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<strong>Nuytsia</strong><br />

WESTERN AUSTRALIA’S JOURNAL OF SYSTEMATIC BOTANY<br />

All enquiries and manuscripts should be directed to:<br />

The Managing Editor – NUYTSIA<br />

Western Australian Herbarium<br />

Dept of Environment and Conservation<br />

Locked Bag 104 Bentley Delivery Centre<br />

Western Australia 6983<br />

AUSTRALIA<br />

The classification of<br />

Australian species currently<br />

included in Helipterum<br />

and related genera<br />

(Asteraceae:<br />

Gnaphalieae): Part 1<br />

<strong>Nuytsia</strong> 8: 3 (<strong>1998</strong>)<br />

Telephone: +61 8 9334 0500<br />

Facsimile: +61 8 9334 0515<br />

Email: nuytsia@dec.wa.gov.au<br />

Web: science.dec.wa.gov.au/nuytsia<br />

All material in this journal is copyright and may not be reproduced except with the written permission of the publishers.<br />

© Copyright Department of Environment and Conservation<br />

ISSN 0085–4417<br />

Paul G. Wilson


Paul G. Wilson, Helipterum: Part 1<br />

The classification of Australian species currently included in Helipterum<br />

and related genera (Asteraceae: Gnaphalieae): Part 1<br />

Paul G. Wilson<br />

Western Australian Herbarium, Department of Conservation and Land Management,<br />

PO Box 104, Como, Western Australia 6152<br />

Abstract<br />

Wilson, Paul G. The classification of Australian species currently included in Helipterum and related<br />

genera (Asteraceae: Gnaphalieae): Part 1. <strong>Nuytsia</strong> 8(3): 379-438 (1992). The name Helipterum DC. is<br />

recognised as being illegitimate and not applicable to Australian taxa. The Australian species currently<br />

included in that genus and in the related genera Cephalipterum, Gilberta, Triptilodiscus and Pterochaeta<br />

are reclassified. One genus, Haptotrichion, is described as new. Under Rhodanthe 11 sections are<br />

recognised of which 1 is new and 10 are new combinations. Eight new species of Rhodanthe and one<br />

new species of Haptotrichion are described, these are R. ascendens, R. collina, R. cremea, R. gossypina,<br />

R. nullarborensis, R. psammophila, R. rufescens, R. sphaerocephala and H. colwillii. A number of new<br />

species combinations are made. Descriptions are given for each of the recognised genera and sections.<br />

Contents Page<br />

Introduction 380<br />

Rhodanthe 383<br />

Rhodanthe sect. Rhodanthe<br />

383<br />

Rhodanthe sect. Monencyanthes 384<br />

Rhodanthe sect. Leiochrysum 385<br />

Rhodanthe sect. Achyroclinoides 399<br />

Rhodanthe sect. Citrinae 406<br />

Rhodanthe sect. Synachryum 408<br />

Rhodanthe sect. Helipteridium 412<br />

Rhodanthe sect. Helichrysoides 412<br />

Rhodanthe sect. Actinaria 414<br />

Rhodanthe sect. Anisolepis 415<br />

Rhodanthe sect. Polyphyllum 416<br />

379


380 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Contents Page<br />

Cephalipterum 417<br />

Erymophyllum 418<br />

Hyalosperma 419<br />

Gilberta 419<br />

Triptilodiscus<br />

420<br />

Haegiela 421<br />

Pterochaeta 422<br />

Haptotrichion 422<br />

Acknowledgements 426<br />

References 426<br />

Index to Scientific Names 427<br />

Index to name changes 436<br />

Introduction<br />

As has been pointed out by many syantherologists (Hilliard & Burtt 1981, Anderberg 1989, 1991,<br />

Wilson 1989a), the genera Helipterum Lindley (1836), Helipterum DC. (1838) and Helichrysum Miller,<br />

in the strict sense, do not occur in Australia and, in any event, the first name is illegitimate and is to be<br />

typified by a South African species currently placed in Helichrysum (Wilson 1989a), while the second<br />

is illegitimate and is to be typified by a South African species of Syncarpha DC. (Nordenstam 1989).<br />

The difference between the genera Helipterum DC. and Helichrysum, as enunciated by A. de Candolle<br />

(1838), is that in Helipterum the pappus bristles are plumose whereas in Helichrysum they tend to be<br />

barbellate, or plumose only at the apex. A strict interpretation of this distinction can lead to a placement<br />

of related species into separate genera, although the distinction has meant that closely related species<br />

have usually been placed together in either Helipterum or in Helichrysum.<br />

In order to classify the species currently included in Helipterum and Helichrysum it has been<br />

necessary to study related genera of the Gnaphalieae (sensu Anderberg 1989) such as Leptorhynchus,<br />

Ixiolaena, Waitzia, Podolepis, Podotheca, Myriocephalus, as well as several monotypic genera.<br />

The revised classification has resulted in the recognition of additional genera and the rearrangement<br />

of the species currently in Helipterum and Helichrysum. Some of the resulting genera are being revised<br />

by other botanists whose work I do not wish to compromise by producing a comprehensive classification<br />

lacking in the information that they will be able to provide. Therefore in this paper I am formally treating<br />

a portion of the Helipterum complex while indicating the genera into which other species currently<br />

included in Helipterum are likely to be transferred.


Paul G. Wilson, Helipterum: Part 1<br />

The classification that follows for a portion of the Helipterum complex in Australia attempts to<br />

associate closely related species and to segregate as distinct genera those species or groups of species<br />

that exhibit marked morphological discontinuity. In Rhodanthe I have not been successful in interpreting<br />

the phylogenetic significance of the varied morphological forms that are adopted by homologous<br />

organs. In the achene, for example, the pericarp and testa can have many different textures and structures<br />

(Short et al. 1989), yet to generically separate taxa that have dissimilar achenes can lead to the separation<br />

of species which otherwise have markedly similar florets and similar bract morphologies. For this reason<br />

I have adopted a sectional rank for taxa that may later be shown to deserve generic rank and which were<br />

sometimes so accorded by early 19th century taxonomists.<br />

I have not attempted to place all the genera of the 'Angiantheae’ sensu Bentham (1867) in context<br />

with those of the Helipterum complex but it is obvious that a close relationship often exists. This type<br />

of synoptic work will have to be done by others; my principal concern is to provide a classification that<br />

can be used for a 'Flora of Australia’ treatment and, from the pressure of time alone, many interesting<br />

phylogenetic avenues must remain unexplored.<br />

This paper considers the Rhodanthe, Hyalosperma and Triptilodiscus groups of species, most of<br />

which have, in general, been placed in Helipterum. These groups can be informally categorized as<br />

follows.<br />

Rhodanthe group: Inner involucral bracts with broad flat claws; anthers with fine filamentous tails;<br />

achenes with normal (mostly not thickened) duplex hairs. Species in this group, with the exception of<br />

Rhodanthe citrina, that have been examined, do not form ectomycorrhiza (Warcup 1990).<br />

Hyalosperma group: As above, however, achenes lacking normal duplex hairs but with 2-celled bulbous<br />

papillae that are frequently myxogenic; pappus shed entire. Species in this group that have been<br />

examined form ectomycorrhiza (Warcup op.cit.)<br />

Triptilodiscus group: Species in this group have achenes with 2-celled tooth-like trichomes and oblong<br />

or boat-shaped claws to the involucral bracts. Other characters suggest that the included genera may<br />

not be closely related and that the group is polyphyletic; this therefore becomes a grouping of<br />

convenience. In only Triptilodiscus has the mycorrhizal association been investigated; it was found<br />

to form ectomycorrhiza (Warcup op.cit.).<br />

The genus Rhodanthe, which is by far the largest of the segregate genera, is circumscribed to contain<br />

the majority of the Australian species previously included in Helipterum. Some of the remaining species<br />

are listed under other genera in this paper. Those species excluded from the present treatment are as<br />

follows:<br />

Helipterum albicans (Cunn.) DC., Helipterum fitgibbonii F. Muell., H. molle (DC.) Paul G. Wilson, and<br />

H. stipitatum (F. Muell.) Benth. have been transferred to Leucochrysum (DC.) Paul G. Wilson, (see<br />

Wilson 1992a) a genus closely related to Waitzia.<br />

Helipterum niveum Steetz is closely related to Helichrysum obtusifolium Sonder; both species will be<br />

segregated, with others, in a genus to be described.<br />

Helipterum saxatile Paul G. Wilson is closely related to Helichrysum podolepidium F. Muell., these two<br />

species will, with others, be placed in a genus to be described.<br />

381


382 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Helipterum craspedioides is conspecific with Myriocephalus morrisonianus and is possibly best<br />

recognised as a member of the genus Polycalymma (but see Short et al. 1989 and Anderberg 1991).<br />

Helipterum adpressum W. Fitzg. (= Helichrysum puteale S. Moore) and Helipterum pterochaetum<br />

(F. Muell.) Benth. have been transferred to the genus Chrysocephalum Walp. as C. puteale<br />

(S. Moore) Paul G. Wilson and C. pterochaetum F. Muell. (see Anderberg 1991 and Wilson, Short &<br />

Orchard 1992).<br />

The species that are here placed in Rhodanthe form a polymorphic assemblage that is divided into<br />

eleven sections. Some of the sections that have obvious problems associated with them are briefly<br />

discussed below.<br />

The sect. Monencyanthes, in which the achenes are persistent and the capitula caducous, contains<br />

three species. These species probably attained their specialised condition independently for each has<br />

features that suggest derivation from a different ancestor possibly of the Achyroclinoides type.<br />

The species in sect. Leiochrysum are diverse in vegetative and floral morphology. The organs appear<br />

to vary in morphology independently from each other and I have been unable to correlate characters<br />

in a manner that would achieve a more natural classification.<br />

The sect. Helichrysoides appears to be a monophyletic group which, at one extreme has affinity to<br />

the genus Podotheca and, at the other, to species in sect. Leiochrysum. Its correct taxonomic status<br />

is uncertain (see also Anderberg 1991).<br />

Each of the sections Anisolepis (1 sp.), Actinaria (2 spp.), and Helipteridium (1 sp.), has very<br />

distinctive involucral bracts but each shows little difference in floral morphology to certain species in<br />

sect. Achyroclinoides; I have therefore included these sections in Rhodanthe rather than give them<br />

generic status.<br />

The sect. Citrinae, with the single species R. citrina, has been difficult to place. Due to the beaked<br />

apex to its achene this species has been previously included in Waitzia. It differs radically from Waitzia<br />

in that the involucral bracts have flat claws with an undivided stereome (fide Anderberg 1991), the style<br />

apex is truncate, and the achenes pilose. It differs from other species of Rhodanthe in sometimes having<br />

the outer florets female (not hermaphrodite) and in being an ectomycorrhizal species (Warcup 1990). Both<br />

these characters suggest a relationship with species such as Chrysocephalum apiculatum (Labill.) Steetz<br />

and Leucochrysum albicans (Cunn.) Paul G. Wilson, but the morphology of the achenes and corollas<br />

of the three species is quite different. It possibly deserves recognition as a distinct genus.<br />

Rhodanthe polyphyllum, which is placed in the monotypic section Polyphyllum, is a further anomaly<br />

since it has stout, branched anther tails and clustered, filiform leaves. It appears to have no close relative<br />

in the Helipterum - Helichrysum complex. Nevertheless I have included it in Rhodanthe with which it<br />

appears to have greatest affinity. Again, it may deserve recognition as a distinct genus.


Paul G. Wilson, Helipterum: Part 1<br />

Catalogue of taxa<br />

In this Catalogue I have listed the accepted names and their synonyms in the Rhodanthe,<br />

Hyalosperma and Triptilodiscus groups. For new taxa and for all genera and sections I have provided<br />

descriptions. Where it has been necessary I have lectotypified names.<br />

Rhodanthe Lindley, Bot. Reg. t.1703(1834).<br />

The Rhodanthe group<br />

Rhodanthe Lindley<br />

Helichrysum sect. Rhodanthe (Lindley) Baillon, Hist. Pl. 8:314(1886). Type: R. manglesii Lindley<br />

Roccardia Necker ex Voss, Vilm. Blumengärtn. ed.3, 1:530(1895) nom. illeg. non Necker ex Raf.(1838).<br />

Type: R. manglesii (Lindley) Voss [=Rhodanthe manglesii Lindley].<br />

Annual or short-lived perennial herbs, woolly, puberulous, pilose, glandular, or glabrous. Leaves<br />

mostly alternate, simple, entire, rarely terete. Capitula homogamous or heterogamous, solitary or<br />

clustered, radiant or not; subtending leaves, when present, grading into the outer bracts. Involucral<br />

bracts multi-seriate, glumaceous or scarious, rarely with a herbaceous tip; claws oblong with narrow or<br />

broad undivided stereome (Anderberg 1991). Receptacle glabrous, or pilose around alveolae,<br />

predominantly ebracteate. Florets bisexual or the innermost male, otherwise homomorphic. Corolla<br />

tubular at base, usually turbinate above, glabrous or sparsely puberulous; lobes 5, equal or not, inner<br />

epidermis with cells not oblong in transverse rows. Anther tails filamentous and weak. Style apex<br />

truncate to ellipsoid or deltoid. Achene with normal duplex hairs; carpopodium annular or absent.<br />

Pappus bristles barbellate to plumose.<br />

1. Rhodanthe Lindley sect. Rhodanthe<br />

Annual erect herb, glabrous. Leaves cauline, sessile, suborbicular, base decurrent on each side of<br />

stem. Capitula heterogamous, solitary on slender branches. Involucre turbinate; bracts multi-seriate,<br />

papery, petaloid; claw with scarious margin and narrow-oblong thin stereome. Receptacle rounded,<br />

smooth, glabrous. Florets numerous, mainly bisexual but the innermost male, 5-merous. Corolla<br />

actinomorphic, glabrous; upper portion cup-shaped; lobes not papillose; veins extending into base of<br />

lobes. Stamens: anther appendage cordate, acute, the cells narrow-oblong towards apex, equilateral near<br />

base; anther tails filamentous, weak; collar narrow-oblong. Style apex narrow-deltoid, vascular trace not<br />

extending to tip. Achene obovoid, dorsiventrally compressed, densely silky to base; carpopodium<br />

absent; pericarp thin, translucent, in medial position in relation to the cotyledons; testa thin, translucent,<br />

± fused to pericarp, the vascular strand extending almost completely around seed; crystals absent.<br />

Pappus persistent; bristles plumose, the cilia all acute.<br />

A monotypic section that is most closely related to species in sect. Leiochrysum. It is distinctive<br />

in being glabrous, in having suborbicular sessile decurrent leaves, and possessing an anther appendage<br />

and a style apex that are unique in the genus.<br />

383


384 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Rhodanthe manglesii Lindley, Bot. Reg. 20: t.1703(1834). - Helipterum manglesii (Lindley) Benth., Fl.<br />

Austral. 3:640(1867). - Helichrysum manglesii (Lindley) Baillon, Hist. Pl. 8:314(1886). - Argyrocome<br />

manglesii (Lindley) Kuntze, Revis. Gen. Pl. 1:309(1891). - Roccardia manglesii (Lindley) Voss, Vilm.<br />

Blumengärtn. 3rd edn, 1:531(1895). Type: Swan River Colony, leg. J. Drummond, n.v.<br />

Rhodanthe atrosanguinea J. Drumm. ex hort., Gard. Chron. no. 27:622(1861). Type: Limestone Hills,<br />

Champion Bay, W.A., J. Drummond, n.v.<br />

Rhodanthe manglesii var. sanguinea Hook., Bot. Mag. 87:t.5283(1861). - Roccardia manglesii f.<br />

sanguinea (Hook.) Voss, Vilm. Blumengärtn. 3rd edn, 1:531(1895). Type: Western Australia,<br />

J. Drummond, n.v.<br />

Roccardia manglesii f. ligulosa Voss, Vilm. Blumengärtn. 3rd edn, 1:531(1895). Type: not indicated.<br />

Helipterum cryptanthum O. Sarg., J. Bot. 61:285(1923). Type: Mt Brown, York, Western Australia,<br />

O. Sargent 1404 (holo: BM; ?iso: MEL 109635 Sargent 1404a).<br />

2. Rhodanthe sect. Monencyanthes (A. Gray) Paul G. Wilson, comb. nov.<br />

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

Helipterum sect. Monencyanthes (A. Gray) Benth., Fl. Austral. 3:648(1867). Type: Monencyanthes<br />

gnaphalioides (Hook.) A. Gray<br />

Annual herbs, variably woolly. Leaves cauline, linear to narrow-oblong, entire. Capitula heterogamous,<br />

sessile or subsessile in terminal glomerules, caducous. Involucre ellipsoidal, narrow-cylindrical, or cupshaped,<br />

not radiant; bracts scarious, woolly ciliate, the inner with linear hard claw. Receptacle<br />

insignificant or rounded, with or without receptacular bracts. Florets 1-c. 12, bisexual or male, 5-merous.<br />

Corolla actinomorphic, cylindrical or narrow-turbinate, glabrous. Stamens: anther appendage broad<br />

ovate, obtuse, the cells narrow-oblong, thin-walled; anther tails filamentous, weak. Style apex truncate,<br />

papillose. Achene ellipsoidal, c. 2.5 mm long, persistent on receptacle, sparsely short pilose; carpopodium<br />

absent; pericarp membranous, hyaline, and with crystals (M. maryonii) or outer layer with thickened<br />

brown radial walls and crystals absent, nerves medial or medial-oblique in relation to cotyledons (Short<br />

et al. 1989); testa membranous and fused to pericarp or somewhat leathery and free (M. maryonii),<br />

crystals present. Pappus persistent, the bristles plumose weak with a few long cilia.<br />

Notes. This taxon was treated in a very broad sense as a section of Helipterum by Bentham (1867)<br />

so as to include some species now placed in Chrysocephalum, Rhodanthe sect. Achyroclinoides, and<br />

Hyalosperma. I have restricted it to those species with a woolly indumentum, and caducous capitula<br />

with persistent shortly hirsute achenes that have a persistent pappus. The included species, though<br />

superficially similar, probably arose independently from different species with caducous achenes<br />

since in the microscopic structure of the corolla each displays a significantly different morphology.<br />

It is interesting that in the achene of R. maryonii the pericarp bears crystals (presumably of calcium<br />

oxalate) whereas in all other species of Rhodanthe that have crystals in the achene they are found only


Paul G. Wilson, Helipterum: Part 1<br />

in the testa. Short et al. (1989) record that of the members of the Inuleae that they examined, crystals<br />

in the pericarp were only found in Craspedia aff. pleiocephala.<br />

Rhodanthe moschata (Cunn. ex DC.) Paul G. Wilson, comb. nov.<br />

Gnaphalium moschatum Cunn. ex DC., Prod. 6:236(1838). - Helipterum moschatum (DC.) Benth., Fl.<br />

Austral. 3:648(1867). Argyrocome moschata (DC.) Kuntze, Revis. Gen. Pl. 1:309 (1891). Type: Molle’s<br />

Plains, Lachlan River, Interior of New South Wales, 1817, A. Cunningham (holo: G-DC photo seen).<br />

Calocephalus gnaphalioides Hook. in T. Mitch., J. Exped. Int. Trop. Austral. 378(1849). - Monencyanthes<br />

gnaphalioides (Hook.) A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 6:230(1852). Type: T.L. Mitchell, n.v.<br />

This species is often confused with R. tietkensii which may be distinguished by its more numerous<br />

florets (c. 10 to a head), and absence of prominent glandular hairs on the corolla lobes. In R. moschata<br />

there are c. 3 florets to a capitulum (1 or 2 bisexual) and prominent glands on the lobes.<br />

A collection from the Simpson Desert, Northern Territory (G. Chippendale, NT 6577, herb. AD)<br />

appears to be a hybrid between R. tietkensii and R. moschata. The herb. NT duplicate of this collection<br />

is of true R. moschata. Rhodanthe tietkensii has been found in the same area.<br />

Material collected in northern Eyre Peninsula, South Australia, differs from the common variant in<br />

having milky white (not scarious) involucral bracts.<br />

Rhodanthe uniflora (J. Black) Paul G. Wilson, comb. nov.<br />

Helipterum uniflorum J. Black, Trans. & Proc. Roy. Soc. South Australia 41:651 t.43(1917). Lectotype<br />

(here chosen): Koppermanna, 7 Oct. 1916, S.A. White (AD 98625090 p.p.), isolecto: MEL 110472, NSW<br />

181468.<br />

Rhodanthe maryonii (S. Moore) Paul G. Wilson, comb. nov.<br />

Helipterum maryonii S. Moore, J. Linn. Soc. Bot. 45:182(1920). Type: Mulline, J.E.C. Maryon (holo: BM;<br />

iso: MEL 110713, NSW 181424).<br />

3. Rhodanthe sect. Leiochrysum (DC.) Paul G. Wilson, comb. nov.<br />

Helipterum sect. Leiochrysum DC., Prod. 6:216(1838). Lectotype (here designated): Helipterum<br />

polygalifolium DC. [=Rhodanthe polygalifolia (DC.) Paul G. Wilson].<br />

Helipterum sect. Sericophorum DC., Prod. 6:216(1838). Lectotype (here designated): Helipterum<br />

humboldtianum (Gaudich.) DC. [=Rhodanthe humboldtiana (Gaudich.) Paul G. Wilson].<br />

Pteropogon DC., Prod. 6:245(1838). - Pteropogon sect. Facelioides A. Gray, Hooker’s J. Bot. Kew Gard.<br />

Misc. 4:267(1852) nom. illeg. - Helipterum sect. Pteropogon (DC.) Benth., Fl. Austral. 3:639(1867). Type:<br />

P. pygmaeus DC.<br />

385


386 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

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

Acroclinium A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:270(1852). - Helichrysum sect. Acroclinium<br />

(A.Gray) Baill., Hist. Pl. 8:174, 314(1882); F. Muell., Fragm. 3:32(1862) nomen. Lectotype (here designated):<br />

A. multicaule A. Gray [=Rhodanthe chlorocephala (Turcz.) Paul G. Wilson].<br />

Griffithia J. Black, Trans. & Proc. Roy. Soc. South Australia 37:122(1913). Type: G. helipteroides J. Black<br />

[=Rhodanthe oppositifolia (S. Moore) Paul G. Wilson].<br />

Annual herbs (rarely short-lived perennial), glabrous or variously pubescent. Leaves linear to<br />

oblong, entire. Capitula heterogamous or homogamous, solitary or clustered. Involucre globular to<br />

cylindrical or turbinate, radiant or not; bracts multiseriate, scarious, papery, or thinly cartilaginous;<br />

stereome oblong to linear, thin; claw of inner involucral bracts flat, scarious. Receptacle rounded to<br />

conical, smooth or pitted, glabrous or puberulous. Florets 5 to numerous, bisexual, or the innermost male,<br />

