pod-articles finely
transversely reticulate, black at maturity. It is
certainly close to A. rugosus subsp. reticulatus
Verde, from Africa.

6. Alysicarpus ovalifolius (Shumach.)
Leonard, Bull. Jard. Bot. Etat. Brux. 24:
881.11(1954), FI. Congo Beige Ruanda-
Burundi 5: 226. t. 13B (fruit) & t. 18
(1954); Verdcourt, FI. Trop. East Africa.
Leguminosae: Papilion. 493 (1971).
Hedysarum ovalifolium Schumach,
Beskrwelse af Guineeske Planter 359
(1829). Type: Africa: ‘Guinea’ [Ghana:
Ada, fide Verdcourt], Thonning (holo: C,
n.v., photograph in Leonard, loc. cit.).

Annual with main stem erect, sometimes to 1
m tall, lower branches prostrate or often whole
plant ± prostrate. Stems strigose with scattered
yellowish ascending hairs, occasionally with
minute weak uncinate hairs, rarely glabrous;
stipules narrowly triangular, 5-12 mm long.
Leaflet orbicular, broadly oblong or obovate
to narrowly ovate, rounded, truncate or slightly
retuse at apex, subcordate or cordate at base,
(8-) 15-40 (-50) mm long, 10-20 (-25) mm,
1.2-2.5 times longer than wide, glabrous above,
minutely uncinately pubescent with some
appressed hairs on veins beneath; petiolules
(0.5-) 1 (-1.5) mm long; stipels cartilaginous,
obtuse about as long as petiolules; petioles 3-9
(-12) mm long. Inflorescence open to
moderately dense racemes, terminal and in
upper axils, forming an often leafy panicle;
rachis glabrous in fruit, pedicels paired 0.5-2
mm long; bracts 2-4 mm long, concave,
acuminate. Llowers orange to red; calyx c. 4.5
mm long the tube 2 mm long, upper lobe c. 2
mm long, shortly bifid, the rest c. 2.5 mm long;
corolla: standard obovate, c. 6 mm long and 4
mm, wings clawed, c. 5 mm long, 1.2 mm wide;
keel petals slightly longer than wings, not
clawed, c. 5.5 mm long, 1 mm wide; ovary
uncinately pubescent. Pods + cylindrical, not
constricted between articles, to 24 mm long,
with up to 8 articles each 2-3.5 mm long, 1.5-
2 mm wide, obscurely reticulately veined

glabrous or minutely uncinately pubescent;
seeds obloid, smooth, yellow or pale brown,
variable in size, 1.7-3.2mmlong, 1-2.5 mm wide.

Selected specimens: Western Australia. Mitchell
Plateau, Old Mining Camp, 14°50’S 125°51’E, Apr
1988, Dunlop 7906 (BRI, DNA, MEL, NSW, PERTH).
Northern Territory. Melville Is., Garden Point barge
landing, 11°24’S 130°25’E, Mar 1994, Cowie 4713
(BRI, DNA); Fog Dam area, about 40 miles [64 km] SE
of Darwin, May 1959, Chippendale NT 6189 (BRI,
DNA); ‘Malapunya’ Station, 16°58’S 135°49’E, Mar
1981, Maconochie 2628 (BRI, CANB, NT).
Queensland. Burke District: ‘Myra Vale’, Normanton,
Mar 1944, Bell s.n. (BRI). Cook District: ‘Heathlands’,
Mar 1992, Johnson 5200 & Sharpe (BRI); 4.8 km N
of Little Laura River, Apr 1983, Clarkson 4782 (BRI).
North Kennedy District: Townsville, Feb 1918, White
s.n. (BRI); Sinclair Bay, 20°05’S 148°26’E, Mar 1994,
Batianoff 9403263 (BRI). Port Curtis District:
Cedarvale, 23°56’S 150°54’E, Partridge s.n. (BRI)

Distribution and habitat: Alysicarpus
ovalifolius is a native of Africa, naturalised in
Australia. The earliest collection seen is from
Townsville in 1918. At present it occurs
sporadically in the Kimberley region of Western
Australia, the north of the Northern Territory
(common around Darwin) and sporadically
along the Queensland coast from Thursday Is.
to a little south Rockhampton where it was
recorded in cultivated land. It is found usually
in disturbed situations on soils of various types,
probably most commonly on sand. It is reported
to be eaten readily by cattle and to make
excellent hay.

Affinities'. The species is closely related to
A. vaginalis and the two are often confused.
They are easily identified in the field but a small
proportion of herbarium specimens is difficult
to identify. Verdcourt (1971) mentioned
intermediates between A. ovalifolius and
A. vaginalis but none has been observed in
Australia. Meeuwen (1961) and Dunlop et al.
(1995) did not accept A. ovalifolius as distinct
from A. vaginalis, though the latter, in a note,
distinguished the two species quite well. In a
well presented paper, Endo & Ohashi (1990)
illustrated that attributes of the pod provided
reliable distinctions between the two. As noted
by them (op. cit.), A. ovalifolius occurs in the
Philippines and the Lesser Sunda Is. and
specimens from New Guinea (near Port
Moresby, Gillison 22113, and near Lae,
Streimann & Kauro NGL 27882 - both BRI)
have also been seen. Leaflets of depauperate

114

Austrobaileya 6 (1): 107-116 (2001)

plants of A. ovalifolius may be only 7 mm long
and 6 mm wide, but the open inflorescences,
attributes of the pod and the presence of long
hyaline antrorsely curved hairs on the stems
distinguish it from A. vaginalis.

7. Alysicarpus schomburgkii Schindler, Repert.
sp. nov. reg. veg. 22:269 (1926). Syntypes:
Port Darwin, Schomburgk 84 (iso: K); Port
Darwin, Foelsche 97 (iso: MEL); Palmer
River, Wycliffe (iso: BM, MEL ?); Thursday
Is., Jaheri s.n. (iso: K).

A. ludens auct. non. (Baker) Backer; Backer,
Schoolflora voor Java: 349 (1911).

A. rugosus var. ludens auct. non Baker;
Domin, Bibliotheca Bot. 89: 217 (1926).

Annual 60 cm, rarely to 1 m tall, main stem
erect, basal branches ascending. Young stems
angular throughout, glabrous or with appressed
hairs on the angles or occasionally terete with
hairs on ribs; stipules 2-12 (-14) mm long.
Leaflet elliptic, narrowly ovate or narrowly
obovate, obtuse at base of plant, becoming
lanceolate, acute in upper part, rounded at base,
15-30 mm long, 3-7.5 mm wide, 2.5-7 times
longer than wide becoming 30-145 mm long,
2.2-9 mm wide, 9-20 (-40) times longer than
wide, glabrous on upper surface, stiff ±
appressed hairs scattered on veins beneath and
sometimes with sparse minute uncinate hairs;
petiolules c. 0.5; stipels shorter than petiolules,
somet im es absent; petioles 1.5-3 (-5.5) mm
long. Inflorescence terminal, moderately dense;
rachis with indumentum of minute uncinate
hairs and usually longer stiff erect yellowish
hairs; bracts ovate acute, 3-6 mm long, 1.5-3
mm wide, subtending 2 flowers; pedicels 1.5-
3 (-5.5) mm long. Llowers yellow; calyx 4-6
(-8) mm long, the tube c. 1 mm long, lobes
overlapping at base slightly at anthesis, upper
lobe slightly bifid at tip; corolla: standard
obovate, obtuse, 3.5-6 mm long, 2.5-4 mm
wide; wings 4-5.5 mm long including claw c.
1 mm long; keel petals as long as or slightly
shorter than wings, claw 1-1.5 mm long; ovary
glabrous or minutely puberulent towards the
tip. Pod to c. 8 mm long of 2-4 articles, deeply
constricted between them, each 1.5-2.2 mm
long, 1.5-2 mm wide, transversely reticulately

veined (not as closely or deeply as in
A. rugosus), glabrous or sparsely puberulent,
blackish when mature; seeds smooth, brown
subglobular 1.4-1.6 mm long, 1.3-1.4 mm wide.

Selected specimens: Western Australia, near
Manning gorge, 16°39’S 125°55’E, Jun 1978, George
15189 (PERTH). Northern Territory. Darwin, Oct
1946, Blake 17315 (BRI, CANB, MEXU, NT);
Litchfield N.P., 13°03’S 130°50’E, Mar 1995, Cowie
5257 & Taylor (BRI, CANB, DNA, PERTH); Jabiru,
12°40’S 132°53’E, Feb 1973, Dunlop 3351 (BRI,
CANB, DNA, MEL, MO). Queensland. Cook District:
Milman Hill, Thursday Is., 10°35’S 142°13’E, Apr 1986,
Clarkson 6421 (BRI, DNA, K, L, MBA, MO, NSW,
PERTH, QRS); Mareeba Mining Lease, c. 15 km SE of
Mareeba, 17°15’S 145°30’E, Apr 1972, Staples
030472/5 (BRI, CANB).

Distribution and habitat: Alysicarpus
schomburgkii is most common in the extreme
north-west of the Northern Territory, with
scattered occurrences in the Kimberley region
of Western Australia and in the eastern tropical
part of Queensland from Thursday Is. to about
Proserpine on the central coast. It has been
recorded from shallow skeletal soils, sands, red
earths derived from laterite and rarely, heavy
alluvial soil, occasionally near seasonally
flooded depressions.

Affinities: The affinities of the species are
with A. rugosus and, less closely, A. muelleri,
both of which have pods less constricted
between the articles which are also larger. The
articles of A. rugosus have a closer, more
sharply defined reticulum, and become much
darker than those of A. schomburgkii when
mature. Specimens examined suggest that
plants from Queensland have smaller flowers
than those from the Northern Territory. Their
standards are 3.5-4 mm long (N.T.: 4.5 mm or
more) and have 2 or 3 ovules (N.T. usually 4,
occasionally 3).

Backer and Domin independently
identified specimens from Australian as the
Indian taxon currently accepted as A. heyneanus
var. ludens (Baker) A. Pramanik & Thoth.
(Pramanik & Thothathri 1983).

8. Alysicarpus suffruticosus Pedley sp. nov.
affinis A. vaginali (L.) DC. a quo planta
suffruticosa, petiolis brevioribus, foliolis
utrinque villosis, et fortasse coloribus
florium differt. Typus: Western Australia.

Pedley, Alysicarpus in Australia

115

4 km NNW of Milliwindi road turn-off,
opposite Mt Hart outcamp, 17°06’S
124°12’E, 15April 1988 ,R.J. CranfieldMOl
(PERTH); iso (n.v.): CANB).

Compact subshrub to 30 cm tall with stout
taproot. Stem + terete, with dense spreading
hairs c. 0.8 mm long and scattered weak
uncinate hairs; stipules 3-6 mm long, 1.5-2 mm
wide, acute, closely longitudinally veined,
spreading and lacinate when old. Leaflet oblong
or ovate, rounded or slightly retuse at the apex,
rounded or slightly cordate at the base, 15-30
mm long, 8-16 mm wide, 1.2-2.5 t im es longer
than wide, villose with weak hairs on both
surfaces (or hairs somewhat shorter beneath,
or less dense above) strongly reticulately veined
beneath; petiolules c. 0.6 mm long; stipels about
half as long as petiolules, often obscure; petioles
2-3 (-5) mm long. Inflorescences terminal to
c. 80 mm long; rachis with uncinate hairs; bracts
ovate acuminate, c. 4 mm long; secondary
bracts c. 2 mm long, oblong, both early
decidous. Flowers ‘pale pink’ (teste Cranfield)
or ‘yellow and rust red’ (teste Sands); calyx c.
5 mm long, the tube 1.5 mm, lobes narrow,
acute, not overlapping at the base, the upper
slightly shorter than the others, bifid at apex,
indumentum of minute uncinate hairs with 0.8
mm long hyaline hairs on margin of lobes;
corolla: standard obovate, obtuse, c. 8 mm long,
5 mm wide; wings 6.3 mm long (including claw
1 mm long), 1.3 mm wide; keel petals 5.7 mm
long; ovary with indumentum of long appressed
hairs, ovules 5, style truncate. Pods and seeds
not seen.

Selected specimen: Western Australia. Old Settlement,
Sunday Is., 16°24’S 123°11’E, Apr 1992, Mitchell 2231
(PERTH).

Distribution and habitat: Alysicarpus
sujfruticosus is known from the two localities
cited, in the Kimberley region of Western
Australia. It has been recorded from sandy soil
in grassland and beneath Livistona sp.

Affinities: The species is allied to A. vaginalis
and A. aurantiacus but is easily distinguished
from both by its growth form and the villosity
of its leaflets.

Etymology. The epithet is Latin meaning
somewhat woody, an allusion to perennial
shrubby habit of the species best shown in the
Sands specimen cited.

9. Alysicarpus vaginalis (L.) DC., Prodr. Syst.
Veg. 2: 353 (1825); Meeuwen,
Reinwardtia 6: 87 (1961) pro parte (incl.
A. ovalifolius)', Leonard, FI. Congo Beige
Ruanda-Urundi 5: 224 (1954); Verdcourt,
FI. Trop. East Africa. Leguminosae:
Papilion. 493 (1971), Man. New Guinea
Leg. (1979) pro parte (incl.
A. aurantiacus)', Nguyen, FI. Camb. Laos
Vietnam 23: 128 (1987); Pedley, Rev.
Handb. FI. Ceylon 10: 152 (1996).
Lectotype: Sri Lanka (Ceylon), herb.
Hermann 1:27, No. 287 (BM), fide
Verdcourt in Turland & Jarvis, Taxon
46:473 (1997).

Prostrate perennial. Stems glabrous or sparsely
minutely uncinate pubescent; stipules 4-12(-
15) mm long, often longer than the petioles.
Leaflet broadly ovate or oblong (orbicular when
young), rarely ovate, rounded at apex, cordate
at base, (5-) 8-35 mm long, 4-12 (-14) mm
wide 1.3-2.5 (-4) times longer than wide,
glabrous above, sparsely uncinately pubescent
beneath with scattered appressed hairs,
sometimes only on veins; petiolule c. 0.3 mm
long, stipels about as long as petiolules; petiole
4-10 mm long. Inflorescence terminal, dense
at anthesis; rachis uncinately pubescent; bracts
narrowly ovate, 4-5 mm long; pedicels paired,
c. 1.5 mm long. Flowers bluish, purplish or
mauve to wine red, not yellow or orange; calyx
4.5-5.5 mm long, tube 1.7-2.5 mm long, lobes
subequal 2.3-3 mm long, the upper shortly bifid
at apex, somewhat shorter than the other;
corolla: standard ovate c. 6 mm long, 3-4 mm
wide, wings as long as or slightly shorter than
keel clawed, 4.5-6 mm long, 1-1.5 mm wide;
keel petals not clawed, 5-6 mm long, 1.2-1.5
mm wide; ovary sparsely uncinately
puberulent. Pods to 18 mm long with up to 8
articles, not contracted between them; articles
drying black, obscurely coarsely reticulately
veined when mature, 2-3 mm long, 2-2.3 mm
wide; seeds obloid, c. 2 mm long, 1.3 mm wide.

Selected specimens: Northern Territory, c. 12°40’S
131°20’E, Sep 1946, Blake 17004 (BRI); Bamboo
Point, Mary River, Apr 1972, Barrett 8 (DNA, K).

116

Austrobaileya 6 (1): 107-116 (2001)

Queensland. Cook District: Thursday Is., 10°35’S
142°13’E, Apr 1986, Clarkson 6444 (BRI, DNA, K, L,
MBA, MEL, MO, NSW, PERTH, QRS); Rutland Plains
near mouth of Mitchell River, Jun 1943, Whitehouse
s.n. (BRI). North Kennedy District: Nettle Creek, 10
km E. of Mt. Garnet, Jul 1990, Silcock SI050 (BRI).
South Kennedy District: Slade Point, Mackay, 21°05’S
149°14’E, Apr 1975, McDonald 1304 & Batianoff
(BRI). Leichhardt District: Boothill Creek, 10 miles
[16 km] S of Nebo turnoff on [Old] Bruce Highway, c.
21°50’S 148°55’E, Jun 1958, Pedley 270 (BRI). Port
Curtis District: North Rockhampton, Feb 1980, Stanley
557 (BRI).

Distribution and habitat: Alysicarpus
vaginalis is widely spread in the tropics of the
Old World but is probably a 20th century
introduction to Australia. It is rare in the
Kimberley region of Western Australia and the
north of the Northern Territory but is more
common in Queensland where it occurs in
sandy soils in eucalypt communities. It is,
however, essentially an urban weed, occurring
in lawns and on road verges in Gladstone,
Rockhampton, Mackay, Townsville and Cairns.

Affinities: The species is frequently confused
with A. ovalifolius. The latter is a less
conspicuously prostrate plant with much larger
leaflets (when well grown) and always with a
more open branched inflorescence. The
differences between A. vaginalis and the
indigenous A. aurantiacus are more subtle.
A. aurantiacus has more elongate leaflets, a
more open inflorescence, though not branched
as in A. ovalifolius, and orange or yellow
flowers. Herbarium specimens of the two are
easily separated: A. aurantiacus dries greyish
while A. vaginalis is either brown or retains a
greenish tinge.

Acknowledgments

I am grateful to the officers in charge of the
Australian herbaria, CANB, DNA, MEL, and
PERTH for extended loans of specimens, and
to the Director, Royal Botanic Gardens, Kew
and to the Director of The Natural History
Museum, South Kensington for allowing me
access to the collections under their care.
Members of the Legume Section at the
Herbarium, Kew were particularly generous in
their assistance.

References

Au, S.I. (1977). Alysicarpus. In Papilionaceae. Flora
of West Pakistan No. 100: 341-346

Backer, C.A. (1911). Schoolflora voor Java p. 349.
Weltevreden: Visser & Co.

Bentham, G. (1864). Alysicarpus. Flora Australiensis 2:
238-240. London: Lovell Reeve & Co.

Domin, K. (1926). Alysicarpus. In Beitrage zu Flora und
Pflanzengeographie Australiens. Biblioth. Bot.
89: 216-217.

Dunlop, C.R., Leach, G.J. & Cowie, I.D. (1995). Flora of
the Darwin Region, vol. 2. Northern Territory
Botanical Bulletin no. 20.

Endo, Y. & Ohashi, H. (1990). New distinctions between
Alysicarpus vaginalis and A. ovalifolius
(Leguminosae). Natural History Research (Chiba)
1: 43-48.

Hutchinson, J. (1964). Alysicarpus. In The Genera of
Flowering Plants 1: 482. Oxford: Clarendon
Press.

Leonard, J. (1954). Notulae Systematicae, XV.
Papilionaceae-Hedysareae Africanae. Bulletin du
Jardin Botanique de I’Etat, Bruxelles 24: 63-
106.

Pedley, L. (1999). Desmodium Desv. (Fabaceae) and related
genera in Australia: a taxonomic revision.
Austrobaileya 5: 209-261.

Pramanik, A. & Thothathri, K. (1983). Taxonomic notes
on the genus Alysicarpus Desv. (Leguminosae).
Bulletin of the Botanical Survey of India 24:
112-113.

Pramanik, A. & Thothathri, K. (1986). On the status of
Desmodiastrurn Prain (Fabaceae). Journal of the
Indian Botanical Society 65: 373-379.

Verdcourt, B. (1971). Alysicarpus. In Milne-Redhead, E.
& Polhill, R.M. (ed.): Leguminosae part 3, subf.
Papilionoideae (3). Flora of Tropical East
Africa 491-501. Crown Agents for Overseas
Governments and Administrations.

White, C.T. & Francis, W.D. (1920). Contributions to the
flora of Queensland. Botany Bulletin (Queensland
Department of Agriculture and Stock) 22: 15.

Eucalyptus broviniensis (Myrtaceae), a new critically
endangered species from south-eastern Queensland

A.R. Bean

Summary

Bean, A.R. (2001). Eucalyptus broviniensis (Myrtaceae), a new critically endangered species from south¬
eastern Queensland. Austrobaileya 6(1): 117-119. E. broviniensis, a new gum-barked species is described,
illustrated and diagnosed against the closely related E. hallii Brooker. The new species is confined to a
very small area near Mundubbera in south-east Queensland, and its conservation status is assessed as
‘critically endangered’.

Keywords: Eucalyptus, Myrtaceae, taxonomy, Eucalyptus broviniensis, Queensland flora.

A.R. Bean, Queensland Herbarium, Enviommental Protection Agency, Brisbane Botanic Gardens Mt
Coot-tha, Mt Coot-tha Road, Toowong, Queensland, 4066.

Introduction

Brooker (1975) described Eucalyptus hallii , a
restricted but common eucalypt of the sandy
coastal lowlands between Maryborough and
Bundaberg in southern Queensland. A recently
discovered, very rare, gum-barked eucalypt is
named here as Eucalyptus broviniensis. While
it grows in a vastly different habitat and is
geographically disjunct, it is clearly a very close
relative of E. hallii.

