Nuytsia : bulletin of the Western Australian Herbarium

VOLUME 8
NUMBER 3

^ DEPARTMENT OF
CONSERVATION AND
LAND MANAGEMENT

UME 8 NUMBER3 199

Western Australian Herbarium
Department of Conservation and Land Management
Como, Western Australia

CONTENTS

Page

Calolhamnus superbus T.J. Hawkeswood & F.H. Mollemans (Leptospermoideae:

Myrtaceae), a new species from south-west Western Australia. By T.J. Hawkeswood

and F.H. Mollemans 311

A new species of Guichenotia (Sterculiaceae) from south western Australia.

By Gregory J. Keighery 319

Four new Drosera taxa from south western Australia. By Allen Lowrie and

Neville Marchant 323

New names and combinations for some Melaleuca (Myrtaceae) species and
subspecies from the south-west of Western Australia considered rare or threatened.

By F.C. Quinn, K.J. Cowley, B.A. Barlow and K.R. Thiele 333

Three new species of Euphorbia L. subgenus Chamaesyce Rafinesque

(Euphorbiaceae) from central and northern Australia. By B.G. Thomson 351

Hht Lawrencella complex (Asteraccac: Gnaphalieae: Angianthinae) of Australia.

By Paul G. Wilson 361

The classification of Australian species currently included in Heliplerum and related

genera (Asteraceae; Gnaphalieae): Part 1. By Paul G. Wilson 379

The classification of Australian .species currently included in Heliplerum (Asteraceae:
Gnaphalieae): Part 2 Leucochrysum. By Paul G. Wilson 439

The classification of some Australian species currently Included in Heliplerum
and Helichrysum (Asteraceae: Gnaphalieae): Part 3 Anemocarpa and

Argenlipallium, two new genera from Australia. By Paul G. Wilson 447

The classification of the genus Wailzia Wendl. (Asteraceae: Gnaphalieae).

By Paul G. Wilson 461

A new species of Acomis from the Northern Territory and a new combination in

the genus Thisellonia (Asteraceae: Gnaphalieae). By Paul G. Wilson 479

Correction: Nuytsia 8(2):228 484

Publication date of Nuytsia Volume 8 Number 2 484

Editor

Kevin F. Kenneally
Editorial Advisory Committee
N.S. Lander
T.D. Macfarlane
N.G. Marchant
Paul G. Wilson

Editorial Assistant
J.W. Searle

Western Australian Herbarium,
Department of Conservation and Land Management,
PO Box 104, Como, Western Australia 6152

Cover

Nuytsia floribunda (Labill.) R. Br. ex Fenzl (Loranthaceae) — the Western Australian Christmas Tree is one of the few
aborescent mistletoes in the world. This endemic tree is a semi-parasite common in sandy soil from the Murchison River to
Israelite Bay. The journal is named after the plant, which in turn commemorates Pieter Nuijts, an ambassador of the Dutch East
India Company, who in 1627 accompanied the "Guide Zeepard” on one of the first explorations along the south coast of
Australia.

Cover design by Sue Marais

Photograph A.S. George

Nuytsia 8(3): 31 1-318 (1992)

311

Calothamnus superbus T.J. Hawkeswood & F.H. Mollemans
(Leptospermoideae: Myrtaceae), a new species from south-west

Western Australia

TJ. Hawkeswood' & F.H. Mollemans^

'49 VennerRoad, Annerley, Brisbane, Queensland 4103
^76-6228 Plumeria Road, Kailua-Kona, Hawaii 96740 USA
[Current address: PO Box 734, Victoria Park, Western Australia 6100]

Abstract

Hawkeswood, T. J. &Mollcmans, F,H,Ca/o//!t7WrtHd'5M;jcrftM^T.J. Hawkeswood &F,H. Mollemans
(Leptospermoideae: Myrtaceae), a new species from south-west Western Australia. Nuytsia 8(3):
311-318 (1992). Anew 5-merous species of Calothamnus, C. superbus Hawkeswood & Mollemans,
is described from sandplains in the Pigeon Rocks area, south-west Western Australia. Its affinities
with the closely related species, C. aridus T.J. Hawkeswood, are outlined, and ecological data
provided. The plant is not represented in any National Parks or Nature Re.serves but its survival is
probably ensured due to its isolated location.

Introduction

During 1990, an apparently undescribcd 5-merous taxon of Calothamnus (Leptospermoideae:
Myrtaceae), was located by the second author in aremote area near Pigeon Rocks, south-west Western
Australia. Further research and collections have indicated that this taxon is indeed distinct and
previously undescribed. The new species is described below. The terminology used in the description
and general format follows that of Hawkeswood (1984a,b, 1987).

Taxonomy

Calothamnus superbus T.J. Hawkeswood & F.H. Mollemans, sp. nov. (Figures 1-4)

Frutex erectus ad 2.5 m altus. Folia linearia, terctia, erecta, (10)13-15(18) cm longa, L2-L6 mm
lata, mucronata, glabra. Flores fasciculati vel breviter spicati. Calycis tubus pleraque4-5 mm longis,
glabris; calycis-lobi 1-1.5 mm longi. Petala 3-5 mm longa, ferruginea. Unguis staminalis 20-25 mm
longus, 1. 8-2.1 mm latus,glaber; filamentis marginalibus 12-17. Frutus globulus vel ± cylindraceus,
5-7 mm longus, 7-9 mm latus, glaber. Semina pleraque 1-1.4 mm longa, ferruginea.

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Nuytsia Vol. 8, No. 3 (1992)

Figure 1. T.J, Hawkeswood & F.H. Molleman.s - Holotype (F.M. & M.P. Mollemans 3078).

T.J. Hawkeswood & F.H. Mollemans, Calothamnus superbus

313

Figure 2. Habit of Calothamnus supcrbus T.J. Hawkeswood & F.H. Mollemans
in mixed shrubland and heath near Pigeon Rocks. (Photo: F.H. Mollemans).

Typus: Track from Emu Fence, east towards Pigeon Rocks Tank & west of Clampton Mineral Prospect
(prcciselocalitywithheld),alt.c.510m;abund;uice uncommon; 11 July 1990, F./f. &.M.P. Mollemans
3078 (holo: PERTH). (Figure 1)

Erect, wide, compact to spreading, multi-stemmed, diffuse branching, straggly to somewhat
compact, glabrous shrubs to 2.5 metres high, with thick stems and hard grey bark, often splitting at
or near the bases of the trunks. Y oung branches mostly glabrous. Leaves sessile, linear, terete, erect,
rigid, ( 1 0) 13- 1 5( 1 8) cm long, 1 .2- 1 .6 m m wide, glabrous, rough , pungent, narrowed into a thin, sharp,
straight apex 4-7 mm long, bright pale green to yellowish-green; oil glands very prominent and
exserted into the epidermis, giving rise to the rough leaf surface. Flowers 2-5 (mostly 3) arranged
in a cluster or 5-8(10) in a loose spike amongst leaves on younger branches. Calyx-tube narrow
campanulate to almost cylindrical, 4-5 mm long, glabrous; oil glands prominent; rhachis usually
dilated at the base of the calyx-tube; calyx-lobes deltoid, mostly acute, concave, 1-1.5 mm long,
glabrous; margins thin, scarious, partially ciliate. Pcm/j'obovatc, concave, obtuse, 3-5 mm long, thin,
mostly glabrous, but with a few simple, scattered hairs, orange-brown; margins thin, glabrous.
Staminal claws ± equal, free, 20-25 mm long, 1.8-2. 1 mm wide, glabrous, orange-red in lower half
to two-thirds, pink-red in upper portion; marginal filaments 12-17; anthers linear, 1-1.4 mm long,
brown. Style slender 15-25 mm long, glabrous, pink-red, stigma small, often persistent in young fruit.
Summit of ovary densely pubescent. Fruit globular or depressed globular to almost cylindrical
(rarely), 5-7 mm long, 7-9 mm wide, smooth or with irregular bulges, ribs and/or striations, mostly
truncate or very shortly 5-lobed (calyx-lobes persistent in young fruit and usually ab.senl in older, worn
fruit). Fertile iecfJs linear-oblong, 1.0- 1.4(1. 5) mm long, angular, cuneate, often obliquely truncate,
glabrous; testa dark reddish-brown . Ovulodes similar to fertile seeds, mostly 1 .0- 1 .2 mm long, slightly
paler in colour, usually obliquely truncate, glabrous.

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Nuytsia Vol. 8, No. 3 (1992)

Figure 3. Close-up of a branch of Calolhamrms superbus T.J. Hawkeswood
& F.H. Mollemans in mixed shrubland and heath near Pigeon Rocks showing a cluster of
young fruiting capsules. (Photo: F.H. Mollemans).

Other- specimens examined. WESTERN AUSTRALIA: Pigeon Rocks (Clamptons Vermin Fence
track, east of vermin fence), 12 July \99\,F.H. Mollemans 3S12 (PERTH); Pigeon Rocks (Clamptons
Vermin Fence track, east of vermin fence), 13 July 1991, F.H. Mollemans 3813 (PERTH).

Distribution. Known only from two sites in the Pigeon Rocks area (29‘^52'S, 1 18°39’E), south-west
Western Australia (Figure 4). C. superhus occurs c. 100 km south of the most southerly known
occurrence of C. aridus, and the northernmost distribution of C. gilesii is sympatric with that of
C. superhus. The sympatric distribution of C. superbus and C. o/Zej/i caused initial confusion during
the return collection trip in July 1991 (because of morphological resemblance), however, it was
determined during a 3 km transect on foot along Clamptons Vermin Fence track that C. gilesii and
C. superhus are not sympatric at the local scale as they do not grow together.

Habitat. Grows in pale yellow brown sand with scattered ferruginous pebbles of laterite in mixed
sandplain heath wilh Acacia (Mimosaceae), Grevillea (Proteaceae) and/4//ocfliMa/'i/ifl (Casuarinaceae)
or in pale yellow brown sand in mixed shrubland and heath with Grevillea (Proteaceae), Melaleuca
(Myrtaceae) and Boronia ternala (Rutaceae).

Floweringand fruiting period. Flowering in July 1990 when the initial collection was made, but plants
were in fruit in July 1991; this taxon is therefore thought to flower opportunisticly depending on
rainfall.

Conservation status. 2V using the criteria of Briggs & Leigh (1988). Calothamnus superbus is not
represented in any National Park or Nature Reserves but its isolated location should afford a
significant degree of protection from land clearing, residential development and illicit flower
collection at least in the near future. However, at one of the sites where C. superbus was found
{Mollemans 3812), there was evidence of a recent fire so that bushfires may prove to be a possible

T.J. Hawkeswood & F.H. Mollemans, Calothamnus superbus

315

Figure 4. Distribution of C. superbus T.J. Hawkeswood & F.H.Mollenians.

and serious threat especially considering the small size of known C. superbus populations, viz. only
about 25 plants were counted by Mollemans at the first site (Mo/Zmanx 3812)andabout ISattheother
site {Mollemans 3813). The degree of susceptihility of C. superbus to fires is not known and perhaps
fires and their effects should be closely monitored in the future.

Etymology. The specific epithet superbus, from Latin, meaning impressive or exalted, refers in part
to the distinctive pale green to yellow-green, erect foliage of this species and to the plants prominence
in the localized area within which it occurs.

Discussion

There are about twenty 5-merous species of Calothamnus, and C. superbus is most closely related
to C. aridus T.J. Hawkeswood (Hawkeswood 1984a). A comparison of some taxonomic and
ecological features of these two species is provided in Table 1.

Morphological dijferences. C. superbus and C. aridus can be clearly distinguished on the basis of
the four leaf characters given in Table 1. Differences in the morphology and size of the calyx-tube,
petal pubescence, width of the staminal claws and the number of filaments per staminal claw also
distinguish C. superbus from C. aridus. Of these characters, leaf size (length and width), pubescence
of calyx-tube and the number of marginal filaments per staminal claw are the most important. Fertile
seed size differences are also highly significant (Table 1).

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Nuytsia Vol. 8, No. 3 (1992)

Table 1. Comparison of some important taxonomic and ecological features of Calothamnus aridus
T.J. Hawkeswood and C. superbus TJ. Hawkeswood & F.H. Mollemans

Character

C. aridus

C. superbus

Leaf length (mm)

(5)7-10(12)

(10)13-15(18)

Leaf width (mm)

0.6-L2

1.2-1.6

Leaf colour

Dark green to
olive green

Bright pale green
to yellow-green

Leaf apex length (mm)

2-3

4-7

Calyx-tube length (mm)

(2)2.5

4-5

Calyx-tube

Pubescent

Glabrous

Petals

Densely pubescent

Few scattered hairs

Petal margins

Ciliate

Glabrous

Staminal claw width (mm)

1.0-1.5

L8-2.1

Number of marginal
filaments per staminal claw

10-12

12-17

Fertile seed length (mm)

L5-2.0

L0-L4(1.5)

Habitat

Red sand, Triodia-
Eucalyptus woodland;
Triodia sandplain

Yellow sand, mixed
heath, sandplain

Fruit size has been important in delineating and differentiating other Calothamnus taxa (Hawkeswood
1984a,b, 1987), but this character is not so readily applicable in distinguishing C. rapcrfmr and
C. aridus. The two known populations of C. superbus have variable fruit size with many fruits larger
than those of C. aridus, but there is an overlap in size range of fruits of the two species, so this character,
if used in isolation, is not reliable in differentiating the two taxa.

Non-flowering C. superbus plants resemble those of C. gilesii¥. Muell but the fruit of the latter
species are not depressed-globular in shape. In addition, the fruits of C. gilesii are much larger than
those of C. superbus, and C. gilesii is not so tall or broad in habit, nor does it have the superb ascending
leaves of C. superbus.

Ecological dijferences. Calothamnus superbus and C. aridus have the most inland distribution of any
species in the genus. However, the habitats in which they occur are quite different (Table 1) and the
soil type and other environmental factors may have been important in the evolution and genetic
integrity of these two species.

T.J. Hawkeswood & F.H. Mollemaiis, Calothamnus superbus

317

Possible evolutionary relationships. C. superbus, like all species of Calothamnus, other than
C. aridus, lacks pubescent petals. This is an important difference between C. superbus and C. aridus,
and maintains C. aridus as the only species of Calothamnus which possesses petals covered in short,
appressed, simple hairs (see Hawkeswood 1984a). Despilethisdifferencc.itis very clear, on the basis
of other characters (see Table T) that C. superbus is very closely related to C. aridus.

Hawkeswood (1984a), noted that C. aridus was not closely related to any other Calothamnus
species. But the new species C. superbus, because of its lack of petal pubescence and its very close
relationship to C. aridus, appears to provide a link between C. aridus and other Calothamnus species
which has not previously been evident. However, absence of petal indumentum is only one common
character among a number of characters in which C. superbus (in common with C. aridus) differs from
other species of Calothamnus, so the apparent link between C. aridus and other species of
Calothamnus provided by C. superbus is in fact only superficial.

C. aridus has an arid zone distribution. The presence of petal indumentum in this species and its
absence IVom other Calothamnus species, including C. superbus, which occur nearer to or within the
floristically diverse transitional rainfall zone of Western Australia where marked spcciation has
occurred (see Hopper 1979), suggests two things. Firstly, that petal indumentum is not required by
Calothamnus species growing in marginal to higher rainfall areas, but may have significant adaptive
value in the arid zone. Petal pubescence in C. aridus probably developed as aresponse to developing
aridity since the separation of Australia and Antarctica. Secondly, based on our knowledge of the
floristic anatomy and distribution of Calothamnus, the evolutionary relationship of C. superbus to
C. aridus and to other members of the genus appears to be comprised of a number of lineages arising
from a common ancestor, which reflect evolution in response to local and regional influences.
Determination of the actual arrangement of lineages in would require a detailed cladistic

analysis (e.g. Weston et al. 1984), which is beyond the scope of this paper. However, in a hypothetical
cladogram it is likely that C. superbus and C. aridus would form a common clade, which is generally
removed from all other species of Calothamnus presently known, apart perhaps for a tenuous link with
C. gilesii.

Acknowledgements

We would like to thank Jim Armstrong, Neville Marchant and the staff of the Western Australian
Herbarium (PERTH) for assistance in work associated with the preparation of this paper. The second
author made the initial collection, the type, of Calothamnus superbus while employed on contract with
the Department of Conservation and Land Management (CALM) on a survey of rare flora in CALM’s
Merrcdin District, a project funded jointly by the Australian National Parks and Wildlife Service and
CALM. F. Mollemans wishes to acknowledge the funding bodies and to thank the project supervisors
DrD.J. Coates and MrP. Brown for their support and encouragement during that project. Constructive
criticisms by an anonymous referee pointed out aspects of this paper requiring improvement.

318

Nuytsia Vol. 8, No. 3 (1992)

References

Briggs, J.D. and Leigh, l.H. (1988). "Rare or Tlireatened Australian Plants." Revised edn. (Australian National Parks and Wildlife
Service, Canberra.)

Hawkeswood, T.J. (1984a). Nine new species of Catotliamruis Labi 11. (Myrtaceae: Leptospennoideae) from Western Australia.
Nuytsia 5:123-153.

Hawkeswood, T.J. (1984b). Calothamnus accedens T.J. Hawkeswood (Myrtaceae), a rare and endangered new species from
Western Australia. Nuytsia 5:305-310.

Hawkeswood, T.J. (1987). A taxonomic revision of the genus Calothamnus Labill. (Myrtaceae: Leptospermoideae). Part 1. The
4-merous species. Nuytsia 6:67-126.

Hopper, S.D. (1979). Biogeographical aspects of speciation in the south-west Australian flora. Annual Rev. Ecol. Syst.
10:399-422.

Weston, P.H., Carolin, R.C. and Annstrong, J.A. (1984). A cladistic analysis of Boronia Sm. and Boronella Baill. (Rutaceae).
Austral. J. Bot. 32:187-203.

Nuytsia 8(3): 319-321 (1992)

319

A new species of Guichenotia (Sterculiaceae) from south western

Australia

Gregory J. Keighery

Department of Conservation and Land Management, Western Australian Wildlife Research Centre,
PO Box 51, Wanneroo, Western Australia 6065

Abstract

Keighery , Gregory , J. A new species of Guichenotia (Sterculiaceae) from south western Australia.
Nuytsia 8(3): 319-321 ( 1 992). A new species of Guichenotia is described: G. alba Keighery confined
to heathland between Cataby and Three Springs north of Perth. A key to all species is included.

Introduction

Guichenotia J. Gay (Sterculiaceae) is a small genus of 7 species, all of which are confined to south
western Australia. The genus is distinguished from the 7 other genera of the tribe Lasiopetalae by
having leafy stipules (giving the leaves a whorl like appearance) and the sepals being prominently
ribbed after flowering. Members of the genus also have distinctive pendant "bell like" flowers.
Members of the genus are widely cultivated in Australia (Wrigley and Fagg 1979) and to a limited
extent overseas. This distinctive taxon was segregated from the widespread Guichenotia sarotes,
when the author sorted into single taxa the taxonomically poorly known groups recorded for the Mount
Lesueur report. This species was li.sted in the subsequent report (Griffin et al. 1990) as Guichenotia
sp. {EA. Grijfin 858). The purpose of this paper is to provide a valid name for this taxon.

Key to species of Guichenotia

1. Petals absent G. apetala

1. Petals present 2

2. Style glabrous in upper half 3

3. Stipules considerably smaller than leaves G. angustifolia

3. Stipules considerably larger than leaves 4

4. Raceme of 4-8 flowers, calyx c. 1 cm long G.ledifolia

4. Raceme of 1-3 flowers, calyx, 2-3 cm long G. macrantha

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Nuytsia Vol. 8, No. 3 (1992)

2. Style stellate-tomentose in upper half 5

5. Stipules leaf like, but smaller than leaves, calyx large, 10-15 cm long 6

5. Stipules small or absent, cordate, calyx small, 5-6 cm long G. micrantha

6. Flowers white; spreading (often almost prostrate) multi-stemmed shrub G. alba

6. Flowers pink; single-stemmed, erect shrub G. sarotes

Guichenotia alba Keighery sp. nov. (Figure 1)

Guichenotia alba Keighery; a G. sarotes flores alba decolor brunneolus, laxi-multi caulis e basi
ramosae statim dignoscenda.

Typus: 5 kilometres south of Cataby, at the intersection of Mimmegarraroad and the Brand Highway,
Western Australia(30°45’S115°30’E), 10 July 1988, G.7./c'eig/;co’10270(holo:PERTH; iso: CANB).

A slender lax few-branched multi-stemmed from a woody rootstock, stems to 40 cm long,
plantto 10cm hdl. Young branches densely stellate-tomentose, becoming glabrous with age. Leaves
alternate, petiole c. 0.5 mm long, lamina (9)13-22 mm long with revolute margins, densely stellate-
tomentose when young becoming glabrous and reticulately veined above. Stipules leaf-like,
approximately 1/2 -2/3 length of the leaves. Racemes simple, loose, in upper axils, 1-2 flowered,
pedicel 6-9 mm, recurved, peduncle 3-6 mm, both densely stellate-tomentose. Bracts and bracteoles
linear-obovate, apex acute. Co/y.r campaji ulate, white outside, pale green inside, lobes 1 1- 13 cm long,
with 3 central veins outside reticulate veined, stellate hairs on the veins, apex acute. Petals small and
scale-like. Stamens 2-2.5 mm long, red-brown purple, apex pore area white. Ovary 1-1.5 mm long,
covered with red glandular hairs. Style ± 2 mm long, with a ring of stellate hairs below stigma. Fruit
crustaceous, loculi are seeded, 4-5 mm diameter. Seeds pale brown, smooth, strophiolate, ± 4 mm
long.

Other specimens examined. WESTERN AUSTRALIA: Cockleshell Gully, W.E. Blackall 3554
(PERTH); Cockleshell Gully, W.E. Blackall 3619 (PERTH); Three Springs, W.E. Blackall 4390
(PERTH); Diamond of the Desert Spring, C.A. Gardner 9103 (PERTH); Cockleshell Gully to Lake
Logue, C.A. Gardner 9382 (PERTH); One kilometre west of Brand Highway on Mimegarra Road,
R. Hnatiuk 1465 (PERTH); 3.5 kilometres ea.st of Lake Indoon, E.A. Griffin 858 (PERTH).

Distribution. Restricted to the northern heathlands of southern Western Australia, between Three
Springs and Cataby.

Habitat. Guichenotia alba grows in low open heath usually on sandy clay or duplex soils in
depressions which are winter wet.

Flowering time. July to August. Mature fruits in November.

Conservation status. Not considered rare or endangered, but surveys are needed to determine this
species occurrence in reserves in the region.

Affinities. This species is closely related to G. sarotes (under which the collections were previously
placed), but can be immediately distinguished (when alive) by the white not pink flowers.

Gregory J. Keighery, Guichenotia

321

Figure 1. Guichenotia alba K&'ighery a-abaxial leaf surface, b-adaxial surface, c- habit, d - flowerfrom below, e - flower,
f - detail of flower with calyx removed, g - anther. Drawn from the type.

Additionally it differs in being an open, spreading (often almost prostrate) multi-stemmed shrub
whereas G. sarotes is a single-stemmed erect shrub to 1.5 metres tall.

Etymology. The species name refers to the pale white flowers, which are unique for the genus.

Acknowledgements

The distinctiveness of this taxon was first recognised after it was collected during the Banksia
sandplain survey by the W.A. Wildflower Society.

References

Griffin, E.A., Hopper, S.D. and Hopkins, A.J.M. (1990). "Flora" In Nature Conservation, Landscape and Recreation values of
the Lesueur area. Eds Burbidge, A.A., Hopper, S.D. and Van Leeuwin, S. E.P.A. Bulletin 424. pp. 39-70.

Wrigley, J.W. and Fagg, M. (1979). Australian Native Plants. (Collins, Sydney.)

Nuytsia 8(3): 323-332 (1992)

323

Four new Drosera taxa from south western Australia

Allen Lowrie' and Neville Marchant^

■6 Glenn Place, Duncraig, Western Australia 6023
^Western Australian Herbarium, Department of Conservation and Land Management,
PO Box 104, Como, Western Australia 6152

Abstract

Lowrie, Allen and Marchant, Neville. Four new Drosera taxa from south western Australia.
Nuytsia 8(3): 323-332 (1992). Three new species and one new subspecies oi Drosera (Droseraceae)
are described, D. browniana and D. stolonifera subsp. monticola (both tuberous Drosera) and
D. grievei and D. sargentii (both pygmy Drosera), all endemic to south western Australia. The
distinguishing characters of each are presented as well as their relationships and an indication of
habitat preferences and conservation status.

Introduction

The south west of Western Australia is renowned for its richness of vascular plant species and its
high degree of endemism. Fifty-four species of Drosera were recorded in Australia by Marchant and
George ( 1982); of these, 42 species were then known to occur in the south west of Western Australia.
The number of species now known to occur in this region is 68.

Since the publication of the first two volumes of Carnivorous Plants of Australia (Lowrie 1987,
1989), Allen Lowrie has made extensive field studies and has discovered a number of new Drosera
taxa. As well as extensive field studies, he has propagated and cultivated all known species. Neville
Marchant has made extensive herbarium studies including examination of almost all Drosera type
specimens. In an endeavour to combine information from these varied sources the present authors
have collaborated to circumscribe and publish new taxa.

Descriptions

1. Drosera browniana A. Lowrie & N. Marchant, sp. nov. (Figure 1)

Drosera bulbosa Hook, subsp. biilbosa affinis sed flore in statu fructificanti erecto, petalis
obovatis apice truncato-crenato, extus roseis, intra albis, 7-10 mm longis, 4-6.5 mm latis.

Typus: 0.9 km south of the rock cairn on Hatters Hill goldmine, 0. 1 km west of the road on the summit
of a granite outcrop, Western Australia, Allen Lowrie 99, 2/9/90 (holo: PERTH; iso: MEL, RSA).

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Nuytsia Vol. 8, No. 3 (1992)

Figure 1. Drosera browniana A. Lowrie and N. Marchant

A - mature plant in flower, B - sepal; C - petal; D - ovary with styles; E - part of style and stigmas, enlarged;
F - leaf lamina; G - T.S. of leaf lamina as indicated on F. Scale bar for all = 1 mm

Allen Lowrie and Neville Marchant, Four new Drosera

325

Tuberous herb. Underground stem 4-6 cm long. Leaves all in a flat basal rosette; lamina obovate,
18 mm long, 10-12 mm wide; petiole 7 mm long, 3 mm wide. Scapes 1-20, 4-10 cm long, single-
flowered, erect in fruit. Calyx lobes ovate, united at the base, margin entire, apex serrate, 4.5 mm
long, 2 mm wide. Petals distally pink, proximally white, obovate, apex truncate-crenate, 7-10 mm
long, 4-6.5 mm wide. Ovary ovoid, 1.5 mm long and 1.3 mm in diameter at anthesis. Styles 3,1 mm
long, branched repeatedly from near the base into numerous filiform segments, forming a dense,
rounded tuft; stigmas subclavate.

Affinities. Drosera hrowniana possesses a tuber with a vertical stolon with prophylls and a rosette of
flat leaves with a broad petiole; stipules are absent. Thus, it belongs in Drosera subg. Ergaleium DC.
sect. Erythrorhizae (Planchon) Diels.

D. macrophylla Lindley andD. bulbosa Hook, and its subspecies are morphologically similar to
D. browniana. The three taxa can easily and reliably be differentiated by the characters which are
presented in the following synoptic key:-

1. Leaves of the rosette up to 12 cm long at maturity

2. Scapes 4-6-flowered; sepals entire D. macrophylla

2. Scapes 1 -flowered; sepals serrate D. bulbosa subsp. major

1. Leaves of the rosette less than 3 cm long at maturity

3. Styles divided; scapes erect in fruit; petals pink and white;

flowers August-September D. browniana

3. Styles undivided; scapes secund in fruit; petals white; flowers

May-June D. bulbosa subsp. bulbosa

Distribution. Drosera browniana is a widely scattered species and is quite common on soils associated
with the narrow greenstone belt from Mt Holland (Lat. 32° lO’S, Long. 1 19° 44’E), to Hatters Hill
(Lat. 32° 49' S, Long. 119° 59' E), a distance of 85 km.

Habitat and phenology. Drosera browniana grows in small depressions which are filled with brown
loam soil on weathered granitic rock outcrops. The outcrops are generally only a metre or so higher
than the surrounding country and aie usually concealed in thickets of gimlet trees (Eucalyptus
species). Rainfall in this region is generally low and restricted to late autumn and winter (May to
August).

The leaves of Drosera browniana appear from the tubers in June and flowering occurs in August
and September. In the dry period of the year Drosera browniana survives as dormant underground
tubers until the onset of winter rains.

Other collections. WESTERN AUSTRALIA: Hatters Hill, intersection of pipeline track and main
Hatters Hill bypass road. Western Australia, Lat. 32° 49’ S, Long. 119° 59’ E, Paul G. Armstrong,
24 Aug. 1990 (PERTH).

Conservation status. A relatively widespread species in native bushland east of cultivated areas and
not under current threat.

Etymology. This species is named after Andrew Brown, an enthusiast of native orchids and officer
of the Department of Conservation and Land Management who discovered the first population.

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2. Drosera stolonifera Endl. subsp. monticola A. Lowrie & N. Marchant, subsp. nov. (Figure 2)
r)'pMi. SummitofToolbrunupPeak,StirlingRange National Park,P. Marti; 14/1 1/89 (holo:PERTH).

Drosera stolonifera Endl. subsp. compacta N. Marchant affinis sed stolonibus et tuberibus
hypogaeis in statu mature persaepe praesentibus, caulis paile hypoggaeis 3.5 cm longis, stolonibus
adventitiis parum abbreviatis, petalis roseis usque ad 4 mm longis, ovario elliptico.

Tuberous herb with a vertical stolon up to 3.5 cm long which may branch to produce additional
terminal tubers. Above ground stem more or less erect, 2-7 cm long; leaves basal and cauline, stem
usually not developed in non-flowering specimens. Basal leaves rosetted, lamina spathulate, 4 mm
long, 6 mm wide, more or less flat; petiole flattened, 4 mm long, 1.5 to 2 mm wide, dilated in upper
part. Cauline leaves usually scattered, rarely sub-opposite or whorled, lamina flabellate, 5 mm long,
8 mm wide, distinctly concave; petiole 10 mm long, 1 mm wide, channelled above. Flowers terminal.
Scape few-flowered, 5 cm long, glabrous; pedicels 8 mm long. Calyx lobes united at the base, ovate,
4 mm long, 2 mm wide, lower margin entire, upper margin and apex irregularly dentate-crenate.
Petals obovate, 8 mm long, 5 mm wide, pale-pink with dark-pink closely spaced flabellate veining.
Ovary ellipsoid, 1.3 mm long, 1.2 mm in diameter at anthesis. Styles 3,0.8 mm long, each divided
into many terete segments, half of them forming an irregular horizontally spreading whorl, the
remainder erect; stigmas terete.

Distribution. Drosera stolonifera subsp. monticola is common on the summits of Toolbrunup Peak
and Bluff Knoll. It is likely to occur on the summits of other peaks in the eastern part of the Stirling
Range.

Habitat. Restricted to winter moist, black or brown loamy soils on ledges and in shallow depressions,
often growing with moss. Only recorded from the upper slopes and summits which are frequently
cloud-covered and are usually very moist.

Flowering period. Only recorded for November. The flowering period is probably between October
and December.

Affinities. Drosera stolonifera subsp. monticola belongs in subg. Ergaleium DC. sect. Stolonifera
DeBuhr because it possesses a tuber and has leaves in a basal rosette as well as cauline leaves.

Drosera stolonifera subsp. compacta N. Marchant is considered to be the taxon most similar to
Drosera stolonifera subsp. monticola. The former subspecies occurs in heathland from the Stirling
Range to the south coast and can be readily separated from the new subspecies by its flower colour,
stolon morphology and petiole length. The morphological differences between the two taxa are
presented in the following synoptic key:-

1. Petals pink; additional below-ground stolons and tubers
often present at maturity; petioles up to 4 mm long; ovary

ellipsoid D. stolonifera subsp. monticola

1. Petals white; additional below-ground stolons and tubers
absent, the lower stem prostrate; petioles 5-30 mm long;
ovary subglobose

D. stolonifera subsp. compacta

Allen Lowrie and Neville Marchant, Four new Drosera

327

Figure 2. Drosera stolonifera subsp. monticola A. Lowrie and N. Marchant

A - mature plant in flower; B - mature non-flowering plant; C - sepal; D - petal; E - ovary with styles;

F - part of style and stigmas, enlarged; G - cauline leaf lamina; H - basal leaf lamina; 1 - T.S. of cauline
leaf petiole as indicated on G. Scale bar for all = 1 mm

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Conservation status. Relatively common in the Stirling Range National Park where it is apparently
not under threat.

Etymology.This new subspecies is restricted to the upper parts of the Stirling Range hence the epithet,
monticola, referring to its mountain habitat.

3. Drosera grieve! A. Lowrie & N. Marchant, sp. nov. (Figure 3)

Drosera paleacea DC. subsp. paleacea affinis sed scapo uno usque ad 3 cm longo, 5-10-floris,
sepalis perlato-obovatis 2.5 mm longis, 2.5 mm latis.

Typus: At the east end of Baanga Hill road, near the junction of Sennetts Lake Road, c. 20 km east
of Lake King township. Western Australia, A Lowrie 25/9/89 (holo; PERTH; iso: MEL).

Fibrous rooted herb forming a compact convex rosette to 1.2 cm diameter. Stem to 2 cm long,
covered with the withered remains of the previous seasons growth. Leaves 20-30. Lamina orbicular,
subpeltatc, 0.7 mm diameter; petiole 3.6 mm long, 0.5 mm wide at the ba.se, dilated to 0.8 mm wide
in the lower part, 0.2 mm wide at the apex. Stipule bud ovoid, 3 mm long, 2 mm diameter, with setae.
Stipules 3-lobed, 5 mm long, 4 mm wide; the central lobe divided into 2 segments, each segment in
turn divided into two segments narrowing to setae, lateral lobes with a short segment on the outer
margin near the base, and two asymmetrically bifurcate apical lobes. Inflorescence racemose; scape
1, to 3 cm long, covered with minute short broad glands; flowers 5 to 10; pedicels 2.5 mm long, semi-
erect in fruit. Calyx lobes very broadly obovate, united at the base, 2.5 mm long, 2.5 mm wide, apex
irregularly dentate. Petals white, oblong,4.0-4.5 mm long, 2 mm wide. Ovary turbinate, 0.7 mm long,
1mm diameter at anthesis. Styles 4, spreading horizontally, 0.5 mm long, 0. 1 mm diameter; stigmas
falcate, 1.2 mm long, 0.15 mm diameter, apex subacute. (Figure 3)

Distribution. Known only from two locations, 30 km apart, in the eastern wheatbelt south-east of
Hyden.

Habitat. In clayey sand on heathland under and between low shrubs.

Affinities. D. grievei belongs in Drosera subg. Rorella (DC.) Diels, sect. Lamprolepis Planchon
because it has rosetted leaves, prominent stipules, many-flowered scapes and it reproduces by
propagules. D. paleacea DC. subsp. paleacea is considered the closest relative of Drosera grievei
but differs from it in the features presented in the synoptic key below:-

1. Scapes 5-10-flowered; sepals very broadly obovate, 2.5 mm

long, 2.5 mm wide. Petals oblong, 4-4.5 mm long D. grievei

1. Scapes 30 or more-flowered; sepals obovate, 1.5 mm long,

0.7 mm wide. Petals obovate, 3-4 mm long D. paleacea subsp. paleacea

Conservation status. Threatened. Currently known from only two roadside locations in an area which
has been extensively cleared for farmland.

Allen Lowrie and Neville Marchant, Four new Drosera

329

Figure 3. Drosera grievei A. Lowrie and N. Marchant

A - mature plant in flower; B - sepal; C - petal; D - ovary with styles; E - part of style and stigma, enlarged.
F - leaf lamina; G - T.S. of leaf petiole as indicated on F. Scale bar for all = 1 mm

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Etymology. Drosera grievei is named in honor of Brian James Grieve (1907- ), Professor Emeritus,
who has provided the only comprehensive means of identifying the rich flora of southern Western
Australia. From 1954 Professor Grieve has produced the outstanding series of books "How to Know
Western Australian Wildflowers".

4. Drosera sargentii A. Lowrie & N. Marchant, sp. nov. (Figure 4)

Drosera parvula Planchon affinis sed stipula-gemma conicis, 6 mm longa, 4 mm diam., setis
longissimis dense obtectis.

Typus: At the junction of Stockyard Road and Merivale Road, in the S.E. corner, c. 20 km E of
Esperance, Western Australia, A//c/i Lowrie 22/11/89 (holo: PERTH; iso: MEL).

Fibrous rooted herb forming a compact convex rosette up to 1.5 cm diameter. Stem to 2 cm long,
covered with the withered remains of the previous seasons growth. Leaves 20-50. Lamina orbicular,
almost peltate, 0.8 mm diameter; petiole 3.8 mm long, 0.5 mm wide from the base for a third of the
length, the upper part 0.2 mm wide. Stipule bud conical, 6 mm long, 4 mm diameter, with extremely
long setae. Stipules shortly 3-lobed, 1 .5 mm long, 1 .7 mm wide, lobes truncate, the central lobe with
5 short fringes, the lateral lobes with 3 short fringes and a 5 mm long seta adjacent to the mid-lobe.
Inflorescence racemose; scapes 1 to 2, up to 5 cm long, glabrous; flowers 20 to 40; pedicels 1 mm
long, pendulous in fruit. Calyx lobes glabrous, ovate, united at the base, 1.5 mm long, 0.9 mm wide,
margin entire, apex slightly serrate. Petals white, obovate, 3 mm long, 2 mm wide, entire. Ovary
obovoid, 0.4 mm long, 0.5 mm diameter at anthesis. Styles 3. while, semi-erect, 0.3 mm long, 0.1
mm diameter; stigmas while, falcate-clavate, 1.1 mm long, 0.1 mm diameter at the base, dilated to
0.15 mm diameter near the rounded apex.

Distribution. Drosera sargentii is a common species in the coastal regions between Esperance and
Cape Le Grand National Park.

Habitat. In deep, while siliceous soil on heathland. At many locations it grows with the widespread,
white-flowered species Drosera scorpioides.

Flowering period. November-December.

Affinities. D. sargentii belongs in Drosera subg. Rorella (DC.) Diels, section Lamprolepis Planchon
because it has rosetted leaves, prominent stipules, many-flowered scapes and it reproduces by
propagules. D. pan’ula Planchon is considered here to be species most similar Xo Drosera sargentii.
The two taxa differ in the features presented in the following synoptic key;-

1. Stipule bud forming a conical tuft, densely covered with extremely long

setae; pedicels pendulous in fruit; sepals 1.5 mm long; petals 3 mm long D. sargentii

1. Stipule bud forming an ovoid tuft, with short setae covering the lower part
and long setae at the apex; pedicels semi-erect in fruit; sepals 3-4 mm long;
petals 4-6 mm long D. parvula

Conservation status. A common and apparently widespread species not under current threat.

Allen Lowrie and Neville Marchant, Four new Drosera

331

Figure 4. Drosera sargeiitii A. Lowrie and N. Marcliant

A - mature plant in flower; B - sepal; C - petal; D - ovary with styles; E - part of style and stigma, enlarged;
F - leaf lamina; G - T.S. of leaf petiole as indicated on F. Scale bar for all = 1 mm

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Etymology. Drosera sargentii i.s named in honour of Oswald Hewlett Sargent (.1880-1952), a
Pharmacist of York, Western Australia, who made a major contribution to Western Australian botany
including studies on the genus Drosera.

This new species of pygmy sundew was discovered by Allen Lowrie and Steve Rose in 1 988 when
it had just finished flowering. The following season, flowering specimens were found and brought
into cultivation for closer study.

References

Lowrie, A. (1987). Carnivorous Plants of Australia. Volume 1. (University of Western Australia Press, Nedlands, Western
Australia.)

Lowrie, A. (1989). Carnivorous Plants of Australia. Volume 2. (University of Western Australia Press, Nedlands, Western
Australia.)

Marchant, N.G. (1982). Droseraceae. In George, A.S. (ed.) Flora of Australia. Volume 8:383-385. (Australian Government
Publishing Service, Canberra.)

Nuytsia 8(3); 333-350 (1992)

333

New names and combinations for some Melaleuca (Myrtaceae) species
and subspecies from the south-west of Western Australia considered

rare or threatened

F.C. Quinn'^, K.J.Cowley'', B.A. Barlow* and K.R. Thiele®

‘Australian National Herbarium, Division of Plant Industry, CSIRO, GPO Box 1600, Canberra, ACT 2601
^School of Botany, University of Melbourne, Parkville, Victoria 3052

Abstract

F.C. Quinn, K.J. Cowley, B.A. Barlow and K.R. Thiele. New names and combinations for some
Melaleuca (Myrtaceae) species and subspecies from the south-west of Western Australia considered
rare or threatened. Nuytsia 8(3): 333-350 (1992). The paper describes or makes new combinations
for several species and subspecies of Melaleuca considered to be rare or threatened. Six species and
two subspecies, Melaleuca araucarioides, M.fissurata, M. ordinifolia, M. pomphostoma, M. ringens,
M. sculponeata, M. viminea subsp. appressa andM. huegelii subsp. prisiicensis are described as new.
M. tenella Bentham is reduced to a subspecies of M. incana R. Br. and M. densa var. pritzelii Domin
is raised to specific rank.

Introduction

Like many other medium-sized and large genera of the Autochthonous Sub-element of the
Australian flora, Melaleuca has a major centre of species richness in the South west Botanical Province
of Western Australia. It is probable that the genus comprises 250-300 species, of which about half
probably occur in the Southwest Botanical Province. As a consequence, much of the systematic
research necessary to provide a comprehensive inventory of the genus is concentrated on this region.
Our research indicates that many of the species in this area have relatively small distribution areas.

Among the many taxa which have been distinguished, there are several which appear to require
documentation as rare or threatened. The purpose of this paper is to make names available for a
number of these species and subspecies, as a step in expediting full documentation of the conservation
status of each. Some of these have already been listed, as undescribed species, in the work on rare
or threatened Australian plants by Briggs and Leigh (1988). Others are recognised as new for the first
time in this paper, and conservation codes are suggested for them also.

The recommendations for conservation status suggested here are in accordance with the criteria
set out by Briggs and Leigh (1988). With respect to the species dealt with here, the following
designations are relevant:

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Nuytsia Vol. 8, No. 3 (1992)

Code

Explanation

2

3

K

C

E

V

geographic range less than 100 km

geographic range more than 100 km

poorly known

population reserved

endangered

vulnerable

adequacy of reserve unknown

Of the ten taxa dealt with in this paper, the codes which have been suggested include 2K ( 1 taxon),
2KC- (1), 2E (2), 2V (1), 3K (2), 3KC (1), and 3KC- (2).

All of the species and subspecies dealt with here occur in the Southwest Botanical Province of
Western Australia. The four letter codes used in qualifying the geographic occurrences of the taxa
refer to the natural regions of Australia delineated by Barlow (1987). These codes are useful in
quantifying the distributions of plant species on an Australia-wide basis. Because these taxa all occur
in an area where there is now a very complex mosaic of natural vegetation and cleared agricultural
land, there is clearly an urgent need for additional field work in order to clarify the conservation status
of each taxon.

Taxonomy

The taxa dealt with below have been arranged in alphabetical order. Because our revision of
Melaleuca is still in progress, we have not yet finalised an infra-generic classification of the genus.
The classification of Bentham ( 1 867) appears to be artificial, and we have therefore avoided referring
species to Bentham's sections. The taxa dealt with here belong to a number of distinct species groups
within the genus. These species groups all have geographie distributions which are essentially
temperate Australian. They also all have centres of species richness in the south west province of
Western Australia.

1. Melaleuca araucarioides Barlow, sp. nov. (Figure la)

Speeies nova M. hlaeriifoliae Turez. proxima, sed foliis ternatis, stylis brevioribus, fructibus
plerumque minoribus differ!.

Holotypus: Western Australia: Esperance Region: About 1 km W of Kog’s Corner on the Cape Riche
to Highway road, 34°36’S 1 18°45'E, Cowley & Quinn 15 1, 7.x. 1988 (CANB 383935). Isotypi: BRl,
K, MEL. PERTH.

Shrub to 1.5 m high, young shoots lanuginose and the rachis occasionally tomentose, otherwise
glabrous. Leaves ternate, semi-appressed and crowded in 6 regular rows along the axis; lamina
narrowly elliptic to narrowly obovate, slightly rounded abaxially and flat or channelled adaxially,
L7-3.5 mm long, 0.7-1. 8 mm wide, obtuse and inflexed at the apex, slightly reflexed and attenuate
at the biise into a petiole 0.3-0.8 mm long; venation obscure; glands obscure or slightly pustulate.
Inflorescence a basal or terminal sp ike or head of 8- 1 7 crowded monads, to 1 0 mm long, term inal heads
often predominantly male flowers; bracts 0.8- 1.3 mm long, 0.4-0.8 mm wide, persistent to flower

F.C. Quinn et al.. Rare or threatened Melaleuca

335

senescence. Hypanlhium cup-shaped, rounded al (he base, 0.8- 1 .2 mm long. Sepals 0.5-0.8 mm long,
membranous at the margin, persistent to mature fruits. Stamens 1.8-5 mm long including a claw
0.8- 1.4 mm long, 3-4(5) per bundle, filaments cretun to yellow. Style to 5.5 mm long; ovules c. 22 per
loculus. Fruits shortly bell-shaped, compressedby mutual pressure, 2.5-3.5 mm long, 2-4.3 mm diam.
with thickened, rounded and outspread persistent sepals.

Distribution and ecology. Confined to the Ongerup-Cape Riche area, NW of Albany (ESPR) with a
geographic range on current knowledge of less than 100 km. Occurs in well drained sandy or loamy
soils in heaths or open eucalypt woodlands. Flowering has been recorded in July and October.
Figure 2a.

Distinction from related species. Melaleuca araucarioides is closely related to M. blaeriifolia, from
which it can readily be distinguished by its ternate phyllotaxy, shorter style and smaller fruits. In
M. blaeriifolia the style is 8-9 mm long and the fruits arc 3.5-5 mm in diameter and 3.5-4 mm long.
The very regular arrangement of the leaves into 6 rows is particularly distinctive.

Conservation status. We recommend that the conservation status of this species should be Poorly
Known (code 2K of Briggs and Leigh 1988) as it is only known from eight collections, only two of
which have been made in the last fifteen years. It also appe;irs to be geographically restricted and from
our field ob.servations is probably rare.

Etymology. The specific epithet alludes to the superficial resemblance of the leafy shooLs to those of
the gymnosperm genus Araucaria Juss.

Figure 1. a, Melaleuca araucarioides. portion of plant {Cowley & Quinn 151). b, M.fissurata, portions of plant (left piece,
van der MoezelPGW \20-. right piece, VVt/ion 7000). Scale bars = 1 cm

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Nuytsia Vol. 8, No. 3 (1992)

Additional specimens examined. WESTERN AUSTRALIA: c. 6 km S of Jerramungup on road to
Albany, Cowley & Quinn 146, 7.X.1988 (CANB); 10 miles (16 km) N of Cape Riche, Gardner s.n.,
12.x. 1942 (PERTH); 8 miles (13 km) SE of Ongerup, Aewliey 1277, 12.vii.l964 (PERTH); 10 km
S of Boxwood Hills, Newbey 4264, 19.vii.l974 (PERTH); Kojaneerup East Road, Pfeiffer s.n.,
18.vii.l969 (PERTH); cultivated at Glenmorgan, Qld, from seed collected at Cape Riche, Gordon
2358, X.1962 (PERTH); Nova Hollandia, Drummond 5th coll. 167, no date (KW, photo in CANB).

2. Melaleuca fissurata Barlow, sp. nov. (Figure lb)

Species nova M. lateriflorae Benth. proxima, sed fructibus aspri fissuratis, foliis carinatis
retroflexis brevioribus differ!.

Holotypus: Western Australia: Esperance Region: 29 km SS W of Tadpole Lake, c. 65 km ENE of
Lake King, 32°32’S 119°54’E, Aew/iey 5535, 13.viii.l979 (PERTH).

Low shrub to 1.5 m tall, rough-barked, the inflorescence minutely pubescent, otherwise glabrous.
Leaves spirally arranged, spreading; lamina broadly elliptic, somewhat thickened and keeled
abaxially, 4.0-5.0 mm long, 1. 5-2.0 mm wide, obtuse at the apex, broadly tapering to a stout petiole
0.5-0.7 mm long, veinless or with 1-2 faint, fine lateral veins on either side of the keel; glands obscure
or prominent, scattered. Inflorescence a lateral cluster of 1-5(10) flowers on old wood; bracts ovate,
c. 1 mm long, c. 0.8 mm wide, persistent to anthesis. Hypanthium broadly cup-shaped, 1.2-1.5 mm
long, glabrous, smooth or verrucose. Sepals rounded-triangular, 1.2-1. 4 mm long, deciduous before
fruit maturity. Stamens white, 5.0-10.0 mm long including a broad claw c. 1.5 mm long, 10-12 per
bundle. Style c. 5.5 mm long; ovules not seen. Fruit cup-shaped, c. 4 mm long, 5.5-8.0 mm diam.
with a rough, fissured, dark bark-like surface; valves enclosed.

Distribution and ecology. Known from Mt Ney to Lake King, N of Esperance in the South West
Botanical Province of Western Australia with a geographic range on current knowledge of c. 260 km.
Occurs in shrub mallee or woodland on sand or sandy loam usually over clay or clay loam. Flowering
is predominantly in July and August. (Figure 2b)

Distinction from related species. A/e/a/eMca/issurata is most closely related to M. /atenyiora differing
markedly in its very roughly textured, fissured, corky fruit and keeled, retrollexed, shorter leaves.
M. lateriflora has erect, sometimes slightly channelled leaves (5)7-20(25) mm long. The two species
overlap for a small part of their ranges but, maintain their morphological distinction, reinforcing the
appropriateness of specific rank for Melaleuca fissurata.

Conservation status. We support Briggs and Leigh (1988) who cited this species asM. sp.2 (Mt. Ney-
Lake King) in assigning a conservation status of Poorly Known (code 3K) as it is only known from
7 collections. This species appears to occur in small populations in apparently localised and specific
habitats making it very vulnerable to disturbance. Further field work is needed to confirm its status,
particularly to examine whether or not the Lake King population is disjunct from other known
populations.

Etymology. The specific epithet is derived from the Latin fissuratus (fissured) and refers to the surface
texture of the fruit.

F.C. Quinn et al.. Rare or threatened Melaleuca

337

Figure!. Distributions of the species shown as occurrences in 1° cells. n,M. araucarioides. b,M.fissurata. c,M. huegelii
subsp. pristicensis with the outline showing the range of the typical species, d, M. incana subsp. tenella with the outline
showing the range of the typical species. e, M. ordinifoUa. f, M. pomphostoma.. g, M. pritzelii. h, M. ringens.
i, M. sculponeala. j, M. viminea subsp. appressa with the outline showing the range of the typical species.

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Nuytsia Vol. 8, No. 3 (1992)

Additional specimens examined. WESTERN AUSTRALIA; 33 km WSW of Lake King township,
WilsonlOOO, lO.viii. 1968 (CANB, PERTH); 13km ENEofScaddenonCoxRd, vwirferA/oeze/j.n.,
Lvii.1984 (PERTH); Ridley Rd, 5.4 km from Ridley-Truslove Rd junction, c. 20 km NEof Scadden,
van derMoezel 120, 17-18.viii.1982 (CANB, PERTH); 9.2 km due SSW of Mt Ridley, BMigman &
McNee 1863, 8.viii.l983 (PERTH); 15 km due SW of Mt Ney, 5 km NE of Burdett Rd on Mt Ney
Rd, Burgman & McNee 1657, 3.viii.l983 (CANB, PERTH); 30.75 km N of Mt Beaumont, 9.58 km
NW of Mt Ney Rd on Clyde Rock Rd, Burgman & Layman 3061, no date (PERTH).

3. Melaleuca huegelii subsp. pritsicensis Barlow, subsp. nov. (Figures 3a-e)

Subspecies nova M. huegelii Endl. subsp huegelii proxima, sed rhachidi breviore, floribus
malvinis paucioribus differt.

Hoiotypus: Western Australia: Murchison Region: Tamala Station, 26°42’S 113°43’E,5rarr/6800,
11.X.1973 (PERTH). Isotypi: PERTH, CANB.

Spreading shrub or tree 0.9-2 (rarely to 5) m tall, inflorescence axis pubescent at flower bases only
and the bracteoles, hypanthium, anthers and stems with few long hairs, otherwise glabrous. Leaves
spirally arranged; lamina ovate or triangular, flat to weakly keeled, 1. 8-4.0 mm long, L5-2.5 mm
wide, acute to acuminate at the apex , slightly contracted and sessile or weakly peltate at the base; veins
several, sometimes obscure; glands fine, in rows. Inflorescence subbasal on lateral stems or terminal
on main axes, a spike of 1-12 triads spirally arranged on an axis (5)10-20(30) mm long; triads with
the central flower higher than the laterals, forming a triangular group; bracts broadly triangular, c.

2.5 mm long, c. 2 mm wide, foliose, green to scarious, deciduous; bracteoles nttrrowly triangular, c.

1.5 mm long, (0.2)0.6- 1 .2 mm wide, deciduous. Hypanthium cup-shaped, c. 2 mm long, c. 2 mm wide,
ribbed. Sepals triangular, 0.8- 1.0 mm long, persistent to mature fruit. Stamens mauve to pink, 2-3
mm long including a claw 3-5 mm long and 0.5-0.6 mm wide, 9-13 per bundle. Style 8-9 mm long;
ovules c. 12 per loculus. Fruit globular, c. 3 mm long, 3. 5-4.0 mm diam., weakly lobed, smooth to
papery in texture; valves deeply reces.sed.

