Volume 9 


Number 1 2013 


AUSTROBAIIEYA 

A Journal of Plant Systematics 

and 

Conservation Biology 


Queensland Herbarium 

Department of Science, Information Technology, Innovation and the Arts 


Queensland 

Government 


Volume 9 


Number 1 2013 


A Journal of Plant Systematics 

and 

Conservation Biology 


Queensland Herbarium 

Department of Science, Information Technology, Innovation and the Arts 


iSh QUEENSLAND 
ffilr GOVERNMENT 


Editorial Committee 


P.I.Forster (editor) 

P.D.Bostock (technical advisor) 

G.P.Guymer (technical advisor) 

Graphic Design 

Will Smith 

Desktop Publishing 

Aniceta Cardoza 

Austrobaileya 

Vol. 1, No. 1 was published on 1 December 1977 
Vol. 8, No. 4 was published on 21 December 2012 

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ISSN 0155-4131 


© Queensland Herbarium 2013 

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Austrobaileya is the journal of the Queensland Herbarium and publishes peer-reviewed research 
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Austrobaileya 9 ( 1 ): 1-154 ( 2013 ) 


Contents 

A taxonomic revision of Garcinia L. (Clusiaceae) in Australia, including four 
new species from tropical Queensland 

W.E. Cooper .1-29 

Reinstatement and revision of Sphaeromorphaea DC. and Ethuliopsis 
F.Muell. (Asteraceae: Plucheinae ) 

A.R.Bean .30-59 

Reinstatement of intraspecific taxa for Bosistoa pentacocca 

(F.Muell.) Baill. (Rutaceae) with a new combination B. pentacocca 
subsp. connaricarpa (Domin) P.I.Forst. 

P. I. Forster .60-65 

Three new species of Pluchea Cass. (Asteraceae: Inuleae-Plucheinae) from 
northern Australia 

A.R.Bean .66-74 

Cryptocarya cercophylla W.E.Cooper (Lauraceae), a new species from 
Queensland’s Wet Tropics 

W.E.Cooper .75-79 

Violaperreniformis (L.G.Adams) R. J.Little & G.Leiper, stat. nov., with notes 
on Australian species in Viola section Erpetion (Violaceae) 

R. J.Little & G.Leiper .80-101 

Pterostylis caligna M.T.Mathieson (Orchidaceae), a new species from 
northern Queensland 

M.T.Mathieson .102-106 

Two new subspecies of Eucalyptus tereticornis Sm. (Myrtaceae) 

A. R.Bean .107-113 

A new fern species for Queensland: Diplazium squamuligerum (Rosenst.) 

Parris (Woodsiaceae) 

D.J.Ohlsen & A.R.Field .114-125 

New records for Queensland in Lindernia All. (Linderniaceae) 

B. S.Wannan .126-129 

Status and lectotypification of Craspedia gracilis Hook.f. (Asteraceae: 

Gnaphalieae) 

A.C.Rozefelds .130-135 

Craspedia uniflora G.Forst. (Asteraceae) is a New Zealand endemic and not 
part of the Australian flora 

A.N.Schmidt-Lebuhn .136-139 

Cullen spicigerum (Domin) A.E.Holland (Fabaceae), a new combination and 
reinstatement of a north Queensland species 

A. E. Holland .140-145 


Celtis strychnoides Planch, an earlier name for Celtis australiensis Sattarian 
(Ulmaceae) 

G.P.Guymer .146-147 

Reinstatement of Rapanea leucobrachya P.Royen (Myrsinaceae) from New 
Guinea, with a new combination as Myrsine leucobrachya (P.Royen) 

PI.Forst. 

P.I.Forster. .148-149 

Emmenosperma pancherianum Baill. (Rhamnaceae) newly recorded for 
Australia 

A. R. Be an .150-152 

Euphorbia operta Halford & W.K.Harris (Euphorbiaceae), a new name for 
Euphorbia occulta Halford & W.K.Harris, nom. illeg. and lectotypification 
of the name Euphorbia australis var. subtomentosa Domin 

D.A.Halford & W.K.Harris .153-154 


A taxonomic revision of Garcinia L. (Clusiaceae) in Australia, 
including four new species from tropical Queensland 

W.E. Cooper 
Summary 

Cooper, W.E. (2013). A taxonomic revision of Garcinia L. (Clusiaceae) in Australia, including four 
new species from tropical Queensland. Austrobaileya 9(1): 1-29. The genus Garcinia is revised for 
Australia with nine native and three naturalized species included. Garcinia jensenii W.E.Cooper, 
G. leggeae W.E.Cooper, G. russellii W.E.Cooper and G. zichii W.E.Cooper are newly described and 
illustrated. All species are fully described, together with notes on habitat and distribution. A key to 
Garcinia species in Australia is provided. 

Key Words: Clusiaceae, Garcinia , Garcinia brassii , Garcinia dnlcis , Garcinia gibbsiae, Garcinia 
jensenii , Garcinia leggeae , Garcinia livingstonei, Garcinia mangostana , Garcinia mestonii , 
Garcinia riparia , Garcinia russellii , Garcinia zichii , Garcinia warrenii, Garcinia xanthochymus , 
Australia flora, Queensland flora. Northern Territory flora, new species, taxonomy, identification 
keys, rainforest, naturalised species 

W.E. Cooper, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor 
Road, Smithfield, Queensland 4878, Australia. 


Introduction 

Garcinia L. (Clusiaceae) is a pantropical 
genus of about 260 species (Stevens 2007) 
of mostly dioecious shrubs and medium 
sized trees found in Africa, Madagascar, 
Comoros, the Indian subcontinent (including 
Bangladesh, Bhutan, India & Sri Lanka), 
SE Asia (including Cambodia, China, 
India, Laos, Myanmar, Nepal, Thailand 
and Vietnam), Malesia (including Malaysia, 
Indonesia, the Philippines and New Guinea), 
Solomon Islands, Australia, New Caledonia, 
Fiji and South America. Species are restricted 
to various forms of rainforest between the 
lowlands and the highlands, the exception 
being Garciniaamplexicaulis Vieil 1. exPierre, 
which occurs in exposed areas and secondary 
regrowth thickets on degraded and mined 
areas in New Caledonia. Within Australia, 
12 species (nine native, three naturalised) of 
Garcinia occur in Queensland’s rainforests 
and gallery forests from the Torres Strait 
islands south to the Kirrama Range in the 
Wet Tropics bioregion, with one species (G. 
warrenii) on Melville Island in the Northern 
Territory where it occurs in monsoon 


Accepted for publication 9 September 2013 


vine forest. Seven species are endemic to 
Queensland. 

Garcinia plants have been recorded as 
having white, cream or yellow exudate (Nazre 
2006); whereas, some Australian species also 
have clear or slightly milky exudate. Bracts 
and glands are a feature of this genus and 
there are three different types on most species, 
all are insignificant in species delimitation 
(Nazre 2006). Fruits are baccate with a single 
ovule per carpel, a unique combination in 
Clusioideae and a single ovule per carpel 
is a synapomorphy for the genus (Sweeney 
2008: 1298). However, two Australian 

species, Garcinia gibbsiae S.Moore and G. 
russellii W.E.Cooper produce dry berries and 
commonly contain up to 4 ovules per locule 
and therefore the presence of a single ovule 
per carpel is no longer a diagnostic feature for 
the genus. 

The first record of Garcinia from 
Australia was of G. mestonii F.M.Bailey (as 
‘mestonf ) (Bailey 1889), followed quickly by 
G. warrenii F.Muell. (Mueller 1891). The last 
overall flora account of Australian Garcinia 
enumerated only three species (G. cherryi 
F.M.Bailey [Bailey 1902; subsequently 
transferred to Ternstroemia (Theaceae) as T. 


2 

cherryi (F.M.Bailey) J.F.Bailey & C.T.White], 
G. mestonii and G. warrenii ) (Bailey 1899). 
Additional species were added by Moore 
(1917) and White (1933, 1936) and names for 
Malesian taxa have been variously misapplied 
to some species; however, the genus has been 
little studied taxonomically in the intervening 
years to the present. Consequently, putatively 
undescribed species known only by phrase 
names have been recognised in the last twenty 
years and nine native species were listed for 
Australia by Cooper & Cooper (2004) and 
CHAH (2007). 

Examination of both live populations 
of plants and herbarium material, coupled 
with preliminary phylogenetic analyses of a 
limited DNA sequence dataset indicate that 
four undescribed species occur in Australia. 
Previously, Garcinia species endemic to 
Australia have been named in honour of 
worthy recipients and this revision continues 
the tradition by honouring four people 
who have made valuable contributions to 
Queensland botany. G. jensenii W.E.Cooper 
is described for material from Cape York 
Peninsula previously assigned to G. riparia 
A.C.Sm. at CNS. G. leggeae W.E.Cooper is 
described for all of the CNS and some of the 
BRI material of G. sp. (Claudie River) (the 
remaining BRI material is G. jensenii). G. 
zichii W.E.Cooper is described for specimens 
known as G. sp. (Davies Creek) and G. 
russellii had not been recognised by any 
herbaria but was listed in Cooper & Cooper 
(2004) as G. sp. (Mossman). 

Lineages, clades and associated 
morphology 

A phylogenetic analysis of DNA sequence data 
resolved two major lineages within Garcinia 
(Sweeney 2008). Australia is represented by 
12 species (nine native and three naturalised) 
distributed among 6 clades. Two species 
occur in Clades 1 & 2 within Lineage A, 
and ten species occur in Clades 4, 5, 6 & 9 
within Lineage B. Lineage A is distinguished 
by the presence of floral nectaries. Lineage B 
is distinguished in part, by the lack of disks, 
appendages and rings (nectaries) (Sweeney 
2010: 158). 


Austrobaileya 9(1): 1-29(2013) 

Sweeney (2008: 1299) in reference to 
Lineage B, stated that Clades 2 and 4 usually 
lack a well-developed pistillode. However, 
using Sweeney’s evidence (2008: 1292 and in 
lift .), Clade 2 is part of Lineage A and Clade 

4 does have a well-developed pistillode. It is 
Clade 7 and some species within Clades 5 and 
9, which lack a well-developed pistillode. 

Clade 1 (Garcinia dulcis (Roxb.) Kurz 
and G. xanthochymus Hoolc.f. [the latter 
naturalised in Queensland]) is distinguished 
by 5-merous flowers. Clade 2 (G. Iivingstonei 
[naturalised in Queensland]) is distinguished 
by free stamens. Clade 4 (G. jensenii and G. 
zichii ) is distinguished by fleshy bilocular 
flowers and fruit, which are capped with a 
disk-like stigma, unlike the woody pericarp 
described by Sweeney (2008). Clade 5 
(G.mestonii and G. leggeae ) is distinguished 
by an androecium of fused filaments creating 
a solitary fleshy receptacle that is not adnate 
to the petals. Sweeney (2008) stated an 
additional possible synapomorphy for Clade 

5 was the fruit, which usually have furrows 
along the septal radii, and for Clade 9, the 
presence of a distinctive adaxial, exudate- 
containing canal pattern in the leaves, 
consisting of highly branched, anastomosing 
canals. In Australian material; however, these 
characters poorly predict clade membership 
because furrows (or at least grooves) along 
the septal radii (putatively synapomorphic for 
Clade 5) occur in two species from Clade 9 
but not in the Clade 5 species. Furthermore, 
intricately branched exudate-containing 
canals (putatively synapomorphic for Clade 9: 
G. brassii C.T.White, G. gibbsiae, G. rnsselli 
and G. warrenii) are found in G. dulcis and 
G. Iivingstonei (Clades 1 & 2). Clade 6 (G. 
mangostana L.) has no clear morphological 
synamorphies but is usually distinguished 
by a fungiform pistillode surrounded by an 
undivided 4-lobed androecium (Sweeney 
2008). Clade 9 in Australian plants is 
distinguished by an androecium that is adnate 
to the petals. 

Materials and methods 

The study is based upon the examination of 
521 specimens or photos from BRI, CANB, 
CNS, K, MEL, NSW and P. All specimens 


Cooper, Garcinia in Australia 

cited have been seen by the author and all 
native species have been observed in the field. 
Naturalised species have been seen in the field 
and/or cultivation. 

Measurements of the floral parts and 
fruits are based on material preserved in 70% 
ethanol as well as fresh material from the 
field; d.b.h. refers to the forestry logging term 
of diameter at breast height. Abbreviations in 
the specimen citations are: EP (Experimental 
Plot), FR (Forest Reserve), LA (Logging 
Area), NP (National Park), SF/SFR (State 
Forest/State Forest Reserve) and TR (Timber 
Reserve). The superscript 1 refers to online 
images of specimens. 

Species are arranged sequentially 
following the disposition of taxa outlined by 
Sweeney (2008). 

Taxonomy 

Garcinia L. Sp. PI. 1: 443 (1753). Type 
species: G. mangostanaV. 

Rheedia L., Sp. PI. 1193 (1753). Type species: 
Rheedia laterifolia L. [=Garcinia humilis 
(Vahl) C.D.Adams], 

Cambogia L., Gen. PI. , ed. 5:225 (1754). 
Type species: Cambogia gummi-gutta L. 
[=Garcinia gummi-gutta (L.) N.Robson], 

Oxycarpus Lour., FI. Cochinch. 647 (1790). 
Type species: Oxycarpus cochinchinensis 
Lour. [=Garcinia cochinchinensis (Lour) 
Choisy], 

Verticillaria Ruiz & Pav., FI. Peruv. Prodr. 
81: t. 15. (1794). Type species: Verticillaria 
acuminata Ruiz & Pav. 

Xanthochymus Roxb., PI. Coromandel 2: 51 
(1798). Type species: Xanthochymuspictorius 
Roxb. [=Garcinia xanthochymus Hook.fi], 

Brindonia Thouars, Diet. Sci. Nat. 5: 339 
(1804). Type species: Brindonia indica 
Thouars. [=Garcinia indica (Thouars) 
Choisy], 

Stalagmitis Murray, Commentat. Soc. Regiae 
Sci. Gott. 9: 173 (1789). Type species: 
Stalagmitis cambogioides Murray \=Garcinia 
ovalifolia 01 iv.]. 


3 

Hebradendron Graham, Companion Bot. Mag. 
2: 199 (1837). Type species: Hebradendron 
cambogioides Graham [=Garcinia morella 
Desr.]. 

Discostigma Hassk., Flora 25(2, Beibl .): 33 
(1842). Type species: Discostigma rostrata 
Hassk. [=Garcinia rostratum (Hassk.) 
Hook.fi], 

Rhinostigma Miq., FI. Ned. Ind., Eerste. Bijv. 
3: 495 (1861). Type species: Rhinostigma 
parvifolia Miq. [=Garciniaparvifolia Miq.]. 

Pentaphalangium Warb., Bot. Jahrb. Syst. 13: 
382 (1891). Type species: Pentaphlangium 
crassinerve Warb. \=Garcinia crassinervis 
(Warb.) Kosterm.]. 

Garcinia in Australia: Dioecious or 
monoecious (agamospermous in the 
naturalised G. mangostana ), mostly subcanopy 
trees but also shrubs or canopy trees to 30 m; 
exudate, yellow, white or colourless, opaque 
or translucent. Branchlets square, ribbed 
or terete; petioles mostly channelled and 
ligulate. Interpetiolar gland-like bracts at 
nodes on stems and inflorescences, deltoid, 
peltate, tiny, caducous, brown. Leaves 
simple, opposite or rarely whorled, coriaceous 
or chartaceous, shape variable, glabrous; 
margin entire (or crenate in the naturalised 
G. livingstonei), mostly recurved; venation 
brochidodromous or paxillate; primary vein 
raised or flush; exudate-containing canals 
as parallel or random lines, branched or 
unbranched, continuous or broken dashes 
and dots. Inflorescences terminal, axillary or 
ramiflorous; bracts opposite, leaf-like with a 
primary vein, occurring at peduncle apex on 
racemose or paniculate species. Bracteoles 
mostly opposite, convex or cupular, occurring 
at pedicel base on most species. Flowers 
solitary or in cymes, compact panicles, 
racemes or fascicles, up to 66-flowered; 
actinomorphic, hypogynous, unisexual, may 
be fragrant, sessile or pedicellate; sepals 4 or 
5, free or joined; petals 4 or 5, free, convex, 
cupular, erect to almost explanate. Male 
flowers: androecium of fused stamens as 
phalanges, free stamens or fused filaments 
as a receptacle, pistillode may be present; 
anthers mostly bithecate, sessile or with a 


4 


short filament. Female flowers: staminodal 
phalanges present or absent; ovary superior, 
2-13-locular, 1-4 ovules per locule; stigma 
sessile or with a short style, discoid or lobed. 
Fruit a fleshy or dry berry, sometimes 
with seams or furrows along septal radii, 


Austrobaileya 9(1): 1-29 (2013) 

2-13-locular; seeds 1-13, 1-4 per locule; 
aril present in most species. Germination is 
hypogeal or epigeal. 

Etymology : The generic epithet Garcinia 
is named for the Dutch army doctor and 
naturalist Laurentius Garcin (1683-1751). 


Key to Garcinia species in Australia 


1 Stigma on flowers and fruit disk-like and entire.2 

1. Stigma on flowers and fruit rayed, lobed or crenate. 4 

2 Leaves always opposite; flowers with stamens united as phalanges and 

surrounding a fungiform pistillode; fruit red or blackish. 3 

2. Leaves mostly 3-whorled (occasionally opposite or 4-whorled); flowers 

with free stamens, pistillode absent; fruit yellow to orange. 3 . *G. livingstonei 

3 Fruit much longer than wide, occurs in the Wet Tropics area.5. G. zichii 

3 . Fruit equidimensional, occurs north from the Mcllwraith Range. 4 . G. jensenii 

4 Seeds and ovules usually more than one per locule.5 

4 . Seeds and ovules only one per locule.6 

5 Leaves peltate.11. G. russellii 

5. Leaves never peltate.10. G. gibbsiae 

6 Exudate in stems and leaves white; petals 5.7 

6. Exudate in stems and leaves yellow; petals 4.8 

7 Secondary veins 20-25 pairs.2. *G. xanthochymus 

7. Secondary veins 9-14 pairs.1. G. dulcis 

8 Leaves with 10 pairs of secondary veins; exudate-containing canals as 

short streaks, dashes and dots, unbranched.7. G. mestonii 

8. Leaves with 12 or more pairs of secondary veins; exudate- 

containing canals as continuous long lines.9 

9 Leaves with 2 distinct intramarginal veins.8. *G. mangostana 

9. Leaves with 1 intramarginal vein or intramarginal vein absent or not obvious 10 

10 Leaves almost membranous; petals up to 3 mm long.6. G. leggeae 

10. Leaves coriaceous; petals 5 mm or more long.11 

11 Leaves with tertiary veins absent.9. G. brassii 

11. Leaves with tertiary veins present.12 

12 At least some or most tertiary veins diminishing and not anastomosing . . . . 9. G. brassii 

12. All tertiary veins anastomosing.12. G. warrenii 


1. Garcinia dulcis (Roxb.) Kurz, J. Asiat. 
Soc. Bengal, Pt. 2, Nat. Hist. 43: 88 (1874); 
Xanthochymus dulcis Roxb., PI. Coromandel 
3: 66 (1820). Type: “ex Molucca Islands, cult. 
In Indian Botanic Garden, Calcutta” n.v.,fide 
Maheshwari (1964: 116). 


Illustrations : Hooker (1831: t. 3088); Wight 
(1840: 1: t. 192); van Nooten (1880: t. 15); 
Christophel & Hyland (1993: 81); Cooper & 
Cooper (2004: 123); CANBR (2010); Hyland 
etal. (2010); Sompong (2007: 248-250). 


5 


Cooper, Garcinia in Australia 

Tree to 20 m, dioecious; exudate white in 
stems and petioles, yellow in fruit; bark with 
fine vertical fissures, becoming thin flakey 
on older trees; branchlets square, ribbed, 
pendulous, glabrous. Leaves discolorous, 
coriaceous, glabrous; petioles 10-30 mm 
long, channelled, ligulate; lamina elliptic, 
ovate or oblong-ovate, 75-280 mm long, 
32-180 mm wide; base attenuate, cuneate, 
obtuse, cordate or asymmetrical; apex shortly 
acuminate, acute or obtuse; margin recurved; 
venation brochidodromous, primary vein 
± flush on upper surface and protruding on 
abaxial surface; secondary veins 9-14 pairs, 
raised on both sides in dried specimens, angle 
of divergence from primary vein 30-60°, 
forming loops within 1.5-5 mm from margin; 
exudate-containing canals on upper side 
are numerous many-branched wriggly lines 
running more or less parallel to secondary 
veins, on the underside they are numerous 
parallel branched or unbranched lines 
which run at angles between being parallel 
to primary vein and margin, sometimes 
running parallel to secondary veins. Male 
and female inflorescences similar, axillary 
or ramiflorous, a raceme or a fascicle on 
a pulvinus, 1-many-flowered; bracteoles 
persistent, clustered at base of pedicels, 
ovate, convex, keeled, 1-2.25 mm long, 
1-2 mm wide, base truncate, apex obtuse 
or acute, entire, minutely hairy especially 
around margin, sparsely hairy abaxially. 
Male flowers: fragrant, diameter 11-12 mm, 
glabrous; pedicels 7-14 mm long; sepals 5, 
free, orbicular, 2-4 mm long, 3-4 mm long, 
green, white or cream, margin minutely 
ciliate; petals 5, free, erect, imbricate, 
orbicular, convex, 5-7 mm long, 5-7 mm 
wide, membranous, green, lime-yellow, cream 
or yellow, glabrous, margin thin and minutely 
ciliate; androecium 5-phalangiate, glabrous; 
phalanges antepetalous, strap-shaped at 
base and flabellate towards apex, introrse, 
8-9 mm long, 0.8-1.4 mm wide, apically 
branched into 5-10 filaments, filaments 0.3- 
0.7 mm long, 1 anther per filament; anthers 
bithecate, introrse, broadly-globose, c. 0.2 
mm long and 0.3 mm wide; disk diameter 
c. 4 mm, pitted or honeycombed; pistillode 
rudimentary, conical-attenuate c. 0.5 mm 


long. Female flowers: fragrant, diameter 
8-9 mm; pedicels 9-17 mm long; sepals 5, 
free, imbricate, orbicular, convex, 3-4.5 mm 
long, 4.5-6.5 mm wide, green or yellowish, 
glabrous, margin minutely ciliate; petals 5, 
free, imbricate, orbicular, convex, 7-8 mm 
long, 7-8 mm wide, green, cream or yellow, 
glabrous, margin minutely ciliate; staminodal 
phalanges 5, antepetalous, solitary and 
filament-like or 2 or 3-fused, distally 
indexed, 2-3.5 mm long, 2 or 3-branched 
at apex, filaments 0.9-1.4 mm; antherodes 
1-3 per phalange, introrse, bilobed, flattened, 
0.2-0.3 mm long, c. 0.3 mm wide; nectaries 
alternating between phalanges, c. 2 mm long 
and 2 mm wide, pitted; disk c. 4 mm wide; 
ovary globular, c. 5 mm long and 4 mm wide, 
sessile, 5-locular, locules uniovulate; stigma 
5-branched, style c. 1.8 mm long; stylar arms 
5, c. 2.5 mm long. Fruiting pedicel c. 15 
mm long, 5 sepals persistent at base; fruit 
a fleshy berry, ovoid, globular or depressed 
globular, apex often beaked, 35-50 mm long, 
45-53 mm wide, yellow, glabrous; stigma 
5-rayed, diameter 2-3 mm; seeds usually 5, 
c. 30 mm long and 20 mm wide; testa brown, 
smooth; aril fleshy, cream or yellow. Claudie 
mangosteen, gourka, mundu tree. Fig. 1A & 
B, 2B. 

Additional selected specimens (from 47 examined): 
Malaysia. Kelantan: Pasir Putih, Semarak Bt., Sep 
1992, FRI37081 (CNS). Papua New Guinea. West New 
Britain District: 32 km SW of Linga Linga Plantation, 
May 1973, Isles & Katik NGF32248 (CNS). Milne Bay 
Province: Esirava Creek, Tagula Island, Mar 1979, 
Damas LAE74527 (CNS). Australia. Queensland. 
Cook District: Banks Peak, Moa Island, Torres Strait, 
Jul 2008, Fell 9722 & Stanton (BRI); Moa (Banks) 
Peak, Moa Island, Torres Strait, Feb 1989, Gray 4993 
(CNS); Escarpment of Great Dividing Range, 14.6 km 
NE of Heathlands Ranger Base, Heathlands D & O 
Reserve, Oct 1993, Fell 3746 & Stanton (BRI); CSIRO 
Plot, West Claudie River, Dec 2011, Cooper 2216 & 
Venables (CNS); Gordon Creek, Oct 1972, Hyland 6416 
(CNS); Claudie River, Oct 1981, Hyland 11181 (CNS); 
Iron Range, base of Lamond Hill, Sep 1988, Sankowsky 
861 (CNS); Nesbit River, Silver Plains Holding, Oct 
1999, Hyland 16253 (CNS); Nesbit River, Silver Plains 
Holding, Oct 1999, Hyland 16254 (CNS); Junction of Leo 
Creek and Nesbit River, Aug 1993, Fell 3385, Jensen & 
Barnes (BRI); Chester River Scrub, Jun 1992, Forster 
PIF10440 & Tucker (BRI); TR 14, Leo Creek Road, Dec 
1979, Hyland 10132 (CNS); TR 14, Leo Creek Mine area, 
Mcllwraith Range, Jun 1992, Forster P1F10236, Tucker 
& Kenning (BRI); Rocky River crossing, Oct 1996, 
Cooper 1016 & Cooper (CNS); Rocky River catchment. 


A ustrobaileya 9(1): 1-29(2013) 


Fig. 1. Flowers. A. Garcinia dulcis showing rayed stigma at ovary apex {Hyland 16253 [CNS]). B. G. dulcis S, 
showing apically branched phalanges {Hyland 16254 [CNS]). C. G. mestonii S, showing small opening and anthers 
{Sankowsky 2326 [BRI]). D. G. jensenii showing 3-flowered inflorescence and yellow stigma at apex of fungiform 
gynoecium {Cooper 2225, Zdenek, Venables & Pritchard [CNS]). E. G. leggeae S showing anthers {Cooper 2175, 
Jensen & Kemp [CNS]). F. G. mestonii showing small opening and stigma {Forster PIF24028 [CNS]). G. G. 
zichii % showing viscous fungiform gynoecium {Sankowsky 2547 [BRI]). H. G. zichii 3 , showing viscous fungiform 
pistillode and anthers at apices of 4-phalangiate androecium {Dowe 1159 [CNS]). Photos: A, B, F, CSIRO; C, G, 
G.Sankowsky, D, B. Venables, E, R. Jensen, H, J.Dowe. 


Cooper, Garcinia in Australia 


7 


Fig. 2. Exudate-containing canals on leaf undersides shown crossing secondary venation. A. Garcinia brassii 
showing as pale lines {Cooper 2163 & Ford [CNS]). B. G. dulcis showing as pale lines and reticulations {Cooper 
2216 & Venables [CNS]). C. G.jensenii showing as dark lines {Cooper 2194, Jensen & Kemp [CNS]). D. G. mestonii 
showing as pale wriggly dashes and dots {Cooper 2203 & Jensen [CNS]). E. G. gibbsiae showing as pale lines {Cooper 
2212 [CNS]). F. G. leggeae showing as dark lines {Cooper 2187, Jensen & Kemp [CNS]). G. G. warrenii showing as 
dark lines {Cooper 474 [CNS]). H. G. livingstonei showing as dark lines {Cooper 2218 [CNS]). Photos: A, B, C, D, E, 
G, H W.Cooper ; F, R. Jensen. 


8 

Silver Plains, Aug 1997, Cooper & Jensen 44 (CNS); 
Upper Massey (Massy) Creek, Oct 1962, Smith 11719 
(BRI); Melville Range 4 km N of Abbey Peak, Nov 1981, 
Irvine 2195 (CNS). 

Distribution and habitat : Garcinia dulcis is 
widespread from Australia and New Guinea to 
South East Asia and India. Within Australia, 
it is restricted to north Queensland where it 
occurs in rainforest (complex, semideciduous 
notophyll and mesophyll vine forest) from the 
Melville Range on Cape York Peninsula to the 
Torres Strait islands, altitude 10-500 m (Map 
1 ) 

Phenology : Flowers have been recorded 
from September to February; fruit has been 
recorded from August to October and January 
to March. 

Etymology : The specific epithet is derived 
from the Latin dulcis (sweet or pleasant) 
referring to the sour but palatable aril. 

2. *Garcinia xanthochymus Hook.fi ex 
T.Anders., FI. Brit. India 1: 269 (1874); 
Xanthochymus pictorius Roxb., PI. 
Coromandel 2: 51, t. 196 (1805); Stalagmitis 
pictoria (Roxb.) G.Don., Gen. Syst. 1: 620 
(1831); Xanthochymus tinctorius DC., Prodr. 
1: 562 (1824) [sphalm. for pictorius] ; Garcinia 
tinctoria (DC.) W.F.Wight, Bull. Bur. PI. 
Industr. U.S.D.A. 137: 50 (1909). Type: India: 
Circars, s.dat., Roxburgh s.n. (holo: BR, herb. 
Mart.). 

Illustrations : Roxburgh (1805, 2: t. 196) as 
Xanthochymus pictorius Roxb.; Kirtikar & 
Basu (1918, 1: t. 104); Sweeney (2008: 1289, 
K; 2010: 158, 160, 162); Sharma et al. (2012: 
8,9). 

Shrub or tree to 20 m, dioecious; exudate 
yellow or milky; bark dark brown, with 
short vertical fissures, flaky on older trees; 
branchlets square, mostly pendulous, twigs 
distinctly 6-8-ribbed. Leaves discolorous, 
coriaceous, glabrous, shiny; petioles 10-25 
mm long, ligulate; lamina oblong, linear- 
oblong, oblong-lanceolate, oblong-ovate 
or ovate, 120-410 mm long, 38-120 mm 
wide, base cuneate or rounded; apex acute, 
acuminate or apiculate; margin recurved; 
venation brochidodromous, primary vein 
slightly raised on upper side and distinctly 


Austrobaileya%\). 1-29(2013) 

raised on underside; secondary veins 20-25 
pairs raised on both sides, angle of divergence 
from primary vein 50-60°, forming loops 
within margin, intramarginal vein 0.5-5 
mm from margin; exudate-containing 
canals sparse on the underside as faint long 
lines running at angles between parallel 
to the margin and the primary vein. Male 
flowers: not seen for Australian material, 
recorded as: diameter 15 mm, fascicles 
4-8-flowered; sepals 5; petals 5, diameter c. 
8 mm; androecium 5-phalangiate; phalanges 
strap-shaped at base and flabellate towards 
apex, indexed, apically branched into 3-6 
filaments; anthers 2-celled, introrse; nectaries 
in between phalanges. Female inflorescences 
axillary or ramiflorous, a reduced raceme, 
fascicle-like, 2-14-flowered. Female flowers: 
fragrance not recorded; diameter 10—12(— 
20) mm; pedicels (18-)24-36 mm long; 
sepals 5, free, suborbicular, convex, 3.5-4 
mm long, 6-7 mm wide, green or whitish, 
glabrous, margin minutely ciliate; petals 
5, free, orbicular, convex, erect, 7.5-8 mm 
long, 10-11 mm wide, white, cream or pale 
yellow, margin minutely ciliate; staminodal 
phalanges 5, antepetalous, solitary and 
slender or 3 or 4-fused and strap-shaped at 
base and flabellate towards apex, indexed, c. 

3 mm long, branched into 3 or 4 filaments at 
apex, filaments 0.5-0.7 mm long, antherodes 
solitary or 3 or 4 per phalange, introrse, 
bilobed, flattened, c. 0.3 mm long and 0.5 
mm wide; nectaries alternating between 
phalanges, c. 2 mm long and 2.5 mm wide, 
pitted; disk c. 5 mm wide; ovary globular, c. 

4 mm long and 6 mm wide, sessile, 5-locular, 
locules uniovulate; stigma 5-branched, style 
c. 1.7 mm long; stylar arms 5, 2.5-3 mm 
long. Fruiting pedicel to 40 mm long, sepals 
persistent at base; fruit a fleshy berry, often 
asymmetrical with beak not centred, ovoid 
or globose, beaked, 38-65 mm long, 35-50 
mm wide, yellow or orange; stigma 5-rayed, 
diameter 4-5 mm; seeds 1-5 (usually 1 or 
2), 25-35 mm long, 15-18 mm wide, testa 
brown, veined; aril fleshy, yellow or orange. 
sour mangosteen, false mangosteen , yellow 
mangosteen, gamboge tree , eggtree. 

Additional selected specimens (from 6 examined): 
Australia. Queensland. Cook District: Tributary of 


Cooper, Garcinia in Australia 

Bushy Creek, Rumula, Oct 1998, Holmes 66 (CNS); 
Cultivated, Cairns Botanic Gardens, Feb 2013, Cooper 
2217, Cooper & Venables (CNS); Wadda Creek, East 
Palmerston, Sep 1941, Wilson s.n. (CNS); Patrick Road 
off Dunne Road, East Palmerston, Dec 2005, Jensen 
1486 & Cauley (BRI); Tully, Dec 1945, Shearer s.n. 
(CNS). North Kennedy District: Delta, 3 miles [5 km] 
W of Bowen, Nov 1971, Wright s.n. (BRI [AQ313]). 
Moreton District: Rosemount Road, Nambour, Nov 
1989, Brufords.n. (BRI [AQ458316]). 

Distribution and habitat : Garcinia 
xanthochymus is a native of Bangladesh, 
Bhutan, Cambodia, China, India, Laos, 
Myanmar, Nepal, Thailand and Vietnam. It 
is cultivated for its edible sour fruit and has 
become naturalised in a few locations in 
Queensland from the Rumula area south to 
Nambour (Map 6), altitude 20-500 m. 

Phenology : Flowers have been recorded from 
February to May. Fruit has been recorded from 
February to May, November and December. 

Typification : Garcinia xanthochymus was 
named by Anderson in Hooker’s Flora of 
British India (1874); however, nomenclature 
for this species commenced nearly 70 years 
earlier when Roxburgh (1805) described 
Xanthochymus pictorius Roxb. (non Garcinia 
pictoria Roxb.). De Candolle (1824) did 
not help the situation by totally misprinting 
(or perhaps ‘correcting’) the epithet as X. 
tinctorius DC. (Maheshwari 1964; Singh 
1993). Anderson (1874) interpreted de 
Candolle’s (1824) account as a misspelling, so 
coined the new name G. xanthochymus with 
Roxburgh’s X. pictorius and de Candolle’s X. 
tinctorius both in synonymy. 

Much later, D’Arcy (1980) overlooked the 
name Garcinia pictoria Roxb. and published 
the superfluous new combination G. pictorius 
(Roxb.) D’Arcy based on Xanthochymus 
pictorius Roxb. 

Anderson (1874) did not unambiguously 
specify a type collection for Garcinia 
xanthochymus , but did list X. pictorius 
Roxb. and the misprinted X. tinctorius DC. 
in synonymy, together with references to 
other literature and one specimen in the 
Wallich Catalogue (no. 4837). Under the 
distribution notes for this species Anderson 
(1874) cited a single specimen collected by 
Roxburgh from the Circars. Maheshwari 


9 

(1964) listed the type for G. xanthochymus 
as this Roxburgh collection (deposited in 
BR) and this interpretation appears to have 
been universally accepted by later authors 
(Kostermans 1980; Singh 1993). 

Notes : Sharma et al. (2012) suggested that 
G. xanthochymus produces bisexual flowers; 
however they concluded that functional 
bisexuality had not been confirmed. 
Agamospermy is known or suspected for 
ten Garcinia species (not including G. 
xanthochymus) (Richards 1990a). Separate 
male and female flowers have been observed 
and illustrated by Sweeney (2010). 

Etymology : The specific epithet is derived 
from the Greek xanthos (yellow) and chymos 
(sap); referring to the exudate. 

3. *Garcinia livingstonei T.Anders., Bot. J. 
Linn. Soc. 9: 263 (1866). Type: cult. In hort. 
bot. Calcutta, ex Mozambique, “in rupibus 
schistosis prope flumen Zambesi”, s.dat., Kirk 
s.n. (holo: CAL);, fide Robson (1960: 400). 

Illustrations : Sweeney (2008: 1289, Q). 

Shrub or tree to 18 m, monoecious; exudate 
yellow or reddish; bark grey and rough with 
numerous vertical fissures; branches terete, 
arching. Leaves discolorous, opposite or 
3-4-whorled, coriaceous, glabrous; petioles 
2-7 mm long, channelled, ligulate; lamina 
oblong, elliptical or oblanceolate, 35-110 
mm long, 20-55 mm wide; base cuneate, 
rounded or cordate; apex acute, rounded, 
emarginate or apiculate; margin crenate or 
entire, recurved; venation brochidodromous, 
primary vein raised in a groove adaxially 
and distinctly raised abaxially; secondary 
veins 9-15 pairs, pale yellow, raised on both 
surfaces, angle of divergence from primary 
vein 60-70°; intramarginal vein merges 
with margin; exudate-containing canals on 
upperside are numerous branched lines ± 
parallel to secondary veins, on the lower side 
they are branched lines running at angles 
between the primary vein and margin. Male 
and female inflorescences: similar, axillary 
or ramiflorous, a fascicle, 2-7-flowered; 
bracteoles ± quadrate or rhombic, slightly 
convex, 1-1.5 mm long, 1-1.5 mm wide, 
base truncate, apex acute, margin 3-toothed, 


10 


glabrous. Male flowers: fragrant, diameter 
8-10 mm; pedicels 6.5-8 mm long, softly 
muricate; sepals 4,2-whorled, free, D-shaped, 
c. 1 mm long and 1.8 mm wide, green, margin 
entire; inner pair orbicular, convex, c. 2 mm 
long and 2.5 mm wide, green or yellowish- 
green, margin entire; petals 5, free, orbicular, 
reflexed, 4.5-5.5 mm long, 3.5-4.5 mm wide, 
membranous, green or greenish-yellow, 
glabrous; androecium a globose yellow disk 
with c. 30 free stamens radiating like needles 
in a pin cushion, disk sunken directly below 
each stamen, filaments 1.5-1.7 mm long; 
anthers bithecate, globose, diameter c. 0.3 mm; 
pistillode absent. Female flowers: fragrant, 
diameter c. 7 mm; pedicels 16-20 mm long, 
softly muricate; sepals 4, free, green; outer 
pair D-shaped, c. 1.2 mm long and 1.4 mm 
wide; inner pair ovate, c. 2.5 mm long and 2 
mm wide, glabrous; petals 5, free, orbicular 
or obovoid, c. 8 mm long and 6 mm wide, 
green or greenish-yellow; disk oblate, yellow, 
with c. 26 free staminodes, diameter c. 3.5 
mm, sunken directly below each staminode; 
staminodes c. 2 mm long; antherodes 1 per 
staminode, ± globose, bilobed, diameter c. 
0.5 mm; ovary obovoid, c. 3 mm long and 
3 mm wide, 2-locular, locules uniovulate; 
stigma domed, not rayed. Fruiting pedicel: 
not seen for Australian material, recorded as: 
at least as long as fruit, staminodes usually 
persistent at base, sepals absent at base; fruit 
(not seen for Australian material) a fleshy 
berry, globular or obovoid, 10-30 mm long, 
10-30 mm wide, orange or reddish-orange; 
seeds 1 or 2, cylindrical, 8-20 mm long; aril 
fleshy, orange. African mangosteen , lowveld 
mangosteen, imbe. Fig. 2H. 

Additional selected specimens (from 54 examined): 
Senegal. Ouassadou, Jan 1959, Berhaut 4302 (P 1 ). 
Mali. Bankoumana, Nov 2005, Birnbaum 920 (P 1 ); 
Tienfala (25 km E Bamako), Dec 1978 (P 1 ). Cameroon. 
Southwest Province: Mt Kupe, Feb 1986, Thomas & 
McLeod 5480 (P 1 ). Comoros. Mayotte, Sohoa, May 
1997, Pascal 934 (P 1 ); Itsamia, Nov 1999, Labat 3223 
& Yahaya, Dloubeiri & Mindhiri (P 1 ). Kenya. Sep 
1957, Verdcourt 1936 (P 1 ). Tanzania. 40 km west of 
Lindi, Sep 1934, Schlieben 5402 (P 1 ); Zanzibar, Nov 
1889, Sacleux 886 (P 1 ); Pwani Coast Region, Matumbi 
Hills, Oct 1997, Phillipson 4960, Sitoni & Kibure (P 1 ). 
Zimbabwe. Urungwe District, Oct 1964, Bingham 1370 
(P 1 ). Mozambique. Gaza, Nov 1970, Correia 2015 (P 1 ); 
Lugela-Mucuba, Namagoa, s.dat., Faulkner Kew 77 (P 1 ). 
Australia. Queensland. Cook District: Cooktown, 


Austrobaileya 9(1): 1-29 (2013) 

Anzac Memorial Park, Sep 2008, McKenna 267 (CNS); 
Helen St, Cooktown, May 2013, Cooper 2218 & Cooper 
(CNS); Trevethan Creek, Cooktown Development Road, 
May 1998, Jago s.n. & Wannan (CNS). 

Distribution and habitat: Garcinia 
livingstonei is a native of Africa with a 
widespread distribution, occurring on the 
mainland from West Africa (including 
Senegal, Mali, Nigeria and Cameroon) 
through to Somalia and East Africa (including 
Kenya and Tanzania) and south to Zimbabwe, 
Angola and South Africa; as well as the 
Comoros islands. In Australia it has been 
cultivated and a recent record ( Jago s.n. & 
Wannan CNS) from Trevethan Creek near 
Cooktown indicates possible naturalization 
(Map 1) 

Phenology: Flowers from Australian material 
have been recorded in May and no fruiting 
records exist. 

Notes: Cultivated plants of Garcinia 
livingstonei in Cooktown, which had been 
poisoned, were sprouting vigorous suckers 
from the base of otherwise dead trees. 

One tree at Cooktown {Cooper 2218 ) had 
both female and male flowers confirming that 
this species is monoecious. 

Etymology: The specific epithet is in honour 
of the explorer David Livingstone (1813— 
1873), who was amongst the first to collect 
this species. 

4. Garcinia jensenii W.E.Cooper sp. nov. 
Similar to Garcinia hunsteinii Lauterb. but 
differs in the leaf apex (bluntly acute versus 
acuminate), secondary veins (7-10 pairs 
versus c. 50 pairs), and secondary vein angle 
(c. 45° versus c. 80°). Typus: Australia: 
Queensland. Cook District: about 3 km N 
of 3-ways, Iron Range National Park, 29 July 
2013, W.Cooper 2225, C.Zdenek, B.Venables 
& J.Pritchard (holo: CNS [2 sheets + spirit], 
iso: 13 sheets to be distributed to A, BRI, 
CANB, FRI, K, KEP, KL, MEL, MO, NSW, 
NY, SING, US). 

Garcinia riparia auct. non A.C.Sm.; Hyland 
etal. (2003, 2010); CHAH (2007). 

Garcinia sp. (Claudie River L.J.Brass 19658) 
in part; Jessup (2007: 54, 2010: 44). 


Cooper, Garcinia in Australia 

Illustrations : Christophel & Hyland (1993: 
81) as G. riparia vel aff; Hyland et al. (2003, 
2010), CHAH (2007) as G. riparia; Zodiac 
Publications (undated), as Garcinia sp. 
Claudie River. 

Tree to 15 m, dioecious; exudate white or 
colourless; dbh to 150 mm; bark slightly 
flakey with fine vertical fissures, lenticels 
horizontal and more prevalent on younger 
trees; branchlets square, young twigs 
narrowly 4-winged. Leaves discolorous, 
coriaceous, glabrous; petioles 5-7 mm long, 
channelled, ligulate; lamina narrowly to 
broadly elliptic, ovate-elliptic or obovate, 
52-160 mm long, 20-72 mm wide; base 
cuneate, attenuate or decurrent; apex acute, 
retuse-acute or rounded-acute; margin entire; 
venation brochidodromous, primary vein on 
upperside flush or slightly raised, distinctly 
raised on underside; secondary veins 7-10 
pairs, slightly raised on both surfaces, angle of 
divergence with primary vein c. 45°, forming 
intramarginal loops 0.4-1.4 mm from margin; 
exudate-containing canals numerous and 
conspicuous on the underside as unbranched 
lines ± parallel to each other and running at 
angles between primary vein and margin. 
Male inflorescences axillary, a fascicle or 
raceme, 5-many-flowered, rachis up to 3.5 
mm long; bracts at pedicel bases broadly 
ovate, apex acute, base truncate, convex, c. 1.6 
mm long and 1 mm wide, glabrous; bracteoles 
present or absent, part way along pedicels, 
solitary or paired, broadly ovate, convex, c. 
1 mm long and 1 mm wide, glabrous. Male 
flowers: fragrance not recorded, diameter 
7-8 mm; pedicels 4.5-12 mm long; sepals 4, 
2-whorled, colour not recorded but probably 
white or green white like females, glabrous; 
outer pair joined, D-shaped, convex, may 
be keeled, c. 1.6 mm long and 1.8 mm wide; 
inner pair free, petal-like, orbicular, convex, 
c. 4 mm long and 3 mm wide; petals 4, free, 
imbricate, orbicular, convex, c. 4 mm long 
and 3-4 mm wide, membranous, colour not 
recorded but probably white like females, 
glabrous; androecium 4-phalangiate, 
glabrous; phalanges antepetalous, strap¬ 
shaped, slightly ribbed, erect or distally 
indexed, c. 3.2 mm long and 2-3 mm wide, 
apex divided into 2-7 short filaments; 


ll 

anthers 15-30 per phalange, completely 
enveloping phalange apex, D-shaped, circular 
or irregular, margin lobed, erect or introrse, 
diameter 0.5-6 mm; pistillode fungiform, 
c. 3 mm long and 2 mm wide at apex, apex 
colour not recorded but probably yellow like 
females. Female inflorescences terminal 
or axillary, supra-axillary or ramiflorous, a 
solitary flower, fascicle or condensed raceme, 
3-9-flowered; peduncles 1.2-2 mm long; 
bracteoles triangular, keeled, c. 1 mm long 
and 1 mm wide, entire, persistent. Female 
flowers: fragrance not detected, diameter 
4.5-6.5 mm; pedicels 3.5-4.5 mm long; 
sepals 4, 2-whorled, D-shaped, convex, white 
or palest green, glabrous; outer pair joined, c. 

