Muelleria

Volume 7
Number 1
April 1989

NATIONAL HERBARIUM OF VICTORIA if
LANDS & FORESTS DIVISION ^

DEPARTMENT OF CONSERVATION, FORESTS AND LANDS

Muelleria

Volume 7, Number 1

April 1989

CONTENTS

Volume 7, Number 1

Page

Two new lichens; Cladonia paeminosa and C. humilis var. bourgeanica

— Alan W. Archer

Eucalyptus cadens (Myrtaceae), a new swamp gum from the Warby Range,
North-East Victoria

— J. D. Briggs and M. D. Crisp

Azorella Lamarck (Apiaceae) on Heard and Macquarie Islands, with
description of a new species

— A. E. Orchard

Notes on Hovea R. Br. (Fabaceae): 2

— J. H. Ross

A revision of Podotheca Cass. (Asteraceae: Gnaphaliinae)

— P. S. Short

Notes on the fruit anatomy of Australian members of the Inuleae
(Asteraceae)

— P. S. Short, K. E. Wilson and J. Nailon

Two new species of Pomaderris Labill. (Rhamnaceae) from South-Eastern
Australia

— Neville G. Walsh •. •••••

A new subspecies of Grevilleu ucunthifolici (Proteaceae: Grevilleoideae) from
South-Eastern New South Wales

— R. O. Markinson and D. E. Albrecht

A new species of Tetranhena R.Br. (Poaceae) from Victoria and New South
Wales

— Neville G. Walsh

Ferdinand Mueller, Government Botanist: the role of William Hooker m his
appointment

— Helen M. Cohn

New genera and species of Australian Inuleae (Asteraceae)

— P. S. Short ;••••••

OlcGfici ciTchcTi (Asteraceae I Astereae), a new name for a familiar species irom
Tasmania

— N. S. Lander

Olearia astroloba (Asteraceae: Astereae), a new species endemic to Victoria

— N. S. Lander and N. G. Walsh ...........

Valid publication of the names Torrenticola and T. queenslandica
(Podostemaceae)

— Helen I. Aston

Acacia cacfulcsccns, a new species of Acacia section Phyllodincac trom

Victoria

— B. R. Maslin and A. B. Court

Notes on Hovea R. Br. (Fabaceae); 3

— J. H. Ross

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15

21

39

57

81

89

95

99

103

117

123

127

131

135

Editor; D. B. Foreman

Published by the National Herbarium of Victoria (MEL).
Royal Botanic Gardens, South Yarra, Victoria 3141, Australia.

The date of distribution of Volume 6, number 6 was 29 March 1988.

TWO NEW LICHENS: CLADONIA PAEMINOSA AND C. HUMILIS VAR.

BOURGEANICA

by

Alan W. Archer*

ABSTRACT

Archer, Alan W. Two new lichens: Cladonia paeminosa and C. humilis var.
bourgeanica. Muelleria (1): 1-5 (1989). — Cladonia paeminosa A. W. Archer and
Cladonia humilis (With.) Laundon var. bourgeanica A. W. Archer are described as
new. Both taxa contain fumarprotocetraric and bourgeanic acids and occur in
Australia. C. humilis var. bourgeanica also occurs in Europe and North and South
America.

INTRODUCTION

A recent examination of undetermined specimens in the lichen genus Cladonia
(Ascomycetes, Lecanorales) in Herbarium collections from South Australia (AD) and
Tasmania (HO) revealed specimens that were not referrable to any known taxa;
similar specimens had been collected by the author in Victoria, New South Wales and
the Australian Capital Territory. These specimens which resemble Cladonia
scabriuscula (Del. in Duby) Nyl. and contain fumarprotocetraric and bourgeanic acids
are here described as a new species.

A recent typification (Laundon 1984) of the lichens described by William
Withering in the eighteenth centu^ showed that the name Cladonia conista C. Robb,
ex Allen was illegitimate as the epithet conista belonged to the synonymy of Cladonia
humilis (With.) Laundon. The name ‘C. conista' has referred to a sorediate, scyphose
taxon containing fumarprotocetraric and bourgeanic acids which Laundon (loc. cit.)
referred to as an undescribed variety of Cladonia humilis. This variety is here formally
named and described.

METHOD

The lichen compounds present in the specimens examined were identified by
thin-layer chromatography of acetone extracts, using the mobile phases described by
Culberson (Culberson 1972) and the separated compounds were detected with
sulphuric acid (Culberson 1972) and MBTH (Archer 1978). The presence of
bourgeanic acid (substance H) was confirmed with the micro-crystal test described by
Thomson (Thomson 1967) and by mass spectrometry (c/ Bodo et al. 1973).

TAXONOMY

Cladonia paeminosa A. W. Archer, sp. nov.

Sicut Cladonia scabriuscula sed podetiis subsimplicibus, basibus podetiorum ecorticatis et acidum
bourgeanicum continens.

Primary squamules small, inconspicuous, persistent, 1 x 2 mm, subdigitately
lobed, margins smooth, green above, white below. Podetia arising from the squamules,
15-50(-100) mm tall, 0-5-1 -5 mm diam., escyphose, simple or dichotomously
branched, axils open, tapering towards the apices, the apices simple or bifurcating,
subulate or with terminal pycnidia; ecorticate, or corticate at the base and becoming
ecorticate, with tiny corticate patches scattered along the podetia, minutely
squamulose, esorediate. Apothecia pale brown, terminal, convex, (0-2-) 0-5- 1-0 mm
diam. Ascospores eight per ascus, colourless, simple, ellipsoid, 12-14 pm long, 3-4 pm

•Division of Analytical Laboratories, Department of Health, P.O. Box 162, Lidcombe, New South Wales,
Australia 2141.

1

2

Fig. la. Cladonia paeminosa. Holotype. Scale in millimetres.

Fig. lb. Cladonia paeminosa. Holotype. Close-up of podetia. Bar= 1 millimetre.

wide. Pycnidia terminal, black, subconical, 0-3 mm x 0-3 mm; conidia not seen.
Phallus K— , KC— , Pd+ red; containing fumarprotocetraric and bourgeanic acids
(Figs 1-2).

Typus: Australia, Victoria, near Mirimbah, by side of Mt Stirling Road, c. 30 km E. of
Mansfield, 146° 25' E., 37° 07' S., alt. c. 700 m, 9.xi.l986, A. Archer 2027
(Holotypus; MEL 1050876. Isotypi: ANUC, CBG, H, NSW).

Further Specimens Examined:

South Australia — 14 km SE. of Mt Burr Township, 7.ix.l966, 1.B. Wilson 516 (AD 9255); Marshes
Swamp, 24.vii.1980, N. Donner 6997 (AD 4862); ibidem, 23.x. 1982, N. Donner 9577 (AD 9235).

New South Wales — 4 km S. of Termed, 7.iii.l970, E. Dahl (CANB 227416); Black Flat, Currowan
State Forest, 15 km NW. of Batemans Bay, 150° 04' E., 35° 36' S., alt. c. 200 m, 28. v. 1983, .4. Archer 2113
(NSW).

Australian Capital Territory — Eishing Gap Trail, 33 km SW. of Canberra, 148° 52' E., 36° 29' S., alt.
900 m, l.iv.l983, A. Archer 2098 (MEL 1050870, NSW).

Victoria — on earth near sea, Mentone, 15.v. 1886, F.R.M. Wilson (NSW); on mossy earth,
Cunningham, iii. 1890, F.R.M. Wilson (NSW); Narbethong, Healesville, viii. 1906, Mrs Goodyear {NSAN)\ by
side of Howqua River, near Sheepyard Elat, c. 30 km SE. of Mansfield, 1 46° 20' E., 37° 1 2' S., alt. c. 500 m,
3.ix.l986, /I. Archer 2069 (NWh 1050874, NSW).

Tasmania — St Mary’s Pass, G. Bratt 73/646 (HO 54009); Prosser River Gorge, J. Gilbert, s.n. (HO
60742); Drip Palls, M. Westbrook, s.n. (HO 50617).

Discussion

The specific epithet 'paeminosa'' refers to the rough surface of the podetia. C.
paeminosa (Fig. la) is morphologically similar to C scabriuscula (Del. in Duby) Nyl.
but differs from that species in often lacking a well-defined corticate area at the base
and on the lower part of the podetia. C. paeminosa has small irregular corticate patches
along the podetia (Fig. lb) and contains the aliphatic acid bourgeanic acid which is
lacking in C. scabriuscula. Both species contain the depsidone fumarprotocetraric
acid. C. scabriuscula has been reported to contain the depside atranorin (Culberson

3

Fig. 2. Cladonia humilis var. bourgeanica. Holotype. Scale in millimetres.

1970) or the triterpene ursolic acid (Thornson 1968), in additiori to fuinar-
protocetraric acid. Specimens with atranorin are common in Chile (Ahti &
Kashiwadani 1984) but only one Australian specimen (AD 4868) was found with
atranorin. Ursolic acid was not found in any of the Australian specimens of C.
scdbfiuscula examined. C. scubriusculci often occurs in large clumps with the much
branched podetia dying at the base; in contrast, C. paeminosa is much less branched
and often remains attached to the primary thallus on the soil on which it grows.
Apothecia were seen in only one specimen of C. paeminosa, from South Australia. C.
scabriuscula is a cosmopolitan species, widely distributed in Australia and occurs in all
states whereas C. paeminosa is so far known only from south-eastern Australia and
Tasmania. C. paeminosa belongs to the group Cladonia sub-group Furcatae in the
recently proposed subgeneric classification of the genus Cla(^onia (Huovinen & Ahti
1982).

Cladonia humilis (With.) Laundon var. bourgeanica A. W. Archer, var. nov.

Cladonia conista auct.

Sicut Cladonia humilis var. humilis sed acidum bourgeanicum continens vice atranorinum.

Primary squamules persistent, conspicuous, 1-2 mmx 1-5 mm, rounded, lobed,
margins slightly crenate, green above, white below, esorediate. Podetia growing from
the upper surface of the primary squamules, simple, scyphose, lacking marginal or
central proliferations, 5-20 mm tall, scyphi 3-6 mm diam., the podetial stalk
corticate, the cortex areolate at the base, becoming sub-verrucose, rarely rninutely
squamulose at the base; the scyphi ecorticate and farinose sorediate, the interior

4

closed, ecorticate and farinose sorediate, margins entire or minutely denticulate,
becoming slightly revolute in older specimens. Apothecia and pycnidia not seen.
Thallus K— , KC— , Pd+ red; containing fumarprotocetraric and bourgeanic acids.

Typus: Australia, New South Wales, Six Foot Track, Binomea Ridge, 2 km N. of
Jenolan Caves, 150° 02' E., 33° 48' S., alt. 1 100 m, 13.iii.l987, A. Archer 2086.
Holotypus: MEL 1050873; Isotypus: NSW.

Further Specimens Examined:

Australian Capital Territory — Kambah Pool, 28. vi. 1 970, £■. Dahl (CANB 227965); near Honeysuckle
Creek, 20 km SSW. of Canberra, 148° 58' E., 35° 35' S., alt. 1 100 m, 2.iv. 1983,/!. Archer 2097 {fiS'fjy.hy side
of Two Sticks Rd, 35 km W. of Canberra, 148° 48' E., 35° 19' S., alt. 1 100 m, 30.X.1985, A. Archer 1822A
(NSW); near Tidbinbilla River, c. 27 km SW. of Canberra, 148° 55' E., 35° 28' S., alt. 900 m, 21.iv.l986, .4.
Archer 1935 (NSW).

Victoria — by side of Mt Stirling Rd, Mirimbah, c. 30 km E. of Mansfield, 146° 25' E., 37° 07' S. alt.
700 m, 5.X1.1986, A. Archer 2005 (H, MEL 1050879); ibidem 9.xi.l986, A. Archer 2024 (MEL
1050878).

Tasmania — Table Mountain, G.C. Bratt 72/930 (HO 53023); Finger Post Track, A.M. Gray 36 (HO
69121); Mt Wellington, D. Ratkowsky L129 (BM, NSW).

Discussion:

The combination Cladonia conista (Ach.) C. Robb, ex Allen (Allen 1930) was
based on Cenomyce fimbriata var. conista Ach.; a specimen collected by Fldrke was
chosen as lectotype (Ahti 1 966). The basionym of the epithet was later reported (Ahti
1980) to be superfluous and hence illegitimate and the combination C. conista (Nyl.)
C. Robb, ex Allen was proposed, based on C. fimbriata f. conista Nyl. This com-
bination was rejected as the basionym was published without a description (Laundon
1 984), thus leaving the bourgeanic acid containing variety of C. humilis lacking a valid
name. The epithet "bourgeanica’ is here proposed for this chemical variety.

C. humilis var. bourgeanica (Fig. 2) is distinguished chemically from var. humilis
by the presence of the fatty acid bourgeanic acid in place of the depside atranorin. Both
varieties are distinguished from the somewhat morphologically similar C. fimbriata
(L.) Fries by the presence of bourgeanic acid or atranorin and the corticate podetial
stalk, absent in C. fimbriata. Two specimens of C. humilis, containing both atranorin
and bourgeanic acid with fumarprotocetraric acid, have been reported, one from Chile
(Ahti & Kashiwadani 1984) and a second specimen from Argentina (T. Ahti, in litt.,
1987).

C. humilis var. bourgeanica is a widely distributed taxon and occurs in Europe,
North America and South America. In Australia it has been found only in
south-eastern Australia and Tasmania, in contrast to var. humilis which occurs in all
States except Queensland. C. humilis var. bourgeanica has also been reported, as C.
conista, from New Zealand (Martin 1958); the specimen on which this report was
based [Mt Cargill, Dunedin, Otago, W. Martin 4414 (CHR 385665)] was found to
contain only fumarprotocetraric acid and is identified as C chlorophaea (Floerke ex
Sommerf.) Sprengel.

C. humilis var. bourgeanica belongs to the infra-generic group Cladonia subgroup
C. grayi and allies (Huovinen & Ahti 1982).

ACKNOWLEDGEMENTS

The author is grateful to Professor T. Ahti, Helsinki, for much helpful discussion
and to the National Herbarium of New South Wales for arranging the loan of
specimens, and acknowledgement is made to the Director, Division of Analytical
Laboratories, for permission to publish this paper.

REFERENCES

Ahti, T. ( 1 966). Correlation of the chemical and morphological characters in Cladonia chlorophaea and allied
lichens. Annls. hot. fenn. 3: 380-390.

Ahti, T. (1980). Noraenclatural notes on Cladonia species. Lichenologist 12: 125-133.

5

Ahti, T. & Kashiwadani, H. (1984). The lichen genera Cladia, Cladina and Cladonia in southern Chile. In
Inoue, H. (ed.), ‘Studies on Cryptogams in Southern Chile’ pp. 125-151. (Kenseisha Ltd: Tokyo.)

Allen, A. F. (1930). Cladoniae from the valley of the Cap Chat, Gaspe Peninsula. Rhodora 32: 91-114.

Archer, A. W. (1978). 3-Methyl-2-benzothiazolone hydrazone hydrochloride as a spray reagent for phenolic
lichen compounds. J. Chromatogr. 152: 290-292.

Bodo, B., Hebrard, P., Mohlo, L. & Mohlo, D. (1973). Un nouvel acide aliphatique des lichens Desmaziera
evernioides et Ramalina bourgeanica. Tetrahedron Letters 18: 1631-1634.

Culberson, C. F. (1970). ‘Chemical and Botanical Guide to Lichen Products’. (University ofNorth Carolina:
Chapel Hill, N.C.)

Culberson, C. F. (1972). Improved conditions and new data for the identification of lichen products by a
standardised thin-layer chromatographic method. J. Chromatogr. 12: 113-125.

Huovinen, K. & Ahti, T. (1982). Biosequential patterns for the formation of depsides, depsidones and
dibenzofurans in the genus Cladonia (lichen forming ascomycetes). Annls. hot. fenn. 19: 225-234.

Laundon, J. R. (1984). The typification of Withering’s neglected lichens. Lichenologist 16: 21 1-239.

Martin, W. (1958). The Cladoniaceae of New Zealand. Trans. R. Soc. N.Z. 85: 603-632.

Thomson, J. W. (1967). ‘The Lichen Genus Cladonia in North America’. (University of Toronto:
Toronto.)

Manuscript received 11th September 1987.

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EUCALYPTUS CADENS (MYRTACEAE), A NEW SWAMP GUM FROM THE
WARBY RANGE, NORTH-EAST VICTORIA

by

J. D. Briggs' and M. D. Crisp^

ABSTRACT

Briggs, J. D. and Crisp, M. D. Eucalyptus cadens (Myrtaceae), a new swamp gum from
the Warby Range, North-East Victoria. Muelleria 1 (1): 7-13 (1989). — A new
eucalypt, known only from a single locality in the Warby Range in NE. Victoria, is
described and named. It is related to the Swamp Gums (the informal E. subser.
Ovatinae), particularly E. aggregata Deane & Maiden, but is readily distinguished by
its glaucous new growth. Often mature trees lean or fall over but continue to thrive in
their boggy habitat which surrounds permanent springs.

INTRODUCTION

The first record of this species appears to be a collection made in 1 979 by Mr A. C
Beauglehole and Miss C. D. Nason on the late Miss Nason’s property in the Warby
Range. The same collectors obtained more material in 1985, and subsequently the
population was listed by Beauglehole (1986) under the name E. yarraensis Maiden &
Cambage. Miss Nason had also mentioned the existence of this unusual eucalypt to Mr
J. L. Briggs, a local beekeeper, who recently visited the site and sent specimens to one
of us (J. D. Briggs, his nephew) for identification. As the specimens did not match any
species known to us, we undertook further investigations including field studies,
collection of additional specimens and the growing of seedlings. The distinctiveness of
the population combined with a lack of segregation in the seedlings convinced us that
this was a new species. In this paper, we describe and name the new eucalypt and
discuss its affinities and conservation status.

TAXONOMY

Eucalyptus cadens J. Briggs & Crisp, sp. nov.

E. aggregata Deane & Maiden arte simulans sed surculis manifeste glaucis, foliis maturis
glaucescentibus, cortice pro parte maxima laevi praeter in parte inferna trunci differt.

Holotypus: Victoria, eastern foot of the Warby Range, between Wangaratta and
Glenrowan, J. D. Briggs 2068 & J. L. Briggs, 8.x. 1986, 2 sheets (CANB 370885-6).
IsoTYPi: CBG, HO, MEL, NSW.

Spreading tree, often leaning, or fallen and continuing to grow from existing
shoots as well as producing new vertical stems from the old trunk; standing trees
8-25 m tall, to 1 m d.b.h.; forming a lignotuber; bark smooth, decorticating in short
ribbons, greenish grey above a 1-10 m hard, dark-grey, finely furrowed stocking; oil
glands abundant in bark; new shoots and leaves glaucous, weathering to grey-green at
maturity; crowns frequently containing mostly intermediate leaves. Cotyledons
bilobed. ^Seedling leaves decussate to 5th-7th node, subsessile, spreading, elliptic to
oblong, mostly narrow, slightly concave above, obtuse or rounded at apex, abruptly
tapered at base, 10-50x3-22 mm, grey-green; leaves (above 12th node)

similar but not opposite, ascending, narrow-elliptic to linear, ± acute, more tapered at
base, glaucescent; seedling stems terete, red. Intermediate leaves petiolate (1-5 mm),
narrow-elliptic to elliptic, ±flat, acute or obtuse, to 200 x65 mm, conspicuously
glandular; late intermediate leaves narrow-ovate to elliptie, to 1 50 x 34 mm including
petiole to 20 mm. Adult leaves mostly pendulous, narrow-elliptic, slightly falcate,
tapering to both ends (often abruptly to apex), 70- 1 1 5 x 9-20 mm including 5- 1 4 mm

1 CSIRO, Division of Plant Industry, P.O. Box 1600, Canberra, A.C.T., Australia 2601.

2 Australian National Botanic Gardens, P.O. Box 1777, Canberra, A.C.T., Australia 2601.

7

8

petiole, with abundant oil glands; midrib prominent, venation obscure, intramarginal
vein c. 0-75 mm from leaf margin. Unit inflorescences one per axil, 7-flowered;
peduncles terete to slightly angular, 4-6 mm long, c. 1 mm thick. Buds broad-fusiform,
up to 8x3 mm including 0-5-2 mm pedicel; hypanthium obconical, c. 3x3 mm,
slightly contracted at base; inner operculum conical, slightly beaked, about equal in
length and fractionally narrower than hypanthium; outer opercululm shed early and
intact, leaving scar. Stamens: filaments inflexed in bud, white; anthers versatile,
obovoid, opening in vertical slits, c. 0-6 mm long. Ovary with 3 or 4 locules; ovules in 4
longitudinal rows on lower part of placenta; style clavate, 2-5-3 mm long; stigma blunt
with a lobed surface, papillae short. Fruit densely clustered and appearing sessile, very
broadly obconical, slightly contracted at base, 3-5-4-5 x 4-6-6 mm; pedicel 0-1-5 mm
long; disc slightly raised, narrow (c. 0-5 mm wide); valves opening widely, exserted,

0- 5-1 mm, rarely level with rim. Seed irregularly ellipsoid, depressed, smooth-edged,

1- 0-1 -7 mm long, 0-6-1 -0 mm broad, c. 0-3 mm thick, shallowly reticulate,
dullish-black, hilum ventral. (Figs 1-2).

Flowering Period:

Late March to early May.

Etymology:

The specific epithetic is from the Latin verb cadere, to fall down, and refers to the
tendency of mature trees to lean or fall over in their boggy habitat.

Distribution:

Known only from a single locality in the Warby Range of North-East Victoria
where two stands, one of about 550 trees and the other of 45 trees, survive on 5 hectares
of private property around the margins of two permanent springs.

Other Specimens Examined:

Victoria — Eastern foot of the Warby Range about 6 km N. of Glenrowan, 9.xi. 1 985, A. C. Beauglehole
81544 & CD. Nason (MEL); iv.l986, J.L. Briggs I, 2, 3& CD. Nason (CANB, CBG, MEL); 8.X.1986, J.D.
Briggs 2065 & J.L. Briggs (CMiB, CBG, MEL, NSW); 8.x. 1986, J.D. Briggs 2069 <6 J.L. Briggs (AD, CANB,
CBG, HO, MEL, MELU, NSW); 8.x. 1 986, J.D. Briggs 2070 & J.L. Briggs (CANB, CBG); 30.xii. 1 986, J.D.
Briggs 2166 & J.L. Briggs (AD, CANB, CBG, HO, MEL, MELU, NSW); 30.xii. 1986, J.D. Briggs 2168 & J.L.
Briggs (CANB, CBG, HO, MEL, NSW); J. Chappill 493, 494, 495, and seedlings grown from these
(MELU).

Habitat:

The population of E. cadens is at the eastern foot of a steep hill on the very gently
sloping floor of an open valley. There, it grows in seasonally waterlogged and
permanently moist conditions in an area surrounding perennial springs. The soil is a
grey sandy-clay loam overlying white gravelly clay. Eucalyptus cadens forms an almost
pure stand in an open-forest/woodland community, where over half of the trees either
have a significant lean or have fallen completely but continue to grow. A few
specimens of E. blakelyi Maiden are scattered through the stand and about 5 trees of a
form of E. camphora R. Baker occur in one edge of the larger stand. The understorey is
predominantly a dense sedge and grass sward dominated by Carex appressa R. Br. and
Juncus sp. (probably J. sarophorus L. Johnson), but with scattered shrubs of Acacia
melanoxylon R. Br., Leptospermum juniperinum Smith and Viminaria juncea (Schrad
& Wendl.) Hoffsgg. also present.

The slopes surrounding the site support E. blakelyi - E. bridgesiana R. Baker -
E. polyanthemos Schauer - E. albens Benth. - E. macrorhyncha F. Muell. - E. mellio-
dora Cunn. ex Schauer woodland and the adjacent flat areas have been heavily cleared
and converted to pasture.

Conservation Status:

Endangered, coded 2E (criteria from Leigh et al. 1981). Eucalyptus cadens is
known only from the one locality where about 600 trees occupy a total area of about
5 ha on two private properties. The area is subject to cattle grazing which largely

9

Fig. 1. Eucalyptus cadens. a — branchlet. b — cotyledons, c — bud. d — tangential longitudinal section of
bud, showing inflexion of stamens, e — anther, anterior view, f — anther, posterior view, g —
post-flowering hypanthium and style, h — unit infructescence. a,h from J. D. Briggs 2068 (the type);
c-g from J. L. Briggs 2\ b from a living seedling, grown from J. L. Briggs 2.

prevents seedling regeneration. The stand occupies potential agricultural land and
surrounds valuable permanent water supplies. There is a serious risk that much or all
of the species could be cleared when the properties come under new ownership. The
need for special protection of the species has become urgent in view of the recent death
of Miss Nason who had been keen to preserve the large portion of the population which
is on her land and whose family had owned the property since last century.

10

Fig. 2. Eucalyptus cadens. a — seedling, ex J. L. Briggs 2, showing gradation from elliptic seedling leaves to
the distinctly narrower juvenile leaves, b — typical habitat with a few standing trees and the bushy
crowns of several fallen specimens evident, c — a fallen tree continuing to thrive in its boggy habitat.
The paler glaucous new growth is a distinctive feature of this species, d — rough persistent bark forming
a stocking to 1 0 m at the base of the trunks, e — branches showing the smooth bark which decorticates
in ribbons.

11

No similar springs supporting stands of E. cadens were known to Miss Nason or
other long-term residents of the area. Whilst a few scattered individuals of E. cadens
may survive on private property along the edge of the Warby Range, substantial stands
are unlikely to be found in view of the very limited potential habitat and the extent to
which clearing has already occurred in the area.

Affinity:

Eucalyptus cadens clearly belongs to the informal E. sect. Maidenaria (Pryor &
Johnson 1971). The strongest supporting evidence is the presence of oil glands in the
bark, a feature which has been found to be largely confined to species in E. sect.
Maidenaria (Crisp 1988 and references therein). Additional support for this
placement comes from the bilobed cotyledons, more or less sessile juvenile leaves
(Boland et al. 1984) and general appearance of the species.

Within E. sect. Maidenaria, placement in the informal E. ser. Ovatae (Pryor &
Johnson 1 97 1) is suggested by the subsessile seedling leaves, as opposed to the strictly
sessile seedling leaves of E. ser. Viminales. Moreover, E. cadens bears a close
resemblance to the Swamp Gums (informal E. subser. Ovatinae Pryor & Johnson
1971), particularly with respect to the obconical fruit and swampy habitat.

Despite its close resemblance to the Swamp Gums, E. cadens is immediately
distinguished by its glaucous new growth, which weathers to a grey-green colour at
maturity. Under a dissecting microscope, the leaf cuticle has a dull appearance. None
of the Swamp Gums develop glaucescence, and under the microscope their cuticle
often appears somewhat lustrous. Nevertheless, there is little doubt that E. cadens
closely related to this group, and Table 1 presents some comparisons.

Eucalyptus cadens resembles E. aggregata more closely than any other species. At
all developmental stages except very young seedlings, both these species, together with
E. rodwayi, have narrower leaves than the other Swamp Gums (Table 1). They are
particularly close in bud and fruit morphology. Interestingly, the isolated Victorian
population of E. aggregata (at Woodend) shows the closest similarity to E. cadens,
particularly in bud and fruit shape. Despite this similarity, E. aggregata may be readily
distinguished from E. cadens by its shiny green leaves, persistence of rough bark to the
small branches and more gradually tapered adult leaves.

Most authors (e.g. Johnson 1962; Pryor and Johnson 1979; Simmons 1985)
regard Eucalyptus rodwayi R. Baker and H. G. Smith, a Tasmanian endemic, as being
closely related to E. aggregata. It is distinguished from E. cadens using similar
character differences to those exhibited by E. aggregata.

It has been suggested to us (J. Chappill, pers. comm.) that E. cadens may be closely
related to E. camphora R. Baker, especially to the type population near Rylstone, New
South Wales. In a current, uncompleted study, R. Coorey (pers. comm.) has recognised
four taxa within E. camphora. Two of these are represented in Table 1: the type
population, and the most southern form, which is the only form occurring in Victoria.
A few trees of the latter are growing at one end of the E. cadens population. All four
forms of E. camphora appear to be united as a natural group by the presence of
emarginate intermediate leaves. Neither E. cadens nor any other Swamp Gum species
shows this character (Table 1). All forms of E. camphora h5ve, on average, longer
pedicels than E. cadens (examples in Table 1). The Victorian form of E. camphora
differs further from E. cadens by its very broad somewhat lustrous leaves. In the type
population of E. camphora, the leaves are much narrower; nevertheless, they are
broader than and differently shaped from those of E. cadens, especially at the
intermediate stage (Table 1). We have examined Chappill’s adult and seedling
specimens (C/irzpp/7/ 493-5) and all except one segregate seedling (out of eighteen seen)
fall within our circumscription of the species. In particular, the late juvenile leaves
were typically narrow and there was no evidence of the broad, emarginate
intermediate leaves that characterise E. camphora.

Beauglehole (1986) identified the population of E. cadens as "E. yarraensis'.
However, the latter species does not appear to be so closely related. It differs in having
rough bark persistent to the small branches, glossy green leaves which are considerably

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13

broader at the intermediate stages, a more or less hemispherical hypanthium, and
usually longer pedicels (Table 1).

There is a history of confusion between the Swamp Gums and the closely related
E. aromaphloia Pryor & J. H. Willis (Simmons & Brown 1986). Recently, Chappill et
al. (1986) have demonstrated that E. aromaphloia, as previously circumscribed,
comprises four distinct taxa. However, the entire E. aromaphloia complex may be
readily distinguished from E. cadens by its rou^ bark which extends to the small
branches, manifestly attenuate adult leaf and hemispherical fruit with a strongly raised
disc.

ACKNOWLEDGEMENTS

The authors wish to thank Mr J. L. Briggs for collecting specimens and drawing
their attention to this new species and thank the late Miss Nason for her permission to
study this species on her property. We also thank our colleagues Jennifer Chappill,
Ken Hill, Lawrie Johnson and Lindsay P^or for their comments. Mr D. Fortescue is
thanked for his work in producing the line drawings of the new species. The work
relating to the study and description of E. cadens was supported by grants from the
Australian Heritage Commission and the Nell and Hermon Slade Trust.

REFERENCES

Beauglehole, A. C. ( 1986). The Distribution and Conservation of Vascular Plants in the Murray Valley Area,
Victoria. (E. Davis and Sons: Portland, Victoria.) „ ^ ^

Boland, D. J., Brooker, M. I. H., Chippendale, G. M., Hall, N., Hyland, B. P. M., Johnston, R. D., Kleinig, D.
A & Turner, J. D. (1984). Forest Trees of Australia 687 pp. (Nelson/CSIRO: Melbourne.)

Chappill, J., Ladiges, P. Y. & Boland, D. (1986). Eucalyptus aromaphloia Pryor & Willis — a redefinition of
geographical and morphological boundaries. Aust. J. Bot. 34: 395-412.

Crisp, M. D. (1988). Eucalyptus recurva (Myrtaceae), a new species from the Southern Tablelands of New
South Wales. Te/opea 3(2): 223-230. o u i. ms

Johnson, L. A. S. (1962). Studies in the Taxonomy of Eucalyptus. Contr. N.S.W. Nat. Herb. 3(3): 105.

Lei^, J., Briggs, J. & Hartley, W. (1981). Rare or Threatened Australian Plants. Australian National Parks
and Wildlife Service Special Publication No. 7, 178 pp. , ,, .

Pryor, L. D. & Johnson, L. A. S. (1971). A Classification of the Eucalypts. (Australian National University
Press* )

Simmons, D. (1985). Observations on Disjunct Populations of Eucalyptus aggregata in Victoria and New
South Wales with notes on the status of E. rodwayi. Victorian Nat. 103(4): 131-6.

Simmons, D. & Brown, G. (1986). The distribution of some Eucalyptus species (the Swamp Gums) in the
Yarra Valley, Victoria. Victorian Nat. 103: 19-25.

Manuscript received 6 January 1988

AZORELLA Lamarck (APIACEAE) ON HEARD AND MACQUARIE ISLANDS,
WITH DESCRIPTION OF A NEW SPECIES, A. MACQUARIENSIS

by

A. E. Orchard*

ABSTRACT

Orchard, A. E. Azorella Lamarck (Apiaceae) on Heard and Macquarie Islands, with
description of a new species, A. macquariensis. Muelleria 7(1): 15-20 (1989). —
Azorella macquariensis, a new species endemic to Macquarie Island has been
recognized. Hitherto it had been confused with A. selago which occurs on Heard Island
and several other subantarctic islands but not on Macquarie Island. A key, comparable
descriptions, and illustrations of the salient features are provided for the two species,
and some of the specimens examined are listed.

INTRODUCTION

Azorella is a genus of 33-70 (Allan 1961; Mathias & Constance 1971; Airy-Shaw
1973) species of perennial herbs, often mat or cushion-forming with their main centre
of speciation in the Andes of South America, but with a few taxa extending to the
Falkland and sub-Antarctic islands.

One of the most southerly, and the most widespread species is A. selago, described
by Hooker ( 1 845) from a number of specimens collected on the voyage of the ‘Erebus’
and ‘Terror’. It seems that he relied mainly on material from Kerguelen Island
collected by himself and Anderson in drawing up his description, but also had
available specimens collected by Darwin from Tierra del Fuego, by King from Port
Famine, and by himself from Hermit Island. At the end of the discussion he
mentioned, almost as an after-thought ‘I have also seen specimens sent from
McQuarrie’s Island by Mr. Frazer’. (The Hermit Island collection was subsequently
chosen as lectotype by Moore (1968).) Consequently, from the beginning this species
was perceived to be widespread and circumpolar in distribution. This opinion was
reinforced by subsequent reports of A. selago from Marion Island, Crozet Island and
Heard Island (summarised by Greene and Greene 1 963) and from Prince Edward and
McDonald islands (summarised by Greene & Walton 1975). Chastain (1958)
described polymorphism in A. selago on Kerguelen Island, adding to the character-
isation of the species as widespread and variable.

Recently, comparison of material from Heard and Macquarie Islands has
convinced me that two taxa are involved. While Heard Island specimens clearly match
Hooker’s original description and illustration, modern descriptions of Fuegian and
Falkland Island plants (Moore 1968, 1983) and specimens from Kerguelen Island, the
Macquarie Island plants can be distinguished by a number of characters, principally of
the leaves, but also of the inflorescence and fruit. This species is described below, along
with a comparable account of A. selago.

Key to the Species of Azorella on Heard and Macquarie Islands
1 . Leaf lamina 3(-5)-lobed, the lobes divided ± to the base, acute, pungent with a long
setose point; wings of the petiole reduced to a truncate ridge at the base of the
lamina; flowers solitary or sometimes paired; fruits up to 1-7 mm long, hidden in

upper leaves, sepals ± deciduous 1. A. macquariensis

1. Leaf lamina (4-) 5-6(-7)-lobed, the lobes divided only halfway to the base, blunt,
with at most a tiny mucro; wings of the petiole produced into auricles at the base of
the lamina; flowers usually in groups of 3; fruits 1-7-2-Omm long, at least
j-exserted from the leaves on pedicels 2 mm long, sepals persistent. 2. A. selago

‘Tasmanian Herbarium, G.P.O. Box 252C, Hobart, Tasmania, Australia 7001.

15

16

1 . Azorella macquariensis Orchard, sp. nov.

Azorella selago auctt. non J. Hooker (1846): J. Hooker, FI. Antarc. 2(1846)
284-5(quoad specimen Frazeris); Cheeseman, Vase. FI. Macq. Is. (1919) 26; Taylor,
FI. Veg. Soils Macq. Is. (1955) 128-30; Allan, FI. New Zeal. 1(1961) 451-2; Copson,
Atlas Vase. FI. Macq. Is. (1984) 37; and all other references to ‘A. selago' in so far as
Macquarie Island is concerned.

Illustrations: Taylor, FI. Veg. Soils Macq. Is. (1955) PI. 16, 17, 18, 29 (as A.
selago).

Herbae perennes pulvini formantes. Folia alterna arete imbricata; petioli 3-4(- 1 0) mm longi, late alati,
vaginantes, jugo incrassato truncate ad apicem; lamina 3(-5)-partita, lobis lanceolatis liberis fere basi
acutis acumine setose, glabra vel sparsim setosa in superficie. Flores solitarii, interdum binati,
hermaphroditi. Fructus cinnamomei, 1 -3- 1 -7 mm longi, inter folia supera occulta, stylis persistentibus,
sepalis ± deciduis. (Fig. 1).

Typus: Macquarie Island, Pyramid Lake, north side, in Azorella/Festuca/
Rhacomitrium crispulum fellfield, alt. 190 m, 4.xi. 1981, R. D. Seppelt 12039.
Holotypus: HO 67713.

Perennial herb forming extensive tight mats, cushions, or in exposed situations,
buttons; main branches prostrate, woody, to 5 mm in diameter; lateral shoots erect,
herbaceous, freely branched, crowded, 3-5(-l 5) cm tall clothed in the remains of old
leaves. Leaves alternate, closely imbricate and appressed to stems, persistent; petioles
white, 3-4(-10) mm long, thickened at the apex, 3(-5)-veined, with a membranous
wing outside the veins, the wings fused to form a sheath around the stem in the lower
y-y and produced above to form a very short truncate ridge-like ‘ligule’ at the base of
the lamina; lamina 3(-5)-partite, the lobes divided almost to the base in young leaves,
lanceolate, l-7-2-0(-4-0) mmlong,0-6-0-9(-l -4) mm wide, thick and subfleshy, acute
with a terminal setose apiculum, thickened margins, glabrous or with l-3(-5)
bristle-like hairs 1 -4-4-0 mm long on the adaxial surface. Flowers terminal, solitary or
sometimes paired, hermaphrodite, peduncles short with the flowers enclosed by the
upper leaves. Involucral bracts 2, fused at the base to form a small cup, leaf-like or
lanceolate. Sepals 5, white, linear, 0-5-0-9 mm long, unequal. Petals 5, pale
reddish-brown, l-5-2 0mm long, incurved, slightly hooded, acute. Stamens 5,
1 -7-3 0 mm long. Styles 2, 0-75-1 0 mm long, with a swollen stylopodium at the base.
Ovary slightly laterally compressed. Fruit yellow-brown, ± sessile or on a pedicel to
1 mm long and therefore hidden amongst the upper leaves; body of fruit obovoid and
slightly flattened laterally, T3-l-7mm long, 10-1-5 mm wide, O-9-l-Omm thick,
weakly ribbed; styles persistent, sepals ± deciduous.

This species is confined to Macquarie Island where it dominates the feldmark
community and other exposed windswept situations, forming extensive cushions and
tight mats. Flowering occurs from December to February and fruiting from January to
April. A detailed account of the ecology of the species is given by Taylor ( 1 955), of the
process of cushion formation by Ashton and Gill (1965) and of detailed distribution by
Copson (1984), all under the name A. selago.

Selected Specimens Examined (total 40):

MacQuarie Island — ‘Featherbed’ terrace, 7.xii.l948, Laird s.n. (AD, AK, BISH, CHR, HO 86261,
MEL); eastern side of :)awyer Creek Valley, 21.i.l981, Seppelt 11939 (HO); north side of Pyramid Lake,
4.xi.l981, Seppelt 12039 (HO)-, SW. side of Green Gorge, 4.i.l982, Seppelt 12390 (HO)-, near Flynn Lake,
29.xi.1950, Taylor s.n. (MEL 689443); Gadgets Gully, 3.ii.l951, Taylor s.n. (MEL 689445); North Mt,
4.iii.l951, Taylor s.n. (MEL 689450); Plateau, xi.l976, Tyler s.n. (HO 30818).

Notes:

This species is most obviously distinguished from^4. selago s.str. by its small size
and by the shape of its leaves. Upper (current year) leaves of A. macquariensis are
usually 3-lobed with the lamina divided almost to the base, and the lobes are acute and
bristle-tipped. In lower, older, leaves the bristle may be lost but the lobes
remain ± acute, rather than blunt and rounded as in A. selago. In particularly robust
plants or those growing in shaded places the leaves are larger and sometimes up to

17

Fig. 1 . Azorella macquariensis. A — Young 5-lobed leaf, abaxial view. B The same, lateral view. C ^ ^der
leaf from a robust (’shaded) branch, adaxial view, stem removed to show sheath and hgule . U —
Young 3-lobed leaf, glabrous form. E — Inflorescence showing a pair of flowers in an mvolucral cup. F
— Rower at anthesis. G — The same, 2 petals removed. H — Petal I — Stamen. J
view 2 sepals still attached. K — The same, commissural view. (A-C, E from Seppelt 12039\ D, F-1
from’ Tyler i n HO 30818; J, K from Taylor s.n., MEL 689445; all bar scales represent 1 mm.)

18

5-lobed, resembling those of A. selago, but the species can still be distinguished by the
characters of the tips of the leaf lobes, and also by the shape of the ‘ligule’. This is
formed as a prolongation of the membranous wings of the petiole. In A. tncicQucifiensis
the ‘ligule’ is short, little more than a ridge, and truncate or slightly rounded. In A.

consists of two distinct, rounded auricles. There are also supporting
differences in the flowers and fruits. In A. macquariensis the flowers are typically
solitary at the tips of the new shoots, sometimes paired, whereas in A. selago they are
borne m groups of three. Taylor (1955) mentioned that ‘ripe seeds are shed from
January to April’ in the Macquarie Island species, without further comment. However,
fruits are very scarce on existing collections. It is not clear whether this reflects a failure
of fruit formation or whether fruits in this species are shed rapidly and easily, not
surviving on herbarium specimens. The few surviving fruits are hidden amongst the
upper leaves on short pedicels and very difficult to hnd. On A. selago on the other hand
fruits seem to set readily and persist for some time on the plants. Their peduncles
elongate and they are held at least half exserted and often fully exserted above the
upper leaves. The sepals in A. selago persist on the fruit, white those of A
macquariensis are mostly shed.

2. Azorella selago J. Hooker, Flora Antarctica 2 (1845) t. 99, (1846) 284.

[Typus: ‘Tierra del Fuego, south part, C. Darwin, Esq. Port Famine, Capt. King.
Hermit Island, towards the top of the mountains, J.D.H. Kerguelen’s Land, covering
ffie ground near the sea, Anderson, J.D.H.’. Lectotypus (Moore 1968); Hooker 8
Tierra del Fuego, Isla Hermite, K (n.v.) Syntypi (?): Anon. (‘Erebus’ & ‘Terror’
Exped.), Cape Cumberland, Kerguelen Is., 1 840 (HO 1 08900 p.p.). — Anon. (‘Erebus’
& ‘Terror’ Exped.), Christmas Harbour, Kerguelen Is., 1840 (HO 108899 pp)l
Chastain, FI. Veg. lies Kerguelen (1958) 89-91; Moore, Vase. FI. Falkland Is. (1968)
Walton, Polar Record 17 (1975) 473-84; Moore, FI. Tierra del Fuego

( 1 983) 1 75—6.

Illustrations: J. Hooker, FI. Antarct. ( 1 845) PI. 99; Chastain FI. Veg. lies Kerguelen
(1958) PI. 2,3,23: Moore, Vase. FI. Falkland Is. (1968) Fig. 14c.

Perennial herbs forming extensive tight mats and cushions; main
branches ± prostrate, woody, 2-3 mm in diameter; lateral shoots erect, herbaceous,
freely branched, crowded, 3-15 cm tall, clothed in the remains of old leaves. Leaves
alternate, closely imbricate and appressed to the stems, persistent; petioles white,
3-3-5-0(-10) mm long with 5 longitudinal veins, the outermost weak, with a
membranous wing outside the veins, the wings fused in the lower ‘A-’/j to form a sheath
around the stem and produced above into free auricles at the base of the lamina;
lamina ± reniform, 21-4-2(-6-7) mm in radius, very thick, leathery, divided to about
'h (or less) of its depth into (4-)5-6(-7) lobes; lobes blunt, tips rounded or with at most
a tiny blunt apiculum (outermost lobes sometimes acute or long-apiculate), ± flat on
abaxial face, distinctly keeled on adaxial face, with thickened margins, glabrous or
with a few coarse bristles 1 •5-2-0(-3-0) mm long on adaxial surface, arising from the
veins. Flowers terminal, in groups of 3, ± enclosed by the upper leaves. Involucral
bracts 2, lobed, leaflike, fused at the base to form a small cup. Sepals 5, deltoid,
0-4-0-6 mm long, 0-4 mm wide, spreading, weakly midribbed. Petals 5, 1-5-1 -7 mm
long, incurved, slightly hooded. Stamens 5, 2-0 mm long. Styles 2, M-1-8 mm long,
curved, with a swollen stylopodium at the base. Ovary ovoid, c. 2 mm long. Fruit
olive-brown to yellow-brown, on a pedicel 2 mm long, at least '/ 2 -exserted from the
upper leaves; body of fruit ovoid to obovoid, slightly flattened laterally, 1-7-2-0 mm
long, 1-3-2-Omm wide, 1-3 mm thick, weakly ribbed; styles and sepals persistent.
(Fig. 2).