Distribution and ecology. Restricted to Dirk Hartog Island and around Tamala Station, in the south
end of Shark Bay in South West Botanical Province of Western Australia (MRCH). Although this
subspecies has a geographic range on current knowledge of more than 100 km, existing records show
that the distribution may be di.sjunct. It occurs in heathlands in sand. Flowering is predominantly in
September and October. (Figure 2c)

Distinction as a subspecies. In Af. huegelii, the triangular rather than linear arrangement of the flowers
in each triad is distinctive, and the species may be more closely related to other species which have
this pattern, including M. adnata Turez. and M. eleuterostachya F. Muell., than with other species
having more or less peltate leaves. The populations from the northern part of the range can be readily
distinguished in several characters. Strong ecogeographic divergence is indicated, and two
subspecies have been distinguished accordingly. The new subspecies differs consistently from the
typical one in its much shorter and fe wer-tlowcrcd inflorescence, and in the mauve to pink rather than
white or cream staminal filaments. The typical M. huegelii has inflorescence axes of (25)40-80(130)
mm and usually 50- 140 triads in each flower .spike. The sharp distinction between the two subspecies
suggests that substantial genetic divergence has occurred. The differences shown by subsp.
pristicensis are presumably adaptive, reflecting a lower level of resource allocation to seed production
and perhaps a different pollinator.

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Figure 3. Melaleuca hnegclU subsp. pristicensis. a, portions of plant (George 11566). b-e. Scanning Electron Micrographs
(SEMs), showing variations in leaf fomi. b, abaxial surface of leaf from indetemiinate shoot (George 11566).
c, indetemiinate shoot (George 11566). d, abaxial surface of leaf from lateral shoot (George 11566). e, lateral shoot
(George 1 1566). Scale bars = 1 cm

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Conservation status. We recommend that the conservation status of this speices should be Poorly
Known (code 3K of Briggs and Leigh, 1988). Even though the geographic range for this species is
more than 100 km, it seems to occur only in small populations at each locality in apparently localised
and specific habitats.

Etymology. The subspecific epithet is derived from pristix (shark) and refers to its occurrence in the
Shark Bay area.

Additional specimens e.xaminecl. WESTERN AUSTRALIA: MRCH: just S of Cape Inscription, Dirk
Hartog Island, George 1 1 566, 5. ix. 1972 (C ANB , K, MEL); inland of Sandy Point, Dirk Hartog Island,
jB card 709 1,18.x. 1974 (PERTH) ; bet ween Carraran gandTamala.R card 7079, 16.x. 1974 (PERTH) ;
16 miles SW of Tamala, Demarz 5169, 8.ix.l974 (PERTH); 22 km on Tamala Kd, Demarz 8998,
2.x. 1981 (PERTH); 3 miles W of Tamala, Davies s.n., 13.x. 1960 (PERTH).

4. Melaleuca incana subsp. tenella (Bentham) Barlow, comb, et stat. nov. (Figure 4a)

Melaleuca renei/a Bentham , FI. Aust. 3 : 1 60 ( 1 867) ; Myrtoleucodendron tenellum (Bentham) Kuntze,
Revis. Gen. PI. 241 (1891). Type: Western Australia: Phillips River, Maxwell s.n. (holo: K; iso:
CANB, MEL 602187, MEL 602188).

Shrub 1 m, the inflorescence axis tomentose, otherwise glabrous. Leaves ternate or displaced
ternate, often slightly keeled, reflexed from the petiole and incurved towards the axis; lamina narrowly
ovate, flat to compressed, (3.5)7-9 mm long,0.5-0.9 mm wide, obtuse and often thickened at the apex,
attenuate at the base into a petiole 0.5- 1 mm long; venation obscure or only the midvein evident; glands
10-15 on the abaxial surface towards the centre of the lamina, uniformly large. Inflorescence a basal
or terminal spike or head of 10-35 crowded monads, 5-25 mm long, with the terminal heads often
predominantly of male flowers; bracts 1.2-2 mm long, 0.9-1. 3 mm wide, persistent to anthesis.
Hypanthium cylindrical with an elongated base in hermaphrodite flowers, conical in male flowers,
1.5-2 mm long. Sepals 0.5-0.7 mm long, membranous at the margins, persistent to mature fruit.
Stamens 3.5-6 mm long including a claw 0.5-1 mm long, (3)4-8(10) per bundle with filaments white
to yellow. Style 5-8 mm long; ovules c. 25-32per loculus. Fruits shortly bell-shaped with an elongated
base, sometimes compressed by mutual pressure, 1.8-4 mm long, 3-5 mm diam., with persistent
rounded thickened outspread sepals.

Distribution and ecology. Found on the coast and in adjacent inland areas from near Esperance to Cape
Le Grande National Park in the south-west of Western Australia (ESPR). Associated with swampy
and moist areas. Flowering is from August to October. (Figure 2d)

Distinction as a subspecies. There is a disjunction in the distribution of M. incana which corresponds
closely with the occurrence of two distinct morphological forms (Figure 2d). Whilst the more easterly
of these was described as a distinct species, M. tenella, by Bentham, the differences appear to be
relatively minor consequences of ccogeographical divergence, and there is some evidence of
introgression (see below). The two entities are accordingly treated as subspecies. Melaleuca incana
subsp. tenella differs from subsp. incana in the glabrous, smaller leaves, as well as in having slightly
shorter stamens, more obscure venation and fewer leaf glands, as summarized in the description
above. M. incana subsp. incana usually has pubescent leaves that are (3.5)7-15(17) mm long and
1-1 .5(3) mm wide and have c. 40-100 small and c. 20 larger leaf glands, while the stamens range from
(3.5)6-8.5 mm long.

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341

Figure 4. a. Melaleuca incana subsp. lenella, iwrtions of plant (Jackson 1273). b, M. ordinifolia , portion of plant
(Burbidge 2451). c, M. poniphosloma, portions of plants (llowering piece, Wittwer 4\5\ fruiting piece, Newbey 4913).
d, M. pritzelii, portion of plant (Cowley & Quinn 108). Scale bars = 1 cm

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There ai'e a few collections from within the geographical range of M. incana subsp. incana which
resemble M. incana subsp. tenella. However, they all show minor differences from the morphology
of subsp. tenella in its disjunct eastern area, and have not been included in this subspecies. It is possible
that the divergence of the two subspecies is relatively old, and may have had a different geographical
basis from that which now di.stinguishes them. Whilst it may not be relevant to this hypothesis, it is
notable that the type localities of M. incana and its subsp. tenella are both outside the known present-
day ranges of the two subspecies as indicated by more recent collections.

Misapplied name. The name M. microphylla has sometimes been misapplied to M. incana subsp.
tenella, which differs from that species in its tomentose rachis and ternate leaves.

Conservation status. We recommend the conservation status of this subspecies should be Poorly
Known (code 2KC- of Briggs and Leigh, 1988) on the basis that there are herbarium specimens
collected from proclaimed reserves but from populations of unknown size. Melaleuca incana subsp.
tenella is poorly known and appears to have a restricted geographic range. This subspecies is known
to be represented by populations within a national park and another reserve, though the adequacy of
reservation is unknown.

Additional specimens examined. WESTERN AUSTRALIA: ESPR; 0.4 km N of Fisheries Rd on
Coolinup Rd, Reserve 27354, Burgman 4407, x.1984 (PERTH); c. 50 km E of Esperance, Kuchel
1674, 1 Lix.1964 (AD, CANB); 7 miles from Esperance at junction ofRavensthorpe and Norseman
roads, Wrigley s.n., 2.xi.l968 (CBG); 6 miles N of Esperance, leg. ign. s.n., 18.ix.l950 (AD); Buyi
Billanak Homestead, c. 12 km SE of Condingup Peak, Jackson 1273, 21. ix. 1968 (CANB); 3 km N
of Duke of Orleans Bay, Wilson 8108, 30.ix.l968 (CANB, PERTH); N of Mount Le Grand, Eichler
21229, 1 Lix.1971 (CANB, AD); near "yate" swamp, 3.4 miles E of Cape Le Grand National rd, N
of Lucky Bay, Weston 9576, 3.x. 1974 (CANB, PERTH); CapeLe Grande, Lm////7z 3565, 23.viii. 1964
(PERTH); between Cape Le Grande and Lucky Bay, Gardner 14120, 2.ix.l962 (CANB, PERTH).

5. Melaleuca ordinifolia Barlow, sp. nov. (Figure 4b)

Species nova M. apodocephalae Turcz. proxima, sed foliis decussatis brevioribus, inflorescentiis
monadibus paucioribus bracteis minoribus, sepalis longioribus, staminibus longioribus pluribus
differt.

Holotypus: Western Australia; Esperance Region; near Hamersley River crossing, Ongerup-
Ravensthorperoad,33°46’S 119°36’E,Ntm75C7 5075, 28.ix. 1977 (CANB 313531). Isotypv.CAEiB,
PERTH.

Low slirub usually less than 1 m tall, the young shoots moderately pubescent with crisped hairs and
soon glabrescent and the inflorescence axis and hypanthium white-pube.scent, otherwise glabrous.
Leaves decussate, nearly erect at 30°, crowded in 4 regular longitudinal series; lamina narrowly ovate,
compressed concavo-convex, straight or slightly incurved, (4.0)4.5-5.5(7.0) mm long, 1.0- 1.5 mm
wide, obtuse at the apex, slightly contracted at the ba.se into a stout petiole c. 0.5 mm long; veins and
glands obscure. Inflorescence a lateral cluster of 1-3 flowers on old wood; bracts triangular to ovate,
to 1.5 mm long and 1. 0 mm wide, persistent to anthesis. Hypanthium broadly cylindric,c. 1.5 mm long.
Sepals obtusely triangular, thick, erect, c. 1 .0 mm long, deciduous before fruit maturity. Stamens white,
5.0-5. 5 mm long Including a very short clawc. 0.5 mm long, 8-12 per bundle. Style 6. 5-7. 5 mm long;

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343

ovules 27-31 per loculus. Fruit shortly cup-shaped, 34 mm long, 4-5 mm diam., dark-coloured,
somewhat rough and fissured below a papery epidermis, slightly sinuate and not narrowed at the
aperture; valves enclosed.

Distribution and ecology. Found in the south-west of Western Australia from Cranbrook to the
Hammersley River (LUWN, ESPR). The range is possibly disjunct. Occurs in mallee shrubland on
loams and clays. Flowering is predominantly from August to October. (Figure 2e)

Distinction fromrelated species. Melaleuca ordinifolia is probably closely related to M. apodocephala
and M. 6rcvi/b//a Turcz., although the latter two have spiral phyllotaxy while the former has decussate
phyllotaxy . Melaleuca ordinifolia is entirely sy mpatric with M. brevifolia and with M. apodocephala
subsp. apodocephala, and appears to completely maintain its identity in the field. It is readily
distinguished from most other decussate-leaved species by its short, compressed, channelled leaves
crowded into four regular rows and by its rough-textured fruit surface.

Conservation status. We recommend that the conservation status of this species, listed by Briggs and
Leigh (1988) as M. sp. 1 (Cranbrook-Hammersley River), should be Poorly Known (code 3KC- of
Briggs and Leigh, 1988) on the basis of a herbarium specimen collected from Stirling Range National
Park but from a population of unknown size. Although Melaleuca ordinifolia appears to have a
geographic range more than 100 km, it seems to only occur in highly specific and localised habitats.
Of the 7 collections seen there is only one collection since 1977 and this was from the Stirling Range
National Park.

Etymology. The specific epithet is derived from the Latin ordinis (methodical arrangement) and
alludes to the regular arrangement of the leaves.

Additional specimens examined. WESTERN AUSTRALIA; LUWN: nearPootenup,/VeH’()ey 1861,
24. ix. 1965 (PERTH); E from Solomons Well, Stirling Range, Morrison s.n., 28. ix. 1902 (PERTH);
sandplain S of Pootenup, Burbidge 2457, lLix.1947 (CANB); Cranbrook, Diels s.n., no date
(PERTH); c. 500 m W of junction of Salt River Rd with Red Gum Pass Rd on Salt River Rd, Stirling
Range National Park, Cowley & Quinn 169, 9.x. 1988 (CANB); cultivated at Glenmorgan, Qld, from
seed collected at Albany, W.A., Gordon 6040, x.i962 (PERTH).

6. Melaleuca pomphostoma Barlow, sp. nov. (Figures 4c)

Species nova M. bracteosae Turez. proxima, .sed stomatibus prominentibus staminibus pluribus
longioribus ungue staminum longiore stylo longiore differt.

Holotypus: Western Australia: Esperance Region: Below W end of Eyre Range, c. 33°51' 119°59',
George 9296 (PERTH). Isolypus: CANB.

Dense shrub to 1.5 m high, young shoots and rachis tomentose, and branchlets minutely
lanuginose, otherwise glabrous. Bark grey, rough and thick. Leaves spirally arranged, spreading;
lamina narrowly elliptical to narrowly ovate, compressed especially towards the apex, (3)5-8(10) mm
long, 1 -2 mm wide, covered with very numerous white pustulate stomata, obtuse at the apex, attenuate
at the base into a petiole 0.5-1 mm long; venation obscure; glands obscure. Inflorescence a basal or
terminal spike of up to 10 crowded monads, up to 15 mm long; bracts and bracteoles to 1 mm long
and 1mm wide, persistent to anthesis. Hypanthium cylindrical with asomewhat elongated base, 2.5-3

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Figure 5. SEMs showing leaf surface characlers. a, c, e, abaxial whole leaf surface, b, d, f, abaxial leaf surface showing
stomata, a.b, M. bracteosa (Barlow 423\). c, d, M. pomphostoma (George 9296), e, f, M. bracteosa x M. pomphostoma
(Short, Amarena & Fuller 2696). Scale bars = 1 cm

F.C. Quinn et at.. Rare or threatened Melaleuca

345

mm long. Sepals 1-1.5 mm long, membranous at the margins, persistent to mature fruits. Stamens
(8)10-14mmlongincludingaclaw 4.5-8 mm long, 12- 18 per bundle, with filaments greenish-yellow.
Style 9-15 mm long; ovules 30-50 per loculus. Fruits broadly bell-shaped with an elongated base,
sometimes compressed by mutual pressure, 3-5 mm long, 4-7 mm diam., usually with thickened and
outspread sepals.

Distribution and ecology. Distributed in a small area in south-western Western Australia, near
Ravensthorpe and the Eyre Range, with most of the collections being from the eastern end of
Fitzgerald River National Park (ESPR). Flowering has been recorded in April, May and August.
(Figure 20

Distinctionfi'om related species. Melaleuca pomphostoma is most closely related to M. bracteosa and
differs from that species mainly in its conspicuous stomata, more numerous, longer stamens and
different leaf shape. Although barely visible with a hand lens, the differences between the stomata
of M. pomphostoma and M. bracteosa are striking and consistent. One putative hybrid has been
recorded between M. pomphostoma and M. bracteosa, and the stomata of that specimen are
intermediate between the two putative parents (Figure 5). M. bracteosa has (3)7-11 stamens per
bundle and they are 3.5-5 mm long. Its keeled leaves range from being very narrowly elliptic to very
narrowly obovate.

Conservation status. We recommend that the conservation status of this species should be Poorly
Known (code 2KC- of Briggs and Leigh, 1988) on the basis of herbarium specimens collected from
Fitzgerald River National Park but from populations of unknown size. The species is known from
only 6 collections and has a geographic range of less lhan 100 km. It is represented by populations
within a national park, although adequacy of reservation is unknown.

Etymology. The specific epithet is derived from the Greek pomphos (blister) and stoma (mouth), and
alludes to the distinctive pustular stomata on the lamina.

Additional specimens examined. WESTERN AUSTRALIA: ESPR: 1.9 km Eof jnctn of MoirRd &
TrackjoiningRavenslhorpe/HopetounRd,c.7kmSofRavensthorpe,Fox86/168,6,ii.l986(CANB);
Ravensthorpe, cultivated in Qld by D.M. Gordon, Gray 2 1 29, 196 1 (PERTH); Phillips River Reserve,
Eyre and Whoogarup Range Area, Kessell 870, 20.V.1969 (PERTH); 7 km W of East Mt Barren,
Newbey 4913, 2.xi.l975 (PERTH); 98 (28?) miles from Ravensthorpe, Witlwer 415, 27.viii.1965
(PERTH).

7. Melaleuca pritzelii (Domin) Barlow, comb, et stat. nov. (Figure 4d)

Melaleuca densa var. pritzelii Domin, Mem. Soc. R. Sci. Boheme (Vestn. Krai. Ceske Spolecn. Nauk)
2: 90 (1923). Type: NW Plantaginet, in arenosis, Pritzel 696 (holo: ?PR, not seen; iso: BRI 230687,
E, K, PERTH).

Shrub to 1.2 m tall, with the inllorescence axis and hypanthium tomentose and young shoots
tomentose soon glabrescent, otherwise glabrous. Leaves decussate and sometimes ternate, strongly
spreading; lamina broadly ovate or sometimes obovate, flat but keeled, L2-2.5(4.1) mm long,
1 .2-2. 0(3. 5) mm wide, acute at the apex, sessile; midvein sometimes prominent but venation obscure;
glands punctate. Inflorescence lateral on old wood or term inal (usually male flowers only), a cluster
of 10-15 flowers with axis rarely growing on after anthesis; bracts broadly ovale, L6-2.0 mm long.

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0.8-1.3 mm wide, early deciduous. Hypanthium funnel-shaped, 1. 4-1.9 mm long. Sepals transversely
ovate, scarious, glandular, 0.8- 1 .0 mm long, persistent to mature fruit. Stamens light cream, 3.6-6.5
mm long including a claw (1.2)1. 5-2.5 mm long, 2-4 per bundle. Style 5.5-8.2 mm long; ovules
22-27 per loculus. Fruit cup-shaped, 2.0-2.4 mm long, 3.6-4.0 mm diam., papery in texture; valves
deeply recessed below the aperture.

Distribution and ecology. Distributed in south-western Western Australia from near Ongerup to near
Pootenup (BENC, LUWN, ESPR). Occurs in mallee heath or tall shrubland on shallow, poorly drained
sands over clay. Flowering occurs in August and September. (Figure 2g)

Distinction from related species. Melaleuca pritzelii is probably notclosely related to M. ^fe/!rflR.Br.
even though Domin first described it as a variety of the latter species. It is quite distinctive in its
combination of leaf and innorescence characters and its status as a distinct species is clearly indicated.

Conservation status. We support Briggs & Leigh (1988) who cited this species as M. sp. 4 (Ongerup
and Pootenup) in assigning a conservation status to this species of Endangered (code 2E). An
extensive search in 1986 by B.A.B. in an area near Ongerup revealed only 2 plants. At other sites
visited by F.C.Q. and K.J.C. in 1988 only one or a few plants were found. As each of the populations
is apparently limited to a few plants, the risk of this species disappearing altogether is very high.

Additional specimens e.xamined. WESTERN AUSTRALIA; BENC: 19 km E Katanning, Co/c &
McDonald 201 IH, 15.ii.l986 (CANB); 19 km E of Katanning on Douglas Rd towards Nyabing,
Cowley & Quinn 108, 3.x. 1988 (CANB). LUWN: Gordon River, Tambellup, i./r, i. 1964

(PERTH); near Pootenup, NcM’/jcy 1862, 24.ix. 1965 (PERTH); 11-14 km by road from Kendenup on
Red Gum Pass Rd, Barlow 422S, 24.x. 1986 (CANB); c. 10.3 km S of Stirling Range Rd junction with
Red Gum Pass Rd on Red Gum Pass Rd.,CoH7cy&QMJ>i« 168,9.x. 1988 (CANB). ESPR: ll-13km
E of Ongerup, Barlow 4268, 25.x. 1986 (CANB); 11 km E of Ongerup, Newbey 4781, 5.ix.l975
(PERTH); 23 km E of Ongerup, Newbey 4292, 15.viii.l974 (CANB, PERTH); 7.5 miles NE of
Ongerup, jVcH’foey 342, 19.viii.l962 (PERTH); 10 miles E of Ongerup, Newbey 383, 26.viii.1962
(PERTH).

8. Melaleuca ringens Barlow, sp. nov. (Figure 6a)

Species nova M. vimineae Lindley proxima, sed unguibus staminum brevioribus, ramulis
tomentosis, intlorescentiis semper terminalibus, foliis plerumque latioribus differt.

Holotypus: Western Australia: Leeuwin Region: Point d’Entrecasteaux, 300 m below the lighthouse,
34°50’S ll6°00^E,Cowley&Quinn 183, 10.x. 1988 (CANB 383963). /xotyp/: AD,BRI,G,K,MEL,
PERTH.

Shrub to 3 m high, the rachis and young shoots tomentose, the stems sparsely pubescent, otherwise
glabrous. Leaves spirally arranged, densely crowded and spreading; lamina elliptic, Hat, 4.5-6.6 mm
long, 1. 8-3.0 mm wide, obtuse to almo.st acute at the apex, truncate at the base with a petiole 0.9- 1.1
mm long; midvein distinct with other venation obscure; glands obscure. Inflorescence of 10-60
monads densely arranged into terminal spikes 9-30 mm long; bracts L5-2.5 mm long, L0-L5 mm
wide, early deciduous. Hypanthium cylindrical with an elongated ba.se, c. 1.8 mm long. Sepals 0.8-L0
mm long, entire or narrowly membranous at the margins, persistent to mature fruit. Stamens 4. 8-7.0
mm long including a claw 0.9- 1.3 mm long. 7-11 per bundle with filaments cream. Style 8-11 mm

F.C. Quinn el al.. Rare or threatened Melaleuca

347

long; ovules 20-25 per loculus. Fruits cylindrical to bell-shaped, compressed by mutual pressure,
4-5 mm long, 4-7 mm diam.,with thickened erect or outspread sepals.

Distribution and ecology. Known only from Point d’Entrecasteaux, Western Australia (LUWN).
Occurs in sands over limestone on exposed high ridges or clifftops. Flowering has been recorded in
September and October. (Figure 2h)

Distinction from related species. Melaleuca ringens is probably closely related to M. viminea from
which it differs in its strictly terminal inflorescences, shorter staminal claw, tomentose branchlets and
wider leaves. M. vimineahas leaves 0.6-2.0 mm wideand a staminal claw of more than 1.5 mm long.
Melaleuca ringens also resembles some specimens of M. densa, but can be distinguished by its spiral
leaves and strictly terminal inflorescences.

Conservation status. We recommend that the con.servation status ofthis species should be Vulnerable
(code2V of Briggs and Leigh, 1988). Due to its very limited known distribution (only found at Point
d’Entrecasteaux), M. ringens may be vulnerable to localised threats. The species should, however,
be further surveyed in an effort to locate addition.al populations.

Etymology. The specific epithet is derived from the Latin ringens (gaping), and refers to the very open
fruit aperture.

Additional specimens examined. WESTERN AUSTRALIA: LUWN: Point D’Entrecasteaux, at the
lighthouse, Cowley & Quinn 183A, 10.x. 1988 (CANB); Point D'Entrecasteaux, at the lighthouse,
Eichler 23052, 22.viii.1982 (CANB, PERTH); Point D’Entrecasteaux, along the road to the
lighthouse just below the top of the hill, Eichler 23054, 22.ix.1982 (CANB); locality not known,
voucher for computability tests PERTH Oct. 1984, parent #28, Kenneally s.n. (= Barlow 3835),
9.x. 1984 (CANB); Point D’Entrecasteaux, 7VeH'/;ey 3152, 3 l.i.l968 (PERTH); cultivated from seed
collected from Point d’Entrecasteaux, Newhey 3655, 4.xi.l972 (PERTH).

9. Melaleuca sculponeata Barlow, sp. nov. (Figures 6b, c)

Species nova M. brevifoliae Turez. proxima, sed foliis decussatis basi peltatis, glandulis non
prominentibus differt.

Holotypus: Western Australia: Esperance Region: "Giraween" farm, c. 65 km W of Ravensthorpe,
33°27’S 119°20’E, nnatiuk 800128, 3.X.1980 (PERTH).

Shrub 0.4 m tall, lignotuberous, multistemmed, the young shoots tomentose and the inflorescence
axis with short crisped hairs, otherwise glabrous. Leaves decussate, erect; lamina narrowly elliptic
or oblong, convex abaxially, concave adaxially, 1. 9-3.1 mm long, 0.8- 1.1 mm wide, obtuse or very
shortly mucronate at the apex, obtuse at the ba.se, sessile and attached pcltately near the base; venation
obscure; glands 6-8 visible on the abaxial surface, more or less in two rows, not prominent.
Inflorescence a lateral cluster of c. 10 flowers on old wood. Fruit dcpres.sed-globular, slightly
compressed by mutual pressure, 2-2.5 mm long, 3-4 mm diam., smooth, contracted and entire at the
aperture, 2- or 3-locular with 25-30 seeds per loculus; valves deeply enclosed. Other characters
unknown.

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Nuytsia Vol. 8, No. 3 (1992)

Figure 6. a, Melaleuca rinf-ens. portions of plant.s (flowering piece, Eichler 23054; fruiting piece, Cowley & Quinn 183A).
b,c, M. iculponeala. b, portion of plant {llnaliuk 800128). c, SKM showing abaxial surface of leaf (Hnaliuk 800128).
d, M. viminea subsp. appressa, portions of plant (Newbey 4864). Sc.ale bars = 1 cm

F.C. Quinn et al.. Rare or threatened Melaleuca

349

Distribution and ecology. Known from the type locality only, near Ravensthorpe (ESPR). Recorded
from regenerating mallee heath on light grey sand over clay. (Figure 2i)

Distinction from related species. Although known only from a single non-flowering specimen,
M. sculponeata appears to be distinct from all other species in the genus in its combination of peltate
leaves (Figure 6c) and determinate lateral inflorescences. It is possibly allied to M. brevifolia,
differing in the decussate, peltate leaves which do not have prominent, raised glands.

Conservation status. We recommend that the conservation code for this species, listed by Briggs and
Leigh (1988) as M. sp. 3 (W of Ravensthorpe), should be Endangered (code 2E of Briggs and Leigh,
1988). Melaleuca sculponeata is known from only one locality on a roadside verge surrounded by
wheatfields. It is from an area reasonably well explored botanically and now extensively cleared for
agricultural purposes, and therefore is under serious threat owing to loss of habitat.

Etymology. The specific epithet is derived from the Latin sculponea and alludes to the resemblance
of the detached leaf to the shape of a wooden shoe.

10. Melaleuca viminea subsp. appressa Btu'low, subsp. nov. (Figure 6d)

Subspecies nova M. vimineat Lindley subsp. vimineae proxima, sed foliis aggregatis appressis
apicibus inflexis, staminibus brevioribus, stylis brevioribus differt.

Holotypus: Western Australia: E.sperance Region: 16 km Eof Ongerup, 33°57’S ll^°31'E,Newbey
4864,21.x.l975 (CANB 285751).

Shrub 1. 3-4.5 m high, the young shoots sparsely lanuginosc, the hypanthium and rachis usually
tomentose, otherwise usually glabrous. Leaves spiral (rarely ternate), moderately densely arranged,
appressed; lamina linear to narrowly elliptic or narrowly ovate, flat or slightly compressed, 5-9 mm
long, 0.6-2.0 mm wide; obtuse and slightly inflexed at the apex; attenuate at the base into a petiole
0.8-1. 2 mm long; venation usually obscure, rarely with the midvein evident; glands usually obscure,
rarely pustulate. Inflorescence of 10-50 monads relatively densely arranged into both long basal
spikes and terminal spikes or heads, 8-40(50) mm long, often with the terminal heads predominantly
of male flowers; bracts 1.5-4 mm long, 1-2 mm wide, persistent to anthesis. Hypanthium cylindrical
with an elongated base in hermaphrodite flowers, conical in male flowers, 1-2.5 mm long. Sepals
triangular, 0.5-1 mm long, membranous at themargins, sometimes persistent in mature fruit. Stamens

5- 5.5 mm long including a claw 2-3 mm long, (5)8-1 1(13) per bundle, with filaments cream. Style

6- 7 mm long; ovules 50-80 per loculus. Fruits shortly cylindrical to bell-shaped, 3-4 mm long, 2-4
mm diam., bluntly toothed at the rim.

Distribution and ecology. Known from near Ongerup, MtBurdett and NW of Skeleton Rock(BENC,
ESPR), with a possibly disjunct distribution. Occurs near creeks or wet depressions in clayey soils,
possibly associated with granite. Flowering has been recorded in September and October. (Figure 2j)

Distinction as a subspecies. Melaleuca viminea is an extremely variable species. Much of this
variation has no clear geographic basis, and is presumably the result of an unusual level of
heterozygosity for morphological character states. However, within the total range of variation two
more or less homogeneous entities with individual geographic integrity stand out from the general
residual range of polymorphy, and three subspecies have accordingly been recognised. One of these.

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Nuytsia Vol. 8, No. 3 (1992)

described here as subsp. appressa, is considered to merit coding as a threatened taxon. Melaleuca
viminea subsp. appressa differs from the other subspecies by its oppressed leaves wh ich are incurved
at the apex, and its distinctly shorter floral parts. The collection from Mt Burdctt has slightly shorter,
wider leaves which are more incurved at the apex than the other collections, but is clearly referrable
to this subspecies.

Conservation status . We recommend that the conservation status of this subspecies should be Poorly
Known (code 3KC- of Briggs and Leigh, 1988) on the basis of a herbarium specimen collected from
Mt Burdett Nature Reserve but from a population of unknown size. Subspecies appressa is known
from only 4 collections and has a possibly disjunct distribution over more than 100 km. One
population occurs within a proclaimed Nature Reserve although adequacy of reservation is unknown.

Etymology. The sub.specific epithet is derived from the Latin appressus, and refers to the appressed
leaves of this taxon.

Additional specimens examined. WESTERN AUSTRALIA: BENC: c. 2 km SW of junction of Emu
Fence Road and Brennand Road, Barlow & Fax 4133, 8.x. 1986 (CANB). ESPR: c. 1 km past sign
for Mt Burdett Nature Reserve on road to Mt Burdett & Mt Wittenoom (Kendall’s Road), Cowley &
Quinn 35, 28. ix. 1988 (CANB); 16 km E of Ongerup, Newhey 4610, 25.xi.1974 (PERTH).

Acknowledgements

The conservation coding for all taxa was determined during consultation with John Briggs. We
thank Emile Brunoro, David Coltman and Julie Faulkner for assistance with macrophotography.
Scanning electron microscopy was carried out in the facility of the Division of Entomology, CSIRO.
Advice on the latin diagnoses was provided by Ms Jane Bellemore ( ANU). The Directors of AD, BRI,
CANB, MEL, NSW and PERTH kindly made specimens available for study. Support of the
Australian Biological Resources Study is gratefully acknowledged.

References

Barlow, B.A. (1986). A revised natural regions map for Australia. Branonia 8:387-92.

Barlow, B. A. (1987). Contributions to a revision of AJc/u/fuca (Myrlaceae): 1-3. Brunonia 9:163-77.

Barlow, B.A. and Cowley, K.J. (1988). Contributions to a tevhionof Melaleuca (Myrtaceae): 4-6. Austral. Syst. Bot. 1:95-126.

Briggs, J.D. and Leigh, J.H. (1988). Rare or Threatened Australian Plants. Australiaji National Parks and Wildlife Service
Special Publication No. 14. (Pirie Printers Sales, Fyshwick, ACT.)

Cowley, K.J., Quinn, F.C., Barlow, B.A. and Craven, L.A. (1990). Contributions to a Revision of Melaleuca (Myrlaceae): 7-10.
Austral. Syst. Bot. 3:165-202.

Nuytsia 8(3): 351-360 (1992)

351

Three new species of Euphorbia L. subgenus Chamaesyce Rafinesque
(Euphorbiaceae) from central and northern Australia

B.G. Thomson

Northern Territory Herbarium, Conservation Commission of the Northern Territory,
PO Box 1046, Alice Springs, Northern Territory 0871

Abstract

Thomson, B.G. Three new species of Euphorbia L. subgenus Chamaesyce Rafinesque
(Euphorbiaceae) from central and northern Australia. Nuytsia 8(3): 351-360 (1992). Euphorbia
centralis from central Australia, E. maconochieana from the Victoria River and E. kimberleyensis
from the northern Kimberley region are described, illustrated and their affinities with related species
discussed. E. vaccaria Baillon is lectotypified.

Introduction

Euphorbia L. is a worldwide genus of primarily sub-tropical and temperate distribution. It
contains approximately 2000 species (Willis 1966) and is represented in Australia by an estimated
45 species.

Bentham (1873) included 18 species in his revision of the genus. These he treated in 2 sections;
Eremophila Boiss. with 1 species and Anisophyllum (the correct author being Roeper rather than
Roxborough as stated) with 17. Bentham later transferred the single species that he had placed in
section Eremophila to section Eremophyton Boiss. (Bentham & Hooker 1880). This section has since
been given sub-generic status by Wheeler (1943) and now includes five Australian species (Hassall
1977).

Section Anisophyllum had also been given sub-generic status (Gaucher 1898), however, the name
at this level of classification was preoccupied. The name correctly applied at sub-generic rank to this
taxon is Chamaesyce Ral'ine.sque 1817 (Croizat 1938, Wheeler 1943).

In 1821 S.F. Gray elevated Chamaesyce to generic status in 'A Natural Arrangement of British
Plants'. Since this time many researchers have supported its retention at this level (Croizat 1938,
Webster 1975) and many have rejected it (Wheeler 1941, Radcliffe-Smith 1975). Ha.s.sall (1976)
appraised the situation as it related to Australian species and supported the retention of Chamaesyce
as a distinct genus. This change was not. however, widely accepted as evidenced by the treatments
of Euphorbiaceae in many subsequent state and regional floras.

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Nuytsia Vol. 8, No. 3 (1992)

Figure 1. Distribution of Euphorbia centralis (□), E. maconochieana (A), and E. kimberleyensis (O).

Most authors agree that the taxon Chamaesyce is a natural one. The difficulties arise, however,
in concisely defining the group. As Wheeler (1941) points out, it is with the greatest difficulty that
Chamaesyce can be defined so as to exclude all members of other taxa and, at the same time, include
all members of Chamaesyce. Although this statement was applied by Wheeler to the characters
detailed by Croizat (1938) and earlier authors, it can be just as readily applied to the more modern
arguments citing carbon fixation pathways (Webster et al. 1975) and cytological evidence (Hassall
1976).

A most important feature that .serves to define Chamaesyce is the unique growth habit whereby
the main stem axis aborts its apical meristem just above the first pair of true leaves. All subsequent
growth arises sympoidally from the region of the cotylendonary nodes (Croizat 1938). This character
is clearly seen in many of the prostrate .species. Many, however produce only a single, erect stem and,
in these cases, the interruption of the primary axis becomes particularly difficult if not impossible to
observe in the mature plant. Croizat (1938) illustrates this feature. Thus, while the character is
diagnostic of Chamaesyce, it is not particularly helpful in terms of practical taxonomy.

In this study and in future work, including the preparation of the treatment for the Flora of
Australia, I follow Wheeler (1941) in accepting Chamaesyce at a sub-generic level.

The species dealt with below, are members of sub-genus Chamaesyce Rafinesque and are the first
of a number of new taxa that have been revealed as aresult of studies undertaken to provide an account
for the Flora of Australia.

B.G. Thomson, Three new species of Euphorbia

353

Descriptions and lectotypification

1 . Euphorbia centralis B.G. Thomson, sp. nov. (Figure 2)

E. australi Boissier affinis, ab qua foliis late ovatis vel orbiculatus, serratis, appendicis glandis
distincte laciniata, suffruticoso habitu distingucnda.

Typus: 3 km SW of Alice Springs, Northern Territory, 23° 35' S, 133° 50' E, 18 January 1990,
B.G. Thomson 3408 (holo: DNA; iso: AD, BRl, CANB)

Erect or occasionally decumbent annual orperenniaUu/j-s/irM^j to 30 cm tall, multi-stemmcd from
the base. Sfcmi puberulous to pilose with hairs 0.2-1 mm long. triangular, 0.2-0.4 mm long,

margins lacerate and with ciliate hairs. Leaf lamina broad-ovate to orbicular, 3-5.5 x 1.5-5 mm,
surfaces puberulous to pilose, margins serrate in the upper half, base oblique, apex obtuse. Petioles
0.5- 1.3 mm long. Inflorescence in form of cyathia, solitary, axillary, throughout the plant or in upper
parts only. Cyat/tmturbinate, 0.8-1. 1 xO.8-1.1 mm, puberulous to hispid. Cyathial lobes ix'anguXas,
margins ciliate with stiff, white hairs. Glands oblong, concave, 0.2-0.4 x 0. 1-0.3 mm. Gland
appendages 0.6-1. 3 x 0.3-0.5 mm, light green, cream, pink or red, margins strongly laciniate.
Androphores level with the cyathial rim or exserted. Gynophore glabrous or pubescent, 1.3-2 mm
long. Capsule 1.5-2 x 1. 8-2.2 mm, muricate to hispid. Styles 0.3-0.4 mm long, glabrous. Seeds
narrowly ovoid, 0.9- 1.5 x 0.5-0.6 mm, telragonous to tetraquetrous, rugose, brown (or mottled creamy
brown with mucilage coat intact).

Selected Specimens. NORTHERN TERRITORY: James Range, 3 km N of Hugh River,
A.C. 24586(CANB,DNA);Hatches Creek Mine,20°55'S, 135° 12'E,D.F. McKey302

(DNA); 7 miles NNEof Willowra Homestead, 21° 15' S 132° 3TE,P.K. Latz 1245 (DNA); 'Victory
Downs' Station, 25° 59' S 132° 10' E, P.K. Latz 5099 (BRI, DNA); 22 miles SSW of Georgina Downs
Station, 21° 22' S, 137° 24' E, R.A. Perry 3471 (CANB, DNA); 5 km WNW of Supplejack Bore,
'Elkedra' Station, 21° 10' S, 135° 30' E (DNA); 6 km E of John Hayes Rockhole, Trephina National
Park, 23° 3 1’ S, 134° 22' E, B.G. Thomson 1496 (DNA); 1 km S of Muranji Rockhole, Mt Winter,
23° 49' S, 130° 53' E, B.G. Thomson 1552 (AD, DNA, NT); eastern end of Harts Range, 23° 03' S,
134° 39'E,B.G. Thomson 2413 (DNA, MEL, NT); MtZeil, 23° 24' S, 132° 23'E,B.C. Thomson2131
(DNA).

SOUTH AUSTRALIA: Musgrave Range, 1 mile E of Mt Woodroffe, R. Hill 8c T.RJI. Lothian 712
(AD, DNA); Kalka, Tomkinson Ranges, 26° 07' S. 129° 10' E, A. Kalotas 831 (DNA).
QUEENSLAND: 3 miles NW of 'Buckingham Downs' Station, M. Lazarides 4345, (CANB, DNA).

Distribution. E. centralis occurs from the Tennant Creek region (NT) to the Musgrave Ranges (SA)
and from Mt Isa (Qld), west to the Docker River region (NT). (Figure 1)

Habitat. Found commonly on limestone, qutirtzite and sandstone ranges and low hills.

Affinities. This species is closely allied to£. australis Boissier but differs in having strongly laciniate
gland appendages and more orbicular, deeply serrate leaves. Mature plants exhibit a rounded, shrub-
like habit, the lower leaves withering but often persisting on the stems.

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Nuytsia Vol. 8, No. 3 (1992)

Conservation status. E. centralis is common and well represented in collections from central
Australia and can not be considered as rare or endangered.

Etymology. The specific epithet relates to the species’ central Australian distribution.

2. Euphorbia vaccaria Boiss., Adansonia 6: 286 (\?,66).Type citation: 'Exs. F. Mueller, Victoria
River (herb.!). - Id. "Rocky high hills Hierson island, Nickol hay." Lectotype (here chosen): Nickol
[Nichol] Bay.E. Walcott (MEL 1551017).

Typification. Baillon (1866) described Euphorbia vaccaria and cited material collected by Mueller
from the Victoria River and by an unknown collector from Rocky High Hills, Hcirson (Hearson)
Island, Nickol (Nichol) Bay. In the course of this study, three specimens collected by Mueller from
the Victoria River and named by him as E. vaccaria were examined. These specimens were located
in MEL (MEL 1551016, 1551015, 1551018). Also located in MEL was the Heirson (Hearson) Island
specimen bearing an original, hand-written label including the details cited by Baillon and the
collector’s initials. A comparison of handwriting confirmed that this specimen was one collected by
Pemberton Walcott who visited Nickol (Nichol) Bay with the F.T. Gregory Expedition from May to
July 1861. According to Baillon, who worked in Paris, these specimens were received on loan from
Mueller. A search in Paris failed to locate any further material.

On clo.se inspection of these syntypcs it was apparent that they represented two .separate taxa,
the material collected by Mueller being quite different from Walcott’s specimen. Baillon’s
protologue describes E. vaccaria as villous and, in this respect his description matches Walcott’s
specimen but is at odds with the Mueller collections. Mueller’s specimens have a short, puberulous
indumentum which is only visible under magnification. In no way could they be considered as villous
even allowing for the broadest interpretation of the term.

Although the Walcott specimen is cited in the protologue without the collector’s name, Baillon
has included sufficient details to reliably establish it as a syntype. The same, however, can not be said
for the Mueller material. Although the three specimens examined were collected from the Victoria
River and bear label data that agree with the protologue, it is not clear which of these specimens were
seen by Baillon. Baillon apparently did not annotate any of the material that he examined on loan
from Mueller.

In consideration of the above points, I have chosen the Walcott specimen (MEL 155 1017) as the
lectotype for E. vaccaria. The Mueller material represents a new species described below as
E. maconochieana.

3. Euphorbia maconochieana B.G. Thomson sp. nov. (Figure 3)

E. schultzio Benth. affinis, ab qua appendicis glandis petaloidea grandi, aba, integra, capsula
parviore, marginibus folii integris distinguenda.

Typus: Cahills Crossing; Victoria River crossing on the Top Springs to 'Victoria River Downs' road,
N.T., 16° 20' S, 131° 07' E, B.G. Thomson 3486 (holo: DNA; iso: AD, BRI, PERTH).

B.G. Thomson, Three new species of Euphorbia

355

Figure 2. Euphorbia centralis A - habit, B - leaf, C - cyathia and capsule, D - seed. From B.G. Thomson 3408.

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Nuytsia Vol. 8, No. 3 (1992)

Prostrate, annual herb. Stems puberulous with curved or occasionally straight hairs to 0.2 mm
long. Stipules subulate, 0.2-0.5 mm long, often divided into several filiform segments, margins
ciliate. Leaf lamina elliptic to obovate, 6-15 x 2-7 mm, upper surface glabrous to puberulous, lower
surface puberulous, margins entire, base strongly oblique, apex obtuse. Petioles 0.5-1 mm long.
Inflorescence in form of cyathia, solitary, axillary, throughout the plant or occasionally in upper parts
only. Cyathia turbinate, 1.3-1. 9 x 1. 2-1.7 mm, densely muricate, puberulous. Cyathial lobes
triangular, margins ciliate with stiff, white hairs. Glands oblong, flat to concave, 0.3-0. 5 x 0. 3-0.4
mm, pink to dark red. Gland appendages 0. 1-0.5 x 0.3-0.9 mm, white, prominent on living material,
margins entire. Androphores exserted above the rim of the cyathia. Gynophore scabrous 1 .0-1.6 mm.
Capsule 1.7-2.1X 1. 9-2.2 mm, densely muricate and often puberulous with short, stout hairs. Styles
0. 3-0.5 mm long, smooth or minutely scabrous. Seeds ovoid, 1. 1-1.4 x 0.7-0. 8 mm, tetragonous,
lightly rugose, light brown (or creamy brown with mucilage coat intact).

Specimens examined. NORTHERN TERRITORY: 51 miles E of 'Victoria River Downs', 16° 23' S,
131°29'E,C. Chippendale 6095 (DNA); lOmilesNNEof'Wavehill' Station, 17° 17' S, 131° 10' E,
M. Lazarides 6278 (DNA); Victoria River Crossing on Top Springs/Timber Creek Road, 16° 20' S,
131° 06' E, M.O. Parker 1009 (DNA); 4 miles S of 'Willeroo' outstation, 15° 19' S, 131° 35' E,
R.A. Perry &M.Lazarides2025 (DNA); 46 miles SW of'Birrimbah'Out.station, 16°30'S, 131° 52' E,
R.A. Perry & M. Lazarides 2078 (DNA); 4 km S of No. 10 Bore, Rosewood Station, 16° 16' S,
129° 27' E, B.W. Strong 989 (DNA); Victoria River, F. Mueller s.n. (MEL 1551015, 1551016,
1551018).

WESTERN AUSTRALIA; near Oscar Range road to Leopold N, A.J. Ewart s.n. (PERTH); Drain 7,
Packsaddle Creek, Kimberley, 15° 49' S, 128° 41' E, K.F. Kenneally 1941 (PERTH); Kunnunurra,
E.M. Scrymgeour 1673 (PERTH); Smoke Creek, SW of Lake Argyle, 16°45' S, 128°30' E,
A.S. Weston 12185 (PERTH).

Distribution. This species is known from the Victoria River region in the Northern Territory and
adjacent Lake Argyle in Western Australia. (Figure 1)

Habitat. Normally found in heavily textured, alluvial soils.

Affinities. E. maconochieana is closely related to E. schultzii Benth. and shares an almost identical
indumentum characterised by short (0.2 mm or less), thick, incurved hairs. It is separated from
E. schultzii by the following key.

1. Gland appendages entire, white; capsules 1.7-2. 1 mm long,

1.9-2.2 mm wide; leaf margins entire; prostrate annual E. maconochieana

1. Gland appendages denticulate, pink to red; capsules 2.3-2.8 mm
long, 3.0-3.5 mm wide; leaf margins serrate; erect, decumbent

or prostrate annual or occasionally perennial E. schultzii

Conservation status. This species, although rather poorly collected, is not considered as rare or
endangered in any part of its range.

Etymology. The species is named in honour of the late Mr J.R. Maconochie, a close friend and former
colleague at the Northern Territory Herbarium.

B.G. Thomson, Three new species of Euphorbia

357

Figure 3. Euphorbia maconochieana A - habit, B - leaf, C - cyathia and capsule, D - seed. From B.G. Thomson 3486.

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Nuytsia Vol. 8, No. 3 (1992)

4. Euphorbia kimberleyensis B.G. Thomson sp. nov. (Figure 4)

E. schizolepi F. Muell. ex Boiss. affinis, ab qua caulibus et foliis glabris vel glabratis, capsula et
semine parvioribus et stylo glabro distinguenda.

Typus: Palm Woodland, Mitchell Plrteau, West Kimberley, 14° 50' S, 125° 50’ E, 15 June 1976,
K.F. Kenneally A92\ (holo; PERTH; iso: CANB)

Annual herb, prostrate or with ascending or decumbent stems, to 20 cm tall. Stems glabrous or
puberulous. Stipules subulate, 0.2-0.5mm long, margins lacerate. Leaf lamina elliptic to obovate,
6-16 X 5-12 mm, surfaces glabrous, margins entire, base oblique, apex apiculale, often carinate.
Petioles 1 -2 mm long. Inflorescence in form of cyathia, solitary, axillary, throughout the plant or in
clusters of 2-4 on lateral brachlets due to shortening of internodes. Cyat/j/n turbinate, 1. 5-2.0 x
1. 2-1.5 mm, glabrous. Cyathial lobes narrow-triangular, margins conspicuously fimbriate. Glands
orbiculartooblong, flat or concave, 0.6- 1.1 xO.4-0.7 mm, red. Gland appendages 0.5-2.0 xO.S-2.0
mm, white or pink, margins denticulate. Gynophore 3.0-4 .5 mm long, glabrous. Capsule 3. 2-3.7 x
3.0-3.5 mm, smooth or tuberculate, glabrous or rarely with sparse pubescence. Seeds ovoid,
tetragonous, 1.9-2.3 x 1.2- 1.4 mm, rugose with flat topped ridges, cream to brown.

Specimens examined. WESTERN AUSTRALIA: Near Lone Dingo VT, 9 km S W of WarrenderHill,
14° 30' S, 1 25° 45' E, J. J. Alford 55 1 (PERTH); Kalumburu Road, 108.9 km by road N of Gibb River
and Ellenbrae Road, 15° 23' S, 126° 12’ E, Aplin et al. 721 (PERTH); base of Mt Behn, West
Kimberley, W.V. Fitzgerald 685 (PERTH); Inglis Gap, King Leopold Range, W.V. Fitzgerald 755
(PERTH); 1.6 km along Surveyors Fall Track, N of Mitchell Plateau Mining Camp, N Kimberley,
14°40'S, 125°47'E,A„S. George 14478 (PERTH); Port Warrender, Mitchell Plateau, W Kimberley,
14° 34’ S, 125° 16’ E, K.F. Kenneally 5254 (PERTH); area of Carson Volcanics towards Port
Warrender,offthelaleriteplateau.WKimberley, 14°34’S, 125°50’E,K.F. Kenneally 6704 (PERTH);
Crocodile Creek, 5 km E of west end of Koolan Island, W Kimberley, 16° 10’ S, 123°41’ E,
K.F. Kenneally 9719 (PERTH).

Distribution. From Derby, north-east to the Drysdale River, Western Australia. (Figure 1)

Habitat. Found commonly in association with open woodland communities on lateritic red soil or
sandstone.

Affinities. This species is closely related to E. schizolepis F. Muell ex Boiss. and, in particular to E.
schizolepis var. glabra Benth. Bentham ba.ses his variety on material collected by Mueller in the Gulf
of Carpentaria and describes it as perfectly glabrous with the gland appendages much less lobed. A
search in K failed to locate any material of Bentham's variety. I have, however, examined a specimen
in MEL which matches the protologue and is probably part of the type collection. This material is
clearly not referable to E. kimberleyensis, differing in its lai ger cyalhium and glaucous upper stems
which are commonly seen on the more glabrescent forms of E. schizolepis. E. kimberleyensis is also
restricted to the far north of Western Australia and does not occur in the 'Gulf of Carpentaria’ which
is the rather generalised collection locality given by Bentham for his variety glabra.

B.G. Thomson, Three new species of Euphorbia

359

Figure 4. Euphorbia kimberleyensh A - habit, B - leaf, C - cyathia and capsule, D - seed. From K.F. Kenneally 4921

360

Nuytsia Vol. 8, No. 3 (1992)

E. kimberleyensis may be distinguished from E. schizolepis by the following key.

1. Herbaceous, prostrate or weakly ascending annual, glabrous or
occasionally with puberulous stems; capsules 3.2-3.7 mm long
and 3.0-3.5 mm wide, smooth or tuberculate, glabrous or rarely

with sparse pubescence; styles glabrous; seeds 1. 9-2.3 mm long E. kimberleyensis

1. Erect annual or perennial, pilose to villous, rarely glabrescent;
capsules 4.0-6.0 mm long and 4.5-6.5 mm wide, surface obscured
by densely tomentose indumentum; styles distinctly pubescent;

seeds 2.7-3.5 mm long E. schizolepis

Conservation status. This species appears to be common throughout its limited range.
Etymology. The specific epithet relates to the species distribution within the Kimberley region.

Acknowledgements

I would like to thank G. Leach and C. Dunlop for their critical review of the manuscript and
assistance with the latin diagno.ses. K.L. Wilson, as Australian Botanical Liaison Officer, searched
collections held at K and P and also provided advice regarding overseas collections. I also wish to
thank the Curator of the W. A. Herbarium (PERTH) for access to the collection. The drawings were
prepared by Milton Andrews.

These species were examined in the field during a collecting trip to N.W. Western Australia which
was made possible through Australian Biological Resources Study funding.

References

Bentham, G. (1873). "Flora Au.straliensis'' Vol.6. (Lovell Reeve, London.)

Bentham, G. and Hooker, J.D. (1880). "Genera Plantarum" Vol. 3. (Lovell Reeve, London.)

Croizat, L. (1938). Euphorbiaceae in Degener, O. "Flora Hawaiiensis" (Degener, U.S.A.)

Gaucher, L. (1898). Etude Anatomique du Genre Euphorbia L.

Hassall, D.C. (1976). Numerical and Cytolaxonomic Evidence for Generic Delimitation in Australian Euphorbieae. Austral. J.
Bot. 24:633-640.

Hassall, D.C. (1977). 'Hie Genus Euphorbia in Australia. Austral. J. Bot. 25:429-453.

Rafinesque, C.S. (1817). Second Decade of Undescribed American Plants. Amer. Monthly Mag. 2: 1 1 9.

Webster, G.L. (1975). Conspectus of a New Classification of the Euphorbiaceae. Taxon 24:593-601.

Webster, G.L., Brown, W.V. and Smith, B.N. (1975). Systematics of Photosynthetic Carbon Fixation Pathways in Euphorbia.
Taxon 24:27-33.

Wheeler, L.C. (1941). Euphorbia subgenus chaniaesyce in Canada and the United States, exclusive of Southern Florida. Rhodora
43:97-286.

Wheeler, L.C. (1943). The Genera of the Living Euphorbieae. Am. Midi. Nat. 30:456-503.

Willis, J.C. (1966). "A Dictionary of the Flowering Plants and Ferns" Edn 7. (Cambridge at tire University Press, London.)

Nuytsia 8(3): 361-377 (1992)

361

The Lawrencella complex (Asteraceae: Gnaphalieae; Angianthinae)

of Australia

Paul G. Wilson

Western Australian Herbarium, Department of Conservation and Land Management,
PO Box 104, Como, Western Australia 6152

Abstract

Wilson. Paul G. ThsLawrencellacomplcx (Asteraceae:Gnaplialieae:Angianthinae) of Australia.
Nuytsia8(3); 361-377 (1992). The Lflwm!ce//a complex includes the monotypicgenera5e///c?a and
Schoenia, several species included by Bentham in Helichrysum sect. Lawrencella, and one species
previously placed in both Helichrysum and Podolepis. It is considered to be a natural group clearly
distinct from other members of the Angianthinae. Three genera are here recognised: Bellida,
Lawrencella, and Schoenia. One subspecies is described as new; four new species combinations are
made.