1 mm long and 2 mm wide, coriaceous; inner 
pair free, petal-like, orbicular, convex, 1.2-2 
mm long and 2-3 mm wide, membranous; 
petals 4, free, imbricate, orbicular, convex, 
c. 2 mm long, 2-3 mm wide, margin thin, 
white, glabrous; gynoecium fungiform, 
unlobed, glabrous; ovary obovoid and 
laterally compressed, c. 1.2 mm long and 1.8 
mm wide, 2-locular, uniovulate; stigma disc¬ 
like, entire, diameter c. 2.5 mm, yellow, not 
viscous. Fruiting pedicel c. 5 mm long; 2 
joined sepals persistent at base; fruit a fleshy 
berry, globose or ellipsoid-globose, 12.5-15.5 
mm long, 11-16 mm wide, pink or reddish; 
stigma at apex, diameter 3-3.8 mm; seeds 1 
or 2, c. 10 mm long and 5 mm wide; testa and 
aril not recorded. Fig. ID, 2C. 

Additional selected specimens (from 23 examined): 
Australia. Queensland. Cook District: Widul Island, 
4 km west of Mabuyang Island Airfield, 73 km NNW of 
Horn Island Airfield, Apr 2009, Fell 10037 (CNS); Ulu 
(Saddle Islet), 62 km NE of Horn Island Airfield, Torres 
Strait, Feb 2012, Fell 10712 (CNS); Eet Hill Track, Moa 
Island, Torres Strait, May 1987, Budworth 1185 (BRI); c. 

2 km N of St Paul’s community, Moa Island, 46 km NNE 
of Horn Island Airfield, Torres Strait, Nov 2007, Fell 
8955 (CNS); Mew River, Muddy Bay, Cape York, Jun 
1994, Forster PIF15300 & Tucker (BRI); Lake Boronto - 
Newcastle Bay, Cape York Peninsula, Sep 1974, Tracey 
14349 (BRI); Between Lockerbie & Somerset, Feb 1980, 
Hyland RFK3984 (CNS); Lockerbie Scrub, Sep 1974, 
Tracey 14306 (BRI); 5.5 km NE of Moreton, Jun 1994, 
Fell 4457 & fiMdt(BRI); Olive River, N bank, Sep 1974, 
Traceyl4373 (BRI); Nelson Creek, Aug 2011, Cooper 
2145, Venables & Murphy (CNS); Iron Range NP, 
southside of Pascoe River, Sep 2004, Sankowsky 2517 & 
Sankowsky (BRI); about 3 km N of 3-ways, Iron Range 
NP, Dec 2012, Cooper 2194, Jensen & Kemp (CNS); 
Iron Range, Jul 1923, Brass 19658 (BRI); about 2.8 km 


12 

N of 3-ways Iron Range NP, Jun 2013, Cooper 2219, 
Cooper & Venables (CNS); Chester River, Jun 1977, 
Hyland RFK3555 (CNS); TR 14, Leo Creek Mine area, 
Mcllwraith Range, Jun 1992, Forster PIF10160, Tucker 
& Kenning (BRI); Mcllwraith Range, E foothills. Rocky 
River, Oct 1969, Webb 9345 & Tracey (BRI). 

Distribution and habitat : Garcinia jensenii 
grows on red soils derived from a mixture 
of basic igneous rocks and ferrugineous 
sandstone. It is as an understory tree in 
rainforest (semideciduous notophyll/ 
mesophyll vine forest) from the Torres Strait 
islands and down the eastern side of Cape 
York Peninsula to the Mcllwraith Range at 
altitudes between sea level and 500 m (Map 
2 ). 

Garcinia jensenii occurs in rainforest 
dominated by Archidendron hirsutum 
I. C.Nielsen, Buchanania arborescens 
(Blume) Blume, Calamus caryotoides 

A. Cunn. ex Mart., Dillenia alata (R.Br. 
ex DC.) Martelli, Euonymus australiana 
F.Muell., Hydriastele costata F.M.Bailey, 
Maranthes corymbosa Blume, Planchonella 
xylocarpa C.T.White, Ptychosperma elegans 
(R.Br.) Blume, Syzygium bamagense 

B. Hyland & Syzygium pseudofastigiatum 
B. Hyland. 

Phenology : Flowers have been recorded in 
July and August; fruits have been recorded in 
December and February. 

Affinities : Within Australia, Garcinia jensenii 
is most similar to G. zichii W.E.Cooper. Both 
species have similarly shaped leaves and 
exudate canal pattern, a 2-locular ovary and 
1 or 2-seeded fruit. G. jensenii has leaves 
with 7-10 pairs of secondary veins, flowers 
with white petals and fruit that are globose 
or ellipsoid-globose. G. zichii has leaves 
with 12—18 pairs of secondary veins, flowers 
with green or yellow petals and fruit that are 
distinctly ellipsoid. 

Notes: In Queensland, Garcinia jensenii 
has been confused with G. riparia that is 
considered to be endemic to New Guinea; 
however, the two species are significantly 
different morphologically. G. jensenii 
has white or colourless exudate, flowers 
pedicellate, ovary 1 or 2-locular, fruit not 
ribbed and 1 or 2-seeded. G. riparia has yellow 


Austrobaileya 9(1): 1-29 (2013) 

exudate, flowers sessile with a 7-10-locular 
ovary, fruit ribbed and many-seeded. 

Etymology: The specific epithet honours 
botanist Rigel Jensen (1959-) acknowledging 
his contribution to botany in Australian and 
New Guinea, as well as his enthusiastic 
assistance with all of my botanical projects. 

5. Garcinia zichii W.E.Cooper sp. nov. 
Similar to Garcinia hunsteinii but differs in 
the leaf apex (acute versus acuminate), lateral 
veins (7-10 pairs versus c. 50 pairs), lateral 
vein angle (c. 45° versus c. 80°) and the fruit 
shape (ellipsoid versus globose). Typus: 
Australia: Queensland. North Kennedy 
District: Tully Falls National Park, 2.2 km 
along Tully Falls Road from Charmillan 
Creek bridge, 8 November 2011, A. Ford 5908 
& M. Torello-Raventos (holo: CNS [2 sheets + 
spirit], iso: 10 sheets to be distributed to BRI, 
CANB, K, KL, L, MEL, MO, NSW, SING). 

Garcinia sp. aff. G. hunsteinii ; Hyland 
(1982); http://www.anbg.gov.au/cgi-bin/apni 
Accessed January 2013. 

Garcinia sp. (Davies Creek JGT 14745); 
Hyland etal. (2003). 

Garcinia sp. (Davies Creek); Cooper & 
Cooper (2004: 124). 

Garcinia sp. (Davies Creek J.G.Tracey 14745); 
Jessup (2007: 49, 2010: 44). 

Garcinia sp. Davies Creek (J.G.Tracey 14745); 
CHAH (2007); Hyland et al. (2010). 

Garcinia sp. (=AFO/1268); http://www.anbg. 
gov.au/cgi-bin/apni Accessed January 2013. 

Illustrations: Christophel & Hyland (1993: 
81) as G. sp. aff. G. hunsteinii ; Hyland et al. 
(2003) as G. sp. (Davies Creek JGT 14745); 
Cooper & Cooper (2004: 124) as G. sp. 
(Davies Creek); Hyland et al. (2010), as G. sp. 
Davies Creek (J.G.Tracey 14745). 

Small tree or canopy tree to 30 m, dioecious; 
exudate colourless, white or slightly milky, 
meager; dbh to 20 cm, may have small 
buttresses; bark rough, fissured, slightly 
flaky, lenticels pale orange; branchlets square, 
deeply grooved between last 2 pairs of 
leaves, becoming terete. Leaves discolorous, 


Cooper, Garcinia in Australia 

coriaceous, glabrous; petioles 5-10 mm long, 
channelled, ligulate; lamina elliptical or rarely 
obovate, 35-110 mm long, 15-35 mm wide; 
base attenuate, cuneate or decurrent; apex 
bluntly acute; margin flat or may be recurved; 
venation brochidodromous, primary vein 
distinctly raised on upperside, slightly raised 
near base on underside and flush or slightly 
depressed towards apex; secondary veins 
obscure, 12-18 pairs, flush on both sides, 
angle of divergence with primary vein c. 
60°, forming intramarginal loops c. 0.5 mm 
from margin; exudate-containing canals 
numerous and conspicuous on the underside 
as unbranched lines ± parallel to each other 
and running at angles between primary vein 
and margin. Male inflorescences axillary 
or rarely terminal, an umbel or rarely a 
cyme, 2-11-flowered (mostly 3-5-flowered), 
peduncles 0.8-2.5 mm long; bracts at 
peduncle bases and bracteoles at pedicel 
bases similar, triangular or broadly rhomboid, 
convex, keeled, 0.8-2.6 mm long, muricate, 
apex serrate with 3-7 teeth, caducous. Male 
flowers: not fragrant, diameter 7.5-8.5 mm, 
glabrous; pedicels 3.5-5 mm long; sepals 4, 
outer pair joined, inner pair free, imbricate, 
D-shaped, convex but not keeled, 2-4 mm 
long, 2-4 mm wide, coriaceous-membranous, 
yellow or green, glabrous, margin thin; petals 
4, free, orbicular, convex, not keeled, becoming 
reflexed but remaining incurved at apex, 3-4.5 
mm long, 3-4.5 mm wide, membranous, 
green; androecium 4-phalangiate; phalanges 
antepetalous, strap-shaped, ribbed, erect and 
becoming slightly reflexed, 1-4 mm long, 
1-2 mm wide, apex undivided or divided into 
two short arms; anthers c. 20 per phalange, 
completely enveloping phalange apex, 
oblong, 0.8-1 mm long, c. 0.3 mm wide; 
pistillode fungiform, 2-2.5 mm long, 2-2.8 
mm wide at apex, apex dark orange-red or 
reddish-brown and quickly becoming viscous 
once flowers are open. Female inflorescences 
axillary, an umbel, 3-5-flowered, peduncles 
1-2 mm long; bracteoles triangular, c. 1.2 
mm long and 1 mm wide, muricate, margin 
3-5-toothed, caducous. Female flowers: 
occasionally fragrant, diameter 6-8 mm; 
pedicels 2-5 mm long; sepals 4, D-shaped or 
deltoid, convex, not keeled or slightly keeled, 


13 

coriaceous-membranous, yellow or green, 
margin thin; outer pair joined, c. 2 mm long, 
2-3 mm wide; inner pair free, 2-4 mm long, 
2.5-4 mm wide; petals 4, free, imbricate, 
becoming reflexed, orbicular, convex, not 
keeled, 2.2-4.2 mm long, 2.8-4.2 mm wide, 
coriaceous-membranous, yellow or green, 
margin thin; gynoecium fungiform, glabrous; 
ovary ellipsoid, 0.6-1.5 mm long, 4-7 mm 
wide, 2-locular, uniovulate; stigma disc-like, 
entire, diameter 4-5 mm, yellow, orange, 
rusty red or brownish-red, quickly becomes 
viscous once flowers are open. Fruiting 
pedicel 2.5-5 mm long; sepals absent from 
base; fruit a fleshy berry, ellipsoid, 25-37 
mm long, 10-18 mm wide, red to dark red or 
blackish; stigma at apex, diameter 4-6 mm; 
seeds one or two, surface slightly undulating, 
22-32 mm long, 11-12 mm wide; testa brown; 
aril meagre, clear, gelatinous, baconwood , 
marblewood. Fig. 1G & H. 

Additional selected specimens (from 118 examined): 
Australia. Queensland. Cook District: Cedar Bay 
NP, western slopes of Mt Finnigan, Oct 1999, Forster 
PIF25016 & Booth (BRI); Mt Hemmant just N of Noah 
Creek in Cape Tribulation area, Jul 1973, Webb 11967 
& Tracey (BRI); Mt Windsor NP, Nov 2008, Forster 
PIF34675, Jensen & McDonald (BRI); Mt Misery, Jun 
1992, Forster PIF10759, Sankowsky & Tucker (BRI); 
Pinnacle Rock Track, 4.5 km W of Karnak, Jun 1992, 
Forster PIF10700, Sankowsky & Tucker (BRI); TR 66, 
Mt Lewis, Jun 1997, Forster PIF21239, Jensen & Tucker 
(BRI); Mt Lewis Road, Aug 1957, Smith 10094 (CANB); 
Davies Creek, SF 607, Aug/Sep 1971, Tracey 14745 
(BRI); SFR 607, Emerald LA, Aug 1982, Gray 2701 
(CNS); Mt Haig, Oct 2012, Cooper 2147 & Ford (CNS); 
Mt Haig, CSIRO plot E27, Oct 2012, Cooper 2148 & Ford 
(CNS); Lake Morris Road at start of track to tower 20, 
Sep 2012, Dowe 1159 (CNS); Copperlode Dam, Cairns, 
Oct 1970, Gittins 2196 (NSW); Palmerston Ridge Track, 
Wooroonooran NP, Feb 1995, Hunter 1109 (BRI); Breeden 
property, Hughes Road, Topaz, Aug 1997, Cooper 1152 
& Cooper (CNS); Track to Phoenix Battery, Towalla, Jul 
1997, Cooper 1132 & Cooper (CNS); Elinjaa LA, Oct 
1974, Irvine 961 (CNS); Tully Falls NP, 2.2 km along 
Tully Falls road from Charmillan Creek bridge, Nov 2011, 
Ford5909 & Torello-Raventos (CNS); SFR 344, Douglas 
Creek, 0.7 km along Carron Creek Road from Kirrama 
Road, Jun 2002, Ford 3510 & Holmes (CNS). 

Distribution and habitat : Garcinia zichii 
is endemic to the Wet Tropics bioregion of 
north-east Queensland from Mt Finnigan 
to the Kirrama Range (Map 2) at altitudes 
between 150-1300 m. It occurs in rainforest 
(evergreen simple to complex notophyll/ 


14 

mesophyll vine forest) on soils derived from 
granite and rhyolite. 

Garcinia zichii grows often in 
association with Beilschmiedia oligandra 
L.S.Sm., Cryptocarya angulata 

C.T.White, Elaeocarpus elliffii B.Hyland 
& Coode, Flindersia pimenteliana 

F. Muell., Planchonella euphlebia (F.Muell.) 
W.D.Francis, Syzygium endophloium B.Hyland 
& S. wesa B.Hyland. 

Phenology : Flowers have been recorded in 
October and November; fruits have been 
recorded from June to November. 

Affinities : Within Australia, Garcinia zichii 
is most similar to G. jensenii. Both species 
have similarly shaped leaves and exudate 
canal pattern, a 2-locular ovary and 1 or 
2-seeded fruit. G. zichii has leaves with 12- 
18 pairs of secondary veins, flowers with 
green or yellow petals and fruit that are 
distinctly ellipsoid. G. jensenii has leaves 
with 7—10 pairs of secondary veins, flowers 
with white petals and fruit that are globose or 
ellipsoid-globose. G. zichii is also similar to 

G. hunsteinii Lauterb. from New Guinea. G. 
zichii has colourless, slightly milky or white 
latex, leaf apex bluntly acute and fruit much 
longer than wide, whereas G. hunsteinii has 
white or yellow latex, leaf apex acuminate 
and globose fruit. 

Notes : There is one specimen (Cooper 1152) 
as well as an observation by Tim Hawkes 
pers. comm., of fruit dehiscing, which was 
presumably caused by recent abundant 
rainfall. This phenomenon is well documented 
within horticulture for other plant groups, 
occurring after superfluous moisture is 
applied to orchard plantings. The abundant 
water causes osmotic absorption through the 
epicarp and the disintegration of the cuticle 
(Opara et al. 1997: 227). 

The exudate-containing canals within 
the leaves (a useful distinguishing character 
in Australian Garcinia) of this species are 
indistinguishable from G. jensenii (Fig. 2C). 

Etymology : The specific epithet honours 
Frank Zich (1968-), a most competent and 


Austrobaileya 9( 1): 1-29(2013) 

obliging curator of the collection at the 
Australian Tropical Herbarium (CNS). 

6. Garcinia leggeae W.E.Cooper sp. nov. 
Similar to Garcinia riparia but differs by 
flower diameter (5 mm versus 15 mm), stamen 
number (5-9 versus 40-50), fruit shape (ovoid 
versus obovoid, oblate or patelliform), fruit 
size (20-27 mm long and 29-45 wide versus 
c. 70 mm long and c. 50 mm wide), sepal size 
on fruit (c. 1.5 mm long versus c. 5 mm in G. 
riparia). Typus: Australia: Queensland. Cook 
District: Gordon Creek, Iron Range National 
Park, 5 December 2012. W.Cooper 2187, 
R. Jensen & J.Kemp (holo: CNS [2 sheets + 
spirit], iso: 7 sheets to be distributed to BRI, 
CANB, K, KEP, L, MO, SING). 

Garcinia sp. (Claudie River LJB 19658) in 
part; Hyland et al. (2003). 

Garcinia sp. (Claudie River); Cooper & 
Cooper (2004: 123). 

Garcinia riparia auct. non A.C.Sm.; Cooper 
& Cooper (2004: 123). 

Garcinia sp. (Claudie River L.J.Brass 19658) 
in part; Jessup (2007: 54, 2010: 44). 

Garcinia sp. Claudie River (L.J.Brass 19658) 
in part; CHAH (2007); Hyland et al. (2010). 

Illustrations : Christophel & Hyland (1993: 
81) as G. riparia vel aff.; Cooper & Cooper 
(1994: 146-147) as G. riparia vel. aff: y Hyland 
et al. (2003) as G. sp. (Claudie River LJB 
19658); Cooper & Cooper (2004: 123) as G. 
sp. (Claudie River) & as G. riparia. 

Shrub or small tree to 6 m, dioecious; exudate 
yellow; dbh to 100 mm, often multi-stemmed; 
bark grey or greyish-brown, fissured and 
flakey, lenticels numerous; branchlets 
square, glabrous. Leaves discolorous, almost 
membranous, shiny on upperside and less so 
on underside, glabrous; petioles 5-8 mm long, 
channelled, not ligulate; lamina elliptical, 
30-145 mm long, 25-57 mm wide, base 
cuneate; apex acuminate or caudate, rarely 
acute or obtuse; margin narrowly recurved; 
venation brochidodromous, primary vein 
flush on upperside and raised on underside; 
secondary veins 12-16 pairs, slightly raised 
on both surfaces, becoming puckered in dry 


Cooper, Garcinia in Australia 

specimens, angle of divergence from primary 
vein c. 45°, intramarginal vein 0.6-1.2 mm 
from margin; exudate-containing canals 
numerous, continuous parallel or intertwined 
lines running at angles between + parallel 
to primary vein and diverging up to c. 45° 
Male inflorescences axillary or ramiflorous, 
a fascicle seated on a pulvinus, 1-3-flowered; 
bracteoles persistent, triangular, c. 0.7 mm 
long and 0.8 mm wide, base truncated, 
apex acute, glabrous, margin entire. Male 
flowers: fragrant, diameter 5-6.5 mm, 
glabrous, sessile; sepals 4, 2-whorled, free, 
ovate, convex, entire, c. 1.5 mm long and 1 
mm wide, membranous, green or yellowish, 
glabrous, margin thin; petals 4 (rarely 5), 
free, explanate, oblong-elliptic, sometimes 
slightly convex, 2.5-3 mm long, c. 1.5 mm 
wide, membranous, cream or yellow, margin 
entire or minutely ciliate; androecium a fleshy 
receptacle not adnate to petals, c. 0.5 mm 
long; anthers 5-9, sessile or almost sessile, 
erect or extrorse, quadrate, c. 0.4 mm long 
and 0.4 mm wide, bithecate; pistillode absent. 
Female inflorescences axillary or ramiflorus, 
a solitary flower, sessile; bracteoles persistent, 
triangular, convex, c. 0.8 mm long and 
0.8 mm wide, base truncated, apex acute, 
glabrous, margin entire. Female fiowers: 
fragrant, diameter c. 5 mm, sessile; sepals 
4, 2-whorled, free, ovate, convex, 2-3 mm 
long, 1.5-2 mm wide, membranous, green 
or yellowish, glabrous, margin thin, entire; 
petals 4, free, spreading widely, oblong- 
elliptic, sometimes slightly convex, c. 3 mm 
long and 1.5 mm wide, membranous, cream 
or yellow, glabrous, margin entire and thin; 
staminodal phalanges 4, adnate to petals, 
filament-like with a globose apex, distally 
inflexed, c. 1.4 mm long; antherodes 1 per 
phalange, c. 0.8 mm long; ovary urceolate, 
c. 2.5 mm long and 1.5 mm wide, sessile, 
6-13-locular, uniovulate; stigma sessile, 
diameter c. 1.5 mm. Fruiting pedicel 0.7-1 
mm long, sepals persistent at base c. 1.5 
mm long, fruit a fleshy berry, ovoid (oblate, 
patelliform or asymmetrical when some 
ovules infertile), 20-27 mm long, 29-45 mm 
wide, pink-red to dark red, 6-13 septum lines, 
6-13 segments; seeds 6-13, hairy, c. 11 mm 
long and 7-8.5 mm wide (not including hairs); 


15 

testa brown; aril fleshy, translucent whitish or 
pinkish. Figs. IE, 2F. 

Additional selected specimens (from 22 examined): 
Queensland. Cook District: Claudie River, Nov 1977, 
Hyland RFK3623 (CNS); ditto , Jan 1982, Hyland 11515 
(CNS); ditto , Dec 1982, Hyland 12412 (CNS); ditto , 
Dec 1982, Hyland 12413 (CNS); West Claudie River, 
Iron Range, Aug 1997, Jensen 874 (CNS); Iron Range, 
Sep 1962, Volck AF02600 (CNS); Iron Range NP, 
CSIRO EP/42, Sep 2008, Ford 5422, Metcalfe, Murphy 
& Bradford (CNS); Claudie River EP/42, July 1978, 
Unwin 625 (CNS); Claudie River, Iron Range, May 
1992, Cooper 279 & Cooper (CNS); Claudie River 
area, Oct 1968, Webb 8335 & Tracey (BRI); Claudie 
River, Iron Range NP, Dec 2012, Cooper 2168, Jensen & 
Kemp (CNS); Gordon Creek, Iron Range NP, Dec 2012, 
Cooper 2171 , 2175, 2177, 2186, 2188, Jensen & Kemp 
(CNS); ditto , Jun 2014, Cooper 2220 & Cooper (CNS); 
Gordon Creek, 10 km ENE of Mt Tozer, Iron Range NP, 
May 1992, Fell 2543 (BRI). 

Distribution and habitat : Garcinia leggeae 
is endemic to rainforest (evergreen mesophyll 
vine forest) in the Iron Range area, mostly 
on the Claudie River and Gordon Creek 
floodplains but occasionally on hillslopes to 
an altitude of 75 m (Map 3). 

Garcinia leggeae grows as an understory 
shrub or small tree on alluvial soils derived 
from basalt, frequently under a canopy 
of Blepharocarya involucrigera F.Muell., 
Calamus australis Mart., Garcinia dulcis , 
G. warrenii, Lasjia claudiensis C.L.Gross & 
B. Hyland, Myristica insipida R.Br., Nauclea 
orientalis (L.) L., Pandanus zea H.St.John, 
Syzygium bamagense B.Hyland & Tetrameles 
nudiflora R.Br. 

Phenology: Flowers have been recorded in 
June and December; fruits have been recorded 
in May, August and September. 

Affinities: Garcinia leggeae is similar to G. 
riparia from New Guinea. Compared with G. 
riparia, G. leggeae is a more diminutive plant, 
having much smaller flowers (5 mm versus 15 
mm) with fewer stamens (5-9 versus 40-50), 
smaller fruit (20-27 mm long and 29-45 wide 
versus c. 70 mm long and c. 50 mm wide) and 
smaller sepals on ripe fruit ( c . 1.5 mm long 
versus c. 5 mm). 

Notes: Plants are frequently multi-stemmed, 
presumably caused by the forces of floodwater 
and floating debris during the wet season. 


16 

Some leaves on most herbarium 
collections, and all plants in the field seen 
by the author, are marked with black spots 
(diameter c. 5 mm), which appear to be caused 
by insects rather than fungi. 

Etymology : The specific epithet honours 
ornithologist and ecologist Sarah Legge 
(1969-) who collected this species as well as 
many other valuable specimens for illustration 
in the fruit book (Cooper & Cooper 2004), 
while working in the Iron Range area. 

7. Garcinia mestonii F.M.Bailey, Report on 
New Plants, Preliminary to General Report 
on Botanical Results on Meston’s Expedition 
to the Bellenden-Ker Range 2 (1889), as 
‘mestoni ’■ Type: Australia: Queensland. 
Cook District: Bellenden-Ker Range at an 
altitude of 2000 feet, [June 1889], [A.Meston 
& D.Whelan s.n .] (holo: BRI [AQ340247] 1 ). 

Illustrations : Christophel & Hyland (1993: 
81); Cooper & Cooper (2004: 123), Hyland et 
al. (2010). 

Shrub or small tree to 6 m, dioecious (rarely 
monoecious); exudate yellow, meagre; bark 
nondescript with irregular-shaped lenticels; 
branchlets square when young, becoming 
terete. Leaves discolorous, coriaceous, 
glabrous; petioles 6-11 mm long, channelled, 
not ligulate; lamina elliptic, ovate, lanceolate, 
obovate, oblanceolate, 35-80 mm long, 
12-34 mm wide; base attenuate, cuneate or 
oblique; apex caudate or acuminate; margin 
may be recurved in dry specimens; venation 
brochidodromous, primary vein + flush on 
upperside and raised on underside; secondary 
veins 10 pairs, angle c. 70° to primary 
vein, forming loops c. 1 mm from margin; 
exudate-containing canals numerous as short 
streaks, dashes and dots (more conspicuous 
on upperside), running at angles between 
45° and parallel to primary vein or margin. 
Male inflorescences axillary, terminal or 
ramiflorous, a 2-5-flowered umbel or fascicle; 
bracts oblong, convex, 1.5-5.5 mm long, 0.5- 
0.8 mm wide, glabrous, apex obtuse, base 
truncate, caducous; bracteoles triangular, c. 
0.8 mm long and 0.6 mm wide, caducous; 
peduncles absent or up to 4 mm long. Male 
flowers: not fragrant, diameter c. 4 mm; 


Austrobaileya%\). 1-29(2013) 

pedicels 1.5-2.5 mm long; sepals 4,2-whorled, 
outer pair joined, D-shaped, cupular, c. 4 
mm long and 4.5 mm wide, green, glabrous, 
margin thin; petals 4, orbicular, cupular with 
apex recurved, c. 4.8 mm long and 4.8 mm 
wide, membranous, green or cream, glabrous, 
margin thin; androecium a fleshy receptacle 
not adnate to petals, c. 2.5 mm long and 3.2 
mm wide; anthers c. 16, triangular, sessile or 
almost sessile, 0.6-1 mm long, 0.6-1 mm wide 
at base, bithecate; pistillode absent. Female 
inflorescences axillary or terminal, solitary 
flowers or 2-3-flowered umbels or fascicles; 
bracts 4-whorled, oblong, 1.5-3.2 mm long, c. 
1 mm wide, caducous; bracteoles triangular, 
c. 1 mm long and 0.6 mm wide; peduncle c. 
1 mm long. Female flowers: not fragrant, 
pedicels c. 2.5 mm long, diameter 6-7.5 mm; 
sepals 4, 2-whorled, outer pair joined, inner 
pair free, D-shaped, cupular, 3.8-4.4 mm 
long, 4.3-5.5 mm wide, membranous, green 
or cream, glabrous; petals 4, free, orbicular, 
cupular with apex recurved, c. 5 mm long 
and 4 mm wide; staminodal phalanges adnate 
to petals, narrowly or broadly triangular, 
apically 2 or 3-branched, distally indexed, 
c. 2 mm long, antherodes 2 per branch; 
ovary oblate, 2.2-2.8 mm long, 4-4.5 mm 
wide, sessile, 3-9-locular, uniovulate; style 
c. 0.5 mm long, stigma diameter c. 2.5 mm, 
3-9-rayed. Fruiting pedicel 1-3 mm long; 4 
sepals persistent at base; fruit a fleshy berry, 
depressed globular, 31-50 mm long, 40-60 
mm wide, cream or greenish-cream; stigma 
3-9-rayed, diameter 3.5-4 mm; seeds 13-25 
mm long, c.10 mm wide, testa brown; aril 
fleshy, cream. Meston’s mangosteen. Fig. 1C 
& F, 2D. 

Additional selected specimens (from 26 examined): 
Australia. Queensland. Cook District: Little Cooper 
Creek, Cow Bay, Cape Tribulation area, Oct 1994, 
Small s.n. (CNS [QRS104526]); Behana Creek, S of 
Gordonvale, Jul 2005, Halford Q8446 & Jensen (BRI); 
Divide between Babinda Creek and East Mulgrave, May 
2002, Jago 6211 (CNS); Mt Harold, Aug 1985, Irvine 
2322 , 2323 (BRI, CNS); Goldsborough Road, Mulgrave 
River Valley, Jul 1992, Tucker s.n. (CNS [QRS105577]); 
Goldsborough Road, Jul 1992, Sankowsky 1332 (CNS); 
East Mulgrave River, Nov 1995, Jensen 483 (CNS); 
Ridgeline NW of airplane wreck, Bellenden Ker, Jun 
1995, Hunter 5308 (BRI); Mt Bellenden Ker, c. 7/8 mile 
SE of centre peak, Jun 1969, Smith 14691 (BRI); Harvey 
Creek, N of Bellenden Ker, Dec 1998, Forster PIF24028 
(CNS); Harvey Creek, Dec 2012, Cooper 2203, 2204 & 


Cooper, Garcinia in Australia 

Jensen (CNS); Cultivated at Tolga, ex Harvey Creek, 
Dec 2003, Sankowsky 2326 & Sankowsky (BRI); 
Sankowsky’s garden, Tolga [ex Harvey Creek], Jan 2013, 
Cooper 2209, Sankowsky & Jensen (CNS); NPR 226, 
Parish of Bellenden Ker, Dec 1983, Hyland 25062 RFK 
(BRI, CNS); Ridgetop S of Tower 6, Bellenden Ker, Jun 
1995, Hunter 5292 (BRI); E slopes of Bellenden Ker 
near Station 3 on the Bellenden Ker Cable Car Track, 
Jan 1981, Irvine 2056 (BRI, CNS); Bellenden Ker, s.dat., 
Meston 1 (BRI [AQ166293]); Bellenden Ker Range, 
Mar 1922, White s.n. (BRI [AQ166291]); Bellenden 
Ker, below summit, Jun 1949, Smith 4236 (BRI); 
Wooroonooran NP, headwaters of East Mulgrave River, 
5 km W of Bellenden Ker township, Nov 2000, Forster 
PIF26412, Booth & Jensen (BRI). 

Distribution and habitat: Garcinia mestonii 
is endemic to the Wet Tropics bioregion 
of north-east Queensland between Cape 
Tribulation and the Mt Bellenden Ker Range 
(Map 3). It occurs in rainforest (evergreen, 
simple/complex notophyll/mesophyll vine 
forest) on granitic and metamorphic soils at 
altitudes between 30 and 1340 m. 

Phenology: Flowers have been recorded 
in December and January; fruit has been 
recorded from August to February. 

Typification: The type collection at BRI has 
minimal original labelling attached to the 
sheet, apart from a tattered scrap of paper with 
the number 18 and the words ‘Meston’s Apple’. 
No specific collectors or date of collection 
are given; although a copy of the protologue 
is attached. In the protologue Bailey (1889) 
noted that it was collected by Messrs. Meston 
and Whelan on the first ascent of Bellenden 
Ker. As noted by Dowe & Broughton (2007) 
this was in June 1889. 

Archibald Meston is well known as the 
leader and organiser on the 1899 and 1904 
Bellenden Ker expeditions, but was also 
a significant collector of plant specimens, 
particularly on the latter expedition when 
F.M.Bailey was absent (Dowe & Broughton 
2007, 2008). The co-collector of the type of 
Garcinia mestonii was Daniel Whelan, a 
Sergeant in the infamous [Queensland] Native 
Mounted Police (Richards 2008) and who led 
the force of troopers allocated as protection 
on the 1889 expedition. Whelan was honoured 
by Bailey with the Proteaceous tree Lasjia 
whelanii (originally as Helicia whelanii 
F.M.Bailey) and the fern Blechnum whelanii 


17 

F.M.Bailey. The ‘Whelanian Pools Camp’ on 
Bellenden Ker was also named for him (Dowe 
& Broughton 2007). 

Notes: Garcinia mestonii has been recorded 
as dioecious in the field; however, one plant 
cultivated by G & N Sankowsky at Tolga 
is distinctly monoecious with about 50% 
of flowers being male and 50% female. 
Individual branches on this tree appear to 
bear flowers of one sex only. 

Etymology: The specific epithet honours 
Archibald Meston (1851-1924), a botanical 
collector, journalist, civil servant, explorer 
and member for Rosewood in the Queensland 
Legislative Assembly from 1878-1882. 

8. * Garcinia mangostana L ., Sp. PI. 443 
(1753). Type: ‘Mangostan’ in L.Garcin, 
Philos. Trans. 38: t. 1 (1734); fide Hammel 
(1993: 28). 

Illustrations: Fleming ( c. 1795-1805); 

Hooker (1855: t. 4847); van Nooten (1863); 
Marsden (2005: plate 3); Nazre (2006: 41, 45, 
49, 52, 54, 57, 63, 181); Te-chato (2007: 248, 
249, 250). 

Tree to 20 m, agamospermous; exudate 
yellow; bark fissured and flaky; branchlets ± 
terete with some ribbing, glabrous. Leaves 
discolorous, coriaceous, glabrous, drying 
reddish-brown; petioles 10-30 mm long, not 
channelled on fresh specimens and slightly 
channelled on dried specimens, ligulate; 
lamina elliptic, broadly elliptic, ovate or 
oblanceolate, 100-250 mm long, 45-105 mm 
wide, base cuneate or obtuse; apex acute, 
shortly acuminate or obtuse; margin narrowly 
recurved; venation brochidodromous, 
primary vein distinctly raised on both sides; 
secondary veins 15-17 pairs, prominent, 
angle of divergence 50-70° from primary 
vein; 2 intramarginal veins, outermost 
0.5-2 mm from margin, innermost 3-7 mm 
from margin; exudate-containing canals 
on underside as numerous branched lines 
running at angles between parallel to primary 
vein and margin, on upperside as numerous 
branched lines mostly running parallel to 
secondary venation. Male flowers unknown. 
Female inflorescences terminal, a solitary 
flower, pairs or rarely in 3’s. Female flowers: 


18 

diameter 25-60 mm; pedicels 1.5-20 mm 
long; sepals, 4, 2-whorled, c. 20 m long, 
green, cream, yellowish, pinkish or reddish, 
cupular, coriaceous; petals 4, free, erect 
to spreading, broadly obovate or broadly 
orbicular, c. 25 mm long, cream, pink or 
reddish, glabrous, thickly fleshy; staminodes 
up to 20 shorter than ovary, adnate to petals, 
free, filament-like, c. 5 mm long, caducous; 
antherodes 1 per filament; ovary broadly 
ovoid or ellipsoid or globose, 12-16 mm, 
sessile, 4-8-locular, locules uniovulate; 
stigma mass sessile, domed, diameter 8-12 
mm, 4-8-lobed. Fruiting pedicel with 4 
green to reddish sepals persistent at base; 
fruit a dry berry with a thick corky-leathery 
pericarp c. 9 mm wide, globose or depressed 
globose, diameter 35-70 mm, purple to 
blackish, glabrous; sepals persistent; stigma 
persistent, 4-8-rayed; seeds 4-8, 15-25 mm, 
testa brown; aril thickly fleshy and white. 
mangosteen, Queen of fruit. 

Additional selected specimens (from 5 examined): 
Queensland. Cook District: Cultivated, Stanton 
property. Intake Road, Redlynch, Cairns, Jul 2013, 
Cooper 2222 (CNS); North Johnstone River, off the 
Cooroo Lands Road, W of Innisfail, Jul 2007, Bean 
26749 (BRI); Dunk Island, Aug 1963, Scanlan AFO 
2812 (CNS); Dunk Island, Aug 1964, Scanlan AFO 3490 
(CNS); Tully, Sep 1941, Shearer s.n. (CNS). 

Distribution and habitat: Garcinia 
mangostana is unknown in the wild but 
is likely to have originated on Peninsula 
Malaysia (Richards 1990b). In Australia 
it is cultivated in the tropical lowlands of 
Queensland, and a recent collection ( Bean 
26749), in the Innisfail area, confirms 
naturalisation within Queensland rainforest. 
Three old collections from Dunk Island and 
Tully also indicate possible naturalisation 
(Map 3). 

Phenology : Flowers and fruit have not been 
collected from naturalised plants; however, 
Peter Stanton and Allen Sheather have 
cultivated plants in the Cairns and Daintree 
areas and record flowers from September to 
March; fruit December to July. 

Notes: Garcinia mangostana is thought to be 
a hybrid of multiple origins (Richards 1990b). 
Within tropical Asia, Mangosteens have been 
widely cultivated over hundreds of years for 


Austrobaileya 9(1): 1-29(2013) 

their delicious arils and are referred to as the 
Queen of Fruit. Several authors including 
Richards (1990a), suggest that male flowers 
are probably always absent and the species 
is an obligate agamosperm, forming fruit 
without fertilisation. 

Etymology: The specific epithet is derived 
from Latinising the Malay name manggis. 

9. Garcinia brassii C.T.White, Proc. Roy. Soc. 
Queensland 47: 52 (1936). Type: Australia: 
Queensland. Cook District: Thornton Peak, 
in low scrub at the summit, 14 March 1932, 
L.J. Brass 2277 (holo: BRI). 

Illustrations: Christophel & Hyland (1993: 
81); Cooper & Cooper (2004: 123); Hyland et 
al. (2010). 

Tree to 18 m, dioecious; exudate yellow; bark 
with fine vertical fissures; branchlets square 
becoming terete, new twigs mostly red, 
glabrous. Leaves discolorous, coriaceous, 
glabrous; petioles 5-12 mm long, shallowly 
channelled on younger growth and on dried 
specimens, ligulate; lamina obovate or 
elliptical, 25-125 mm long, 16-63 mm wide, 
base attenuate or cuneate; apex acute, obtuse 
or emarginate; margin recurved; venation 
brochidodromous, primary vein ± flush on 
upperside but may be raised near base on dry 
specimens; secondary veins raised on both 
sides, 12-35 pairs, angle of divergence from 
primary vein 65-80°, forming loops 0.3-1.2 
mm (rarely 4 mm) from margin, intramarginal 
vein often hidden in the recurved margin; some 
tertiary veins or veins forking off secondary 
veins may be present, if so, most diminish and 
do not reach the intramarginal vein; rarely are 
reticulations of similar strength to secondary 
veins present and if they do they are few and 
near the margin; exudate-containing canals 
on underside as numerous parallel lines, 
branched, straight, wriggly or curled, running 
at angles radiating between primary vein 
and margin. Male inflorescences terminal, 
a panicle, 3-18-flowered, 19-28 mm long; 
peduncles 10-21 mm long; bracts obovate 
or elliptical, keeled, 5-7 mm long, 2-3 mm 
wide (these may expand into small leaves up 
to 55 mm long and 21 mm wide), caducous; 
bracteoles 2-2.5 mm long and c. 2 mm wide. 


Cooper, Garcinia in Australia 

Male flowers: fragrant, diameter 15-28 mm; 
pedicels absent or up to 6 mm long; sepals 
4, 2-whorled, dark red, glabrous, margin 
thin; outer pair joined, D-shaped or ovate- 
orbicular, convex, 4-6.5 mm long, 4-6.5 mm 
wide, coriaceous; inner pair free, orbicular or 
oblong-orbicular, convex, 7-8 mm long, 6-7 
mm wide; petals 4, free, erect or spreading, 
imbricate, orbicular, convex, 9-14 mm 
long, 8-14 mm wide, waxy-coriaceous, 
cream, yellow, pink or reddish, unmarked 
or with dark red markings, glabrous, margin 
thin; androecium 4-phalangiate, glabrous; 
phalanges adnate to petals, obovate or elliptical 
at anthesis, 5-8 mm long, 5-6 mm wide; 
anthers several to numerous per phalange, 
sparsely to densely clustered over phalange, 
bilobed, c. 0.5 mm long and 0.5 mm wide; 
pistillode rudimentary, attenuate-conical, 

1- 3.5 mm long. Female inflorescences 
terminal, a solitary flower 10.5-12 mm long; 
peduncle 1.5-3 mm long; bracts at apex of 
peduncle triangular, keeled, 3-7 mm long, 

2- 3 mm wide, caducous; bracteoles 1.5-2 
mm long and 2 mm wide. Female flowers: 
fragrant, diameter 13-18 mm; pedicels absent 
or up to 1.5-3.5 mm long; sepals 4,2-whorled, 
coriaceous, green and mostly blushed dark- 
red towards apex, glabrous, margin thin; 
outer pair joined, oblong, convex, c. 5 mm 
long and 4 mm wide; inner pair free, broadly 
ovate or orbicular, convex, 6-8 mm long, 5-7 
mm wide; petals 4, free, erect to spreading, 
orbicular, convex, 8-10 mm long, 8—10 mm 
wide, waxy-coriaceous, cream or yellowish, 
often with red markings and often ageing 
to entirely reddish, glabrous, margin thin; 
staminodal phalanges 4, erect, adnate to 
petals, strap-shaped with flabellate apex, c. 3.5 
mm long and 4 mm wide, apically branched 
into 5-8 individual filaments c. 1 mm long; 
antherodes 1 or 2 per filament, 0.5- 0.7 mm 
long; gynoecium globose, 6-7.5 mm long, 
4-5 mm wide, glabrous; ovary ovoid, 5-6 mm 
long, 4-5 mm wide, sessile, 4-locular, locules 
uniovulate; stigma mass sessile, domed, 
diameter 4.5-5.5 mm, cream becoming 
reddish with age or exposure. Fruiting 
pedicels 2-3.5 mm long, 4 sepals persistent at 
base, fruit a fleshy berry, depressed globular, 
22-45 mm long, 27-50 mm wide, dark red 


19 

to blackish, glabrous; stigma shallowly 3 or 
4-lobed, diameter 5.5-6.5 mm; seeds 1-4, 
12-22 mm long, 10-18 mm wide; testa brown, 
smooth; aril fleshy, dark red. Fig. 2A, 3A-C. 

Additional selected specimens (from 36 examined): 
Queensland. Cook District: Near peak of walking 
track to Mt Sorrow, 3 km W of Cape Tribulation, 
Daintree NP, Sep 2009, Worboys 825 (CNS); NPR 133, 
Daintree, summit of Mt Sorrow track, Dec 1998, Ford 
2147 (CNS); VCL Noah, Mt Hemmant, Oct 1975, Hyland 
8490 (CNS); Thornton Peak, Jul 2008, Jago 7152, Ford 
& Keith (BRI); Thornton Peak, Sep 1937, Brass & White 
280 (BRI, K 1 ); Thornton Peak, in valley between N 
and S peaks near campsite on Hilda Creek, Sep 1984, 
Clarkson 5588 (CNS); Pinnacle Rock Track, 4.5 km W 
of Karnak, Jun 1992, Forster PIF10708, Sankowsky & 
Tucker (CNS); Along track to Devils Thumb, Sep 1997, 
Jago 4496 (BRI); Mt Misery, E of Mt Spurgeon, 15.7 km 
NE of Mt Carbine, Nov 1988, Jessup GJM861, Guymer 
& McDonald (BRI); Mossman Bluff c. 10 km W of 
Mossman, Dec 1998, Fell 1683 (BRI); Near Schiller’s 
hut, Mt Spurgeon, Sep 1972, Webb 13438 & Tracey 
(CNS & BRI); 27.3 km from Rex Highway on Mt Lewis 
Road, Nov 1990, Holland 31 & Hind (NSW); Mt Lewis 
Road S[South] Mary LA, Nov 1988, Jessup GJM1526, 
Guymer & McDonald (BBS), Mt Lewis FR, 28 km along 
Mt Lewis Road, Jan 2002, Forster PIF28164, Booth & 
Jensen (BRI); Mt Lewis, on track to Rhododendrons, 
Sep 2012, Cooper 2163 & Ford (CNS); ditto , Dec 2012, 
Cooper 2207 & Ford (CNS); Mt Lewis, CSIRO plot 
EP18, Sep 2012, Cooper 2162, 2164 & Ford (CHS). 

Distribution and habitat : Garcinia brassii 
is endemic to the Wet Tropics bioregion of 
north-east Queensland from Mt Sorrow to 
Mossman Bluff and west to the Mt Lewis -Mt 
Spurgeon areas (Map 1), at altitudes between 
500-1300 m. It occurs in rainforest (simple 
notophyll vine forest, complex notophyll 
vine forest and simple microphyll vine-fern 
thicket) on granitic soils. 

Phenology : Flowers have been recorded in 
June, August to November; fruit has been 
recorded in December and January. 

Etymology : The specific epithet is in honour 
of botanist Leonard J. Brass (1900-1971), 
collector of the type specimen. 

10. Garcinia gibbsiae S.Moore, J. Bot. 55: 
302 (1917). Type: Australia: Queensland. 
Cook District: Bellenden Ker, forest on long 
ridge, March 1914, L.S.Gibbs 6306 (holo: 
BM 1 ; iso: K 1 ). 