Extends from Tierra del Fuego to Falkland, Marion, Crozet, Kerguelen and
Heard Islands. In Tierra del Fuego it is found in feldmark communities from
450-1 100 m (Moore 1983). On Kerguelen Island the species grows in similar habitats,
but at lower altitudes (Chastain 1958). On Heard Island it is the dominant species.

19

Fig. 2. Azorella selago. A — Leaf with bristles, adaxial view, stem removed to show sheath and ‘ligule’. B —
Leaf glabrous form, abaxial view, sheath split and flattened. C The same, adaxial view. D
Inflorescence showing group of 3 old flowers/young fruits in involucral cup. E — Flower. F — Fruit,
dorsal view. G - The same, commissural view. (A, D, F, G from McGregor I2\ B, C, E from Bratt P7\
all bar scales represent 1 mm.)

20

^^ZithfjO)^ between the limit of seaspray and c. 100m

Specimens Examined:

Bran prfra (»0); f°°thills of Chateau Range, 17.ii.l971,

isratt r/ & Fo (HO), Hill of the Drumlins, 15. in. 1971, Bratt P21 (HO).

Heard Island — Old ANARE Station, Atlas Cove, 12.iii.l983 Cooson 103- Atlas Cove 8 li to/;^
"• ' ^50, Kenny s.n. (MEL 1554181); Atlas Cove, 20.ii. 1983, McGregor 12 (HO)'
104164); Fairchild Beach, 23.xii.1986, Scon s.n. (HO 104165)- Atlas
8.U fm ‘ ‘O'*’ ®3); near former ANARE base camp. Atlas Cove,

ACKNOWLEDGEMENTS

u Srsteful to Jenny Scott for the collection of Azorella and other genera on

Heard Island during the 1986/87 expedition, and to Dr R. D. Seppelt for earlier
collections Azorella from Macquarie Island. The Australian Biological Resources
S udy supported the curation of the important Laird collection of Macquarie Island
plants, and Dr J. H. Ross (MEL) kindly arranged the loan of the Taylor
specimens. ^

REFERENCES

Air Shaw, H. K. (ed.) (1973). J C Willis, ‘A Dictionary of the Flowering Plants and Ferns’, 8th edn.
(University Press: Cambridge.)

a"i?>'’' of New Zealand’, vol. 1. (Government Printer; Wellington.)

23^^^^ Pattern and process in a Macquarie Island feldmark. Proc. Roy. Soc.

Kerguelen. Polymorphisme des esp6ces australes.

Copson, G. R ( 1984). An annotated atlas of the vascular flora of Macquarie Island. ANARE Research Notes

lo! 1 — /U.

Greene, S. W & Greene, D. M. (1963). Check list of the sub-Antarctic and Antarctic vascular flora Polar
Kecora II: 411-418.

Greene, Walton, E).W H ( 1 975). An annotated check list of the sub-Antarctic and Antarctic vascular

rlora. Polar Record 17: 473-484.

u’aa ^(**‘*5-6) Flora Antarctica. II. t. 99 & pp. 284-285, in The Botany of the Antarctic Voyage of
H.M. Discovery Ships ‘Erebus’ and ‘Terror’. (Reeve: London.)

Mathias, M. E. &Constance, L. (1971). Umbelliferae, in Lasser, T., ‘Flora de Venezuela’ Vol 3 no 93-168
(Instituto Botanico: Caracas.) •

Moore, D^2M. (1968). The vascular flora of the Falkland Islands. British Antarc. Sun. Sci. Rep. 60:

Moore, D. M. (1983). ‘Flora of Tierra del Fuego’. (Nelson; Oswestry & Missouri Botanical Garden-
St. Louis.)

Taylor, B. W. (1955). The flora, vegetation and soils of Macquarie Island. ANARE Rep., Bot. 2: 1-192.

Manuscript received 1 March 1 988

NOTES ON HOVEA R. Br. (FAB ACE AE): 2

by

J. H. Ross*

ABSTRACT

Ross, J. H. Notes on Hovea R. Br. (Fabaceae): 2. Muelleria 7(1): 21-38 (1989). The six
Western Australian species of the endemic Australian genus Hovea are revised.
Descriptions, a key to the identification of the species, illustrations, and distribution
maps are provided, together with notes on ecological preferences and relation-
ships.

INTRODUCTION

The genus Hovea is represented in Western Australia by six species which are
distributed largely within the South-West Botanical Province as defined by Beard
(1980) with outliers in the Coolgardie Botanical District of the South-Western
Interzone. The Western Australian species are isolated geographically from the
remaining species in the genus and none is allied closely to any of the eastern or the
northern species. Some specimens of H. trisperma with narrow leaves are superficially
similar to specimens of H. linearis (Sm.) R. Br. from New South Wales but the two
differ in many significant respects. In contrast to the eastern species, the Western
Australian species are relatively straightforward taxonomically.

The Western Australian species do not form a homogeneous group and, apart
from the obvious close relationship that exists between H. stricta and H. purigens, their
affinities are not obvious. Some of the xeromorphic adaptations exhibited in the
Western Australian species such as the rigid pungent-pointed leaves of H. pungens and
H. stricta, the coriaceous leaves with sinuate prickly-toothed margins of H.
chorizemifolia, and the short lateral spine-tipped shoots and minute leaves of H.
acanthoclada are not found elsewhere in the genus.

The flowers of the Western Australian species are a deep intense blue or
purplish-blue (except for the occasional white-flowered variants which occur
throughout the range of the genus) in contrast to those of the eastern species which are
usually a paler purplish-blue, pinkish-purple or insipid mauve. The inner surfaces of
the pod valves of many of the eastern species are pubescent whereas those of the
Western Australian species are all glabrous.

Key to the Western Australian species

1. Branches armed with short lateral spreading spine-tipped leaf-bearing shoots;

leaves minute, up to 7 mm long H. acanthoclada

1 . Branches lacking spine-tipped shoots; leaves larger than above:

2. Leaves rigid, the margins usually strongly revolute and apices mostly
pungent-pointed:

3. Leaves sessile or on petioles up to 0-5 mm long, typically inserted on the
branch at an angle of more than 45° and almost at right angles to it, the lamina
arching outwards or spreading laterally and often slightly reflexed; seeds

mottled 5. 77. pungens

3. Leaves on short but distinct petioles 0-75-1 -5 mm long, typically inserted on
the branch at an acute angle (less than 45°) the lamina arching outwards and
upwards with the apex pointing towards the apex of the branch; seeds
uniform yellowish or olive-brown 6. 77. stricta

•National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

21

22

2. Leaves scarcely or distinctly coriaceous but not rigid, margins not strongly
revolute, apices not pungent-pointed, or, if so, then margins usually sinuate and
prickly-toothed:

4. Leaf margins usually distinctly sinuate and pungent-pointed, occasionally
almost entire and almost lacking teeth but the upper stamen-filament free

from the tube 2. H. chorizemifolia

4. Leaf margins entire, crisped or almost crenulate but not pungent-pointed,
upper stamen-filament usually united to the tube:

5. Subshrub to 0-6 m high, stems often weak and sprawling unless supported;
exterior of calyx clothed with appressed to slightly spreading hairs, the
hairs mainly silvery-white or sometimes with an understorey of shorter
rust-coloured hairs; lower surface of leaf with appressed, spreading or

asymmetrically biramate hairs, the hairs often crimped; ovules 2-7

3. H. trisperma

5. Slender erect shrub or tree to 3 m high; exterior of calyx clothed with
appressed hairs, the hairs all short and rust-coloured or sometimes with
paler longer hairs interspersed; lower surface of leaf with predominantly
or exclusively medifixed or asymmetrically biramate hairs; ovules 2 (very
rarely 3 or 4) 4. //. elliptica

1. Hovea acanthoclada F. Muell., Fragm. 4: 15 (1863); Benth., FI. Austral. 2: 174
(1864). Lectotype (here selected): Western Australia, Phillips River, Maxwell s.n.
(MEL).

Daviesia? acanthoclada Turcz., Bull. Soc. Imp. Naturalistes Moscou 26: 262
(1853). Lectotype (here selected): Western Australia, Drummond 5th coll. No. 90.
(KW; Isolectotype: BM, G, K, NSW, PERTH, W).

Rigid divaricate shrub to 2 m high, single or many-stemmed, armed with short
lateral spreading spine-tipped shoots which bear leaves and inflorescences; branches
densely clothed with appressed to slightly spreading antrorse hairs, some of which are
often asymmetrically biramate. Leaves solitary or appearing fascicled in twos or
threes; lamina oblong, narrow-obovate, ovate or rotund, 1-5-7 mm long, 0-9-2 mm
wide, rounded, obtuse or retuse apically, with or without a short mucro, margins
revolute, glabrous above, sparingly to densely clothed with appressed often
asymmetrically biramate hairs below; petiole up to 0-5 mm long. Inflorescence
axillary, sessile or on peduncles up to 1 -2 mm long. Flowers solitary or paired,
pedicellate, the pedicels 1-5-4 mm long, clothed with appressed hairs; bracteoles
± 1 mm long, inserted at the base of or a short distance below the base of the calyx,
much shorter than the calyx-tube, pubescent like the pedicel and bract; bract ± 1 mm
long, inserted 1-2 mm below the bracteoles. Calyx turbinate-campanulate, densely
clothed with short appressed hairs; 2 upper lobes 4- 5-5 -5 mm long including the tube
3-3-5 mm long, ± truncate except for an acute tip; the lower 3 lobes 1 -5-2-5 mm long.
Standard 8-10-2 mm long, 8-10 mm wide, purplish-blue with a basal whitish
horse-shoe shaped flare; wings 7-9 mm long, 2 -9-3 -7 mm wide, purplish blue; keel
5-7-6-5mm long, 2-2-3 mm wide, purplish-blue apically, white basally.
Stamen-filaments 4-5-6 mm long. Ovary 1-2-1 -5 mm long, 2-ovulate, glabrous, on a
stipe up to 1 mm long. Pods shortly stipitate, the stipe almost as long as the calyx,
obliquely to transversely globular, ovoid or ellipsoid, 0-7-0-9cm long, 0-9-1 -1 cm
wide, glabrous. Seeds elliptic, plump, 4-4-8 mm long, 2-3-3 mm wide, 2-2-2-7 mm
thick, dark brown, aril extending for more than half the length of the seed (Fig.
Id-f).

Confined to southern Western Australia where there are two disjunct centres of
distribution: one in the vicinity of Kalgoorlie and the other further south from the
Fitzgerald River area to east of Ravensthorpe and southwards to Hopetoun (Fig. 2).
Recorded from rocky outcrops, scree slopes, laterite and loam over granite in mallee,
in tall Eucalyptus woodland and in association with Acacia, Casuarina and Hakea
species.

23

Representative Specimens (total number examined 23):

Western Australia — Kalgoorlie, 16.viii.l963, T. E. H. Aplin 2259 (PERTH); Mt Short, N. of
Ravensthorpe, 30. viii. 1 963,^. S. George 5711 (PERTH); E. side of Fitzgerald River valley, above Roes Rock,
18.xii.l970, S. George /0552 (PERTH); 13 km S. of Ravensthorpe towards Hopetoun, 27.vii.1983, G. J.
Keighery 6223 (PERTH); 1 7 km due NE. of Ravensthorpe, nearWoodenup Creek, 30.viii. 1980, B. R. Maslin
4 773 (PERTH).

Typification:

Turczaninow based his description of Daviesia? acanthoclada on Drummond 5th
coll. No. 90 and, as a description of the pods and seeds is included in the protologue, it
is clear that he saw a fruiting specimen. It would appear reasonable to assume that the
Drummond specimen that Turczaninow saw is the one housed in KW but the
specimen in KW lacks pods and seeds; there is no means of knowing whether or not
pods and seeds were present in Turczaninow’s time and have since been lost or
whether he examined a specimen with pods and seeds elsewhere. Specimens of
Drummond 5th coll. No. 90 in G (2 sheets), K (2 sheets), NSW, PERTH (ex K) and W
bear pods and seeds. Despite the lack of pods and seeds, I here select the Drummond
collection in KW as the Lectotype of D. ? acanthoclada, the collections in G, K, N S W ,
PERTH and W being treated as Isolectotypes.

Mueller, Fragm. 4: 15 (1863), based his description of H. acanthoclada as far as
one can tell on a Maxwell collection from Phillips River. There is no reference by
Mueller, either direct or implied, to the earlier name Daviesia? acanthoclada Turcz.
based on Drummond 5th collection No. 90.

It has been customary to regard H. acanthoclada as a new combination by Mueller
based on Daviesia? acanthoclada but H. acanthoclada Mueller is treated here as a new
name as I believe that when Mueller described H. acanthoclada he thought that he was
providing a name to a taxon that previously was without a name. Perusal of the pages
of Fragm. 4 following the description of H. acanthoclada shows that Mueller was
meticulous about citing references to earlier published descriptions if he was aware
that they existed. The absence of any reference whatsoever by Mueller to D.?
acanthoclada therefore seems to be of significance.

Circumstantial evidence supports the view that Mueller was unaware ot
Turczaninow’s earlier name. In the Appendix to the 1864—65 Annual Report as
Director of the Botanic Gardens, Bull. Soc. Imp. Naturalistes Moscou is not listed by
Mueller among the holdings of the library. It is probably unlikely that he would have
omitted the journal from his list inadvertently. Records in the State Library of Victoria
indicate that the journal was not acquired by the State Library prior to 1 869 and there
was no other library in Victoria likely to have contained the publication. There is no
specimen of Drummond’s 5th collection No. 90 in MEL, so Mueller could not have
seen a named collection. All of the incoming correspondence to Mueller was destroyed
in the 1 930s so there is no means of knowing whether he was aware of the name from
that source.

Mueller’s choice of the same specific epithet as Turczaninow does not seem to be
too great a coincidence for credibility. The occurrence of spine-tipped branches occurs
nowhere else in the genus so the presence of this character would suggest itself as a
fairly obvious specific epithet. Furthermore, Mueller had used the same epithet for
Acacia acanthoclada in Fragm. 3: 127 (1863) published shortly before H.

acanthoclada. , . • • .

H. acanthoclada differs from all of the other species in having spine-tipped
branches and minute leaves. It is reminiscent of some of the xeromorphic Bossiaea
species.

2 Hovea chorizemifolia DC., Prodr. 2:116(1825) (as chorizemaefolia)-, Sweet, Hortus
Brittanicus 2: addenda 475 (1827); Lindley in Edwards’s Bot. Reg. 18: t. 1524 (1832)
(as chorozemaefolia)', Benth., FI. Austral. 2: 174 (1864).

Plagiolobium chorizemifolia (DC.) Sweet, H. Australasica t. 2 (1827); Meissn. in

24

Uhm., PL Preiss. 1: 80 (1844); Wheeler, FI. Perth Region 268 (1987). Lectotype
(here selected): Western Australia, collector unknown, (G-DC)

H. ihcifolia Sweet, Hortus Britannicus 1:111 (1826) nomen nudum
Plagiolobium ilicifohum Sweet, FI. Australasica t. 2 (1827) partly but excluding
the pod; Meissn m Lehm., PI. Preiss. 1: 80 (1844). Neotype: Western Australia!
Drummond 182 (MEL; Isoneotype: BM, K, NSW, W).

H. ilicifolia Cunn. in Lindley, Edwards’s Bot. Reg. 30: t. 58 (1844). Lectotype
(here selected): Western Australia, King Georges Sound, A. Cunningham (CGE).

1 o c chorozemaefolium var. subintegrum Meissn. in Lehm., PI. Preiss. 1 :
80 (1844). Syntypes: Western Australia, between Greenmountain and Mahogany
Creek, 12.ix.l839, Preiss 1052 (HBG, K, LD, MEL, NY, W); Western Australia,
Drummond 181 (BM, K, MEL, NSW, W).

Plagiolobium chorozemaefolium var. dentatum Meissn. in Lehm., PI. Preiss. 1 : 80
(1844). Lectotype (here selected): Western Australia, Mount Melville, 25.ix.1840.
Preiss 1058 (LD; Isolectotype S).

Shrub or subshrub to 0-6 m high with one or several stems arising from the base
stems branched or sometimes simple, sparingly to densely clothed with appressed
antrorse or spreading often rusty hairs, glabrescent. Leaves', lamina ovate, elliptic to
lanceolate, those on lower leaves often shaped differently to those on upper leaves
margins usually distinctly sinuate and prickly-toothed but occasionally almost entire
and almost lacking teeth, 1-5-8 cm long including a pungent point up to 0-6 cm long,
0-8-4 cm wide (including the teeth), coriaceous, glabrous throughout or with scattered’
hairs on lower surface of midrib and/or lamina, venation sometimes prominently
reticulate; petiole 0-5-3-5 mm long. Stipules subulate, up to 4-5 mm long, 0-45 mm
wide. Inflorescence axillary, sessile or on peduncles up to 1 - 5 mm long, 2-6 flowered.
Flowers pedicellate, the pedicels 2-5 mm long, densely pubescent with long straightish
hairs scattered in amongst shorter curled hairs; bracteoles subulate, up to
3-5 X 0-6 mm, inserted ± 1 mm below the calyx, pubescent when young but gla-
brescent; bract subulate, up to 3x0-6 mm, inserted 1-3-5 mm below the bracteoles.
Calyx densely clothed with long straightish hairs and shorter curled or crinkled hairs; 2
upper lobes 5—6 mm long including the tube 2-2-5 mm long, up to 7 mm across,
emarginate; the 3 lower lobes 1 -2-2-5 mm long. Standard 10-4-14 mm longi
10-13 mm wide, purplish-blue with a basal whitish horse-shoe shaped flare; wings
8-9-5 mm long, 4-4-5 mm wide, purplish-blue; keel 4-7 mm long, 2-5-3 mm wide.
Stamen-filaments 4-4-5 mm long, staminal sheath open on the upper side and
sometimes also on the lower, upper filament usually free. Ovary sessile, 1-1-5 mm
long, 2-ovulate, glabrous. Pods shortly stipitate, globular, ovoid or ellipsoid,
0-8-1 -1cm long, 0-8-1 cm wide, glabrous. Seeds elliptic, 4-5-5-5 mm long,
3-2-3-7 mm wide, olive-brown or brown, aril extending for more than half the length
of the seed. (Fig. 1 g-i).

Confined to the Darting, Avon and Eyre Botanical Districts of the Southwestern
Botanical Province of Western Australia as defined by Beard (1980) occurring from
the vicinity of Bindoon north-east of Perth south-eastwards to York, Narrogin and the
Bremer River north-east of Albany (Fig. 2). The label accompanying one specimen
(MEL 6671 33) alleges that the specimen was collected by Maxwell in 1 875 ‘near Cape
And’. This is considerably further east than any other records of the species and
suggests that the label may not belong with the specimen.

Recorded frc)m laterite, gravel, granite outcrops, sand and in karri forests in sandy
soils rich in organic matter. In the northern part of its range and in the Darling Range
H. chorizemifolia often grows in association with H. trisperma whereas in the karri
forests it is often associated with H. elliptica.

Representative Specimens (total number examined 246):

Western Australia — 5 km S. of Margaret River township on Caves Road, 28.x. 1983, M.G. Corrick
8966 (MEL); Bindoon, 3.ix.l964, J. Galbraith 56IA (MEL). Dwellingup, 25.ix.1942, C.A. Gardner 6484
(PERTH); Greenmount, Darling Range, 23.vii.1898, A. Morrison (AD, MEL, PERTH); Knoll Drive
Nornalup-Walpole National Park, 9.ix.l971, S. Paust 346 (PERTH); York, 4.vi.l905, OH. Sargent

25

Fig. 1 . Hovea elliptica. a — fruiting twig, X 1 . b — seed, side view, X 6. c — seed, hilar view, X 6. H.
acanlhoclada. d — flowering twig, X 1. e — seed, side view, X 6. f — seed, hilar view, X 6. H.
chorizemifolia. g — fruiting twig, X 1 . h — seed, side view, X 6. i — seed, hilar view, X 6. a-c from R.A.
Kilgour 520 (MEL); d from G J. Keighery 6223 (PERTH); e-f from A.S. George 70552 (PERTH); g from
M.G. Corrick 9597 (MEL); h and i from F. Mueller s.n. (MEL 667151).

26

Fig. 2. The known distribution of Hovea acanthoclada (•) and H. chorizemifolia (a).

Typification:

There are in De Candolle’s herbarium in Geneva four specimens of H.
chorizemifolia. The specimen bearing a label which reads ‘aff. chorizema. Nouvell(e)
Holland cote orient Mus de Paris 1491’ pinned to the sheet is selected from among
these specimens as the Lectotype of H. chorizemifolia. De Candolle recorded that the
species came from eastern Australia but this is an error.

Plagiolobium ilicifolium, described by Sweet in a footnote, was based on a
specimen in A. B. Lambert’s herbarium collected by Captain King at King Georges
Sound. Unfortunately I have not succeeded in locating this specimen but it is clear that
if the pod was described correctly as pubescent internally and externally it did not
belong with the specimen. The alternative possibility that neither the specimen nor the
pod is referrable to H. chorizemifolia is considered less likely as the leaf illustrated in
t. 2 falls within the range of variation of this species. As it is considered that the leaf
alone illustrated in t. 2 is inadequate for the purpose of serving as a lectotype,
Drummond 182 (MEL) is selected here as the Neotype of P. ilicifolium. Isoneotypes
are housed in BM, K, NSW and W.

It is clear from the protologue that H. ilicifolia Cunn. was recognized as a species
distinct from Plagiolobium ilicifolium Sweet because it had glabrous pods in contrast
to the pods of the latter species which had been described as pubescent. H. ilicifolia was
based on a plant or plants raised by R. Mangles from seed from Swan River and on
material collected by A. Cunningham at King Georges Sound. There is in Lindley’s
herbarium at CGE a sheet consisting of three twigs, two of which bear persistent
calyces, an envelope containing the remnants of a pod and a mature and an immature
seed, and a label just above the bottom right hand comer which reads "Hovea ilicifolia
C. King Georges Sound. 1822’. Although the collector is not indicated, I am satisfied
that this is one of the Cunningham specimens alluded to by Lindley. In Herbarium
Benthamianum at K there is a sheet containing four different collections including
some of Cunningham’s. The flowering specimen second from the right, which is
mounted with the apex at the foot of the sheet, has a label attached to its base with

27

‘Hovea ilicifolia see Plagiolobium' written on it. The label at the foot of the sheet which
covers the apex of the shoot indicates that the specimen was collected by Cunningham
at King Georges Sound. These two collections are regarded as Syntypes of H.
ilicifolia. I here seleet the fruiting specimen in CGE as the Lectotype of H. ilicifolia
Cunn..

Notes:

The upper stamen-filament in H. chorizemifolia is usually free from the others, a
feature which occurs only sporadically and irregularly in other speeies.

The leaves typically have distinctly sinuate pungent-pointed margins and a
pungent apex and are very characteristic. However, leaf size and leaf shape, especially
the shape of the margins, vary quite markedly and occasional specimens occur in
which the margins are only slightly sinuate or undulate and possess only a few marginal
teeth, for example, J.H. Willis s.n. (MEL 1532104). Some of these specimens
(corresponding to var. subintegrum recognized by Meissner), which occur within the
distributional range of both H. chorizemifolia and H. elliptica, have been confused in
the past with H. elliptica or considered as intermediates between the two species but
they are referrable to H. chorizemifolia. These specimens have the bract and bracteoles
of H. chorizemifolia rather than those of H. elliptica, the upper stamen-filament is
always free as in H. chorizemifolia and the lower surfaces of the leaves lack the
distinctive asymmetrically biramate or medifixed hairs which are characteristic of
elliptica. Even although the two species often grow together their habits are quite
different. Leaf shape in H. chorizemifolia is clearly more variable than previously
realized, but, despite this, there does not appear to be a means of dividing up the range
of variation satisfactorily.

White-flowered variants occur occasionally, for example, Mrs W.A. Ross
(PERTH) from Waroona.

3. Hovea elliptica (Sm.) DC., Prodr. 2: 1 15 (1825); Sweet, Hortus Britannicus 1:111
(1826); Benth., FI. Austral. 2: 175 (1864).

Poiretia elliptica Sm., Trans. Linn. Soc. Lond. 9: 305 ( 1 808); Phusicarpos elliptica
(Sm.) Poiret in Lamarck & Poiret, Encycl. meth. Bot. suppl. 4: 400 ( 1 81 6). Lectotype
(here seleeted): Western Australia, King Georges Sound, 1 803, Menzies (LINN, sheet
1190.2). ,

Platychilum celsianum Delaunay, Herb. Amat. t. 1 87 ( 1 8 1 5); Goodia simplicijolia
Spreng., Syst. Veg. ed. 16, 4(2): 267 (1827). Lectotype (here selected): Delaunay,
Herb. Amat. t. 187.

H. celsii Bonpl., Descr. PI. Malmaison t. 51 (1816). Lectotype (here selected):
Descr. PI. Malmaison t. 51.

Slender shrub or tree to 3 m high, often single-stemmed, young branches densely
clothed with appressed to slightly spreading hairs, the hairs predominantly or
exclusively medifixed or asymmetrically biramate, often rust-coloured. Leaves.
lamina almost flat, elliptic, ovate-elliptic, obovate-elliptic to obovate or fusiform,
(l-5-)2-5-10(-14)cm long, (0-5-)l-3-2(-6) cm wide, obtuse, emarginate, retuse or
mucronate apically, glabrous above and reticulate, the venation usually prominent,
lower surface and midrib sparingly to densely elothed with predominantly or
exelusively medifixed or asymmetrically biramate hairs; petiole 0-8—1 cm long,
sparingly to densely clothed with medifixed or asymmetrically biramate hairs. Stipules
narrow-triangular, up to 1 mm long and 0-5 mm wide, sparingly to densely clothed
with medifixed or asymmetrically biramate hairs. Inflorescence axillary, sessile or a
pedunculate raceme, sometimes auxotelic, 1—7 flowered. Flowers pedicellate, the
pedicels 4-9 mm long, densely clothed with rusty appressed hairs; bracteoles
1-1-5 mm long, up to 0- 5 mm wide, inserted at the base of the calyx and appressed to it
or inserted up to 1 mm below the calyx and free from it, densely clothed with appressed
rusty hairs; bract 1-1-5 mm long, up to 0-5 mm wide, inserted at base of pedicel and
5-8 mm below the bracteoles, densely clothed with rusty appressed hairs. Calyx

28

densely clothed with appressed hairs, the hairs either all short and rusty or sometimes
with longer paler hairs interspersed: 2 upper lobes 5- 1-6-4 mm long including the tube
1 -8-2-9 mm long, 6-8-1 mm across, emarginate; the 3 lower lobes 1 -3-2-5 mm long.
Standard 10-5-14-5 mm long, 11-5-17 mm wide, purplish-blue with a basal whitish
horse-shoe shaped flare; wings 7-8-9-6mm long, 3-5-5-5 mm wide, auricled,
purplish-blue; keel petals 7-2-8- 1 mm long, 2-6-3-2 mm wide, the inner margin
sometimes papillate, purplish-blue. Stamen-filaments 5-5-6-2 mm long. Ovary
1-2-1 -6 mm long, on a stipe 1-1-5 mm long, ovules 2 (rarely 3 or 4), glabrous; stigma
papillate. Pods stipitate, the stipe about as long as the calyx, obliquely to transversely
globular, ovoid or ellipsoid, 0-8-l-2cm long, 0-8-l-3cm wide, 0-7-1 cm thick,
glabrous. Seeds elliptic, plump, 3-8-5- 1 mm long, 2-8-3- 1 mm wide, 2-2-7 mm thick,
olive-brown or brown, the aril extending for more than half the length of the seed.
(Fig. la-c).

Confined to the Menzies and Warren subdistricts of the Darling Botanical
District and the western Eyre Botanical District of the Southwestern Botanical
Province of Western Australia as defined by Beard (1980) occurring from the Preston
River in the north to Cape Naturaliste and Cape Leeuwin in the west and
south-eastwards to the Bremer River (Fig. 3). The label accompanying one specimen
(MEL 666464) alleges that the specimen was collected by Maxwell in 1 875 ‘near Cape
Aird’. This is considerably further east than any other records of the species and
suggests that the label may not belong with the specimen.

Recorded from laterite, gravel, granite outcrops, clay loam, stabilized sand dunes,
sandy loam and sandy soils rich in organic matter. A common understorey plant, often
with Bossiaea laidlawiana and H. cfiorizemifolia, in karri-jarrah-marri forests and
woodland.

Representative Specimens (total number examined 222):

Western Australia — 5 km S. of Margaret River township on Caves Road, 28.x. 1983, M.G. Corrick
8959 (CBG, HO, MEL, NSW, PERTH). Davidson’s Road, near comer of Coronation Road, W. of
Manjimup, 10.x.l984,MG. Corrick 92 38 (Um.). Willyung Hill, about 12km N. of Albany, 23.ix. 1984, D.fi.

Fig, 3. The known distribution of Hovea elliptica.

29

Foreman 829 (AD, CBG, MEL, PERTH). Northern slope of Castle Rock, Porongurup Range, 29.ix.1966,
T.B. Muir 3966 (MEL). Bremer River, 1884, W. Webb (MEL 66439).

Notes:

In stature H. elliptica is by far the largest of the Western Australian species. Some
specimens of H. trisperma with large elliptic leaves are sometimes superficially similar
to and have been confused with H. elliptica. Apart from differences in habit, H.
elliptica invariably has 2 ovules per ovary in contrast to the specimens of H. trisperma
which have 3-6 ovules per ovary, the indumentum on the lower surface of the leaves is
different and the shape of the lamina itself is different.

Occasional white-flowered specimens of H. elliptica occur.

4. Hovea trisperma Benth. in Endl. et ai, Enum. PI. Huegel 37 (1837); Meissn. in
Lehm., PI. Preiss. 1: 79 (1844); Benth., FI. Austral. 2: 175 (1864); Wheeler, FI. Perth
Region 269 (1987). Lectotype (here selected): Western Australia, King Georges
Sound, Huegel s.n. (W).

H. lanceolata var. linearis Lindley in Edwards’s Bot. Reg. 17: t. 1427 (1831).
Lectotype (here selected): Fragment from cultivated plant in Low’s nursery, 1831,
(CGE).

H. manglesii Lindley in Edwards’s Bot. Reg. 24: t. 62 (1838). Lectotype (here
selected): Specimen from plant cultivated by R. Mangles, 1837, (CGE).

H. crispa Lindley in Edwards’s Bot. Reg. 25: misc. 19 (1839). H. trisperma var.
crispa (Lindley) Benth., FI. Austral. 2: 176 (1864). Lectotype (here selected):
Specimen from plant cultivated by R. Mangles (CGE).

H. grandiflora Drummond, Hooker’s J. Bot. 2: 365 (1840). H. trisperma var.
grandiflora (Drummond) Benth., FI. Austral. 2: 176 (1864). Lectotype (here
selected): Western Australia, Drummond (K).

H. splendens Paxton, Paxton’s Mag. Bot. 10: 70, plate facing p. 103 (1843).
Lectotype (here selected): Paxton’s Mag. Bot. 10: plate facing p. 103.

H. elliptica sensu Meissn. in Lehm., PI. Preiss. 1: 79 (1844) non (Sm.) DC.

Subshrub to 0-6 m high with one or several stems arising from the base, stems
often weak and sprawling unless supported by surrounding vegetation, young
branches sparingly to densely clothed with appressed medifixed or asymmetrically
biramate antrorse or spreading hairs, the hairs sometimes crimped or twisted. Leaves:
lamina ovate, obovate, elliptic, elliptic-oblong, lanceolate, linear-oblong or on basal
leaves occasionally almost rotund, those on lower leaves often shaped differently to
those on upper leaves, (0-8-)2-8(-13) cm long, (0-3-)0-7-3-6 cm wide, obtuse, acute
or shortly mucronate apically, margins slightly to distinctly recurved, entire, crisped or
almost crenulate, upper surface pubescent especially when young or glabrous,
sometimes prominently reticulate, lower surface sparingly to densely clothed with
appressed to spreading or asymmetrically biramate often crimped hairs; petiole

1- 3 mm long. Stipules subulate, up to 2 mm long, pubescent, sometimes persisting.
Inflorescence axillary, sessile or on short peduncles, 1 -6-flowered. Flowers pedicellate,
the pedicels 1-7 mm long, densely clothed with appressed to spreading hairs;
bracteoles subulate, 1-5-3 mm long, inserted at the base of or up to 2 mm below the
calyx, pubescent; bract subulate, up to 3 mm long, inserted at the base of the pedicel.
Calyx densely clothed with appressed antrorse to slightly spreading hairs: 2 upper
lobes 6-1-1 1-7 mm long including the tube 2-4-5 mm long, emarginate; the 3 lower
lobes 1 -4-2 mm long. Standard 10-8-20-2 mm long, 10-5-25 mm wide, purplish-blue
with a basal white horseshoe shaped flare; wings 9-14-8 mm long, 3-1-8 mm wide,
purplish-blue; keel 6-11-6 mm long, 2-3-4 mm wide. Stamen-filaments 5-8 mm long,
staminal sheath open on upper side and sometimes also on lower, occasionally the
upper filament free. Ovary subsessile or on a stipe up to 2 mm long, 2-7-ovulate,
glabrous. Pods shortly stipitate, the stipe about half as long to as long as the calyx-tube,
globular, ovoid or ellipsoid, sometimes transversely so, 0-8-1 -2 cm long, 0-8-1 -2 cm
wide, C-65-1 cm thick, glabrous. Seeds elliptic, 4-6 mm long, 2-6-3-8 mm wide,

2- 2-7 mm thick, uniform olive- to dark brown, aril collar-like with a raised upper lip,
about j to f as long as the seed, margin sometimes slightly frilled.

30

Confined to the Irwin, Avon, Darling and Eyre Botanical Districts of the South-
western Botanical Province of Western Australia as defined by Beard ( 1 980) occurring
from Mogumber in the north southwards to Augusta and eastwards with several large
disjunctions to Duke of Orleans Bay east of Esperance. There are three early records
from two localities far removed from the general range of distribution and
confirmation of the existence of the species at these disjunct localities is required. The
labels accompanying two specimens (NSW 58717, NSW 58737) indicate that they
were collected by L. C. Webster in 1899 and 1900 respectively at Coolgardie, and the
label accompanying a third specimen indicates that it was collected by F. Stoward
(PERTH) at Geraldton in Sept. 1917. These localities are respectively much further
west and north than those of any other records (Fig. 4).

Recorded from sandy soil in heath, tall Banksia, Xanthorrhoea or Casuarina -
marri woodland, and in laterite, gravel and clay loam in Eucalyptus woodland or
mallee. Occasionally found in wet low-lying areas in association with Melaleuca. Often
growing in association with H. chorizemifolia in areas where the distribution of the
two species overlaps.

Representative Specimens (total number examined 325):

Western Australia — Logue Brook Dam, hillside above Clarke Brook, 9.x. 1984, M.G. Corrick 9212
(MEL 669610); near Brockton Highway, ca. 1 1 km E. of Kelmscott, lO.x.1985, M.G. Corrick 9635 (MEL
677476); near Gingin, 17.vii.l920, C.A. Gardner 574 (MEL 1529399, PERTH); Wharton Beach, Duke of
Orleans Bay, 30.ix.l985, C.W. Huggins s.n. (MEL 1540176); Bayswater, 22.vi.1907, A. Morrison
(PERTH).

Typification:

The Lectotype of H. trisperma Benth. as here selected consists of two shoots, the
left hand one in flower and the right hand one with young pods, mounted on a sheet in
W bearing a label in the lower right hand corner which reads ‘111 Hovea trisperma
King George’s Sound Hiigel’. The dehisced pods in an envelope bearing the number

31

40 1 mounted on the top left hand corner of the sheet do not belong with the mounted
specimens on this sheet and are excluded from H. trisperma.

Lindley based his description of H. lanceolata var. linearis on a plant in
cultivation in the nursery of Messrs Low and Co. of Clapton in March 1831.
Fortunately a small fragment of the plant from Low’s nursery is preserved in Lindley’s
herbarium at CGE and it is selected here as the lectotype of H. lanceolata var. linearis.
The fragment is mounted in the lower right hand corner of the sheet, the remainder of
the sheet being occupied by a specimen collected at the Vasse River by Mrs G. Molloy
in 1839.

The description of H. manglesii Lindley was based on a specimen cultivated by
Robert Mangles of Sunning Hill in Jan. 1837 and on specimens collected in Western
Australia subsequently (possibly by Mrs G. Molloy) and communicated to his brother,
Capt. James Mangles. The specimen cultivated by Robert Mangles in 1837 and a wild
specimen from Western Australia, together with one collected by Drummond in 1839,
are mounted on a sheet in Lindley’s herbarium in CGE. Although Lindley mentioned
having received wild specimens from Capt. Mangles I have only traced the one
referred to. I here select the specimen cultivated by Robert Mangles in 1837 mounted
in the lower right hand corner of the sheet as the Lectotype of H. manglesii. As noted
by Lindley, the leaves in this specimen are almost twice as wide as those on the
specimen collected in the wild. Lindley commented that H. manglesii dilfered from H.
trisperma in having a sessile ovary but the ovary illustrated in Edwards’s Bot. Reg. 24;
t. 62 is shortly stipitate.

Lindley based his description of//, crispa on a plant cultivated by Robert Mangles
of Sunning Hill. A specimen preserved in Lindley’s herbarium at CGE, mounted on a
sheet to the right of a specimen collected by Drummond in 1 839, is here selected as the
Lectotype of H. crispa. In the protologue Lindley described the ovary of H. crispa as
4-seeded. Although 3 or 4 ovules are recorded in H. crispa the ovaries are usually
2-ovulate and the ovary of a flower on the lectotype was found to have 2 ovules.

H. grandijlora Drummond was described from material collected between the
Darling Range and Toodyay. The actual locality given was ‘the west side of a hill,
which the road crosses about a mile to the east of the watering-place called Goolgoil, by
the natives . . .’. I have not succeeded in tracing this locality and efforts by Mrs R.
Paynter of Toodyay and Mr and Mrs B. H. Smith of Manmanning have likewise been
unsuccessful. There are in Herbarium Hookerianum at K two sheets of material
bearing three different collections collected by Drummond which are referrable to H.
grandijlora. All three collections are labelled Swan River, Drummond, and although
none is named H. grandijlora at least two of the collections match the brief description
and undoubtedly represent type material. I here select the collection to the right of the
‘Herbarium Hookerianum’ stamp on the right hand side of the sheet containing two
different collections as the Lectotype of H. grandijlora. The second collection on the
sheet bears very much smaller leaves. The Lectotype consists of two flowering
branches, the one on the right mounted with the base at the top of the sheet and the
apex at the foot. The shape of some of the leaves on the second sheet of Drummond
material matches the description in the protologue better but the specimen is a poor
one and consequently was not chosen as Lectotype. Drummond specimens are
housed in several other herbaria, for example BM, E, MEL and W, but there is no
means of knowing whether or not they formed part of the type collection. E)espite the
absence of the name ‘grandijlora' on either of the sheets in K I am satisfied that
Bentham would have been aware of Drummond’s name and that Bentham’s var.
grandijlora was based upon H. grandijlora Drummond.

H. splendens Paxton was based on a plant cultivated by Mr Knight, Chelsea, from
seed collected in Swan River Colony and sent to Capt. Mangles. I have not succeeded
in locating a specimen upon which Paxton’s plate was based and consequently now
select the plate facing p. 103 in Paxton’s Mag. Bot. 10 (1843) as the Lectotype of H.
splendens. I am not absolutely certain of the identity of H. splendens as I have seen no
specimen that exactly matches the illustration or description but it is a probable
synonym of H. trisperma. Bentham (1864, 2: 176) was of the opinion that it was ‘very
near’ var. crispa.

32

Notes:

H. trisperma is a variable species within which three main variants are evident
and these correspond to the varieties trisperma, crispa and grandiflora recognized by
Bentham (1864). Variety crispa was characterised by Bentham as having ovate to
lanceolate often slightly cordate leaves with crisped or sometimes almost crenulate
margins, more pubescent calyces and ovaries with 2-4 ovules whereas var. grandiflora
was characterised by being nearly glabrous, having more coriaceous leaves with entire
margins and strongly reticulate surfaces, large flowers and often 6 ovules per ovary.
These differential tendencies enabled the material available to Bentham to be sorted
readily into the three varieties but the abundant material now available and field
observations indicate that each variant is far more variable than was apparent to
Bentham and that the distinctions between them break down as the characters
typifying each variant are not necessarily always associated together.

In their typical forms each variant is quite distinctive but specimens exhibiting a
mixture of characters are difficult to place with certainty. For example, specimens
which on the basis of flower size are referrable to var. grandiflora sometimes have as
few as 4 ovules per ovary and linear-oblong leaves characteristic of typical var.
trisperma. Conversely, specimens with 5 or 6 ovules per ovary and narrow-ovate or
elliptic coriaceous leaves with prominent reticulate venation characteristic of var.
grandiflora do not always have large flowers. Flowers in such specimens can be as small
as those in other variants. Specimens with large flowers are not homogeneous in
appearance because of the variation in the other characters. There is no discontinuity
in flower size, pedicel length or in the number of ovules per ovary between the variants
although there is a tendency for var. crispa to have 2 ovules per ovary (sometimes 3 or
4), var. trisperma to have 3 or 4 ovules (sometimes 2) and var. grandiflora 5 or 6 (some-
times 4 or 7). There is no significant difference in the flowering times of the variants.
The main differential tendencies may be seen in Table 1.

Crispa appears to be less well differentiated than grandiflora. It is not uncommon
in the Darling Range to find a range of variation within a population and plants with
the characteristic spreading pubescence and crisped margins to the leaves {crispa)
growing next to plants which lack the spreading pubescence and crisped margins to the

Table 1. The differential tendencies of the three main variants of H. trisperma

Typical trisperma

Typical crispa

Typical grandiflora

Pubescence of
branchlets

Hairs mostly appressed
and antrorse but
sometimes spreading,
often crimped

Hairs spreading, usually
almost at right angles to
branchlet, sometimes
crimped

Hairs mostly appressed
and antrorse, sometimes
crimped, at times
glabrescent

Shape of leaf lamina

Linear-oblong, narrow-
ovate, ovate or elliptic

Ovate, elliptic, elliptic-
oblong, narrow-ovate or
almost rotund

Ovate, narrow-ovate,
obovate, elliptic,
elliptic-oblong,
linear-oblong

Leaf margin

Entire

Crisped to almost
crenulate

Entire, occasionally
almost crenulate

Pubescence of lower
surface of leaf

Hairs more or less
appressed or sometimes
spreading, crimped

Hairs spreading, usually
at right angles to the
lamina

Hairs appressed or
sometimes spreading,
sometimes crimped,
glabrescent

Pedicel length

1-4 mm

2-4-5 mm

2-7 mm

Length of 2 upper calyx
lobes (including tube)

6-5-7-6 mm

6-6-81 mm

6-1-1 1 -7 mm

Length of standard

1 1-5-13-8 mm

10-8-14-8 mm

12-8-20-2 mm

Length of wings

9-1 1 mm

9-2-12-4 mm

9-5-14-8 mm

Length of keel

6-8T mm

8-11-6 mm

6-9-10-3 mm

Number of ovules per
ovary

Usually 3 or 4,
sometimes 2

Usually 2, sometimes 3
or 4

Usually 5 or 6,
sometimes 4 or 7

33

leaves (trisperma). The degree of crisping of the leaf margin ean vary quite markedly
on a single plant.