Introduction

For many years botanists have recognised that the Australian species placed in Helichrysum are
not congeneric with the South African and European taxa of that genus (e.g. Merxmullercfa/. 1977;
Hilliard & Burtt, 1981), they have further recognised that the genus as circumscribed in Au.stralia is
an artificial assemblage of species and that the closest affinities of these species is frequently to species
in other genera rather than to those within the genus (see Haegi 1986). Recently Anderberg (1991)
has reviewed the tribe Gnaphalieae. He has removed the Australian species that had previously been
included in Helichrysum and placed them in Chrysocephalum Walp., Bracteantha A. Anderb. &
L. Haegi, Ozothamnus R.Br., Schoenia Steetz, and the ’Lawrencella' complex which, he noted,
required further study. The genus Bellida he treated as a distinct monotypic taxon possibly related
to the Waitzia group.

A collaborative comprehensive classification of the Helichrysum-Helipterum complex in Australia
is in preparation. Prior to this treatment a number of segregate genera are being described or
recognised in prep:iration for a treatment of the family Asteraceae in volume 38 of the Flora of
Australia. In view of this forthcoming publication, and in order to save lime and avoid repetition, for
some of the taxa investigated only brief descriptive details are provided.

For a long time some or all of the species that are here included in the Lawrencella complex have
been considered to be related. The complex was recognised by Bentham (1867) as a section of
Helichrysum, with the exclusion of Schoenia since its single .species he considered to be generically

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Nuytsia Vol. 8, No. 3 (1992)

distinct due to its flattened achenes; he noted, however, that this species was otherwise similar to
species in Helichrysum sect. Lawrencella. Mueller (1889) recognised the relationship between the
one species of Schoenia and the species included by Bentham in sect. Lawrencella for he li.sted them
together as species of Helichrysum, as did Black (1929, 1957). Earlier Black (1915) had suggested
that if Schoenia is to be recognised as a genus distinct from Helichrysum then H. ayersii F. Muell.
must be included in it. Haegi (1986) placed Schoenia cassiniana (as Helichrysum cassinianum),
Helichrysum ayersii, H. davenportii F. Muell., and H. semifertile F. Muell. in sect. Lawrencellabut
noted that this section should have generic rank since the included species were very distinct from
other members of Helichiysum. In a recent paper Anderberg (1991) indicated the close affinity
between Lawrencella and Schoenia and recognised that further work was required before the status
of related taxa could be clarified. In the 'Lawrencella' complex he placed Helichrysum ayersii,
H. davenportii, H. lindleyi Eichler (= Lawrencella rosea Findley), H. filifolium (with its close
relatives here referred to as the 'Xanthochrysum' group), H. spiceri F. Muell. and H. obtusifolium
F. Muell. & Sonder. I concur with his suggestions except that I exclude the last two species which
I consider to generically distinct.

Tribal classification

The species included in the Lawrencella complex have traditionally been placed in the tribe
Inuleae. Work on the tribal classification by Anderberg (1989) has indicated that this tribe should
be divided into three, the Inuleae, the Gnaphalieae, and the Plucheae. The Gnaphalieae, as defined
by Anderberg, comprises the taxa that were included in the two subtribes Gnaphaliinae and
AthrixiinaebyMerxmullereta/.(1977). In arecent comprehensive treatmentofthetribe Gnaphalieae
Anderberg (1991) has recognised a number of subtribes including the subtribe Angianthinae Benth.
in which Lawrencella and its relatives are placed. Anderberg expressed uncertainty as to the
systematic position of Bellida but, as indicated above, I consider that it should be included in the
Angianthinae and that it is closely related to Lawrencella.

Taxonomy

The Lawrencella complex

Many of the characters that have been exam ined and used to delimit the genera have been discussed
elsewhere (Wilson 1989). There are many morphological and anatomical characters associated with
the plants that are of value in determining relationships, however, not all of these characters can be
so described that they are readily comprehended. Thus the texture of the corolla, the shape of its
constituent cells and the thickening of the cell walls are characters that are of value in suggesting
affinity, though sometimes of limited use in formal descriptions because of their cryptic nature. I have
expanded below on some of the characters that I consider to be important when assessing affinities.

A number of characters that are found in the achene are useful in serving either to distinguish the
Lawrencella complex from other taxa that have been included in Helichrysum, or to distinguish the
putative genera or infrageneric taxa within the group. Those characters that are used in this paper are
as follows:

Paul G. Wilson, Lawrencella complex

363

Testa ; vascular sl/ and (Figures 2-7) . The testa always has a solitar>' vascular strand that may terminate
before the apex of the seed proper, or may term inatc at the apex of the seed (particularly in those cases
where the apex is in the form of a short sterile apiculum), or may continue around the seed. This
character appears to be constant in many sections or genera of the Australian Angianthinae as recently
recognised. Thus in the Helipterum albicans group (Leucochrysum Paul G, Wilson, 1992b) the
vascular strand consistently terminates in the short sterile apex of the seed. In Bracteantha it passes
over the apex to the other side; while in WaiVzia Wendl., Chrysocephalum,Leptorhynchus Less., and
in the Helichrysum datum group it ceases before reaching the apex. With regard to those taxa here
referred to the Lawrencella complex it terminates in a sterile apex in Bellida, Helichrysum
davenportii, and Lawrencella rosea, while it passes over the apex to the other side in H. ayersii,
Schoenia cassiniana, and in species of the 'Xanthochrysum' group.

Testa: epidermis. Examination of the cells of the outer layer of the testa in numerous species
previously included in Helipterum and Helichrysum has shown that they in general have the same
shape in species that are now recognised as being closely related but may vary in shape between
species-groups and between genera. Thus in Chrysocephalums.str. the cells are linear, in 5; actea«t/ia
they are almost square with straight margins, while in Waitzia and Leucochrysum they are short-
oblong with corrugate margins. In the Lawrencella complex they are ± equilateral in all taxa except
for H. davenportii and Bellida where they are short-oblong (Figures 2-7).

Corolla. The shape and texture of the tube and lobes, the indumentum, the shape and thickening of
the cells of the inner epidermis, the papillosity of the inside of the lobes and throat, and the extent of
the vasculature, are all characters that must be assessed for their generic significance. In the
Lawrencella complex the corolla is regular with cylindrical tube and campanulate limb, the cells of
the inner epidermis of the lobes are ± equilateral and of the throat ± straight, the vascular strands reach
to the apex of the lobes, while the hairs on corolla tube are biseriate and gland-tipped. InBracteantha,
Chrysocephalum, Ozothamnus, and in the Helichrysum obtusifolium group the corolla is narrow-
cylindrical and almost glabrous with vascular strands of the last three taxa not passing into the lobes.

Anthers (Figure 1). Of significance in the anther are the shape and length of the collar; the length,
texture and branching of the tails; the shape and texture of the anther appendage, and the arrangement,
shape, and thickening of its constituent cells. The shape of the anther appendage has frequently been
noted but the constituent cells offer further characters that are of value when assessing affinities.

In the Lawrencella complex the appendage is acutely cordate or broad-ovate, the proximal cells
± equilateral with thick walls, the distal cells oblong with thinner walls, and the marginal cells small
and equilateral, the collar is narrow-oblong and the anther tails very weak and filamentous and
extending beyond collar. This situation issimilar to that found in /?/(o<7anr/i<;Lindleys./.andcontrasts
with the other genera that have been traditionally included in Helichrysum', in these other genera (e.g.
Bracteantha, Chrysocephalum, and Ozothamnus) the proximal cells of the appendage are the same
as the distal and the tails are firm with the cell walls somewhat thickened.

Style (Figure 1). Of significance are the length and thickness of the arms, the shape and degree of
papillosity of the apex, and the thickness andextentofthe vascular .strand. In the Lawrence/Za complex
the arms are long, the apex deltoid with a stout vascular strand extending to its tip. This contrasts with
the other taxa of Australian 'Helichrysum' in which the apex is rounded, truncate, or acuminate, and
of the same width as, or only slightly wider than, the style arm.

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Nuytsia Vol. 8, No. 3 (1992)

Pappus (Figures 2-7). In the Lawrencella complex the bristles are linear, firm, prominently
barbellate, without spreading basal cilia, they are persistent or shed entire. In Bellida, H. lindleyi and
H. davenportii the pappus lengthens during anthesis. In other members of 'Helichrysiim ' the bristles
are filiform or, if broadened at base, eventually break just above the base to leave a persistent corona
on the achene.

Achene apex (Figures 2-7). The apex of the achene forms a shallow rim around the base of the pappus
in Helichtysum cassinianum and in H. ayersii, in the other species of the Lawrencella complex the
achene apex narrows and passes smoothly into the pappus base.

Achenial hairs (Figures 2-7). Normal duplex hairs, when present, are thick-walled; two-celled
papillose hairs are absent. The pericarp is velutinous with minute acicular unicellular papillae in
Bellida, H. lindleyi. and H. davenportii, in at least some of the achenes. In H. davenportii acicular
multicellular hairs are sometimes also present. The achenes of H. cassinianum, H. ayersii, and the
'Xanthochrysum' group lack these papillae or hairs but bear scattered multicellular gland-tipped hairs
in addition to normal duplex hairs.

Vasculature of the achene and seed (Figures 2-7). The position of the two vascular strands in the
pericarp in relation to the solitary vascular strand in the testa and to the position of the cotyledons of
the embryo was suggested by Short et al. (1989) to be of generic significance.

Figure 1. Anther apicula (A-D) and style apices (E-H) of Lawrencella davenportii (A & E), Bellida graminea (B & F),
Ozothamnus lepidophyllus (C & G), and Chrysocephalum scmicalvurti (D & H).

Paul G. Wilson, Lawrencella complex

365

The strands of the pericarp and testa are laterally placed (with regard to cotyledons) in Bellida,
Helichrysum lindleyi, and II. davenportii, while in H. cassinianum,H . oyera'i , and in the 'Xanthochrysum'
group they are medially placed.

Carpopodium. The carpopodium is small or insignificant in all species of the Lawrencella complex;
however, in Bellida, H. lindleyi, and H. davenportii the achene has a prominent hollow base while
in H. cassinianim, H. ayersii, and in the Xanthochrysum' group the base is not excavated.

Receptacle. The receptacle is smooth and glabrous in all the taxa of Iho Lawrencella complex except
for the Xanthochrysum' group in which it is glandular papillose.

A summary of some of these characters for the taxa of the Lawrencella complex is listed in Tablel .

B L D C A X

Pericarp v.s.

lateral

+

+

+

medial

+

+

+

Testa v.s.

to apex

+

+

+

circum.

+

+

+

Achene base

minute

+

+

+

excav.

+

+

Achene apex

rimmed

+

+

not rimmed

+

+

+

+

Pericarp

thick

+

+

+

thin

+

+

+

velutinous

+

+

+

not vel.

+

+

+

Gland, hairs

present

+

+

+

absent

+

+

+

Pappus

elongating

+

+

+

not elong.

+

+

+

Receptacle

papillose

+

smooth

+

+

+

+

+

Table 1. Comparison of some characters of the achene and capitulum in Bellida (B), Helichrysum
lindleyi (L), H. davenportii (D), H. cassinianum (C), H. ayersii (A), and the 'Xanthochrysum'
group (X).

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Chemical data

Few Australian species of Heliclirysum have been chemically investigated and the taxonomic
significance of the results of those that have is not very informative. However, in a paper by Jakupovic
et al. (1989) the results of an analysis of Helichrysum davenportii, H. lindleyi, and Bellida graminea
were presented along with analyses of H. ambiguum Turcz. (= Chrysocephalum), H. bilobum Wakef.
(= Ozothamnus), and H. leucopsideum DC. From this investigation some tentative conclusions were
drawn.

In Helichrysum davenportii was found isokaurenic acid and a thiophene derivative numbered 37.
In H. lindleyi the thiophenes numbered 37 and 38 and some triterpenes were found, while in Bellida
graminea were found thiophene acetylenes and the desoxy derivative of isokaurenic acid numbered
37a and the thiophene numbered 38. These compounds were not found in the three other species of
Australian 'Helichrysum' .

The chemical analyses therefore suggests that the three 'Lawrencella' species are more closely
related to each other than they are to the remaining 'Helichrysum' species This lends support to the
opinion, based on morphological considerations, that the 'Lawrencella' complex is a natural
assemblage of taxa.

Mycorrhizal data

It has been demonstrated by Warcup (1990) that Australian species in the tribe Inuleae can be
almost equally divided into tho.se that form only vesicular-arbuscular mycorrhiza ( VAM species) and
those that form both ectomycorrhiza and vesicular-arbuscular mycorrhiza (Ecto species). The
Australian taxa placed in 'Helichrysum' can also be separated into Ecto and VAM species. The
presence of both VAM and Ecto species in the same gen us is unusual since normally genera have been
found to contain species that are either all ectomycorrhizal or all non-ectomycorrhizal. When the
'Helichrysum' species are segregated into those genera recognised by Anderberg (1991) and by myself
the situation changes, for then each of the segregate genera contains, as far as is known, only
ectomycorrhizal or only non-ectomycorrhizal species.

Warcup (1990) examined the mycorrhiza of the following species in ff\& Lawrencella complex;
Helichrysum ayersii, H. cassinianum, H. lindleyi, H. davenportii, H. subulifolium, and Bellida
graminea-, he found them all to be non-ectomycorrhizal. This is also the situation in Rhodanthe s.l.
(Wilson 1992a) and Helichiysum subg. Ozothamnus (Ozothamnus R.Br.); it contrasts with the
situation in the Helichrysum bracteatum group (Bracteantha), the H. apiculatum group
(Chrysocephalum), and in the//, leucopsideum group all of which are ectomycorrhizal.

Generic circumscription

On the basis of the characters that are found in the achene (see above) a close relationship is
suggested between H. cassinianum, H. ayersii, and the species of the 'Xanthochrysum' group on the
one hand, and between Bellida, H. lindleyi and H. davenportii on the other. This correlates with the
nature of the pappus which in H. lindleyi, //. davenportii, and Bellida elongates during anthesis to
extend beyond the corolla while in the other genera it remains more or less equal to the corolla.

Paul G. Wilson, Lawrencella complex

367

The number of genera that should be recognised in i\\e,LawrenceUa complex is difficult to assess,
partly because of the few species involved and their morphological diversity (apart from those in the
'Xanthochrysum' group). Of the taxa represented, BeZ/rVifl appears to be morphologically distinct and
I am continuing to recognise it as a monotypic genus. However, the morphology of the epidermal
cells of its testa suggests XhaiBellida and H. davenportii are more closely related to each other than
either is to H. Undleyi. With the exclusion oiBellida I am dividing the taxa into two genera based
on characters observed in the achene and pappus. These two genera consist of 1 ) Helichrysiim Undleyi
and H. davenportii, and 2) H. ayersii, H. cassinianum, and species of the Xanthochrysum' group.

Key to genera and species of the Lawrencella complex

1. Vascular strands of pericarp in lateral position in relation to cotyledons;
base of achene excavated

2. Apex of achene extended to form a pair of cup-shaped protuberances;

ray bracts absent Bellida

2. Apex of achene truncate; ray bracts present {Lawrencella)

3. Leaves terete, cauline Lawrencella Undleyi

3. Leaves flat, sub-basal Lawrencella davenportii

1. Vascular strands of pericarp in medial position in relation to coytledons;
base of achene not excavated (Schoenia)

4. Leaves flat (Schoenia subgroup)

5. Ray bracts present, pink or white Schoenia cassiniana

5. Ray bracts absent Schoenia ayersii

4. Leaves terete (Xanthochrysum subgroup)

6. Terminal barbs of pappus bristles densely clustered, clavate
7. Involucre narrow-cylindrical or narrow-turbinate; lamina of

ray bracts 3-4 mm long; plant cottony Schoenia ramosissima

7. Involucre hemispherical; plant shortly hirsute Schoenia macivorii

6. Terminal barbs of pappus bristles distinct, acute Schoenia filifolia

Bellida Ewart, Proc. Roy. Soc. Victoria 19:34(1907).

Type: Bellida graminea Ewart

Annual erect herb to 12 cm high with several leafless scapes arising from base, glabrous except
for white villi at base of .stems. Leaves basal, filiform. Capitula solitary, terminal. Involucre
turbinate, not radiant; bracts 3-seriate, hyaline with a narrow brown stereomereachingtoapex, central
nerve continuing as a piliferous tip. Receptacle hemispherical, glabrous. Florets numerous,
actinomorphic, outer bisexual, inner male. Corolla; tube shortly cylindrical, pilose; limb barrel-
shaped, glabrous; lobes 5, papillose within, sparsely pilose outside, vascular strands extending to
apex, cells of inner epidermis ± equilateral. Anthers; appendage narrow-cordate, proximal cells ±
equilateral and thick-walled, medial and distal cells oblong and thin-walled, marginal cells small;
collar narrow-oblong; tails extremely fine and exceeding collar. Style apex deltoid, acute to
acuminate, vascular strand stout, extending to apex. Achene compressed clavate, stipitate with

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Nuytsia Vol. 8, No. 3 (1992)

Figure!. Bellida graminea. A - achene with pappus and corolla. B - T.S. achene (semidiagramatic). C- achenial single celled
papillae and duplex hairs. D - portion of pappus bristles. E - seed (i.e. te.sta). F - epidermal cells of testa, (v.s. - vascular strand,
p - pericarp, t - testa, e - embryo.) From Kings Park 793/86.

excentric hollow base, moderalely to sparsely hirsute with duplex hairs and densely and minutely
velutinous with acute single-celled papillae (in smooth achenes these absent), not my xogenic, at apex
expanded into two cup-shaped extensions that bear the pappus; pericarp deeply and transversely
rugose (or occasionally smooth), thick and cartilaginous, a vascular strand in each margin of achene.
Seed compressed turbinate, free from pericarp; testa thin, vascular strand extending to apex in medial
position and at right-angles to vascular strands of pericarp. Pappus at anthesis about half as long as
the corolla, in fruit almost twice as long; bristles filiform, prominently dentate, reddish towards apex,
united into groups that are firmly fixed to each of the extensions of the achene, persistent.

A monotypic genus.

BellidagramineaEwart,Proc. Roy. Soc. Victoria 19:35(1907). Type.' Jibberding,Western Australia,

1905, M. Koch s.n. (iso: PERTH). (Figures 1 & 2)

Distribution. Found only in southern Western Australia.

Moore (19 17) indicated a relationship of Bellida to Helichrysim davenportii and treated them as
being congeneric, while Anderberg (1989) has suggested ihal Bellida may not even be a member of
the Gnaphalieae. 1 consider that the similarity of floral and fruit morphology between Bellida and
Lawrencella indicate that its position here is correct. The similarity is strongest to Lawrencella
davenportii which, if a true reflection of phylogeny, indicates that Lawrencella as treated here is
paraphyletic.

Paul G. Wilson, Lawrencella complex

369

Lawrencella Lindley, Sketch Veg. Swan R. Col. 23(1839).

HelichrysumstctLawrencella (Lindley) Benth.,Fl. Austral. 3:613(1867). Type: Lawrencella rosea
Lindley

Annual herbs, glandular puberulous or pilose. Stem single, erect, branched above. Leaves cauline
or sub-basal, opposite or alternate, terete or flattened. Capitula tenninal to branches or leafless scapes,
radiant. Involucre hemispherical; bracts 3-6-seriate. Outerand intermediate bracts narrow-triangular
to narrow-oblong, chartaceous, ciliate; stereome narrow-triangular, thin: innermost bracts: claw
oblong, hyaline with an oblong flat stereome; lamina elliptic, pink. Receptacle cushion shaped,
smooth, glabrous. Florets numerous, outer bisexual, inner male. Corolla actinomorphic, tubular
below, narrowly campanulate above, glandular hispidulous; cells of throat with straight walls; lobes
5, papillose within, vascular strands extending to tip. Anther appendage broad-ovate; proximal cells
equilateral, thick- walled, distal cells oblong, thin-walled, marginal cells small; collar narrow-oblong;
anther tails filamentous exceeding collar. Style apex deltoid, vascular strand, extending to tip.
Achenes polymorphic; base excavated forming a hollow extension of the pericarp, pericarp hard and
thick, smooth or rugose, glabrous or velulinous with minute aciformc 1-celled papillae or scabrous
with flattened acuminate multicelled trichomes or sparsely hispid with stiff duplex hairs; seed narrow-
fusiform with a short flattened sterile tip, free from pericarp; testa membranous, vascular strand
laterally placed with reference to cotyledons, terminating in the flattened tip. Pappus persistent,
colourless or pink, shorter than corolla at anthesis but lengthening in fruit; bristles firm, linear-
acuminate, prominently dentate, united in groups towards base.

Two species endemic to Australia.

Figures. Lawrencella davenportii. A - achene with pappus. B - T.S. achene (.semidiagramatic). C - trichomes from achene
(left papilla from surface of achene. centre .and right from near apex of achene). D - Apex of p.appus bristle. E- seed (i.e. testa).
F - epidemial cells of testa, (v.s. - vascular strand, p - pericarp, t - testa, e - embryo.) From AS. George 3924,

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NuytsiaVol.S, No. 3 (1992)

Lawrencella davenporfii (F.Muell.) Paul G. Wilson, comb. nov. (Figures 1 & 3)

Helichrysum davenportii F.Muell., Fragm. 3:32(1862). - Helichrysum lawrencella var. davenportii
(F.Muell.) Benth., FI. Austral. 3:616(1867). - Helichtjsum roseum (Lindley) Druce var. davenportii
(F.Muell.) Domin, Vestn. Krai. Ceske Spolecn. Nauk. Tr. Mat.-Pir. 2:120(1923). Type citation: 'In
Australia centrali ad flumen Neales. J.Macd. Stuart.' Lectotype (here chosen): Neales River (MEL
579934; isolecto: MEL s.n.).

Bellida major S.Moore, J. Bot. 55:100(1917). Type citation: 'Western Australia, Mulline;
J.E.C. Maryon, 1916.' Type n.v.

Distribution. Found in Western Australia, Northern Territory, and South Australia.

Lawrencella rosea Lindley, loc.cit. (Figures 1 & 4)

HelichrysumlawrencellaBen\h.,F\.A\is\iai.3:616{lS67)nom.illeg.-Helichrysumroseum(Lmd]ey)
Druce, Bot. Soc. Exch. Club Brit. Isles 4:626(1917); Domin, Vestn. Krai. Ceske Spolecn. Nauk. Tr.
Mat.-Prir. 2:120(1923) comb.illeg. non (Hook.) Baillon (1886). - Helichrysum lindleyi H. Eichler,
Taxon 12:295(1963). Type.' Vasse River, on the South West coast of New Holland, 1^39, Mrs Molloy
(holo: CGE photo seen).

Distribution. Southern Western Australia.

1

Figure4. Lawrencella lindleyi. A -achene with pappus. B - L.S. achene. C-T.S. achene(semidiagramatic). D- achenial duplex
hair. E - Apex of pappus bristle. F - seed (i.e. testa). G - epidermal cells of testa, (v.s. - vascular strand, p - pericarp, t - testa,
e - embryo.) From P.G. Wilson 12353.

Paul G. Wilson, Lawrencella complex

371

Schoenia Steetz in Lehm., Pl.Preiss. 1:480(1845).

Pleropogon sect. Schoenia (Steetz) F.Muell., Linnaea 25:415(1853). - Helichrysum sect. Schoenia
(Steetz) Baillon, Hist.Pl. 8:175(1882). Lectotype (here chosen): Schoenia oppositifolia Steetz
[= S. cassiniana (Gaudich.) Steetz].

Xanthochrysumi:wxcz.,'&\i\\. Soc. Imp. Naturalistes Moscou 24/1 : 199(185 1). Type:Xanthochrysum
filifolium Turcz.

Helipterum sect. Geniosperma A.Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:230(1852). Type:
Helipterum tenellum A.Gray

P/eropogon sect. //c///;t£’ro/;i/5F.Muell., Linnaea 25:415(1853). Lectotype (herechosen):Prera/?og£)/i
ramosissimus F. Muell.

Annual erect herbs, hirtellous with short uniscriate hairs that, when young, bear a curled
filamentous apiculum producing a woolly cover, or shortly hirsute, or glandular puberulous. Leaves
opposite or alternate, oblong or terete. Capitula solitary or corymbiform. Involucre hemispherical
to narrow-cylindrical, radiant or not; bracts c. 5-seriate, sometimes woolly ciliate, otherwise glabrous
or puberulous on the stereome of the outer bracts; outer bracts scarious-ovate, glossy with a triangular
green flat stereome. Inner bracts present or absent: claw oblong, scarious with an oblong stereome;
lamina elliptic, pink or yellow (or white). Receptacle convex, glabrous or glandular papillose. Rorets
actinomorphic, the outer bisexual and the inner male. Corolla narrow-cylindrical with narrow-
turbinate limb, 5-lobed, sparsely glandular-puberulous; lobes papillose or colliculate within, vascular
strands extending to tip. Anthers: appendage cordate, proximal cells ± equilateral and thick-walled,
medial and distal cells oblong and thin-walled, marginal cells small and equilateral; collar mu'row-
oblong; tails extremely fine and exceeding the collar. Style apex deltoid, acute, papillose, vascular
strand stout and extending to tip. Achene terete to compressed-obovoid, sparsely to densely hirsute
with somewhat rigid hairs, short gland-tipped hairs also present; apical margin sometimes raised and
surrounding base of pappus; pericarp cartilaginous, thin and crustaceous, or papery, vascular strands
in centre of ventral and dorsal faces and in medial position; carpopodium annular, insignificant; base
not excavated; seed free from or adherent to pericarp. Testa thinly coriaceous, vascular strand 2/3
orcompletelyencircling seed in themedial position. Pappus bristles narrow-linearat base and filiform
towards apex, dentate, colourless or yellow, free but united in a short ring at base, persistent or
deciduous (persistent on the sterile achenes).

Five species endemic to Australia.

Schoenia subgroup

Schoenia ayersii (F.Muell.) J. Black, Trans. & Proc. Roy. Soc. S. Australia 39:840(191 5). (Figures 1
&5)

Helichrysum ayersUT. Muell., Fragm. 8; 167(1874). Type citation: 'In vicinia montis Olgae; Gosse'.
Type: Gosse’s Exped., 1873 (holo: MEL).

Podolepis georgei Diels, Bot. Jahrb. Syst. 35:619(1905). Type citation: 'Hab. in distr. Austin pr.
Murrinmurrin, unde misit cl. W.J. George (Hb. Berol.!)'. Type non vidi.

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Nuytsia Vol. 8, No. 3 (1992)

Distribution. Found in southern Western Australia, Northern Territory, and South Australia.

A species readily distinguished from Podolepis, in which genus it was placed by both Diels (1905)
and Davis (1957), by its triangular style apex and its turbinate rugose achenes with coarse hairs.

Schoeniacassiniana(Gaudich.)SteetzinLehm.,Pl.Preiss. \-A?,\{\%A5). - Helichrysumcassinianum
Gaudich. in Freyc., Voy. Uranie 466 t.87(1830). - Helipterum cassinianum (Gaudich.) DC., Prod.
6:l\6{\%'i%).-Pteropogoncassinianus{Gwi(l\dn) F. Muell.,Linnaea 25:415(1853). LcGo/ype (here
chosen): Baie des Chiens-Marins, C.Gaudichaud (P photo seen, isolecto G-DC photo seen).
(Figures 1 & 6)

Schoenia oppositifolia Steetz in Lehm., PI. Preiss. 1:480(1845). - Pteropogon oppositifolius (Steetz)
F.Muell.,Linnaea 25:415( 1853). Type citation: 'Speciminaintegraleg. cl. Roe in Australasia australi-
occidentali inter flumen Swan-river et sinum regis Georgii 111. (V.s. in herb, aulico Vindobonnensi!)',
non vidi.

Distribution. Found in Western Australia, Northern Territory, and South Australia.

Figures. Schoenia ayeKsii. A - achene with pappu.s. B - L.S. - achene. C - T.S. achene (semidiagramatic). D - achenial duplex
hair. E - apex of pappus bristle. F - seed (i.e. testa). G - epidenual cells of testa, (v.s. - vascular .strand, p - pericarp, t - testa,
e - embryo.) From H. Demarz 6588.

Paul G. Wilson, Lawrencella complex

373

Figure 6. Schoenia cassiniana. A - achene with pappus. B - T.S. acliene (scmidiagramatic). C - achenial duplex hair. D - apex
of pappus bristle. E - seed (i.e. testa). F- epidennal cells of testa, (v.s. - vascular strand, p- pericarp, t - testa, e - embryo.) From
P.G. Wilson 12600.

Xanthochrysum subgroup

Schoenia filifolia (Turcz.) Paul G. Wilson, comb. nov. (Figures 1 & 7)

Xanthochrysum filifolium Turcz.. Bull. Soc. Imp. Naturalistes Moscou 24/1:199 t.4(1851). -
Helichrysumfilifolium (Turcz.) F. Muell.,Fragm. 3: 134(1863). - Type: Western Australia,/. Drummond
3rd coll. no. 1 19 (holo: KW photo seen; iso: K, MEL, NSW).

Helipterum tenellum A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:231(1852) nom. illeg. non Turcz.
(1851). Type: 'Swan River, Drummond' (holo: K).

Helichrysum subulifoliumV . MuelL.Fragm. 3:134(1863). Type citation: 'Ad sinum Champion Bay
Australiae occidentalis. P. Walcott.' Type: Champion Bay, P. Walcott (holo: MEL; iso: K).

Helichrysum turbinatum W. Fitzg., J. W. Austral. Nat. Hist. Soc. No.l:23(May 1904). - Helichrysum
pseudoturbinatum C. Gardner, Enum. PI. Austral. Occ. 133(1931) nom. illeg. Type: Nannine, Sept.
1903, W.F. Fitzgerald (iso: PERTH).

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Nuytsia Vol. 8, No. 3 (1992)

nooo

0.2 mm

Figure?. Schoeniafilifolia&uh^p.suhuUfolia, A - achene with pappus. B - T.S. achene (semidiagramatic). C - achcnial duplex
hair. D - apex of pappus bristle. E - seed (i.e. testa). F - epidennal cells of testa, (v.s. - vascular strand, p - pericarp, t - testa,
e - embryo.) From A. Morrison, 30.ix.l904.

Key to subspecies

1. Involucre turbinate to cylindrical; ray 3-6 mm long

2. Plant single-stemmed; involucre turbinate subsp.//7i/o/;a

2. Plant multi-stemmed; involucre cylindrical subsp. arenicola

1. Involucre hemispherical; ray 7-10 mm long subsp. subulifoUa

subsp. filifolium

Helipterum tenellum A. Gray, l.c.

Helichrysum turbinatum W. Fitzg., l.c. - Helichrysiim pseudoturbinatum C. Gardner, l.c., nom. illeg.

Distribution and habitat. Inland south-west Western Australia from Mullewa to Kalgoorlie and Lake
Barker in saline areas.

Paul G. Wilson, Lawrencella complex

375

The typical variant of suhsp. filifolium has obovoid achcnes and yellow ray laminae 4-6 mm long
whereas the variant represented by the type of H. tiirhinatum, which is immature and only known from
that collection, possibly differs in having short (c. 3 mm long) white laminae to the inner involucral
bracts, and narrow, quadrangular, very sparsely hispidulous achenes. Its type was collected 'in
crevices of rocks along the shores of a salt lake'.

In the above taxonomy suhsp.filifolia remains somewhat variable in morphology even with the
segregation of the other two subspecies, however, it is not at present reasonable to describe further
infraspecific taxa since only a few collections have been made.

subsp. arenicola Paul G. Wilson, subsp. nov.

Herba erecta supra basim ramosa ad 30 cm alta. Inflorescentia laxe corymbosa. Involucrum
cylindraccum c. 7 mm altum; bractea intima laminis ovatis luteis c. 5 x 2.5 mm. Achenium teres, pilis
duplicibus crassis c. 0.2 mm longis.

Typus: Western Australia, 5 miles S.E. of Carnarvon, 4 Sept. 1959, N.T. Biirbidge 6502 (holo:
PERTH; iso: BRI, CANB).

Annual erect herb branching above, to 30 cm high. Inflorescence of open corymbs. Involucre
cylindrical, c. 7 mm high; innermost bracts a with yellow ovate lamina c. 5 x 2.5 mm. Achenes terete;
duplex hairs thick, c. 0.2 mm long.

Additional collections seen. WESTERN AUSTRALIA: Champion Bay, 1889, Mrs Forrest (MEL
110575): Carnarvon, Sept./October 1964, T.Nysen (PERTH).

Distribution and habitat. Evidently confined to sandhills in the Carnarvon area of Western Australia.
The collection from Champion Bay, if the locality data is correct, indicates that it was once found as
far south as Geraldton.

Derivation of epithet. The epithet arenicola (sand dweller) refers to the plant’s habitat preference.

This subspecies differs from subsp. filifolia in being single-stemmed (not with several major
branches), in having a cylindrical involucre (not turbinate or hemispherical), and in having terete
achenes with short (c. 0.2 mm long) duplex hairs (not obovoid and with hairs c. 0.4 mm long).

subsp. subulifulia (F. Muell.) Paul G.Wilson, comb, et stat. nov.

Helichrysum siibul ifolium F. Muell., Fragm. 3:134(1863). Type citation: 'Ad sinum Champion Bay
Australiae occidentalis. P.Walcott.' Type: Champion Bay, P. Walcott (holo: MEL; iso: K).

Distribution. Western Australia, Geraldton district.

The typical variant of this subspecies differs from suhsp.filifolia in having much larger capitula
and larger achenes with longer and more dense hairs, but some collections suggest that the two
subspecies grade into each other.

376

Nuytsia Vol. 8, No. 3 (1992)

Schoenia macivorii (F. Muell.) Paul G. Wilson, comb nov.

Helichrysum macivorii F. Muell., S. Sci. Rec.3:99(1883). Type: Gascoyne River, 1882, J. Forrest
(iso: MEL two sheets, PERTH).

Distribution and habitat. Gascoyne River area on sand or loam.

Schoenia ramosissima (F. Muell.) Paul G. Wilson, comb. nov.

PteropogonramosissimusF.Mucl\.,Limtita25:4l2(li53).-HelichrysimsemifertileF.Muell.,R&p.
PI. Babbage Exped. 14(1859) - Helichrysum ranwsissimum (F. Muell.) Druce, Bot. Exch. Club Brit.
Isles 4:626(1917) nom. illeg. non Helichrysum ramosissimum Hook.(1848). Type citation: 'In
planitiebus arenoso-argillaceis inter monies Flinders-range et sinum Spencers-gulph etiam non procol
a rupe Cudnaka.' Lectotype (here chosen): In planitiebus valibusque sterilibus prope Cudnaka et
WalIendunga,Oct. [18]51,DrM[ueller] (MEL 604822). Para/ec/ofypex.' Cudnaka, Nov. Holl.austr.
inter., F. Mueller (MEL); Inter montis Flinder’s & Sinum Spenceri, Oct. F .Mueller (MEL).

Helichrysum semifertile var. xanthoglossum F. Muell., Rep. PI. Babbage Exped. 14(1859). Type
citation: 'Wonnomulla, Elizabeth Creek'. Type: Smith’s Waterhole, anon, (holo: MEL).

Distribution. Found in Northern Territory, Queensland, New South Wales and South Australia.

This species has a variant with white and a variant with yellow bract ray laminae. All of the
syntypes of Pteropogon ramosissimus have white laminae and evidently for this reason Mueller
described as a variety a yellow-rayed plant collected on the 'Babbage' expedition.

The presumed holotype of//, semifertile vas.xanthoglossum'istismtiW specimen mounted in a blue
packet that is labelled 'Pteropogon ramosissimus fi flavisimus. Smith’s Waterhole'; it bears no other
information. Smith’s Waterhole is one of the localities visited by B. H. Babbage on his expedition
in 1858. The locality is also referred to in the 'Report' by the Aboriginal name Wirra-Wirralu which
is staled to be near Elizabeth Creek, the place name mentioned by Mueller in his protologue to var.
xanthoglossum. Since the specimen agrees with Mueller’s description, and is the only collection in
herb MEL of that species from the 'Babbage' expedition, 1 am confident that it is the type.

Acknowledgements

I am grateful to Arne Anderberg, Laurie Haegi and Jim Armstrong, who made many constructive
comments on an early manuscript, and to Margaret Menadue who provided the illustrations. The
figures of some of the transverse sections of achenes were based on slides kindly provided by Phil
Short.

Paul G. Wilson, Lawrencella complex

377

References

Anderberg, A.A. (1989). Phylogeny and reclassification of the tribe Inuleae (Asteraceae). Canadian J. Bot. 67:2277-2296.
Anderberg, A.A. (1991). Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Botanica 104:1-195.
Bentham, G. (1867). "Flora Australiensis", vol. 3. (Lovell Reeve & Co., London.)

Black, J.M. (1915). Scientific notes on an expedition into the north-western regions of South Australia (j) Botany. Trans. &Proc.
Roy. Soc South Australia 39:823-842.

(1929). "Flora of South Australia." (Government Printer, Adelaide.)

(1957). "Flora of South Australia." Edn 2. (Government Printer, Adelaide.)

Haegi, L. (1986). Helichrysum Miller. !n J.P. Jessop & H.R. Toelken (eds). "Flora of South Australia." Edn 4. (Government
Printer, Adelaide.)

Hilliard, O.M. & Burtt, B.L. (1981). Some generic concepts in Compositae - Gnaphaliinae. J. Linn. Soc., Bot. 82:181-232.
Jakupovic.J. etal. (1989). Diterpenes and other constituents from Auslralian//e//cfi/ 7 X«m and related species. Phytochemistry
28:543-551.

Merxmuller, H., Leins, P. & Roessler, H. (1977), Inuleae - systematic review. In V.H. Heywood, J.B. Harbome & B.L. Turner
(eds) "The Biology and Chemistry of Compositae.” (Academic Press, London.)

Mueller, F. (1889). "Second Systematic Census of Australian Plairts." (Govenunent Printer, Melbourne.)

Short, P.S., Wilson, K.E. & Nailon, J. (1989). Notes on the fmit anatomy of Australian members of the Inuleae (Compositae).
Muelleria 7:57-79.

Warcup, J.H. (1990). The mycorrhizal associations of Australian Inuleae (Asteraceae). Muelleria 7:179-187.

Wilson, P.G. (1989). A revision of the genus T/ya/ospcrma (Asteraceae: Inuleae: Gnaphaliinae). Nuytsia 7:75-101.

Wilson, P.G. (1992a). The classification of the Australian species currently included in Helipterum and related genera
(Asteraceae: Gnaphalieae): Part 1. Nuytsia 8:379-438.

Wilson, P.G. (1992b), The classification of Australian species currently included in Helipterum (Asteraceae: Gnaphalieae) Part
2 Leucochrysum. Nuytsia 8:439-445.

Zdero, C. el at. (1989). Sesquiterpene lactones and other constituents from Australian Helipterum species. Phytochemistry
28:517-526.

Nuytsia 8(3): 379-438 (1992)

379

The classification of Australian species currently included in
Helipterum and related genera (Asteraceae: Gnaphalieae): Part 1

Paul G. Wilson

Western Australian Herbarium, Department of Conservation and Land Management,
PO Box 104, Como, Western Australia 6152

Abstract

Wilson, Paul G. The classification of Australian species currently included in Helipterum and
related genera (Asteraceae: Gnaphalieae): Part 1. Nuytsia 8(3): 379-438 (1992). The name
Helipterum DC. is recognised as being illegitimate and not applicable to Australian taxa. The
Australian species currently included in that genus and in the related genera Cephalipterum, Gilherta,
Triptilodisciis and Pterochaeta are reclassified. One genus, Haptotrichion, is described as new.
\}nd&x Rhodanthe 11 sections are recognised of which 1 is new and 10 are new combinations. Eight
new species oiRhodanthe and one new species of Haptotrichion are described, these are /?. ascendens,
R. collina,R. cremea.R. gossypina,R. nullarborensis,R.psammophila,R. rufescens,R. sphaerocephala
and H. colwillii. A number of new species combinations are made. Descriptions are given for each
of the recognised genera and sections.

Contents

Page

Introduction

380

Rhodanthe

383

Rhodanthe sect. Rhodanthe

383

Rhodanthe sect. Monencyanthes

384

Rhodanthe sect. Leiochrysum

385

Rhodanthe sect. Achyroclinoides

399

Rhodanthe sect. Citrinae

406

Rhodanthe sect. Synachryum

408

Rhodanthe sect. Helipteridium

412

Rhodanthe sect. Helichrysoides

412

Rhodanthe sect. Actinaria

414

Rhodanthe sect. Anisolepis

415

Rhodanthe sect. Polyphyllum

416

380

Nuytsia Vol. 8, No. 3 (1992)

Contents

Page

Cephalipterum

417

Erymophyllum

418

Hyalosperma

419

Gilberta

419

Triptilodiscus

420

Haegiela

421

Pterochaeta

422

Haptotrichion

422

Acknowledgements

426

References

426

Index to Scientific Names

427

Index to name changes

436

Introduction

As has been pointed out by many syantherologists (Hilliard &Burtt 1981, Anderberg 1989, 1991,
Wilson 1989a), the genera Helipterum Lindley (1836), Helipterum DC. (1838) and Helichrysum
Miller, in the strict sense, do not occur in Australia and, in any event, the first name is illegitimate
and is to be typified by a South African species currently placed in Helichrysum (Wilson 1989a), while
the second is illegitimate and is to be typified by a South African species of Syncarpha DC.
(Nordenstam 1989).

The difference between the genera Helipterum DC. and Helichrysum, as enunciated by
A. de Candolle (1838), is that in f/e/ipteram the pappus bristles are plumose whereas in
they tend to be barbellate, or plumose only at the apex. A strict interpretation of this distinction can
lead to a placement of related species into separate genera, although the distinction has meant that
closely related species have usually been placed together in either Helipterum or in Helichrysum.

In order to classify the species currently included in Helipterum and Helichrysum it has been
necessary to study related genera of the Gnaphalieae {sensu Anderberg 1989) such as Leptorhynchus,
Ixiolaena, Waitzia, Podolepis, Podotheca, Myriocephalus, as well as several monotypic genera.

The revised classification has resulted in the recognition of additional genera and the rearrangement
of the species currently in Helipterum and Helichiysum. Some of the resulting genera are being
revised by other botanists whose work 1 do not wish to compromise by producing a comprehensive
classification lacking in the information that they will be able to provide. Therefore in this paper 1
am formally treating a portion of the Helipterum complex while indicating the genera into which other
species currently included in Helipterum are likely to be transferred.

Paul G. Wilson, Helipterum: Part 1

381

The classification that follows for a portion of the Helipterum complex in Australia attempts to
associate closely related species and to segregate as distinct genera those species or groups of species
that exhibit marked morphological discontinuity. In Rhodanthe I have not been successful in
interpreting the phylogenetic significance of the varied morphological forms that are adopted by
homologous organs. In the achene, for example, the pericarp and testa can have many different
textures and structures (Short etal. 1989), yet to generically separate taxa that have dissimilar achenes
can lead to the separation of species which otherwise have markedly similar florets and similar bract
morphologies. For this reason I have adopted a sectional rank for taxa that may later be shown to
deserve generic rank and which were sometimes so accorded by early 19th century taxonomists.

I have not attempted to place all the genera of the 'Angiantheae' sensu Bentham (1867) in context
with those of the Helipterum complex but it is obvious that a close relationship often exists. This type
of synoptic work will have to be done by others; my principal concern is to provide a classification
that can be used for a 'Flora of Australia' treatment and, from the pressure of time alone, many
interesting phylogenetic avenues must remain unexplored.

This paper considers the Rhodanthe, Hyalosperma and Triptilodiscus groups of species, most of
which have, in general, been placed in Helipterum. These groups can be informally categorized as
follows.

Rhodanthe group: Inner involucral bracts with broad flat claws; anthers with fine filamentous tails;
achenes with normal (mostly not thickened) duplex hairs. Species in this group, with the exception
of Rhodanthe citrina, that have been examined, do not form ectomycorrhiza (Warcup 1990).

Hyalosperma group: As above, however, achenes lacking normal duplex hairs but with 2-celled
bulbous papillae that are frequently myxogenic; pappus shed entire. Species in this group that have
been examined form ectomycorrhiza (Warcup op.cit.)

Triptilodiscus group: Species in this group have achenes with 2-celled tooth-like trichomes and
oblong or boat-shaped claws to the involucral bracts. Other characters suggest that the included
genera may not be closely related and that the group is polyphy letic; this therefore becomes a grouping
of convenience. In only 7’n/;///or//Tc«s’ has thcmycorrhizal association been investigated; it was found
to form ectomycorrhiza (Warcup op.cit.).

The genus Rhodanthe, which is by far the ku'gest of the segregate genera, is circumscribed to
contain the majority of the Australian species previously included in Helipterum. Some of the
remaining speciesare listed under other generain this paper. Those species excluded from the present
treatment are as follows:

Helipterum albicans (Cunn.) DC., Helipterum fitgibboniiV . Muell., //. molle (DC.) Paul G. Wilson,
and H. stipitatum (F. Muell.) Benth. have been transferred to Leucochrysum (DC.) Paul G. Wilson,
(see Wilson 1992a) a genus closely related to Wailzia.

Helipterum niveum Steetz is clo.sely related to Helichrysum obtusifolium Sonder; both species will
be segregated, with others, in a genus to be described.

Helipterum saxatile Paul G. Wilson is closely related to Helichrysum podolepidium F. Muell., these
two species will, with others, be placed in a genus to be described.

382

Nuytsia Vol. 8, No. 3 (1992)

Helipterum craspedioides is conspecific with Myriocephalus morrisonianus and is possibly best
recognised as a member of the genus Polycalymma (but see Short et al. 1989 and Anderberg 1991).

Helipterum adpressum W. Fitzg. (= Helichrysum puteale S. Moore) and Helipterum pterochaetum
(F. Muell.) Benth. have been transferred to the genus Chrysocephalum Walp. as C. puteale
(S. Moore) Paul G. Wilson and C. pterochaetum F. Muell. (see Anderberg 199 1 and Wilson, Short
& Orchard 1992).

The species that are here placed in Rhodanthe form apolymorphic assemblage that is divided into
eleven sections. Some of the sections that have obvious problems associated with them are briefly
discussed below.

The sect. Monencyanthes, in which the achenes are persistent and the capitula caducous, contains
three species. These species probably attained their specialised condition independently for each has
features that suggest derivation from a different ancestor possibly of the Achyroclinoides type.

The species in sect. Leiochrysum are diverse in vegetative and floral morphology. The organs
appear to vary in morphology independently from each other and 1 have been unable to correlate
characters in a manner that would achieve a more natural classification.

The sect. Helichrysoides appears to be a monophyletic group which, at one extreme has affinity
to the genus Podotheca and, at the other, to species in sect. Leiochrysum. Its correct taxonomic status
is uncertain (see also Anderberg 1991).

Each of the sections Anisolepis (1 sp.), Actinaria (2 spp.), and Helipteridium (1 sp.), has very
distinctive involucral bracts but each shows little difference in floral morphology to certain species
in sect. Achyroclinoides-, 1 have therefore included these sections in Rhodanthe rather than give them
generic status.

The sect. Citrinae, with the single species R. citrina. has been difficult to place. Due to the beaked
apex to its achene this species has been previously included in Waitzia. It differs radically from
Waitzia in that the involucral bracts have flat claws with an undivided .stereome (fide Anderberg 1991),
the style apex is truncate, and the achenes pilose. It differs from other species of Rhodanthe in
sometimes having the outer florets female (not hermaphrodite) and in being an ectomycorrhizal
species (Warcup 1990). Both these characters suggest a relationship with species such as
Chrysocephalum apiculatum (Labill.) Steetz and Leucochrysum albicans (Cunn.) Paul G. Wilson,
but the morphology of the achenes and corollas of the three species is quite different. It possibly
deserves recognition as a distinct genus.

Rhodanthe polyphyllum, which is placed in the monotypic section Polyphyllum, is a further
anomaly since it has stout, branched anther tails and clustered, filiform leaves. It appears to have no
close relative in the Helipterum - Helichrysum complex . Nevertheless I have included it in Rhodanthe
with which it appears to have greatest affinity. Again, it may deserve recognition as a distinct genus.

Paul G. Wilson, Helipterum: Part 1

383

Catalogue of taxa

In this Catalogue I have listed the accepted names and their synonyms in the Rhodanthe,
Hyalosperma and Triptilodiscus groups. For new taxa and for all genera and sections I have provided
descriptions. Where it has been necessary I have lectotypified names.

The Rhodanthe group

Rhodanthe Lindley

Rhodanthe Lindley, Bot. Reg. t.l703(1834).

Helichrysum sect. Rhodanthe (Lindley) Baillon, Hist. PI. 8:314(1886). Type: R. manglesii Lindley

Roccardia Necker ex Voss, Vilm. Blumengartn. ed.3, 1:530(1895) nom. illeg. non Necker ex
Raf.(1838). Type: R. manglesii (Lindley) Voss [=Rhodanthe manglesii Lindley].

Annual or short-lived perennial herbs, woolly, pubcrulous, pilose, glandular, or glabrous. Leaves
mostly alternate, simple, entire, rarely terete. Capitula homogamous or heterogamous, solitary or
clustered, radiant or not; subtending leaves, when present, grading into the outer bracts. Involucral
bracts multi- seriate, glumaceous or scarious, rarely with a herbaceous tip: claws oblong with nanow
or broad undivided stereome (Andcrberg 1991). Receptacle glabrous, or pilose around alveolae,
predominantly ebracteate. Florets bisexual or the innermost male, otherwise homomorphic. Corolla
tubular at base, usually turbinate above, glabrous or sparsely puberulous; lobes 5, equal or not, inner
epidermis with cells not oblong in transverse rows. Anther tails filamentous and weak. Style apex
truncate to ellipsoid or deltoid. Achene with normal duplex hairs; carpopodium annular or absent.
Pappus bristles barbellate to plumose.

1. Rhodanthe Lindley sect. Rhodanthe

Annual erect herb, glabrous. Leaves cauline, sessile, suborbicular, base decuirent on each side
of stem. Capitula heterogamous, solitary on slender branches. Involucre turbinate; bracts multi-
seriate, papery, petaloid; claw with scarious margin and narrow-oblong thin stereome. Receptacle
rounded, smooth, glabrous. Florets numerous, mainly bisexual but the innermost male, 5-mcrous.
Corolla actinomorphic, glabrous; upper portion cup-shaped; lobes not papillose; veins extending into
base of lobes. Stamens: anther appendage cordate, acute, the cells narrow-oblong towards apex,
equilateral near base; anther tails filamentous, weak; collar niurow-oblong. Style apex narrow-
deltoid,va.scu!arlracc notextending to tip. Achene obovoid.dorsiventrally compressed, densely silky
to base; carpopodium absent; pericarp thin, translucent, in medial position in relation to the
cotyledons; testa thin, translucent, ± fused to pericarp, the vascular strand extending almost
completely around seed; crystals absent. Pappus persistent; bristles plumose, the cilia all acute.

A monotypic section that is most closely related to species in sect. Leiochrysum. It is distinctive
in being glabrous, in having suborbicular sessile decurrent leaves, and possessing an anther appendage
and a style apex that are unique in the genus.

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Nuytsia Vol. 8, No. 3 (1992)

RhodanthemanglesiiLindley,Bot.Reg.20: 1.1103(1^34). - Helipterummanglesii {Und\cy)B£nth.,
FI. Austral. 3:640(1867). - Helichrysum manglesii (Lindley) Baillon, Hist. PI. 8:314(1886). -
Argyrocome manglesii (Lindley) Kuntze, Revis. Gen. PI. 1:309(1891). - Roccardia manglesii
(Lindley) Voss, Vilm. Blumengartn. 3rd edn, 1:531(1895). Type: Swan River Colony, leg.
J. Drummond, n.v.

Rhodanthe atrosanguinea J. Drumm. ex hort., Card. Chron. no. 27 :622(1 861). Type: Limestone Hills,
Champion Bay, W.A., J. Drummond, n.v.

Rhodanthe manglesii var. sanguinea Hook., Bot. Mag. 87:1.5283(1861). - Roccardia manglesii f.
sanguinea (Hook.) Voss, Vilm. Blumengartn. 3rd edn, 1:531(1895). Type: Western Australia,
J. Drummond, n.v.

Roccardiamanglesiif.ligulosaVoss,Vi\m.B\iimer\gMr].3rdedn, 1:531(1895). Type.' not indicated.

Helipterum cryptanthum O. Sarg., J. Bot. 61:285(1923). Type: Mt Brown, York, Western Australia,
0. Sargent 1404 (holo: BM; ?iso: MEL 109635 Sargent 1404a).

2. Rhodanthe sect. Monencyanthes (A. Gray) Paul G. Wilson, comb. nov.

Monencyanthes A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:229(1852).

Helipterum sect. Monencyanthes (A. Gray) Benth., FI. Austral. 3:648(1867). Type: Monencyanthes
gnaphalioides (Hook.) A. Gray

Annual herbs, variably woolly. Leaves cauline, linear to narrow-oblong, entire. Capitula
heterogamous, sessile or subsessile in terminal glomerules, caducous. Involucre ellipsoidal, narrow-
cylindrical, or cup-shaped, not radiant; bracts scarious, woolly ciliate, the inner with linear hard claw.
Receptacleinsignificantorrounded, with or without receptacular bracts. Florets 1-c. 12, bisexual or
male, 5-merous. Corolla aclinomorphic, cylindrical or narrow-turbinate, glabrous. Stamens: anther
appendage broad ovate, obtuse, the cells narrow-oblong, thin- walled; anther tails filamentous, weak.
Style apex truncate, papillose. Achene ellipsoidal, c. 2.5 mm long, persistent on receptacle, sparsely
short pilose; carpopodium absent; pericarp membranous, hyaline, and with crystals (M . maryonii) or
outer layer with thickened brown radial walls and crystals absent, nerves medial or medial-oblique
in relation to cotyledons (Short et al. 1989); testa membranous and fused to pericarp or somewhat
leathery and free (M. maryonii), crystals present. Pappus persistent, the bristles plumose weak with
a few long cilia.