20 


Austrobaileya 9(1): 1-29 (2013) 


Fig. 3. Flowers. A. Garcinia brassii showing stigma and apically branched staminodal phalanges around the ovary 
(i Cooper 2163 & Ford [CNS]). B. G. brassii $, showing 4-phalangiate androecium prior to anthesis ( Cooper 2164 
& Ford [CNS]). C. G. brassii S, showing 4-phalangiate androecium post anthesis ( Cooper 2164 & Ford [CNS]). D. 

G. gibbsiae showing stigma and apically branched staminodal phalanges around ovary {Cooper 2213 & Cooper 
[CNS]). E. G. gibbsiae S, showing androecium of 4 fused phalanges {Cooper 2212 & Cooper [CNS]). F. G. warrenii 
& showing anthers on 4-phalangiate androecium post anthesis {Cooper 2158 & Hawkes [CNS]). G. G. russellii 
showing stigma and apically branched staminodal phalanges around ovary {Cooper 2150, Russell & Hawkes [CNS]). 

H. G. russellii 3 , showing androecium of 4 fused phalanges {Cooper 2149, Russell & Hawkes [CNS]). I. G. warrenii 
showing stigma and apically branched staminodal phalanges around the ovary {Cooper 2159 & Hawkes [CNS]). 

Photos: A, D, E, W.Cooper, B, C, F, G, H, I, T.Hawkes. 


Cooper, Garcinia in Australia 

Illustrations : Christophel & Hyland (1993: 
81); Cooper & Cooper (1994: 288); Cooper & 
Cooper (2004: 123); Hyland et al. (2010). 

Tree to 12 m, dioecious; exudate bright 
lime-yellow; bark with fine vertical fissures 
and irregular lenticels; branchlets shallowly 
ribbed, horizontal, erect or pendulous, 
glabrous. Leaves discolorous, coriaceous, 
glabrous; petioles 8-20 mm long, channelled, 
ligulate; lamina oblong-elliptic, ovate, 
oblanceolate, elliptical or elliptic-obovate, 65- 
200 mm long, 30-93 mm wide, base cuneate 
or obtuse, apex acute or obtuse, margin 
thickened and sometimes remotely recurved; 
venation paxillate, primary vein raised on 
both sides; secondary veins numerous (more 
than 35), + flush, parallel, angle of divergence 
with primary vein 70-90°, intramarginal vein 
on the margin; exudate-containing canals 
similar on both sides as sparse lines, branched 
or unbranched, mostly running at a slightly 
narrower angle than the secondary veins 
and sometimes crossing over them. Male 
inflorescences terminal or rarely axillary, 
a solitary flower, pairs, or a 2-8-flowered 
raceme, up to 38 mm long; peduncle 3-22 mm 
long; bracts narrowly triangular, convex, 5-10 
mm long, 3-4 mm wide, glabrous; bracteoles 
triangular, convex, c. 2 mm long, glabrous. 
Male flowers: not fragrant, diameter 17-46 
mm; pedicel absent or up to 2.7 mm long; 
sepals 4, 2-whorled, convex, may be slightly 
keeled, green, margin thin; outer pair joined, 
D-shaped, 5-8 mm long, 7-9 mm wide; inner 
pair free, orbicular, 5.5-7 mm long, 8-10.5 
mm wide; petals 4 (rarely 3), free, orbicular, 
convex or flat, 11-20 mm long, 11-20 mm 
wide, thick, waxy, yellow (sometimes blushed 
with pink), glabrous, margins thin and mostly 
incurved; androecium of 4 fused phalanges 
creating a solitary receptacle, rhombic, adnate 
to petals, divided into 4 shallow lobes at apex, 
diameter 10-13 mm; anthers numerous, 
densely clustered over receptacle, subsessile, 
strap-shaped or wedge-shaped, bithecate, 
0.9-1.3 mm long, 0.5-1 mm wide, glabrous; 
pistillode absent. Female inflorescences 
terminal, solitary or 2-flowered; peduncles 
1-2.5 mm long; bracts triangular, keeled, 
3-5 mm long, 1.5-2 mm wide, caducous; 
bracteoles triangular, orbicular or ovate, 


21 

slightly keeled, c. 3 mm long and 2 mm wide, 
caducous. Female flowers: faintly fragrant, 
diameter 15-22 mm, pedicels 1.5- 2.5 mm 
long; sepals 4, convex, green, glabrous, 
margin entire and membranous; outer pair 
narrowly joined, D-shaped, 5-6.5 mm long, 
8-10 mm wide; inner pair free, orbicular, 
6.5-8 mm long, 7.5-12 mm wide; petals 
4, free, orbicular, convex, 14-17 mm long, 
11-15 mm wide, waxy-coriaceous, yellow, 
glabrous, margin thin; staminodal phalanges 
4, free or adjacent pairs fused, adnate to 
petals, strap-shaped, erect, 2.5-5 mm long, 
3-3.5 mm wide, apically branched into 6-9 
individual filaments c. 1 mm long; antherodes 
1 or 2 per filament, erect or introrse, oblate, 
c. 1 mm long and 0.6 mm wide; gynoecium 
urn-shaped, 7-9-locular, surface irregularly 
lobed, glabrous; ovary globular, 4-5 mm long, 
5-6 mm wide, sessile, 7-9-locular, 1-4 ovules 
per locule; stigma sessile, domed, 7-9-rayed, 
diameter 5-6 mm. Fruiting pedicel 2-4 
mm long, 4 sepals persistent at base; fruit a 
semi-dry berry, top-shaped, broadly-ovoid 
or truncated-globose, base truncated, apex 
acute, beaked or rarely truncated, 30-62 mm 
long, 30-67 mm wide, green, septal radii 
conspicuous as 7-9 shallow ribs or lines; 
stigma persistent at apex, 7-9-rayed; locules 
7-9, septa thick, coriaceous and entire; seeds 
1-4 per locule, ± globular, 10-18 mm long, 
7-11 mm wide, testa brown, aril absent. 
mountain mangosteen. Figs. 2E, 3D & E. 

Additional selected specimens (from 29 examined): 
Australia. Queensland. Cook District: Mt Bellenden 
Ker, east slopes, April 1948, Brass 18330 (BRI); Mt 
Bellenden Ker, c. 7/8 mile SE of the centre peak, June 
1969, Smith 14690 (BRI); Near junction of E and W 
Mulgrave River in SFR 310, Upper Goldsborough LA, 
19.1 km SSE Little Mulgrave town, Nov 1988, Jessup 
GJM1746, Guymer & McDonald (BRI); Old Mailman’s 
Track, W of Windin Creek, Feb 1995, Hunter 849 (BRI); 
SFR 755, Gosschalk LA, EP/34, Nov 1976, Unwin 117 
(CNS); SFR 310, Windin LA, Apr 1973, Hyland 6702 
(CNS); Lamonds [Lamins] Hill, Atherton Tableland, 
Oct 1949, Stephens NQ Nat. Club No. 13312 (BRI); SFR 
310, Swipers LA, Oct 1963, Hyland 278RFK (CNS); 
Headwaters of Jimpee Creek, s.dat ., Hunter 4522 (CNS); 
TR 1230, Boonjee LA, Oct 1979, Moriarty 2678 (CNS); 
ditto , May 1982, Gray 2550 (CNS); TR 1230, Boonjee 
LA, Boonjee, walking track to Stockwellia site, June 
1992, Tucker & Sankowsky s.n. (BRI [AQ546523]); 
Wooroonooran NP, track to Chuck Lunga Creek, E of 
Boonjee, Forster PIF19286 & Tucker (BRI, CNS); Mt 
Bartle Frere, 1.8 km WSW Bobbin Bobbin Falls, 4.4 km 


22 

NNE Boonjee, Nov 1988, Jessup GJM1049, Guymer & 
McDonald (BRI); Boonjie LA near Bartle Frere track, 
0.9 km SE of Bobbin Bobbin Falls, Nov 1988, Jessup 
GJM269, Guymer & McDonald (BRI); Westcott Road, 
Topaz, Jul 1992, Cooper 320 & Cooper 320 (CNS); ditto , 
Mar 2013, Cooper 2212, 2213 & Cooper (CNS); Topaz, 
Dec 1970, Kitchener s.n. (CNS); Downey Creek SA 34, 
24.6 km SE of Millaa Milla, Oct 1988, Jessup GJM2217, 
Guymer & McDonald (BRI). 

Distribution and habitat : Garcinia gibbsiae 
is endemic to the Wet Tropics bioregion of 
north-east Queensland where it occurs in 
rainforest (complex mesophyll vine forest and 
simple notophyll vine forest) on basalt soil 
from the eastern slopes of the Mt Bellenden 
Ker Range to Downey Creek near Innisfail 
and west to the Boonjie-Topaz area (Map 4), 
at altitudes between 120 and 1160 m. 

Phenology : Flowers have been recorded from 
February to June; fruit has been recorded 
from July to January. 

Affinities : Garcinia gibbsiae appears to be 
most similar to G. russellii. Both species 
have tough leathery leaves with numerous 
parallel secondary veins and green fruit with 
up to three or four seeds in each locule. G. 
gibbsiae is a moderately to densely branched 
shrub or small tree, with bright lime-yellow 
exudate, leaf bases which are either cuneate 
or obtuse and never peltate or cordate, male 
inflorescences solitary or a few-branched 
raceme and petals yellow, fruit with spacious 
locules not completely filled by seeds; locular 
septa thick, fleshy and entire. G. russellii is a 
slender and sparsely branched shrub or small 
slender tree with colourless, milky or white 
exudate, peltate leaves usually with cordate 
bases, male inflorescences a 9- or more- 
flowered panicle and petals cream or white, 
fruit with seeds tightly packed within each 
locule which mostly cause the exocarp to 
bulge; locular septa thin, pithy and incomplete. 

Etymology : The specific epithet honours 
British botanist Lilian Suzette Gibbs (1870— 
1925). 

11. Garcinia russellii W.E.Cooper sp. nov. 
Similar to Garcinia gibbsiae but differs in the 
exudate colour (clear or slightly milky versus 
bright lime-yellow), leaves (peltate versus not 
peltate), male inflorescences (nine or more 


Austrobaileya%\). 1-29(2013) 

flowers versus up to 8-flowered but usually 
less); fruit septa (thin, pithy and incomplete 
versus thick, fleshy and entire). Typus: 
Australia: Queensland. Cook District: North 
of Mossman River, Daintree National Park, 9 
July 2011, W.Cooper 2137, R.Russell, R. Jensen 
& R.Jago (holo: CNS [1 sheet + spirit]; iso: 3 
sheets to be distributed to BRI, CANB, MO). 

Garcinia sp. (Mossman); Cooper & Cooper 
(2004: 124). 

Illustrations : Cooper & Cooper (2004: 124) 
as Garcinia sp. (Mossman). 

Shrub or small tree to 6 m, sparsely branched 
with branches mostly pendulous, dioecious; 
exudate opaque white, colourless or slightly 
milky; bark with a few horizontal raised rings 
at old interpetiolar scars, dotted with small 
pustules; branchlets slightly 4-sided on new 
growth and becoming terete on older growth, 
glabrous. Leaves discolorous, coriaceous, 
glabrous, peltate; petioles 10-25 mm long, 
attached on underside of lamina; lamina 
oblong or ovate-oblong, 110-295 mm long, 
45-100 mm wide, base cordate or rounded, 
apex acute and often almost aristulate, 
margin entire and strongly recurved; venation 
paxillate, primary vein distinctly raised on 
both sides, green on upperside and yellow on 
underside; secondary veins c. 100, raised on 
both sides, angle of divergence from primary 
vein 70-90°, intramarginal vein on the 
margin; exudate-containing canals sparse on 
both sides as unbranched or branched lines, 
mostly running at a slightly narrower angle 
than the secondary veins and sometimes 
crossing over them. Male inflorescences 
terminal, a panicle, 9-39-flowered, 30-62 
mm long; peduncle 5-11 mm long; bracts 
narrowly triangular, convex, 8-9 mm long, c. 
2.5 mm wide, glabrous; bracteoles triangular, 
convex, 2-3 mm long, 1.5-2 mm wide. Male 
flowers: not fragrant, diameter 13-27.5 mm, 
pedicels 2.5-5 mm long; sepals 4, 2-whorled, 
orbicular, convex, may be slightly keeled; 
outer pair joined, with margin entire or 
irregularly lobed, 4-6 mm long, 7-8 m wide, 
green, glabrous; inner pair free, with margin 
recurved, entire or irregularly lobed, 4-8 mm 
long, 4-10 mm wide, pale green, glabrous; 
petals 4, free, orbicular or oblong-orbicular, 


Cooper, Garcinia in Australia 

convex, 8.5-15 mm long, 7.5-15 mm wide, 
white or cream on the inner surface and cream 
or pale yellow on the outer surface, glabrous; 
androecium of 4 fused phalanges creating a 
solitary receptacle, rhombic, adnate to petals 
and flower base, undivided or divided into 
4 shallow lobes at apex, diameter 5.5-11 
mm; anthers numerous, densely clustered, 
subsessile, strap-shaped or broadly wedge- 
shaped, bithecate, c. 2 mm long and 0.5 mm 
wide, glabrous; fungiform pistillode present 
in some flowers, c. 4.5 mm long, diameter 
at apex c. 2 mm. Female inflorescences 
terminal, a solitary flower (rarely pairs); 
bracteoles narrowly triangular, convex, c. 8.5 
mm long and 2.5 mm wide at base, glabrous, 
caducous. Females flowers: not fragrant, 
diameter 16-25 mm; pedicel c. 2 mm long; 
sepals 4, free, D-shaped, base truncate, apex 
obtuse, cream; outer pair narrowly joined, 
c. 6.3 mm long and 7.2 mm wide; inner pair 
free, c. 8.3 mm long and 10 mm wide; petals 
4, oblong-orbicular, convex, c. 13.5 mm 
long and 10.5 mm wide, cream; staminodal 
phalanges 4, adnate to petals, somewhat 
square, apex rounded, antherodes 5-8 per 
phalange, c. 1 mm diameter; gynoecium 
urn-shaped, c. 6 mm long; ovary ovoid or 
top-shaped, 6-8-locular; stigma domed, 
6-8-rayed, diameter c. 3.5 mm; ovules 1-4 
per locule. Fruiting pedicels c. 2 mm long; 
4 sepals persistent at base; fruit a semi-dry 
berry, top-shaped or broadly ovoid, mostly 
beaked, beak up to 20 mm long, 6-8 shallow 
septum-lobes or septum-lines, 60-72 mm 
long, 50-58 mm wide, green, dull (shiny 
when unripe), stigma remnants 6-8-rayed, 
diameter 5-6 mm; locules 6-8; septa thin, 
transparent, pithy and incomplete; seeds 1-3 
per locule, wedge-shaped, 20-22.5 mm long, 
18-23 mm wide; testa cream to brown; aril 
cream and netted, not fleshy. Fig. 3G & H, 4. 

Additional selected specimens (from 12 examined): 
Queensland. Cook District: Finlayvale, NW of 
Mossman, May 2012, Hawkes s.n. (CNS); Mossman 
Gorge NP on circuit track, Dec 1989, Jessup GJD3330, 
Guymer & Dillewaard (BRI); above the north bank of 
the Mossman River west of Mossman, Jan 2003, Jago 
6382, Russell & Darwin (BRI); N of Mossman River 
in Daintree NP, Dec 2002, Russell 11 (CNS); North of 
Mossman River, Daintree NP, Jul 2011, Cooper 2138, 
Russell, Jensen & Jago (CNS); ditto , May 2012, Cooper 
2149, 2150, 2151, Russell & Hawkes (CNS); ditto , Nov 


23 

2012, Cooper 2167, Russell, Ford & Sheather (CNS); 
ditto , Dec 2012, Cooper 2205, 2206, Russell & Sheather 
(CNS). 

Distribution and habitat : Garcinia russellii 
is endemic to the Wet Tropics bioregion of 
north-east Queensland where it is currently 
known to occur in rainforest (evergreen 
mesophyll vine forest) on granitic soils in 
the Mossman Gorge section of Daintree NP 
and at Finlayvale near Mossman (Map 4) at 
altitudes between 150 and 300 m. 

At the type locality Garcinia russellii 
grows as a slender understory tree in 
rainforest under a canopy of Backhousia 
bancroftii F.M.Bailey & F.Muell. ex 

F. M.Bailey, Flindersia bourjotiana F.Muell., 
Linds ay omyrtus racemoides (Greves) Craven, 
Hicksbeachia pilosa P.H.Weston & Calamus 
caryotoides A.Cunn. ex Mart. 

Phenology : Flowers have been recorded in 
May and July; fruits have been recorded in 
December. 

Notes : Male flowers within the same 
inflorescence can be inconsistent with some 
flowers having a fungiform pistillode and 
others not having any indication of a pistillode. 

The exudate-containing canals within the 
leaves of this species are indistinguishable 
from G. gibbsiae (Fig. 2E). 

Affinities : Garcinia russellii appears to be 
most similar to G. gibbsiae. Both species 
have tough coriaceous leaves with numerous 
parallel secondary veins and green fruit 
with up to three or four seeds within each 
locule. G. russellii is a slender and sparsely 
branched small slender tree with colourless, 
milky or white exudate, peltate or almost 
peltate leaves with distinctly cordate bases, 
male inflorescences are a 9- or more-flowered 
panicle and petals cream or white, fruit with 
seeds tightly packed within each locule which 
mostly cause the exocarp to bulge; locular 
septa thin, pithy and incomplete. Whereas, 

G. gibbsiae is a moderately to densely 
branched small to medium-sized tree, bright 
lime-yellow exudate, leaf bases which are 
either cuneate or obtuse and never peltate 
or cordate, male inflorescences solitary or a 
few-branched raceme and petals yellow; fruit 


24 


Austrobaileya 9(1): 1-29(2013) 


Fig. 4. Garcinia russellii fruit A. showing longitudinal cross-section and multi-carpelled locules, thin transparent 
septa and netted aril. B. showing beaked fruit, bracts and terminal attachment to branch (Cooper 2206, Russell & 
Sheather [CNS]). Photos: A, W.Cooper, B, T.Hawkes. 


with seeds usually in spacious locules; locular 
septa thick, fleshy and entire. 

Etymology : The specific epithet is in honour 
of naturalist and conservationist Rupert 
Russell (1939-). 

12. Garcinia warrenii F.Muell., Viet. Nat. 
8: 109 (1891). Type: Australia: Queensland. 
Cook District: Coen River, in 1891, S. 
Johnson s.n. (holo: MEL n.v.\ iso: BRI 
[AQ0340253 1 ], K 000677819 1 ). 

Garcinia kajewskii C.T.White, Contr. Arnold 
Arbor. 4: 72 (1933). Type: Queensland. Cook 
District: Daintree River, 30 November 1929, 
S.F.Kajewski 1418 (holo: BRI [AQ0340243 1 ]; 
iso: S 11-34563 1 ). 

Illustrations: Christophel & Hyland (1993: 
82); Cooper & Cooper (1994:288); Hyland et 
al. (2003); Cooper & Cooper (2004: 123). 

Tree to 25 m, dioecious; exudate yellow; dbh 
to 35 cm; bark tessellated and flakey in large 
trees, vertically fissured and with vertical 
lenticels; branchlets square orterete andmostly 
horizontal, glabrous. Leaves discolorous, 
coriaceous, glabrous; petioles 8-30 mm 
long, not channelled or becoming shallowly 


channelled in dry specimens, ligulate; lamina 
elliptic, oblong-elliptic, ovate, obovate, 
orbicular or oblanceolate, 40-230 mm long, 
28-123 mm wide; base attenuate, cuneate or 
obtuse; apex acuminate, acute, obtuse, retuse 
or emarginate; margin not recurved or rarely 
slightly recurved; venation brochidodromous, 
often with a second set of loops near margin, 
primary vein on upperside raised towards 
base and more or less flush towards apex; 
secondary veins 24-50, angle of divergence 
from primary vein 60-90°, forming an 
intramarginal vein 0.5-4 mm from margin; 
exudate-containing canals difficult to see on 
fresh leaves, distinct on underside of dried 
leaves as numerous unbranched or branched 
straight or wriggly + parallel lines, sometimes 
a second set of canals is visible as more 
widely spaced long straight lines running at 
angles 45-80° to primary vein and across the 
other canal set. Male inflorescences terminal 
or rarely axillary, a panicle, 8-66-flowered, 
15-120 mm long; peduncle 5-38 mm long; 
bracts (which sometimes become full- 
sized leaves) at peduncle apex, triangular, 
becoming incurved, glabrous, 8.5-10 mm 
long, 2-3 mm wide, green with a yellowish 
primary vein, caducous; bracteoles triangular, 


25 


Cooper, Garcinia in Australia 

incurved, 2.5-3 mm long, 2.5-3 mm wide, 
green, caducous. Male flowers: fragrant 
with a clove-like smell, diameter 10-15 mm; 
pedicels absent or up to 4 mm long; sepals 
4, 2-whorled, outer pair joined, D-shaped, 
convex, 4-5 mm long, 4-5mm wide, green; 
petals 4, free, erect, imbricate, orbicular or 
oblong-obovate, may be convex, 5-7 mm 
long, 4.5-6 mm wide, cream or yellow, 
glabrous, margin usually recurved and thin; 
androecium 4-phalangiate; phalanges adnate 
to petals, obovate-elliptic at anthesis, c. 3 
mm long, 2-2.5 mm wide; anthers several, 
most are clustered around phalange margin, 
bilobed, bithecate, c. 0.4 mm long and 0.2 
mm wide, glabrous; pistillode rudimentary, 
attenuate-conical, c. 2.5 mm long. Female 
inflorescences terminal, a solitary flower 
or a 2 or 3(rarely 4)-flowered raceme, 17-25 
mm long; peduncles 4-12 mm long; bracts at 
apex of peduncle triangular, keeled, 6-11 mm 
long, 3-5 mm wide, green with a yellowish 
primary vein, glabrous, caducous; bracteoles 
triangular, 3-4 mm long, 3-4 mm wide, 
green, caducous. Female flowers: fragrant 
with a clove-like smell, diameter 14-16 mm; 
pedicels absent or 3-5 mm long; sepals 4, 
2-whorled, outer pair joined, inner pair free, 
D-shaped, convex, 5-6.5 mm long, 8-9 mm 
wide, green, glabrous, margin thin; petals 
4, free, orbicular, convex, 9-11 mm long, 
10-12 mm wide, cream or yellow, sometimes 
blushed with pink, glabrous, margin thin; 
staminodal phalanges 4, erect, adnate to 
petals, strap-shaped, c. 5 mm long and 3.5 
mm wide, apex divided into 4-6 filaments 
with 1 or 2 antherodes on each side; filaments 
1-1.5 mm long; antherodes obovate or 
rhomboid, c. 1 mm long; gynoecium oblong- 
globose, c. 8.5 mm long and 6-8 mm wide; 
ovary globose or 4-sided, 7-8 mm long and 
6-8.5 mm wide, sessile, 4-locular, uniovulate; 
stigma mass sessile, domed, diameter 7-8 
mm, cream becoming pink or reddish with 
age or exposure. Fruiting pedicel 3-5 mm 
long, 4 sepals persistent at base; fruit a fleshy 
berry, globular or depressed globular, often 
shortly beaked, 40-58 mm long, 50-75 mm 
wide, yellow or dark red to blackish; stigma 
shallowly 3 or 4-lobed or rayed, diameter 
8-9.5 mm; seeds 1-4, 32-41 mm long, 23-35 


mm wide; testa brown, smooth; aril fleshy, 
dark red, purple or cream, native mangosteen, 
Warren’s mangosteen. Fig. 2G, 3F & I. 

Additional selected specimens (from 144 examined): 
Papua New Guinea. Western District: Tarara, Wasi 
Kussa River, Dec 1936, Brass 8430 (BRI). Australia. 
Northern Territory. Jessie River, Melville Island, Aug 
2000, Harwood907 (CNS). Queensland. Cook District: 
Gabba Island, Torres Strait, Mar 2003, Waterhouse 6592 
(BRI); lama (Yam) Island, Torres Strait, Nov 2007, Fell 
8855 & Stanton (BRI); Pajinka Lodge, Feb 2001, Cooper 
1453 & Jensen (CNS); Escarpment of the Great Dividing 
Range, 18.2 km northeast of Heathlands Ranger Base, 
Heathlands D and O Reserve, Oct 1993, Fell 3759, Stanton 
& Dibella (BRI); Wattle Hills, Sep 2002, Sankowsky 
1836 & Sankowsky (BRI); Stone Crossing, Wenlock 
River, Oct 1980, Hyland 1075 (CNS); Hey Point, Aug 
1985, Gunness 1899 (BRI); Aurukun, banks of Watson 
River, Mar 2001, Smith 4609 (BRI); Ling Creek Spring, 
Steve Irwin Wildlife Reserve, c. 16km due NE of Stone 
Crossing, Wenlock River, Sep 2009, McDonald 2009- 
09-034 & Addicott (BRI); 12.6 km WNW of Lockhart 
River Community, Mar 1994, Fell 4194 & Stanton (BRI); 
Nesbit River, Aug 1948, Brass 19915 (BRI); Rocky River 
Crossing, Oct 1996, Cooper 1011 & Cooper (CNS); Gap 
Creek, c. 38 km S by E of Cooktown, Aug 1959, Smith 
10744 (BRI); Pilgrim Sands, Cape Tribulation, Jan 1993, 
Cooper 474 & Cooper (CNS); Clacherty Road, Julatten, 
Aug 2012, Cooper 2158 & Hawkes (CNS); Euluma 
Creek Road, Julatten, Aug 2012, Cooper 2159 & Hawkes 
(CNS); SFR 843, Parish of Bellenden Ker, Frenchman 
Creek, Sep 1986, Gray 4333 (CNS); Babinda, Aug 2003, 
Jago 119 (CNS). 

Distribution and habitat: Garcinia warrenii 
occurs in Australia and New Guinea. 
Within Australia it is recorded from north 
Queensland where it occurs in rainforest 
(complex mesophyll vine forest, mesophyll 
vine forest, complex notophyll vine forest, 
notophyll vine forest and simple microphyll 
vine-fern forest) and vine thickets on granitic, 
basaltic and rarely metamorphic soils from 
the Torres Strait islands to Babinda, altitude 
0-1050 m. There are also two records from 
Melville Island, in the Northern Territory 
(Map 5) 

Phenology : Flowers have been recorded in 
July, August and September; fruit has been 
recorded in October, November, December, 
January and February. 

Etymology : The specific epithet is in honour 
of Dr William Henry Warren (1852-1926), 
Professor of Engineering at the University of 
Sydney. 


26 

Excluded Names 

Garcinia cherryi F.M.Bailey, Queensl. FI. 
6: 103 (1902). Basionym of Ternstroemia 
cherryi (F.M.Bailey) Merr. ex J.F.Bailey & 
C.T.White (Theaceae). 

Acknowledgements 

Christina Zdenek, Brian Venables and 
John Pritchard are sincerely thanked for 
making several trips to check for flowers 
from the female tree of Garcinia jensenii. I 
am especially grateful to Rupert Russell for 
showing me G. russellii in the field, where 
he initially discovered them and for assisting 
on numerous trips across the Mossman 
River during varying water levels to finally 
collect enough fertile material for a formal 
description. Andrew Ford, Rigel Jensen and 
Jeanette Kemp have been invaluable in the 
field for their good company and sources 
of expertise. I also thank Frank Zich and 
Darren Crayn for support and access to the 
Australian Tropical Herbarium (CNS); Paul 
Forster for the complicated typification for G. 
xanthochymus and notes on the typification of 
G. mestonii, as well as other assistance; Ben 
Trupperbaumer for old German translations 
and Peter Bostock for translation of Latin 
descriptions as well as creating the maps. 
Bill Cooper, David Fell, Mark Harrington, 
Mel Harrison, Tim Hawkes, Bob Jago, Sarah 
Legge, Stephen McKenna, Steve Murphy, 
Garry Sankowsky, Allen Sheather, Peter 
Stanton & Gary Wilson are thanked for much 
appreciated assistance in various forms. 
Darren Crayn made valuable suggestions to 
an earlier manuscript. The curators of BRI, 
CANB, MEL & NSW are thanked for loans 
and/or photos of relevant material. Permits 
to collect were issued by the Queensland 
Department of Environment and Heritage 
Protection. 

References 

Anderson, T. (1874). Guttiferae. In J.D. Hooker, Flora 
of British India 1: 258-278. L.Reeve & Co.: 
London. 

Bailey, F.M. (1889). Report on new plants, preliminary to 
general report on botanical results on Meston’s 
expedition to the Bellenden-Ker Range, by 
F.M. Bailey, F.L.S., Colonial Botanist , p. 2. 
Department of Agriculture: Brisbane. 


Austrobaileya 9(1): 1-29 (2013) 

- (1899). Guttiferae. In The Queensland Flora , 

Part 1, pp. 102-105. H.J.Diddams & Co: 
Brisbane. 

- (1902). The Queensland Flora , Part 6, p. 103. 

H.J.Diddams & Co: Brisbane. 

Canbr (Centre for Australian National Biodiversity 
Research) (2010). Australian Tropical Rain 
Forest Plants Edition 6 (online version), viewed 
20 June 2012. http://www.anbg.gov.au/cpbr/cd- 
key s/rfk/index. html 

Chah (2007). Australian Plant Census. http://www. 
anbg.gov.au/cgi-bin/apclist; Australian Plant 
Names Index, http://www.anbg.gov.au/cgi-bin/ 
apni Accessed November 2011. 

Christophel, D.C. & Hyland, B.P.M (1993). Leaf 
Atlas of Australian Rain Forest Trees. CSIRO 
Publishing: Melbourne. 

Cooper, W. & Cooper, W.T. (1994). Fruits of the Rain 
Forest. Geo: Sydney. 

- (2004). Fruits of the Australian Tropical 

Rainforest. Nokomis Editions: Melbourne. 

D’arcy, W.G. (1980). Flora of Panama. Family 123. 
Guttiferae. Annals of the Missouri Botanical 
Garden 67: 969-1043. 

De Candolle, A.P (1824). Guttiferae. In Prodromus 
Systematis Naturalis Regni Vegetabilis 1: 557- 
564. Treuttel & Wtirtz: Paris. 

Dowe, J.L. & Broughton, A.D. (2007). F.M.Bailey’s 
ascent of Mt Bellenden-Ker in 1889, and notes 
on the publication priority of new vascular plant 
species from the Expedition. Austrobaileya 7: 
555-566. 

- (2008). F.M.Bailey’s taxonomy of A.Meston’s 

collections from the Bellenden Ker Expedition 
of 1904. Austrobaileya 7: 677-679. 

Fleming, (circa 1795-1805). Fleming Indian Drawings 
Collection circa 1795-1805, Natural History 
Museum, London/Science Photo Library http:// 
www.sciencephoto.com/media/428689/enlarge 
Accessed July 2013 

Hammel, B. (1993). In C.E. Jarvis, F.R. 
Barrie, DM. Allan & J.L. Reveal. 
A list of Linnean generic names and their types. 
Regnum Vegetabile 127: 28. 

Hooker, W.J. (1831). Xanthochymus dulcis. Sweet- 
fruited Xanthochymus. Curtis’s Botanical 
Magazine 58: t. 3088. 

- (1855). Garcinia mangostana. Mangosteen, or 

Mangostan. Curtis’s Botanical Magazine 81: t. 
4847. 

Hyland, B.P.M. (1982). A revised card key to rainforest 
trees of North Queensland. CSIRO: Melbourne. 


27 


Cooper, Garcinia in Australia 

Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray, 
B. & Elick, R.W. (2003). Australian Tropical 
Rain Forest Plants. Trees, Shrubs and Vines. 
CD-ROM. CSIRO Publishing: Melbourne. 

Hyland, B.P.M., Whiffin, T., Zich, F. Edition 6. (2010). 
Australian Tropical Rain Forest Plants, CSIRO. 
http://www. anbg.gov.au/cpbr/cd-keys/rfk/ 
index.html Accessed October 2011. 

Jessup, L.W. (2007). Clusiaceae. In P.D. Bostock & A.E. 
Holland (eds.). Census of the Queensland Flora 
2007, p. 54. Environmental Protection Agency, 
Brisbane. 

- (2010). Clusiaceae. In P.D. Bostock & A.E. 

Holland (eds.). Census of 

the Queensland Flora 2010, p. 44. 

Environmental Protection Agency: Brisbane. 

Kirtikar, K.R. & Basu, B.D. (1918). Indian Medicinal 
Plants. Sudhinra Nathbasu: Bahadurganj. 

Kostermans, A.J.G.H. (1980). Clusiaceae (Guttiferae). 
In M.D. Dassanayake (ed.), Flora of Ceylon 1: 
72-110. Amerind Publishing Co. Pvt. Ltd.: New 
Delhi. 

Maheshwari, J.K. (1964). Taxonomic studies on Indian 
Guttiferae III. The genus Garcinia L. s.l. 
Bulletin of the Botanical Survey of India 6: 
107-135. 

Marsden, W. (2005). The History of Sumatra containing 
an Account of the Government, Laws, Customs 
and Manners of the Native Inhabitants. 
ebook #16768 http://www.gutenberg.org/ 
files/16768/16768-h/16768-h.htm#sumatra-03 
Accessed July 2013. 

Moore, S. (1917). A contribution to the phytogeography 
of Bellenden-Ker. II. Systematic account. 
Journal of Botany 55: 302-310. 

Mueller, F.J.H. (1891). Descriptions of new Australian 
plants with occasional other annotations 
(continued). The Victorian Naturalist 8(7): 109. 

Nazre, M. (2006). Taxonomic Revision and Molecular 
Studies of Garcinia Section Garcinia 
(Guttiferae ), unpublished PhD thesis. The 
University of Edinburgh. Opara, L.U., 
Studman, C.J. & Banks, N.H. (1997). Fruit 
Skin Splitting and Cracking. In J. Janick (ed.), 
Horticultural Reviews 19(4): 227. 

Richards, A.J. (1990a). Studies in Garcinia , dioecious 
tropical forest trees: agamospermy. Botanical 
Journal of the Linnean Society 103: 233-250. 

- (1990b). Studies in Garcinia , dioecious tropical 

forest trees: the origin of mangosteen. Botanical 
Journal of the Linnean Society 103: 301-308. 

Richards, J. (2008). The secret war: a true history of 
Queensland’s Native Police. University of 
Queensland Press: St Lucia. 


Robson, N.K.B. (1960). Guttiferae. In A.W. Excell & H. 
Wild (eds.). Flora Zambesiaca 1(2): 378-404. 
Crown Agents for Overseas Governments & 
Administrations: London. 

Roxburgh, W. (1805). Plants of the Coast of Coromandel 
2: 51, t. 196. W. Bulmer 
and Co.: London. 

- (1820). Plants of the Coast of Coromandel 3: 66. 

W. Bulmer and Co.: London. 

Sharma, B.P., Singh, T.D., Handique, P.J., Devi, H.S. 
(2012). Unambiguous identification of Garcinia 
xanthochymus (Clusiaceae) in Manipur, India. 
Nebio 3(3): 7-10. 

Singh, N.P (1993). Clusiaceae. In B.D. Sharma & 
M. Sanjappa (eds.). Flora of India 86-159. 
Botanical Survey of India: Calcutta. 

Sompong, T. (2007). Floral and fruit morphology of 
some species in Garcinia spp. Songklanakarin 
Journal of Science and Technology 29: 245- 
252. 

Stevens, P.F. (2007). Clusiaceae - Guttiferae. In K. 
Kubitzki (ed.). The Families and Genera of 
Vascular Plants. 9: 48-66. Springer-Verlag: 
Berlin/Heidelberg. 

Sweeney, P.W. (2008). Phylogeny and floral diversity in 
the genus Garcinia (Clusiaceae) and relatives. 
International Journal of Plant Sciences 
169:1288-1303. 

- (2010). Floral anatomy in Garcinia nervosa 

and G. xanthochymus (Clusiaceae): A first step 
toward understanding the nature of nectaries in 
Garcinia. Bulletin of the Peabody Museum of 
Natural'History 51: 157-168. 

Te-chato, S. (2007). Floral and fruit morphology of 
some species in Garcinia spp. Songklanakarin 
Journal of Science Technology 29: 245-252. 

van Nooten, B.H. (1880). Fleurs, fruits et feuillages 
choisis de l ’ille de Java: peintes d ’apres nature. 
C.Muquardt: Bruxelles. 

White, C.T. (1933). Ligneous plants collected for the 
Arnold Arboretum in north Queensland by 
S.F. Kajewski in 1929. Contributions from the 
Arnold Arboretum of Harvard University 4: 
1-113. 

- (1936). Contribution to the Queensland flora. 

No. 5. Proceedings of the Royal Society of 
Queensland 47: 51-84. 

Wight, R. (1840). Xanthochymus dulcis. leones 
Plantarum Indiae Orientalis. 1: t. 192. 
J.B.Pharoah: Madras. 

Zodiac Publications (undated). Australian Tropical 
Plants version 4.1 DVD ROM. Zodiac 
Publications. 


28 


Austrobaileya 9(1): 1-29(2013) 


Map 1. Distribution of Garcinia dulcis • (within 
Australia), G. brassii A and *G. livingstonei M. 


Map 3. Distribution of Garcinia leggeae A, G. mestonii 
■ and *G. mangostana •. 


Map 2. Distribution of Garcinia jensenii A and G. 
zichii •. 


Map 4. Distribution of Garcinia gibbsiae A and G. 
russellii ■. 


Cooper, Garcinia in Australia 


29 


Map 5. Distribution of Garcinia warrenii • (within Australia). 


Map 6. Distribution of *Garcinia xanthochymus •. 


Reinstatement and revision of Sphaeromorphaea DC. 
and Ethuliopsis F.Muell. (Asteraceae: Plucheinae) 


A.R. Bean 
Summary 

Bean, A.R. (2013). Reinstatement and revision of Sphaeromorphaea DC. and Ethuliopsis F.Muell. 
(Asteraceae: Plucheinae). Austrobaileya 9(1): 30-59. Sphaeromorphaea DC. and Ethuliopsis 
F.Muell., historically included in Epaltes Cass., are reinstated and taxonomically revised. 
Sphaeromorphaea occurs in Australia, Indian Subcontinent, Indochina, China, eastern Asia, 
Malesia, Papuasia, northwestern Pacific and southwestern Pacific, and comprises six species. Three 
new species; S. ephemera A.R.Bean, S. major A.R.Bean and S. subintegra A.R.Bean are described, 
and two new combinations; S. harrisii (F.Muell.) A.R.Bean and S. littoralis (Retz.) A.R.Bean are 
made. Ethuliopsis occurs only in Australia and comprises the single species E. cunninghamii (Hook.) 
F.Muell. A neotype is chosen for Epaltes australis Less., and lectotypes are chosen for Artemisia 
littoralis Retz., Ethulia cunninghamii Hook, and Epaltes harrisii F.Muell. Descriptions, illustrations 
and distribution maps are provided for all taxa. Diagnostic morphological characters and ecological 
aspects are discussed, and identification keys are provided to the species and to the Australian genera 
of Subtribe Plucheinae. 

Key Words: Asteraceae, Plucheinae, Epaltes, Ethuliopsis, Sphaeromorphaea, Sphaeromorphaea 
ephemera, Sphaeromorphaea harrisii, Sphaeromorphaea littoralis, Sphaeromorphaea major, 
Sphaeromorphaea subintegra, Australia flora, India flora, China flora, taxonomy, new species, 
identification key, distribution maps 

A.R. Bean, Queensland Herbarium, Department of Science, Information Technology, Innovation 
and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. 
E-mail: Tony.Bean@science.dsitia.qld.gov.au 


Introduction 

Sphaeromorphaea DC. and Ethuliopsis 
F.Muell. are often regarded as synonyms of 
Epaltes Cass. (Cooke 1986; Bremer 1994; 
Anderberg & Eldenas 2007; GRIN 2012) and 
their histories are intertwined. 

Cassini (1818) named the genus Epaltes 
with a single species Epaltes divaricata (L.) 
Cass., native to the Indian subcontinent, 
China and Indochina. C.F. Lessing added 
Epaltes australis Less., E. mexicana Less, 
and E. hirsuta Less, in 1830 and 1832. In the 
ensuing century, nearly 20 Epaltes species 
were described from all tropical parts of the 
world. 

Sphaeromorphaea was named by De 
Candolle (1838) with three species at that 
time, two originating from southern Asia and 
one from Australia. No other species were 
subsequently described under that generic 


Accepted for publication 16 July 2013 


name; however, the combination S. australis 
(Less.) Kitam., based on Epaltes australis 
Less., was made in 1936. 

In Australia, Mueller (1861) proposed 
a new genus, Ethuliopsis , to accommodate 
Ethulia cunninghamii Hook. Bentham (1867) 
relegated Ethuliopsis to synonymy with 
Epaltes , while maintaining Epaltes australis ; 
the two species he placed in Epaltes (E. 
australis , E. cunninghamii (Hook.) Benth.) 
were to remain there for well over a century. 

Bentham (1873) dealt incompletely with 
Sphaeromorphaea , saying correctly only that 
it is, in part, synonymous with Centipeda 
Lour. One of the three named species, S. 
centipeda DC., is a nomenclatural synonym 
of Centipeda minima (L.) A.Braun & Asch. 
Hooker (1882) accepted Sphaeromorphaea 
with a single species (for India), S. russeliana 
DC., and placed the genus next to Centipeda 
in his systematic arrangement. 


Bean, Sphaeromorphaea and Ethuliopsis 

Mueller described three further species 
under Epaltes (E. harrisii F.Muell., E. tatei 
F.Muell. and E. pleiochaeta F.Muell.) from 
Australia; Moore & Betche (1893) listed 
E. pleiochaeta as an accepted species, and 
Bailey (1900) listed E. harrisii as an accepted 
species, but neither name ever came into 
general usage. Epaltes tatei is an accepted 
species that was transferred to a new genus, 
Haegiela P.S.Short & Paul G.Wilson (Short 
& Wilson 1990), belonging to the Tribe 
Gnaphalieae. 

Contemporary Asian flora treatments 
have used either Epaltes australis or 
Sphaeromorphaea australis for the taxa 
found there. For Taiwan, Li (1978) accepted 
S. australis , but Peng (2004) reversed this, 
with Epaltes australis then the accepted 
name; Naithani (1995) accepted S. australis , 
placing the genus near Centipeda in the 
Tribe Anthemideae\ Hu & Wong (2009) used 
E. australis as the accepted name, with S. 
australis given as a synonym. 

In recent decades Epaltes has been regarded 
as a heterogenous genus incorporating any 
Plucheoid taxa whose achenes lack pappus 
bristles (Leins 1971; Merxmueller et al. 1978; 
Cooke 1986; Anderberg 1991). The present 
author agrees that Epaltes is paraphyletic, and 
that the Australian species currently referred 
to that genus belong in Sphaeromorphaea 
DC. or Ethuliopsis F.Muell. 

Dunlop (2000) was the first author to 
accept the genus name Sphaeromorphaea (and 
the species S. australis ) in an Australian flora 
treatment and some major Australian herbaria 
subsequently adopted Sphaeromorphaea 
for their state in plant checklists or censuses 
(Walsh & Stajsic 2007; Short et al. 2011). 
Short & Wilson (1990) foreshadowed the 
acceptance of the genus name Ethuliopsis , 
but the Australian Plant Census (APC 2012) 
continued to list it as a synonym of Epaltes. 

Sphaeromorphaea and Ethuliopsis 
are revised in this paper. Six species of 
Sphaeromorphaea are enumerated, including 
three new species and two new combinations; 
all of these species have previously been 
referred to a broadly circumscribed Epaltes 


31 

australis. They occur in Australia, Indian 
subcontinent, Indochina, China, eastern 
Asia, Malesia, Papuasia, northwestern Pacific 
and southwestern Pacific. Ethuliopsis is 
monotypic, and is endemic to Australia. 

Materials and methods 

This revision is based on a morphological 
study of herbarium specimens from A, 
AD, AMES, BM, BRI, C, CANB, GH, K, 
LD, MEL, PE, PERTH and TAI. Images of 
specimens, mainly types, from BM, C, CAL, 
G-DC, K, M and P have been examined. In 
addition, the author has collected and studied 
plants in the field in Queensland, New South 
Wales and the Northern Territory. 

All measurements were made from 
dried material, using a stereo microscope 
equipped with a graticule. Commonly used 
abbreviations in the specimen citations are 
HS for Homestead, NP (National Park), SF 
(State Forest). 

The distribution maps were compiled 
using DIVA-GIS Version 7.5.0, using label 
data of specimens from the herbaria listed 
above. 

Geographical regions are listed according 
to the scheme of Brummitt (2001); for the 
generic distributions, the areas cited are 
Regional Names; for the specimen citations, 
the areas cited are Level 3 Names. Specimen 
citations within Queensland are further 
subdivided into Pastoral districts. 

Relationships 

Anderberg (1991) provided a cladistic 
analysis of the subtribe Plucheinae based 
on morphological characters. He reinstated 
the African genus Litogyne Harv., and 
transferred Epaltes gariepina (DC.) Steetz to 
it, based mainly on the very different shape 
of the receptacle. Litogyne has since gained 
some acceptance as a distinct genus (Retief & 
Herman 1997; Beentje 2002). 

Anderberg (1991) studied material of the 
following Epaltes species: E. australis , E. 
brasiliensis (Link) DC., E. cunninghamii 
and E. mattfeldii. Interpretation of his 
data is fraught because E. australis is here 


32 

regarded as belonging to Sphaeromorphaea, 

E. cunninghamii is regarded as belonging 
to Ethuliopsis , and the generic status of E. 
brasiliensis and E. mattfeldii is uncertain. 
The type species of Epaltes (E. divaricata ) 
was not included in the analysis. 

Anderberg (1991) placed Thespidium 

F. Muell. ex Benth. and Coleocoma F.Muell. 
in the ‘Coleocoma group’, while Epaltes was 
placed in the ‘Pluchea group’. The present 
author is of the opinion that while Epaltes 
sens. str. and Ethuliopsis probably do belong 
in the ‘Pluchea group’, Sphaeromorphaea 
belongs to the ‘Coleocoma group’ and is 
most closely related to the monotypic genus 
Thespidium. The persistent ebarbellate 
bristles formed on the achenes of some 
Sphaeromorphaea spp. are very similar to 
those of Thespidium basiflorum (F.Muell.) 
Benth., and the cylindrical longitudinally- 
ribbed achenes are a feature of both genera. 