To some extent the morphological variation in H. trisperma is associated with
geographical distribution. Typical var. trisperma is the most widespread of the
variants occurring from near Gingin in the north as far inland as Kellerberrin
southwards to Busselton and south-east to a little east of Esperance, var. grandiflora is
recorded from Mogumber (with an outlier at Geraldton) southwards to Augusta with
an outlier west of Ravensthorpe, and crispa oceurs chiefly in the Darling Range
north-east, east and south of Perth with outliers at Yanchep, Bindoon, Cunderdin,
Northam, York and near Crossman. The distributional range of var. crispa falls
entirely within that of var. trisperma and that of var. grandiflora overlaps that of var.
trisperma except in the extreme north and south-west.

Although acknowledging the existence of these differential tendencies within H.
trisperma the nature of the variation is such that I do not propose to recognize the
variants formally. Names at varietal rank are available should anyone wish to use
them.

Some specimens of H. trisperma with large elliptic leaves have been confused with
H. elliptica. The two species differ in habit, in the nature of the indumentum on the
lower surface of the leaves and H. elliptica invariably has 2 ovules per ovary whereas
the specimens of H. trisperma which have been confused have 3-6 ovules per
ovary.

Occasional specimens of H. trisperma with linear-oblong leaves are superficially
similar to specimens of H. linearis from eastern Australia. The latter differs in the
nature of the indumentum on the branchlets and lower surfaces of the leaves, in having
smaller flowers and pubescent pods.

5. Hovea pungens Benth. in Endl. etai, Enum. PI. Huegel 37 (1837); Bot. Arch. 2: t. 7
(1837); Paxton, Paxton’s Mag. Bot. 6: 101 (1839); Meissn. in Lehm., PI. Preiss. 1: 78
(1844); Benth., FI. Austral. 2: 176 (1864); Wheeler, FI. Perth Region 269 (1987).
Lectotype (here selected): Western Australia, King Georges Sound, Huegel s.n.
(W).

H. ulicina Meissn., Bot. Zeitung 13: 30 (1855). H. pungens var. ulicina (Meissn.)
Benth., FI. Austral. 2: 176 (1864). Lectotype (here selected): Western Australia,
Drummond, coll. 6 no. 26 (NY; Isolectotype: CGE, MEL).

H. pungens var. major Paxton, Paxton’s Mag. Bot. 10: 51 (1843). Lectotype
(here selected): Mag. Bot. 10: plate facing p. 51. A probable synonym.

Shrub or subshrub to 1-8 m high, single or several-stemmed, often much
branched; branchlets densely clothed with a mixture of straight, curled, slightly
crinkled or asymmetrically biramate appressed to slightly spreading or antrorse hairs.
Leaves sessile or subsessile, typically inserted on the branch at an angle of more than
45° and often almost at right angles to it, the lamina arching outward or spreading
laterally, often reflexed: petiole up to 0-5 mm long; lamina linear to narrowly
elliptic-oblong or ovate-lanceolate, the margins strongly revolute and the lamina
occasionally almost subterete, 0-5-3 cm long including the pungent tip,
0-l-0-3(-0-4) cm wide (excluding the inrolled margins), rigid, narrowed apically and
terminating in a distinct pungent point, rounded or cordate basally, reticulate and
glabrous above, lower surface with hairs usually confined to the midrib, some of the
hairs usually asymmetrically biramate. Stipules setaceous, 2-5-5 mm long, often
diverging, typically persisting for some time. Inflorescence axillary, sessile or on
peduncles up to 1 mm long. Flowers solitary or in 2s or 3s, pedicellate, the pedicels
0-3-0- 9 cm long, densely clothed with a mixture of short curled or asymmetrically
biramate hairs and longer spreading hairs; bracteoles subulate, 1 -5-3-5 mm long,
inserted at the base of or a short distance below the base of the calyx, shorter than the
ealyx-tube, pubescent. Calyx densely clothed with short and longer appressed to
slightly spreading antrorse hairs: 2 upper lobes 5-8-6-9 mm long including the tube
2-3-5 mm long, the 3 lower lobes l-3-2-5mm long. Standard 12-16-8 mm long,
12-16 mm wide, deeply emarginate apically (to 6 mm), purplish-blue with a basal

34

Fig. 5. a — fruiting twig, X 1. b — portion of stem showing point of attachment of a leaf, X 3.

c seed, hilar view, X 6. d — seed, side view, X 6. H. stricta. e — fruiting twig, X 1. f — portion of
stem showing point of attachment of a leaf, X 3. g — seed, side view, X 6. h — seed, hilar view, X 6. a-d
from R. Pullen 9676 (MEL); e and f from M.G. Corrick 9296 (MEL); g and h from B.H. Smith 646B
(MEL).

35

white horse-shoe shaped flare; wings 9-8-12-5mm long, 3-5-6-2mm wide,
purplish-blue; keel 4-2-6-7 mm long, 2-2-7 inm wide. Stamen-filaments 3-8-6-1 mm
long. Ovary 1-1-3 mm long, on a stipe 0-4-1 mm long, 2-ovulate, glabrous. Pods
shortly stipitate, the stipe not exceeding the calyx, obliquely to transversely globular,
ovoid or ellipsoid, 0-6-0-9 cm long, 0-7-0-9 cm wide, glabrous. Seeds elliptic, plunip,
4- 1-5-5 mm long, 2-3-2 mm wide, 2-3 mm thick, dark brown or olive-brown with
paler mottling, hilum elliptic, j to f as long as the seed, aril collar-like with a raised
lateral lip, yellow. (Fig. 5 a-d).

Found in the Irwin Botanical District, the Drummond and Dale subdistricts of
the Darling Botanical District, the extreme west (east of Toodyay) and south (Stirling
Range) of the Avon Botanical District and the Eyre Botanical District of the
Southwestern Botanical Province of Western Australia with an outlier at Coolgardie in
the Coolgardie Botanical District of the Southwestern Interzone as defined by Beard
(1980). Within the Southwestern Botanical Province the distribution of the species is
interrupted by several disjunctions. It occurs in the northern sandplains and heaths
from Burma Rd NW. of Mingenew in the north southwards to the Swan coastal plain
and Darling scarp to the vicinity of Armadale and eastwards in the Wheatbelt to
Wongamine. The species also occurs in the Stirling Ranges and eastwards in the
Transitional Woodland with some further disjunctions to about 100 km E. of
Esperance (Fig. 6). The three most northern occurrences of the species at Burma Rd,
near Lake Indoon and Mt Lesueur are separated by some distance from the nearest
populations near Mogumber and occur within the distributional range of the closely

allied H. stricta. , c

Recorded from sandy soil and coastal limestone in low heath, shallow pockets ot
soil on and around granite outerops, and from laterite, gravel, clay and loamy soils in
heath, jarrah forest and woodland.

Fig. 6. The known distribution of Hovea pungens (A) and H. stricta (•).

36

Representative Specimens (total number examined 157):

Western Australia — Coolgardie, 1902, W.E. Blackall s.n. (PERTH); 10 km due SSE. of Mt Burdett,
2.viii.l983, M.A. Burgman 1637 & S. AfcVee (PERTH); 1-5 km N. of Mt Lesueur, 20.vii.l979, E.A. Griffin
1966 (PERTH); Foot of Bluff Knoll, Stirling Range, x. 1984, R. & M. Hamilton 32 (MEL, PERTH);
Wongamine Reserve, 19 km NW. of Northam, 19.viii.l983, B.H. Smith 243 (AD, CBG, HO, MEL,
PERTH).

Notes:

H. pungens is very closely allied to H. stricta. Bentham (1864) differentiated the
two species on the basis of the shape of the leaf apices, those of H. pungens being
described as ‘pungent-pointed’ in contrast to those of H. stricta which were described
as ‘obtuse with a small scarcely pungent point’. This may well have been true of the
limited material available to Bentham but the material now available indicates that
this character does not enable the two species to be differentiated.

The two species may be distinguished chiefly by the disposition of the leaves on
the branches, the shape of the leaves and by the seeds. Very few seeds of H. stricta were
available for study but as far as can be judged the seeds of the two species differ in
colour, in the size of the hilum and the shape of the aril. The seeds of H. stricta are a
uniform yellowish or olive-brown, the hilum is less than j as long as the seed and the
aril is shortly columnar whereas those of H. pungens are mottled, the hilum is about j
to I the length of the seed and the aril is not as tall. It is unfortunate that seeds are
seldom available as it restricts the use of seed as a distinguishing character. The almost
invariable absence of seeds means that the two species are differentiated usually on
vegetative characters. Despite this, most specimens can be sorted into two species
quite readily as each has a different ‘look’ about it although the differences are difficult
to express in words.

The leaves in H. pungens are sessile or on petioles up to 0-5 mm long and typically
are inserted on the branch at an angle >45° and often almost at right angles to it, the
lamina arching outwards or spreading laterally and frequently slightly reflexed. In H.
stricta the leaves are shortly but distinctly petiolate on petioles 0-75-T5 mm long and
typically are inserted on the branch at an acute angle (<45°), the lamina arching
outwards and upwards with the apex pointing towards the apex of the branch. These
differences in leaf shape are not absolute but the leaf lamina in H. stricta tends to be
broader than in H. pungens. These apparently trivial characters do nevertheless enable
most specimens to be sorted into the two species without much difficulty in the
absence of seeds.

H. stricta tends to have a denser shaggier indumentum on many of its parts and
the stipules are smaller and less persistent than in H. pungens. In addition, tfie growth
form of the two species tends to differ, H. stricta often being smaller and more
sparingly branched.

The application of the name H. pungens var. major Paxton, published a year
before H. stricta, is not entirely certain. The plant illustrated is neither typical of H.
pungens nor of H. stricta which is not surprising seeing that it was grown under glass.
The diagnostic characters used to differentiate the two species are not readily evident
from the illustration but it seems probable that the plant was referrable to H. pungeris
rather than to H. stricta. A cultivated specimen of H. pungens from South Australia
shows an approach to the plant illustrated. Rather than treat H. pungens var. major as a
name of uncertain application it is regarded here as a probable synonym of H.
pungens.

6. Hovea stricta Meissn. in Lehm., PI. Preiss. 1: 79 (1844); Benth., FI. Austral. 2: 176
(1864). Lectotype (here selected): Western Australia, ‘in arenosis sylvae districtus
Sussex’, Dec. 1839, Preiss J05 7 (LD; Isolectotype: MEL, NY).

H. stricta var. major Meissn., Bot. Zeitung 13: 30 (1855). Lectotype (here
selected): Western Australia, Drummond coll. 6 no. 27 (NY; Isolectotype BM, CGE,
K, MEL, NSW, W).

Shrub to 1 m high, single or several-stemmed, usually sparingly branched;
branchlets densely clothed with a mixture of straight, curled, slightly crinkled or

37

twisted villous or asymmetrically biramate spreading to slightly antrorse hairs. Leaves
shortly but distinctly petiolate, typically inserted on the branch at an acute angle (less
than 45°), the lamina arching outward and upward with the apex pointing towards the
apex of the branch but sometimes inserted almost at right angles to the branch and
spreading laterally: petiole 0-75- 1-5 mm long; lamina linear to narrowly
elliptic-oblong or ovate-lanceolate, the margins strongly revolute, (0-5-)l-2-4-2 cm
long including the pungent tip, 0-1 5-0- 75 cm wide (excluding the inrolled margins),
rigid, obtuse apically and scarcely mucronate or narrowed and with a distinct pungent
point, rounded or cordate basally, reticulate and glabrous or with scattered hairs
above, lower surface with scattered hairs throughout or hairs confined to the midrib or
sometimes glabrous, some of the hairs usually asymmetrically biramate. Stipules
setaceous, up to 2-5 mm long, sometimes not persisting. Inflorescence axillary, sessile
or on peduncles up to 1 -2 mm long. Flowers solitary or in 2s or 3s, pedicellate, the
pedicels 0-4-1 • 1cm long, densely clothed with a mixture of short curled or
asymmetrically biramate hairs and longer spreading hairs; bracetoles subulate,

1 -2-3 mm long, inserted at the base of or a short distance below the base of the calyx,
shorter than the calyx-tube, pubescent. Calyx densely clothed with short hairs and
longer straighter or crisped spreading antrorse hairs: 2 upper lobes 4-4-7-5 mm long
including the tube 2-5-4 mm long, the 3 lower lobes l-6-2-5mm long. Standard
14-5-17 mm long, 11-8-17-3 mm wide, deeply emarginate apically (to 6-5 mm),
purplish-blue with a basal white horseshoe shaped flare; wings 11-5-13-5 mm long,
5-6-9 mm wide, purplish-blue; keel 7-7-7 mm long, 2-4-2-9 mm wide.
Stamen-filaments 4-2-6-2 mm long. Ovary 1-2-1 -5 mm long, on a stipe ± 1 mm long,
2-ovulate, glabrous. Pods shortly stipitate, the stipe not exceeding the calyx, globular,
ovoid or transversely ellipsoid, 0-9-1 -1 cm long, 0-9-1 cm wide, glabrous. Seeds
elliptic, plump, 4-4-5 mm long, 2-8-3 mm wide, 2-5-3 mm thick, uniform yellowish-
or olive-brown, hilum oval or elliptic, less than j the length of the seed, aril collar-like,
shortly columnar, with a raised lateral lip, yellow (Fig. 5 e-h).

Confined to the Irwin Botanical District and the Drummond subdistrict of the
Darling Botanical District of the Southwestern Botanical Province of Western
Australia as defined by Beard (1980) where there are two disjunct centres of
distribution: one in the northern sandplains and heaths from west of Three Springs m
the north southwards to the Moore River National Park, and the other from the
Busselton area and Cape Naturaliste (Fig. 6). With one exception, all of the specimens
from the southern centre of distribution were collected last century and cannot be
localised accurately so that the extent and range of the populations is not clear.

In the northern part of its range the species occurs usually on white sand or sand
over laterite in low heath, scrub or shrubland. The ecological preferences of the
southern populations are not known as none of the specimens is accompanied by
ecological data.

Representative Specimens (total number examined 44):

Western Australia — 4-8 km SW. of Mt Lesueur, 24.vii.1969, M.I.H. Brooker 1938 (PERTH);
Coorow-Green Head Rd, 16 km E. of Rose Thompson Rd, 6.ix.l984, D.B. Foreman 565 (MEL, PERTH);
48 km W. from Three Springs, 27.viii.1948, C.A. Gardner 9137 (PERTH); 16 km S. of Regans Ford,
26.viii. 1 964, K. Newby 1386 (PERTH); Vasse River, Oldfteld(MBL 666572); 39 km NNE. of Jurien on Green
Head-Jurien Rd, 28.xi.1974, R. Pullen 9676 (CANB, MEL).

Notes: . . ■

The arils of the seeds of//, stricta are somewhat atypical for the genus in being less
than three times longer than broad. This raises questions about the placement of the
species in Hovea or the usefulness of this character in diagnosing the genus.
Admittedly this observation is based upon the seeds of only one specimen of//, stricta
which highlights the paucity of fruiting specimens in herbaria, especially those with
mature pods and seeds, in many genera of Fabaceae and other families, a fact lamented
by Bentham ( 1 864) when discussing Acacia. Fruiting specimens of Hovea are collected
far less frequently than flowering specimens, probably because the fruits are generally

38

inconspicuous and the plants are overlooked in contrast to the bright flowers which
attract attention. More mature seeds of H. stricta are required to confirm whether or
not the arils are always as observed on the fruiting specimen referred to above. H.
stricta is very closely allied to H. pungens, the seeds of which are typical of Hovea. In
view of this, and in the absence of any other correlated differential character, it would
appear inappropriate to consider excluding H. stricta from Hovea on the basis of its
seeds being different.

For a discussion of the differences between H. stricta and H. pungens see the notes
under the latter species.

ACKNOWLEDGEMENTS

I am most grateful to Mr N. S. Lander, Mr C. R. Dunlop and Dr G. P. Guymer for
answering enquiries and for photographing type specimens while serving as Australia
Botanical Liaison Officer at the Elerbarium, Royal Botanic Gardens, Kew; to Miss
H. M. Cohn, Librarian, National Herbarium of Victoria, for checking State Library
records relating to the holdings of Bull. Soc. Imp. Naturalistes Moscow, to my
colleagues Mrs M. G. Corrick, Dr D. B. Foreman, Mrs R. A. Kilgour and Dr P. S.
Short and to Mr & Mrs B. H. Smith, Manmanning, Mr & Mrs R. Hamilton,
Kalamunda, Mr & Mrs C. E. Woolcock, Portland, and Mr C. W. Huggins, Anglesea,
for making special collections; to my colleague Mr R. H. Barley for preparing the black
line drawings which accompany the text; to Ms J. Hurse for technical assistance; to Dr
M. D. Crisp, Herbarium, Australian National Botanic Gardens, Canberra, for the loan
of colour transparencies of type material of Daviesia? acanthoclada Turcz.; to the
Directors/Curators of AD, BM, BRI, CGE, E, HBG, HO, K, LD, NSW, NY, PERTH,
PR, PRC and W for the loan of specimens and/or for working facilities in their
institutions; to Mrs J. Thomas for typing the manuscript; and for an Australian
Biological Resources Study grant which greatly facilitated this project.

REFERENCES

Beard, J. S. ( 1 980). A new phytogeographic map of Western Australia. Western Australian Herb. Res. Notes 3:
37-58.

Bentham, G. (1864). Leguminosae in Flora Australiensis, Vol. 2. (Lovell Reeve & Co: London.)

Manuscript received 8 March 1988

A REVISION OF PODOTHECA Cass. (ASTERACEAE:
INULEAE: GNAPHALIINAE)

by

P. S. Short*

ABSTRACT

Short, P. S. A revision of Podotheca Cass. (Asteraceae: Inuleae: Gnaphaliinae).
Muelleria 7(1): 39-56 (1989). The endemic Australian genus Podotheca Cass, is
revised. Six species are recognized and three of them, P. pritzelii P. S. Short, P. uniseta
P. S. Short and P. wilsonii P. S. Short are described as new. P. fuscescens (Turcz.)
Benth. and P. pollackii (F. Muell.) Diels are excluded from the genus.

GENERIC HISTORY

Conservation of the Name Podotheca Cass

Labillardiere ( 1 806) described and named for the first time the genus Podosperma
Labill. He attributed a single species, P. angustifolia Labill., to the genus and based his
description on material gathered in Western Australia. The previous year the name
Podospermum DC. (1805) had been erected. Both names apply to different genera of
the Compositae and with the similarity of the names confusion soon reigned. Lessing
(1832) incorrectly spelt Podospermum DC. as "Podosperma' and this mistake was
perpetuated by Kunth (1838) and Schultz ( 1834). Prior to the aforementioned mistake
Cassini (1822) had erected the name Podotheca Cass., indicating that it was the
equivalent of Podosperma Labill. No reason for the name change was given and the
required new combination for the species was not made. Four years later Cassini
(1826) suggested that the name Phaenopoda Cass, be adopted instead oi Podosperma
and Podotheca. Lessing (1832) adopted the name Podotheca and effected the
combination Podotheca angustifolia (Labill.) Less.

Graham (1842) described Podotheca gnaphalioides Grab, and Steetz (1845) also
adopted the name Podotheca. Bentham (1867) followed suit and in citing generic
synonyms referred to Lophoclinium Endl. (1843), Phaenopoda Cass, and "Podosper-
mum Labill.’.

Podosperma Labill., not Podotheca Cass., was subsequently used by many authors
(e.g. Mueller 1882, 1889 — as Ewart 1931, Black 1957; Curtis 1963).
Others (e.g. Hoffman 1894; Rodway 1903; Maiden & Betche 1916) retained
Podotheca.

Because of the confusion that had occurred with both Podospermum DC. and
Podosperma Labill. Eichler (1964) proposed that the name Podotheca be conserved
against Podosperma Labill. [Submittance of this proposal was partly influenced by the
recognition in Australia of species referable to Podosperma and Podospermum, the
latter being represented by the weed P. laciniatum (L.) DC. It now seems that
Podospermum is generally considered to be synonymous with Scorzonera L.] The
Committee for Spermatophyta (1967) reported that in their opinion Podosperma
Labill. should be regarded as illegitimate (Art. 75) due to the similarity to
Podospermum DC., thus making the conservation of Podotheca unnecessary.
However, it was also felt that, as Art. 7 5 and the examples cited within left the question
of the legitimacy of Podosperma open to dispute, it was in the interests of
nomenclatural stability to recommend conservation of Podotheca against
Podosperma. The General Committee of Botanical Nomenclature (1968) approved
the proposal.

*National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

39

40

Generic Delimitation

Prior to Bentham’s ( 1 867) account of Podotheca three species had been referred to
the genus, i.e. P. angustifolia, P. gnaphalioides and P. pygmaea A. Gray (1851).
Bentham transferred two further species, Ixiolaena chrysantha Steetz and Helipterum
fuscescens Turcz., to Podotheca. He noted that ‘the genus is limited to Australia,
differing from Helipterum chiefly in the stipitate achenes, and generally in the
involucre, which however is less foliaceous in P. fuscescens than in the other species’
(Bentham 1867, p. 601). Mueller (1882) retained the name Podosperma in preference
to Podotheca and described a further new species, P. pollackii F. Muell. In the same
publication he suggested that P. fuscescens, because of the outer scarious bracts of the
involucre, should again be returned to Helipterum. Diels (1904) effected the
combination Podotheca pollackii (F. Muell.) Diels.

Despite Mueller’s comment that P. fuscescens be referred to Helipterum the
species is commonly retained in Podotheca. All species of Podotheca occur in Western
Australia and Grieve & Blackall (1975) recognized six species, i.e. P. angustifolia, P.
chrysantha, P. fuscescens, P. gnaphalioides, P. pollackii and P. pygmaea. In this
revision P. fuscescens and P. pollackii are excluded from Podotheca s. str. , three further
species {P. pritzelii, P. uniseta and P. wilsonii) are described as new, and P. pygmaea is
reduced to synonymy under P. angustifolia.

Merxmiiller et al. (1977) referred Podotheca to their ‘group 16’ or ’’Schoenia
group’ of the Gnaphaliinae sensu amplo. This group is characterized by the triangular
hairy appendage of the style arms and includes some members of Australian
Helichrysum Miller and Helipterum DC. plus genera such as Millotia Cass, and
Waitzia Wendl. Podotheca s. str. is readily distinguished from other members of this
group by the involucral bracts, which are arranged in several series with the outer ones
leaf-like, the pappus of usually one, five or ten bristles and the long, bisexual florets of
the large capitula. The fruit has a prominent stipe (Fig. 2a) but this feature is not
exclusive to species of Podotheca s. str. Chromosome number determinations by
Turner (1970) suggest a base number of x= 13 for the genus.

Podotheca fuscescens is excluded from Podotheca s. str. on a number of grounds.
The involucral bracts are quite dissimilar to those of Podotheca s. str. The outer
leaf-like bracts are few in number and the inner bracts have white, opaque tips which
are absent in species of Podotheca s. str. Other differences occur in the style
appendages, which are more or less truncate and long papillate, and the pappus of c.
12-14 plumose bristles. P. fuscescens has strong affinities with Helipterum
oppositifolium S. Moore and H. strictum (Lindl.) Benth. (P. G. Wilson, in litt.,
1987).

Podotheca pollackii is readily distinguished from members of Podotheca s. str. by
virtue of the spike-like arrangement of the capitula. As with H. fuscescens the
involucral bracts of this species differ from those in Podotheca s. str. and the fruit are
anatomically different. In Podotheca s. str. the vascular bundles of the pericarp are
oblique to the cotyledons and the sclerenchymatous layer in the pericarp is mainly one
cell wide. In P. pollackii the vascular bundles are in the plane of the adaxial surface of
the cotyledons and the sclerenchymatous layer is 2-4 cells wide. Unlike members of
Podotheca s.str. fresh specimens produce a fetid odour when the leaves are crushed and
the anthers are black. The style and anther morphology suggest the retention of P.
pollackii in the ‘Schoenia group’. Its strongest affinities are with Helipterum battii F.
Muell., H. charsleyae F. Muell. and H. spicatum (Lindl.) Benth. (P. G. Wilson, in litt.,
1987).

MATERIALS AND METHODS

Descriptions of taxa were made from dried herbarium material and from
specimens stored in 70% ethanol. Shapes were defined using the terms given by the
Systematic Association Committee for Descriptive Terrninology (1962).

Specimens were examined from the following herbaria: AD, BM, E, K, LD, MEL,
PERTH, S and W.

41

Pollen: ovule ratios (P/Os) were obtained following the method previously
outlined (Short 1981). In all but P. chrysantha P/Os were obtained on a population
basis, the number of pollen grains being determined for at least 1 5 florets, with each
floret sampled from different plants. Measurements pertaining to anther characteris-
tics (see Short 1985, fig. 1) were also obtained on a population basis.

Fruit sections were obtained following the fixation of dry, mature fruit with 5%
glutaraldehyde in Pipes buffer (O’Brien & McCully 1981) and the subsequent
infiltration and embedding in L. R. White (London Resin Co.). Sections were stained
in toluidine blue (pH 4-5).

TAXONOMY

Podotheca Cass., Diet. Sci. Nat. 23: 561 (1822), nom. cons.-. Less., Syn. gen. Compos.
272 ( 1832); DC., Prod. 6: 1 59 ( 1838); Steetz in Lehm. PI. Preiss. 1 : 448 ( 1 845); Benth.,
FI. Austr. 3: 600 (1867) p.p., excluding P. fuscescens-, Benth. in Benth. & Hook, f..
Genera PI. 2: 315 (1873) p.p.; Hoffman in Engler & Prantl., Naturl. Pflanzenfam.

I V(5): 1 90 ( 1 890); Eichler, Regnum Veg. 34: 62 ( 1 964); Taxon 16:229(1967); Grieve
& Blackall, W. Aust. Wildfls 824 (1975); Cooke in Jessup & Toelken, FI. S. Aust. 3:
1573 (1986); Lander in Marchant et ai, FI. Perth Region 699 (1987). — Podosperma
LabilL, Nov. Holl. PI. Sp. 2: 35, t. 177 (1806); J. M. Black, FI. S. Aust. 1st ed. 636
(1929), 2nd ed. 914 (1957); W. M. Curtis, Stud. FI. Tas. 341 (1963); J. H. Willis,
Handb. PI. Viet. 2: 719 (1973). — Phaenopoda Cass., Diet. Sci. Nat. 42: 84 (1826),
nom. illeg. Type: Podotheca angustifolia (Labill.) Less.

Lophoclinium Endl., Bot. Zeitung (Berlin) 1: 457 (1843), p.p. excluding L. album.
Lectotype: Not chosen, see note 2 under P. angustifolia.

Annual herbs. Major axes decumbent to erect, sometimes cottony, with flat,
septate hairs and/or glandular hairs. Stem simple or forming major branches at basal
and/or upper nodes. Leaves mainly alternate but the lowest pair(s) opposite, all leaves
sessile, sometimes succulent, entire, leaf bases ± decurrent, all leaves with fiat, septate,
non-glandular hairs usually mixed with glandular hairs. Capitula homogamous, ovoid
to lanceoloid or obovoid to very broadly obovoid or ± oblong. Involucral bracts
14-c. 60, multiseriate, the outermost leaf-like, the inner mainly hyaline. Receptacle
flat or concave, glabrous, base of pedicels prominent. Florets c. 10-300 per capitulum,
all tubular, bisexual, 5-merous; style appendix triangular, papillose; stamens 5, anthers
tailed and each with a sterile, apical appendage. Cypselas homomorphic, obovoid,
pubescent; pericarp with a layer of sclerenchyma which is usually one cell wide but
sometimes two cells wide around the two vascular bundles, the bundle orientation
medial or oblique; testa of 2 ± distinct layers, the outer with crystals, vascular bundles
absent; endosperm one cell wide; carpopodium oblique; pedicel or stipe prominent.
Pappus usually of 1 , 5 or 10 bristles c. the length of the corolla tube, bristles barbellate
to plumose, joined at the base.

Chromosome numbers: n = 1 3, 26.

Distribution (Fig. 1):

All six species occur in Western Australia and five are endemic to that state. P .
angustifolia extends across the southern part of the mainland and reaches islarids in
Bass Strait. Three species in Western Australia seem to only occur on the margins of
salt lakes. Two, P. pritzelii and P. uniseta, are restricted to the Avon Lake system and
the Monger Lake system respectively (Bettenay & Mulcahy 1972; Mulcahy & Bettenay
1972). (Lake Moore, as suggested in 1973 by Beard, is considered to belong to the
Monger system.) In contrast P. wilsonii occurs in three different Drainage Divisions,
the Murchison, South-West and Eucla Divisions (Mulcahy & Bettenay l.c.).

Reproductive Biology, Cytology and Evolution:

The use of pollen: ovule ratios (P/Os) in the determination 9 f plant breeding
systems has been previously discussed (Short 1981). The values obtained for five of the
six species of Podotheca, ranging from one to several thousand (Table 1), suggests that

42

Fig. 1. Distribution of species of Podotheca.

they are outbreeders (i.e. cross-fertilization is common). In contrast P. angustijolia,
with an average P/O of 1 84, is considered to be an inbreeder (i.e. self-fertilization is
common). The low P/O in P. angustifolia is correlated with inconspicuous florets,
compared to other species of Podotheca, and small anthers (Table 1).

Chromosome number determinations are only available for three of the six
species (Table 1). All were recorded by Turner (1970) and suggest a base number of
X = 1 3 for the genus. That polyploidy is recorded in P. angustifolia and no other species
is not surprising when it is considered that it is the only one classed as an
inbreeder.

43

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By virtue of the shape of the capitulum two groups can be discerned within
Podotheca. Thus P. chrysantha and P. wilsonii, with obovoid to very broadly obovoid
(rarely ± oblong) capitula, are readily distinguished from all other species which have
ovoid or narrowly ovoid capitula. The dense vestiture of stalked, glandular hairs on
the bracts also suggests that P. chrysantha and P. wilsonii are more closely related to
one another than other species in the genus although such a vestiture does occur in a
few specimens of/*, gnaphalioides. The four remaining species are closely related, with
three (P. gnaphalioides, P. pritzelii and P. uniseta) mainly dilfering in the number of
pappus bristles, the presence or absence of succulent bracts and the distribution of
septate hairs. Ecological differences also exist, with P. uniseta and P. pritzelii occurring
in more saline habitats than P. gnaphalioides. (For further notes on morphological and
ecological differences see under treatment of species.) P. angustifolia, although clearly
having strong affinities with the proceeding three species, is readily distinguished by
virtue of its inbreeding nature, correlated as it is with small florets and anthers. The
widespread occurrence of P. angustifolia is consistent with the distribution observed
for many other inbreeding species of the Inuleae (Short 1981).

On several previous occasions (Short 1981,1 986) attention has been drawn to the
importance of the salt lake systems of south-west Western Australia to speciation in a
number of inuloid genera. The apparent restriction of P. pritzelii and P. uniseta to
separate lake systems further highlights their importance.

Key To Species of Podotheca

1 . Pappus bristles 5 per floret

2. Capitula obovoid to very broadly obovoid, rarely ± oblong; outer involucral
bracts with stalked, glandular hairs not flat, septate hairs; pappus bristles usually

white or pale yellow but sometimes the upper part pink 2. P. wilsonii

2. Capitula ovoid to narrowly ovoid; bracts glabrous or with flat, septate hairs,
sometimes with stalked glandular hairs; pappus bristles white or pale yellow,
never pink

3. Florets inconspicuous, barely exerted above the bracts; anthers 0-4- 1 T 5 mm

long 6. P. angustifolia

3. Florets conspicuous, exerted well beyond the bracts; anthers 1-24-2-61 mm
long

4. Leaves and bracts at most semi-succulent, dark green or purple; glandular
hairs usually present on major axes; outer involucral bracts usually with

septate hairs, rarely glabrous 3. P. gnaphalioides

4. Leaves and bracts succulent, pale-green, rarely purplish; glandular hairs
absent from major axes; outer involucral bracts glabrous . 4. P. pritzelii
1 . Pappus bristles not 5 per floret

5. Pappus bristles 1 per floret 5. P. uniseta

5. Pappus bristles (9)10(1 1) per floret 1. P. chrysantha

1. Podotheca chrysantha (Steetz) Benth., FI. Austr. 3: 602 (1867); Grieve & Blackall,
W. Aust. Wildfls 824 (1975); Lander in Marchant et ai, FI. Perth Region 700 (1987).
— Ixiolaena chrysantha Steetz in Lehm., PI. Preiss. 1: 459 (1845). — Podosperma
chrysantha (Steetz) F. Muell., Fragm. 12: 22 (1882). Type: ‘In arenosis sylvae prope
oppidulum Perth, d. 23. Sept. 1839. Herb. Preiss. No. 105’. Lectotype (here chosen):
Preiss 105, In Nova Hollandia (Swan River Colonia), in arenosis sylvae prope
oppidulum Perth, s. dat. (MEL 1553907, ex herb. Steetz). Isolectotypes: GH, LD,
MEL 1553905 (ex herb. Sonder), MEL 1553906 (ex herb. Sender), MEL 1543871, P
(three sheets, one ex herb. E. Drake, one ex herb. Schultz-Bip.), S, W (three sheets).
(See note 1 below.)

Annual herbs. Major axes erect, 6-35 cm long, with stalked, glandular hairs,
brown orbrown purple. Leaver lanceolate or ± linear, 0-5-8-5 cm long,c. OT-0-65 cm
wide, apex often incurved, with stalked, glandular hairs, green. Capitula obovoid to
very broadly obovoid, 1-2-5 cm long, 0-4-3 cm diam. Involucral bracts 14-55 per

45

Fig. 2. a — Carpopodium of P. angustifolia (Short 2689). b — Apex of fruit of P. uniseta showing the
single pappus bristle (Short 2642).

Fig. 3. Transverse section of fruit of P. angustifolia (Short 2689). Sclerenchyma = hatching, testa = dots,
endosperm = black shading.

capitulum, narrowly elliptic, lanceolate, oblanceolate, narrowly obtrullate or narrowly
to linear triangular, 6-5-14 mm long, 0-8— 4-2 mm wide; outer bracts herbaceous, the
surface with stalked, glandular hairs, sometimes with narrow hyaline margins with
long-ciliate hairs; inner bracts hyaline except for an opaque midrib, glabrous or with
long-ciliate hairs on the margins. Florets 10-320 per capitulum, yellow; corolla tube
7-5-12-5mm long. Stamens 5; anthers l-74-2-24mm long; microsporangia
1.24-1-77 mm long; apical appendages 0-39-0-52 mm long. Pollen grains c. 500-600
per anther. Cypselas 2-3-3 mm long, 0-6-0-9 mm diam. Pappus bristles (9-)10(-l 1),
smooth near the base, otherwise barbellate.

Chromosome number: n= 13 (Turner 1970).

46

Distribution (Fig. 1); ,

Western Australia between latitudes c. 29° 30' S. and 35° S. and west of longitude
c. 1 1 7° E. A collection by Oldfield (K) is labelled as coming from the Murchison River
but this seems to be erroneous.

Ecology: .. ^ j

Favours sandy soil. Collectors’ notes include ‘low woodland . . . ot Banksia, ridge
of light yellow sand’, ^Banksia - coastal Blackbutt association’ and ‘closed heath,
shallow sand over limestone on ridge’. Flowers from late August to December.

1. The selection of MEL 1553907 as the lectotype of Ixiolaena chrysantha is
consistent with the argument previously put (Short & Sinkora 1988) that in the case of
names originally coined by Steetz that specimens in his own herbarium should
generally be chosen as lectotypes.

Selected Specimens Examined (Total c. 50):

Western Australia — near Boongarra, 17.x. 1978, Hnatiuk 780145 PERTH); Yanchep National Park,
17 X 1963 James 19 (PERTH); 18 km E. of Lancelin, 17.X.1981, Keighery 4140 (PERTH); Bayswater,
6.X.1897, Morrison s.n. (E, K, MEL 1543881, PERTH); Capel, 18.ix.l949, Royce 3122 (PERTH).

2. Podotheca wilsonii P. S. Short, sp. nov.

Herba annua. Axes majores ascendentes vel erecti, 7-5-45 cm longi, pilis stipitatis glandulis. folia
linearia vel lanceolata, 0-5-13-5 cm longa, O-l-l-l cm lata, apicibus saepe mcurvatis, pilis stipitatis
glandulis, viridia usque purpurea. Capitula obovoidea usque perlate obovoidea, raro ± oblonga,
l-5-3-5cm longa, 0-7-3 cm diametro. Bracteae involucrales 26-75, ovatae usque lanceolatae,
oblanceolatae, anguste trullati usque trullati, triangulares vel ± lineares, 7-6-24 mm longae, 1^5-5 mm
latae; bracteae exteriores herbaceae, pilis stipitatis glandulis, saepe marginibus angustis hyalinis pnis
longis ciliatis ferentibus; bracteae interiores hyalinae praeter costa opaca, glabrae. Floscuti 44-294,
lutei- corolla tubus 14-5-25-1 mm longa. Stamina 5; antherae 1-96-2-46 mm longae, unaquaeque
pollinibus c. 400-660. Cypselae 1 -7-2-1 mm longae, 0-55-0-65 mm diametro. Pappi setae 5 laeves
usque barbellatae in parte inferna 1/3-1/2, in supera 1/2-2/3 subplumosae, plerumque albae vel
luteolae sed interdum in supera 1/3 roseae.

Holotypus: FLammersley Lakes, c. 16 km S. of Mt Jackson Homestead, f- 30

11 9° 01' E. In sand amongst samphire, and Frankenia. 7.x. 1983, Short 1995

(MEL 689073). Isotypi: AD, BRI, CANB, CBG, HO, K, NSW, NT, PERTH.

Annual herbs. Major axes ascending to erect, 7-5-45 cm long, with stalked
glandular hairs, purple. Leaves linear or lanceolate, 0-5-1 3-5 cm long, O-l-l-l cm
wide, the apex often incurved, with stalked glandular hairs, green to purple. CapiMa
obovoid to very broadly obovoid, rarely ± oblong, 1 -5-3-5 cm long, 0-7-3 cm diarn.
Involucral bracts 26-75 per capitulum, ovate to lanceolate, oblanceolate, narrowly
trullate to trullate, triangular or ± linear, 7-6-24 mm long, 1-5-5 mm wide, outer
bracts herbaceous, the surfoce with stalked glandular hairs, often with narrow hyaline
margins with long-ciliate hairs; inner bracts hyaline except for the opaque midrib,
glabrous. Florets 44-294 per capitulum, yellow; corolla tube 14-^5-25-1 mm long
Stamens 5; anthers 1-96-2-46 mm long; microsporangia 1-45-1-88 mm long; apical
appendages 0-35-0-62 mm long. Pollen grains c. 400-660 per anther. Cyp.se/fl3
1 -7-2- 1 mm long, 0-5-0-6 mm diam. Pappus of 5 bristles, bristles smooth to barbellate
in the lower 1/3-1/2, the upper 1/2-2/3 subplumose, commonly white or pale yellow
but sometimes the upper c. 1/3 pink. (Fig. 4)

Distribution (Fig. 1): , 3i»3n'q and

Western Australia. Occurs between latitudes c. 25 30 S. and 31 30 b. and

longitudes 118°E. and 124°30'E.

IE C L Y^ *

Collectors’ notes suggest that the species is restricted to saline, generally sandy
soil. A single collection records that specimens were collected in a clay depression.

47

051 m 0£l OZl oil QOl 06

ue 0/ 1)9 (ft ot oe 0^ 01

Mli'NVl HIKH\KIIM''I WtloKlNiMl

N S I IO\ \1 HI «H\MH \1 (»>

\ K lOKJ \ (Ml 1 <, VI t K VI I \

Fig. 4. Holotype sheet of P. wilsonii (Short 1995).

Notes include ‘Growing in red-brown sand in open areas between shrubs oi Melaleuca.
On edge of salt lake’ and ‘In sand amongst samphire, Atriplex and Frankenia'.

Notes:

1. The specific epithet honours Paul G. Wilson of PERTH.

2. Three of the thirteen collections examined contain plants in which the upper

48

part of the pappus bristles are pink. A single collection, Demarz 4613, has some
specimens with pink bristles, others with yellow bristles.

3. Podotheca wilsonii has close affinities with P. chrysantha but the latter species
has ten, rarely nine or eleven, bristles per floret, is always an erect herb and displays a
preference for non-saline habitats.

Selected Specimens Seen (Total 1 3):

Western Australia — Lake Carey, 7.x. 1973, Demarz 4613 (PERTH); 41 miles N. of Bulga Downs,
24.ix.1975, Demarz 5642 (PERTH); c. 16 km S. of Mt Jackson Homestead, 5.xi.l983, Short 2298 & Haegi
(AD, MEL, PERTH); 30 km NE. of Nambi Homestead, 28.viii.1968, Wilson 7482 (PERTH); southern
margins of Lake Rason, 13.ix.l984, Wilson 72/77 (PERTH).

3. Podotheca gnaphalioides Grab., Bot. Mag. t. 3920 (1842); Steetz in Lehm. PI.
Preiss. 1; 449 (1845); Benth., FI. Austr. 3; 601 (1867); Grieve & Blackall, W. Aust.
Wildlfs 824 (1975); Lander in Marchant et ai, FI. Perth Region 699 (1987). —
Podosperma gnaphalioides (Grab.) F. Muell., Fragm. 12: 22 (1882). Type: ‘raised at
the nursery garden of Messrs. James Dickson & Sons, Edinburgh, in spring, 1841, from
a collection of Swan River seeds, communicated the year before by Mr. Murray,
Lintrose . . . struck from cuttings by Mr. Kelly . . . of Messrs. Dickson’s establishment.’
Lectotype (here chosen): Anon, s.n.. Swan River, cult., s. dat. (K). (See note 1
below.)

Lophoclinium manglesii Endl., Bot. Zeitung (Berlin) 1: 457 (1843). Type: ‘Nova
Hollandia austro-occidentalis’. Possible Syntypes and Isosyntypes: Preiss 107.
LD, MEL 1543867 (ex herb. Sond.), MEL 691442 (ex herb. Steetz), P, W. (See note 2
under P. angustifolia.)

Podotheca pygmaea A. Gray, Hook. J. Bot. Kew Gard. Misc. 4: 227 ( 1 85 1 ). Type:
‘Swan River, Drummond’. Lectotype (here chosen): Drummond 64, Swan River, N.
Holl., s. dat. (K). (See note 2 below.)

Annual herbs. Major axes ± prostrate to erect, 6-55 cm long, with flat, septate,
non-glandular hairs and often stalked glandular hairs, green to purple. Capitula ovoid
to lanceoloid, 2-5 cm long, 0-3-1 -5 cm diam. Involucral bracts 11-43 {c. 60) per
capitulum, ± ovate to lanceolate or narrowly to linear triangular or ± oblanceolate or
narrowly obtrullate, 8-6-36-7 mm long, 0-9-3-8 mm wide; outer bracts herbaceous,
sometimes semi-succulent, dark green or purple green, usually with narrow, hyaline
margins with long-ciliate hairs, the outer surface with flat, septate hairs and/or stalked,
glandular hairs, rarely glabrous; inner bracts hyaline except for an opaque midrib,
glabrous or with, long-ciliate hairs on the margins. Florets 10-204 per capitulum,
yellow or yellow-orange, corolla tube 20-7-30 mm long. Stamens 5; anthers
1-24-2-61 mm long; microsporangia 1-37-2-14 mm long; apical appendages
0-37-0-66 mm long. Pollen grains c. 300-500 per anther. Cypselas 1 -9-2-7 mm long,
0-5-0-75 mm diam. Pappus of 5 bristles, each bristle usually smooth at the base,
grading to plumose, sometimes barbellate or ± plumose near the base.

Chromosome number: n= 13 (Turner 1970).

Distribution (Fig. 1):

South-west of Western Australia, including Dirk Hartog Island.

Ecology:

Occurs in a variety of habitats and tends to favour sandy soils but has also been
found growing in clay loam. It apparently has some salinity tolerance, with plants
having been gathered on the edge of saline depressions. However it is generally
restricted to areas above the samphire zone. Collectors’ notes include: ‘Growing in
sand with Melaleuca, occasionally just extending to area with Gunniopsis on edge of
saline depression’, ‘closed heath, shallow sand over limestone’, ‘open forest
Eucalytpus wandoo, clay loam in drainage line’, ‘Open heath, orange-brown sandy
loam over outcropping ferrugineous sandstone’, ‘Sandplain with heath c.lm tall
dominated by Leptospermum sp., Casuarina cutivalvus and Acacia spp.’, ‘In very

49

sandy loam on edge of granite outcrop’ and ‘mallee eucalypt scrub, red-brown

loam’.