Notes. This taxon was treated in a very broad sense as a section of Helipterum by Bentham (1867)
so as to include some species now placed in Chrysocephalum, Rhodanthe sect. Achyroclinoides, and
Hyalosperma. I have restricted it to those species with a woolly indumentum, and caducous capitula
with persistent shortly hirsute achenes that have a persistent pappus. The included species, though
superficially similar, probably arose independently from different species with caducous achenes
since in the microscopic structure of the corolla each displays a significantly different morphology.
It is interesting that in the achene of/?, maryonii the pericarp bears crystals (presumably of calcium
oxalate) whereas in all other species of Rhodanthe that have crystals in the achene they are found only

Paul G. Wilson, Heliplerum: Part 1

385

in the testa. Short e/ a/. (1989) record that of the members of the Inuleae that they examined, crystals
in the pericarp were only found in Craspedia aff. pleiocephala.

Rhodanthe moschata (Cunn. ex DC.) Paul G. Wilson, comb. nov.

Gnaphalium moschatum Cunn. ex DC., Prod. 6:236(1838). - Helipterum moschatum (DC.) Benth.,
FI. Austral. 3:648(1867). Argyrocome moschata (DC.) Kuntze, Revis. Gen. PI. 1:309 (1891). Type:
Mode’s Plains, Lachlan River, Interior of New South Wales, 1817,/l.CMn«/>i^/iam(holo:G-DCphoto
seen).

Calocephalus gnaphalioides Hook, in T. Mitch., J. Exped. Int. Trop. Austral. 378(1849). -
Monencyanthes gnaphalioides (Hook.) A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 6:230(1852).
Type: T.L. Mitchell, n.v.

This species is often confused with R. tietkensii which may be distinguished by its more numerous
florets (c. 10 to a head), and absence of prominent glandular hairs on the corolla lobes. In R. moschata
there are c. 3 florets to a capitulum (1 or 2 bisexual) and prominent glands on the lobes.

A collection from the Simpson Desert, Northern Territory (G. Chippendale, NT 6577, herb. AD)
appears to be a hybrid between R. tietkensii and R. moschata. The herb. NT duplicate of this collection
is of true R. moschata. Rhodanthe tietkensii has been found in the same area.

Material collected in northern Eyre Peninsula, South Australia, differs from the common variant
in having milky white (not scarious) involucral bracts.

Rhodanthe uniHura (J. Black) Paul G. Wilson, comb. nov.

Helipterum uniflorum J. Black, Trans. & Proc. Roy. Soc. South Australia 41:651 t.43(1917).
Lectotype (here chosen): Koppermanna, 7 Oct. 1916, S.A. White (AD 98625090 p.p.), isolecto:
MEL 110472, NSW 181468.

Rhodanthe maryonii (S. Moore) Paul G. Wilson, comb. nov.

//e/ipterH/nwaryoniiS. Moore, J. Linn. Soc. Bot. 45: 182(1920). Type:Mu\lino,J.E.C.Maryon(holo:
BM; iso: MEL 110713, NSW 181424).

3. Rhodanthe sect. Leiochrysum (DC.) Paul G. Wilson, comb. nov.

Helipterum sect. Leiochrysum DC., Prod. 6:216(1838). Lectotype (here designated): Helipterum
polygalifolium DC. [^Rhodanthe polygalifolia (DC.) Paul G. Wilson],

Helipterum sect. Sericophorum DC., Prod. 6:216(1838). Lectotype (here designated): Helipterum
humboldtianum (Gaudich.) DC. [=Rhodanthe humboldtiana (Gaudich.) Paul G. Wilson].

Pteropogon DC., Prod. 6:245( 1 838). - Pteropogon sect. Facelioides A. Gray, Hooker’s J. Bot. Kew
Gard. Misc. 4:267(1852) nom. illeg. - Helipterum sect. Pteropogon (DC.) Benth., El. Austral.
3:639(1867). Type: P. pygmaeus DC.

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Nuytsia Vol. 8, No. 3 (1992)

Xyridanthe Lindley, Sketch Veg. Swan Riv. Col. 23(1839). Type: X. stricta Lindley

Acroclinium A. Gray, Hooker’s J.Bot.Kew Card. Misc. 4:270(1852). -Wc//c/(ryiMm sect. Acrod/VuMw
(A.Gray) Baill., Hist. PI. 8:174, 314(1882); F. Muell., Fragm. 3:32(1862) nomen. Lectotype (here
designated): A. multicaule A. Gray [=Rhodanlhe chlorocephala (Turcz.) Paul G. Wilson].

Grijfithia J. Black, Trans. & Proc. Roy. Soc. South Australia 37:122(1913). Type: G. helipteroides
J. Black [=Rhodanthe oppositifolia (S. Moore) Paul G. Wilson].

Annual herbs (rarely short-lived perennial), glabrous or variously pubescent. Leaves linear to
oblong, entire. Capitulaheterogamousorhomogamous, solitary or clustered. Involucre globular to
cylindrical or turbinate, radiant or not; bracts multiseriate, scarious, papery, or thinly cartilaginous;
stereome oblong to linear, thin; claw of inner involucra! bracts flat, scarious. Receptacle rounded to
conical, smooth or pitted, glabrous or puberulous. Florets 5 to numerous, bisexual, or the innermost
male, 5-merous. Corolla actinomorphic to zygomorphic, almost glabrous to variously pubescent,
cylindrical or expanded above, cells of inner epidermis of lobes equilateral to broad-oblong, smooth
or papillose. Stamens: anther appendage narrow-deltoid to ovate or oblong, cells narrow-oblong;
anther tails filamentous, weak; collaroblongornarrow-oblong. Styleapex truncate to broadly deltoid,
prominently papillose. Acheneobovoid, compressed obovoid, or ellipsoid, L5-3.5 mm long, densely
silky or moderately short pilose; carpopodium a short glabrous slightly tumid hollow ba.se or absent;
pericarp crustaceous, thinly coriaceous, or papery , vascular strands in medial or oblique position; testa
membranous to thinly coriaceous, free from or fused to pericarp, the vascular strand 1/2 to 3/4
encircling seed; crystals in testa present or absent. Pappus persistent or caducous; bristles linear-
lanceolate, plumose, the terminal cilia frequently rounded or clavate.

Note 1 . Candolle (1838) placed llelipieniinpolygalifaliummd //. dijfusiim in his section Leiochrysum,
I have selected the first of the.se as the lectotype since it better agrees with his sectional description.

Note 2. Gray (1852) included two species in Pteropogon sect. Facelioides of which one was
P. pygmaeus, the type of Pteropogon. His sectiomd name is therefore superfluous.

Rhodanthe anthemoides (Sprengel) Paul G. Wilson, comb. nov.

Helichrysum anthemoides Sieber ex Sprengel, Syst. Veg. 3:484(1826). - Helipterum anthemoides
(Sprengel) DC., Prod. 6:216(1838). - Argyrocome anthemoides (Sprengel) Kuntze, Revis. Gen. PI.
1:309(1891). - Roccardia anthemoides (Sprengel) Voss, Vilm. Blumengartn. 3rd edn, 1:531(1895)
' anthemodes' . Type: 'Nov Holl.' probably F.W. Sieber 344 (iso: G-DC photo seen, K, MEL 109048,
604823).

Helipterum punctatum DC., Prod. 6:216(1838). Type: Van-Diemen Land, R.Gunn 239 (holo: G-DC
photo seen; ?iso: MEL 109067 p.p.).

Rhodanthe chlorocephala (Turcz.) Paul G. Wilson, comb. nov.

Schoenia chlorocephala Turcz., Bull. Soc. Imp. Naturalistes Moscou 24/l:193(March 1851). -
Helipterum chlorocephalum (Turcz.) Benth.,Fl. Austral. 3:641(1867). - Argyrocome chlorocephala

Paul G. Wilson, Helipterum: Part 1

387

(Turcz.) Kuntze, Revis. Gen. PI. 1:309(1891). - Roccardia chlorocephala (Turcz.) Voss, Vilm.
Blumengartn. 3rd edn, 1:530(1896). Type: Western Australia, J. Drummond 4th coll. n. 199 (iso:
MEL 109146, 109147, 109148).

Rhodanthe chlorocephala subsp. chlorocephala

[Acroclinium multicaule A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:271(1852) as to var. 6 not as
to lectotype. Syntype: Swan river, J. Drummond (K).]

I am applying the epithet subsp. chlorocephala in a narrow sense so as to restrict it to the form
represented by the type collection of this taxon. This type was collected from an unknown locality
in the south-west of Western Australia: it was included in Drummond’s fourth collection which was
despatched to England in July 1847. According to Erick.son (1969) the fourth collection consisted
of plants gathered from King George Sound, Stirling Range, Porongorups, Mt Manypeaks, Cape
Riche, West Mt Barren, and from north and east of Moore River. Of these various localitiesR/ioi/ant/ie
chlorocephala is only known from the Moore River area but the only other specimens 1 have been
able to match with the type me those that represent the original material of Acroclinium multicaule
A.Gray var. 6 which is an apparently undated and unnumbered collection of Drummond’s in herb.
K. It is possible that the specimens that form the bases of R. chlorocephala and A. multicaule var.
6 are from the same Drummond collection.

Rhodanthe chlorocephala subsp. rosea (Hook.) Paul G. Wilson, comb, et subsp. nov.

Acroc/m/Mmrarmn Hook., Bot.Mag. 1.4801(1854). -7/e/;c/!r>’rHmgray/F.Muell.,Fragm. 5:200(1866).
- Helipterum roseum (Hook.) Benth., FI. Austral. 3:640(1867). - Argyrocome rosea (Hook.) Kuntze,
Revis. Gen. PI. 1:309(1891). - Roccardia rosea (Hook.) Voss, Vilm. Blumengartn. 3rd edn,
1:53 1( 1896). Type citation; 'A native of the interior of South-west Australia, between the Moore and
Murchison rivers from whence the seeds were sent in 1853 by Mr. James Drummond, together with
dried specimens (No. 1 57 of his Herbarium ).' Lectotype (here chosen): J. Drummond 6th coll. no. 157
(lecto: K; isolccto: NSW 180743). Syntype: Hort. Kew. 1854 from Drummond S.W. Australia (K).

Acroclinium multicaule A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:271(1852). - Helichrysum
drummondii F. Muell., Fragm. 5:200(1866). Lectotype (here chosen): Swan River, J. Drummond
(lecto: ?K n.v.; ?iso: P, 'J. Drummond 349’, annotated by A. Gray in 1851).

Acroclinium roseum var. grandiflorum Nicholson, 111. Diet. Gard. 1:18(1884). - Helipterum roseum
var. grandiflorum (Nicholson) Chitt., Diet. Gardening 2nd edn, 2:978(1951). Type: not indicated.

Acroclinium roseum var. album hort. ex Nicholson, 111. Diet. Gard. 1: 18( 1884). - Helipterum album
(Nicholson) L. Bailey, Cycl. Amer. Hort. 2:726(1900). - Helipterum roseum f. album (Nicholson)
Moldcnke, Phytologia 2:312(1947). Type: not indicated.

Helipterum roseum var. nigropapposum Ostenf., Biol. Meddel. Kongel. Danske Vidensk. Selsk.
3/2:140(1921). Type citation: 'Perth, King’s Park, in open sandy places (No. 874; 10.Sept.l914).'(iso:
MEL 110494).

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Nuytsia Vol. 8, No. 3 (1992)

[Helipterum troedelii var. patens Ewart, J. White & B. Rees, Proc. Roy. Soc. Victoria 22
n.s.: 15(1909). - Helipterum roseum var. patens (Ewart et al.) J. Black, Trans. & Proc. Roy. Soc. South
Australia 45:21(1921) as to syntype 'Fraser Range, 1891, R. Helms', not as to lectotype. Syntype:
FraserRange,4 Oct.l891,y?. Helms (MEL 110705, AD 96343013, NSW 180742).]

The involucral bracts in this subspecies can be pink or white.

Note. Asa Gray, mdar Acroclinium multicaule, described a variety a in which the ray bracts were
white, and a variety 6 in which the ray bracts were 'subaeruginosis'. The lectotype cited above is of
the var. a which corresponds to R. chlorocephala subsp. rosea, var. B is referred to subsp.
chlorocephala.

Rhodanthe chlorocephala subsp. splendida (Hemsley) Paul G. Wilson, subsp. et comb. nov.

Helipterum splendidum Hemsley, Bot. Mag. t.7983(1904). - Helipterum roseum var. album Ewart
et al., Proc. Roy. Soc. Victoria n.s. 23:59(1910) nom.illeg. Type citation: 'Kew is indebted to
Mr G.F. Berthoud, of Waroona, near Drakesbrook, W. Australia, for dried specimens and seeds of
this handsome plant, which he labelled:- "Native of North-western Districts." There are also
herbarium specimens collected in open plains at Menzies, about a hundred miles north of Coolgardie,
in about 29'" 30’S. lat. and 121° E. long., by Cecil Andrews.' Lectotype (here chosen): North-western
Districts of Western Australia, 26.x. 1903, G.F. Berthoud (lecto: K; isolecto: MEL 110704).

Rhodanthe chlorocephala is extremely variable. At one extreme is found in the Shark Bay area
of Western Australia a large flowered variant with long (to 30 mm) cream-coloured limbs to the inner
involucral bracts. At the other extreme is found a variant in the Nullarbor area of South Australia and
Western Australia with small heads and short (c. 5 mm) white limbs. Between these extremes is a
continuity of forms. In addition to the differences in size of the heads and bracts, and the presence
of a black spot at the base of the limb in the large cream bracted variant, there are found colour forms
that range from white to cream and various shades of pink. Hooker noted that in cultivation
Acroclinium roseum had much larger heads than the wild plant from which the seed was obtained and
this difference is apparent when comparing the Drummond herbarium material with Hooker s
illustration.

The variant described by Ostenfeld as Helipterum roseum var. nigropapposum was collected in
Perth; it is almost identical to Drummond’s collection no. 157 (from between the Moore and
Murchison Rivers) on which the name Acroclinium roseum was based.

The name A. multicaule A. Gray was based on two variants, as is noted above, of which one
corresponds to subsp. chlorocephala and the other to subsp. rosea.

The name A. roseum var. grandiflorum Nicholson was based on a large-headed rose-coloured
variant while var. album Nicholson was based on a white variant, both variants are presumably forms
of subsp. roseum.

Ewart et al. (1909) based the name H. troedelii var. patens on two collections of which one, the
lectotype, is Rhodanthe troedelii s.str. while the other (from the Fraser Range in Western Australia)
is of the small 'Nullarbor’ variant of subsp. rosea. It was the latter specimen that J.M. Black first saw

Paul G. Wilson, Heliplerum: Part 1

389

and therefore transferred the variety to H. roseum in the first edition of the Flora of South Australia
(1929).

Rhodanthe collina Paul G. Wilson, sp. nov. (Figure 1)

Annua erecta ad 20 cm alta. Caules graciles, atro-rubri, sparse puberuli et glanduloso punctati.
Folia caulina altcrna. aliquantum coriacea linearia vel anguste oblonga, integra, 8-15 mm longa,
obtusa, margine saepe recurva, supra glabra, infra glanduloso punctata. Capitula heterogamia,
solitaria, terminalia. Involucrum hemispheric urn, c. 5 mm altum, radians. Bracteae multi-seriatae,
glumaceae glabrae; bracteae intimae: ungue late elliptico c. 5 mm longo, stereomate lineari, duro,
lamina oblongo-elliptica c. 6 mm longa, alba. Receptaculum conicum, glabrum. Flosculi numerosi,
bisexuales vel masculi. Corolla actinomorpha, anguste cylindracea, c. 3.5 mm longa; lobi 5,
triangulares, c. 0.4 mm longi; intra dense papillosi. Antherarum loculi c. 1 mm longi; appendix
oblongo-ellipticus, acutus c. 0.4 mm longus, margine minute erosa. cellulis anguste oblongis, tcnui-
parietibus; caudi tenues, debiles. Styli rami apicem versus aliquantum latiores, apice rotundato longe
papilloso, nervo gracili in medio rami terminanti. Achenium obovoideum c. 2 mm longum, atro-
rubiginosum, dense albo-hirsutum, pilis crasso-parietibus; carpophorum annulare; pericarpium
crustaceum; testa membranacea, atro-violaceum, ad pericarpio ± adnatum. Pappus in statu integro
tarde deciduus; setae lineari-acuminatae, breviter plumosae, ciliis terminalibus clavatis.

Typus: Paynes Find - Thundelarra road near Monger Lake, 29° 03' S, 1 17° 17' E, low hill of quartzite
boulders, 27 Sept. 1986, Paul G . Wilson 1 2300 (holo; PERTH; iso: AD, BRI, CANB , K, MEL, NSW).

Annual erect herb to 20 cm high branching at and above base. Major axes slender, dull reddish
brown with a few small septate hairs and sessile globular glands. Leaves cauline, alternate, somewhat
leathery, linear to narrow-oblong or spatulate, entire, 8 x 0.5 - 15 x 2 mm, obtuse, margin often
recurved (at least when dry), glabrous above, spar.sely covered below with sessile globular glands and
with a few .shorter septate hairs on midrib. Capitula hctcrogamous, solitary on slender leafless
terminal peduncles. Involucre hemispherical, c. 5 mm high, radiant; bracts glumaceous, glabrous;
outer and intermediate bracts elliptic, to 6 mm long, pale brown, with a short flat stereome; inner
bracts: claw broad-elliptic, hyaline, c. 5 mm long with a hard linear stereome c. 2/3 its length; lamina
oblong-elliptic, obtuse, c. 6 mm long, 2.5 mm wide, white. Receptacle conical, glabrous. Florets
numerous, the outer bisexual and the inner male. Corolla narrow-tubular, somewhat constricted
below middle, c. 3 .5 mm high, yellow, sparsely puberulous towards base, wall thickened in lower half,
vascular strandsextending to base of lobes; lobes 5, short erect, triangular, c. 0.4 mm long, all densely
papillose within. Stamens: anther loculi c. 1 mm long; anther appendage oblong-elliptic, acute, c.
0.4 mm long, minutely eroseon margin, cells narrow-oblong, walls unthickened except fora marginal
row of small cells with thickened walls; anther tails very thin and delicate. Style branches broadened
towards tip and with a rounded long-papillose apex; vascular strand very slender and terminating
about half way along style branch. Achene obovoid, c. 2 mm long, very dark reddish brown, densely
white-hirsute with thich-walled hairs, myxogenic; carpopodium annular; pericarp crustaceous with
lignified reddish brown radial walls: testavery thin and weak, dark mauve, with scattered flat crystals,
± fused to pericarp. Pappus tardily deciduous as a whole; bristles linear-acuminate. ± equal to corolla,
united in a short ring at base, shortly plumose, the terminal cilia clustered and clavate.

Distribution. Near Yalgoo, central western Western Australia, in the far western area of the Austin
Botanical District.

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NuytsiaVoI. 8, No. 3 (1992)

Figure 1. Rhodanihe collina. A - Habit. B - Undersurface of leaf. C - Multicellular hispidulous and globular glandular
trichomes from undersurface of leaf. D - Capitulum. E - Inner, intennediate, and outer involucral bracts. F - Receptacle.
G - Floret. H - Corolla. I - Ii\ner surface of apex of corolla. J - Apex of pappus bristle. K- Anther with enlargement of margin
of appendix (tails drawn recurved for space exigencies). L - Apex of style with two papillae enlarged. M - Achene with apex
of duplex hair enlarged. N - Achene showing position of seed and of Us vascular strand. O - Enlargement of epidennis of
pericarp. P • Enlargement of epidennis of testa showing crystals. From B.H. Smith 460.

Paul G. Wilson, Heliplerum: Part 1

391

Specimens examined. WESTERN AUSTRALIA: 112milepcgonlhe YalgooRoad,A.M.A^/i^))'2976
(AD, PERTH); Ml Gibson, B.H. Smith 460 (MEL, PERTH); Mingenew Hill, 29 Aug.1957,
R.F. Watson (PERTH).

Habitat. On rocky bilks in exposed situations.

Notes. This species would appear to have no close relatives although in habit and in indumentum it
is similar to Rhodanthe diffusa. The unique anther appendages, style apices, and achenes combine
to make it a very distinct species.

Rhodanthe corynibillora (Schldl.) Paul G. Wilson, comb. nov.

Helipterum corymbiflonim Schldl., Linnaea 21:448(1848); Sender, Linnaea 25;519(1853). -
Argyrocome corymbiflora (Schldl.) Kuntze, Revis. Gen. PI. 1:309(1891). - Roccardia corymbiflora
(Schldl.) Voss, Vilm. Blumengartn. 3rdedn, 1:532(1895). Type citation: 'DrBehr’ [South Australia]
(?holo: MEL 604825).

Note. Sender, /.c., cited the Behr collection as coming from 'Fiedler’ s section, Novemb.' This locality
is in the Hundred of Moorooroo, County Light, near the present town of Nuriootpa (South Australia).
The putative type sheet in herb. MEL has on it three specimens, two of which arc somewhat etiolated
whole plants while one is a single stem off a mature plant. A note on the sheet by Sonder states 'culmi
e seminibus plantae behrianae! ' It would appear likely that the two whole plants represent those raised
by Sonder while the single stem is the type specimen collected by Behr since it corresponds to the
following comment made by Schlechtendal, /.c., 'Unicum specimen vidi, ramum 5 poll, longum'.

Rhodanthe cremea Paul G. Wilson, sp. nov. (Figure 2)

Annua erecta, c. 25 cm alta, sparse lanosa, pilis glandulosis absentibus. Axes majores graciles,
solitarii vel numerosi, baud ramosi, apicem versus lanosi aliter glabri. Folia caulina alterna, Integra,
linearia vel anguste obovata, acuta vel obtusa, ad 30 mm longa, apicem versus caulorum absentia.
Capitula solitaria ad caules terminalia posita. Bracteae exteriores scariosae vel chartaceae, latissimc
ovatae vel latissime ellipticae, nilidae, ad 7 mm longae, pallido brunneae; bracteae intermediae late
ellipticae, ad 15 mm longae, ad apicem malvinae; bracteae intimae: unguis late obovatus, stereomate
oblongo, vix incrassato, nervo singular! ad apicem extenso; lamina petaloidea, anguste elliptica, ad
20 mm longa, pallido cremea vel ad basim alropurpurea. Rcceptaculum discoideum, margine areolae
pilosa aliter glabrum. Flosculi numerosi, biscxuales vel flosculis intimis masculinis. Corolla: tubus
anguste cylindraccus, c. 2 mm longus; faux valde zygomorpha anguste turbinata, c. 1 mm longa; lobi
5, c. 1 .5 mm longi, lobis tribus abaxialis recurvis, cremeis, profunde fissa, intra laevibus, lobis duobus
adaxiali ercctis,ad apicem connatis,rubiginosis, intra papillosis.nerviscorollae ad apicem lobulorum
extensis. Antherarum appendix stcrilis late elliptica, obtusa, 0.2 mm longa; cellulae oblongae,
parietibus tenuibus, cellulis parietibus incrassatis immixtis; cellulae marginales manifeste discretae.
Antherarum caudac filamcniosae, debiles. Styli apex truncatus. nervo prom inenti ad apicem extenso.
Achenium compre.sse turbinatum, c. 2.5 mm longum, facie abaxiali scriceum, adaxiali hirsutum;
carpophorum absens: pericarpium chartaceum; testa mcmbranacea ad pcricarpium adnata, nervo in
positione medio cingenti. Pappi setae corollam leviter excedentes; in parte iuferiore persistentes,
anguste ellipticae; in parte superiore caducae, filamentosae, apice versus plumosae, ad extremum
penicillatae ciliis clavatis ornatis, aliter denticulatae.

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NuytsiaVol.8, No. 3 (1992)

Typus: Western Australia, 6.5 km east of the turnoff to Useless Loop; abundant on flats, dominating
astony c\aydepTQSs\onwith Acaciatetragonophyllaand Acacia sp., 16.viii.l986,f?.C. Chinnock6160
(holo: AD; iso: K, MEL, PERTH).

Erect annual to 25 cm high, branching at base; major axes slender, solitary or several, simple,
woolly near apex otherwise glabrous, glandular hairs absent. Leaves cauline, alternate, entire, linear
to narrow-obovate, acute to obtuse, to 30 mm long, absent towards apex of stem. Capitula solitary,
terminal to stems. Involucre hemispherical, c. 6 mm high. Bracts multiseriate, scarious or papery,
glabrous; outer bracts scarious, very broadly ovale or broadly elliptic, to 7 mm long, pale brown;
intermediate bracts broad-elliptic, to 15 mm long, pale brown with mauve apex; inner bracts: claw
broad-obovate, stereome broad-oblong, scarcely lhickened,with a single nerve extending to apex,
lamina petaloid, narrow-elliptic, to 20 mm long, pale cream, often with a dark purple band at base.
Receptacle disc-shaped, c. 5 mm broad, pilose on margin of areolae otherwise glabrous. Florets
numerous, bisexual or the inner ones male. Corolla almost glabrous: tube cylindrical, c. 2 mm long;
throat narrow-turbinate, strongly zygomorphic, c. 1 mm long; lobes 5, c. 1.5 mm long, the three
abaxial lobes deeply divided, recurved, cream coloured, smooth within, the 2 adaxial lobes fused to
near apex, dark reddish brown, papillose within; vascular strands extending to tip of lobes. Anther
appendage broad-elliptic, obtuse, 0.2 mm long; cells oblong, thick-walled cells interspersed among
thin-walled cells; marginal cell row clearly demarcated; anther tails filamentous and weak. Style tip
truncate, the vascular strand stout and extending to apex. Achene compressed turbinate, c. 2.5 mm
long, silky on abaxial face and hirsute on adaxial face; carpopodium absent; pericarp papery; testa
membranous, fused to pericarp, the vascular strand encircling seed in medial position. Pappus bristles
slightly exceeding corolla; lower half narrow elliptic, barbellate and persistent; upper half filamentous
and deciduous, towards apex plumose with the tenninal cilia penicillate and clavate, towards base
barbellate.

Distribution. Western Australia, from the south end of Hamelin Pool, Shark Bay, eastwards to the
North West Coastal Highway.

Specimens seen. WESTERN AUSTRALIA: 493 mile peg North West Highway, T.E.H. Aplin 3282
(PERTH); 5 miles west of Hamelin Station, J.W. Green 1444 (PERTH); 22 km west of Ovcrlander
Roadhouse on Denham road, N.S. Lander 1322 (PERTH); 28 km NNW of Overlander Roadhouse,
North West Coastal Highway, E.N.S. Jackson 3121 (AD pro parte)’, 3 km from Overlander
Roadhouse, P.S. Short 422 (AD pro parte)’, 20 km north of Billabong Roadhouse, P.S. Short 2453
(PERTH).

Habitat. Usually found in red sand over loam.

This species appears to be related to Rhodanthe chlorocephala. It superficially very closely
resembles /?. chlorocephala subsp. splendida which grows intermixed with R. cremea near Shark Bay
and some herbarium collections consist of material of both species. Rhodanthe chlorocephala may
be distinguished by the spherical glandular hairs on leaves and stems, the absence of woolly hairs,
the regular corollas, and the evenly silky achenes.

Those variants of Cephalipterum drummondii and of R. chlorocephala subsp. splendida that have
large white involucral bracts often resemble R. cremea in having a dark purple band at the base of
these bracts. Since the three species grow together in the Hamelin Pool region it is likely that they
attract the same pollen vectors.

The epithet cremea (cream coloured) refers to the colour of the inner involucral bracts.

Paul G. Wilson, Helipierum: Part 1

393

0-2 mm 0.5 mm

Figure 2. Rhodanthe cremea. A - Habit. B - Capituluni. C - Corolla opened out. D - Bisexual floret. E - Corolla. F- Male
floret. G - Receptacle. H - Pappus bristles. I - Style branches. J - Anther. K - Leaf. L - Leaf hairs. M - Outer and inner
involucral bracts. N- Achene. O - T.S. achene. From 7.VL. 1444.

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Rhodanthe diffusa (Cunn. ex DC.) Paul G. Wilson, comb. nov.

Helipterum dijfusum A.Cunn. ex DC., Prod. 6:216(1838). Type: Forest grounds at the base of Peels
range. Interior of New South Wales, June 1817, A. Cunningham (holo: G-DC photo seen).

Rhodanthe diffusa (DC.) Paul G. Wilson subsp. diffusa

This taxon represents the yellow-rayed variant.

Rhodanthe diffusa subsp. leucactina (F. Muell.) Paul G. Wilson, comb. nov.

Helipterum polygalifolium var. leucactinum F. Muell., Fragm. 10:107(1877). Type citation:
'Darling’s River (Andrae),Warrego (Bailey), Curruwillughi (Dalton), Ballandool (Looker).' Lectotype
(here chosen): Warrego district, F.M. Bailey (MEL 1 10257).

The taxon represents the white-rayed variant.

Note. The only substantial difference between the two subspecies is in the colour of the radiant
involucral bracts, however the stems of northern variant of subsp. leucactina are sometimes branched
above and have a minutely glandular hispidulous indumentum beneath the capitula rather than
cottony. The typical subspecies is found in south-eastern New South Wales between Hillston and
Narrandera while the subsp. leucactina is found further north from Nyngan north to south-eastern
Queensland. Both subspecies have been confused with other species: the yellow subspecies was
synonymized by Bentham (1867) (and subsequently confused in some herbaria) with Helipterum
polygalifolium, while the white has been confused with H. anthemoides. Rhodanthe polygalifolia
may be readily distinguished by being glabrous apart from the minute globular glands on the adaxial
surface of the leaves (in . diffusa the leaves bear glands and multiseptate hairs on the stems and on
both surfaces of the leaves), while /?. anthemoides may be distinguished by being a perennial, by the
generally thicker leaves and embedded globular glands, and by the reddish brown linear midrib on
the claws of the involucral bracts.

Rhodanthe fuscescens (Turez.) Paul G. Wilson, comb. nov.

Helipterum fuscescens Turez., Bull. Soc. Imp. Naturalistes Moscou 24/2:80(1851). - Podotheca
fuscescens (Turez.) Benth., FI. Austral. 3:602(1867). - Podosperma fuscescens (Turez.) F. Muell,
Fragm. 12:22(1^82). - Argyrocome fuscescens (Turez.) Kuntze, Revis. Gen. PI. 1:309(1891). Type:
Western Australia,/. Drummond 5th coll, (suppl.) n.64 (holo: KW photo seen; iso: K, MEL 1 10215,
PERTH).

Acrocliniumphyllocephalum A. Gray, Hooker’s J. Bot. Kew G;ud. Misc.4:27 1(1852). Type citation:
'South-west Australia, Dnww/o/i/ (received in 1850)' n.v.

This species is very similar to Rhodanthe oppositifolia from which it differs in having leaves that
subtend the capitula and in having very short and brown involucral bracts. Rhodanthe fuscescens is
known only from the Drummond types and from E. Pritzel 695. The Drummond sheets on which the
names H. fuscescens and A . phyllocephalum were based are probably duplicates of the same collection
(Short 1983); both sheets lack locality data. The collection Pritzel 695, dated 1901, is almost identical
to the Drummond; it came from near Cranbrook in south-west Western Australia which is about 400
km from the nearest locality for R. oppositifolia.

Paul G. Wilson, Ileliplerum: Part 1

395

Rhodanthe gossypina Paul G. Wilson, sp. nov.

Herba annua erecta gossypina. Folia caulina. alterna, linearia vel anguste oblonga, acuminata,
2-4 cm longa. Capitula homogama in corymbis tcrminalibus ordinata. Involucrum turbinatum, c. 4
mm altum, radians; bracteae c. 3-seriata; bracteae intermediae ellipticae c. 5 mm longae, hyalinae,
lanato ciliatae, ad apice brunneolae, stereomate anguste oblongo, viridi, in parte superiore gossypino;
bracteae intimae; unguis late obovatus c. 4 mm longus, hyalinis, gossypino ciliatus, .stereomate
anguste oblongo, gossypino, in parte superiore pilis resinosis rubiginosis omatum; lamina patula,
ovata, c. 4 mm longa, alba. Receptaculum conicum, c. 0.7 mm diam. Flosculi c. 15, bisexuales.
Corolla c. 3 mm longa, 5-lobata; lobi inequales, oblongo-triangulares, intra Faeves, lobo abaxiali c.
1.5 mm longo, ceteris c. 1 mm longis; nervi corollae in tubo terminantes. Antherae; appendix sterilis
elliptica c. 0.4 mm longa. parietibus cellularum nec incrassatis; caudis filamentis, debiles. Styli apex
± truncatus. Achenium ellipsoideum, 1 .5-3 mm longum, subtiliter pilosum; carpophorum annulare;
pericarpium cbartaceum, translucens; tesla aliquanlum coriacca, brunnea, ad pericarpium adnata,
nervo c. 2/3 cingenti. Pappus pcrsistens; setae lineari-acuminatae corollam aequantes, omnino
pilosae.

Typus: Queensland: Mitchell district, 36.9 km SE of Winton on Longreach road, 19 Sept. 1984,
RJ. Chinnock 6106 (holo: AD; iso: PERTH).

Annual erect cottony herb; stems slender, single or branching at base, to 20 cm high. Leaves
cauline, alternate, linear to narrow-oblong, acuminate, 2-4 cm long, 1-5 mm wide. Capitula
homogamous, in tenninal corymbs. Involucre turbinate, c. 4 m high, radiant; bracts c. 3-scriate;
intermediate bracts elliptic, c. 5 mm long, hyaline, woolly ciliate, brown-tinged towards apex, with
narrow-oblong green stereome cottony in upper half, c. 2 mm long; innermost bracts: claw
broad- obovate, c. 3.7 mm long, 2 mm wide, hyaline, cottony ciliate, stereome narrow-oblong
extending to near apex, cottony and with dark reddish brown resinous hairs in upper half; lamina
spreading, ovate, c. 4 mm long, 2 mm wide, white. Receptacle conical, c. 0.7 mm diameter Florets
c. 15, all bisexual. Corollae. 3 mm long, narrow-cylindrical below, somewhat broadcrabove.sparsely
puberulous in lower half, yellow, vascular strands terminating well below lobes; lobes oblong-
triangular, 0.5-1 mm long, not papillose within, glabrous outside, the abaxial lobe more deeply
divided (up to 1 .5 mm long). Anthers: appendage elliptic, c. 0.4 mm long, smooth, cells not thickened,
very pale brown or the marginal ones clear; tails filamentous, weak. Style apex ± truncate to slightly
rounded, shortly papillose. Achene ellipsoidal, 1 .5-3 mm long, pilose with fine very pale brown hairs;
carpopodium a short glabrous ring: pericarp translucent, papery, very pale brown; testa somewhat
leathery, brown, fused to pericarp when mature: va.scul:ir strand of testa extending c. 2/3 around seed
in medial position. Pappus bristles linear-acuminate, ± equal to corolla, united in a ring at base,
persistent, pilose throughout, the terminal cilia somewhat thicker and rounded at apex.

Distribution. Southeastern Northern Territory, southwestern Queensland, extreme northeastern
South Australia and northwestern New South Wales.

Specimens examined (selection only). QUEENSLAND: 100 km WNW of Charleville, C. Sandercoe
369 (BRI); Near Windorah, S.T, Blake 12079 (BRI).

NEW SOUTH WALES: 17.5 km from Louth on Telpa road, C.W.E. Moore 7794 (CANB).
SOUTH AUSTRALIA: Kudriemitchie Waterhole, L.D.Williams 8156 (AD).

NORTHERN TERRITORY: 7.5 mi N of Tobermorey homestead, G. Chippendale (NT 2961).

Habitat. In open situations on heavy soil that is frequently subject to flooding.

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Flowering time. June to September.

This species is similar to Rhodanthe corymbiflora and R. microglossa with which two species it
has been confused in herbaria. It differs from the former in having a smaller involucre with the bract-
lamina about half as long, and in the possession of dark resinous hairs on the stereome of the inner
bracts. From the latter it differs in having a more open inflorescence, dark resinous hairs on the inner
bracts, much longer radiating laminae, and in having a smooth throat to the corolla.

Rhodanthe gossypina was recognised as a distinct taxon by C.T. White (herb. BRI) who evidently
intended to describe it as a variety of Helipterum corymbiflorum but never did so (R.W. Johnson,
pers. comm.).

The solitary collection from New South Wales, cited above, comes from a site geographically
isolated from the other locations. It is morphologically distinct for the stereomes of the inner
involucral bracts do not have glandular hairs and the lobes of the corolla are equal in length. For these
reasons it may be found to represent a distinct species.

The specific epithet refers to the cottony indumentum that covers the plant.

Rhodanthe humboldtiana (Gaudich.) Paul G. Wilson comb, nov,

Helichrysum humboldtianmi Gaudich. in Freyc., Voy. Uranie 465(1830) t.88(1829). - Helipterum
humboldtianum (Gaudich.) DC., Prod. 6:216(1838). - Schoenia humboldtiana (Gaudich.) Walpers,
Repert. Bot. Syst. 6:244(1846). - Pteropogon humboldtianus (Gaudich.) F. Muell., Linnaea
25:415(1853). - Argyrocome humboldtiana (Gaudich.) Kuntze, Revis. Gen. PI, 1:309(1891). -
Roccardia humboldtiana (Gaudich.) Voss, Vilm. Blumengartn. 3rd edn, 1 :532( 1 896). Type citation:
'In Novae-Hollandiae oraoccidentale (baiedes Chiens-Marins)', 1830, Gaudichaud. Lectotype (here
chosen): P photo seen; isolecto: G-DC photo seen.

HelipterumsandfordiiHook.,Bol. Mag. t.5350(1862). Type:J. Drummond 160, between Moore and
Murchison rivers; 1860,RM/ gm; cult. Thompson from seeds from Major Sandford,W. A./iOiyn/ypci'.-
J. Drummond 160 (MEL 109493, NSW).

Helipterum largiflorens F. Muell., Fragm. 3:135(1863). Type citation: 'Ad sinum Champion bay et
flumen Murchison. Walcott et Oldfield.' Lectotype (here chosen): Champion Bay, A. Oldfield (MEL
109501).

Helipterum clementii Domin, Biblioth. Bot. 89:667(1929). Type citation: 'Nordwest-Australien:
zwischen Ashburton- und De Gray River, E. Clement'. Type n.v.

Rhodanthe microglossa (Maiden & E. Betche) Paul G. Wilson, comb. nov.

Helipterum microglossum Maiden & E. Betche, Proc. Linn. Soc. New South Wales 22:151(1897).
Type: Tarella, New South Wales, Aug. 1887, W. Bduerlen 131 (holo: NSW 181467).

Helipterum coiymbiflorum var. microglossa F. Muell. ex Benth., FI. Austral. 3:647(1867). -
H. microglossa (Benth.) Tate, Trans & Proc. Roy. Soc. South Australia 22:121(1898). Lectotype
(here chosen): Goyinga mountains, 6 & 7 Nov. 1860, Victorian Expedition (MEL 110255).

Paul G. Wilson, Helipterum: Pait 1

397

Note. It was unclear from the paper by Maiden and Betche whether they intended to describe a new
species based on the Bauerlen collection or whether they intended to make a new combination based
on Helipterum corymbiflorum var. microglossa\ I have assumed the former intention, partly because
the epithet was changed from a substantive to an adjective and partly because the format used was
the same as that for other new species described in the paper. Ralph Tate (1898 loc.cit.) claimed to
have published the species combination in the Trans. &Proc.Roy. Soc. South Australia 6; 104(1883),
however, in that paper the name Helipterum microglossa was not validly published since it appeared
without citation of either author or basionym, and without a description.

Rhodanthe oppositifolia (S. Moore) Paul G. Wilson, comb. nov.

Helipterum oppositifolium S. Moore, J. Bot. 35165(1897). Type: Near Coolgardie, Sept. 1895,
S. Moore (holo: BM; iso: MEL 110309).

Grijfithia helipteroides J. Black, Trans. & Proc. Roy. Soc. South Australia 37:122(1913). Type
citation: 'Gawler Ranges, Eyre Peninsula {S.A.White, September, 1912)' (holo: AD; iso: K),

subsp. oppositifolia

subsp. ornata Paul G. Wilson, subsp. nov.

Folia lineari-acuminata, c. 30 x 2 mm. Involucrum hemisphcricum, c. 6 mm altum; lamina
bracteae interiorum elliptica, 10 mm longa, flava.

Typus: Western Australia: 3 km NW of Tamala Homestead, 26° 40' S, 1 13° 42' E, at foot of calcrete
ridge on edge of samphire flat, powdery white clay loam on sheet calcrete, 25 August 1991,
S.D. Hopper 8112 (holo: PERTH; iso: CANB, K, MEL, S).

Leaves linear-acuminate, c. 30 x 2 mm. Involucre hemispherical, c. 6 mm high; inner bracts with
a prominent elliptic yellow lamina c. 10 mm long.

Distribution. Only known from the southern margin of Freycinet Estuary in Tamala station. Western
Australia. Found growing on calcrete rises on edge of saline flats.

Additional specimens seen. WESTERN AUSTRALIA; 13.5 km W of Tamala homestead road on
Useless Loop road. S.D. Hopper ?,l] 3 (PERTH); Tamala Station, 20 July 1988, Ph. Moral 8212 (P).

This subspecies differs from the typical in having larger capitula and much larger lamina to the
inner involucral bracts which in the typical subspecies are only 2-3 mm long. Subsp. oppositifolia
is also found in the Shark Bay region but evidently not in association with subsp. ornata.

Rhodanthe polygalifoliu (A.Cunn. ex DC.) Paul G. Wilson, comb. nov.

Helipterum polygalifolium A.Cunn. ex DC., Prod. 6;216(1838). - Argyrocome polygalifolia (DC.)
Kuntze, Revis. Gen. PI. 1:309(1891). Type.- Molle’s Plains, Lachlan river. New South Wales, 1817,
A. Cunningham (holo: G-DC photo seen).

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Nuytsia Vol. 8, No. 3 (1992)

Rhodanthe propinqua (W. Fitzg.) Paul G. Wilson, comb. nov.

Heliplerum propinquum W. Fitzg., J. W. Austral. Nat. Hist. Soc. No. 1 :24(May 1 904). Lectotype (here
chosen); Nannine, Western Australia, Sept. 1903, W.V. Fitzgerald (NSW 181422; isolecto; NSW
181423, PERTH).

Rhodanthe pygmaea (DC.) Paul G. Wilson, comb. nov.

Pteropogon pygmaeus DC., Prod. 6:245(1838). - Pteropogon australis Nees, Linnaea 16:223(1842)
nom. illeg., based on above. - Helipterum pygmaeum (DC.) F. Muell., Rep. Babbage’s Exped.
14(1859); Benth., FI. Austral. 3;647(1867). Type: Mode’s plains, Lachlan river. Interior west from
Port Jackson, Long. 145°E, 'aremarkablepigmy plant’, July 1817, A. Cunningham 101 (holo:G-DC
photo seen).

Pteropogon drummondii A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4;267(1852). - Helipterum
pygmaeumvsr. occidentale Benth., FI. Austral. 3:647(1867). -Helipterumpygmaeum\es. drummondii
(A. Gray) Ostenfeld, Dansk Bot. Ark. 3:142(1921) nom. illeg. - Helipterum drummondii (A. Gray)
Ostenfeld, l.c. Type: Western Australia, 1849. J. Drummond (holo: K).

Pteropogon intermedius F.Muell., Linnaea 25:411(1853). Type citation: 'In collibus graminosis
montem Rufus versus, ante Torrens tenus, ubertim rarus.' Lectotype (here chosen): Ad rivum
Broughton, Oct. 1851, F. Mueller (MEL 604821).

Rhodanthe rubella (A. Gray) Paul G. Wilson, comb. nov.

AcrocliniumrubellumA. Gray, Hooker’s J. Bot. Ke.wGard.Misc.4:ni(lS52).-Heliplerumrubellum
(A. Gray) Benth., FI. Austral. 3:641(1867). - Argyrocome rubella (A. Gray) Kuntze, Revis. Gen. PI.
1:309(1891). -Roccfl/ c/m rubella (A. Gray) Voss, Vilm. Blumengartn. 3rd edn, 1:530(1896). Type
citation: 'Between Swan River and King George Sound, Drummond.'Lectotype (here chosen):
J. Drummond 347 (K, photo seen).

Rhodanthe rufescens Paul G. Wilson, sp. nov.

Herba annua decumbens, spimse et minute glanduloso-puberula; axes majores numerosi ad 10 cm
longi. Folia caulina alterna sessilia, anguste oblonga, ad 10 mm longa, integra, obtusa. Capitula
homogama, ad ramulos terminalia posita, nec radiantia. Involucrum breviter cylindraceum, 4-5 mm
altum. Bracteae c. 3-seriatae, scariosae, supra stereoma sparsissime et minute glanduloso puberulae
aliter glabrae; bracteae exteriores ovatae, c. 2 mm longae, stramineae; bracteae interiores obovatae,
ad 5 mm longae, ad apicem rotundatae, versus basim stramineae, versus apicem violaccae vel margine
apicali hyalino. Receptaculum rotundatum c. 0.7 mm diam., glabrum. Flosculi c. 7, bisexuales,
actinomorphi. Corolla c. 3.5 mm longa, infra medium cylindracea, supra medium anguste cupulata;
lobi 5, triangulares, c. 1 mm longi, intraad basim papillosi; nervi ad basim loborum extensi. Antherae
loculis c. 1 mm longis; appendix c. 0.3 mm longus ad apicem rotundatus, cellulis oblongis nec
incrassatis; caudae filamentac, debiles. Styli apex truncatus, nervo ad basim apicis extenso.
Achenium (in statu immaturo) sericeum, carpophore absenti, pcricarpio tenui, translucenti, testa
tenuiter coriacea. Pappi setae lineares acuminatae, c. 2 mm longae, omnino plumosae, ciliis omnino
acuminatis.

Paul G. Wilson, Heliplerum: Part 1

399

Typus: Queensland, Gregory South District, Noccundra, about 130 km west-north-west of
Thargomindah, 'herbage', 11 Aug. 19^1 ,P.Warhursi s.n. (holo: BRI 410267).

Annual (? decumbent) herb with many somewhat wiry stems to 10 cm long arising from base.
Branches and leaves sparsely covered with minute sessile or shortly stipitate globular glands,
otherwise glabrous. Leaves cauline, alternate, sessile, narrow-oblong, to 10 mm long, 1-2 mm wide,
entire, obtuse. Capitula homogamous, terminal to slender lateral and main branches, sometimes
subtended by a foliaceous bract. Involucre shortly cylindrical, 4-5 mm high, not radiant; bracts
scarious, glabrous apart from a few minute glandular hairs on stereome,c. 3-seriate; outer bracts ovate
c. 2 mm long, straw-coloured, glabrous; inner bracts obovate, to 5 mm long, rounded at apex, straw-
coloured towards base, violei brown towards apex or with the tip hyaline, stereome oblong, 2/3 length
of bract, flat, pale green. Receptacle rounded, c. 0.7 mm diameter, glabrous. Florets c. 7, all
apparently bisexual. Corolla tube c. 3. 5 mm high, cylindrical below, narrowly cup-shaped above, very
sparsely puberulous, yellow; lobes 5, ovate, triangular, c. 1 mm long,papillose within in abroad ring
around base of lobes; vascular strands extending to base of lobes. Stamens: anther appendage c. 0.3
mm long rounded at apex, cells oblong, not thickened; anther tails weak and filamentous. Style tip
truncate, papillose; va.scular strand prominent and extending to ba.se of tip. Achene (not seen mature)
silky villous all over; carpopodium absent; pericarp thin, translucent; testa thinly coriaceous, with
crystals. Pappus bristles linear-acuminate, c. 2 mm long at anthesis, plumose to tip, tenninal cilia
not thickened.

Distribution. Only known from the type locality in south-west Queensland.

The specific epithet refers to the reddish colour of the terminal portion of the inner involucral
bracts.

Rhodanthe stricta (Lindley) Paul G. Wilson, comb. nov.

Xyridanthe stricta Lindley, Sketch Veg. Swan Riv. Col. 23(1839). - Helipterum strictum (Lindley)
Benth., FI. Austral. 3:646(1867), Type: Swan River, 1839,7. Drummond (holo: CGE photo seen).

Pteropogon platyphyllus F. Muell., Linnaea 25:413(1853). Type citation: 'Ad margines rupestres
rivulorum ad Cudnaka rariusve ibidem in planitiebus sterilibus.' Lectotype (here chosen): 'Ad
marginis rupestris rivi Cudnaka', Oct. 185 1, F. Mueller (lecto: MEL 1 10685; isolecto: MEL 604829).

Helipterumstriclumvtu.stenocephalaS. Moore, J. Linn. Soc. 34:200(1899). Type.- Near Coolgardie,
August 1895, S. Moore (holo: BM photo seen).

4. Rhodanthe sect. Achyroclinoides (A. Gray) Paul G. Wilson, comb. nov.

Pteropogon sect. Ac/;yroc/i>ioi7er A. Gray, Hooker’s J.Bot.KewGard.Misc. 4:268(1852). Lectotype
(here designated): Pteropogon corymbosus A. Gray.

[Cassinia sect. Cassiniola E. Muell., Fragm. 3:139(1863) nomen sub Cassinia cuprea F. Muell.
(= Pteropogon polycephalus A. Gray)].

Plant variously pubescent to sub-glabrous, without sessile glands on leaves. Style apex truncate.
Achene small, 1-1.5 mm long, moderately to sparsely short hirsute; carpopodium annular, glabrous;

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Nuytsia Vol. 8, No. 3 (1992)

pericarp crustaceous, thin, brown, the radial cell walls thickened; testa membranous, free or fused to
pericarp. Pappus caducous.

Note. Asa Gray (1852) included Pteropogon polycephalus A. Gray, P. corymbosus A. Gray and
P. laevis A. Gray in his section AchyrocUnoides. These three species are all recognised here as being
members of this section.

Rhodanthe ascendens Paul G. Wilson, sp. nov.

Herba annua ascendens ad 10 cm alia, modice lanosa. Folia caulina, alterna, anguste obovata,
plana, 10-15 mm longa, c. 2 mm lata, sessilia. Inflorcscentia corymbiformia, compacta. Capitula
homogama, breviter pedunculata. Involucrum cupulalum c. 5 mm allum, 2.5 mm latum, nec radians;
bracteae glumaceae, 3-4-serialae, nitidae, bracteis intermcdiis interioribusque ellipticis, 4-5 mm
longis, ad apicem rotundatis, ad basim sparse ciliatis aliter glabris. Receptaculum convexum,
glabrum, c. 1 mm latum, bracteis receptaculi absentibus. Flosculi c. 20, bisexuales, actinomorphi.
Corolla c. 3 mm longa, glabra, nervis ad apicem tubi extensis; lobi deltoidei, c. 0.5 mm longi, lobis
tribus parte interiore ad basim papillosis, ceteris glabris. Anthcrae c. 1 mm longae; apex anguste
ovatus obtusus, c. 0.3 mm longus, cellulis anguste oblongis; caudae filamentae, debiles. Styli apex
truncatus,nervoprominentiadapicemextenso. Achenium, in statu immaturo,angustecylindraceum,
1 mm longum, breviter hirtellum; carpophorum annulare, glabrum; pericarpium tenue, translucens;
testa membranacea. Pappus corollam aequans, in statu integro caducae; setae c. 15, filiformae, ad
basem breviter connatae, barbellatae.

TypMj.-Western Australia, Gascoyne Junction; florets yellow; roadside verge, clay soil; 20 Aug. 1986,
P.S. Short 2531 (holo; PERTH; iso: MEL n.v.).

Annual ascendent herb to 10 cm high, branching at base, moderately and loosely woolly. Leaves
sessile,alternate,cauline, narrow obovate, obtuse, flat, 10- 15mmlong,c.2mmwide. Inflorescence
corymbiform, compact. Capitula homogamous, very shortly pedunculate. Involucre cup-shaped, c.
5 mm high, not radiant; bracts glumaceous. 3-4-seriate, glossy, intermediate and inner bracts elliptic,
to 4.5 mm long, rounded at apex, sparsely ciliate at base otherwise glabrous. Receptacle convex,
glabrous, without receptacular bracts, c. 1 mm diameter. Florets c. 20, bisexual, actinomorphic.
Corolla c. 3 mm long, yellow, narrow-tubular in lower half, narrow-turbinate above, glabrous,
vascular strands extending to apex of tube; lobes broad-triangular, c. 0.5 mm long, three of the lobes
papillose within at base the others glabrous; cells of throat strongly undulate. Stamens: anther loculi
c. 1 mm long; tip oblong-ovate, obtuse, c. 0.3 mm long, the cells narrow-oblong; tails filamentous,
weak. Style arms truncate, papillose, the vascular strand prominent and extending to base of tip.
Achene (in immature state) narrow-cylindrical, 1.0 mm long, shortly hirtellous with duplex hairs c.
0.1 mm long, bifid at tip, myxogenic; carpopodium a narrow glabrous ring; pericarp thin and
translucent, weak; testa membranous, free from pericarp, vascular strand extending to apex of seed;
endosperm persistent and forming a sheath around the embryo. Pappus bristles c. 15, ± equal to
corolla, very shortly united in a ring at base, barbellate throughout, the terminal teeth rounded,
caducous as a whole.

Distribution. Found only in the Carnarvon Botanical District (Beard 1980) near Gascoyne Junction,
Western Australia.

Additional specimen examined. WESTERN AUSTRALIA: Middalya Station, 19S9, D.C.Norbury 1
(PERTH).

Paul G. Wilson, Helipterum: Part 1

401

Habitat. The type was found in clay soil on a roadside verge.