5-merous. Corolla actinomorphic to zygomorphic, almost glabrous to variously pubescent, cylindrical<br />

or expanded above, cells of inner epidermis of lobes equilateral to broad-oblong, smooth or papillose.<br />

Stamens: anther appendage narrow-deltoid to ovate or oblong, cells narrow-oblong; anther tails<br />

filamentous, weak; collar oblong or narrow-oblong. Style apex truncate to broadly deltoid, prominently<br />

papillose. Achene obovoid, compressed obovoid, or ellipsoid, 1.5-3.5 mm long, densely silky or<br />

moderately short pilose; carpopodium a short glabrous slightly tumid hollow base or absent; pericarp<br />

crustaceous, thinly coriaceous, or papery, vascular strands in medial or oblique position; testa<br />

membranous to thinly coriaceous, free from or fused to pericarp, the vascular strand 1/2 to 3/4 encircling<br />

seed; crystals in testa present or absent. Pappus persistent or caducous; bristles linear-lanceolate,<br />

plumose, the terminal cilia frequently rounded or clavate.<br />

Note 1. Candolle (1838) placed Helipterum polygalifolium and H. diffusum in his section Leiochrysum,<br />

I have selected the first of these as the lectotype since it better agrees with his sectional description.<br />

Note 2. Gray (1852) included two species in Pteropogon sect. Facelioides of which one was P. pygmaeus,<br />

the type of Pteropogon. His sectional name is therefore superfluous.<br />

Rhodanthe anthemoides (Sprengel) Paul G. Wilson, comb. nov.<br />

Helichrysum anthemoides Sieber ex Sprengel, Syst. Veg. 3:484(1826). - Helipterum anthemoides<br />

(Sprengel) DC., Prod. 6:216(1838). - Argyrocome anthemoides (Sprengel) Kuntze, Revis. Gen. Pl.<br />

1:309(1891). - Roccardia anthemoides (Sprengel) Voss, Vilm. Blumengärtn. 3rd edn, 1:531(1895)<br />

'anthemodes’. Type: 'Nov Holl.’ probably F.W. Sieber 344 (iso: G-DC photo seen, K, MEL 109048, 604823).<br />

Helipterum punctatum DC., Prod. 6:216(1838). Type: Van-Diemen Land, R.Gunn 239 (holo: G-DC photo<br />

seen; ?iso: MEL 109067 p.p.).<br />

Rhodanthe chlorocephala (Turcz.) Paul G. Wilson, comb. nov.<br />

Schoenia chlorocephala Turcz., Bull. Soc. Imp. Naturalistes Moscou 24/1:193(March 1851). -<br />

Helipterum chlorocephalum (Turcz.) Benth., Fl. Austral. 3:641(1867). - Argyrocome chlorocephala


Paul G. Wilson, Helipterum: Part 1<br />

(Turcz.) Kuntze, Revis. Gen. Pl. 1:309(1891). - Roccardia chlorocephala (Turcz.) Voss, Vilm. Blumengärtn.<br />

3rd edn, 1:530(1896). Type: Western Australia, J. Drummond 4th coll. n. 199 (iso: MEL 109146, 109147,<br />

109148).<br />

Rhodanthe chlorocephala subsp. chlorocephala<br />

[Acroclinium multicaule A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:271(1852) as to var. ß not as to<br />

lectotype. Syntype: Swan river, J. Drummond (K).]<br />

I am applying the epithet subsp. chlorocephala in a narrow sense so as to restrict it to the form<br />

represented by the type collection of this taxon. This type was collected from an unknown locality in<br />

the south-west of Western Australia; it was included in Drummond’s fourth collection which was<br />

despatched to England in July 1847. According to Erickson (1969) the fourth collection consisted of<br />

plants gathered from King George Sound, Stirling Range, Porongorups, Mt Manypeaks, Cape Riche,<br />

West Mt Barren, and from north and east of Moore River. Of these various localities Rhodanthe<br />

chlorocephala is only known from the Moore River area but the only other specimens I have been able<br />

to match with the type are those that represent the original material of Acroclinium multicaule A.Gray<br />

var. ß which is an apparently undated and unnumbered collection of Drummond’s in herb. K. It is possible<br />

that the specimens that form the bases of R. chlorocephala and A. multicaule var. ß are from the same<br />

Drummond collection.<br />

Rhodanthe chlorocephala subsp. rosea (Hook.) Paul G. Wilson, comb. et subsp. nov.<br />

Acroclinium roseum Hook., Bot.Mag. t.4801(1854). - Helichrysum grayi F. Muell., Fragm. 5:200(1866).<br />

- Helipterum roseum (Hook.) Benth., Fl. Austral. 3:640(1867). - Argyrocome rosea (Hook.) Kuntze, Revis.<br />

Gen. Pl. 1:309(1891). - Roccardia rosea (Hook.) Voss, Vilm. Blumengärtn. 3rd edn, 1:531(1896). Type<br />

citation: 'A native of the interior of South-west Australia, between the Moore and Murchison rivers from<br />

whence the seeds were sent in 1853 by Mr. James Drummond, together with dried specimens (No. 157<br />

of his Herbarium).’ Lectotype (here chosen): J. Drummond 6th coll. no. 157 (lecto: K; isolecto: NSW<br />

180743). Syntype: Hort. Kew. 1854 from Drummond S.W. Australia (K).<br />

Acroclinium multicaule A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:271(1852). - Helichrysum<br />

drummondii F. Muell., Fragm. 5:200(1866). Lectotype (here chosen): Swan River, J. Drummond (lecto:<br />

?K n.v.; ?iso: P, 'J. Drummond 349’, annotated by A. Gray in 1851).<br />

Acroclinium roseum var. grandiflorum Nicholson, Ill. Dict. Gard. 1:18(1884). - Helipterum roseum var.<br />

grandiflorum (Nicholson) Chitt., Dict. Gardening 2nd edn, 2:978(1951). Type: not indicated.<br />

Acroclinium roseum var. album hort. ex Nicholson, Ill. Dict. Gard. 1:18(1884). - Helipterum album<br />

(Nicholson) L. Bailey, Cycl. Amer. Hort. 2:726(1900). - Helipterum roseum f. album (Nicholson)<br />

Moldenke, Phytologia 2:312(1947). Type: not indicated.<br />

Helipterum roseum var. nigropapposum Ostenf., Biol. Meddel. Kongel. Danske Vidensk. Selsk.<br />

3/2:140(1921). Type citation: 'Perth, King’s Park, in open sandy places (No.874; 10.Sept.1914).’ (iso:<br />

MEL 110494).<br />

387


388 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

[Helipterum troedelii var. patens Ewart, J. White & B. Rees, Proc. Roy. Soc. Victoria 22 n.s.:15(1909).<br />

- Helipterum roseum var. patens (Ewart et al.) J. Black, Trans. & Proc. Roy. Soc. South Australia<br />

45:21(1921) as to syntype 'Fraser Range, 1891, R. Helms’, not as to lectotype. Syntype: Fraser Range,<br />

4 Oct.1891, R. Helms (MEL 110705, AD 96343013, NSW 180742).]<br />

The involucral bracts in this subspecies can be pink or white.<br />

Note. Asa Gray, under Acroclinium multicaule, described a variety a in which the ray bracts were white,<br />

and a variety ß in which the ray bracts were 'subaeruginosis’. The lectotype cited above is of the var.<br />

a which corresponds to R. chlorocephala subsp. rosea, var. ß is referred to subsp. chlorocephala.<br />

Rhodanthe chlorocephala subsp. splendida (Hemsley) Paul G. Wilson, subsp. et comb. nov.<br />

Helipterum splendidum Hemsley, Bot. Mag. t.7983(1904). - Helipterum roseum var. album Ewart et al.,<br />

Proc. Roy. Soc. Victoria n.s. 23:59(1910) nom.illeg. Type citation: 'Kew is indebted to Mr. G.F. Berthoud,<br />

of Waroona, near Drakesbrook, W. Australia, for dried specimens and seeds of this handsome plant,<br />

which he labelled:- "Native of North-western Districts." There are also herbarium specimens collected<br />

in open plains at Menzies, about a hundred miles north of Coolgardie, in about 29° 30’S. lat. and 121° E.<br />

long., by Cecil Andrews.’ Lectotype (here chosen): North-western Districts of Western Australia,<br />

26.x.1903, G.F. Berthoud (lecto: K; isolecto: MEL 110704).<br />

Rhodanthe chlorocephala is extremely variable. At one extreme is found in the Shark Bay area of<br />

Western Australia a large flowered variant with long (to 30 mm) cream-coloured limbs to the inner<br />

involucral bracts. At the other extreme is found a variant in the Nullarbor area of South Australia and<br />

Western Australia with small heads and short (c. 5 mm) white limbs. Between these extremes is a<br />

continuity of forms. In addition to the differences in size of the heads and bracts, and the presence of<br />

a black spot at the base of the limb in the large cream bracted variant, there are found colour forms that<br />

range from white to cream and various shades of pink. Hooker noted that in cultivation Acroclinium<br />

roseum had much larger heads than the wild plant from which the seed was obtained and this difference<br />

is apparent when comparing the Drummond herbarium material with Hooker’s illustration.<br />

The variant described by Ostenfeld as Helipterum roseum var. nigropapposum was collected in<br />

Perth; it is almost identical to Drummond’s collection no. 157 (from between the Moore and Murchison<br />

Rivers) on which the name Acroclinium roseum was based.<br />

The name A. multicaule A. Gray was based on two variants, as is noted above, of which one<br />

corresponds to subsp. chlorocephala and the other to subsp. rosea.<br />

The name A. roseum var. grandiflorum Nicholson was based on a large-headed rose-coloured variant<br />

while var. album Nicholson was based on a white variant, both variants are presumably forms of subsp.<br />

roseum.<br />

Ewart et al. (1909) based the name H. troedelii var. patens on two collections of which one, the<br />

lectotype, is Rhodanthe troedelii s.str. while the other (from the Fraser Range in Western Australia)<br />

is of the small 'Nullarbor’ variant of subsp. rosea. It was the latter specimen that J.M. Black first saw


Paul G. Wilson, Helipterum: Part 1<br />

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

Rhodanthe collina Paul G. Wilson, sp. nov. (Figure 1)<br />

Annua erecta ad 20 cm alta. Caules graciles, atro-rubri, sparse puberuli et glanduloso punctati. Folia<br />

caulina alterna, aliquantum coriacea linearia vel anguste oblonga, integra, 8-15 mm longa, obtusa, margine<br />

saepe recurva, supra glabra, infra glanduloso punctata. Capitula heterogamia, solitaria, terminalia.<br />

Involucrum hemisphericum, c. 5 mm altum, radians. Bracteae multi-seriatae, glumaceae glabrae; bracteae<br />

intimae: ungue late elliptico c. 5 mm longo, stereomate lineari, duro, lamina oblongo-elliptica c. 6 mm longa,<br />

alba. Receptaculum conicum, glabrum. Flosculi numerosi, bisexuales vel masculi. Corolla actinomorpha,<br />

anguste cylindracea, c. 3.5 mm longa; lobi 5, triangulares, c. 0.4 mm longi; intra dense papillosi.<br />

Antherarum loculi c. 1 mm longi; appendix oblongo-ellipticus, acutus c. 0.4 mm longus, margine minute<br />

erosa, cellulis anguste oblongis, tenui-parietibus; caudi tenues, debiles. Styli rami apicem versus<br />

aliquantum latiores, apice rotundato longe papilloso, nervo gracili in medio rami terminanti. Achenium<br />

obovoideum c. 2 mm longum, atro-rubiginosum, dense albo-hirsutum, pilis crasso-parietibus; carpophorum<br />

annulare; pericarpium crustaceum; testa membranacea, atro-violaceum, ad pericarpio ± adnatum.<br />

Pappus in statu integro tarde deciduus; setae lineari-acuminatae, breviter plumosae, ciliis terminalibus<br />

clavatis.<br />

Typus: Paynes Find - Thundelarra road near Monger Lake, 29° 03' S, 117° 17' E, low hill of quartzite<br />

boulders, 27 Sept. 1986, Paul G. Wilson 12300 (holo: PERTH; iso: AD, BRI, CANB, K, MEL, NSW).<br />

Annual erect herb to 20 cm high branching at and above base. Major axes slender, dull reddish brown<br />

with a few small septate hairs and sessile globular glands. Leaves cauline, alternate, somewhat leathery,<br />

linear to narrow-oblong or spatulate, entire, 8 x 0.5 - 15 x 2 mm, obtuse, margin often recurved (at least<br />

when dry), glabrous above, sparsely covered below with sessile globular glands and with a few shorter<br />

septate hairs on midrib. Capitula heterogamous, solitary on slender leafless terminal peduncles.<br />

Involucre hemispherical, c. 5 mm high, radiant; bracts glumaceous, glabrous; outer and intermediate<br />

bracts elliptic, to 6 mm long, pale brown, with a short flat stereome; inner bracts: claw broad-elliptic,<br />

hyaline, c. 5 mm long with a hard linear stereome c. 2/3 its length; lamina oblong-elliptic, obtuse, c. 6 mm<br />

long, 2.5 mm wide, white. Receptacle conical, glabrous. Florets numerous, the outer bisexual and the<br />

inner male. Corolla narrow-tubular, somewhat constricted below middle, c. 3.5 mm high, yellow, sparsely<br />

puberulous towards base, wall thickened in lower half, vascular strands extending to base of lobes; lobes<br />

5, short erect, triangular, c. 0.4 mm long, all densely papillose within. Stamens: anther loculi c. 1 mm long;<br />

anther appendage oblong-elliptic, acute, c. 0.4 mm long, minutely erose on margin, cells narrow-oblong,<br />

walls unthickened except for a marginal row of small cells with thickened walls; anther tails very thin and<br />

delicate. Style branches broadened towards tip and with a rounded long-papillose apex; vascular strand<br />

very slender and terminating about half way along style branch. Achene obovoid, c. 2 mm long, very<br />

dark reddish brown, densely white-hirsute with thich-walled hairs, myxogenic; carpopodium annular;<br />

pericarp crustaceous with lignified reddish brown radial walls; testa very thin and weak, dark mauve, with<br />

scattered flat crystals, ± fused to pericarp. Pappus tardily deciduous as a whole; bristles linearacuminate,<br />

± equal to corolla, united in a short ring at base, shortly plumose, the terminal cilia clustered<br />

and clavate.<br />

Distribution. Near Yalgoo, central western Western Australia, in the far western area of the Austin<br />

Botanical District.<br />

389


390 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Figure 1. Rhodanthe collina. A - Habit. B - Undersurface of leaf. C - Multicellular hispidulous and globular glandular<br />

trichomes from undersurface of leaf. D - Capitulum. E - Inner, intermediate, and outer involucral bracts. F - Receptacle.<br />

G - Floret. H - Corolla. I - Inner surface of apex of corolla. J - Apex of pappus bristle. K - Anther with enlargement of<br />

margin of appendix (tails drawn recurved for space exigencies). L - Apex of style with two papillae enlarged. M - Achene<br />

with apex of duplex hair enlarged. N - Achene showing position of seed and of its vascular strand. O - Enlargement of<br />

epidermis of pericarp. P - Enlargement of epidermis of testa showing crystals. From B.H. Smith 460.


Paul G. Wilson, Helipterum: Part 1<br />

Specimens examined. WESTERN AUSTRALIA: 112 mile peg on the Yalgoo Road, A.M. Ashby 2976<br />

(AD, PERTH); Mt Gibson, B.H. Smith 460 (MEL, PERTH); Mingenew Hill, 29 Aug.1957, R.F. Watson<br />

(PERTH).<br />

Habitat. On rocky hills in exposed situations.<br />

Notes. This species would appear to have no close relatives although in habit and in indumentum it is<br />

similar to Rhodanthe diffusa. The unique anther appendages, style apices, and achenes combine to make<br />

it a very distinct species.<br />

Rhodanthe corymbiflora (Schldl.) Paul G. Wilson, comb. nov.<br />

Helipterum corymbiflorum Schldl., Linnaea 21:448(1848); Sonder, Linnaea 25:519(1853). - Argyrocome<br />

corymbiflora (Schldl.) Kuntze, Revis. Gen. Pl. 1:309(1891). - Roccardia corymbiflora (Schldl.) Voss, Vilm.<br />

Blumengärtn. 3rd edn, 1:532(1895). Type citation: 'Dr Behr’ [South Australia] (?holo: MEL 604825).<br />

Note. Sonder, l.c., cited the Behr collection as coming from 'Fiedler’s section, Novemb.’ This locality<br />

is in the Hundred of Moorooroo, County Light, near the present town of Nuriootpa (South Australia).<br />

The putative type sheet in herb. MEL has on it three specimens, two of which are somewhat etiolated<br />

whole plants while one is a single stem off a mature plant. A note on the sheet by Sonder states 'culmi<br />

e seminibus plantae behrianae!’ It would appear likely that the two whole plants represent those raised<br />

by Sonder while the single stem is the type specimen collected by Behr since it corresponds to the<br />

following comment made by Schlechtendal, l.c., 'Unicum specimen vidi, ramum 5 poll. longum’.<br />

Rhodanthe cremea Paul G. Wilson, sp. nov. (Figure 2)<br />

Annua erecta, c. 25 cm alta, sparse lanosa, pilis glandulosis absentibus. Axes majores graciles,<br />

solitarii vel numerosi, haud ramosi, apicem versus lanosi aliter glabri. Folia caulina alterna, integra, linearia<br />

vel anguste obovata, acuta vel obtusa, ad 30 mm longa, apicem versus caulorum absentia. Capitula<br />

solitaria ad caules terminalia posita. Bracteae exteriores scariosae vel chartaceae, latissime ovatae vel<br />

latissime ellipticae, nitidae, ad 7 mm longae, pallido brunneae; bracteae intermediae late ellipticae, ad 15<br />

mm longae, ad apicem malvinae; bracteae intimae: unguis late obovatus, stereomate oblongo, vix<br />

incrassato, nervo singulari ad apicem extenso; lamina petaloidea, anguste elliptica, ad 20 mm longa,<br />

pallido cremea vel ad basim atropurpurea. Receptaculum discoideum, margine areolae pilosa aliter<br />

glabrum. Flosculi numerosi, bisexuales vel flosculis intimis masculinis. Corolla: tubus anguste<br />

cylindraceus, c. 2 mm longus; faux valde zygomorpha anguste turbinata, c. 1 mm longa; lobi 5, c. 1.5 mm<br />

longi, lobis tribus abaxialis recurvis, cremeis, profunde fissa, intra laevibus, lobis duobus adaxiali erectis,<br />

ad apicem connatis, rubiginosis, intra papillosis, nervis corollae ad apicem lobulorum extensis. Antherarum<br />

appendix sterilis late elliptica, obtusa, 0.2 mm longa; cellulae oblongae, parietibus tenuibus, cellulis<br />

parietibus incrassatis immixtis; cellulae marginales manifeste discretae. Antherarum caudae filamentosae,<br />

debiles. Styli apex truncatus, nervo prominenti ad apicem extenso. Achenium compresse turbinatum,<br />

c. 2.5 mm longum, facie abaxiali sericeum, adaxiali hirsutum; carpophorum absens; pericarpium chartaceum;<br />

testa membranacea ad pericarpium adnata, nervo in positione medio cingenti. Pappi setae corollam leviter<br />

excedentes; in parte inferiore persistentes, anguste ellipticae; in parte superiore caducae, filamentosae,<br />

apice versus plumosae, ad extremum penicillatae ciliis clavatis ornatis, aliter denticulatae.<br />

391


392 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Typus: Western Australia, 6.5 km east of the turnoff to Useless Loop; abundant on flats, dominating a<br />

stony clay depression with Acacia tetragonophylla and Acacia sp., 16.viii.1986, R.C. Chinnock 6760<br />

(holo: AD; iso: K, MEL, PERTH).<br />

Erect annual to 25 cm high, branching at base; major axes slender, solitary or several, simple, woolly<br />

near apex otherwise glabrous, glandular hairs absent. Leaves cauline, alternate, entire, linear to narrowobovate,<br />

acute to obtuse, to 30 mm long, absent towards apex of stem. Capitula solitary, terminal to stems.<br />

Involucre hemispherical, c. 6 mm high. Bracts multiseriate, scarious or papery, glabrous; outer bracts<br />

scarious, very broadly ovate or broadly elliptic, to 7 mm long, pale brown; intermediate bracts broadelliptic,<br />

to 15 mm long, pale brown with mauve apex; inner bracts: claw broad-obovate, stereome broadoblong,<br />

scarcely thickened,with a single nerve extending to apex, lamina petaloid, narrow-elliptic, to 20<br />

mm long, pale cream, often with a dark purple band at base. Receptacle disc-shaped, c. 5 mm broad, pilose<br />

on margin of areolae otherwise glabrous. Florets numerous, bisexual or the inner ones male. Corolla<br />

almost glabrous: tube cylindrical, c. 2 mm long; throat narrow-turbinate, strongly zygomorphic, c. 1 mm<br />

long; lobes 5, c. 1.5 mm long, the three abaxial lobes deeply divided, recurved, cream coloured, smooth<br />

within, the 2 adaxial lobes fused to near apex, dark reddish brown, papillose within; vascular strands<br />

extending to tip of lobes. Anther appendage broad-elliptic, obtuse, 0.2 mm long; cells oblong, thickwalled<br />

cells interspersed among thin-walled cells; marginal cell row clearly demarcated; anther tails<br />

filamentous and weak. Style tip truncate, the vascular strand stout and extending to apex. Achene<br />

compressed turbinate, c. 2.5 mm long, silky on abaxial face and hirsute on adaxial face; carpopodium<br />

absent; pericarp papery; testa membranous, fused to pericarp, the vascular strand encircling seed in<br />

medial position. Pappus bristles slightly exceeding corolla; lower half narrow elliptic, barbellate and<br />

persistent; upper half filamentous and deciduous, towards apex plumose with the terminal cilia penicillate<br />

and clavate, towards base barbellate.<br />

Distribution. Western Australia, from the south end of Hamelin Pool, Shark Bay, eastwards to the North<br />

West Coastal Highway.<br />

Specimens seen. WESTERN AUSTRALIA: 493 mile peg North West Highway, T.E.H. Aplin 3282<br />

(PERTH); 5 miles west of Hamelin Station, J.W. Green 1444 (PERTH); 22 km west of Overlander<br />