Austrobaileya 9(1): 30-59 (2013) 

Leins (1971) had previously proposed a close 
relationship between Epaltes australis and 
Thespidium based on a study of the pollen 
grains and style morphology. 

The molecular study of Anderberg et al. 
(2005) placed Coleocoma , Streptoglossa 
Steetz and Epaltes cunninghamii (i.e. 
Ethuliopsis) as part of a largely unresolved 
clade dominated by species of Pluchea Cass. 

The presence of a coronal pappus 
is a uniting character for all species of 
Sphaeromorphaea and Ethidiopsis. Coronal 
pappus is not common in Asteraceae, being 
confined to some genera in Tribe Anthemideae 
and Tribe Inuleae (Heywood & Humphries 
1977; Mukherjee & Sarkar 2001). 

The species of Sphaeromorphaea and 
Ethuliopsis differ from Epaltes sens. str. (E. 
divaricata) in a number of significant ways 

(Tables 1 & 2): 


Table 1. Morphological character differences between Sphaeromorphaea and Epaltes 
divaricata 


Characters 

Sph aeromorph aea 

Epaltes divaricata 

stems 

not winged 

winged 

capitula 

borne usually in pairs 

borne singly 

involucral bracts 

mostly obtuse, cartilaginous, 
incurved 

all acute, herbaceous, straight or 
excurved 

marginal florets 

conical or lageniform, or 
filiform with an expanded 
base 

filiform, not significantly 
expanded at the base 

style of disc florets 

rather slender, not divided 

style arms short and broad, 
distinctly divided 

pappus 

a stiff persistent corona, 
bristles sometimes present 

no corona, no bristles 

achenes of disc florets 

full sized, often 
indistinguishable from 
marginal-floret achenes 

not formed; ovary vestigial 

achene shape 

cylindrical 

ellipsoidal to obovoid 

carpophore 

prominent white ring 

small, obscure 


Bean, Sphaeromorphaea and Ethuliopsis 33 

Table 2. Morphological character differences between Ethuliopsis and Epaltes divaricata 


Characters 

Ethuliopsis 

Epaltes divaricata 

stems 

not winged 

winged 

sexuality 

subdioecious 

bisexual 

receptacle 

hemispherical (in 
heterogamous capitula) 

flat 

style of disc florets 

rather slender, not divided 

style arms short and 
broad, distinctly divided 

pappus 

a persistent cylindrical 
corona, bristles present 
on disc florets 

no corona, no bristles 

achene shape and ribbing 

lunate; 1-ribbed 

ellipsoidal to obovoid; 
several-ribbed 

carpophore 

prominent white ring 

small, obscure 


In Ethuliopsis , some plants bear 
homogamous capitula (with disc florets only), 
while other plants bear heterogamous capitula 
(nearly all marginal florets, but with a few 
disc florets). In Epaltes divaricata , all capitula 
on all plants are heterogamous. 

Ecology and distribution 

Ethuliopsis cunninghamii is an erect annual 
herb reaching one metre in height. It is found 
on alluvial flats with heavy clay soils, in 
swamps, or beside river channels and lakes. 
It is endemic to arid and semi-arid eastern 
Australia. 

All Sphaeromorphaea species are herbs 
or small shrubs 15-60 cm high or wide. 
Some species are seemingly annual (e.g. S. 
ephemera ), while other species can develop a 
substantial taproot and probably live for two or 
three years, with older stems dying, replaced 
by new stems shooting from near the base of 
the plant. They prefer sunny areas with bare 
soil in grasslands, sedgelands, woodlands 
or forests. They appear to have no edaphic 
preference, occurring on soils as diverse as 
pure beach sand and heavy cracking clays. 

All Sphaeromorphaea species inhabit 
places where the soil is intermittently or 
permanently moist or wet. Poor drainage is 


readily tolerated, but they will not persist 
where there is standing water. They can most 
often be found on alluvial flats, in swamps, 
or beside rivers and lakes; however, some 
species can also occur in hilly terrain, where 
they occupy the wetter microhabitats, such as 
depressions, drains and gullies. 

Most species display a considerable 
tolerance for high levels of salt in the soil or 
in the air. One species (Sphaeromorphaea 
major A.R.Bean) is confined to marine couch 
grasslands, and most other species sometimes 
inhabit salt affected areas. S. australis can 
grow on coastal headlands, exposed to salty 
winds. Label data suggest that S. subintegra 
A.R.Bean sometimes grows on serpentinite, 
a substrate well known to be toxic to many 
plant species. 

Waterbirds are well known as dispersal 
agents for the seeds of wetland plants (e.g. 
Figuerola & Green 2002; Soons et al. 2008), 
and although there is no literature documenting 
the dispersal of Sphaeromorphaea or 
Ethuliopsis , it seems reasonable to assume 
that waterbirds play a major role. This would 
help explain the distribution of S. littoralis , 
which is widespread from southern Australia 
to southern Asia. Its apparent absence 
from Indonesia may be partly due to the 


34 

predominance of rainforest communities that 
are not suitable habitat for the light-loving 
Sphaeromorphaea. 

It is interesting to note that Sphaeromorphaea 
and the ecologically comparable genus 
Centipeda (taxonomy revised by Walsh 
[2001]) have similar distribution patterns, to 
the extent of possessing a single widespread 
species shared by southern Asia and Australia, 
and a centre of diversity in Australia. 

Some diagnostic morphological characters 

1. Resin globules 

Ethuliopsis and most species of 
Sphaeromorphaea have resin globules on the 
lower and often on the upper leaf surfaces, and 
on the corollas and achenes. They are shining, 
transparent and globose, but may be difficult 
to see in very old herbarium specimens, as 
with age they degrade and deflate. Their 
presence, distribution and sometimes their 
numbers are diagnostic. S. major does not 
have resin globules. 

2. Leaf and stem indumentum 

Sphaeromorphaea major has glabrous leaves 
and stems. All other species bear crisped, 
multicellular, eglandular hairs, at least on 
young developing leaves and stems. For S. 
ephemera and Ethuliopsis cunninghamii, 
leaves are glabrous at maturity; S. australis 
may be glabrous or sparsely hairy. For S. 
littoralis, S. harrisii and S. subintegra, at least 
some hairs persist on mature leaves. Leaf 
indumentum is particularly variable in S. 
littoralis and S. harrisii, sometimes sparsely 
hairy and sometimes densely hairy on fully 
expanded leaves. 

3. Leaf shape and margins 

Sphaeromorphaea species have leaves that 
at first glance appear petiolate, but they are 
in fact sessile, as the lamina does extend to 
the stem in all taxa. In most species, the leaf 
shape is more or less obovate, but S. major 
is distinguishable by being especially narrow¬ 
leaved. No species has entire margins; in S. 
subintegra the margins are denticulate with 
teeth less than 0.5 mm long, and hence may 
appear entire without close inspection. In 


Austrobaileya 9(1): 30-59 (2013) 

S. australis, some large teeth or lobes (>1 
mm long) are usually present, often on the 
basal part of the lamina (the pseudo-petiole). 
Sphaeromorphaea littoralis and S. harrisii 
similarly are often conspicuously toothed on 
some leaves, with the teeth extending to the 
basal half of the lamina. 

Ethuliopsis cunninghamii leaves are not 
pseudo-petiolate, and the lamina is broad 
almost to the base. The leaf margins are 
denticulate to dentate. 

4. Marginal floret shape 

The shape of the marginal florets is 
an important diagnostic character for 
Sphaeromorphaea. In S. littoralis, S. 
subintegra and S. ephemera, the corolla is 
consistently conical, tapering more or less 
evenly from the base to the very narrow apex 
(e.g. Fig. 3E); in S. australis and S. harrisii, 
the corolla is consistently lageniform i.e. 
swollen in appearance, broadest above the 
base, then tapering rapidly towards the apex 
(Figs. 1G, 2C); in S. major and Ethuliopsis 
cunninghamii, the corolla of the marginal 
florets is filiform (Figs. 1C, 4H). 

5. Achene features 

All Sphaeromorphaea achenes are 
cylindrical, providing an instant distinction 
from Ethuliopsis whose achenes are 
lunate. Ethuliopsis achenes have a single 
adaxial longitudinal rib, although several 
anastomosing veins on the surface are often 
also visible. The number of longitudinal ribs 
for Sphaeromorphaea species varies from 
5-14, and is moderately diagnostic for each 
species. In S. littoralis and S. harrisii, there is 
a whorl of antrorse to appressed twin hairs at 
the base of the achene, with a few twin hairs 
sometimes occurring also on the achene body. 
In other species, the twin hairs are nearly 
always absent, although S. australis and S. 
subintegra may occasionally have 1-4 twin 
hairs at the achene base. 

6. Pappus corona 

All species of Sphaeromorphaea and 
Ethuliopsis have a stiff persistent coronal 
pappus. In Ethuliopsis cunninghamii and 
in Sphaeromorphaea major, the corona is 


Bean, Sphaeromorphaea and Ethuliopsis 

erect and shortly cylindrical with an entire or 
fimbriate margin, while in the other species 
of Sphaeromorphaea , it is annular or disc¬ 
like, transverse or nearly so, with an entire, 
pentagonal or erose margin (Fig. 3G). The 
width (and therefore the visibility) of this 
corona varies with the species; in S. littoralis , 
S. harrisii and S. subintegra it is relatively 
broad and easy to see, while in S. australis and 
S. ephemera , it is very narrow and difficult to 
detect. 

7. Pappus bristles 

In some species of Sphaeromorphaea 
pappus bristles emerge from the margin of 
the corona. They are consistently ascending, 
terete, persistent, and shorter than the corolla. 
With the exception of S. major , they lack 
barbellae. Sometimes only a single bristle is 
produced, but there may be as many as five. 
The length and number of bristles produced 
may differ according to the type of floret; 
S. major can produce up to five bristles on 
the disc achenes, but not more than three on 
the marginal achenes. Pappus bristles occur 
frequently in S. harrisii and S. major , but 
only rarely in S. littoralis and S. subintegra ; 
they have not been observed for S. australis 
and S. ephemera. Ethuliopsis consistently 
produces slightly flattened barbellate bristles 
on the disc florets, but never any bristles on 
the achenes. 

In summary, Sphaeromorphaea is 
characterised by the hemispherical to oblate 
capitula borne singly or in pairs opposite 
a leaf; the cartilaginous incurved glabrous 
involucral bracts; the presence of a coronal 
pappus (sometimes produced into slender 
persistent bristles); the cylindrical achenes, 
formed from all florets (though achenes 
produced from disc florets may not be fertile), 
glabrous or with straight twin hairs mainly at 
the base. Ethuliopsis is characterised by the 
subdioecious habit; the vestigial ovary of the 
disc florets; white corollas; barbellate pappus 
bristles on the disc florets; and the glabrous 
one-ribbed lunate achenes with a cylindrical 
coronal pappus, obliquely placed on the 
achene. 


35 

Taxonomy 

Sphaeromorphaea DC., Prodr. 6: 140 (1838). 
Type: S. russeliana, fide J.R. Drummond in 
Merrill & Rolfe (1908: 126) [=S. littoralis ]. 

Annual or perennial herbs with a well 
developed rootstock. Latex absent. 
Indumentum of crisped, multicellular, 
eglandular hairs. Sessile to very shortly 
stalked resin globules frequently present 
on stems, leaves, involucral bracts, corolla 
and achenes, initially shiny and transparent, 
becoming opaque and deflated with age. 
Stems terete, not winged. Leaves alternate, 
sessile, spreading; margins denticulate, 
dentate or lyrate; base attenuate, not 
decurrent. Capitula axillary or leaf-opposed, 
solitary or in pairs on often short peduncles, 
hemispherical to depressed-globose, 
disciform, heterogamous. Involucral bracts 
subequal in length, cartilaginous, incurved, 
in 3-4 rows, innermost row conspicuously 
narrower. Receptacle flat, glabrous, without 
paleae. Marginal florets fertile, female, 
without rays; corolla without multicellular 
hairs, resin globules present or absent, white 
or pink to purple at anthesis; lobes 3, tiny; 
style with bulbous base, branches divergent. 
Disc florets bisexual, usually producing 
achenes, but in some species infertile; corolla 
cylindrical to narrowly campanulate, without 
multicellular hairs, resin globules present or 
absent, pink to maroon; lobes 4 or 5; anthers 
tailed, apically obtuse; style bulbous at the 
base, divided or undivided at apex, sweeping 
hairs obtuse and extending well down the 
shaft. Achenes cylindrical, longitudinally 
ribbed; carpopodium ring-like, prominent, 
white. Pappus comprising a stiff, transverse 
or erect annular corona, sometimes produced 
into short slender persistent bristles. Twin 
hairs (when present) straight, antrorse to 
appressed, in a basal whorl, and occasionally 
on body of achene. 

Six species, occurring in Australia, Indian 
subcontinent, Indochina, China, eastern Asia, 
Malesia, Papuasia, northwestern Pacific and 
southwestern Pacific. 


36 


Austrobaileya 9(1): 30-59 (2013) 

Key to the species of Sphaeromorphaea using microscopic characters 

1 Corolla of marginal florets lageniform.2 

1. Corolla of marginal florets conical or filiform.3 

2 Coronal pappus 0.02-0.05 mm wide, obscure; leaves glabrescent; marginal 

florets with several resin globules; pappus bristles always absent; achene 

hairs usually absent, or rarely 1-4 present at base.2. S. australis 

2. Coronal pappus 0.05-0.08 mm wide, obvious; leaves persistently hairy; 

marginal florets with few or no resin globules; pappus bristles frequently 

present; hairs 5-20 at base of achene.3. S. harrisii 

3 Fully expanded leaves sparsely to densely hairy; achenes with several resin globules ... 4 

3. Fully expanded leaves glabrous; achenes without resin globules.5 

4 Leaves dentate or denticulate, teeth present on basal half of leaf; peduncles 

2-14 mm long; achenes with 10-40 hairs, mainly at the base.5. S. littoralis 

4. Leaves denticulate, teeth absent from basal half of leaf; peduncles 1.5-3.5 

mm long; achene hairs usually absent, or rarely 1-4 present at base . . . 6. S. subintegra 

5 Pappus bristles absent; young stems and leaves sparsely hairy; fertile 

achenes 0.45-0.7 mm long.4. S. ephemera 

5. Pappus bristles present on some florets; all stems and leaves glabrous; 

fertile achenes 1.0-1.3 mm long.1. S. major 

Field key to the species of Sphaeromorphaea 

1 Leaves narrow, 5.5-14 times longer than broad, without resin globules.1. S. major 

1. Leaves broader, 2.3-5.3 times longer than broad, resin globules present.2 

2 Older leaves glabrous, bright green.3 

2. Older leaves sparsely to densely hairy, green to grey-green.4 

3 Annual; achenes without resin globules, marginal florets conical.4. S. ephemera 

3. Perennial; achenes with resin globules, marginal florets lageniform.2. S. australis 

4 Teeth on leaf margin very small on all leaves and absent from basal half. . 6. S. subintegra 

4. Teeth on leaf margin > 1 mm long on some leaves, teeth present on basal half.5 

5 Most stems erect; capitula almost sessile; pappus bristles frequently 

present; leaves deeply and irregularly lobed.3. S. harrisii 

5. Most stems sprawling; capitula conspicuously pedunculate; pappus 

bristles rarely present; leaves shortly lobed, lobes fairly regular.6 

6 Leaves grey-green; achenes with numerous twin hairs at base and with 

conspicuous corona.5. S. littoralis 

6. Leaves bright green; achenes lacking twin hairs at base (or just a few 

present) and with very narrow corona.2. S. australis 


Bean, Sphaeromorphaea and Ethuliopsis 37 

Diagnostic characters for Sphaeromorphaea species 


Achenes with 5^10 twin hairs, mainly at base 

S. harrisii, S. littoralis 

Achenes with 1-4 twin hairs at base, or twin 
hairs absent 

S. australis, S. ephemera, S. subintegra, 

S. major 

Achene ribs 5-10 

S. australis, S. harrisii, S. littoralis, S. 
major 

Achene ribs 10-14 

S. australis, S. ephemera, S. littoralis, S. 
subintegra 

Corona transverse, prominent, >0.05 mm wide 

S. harrisii, S. littoralis, S. subintegra 

Corona transverse, obscure, < 0.05 mm wide 

S. australis, S. ephemera 

Corona erect, entire or fringed 

S. major 

Marginal floret lageniform 

S. australis, S. harrisii 

Marginal floret conical 

S. ephemera, S. littoralis, S. subintegra 

Marginal floret filiform 

S. major 

Pappus bristles frequently present 

S. harrisii, S. major 

Pappus bristles rarely present 

S. littoralis, S. subintegra 

Pappus bristles absent 

S. australis, S. ephemera 


1. Sphaeromorphaea major A.R.Bean 
sp. nov. affinis S. anstrali sed acheniis 
longioribus costis longitudinalibus 
paucioribus, flosculorum discoideorum 
corolla 2-2.5 mm longa (in S. australi 
1-1.3 mm) et foliis acheniis corollisque 
globulis resinae carentibus differens. Typus: 
Queensland. Port Curtis District: 100 
metres from Huttonvale track, SE of Sabina 
Point, Shoalwater Bay Training Area, 8 April 
2011, A.R. Bean 30830 & D. Halford (holo: 
BRI [1 sheet + spirit]; iso: BM, CANB, MEL, 
MO, distribuendi). 

Prostrate or procumbent perennial shrub to 30 
cm high and 60 cm across. Stems glabrous. 
Leaves linear to narrowly oblanceolate, 11-85 
x 2-15 mm, 5.5-14 times longer than broad, 
glabrous, concolorous, resin globules absent; 
margins dentate to denticulate, with 3-6 
pairs of teeth 0.1-07 mm long; apex acute 
or obtuse. Capitula solitary or in pairs, leaf 
opposed, hemispherical, 3-4.5 mm long, 

4.5- 8 mm wide; peduncles 2-28 mm long. 
Involucral bracts incurved, glabrous; median 
bracts ovate, 27-3.2 x 1.3-1.8 mm, apex 
acute or obtuse. Marginal florets 75-250, 
in several rows, female; corolla cylindrical, 

1.5- 2.4 mm long, apex very narrow, white, 


without resin globules. Disc florets 16-68, 
functionally male; corolla cylindrical to 
narrowly campanulate, 2-2.5 mm long, 0.25- 
0.35 mm wide near base, 0.4-0.5 mm wide 
near apex, pink, resin globules absent; lobes 
4 or 5, triangular, each 0.4-0.5 mm long; 
style undivided. Achenes formed from all 
florets, but those from disc florets infertile. 
Marginal achenes fertile, cylindrical, 1-1.3 
mm long, 0.25-0.3 mm wide, brown; ribs 
5-10, white; twin hairs absent or 1-4, resin 
globules absent. Pappus corona almost 
erect, 0.02-0.05 mm high, margin entire or 
fimbriate; pappus bristles absent or with 1-3 
bristles up to 1.9 mm long, and (in inland 
populations) with numerous short bristles. 
Disc achenes infertile, cylindrical, 0.8-17 
mm long, 0.3-0.4 mm wide, white; ribs 
absent; pappus corona erect, white, margin 
entire or fimbriate; pappus bristles absent or 
with 1-5 barbellate bristles arising from the 
corona, up to 1.9 mm long. Fig. 1A-D. 

Additional specimens examined : Queensland. Port 
Curtis District: Broad Sound, Sep 1802, Brown 
s.n. (CANB, MEL); Bay of Inlets, Jun 1770, Banks 
& Solander s.n. (BM); 2 km NW of Seahound Hard 
ramp, Shoalwater Bay Training Area, N of Yeppoon, 
Feb 2012, Bean 31610 & Mathieson (AD, B, BRI, US, 
W, to be distributed); Ramsay Crossing, squatters 


38 


Austrobaileya 9(1): 30-59 (2013) 


Fig. 1. A-D. Sphaeromorphaea major. A. Flowering branchlet x0.5. B. disc floret and infertile achene with pappus 
bristles xl6. C. marginal floret x32. D. achene x32. E-I, S. australis. E. flowering branchlet xl. F. disc floret x32. G. 
marginal floret x32. H. achene, lateral view x32.1. achene, plan view x32. A-D from Bean 30830 & Halford (BRI); E 
from Bean 9548 (BRI); F-I from Kemp 3362H& Kutt (BRI). Del. W.Smith. 


Bean, Sphaeromorphaea and Ethuliopsis 

camp, 40 km NW of Gladstone, near Curtis Island, Dec 
2004, Batianoff 0412502 & Halford (BRI); Targinnie 
district, 18.5 km NW of Gladstone, Nov 2011, Fell DGF 
QEQ79/1 (BRI). Mitchell District: Bowen Downs, 
in 1873, Birch s.n. (MEL); Second Spring, Edgbaston 
Reserve, NE of Aramac, Apr 2012, Bean 31643 (A, BRI, 
DNA, G, HO, MEL, NSW, to be distributed); Measuring 
Spring, Edgbaston Reserve, NE of Aramac, Apr 2012, 
Bean 31646 (BRI, CANB); Edgbaston, Mar 1998, 
Fensham 3464 (BRI). South Kennedy District: Little 
Moses Spring, Doongmabulla Station, 165 km NW of 
Clermont, May 2012, Danielsen 1555 (BRI). 

Distribution and habitat: Sphaeromorphaea 
major is endemic to Queensland. It is 
known from some coastal locations near 
Rockhampton, and three inland locations 
at Edgbaston Reserve, Bowen Downs and 
Doongmabulla station (Map 1). On the 
coast, it is confined to those salt marsh areas 
(inundated by spring tides and dominated 
by grassland of Sporobolus virginicus (L.) 
Kunth) that apparently receive freshwater 
seepage from adjacent eucalypt or melaleuca 
forest. At the inland locations, it grows on 
the margins of artesian springs in open 
salt-encrusted areas dominated by Cyperus 
laevigatus L., but not where there is surface 
water. 

Phenology: Flowers and fruits have been 
recorded from September to April. 

Notes: Banks and Solander were the first to 
collect this species, from ‘Bay of Inlets’ (near 
Stanage Bay) in 1770, and Solander gave it 
the manuscript name Cotula glabrata. Robert 
Brown then collected it in 1802; the label on 
his specimen at BM [Bennett 2131] states that 
the plant was seen “in paludosis salsis Port 
1, Keppel Bay, Shoalwater Bay [&] Broad 
Sound”. ‘Port 1’ and ‘Keppel Bay’ are near the 
southern and northern ends of Curtis Island 
respectively (Vallance et al. 2001), places 
where there are no modern collections. The 
area covered by Brown’s excursions at ‘Broad 
Sound’ (between the present-day towns of 
Ogmore and Carmila) is also without modern 
collections. For this species, there was a time 
lag of 243 years between the first herbarium 
collection and formal description. This lag 
figure is surely the highest on record for an 
Australian plant taxon, and it may well be a 
record not easily broken. Fontaine et al. (2012) 
found that, for species described in 2007, the 


39 

average time lag (or ‘shelf life’) was around 21 
years for all species, and around 32 years for 
plant species. 

Sphaeromorphaea major differs from the 
sympatric S. australis by the achenes 1-1.3 
mm long (0.65-0.9 mm long in S. australis ); 
marginal florets 1.7-2.2 mm long (0.5-0.9 
mm long in S. australis)-, leaves without 
resin globules (always present in S. australis, 
though sometimes hard to see); no resin 
globules on achenes or corolla (always present 
in S. australis ). 

The Edgbaston populations have a coronal 
pappus with a strongly fimbriate margin, 
while the coastal populations have an entire 
corona margin. The Doongmabulla specimen 
is intermediate with regard to this character, 
with a weakly fimbriate coronal pappus. There 
seem to be no other consistent morphological 
differences between the coastal and inland 
populations. 

Sphaeromorphaea major can be 
distinguished by its narrow leaves and often 
very long peduncles. It is the only species in 
which the leaves lack eglandular multicellular 
hairs and resin globules. Furthermore, its disc 
florets are functionally male and the achenes 
produced are infertile, while in the other 
species the “disc achenes” are usually fully 
fertile. 

Conservation status: Sphaeromorphaea 
major is known from about 400 plants at 
Edgbaston Reserve near Aramac, on the 
margins of three springs. The population 
size at Doongmabulla station is unknown. 
Its continued existence at Bowen Downs is 
unknown. The author has observed several 
hundred plants at Shoalwater Bay Training 
Area, and Brown’s recordings of it suggest 
that there is a strong possibility of extending 
the known distribution. However, in view of 
the very specialised habitat, the total area 
covered is unlikely to exceed 40 square km. 
Applying the Red Fist criteria (IUCN 2001), a 
status of ‘Near Threatened’ is recommended. 

Etymology: The specific epithet refers to the 
disc florets, which are larger in this species 
than in any other species of Sphaeromorphaea. 


40 

2. Sphaeromorphaea australis (Less.) 
Kitam., Acta Phytotax. Geobot. 5(4): 276 
(1936), 6(2): 80 (1937); Epaltes australis Less., 
Linnaea 5: 148 (1830), Syn. Gen. Compos. 206 
(1832); Erigerodes australe (Less.) O.Kuntze, 
Revis. Gen. PL 1: 335 (1891). Type: not cited 
(B, t). Neotype (here designated): Australia: 
New South Wales. Myall Creek road, 
Bungawalbin National Park, c. 22 km SSE of 
Casino, 30 December 2011, A.R. Bean 31450 
(neo: BRI [1 sheet + spirit]; iso: CANB, E, K, 
MEL, NSW, RSA, US, W, to be distributed). 

Sphaeromorphaea petiolaris DC., Prodr. 
6: 140 (1838). Type: [Australia: New South 
Wales.] Port Jackson, undated [in 1819], C. 
Gaudichaud (syn: G-DC, image!; P, image!); 
Port Jackson, undated [in 1823], F.W. Sieber 
697 (syn: G-DC, image!). 

Illustrations : Logan River Branch SGAP 
(2005: 130); Stephens & Sharp (2009: 44), 
both as Epaltes australis. 

Prostrate to ascending perennial herb, to 30 
cm high. Stems with sparse multicellular hairs 
when young, glabrescent. Leaves spathulate, 
obovate or oblanceolate, 26-70 x 7-21 mm, 
2.6-5.3 times longer than broad, discolorous, 
resin globules present on both surfaces; 
young leaves with sparse multicellular hairs, 
often absent at maturity; margins dentate 
to denticulate, with 4-8 pairs of teeth up to 
5 mm long, the longer lobes frequently on 
the attenuate basal part of the lamina; apex 
obtuse or acute. Capitula solitary or in pairs, 
leaf opposed, hemispherical to oblate, 2-3 
mm long, 4-5.5 mm wide; peduncles 1-6 
mm long. Involucral bracts incurved; median 
bracts ovate, 1.5-1.9 x 0.8-1.2 mm, glabrous 
or with a few resin globules on outer surface, 
apex acute. Marginal florets 150-300, in 
several rows, female; corolla lageniform, 
0.5-0.9 mm long, pink to purple, resin 
globules few to numerous, apex very narrow. 
Disc florets 13-34, bisexual or functionally 
male; corolla narrowly campanulate, 1-1.3 
mm long, 0.25-0.3 mm wide near base, 0.4- 
0.45 mm wide near apex, pink to maroon, 
resin globules present throughout; lobes 
4, triangular, each c. 0.2 mm long; style 
undivided. Achenes formed from all florets, 
but those from disc florets sometimes infertile, 


Austrobaileya 9(1): 30-59 (2013) 

white to pale yellow. Marginal achenes fertile, 
cylindrical, 0.65-0.9 mm long, 0.2-0.25 mm 
wide, brown, ribs 8-12, white, twin hairs 
absent or 1-4 at base; resin globules few to 
numerous, shining. Pappus corona transverse, 
obscure, translucent, 0.02-0.05 mm wide, 
with numerous radial striations, margin entire 
to erose; pappus bristles absent. Fig. 1E-I. 

Additional selected specimens examined : Taiwan. 
Tamsuy [Tan-shui], 1864, Oldham s.n. (GH); Houlung, 
Miaoli County, Aug 1968, Kao 7364 (TAI). New 
Caledonia. Koutio-Koudta, Dumbea, Sep 1971, McKee 
24355 (NOU). Australia: Western Australia. 1 km 
N of Serpentine, Feb 1985, Keighery 7182 (PERTH); 
Lambkin Reserve, Serpentine, Apr 1996, Keighery 
14346 (PERTH). Queensland. Cook District: Rocky 
Isles, Great Barrier Reef, c. 18 km S of Cooktown, 
Jun 1969, Done s.n. (BRI [AQ7513]). North Kennedy 
District: Alva Beach, Jun 1949, Smith 4400 (BRI). South 
Kennedy district: Western edge of Lake Buchanan, 
Yarrowmere Station, Mar 1998, Kemp 3362H & Kutt 
(BRI). Mitchell District: Myross, east of Aramac, 
May 2000, Fensham 3885 (BRI). Leichhardt District: 
Baralaba - Woorabinda Road, 1.8 km W of Dawson 
Range, Mar 2005, Bean 23551 (BRI); Humboldt, 45 km 
NE of Rolleston, Jan 1996, Bean 9548 (BRI). Port Curtis 
District: Livingstone Shire Vegetation Survey, site 227 
behind Lammermoon Beach, Sep 1977, Batianoff 806 
& McDonald (BRI, CANB). Burnett District: Allies 
Creek SF, Compartment 146, W of Middle Creek Road, 
Apr 2004, Halford Q8271 & Jessup (BRI). Wide Bay 
District: Old Hollett Road, c. 5 km SW of Noosaville, 
Dec 2003, Bean 21577 (BRI). Maranoa District: 
Around barracks, Carellen, c. 90 km WNW of Bollon, 
Mar 2008, Bean 27456 (BRI). Darling Downs District: 
Girraween NP, 22 km S of Stanthorpe, Apr 2001, 
Batianoff 210534 & Collyer (BRI). Moreton District: 
Hay’s Landing, Wivenhoe Dam, N of Fernvale, Jan 2001, 
Bean 17303 (BRI). New South Wales. Tenterfield, May 
1921, Cheel s.n. (NSW); Pilliga, Nov 1932, Rupp 15 
(NSW); Trial Bay to Laggers Point, NE of Kempsey, Jan 
1953, Constable s.n. (NSW, US); c. 4 km by road W then 
N of Crowdy Head on road to Diamond Head, Mar 1981, 
Haegi 2026 (AD, MEL, NSW); 3 miles [5 km] along 
Currambean Creek Road, 5 miles [8 km] S of Nowra, 
Mar 1951, Ford s.n. (NSW); Huskisson, Jervis Bay, Apr 
1916, Rodway 4509 (NSW). 

Distribution and habitat : Sphaeromorphaea 
australis occurs naturally in Australia and in 
New Caledonia. In Australia it extends from 
Ulladulla, New South Wales to Townsville, 
Queensland (with an outlier further north 
near Cooktown), and up to 600 km inland 
e.g. Lake Buchanan, Bollon, Pilliga (Map 2). 
It is naturalised in Taiwan (first collection in 
1864) and near Perth, Western Australia (first 
collection in 1985) (Map 3). In New South 


Bean, Sphaeromorphaea and Ethuliopsis 

Wales and southern Queensland, it occupies 
a range of habitats in forest dominated by 
Eucalyptus spp., Acacia harpophylla F.Muell 
exBenth. ox Melaleuca spp., and sometimes in 
marine couch grassland adjacent to mangrove 
trees. Towards the northern end of its native 
range, the species appears to be increasingly 
confined to saline habitats, such as coastal salt 
marsh, artesian springs or the edges of saline 
inland lakes. Soil ranges from white sands to 
dark heavy clays. 

Phenology : Flowers and fruits can be found 
in any month of the year. 

Typification: Lessing (1830) catalogued all 
Asteraceae specimens held at the Berlin 
Botanical Museum (B), wherein he provided a 
three word description of Epaltes australis as 
a footnote, without mention of the specimen(s) 
upon which the name is based. Soon after, in 
his Synopsis Generum Compositarum , he 
(Lessing 1832) expanded the description of 
E. australis slightly, again without referring 
to the specimen(s) used. He did not explicitly 
give the country of origin, but did state that 
the genus is found in “Asiae vel Novae- 
Hollandiae vel Americae aequinoctialis” The 
other three species treated ( E. divaricata, E. 
mexicana, E. hirsuta) were known to him 
from India, Mexico and India respectively, so 
it follows that E. australis is the Australian 
species. 

While it is unclear from reading A.P. 
de Candolle’s treatment of E. australis 
(de Candolle 1836) whether he viewed the 
specimen that Lessing used to describe it, in 
hindsight it is obvious that he did not. Candolle 
cited a specimen for E. australis , collected by 
Cunningham from the Lachlan River in New 
South Wales and thanks to a paper by O.W. 
Sonder (1853), we know this is not the correct 
application of the name Epaltes australis. 
Sonder examined both Lessing’s specimen 
at B and the Cunningham specimen cited by 
de Candolle, and he indicated unambiguously 
(Sonder 1853: 482) that E. australis sensu de 
Candolle was not that of Lessing, and that 
Ethulia cunninghamii Hook, is synonymous 
with E. australis sensu de Candolle. He 
further stated (p. 485) that the cited specimens 
of Sphaeromorphaea petiolaris DC. belong to 


41 

the same taxon as Lessing’s (type) specimen 
of Epaltes australis. However, Sonder did not 
impart any information about the collector or 
provenance of Lessing’s type specimen. 

Nearly all material at B was destroyed 
during World War II (Hiepko 1987), including 
the type of E. australis , and the absence of 
information about the collector, date of 
collection or provenance of the type (except 
‘Novae-Hollandiae’) means that there is no 
starting point to search for potential duplicate 
specimens. As no known type material exists 
for E. australis , a neotype is here chosen from 
material collected in “Novae-Hollandiae”, 
i.e. Australia, and from the taxon that occurs 
around Sydney, the most likely collection site 
of the original specimen. 

Notes : Sphaeromorphaea australis is 
distinguished from other species in the genus 
by its dark green leaves, the frequent presence 
of conspicuous lobes on the basal attenuate 
part of the lamina; the lageniform marginal 
florets; the narrow and inconspicuous coronal 
pappus; the lack of pappus bristles; and the 
absence (usually) of hairs at the base of the 
achene. Leaves are often glabrous at maturity, 
but specimens from the more westerly areas 
have leaves that often retain some hairs at 
maturity. 

In Queensland, its distribution overlaps 
extensively with Sphaeromorphaea 
subintegra, but both species maintain their 
identities without any sign of intergradation 
or hybridization. S. subintegra may be 
differentiated by its pale green leaves with 
persistent (though often sparse) hairs and 
denticulate margins (teeth <0.5 mm long), 
the conical marginal florets, and the relatively 
broad and conspicuous coronal pappus. 

Conservation status : A common and 
widespread species. 

3. Sphaeromorphaea harrisii (F.Muell.) 
A.R.Bean comb, nov.; Epaltes harrisii 
F.Muell., Fragm. 11: 101 (1880); Erigerodes 
harrisii (F.Muell.) O.Kuntze, Revis. Gen. PL 
1: 335 (1891). Type: Australia: Queensland. 
Cook District: Possession Island, in 1880, 
C.C. Harris s.n. (lecto [here designated]: K 


42 

000373331; isolecto: MEL 2159948, NSW 
582206). 

Illustrations : Britten (1901: t. 158), as 
Epaltes australis ; Dunlop (2000: 185), as 
Sphaeromorphaea australis. 

Erect annual or perennial herb to 35 cm high. 
Stems with dense multicellular hairs when 
young, hairs persistent. Leaves spathulate to 
oblanceolate, 22-61 x 7-21 mm, 2.5-3.8 times 
longer than broad, discolorous or concolorous, 
resin globules present on both surfaces; young 
leaves with dense multicellular hairs, fully 
expanded leaves sparsely to densely hairy; 
margins dentate to lyrate, rarely denticulate, 
with 6-10 pairs of teeth up to 5 mm long; 
apex acute. Capitula solitary or in pairs, 
hemispherical to cupular, 3-3.5 mm long, 
4.5-6 mm wide; peduncles 1-3 mm long. 
Involucral bracts incurved; median bracts 
ovate, 2-2.5 x 0.7-1.1 mm, glabrous, apex 
acute. Marginal florets 200-300, in several 
rows, female; corolla lageniform, 0.9-1.3 mm 
long, maroon to purple, resin globules few or 
none, apex very narrow. Disc florets 15-31, 
bisexual; corolla cylindrical, 1.25-1.4 mm 
long, c. 0.3 mm wide near base, c. 0.35 mm 
wide near apex, pink to maroon, resin globules 
scattered throughout; lobes 4, triangular, 
0.2-0.25 mm long; style undivided; pappus 
bristles sometimes present. Achenes formed 
from all florets, but those from the disc florets 
sometimes infertile, white to pale yellow, not 
ribbed. Marginal achenes fertile, cylindrical, 
0.7-1.1 mm long, 0.2-0.3 mm wide, brown, 
ribs 6-10, white, twin hairs 5-20 at base; resin 
globules numerous. Pappus corona transverse, 
pale and translucent, 0.05-0.08 mm wide, 
margin entire to erose; pappus bristles absent 
or present, slender, without barbellae, 0-6 per 
floret, 0.1-1 mm long. Fig. 2A-E. 

Additional selected specimens examined : Papua 
New Guinea. Western Province: Arufi, Wassi Kussa 
River, Jul 1968, Henty & Katik NGF38641 (A, BRI, 
CANB, K, L); Bula Plains, Morehead subdistrict, Nov 
1972, Henty & Foreman NGF49355 (A, BRI, CANB, 
K, L, LAE); Garmari on the Bula Plains, c. 30 km S of 
Bensbach Lodge, Apr 1997, Mitchell 4696 (BISH, BRI, 
LAE); Balamuk, Bensbach River, c. 9 km S of Bensbach 
Wildlife Lodge, May 1992, Waterhouse BMW2503 
& Obedi (BRI). Australia: Northern Territory. 
Nhulunbuy Lagoon, Oct 1993, Egan 2804 (DNA). 
Queensland. Cook District: Saibai Island, Sep 1994, 


Austrobaileya 9(1): 30-59 (2013) 

Wannan 130 (BRI); [North] Possession Island [Iem Islet], 
Sep 1792, Smith & Wiles s.n. (BM); Possession Island, in 
1880, Harris s.n. (MEL2162771); Thursday Island, June 
1897, Bailey s.n. (BRI [AQ247412]); 1 km S of Seisia, 250 
m from beach, Nov 1999, Wannan 1412 (BRI, CANB, 
NSW); New Holland, in 1770, Banks & Solander s.n. 
(BRI [AQ450752]; MEL); Bolt Head, Temple Bay, Jul 

1991, Forster PIF8985 (BRI, DNA, MEL); Coen River 
[Pennefather River], Gulf of Carpentaria, Nov 1802, 
Brown s.n. (CANB); 8.5 km NW of Weipa Mission, 
Dec 1974, Specht W243 & Salt (BRI); Evans Landing 
rubbish dump, Weipa, Aug 1996, Waterhouse BMW3918 
(BRI); 11 km N of the Chester River Crossing, W of the 
Embley Range, Silver Plains Station, Jun 1992, Forster 
PIF10539 et al. (BRI); 18.6 km from Running Creek 
on the track to Old Port Stewart, Jun 1993, Clarkson 
10103 & Neldner (BRI, DNA, L, MEL); Lakefield NP, 
29.4 km by road from Hann River Crossing, Aug 2011, 
McDonald KRM11779 (BRI); East of mouth of Muck 
River, Cape Melville NP, Jul 1998, Bean 13666 (BRI); 
Lakefield NP, 22.8 km by road N of Ranger base, Aug 
2011, McDonald KRM11776 (BRI); Lizard Island, Nov 
1997, Muir s.n. (BRI [AQ667812]); Lizard Island, Dec 
1974, Specht LI420 & Specht (BRI). North Kennedy 
District: Edmund Kennedy NP, near Cardwell, Jan 

1992, Bean 3912 (BRI, DNA, MEL). 

Distribution and habitat : In Australia 
Sphaeromorphaea harrisii occurs along 
the east coast of Queensland from Cardwell 
to Cape York and on several of the Torres 
Strait islands as well as in the north-eastern 
Northern Territory. It is also found in southern 
Papua New Guinea (Map 4). It grows on a 
wide range of soil types, from sand to black 
clay, on floodplains and riverbanks where 
Melaleuca spp. or Lophostemon suaveolens 
(Sol. ex Gaertn.) Peter G.Wilson & J.T.Waterh. 
are dominant. It also occurs at the base of 
sand dunes, and in Sporobolus virginicus 
grassland adjacent to mangrove communities. 

Phenology : Flowers and fruits can be found 
in any month of the year. 

Typification: The original material of Epaltes 
harrisii (now distributed in K, MEL and 
NSW) comprises numerous very young 
plants that bear only one or two capitula. Most 
capitula are immature; only the material at K 
has florets at anthesis, hence the K sheet has 
been selected as the lectotype. 

There is a second gathering of S. harrisii 
at MEL (MEL2162771) with the same label 
details as the type. This sheet comprises 
three mature plants bearing many flowers and 
mature achenes. Judging by the description 


Bean, Sphaeromorphaea and Ethuliopsis 


43 


Fig. 2. A-E, Sphaeromorphaea harrisii. A. Flowering branchlet *1. B. disc floret and achene with pappus bristle x32. 
C. marginal floret x32. D. achene, lateral view x 32. E. achene, plan view *32. F-J, S. ephemera. F. flowering branchlet 
xl. G. disc floret and achene x32. H. marginal floret x32.1. achene, lateral view x32.1. achene, plan view x32. A, from 
Forster PIF10539 et al. (BRI); B-E from McDonald KRM11779 (BRI); F-J from Forster PIF22217 & Booth (BRI). 
Del. W.Smith. 


44 

in the protologue, Mueller did not use this 
collection when naming the species and hence 
it cannot be considered a type. 

Notes : Sphaeromorphaea harrisii is 
distinctive in the erect stems, the lageniform 
marginal florets with few or no resin globules; 
capitula not as broad as in other species; the 
very short peduncles (often c. 1 mm long); 
the frequent presence of pappus bristles; and 
the achenes with few (6-10) ribs. It is perhaps 
closest to S. australis , but differs from that 
species by the more upright habit, persistently 
hairy leaves, the longer corolla of the marginal 
florets, the broader coronal pappus and the 
frequent presence of pappus bristles. 

Pappus bristles are present on most S. 
harrisii collections, but completely lacking 
in others (including the type). There is no 
geographical basis to this variation. For 
example, Specht LI420 & Specht, from Lizard 
Island, is without pappus bristles; while Muir 
s.n., also from Lizard Island, has numerous 
pappus bristles. 

Conservation status : A common and 
widespread species. 

Etymology : This species was named for 
Charles Christopher Harris (1844-1887), an 
amateur naturalist. He was borne in Maine, 
U.S.A., and immigrated to Australia in the 
1860s. He arrived at Cooktown in 1873 or 
1874, and resided there until his death. He was 
accidently killed while on board the sailing 
vessel ‘Spey’ on 30 th April 1887, when the 
boom struck him on the head (M.Scully pers. 
comm.; National Library of Australia (2009- 
onwards)). 

4. Sphaeromorphaea ephemera A.R.Bean 
sp. nov. affinis S. australi sed habitu annuo, 
flosculis foemineis conicis et acheniis 
brevioribus globulis resinae carentibus 
differens. Typus: Australia: Queensland. 
Gregory North District: Bladensburg 
National Park, S of Winton, Opalton Road, 
19 March 1998, P.I. Forster PIF22217 & R. 
Booth (holo: BRI). 

Prostrate to ascending annual herb, to 15 cm 
high. Stems with sparse multicellular hairs 
when young, glabrescent. Leaves spathulate 


Austrobaileya 9(1): 30-59 (2013) 

to oblanceolate, 14-53 x 5-17 mm, 2.8-4.8 
times longer than broad, discolorous, resin 
globules present on both surfaces; young 
leaves with sparse multicellular hairs, usually 
absent at maturity; margins dentate to 
denticulate, with 3-12 pairs of teeth mostly < 
0.5 mm long, but with some up to 2 mm long; 
apex obtuse. Capitula solitary or in pairs, leaf 
opposed, hemispherical, 1.5-2 mm long, 4-5 
mm wide; peduncles 1-7 mm long. Involucral 
bracts incurved; median bracts ovate, 1.4- 
2.3 x 0.5-0.9 mm, glabrous or with a few 
resin globules on outer surface, apex acute. 
Marginal florets 100-200, in several rows, 
female; corolla conical, 0.6-0.7 mm long, 
pink to purple, broadest at base, few to several 
resin globules present, apex narrow. Disc 
florets 13-24, bisexual; corolla cylindrical to 
narrowly campanulate, 1-1.1 mm long, 0.25- 
0.3 mm wide near base, 0.25-0.35 mm wide 
near apex, pink to maroon, resin globules 
present throughout; lobes 4, each c. 0.2 
mm long; style undivided. Achenes formed 
from all florets, cylindrical, tapered at both 
ends, 0.45-0.7 mm long, 0.15-0.2 mm wide, 
brown, ribs 10-14, white, twin hairs absent; 
resin globules absent, or rarely 1 or 2. Pappus 
corona transverse, obscure, translucent, 
0.01-0.04 mm wide, with numerous radial 
striations, margin entire; pappus bristles 
absent. Fig. 2F-J. 

Additional specimens examined'. Queensland. South 
Kennedy District: Cudmore, Apr 1997, Fensham 3130 
(BRI). Mitchell District: Lake Inveresk, 70 km NNE 
of Muttaburra, Mar 2004, Cumming 22432 & Thompson 
(BRI). Warrego District: Mariala NP, 52.5 km east 
by road from Adavale, Aug 2009, Forster PIF35727 & 
Thomas (BRI); Road reserve, south of Biloola HS, 2 
km off Adavale Road, west of Charleville, May 2010, 
Wang JW0180 (BRI); 60 km NW [actually WSW] of 
Charleville, off the Diamantina Developmental Road, 
Jun 1976, Purdie 373D (BRI). 