Rhodanthe ascendens is very similar to/?, nullarborensis from which it differs most obviously in
having broader leaves and larger capitula. In the details of the florets it differs in having barbellate
(not shortly plumose) pappus bristles and in having three (not two) of the corolla lobes papillose at
their base, but this latter character may be a variable one. The achene of R. nullarborensis has a thin
crustaceous pericarp with lignificd radial walls while in its immature state the pericarp ofR. ascendens
shows no sign of such thickening but this may develop later.

Rhodanthe condensata (F. Muell.) Paul G. Wilson, comb. nov.

Helipterum condensatum F. Muell., Fragm. 3:136(1863). - Argyrocome condensata (F. Muell.)
Kuntze, Revis. Gen. PI. 1:309(1891). Type: Murchison River, Western Australia, A. Oldfield (holo:
MEL 1 10770).

Rhodanthe corymbosa (A. Gray) Paul G. Wilson, comb. nov.

Pteropogon corymbosus A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:268(1852). - Helipterum
corymbosum (A. Gray) BenIh.,Fl. Austral. 3:649(1867). - Argyrocome corymbosa (A. Gray) Kuntze,
Revis. Gen. PI. 1:309(1891). Type citation: 'Swan river, Drummond. - Darling Range, South-west
Australia, Collie.' Isosyntype: J. Drummond 364 (MEL 109605).

Helipterum album Ewart, J. & Proc. Roy. Soc. New South Wales 42:189(1909). Type: Wooroloo,
Western Australia, 1906, M.Koch 1553 (?holo: NSW 181466; iso: PERTH).

Rhodanthe forrestii (F. Muell.) Paul G. Wilson, comb. nov.

Helipterum forrestUY. Muell., S. Sci. Rec. 2:273(1882). - Argyrocome forrestii {¥. Muell.) Kuntze,
Revis. Gen. PI. 1:309(1891). Type citation: 'In the neighbourhood of the Gascoyne-River; J. Forrest.'
Lectotype (here chosen): Gascoyne river, 1882, J. Forrest (lecto: MEL 50292; isolecto: K, MEL
50290, 50291, PERTH).

This species differs from R. polycephala in having the leaves narrowed at the base and not
decurrent and in having the capitula turbinate and not narrow-cylindrical. Further collecting may
show that R. polycephala is only a northern variant of R. forrestii.

Rhodanthe haigii (F. Muell.) Paul G. Wilson, comb. nov.

Helipterum haigUY. Muell., Fragm. 10:107(1877), - Argyrocome haigii (F. Muell.) Kuntze, Revis.
Gen. PI. 1:309(1891). Type: Eucla, Richards (holo: MEL 110217; iso; PERTH).

Helipterum mullinense S. Moore, J. Linn. Soc. Bot. 45:181(1920). Type: Mulline, /.E.C. Maryon
(?iso: MEL 110310).

Note. This species is similar toR. /iMm6o/r/r/rt/irt with which it was confused by Black (1957) and Haegi
(1986). Apart from the presence of larger rays to the involucral bracts, R. humboldliana may be readily
distinguished by its silky pilose achenes for in R. haigii Xht achenes are shortly hirsute. Only R. haigii
occurs in South Australia while both species are found in Western Australia.

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Nuytsia Vol. 8, No. 3 (1992)

Rhodanthe laevis (A. Gray) Paul G. Wilson, comb. nov.

Pteropogon laevis A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:269(1852). - Helipterum laeve
(A. Gray) Benth., FI. Austral. 3:649(1867). - Argyrocome laevis (A. Gray) Kuntze, Revis. Gen. PI.
1:309(1891). Type: Swan River, 1843,7. Drummond 366 (or 356) (holo:K).

Rhodanthe nullarborensis Paul G. Wilson, sp. nov. (Figure 3)

Herba annua ad 30 cm alta, modice lanosa, aromatica. Folia caulina, alterna, linearia, plana, 10-15
mmlonga, 1-1.5 mm lata, sessilia. Inflorescentia corymbiformia, compacta. Capitula homogama,
breviterpedunculata. Involucrum cylindraceum vel anguste turbinatum, 3-4 mm attum, nec radians;
bracteae glumaceae, 3-4-seriatae, nitidae, bracteis intermediis intcrioribusque oblongis, 3-4 mm
longis, ad apicem rotundatis, sparse ciliatis aliter glabris. Receptaculum convexum, glabrum, c. 0.4
mm latum. bracteis receptaculiabsentibus.Flosculic. 12,bisexuales,actinomorphi. Corolla c. 3 mm
longa, fere glabra, nervis ad apicem tubi extensis; lobi deltoidei, c. 1 mm longi, lobis duobus parte
interiore ad basim papillosis, ceteris glabris. Antherae e. 1 mm longae; apex anguste ovatus obtusus,
c. 0.3 mm longus. cellulis luiguste oblongis; caudae filamentae, debiles. Styli apex truncatus, nervo
prominent! ad apicem extenso. Achenium anguste obovoideum, 1 mm longum, breviter hirtellum;
carpophorum annulare, glabrum; pericarpium tenuiter crustaceum, atro rubiginosum, parietibus
radialibus cellularum incrassatis; testa diaphana, ad pericarpum adnata. Pappus corollam aequans,
in statu integro caducae; setae c. 12, filiformae, ad basem breviter connatae, breviter plumosae.

rypwi.- Forrest, Western Australia, 'heavily scented, flowerheads yellow', 30 Aug. 1930,£.R.L. Johnson
75 (holo; PERTH; iso: AD, K, MEL).

Annual erect herb to 30 cm high, branching at base, moderately and loosely woolly, strongly
scented. Leavesnumerous, alternate, cauline, linear, Bat, 10- 15 mm long, 1 - 1.5 mm wide, sessile.
Inflorescence corymbiform, compact. Capitula homogamous, very shortly pedunculate. Involucre
cylindrical to narrow-turbinate, 3-4 mm high, not radiant; bracts glumaceous, 3-4-seriate, glossy,
intermediate and inner bracts oblong, 3-4 mm long, rounded at apex, sparsely ciliate, otherwise
glabrous. Receptacle convex, glabrous, without receptacular bracts, c. 0.4 mm diameter. Florets c.
12, bisexual, actinomorphic. Corolla c. 3 mm long, yellow, n;uTow-tubul.ar in lower half, narrow-
turbinate above, glabrous or rarely very sparsely glandular puberulous, vascular strands extending to
apex of tube; lobes broad-triangular, c. 1 mm long, two of the lobes papillose within at base, the others
glabrous. Stamens: anther loculi c. 1 mm long; tip narrow-ovate, obtu.se, c. 0.3 mm long, the cells
narrow-oblong; tails filamentous, weak. Style arms truncate, papillose, the vascular strand prominent
and extending to base of tip. Achene narrow-obovoid, 1.0 mm long, shortly hirtellcus with duplex
hairs c. 0.07 mm long, minutely bifid at tip, myxogenic; carpopodium a narrow glabrous ring; pericarp
thinly crustaceous, dark rcddi.sh brown, radial cell walls thickened; testa extremely thin and fused to
pericarp, the vascular strand extending to apex of seed; endosperm persistent and forming a sheath
around the embryo. Pappus bristles c . 1 2, ± equal to corolla, very shortly united in a ring at base, shortly
plumose throughout, caducous as a whole.

Distribution. Found only in the Nullarbor Region of Western Australia.

Specimens examined (selection only). WESTERN AUSTRALIA: Kanowna, T.E.H. Aplin 5745
(PERTH); 30 km S of Rawlinna.R.i. Chinnock 1 1262 (AD); 70 mi NNW of Reid, A.S, George 8491

Paul G. Wilson, HcUplerum: Part 1

403

Figure 3. Rhodanthe nuUarborensis. A - Habit. B - Teraiinal cluster of capitula. C - Leafy bract. D - Leaf. E - Outer,
intemiediale, and inner involucral bracts showing enlargement of glandular hair. F- Floret. G - Inner surface of apex of corolla
indicating that only two lobes are papillose. H - Anther. 1- Style apex. J - Pappus bristle. K-Achene. L - Duplex hair from
achene. M - Surface of pericarp. From GJ. Keighery 1623.

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Nuytsia Vol. 8, No. 3 (1992)

(PERTH); 17 km NW of Koonjarra, D.W. Goodall 2511 (PERTH); 49 km NNE of Balladonia,
G.J. Keighery 7623 (PERTH).

Habitat. In shallow open depressions (dongas) that have a layer of heavy soil over limestone.
Flowering time. July to October.

Rhodanthe nullarborensis resembles H. tietkensii with which it has been confused in herbaria.
The latter species may be recognised by its larger leaves, its larger and more hairy capitula, and by
the presence of receptacular bracts.

Johnson & Baird ( 1970) in referring to this species as Helipterum tietkensii state that it is heavily
scented and that it forms dense colonies in the dongas.

Rhodanthe polycephala (A. Gray) Paul G. Wilson, comb. nov.

Pteropogon polycephalus A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:268(1852). - Helipterum
polycephaliim (A. Gray) Benth., FI. Austral. 3:649(1867). - Argyrocome polycephala (A.Gray)
Kuntze, Revis. Gen. PI. 1:309(1891). Type citation: 'Swan river, Drummond', n.v.

Cassinia cuprea F. Muell., Fragm. 3:139(1863). Lectotype (here chosen): Near the Murchison,
A. Oldfield (MEL 110308).

Rhodanthe psammophila Paul G. Wilson, sp. nov. (Figure 4)

Herba erecta ad 40 cm alta. Folia caulina alternata. anguste oblonga vel linearia, ad 6 cm longa,
sessilia, parce decurrentia, lanata et minute stipitato-glandulosa. Capitula heterogama, sessilia,
glomerata. Involucrum anguste turbinatum, c. 4 mm altum; bracteae 4-6-seriatae, obovatae, 3-4 mm
longae, apice albo, rotundato, margine eroso; unguis bractcarum interiorum dense lanato- ciliatus.
Flosculi semper quinque,unofertili,quatuormasculinis. Corolla actinomorpha, sparse puberula; lobi
intra papillosi. Achenium compresso-obovoideum, c. 2.5 mm longum, sparse et minute puberulum;
carpophorum annulare; pericarpium tenuiter crustaceum; testa membranacea pericarpium adnata.
Pappus achenii fertile tarde deciduus; setae c. 13, anguste lineares, acuminatac, c. 2.0 mm longae,
sparse ciliatae pilis gracilibus acuminatis. Achenium bracteis intcrioribus consociatis deciduum.

Typui.- Western Australia, 20 km north ofCarn;trvon,redsanddune,25 September 1987,PaM/ G. Wilson
12604 (holo: PERTH; iso: AD, CANB, K, MEL, NSW, S).

Erect shortly woolly annual branching at and above base, to40 cm high. Leaves cauline, alternate,
narrow-oblong to linear, acuminate, to 6 cm long, sessile, slightly decurrent along the margins, woolly
all over and with minute shortly stipitate reddish brown globular glands. Capitula heterogamous,
sessile in dense clusters, these shortly stipitate forming subglobular heads on slender peduncles 1-3
cm long. Involucre narrow-turbinate, c. 4 mm high; bracts 6-seriate, obovate, 3-4 mm long, rounded
at apex, hyaline in lower 2/3, white petaloid (with flat cells) and wrinkled above with erose margin;
claw of inner bracts densely long woolly ciliate on margin, the wool enveloping the ovaries; stereome
linear, hard, green, slightly shorter than claw, passing downwards into a very short hard stipe.
Receptacle hemispherical, c. 3 mm high and wide, glabrous, smooth. Florets consistently 5 only one

Paul G. Wilson, Uelipterum: Part 1

405

Figure 4. Rhodanthe psammophila . A -Habit. B - Cluster of capitula. C - Capitulum. D - Florets with and without corolla.
E - Carpopodium. F - Pappus bristle. G - Corolla. H - Style apex. I - Achene with pappus. J - Pericarp. K - Duplex
myxogenic achenial hairs. From Paul G. Wilson 12604.

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Nuytsia Vol. 8, No. 3 (1992)

of which is bisexual, the others male. Corolla actinomorphic, yellow, tube c. 2.5 mm long, sparsely
puberulous, narrow-tubular below, narrow-turbinate above; lobes oblong, obtuse, c. 0.5 mm long, the
vascular strands extending to tips, all papillose within along entire length. Stamens: anther loculi c.
1 mm long; appendage ovate with narrow-oblong cells, the marginal ones smaller and equilateral; tails
weak, slightly exceeding collar. Style apex truncate, penicillate, the vascular strand extending to
apex. Achene compressed obovoid, c. 2.5 mm long, sparsely and minutely puberulous with blunt
duplex hairs c. 0.06 mm long; carpopodium a very short glabrous ring; pericarp thinly crustaceous,
pale reddish brown, the cells of epidermis amoeboid, thickened on radial walls; testa membranous,
vascular strand beneath a pericarpial strand and passing over apex of seed; endosperm persistent
around embryo. Pappus (of fertile achene) tardily deciduous; bristles c. 13, narrow-linear, acuminate,
c. 2.0 mm long, sparsely ciliate with slender acuminate hairs. Achene deciduous along with inner
involucral bracts.

Distribution. Western Australia, near the coast between Carnarvon and Onslow, and inland to the
Kennedy Range.

Specimens seen (selection only). WESTERN AUSTRALIA: Kennedy Range, R.J. Cranfield 1913
(PERTH); 10 mi S of Onslow, A.S. George 1 150 (PERTH); southern margins of Lake McLeod,
P.S. Short 2491(MEL); 12 mi NW of Mt Sandiman, B.L. Turner 5401 (PERTH); Brown Range,
B.L. Turner 5411 (PERTH).

Habitat. Grows only on sand dunes, both inland and coastal.

Chromosome number: n = 5 fide B.L. Turner (1970) as Helipterum condensatum.

Rhodanthe psammophilaissupeTficinWy simihuioR. coAic/cnxfltawith which it has been confused
in herbaria. The latter species may be distinguished by the absence of glandular hairs on the leaves,
by the presence of 10-13 florets to a capitulum, and by the persistent pappus. The cells of the white
lamina of the involucral bracts of R. condensala are raised at their tips giving it a slightly rough surface
whereas in R. psammophila the lamina is smooth. Rhodanthe condensata is found from Shark Bay
south to the Murchison River which is south of the distribution of R. psammophila.

The relationships of this species are unclear. In some ways, such as in the structure of the floret,
it is similar to Rhodanthe maryonii, but it differs from that species in having a caducous achene and
a 1 -celled thick crustaceous pericarp (characters that, in this context, may be of little phyllogenetic
significance).

Rhodanthe tietkensii (F. Muell.) Paul G. Wilson, comb. nov.

Helipterum tietkensii F. Muell., Fragm., 8:227(1874). - Argyrocome tietkensii (F. Muell.) Kuntze,
Revis.Gen.Pl. 1:309(1891). Ty’pe.- Between the Alberga and Mt Olga, 1873/4, £. G/7ex(holo: MEL
110506).

5. Rhodanthe sect. Citrinae Paul G. Wilson, sect. nov.

Herba annua erecta sparse lanosa, non glandulosa. Folia caulina alterna linearia vel anguste
oblonga. Capitulahomogamavel heterogama.terminalia. Involucrum radians; bracteaemultiseriatae,
omnes chartaceae et petaloideae, exteriores sessiles, interiores unguiculatae; unguis oblongus vel

Paul G. Wilson, Uelipterum: Part 1

407

ellipticus, hyalinus, lanato-cilialus, stereomate lineari applanato. Receptaculum glabrum, planum.
Rosculi numerosi, homomorphi, aclinomorphi, extra feminei vel bisexuales, intra bisexuales.
Corolla infra lubulosa supra anguste lurbinata, 4-5-loba, sparse pilosa; lobi glabri, intra laeves.
Antherae: appendix anguste ovatus, tenuis, cellulisoblongisparietibustenuibustcaudaefilamentosae.
Styli apex truncatus. Achenium anguste ellipsoidcum, c. 1 mm longum, plcrumque in rostrum
glabrum attenualum,breviterpilosum,pilis myxogeneis; pericarpium debile, cellulis epidermidis ±
cubicis, parietibus tenuibus; testa chartacea, pallido brunea ad pericarpio affixa, crystallis applanatis
munitis. Pappi setae imo basi connatae, breviter plumosae vel denliculatae, caducae.

Holotypus: Rhodanthe citrina (Benth.) Paul G. Wilson

Annual erect herbs, sparsely woolly, not gland dotted. Leaves cauline, alternate, linear or
narrow-oblong. Capitula homogamous or heterogamous, terminal. Involucre radiant; bracts
multiseriate, all glumaceous and pclaloid, outer sessile, the inner with an oblong or elliptic hyaline
woolly ciliate claw. Receptacle glabrous, flat. Florets numerous, homomoiphous, regular, the outer
female or bisexual, the inner bisexual. Corolla tubular below, narrow-turbinate above, 4-5-lobed,
sparsely pilose; lobes glabrous, smooth within. Anther appendix narrow-ovate, c. 0.2 mm long, thin,
cells oblong with thin walls; tails filamentose. Style apex truncate. Achene narrow-ellipsoid, c. 1
mm long, frequently narrowed at apex into a beak; shortly pilose with duplex myxogenic hairs;
pericarp with epidermis of thin-walled cubical cells, other tissue flallened and inconspicuous,
vascular strands lateral (slightly oblique); testa papery, pale brown, adnate to pericarp, sparsely
furnished with broad, flat crystals or these absent when mature, vascular strand ascending to apex and
adjacent to a pericarp ial strand. Pappus caducous; setae connate into a ring at base, shortly plumose
or barbellate.

Rhodanthe citrina (Benth.) Paul G. Wilson, comb. nov.

Leptorhynchos citrinus Benth. in Endl. et al., Enum. PI. Hueg. 64(1837). Waitzia citrina (Benth.)
Steetz in Lehm., PI. Preiss. 1:454(1845). Type citation: 'Swan-River, (Hugel.).'n.v.

Waitzia brevirostris Steetz in Lehm., op.cit. 1 :45 1(1845). Type: In arenosis sylvae supra urbiculam
"Perth", 7 Oct.1839, L. Preiss 15. Lectotype (here chosen): MEL 1585201; isolecto: LD, MEL
1585197.

Waitzia sulpimrea Steetz in Lehm., op.cit. 1:4553(1845). Lectotype (here chosen): In arenosis inter
frutices prope urbiculam "Guildford", 14 Oct. 1839, L. Preiss! (lecto: MEL 1585196; isolecto: LD).

Waitzia steetziana Lehm., op.cit. 1:454(1845). - Waitzia tenella Hook., Bot. Mag. t.5342(1862)
nom.illeg. Type: In solo limoso sylvae prope praedium rusticum "Maddington", 31 Oct.1839,
L. Preiss 6 (synlypesand isosyntypes: LD, MEL 1585194, 1585195, 1585198).

Waitzia dasycarpa Turez., Bull. Soc. Imp. Naturalistes Moscou 24/2:77(1851). Type: Swan River
Colony,/. Drummond 5th coll. no. 65 (holo: KW photo seen; iso: PERTH).

Helichrysum oldfieldii F.Muell., Fragm. 3:134-135(1863). Lectotype (here chosen): Murchison
River, A. Oldfield (MEL 108299).

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Nuytsia Vol. 8, No. 3 (1992)

Distribution. Western Australia south of 11° lat., southern Northern Territory, northern South
Australia, north western New South Wales.

Note: The genus Waitzia, as previously circumscribed, was an unnatural assemblage of species in
the Helipterum complex that had in common a beak-like apex to the achene. In the strict sense (see
Wilson 1992b) Waitzia contains species with the following characters: stipe-like claws to the
involucral bracts, tooth-like papillae (that consist of 2 overtopping cells) on the achenes, and ellipsoid
apices to the styles. In Rhodanthe citrina the inner involucral bracts have flattened scarious claws,
the achenes have normal slender duplex hairs, and the style apices are truncate; it is thus distinct from
Waitzia but similar to some species of Rhodanthe sect. Achyroclinoides. The section Citrinae differs
from sect. Achyroclinoides principally in the nature of the involucral bracts which are all scarious and
petaloid, the outer sessile and the inner stipitate on a broad hyalineciliateclaw; none of them is radiant.
The appearance of the capitula is similar to that found in some species of Chrysocephalum
(Helichrysuni sect. Chrysocephalum), such as Chrysocephalum apiculatum (Labill.) Steetz, and the
two species are sometimes confused.

In Rhodanthe citrina the rostrum to the achene is variable in length and in some specimens is
completely absent; that organ, in any event, does not seem to be necessarily of generic significance
since it merely represents an upwards extension of the pericarp.

It has been shown by Warcup (1990) that Waitzia citrina forms ectomycorrhizal associations
whereas other Rhodanthe species that have been tested do not. It is possible that the seed with which
Warcup worked was either Hyalosperma cotula (Benth.) Paul G. Wilson or Chrysocephalum
apiculatum, both of these form ectomycorrhiza and both resemble W. citrina. No voucher material
can be traced.

Two other species previously placed in Waitzia, i.e. W. paniculata (Steetz) Benth. and IT. con/ca
B.Turner, are also anomalous in that genus and are here transferred to Pterochaeta and Haptotrichion
respectively.

6. Rhodanthe sect. Synachyrum (A. Gray) Paul G. Wilson, comb. nov.

Helipterum sect. Synachyrum A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:231(1852). Type:
Helipterum floribundum DC.

[Helipterum sect. Sericophorum DC., Prod. 6:216(1838) p.p. min. excluding lectotype.]

Plants cottony. Intermediate involucral bracts with thick narrow-oblong stereome not extending
into lamina; cottony. Corolla with vascular strands not extending to tips of lobes. Style apex truncate.
Achene densely silky; carpopodium absent; pericarp thinly coriaceous or papery; testa weak, free
from pericarp, containing oblong imbricate crystals that form a complete cover to seed (see Figure
5). Pappus bristles narrow-oblong and sometimes partially united into a tube at base, caducous.

Rhodanthe floribunda (DC.) Paul G. Wilson, comb. nov. (Figure 5)

Helipterum floribundum DC., Prod. 6:217(1838). Type: Biuren forests at the foot of Peel’s Range,
Interior of N.S.Wales, June \S\1 , A.Cunningham 107 (holo: G-DC photo seen).

Paul G. Wilson, Helipierum: Part 1

409

C0009

Figure 5. Rhodanthe floribunda. A - Pappus. B - Achene. C - Testa with imbricate scale-like crystals.

Helipierum chionolepis F. Muell., Linnaea 25:416(1853). Type citation: 'Juxta montes Flinders-
range et Cudnaka locis sterilibus.' Lectotype (here chosen): MEL 1 10120 with labels 'Cudnaka' and
'Prope montis Flinders range, Oct.51' leg. F. Mueller; syntype: Cudnaka, F. Mueller (MEL 604828).

Helipierum cirratum Morrison, J.Bot. 50:168(1912). Type: Between Globe Hill and Uaroo,
Ashburton River, 1 Oct. 1905, A. Morrison (? iso: PERTH).

Helipierum floribundiim var. luhulipappum}. Black, Trans. &Proc.Roy. Soc. South Australia 36:23
t.2(1912). Type: Oodnadatta, Scpt.191 1, F.C. Slaer per Mrs Mellor (holo: AD 98625088 p.p.).

Rhodanthe sphaerocephala Paul G. Wilson, sp. nov.

Annua erecta ad 25 cm alta. Rami lanosi, omnino foliacei. Folia alterna sessilia, angustissimo
oblonga, c. 10 mm longa, sparse lanosa. Capitula homogama, solitaria, terminalia, sphaeroidea, c.
10mm diam. Bracteaeinvolucrimulti-seriatae,aliquantumcorrugatae,nilidac. Bracteae intermediae
in ambito obovatac, c. 5 mm longae; unguis latissimeoblongus, hyalinus; stereomaspatulatum, parte
terminaliincrassataet dense lanosa; lamina lateelliptica,integra,straminea. Receptaculumeonicum,
tuberculatum. Floscull numerosi, bisexuales, actinomorphi. Corolla c. 4 mm longa, glabra; lobi 5,
anguste oblongi, c. 1.5 mm longi, intra laeves; nervi corollae ad apice tubi extensi. Antherae:
appendix sterilis ovala, aeuta, c. 0.6 mm longa, cellulis oblongis, pai ielibus nec incrassatis; caudis
filamentis debiles. Styli apex truncatus, nervo prominenti ad basem apicis extenso. Achenium
ellipsoideum c. 2 mm longum, dense sericeum, pilis in lobos inaequales terminantibus; pericarpum
membranaeeum, translucens; testa tenuiter eoriacea, pallide brunea, cellulis crystallis oblongis

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Nuytsia Vol. 8, No. 3 (1992)

repletis. Pappi setae librae, caducae, in parte inferioreangusto ellipticae, breviterplumosae, in parte
superiore filiformae, sparse plumosae, ad extremum plumoso-penicillatae.

rjpMs.- BeleleStation.c. 56 km WNW of Meekatharra, Western Australia, 30Oct. \965, D.W. Goodall
3366 (holo: PERTH; iso: CANS).

Erect annual to 25 cm high branching at and shortly above base. Stems loosely woolly, leafy
throughout. Leaves cauline, alternate, sessile, very narrow-oblong, c. 10 mm long, 1 mm wide,
sparsely woolly with crinkly hairs. Capitula homogamous, solitary, terminal to long branches, almost
spherical, c. 10 mm diameter. Involucral bracts multi-seriate, somewhat crinkly, glossy, straw-
coloured, similar throughout; Intermediate bracts obovate in outline, c. 5 mm long; claw very broad-
oblong, hyaline, woolly ciliate on distal margins, stereome spatulate with the terminal portion
thickened and bearing a dense tuft of wool on abaxial surface, lamina broad-elliptic, entire, .somewhat
wrinkled, glabrous, glossy, straw-coloured. Receptacle ovoid, c. 3 mm high, 2.5 mm wide, glabrous,
tuberculate with raised achenial scars. Florets numerous, homogamous, bisexual, actinomorphic.
Corolla c. 4 mm long to base of lobes; tube narrow-cylindrical, glabrous; limb narrow-campanulate,
glabrous; lobes 5, narrow-oblong, c. 1.5 mm long with a few glandular hairs (with large oblongoid
terminal cells) on abaxial side, smooth within, vascular strands reaching to base of lobes. Anther
appendage ovate, acute, c. 0.6 mm long, cells oblong with unthickened walls, marginal cells not
differentiated; anther tails filamentous, weak. Style tip truncate, shortly penicillate, the vascular
strand stout, not extending into apex. Achene ellip.soidal, c. 2 mm long, densely long-silky to base,
the duplex hairs terminating in two very uneven arms with one very short (c. 0.005-0.01 mm) and the
other long (c. 0.3 mm), not myxogenic; pericarp thin, translucent; testa thinly coriaceous, pale brown,
the outer cells filled with flat broad-oblong crystals that form an armour-like covering to seed. Pappus
bristles slightly shorter than corolla, free, caducous, linear-elliptic and shortly plumose in lower half,
filiform and sparsely plumose in upper half, plumose tufted at end.

This species is known only from the type collection for which ecological information is not
available.

Rhodanthe sphaerocephala would appear to have no close relative but the morphology of its florets
and fruits suggests affinity to other members of the section Synachyrum.

Rhodanthe sterilescens (F. Muell.) Paul Wilson, comb. nov.

Helipterum sterilescens F. Muell., S. Sci. Rec. 2:274(1882). - Argyrocome sterilescens (F. Muell.)
O. Kuntze, Rev. Gen. PI. 1: 309 (1891). Type: Gascoyne River, W.A., 1882, Pollack (holo:
MEL 110428; iso: PERTH).

This species bears dense clusters of capitula that give it a similar appearance to Cephalipterum
drummondii with which it has been frequently confused.

Rhodanthe sterilescens differs from C. drummondii in the following significant characters: sessile
capitula at base of plant in addition to terminal (not all terminal); capitula in a compact cyme (not
dense umbel); indumentum woolly (not of curled filiform hairs with septate base and glandular
stipitate hairs); achenial hairs slender of the normal duplex type (not clavate helically coiled and

Paul G. Wilson, Helipterum: Part 1

411

barbed woolly); achcnes free, three fertile and silky, the rest sterile and glabrous (not coalescent with
1 or two fertile and the remainder sterile but hairy); achene narrowed at apex (not broad and lobed
at apex). The two species are similar in floral characters, in the seed morphology, and in form of the
crystals present in the testa.

Rhodanthe stuartiana (Sond. & F. Muell.) Paul G. Wilson, comb. nov.

Helipterum sluarlianiim Sond. & F. Muell., Linnaea 25; 518 (1853). - Helipterum floribundum var.
stuartianum (Sond. & F. Muell.) Benth., FI. Austral. 3:642(1867). Type citation: 'Ad fl. Murray leg.
Stuart.' Lectolype (here chosen): Ad fl. Murray, F. Mueller (MEL 604830).

The type of Helipterum stuartianum is stated to have been collected by Stuart. A herbarium
O. W. Sonder specimen (MEL 604830) labelled 'Ad fl. Murray' was collected by F. Mueller and bears
in (?)Sonder’s handwriting the phrase 'Antherae basi bisetae'; these words are used in the published
description. A further collection from 'Near the Murray R.' no. 785 (MEL 1539201) was evidently
from Stuart. It bears the note 'This plant did not occur in my collection at Adelaide but 1 found seed
of it amongst those you procured from Stephens' [? William Stephens, a garden’s collector]. I have
selected the Mueller collection (MEL 604830) as the lectotype since it was evidently studied by
Sonder.

Rhodanthe troedelii (F. Muell.) Paul G. Wilson, comb. nov.

Helipterum troedelii F. Muell., Victorum Naturalist 7:77(1890). Type citation: 'Near the Barrier-
Ranges; Mrs Irvine, at Leight’s Creeks, beyond Beltana; Mrs Richards.' Lectotype (here chosen);
Leigh’s Creek, \?&1 , Mrs Richards {\tc\o\ MEL 110636; isolecto: MEL 696324, NSW 181421).

Helipterum troedelii vw.patensEwait, J. White & B. Rees.Proc. Roy. Soc. Victoria 22 n.s.: 15(1909)
pro parte as to lectotype. - H.roseumvta. patens (Ewart et al.) J. Black, Trans. & Proc. Roy. Soc. South
Australia 45:21(1921). Type citation: 'Mt. Lyndhurst, M. Koch, No. 1644 (1899); Fraser Range, W.
Austr.,R. Helms, 1891.’ Lccto/ypc.' MtLyndhurst.S. Australia, M.Koc/! 1644(1899), AD 976501 28,
fide D. Cooke in Jessop & Toelken, Fl. South Australia edn 4, 1549(1986).

The syntype material of H. troedelii var. patens consists of two species. The collection of Max
Koch represents Rhodanthe troedelii while the collection of R. Helms represents the 'Nullarbor'
variant of/?, chlorocephala. J.M. Black (1921 /.c.) indicated that he had examined duplicate material
of the Helms syntype and recognised it as being a variant of Helipterum roseiim, he therefore
transferred var. patens to that species (which is a synonym of Rhodanthe chlorocephala). However,
D. Cooke (1986 l.c.) lectolypified the name on a Max Koch collection in herb. AD which had been
received by Black as a donation from MEL and is probably a duplicate of the Max Koch syntype (it
does not bear the month of collection that is present on the syntype). The AD specimen had been
annotated by Ewart as 'typical //. Troedelii'. In herb. MEL the Max Koch syntype has had the epithet
'var. patens' deleted, presumably by Ewart, while the Helms syntype has been labelled 'type'. It would
therefore appear that subsequent to its publication Ewart intended to apply the epithet var. patens to
the element represented by the Helms collection which was also the way in which Black applied the
name (as a synonym off/, roseum) in 1921 and in the Flora of South Australia (1929).

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7. Rhodanthe sect. Helipteridium (A. Gray) Paul G. Wilson, comb. nov.

Helipterum sect. Helipteridium A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:231(1852). Type:
Helipterum discoideim A. Gray

Helipterum sect. Aglossum F. Muell., Fragm. 3:137(1863) nomen.

Annual erect herb, sparsely woolly with filiform hairs, and with curled uniseriate septate hairs,
eglandular. Major axes slender. Leaves alternate, flat, sessile. Capitula heterogamous, solitary,
terminal. Involucre hemispherical, not radiant. Receptacle glabrous with prominent alveolae.
Involucral bracts c. 4-seriate, glabrous, glossy, uniform; claw broad-oblong with a broad herbaceous
stereome and narrow scarious margins; lamina short, erect, scarious. Florets bisexual or the inner
male. Corolla slightly zygomorphic; lube cylindrical, pubescent with gland-tipped hairs; throat
turbinate, glabrous; lobes 5, unequal, papillose within, vascular strands extending to apex of lobes.
Anther appendix deltoid, cells short and irregular; anther tails filamentous, weak. Style apex truncate
with a subulate coalescence of epidermal cells in the centre. Achene ellipsoid, silky pilose with
normal duplex hairs; carpopodium annular; pericarp brittle; testa soft with numerous oblong crystals,
lower 2/3 surrounded by a network of vascular strands. Pappus bristles linear-acuminate, plumose.

A monotypic section.

Rhodanthe heterantha (Turcz.) Paul G. Wilson, comb. nov.

Helipterum heteranthum Turcz., Bull. Soc. Nat. Mosc. 24/1 :198 (1851). Type: Nova Hollandia,
J. Drummond 4x\\ coll. n.214 (holo: KW photo seen; iso: K, MEL 110231, 110233).

Helipterum discoideum A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:231(1852). Type citation:
'Variat a, involucro pallido; 6, involucro sanguineo. Swan River (6, Swan River to King George’s
Sound), Drummond.' {syn: K 'J. Drummond 96', photo seen).

Helipterum anactinumF. Muell., Fragm.3:137(1863). Type citation: 'Adsinum orarium Champion
Bay. Walcott et Oldfield.' Lccto/>’Pe(herechosen): Champion Bay, A. C)/r///e/£/(lecto: MEL 110229).

Helipterum heteranthum van majus Benth., FI. Austral. 3:643(1867). Type: Between Moore and
Murchison rivers, 7. Drummond 6th coll. n.l52 (holo: K; iso: MEL 110226, NSW).

Helipterum pachychaetum W.Fitzg., J. W. Austral. Nat. Hist. Soc. No.2:25(May 1905). Lectotype
(here chosen): Jacup, 50 miles west of Phillips River, Oct.1903, C.R.P. Andrews (lecto: PERTH;
isolecto: NSW).

Helipterum heteranthum var. mmor Ewart, J. White & Tovey, J. Roy. Soc. N.S. Wales 42: 190(1908).
Lectotype (here chosen): Cowcowing, W.A., Oct. 1904, A7. Koch 1108 (lecto: MEL 110220; isolecto:
MEL 110221).

8. Rhodanthe sect. Helichrysoides (A. Gray) Paul G. Wilson, comb. nov.

Pteropogon sect. Helichrysoides A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:267(1852). Type:
Pteropogon spicatus Steetz

Paul G. Wilson, Helipterum: Part 1

413

Annual erect herbs, woolly or glandular puberulous. Leaves cauline, alternate, filiform to
obovate. Capitula homogamous or heterogamous, clustered, paniculate or condensed in spike-like
inflorescences. Involucre turbinate to cylindrical, not radiant; bracts multiseriate, glabrous or
glandular pubescent, scarious or with herbaceous tips; stcreome small, basal. Receptacle convex to
broad-conical, glabrous or scabrid. Florets 5-30, bisexual, or the Innermost male. Corolla
actinomorhpic , sparsely puberulous, narrow-cylindrical below, narrow-campanulate above; lobes
ovate, papillose or smooth within, vascular strands extending to tips. Stamens: anther apex ovate the
distal cells slightly raised at tip; tails filamentous, weak. Style apex broad to narrow -deltoid,
puberulous, vascular strand extending to tip. Achene narrow-obovoid to narrow-turbinate, pilose
almost to base; carpopodium an extremely short ring; pericarp rough, thick, brittle, dark brown; testa
free from pericarp, membranous, vascular strand extending almost completely around seed; crystals
absent. Pappus persistent; bristles thick at base, plumose, the cilia all acute.

The section f/e/ic/iryso/r/es is sim ilar to sectLeiochrysim but differs in the nature of the in vol ucral
bracts (foliaceous at apex except in R. spicata), the presence of vascular strands at the apex of the
corolla lobes, the deltoid style apex, and the thick brittle pericarp. It is also similar to species in the
genus Podotheca in the morphology of the achene, the deltoid style apex, and the foliaceous apex to
the involucral bracts. In the latter genus the chromosome number is n=13, 26 (Short 1989) whereas
in Rhodanthe sect. Helichrysoides the three species R baiiii, R. pollackii, and R. spicata, have a
chromosome number of n = 10 (B. Turner in sched.y, the number for R. charsleyae has not been
recorded. Ware up (1990) has shown that/?, battii and/?, spicata do not form ectomycorrhiza whereas
species of Podotheca that have been tested do; this again suggests that the species in sect.
Helichrysoides should not be placed in Podotheca.

Rhodanthe battii (F. Muell.) Paul. G. Wilson, comb. nov.

Helipterum batt// F.Muell., Victorian Naturalist 10:144(1893). Type: Between Dundas-Hills and
Lake-Lefroy, 1893,7.0. Ball (holo: MEL 110727).

[Podotheca pollackii unci, non (F.Muell.) Diels: Diels &Pritzel, Bot. Jahrb. Syst. 35:615-623(1905)
as to specimen cited and description.]

Rhodanthe pollackii (F. Muell.) Paul G. Wilson, comb. nov.

Podosperma pollackii F. Muell., Fragm. 12:21(1882); - Podotheca pollackii (F. Muell.) Diels, Bot.
Jahrb. Syst. 35:617(1905). Type citation: 'In vicinia fluminis Gascoyne-River; Forrest et Pollack.'
Syntype: Gascoyne River, 1882, J. Forrest (PERTH).

Rhodanthe charsleyae (F. Muell.) Paul G. Wilson, comb. nov.

Helipterum charsleyaeF.'Mnc\\.,¥xugm. 8:168(1874). Type citatiom'In vicinia lacusAmadei, Giles;
prope lacum Lefroyi, Forrest.' Lectotype (here chosen): Lake Lefroy, State Well, A. Forrest (MEL
110714).

Rhodanthe spicata (Steetz) Paul G. Wilson, comb. nov.

Pteropogon spicatus Steetz in Lehm.,Pl.Prciss. 1:479(1845); A. Gray, Hooker’s J. Bot. KewGard.
Misc. 4:268(1852). - Helipterum spicatum (Steetz) Benlh., FI. Austral. 3:647(1867). Type citation:

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'In limoso-calculosis illustribus lateris orientalis montis Lehmann', 4 Sept. 1839, L. Preiss 24.
Lectotype (here cho.scn): MEL 110492; i.solecto MEL 1 10493, MEL 604832.

Helipterum monencyanlhoides F. Muell., Fragm. 3:137(1863). - H. spicatum var. paUens Benth.,
FI. Austral. 3:648(1867). Type citation: 'Ad rivos Tom [i.e. Tone] River et Salt River', G. Maxwell.
Lectotype (here chosen): Banks and valleys of Salt River, G.Maxwell 75 (MEL 1 10707).

Calocephalus globosus M.Scott & Hutch., Kew Bull. 1916:36(1916). Type: Kauring, on York -
Greenhills line. Western Australia, F. Steward 505 (holo: K photo seen).

9. Rhodanthe sect. Actinaria Paul G. Wilson, sect. nov.

Herha annua erecta sparse glanduloso puherula. Folia caulina, alterna, applanata. Capitula
homogama, solitaria, terminalia. Involucrum cupulatum ad hemisphaericum, radians; hracteae
manifesto dimorphae, multiseriatae; hractea exteriora angu.ste linearia. acuminata, glanduloso
puherula; hracteae interiores ungue anguste ohlongo, lamina ovata, petaloidca. Receptaculum
pulvinatum, glahrum, laeve. Flosculi numerosi, hi.sexuales, homomorphi, actinomorphi. Corolla
infra tuhulosa supra anguste turhinata, pallido flava. 5-loha; lohi glahri, extra .sparse pilosi, intra
laeves. Antherae: appendix anguste ovatus, ccllulis anguste ohlongis parietihus tenuihus; caudae
parte proximali firmae, parte distali filamentosae dehiles. Styli apex ellipsoideus, nervo carenti.
Achenium cylindraceum,c. 1 mm longum,modicepilosum,pilorum longitudinedissimili (0.02-0.2
mm); carpophorum annulare; pericarpium unicellulam crassum, crustaceum; testa memhranacea,
unicellulam crassum, translucens, crystallis carentihus. Pappi setae filiformes,imo hasi connatae, in
statu integro caducae, hreviter plumosae.

Typus: Rhodanthe margarethae (F. Muell.) Paul G. Wilson

Erect annual herhs, branched, sparsely glandular puberulous. Branches slender, reddish. Leaves
cauline, alternate, thin. Capitula homogamous, solitary, terminal to branches. Involucre cup-shaped
to hemispherical, c. 2 cm diameter, radimit; bracts distinctly dimorphic; outer bracts narrow-linear,
acuminate, c. 4 mm long, glandular puberulous; inner bracts with erect narrow-oblong woolly ciliate
claw and spreading ovate pelaloid while or pale yellow lamina, the stercome narrow-oblong, thick
and glandular puberulous. Receptacle cushion-shaped, c. 5 mm diameter, smooth, papillose around
alveolae. Florets numerous, bisexual, 5-merous, actinomorphic. Corolla narrow-tubular below,
expanded above, pale yellow, very sparsely puberulous outside, lobes smooth within, cells of inner
epidermisof lobes oblong. Stamens: anther tip narrow-ovate acute, cells narrow-oblong, the marginal
cells forming an indeterminate row; tuilhcr tails firm, equal to collar. Style apex elliptic, shortly
papillose, without vasculature. Achene cylindrical c. 1 mm long, moderately short pilose, hairs
rounded at apex and of different lengths (0.02-0.2 mm); carpopodium a short glabrous ring; pericarp
thin and crustaceous, brown; testa thin, translucent, without crystals. Pappus caducous as a whole;
bristles filiform, shortly plumose throughout, connate in ring at base.

A section of two morphologically similar species that are restricted in their distribution to the
northwestern region of Western Australia.

The section Actinaria differs from other members of the Rhodanthe complex in having narrow-
linear outer involucral bracts, elliptic style apices that are without vasculature, filiform pappus
bristles, and short blunt achenial hairs that are of different lengths. The habit of the two included

Paul G. Wilson, Helipterum: Part 1

415

species is similar to that of R. manglesii in section Rhodanthe, as was noted by Mueller in his
description of H. margarethae, but in floral characters the sections are very distinct.

Rhodanthe margarethae (F. Muell.) Paul G. Wilson, comb. nov.

Helipterum margarethae F. Muell., Fragm. 11:48(1878); F. Muell., Syst. Census Austral. PI.
80(1882) 'Margaritae'\ F. Muell., Sec. Syst. Census Austral. PI. 136(1889) 'Margaritae'; Domin,
B iblioth . Bot. 89: 667( 1 929); Ar gyro come margarethae (F. Muell.) Kuntze, Revis. Gen. PI . 1 : 309( 1 89 1 )
Margaritae . Type citation:'Ad amnem Jones’s Creek ct flumen George's River prope Nickol-Bay;
J. Forrest'. Lectotype (here chosen); Jone's Creek and George River south of Roebourne, 1878,
J. Forrest (MEL 1 10234).

Distribution. Pilbara region (Fortescue Botanical District) of Western Australia.

I have not found a syntype that was collected by J. Forrest near Nickol Bay, however, there is a
specimen (MEL 110235) labelled by Mueller 'Helipterum Margarethae, Nickol River, A. Forrest'
which is possibly the collection referred to by him since Alexander Forrest was with John Forrest on
the expedition to the Nickol Bay area in 1878. This A. Forrest collection is oi Rhodanthe frenchii.
Mueller describes the leaves as 'amplexantibus' and the involucral bracts as 'Candida', that is, pure
white, terms that apply to R. margarethae as lectotypified by the 'Jone’s creek and George River'
specimen, but not to the Nickol River collection.

The epithet 'margarethae' honours Margaret Forrest (1845-1929), wife of John Forrest the
explorer and statesman. Mueller evidently considered a preferred latinization of her name to be
'margaritae' for he used this variant in 1882 and 1889 as did Otto Kuntze in 1891.

Rhodanthe frenchii (F. Muell.) Paul G. Wilson, comb. nov.

Helipterum frenchii F. Muell., S. Sci. Rec. 3:34(1883). Type citation: 'Near Menilayalya in the
vicinity of Shark-Bay; J. Forrest' (holo; MEL 50293 'Head of Menilyalya R., 1882, J. Forrest').

[Helipterum margarethae F. Muell., Fragm. 1 1 :48(1878) pro parte as to the Nickol River collection,
not as to lectotype]

Distribution . Northern Carnarvon and north-west Ashburton Botanical districts of Western Australia.

10. Rhodanthe sect. Anisolepis (Steetz) Paul G. Wilson, comb, et stat. nov.

Anisolepis Steetz in Lehm., Pl.Prciss. 1:446(1845). Type: Anisolepis pyrethrum Steetz

Annual erect herb, glabrous orsparsely and minutely puberulous. Stem simple or branched above,
towards the base thickened and aerenchymatous. Leaves cauline, simple, entire, alternate or the lower
opposite, if submerged filiform, otherwise oblong and somewhat fleshy, c. 5 mm long, towards the
apex becoming ovateand with scarious margins. Capitulahomogamous, solitary and terminal to stem
and branches. Involucre at first turbinate, becoming hemi.spherical with age, often subtended by the
uppermost leaves; bracts 2-seriatc, all radiant and with claws; claw erect, narrow-deltoid, c. 1 mm

long, margin scarious, stereomeprominent.iuirrow-oblong, thick andraised, sparsely glandular, limb

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Nuytsia Vol. 8, No. 3 (1992)

petaloid, radiating, white, elliptic, c. 5 mm long, eventually breaking away from the persistent claw.
Receptacle narrow-conical, deeply foveolate, fimbrilliferous around foveolae. Florets numerous,
bisexual. Corolla tube cylindrical, c. 1.5 mm long, sub-campanulate above, shortly 5-lobed; lobes
triangular, abaxial surface with a few hairs that have large ovoid gland-cells at their tips, two of the
lobes with adaxial surface papillose at base, the others glabrous, margins thickened; cells of inner
epidermis of lobes irregularly oblong, of the neck oblong and undulate on margin. Anthers included;
terminiU appendage broad-oblong, obtuse, thin, cells oblong, thin-walled; tails weak and filamentous.
Style tips rounded to acuminate; vascular strand extending to base of tip. Achene barrel-shaped, c.
1 mm long, dark reddish brown, pilose with fine hairs, not myxogenic; carpopodium very short,
annular; pericarp very thinly crustaceous, pale brown; testa mauve, thinly coriaceous, vascular strands
not apparent; crystals absent. Pappus c. 1 mm long of c. 10 narrowly triangular serrate scales united
in lower half, tardily deciduous as a whole.

Rhodanthe pyrethrum (Steetz) Paul G. Wilson, comb. nov.

Anisolepis pyrethrum Steetz in Lehm., PI. Preiss. 1;447(1845). - Helipterum pyrethrum (Steetz)
Benth. , FI. Austral. 3;642(1867).- Argy/'ocomcpyre/Ziram (Steetz) Kuntze.Revis. Gen. PI. 1;309(1891).
Type citation; 'In depressis uliginosis sylvae supra oppidulum Perth, d. 26.Sept.1839. Herb. Preiss.
No. 14.' Lectotype (here chosen): MEL 108293; isolectp: MEL 108294.

There is some variability within this species but it is unclear as to how much is due to growth
conditions. Collections made to the north of Perth have the lower (underwater) leaves linear-
acuminate and opposite, whereas collections made to the south of Perth have the underwater leaves
filiform and scattered. There are also slight differences between different populations in the size of
achenes, in the distribution of the globular glands, and in the branch ing of the inflorescence. However,
all these characters show such variability over the total species range that specific or infraspecific
discrimination docs not appear practicable.

Rhodanthe pyrethrum is similar in some features to the species in Rhodanthe sect. Achyroclinoides,
noticeably in the size and morphology of the achene. It differs from this section and from other
sections oi Rhodanthe, in the nature of the invol ucral bracts which are all radiant and wh ich have claws
that possess a very thick narrow-oblong stereome, in the laciniate scales of the pappus, and in the
rounded to acuminate style apices.

11. Rhodanthe sect. Polyphyllum Paul G. Wilson, sect. nov.

Herba annua, modice lanosa, glandis sessilibus globosis sparse ornata. Folia numerosa, alterna,
filiformia. Inflorescentiumcorymbosum. Capitulahomogama,radianlia. Involucrumlateturbinatum
c. 5 mm ahum; bracteae intermediae scariosae, sparse lanosae, stereomate anguste oblongo, crasso;
bracteae intimae similis sed limbo albo ornato. Flores 8-12, bisexuales; corolla tubularis, sparse
glanduloso puberula, lobis 5, brevibus, intra glabris, nervis adapicem extensis. Antherum appendice
late ovato, crasso, caudis collum aequantibus. crassis, ramosis. Styli apex truncati, nervo crasso sub-
apice extenso. Achenium doleiforme, truncatum, modice hirtellum; carpopodium crassum; pericarpium
crustaceum, rugosum; testa chartacea, libra, nervo 3/4 circumdato; nervis pericarpi et testae in posite
laterali locatis. Pappus corollam aequans; setae filamentosae, brevitcr plumosae, caducae.

Typus: Rhodanthe polyphylla (F. Muell.) Paul G. Wilson

Paul G. Wilson, Heliplerum: Part 1

417

Annual erect herb, somewhat woolly and with scattered minute globular reddish brown glandular
hairs, single stemmed. Leaves numerous, filiform. Inflorescence a terminal corymb. Capitula
homogamous, shortly pedunculate, radiant. Involucre broadly turbinate c. 5 mm high; bracts c.
4-seriate; outer and intermediate bracts scarious, sparsely woolly with a thick narrow-oblong
stereome; innermost bracts similar but with a short oblong white limb. Florets 8-12, bisexual; corolla
narrow-tubular, slightly broader above, sparsely glandular puberulous; lobes 5 short, glabrous within,
sparsely glandular puberulous outside, cells of inner epidermis narrow-oblong, vascular strands
passing to apex of lobes. Anther appendage broad-ovate, thick, cells unevenly oblong; anther tails
stout, branched, equal to collar. Style apex truncate, vascular strand thick and extending to just below
apex. Achene barrel-shaped with a truncate apex, moderately hirtellous with slender duplex hairs;
carpopodium thick; pericarp crustaceous, rough; testa papery, free from pericarp, vascular strand
passing over apex of seed; vascular strands of pericarp and of testa in lateral position (in relation to
cotyledons). Pappus equal to corolla; bristles filamentous, shortly plumose, shortly united at base,
caducous.

Note. A number of features make this monotypic section unique in the Helichrysum complex; these
are: 1) the numerous caulinc, lanate, filiform leaves, 2) the short, stout, branched anther tails, and 3)
the thick anther appendage with unevenly shaped oblong cells. Its position in relation to the other
sections of Rhodanthe and to genera in the Casssinia complex is unclear. It is likely that this section
should be recognised as a distinct genus.

Rhodanthe polyphylla (F.Muell.) Paul G. Wilson, comb. nov.

HelipterumpolyphyllumF. Muell.,Fragm. 1:35(1858). -Argyracowe/Jo/yp/jy/Za (F.Muell.) Kuntze,
Revis. Gen. PI. 1:309(1891). Type citation: 'In planiticbus basalticis a fluvio Brisbane usque ad
monies Peak Range. - In tractu Kent's Plains. W. Hill.' Lectotype (here chosen): Brisbane River, Peak
Range, F. Mueller (MEL 109740); syntype: Moreton Bay, Kents Plains, anon.(MEh 109757).

Distribution. Eastern Queensland and north-east New South Wales.

Cephalipterum A. Gray

Cephalipterum A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:271(1852). Type: Cephalipterum
drummondii A. Gray

Annual erect herb, indumentum of curled filiform hairs septate at base with glandular slipitale
hairs below capitula. Leaves caulinc and basal, linear to obovate. Capitula heterogamous, very
shortly pedunculate in dense terminal umbels. Involucre cup-shaped; bracts multiseriate, scarious,
the innermost with a white or yellow lamina. Receptacle small, conical, glabrous. Florets numerous;
1-2 outer bisexual, remainder male. Corolla narrow-urccolate, shortly 5-lobed, glabrous, lobes
smooth within; va.scular strands terminating in tube. Anthers: appendix ovate with narrow-oblong
cells; collar short and broadly expanded at ba.se; tails weak; style apex truncate, somewhat broader
than style branch, prominently papillose. Achcnebroadly compressed obovoid with abroad-circular
apex that has a hard raised lip on the abaxial margin; sterile achenes hard, broad-obovoid, the apical
surface vertical; attachment small, carpopodium absent; indumentum (fertile and sterile achenes) of
dense white clavatc helically coiled duplex hairs on abaxial surface, of woolly clawed simple hairs
on adaxial surface; pericarp thick and hard; testa thinly coriaceous closely covered all over with broad-

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Nuytsia Vol. 8, No. 3 (1992)

oblong imbricate cryslals; vascular strands of pericarp and testa medial in relation to the broad-
obovate embryo. Pappus bristles free, caducous, linear to filiform, shortly plumose in upper half, the
terminal cilia congested, thickened and recurved.

Cephalipterum drummondii A. Gray, op.cii 111. Type citation: 'Swan River, Drummond.' n.v.

Cephalipterum drummondii f. major Diels & Pritz., Bot. Jahrb. 35:615(1905) nom.illeg. (based on
type/orma).

Cephalipterum drummondii f. minor Diels & Pritz., l.c.

Type citation: 'hab. in distr. Coolgardie pr. Kanowna flor. m. Aug. (W.V. Fitzgerald in hb. Berk); in
distr. Austin pr. Murrinmurrin fl. m. Nov. (W.J. George).' Neotype (here chosen): Kanowna, August
1898, W.V. Fitzgerald (PERTH).