Roadhouse on Denham road, N.S. Lander 1322 (PERTH); 28 km NNW of Overlander Roadhouse, North<br />

West Coastal Highway, E.N.S. Jackson 3121 (AD pro parte); 3 km from Overlander Roadhouse, P.S.<br />

Short 422 (AD pro parte); 20 km north of Billabong Roadhouse, P.S. Short 2453 (PERTH).<br />

Habitat. Usually found in red sand over loam.<br />

This species appears to be related to Rhodanthe chlorocephala. It superficially very closely<br />

resembles R. chlorocephala subsp. splendida which grows intermixed with R. cremea near Shark Bay<br />

and some herbarium collections consist of material of both species. Rhodanthe chlorocephala may be<br />

distinguished by the spherical glandular hairs on leaves and stems, the absence of woolly hairs, the<br />

regular corollas, and the evenly silky achenes.<br />

Those variants of Cephalipterum drummondii and of R. chlorocephala subsp. splendida that have<br />

large white involucral bracts often resemble R. cremea in having a dark purple band at the base of these<br />

bracts. Since the three species grow together in the Hamelin Pool region it is likely that they attract the<br />

same pollen vectors.<br />

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


Paul G. Wilson, Helipterum: Part 1<br />

Figure 2. Rhodanthe cremea. A - Habit. B - Capitulum. C - Corolla opened out. D - Bisexual floret. E - Corolla. F - Male<br />

floret. G - Receptacle. H - Pappus bristles. I - Style branches. J - Anther. K - Leaf. L - Leaf hairs. M - Outer and inner<br />

involucral bracts. N - Achene. O - T.S. achene. From J.W. Green 1444.<br />

393


394 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Rhodanthe diffusa (Cunn. ex DC.) Paul G. Wilson, comb. nov.<br />

Helipterum diffusum A.Cunn. ex DC., Prod. 6:216(1838). Type: Forest grounds at the base of Peels range,<br />

Interior of New South Wales, June 1817, A. Cunningham (holo: G-DC photo seen).<br />

Rhodanthe diffusa (DC.) Paul G. Wilson subsp. diffusa<br />

This taxon represents the yellow-rayed variant.<br />

Rhodanthe diffusa subsp. leucactina (F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum polygalifolium var. leucactinum F. Muell., Fragm. 10:107(1877). Type citation: 'Darling’s<br />

River (Andrae), Warrego (Bailey), Curruwillughi (Dalton), Ballandool (Looker).’ Lectotype (here<br />

chosen): Warrego district, F.M. Bailey (MEL 110257).<br />

The taxon represents the white-rayed variant.<br />

Note. The only substantial difference between the two subspecies is in the colour of the radiant<br />

involucral bracts, however the stems of northern variant of subsp. leucactina are sometimes branched<br />

above and have a minutely glandular hispidulous indumentum beneath the capitula rather than cottony.<br />

The typical subspecies is found in south-eastern New South Wales between Hillston and Narrandera<br />

while the subsp. leucactina is found further north from Nyngan north to south-eastern Queensland.<br />

Both subspecies have been confused with other species: the yellow subspecies was synonymized by<br />

Bentham (1867) (and subsequently confused in some herbaria) with Helipterum polygalifolium, while<br />

the white has been confused with H. anthemoides. Rhodanthe polygalifolia may be readily distinguished<br />

by being glabrous apart from the minute globular glands on the adaxial surface of the leaves (in R. diffusa<br />

the leaves bear glands and multiseptate hairs on the stems and on both surfaces of the leaves), while<br />

R. anthemoides may be distinguished by being a perennial, by the generally thicker leaves and embedded<br />

globular glands, and by the reddish brown linear midrib on the claws of the involucral bracts.<br />

Rhodanthe fuscescens (Turcz.) Paul G. Wilson, comb. nov.<br />

Helipterum fuscescens Turcz., Bull. Soc. Imp. Naturalistes Moscou 24/2:80(1851). - Podotheca fuscescens<br />

(Turcz.) Benth., Fl.Austral. 3:602(1867). - Podosperma fuscescens (Turcz.) F. Muell., Fragm. 12:22(1882).<br />

- Argyrocome fuscescens (Turcz.) Kuntze, Revis. Gen. Pl. 1:309(1891). Type: Western Australia,<br />

J. Drummond 5th coll. (suppl.) n.64 (holo: KW photo seen; iso: K, MEL 110215, PERTH).<br />

Acroclinium phyllocephalum A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:271(1852). Type citation:<br />

'South-west Australia, Drummond (received in 1850)’ n.v.<br />

This species is very similar to Rhodanthe oppositifolia from which it differs in having leaves that<br />

subtend the capitula and in having very short and brown involucral bracts. Rhodanthe fuscescens is<br />

known only from the Drummond types and from E. Pritzel 695. The Drummond sheets on which the names<br />

H. fuscescens and A. phyllocephalum were based are probably duplicates of the same collection (Short<br />

1983); both sheets lack locality data. The collection Pritzel 695, dated 1901, is almost identical to the<br />

Drummond; it came from near Cranbrook in south-west Western Australia which is about 400 km from<br />

the nearest locality for R. oppositifolia.


Paul G. Wilson, Helipterum: Part 1<br />

Rhodanthe gossypina Paul G. Wilson, sp. nov.<br />

Herba annua erecta gossypina. Folia caulina, alterna, linearia vel anguste oblonga, acuminata, 2-4<br />

cm longa. Capitula homogama in corymbis terminalibus ordinata. Involucrum turbinatum, c. 4 mm altum,<br />

radians; bracteae c. 3-seriata; bracteae intermediae ellipticae c. 5 mm longae, hyalinae, lanato ciliatae, ad<br />

apice brunneolae, stereomate anguste oblongo, viridi, in parte superiore gossypino; bracteae intimae:<br />

unguis late obovatus c. 4 mm longus, hyalinis, gossypino ciliatus, stereomate anguste oblongo,<br />

gossypino, in parte superiore pilis resinosis rubiginosis ornatum; lamina patula, ovata, c. 4 mm longa,<br />

alba. Receptaculum conicum, c. 0.7 mm diam. Flosculi c. 15, bisexuales. Corolla c. 3 mm longa, 5-lobata;<br />

lobi inequales, oblongo-triangulares, intra laeves, lobo abaxiali c. 1.5 mm longo, ceteris c. 1 mm longis;<br />

nervi corollae in tubo terminantes. Antherae: appendix sterilis elliptica c. 0.4 mm longa, parietibus<br />

cellularum nec incrassatis; caudis filamentis, debiles. Styli apex ± truncatus. Achenium ellipsoideum,<br />

1.5-3 mm longum, subtiliter pilosum; carpophorum annulare; pericarpium chartaceum, translucens; testa<br />

aliquantum coriacea, brunnea, ad pericarpium adnata, nervo c. 2/3 cingenti. Pappus persistens; setae<br />

lineari-acuminatae corollam aequantes, omnino pilosae.<br />

Typus: Queensland: Mitchell district, 36.9 km SE of Winton on Longreach road, 19 Sept. 1984,<br />

R.J. Chinnock 6106 (holo: AD; iso: PERTH).<br />

Annual erect cottony herb; stems slender, single or branching at base, to 20 cm high. Leaves cauline,<br />

alternate, linear to narrow-oblong, acuminate, 2-4 cm long, 1-5 mm wide. Capitula homogamous, in<br />

terminal corymbs. Involucre turbinate, c. 4 m high, radiant; bracts c. 3-seriate; intermediate bracts elliptic,<br />

c. 5 mm long, hyaline, woolly ciliate, brown-tinged towards apex, with narrow-oblong green stereome<br />

cottony in upper half, c. 2 mm long; innermost bracts: claw broad-obovate, c. 3.7 mm long, 2 mm wide,<br />

hyaline, cottony ciliate, stereome narrow-oblong extending to near apex, cottony and with dark reddish<br />

brown resinous hairs in upper half; lamina spreading, ovate, c. 4 mm long, 2 mm wide, white. Receptacle<br />

conical, c. 0.7 mm diameter Florets c. 15, all bisexual. Corolla c. 3 mm long, narrow-cylindrical below,<br />

somewhat broader above, sparsely puberulous in lower half, yellow, vascular strands terminating well<br />

below lobes; lobes oblong-triangular, 0.5-1 mm long, not papillose within, glabrous outside, the abaxial<br />

lobe more deeply divided (up to 1.5 mm long). Anthers: appendage elliptic, c. 0.4 mm long, smooth, cells<br />

not thickened, very pale brown or the marginal ones clear; tails filamentous, weak. Style apex ± truncate<br />

to slightly rounded, shortly papillose. Achene ellipsoidal, 1.5-3 mm long, pilose with fine very pale brown<br />

hairs; carpopodium a short glabrous ring; pericarp translucent, papery, very pale brown; testa somewhat<br />

leathery, brown, fused to pericarp when mature; vascular strand of testa extending c. 2/3 around seed<br />

in medial position. Pappus bristles linear-acuminate, ± equal to corolla, united in a ring at base, persistent,<br />

pilose throughout, the terminal cilia somewhat thicker and rounded at apex.<br />

Distribution. Southeastern Northern Territory, southwestern Queensland, extreme northeastern South<br />

Australia and northwestern New South Wales.<br />

Specimens examined (selection only). QUEENSLAND: 100 km WNW of Charleville, C. Sandercoe 369<br />

(BRI); Near Windorah, S.T. Blake 12079 (BRI).<br />

NEW SOUTH WALES: 17.5 km from Louth on Telpa road, C.W.E. Moore 7794 (CANB).<br />

SOUTH AUSTRALIA: Kudriemitchie Waterhole, L.D.Williams 8156 (AD).<br />

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

Habitat. In open situations on heavy soil that is frequently subject to flooding.<br />

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396 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Flowering time. June to September.<br />

This species is similar to Rhodanthe corymbiflora and R. microglossa with which two species it has<br />

been confused in herbaria. It differs from the former in having a smaller involucre with the bract-lamina<br />

about half as long, and in the possession of dark resinous hairs on the stereome of the inner bracts. From<br />

the latter it differs in having a more open inflorescence, dark resinous hairs on the inner bracts, much<br />

longer radiating laminae, and in having a smooth throat to the corolla.<br />

Rhodanthe gossypina was recognised as a distinct taxon by C.T. White (herb. BRI) who evidently<br />

intended to describe it as a variety of Helipterum corymbiflorum but never did so (R.W. Johnson,<br />

pers.comm.).<br />

The solitary collection from New South Wales, cited above, comes from a site geographically isolated<br />

from the other locations. It is morphologically distinct for the stereomes of the inner involucral bracts<br />

do not have glandular hairs and the lobes of the corolla are equal in length. For these reasons it may<br />

be found to represent a distinct species.<br />

The specific epithet refers to the cottony indumentum that covers the plant.<br />

Rhodanthe humboldtiana (Gaudich.) Paul G. Wilson comb. nov.<br />

Helichrysum humboldtianum Gaudich. in Freyc., Voy. Uranie 465(1830) t.88(1829). - Helipterum<br />

humboldtianum (Gaudich.) DC., Prod. 6:216(1838). - Schoenia humboldtiana (Gaudich.) Walpers,<br />

Repert. Bot. Syst. 6:244(1846). - Pteropogon humboldtianus (Gaudich.) F. Muell., Linnaea 25:415(1853).<br />

- Argyrocome humboldtiana (Gaudich.) Kuntze, Revis. Gen. Pl. 1:309(1891). - Roccardia humboldtiana<br />

(Gaudich.) Voss, Vilm. Blumengärtn. 3rd edn, 1:532(1896). Type citation: 'In Novae-Hollandiae ora<br />

occidentale (baie des Chiens-Marins)’, 1830, Gaudichaud. Lectotype (here chosen): P photo seen;<br />

isolecto: G-DC photo seen.<br />

Helipterum sandfordii Hook., Bot. Mag. t.5350(1862). Type: J. Drummond 160, between Moore and<br />

Murchison rivers; 1860, Burgess; cult. Thompson from seeds from Major Sandford, W.A. Isosyntypes:<br />

J. Drummond 160 (MEL 109493, NSW).<br />

Helipterum largiflorens F. Muell., Fragm. 3:135(1863). Type citation: 'Ad sinum Champion bay et flumen<br />

Murchison. Walcott et Oldfield.’ Lectotype (here chosen): Champion Bay, A. Oldfield (MEL 109501).<br />

Helipterum clementii Domin, Biblioth. Bot. 89:667(1929). Type citation: 'Nordwest-Australien: zwischen<br />

Ashburton- und De Gray River, E. Clement’. Type n.v.<br />

Rhodanthe microglossa (Maiden & E. Betche) Paul G. Wilson, comb. nov.<br />

Helipterum microglossum Maiden & E. Betche, Proc. Linn. Soc. New South Wales 22:151(1897). Type:<br />

Tarella, New South Wales, Aug. 1887, W. Bäuerlen 131 (holo: NSW 181467).<br />

Helipterum corymbiflorum var. microglossa F. Muell. ex Benth., Fl. Austral. 3:647(1867). -<br />

H. microglossa (Benth.) Tate, Trans & Proc. Roy. Soc. South Australia 22:121(1898). Lectotype<br />

(here chosen): Goyinga mountains, 6 & 7 Nov. 1860, Victorian Expedition (MEL 110255).


Paul G. Wilson, Helipterum: Part 1<br />

Note. It was unclear from the paper by Maiden and Betche whether they intended to describe a new<br />

species based on the Bäuerlen collection or whether they intended to make a new combination based<br />

on Helipterum corymbiflorum var. microglossa; I have assumed the former intention, partly because<br />

the epithet was changed from a substantive to an adjective and partly because the format used was the<br />

same as that for other new species described in the paper. Ralph Tate (1898 loc.cit.) claimed to have<br />

published the species combination in the Trans. & Proc. Roy. Soc. South Australia 6:104(1883), however,<br />

in that paper the name Helipterum microglossa was not validly published since it appeared without<br />

citation of either author or basionym, and without a description.<br />

Rhodanthe oppositifolia (S. Moore) Paul G. Wilson, comb. nov.<br />

Helipterum oppositifolium S. Moore, J. Bot. 35165(1897). Type: Near Coolgardie, Sept.1895, S.Moore<br />

(holo: BM; iso: MEL 110309).<br />

Griffithia helipteroides J. Black, Trans. & Proc. Roy. Soc. South Australia 37:122(1913). Type citation:<br />

'Gawler Ranges, Eyre Peninsula (S.A.White, September, 1912)’ (holo: AD; iso: K).<br />

subsp. oppositifolia<br />

subsp. ornata Paul G. Wilson, subsp. nov.<br />

Folia lineari-acuminata, c. 30 x 2 mm. Involucrum hemisphericum, c. 6 mm altum; lamina bracteae<br />

interiorum elliptica, 10 mm longa, flava.<br />

Typus: Western Australia: 3 km NW of Tamala Homestead, 26° 40' S, 113° 42' E, at foot of calcrete ridge<br />

on edge of samphire flat, powdery white clay loam on sheet calcrete, 25 August 1991, S.D. Hopper 8112<br />

(holo: PERTH; iso: CANB, K, MEL, S).<br />

Leaves linear-acuminate, c. 30 x 2 mm. Involucre hemispherical, c. 6 mm high; inner bracts with a<br />

prominent elliptic yellow lamina c. 10 mm long.<br />

Distribution. Only known from the southern margin of Freycinet Estuary in Tamala station, Western<br />

Australia. Found growing on calcrete rises on edge of saline flats.<br />

Additional specimens seen. WESTERN AUSTRALIA: 13.5 km W of Tamala homestead road on Useless<br />

Loop road, S.D. Hopper 8113 (PERTH); Tamala Station, 20 July 1988, Ph. Morat 8212 (P).<br />

This subspecies differs from the typical in having larger capitula and much larger lamina to the inner<br />

involucral bracts which in the typical subspecies are only 2-3 mm long. Subsp. oppositifolia is also found<br />

in the Shark Bay region but evidently not in association with subsp. ornata.<br />

Rhodanthe polygalifolia (A.Cunn. ex DC.) Paul G. Wilson, comb. nov.<br />

Helipterum polygalifolium A.Cunn. ex DC., Prod. 6:216(1838). - Argyrocome polygalifolia (DC.) Kuntze,<br />

Revis. Gen. Pl. 1:309(1891). Type: Molle’s Plains, Lachlan river, New South Wales, 1817, A. Cunningham<br />

(holo: G-DC photo seen).<br />

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398 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Rhodanthe propinqua (W. Fitzg.) Paul G. Wilson, comb. nov.<br />

Helipterum propinquum W. Fitzg., J. W. Austral. Nat. Hist. Soc. No.1:24(May 1904). Lectotype (here<br />

chosen): Nannine, Western Australia, Sept. 1903, W.V. Fitzgerald (NSW 181422; isolecto: NSW 181423,<br />

PERTH).<br />

Rhodanthe pygmaea (DC.) Paul G. Wilson, comb. nov.<br />

Pteropogon pygmaeus DC., Prod. 6:245(1838). - Pteropogon australis Nees, Linnaea 16:223(1842) nom.<br />

illeg., based on above. - Helipterum pygmaeum (DC.) F. Muell., Rep. Babbage’s Exped. 14(1859); Benth.,<br />

Fl. Austral. 3:647(1867). Type: Molle’s plains, Lachlan river, Interior west from Port Jackson, Long. 145°E,<br />

'a remarkable pigmy plant’, July 1817, A. Cunningham 101 (holo: G-DC photo seen).<br />

Pteropogon drummondii A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:267(1852). - Helipterum pygmaeum<br />

var. occidentale Benth., Fl. Austral. 3:647(1867). - Helipterum pygmaeum var. drummondii (A. Gray)<br />

Ostenfeld, Dansk Bot. Ark. 3:142(1921) nom. illeg. - Helipterum drummondii (A. Gray) Ostenfeld, l.c.<br />

Type: Western Australia, 1849, J. Drummond (holo: K).<br />

Pteropogon intermedius F.Muell., Linnaea 25:411(1853). Type citation: 'In collibus graminosis montem<br />

Rufus versus, ante Torrens tenus, ubertim rarus.’ Lectotype (here chosen): Ad rivum Broughton, Oct.<br />

1851, F. Mueller (MEL 604821).<br />

Rhodanthe rubella (A. Gray) Paul G. Wilson, comb. nov.<br />

Acroclinium rubellum A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:271(1852). - Helipterum rubellum<br />

(A. Gray) Benth., Fl. Austral. 3:641(1867). - Argyrocome rubella (A. Gray) Kuntze, Revis. Gen. Pl.<br />

1:309(1891). - Roccardia rubella (A. Gray) Voss, Vilm. Blumengärtn. 3rd edn, 1:530(1896). Type citation:<br />

'Between Swan River and King George Sound, Drummond.’ Lectotype (here chosen): J.Drummond 347<br />

(K, photo seen).<br />

Rhodanthe rufescens Paul G. Wilson, sp. nov.<br />

Herba annua decumbens, sparse et minute glanduloso-puberula; axes majores numerosi ad 10 cm<br />

longi. Folia caulina alterna sessilia, anguste oblonga, ad 10 mm longa, integra, obtusa. Capitula<br />

homogama, ad ramulos terminalia posita, nec radiantia. Involucrum breviter cylindraceum, 4-5 mm<br />

altum. Bracteae c. 3-seriatae, scariosae, supra stereoma sparsissime et minute glanduloso puberulae<br />

aliter glabrae; bracteae exteriores ovatae, c. 2 mm longae, stramineae; bracteae interiores obovatae,<br />

ad 5 mm longae, ad apicem rotundatae, versus basim stramineae, versus apicem violaceae vel margine<br />

apicali hyalino. Receptaculum rotundatum c. 0.7 mm diam., glabrum. Flosculi c. 7, bisexuales,<br />

actinomorphi. Corolla c. 3.5 mm longa, infra medium cylindracea, supra medium anguste cupulata;<br />

lobi 5, triangulares, c. 1 mm longi, intra ad basim papillosi; nervi ad basim loborum extensi. Antherae<br />

loculis c. 1 mm longis; appendix c. 0.3 mm longus ad apicem rotundatus, cellulis oblongis nec<br />

incrassatis; caudae filamentae, debiles. Styli apex truncatus, nervo ad basim apicis extenso.<br />

Achenium (in statu immaturo) sericeum, carpophoro absenti, pericarpio tenui, translucenti, testa<br />

tenuiter coriacea. Pappi setae lineares acuminatae, c. 2 mm longae, omnino plumosae, ciliis omnino<br />

acuminatis.