Distribution and habitat : Endemic to 
western Queensland, between Adavale and 
Winton (Map 1). Most occurrences are 
along drainage lines under shrubland of 
Acacia aneura F.Muell. ex Benth., often with 
emergent Eucalyptus populnea F.Muell., and 
in red clayey soils. One site is on the margins 
of a salt lake, with fine sandy soil. 

Phenology : Flowers and fruits have been 
recorded from March to August. 


Bean, Sphaeromorphaea and Ethuliopsis 

Affinities : Sphaeromorphaea ephemera is 
apparently related to S. australis, but differs 
by the conical marginal florets, the annual 
habit, and the shorter achenes that lack resin 
globules. S. ephemera differs from S. littoralis 
by the glabrescent leaves, the shorter achenes 
that lack resin globules, the absence of twin 
hairs on the achenes, and the narrower pappus 
corona. 

Conservation status : Data deficient. While the 
species is known from just a few specimens, it 
is potentially common. 

Etymology : From the Greek ephemeros, 
meaning ‘short-lived’. 

5. Sphaeromorphaea littoralis (Retz.) 
A.R.Bean comb, nov.; Artemisia littoralis 
Retz., Observ. Bot. 5, p. 28 (1789); Epaltes 
littoralis (Retz.) Less., Linnaea 6: 151 (1831). 
Type: seashores of eastern India, undated 
[1768-1785], J.G. Koenig s.n. (lecto, here 
designated: C 10007263, image!). 

Epaltes hirsuta Less., Syn. Gen. Compos. 
206 (1832). Type: ‘In India orientali lecta in 
herbario Thunbergiano’, J.P. Rottler s.n., n.v. 

Cotula quinqueloba Blanco, FI. Filip. [F.M. 
Blanco] 626 (1837), nom. illeg. non L.f. (1782). 

Sphaeromorphaea russeliana DC., Prodr. 6: 
140 (early Jan 1838); in Deless., Icon. Sel. 4, t. 
49 (Feb 1838); Sphaeromorphaea russeliana 
DC. var. russeliana. Prodr. 6: 140 (early 
Jan 1838); in Deless., Icon. Sel. 4, t. 49 (Feb 
1838); Centipeda russeliana (DC.) F.Muell. 
as ‘ russellianaFragm. 8: 142 (1874). Type: 
India. The Circars, undated [1781-1790], P. 
Russell s.n. (syn: G-DC, image!; syn: K-W, 
image!). 

Cotula russeliana Wal 1., Ntimer. List[Wallich] 
3240 (1831), nomen nudum 

Illustrations : Gagnepain (1924: 557), as 
Sphaeromorphaea russeliana, Cunningham 
et al. (2011: 685), as Epaltes australis. 

Prostrate to sprawling perennial herb, to 25 
cm high. Stems with dense multicellular hairs 
when young, hairs persisting at maturity. 
Leaves spathulate to oblanceolate or obovate, 
9-60 x 4-15 mm, 2.3-4.8 times longer than 
broad, concolorous, resin globules present on 


45 

both surfaces; young leaves densely hairy, 
hairs sparse to dense at maturity; margins 
dentate to denticulate, with 4-14 pairs of teeth 
usually less than 1 mm long, but occasionally 
up to 2 mm long; apex acute. Capitula solitary 
or in pairs, hemispherical, 2.5-3.5 mm long, 
3.5-7 mm wide; peduncles 2-14 mm long. 
Involucral bracts incurved, glabrous; median 
bracts ovate, 1.6-2.3 x 0.8-1.2 mm, glabrous 
or with few to many resin globules on outer 
surface, apex acute. Marginal florets c. 200, 
in several rows, female; corolla conical, 0.7- 
1.2 mm long, pink to purple, resin globules 
numerous, apex very narrow. Disc florets lb- 
40, bisexual; corolla cylindrical, 1.1-1.4 mm 
long, 0.25-0.4 mm wide, pink, resin globules 
present throughout; lobes 4, triangular, each 
0.2-0.3 mm long; style undivided, arms 
parallel. Achenes formed from all florets, 
0.7-1.3 mm long, 0.25-0.35 mm wide, brown, 
ribs 8-14; twin hairs 10-40, mainly at base 
but with some scattered above; resin globules 
numerous, shining. Pappus corona transverse, 
circular or somewhat pentagonal, 0.05-0.09 
mm wide, with numerous radial striations, 
margin entire, erose or rarely toothed; pappus 
bristles usually absent, rarely present. Fig. 3. 

Additional selected specimens examined : India. 
Hazaribagh, Apr 1884, Clarke 34806 (K); Eastern India, 
s.dat ., Koenig s.n. (LD); Visakhapatnam, Waltan Beach 
near university, Sep 1969, Rao 7101 (CAL). China 
Southeast. Chung Chi college, Hong Kong, Apr 1972, 
Hu 11888 (A, K, US); Nanning, Guangxi province, Nov 
1935, Liang 67192 (PE); Yongning, Guangzi province, 
Jul 1958, ZhongA62026 (PE). Hainan: Yaichow, Hainan 
Island, Sep 1933, Liang 62870 (GH, US); Danzhou 
county, Hainan, May 1928, Tsang 16813 (PE, US). 
Thailand. Nawng Kai, Chaiyaburi, Feb 1924, Kerr 8543 
(BM); Klong Hoy Kong, Songkla province. May 1986, 
Maxwell 86-305 (A); Kao Sabab, Chantaburi, Feb 1935, 
Seidenfaden 2735 (C). Vietnam. Near beach, Tourane, 
May-Jul 1927, Clemens & Clemens 3079 (BM, K, US); 
Phu Yen province, Jan 1936, Petelot 6285 (GH, US); Nha 
Trang, s.dat., Poilane 6046 (K, US). Malaysia. Near 
Kampong Padangnegeri, Kuala Trengganu, Jul 1953, 
Sinclair 39806 & Salleh (BM); Tumpat, Kota Bahru, 
Feb 1917, Ridley 535 (K). Philippines. Manila, Jan 1915, 
Merrill Species Blancoanae No 773 (AMES, BM, CAL, 
GH, NSW, US); Caloocan, province of Rizal, Luzon, 
Nov 1903, Merrill 3655 (US). Marianas, southern end of 
Cabras Island, Guam, Mar 1966, Evans 1673 (US); Waste 
ground just inland from Asan Point, Guam, Jun 1983, 
Raulerson 4561 (US). Australia: Western Australia. 
Piccaninny Creek Gorge, 15 km SE of Bungle Bungle 
Outcamp, Bungle Bungle Range, Apr 1985, Blackwell 
BB297 (PERTH); King Edward River, old CRA 


46 


Austrobaileya 9(1): 30-59 (2013) 


Fig. 3. Sphaeromorphaea littoralis. A. Flowering branchlet xl.5. B. leaf surface with resin globules and eglandular 
multicellular hairs x24. C. flowering capitulum x8. D. disc floret and achene x32. E. marginal floret x32. F. achene, 
lateral view x32. G. achene, plan view x32. A-B from Bean 23777 (BRI); C from Brass 8806 (BRI); D-G from 
Badman 1473 (AD). Del. W. Smith. 


campsite, 1 km S of track to old Mitchell River Station, 
Jun 1988, Edinger 541 (DNA, PERTH); Drysdale River 
above Mogurnda Creek, Drysdale River NP, Aug 1975, 
George 13570 (CANB, PERTH). Northern Territory. 
1.5 km NE of Fish River Gorge in China Wall, Barkly 
Tableland, Jun 1974, Kanis 1823 (CANB, DNA, L, 
US); SE corner of entry to Lake Surprise, Mar 1973, 
Maconochie 1672 (DNA, MO, NSW); Mary Ann Dam, 
c. 5km NE of Tennant Creek, May 2005, Bean 23777 


(BRI); c. 12 miles [c.19 km] NE Finniss River HS, Oct 
1971, Must 842 (CANB, DNA, NSW); Near mouth 
of Foelsche River, Jul 1948, Perry 1829 (AD, BRI, 
CANB, MEL, NSW, US). South Australia. Carruranna 
Waterhole, Cooper Creek, 32 km WNW of Etadunna 
HS, Feb 1984, Badman 1473 (AD, BRI, CANB, MEL, 
NSW); c.lkmN of Big Bend S of Swan Reach, May 
1973, Donner 4113 (AD, BRI); Sunnyside opposite 
Mypolonga on riverbank, Dec 1961, Symon 1926 (AD, 


Bean, Sphaeromorphaea and Ethuliopsis 

BRI). Queensland. Cook District: Cumberland, Gilbert 
River, May 1937, Brass 8806 (BRI). Burke District: 
eastern shore of Lake Moondarra, 16 km NNE of Mt Isa, 
Oct 1997, Fraser 245 (A, CANB). Mitchell District: 
Yalleroi, Mar 1946, Clemens s.n. (BRI [AQ247452]; 
GH). Gregory North District: SE edge of Lake 
Machattie, c. 70 km SE of Bedourie, May 2007, Bean 
26406 (BRI). Warrego District: Currawinya NP, Lake 
Numalla, Mar 1996, Forster PIF18688 (BRI). New 
South Wales. Cobar, Sep 1910, Abrahams 387 (NSW); 
4 miles [6 km] S of Gwabegar, between Baradine and 
Pilliga, Nov 1967, McGillivray 2746 (NSW); Cobham 
Lake, c. 32 km SSE of Milparinka, Mar 1972, Milthorpe 
711 (NSW). Victoria. Dimboola, Feb 1901, D’Alton s.n. 
(NSW); Post No 7 on Hattah Nature Drive, Hattah NP, 
Jan 1989, DAubert 483 (MEL, NSW); On SW side of 
Lake Lockie, 465 m NNE (at 188 degrees) from the E end 
of Lockie Track, Mar 2007, Stajsic 4364 (MEL, NSW). 

Distribution and habitat : Sphaeromorphaea 
littoralis is widely distributed in southern 
Asia, with native occurrences in India, 
Thailand, Vietnam, Malaya, Philippines 
and southeastern China. It is naturalised on 
the island of Guam (Map 6). In Australia, 
it is indigenous in the Kimberley region of 
Western Australia, South Australia, Northern 
Territory, western Queensland, western New 
South Wales and north-western Victoria 
(Map 5). The species inhabits sunny places 
on the margins of creeks and dams, saline 
coastal flats and swamps, and in Australia 
extends into areas receiving very low annual 
rainfall. Soils are variable. 

Phenology : Flowers and fruits may be found 
at any time of the year. 

Typification: I am aware of just two specimens 
of Artemisia littoralis that were collected by 
J.G. Koenig - one at C and one at LD. The 
LD specimen is notable for the presence of 1 
or 2 pappus bristles, up to 0.6 mm long, on 
many of its achenes. The specimen at C lacks 
these bristles, and it is therefore deduced 
that these two specimens were from separate 
gatherings. The C specimen is superior in 
quality, and pappus bristles are not mentioned 
in the protologue; hence it is chosen as the 
lectotype. 

Type material of Epaltes hirsuta was sought 
from all herbaria that are listed as having 
collections of C.P. Thunberg or J.P Rottler 
(HUH 2012), but without success. However, 
I consider it a synonym of S. littoralis ; the 
words “capitulis solitariis vel geminis” given 


47 

in the protologue strongly suggests that E. 
hirsuta is a Sphaeromorphaea , and the “tota 
hirsuta” accords with the hairy Indian form of 
S. littoralis. 

Notes: Sphaeromorphaea littoralis differs 
from S. australis by the grey-green hairy 
leaves (green glabrous leaves in S. australis)', 
the conical-shaped marginal florets that have 
numerous resin globules (lageniform florets 
with few resin globules for S. australis), the 
prominent corona on the achenes; and the 
numerous hairs at the base of the achene. 

Sphaeromorphaea littoralis differs from 
S. subintegra by the longer teeth on the leaf 
margins, presence of teeth on the basal half of 
the leaves, fully expanded leaves moderately 
to densely hairy, mostly longer peduncles (to 
14 mm, versus 1.5-3.5 for S. subintegra), and 
the presence of numerous antrorse hairs at the 
base of the achene. 

Australian specimens of S. littoralis tend 
to have larger leaves than is usual for Asian 
specimens. Indian and northern/central 
Australian specimens are generally densely 
hairy, while specimens from south-east Asia 
(e.g. Thailand, China) and around the Murray 
River in southern Australia are generally only 
sparsely hairy. According to Cunningham et 
al. (2011), it is completely ignored by domestic 
livestock. 

The collector of the type of S. russeliana 
was Patrick Russell, usually spelt with 
two “l’s” (Hawgood 1994). However, early 
references (Wallich 1831; Wight 1831; De 
Candolle 1838) cited his surname as ‘Russel’, 
while Lowndes (1834) cited his surname 
as ‘Russel’ or ‘Russell’. The spelling of the 
epithet ‘russeliana’ by de Candolle (1838) was 
clearly intentional and that spelling must be 
maintained (Art. 60). 

Wallich (1831) included a Russell 
manuscript name (Artemisia chrysanthemum) 
as a synonym of his Cotula russeliana [Cat. 
3240], The latter, also a nomen nudum, 
was later validated as Sphaeromorphaea 
russelliana DC. The epithet chrysanthemum 
means ‘golden-yellow flowers’, and so some 
later accounts (Hooker 1882; Li 1978; Naithani 
1995) and some specimen labels have stated 


48 

that Sphaeromorphaea has yellow flowers. 
All taxa and populations of Sphaeromorphaea 
known to the present author have pink, purple, 
maroon or (rarely) white florets, and the 
occurrence of yellow flowered populations is 
considered highly unlikely. It is probable that 
Russell’s manuscript name referred to another 
species e.g. Grange a maderaspatana (L.) 
Poir., and Wallich merely misapplied it. 

Sphaeromorphaea littoralis is naturalised 
at Guam, being first recorded in 1966 (Fosberg 
& Sachet 1980), adjacent to the Naval Base. 

Conservation status : A common and 
widespread species. 

6. Sphaeromorphaea subintegra A.R.Bean 
sp. nov. affinis S. australi sed infoliis dentibus 
aequabiliter parvis, pappo coronali comparate 
lato et conspicuo, pilis persistentibus in 
foliis maturis et corolla foeminea conica (in 
S. australi lageniformi) differens. Typus: 
Australia: Queensland. South Kennedy 
District: Lambert Beach, Slade Point, 
Mackay, 16 July 1992, G.N. Batianoff9207103 
& H.A. Dillewaard (holo: BRI; iso: MEL). 

Prostrate to ascending perennial herb, to 
25 cm high. Stems with sparse to dense 
multicellular hairs when young, hairs ± 
persistent. Leaves spathulate to obovate, 
14-50 x 4-19 mm, 2.4-37 times longer than 
broad, discolorous, resin globules present on 
both surfaces; young leaves with moderately 
dense multicellular hairs, some hairs 
persisting at maturity; margins denticulate, 
with 3-12 pairs of small teeth <0.5 mm long; 
apex obtuse. Capitula solitary or in pairs, 
leaf-opposed, hemispherical, 2-3 mm long, 
5-6 mm wide; peduncles 1.5-3.5 mm long. 
Involucral bracts incurved; median bracts 
1.5-2.5 x 0.8-0.9 mm, glabrous or with a few 
resin globules on outer surface, apex acute. 
Marginal florets 150-300, in several rows, 
female; corolla conical, 0.6-0.9(-l) mm long, 
mauve to purple, broadest at base, with few to 
numerous resin globules; apex narrow. Disc 
florets 22-44, bisexual; corolla cylindrical 
to narrowly campanulate, 0.9-1.3 mm long, 
0.25-0.35 mm wide near base, 0.25-0.4 mm 
wide near apex, pink to maroon, resin globules 
numerous; lobes 4, triangular, each 0.15-0.25 


Austrobaileya 9(1): 30-59 (2013) 

mm long; style undivided. Achenes formed 
from all florets, cylindrical, 0.7-0.9 mm long, 
0.25-0.3 mm wide, brown, ribs 10-14, twin 
hairs absent or 1-4 at base; resin globules 
numerous, shining. Pappus corona transverse, 
obvious, white to grey, 0.05-0.08 mm wide, 
margin entire to erose; pappus bristles absent 
or very rarely present. Fig. 4A-E. 

Additional selected specimens examined : Papua New 
Guinea. Milne Bay Province: Narian, Misima Island, 
Aug 1956, Brass 27606 (A); Joe’s Landing, Sudest Island, 
Aug 1956, Brass 27780 (A, US). Western Province: 
Daru, near airstrip, Jun 1973, Henty NGF49517 (A, 
BRI); Daru Island, Sep 1963, Womersley NGF17786 
(BRI). New Caledonia. Presqu’ile de Ouano, Sep 1971, 
Raynal 16504 (NOU). Australia: Queensland. Cook 
District: Booby Island, Great Barrier Reef, c. 34 km 
from Thursday Island, Jul 1969, Heatwole s.n. (BRI 
[AQ8142]); East of the Mossman to Daintree Road north 
of Wonga Beach, Sep 2002, Jago 6297 (BRI); Mareeba, 
Jan 1918, White s.n. (BRI [AQ247418]). North Kennedy 
District: E of Mount Garnet, Feb 2008, Wannan 4999 
(BRI, NSW); Little Ramsay Bay, Hinchinbrook Island, 
Apr 2012, Mathieson MTM1327 (BRI, NSW); Snake 
Creek, c. 95 km NW of Charters Towers on Gregory 
Developmental Road, Apr 1991, Batianoff SC9104010 
(BRI). South Kennedy District: Brampton Island near 
airstrip, Sep 1986, Batianoff 5123 (AD, BRI); Black 
Wattle Creek Crossing, on Gregory Development Road, 
May 1991, McFadyen 453 (BRI); Collinsville Coal Pty 
Ltd mine lease area, near Collinsville and Scottville, Apr 
1984, Thompson 587 (BRI). Leichhardt District: 8.8 km 
from Mt Dillingen peak and c. 87 km NNE of Clermont, 
Feb 1999, Bailey LB65 (BRI); 45.8 km from Taroom, on 
road to Bauhinia Downs, May 2010, Bean 29617 (BRI). 
Port Curtis District: 200m down road to St Lawrence 
from Bruce Highway, Jul 1998, Thompson 1317 (BRI); 
Facing Island, Oct 2004, c. 0.7 km NW of Catcombe, 
Halford Q8670 (BRI). Burnett District: About 3.3 km 
SE of Culcraigie along Culcraigie - Eidsvold Road, Apr 
1997, Pollock ABP391 (BRI); Toondahra HS yards, Jun 
1989, Forster PIF5076 (BRI, US). Wide Bay District: 
Near Maroondan, E of Gin Gin, Nov 1996, Bean 11229 
(BRI). 

Distribution and habitat : Sphaeromorphaea 
subintegra is recorded from southern lowland 
parts of Papua New Guinea, New Caledonia, 
and in Australia is widespread in eastern 
Queensland from the Torres Strait islands 
to Mundubbera (Map 7). It grows on wet 
or moist microhabitats in open forest and 
woodland dominated by Eucalyptus or 
Melaleuca , often on alluvium, but also on 
hillsides and plateaux. Soils vary from sand 
to reddish basaltic clay, and including soils 
derived from serpentinite. It has not been 
recorded from salt marsh communities. 


Bean, Sphaeromorphaea and Ethuliopsis 


49 


Fig. 4. A-E, Sphaeromorphaea subintegra A. Flowering plant *0.6. B. disc floret and achene *32. C. marginal floret 
x32. D. achene, lateral view x32. E. achene, plan view x32. F-J, Ethuliopsis cunninghamii. F. flowering branchlet xl. 
G. disc floret xl6. H. marginal floret x32. L achene, abaxial view x32. J. achene, adaxial view x32. A, from Bean 29617 
(BRI); B-E from McFadyen 453 (BRI); F, from Cowan 81 (BRI); G, from Mitchell 824 (BRI); H-J from Emerson s.n. 
(BRI [AQ501325]). Del. W.Smith. 


50 

Phenology : Flowers and fruits can be found 
in any month of the year. 

Notes : Sphaeromorphaea subintegra differs 
from S. australis by the consistently small 
teeth on the leaves (no more than 0.5 mm long), 
the relatively broad and conspicuous coronal 
pappus, persistent hairs on mature leaves, and 
the marginal corolla conical in shape. Field 
observations and herbarium specimen labels 
suggest that where the distributions of this 
species and S. australis overlap, S. subintegra 
inhabits the more mesic sites (where fresh 
water is more readily available), whereas S. 
australis inhabits salt-affected sites, including 
salt marsh or headlands near the ocean, or 
sites that may dry out more quickly. No 
intergradation between these two species has 
been observed in the field or in the herbarium, 
even though they sometimes occur less than a 
kilometre from each other. 

S. subintegra differs from S. littoralis by 
the lobing of the leaves (more conspicuously 
lobed in S. littoralis ) and the very few twin 
hairs on the achene, if any (numerous twin 
hairs in S. littoralis). 

The oldest herbarium specimen of S. 
subintegra that I have seen is one collected by 
Ernst Betche at Cooktown in 1881. 

Conservation status : A common and 
widespread species. 

Ethuliopsis F.Muell., Fragm. 2: 154 (1861); 
Epaltes sect. Ethuliopsis (F.Muell.) F.Muell., 
Fragm. 10: 100 (1877). Type: E. dioica 
F.Muell. (= E. cunninghamii). 

Gynaphanes Steetz in Peters, Naturw. Reise 
Mossambique [Peters] 6(2): 457 (1864). 
Type: Gynaphanes australis (= Ethuliopsis 
cunninghamii). 

Perennial herbs. Latex absent. Stems terete, 
not winged. Leaves alternate, sessile, 
toothed, spreading, not decurrent. Capitula 
pedunculate, in terminal panicles; on some 
plants disciform and heterogamous; on 
other plants homogamous, with disc florets 
only. Involucral bracts variable in length, 
cartilaginous, in 3-6 rows. Receptacle 
glabrous, without paleae. Marginal florets 


Austrobaileya 9(1): 30-59 (2013) 

fertile, female, without rays; corolla lobes 
3, tiny; style base not bulbous, branches 
divergent. Disc florets bisexual, functionally 
male, ovary vestigial, not producing achenes. 
Anthers tailed, apically obtuse. Style base not 
bulbous, style undivided at apex; sweeping 
hairs obtuse, extending well down the shaft. 
Achenes lunate, with a single longitudinal rib 
on adaxial side; carpopodium annular, white, 
prominent. Pappus on the marginal florets 
comprising a persistent erect cylindrical 
corona; on the disc florets comprising 3-6 
barbellate bristles. 

A monotypic genus, endemic to Australia. 

Ethuliopsis cunninghamii (Hook.) F.Muell., 
Key to the System of Victorian Plants 1: 
313 (1888); Ethulia cunninghamii Hook, in 
T.Mitch., J. Exped. Trop. Australia 62 (1848); 
Ethuliopsis dioica F.Muell, Fragm. 2: 155 
(1861), nom. illeg .; Gynaphanes australis 
Steetz in Peters, Naturw. Reise Mossambique 
[Peters] 6(2): 458 (1864), nom. illeg .; Epaltes 
cunninghamii (Hook.) Benth., FI. Austral. 3: 
530 (1867); Erigerodes cunninghamii (Hook.) 
O.Kuntze, Revis. Gen. PI. 1: 335 (1891). Type 
citation: “We saw... Also an ETHULIA, which 
may, on further examination, constitute a new 
genus; it was found by Allan Cunningham on 
the Lachlan” Type: Australia: New South 
Wales. Swampy banks of the Lachlan River, 
26 June 1817, A. Cunningham 290/1817 (lecto 
[here designated]: K 000373334; isolecto: 
G-DC). 

Illustrations : Cunningham et al. (2011: 685), 
as Epaltes cunninghamii. 

Erect herb, to 100 cm high. Stems glabrous. 
Leaves oblong to oblanceolate, 20-69 x 

4.5- 17 mm, 3.6-8.5 times longer than broad, 
bright green, concolorous, resin globules 
sometimes present on lower surface; glabrous 
or with sparse multicellular hairs; margins 
denticulate to dentate with 5-13 pairs of 
teeth each 0.3-3 mm long; apex obtuse or 
acute; base cuneate to obtuse. Capitula in 
small terminal panicles. Peduncles 0-4.5 mm 
long, glabrous or sparsely glandular-hairy. 
Homogamous capitula. Capitula cylindrical 
to campanulate at anthesis, 3.5-5 mm long, 

2.5- 5.5 mm wide. Involucral bracts flat, 


Bean, Sphaeromorphaea and Ethuliopsis 

glabrous, outer ones much shorter than inner; 
median involucral bracts 3.1-3.5 x 0.6-1.1 
mm, apex obtuse, margin entire. Receptacle 
flat. Marginal florets absent; disc florets 20- 
35, bisexual; corolla narrowly campanulate, 
2-3.1 mm long, 0.2-0.3 mm wide near 
base, 0.5-1 mm wide near apex, white, 
resin globules present throughout; lobes 5, 
0.25-0.3 mm long; style undivided; ovary 
vestigial; pappus bristles 2-6, 1.9-2.6 mm 
long, barbellate, slightly flattened, often bent 
or twisted near apex. Heterogamous capitula: 
Capitula globose at anthesis, 2.5-3 mm long, 
2.5-3 mm wide. Involucral bracts incurved, 
glabrous; outer ones much shorter than 
inner; median bracts 2-2.7 x 1-1.2 mm, apex 
obtuse, glabrous, margin entire or denticulate. 
Receptacle hemispherical. Marginal florets 
100-200, in several rows, female; corolla 
filiform, 0.8-1.3 mm long, white, cylindrical, 
multicellular hairs absent, few to numerous 
resin globules present; lobes 3, tiny; styles 
slender, branched. Disc florets 2-16, bisexual; 
corolla narrowly campanulate, 1.2-1.5 mm 
long, 0.2-0.3 mm wide near base, c. 0.5 mm 
wide near apex, white, resin globules present 
throughout; lobes 5, 0.25-0.3 mm long; style 
undivided; ovary vestigial; pappus bristles 
2-6, 0.8-1.1 mm long, barbellate, slightly 
flattened, often bent or twisted near apex. 
Achenes formed only from marginal florets, 
lunate, 0.6-0.7 mm long, c. 0.2 mm wide, 
with 1 conspicuous longitudinal rib, and 
several raised anastomosing veins sometimes 
visible; twin hairs absent; resin globules 
absent. Pappus corona obliquely placed, 
erect, cylindrical, white, 0.03-0.05 mm high, 
margin entire; pappus bristles absent. Fig. 
4F-J. 

Additional selected specimens examined: Australia: 
South Australia. Katarapko Reserve, N of Loxton, Jul 
1985, Alcock 10249 (AD, MEL, NSW, NY); 7 km W of 
Mundondna HS, May 1984, Badman 1022 (AD, CANB, 
MEL, NSW); Waukatanna waterhole. Cooper Creek, 
Apr 1985, Badman 1646 (AD, BRI, MEL); 50 km SE of 
Coongie, Lake Eyre basin, Oct 1986, Conrick 1939 (AD, 
BRI, NSW); Pandiburra Bore, Aug 1975, Donner 5177 
(AD, US); Neals River, 56 km S of Oodnadatta on track 
to William Creek, Nov 1989, Nordenstam & Anderberg 
972 (AD, MEL, NSW, PERTH); Engenina Creek, on 
the Coober Pedy - William Creek Road, Sep 1990, 
Wilson 789 & Rowe (AD, MEL, NSW). Queensland. 
Gregory North District: Cuttaburra Channels, Eyre 
Development Road, S of Bedourie, May 2007, Bean 


51 

26385 (BRI, MEL); 116 km N of Birdsville, Sep 1989, 
Cowan 81 & Bushel1 (BRI). Warrego District: c. 15 
km W of Quilpie, Nov 1975, Henderson H2384 (BRI). 
Gregory South District: Thylungra, c. 75 miles [c. 121 
km] NW of Quilpie, Oct 1955, Everist 5768 (BRI); 1.3 
km S of Plevna Downs HS gate, W of Eromanga, Aug 
2010, Bean 30103 (BRI); Lake Yamma Yamma, Sep 
1991, Mitchell 824 (BRI); Windorah, ‘Tanbar’, Cooper 
Creek, Nov 1990, Emerson s.n. (BRI [AQ 501325]). New 
South Wales. Yantara Lake, Oct 1887, Baeuerlen s.n. 
(NSW); 15.1 km SW of Goodooga, Dec 1998, Bean 14508 
(BRI, NSW); Near Barrier Range, Feb 1861, Beckler s.n. 
(MEL); Delalah Downs, Aug 1972, Cunningham 476 
(BRI, NSW); Willandra Creek, c. 28 miles [c. 43 km] SE 
of Ivanhoe, Nov 1973, Salasoo 5427 (NSW). Victoria. 
Kings Billabong Wildlife Reserve, near Red Cliffs, Aug 
1975, Beauglehole 50256 (AD, MEL); Kings Billabong, 
Red Cliffs area. May 1969, Henshall s.n. (MEL); W 
of Neds Corner HS, Mullaroo Creek, 5.5 km due N of 
Lake Wallawalla, Lindsay Island, Dec 1981, Browne 73 
(MEL); Lake Lalbert, Dec 1853, Mueller s.n. (MEL). 

Distribution and habitat : Ethuliopsis 
cunninghamii is widely distributed in 
south-western Queensland, western New 
South Wales, eastern South Australia and 
north-western Victoria (Map 8), in areas 
that receive less than about 600 mm annual 
rainfall. It grows in heavy clay soil on alluvial 
flats, and around ephemeral swamps and 
lakes. Associated species include lignum 
{Muehlenbeckia flondenta Meisn.), gidgee 
(Acacia cambagei R.T.Baker), coolabah 
{Eucalyptus coolabah Blakely & Jacobs), and 
bluebush ( Chenopodium auricomum Lindl.). 

Phenology : Flowers and fruits have 
been recorded from March to December, 
but flowering and fruiting is most likely 
synchronised to rainfall events rather than 
season. 

Notes : Ethuliopsis is distinct from both 
Epaltes sens. str. (E. divaricata) and from 
Sphaeromorphaea , and would appear to 
be more closely related to some members 
of the large genus Pluchea. The latter is 
a very complex and possibly paraphyletic 
assemblage (Anderberg et al. 2005), and none 
of the Pluchea species in Australia or south¬ 
east Asia have characters similar to those 
exhibited by Ethuliopsis. 

Ethuliopsis cunninghamii is sub- 
dioecious. Some plants have homogamous 
capitula that bear only disc florets. These 
florets are bisexual, but do not produce 


52 

achenes and the ovary is vestigial, hence 
they are effectively male. The capitula on 
these plants are campanulate or cylindrical, 
the median involucral bracts are 3.1-3.5 mm 
long, and the corolla of the disc florets is 2-3.1 
mm long. Other plants have heterogamous 
capitula that bear predominantly marginal 
florets, with a few disc florets in the centre. 
The capitula on those plants are globose, the 
median involucral bracts are 2-2.7 mm long, 
and the corolla of the disc florets is 1.2-1.5 
mm long. Hence, specimens of the “male” and 
“female” plants can be readily distinguished 
by capitulum shape and size, or by floret 
size. The corolla of both marginal and disc 
florets is white, unlike Sphaeromorphaea 


Austrobaileya 9(1): 30-59 (2013) 

spp. (except marginal florets of S. major) and 
Epaltes divaricata which are pink to purple. 

There is one herbarium specimen 
reportedly collected from the Northern 
Territory. It is at MEL, and has a handwritten 
label written in ink stating “118. Epaltes 
cunninghamii B., 1885/6, Lieut. Dittrich, 
Lindsay’s Exped.” Someone else has later 
written, in pencil, “finke river”. As this species 
has never otherwise been recorded from the 
Northern Territory, it seems likely that the 
locality is in error, and that the specimen was 
in fact collected in South Australia. 

Conservation status : A common and 
widespread species. 


Key to the Australian genera of Subtribe Plucheinae 


1 Capitula grouped into secondary heads or glomerules.2 

1. Capitula all separate.3 

2 Pappus absent.Sphaeranthus 

2. Pappus present.Pterocaulon 

3 Pappus of the marginal florets consisting of a small corona, sometimes 

with 1-5 bristles arising from it.4 

3. Pappus of all florets with 10 or more bristles, corona absent.5 

4 Achenes cylindrical, with 5-14 longitudinal ribs; capitula all 

heterogamous, hemispherical to oblate; corolla pink 

to purple, rarely white.Sphaeromorphaea 

4. Achenes lunate, with 1 longitudinal rib; capitula heterogamous or 

homogamous, campanulate or globose; corolla white.Ethuliopsis 

5 Pappus bristles fused into a tube at their base.Coleocoma 

5. Pappus bristles all free.6 

6 Pappus bristles broad-based, persistent, ebarbellate.Thespidium 

6. Pappus bristles slender throughout, somewhat caducous, barbellate.7 

7 Disc florets 3-merous; anthers not tailed.Allopterigeron 

7. Disc florets 4-merous or 5-merous; anthers tailed.8 

8 Corolla of marginal florets irregularly lobed to distinctly ligulate; achenes 

densely sericeous.Streptoglossa 

8. Corolla lobes of marginal florets equal; achenes glabrous or sparsely hairy.Pluchea 


53 


Bean, Sphaeromorphaea and Ethuliopsis 

Excluded names in Epaltes and 
Sph aeromorph aea 

Epaltes alata (Sond.) Steetz in Peters, 
Naturw. Reise Mossambique [Peters] 6(Bot., 
2): 452 (1864) = Litogyne gariepina (DC.) 
Anderb. 

Epaltes gariepina (DC.) Steetz in Peters, 
Naturw. Reise Mossambique [Peters] 6(Bot., 
2): 451 (1864) = Litogyne gariepina (DC.) 
Anderb. 

Epaltespleiochaeta F.Muell., Fragm. 10: 100 
(1877) = Pluchea baccharoides (F.Muell.) 
Benth. 

Epaltes tatei F.Muell., Trans. & Proc. Roy. 
Soc. South Australia 6: 31 (1883) = Haegiela 
tatei (F.Muell.) P.S.Short & Paul G.Wilson. 

Sphaeromorphaea centipeda DC., Prodr. 6: 
140 (1838). An illegitimate name based on 
Artemisia minima L. (= Centipeda minima). 

Centipeda sect. Sphaeromorphaea 
C.B.Clarke, Compos. Ind. 151 (1876) is 
typified by Centipeda orbicularis nom. illeg. 
(= Centipeda minima). This sectional name 
is therefore a synonym of Centipeda. 

Sphaeromorphaea russeliana var. glabrata 
DC., Prodr. 6: 140 (1838). Type: NW India, 
J.F. Royle s.n. (syn: K 000373336, image!) = 
Centipeda minima subsp. minima. A Royle 
specimen at G-DC [G00318749] identified 
as var. glabrata is evidently from a different 
gathering. It bears the number 260, and the 
locality Cachemire [?], rather than the ‘prov. 
bor.-occid.’ as stated in the protologue. This 
latter specimen is not considered a type. It 
represents Sphaeromorphaea littoralis. 

Acknowledgements 

I thank the Directors of A, AD, AMES, BM, 
C, CANB, GH, K, LD, MEL, NOU, PE, 
PERTH and TAI for the loan of specimens; the 
Directors of CAL and M for sending images 
of specimens, V. Fonjallaz (G) for sending 
images of type specimens, J. Hunnex (BM) 
for sending images of historical specimens; 
and lb Friis and Hans Hansen (C) for 


assistance with the type of Artemisia littoralis. 
I am grateful to Philip Sharpe for translating 
some German text, and Jian Wang (BRI) 
for interpreting Chinese specimen labels; to 
Keith McDonald and Mike Mathieson for 
collecting Sphaeromorphaea material from 
Queensland. I also thank Melinda Peters 
(HUH) and Nicola Biggs (K) for friendly and 
efficient assistance; Murray Haseler and Steve 
Heggie for enabling access to Edgbaston 
Reserve; Marge Scully of the Cooktown 
Historical Society for providing information 
about C.C. Harris; and Dr Barbara Hawgood 
for kindly sending a copy of her article on 
Patrick Russell. Peter Bostock (BRI) kindly 
provided the Latin diagnoses, and Will Smith 
(BRI) provided the illustrations and put the 
finishing touches on the distribution maps. 

References 

Anderberg, A. A. (1991). Taxonomy and phylogeny of the 
tribe Plucheeae (Asteraceae). Plant Systematics 
and Evolution 176: 145-177. 

Anderberg, A.A., Eldenas, P., Bayer, R.J. & Englund, 
M. (2005). Evolutionary relationships in the 
Asteraceae tribe Inuleae (incl. Plucheeae) 
evidenced by DNA sequences of ndhF; with 
notes on the systematic positions of some 
aberrant genera. Organisms, Diversity & 
Evolution 5: 135-146. 

Anderberg, A.A. & Eldenas, P. (2007). Tribe Inuleae. 
In: K. Kubitzki (ed.). The Families and Genera 
of Vascular Plants , Volume VIII, Asterales. 
Springer: Berlin, Heidelberg, New York. 

Apc (2012). Australian Plant Census. IBIS database. 
Centre for Australian National Biodiversity 
Research, Council of Heads of Australasian 
Herbaria. http: //ww w. chah. gov. au/apc/index. 

html Accessed 16 December 2012. 

Bailey, EM. (1900). The Queensland Flora, Part III. J. 
Diddams & Co.: Brisbane. 

Beentje, H.J. (2002). Flora of Tropical East Africa , 
Compositae Part 2. Royal Botanic Gardens: 
Kew. 

Bentham, G. (1867). Compositae. In Flora Australiensis 
3: 447-680. L.Reeve & Co.: London. 

- (1873). Compositae. In G. Bentham & J.D. 

Hooker, Genera Plantarum 2(1): 163-537. 
L.Reeve & Co.: London. 

Bremer, K. (1994). Asteraceae: Cladistics and 
Classification. Timber Press: Portland, Oregon. 


54 

Britten, J. (1901). Illustrations of Australian plants 
collected in 1770 during Captain Cook’s voyage 
round the world in H.M.S. Endeavour /by 
the Right Hon. Sir Joseph Banks and Daniel 
Solander, with determinations by James 
Britten. Australian Plants, Part II. Trustees of 
the British Museum: London. 

Brummitt, R.K. (2001). World Geographical Scheme 
for recording Plant Distributions, Edition 2. 
International Working Group on Taxonomic 
Databases for Plant Sciences: Pittsburgh, http:// 
www.tdwg.org/standards/109 Accessed 29 
December 2012. 

Cassini, H. (1818). Aper^u des genres nouveaux formes 
par M. Henri Cassini dans la famille des 
Synantherees. Bulletin des Sciences, par la 
societe Philomatique de Paris 1818: 139-142. 

Cooke, D.A. (1986). Compositae (Asteraceae). In J.P 
Jessop & HR. Toelken (eds.). Flora of South 
Australia, Pt. III. Polemoniaceae-Compositae, 
pp. 1423-1658. South Australian Goverment. 
Printing Division: Adelaide. 

Cunningham, G.M., Mulham, W.E., Milthorpe, PL. 
& Leigh, J.H. (2011). Plants of Western New 
South Wales, 2 nd edition. CSIRO Publishing: 
Collingwood, Victoria. 

De Candolle, A.P (1836). Prodromus systematis 
naturalis regni vegetabilis 5:461-462. Treuttel 
& Wurtz: Paris 

- (1838). Prodromus systematis naturalis regni 

vegetabilis 6: 140. Treuttel & Wurtz: Paris 

Dunlop, C.R. (2000). Asteraceae. In I.D. Cowie, 
P.S. Short & M. Osterkamp Madsen (eds.). 
Floodplain Flora - A flora of the coastal 
floodplains of the Northern Territory, Australia, 
pp. 180-189. Flora of Australian Supplementary 
series No. 10. Australian Biological Resources 
Study: Canberra. 

Figuerola, J. & Green, A.J. (2002). Dispersal of 
aquatic organisms by waterbirds: a review of 
past research and priorities for future studies. 
Freshwater Biology 47: 483-494. 

Fontaine, B., Perrard, A. & Bouchet, P. (2012). 21 years 
of shelf life between discovery and description 
of new species. Current Biology 22: 943-944. 

Fosberg, F.R. & Sachet, M.-H. (1980). Flora of 
Micronesia, 4: Caprifoliaceae to Compositae. 
Smithsonian Contributions to Botany, Number 
46. Smithsonian Institution Press: Washington. 

Gagnepain, F. (1924). In H. Lecomte (ed), Flore generale 
de Undo Chine 3(5): 557-664. Masson & Co.: 
Paris. 


Austrobaileya 9(1): 30-59 (2013) 

Grin (2012). USDA, ARS, National Genetic Resources 
Program. Germplasm Resources Information 
Network - (GRIN) [Online Database], Accessed 
29 January 2012. 

Hawgood, B. J. (1994). The life and viper of Dr Patrick 
Russell MD FRS (1727-1805): Physician and 
Naturalist. Toxicon 32: 1295-1304. 

Heywood, V.H. & Humphries, C.J. (1977). Anthemideae 
- systematic review. In V.H. Heywood et 
al. (eds.). The Biology and Chemistry of the 
Compositae 2: 851-898. Academic Press: 
London. 

Hiepko, P. (1987). The collections of the Botanical 
Museum Berlin-Dahlem (B) and their history. 
Englera 7: 219-552. 

Hooker, J.D. (1882). Compositae. In The Flora of British 
India 3: 219-419. L. Reeve & Co.: London. 

Hooker, W.J. (1831). In R. Wight, Illustrations of Indian 
Botany, principally of the southern parts of the 
peninsula. Botanical Miscellany, London 2: 
90-97. 

Hu, S. & Wong, Y. (2009). Asteraceae. In Flora of Hong 
Kong 3: 245-328. Government of the Hong 
Kong Special Administrative Region: Hong 
Kong. 

Huh (2012) Index of Botanists. Harvard University 
Herbaria. http://kiki. huh. harvard.edu/databases/ 
botanist_index.html Accessed 10 October 2012. 

Iucn (2001). Red List Categories and Criteria: Version 
3.1. IUCN Species Survival commission. IUCN, 
Gland, Switzerland and Cambridge, U.K. 

Leins, P. (1971). Pollensystematische Studien an Inuleen 
I. Botanische Jahrbiicher fur Systematik, 
Pflanzengeschichte und Pflanzengeographie 
91: 91-146. 

Lessing, C.F. (1830). De Synanthereis Herbarii Regii 
Berolinensis, dissertatio secunda: Nassauvieae. 
Linnaea 5: 1-236. 

- (1832). Synopsis Generum Compositarum 

earumque dispositionis novae tentamen 
monographiis multarum capensium interjectis. 
Dunckeri & Humblotii: Berlin. 

Li, H. (1978). Compositae. In Flora of Taiwan 4: 768- 
965. Epoch Publishing Co., Ltd: Taipei. 

Logan River Branch Sgap (2005). Mangrove to 
Mountains, a field guide to the native plants of 
south-east Queensland, Volume 2. Logan River 
Branch SGAP: Browns Plains. 

Lowndes, W.T. (1834). A bibliographer’s manual 
of English literature. Volume 3. William 
Pickering: London. 


55 


Bean, Sphaeromorphaea and Ethuliopsis 

Merrill, E.D. & Rolfe, R. A. (1908). Notes on Philippine 
Botany. The Philippine Journal of Science 3: 
95-127. 

Merxmueller, H., Leins, P. & Roessler, H. (1978). 
‘Inuleae - systematic review’. In V.H. Heywood 
et al. (eds.). The Biology and Chemistry of the 
Compositae, pp. 577-602. Academic Press: 
London, New York, San Lrancisco. 

Moore, C. & Betche, E. (1893). Handbook of the Flora of 
New South Wales. Government Printer: Sydney. 

Mueller, F. (1861). Ethuliopsis. Fragmenta 
Phytographiae Australiae 2(16): 154. 

Government Printer: Melbourne. 

Mukherjee, S.K. & Sarkar, A.K. (2001). Morphological 
diversity of pappus in the subfamily Asteroideae 
(Asteraceae). In J.K. Maheshwari (ed.). Recent 
Researches in Plant Anatomy and Morphology , 
pp. 275-294. Scientific Publishers: Jodhpur, 
India. 

Naithani, B.D. (1995). Tribe Anthemideae. In Flora 
of India 12: 3-75. Botanical Survey of India: 
Calcutta. 

National Library Of Australia (2009-onwards). 
TROVE, http://trove.nla.gov.au/ Accessed 
November 2012. 

Peng, C. (2004). Digital Flora of Taiwan. http://www. 
efloras.org/flora_page.aspx?flora_id=100 
Accessed 29 January 2012. 

Retief, E. & Herman, PP J. (1997). Plants of the northern 
provinces of South Africa: keys and diagnostic 
characters. Strelitzia 6: 1-681. 

Short, P.S. & Wilson, P.G. (1990). Haegiela , a new 
genus of Australian Asteraceae (Inuleae: 
Gnaphaliinae), with notes on the genus Epaltes 
Cass. Muelleria 7: 259-265. 


Short, P.S., Albrecht, D.E., Cowie, I.D., Lewis, D.L. & 
Stuckey, B.M. (2011). Checklist of the Vascular 
Plants of the Northern Territory, May 2011. 
Northern Territory Herbarium: Department 
of Natural Resources, Environment, The Arts 
and Sport, http://www.nt.gov.au/nreta/wildlife/ 
plants_herbarium/pdf/200701nt_checklist.pdf 
Accessed 3 March 2012. 

Sonder, O.W. (1853). Compositae. In Plantae 
Muellerianae, Beitrag zur Flora Sudaustraliens, 
aus den Sammlungen des Dr. Ferd. Muller. 
Linnaea 25: 450-656. 