Note. The syntype material in herb. B is destroyed, 1 have therefore chosen as neotype a probable
isosyntype in PERTH.

This genus is similar in many features to species in Rhodanthe sect. Synachyrum, in particular in
possessing a dense imbricate crystalline cover to the seed. Only one or two seeds in the capitulum
are fertile but these coalesce to the sterile achenes by means of a dense mass of barbed hairs on their
adaxial surfaces which cause the entire cluster to be dispersed as one unit from which one seedling
emerges with the sterile achenes forming a persistent ring around the base of the root. The fertile
achenes bear at their tip a raised rim on their abaxial margin; this character is unique in the Rhodanthe
complex.

A number of variants of this species occur which vary in the size and colour of the in volucral bracts
with sometimes two or more variants being present at the same locality. It is probable that the variants
have edaphic preferences but speculation on this matter has still to be confirmed. Some of these
variants may warrant recognition; Turner (1970) has recorded chromosome counts of n =12 and
n = 14 for the species, however, the voucher material (PERTH) for the counts appears to be
morphologically identical.

Erymophyllum Paul G. Wilson
Erymophyllum Paul G. Wilson, Nuytsia 7; 105(1989).

Type: Erymophyllum gracile (A. Gray) Paul G. Wilson.

Pteropogon sect. Helipteroides A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4;269( 1852). Lectotype:
Pteropogon gracilis A. Gray (Wilson 1989b).

For list of species see Wilson 1989b.

Paul G. Wilson, Helipteruni: Part 1

419

The Hyalosperma group
Hyalosperma Steetz

Hyalosperma Steetz in Lehm., PI. Preiss. 1:476(1845). Lectotype: Hyalosperma glulinosum Steetz
(Wilson 1989a).

Helipterum sect. Pachypterum Steetz in Lehm., PI. Preiss. 1:473(1845). Lectotype: Helipterum
cotula (Benth.) DC. (Wilson 1989a).

Pteropogon sect. Pteropogonopsis A. Gray, Hooker’s J. Bot. Kew Card. Misc. 4:269(1852). Type:
Pteropogon demissus A. Gray.

For description and list of species see Wilson 1989a.

Gilberta Turcz.

Gilberta Turcz., Bull. Soc. Nat. Mosc. 24/1:192 (pre 27 March 1851). Type: Gilberta tenuifolia
Turcz.

Antheidosorus A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 3:98 (April 1851). Type: Antheidosorus
gracilis A. Gray.

Erect annual, moderately arachnoid. Leaves alternate, filiform. Capitulaheterogamous, terminal
to short lateral branches, nutant in fruit. Involucre turbinate to hemispherical, c. 4 mm high, radiant;
outer bracts very broadly ovate, hyaline, slightly woolly at base; intermediate bracts with a broad
hyaline slightly woolly claw and a short spreading yellow limb; innermost bracts obovate, hyaline,
pale yellow, not radiating; stereome of ray bracts short and triangular. Receptacle flat, covered with
scarious narrow-oblong receptacular bracts of which c. 5 surround each floret. Florets actinomorphic,
theouterrow bisexual and the remainder male. Bisexual florets: Corolla glabrous, the vascular strands
terminating at apex of throat; tube narrowly cylindrical; throat very short and turbinate; lobes 5,
oblong, acute, prominent, smooth within, the cells narrow-oblong with straight walls. Anthers
exserted; terminal appendage ovate, c. 0.2 mm long, cells narrow-oblong with thin walls; tails weak
and filamentous. Style apex narrow-ovate to deltoid, densely papillose, the vascular strand slender
and extending to base of apex. Achcne compressed obovoid, c. 1.5 mm long, minutely colliculate;
carpopodium minute (scarcely apparent); pericarp thick and transparent, strongly myxogenic; testa
thinly coriaceous, brown, without crystals, vascular strands not apparent. Sterile achencs linear.
Pappus of 5 unequal linear-acuminate shortly plumose bristles of which one or two have clavate
terminal cilia; pappus of fertile florets weak and caducous as a whole, of male florets indurated at base
and persistent on achene.

Gilberta tenuifolia Turcz., o/;.c;7.193. Type: Western Australia, 7. Gilbert 111 (holo: KW).

Antheidosorus gracilis A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 3: 174 (June 1851). -Myriocephalus
gracilis (A.Gray) Benth., FI. Austral. 3:559(1867). Type: Swan river Colony, J. Drummond, n.v.

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Nuytsia Vol. 8, No. 3 (1992)

Helipterum verecundum S. Moore, J. Linn. Soc. 34:200(1899). Type citation: 'Juxta Coolgardie
repperi mens. Aug. florentem', S. Moore, n.v.

Asa Gray (1851) suggested that, due to the presence of receptacular bracts, this species had
affinities to Myriocephalus. However, as has been pointed out by Short et al. (1989), Gilberta has
little in common with M. appendiculatus Benth., the type of Myriocephalus. The florets and achenes
of Gilberta tenuifolia arc, in fact, very similar to those of Hyalosperma, particularly to H. ghitinosum
(Steetz) Paul G. Wilson. Both of these species have a similar corolla with narrow-oblong cells in
the inner epidermis of the lobes, almost identical anthers and style apices, a similar pappus, and very
similar achenes in which the pericarp forms a pellucid myxogenic layer around the seed. The leaves
and indumentum of Gilberta are also of the same form as those found in Hyalosperma. Gilberta differs
most obviously in the arrangement of the capitula and in the presence of capitular bracts around each
floret; it also has glabrous corolla lobes whereas in //. glutinosum the corolla lobes have hairs that
possess a large ovoid terminal gland.

According to Warcup (1990) both Gilberta and Hyalosperma form an ectomycorrhizal association
while four species currently placed in Myriocephalus that have been tested do not.

The two names, Gilberta and Antheidosorus, were evidently published within a few days of each
other. A copy of the Bull. Soc. Imp. Naturalistes Moscou 24/1 was lodged with the Censor on
17 January 1851 (Old Style Calendar) and presented to the Society on 15 March 1851 (O.S.) which
is equivalent to 27 March 185 1 (New Style). I have taken the date on which it was presented to the
Society as being the date of publication (see Stafleu 1969,Marchant 1989). According to Stafleu and
Cowan ( 1 979), p.98 of Hooker’s J. Bot. Kew Card. Misc. vol. 3 was published in April 185 1 and p. 174
in June 1851.

The spelling Gilberta (and not Gilbertia) was deliberately used by Turezaninow in order that it
could be better distinguished from Gilibertia Ruiz & Pav., a genus in the Araliaceae.

The Triptilodiscus group
Triptilodiscus Turez.

Triptilodiscus Turez., Bull. Soc. Imp. Naturalistes Moscou 24/2:66(Aug.-Nov. 1851). Type:
T. pygmaeus Turez.

Dimorpholepis A. Gray in Hooker, Icon. PI. 9:t.856(Sept.-Nov. 1 85 1); A. Gray, Hooker’s J. Bot. Kew
Card. Misc. 4:227(Aug.l852). Type: D. australis A. Gray.

Duttonia F. Muell., Linnaea 25:409(1853). Type: D. sessiliceps F. Muell.

Annual herb with several major axes, sparsely pilose with slender hairs. Leaves alternate linear
to narrow-oblong. Capitula heterogamous, terminating branches and subtended by c. 5 leaves of
which one or more may bear a shoot that continues growth to give a monochasial conflorescence.
Involucre subglobular c. 6 mm high, not radiant. Outer bracts scarious, hyaline, narrow-elliptic, ±
equal to head, long-ciliate with linear stereome. Inner bracts narrow-oblong, slightly exceeding outer

Paul G. Wilson, Helipterum: Part 1

421

bracts, cartilaginous and prominently thickened with narrow scarious shortly ciliate margin and
scanous ciliate apex, somewhat boat-shaped, hispidulous abaxially. Receptacle broad-conical,
smooth, glabrous. Florets discoid, a few outer female, remainder bisexual. Female florets: corolla
very narrow-tubular 2-2.5 mm long, narrow-urceolate at apex, eventually curved outwards, very
shortly 3-dentate, minutely and sparsely glandular pubcrulous with uniseriate hairs: inner cells of
urceolate portion with sinuate margins, lobes densely papillose within; pappus a short corona of
barbellate scales, sometimes with one shortly plumose bristle. Bisexual florets: corolla narrow-
tubular , urceolate above, minutely 4-dentate, c. 2.5 mm long, at first erect eventually curved outwards,

otherwise as in female florets; anther-appendage oblong, very thin, cells narrow-oblong, un thickened;
anther tails filamentous, shorter than collar; style apex truncate; achenenarrow-cylindrical,c. 1.8 mm
long, minutely puberulous with 2-celled tooth-like hairs rounded at apex; pericarp diaphanous; testa
leathery, smooth, without crystals; boss a minute narrow ring; pappus bristles 3-4, linear-acuminate,
shortly plumose, c. 2.5 mm long, persistent,

Triptilodiscus pygmaeus Turcz., l.c. - Helipterum. pygmaeum (Turcz.) Druce in Hayward & Druce,
Advent. FI. Tweedside 103(1919) comb, illeg. Type: Western Australia,y. Drummond 5th coll, n.54
(iso; MEL 109204).

Dimorpholepis australis A. Gray, l.c. - Helipterum dimorpholepis Benth., FI. Austral. 3:650(1867)
nom. illeg. -ArgyrocomedimorpholepisKmtz^,KQV\s. Gen. PI. Helipterum australe
(A. Gray) Druce, Bot.Exch. Club Brit. Isles 1916,4:627(1917). Type citation: 'Hab. South-western
Australia, Drummond. Also in the interior of Eastern Australia, at Bathurst Plains, Fraser; and
Nangers, Captain M'Arthur.' SyntypetJ. Drummond 54 (K, photo seen).

Duttonia sessiliceps F. Muell., Linnaea 25:410(1853). Type citation: 'In pratis prope rivum Rocky-
creek.' Lectotype (here chosen); Rocky Creek, Oct. 1851, F. Mueller (lecto: MEL 109152; isolecto-
MEL 109197).

There is uncertainty over the publication dates of Triptilodiscus Turcz. and Dimorpholepis
A. Gray. The former name was published in the Bull. Soc. Imp. Naturalistes Moscou vol.24 part 2
no. 3 which was sent to the Censor on 15 August 185 1 and presented to the Society on 5 November
1851 (New Style calendar). The latter name was published as tab. 856 of the leones Plantarum which
appeared between September and November 1851 or possibly later, //Jc C. Jeffrey in litt. I have
adopted the Turezaninow name, as did Laurie Haegi (1986), since there is the greater likelihood of
it having priority.

This genus is closely related to Haegiela which shares the same branching habit of the
con florescence, as well as the .same hair type, and similar corolla and achene characters. Haegiela
differs principally in the absence of a pappus, the presence of crystals in the testa, and the presence
of unthickened inner involucral bracts.

Haegiela P. Short & Paul G. Wilson

Haegiela P. Short & Paul G. Wilson, Muelleria 7:259(1990). Type: Haegiela tatei (F. Muell.)
P. Short & Paul G. Wilson

For description and discussion see Short & Wilson, op.cit.

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Nuytsia Vol. 8, No. 3 (1992)

Pterochaeta Steetz

Pterochaeta Steetz in Lehm., PI. Preiss. 1:456(1845). Type: Pterochaeta paniculata Steetz

Annual woolly herb with one to several major axes. Leaves alternate, oblong. Capitula
homogamous, shortly pedunculate, racemosely arranged, subtended by two leafy bracts. Involucre
broad urceolate; bracts multiseriate, long ciliate, woolly, pale yellow; outer bracts ovate, scarious,
attenuate at base with a minute hard green stereome. shortly apiculate with a pale fawn lamina;
intermediate bracts broad-ovate scarious with a narrow-oblong hard claw (stereome) and a short
oblong lamina; innermost bracts very short on a narrow-oblong hard naviculiform claw; receptacle
naked; florets c. 20, outer female, inner bisexual, actinomorphic. Corolla cylindrical, shortly 4-lobed
in female. 5-lobed in bi.sexual, glabrous; lobes glabrous within, thickened on margin; cells of throat
undulate. Anther c. 0.7 mm long, delicate, appendage short, thin; tails filamentous. Style apex
truncate. Achene narrow-ellipsoid, shortly beaked, covered with 2-celled tooth-like papillae;
pericarp thin and weak, transparent, myxogenic, vascular strands laterally placed in relation to the
cotyledons; testa leathery, brown rugulose, cells very thick walled and deeply undulate with scattered
narrow-oblong crystals, vascular strand laterally placed and confined to near base of seed; carpopodium
minute, annular. Pappus equal to corolla; bristles plumose, persistent.

One species endemic to southern Western Australia.

Pterochaeta paniculata Steetz in Lehm., op.cit. 455. - Waitzia paniculata (Steetz) Benth.,
FI. Austral. 3:637(1867). Lcr/otypc (here chosen): In limoso-arenosis planitiei baud longe a praedio
rustico "Maddington",et sinu regisGeorgiillL, Nov.[18]39 et [I8]40. L. Preiss. No.35. (lecto: MEL
1585199; isolecto: LD).

The genus Waitzia differs from Pterochaeta most obviously in habit, in the nature of the anther
apiculum and anther tails, in having a stout deltoid style apex with a vascular strand extending to the
tip, and in the form of the achene. The vascular strands of the pericarp in Waitzia are medially placed
in relation to the cotyledons whereas in Pterochaeta they are laterally placed.

Haptotrichion Paul G. Wilson, gen. nov.

Herba annua erecta pilis .septatis glandulosis pubescens. Folia caulina alterna linearia. Capitula
homogama.solitariaradiantia. lnvolucrumhemisphericum;bracteaec.4-seriatae;bracteaeexteriores
elliptica hyalina longe ciliata, stereomate anguste oblongo piano; bracteae intimae ungue anguste
elliptico longe ciliato, stereomate anguste oblongo, lamina elliptica flava. Receptaculum minute
glanduloso papillosum. Flosculinumerosi.bisexuales.actinomorphi. Corolla tubo graciliglanduloso
piloso, limbo campanulalo glabro; lobi 5, ovati, nervis ad apicem ex tensis. Antherae: appendix ovatus
tenuis, cellulis marginalibus manifeste discretis, cellulis basalibus ± equilateralibus, cetera anguste
oblongis; caudae filamento.sae collum superantes. Styli apex truncatus. Achenium angusto
ellipticum compressum, ad apicem in rostro productum, papillis 2-cellulis dcntoidcis ornatis;
pericarpium: stratum exterius cellulis cubicisparietibustcnuibus; stratum interiuscellulisincrassatis,
lignosis, sclerenchymatis; testa tenuissima, nervo circumnexo. Pappus persistens, setis filiformibus,
denticulatis, basi versus in tubo conjunctis.

Typus: Haptotrichion conician (B. Turner) Paul G. Wilson

Paul G. Wilson, Helipterum: Part 1

423

Annual erect herbs pubescent with gland-tipped septate hairs. Leaves cauline, alternate, linear.
Capitula solitary and terminal to long branches, radiant. Involucre hemispherical; bracts c. 4-seriate;
outer bracts elliptic, hyaline, long-ciliate with a brown nanow-oblong flat stereome; innermost bracts
with a narrow-elliptic hyaline long-ciliate claw, a narrow-oblong stereome, and a yellow elliptic
lamina. Receptacle minutely glandular papillose. Florets numerous, bisexual, actinomorphic.
Corolla: tube slender, glandular pilo.se; limb campanulatc, glabrous; lobes five, ovale, vascular
strands extending to tips. Anther; appendage ovate, thin, marginal cells differentiated, basal cells +
equilateral, medial and distal cells narrow-oblong; tails slender delicate, extending to beyond collar.
Style apex truncate; vascular strand prominent, extending almost to tip. Achene narrow-elliplic,
compressed, extending into a terminal beak, minutely papillose with 2-celled tooth-like trichomes;
carpopodium annular, short; pericarp: outer layer of 1-cell thick layer of thin-walled cubical cells;
inner layer of 2-3 rows of thick walled sclerenchymatous cells; testa extremely thin, more or less
adherent to pericarp; endosperm thin, free from testa; vascular strands of pericarp lateral (in relation
to cotyledons); vascular strand of testa passing almost completely around seed in lateral position.
Pappus bristles filiform, barbellate, united below to form a cup-shaped base, entire or slit on one side,
persistent.

Haptotrichion conicum had been placed in Waitzia since it possessed beaked achenes. It differs
from Waitzia most obviously in 1) having a different type of indumentum (not cottony), 2) having
broad flat hyaline claws to the inner involucral bracts (not terete and firm), 3) in having truncate style
apices (neither ellipsoid nor deltoid), and 4) in having delicate filamentous anther tails (not slender
and firm). Haptotrichion further differs from Waitzia in the anatomy of the achene for its pericarp
has an outer single-celled layer of thin-walled cube like cells and an inner layer of several rows of
thick- walled sclerenchymatous cells, while the testa is extremely thin and delicate. In Waitzia the
pericarp consi.sts of a thin layer of hyaline linear cells while the testa is thick, rugose, and made up
of thick-walled collenchyma. The vascular strand in the testa of Haptotrichion passes almost
completely around the seed but in Waitzia it is confined to the base.

From Pterochaeta this genus differs in the manner of branching of the stem and inflorescence
(q.v.), in the nature of the involueral bracts, the nature of the corolla, and the nature of the anther
apiculum. In Pterochaeta, Haptotrichion and Waitzia the achenes are beaked and bear tooth-like
2-celled trichomes but in other chiuracters the genera are very distinct.

The genus consists of two species, both endemic to the Carnarvon District (Beard 1 980) of Western
Australia.

Haptotrichion colwillii Paul G. Wilson, sp. nov. (Figure 6)

Herba erecta ad 25 cm alta. Rami glanduloso puberuli sub capitula pilosi. Folia linearia, 8-15
mm longa, glanduloso puberula. Involucrum hemisphaericum, c. 10 mm altum et latum; bracteis
exterioribus late ellipticis, hyalinis, ciliatis; bracteis intimis ungue anguste ciliptico, c. 9 mm longo,
hyalino, ciliato, limbo anguste obovaloobtuso, 9-15 mm longo, luteo. Corolla c. 5 mm longa, tubo
glanduloso hirtello. limbo anguste turbinato, glabro, intra sub lobis papilloso. Achenium anguste
ellipsoideum c. 2.5 mm longum, denticulalum; rostrum gracile, laeve, ad 3 mm longum. Pappus
persistens, .setis in basi connatis, tubiformibus.

TypMS.- 7 km S of Overlander Roadhouse [c. 170 km S of Carnarvon], Western Australia, 1 Sept. 1985,
H. Demarz 10702 (holo: PERTH).

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Nuytsia Vol. 8, No. 3 (1992)

Figure 6. Haptotrichion colwillii. A - Habit. B - Florets. C - Pappus bristle apices. D - Hair of corolla. E - Style apex.
F- Outer, intermediate, and innerinvolucral bracts. G - Anther. H- Achene with pappus. I- Achene. J- Achene papillae.
K - Capitulum. L & M - Hairs of stem. N - Leaf. From H. Demarz 10702.

Paul G. Wilson, Helipterum: Part 1

425

Erect herb to 25 cm high, branching at base. Principal axes slender, to 20 cm high, simple or
sparsely branched, glandular puberulous, pilose beneath capilulum. Leaves linear, 8-15 mm long,
c.0.7 mm wide, glandular puberulou.s,marginsslightlyrecurved. Involucre hemispherical, c. 10mm
high and wide. Outer bracts broad-elliptic, hyaline, ciliale; stereome narrow-triangular, brown,
glandular puberulous. Innermost bracts: clawnarrow-elliptic,c. 9 mm long, hyaline, ciliate, stereome
very narrow-oblong, glabrous except for the glandular puberulous apex; limb narrow-obovate, obtuse,
9-15 mm long, yellow. Corollac. 5 mm long; tube narrow, glandular hirtcllous with multicelled hairs;
limb narrow-turbinate, glabrous, shortly 5-lobed, papillose within below junction of lobes. Achene
compressed narrow-ellipsoid, c. 2.5 mm long, barbellate with 2-celledpapillae; beak slender, smooth,
to 3 mm long. Pappus more or less equal to corolla, persistent, bristles united towards base to form’
a tube c. 2 mm long.

Additional specimen examined. WESTERN AUSTRALIA: Hamelin Pool, 16 Sept 1984 J Colwill
s.n. (PERTH).

Habitat. Found on red sand over limestone (fide H. Demarz in sched.).

Distribution. Only known from near Hamelin Pool, c. 170 km south of Carnarvon, Western Australia.

Notes. Haptotrichion colwillii is very similar to //. conicum which is known from an area between
Carnarvon and Gascoyne Junction. The latter species differs most obviously in 1) the leaves ending
in a rounded 'capitate' tip, 2) the receptacle being narrow-conical (not rounded), and 3) the pappus
sheath being split along one side.

Haptotrichion colwillii is named after John Colwill, a Western Australian horticulturalist who has
a particular interest in the native Asteraceae and who first collected this species.

The illustration is drawn from a plant raised from seed collected by Herbert Demarz and grown
at Kings Park Botanic Garden, Perth. I should like to thank the staff of Kings Park for their unstinted
assistance in my study of this and other annual species of the Rhodanthe complex.

Haptotrichion conicum (B. Turner) Paul G. Wilson, comb. nov.

Wai7zmco/!/caB.Turner,Sida2:428(I966). Ty/jc.- 11 miles west of Gascoyne Junction, 24 Aug. 1965,
BL. Turner 5405 (holo: MEL 598263; iso; MEL 598265, PERTH).

[Waitziapodolepis auct. non (Gaud.) Benth.: F. Muell., Zeitschrift des allgem . osterreich. Apotheker-
Vereines 34 (no.36):933-936 (1896); F. Muell., PI. Indig. Sharks Bay 16(1883).]

Distribution. Between Carnarvon and Gascoyne Junction, Western Australia.

A de.scription of this species was provided by Mueller (1883) based on a collection made by
J. Polak (or Pollack) in 1882 from the Ga.scoyne River region (MEL 1584941). Mueller assumed that
the plant he was describing was Waitziapodolepis (Gaudich.) Benth.: 'this plant represents evidently
the genuine species, illustrated by Gaudichaud', even though Mueller had seen no authentic material
of that species. In 1 896 Mueller again included the Pollack specimen under W. podolepis but without
comment. In 1905 Diels and Pritzel slated that the Pollack collection was not W. podolepis, however,
they did not give it a name.

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Nuytsia Vol. 8, No. 3 (1992)

Acknowledgements

I thank the various Australian herbaria who sent on loan much of their material of Helipterum for
study. Kings Park Botanic Garden allowed me to have access to their live collection and provided
me with seed of a n umber of species. S teve Hopper kindly searched for and collected from different
populations of rare Rhodanthe species near Shark Bay. Photographs of relevant Asteraceae types in
herb. KW were made available to me by my colleague Neville Marchant. Discussions and
correspondence with Philip Short and Arne Anderberg have been stimulating and of considerable
assistance. The drawings were prepared with much care by Margaret Menadue.

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Erickson, R. (1969). "Tlie Drummonds of Hawthomdcn." (Lamb Patterson, Perth.)

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3:172-178.

Haegi,L. (1986). Helichrysum Miller, /n Jessop, J.P. & Toelken, H.R. "Fora of South Australia." (South Australian Government,
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Hilliard, O.M. (1983). "Flora of Southern Africa." Vol. 33 Asteraceae part 7 Inuleae fasc. 2 Gnaphaliinae. 1-325. (Department
of Agriculture, Pretoria.)

Hilliard, O.M. & Burlt, B.L. (1981). Some generic concepts in Compositae-Gnaphaliinae. J. Linn. Soc., Bot. 82:181-232.
Hnatiuk, R.J. (1990). Census of Australiiur vascular plants. (Bureau of Flora and Fauna, Canberra.)

Marchant, N.G. (1990). The contribution of the Russian botanist Turczaninov to Australian plant taxonomy. In P.S. Short (ed),
"History of systematic botany in Australasica."(Australian Systematic Botany Society Inc., Melbourne.)

Nordenstam, B. (1989). A synopsis of the genus Syncarpha (Compositae-Gnaphalieae). Compositae Newsletter No 17:2-6.

Short, P.S. (1983). A revision o[ Angianlluis Wendl., sensu lato (Compositae: Inuleae: Gnaphaliinae), 1. Muelleria 5:143-183
(1983).

Short, P.S. (1989). A revision of Podotheca Cass. (Asteraceae: Inuleae: Gnaphaliinae). Muelleria 7:39-56.

Short, P.S. & Sinkora, D.M. (1988). The botanist Joachim Steetz (1804-1862). Muelleria 6:449-494.

Short, P.S., Wilson, K.E. & Nailon, J. (1989). Notes on the fruit anatomy of Australian members of the Inuleae (Compositae).
Muelleria 7:57-79.

Stafleu, F.A. (1969). Turczaninow’s Flora Baicalensi Dahurica. Taxon 18:563-565.

Stafleu.F.A. & Cowan, R.S. (1979), "Taxonomic literature, a .selective guide to botanical publications and collections with dates,
commentaries and types." Vol, 2. (Bonn, Scheltema & Holkema, Utrecht.)

Turner, B.L. (1970). Chromosome numbers in the Compositae. Xll Australian species. Amer. J. Bot. 57:382-389.

Warcup, J.H. (1990). Tlie mycorrhizal associations of Australian Inuleae (Asteraceae). Muelleria 7:179-187.

Wilson, P.G. (1989a). A revision of the genus Hyalosperma (Asteraceae: Inuleae: Gnaphaliinae). Nuytsia 7:75-101.

Wilson, P.G. (1989b). Erymophyllum (Asteraceae: Inuleae: Gnaphaliinae), a new Australian genus in the Helipierum complex.
Nuytsia 7:103-1 16.

Wilson, P.G. (1992a). The classification of Australian species currently included in Helipterum (Asteraceae: Gnaphalieae): Part
2 Leucochrysum. Nuytsia 8:439-445.

Wilson, P.G. (1992b). Tlie classification of the genus Wailzia Wendl. (Asteraceae: Gnaphalieae). Nuytsia 8:461-477.

Wilson, P.G., Short, P.S., & Orchard, A.E. (1992). Some nomencltitural ch.'inges in the Angianthinae and Cassiniinae
(Asteraceae: Gnaphalieae). Muelleria 7:519-524.

Paul G. Wilson, Helipterum: Part 1

427

Index to Scientific Names

New names are given bold type. Other accepted names are in roman. Synonyms are in italics.
Acroclinium A. Gray 386

multicaule A. Gray 387
phyllocephalum A. Gray 394
roseum A. Gray 387

var. album G. Nicholson 387
var. grandiflorum G. Nicholson 387
rubellum A. Gray 398
Anisolepis Steetz 415

pyrethmm Steetz 416
Antheidosorus A. Gray 419
gracilis A. Gny 419
Argyrocome Gaertn.

charsleyae (F. Muell.) Kuntze 413
chlorocephala (TuTcz.)Kmlz£ 386
condensafa (F. Muell.) Kuntze 401
co/7TOZ)//7ora (Schldl.) Kuntze 391
corymbosa (A. Gray) Kuntze 401
dimorpholepis (Benth.) Kuntze 421
floribunda (DC.) Kuntze 408
forrestii{F. Muell.) Kuntze 401
frenchii (F. Muell.) Kuntze 415
fuscescens (Turcz.) Kuntze 394
haigii (F. Muell.) Kuntze 401
heterantha (Turcz.) Kuntze 412
humboldtiana (Gaudich.) Kuntze 396
laevis (A. Gray) Kuntze 402
nunglesH (Lindley) Kuntze 384
margarefhae (F. Muell.) Kuntze 415
moschala (DC.) Kuntze 385
polycephala (A. Gray) Kuntze 404
polygalifolia (DC.) Kuntze 397
polyphylla (F. Muell.) Kuntze 417

428

NuytsiaVol. 8, No. 3 (1992)

pygmaea (DC.) Kuntze 398
pyrethrum {SXeciz) Kmize 416
rosea (Hook.) Kuntze 387
rubella (A. Giay) Kuntze 398
spicata{¥. Muell.) Kuntze 413
sterilescens (F. Muell.) Kuntze 409
stricta (Lindley) Kuntze 399
(ielkensii (F. Muell.) Kuntze 406
Calocephalus

glohosus Hutch. 414
gnaphalioides Hook. 385
Cassinia

cuprea F. Muell. 399, 404
Cephalipterum A. Gray 417

drummondii A. Gray 417

f. major Diels & E. Pritzel 418
f. minor Diels & E. Pritzel 418
Dimorphotepis A. Gray 420
australis A. Gray 421
Duttonia F. Muell. 420

sessilicepsF. Muell. 421
Erymophyllum Paul G. Wilson 418

gracile (A. Gray) Paul G. Wilson 418
Epaltes Cass.

latei F. Muell. 421
Gilberta Turcz. 419

tenuifolia Turcz. 419
Gnaphalium

moschatum DC. 385
Grijfithia J. Black 386

helipteroides J. Black 397
Haegiela P. Short & Paul G. Wilson 421

tatei (F. Muell.) P. Short & Paul G. Wilson 421

Paul G. Wilson, Helipierum: Part 1

429

Haptotrichion Paul G. Wilson 422

colwillii Paul G. Wilson 423, 424
conicum (B. Turner) Paul G. Wilson 425
Helichrysum Miller

sect. Acroclinium (A. Gray) Baillon 386
sect. Rhodanthe (Lindley) Baillon 383
anthemoides Sprengel 386
drummondii F. Muell. 387
gray/F. Muell. 387
huniboldtianum Gaudich. 396
manglesii (Lindley) Baillon 384
obtusifolium Sander 381
oldfieldii F. Muell. 407
Heliptemm DC.

sect. Ag/ossMm F. Muell. 412

sect. Helipteridium A. Gray 412

sect. Leiochrysum DC. 385

sect. Monencyanthes (A. Gray) Benlh. 384

sect. P achy pterum Sleetz 419

sect. Pleropogon (DC.) Benth. 385

sect. Sericophorum DC. 385, 408

sect. Synachyrum A. Gray 408

adpressum W. Fitzg. 436

albicans (Cunn.) DC. 436

album (Nicholson) L. Bailey 387

album Ewart 401

anactinumF. Muell. 412

anthemoides (Sprengel) DC. 386

austral e (A. Gray) Druce 421

battii F. Muell. 413

charsleyae F. Muell. 413

chionolepis F. Muell. 409

chlorocephalum (Turcz.) Benth. 386

cirratum Morrison 409

430

Nuytsia Vol. 8, No. 3 (1992)

clementii Domin 396
condensatumV. Mvi&W. 401
corymbiflorum Schldl. 391

\ai microglossa Benlh. 396
corymbosum (A. Gray) Benth. 401
colu/a (Benth.) DC. 419
craspedioides W.V. Fitzg. 436
cryptanthum O. Sarg. 384
dijfusum DC. 394
dimorpholepis Benth. 421
discoideum A. Gray 412
drummondii (A. Gray) Ostenf. 398
fitzgibboniiY. Mn&W. 436
floribundum DC. 408. 409

var. stuartianum (Sond.) Benth. 411
var. tubulipappum J. Black 409
F. Muell. 401
/7(?«c/ii7 F. Muell. 415
fuscescens Turcz. 394
/la/gn F. Muell. 401
heteranthim Turcz. 412

var. minor Ewart 412
var. majus Bcnih. 412
humboldlianum (Gaudich.) DC. 396
laeve (A. Gray) Benth. 402
largiflorens F. Muell. 396
manglesii (Lindley) Benth. 384

var. nuiculalum (J. Drumm. ex hort.) L.H. Bailey 384
margarethae F.MueW. 415
margarilae F.MucW. 415
maryo/u'/ S. Moore 385
microglossa (Benth.) Tate 396
microglossum Maiden & E. Betche 396
mode (DC.) Paul G. Wilson 437

Paul G. Wilson, Helipterum: Part 1

431

monencyanthoiclesY. MueW. 414
moschatum (DC.) Benth. 385
mullinense S. Moore 401
niveum Steetz 437
oppositifolium S. Moore 397
pachychaetum W. Fitzg. 4 1 2
polycephalum (A. Gray) Benth. 404
polygalifoUum DC. 397

var. leucactinum F. Muell. 394
polyphyllum¥.y[\it\\. 417
propinguiim W. Fitzg. 398
pterochaetum (F. Muell.) Benth. 437
pane ta turn DC. 386
pygmaeum (DC.) F. Muell. 398

var. drummondii (A. Gray) Ostenf. 398
var. occideniale Benth. 398
pygmaeum (Turez.) Druce 421
/tyret/irwm (Steetz) Benth. 416
roseum (Hook.) Benth. 387

f. album (Nicholson) Moldenke 387
var. album Guilfoyle 387
var. album Ewart el al. 388
var. grandiflorum (Nicholson) Chitt. 387
var. nigropapposum Ostenf. 387
vtu-. patens (Ewart el al.) J. Black 388, 411
rubellum (A. Gray) Benth. 398
sandfordii Hook. 396
s/r/cfltMw (Steetz) Benth. 413
var. /;a//crtj Benth. 413
splendidum Hemsley 388
sterilescens F. Muell. 409
stipitatum (F. Muell.) Benth. 438
strictum (Lindley) Benth. 399

var. stenocephala S. Moore 399

432

Nuytsia Vol. 8, No. 3 (1992)

stuartianum Sond. 411
tietkensii F. Muell. 406
iroedelii F. Muell. 411

var patens Ewart et al. 388,411
uniflorum J. Black 385
verecundumS. Moors, 420
Hyalosperma Steetz 419

glutinosum Steetz 419
Leptorhynchos Less.

citrinus Benth. 407
Monencyanlhes A. Gray 384

gnaphalioides (Hook.) A. Gray 384, 385
Myriocephalus Benth.

graci/A (A. Gray) Benth. 419
Podosperma Labill.

fuscescens (Turcz.) F. Muell. 394
pollackii F. Muel\. 413
Podotheca Cass.

fuscescens (Turcz.) Benth. 394
pollackii (F. Muell.) Diels 413
Pterochaeta Steetz 422

paniculata Steetz 422
Pteropogon DC. 385

sect. Achyroclinoides A. Gray 399
sect. F acelioides A. Gray 385
sect. Helichrysoides A. Gray 412
sect. Helipteroides A, Gray 418
sect. Pteropogonopsis A. Gray 419
australis Nees 398
corymbosus A.. Grny 401
demissus A. Gray 419
drummondii A. Gray 398
humboldtianus (Gaudich.) F. Muell. 396

Paul G. Wilson, Heliplerum: Part 1

433

intermecHus F, Muell. 398
laevis A. Gray 402
platyphyllus F. Muell. 399
polycephalus A. Gray 414
pygmaeum DC. 398
spicatus SXQtiz 413
Rhodanthe Lindley 383

sect. Achyroclinoides (A. Gray) Paul G. Wilson 399
sect. Actinaria Paul G. Wilson 414
sect. Anisolepis (Sleetz) Paul G. Wilson 415
sect. Citrinae Paul G. Wilson 406
sect. Helichrysoides (A. Gray) Paul G. Wilson 412
sect. Helipteridium (A. Gray) Paul G. Wilson 412
sect. Leiochrysum (DC.) Paul G. Wilson 385
sect. Monencyanthes (A. Gray) Paul G. Wilson 384
sect. Polyphyllum Paul G. Wilson 416
sect. Rhodanthe 383

sect. Synachyrum (A. Gray) Paul G. Wilson 408
anthemoides (Sprengel) Paul G. Wilson 386
ascendens Paul G. Wilson 400
atrosanguinea J. Drumni. ex hort. 384
battii (F. Muell.) Paul G. Wilson 413
charsleyae (F. Muell.) Paul G. Wilson 413
chlorocephala (Turcz.) Paul G. Wilson 386
subsp. chlorocephala 387
subsp. rosea (Hook.) Paul G.Wilson 387
subsp. splendida (Hemsiey) Paul G. Wilson 388
citrina (Benth.) Paul G. Wilson 407
collina Paul G. Wilson 389, 390
condensata (F. Muell.) Paul G. Wilson 401
corymbiflora (Schldl.) Paul G. Wilson 391
corymbosa (A. Gray) Paul G. Wilson 401
cremea Paul G. Wilson 391, 393

434

Nuytsia Vol. 8, No. 3 (1992)

diffusa (DC.) Paul G. Wilson 394
var. diffusa 394

var. leucactina (F. Muell.) Paul G Wilson 394
floribunda (DC.) Paul G. Wilson 408
forrestii (F. Muell.) Paul G. Wilson 401
frenchii (F. Muell.) Paul G. Wilson 415
fuscescens (Turcz.) Paul G. Wilson 394
gossypina Paul G. Wilson 395
haigii (F. Muell.) Paul G. Wilson 401
heferantha (Turcz.) Paul G. Wilson 412
humboldtiana (Gaudich.) Paul G. Wilson 396
laevis (A. Gray) Paul G. Wilson 402
maculata J. Drumm. ex hort. 384
manglesii Lindley 384

var. Regel 384
subvar. alba hort ex Vilm. 384
var. ntaculala (J. Drumm. ex hort.) Vilm. 384
var. maculata Hook. 384
var. sail guinea Hook. 384
margarethae (F. Muell.) Paul G. Wilson 415
tnaryonii (S. Moore) Paul G. Wilson 385
microglossa (Maiden & E. Betche) Paul G. Wilson 396
moschata (DC.) Paul G. Wilson 385
nullarborensis Paul G. Wilson 402, 403
oppositifolia (S. Moore) Paul G. Wilson 397
subsp. oppositifolia 397
subsp. ornata Paul G.. Wilson 379
pollackii (F. Muell.) Paul G. Wilson 413
polycephala (A. Gray) Paul G. Wilson 404
polygalifolia (DC.) Paul G. Wilson 397
polyphylla (F. Muell.) Paul G. Wilson 417
propinqua (W. Fitzg.) Paul G. Wilson 398
psammophila Paul G. Wilson 404, 405
pygmaea (DC.) Paul G. Wilson 398

Paul G. Wilson, Helipterum: Part 1

435

pyrethrum (Stectz) Paul G. Wilson 416
rubella (A. Gray) Paul G. Wilson 398
rufescens Paul G. Wilson 398
sphaerocephala Paul G. Wilson 409
spicata (Steetz) Paul G. Wilson 413
sterilescens (F. Muell.) Paul G. Wilson 409
stricta (Lindley) Paul G. Wilson 399
stuartiana (Sond.) Paul G. Wilson 411
tietkensii (F. Mucll.) Paul G. Wilson 406
troedelii (F. Muell.) Paul G. Wilson 411
uniflora (J. Black) Paul G. Wilson 385
Roccardia Voss 383

anthemoides (Sprengel) Voss 386
c/)/or£)cc/;/)a/a (Turcz.) Voss 387
corymbiflora (Schldl.) Voss 391
humboldtiana (Gaudich.) Voss 396
manglesii (Lindley) Voss 384
f. alba Voss 384
f. ligulosa Voss 384
f. maculata Voss 384
f. sanguinea Voss 384
rosea (Hook.) Voss 387
rubella (A. Gray) Voss 398
Schoenia Steetz

chlorocephala Turez. 386
huniboldliana (Gaudich.) Walp. 396
Triptilodiscus Turez. 420
pygmaeus Turez. 421
Waitzia Wendl.

brevirostris Steetz 407
citrina (Benth.) Steetz 407
conica B. Turner 425
dasycarpa Turez. 407
paniculata (Steetz) Benth. 422

436

Nuytsia Vol. 8, No. 3 (1992)

steetziana Lehm. 407
sulphurea Steetz 407
tenella Hook. 407
Xyridanthe Lindley 386

stricta Lindley 399

Name changes in Australian species of Helipte rum (see also Wilson 1989a, b and 1992a,b)

Taxa listed in Hnatiuk (1990)

H. adpressum
H. albicans

H. albicans subsp. albicans
H. albicans var. biiffaloensis

H. albicans var. incanum

H. albicans subsp. alpinum

H. albicans var. graminifolium
H. anthemoides

H. australe
H. battii
//. charsleyae

H. chlorocephalum

H. condensatum

H. corymbiflorum
H. corymbosum
H. cotula
H. craspedioides
H. demissum
H. dijfusum

H. fitzgibbonii

Taxa accepted by Wilson

Chrysocephalum puteale (S. Moore) Wilson

Leucochrysum albicans (A.Cunn.) Wilson

Leucochrysum albicans subsp. albicans

Leucochrysum albicans var. buffaloensis (Wilson)
Wilson

Leucochrysum albicans subsp. albicans var. tricolor
(DC.) Wilson

Leucochrysum albicans subsp. alpinum (F. Muell.)
Wilson

Leucochrysum graminifolium (Wilson) Wilson

Rhodanthe (Leiochrysum) anthemoides (Sprengel)
Wilson

Triptilodiscus pygmaeus Turcz.

Rhodanthe (Helichrysoides) battii (F. Muell.) Wilson

Rhodanthe (Helichrysoides) charsleyae (F. Muell.)
Wilson

Rhodanthe (Leiochrysum) chlorocephala (Turcz.)
Wilson subsp. chlorocephala

Rhodanthe (Achyroclinoides) condensata (F. Muell.)
Wilson

Rhodanthe (Leiochrysum) corymbiflora (Schldl.) Wilson

Rhodanthe (Achyroclinoides) corymbosa (A. Gray) W ilson

Hyalosperma cotula (Benlh.) Wilson

Possibly referrable to Polycalymma

Hyalosperma demissum (A. Gray) Wilson

Rhodanthe (Leiochrysum) diffusa (Cunn. ex DC.) Wilson
var. diffusa

Leucochrysum fitzgibbonii (F. Muell.) Wilson

Paul G. Wilson, Helipterum: Part 1

437

H. floribundum

Rhodanthe (Synachyrum) floribunda (DC.) Wilson

H. forrestii

Rhodanthe (Achyroclinoides) forrestii (F. Muell.) Wilson

H. frenchii

H. fuscescens

Rhodanthe (Actinaria) frenchii (F. Muell.) Wilson

Rhodanthe (Leiochrysum) fuscescens (Turcz.) Wilson

H. gracile

Erymophyllum tenellum (Turcz.) Wilson

H. haigii

Rhodanthe (Achyroclinoides) haigii (F. Muell.) Wilson

H. heteranthum

Rhodanthe (Helipteridium) heterantha (Turcz.) Wilson

H. humboldtianum

Rhodanthe (Leiochrysum) humboldtiana (Gaudich.)
Wilson

H. hyalospermum

Hyalosperma glutinosum Steetz subsp. glutinosum

H. involucratum

Erymophyllum ramosum (A. Gray) Wilson subsp.
involucratum (F. Muell.) Wilson

H. jessenii

Hyalosperma semisterile (F. Muell.) Wilson

H. laeve

Rhodanthe (Achyroclinoides) lacvis (A. Gray) Wilson

H. manglesii

Rhodanthe (Rhodanthe) manglesii Lindley

H. margarethae

H. maryonii

Rhodanthe (Actinaria) margarethae (F. Muell.) Wilson

Rhodanthe (Monencyanthes) maryonii (S. Moore) Wilson

H. microglossum

Rhodanthe (Leiochrysum) microglossa (Maiden & Betche)
Wilson

H. molle

Leucochrysum molle (Gunn, ex DC.) Wilson

H. moschatum

Rhodanthe (Monencyanthes) moschata (Cunn. ex DC.)
Wilson

H. niveum

Distinct genus; affinities with Helichrysum obtusifolium
Sond.

H. oppositifolium

Rhodanthe (Leiochrysum) oppositifolia (S . Moore) W ilson

H. polycephalum

Rhodanthe (Achyroclinoides) polycephala (A. Gray)
Wilson

H. poly gal ifolium

Rhodanthe (Leiochrysum) polygalifolia (Cunn. ex DC.)
Wilson

H. polyphyllum

Rhodanthe (Polyphyllum) polyphylla (F. Muell.) Wilson

H. praecox

Hyalosperma praecox (F. Muell.) Wilson

II. propinquum

H. pterochaetum

Rhodanthe ((Leiochrysum) propinqua (W. Fitzg.) Wilson

Chrysocephalum pterochaetum F. Muell.

H. pygmaeum

Rhodanthe (Leiochrysum) pygmaea (DC.) Wilson

II. pyrethrum

Rhodanthe (Anisolepis) pyrethrum (Steetz) Wilson

H. roseum

Rhodanthe (Leiochrysum) chlorocephala subsp. rosea
(Hook.) WiLson

Nuylsia Vol. 8, No. 3 (1992)

438

H. rubellum
H. saxatile
H. semisterile
H. spicatum
H. splendidum

H. sterilescens
H. stipitatum
H. stoveae
H. strictum
H. stuartianum
H. tenellum
H. tietkensii
H. troedelii
H. uniflorum
H. venustum

H. zacchaeus

Rhodanthe (Leiochrysum) rubella (A. Gray) Wilson
Distinct genus

Hyalosperma semisterile (F. Muell.) Wilson

Rhodanthe (Helichrysoides) spicata (Steetz) Wilson

Rhodanthe (Leiochrysum ) chlorocephala subsp. splendida
(Hemsley) Wilson

Rhodanthe (Synachyrum) sterilescens (F. Muell.) Wilson

Leucochrysum stipitatum (F. Muell.) Wilson

Hyalosperma stoveae (D.A. Cooke) Wilson

Rhodanthe (Leiochrysum) stricta (Lindley) Wilson

Rhodanthe (Synachyrum) stuartlana (Sond.) Wilson

Erymophyllum tenellum (Turcz.) Wilson

Rhodanthe (Achyroclinoides) tietkensii (F. Muell.) Wilson

Rhodanthe (Synachyrum) troedelii (F. Muell.) Wilson

Rhodanthe (Monencyanthes) uniflora (J. Black) Wilson

Hyalosperma glutinosum Steetz subsp. venustum
(S.Moore) Wilson

Hyalosperma zacchaeus (S. Moore) Wilson

Nuytsia 8(3); 439-446 (1992)

439

The classification of Australian species currently included in
Helipterum (Asteraceae: Gnaphalieae): Part 2 Leucochrysum

Paul G. Wilson

Western Australian Herbarium, Department of Conservation and Land Management,
PO Box 104, Como, Western Australia 6152

Abstract

Wilson, Paul G. The classification of Australian species currently included in Helipterum
(Asteraceae: Gnaphalieae): Peat 2 Leucochrysum. Nuytsia 8(3): 439-446 (1992). A new endemic
Australian genus, Leucochrysum (DC.) Paul G. Wilson, is described to accommodate Helipterum
stipitatum, H.fitzgibbonii, and most of the taxa previously referred to the H. albicans group. New
combinations arc effected to accommodate the five species and five infraspecific taxa.

Introduction

The application of the name Helipterum DC. was discussed in a previous paper (Wilson 1989),
while the fact that it is an unnatural genus as currently circumscribed has been indicated by a number
of workers in the group (e.g . Hilliard & B urtt 1981, Anderbcrg 1 989, 1 99 1 .Wilson 1989). In this paper
I have segregated from Helipterum some .species that are closely related to each other and that 1
consider to be clearly generically distinct from other species currently included in Helipterum or
Helichrysum, or in any of the genera recently separated from them. Since this paper is a precursor
to a treatment in the Flora of Australia, and since only new combinations are involved, descriptions
are not provided for the .specific or infraspecific taxa.

Historical

When discussing the rationale for accepting the as then unpublished name Helipterum, that had been
proposed in litt. by Augustin de Candolle, Bentham (1837) suggested that Helichrysum colula Bcnth.,
H. anthemoides Spreng., andpossibly H. incanum Hook. , should be placed in Helipterum , either treating
it as a section of Helichrysum or as a new genus. De Candolle (1838) subsequently published the
name Helipterum as a genus and partly followed Bentham’s suggestion of species affinities by
placing H. incanum and H. cotula in his new section Leucochrysum along with two other species
that are now recognised (Wilson 1960) as being conspecific with H. incanum. Bentham (1867)
placed these species and a number of others, including H. stipitatum, in Helipterum sect.
Euhelipterum. Subsequent Australian botanists have included in Helipterum, without reference to a

440

Nuytsia Vol. 8, No. 3 (1992)

section, all the species here placed in Leiicochrysiim. Recently Anderberg (1991) noted that most of
the Australian species of Heliptenim should be placed in Rhodanthe Lindley but he indicated that 'a
group of five species including H.fitzgibbonii, H. albicans. H. stipitatum, //. moUe, and H. saxalilis
share a number of apomorphic features with the genus Waiizia' and 'deserve to be considered in an
overall study of the species of the Waiizia generic group.' 1 agree with Anderberg’s assessment of
relationships (except that I exclude H. saxatile) and recognise the close affinity between Waiizia and
Leucochrysum.

Generic affinities

The genus Leucochrysum differs from other Australian species-groups in Heliptenim in having
stipe-like claws to the involucral bracts (Figure lA), a glabrous, translucent two-layered pericarp
(Short e! al. 1989, Figures 7b, 11c), a firm testa that is distinct from the pericarp, straight firm anther
tails, and abroad deltoid or broad ovoid style apex with a thick vascular strand that forms aclub-shaped
mass in the tip (Figure 1 G). Short e! al. ( 1 989) in a discussion on the fruit anatomy of various members
of the Australian Gnaphalieae noted that the achene anatomy of Heliplerum albicans differed from
other members oi Heliplerum that had been studied and suggested that it and its close relatives should
be recognised as a distinct genus.

The capitula of the Leucochrysum species resemble those found in a number of Australian species
currently placed in Helichrysum, such as Helichrysum elalum DC., H. boormanii Maiden & Betche,
and H . colliniim DC. that form part of the section Blepharolepis. These species also have a terete claw
to the involucral bracts; however, the species differ in having a pericarp with paired myxogenic cells
that are formed by a periclinal division (anticlinal in other species-groups), a thin smooth testa with
narrow-oblong Ihin-wallcd cells, a rounded to long-acuminate style apex with a slender vascular
strand that terminates below the apex, an antherappendage with thick-walled cells, barbellate pappus
bristles (shortly plumose in Leucochrysum), and undivided stereome to the involucral bracts.

The indumentum, involucre, corolla, anthers, and style of Leucochrysum (see below) are
morphologically similar to those found in Waiizia and it is to this genus Ihtxi Leucochrysum is most
closely related, as has been recognised by Anderberg (see above).

The species of Waiizia s.sir. (Wilson 1992) differ in having an elongated neck to the achene,
2-celled achenial papillae in which the lower cell overtops the upper, and pappus bristles that are
barbellate and are shed as a unit from the achene. They also have branched stems with the capitula
terminating the lateral branches.

In a treatment of the Heliplerum albicans group (Wilson 1960) the superficially similar species
Heliplerum saxalile was described. It is not here included in the Heliplerum albicans group since it
differs in having concavo-convex claws to the involucral bracts, a slender cylindrical corolla with
strongly undulate thickened cell walls of the inner epidermis and vascular strands extending to the
apex of the lobes, a pericarp with papillose paired (by ;ui ticlinal division) slime cells, a small rounded
style apex, and laneeolate anther appendages with thick cell walls. It is apparently related to
Helichrysum podolepidium F. Muell. [Chiysocephalumpodolepidium (F. Muell.) Anderberg] whose
taxonomic position in relation to genera in the Heliplerum complex will be discussed in a separate
paper.

Paul G. Wilson, Helipterum: Part 2 Leucochrysum

441

Taxonomic treatment

Leucochrysum (DC.) Paul G. Wilson, gen. et comb. nov.

Helipterum sect. Leucochrysum DC., FYod. 6:215(1838). Lectotype (here chosen); Helipterum
albicans (Cunn.) DC. (see below).

Annual or perennial erect herbs branching at or above the base, sparsely to densely glandular
stipitate on branches and leaves, frequently woolly. Leaves alternate, simple, entire, slender terete
to oblong or obovate. Capitula solitary on slender leafless peduncles arising from apex of branches.
Involucre hemispherical; intermediate bracts with terete glandular stipe-like claws and petaloid
laminae; innermost two series with lineart flat claws and very small laminae; the outer bracts in some
species sessile. Receptacle flat, epaleatc. Florets bisexual, actinomorphic. Corolla tubular below,
narrowly turbinate or narrowly cup-shaped above, shortly 5-lobed, very sparsely puberulous; vascular
strand terminating below lobes. Anther appendage ovate, cells narrow-oblong, walls somewhat
thickened; marginal cellsdifferentiated;anthertails firm, slightly branchedat ends. Styleapex deltoid
or broad-ovate, the vascular strand thick and clavate in the apex. Achene ellip.soidal; surface
prominently undulate or plane; carpopodium very short, 1 cell high; pericarp thick, 2-layered,
glabrous, translucent, cells linear, vascular strands laterally placed with respect to cotyledons; testa
free from pericarp, somewhat thick, epidermal cells + equilateral or oblong, with straight walls;
vascular strand laterally placed terminating in tip of seed. Pappus bristles equal to corolla, broad and
weakly connate at base, shortly plumose with acute cilia, eventually breaking shortly above base.

Five species endemic to temperate Australia. For distribution notes on taxa other than
L. stipitatum see Wilson (1960).

Hotel. De Candolle included the following species in his section Leucochrysum: Helipterum incanum
(Hook.) DC., //. bicolorum DC., //. albicans (Cunn.) DC., and //. cotula (Benth.) DC. The first three
taxa are now considered to be conspecific and are referred here \o Leucochrysum albicans while
H. cotula is now (Wilson 1989) treated as a member of the genus Hyalosperma. In the sectional
description de Candolle stated that the involucral bracts were stipitate: this is correct for //. albicans
but not for H. cotula and for this reason it would be inappropriate to de.signate the latter as lectotype.

Note 2. The concept that Bentham (1837) validly published the name Helichrysum. sect. Helipterum,
basing it on Helichrysum cotula Benth., //. anthemoides Spreng., and (with doubt) H. incanum Hook,
is discussed by Wilson (1989) but not accepted.