Taxonomy

Eucalyptus broviniensis A.R.Bean sp. nov.
affinis E. hallii autem fructibus
grandioribus, foliis adultis 2.5-4.5 cm
latis, glandulis oleosis paucioribus
praeditis, et pedunculis longioribus
differt. Typus: Queensland. Burnett
District: 10 km ESE of Brovinia, S.F.
132, S of Mundubbera, 7 April 1991, A.R.
Bean 11911 (holo: BRI; iso: CANB).

Small tree to 10 metres high, lignotuberous.
Bark deciduous, dull, granular, mottled light
and dark grey. Juvenile leaves alternate, ovate,
c. 10 x 5 cm, petiolate. Intermediate leaves
similar to adult leaves. Adult leaves lanceolate,
10-14.5 x 2.5-4.5 cm, alternate, leathery,
concolorous, dull; penninerved, lateral veins at
40-60° to the midrib; reticulation dense, with
sparse small island oil glands, one per areole

Accepted for publication 25 May 2001

or absent from some areoles; petioles 20-30
mm long. Inflorescences axillary, unbranched,

7- flowered; peduncles thick, more or less terete,

8- 22 mm long. Mature buds ovoid, 6-9 mm
long, 4.5-6.5 mm in diameter, sessile or with
pedicels up to 2 mm long. Hypanthium with
1-3 prominent longitudinal ribs; operculum
scar present; inner operculum hemispherical,
smooth, thick; stamens c. 4.5 mm long, white,
mostly inflexed but with a few erect, all fertile,
in 3 or 4 whorls on narrow staminophore.
Anthers oblong, versatile, dorsifixed, opening
in longitudinal slits. Style terete, with broad
conical base, stigma blunt. Ovary 3 or 4 locular,
ovules in 6-8 indistinct vertical rows. Fruits
obconical, 5.5-8.5 mm long, 7-11 mm in
diameter, disc annular, valves 3 or 4, exserted.
Seeds ellipsoidal to cuboid, 0.75-1 mm long,
not toothed, surface faintly reticulate, hilum
terminal. Chaff pale brown to yellow, mostly
cuneate. Fig. 1.

Specimens examined: Queensland. Burnett District:
Brovinia S.F., S of Mundubbera, Dec 1998, Bean 14439
(AD, BRI, NSW); S.F. 132, 10 km ESE of Brovinia, Nov
1997, Bean 12581 (BRI).

Distribution and habitat: E. broviniensis is
confined to a small area within Brovinia State
Forest south of Mundubbera. It grows on the
edges of a lateritised plateau, in heathy eucalypt
woodland with Eucalyptus exserta F.Muell.,
E. cloeziana F. Muell., E. virens Brooker &
Bean and Triodia scariosa N.T.Burb.

118

Austrobaileya 6 (1): 117 -119 (2001)

Fig. 1. Eucalyptus broviniensis. A, fruits x 2. B, fruits x 2. C, buds x 2. D, longitudinal section of bud, showing ovules,
style and stamens x 8. E, adult leaf x 1. F, intermediate leaf x 1. A-C, E, Bean 11911; B,F, Bean 12581; D, Bean 14439 (all
BRI). Del. W. Smith.

Bean, Eucalyptus broviniensis

Phenology: Flowers in summer.

Affinities: E. broviniensis is most closely
related to E. hallii, and the main differences
are detailed in the following table. Both species
have dull deciduous bark which is off-white to
grey for much of the year, shedding to bright
orange in summer. This bark type is virtually
identical in texture, colour and pattern to that
of the related species E. bancroftii (Maiden)
Maiden and the less closely related species

119

Etymology: The specific epithet refers to the
Brovinia State Forest where the species is
endemic.

Acknowledgements

I am grateful to Martin Ambrose for assistance
in the field, Les Pedley for the Latin diagnosis,
and Will Smith for the illustration.

E. broviniensis

E. hallii

Adult leaf width

2.5-4.5 cm

1.5-3 cm

Oil gland density (adult lvs)

sparse

moderately dense

Peduncle length

8-22 mm

4-8 mm

Fruit diameter

7-11 mm

5-7 mm

E. propinqua H.Deane & Maiden and E. major
(Maiden) Blakely.

The classification of E. hallii within
Eucalyptus has been somewhat difficult. It was
placed in E. ser. Subexsertae Blakely
(Chippendale 1988), and in informal E. ser.
Albae of Brooker & Kleinig (1994). However
it differs significantly from all other members
of these series by its dull, thick, granular bark,
which is bright orange when newly exposed
following decortication. Brooker (2000) created
a monotypic series {E. ser. Connexentes ) for
E. hallii, hence recognizing its distinctive suite
of characters. The present author agrees with
the removal of E. hallii (with E. broviniensis)
to a separate series, but recommends their
placement under E. sect. Liberivalvae (Blakely)
Brooker. E. ser. Connexentes has many features
in common with E. sect. Liberivalvae e.g. bark
type, leaf venation, leaf oil glands, fruit
morphology; but differs mainly in seed
morphology and operculum shape.

Conservation status: E. broviniensis is known
from only about 20 individuals at the type
locality. It is under threat from increased fire
frequency. In recent years, fires have destroyed
a number of mature stems. Applying IUCN
guidelines (Anon. 1994), a category of
‘critically endangered’ is proposed (Criteria B1,
B2, C2, D).

References

Anonymous, (1994). IUCN Red List Categories.
International Union for Conservation of Nature
and Natural Resources: Switzerland.

Brooker, M.I.H. (1975). ANew Species of Eucalyptus from
Queensland. Australian Forest Research 7: 11-
14.

-(2000). A New Classification of the Genus

Eucalyptus L’Her. (Myrtaceae). Australian
Systematic Botany 13: 79-148.

Brooker, M.I.H. & Kleinig, D.A. (1994). Field Guide to
Eucalypts, Volume 3. Sydney: Inkata Press.

Chippendale, G.M. (1988). Eucalyptus, Angophora
(Myrtaceae). In A.S. George (ed), Flora of
Australia Vol. 19. Canberra: Australian
Government Publishing Service.

Proiphys infundibularis (Amaryllidaceae), a new species from the
Townsville region of Queensland
D.L. Jones & J.L. Dowe

Summary

Jones, D.L. & Dowe, J.L. (2001). Proiphys infundibularis (Amaryllidaceae), a new species from the
Townsville region of Queensland. Austrobaileya 6 (1): 121-126. Proiphys in Australia comprises four
species, P. amboinensis (L.) Herbert, P. cunninghamii (Aiton ex Lindl.) Mabb., P alba (R.Br.) Mabb. and
P. infundibularis D.L.Jones & Dowe sp. nov., all occurring in eastern Queensland with P. alba also found
in northern Western Australia and P. cunninghamii in northern New South Wales. A key is provided for
identification of the Australian species of the genus. Three of the four species (not including P. alba ) are
illustrated. Notes on the habitat and ecology of P. infundibularis are included.

Keywords: Amaryllidaceae, Proiphys, Proiphys infundibularis, new species, tunicate bulbs, Queensland
flora.

D.L.Jones, Centre for Plant Biodiversity Research, Australian National Herbarium, GPO Box 1777,
Canberra, A.C.T., Australia, 2601.

J.L.Dowe, Department of Tropical Plant Sciences, James Cook University, Townsville, Qld, 4811,
Australia.

Introduction

Proiphys Herb, is a genus of bulbous,
seasonally deciduous geophytes distributed in
South-east Asia, Melanesia and Australia
(Backer & Bakhuizen van den Brink 1968;
Telford 1987). In Australia Proiphys is one of
the few indigenous bulbous genera to have
tunicate bulbs. Plants commonly grow in
colonies and their growth and flowering is
linked closely with the wet season, dying back
to dormant tubers during the dry season. They
are frequently cultivated as ornamentals,
especially in tropical regions.

Three species of Proiphys were previously
recorded for Australia (Telford 1987):
P. amboinensis (L.) Herb, distributed from Cape
York to near Cardwell, Queensland,
P. cunninghamii (Aiton ex Lindl.) Mabb.
occurring in south-eastern Queensland and
north-eastern New South Wales and P. alba
(R.Br.) Mabb., occurring on northern Cape
York Peninsula, Queensland and the Kimberley
region of Western Australia. A fourth Australian
species, from the Townsville region in
Queensland is described here.

Accepted for publication 6 July 2001

Materials and Methods

All species dealt with in this paper were
examined in the field and as cultivated plants
in the living collection of the Australian
National Botanic Gardens in Canberra.
Measurements cited here were made from both
living material and herbarium collections.

Taxonomy

Proiphys infundibularis D.L.Jones & Dowe,
sp. nov., P. amboinensi (L.) Herb, affinis
sed pedicellis ad 25 mm longis, floribus
infundibuliformibus, lobis corollae
anguste imbricatis, lobis coronae ad 16
mm longis, et filamenti staminum ad 30
mm longis, differt. Typus: Queensland.
North Kennedy District: 22 km W of
Townsville, lower slopes of Hervey
Range, alt. c. 250 m, 2 Oct 1998,

J. L.Dowe 521 (holo BRI; iso CANB, JCT,

K, QRS).

Bulbous herb growing in loose colonies. Bulb
ovoid, 3-6 cm long, 4-6.5 cm wide. Leaves 1-
4 per bulb, erect; petiole 15-50 cm long, 4-9
mm wide, broadly channelled distally; lamina
broadly ovate to oblong-ovate or ovate-elliptic,
10-30 cm long, 8-28 cm wide, bright green,
shiny, with veins slightly lighter, the midvein
prominent, slightly sunken; base shallowly

122

Austrobaileya6(1): 121-126 (2001)

cordate; apex acute to shortly acuminate.
Inflorescence an erect umbel; scape 35-90 cm
long, 8-14 mm wide, fleshy, bright green;
umbel 5-14-flowered; involucral bracts 2,10-
35 mm long, 4-7 mm wide, papyraceous,
whitish coloured; pedicels 15-25 mm long, c.
1.5 mm diam., fleshy, pale green. Flowers
infundibuliform, 4-5 cm long, 4-4.5 cm diam.,
white, fragrant, each lasting 2-4 days; perianth
tube 20-25 mm long, nectariferous; lobes
obovate to obovate-spathulate, 28-38 mm long,
11-16 mm wide, obtuse to apiculate, obliquely
erect, with margins shortly imbricate. Corona
14-16 mm long, the expanded filaments fused
for 8-11 mm with adjacent filaments; lobes
linear, tapered upwards, 8-10 mm long,
acuminate. Stamens 6, protruding from the
perianth; filaments 18-30 mm long, white;
anthers linear, 4.5-6.5 mm long, yellow. Style
protruding from the perianth, 50-55 mm long,
white; stigma c. 1.5 mm wide. Capsule oblong
to subovoid, 20-35 mm long, 10-20 mm wide,
greenish coloured to yellow. Fig. 1.

Additional specimens examined: Queensland. North
Kennedy District: lower slopes of Hervey Range, alt. c.
250 m, Jan 1998, Jones 15703 & Dowe (BRI, CANB);
Magnetic Island, Gustav Ck, among granite boulders, Oct
1998, Corbett (CANB).

Distribution and ecology : P. infundibularis is
known only from near-coastal locations

between Townsville and Home Hill in
Queensland, with populations of it occurring
on the eastern lower slopes of Hervey Range,
southern and northern footslopes of Mt Stuart,
eastern footslopes of Mt Elliot, Magnetic Island
(Gustav Creek) and hills to the northwest of
Ayr and Home Hill. It grows close to streams
and in small sheltered gullies, in vine thickets,
and among rocks and large granite boulders on
open sites. Soils are mainly grey gravelly loams
derived from granite. Altitude ranges from 100
to 300 m.

Phenology : Plants flower from October to
January and fruit from October to February.

Recognition : Table 1 presents a comparison of
some diagnostic morphological characters of
the Australian species of Proiphys. Proiphys
infundibularis is readily recognised by its erect
funnel-shaped flowers with imbricate corolla
lobes and long staminal filaments (Figs 1 & 2).
The leaves of P. infundibularis and
P. amboinensis are almost identical although
those of the former species are often smaller
than those of the latter. However, the flowers
of P. amboinensis are readily distinguished
from those of P. infundibularis by their
spreading corolla lobes set widely apart and
with short staminal filaments (Fig. 3). Proiphys
cunninghamii can immediately be distinguished

Table 1. Comparison of morphological characters of Proiphys alba, P. amboinensis ,
P. cunninghamii and P. infundibularis.

Character

P. alba

P. amboinensis

P. cunninghamii

P. infundibularis

leaf shape

elliptic-oblong

ovate-cordate

ovate-cordate

ovate-cordate

leaf size

10-35 x 2-11 cm

18-35 x 15-35 cm

10-25 x 8-13 cm

10-30 x 8-28 cm

involucral

bracts

20-30 mm long

30-100 mm long

15-50 mm long

10-35 mm long

umbels

10-30 flowered

5-25 flowered

5-12 flowered

5-14 flowered

pedicels

7-30 mm long

15-45 mm long

25-35 mm long

15-25 mm long

flowers

erect

erect

porrect to drooping

erect

flower size

6-24 mm long

25-40 mm long

15-18 mm long

40-50 mm long

corona lobes

5-12 mm long

7-10 mm long

15-20 mm long

14-16 mm long

filament

length

3-12 mm

7-12 mm

12-16 mm long

18-3 0 mm

Jones & Dowe, P roiphys infundibularis in the Townsville Region

123

Fig. 1. Proiphys infundibularis, Hervey Range. A. flowering plant; B. flower, front view; C. flower, side view;
D. stamen; E. corona segment. All from Jones DLJ 15703 & Dowe (CANB).

124

Austrobaileya6(1): 121-126 (2001)

from P. infundibularis by its porrect to drooping
flowers, broadly imbricate corolla lobes that are
spreading at right angles to the perianth tube
and short staminal filaments (Fig. 4). Proiphys
alba is readily distinguished from all other
Australian species of Proiphys by its greyish
green elliptic-oblong leaves.

Notes: Initially there was speculation that this
new species was a natural hybrid between
P. cunninghamii and P. amboinensis. However
neither species occurs in the Townsville region
and the new species has unique characters and
does not appear intermediate in any way.

Proiphys amboinensis appears to be most
closely allied to the new species having leaves
of similar shape but achieving larger
dimensions and with morphologically different
flowers.

Etymology: From infundibularis, Latin for
funnel-shaped, in reference to the shape of the
flowers which readily distinguishes this species
from all other Australian species of Proiphys.

Conservation Status: Uncommon, but
conserved in Magnetic Island National Park and
Cape Bowling Green National Park.

Acknowledgements

We thank Alex George for the Latin diagnosis
and Marion Garratt for technical assistance with
the drawing. Larry Corbett is thanked for
assistance with field collections, and Allen King
for first bringing this species to our attention.
Karina FitzGerald, Laurie Adams and Rogier
de Kok commented on the manuscript.

Key to Australian Species of Proiphys

1. Leaves with a broadly ovate lamina to 35 cm wide;

perianth tube more than 20 mm long.2

Leaves with an ovate or elliptic lamina to 13 cm wide;
perianth tube less than 15 mm long. 3

2. Flowers porrect, campanulate; corolla lobes widely separated;

filaments 7-12 mm long.P.amboinensis

Flowers erect, infundibuliform; corolla lobes narrowly imbricate;
filaments 18-30 mm long.P. infundibularis

3. Leaf lamina ovate, bright green; corolla lobes recurved at

right angles to the perianth tube; corona to 20 mm long. P. cunninghamii

Leaf lamina elliptic, grey green; corolla lobes obliquely erect;
corona to 12mm long

P. alba

Jones & Dowe, P roiphys infundibularis in the Townsville Region

125

Fig. 2. Proiphys infundibularis, cultivated ex Hervey Range, Queensland (DLJ 15703).

Fig. 3. Proiphys amboinensis, cultivated ex Cape York Peninsula, Queensland.

126

Austrobaileya 6 (1): 121-126(2001)

Fig. 4. Proiphys cunninghamii, cultivated ex Glenugie Peak, New South Wales.

References

Backer, C.A. & R.C.Bakhuizen van den Brink (1968).

Flora of Java 3: 139-140. Groningen, The
Netherlands: Wolters - Noordhoff N.V.

Telford, I.R. (1987) Proiphys. Liliaceae. In A.S. George
(ed.). Flora of Australia 45: 376-379. Canberra:

Australian Government Publishing Service.

Drupe - a term in search of a definition

H. Irevor Clifford and Mary E. Dettmann

Summary

Clifford H. Trevor and Dettmann, Mary E. (2001). Drupe - a term in search of a definition,
Austrobaileya 6(1): 127-131. The term drupe as defined in modem Australian Floras has several
meanings and is often employed inconsistently. Reasons for the various usages are offered and a
new definition of the term proposed.

Key words: drupe, drupaceous, exocarp, epicarp, mesocarp, endocarp

H. Trevor Clifford, Honorary Associate, Queensland Herbarium, Environmental Protection Agency,
Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road, Toowong 4066.

Mary E. Dettmann, Botany Department, The University of Queensland, 4072.

Introduction and discussion

The need for a precise definition of the term
drupe arose during a study of fossils described
as “endocarp species” and presumed to
represent the inner woody layers of drupes.
Reference to recent Australian Floras revealed
that there is a diversity of opinion as to the
definition of the term drupe. Some authors
(Carolin & Tindale 1994; McCusker 1981;
Harden 1990; Beadle 1971; Walsh & Entwistle
1994) restrict the term to the product of a
monocarpellary ovary whereas others (Stanley
&Ross 1983; Jessop & Toelken 1986; Curtis
& Morris 1975) ignore carpel number.
Furthermore, there is inconsistency in the use
of the term drupe especially by those who claim
it develops from a monocarpellary ovary.

Thus, Walsh & Entwistle (1994) qualify
their definition of drupe as monocarpellary by
citing three examples of which only the peach
is appropriate. The other two examples are the
bicarpellary olive, and Nitraria billardieri DC.
which is tricarpellary. It is ironic that in this
context the olive is an inappropriate example
of a drupe because Linnaeus (1751), when he
coined the term, presumably assumed his
readers would know that dryppa [Greek] and
drupa [Latin] were words that in classical times
referred to ‘an over-ripe wrinkled olive’ and so
did not refer to the plant by name in either his
text (p.53) or accompanying figure (Tab. VIII,
Fig. 157)

Accepted for publication 25 May 2001

It should be noted that Linnaeus (1753)
regarded both the peach and olive as
monogynous. He was not familiar with the term
carpel for it was not coined until 1817 almost
forty years after his death. Originally employed
for divisions of a fruit the term now applies to
a gynoecium in any stage of its development
(Stearn 1992).

The adjectival term drupaceous is also
widely used in Australian literature but usually
without adequate definition. Exceptions are
Beadle (1971) and Carolin & Tindale (1994) who
having accepted the drupe as monocarpellary,
defined drupaceous as applying to fruits ‘with
the structure of a drupe but derived from more
than one carpel’. An alternative approach to
the problem of incorporating carpel number into
the definition of drupe is that of Takhtajan (1991)
who proposed that such a fruit be described as
apocarpous or syncarpous according to the
ovary type from which it develops. Although
the definitions of drupe may vary in their
concept of the number of carpels involved, all
acknowledge the fruit contains a stone or stones
derived solely from the endocarp.
Monocarpellary ovaries have a single stone
enclosing one (peach) or more than one seed
(some Persoonia species). In contrast those that
are syncarpous have either several one-seeded
stones (. Leucopogon species) or a single stone
with one ( Semecarpus australiensis Engl.) or
more loculi of which one or more usually
contain seeds the remainder being empty and
sometimes compressed ( Elaeocarpus species).

128

Austrobaileya 6(1): 127-131 (2001)

Alternative names for stone are shell,
pyrene, putamen and endocarp each of which
has been more or less widely used
interchangably in the literature. Likewise both
epicarp and exocarp have been employed as
alternative terms for the skin or outermost layer
of the drupe. For example, Jessup (1985), who
has written extensively on genera occurring in
both Australia and south-east Asia has followed
Dutch terminology in referring to the outermost
layer of the drupe as the exocarp (Hou 1978).
In contrast most Australian glossaries refer to
the tissue as the epicarp. Exceptions are in the
glossaries of Stanley & Ross (1983), Walsh &
Entwistle (1994), Harden (1990) and Jessop &
Toelken (1986) who define the exocarp as the
epicarp plus mesocarp in which sense the term
is also used by some palaeobotanists including
(Reid and Chandler 1933: p.30).

Furthermore, terminological confusion
may also arise when portions of drupes are
defined in terms of other fruit types as when
Curtis & Morris (1975) defined the pyrene as
‘a nutlet of a several-seeded drupe’. The
incorporation of nutlet in the definition could
be regarded as compromising the integrity of
both fruit types, notwithstanding that in their
definition the authors made it clear that in
popular usage the term nutlets is widely applied
to fruit segments which technically are not small
nuts.