Soons, M.B., Vlugt, C. Van Der, Lith, B. Van, Heil, 
G.W. & Klaassen, M. (2008). Small seed size 
increases the potential for dispersal of wetland 
plants by ducks. Journal of Ecology 96: 619— 
627. 

Stephens, K. & Sharp, D. (2009). Flora of North 
Stradbroke Island. Queensland Herbarium: 
Brisbane. 

Vallance, T.G., Moore, D.T. & Groves, E.W. (2001). 
Nature’s Investigator, The Diary of Robert 
Brown in Australia. Australian Biological 
Resources Study: Canberra. 

Wallich, N. (1831). Numerical List of Dried Specimens 
of plants in the Muse um of the Honl. East India 
Company , numbers 2604-4877. East India 
Company: London. 

Walsh, N.G. (2001). A revision of Centipeda 
(Asteraceae). Muelleria 15: 33-64. 

Walsh, N.G. & Stajsic, V. (2007). A Census of the 
Vascular Plants of Victoria , 8 th edition. Royal 
Botanic Gardens, Melbourne, http://www.rbg. 
vic.gov.au/viclist/ Accessed 3 March 2012. 


56 


Austrobaileya 9(1): 30-59 (2013) 


Map 2. Australian distribution (native and naturalised) of Sphaeromorphaea australis. 


Bean, Sphaeromorphaea and Ethuliopsis 


57 


Map 3. World distribution (native and naturalised) of Sphaeromorphaea australis. 


Map 4. World distribution of Sphaeromorphaea harrisii. 


58 


Austrobaileya 9(1): 30-59 (2013) 


Map 6. World distribution (native and naturalised) of Sphaeromorphaea littoraJis. 


Bean, Sphaeromorphaea and Ethuliopsis 


59 


160°E 


Map 7. World distribution of Sphaeromorphaea subintegra. 


25°S 


30°S 


35°S 


150°E 


Map 8. Distribution of Ethuliopsis cunninghamii. 


Reinstatement of intraspecific taxa for Bosistoapentacocca 
(F.Muell.) Baill. (Rutaceae) with a new combination B. 
pentacocca subsp. connaricarpa (Domin) P.I.Forst. 

Paul I. Forster 

Summary 

Forster, P.I. (2013). Reinstatement of intraspecific taxa for Bosistoa pentacocca (F.Muell.) 
Baill. (Rutaceae) with a new combination B. pentacocca subsp. connaricarpa (Domin) RI.Forst. 
Austrobaileya 9(1): 60-65. Geographic variation in morphology of Bosistoa pentacocca is discussed 
and the recognition of subspecies is proposed with the new combination B. pentacocca subsp. 
connaricarpa (Domin) RI.Forst. Typification of the name Bosistoa connaricarpa Domin is reviewed 
with the holotype designated as being at K. 

Key Words: Rutaceae, Bosistoa , Bosistoa pentacocca subsp. connaricarpa , Australia flora, 
Queensland flora, identification key, taxonomy, subspecies, John Dallachy 

P.I.Forster, Queensland Herbarium, Department of Science, Information Technology, Innovation 
and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. 
Email: Paul. Forster@science. dsitia.qld.gov. au 


Introduction 

Bosistoa pentacocca (F.Muell.) Baill. as 
defined by Hartley (2013), is a variable 
species from eastern Australia with numerous 
geographically discrete population centres 
(Map 1). This revised his initial analysis of 
these same populations as comprising one 
species with three varieties, with each variety 
based on discrete population centres and 
defined on leaf morphology (Hartley 1977). 
In this instance Hartley’s use of variety was 
equivalent to that of subspecies (sensu Forster 
2005; Duretto & Forster 2007) wherein 
the relevant taxon is confined to a discrete 
geographic region and is morphologically 
distinct. Hartley’s (2013) more conservative 
taxonomic conclusion was based on the greater 
number of collections now available and that 
the previously perceived differences between 
taxa were no longer valid. Diverse chemical 
evidence also tends to support a single 
species, albeit with some variation between 
populations (Auzi et al. 1997; Brophy et al. 
2007), although both these studies sampled 
only a small number of populations. 


Accepted for publication 26 August 2013 


Notwithstanding Hartley’s (2013) 
conclusion, there remain clear morphological 
differences between the most southern and 
the other population centres, principally 
the number of leaflets in each leaf and the 
number of secondary (lateral) veins per side 
of the primary (midrib) vein in each leaflet. 
These differences are easily discernible both 
in the field and on herbarium material and 
persist in material of known origin cultivated 
in a common garden. Although occasional 
collections exhibit divergence towards the 
other morphotype, the overall differences 
are consistent within a given geographic 
area (Map 1). My conclusion is that two 
intraspecific taxa can still be recognised 
within Bosistoa pentacocca (maintaining 
the status quo of Forster 1997, 2002, 2007, 
2010; Harden et al. 2006); however, I prefer 
the rank of subspecies rather than variety 
as the latter taxonomic category is more 
suited to within population variation. The 
existence of some populations (see below) 
exhibiting intermediate combinations of 
characters are to be expected and demonstrate 
continuity as required by Stebbins (1950), viz. 
“subspecies....connected....by a series of 
intergrading forms”. 


Forster, Bosistoa pentacocca 

If a metapopulation concept (Hanski 
& Gilpin 1991; Freckleton & Watkinson 
2002) is applied to Bosistoa pentacocca 
then it can be hypothesised that the species 
consists of at least six, geographically 
discrete, metapopulations with B. pentacocca 
subsp. pentacocca existing as a single 
metapopulation (1 on Map 1). Each of the 
metapopulations comprise few to many 
populations or subpopulations in ‘core’, 
possibly refugial areas, with occasional outlying 
populations/subpopulations. Applying this 
hypothesis, the ‘core’ area for B. pentacocca 
subsp. pentacocca exists within the Border 
Ranges (McPherson/Macleay overlap) centre 
of plant species richness and endemism of 
southeast Queensland and northeast New 
South with a concentration of populations 
on the basalt flows associated with the Mt 
Warning caldera and an ‘outlier’ population 
at Mt French, Boonah. Five metapopulations 
can be hypothesised for B. pentacocca subsp. 
connaricarpa (viz. 2. Mt Dryander - Conway 
Range area; 3. Eungella - Kuttabul area; 4: 
Rockhampton area; 5. Bulburin - Miriam 
Vale area; 6. Mt Urah - Cold Creek area (Map 
1). These proposed metapopulations remain 
to be tested using molecular data, although it 
is perhaps likely that most are refugial in a 
genetic sense (i.e. with little existence of active 
gene flow apart from within catchments). 


61 

Materials and methods 

Populations of Bosistoa pentacocca 
throughout its known range in Queensland 
were examined in the field. Herbarium 
collections at BRI and CANB were reviewed, 
plus online or scanned images of selected 
specimens at K and MEL. 

Common abbreviations in the specimen 
citation text include NP (National Park), SF 
(State Forest) and TR (Timber Reserve). 

Taxonomy 

Bosistoa pentacocca (F.Muell.) Baill., Hist. 
PI. 4: 470 (1873); Euodiapentacocca F.Muell., 
Fragm. 3: 41 (1862); B. sapindiformis F.Muell. 
ex Benth., FI. Austral. 1: 359 (1863), nom. 
illeg.', Acradenia bosistoi F.Muell., Pap. 
& Proc. Roy. Soc. Tasmania 8: 8 (1868) 
& Fragm. 6 167 (1868), nom. illeg. Type: 
New South Wales. Clarence River, s.dat., H. 
Beckler s.n. (holo: MEL 67499, online image!; 
iso: K 000717477, online image!, P 04355028, 
online image!). 

The species description is as per Hartley 
(2013) and two subspecies are recognised here. 
The specimens cited are representative of the 
six proposed metapopulations (numbered as 
above). 


Key to the subspecies of Bosistoa pentacocca 

1 Leaves with 3-5(-6) leaflet pairs; leaflets with 16-30 secondary (lateral) 

veins per side of primary (midrib) vein.B. pentacocca subsp. pentacocca 

1. Leaves with 1 or 2 (rarely 3) leaflet pairs; leaflets with 13—16(—18) 
secondary (lateral) veins per side of primary (midrib) 


vein. 

Bosistoa pentacocca subsp. pentacocca 

B. pentacocca var. pentacocca Hartley, J. 
Arnold Arbor. 58: 423 (1977). 

Leaves imparipinnate, 14-45 cm long, with 
3-5 (rarely 6) leaflet pairs; terminal leaflet 
usually oblanceolate, more rarely elliptic- 
oblong, subterminal leaflets oblanceolate, 
elliptic-oblong to narrowly elliptic, margins 
of all leaflets crenate to coarsely serrate, 
rarely entire, with 16-30 secondary (lateral) 
veins per side of primary (midrib) vein. 


.B. pentacocca subsp. connaricarpa 

Selected specimens examined'. Queensland. Moreton 
District: Ormeau, corner of Barrenjoey & Upper 
Ormeau Roads, Jan 1995, Forster PIF16086 & Leiper 
(BRI); NW slopes of Mt French, Apr 1982, Bird s.n. 
(BRI [AQ339215]); Mt French, NW slopes, 7.8 km 
WNW of Boonah, Jan 1983, Guymer 1847A & Jessup 
(BRI, CANB, MEL, NSW); Tambourine Mt, in Guanaba 
Gorge, Mar 1947, Blake 17387 (BRI); Numinbah Forest 
Reserve, c. 15 km SW of Nerang, southern slope of 
Pages Pinnacle, Jan 2006, Halford Q8958 (BRI); Upper 
Currumbin Creek, Tomewin Range, Oct 1981, Williams 
81250 (BRI); Nicholl Scrub NP, Currumbin Valley, Feb 
2011, Forster PIF37846 & Fernando (BISH, BRI, MEL); 
Egg Rock/Prison Farm track Nixons Creek, Lamington 


62 

NP, Nov 1993, Grimshaw G168 (BRI); Cave Creek, 
Natural Bridge [Springbrook] NP, Numinbah Valley, 
Mar 1978, Field s.n. (BRI [AQ264150], New South 
Wales. North Coast: Booyong Recreation Reserve, Big 
Scrub Remnant, Feb 1987, Floyd s.n. (BRI [AQ495572]). 

Distribution and habitat : Bosistoa 
pentacocca subsp. pentacocca is endemic 
to southeast Queensland (northern limit at 
Ormeau) and northeast New South Wales 
(south to the Clarence River) (Map 1). A 
western, outlying population occurs at Mt 
French, Boonah. It is found in rainforest 
(araucarian notophyll to microphyll 
vineforest) on diverse substrates (basalts, 
metasediments) predominantly on alluvium 
and colluvium along ephemeral watercourses. 

Notes : The most commonly encountered 
morphological state for this subspecies has 
plants with leaves with 4 or 5 leaflet pairs and 
predominantly oblanceolate leaflets that are 
always toothed. Occasionally the leaflets are 
more or less entire, but they are usually mixed 
in with toothed leaflets on the same branch. 
Fertile branchlets generally have leaflets at the 
larger end of the recorded dimensions whereas 
in Bosistoa pentacocca subsp. connaricarpa , 
the leaflets are usually much smaller and tend 
to be elliptic in shape. 

Bosistoa pentacocca subsp. connaricarpa 
(Domin) P.I.Forst., comb, et stat. nov.; B. 
connaricarpa Domin, Feddes Repert. Spec. 
Nov. Regni Veg. 12: 390 (1913); B. pentacocca 
var. connaricarpa (Domin) T.G.Hartley, J. 
Arnold Arbor. 58: 424 (1977). Type citation: 
“Queensland: Rockhampton, leg. Dallachy”. 
Type: Queensland. Port Curtis District: 
Rockhampton, 12 December [1862], [J. 
Dallachy s.n.] (holo: K, image!). 

Bosistoa pentacocca var. dryanderensis 
T.G.Hartley, J Arnold Arbor. 58: 425 (1977). 
Type: Queensland. North Kennedy District: 
Mt Dryander, 11 August 1972, V.K. Moriarty 
968 (holo: CANB; iso: BRI). 

Leaves imparipinnate or pinnately trifoliate, 
9-30 cm long, with 1 or 2, rarely 3 leaflet 
pairs; terminal leaflet usually elliptic, 
subterminal leaflets oblanceolate to elliptic 
or narrowly elliptic, margins of all leaflets 
entire, or crenate to coarsely serrate, with 
13—16(—18) secondary (lateral) veins per side 


Austrobaileya 9(1): 60-65 (2013) 
of primary (midrib) vein. 

Additional selected specimens examined: Queensland. 
North Kennedy District: 2 *Portello’s Crossing, 
Dryander Creek, base of Mt Dryander, Feb 1994, Forster 
PIF14859 & Bean (A, BRI, CNS, MEL); 2 Sawmill 
Beach, Whitsunday Island [Whitsunday Islands NP], 
Sep 1990, Batianoff9009838 & Batianoff (BRI; BISH, 
LAE o.v.); impulse Creek, SF299 Conway, May 1991, 
Forster PIF8300 & McDonald (BRI, CNS, K, MEL). 
South Kennedy District: 3 Mt Blackwood, S side c. 

5 km from Kuttabul, Jun 1984, Champion 68 (BRI); 
3 Mt Adder, southern base. Mount Jukes NP, Apr 1991, 
Forster PIF8115 & McDonald (BRI, CANB, CNS, K, L); 
3 Camerons Pocket, 12 km W of Calen [Mia Mia SF], Jan 
2003, Hambleton 4 (BRI); 3 23 km S of Eungella Dam 
Road on Cockies Creek Road, Crediton SF, Sep 2002, 
Gumming 21133 (BRI). Port Curtis District: 4 Black 
Creek, SW base of Mt Sleipner, Mar 1989, Forster 
P1F4999 & Hoy (BRI, CANB, CNS, K, L, MEL, MO, 
NSW, US); 5 TR115 [now Castle Tower NP], Mt Coulston, 
9 km NNW of Bororen, Sep 1989, Gibson TOI828 (BRI); 
5 Bindaree, NE of Miriam Vale & just S of Arthurs Seat, 
Jun 1995, Meter RM911 (BRI); 5 TR122, Wietalaba [now 
Wietalaba NP], 32 km S of Calliope, Oct 1989, Gibson 
TO1740 (BRI, CANB); 5 TR202 [now Bulburin NP], Feb 
1995, Forster P1F16176 (BRI, CNS, MEL); 5 2.5 km N 
of Glassford Vale, TR353 [now Dawes NP], Sep 1989, 
Forster PIF5674 & Bean (BRI, CANB); 5 Bulburin NP, 
Pine Creek headwaters, Jan 2009, Forster PIF34931 

6 Brushe (BRI, MEL); 5 Bulburin NP, Cassilus Creek, 
Jan 2009, Forster P1F34944 et al. (BRI, CNS). Burnett 
District: 5 SF 695 Kalpowar, Mt Fort William, Mar 
2000, Forster PIF25439 & Booth (BRI, CNS). Wide 
Bay District: 6 SF50 Glenbar, 1 km WSW of Mt Urah 
summit [now Glenbar NP], Feb 1993, Forster PIF 13130 
& Machin (BRI); 6 Glenbar NP, Cold Creek catchment, 
8 km NW of Miva, Dec 2008, Forster PIF34782 et al. 
(BRI). * superscript number refers to the metapopulation 
in Map 1 

Distribution and habitat : Bosistoa 
pentacocca subsp. connaricarpa occurs in 
eastern Queensland in five metapopulations 
from Mt Dryander to northeast of Gympie 
(Map 1). It invariably occurs in rainforest 
(microphyll to notophyll vineforest, often with 
Araucaria cunninghamii Aiton ex A.Cunn.) 
on diverse substrates (andesites, basalts, 
granites, granodiorites), often on alluvium or 
colluvium along ephemeral watercourses. 

The type of B. pentacocca subsp. 
connaricarpa is from ‘Rockhampton’; and 
the subspecies has only been recollected from 
the Mt Sleipner locality in this general area, 
thus this proposed metapopulation (4 on Map 
1) consists of a single population. By contrast 
the other metapopulations are more diverse (2 
has at least seven populations, 3 has at least 


Forster, Bosistoa pentacocca 

four populations, 5 has at least 13 populations 
and 6 has at least two populations). 

Typification : Domin (1913) cited a 

single collection in the protologue for 
Bosistoa connaricarpa, viz. “Queensland: 
Rockhampton, leg. Dallachy”. Earlier he 
had stated “I give descriptions of new forms 
according to specimens preserved in the 
herbarium of the Royal Botanic Gardens at 
Kew” when basing names on collections other 
than his own from Australia (Domin 1911). It 
can be concluded from this that a holotype for 
this name should be at Kew. 

Hartley (2013) gives the type of this taxon as 
“Rockhampton, Qld, 5 Feb. 1863, J.Dallachy, 
iso: MEL”. Earlier he stated “Type: Dallachy, 
February 5,1863, Queensland, Rockhampton” 
with an isotype at MEL (Hartley 1977). There 
are two sheets at MEL (67497 & 67503) of 
this collection, both labelled as an isotype by 
Hartley in January 1977. Both of these sheets 
have original collection labels from Dallachy 
with his collection number #297 and the 
locality of “Devils Creek”, but no mention of 
Rockhampton. 

There exists atK a sheet with a ‘ Phytologic 
Museum of Melbourne’ label stating “Bosistoa 
sapindiformis F.M. Rockhampton” (although 
the specimen can now be clearly identified as 
Bosistoa pentacocca subsp. connaricarpa). 
The handwriting on this label is Mueller’s. 
Whilst there is no mention of a collector or 
specific date on this label, further evidence 
can be found in a field label attached to the 
bottom left corner of the sheet. This field 
label is written in Dallachy’s hand using blue 
crayon and amongst various points on plant 
habit, states “12 Dec”. Dallachy is known to 
have made collections in the Rockhampton 
area “at least from the 5 th [of] January 1863” 
to “at least April 16 th ” (Blake 1954) and this 
field label indicates that he was active there 
three weeks earlier. Prior to the Rockhampton 
area, Dallachy collected about Brisbane 
(Blake 1954). 

It is unlikely that Domin saw any of the 
MEL sheets discussed and although labelled 
as isotypes by Hartley they are not duplicates 
of the K material as they were collected on 


63 

a different date. Mueller distributed material 
sent to him by collectors to many herbaria, 
often with inadequate or different labels 
to that of those that were retained in MEL. 
Collectors such as Dallachy also often 
recollected material and it can be difficult to 
ascertain the specific collections subsequently 
cited by Mueller and others as they were 
generally unmounted in the 1800s and often 
with multiple sheets. 

Precisely where Dallachy made his 
“Rockhampton” collection can probably 
never be resolved; however, it is likely that he 
returned to the same vicinity on subsequent 
occasions and so the collections by him of this 
plant in K and MEL may be from the same 
spot. His locality “Devils Creek” is possibly 
equivalent to Frenchmans Creek in a modern 
context for the Rockhampton area as this 
ephemeral watercourse flows into the Fitzroy 
River at Devils Elbow. The known locality 
for Bosistoa pentacocca at the southwestern 
base of Mt Sleipner is on the eastern side of 
Black Creek that is east of Mt Berserker and 
only about 3 km in a direct line from where 
Dallachy may have collected his specimens. 
Suitable habitat still exists in the upper 
reaches of Frenchmans Creek and should 
be explored to determine if B. pentacocca 
populations persist. 

Notes : Several populations of this 
subspecies occasionally exhibit characteristics 
of B. pentacocca subsp. pentacocca. These 
are at Kalpowar, Bulburin and Mt Urah (all 
at the southern end of the northern subspecies 
range) where the leaves can occasionally have 
three leaflet pairs; however, these are mixed 
in with leaves predominantly of 1 or 2 leaflet 
pairs. 

Acknowledgements 

I am very grateful to David Goyder (Royal 
Botanic Gardens Kew) for locating the 
Dallachy material in the general herbarium 
and Lucia Lopez Poveda for photographing 
material. Elucidation of Dallachy’s 
handwriting (scrawl) was assisted by 
Peter Bostock, Gordon Guymer and Will 
Smith (BRI). The population of Bosistoa 
pentacocca at Black Creek was located with 


64 


the assistance of Dianne & Neil Hoy in 1989. 
Useful discussions of possible localities for 
Dallachy’s ‘Devils Creek’ were held with Neil 
Hoy and Susan Cunningham. Map 1 was 
prepared by Will Smith. 

References 

Auzi, A.A., Forster, P.I., Gray, A.I., Hartley, T.G., 
Waigh, R.D. & Waterman, P.G. (1997). 
Phloroglucinol derivatives from leaves of 
Bosistoapentacocca. Phytochemistry 45: 1673— 
1678. 

Blake, S.T. (1954). Some pioneers in plant exploration 
and classification. Proceedings of the Royal 
Society of Queensland. 66: 1-19. 

Brophy, J.J., Goldsack, R.J. & Forster, PI. (2007). 
Essential oils from the leaves of Bosistoa 
F.Muell. ex Benth. (Rutaceae). Journal of 
Essential Oil Research 19: 249-254. 

Domin, K. (1911). First contribution to the Flora of 
Australia. Feddes Repertorium Specierum 
Novarum Regni Vegetabilis 9: 550-553. 

- (1913). Eighth contribution to the Flora of 

Australia. Feddes Repertorium Specierum 
Novarum Regni Vegetabilis 12: 388-390. 

Duretto, M.F. & Forster, PI. (2007). A taxonomic 
revision of the genus Zieria Sm. (Rutaceae) in 
Queensland. Austrobaileya 7: 473-544. 

Forster, PI. (1997). Rutaceae. In R. J.F. Henderson (ed.), 
Queensland Plants - Names and Distribution , 
184-188. State of Queensland, Department of 
Environment: Brisbane. 

- (2002). Rutaceae. In R.J.F. Henderson (ed). 

Names and Distribution of Queensland Plants, 
Algae and Lichens , pp. 177-181. Queensland 
Government, Environmental Protection 
Agency: Brisbane. 

- (2005). New species of Philotheca Rudge 

(Rutaceae) from Queensland. Austrobaileya 7: 
175-181. 


Austrobaileya 9(1): 60-65 (2013) 

Freckleton, R.P & Watkinson, A.R. (2002). Large-scale 
spatial dynamics of plants: metapopulations, 
regional ensembles and patchy populations. 
Journal of Ecology 90: 419-434. 

Hanski, I. & Gilpin, M. (1991). Metapopulation 
dynamics: brief history and conceptual domain. 
Biological Journal of the Linnean Society 42: 
3-16. 

Harden, G., McDonald, W.J.F. & Williams, J. (2006). 
Rainforest Trees and Shrubs. A field guide to 
their identification. Gwen Harden Publishing: 
Nambucca Heads. 

Hartley, T.G. (1977). A revision of the genus Bosistoa 
(Rutaceae). Journal of the Arnold Arboretum 
58: 416-436. 

- (2013). Bosistoa. In A.J.G. Wilson (ed.). Flora of 

Australia 26: 48-52. ABRS/CSIRO Publishing: 
Melbourne. 

Stebbins, G.L. (1950). Variation and Evolution in Plants. 
Columbia University Press: New York. 


- (2007). Rutaceae. In P.D. Bostock & A.E. 

Holland (eds.), Census of the Queensland Flora 
2007 ; pp. 180-183. Queensland Herbarium, 
Environmental Protection Agency: Brisbane. 

- (2010). Rutaceae. In P.D. Bostock & A.E. 

Holland (eds.), Census of the Queensland Flora 
2010 , pp. 174-178. Queensland Herbarium, 
Department of Environment and Resource 
Management: Brisbane. 


Forster, Bosistoa pentacocca 


65 


Map 1 . Distribution of Bosistoa pentacocca in eastern Australia. Numbered areas refer to proposed metapopulations 
(1 -B.pentacocca subsp. pentacocca A, outlier population at Mt French arrowed; 2-6 -B. pentacocca subsp. 
connaricarpa •. 


Three new species of Pluchea Cass. (Asteraceae: 
Inuleae-Plucheinae) from northern Australia 

A.R. Bean 

Summary 

Bean, A.R. (2013). Three new species of Pluchea Cass. (Asteraceae: Inuleae-Plucheinae) from 
northern Australia. Austrobaileya 9(1): 66-74. Three new species of Pluchea Cass, are described; 
P. longiseta A.R.Bean from northern Western Australia, P. mesotes A.R.Bean from the Northern 
Territory, and P. alata A.R.Bean from central Queensland. All species are illustrated and distribution 
maps are provided. A revised key to the Australian of Pluchea species is provided. 

Key Words: Asteraceae, Pluchea , Pluchea alata , Pluchea longiseta , Pluchea mesotes , Australia flora, 
taxonomy, identification key 

A.R. Bean, Queensland Herbarium, Department of Science, Information Technology, Innovation 
and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. 
E-mail: Tony.Bean@science.dsitia.qld.gov.au 


Introduction 

Pluchea Cass, was revised for Australia by 
Hunger (1997), with seven species. Bean (2011) 
named two additional species of Pluchea 
from Queensland. Further investigations 
have revealed three additional species, 
which are named here: P. alata is endemic 
to the Aramac area of central Queensland, P. 
longiseta occurs in coastal and near-coastal 
parts of north-western Western Australia, and 
P. mesotes occurs in coastal parts of north¬ 
eastern Northern Territory. 

Materials and methods 

This study is based on herbarium specimens 
from BRI, DNA, MEL and PERTH, and (for 
P. alata) field investigations. Measurements of 
florets were made from material preserved in 
spirit or reconstituted in boiling water, while 
leaves, bracts and achenes were measured 
from dried specimens. 

Common abbreviations in the text include 
NT (Northern Territory), Qld (Queensland), 
SA (South Australia), WA (Western Australia). 


Accepted for publication 25 July 2013 


Taxonomy 

Pluchea alata A.R.Bean sp. nov. affinis P. 
rubelliflorae (F.Muell.) B.L.Rob. sed capitulo 
cylindrico sessili vel subsessili, flosculis 
marginalibus prope horum apices expansis, 
setis pappi in flosculis disci paucioribus et 
alis caulis usque ad 1.2 mm latis differens. 
Typus: Queensland. Mitchell District: First 
Spring, Edgbaston Reserve, NE of Aramac, 6 
April 2012, A.R. Bean 31641 (holo: BRI; iso: 
CANB, MEL). 

Woody shrub to 70 cm high and 120 
cm across. Stems erect to spreading, 
well-branched, glabrous throughout, 
conspicuously winged, wings 0.5-1.2 mm 
wide. Leaves sessile, elliptical, spreading, 
slightly fleshy, decurrent, 15-30 x 4.2-7 mm, 
3.5-4.5 times longer than broad, eglandular, 
glabrous; margins denticulate; apex acute. 
Capitula cylindrical, 5-6.5 x 2.5-3 mm, 
in terminal clusters of 2-4; peduncles 0-1 
mm long. Involucral bracts in several rows; 
outer bracts ovate, 1.2-2 x 0.4-0.6 mm, with 
apex acute, outer surface glabrous, margins 
entire, not ciliate and glabrous; median 
bracts lanceolate, 2-3.5 x 0.3-0.6 mm, 
apex acute, outer surface glabrous, margins 
glabrous; inner bracts linear, 3.7-4 x 0.8- 
0.9 mm, apex acute, outer surface glabrous, 
upper margins not ciliate, entire. Receptacle 
epaleate, verrucose, glabrous, flat, 1-1.2 


Bean, Pluchea 


67 


Fig. 1. Pluchea alata. A. flowering branchlet xl.5. B. leaf showing decurrent base and winged stem *4. C. flowering 
capitulum xl2. D. disc floret xl6. E. outer floret x24. F. achene x48. A & B from Bean 31648 (BRI); C-E from Bean 
31641 (BRI); F from Fairfax s.n. (BRI [AQ875934]). 


68 

mm diameter. Outer florets 55-75, female; 
corolla filiform, but expanded (0.2-0.25 mm 
wide) towards apex, 2.8—3.1 mm long, white; 
lobes 3 or 4, equal, triangular, 0.15-0.25 
mm long, glabrous; style bifid, exserted, 
glabrous; pappus corona present, erect, 
cylindrical, c. 0.03 mm long; pappus bristles 

4- 12, 2.4-2.8 mm long, caducous, in a single 
whorl, not coherent at base, all of similar 
length, barbellate throughout;. Disc florets 

5- 13, ovary fully developed but infertile, 
pink; corolla cylindrical, 3-3.4 mm long, 
pink; lobes 5, triangular, 0.4-0.6 mm long, 
glabrous; anthers c. 1.5 mm long, including 
tail c. 0.15 mm long; style undivided, with 
obtuse sweeping hairs extending below 
the furcation;. Achenes developing only 
from outer florets; narrowly-ellipsoidal, 
0.7-0.85 mm long, brown, glabrous; surface 
obscurely longitudinally striate; carpopodium 
prominent, white. Fig. 1. 

Additional specimens examined: Queensland. 

Mitchell District: Bowen Downs, in 1874, Birch s.n. 
(MEL); Big Spring, Edgbaston Station, N of Aramac, 
Apr 2005, Fairfax s.n. (BRI [AQ875934]); Edgbaston, 
NE of Aramac, Sep 2006, Fensham 5549 (BRI); 
Edgbaston Station 33 km NNE of Aramac, Apr 2010, 
Thompson MUT410 et al. (BRI, PE); Measuring Spring, 
Edgbaston Reserve, NE of Aramac, Apr 2012, Bean 
31648 (BRI, NSW, US). 

Distribution and habitat: Pluchea alata is 
endemic to Queensland and is known from 
Edgbaston Reserve, NE of Aramac, and an 
old record from nearby Bowen Downs station 
(Map 1). It inhabits the margins of artesian 
springs, on saline soils. 

Phenology : Flowers and fruits have been 
recorded for April and September. 

Affinities: This species is allied to Pluchea 
rubelliflora , but differs by the woody stems, the 
relatively broader leaves (length 3.5-47 times 
width versus 4.5-12 times for P. rubelliflora ), 
the more or less cylindrical capitula that are 
sessile or with peduncles up to 1 mm long 
(hemispherical capitula on peduncles (1—)2—10 
mm long for P. rubelliflora ); the outer florets 
expanded towards apex, with the lobes 0.15— 
0.25 mm long (florets not expanded and with 
lobes 0.08-0.12 mm long for P. rubelliflora ); 
and the disc florets with 3-6 pappus bristles 
(6-18 bristles for P. rubelliflora). The stem 


Austrobaileya 9(1): 66-74 (2013) 

wings are up to 1.2 mm wide, whereas for P. 
rubelliflora in the Aramac - Barcaldine area, 
the stem wings are 0.2-0.4 mm wide. 

Note: ‘Bowen Downs’ was a much larger 
station when this species was collected there 
in 1874. In view of the habitat occupied by this 
species, Birch’s collecting locality was most 
likely in the Lake Huffer area (R. Fensham, 
pers. comm. Feb 2013). 

Conservation status: The species is known 
from fewer than 100 plants at the type locality. 
Applying the Red List criteria (IUCN 2012), 
a category of Endangered is recommended 
(Criterion D). 

Etymology: The epithet is from the Latin 
alatus meaning winged, in reference to the 
prominently winged stems of this species. 

Pluchea longiseta A.R.Bean sp. nov. ab 

omnibus aliis speciebus Plucheae foliis 
carnosis, flosculis achenisque longioribus 
et pappi setis in verticillis duobus tribusve 
dispositis differens. Typus: Western 
Australia. North shore of Willie Creek inlet, 
Dampierland Peninsula, 6 September 1985, 
K.F. Kenneally 9526A (holo: BRI; iso: CANB, 
DNA, PERTH). 

Pluchea sp. A; Wilson (1992: 950). 

Pluchea sp. B; Wilson (1992: 950). 

Pluchea sp. B Kimberley Flora (K.F. 
Kenneally 9526A); Western Australian 
Herbarium (1998-). 

Woody shrub to 60 cm high and 80 cm 
across. Stems terete, erect to spreading, 
well-branched, glabrous or with very sparse 
multicellular eglandular hairs near growing 
point. Leaves sessile, narrowly-oblanceolate 
to linear, spreading, fleshy, not decurrent, 
28-52 x 2-5.5 mm, 9-14 times longer than 
broad, eglandular, multicellular hairs sparse 
or absent; margins entire; apex acuminate or 
apiculate. Capitula in small terminal clusters, 
narrowly campanulate to cylindrical, 12-15 
x 8-12 mm; peduncles 2-27 mm long, with 
1-3 lanceolate bracts near the apex, each 2-4 
mm long. Involucral bracts in several rows; 
outer bracts ovate, 2.5-5.5 x 1.5-3 mm, apex 
acuminate, outer surface glabrous, margins 


Bean, Pluchea 


69 


entire and ciliate; median bracts lanceolate, 
6.5-9.5 x 2.3-3 mm, apex acute to acuminate, 
outer surface glabrous, margins glabrous or 
with sparse eglandular hairs; inner bracts 
linear, 10.5-12 x 0.7-1.1 mm, apex laciniate, 
outer surface glabrous, upper margins 
glabrous, not ciliate. Receptacle epaleate, 
verrucose, glabrous, flat to concave, 1.7-2 
mm diameter. Outer florets 100-200, female; 
corolla filiform, 8-9.5 mm long, pink; lobes 3, 
equal, triangular, 0.2-0.3 mm long, glabrous; 
style bifid, exserted, glabrous; pappus 8.2-9.2 
mm long, pappus bristles 25-40, in 2 or 3 
whorls, coherent at base, all of similar length. 
Disc florets 12-22, ovary fully developed but 
infertile, pale yellow; corolla cylindrical, 7.9- 
9.3 mm long, pink; lobes 4(-5), triangular, 
0.5-0.6 mm long, glandular; anthers c. 3 mm 
long, including tail c. 0.2 mm long; style bifid, 
with obtuse sweeping hairs extending below 
the furcation. Achenes narrowly-ellipsoidal, 
1.3-1.5 mm long, brown, with numerous 
appressed twin hairs; longitudinal ribs 4-6; 
carpopodium prominent, white. Fig. 2. 

Additional specimens examined : Western Australia. 
Montgomery Island, E of Koolan Island, May 1998, 
Mitchell 5381 (DNA, PERTH); Cygnet Bay, Nov 1906, 
Fitzgerald 1706 (PERTH); Buckley Plain, north of town 
boundary, Broome, Feb 2005, Byrne 1374 (PERTH); 
Buckley Plain, 6 km NE of Broome, Nov 1987, Foulkes 
442 (DNA); c. 9 km NE of Mandora Homestead, 
between Broome and Port Hedland, Aug 1997, Mitchell 
PRP1787 (PERTH); Loam 2 site, Mandora Marsh area, 
Oct 1999, Duero WEL2-10 et al (DNA, PERTH); NW 
Island, Montebello Islands, Oct 2000, Kenneally 11545 
(PERTH); Hermite Island, Montebello Islands, Oct 
2000, Kenneally 11573 (PERTH); below tip at side of 
DSI lease, Dampier Island, Sep 1987, Glennon 397K 
(PERTH); Boodie Island, S of Barrow and Middle 
Islands, Sep 2004, Long VL360-3 (PERTH); Duffers 
Bore, 24 km N of Dampier Downs, Aug 1976, Kenneally 
5782 (DNA, PERTH); between T-bone Bay and hill 
towards lighthouse, Oct 1975, Weber 4949 (PERTH); 
Willie Creek marsh, Jan 1985, Foidkes 97 (PERTH); 
just N of Dragon Tree Soak, Great Sandy Desert, Aug 
1977, George 14747 (BRI, PERTH); 80 Mile Beach, N of 
Wooroo Creek, Jul 1941, Burbidge 1300 (PERTH). 

Distribution and habitat : This species is 
endemic to Western Australia, mainly in 
coastal areas from the Montgomery Islands 
in the western Kimberley region, to the Cape 
Range, but also in a few scattered places 
away from the coast (Map 1). It inhabits 
samphire communities behind frontal dunes, 


saline marshy plains, and the margins of salty 
lagoons. Soils are usually pale and clayey but 
can have a sandy surface layer. 

Phenology : Flowers and fruits are recorded 
from May to February, with most collections 
made in September and October. 

Affinities : Because of the relatively large 
floral parts and the broad involucral bracts, 
this species resembles members of the genus 
Streptoglossa Steetz, and some specimens 
have previously been identified as such. But 
Streptoglossa species have ligulate outer 
florets and achenes with very dense twin hairs 
that conceal the surface. Pluchea longiseta has 
the tubular outer florets and almost glabrous 
achenes that are typical of this genus. 

Pluchea longiseta does differ from the usual 
characteristics of Pluchea in that the pappus 
bristles are arranged in two or three whorls. 
Pluchea usually has pappus bristles in a single 
whorl. P. longiseta can be distinguished from 
other Australian species of Pluchea also by 
the fleshy leaves, and the longer achenes, 
pappus and florets. 

Conservation status : This species is 
relatively widespread, but no surveys have 
been carried out to determine population 
sizes. Its habitat is potentially under threat 
from port development and infrastructure 
in some areas. A category of Data Deficient 
is appropriate under the IUCN guidelines 
(IUCN 2012). 

Etymology: The epithet longiseta refers to the 
pappus bristles that are longer in this species 
than in any other Australian Pluchea. 

Note: According to Western Australian 
Herbarium (1998-), the taxon “Pluchea sp. A” 
listed in Wilson (1992) is synonymous with 
“Pluchea sp. B”. 


70 


Austrobaileya 9(1): 66-74 (2013) 


Fig. 2. Pluchea longiseta. A. flowering branchlet xl. B. flowering capitulum x4. C. disc floret x8. D. outer floret x8. E. 
achene x32. A from Byrne 1374 (PERTH); B-D from Kenneally 9526A (PERTH); F from Kenneally 11573 (PERTH). 


Bean, Pluchea 


71 


Pluchea mesotes A.R.Bean sp. nov. affinis 
P. indicae (L.) Less, sed foliis angustioribus 
dentibus marginalibus paucis vel nullis, 
pedunculis capitulisque longioribus et pappi 
setis in quoque flosculo paucioribus differens. 
Typus: Northern Territory. Connexion Island, 
off Groote Eylandt, 29 September 1988, P.K. 
Latz 10901 (holo: BRI; iso: DNA). 

Multi-stemmed annual shrub to 100 cm 
high. Stems terete, erect to spreading, well- 
branched, with sparse to moderately dense 
multicellular eglandular hairs. Leaves sessile, 
narrowly-elliptical to spathulate, spreading, 
chartaceous, not resinous, glands absent or 
sparse, multicellular eglandular hairs sparse, 
bases not decurrent, 24-55 x 4-16 mm, 3.4- 
6.2 times longer than broad, margins entire 
or occasionally with 1-3 pairs of teeth, apex 
acute. Capitula in small terminal clusters, 
narrowly campanulate, 5.5-7 x 5-5.5 mm; 
peduncles 0-7 mm long, with 1-3 lanceolate 
bracts near the apex, each 1-2.5 mm long. 
Involucral bracts in several rows; outer bracts 
broadly ovate, 1.6-2.7 x 1—1.4 mm, with 
apex acute, outer surface glabrous, upper 
margins ciliate; median bracts lanceolate, 
3.2-3.8 x 0.8-1 mm, apex acute, outer surface 
glabrous, upper margins ciliate; inner bracts 
linear, 4.8-5.2 x 0.2-0.3 mm, upper margins 
ciliate (cilia 0.05-0.1 mm long), outer surface 
glabrous. Receptacle epaleate, verrucose, 
densely hairy, flat, 1.4-1.7 mm diameter. 
Outer florets 60-120, female; corolla filiform, 
4-4.5 mm long, colour unknown; lobes 3, 
equal, triangular, 0.1-0.15 mm long, glabrous; 
style bifid, exserted, glabrous; pappus 3.8-4.1 
mm long, pappus bristles 12-15, in a single 
whorl, coherent at base, all of similar length; 
Disc florets 2-5, ovary fully developed but 
infertile; corolla cylindrical, 4-4.3 mm long, 
white to pale yellow; lobes 5, triangular, 0.4- 
0.7 mm long, glandular; anthers c. 1.9 mm 
long, including tail c. 0.3 mm long; style bifid, 
with obtuse sweeping hairs extending below 
the furcation. Achenes narrowly-ellipsoidal, 
0.9-1 mm long, brown, with scattered 
appressed twin hairs; longitudinal ribs 2-5; 
carpopodium prominent, white. Fig. 3. 


Additional specimens examined : Northern Territory. 
Bennett Bay, Eastern Arnhem Land, Nov 1987, Russell- 
Smith 4219 & Lucas (BRI, DNA); Roper River, Sep 1867, 
Gulliver s.n. (MEL). 

Distribution and habitat: This species is 
apparently confined to the coastal parts of 
north-eastern Northern Territory (Map 1). 
It is recorded from “dune thicket”, and from 
“coral-shell gravel above mangroves”. 

Phenology: Flowers and fruits have been 
recorded for September and November. 

Affinities: Pluchea mesotes is related to P. 
indica but differs by the leaves 3.4-6.2 times 
longer than broad, that are without teeth or 
sometimes with a few small teeth (leaves 
1.7-3.3 times longer than broad and with 
numerous prominent teeth for P. indica ); the 
peduncles 0-7 mm long (versus 0-1 mm long 
for P. indica}, the longer capitula 5.5-7 mm 
long (versus 4-5.5 mm forP. indica)', the cilia 
on the upper margins of the inner involucral 
bracts 0.05-0.1 mm long (versus 0.2-0.3 mm 
long for P. indica)', and the 12-15 pappus 
bristles (versus 17-25 for P. indica). P. indica 
occurs in the Darwin - Kakadu area, and on 
Cape York Peninsula, but apparently not in 
the region occupied by P. mesotes. 

Conservation status: A category of Data 
Deficient is appropriate under the IUCN 
guidelines (IUCN 2012). 

Etymology: From the Greek mesotes, 
meaning ‘in the middle position’. This is 
reference to the geographical position of this 
species in Australia compared with P. indica. 
The epithet is treated as a noun in apposition. 

Acknowledgements 

I am grateful to the Directors of DNA, 
MEF and PERTH for the loan of specimens, 
to Peter Bostock for the Fatin diagnoses 
and to Wifi Smith for the illustrations and 
distribution map. Thanks to Murray Haseler 
for organising access to Edgbaston Reserve, 
and Rod Fensham for elucidating Birch’s 
probable collecting locality for P. alata. 


72 


Austrobaileya 9(1): 66-74 (2013) 


Key to the Pluchea species in Australia 


1 Leaves and stems glabrous (disregarding sessile glands) or with very few scattered hairs . 2 

1. Leaves and stems with frequent to abundant eglandular hairs or stalked glandular hairs. . 8 


2 Pappus bristles 8.2-9.2 mm long, in 2 or 3 whorls; leaves fleshy WA .P. longiseta 

2. Pappus bristles 1.8-5 mm long, in a single whorl; leaves not fleshy.3 

3 Leaves narrow, 4.5-23 times longer than wide.4 

3. Leaves broad, 1.5-47 times longer than wide.6 

4 Leaves with dark punctate glands; stems not winged.5 

4. Leaves without punctate glands; stems narrowly winged NT, Qld, SA, WA. . P. rubelliflora 

5 Capitula 6-7 mm long; disc florets 5-lobed; leaves 7-24 mm long; 

margins of involucral bracts fimbriate, eglandular NSW, Qld .P. baccharoides 

5. Capitula 7-10 mm long; disc florets 4-lobed; leaves 18-54 mm long; 

margins of involucral bracts with stalked glands Qld .P. punctata 

6 Stems winged; leaf glands absent Qld .P. alata 

6. Stems not winged; leaf glands present.7 

7 Leaves not resinous; capitula 4-5.5 mm long, clustered into sessile groups 

of (1—)2—4 NT, Qld .P. indica 

7. Leaves resinous; capitula 6-8.5 mm long, usually solitary; some peduncles 

on any given plant more than 3 mm long WA .P. tetranthera 

8 Hairs on leaves extremely dense, obscuring leaf surfaces NT, Qld, 

WA .P. ferdinandi-muelleri 

8. Leaf surfaces readily visible.9 


9 Hairs on leaves and stems predominantly gland tipped NSW, NT, Qld, SA, WA . . P. dentex 

9. Hairs on leaves and stems all or mostly eglandular.10 

10 Leaves narrowly-elliptic or oblanceolate, 3.4-6.4 times longer than wide.11 

10. Leaves obovate, 1.5-3.3 times longer than wide.12 


11 Leaf margins denticulate throughout; leaf surfaces with abundant, yellow 

glands; receptacle glabrous Qld .P. xanthina 

11. Leaf margins entire or with 1 or 2 pairs of teeth; leaf surface glands absent 

or sparse, not yellow; receptacle hairy NT .P. mesotes 


12 Capitula 5.5-75 mm long, not clustered, at least some peduncles more 

than 2 mm long; leaves often resinous NSW, NT, Qld, SA .P. dunlopii 

12. Capitula 4-5.5 mm long, clustered into sessile groups of (l-)2-4; leaves 

not resinous NT, Qld .P. indica 


Bean, Pluchea 


73 


Fig. 3. Pluchea mesotes. A. flowering branchlet x0.5. B. leaf *2. C. flowering capitulum x6. D. disc floret xl6. E. 
achene x32. A, C-E from Latz 10901 (BRI); B from Russell-Smith 4219 & Lucas (DNA). 


74 

References 

Bean, A.R. (2011). Two new species of Pluchea Cass. 
(Asteraceae: Plucheinae ) from Queensland, 
Australia. Austrobaileya 8: 340-346. 

Hunger, S. (1997). A survey of the genus Pluchea 
(Compositae, Plucheeae) in Australia. 
Willdenowia 27: 207-223. 

Iucn (2012). IUCN Red List Categories and Criteria , 
Version 3.1, 2 nd edn. IUCN - The World 
Conservation Union: Gland. 


Austrobaileya 9(1): 66-74 (2013) 

Western Australian Herbarium (1998-). FloraBase — 
the Western Australian Flora. Department 
of Environment and Conservation, http:// 
florabase.dec.wa.gov.au/Accessed 17 Feb 2013. 

Wilson, A.J.G. (1992). Pluchea. In J.R. Wheeler (ed.). 
Flora of the Kimberley Region. Department of 
Conservation and Land Management: Como, 
Western Australia. 