Key to species

1. Stem much branched, the lateral branches terminating in capitula; outer

bracts dark reddish brown, not needle-like L.fitzgibbonii

1. Stem branched only near base; peduncles long and arising from near base
of plant; outer bracts pale or needle-like

2. Outer involucral bracts needle-like L. stipitatum

2. Outer involucral bracts with an obvious flattened lamina

442 Nuytsia Vol. 8, No. 3 (1992)

3. Leaves filiform, tightly revolute, congested, glabrescent L. graminifolium

3. Leaves oblong or obovate or, if filiform, not tightly revolute,
congested or loosely arranged along stem, woolly
4. Plant annual; lamina of intermediate involucral bracts broad-ovate to
suborbicular, pale yellow; leaves linear to narrow-oblong or narrow-

obovate L. molle

4. Plant perennial; lamina of intermediate involucral bracts very narrow-

obovate to obovate, if debate then with red streaks L. albicans

Leucochrysum albicans (Cunn.) Paul G. Wilson, comb. nov.

Helichrysum albicans Cunn. in Field, Geog. Mem. New South Wales 359(1825) 'Elichrysum' . -
Heliptermi albicans (Cunn.) DC., Prod. 6:215(1838).- Argyrocome albicans (Cunn.) Kuntze, Revis.
Gen. PI. 1:308(1891). - Roccardia albicans (Cunn.) Voss, Vilm. Blumeng. ed. 3, 1:532(1895).
Lectotype: Forest Land, Cox’s River, 9 Oct. 1822, A. Cunningham 71 (K, ?iso: ME\J)fide Wilson
(1960).

Helichrysum incanum Hook., Bot. Mag, t.288 1(1829). -Aphelexis incana (Hook.) G. Don in Sweet,
Hort. Brit, ed.3, 379(1839). Lectotype: op. cit. t. 2881, /b/c Wilson (1960).

Helipterum incanum DC., Prod. 6:215(1838). Type: Van diemen,7?.C. Gunn 108, comm. Lindley in
1834 (holo: G-DC, photo seen), nom. superfl.

Helipterum bicolorum Cunn. ex DC., Prod. 6:215(1838). Lectotype: Hills of the Lachlan River,
31 July 1817 (holo: G-DC, photo seen) fide Wilson (1960).

Key to infraspecific taxa

1. Leaves obovate, densely woolly subsp. alpinum

1. Leaves filiform to narrow-oblong or narrow-obovate, thin, moderately

woolly subsp. albicans

2. Inner involucral bracts white var. tricolor

2. Inner involucral bracts yellow

3. Intermediate involucral bracts ovate to oblong, obtuse to acute var. albicans

3. Intermediate involucral bracts broadly ovate to debate or orbicular,

acute var. buffaloensis

subsp. albicans
var. albicans

Helipterum incanum vm. flavidiceps F. Muell., Rep. PI. Babbage’s Exped. 14(1859). Lectotype:
Subalpine plains on the Snowy River, F. Mueller (K, iso: MEG) fide Wilson (1960).

Paul G. Wilson, Helipterum: Part 2 Leucochrysum

443

Helipterum incanum var. auriceps F.Muell., Rep. PI. Babbage’s Exped.l4(1859). Type citation: 'In
many of the arid plains and mountains of Australia Felix and in the northern parts of South Australia'.
Lectotype not designated.

//e///;tcrM/ninc(7n«wvar.//7i/o//«mF.Muell.,Rep.Pl.Babbage’sExped.l4(1859). Lec/otype.- Station
Peak, You Yangs Mts, Victoria, F. Mueller (MEL, iso: K)fide Wilson (1960).

var. buffaloensis (Paul G, Wilson) Paul G. Wilson, comb. nov.

Helipterum albicans var. hiijfaloensis Paul G. Wilson, Trans. Roy Soc. South Austral. 83: 170(1 960).
Type: Mt Buffalo, Victoria, 10 Nov. 1955, E. Gauba (holo: GAUBA).

var. tricolor (DC.) Paul G. Wilson, comb. nov.

Helipterum incanum var. tricolor DC., Prod. 6:215(1838), Type: based on Helichrysum incanum
Hook.

Helichrysum incanum Hook., Bol. Mag. t. 2881(1829). - Helipterum albicans var. incanum (Hook.)
Paul G. Wilson, Trans. Roy. Soc. South Austral. 83:170(1960).

Helipterum incanum DC., Prod. 6:215(1838), nom. superfl.

Helipterum bicolorum DC., Prod. 6:215(1838). - Roccardia albicans f. bicolorum (DC.) Voss, Vilm.
Blumeng. ed. 3, 1:532(1895). Type: see above.

Helipterum incanum var. purpureo-album F. Muell., Rep. PI. Babbage's Exped. 14(1859). -
H. albicans f. purpureo-album (F. Muell.) Paul G. Wilson, Trans. Roy. Soc. South Austral.
83:172(1960). Lectotype: Maneroo Plains. 1855, F. Mueller (MEL), fide Wilson l.c.

Helipterum albicans f. grampianum Paul G. Wilson, op. cit. 172. Type: Skipton Plains, 1860,
W.J. Whan 39 (holo: MEL; iso: NSW).

subsp. alpinum (F.Muell.) Paul G. Wilson, comb. nov.

var. F.Muell. , Rep. PI. Babbage’s Exped. 14( 1859).- /7.fl/6ica«5subsp.

alpinum (F. Muell.) Paul G. Wilson, op.cit. 174. Type: Summit of the Australian Alps, F. Mueller
(holo: MEL; iso: K, NSW).

Note: When revising the Helipterum albicans complex (Wilson 1960), I assumed that Helipterum
incanum DC.(1838) represented a new combination ba.sed on Helichrysum incanum Hook. (1829),
a name that de Candolle had placed in synonymy under Helipterum incanum var. tricolor DC. After
discussion with my colleague Gillian Perry I have decided that this assumption was incorrect. I now
consider that Helipterum incanum DC. was published as a new species; the name was. however,
superfluous and illegitimate since a previously published species name was cited under one of its
varieties (i.e. under var. tricolor). The epithet var. tricolor is therefore legitimate but it does not create
the autonym 'var. incanum' since the species name to which it was applied was not legitimate. The
epithet var. tricolor therefore has priority over its synonym var. incanum (Hook.) Paul G. Wilson
(1960).

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Nuytsia Vol. 8, No. 3 (1992)

Figure 1. Leucochrysum stipitaium. A - Intermediate involucral bract. B - Achene with pappus. C - Apex of pappus bristle.
D - Achene after pappus bristles have broken near base. E - Anther. F - Terminal portion of anther appendage. G- Style branch
and apex showing thick vascular strand. From //. Demarz 2786.

Leucochrysum fitzgibbonii (F. Muell.) Paul G. Wilson, comb. nov.

Helip.enim fitzgibbonii F. Muell. Victorian Nat. 7:38 (July 1890); Trans. & Proc. Roy. Soc. South
Australia 13:170 (Dec. 1890); Bot. Centralbl. 43:276(1890), op.cil. 45:122(1891). Type citation:
'Tempe Downs, R.Thornton; near Georgina-River, All'r. Henry: Finke-river, Rev. H. Kempe; Lady
Charlotte’s Waters and west of Eringa, W. Tietkens; Nullarbor-Plains, J. Batt; Mount Moore, Edwin
Merrall; remotest eastern sources of Swan-River, Miss Alice Eaton.' Lectotype (here chosen): Einke
River, Kempe 408 (MEL 1 10456).

Distribution. Central Western Australia, far north South Australia, and southern Northern Territory.

Leucochrysum graminifolium (Paul G. Wilson) Paul G. Wilson, comb. nov.

Helipterumalbicansvar. graminifoliim'Piml G. Wilson, Trans. Roy. Soc. South Austral. 83: 17 1( 1960).
Type: Clarence-Wolgan Road, New South Wales, 26 Feb. 1939, W.F. Blakely, J. and
W.J. Buckingham 3306 (holo: NSW; iso: AD).

This species is evidently found on sandstone cliffs in a small area c. 40 km north of Lithgow in
New South Wales.

Paul G. Wilson, Helipteriim: Part 2 Leucochrysum

445

Although very similar to narrow-leaved variants of L. albicans it differs significantly in its almost
silky indumentum, in its tightly revolute leaves that (unlike those of L. albicans) lackchlorenchymatous
tissue beneath the thickened midrib, and in having 2-celled myxogenic papillae on the achene (these
absent in L. albicans).

Leucochrysum molle (Cunn. ex DC.) Paul G. Wilson, comb. nov.

Helichrysum molle A. Cunn. ex DC., Prod. 6:194(1838). - Helipterum molle (Cunn. ex DC.)
Paul G. Wilson, Trans. Roy. Soc. South Austral. 83:175(1960). - Gnaphalium molle (Cunn. ex DC.)
Schultz-Bip., Bot. Zeitung (Berlin) 3:171(1845) nom. illeg. Lectotype: Mode’s Plains, Lachlan
River, New South Wales, July 1817, A. Cunningham (G-DC photo seen) fide Wilson (1960).

Wailzia brachyrrhyncha F. Muell., Linnaea 25:407 (April 1853). - Helipterum brachyrrhynchum
(F. Muell.) Sonder, Linnaea 25:517 (June 1853). - Helichrysum brachyrrhynchum (F. Muell.)
Baillon, Diet. Bot.3:27(1891), - Roccardia albicans f. brachyrrhyncha (F. Muell.) Voss, Vilm.
Blumeng.3rdedn, 1:532(1895). Lec/oA’/^c.’Cudnaka.South Australia, F.yV/M<?//cr(MEL)//f/e Wilson
(1960).

Helipterum incanum var. brachylepis F. Muell., Rep. PI. Babbage’s Exped. Type citation: 'Flinders
Ranges,&c.'Lectotype{hexechosen):Cudnaka.F.Mueller(MEL,ho\otypeo{Wailziabrachyrrhyncha).

Leucochrysum stipitatum (F. Muell.) Paul G. Wilson, comb. nov. (Figure 1)

Helipterum stipitatum (F. Muell.) Bcnth., FI. Austral. 3:643(1867). - Helichrysum stipitatum
F. Muell., Fragm. 3:133(1863). - Argyrocome stipilata (F. Muell.) Kuntze, Rcvis. Gen. PI. 1:309
(1891). Type: Finke River, Central Australia,/. Macd. Stuart (holo: MEL 110722).

Distribution. Central Western Australia east to central and northern South Australia, southern
Northern Territory, and south west Queensland.

Acknowledgements

1 thank my colleague Gill Perry for helpful di.scussions on some questions of nomenclature, Phil
Short for his useful comments on the manuscript, Margaret Menadue for providing the illustration,
and the curators of the herbaria of Adelaide (AD), Melbourne (MEL), and Sydney (NSW) for the loan
of type material.

References

Anderberg, Ame A. (1989). Phylogeny and reclassification of the tribe Inuleae (Asteraceae). Canadian J. Bot. 67:2277-2296.
Anderberg, Ame A. (1991). Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Botanica 104:1-195.
Benlhani, G. (1837). Compositae. In S.L. Endlicher el al., "Enumeratio Planlanuii." (Beck University Biblio., Vienna.)
Bentham, G. (1867). "Flora Ausiraliensis," vol. 3. (I.ovell Reeve & Co., London.)

446

Nuytsia Vol. 8, No. 3 (1992)

Candolle, A.P. de (Jan. 1838). "Prodromus systemalis naturalis regni vegetabilis." Vol. 6. (Treuttel & Wurtz, Paris.)
Hilliard, O.M. & Burtl. B.L. (1981). Some generic concepts in Compositae - Gnaphaliinae. J. Linn. Soc. Bot. 82:181-232.

Short, P.S., Wilson, K.E. and Nailon J. (1989). Notes on the fmit anatomy of Australian members of the Inuleae (Compositae).
Muelleria 7:57-79.

Turner, B.L. (1970). Chromosome numbers in the Compositae. XII Australian species. Amer. J. Bot. 57:382-389.

Wilson, Paul G. (1960). A consideration of the species previously included wilh'm Heliplerum albicans (A.Cunn.)DC. Trans.
Roy. Soc. South Australia 83:163-177.

Wilson, Paul G. (1989). A revision of the genus Hyalosperma (Asteraceae: Inuleae: Gnaphaliinae). Nuytsia 7:75-101.

Wilson, Paul G. (1991). The classification of the Australian species currently included in WW(ptcr«m(Asteraceae:Gnaphalieae):
Parti. Nuytsia 8:379-438.

Nuytsia 8(3): 447-460 (1992)

447

The classification of some Australian species currently included in
Helipterum and Helichrysum (Asteraceae: Gnaphalieae): Part 3
Anemocarpa and Argentipallium, two new genera from Australia

Paul G. Wilson

Western Australian Herbarium, Department of Conservation and Land Management,
PO Box 104, Como, Western Australia 6152

Abstract

Wilson, Paul G. The classification of some Australian ,species currently included in Helipterum
andHelichrysum (Asteraceae; Gnaphalieae): Part 3 Anemocarpa and Argentipallium, two new genera
from Australia. Nuytsia 8(3): 447-460 (1992). Two new genera Anemocarpa ax\d Argentipallium txre
described in the Gnaphalieae: Angianthinae. The first genus is based on a new species
Anemocarpa calcicola from Western Australia and includes two other species: A. podolepidium
[Helichrysum podolepidium F. Muell.], and A. saxalilis [Helipterum saxatile Paul G. Wilson]; it is
apparently most closely related to the generaLeucochrysum (DC.) Paul G. Wilson and Chrysocephalum
Walp. The second genus is based on Argentipallium obtusifolium (Sonder) Paul G. Wilson
[Helichrysum obtusifoliumSonder] and includes five other species: A. blandowskianum [Helichrysum
blandowskianum Sonder], A. dealbatum [Helichrysum dealhatum Labill.], A. niveum [Helipterum
niveum Steetz], A. spiceri [Helichrysum spiceri F. Muell.], and A. tephrodes [Ozothamnus tephrodes
Turez.]; it is evidently most closely related to Ozothamnus R.Br.

Introduction

The circumscription of the genera Helipterum DC. and Helichrysum Miller in Australia has been
discussed in several recent papers (Hilliard & Burtt 1981, Wilson 1989, Anderberg 1991), where it
has been demonstrated that these genera, in the strict sense, do not occur in Australia and that the
Australian species previously included in them should be placed in a number of distinct genera.
Anderberg, op. cit., further indicated that the genera Lawrencetla Lindlcy and Rhodanthe Bindley
were polyphylelic as circumscribed in his treatment and needed further study.

In this paper 1 am describing two new genera. The ’Cirsi, Anemocarpa, includes one species placed
by Anderberg in the 'Rhodanthe' assemblage and one placed by him in Chrysocephalum Walp. The
second, Argentipallium, contains six species, four of which were placed by Anderberg in the
Lawrencella 'Scorpioide s’ eomplex. Prior to Anderberg's assessment the species involved in the two
new genera had been treated as belonging to either Helichrysum or Helipterum. Eventually it is
probable that further species currently placed in Helichrysum will be trtinsfcrred to one or other of
the new genera. The correct circumscription and nomenclature of these additional species has still

448

Nuytsia Vol. 8, No. 3 (1992)

to be elucidated, a task that will entail further research and will involve the study of material in
Australian and overseas herbaria; for these reasons only those species are treated whose names are
required in regional floras and whose taxonomy and nomenclature is clear.

Methods

Dissections of achenes, florets, and involucral bracts of representative Australian species of
'Helipterum', 'Helichrysum', and of related genera have been mounted in Hoyer’s solution (King &
Robinson 1970) and examined under the stereo and high-powered microscope. Achenes have also
been embedded in Spurr’s low-viscosity embedding media and sections stained in Toluidine Blue.
In addition microscope slides were studied of transverse sections of achenes, some of which were
prepared by P.S. Short, K.E. Wilson and J. Nailon for their paper (1989) on achene anatomy while
others were prepared for me by my colleagues R. Cranfield and C. Parker.

Morphological studies

The morphological characters used in discriminating the genera and species have been described
(or references given to their description) in a number of earlier papers (e.g. Wilson 1989, 1992b);
supplementary notes are here provided on some of these characters.

Achenial hairs. The nature of the hairs or papillae on the achenes was studied. Various types of hairs
and papillae found in the Gnaphalieae have been described by Anderberg (1991) and their taxonomic
significance assessed. In the 'Helichrysum elatum' group an additional 'papilla' type has been
observed, in this two cells of the pericarp epidermis are laterally in.stead of vertically placed; they are
transparent and often myxogenic. Papillae of this type are referred to as 2-celled fenestrae* . In some
species of this group the fenestrae apparently consist of only one longitudinally elongated cell, this
cell is usually not myxogenic and is not raised above the surface of the achene.

In Anemocarpa low, rounded, 2-celled papillae are present with the lower cell slightly exceeding
the upper. This papilla type, which was described by Anderberg (1991), is comparable to that found
in the Waitzia group of genera.

In Argentipalliuni short narrow-obovoid duplex hairs are present; these are similar to the more
elongated duplex hair type that is found in Ozothamnus.

Achenial nervation. The vasculature of the achene can be determined by mounting it in Hoyer’s
solution. Where the testa is free from the pericarp, or where it can be readily separated from it, the
vasculature is preferably examined in an achene in which the seed (i.e. testa, endosperm, and embryo)
has been removed. In all Australian members of the Gnaphalieae where this organ has been studied,
with the exception of Myriocephalus guerinae F. Muell. (see Short et al. 1989), there are two vascular
strands in the pericarp. In the testa there is always one strand and this passes for a varying degree
longitudinally around the seed; it is best observed by studying a whole mount of the testa from which
the embryo and endosperm (which is usually ± free from the testa) have been removed.

* This 'papilla' type is also found in Bracieaniha as is noted by E.E. Georgeadou Tvrlkovic-Saltin (1982) who refers to them as
'2-celled idioblasts'. It is anticipated that these will be more fully described, and their taxonomic significance assessed by her
in a forthcoming paper.

Paul G. V*/\]son, Anemocarpa and Argentipallium

449

The position of the vascular strands of the testa and pericarp relative to the cotyledons of the
embryo is of generic significance (Short etal. op. cit.) and may be determined by examining transverse
sections of the achene, by examining cleared whole mounts (if these are sufficiently transparent), or
by dissecting the achene. Their position has been observed in representative species of all of the genera
or species-groups discussed in this paper.

In Anemocarpa the achenial vascular strands arc laterally placed in relation to the cotyledons as
they are in its putative closest relatives Waitzia Wendl. and Leucochrysum (DC.) Paul G. Wilson.
They are also laterally placed in species of the Helichrysum elatum group.

In ArgentipaUium the vascular strands arc medially placed, which is also the case in Ozothamnus

R.Br.

Involucral bracts. The cartilaginous basal portion of the involucral bracts, the stcreome, was first
critically described by Drury (1970). He pointed out that in some genera the stereome is not entire
and that unthickened translucent areas, here referred to as 'fenestrae', are present between the
thickened opaque regions. This character has been used by Hilliard and B urtt ( 1 98 1 ) and by Anderbcrg
(1991) to assist in the segregation of species into discrete genera in the Gnaphalium - Helichrysum
complex. The Au.stralian .species of this complex are also amenable to this procedure. The species
of the genus ArgentipaUium have fenestrate stercomes whereas species of Anemocarpa and of the
Helichrysum elatum group do not. In the genus Ozothamnus, as currently circumscribed (Anderberg
1991), some species appear to have fenestrate stereomes while others do not.

Pappus. In Anemocarpa the pappus bristles are free and persistent but they eventually break c. 1 mm
above the base in the same manner as do the bristles in species of the Helichrysum elatum group and
of Leucochrysum. The bristles are barbellate upwards with the terminal barbellae congested and
prominent but not thickened.

In ArgentipaUium the bristles are very shortly united, are persistent, and do not break above base.
They are barbellate upwards with the terminal barbellae clavate and thickened. This situation is
similar to that found in Ozothamnus.

Mycorrhizal associations. Ithas been shown by Warcup (1990) that only about half of the Australian
species that had been previously included in either H elipterum or Helichrysum form ectomycorrhiza.
Warcup pointed out that this was unusual since, in general, species of a composite genus are usually
either all ectomycorrhizal or all non-ectomycorrhizal. However, when the species are segregated into
the genera recognised by Anderberg (1991) and by myself these segregate genera have so far been
found to contain only ectomycon hizal or only non-ectomycorrhizal species. Unfortunately only a
few species involved in the present study have been tested.

Helichrysum podolepidium F. Muell. (Anemocarpa) was found not to form ectomycorrhiza. This
contrasts with species in the putatively closely related genera Leucochrysum and Waitzia that form
ectomycorrhiza. It also contrasts with species of Chrysocephalum in which genus H. podolepidium
was placed by Anderberg (1991).

Helichrysum dealbatum Labill. and H. blandowskianum Sonder (both ArgentipaUium) form
ectomycorrhiza, while species in the morphologically similar genus Ozothamnus do not.

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Nuytsia Vol. 8, No. 3 (1992)

Generic studies

In a recent paper Anderberg (1991) recognised the genera Chrysocephalum Walp., Bracteantha
Anderb., Ozothamnus R.Br., and iMwrenceUa Lindley sensu lato as segregates of (Australian)
Helichrysum. I have reclassified the taxa in the Lawrencella complex (Wilson 1992a) and most of
the Australian representatives of Heliplerum (Wilson 1992a,b). This leaves a number of Australian
species of // elichrysim and of Helipterum that still require generic classification; they are as folio ws:-

1) The 'Helichrysum datum' group. Species in which the involucral bracts have terete or semi-terete
stipes and are not fenestrate, the anther apicula have thick cell walls, the style appendages are ovate
to acuminate, and the achencs are glabrous with a smooth crustaceous pericarp in which the vascular
strands are laterally positioned. The epicarp has longitudinal 2-celled fenestrae (idioblasts) of the
Bracteantha type. The group is represented by such species as //. elatum DC., //. boormanii Maiden
& Betche, and H. adenophoriim F. Muell. It corresponds to Helichrysum sect. Blepharolepis DC. pro
parte and, from the few species investigated (Warcup 1990), is evidently not ectomycorrhizal.

2) The 'Helipterum saxatile' group. Species similar in mo.st characters to those in group 1 but with
thin anther apicula, rounded style apex, and papillose achenes (the two cells of the papillae vertically
positioned) with a thin pericarp. This group is represented by Helipterum saxatile Paul G. Wilson,
Helichrysum podolepidium F. Muell., and an undescribed species. The species are here placed in the
new genus Anemocarpa. It is possible that it will have to be enlarged to include species at present
in group 1.

3) The 'Helichrysum dealbatum' group. Species with a flattened claw to the involucral bracts,
fenestrate stereome, thin anther apicula, truncate to rounded style apices, short clavate to obovoid
duplex hairs on the achenes, a diaphanous pericarp, and a leathery testa, with the achenial vascular
strands medially positioned. These species form ectomycorrhiza. To this group belong Helichrysum
dealbatum Labill., H. spiceri F. Muell., Ozothamnus tephrodes Turez., Helichrysum obtusifolium
Sonder, and Helipterum niveum Steetz; they are here placed in the new gems Argentipallium.

4) H. oligochaetumF. Muell. This species appears to have no near relatives in the 'Helich?jsum' group
and is possibly best placed in close association with Rhodanthe.

5) Helichrysum pumilum J.D. Hook. A species with the habit of Leucochrysum albicans (A.Cunn.)
Paul G. Wilson but with florets similar to some species of Rhodanthe particularly in having villous
achenes; it differs in having heterogamous capitula, thick branched anther tails, deeply stained
corollas, and barbellate pappus bristles.

6) Helichrysum leucopsideum DC. A rhizomatous species with the appearance of certain species in
the 'Helichrysum elatum 'group. Its florets are similar to those in Bracteantha but have a rounded style
apex, a smooth glabrous cylindrical achenc without fenestrae or myxogenic cells, and adoublc pappus
that is persistent. The vascular strands of the pericarp and testa are medially orientated relative to the
cotyledons whereas in Bracteantha they are laterally orientated (Short 1989). This is the only
Australian species in the 'Helipterum - Helichrysum' complex in which the pappus is double and it
is the only species that is recorded as being rhizomatous.

Paul G. Wilson, Anemocarpa and ArgentipalUum

451

Key to genera

The genera discussed in this paper may be distinguished by the following synoptic key:-

1. Annual herbs; involucral bracts with flattened proximal portion,
stereome not fenestrate: anther appendage with proximal cells
equilateral and thick-walled, distal cells narrow-oblong and
thin-walled; style apex deltoid; pappus bristles evenly barbellate
or cilia longer near base; achene villous or hispid with large

thick duplex hairs; ectomycorrhiza absent Lawrencella group

2. Base of achene prominently excavated; vascular strands (v.s.) of
pericarp and testa lateral

3. Radiant petaloid involucral bracts absent; leaves terete, strictly basal Bellida

3. Radiant petaloid involucral bracts present; leaves various, sub-basal

or caulinc Lawrencella

2. Base of achene minute, scarcely excavated; v.s. of pericarp and testa

medial Schoenia

1. Perennial herbs or shrubs; involucral bracts with flattened or stipe-like
proximal portion, fenestrate or not; anther appendage with proximal
cells narrow-oblong and similar to distal cells; style apex truncate or
rounded to acuminate; pappus bristles barbellate or plumose, smooth
near base; achene glabrous, papillo.se, or with very small or weak
duplex hairs; ectomycorrhiza present or absent

4. Achenes with short clavate duplex hairs; pericarp thin, transparent,
v.s. medial; involucral bracts papery, fenestrate; anther tails long
and slender exceeding the collar; pappus persistent; ectomycorrhiza

present; perennial herbs ArgentipalUum

4. Achenes glabrous or minutely papillose, or with .short slender duplex
hairs; v.s. of pericarp medial or lateral; involucral bracts glumaceous
or papery, not fenestrate; pappus persistent or caducous;
ectomycorrhiza present or ab.scnt: shrubs or perennial herbs

5. Shrubs; capitula small and clustered; involucral bracts papery; style
apex truncate; achene with short narrow duplex hairs or glabrous,

v.s. of pericarp medial; pappus persistent; ectomycorrhiza absent Ozothamnus

5. Perennial herbs; capitula solitary or clustered; style various; achenes
smooth or minutely papillose, v.s. of pericarp lateral; pappus
persistent or caducous; ectomycorrhiza present or absent

6. Intermediate involucral bracts with a flattened proximal portion;
ectomycorrhiza present

7. Achenes with papillae consisting of two cells of which the lower
overtops the upper; involucral bracts papery, strongly ciliate;

style apex truncate; pericarp ciu-tilaginous; pappus persistent Chrysocephalum

7. Achenes smooth and glabrous; involucral bracts glumaceous,

entire; style apex acute; pericarp thick and hard; pappus caducous Bracteantha

6. Involucral bracts with a stipc-like proximal portion; pappus
persistent; ectomycorrhiza pre.sent or absent

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Nuytsia Vol. 8, No. 3 (1992)

8. Pappus bristles barbellate; style apex rounded to acuminate,
not deltoid: pericarp papery or brittle; ectomycorrhiza absent
9. Pericarp papery; style apex rounded; carpopodium

prominent, several cells high Anemocarpa

9. Pericarp brittle; style apex ovate to acuminate;

carpopodium minute, 1 cell high 'Helichrysum elatum' group

8. Pappus bristles plumose; style apex broad-deltoid;
pericarp thick, cartilaginous; carpopodium minute,

1 cell high; ectomycorrhiza present Leucochrysum

Anemocarpa Paul G. Wilson, gen. nov.

Herbae perennes, non-ectomycorrhizales, lanatae. Folia alterna, sessilia. Capitula solitaria.
Involucrumhemisphericum. Bracteaeinvolucraleschartaceae,angu.steellipticae;bracteaeinterioris
parte inferiores sem iteretibus, stereomate efenestrato, superiores anguste ellipticae albae. Receptaculum
convexum, epaleaceum, glabrum. Flosculi homogami, disciformes. Corolla anguste cylindrica sed
versus apicem anguste turbinata, breviler 5-lobata, cellulis faucis undulatis. Caudae antherarum
graciles sed firmac. Styli apex rotundatus, nervo prominenti fere ad apicem extenso. Achenium
ellipsoideum, glabrum; pericarpum translucens, papillis 2-cellulis ornatum; testa membranacea,
crystallina, nervo lateraliter posito; carpopodium multicelluloso altum. Pappi setae persistentes
barbellatae, cellulis apicalibus acutis.

Typus: Anemocarpa calcicola Paul G. Wilson

Perennial herbs. Ectomycorrhiza not formed. Plant covered with a woolly indumentum. Leaves
alternate, simple, entire, sessile. Capitula solitary and terminal to stems or branches, homogamous,
disciform. Involucre hemispherical. Involucral bracts papery, narrow-elliptic; outer bracts pale
brown without stipe; inner bracts with proximal portion semi-terete and stipe-like with thickened
stereome, not fenestrate, distal portion a white papery lamina, the central vascular strand passing into
the lamina; innermost bracts with a linear flattened proximal portion grading into a very short truncate
lamina. Receptacle convex, epaleate, glabrous. Corolla narrow-cylindrical, narrow-turbinate
towards the apex, regularly shortly 5-lobed, glabrous and smooth within, sparsely glandular pilose
outside; cells of inner epidermis of throat undulate. Anther apiculum narrow-triangular; constituent
cells narrow-oblong with thickened walls; tmls slender, firm, ± equal to narrow-oblong collar. Style
apex rounded, shortly papillate; vascular strand prominent and extending to near tip. Achene
ellipsoid, glabrous; pericarp diaphanous to papery and translucent, epidermis with 2-celled myxogenic
papillae, the cells separated by a transverse wall, the tower cell slightly exceeding the upper; testa
membranous, pale brown, adnate to pericarp, epidermal cells broad-oblong with straight walls,
crystals present, vascuhu" strand in lateral position (in relation tocotyledons) and not extending to apex
of seed; carpopodium prominent, several cells high. Pappus bristles, free, persistent but eventually
breaking c. 1 mm above base, equal to corolla, slender, smooth towards base, barbellate upwards, the
terminal barbellae congested, prominent, and acute.

Etymology. The generic name is derived from the Greek words anema, without a thread, and carpos,
fruit, with reference to the absence of hairs on the achenes.

Paul G. Anemocarpa and Argendpallium

453

This genus appears to be most closely related to Leucochiysum. It shares with that genus lateral
vascular strands in the achene, a similar type of pappus, and similar involucral bracts. The genera
differ from each other in the shape of the style apex, in the achene morphology (particularly the
presence of a prominent carpopodium in Anemocarpa), in the presence of 2-celled papillae in
Anemocarpa and their absence in Leucochrysum, and evidently also in their mycorrhizal status
(Warcup 1990).

As is mentioned earlier, a number of species additional to those listed here will probably have to
be transferred to this genus from Helichrysum. However, taxonomic studies on these species is
continuing and will be the subject of a later paper. Only those species whose nomenclature and
circumscription is clear are presently considered.

Key to species

1. Involucre lacking ray bracts A. podolepidium

1. Involucre with prominent white ray bracts

2. Leaves mostly 5-7 mm long (south coast W.A.) A. calcicola

2. Leaves mostly 20-40 mm long (central Australia) A. saxatilis

Anemocarpa calcicola Paul G. Wilson, sp. nov. (Figures 1, 2a)

Rami graciles. Folia anguste elliptica, 5-7(12) mm longa, lanata, parum crenulata, marginibus
revoluta. Capitula hemispherica, ad 30 mm diam.; bracteae exteriores ovatae, malvinae vel pallide
bruneae; interiores lamina anguste elliptica, alba ad 12 mm longa.

Typus: 10 km north of Eyre, 32° 09’S, 126° 17’E, Western Australia, 1 Oct. 1984, G.y.A'c/g/jcry 7533
(holo: PERTH; iso: CANB, K).

Erect perennial to 30 cm high and wide, with a white woolly indumentum on leaves and branches.
Branches slender. Leaves cauline, alternate, sessile, narrowly elliptic, 5-7(12) mm long, sparse and
small towards the capitula; margins revolute, slightly crenulate; upper surface sparsely woolly; lower
surface densely woolly. Capitula hemispherical, to 30 mm diameter; outer bracts sessile, ovate, pale
brown or tinged with mauve, inner with ikutow elliptic white lamina to 12 mm long, spreading but
not reflexed on semiterete claw. Florets numerous. Corolla c. 3 mm long; lobes deltoid c. 0.4 mm
long. Anther loculi c. 1 mm long; apiculum c. 0.3 mm long. Achene ellipsoid, 1.5 mm long.

Specimens examined (selection only). WESTERN AUSTRALIA; 6km north ofEyre,Demar 2 9760;
Baxters Cliffs south of Caiguna, 24 August 1983, Fitzgerald', Twilight Cove, George 8558; 2 km
north-west of Toolina Rockholc, Keighery & Alford 635; Telegraph Pass, Hampton Scarp, Newbey
11762; Pt Dover, Wilson 5917. (All PERTH).

Distribution. Near the south east coast of Western Australia from Toolina Cove (125° E) to Eyre
(126° 18’E).

Habitat. Sandy loam over limestone in mallee shrubland or coastal heath.

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Nuytsia Vol. 8, No. 3 (1992)

Figure 1. Anemocarpa calcicola. A - habit; B - leaf; C - outer, inteniiediate and imier involucral bracts; D - floret; E - apex of
pappus bristle; F- style apex; G- anther; H - achene; I - surface and side views of pericarp showing myxogenic2-celled papillae;
J - seed showing vascular strand. From P.G. Wilson 7693.

Paul G. Wilson, Anemocarpa and Argentipallium

455

Etymology. The specific epithet is derived from the Latin words calx, chalk, and cola, an inhabitant,
with reference to the substrate over which the plant is found.

This species is superficially similar to and has been confused with Argentipallium obiusifolium
(Sonder) Paul G. Wilson (as Helichrysum obtusifolium Sonder), from the latter it differs in the
indumentum being woolly (not thick silky), in the involucral bracts having a semi-terete stipe (not
a flattened basal portion), in the lamina of the innermost bracts being very short and truncate (not large
and similar to the inner bracts), in the achenes having low rounded 2-celled papillae (not short duplex
hairs), and in the anther apiculum being narrow-triangular with thickened cells (not ovate with thin-
walled cells). Whereas A/I cOTOcarpo cakicola is found in south-east Western Australia Argc/m>a///M/ 7 i
obtusifolium is found in southern eastern Australia to as far west as Eyre Peninsula.

Another superficially similar taxon that has been confused with 11. obtusifolium is Argentipallium
niveum (Steetz)Paul G. Wilson [Helipterum niveum Steetz], It is found in the south-west of Western
Australia and is readily distinguished from otherwise similar species by its axillary clusters of linear
leaves.

Anemocarpa podolepidium (F. Muell.) Paul G. Wilson, comb. nov.

Helichrysum podolepidium F. Muell., Rep. PI. Babbage Exped. 13(1859). - Chrysocephalum
podolepidium (F. Muell.) Andcrb., Op. Bot. 104:119(1991). Type: Wirrawirraloo, 31° 11' S,
136° 41' E, South Australia, 1858, B.H. Babbage.

Distribution. Found in southern Northern Territory, north-eastern South Australia, south-west
Queensland, and north-west New South Wales.

Anemocarpa saxatilis (Paul G. Wilson) Paul G. Wilson, comb. nov.

Helipterum saxatile Paul G. Wilson, Trans. Roy. Soc. South Australia 83:166(1960). Type: Near
Hermannsburg, 24 Aug. 1956, G. Chippendale 2632 (holo: NT; iso: CANB, NT).

Distribution. Found in southern Northern Territory, northern South Australia, south-westQueensland,
and north-west New South Wales.

Argentipallium Paul G. Wilson, gen. nov.

Herbae perennes cum fungis in ectomycorrhiza consociatae, dense sericeae. Folia alterna integra
sessilia, linearia vel elliptica. Capitula solitaria. Involucrum hemisphericum vel late turbinatum;
bracteae exteriores, chartaceae, anguste ellipticae; bracteae interiores parte inferiores oblongo
indurato, lamina chartacea anguste elliptica alba ornatac, stereomate fcncstrato; bracteae intimis
interioribus similaribus. Receptaculum convexum epaleaccum glabrum. Flosculi homogami vel
heterogami cum pauci flosculi feminei, disciformes. Corolla anguste cylindrica vel supra anguste
turbinata, 5-lobata. Caudae anthcrarum graciles, debiles vel firmae. Styli apex truncatus vel
rotundatus. Acheniumangustecylindricum,laeve,pilisbrcvissimisobovoideisornatum;pericarpium
diaphanum; testa membnuiacea palido brunea, nervo in posiio laterali ad apicem seminis extenso.
Pappi setae ad basim concretae, ± persistentes, barbellatae, in flosculis femineispaucae vel absentes,
barbellis ad apicem plerumque clavalis.

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Nuytsia Vol. 8, No. 3 (1992)

Figure 2. Transverse sections of achenes of A - Anemocarpa calcicola and B - Argentipallium obtusifoUum, both semi-
diagrammatic. A from H. Demarz 9760 and B from D. Hunt 3041.

Typus: Argentipallium obtusifoUum (Sonder) Paul G. Wilson

Perennial herbs. Ectomycorrhiza formed. Plant covered with a dense silvery silky indumentum.
Leaves alternate, simple, entire, sessile, linear to elliptic. Capitula solitary and terminal to stems or
branches or in loose terminal panicles, homogamous or heterogamous, disciform. Involucre
hemispherical to broad-turbinate. Involucral bracts papery, narrow-elliptic; outer bracts pale brown
without lamina; inner bracts with proximal portion flat and grading into the white lamina; stereome
flat, fenestrate, grading into the surrounding bract tissue, the vascular tissue not passing into the
lamina; innermost bracts similar to the inner bracts. Receptacle convex, epaleate, glabrous. Female
florets few or absent, with filiform corollas and few or no pappus bristles; achcne as in disc llorets.
Disc florets perfect. Corolla narrow-cylindrical or upper portion turbinate, regularly 5-lobed;
vascular strands not passing into lobes. Anther apiculum oblong to elliptic, thin; constituent cells
narrow-oblong, not thickened, the marginal ones differentiated or not. Anther tails slender, shortly
branched, weak or firm, equal to or exceeding collar. Anther collar narrow-oblong. Style apex
truncate or rounded, shortly papillate; vascular strand either slender and ending well below apex or
thick and extending to near tip. Achene terete; surface smooth; indumentum sparse or dense, of .short
(0.05-0.15 mm long) obovoid duplex hairs; pericarp diaphanous; testa membranous, pale brown,
adnate to pericarp, cells broad-oblong, vascular strand in lateral position (in relation to cotyledons)
and extending to apex of seed; carpopodium several cells high. Pappus bristles free or fused towards
base in small clusters, very shortly united in a ring at base, ± persistent, slender, smooth towards base,
barbellate upwards, the terminal barbellae usually clavate.

This genus is similar in floral characters to Ozothamnus. The latter genus differs principally in
habit (its species being either shrubs or subshrubs), in the size and arrangement of the capitula, and
in not being ectomycorrhizal. The achenes of Ozothamnus have short narrow duplex hairs whereas
in Argentipallium they are very short and clavate to obovoid. In Ozothamnus there is considerable

Paul G. Wilson, Anemocarpa and Argeniipallium

457

variation in the form and length of the anther tails, whereas in Argeniipallium the anther tails are long,
stout (weak in A. niveum), and shortly branched.

Etymology. The name is derived from the Latin words argenteus, silvery and pallium, a mantle, in
allusion to the silvery indumentum that characteristically covers the leaves and branches of species
of this genus.

Key to species

1. Stem leaves small, linear, clustered in axillary fascicles A. niveum

1. Stem leaves solitary

2. Tufted herb; flower head solitary on scape-like stems A. dealbatum

2. Open perennial or small shrub; flower heads clustered or solitary
3. Flower heads to 15 mm diameter, frequently corymbose;

indumentum woolly.

4. Leaves thick-woolly all over A. blandowskianum

4. Leaves closely woolly below, glabrescent above A. spiceri

3. Flower heads 10-25 mm diameter, solitary; indumentum a
compact silky sheen

5. Leaves narrow-oblong to linear, 5-20(60) mm long; branches
never spinescent; flower heads 15-25 mm diameter, not

subtended by leaves; achenes sparsely papillose A. obtusifolium

5. Leaves linear or scale-like, to 8 mm long; branches sometimes
spinescent; flower heads to 15 mm diameter, subtended by

reduced foliage leaves; achenes densely papillose A. tephrodes

Argentipallium blandowskianum (Steetzex Sender) Paul G. Wilson, comb. nov.

Helichrysum blandowskianum Steetzex Sender, Linnaea25;5 12 (1853). Type: 'Encounter Bay' n.v.

Helichrysum blandowskianum var. dichroon Sender, Linnaea 25:512 (1852). Type: 'Nov.-Holl.
austro-occidentalis’ n.v.

Distribution. Found in south eastern South Australia and western Victoria.

I have not been able to find the types of the above two names but the original descriptions are clear
and there is little doubt about their application.

This species differs from the other species of the genus in having a semi-terete stipe to the
intermediate involucral bracts, in having a longer and more slender corolla with large ellipsoid glands
on the lobes, and in the achene having a translucent pericarp with thickened radial walls and a thin
weak testa. It has .some similarities to the Helichrysum elatum group but differs profoundly in that
the bract-stipe does not bear the glandular hairs that are found in H. elatum, and the achenial nerves
are medially (not laterally) placed. In addition it is ectomycorrhizal whereas species of the//, elatum
group are not (Warcup 1990).

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Nuytsia Vol. 8, No. 3 (1992)

Argentipallium dealbatum (Labill.) Paul G. Wilson, comb. nov.

Helichrysum dealbatum Labill., Nov. Holl. PI. Sp. 2:45 t. 190 (1806). - Gnaphalium niveum Poir.,
Encycl. Meth. Bot. Suppl. 2:808(1812) nom. illeg. non L. (1753) - G. dealbatum (Labill.) Schultz-
Bip., Bot. Zeitung 3:171(1875) nom. illeg. non Thunb. (1800). Type citation: 'Habitat in capiteVan-
Diemen'. Type n.v.

Distribution. Southern Victoria and Tasmania.

Argentipallium niveum (Steetz) Paul G. Wilson, comb. nov. (Figure 3)

Helipterum niveum Steetz in Lehm., PI. Preiss. 1:475(1845). Type: In Nova Hollandia (Swan River
Colonia) in solo sublimoso inferioris sinus regis Georgii III, Oct. 1840, L. Preiss 11 (holo: MEL).

[Helichrysum obtusifolium auct. non Sonder: Benth., FI. Austral. 3:619(1867) p.p.; Grieve &
Blackall, How to know Western Australian Wildflowers 840(1975).]

Distribution. South-west region of Western Australia.

Chromosome number: n-\2 fide Turner (1970) as Helipterum obtusifolium.

Argentipallium obtusifolium (Sonder) Paul G. Wilson, comb. nov.

Helichrysum obtusifolium Sonder, Linnaea 25:513(1853). Type: Encounter Bay, South Australia,
November, F. Mueller (holo: MEL 604819).

Distribution. Southern South Australia, western and southern Victoria, south coast of New South
Wales, and northern Tasmania.

Note. There is considerable difference in the habit and capitulum shape and size in different
geographical variants of this species. In Kangaroo Island, South Australia, the plants are small, 6-12
cm high, the involucres short (3-4 mm high) and urceolate, the laminae to the involucral bracts 3-4
mm long, and the pappus bristles white with clavate cilia at the apex. This variant resembles the
Western Australian species, A. tephrodes, which differs most obviously in having smaller leaves luid
densely papillose achenes. A rather similar plant is also found on the Fleurieu Peninsula. In south-
eastern South Australia, Victoria, and in New South Wales the plants are larger with longer leaves,
larger and hemispherical involucres, laminae to 10 mm long, and the pappus bristles with translucent
acute terminal cilia. However, since the variants appear to grade into each other I have not attempted
to provide an infra-specific classification.

Argentipallium spiceri (F. Muell.) Paul G. Wilson, comb. nov.

Helichrysumspiceri'F.y[u&\\.,Vxt\gm. 1 1:47(1878). Type.' Near Longley, Tasmania, A. S/wxon (holo:
MEL).

[Helichrysum obtusifolium auct. non F. Muell.: Rodway, Tasmanian Flora 87(1903) p.p.]

Distribution. Southern Tasmania.

Paul G. 'fJWs.on, Anemocarpa and Argentipallium

459

Figure 3. Argentipallium niveum. A - habit; B - inner, intemiediate and outer involucral bracts; C - floret; D - apex of pappus
bristle; E - style apex; F - anther; G - achene; H - duplex hair on pericarp. From P.G. Wilson 7871.

460

Nuytsia Vol. 8, No. 3 (1992)

Argentipallium tephrodes (Turcz.) Paul G. Wilson, comb. nov.

Ozothamnus tephrodes Turcz., Bull. Soc. Imp. Naturalistes Moscou 24(2):79(1851). - Helichrysum
obtusifolium var. tephrodes (Turcz.) Benth., FI. Austral. 3:619(1 867). Type: J. Drummond 5th coll,
no. 385 (holo: KW, photo seen; iso: MEL, PERTH).

Helichrysum obtusifolium var. squamiger Benth., FI. Austral. 3:619(1867). Type citation: 'Near
Oldfield river. Maxwell'. Syntypes: In tea tree salt flats near Oldfield R., G. Maxwell (MEL);
Melaleuca flats, near Oldfield R., G. Maxwell (MEL).

Distribution. Southern Western Australia from Fitzgerald River east to Israelite Bay.

Acknowledgements

I am grateful to Philip Short (MEL) for making available to me slides of transverse sections of
achenes of various species of 'Helipterum' and 'Helichrysum', to Ray Cranfield and Cheryl Parker for
preparing slides of sections of achenes, and to the herbaria AD, HO, and MEL for the loan of
specimens.

References

Anderberg, A.A. (199t). Taxonomy and phytogeny of the tribe Gnaphalieae (Asteraceae). Opera Botanica 104:t-195.
Drury, D.G. (t970). A fresh approach to the classification of the genus Giiaplialium with particular reference to the species
present in New Zealand. (Inuleae-Coniposilae). New Zealand Journal of Botany 8:222-248.

Candolle, A.P. de (1838). "Prodromus systeinalis naturalis regni qvegeuibilis." Vol. 6. (Treuttel & Wurtz, Paris.)

Hilliard, O.M. & Burtt. B.L. (1981). Some generic concepts in Compositae-Gnaphaliinae. J. Linn. Soc. Bot. 82:181-232.
King, R.M. & Robinson, H. (1970). Tlie new synantherology. Taxon 19:6-11.

Short, P.S., Wilson, K.E. & Nailon, J. (1989). Notes on the fruit anatomy of Au.stralian members of the Inuleae (Compositae).
Muelleria 7:57-79.

Turner, B.L. (1970). Chromosome numbers in the Compositae. XII Australian species. Amer. J. Bot. 57:382-389.
Tvrtkovic-Sahin, E.E. Georgiadou (1982). The systematics of the genus Helichrysum Miller in the countries bordering the
Mediterranean. Thesis, University of Reading

Warcup, J.H. (1990). The mycorrhizal associations of Australian Inuleae (Asteraceae). Muelleria 7:179-187.

Wilson, Paul G. (1989). A revision of the genus Hyatosperma (Asteraceae: Inuleae: Gnaphaliinae). Nuytsia 7:75-101.
Wilson, Paul G. (1992a). Tlie LaH'rencella complex (Asteracetie: Gnaphalieae) of Australia. Nuytsia 8:361-377.

Wilson, Paul G. (1992b). The classification of the Australian species currently included in Helipterum and related genera
(Asteraceae: Gnaphalieae): Part 1. Nuytsia 8:379-438.

Wilson, Paul G. (1992c). Leucochrysum (Asteraceae: Gnaphalieae), a new Australiiui genus in the //cZ/ptcruni complex. Nuytsia
8:439-445.

Nuytsia 8(3); 461-477 (1992)

461

The classification of the genus Waitzia Wendl. (Asteraceae:

Gnaphalieae)

Paul G. Wilson

Western Australian Herbarium, Department of Conservation and Land Management,
PO Box 104, Como. Western Australia 6152

Abstract

Wilson, Paul G. The classification of the genus Waitzia Wendl. (Asteraceae: Gnaphalieae).
Nuytsia 8(3): 46\-An (1992). The circumscription of Waitzia is discussed and a narrow generic
concept proposed. Five species and two infraspecific taxa are recognised. A key is provided and full
synonymy given. One new species combination, W. nitida, is made: two new varieties are described.
Waitzia citrina, W. conica and W. paniculata are excluded from the genus.

Introduction

During an ongoing investigation into the taxonomy of the Australian species placed in Helipterum
DC. and Helichrysum Miller it became evident that the genus Waitzia Wendl. (1808) should also be
studied since some of the species included in it appeared to be closely related to species of Helipterum
and Helichysum. Further investigation indicated that as currently accepted it was polyphyletic.
However, with three of the taxa transferred to other genera it consists of five closely related species.
The characters that have been used to discriminate these related species in Floras and in revisional
papers over the past 150 years have not permitted them to be correctly distinguished. Moreover an
examination of specimens in Australian and some ovenseas herbaria, including specimens studied by
George Bentham (1867) and Ferdinand Mueller (1896), indicated that botanists have always had
confused concepts of the taxa. 1 have therefore investigated the taxonomy of the genus and have
endeavoured to place it in the context of its related taxa.

Since the family Asteraceae will shortly be written up and published in vols 37 and 38 of the Flora
of Australia 1 have only provided descriptions for the infraspecific taxa. Because of the absence of
species descriptions 1 have included in the key rather more detail than is strictly necessary for the
purpose of identification.

Historical

Waitzia Wendl. (1808) was the third generic name published in the Helipterum - Helichrysum
complex that was based on an Australian plant, the two earlier names being PodolepisLuhiil. (1806)
and Podosperma Labill. (1806) {nom.rej. = Podotheca Cass.), both of which arc relatively distinct

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Nuytsia Vol. 8, No. 3 (1992)

from the remainder of the complex (Short 1 989) and do net affect the application of Wendland’s name.
The delimitation of Waitzia is thus clearly of particular significance, since, depending on its
circumscription, it could encompass a few or numerous species in the Helipterum - Helichrysum
complex.

The name Waitzia Wendl. (1808) was based on W. corymbosa, a plant cultivated in the royal
gardens of Herrenhausen in Germany. The origin of the plant was indicated as doubtfully 'Neu-
Holland'. The plant illustrated and described is of a species found only near the west coast of Western
Australia from about 1 60 km north of Geraldton to Shark Bay . It is probably from the Shark Bay region
that the seed originated since this is the only portion of that general area that had been visited by plant
collectors prior to 1808. The generic name was ignored by early 19lh century botanists including de
Candolle (1838) and Bentham (1837) but recognised by Steetz (1845) who correctly synonymized
under it Viraya Gaudich. (1830) and Morna Bindley (1837). Bentham (1837) treated a number of
species under Lcptor/iy/jc/jo.? Less. (1832) which were later accepted as such by de Candolle (1838)
but which Steetz (1844) transferred to Waitzia and Bentham (1867) later accepted as belonging here.

The genus Viraya was described by Gaudichaud (1830) who based it on a plant collected from
Shark Bay. The name was considered by de Candolle (Jan. 1838) to be a later homonym of Vireya
Blume (1826), Vireya Raf. (1814), and Virea Adans. (1763). He also considered that the name should
be spelt Vireya (since it honoured J.-J. Virey) and would then be a precise homonym of Blume’s
generic name. De Candolle therefore placed Viraya Gaudich. ( 1 830) in synonymy under Leptorhynchos
Less. (1832) even though the latter name was published two years after the former. Under
Leptorhynchos two informal sections were recognised by de Candolle; the first 'section' he associated
with the name 'Aphanorhynchos Less. syn. 273' and placed under it those species with short achene
beaks (which included the two syntype species of Leptorhynchos), while the second 'section' he
associated with the name L/ravw Gaudich. and placed under it species with long beaks (which included
species now placed in Waitzia). Lessing does not appear to have published the name to which his
authorship was attached and its use by de Candolle was therefore possibly the result of personal
communication between the two botanists for it is known that de Candolle widely distributed draft
copies of his Compositae 'Prodromus' treatment some years before its publication (Wilson 1989). Use
of the name by de Candolle may also have been due to the long illness that he suffered about this time.
His illness delayed publication of the Compositae treatment and was probably responsible for
numerous errors in its text (Bentham 1873b). The monotypic genus Morna Bindley (1837) was also
recognised by de Candolle (1838) but with an indication of uncertainty since he had seen no material.
Later Endlicher (June 1838) formally published the name 'Leptorhynchus a. Aphanorhynchus'
(without indication of rank) based on Chrysocoma squamata Labill. (the type of Leptorhynchos).
Endlicher also adopted de Candolle’s other informal section but called it 'Leptorhynchus b. Morna',
not 'b. Viraya', since he similarly considered the name Viraya Gaudich. to be illegitimate (a later
homonym) while he recognised Morna Bindley to be a synonym of it.

Meissner ( 1 839) recognisedA/o/vm andLeplorhynchos as separate genera and clearly discriminated
the two. However Reichenbach (1841) once more united them but under the name Aphanorrhynchus
which he published as a replacement for Leptorhynchos Less, since the latter, he explained, was
illegitimate having been used already for a genus of insects; he again recognised Morna as an
infrageneric group. These infrageneric names were given the formal rank of sectio by Pfeiffer (1 873).

Therefore, until 1 845 , botanists recognised only one genus for the taxa now pi\nced\nLeptorhynchos
and in Waitzia (the recognition oiMornahy de Candolle and Meissner being due to lack of personal
knowledge of its type species). Since the generic name Waitzia had been overlooked, and because

Paul G. Wilson, Waitzia

463

Viraya Gaudich. (1830) was considered to be a later homonym, the name Le/jtor/;y/ic/jo5 (1832) was
by some botanists accepted as correct; while those who considered that plant names should not
duplicate names already applied to animals the name Aphanorrhychus Reichb. (1841) was accepted.