Paul G. Wilson, Helipterum: Part 1<br />

Typus: Queensland, Gregory South District, Noccundra, about 130 km west-north-west of Thargomindah,<br />

'herbage’, 11 Aug. 1987, P.Warhurst s.n. (holo: BRI 410267).<br />

Annual (? decumbent) herb with many somewhat wiry stems to 10 cm long arising from base.<br />

Branches and leaves sparsely covered with minute sessile or shortly stipitate globular glands, otherwise<br />

glabrous. Leaves cauline, alternate, sessile, narrow-oblong, to 10 mm long, 1-2 mm wide, entire, obtuse.<br />

Capitula homogamous, terminal to slender lateral and main branches, sometimes subtended by a<br />

foliaceous bract. Involucre shortly cylindrical, 4-5 mm high, not radiant; bracts scarious, glabrous apart<br />

from a few minute glandular hairs on stereome, c. 3-seriate; outer bracts ovate c. 2 mm long, strawcoloured,<br />

glabrous; inner bracts obovate, to 5 mm long, rounded at apex, straw-coloured towards base,<br />

violet brown towards apex or with the tip hyaline, stereome oblong, 2/3 length of bract, flat, pale green.<br />

Receptacle rounded, c. 0.7 mm diameter, glabrous. Florets c. 7, all apparently bisexual. Corolla tube c.<br />

3.5 mm high, cylindrical below, narrowly cup-shaped above, very sparsely puberulous, yellow; lobes<br />

5, ovate, triangular, c. 1 mm long, papillose within in a broad ring around base of lobes; vascular strands<br />

extending to base of lobes. Stamens: anther appendage c. 0.3 mm long rounded at apex, cells oblong,<br />

not thickened; anther tails weak and filamentous. Style tip truncate, papillose; vascular strand prominent<br />

and extending to base of tip. Achene (not seen mature) silky villous all over; carpopodium absent;<br />

pericarp thin, translucent; testa thinly coriaceous, with crystals. Pappus bristles linear-acuminate, c. 2<br />

mm long at anthesis, plumose to tip, terminal cilia not thickened.<br />

Distribution. Only known from the type locality in south-west Queensland.<br />

The specific epithet refers to the reddish colour of the terminal portion of the inner involucral bracts.<br />

Rhodanthe stricta (Lindley) Paul G. Wilson, comb. nov.<br />

Xyridanthe stricta Lindley, Sketch Veg. Swan Riv. Col. 23(1839). - Helipterum strictum (Lindley) Benth.,<br />

Fl. Austral. 3:646(1867), Type: Swan River, 1839, J. Drummond (holo: CGE photo seen).<br />

Pteropogon platyphyllus F. Muell., Linnaea 25:413(1853). Type citation: 'Ad margines rupestres<br />

rivulorum ad Cudnaka rariusve ibidem in planitiebus sterilibus.’ Lectotype (here chosen): 'Ad marginis<br />

rupestris rivi Cudnaka’, Oct.1851, F. Mueller (lecto: MEL 110685; isolecto: MEL 604829).<br />

Helipterum strictum var. stenocephala S. Moore, J. Linn. Soc. 34:200(1899). Type: Near Coolgardie,<br />

August 1895, S. Moore (holo: BM photo seen).<br />

4. Rhodanthe sect. Achyroclinoides (A. Gray) Paul G. Wilson, comb. nov.<br />

Pteropogon sect. Achyroclinoides A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:268(1852). Lectotype<br />

(here designated): Pteropogon corymbosus A. Gray.<br />

[Cassinia sect. Cassiniola F. Muell., Fragm. 3:139(1863) nomen sub Cassinia cuprea F. Muell.<br />

(= Pteropogon polycephalus A. Gray)].<br />

Plant variously pubescent to sub-glabrous, without sessile glands on leaves. Style apex truncate.<br />

Achene small, 1-1.5 mm long, moderately to sparsely short hirsute; carpopodium annular, glabrous;<br />

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400 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

pericarp crustaceous, thin, brown, the radial cell walls thickened; testa membranous, free or fused to<br />

pericarp. Pappus caducous.<br />

Note. Asa Gray (1852) included Pteropogon polycephalus A. Gray, P. corymbosus A. Gray and P. laevis<br />

A. Gray in his section Achyroclinoides. These three species are all recognised here as being members<br />

of this section.<br />

Rhodanthe ascendens Paul G. Wilson, sp. nov.<br />

Herba annua ascendens ad 10 cm alta, modice lanosa. Folia caulina, alterna, anguste obovata, plana,<br />

10-15 mm longa, c. 2 mm lata, sessilia. Inflorescentia corymbiformia, compacta. Capitula homogama,<br />

breviter pedunculata. Involucrum cupulatum c. 5 mm altum, 2.5 mm latum, nec radians; bracteae<br />

glumaceae, 3-4-seriatae, nitidae, bracteis intermediis interioribusque ellipticis, 4-5 mm longis, ad apicem<br />

rotundatis, ad basim sparse ciliatis aliter glabris. Receptaculum convexum, glabrum, c. 1 mm latum,<br />

bracteis receptaculi absentibus. Flosculi c. 20, bisexuales, actinomorphi. Corolla c. 3 mm longa, glabra,<br />

nervis ad apicem tubi extensis; lobi deltoidei, c. 0.5 mm longi, lobis tribus parte interiore ad basim<br />

papillosis, ceteris glabris. Antherae c. 1 mm longae; apex anguste ovatus obtusus, c. 0.3 mm longus,<br />

cellulis anguste oblongis; caudae filamentae, debiles. Styli apex truncatus, nervo prominenti ad apicem<br />

extenso. Achenium, in statu immaturo, anguste cylindraceum, 1 mm longum, breviter hirtellum;<br />

carpophorum annulare, glabrum; pericarpium tenue, translucens; testa membranacea. Pappus corollam<br />

aequans, in statu integro caducae; setae c. 15, filiformae, ad basem breviter connatae, barbellatae.<br />

Typus: Western Australia, Gascoyne Junction; florets yellow; roadside verge, clay soil; 20 Aug. 1986,<br />

P.S. Short 2531 (holo: PERTH; iso: MEL n.v.).<br />

Annual ascendent herb to 10 cm high, branching at base, moderately and loosely woolly. Leaves<br />

sessile, alternate, cauline, narrow obovate, obtuse, flat, 10 - 15 mm long, c. 2 mm wide. Inflorescence<br />

corymbiform, compact. Capitula homogamous, very shortly pedunculate. Involucre cup-shaped, c. 5<br />

mm high, not radiant; bracts glumaceous, 3-4-seriate, glossy, intermediate and inner bracts elliptic, to<br />

4.5 mm long, rounded at apex, sparsely ciliate at base otherwise glabrous. Receptacle convex, glabrous,<br />

without receptacular bracts, c. 1 mm diameter. Florets c. 20, bisexual, actinomorphic. Corolla c. 3 mm long,<br />

yellow, narrow-tubular in lower half, narrow-turbinate above, glabrous, vascular strands extending to<br />

apex of tube; lobes broad-triangular, c. 0.5 mm long, three of the lobes papillose within at base the others<br />

glabrous; cells of throat strongly undulate. Stamens: anther loculi c. 1 mm long; tip oblong-ovate, obtuse,<br />

c. 0.3 mm long, the cells narrow-oblong; tails filamentous, weak. Style arms truncate, papillose, the<br />

vascular strand prominent and extending to base of tip. Achene (in immature state) narrow-cylindrical,<br />

1.0 mm long, shortly hirtellous with duplex hairs c. 0.1 mm long, bifid at tip, myxogenic; carpopodium a<br />

narrow glabrous ring; pericarp thin and translucent, weak; testa membranous, free from pericarp, vascular<br />

strand extending to apex of seed; endosperm persistent and forming a sheath around the embryo. Pappus<br />

bristles c. 15, ± equal to corolla, very shortly united in a ring at base, barbellate throughout, the terminal<br />

teeth rounded, caducous as a whole.<br />

Distribution. Found only in the Carnarvon Botanical District (Beard 1980) near Gascoyne Junction,<br />

Western Australia.<br />

Additional specimen examined. WESTERN AUSTRALIA: Middalya Station, 1989, D.C. Norbury 1<br />

(PERTH).


Paul G. Wilson, Helipterum: Part 1<br />

Habitat. The type was found in clay soil on a roadside verge.<br />

Rhodanthe ascendens is very similar to R. nullarborensis from which it differs most obviously in<br />

having broader leaves and larger capitula. In the details of the florets it differs in having barbellate (not<br />

shortly plumose) pappus bristles and in having three (not two) of the corolla lobes papillose at their base,<br />

but this latter character may be a variable one. The achene of R. nullarborensis has a thin crustaceous<br />

pericarp with lignified radial walls while in its immature state the pericarp of R. ascendens shows no sign<br />

of such thickening but this may develop later.<br />

Rhodanthe condensata (F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum condensatum F. Muell., Fragm. 3:136(1863). - Argyrocome condensata (F. Muell.) Kuntze,<br />

Revis. Gen. Pl. 1:309(1891). Type: Murchison River, Western Australia, A. Oldfield (holo: MEL 110770).<br />

Rhodanthe corymbosa (A. Gray) Paul G. Wilson, comb. nov.<br />

Pteropogon corymbosus A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:268(1852). - Helipterum corymbosum<br />

(A. Gray) Benth., Fl. Austral. 3:649(1867). - Argyrocome corymbosa (A. Gray) Kuntze, Revis. Gen. Pl.<br />

1:309(1891). Type citation: 'Swan river, Drummond. - Darling Range, South-west Australia, Collie.’<br />

Isosyntype: J. Drummond 364 (MEL 109605).<br />

Helipterum album Ewart, J. & Proc. Roy. Soc. New South Wales 42:189(1909). Type: Wooroloo, Western<br />

Australia, 1906, M.Koch 1553 (?holo: NSW 181466; iso: PERTH).<br />

Rhodanthe forrestii (F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum forrestii F. Muell., S. Sci. Rec. 2:273(1882). - Argyrocome forrestii (F. Muell.) Kuntze, Revis.<br />

Gen. Pl. 1:309(1891). Type citation: 'In the neighbourhood of the Gascoyne-River; J. Forrest.’ Lectotype<br />

(here chosen): Gascoyne river, 1882, J.Forrest (lecto: MEL 50292; isolecto: K, MEL 50290, 50291, PERTH).<br />

This species differs from R. polycephala in having the leaves narrowed at the base and not decurrent<br />

and in having the capitula turbinate and not narrow-cylindrical. Further collecting may show that<br />

R. polycephala is only a northern variant of R. forrestii.<br />

Rhodanthe haigii (F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum haigii F. Muell., Fragm. 10:107(1877). - Argyrocome haigii (F. Muell.) Kuntze, Revis. Gen.<br />

Pl. 1:309(1891). Type: Eucla, Richards (holo: MEL 110217; iso: PERTH).<br />

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

MEL 110310).<br />

Note. This species is similar to R. humboldtiana with which it was confused by Black (1957) and Haegi<br />

(1986). Apart from the presence of larger rays to the involucral bracts, R. humboldtiana may be readily<br />

distinguished by its silky pilose achenes for in R. haigii the achenes are shortly hirsute. Only R. haigii<br />

occurs in South Australia while both species are found in Western Australia.<br />

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402 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Rhodanthe laevis (A. Gray) Paul G. Wilson, comb. nov.<br />

Pteropogon laevis A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:269(1852). - Helipterum laeve<br />

(A. Gray) Benth., Fl. Austral. 3:649(1867). - Argyrocome laevis (A. Gray) Kuntze, Revis. Gen. Pl.<br />

1:309(1891). Type: Swan River, 1843, J. Drummond 366 (or 356) (holo:K).<br />

Rhodanthe nullarborensis Paul G. Wilson, sp. nov. (Figure 3)<br />

Herba annua ad 30 cm alta, modice lanosa, aromatica. Folia caulina, alterna, linearia, plana, 10-15 mm<br />

longa, 1-1.5 mm lata, sessilia. Inflorescentia corymbiformia, compacta. Capitula homogama, breviter<br />

pedunculata. Involucrum cylindraceum vel anguste turbinatum, 3-4 mm altum, nec radians; bracteae<br />

glumaceae, 3-4-seriatae, nitidae, bracteis intermediis interioribusque oblongis, 3-4 mm longis, ad apicem<br />

rotundatis, sparse ciliatis aliter glabris. Receptaculum convexum, glabrum, c. 0.4 mm latum, bracteis<br />

receptaculi absentibus. Flosculi c. 12, bisexuales, actinomorphi. Corolla c. 3 mm longa, fere glabra, nervis<br />

ad apicem tubi extensis; lobi deltoidei, c. 1 mm longi, lobis duobus parte interiore ad basim papillosis,<br />

ceteris glabris. Antherae c. 1 mm longae; apex anguste ovatus obtusus, c. 0.3 mm longus, cellulis anguste<br />

oblongis; caudae filamentae, debiles. Styli apex truncatus, nervo prominenti ad apicem extenso.<br />

Achenium anguste obovoideum, 1 mm longum, breviter hirtellum; carpophorum annulare, glabrum;<br />

pericarpium tenuiter crustaceum, atro rubiginosum, parietibus radialibus cellularum incrassatis; testa<br />

diaphana, ad pericarpum adnata. Pappus corollam aequans, in statu integro caducae; setae c. 12,<br />

filiformae, ad basem breviter connatae, breviter plumosae.<br />

Typus: Forrest, Western Australia, 'heavily scented, flower heads yellow’, 30 Aug. 1930, E.R.L. Johnson<br />

75 (holo: PERTH; iso: AD, K, MEL).<br />

Annual erect herb to 30 cm high, branching at base, moderately and loosely woolly, strongly scented.<br />

Leaves numerous, alternate, cauline, linear, flat, 10 - 15 mm long, 1 - 1.5 mm wide, sessile. Inflorescence<br />

corymbiform, compact. Capitula homogamous, very shortly pedunculate. Involucre cylindrical to<br />

narrow-turbinate, 3-4 mm high, not radiant; bracts glumaceous, 3-4-seriate, glossy, intermediate and<br />

inner bracts oblong, 3-4 mm long, rounded at apex, sparsely ciliate, otherwise glabrous. Receptacle<br />

convex, glabrous, without receptacular bracts, c. 0.4 mm diameter. Florets c. 12, bisexual, actinomorphic.<br />

Corolla c. 3 mm long, yellow, narrow-tubular in lower half, narrow-turbinate above, glabrous or rarely very<br />

sparsely glandular puberulous, vascular strands extending to apex of tube; lobes broad-triangular, c.<br />

1 mm long, two of the lobes papillose within at base, the others glabrous. Stamens: anther loculi c. 1 mm<br />

long; tip narrow-ovate, obtuse, c. 0.3 mm long, the cells narrow-oblong; tails filamentous, weak. Style<br />

arms truncate, papillose, the vascular strand prominent and extending to base of tip. Achene narrowobovoid,<br />

1.0 mm long, shortly hirtellous with duplex hairs c. 0.07 mm long, minutely bifid at tip, myxogenic;<br />

carpopodium a narrow glabrous ring; pericarp thinly crustaceous, dark reddish brown, radial cell walls<br />

thickened; testa extremely thin and fused to pericarp, the vascular strand extending to apex of seed;<br />

endosperm persistent and forming a sheath around the embryo. Pappus bristles c. 12, ± equal to<br />

corolla,very shortly united in a ring at base, shortly plumose throughout, caducous as a whole.<br />

Distribution. Found only in the Nullarbor Region of Western Australia.<br />

Specimens examined (selection only). WESTERN AUSTRALIA: Kanowna, T.E.H. Aplin 5745<br />

(PERTH); 30 km S of Rawlinna, R.J. Chinnock 11262 (AD); 70 mi NNW of Reid, A.S. George 8491


Paul G. Wilson, Helipterum: Part 1<br />

Figure 3. Rhodanthe nullarborensis. A - Habit. B - Terminal cluster of capitula. C - Leafy bract. D - Leaf. E - Outer,<br />

intermediate, and inner involucral bracts showing enlargement of glandular hair. F - Floret. G - Inner surface of apex of<br />

corolla indicating that only two lobes are papillose. H - Anther. I - Style apex. J - Pappus bristle. K - Achene. L - Duplex<br />

hair from achene. M - Surface of pericarp. From G.J. Keighery 7623.<br />

403


404 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

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

7623 (PERTH).<br />

Habitat. In shallow open depressions (dongas) that have a layer of heavy soil over limestone.<br />

Flowering time. July to October.<br />

Rhodanthe nullarborensis resembles H. tietkensii with which it has been confused in herbaria. The<br />

latter species may be recognised by its larger leaves, its larger and more hairy capitula, and by the<br />

presence of receptacular bracts.<br />

Johnson & Baird (1970) in referring to this species as Helipterum tietkensii state that it is heavily<br />

scented and that it forms dense colonies in the dongas.<br />

Rhodanthe polycephala (A. Gray) Paul G. Wilson, comb. nov.<br />

Pteropogon polycephalus A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:268(1852). - Helipterum<br />

polycephalum (A. Gray) Benth., Fl. Austral. 3:649(1867). - Argyrocome polycephala (A.Gray) Kuntze,<br />

Revis. Gen. Pl. 1:309(1891). Type citation:'Swan river, Drummond’, n.v.<br />

Cassinia cuprea F. Muell., Fragm. 3:139(1863). Lectotype (here chosen): Near the Murchison, A. Oldfield<br />

(MEL 110308).<br />

Rhodanthe psammophila Paul G. Wilson, sp. nov. (Figure 4)<br />

Herba erecta ad 40 cm alta. Folia caulina alternata, anguste oblonga vel linearia, ad 6 cm longa, sessilia,<br />

parce decurrentia, lanata et minute stipitato-glandulosa. Capitula heterogama, sessilia, glomerata.<br />

Involucrum anguste turbinatum, c. 4 mm altum; bracteae 4-6-seriatae, obovatae, 3-4 mm longae, apice<br />

albo, rotundato, margine eroso; unguis bractearum interiorum dense lanato- ciliatus. Flosculi semper<br />

quinque, uno fertili, quatuor masculinis. Corolla actinomorpha, sparse puberula; lobi intra papillosi.<br />

Achenium compresso-obovoideum, c. 2.5 mm longum, sparse et minute puberulum; carpophorum<br />

annulare; pericarpium tenuiter crustaceum; testa membranacea pericarpium adnata. Pappus achenii<br />

fertile tarde deciduus; setae c. 13, anguste lineares, acuminatae, c. 2.0 mm longae, sparse ciliatae pilis<br />

gracilibus acuminatis. Achenium bracteis interioribus consociatis deciduum.<br />

Typus: Western Australia, 20 km north of Carnarvon, red sand dune, 25 September 1987, Paul G. Wilson<br />

12604 (holo: PERTH; iso: AD, CANB, K, MEL, NSW, S).<br />

Erect shortly woolly annual branching at and above base, to 40 cm high. Leaves cauline, alternate,<br />

narrow-oblong to linear, acuminate, to 6 cm long, sessile, slightly decurrent along the margins, woolly<br />

all over and with minute shortly stipitate reddish brown globular glands. Capitula heterogamous,<br />

sessile in dense clusters, these shortly stipitate forming subglobular heads on slender peduncles 1-3<br />

cm long. Involucre narrow-turbinate, c. 4 mm high; bracts 6-seriate, obovate, 3-4 mm long, rounded<br />

at apex, hyaline in lower 2/3, white petaloid (with flat cells) and wrinkled above with erose margin;<br />

claw of inner bracts densely long woolly ciliate on margin, the wool enveloping the ovaries; stereome<br />

linear, hard, green, slightly shorter than claw, passing downwards into a very short hard stipe.<br />

Receptacle hemispherical, c. 3 mm high and wide, glabrous, smooth. Florets consistently 5 only one


Paul G. Wilson, Helipterum: Part 1<br />

Figure 4. Rhodanthe psammophila. A - Habit. B - Cluster of capitula. C - Capitulum. D - Florets with and without corolla.<br />

E - Carpopodium. F - Pappus bristle. G - Corolla. H - Style apex. I - Achene with pappus. J - Pericarp. K - Duplex<br />

myxogenic achenial hairs. From Paul G. Wilson 12604.<br />

405


406 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

of which is bisexual, the others male. Corolla actinomorphic, yellow, tube c. 2.5 mm long, sparsely<br />

puberulous, narrow-tubular below, narrow-turbinate above; lobes oblong, obtuse, c. 0.5 mm long, the<br />

vascular strands extending to tips, all papillose within along entire length. Stamens: anther loculi c. 1<br />

mm long; appendage ovate with narrow-oblong cells, the marginal ones smaller and equilateral; tails<br />

weak, slightly exceeding collar. Style apex truncate, penicillate, the vascular strand extending to apex.<br />

Achene compressed obovoid, c. 2.5 mm long, sparsely and minutely puberulous with blunt duplex hairs<br />

c. 0.06 mm long; carpopodium a very short glabrous ring; pericarp thinly crustaceous, pale reddish brown,<br />

the cells of epidermis amoeboid, thickened on radial walls; testa membranous, vascular strand beneath<br />

a pericarpial strand and passing over apex of seed; endosperm persistent around embryo. Pappus (of<br />

fertile achene) tardily deciduous; bristles c. 13, narrow-linear, acuminate, c. 2.0 mm long, sparsely ciliate<br />

with slender acuminate hairs. Achene deciduous along with inner involucral bracts.<br />

Distribution. Western Australia, near the coast between Carnarvon and Onslow, and inland to the<br />

Kennedy Range.<br />

Specimens seen (selection only). WESTERN AUSTRALIA: Kennedy Range, R.J. Cranfield 1913<br />

(PERTH); 10 mi S of Onslow, A.S. George 1150 (PERTH); southern margins of Lake McLeod, P.S. Short<br />

2491(MEL); 12 mi NW of Mt Sandiman, B.L. Turner 5401 (PERTH); Brown Range, B.L. Turner 5411<br />

(PERTH).<br />

Habitat. Grows only on sand dunes, both inland and coastal.<br />

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

Rhodanthe psammophila is superficially similar to R. condensata with which it has been confused<br />

in herbaria. The latter species may be distinguished by the absence of glandular hairs on the leaves,<br />

by the presence of 10-13 florets to a capitulum, and by the persistent pappus. The cells of the white lamina<br />

of the involucral bracts of R. condensata are raised at their tips giving it a slightly rough surface whereas<br />

in R. psammophila the lamina is smooth. Rhodanthe condensata is found from Shark Bay south to the<br />

Murchison River which is south of the distribution of R. psammophila.<br />

The relationships of this species are unclear. In some ways, such as in the structure of the floret,<br />

it is similar to Rhodanthe maryonii, but it differs from that species in having a caducous achene and a<br />

1-celled thick crustaceous pericarp (characters that, in this context, may be of little phyllogenetic<br />

significance).<br />

Rhodanthe tietkensii (F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum tietkensii F. Muell., Fragm., 8:227(1874). - Argyrocome tietkensii (F. Muell.) Kuntze, Revis.<br />

Gen. Pl. 1:309(1891). Type: Between the Alberga and Mt Olga, 1873/4, E. Giles (holo: MEL 110506).<br />

5. Rhodanthe sect. Citrinae Paul G. Wilson, sect. nov.<br />

Herba annua erecta sparse lanosa, non glandulosa. Folia caulina alterna linearia vel anguste<br />

oblonga. Capitula homogama vel heterogama, terminalia. Involucrum radians; bracteae multiseriatae,<br />

omnes chartaceae et petaloideae, exteriores sessiles, interiores unguiculatae; unguis oblongus vel


Paul G. Wilson, Helipterum: Part 1<br />

ellipticus, hyalinus, lanato-ciliatus, stereomate lineari applanato. Receptaculum glabrum, planum.<br />

Flosculi numerosi, homomorphi, actinomorphi, extra feminei vel bisexuales, intra bisexuales. Corolla infra<br />

tubulosa supra anguste turbinata, 4-5-loba, sparse pilosa; lobi glabri, intra laeves. Antherae: appendix<br />

anguste ovatus, tenuis, cellulis oblongis parietibus tenuibus; caudae filamentosae. Styli apex truncatus.<br />

Achenium anguste ellipsoideum, c. 1 mm longum, plerumque in rostrum glabrum attenuatum, breviter<br />

pilosum, pilis myxogeneis; pericarpium debile, cellulis epidermidis ± cubicis, parietibus tenuibus; testa<br />

chartacea, pallido brunea ad pericarpio affixa, crystallis applanatis munitis. Pappi setae imo basi<br />

connatae, breviter plumosae vel denticulatae, caducae.<br />

Holotypus: Rhodanthe citrina (Benth.) Paul G. Wilson<br />

Annual erect herbs, sparsely woolly, not gland dotted. Leaves cauline, alternate, linear or<br />

narrow-oblong. Capitula homogamous or heterogamous, terminal. Involucre radiant; bracts multiseriate,<br />

all glumaceous and petaloid, outer sessile, the inner with an oblong or elliptic hyaline woolly ciliate claw.<br />