Notes:

1 . It could be expected that type material examined by Graham is housed at E.
However, a visit to E in August 1985 failed to reveal syntype material of P.
gnaphalioides, there only being a photograph of the K specimen selected as the
lectotype.

2. When describing P. pygmaea Gray referred to the ‘whole plant scarcely above
an inch high’, suggesting that he had only seen a single specimen. The lectotype sheet
contains two small plants. Despite this it seems that this is the material examined by
Gray. The sheet is annotated with 'pygmaea n. sp.' in his hand and no other possible
type material has been located at GH (M. Canosa, in Hit., 1987) or any other
herbarium.

3. Podotheca gnaphalioides is a polymorphic species in regard to its habit, which
varies from prostrate to erect, and in the vestiture of the leaves and bracts. The most
common form of the species has ascending to erect major axes and the outer involucral
bracts have a vestiture of septate hairs and occasional, seemingly sessile or shortly
stalked, glandular hairs (e.g. Aplin 3362, Short 1602, Short 1722). A few collections
from drier and inland localities are of specimens with prostrate major axes and with
bracts which are glabrous or with few septate hairs (e.g. Blackall 453 from Sandstone).
Other collections (e.g. Willis s.n. MEL 1555706, Selk 1705) are characterized by
having bracts with a dense vestiture of stalked, glandular hairs as in P. chrysantha and
P. wilsonii. Such specimens occur in the Jurien Bay-Yanchep region.

Formal recognition of the entities does not seem warranted but additional
collections may prove otherwise.

Selected Specimens Examined (Total c. 120):

Western Australia — Dirk Hartog Island, 2.ix. 1972, George 1 1383 (PERTH); 15-5 km W. of Mullewa,
l.ix.l982, Short 1602 (MEL, PERTH); Anderson Rocks, 13.ix.l982, Short 1722 MEL, PERTH); Caroling
Rocks, 6.X.1983, Short 1971 (MEL); E. edge of Lake Moore, 15.ix.l986, Short 2930 (AD, MEL,
PERTH).

4. Podotheca pritzelii P. S. Short, sp. nov.

Herba annua. Axes majores ascendentes vel erecti, c. 5-25 cm longi, pilis planis septatis. Folia lineari
vel lanceolata 1-3-5 cm longa, 0- 1-0-25 cm lata, succulenta, pallens viridia vel purpurascentia, pilis
planis septatis. Capitula ovoidea usque lanceoloidea vel cylindrica, T9-2-6 cm longa, 0-26-0-8 cm
diametro. Bracteaea involucralis 15-32, ovatae usque lanceolatae vel anguste triangulares usque
lineares triangulares vel obovatae usque oblanceolatae, 7-5-22 mm longae, 1 -5-3-5 mm latae; bractae
exteriores herbaceae, succulentae, pallentes virides vel purpurae, marginibus angustis hyalinis pilis
longis ciliatis, pili septati absens; bracteae interiores hyalinae praeter costa opaca, glabrae. Flosculi
19-73; corolla tubes 14-19 mm longa. Stamina 5; antherae 1-4-1-83 mm longae, unaquaeque
pollinibus c. 400. Cypselae 1-5-1 -6 mm longae, 0-45-0-6 mm diametro. Pappi setae laeves usque
barbellatae in parte inferna 1/3-1/2, superae 1/2-2/3 plumosae.

Holotypus: Lake Ninan, at junction of Brennan Road with the Wongan
Hills-Yereeoin road. 30° 56' S., 1 16° 39' E. Growing in sand amongst samphire and
Melaleuca. 25.x. 1 983, Short 2214B (MEL 1 524328). Isotypi: AD, BRI, CANB, CBG,
HO, K, NSW, PERTH.

Annual herbs. Major axes ascending to erect, 5-25 cm long, with flat, septate
hairs. Leaves linear or lanceolate, 1-3-5 cm long, 0-l-0-25cm wide, succulent,
pale-green or purplish, with flat, septate hairs. Capitula ovoid to lanceoloid or
cylindrical, l-9-2-6cm long, 0-26-0-8 cm diam. Involucral bracts 15-32 per
capitulum, ovate to lanceolate or narrowly to linear triangular or obovate to
oblanceolate, 7-5-22 mm long, 1 -5-3 -5 mm wide; outer braets herbaceous, succulent,
pale green or purple, with narrow hyaline margins with long-ciliate hairs, flat septate
hairs absent; inner bracts hyaline except for opaque midrib, glabrous. Florets 19-73
per capitulum, yellow-orange; corolla tube 14-19 mm long. Stamens 5; anthers
l-4-l-83mm long; microsporangia 1-05-1-46 mm long; apical appendages 0-35-

50

S^llONAl HFRB-\RH '.t ('h 'U umi A i\lFl i

-t •;••

N \ MON \f IH HU\HM M (>!■
\ l< lORI \ (M! n. \t vl K \M \

Fig. 5. Holotype sheet of P. pritzelii (Short 2214B).

0-43 mm long. Pollen grains c. 400 per anther. Cypselas l-5-l-6mm long, 0-45-
0-6 mm diam. Pappus 5 bristles, each smooth tobarbellate in the lower 1/3- 1/2, the
upper 1/2-2/3 plumose. (Fig. 5).

Distribution (Fig. 1):

Western Australia. Only known with certainty from the vicinity of Wongan Hills.
A collection, Gardner 7467, said to come from Lake Annean, Nannine, seems to be

51

erroneously labelled. The locality seems more likely to be Lake Ninan, the type
locality. A further collection, Kenneally 5 799 from the Mortlock Flora Reserve and
containing somewhat immature specimens, may be of this species.

Lake Ninan is in the Monger Lake System (Bettenay & Mulcahy 1972).

Ecology:

The only ecological data available comes from the type collection (see above)
which indicates that the species grows in saline sand amongst samphire and
Melaleuca.

Notes:

1 . The specific epithet commemorates Ernst Georg Pritzel, a German botanist
who, in 1900-1901, collected with Ludwig Diels in Western Australia. The first
collection of this species was gathered by Pritzel.

2. In the field P. pritzelii is readily distinguished from P. gnaphalioides. The
leaves and bracts are manifestly succulent and are usually a distinct pale green,
although sometimes the bracts may be purple. Larger individuals frequently branch at
the upper nodes, an uncommon feature in P. gnaphalioides. There is also a tendency
for the largest capitula in P. pritzelii to be smaller than those of robust specimens of P.
gnaphalioides, a situation reflected by the bract and floret number per capitulum. The
bracts of this species also lack flat, septate hairs on the outer surface, an uncommon
condition in P. gnaphalioides.

Differentiation of P. pritzelii from P. gnaphalioides can be difficult from
herbarium specimens, a situation not helped when habitat notes are lacking. Although
the pale green colour of the leaves and bracts may more or less remain the original
succulent nature is often not apparent in dried specirnens.

Apart from morphological differences P. pritzelii is found in a different habitat
from that frequented by P. gnaphalioides. The latter species was not observed at Lake
Ninan when the type collection of P. pritzelii was gathered. As noted above P.
gnaphalioides barely encroaches into the samphire dominated zone of saline lakes
(also see under P. uniseta).

Specimens Examined:

Western Australia — ?Lake Annean, Nannine, x. 1945, Gardner 7467 (PERTH); Wongan Hills,
13.X.1903, Morrison 13058 (K, PERTH — 2 sheets); District Avon, in apertis arenosis, x.1901, Pritzel 775
(BM, E, K, PERTH).

5. Podotheca uniseta P. S. Short, sp. nov.

Herba annua. Axes majores ascendentes vel erecti, c. 5-25 cm longi, pilis planis septatis. Folia linearia
vel lanceolata, 1-4-5 cm longa, 0-1-0-35 cm lata, succulenta. Capitula anguste ovoidea vel cylindrica,
2-2-8 cm longa, 0-26-0-9 cm diametro. Bracteae involucralis 23-35, ovatae usque lanceolatae vel
anguste triangulares usque lineares triangulares vel lineares vel obovatae usque oblanceolatae,
4-5-22 mm longae, 0-6-3-2 mm latae; bracteae exteriores herbaceae interdum semisucculentae,
marginibus angustis hyalinis pilis longis ciliatis, pili septati absens vel on bracteae eximae verticillus;
bracteae interiores hyalinae praeter costa opaca, glabrae. Flosculi 26-67; corolla tubus 14-8-20-2 mm
longa. Stamina 5; antherae 1-35-1-94 mm longae, unaquaeque pollinibus c. 300-600. Cypselae
1 - 8- 1 -9 mm longae, 0- 5 5-0-65 mm diametro. Pappus annulus seta uno, seta laevi vel barbellata in parte
inferna 1/2-2/3, in supera 1/3-1/2 plumosa.

Holotypus: E. edge of Lake Moore (c. 58 km from Paynes Find along Cleary road),
29° 40' S., 117° 43' E. Growing amongst Gunniopsis & Halosarcia. Sand. 15.ix.l986,
Short 2929 (MEL 689074). Isotypi: AD, BRI, CANB, HO, K, NSW, PERTH.

Annual herbs. Major axes ascending to erect, c. 5-25 cm long, with flat, septate
hairs. Leaves ± linear or lanceolate, 1-4-5 cm long, 0-1-0-35 cm wide, succulent,
green, red or purple. Capitula ± narrowly ovoid or ± cylindrical, 2-2-8 cm long,
0-26-0-9 cm diam. Jnvolucral bracts 23-35 per capitulum, ovate to lanceolate or
narrowly to linear triangular or ± linear or ±obovate to oblanceolate, 4-5-22 mm
long, 0-6-3-2 mm wide; outer bracts herbaceous, sometimes semisucculent, green or
tinged purple, with narrow, hyaline margins with long-ciliate hairs, septate hairs

52

absent or only present on bracts of the outermost whorl; inner bracts hyaline except for
opaque midrib, glabrous. Florets 25-61 per capitulum, mainly yellow but upper part of
corolla tube usually purple; corolla tube 14-8-20-2mm long. Stamens 5; anthers
l-35-l-94mm long; microsporangia 0-97-l-59mm long; apical appendages
0-28-0-49 mm long. Pollen grains c. 300-600 per anther. Cypselas 1-8-1 -9 mm long,
0-55-0-65 mm diam. Pappus a basal annulus with a single bristle, the bristle smooth or
barbellate in the lower 1/2-2/3, the upper 1/3-1/2 plumose. (Figs, lb, 6).

Distribution (Fig. 1);

Western Australia. Only known from the margins of Lake Monger, Lake Moore
and a saline flat south of Mora wa. All locations fall within the Avon drainage system as
defined by Beard (1973).

Ecology:

Commonly found in the samphire zone surrounding salt lakes. Collectors’ notes
include ‘Sandy rise in samphire flat with other chenopodiaceous shrubs and scattered
Eremophila. Sandy to very sandy pale red loam forming weak erust in places; coarse
sand frequently on surface’ and ‘Growing amongst samphire c. 20 m above salt pan,
just extending into Melaleuca shrub zone. In white sand.’.

Notes:

1 . The single pappus bristle is the most distinctive feature separating this species
from P. gnaphalioides and P. pritzelii. It is virtually indistinguishable from the latter
on other features although the herbaceous bracts of P. pritzelii are more succulent and
usually a bright green. The distinction of herbarium specimens of P. gnaphalioides
from P. uniseta on morphological features other than the pappus seems untenable.
However, distinct ecological differences have been observed where the two species
occur in the same locality. At the type locality, where P. uniseta was observed to grow
amongst samphire and Gunniopsis, P. gnaphalioides {Short 2930) was reeorded as
primarily growing under Melaleuca. Only oecasionally did it extend to the outer limits
of the samphire zone where individuals of both species grew. In the field it is evident
that plants of P. uniseta tend to be smaller, have more succulent bracts and are
coloured a deeper purple than specimens of P. gnaphalioides. Putative hybrid
individuals have never been observed at such sites.

Specimens Examined:

Western Australia — 93-5 km N. of Cleary, 13.xi.l983, Haegi 2642 & Short (AD, MEL, PERTH);
Mongers Lake, 3.ix. 1982, Short 1634B (AD, CANB, HO, NSW, PERTH); Mongers Lake, 23.x. 1983, Short
2I79B (MEL); 5 km S. of Morowa, 16,ix.l986, Short 2960 (MEL, PERTH).

6. Podotheca angustifolia (Labill.) Less., Syn. gen. Compos. 272 (1 832); DC., Prod. 6:
159 (1838); Steetz in Lehm. PI. Preiss. 1: 448 (1845); Benth., FI. Austr. 3: 601 (1867);
Grieve & Blackall, W. Aust. Wildfls 824 (1975); Cooke in Jessop & Toelken, FI.
S. Aust. 3: 1573 (1986); Lander in Marchant et ai, FI. Perth Region 700 (1987). —
Podosperma angustifoliaGdbi\\.,'Ho\ . Holl. PI. Sp. 2: 35, t. 177 (1806); J. M. Black, FI.
S. Aust. 1 St ed. 636 ( 1 929), 2nd ed. 9 1 4 ( 1 957); W. M. Curtis, Stud. FI. Tas. 34 1 ( 1 963);
J . H. Willis, Handb. PI. Viet. 2:719(1973). — Phaenopoda angustifolia (Labill.) Cass.,
Diet. Sci. Nat. 42: 84 (1826). Type: ‘Habitat in terra Van-Leuwin.’ Syntypes: FI
(n.v.), P. (see note 1 below).

Lophoclinium citrinum Endl., Bot. Zeitung (Berlin) 1: 457 (1843). Type: ‘Nova
Hollandia austro-occidentalis’. Possible Syntypes & Isosyntypes: Preiss 106, LD,
MEL 1543637 (ex herb. Steetz), MEL 1543638 (ex herb. Sonder), MEL 691441, P (3
sheets), W (2 sheets). (See note 2 below.)

Annual herbs. Major axes ascending to erect, 2-30 cm long, sometimes cottony,
always with some flat, septate, non-glandular hairs and short, usually conspicuously
stalked, glandular hairs. Leaves ± linear, lanceolate or oblanceolate, 0-6-3(9) cm long,
0-1-0- 3(0- 7) cm wide, with flat, septate, non-glandular hairs and stalked glandular

53

N\nOS\l mKBvKlIM OI-
3 ri loKU (\n I ). u j\

Fig. 6. Hoiotype sheet of P. uniseta {Short 2929).

nairs, sometimes ± succulent. Capitula ovoid to lanceoloid or ± cylindrical,
1 •2-4-5 cm long, 0-3-1 -2 cm diam. Involucral bracts 14-28 per capitulum, ovate to
lanceolate or narrow^ly triangular to linear triangular or ± linear, 7-28 mm long,
1-3 mm wide; outer bracts herbaceous, sometimes ± succulent, green to purple,
usually with narrow hyaline margins with long-ciliate hairs, the margins not extending
to the usually recurved apex, the outer surface with flat, septate hairs and some
glandular hairs; inner bracts hyaline except for an opaque midrib, glabrous or with
long-ciliate hairs on the margin. Florets 9-73 per capitulum, usually yellow but the
upper part of the corolla tube often purple; corolla tube 14-21 mm long. Stamens 5;
anthers 0-65-1-15 mm long; microsporangia 0-33-0-82 mm long; apical appendages
0-23-0-46 mm long. Pollen grains 20-60 per anther. Cypselas 2-2-4 mm long.

54

0-4-0-7 mm diam. Pappus of 5 bristles, each bristle smooth at the base, grading to
plumose in the upper \ 12-21 3>.

Chromosome numbers: n= 13,26 (Turner 1970).

Distribution (Fig. 1):

Western Australia (south of a line extending from Pt Quobba through Menzies to
the south coast and including islands such as Rottnest Island and North Island and
West Wallabi Island in Houtman Abrolhos), South Australia (S. of c. 30° S.),
south-west New South Wales, Victoria and islands of Bass Strait.

Ecology:

Occurs in an array of coastal and inland habitats. Collectors’ notes include ‘Open
scrub . . . dominated by Leucopogon parvijlorum, in swale near inland margin of
dunes’, ‘in open Melaleuca cuticularis swamp ... in low lying winter wet grey sandy
clay with clay subsurface’, ‘Calcareous soils. Agonis flexuosa open woodland’, ‘on
white sand ridge between trees of Casuarina', ‘in sandy loam amongst Melaleuca trees
inimediately above saline depression and extending into open salmon gum woodland
with an understorey of Atriplex, Eremophila, Olearia and Acacia shrubs. Sometimes
powing with P. gnaphalioides' , ‘Open scrub . . . Banksia - Acacia, deep sandy silt’,
‘Moderately exposed base of small granite mountain. Variable-drained, shallow,
arkosic loamy sand’ and ‘sand ridge with low mallee eucalypts and Callitris.
Associated herbs include Gnaphalium sp. & Waitzia acuminata’ .

Notes:

1 . Type material of P. angustifolia is presumably held at FI but is not available for
loan. Both the description and illustration leave little doubt as to the application of the
name. A single specimen in P is probably syntype material. It was at one time in the
herbarium of E. Drake and is labelled in an unknown hand as having been collected by
Labillardiere. It consists of a single capitulum and dissection of one of several poorly
preserved florets revealed anthers barely 1 • 1 mm long. The original illustration shows
a plant in which the capitula are somewhat open and in this respect the P specimen is
also a good match.

A further specimen in P attributed to the D’Entrecasteaux expedition and said to
be of P. angustifolia is considered to be erroneously labelled. It is not a match for
Labillardiere’s description or illustration and in fact is ofP. gnaphalioides. The notion
that the label is incorrect is supported by the fact that the D’Entrecasteaux expedition,
of which Labillardiere was a member, explored the southern coastline of Western
Australia. P. gnaphalioides, unlike P. angustifolia, barely encroaches upon the
southern coast.

2. Endlicher (1843) described the genus Lophoclinium and included in it three
species, i.e. L. manglesii Endl., L. citrinum Endl. in [sect.] Lophoclinium (as
’Eulophoclinium’} and L. album Endl. in [sect.] Brachycallyma Endl. (Endlicher did
not indicate the rank of the infrageneric names.) It follows that Lophoclinium should
be lectotypified by L. manglesii or L. citrinum, but due to uncertainty as to the
typification of both names, neither has been chosen.

Endlicher, apart from indicating that the species were from ‘Nova Hollandia
austro-occMentalis’, gave no information pertaining to the identity of the specimens
and their collector(-i). Bentham (1867) reduced L. manglesii to synonymy under P.
angustifolia and L. citrinum was placed under P. gnaphalioides. He also suggested that
L. album was conspccific with P. fuscescenshui was clearly uncertain as to its identity.
Specimens from W have been examined and it seems that both L. manglesii and L.
citrinum were described from specimens gathered in Western Australia by Ludwig
Preiss. Thus single specimens of both Preiss 106 (of P. angustifolia and collected from
Rottnest Island) and Preiss 107 (of P. gnaphalioides and from Lake Mathilda) are
labelled, in what I suspect is Enlicher’s hand, as L. citrinum and L. manglesii
respectively. The labels were compared with a sample of Endlicher’s handwriting
contained in a file compiled by Mr A. Court and housed at MEL. However, I have
some doubts as to the handwriting on the Preiss collections. It is not too dissimilar to

55

that of Lehmann. Furthermore, if the Preiss collections are syntypes then the
application of the names L. citrinum and L. manglesii differ from that applied not just
be Bentham but, as evident from determinations on sheets in W, by other botanists.
Because of such doubts I have refrained from selecting lectotypes of both names,
merely suggesting that the Preiss collections are syntypes or isosyntypes.

The application of the name Lophoclinium album remains unresolved as no
specimens bearing this name arrived with the loan of Pddotheca from W. However, it
is clear that this name does not apply to members of Podotheca s. str. as the plant was
described as having a single row of involucral bracts and white florets, each with a
pappus of about 1 5 bristles.

Selected Specimens Examined (Total c. 320)

Western Australia — 4-7 km E. of Yellowdine, 1 8.ix. 1 982, Short 1 752 (MEL); 1 6-5 km SW. of Nannup,
1 .xi. 1 983, Short 2268 & Haegi (AD, MEL, PERTH).

South Australia — Danggali Conservation Park, 12.xi.l980, Conn 910 (AD); W. of Lake Newland,
, Donner 2448 (SD).

New South Wales — 5-5 km N. of Wentworth-Renmark Rd on road to Belmore, 10.ix.l980,
Christensen 54 (AD).

Victoria — King Island, xi.l887, French s.n. (MEL 1543649).

ACKNOWLEDGEMENTS

I thank Mr J. Nailon for the SEM photographs and Miss K. E. Wison for general
technical assistance, including the preparation of the fruit sections and illustrations.
Mr P. G. Wilson kindly provided comments on the affinities of P. fuscescens and P.
pollackii and Dr W. R. Barker assisted with the Latin descriptions.

This paper is partly the result of a broad survey of fruit anatomy of species of
Australian Gnaphaliinae, a survey funded by Australian Biological Resource Study
grants (1982-1987).

REFERENCES

Beard, J. S. (1973). ‘The elucidation of palaeodrainage patterns in Western Australia through vegetation
mapping’. Vegetation Survey of Western Australia, Occasional Paper 1. (Vegmap Publications:
Perth.)

Bentham, G. (1867). ‘Flora Australiensis’, vol. 3. (Reeve: London). Podotheca, pp. 600-602.

Bettenay, E. & Mulcahy, M, J. (1972). Soil and landscape studies in Western Australia. (2) Valley form and
surface features of the South-West Drainage Division. J. Geol. Soc. Aust. 18: 349-357.

Black, J. M. (1957). ‘Flora of South Australia’, pt 4, ed. 2. (Govt. Printer: Adelaide.) Podosperma,
p. 1127.

Cassini, H. (1822). Podotheca. Dictionnaire des sciences naturelles 23: 561.

Cassini, H. (1826). Phaenopoda. Dictionnaire des sciences naturelles 42: 84.

Committee for Spermatophyta (1967). Conservation of generic names, VII. Taxon 16: 226-229.

Curtis, W. M. (1963), ‘The Student’s Flora of Tasmania’, pt 4, ed. 2. (Govt. Printer: Tasmania.) Podosperma,
p. 341.

Diels, L. (1904-1905). Fragmenta phytographiae Australiae occidentalis. Bot. Jab. 35: 55-662. [Podotheca
pollackii p. 617, (1804)].

Eichler, H. ( 1 964). Proposal for the conservation of 9009 Podotheca Cass. (1822) versus Podosperma Labill.
(1806). Regnum Veg. 34: 62-64.

Endlicher, S. (1843). Plantae Hortenses. Bot. Zeitung (Berlin) 1: 457-459.

Ewart, J. (1931). ‘Flora of Victoria’. (University Press: Melbourne.) Podosperma, p. 1 127.

General Committee on Botanical Nomenclature (1968). Conservation of generic names, IX. Taxon 17:
325-329.

Graham, R. (1842). Podotheca gnaphalioides. Bot. Mag. t. 3920.

Gray, A. (1851). Characters of some south-west Australian Compositae, principally of the subtribe
Gnaphalieae. Hook. J. Bot. Kew Card. Misc. 4: 227.

Grieve, B. J. & Blackall, W. E. (1975). ‘How to know Western Australian Wildflowers’. pt 4. (University of
Western Australia). Podotheca, pp. 824-825.

Hoffman, O. (1894). Podotheca. In Engler, A. & Prantl, K. ‘Die naturlichen Pflanzenfamilien’. (Wilhelm
Engelmann: Leipzig.) 4(5): 190-191.

Kunth, C. S. (1838). ‘Flora berolinensis’, ed. 2, vol. 1. (Berlin). 'Podosperma’, p. 379.

Labillardiere, J. H. H. de (1806). ‘Nova Hollandiae plantarum specimen’, vol. 2. (Paris). Podosperma,
pp. 35-36, t. 177.

Lessing, C. H. (1832). ‘Synopsis generum Compositarum’. (Berlin). 'Podosperma', Podotheca, pp. 133,
272.

56

Maiden, J. H. & Betche, E. ( 1 9 1 6). ‘A census of New South Wales Plants’. (Govt. Printer: Sydney.) Podotheca,

p. 201.

Merxmiiller, H., Leins, P. & Roessler, H. (1977). Inuleae — systematic review. In Heywood, V. H., Harbome,
J. B. & Turner, B. L. (eds), ‘The biology and chemistry of the Compositae’. (Academic Press: London.)
1: 577-602.

Mueller, F. J. H. ( 1 882). ‘Fragmenta phytographiae Australiae’, vol. 1 2. (Melbourne.) Podosperma pollackii,

pp. 21-22.

Mueller, F. J. H. ( 1 889). ‘Second systematic census of Australian plants. Part 1 . Vasclares’. (McCarron, Bird
& Co.: Melbourne). Podosperma, pp. 134-135.

Mulcahy, M. J. & Bettenay, E. ( 1 972). Soil and landscape studies in Western Australia. ( 1 ) The major drainage
divisions. J. Geol. Soc. Aust. 18: 349-357.

O’Brien, T. P. & McCully, M. E. (1981). ‘The study of plant structure. Principles and selected methods’.

(Termarcarphi Pty Ltd: Wantima, Victoria.)

Oliver, F. W. (1913). ‘Makers of British Botany’. (Cambridge University Press.)

Rodway, L. (1903). ‘The Tasmanian Flora’. (Govt. Printer: Hobart.) Podotheca, p. 96.

Schultz, C. H. (1834). Drei neue Pflanzengattungen. Flora 17: 465-479.

Short, P. S. (1981). Pollen-ovule ratios, breeding systems and distribution patterns of some Australian
Gnaphaliinae (Compositae: Inuleae). Muelleria 4: 395-417.

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6: 9-22.

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382-389.

Manuscript received 30 March 1988

NOTES ON THE FRUIT ANATOMY OF AUSTRALIAN MEMBERS OF THE

INULEAE (COMPOSITAE)

by

P. S. Short*, K. E. Wilson* and J. Nailon^

ABSTRACT

Short, P. S., Wilson, K. E. & Nailon, J. Notes on the fruit anatomy of Australian
members of the Inuleae (Asteraceae). 7(1): 57-79(1989). — Observations of

the anatomy and morphology of Australian inuloid genera are presented. Genera
examined include Angianthus Wendl. s. lat., Helichrysum Miller, Helipterum DC.,
Ixiolaena Benth., Millotia Cass., Myriocephalus Benth., Podolepis Labill.,
Scyphocoronis A. Gray and Toxanthes Turcz. Comments on generic delimitation are
made.

INTRODUCTION

As an adjunct to revisionary studies (by Short) of Australian genera a study of the
fruit anatomy and morphology of species of Inuleae was commenced in 1982. Initial
studies involved an assessment of the value of fruit anatomy in ascertaining specific
and generic limits, an assessment carried out through the study of the segregate genera
of Angianthus Wendl. 5. lat. previously recognized by Short (1983). Following
encouraging results a more general survey was carried out.

It is clear from studies that fruit characters are of much value in ascertaining
generic limits within the Inuleae, and descriptions of fruit are incorporated in
revisions of Blennospora A. Gray (Short 1987), Podotheca (Short 1989) and
Pogonolepis Steetz (Short 1986a). Forthcoming revisions of Calocephalus R. Br.,
Gnephosis Cass, and Chthonocephalus Steetz (all sensu Benth. 1867) will also include
descriptions of the fruit anatomy of many of the segregate genera that will be
recognized. In this paper results of the studies of Angianthus 5. lat. and miscellaneous
observations of other inuloid genera are presented.

MATERIALS AND METHODS

The extent of the observations of the fruit anatomy and morphology of species in
this paper vary considerably. In some cases anatomical comments are the result of the
examination of median-transverse sections (TS) and median-longitudinal sections
(LS) of fruit, in others only TS have been studied. Similarly not all species have been
examined with a scanning electron microscope (SEM), surface features sometimes
being only examined by light microscope using a 40X objective. The bracketed
abbreviations LS, TS & SEM accompanying the generic headings below are used to
indicate the extent of observations.

Voucher specimens are housed in either AD or MEL, the majority being in the
latter herbarium. The appendix lists the specimens examined, giving species name,
author citation and collector’s name and number.

Immature fruit were fixed in the field in 4:3: 1 (chloroform:absolute
ethanol: glacial acetic acid) and subsequently stored in 70% ethanol. Mature, dry fruit
were rehydrated and then fixed in 5% glutaraldehyde in 0-03 M Pipes buffer.
Following fixation and dehydration both immature and mature fruit were infiltrated
with G.M.A., Spurr’s or L. R. White (London Resin Co.). Infiltration of L. R. White
over several weeks, with repeated two hour vacuum infiltrations, proved to be the
most effective. Two micron thick sections were stained with toluidine blue (pH 4-5)
and mounted in histomount.

’National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.
^Department of Botany, Monash University, Clayton, Victoria, Australia 3168.

57

58

Fig. 1 . The position of the vascular bundles in the pericarp in relation to the cotyledons.

To assist in the identification of vascular bundles in the pericarp fruit were
cleared and stained in a solution of 1% basic fuchsin in 10% KOH (Wilcox 1977).

Terminology is generally self-explanatory and follows Bruhl (1984); the terms
lateral, medial and oblique have been used to indicate the position of the vascular
bundles in the pericarp and testa in relation to the cotyledons. In fruit with lateral
vascular bundles the bundles occur in the plane of the upper surface of the cotyledons.
In fruit with medial vascular bundles the bundles are opposite the lower surface of the
cotyledons. If the position is intermediate between lateral and medial then bundles are
considered to be oblique (Fig. 1 ). Placing the position of the bundles into one of the
three categories did not usually present a problem. However, in some species any two
bundles in a fruit were not opposite one another as illustrated in Fig. 1 , but occupied
different positions, e.g. one in a lateral position, the other in an oblique position. In a
few species bundle position could not be determined as individual seedling leaves
could not be discerned.

RESULTS AND DISCUSSION

General

Fruit anatomy, and to some extent morphology, has been generally ignored in
taxonomic work in the Australian Asteraceae. A notable exception is the recent work
on the Cotuleae (Bruhl 1 984; Bruhl & Quinn 1 988). Such studies are useful in deciding
generic limits, a fact underlined by studies of most genera examined in this paper. This
conclusion, plus the comments made below on the affinities of species and generic
limits have been only reached following consideration of numerous morphological
features of the taxa under discussion, not just fruit characteristics.

Fruit characteristics which are often indicative of species groups are;

a) presence or absence and structure of thick-walled tissue in the pericarp and/or
testa, e.g. compare Helipterum albicans with other species of Helipterum.

b) presence or absence of a carpopodium, e.g. compare Cephalosorus and
Dithyrostegia with other members of Angianthus s. lat. (also see Haque &
Godward 1984).

c) structure of the epidermis including trichomes, e.g. compare Cephalosorus and
Pogonolepis, with fruit virtually enveloped in myxogenic cells, with other
members of Angianthus s. lat.

d) a crystalline pericarp, e.g. see Craspedia. Ciystals are commonly found in the
testa of most, if not all, inuloid species examined but not in the pericarp.

The presence or absence of vascular bundles in the testa may also be indicative of

species groups, e.g. compare Podolepis georgei with other species of Podolepis s. lat.
However, the reliability of our data is open to question. We stress that our
observations of vascular bundles in the testa were only made from medial transverse
sections. Ideally, cleared seed should be examined. Paul Wilson (pers. comm. 1988),
following examination of cleared specimens, has recorded different results to us for

59

Table 1. Fruit characteristics in species of Helipterum.

( + = present; - = absent or not applicable; L = lateral; M = medial; Ob = oblique; ? = number of vascular
bundles not clear or position not known because cotyledons absent from section; ( )♦ = number of vascular
bundles observed in cleared fruit (see text, p. 59).

Layer of
collenchyma
or

sclerenchyma
in pericarp

Number of
vascular
bundles in
pericarp in
TS

Position of
vascular
bundles in
pericarp

Number of
vascular
bundles in
testa in TS

Position of
vascular
bundles in
testa

Lateral
thickening of
epidermal
cell walls

H. albicans

+

2

L

0

—

-

H. charsleyae

+

2

Ob

Ob

—

H. chlorocephalum

—

2

M

1

M

—

H. Jloribundum

—

2

9

1

9

—

H. maryonii

—

2

Ob/M

0(1)*

—

—

H. microglossum

—

2

9

1(?2)

9

9

H. moschatum

—

2

M/Ob

0(1,2)*

—

+

H. pygmaeum

2

M

1(?2)

M

+

H. strictum

—

2

Ob

2

Ob

—

H. stuartianum

—

2

M

0

—

—

H. uniflorum

—

2

Ob

0(1)*

—

+

some species of Helipterum (see Table 1). He has also recorded the presence of a
vascular bundle in the testa of Triptilodiscus. We observed none in TS. The
discrepancy in the data partly reflects the occasional difficulty in recognizing bundles
in TS. However, it also reflects variation in the location of vascular bundles and the
extent of vascularization in the testa. Observations by Paul Wilson indicate that a
single vascular strand enters the testa through the funicle and penetrates a varying
distance into the testa. Very often strands do not reach the apex of the ovule and the
extent to which strands penetrate the testa may also vary within the one species. On
entry to the ovule the strand may remain undivided or divide into two.

The position of the vascular bundles in the pericarp or testa is rarely constant in
any species group, e.g. in Angianthus s. str. On the other hand, in Podotheca s. str. only
oblique bundles occur in the pericarp of the five species examined (Short 1989).

The possible occurrence of a reticulate venation pattern in the pericarp has been
alluded to for Craspedia aff. pleiocephala. Of all species included in the general survey,
including species in genera under revision and not included here, only Gnephosis
gynotricha Diels definitely has reticulate venation. This pattern is absent from its close
relative G. macrocephala Turcz.

In some species it was difficult, from transverse sections, to differentiate with
certainty the testa from the pericarp. Sometimes this problem was overcome through
the examination of sections of immature fruit. With many sections a clear gap exists
between the layers of tissue considered to be testa and pericarp but in some species the
tissue layers are all but obliterated in mature fruit. Thus it is possible that mistakes
have been made in our interpretation of these tissues. Paul Wilson {in litt. 1987) has
noted that, in Blennospora, a layer of sclerenchyma previously deemed to be in the
pericarp (Short 1987) is more likely to be the outer layer of the testa. However,
although the correct elucidation of testa and pericarp is critical for broad comparisons
of composite genera, it is not so important for discerning species groups within
unnatural genera. Differences in characters such as the presence or absence of
sclerenchyma in the combined fruit/seed wall and the total number of vascular
bundles are still useful features for indicating species affinities.

Actinobole Fenzl ex Endl. (LS, TS, SEM)

Fruit sections were obtained for each of the four species recognized in the genus
(Short 1 985). They are seemingly identical. All possess a proininent stylopodium, lack
a distinct carpopodium, are covered with twin-myxogenic hairs, and have two oblique
vascular bundles in the pericarp. Vascular bundles are lacking from the testa (Figs
2a-b, 5d).

60

Fig. 2. Surface features of fruit. a,b — Actinobole condensatum, general view and apex showing the prominent
stylopodium (Short 1005). c — Angianthus acrohyalinus, upper half of fruit with the position of the
vascular bundle visible (Short WOO). d,e — Angianthus aff. micropodioides, general view and
myxogenic hairs (Short 2664). f — Angianthus atf. milnei, carpopodium (Short W12). g —
Cephalosorus carpesioides, general view (Short 2043). h — Chondropyxis halophita, trichomes (Short
2565).

61

Angianthus Wendl. s. lat.

In a recent revision (Short \ oi Angianthus s. lat. eight genera were reinstated.

Genera were recognized on the basis of a number of characters, including leaf
morphology and the number, arrangement and morphology of the bracts of both the
general involucre and capitula. Fruit characteristics were also used but additional
notes on fruit structure are reported here.

Angianthus Wendl. s. str. (TS, SEM)

Species of Angianthus have a thin pericarp containing no thick-walled tissue. In
all species it is difficult to differentiate the testa from the pericarp. In some sections a
more or less well-defined, one-cell wide layer could be discerned. The cells have
slightly thickened walls and it is probably the outer layer of the testa. Vascular bundles
were not observed in the testa but two occur in the pericarp. The vascular bundles are
oblique in A. cunninghamii andT. acrohyalinus hni in A. drummondii they are almost
medial. In both A. brachypappus and A. milnei one vascular bundle is in the lateral
position, the other is oblique. Due to the thinness of the pericarp the position of the
vascular bundles is often visible on the fruit surface as a small ridge (Fig. 2c). A
carpopodium is well developed in all species (Figs 2d, f), and apparently twin-celled,
myxogenic papillae occur on the fruit surface of most species (Figs 2c-f).

Cephalosorus A. Gray (LS, TS, SEM)

Cephalosorus is a monotypic genus. Fruit of C. carpesioides have an outer layer of
myxogenic cells (Figs 2g, 5c). Two, more or less lateral vascular bundles occur in the
pericarp. Vascular bundles were not observed in the testa, and thick-walled tissue is
absent from both the testa and pericarp. A carpopodium is absent or poorly developed
with the dehiscence region being in a pronounced basal hollow due to the large,
myxogenic cells enveloping the fruit.

Dithyrostegia A. Gray (SEM)

Dithyrostegia contains two species, D. amplexicaulis and D. gracilis P. Short.
Unlike all other species once referred \o Angianthus s. lat. both have fruit enveloped by
elongate, somewhat rigid hairs (Figs 3a, b). A carpopodium is absent.

Epitriche Turcz. (LS, TS, SEM)

Fruit of the only species, E. cuspidata, was not closely examined before the
mounted sections had considerably faded. However, longitudinal sections revealed an
unusual feature, that of a several cell-wide cap of sclerified tissue at the apex of the
fruit. The pericarp, of which the cap is part, was barely sclerified in the TS of the
fruit.

Hyalochlamys A. Gray (TS)

Hyalochlamys globifera A. Gray, the only member of the genus, has a more or less
obpyriform, glabrous fruit with a well developed carpopodium. The pericarp is thin,
lacks thick-walled tissue and contains two vascular bundles.

Pleuropappus F. Muell. (TS, SEM)

Angianthus phyllocalymmeus was referred (Short l.c.), with some apprehension,
to Pleuropappus. Not only does the species superficially resemble many species of
Angianthus s. str., but it also has four capitular bracts which are arranged in the same
manner as in Angianthus. However, Pleuropappus can be distinguished by the fruit,
which are oblique to the corolla tube, the oblique pappus, and the presence and
arrangement of about four capitulum-subtending bracts. The oblique orientation of
the fruit does not occur in any other member of Angianthus s. lat., although this same
feature is found in Rutidosis (see below).

Transverse sections of fruit show no major anatomical differences between P.
phyllocalymmeus and members of Angianthus s. str. The fruit wall is thin and the two
vascular bundles, which only occur in the pericarp, are oblique (Fig. 12c). A
carpopodium is also well developed in Pleuropappus. These observations do not
support the reinstatement of the genus.

62

Fig. 3. Surface features of fruit. a,b — Dithyrostegia amplexicaulis, general view and base, note absence of
carpopodium (Short 2398). c,d — Gnephosis pygmaea, general view and trichomes (Short 2383). e,f —
Helipterum charsleyae, general view and carpopodium (Short 2003). g,h — Millotia myosotidifolia,
general view and trichomes (Short 2I28B).

63

Pogonolepis Steetz (LS, TS, SEM)

The outermost layer of the pericarp of species of Pogonolepis is composed of
myxogenic cells (see Fig. 7, Short 1986a). There is also a pronounced carpopodium.
Layers of thick-walled tissue are absent from both the testa and pericarp. Two vascular
bundles, more or less in the lateral position, occur in the pericarp (Fig. 10b). Vascular
bundles were not observed in the testa. Pogonolepis and Cephalosorus are the only
genera in Angianthus s. lat. to have species with the myxogenic cells covering the
surface of the fruit. Despite this similarity they are not closely related, differing in
many inflorescence and leaf characters.

Siloxerus Labill.

This genus, containing three species, was not closely examined for this paper.
However, the cypselae are readily distinguished from those of other segregate genera
by their colour alone. Beside the purple colour the small fruit are sparsely to densely
papillose and lack a carpopodium.

In the revision of Angianthus s. lat. (Short l.c.) three species, each clearly without
close affinities with Angianthus s. str., were treated as species of uncertain affinity.
Each is now referred to a different genus, viz Fitzwillia P. Short, Lemooria P. Short
and Sondottia P. Short (Short 1 989). Fruit of F. axilliflora have some similarities with
those in Angianthus s. str. in that in TS the fruit wall is thin, thick-walled tissue is
lacking in either the testa or pericarp and there are two, oblique/medial vascular
bundles in the pericarp. However, a carpopodium is absent, a capping of sclerenchyma
occurs at the fruit apex and the entire surface is covered with long, intertwined hairs.
Fruit sections of S', connata (W. V. Fitzg.) P. Short have not been obtained. The fruit
surface in this species is glabrous and there is a well developed carpopodium. Fruit
sections of L. burkittii faded before they were closely examined. Nevertheless it is
evident that this species has two vascular bundles in the pericarp and that a few
papillae are scattered over the fruit surface. Thick-walled tissue seems to be absent
from the pericarp and testa.

Chondropyxis D. Cooke (SEM)

Chondropyxis halophila was included by its author (Cooke 1986) in the
Anthemideae but with the comment that, along with Ceratogyne Turcz.,
Dimorphocoma F. Muell. & Tate, Elachanthus F. Muell. and Isoetopsis Turcz. it
formed a group with closer affinities to Astereae than Anthemideae. Bremer (1987)
has suggested that Isoetopsis is a member of the Inuleae (subtribe Pluchinae). If this is
the case then Chondropyxis possibly should be placed in the Inuleae. It is included in
the current paper because of the distinctive hooked hairs which occur on the fruit (Fig.
2h), a form of trichome which does not seem to be widely distributed in the tribe.

Craspedia Forst. f (TS)

Fruit of two species currently included within Craspedia have been examined.
One, Short 853, is a New Zealand member of the C. uniflora complex. The other
species, Short 837, possibly C. pleiocephala, was collected in South Australia.

The New Zealand taxon has fruit with a thin pericarp (two cells wide) which lacks
thick-walled cells and contains two vascular bundles in the lateral/oblique position.
The testa lacks thick-walled tissue and vascular bundles were not observed. Externally
the fruit are minutely papillate, occasionally have a sparse cover of long trichomes,
and possess a prominent annular carpopodium.

Craspedia afli pleiocephala has two, oblique vascular bundles in the perica^ (Fig.
5a). Two to three scattered vessels were observed in some sections suggesting the
presence of a reticulate venation system. However, the most striking aspect of this
species was the presence of large, usually more or less oblong crystals in the pericarp.
The crystals are not scattered but make up much of the pericarp, cell detail being
obscured. Externally the fruit are silky hairy and lack a prominent, annular
carpopodium.

Craspedia, as currently delimited, is a heterogeneous group. Arid zone taxa such
as C. pleiocephala and C. chrysantha (Schdl.) Benth. certainly have a superficial

64

Fig 4. Surface features of fruit. a,b,c — Millotia tenuifolia, general view, myxogenic papillae and
carpopodiura {Short 1753). d,e,( — Scyphocoronis incurva, general view, base lacking carpopodium
and hollow apex {Short 2001). g,h — Toxanthes muetteri, general view (corolla attached) and
carpopodium {Short 1251).

65

O 0

D

■ Endosperm
E3 Testa

ID Sclerenchyma
BCollenchyma
c Vascular bundle
s Space

Distinction of cell types
in pericarp/testa

0.2mm

Fig. 5. Transverse sections of fruit, a — Craspedia aff. pleiocephala {Short 837).b — Helichrysum bracteatum
{Short 3022). c — Cephalosorus carpesioides {Short 2403). d — Actinobole condensatum {Short

1005 ).

66

resemblance to Cvaspedia s. str. [which includes the many, mainly alpine, species
commonly referred to the C. uniflora and C. glauca (Labill.) Spreng. complexes] and
all are united in Craspedia s. lat. by the presence of receptacular scales. However,
cytological evidence and that presented above does not support the current circum-
scription of the genus. Craspedia pleiocephala has a haploid chromosome number of
« = 6 (Short 1981,1 986b) whereas members of Craspedia s. str. have a base number of
x = 1 1 (Turner 1970; Beuzenberg & Hair 1984). The observed differences in the fruit
are certainly major, with the largely crystalline pericarp seen in C. aff. pleiocephala not
being observed in any of the other genera examined.