In 1845 Steetz resurrected the name Waitzia Wendl. (1808) and clearly distinguished the genus
from Leptorhynchos on the lines previously adopted by Meissner. He also described the monotypic
genus Plerochaeta which he considered to be intermediate between Waitzia and Helipterum. This
name was synonymized under Waitzia by Bentham (1867) along with Viraya and Mania.

Changes made since Bentham (1867) have been at the species level. In 1883 Mueller misapplied
the name Waitzia podolepis (Gaudich.) Benth. to a plant collected by 'Polak' near the Gascoyne River
and provided a description. The species represented by this collection has since been described as
Waitzia conica B.Turner(1966). Mueller posthumously (1896) published corrections to the various
Waitzia species names used by Bentham and these amendments were later accepted by Diels and
Pritzel (1905) who also discu.ssed the ecology and distribution of the various taxa. They indicated
that W. steetzianai'i.e. W. citrina) was distinct from Waitzia .^..^tr. and a derivative of Helipterum while
W.paniculata they considered to be of a very different origin again.

I agree with Diels and Pritzel that both Waitzia citrina and W. paniculata are anomalous in this
genus and, along with the subsequently described W. conica, I have transferred them to other genera
(Wilson 1992a). WaitziapaniculataislramferredloihcmonotypicgenusPlerochaetaosP. paniculata
Steetz, W. citrina to the expanded genus Rhodanthe as R. citrina (Benth.) Paul G. Wilson, and
W. conica to the new genus Haptotrichion as H. conicum (B. Turner) Paul G. Wilson.

Generic affinities

The genus Waitzia was placed in the tribe Gnaphalieae subtribc Helichryseae by Bentham (1867)
and later (1873a) in the tribe Inuleae ('Inuloideae') sublribe Gnaphalieae series Helichryseae. The
latter classification at tribal level has been accepted until recently. Anderberg (1989) provided a
history of the various classifications and proposed a reassessment of the lnule.ae sensu lato in which
the Gnaphalieae s.str. is recognised as a distinct tribe. The genus Waitzia, Anderberg concluded,
should be placed in the Gnaphalieae s.str. to form part of the 'Waitzia clade' along with the genera
Leptorhynchos, Podolepis, Asteridea, and some Australian Helipterum and Helichrysum species.
Anderberg later (1991) indicated that Waitzia should be placed in the Waitzia group of the subtribe
Angianthinae along with Gratwickia, Chrysocephalum, Leptorhynchos, Asteridea, Podolepis, and
Triptilodiscus all of which had involucral bracts with divided stereomes; he pointed out that as
currently circumscribed Waitzia was polyphyletic and that W. citrina differed from the other species
in having an undivided stercome and in lacking the diagnostic trichomes on the achenes.

I concur with Anderberg as to the relationship of Waitzia sensu stricto to these genera and accept
his tribal :uid subtribal classification.

The position of Waitzia within the 'Waitzia group' requires an explanation. The genus Waitzia had
beena.ssociated by Bentham (1867, lH73a)andhy later authors whhLeptorhynchosLess., Helipterum
DC., Helichrysum Miller, Ixiolaena Benth., and Asteridea Bindley (as Athrixia).

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Nuytsia Vol. 8, No. 3 (1992)

The two genera Leptorhynchos and Asteridea form a close knit group the species of wh ich possess
organs that share a number of similar characters. The nan ow elliptic achenes (with or without a neck)
have a thin translucent pericarp with 2-celled tooth-like papillae with one cell overtopping the other
(Anderberg 1989), a thick soft testa free from the pericarp, and persistent barbellate pappus bristles;
the corolla tubes are curved and overhang the involucre with the vascular strands passing to the apex
of the corolla lobes; the apical appendages of the anther are small (c. 0.2 mm long) and are made up
of unthickened cells, and the style apices are relatively small and truncate.

Waitzia, in the strict sense, consists of plants with linear leaves, an arachnoid and glandular-
stipitate indumentum, capitula with scarious bracts that have terete glandular stipes (linear in the
innermost two series), apical appendages of anther relatively large (ovate, c. 0.5 mm long, with
narrow-oblong unthickened cells), deltoid to narrowly elliptic style apices in which the stout vascular
strand extends to the tip, achenes with slender necks and 2-celled tooth-like papillae in which the lower
cell overtops the upper, pericarp thin and transparent, testa thick and ruminate with oblong crystals
and with oblong epidermal cells that are regularly corrugate on their margins, pappus bristles
barbellate and shortly united at the base with the pappus eventually deciduous as a whole from the
apex of the achene neck.

The genera Heiipterum .and Helichrysum (in Australia) are a polyphyletic assemblage of species
(Anderberg 1989, Wilson 1989) and it is to .some members of these genera, in particular to the
Heiipterum albicans group of species, that Waitzia is clearly allied. The Heiipterum albicans group,
which is referred to the genus Leucochrysum (DC.) Paul G. Wilson (Wilson 1992b), consists of
Heiipterum albicans (A.Cunn.) DC., H. fitzgibbonii F. Muell., H. albicans var. graminifolium
Paul G. Wilson, H. molle (DC.) Paul G. Wilson, .and H. stipitatum (F.Muell.) Benth. These five
species resemble Waitzia in indumentum, leaf-shape, and in the morphology of the involucral bracts,
corolla, anthers, and style apices. The species differ from Waitzia s.str. in habit, in having plumose
pappus bristles that are shed by bre.aking shortly above the base, in the lack of an achene neck, and
in the absence of 2-cellcd tooth-like achene papillae.

rvlorphological characters

The characters used in the past to distinguish the species of Waitzia sect. Waitzia have been based
largely on the colour and shape of the involucral bracts. This has led to some confusion since two
of the species have both white and yellow variants and a range of bract shapes. It has therefore been
necessary to incorporate other characters into the discrimination of the species. These additional
characters are found in the indumentum, capitulum shape, and in the .'irrangement, relative size, and
surface texture of the bracts.

Indumentum. All species have some septiate-flagellate hairs (Ramayya 1962) on leaves and stem. The
flattened oblong cells of the bi.seriate to uniseriate proximal portion of the hairs vary in number with
species and with their position on the plant while the uniseriate filiform distal portion varies
considerably in length. Depending on the density of the hairs and the lengths of the proximal and distal
portions the indumentum can be rough or smooth and thinly to densely arachnoid.

Most of the species also have biseriate or uniseriate capitate glandular hairs on the leaves and stem,
or these may be restricted to the branches of the inflorescence. In Waitzia acuminata and W. nitida

Paul G. Wilson, Wailzia

465

they are absent (except on the stipes of the involucral bracts). Both types of hairs are sometimes shed
above the base to leave a short hard projection that produces a rough surface to the leaf. The glandular
hairs on the stipes of the bracts have a single globular cell at the apex.

Sessile globular glandular hairs arc present on the leaves and branches of some species in addition
to the longer hairs.

Leaf shape. In all species the leaves are sessile, linear to very narrowly oblong (or the ba.sal leaves
very narrow-obovate) and usually with the margin recurved, at least on drying. The upper leaves may
be slightly decurrent and sometimes semiamplexlcaul. Variation within a species is considerable and
probably largely environmentally induced since plants in cultivation have, in general, broader and
flatter leaves than their wild counterparts.

Involucre. This can be hemispherical, cup-shaped or, in W. acuminata, turbinate at the base with the
involucral bracts descending along the peduncle.

Involucral bracw (see Figure 1). The bracts, except for the innermost two series, have stipe-likeclaws
that are shortly glandular pilose and arachnoid; the lamina varies from ovate-cordate to narrowly
triangular, obtuse to acute or acuminate, entire to fimbriatc-serrate. The two innermost series of bracts
have linear claws with a shortly glandular pilose stereome and a narrow scarious margin; the lamina
is very small, scarious, and white or coloured.

The surface of the laminae may be smooth or minutely roughened due to the emergent distal ends
of some of the epidermal cells. In W. corymbosa the two innermost series of involucral bracts lengthen
during anthesis to well exceed the intermediate and outer bracts. The outer and intermediate
involucral bracts may remain erect, or become reflexed or spreading during anthesis.

Pappus (see Figure 2). The pappus bristles are barbellate (or pilose at the base) and eventually
deciduous as a whole from the achene neck. They are generally colourless, white, or faintly tinged
with yellow near the base, except in W. nitida in which the distal half of the pappus is always pale
yellow.

Mycorrhizal relationship. Waitzia species form ectomycorrhiza as do species in Leucochrysum and
in Leptorhynchos (Warcup 1990). In this they differ from species in the Rhodanthe complex.

Achene venation. The species ofWaitzia have a very short vascular strand in the testa which contrasts
with the situation in Leucochrysum in which the strand passes to the apex of the seed. The vascular
strands of the testa and pericarp are arranged in a lateral position in relation to the cotyledons which
is the same position as is found in Leucochrysum.

Chromosome numbers. Turner (1970) has recorded a count of n = 10 for W. nitida and of n = 12 for
W. suaveolens. No count has been published for the species now placed in Leucochrysum.

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Nuytsia Vol. 8, No. 3 (1992)

Hybridization

Hybridization between species appears to be rare in Waitzia and only two possible cases have been
observed.

A collection from Kojonup {Meebold 7323, PERTH) consists of specimens that appear to have
been gathered from a hybrid swarm involving W. acuminata subsp. acuminata and W. suaveolens var.
suaveolens.

Some specimens gathered between Kalbarri and Geraldton appear to be intermediate between
W. suaveolens var. suaveolens tvni W. acuminata subsp. albicans. They have bracts that are only very
faintly scabridulous and are somewhat intermediate in shape between those of the putative parents,
while some of the specimens lack the stipitate glands that are present in W. suaveolens but absent in
W. acuminata. Examples are: R.C.Wemm 11 1 IB, D. & N. McFarland 1203, M.G.Corrick 8122, and
A.J. Cough 269, all PERTH.

Both of the putative parents are found near the area where these specimens were collected.

Waitzia Wendl.

Wendl., Coll. PI. 2:13 1.42(1808). T: Waitzia corymhosa Wendl.

Viraya Gaudich. in Freyc., Voy. Uranie 466 1.89(1830); Post et Kuntze, Lexicon Gen. Phan.
590(1903) 'Vireya' pro syn. sub Waitzia. - Leptorhynchos sect. Viraya (Gaudich.) DC. ex Pfeiffer,
Norn. Bot. 2:86(1874). T: Viraya podolepis Gaudich.

Morna Lindley, Bot. Reg. 1.1941(1837). - Aphanorrhynchus 2. Morna (Lindley) Reich., Deut. Bot.
Herb.-Buch. 1:90(1841) nom.illeg. - Aphanorrhynchus sect. Morna (Lindley) Reich, ex Pfeiffer,
Norn. Bot. 1:229(1873) nom.illeg. - Leptorhynchos b. Morna (Lindley) Endl., Gen. PI. 445(1838). -
Leptorhynchos sect. Morna (Lindley) Endl. ex Pfeiffer, Norn. Bot. 1:229(1873). T: Morna nitida
Lindley.

Aote.- The generic ntsmt Aphanorrhynchus (Endl.) Reich. (184 1) is ultimately basedonLc/;to/7iy«c/i05
Less. (1832). The latter name was considered by Reichenbach to be illegitimate since there existed
an insect genus Leptorrhynchus.

Annual erect herbs, sparsely to moderately cobwebby with septate flagellate hairs with or without
stalked glandular hairs (that may leave a hard base when shed to form a scabrid surface). Leaves
alternate, linear to narrow-oblong, sessile, margin mostly recurved on drying. Inflorescence terminal,
cymose, the uppermost leaves grading into linear capitula-subtending bracts (or these with scarious
narrow-elliptic laminae) that grade into the outer involucral bracts. Involucre ± hemispherical to
turbinate. Outer and intermediate bracts multi.seriate, stipitate; stipe terete with divided stereome,
with stalked glandular hairs and ± cobwebby; lamina scarious, ovate or narrow-triangular, entire to
serrate, smooth or minutely scabridulous from the emergent distal apices of the epidermal cells, white
to violet, yellow or gold, erect or at length spreading or reflexed on the stipe. Innermost bracts erect,
shorter than or exceeding intermediate bracts; stipe linear with narrow scarious margin and prominent

Paul G, Wilson, Waitzia

467

Figure 1. Intemiediate and inner involucral bracts of Waitzia species. A - W. nitida. B - W. podolepis. C - W. corymbosa.
D - W. acuminata var. acuminata. E - W. suaveolens subsp. suaveolens. (a - intemiediate bract; b - imier bract.)
/KimmR.SaffreyXM. B from W. Blackali 563. C from P Wilson 1219&. D from P. W(7io« 12036. E from 7. AToc/i 465.

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Nuytsia Vol, 8, No. 3 (1992)

midrib with stalked glandular hairs; lamina very short otherwise either similar to the intermediate
bract-laminae or hyaline and minute. Receptacle rounded, glabrous. Florets numerous, bisexual,
actinomorphic. Corolla tube narrow-cylindrical, turbinate above, prominently 5-lobed, glabrous
except for short glandular hairs on outside of lobes, smooth within; cells of inner epidermis of throat
regularly and prominently undulate; vascular strands extending to base of lobes. Anthers: appendage
narrow-ovate, c. 0.5 mm long; cells narrow-oblong, thin-walled; tails slender, firm, ± equal to collar.
Style apex deltoid to narrow elliptic; vascular strand stout and extending to tip. Achene compressed
ellipsoid with slender neck, almost smooth or barbellate with myxogenic 2-celled papillae; neck
glabrous, barbellate or shortly pilose. Pericarp thin, translucent, collenchymatous, of narrow-oblong
cells, in most species myxogenic; vascular strands laterally oriented with reference to cotyledons.
Testa free from pericarp, weakly coriaceous, brown, with scattered crystals; apex laterally flattened
and sterile; outer epidermis of narrow-oblong cells prominently and regularly undulate on margin;
vascular strand confined to base or to near base of seed, laterally orientated. Pappus bristles numerous,
shortly united at base, dentate, sometimes pilose at base, eventually deciduous as a whole from achene
neck.

Five species endemic to Australia.

Key to species and infraspecific taxa

1. Inner involucral bracts yellow, outer yellow, orange or straw-coloured

2. Pappus yellow in upper half; upper leaves and branches cobwebby,

not glandular; bracts yellow 3. W. nitida

2. Pappus white or colourless in upper half; upper leaves and
branches cobwebby or shortly glandular pilose; intermediate
bracts yellow, outer bracts yellow or orange

3. Upper leaves and branches cobwebby, bracts minutely
scabridulous and descending down peduncle (Figure lA),

capitula not subtended by linear herbaceous bracts 1. IT. acuminata var. acuminata

3. Upper leaves and branches shortly glandular pilose;
bracts smooth; capitula subtended by linear herbaceous

bracts 5. W. suaveolens vwt.flava

1. Inner involucral bracts white, outer bracts white, pink, orange,
or brown

4. Innermost bracts with pale brown mottled laminae

5. Innermost bracts long exceeding the pink and white

intermediate bracts; lamina of innermost variably crinkled,
prominently ciliolate (Figure 1C); outer and intermediate

bract-laminae soon reflcxed on stipes, long acuminate 2. W. corymbosa

5. Innermost bracts ± equal to the intermediate bracts, lamina
crinkled, minutely dentate, not ciliolate (Figure IB); outer
and intermediate bract-lamina while or straw coloured, not or

tardily spreading, acute to obtuse 4. W. podolepis

4. Innermost bracts with white or hyaline laminae

Paul G. Wilson, Waitzia

469

6. Capitula not obviously subtended by linear herbaceous
bracts. Involucral bracts acuminate, slightly rough
(Figure ID); upper branches and leaves somewhat

cobwebby not shortly glandular pilose i,w. acuminata var. albicans

6. Capitula subtended by linear herbaceous bracts.

Involucral bracts acuminate to acute or obtuse
(Figure IB); upper branches and leaves shortly

glandular pilose with or without cobwebby hairs 5. W. suaveolens var. suaveolens

t

1. Waitzia acuminata Steetz in Lehm., PI. Preiss. 1:453(1845). (Figure 2). Type citation: 'In
Australasia orientali-occidentali leg. c. Preiss. Herb. Preiss. sine No. Roe! Drummond! (V.s. in herb,
aulico Vindobonnense!)'. Lectotype (here chosen): Roe s.n. (W); syntypes: Preiss s.n. (LD
MEL 221975), Drummond 286 (W).

[Waitzia coijmbosatuicX. non Wendl.: Benth.,Fl. Austral. 3:635(1867); Chittenden, Diet Gardening
edn 2, 4:2258(1977)]

var. acuminata (Figures ID, 2)

Waitzia discolor Twez., Bull. Soc. Imp. Naturalistes Moscou 24/1:194(1851). T: J. Drummond Aih
coll. n. 198 (holo: KW photo seen; iso: MEL 1585193).

Involucral bracts orange-yellow.

Distribution. Western Australia south of the 23° latitude, southern Northern Territory, South
Australia, north western Victoria, western New South Wales, and extreme southern central and
western Queensland.

var. albicans Paul G. Wilson, subsp. nov.

Bracteae involucri albae vel bracteis exterioribus violaceo-rubrae.

Typus\ Western Australia, 21 km from Northampton towards Port Gregory, "buds deep pink,
becoming more pale as head opens and eventually white, disc florets yellow, numerous plants amongst
tow scrub on rocky outcrop', 6 Oct. 1972, S.Paust 1258 (holo: PERTH; iso: AD CANB K MET,
NSW, S). > . .

Involucral bracts white or the outer violet-red.

Distribution. Western Australia, near the west coast between York and Hamelin Pool (c 31° 30’S
to 26° 30’S).

The distribution of the two varieties overlaps between Hamelin Pool and Geraldton but I have seen
no evidence to suggest that in this area they grow together in the same locality. However, collections
made by J.H.Gregory at Northam in 1901 and by Keighery & Alford c. 17 km west of York in 1985
(both PERTH) are of a mixture of both varieties. These two localities are at the southernmost

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Nuytsia Vol. 8, No. 3 (1992)

Figure 2. A & C - Wailzia suaveolens. B - IV. acuminata. D - W. corymbosa. E-W. podolepis. F - IV. nitida. (a - acliene with
pappus; b - apex of pappus bristle; c - barbellate hairs of achene neck; d - seed removed from pericarp; e - surface view of testa
showing undulate cells and calcium oxalate crystals; f- basal portion of testa; g- achene; h- anther; i- anther appendage;] - style
apex; k - surface view of cells of pericarp.)

A & C from R. floyce 4731. B from A. S. George 8437. D from P. Wilson 12198. E from //. Demarz 8996. F from R. Saffrey 147.

Paul G. Wilson, Waitzia

471

distribution of var. albicans and the westernmost distribution (at that latitude) of var. acuminata.
Where the two varieties grow together they differ only in the colour of the involucral bracts.

2. Waitzia corymbosa Wendl., Coll. PI. 2:13, t.42(1808). - W. corymbosa f. wendlandiana Steetz,
PI. Preiss. 1:451(1845) excluding specimen cited, nom.illeg. - W. corymbosa var. wendlandiana
(Steetz) Diels & Pritz., Bot. .lahrb. 35:626(1905) nom. illeg. Type citation: 'Das Vaterland: Neu-
Holland?' Leclotype (here chosen): Wendland op.cit. t. 42. (Figures 1C, 2D)

Distribution. Shark Bay south to Kalbarri.

The name Waitzia corymbosa has been incorrectly u.sed by previous authors. The species was
unknown to Bentham (1867) who misapplied the name to plants of W. acuminata, and unknown to
Mueller (1896) who misapplied the name principally to collections of Waitzia suaveolens var.
suaveolens. Diels and Pritzel (1905) considered it to be conspecific with W. nivea (i.e. to
W. suaveolens). The name was omitted by Grieve and Blackall (1975) who included the species under
the misapplied name W. podolepis.

The specimen on which Wendland based his description could not be found in the Wendland
herbarium (herb. GOET),/iVic G. Wagenitz. pers. comm. 1 have therefore designated the illustration
that accompanied the description as the Icctotype.

3. Waitzia nitida (Bindley) Paul G. Wilson, comb. nov. (Figures lA, 2F)

Morna nitida Bindley, Bot. Reg. t.l941(l March 1837). Type citation: 'inhabiting the dry country
about the Swan River, whence it was introduced in the year 1835, by Sir James Stirling. The first time
it was publicly seen in this country was at one of the great exhibitions held in the Garden of the
Horticultural Society in 1836, when the judges awarded to Robert Mangles, Esq. who exhibited it,
a Knightian Medal.' Lectotype (here chosen): Right-hand specimen on sheet labelled 'TYPE. Morna
nitida Bindl.' (CGE, photo seen). See note below.

Leptor/iync/joiaMrcMiBenth.inEndl. e/a/.,Enum.Pl.Hugel64(April I'&'il). -Waitzia aureai^QnW.)
Steetz in Behm., PI. Preiss. 1:452(1845). Type citation: 'King Georges Sound et Swan-River.
(Hugel.". Lcctoty/;e (here chosen): 'Freemantle', //age/ (W,isolectotype:K);iy«t>’pe.- 'King George’s
Sound', lliigel (W).

? Waitzia grandiflora W. Thompson in T. Moore, Florist & Pomologist 4:41 & tab.(March 1865);
Naudin, Revue Hort. (Paris) 185( 1865);Chittenden, Diet. Gardeningcdn 2, 4:2258(1977). Typification:
'I received this fine species from Dr. F. Mueller, of Melbourne'.

Note: Bindley initially (1 March 1837) stated that Morna nitida was introduced in 1835 by James
Stirling, the governor of Swan River Colony. He subsequently (Bot. Reg. 23: sub tab. 1944, 1 April
1837) corrected this .statement by adding a note written by Donald Mackay, the gardener of Robert
Mangles. In this note Mackay indicated that seeds of Morna nitida were received by Robert Mangles
early 1836 and sown on 2 February 1836. A sheet in CGE has a label stating that it is the type of AJorna
nitida\ the sheet bears two specimens the smaller of which has 'R. Mangles Esq.' written beside it while
the other has a printed label attached which reads 'Swan River/ Capt. James Mangles, R.N.'. James
was the brother of Robert Mangles in whose garden the plant was first grown. I have designated as
lectotype the right-hand specimen with the James Mangles label but it is likely that the specimens had
a common origin.

472

Nuytsia Vol. 8, No. 3 (1992)

Waitzia grandiflora was described as having a capitulum much larger than that of W. aurea (i.e.
of W. nitida) and to be almost glabrous. It is probably a variant of W. nitida. No material under the
name W^. grandiflora has be located in Australian herbaria nor in K or BM (fide G. Leach pers. comm.).
Although the name is still used in horticulture in Europe (see above) I have been unable to discover
to what species it is being applied.

Chromosome number: n=lO fide Turner (1970).

Distribution. Western Australia: Shark Bay (? Broome) south to Albany and Ravensthorpe and east
to Merredin (? Kalgoorlie). Two collections are labelled as having come from localities that are
outside the distribution range as otherwise recorded for this species; they are as follows: E, Kelso,
1900, Broad Arrow, north ofKalgoorlie (PERTH); Roebuck Bay (near Broome), pre 1867, £>/- Marten
(MEL 1585029).

Waitzia nitida shows very little variation throughout its range and does not intergrade with any
other species.

The concept of W. n itida (as W. aurea) held by previous authors including S teetz ( 1 845), Bentham
( 1867), Mueller (1896), Diels & Pritzel (1905), and Grieve & Blackall (1975) also included specimens
of W. suaveolens var.flava which may be most readily distinguished by its colourless pappus.

4. Waitzia podolepis (Gaudich.) Benth., FI. Austral. 3:637(1867). - Viraya podolepis Gaudich. in
Freyc., Voy. Uranie466 t.S9(lS30).- Leptorhynchos podolepis (Gaudich.) DC., Prod. 6:160(1838).
Lee to type (here chosen): Collected by Gaudichaud at S hark Bay (P 'Bale de Ch. Marin, C. Gaudichaud').
(Figures IB, 2E)

[W. corymbosa auct. non Wendl.: F. Mueller, Z. Allg. osterr. Apotheker-Vereines 34(36):935(1896)

p.p.]

Distribution. Western Australia: Irwin River north to Shark Bay.

An accurate description of this species was provided by Diels and Pritzel (1905) based on an
original collection in herb. B which evidently came from Kunth’s herbarium. They pointed out that
Mueller (1896) had misapplied the name Waitzia podolepis to a J. Pollack collection of an
undescribed species that was new to them. This unknown species was described by Turner (1966)
under the name Waitzia conica. The microfiche photograph of the possible type specimen seen by
de Candolle (G-DC) and cited by him in the Prodromus l.c., is not sufficiently clear to confirm its
identity, however, the description provided by de Candolle is accurate.

5. Waitzia suaveolens (Benth.) Druce, Bot. Exch. Club Soc. Brit. Isles 4:652( 19 17). - Leptorhynchos
suaveolens Benth. in Endl. et al., Enum. PI. Hiigel 64(1837). - Type citation: 'Swan-River (Hiigel.)'
non vidi.

No type material of this name has been located in the herbaria W, BM, and K. The application
of the name is based on the description and on contemporary collections made around Perth in the
areas visited by Hugel.

Paul G. Wilson, Waitzia

473

var. suaveolens (Figures IE, 2A , 2C)

Morna nivea Lindley, Bot. Reg. 24:t.9(Feb.l838). - Waitzia nivea (Lindley) Benth., FI. Austral.
3:636(1867). Type citation: 'raised from Swan River seeds in the garden of Robert Mangles, Esq.,
of Sunning Hill.' (holo: CGE 'Hort. Mangles July 1837' photo seen).

Waitzia corymbosa f. benthamianaSlQQtz in Lehm., PI. Preiss. 1 :45 1 ( 1 845). Based on Leptorhynchos
suaveolens Benth., Morna nivea Lindley, L.Preiss 12, Hiigel s.n., and F £auer s.n. Lectotype (here
chosen): Leptorhynchos suaveolens Benth..

? Helichrysum rigidulum DC., Prod. 6: 193(1838). T: Nova Hollandia, Cult. 1832, Sweet (holo: G-DC
photo seen).

Waitzia odontolepis Turcz., Bull. Soc. Imp. Naturalistes Moscou 24/1:77(1851); Chittenden, Diet.
Gardening edn 2, 4:2258(1977). T: Western Australia, J. Drummond 5th coll. no. 382 (holo: KW
photo seen; iso; MEL).

[Waitzia corymbosa WiC\..x\on'WQn(\\.:¥. Muell.,Z. Allg.Osterr. Apotheker-Vereines34(36);935(1896)
p.p. maj.; Diels & Pritzel, Bot. Jahrb. 35:626(1905).]

[Waitzia corymbosa f. wendlandiana auct. non Steetz: Steetz in Lehm., PI. Preiss. 1:451(1845)]

Upper branches and leaves cobwebby or glandular stipitate. Corymbs congested or loose. Outer
involucral bracts acuminate to acute, white, straw coloured, or purplish pink; inner involucral bracts
obtuse to acute, white. Pappus colourless.

Chromosome number: n = 12 fide Turner (1970), see comment below.

Distribution. South west Western Australia: Geraldton to the south coast and east to Esperance.

Note 1 . The description oi Helichrysum rigidulum by de Candolle is not sufficiently detailed, nor the
microfiche photograph of the type in G-DC sufficiently clear, for me to be sure that this name is
synonymous with W. suaveolens.

Note 2. A western entity of var. suaveolens has obtuse inner involucral bracts whereas an eastern entity
has acute (or even acuminate) inner bracts, however, the extremes of the two (which also differ in the
degree of glandular indumentum, size of capitula, and openne.ss of inflorescence) grade into each
other. The easlcrn more glandular entity has smaller capitula and more open inflorescences, it is in
some areas virtually identical with var.flava except in colour of bracts.

var. flava Paul G. Wilson, var. nov.

[Waitzia aurea auct. non (Benth.) Steetz: Benth., FI. Austral. 3:636(1867) p.p; F. Muell, Z. Allg.
Osterr. Apotheker-Vereines 34(36):935(1896) p.p; Grieve & Blackall, How to know Western
Australian Wildflowers pt 4:844(1975) /;./;.]

Bracteae involucri flavae, anguste triangulares acutae vel acuminatae.

474

Nuytsia Vol. 8, No. 3 (1992)

Typus: Mt Ridley, Western Australia, granitic slope, 7 August 1970, K.M. Allan 360 (holo: PERTH;
iso: CANB,K).

Upper branches and leaves glandular stipitate. Corymbs loose. Involucral bracts narrowly
triangular, acute or the inner acuminate, smooth, yellow. Innermost bracts with lamina ovate to
oblong, scarious or pale yellow. Pappus colourless.

Distribution. Norseman area south-east to Balladonia and west to Stirling Range.

Specimens of vtxr.flava have been generally referred in herbaria and in literature to W. aurea. The
latter species differs in having an cobwebby indumentum without glandular hairs, in having obtuse
bracts (not acute to acuminate) and in having yellow pappus bristles (not colourless); in addition the
capitula of W. aurea are larger than those of yar.flava (but of a similar size to the capitula of the western
entity of var. suaveolens).

As noted above, vta.flava differs from some forms of the eastern entity of var. suaveolens almost
solely in the colour of the bracts. The two varieties may sometimes be found growing together and
it was from such a mixed population that Turner, l.c., recorded the chromosome number of n = 12.

Excluded Names

Waitzia brachyrrhyncha F.Muell., Linnaea 25:407(1853) = Leucochrysum molle (Cunn. ex DC.)
Paul G. Wilson, Nuytsia 8:444(1992).

Waitzia brevirostris Steetz in Lehm., PI. Preiss. 1:451(1845) = Rhodanthe citrina (Benth.)
Paul G. Wilson, Nuytsia 8:407(1992).

Waitzia citrina (Benth.) Steetz in Lehm., PI. Preiss. 1;454(1845) = Rhodanthe citrina (Benth.)
Paul G. Wilson, op. cit.

Waitzia conica Turner, Sida 2:428(1966) = Haptotrichion conicum (B. Turner) Paul G. Wilson,
Nuytsia 8:425(1992).

Waitzia dasycar pa JuTcz., Bull. Soc. Nat. Moscou 24:71 (IS51) = Rhodanthe citrina (Benth.)
Paul G. Wilson op. cit.

Waitzia paniculata (Steetz) Benth., FI. Austral. 3:(1867) = Pterochaeta paniculata Steetz (see
Nuytsia 8:422(1992).)

Waitzia steetziana Lehm., PI. Preiss. 1:454(1845) = Rhodanthe citrina (Benth.) Paul G. Wilson,
op. cit.

Wa/L/axM/plfMrca Steetz in Lehm., PI. Preiss. l:453(1845)=R/mf(flnt/;ccitn/ia (Benth.) Paul G. Wilson,
op. cit.

Waitzia tenella Hook., Bot. Mag. t.5342(1862) = Rhodanthe citrina (Benth.) Paul G. Wilson, op. cit.

Paul G. Wilson, Wailzia

475

Acknowledgements

I am grateful to the staff at the herbaria of Lund (LD) Paris (P) and Vienna (W) for the loan of
material, to Kew (K) for the donation of cibochromes of some type specimens, and to the National
Herbarium at Melbourne (MEL) for providing me with facilities and access to its collections. Greg
Leach, while Australian Botanical Liaison Officer, meticulously searched out and provided me with
details of types held in Kew (K), Cambridge (CGE) and the Natural History Museum (BM). Arne
Anderberg kindly read my manuscript and suggested a number of improvements. Phil S hort suggested
a number of alterations and additions which I have adopted. I thank Margaret Menadue for providing
the illustrations.

References

Anderberg, Arne A. (1989). Phylogeny and reclassification of the tribe Inuleae (Asteraceae). Canadian J. Bot. 67:2277-2296.
Anderberg. Arne A. (1991). Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Botanica 104:1-195.
Bentham, G. (1837). Compositae. In S.L. Endlicher et al., "Enumeratio Pltintarum." (Beck University Biblio., Vienna.)
Bentham, G. (1867). "Flora Australiensis." vol. 3. (Lovell Reeve & Co.. London)

Bentham, G. (1873a). Compositae. In "Genera plantamm" Vol. 2. Edited by G. Bentham and J.D. Hooker. (Reeve & Co.,
London.)

Bentham, G. {1873b). Notes on the classification, history, and geographical distribution of Compositae. J. Linn. Soc. Botany
13:335-577.

Candolle, A.P. de (Jan. 1838). "Prodromus syslematis n.aturalis regni vegetabilis." Vol. 6. (Treuttel & Wurtz, Paris.)

Diels, L. & Pritzel, E. ( 1904-1905). Fragmenta Phytographiae Au.straliae occidentalis. Bot. Jahbr. 35:55-662.
Endlicher.S.L. (June 1838). Genera plantarum secundum ordinesnaturalesdisposita. Vol. 1. (Beck Univ. Biblio., Vindobonae.)

Gaudichaud-Beauprd, C. (1826-1830). Botanique. In L. de Freycinet, "Voyage autour du monde,...sur les corvettes de S.M.
rUranie et la Physicienne." (Pillet, Paris.)

Grieve, B. J. & Blackall.W.E. (1975). "How to know Western Australian WildlJowers." Part IV. (University of Western Australia
Press, Perth.)

Meisner, C.F. (1839). "Plantarum vascularium genera." Vol. 1. (Weidmannia, Leipzig.)

Mueller, F. (1883). "The plants indigenous around Sharks Bay and its vicinity.” (Government Printer, Perth.)

Mueller, F. (1896). Bemerkungcn liber das Genus Wailzia und seine Arlen. Zeitschrift des allgem. dsterreich. Apotheker-
Vereines 34 (no.36):933-936.

Pfeiffer. L.K.G. (1873-1874). "Nomenclator botanicus." 2 vols. (Fischer, Cassellis.)

Ramayya, N. (1962). Studies on the trichomes of some Compositae 1. General structure. Bull. Bot. Surv. India 4:177-188.
Reichenbtich, H.G.L. (1841). "Dcrdeutsche Botaniker." Vol. 1. (Arnold. Dresden & Leipzig.)

Short, P.S. (1989). A revision of Podoiheca Cass. (Asteraceae: Inuleae: Gnaphaliinae). Muelleria 7:39-56.

Steetz, J. (1845). Compositae. /n J.G.C. Lehmann, "Plantae Preissianae." Vol. 1:417-490. (Meissner, Hamburg.)

Turner, B.L. (1966). Wailzia conica (Compositae-Inuleae), a new species from Western Australia. Sida 2:428-430.

Turner, B.L. (1970). Chromosome numbers in the Compositae. XII Australian species. Amer. J. Bot. 57:382-389.

Wilson, Paul G. (1989). A revision of the genus Hyaiosperma (Asteraceae: Inuleae: Gnaphaliinae). Nuytsia 7:75- 101.

Wilson, Paul G. (1992a). The classific.alion of the Australian species currently included in Heitpleriim and related genera
(Asteraceae:Gnaphalieae): Part 1. Nuytsia 8:379-438.

Wilson, Paul G. (1992b). 'Hie classification of Australi.m species currently included in llclipleritm (Asteraceae:Gnaphalieae):
Part 2 Leucochrysam. Nuytsia 8:439-445.

476

Nuytsia Vol. 8, No. 3 (1992)

Index

New names in bold, other accepted names in roman, synonyms in italics.

Aphanorrhynchos Less.. 462
Aphanorrhynchus (Endl.) Reich. 463, 466
sect. Morna (Lindley) Pfeiffer 466
Chrysocorm squamala Labill. 462
Helichrysum Miller

rigidulum DC. 473
Helipterum DC.

molle (DC.) Paul G. Wilson 464
Leptorhynchos Less. 462
Benth. 471

podolepis (Gaudich.) DC. 472
iwaveo/e/u Benth. All.Al'i
Morna Lindley 466

nitida Lindley 47 1
nivea Lindley 473
Pterochaeta Steetz

paniculata Steetz 463
Rhodanthe Lindley
Viraya Gaudich. 466

podolepis Gaudich. 472
Virea Adans. 462
Vireya Blume 462
Vireya Raf. 462
Waitzia 466

acuminata Steetz 469

var. acuminata 469
var. albicans Paul G. Wilson 469
au/ea (Benth.) Steetz 471
brachyrrhyncha F.Muell. 474
brevirostris Steetz 474
citrina (Benth.) Steetz 463,474
co/iicfl B. Turner 463,474
corymbosa Wendl. 471,472,473
f. benthamiana Steetz 473
{. wendlandiana Steelz 471,473
var. wendlandiana (Steetz) Diels & Pritz. 471

Paul G. Wilson, Waitzia

All

dasycarpa Turcz. 474
discolor Turcz. 469
grandiflora W. Thompson 471
nitida (Lindley) Paul G. Wilson 471
nivea (Lindley) Benth. 473
odontolepis Turcz. 473
podolepis (Gaudich.) Benth. 472
paniculata (Steetz) Benlh. 463, 474
steetziana Lehm. 474
suaveolens (Benth.) Druce 472
var. suaveolens 473
var. Hava Paul G. Wilson 473
sulphurea Steetz. 474
tenella Hook. 474

Nuytsia 8(3): 479-483 (1992)

479

A new species of Acomis from the Northern Territory and a new
combination in the genus Thiseltonia (Asteraceae: Gnaphalieae)

Paul G. Wilson

Western Australian Herbarium, Department of Conservation and Land Management,
PO Box 104, Como, Western Australia 6152

Abstract

Wilson, Paul G. A new species of Acomis from the Northern Territory and a new combination
in the genus Thiseltonia (Asteraceae: Gnaphalieae). Nuytsia 8(3): 479-483 (1992). The recognition
of the genera Rulidosis, Acomis, and Thiseltonia is discussed. A new species, Acomis kakadu, is
described from Kakadu National Park, Northern Territory, and a new combination, Thiseltonia
gracillima, is made for the plant previously known as T. dyeri.

Introduction

The name Acomis was published by F. Mueller in 1864 and validated along with Acomis macra
F. Muell. by a generico-specific description, but nowhere in the protologue did Mueller indicate that
a new genus was involved and, in fact, both Bentham (1873) and Mueller (1882 & 1889) attributed
the publication ofAcomw to an earlier (1860) paper by Mueller where, underadescriptionofT^unJoiB
acoma, he indicated that he had used the manuscript name Acomis rutidosea for this species on
herbarium sheets but that he was now placing it in Rutidosis section Acomis. However, since the
sectional name was not accompanied by a sectional description it was not validly published. Possibly
due to the casual manner of its publication the name Acomis was incorrectly attributed to Bentham
(1867) by Ffur et al. (1979) and by Anderberg (1991) while the last author incorrectly indicated its
type as being A. acoma (F.Muell) Druce {=Rutidosis acoma F.Muell.). The correct choice of type is
critical since the type that has been designated by Anderberg (1991) and the type designated by me
have different style shapes and, as is discussed below, with a revised classification could be placed
in different genera.

Mueller later (1893) included Acomis in Humea Sm. (as sect. Acomis) along with Haeckeria
F. Muell. (1853) (as sect. Haeckeria) and Piihocarpa Bindley (1839) (as sect. Pithocarpa) but he
retained Rutidosis as a distinct genus. Mueller also indicated that he was placing Humea gracillima
in sect. Acomis although he had not at that time validly publish the species name.

De Candolle (1838) recogni.sed that the types of Humea Sm. (Dec. 1804) and Calomeria Vent.
(Oct. 1804) wereconspecific and he therefore synonymized the latter name under the former. Heine
(1967) accepted de Candolle’s synonomy but noted that Humea was published shortly after
Calomeria, he therefore made a number of new combinations in the latter genus based on those names

480

Nuytsia Vol. 8, No. 3 (1992)

in Humea recognised by Mueller (1893) and those described later based on African material. All
recent authors (e.g. Lewis &Summerhayes 1951, Grieve 1975, Anderberg 1989, 1991) have followed
Bentham (1867) in regarding Pithocarpa as a distinct genus. Willis (1967), in discussing a number
of taxa related to Cassinia, recognised Calomeria and Haeckeria as separate genera distinct from
Acomis and Rutidosis; his generic taxonomy has been accepted in Australia. The African plants that
had been placed in Calomeria are now considered to be epappose species of Helichrysum (Hilliard
and Bunt 1973, Hilliard 1983).

In 1896 Mueller and Tate formally published Humea gracilUma which Mueller (1893) had
previously mentioned by name only; they suggested that it was 'closely allied to Acomis macra'. The
former taxon was subsequently described independently by Hemsley (1905) as a new genus and
species, Thiseltonia dyeri. It differs from the species currently included in Acomis in being a delicate
minutely glandular-puberulous annual, in having the outer florets female with very slender 4-lobed
corollas, in having (in the bisexual florets) truncate style appendages with a central subulate apex,
and in having a diaphanous papillose peric:up that is united to the pale brown papery testa. In Acomis
the species are more or less woolly, the florets are all bisexual, and the pericarp is thick, crustaceous
and free from the membranous testa.

The species currently placed in Acomis, Rutidosis, and Thiseltonia have involucral bracts with
similar clearly demarcated stereomes which completely enclose the vascular strands, they also have
similar corolla shapes, and similar small anthers in which the tails are fine and difficult to discern.

Although Bentham ( 1 867, 1873) recognised both Acomis and Rutidosis DC. ( 1 838) he considered
their separation to be artificial since it was based solely on the presence (in Rutidosis) or absence (in
Acomis) of a pappus. The species of the two genera can also be divided into two groups based on their
style morphology. In one group the style appendage is truncate while in the other it is narrowly
triangular. In the former group are found the type species of both Rutidosis tmd Acomis , viz. Rutidosis
helichrysoides DC. and Acomis macra F. Muell.; in the latter group are found Rutidosis leucantha
F.Muell. and Acomis acoma (F.Muell.) Druce. Further work may indicate that Acomis and Rutidosis
are best treated as being congeneric, or it may suggest that the generic circumscription within the
complex shouldbe based more on the morphology of the style apex and less on the presence or absence
of a pappus; if the former taxonomy were decided on then the correct name for the complex would
be Rutidosis, if the latter (hen a new generic name would be required for those species with acuminate
style apices including the single Acomis species described in this paper.

Anderberg (1991) comments on the close relationship between AcomA and Rutidosis and on the
marked similarity of these genera to Leptorhynchos tuid Chrysocephalum. I agree with the first
observation but not the second since I consider that the morphology of the achenes and of the
involucral bracts in the first pair of genera is so obviously distinct from the morphology of the achenes
and bracts in the second pair as to provide a clear generic separation.

Acomis

Acomis kakadu Paul G. Wilson, sp. nov. (Figure 1)

Herba erecta ad 40 cm alta, gossypina. Folia linearia, 2-4(8) cm longa. Involucrum hemisphericum;
bracteae multiseriatae, homomorphicae; lamina ovata, alba, 4-8 mm longa. Receptaculum conicum.

Paul G. Wilson, Acomis and Thiseltonia

481

villosum. Flosculi numerosi, homogami. Corolla c. 2.5 mm longa, apicem versus late turbinata. Styli
angustissime triangulares. Achenium doliiforme, c. 1.4mm longum;pericarpumcrustaceum,colliculis
et papillis globosis dense ornatum, bruneum.

Typus: Kakadu National Park, 1 0 km west-south-west of Jabiru East, 1 5 May 1 980, L.A . Craven 5466
(holo: CANB 307405; iso: CANB 307406, PERTH).

Erect annual herb to 40 cm high. Stem single, slender, grey cottony, giving off branches above
the base. Leaves alternate, narrow linear, 2-4(8) cm long, 1 mm wide, very sparsely cottony.
Inflorescence corymbose, peduncles 1-3 cm Icng, cottony, bearing a few foliaceous bracts with
scarious apices that grade into the involucral bracts. Involucre hemispherical. Involucral bracts
multiseriate, loosely arranged, homomorphic: claw broad-oblong, c. 2 mm long, 1 mm wide, sparsely
long-ciliate, filled by a green flat stereome except for the narrow translucent margin; abruptly
separated from the lamina by a horizontal fold; va.scular strand branched, not extending beyond the
stereome; lamina ovate, white, 4-8 mm long. Receptacle conical, c. 3 mm high, smooth, white-
villous. Florets numerous, homogamous, actinomorphic, equal to involucre. Corolla narrow-
cylindrical below and very sparsely pilose, broadly turbinate above, in all c. 2.5 mm long; lobes 5,
triangular, c. 1 mm long, smooth within, very sparsely and minutely glandular puberulous outside,
vascular strands extending to near tips. Anther loculi c. 1 mm long; appendage broad ovate, obtuse,
c. 0.25 mm long, thickened atid obtuse at base; tails absent; collar narrow-oblong, slightly broader
at base, 0.3 mm long. Style apex very mirrowly triangular, c. 1 mm long, prominently papillose.
Achene compressed barrel-shaped, c. 1.4 mm long, myxogenic; carpopodium absent; pericarp
crustaceous, brown, minutely colliculatc, the colliculi interspersed with larger duplex, rounded, clear
papillae: testa free from pericarp, papery, very pale brown, vascular strand slender and extending
around apex of seed and half way down other side. Pappus absent.

Specimens examined. NORTHERN TERRITORY: 12 miles south east of Mt Brockman, TV. Byrnes
2705 (NT); 18 km north-north-east of Jabiru East, L.A. Craven 6347 (CANB); 26 km south-south-
east of Jabiru East, M. Lazarides 9138 (CANB).

Distribution. Only known from the Kakadu National Park in the Northern Territory.

Habitat. Open Eucalyptus or Acacia woodland on sandstone plateau.

Etymology. The species is named after the National Park in which it is found.

This species does not appear to be closely related to any other species in the genus. The only other
member that has acuminate style branches is A. acoma which differs markedly in having broad flat
leaves, a naked convex receptacle, and finely tailed anthers.

Thiseltonia

Thiseltonia gracillima (F.Muell. & Tate) Paul G. Wilson, comb. nov.

7/Hmca granV/TmaF. Muell. & Tate, Trans. & Proc. Roy. Soc. South Australia 16:367(1896);
F. Muell., Victorian Nat. 9:144(1893) nomen. - Calomeria gracillima (F. Muell. & Tate) Heine!

482

Nuytsia Vol. 8, No. 3 (1992)

Figure 1. Acomis kakadu. A & B - habit. C - cluster of capitula. D - receptacle. E - intennediate involucral bract. F - floret.
G - two corolla lobes showing nervation. H-anther. I-styleamis. J- achene. A-I from L. 5466, J from L. Craven 6347.

Paul G. Wilson, /icowzi' and Thisellonia

483

Adansonia sen 2, 7 : 1 38( 1967). Leciotype (here chosen): Elder Exploring Expedition, Victoria Desert
Camp44 [c.27°S, 127°E, Western Australia], 7 September 1 891, (MEL, iso: NSW 179949).

Thisellonia dyeri Hemsley, Hooker’s Icon. PI. 28: tab. 2781(1905). Typification: 'West Australia:
Dedari, twenty-four miles west of Coolgardie, at 1,400 feet above sea-level, G.H. Thiselton-Dyer'.
Type n.v.

Hemsley, op. c//., suggested that Thisellonia was related to Pilhocarpa, while recently Anderberg
(1990) has proposed a close affinity to Hyalosperma S teetz to which in general appearance Thisellonia
undoubtedly corresponds. However, as is mentioned above, the corolla, anthers, style appendages,
andinvolucral bractsofr/j/^elton/o all indicateaclose relationship ioAcomis,m affinity thatMueller
(1893) had previously indicated when he placed the taxon in Hiimea sect. Acomis.

Acknowledgements

I thank Clyde Dunlop for passing to me the interesting species of Acomis and for indicating its
possible relationship; I am also grateful for his critical comments on the manuscript. I thank the
Curator of the Australian Herbarium (CANB) for sending on loan material of A. kakadu. The
illustration was kindly prepared by Margaret Menadue.

References

Anderberg, A. (1989). Phylogeny and reclassification of the tribe Inuleae (Asteraceae). Canadian J. Bot. 67:2277-2296.
Anderberg, A. (1991). Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Botanica 104:1-195.
Bentham. G. (1867). "Flora Australiensis," vol. 3. (Lovell Reeve & Co., London.)

Bentham, G. (1873). In G. Bentham & J.D. Hooker, "Genera Plantarum" Vol. 2. (Lovell Reeve & Co., London.)

Candolle, A.P. de (1838), "Prodromus systematis naturalis regni vegetabilis." Vol. 6. (Treuttel & Wurtz, Paris.)

Farr, E.R., Leussink, J.A. & StaOeu, F.A. (1979). "Index nominum genericorum." Vol. 1. (Bohn, Scheltema & Holkema
Utrecht.)

Grieve, B.J.&Blackall,W.E. (1975). "HowtoknowWestem AustralianWildflowers."PartIV. (University of Western Australia
Press, Perth.)

Heine, R. (1967). Ca/ooieria in "Ave Caesar, botanici te salutant." Adansonia ser. 2, 7:137-139.

Hemsley, W.B. (1905). Thisellonia dyeri in Hooker's leones Plantamm 28: tab. 2781.

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1:325. (Department of Agriculture, Pretoria.)

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Edinburgh 32:303-387. '

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Mueller, F. (1860). Rmidosis acoma. Fragmenta Phytographiae Australiae 2:89.

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Willis, J.H. (1967). Systematic notes on the indigenous Australian flora. Muelleria 1:117-163(1967).

484

Nuytsia Vol. 8, No. 3 (1992)

CORRFXTION

Correction to 'Taxonomic review of the Grevillea drummondii Meissn. species group
(Proteaceae)' by G.J. Keighery, Nuytsia 8(2): 225-230 (1992)

In the above paper a new species Grevillea fuscolutea Keighery, sp. nov. was described. In the
heading on page 228 the spelling 'fusculotea' was printed, but this was a typographical error and should
have read ’fuscolutea'. The latter spelling was used elsewhere in the paper. Article 73.1 of the
'International Code of Botanical Nomenclature' allows for correction of typographic errors. The
spelling Grevillea fuscolutea is correct and the spelling 'fusculotea' rejected. - Editor.

Publication date of Nuytsia Volume 8 Number 2: 2 April 1992

0980 0892 650

Notes for Authors

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Herbarium. The Herbarium reserves the right to reject papers.

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(1) Taxon name, synonymy (if any) and type details (forpreviously published taxa).

(2) Latin(fornewtaxa-indented).

(3) TypMj.Tfornewtaxa-notindented).

(4) Englishdescription(indented).

(5) Other specimens examinedoi Selected specimens examined as appropn3le,pieleTsMy including
number of collections examined.

(6) Distribution.

(!) Habitat

(8) Floweringperiod.

C?) Fruiting period.

(10) Typification (discussion).

(1 1) AffinitiesorRelationships.

(12) Discussion or Comments oiNotes.

(13) Conservation status.

(14) Etymology.

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(1) Authorabbreviations— Anon.(1980).DraftindexofAuthorAbbreviationsCorapiledattheHerbarium,RoyalBotanicGarrdens,
Kew. (HMSO: London.)

( 2 ) Booktitlesinliteraturecitations — Stafleu,F.A.&Cowan,R.S. (1976-83). TaxonomicLiterature.Edn2.(I.A.P.T.:Utrecht)(but
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(Department of Agriculture: Perth.)

(3) JoumaltitlesinliteraturecitationsandiEferencelists — Lawrence, G.H.M. era/. (1968).B-P-H(Botanico-Periodicum-Huntianum).
— Gxeenloc .cit.

Figures. Numbers should follow a single sequence including maps.

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reported (e.g. arevision). Amethods section shouldinclude the method ofdrawingupthedescriptionsfrom specimens, extent of search
fortypes, and discussion of concepts forchoice of taxonomic categories. Adiscussion section should be considered, which would

include some or all of thefollowing:asummaty oflhe findings, emphasising the most significant; interpretationoftheresultsin the light

of other relevant work; statement of new problems which have arisen; advising of aspects which ate to be followed up; suggestion of
topics whichothersmightusefuUy pursue; prediction and speculation.

CONTENTS

Calothamnus superbus TJ. Hawkeswood & F.H. Mollemans
(Leptospermoideae; Myrtaceae), a new species from
south-west Western Australia.

By TJ Hawkeswood and F.H. Mollemans

31T

A new species of Guichenotia (Sterculiaceae) from
south western Australia.

By Gregory J. Keighery

319

Four new Drosera taxa from south western Australia.

By Allen Lowrie and Neville Merchant

323

New names and combinations for some Melaleuca
(Myrtaceae) species and subspecies from the south-west of
Western Australia considered rare or threatened.

By F.C. Quinn, KJ. Cowley, B.A. Barlow and K.R. Thiele

333

Three new species of Euphorbia L. subgenus Chamaesyce
Rafinesque (Euphorbiaceae) from central and
northern Australia.

By B.G. Thomson

351

The Lawrencella complex

(Asteraceae: Gnaphalieae; Angianthinae) of Australia.

By Paul G. Wilson

361

The classification of Australian species currently included in
Helipterum and related genera
(Asteraceae; Gnaphalieae): Part 1.

By Paul G. Wilson

379

The classification of Australian species currently included in
Helipterum (Asteraceae: Gnaphalieae): Part 2 Leucochrysum.
By Paul G. Wilson

439

The classification of some Australian species currently
included in Helipterum and Helichrysum (Asteraceae:
Gnaphalieae); Part 3 Anemocarpa and Argentipallium,
two new genera from Australia.

By Paul G. Wilson

447

The classification of the genus Waitzia Wendl.
(Asteraceae: Gnaphalieae).

By Paul G. Wilson

461

A new species of Acomis from the Northern Territory
and a new combination in the genus Thiseltonia
(Asteraceae: Gnaphalieae).

By Paul G. Wilson

479

Correction: Nuytsia 8(2):228

484

Publication date of Nuytsia Volume 8 Number 2

484

ISSN 0085-4417

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