Map 1 . Distribution of Pluchea longiseta •, P.mesotes A and P. alata M. 


Cryptocarya cercophylla W.E.Cooper (Lauraceae), 
a new species from Queensland’s Wet Tropics 

W.E. Cooper 

Summary 

Cooper, W.E. (2013). Cryptocarya cercophylla W.E.Cooper (Lauraceae), a new species from 
Queensland’s Wet Tropics. Austrobaileya 9(1): 75-79. Cryptocarya cercophylla W.E.Cooper is 
described and illustrated. Notes on habitat, habit and distribution are provided. The new species is 
restricted to a small area near Boonjee in the Wet Tropics of north-eastern Queensland. 

Key Words: Lauraceae, Cryptocarya , Cryptocarya cercophylla , rainforest, Australia flora, 
Queensland flora. Wet Tropics bioregion, new species 

W.E. Cooper, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor 
Road, Smithfield, Queensland 4878, Australia 


Introduction 

Cryptocarya R.Br. (Lauraceae) is a 
pantropical genus of200-250 species of small 
to large trees, distributed from Africa, Asia, 
Malesia, Australia, the Pacific and South 
America (Le Cussan & Hyland 2007). In 
Australia, 48 species are now recorded, 39 or 
40 are endemic and 36 of those occur in the 
rainforests of tropical Queensland. 

Many Cryptocarya species are very 
restricted in distribution. Most species in 
Australia occur in various forms of rainforest, 
ranging from wet evergreen rainforest 
to gallery forest and other more seasonal 
rainforests, with a few also occurring in wet 
sclerophyll forest. They occur in a variety of 
substrates from sea level to 1600 m. 

In February 1981, Laurence Jessup with 
J. Geoff Tracey and Tony Irvine collected 
the first specimens {Jessup 319, Tracey & 
Irvine and Tracey 14993 ) of the species 
described below as Cryptocarya cercophylla 
W.E. Cooper within the Boonjee Logging 
Area in the Russell River headwaters (now 
part of Wooroonooran National Park). The 
plant was originally thought to be a species 
of Icacinaceae. Flowering specimens were 
subsequently collected by Jessup, Gordon 
Guymer and Bill McDonald in October 1988, 


Bernie Hyland in July 1990, Bob Jago in 
August 1994 and Rigel Jensen in November 
1994. By this time the plant was recognised as 
a species of Cryptocarya. Visiting Nigerian 
researcher, Olusegan Osunkoya with direction 
from Jensen, collected the first specimen with 
one fruit, in February 1994. Hyland et al. 
(2003) listed this entity under the phrase name 
Cryptocarya sp. (Boonjee BH 25794RFK), 
whereas the Queensland Herbarium listed it 
as Cryptocarya sp. (Boonjie L.W. Jessup+ 319) 
(Jessup 2007). However, it was not treated in 
either Hyland (1989) or the Flora of Australia 
(Le Cussan & Hyland 2007). With advice 
from Andrew Ford, a tree was located in 
January 2013 with enough fruiting material to 
enable a complete description of this species 
as Cryptocarya cercophylla W.E.Cooper. 

Materials and methods 

The study is based upon the examination of 
herbarium material from CNS together with 
relevant field observations. All specimens 
cited have been seen by the author. 

Measurements of the floral parts and 
fruits are based on material preserved in 
70% ethanol and fresh specimens in the field. 
Abbreviations in the specimen citations are: 
LA (Logging Area), NP (National Park), 
SFR (State Forest Reserve) and TR (Timber 
Reserve). 


Accepted for publication 16 July 2013 


76 

Taxonomy 

Cryptocarya cercophylla W.E.Cooper sp. 
nov. Similar to Cryptocarya meissneriana 
Frodin but differs in the leaf apex (distinctly 
caudate, with acumen 8-20 mm long and 

1- 2 mm wide, versus shortly acuminate or 
caudate with acumen up to 10 mm long and 

2- 4 mm wide), lateral veins (3-6 versus 7-10), 
tepals (erect and not opening widely versus 
opening widely at anthesis), and anther apex 
(acute versus truncate). Typus: Australia: 
Queensland. Cook District: Timber Reserve 
1230, Bartle Frere, Boonjee Logging Area 
[east of Malanda], 17 November 1994, R. 
Jensen 72 (holo: CNS [2 sheets + spirit], iso: 
BRI, CANB, MO distribuendi). 

Cryptocarya sp. (Boonjee BH 25794RFK); 
Hyland etal. (2003). 

Cryptocarya sp. (Boonjee); Cooper & Cooper 
(2004: 144). 

Cryptocarya sp. (Boonjie L.W.Jessup+ 319); 
Jessup (2007: 92, 2010: 97). 

Cryptocarya sp. Boonjee (B.Hyland 
25794RFK); Hyland et al. (2010). 

Illustrations : Hyland et al. (2003), as 
Cryptocarya sp. (Boonjee BH 25794RFK); 
Cooper & Cooper (2004: 144) as Cryptocarya 
sp. (Boonjee). Hyland et al. (2010), as 
Cryptocarya sp. (Boonjee B.Hyland 
25794RFK). 

Small tree to 10 m, poorly formed and often 
with coppice shoots, stems to 12 cm diameter 
at breast height, not buttressed. Bark pale 
with sparse vertical elongated lenticels and 
fine fissures; twigs terete, green. Leaves 
simple, alternate, petiolate, discolorous, 
glabrous; petioles 7-13 mm long, grooved 
and yellowish; lamina ovate or elliptical, 
53-100 mm long, 22-45 mm wide, thinly 
leathery, upper side glossy dark green, lower 
side shiny (less so than the upper surface), 
paler green, base attenuate or cuneate, apex 
caudate, acumen 8-20 mm long, 1.3-2 mm 
wide; margins entire and slightly recurved, 
mostly sinuous; venation brochidodromous, 
midrib depressed on upper side; secondary 
lateral veins indistinct, 3-6 pairs at 45-90° to 
the primary midrib, oil dots numerous (visible 


Austrobaileya 9(1): 75-79 (2013) 

with a lens). Inflorescences axillary or rarely 
terminal, racemose or rarely paniculate, 
4-7-flowered, 10-28 mm long, not exceeding 
leaves, rachis ± glabrous with occasional 
minute simple hairs; bracts triangular, c. 
0.5 mm long and 0.2 mm wide, sparsely and 
minutely hairy, caducous. Flowers bisexual, 
actinomorphic, not fragrant, 3-merous, c. 
2.5 mm long and 1.75 mm diameter; pedicels 
0.6-2 mm long, sparsely and minutely hairy; 
perianth tube 2.5-2.8 mm long, c. 1.5 mm 
wide at apex, inner surface hairy, outer surface 
glabrous; tepals 6, in 2 whorls of 3, glabrous 
outside, sparsely hairy inside, creamy-green 
or cream, erect at anthesis; outer tepals 1.2- 
1.7 mm long, 1.2-1.5 mm wide; inner tepals c. 
1.7 mm long and 1.6 mm wide; stamens with 
filaments sparsely hairy, c. 5 mm long; gland 
heads conical, glabrous, c. 1 mm long and 0.4 
mm wide, stalks sparsely hairy or glabrous 
c. 0.2 mm long; anthers 6 introrse and 3 
extrorse, 2-locular, inner and outer anthers 
c. 0.5 mm long and 0.5 mm wide, glabrous; 
staminodes 3, head cordate, sessile, c. 0.7 mm 
long; ovary glabrous to sparsely hairy, 0.8-1 
mm long, c. 0.5 mm wide; style glabrous, 
1.2-1.6 mm long. Fruit a nut, appearing as 
an inferior drupe, elliptical, rarely ovoid or 
obovoid, mostly asymmetric at base with 
one side slightly lobed, 16.5-20 mm long, 
10-14.5 mm wide, ribbed, glabrous, red to 
shiny black at maturity, epicarp 0.4-0.6 mm 
thick, mesocarp + epicarp c. 1.8 mm thick, 
mesocarp green, stigma persistent at apex 
with diameter c. 1.5 mm, pedicel 1-2.8 mm 
long; seed solitary, 16-19.5 mm long, 8-12.5 
mm wide; endocarp beaked at base, cream, 
ribbed; cotyledons white, uniform in texture 
(not ruminate), radicle apical. Figs. 1 & 2. 

Additional selected specimens (from 25 examined): 
Queensland. Cook District: Tolga, cultivated, Jan 1995, 
Sankowsky 1437 (CNS); Atherton, cultivated, Jul 1990, 
Gray 5210 (CNS); Butchers Creek, c. 20 km ESE of 
Yungaburra, Stockwellia site, Jan 1985, Morawetz, Waha 
& Weber 114-30185 (CNS); TR 1230 Boonjie LA, Feb 
1981, Jessup 319, Tracey & Irvine (BRI); Wooroonooran 
NP, Stockwellia site, July 2000, Forster PIF25913, 
Booth & Jensen (BRI); TR 1230, Bartle Frere, Boonjee 
LA, July 1990, Hyland 14052 (CNS), 14053 (CNS), 
25793RFK (CNS), 25794RFK (CNS), 25795RFK (CNS); 
TR 1230, Boonjee LA, the headwaters of the Russell 
River, Feb 1981, Tracey 14993 (CNS); Wooroonooran 
NP, Stockwellia area south of road to Mt Bartle Frere, 
Site 216, Dec 2009, Ford 5636 & Cooper (BRI, CNS); 


Cooper, Cryptocarya cercophylla 

Stockwellia site, Boonjee, Wooroonooran NP, Jul 2011, 
Cooper 2136 & Ford (CNS), Oct 2012, Cooper 2165 
(CNS); SFR 755, Bartle Frere, Gosschalk LA, Sep 1994, 
Gray 5774 (CNS); TR 1230, Boonjee LA, Feb 1994, 
Osunkoya 1 (CNS); Wooroonooran NP, along Russell 


77 

River track between lookout and Chucklunga track, Sep 
2012, Ford 6082 (CNS); Wooroonooran NP, Donkey 
Track near Zig Zag, Mar 2003, Forster PIF29259 & 
Cooper (BRI); Wooroonooran NP, along Chucklunga 
track, Jan 2013, Cooper 2210 & Ford (CNS); Palmerston 
Ridge Track, Feb 1995, Hunter 1154 (BRI). 


Fig. 1. Cryptocarya cercophylla. A. stem with leaves showing main lateral veins *0.5. B. leaf showing primary, 
secondary and some tertiary venation xi. C. inflorescence showing open flower, buds and bracts on pedicels x4. D. 
flower (side view) x8. E. flower (side view with 3 tepals removed) *12. F. staminode x32. G. anther (inner whorl) x32. 
H. anther (outer whorl) x32.1. gland showing head and stalk xl. A & B from Forster PIF29259 & Cooper (BRI); C-I 
from Jensen 72 (CNS). Del. W. Smith. 


78 


Austrobaileya 9(1): 75-79 (2013) 


Fig. 2. Cryptocarya cercophylla fruiting stem (i Cooper 2210 & Ford [CNS]). Photo: A. Ford. 


Distribution and habitat : Cryptocarya 
cercophylla is endemic to Wooroonooran 
National Park within the Wet Tropics 
bioregion of north-eastern Queensland, where 
it is known to occur in a small area between 
what is locally referred to as the Stockwellia 
site at Boonjie and the nearby Russell River, 
at altitudes 580-680 m. The species may well 
occur in a broader area of the western Russell 
River catchment (Map 1). 

Cryptocarya cercophylla grows as an 
understory tree in a high rainfall area of 
notophyll rainforest on clay soils derived from 
metasediments. Tree species that it co-occurs 
with are Stockwellia quadrifida D.J.Carr, 
S.G.M.Carr & B.Hyland, Beilschmiedia 
oligandra L.S.Sm., Ceratopetalum virchowii 
F.Muell., Elaeocarpus johnsonii F.Muell 
Endiandra dichrophylla F.Muell, Flindersia 
bourjotiana F.Muell, Franciscodendron 
laurifolium (F.Muell) B.Hyland & Steenis and 
Peripentadenia mearsii (C.T.White) L.S.Sm., 
Bobea myrtoides (F.Muell.) Valeton and 
Aceratium doggrellii C.T.White. It appears to 
favour areas disturbed by landslips. 


Phenology : Flowers have been recorded in 
July, October, November and December: 
fruits have been recorded in January and 
February. 

Notes : In cultivation, Cryptocarya 

cercophylla has flowered as a shrub at 1.5 m 
tall; however, flowering collections in the wild 
are only known from trees 6 m and above. 

Fallen stems produce adventitious 
roots, which appear to eventually grow into 
individual trees following decomposition 
of the fallen stem. Evidence of such a 
phenomenon can be seen as a number of 
‘individuals’ occurring close together (3-5 m) 
and in a straight line. 

Affinities: Cryptocarya cercophylla appears 
to be most similar to C. meissneriana from 
southeast Queensland and northeast New 
South Wales but differs from this species in 
the leaf apex (distinctly caudate, with acumen 
8-20 mm long and 1-2 mm wide, versus 
shortly acuminate or caudate with acumen 
up to 10 mm long and 2-4 mm wide), leaf 
venation (3-6 secondary lateral veins versus 


Cooper, Cryptocarya cercophylla 

7-10 secondary lateral) and the tepals erect 
and not opening widely at anthesis (versus 
tepals opening widely at anthesis). 

Conservation status : Cryptocarya 

cercophylla is a geographically restricted 
species and can be classified as Vulnerable 
(Criterion D2) using the criteria of the IUCN 
( 2001 ). 

Etymology : The specific epithet is derived 
from the Greek, kerkos (tail), and phyllon 
(leaf), in reference to the distinctive caudate 
or tail-like leaf apices. 

Acknowledgements 

Frank Zich and Darren Crayn are thanked 
for support and access to CNS herbarium, 
Andrew Ford and Cath Moran for assistance 
and good company in the field and especially 
Andrew for finding the elusive fruiting tree, 
Peter Bostock and Will Smith for the map and 
illustrations. Laurie Jessup enlightened me 
regarding the discovery of the plant. Darren 
Crayn and Andrew Ford made worthwhile 
suggestions to an earlier manuscript. Permits 
to collect in National Parks and State Forests 
were issued by the Queensland Department of 
Environment and Resource Management. 

References 

Cooper, W. & Cooper, W.T. (2004). Fruits of the 
Australian Tropical Rainforest. Nokomis 
Editions: Melbourne. 

Hyland, B.P.M. (1989). A revision of Lauraceae in 
Australia (excluding Cassytha ). Australian 
Systematic Botany 2: 135-367. 

Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray, 
B. & Elick, R.W. (2003). Australian Tropical 
Rain Forest Plants. Trees, Shrubs and Vines. 
CD-ROM. CSIRO Publishing: Melbourne. 

Hyland, B.P.M., Whiffin, T. & Zich, F. (2010). Australian 
Tropical Rain Forest Plants Edition 6 (online 
version). CSIRO. http://www.anbg.gov.au/cpbr/ 
cd-keys/rfk/index.html Accessed October 2011. 

Iucn (2001). IUCN Red List of Categories and 
Criteria: Version 3.1. IUCN Species Survival 
Commission: Gland (Switzerland)/Cambridge 
(United Kingdom). 


79 

Jessup, L.W. (2007). Lauraceae. In P.D. Bostock & A.E. 
Holland (eds.). Census of the Queensland Flora 
2007, p. 54. Environmental Protection Agency, 
Brisbane. 

- (2010). Lauraceae. In P.D. Bostock & A.E. 

Holland (eds.). Census of the Queensland Flora 
2010, p. 92. Environmental Protection Agency, 
Brisbane. http://www.derm. qld.gov. au/wildlife- 
ecosystems/plants/pdf/qld-flora-census.pdf 
Accessed October 2011. 

Le Cuss an, J. & Hyland, B.P.M. (2007). Lauraceae. In 
A.S. George (ed.). Flora of Australia. 2: 106- 
223. Australian Biological Resources Study: 
Canberra. 


Map 1. Distribution of Cryptocarya cercophylla in 
north-east Queensland. Shaded area on map indicates 
Wooroonooran National Park. 


Violaperreniformis (L.G.Adams) R.J.Little & 

G.Leiper, stat. nov with notes on Australian 
species in Viola section Erpetion (Violaceae) 

R. John Little 1 & Glenn Leiper 2 

Summary 

Little, J. & Leiper, G. (2013). Viola perreniformis (L.G.Adams) R.J.Little & G.Leiper, stat. nov., 
with notes on Australian species in Viola section Erpetion (Violaceae). Austrobaileya 9(1): 80-101. 
Viola perreniformis (L.G.Adams) R.J.Little & G.Leiper is recognized as a distinct species in section 
Erpetion in Australia. A description of the species is presented. Leaf and floral characters of four 
closely related species in section Erpetion, V. banksii Thiele & Prober, V. hederacea Labill. subsp. 
hederacea, V. perreniformis stat. nov., and V. silicestris Thiele & Prober, are compared and discussed. 
Various aspects of the biology of these species are discussed and provisional keys are presented based 
on floral and leaf characters. Comments are also provided for four other species in section Erpetion. 
V. cleistogamoides (L.G.Adams) Seppelt, V eminens Thiele & Prober, V. fnscoviolacea (L.G.Adams) 
T.A. James, and V. sieberiana Sprengel, and a putative hybrid, V x zophodes Thiele & Prober. 

Key Words: Violaceae, Viola , Viola section Erpetion, Viola banksii, Viola betonicifolia, Viola 
caleyana, Viola cleistogamoides, Viola eminens, Viola fuscoviolacea, Viola hederacea subsp. 
hederacea, Viola perreniformis, Viola sieberiana, Viola silicestris, Viola x zophodes, anther gland, 
cleistogamy, nectary 

'R. John Little, MagnaFlora, 16 Pebble River Circle, Sacramento, California 95831, U S A. Email: 
bugsandbunnies4u@sbcglobal. net 

2 Glenn Leiper, 30 Tweedvale Street, Beenleigh, Queensland 4207, Australia. Email: leiper30@ 
bigpond.com 


Introduction 

Viola hederacea Labill. subsp. perreniformis 
L.G.Adams was one of eight subspecies 
recognized by Adams in the V hederacea 
complex in Australia (Adams 1982a,b). This 
was the first attempt to classify the diversity of 
forms that had been assigned to V hederacea 
sensu lato in section Erpetion (Sweet) Benth. 
& Hook.fi Later, James (1990) recognized 
seven forms (A through G) of V hederacea 
in New South Wales based on leaf shape and 
pubescence, and in addition, elevated three 
of Adams’ subspecies to species rank. Thiele 
& Prober (2003) summarized the taxonomic 
status of each subspecies of V hederacea 
described by Adams with all subspecies 
except subsp. perreniformis and subsp. 
curtisae L.G.Adams having been elevated to 
species rank. 


In the original description, Viola hederacea 
subsp. perreniformis was distinguished from 
V hederacea subsp. hederacea by the erect, 
soft, short pubescence, by the corolla being 
more or less concolorous, pale (lazuline) blue- 
violet, with the anterior petal white at the base 
with violet-coloured veins, and distinguished 
from other V. hederacea subspecies by 
suborbicular leaves with a deeply cordate base 
(Adams 1982a). The epithet ‘perreniformis’ 
means ‘very reniform.’ Interestingly, the 
adjective ‘reniform’ with respect to leaves of 
subsp. perreniformis was not mentioned in 
the Latin diagnosis and appears only in the 
description of subspecies (Adams 1982b). 
The leaves of subsp. hederacea were stated 
by Adams (1982b) to be variable but never 
suborbicular (authors’ emphasis). Adams 
reported that subsp. perreniformis occurred in 
coastal regions between Cairns, Queensland 
(Qld) and Sydney, New South Wales (NSW). 


Accepted for publication 26 September 2013 


81 


Little & Lei per, Viola perreniformis 

Some Australian botanists have considered 
Viola hederacea subsp. perreniformis as 
warranting specific status (e.g. Thiele & Prober 
2003), but the nomenclatural change has not 
been made. Based on a study of the holotype 
and isotype as well as other collections of 
the V hederacea complex at BRI, CANB, 
and NSW, and recent field observations by 
the authors, we believe subsp. perreniformis 
differs sufficiently from subsp. hederacea 
to warrant recognition at species rank, as V 
perreniformis. The close relationship among 
the four taxa, V banksii, V hederacea subsp. 
hederacea, V perreniformis and V silicestris, 
can be seen in the morphology of their leaves 
and flowers. 

Long regarded as a difficult taxon because 
of its many leaf forms, recent studies of 
the Viola hederacea complex in Australia 
in section Erpetion have begun to clarify 
relationships among taxa in this group. In 
2003, Thiele & Prober presented the most 
comprehensive study and summary to date of 
species in section Erpetion (Thiele & Prober 
2003), which included a circumscription 
of species in the V hederacea complex, 
descriptions of several new species including 
V banksii , and a key to most members of 
the complex. Their circumscription of V 
hederacea did not include V hederacea 
subsp. perreniformis , which they believed 
to be a taxon distinct from V hederacea. 
Thiele & Prober (2006) described V 
silicestris as a new species and discussed 
similarities and differences of it compared 
to four other species in section Erpetion. V. 
cleistogamoides (L.G.Adams) Seppelt, V 
fuscoviolacea (L.G.Adams) T.A.James, V. 
hederacea and V sieberiana Sprengel. 

During our study of Viola perreniformis, 
we found that V banksii, V. hederacea subsp. 
hederacea , and V silicestris were the taxa most 
often confused with Vperreniformis stat. nov. 
In this paper we examine and compare these 
four species (collectively referred to herein as 
the “four taxa”) with new and existing data, 
document previously unreported observations 
of various morphological features, and 
suggest topics for future study. The taxonomy 
of section Erpetion is challenging and far 


from complete. The four taxa and other 
species in the section appear to have many 
unique features not known to occur in other 
Viola species. 

Materials and methods 

In November and December 2011 and January 

2012, the first author examined herbarium 
specimens in the Viola hederacea complex 
at the herbaria BRI and NSW, including 
specimens identified as subsp. perreniformis. 
Both authors conducted numerous field studies 
in Qld and NSW during this timeframe. In 

2013, the first author continued study of 
species in the V. hederacea complex at the 
herbaria CANB and NSW, and both authors 
examined specimens at BRI identified as 
subsp. perreniformis and subsp. hederacea. 
Scanned images were examined of slides of 
the type of V hederacea at CANB. 

At BRI, all specimens previously 
determined as subsp. perreniformis were 
compared with the isotype housed at BRI and 
an image of the holotype housed at CANB. 
Specimens were then sorted into three 
groups: 1) specimens that matched or closely 
resembled the holotype and isotype (these 
specimens are listed below under Additional 
specimens examined)', 2) specimens generally 
similar to the holotype and isotype, but with 
leaves too immature or in too poor a condition 
to make a definitive determination; and 3) 
specimens which, in the authors’ opinion, 
did not fit the concept of the perreniformis 
taxon, most of which are referable to subsp. 
hederacea. Over 1000 high-resolution, 
digital photographs were taken of herbarium 
specimens to facilitate further study. 

Field observations of Viola perreniformis 
were made by both authors in February 2013 
at Nightcap National Park (NP) in NSW. The 
second author made additional observations 
and collections of this taxon in March, April, 
and May 2013 at Crows Nest NP and in 
March 2013 at Moogerah Peaks NP, both in 
Qld. In June 2013, the second author collected 
fresh flowers of V banksii from Springbrook 
NP (possibly a horticultural escape) and 
Venman Bushland NP and V hederacea from 
Springbrook NP, all sites in Qld; the anther 


82 

glands on these flowers were then observed 
with a stereozoom microscope. Using a 
stereozoom microscope with a camera 
attachment, the first author obtained micro¬ 
photographs of flowers and anther glands of 

V banksii and seeds of V perreniformis and 

V betonicifolia. 

The species description is based on a 
composite of observations and measurements 
of leaves, petioles, peduncles, and flowers 
from a) the holotype and isotype; b) from 
herbarium specimens at BRI, and c) from 
plants collected by the authors in 2012 and 
2013 and photographs taken of fresh material. 
The number of ‘crenations’ (rounded lobes) 
on leaf margins were recorded only from 
one side of a leaf because on herbarium 
specimens, overlapping leaves often prevented 
observation of the entire margin. 

As used herein in reference to pubescence, 
the term pilose means soft, thin, straight, and 
clearly separated hairs; the type of hairs on 
leaf surfaces of Viola perreniformis. Villous 
means ± densely covered with soft, thin, long, 
straight or sometimes shaggy hairs, not clearly 
separated; the type of hairs often found on 
petioles and peduncles of V perreniformis. 

Common abbreviations used in specimen 
citation include CP (Conservation Park), NP 
(National Park) and SF (State Forest). 

Taxonomy 

Viola perreniformis (L.G.Adams) R.J.Little 
& G.Leiper, stat. nov.; Viola hederacea 
Labill. subsp. perreniformis L.G.Adams, FI. 
Aust. 8: 386 (1982). Type: Queensland. Port 
Curtis District: c. 40 km WSW of Gladstone, 
Many Peaks Range, 2 June 1977, M.D. Crisp 
2751 & R.K. Ellyard (holo: CANB, image!; 
iso: BRI). 

Plants perennial, often appearing acaulescent, 
3-6.5 cm high, stoloniferous, rooted sections 
of the stolon forming rosettes of leaves on short, 
fleshy rhizomes. Stems sometimes present, 
1-8 cm long. Leaves: stipules narrow-ovate 
or lanceolate, margin irregularly fimbriate, 
apex acute, long-aristate; petiole 9-23 cm 
long, sparsely or densely pilose or villous, 
occasionally glabrous; blade reniform, sub- 


Austrobaileya 9(1): 80-101 (2013) 

reniform, or ovate (outline ovate, orbicular, 
suborbicular, or semicircular), 5.5-23 mm 
long, 7-27 mm wide, base cordate or deeply 
cordate, occasionally subtruncate or truncate, 
often with distinct sinus between lobes; 
margins ciliate or eciliate, uniformly or 
irregularly crenate from basal lobes to apex, 
4-9(-10) lobes per side, sometimes poorly 
defined, occasionally some lobes ± serrate, 
apex acute, obtuse, or rounded, occasionally 
± truncate, the midvein terminating in a 
blunt mucro; abaxial surfaces of mature and 
immature leaves sparsely or densely pilose, 
with hairs scattered on 1° and 2° veins or over 
entire surface, occasionally glabrous, small 
or immature leaves sometimes tomentose; 
adaxial surfaces of mature and immature 
leaves sparsely or densely pilose, with hairs 
scattered over entire surface, occasionally 
glabrous; blades bi-colored, darker green 
adaxially, lighter abaxially. Flowers solitary, 
peduncles arising from a fleshy, usually 
vertical rhizome, extending above the leaves, 
25-77 mm long, sparsely or densely pilose 
or villous, occasionally glabrous; bracteoles 
narrowly linear or aciculate, 2.5-5 mm 
long, opposite or alternate, often above the 
middle during capsule development; sepals 
lanceolate, margin entire, scarious, eciliate, 
apex acute, usually gland-tipped, auricles not 
prominent, c. 0.5 mm long; upper petals erect 
in young and reflexed in mature flowers, pale 
blue-violet on both surfaces, white basally, 
occasionally with a faint or prominent violet- 
coloured blotch distal to the proximal white 
area; lateral petals spreading in new and 
twisted in mature flowers, pale blue-violet 
on both surfaces, occasionally appearing 
white distally, white basally with a green 
and white, ± 1-2 mm long, ± 1-2 mm wide 
indurate area at the base, often with a faint or 
prominent violet-coloured blotch distal to the 
proximal white area, with faint or prominent 
violet-coloured veins, usually with a dense 
patch of white clavate and cylindrical beards 
on proximal third of petal, often with some 
tips violet-coloured, distal beards sometimes 
entirely violet-coloured; lowest petal obovate 
to oblong, usually white on proximal half 
with a prominent green and white, 1-2 mm 
long, 1-2 mm wide indurate area at the base, 


83 


Little & Lei per, Viola perreniformis 

the distal half of petal pale blue-violet, the 
veins violet-coloured, prominent on adaxial 
surface, faint or prominent on abaxial 
surface, usually not extending to distal 
margin of petal, the midvein and the two 
adjacent veins distinct, prominent, parallel, 
± same length, scarcely or not anastomosing 
(without evident, interconnecting 2° veins), 
5-9 mm long, apex retuse, truncate, rounded, 
occasionally with a short, sharp or rounded 
point between two lobes, spur not present; 
style geniculate near attachment to ovary, ± 
terete, style head glabrous, not differentiated 
from style shaft; anthers cream-coloured, 
anther glands pale green, smooth, c. 14-2/3 as 
long as anther, prominent, margins of terminal 
anther appendages not hairy; pollen cream- 
coloured. Capsules 5-6 mm long, 3-valved, 
often purple-spotted, glabrous. Seeds ovoid, 
surface uneven to irregularly rugose, 1—1.2 
mm long, shiny or dull dark brown mottled 
with patches of light brown or tan, elaiosome 
not present. 

Additional selected specimens examined : Queensland. 
Cook District: Intersection of road to radio tower and 
Longlands Gap Road, Longlands Gap SF, Aug 2007, Zich 
617 & Quinn (BRI, CANB). North Kennedy District: 
Mt Fox, Dec 1949, Clemens s.n. (BRI [AQ115414]). 
South Kennedy District: Dalrymple Heights & vicinity, 
Jul 1947, Clemens s.n. (BRI [AQ115415]). Leichhardt 
District: Carnarvon Creek Gorge, 70 miles [112 
km] NW of Injune, May 1962, Johnson 2411 (BRI); 
Blackdown Tableland, c. 1 km E of main N-S road 
crossing, Jan 1973, Henderson H1392 & Sharpe (BRI). 
Port Curtis District: c. 60 km SW of Gladstone, 8 km 
SE of Amys Peak, Jun 1977, Crisp 2793 (BRI); Kroombit 
SF 316, Aug 1984, Gibson 634 (BRI); Castletower Mt, 
Oct 1988, Randall 591 (BRI); Portion 10 V, Oaky Creek, 
Castletower, Apr 1989, Gibson TOI482 (BRI). Wide Bay 
District: Mt Woowonga, Woowonga Range SF, Jan 
1982, Forster PIF1153 (BRI); Tinana Creek at Missings 
Crossing, May 2001, Stephens 138903A(11) (BRI); Mt 
Boulder, SE of Gympie, Oct 1993, Bean 6831 (BRI); 
Mapleton Logging Area, Mapleton SF, W of Mapleton, 
Mar 1998, Bean 13121 (BRI, CANB, HO, MEL, NSW). 
Moreton District: Mt Mee SF 893,2 km SE of Forestry 
office, Sep 1997, Halford Q3357 & Boyle (BRI); Alice 
Creek, 7.5 km ESE of Murphys Creek Rail Siding, Aug 
1990, Forster PIF7100 & Bird (BRI); Mt Edwards NP, 14 
km W of Boonah, Sep 1992, Forster P1F11483 & Reilly 
(BRI); Mt Edwards, Moogerah Peaks NP, c. 14 km W of 
Boonah, Feb 2010, Halford Q9754 (BRI); Mt May, Sep 
1992, Forster PIF11794 et al. (BRI); Southern slopes of 
Mt Greville, SW of Lake Moogerah, Jan 2007, Phillips 
1679 & Phillips (BRI); Between N and S peaks of Mt 
Maroon, W of Rathdowney, Apr 2009, Phillips 2006 & 
Phillips (BRI); Cronins Creek between Mt Barney & Mt 


Ernest, Dec 1932, White 8757 (BRI); Near summit of 
Mt. Lindsay [Lindesay], Sep 1973, Dowling 109 (BRI). 
New South Wales. Terania Creek, 10 km NNE of The 
Shannon [Channon], Feb 1980, Coveny 10628 (BRI); 
Summit of Mt Carnham, W of Grafton, in the Baryugil 
area, Jun 1993, Bryant s.n. (BRI [AQ566756]); Summit 
of Munningyundo Mt, Nymboida NP, between Grafton 
and Glen Innes, Mar 2003, Bean 20105 (BRI); North 
Rocks Road, Nightcap Range, Little 13697 & Leiper 
(BRI). 

Distribution and habitat : The species 
occurs in a series of disjunct localities from 
west of Cairns, north Qld, to Sydney in 
NSW. Recorded elevations of specimens 
examined in this study range from 75 m at 
Oaky Creek, Castletower to 1160 m at Mt 
Lindesay, both in Qld. Of the 14 specimens 
that provide elevation data nearly 60% were 
collected at 500 m or higher. Microhabitat 
situations reported by collectors include: 
moist situations in wet sclerophyll forests, in 
woodlands, in open eucalypt forests, on steep 
slopes, in crevices of rocky cliffs, cascading 
over rock faces with or without water flowing, 
on rhyolite or on rocky brown loam, granite, 
or sandstone; east, south, and west-facing 
slopes (Fig. 1). 

Phenology : Flowering and fruiting occurs 
throughout the year. 

Conservation status : Viola perreniformis is 
widespread, occurs in several National Parks, 
and is not known to be at risk. 

Notes on Viola section Erpetion 

Observations on aspects of morphological 
variation and reproductive biology focus 
primarily on the four taxa Viola banksii, V 
hederacea subsp. hederacea , V. perreniformis 
and V silicestris. Information is also 
presented for four other species in the section, 
V. cleistogamoides, V. eminens Thiele & 
Prober, V fuscoviolacea, and V sieberiana 
and a putative hybrid, V. x zophodes Thiele 
& Prober. Other taxa recognized in section 
Erpetion that are not discussed include V 
hederacea subsp. curtisiae and V improcera 
L.G.Adams. 

Stolons and Rhizomes. All species in section 
Erpetion produce stolons and two small 
bracts are always present between stolon 
nodes. Thiele & Prober (2003) reported that 


84 


Fig. 1. V. perreniformis. Habitat, Crows Nest NP, Qld, 
Mar 2013. Photo: G. Leiper. 


rootstocks of Viola banksii and V hederacea 
were sometimes somewhat swollen and 
bulbous at the stem bases, but they were not 
or scarcely swollen in V silicestris (Thiele 
& Prober 2006). The authors made similar 
observations for these species. 

Stolons in section Erpetion are initially 
produced on the surface. Overtime depending 
on their location, they can become buried 
with eroded soil, leaf litter, etc. Numerous 
herbarium specimens and the author’s recent 
collections show that buried, subterranean 
stolons can eventually become lignified 
in some species. This phenomenon is not 
known to have been previously reported for 
Australian species of Viola in the section. 
Although a buried stolon is technically a 
‘rhizome,’ buried stolons of these species do 
not become thick and fleshy, but retain their 
narrow shape. 

A stolon node situated in a suitable site has 
the capacity to produce a vertically oriented 
rhizome and a rosette of leaves. As a rhizome 
matures it elongates vertically, its diameter 
increases, and it becomes fleshy, similar to 
many other Viola species. New leaves are 
produced distally as older leaves die. Leaf 
abscission scars and the remains of stipules 
are visible on the surface of the rhizome. If 
a vertically oriented rhizome is not buried, 
it can attain the appearance of a miniature 
tree; a large clone can attain the appearance 
of a miniature forest. A similar situation is 
sometimes observed in other Viola species 
e.g. V odorata L. and V riviniana Rchb. If a 


Austrobaileya 9(1): 80-101 (2013) 

vertically oriented rhizome is slowly buried 
with eroded soil, it can survive by elongating 
vertically, a useful strategy in a geologically 
unstable environment. All that may be visible 
at the surface is a rosette of leaves. 

Although empirical data are lacking, it 
is possible that fleshy rhizomes function as 
water storage organs. Enlarged, subterranean 
rhizomes were found by the second author in 
a population of Viola perreniformis plants at 
Crows Nest NP, Qld (Fig. 2). Its gourd-like 
shape suggests it may function as a water 
and/or nutrient storage organ. Perhaps this 
rhizome was initially above ground and then 
subsequently buried; perhaps it was then able 


Fig. 2. V perreniformis. An enlarged, subterranean 
rhizome; length between arrows is c. 22 mm. Crows Nest 
NP, Qld, Mar 2013. Photo: G. Leiper. 


to retain water more efficiently and gradually 
assumed a gourd-like shape. 

Clones of some species can occupy up to 
1 m or more in diameter. Although detailed 
studies of how these species spread via stolons 
have not been made, the first author recorded 
preliminary observations of the stages that a 
single ‘mother’ plant of Viola hederacea went 
through as it matured and spread over time; a 
brief description follows. After several leaves 
in the rosette of the mother plant matured and 
enlarged, five stolons developed that spread 
out in all directions. A rosette of miniature 
leaves formed at the distal end of each 
elongating stolon. If the stolon tip contacted 
suitably moist soil, a white, root-like structure 
developed and grew into the soil. After 
several days, the leaves in the newly rooted 
rosette increased in size and simultaneously 


85 


Little & Lei per, Viola perreniformis 

several new stolons formed and spread out in 
all directions and as they elongated, rosettes 
of miniature leaves formed at the tip of each 
new stolon. This process was repeated over 
and over in all directions from the initial 
mother plant until there was a mass of stolons 
and leaves c. 1 m in diameter. If a node didn’t 
contact moist soil, the stolon continued to 
elongate, developing rosettes of leaves at 
each node. Because the available evidence 
suggests seeds of the four taxa may not be 
ant dispersed (see below) and they may not 
produce cleistogamous flowers (see below), 
vegetative reproduction through vigorous 
stolon growth may be an important strategy 
for survival in section Erpetion. 

Stems. A valuable taxonomic character 
commonly used to separate Viola species in 
European, North American, and Asian floras, 
is whether the plant has stems (caulescent) or 
does not (acaulescent). Stems in caulescent 
Viola species are usually upright structures 
that originate from a fleshy or somewhat 
woody rhizome and possess alternating 
leaves subtended by stipules. Peduncles that 
produce chasmogamous and in many species, 
cleistogamous flowers, arise from axillary 
positions on the stem. In caulescent species 
in North America, the internode distance is 
usually short early in the growing season, 
elongating later in the season. Peduncles in 
caulescent species are normally produced 
only from stems and not directly from a 
rhizome. Caulescent Viola species in North 
America have both basal and cauline leaves, 
although a few species have only cauline 
leaves and no basal leaves. When present, 
basal leaves arise directly from the rhizome 
and are taxonomically useful in some species. 
In all acaulescent species known outside of 
Australia, the leaves and peduncles arise 
directly from a rhizome. 

Of interest relative to the foregoing 
is that all Australian species in section 
Erpetion produce stolons. By analogy with 
the acaulescent, stoloniferous species in 
Europe and North America, species in 
section Erpetion would be assumed to be 
acaulescent. However, the Viola treatments 
in most Australian floras e.g. Flora of 


Australia (Adams 1982b), Flora of South-East 
Queensland (Stanley & Ross 1983), Flora 
of New South Wales (James 1990), Flora of 
Victoria (Entwisle 1996), Electronic Flora 
of South Australia (2007), Flora of Tasmania 
Online (Duretto 2009), and New South Wales 
Flora Online (James 2013), state that stems are 
present for all species in the section. “Stems 
short” and “stems short, erect” are common 
descriptions in these floras for stems of species 
in section Erpetion. Also of interest is that the 
peduncles of all Australian species in section 
Erpetion , with two known exceptions, appear 
to develop only from rhizomes produced 
at a stolon node, similar to acaulescent, 
stoloniferous species elsewhere in the world. 
Thus, at present it appears that stems do not 
normally occur in section Erpetion , such as 
those typically found in caulescent species, 
e.g. V caleyana Don in Australia. 

A remarkable exception was documented 
by Thiele & Prober (2006) when they 
reported that Viola silicestris often develops 
“a caulescent growth habit when growing in 
sheltered sites amongst other vegetation, with 
elongate, weakly erect, scrambling aerial 
stems to 30 cm high and scattered leaves. Plants 
in exposed sites have contracted stems and 
fasciculate leaves”. In northern NSW and Qld, 
we found stems on V silicestris plants that had 
developed at stolon nodes and peduncles that 
developed from axillary nodes on the upright 
stems. Peduncles may also develop directly 
from rhizomes at stolon nodes, but this has 
not been verified. Stems can be short, e.g. 1.5 
cm, or greatly elongated, 30-40 cm (Thiele & 
Prober 2006). The authors found a population 
in Springbrook NP, Qld, where clearly 
discernable stems occurred at 6 of 8 nodes 
on a stolon 75 cm long. At another location 
in Springbrook NP an 85 cm long stolon had 
discernable stems at 3 of 10 nodes. It may be 
that stem development in this species is more 
common, perhaps overlooked when stems are 
short and hidden in a rosette of leaves. The 
development of a stem from a stolon node 
has not previously been documented in other 
Viola species known to the authors and may 
be unique to section Erpetion. Acaulescent, 
stoloniferous species such as V odorata 
typically root at a node and form a rosette 


86 


Austrobaileya 9(1): 80-101 (2013) 


of leaves similar to the four taxa, but erect 
stems are never produced by such species at a 
stolon node. The growth habit of V silicestris 
may represent a transitional stage from an 
acaulescent to a caulescent habit. 

Viola perreniformis also produces stems. 
In 2013 the authors found plants with stems 
in Nightcap NP in NSW (Little 13697 & 
Leiper [BRI]) (Fig. 3). In addition, possible 
evidence of stems have been observed on 
several herbarium specimens at BRI that were 
collected in Qld and identified by the authors 
as V perreniformis. Stems were not observed 
in populations of V perreniformis at Crows 
Nest NP or on Mt Edwards (Moogerah Peaks 
NP) in Qld. Although phenotypically similar, 
it is possible these populations are genetically 
different and never produce stems, or stems 
were produced but overlooked, or perhaps 
microhabitat conditions precluded stem 
development. 

Thiele & Prober (2006) noted that Viola 
hederacea and V sieberiana, in contrast 
with V silicestris , never (author’s emphasis) 
develop a caulescent growth habit, even when 
growing amongst dense vegetation. Our 
observations of herbarium specimens and of 
living populations of V banksii in Qld have 
not revealed evidence of stems. In V banksii , 
V hederacea and V sieberiana , leaves and 
peduncles appear to arise only from a rhizome 
and never from a caulescent stem. These 
observations suggest these taxa could be 
acaulescent, although additional observations 
are needed. No data are available for V 
cleistogamoides or V fuscoviolacea regarding 
presence/absence of stems. 

Leaves. The size and shape of leaves of 
most Viola species can vary greatly due to 
many factors, such as time of year (spring 
versus summer), the amount of soil moisture 
and other edaphic factors, and whether in 
sun or shade (Little & McKinney in press). 
Thiele & Prober (2006) appear to have been 
the first to document similar observations in 
reference to Australian violets, stating “This 
is a problem throughout the section, with 
reduced leaves of all species converging in 
shape [when immature] while fully developed 
leaves from sheltered sites are usually highly 


Fig. 3. V perreniformis. Pressed specimen showing 
stem (arrow) arising from a stolon node; 3 peduncles 
from the stem. Nightcap NP, NSW. Feb 2013. Photo: J. 
Little. 

distinctive”. They noted that most herbarium 
specimens fail to adequately represent the 
local variation present in leaves of species in 
section Erpetion and that the location of the 
plant, e.g. sun or shade, can have a profound 
effect on leaf size. We concur and emphasize 
that mature, fully developed leaves can 
facilitate identification of the four taxa. 
However, plants may have mature leaves that 
are not fully expressed, i.e. not typical, if they 
are growing in difficult sites (pers. comm., K. 
Thiele and observations by the authors). Thus, 
having fully developed, mature leaves may 
be essential for identification. In addition, 
it is necessary to consider all the leaves in a 
population and not focus on a few variants 
because in most populations of the four taxa, 
a few leaves can often be found at different 
stages of growth that are more typical of 
one of the other species. Also important to 
note is that various combinations of the four 
taxa are sympatric and it is possible that the 
stolons of two or more species could become 
intertwined in a given population resulting in 
some confusion if the leaves of one species 


Little & Lei per, Viola perreniformis 

are associated with the flowers of another. 

The leaves of all four taxa are darker green 
adaxially compared to lighter coloured abaxial 
surfaces (Fig. 4). The overall outline of Viola 
banksii and V perreniformis leaves remain ± 
proportionally constant during development 
from immature to fully developed, whereas 
the leaves of V hederacea and V silicestris 
expand laterally as the blade matures, with 
the blade becoming much wider than long. 

The information in Table 1 is an attempt 
to qualitatively summarize and contrast 
morphological leaf characters in the four 
taxa typically found in fully developed 
leaves. Some characters for a given species 


87 

appear genetically fixed and do not vary, 
e.g. fully developed Viola silicestris leaves 
never have an orbicular outline; the outline 
of V banksii and V. perreniformis leaves are 
never ± rectangular and their shapes are never 
falcate; a distinct sinus never occurs in fully 
developed V hederacea or V silicestris leaves, 
etc. As noted in the table, other morphological 
characters are only ‘sometimes’ present 
in a given leaf or population. ‘Usually’ 
in the table indicates the norm for the 
species, recognizing that variation occurs 
in that character. Immature leaves of some 
species have characters associated with 
fully developed leaves of other species (see 
discussions below). 


Table 1. Qualitative summary of some leaf characters typically found in fully developed 
leaves of some Viola species 


Leaf characters 

V banksii 

V perreniformis 

V hederacea subsp. 
hederacea 

V silicestris 

outline orbicular 

usually 

sometimes 

no 

no 

outline ovate 

no 

sometimes 

no 

no 

outline ± rectangular 

no 

no 

sometimes 

sometimes 

outline semi-circular 

no 

sometimes 

usually 

no 

shape distinctly 
reniform 

sometimes 

usually 

no 

no 

shape sub-reniform 
(i.e. cordate lobes not 
well developed) 

no 

sometimes 

sometimes 

sometimes 

shape falcate 

no 

no 

no 

sometimes 

leaf margins crenate 
from base near netiole 

to apex 

no 

usually 

no 

no 1 

leaf base cordate 

yes 

usually 

no 

no 

leaf base truncate 

no 

sometimes 

usually 

usually 

distinct basal lobes 
present 

yes 

usually 

no 

no 

distinct sinus present 
formed by lobes 

yes 

usually 

no 

no 

leaves much wider 
than long 

no 

no 

yes 

yes 


The distal leaf margins of V silicestris are occasionally ± crenate, but the basal part of the blade is never 
crenate. 