Helichrysum Miller (TS)

Helichrysum, in the broad sense, occurs in Eurasia, Africa, Madagascar, Australia
and New Zealand and contains about 500 species (Merxmuller et al. 1977). The
artificiality of the genus is widely recognized, with Bentham ( 1 867, p. 1 62) noting that
‘the limits to be assigned to the group are very uncertain, as it is connected with so
many others by insensible gradations’. Using features such as habit, shape of the
involucre and receptacle, bract morphology, pappus structure, and vestiture of the
cypsela, Bentham {l.c.) erected six Sections to accommodate Australian species of
Helichrysurn. Given the array of morphological variation exhibited there seems no
doubt that in years to come most, in fact probably all, Australian species currently
placed in Helichrysum will be referred to a number of segregate genera, e.g.
Chrysocephalum Walp. and Lawrencella Lindley. Only four species of Helichrysum
were examined in this study but observations of fruit anatomy support this
contention.

Helichrysum apiculatum, a name applied to a large number of taxa deserving
specific recognition, was referred by Bentham to Sect. Chrysocephalum (Walp.) Benth.
Fruit of the taxon examined displayed a thin pericarp with two, oblique vascular
bundles (Fig. 1 If). Thick-walled tissue was absent from the pericarp and testa, and
vascular bundles were not observed in the testa. There is a prominent, annular
carpopodium.

Helichrysum bracteatum in Sect. Xerochlaena Benth., is another polymorphic
species. The fruit examined have a pericarp which is 14-15 cells wide and primarily
consists of collenchyma, but there is a gradation to sclerenchyma, with the innermost

3- 5 cell wide layer being solely of the latter tissue. Two lateral vascular bundles occur
in the pericarp and two more occur in the testa (Fig. 5b).

Helichrysum leucopsideum, also placed by Bentham in Sect. Xerochlaena, has a

4- 8 cell wide collenchymatous pericarp (Figs 7a, 1 1 b). Sclerenchyma is absent. As with
H. bracteatum this species has two vascular bundles in the pericarp, but they have a
medial, not a lateral orientation. A single vascular bundle was observed in the
testa.

Helichrysum cassinianum was referred by Steetz (1845) to Schoenia Steetz and
this was followed by Bentham (1867). Haegi (1986) referred it to Sect. Lawrencella
(Lindley) Benth. of Helichrysum, commenting that ‘further studies are likely to
confirm the placement of all these related species in a distinct genus Lawrencella
Lindley’ (p. 1 526). The large, coarsely silky, compressed fruit readily distinguish it
from other species examined, but it has similarities with H. leucopsideum in that there
are two medial vaseular bundles and a layer of collenchyma (2-4 cells wide) in the
periearp. Unlike H. leucopsideum two vascular bundles were seen in the testa (Figs 6,
llg).

Helipterum DC. (TS, SEM)

Despite its illegitimaey the generic name Helipterum DC. is commonly applied to
about 1 00 species from Africa and Australia. As with Helichrysum it is well known to
be an unnatural genus (e.g. Short 1983, Wilson 1987), a fact underlined by our
examination of the fruit anatomy of 1 1 Australian species.

The fruit section of H. albicans subsp. albicans var. buffaloensis is markedly
different from that obtained for other species of Helipterum. It has two layers, eaeh
one-cell wide, of sclerenchyma and two, lateral vascular bundles in the pericarp (Figs

67

Fig. 6. Transverse section of fruit of Helichrysum cassinianum (Short 1575).

68

o

o

Fig. 7. Transverse sections of fruit, a — Helichrysum leucopsideum (Short 1554). b — Heliplerum albicans
subsp. albicans var. buffaloensis (Short 1403). c — Helipterum pygmaeum (Short 2973). d —
Helipterum charsleyae (Short 2003).

69

7b, 1 Ic). Vascular bundles were not observed in the testa. The H. albicans species
group, revised by Wilson ( 1 960), is undoubtedly generically distinct from other species
currently included in Helipterum.

Helipterum charsleyae is also one of several species which probably constitute a
distinct genus (Short 1989). Unlike other species examined it has a layer of
sclerenchyma, 4-6 cells wide, in the pericaip. There are two vascular bundles in both
the pericarp and testa (Fig. 7d). The fruit has a prominent carpopodium and is
enveloped in long trichomes with bifid apices (Figs 3e, f).

Pericarps of fruit of the remaining nine species examined lack at least a complete
layer of collenchyma or sclerenchyma, but in H. pygmaeum there is a scattering of
thick-walled cells (not depicted in Fig. 7c) in what appears to be the inner layer of the
pericarp. However, in this species and H. microglossum, H. strictum and H. uniflorum
the distinction between testa and pericarp was not obvious in TS of mature fruit.
Species differ from one another in characters such as the number of vascular bundles
in the testa, the vestiture of the cypsela and the presence or absence of lateral
thickening of the cell walls in the epidermis (Fig. 1 le; see table 1).

IxiOLAENA Benth. (TS)

Bentham (1837), when describing the new genus Ixiolaena, recognized a single
species, I. viscosa Benth. Subsequently he referred a further four species, i.e. 7.
brevicompta, I. leptolepis, /. supina and 7 tomentosa to the genus (Bentham 1867).
Moore (1903) described a further species, 7 websteri, and recently Haegi (1986)
described 7. chloroleuca and 7. pluriseta.

Specimens of 7 websteri and 7 pluriseta have not been seen, but studies of the
other species currently included in the genus suggest that Ixiolaena s. str . includes only
7. viscosa. The habit, the presence of white laminae on the middle bracts of the
involucre, and the lower petiolate leaves set this species apart from all others. Fruit of
7 viscosa also differ from those described below in that they are not angular, are
brown-black and have prominent, elongate papillae scattered over much of the
surface. This species is also restricted to the comparatively high rainfall areas of
south-western Australia, its supposed congeners are primarily species of eremaean
Australia. Ixiolaena websteri is apparently a rhizomatous, branched shrub and has
female as well as bisexual forets (Grieve & Blackall 1975; Moore l.c.) suggesting it is
not closely related to other species currently included in Ixiolaena. A number of
inflorescence, fruit, and vegetative characters suggest that the remaining species form
a natural group, to which a further species, Leptorhynchos panaetioides (DC.) Benth.
shows affinities.

Fruit of four species, 7. brevicompta, I. chloroleuca, I. leptolepis and 7. tomentosa
have been examined (Figs 8a-d, 1 li). In many ways the cypselae of each species are
very similar. All are brown, more or less oblong and have three to five somewhat
obscure to prominent longitudinal ribs. An annular carpopodium occurs in all species,
although in 7 chloroleuca it is almost obscured by basal, myxogenic cells. In transverse
section the pericarp displays an outer sclerenchymatous layer and has two vascular
bundles. A single vascular bundle was noted in the testa, and the outer one-cell wide
layer of the testa consists of cells with pronounced, more or less, U-shaped thickening
(Fig. Hi).

The fruit of the aforementioned species differ from each other in a number of
ways, including the prominence of the ribbing, vestiture, and the position of the
vascular bundles in the pericarp. 7. brevicompta has several attributes which set it apart
from the others. The cypselae of this species has a vestiture of glandular hairs on the
upper c. f of the body. Other species have glabrous fruit or only a few glandular hairs at
the apex. The vascular bundles of the pericarp are more or less in the lateral position,
not the medial position as occurs in 7. chloroleuca and 7. tomentosa. (Their position
could not be determined in 7. leptolepis.) I. brevicompta also lacks, or has very few,
two-celled myxogenic cells embedded in the sclerenchymatous epidermal layer of the
pericarp. Such cells are common in the other species. Although obvious in transverse
section their presence is not readily observed under low magnification.

70

Fig. 8. Transverse sections of fruit, a — Ixiolaena tomentosa (Short 2988). b — Ixiolaena brevicompta (Short
3018). c — Ixiolaena leptolepis (Short 2990). d — Ixiolaena chloroleuca (Short 3003). e —
Myriocephalus stuartii (Czorney 1160).

However, after soaking in water for 24 hours, fruit with these cells are found to be
encased in a layer of mucilage. Despite prolonged soaking, such a layer was never
evident in I. brevicompta. The latter species also differs from others by having the
mature ovule contained in the lower j of the fruit, not virtually filling the whole
fruit.

71

Longitudinal sections of I. brevicompta were not obtained but the upper } of the
cypsela consists of sclerenchyma, i.e. it is a continuation of the outer layer of the
pericarp. (If the upper part of the fruit was more narrow it would no doubt be deemed
to be a ‘beak’ as in Leptorhynchos, Millotia etc.).

A further difference between species occurs in the width of the sclerenchymatous
layer. In 7. chloroleuca the layer is 3-4 cells wide, in other species it is only a single cell
wide. The outermost layer of sclerenchymatous cells in 7 chloroleuca is composed of
much larger cells than those of the inner layers and is comparable in structure to that
observed in other species.

Leptorhynchos Less. (TS)

As currently defined Leptorhynchos is a genus of about ten species. Transverse
sections of fruit of the single species examined, 7. tetrachaetus, reveal a pericarp which
lacks a layer of sclerenchyma and has two vascular bundles which are probably in the
oblique position. A further layer, probably the outer, one-cell wide layer of the testa
consists of cells with U-shaped thickening. Vascular bundles were not observed in the
testa. Externally the fruit is papillose and has an annular carpopodium.

Haegi (1986) noted the difficulty of distinguishing some species from groups of
species in Helichrysum and Waitzia Wendl. We have noted above that L. panaetioides
has strong affinities with many species currently placed in Ixiolaena.

Millotia Cass., Scyphocoronis A. Gray & Toxanthes Turcz. (LS, TS, SEM)

Schodde ( 1 963), in a revision of Millotia, noted a close relationship between this
genus, and Scyphocoronis and Toxanthes. He reported that S. majus (Turcz.) Druce is
almost morphologically identical to T. muelleri except for the hollow cup-like apex of
the fruit. He further noted that an undescribed species had the habit, indumentum,
and involucre of Toxanthes-, the fruit of Scyphocoronis-, and the alternate leaves,
peduncles, floret form, free bracts and peduncles typical of some species of Millotia.
(This species has since been named S. incurva.) Despite the intricacy of the
relationships each genus was maintained.

Sections, both TS and LS, show a marked similarity in the fruit of species
examined. All possess a layer of sclerenchyma in the pericarp (Fig. 12h). The seed is
always enclosed in the lower portion of the fruit, and whether or not it tapers to a
distinct beak or forms a hollow apex, the upper part of the fruit is forrned of
sclerenchyma, this being a continuation of the same layer of tissue surrounding the
seed. Of the species examined all but M. myosotidifolia have only two vascular bundles
in the pericarp; the latter has four.

There is some variation in two aspects, i.e. the carpopodium and the yestiture
(Figs 3g-h, 4a-h). A carpopodium is best developed in species of Millotia and is absent
or at least less well developed in Toxanthes and Scyphocoronis. However, it is not well
developed in all Millotia species, e.g. in M. greevesii F. Muell. and M. macrocarpa
Schodde it is much less prominent than in M. myosotidifolia. More marked are
differences in the vestiture of the fruit. Distinct, apparently two-celled, myxogenic
papillae occur on the surface of species of Millotia. In M. myosotidifolia and M.
inopinata Schodde they are particularly elongated and appressed (Fig. 3h). Papillae of
this type seem to be absent from Scyphocoronis and Toxanthes but trichomes do occur
over much of the fruit surface in these genera. Their structure could not be readily
determined, however, at least some of the trichomes are glandular. Most importantly
such glandular trichomes have been observed towards the apex of fruit of M. greevesii,
M. macrocarpa and M. tenuifolia. They were not observed in M. myosotidifolia.

Unless considerable emphasis is placed on the difference in fruit vestiture then
the anatomical work adds little support to Schodde’s contention that the three genera
be maintained.

Myriocephalus Benth. (TS, SEM)

As previously noted (Short 1983; Wilson 1987) the genus Myriocephalus, as
circumscribed by Bentham (1867), is an unnatural group, a fact underlined by our
studies of fruit anatomy of some species. M. gracilis differs from all other species

72

currently included in the genus by bract characters and fruit anatomy. The fruit is
covered by a semi-transparent layer of myxogenic cells, and has two medial or oblique
vascular bundles in the pericarp. A layer of sclerenchyma, observed in the pericarp of
the sectioned species of Myriocephalus, is absent or only partially developed in the

73

Fig. 1 0. Transverse sections of fruit, a — Rutidosis helichrysoides (Short 2999). b — Pogonolepis stricta (Short
2086). c — Podotepis arachnoidea ( Willis s. n . , MEL 1 554024). d — Podolepis gracilis ( Willis s . , MEL
611247).

fruit of M. gracilis and seems to be part of the testa, not the pericarp. The affinities of
M. gracilis are with species of Helipterum s. lat.

Myriocephalus helichrysoides A. Gray, M. nudus, M. plurijlorus (J. M. Black) D.
Cooke, M. rhizocephalus (DC.) Benth. and M. rudallii (F. Muell.) Benth. plus
Gnephosis pygmaea form a distinct group, with all species having similar fruit and

74

pappus characteristics. All have more or less narrow ellipsoid fruit with a sparse
covering of straight twin hairs, and an annular carpopodium. A pappus is either absent
or consists of a single bristle. The only species sectioned, M. nudus and G. pygmaea,
display a testa with a crystalline layer, and a pericarp with a single, or mainly single,
cell-wide layer of sclerenchyma (Fig. 1 lb) and two lateral vascular bundles. Three
species, G. pygmaeus, M. nudus & M. rhizocephalus, have been viewed with the SEM.
The fruit surface of each species displays a wave-like pattern caused by thickening in
cells of the epidermis (Fig. 3d). Although showing similar fruit morphologies the
species of this group have not been thoroughly examined in regard to inflorescence
structure, and given the habit differences displayed by the species (e.g. compare M.
rhizocephalus with M. helichrysoides they may prove to contain disparate taxa.
However, it would not surprise if this group, along with M. appendiculatus Benth.,
were eventually regarded as Myriocephalus s. sir.

Wilson (1987) suggested that a group of species, i.e. M. stuartii, M. morrisonianus
Diels (conspecific with Helipterum craspedioides W. Fitzg.) and M. guerinae should be
referred to Polycalymma F. Muell. & Sond. He noted that all had a similar
inflorescence structure. Fruit of H. craspedioides have not been examined (MEL
material on loan) but sections of M. stuartii and M. guerinae fruit were obtained (Figs
8e, 9, 12a, b). Both have fruit with a prominent carpopodium, a long silky
indumentum of twin hairs, a layer of sclerenchyma in the pericarp, and two vascular
bundles in the testa. However, there are major structural differences, including general
size and shape, and the number of vascular bundles in the pericarp. There are four
vascular bundles in the periearp of fertile fruit of M guerinae but only two occur in M.
stuartii. [Paul Wilson (pers. comm., 1 988) has recorded that only two vascular bundles
occur in the unfertilized ovary of M. guerinae.] The most obvious difference pertains
to two depressions occurring in the lower of the flat surface of the fruit of M.
guerinae. In sectioned fruit, these regions are seen to consist of parenchyma and,
presumably as a result of infiltration, are not seen as marked depressions (Fig. 12a).
These regions lack the long trichomes which otherwise envelop the fruit. The major
anatomical differences between the speeies does not support the inclusion of M.
guerinae with M. stuartii in Polycalymma.

Pluchea Cass. (TS)

Fruit of only P. tetranthera have been examined. Cypselae are glabrous and have a
prominent annular carpopodium. Transverse sections revealed a thin pericarp, which
lacks a thick-walled layer of tissue, and contains two vascular bundles in the
lateral/oblique position. It was difficult to distinguish testa from pericarp, and what
was believed to be testa was highly stained with cellular detail being almost obscured.
Vascular bundles were not observed in the testa and cells with thickened walls also
seemed to be absent from this tissue.

PoDOLEPis Labill. (TS, SEM)

Asteridea Lindley and Podolepis are the only inuloid genera in Australia which
contain species with marginal ligulate florets. However, not all Podolepis species
possess such florets, and despite statements to the contrary by Turner (1967) Podolepis
is clearly an unnatural genus. Members of the genus are only loosely grouped together
by virtue of their scarious bracts and the usually conspicuous outer florets of the
capitulum. Close examination reveals an array of floret types (including capitula with
bisexual florets only; or with ligulate, irregularly ligulate or bilabiate outer female
florets as well as inner bisexual florets), bract morphologies, and habit differences.
Various chromosome numbers (« = 3, 7, 8, 9, 10, 11, 12; Turner 1967; Henderson
1969) also occur. Variation in the morphology of both bracts and corolla suggest that
Siemssenia Steetz [including P. capillaris (Steetz) Diels and P. microcephala] and
Panaetia Cass, [probably comprising Podolepis lessonii, P. muelleri (Sonder) G. L.
Davis, P. davisiana D. Cooke and P. tepperi (F. Muell.) D. Cooke] could be reinstated.
Similarly both P. georgii and P. kendallii (F. Muell.) F. Muell. should be excluded from
Podolepis s. str. Davis (1956) noted that both were atypical of the genus, and it has

75

Fig. 11. Transverse sections of fruit, a — Cephalosorus carpesioides (Short 2403). b — Gnephosis pygmaea
(Short 2385). c — Helipterum albicans subsp. albicans var. buffaloensis (Short 1403). d — Helipterum
charsleyae (Short 2003). e — Helipterum pygmaeum (Short 2973). f — Helichrysum apiculatum (Short
3027). g — Helichrysum cassinianum (Short 1575). h — Helichrysum leucopsideum (Short 1554). i —
Ixiolaena chloroleuca (Short 3003).

76

0.2mm

Fig. 12. Transverse sections of fruit, a — Myriocephalus guerinae {Short 540). b — Myriocephalus stuartii
(Czorney 1160). c — Pleuropappus phyllocalymmeus (Short 841). d — Podolepis arachnoidea (Willis
s.n., MEL 1554024). e — Podolepis gracilis (Willis s.n., MEL 61 1247). f — Podolepis microcephala
(Short 2050). g — Podolepis rugata (Short 1355). h — Toxanthes muelleri (Short 1251). i —
Triptilodiscus pygmaeus (Short 3014).

77

been noted that P. georgi is possibly conspecific with Helichtysum ayersii F. Muell.
(Grieve & Blackall 1975).

Davis {I.C., p. 249) stated that, with the exception of P. kendallii and P. georgei, in
which the fruits are ‘distinctive and diagnostic’, the similarity in all other species ‘is
almost an occasion for comment’. Externally the pericarp of most species is somewhat
similar, with the raised mucilage containing cells giving fruit a papillate appearance. A
well developed carpopodium is also present in most, if not all, species. However, there
are internal differences, the most obvious pertaining to the presence or absence of a
layer of sclerenchyma in the pericarp, and the thickness of such layers when present. Of
the ten species examined P. gardneri, P. gracilis (Figs lOd, 12e), P. lessonii and P.
microcephala (Fig. 12f) all lack a layer of sclerenchyma whereas P. arachnoidea (Figs
10c, 12d), P. auriculata, P. canescens, P. georgei, P. robusta and P. rugata (Fig. 12g)
possess one. The orientation of the vascular bundles in the pericarp varies from medial
in P. robusta and P. gardneri to almost lateral in P. rugata. In other species examined
the orientation is oblique. (Orientation of the vascular bundles could not be
determined in P. georgei & P. canescens.) With the exception of P. georgei vascular
bundles were not observed in the testa of any species. In the latter there are two bundles
within the testa, with each one being opposite a vascular bundle in the pericarp.

Although incomplete, observation of the fruit structure of species of Podolepis
support the notion that it is an unnatural genus and that Siemssenia and Panaetia
should be reinstated.

Rutidosis DC. (TS)

Transverse sections of fruit of only a single species, R. helichrysoides, the type of
the genus have been obtained (Fig. 10a). This species has an external layer of
sclerenchyma in the pericarp, the layer being usually two, but sometimes three or four
cells wide. Two vascular bundles (their position not determined) occur in the pericarp,
but bundles were not observed in the testa. Little cell detail could be discerned in a
narrow, highly stained testa but thick-walled cells seemed to be absent. Externally the
cypselae are covered with two-celled papillae which have their bases in the
sclerenchymatous. tissue. There is a prominent carpopodium.

The species of Rutidosis display an array of vegetative and inflorescence
characters which suggest that the genus is heterogeneous. Haegi (1986) retained R.
multiflora (Nees) Robinson in Rutidosis but indicated that were it not for the similar
fruit it would have been excluded. Fruit of the former species and R. helichrysoides are
similar in that both are obliquely attached to the corolla tube but an apparent absence
of a carpopodium has been noted in R. multiflora. If a carpopodium is present then it is
extremely small and concealed by basal papillae. The presence or absence of a
carpopodium is frequently indicative of different genera, thus supporting the
contention that R. multiflora should be excluded from Rutidosis.

Triptilodiscus Turcz. (TS)

For many years Triptilodiscus has been included in Helipterum DC. The single
species, T. pygmaeus [syn. H. australe (A. Gray) Druce], is readily distinguished from
species of Helipterum by a number of features of the inflorescence and the
reinstatement of the genus by Haegi (1986) is generally accepted. The fruit has two
medial vascular bundles in the pericarp but, in TS, bundles were not observed in the
testa. Paul Wilson (pers. comm. 1988) has recorded a single bundle in the testa. The
pronounced thickening of the outer walls of the testa (Fig. 12i) in this species has not
been observed in species currently included in Helipterum.

ACKNOWLEDGEMENTS

This work was funded by Australian Biological Resources Study grants to PSS
from 1 982 to 1 987. We thank Ms R. McFarlane, Mr T. Dewan and Mr A. J. Spies for
technical assistance and Dr D. B. Foreman, Mr J. Bruhl and Mr Paul G. Wilson for
constructive criticism of the manuscript. Mr J. Bruhl and Dr C. J. Humphries kindly
helped with the interpretation of fruit structure in the early stages of the survey.

78

APPENDIX

Species and voucher specimens from which fruit sections were obtained. All collections gathered by

P. S. Short unless otherwise indicated.

Genus and species

Collection Genus and species

Collection

Actinobole a. Gray
A. condensatum (A. Gray) P. Short

1005

A. drummondiana P. Short

2032

A. oldfieldiana P. Short

2013

A. uliginosum (A. Gray) H. Eichler

940

Angianthus

A. acrohyalinus Morrison
A. brachypappus F. Muell.

2045

Barker s.n., MEL 594984

A. conocephalus (J. Black) P. Short

Batt s.n..

MEL 84412

A. cunninghamii (DC. Benth.

Oldfield s.n..

MEL 84418

A. drummondii (Turcz.) Benth.

1102

A. glabralus P. Short

838

A. aff. micropodioides (Benth.) Benth.

Haegi 2664

A. milnei Benth.

2089

A. aff. milnei

1012

A. preissianus (Steetz) Benth.

842

A. tomentosus Wendl.

845

Cephalosorus a. Gray
C. carpesioides (Turcz.) P. Short

2403

Chondropyxis D. Cooke
C. halophila D. Cooke

Haegi 2565

Craspedia Forst. f.

C. aff. pleiocephala F. Muell.

837

C. unijlora Forst. f.

853

Dithyrostegia a. Gray
D. amplexicaulis A. Gray

2398

Epitriche Turcz.

E. demissus (A. Gray) P. Short

2391

Fitzwillia P. Short

F. axillifiorus (W.V. Fitzg. ex Ewart & J. White)

P. Short

2188

Gnephosis Cass.

G. pygmaea (A. Gray) Benth.

2385

Helichrysum Miller
H. apiculalum (Labill.) D. Don

3027

H. bracteatum (Vent.) Andrews

3022

H. cassinianum Gaudich.

1575

H. leucopsideum DC.

1554

Heupterum DC.

H. albicans (A. Cunn.) DC. subsp. albicans var.

buffaloensis P.G. Wilson

1403

H. charsleyae F. Muell.

2003

H. chlorocephalum (Turcz.) Benth.

1746

H. fioribundum DC.

3025

H. maryonii S. Moore

2644

H. microglossum (F. Muell. ex Benth.) Maiden &

Betche

2992

H. moschatum (Cunn. ex DC.) Benth.

2985

H. pygmaeum (DC.) Benth.

2973

H. strictum (Lindley) Benth.

1581

H. stuartianum Sonder & F. Muell. ex Sonder

2982

H. uniflorum J. Black

3002

Hyalochlamys
H. globifera A. Gray

1040

IxiOLAENA Benth.

I. brevicompta F. Muell. 3018

1. chloroleuca Haegi 3003

I. leptolepis (DC.) Benth. 2990

1. tomentosa Sonder & F. Muell. ex Sond. 2988

Lemooria P. Short
L. bukittii (Benth.) P. Short

1761

Leptorhynchos Less.

L. tetrachaetus (Schldl.) J. Black

3015

Millotia Cass.

M. myosotidifolia (Benth.) Steetz
M. tenuifolia Cass.

2128B

1753

Myriocephalus Benth.

M. gracilis (A. Gray) Benth.
M. guerinae F. Muell.

M. nudus A. Gray
M. stuartii (Sonder) Benth.

1018
540
1004/1569
Czorney 1160

Pluchea Cass.

P. tetranthera F. Muell.

3008

Pleuropappus F. Muell.

P. phyllocalymmeus F. Muell.

841

Podolepis Labill.

P. arachnoidea (Hook.) Druce

Willis s.n..

MEL 1554024

P. auriculata DC.

2033

P. canescens Cunn. ex DC.

2142

P. gardneri G. L. Davis

Smith 66/494

P. georgei Diels

Nelson 1754

P. gracilis (Lehm.) Grab. Willis s.n.

, MEL 61 1247

P. lessonii (Cass.) Benth.

2404

P. microcephala Benth.

2060

P. robusta (Maiden & Betche) J. H. Willis

Forbes 252

P. rugata Labill.

1355

PoGONOLEPis Steetz

P. muelleriana (Sonder) P. Short

843

P. stricta Steetz

2086

Rutidosis DC.

R. helichrysoides DC.

2999

ScYPHOcoRONis A. Gray

5. incurva D. Cooke

2001

Toxanthes Turcz.

T. muelleri (Sonder) Benth.

1251

T. sp. aff. muelleri

2353

Triptilodiscus Turcz.

T. pygmaeus Turcz.

3014

79

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237-253.

Short, P. S. (1986b). In Love, A. (ed.). Chromosome number report XCIl. Taxon 35: 610.

Short, P. S. (1987). A revision of Blennospora A. Gray (Compositae: Inuleae: Gnaphaliinae). Muelleria 6:
349-358.

Short, P. S. (1989a). A revision of Podotheca Cass. (Asteraceae: Inuleae: Gnaphaliinae). Muelleria 7:
39-56.

Short, P. S. (1989b). New genera and species of Australian Inuleae (Asteraceae) Muelleria 7: 103-1 16,
Steetz, J. (1845). Schoenia. In Lehmann, J. G. C. (ed.), ‘Plantae Preissianae’, vol. 1. (Hamburg), pp.
480-481.

Turner, B. L. (1967). Chromosome survey of Podolepis (Compositae-lnuleae). Aust. J. Bot. 15: 445-449.
Wilcox, B. H. ( 1 977). ‘A systematic study of the Leucanthemum-Chrysanthemum complex in North Africa’
Ph.D. thesis. University of Reading.

Wilson, P. G. (1960). A consideration of the species previously included within Helipterum albicans
(A. Cunn.) DC. Trans. Roy. Soc. S. Aust. 83: 163-177.

Wilson, P. G. (1987). Quinqueremulus linearis, a new genus and species in the Australian Asteraceae (tribe
Inuleae). Nuytsia 6: 1-5.

Manuscript received 30 March 1988

9.^

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TWO NEW SPECIES OF POMADERRIS Labill. (RHAMNACEAE) FROM
SOUTH-EASTERN AUSTRALIA

by

Neville G. Walsh*

ABSTRACT

Walsh, Neville G. Two new species of Pomaderris Labill. (Rhamnaceae) from
south-eastern Australia. Muelleria 7(1): 81-87 (1988). — Pomaderris gilmourii from
New South Wales and P. humilis from Victoria are described as new species and notes
on distribution, ecology and their relationship to other members of the genus are
provided. Two varieties of P. gilmourii are recognised.

INTRODUCTION

The genus Pomaderris contains a number of species which are morphologically
ill-defined and nomenclaturally confused. In the course of the preparation of a general
revision of the genus, a number of apparently clearly defined, undescribed taxa have
been encountered. As sufficient data pertaining to these taxa are gathered which
confirm their distinctness, they will be described in order that reference to them is
facilitated more quickly than would be the case if their publication were delayed until a
generic revision is completed.

This is the second recent paper describing new species in Pomaderris. See also
Muelleria 6: 6 (1988).

TAXONOMY

Pomaderris gilmourii N. G. Walsh, sp. nov.

Frutexad 4 m altum. Ramuli glabrescentes. Folia obovata, oblanceolata vel anguste elliptica, 8-30 mm
(raro ad 40 mm) longa, 4- 1 3 mm lata, apice acuto ad obtusum vel rotundato, pinnatinervia, nervorum
3-5 pares, supra glabra vel pubescentia secus costae, infra tomentosa trichomatibus stellatis. Stipulae
subulatae ad 4 mm longas, caducae. Inflorescentiae terminates, laxe paniculatae, pyramidales vel
rotundatae, 2-5 cm diametro, aliquantum abundans. Flores apetala, pedicellis 1-4 mm longis. Sepala
oblonga, apice acuto, pagina externa pilis brevis argenteis, interne glabra. Filamenta staminum
1-1-5 mm longa, Antherae oblongae vel ellipticae, 0-5-1 mm longae. Stylus c. 1 mm longus, trilobus,
divisus basi fere.

Typus: New South Wales — South Coast, Deua National Park, Prominence 1-9 km
north from Coondella trig, point, c. 16 km WSW. from Moruya, 35° 55' 50" S.,
1 49° 54' 20" E. Alt. 480 m, 7.xii.l987, A.G. Walsh 7559(Holotypus: MEL 1557601.
IsoTYPi: BRI, CBG, HO, NSW).

Shrub to 4 m high. Branchlets glabrescent, but covered when young by
semi-appressed to appressed simple hairs or tufted trichomes, with or without an
underlying hoary layer. Petiole 2-8 mm long. Lamina obovate, oblanceolate or
narrowly elliptic, 8-30 (rarely to 40) mm long, 4- 1 3 mm wide; apex acute to obtuse or
rounded; penninerved with 3-5 pairs of lateral veins which are inconspicuous above;
upper surface glabrous or with a line of short hairs above the midvein; lower surface
densely covered with fine stellate trichomes, with or without a superficial layer of
appressed, shining simple hairs. Stipules subulate, to 4 mm long, soon deciduous.
Inflorescences loosely paniculate, pyramidal or rounded, mostly 2-5 cm diam.,
terminal on the main axis and short lateral branches and rather prolific. Pedicels
1 -4 mm long. Sepals oblong, acute at apex, covered with short silver-grey hairs
externally, glabrous and cream-coloured on the inner face. Petals absent. Stamens
alternating with sepals; filament 1-1-5 mm long; anther c. 0-5-1 mm long. Ovary

‘National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

81

82

inferior, trilocular, the summit densely covered by erect simple hairs. Style c. 1 mm
long, divided almost to the base into 3 spreading arms. Capsules not seen.

Although the species is known from only, a few collections, specimens can be
readily segregated into two entities. These are here described as varieties although
future collections of the species may indicate that a reassessment of the taxonomic
rank of the two entities is warranted.

Key to the Varieties of Pomaderris gilmourii

1 . Indumentum of leaf undersurfaces, petioles, branchlets etc. with a dense superficial
layer of appressed simple hairs, imparting a shining appearance to those surfaces;
leaf margins thickened and glabrous, apparent as a distinct border on undersurface.

Sepals c. 1-5 mm long var. gilmourii

1. Indumentum not including a dense superficial layer or appressed simple hairs,
surfaces appearing dull; leaf margins neither thickened or glabrous, not apparent as
a border on either surface. Sepals c. 1 mm long var. cana

P. gilmourii N. G. Walsh var. gilmourii

Folia incrassata margine vel recurvata, supra glabrum omnino, infra cum pilis adpressis sericeis
nitentibus. Sepala c. T5 mm longa. Antherae oblongae c. 1 mm longae. Stylus sparsim pubescens infra
divisuram.

Leaves with margins thickened or minutely recurved, appearing bordered from
beneath; undersurface with fine stellate trichomes overlain by appressed, shining,
silky, simple hairs; upper surface quite glabrous. Sepals c. 1 ■ 5 mm \ong. Anthers oblong
c. 1 mm long. Style sparsely pubescent below point of division, glabrous above.
(Fig. l,a-d).

Other Specimens Examined (Total examined, 7):

New South (Tate — South Coast: Deua National Park, c. 6km N. of'Bendethra’, 35° 54' S., 149° 43' E
Alt. 700 m, 28.iii.1985, P. Gilmour 4988 (MEL 681685 and 671 106, Dupl. CBG 8505373). South Coasf
Deua National Park, c. 2km N. of Coondella trig., 35°56'S., 149° 54' E. Alt. 450 m, 25.xi.1985, />. Gilmour
8333 (MEL 68 1 668, Dupl. CBG 8505382). Deua National Park, near Mt Donovan, AMG 8826-62429 1 . Alt.
540 m, 28.111.1985, P. Beesley 376&D. fimns (MEL 672076). From type locality, 7.xii.l987,Af.G. Walsh 1877
and /590 (MEL 1557604 and 1557609 respectively).

P. gilmourii var. cana N. G. Walsh var. nov.

Folia piano margine, supra pubescentia secus costae. Sepala c. 1 mm longa. Antherae ellipticae,
c. 0-5 mm longae. Stylus pubescens ultra divisuram.

Typus; New South Wales — South Coast, Deua National Park. Prominence 1-9 km
north from Coondella trig, point, c. 16 km WSW. from Moruya, 35° 55' 50" S.,
149° 54' 20" E. Alt. 480 m, 7.xii.l987, N.G. Walsh 7576 (Holotypus: MEL 1557595.
IsoTYPi: BRI, CBG, HO, K, NSW).

Leaves with plane margins, not appearing bordered from beneath; tomentum of
lower surface of stellate trichomes only, or if a few simple hairs present, then these
neither appressed nor shining; upper surface minutely pubescent above the midvein.
Sepals c. 1 mm long. Anthers elliptic c. 0-5 mm long. Style pubescent beyond the point
of division. (Fig. 1, e-g).

Other Specimens Examined (Total examined, 2):

New South Wales — South Coast: Deua National Park, c. 2 km N. of Coondella trig., 35°56'S.,
149° 54' E. Alt. 450 m, 25.xi.1985, P. Gilmour 5327 (MEL 684706, 681669, Dupl. CBG 8505378).

Distribution and Conservation Status;

P. gilmourii var. gilmourii is currently known only from Deua National Park and
its immediate environs, within an area of only about 1 5 km diameter. The land system
supporting P. gilmourii is poorly known however and further populations may be

83

Fig. 1. Pomaderris gilmourii var. gilmourii. a — branchlet in bud and flower, X 1. b — undersurface of leaf
X 3; inset X 9. c — flower X 8. d — style X 1 2. Z". gilmourii var. cana. e — undersurface of leaf X 3; inset
X9. f — flower X8. g — style X 12. a-d drawn from N.G. Walsh 1889 (type); e-g drawn from N.G.
Walsh 1876 (type).

84

expected to be discovered in the course of detailed survey of the general area. P.
gilmourii var. cana is known only from a few plants at the type locality where it occurs
with the typical variety. The species conservation status has been assessed as 2RC-t
(Briggs & Leigh 1988), that is the species is rare (‘R’), represented within a
conservation reserve (‘C’), the population size is unknown but all known plants
are reserved (‘t’).

Habitat:

P. gilmourii occurs on skeletal soils derived from rhyolite, an igneous rock, in this
area formed as part of the Comerong Volcanic series (Gilligan 1974). Most sites are
exposed, on or atop steeply sloping rock faces supporting shrubland or open woodland,
although one collection {Gilmour 4988) is apparently from a more sheltered site
supporting open forest. Recorded altitudes range from 450 m to 700 m. Associated
species typically include a number of similarly localised, rhyolite-endemic species
such as the recently described Prostanthera porcata, Westringia saxicola and an
undescribed Leptospermum as well as other species characteristic of clifftop shrubland
and woodland communities, e.g. Eriostemon trachyphyllus, Platysace lanceolata,
Hakea dactyloides, H. macreana, Kunzea ambigua and Eucalyptus stenostoma.

Notes:

P. gilmorii is a very distinctive species unlikely to be confused with any other
Pomaderris in the eastern states. However, sterile specimens somewhat resemble P.
myrtilloides, a species of calcareous, mostly coastal sites in southern Western Australia
and it was to this species which I very hesitantly referred the first (sterile) material of P.
gilmourii I saw. Unlike P. gilmourii though, the western species is not apetalous and
has a style which is barely cleft. The indumentum of P. gilmourii var. gilmourii is
suggestive of that of P. ledifolia, a species also typically associated with exposed, rocky
peaks, but the characteristic very narrow leaves and crowded, petalous flowers of that
species readily distinguish it from P. gilmourii. P. cinerea, a species endemic to the
south coast region of New South Wales and which also occurs within Deua National
Park shares with P. gilmourii a number of features which may indicate a closer
relationship than the general appearance of the two species suggests. Both species
flower later in the year than any other Pomaderris in the broad area, both are apetalous
and perhaps most significantly, both species have a pubescent style, a feature not
known from other members of the genus. The general nature of the indumentum of P.
cinerea and its overall dusky appearance are reflected to some extent in P. gilmourii
var. cana.

The specific epithet honours Mr Phil. Gilmour, formerly of Canberra, who first
collected this species and whose collections comprise the majority of herbarium
specimens of it. His botanical surveys of the largely unexplored south coast area of
New South Wales in general and the rugged and largely inaccessible rhyolite country in
particular, have unearthed a number of new species and improved our knowledge of
many rare and restricted plants.

The varietal epithet cana refers to the dull, greyish appearance of that variety, in
contrast to the brighter overall aspect of the typical variety.

Pomaderris humilis N. G. Walsh, sp. nov.

Frutex decumbens vel infirme ascendens, plerumque ad 0-5 m altum. Ramuli stellato-tomentosi cum
pilis simplicibus vel trichomatibus caespitosis longioribus. Folia elliptica vel ovata rare obovata,
10-50 mm longa, 7-25 mm lata, apice rotundato versus late acutum, pinnatinervia, nervorum 5-7
pares, supra pilosa cum pilis simplicibus suberectis raro cum stellatis vel bifidis trichomatibus
paucibus, c. 0-2 mm longis, infra stellato-tomentosa cum pilis simplicibus longioribus praesertim in
costis et nervis. Stipulae lanceolatae, ad 4 mm longas, caducae. Injlorescentia terminales, paniculatae,
pyramidales plerumque 3-5 cm diametro. Sepala oblonga, 2-2-5 mm longa, apice acuto, pagina
externa stellato-tomentosa cum pilis simplicibus longioribus, interne glabra. Petala spathulata,
breviora parum sepalis, margine apicali irregulatim crenato. FUamenta staminum 2-2-5 mm longa.
Antherae oblongae, c. 1 mm longae. Stylus 1-5-2 mm longus, integer fere vel divisus j parte sui
longitudinis. Capsulae ovoideae, c. 3-5x2-5 mm.

85

Typus: Victoria — Gippsland, Holey Plains State Park, Holey Hill, beside ‘Banksia
Forest’ walk, 0-3 km WNW. from fire tower on summit, 14 km south-east from
Rosedale, 38°13'S., 146°57'E. Alt. c. 170m, l.x.1986, N.G. Walsh 1616
(Holotypus: MEL 1556333. Isotypi: CANB, HO).

Decumbent or weakly ascending shrub, mostly to c. 0-5 m high, rarely to 1-6 m
high. Branchlets and petioles closely stellate-tomentose with an overlying layer of
longer, simple hairs or tufted trichomes. Lamina elliptic or ovate, rarely obovate,
10-50 mm long, 7-25 mm wide; apex rounded to broadly acute; penninerved with 5-7
pairs of lateral veins; upper surface pilose with erect or slightly antrorsely inclined
simple hairs c. 0-2 mm long, rarely also with a few bifid or stellate trichomes; lower
surface pale, with a dense mat of minute stellate trichomes and some longer
(c. 0-5 mm) simple hairs, those of the midrib and lateral veins longer (to 1 mm) and
more numerous. Stipules lanceolate, to 5 mm long, soon deciduous. Inflorescences
paniculate, pyramidal, mostly 3-5 cm diam., terminal. Pedicels 2-3 mm long. Sepals
spreading, oblong, 2-2-5 mm long, acute at apex, covered externally with a dense
stellate tomentum with longer simple hairs, these dense on the thalamus tube; inner
face creamy yellow, glabrous. Petals erect, pale yellow, spathulate, slightly shorter than
sepals, the apieal margin irregularly crenate. Stamens opposite the petals; filament
2-2-5 mm long; anther oblong, c. 1 mm long. Ovary inferior, trilocular, the summit
densely covered by erect hairs. Style 1-5-2 mm long, almost entire or divided in the
upper one-third into three spreading arms. Capsules ovoid, c. 3-5 x 2-5 mm. Seeds
elliptic, dorsally rounded, c. 2-5 x 1-5 mm. (Fig. 2).

Other Specimens Examined (Total examined, 6):

Victoria — from type locality: 2.ix.l949, 7.D. Sherwood s.n. (MEL 55064); 6.iii.l973,.4.C. Beauglehote
41615 (MEL 517247, Dupl. NSW); 28.ix.I974, J.H. Willis s.n. (MEL 91867); 14.vi.l986, N.G. Walsh 1602
(MEL 1 549453, Dupl. NSW, HO); Gippsland, Rosedale, Cheshum Rd, c. 8 miles south of town, ofFLimepit
Rd, 18.x. 1969, B. Thompson s.n. (MEL 504689).

Distribution and Conservation Status:

Known with certainty only from the type area, the population there consisting of
an estimated 2000-5000 plants. A specimen in early bud [Beauglehole 37541 (MEL
5 1 7249, Dupl. NSW)] from near Bruthen (c. 1 00 km ENE. from the type locality), may
be P. humilis but the buds are insufficiently developed for a confident assignation of
the specimen. Surveys of the area during the 1986 and 1987 flowering seasons failed to
locate plants matching the Beauglehole (37541) specimen. Both the Bruthen site and
Holey Hill are formed on late Tertiary marine sediments (Geological Survey of
Victoria 1971, 1977). Local edaphic and general conditions in the two areas are very
similar and the two sites have many species in common.

The conservation status of P. humilis has been assessed as 2RCat (Briggs & Leigh
1 988), that is the species is rare, represented in a conservation reserve, the population
exceeds 1000 individuals (a) and the total population occurs in a conservation reserve.
A number of plants of P. humilis at the northern fringe of the population were
apparently removed in the early 1 970s in a clearing operation to make way for a Pinus
radiata plantation (Mrs B. Thompson, South Traralgon, pers. comm.).

Habitat:

P. humilis occurs on coarse, sandy soil derived from late Tertiary marine
deposits. The vegetation type is a Eucalyptus consideniana-E. globoidea open forest
with a tall shrub layer dominated by Banksia serrata and a heathy low shrub and
ground layer including Brachyloma daphnoides, Monotoca scoparia, Banksia
marginata, Grevillea chrysophaea and Lepidosperma concavum.

Notes:

Pomaderris humilis has in the past (e.g. Beauglehole 1 980) been referred to P. aff.
velutina. It bears a superficial resemblance to P. velutina J. H. Willis in the somewhat
velvety upper surfaces of the leaves, but the hairs on the upper surfaces of the leaves of

86

Fig. 2. Pomaderris humilis. a — flowering branchlet X 1. b — stipules and leaf base X4. c — upper surface of
leaf X 1 0. d — lower surface of leaf X 1 0; flower X 6. a, c, d and e drawn from the holotype; b drawn from
Beauglehote 41615 (MEL 517247).

87

P. humilis differ in being predominantly simple and substantially longer and sparser.
Furthermore P. velutina has few, thread-like simple hairs confined to the midrib and
larger veins on the undersurfaces of the leaves and flowers with relatively broad petals
and deeply trifid styles, whereas P. humilis has copious simple hairs on both the veins
and intemerves of the lower leaf surface, and flowers with narrow petals and shallowly
cleft styles.

P. humilis is probably more closely allied to P.^aurea Wakefield and the
widespread P. lanigera (Andrews) Sims. From the former it differs conspicuously in
the simple pubescence of the upper leaf surface and narrow petals and from P. lanigera
in the finer tomentum on the upper surface of the leaves, the sparser, less spreading
pubescence beneath, the narrow petals, shallowly cleft style and the relatively small,
pyramidal inflorescence {P. lanigera characteristically has a broad, hemispherical or
flat-topped inflorescence).