Receptacle glabrous, flat. Florets numerous, homomorphous, regular, the outer female or bisexual, the<br />

inner bisexual. Corolla tubular below, narrow-turbinate above, 4-5-lobed, sparsely pilose; lobes<br />

glabrous, smooth within. Anther appendix narrow-ovate, c. 0.2 mm long, thin, cells oblong with thin walls;<br />

tails filamentose. Style apex truncate. Achene narrow-ellipsoid, c. 1 mm long, frequently narrowed at<br />

apex into a beak; shortly pilose with duplex myxogenic hairs; pericarp with epidermis of thin-walled<br />

cubical cells, other tissue flattened and inconspicuous, vascular strands lateral (slightly oblique); testa<br />

papery, pale brown, adnate to pericarp, sparsely furnished with broad, flat crystals or these absent when<br />

mature, vascular strand ascending to apex and adjacent to a pericarpial strand. Pappus caducous; setae<br />

connate into a ring at base, shortly plumose or barbellate.<br />

Rhodanthe citrina (Benth.) Paul G. Wilson, comb. nov.<br />

Leptorhynchos citrinus Benth. in Endl. et al., Enum. Pl. Hueg. 64(1837). Waitzia citrina (Benth.) Steetz<br />

in Lehm., Pl. Preiss. 1:454(1845). Type citation: 'Swan-River. (Hügel.).’n.v.<br />

Waitzia brevirostris Steetz in Lehm., op.cit. 1:451(1845). Type: In arenosis sylvae supra urbiculam<br />

"Perth", 7 Oct.1839, L. Preiss 15. Lectotype (here chosen): MEL 1585201; isolecto: LD, MEL 1585197.<br />

Waitzia sulphurea Steetz in Lehm., op.cit. 1:4553(1845). Lectotype (here chosen): In arenosis inter<br />

frutices prope urbiculam "Guildford", 14 Oct.1839, L. Preiss 7 (lecto: MEL 1585196; isolecto: LD).<br />

Waitzia steetziana Lehm., op.cit. 1:454(1845). - Waitzia tenella Hook., Bot. Mag. t.5342(1862) nom.illeg.<br />

Type: In solo limoso sylvae prope praedium rusticum "Maddington", 31 Oct.1839, L. Preiss 6 (syntypes<br />

and isosyntypes: LD, MEL 1585194, 1585195, 1585198).<br />

Waitzia dasycarpa Turcz., Bull. Soc. Imp. Naturalistes Moscou 24/2:77(1851). Type: Swan River Colony,<br />

J. Drummond 5th coll. no. 65 (holo: KW photo seen; iso: PERTH).<br />

Helichrysum oldfieldii F.Muell., Fragm. 3:134-135(1863). Lectotype (here chosen): Murchison<br />

River, A. Oldfield (MEL 108299).<br />

407


408 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Distribution. Western Australia south of 22° lat., southern Northern Territory, northern South Australia,<br />

north western New South Wales.<br />

Note: The genus Waitzia, as previously circumscribed, was an unnatural assemblage of species in the<br />

Helipterum complex that had in common a beak-like apex to the achene. In the strict sense (see Wilson<br />

1992b) Waitzia contains species with the following characters: stipe-like claws to the involucral bracts,<br />

tooth-like papillae (that consist of 2 overtopping cells) on the achenes, and ellipsoid apices to the styles.<br />

In Rhodanthe citrina the inner involucral bracts have flattened scarious claws, the achenes have normal<br />

slender duplex hairs, and the style apices are truncate; it is thus distinct from Waitzia but similar to some<br />

species of Rhodanthe sect. Achyroclinoides. The section Citrinae differs from sect. Achyroclinoides<br />

principally in the nature of the involucral bracts which are all scarious and petaloid, the outer sessile and<br />

the inner stipitate on a broad hyaline ciliate claw; none of them is radiant. The appearance of the capitula<br />

is similar to that found in some species of Chrysocephalum (Helichrysum sect. Chrysocephalum), such<br />

as Chrysocephalum apiculatum (Labill.) Steetz, and the two species are sometimes confused.<br />

In Rhodanthe citrina the rostrum to the achene is variable in length and in some specimens is<br />

completely absent; that organ, in any event, does not seem to be necessarily of generic significance since<br />

it merely represents an upwards extension of the pericarp.<br />

It has been shown by Warcup (1990) that Waitzia citrina forms ectomycorrhizal associations<br />

whereas other Rhodanthe species that have been tested do not. It is possible that the seed with which<br />

Warcup worked was either Hyalosperma cotula (Benth.) Paul G. Wilson or Chrysocephalum apiculatum,<br />

both of these form ectomycorrhiza and both resemble W. citrina. No voucher material can be traced.<br />

Two other species previously placed in Waitzia, i.e. W. paniculata (Steetz) Benth. and W. conica<br />

B.Turner, are also anomalous in that genus and are here transferred to Pterochaeta and Haptotrichion<br />

respectively.<br />

6. Rhodanthe sect. Synachyrum (A. Gray) Paul G. Wilson, comb. nov.<br />

Helipterum sect. Synachyrum A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:231(1852). Type: Helipterum<br />

floribundum DC.<br />

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

Plants cottony. Intermediate involucral bracts with thick narrow-oblong stereome not extending into<br />

lamina; cottony. Corolla with vascular strands not extending to tips of lobes. Style apex truncate. Achene<br />

densely silky; carpopodium absent; pericarp thinly coriaceous or papery; testa weak, free from pericarp,<br />

containing oblong imbricate crystals that form a complete cover to seed (see Figure 5). Pappus bristles<br />

narrow-oblong and sometimes partially united into a tube at base, caducous.<br />

Rhodanthe floribunda (DC.) Paul G. Wilson, comb. nov. (Figure 5)<br />

Helipterum floribundum DC., Prod. 6:217(1838). Type: Barren forests at the foot of Peel’s Range,<br />

Interior of N.S.Wales, June 1817, A.Cunningham 107 (holo: G-DC photo seen).


Paul G. Wilson, Helipterum: Part 1<br />

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

Helipterum chionolepis F. Muell., Linnaea 25:416(1853). Type citation: 'Juxta montes Flinders-range et<br />

Cudnaka locis sterilibus.’ Lectotype (here chosen): MEL 110120 with labels 'Cudnaka’ and 'Prope montis<br />

Flinders range, Oct.51’ leg. F.Mueller; syntype: Cudnaka, F.Mueller (MEL 604828).<br />

Helipterum cirratum Morrison, J.Bot. 50:168(1912). Type: Between Globe Hill and Uaroo, Ashburton<br />

River, 1 Oct. 1905, A.Morrison (? iso: PERTH).<br />

Helipterum floribundum var. tubulipappum J. Black, Trans. & Proc. Roy. Soc. South Australia 36:23<br />

t.2(1912). Type: Oodnadatta, Sept.1911, F.C. Staer per Mrs Mellor (holo: AD 98625088 p.p.).<br />

Rhodanthe sphaerocephala Paul G. Wilson, sp. nov.<br />

Annua erecta ad 25 cm alta. Rami lanosi, omnino foliacei. Folia alterna sessilia, angustissimo<br />

oblonga, c. 10 mm longa, sparse lanosa. Capitula homogama, solitaria, terminalia, sphaeroidea, c.<br />

10 mm diam. Bracteae involucri multi-seriatae, aliquantum corrugatae, nitidae. Bracteae intermediae<br />

in ambito obovatae, c. 5 mm longae; unguis latissime oblongus, hyalinus; stereoma spatulatum, parte<br />

terminali incrassata et dense lanosa; lamina late elliptica, integra, straminea. Receptaculum conicum,<br />

tuberculatum. Flosculi numerosi, bisexuales, actinomorphi. Corolla c. 4 mm longa, glabra; lobi 5,<br />

anguste oblongi, c. 1.5 mm longi, intra laeves; nervi corollae ad apice tubi extensi. Antherae:<br />

appendix sterilis ovata, acuta, c. 0.6 mm longa, cellulis oblongis, parietibus nec incrassatis; caudis<br />

filamentis debiles. Styli apex truncatus, nervo prominenti ad basem apicis extenso. Achenium<br />

ellipsoideum c. 2 mm longum, dense sericeum, pilis in lobos inaequales terminantibus; pericarpum<br />

membranaceum, translucens; testa tenuiter coriacea, pallide brunea, cellulis crystallis oblongis<br />

409


410 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

repletis. Pappi setae librae, caducae, in parte inferiore angusto ellipticae, breviter plumosae, in parte<br />

superiore filiformae, sparse plumosae, ad extremum plumoso-penicillatae.<br />

Typus: Belele Station, c. 56 km WNW of Meekatharra, Western Australia, 30 Oct. 1965, D.W. Goodall<br />

3366 (holo: PERTH; iso: CANB).<br />

Erect annual to 25 cm high branching at and shortly above base. Stems loosely woolly, leafy<br />

throughout. Leaves cauline, alternate, sessile, very narrow-oblong, c. 10 mm long, 1 mm wide, sparsely<br />

woolly with crinkly hairs. Capitula homogamous, solitary, terminal to long branches, almost spherical,<br />

c. 10 mm diameter. Involucral bracts multi-seriate, somewhat crinkly, glossy, straw-coloured, similar<br />

throughout; intermediate bracts obovate in outline, c. 5 mm long; claw very broad-oblong, hyaline,<br />

woolly ciliate on distal margins, stereome spatulate with the terminal portion thickened and bearing a<br />

dense tuft of wool on abaxial surface, lamina broad-elliptic, entire, somewhat wrinkled, glabrous, glossy,<br />

straw-coloured. Receptacle ovoid, c. 3 mm high, 2.5 mm wide, glabrous, tuberculate with raised achenial<br />

scars. Florets numerous, homogamous, bisexual, actinomorphic. Corolla c. 4 mm long to base of lobes;<br />

tube narrow-cylindrical, glabrous; limb narrow-campanulate, glabrous; lobes 5, narrow-oblong, c. 1.5 mm<br />

long with a few glandular hairs (with large oblongoid terminal cells) on abaxial side, smooth within,<br />

vascular strands reaching to base of lobes. Anther appendage ovate, acute, c. 0.6 mm long, cells oblong<br />

with unthickened walls, marginal cells not differentiated; anther tails filamentous, weak. Style tip<br />

truncate, shortly penicillate, the vascular strand stout, not extending into apex. Achene ellipsoidal, c.<br />

2 mm long, densely long-silky to base, the duplex hairs terminating in two very uneven arms with one<br />

very short (c. 0.005-0.01 mm) and the other long (c. 0.3 mm), not myxogenic; pericarp thin, translucent;<br />

testa thinly coriaceous, pale brown, the outer cells filled with flat broad-oblong crystals that form an<br />

armour-like covering to seed. Pappus bristles slightly shorter than corolla, free, caducous, linear-elliptic<br />

and shortly plumose in lower half, filiform and sparsely plumose in upper half, plumose tufted at end.<br />

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

Rhodanthe sphaerocephala would appear to have no close relative but the morphology of its florets<br />

and fruits suggests affinity to other members of the section Synachyrum.<br />

Rhodanthe sterilescens (F. Muell.) Paul Wilson, comb. nov.<br />

Helipterum sterilescens F. Muell., S. Sci. Rec. 2:274(1882). - Argyrocome sterilescens (F. Muell.)<br />

O. Kuntze, Rev. Gen. Pl. 1: 309 (1891). Type: Gascoyne River, W.A., 1882, Pollack (holo: MEL 110428;<br />

iso: PERTH).<br />

This species bears dense clusters of capitula that give it a similar appearance to Cephalipterum<br />

drummondii with which it has been frequently confused.<br />

Rhodanthe sterilescens differs from C. drummondii in the following significant characters: sessile<br />

capitula at base of plant in addition to terminal (not all terminal); capitula in a compact cyme (not<br />

dense umbel); indumentum woolly (not of curled filiform hairs with septate base and glandular<br />

stipitate hairs); achenial hairs slender of the normal duplex type (not clavate helically coiled and


Paul G. Wilson, Helipterum: Part 1<br />

barbed woolly); achenes free, three fertile and silky, the rest sterile and glabrous (not coalescent with<br />

1 or two fertile and the remainder sterile but hairy); achene narrowed at apex (not broad and lobed at apex).<br />

The two species are similar in floral characters, in the seed morphology, and in form of the crystals present<br />

in the testa.<br />

Rhodanthe stuartiana (Sond. & F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum stuartianum Sond. & F. Muell., Linnaea 25: 518 (1853). - Helipterum floribundum var.<br />

stuartianum (Sond. & F. Muell.) Benth., Fl. Austral. 3:642(1867). Type citation: 'Ad fl. Murray leg. Stuart.’<br />

Lectotype (here chosen): Ad fl. Murray, F. Mueller (MEL 604830).<br />

The type of Helipterum stuartianum is stated to have been collected by Stuart. A herbarium<br />

O.W. Sonder specimen (MEL 604830) labelled 'Ad fl. Murray’ was collected by F. Mueller and bears in<br />

(?)Sonder’s handwriting the phrase 'Antherae basi bisetae’; these words are used in the published<br />

description. A further collection from 'Near the Murray R.’ no. 785 (MEL 1539201) was evidently from<br />

Stuart. It bears the note 'This plant did not occur in my collection at Adelaide but I found seed of it amongst<br />

those you procured from Stephens’ [? William Stephens, a garden’s collector]. I have selected the<br />

Mueller collection (MEL 604830) as the lectotype since it was evidently studied by Sonder.<br />

Rhodanthe troedelii (F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum troedelii F. Muell., Victorian Naturalist 7:77(1890). Type citation: 'Near the Barrier-Ranges;<br />

Mrs Irvine, at Leight’s Creeks, beyond Beltana; Mrs Richards.’ Lectotype (here chosen): Leigh’s Creek,<br />

1887, Mrs Richards (lecto: MEL 110636; isolecto: MEL 696324, NSW 181421).<br />

Helipterum troedelii var. patens Ewart, J. White & B. Rees, Proc. Roy. Soc. Victoria 22 n.s.:15(1909) pro<br />

parte as to lectotype. - H.roseum var. patens (Ewart et al.) J. Black, Trans. & Proc. Roy. Soc. South<br />

Australia 45:21(1921). Type citation: 'Mt. Lyndhurst, M. Koch, No.1644 (1899); Fraser Range, W. Austr.,<br />

R. Helms, 1891.’ Lectotype: Mt Lyndhurst, S. Australia, M. Koch 1644 (1899), AD 97650128, fide D. Cooke<br />

in Jessop & Toelken, Fl. South Australia edn 4, 1549(1986).<br />

The syntype material of H.troedelii var. patens consists of two species. The collection of Max<br />

Koch represents Rhodanthe troedelii while the collection of R. Helms represents the 'Nullarbor’<br />

variant of R. chlorocephala. J.M. Black (1921 l.c.) indicated that he had examined duplicate material<br />

of the Helms syntype and recognised it as being a variant of Helipterum roseum, he therefore<br />

transferred var. patens to that species (which is a synonym of Rhodanthe chlorocephala). However,<br />

D. Cooke (1986 l.c.) lectotypified the name on a Max Koch collection in herb. AD which had been<br />

received by Black as a donation from MEL and is probably a duplicate of the Max Koch syntype (it<br />

does not bear the month of collection that is present on the syntype). The AD specimen had been<br />

annotated by Ewart as 'typical H. Troedelii’. In herb. MEL the Max Koch syntype has had the epithet<br />

'var. patens’ deleted, presumably by Ewart, while the Helms syntype has been labelled 'type’. It would<br />

therefore appear that subsequent to its publication Ewart intended to apply the epithet var. patens to<br />

the element represented by the Helms collection which was also the way in which Black applied the<br />

name (as a synonym of H. roseum) in 1921 and in the Flora of South Australia (1929).<br />

411


412 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

7. Rhodanthe sect. Helipteridium (A. Gray) Paul G. Wilson, comb. nov.<br />

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

discoideum A. Gray<br />

Helipterum sect. Aglossum F. Muell., Fragm. 3:137(1863) nomen.<br />

Annual erect herb, sparsely woolly with filiform hairs, and with curled uniseriate septate hairs,<br />

eglandular. Major axes slender. Leaves alternate, flat, sessile. Capitula heterogamous, solitary, terminal.<br />

Involucre hemispherical, not radiant. Receptacle glabrous with prominent alveolae. Involucral bracts<br />

c. 4-seriate, glabrous, glossy, uniform; claw broad-oblong with a broad herbaceous stereome and narrow<br />

scarious margins; lamina short, erect, scarious. Florets bisexual or the inner male. Corolla slightly<br />

zygomorphic; tube cylindrical, pubescent with gland-tipped hairs; throat turbinate, glabrous; lobes 5,<br />

unequal, papillose within, vascular strands extending to apex of lobes. Anther appendix deltoid, cells<br />

short and irregular; anther tails filamentous, weak. Style apex truncate with a subulate coalescence of<br />

epidermal cells in the centre. Achene ellipsoid, silky pilose with normal duplex hairs; carpopodium<br />

annular; pericarp brittle; testa soft with numerous oblong crystals, lower 2/3 surrounded by a network<br />

of vascular strands. Pappus bristles linear-acuminate, plumose.<br />

A monotypic section.<br />

Rhodanthe heterantha (Turcz.) Paul G. Wilson, comb. nov.<br />

Helipterum heteranthum Turcz., Bull. Soc. Nat. Mosc. 24/1 :198 (1851). Type: Nova Hollandia,<br />

J. Drummond 4th coll. n.214 (holo: KW photo seen; iso: K, MEL 110231, 110233).<br />

Helipterum discoideum A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:231(1852). Type citation: 'Variat<br />

a, involucro pallido; ß, involucro sanguineo. Swan River (ß, Swan River to King George’s Sound),<br />

Drummond.’ (syn: K 'J. Drummond 96’, photo seen).<br />

Helipterum anactinum F. Muell., Fragm.3:137(1863). Type citation: 'Ad sinum orarium Champion Bay.<br />

Walcott et Oldfield.’ Lectotype (here chosen): Champion Bay, A. Oldfield (lecto: MEL 110229).<br />

Helipterum heteranthum var. majus Benth., Fl. Austral. 3:643(1867). Type: Between Moore and<br />

Murchison rivers, J. Drummond 6th coll. n.152 (holo: K; iso: MEL 110226, NSW).<br />

Helipterum pachychaetum W.Fitzg., J. W. Austral. Nat. Hist. Soc. No.2:25(May 1905). Lectotype (here<br />

chosen): Jacup, 50 miles west of Phillips River, Oct.1903, C.R.P. Andrews (lecto: PERTH; isolecto: NSW).<br />

Helipterum heteranthum var. minor Ewart, J. White & Tovey, J. Roy. Soc. N.S.Wales 42:190(1908).<br />

Lectotype (here chosen): Cowcowing, W.A., Oct.1904, M. Koch 1108 (lecto: MEL 110220; isolecto: MEL<br />

110221).<br />

8. Rhodanthe sect. Helichrysoides (A. Gray) Paul G. Wilson, comb. nov.<br />

Pteropogon sect. Helichrysoides A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:267(1852). Type:<br />

Pteropogon spicatus Steetz


Paul G. Wilson, Helipterum: Part 1<br />

Annual erect herbs, woolly or glandular puberulous. Leaves cauline, alternate, filiform to obovate.<br />

Capitula homogamous or heterogamous, clustered, paniculate or condensed in spike-like inflorescences.<br />

Involucre turbinate to cylindrical, not radiant; bracts multiseriate, glabrous or glandular pubescent,<br />

scarious or with herbaceous tips; stereome small, basal. Receptacle convex to broad-conical, glabrous<br />

or scabrid. Florets 5-30, bisexual, or the innermost male. Corolla actinomorhpic , sparsely puberulous,<br />

narrow-cylindrical below, narrow-campanulate above; lobes ovate, papillose or smooth within, vascular<br />

strands extending to tips. Stamens: anther apex ovate the distal cells slightly raised at tip; tails<br />

filamentous, weak. Style apex broad to narrow-deltoid, puberulous, vascular strand extending to tip.<br />

Achene narrow-obovoid to narrow-turbinate, pilose almost to base; carpopodium an extremely short<br />

ring; pericarp rough, thick, brittle, dark brown; testa free from pericarp, membranous, vascular strand<br />

extending almost completely around seed; crystals absent. Pappus persistent; bristles thick at base,<br />

plumose, the cilia all acute.<br />

The section Helichrysoides is similar to sect. Leiochrysum but differs in the nature of the involucral<br />

bracts (foliaceous at apex except in R. spicata), the presence of vascular strands at the apex of the corolla<br />

lobes, the deltoid style apex, and the thick brittle pericarp. It is also similar to species in the genus<br />

Podotheca in the morphology of the achene, the deltoid style apex, and the foliaceous apex to the<br />

involucral bracts. In the latter genus the chromosome number is n=13, 26 (Short 1989) whereas in<br />

Rhodanthe sect. Helichrysoides the three species R. battii, R. pollackii, and R. spicata, have a<br />

chromosome number of n = 10 (B. Turner in sched.); the number for R. charsleyae has not been recorded.<br />

Warcup (1990) has shown that R. battii and R. spicata do not form ectomycorrhiza whereas species of<br />

Podotheca that have been tested do; this again suggests that the species in sect. Helichrysoides should<br />

not be placed in Podotheca.<br />

Rhodanthe battii (F. Muell.) Paul. G. Wilson, comb. nov.<br />

Helipterum battii F.Muell., Victorian Naturalist 10:144(1893). Type: Between Dundas-Hills and Lake-<br />

Lefroy, 1893, J.D. Batt (holo: MEL 110727).<br />

[Podotheca pollackii auct. non (F.Muell.) Diels: Diels & Pritzel, Bot. Jahrb. Syst. 35:615-623(1905) as<br />

to specimen cited and description.]<br />

Rhodanthe pollackii (F. Muell.) Paul G. Wilson, comb. nov.<br />

Podosperma pollackii F. Muell., Fragm. 12:21(1882); - Podotheca pollackii (F. Muell.) Diels, Bot. Jahrb.<br />

Syst. 35:617(1905). Type citation: 'In vicinia fluminis Gascoyne-River; Forrest et Pollack.’ Syntype:<br />