The most recent classification of fruits
(Spjut 1994) described the drupe as ‘a fleshy
pericarpium or fruit with one or more stones’.
This definition lacks any reference to either
carpel number or the origin of the woody or
bony tissue but is misleading in that it could be
taken to suggest the stone is not part of the
pericarp. In what follows there is further
confusion in that he also states ‘Drupes may
consist of one or more stones’, although it is
possible that ‘contain’ rather than ‘consist of’
was intended. Furthermore, by defining a drupe
with reference to the pericarp, Spjut has
restricted the term to fruits which develop from
superior ovaries. The pericarp, as widely
understood, refers to a fruit wall that has
developed from that of an ovary rather than an
ovary plus its associated receptacle.

By defining the term drupe solely on its

mature anatomy, Spjut (1994) has adopted a
descriptive definition of the term. In contrast,
those who relate the tissues of the fruit wall to
those of the ovary from which it developed have
an interpretive definition which, in most
instances, is unsupported by developmental
anatomical studies. This situation is unfortunate
for, as stressed by Roth (1977), the anatomical
development of relatively few fruits has been
documented.

Therefore, it is not surprising that
definitions of the term drupe based on surmize
as to the developmental pathways of their
anatomy should lead to conflicting
interpretations of their anatomy. By way of
example, the drupe as defined by Jessop &
Toelken (1986), has a pericarp which ‘ consists
of three layers (1) the epicarp or skin, (2) the
mesocarp or juicy layer and (3) the bony
endocarp or stone’. Such a definition prescribes
that the mesocarp be juicy and the endocarp
bony or woody. However, as was long ago
observed by Hill (1934) the mesocarp of the
fruit of Pleiogynium timorense (DC.) Leenh. is
differentiated into two layers, the inner of which
is thick and woody and the outer thin and fleshy
(Fig.l A). With the fruit of the closely related
Rhodosphaera rhodanthema (F.Muell.) Eng.
the outer fleshy tissue is so thin that Jessup
(1985) thought it appropriate to describe the
entire mesocarp as woody.

Although such a fruit does not match any
current definition of drupe it is only a somewhat
more extreme example of the situation which
obtains with Pleiogynium timorense. The
inadequacy of comparative anatomy for
determining fruit type is indicated by the
diversity of terms currently applied to the fruit
of Hicksbeachia pinnatifolia F.Muell. These
include the following - typical drupe (Rao
1971), indehiscent drupe (Stanley & Ross
1983), hard drupe (Beadle 1971), drupaceous
(Johnson & Briggs 1975), nut (Filla 1926) and
fleshy indehiscent follicle (Floyd 1978).

Anatomical studies of the developing
ovary wall of this species (Strohschen 1986)
have revealed that the woody layer of the fruit
derives from the inner mesocarp and not the
endocarp and so the fruit is not a drupe. Nor is
it a nut or follicle. Perhaps it was this uncertainty

Clifford & Dettman, Drupe

129

Fig. 1. Stones of Pleiogynium timorense (A) and Elaeocarpus grandis (B) in vertical section, a, axis; e, endocarp; en,
endosperm; em, embryo; ex, exocarp; f, fibres; im, inner mesocarp; om, outer mesocarp; 1, locule; s, seed. A, redrawn from
Hill (1933).

as to the nature of the fruit that led Weston
(1995) to describe but not name fruits of
Hicksbeachia, Athertonia and Catalepidia in
his Flora of Australia treatment of these genera.

A cursory survey of Australian species
whose fruits are drupes derived from
syncarpous ovaries has indicated that many
have a woody inner mesocarp in which are
encased the woody endocarps surrounding
each seed as with Pleiogynium timorense. A
notable example is Elaeocarpus grandis F.
Muell. (Fig. IB) whose stones resemble
permineralised fossils described from Oligocene
sediments of central Queensland under the
name Elaeocarpus spackmaniorum Rozefelds
(Rozefelds 1990). A further complication in
terminology arises when the term drupe is
applied to the fruit of Chrysanthemoides
monilifera (L). Norlindh which is derived from
an inferior ovary and so derived in part from a
receptacle, a situation not allowed for by any
definition of drupe.

Accordingly, there is little agreement
amongst taxonomists as to the fruit type of
C. monilifera with some describing it as a drupe
(Stanley & Ross 1986, Short 1999) others as
drupe-like (Brown 1992) or a cypsela (Carolin &
Tindale 1994). Obviously, the term drupe has
been applied to a diversity of fruits without
regard to adequate knowledge of their
developmental anatomy. Nonetheless,
inconsistency between the usage of a term and
its application in the subsequent text should be
avoided.

Conclusion

The term drupe because of its long and useful
history in descriptive taxonomic botany
deserves to be retained subject to agreement
as to its definition. Until more information is
available on the development of fleshy fruits
with a stone or stones surrounding the seeds it
would be helpful to have a definition of drupe
based solely on anatomy.

The following definition of drupe is
proposed:-

Drupe: A fruit whose wall has three
distinct tissues - an outer membranous or
chartaceous skin enclosing a zone of more or
less fleshy tissue surrounding one or more
woody, bony or parchment-like stones each of
which if fertile contains one or more seeds. The
term stone may be replaced by pyrene. Drupes
may be either apo- or syncarpous, if the latter
the stone may be unilocular or plurilocular and
with respect to the receptacle the drupe may be
superior or inferior. Plurilocular drupes may
contain several pyrenes or these may be
embedded in a woody tissue derived from the
innner mesocarp to form a single stone.

With the passage of time the stones of
plurilocular drupes may separate passively
{Elaeocarpus species) or explosively into
segments ( Petalostigma species).

In combination these characters could be
employed, to define a range of drupes, some
examples of which from the Australian flora are

130

Austrobaileya 6 (1): 127-131 (2001)

listed below.

1. Persoonia apocarpous

2. Pleiogynium syncarpous, superior,
plurilocular

3. Elaeocarpus syncarpous, superior, 2-
plurilocular

4. Semecarpus syncarpous , superior,
unilocular

5. Chrysanthemoides syncarpous, inferior,
unilocular

6. Coelospermum syncarpous, inferior, 4-
locular

The drupes of these genera differ
significantly in structure and so it could be
argued merit different names. Nonetheless, until
the developmental anatomy of fruits with flesh
surrounding a stone or stones is better known
it would be premature to replace the term drupe
with a plethora of names. Instead it would
appear preferable to retain the word drupe,
defined soley on anatomical characters, and
qualify its use with appropriate descriptive
adjectives.

Acknowledgements

The authors are grateful to Laurie Jessup for
his helpful co mm ents on an early draft of this
paper and to Jeremy Bruhl for assistance in
supplying photocopies of the relevant pages of
Philosophia Botanica.

References

Beadle, N.C.W. (1971). Students Flora of North Eastern
New South Wales. Part 1. Armidale: University
of New England.

Brown, F. A. (1992). Asteraceae. In Harden, G.J. (ed.), Flora
of New South Wales. 3. Kensington: New South
Wales University Press.

Carolin, R.C. & Tindale, M.D. (1994). Flora of the Sydney
Region. Chatswood: Reed.

Curtis, W.M. & Morris, D.I. (1975). The Student’s Flora
of Tasmania 1. 2nd Edition. Tasmania:
Government Printer.

Ding Hou (1978). Anarcardiaceae. In C.G.G.J. van Steenis
(ed.). Flora Malesiana ser. 1,8:397-548.
Djakarta: Noordhoff-Kolff.

Filla, F. (1926). Das Perikarp der Proteaceae. Flora
120:99-142.

Floyd, A.G. (1978). New South Wales rainforest trees. Pt
VII. Forestry Commission of New South Wales.
Research note, no 35.

Harden, G.W. (1990). Flora of New South Wales. 1.
Kensington: New South Wales University Press.

Hill, A.W. (1933). The method of germination of seeds
enclosed in a stony endocarp. Annals of Botany
47:873-887.

Jessop, J.P. & Toelken, H.R. (1986). Flora of South
Australia. Pt 1. Adelaide: South Australian
Government Printing Division.

Jessup, F.W. (1985). Anacardiaceae. Flora of Australia
25:170-187. Canberra: Australian Government
Publishing Service.

Johnson F.A.S. & Briggs, B.G. (1975). On the Proteaceae
- the evolution and classification of a southern
family. Botanical Journal of the Linnean
Society 70:83-182.

Finnaeus, C. (1751). Philosophia Botanica. Apud Godofr.
Kiessewetter, Stockholmiae. Repr.(1966). Fehre:
Cramer.

-(1753). Species Plantarum, Tomus 1. Impensis

Faurentii Salvii, Holmiae. Repr. [1957] , with an
Introduction by W.T. Steam. Fondon: Ray Society.

McCusker, A. (1981). Glossary In George, A.S. (ed.). Flora
of Australia Vol.l: pp. 169-198. Canberra:
Australian Government Publishing Service.

Rao, C.V. (1971). Proteaceae. New Delhi: C.S.I.R.

Reid, E.M. & Chandler, M.E.J. (1933). The London Clay
Flora. Fondon: British Museum [Nat. Hist.].

Roth, I. (1977). Fruits of angiosperms. Berlin: Gebruder
Borntraeger.

Rozefelds, A.C. (1990). A mid Tertiary r a inforest flora from
Capella, central Queensland. In Douglas, J. G. &
Christophel, D.C. (eds) Proceedings of the Third
International Organization of Palaeobotany
Symposium 1988,pp. 123-136. Melbourne: A-Z
Printers.

Short, PS. (1999). Chrysanthemoides. In Walsh, N.G.
& Entwistle, T.J. eds Flora of Victoria, Volume
4. Melbourne: Inkata Press.

Clifford & Dettman, Drupe

131

Spjut, R.W. (1994). A Systematic Treatment of Fruit
Types. Memoirs of the New York Botanical
Garden, pp. 69-70.

Stanley, T.D. & Ross, E.M. (1983). Flora of south-east
Queensland. 1. Brisbane: Queensland Department
of Primary Industries.

Stearn, W.T. (1994). Botanical Latin, ed.4. Newton Abbot:
David and Charles.

Strohschen, B. (1986). Contributions to the biology of
useful plants. 5. Anatomical studies of fruit
development and fruit classification of the monkey
nut ( Hicksbeachia pinnatifolia F.Muell.).
Angewandte Botanik 60:249-256.

Takhtajan, A. (1991). Evolutionary trends in flowering
plants. New York:Columbia University Press.

Walsh, N.G. & Entwistle, T. J. (1994). Flora of Victoria
2. Chatswood: Enkata Press.

Weston, P.H. (1995). Hicksbeachia, Athertonia,
Catalepidia, in Flora of Australia 16: 410-416.
Melbourne: CSIRO Australia.

A new species of Myriophyllum L. (Haloragaceae)
from artesian springs in Queensland
D. Halford & R.J. Fensham

Summary

Halford, D.& Fensham, R.J. (2001). A new species of Myriophyllum L. (Haloragaceae) from artesian
springs in central Queensland. Austrobaileya 6 (1): 133-137. M. artesium Halford & Fensham is described,
illustrated and diagnosed against related species. Notes on habitat and distribution are provided.

Key words: Myriophyllum , taxonomy, Australian flora, Myriophyllum artesium, Haloragaceae

D. Halford & R.J. Fensham, Queensland Herbarium, Environmental Protection Agency Mt Coot-tha
Road, Toowong, Qld 4066, Australia.

Introduction

Since 1997, the second author has been
undertaking vegetation surveys of artesian
mound springs in Queensland. In the course
of this work a number of specimens of
Myriophyllum were collected that did not match
any species previously accounted for in
Orchard’s (‘1985’ 1986) revision of Australian
Myriophyllum. Further research has since
indicated that this Myriophyllum is a distinct,
previously undescribed taxon. The new species
is described here.

Taxonomy

Myriophyllum artesium Halford & Fensham,
sp. nov., quoad habitum et folia
monomorpha et aliquantum sculpturam
fructuum aspectu M. implicati Orchard,
quoad folia opposita et caules comparate
crassos M. pedunculato J. Hooker simile,
autem ab illo foliis oppositis plerumque
latioribus oblanceolatis vel anguste
ellipticis in ambitu et caulibus florentibus
crassioribus et mericarpiis pallido-
brunneis et floribus masculis sub anthesi
pedicellis longioribus differt, et ab hoc
sepalis florum masculorum nullis et forma
sculturaque mericarpiorum et forma
foliorum facile distinguendum. Typus:
Queensland. South Kennedy District:
Doongmabulla, NW of Clermont, 3 Feb
1998, R.J. Fensham 3355 (holo: BRI).

Accepted for publication 9 March 2001

Myriophyllum sp. (Aramac B.A. Wilson
110) in Henderson (1997).

Creeping mat-forming herb 15 cm high, with
erect flowering stems unbranched, arising from
a tangled mat of prostrate rhizomatous stems.
All stems slender, c. 0.8 mm in diameter;
prostrate stems rooting freely at the nodes.
Leaves monomorphic, all opposite (rarely, a
few alternate on fast growing stems),
oblanceolate to narrowly obovate or narrowly
elliptic to elliptic, 3.3-5.5 mm long, 1.2-2.9
mm wide, obtuse or attenuate at base; margins
entire; tip blunt with a small red terminal gland;
midrib obscure. A very small (0.1 mm)
hydathode present on each side of the base of
the leaf. Plants dioecious (individual stems
either male or female). Inflorescence a simple
spike with the flowers borne singly in the axils
of the leaves. Bracteoles sexually dimorphic.
Bracteoles of male flowers ovate to broad ovate,
0.6-1 mm long, 0.4-0.6 mm long; + margins
entire; tip acute to obtuse. Bracteoles of female
flowers ovate to lanceolate, 0.6-0.8 mm long,
0.2-0.3 mm wide; margins entire; tip acute.
Male flowers 4-merous, on pedicels 2-3.5 mm
long at anthesis. Sepals absent. Petals 4,
maroon flush distally, 1.8-3 mm long, 0.9-1
mm wide, hooded, not keeled or unguiculate,
tip rounded. Stamens 8; filaments 1-1.2 mm
long; anthers linear-oblong, 1.4-1.7 mm long,
c. 0.4 mm wide, non-apiculate. Styles and
ovary absent. Female flowers 4-merous,
sessile. Sepals, petals and stamens absent.
Styles 4, sessile; stigmas white, fimbriate.

134

Austrobaileya 6(1): 133-137 (2001)

Ovary ± cubic, 0.5-0.6 mm long, 0.4-0.5 mm
wide, rounded on angles, weakly verrucose.
Fruit sessile, pale brown, ± cubic (slightly
longer than wide), 0.7-0.9 mm long, 0.7 mm
diameter. Mericarps separating freely at
maturity, + cylindrical (slightly wider near the
base). 0.6-0.8 mm long, 0.3-0.4 mm diameter,
rounded at base, slightly obliquely truncate at
apex, sparsely papillose on dorsal surface. Fig.
1 .

Additional specimens’. Queensland. Gregory South
District: Elizabeth Springs, ca 100 km SW of Boulia, Feb
1999, Fensham 3669 (BRI); Mitchell District: Edgbaston
E of Aramac, Feb 1998, Fensham 3347 (BRI); Edgbaston
Station, 40 kmNE of Aramac, Nov 1994, Wilson 110 (BRI).
Leichhardt District: Karalee, Taroom district, Jul 1996,
Fensham 2878 (BRI). Gregory South District: ca 35 km
WSW of Quilpie, Mar 1997, Connolly [AQ522039] (BRI).
Warrego District: Twomanee Plain, Granite Springs, ca
55 km SW of Eulo, Feb 1999, Fensham 3659 (BRI); Yowah
Ck, Bundoona ca 40 km NW of Eulo, Feb 1999, Fensham
3681 (BRI); loc. cit., Fensham 3685 (BRI); Werewilka, ca
70 km SW of Eulo, Feb 1999, Fensham 3658 (BRI).

Distribution and habitat : Myriophyllum
artesium occurs in central and southern

Queensland, where it has been collected from
the Boulia, Clermont - Aramac, Taroom,
Quilpie, and Eulo districts Map 1. It is generally
restricted to the wetlands associated with
springs emanating from the Great Artesian and
associated basins. There are also two
collections where the species was recorded
having colonised a creek-line fed by permanent
artesian water emanating from a running bore.

Phenology: Flowers have been recorded in
February and November; fruits have been
recorded in February.

Affinities : M. artesium resembles

M. implicatum Orchard in its habit,
monomorphic leaves and to some extent its fruit
sculpturing. However, it differs in having
opposite leaves which are oblanceolate to
narrowly obovate or narrowly elliptic to elliptic
in outline and generally broader, stouter
flowering stems, light brown mericarps, and
longer pedicels on male flowers at anthesis.
M. artesium is similar to M. pedunculatum J.
Hooker and M. amphibium Labill. in having

Table 1. Comparison of Myriophyllum artesium with similar species, M. implicatum,
M. pedunculatum and M. amphibium.

Character

M. artesium

M. amphibium

M. pedunculatum

M. implicatum

sexuality

dioecious

monoecious

monoecious

monoecious

leaf arrangement

opposite

opposite

opposite

alternate

leaf shape

oblanceolate to
narrowly obovate
or narrowly
elliptic to elliptic

oblanceolate to
obovate

linear to terete

linear

leaf width (mm)

1.2-2.9

( 1 • 5—)2—3 (—5)

0.2-0.3(-0.4)

0.6-0.7

stem diameter
(mm)

c. 0.8

c. 1

0.9-1.1

c. 0.4

pedicel length
of male flowers
at anthesis (mm)

2-3.5

<0.5

1-4.5

0.6-0.7

sepals on male
flowers

absent

present

present

absent

mericarp shape

± cylindrical

ovoid to lacrimiform

ovoid to obpyriform

cylindrical

mericarp

sculpturing

papillate

verrucose-papillose

smooth or verrucose

papillate

mericarp

colour

light brown

deep purplish black

reddish purple to black

purplish-red

Fig. 1 . Myriophyllum artesium Halford & Fensham. A. branchlet x 2. B. female flower x 40. C. male flower x 8. D. dorsal
view of mericarp x 40. E. lateral view of mericarp x 40. F. whole plant x 0.5. A-C from Fensham 3685 (BRI); D-F from
Fensham 3355 (BRI). Del. W. Smith.

136

Austrobaileya 6(1): 133-137(2001)

opposite leaves and relatively robust stems.
However, M. artesium is easily distinguished
from them by being dioecious; the absence of
sepals in male flowers; the mericarp shape and
the sculpturing on the dorsal surface. These
and other differences are summarized in Table
1 .

M. artesium will key to couplet 44 with
M. lophatum and M. austropygmaeum in
Orchard’s (1990) key. It can be distinguished
from both of these species by the papillate
sculpturing on the mericarps and shorter
pedicels on male flowers at anthesis (2.0-3.5
mm long for M. artesium ; greater than 4.0 mm
long for M. lophatum and M. austropygmaeum).

Notes: There are a suite of taxa that are endemic
to artesian springs in Queensland including
Eriocaulon carsoni, Eryngium fontanum,
Sporobolus pamelae and other undescribed
taxa. Myriophyllum artesium is the most
widespread of these species and the only one

known to also occur outside the wetland habitat
of natural springs.

Conservation status: Myriophyllum artesium
is a Queensland endemic species known from
17 spring complexes and two artesian bore
drains. There are a number of threatening
processes that have had a dramatic impact on
the natural springs of the Great Artesian Basin.
These include drastically diminished flows as
a result of pressure draw-down because of
artesian bores, excavation, eradication of
wetlands by pumping, introduction and spread
of ponded pasture species such as Brachiaria
mutica, trampling by domestic stock and
rooting by pigs. Many of these threats are
ongoing. The recommended conservation
status for this species as defined under the
Queensland Nature Conservation Act 1992 is
Vulnerable (V).

Etymology : The specific epithet artesium is
derived from the latinization of the term artesian

Halford & Fensham, Myriophyllum

137

and refers to the habitat in which this species
grows.

Acknowledgements

The authors wish to thank Will Smith for the
illustration, Peter Bostock for the map, Les
Pedley for the Latin diagnosis and Tony Orchard
for his constructive comments.

References

Henderson, R.J.F. (ed.) (1997). Queensland Vascular
Plants: Names and Distribution. Brisbane:
Queensland Department of Environment.

Orchard, A.E. (‘1985’ 1986). Myriophyllum
(Haloragaceae) in Australasia. II. The Australian
species. Brunonia 8(2): 173-291.

-(1990). Myriophyllum, Haloragaceae. In A.S.

George (ed), Flora of Australia 18: 59-84.
Canberra: Australian Government Publishing
Service.

Pappus morphology and terminology in Australian and New Zealand
thistles (Asteraceae, tribe Cardueae)

A.R. Bean

Summary

Bean, A.R. (2001). Pappus morphology and terminology in Australian and New Zealand thistles
(Asteraceae, tribe Cardueae). Austrobaileya 6 (1) 139-152. 23 pappus characters (including several
newly recognised) and five achene characters have been used to compile comprehensive morphological
descriptions for 30 species of mostly naturalised thistles occurring in Australia and New Zealand.
This represents the first detailed English account of pappus morphology for tribe Cardueae.
Some new pappus terminology for the tribe is introduced, and standardised definitions are suggested.
A key to the described species is provided, based on pappus and achene morphology only.

Key words: Asteraceae, Compositae, pappus, morphology, terminology, thistles, Australia, New Zealand,
tribe Cardueae.