From virtually all eastern state Pomaderris species, P. humilis is distinguished by
its largely decumbent growth habit, whence the specific epithet is derived. This growth
habit has been maintained in cultivated plants growing in East Malvern (Victoria) for
more than two years. Occasional wild-growing plants occurring amongst dense
vegetation may attain 1-6 m in height but this growth is usually rather spindly. The
growth habit of the plant near Bruthen from which Beauglehole 37541 was collected is
unknown.

P. humilis is the taxon referred to as "Pomaderris sp. (Holey Hill)’ in Forbes and
Ross (1988).

ACKNOWLEDGEMENTS

I wish to thank Mr Phil Gilmour for his efforts in procuring specimens of P.
gilmourii on my behalf; Jan, Kate and Rick Walsh for their cheerful company in the
field; Mrs Bon Thompson of Traralgon South for information pertaining to P. humilis-,
and my colleague. Miss Anita Podwyszynski who prepared the illustrations.

REFERENCES

Beauglehole, A. C. (1980). ‘Victorian Vascular Plant Checklists’. (Western Victorian Field Naturalists Clubs
Association: Portland.)

Briggs, J. D. & Leigh, J. H. (i988). Rare or threatened Australian plants: The 1988 revised edition. Austral.
Natl Parkes & Wildlife Serv. Special Publ. 14.

Forbes, S. J. & Ross, J. H, (1988). ‘A census of the vascular plants of Victoria. Second edition’. (National
Flerbarium of Victoria: Melbourne.)

Geological Survey of Victoria (1971 and 1977). Australia 1:250,000 geological map series, Warragul and
Baimsdale, sheets SJ55-10 and SJ55-7 respectively. (Govt. Printer: Melbourne.)

Gilligan, L. B. (1974). Canberra 1:250,000 metallogenic map. SI-55-16. Geol. Surv. N.S.W., Sydney.

Manuscript received 1 3 May 1 988

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A NEW SUBSPECIES OF GREVILLEA ACANTHIFOLIA
(PROTEACEAE:GREVILLEOIDEAE) FROM SOUTH-EASTERN
NEW SOUTH WALES

by

R. O. Makinson* and D. E. Albrecht^

ABSTRACT

Makinson, R. O. & Albrecht, D. E. A new subspecies of Grevillea acanthifolia A. Cunn.
(Proteaceae : Grevilleoideae) from south-eastern New South Wales. Muelleria 7(1):'
89-93 (1989). — A new subspecies of Grevillea acanthifolia spp. paludosa R.
Makinson & D. Albrecht, from the Nalbaugh Pleateau is described and illustrated,
with notes on distribution and habitat, and a key to subspecies of G. acanthifolia.

INTRODUCTION

Material collected by P. Gilmour and D. Albrecht on the Nalbaugh Plateau in
February 1 987 was found to be closely related to G. acanthifolia, ssp. acanthifolia and
to G. acanthifolia ssp. stenomera (F. Muell. ex Benth.) McGillivray, which occur
respectively in the Central and Northern Tablelands of New South Wales, and shares
features with each that make subspecific rank appropriate.

Terminology and presentation follows that used by D. J. McGillivray in his
revision of Grevillea (in prep.).

TAXONOMY

Grevillea acanthifolia A. Cunn. ssp. paludosa R. Makinson et D. Albrecht, ssp. nov.

a Grevillea acanthifolia A. Cunn. subsp. acanthifolia lobis foliorum angustioribus et inflorescentiis
brevioribus, a G. acanthifolia ssp. stenomera (F. Muell. ex Benth.) McGillivray ramulis subsericeis et
statura excelsiore, ab ambosus subspeciebus pilis adpressis plerumque praesentibus infra laminarum
foliorum affinibus costae et donatoribus pollinis conicis validus et erectis plus differt. (Fig. 1).

Typus: c. 2-5 km WNW. ofMt Wog Wogtrig., Nalbaugh National Park, 37° 05' 15" S.,
149° 24' E., rare (12 plants seen) in peaty drainage line. Shrub c. 3 m high and 5 m
diameter. Associated spp. include Boronia deanei, Gahnia radula, Leptospermum
lanigerum, Tetrarrhena acuminata, Schoenus maschalinus, Gleichenia dicarpa,
Bauera rubioides. Acacia costiniana, 22.ii.1987, D.E. Albrecht 3078 & P. Gilmour
(bud, 11., fr.). (Holotypus: MEL 1556925; Isotypus: NSW).

Prickly spreading shrub to 3 m high and 5 m wide, with ascending to decumbent
branches and entangled foliage. Branchlets angular in cross-section, longitudinally
ridged, subsericeous (sometimes the ridges almost glabrous). Leaves subsessile
(appearing petiolate — laminal tissue below basal lobe-pair very narrow), deeply
pinnatipartite, (35-)40-60 mm long, 40-70 mm wide (when pressed flat), with 5-9
divaricate primary lobes, primary lobes usually 3-7-partite; secondary lobes
divaricate, usually entire or occasionally bipartite; ultimate lobes subulate to
sublinear, (10-)12-18(-20) mm long, 1 •5-2-5 mm wide, rigid, acute, cuspidate with a
pungent point 1-2 mm long; margin refracted, usually some undersurface of lamina
visible on either side of the midribs of leaf and lobes; upper surface glabrous; lower
surface subsericeous to glabrous on either side of the midribs; midribs glabrous or with
scattered more or less appressed hairs; texture coriaceous. Inflorescences terminal,
profuse, conspicuous, erect, pedunculate, simple, many-flowered, secund, acropetal,
30-50 mm long; peduncles 5-10 mm long, tomentose; peduncle and rhachis
subvillous with very pale spreading hairs, hairs darker and more appressed below the

* National Herbarium of New South Wales, Mrs Macquaries Road, Sydney, NSW, Australia 2000.
♦National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

89

90

Fig. 1. Grevillea acanthifolia ssp. paludosa. a — Flowering branch, XO-8. b — Undersurface of leaf, X2 and
(inset) showing distribution of hairs, c — Mature fruit, X2-5. d — Inner face of mature seed, X4. e —
Mature flower, X3-5. f — Section through mature flower, X3-5. g — Floral bract, X5. Drawn from
holotype, (D.E. Albrecht 3078 and P. Gilmour).

91

lowermost peduncular bract or bract scar; floral bracts broadly ovate, apiculate,
2-3-5 mm long, 2-5-4-5 mm wide, outer surface tomentose, inner surface glabrous
except for few to many ascending hairs along the midline or towards the apex, bracts
conspicuous on young inflorescences, usually deciduous when buds are c. 6-10 mm
long. Pedicels 1-0-1 -5 mm long, villous; torus oblique at c. 15°-25° to the transverse,
1 -2-1 -5 mm from ventral to dorsal side. Perianth narrowly and obliquely ovoid below
the subapical curve, outer surface subsericeous to tomentose below the curve, villous
on the ellipsoid limb, inner surface glabrous; nectary U-shaped, partly enclosed within
the torus, pistil 21-24 mm long; stipe 0-5- 1-2 mm long, subsericeous; ovary
1-2-1 -6 mm long, villous; style glabrous; pollen-presenter a more or less erect cone
with a basal rim, ± ovate or round in plan view, 0-3- 1-0 mm long and wide, base
oblique at c. 10°-20° to the transverse, cone 0-8-1 -0 mm high. Fruit a follicle,
appressed to the rhachis by inward curving of the stipe, obliquely ellipsoid,
8-5-10-5 mm long, c. 6-7-5 mm wide, c. 5-6 mm thick; style persistent; surface
villous-velutinous, with a short dense indumentum of stout erect hairs, indumentum
with purplish longitudinal stripes and blotches (formed by hairs with dark cell
contents), and an emergent loose indumentum of longer pale spreading hairs; pericarp
c. 0-3-0-4mm thick, weakly crustaceous. Seeds {Albrecht 3078) obliquely ellipsoid,
6-7 mm long, c. 3-5 mm wide, 1-5-2 mm thick; outer face convex, somewhat rugose;
inner face with a peripheral ridge of waxy material, and an inner ridge of harder tissue
surrounding a central elliptical area c. 2-5-3-5 mm long and 1-1-5 mm wide; apical
elaiosome subtriangular, c. 1 mm long, of waxy material similar to and connecting
with the peripheral ridge on the inner face. Buds with limb blue-grey, overlain with
cream hairs; tepals below the limb cream-green in early bud, becoming blue-grey near
(and after) anthesis. Ovary covered with white hairs. Style (emergent from dorsal side
of perianth in late bud) rich puce-pink in bud, paling slightly at and after anthesis.
Style-end and pollen-presenter green.

Distribution and Conservation Status:

G. acanthifolia ssp. paludosa is known by only 41 plants in two drainage lines on
Nalbaugh Plateau (which extends from Mt Wog Wog to White Rock Mountain),
c. 45 km inland from Eden, south-east New South Wales. Nalbaugh Plateau is situated
within Nalbaugh National Park, hence all known individuals are secured within a
biological reserve. Figure 2 shows the locations of populations seen on the plateau.

Fig. 2. Occurrences of G. acanthifolia ssp. paludosa on Nalbaugh Plateau (•);
boundaries.

swamp

92

Ecological Notes:

Most plants (34 of the 41 known individuals) were found in a structurally and
floristically heterogeneous swamp community in the headwaters of the White Rock
River catchment at an elevation of 975-1005 m. The swamp system is underlain by
Nalbaugh granite and has a distinctive microtopography consisting of peat and
Sphagnum, with flat depressions; occasional shallow pools; emergent better drained
and densely vegetated hummocks, and occasional runnels of quartzitic gravel. Depth
of the surface layer of peat at one population was 50 cm. Plants of G. acanthifolia ssp.
paludosa appeared to be restricted to the hummocks, which were shared with and
surrounded by, Leptospermum scoparium, Gahnia sieberiana, Boronia deanei,
Sprengelia incarnata, Gleichenia dicarpa, Pultenaea subumbellata and Bauera
rubioides. The height of the community ranged from one to three metres and is
surrounded by a dense Leptospermum lanigerum dominated scrub to c. 10 m high.

Most of the remaining few plants were growing in dense Leptospermum
lanigerum thickets that fringe the free-flowing creek downstream from the swamps.
The substrate in these localities differed in being rocky and gravelly with loam pockets.
Two isolated plants were seen in similar streamside vegetation on the separate Wog
Wog River catchment; the swamp upstream of this at present lacks any plants despite
its close similarities to the White Rock River swamps.

The age structure of populations was relatively uniform. Judging from the stem
diameter and size of plants (the smallest seen was 2 m high and c. 1-5 m diameter),
most plants appear to be quite old. Although most plants appeared to have set a
considerable amount of seed, no seedlings were observed.

Like other closely related taxa, G. acanthifolia ssp. paludosa is probably fire
sensitive and principally regenerates from seed. It may however have the potential to
resprout as occasional post fire resprouting from the base of the stem has been
observed in populations of G. acanthifolia ssp. acanthifolia in swamps of the Lithgow
area (D. H. Benson, pers. comm.).

The introduced honey bee {Apis mellifera) and two species of honeyeater (the
White-eared Honeyeater and the New Holland Honeyeater) were seen feeding on the
flowers of the Grevillea.

Key to the Subspecies of G. acanthifolia

1 . Ultimate leaf lobes linear or subulate; most of undersurface of leaf enclosed by
refracted margin; inflorescences usually <5 cm long.

2. Undersurface of leaves and lobes glabrous except for scattered appressed hairs
on the midribs; branchlets glabrous or occasionally loosely tomentose; stipe
>1-5 mm long; pollen-presenter oblique, broadly conical; low spreading shrub

usually < 1 m high ssp. stenomera

2. Undersurface of some or all leaves and lobes with white appressed hairs on the
lamina either side of the midrib; branchlets subsericeous (ridges sometirnes
glabrous); stipe <1-5 mm long; pollen-presenter an erect cone; erect spreading

shrub attaining 3 m in height ssp. paludosa

1 . Ultimate leaf lobes cuneate or ovate, rarely linear; most of undersurface of leaf

lobes exposed, margin recurved; inflorescences usually > 5 cm long

ssp. acanthifolia

Affinities

This new subspecies closely resembles in leaf division and lobe shape ssp.
stenomera from the eastern part of the New England area of New South Wales, but the
latter is a less hairy shrub and does not appear to much exceed 1-5 m in height.

The erect, strongly conical pollen-presenter of ssp. paludosa is not shared with
either ssp. stenomera or ssp. acanthifolia, which usually have oblique, broadly conical
to convex pollen-presenters, although occasional flowers of the latter subspecies have
pollen-presenters oblique at as little as c. 1 0°- 1 5° (more usually 20°-30°). The basal rim
or ‘brim’ of the pollen-presenter in ssp. paludosa is somewhat oblique (to 20°) but the
axis of the cone in relation to the upper few millimetres of the style is quite erect.

93

Fig. 3. Distribution of G. acanthifolia subsp. paludosa and some related taxa in New South Wales. G.

acanthifolia ssp. acanthifolia □, G. acanthifolia ssp. stenomera 0, G. acanthifolia ssp. paludosa ■,

G. rivularis t.

Like ssp. paludosa, the other two subspecies of G. acanthifolia are also usually
found in swampy heath communities, although ssp. stenomera is occasionally found
on drier woodland slopes.

In habit and leaf shape, ssp. paludosa is strongly reminiscent of the closely related
G. rivularis L. Johnson & McGillivray which also grows ‘with its feet wet’, in the flood
zone of a creek near Fitzroy Falls on the N.S.W. Central Tablelands. G. rivularis is
however an almost glabrous plant.

Figure 3 shows the distribution of ssp. paludosa in relation to these closely related
taxa. The great geographic separation of these taxa implies a long term separation of
the original stock and subsequent divergence to the present taxa. It is interesting to
speculate why ssp. paludosa occurs where it does and why it is so rare. Although the
Nalbaugh peat deposits may not span the last ice age (P. Cell, pers. comm.), analysis of
the pollen profiles of cores extracted from the Nalbaugh swamps may provide some
additional informative data on fire history of the plateau and relative changes in the
abundance of ssp. paludosa over the last few thousand years.

Further Specimen Seen:

New South Wales: Southern Tablelands — Plateau between Mt Wog Wog and White Rock Mountain,
‘Long Swamp’, catchment of White Rock River, 8.xi.l987, R.O. Makinson 322(bud-fl.) (NSW, BRI, CANB,
CBG, MEL, NE).

ACKNOWLEDGEMENTS

We wish to thank Mr Don McGillivray for comments on the specimens; Mr P.
Gilmour and Mr W. M. (Bill) Molyneux for assistance with field work; Mr D. H.
Benson, Mr N. G. Walsh and Mrs K. L. Wilson for comments on the manuscript;
Miss A. M. Podwyszyinski for the illustration; and Mr N. G. Walsh for the Latin
diagnosis.

Manuscript received 14 June 1988

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A NEW SPECIES OF TETRARRHENA R. Br. (POACEAE) FROM VICTORIA

AND NEW SOUTH WALES

by

Neville G. Walsh*

ABSTRACT

Walsh, Neville G. A new species of Tetrarrhena R. Br. (Poaceae) from Victoria and
New South Muelleria 7(1): 95-98 (1989). — Tetrarrhena turfosa is described as
a new species with notes on distribution and ecology. Its relationship to other members
of the genus is discussed.

INTRODUCTION

In the course of preparing the account of Poaceae for a forthcoming Flora of
Victoria, several apparently unnamed taxa have been encountered. The majority of
these are in groups currently under study by others and should, in due course, be dealt
with by them. The species described herein has long been recognised as being distinct
but has evaded formal recognition. As it seems no specialists are presently dealing with
Tetrarrhena, the opportunity is here taken to validate the status of a sixth member of
the genus.

TAXONOMY

Tetrarrhena turfosa N. G. Walsh, sp. nov.

Gramen perenne, rhizomatosum, caespitosum vel ascendens, 0-2-1-3 m altum. Folia erecta, laevia et
glabra. amplexicaules. L/gw/aeciliatae, adO-5 mmlongae. Lam/'naeinvolutae, 2-7 cmlongae,

0-3-0-8 mm latae, obtusae interdum inflatae apicibus. Inflorescentia racemosa, angusta, erecta, spicam
simulans, 1-3 cm longa. Spiculae 3-10, subsessiles, saepe purpuratae, 4-8-6-8 mm longae. Glumae
subaequales, ovatae, 1-2 mm longae. Lemma sterilis infemum longitudine circa j partes lemmatis
sterilis superni, ambo oblongae, obtusae, carinatae vix, nervi 5-7 ellevati manifeste, scabri. Lemma
fertilis aequans fere lemma sterilem supemum, carinatum, scaberulum. Palea aequans fere lemma
sterilem. Antherae quatuor, circa 3 mm longae.

Typus: Victoria — Western. Grid D 18. Grampians, 3 miles (6-4 km) SW. of Halls
Gap, 0- 1 5 miles (0-24 km) west of junction with Mt Rosea Track, along watercourse.
Associated species include: Pultenaea subumbellata, Sprengelia, Selaginella, Restio
complanatus, Lepidosperma spp. Gymnoschoenus, Gahnia sieberiana, 18.1.1969,
Beauglehole 30309 (Holotypus: MEL 597060. Isotypi: AD, BRI, CANB, HO,
NSW).

A rhizomatous, perennial grass, forming compact tufts, commonly to c. 0-6 m
high in exposed sites, or with leafy, branched, ascending strands to 1 - 3 m high amongst
taller vegetation. Leaves erect, smooth and ^abrous. Sheaths tightly encircling stem.
Lamina tightly involute, 2-7 cm long, 0-3-0-8 mm diameter, terminating in a blunt,
sometimes slightly swollen tip. Ligule a dilate rim to 0-5 mm long, sometimes with a
few marginal hairs to 1 mm long. Inflorescence an erect, spike-like raceme 1-3 cm
long. Spikelets 3-10 per raceme, 4-8-6-8 mm long, subsessile, often purplish. Glumes
subequal, the upper usually slightly larger, ovate, 11-2 mm long, smooth and
glabrous. Lower sterile lemma about j as long as upper, both oblong, blunt, hardly
keeled, the 5-7 nerves prominently raised and scabrid. Fertile lemma almost equal to
upper sterile lemma, keeled, uniformly scaberulous, obscurely 5-7 nerved. Palea
about as long as lower lemma, membranous. Anthers 4, about 3 mm long. (Fig. 1).

Selected Specimens Examined (Total number examined 43):

New South Wales — Barrington Tops, swamps and grasslands, 7.i.l934, Vickery (NSW 115676).

•National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

95

96

Fig. 1. Tetrarrhena turfosa. & — habn, X^; inset — leaf tip X 5. b — spikelet XlO.from Beauglehole 30309
(type). Inset — distribution of T. turfosa (n.b. each dot may represent more than one collection); places
represented are A “Ararat, M = Melbourne, B==Mt. Bogong, 0 = Orbost, W = Wollongong,
S = Sydney, N = Newcastle.

97

Centennial Park, vi.l897, Cornfield s.n. (NSW 115689). Katoomba, swampy places, alt. 3100 ft, 22. i. 1939,
Blake i3920(NSW 1 15671). Bridal Veil Falls, Blackheath, alt. 3450 ft. Constable s.n. (NSW 56320). 1-5 miles
N. of Hilltop near Mittagong, 6.ii.l965, McBarron 10538 (NSW 1 15680).

Victoria — Mt Buffalo, Crystal Brook, near ‘Tucker Box’ corner, 21. ii. 1963, Willis s.n. (MEL 113885).
Bogong High Plains, 20.i.l940, Patton s.n. (MEL 597061). Nunniong Plateau, c. 0-5 ml. SW. of Reedy River
Chasm, 3.ii.l973, Beauglehole 41349 (MEL 527466). Mallacota-Wingan coast, 0-9 ml. NE. of Little River
Mouth, 22.xii. 1 969, Beauglehole 32695 (MEL 564478). Erica district, beside Beynons Road, c. 3 km south of
Morgans Mill, 10.i.l980, Scarlett 80-3 (MEL 596706).

Distribution and Conservation Status (Fig. 1):

Occurs mostly on and seaward from the Dividing Range from as far north as the
Barrington Tops area in New South Wales south to the Victorian border and west to
The Grampians in south-western Victoria. The species is locally common in
appropriate habitats, although some populations (e.g. near Mt Wog Wog in
south-eastern NSW, and subalpine to subalpine sites in Victoria) are disjunct and
would appear to be quite small. Fortunately the species is well reserved in national
parks in both states and its conservation status is therefore considered to be
secure.

Ecology;

The grass is invariably associated with heathy and sedge-rich vegetation in
swamps and fringing watercourses from near sea-level in eastern Victoria to subalpine
situations (to c. 1650 m) in both states. Soils are typically sodden and peaty, chiefly
derived from or formed upon sandstone, but on Mt Buffalo and the Bogong High
Plains the parent materials are granite and basalt respectively. Commonly associated
plants are typified by those species accompanying the type collection, i.e. with strong
representation of the Epacridaceae, Cyperaceae, Restionaceae and, particularly at
higher altitudes. Sphagnum mosses. The main flowering and fruiting period is from
November to February.

Notes:

T. turfosa is the taxon first recorded as an apparently undescribed species by
Willis (1970) and subsequently by Beauglehole (1980) and Forbes and Ross (1988) as
Tetrarrhena sp. It would appear to be most closely allied to T. acuminata R. Br. (near
which it occasionally occurs) and the recently described T. oreophila D. I. Morris of
Tasmanian alps and subalps to which it bears a strong superficial resemblance. From
T. acuminata, it differs primarily in the shorter (<7 mm) spikelets, the obtuse, not
acuminate sterile lemmas and the smooth, inrolled, not scabrous or flat leaf
blades.

From T. oreophila it is distinguished by the obtuse, strongly scabrous and
prominently 5-7 nerved sterile lemmas, in contrast to those of the Tasmanian
endemic which are acute to acuminate, minutely scaberulous or almost smooth and
lacking prominent nerves.

Material of T. turfosa at NSW had been segregated as an ecological variant of T.
juncea R. Br. Examination of the types from BM and K (including the type of T.
tenacissima, a later synonym), and all specimens of T. juncea at MEL and NSW
confirms that there is no continuity of variation from that species to T. turfosa. T.
juncea, a forest species, is infamous for its harshly scabrous, leaves and wiry stems and
differs otherwise from T. turfosa in its relatively longer glumes, and more tapered,
virtually smooth lemmas. Both T. juncea and T. oreophila are atypical in the genus
(and belie the feature on which the generic name is conferred) in possessing 6 and 2 (or
1) anthers respectively rather than the ‘typical’ tetrandrous condition.

The two other species of Tetrarrhena are T. laevis R. Br., confined to the
south-west of Western Australia and T. distichophylla R. Br., which typically has
pubescent spikelets and leaves and is a species of poor, dryish country in southern
Victoria, south-eastern South Australia and Tasmania.

The specific epithet "turfosa’ (from a peat bog) pertains to the species well-defined
preference of habitat.

98

ACKNOWLEDGEMENTS

I am grateful to the Curators of BM and K for the loan of type material, to the
Direetor and staff of NSW for assistanee during a visit to that institution, to my
colleague Mr David Albrecht who examined collections of Tetrarrhena at HO, and
Miss Anita Podwyszynski who prepared the illustration.

REFERENCES

Beauglehole, A. C. (1980). ‘Victorian Vascular Plant Checklists’. (Western Victorian Field Naturalists Clubs
Association: Portland.)

Brown, R. (1810). Prodromus Florae Novae Hollandiae. (Taylor: London.)

Forbes, S. J. & Ross, J. H. (1988). ‘A census of the vascular plants of Victoria. Second edition’. (National
Herbarium of Victoria: Melbourne.)

Morris, D. I. (1977). Miseellaneous notes on endemic Tasmanian plants. Records of the Queen Victoria
Musdum 55: 3-5.

Willis, J. H. (1971). ‘A handbook to plants in Victoria. Volume 1. Ferns, Conifers and Monocotyledons’.
(Melbourne University Press: Carlton.)

Manuscript received 21 June 1988

FERDINAND MUELLER, GOVERNMENT BOTANIST: THE ROLE OF
WILLIAM HOOKER IN HIS APPOINTMENT

by

Helen M. Cohn*

ABSTRACT

Cohn, Helen M. Ferdinand Mueller, Government Botanist: the role of William
Hooker in his appointment. Muelleria 7(1): 99-102 (1989) — Contemporary
commentary makes no mention of William Hooker being involved in Mueller’s
appointment as Government Botanist in Victoria. The only obituary that makes this
claim is unreliable. Later writers olfer no support for their contention that Hooker
recommended Mueller for the post. Circumstances of Mueller’s training, early
ernigration to Australia and botanical researches in Australia before settling in
Victoria suggest that the possibilities for him to have established any connection with
Hooker were too limited for Hooker to be in a position to recommend him. The
Colonial Office disclaimed official knowledge of Mueller’s appointment, an unlikely
occurrence had Hooker been involved. Statements made by Mueller and Hooker
themselves confirm that Hooker had no role in Mueller’s appointment.

INTRODUCTION

It is generally believed that Ferdinand Mueller’s appointment by Charles La
Trobe to the position of Government Botanist to the Colony of Victoria in 1 853 was at
the recommendation of Sir William Hooker, then Director of the Royal Botanic
Gardens, Kew . This would not have been unusual or unexpected given Hooker’s
position of influence with regard to the placement of suitably qualified people in
colonial botanical posts. However, an examination of contemporary and later writings
reveals little ground to support this contention. Similarly, the circumstances of
Mueller’s early career suggest that it was most unlikely that Hooker made any such
recommendation.

BIOGRAPHICAL SOURCES

There are very few contemporary biographical notes about Mueller which shed
any light on this question. Mueller was extremely reticent about himself, his letters and
publications being confined almost exclusively to his work. It was left to other people
to make public the details of his career. Among the earliest general biographical notes
is that written by Joseph Knapp (1877). In it he states: ‘In demselben Jahre [1852]
folgte er dem Rufe des Gouvemeurs C. Latrobe als Regierungs-Botaniker der Colonie
Victoria in den Staatdienst . . .’. No mention is made here of any intercession on the
part of William Hooker.

On Mueller’s death in 1896 a large number of obituaries and memorial notices
was published. These also are remarkably silent on the circumstances of Mueller’s
appointment. Baldwin Spencer, Professor of Biology at the University of Melbourne
and like Mueller one of the leading figures in the still small world of Victorian science,
wrote from personal acquaintance with Mueller (Spencer 1 896). Spencer makes no
mention of William Hooker but gives La T robe all the credit: ‘Evidently his reputation
as a botanist had preceded him, for in the same year Governor Latrobe appointed him
Government Botanist . . .’. Other memorialists, like Spencer, credit Mueller’s
appointment solely to La Trobe (Battye 1897; Warburg 1897), or make no mention of
the underlying reasons (McOwan 1896; Anon. 1896).

•National Herbarium of Victora, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

99

100

The only obituary that attributes to William Hooker any role in Mueller’s
appointment is that written by William’s son Joseph, his successor as Director at Kew
(Hooker 1897). In it he says:

Among his [Mueller’s] English correspondents was Sir W. Hooker, who interested
himself in his favour with Mr Goulburn, then on the point of leaving England as
Lieutenant-Governor of Victoria, and who was desirous of having the vegetable
resources of that colony turned to the best account. This resulted in the creation of
a Department of Botany in the Public Service of Victoria, and the appointment of
Mueller to its directorship.

No ‘Mr Goulburn’ occupied any senior position in the Victorian Civil Service at
that time. It seems probable that Joseph Hooker was referring to Frederick Goulburn
who came to the colonies as Colonial Secretary (not Lieutenant-Governor) to New
South Wales (not Victoria) and died in 1837. There are other errors of fact in the
obituary: Mueller is said to have arrived in Victoria from South Australia in 1 848 (the
actual date was 1852) expressly to visit the Victorian alps, and to have been one of the
three founders of the Royal Society of Victoria. The Society came into being from the
amalgamation of two earlier societies having between them a total of 21 founding
members. In 1 897 when he wrote this tribute to Mueller, Joseph Hooker was 80 years
old, trusting to his memory to recall events of 45 years previously. Joseph Hooker
cannot be considered a reliable witness to his father’s role in Mueller’s
appointment.

Later biographers unequivocally state that William Hooker was involved in
Mueller’s appointment. Mueller was variously said to have been ‘well accredited by
the eminent botanist. Sir William Hooker’ (Daley 1924: 26) and ‘recommended’ by
him (Hardy 1945), and La Trobe to have been ‘persuaded’ by Hooker to appoint
Mueller (Roach 1921). The two major biographies of Mueller claim that Hooker had
‘spoken highly of the merits of Dr Ferdinand Mueller’ (M. Willis 1949: 19), and that
approval of the appointment lay with Hooker (Kynaston 1981: 80). Other writers have
simply followed suit, for example J. Willis ( 1 97 1 ) and Powell ( 1 982). However not one
of these writers offers any supporting evidence.

MUELLER IN EUROPE

It is highly unlikely that William Hooker knew very much about Mueller at the
time of his appointment. Mueller left Germany at the age of 22, having just qualified
and having had little if any time to establish a reputation for himself outside his own
country. His outward journey to South Australia on the ‘Herrmann von Beckerath’
seems to have offered him no opportunity to meet either of the Hookers or any of the
other leading botanical personalities in England, so that he was virtually unknown to
the British botanical establishment. For the first five years of his time in Australia
before settling in Victoria, the specimens and papers he sent back to Europe were to his
German colleagues rather to anyone in England. This is reflected in the pattern of his
early publications.

By the end of 1853 Mueller’s European publications comprised eight papers in
German scientific journals and only two in England (Churchill, Muir & Sinkora 1978
& 1984). Taking into account the length of time for mail to reach Europe and the
vagaries of the editorial and publication process, publications in Europe or England in
1853 represent work done some time before that. In the absence of any scientific
publishing outlet in the Australian colonies, Mueller had naturally turned to the
contacts he had at home to assist with the publication of his scientific researches.

The two papers published in England were actually translations of papers
published in Germany in the same year (Mueller 1853c & 1853d). They appeared in
William Hooker’s own Journal of Botany and Kew Garden Miscellany (Mueller 1 853a
& 1 853b). However, they were not sent to Hooker. Rather they were sent to Richard
Kippist, then Librarian to the Linnean Society of London, who translated them from
German and read them to meetings of the Society on the 7th and 2 1 st December 1852
respectively. Hooker had an unhappy history of publishing a succession of journals
which struggled and finally collapsed under the weight of a small buying public.

101

dwindling copy and heavy production costs (Brock 1980). It is not hard to see why he
would have seized the opportunity to print these two short papers in the fourth and last
of his journals. What is clear from this is that Mueller at that point had no thought of
relying on Hooker as he did on his German contacts.

Even had William Hooker been well acquainted with Mueller it is unlikely he
would have recommended him ahead of a British botanist. Throughout his career at
Kew Hooker was instrumental in placing British botanists in key botanical posts, all of
which were official, all over the world. In this he took his ‘right’ of patronage very
seriously. There is the celebrated case of Charles Moore, who was appointed to the
Botanic Gardens in Sydney ahead of Hooker’s own nominee and to Hooker’s
considerable disgust (Gilbert 1986). His influence in colonial botanical matters was
well known. G. W. Francis at the Adelaide Botanic Gardens appealed to Hooker for
support should rival elaims be made for his job as Director there (Best 1986).

COLONIAL APPOINTMENT

Finally, had William Hooker recommended or approved Mueller’s appointment
the neeessary correspondence would have passed through the Colonial Office in
London. Minuted on a letter from Governor Hotham to the Colonial Office dated 3 1 st
January 1855 is the following revealing comment:

I do not recall that Dr Mueller’s name ever came officially before this Department
before. He is here styled Government Botanist, and he certainly was appointed in
the Colony. I recollect, however, Mr La Trobe mentioning Dr Mueller very
favorably in a private letter. He would be able to furnish any information
regarding Dr Mueller. (CO 309 v.31 204)

The most positive indication that William Hooker played no part in Mueller’s
appointment eomes from Mueller himself. He introdueed himself to Hooker in a letter
dated 3rd February 1853, a mere seven days after being appointed Government
Botanist. He had already embarked on the first of his exploring journeys through
Victoria.

As a highly esteemed promoter of botanical science throughout the world, you
will, I trust. Sir William, not without some interest receive the intelligence, that
his Excell, our scientific Governor Latrobe has been pleased to entrust to me the
newly created office of a government botanist for this province, an appointment
that I joyfully accept, as it now enables me at length to devote my time henceforth
exelusively to the study of the indigenous plants ... Of my botanical labours in
South Australia, I suppose little came to your notice . . . (Kew Correspondence
V.74, 135)

Hooker replied that he was ‘most agreeably surprised’ to have received the
‘welcome intelligence’ of his engagement by the government. ‘This is exactly as it
should be and I shall write to the Governor by this day’s post to thank him for his
service thus rendered to our favourite science’ (Argus 31.10.1853 p. 5).

ACKNOWLEDGEMENTS

The letter from Mueller to William Hooker is quoted with permission from the
Royal Botanic Gardens, Kew.

REFERENCES

Colonial Office series 309: inwards correspondence to the Secretary of State from Victoria.

Royal Botanic Gardens, Kew. Correspondence vol. 74: Australian letters 1851-58.

Anon. (1896). Baron von Mueller. Gard. Chron. 3s. 20: 464-466.

The Argus, Melbourne.

Battye, J. S. (1897). Memoirs of the late Baron von Mueller. J. Proc. Mueller Bot. Soc. W. Austral. 1(5):
5-12.

Best, B. J. (1986). George William Francis, first Director of the Adelaide Botanic Garden. (B. J. Best:
Adelaide.)

Brock, W. H. (1980). The development of commercial science journals in Victorian Britain. In Meadows,
A. J. (ed.). Development of science publishing in Europe. (Elsevier: Amsterdam.) pp. 95-122.

102

Churchill, D. M., Muir, T. B. & Sinkora, D. M. (1978). The published works of Ferdinand J. H. Mueller
(1825-1896). A/ue//eria 4: 1-120.

Churchill, D. M., Muir, T. B. & Sinkora, D. M. (1984). The published works of Ferdinand J. H. Mueller
(1825-1896). Supplement Muelleria 5: 229-248.

Daley, C. (1924). Baron Sir Ferdinand von Mueller, K.C.M.G., M.D., F.R.S., botanist, explorer and
geographer. Victorian Hist. Mag. 10(1); 23-75.

Gilbert, L. A. (1986). The Royal Botanic Gardens Sydney. (Oxford University Press: Sydney.)

Flardy, A. D. (1945X Excerpts from letters of Baron Sir Ferdinand von Mueller. Victorian Hist. Mag. 21(1):
i-23.

Hooker, J. D. (1897). [Obituary]. Proc. Roy. Soc. London 63:xxxii-xxxvi.

Knapp, J. A. (1877). Baron Ferdinand von Mueller: eine biographische Skizze. Z. Allg. Oesterr. Apotheker
Vereines 15(36): 597-617.

Kynaston, E. (1981). ^ man on edge: a life of Baron Sir Ferdinand von Mueller. (Allen Lane: Ringwood).

MacOwan, P. ( 1 896). The late Baron Sir Ferdinand von Mueller, C.M.G., F.R.S., etc. Agric. J. Cape of Good
Hope 9(24); 627-628.

Mueller, F. (1853a). The flora of South Australia, displayed in its fundamental features, and comparatively.
Hook. J. Bot. Kew Gard. Misc. 5; 65-72.

Mueller, F. ( 1853b). The vegetation of the districts surrounding Lake Torrens. Hook. J. Bot. Kew Gard. Misc.
5: 105-109.

Mueller, F. (1853c). Siidaustraliens Flora, in ihren Grundziigen und vergleichend dargestellt. Hamburger
Garten-Blumenzeitung 9: 267-273.

Mueller, F. (1853d). Die Vegetation der Gegenden urn den Torrens-See. Hamburger Garten-Blumenzeitung
9: 340-343.

Powell, J. M. (1982). National identity and the gifted immigrant Baron von Mueller, scientist and public
servant, 1825-96. Journal of Intercultural Studies 3(1): 56-61.

Roach, B. S. U921). Ferdinand von Mueller. South Australian Naturalist 2; 76-82.

Spencer, W. B. (1896). Baron von Mueller. Victorian Naturalist 13: 87-92.

Warburg, O. (1897). Ferdinand von Mueller. Ber. Deutsch. Bot. Ges. 15; 56-70.

Willis, J. H. (1971). The genesis of botanical science in Victoria, 1851-1875. Victorian Hist. Mag. 42(2):
699-703.

Willis, M. ( 1 949). By their fruits: a life of Ferdinand von Mueller, botanist and explorer. (Angus & Robertson:
Sydney.)

Revised manuscript received 4 July 1988

NEW GENERA AND SPECIES OF AUSTRALIAN INULEAE (ASTERACEAE)

by

P. S. Short*

ABSTRACT

Short, P. S. New genera and species of Australian Inuleae (Asteraceae). Muelleria 7( 1 ):
103-116 (1989). — Five new genera, Dielitzia Short, Feldstonia Short, Fitzwillia
Short, Lemooria Short and Sondottia Short are described. All but Sondottia are
monotypic. A new species of Dithyrostegia A. Gray is described. New species and new
combinations are; Dielitzia tysonii Short, Dithyrostegia gracilis Short, Feldstonia
nitens Short, Fitzwillia axilliflora (W.V. Fitzg. ex Ewart & J. White) Short, Lemooria
burkittii (Benth.) Short, Sondottia connata (W.V. Fitzg.) Short and S. glabrata
Short.

INTRODUCTION

In a revision of Angianthus Wendl. (sensu Bentham 1 867) I (Short 1 983) excluded
three species from Angianthus s. str., viz. A. axilliflorus W.V. Fitzg. ex Ewart & J.
White, A. connatus W.V. Fitzg. and A. burkittii (Benth.) J.M. Black. At the time 1
suggested that they may represent monotypic genera or have affinities with other
genera which had not been examined. Subsequent investigations have failed to reveal
such affinities. Thus, in this paper, each is formally referred to a new genus. Several
further taxa, of which I have known for some years, are also described.

Collections from all major Australian herbaria have been examined.

TAXONOMY

Dielitzia Short, gen. nov.

Herba annua, caespitosa, glomerulis foliis circumcinctis; glomeruli sessiles vel in axibus brevibus,
tomentosis majores. Folia sessilia, integra, infima opposita, superne alterna, sublinearia, tomentosa.
Glomeruli late ellipsoidei usque oblati; bracteae glomerulos subtendentes involucrum conspicuum
glomerulis aequilongum facientes, cartilagineae; receptaculum cupulatum, nudum. Capitula c. 4-15.
Bracteae intra capitulum biseritatae; bracteae exteriores 1-4, setaceae, longo-plumosae; bracteae
interiores (2)3-4, hyalinae, marmoratae, apicibus pilis rectis subrigidis praeditis. Flosculi 1 in quoque
capitulo, tubulare, hermaphroditi. Corolla 4 vel 5-lobata. Styli rami truncati, apicibus papillatis.
Stamina 4 vel 5; antherae ad basim caudatae, ad apicem appendicibus sterilibus. Cypselae
subobovoidae, papillatae; carpopodium absens. Pappus setaceus.

Typus: D. tysonii Short

Annual herb, tufted, of 1-20 compound heads surrounded by leaves, the
compound heads sessile or terminating short, tomentose major axes. Leaves sessile,
entire, ± linear, at least the lowermost opposite, the upper alternate, tomentose.
Compound heads broadly ellipsoid to obloid; bracts subtending the compound heads
forming a conspicuous involucre the length of the head, the bracts mainly
cartilaginous. General receptacle cup-like, naked. Capitula c. 4-15 per compound
head. Capitular bracts in 2 rows; outer bracts 1-4, bristle-like, long-plumose; inner
bracts (2)3-4, ± hyaline, with brown or blackish marbling, apices with straight, ± rigid
hairs f- ^ the total length of the bracts. Florets 1 per capitulum, bisexual. Corolla 4 or 5
lobed. Style branches truncate; apices papillate. Stamens 4 or 5; anthers caudate, each
with a sterile apical appendage; filament collar straight in outline and not thicker than
the filament. Cypselas ± obovoid, minutely papillate; carpopodium absent. Pappus
setaceus.

‘National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

103

104

Distribution (Fig. 1):

Monotypic. Restricted to inland Western Australia between latitudes c. 24° S.
and 29° S. and longitudes c. 115°E. and 123° E.

Etymology:

The generic name is an anagram derived from the surnames and commemorating
botanists F. L. E. Diels (1874-1945) and E. G. Pritzel (1875-1946).

Fig. 1. Distribution of Dielitzia tysonii, Feldstonia nitens, Dithyrostegia gracilis, Fitzwillia axilliflora,
Sondottia connatCy S. giabrata and Lemooria burkittii.

105

Notes:

With the exception of Isoetopsis the habit alone readily distinguishes this taxon
from all other Australian Asteraceae. The cartilaginous bracts of the general involucre,
the cup-like general receptacle and the structure of the capitular bracts (particularly
the marbled, inner bracts) are features unique to the genus.

Although the general receptacle is described as being glabrous the bristle-like
bracts deemed to be the outer capitular bracts could possibly be interpreted as
receptacular bracts.

The marbling of the inner bracts is best observed in spirit collections. In
herbarium specimens bracts may appear to be more or less uniform in colour.

Dielitzia tysonii Short, sp. nov.

Herba annua, caespitosa, glomerulis 1-20, ab foliis circumcinctis, sessilibus vel in axibus majoribus
usque ad 1-5 cm longibus. Folia sublinearia, c. 1-8 cm longa, 0 08-0T5 cm lata, ad basim dilatatae,
submucronatae, semisucculentae, tomentosae. Glomeruli late ellipsoidei usque obloidei, 4-6 mm
longi, 3-7 mm diametro; bracteae glomerulos subtendentes c. 8-12, praecipue cartilagineae sed
apicibus hyalinis. Capitula c. 4-15. Bracteae intra capitulum longitudine c. y-|- flosculi aequanti.
Flosculi 1 in quoque capitulo. Stamina 4 vel 5; antherae 0-53-0-64 mm longae; microsporangia
0-35-0-48 mm longa; appendices apicales 0'35-0-48 mm longae; pollinis grana in quaque anthera
28-88. Cypselae subobovoidea, 1-1-3 mm longae, 0-5-0-6 mm diametro. Setae pappi c. 10,
barbellatae, ad bases conjunctae, longitudine c. j-y corollae aequanti. (Fig. 2).

Holotypus: Western Australia, 17-2 km NE. of Nallan on Yarrabubba road,
23.viii.1986, Lander 1389, Fuhrer & Short {ULL 1556923). Isotypi: AD, CANB, K,
PERTH, S.

Annual herb, tufted, of 1-20 compound heads surrounded by leaves, the
compound heads ± sessile or on major axes to 1 -5 cm long. Leaves ± linear, c. 1 -8 cm
long, 0-08-0- 1 5 cm wide, expanded at the base, ± mucronate, semisucculent,
tomentose. Compound heads broadly ellipsoid to obloid, 4-6 mm long, 3-7 mm
diam.; bracts subtending compound heads c. 8-12, mainly cartilaginous but with
hyaline apices. Capitula c. 4-15 per compound head. Capitularhracts c. f- 1 the length
of the florets. Florets 1 per capitulum. Stamens 4 or 5; anthers 0-53-0-64 mm long;
microsporangia 0-35-0-48 mm long; apical appendages 0-1 6-0-25 mm long; pollen
grains 28-88 per anther. Cypselas ±obovoid, 1-1-3 mm long, 0-5-0-6 mm diam.
Pappus bristles c. 1 0, barbellate, of unequal length, fused at the base, c. }- j the length
of the corolla.

Distribution (Fig. 1):

See generic treatment.

Ecology & Reproductive Biology;

Habitat notes suggest a preference for sandy loam to clay soil. Collectors’ notes
include: ‘Growing in open Acacia shrubland. Brown sandy loam with gravel. With an
array of ephemeral composites including Gnephosis burkittii, Podolepis, Cephalip-
terum drummondii, Isoetopsis graminifolia, Brachyscome & Calotis', ‘Open
Acacia/Cassia scrub. Sandy loam covered with ironstone gravel’ and ‘Growing on
saline clay flat’.