Gascoyne River, 1882, J. Forrest (PERTH).<br />

Rhodanthe charsleyae (F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum charsleyae F. Muell., Fragm. 8:168(1874). Type citation:'In vicinia lacus Amadei, Giles; prope<br />

lacum Lefroyi, Forrest.’ Lectotype (here chosen): Lake Lefroy, State Well, A. Forrest (MEL 110714).<br />

Rhodanthe spicata (Steetz) Paul G. Wilson, comb. nov.<br />

Pteropogon spicatus Steetz in Lehm., Pl.Preiss. 1:479(1845); A. Gray, Hooker’s J. Bot. Kew Gard.<br />

Misc. 4:268(1852). - Helipterum spicatum (Steetz) Benth., Fl. Austral. 3:647(1867). Type citation:<br />

413


414 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

'In limoso-calculosis illustribus lateris orientalis montis Lehmann’, 4 Sept.1839, L. Preiss 24.<br />

Lectotype (here chosen): MEL 110492; isolecto MEL 110493, MEL 604832.<br />

Helipterum monencyanthoides F. Muell., Fragm. 3:137(1863). - H. spicatum var. pallens Benth.,<br />

Fl. Austral. 3:648(1867). Type citation: 'Ad rivos Tom [i.e. Tone] River et Salt River’, G.Maxwell.<br />

Lectotype (here chosen): Banks and valleys of Salt River, G.Maxwell 75 (MEL 110707).<br />

Calocephalus globosus M.Scott & Hutch., Kew Bull. 1916:36(1916). Type: Kauring, on York - Greenhills<br />

line, Western Australia, F. Stoward 505 (holo: K photo seen).<br />

9. Rhodanthe sect. Actinaria Paul G. Wilson, sect. nov.<br />

Herba annua erecta sparse glanduloso puberula. Folia caulina, alterna, applanata. Capitula<br />

homogama, solitaria, terminalia. Involucrum cupulatum ad hemisphaericum, radians; bracteae manifeste<br />

dimorphae, multiseriatae; bractea exteriora anguste linearia, acuminata, glanduloso puberula; bracteae<br />

interiores ungue anguste oblongo, lamina ovata, petaloidea. Receptaculum pulvinatum, glabrum, laeve.<br />

Flosculi numerosi, bisexuales, homomorphi, actinomorphi. Corolla infra tubulosa supra anguste<br />

turbinata, pallido flava, 5-loba; lobi glabri, extra sparse pilosi, intra laeves. Antherae: appendix anguste<br />

ovatus, cellulis anguste oblongis parietibus tenuibus; caudae parte proximali firmae, parte distali<br />

filamentosae debiles. Styli apex ellipsoideus, nervo carenti. Achenium cylindraceum, c. 1 mm longum,<br />

modice pilosum, pilorum longitudine dissimili (0.02-0.2 mm); carpophorum annulare; pericarpium<br />

unicellulam crassum, crustaceum; testa membranacea, unicellulam crassum, translucens, crystallis<br />

carentibus. Pappi setae filiformes,imo basi connatae, in statu integro caducae, breviter plumosae.<br />

Typus: Rhodanthe margarethae (F. Muell.) Paul G. Wilson<br />

Erect annual herbs, branched, sparsely glandular puberulous. Branches slender, reddish. Leaves<br />

cauline, alternate, thin. Capitula homogamous, solitary, terminal to branches. Involucre cup-shaped to<br />

hemispherical, c. 2 cm diameter, radiant; bracts distinctly dimorphic; outer bracts narrow-linear,<br />

acuminate, c. 4 mm long, glandular puberulous; inner bracts with erect narrow-oblong woolly ciliate claw<br />

and spreading ovate petaloid white or pale yellow lamina, the stereome narrow-oblong, thick and<br />

glandular puberulous. Receptacle cushion-shaped, c. 5 mm diameter, smooth, papillose around alveolae.<br />

Florets numerous, bisexual, 5-merous, actinomorphic. Corolla narrow-tubular below, expanded above,<br />

pale yellow, very sparsely puberulous outside, lobes smooth within, cells of inner epidermis of lobes<br />

oblong. Stamens: anther tip narrow-ovate acute, cells narrow-oblong, the marginal cells forming an<br />

indeterminate row; anther tails firm, equal to collar. Style apex elliptic, shortly papillose, without<br />

vasculature. Achene cylindrical c. 1 mm long, moderately short pilose, hairs rounded at apex and of<br />

different lengths (0.02-0.2 mm); carpopodium a short glabrous ring; pericarp thin and crustaceous,<br />

brown; testa thin, translucent, without crystals. Pappus caducous as a whole; bristles filiform, shortly<br />

plumose throughout, connate in ring at base.<br />

A section of two morphologically similar species that are restricted in their distribution to the<br />

northwestern region of Western Australia.<br />

The section Actinaria differs from other members of the Rhodanthe complex in having narrowlinear<br />

outer involucral bracts, elliptic style apices that are without vasculature, filiform pappus<br />

bristles, and short blunt achenial hairs that are of different lengths. The habit of the two included


Paul G. Wilson, Helipterum: Part 1<br />

species is similar to that of R. manglesii in section Rhodanthe, as was noted by Mueller in his description<br />

of H. margarethae, but in floral characters the sections are very distinct.<br />

Rhodanthe margarethae (F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum margarethae F. Muell., Fragm. 11:48(1878); F. Muell., Syst. Census Austral. Pl. 80(1882)<br />

'Margaritae’; F. Muell., Sec. Syst. Census Austral. Pl. 136(1889) 'Margaritae’; Domin, Biblioth. Bot.<br />

89:667(1929); Argyrocome margarethae (F. Muell.) Kuntze, Revis. Gen. Pl. 1:309(1891) 'Margaritae’.<br />

Type citation:'Ad amnem Jones’s Creek et flumen George's River prope Nickol-Bay; J. Forrest’. Lectotype<br />

(here chosen): Jone’s Creek and George River south of Roebourne, 1878, J. Forrest (MEL 110234).<br />

Distribution. Pilbara region (Fortescue Botanical District) of Western Australia.<br />

I have not found a syntype that was collected by J. Forrest near Nickol Bay, however, there is a<br />

specimen (MEL 110235) labelled by Mueller 'Helipterum Margarethae, Nickol River, A. Forrest’ which<br />

is possibly the collection referred to by him since Alexander Forrest was with John Forrest on the<br />

expedition to the Nickol Bay area in 1878. This A. Forrest collection is of Rhodanthe frenchii. Mueller<br />

describes the leaves as 'amplexantibus’ and the involucral bracts as 'candida’, that is, pure white, terms<br />

that apply to R. margarethae as lectotypified by the 'Jone’s creek and George River’ specimen, but not<br />

to the Nickol River collection.<br />

The epithet 'margarethae’ honours Margaret Forrest (1845-1929), wife of John Forrest the explorer<br />

and statesman. Mueller evidently considered a preferred latinization of her name to be 'margaritae’ for<br />

he used this variant in 1882 and 1889 as did Otto Kuntze in 1891.<br />

Rhodanthe frenchii (F. Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum frenchii F. Muell., S. Sci. Rec. 3:34(1883). Type citation: 'Near Menilayalya in the vicinity<br />

of Shark-Bay; J. Forrest’ (holo: MEL 50293 'Head of Menilyalya R., 1882, J. Forrest’).<br />

[Helipterum margarethae F. Muell., Fragm. 11:48(1878) pro parte as to the Nickol River collection, not<br />

as to lectotype]<br />

Distribution. Northern Carnarvon and north-west Ashburton Botanical districts of Western Australia.<br />

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

Anisolepis Steetz in Lehm., Pl.Preiss. 1:446(1845). Type: Anisolepis pyrethrum Steetz<br />

Annual erect herb, glabrous or sparsely and minutely puberulous. Stem simple or branched above,<br />

towards the base thickened and aerenchymatous. Leaves cauline, simple, entire, alternate or the lower<br />

opposite, if submerged filiform, otherwise oblong and somewhat fleshy, c. 5 mm long, towards the<br />

apex becoming ovate and with scarious margins. Capitula homogamous, solitary and terminal to stem<br />

and branches. Involucre at first turbinate, becoming hemispherical with age, often subtended by the<br />

uppermost leaves; bracts 2-seriate, all radiant and with claws: claw erect, narrow-deltoid, c. 1 mm<br />

long, margin scarious, stereome prominent, narrow-oblong, thick and raised, sparsely glandular, limb<br />

415


416 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

petaloid, radiating, white, elliptic, c. 5 mm long, eventually breaking away from the persistent claw.<br />

Receptacle narrow-conical, deeply foveolate, fimbrilliferous around foveolae. Florets numerous,<br />

bisexual. Corolla tube cylindrical, c. 1.5 mm long, sub-campanulate above, shortly 5-lobed; lobes<br />

triangular, abaxial surface with a few hairs that have large ovoid gland-cells at their tips, two of the lobes<br />

with adaxial surface papillose at base, the others glabrous, margins thickened; cells of inner epidermis<br />

of lobes irregularly oblong, of the neck oblong and undulate on margin. Anthers included; terminal<br />

appendage broad-oblong, obtuse, thin, cells oblong, thin-walled; tails weak and filamentous. Style tips<br />

rounded to acuminate; vascular strand extending to base of tip. Achene barrel-shaped, c. 1 mm long,<br />

dark reddish brown, pilose with fine hairs, not myxogenic; carpopodium very short, annular; pericarp<br />

very thinly crustaceous, pale brown; testa mauve, thinly coriaceous, vascular strands not apparent;<br />

crystals absent. Pappus c. 1 mm long of c. 10 narrowly triangular serrate scales united in lower half, tardily<br />

deciduous as a whole.<br />

Rhodanthe pyrethrum (Steetz) Paul G. Wilson, comb. nov.<br />

Anisolepis pyrethrum Steetz in Lehm., Pl. Preiss. 1:447(1845). - Helipterum pyrethrum (Steetz) Benth.,<br />

Fl. Austral. 3:642(1867). - Argyrocome pyrethrum (Steetz) Kuntze, Revis. Gen. Pl. 1:309(1891). Type<br />

citation: 'In depressis uliginosis sylvae supra oppidulum Perth, d. 26.Sept.1839. Herb. Preiss. No.14.’<br />

Lectotype (here chosen): MEL 108293; isolecto: MEL 108294.<br />

There is some variability within this species but it is unclear as to how much is due to growth<br />

conditions. Collections made to the north of Perth have the lower (underwater) leaves linear-acuminate<br />

and opposite, whereas collections made to the south of Perth have the underwater leaves filiform and<br />

scattered. There are also slight differences between different populations in the size of achenes, in the<br />

distribution of the globular glands, and in the branching of the inflorescence. However, all these<br />

characters show such variability over the total species range that specific or infraspecific discrimination<br />

does not appear practicable.<br />

Rhodanthe pyrethrum is similar in some features to the species in Rhodanthe sect. Achyroclinoides,<br />

noticeably in the size and morphology of the achene. It differs from this section and from other sections<br />

of Rhodanthe, in the nature of the involucral bracts which are all radiant and which have claws that<br />

possess a very thick narrow-oblong stereome, in the laciniate scales of the pappus, and in the rounded<br />

to acuminate style apices.<br />

11. Rhodanthe sect. Polyphyllum Paul G. Wilson, sect. nov.<br />

Herba annua, modice lanosa, glandis sessilibus globosis sparse ornata. Folia numerosa, alterna,<br />

filiformia. Inflorescentium corymbosum. Capitula homogama, radiantia. Involucrum late turbinatum c.<br />

5 mm altum; bracteae intermediae scariosae, sparse lanosae, stereomate anguste oblongo, crasso;<br />

bracteae intimae similis sed limbo albo ornato. Flores 8-12, bisexuales; corolla tubularis, sparse<br />

glanduloso puberula, lobis 5, brevibus, intra glabris, nervis ad apicem extensis. Antherum appendice<br />

late ovato, crasso, caudis collum aequantibus, crassis, ramosis. Styli apex truncati, nervo crasso subapice<br />

extenso. Achenium doleiforme, truncatum, modice hirtellum; carpopodium crassum; pericarpium<br />

crustaceum, rugosum; testa chartacea, libra, nervo 3/4 circumdato; nervis pericarpi et testae in posite<br />

laterali locatis. Pappus corollam aequans; setae filamentosae, breviter plumosae, caducae.<br />

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


Paul G. Wilson, Helipterum: Part 1<br />

Annual erect herb, somewhat woolly and with scattered minute globular reddish brown glandular<br />

hairs, single stemmed. Leaves numerous, filiform. Inflorescence a terminal corymb. Capitula homogamous,<br />

shortly pedunculate, radiant. Involucre broadly turbinate c. 5 mm high; bracts c. 4-seriate; outer and<br />

intermediate bracts scarious, sparsely woolly with a thick narrow-oblong stereome; innermost bracts<br />

similar but with a short oblong white limb. Florets 8-12, bisexual; corolla narrow-tubular, slightly broader<br />

above, sparsely glandular puberulous; lobes 5 short, glabrous within, sparsely glandular puberulous<br />

outside, cells of inner epidermis narrow-oblong, vascular strands passing to apex of lobes. Anther<br />

appendage broad-ovate, thick, cells unevenly oblong; anther tails stout, branched, equal to collar. Style<br />

apex truncate, vascular strand thick and extending to just below apex. Achene barrel-shaped with a<br />

truncate apex, moderately hirtellous with slender duplex hairs; carpopodium thick; pericarp crustaceous,<br />

rough; testa papery, free from pericarp, vascular strand passing over apex of seed; vascular strands of<br />

pericarp and of testa in lateral position (in relation to cotyledons). Pappus equal to corolla; bristles<br />

filamentous, shortly plumose, shortly united at base, caducous.<br />

Note. A number of features make this monotypic section unique in the Helichrysum complex; these are:<br />

1) the numerous cauline, lanate, filiform leaves, 2) the short, stout, branched anther tails, and 3) the thick<br />

anther appendage with unevenly shaped oblong cells. Its position in relation to the other sections of<br />

Rhodanthe and to genera in the Casssinia complex is unclear. It is likely that this section should be<br />

recognised as a distinct genus.<br />

Rhodanthe polyphylla (F.Muell.) Paul G. Wilson, comb. nov.<br />

Helipterum polyphyllum F. Muell., Fragm. 1:35(1858). - Argyrocome polyphylla (F. Muell.) Kuntze,<br />

Revis. Gen. Pl. 1:309(1891). Type citation: 'In planitiebus basalticis a fluvio Brisbane usque ad montes<br />

Peak Range. - In tractu Kent’s Plains. W. Hill.’ Lectotype (here chosen): Brisbane River, Peak Range,<br />

F. Mueller (MEL 109740); syntype: Moreton Bay, Kents Plains, anon.(MEL 109757).<br />

Distribution. Eastern Queensland and north-east New South Wales.<br />

Cephalipterum A. Gray<br />

Cephalipterum A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:271(1852). Type: Cephalipterum drummondii<br />

A. Gray<br />

Annual erect herb, indumentum of curled filiform hairs septate at base with glandular stipitate<br />

hairs below capitula. Leaves cauline and basal, linear to obovate. Capitula heterogamous, very<br />

shortly pedunculate in dense terminal umbels. Involucre cup-shaped; bracts multiseriate, scarious,<br />

the innermost with a white or yellow lamina. Receptacle small, conical, glabrous. Florets numerous;<br />

1-2 outer bisexual, remainder male. Corolla narrow-urceolate, shortly 5-lobed, glabrous, lobes<br />

smooth within; vascular strands terminating in tube. Anthers: appendix ovate with narrow-oblong<br />

cells; collar short and broadly expanded at base; tails weak; style apex truncate, somewhat broader<br />

than style branch, prominently papillose. Achene broadly compressed obovoid with a broad-circular<br />

apex that has a hard raised lip on the abaxial margin; sterile achenes hard, broad-obovoid, the apical<br />

surface vertical; attachment small, carpopodium absent; indumentum (fertile and sterile achenes) of<br />

dense white clavate helically coiled duplex hairs on abaxial surface, of woolly clawed simple hairs<br />

on adaxial surface; pericarp thick and hard; testa thinly coriaceous closely covered all over with broad-<br />

417


418 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

oblong imbricate crystals; vascular strands of pericarp and testa medial in relation to the broadobovate<br />

embryo. Pappus bristles free, caducous, linear to filiform, shortly plumose in upper half, the<br />

terminal cilia congested, thickened and recurved.<br />

Cephalipterum drummondii A. Gray, op.cit 272. Type citation: 'Swan River, Drummond.’ n.v.<br />

Cephalipterum drummondii f. major Diels & Pritz., Bot. Jahrb. 35:615(1905) nom.illeg. (based on type<br />

forma).<br />

Cephalipterum drummondii f. minor Diels & Pritz., l.c.<br />

Type citation: 'hab. in distr. Coolgardie pr. Kanowna flor. m. Aug. (W.V. Fitzgerald in hb. Berl.); in distr.<br />

Austin pr. Murrinmurrin fl. m. Nov. (W.J. George).’ Neotype (here chosen): Kanowna, August 1898,<br />

W.V.Fitzgerald (PERTH).<br />

Note. The syntype material in herb. B is destroyed, I have therefore chosen as neotype a probable<br />

isosyntype in PERTH.<br />

This genus is similar in many features to species in Rhodanthe sect. Synachyrum, in particular in<br />

possessing a dense imbricate crystalline cover to the seed. Only one or two seeds in the capitulum are<br />

fertile but these coalesce to the sterile achenes by means of a dense mass of barbed hairs on their adaxial<br />

surfaces which cause the entire cluster to be dispersed as one unit from which one seedling emerges<br />

with the sterile achenes forming a persistent ring around the base of the root. The fertile achenes bear<br />

at their tip a raised rim on their abaxial margin; this character is unique in the Rhodanthe complex.<br />

A number of variants of this species occur which vary in the size and colour of the involucral bracts<br />

with sometimes two or more variants being present at the same locality. It is probable that the variants<br />

have edaphic preferences but speculation on this matter has still to be confirmed. Some of these variants<br />

may warrant recognition; Turner (1970) has recorded chromosome counts of n =12 and n = 14 for the<br />

species, however, the voucher material (PERTH) for the counts appears to be morphologically identical.<br />

Erymophyllum Paul G. Wilson<br />

Erymophyllum Paul G. Wilson, <strong>Nuytsia</strong> 7:105(1989).<br />

Type: Erymophyllum gracile (A. Gray) Paul G. Wilson.<br />

Pteropogon sect. Helipteroides A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:269(1852). Lectotype:<br />

Pteropogon gracilis A. Gray (Wilson 1989b).<br />

For list of species see Wilson 1989b.


Paul G. Wilson, Helipterum: Part 1<br />

The Hyalosperma group<br />

Hyalosperma Steetz<br />

Hyalosperma Steetz in Lehm., Pl. Preiss. 1:476(1845). Lectotype: Hyalosperma glutinosum Steetz<br />

(Wilson 1989a).<br />

Helipterum sect. Pachypterum Steetz in Lehm., Pl. Preiss. 1:473(1845). Lectotype: Helipterum cotula<br />

(Benth.) DC. (Wilson 1989a).<br />

Pteropogon sect. Pteropogonopsis A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 4:269(1852). Type:<br />

Pteropogon demissus A. Gray.<br />

For description and list of species see Wilson 1989a.<br />

Gilberta Turcz.<br />

Gilberta Turcz., Bull. Soc. Nat. Mosc. 24/1:192 (pre 27 March 1851). Type: Gilberta tenuifolia Turcz.<br />

Antheidosorus A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 3:98 (April 1851). Type: Antheidosorus gracilis<br />

A. Gray.<br />

Erect annual, moderately arachnoid. Leaves alternate, filiform. Capitula heterogamous, terminal to<br />

short lateral branches, nutant in fruit. Involucre turbinate to hemispherical, c. 4 mm high, radiant; outer<br />

bracts very broadly ovate, hyaline, slightly woolly at base; intermediate bracts with a broad hyaline<br />

slightly woolly claw and a short spreading yellow limb; innermost bracts obovate, hyaline, pale yellow,<br />

not radiating; stereome of ray bracts short and triangular. Receptacle flat, covered with scarious narrowoblong<br />

receptacular bracts of which c. 5 surround each floret. Florets actinomorphic, the outer row<br />

bisexual and the remainder male. Bisexual florets: Corolla glabrous, the vascular strands terminating at<br />

apex of throat; tube narrowly cylindrical; throat very short and turbinate; lobes 5, oblong, acute,<br />

prominent, smooth within, the cells narrow-oblong with straight walls. Anthers exserted; terminal<br />

appendage ovate, c. 0.2 mm long, cells narrow-oblong with thin walls; tails weak and filamentous. Style<br />

apex narrow-ovate to deltoid, densely papillose, the vascular strand slender and extending to base of<br />

apex. Achene compressed obovoid, c. 1.5 mm long, minutely colliculate; carpopodium minute (scarcely<br />

apparent); pericarp thick and transparent, strongly myxogenic; testa thinly coriaceous, brown, without<br />

crystals, vascular strands not apparent. Sterile achenes linear. Pappus of 5 unequal linear-acuminate<br />

shortly plumose bristles of which one or two have clavate terminal cilia; pappus of fertile florets weak<br />

and caducous as a whole, of male florets indurated at base and persistent on achene.<br />

Gilberta tenuifolia Turcz., op.cit.193. Type: Western Australia, J. Gilbert 277 (holo: KW).<br />

Antheidosorus gracilis A. Gray, Hooker’s J. Bot. Kew Gard. Misc. 3:174 (June 1851). - Myriocephalus<br />

gracilis (A.Gray) Benth., Fl. Austral. 3:559(1867). Type: Swan river Colony, J. Drummond, n.v.<br />

419


420 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Helipterum verecundum S. Moore, J. Linn. Soc. 34:200(1899). Type citation: 'Juxta Coolgardie<br />

repperi mens. Aug. florentem’, S. Moore, n.v.<br />

Asa Gray (1851) suggested that, due to the presence of receptacular bracts, this species had affinities<br />

to Myriocephalus. However, as has been pointed out by Short et al. (1989), Gilberta has little in common<br />

with M. appendiculatus Benth., the type of Myriocephalus. The florets and achenes of Gilberta<br />

tenuifolia are, in fact, very similar to those of Hyalosperma, particularly to H. glutinosum (Steetz)<br />

Paul G. Wilson. Both of these species have a similar corolla with narrow-oblong cells in the inner<br />

epidermis of the lobes, almost identical anthers and style apices, a similar pappus, and very similar<br />

achenes in which the pericarp forms a pellucid myxogenic layer around the seed. The leaves and<br />

indumentum of Gilberta are also of the same form as those found in Hyalosperma. Gilberta differs most<br />

obviously in the arrangement of the capitula and in the presence of capitular bracts around each floret;<br />

it also has glabrous corolla lobes whereas in H. glutinosum the corolla lobes have hairs that possess<br />

a large ovoid terminal gland.<br />

According to Warcup (1990) both Gilberta and Hyalosperma form an ectomycorrhizal association<br />

while four species currently placed in Myriocephalus that have been tested do not.<br />