A.R.Bean, Queensland Herbarium, Enviornmental Protection Agency, Brisbane Botanic Gardens Mt
Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Introduction

The morphology of pappus in the Asteraceae
is very useful taxonomically, as it offers a
wealth of characters.

Dittrich presented a comprehensive
account of fruiting morphology, covering
numerous genera of the subtribes Centaureinae
(Dittrich 1968) and Carduinae (Dittrich 1970).
He also provided keys to genera based solely
on achene and pappus characteristics. Petit
(1997), in his cladistic analysis of the whole of
the Cardueae, included further salient
observations about pappus and achene
morphology. Few other workers have paid
much attention to the diversity of carduine
pappus morphology.

Members of Asteraceae tribe Cardueae
(.sensu Bremer 1994), commonly known as the
“thistles”, are well represented in Australia
(with c. 32 species in 15 genera) and New
Zealand (with c. 21 species in 9 genera
(Garnock-Jones 1988)). These all belong to
either subtribe Carduinae or subtribe
Centaureinae. Most are naturalised and
originate from Europe. Australia has two
indigenous species ( Hemisteptia lyrata and
Stemmacantha australis ), while New Zealand
is without indigenous species.

Accepted for publication 18 May 2001

This paper, the first of its kind in English,
documents further details of pappus
morphology not recorded by Dittrich ( loc. cit.)
or Petit {loc. cit.), and provides definitions for
several terms (some new, some already
established) as they apply to tribe Cardueae. It
provides comprehensive, directly comparable
measurements and observations of pappus
characters for the great majority of Australian
and New Zealand species, as well as a few
obvious features of the achenes. A dichotomous
key to these taxa is provided using only pappus
and achene characters.

It is hoped that this paper will provide
some impetus for a more wide-ranging study
of pappus morphology, and that others will
attempt to improve the standardisation of
pappus terminology in this and other tribes of
Asteraceae.

Materials and methods

30 taxa have been included in this study.
Specimens from all major Australian herbaria
and from Christchurch (CHR) in New Zealand
have been examined (Appendix 1), using a light
microscope (x 40) with graticule.

2-5 specimens per taxon were used, at
least two pappi and two achenes were measured
per specimen, and 23 pappus characters and 5
achene characters were recorded for each
pappus and achene.

140

Austrobaileya 6 (1): 139-152 (2001)

Proposed definitions for Tribe Cardueae

A plethora of terms have been used to describe
Asteraceae pappus, viz. scale, seta, bristle,
paleaceous, capillary, scabrid, plumose,
barbellate, filamentous etc. Many of these terms
are listed and loosely defined by Jackson
(1928), but a lack of definition has resulted in
a variety of terms being used for what amounts
to the same structure, or conversely the one term
has been applied to a wide range of structures.
For this reason, appropriate pre-existing terms
have been selected here, and more explicitly
defined, to allow uniformity of interpretation,
while reinforcing common usage. Also, a
number of new terms have been introduced.

These definitions have been applied to
members of tribe Cardueae only, and it should
be realised that some of them may not be readily
applicable to other tribes in Asteraceae.

a) . The term “monomorphic pappus” is used

for a pappus where all the elements are
the same or similar, or they vary in length
or width only (e.g. Fig. 1A). The term
“dimorphic pappus” is used where there
are two distinct types or groups of pappus
elements forming an entire whorl or
partial whorl (e.g. Fig. IE). They must
be discernible by position, orientation or
major morphological trait. Discordant
elements within a whorl e.g. the few long
broad bristles in Onopordum acanthium,
are not considered sufficient to
constitute a distinct pappus type.

b) . A “pecten” (plural “pectines”) is the term

used here for the regularly or irregularly
spaced antrorse side-branches (or lateral
projections) of a bristle. They are formed
by an extension of the outer epidermal
cells of the bristle (Fig. 1G, 1H).

c) . A “bristle” is defined here as a pappus

element which is more than 5 times longer
than wide, is parallel-sided or slightly
tapered apically or at both ends, and
possesses pectines. A bristle with a length
exceeding 100 t im es its width (excluding
pectines) is called a “capillary bristle”
e.g. Fig. 2A; a bristle with a length 5-50
times its width (excluding pectines) is

called an “ensiform bristle” e.g. Fig. 2J;
any rigid or non-parallel-sided pappus
element with or without pectines is called
an “awn” e.g. Fig. 2M.

d) . A “plumose” bristle is one where the length

of the average mid-bristle pecten is 10-
30% of the (capillary) bristle length (e.g.
Fig. 2F). In plumose bristles, the pectines
are usually irregularly spaced. A “sub-
plumose” bristle is one where the length
of the average mid-bristle pecten is 2-10%
of the bristle length. A “barbellate”
bristle is one where the length of the
average mid-bristle pecten is less than 2%
of the bristle length (e.g. Fig. 2A). In
barbellate bristles, the pectines are nearly
always regularly spaced along the bristle.

e) . The term “fibrillas”, used by Petit (1997) is

adopted here. Fibrillas are short, erect,
uniseriate trichomes, often densely
clustered, and borne on or near the top of
the cylinder, forming the innermost
pappus whorls in a few species (Fig. II).

f) . The “cylinder” (equivalent to the pappus-

cover of Dittrich (1970)), is a smooth
ring-like structure composed of pericarpal
tissue, and readily detaching from the
achene (Fig. IF, II). The cylinder forms
the connate base of the pappus in many
species, and is apparently associated only
with capillary bristles. The bases of
bristles are individually obliquely inserted
onto the outer face of the cylinder,
unifying the entire pappus. In a few
species with connate pappus, the cylinder
is absent, and the bristles are merely fused
together at their bases.

List of characters used for descriptions of

Australian and New Zealand taxa

1. pappus present or absent. A few species of
Centaurea lack a pappus. In Carthamus
tinctorius, the pappus is infrequently present.

2. pappus monomorphic or dimorphic. A term
of convenience which does not indicate
relationship. For example, the dimorphic
pappus of Silybum is clearly not homologous
with that of Carthamus.

3. number of whorls in pappus.

Bean, Pappus in thistles

141

Fig. 1 . A-C. Achene and pappus of various species x 2 (A. Stemmacantha australis, B. Picnomon acarna, C. Carduus
pycnocephalus). D. achene of Hemisteptia lyrata showing persistent ensiform bristles on outer edge x 12. E. achene and
pappus of Carthamus lanatus, some bristles removed to reveal inner whorl x 4. F. basal portion of pappus of Cirsium
vulgare, showing cylinder, and bristles inserted at different points x 24. G-H. insertion of pectines. G. lateral only ( Cirsium
vulgare ) x 48. H. lateral and dorsal ( Hemisteptia lyrata) x 48.1. basal portion of pappus of Silybum marianum, showing
cylinder and fibrillas x 12. A, Bean 11348 (BRI); B, Tann s.n. (MEL); C, Bean 15601 (BRI); D, Bean 14133 (BRI); E,
Blake 5164A (BRI); F,G, Dillewaard 590 & Stanley (BRI); H, K.McDonald s.n. (BRI); I, Bean 15829 (BRI). Del. W.
S mi th.

142

Austrobaileya 6 (1): 139-152 (2001)

Fig. 2. Individual bristles of various species. A. Carduus pycnocephalus x 10. B. Carduus tenuiflorus x 12. C. Carduus
thoermeri x 6. D. Onopordum illyricum x 12. E. Picnomon acarna (inner bristle) x 6. F. Picnomon acarna (outer bristle)
x 6. G. Hemisteptia lyrata x 12. H. Centaurea melitensis x 24. I. Arctium minus x 24. J. Carthamus lanatus x 12. K.
Carthamus dentatus x 6. L. Mantisalca salmantica (bristle from outer pappus) x 24. M. Mantisalca salmantica (inner
pappus) x 24. A, Bean 15738 (BRI); B, Whinray 607 (AD); C, Bean 15914 (BRI); T>,Alcock4911 (AD); E,F, Symon 2095
(AD); G, Jones 2908 (CANB); H, Bean 15890 (BRI); I, Grace s.n. (HO); J, Blake 5164A (BRI); K, Howlett s.n. (MEL);
L,M, Bean 15637 (BRI). Del. W. Smith.

143

Bean, Pappus in thistles

4. pappus element type (bristles or awns).

5. bristle type (capillary or ensiform).

6. number of bristles/awns per pappus.

7. pappus deciduous from achene or persistent.

8. pappus connate (bristles fused at their bases
into a ring) or free (bristles individually
attached to achene).

9. cylinder present or absent.

10. length of cylinder.

11. bristle whorls uniform or varying in length.
When bristle whorls vary in length, it is
apparently always the inner whorls that are
longer.

12.1-3 bristles within a whorl longer and
thicker than remainder. This character was
noted for Onopordum by Tamamschyan
(1998).

13. bristle length (if varying, longer or longest
whorls have been measured).

14. bristle width (excluding pectines). This is
the width measured about half way along
the bristle.

15. bristles expanded at base or not. This
character applies only to the innermost
whorl where an expanded spathulate bristle
base occurs in some species (Dittrich 1968).
The bristles otherwise taper gradually from
proximal to distal end.

16. bristles not fasciculate or some bristles
fasciculate. In Onopordum acaulon, some
bristle pairs (or triplets) are laterally fused
for one-fifth to one-half of their length.

17. pecten type - plumose, sub-plumose or
barbellate.

18. bristles plumose throughout on all whorls,
or with barbellate apical section (Fig. 2E,
2F). The inner whorls of a pappus bearing
plumose capillary bristles often have a
barbellate apical section, where the pectines
are abruptly reduced. This section is often
twisted or bent relative to the rest of the

bristle, and may be slightly thickened. This
character has been noted for Cirsium b y
Davis and Parris (1975) and by Charadze
(1998).

19. length of barbellate apical section.

20. bristle apex. Generally acute; some
ensiform bristles may have a toothed or
obtuse apices.

21. pecten insertion. In some species, pectines
are inserted only laterally onto the bristles
(Fig. 1G). Some species also have pectines
inserted onto the dorsal surface of the bristle
(Fig. 1H). Rarely, pectines are inserted onto
the ventral surface. Ventral pectines were
noted for Cynara by Dittrich (1970) and
Wiklund (1992).

22. pecten length. This is often relatively
uniform, but where there is variability,
pectines occurring about midway along
bristle have been measured.

23. fibrillas absent or present (Fig. II). Fibrillas
occur in only a few species. They were
noted for Silybum by Dittrich (1970).

24. achene length. Measured along medial axis.

25. achene maximum width.

26. achene ribbing (longitudinally ribbed or
transversely rugose or smooth).

27. achene indumentum. Glabrous or with
sparse ciliate hairs.

28. apical rim. Absent in some species;
otherwise entire, dentate or sinuate.

Pappus and achene descriptions for
Australian and New Zealand taxa

Taxa are arranged in systematic order,
following Petit (1997). These descriptions are
based entirely on measurements of the
specimens cited in Appendix 1. None has been
augmented by descriptions given in Flora
treatments or revisionary works.

(i) Subtribe Carduinae

144

Austrobaileya 6 (1): 139-152 (2001)

1. Hemisteptia lyrata (Bunge) Fisch. &

C.A.Mey.

Pappus dimorphic. Inner pappus
comprising one whorl of capillary
bristles, 14-18 in number, deciduous,
basally connate, cylinder absent; bristles
± uniform in length, 7-10 mm long, 0.05-
0.1mm wide, not expanded at base,
plumose throughout, barbellate apical
section absent, apex acute; pectines borne
laterally and dorsally, 1.3-2.2mm long at
mid-bristle. Outer pappus comprising one
part-whorl of 7-10 sub-plumose ensiform
bristles, persistent, free, each 0.2-0.3 mm
long and c. 0.05 mm wide, apex obtuse.
Achenes 2.0-2.8mm long, 0.8-1.1mm
wide, glabrous, conspicuously longitudinally
ribbed, apical rim entire. (Fig. ID, 1H,
2G).

2. Carduus nutans L.

Pappus monomorphic, comprising 5-7
whorls of capillary bristles, 130-169 in
number, deciduous, connate to a cylinder
0.5-0.6 mm long; bristles ± uniform in
length, 15-20mm long, 0.05-0.1 mm
wide, not expanded at base, barbellate
throughout, apex acute; pectines borne
laterally only, 0.04-0.1 mm long; fibrillas
absent or very few, not forming a whorl,
each c. 0.6mm long. Achenes 3.3-3.8mm
long, 1.3-1.7 mm wide, glabrous, smooth,
apical rim entire.

3. Carduus pycnocephalus L.

Pappus monomorphic, comprising 6-8
whorls of capillary bristles, 167-190 in
number, deciduous, connate to a cylinder
0.6-0.7 mm long; bristles + uniform in
length, 14-16mm long, 0.04-0.08 mm
wide, not expanded at base, barbellate
throughout, apex acute; pectines borne
laterally only, 0.1-0.13 mm long; fibrillas
absent. Achenes 4.8-5.3mm long, 1.8-2
mm wide, glabrous, smooth, apical rim
entire. (Fig 1C, 2A).

4. Carduus tenuiflorus Curtis

Pappus monomorphic, comprising 6-8
whorls of capillary bristles, 147-195 in
number, deciduous, connate to a cylinder
c. 0.6 mm long; bristles + uniform in
length, 11-13mm long, 0.04-0.08 mm

wide, not expanded at base, barbellate
throughout, apex acute; pectines borne
laterally only, 0.08-0.13 mm long;
fibrillas absent. Achenes 4.0-4.3mm
long, 1.7-2.0 mm wide, glabrous,
smooth, apical rim entire. (Fig. 2B).

5. Carduus thoermeri Weinm.

Pappus monomorphic, comprising 5-7
whorls of capillary bristles, 125-170 in
number, deciduous, connate to a cylinder
0.4-0.6 mm long; bristles ± uniform in
length, 17-23mm long, 0.05-0.08 mm
wide, not expanded at base, barbellate
throughout, apex acute; pectines borne
laterally only, 0.03-0.08 mm long;
fibrillas absent. Achenes 3.5-4.1mm
long, 1.4-1.7 mm wide, glabrous,
smooth, apical rim entire. (Fig. 2C).

6. Silybum marianum (L.) Gaertn.

Pappus dimorphic. Inner pappus
comprising many fibrillas, 0.6-1.2mm
long. Outer pappus comprising 6-7
whorls of capillary bristles, 126-167 in
number, deciduous, connate to a cylinder
0.4-0.8 mm long; bristles ± uniform in
length, 15-20mm long, 0.06-0.18 mm
wide, not expanded at base, barbellate
throughout, apex acute; pectines borne
laterally only, 0.08-0.13 mm long.
Achenes 5.6-6.7mm long, 2.8-3.4 mm
wide, glabrous, smooth, apical rim entire.
(Fig. II).

7. Picnomon acarna (L.) Cass.

Pappus monomorphic, comprising 4-5
whorls of capillary bristles, 95-109 in
number, deciduous, connate to a cylinder
0.4-0.6 mm long; bristles ± uniform in
length, 13-17mm long, 0.07-0.1 mm
wide, not expanded at base, plumose
throughout on outer whorls, otherwise
with barbellate apical section 0.8-2.2 mm
long, apex acute; pectines borne laterally
only, 3.0-5.2mm long at mid-bristle;
fibrillas absent. Achenes 4.6-5.2 mm
long, 2.4-2.6 mm wide, glabrous,
smooth, apical rim entire. (Fig. IB, 2E,
2F).

8. Cirsium arvense (L.) Scop. var. arvense

Pappus monomorphic, comprising 3-4

Bean, Pappus in thistles

145

whorls of capillary bristles, 57-80 in
number, deciduous, connate to a cylinder
0.2-0.3 mm long; bristles ± uniform in
length, 15-22 mm long, 0.05-0.1 mm
wide, not expanded at base, plumose
throughout on outer whorls, otherwise
with barbellate apical section 0.3-1.0 mm
long, apex acute; pectines borne laterally
only, 2.5-5.8 mm long at mid-bristle;
fibrillas absent. Mature achenes not seen.

9. Cirsium brevistylum Cronquist

Pappus monomorphic, comprising 2-4
whorls of capillary bristles, 47-62 in
number, deciduous, connate to a cylinder
0.25-0.3 mm long; bristles ± uniform in
length, 20-25 mm long, 0.06-0.1 mm
wide, not expanded at base, plumose
throughout on outermost whorl,
otherwise with barbellate apical section
0.8-2.0 mm long, apex acute; pectines
borne laterally only, 3.0-5.1 mm long at
mid-bristle; fibrillas absent. Achenes 3.7-
4.0 mm long, 1.4-1.5 mm wide, smooth,
glabrous, apical rim entire.

10. Cirsium palustre (L.) Scop.

Pappus monomorphic, comprising 2-3
whorls of capillary bristles, 35-44 in
number, deciduous, connate to a cylinder
0.2-0.3 mm long; bristles varying in
length with inner exceeding outer, longer
ones 8-11 mm long, 0.05-0.09 mm wide,
not expanded at base, plumose throughout
on outermost whorl, otherwise with
barbellate apical section 0.6-1.7 mm
long, apex acute; pectines borne laterally
only, 0.9-2.3 mm long at mid-bristle;
fibrillas absent. Achenes 3.4-3.7 mm
long, 1.1-1.2 mm wide, smooth, glabrous,
apical rim entire.

11. Cirsium vulgare (Savi) Ten.

Pappus monomorphic, comprising 2-4
whorls of capillary bristles, 38-85 in
number, deciduous, connate to a cylinder
0.3-0.4 mm long; bristles ± uniform in
length, 19-28 mm long, 0.07-0.12mm
wide, not expanded at base, plumose
throughout on outer whorls, otherwise
with barbellate apical section 0.5-1.8 mm
long, apex acute; pectines borne laterally
only, 3-6 mm long at mid-bristle; fibrillas

absent. Achenes 3.5—4.0 mm long, 1.5-1.8
mm wide, smooth, glabrous, apical rim
entire. (Fig. IF, 1G)

12. Ptilostemon afer (Jacq.) Greuter

Pappus monomorphic, comprising 3-4
whorls of capillary bristles, 51-61 in
number, deciduous, basally connate,
cylinder absent; bristles ± uniform in
length, 14-17 mm long, 0.1-0.17 mm
wide, not expanded at base, plumose
throughout, apex acute; pectines borne
laterally only, 1.5-3 mm long at mid¬
bristle; fibrillas absent. Achenes 4.0-4.5
mm long, 2.7-2.9 mm wide, smooth,
glabrous, apical rim absent.

13. Cynara cardunculus subsp.

flavescens Wiklund

Pappus monomorphic, comprising 4-5
whorls of capillary bristles, 66-93 in
number, deciduous, connate to a cylinder
0.3-0.4mm long; bristles varying in
length with inner exceeding outer, longer
ones 26-35mm long, 0.15-0.25mm wide,
not expanded at base, plumose throughout
on outermost whorl, otherwise with
barbellate apical section 3-10 mm long,
apex acute; pectines borne laterally only,
or on inner whorl, pectines borne laterally
and ventrally, 2.5-4.5 mm long at mid¬
bristle; fibrillas absent. Achenes 4.8-5.8
mm long, 3.1-3.8 mm wide, smooth,
glabrous, apical rim absent.

14. Onopordum acanthium L.

Pappus monomorphic, comprising 2
whorls of capillary bristles, 73-119 in
number, deciduous, basally connate,
cylinder absent; bristles varying in length
with inner exceeding outer, usually
including 1-3 stout capillary bristles (9-
11 mm long, 0.15-0.2mm wide) on inner
whorl; of remainder, longer ones 6.0-
8.0mm long, 0.05-0.08mm wide, not
expanded at base, barbellate throughout,
apex acute; pectines borne laterally and
dorsally, 0.07-0.12 mm long. Achenes
4.1-5 mm long, 2.0-2.7 mm wide,
strongly quadrangular, conspicuously
transversely rugulose, glabrous, apical
rim absent.

146

Austrobaileya 6 (1): 139-152 (2001)

15. Onopordum acaulon L.

Pappus monomorphic, comprising 4-5
whorls of capillary bristles, 135-164 in
number, deciduous, basally connate,
cylinder absent; bristles ± uniform in
length, 20-25mm long, 0.07-0.15mm
wide, not expanded at base, a small
proportion fasciculate, i.e. laterally fused
for one-fifth to one-half of their length,
barbellate throughout, apex acute;
pectines borne laterally only, or in some
bristles laterally, dorsally and ventrally,
0.15-0.3 mm long. Achenes 4.0-4.5 mm
long, 2.1-2.5 mm wide, strongly
quadrangular, transversely rugulose,
glabrous, apical rim absent.

16. Onopordum illyricum L.

Pappus monomorphic, comprising 2
whorls of capillary bristles, 61-93 in
number, deciduous, basally connate,
cylinder absent; bristles varying in length
with inner exceeding outer, occasionally
including 1 stout capillary bristle (c.
13mm long, 0.18mm wide) on inner
whorl; of remainder, longer ones 6-11
mm long, 0.05-0.12mm wide, not
expanded at base, barbellate throughout,
apex acute; pectines borne laterally and
dorsally, 0.1-0.2 mm long. Achenes 4.1-
6.5 mm long, 1.8-3.2 mm wide, strongly
quadrangular, conspicuously transversely
rugulose, glabrous, apical rim absent.
(Fig. 2D).