Pollen: ovule ratios were determined for 15 plants from Short 1519. The values
obtained (range = 204-408; x = 329-7; S.D. = 56-7; S.E.x= 14-64) indicate a high
degree of self-pollination (Short 1981).

Etymology:

The specific epithet commemorates Isaac Tyson, a pastoralist who collected the
plant in 1893.

Notes:

Dielitzia tysonii has a marked resemblance in habit to Isoetopsis graminifolia
Turcz. It is readily distinguished in the field from the latter by its hairy, not glabrous,
leaves.

106

H. O

M M >N U H( KM \Htf <,f
^ !* HMM V {\fj ! >. VI -.IK \M V

Fig. 2. Holotype sheet of Dielitzia tysonii.

Selected Specimens Examined (Total 10):

Western Australia — Kennedy Range, 2I.viii.l986, Lander 1368 (MEL, PERTH); c. 39km from
Leonora along road to Laverton, 20. viii. 1982, Short 23/9 (MEL); Mt Gould, 22. viii. 1 986, Short 2552 (MEL,
PERTH); 16 km E. ofYalgoo, 15.ix.l973, Wilson 4/‘/9 (PERTH).

Dithyrostegia A. Gray

For a description of this now ditypic genus see Short (1983, p. 201).

Dithyrostegia gracilis Short, sp. nov.

Herba annua. Axes majores erecti, glabri, ramificatione dichotoma. Folia sessilia, sublinearia vel
lanceolata, 2-9 mm longa, 0-4-2 mm lata, amplexicaules; paginae exteriores glabrae, interiores
sparsim gjandulosae. Gomeruli 3-5-4-S mm longi, 2-5-3-5 mm diametro; bracteae glomerulos

107

subtendentes 2, in dimidio inferiore connatae, glabra. Capitula 6-20. Bracteae intra capitulum 1,
hyalinae, integrae. Florsculi 1 in quoque capitulo. Corolla 5-lobata. Stamina 5; antherae 1-3-1-5 mm
longae; microsporangia 0-96-1-1 mm longa; appendices apicales 0-34-0-38 longae. Cypselae
subobovoideae, c. 1-3-1 -5 mm longae, c. 0-6 mm diametro; carpopodium absens. Setae pappi laeves,
ad basim conjunctae, longitudine c. j corollae tubi aequanti. (Fig. 3).

Holotypus: Western Australia, Yuin Station, 2.ix.l975, Evans s.n. (PERTH).

Fig. 3. Holotype sheet of Dithyrostegia gracilis.

108

Annual herb, to c. 25 cm high. Major axes erect, glabrous; branching
dichotomous. Leaves sessile, ± linear or lanceolate, 2-9 mm long, 04-2 mm wide,
stem-clasping, outer surfaces glabrous, inner surfaces sparsely glandular. Compound
heads 3-5-4-5 mm long, 2-5-3-5 mm diam.; bracts subtending compound heads 2,
connate in c. the lower j, glabrous; general receptacle beset with long hairs. Capitula
6-20 per compound head. Capitular bracts 1, hyaline, enveloping the fruit and the
lower j of the corolla tube, entire. Florets 1 per capitulum; corolla 5-lobed, corolla tube
c. 1-5-2 mm long. Style branches truncate, papillate. Stamens 5; anthers 1 -3-1 -5 mm
long, caudate; microsporangia 0-96-1 -1 mm long; apical appendages 0-34-0-38 mm
long. Cypselas ±obovoid, c. 1-3-1-5 mm long, c. 0-6 mm diam., densely silky hairy,
carpopodium absent. Pappus of smooth bristles fused at the base, c. the length of the
corolla tube.

Distribution (Fig. 1):

Only known from the type collection from south-west Western Australia.

Ecology & Reproductive Biology:

No habitat notes accompany the type specimen.

Anther size and a pollen: ovule ratio of 5,464 determined for a single floret
indicate that the species commonly cross-pollinates.

Etymology:

The specific epithet refers to the slender habit.

Notes:

In a previous treatment of Dithyrostegia (Short 1983) this species was not
described pending further collections. Despite several searches (in 1 982, ’83 & ’86) the
species has not been recollected. That it is specifically distinct from D. amplexicaulis
seems indisputable. The leaves and compound heads are much smaller than in the
latter species, the capitular bracts are entire (not with long hairs at the apex) and the
general receptacle is sparsely hairy (not woolly).

Feldstonia Short, gen. nov.

Herba annua. Axes majores decumbentes usque ad ascendentes, rare erecti, glabri vel sparsim pilosi.
Folia sessilia, integra, infima opposita, supera alterna, linearia vel oblanceolata, glabra vel sparsim
pilosa. Glomeruli late obovoidei usque depresse obovoidei vel depresse ovoidei; involucrum generale
multiseriale, conspicuum; bracteae exteriores praecipue virides, opacae, subcartilagineae, glabrae,
nitentes, marginibus hyalinis; bracteae interiores praecipue hyalinae, glabrae, costis opacis;
receptaculum subconicum, glabrum. Capitula 15-50; bracteae capitula subtendentes (0)1(2),
subrigidae, praecipue, hyalinae glabrae, costis opacis, interdum parte distali leviter constricta, lutea.
Bracteae intra capitulum 4-5, in duo serialibus, praecipue hyalinae, planae usque concavae, costa
opaca, apice glabera vel sparsim pilosa. Flosculi ( 1 )2 in quoque capitulo, tubulare, hermaphroditi, lutei.
Corolla 5-lobata. Styli rami truncati, ad apicem papillati. Stamina 5; antherae ad basem caudatae, ad
apicem appendicibus sterilibus. Cypselae subobovoideae, pubescentes; carpopodium absens. Pappus
setaceus, setas subplumosas usque plumosas, ad basem conjunctae ferens.

Typus: F. nitens Short

Annual herb. Major axes decumbent to ascending, rarely erect, glabrous or
sparsely hairy. Leaves sessile, entire, the lowermost opposite, the upper alternate,
linear or oblanceolate, glabrous or sparsely hairy. Compound heads broadly obovoid to
depressed obovoid or depressed ovoid; general involucre multiseriate, conspicuous;
outer bracts mainly green, opaque, semi-cartilaginous, glabrous, shiny, with hyaline
margins; inner bracts mainly hyaline, glabrous, with opaque midribs; receptacle
± conical, glabrous. Capitula 15-50 per compound head; capitulum-subtending
bracts (0) 1(2), ± rigid, mainly hyaline, glabrous, midrib opaque, with the upper part of
the bract sometimes slightly constricted, yellow. Capitular bracts 4-5, in 2 rows,
mainly hyaline, flat to concave, midrib opaque and glabrous or with sparsely hairy
apices. Florets (1)2, tubular, bisexual, yellow; corolla 5-lobed. Style branches truncate,
apices papillate. Stamens 5; anthers caudate, each with a sterile apical appendage;

109

filament collar straight in outline and not thicker than the filament. Cypselas
± obovoid, pubescent, carpopodium absent. Pappus setaceus, the bristles subplumose
to plumose, united at the base.

Distribution (Fig. 1 ):

Monotypic. Restricted to Western Australia between latitudes c. 28° S. and 30° S.
and longitudes c. 116° E. and 1 1 8° E.

Etymology:

The name Feldstonia is an anagram derived from the surname and com-
memorates Danish botanist C. E. H. Ostenfeld (1873-1931) who published several
papers on Western Australian botany. Although an anagram it is regarded as a
personal generic name and following Recommendation 20A of the ICBN is given the
feminine gender.

Notes:

A number of attributes, including the semicartilaginous bracts of the general
involucre and the combination of features of the fruit, pappus and capitular bracts
readily distinguish the genus from others of the Inuleae.

Feldstonia nitens Short, sp. nov.

Herba annua. Axes majores 6-30 cm longi. Folia linearia vel suboblanceolata, interdum suprema
ovata, 3-40 mm longa, 0- 5-2-5 mm lata, submucronata, glabra vel sparsim pilosa. Glomeruli late
obovoidei usque depresse obovoidei vel depresse ovoidei, 5-8 mm longi, 4- 1 4 mm diametro. Capitula
15-50; bracteae capitula subtendentes ovatae usque lanceolatae, 3-4-4-8 mm longae, 1-2-1 -5 mm
latae. Bracteae intra capitulum 3-8-4-1 mm longae, l-l-l-4mm latae. Flosculi (1)2. Stamina 5;
antherae 1-6-1-9 mm longae, microsporangia 1-2-1-5 mm longa, appendices apicales 0-38-0-45 mm
longae. subobovoideae, 1-4- 1-9 mm longae, 0-6-0-9 mm diametro. Pappi setae c. 10-15, sub-

plumosae usque plumosae, ad basem conjunctae, longitudine r. j corollae tubi aequanti. (Fig. 4).

Holotypus: Western Australia, 1 9-3 km from Yalgoo along the road to Paynes Find.
Acacia scrub. Growing with an array of annuals including Cephalipterum
drummondii, Pogonolepis, Gnephosis and Myriocephalus, 2.ix. 1 982, Short 1615 (MEL
621021). IsoTYPi: AD, BRI, CANB, E, GH, HO, K, NSW, PERTH, S.

Annual herb. Major axes 6-30 cm long. Leaves linear or ± oblanceolate, the
uppermost sometimes ovate, 3-40 mm long, 0-5-2-5 mm wide, mucronate, glabrous
or sparsely hairy. Compound heads broadly obovoid to depressed obovoid or
depressed ovoid, 5-8 mm long, 4-14 mm diam. Capitula 15-50 per compound head;
capitulum-subtending bracts ovate to lanceolate, 3-4-4-8 mm long, 1 -2-1 -5 mm wide.
Capitular bracts 3-8-4-1 mm long, l-l-l-4mm wide. Florets (1)2; corolla tube
2-2-7 mm long. Stamens 5; anthers 1-6-1 -9 mm long; microsporangia 1-2-1 -5 mm
long; apical appendages 0-38-0-45 mm long. Cypselas ± obovoid, 1-4-1 -9 mm long,
0-6-0-9 mm diam. bristles c. 10-15, subplumose to plumose, bases united, c. j

the length of the corolla tube.

Distribution (Fig. 1):

See generic treatment.

Ecology & Reproductive Biology:

Collectors’ habitat notes include: ‘Acacia scrub. Red-brown loam’ and ‘Mallee
scrub {Acacia common). Reddish sandy loam’.

A pollen: ovule ratio of 4,460, determined from a single floret, indicates that the
species commonly cross-pollinates.

Etymology:

The specific epithet refers to the shiny bracts of the general involucre.

Selected Specimens Examined (Total 14):

Western Australia — 9 miles N. of Paynes Find, 23.X.1973, Demarz 4674 (KP, PERTH); 20 km from

no

Hoiorfrf. ^

7-S.SUl-

Dtttiminavit -c^

N*':i/.- i o:' '■

\\llOWI. HF.RIURIl M Of
*K rORII, (MM), \rMR\UA

Fig. 4. Holotype sheet of Feldstonia nitens.

pPf#|l'T«llll''|'«l«N'|i«!!in'jnil|ll»fill|lll'pi|il»(iM||inifili||l»|il«||in|i|||||iii|iii||||iij,^|||i,|,^

Ill

Yalgoo on Paynes Find road, 21.X.1983, Short 2155 (MEL); 86 km NE. of Wubin, 29.ix.1986, Wilson 12350
(MEL, PERTH).

Fitzwillia Short, gen. nov.

Herba annua. Axes majores ascendentes usque erecti, glabri, alis hyalinis; caulis simplex vel e nodis
basalibus superioribusque ramificans. Folia opposita, sessilia, Integra, semisucculenta, glabra,
mucronata. Glomeruli perdepresse obovoidei usque obovoidei; involucrum generate absens;
receptaculum subintegrum. Capitula 5-10; bracteae intra capitulum 4(6), integrae, hyalinae, planae
usque conduplicatae. Flosculi 1 vel 2 in quoque capitulo, tubulares, hermaphroditi, albi. Corolla
5-lobata. Styli rami truncati, ad apicem papillati. Stamina 5; antherae ad basem caudatae, ad apicem
appendicibus sterilibus. Cypselae subobconicae, villosae; pericarpium ad apicem sclerenchymatum,
fascibus vascularibus 2; testa sine fascibus vascularibus; carpopodium absens. Pappus subcyathiformis,
ciliatus.

Typus: F. axilliflora

Annual herb. Major axes ascending to erect, glabrous, with hyaline wings; stem
simple or forming major branches at basal and upper nodes. Leaves sessile, entire,
opposite, erect, glabrous, semisucculent, mucronate. Compound heads broadly
depressed obovoid to obovoid; general involucre absent; receptacle ± entire. Capitula
5-10 per compound head; capitulum-subtending bracts leaf-like, glabrous. Capitular
bracts 4(6), entire, hyaline, flat to conduplicate. Florets 1 or 2 per capitulum, tubular,
bisexual, white; corolla 5-lobed. 5'ty/e branches truncate, apices papillate. Stamens 5;
anthers caudate, each with a sterile apical appendage; filament collar straight in
outline and not thicker than the filament. Cypselas ± obconic, villous; pericarp in
mid-transverse section lacking sclerenchyma but the fruit apex with a capping of
sclerenchyma, with two, medial/oblique vascular bundles; testa thin, lacking vascular
bundles in mid-transverse section; carpopodium absent. Pappus cup-like, ciliate.

Distribution (Fig. 1):

Monotypic. Restricted to south-west Western Australia. Collections have only
been gathered near Cowcowing, Newdegate and Morawa.

Etymology:

The name Fitzwillia is an anagram derived from the names and commemorating
the botanist William V. Fitzgerald (1867-1 929). Although an anagram it is regarded as
a personal generic name and following Recommendation 20A of the ICBN is given the
feminine gender.

Notes:

The fruit provide the most distinctive feature of this genus although a sclerified
capping also occurs in cypselas of Epitriche demissus (A. Gray) Short (Short 1989). The
leaf-like capitulum-subtending bracts and the arrangement of the capitular bracts are
characters unique to the genus. The pale white florets have not been observed in other
Australian inuloid species.

Fitzwillia axilliflora (W.V. Fitzg. ex Ewart & J. White) Short, comb. nov.

Basionym: Angianthus axilliflorus W.V. Fitzg. ex Ewart & J. White, Proc. Roy.
Soc. Viet. 22: 315, pi. 56, figs 1-3 (1910), ('axilijlorus')-, W.V. Fitzg., J. Bot. 50: 21
(1912); Grieve & Blackall, W. Aust. Wildfls 812 (1975); Short, Muelleria 5: 209
(1983). Lectotype {fide Short 1983): Cowcowing, Oct. 1904, Koch 1196 (MEL
541217). Isolectotypes and probable Isolectotypes: AD, BM (same no; but
dated Aug. 1904), MEL 541218, MEL 541219, NSW (2 sheets), PERTH.

Annual herbs, 3-13-5 cm high. Major axes ascending to erect, with 2-4 hyaline
wings; stem simple or forming shorter major axes at basal and upper nodes, all axes
glabrous. Leaver lanceolate or ± linear, c. 4-7 mm long, 0-7-1 -3 mm wide, ± concave,
semisucculent, mucronate, glabrous. Compound heads broadly depressed obovoid to
obovoid, 4-5-7 mm long, 2-5-8 mm diam.; general involucre absent but several
leaf-like bracts present. Capitula 5-10 per compound head; capitulum-subtending

112

bracts ±trullate to narrowly trullate or ovate to lanceolate, 4-4-5 mm long,
1 -3-2-2 mm wide. Capitular bracts 4(5,6), 3-T-4-3 mm long, c. the length of the floret.
Florets 1 or 2, pale white. Stamens 5; anthers 1-3-1 -6 mm long; microsporangia
1-1-2 mm long; apical appendages 0-33-0-44 mm long. Cypselas ±obconic,
1-1-1 -4 mm long, 0-6-0-78 mm diam., villous. Pappus eup-like, with ciliate margins,
c. 0-3-0-5 mm long.

Distribution (Fig. 1):

See generic treatment.

Ecology & Reproductive Biology:

The speeies is apparently confined to the margins of salt lakes where it grows
amongst samphire in sand or sometimes clay loam.

A pollen: ovule ratio of 7,398, determined from a single floret, indicates that the
species commonly cross-pollinates.

Specimens Examined:

Western Australia — Newdegate, 1931, Blackall 1276 (PERTH); 5 km S. of Morawa, 23.x. 1 983, Short
2188 (MEL); 5 km S. of Morawa, 16.ix.l986, Short 2959 (AD, CANB, MEL, PERTH).

Lemooria Short, gen. nov.

Herba annua. Axes majores prostrati, sparsim lanati, pilis glandiferis. Folia ad basem opposita,
superiora altema, sessilia, Integra, glabra vel pilis glandiferis. Glomeruli depresse ovoidei; involucrum
generate conspicuum; bracteae 12-18, marginibus hyalinis, dense lanatae; receptaculum ramosum.
Capitula c. 10-20, sine bracteis subtendentibus. Bracteae intra capitulum (4)5-6(8), in ± 2 serialibus,
ovatae usque lanceolatae vel ellipticae, planae usque concavae, praecipue hyalinae sed costa viridi,
lanatae, marginibus superibus laciniatis. Flosculi 1 vel 2, tubulare, hermaphroditi, lutei. Corolla
5-lobata. Styli rami truncati, ad apicem papillati. Stamina 5; antherae ad basem caudatae, ad apicem
appendicibus sterilibus. Cypse/aesubobovoideae, fuscae, sparsim papillatae, plerumque apicibus pilos
intertextos ferentibus; pericarpium sine sclerenchyma; carpopodium absens. Pappus setaceus; setae
8-12, subplumosae, ad basem conjunctae, longitudine c. y corollae tubi aequanti.

Typus: L. burkittii (Benth.) Short

Annual herb. Major axes prostrate, sparsely woolly, some glandular hairs present.
Leaves opposite at the base, the upper alternate, sessile, entire, glabrous or with
glandular hairs. Compound heads depressed ovoid; general involuere conspicuous,
about equal to or longer than the head, braets 12-18, midribs leaf-like and longer than
the wing-like hyaline margins, densely lanate; receptacle branched. Capitula c. 10-20;
capitulum-subtending braets absent. Capitular bracts (4)5-6(8), in ± 2 rows, ovate to
lanceolate or elliptic, flat to concave, mainly hyaline but with green midrib, lanate, the
upper margins laciniate. Florets 1 or 2, tubular, bisexual, yellow; corolla 5-lobed. Style
branches truncate, apices papillate. Stamens 5; anthers caudate and with sterile apical
appendages. Cypselas ± obovoid, brown, sparsely papillate, usually with apices with
intertwined hairs; pericarp lacking sclerenchyma, vascular bundles two; carpopodium
absent. Pappus setaceus; bristles 8-12, subplumose and united at the base, c. j the
length of the corolla tube.

Distribution (Fig. I ):

Monotypic. Widf spread in semi-arid and arid regions of the Australian mainland
south of c. 26°S.

Etymology:

The name Lemooria is an anagram commemorating the botanist Spencer Le
Marchant Moore (1850-1931). It is of the feminine gender (see note under
Fitzwillia).

Notes:

A unique combination of features, i.e. the capitular bracts and bracts of the
general involucre, the pappus and fruit, distinguishes Lemooria from other inuloid
genera. It is one of the most distinctive Australian composites and the species is readily

113

identified in the field by its reddish, prostrate, somewhat wiry branches terminating in
woolly compound heads.

Lemooria burkittii (Benth.) Short, comb. nov.

Basionym: Gnephosis burkittii Benth., FI. Austr. 3: 570 (1867); Short,
Muelleria 5: 210 (1983). — Angianthus burkittii (Benth.) J.M. Black, FI. S. Aust.
1st ed. 645, pi. 53 (1929), 2nd ed. 925, fig. 1227 (1957). Lectotype {fide Short
1983): Lake Gillies, s. dat., Burkitt s.n. (MEL 541211). Isolectotype: K.

Annual herb. Major axes prostrate, 1-5-10 cm long, reddish, sparsely woolly,
some glandular hairs present. Leaves linear, 0-5-2 cm long, c. 0-03-0-05 cm wide,
glabrous or with glandular hairs. Compound heads depressed ovoid, 0-6- 1 -2 cm diam.;
bracts subtending the heads c. 12-18. Capitula c. 10-20 per compound head.
Capitular bracts (4)5-6(8), ovate to lanceolate or elliptic, 2-8-3-4mm long,
0-6-1 -3 mm wide. Florets 1 or 2 per capitulum; corolla tube 1 -8-2-5 mm long.
Stamens 5; anthers 0-19-0-21 mm long; microsporangia 0-39-0-46 mm long; apical
appendages 0-1 9-0-21 mm long. Cypselas ±obovoid, 0-7-1-2 mm long, 0-5-0-85 mm
diam. Pappus of 8-12 subplumose bristles united at the base.

Distribution (Fig. 1):

See generic treatment.

Ecology & Reproductive Biology:

A common species in both low and tall shrubland formations. Collectors’ notes
include: ‘On desert loam, on bare eroded patches in community oiAtriplex vesicaria,
with some Bassia [Selerolaena] species’. ‘In open Acacia shrubland. Brown sandy loam
with gravel . . . with an array of ephemeral composites’, ‘Tall Acacia open shrubland.
Clay loam’ and ‘In red-brown loam in Dodonaea! Ptilotus shrubland’.

Pollen: ovule ratios determined for Short 756 and Short 1520 range from 446 to
696, values indicative of a high degree of self-pollination (Short 1981).

Selected Specimens Examined (Total c. 120):

Western Australia — 39 km from Leonora along Laverton road, 20.viii.l982, Short 1520 (MEL); E.
edge of Fraser Range, 19.ix.l982, Short 1761 (MEL).

South Australia — 5 km W. of Nonning Hmsd, 30.viii.l968, Copley 2035 (AD); Koonamore,
7.viii.l971, Crisp 283 (AD, CBG).

Victoria — Parish of Olney, 30.viii.l948, Willis s.n. (MEL 84577).

New South Wales — Terrananya Stn, 26.vii.1955, Constable s.n. (NSW 35571, NT 19057); Fowlers
Gap Stn, 30.viii.l973, Cunningham & Milthorpe 1349 (NSW).

Queensland — 65 km N. of Hungerford, s. dat., Hocking s.n. (BRI 192446).

Sondottia Short, gen. nov.

Herba annua. Axes majores aseendentes usque ereeti, glabri usque lanati; caulis simplex vel e nodis
basalibus superioribusque ramificans. Folia opposita, connata, sessilia, Integra, glabra vel lanata,
submucronata, interdum semisucculenta. Glomeruli late ellipsoidei usque obloidei; bracteae
glomerulos subtendentes earentes, vel 2 vel 4, marginibus hyalinis. Capitula 5-13; bracteae capitula
subtendentes virides, praecipue cartilagineae, apicibus hyalinis et interdum marginibus hyalinis,
glabrae usque dense lanatae. Bracteae intra capitulum 4-6, hyalinae costa opaca, subplanae, glabrae vel
sparsim lanatae. Flosculi 1 in quoque capitula, tubulate, hermaphroditi, lutei. Corolla 5-lobata. Styli
rami truncati, ad apicem papillati. Stamina 5; antherae ad basem caudatae, ad apicem appendicibus
sterilibus. Cypselae subobovoidae, praecipue glabrae sed apicibus longe pilosis, sine carpopodium.
Pappus subcyathiformis, laciniatus.

Lectotypus (here chosen): S. connata (W.V. Fitzg.) Short

Annual herbs. Major axes ascending to erect, glabrous to lanate; stem simple or
forming major branches at basal and upper nodes. Leaves opposite, connate, sessile,
entire, glabrous or lanate, ± mucronate, sometimes semisucculent. Compound heads
broadly ellipsoid to obloid; bracts subtending compound heads 0, 2 or 4, with hyaline
margins (sometimes several leaf-like bracts present); general receptacle a shortly
branched axis. Capitula 5-13 per compound head; capitulum-subtending bracts

114

green, mainly cartilaginous but with hyaline apices and sometimes with very narrow
hyaline margins, glabrous to densely lanate, each bract subtending 1-3 capitula.
Capitularhmcts 4-6, hyaline, with opaque midribs, ± flat, glabrous or sparsely lanate.
Florets 1 per capitulum, tubular, bisexual, yellow; corolla 5-lobed. Style branches
truncate, apices papillate. Stamens 5; anthers caudate, each with a sterile apical
appendage; filament collar straight in outline and not thicker than the filament.
Cypselas ± obovoid, mainly glabrous but with long, intertwined hairs at the apex;
carpopodium annular. Pappus cup-like, laciniate.

Distribution (Fig. 1):

A ditypic genus restricted to Western Australia.

Etymology:

The name Sondottia is of feminine gender and is an anagram derived from the
names and commemorating the botanist Otto Wilhelm Sonder (1812-1881).

Notes:

The cartilaginous capitulum-subtending bracts are apparently unique to this
genus and readily separate it from any other genera with single flowered capitula.
Other distinguishing features include the opposite, connate leaves and the intertwined
long hairs at the apex of the fruit.

Key to Species of Sondottia

1. General involucre absent or several leaf-like bracts at base of head; upper axes

lanate 1 • S. connata

1. General involucre of 2 or 4 bracts with broad, hyaline margins; upper axes
± glabrous 2. S. glabrata

Sondottia connata (W.V. Fitzg.) Short, comb. nov.

Basionym: Angianthus connatus W.V. Fitzg. J. West Aust. Nat. Hist. Soc. 2: 24
(1905); Grieve & Blackall, W. Aust. Wildfls 816 (1975); Short, Muelleria 5: 209
(1983). Lectotype (fide Short 1983); Mingenew, Sept. 1903, Fitzgerald s.n. (NSW
138682). IsoLECTOTYPEs: NSW 138683, PERTH.

Annual herb, c. 3-12 cm high. Mayor oxej ascending to erect, mainly glabrous but
the upper part lanate; stem simple or forming branches at basal and upper nodes.
Leaves linear, c. 5-13 mm long, 0-5-1 -4 mm wide, often semisucculent, mucronate,
usually glabrous but the uppermost leaves sometimes lanate. Compound heads
obovoid, c. 6-5-10 mm long, 3-5-5 mm diam.; general involucre absent but one or
several leaf-like, lanate bracts with small, hyaline apices may be present at the base of
the head. Capitula c. 5-13 per compound head; capitulum-subtending bracts
± elliptic to narrowly elliptic or obovate, 3-6-5-2 mm long, 0-55-1-9 mm wide,
mainly cartilaginous and green but with hyaline apices and, sometimes very narrow,
(<0-l mm) hyaline margins, sparsely to densely lanate, each bract subtending 1-3
capitula. Capitular bracts 5-6, narrowly elliptic or linear, 3-3-8 mm long, 0-3-0- 5 mm
wide, c. the length of the florets, usually mainly hyaline but sometimes the opaque
midrib more prominent, glabrous or sparsely lanate. Florets 1 per capitulum; corolla
tube 2-2-2-6mm long. Stamens 5; anthers 1-4-1 -5 mm long; microsporangia

0- 95-1 mm long; apical appendages 0-43-0-48 mm long. Cypselas ± obovoid,

1- 6-1-85 mm long, 0-5-0-7 mm diam. Pappus cup-like, laciniate, c. 0-2 mm long.

Distribution (Fig. 1):

Restrieted to Western Australia between latitudes c. 21° S. and 30° S. and 1 1 6° E.
and 121° E.

In the revision of Angianthus s. lat. (Short 1 983) it was erroneously recorded that
the species was only known from the type locality. It is moderately common.

115

Ecology & Reproductive Biology:

The species occurs in an array of arid habitats but seems to be most common in
saline environments. Collectors’ notes include: ‘Sandy loam with ironstone gravel,
with scattered Atriplex shrubs’, ‘In sand and very sandy loam amongst Acacia,
Eremophila shrubs’ and ‘Beneath Acacia shrubs amongst samphire. Loam.’

Pollen : ovule ratios have not been obtained for this species but from anther size it
is evident that several thousand pollen grains occur in each floret, suggesting that
cross-pollination commonly occurs (Short 1983).

Selected Specimens Examined (Total 15):

Western Australia — Hospital Rocks, 8.x. 1983, Short 1998 (AD, MEL, PERTH); 26 km S. of Cue,
14.ix.l986, Short 2921 (MEL, PERTH); 6 km S. of Warriedar, 26.ix.1986, Wilson 12293 (MEL, PERTH);
Lake Austin, c. 15 km S. of Cue, 28. ix. 1986, Wilson 12326 (MEL, PERTH).

Sondottia glabrata Short, sp. nov.

Herba annua, usque ad 40 cm alta. Axes majores erecti, glabrati. Folia praecipue linearia, usque ad
c. 10 mm longa et c. 1 mm lata, glabra, summa subovata usque lanceolata basibus hyalinis. Glomeruli
subobovoidei vel ellipsoidei, 6-7 mm longi, 3-4 mm diametro; bracteae glomerulos subtendentes 2 vel
4, subovatae vel ellipticae, 4-5-5 mm longae, 2-5-3-3 mm latae, subglabrae usque lanatae, marginibus
hyalinis 0-7-l-33mm latis. Capitula 4-8; bracteae capitula subtendentes anguste ellipticae vel
oblanceolatae, 3-7-4-3 mm longae, 0-7-1 -3 mm latae, sparsim usque dense lanatae. Bracteae intra
capitulum 4-5, anguste ellipticae vel lanceolatae, 2-8-3-7 mm longae, 0-2-0-5 mm latae, sparsim
lanatae. Flosculi 1 in quoque capitulo. Corollae tubus 2-4-2-7 mm longus. Stamina 5; antherae
l-6-l-9mm longae; microsporangia l-3-l-6mm longa; appendices apicales 0-3-0-34 mm longae.
Cypselae subobovoideae, 1-4- 1-6 mm longae, 0-45-0-6 mm diametro. Pappus subcyanthiformis,
laciniatus, 0-2-0-5 mm longus. (Fig. 5).

Holotypus: Western Australia, c. 6 km S. of Wooramel River along the north-west
coastal highway. ChtnopoA! Acacia shrubland. Compact sandy loam, 1 6.x. 1 983, Short
2088 (MEL 1523448). Isotypus: PERTH.

Annual herb, to c. 10 cm high. Major axes erect, ± glabrous; stem forming
branches at basal and upper nodes. Leaves mainly linear, to c. 10 mm long, c. 1 mm

SI n El 31 tt ai 6 8 r 9 s i> E 6 I ■

NATIONAL HEKB\KILM OK
VICTORIA (MEL). AISIRAL!\

Fig. 5. Holotype sheet of Sondottia glabrata.

116

wide, glabrous, the uppermost ± ovate to lanceolate and with hyaline bases.
Compound heads ±obovoid to ellipsoid, 6-7 mm long, 3-4 mm diam.; bracts
subtending compound heads 2 or 4, c. the length of the head, ± ovate or elliptic,
4-5-5 mm long, 2-5-3-6mm wide, ± glabrous to lanate, with hyaline margins
0-7-1 -33 mm wide. Capitula 4-8 per compound head; capitulum-subtending bracts
narrowly elliptic or oblanceolate, 3-7-4-3 mm long, 0-7-1 -3 mm wide, mainly
cartilaginous and green but with hyaline apices and sometimes with narrow
(<0-l mm) hyaline margins, sparsely to densely lanate, each bract subtending 1
capitulum. Capitular bracts 4-5, narrowly elliptic or lanceolate, 2-S-3-7 mm long,
0-2-0- 5 mm wide, to c. | the length of the florets, mainly hyaline but with an opaque
midrib, sparsely lanate. Florets 1 per capitulum; corolla tubes 2-4-2-7 mm long.
Stamens 5; anthers l-6-l-9mm long; microsporangia l-3-l-6mm long; apical
appendage 0-3-0-34 mm long. Cypselas ±obovoid, l-4-l-6mm long, 0-45-0-6 mm
diam. Pappus cup-like, laciniate, 0-2-0-5 mm long.

Distribution (Fig. 1);

Only known from the vicinity of the Wooramel River, Western Australia.
Ecology & Reproductive Biology:

Habitat notes indicate that the species occurs in chenopod (mainly
Atriplex)l Acacia shrubland.

As with S. connata this species, as indicated by anther size, probably has a
pollen: ovule ratio of several thousand. Therefore cross-pollination is likely to be
common (Short 1983).

Specimens Examined: (2, including type)

Western Australia — e. 28 km S. of Wooramel River along the north-west coastal highway, 16.x 1983
Short 2094 (MEL).

ACKNOWLEDGEMENTS

I thank Dr W. R. Barker for assisting with the Latin descriptions and my
colleagues at MEL for general assistance with the manuscript.

From 1982 to 1987 my work on the Australian Inuleae was partly funded by an
Australian Biological Resources Study Grant.

REFERENCES

Short, P. S. (1981). Pollen-ovule ratios, breeding systems and distribution patterns of some Australian
Gnaphaliinae (Compositae; Inuleae). Muelleria 4: 395-417.

Short, P. S. (1983). A revision of Angianthus Wendl. sensu lato (Compositae; Inuleae: Gnaphaliinae)
Muelleria 5: 143-214.

Short. P.^S. 0989). Notes on the fruit anatomy of Australian members of the Inuleae (Asteraceae). Muelleria

Manuscript received 20 July 1988

OLEARIA ARCHERI (ASTERACEAE: ASTEREAE), A NEW NAME FOR A
FAMILIAR SPECIES FROM TASMANIA

by

N. S. Lander*

ABSTRACT

Lander, N. S. Olearia archeri (Asteraceae: Astereae), a new name for a familiar species
from Tasmania. Muelleria 7(1): 117-121 (1989). — The neotypification by D. I.
Morris of Olearia persoonioides var. lanceolata Benth. is shown to be superseded by
the rediscovery of original material, here designated as the lectotype. A new species,
Olearia archeri, is described and illustrated.

INTRODUCTION

D. I. Morris (1977) raised Olearia persoonioides (DC.) Benth. var. lanceolata
Benth. to specific rank as O. lanceolata (Benth.) D. I. Morris. In the process of doing
this Morris designated a neotype for the variety stating that ‘it has not been possible to
locate Bentham’s type’.

Bentham (1867) cited no material in his protologue of O. persoonioides var.
lanceolata. However, examination of Olearia specimens held at major Australian,
British and Continental herbaria in the course of my ongoing revisionary studies in
this genus has brought to light a specimen at K, originally from the Herbarium
Hookeranum, annotated by J. D. Hooker as "E[urbyid\ persoonioides P lanceolata' and
subsequently annotated by Bentham as 'Olearia persoonioides Benth. var. lanceolata'
(Fig. 1). A small printed label reads ‘FLORA AUSTRALIENSIS, named by Mr
BENTHAM ; a handwritten note gives the relevant volume and page reference.
Although it bears no collector’s name, this specimen bears a typical R. C. Gunn label
with the number 1 142/1842 and was gathered at Mt Wellington on 1 March 1839.

The protologue of O. persoonioides var. lanceolata Benth. distinguishes it from
the type variety as follows: ‘Leaves lanceolate, almost acute. Flower heads fewer, but
scarcely larger. Achenes glabrous.’ The specimen described above matches this
protologue in all details and it is the only original material encountered that is so
annotated as a variety of O. persoonioides by Bentham. Thus its rediscovery
supersedes Morris’ neotypification: it is here designated as the Lectotype of Olearia
persoonioides var. lanceolata Benth. Duplicates (Isolectotypes) of this collection are
held at HO and NSW.

Eurybia persoonioides var. lanceolata J.D. Hook, is validly and legitimately
published (Hooker 1 847) and, as the above specimen is clearly cited by Hooker, it must
be accepted as the holotype. Neither this name or any reference to its place of
publication is given by Bentham (1867) in his treatment of O. persoonioides.

The holotype of Hooker’s Eurybia persoonioides var. lanceolata, and the lectotype
of Bentham’s Olearia persoonioides var. lanceolata, represents a narrow-leaved form
of O. persoonioides of trivial significance taxonomically. It bears little resemblance to
Morris’ neotype which belongs unequivocally to the familiar and distinctive species
hitherto known erroneously as O. lanceolata, which thus stands in need of a new
name.

Olearia archeri Lander, nom. nov.

Species O/ear/a persoonioidi et O. tasmaniae affinis a quibus bracteis involucralibus ferentibus piles
glanduliferos, capitulis pedunculo communi, basibus antherae breviter sagittatis, appendicibus
antherae ovatis, et stylo ferenti pilos T-formes difFert. (Fig. 2).

• Western Australian Herbarium, Department of Conservation & Land Management, P.O Box 104 Como
Western Australia, Australia 6152. ,

117

118

WOLOTYPE

OlVlMyv r« |>iAvt iMK’StHt .v - . .

C,Vl.R.O.. C^satftiu, At.MKAtu ' . . • ,t .

Nw;ai|vi IZ7Q

Dam /3 S3

Fig. 1. Holotype of Eurybia persoonioides var. lanceolata J.D. Hook, and lectotype of Otearia persoonioides
var. lanceolata Benth. Scale in cm.

119

Typus: Cash’s Lookout, Eaglehawk Neck, Tasmania, 10 November 1973, D. & A.
Ratkowsky s.n. (Holotypus: HO. Isotypi: K, Queen Victoria Museum, Launceston
— n.v.)

Missapplied names:

Olearia lanceolata auct. non (Benth.) D. I. Morris (1977) nec F. Muell. ex Sender
( 1 853); D. I. Morris, Rec. Queen Victoria Mus. 55: 1 (J 977). — Olearia persoonioides
var. lanceolata auct. non Benth.; L. Rodway, Tas. FI. 74 (1903); W. Curtis, Stud. FI.
Tasm. 2: 304 (1963).

Shrub to 2 m high. Vestiture of stems, petioles, leaves, peduncles and involucral
bracts pannose with multicellular, T-shaped, eglandular hairs. Stems erect, pinkish
brown to yellowish grey when young, becoming brownish grey, striate and fissured.
Leaves alternate and scattered basally, opposite and crowded apically, ascending,
petiolate; petiole 3-7 mm long; lamina flat, narrowly elliptic, 12-99x3-15 mm,
discolorous, pale yellowish brown abaxially, green adaxially; venation distinct,
reticulate; vestiture pannose abaxially, subglabrous adaxially with multicellular.

Fig. 2. Olearia archeri Lander. A — Habit. B — Ray floret. C — Disc floret. D — Anther tube. E — Stylar
arms with T-shaped hairs. F — T-shaped hairs. Drawn from the holotype.

120

uniseriate, capitate glandular hairs; texture coriaceous; base acute or rounded; margin
entire, flat; apex rounded, muticous. Heads pedunculate, in 3-headed corymbose
clusters forming large, terminal compound-corymbose conflorescenees on a common
peduncle, conspicuously radiate, 15-8-20-2 mm in diameter. Peduncle io 82 mm long
with several leaf-like bracts grading into those of the involucre. Involucre
campanulate; bracts 5-7-seriate, 2-7-8-6x0-8-l-6 mm. Outer involucral bracts
somewhat cymbiform, narrowly triangular; stereome green, basally pannose abaxially
with scattered multicellular, biseriate, capitate hairs, chartaceous; margin fimbriate;
apex rounded. Inner involucral bracts flat, narrowly triangular or elliptic; stereome
green, pannose abaxially and densely glandular basally, chartaceous; margin fimbrate;
apex acute. Receptacle weakly convex. Ray florets 7-9, uniseriate, white,
131-16-4mm long; tube weakly hairy centrally with T-shaped hairs; ligule elliptic,
8-8-1 T2x 3-5-4-2 mm, glabrous, acute, minutely 3-lobed apically; stylar arms
filiform, 1-5- 1-7 mm long. Disc florets 19-27, bisexual, yellow, infundibular,
8-6-9-6mm long, with scattered T-shaped hairs centrally to apically; lobes 5,
2-5-3-2 mm long, acute; anthers 2-5-2-9 mm long, briefly sagitate basally and shorter
than the filament eollar, with ovate sterile appendage; filament collar 0-5-0- 8 mm
long; stylar arms 2-2-2-4 mm long, with scattered T-shaped hairs below the point of
bifurcation, with half-conic sterile appendages bearing botuliform papillae above the
stigmatie lines. Achene narrowly ellipsoid, 3-3-4-7x0-6-10 mm, brown, densely
hairy apically with T-shaped hairs; venation indistinct; carpopodium central or
slightly oblique. Pappus biseriate, with 66-85 barbellate bristles subequal to the
tubular florets, with the barbs longer towards the apex.

Other Specimens Examined (Total number examined 15):

7aiman;a — Franklin Road, 43° OS'S., 147° 01 ' E., Oct. 1974, Af. ^//an 5, n. (HO); Maria Range, Maria
Island, 42° 37' S., 148° 06' E., M/ fiTOW/! 22-#(HO); NicholsCap, 41°45' S., 148° 16' E., May 1984, P Collier
s.n. (HO); Mt Andrew, P. Collier 974 (HO); Nichols Cap, W. of Seymour Beach, 41°45' S., 148° 16' E., Jan.
1979, 5. Harris s.n. (HO); McGregors Peak, W.D. Jackson s.n. (HO); Nichols Cap, 41° 33' S., 148° 06' E , A
Moscal /5i(HO); Franklins Road, Kellevie, 42° 47' S., 147° 48' E., Nov. 1974, D.A. &A. V. Ratkowsky 1317
(CANS, HO, K); Hospital Creek, Kellevie, 42°47'S., 147° 48' E., Dec. 1973, D.A. &A.V. Ratkowsky s n

Prossers Sugar Loaf, 42°s 40' S., 1 47° 49' E., Sept. 1 973, D.A. &A. V. Ratkowsky s.n. (HO); E. of Coppin,
Kellevie, 42° 47' S., 147° 48' E., 1973, D.A. & A. V. Ratkowsky s.n. (K); NE. coast, s.dat., L. Rodway s n [ex
herb. Rodway 334] (HO).

Cultivated — Royal Botanic Gardens, Kew, Richmond, Surrey, England, anno. 1976, s.leg. (K).

Distribution:

On the east coast of Tasmania between 41° and 43° S., from near sea level to an
altitude of 650 m.

Ecology:

In shady situations in open sclerophyll forest on hillsides and stony slopes.
Flowering September to March.

Notes:

The T-shaped hairs found on the stems, dorsal leaf surfaces, involucral bracts,
florets, styles and achenes of this species place it in Olearia sect. Dicerotriche Archer ex
Benth. where it would appear to be closely related to O. persoonioides and O.
tasmariica W. Curtis. Apart from the obvious differences in leaf shape and size, O.
archeri can be distinguished from both these taxa by the glandular hairs found on its
involucral bracts, the clusters of heads on a common peduncle, its basally briefly
sagitate rather than distinctly tailed anther bases, its ovate rather than triangular
anther appendages, and the T-shaped hairs on its styles. A curious feature of both O.
archeri and O. tasmanica is the complete absence of the duplex hairs which
predominiate on the achenes of O. persoonioides.

The specific epithet honours William Archer (1820-1874), the pioneering
botanical collector in Tasmania, whose study of trichome and pappus morphology in
Olearia laid the ground for subsequent delimitation and classification of the genus
(Archer 1861).

ACKNOWLEDGEMENTS

I wish to thank Mr J. J. Rainbird for technical assistance and for preparing the
illustration, and Mr M. I. H. Brooker for providing the Latin diagnosis.

I am indebted to Ms Gillian Perry (PERTH) and an anonymous referee for their
advice on the typification of Olearia persoonioides var. lanceolata Benth.

Much vital background work towards my ongoing revision of Australian Olearia
was carried out during my assignment as Australian Botanical Liaison Officer at Kew
during 1984-5. This work has been further supported by Australian Biological
Resources Study Grants in 1988 and 1989.

REFERENCES

Archer, W. (1861). On the value of hairs, as a character in determining the limits of subordinate groups of
species, considered in connexion with the genera Eurybia (Cass.) and Olearia (Monch), of Compositae.
Proc. Linn. Soc. London 5: 17-25.

Bentham, G. (1867). ‘Flora Australiensis’, vol. 3. (Lovell Reeve & Co.: London.)

Hooker, J. D. (1847). Florae tasmaniae spicilegium. Lond. J. Bot. (Hooker) 6: 106-125.

Morris, D. I. (1977). Miscellaneous notes on endemic Tasmanian plants in the genera Olearia. Ixodia,
Danthonia, Tetrarrhena. Records of the Queen Victoria Museum 55: 1-5.