The two names, Gilberta and Antheidosorus, were evidently published within a few days of each<br />

other. A copy of the Bull. Soc. Imp. Naturalistes Moscou 24/1 was lodged with the Censor on 17 January<br />

1851 (Old Style Calendar) and presented to the Society on 15 March 1851 (O.S.) which is equivalent to<br />

27 March 1851 (New Style). I have taken the date on which it was presented to the Society as being the<br />

date of publication (see Stafleu 1969, Marchant 1989). According to Stafleu and Cowan (1979), p.98 of<br />

Hooker’s J. Bot. Kew Gard. Misc. vol. 3 was published in April 1851 and p.174 in June 1851.<br />

The spelling Gilberta (and not Gilbertia) was deliberately used by Turczaninow in order that it could<br />

be better distinguished from Gilibertia Ruiz & Pav., a genus in the Araliaceae.<br />

The Triptilodiscus group<br />

Triptilodiscus Turcz.<br />

Triptilodiscus Turcz., Bull. Soc. Imp. Naturalistes Moscou 24/2:66(Aug.-Nov. 1851). Type: T. pygmaeus<br />

Turcz.<br />

Dimorpholepis A. Gray in Hooker, Icon. Pl. 9:t.856(Sept.-Nov.1851); A. Gray, Hooker’s J. Bot. Kew Gard.<br />

Misc. 4:227(Aug.1852). Type: D. australis A. Gray.<br />

Duttonia F. Muell., Linnaea 25:409(1853). Type: D. sessiliceps F. Muell.<br />

Annual herb with several major axes, sparsely pilose with slender hairs. Leaves alternate linear<br />

to narrow-oblong. Capitula heterogamous, terminating branches and subtended by c. 5 leaves of<br />

which one or more may bear a shoot that continues growth to give a monochasial conflorescence.<br />

Involucre subglobular c. 6 mm high, not radiant. Outer bracts scarious, hyaline, narrow-elliptic, ±<br />

equal to head, long-ciliate with linear stereome. Inner bracts narrow-oblong, slightly exceeding outer


Paul G. Wilson, Helipterum: Part 1<br />

bracts, cartilaginous and prominently thickened with narrow scarious shortly ciliate margin and<br />

scarious ciliate apex, somewhat boat-shaped, hispidulous abaxially. Receptacle broad-conical, smooth,<br />

glabrous. Florets discoid, a few outer female, remainder bisexual. Female florets: corolla very narrowtubular<br />

2-2.5 mm long, narrow-urceolate at apex, eventually curved outwards, very shortly 3-dentate,<br />

minutely and sparsely glandular puberulous with uniseriate hairs; inner cells of urceolate portion with<br />

sinuate margins, lobes densely papillose within; pappus a short corona of barbellate scales, sometimes<br />

with one shortly plumose bristle. Bisexual florets: corolla narrow-tubular, urceolate above, minutely 4dentate,<br />

c. 2.5 mm long, at first erect eventually curved outwards, otherwise as in female florets; antherappendage<br />

oblong, very thin, cells narrow-oblong, unthickened; anther tails filamentous, shorter than<br />

collar; style apex truncate; achene narrow-cylindrical, c. 1.8 mm long, minutely puberulous with 2-celled<br />

tooth-like hairs rounded at apex; pericarp diaphanous; testa leathery, smooth, without crystals; boss<br />

a minute narrow ring; pappus bristles 3-4, linear-acuminate, shortly plumose, c. 2.5 mm long, persistent,<br />

Triptilodiscus pygmaeus Turcz., l.c. - Helipterum pygmaeum (Turcz.) Druce in Hayward & Druce,<br />

Advent. Fl. Tweedside 103(1919) comb. illeg. Type: Western Australia, J. Drummond 5th coll. n.54 (iso:<br />

MEL 109204).<br />

Dimorpholepis australis A. Gray, l.c. - Helipterum dimorpholepis Benth., Fl. Austral. 3:650(1867) nom.<br />

illeg. - Argyrocome dimorpholepis Kuntze, Revis. Gen. Pl. 1:309(1891). - Helipterum australe<br />

(A. Gray) Druce, Bot. Exch. Club Brit. Isles 1916, 4:627(1917). Type citation: 'Hab. South-western<br />

Australia, Drummond. Also in the interior of Eastern Australia, at Bathurst Plains, Fraser; and Nangers,<br />

Captain M'Arthur.’ Syntype: J. Drummond 54 (K, photo seen).<br />

Duttonia sessiliceps F. Muell., Linnaea 25:410(1853). Type citation: 'In pratis prope rivum Rocky-creek.’<br />

Lectotype (here chosen): Rocky Creek, Oct. 1851, F. Mueller (lecto: MEL 109152; isolecto: MEL 109197).<br />

There is uncertainty over the publication dates of Triptilodiscus Turcz. and Dimorpholepis A. Gray.<br />

The former name was published in the Bull. Soc. Imp. Naturalistes Moscou vol.24 part 2 no. 3 which was<br />

sent to the Censor on 15 August 1851 and presented to the Society on 5 November 1851 (New Style<br />

calendar). The latter name was published as tab. 856 of the Icones Plantarum which appeared between<br />

September and November 1851 or possibly later, fide C. Jeffrey in litt. I have adopted the Turczaninow<br />

name, as did Laurie Haegi (1986), since there is the greater likelihood of it having priority.<br />

This genus is closely related to Haegiela which shares the same branching habit of the conflorescence,<br />

as well as the same hair type, and similar corolla and achene characters. Haegiela differs principally in<br />

the absence of a pappus, the presence of crystals in the testa, and the presence of unthickened inner<br />

involucral bracts.<br />

Haegiela P. Short & Paul G. Wilson<br />

Haegiela P. Short & Paul G. Wilson, Muelleria 7:259(1990). Type: Haegiela tatei (F. Muell.) P. Short &<br />

Paul G. Wilson<br />

For description and discussion see Short & Wilson, op.cit.<br />

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422 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Pterochaeta Steetz<br />

Pterochaeta Steetz in Lehm., Pl. Preiss. 1:456(1845). Type: Pterochaeta paniculata Steetz<br />

Annual woolly herb with one to several major axes. Leaves alternate, oblong. Capitula<br />

homogamous, shortly pedunculate, racemosely arranged, subtended by two leafy bracts. Involucre<br />

broad urceolate; bracts multiseriate, long ciliate, woolly, pale yellow; outer bracts ovate, scarious,<br />

attenuate at base with a minute hard green stereome, shortly apiculate with a pale fawn lamina;<br />

intermediate bracts broad-ovate scarious with a narrow-oblong hard claw (stereome) and a short<br />

oblong lamina; innermost bracts very short on a narrow-oblong hard naviculiform claw; receptacle<br />

naked; florets c. 20, outer female, inner bisexual, actinomorphic. Corolla cylindrical, shortly 4-lobed<br />

in female, 5-lobed in bisexual, glabrous; lobes glabrous within, thickened on margin; cells of throat<br />

undulate. Anther c. 0.7 mm long, delicate, appendage short, thin; tails filamentous. Style apex<br />

truncate. Achene narrow-ellipsoid, shortly beaked, covered with 2-celled tooth-like papillae;<br />

pericarp thin and weak, transparent, myxogenic, vascular strands laterally placed in relation to the<br />

cotyledons; testa leathery, brown rugulose, cells very thick walled and deeply undulate with scattered<br />

narrow-oblong crystals, vascular strand laterally placed and confined to near base of seed; carpopodium<br />

minute, annular. Pappus equal to corolla; bristles plumose, persistent.<br />

One species endemic to southern Western Australia.<br />

Pterochaeta paniculata Steetz in Lehm., op.cit. 455. - Waitzia paniculata (Steetz) Benth.,<br />

Fl. Austral. 3:637(1867). Lectotype (here chosen): In limoso-arenosis planitiei haud longe a praedio<br />

rustico "Maddington", et sinu regis Georgii III., Nov.[18]39 et [18]40. L. Preiss. No.35. (lecto: MEL<br />

1585199; isolecto: LD).<br />

The genus Waitzia differs from Pterochaeta most obviously in habit, in the nature of the anther<br />

apiculum and anther tails, in having a stout deltoid style apex with a vascular strand extending to the<br />

tip, and in the form of the achene. The vascular strands of the pericarp in Waitzia are medially placed<br />

in relation to the cotyledons whereas in Pterochaeta they are laterally placed.<br />

Haptotrichion Paul G. Wilson, gen. nov.<br />

Herba annua erecta pilis septatis glandulosis pubescens. Folia caulina alterna linearia. Capitula<br />

homogama, solitaria radiantia. Involucrum hemisphericum; bracteae c. 4-seriatae; bracteae exteriores<br />

elliptica hyalina longe ciliata, stereomate anguste oblongo plano; bracteae intimae ungue anguste<br />

elliptico longe ciliato, stereomate anguste oblongo, lamina elliptica flava. Receptaculum minute<br />

glanduloso papillosum. Flosculi numerosi, bisexuales, actinomorphi. Corolla tubo gracili glanduloso<br />

piloso, limbo campanulato glabro; lobi 5, ovati, nervis ad apicem extensis. Antherae: appendix ovatus<br />

tenuis, cellulis marginalibus manifeste discretis, cellulis basalibus ± equilateralibus, cetera anguste<br />

oblongis; caudae filamentosae collum superantes. Styli apex truncatus. Achenium angusto<br />

ellipticum compressum, ad apicem in rostro productum, papillis 2-cellulis dentoideis ornatis;<br />

pericarpium: stratum exterius cellulis cubicis parietibus tenuibus; stratum interius cellulis incrassatis,<br />

lignosis, sclerenchymatis; testa tenuissima, nervo circumnexo. Pappus persistens, setis filiformibus,<br />

denticulatis, basi versus in tubo conjunctis.<br />

Typus: Haptotrichion conicum (B. Turner) Paul G. Wilson


Paul G. Wilson, Helipterum: Part 1<br />

Annual erect herbs pubescent with gland-tipped septate hairs. Leaves cauline, alternate, linear.<br />

Capitula solitary and terminal to long branches, radiant. Involucre hemispherical; bracts c. 4-seriate;<br />

outer bracts elliptic, hyaline, long-ciliate with a brown narrow-oblong flat stereome; innermost bracts<br />

with a narrow-elliptic hyaline long-ciliate claw, a narrow-oblong stereome, and a yellow elliptic<br />

lamina. Receptacle minutely glandular papillose. Florets numerous, bisexual, actinomorphic.<br />

Corolla: tube slender, glandular pilose; limb campanulate, glabrous; lobes five, ovate, vascular<br />

strands extending to tips. Anther; appendage ovate, thin, marginal cells differentiated, basal cells ±<br />

equilateral, medial and distal cells narrow-oblong; tails slender delicate, extending to beyond collar.<br />

Style apex truncate; vascular strand prominent, extending almost to tip. Achene narrow-elliptic,<br />

compressed, extending into a terminal beak, minutely papillose with 2-celled tooth-like trichomes;<br />

carpopodium annular, short; pericarp: outer layer of 1-cell thick layer of thin-walled cubical cells;<br />

inner layer of 2-3 rows of thick walled sclerenchymatous cells; testa extremely thin, more or less<br />

adherent to pericarp; endosperm thin, free from testa; vascular strands of pericarp lateral (in relation<br />

to cotyledons); vascular strand of testa passing almost completely around seed in lateral position.<br />

Pappus bristles filiform, barbellate, united below to form a cup-shaped base, entire or slit on one side,<br />

persistent.<br />

Haptotrichion conicum had been placed in Waitzia since it possessed beaked achenes. It differs<br />

from Waitzia most obviously in 1) having a different type of indumentum (not cottony), 2) having broad<br />

flat hyaline claws to the inner involucral bracts (not terete and firm), 3) in having truncate style apices<br />

(neither ellipsoid nor deltoid), and 4) in having delicate filamentous anther tails (not slender and firm).<br />

Haptotrichion further differs from Waitzia in the anatomy of the achene for its pericarp has an outer<br />

single-celled layer of thin-walled cube like cells and an inner layer of several rows of thick-walled<br />

sclerenchymatous cells, while the testa is extremely thin and delicate. In Waitzia the pericarp consists<br />

of a thin layer of hyaline linear cells while the testa is thick, rugose, and made up of thick-walled<br />

collenchyma. The vascular strand in the testa of Haptotrichion passes almost completely around the<br />

seed but in Waitzia it is confined to the base.<br />

From Pterochaeta this genus differs in the manner of branching of the stem and inflorescence (q.v.),<br />

in the nature of the involucral bracts, the nature of the corolla, and the nature of the anther apiculum.<br />

In Pterochaeta, Haptotrichion and Waitzia the achenes are beaked and bear tooth-like 2-celled<br />

trichomes but in other characters the genera are very distinct.<br />

The genus consists of two species, both endemic to the Carnarvon District (Beard 1980) of Western<br />

Australia.<br />

Haptotrichion colwillii Paul G. Wilson, sp. nov. (Figure 6)<br />

Herba erecta ad 25 cm alta. Rami glanduloso puberuli sub capitula pilosi. Folia linearia, 8-15 mm longa,<br />

glanduloso puberula. Involucrum hemisphaericum, c. 10 mm altum et latum; bracteis exterioribus late<br />

ellipticis, hyalinis, ciliatis; bracteis intimis ungue anguste elliptico, c. 9 mm longo, hyalino, ciliato, limbo<br />

anguste obovato obtuso, 9-15 mm longo, luteo. Corolla c. 5 mm longa, tubo glanduloso hirtello, limbo<br />

anguste turbinato, glabro, intra sub lobis papilloso. Achenium anguste ellipsoideum c. 2.5 mm longum,<br />

denticulatum; rostrum gracile, laeve, ad 3 mm longum. Pappus persistens, setis in basi connatis,<br />

tubiformibus.<br />

Typus: 7 km S of Overlander Roadhouse [c. 170 km S of Carnarvon], Western Australia, 1 Sept. 1985,<br />

H. Demarz 10702 (holo: PERTH).<br />

423


424 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Figure 6. Haptotrichion colwillii. A - Habit. B - Florets. C - Pappus bristle apices. D - Hair of corolla. E - Style apex.<br />

F - Outer, intermediate, and inner involucral bracts. G - Anther. H - Achene with pappus. I - Achene. J - Achene papillae.<br />

K - Capitulum. L & M - Hairs of stem. N - Leaf. From H. Demarz 10702.


Paul G. Wilson, Helipterum: Part 1<br />

Erect herb to 25 cm high, branching at base. Principal axes slender, to 20 cm high, simple or<br />

sparsely branched, glandular puberulous, pilose beneath capitulum. Leaves linear, 8-15 mm long,<br />

c. 0.7 mm wide, glandular puberulous, margins slightly recurved. Involucre hemispherical, c. 10 mm<br />

high and wide. Outer bracts broad-elliptic, hyaline, ciliate; stereome narrow-triangular, brown,<br />

glandular puberulous. Innermost bracts: claw narrow-elliptic, c. 9 mm long, hyaline, ciliate, stereome<br />

very narrow-oblong, glabrous except for the glandular puberulous apex; limb narrow-obovate, obtuse,<br />

9-15 mm long, yellow. Corolla c. 5 mm long; tube narrow, glandular hirtellous with multicelled hairs;<br />

limb narrow-turbinate, glabrous, shortly 5-lobed, papillose within below junction of lobes. Achene<br />

compressed narrow-ellipsoid, c. 2.5 mm long, barbellate with 2-celled papillae; beak slender, smooth,<br />

to 3 mm long. Pappus more or less equal to corolla, persistent, bristles united towards base to form<br />

a tube c. 2 mm long.<br />

Additional specimen examined. WESTERN AUSTRALIA: Hamelin Pool, 16 Sept. 1984, J. Colwill s.n.<br />

(PERTH).<br />

Habitat. Found on red sand over limestone (fide H. Demarz in sched.).<br />

Distribution. Only known from near Hamelin Pool, c. 170 km south of Carnarvon, Western Australia.<br />

Notes. Haptotrichion colwillii is very similar to H. conicum which is known from an area between<br />

Carnarvon and Gascoyne Junction. The latter species differs most obviously in 1) the leaves ending<br />

in a rounded 'capitate’ tip, 2) the receptacle being narrow-conical (not rounded), and 3) the pappus sheath<br />

being split along one side.<br />

Haptotrichion colwillii is named after John Colwill, a Western Australian horticulturalist who has<br />

a particular interest in the native Asteraceae and who first collected this species.<br />

The illustration is drawn from a plant raised from seed collected by Herbert Demarz and grown at Kings<br />

Park Botanic Garden, Perth. I should like to thank the staff of Kings Park for their unstinted assistance<br />

in my study of this and other annual species of the Rhodanthe complex.<br />

Haptotrichion conicum (B. Turner) Paul G. Wilson, comb. nov.<br />

Waitzia conica B. Turner, Sida 2:428(1966). Type: 11 miles west of Gascoyne Junction, 24 Aug. 1965,<br />

B.L. Turner 5405 (holo: MEL 598263; iso: MEL 598265, PERTH).<br />

[Waitzia podolepis auct. non (Gaud.) Benth.: F. Muell., Zeitschrift des allgem. österreich. Apotheker-<br />

Vereines 34 (no.36):933-936 (1896); F. Muell., Pl. Indig. Sharks Bay 16(1883).]<br />

Distribution. Between Carnarvon and Gascoyne Junction, Western Australia.<br />

A description of this species was provided by Mueller (1883) based on a collection made by<br />

J. Polak (or Pollack) in 1882 from the Gascoyne River region (MEL 1584941). Mueller assumed that<br />

the plant he was describing was Waitzia podolepis (Gaudich.) Benth.: 'this plant represents evidently<br />

the genuine species, illustrated by Gaudichaud’, even though Mueller had seen no authentic material<br />

of that species. In 1896 Mueller again included the Pollack specimen under W. podolepis but without<br />

comment. In 1905 Diels and Pritzel stated that the Pollack collection was not W. podolepis, however,<br />

they did not give it a name.<br />

425


426 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

Acknowledgements<br />

I thank the various Australian herbaria who sent on loan much of their material of Helipterum for<br />

study. Kings Park Botanic Garden allowed me to have access to their live collection and provided me<br />

with seed of a number of species. Steve Hopper kindly searched for and collected from different<br />

populations of rare Rhodanthe species near Shark Bay. Photographs of relevant Asteraceae types in<br />

herb. KW were made available to me by my colleague Neville Marchant. Discussions and correspondence<br />

with Philip Short and Arne Anderberg have been stimulating and of considerable assistance. The<br />

drawings were prepared with much care by Margaret Menadue.<br />

References<br />

Anderberg, A.A. (1989). Phylogeny and reclassification of the tribe Inuleae (Asteraceae). Canadian J. Bot. 67:2277-<br />

2296.<br />

Anderberg, A.A. (1991). Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Botanica 104:1-195.<br />

Beard, J.S. (1980). A new phytogeographic map of Western Australia. Western Australian Herb. Res. Notes No.3:37-<br />

58.<br />

Bentham, G. (1867). "Flora Australiensis." vol. 3. (Lovell Reeve, London.)<br />

Candolle, A.P. de (1838). 'Prodromus systematis naturalis regni vegetabilis.’ Vol. 6. (Treuttel & Wurtz, Paris.)<br />

Erickson, R. (1969). "The Drummonds of Hawthornden." (Lamb Patterson, Perth.)<br />

Gray, A. (1851). Characters of some Gnaphalioid Compositae of the Division Angiantheae. Hooker’s J.Bot.Kew<br />

Gard.Misc. 3:172-178.<br />

Haegi, L. (1986). Helichrysum Miller. In Jessop, J.P. & Toelken, H.R. "Fora of South Australia." (South Australian<br />

Government, Adelaide.)<br />

Hilliard, O.M. (1983). "Flora of Southern Africa." Vol. 33 Asteraceae part 7 Inuleae fasc. 2 Gnaphaliinae. 1-325.<br />

(Department of Agriculture, Pretoria.)<br />

Hilliard, O.M. & Burtt, B.L. (1981). Some generic concepts in Compositae-Gnaphaliinae. J. Linn. Soc., Bot. 82:181-<br />

232.<br />

Hnatiuk, R.J. (1990). Census of Australian vascular plants. (Bureau of Flora and Fauna, Canberra.)<br />

Marchant, N.G. (1990). The contribution of the Russian botanist Turczaninov to Australian plant taxonomy. In P.S.<br />

Short (ed), "History of systematic botany in Australasica."(Australian Systematic Botany Society Inc., Melbourne.)<br />

Nordenstam, B. (1989). A synopsis of the genus Syncarpha (Compositae-Gnaphalieae). Compositae Newsletter No<br />

17:2-6.<br />

Short, P.S. (1983). A revision of Angianthus Wendl., sensu lato (Compositae: Inuleae: Gnaphaliinae), 1. Muelleria<br />

5:143-183 (1983).<br />

Short, P.S. (1989). A revision of Podotheca Cass. (Asteraceae: Inuleae: Gnaphaliinae). Muelleria 7:39-56.<br />

Short, P.S. & Sinkora, D.M. (1988). The botanist Joachim Steetz (1804-1862). Muelleria 6:449-494.<br />

Short, P.S., Wilson, K.E. & Nailon, J. (1989). Notes on the fruit anatomy of Australian members of the Inuleae<br />

(Compositae). Muelleria 7:57-79.<br />

Stafleu, F.A. (1969). Turczaninow’s Flora Baicalensi Dahurica. Taxon 18:563-565.<br />

Stafleu, F.A. & Cowan, R.S. (1979). "Taxonomic literature, a selective guide to botanical publications and collections<br />

with dates, commentaries and types." Vol. 2. (Bonn, Scheltema & Holkema, Utrecht.)<br />

Turner, B.L. (1970). Chromosome numbers in the Compositae. XII Australian species. Amer. J. Bot. 57:382-389.<br />

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

Wilson, P.G. (1989a). A revision of the genus Hyalosperma (Asteraceae: Inuleae: Gnaphaliinae). <strong>Nuytsia</strong> 7:75-101.<br />

Wilson, P.G. (1989b). Erymophyllum (Asteraceae: Inuleae: Gnaphaliinae), a new Australian genus in the Helipterum<br />

complex. <strong>Nuytsia</strong> 7:103-116.<br />

Wilson, P.G. (1992a). The classification of Australian species currently included in Helipterum (Asteraceae:<br />

Gnaphalieae): Part 2 Leucochrysum. <strong>Nuytsia</strong> 8:439-445.<br />

Wilson, P.G. (1992b). The classification of the genus Waitzia Wendl. (Asteraceae: Gnaphalieae). <strong>Nuytsia</strong> 8:461-477.<br />

Wilson, P.G., Short, P.S., & Orchard, A.E. (1992). Some nomenclatural changes in the Angianthinae and Cassiniinae<br />

(Asteraceae: Gnaphalieae). Muelleria 7:519-524.