17. Onopordum tauricum Willd.

Pappus monomorphic, comprising 2
whorls of capillary bristles, 55-84 in
number, deciduous, basally connate,
cylinder absent; bristles varying in length
with inner exceeding outer, occasionally
including 1 stout capillary bristle (9-
11 mm long, 0.1-0.13 mm wide) on inner
whorl; of remainder, longer ones 6-8.5
mm long, 0.04-0.08mm wide, barbellate
throughout, apex acute; pectines borne
laterally and dorsally, 0.1-0.15 mm long.
Achenes 4.1-4.7 mm long, 1.6-2.4 mm
wide, strongly quadrangular,
conspicuously transversely rugulose,
glabrous, apical rim absent.

18. Arctium lappa L.

Pappus monomorphic, comprising 2-3

whorls of ensiform bristles, 80-150 in
number, deciduous, all free; bristles
varying in length with inner exceeding
outer, longer ones 2.5-3.5 mm long,
0.07-0.1 mm wide, sub-plumose
throughout, apex acute; pectines borne
laterally and dorsally, 0.05-0.1 mm long.
Achenes 6.9-7.3 mm long, 2.5-3.0 mm
wide, faintly longitudinally striate and
rugulose, glabrous, apical rim obscure, or
conspicuous and coarsely sinuate.

19. Arctium minus (Hill) Bernh.

Pappus monomorphic, comprising 2-3
whorls of ensiform bristles, 60-100 in
number, deciduous, all free; bristles
varying in length with inner exceeding
outer, longer ones 2.0-3.3 mm long,
0.07-0.1mm wide, sub-plumose
throughout, apex acute; pectines borne
laterally and dorsally, 0.05-0.1mm long.
Achenes 5.2-6 mm long, 2.2-2.7 mm
wide, faintly longitudinally striate and
rugulose, glabrous, apical rim obscure, or
conspicuous and coarsely sinuate. (Fig.
21 ).

(ii) Subtribe Centaureinae

20. Carthamus dentatus Vahl

Pappus dimorphic. Inner pappus
comprising one whorl of connivent
barbellate bristles, 12-13 in number,
persistent, free, bristles 2.5-14mm long
(usually with 2-6 much shorter than
remainder). Outer pappus comprising 4-
7 whorls of ensiform bristles, 115-155
in number, persistent, free; bristles
varying in length with inner exceeding
outer, longest ones 11-14 mm long, all
0.5-0.9 mm wide, barbellate throughout,
apex acute (inner) to serrate (outer);
pectines borne laterally and dorsally,
0.06-0.12mm long. Achenes 4.5-5.8mm
long, 4.4-5mm wide, angular, not ribbed,
glabrous, apical rim entire. (Fig. 2K).

21. Carthamus lanatus L.

Pappus dimorphic. Inner pappus
comprising one whorl of connivent
barbellate bristles, 11-15 in number,
persistent, free, bristles 0.2-1.5mm long
or 4-11mm long. Outer pappus

Bean, Pappus in thistles

147

comprising 4-7 whorls of ensiform
bristles, 82-108 in number, persistent,
free; bristles varying in length with inner
exceeding outer, longest ones 6.5-11 mm
long, all 0.4-0.6 mm wide, barbellate
throughout, apex acute (inner) to serrate
(outer); pectines borne laterally and
dorsally, 0.08-0.12mm long. Achenes

4.5-5.7mm long, 3.5-4.2mm wide,
angular, not ribbed, glabrous, apical rim
entire to dentate. (Fig. IE, 2J).

22. Carthamus leucocaulos Sm.

Pappus dimorphic. Inner pappus
comprising one whorl of connivent sub-
plumose bristles, 10-13 in number,
persistent, free, bristles 1.0-4mm long
(usually with one or more much longer
than remainder). Outer pappus
comprising 4-6 whorls of ensiform
bristles, 77-95 in number, persistent, free;
bristles varying in length with inner
exceeding outer, longest ones 4.5-5.3 mm
long, all 0.3-0.5 mm wide, barbellate
throughout, apex acute (inner) to serrate
(outer); pectines borne laterally and
dorsally, 0.04-0.08mm long. Achenes 3-
3.5mm long, 2.5-2.8mm wide, angular,
not ribbed, glabrous, apical rim entire.

23. Carthamus tinctorius L.

Pappus absent from most achenes. When
present, pappus dimorphic. Inner pappus
of one partial-whorl of sub-plumose
ensiform bristles, 7-10 in number,
persistent, free, bristles 1.3-2.6mm long.
Outer pappus comprising 4-6 partial-
whorls of ensiform bristles, 13-50 in
number, persistent, free; bristles varying
in length with inner exceeding outer,
longest ones 4.5-6 mm long, all 0.2-0.3
mm wide, barbellate throughout, apex
acute (inner) to serrate (outer); pectines
borne laterally and dorsally, 0.07-
0.12mm long. Achenes 6.5-8mm long,
4.1-4.7mm wide, angular, not ribbed,
glabrous, apical rim entire.

24. Stemmacantha australis (Gaudich.)

Dittrich

Pappus monomorphic, comprising 4-5
whorls of capillary bristles, 89-112 in
number, deciduous, connate to a cylinder

0.3-0.4 mm long; bristles varying in length
with outer whorl somewhat shorter than
remainder, longest ones 18-23 mm long,
0.07-0.15 mm wide (outer whorls), 0.35-
0.4mm wide at base (inner whorl), sub-
plumose throughout, apex acute; pectines
borne laterally and dorsally, 0.8-1.6 mm
long at mid-bristle; fibrillas absent.
Achenes 6.5-8.4 mm long, 2.5-3.1 mm
wide, longitudinally ribbed, glabrous,
apical rim dentate. (Fig. 1A).

25. Mantisalca salmantica (L.) Briq. & Cavill.

Pappus dimorphic. Inner pappus
comprising a single tapering awn,
persistent, annular at base, 3-3.5mm long.
Outer pappus comprising 4-6 whorls of
ensiform bristles, 77-107 in number,
persistent, free; bristles varying in length
with inner exceeding outer, longest ones

2.5- 3.3 mm long, all 0.07-0.1 mm wide,
barbellate throughout, apex acute (inner)
to obtuse (outer); pectines borne laterally
and dorsally, 0.04-0.08mm long. Achenes

3.5- 3.9mm long, 1.3-1.5 mm wide,
lacunose, not ribbed, glabrous, apical rim
shallowly dentate. (Fig 2L, 2M).

26. Centaurea calcitrapa L.

Pappus absent. Achenes 2.7-2.8 mm
long, 1.7-1.8 mm wide, smooth, very
sparsely hairy, apical rim absent.

27. Centaurea jacea L.

Pappus absent. Achenes 2.7-2.9 mm
long, 1.2-1.3 mm wide, faintly
longitudinally ribbed, very sparsely hairy,
apical rim entire.

28. Centaurea melitensis L.

Pappus dimorphic. Inner pappus
comprising one whorl of erect or
connivent barbellate ensiform bristles,
16-17 in number, persistent, free, 0.5-
0.7mm long. Outer pappus comprising 5-
7 whorls of ensiform bristles, 100-150
in number, persistent, free; bristles
varying in length with inner exceeding
outer, longest ones 2.4-3.5 mm long, all
0.05-0.08 mm wide, barbellate
throughout, apex acute (inner) to obtuse
(outer); pectines borne laterally and
dorsally, c. 0.03mm long. Achenes 2.6-

148

Austrobaileya 6 (1): 139-152 (2001)

2.8 mm long, 1.2-1.3 mm wide, smooth,
very sparsely hairy, apical rim minutely
dentate. (Fig. 2H).

29. Centaurea solstitialis L.

Pappus dimorphic. Inner pappus
comprising one whorl of erect or
connivent barbellate ensiform bristles,
17-19 in number, persistent, free, 0.6-
1 .Omm-long. Outer pappus comprising 4-
5 whorls of ensiform bristles, 70-100 in
number, persistent, free; bristles varying
in length with inner exceeding outer,
longest ones 3.3-3.9 mm long, all 0.04-
0.08 mm wide, barbellate throughout,
apex acute; pectines borne laterally and
dorsally, c. 0.04mm long. Achenes 2.3-
2.6mm long, 1.2-1.3mm wide, smooth,
very sparsely hairy, apical rim minutely
dentate.

30. Cnicus benedictus L.

Pappus dimorphic. Inner whorl
comprising one whorl of erect awns, 10
in number, persistent, free, annular at
base, 2.3-3.0mm long, with coarse
irregular trichomes along their length.
Outer pappus comprising one whorl of
erect or spreading awns, 10 in number,
persistent, free, 10-11mm long, apex
acute; pectines absent. Achenes 7.2-
8.5mm long, 3.0-3.1 mm wide,
prominently longitudinally ribbed,
glabrous, apical rim conspicuously
dentate.

Discussion

Bentham (1873) compared bristle width to
pecten length to define the terms ‘plumose’ and
‘barbellate’, and this has been followed in
recent times e.g. Dittrich (1968), Danin (1975),
Anderberg (1991). In my opinion, these terms
relate more to the bristle length, so that a bristle
half as long as another, and with pectines half
as long, is equally plumose, regardless of bristle
width. The terms ‘scabrid, scabrous, rough’ and
‘denticulate’ are not here considered to be
useful, as they are not significantly different
from ‘barbellate’.

While I have not attempted to define a
scale, they are generally considered to have a

small length-breadth ratio, are often scarious,
and frequently broad-based, or with a subulate
apex. Scales have not been observed in
Australian or New Zealand thistles, but are
reported for a few south-west Asian genera e.g.
Xeranthemum, Siebera, Chardinia which were
placed in subtribe Carlininae by Dittrich
(1996), but transferred to subtribe Carduinae
by Petit (1997).

Features of the pappus have been widely
used by botanists to help distinguish or
determine species or genera of Asteraceae, and
the diversity of pappus morphology apparent
in Australian and New Zealand species
demonstrates their high value, but detailed
studies of pappus morphology have been
neglected. By contrast, there are many recent
examples of studies dealing with other
morphological aspects, such as pollen
(Blackmore 1981, Feuer and Tomb 1977),
corolla (Jeffrey 1977), ligules (Baagoe 1977),
style (Jones 1976) and fruits (Kallersjo 1985).

Within Australian and New Zealand
Cardueae at least, there is a considerable
morphological disjunction between capillary
and ensiform bristles (as defined here), so that
there is never any doubt about the application
of these terms. Furthermore, most of those
species with capillary bristles fall readily into
either “plumose” or “barbellate” as defined
here. In fact, only one species ( Stemmacantha
australis ) classifies as “sub-plumose”.For
ensiform bristles, only the “barbellate” and
“sub-plumose” categories apply, and there is
apparently a continuum of forms. Plumose
ensiform bristles are not known.

It is clear from this study that pappus
morphology is more or less fixed at the species
level. At the generic level, the situation is less
clear. Dittrich (1968) suggested that pappus
characteristics often vary between species of
the same genus, and hence have limited value.
There is certainly infrageneric variation in
Onopordum pappus, where some species have
barbellate bristles and some are reported to have
plumose bristles (Feinbrun-Dothan 1978).
Nevertheless, these infrageneric differences in
pappus morphology may point to other
correlated differences in other parts of the plant,
and hence be of phylogenetic significance.

Bean, Pappus in thistles

149

It would be foolish to make sweeping
statements about generic relationships based on
this study, because of the small proportion of
taxa sampled. However, a few observations
may be made:

Carduus, Silybum, Cirsium, Ptilostemon
and Picnomon all have pectines that are borne
laterally only; the last three genera have
plumose bristles. Cynara also seems closely
related, though it has pectines borne ventrally
on the bristles of the inner whorl.

Ptilostemon afer has very closely spaced
pectines compared to the other plumose species
in this study. The number of pectines per bristle
may prove to be another useful pappus
character.

Hemisteptia stands apart from all other
Australian and New Zealand Cardueae , though
it is obviously close to the Asian genus
Saussurea DC.

The Australian representative of
Stemmacantha, with its monomorphic pappus,
deciduous capillary bristles connate to a
cylinder, and the basal detachment area of the
achene, seems closer to subtribe Carduinae than
to subtribe Centaurinae. Its broad-based inner
bristles, while distinctive, do not constitute a
separate pappus type.

Centaurea, Carthamus and Mantisalca
have much in common (persistent barbellate
ensiform bristles, varying in length), and both
Centaurea and Carthamus have a connivent
inner pappus whorl.

The pappus of Onopordum acaulon is
strikingly different from the other three
Australian Onopordum spp. Its bristles are
about twice as long as the other species, and a
small proportion of them are fasciculate,
although this latter feature is not obvious to the
casual observer.

Key to Australian and New Zealand thistle species, based on pappus and achenes

1. Pappus present.2

Pappus absent.25

2. Pappus elements predominantly capillary bristles.3

Pappus elements predominantly ensiform bristles or awns.16

3. Bristles plumose throughout or almost throughout.4

Bristles barbellate or sub-plumose throughout.10

4. Bristles plumose throughout, cylinder absent.5

Apical section of inner bristles barbellate.6

5. Capillary bristles 14-18 in a single whorl.1. Hemisteptia lyrata

Capillary bristles 51-61, in 3-4 whorls.12. Ptilostemon afer

6. Barbellate apical section of inner bristles 3-10 mm long

.13. Cynara cardunculus ssp. flavescens

Barbellate apical section of inner bristles 0.3-2.2 mm long.7

7. Cylinder 0.4-0.6mm long; achenes 4.6-5.2 mm long.7. Picnomon acarna

Cylinder 0.2-0.4mm long; achenes 3.4-4 mm long.8

8. Bristles 8-11 mm long; pectines 0.9-2.3 mm long.10. Cirsium palustre

Bristles 15-28 mm long; pectines 2.5-6 mm long.9

9. Cylinder 0.2-0.3 mm long
Cylinder 0.3-0.4 mm long

8. Cirsium arvense var. arvense, 9. C. brevistylum
. 11. Cirsium vulgare

lO.Bristles sub-plumose
Bristles barbellate....

24. Stemmacantha australis
. 11

150

11 .

12 .

13.

14.

15.

16

17.

18.

19.

20 .

21 .

22 .

23.

24.

25.

Austrobaileya 6 (1): 139-152 (2001)

Pappus dimorphic, inner pappus fibrillate.6. Silybum marianum

Pappus monomorphic, fibrillas rare or absent.12

Bristles varying in length, in two whorls only

.14. Onopordum acanthium, 16. O. illyricum, 17 . O. tauricum

Bristles + uniform in length, in 4-7 whorls.13

Cylinder absent, some bristles fasciculate,

achenes quadrangular. 15. Onopordum acaulon

Cylinder present, bristles not fasciculate,

achenes smooth and rounded.14

Bristles 15-23 mm long; pectines 0.04-0.1 mm long

.2. Carduus nutans, 5. C. thoermeri

Bristles 11-16 mm long; pectines 0.08-0.13 mm long.15

Bristles 11-13 mm long; achenes 4-4.3 mm long.4. Carduus tenuiflorus

Bristles 14-16 mm long; achenes 4.8-5.3 mm long.3. Carduus pycnocephalus

Pappus comprising two whorls of awns, 10 awns per whorl.30. Cnicus benedictus

Pappus wholly or predominantly of ensiform bristles.17

Bristles deciduous.18

Bristles persistent.19

Achenes 6.9-7.3 mm long.18. Arctium lappa

Achenes 5.2-6 mm long. 19. Arctium minus

Inner pappus comprising a single rigid awn.25. Mantisalca salmantica

Inner pappus comprising several connivent bristles.20

Pappus whorls partial only; achenes 6.5-8 mm long.23. Carthamus tinctorius

Pappus whorls complete; achenes 2.6-5.8 mm long.21

Inner pappus of 16-19 bristles; longest bristles of outer pappus

2.4- 3.9 mm long.22

Inner pappus of 10-15 bristles; longest bristles of outer pappus

4.5- 14 mm long.23

Longest bristles 2.4-3.5 mm long; achenes 2.6-2.8 mm long.28. Centaurea melitensis

Longest bristles 3.3-3.9 mm long; achenes 23-2.6 mm long.29. Centaurea solstitialis

Longest bristles 4.5-5.3 mm long; achenes 3-3.5 mm long.22. Carthamus leucocaulos

Longest bristles 6.5-14 mm long; achenes 4.5-5.8 mm long.24

Outer pappus of 115-155 bristles, longest ones 11-14 mm long .. 20. Carthamus dentatus
Outer pappus of 82-108 bristles, longest ones 6.5-11 mm long.25

Achenes 6.5-8 mm long.23. Carthamus tinctorius

Achenes 2.3-2.8 mm long.26

26. Achenes 1.7-1.8 mm wide, smooth.

Achenes 1.2-1.3 mm wide, faintly longitudinally ribbed

26. Centaurea calcitrapa
.27. Centaurea jacea

Bean, Pappus in thistles

151

Acknowledgements

I am grateful to Philip Sharpe for translating
one of Dittrich’s key to genera, to Ailsa Holland
for providing comments on the manuscript, to
the Directors of AD, CANB, CHR, DNA, HO,
MEL, NSW and PERTH for the loan of thistle
specimens, and to Will Smith for the
illustrations. This work was completed with the
assistance of a grant from the Australian
Biological Resources Study.

References

Anderberg, A. A. (1991). Taxonomy and phylogeny of
the tribe Gnaphalieae (Asteraceae). Opera
Botanica 104: 1-187.

Baagoe, J. (1977). Microcharacters in the ligules of the
Compositae. In, The Biology and Chemistry of the
Compositae, Vol. 1. Eds V.H.Heywood,
J.B.Harbome & B.L.Turner. Academic Press:
London.

Bentham, G. (1873). Notes on the classification, history,
and geographical distribution of the Compositae.
Journal of the Linnean Society, Botany 13:

335-577.

Blackmore, S. (1981). Palynology and intergeneric
relationships in subtribe Hyoseridinae
(Compositae: Lactuceae). Botanical Journal of the
Linnean Society 82: 1-13.

Bremer, K. (1994). Asteraceae - Cladistics and
Classification. Timber Press: Oregon.

Charadze, A.L. (1998). Cirsium, in GPanigrahi (ed.), Flora
of the U.S.S.R. Volume XXVIII, English
translation. Bishen Singh Mahendra Pal Singh &
Koeltz Scientific Books: Dehra Dun, India.

Danin, A. (1975). Onopordum, in P.H.Davis (ed.), Flora of
Turkey, Volume 5, Compositae. University Press:
Edinburgh.

Davis, PH. & Parris, B.S. (1975). Cirsium in P.H.Davis
(ed.), Flora of Turkey, Volume 5, Compositae.
University Press: Edinburgh.

Dittrich, M. (1968). Morphologische Untersuchungen an
den Friichten der Subtribus Cardueae-
Centaureinae (Compositae). Willdenowia 5:
67-107.

-(1970). Morphologische und anatomische

Untersuchungen an Friichten der Carduinae
(Compositae) I. Morphologischer Teil. Candollea
25: 45-67.

-(1996). Die Bedeutung morphologischer und

anatomischer Achanen-Merkmale fur die
Systematik der Tribus Echinopeae Cass, und
Carlineae Cass. Boissiera 51: 9-102.

Feinburun-Dothan, N. (1978). Flora Palaestina, Volume
3. Israel Academy of Sciences and Humanities:
Jerusalem.

Feuer, S. & Tomb, A.S. (1977). Pollen morphology and
detailed structure of family Compositae, tribe
Cichorieae, 2: subtribe Microseridinae. American
Journal of Botany 64: 230-45.

Garnock-Jones, P.J. (1988). Asteraceae - Cichorioideae,
in C.J.Webb, W.R.Sykes & P.J.Garnock-Jones
(eds), Flora of New Zealand, Volume 4. Botany
Division CSIR: Christchurch.

Jackson, B.D. (1928). A Glossary of Botanic Terms, 4th
edition. Gerald Duckworth & Co. Ltd.: London.

Jeffrey, C. (1977). Corolla forms in Compositae - some
evolutionary and taxonomic speculations. In, The
Biology and Chemistry of the Compositae, Vol. 1.
Eds V.H.Heywood, J.B.Harbome & B.L.Turner.
London: Academic Press.

Jones, A.G. (1976). Observations on the shape and exposure
of style branches in the Astereae (Compositae).
American Journal of Botany 63: 259-62.

Kallerjo, M. (1985). Fruit structure and generic
delimitation of Athanasia L. and related South
African genera (Asteraceae-Anthemideae). Nordic
Journal of Botany 5: 527-42.

Petit, D.P. (1997). Generic interrelationships of the
Cardueae (Compositae): a cladistic analysis of
morphological data. Plant Systematics and
Evolution 207: 173-203.