Received 21 July 1988

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OLEARIA ASTROLOBA (ASTERACEAE: ASTEREAE), A NEW SPECIES
ENDEMIC TO VICTORIA

by

N. S. Lander* and N. G. Walsh+

ABSTRACT

Lander, N. S. & Walsh, N. G. Olearia astroloba (Asteraceae: Astereae), a new species
endemic to Victoria. Muelleria 7(1); 123-125 (1989). — Olearia astroloba Lander &
N.G. Walsh, a new species known from a single, small population in Gippsland,
Victoria, is described and illustrated. Its conservation status is discussed.

INTRODUCTION

Olearia Moench (Asteraceae: Astereae) in Australia is currently the subject of
ongoing revisionary studies by the first author. A recently discovered species, "Olearia
sp. (Nunniong)’ of Forbes & Ross (1988), would appear to represent one of Victoria’s
most geographically restricted plants. It is described here in order to facilitate
gazetting it for conservation purposes.

Olearia astroloba Lander & N. G. Walsh, sp. nov.

Species Oleariae phlogopappae affinis a qua capitulis solitariis terminalibus sessilibusque, flosculis

tubuliformibus purpureis habentibus lobos abaxialiter pilis stellatis differt. (Fig. 1).

Typus: Victoria, Gippsland, Marble Gully area, 200 m S. from Old Hut Ceeek, c. 6km
E.of‘Bindi’,37°05'05"S., 147° 5 F 30" E., altitude 840 m, 30June 19SS, N.G. Walsh
2086 & D.E. Albrecht (Holotypus; MEL 1557910. Isotypi: AD, PERTH).

Shrub to 0-5 m high. Vestitureof stems, leaves and outer involucral bracts densely
pannose with stellate hairs. Stems erect, smooth, pale green when young, becoming
brown. Leaves alternate, crowded, inclined, sessile; lamina somewhat incurved,
spathulate, 5-18x2-9 mm, somewhat discolourous, greyish green, abaxially paler,
smooth; venation obscure apart from midrib; texture somewhat coriaceous; base
attenuate; margin dentate towards the apex, thickened or revolute; apex obtuse,
muticous. Heads solitary, terminal, sessile, conspicuously radiate, 15-32 mm in
diameter; disc c. 6 mm in diameter. Involucre obconic; bracts 3-4-seriate,
4-0-7-2X l-()-l-3 mm. Outer involucral bracts flat, narrowly triangular; stereome
green; margin chartaceous, entire; apex narrowly acute. Inner involucral bracts flat,
narrowly ovate; stereome green; margin chartaceous, fimbriate, with scattered basally
stellate hairs; apex acute. Receptacle slightly convex. Ray florets c. 20, mostly
uniseriate, female, llT-15-5mm long; tube with abaxial, multicellular, biseriate,
eglandular hairs scattered centrally to apically; ligule narrowly elliptic,
7-4-12-5X 1-8-3-0 mm, violet, glabrous, obtuse and minutely 3-lobed apically;
staminodes absent; styler arms filiform, 1-3- 1-6 mm long. Disc florets c. 12-35,
bisexual, purple, becoming pale basally, infundibular, 5-5-6-5 mm long, with multi-
cellular, biseriate, simple eglandular hairs scattered abaxially; lobes 5, 1-0-1 -5 mm
long, acute, weakly stellate-hairy abaxially; anthers 1-9- 2-2 mm long, narrowly acute
basally and shorter than the filament collar, with narrowly ovate to triangular, sterile
apical appendage; filament collar 0- 3-0-4 mm long; stylar arms 1 - 3-1 - 5 mm long with
half-conic, sterile apical appendages bearing botuliform papillae above the stigmatic
lines. Achene narrowly obovoid, 2- 1-2*5 x 0*7-1 *0 mm, brown or purplish, sericeous
with duplex hairs; venation distinct with 6 ribs; carpopodium slightly oblique. Pappus
biseriate with an inner row of 20-24 free, minutely barbellate bristles subequal to the

STauS! A^a of Conservation & Land Management, P.O. Box 104, Como,

+ National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

123

124

Fig. 1 . Olearia astroloba Under & N. G. Walsh. A — Habit. B — Ray floret. C — Disc floret. D — Anther
tube. E — Stylar arms. F — Multicellular, biseriate, simple, eglandular hair from floral tube. G —
Stellate hair from leaf. Drawn from the original collection, K.C. Norris £ P. Kelly s.n. (MEL

tubular florets and an outer row of many shorter, flattened, minutely barbellate
bristles. Chromosome number. n = 9 (from holotype).

Other Specimens Examined (Total number examined 6):

Victoria — type locality, 30 June 1988, N.G. Walsh 2087 & D.E. Albrecht (MEL 1557922, NSW,
PERTH), N.G. Walsh 2093 (MEL 1 557921); Marble Creek ( = Old Hut Creek) which flows into'Bindi’ below
NW. extremity of Nunniong Plateau 3-5 km NE. from Mt Simson, c. 64 km due N. from Bruthen,
37°05'05" S., 147° 50' 45" E., Victoria, 29 June 1987, K.C. Norris <6 P. Kelly s.n. (MEL 1555686); same
locality as previous, 29 June 1988, N.G. Walsh 2085 & D.E. Albrecht (CANB, MEL 1557919, PERTHy

Distribution:

Olearia astroloba is known from a single population of c. 1 ,000 plants over an area
of c. 40 ha in eastern Victoria. The area is included in the Victorian Alpine Study Area
(Land Conservation Council 1977) and overlaps the boundary between the Victorian
10' Plant Grids W5 and W6 (Churchill & de Corona 1972).

Ecology and Conservation Status:

Soils at the site are skeletal, light reddish loams derived from marble occurring at
a geological interface between Silurian sediments and rhyolitic Snowy River
Volcanics. Plants occur only on slopes of north to north-westerly aspect, of incline
mostly more than 30°, or occasionally on sheer cliff-faces. The predominant
community at the site is a dense shrubland dominated by Pomaderris oraria F. Muell.

125

sens. lat. with occasional emergents of Allocasuarina verticillata (Lam.) L. Johnson
and Eucalyptus nortonii (Blakely) L. Johnson.

The site supports a unique association of rare or disjunctly occurring species,
including Helichrysum adnatum (DC.) Benth., Pimelea flava subsp. dichotoma
(Schldl.) Threlfall and Pultenaea densiflora F. Muell. The Helichrysum is known from
only one other site in Victoria (near Suggan Buggan, c. 50 km to the NE. and the
Pimelea and Pultenaea are not known elsewhere in eastern Victoria, being plants
typically of mallee scrubs in the north-west of the state.

The site lies on unreserved crown land, approximately 2 km from a freehold
grazing property. Its security from a conservation perspective is uncertain as it is
currently the subject of an application for a permit to quarry marble.

Peak flowering of Olearia astroloba appears to be June-July, but occasional
flowers have been observed as late as November.

This species is proving to be amenable to propagation from cuttings and
specimens are being raised at the Royal Botanic Gardens, Melbourne, and at a private
nursery at Swifts Creek in Gippsland near the Marble Gully site.

O. astroloba is regarded as a vulnerable species and its conservation code is
assessed as 2V (Briggs & Leigh, in press).

Notes:

The presence of stellate hairs on the vegetative parts of O. astroloba is typical of
Olearia sect. Asterotriche Archer ex Benth. (Bentham 1 867) where this species would
seem best placed, pending the completion of revisionary studies in the genus. The
characteristic leaf morphology, the solitary, terminal, sessile heads, and the blue
tubular florets with their wealdy stellate-hairy lobes (from which feature the specific
epithet is derived) readily distinguish it from other species in the section. Its affinities
would seem to lie with O. phlogopappa (Labill.) DC. sens. lat. Occasional specimens of
the latter species and of O. asterotricha (F. Muell.) F. Muell. ex Benth. also have stellate
hairs on the lobes of their tubular florets.

ACKNOWLEDGEMENTS

We are grateful to Mr J. J. Rainbird (PERTH) for technical assistance and for
preparing the illustration, to Dr P. S. Short (MEL) for the chromosome count, to Mr
M. I. H. Brooker (CANB) for providing the Latin diagnosis, and to Mr K. C. Norris
and Mr P. Kelly (Bairnsdale Region, Department of Conservation, Forests and Lands)
and Mr D. E. Albrecht (MEL) for their collection and field observations of Olearia
astroloba.

REFERENCES

Bentham, G. (1867, not 1866). ‘Flora Australiensis’, vol. 3. (Lovell Reeve & Co.: London ) Olearia op
463-488.

Briggs, J. D. & Leigh, J. H. (in press). Rare or threatened Australian plants: the 1988 revised edition
Australian National Parks & Wildlife Service Special Publication 14.

Churchill, D. M. & de Corona, A. (1972). ‘The Distribution of Victorian Plants’. (Royal Botanic Gardens &
National Herbarium, and Botany Department, Monash University: Clayton.)

Forbes, S J. & Ross, J. H. (1988). ‘A Census of the Vascular Plants of Victoria’. Edn 2. (National Herbarium
of Victoria: S. Yarra.)

Land Conservation Council (1977). ‘Report on the Alpine Study Area’. (Land Conservation Council'
Melbourne.)

Manuscript received 21 July 1988

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VALID PUBLICATION OF THE NAMES TORRENTICOLA AND
T. QUEENSLANDICA (PODOSTEMACEAE)

by

Helen I. Aston*

ABSTRACT

Aston, Helen I. Valid publication of the names Torrenticola and T. queenslandica
(Podostemaceae). Muelleria 7( 1 ): 1 27- 1 29 ( 1 989). — This article provides the correct
citations for the names Torrenticola and T. queenslandica and discusses the early
publications where these names occurred both invalidly and validly.

CITATIONS

There is considerable variation in the literature as to when, where and by whom
the generic name Torrenticola and the specific name T. queenslandica were validly
published under the International Code of Botanical Nomenclature (Greuter et al.
1988). The correct citations for these names, together with some previously used
incorrect citations, are given here and a discussion of them follows.

Torrenticola Domin ex Steenis, FI. Males, ser. 1,4:66 (1 949).

Torrenticola Domin, Biblioth. Bot. 89^: 150 (1926) [Jan. 1926, not 1925], nom.

prov.

Torrenticola Domin ex Steenis, J. Arnold Arbor. 28: 421 (1947), nom. inval.

Torrenticola queenslandica (Domin) Domin ex Steenis, FI. Males, ser. 1, 4: 68
(1949).

Podostemum queenslandicum Domin, Biblioth. Bot. 89^: 149 (1926) [Jan. 1926,
not 1925], as Podostemon.

Torrenticola queenslandica Domin, op. cit. 1 50, nom. prov., and 89®: tab. 35, figs
7-13 (1928), nom. inval.

Torrenticola queenslandica (Domin) Domin ex Steenis, J. Arnold Arbor. 28: 421
(1947), nom. inval.

DISCUSSION

Domin (1926) published a new species ‘P. ?queenslandicus n. sp.’ under
Podostemum [as Podostemon] Michaux. In his discussion following the species
description he suggested that the new species might actually belong to a separate genus
and provisionally designated this as ‘Torrenticola n. gen.’. In further discussion he
referred to his new species as ‘T. queenslandica'.

Under Artieles 34.1 and 34.2 of the International Code of Botanical
Nomenclature (Greuter et al. 1988) I accept that the specific name Podostemum
queenslandicum was validly published at this time. Both Domin’s wording and the
layout of the printed text show that Domin was actually accepting the new species
within Podostemum although he indicated some taxonomic doubt. The epithet
queenslandicum, at specific rank, is therefore attributable to Domin and originates
from P. queenslandicum Domin (1926).

In contrast, the generic name Torrenticola and the specific combination T.
queenslandica were not validly published by Domin in his 1926 paper. Torrenticola is
not valid according to Article 34.1(b), which states clearly that a name is not validly
published when it is merely proposed in anticipation of future acceptance, i.e. when it
is a so-called provisional name. The name T. queenslandica is not validly published for

•National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

127

128

the same reason and also because the generic name Torrenticola had not been
validated. Article 43.1 indicates that the name of a species is not validly published
unless the name of the genus to which it is assigned is, either simultaneously or
previously, validly published.

Domin (1928) later published a plate with seven detailed figures of his new
species. These figures were captioned "Torrenticola queenslandica Dom.’. Apparently
Domin was accepting by 1928 that the new species warranted the erection of a new
genus and was accepting his former provisional generic name Torrenticola as being
applicable. Article 42.2 allows for an analytical illustration to replace the written
description or diagnosis usually required for valid publication. However, this Article
applies only to publications prior to 1 January 1908 and therefore cannot be invoked
for validation of the names Torrenticola and/or T. queenslandica from Domin’s 1928
plate. The caption apparently satisfies Articles 4 1 .2(b) and 4 1.3(b) which allow for the
names of genera and species to be validly published by a reference (direct or indirect)
to a previously and effectively published description or diagnosis ‘of a genus’ (for
genera) or ‘of a species or infraspecific taxon’ (for species). The indication of the
author’s name in the caption "Torrenticola queenslandica Dom.’ renders the caption
acceptable under Article 32.4 as an indirect reference to Domin’s 1 926 paper (where a
specific description and a generic diagnosis appear). However, Articles 41.2(b) and
41.3(b) do not apply in the present instance as Article 42.1 indicates clearly that any
reference, whether direct or indirect, to an earlier description or diagnosis is not
acceptable in cases of simultaneous validation of a generic and a specific name. For
this reason (and for a further provision of Article 42. 1 ; see next paragraph) the names
Torrenticola and T. queenslandica can not be considered simultaneously validly
published in Domin’s 1928 publication.

Most current publications, including Index Nominum Genericorum (Farr et al.
1979), attribute the valid publication of Torrenticola and T. queenslandica to Van
Steenis ( 1 947) in the Journal of the Arnold Arboretum. In that publication Van Steenis
ciiQd"TorreriticolaqueenslandicaT>om.minBM. Bot. 89^: 149,pl. 35,fig. 7-13. 1925’,
thereby basing his acceptance of both the generic and specific names on Domin’s
papers of 1 926 [not 1 925] and 1 928 and automatically accepting the same type as that
used by Domin. Van Steenis gave no description or diagnosis, nor did he mention the
generic name Torrenticola independently of its use within the specific name. Article
42.1 (‘The names of a genus and a species may be simultaneously validated by
provision of a single description or diagnosis . . .’) may have been applied to Van
Steenis’s publication except that his paper lacks a description or diagnosis and his
reference to Domin’s prior description is unacceptable because Article 42. 1 further
states that, in cases of simultaneous validation, ‘. . . Reference to an earlier description
or diagnosis is not accepted . . .’. The same Article also disallows simultaneous
validation of generic and specific names when any prior name (at any rank) has already
been validly published based on the same type. In the present case Podostemum
queenslandicum Domin (1926) constitutes such a prior name. I can find no
justification for accepting either Torrenticola or T. queenslandica as being validly
published by Van Steenis in the Journal of the Arnold Arboretum cited.

Engler in Engler and Prantl, Nat. Pflanzenfam. edn 2, 1 8a: 484 ( 1 930), referred to
Torrenticola and T. queenslandica but, following Domin’s works which he cited,
retained these under Podostemum [as Podostemon], The names therefore remain
invalid in Engler’s work.

Van Steenis (1949) included Torrenticola and T. queenslandica in his account of
Podostemaceae in Flora Malesiana and this must surely be accepted as the place and
date of valid publication of both names. Here Torrenticola is clearly accepted in both
the key and the text as the name of a distinct genus and Domin’s two papers are cited
under Torrenticola thus: ‘Domin, nom. prov. Bibl. Bot. 89, 2 (vol. 20) ( 1 925) 1 49, t. 35,
f. 7-13’. The citation of p. 149 can be treated merely as a bibliographic error of
citation, p. 1 50 being the actual page on which Domin proposed the provisional name
Torrenticola n. gen. and gave his Latin diagnosis of it. Van Steenis (1949) therefore
satisfies the requirements of Article 36. 1 for a reference to a previously and effectively

129

published Latin description or diagnosis. As he attributed the generic name to Dornin
the complete citation of it is Torrenticola Domin ex Steenis; this may be shortened to

Torrenticola Steenis (Article 46.3). . . , ...

Van Steenis ( 1 949) cited ‘Torrenticola queenslandica Domin, l.c. under the genus
Torrenticola. As we have seen, Domin had only mentioned this combination
provisionally but had validly published Podostemum queenslandicum Domin on
p. 1 49 of his 1 926 paper cited by Van Steenis. Van Steenis ( 1 949) therefore validated a
new combination Torrenticola queenslandica (Domin) Steenis (1949) based on
Podostemum queenslandicum Domin (1926) (Article 49.1). As Domin also had
provisionally suggested the name T. guccnslciHdicci the complete but optional citation
for it (Article 46.3) is T. queenslandica (Domin) Domin ex Steenis (1949).

REFERENCES

Farr, E. R. etal. (eds) (1979). Index nominum genericonim (plantarum). Vol. 3, Pegaeophyton-Zyzygium.

Regnum Veg. vol. 102. . „ i , , o

Greuter, W. et at. (eds) (1988). International code of botanical nomenclature. Regnum Veg. vol. 118.

Revised manuscript received 2 August 1988

ACACIA CAERVLESCENS, A NEW SPECIES OF ACACIA SECTION
PHYLLODINEAE FROM VICTORIA

by

B. R. Maslin' and A. B. Court^

ABSTRACT

Maslin, B. R. & Court, A. B. Acacia caerulescens, a new species of Acacia section
Phyllodineae from Victoria. Muelleria 7(1): 131-134(1989). — Acacia caerulescens, a
new species referable io Acacia section Phyllodineae T>C. and allied to^. obliquinervia
Tind., is described and illustrated. It is endemic in eastern Victoria (Buchan-Lakes
Entrance district) where it is restricted to limestone soils.

TAXONOMY

Acacia caerulescens Maslin & Court, sp. nov.

Acacia obliguinerviae affinis a qua imprimis difFert phyllodiis 4-8 cm longis, (l-)l-5-3cm latis,
I:w = 2-3-5(-6), glaucissimis, costa saepe nec valde excentrica, glande 5-25 mm supra pulvinum
posita versus costam per nervo tenui oblique conjuncta, pedunculis c. 6 mm longis aliquantum
gracilibus, capitalis in alabastro globosis.

Allied to A. obliquinervia but differing principally in the following ways.
Phyllodes 4-8 cm long, (l-)l-5-3 cm wide, 1 :w = 2-3-5(-6), very glaucous, midrib
often not markedly excentric. Gland 5-25 mm above the pulvinus and connected to
midrib by a fine, oblique nerve. Peduncles c. 6 mm long, rather slender. Heads
globular when in bud.

Typus; Beside Wulgulmerang-Buchan road, c. 4-3 km from Buchan by road,
Gippsland, Victoria, 10 Nov. 1985, A.B. Court CBG 8506135 (Holotypus: CBG;
IsoTYPi: AD, B, BRI, G, K, L, NSW, NY, MEL, PERTH, US).

Tree to 1 0- 1 5 m tall, often ± pyramidal. Branches terete but very slightly angled
at extremities, finely and obscurely ribbed, at least the youngest shoots slightly to
moderately pruinose. Phyllodes slightly oblique, obovate to oblanceolate or elliptic to
narrowly elliptic, rather abruptly narrowed into an obtuse apex, 4-8 cm long,
(l-)T5-3 cm wide, 1 :w = 2-3-5(-6), thinly coriaceous, straight or frequently slightly
recurved near the somewhat narrowed base, glabrous, glaucous; midrib apparent,
central or slightly to markedly excentric (i.e. situated closer to the upper margin),
yellowish to light brown; lateral nerves not pronounced, loosely anastomosing;
marginal nerves yellow to light brown; pulvinus 3-5 mm long, wrinkled and brown to
dark brown when dry. Gland solitary, 5-25 mm above the pulvinus and connected to it
by a fine oblique nerve whieh is concurrent with the midrib for a short distance, and
often branched at its point of divergence with one branch extending to the gland and
the other rejoining the midrib. Racemes axillary and terminal, often arranged in
panicles which may reach 9 cm long; raceme axes with 2-8 flower-heads, slightly
flexuose, glabrous, variably pruinose, base ebracteate. Peduncles c. 6 mm long, rather
slender (c. 0-5 mm wide when dry), glabrous, variably pruinose, with 2, basal,
triangular, glabrous bracts (?homologous to stipules) <0-5 mm long, an extremely
reduced phyllode often present between the bracts. Flower-heads globular,
(15-)20-30-flowered, lemon yellow, lightly scented. Bracteoles spathulate to
sub-peltate; claws linear but slightly dilated towards their fimbriolate apices, c. 1 mm
long (equalling calyx); laminae circular to triangular-ovate, apiculate, c. 0-5 mm long
glabrous, brown. Flowers 5-merous. Calyx gamosepalous, f length of corolla, very
shortly divided (for c. j its length or less) into ± broadly triangular, slightly inflexed.

■Western Australian Herbarium, Department of Conservation and Land Management P O Box 104
W. A., Australia 6152. o , ■ ,

^Australian National Botanic Gardens, P.O. Box 1777, Canberra City, A.C.T., Australia 2601
Present address: 71 Miller St., O’Connor, A.C.T., Australia 2601.

Como,

131

132

Fig. i. Acacia caerulescens Maslin & Court, a — Portion of branch, b — Phyllode. c & d — Base of phyllode
showing fine oblique nerve extending from gland (g) to pulvinus (p), note in D that this nerve bifurcates,
e — Legume, f — Seed, a from A.B. Court CBG 8506135 (PERTH isotype); b & d from MEL 615165;
cfrom MEL 615166; e-ffrom J. Kenrick s.n. (PERTH 00721689).

slightly keeled lobes. Petals 2-2-1-5 mm long, glabrous. Legumes oblong to narrowly
oblong, 5-12 cm long, 1-4-2-2 cm wide, flat but rounded over seeds along midline of
legume, slightly constricted between the seeds although occasional deep constrictions
occur on some legumes, chartaceous, straight to very slightly curved, transversely
loosely reticulate, glabrous, light- to medium-brown or purplish brown, pruinose
(pruinosity patchy), stipitate, marginal nerve quite prominent but scarcely thickened.
Seeds longitudinal or more usually longitudinally oblique in the legume, ellipsoid.

133

5-6 mm long, 3-5-4 mm wide, compressed (c. 2 mm thick), dull, black; pleurogram
narrowed but open towards the hilum, not particularly prominent; areole 4 mm long,
2 mm wide; funicle at first straight, c. 2 mm long, normal to margin and closely
appressed to inner wall of legume, then abruptly thickened, li^t red-brown and
once-folded below the yellow, more or less clavate aril, aril constricted at its junction
with the funicle.

Other Specimens Examined:

Victoria — Stony Creek crossing of the Princes Highway, Toorloo Arm, D.E. Albrecht 2276 (CBG,
PERTH); 2 mi [3-2 km] NNE. of Swan Reach on Bruthen Road, A.C. Beauglehole 37683 (MEL); Toorloo
Arm Reserve, Princes Highway, A.C. Beauglehole 37851 (MEL); Cultivated, Buffalo Creek near Myrtleford,
F.E. Bienvenu 0623 (PERTH); Tambo Upper road, L.A. Eell 3149 (MEL); Cultivated, Melbourne Botanic
Gardens, 10 Jan. 1986, J. Kenrick s.n. (PERTH 00512044); Cultivated, near Baileau Library, Melbourne
University, 7 Jan. 1984,/ Kenrick s.n. (PERTH 00721689); Buchan River, Mar. 1854, E. Mueller s.n. (MEL
615167); Buchan, about 1939, E. Robins s.n. (MEL 522743); Murrindal, N.A. Wakefield 4103 (MEL);
Buchan, 1 1 Nov. 1964, J.H. Willis s.n. (MEL 502498); On rocks at the Buchan River, anonymous (MEL
61514); Buchan River, anonymous (MEL 615165); Tambo River, anonymous (MEL 615166).

Distribution:

Restricted to the Lakes Entrance-Buchan district in eastern Victoria where it is
known only from the Swan Reach-Tambo Upper area and Lake Tyers north to the
Murrindal area.

Ecology:

Key associates of the Buchan populations include Eucalyptus melliodora Cunn.
ex Schauer, Acacia falciformis DC. and Themeda triandra Forssk., forming a grassy
woodland. By contrast, the Toorloo Arm population is associated with Eucalyptus
baueriana Schauer and E. globulus ssp. pseudoglobulus (Naudin ex Maiden) Kirkpatr.
open forest with a shrubby understorey dominated by Pomaderris oraria sens. lat.

The population at Toorloo Arm grows on clay over fossiliferous limestone. It
appears that the distribution of this species is strictly associated with limestone
geology.

Conservation Status:

Populations of this species have been fragmented and depleted historically by
land clearance for settlement and agriculture. Remnant populations are currently
threatened by further land clearing, especially on private land, and particularly by
roadworks.

Since 1985 there have been major road widening operations with local
realignments north from Buchan along the Wulgulmerang-Buchan road, to about
half-way between Buchan and Murrindal. These have significantly reduced the
populations in the roadside reserves, which are already remnants adjacent to
agricultural land. The roadworks have extended as far north as all the known roadside
populations of the species.

In the vicinity of the Princes Highway crossing over Toorloo Arm, within the
Lake Tyers Forest Park, there is preserved a significant population containing a
number of mature, 1 0- 1 5 m tall specimens. However, several outstanding individuals
on the east side of the Arm, overlooking the Princes Highway, have been lost recently
through major road reconstruction works at the crossing.

Flowering and Fruiting Period:

Flowering commences in early November but its termination is not known;
immature legumes (resulting from the previous flowering season) are often present
with flowers in November. Legumes with mature seeds have been collected in early
January.

Affinities:

On account of its globular flower-heads and 1 -nerved phyllodes,4. caerulescens is
placed m Acacia section Phyllodineae DC. The species was included by Court (1973,
page 224) and Costermans (1981, page 318) as a variant of A. obliquinervia. The two
species have glabrous, pruinose branchlets and similar phyllode, inflorescence and

134

carpological features. Acacia obliquinervia is most readily distinguished from A.
caerulescens by its grey-green to glaucescent phyllodes which are often longer (5- 1 7 cm
long) and which possess a gland (0-12 mm above pulvinus)ithat does not have an
associated fine oblique nerve extending to the pulvinus. Also, A. obliquinervia has
thicker, shorter peduncles (1-5— 5 mm long) and often oblongoid flower-head buds.
obliquinervia is widespread in Victoria and New South Wales where it occurs in ranges
between 500 m and 1700 m, especially in montane forests (Costermans 1981). The
new species occurs at lower elevations on limestone soils and has a very restricted
geographic range; it is not known to be sympatric with A. obliquinervia.

Cultivation:

According to Elliot and Jones (1982, p. 132) this very ornamental species will
grow in a wide range of soils, is best suited to partial or full sun and withstands frosts
and extended dry periods. It is ideal for gardens and roadsides, windbreaks and shelter
planting. It is very attractive on account of its blue foliage and bright, lemon yellow
heads.

Etymology:

The specific epithet refers to the characteristic blue foliage.

Common Name:

Buchan Blue or Buchan Blue Wattle.

ACKNOWLEDGEMENTS

Mr D. G. Cameron (Botany Department, Latrobe University, Victoria) is
gratefully acknowledged for providing the ecological and conservation status data
presented here. Bill Molyneux (Montrose, Victoria) is thanked for providing
observation on flowering and fruiting phenology. Paul Wilson (W.A. Herbarium) is
thanked for providing the Latin diagnosis.

REFERENCES

Costermans, L. (1981). ‘Native trees and shrubs of south-eastern Australia’. (Rigby: Melbourne.)

Court, A. B. (1973). In Willis, J. H. ‘A handbook to plants in Victoria’. (Melbourne University Press:
Carlton.)

Elliot, W. R. & Jones, D. L. (1982). ‘Encyclopaedia of Australian Plants suitable for cultivation’, vol. 2.
(Lothian Publishing Company: Melbourne.)

Manuscript received 16 August 1988

NOTES ON HOVEA R. Br. (FABACEAE): 3
by

J. H. Ross*

ABSTRACT

Ross, J. H. Notes on Hovea R. Br. (Fabaceae): 3. Muelleria 7(1); 135-139 (1989). H.
arnhemica from the Northern Territory is described as new. H. longifolia R. Br. var.
purpurea (Sweet) Domin subvar. planifolia Domin from south-eastern Queensland is
elevated to species rank as H. planifolia and the necessary combination is made.

HOVEA ARNHEMICA
Hovea arnhemica J.H. Ross sp. nov.

Affinite incerta, forsan H. planifoliae (Domin) J.H. Ross affinis, a qua corolla semper albida, planta
multo minore, caulibus brevibus erectis vel decumbentibus, foliis minoribus lamina plus minusve
plana, inflorescentia sessili, floribus minoribus, et arillo seminis quam semine ultra dimidio breviore,
differt.

Typus: Northern Territory, Arnhem Land, 3 km NW. of Murgenella, 2.iv.l984, C.R.
Dunlop 6673 (Holotypus: MEL; Isotypi: AD, BRI, CANB, CBG, DNA, HO, K,
NSW, PERTH).

Subshrub to 60 cm high with a large lignotuber, multi-stemmed, the stems erect or
decumbent, densely clothed with short curled hairs and longer straighter hairs up to
1 -4 mm long, the hairs faintly or distinctly rusty-brown. Leaves spreading almost at
right angles to the stem or deflexed, lamina more or less flat on upper surface on either
side of the depressed midrib, lamina of upper leaves narrow-ovate or elliptic and of
smaller lower or basal leaves ovate, obovate or subrotund, (0-7)2-6-3 cm long,
(0-5)0-8-l-5 cm wide, apex acute, obtuse or sometimes emarginate, upper surface
densely clothed with short curled white hairs, lower surface densely clothed with
curled and scattered strai^tish white hairs except on midrib and margin where hairs
are tinged with brown; petiole up to 0-3 cm long, densely clothed like the stem. Stipules
subulate, up to 3 mm long, spreading laterally or somewhat recurved and often
persisting for some time, glabrous and glossy above, densely pubescent on lower
surface. Inflorescence axillary, sessile, mostly 2- or 3-flowered, rarely flowers solitary.
Flowers pedicellate, the pedicels up to 1-5 mm long, densely clothed with spreading
hairs; bracteoles narrow-ovate, 2-3-5 mm long, 0-8-1 - 3 mm wide, situated at the base
of the calyx and shorter than or almost as long as the calyx-tube, inner surface with
scattered appressed hairs, outer surface densely clothed with spreading hairs, with a
conspicuous tuft of dark reddish-brown hairs in the axils; bract inserted at the base of
the pedicel, narrow-ovate, 1 -8-2-4 mm long, 1-1-2 mm wide, outer surface densely
clothed with spreading hairs. Calyx densely clothed with short curled and longer
straighter rusty-brown to whitish hairs: 2 upper lobes 5-5-6 mm long including the
tube 1 -5-2-5 mm long, the 3 lower lobes 2-2-7 mm long, acute. Standard 6-6-6 mm
long, 7-8 mm wide, emarginate apically, white except for a greenish-yellow basal flare;
wings 5-7-6 mm long, 2-2-2-8 mm wide; keel 4-8-5-2mm long, 2-2-5 mm wide.
Stamen-filaments 3-5-4-5 mm long. Ovary sessile or very shortly stipitate,
1 -2-1 - 5 mm long, 2-ovulate. Pods sessile or almost so, obliquely ovoid or ellipsoid or
sometimes transversely elliptic, 1-1- 1-4 cm long, 1-1- 1-4 cm wide, densely clothed
with white matted hairs externally, fairly densely clothed with weak white hairs within.
Seeds elliptic, plump, 4-7-5-8 mm long, 3-3-4-2 mm wide, 2-8-3-5 mm thick,
chestnut-brown, hilum linear, the aril less than half the length of the seed, with a raised
lateral lip. (Fig. 1).

•National Herbarium of Victoria, Birdwood Avenue, South Yarra, Victoria, Australia 3141.

135

136

Fig. 1. Hovea arnhemica. A — flowering and fruiting twig, XI. B — surface of portion of stem showing
indumentum, X5. C — calyx opened out (upper lobes on right), X4. D — standard, X4. E — wing >
petal, X 4. F — keel petal, X 4. G — gynoecium, X 9. H — seed, side view, X 4. 1 — seed, hilar view, X 4.
A-H from C. R. Dunlop 6673 (MEL 113278 and 1 13279).

137

Restricted in distribution to Arnhem Land north and north-west of Oenpelli in
the Northern Territory (see Fig. 2). Recorded on deep sandy soil in open Eucalyptus,
Gronophyllum-Eucalyptus-Livistona and Ptychosperma elegans forest. Often locally
common.

Representative Specimens (Total number examined 15):

Northern Territory — 4S km N. of Oenpelli, 15.vii.l961, G.M. Chippendale sah NT 8101 (NT, MEL).
4 km SSW. of Raffles Bay, 19.vii.l961, G.M. Chippendale sub NT 8216 (NT, MEL). Port Essington,
21.iv.l818, A. Cunningham (CGE). Smith Point, Cobourg Peninsula, 8.X.1971, J. Must 827 (DNA, NSW,
NT). 1 km W. of Nabarlek, 12° 18' S., 133° 18' E., 22.iv.1979, M.O. Rankin 2101 (BRI, DNA). Murgenella
road, Cobourg Peninsula, 11°20'S., 132°22'E., 31.V.1983, G. Wightman 444 & C.R. Dunlop (DNA,
MEL).

Notes:

This species was first collected by Cunningham on 21 April 1818 at Port
Essington on the Cobourg Peninsula when he accompanied Lieutenant King in the
‘Mermaid’ (a duplicate in E numbered 270 from BM bears the date 1817 but
presumably this is in error). Cunningham’s material was referred to by Bentham
( 1 864) as being closest to H. longifolia var. pannosa and was included by Domin (1925)
under H. longifolia var. lanceolata subvar. apiculata. Apparently the species was not
re-collected until the early 1 960s.

H. arnhemica is isolated geographically from all of the other species in the genus
and differs from all of them in that the corollas are apparently always white except for a

138

basal greenish-yellow flare on the standard. The affinities of H. arnhemica are not
clear. The species is perhaps allied to H. planifolia from south-eastern Queensland
from which it differs in being a much smaller plant with short erect or decumbent
stems, in haying smaller leaves spreading almost at right angles to the stem or deflexed,
the leaf lamina more or less flat, sessile inflorescences, smaller flowers and seeds with
an aril less than half the length of the seed.

HOVEA PLANIFOLIA

Domin (1925) followed Bentham (1864) in treating H. longifolia R. Br. as an
‘omnibus’ species and recognised within it five varieties and five subvarieties, one of
which was var. purpurea (Sweet) Domin subvar. planifolia Domin. The brief
description of subvar. planifolia was based on specimens (‘viele Exemplare’) collected
by A. Dietrich from the Brisbane River, south-eastern Queensland, and on a specimen
collected by F. Mueller from the same locality in August 1855.

The following syntypes have been located; Brisbane River, A. Dietrich (BRI
345294, HBG (5 sheets), NSW 166516, PR 527088, 527089, PRC, W 108297,
108299); Brisbane River, F. Mueller, August 1855 (K). MEL 667189 collected by
Mueller from the Brisbane River is almost indistinguishable from the specimen in K
but as it was collected in July 1855 rather than in August it is not regarded as a
syntype.

It is clear that subvar. planifolia represents a taxon distinct from H. longifolia and
that it merits specific rank. The opportunity is taken here of raising subvar. planifolia
to specific rank and of providing a description and notes.

Hovea planifolia (Domin) J.H. Ross comb. & stat. nov.

Basionym: H. longifolia var. purpurea (Sweet) Domin subvar. planifolia Domin,
Biblioth. Bot. 22 (89^): 729, fig. 141 (right hand specimen) (1925). Lectotype (here
selected): Queensland, Brisbane River, August 1855, F. Mueller (K).

Shrub 0-5-2(-2-5) m high, often wider than high, usually with several stems or the
single stem branching at or a short distance above the base, branches ascending and
spreading, branchlets densely clothed with short coiled or crinkled hairs and longer
curled, crinkled or straightish hairs up to 9 mm long, the hairs on the young tips
distinctly bright rusty-brown or reddish-brown. Leaves usually held more or less erect
and almost perpendicular to the lateral branches or almost parallel to the vertical
branches, sometimes held at an angle of about 45° but the lower surface usually
conspicuous: lamina arched up slightly on either side of the depressed midrib
and recurved towards the margin or slightly to distinctly V-shaped in section or
sometimes more or less flat, narrow-ovate or elliptic, (2-)4-10(-13) cm long,
(0-5-)0-9-2(-2-7) cm wide, apex rounded, obtuse or subacute, with a short mucro,
upper surface dark green, sparingly to densely clothed throughout with short curled or
twisted hairs or the hairs largely confined to the midrib and decreasing in frequency
towards the margins, lateral veins sometimes prominent and somewhat raised, lower
surface densely clothed with short coiled or curled hairs and longer curled, twisted or
strai^tish hairs, the hairs greyish-white or rusty-brown, lateral veins often prominent
despite the dense indumentum; petiole 0-3-0-9 cm long, densely pubescent like the
branchlet. Stipules subulate, 2-3-2 mm long, 0-3-0-6 mm wide, densely clothed
externally (abaxial surface) with short curled or crinkled hairs and longer spreading
hairs, the hairs usually conspicuously rusty or reddish brown. Inflorescence axiWairy , on
densely pubescent peduncles 0-2-0-8 cm long and usually 3-flowered or occasionally
the axis growing on to form a many-flowered leafy shoot. Flowers pedicellate, the
pedicels up to 2-5 mm long, densely clothed with short coiled or curled hairs and
longer wavy or straightish spreading hairs; bracteoles narrow-ovate, 2-4-3- 1 mm long,
as long as to slightly longer than the calyx-tube, inserted at the base of the calyx,
densely clothed with coiled, curled and longer wavy or straightish spreading hairs, the
hairs usually bright rusty-brown; bract 2-5-3-5 mm long, inserted almost at the same
level as the bracteoles; bract 2 -5-3 -5 mm long, inserted almost at the same level as the
bracteoles; bract and bracteoles with reddish-brown glandular hairs in the axils. Calyx

139

densely clothed with coiled, curled or longer straighter hairs, the hairs silvery-white
throughout or tinged with rusty-brown especially apically: 2 upper lobes ± truncate,
4-5-2 mm long including the tube 2-3-5 mm long, the 3 lower lobes 1-5-2 mm long.
Standard S-6-9-5 mm long, 9-5-1 1 mm wide, pinkish-purple when young but turning
purplish with age, with a greenish-yellow basal flare; wings 5-7-V-5 mm long,

3- 3-4-2 mm wide; keel petals 4-5-5-9 mm long, 2-3-3 mm wide. Stamen-filaments

4- 2-5-5 mm long. Ovary shortly stipitate, 1 -5-2-5 mm long, densely pubescent,
2-ovulate. Pods shortly stipitate but stipe not exceeding the calyx-tube, obliquely or
transversely ovoid or ellipsoid, 1-1-4 cm long, 0-8-1 -6 cm wide, densely clothed with
curled, twisted and wavy hairs externally, sparingly to densely clothed with weak
curled or wavy hairs internally. ellipsoid, plump, 5-5-6-6 mm long, 3-S-4-2 mm
wide, 3 -5-3 -6 mm thick, chestnut brown, hilum linear, the aril with a very small raised
lateral lip and extending for almost the length of the seed.

Restricted in distribution to the Burnett, Darling Downs, Leichhardt, Maranoa
and Moreton districts of south-east Queensland. Recorded most frequently from
laterite, sandstone or shallow sandy soil among sandstone outcrops in open dry
sclerophyll forest.

Representative Specimens (Total number examined 73):

Queensland — Burnett Distr., 44 km from Mundubbera on Durong Rd, 3.x. 1982, P.I. Forster 1408
(BRI); Darling Downs Distr., Barakula State Forest, 15-2 km E. of Chinchilla-Auburn Rd on road to Ballon
Forest Office, 19.viii.l986, Ross 3095 (BRI, MEL); Leichhardt Distr., Blackdown Tableland, 32 km SE.
of Blackwater, 20.iv. 1971, RJ. Henderson, S.B. Andrews (S P. Sharpe 724 (BRI, MEL, NSW); Maranoa Distr.,
12 km NNE. of Mt Moffatt H.S., 13.v.i982, V.J. Nelder & M.B. Thomas 711 (BRI); Moreton Distr.,
Blackstone, c. 8 km SE. of Ipswich, 3.x. 1959, L. Pedley 486 (BRI).

H. planifolia is distinguished readily by the large leaves which are amongst the
largest found in the genus and by the distinctive bright rusty or reddish-brown hairs on
the apices of the young growing shoots. The name H. pannosa Cunn. ex Hook, has been
misapplied to this plant in Queensland.

An attractive open shrub with a generally greenish-grey appearance which would
appear to have considerable horticultural potential.

ACKNOWLEDGEMENTS

I am most grateful to Dr G. P. Guymer for answering enquiries and for
photographing type specimens while serving as Australian Botanical Liaison Officer
at the Herbarium, Royal Botanic Gardens, Kew; to Mr C. R. Dunlop, Darwin
Herbarium, for making a special trip to collect specimens which serve as the type
material of H. arnhemica; to Mr C. R. Dunlop, Dr G. Leach, Mr G. Wightman and
staff of the Darwin Herbarium for organising a most enjoyable field trip to Arnhem
Land; to Mr L. Pedley, Queensland Herbarium, for accompanying me in the field in
south-eastern Queensland; to Mrs Betty Ballingal, Oakey, Queensland, and Mrs Val
Hando, Chinchilla, Queensland, for making special collections; to Mr A. S. George,
Bureau of Flora and Fauna, Canberra, for correcting the Latin diagnosis; to my
colleague Miss A. M. Podwyszynski for preparing the illustration which accompanies
the text; to Ms J. Hurse for technical assistance; to the Directors/Curators of BRI,
CGE, DNA, E, HBG, HO, K, NSW, PR, PRC and W for the loan of specimens and/or
working facilities in their institutions; to Mrs J. Thomas for typing the manuscript;
and for an Australian Biological Resources Study grant which greatly facilitated this
project.

REFERENCES

Bentham, G. (1864). ‘Flora Australiensis’, vol. 2. (Lovell Reeve; London.)

Domin, K. (1925). Beitrage zur Flora und Pflanzengeographie Australiens. Biblioth. Bot. 22 (89^): 729.

Revised manuscript received 11 October 1988

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CONTENTS

Volume 7, Number 1 Page

Two new lichens: Cladonia paeminosa and C. humilis var. bourgeanica

— Alan W. Archer 1

Eucalyptus cadens (Myrtaceae), a new swamp gum from the Warby Range,
North-East Victoria

— J. D. Briggs and M. D. Crisp 7

Azorella Lamarck (Apiaceae) on Heard and Macquarie Islands, with
description of a new species

— A. E. Orchard 1 5

Notes on Hovea R. Br. (Fabaceae): 2

— J. H. Ross 21

A revision of Podotheca Cass. (Asteraceae: Gnaphaliinae)

— P. S. Short 39

Notes on the fruit anatomy of Australian members of the Inuleae
(Asteraceae)

— P. S. Short, K. E. Wilson and J. Nailon 57

Two new species of Pomaderris Labill. (Rhamnaceae) from South-Eastern
Australia

— Neville G. Walsh 81

A new subspecies of Grevillea acanthifolia (Proteaceae: Grevilleoideae) from
South-Eastern New South Wales

— R. O. Markinson and D. E. Albrecht 89

A new species of Tetrarrhena R.Br. (Poaceae) from Victoria and New South
Wfllcs

— Neville G. Walsh 95

Ferdinand Mueller, Government Botanist: the role of William Hooker in his
appointment

— Helen M. Cohn 99

New genera and species of Australian Inuleae (Asteraceae)

— P.S. Short 103

Olearia archeri (Asteraceae: Astereae), a new name for a familiar species from
Tasmania

— N. S. Lander 1 17

Olearia astroloba (Asteraceae: Astereae), a new species endemic to Victoria

— N. S. Lander and N. G. Walsh 123

Valid publication of the names Torrenticola and T. queenslandica
(Podostemaceae)

— Helen I. Aston 127

Acacia caerulescens, a new species of Acacia section Phyllodineae from
Victoria

— B. R. Maslin and A. B. Court 131

Notes on Hovea R. Br. (Fabaceae): 3

— J. H. Ross 135

ISSN 0077-1813

Jean Gordon, Government Printer, Melbourne