Paul G. Wilson, Helipterum: Part 1<br />

Index to Scientific Names<br />

New names are given bold type. Other accepted names are in roman. Synonyms are in italics.<br />

Acroclinium A. Gray 386<br />

multicaule A. Gray 387<br />

phyllocephalum A. Gray 394<br />

roseum A. Gray 387<br />

var. album G. Nicholson 387<br />

var. grandiflorum G. Nicholson 387<br />

rubellum A. Gray 398<br />

Anisolepis Steetz 415<br />

pyrethrum Steetz 416<br />

Antheidosorus A. Gray 419<br />

gracilis A. Gray 419<br />

Argyrocome Gaertn.<br />

charsleyae (F. Muell.) Kuntze 413<br />

chlorocephala (Turcz.) Kuntze 386<br />

condensata (F. Muell.) Kuntze 401<br />

corymbiflora (Schldl.) Kuntze 391<br />

corymbosa (A. Gray) Kuntze 401<br />

dimorpholepis (Benth.) Kuntze 421<br />

floribunda (DC.) Kuntze 408<br />

forrestii (F. Muell.) Kuntze 401<br />

frenchii (F. Muell.) Kuntze 415<br />

fuscescens (Turcz.) Kuntze 394<br />

haigii (F. Muell.) Kuntze 401<br />

heterantha (Turcz.) Kuntze 412<br />

humboldtiana (Gaudich.) Kuntze 396<br />

laevis (A. Gray) Kuntze 402<br />

manglesii (Lindley) Kuntze 384<br />

margarethae (F. Muell.) Kuntze 415<br />

moschata (DC.) Kuntze 385<br />

polycephala (A. Gray) Kuntze 404<br />

polygalifolia (DC.) Kuntze 397<br />

polyphylla (F. Muell.) Kuntze 417<br />

427


428 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

pygmaea (DC.) Kuntze 398<br />

pyrethrum (Steetz) Kuntze 416<br />

rosea (Hook.) Kuntze 387<br />

rubella (A. Gray) Kuntze 398<br />

spicata (F. Muell.) Kuntze 413<br />

sterilescens (F. Muell.) Kuntze 409<br />

stricta (Lindley) Kuntze 399<br />

tietkensii (F. Muell.) Kuntze 406<br />

Calocephalus<br />

globosus Hutch. 414<br />

gnaphalioides Hook. 385<br />

Cassinia<br />

cuprea F. Muell. 399, 404<br />

Cephalipterum A. Gray 417<br />

drummondii A. Gray 417<br />

f. major Diels & E. Pritzel 418<br />

f. minor Diels & E. Pritzel 418<br />

Dimorpholepis A. Gray 420<br />

australis A. Gray 421<br />

Duttonia F. Muell. 420<br />

sessiliceps F. Muell. 421<br />

Erymophyllum Paul G. Wilson 418<br />

gracile (A. Gray) Paul G. Wilson 418<br />

Epaltes Cass.<br />

tatei F. Muell. 421<br />

Gilberta Turcz. 419<br />

tenuifolia Turcz. 419<br />

Gnaphalium<br />

moschatum DC. 385<br />

Griffithia J. Black 386<br />

helipteroides J. Black 397<br />

Haegiela P. Short & Paul G. Wilson 421<br />

tatei (F. Muell.) P. Short & Paul G. Wilson 421


Paul G. Wilson, Helipterum: Part 1<br />

Haptotrichion Paul G. Wilson 422<br />

colwillii Paul G. Wilson 423, 424<br />

conicum (B. Turner) Paul G. Wilson 425<br />

Helichrysum Miller<br />

sect. Acroclinium (A. Gray) Baillon 386<br />

sect. Rhodanthe (Lindley) Baillon 383<br />

anthemoides Sprengel 386<br />

drummondii F. Muell. 387<br />

grayi F. Muell. 387<br />

humboldtianum Gaudich. 396<br />

manglesii (Lindley) Baillon 384<br />

obtusifolium Sonder 381<br />

oldfieldii F. Muell. 407<br />

Helipterum DC.<br />

sect. Aglossum F. Muell. 412<br />

sect. Helipteridium A. Gray 412<br />

sect. Leiochrysum DC. 385<br />

sect. Monencyanthes (A. Gray) Benth. 384<br />

sect. Pachypterum Steetz 419<br />

sect. Pteropogon (DC.) Benth. 385<br />

sect. Sericophorum DC. 385, 408<br />

sect. Synachyrum A. Gray 408<br />

adpressum W. Fitzg. 436<br />

albicans (Cunn.) DC. 436<br />

album (Nicholson) L. Bailey 387<br />

album Ewart 401<br />

anactinum F. Muell. 412<br />

anthemoides (Sprengel) DC. 386<br />

australe (A. Gray) Druce 421<br />

battii F. Muell. 413<br />

charsleyae F. Muell. 413<br />

chionolepis F. Muell. 409<br />

chlorocephalum (Turcz.) Benth. 386<br />

cirratum Morrison 409<br />

429


430 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

clementii Domin 396<br />

condensatum F. Muell. 401<br />

corymbiflorum Schldl. 391<br />

var microglossa Benth. 396<br />

corymbosum (A. Gray) Benth. 401<br />

cotula (Benth.) DC. 419<br />

craspedioides W.V. Fitzg. 436<br />

cryptanthum O. Sarg. 384<br />

diffusum DC. 394<br />

dimorpholepis Benth. 421<br />

discoideum A. Gray 412<br />

drummondii (A. Gray) Ostenf. 398<br />

fitzgibbonii F. Muell. 436<br />

floribundum DC. 408. 409<br />

var. stuartianum (Sond.) Benth. 411<br />

var. tubulipappum J. Black 409<br />

forrestii F. Muell. 401<br />

frenchii F. Muell. 415<br />

fuscescens Turcz. 394<br />

haigii F. Muell. 401<br />

heteranthum Turcz. 412<br />

var. minor Ewart 412<br />

var. majus Benth. 412<br />

humboldtianum (Gaudich.) DC. 396<br />

laeve (A. Gray) Benth. 402<br />

largiflorens F. Muell. 396<br />

manglesii (Lindley) Benth. 384<br />

var. maculatum (J. Drumm. ex hort.) L.H. Bailey 384<br />

margarethae F. Muell. 415<br />

margaritae F. Muell. 415<br />

maryonii S. Moore 385<br />

microglossa (Benth.) Tate 396<br />

microglossum Maiden & E. Betche 396<br />

molle (DC.) Paul G. Wilson 437


Paul G. Wilson, Helipterum: Part 1<br />

monencyanthoides F. Muell. 414<br />

moschatum (DC.) Benth. 385<br />

mullinense S. Moore 401<br />

niveum Steetz 437<br />

oppositifolium S. Moore 397<br />

pachychaetum W. Fitzg. 412<br />

polycephalum (A. Gray) Benth. 404<br />

polygalifolium DC. 397<br />

var. leucactinum F. Muell. 394<br />

polyphyllum F. Muell. 417<br />

propinquum W. Fitzg. 398<br />

pterochaetum (F. Muell.) Benth. 437<br />

punctatum DC. 386<br />

pygmaeum (DC.) F. Muell. 398<br />

var. drummondii (A. Gray) Ostenf. 398<br />

var. occidentale Benth. 398<br />

pygmaeum (Turcz.) Druce 421<br />

pyrethrum (Steetz) Benth. 416<br />

roseum (Hook.) Benth. 387<br />

f. album (Nicholson) Moldenke 387<br />

var. album Guilfoyle 387<br />

var. album Ewart et al. 388<br />

var. grandiflorum (Nicholson) Chitt. 387<br />

var. nigropapposum Ostenf. 387<br />

var. patens (Ewart et al.) J. Black 388, 411<br />

rubellum (A. Gray) Benth. 398<br />

sandfordii Hook. 396<br />

spicatum (Steetz) Benth. 413<br />

var. pallens Benth. 413<br />

splendidum Hemsley 388<br />

sterilescens F. Muell. 409<br />

stipitatum (F. Muell.) Benth. 438<br />

strictum (Lindley) Benth. 399<br />

var. stenocephala S. Moore 399<br />

431


432 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

stuartianum Sond. 411<br />

tietkensii F. Muell. 406<br />

troedelii F. Muell. 411<br />

var patens Ewart et al. 388, 411<br />

uniflorum J. Black 385<br />

verecundum S. Moore 420<br />

Hyalosperma Steetz 419<br />

glutinosum Steetz 419<br />

Leptorhynchos Less.<br />

citrinus Benth. 407<br />

Monencyanthes A. Gray 384<br />

gnaphalioides (Hook.) A. Gray 384, 385<br />

Myriocephalus Benth.<br />

gracilis (A. Gray) Benth. 419<br />

Podosperma Labill.<br />

fuscescens (Turcz.) F. Muell. 394<br />

pollackii F. Muell. 413<br />

Podotheca Cass.<br />

fuscescens (Turcz.) Benth. 394<br />

pollackii (F. Muell.) Diels 413<br />

Pterochaeta Steetz 422<br />

paniculata Steetz 422<br />

Pteropogon DC. 385<br />

sect. Achyroclinoides A. Gray 399<br />

sect. Facelioides A. Gray 385<br />

sect. Helichrysoides A. Gray 412<br />

sect. Helipteroides A, Gray 418<br />

sect. Pteropogonopsis A. Gray 419<br />

australis Nees 398<br />

corymbosus A. Gray 401<br />

demissus A. Gray 419<br />

drummondii A. Gray 398<br />

humboldtianus (Gaudich.) F. Muell. 396


Paul G. Wilson, Helipterum: Part 1<br />

intermedius F. Muell. 398<br />

laevis A. Gray 402<br />

platyphyllus F. Muell. 399<br />

polycephalus A. Gray 414<br />

pygmaeum DC. 398<br />

spicatus Steetz 413<br />

Rhodanthe Lindley 383<br />

sect. Achyroclinoides (A. Gray) Paul G. Wilson 399<br />

sect. Actinaria Paul G. Wilson 414<br />

sect. Anisolepis (Steetz) Paul G. Wilson 415<br />

sect. Citrinae Paul G. Wilson 406<br />

sect. Helichrysoides (A. Gray) Paul G. Wilson 412<br />

sect. Helipteridium (A. Gray) Paul G. Wilson 412<br />

sect. Leiochrysum (DC.) Paul G. Wilson 385<br />

sect. Monencyanthes (A. Gray) Paul G. Wilson 384<br />

sect. Polyphyllum Paul G. Wilson 416<br />

sect. Rhodanthe 383<br />

sect. Synachyrum (A. Gray) Paul G. Wilson 408<br />

anthemoides (Sprengel) Paul G. Wilson 386<br />

ascendens Paul G. Wilson 400<br />

atrosanguinea J. Drumm. ex hort. 384<br />

battii (F. Muell.) Paul G. Wilson 413<br />

charsleyae (F. Muell.) Paul G. Wilson 413<br />

chlorocephala (Turcz.) Paul G. Wilson 386<br />

subsp. chlorocephala 387<br />

subsp. rosea (Hook.) Paul G.Wilson 387<br />

subsp. splendida (Hemsley) Paul G. Wilson 388<br />

citrina (Benth.) Paul G. Wilson 407<br />

collina Paul G. Wilson 389, 390<br />

condensata (F. Muell.) Paul G. Wilson 401<br />

corymbiflora (Schldl.) Paul G. Wilson 391<br />

corymbosa (A. Gray) Paul G. Wilson 401<br />

cremea Paul G. Wilson 391, 393<br />

433


434 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

diffusa (DC.) Paul G. Wilson 394<br />

var. diffusa 394<br />

var. leucactina (F. Muell.) Paul G Wilson 394<br />

floribunda (DC.) Paul G. Wilson 408<br />

forrestii (F. Muell.) Paul G. Wilson 401<br />

frenchii (F. Muell.) Paul G. Wilson 415<br />

fuscescens (Turcz.) Paul G. Wilson 394<br />

gossypina Paul G. Wilson 395<br />

haigii (F. Muell.) Paul G. Wilson 401<br />

heterantha (Turcz.) Paul G. Wilson 412<br />

humboldtiana (Gaudich.) Paul G. Wilson 396<br />

laevis (A. Gray) Paul G. Wilson 402<br />

maculata J. Drumm. ex hort. 384<br />

manglesii Lindley 384<br />

var. alba Regel 384<br />

subvar. alba hort ex Vilm. 384<br />

var. maculata (J. Drumm. ex hort.) Vilm. 384<br />

var. maculata Hook. 384<br />

var. sanguinea Hook. 384<br />

margarethae (F. Muell.) Paul G. Wilson 415<br />

maryonii (S. Moore) Paul G. Wilson 385<br />

microglossa (Maiden & E. Betche) Paul G. Wilson 396<br />

moschata (DC.) Paul G. Wilson 385<br />

nullarborensis Paul G. Wilson 402, 403<br />

oppositifolia (S. Moore) Paul G. Wilson 397<br />

subsp. oppositifolia 397<br />

subsp. ornata Paul G.. Wilson 379<br />

pollackii (F. Muell.) Paul G. Wilson 413<br />

polycephala (A. Gray) Paul G. Wilson 404<br />

polygalifolia (DC.) Paul G. Wilson 397<br />

polyphylla (F. Muell.) Paul G. Wilson 417<br />

propinqua (W. Fitzg.) Paul G. Wilson 398<br />

psammophila Paul G. Wilson 404, 405<br />

pygmaea (DC.) Paul G. Wilson 398


Paul G. Wilson, Helipterum: Part 1<br />

pyrethrum (Steetz) Paul G. Wilson 416<br />

rubella (A. Gray) Paul G. Wilson 398<br />

rufescens Paul G. Wilson 398<br />

sphaerocephala Paul G. Wilson 409<br />

spicata (Steetz) Paul G. Wilson 413<br />

sterilescens (F. Muell.) Paul G. Wilson 409<br />

stricta (Lindley) Paul G. Wilson 399<br />

stuartiana (Sond.) Paul G. Wilson 411<br />

tietkensii (F. Muell.) Paul G. Wilson 406<br />

troedelii (F. Muell.) Paul G. Wilson 411<br />

uniflora (J. Black) Paul G. Wilson 385<br />

Roccardia Voss 383<br />

anthemoides (Sprengel) Voss 386<br />

chlorocephala (Turcz.) Voss 387<br />

corymbiflora (Schldl.) Voss 391<br />

humboldtiana (Gaudich.) Voss 396<br />

manglesii (Lindley) Voss 384<br />

f. alba Voss 384<br />

f. ligulosa Voss 384<br />

f. maculata Voss 384<br />

f. sanguinea Voss 384<br />

rosea (Hook.) Voss 387<br />

rubella (A. Gray) Voss 398<br />

Schoenia Steetz<br />

chlorocephala Turcz. 386<br />

humboldtiana (Gaudich.) Walp. 396<br />

Triptilodiscus Turcz. 420<br />

pygmaeus Turcz. 421<br />

Waitzia Wendl.<br />

brevirostris Steetz 407<br />

citrina (Benth.) Steetz 407<br />

conica B. Turner 425<br />

dasycarpa Turcz. 407<br />

paniculata (Steetz) Benth. 422<br />

435


436 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

steetziana Lehm. 407<br />

sulphurea Steetz 407<br />

tenella Hook. 407<br />

Xyridanthe Lindley 386<br />

stricta Lindley 399<br />

Name changes in Australian species of Helipterum (see also Wilson 1989a,b and 1992a,b)<br />

Taxa listed in Hnatiuk (1990) Taxa accepted by Wilson<br />

H. adpressum Chrysocephalum puteale (S. Moore) Wilson<br />

H. albicans Leucochrysum albicans (A.Cunn.) Wilson<br />

H. albicans subsp. albicans Leucochrysum albicans subsp. albicans<br />

H. albicans var. buffaloensis Leucochrysum albicans var. buffaloensis (Wilson)<br />

Wilson<br />

H. albicans var. incanum Leucochrysum albicans subsp. albicans var. tricolor<br />

(DC.) Wilson<br />

H. albicans subsp. alpinum Leucochrysum albicans subsp. alpinum (F. Muell.)<br />

Wilson<br />

H. albicans var. graminifolium Leucochrysum graminifolium (Wilson) Wilson<br />

H. anthemoides Rhodanthe (Leiochrysum) anthemoides (Sprengel)<br />

Wilson<br />

H. australe Triptilodiscus pygmaeus Turcz.<br />

H. battii Rhodanthe (Helichrysoides) battii (F. Muell.) Wilson<br />

H. charsleyae Rhodanthe (Helichrysoides) charsleyae (F. Muell.)<br />

Wilson<br />

H. chlorocephalum Rhodanthe (Leiochrysum) chlorocephala (Turcz.)<br />

Wilson subsp. chlorocephala<br />

H. condensatum Rhodanthe (Achyroclinoides) condensata (F. Muell.)<br />

Wilson<br />

H. corymbiflorum Rhodanthe (Leiochrysum) corymbiflora (Schldl.) Wilson<br />

H. corymbosum Rhodanthe (Achyroclinoides) corymbosa (A. Gray) Wilson<br />

H. cotula Hyalosperma cotula (Benth.) Wilson<br />

H. craspedioides Possibly referrable to Polycalymma<br />

H. demissum Hyalosperma demissum (A. Gray) Wilson<br />

H. diffusum Rhodanthe (Leiochrysum) diffusa (Cunn. ex DC.) Wilson<br />

var. diffusa<br />

H. fitzgibbonii Leucochrysum fitzgibbonii (F. Muell.) Wilson


Paul G. Wilson, Helipterum: Part 1<br />

H. floribundum Rhodanthe (Synachyrum) floribunda (DC.) Wilson<br />

H. forrestii Rhodanthe (Achyroclinoides) forrestii (F. Muell.) Wilson<br />

H. frenchii Rhodanthe (Actinaria) frenchii (F. Muell.) Wilson<br />

H. fuscescens Rhodanthe (Leiochrysum) fuscescens (Turcz.) Wilson<br />

H. gracile Erymophyllum tenellum (Turcz.) Wilson<br />

H. haigii Rhodanthe (Achyroclinoides) haigii (F. Muell.) Wilson<br />

H. heteranthum Rhodanthe (Helipteridium) heterantha (Turcz.) Wilson<br />

H. humboldtianum Rhodanthe (Leiochrysum) humboldtiana (Gaudich.) Wilson<br />

H. hyalospermum Hyalosperma glutinosum Steetz subsp. glutinosum<br />

H. involucratum Erymophyllum ramosum (A. Gray) Wilson subsp.<br />

involucratum (F. Muell.) Wilson<br />

H. jessenii Hyalosperma semisterile (F. Muell.) Wilson<br />

H. laeve Rhodanthe (Achyroclinoides) laevis (A. Gray) Wilson<br />

H. manglesii Rhodanthe (Rhodanthe) manglesii Lindley<br />

H. margarethae Rhodanthe (Actinaria) margarethae (F. Muell.) Wilson<br />

H. maryonii Rhodanthe (Monencyanthes) maryonii (S. Moore) Wilson<br />

H. microglossum Rhodanthe (Leiochrysum) microglossa (Maiden & Betche)<br />

Wilson<br />

H. molle Leucochrysum molle (Cunn. ex DC.) Wilson<br />

H. moschatum Rhodanthe (Monencyanthes) moschata (Cunn. ex DC.)<br />

Wilson<br />

H. niveum Distinct genus; affinities with Helichrysum obtusifolium<br />

Sond.<br />

H. oppositifolium Rhodanthe (Leiochrysum) oppositifolia (S. Moore) Wilson<br />

H. polycephalum Rhodanthe (Achyroclinoides) polycephala (A. Gray)<br />

Wilson<br />

H. polygalifolium Rhodanthe (Leiochrysum) polygalifolia (Cunn. ex DC.)<br />

Wilson<br />

H. polyphyllum Rhodanthe (Polyphyllum) polyphylla (F. Muell.) Wilson<br />

H. praecox Hyalosperma praecox (F. Muell.) Wilson<br />

H. propinquum Rhodanthe ((Leiochrysum) propinqua (W. Fitzg.) Wilson<br />

H. pterochaetum Chrysocephalum pterochaetum F. Muell.<br />

H. pygmaeum Rhodanthe (Leiochrysum) pygmaea (DC.) Wilson<br />

H. pyrethrum Rhodanthe (Anisolepis) pyrethrum (Steetz) Wilson<br />

H. roseum Rhodanthe (Leiochrysum) chlorocephala subsp. rosea<br />

(Hook.) Wilson<br />

437


438 <strong>Nuytsia</strong> Vol. 8, No. 3 (1992)<br />

H. rubellum Rhodanthe (Leiochrysum) rubella (A. Gray) Wilson<br />

H. saxatile Distinct genus<br />

H. semisterile Hyalosperma semisterile (F. Muell.) Wilson<br />

H. spicatum Rhodanthe (Helichrysoides) spicata (Steetz) Wilson<br />

H. splendidum Rhodanthe (Leiochrysum) chlorocephala subsp. splendida<br />

(Hemsley) Wilson<br />

H. sterilescens Rhodanthe (Synachyrum) sterilescens (F. Muell.) Wilson<br />

H. stipitatum Leucochrysum stipitatum (F. Muell.) Wilson<br />

H. stoveae Hyalosperma stoveae (D.A. Cooke) Wilson<br />

H. strictum Rhodanthe (Leiochrysum) stricta (Lindley) Wilson<br />

H. stuartianum Rhodanthe (Synachyrum) stuartiana (Sond.) Wilson<br />

H. tenellum Erymophyllum tenellum (Turcz.) Wilson<br />

H. tietkensii Rhodanthe (Achyroclinoides) tietkensii (F. Muell.) Wilson<br />

H. troedelii Rhodanthe (Synachyrum) troedelii (F. Muell.) Wilson<br />

H. uniflorum Rhodanthe (Monencyanthes) uniflora (J. Black) Wilson<br />

H. venustum Hyalosperma glutinosum Steetz subsp. venustum<br />

(S.Moore) Wilson<br />

H. zacchaeus Hyalosperma zacchaeus (S. Moore) Wilson

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