Tamamschyan, S.G. (1998). Onopordum, in GPanigrahi
(ed.), Flora of the U.S.S.R. Volume XXVIII,
English translation. Bishen Singh Mahendra Pal
Singh & Koeltz Scientific Books: Dehra Dun,
India.

Wiklund, A. (1992). The genus Cynara L. (Asteraceae-
Cardueae), Botanical Journal of the Linnean
Society 109: 57-123.

152

Austrobaileya 6 (1): 139-152 (2001)

Appendix 1 - Voucher specimens used for
measurements

Arctium

A. lappa : Welsh (HO), Drake (HO); A. minus:
Anon. 10 (HO), Curtis (HO), Grace (HO),
Lampinen 5449 (BRI).

Carduus

C. nutans : Bloomfield s.n. (MEL), Morris s.n.
(HO), Doing s.n. 18/4/67 (CANB);
C. pycnocephalus: Bean 15672 (BRI),
Bean 15738 (BRI); C. tenuiflorus : Vonow
98 (AD), Whinray 607 (AD), Whibley
3723 (AD); C. thoermeri : Bean 15589
(BRI), Bean 15892 (BRI), Bean 15914
(BRI).

Carthamus

C. dentatus: Gray & Medway 7083 (BRI),
Howlett s.n. (MEL); C. lanatus: Adams
s.n. (BRI), Blake 5164A (BRI), Alcock
5556 (AD), Coveny 16368 & Whalen
(BRI); C. leucocaulos: Hunt 1872 (AD),
Overton 767 (AD); C. tinctorius : Moore
s.n. (BRI), A.E.Smith s.n. (BRI).

Centaurea

C. calcitrapa : R.V.Smith 68/29 (BRI); C.jacea:
Bird s.n. 29/4/89 (BRI); C. melitensis:
Bean 15890 (BRI), L.S.Smith 3047
(BRI); C. solstitialis: Bean 14774 (BRI),
Everist 4286 (BRI).

Cirsium

C. arvense var. arvense: Curtis s.n. (HO), Reid
2167 (MEL), Morrison s.n. (CANB),
Scarlett 82-15 (CANB); C. brevistylum:
Worsley s.n. 11/1/72 (CHR), Anon. [CHR
5714B] (CHR); C. palustre : Holm-
Nielsen 330 et al. (BRI); McLaren s.n.
(CHR). C. vulgare: Buchanan 750 (HO),
Cummings 97 (CANB), Jobson 4575 et
al. (BRI).

Cnicus

C. benedictus: Hoffman s.n. (CANB),
Clydesdale s.n. (BRI).

Cynara

C. cardunculus ssp .flavescens: Ising s.n. (AD),
Wilson 3600 (AD).

Hemisteptia

H. lyrata: K.McDonald s.n. (BRI), Jones 2908
(CANB).

Mantisalca

M. salmantica: Bean 15637 (BRI); Mansbridge
s.n. (BRI).

Onopordum

O. acanthium: Cleland s.n. (AD), Ward s.n.

(HO), Fitzpatrick ON09 (CANB),
Michael 29/12/69 (CANB); O. acaulon:
H.W.Andrew (AD), Bartsch 5 (AD),
Beauglehole 87638 & Huebner (MEL),
Symon 1886 (AD); O. illyricum: Alcock
4977 ( AD), Colwill ON017 (CANB),
Somneville ON018 (CANB), Nelligan
ON019 (CANB); O. tauricum : Alcock
6151 (AD), Gebert s.n. (MEL).

Picnomon

P acarna : Symon 2095 (AD), Amtsberg s.n.
(AD), Copley 1106 (AD).

Ptilostemon

P. afer. E.GSmith s.n. (CHR).

Silybum

S. marianum : Bean 15605 (BRI), Bean 15806
(BRI), Bean 15829 (BRI).

Stemmacantha

S. australis: Fensham 1396 (BRI), Bean 11348
(BRI), Bartlam s.n. (BRI).

Cycas cupida (Cycadaceae),
a new species from central Queensland

Paul I. Forster

Summary

Forster, Paul I. Cycas cupida (Cycadaceae), a new species from central Queensland. Austrobaileya
6 (1): 153-160 (2001). The new species Cycas cupida P.I.Forst. from sandstone substrates in
central Queensland is described (together with illustrations) and compared to C. couttsiana K.D.Hill
and C. desolata P.I.Forst. A key to the species of Cycas series Caimsianosae K.D.Hill is presented.

Key words: Cycas, Cycas cupida, Cycas couttsiana, Cycas desolata, Cycas series Caimsianosae

P.I. Forster, Queensland Herbarium, Enviommental Protection Agency, Brisbane Botanic Gardens Mt
Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Introduction

In early November 2000, pressed material and
photographs were sent to me by Tim Perry, then
of the Environmental Protection Agency at
Townsville, of a blue-leaved Cycas from
sandstone substrates well to the south of
Charters Towers in central Queensland. This
population was highly disjunct from other
populations of Cycas and of immediate interest
given the patterns of speciation and taxon rarity
in some of the Australian species. Taxonomic
accounts of the Australian species of Cycas L.
have been presented by Hill (1996, 1998)
wherein some 27 species were enumerated
together with identification keys. This pressed
material differed in a number of ways from
previously described species, although it was
clearly placed in Cycas series Caimsianosae
K.D.Hill which includes the five species
C. cairnsiana F.Muell., C. couttsiana K.D.Hill,
C. desolata P.I.Forst., C. ophiolitica K.D.Hill
and C. platyphylla K.D.Hill and is characterised
by “Basal pinnae not gradually reducing to
spines; pinnae slightly to strongly revolute,
strongly glaucous; hypodermis fully
continuous, often with more than 1 layer of
cells. Cataphylls hard and pungent or soft.
Pollen cones medium, ovoid, orange or brown”
(Hill 1998).

Investigation of this population in
December 2000 enabled collection of a range
of fertile material and documentation of

Accepted for publication 16 July 2001

morphological variation from many individuals
that is never possible only from herbarium
collections. The new species is described in this
paper as Cycas cupida and differentiated from
other taxa in Cycas series Caimsianosae.

Taxonomy

Cycas cupida P.I.Forst., species nova, affinis
speciebus ambabus Cycadi desolatae
P.I.Forst. et C. couttsianae K.D.Hill sed
ab ilia numero majore foliolis in foliis
maturis (174-240 non 90-136), foliolis
e basi foliis non angustis et 90-100%
latitudinis maximae (non 70-80%),
cataphyllis multo longioribus (70-95 mm
non 30-45 mm) dense tomentosis per
longitudinem integram (adversum basin
non nisi), strobilis microsporangiatis
latioribus (10-16 cm diam. non 8-9 cm)
microsporophyllo spina apicali longiore
instructo (5-12 mm non 3-4 mm), et ab
hac caulibus crassioribus (20-30 cm
diam. non 14-20 cm), foliolis e basi foliis
non angustis et 90-100% latitudinis
maximae (non 70-80%), augmento novo
indumento ferrugineo instmcto (adversum
canum), strobilis microsporangiatus
magnioribus (25-33 x 10-16 cm non 15-
20 x 7-9 cm) et megasporophyllis
ferrugineis (adversum grisea) lamina late
triangulari (adversum anguste
triangularem)differens. Typus:
Queensland. South Kennedy District:
Terrace Range, 11 December 2000,

154

Austrobaileya 6(1): 153-160 (2001)

P.I.Forster PIF26540B (male plant) &
R.Booth (holo: BRI [6 sheets and
carpological]).

Stems to 6 m high (rarely multiheaded), 20-30
cm thick. Leaves 72-130 cm long, straight or
sometimes somewhat wavy towards the apex,
strongly keeled in cross-section, the opposing
leaflets inserted at 30-50 degrees to the rhachis,
the rhachis usually terminated by paired
leaflets; petiole 12-26 cm long, 10-11 mm
diameter, strongly blue-grey tomentose,
otherwise green-grey beneath, with 12-26 short
teeth c. 2 mm long and spaced 10-15 mm apart,
rarely spineless. Leaflets 174-240 per leaf,
evenly spaced in lower half of leaf, more
interleaved and wavy in the upper half of leaf;
median leaflets at 40-50 degrees to the rhachis,
150-240 mm long, 5-7 mm wide; glaucous
blue-pruinose, green-grey when covering
removed; more or less flat in cross-section,
decurrent for 0-4 mm, not narrowed at base of
frond (90-100 % of maximum width), midrib
slightly raised above, prominent below. New
growth densely tomentose with ferruginous-
brown indumentum, glabrescent. Cataphylls
pungent, linear, 70-95 mm long, 3-5 mm wide,
densely tomentose for entire length with
ferruginous-brown indumentum. Microsporangiate
cones elongate-ovoid, 25-33 cm long, 10-16
mm diameter, with dense ferruginous-brown
indumentum; microsporophyll fertile zone 14-
48 mm long, 8-15 mm wide; sterile zone 8-19
mm long; apical spine antrorsely recurved, 5-
12 mm long. Megasporophylls 22-30 cm long,
when young densely ferruginous-brown
tomentose, glabrescent and then glaucous blue-
grey; ovules 4 to 6 (usually 4); lamina broadly
triangular, 45-60 mm long, 25-35 mm wide,
strongly dentate with well developed teeth to 3
mm long, apical spine 15-18 mm long. Seeds
ovoid, 35-40 mm long, 32-35 mm diameter,
strongly grey-blue pruinose, green beneath wax
covering, becoming yellow-purple when ripe;
sarcotesta 3-4 mm thick. Figs. 1-6.

Specimens examined: Queensland. South Kennedy
District: Terrace Range, Dec 2000, Forster PIF26540A
(BRI); ditto, PIF26540C (BRI); ditto, PIF26543 (BRI);
ditto, PIF26544 (BRI); ditto, Nov 2000, Perry s.n. (BRI).

Notes: Cycas cupida clearly belongs in Cycas
series Cairnsianosae as defined by Hill (1998).
The new species appears to be closest
morphologically to both C. couttsiana and
C. desolata, both of which are also the closest
geographically, albeit over 200 km to the north.
Cycas cupida differs from C. desolata (Forster
1995) in the greater number of leaflets in mature
leaves (174-240 versus 90-136), the leaflets
at the base of the leaf not narrowed and 90-
100% of maximum width (versus 70-80%), the
much longer cataphylls 70-95 mm (versus 30-
45 mm) that are densely tomentose for the entire
length (versus base only), the thicker
microsporangiate cones (10-16 cm diameter
versus 8-9 cm) with a longer apical spine on
the microsporophyll (5-12 mm long versus 3-
4 mm). Cycas cupida differs from C. couttsiana
(Hill 1998) in thicker stems (20-30 cm diameter
versus 14-20 cm), the leaflets at the base of
the leaf not narrowed and 90-100% of
maximum width (versus 60-80%), the new
growth with ferruginous-brown indumentum
(versus grey-white), the longer and larger
microsporangiate cones (25-33 x 10-16 cm
versus 15-20 x 7-9 cm) and the ferruginous-
brown megasporophylls (versus grey) with a
broadly-triangular lamina (versus narrow-
triangular).

All species in Cycas series Cairnsianosae
have blue-coloured leaves to some extent. In
some species such as C. cairnsiana, this colour
is intense and held for the life of the leaf. Other
species such as C. platyphylla and C. ophiolitica
are blue-coloured when the leaves are young,
however this colour is soon lost. Both
C. couttsiana and C. desolata are somewhat
intermediate in this feature. By comparison
C. cupida appears to retain the blue coloration,
but not to quite the intensity of C. cairnsiana.

The six species in Cycas series
Cairnsianosae may be distinguished in the
following key. This key is designed for leaves
from mature plants and will not work on
juvenile material. It is important to collect
cataphylls and to note indumentum colour on
young leaves and the cataphylls if possible.

Forster, Cycas cupida

155

Fig. 1. Cycas cupida. A. median part of leaf where leaflets are at maximum length, x 0.6. B. cross-section of a median
leaflet, x 4. C. fruiting megasporophyll. x 0.4. D. immature megasporophyll. x 0.6. E. cataphyll. x 1. F. microsporophyll
from near cone apex, x 3. G. & H. microsporophyll from near cone middle, x 3. A-D from Forster PIF26540A (BRI); E
& F from Forster PIF26540B (BRI); G & H from Forster PIF26543 (BRI). Del. W. Smith.

156

Austrobaileya 6 (1): 153 -160 (2001)

Fig. 2. Cycas cupida habit.

Key to the species of Cycas series Cairnsianosae

1. Median leaflets in leaf 2-3 (4) mm wide, margins revolute. C. cairnsiana

Median leaflets in leaf 4-7.5 mm wide, margins recurved to more or less flat.2

2. Basal leaflets in leaf not narrowed (80-100% of median leaflets width).C. cupida

Basal leaflets in leaf narrowed (55-80% of median leaflets width).3

3. New growth with white or grey trichomes; median leaflets 6-7.5 mm wide;.

cataphylls narrow-triangular.C. couttsiana

New growth with orange-brown trichomes; median leaflets 3.5-6 mm wide;
cataphylls linear.4

4. Median leaflets 6-7.5 mm wide; cataphylls soft.C. ophiolitica

Median leaflets 3.8-6 mm wide; cataphylls pungent.5

5 Short teeth at base of leaf petiole present; leaflets more or less straight,

120-260 per leaf; median leaflets 90-170 mm long; indumentum covering

entire cataphyll.C. platyphylla

Short teeth at base of leaf petiole absent; leaflets antrorse, 90-136 per leaf,
median leaflets 180-210 mm long; indumentum only at base of cataphyll.C. desolata

Forster, Cycas cupida

157

Fig. 3. Cycas cupida habit of large plant.

Distribution and habitat: Cycas cupida is
restricted to the Terrace Range, south of
Charters Towers. The Terrace Range is an
isolated low-lying sandstone range with
occasional low outcrops and clifflines of
exposed sandstone rock. The vegetation (Fig.
6) comprises an open woodland dominated by
Corymbia clarksoniana, C. dallachiana and
Eucalyptus crebra with a midstorey of Acacia
bidwillii, A. salicina, Eremophila mitchellii,
Lysiphyllum hookeri, Planchonia careya and
Terminalia oblongata. The cycads are widely
spread in this habitat, being most abundant on
the low hills, but can also be found concentrated
in gullies and creeklines where they may occur

on sandstone colluvium. Occasionally plants
grow in crevices on low sandstone cliffs. The
habitat is seasonally dry and very hot in the
summer months from reflected heat from the
exposed rocks.

Conservation status: This new species is
abundant at the type locality and is widely
distributed throughout the Terrace Range. An
accurate population size is as yet unknown, but
thousands of plants exist with no immediate
threats to the population from land clearing or
agriculture. Under the IUCN Red List
Categories (Anon. 1994), C. cupida fulfils the
criterion of ‘vulnerable’ under Category D2. It

158

Austrobaileya 6(1): 153-160 (2001)

Fig. 4. Cycas cupida female plant with megasporophylls.

is estimated that the area of suitable habitat
comprises less than 60 km 2 and although
diffuse, there is only one large population of
this cycad. Blue-leaved cycads are desirous to
cycad collectors (cf. Forster 1999) and it is
likely that interest in C. cupida will be intense.

assistance; the property owner for access; Peter
Bostock for translation of the diagnosis into
Latin and comments on the manuscript; and
Will Smith for the illustrations.

References

Etymology: The specific epithet is derived
from the Latin cupidus (desirous) and alludes
to the desirability of this cycad to collectors.

Acknowledgements

Thanks to Tim Perry and Step Lawler for
bringing this species to my attention and for
guidance in the field; Ron Booth for field

Anonymous, (1994). IUCN Red List Categories.
Intematonal Union for Conservation of Nature
and Natural Resources: Switzerland.

Forster, RI. (1995). Cycas desolata (Cycadaceae), a new
species from north Queensland. Austrobaileya
4: 345-351.

-(1999). Six blue cycads of desire. In

K.A.W.Williams, Native Plants of Queensland,
4: 388-389. North Ipswich: K.A.W.Williams.

Forster, Cycas cupida

159

Hill, K.D. (1996). A taxonomic revision of the genus
Cycas (Cycadaceae) in Australia. Telopea 7:

1-64.

(1998). Cycadophyta. In P.McCarthy (ed.).
Flora of Australia 48: 597-661. Melbourne:
CSIRO Publications.

160

Austrobaileya 6(1): 153-160 (2001)

Fig. 6. Habitat of Cycas cupida.

New combinations and a new name
in Australian Sapotaceae
L.W. Jessup

Summary

Jessup, Laurence W. (2001). New combinations and anew name in Australian Sapotaceae. Austrobaileya
6(1): 161-163. Five new combinations made as follows, Niemeyera whitei for Amorphospermum whitei
Aubrev, Pouteria queenslandica for Planchonella queenslandica P.Royen, Pouteria myrsinifolia for
Sersalisia myrsinifolia F.Muell., Pouteria myrsinodendron for Chrysophyllum myrsinodendron F.Muell.,
Pouteria cotinifolia var. pubescens for Planchonella cotinifolia var. pubescens P.Royen, while Pouteria
asterocarpon is a new combination made at a new rank for Planchonella pohlmaniana var. asterocarpon
P.Royen, and Pouteria pearsoniorum is a new name provided for Planchonella macrocarpa P.Royen.

Keywords: Niemeyera, Pouteria, Sapotaceae, Australia.

L.W. Jessup, Queensland Herbarium, Enviornmental Protection Agency, Brisbane Botanic Gardens, Mt
Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4006, Australia.

Introduction

In preparing an account of Sapotaceae for the
Flora of Australia, I have encountered several
taxa requiring new combinations or new names.
Pennington (1991) included Amorphospermum
F.Muell. as congeneric with Niemeyera
F.Muell. rejecting the distinction between them
proposed by Aubreville (1962) based on
thickness of the seed coat. One Australian taxon
still requires a new combination under
Niemeyera and this is provided below. In the
same publication, Pennington provided a
detailed discussion of the reasons why
Planchonella Pierre should be included in
Pouteria Aublet and his conclusions are
accepted here. Several taxa currently
recognised at species or varietal rank under
Planchonella have not yet been formally placed
under Pouteria. To remedy this situation the
necessary new combinations are made and new
name provided as follows.

Niemeyera whitei (Aubrev.) Jessup, comb,
nov.

Amorphospermum whitei Aubrev.,
Adansonia ser. 2,5:23, t.2 (1965). Type:
New South Wales, Whian Whian State
Forest, May 1945 C.T. White 13043;
(holo: BRI; iso: BRI, fide G.P. Guymer,

Accepted for publication 13 March 2001

Australian Systematic Botany Society
Newsletter 31:11-13(1982)).

Pouteria queenslandica (P. Royen) Jessup,
comb. nov.

Planchonella queenslandica P.Royen,
Blumea 8: 341, 430 (1957), fig. 32.
Beccariella queenslandica (P.Royen)
Aubrev., Adansonia ser. 2, 3: 335
(1964). Type: Queensland, [South
Kennedy District]: Eungella Mts, H.H.
Haines 136 Q (holo: K).

[Planchonella laurifolia auct. non (A.Rich.)
Pierre, W.D. Francis, Australian Rain-forest
Trees 3 rd edition p.350-353. Canberra,
Australian Government Publishing
Service; P. Royen, Blumea 8: 340, Fig.
31 (1957)].

Note: This tree, found in the rainforests of
eastern Queensland and New South Wales, was
incorrectly known for many years as
Planchonella laurifolia (A.Rich.) Pierre. This
name, based on Sersalisia laurifolia A.Rich.,
applies to a Northern Territory species now
known as Pouteria richardii (F.Muell.) Baehni
(as Pouteria laurifolia (Gomes) Radik, applies
to a different species). The type specimen of
the former name was amongst several
specimens given by Charles Fraser to Captain
Jules Dumont d’Urville following Fraser’s

162

Austrobaileya 6 (1): 161-163 (2001)

expedition to Moreton Bay in 1828. Dumont
d’Urville visited Sydney sometime during the
voyage of the Astrolabe between 1826 and
1829. Unfortunately at least two specimens
amongst this gift to the French visitors did not
originate from Moreton Bay and were
incorrectly included as part of Fraser’s Moreton
Bay collection. These specimens ultimately
became types of two of Achille Richard’s
species’ names, Leucocarpum obscurum
A.Rich. (Celastraceae) and Sersalisia laurifolia
A.Rich. (Sapotaceae). Replicates of these
specimens were independently sent to Kew
with the locality of origin noted on the label as
Melville Island and the communicator as
Charles Fraser. There is no evidence that Fraser
ever went to Melville Island. The British naval
settlement of Fort Dundas was established on
Melville Island in 1824 not long after the
establishment of Port Essington. Fraser sent a
convict gardener, John Richardson, to be in
charge of the garden for the new settlement of
Port Essington (Barker & Barker, 1990). While
there is no evidence that Richardson collected
and dispatched specimens to Fraser,
Richardson’s convict status probably assured
his name would not appear with any such
specimens, their origin being attributed only to
Fraser. Bentham (1863) noted the labelling
error evident in the collections of Denhamia
obscura (A.Rich.)Meisn. ex Walp. ( Leucocarpum
obscurum A.Rich.) but he apparently
overlooked it when he examined and described,
some years later, the Sapotaceae specimens as
Achras laurifolia (A.Rich.)Benth.(Bentham,
1869).

Pouteria myrsinifolia (F.Muell.) Jessup,
comb. nov.

Sersalisia myrsinifolia F.Muell. ‘ Sarsalisia\
Fragm. 5:165 (1866). Type: [Queensland.
Moreton District] : Brisbane River, “20
feet high found on the sides of river,
flowers greenish”, W.Hill [MEF1058112]
(MEF, lecto here selected).

Selected additional specimens seen: Queensland: Sea
Range, Dec 1855, F. Mueller (MEL 1515834) (MEL);
Moreton District: Moreton Bay, W. Hill (MEL1058111,
MEL1058113) (MEL);Breakfast Creek, Moreton Bay, Jul
1843, L. Leichhardt (MEL1058110) (MEL, P); Brisbane
River in 1829, Fraser (K). New South Wales: Sydney
woods, Paris Exhibition, n. 27 and n. 40, 1855, Moore (P).

Note: Practically all herbarium specimens
previously identified as Planchonella
myrsinoides (A. Cunn. ex Benth.) S.T. Blake
ex W.D. Francis or Pouteria myrsinoides (A.
Cunn. ex Benth.) Baehni belong here. As the
lectotype of Achras myrsinoides, basionym of
both of the above two names, as selected by
PS. Green (1986) is taxonomically Pouteria
cotinifolia var. pubescens (see below), another
name is needed for the other plants Bentham
included under that name. As lectotypified here,
Mueller’s name Sersalisia myrsinifolia of 1866
is the earliest legitimate name applying to them
hence a new combination under Pouteria is
required. Thus Pouteria myrsinifolia becomes
the correct name for the plant called for many
years Planchonella myrsinoides. I have
examined at K and MEF all the specimens cited
by Bentham (1869) and all except the
Cunningham collection from Rodd’s Bay are
of P. myrsinifolia.

Pouteria cotinifolia var. pubescens (P.Royen)
Jessup, comb. nov.

Planchonella cotinifolia var. pubescens P.
Royen, Blumea 8: 296, 428 (1957).
Type: Queensland. Leichhardt District:
Duaringa, mixed softwood forest, 23
Nov 1943, C.T. White 12462; holo: L;
iso: BRI.

Pouteria myrsinoides (A. Cunn. ex Benth.)
Baehni, Candollea 9: 303 (1942);
Achras myrsinoides A. Cunn. ex Benth.,
FI. Austral. 4 (16 Dec. 1868) 283.
Sideroxylon myrsinoides (A. Cunn. ex
Benth.) F.Muell., Systematic Census of
Australian Plants 92 (1883);

Planchonella myrsinoides (A. Cunn. ex
Benth.) S.T. Blake ex W.D. Francis,
Australian Rain Forest Trees ed. 2, 358
(1951); Xantolis myrsinoides (A. Cunn.
ex Benth.) Baehni, Boissiera 11: 23, fig
17 (1965); Sapota myrsinoides (F.
Muell. ex Benth.) Radik, ex Holle,
These Erlangen 17 (1892); fide C.
Baehni, Candollea 9: 303 (1942);
Sersalisia myrsinoides (A.Cunn. ex
Benth.) Domin, Biblioth. Bot. 89: 508
(1928). Type: [Queensland. Port Curtis
District]: Rodd’s Bay, A. Cunningham
123 (lecto: K, fide PS. Green (1986));

Jessup, Sapotaceae name

163

Note: These changes affect the name of the New
Caledonian and Lord Howe Island plant called
Pouteria myrsinoides subsp. reticulata (Baill.)
P.S.Green. I hesitate to make a new combination
for this plant under either P. cotinifolia or
P myrsinifolia as I have not examined the New
Caledonian specimens to verify Green’s
conclusion that they and the Lord Howe Island
specimens represent the same subspecies
(Green 1990). Until further work on the
relationships between these Australian and
neighbouring island plants is done it seems best
to apply the name Pouteria howeana (F.Muell.)
Baehni to the Lord Howe Island and New
Caledonian plants.

Pouteria pearsoniorum Jessup, nom. nov.

Planchonella macrocarpa P. Royen,
Blumea 8: 320,429, fig. 27 (1957), non
Pouteria macrocarpa (Martius) D.
Dietrich, Synopsis Plantarum 1:431
(1839) et non Pouteria macrocarpa
(Huber) A.Ducke, Bol. Teen. Inst.
Agron. N. No.8. 11 (1946), nom.
illegit.Type: Queensland. Cook District:
Kaban, Pearson brothers s.n. (holo:
BRI; iso: BRI, L).

Etymology: The epithet honours the Pearson
brothers, collectors of the type specimen.

Note: B. Hyland (pers. comm.) has suggested
that the locality Kaban is more likely to be a
postal address than the place where the type
was actually collected.

Pouteria myrsinodendron (F.Muell.) Jessup,
comb. nov.

Chrysophyllum myrsinodendron F.Muell.,
Fragm. 6:178 (1868). Type: [Queensland.
North Kennedy District]: Herbert
River, J. DallachyP (holo: MEL; iso:
BM ex herb. Hance).

Planchonella obovoidea H.J.Lam, Bull.
Jard. Bot. Buitenzorg ser.3, 7: 207, fig.
56 (1925). Type: [Malesia]. Kai Is.,
Jaheri 134 (holo: ?BO n.v. fide Lam,
op. cit.; iso: L). Pouteria obovoidea
(H.J. Lam) Baehni, Candollea 9: 412
(1942).

Note: Bentham (1869, p.283) referred to
Mueller’s name as representing a possible form
of Pouteria obovata (R.Br.) Baehni which he
accepted as Achras obovata (R.Br.) F.Muell.
ex Benth. This may have obscured the plant’s
true identity from subsequent authors.

Pouteria asterocarpon (P.Royen) Jessup,
comb, et stat. nov.

Planchonellapohlmaniana var. asterocarpon
P.Royen, Blumea 8: 395, 432, fig. 42 e-f.
(1957). Type: Queensland. [Cook
District]: Atherton district, Kemp s.n.
[AQ22582] (holo: BRI).

References

Aubreville, A. (1962) Notes sur les Sapotacees de la
Nouvelle Caledonie. Adansonia n.s. 2: 172.

Barker, R.M & W.R. Barker. (1990) Botanical
contributions overlooked: the role and recognition
of collectors, horticulturalists, explorers and others
in the early documentation of the Australian flora,
in History of Systematic Botany in Australasia
RS.Short ed. Melbourne: Australian Systematic
Botany Society Inc.

Bentham, G. (1863). Celastrineae. In Flora Australiensis
1: 401-402. London: L. Reeve & Co.

-(1869) Sapotaceae. In Flora Australiensis 4: 282

& 283. London: L. Reeve & Co.

Green, P.S. (1986). Notes relating to the floras of Norfolk
and Lord Howe Islands, II. J. Arnold Arbor. 67:
118.

-(1990). Notes relating to the floras of Norfolk and

Lord Howe Islands, III. Kew Bull. 45(2): 251-
252.

Pennington, T.D., (1991). The genera of Sapotaceae.
Richmond: Royal Botanic Gardens, Kew and
New York: The New York Botanical Garden.

New species of Livistona R. Br. (Arecaceae) from
north Queensland and Papua New Guinea

John L. Dowe and Anders S. Barfod

Summary

Dowe, J. L. & Barfod, A. S. (2001). New species of Livistona R. Br. (Arecaceae) from north
Queensland and New Guinea. Austrobaileya 6 (1): 165-174. Livistona concinna, L. surru and L.
tothur are described as new. Livistona concinna is endemic in north Queensland, and L. surru and
L. tothur are endemic in Papua New Guinea. Each species is illustrated.

Keywords: Arecaceae; Livistona - Queensland, New Guinea; Livistona concinna; Livistona surru;
Livistona tothur.

John L. Dowe, Tropical Plant Sciences, James Cook University, Townsville, Qld 4811, Australia

Anders S. Barfod, Department of Systematic Botany, University of Aarhus, Nordlandsvej 68, DK-8240,
Risskov, Denmark

Introduction

Livistona R.Br., with about 33 species, has an
unusually widespread distribution for a palm
genus, occurring from the Horn of Africa and
Yemen, through east Asia to Japan and the
Bonin Islands, throughout south-east Asia and
Malesia to as far east as the Solomon Islands,
and in the north, east and centre of Australia.
Australia has about 18 species. New Guinea
has about seven species, two of which also
occur in northern Australia across Torres Strait,
and others which are endemics that are most
closely related to species in the Philippines,
eastern Indonesia and the Solomon Islands.

The genus Livistona in Australia was
treated by Rodd (1998) as having 16 species
and three subspecies. Dowe and Jones (in press,
Flora of Australia) have recognised 18 species
following minor adjustment to Rodd’s
taxonomy. The genus in New Guinea has had
no recent treatment, although it has received
attention in the past from Beccari (1877,1921)
and Burret (1935, 1939, 1941) who both
described new taxa within it. Rodd (1998)
recently combined one of Beccari’s and three
of Burret’s taxa with those species that are
shared with Australia.

Accepted for publication 20 July 2001

In Australia, many species of Livistona
are ubiquitous in some areas, forming large
colonies that dominate their local environment.
Only a few species are considered rare. In New
Guinea, the distribution ranges are fragmented
and locally most species are rare occuring in
scattered groups on isolated ridges and
mountain slopes.

The new taxon from north Queensland
described here has had some previous
recognition as a distinct species. Jones (1984)
listed it as “ Livistona sp. Cooktown”, and
suggested that it was “similar in many respects
to Livistona benthamii F.M. Bailey, but the
leaves are light green on both surfaces and the
lamina has a broad central area where the
segments are fused”. This ‘tag’ name was
accepted by Irvine (1984) who placed the taxon
in a group that included L. drudei F.Muell. ex
Drude as well as L. benthamii , while Tucker’s
(1988) informal account provided convincing
evidence for the taxon’s distinctiveness.

The two new taxa from New Guinea
described here were discovered during field¬
work conducted within the framework of the
Palms of New Guinea (PONG) project that is
currently being coordinated by the Royal
Botanic Gardens, Kew, England. This project
covers the island of New Guinea, and its aim is
to publish an account of the palm flora of this

166

Austrobaileya 6 (1): 165-174 (2001)

island in the near future. Both of the above new
taxa appear to be confined to Papua New
Guinea despite extensive field-work having
been carried out in Papua (formerly Irian Jaya)
as part of the PONG project. An indication of
the lack of recognition of any diversity within
Livistona in New Guinea was provided by Essig
(1977) who quoted unpublished notes by
Moore stating: “....the species of Livistona in
New Guinea are difficult to separate from
related species in Australia and the Solomon
Islands. Possibly there are no true endemics of
Livistona in New Guinea.”. However, Essig and
Young (1981) described from the West Sepik
Province “specimens of a large Livistona
bearing red fruit that we had seen from the
helicopter, but found only a few sterile
individuals”, and Hay (1984) mentioned an
uncollected taxon in Madang Province:
“.. .isolated stand near the mouth of the Ramu
River has apparently not been collected. Here
the Livistonas are growing in a remarkable rain
forest dominated almost to the exclusion of
dicotyledonous trees by palms...”. These
populations are indeed of the new taxa
described in this account, and illustrate both
the cryptic nature of Livistona in New Guinea
and the need for thorough field-work.

Materials and methods

Field-work was undertaken by the authors in
Queensland and Papua New Guinea, between
1996 and 2000. Specimens were examined at
AAU, BO, BRI, LAE, K, QRS and SING, and
specimens lodged at AAU, BRI, LAE and K.

Taxonomy

Livistona concinna Dowe & Barfod sp. nov.,
quod in inflorescentias individuorum non
fructiferorum ad L. drudei F.Muell. ex
Drude accedit sed ab ea differt
essentialiter inflorescentii fructiferis
validioribus et ramosis ad ordinem
altiorem. Arbor usque ad 30 m alta, velut
dioecia fungens. Lamina foliorum
profunde segmentata, segmentis apice
cemuo. Inflorescentiae non ramosae intra
prophyllum. Inflorescentiae partiales 8
vel 9, individuis non fructiferis 120-180
cm longis, ramosis ad ordinem quartum,

individuis fructiferis 160-250 cm longis,
ramosis ad ordinem quintum, bracteae in
rhachidibus et rachillis glabrae. Flores
hermaphroditi solitarii vel 2-4 fasciculati.
Pedicellus fructus ad 2 mm longus,
fructus globosus, 9-12 mm diametro,
epicarpium anthracinum, punctis
lenticellularibus dispersis, mesocarpium
circa 1 mm crassum, oleosum. Typus:
Queensland. Cook District: 16 km north
of Cooktown, just north of the Cooktown
Airport, Barrett Creek, 15°25’S,
145°11’E, 5 m alt., 17 Oct 2000, J. L.
Dowe 607 (holo: BRI; iso: AAU, K,
QRS).

Solitary, functionally dioecious palm.
Trunk to 30 m tall, with diameter at breast
height 24-35 cm, expanded at the base to 100
cm diam., grey; nodes raised; internodes 2-12
cm wide; petiole bases not persistent. Leaves
50-65 in a globose to hemispherical crown;
petioles 120-300 cm long, glabrous, green
throughout, proximally 5-11 cm wide, 2.2-3.5
cm wide in mid area and distally c. 2.8 cm wide,
triangular in cross-section, adaxially
moderately longitudinally ridged, abaxially
rounded, margins with solitary symmetric black
spines 3-5 mm long congested in the proximal
portion with distal margins unarmed, sharp,
slightly winged; leaf-base fibres not prominent,
coarse, persistent; hastula raised, sharp, papery
on the margins; lamina strongly costapalmate,
glabrous, adaxially mid-green, abaxially
slightly lighter green, glossy on both surfaces,
non-waxy, sub-circular in profile, 155-165 cm
long, about 200 cm wide, folded; segments 60-
78, apices deeply forked, distal portion
pendulous; segment free for about 60% of their
length, and with apical split about 41% of the
length of their free portion; mid-lamina
segments 2.6-4 cm wide at the disjunction, with
apices acuminate, filamentous; longitudinal
veins 9 or 10 each side of midrib, parallel, more
prominent than the transverse veins which are
thin and extend across 2-4 parallel veins;
density of transverse veins about 12 per unit
area of 15 x 10 mm. Inflorescence an
unbranched axis with several partial
inflorescences decreasing in size toward the
apex, sexually dimorphic; non fruit bearing
(functionally male) inflorescences 120-180 cm
long with partial inflorescences 8 or 9; most

Dowe & Barfod, New species of Livistona

167

proximal partial inflorescences branched to the
4th order, or to the 3rd order in the distal partial
inflorescence; partial inflorescences slightly
curved; peduncle dorsi-ventrally compressed,
c. 25 mm wide and 10 mm thick, glabrous; fruit
bearing (functionally female) inflorescences
160-250 cm long with partial inflorescences 8
or 9; most proximal partial inflorescences
branched to the 5 th order, to the 3rd order in
the distal partial inflorescence; partial
inflorescences slightly curved, held horizontal
to semi-pendulous; peduncle dorsi-ventrally
compressed, c. 30 mm wide and 10 mm thick;
peduncule lacking empty bracts; prophyll 27-
35 cm long, ancipitous, glabrous; bracts on the
rachis 30-50 cm long, tubular, tightly
sheathing, not disintegrating or splitting with
age, glabrous, apically acute with margins entire
and lateral splits uneven with one about twice
as deep as the other; rachillae 5-20 cm long,
glabrous. Flowers solitary or in clusters of 2—
4, 1.6-2 mm high and about 2 mm wide, white
to cream; sepals basally fused, cupular, 3-lobed
with lobes triangular and margins hyaline, to
about 1.5 mm high and with apices acute; petals
triangular, slightly asymmetric, 2-2.2 mm long,
1.8-2 mm wide at the base, acute at the apex;
stamens about 1 mm high; filament subulate;
connective very thin; anthers dorsifixed, ovoid,
c. 0.2 mm long, didymous; carpels wedged-
shaped, rounded; styles fused, c. 1 mm high;
stigma erect, trilobed; fruiting pedicel to 2 mm
long. Fruit globose, 9-12 mm diam., shiny
black; stigmatic remains subapical; epicarp
smooth but with scattered lenticellular dots,
drying slightly rugose with integumental scar
extending from the stigmatic remains for about
3 A the length of the fruit toward the base;
mesocarp c. 1 mm thick, moist, oily and gritty
in texture; endocarp thin, crustaceous, light
brown, 0.1-0.2 mm thick. Seed globose to
subglobose; seedcoat intrusion extending to
half or less of the width of the seed, contorted,
light brown, crystalline and spongy in texture;
embryo lateral to sublateral, c. 2 mm long.
Eophyll 5-ribbed. Fig. 1.

Specimens examined : Queensland. Cook District:
Lakefield National Park, Kennedy Bend, 14°50’S,
144°15’E, 50m alt., Oct 1997, Dowe 415 with Smith (JCT);
Lakefield National Park, Twelve Mile Waterhole, Oct 1999,
Dowe 606 with Barfod (BRI, JCT, K); Kennedy River, 5.5
km N of New Laura Station, 15°07’S, 144°18’E, 50 malt.,
riparian forest, Nov 1981, Irvine 2204, 2205 (QRS); near
Cooktown airport, Barrett Ck, Gray 2764 (QRS); 3.3 km

NE of Cooktown Airport, Barrett Creek, NW boundary,
15°25’S, 145°12’E, 0 m alt., riparian evergreen mesophyll
vine forest bordering mangroves, Nov 1981, Irvine 2178,
2179,2180 (QRS); near Cooktown, Barrett Creek, 15°33’S,
145° 12’E, 0 m alt., Oct 1999, Dowe 604 with Barfod (BRI,
JCT, K); Endeavour River, opposite the SW comer of the
CooktownAirport, 15°27’S, 145°11’E, 3 malt., Nov 1981,
Irvine 2181 (QRS); 3.3 km off main road at north side of
CooktownAirport, Aug 1986, Hind 4594 with Hill & Healy
(NSW); Cooktown, S side of Endeavour River, near Anzac
Park, 15°28’S, 145°15’E, 10 m alt., Feb 1996, Dowe 252
(BRI, FTG, JCT). Cultivation. Queensland. Townsville,
Anderson Park Botanic Gardens, Oct 1998, Dowe s.n.
(JCT).

Distribution and habitat : Australia.
Queensland: Flinders Island, drainage area of
the Kennedy River and tributaries, Barrett
Creek and along the Endeavour River north of
Cooktown and Archer Point south of
Cooktown; in seasonally moist open forest,
seasonally inundated Melaleuca swamp, along
creek and river banks, and at mangrove margins
in non-saline environments; soils usually
alluvial. Occurs with Corypha utan Lam. in the
Kennedy River area and with L. muelleri F. M.
Bailey in the vicinity of Cooktown. Most
populations are regularly affected by fire.

Phenology: Flowering December to March;
fruiting April to October.

Etymology : From Latin concinnus, neat, well-
arranged, in reference to the organised and neat
appearance of the petioles in the crown
compared to those in other Livistona species.

Conservation: Adequately conserved in
Lakefield National Park, Endeavour River
National Park and on Flinders Island.

Notes: Specimens of this taxon were tentatively
placed under L. drudei by Rodd (1988). Rodd
cited three specimens {Irvine 2204 & 2205, and
Hind 4594) that were collected in the
distributional range of L. concinna. One
consists of a fallen dead leaf and an old
infructescence. Although there is some gross
resemblance of L. concinna to L. drudei ,
differences are otherwise significant,
particularly in morphology of the inflorescence
and fruit. The inflorescence of non fruit-bearing
individuals is similar to that in L. drudei, but
the fruit-bearing inflorescences are more robust
and branched to a further order in L. concinna
than they are in L. drudei. Although L. concinna
occurs with L. muelleri in some locations, we

168

Austrobaileya 6 (1): 165-174 (2001)

Fig. 1. Livistona concinna. A. Habit. B. Leaf-bases and leaf-base fibres of crown x 0.05. C. Leaf in adaxial view x 0.06.
D. Flower viewed from above. E. Flower in lateral view. F. Flower in LS view showing orientation of the ovule. D-F x
17.5. G. Fruit in lateral view. H. Fruit in LS view showing mesocarp [1], seedcoat intrusion [2], endosperm [3] and embryo
[4], G-H x 2.7. A-C from Dowe 607 (K); D-F from Dowe s.n., 17 Oct. 1998 (JCT); G-H from Dowe 252 (JCT). Del. Lucy
T. Smith.

Dowe & Barfod, New species of Livistona

169

have not detected, at least by visual means,
any putative hybrids between the two.

Livistona surru Dowe & Barfod sp. nov. Species
haec ab L. tothur Dowe & Barfod differt
colori laminae foliorium quasi aequali
utrinque, apice segmentorum foliorum
cernuo, bracteis in pedunculis absentibus,
bracteis in rhachidibus fibrosis et dense
pubescentibus, paginis axium
inflorescentiarum dense

pubescentibus, fructibus rubellis
aurantiacis ad 65 mm diametro, granis
sclerotis longistrorsum parallelis in
epicarpio inclusis. Arbor usque ad 20 m
alta, velut dioecia fungens. Segmenta
laminae foliorum apice cernuo.
Inflorescentiae prophyllo ramosae axibus

3 collateralibus subaequalibus,
unumquemque inflorescentiis partialibus
5-7, individuis fructiferis ad ordinem
secundum divisis, bracteae in rhachidibus
et rhachillis dense pubescentes. Flores 2-

4 fasciculati. Pedicellus fructus 6-12 mm
longus, fructus globosus versus
obovoideus, 55-65 mm diametro,
epicarpium rubellum aurantiacum punctis
lenticellularibus dispersis, mesocarpium
latum et carnosum fibris crassis. Typus:
Papua New Guinea. West Sepik
Province: Miwaute, 03° 25’S, 142° 07’E,
950-1000 m alt., 20 Nov 1996, A. S.
Barfod 390 with M. D. Ferrero & A.
Damborg (holo: AAU; iso: K, LAE).

Solitary, functionally dioecious palm. Trunk to
20 m tall, 18-25 cm diameter at breast height,
erect, slightly broader at the base, light grey,
usually covered by crustaceous lichens; nodes
slightly raised, internodes to 5 cm wide, lacking
longitudinal fissures, petiole bases not
persistent. Leaves 17-29 in a spherical to
vertically oblong crown; petiole 140-180 cm
long, slightly arching, green, proximally about
19 cm wide and triangular in cross section,
adaxially flat, abaxially rounded, glabrous
except for scattered lepidote scales that are
brown in the centre and grey at the margin, more
densely so on the abaxial surface with margins
with single or grouped black spines 5-10 mm
long, largest and more closely inserted in the
proximal portion, becoming smaller and wider
spaced in the distal portion; leaf-base fibres in

2 layers, the outer with thick fibres, the inner
with thin coir-mat like fibres, reddish brown,
persistent until leaf fall then deciduous in
sheets; ligule to 1 m long, to 10 mm thick;
hastula very prominent, raised c. 2 cm, 5 cm
wide, thick edged; lamina sub-circular to ovate,
180-224 cm long, 143-160 cm wide, undulate,
adaxially mid green, abaxially similar green,
pendulous in the distal portions; segments 70-
90, free for 45-80% of their length and apically
split about 6% of length of their free portion;
mid-lamina segments 4.5-7 cm wide at the
disjunction; parallel veins 5 or 6 on each side
of midrib, more prominent than transverse veins
which are thin and extend across 2-6 parallel
veins with a density of about 22 per unit area
of 15 x 10 mm. Inflorescences basally branched
within the prophyll with 3 subequal axes about
120 cm long; each axis bears 5-7 partial
inflorescences which are branched to the 3rd
order on non fruit-bearing plants, to the 2nd
order on fruit-bearing plants but otherwise
similar; prophyll to 37-42 cm long, 12.5-15
cm wide, glabrous, lacerate-fibrous at the apex;
peduncle of individual axes subterete, to 3 cm
wide, lacking bracts; bracts on rachises 40-45
cm long, loosely tubular, fibrous, disintegrating
at the apex with maturity, pubescent throughout
but more densely so toward the apex; rachillae
subterete to angular, 14-24 cm long, densely
covered with long coarse red appressed scales
in the proximal portion, with long white scales
distally; scales less dense to absent in the
extreme distal portions. Flowers in clusters of
2-4; fruiting pedicel 6-12 mm long, c. 3 mm
across, green, with prominent scars from fallen
flowers. Fruit globose to obovoid, 55-65 mm
long, 50-55 mm diam., orange-red, shiny;
epicarp with scattered lenticellular dots and c.

3 mm long lines pointed toward the fruit apex;
stigmatic remains present apically; longitudinal
stripe of suberised epidermal tissue usually
visible for full length of fruit; mesocarp fleshy
with fibres thick, distributed throughout but
more densely aggregated toward the endocarp
and shallowly embedded in the surface of the
endocarp; endocarp to 2 mm thick, bony. Seed
globose to subglobose, 30-40 mm diam.;
endosperm intruded by the seedcoat to about
two-thirds across; intrusion broadly kidney¬
shaped, crystalline/spongy, orange; embryo
lateral. Fig. 2.

170

Austrobaileya 6 (1): 165-174 (2001)

Fig. 2. Livistona surru. A. Habit. B. Leaf in adaxial view x 0.06. C. Infructescence x 0.09. D. Basal portion of partial
infructescence showing rachis bract apex and immature fruit x 0.4. E. Fruit in lateral view. F. Fruit with epicarp removed
exposing mesocarp fibres. G. Fruit in LS view showing mesocarp [1], seedcoat intrusion [2], endosperm [3] and embryo
[4], E-G x 0.4. A, C, D & E from Batfod 390 (K); B, F & G from W.J. Baker 582 (K). Del. Fucy T. Smith.

Dowe & Barfod, New species of Livistona

171

Specimens examined : Papua New Guinea. West Sepik
Province: Miwaute, 142° 07’E, 03° 25’S, 950-1000 m alt.,
Nov 1996, Barfod 399 with Ferrero & Damborg (AAU, K,
LAE); Upper Freida River, Mt Ekwai, 1200-1300 m alt.,
Feb 1998, Ferrero 980029 (LAE). MADANG PROVINCE:
Bosmun village at mouth of Ramu River, 04°07’S,
144°43’E, 10-20 m alt., Nov 1996, Damborg 354 with
Ferrero & Barfod (AAU, LAE, K); Goinbang, near Bosmun
2, mouth of Ramu River between Bogia and Bosmun 2,
Jan 1996, Baker WJB582 (K).

Distribution and habitat: Papua New Guinea.
West Sepik Province, Miwaute area and Mt
Ekwai; Madang Province, in the vicinity of the
mouth of Ramu River.

Phenology: Flowering November to January;
fruiting March to June.

Etymology: From the native Olo language
vernacular name surru, used in the Miwaute
area for this plant.

Ethnobotany: Feaves are used for roof
thatching and umbrellas, stem portions for axe
handles and house frames, and leaf sheath fibres
for brooms and sago strainers.

Notes: Livistona surru is distinguished by its
leaf lamina with almost equal colour on both
surfaces; leaf segment apices pendulous, the
absence of bracts on the inflorescence peduncle,
the presence of densely pubescent, fibrous
bracts subtending the basal branches of the
partial inflorescences, the surfaces of the
inflorescence axes being densely pubescent,
and by its large orange-red fruit to 65 mm
diameter with thick mesocarp fibres and
prominent longitudinally parallel sclerids
embedded in the epicarp.

Livistona tothur Dowe & Barfod sp. nov.
differt a L. surru Dowe & Barfod colore
argenteo glauco in latere abaxiali laminae
foliorum, apice segmentorum foliorum
rigido, praesentia bracteae ad basim
uniuscujusque axium inflorescentiae,
bracteis in rhachidibus chartaceis et
glabris, paginus axium inflorescentiarum
glabris vel moderate pubescentibus,
fructibus rubellis aurantiacis ad 43 mm
diametro. Arbor usque ad 20 m alta, velut
dioecia fungens. Pagina adaxialis laminae
foliorum aeruginosa, pagina abaxialis
argentea glauca, segmenta laminae
foliorum rigida ad apicem, inflorescentiae

prophyllo ramosae axibus 3 collateralibus
subaequalibus, unumquemque
inflorescentiis partialibus 5 vel 6, in
individuis fructiferis ramosis ad ordinem
tertiam, bracteae in rhachidibus et
rhachillis fere glabrae. Flores solitarii
rubrae. Pedicellus fructus 2.5-5 mm
longus, fructus globosus, 35-43 mm
diametro, epicarpium rubellum
aurantiacum, punctis lenticellularibus
dispersis, mesocarpium latum, fibris
tenuibus. Typus: Papua New Guinea.
West Sepik Province: Onake Mts, on road
to Niau Kono from Vanimo, 2°45.89’S,
141°04.06’E, 500 m alt., 26 Nov 1996,
A. Damborg 418 with A. S. Barfod (holo:
AAU; iso: K, FAE).

Solitary, functionally dioecious palm. Trunk to
20 m tall, 15-20 cm diameter at breast height,
erect, slightly swollen at the base, grey, nodes
slightly raised, c. 1 cm wide, dark grey,
internodes to 15 cm long; petiole bases not
persistent. Feaves 24-40 in an open spherical
crown; petiole 150-200 cm long, proximally
about 10 cm wide, distally about 15 mm wide,
adaxially flat, abaxially rounded, arching,
green, glabrous, margins with single, recurved,
green spines 1-2 mm long throughout its length,
but largest and more closely spaced in the
proximal portion; margins lacerate fibrous in
extreme proximal portion; leaf-base fibres
prominent, in 2 layers of more or less equal
coarseness, persistent in sheets, chestnut brown;
ligule to 60 cm long; hastula raised to c. 10
mm, semi-circular; lamina sub-circular, 150-
200 cm long, 120-150 cm wide, adaxially
bluish-green, abaxially silvery glaucous;
segments 60-75, rigid with apex becoming
pendulous only with age or damage, free for
62-85% of their length and apically split for
1-3% of the length of their free portion, 3-4
cm wide at the disjunction; segments not
overlapping in the hastula region; parallel veins
6 or 7 on each side of midrib, more prominent
than transverse veins which are very thin and
extend across 2-7 parallel veins with a density
of c. 33 per unit area of 15 x 10 mm.
Inflorescences basally branched with 3 similar
collateral axes, each about 200 cm long with 5
or 6 partial inflorescences which in fruit¬
bearing plants are branched to the 3rd order;
prophyll c. 30 cm long and 10.5 cm wide,

172

Austrobaileya 6 (1): 165-174 (2001)

Fig. 3. Livistona tothur. A. Habit. B. Leaf in adaxial view x 0.05. C. Mid-section of petiole x 0.2. D. Inflorescence x 0.08.
E. Flower in lateral view. F. Flower in LS view showing orientation of the ovule. E-F x 20. G. Flowers attached to rachilla
x 7. H. Fruit in lateral view. I. Fruit in LS view showing mesocarp [1], seedcoat intrusion [2], and endosperm [3]. H-I x
0.5. A, D, E, F & G from Barfod 510 (K); B from Dowe 516 (JCT); C, H & I from Damborg 418 (K). Del. Lucy T. Smith.

Dowe & Barfod, New species of Livistona

173

yellow and coriaceous proximally, brown and
chartaceous distally, glabrous apart from a
ferrugineous woolly tomentum along the
carinae; peduncle with one tubular, papery and
loosely sheathing bract, densely scaly at the
apex, glabrous; peduncle of individual axes
proximally subterete, becoming dorsi-ventrally
compressed distally with edges angular,
furfuraceous in parts enclosed in bracts,
otherwise patchily furfuraceous or glabrous on
exposed surfaces; bracts on rhachises tubular,
papery, loosely sheathing, glabrous apart from
some irregular pubescence toward the apex;
apices more or less non-fibrous showing only
minor disintegration; rachillae rigid, terete, 6-
12 mm long, to 3 mm diam., glabrous, red,
irregularly disposed. Flowers solitary; sepals
fused, red, tri-lobed with lobes to 1.2 mm long;
petals broadly triangular, about 2 mm long,
basally connate for about half their length,
thick, fleshy, red with apical margins recurved,
the inner surface bearing the impression of the
stamens, the outer surface minutely warty;
stamens much shorter than petals, basally fused
to petals; filaments very short; anthers c. 0.2
mm long, cream; fruiting pedicel 2.5-5 mm
long, c. 2 mm wide. Fruit globose but with a
basal constriction, 35-43 mm diam., orange-
red with stigmatic remains present apically;
epicarp with scattered lenticellular dots, glossy
with a longitudinal stripe of suberised
epidermal tissue usually visible for the full
length of the fruit; mesocarp 13-15 mm thick,
softly fibrous, mealy, orange; endocarp to 2 mm
thick, bony. Seed globose, 22-28 mm diam.;
endosperm deeply intruded by the seedcoat;
intrusion kidney shaped with an orange pulpy
tissue; embryo lateral. Fig. 3.

Specimens examined : Papua New Guinea. West Sepik
Province: Onake Mts, Apol area, on road to Niau Kono
from Vanimo, 02°45.89’S, 141°04.06’E, 500 m alt., Feb
1998, Dowe 516 with Ferrero (JCT, LAE); Onake Mts,
Niau, 2°46.276’S, 141°03.61 l’E, 425 m alt., Mar 2000,
Barfod 510 with Banka & Kjaer (AAU, BRI, JCT, K, LAE).

Distribution and habitat: Papua New Guinea:
West Sepik Province, Onake Mts, 400-600 m
alt., in rainforest on ridges of limestone and
metamorphic rocks.

Phenology: Flowering November to January;
fruiting March to June.

Etymology: From the native Bewani language
vernacular tothur, pronounced ‘tot-her’, and
used in the Niau area for this plant.

Ethnobotany: Bows and roof struts are
fashioned from the petioles, umbrellas are made
from the leaves and salt is extracted from the
ash of burned petioles.

Notes: Livistona tothur is distinguished by the
silver glaucous bloom on the abaxial surface
of its leaf lamina, the leaf segments with rigid
apices, the presence of a bract at the base of
each axis of the inflorescence and papery and
glabrous bracts on the inflorescence rachis,
glabrous or moderately pubescent inflorescence
axes, and orange-red fruit to 43 mm diameter.

Acknowledgments

We thank the PNG Forest Research Institute
for permission to use the drying facilities in the
Lae Herbarium. We appreciate our interaction
with Roy Banka, curator of the living
collections in the Lae Botanic Garden, who
joined us in field studies in 1999 and 2000.
Field-work was funded by the Danish Natural
Science Research Council (Grant 9600861) and
the Carlsburg Foundation (Grant 980298/10 -
1150). This paper was written during Anders
S. Barfod’s sabbatical leave in 1999-2000 while
based at the Queensland Herbarium. We are
grateful to Gordon Guymer, Director of the
Queensland Herbarium, for making this
possible. A special thanks goes to Lucy T. Smith
who skilfully rendered the line drawings.

References

Beccari, O. (1877). Le specie di palme raccolte alia
Nuova Guinea da O. Beccari e dal medesimo
adesso descriite, con note sulle specie dei paesi
circonnvici. Malesia 1: 9-102.

-(1921). Recensione delle Palme del vecchio

mondo. Webbia 5: 11-22.

Burret, M. (1935). Neue Palmen aus Neuguinea II.
Notizbl. Bot. Gart. Berlin-Dahlem 12: 309-
348.

-(1939). Palmae gesammelt in Neu Guinea von

L. J. Brass. J. Arnold Arb. 20: 187-212.

-(1941). Bemerkungen zur Palmengattung

Livistona R. Br. Notizbl. Bot. Gart. Berlin-
Dahlem 15: 319-327.

174

Austrobaileya 6 (1): 165-174 (2001)

Dowe, J. L. and Jones, D. L. (in press). Arecaceae. Flora
of Australia 39.

Essig, F. B.(1977). The palm flora of New Guinea: a
preliminary analysis. Botany Bulletin No. 9. Lae:
Office of Forests.

Essig, F. B. and Young, B. E. (1981). Palm collecting in
Papua New Guinea. II. The Sepik and the north
coast. Principes 25: 3-15.

Hay, A. J. M. (1984). Palmae. In: R. J. Johns and A. J.
M. Hay (eds), A guide to the monocotyledons
of Papua New Guinea, 195-318. PNG: Office
of Forests.

Irvine, A. K. (1984). A guide to Livistona in Queensland.
Palms & Cycads 5: 2-5.

Jones, D. (1984). Palms in Australia. Frenchs Forest:
Reed Books.

Rodd, A. N. (1998). Revision of Livistona (Arecaceae)
in Australia. Telopea 8: 49-153.

Tucker, R. (1988). Palms of subequatorial Queensland.
Milton: Palm & Cycad Societies of Australia.

Austrobaileya 6 (1): 175-176 (2001)

Note

Kentrophora S.M.Wilson & Kraft, a new name for an algal genus in tribe Amansieae

(Rhodomelaceae, Rhodophyceae)

In an account of selected genera of the algal
tribe Amansieae (Wilson & Kraft 2000), the
name Plectrophora S.M.Wilson & Kraft was
published for a genus erected to accommodate
two distinctive species, one occurring in
Western Australia, eastern Queensland, Lord
Howe Island and Norfolk Island, and the other
in Natal, South Africa. Since this generic name
was published, however, it has come to our
attention that H.C. Focke had previously
published the name Plectrophora for a genus
in the phanerogamic family Orchidaceae
(Focke 1848), which therefore renders the more
recent Plectrophora illegitimate and hence
unusable (Greuter etal. 2000, Article 53.1). To
provide a legitimate name for the new algal
genus recognised by Wilson and Kraft in 2000
and acceptable names for the species belonging
in it, the following are proposed.

Kentrophora S.M.Wilson & Kraft, nom. nov.,

Plectrophora S.M.Wilson & Kraft, Aust.
Syst. Bot. 13: 353 (2000), nom. illeg. non
Plectrophora H.Focke (1848, p.212).
Typus: Kentrophora pectinella
(Harvey) S.M.Wilson & Kraft;
Dictyomenia pectinella Harvey

Etymology: Kentrophora from Greek Kevxpov
(.Kentron ), a cock’s spur/porcupine’s quill, and
the feminine adjectival suffix -phora, -carrying,
in reference to the acute spurs borne dorsally
on each of the marginal teeth on the fronds.

1. Kentrophora pectinella (Harvey)
S.M.Wilson & Kraft, comb. nov.

Dictyomenia pectinella Harvey, Trans. Roy.
Irish Acad. 22: 538 (1855); Kuetzingia
pectinella (Harvey) Falkenb., Fauna
und Flora des Golfes von Neapel,
Monographie 26. XYI (Berlin): 454-455
(1901); Plectrophora pectinella

(Harvey) S.M.Wilson & Kraft, Aust. Syst.
Bot. 13: 353 (2000). Type: Australia.
Western Australia: Garden Island,
presumably from drift, Harvey 290 (holo:
TCD; icon: Wilson & Kraft 2000, p.354,
fig. 17 A).

Enantiocladia robinsonii (J.Agardh)
Falkenb., Fauna und Flora des Golfes
von Neapel, Monographie 26. XVI
(Berlin): 441 (1901); Amansia robinsonii
J.Agardh, Lunds Universitets Arsskrift.
Afd. 2,3(6): 174 (1892). Type: Norfolk
Island. [?] Kingston, Isaac Robinson 13
(holo: LD [LD42655-42659a]; icon:
Wilson & Kraft 2000, p.354, fig.l7B
[LD42655]).

2. Kentrophora natalensis (J.Agardh)
S.M.Wilson & Kraft, comb. nov.

Kuetzingia natalensis J.Agardh, Species
Genera et Ordines Algarum 2(3): 1099
(1863); Plectrophora natalensis
(J.Agardh) S.M.Wilson & Kraft, Aust.
Syst. Bot. 13: 358 (2000). Type: South
Africa. Port Natal, Krauss (holo: LD
[LD42572], n.v., fide P. Lassen pers.
comm.; iso: MEL [MEL603052]).

Acknowledgements

We thank Dr Per Lassen, Curator, Botanical
Museum, Lund, for information on the type
specimens of Amansia robinsonii J.Agardh and
Kuetzingia natalensis J.Agardh, and Dr Alan
Millar, Senior Research Scientist, National
Herbarium of New South Wales, Sydney, for
kindly communicating with Dr Lassen on our
behalf. Gea Zijlstra, Nationaal Herbarium
Nederlands, Utrecht University, Utrecht,
Netherlands is also thanked for providing
information regarding Focke’s publication.

Accepted for publication 20 July 2001

176

Austrobaileya 6 (1): 175-176 (2001)

References

Focke, H.C. (1848). Uittreksels uit botanische berigten over
de flora van Suriname [Ornithocephalus falcatus,
Plectrophora iridifolia ]. Tijdschr. Wis-Natuurk.
Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch.
1: 209-212.

Greuter, W., McNeill, J., Barrie, F.R., Burdet, H.M.,
Demoulin, V., Filgueiras, T.S., Nicolson, D.H.,
Silva, P.C., Skog, J.E., Trehane, P., Turland, N.J.
& Hawksworth, D.L. (2000). International Code
of Botanical Nomenclature (Saint Louis Code).
Regnum Vegetabile 138. Konigstein, Germany:
Koeltz Scientific Books.

Wilson, S.M. & Kraft, G.T. (2000). Morphological and
Taxonomic Studies of Selected Genera from the
Tribe Amansieae (Rhodomelaceae, Rhodophyta).
Aust. Syst. Bot. 13: 325-372.

R. J.F. Henderson

Queensland Herbarium, Environmental Protection Agency, Brisbane Botanic Gardens Mt
Coot-tha, Mt Coot-tha Road, Toowong, Queensland 4066

S. M. Wilson and G.T. Kraft

School of Botany, University of Melbourne, Parkville , Victoria 3052