Austrobaileya 9(1): 80-101 (2013) 


Viola banksii: Mature leaves ultimately 
attain an ± orbicular outline and shape or 
sometimes a reniform shape. The apex of 
mature leaves is usually truncate or rounded; 
a distinct mucro at the apex of the midvein is 
sometimes present; the base is cordate, never 
truncate. The two distinct basal lobes often 
overlap in fresh material, but overlapping 
lobes can be obscured when pressed. As the 
leaf matures and expands, the marginal teeth 
become further apart and often become small 
and less obvious on the upper half of the 
leaf. The margin is occasionally ± dentate. 
Occasionally small teeth and/or 3 or 4 small 
lobes (crenations) are present on the lower half 
of the leaf, but never around the entire margin. 
Mature, orbicular-shaped leaves are about 
as wide as long; reniform-shaped leaves are 
slightly wider than long. In small, immature 
leaves a distinct sinus is often present. A 
distinguishing characteristic of this species is 
the tendency for the apex of fully developed 
leaves to be truncate or somewhat rounded 
while also having cordate basal lobes and a 
distinct sinus (Fig. 4). 

Viola hederacea subsp. hederacea: Mature 
leaves have been described as broad-reniform 
or semi-circular, usually truncate at the 
base and a margin with 8—16 teeth, the base 
occasionally with a broad sinus or broadly 
cuneate (Thiele & Prober 2003). Images of 


Fig. 4. V. banksii. Sequence of development from 
new (1) to fully developed leaf (8). The smallest leaves 
often have a distinct sinus. Adaxial surfaces are shown; 
abaxial in 7. Pilose pubescence present on leaves and 
petioles. Photo: J. Little. 

the type specimen show many small and large 
leaves with a distinct cuneate base. We found 
blade outlines of fully developed leaves to be ± 
rectangular (Fig. 5.8). Leaf apices are variable, 
rounded, obtuse, truncate, or occasionally 
convex. When the apex is truncate or convex, 
the outline is arguably not semi-circular, but 
closer to rectangular. The terminal midvein is 
usually just a small point or mucro. The distal 
(upper) leaf margin is ± entire with only a few 
short points where the major veins terminate. 
The base of fully developed leaves is usually 


Fig. 5. V hederacea. Sequence of development from new/immature (1) to fully developed leaf (8). Basal angle increases 
with age until becoming truncate in fully developed leaves. Adaxial surfaces shown. Photo: J. Little. 


89 


Little & Lei per, Viola perreniformis 

truncate and does not form distinct lobes as in 
V banksii or V perreniformis. Occasionally, 
the basal blade margin is curved or arched 
downward between the petiole attachment 
and the outermost edge of the blade, similar 
to V silicestris. Such leaves are sometimes 
described as having a ‘wide’ sinus. The basal 
margin of young and fully developed leaves 
is entire or may have a small point or two 
and often forms a ± straight line between the 
petiole attachment and the outer edge of the 
blade. The angle formed by the leaf base in 
fully developed leaves is obtuse to usually 
truncate (180°). 

Distinguishing features in fully developed 
leaves separating Viola hederacea (Fig. 
5.8) from V banksii and V perreniformis 
are a truncate leaf base and lack of a sinus. 
Fully developed leaves of V hederacea are 
somewhat similar to V silicestris in that both 
can be ± rectangular and are much wider than 
long. Immature V hederacea leaves usually 
have small lobes, but as the leaf matures 
and enlarges, the angle between the lobes 
gradually increases and the lobes ‘disappear’ 
as the base becomes truncate. Occasionally, 
small, immature leaves of V hederacea have 
very distinct lobes and a distinct sinus and 
are very similar to immature leaves of other 
species, e.g. compare Figs. 4.5, 5.3, 6.1. 

Our observations of herbarium and living 
specimens of Viola hederacea suggest that 
leaves that match the description of “broad- 
reniform” are not fully developed or perhaps 
were growing in difficult sites. The lobes 
in broad-reniform leaves are small (=sub- 
reniform) and the width of such blades are 
smaller compared to a fully developed leaf. 
The typical leaf shape of V eminens was 
described as “broad-reniform” (Thiele & 
Prober 2003); however, we have not studied 
this species and our observations that broad- 
reniform leaves are not fully developed pertain 
only to V hederacea. Although V hederacea 
subsp. hederacea was reported to be replaced 
in Qld by V. hederacea subsp. perreniformis 
(Thiele & Prober 2003), many populations 
have been found in Qld that morphologically 
match the description of V hederacea and are 
considered by the authors and others to be 


V hederacea. The second author has found 
several populations in Qld where the flowers 
are somewhat similar to V hederacea, but the 
leaves are cordate or reniform with distinct 
basal lobes. 

Viola perreniformis : Outlines of fully 
developed leaves are ovate, orbicular, 
suborbicular, or semicircular; their shapes 
are reniform or ovate. The apex is acute, 
obtuse, or rounded, occasionally ± truncate. 
The base of mature and immature leaves is 
usually cordate with a distinct sinus formed 
by the lobes of the blade. The sinus is usually 
distinct, but wider than in V banksii. Most 
populations of V perreniformis in Qld 
occasionally have some truncate and/or 
subtruncate leaf bases in addition to typical 
leaves with cordate bases, but these are 
usually associated with immature leaves. 
The marginal crenations (lobes) are usually 
quite uniform, but can be somewhat ‘ragged’ 
and irregularly shaped in some populations. 
Although the number of crenations per 
blade can vary in the same clone and among 
different populations, distinct crenations are 
present on most leaves in a clone. The number 
of crenations on one side varies from four in 
small/immature leaves to nine (seldom ten) 
in the largest/fully developed leaves. Fully 
developed leaves are usually slightly wider 
than long. Distinguishing features are leaves 
with crenate margins, usually around the 
entire blade in fully developed and usually 
also in immature leaves (Fig. 6). 


Fig. 6. V perreniformis. Sequence of development from 
immature (1) to fully developed leaf (5). Leaves at all 
growth stages are lobed. Adaxial surfaces shown. Photo: 
G. Leiper. 


90 


Austrobaileya 9(1): 80-101 (2013) 


Viola silicestris : Fully developed leaves 
have been described as broad, occasionally 
sub-reniform, the margin with (5—)7-12(— 
14) obscure teeth, the apical tooth not 
prominently larger than the rest (Thiele & 
Prober 2006). Here, we describe the overall 
outline of a fully developed blade as broad 
or ± rectangular, often with a falcate shape 
and occasionally sub-reniform. When falcate, 
the upper and lower margins are parallel. 
Occasionally a leaf is found with a shape 
resembling the tail of a diving whale. The 
apex is often truncate, sometimes rounded; 
the terminal midvein usually not much longer 
than the two adjacent veins, but sometimes 
shorter. The terminal midvein is usually just 
a small point or mucro. The blades are not 
distinctly lobed and a distinct sinus is not 
present. The base of fully developed leaves is 
truncate, although sometimes slightly curved 
or arched downward between the petiole 
attachment and the outermost edge of the 
blade. Occasionally, the blade bases curve 
upward from the petiole. The basal margin is 
entire or may have a small point or two. The 
distal leaf margin is ± serrate, subserrate, 
sometimes denticulate, or occasionally ± 
crenate. The outline of immature leaves is 
semi-circular or sub-reniform, smaller leaves 
are sometimes ± ovate; the apex obtuse or 
acute; the upper margin serrate, subserrate, 
or ± denticulate, the leaf base attenuate 
(cuneate) in some populations and truncate in 
others. Distinguishing features of this species 
are leaves that are usually much wider than 
long through all stages of growth from new/ 
immature to fully developed. Fully developed 
leaves have a ± rectangular outline or falcate 
shape and never have distinct lobes or a sinus 
(Fig. 7). 

Indumentum 

Viola banksii: This species was described as 
glabrous (Thiele & Prober 2003). However, 
several specimens at BRI collected in Qld 
have pilose hairs on adaxial, or on adaxial and 
abaxial leaf surfaces (e.g., BRI [AQ11278], 
[AQ115406]). A number of other populations 
in Qld appear to be V banksii based on leaf 
shape, but the plants were variously pubescent 
on one or both surfaces, or leaves and petioles 


Fig. 7. V silicestris. Sequence of development from 
immature (1) to fully developed leaf (5). Leaves are 
much wider than long at all growth stages. Adaxial 
surfaces shown. Photo: G. Leiper. 


were densely pubescent (Table 2). V. banksii 
plants are sold commercially worldwide 
under the name “ Viola hederacea ” (Thiele & 
Prober 2003; Elliot & Jones 2010). The first 
author has maintained a clone of V banksii for 
over 10 years purchased as “ Viola hederacea ” 
from a nursery in Sacramento, California, 
U.S.A. The leaves, petioles, and peduncles 
on this clone are usually pubescent (Fig. 
4), similar to some of the populations noted 
in Table 2. When leaves of V banksii are 
pubescent, the margins are usually ciliate, as 
in Vperreniformis. Based on our observations 
of pubescence on V banksii , we believe this 
character state needs to be recognized in 
descriptions of this taxon. Further study is 
needed of the populations noted in Table 2. 

Viola hederacea subsp. hederacea : Leaves 
were described as glabrous or with scattered 
unicellular hairs on the upper surface 
(Thiele & Prober 2003). Our observations 
of populations in southeast Qld and northern 
NSW are similar, although we have observed 
scattered hairs are sometimes present on 
both leaf surfaces. In addition, this species 
occasionally has hairs proximally on petioles 
and peduncles. In comparing the leaves of 
this taxon with V silicestris , a general tenet 
is that V hederacea leaves are often short- 
hairy, at least on adaxial surfaces, whereas V 
silicestris leaves are usually glabrous. 

Viola perreniformis : Adams (1982b) 
stated that leaves of this species were softly 
pubescent, rarely almost glabrous. The authors 


Little & Lei per, Viola perreniformis 91 

Table 2. Populations of pubescent Viola banksii cf. in Queensland 


Location in Queensland 

Comment 

Pine Ridge CP and Jacobs Well 
(Gold Coast coastal areas) 

Plants appear to be V banksii. At both locations 
leaves have scattered hairs; at Jacobs Well, petioles 
are hairy. 

Point Cartwright (Sunshine Coast) 

Plants appear to be V banksii. Leaves are sparsely 
hairy. 

Alexandra Headland (Sunshine 

Coast) 

Plants appear to be V banksii. Leaves and petioles 
are very hairy. 

Maroochy River CP (Sunshine 

Coast) 

Plants appear to be V banksii. Leaves with scattered 
hairs on both surfaces. 

Ningi (Abbey Museum at the 
southern end of the Sunshine Coast) 

Plants appear to be V banksii. Leaves with scattered 
hairs on both surfaces. 


conducted a brief evaluation of pubescence on 
leaf surfaces, petioles, and peduncles among 
V perreniformis populations in Qld and NSW. 
The evaluation revealed substantial variation 
in pubescence on the same plant, between 
plants in the same population, and among 
different populations. Most populations were 
sparsely to densely pubescent on both leaf 
surfaces although the density of hairs usually 
varied between the abaxial and adaxial 
surfaces on the same leaf. Some populations 
had both glabrous and pubescent leaves on 
the same plant. Leaf margins were ciliate or 
eciliate. The petioles and peduncles of most 
populations were sparsely or densely villous, 
although some populations had both glabrous 
and pubescent petioles and peduncles on the 
same plant. No population was completely 
glabrous, although a population at Crows 
Nest NP in Qld was almost completely 
glabrous, with scattered pilose hairs only on 
the cordate lobes of the adaxial leaf surfaces. 
The pubescence on the abaxial surface of 
smaller leaves on the type specimens is 
considered tomentose, but larger leaves are 
considered densely pubescent. Based on our 
observations, we conclude the absence of hairs 
on leaves, petioles, and/or peduncles is not 
taxonomically useful. However, the presence 
of dense pubescence on leaves, petioles, and/ 
or peduncles can be helpful for identifying V 
perreniformis if appropriate leaf shapes and 
flower colours are also present. When leaves 


of V perreniformis are pubescent, the margins 
are normally ciliate, as in V banksii. 

Viola silicestris : Leaves of this species were 
described as glabrous or occasionally sparsely 
short-pubescent when young (Thiele & Prober 
2006). Our observations of populations in 
southeast Qld and northern NSW are similar. 
The basal margins of young leaves on plants 
in Nightcap NP, NSW, occasionally had 
marginal cilia, but were otherwise glabrous. 
Many leaves on the isotype at BRI are ciliate 
C Thiele 2568 [BRI]). 

Flowers. Petal movements and petal 
twisting : The two upper petals of the four taxa 
are sometimes described as “petals erect or 
reflexed”. Reflexed upper petals and twisted 
lateral petals are common traits among the 
four taxa. Both authors have independently 
observed that petals of Viola banksii reorient 
themselves into a ‘sleep’ mode at dusk, then 
‘reopen’ the next day. For example, as dusk 
approaches, the two completely reflexed 
upper petals of V banksii flowers fold forward 
over the front of the flower (a movement of 
c. 180°) and the lateral petals ‘untwist’ and 
extend straight out in front of the flower. In 
the morning of the next day, the upper petals 
reflex backward (another c. 180°) from their 
forward position and the lateral petals spread 
back laterally to their normal daytime position 
and become twisted. Thus, the upper petals 
are ‘erect’ only for a brief period twice a day 


92 


Fig. 8. V. perreniformis. Erect upper petals; all petals 
pale blue-violet with violet-coloured blotches on upper 
and lateral petals. Moogerah Peaks NP, Qld. 2013. Photo: 
G. Leiper. 

(Fig. 8), at least for the first few days after 
the flower first opens. Initial observations 
indicate that after several days, the petals no 
longer exhibit these nyctinastic movements. 
The first author and W. Cherry (pers. comm.) 
have independently observed nyctinastic 
petal movements in flowers of V silicestris. 
This phenomenon is not known to have been 
previously documented in section Erpetion 
and its occurrence elsewhere in the section is 
unknown. 

Flower colours : Flowers of Viola 
perreniformis and V silicestris have been 
described as “concolorous” whereas flowers 
of V banksii and V hederacea have been 
described as “discolorous” (=bicoloured). 
Concolorous is also applied to the flowers 
of V cleistogamoides, V. fuscoviolacea 
and V sieberiana. The term concolorous 
in reference to the overall appearance of 
a flower is useful as an initial assessment 
to determine the species at hand, e.g., V 


Austrobaileya 9(1): 80-101 (2013) 

perreniformis versus V silicestris. However, 
because the face of V perreniformis flowers 
often has darker coloured blotches on the 
upper and lateral petals, darker veins on the 
lowest and sometimes all petals, and because 
the petal bases are white, the description as 
‘concolorous’ is sometimes not so clear cut. 

The description of flower colours on 
collection labels can assist with taxon 
identification, but can be problematic for 
future researchers if ambiguous. For example, 
flower colours recorded for specimens we 
would consider Viola perreniformis based on 
leaf shape include the following descriptions: 
flowers pale blue; flowers purple; flowers 
sky blue; flowers violet with darker purple 
marks; flowers white and purple streaked; 
lilac-white flowers; mauve and white petals; 
violet flowers. None state whether the petal 
tips are white or not. Because the patterns of 
colouration on the face of flowers in the four 
taxa can be complex and because colours 
facilitate identification, we suggest 1) that 
names for flower colours, veins, and blotches 
be based on a readily available standard, and 
2) that separate descriptions be recorded by the 
collector for the colour, presence or absence 
of darker coloured blotches, and presence 
or absence of darker coloured veins, for the 
upper and lateral petals, and for the lower 
(anterior) petal. For all petals, whether the tips 
are white or not should also be recorded. 

The colours on petals of Viola flowers 
fade rather quickly on herbarium sheets and 
if not quickly pressed flat, the petals shrink 
and become distorted. These factors can 
make it difficult or impossible to determine 
the original colour scheme and/or petal shape 
on herbarium specimens. For example, within 
a month after collecting V perreniformis and 
V silicestris flowers, the distal portions of the 
petals, which had been pale blue when fresh, 
had faded in the press to a whitish colour. 

Viola banksii and V hederacea subsp. 
hederacea : Flowers of V banksii were 
described as strongly discolorous violet- 
and-white (Thiele & Prober 2003). Flowers 
of V hederacea were described as usually 
discolorous violet-and-white (occasionally 
concolorous pale violet or almost white), 


93 


Little & Lei per, Viola perreniformis 

the colours usually not strongly demarcated 
{ibid). In general we concur with these 
descriptions, although some populations 
in Qld that appear to be V banksii based on 
leaf shape have flowers that are not strongly 
demarcated and other populations that 
appear to be V hederacea have flowers that 
appear to strongly demarcated. Among the 
possible explanations is that a greater range of 
variation needs to be recognized, or perhaps 
new taxa are involved. 

Viola perreniformis: Adams noted that 
petal colours of V perreniformis were pale 
(lazuline) blue-violet. We concur and have 
observed the abaxial surfaces of the five 
petals (Fig. 9) and portions of the adaxial 
surfaces (i.e., the ‘face’) are usually a pale 
blue-violet colour (Figs. 8, 10). The adaxial 
surfaces often have distinct purple blotches 
of various sizes on the lateral petals and 
occasionally basally on the upper petals (Fig. 
8). In addition, faint to very distinct violet- 
coloured veins are often present on both 
surfaces of the lateral petals and lowest petal. 
The proximal portions on the adaxial surface 
of the upper and lateral petals are usually 
white, although it may be difficult to detect in 
some flowers without manually spreading the 
petals. The lowest petal is white proximally, 
blue-violet distally, and always has distinct, 
darker, violet-coloured veins (nectar guides). 
We observed substantial variation in the 
amount of pigmentation present on the ‘face’ 
of V perreniformis flowers at Crows Nest NP, 
Moogerah Peaks NP and Nightcap NP. 

Viola silicestris : Flowers were described as 
concolorous pale mauve-blue (rarely somewhat 
discolorous) (Thiele & Prober 2006), similar 
to our observations. At Springbrook NP, the 
authors found populations growing on rhyolite 
which had completely white flowers. These 
flowers did not have any darker coloured veins 
(i.e., no nectar guides) abaxially or adaxially 
on the upper, lateral, or lower petals. 

Venation (nectar guides) on lowest petal: 
Venation of the lowest petal was determined 
to be a useful taxonomic character for 
distinguishing V banksii , V hederacea , and V 
silicestris (Thiele & Prober 2003, 2006). We 
found general congruence with the venation 


Fig. 9. Abaxial surface of V perreniformis flowers wtih 
concolorous petals. Note the green and white indurate 
area at base of lowest petal (arrows). Crows Nest NP, 
Mar 2013. Photo: G. Leiper. 


Fig. 10. V perreniformis. Upper petals fully reflexed, 
lateral petals twisted. Crows Nest NP, May 2013. Photo: 
G. Leiper. 

pattern described for these species in some 
populations, but not in others. 

Green and white indurate area: A small, 1-2 
mm long, 1-2 mm wide, green and white area 
is located at the proximal end of the lowest 
petal of the four taxa (Figs. 9, 11). A narrow, 
white, ere scent-shaped zone usually occurs 
between the coloured part of the petal and the 
green area. A similar green and white area has 
been reported in V eminens and is referred to 
as a “small green blotch” or “V-shaped blotch” 
in these species (Thiele & Prober 2003,2006). 
V cleistogamoides also has a green and white 
area on the lowest petal (Thiele & Prober 
2006), but V fuscoviolacea does not (Thiele & 
Prober 2003). No information is available for 
V. sieberiana. In addition to the colour being 
distinctly different from the rest of the petal, 


94 


Fig. 11. V banksii, face view, lateral and upper petals 
removed. Yellow arrows point to anther glands. Green 
and white indurate area (not in focus) in lowest petal 
indicated by white arrow. Photo. J. Little. 

we found the tissue in the green and white 
area in V banksii and V perreniformis flowers 
to be slightly ‘harder’ (indurate) compared to 
the rest of the petal, a feature not previously 
reported for flowers in section Erpetion. 
In flowers of V banksii , V hederacea, and 
V perreniformis , we also observed a small 
green area at the base of the two lateral 
petals similar in colour to the indurate area 
on the lowest petal, a feature not previously 
reported in other species in section Erpetion. 
It has not been determined if the green area 
on the lateral petals is indurate compared 
to the rest of the petal. Although it may be 
presumed that the green and white indurate 
area in some way facilitates pollination 
and/or pollinator attraction, its purpose or 
function is unknown. When a flower is held 
up to a light source, the centre of the indurate 
area is translucent, perhaps functioning as a 
‘window’ for the benefit of a pollinator. The 
lowest petal of most Viola flowers has the 
same texture throughout their length. For 
example, an indurate area at the base of the 
lowest petal was not reported for any of the 96 
Viola species in China (Chen et al. 2008), nor 
any of the 73 Viola species in North America 
(Little & McKinney in press), and may be 


Austrobaileya 9(1): 80-101 (2013) 

unique to species in section Erpetion. 

Style : The style of the four taxa is similar: a 
thin, straight, ± terete structure, except angled 
downward near the distal end toward the 
lowest petal; the style head is glabrous. The 
base of the style in the four taxa is geniculate 
near the attachment to the ovary, similar to 
many Viola species. The geniculate base 
facilitates the style shaft being easily moved 
when ‘pushed aside’ by insects probing for 
nectar or pollen. The geniculate portion of the 
style functions as a spring, allowing the style 
to be easily moved by the pollinator. After 
the pollinator withdraws, the style returns to 
its original position. In some Viola species 
in Europe, when the style is lifted up by a 
probing insect the anthers are spread apart 
and pollen is released (Beattie 1974). 

Anther glands: The term ‘anther gland’, 
apparently first used by Thiele & Prober (2003), 
is an accurate and appropriate name for the 
glands found in the four taxa. Compared to the 
appendage and nectary found in many Viola 
species, the morphology and presentation of 
the anther glands in the four taxa are totally 
different. A few differences are highlighted. 
First, because the flowers have no spur, the 
anther glands are not protected within one; 
second, the anther glands develop laterally 
from and do not extend beyond the base of 
the anther; third, the nectaries in most Viola 
species are situated adjacent to each other 
whereas in the four taxa the anther glands 
are widely separated. In his description of 
the genus Viola , Adams (1982) stated that the 
anterior two anthers have “basal appendages” 
i.e., outgrowths from each anther, the distal 
portion of which secretes nectar. Although 
correct for species in which the two nectaries 
extend into a spur, e.g. V betonicifolia and V 
caleyana , it is inaccurate for the four taxa in 
section Erpetion because the anther glands are 
never longer than the anther and the nectaries 
are not enclosed in a spur. 

When a flower is viewed face on with a 
hand lens, the two anther glands in flowers 
of the four taxa are seen ‘hanging down’ 
like landing gear on a plane (Fig. 11, 12). 
Remarkably, because the anther glands 
are not protected within the confines of a 


95 


Little & Lei per, Viola perreniformis 

spur, they are ‘exposed’ to the elements and 
potentially, it would seem, to physical damage 
from pollinators. Because the anther glands 
are situated above and immediately adjacent 
to the indurate portion of the lowest petal, one 
might assume that nectar would pool in the 
concavity formed by indurate area. However, 
we have not observed nectar in fresh flowers 
of the four taxa and are not aware of data 
demonstrating that nectar is produced in these 
species. In photographs of anther glands from 
freshly collected flowers of Viola banksii, 
V hederacea and V perreniformis , the 
glands were often ‘glistening,’ but obvious 
liquid nectar was not present. The lack of 
observations of nectar is not, of course, 
evidence that none is produced. Perhaps 
because they are not protected within a spur, 
the anther glands in section Erpetion have 
a different function other than to produce 
nectar. Pollen has been observed adhering to 
the anther glands in the four taxa presumably 
due to secretion of a viscid exudate. 

The shape, colour, and surface texture of 
the anther gland is a useful diagnostic feature 
in section Erpetion (Thiele & Prober 2003). 
Table 3 summarizes information for anther 
glands reported in the literature. Table 4 
summarizes preliminary observations made 
by the authors. The green colour of glands 
in some species is very similar to the colour 
of the indurate area of the lowest petal. The 
glands of some species are reported as pale or 
dull green or whitish. However, it is unknown 
if the colour is initially green and transitions 
to white or whitish over time, or vice-versa, 
or if some flowers typically produce only one 
colour, or if the gland is sometimes or always 
± bi-coloured (e.g. white or pale distally, green 
proximally). A similar situation exists for 
gland surface texture. In a brief comparison 
of newer versus older flowers of Viola banksii , 
the first author found that glands of both 
newer and older flowers were white or whitish 
and both were rugose. However, the glands of 
this species have also been reported as pale 
green and smooth (Tables 3 & 4; Fig. 12). 

The anther glands of Viola silicestris 
were described as “irregular, narrow” and an 
illustration shows them to be sinuous (Fig. 


Fig. 12. V. banksii. Lateral view of one of the two anther 
glands (arrow). Lateral and upper petals removed. Photo: 
J. Little. 


1 in Thiele & Prober 2006). However, in a 
population we sampled in Qld, the glands were 
prominent and curved, not narrow and sinuous. 
We believe that differences reported for shape, 
surface texture, and colour among the four 
taxa suggests that additional observations are 
needed. V hederacea sometimes has purplish 
glands (Thiele & Prober 2003). A pure white 
flower of V silicestris from Springbrook 
N.P was found without an anther gland on 
either of the two lower anthers. A critical 
survey is needed of the morphology and 
development of anther glands, the role they 
play in reproductive biology, and a standard 
terminology developed for species in section 
Erpetion. 

Pollination and cleistogamy. With certain 
exceptions, most species in the section have 
relatively large, showy flowers. Comparably 
sized flowers of Viola species in the northern 
hemisphere attract a wide range of insect 
pollinators such as solitary bees, bee- 
flies, syrphid flies and various Lepidoptera 
including butterflies and skippers. Beattie 
(1971) documented 43 different insect species 
in a Viola pollination study in England. 
The authors have not observed potential 
pollinators visiting flowers of any of the four 
taxa, although pollination of chasmogamous 
flowers is assumed to occur based on the many 
developing and dehisced capsules present in 
most populations and the fact that seeds are 
produced. 


96 

The fully reflexed upper petals of the four 
taxa could be an adaptation that facilitates 
sternotribic pollination by providing 
a platform on which insects can land. 
Sternotribic pollination occurs when an insect 
enters a flower ‘upside down/ in contrast with 
nototribic pollination where the insect enters 
the flower in a supine position (i.e. ‘right side 
up’) (Beattie 1974). 

Typical Viola pollinators seek nectar and/ 
or pollen, although some are only after pollen. 
The authors have observed copious amounts 
of pollen in fresh flowers of each of the four 
taxa. A pollination study conducted with two 
Viola species in Brazil provided evidence that 
both species may be shifting from flowers 
that typically produce nectar, to flowers 
that provide primarily pollen (Freitas & 
Sazima 2003). The first author has observed 
yellow-coloured thrips (Thysanoptera) with 
pollen adhering to their bodies in V banksii 
flowers, randomly and rapidly moving over 
and within dehisced anther sacs and around 


Austrobaileya 9 ( 1 ): 80-101 ( 2013 ) 

the ovary. Thrips were reported in Viola 
flowers (Baker 1935) and the first author has 
also observed them on flowers of several 
Viola species in California. Although thrips 
are often observed with pollen attached to 
them, there is currently no evidence they play 
a role in pollination of Viola. However, due 
to numerous observations of thrips in Viola 
flowers, it seems highly probable they could 
periodically affect pollination. 

Cleistogamy occurs in many species of 
Viola throughout the world and is interpreted 
as a mechanism to help ensure survival when 
pollinators are unavailable. In North America, 
c. 85% of all species are known to produce 
cleistogamous (CL) flowers; only nine species 
are known to never produce CL flowers (Little 
& McKinney in press). In Asia, CL flowers 
are very common in most Viola species (J. 
Leng, pers. comm.). The presence or absence 
of CL flowers is not mentioned in Australian 
floras for any species of Viola. The authors 
have not observed CL flowers in the field or 


Table 3. Summary of anther gland characters reported in the literature for Viola 


Species/Source 

Description 

Surface Texture 

Colour 

V banksii 1 

long, narrow and 
high, scarcely 
flattened 

smooth 

pale green to whitish 

V hederacea subsp. 
hederacea 1 

shorter than the 
anther cells, broad 
at the base and each 
distinctly flattened 
or depressed towards 
the other; short, 
rugose, often purple 

irregularly rugose 

purplish or dull green 

V silicestris 2 

slightly shorter than 
the anther cells, 
irregular, narrow 

(Not reported) 

pale 


'Thiele & Prober 2003. 


2 Thiele & Prober 2006. 


Little & Lei per, Viola perreniformis 

Table 4. Summary of anther gland characters recorded by authors 


97 


Species/Source 

Description 

Surface Texture 

Colour 

V banksii 
Springbrook NP, 
Qld 1 

anther gland prominent, 
‘talk or ‘high’, almost as 
long but not as wide as 
anther, wider proximally, 
slightly curved, ‘top’ of 
gland ± flat 

smooth 

pale green 
basally, ± 
greenish-white 
distally 

Venman 

Bushland, NP, 

Qld 

anther gland not 
prominent, short, c. !4 
length of and not as 
wide as anther, gland 
height ‘short’, not ‘tall’ 
or ‘high’, appears ± ‘flat’ 
entire length of gland 

smooth 

white or whitish 
with a greenish 
tinge 

Plant sold in 

U.S.A. as “V. 
hederacecT 

anther gland prominent, 
‘tall’ or ‘high’ almost 
as long but not as wide 
as anther, much wider 
proximally, curve, ‘top’ 
of gland ± flat 

rugose 

white or whitish 
or pale green 
entire length 

V hederacea 
subsp. hederacea 
Springbrook NP, 
Qld 

anther gland prominent, 
almost as long and 
about as wide as the 
anther, gland is wider 
proximally, arched 
(curved), taller distally 

at lower magnifications, 
the gland appears 
irregularly rugose; at 
higher magnifications the 
surface appears smooth; 
the uneven, rugose 
appearance appears to 
be caused by clumps of 
pollen grains adhering 
to the glandular exudate; 
the normally white pollen 
grains then take on a 
greenish colour 

pale green 
basally, ± 
whitish distally; 
sometimes 
glistening when 
fresh 

V perreniformis 

anther gland prominent, 
c. Vz-% as long as anther 

smooth 

pale green 

V silicestris 
Springbrook NP, 
Qld 

anther gland prominent 
(on a violet-tinged 
anther), almost as long 
as, but narrower than the 
anther, proximal portion 
curved toward other 
gland 

not determined 

yellowish to 
pale 


‘Possibly a horticultural escape. 


98 

on herbarium specimens of the four taxa, nor 
on V sieberiana. Cleistogamous flowers have 
not been observed on herbarium specimens 
of V cleistogamoides or V fuscoviolacea (the 
authors have not observed these species in 
the field). However, detecting CL flowers on 
herbarium specimens is difficult because CL 
flowers can appear similar to undeveloped or 
aborted chasmogamous flowers. 

Viola species exhibit numerous strategies 
that promote reproductive fitness, one of 
which is hybridization that results in the 
formation of amphiploids, diploids, and 
tetraploids. As the authors have found and 
others have reported (Thiele & Prober 2006), 
some of the four taxa and other species in 
section Erpetion are sympatric. For example, 
in NSW sympatric populations occur with V 
perreniformis and V silicestris ; V hederacea 
and V silicestris ; and V fuscoviolacea , V 
sieberiana and V. silicestris ; and in Qld, V 
hederacea and V silicestris. However, among 
the five species mentioned, no hybrids have 
yet been found (Thiele & Prober 2003, 2006). 
An example demonstrating that hybridization 
can potentially occur in section Erpetion was 
reported in Victoria between V eminens and 
V fuscoviolacea when an apparently sterile, 
but vegetatively vigorous putative hybrid, V x 
zophodes , was found (Thiele & Prober 2003). 

Adams (1982a) noted in the key to 
subspecies of Viola hederacea that subsp. 
fuscoviolacea L.G.Adams (=V fuscoviolacea) 
was faintly scented. No other information on 
floral odours of species in section Erpetion is 
known to the authors. 

Seeds and seed dispersal. Seeds of the four 
taxa, like most Viola species, have an ovoid 
outline, a shape that is assumed to facilitate 
ballistic travel after ejection from a capsule 
valve. The seeds of many Viola species 
possess an outgrowth, called an elaiosome or 
caruncle that develops around and extends 
out from the micropyle. Elaiosomes are 
fleshy food bodies of variable size that can 
be attractive to ants. Although the number 
of seeds examined in this study was limited, 
an obvious elaiosome was not detected on 
mature, recently collected seeds of three of 
the four taxa nor on V sieberiana (seeds not 


Austrobail'eya 9(1): 80-101 (2013) 

available for V banksii ). Seed characters 
reported in the literature and observed by the 
authors for the four taxa and V sieberiana 
are summarized in Table 5. The seeds are 
small, 1-2 mm long, although V banksii 
seeds are sometimes up to 2.5 mm long 
(Thiele & Prober 2003). While the first 
author was studying seeds of V sieberiana , 
initial observations suggested that mature 
seeds were reddish-brown, until shiny black 
seeds were found. At least for this species, 
it appears that a reddish-brown colour 
transitions into black. 

The apparent lack of an elaiosome in 
three of the four taxa and Viola sieberiana 
might lead one to suspect their seeds are not 
dispersed by ants. However, seeds of a Viola 
species in Europe with almost no elaiosome 
were reported to be removed by ants (Beattie 
& Lyons 1975). Lengyel etal. (2010) estimated 
that over 70% of Violaceae species throughout 
the world are myrmecochorous. 

Based on numerous observations of 
dehisced capsules split into three valves on 
herbarium specimens and in the field, the 
dispersal mechanism of seeds from capsules 
in the four taxa and other species in section 
Erpetion is assumed to be similar to V 
betonicifolia (Little & Leiper 2012). However, 
studies are needed to determine ballistic 
dispersal distances and to determine if ants 
play any role in dispersing seeds. 

Taxonomic challenges 

There is considerable evidence that Viola 
populations exhibit character divergence 
among local colonies of single species (Beattie 
1976). It should therefore not be too surprising 
to find morphological variation among 
isolated populations of species in section 
Erpetion. Thiele & Prober (2003) noted there 
are several undescribed species in the V 
hederacea complex in NSW. Recognition of 
new taxa may help resolve situations where 
observed character states in a population 
conflict with the current taxonomy of the 
complex. Additional study of the four taxa 
may reveal other morphological characters or 
relationships between characters that provide 
additional or better approaches for taxonomic 


Little & Lei per, Viola perreniformis 

Table 5. Seed characters of five species in Viola section Erpetion 


99 


Species/ Location & 
Source 

Mature Seed Colour 

Surface Texture 

Seed Length 

V banksii 1 * * * V 

glossy purplish-black 

± smooth to 
irregularly rugose 

1.8-2.5 mm 

V hederacea subsp. 
hederacea 1 

dull, mottled cream and 
brown (occasionally 
uniformly reddish-brown) 

± smooth 

1.2-2.0 mm 

Springbrook NP 2 

reddish-brown 

warty-rugose 

1.7-1.8 mm 

V perreniformis 2 
Nightcap NP, NSW 

shiny or dull dark brown 
mottled with patches of light 
brown or tan 

uneven to 
irregularly rugose 

1-1.2 mm 

V sieberiana 2 

shiny black 

warty-rugose 

1.5-1.8 mm 

V. silicestris 3 

glossy purplish-black 

not reported 

1.5-1.7 mm 

Springbrook NP 2 

shiny black 

warty-rugose 

1.6-1.7 mm 


‘Thiele & Prober 2003. 

2 Current authors. 

3 Thiele & Prober 2006. 

recognition. Relationships in section Erpetion 
are largely unknown and will probably require 
DNA sequencing and chromosome counts to 
elucidate taxonomic relationships (Thiele & 
Prober 2006). 

The provisional keys provided herein 
are based on the authors’ observations and 


information in Thiele & Prober (2003, 2006). 
Among the four taxa, the flowers of Viola 
perreniformis are most similar to V silicestris ; 
flowers of V banksii are most similar to V 
hederacea. Fully mature leaves of V banksii 
are in general more similar to Vperreniformis , 
whereas leaves of V hederacea are in general 
more similar to V silicestris. 


Provisional Key to selected species in Viola section Erpetion 
(based on flowers and seeds) 

1 Abaxial surfaces of all petals usually of one colour (concolorous), entire 

surface pale violet, pale blue, or pale lavender, whitish or pale coloured 

proximally; on adaxial surface (face), c. half of upper and lateral 

petals pale violet or pale blue distally; (abaxial and adaxial surfaces of 

V silicestris rarely mostly or entirely white); lowest petal obovate to 

oblong.2 

1. Abaxial surface of all petals usually of two colours (bi-coloured, 
or discolorous), petals white distally and pale violet proximally 
(c. 2/3 of upper petals white distally, pale violet proximally) 
on adaxial surface (face) c. 2/3 of upper petals white distally, violet 
proximally; c. half of lateral petals white distally, violet proximally; 
lowest petal obovate, elliptic, or circular.3 


100 


Austrobaileya 9 ( 1 ): 80-101 ( 2013 ) 


2 Anthers cream-coloured; fully mature seeds shiny or dull dark brown, 

1-1.2 mm long.V. perreniformis 

2. Anthers violet-tinged; fully mature seeds shiny black, 1.5-17 mm long ... V. silicestris 

3 Anther gland smooth or rugose, white or whitish to pale green; the 

midvein and the two adjacent veins on lowest petal usually prominent and 
distinct, often without interconnecting 2° veins; the violet and white areas 
on petals usually strongly demarcated.V. banksii 

3. Anther gland usually rugose, dull green or purplish; the midvein on 

lowest petal often less distinct than the two adjacent veins, often with 
several to many 2° veins connecting them; the violet and white areas on 
petals usually not strongly demarcated.V. hederacea 


Provisional Key to selected species in Viola section Erpetion 
(based on mature, fully developed leaves) 

1 Leaf base usually cordate, sinus and lobes usually present, sometimes 

overlapping, blades about as wide as long (< 2x wider than long), blade 
outline ovate, orbicular, suborbicular, or semi-circular, blade shape 
orbicular, reniform, or ovate.2 

1. Leaf base usually truncate, sinus and lobes not present, blades usually 

much wider than long (> 2x wider than long), blade outline 
semi-circular or ± rectangular, blade shape broad-reniform or 
sub-reniform (but lobes not prominent), or falcate (with the upper 
and lower margins parallel).3 

2 Leaf margin not crenate from basal lobes to apex, margin nearly entire, 

sometimes ± scalloped or sinuous, outline usually orbicular, blade shape 
orbicular or reniform, base cordate, lobes often overlapping in fresh 
material, apex ± truncate, rounded or obtuse, occasionally ± dentate, 
the midvein usually not extending much beyond the length of adjacent 
veins, both leaf surfaces glabrous or with scattered, short hairs, never 
tomentose, leaf blades about as wide as long.V. banksii 

2. Leaf margin regularly or irregularly crenate from basal lobes to apex, 

outline ovate, orbicular, suborbicular, or semicircular, blade shape 
reniform, sub-reniform, or ovate, base cordate, lobes seldom overlapping 


in fresh material, apex usually acute, not truncate, midvein usually 
extending beyond length of adjacent veins, both leaf surfaces often softly 
hairy, occasionally tomentose, rarely sub-glabrous; leaf blades usually 
slightly longer than wide.V. perreniformis 

3 The distal (upper) blade margin ± entire or with only a few short points; 

stems not known to occur at stolon nodes.V. hederacea 

3. The distal (upper) blade margin ± serrate or subserrate, sometimes 
denticulate, occasionally ± crenate; apex often truncate, sometimes 
rounded; stems (1.5-40 cm) sometimes present at stolon nodes .V. silicestris 


101 


Little & Leiper, Viola perreniformis 

Acknowledgements 

The authors thank the curators at BRI, 
CANB, and NSW for access to specimens 
and assistance. We thank RI. Forster for 
encouragement to prepare this paper and 
K.R Thiele for constructive comments on an 
earlier draft. The assistance of B. Lepschi, 
Curator, Australian National Herbarium, 
is greatly appreciated for providing images 
of the holotype of Viola hederacea subsp. 
perreniformis and scans of photographs of the 
type of V hederacea. Thanks to P. Bostock for 
translating the original Latin description and 
for scanning herbarium specimens. Thanks 
to J. Brock (Toowoomba, QLD), B. Neilly 
(Hurstville, NSW), and R. Ollerenshaw 
(Canberra, ACT) for timely processing of 
collection/export permits in early 2013 for the 
first author. 

References 

Adams, L.G. (1982a). Appendix. Violaceae. In A.S. 
George (ed.). Flora of Australia 8: 386. 
Australian Government Publishing Service: 
Canberra. 

Adams, L.G. (1982b). Viola. In A.S. George (ed.). Flora 
of Australia 8: 91-100. Australian Government 
Publishing Service: Canberra. 

Baker, M.S. (1935). Studies in western violets. I. 
Madrono 3: 51-57. 

Beattie, A.J. (1971). Pollination mechanisms in Viola. 
New Phytologist 70: 343-360. 

- (1974). Floral evolution in Viola. Annals of the 

Missouri Botanical Garden 61: 781-783. 

- (1976). Plant dispersion, pollination and gene 

flow in Viola. Oecologia 25: 291-300. 

Beattie, A.J. & Lyons, N. (1975). Seed dispersal in 
Viola (Violaceae): adaptations and strategies. 
American Journal of Botany 62: 714-722. 

Chen, Y., Yang, Q., Ohba, H. & Nikitin, V. (2008). 
Violaceae. In Flora of China 13: 72—111. 
eFloras 2008. Published on the Internet http:// 
www.efloras.org Missouri Botanical Garden: 
St. Louis, MO & Harvard University Herbaria: 
Cambridge, MA. Accessed 28 May 2013. 

Duretto, M.F. (2009). Violaceae. In M.F. Duretto 
(ed.). Flora of Tasmania Online. Tasmanian 
Herbarium, Tasmanian Museum & Art Gallery: 
Hobart, www.tmag.tas.gov.au/floratasmania. 
Accessed 28 May 2013. 


Electronic Flora of South Australia (2007). http:// 
www.flora.sa.gov.au/cgi-bin/speciesfacts_ 
display. cgi?form=speciesfacts&family=&gen 
us=viola&species=&iname=&submit=Display 
Accessed 28 May 2013. 

Elliot, R. & Jones, D. (2010). Encyclopaedia of 
Australian plants suitable for cultivation 9: 441. 
Lothian: Sydney. 

Entwisle, T.J. (1996). Violaceae. In N.G. Walsh & T.J. 
Entwisle (eds.), Flora of Victoria 3: 361-370. 
Inkata Press: Melbourne. 

Freitas, L. & Sazima, M. (2003). Floral biology and 
pollination mechanisms in two Viola species - 
from nectar to pollen flowers? Annals of Botany 
91: 311-317. 

James, T.A. (1990). Violaceae. In G. J. Harden (ed.), Flora 
of New South Wales 1: 435-441. Royal Botanic 
Garden: Sydney. 

- (2013). Family Violaceae, In PlantNET - The 

Plant Information Network System of The 
Royal Botanic Gardens and Domain Trust, 
Sydney, Australia (version 2.0). http://plantnet. 
rbgsyd.nsw.gov.au Accessed 28 May 2013. 

Lengyel S., Gove, A.D., Latimer, A.M., Majer, J.D. 
& Dunn, R.R. (2010). Convergent evolution 
of seed dispersal by ants, and phylogeny 
and biogeography in flowering plants: A 
global survey. Perspectives in Plant Ecology, 
Evolution and Systematics 12: 43-55. 

Little, R.J. & Leiper, G. (2012). Capsule dehiscence 
in Viola betonicifolia Sm. (Violaceae). 
Austrobaileya 8: 624-633. 

Little, R.J. & McKinney, L.E. In press. Violaceae. In 
Flora of North America Editorial Committee 
(eds.). Flora of North America North of Mexico , 
Vol. 6. Flora of North America Association: 
New York & Oxford. 

Stanley, T.D. & Ross, E.M. (1983). Flora of South¬ 
eastern Queensland 2: 99-101. Queensland 
Department of Primary Industries: Brisbane. 

Thiele, K.R. & Prober, S.M. (2003). New species and 
a new hybrid in the Viola hederacea species 
complex, with notes on Viola hederacea Labill. 
Muelleria 18: 7-26. 

- (2006). Viola silicestris , a new species in Viola 

section Erpetion from Australia. Telopea 11: 
99-104. 


Pterostylis caligna M.T.Mathieson (Orchidaceae), 
a new species from northern Queensland 

M.T. Mathieson 

Summary 

Mathieson, M.T. (2013). Pterostylis caligna M.T.Mathieson (Orchidaceae), a new species from 
northern Queensland. Austrobaileya 9(1): 102-106. Pterostylis caligna is described as new. It is a 
tuberous, terrestrial herb known only from the Hann and Windsor Tablelands in northern Queensland 
and is readily distinguished from any other members of the genus by its unique combination of 
characters. The species is illustrated and diagnosed herein. Its conservation status is assessed and a 
status of Vulnerable is recommended. 

Key Words: Orchidaceae, Pterostylis , Pterostylis caligna , Australia flora, Queensland flora, new 
species, taxonomy, conservation status. 

M.T.Mathieson, Queensland Herbarium, Department of Science, Information Technology, Innovation 
and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4006, Australia. 
Email: Michael.Mathieson@science.dsitia.qld.gov.au 


Introduction 

The greenhood orchids of Australia are 
diverse and occur in many habitats throughout 
eastern and southern Australia (Janes & 
Duretto 2010; Jones & Clements 2002) with 
a number of new Pterostylis species having 
been described in recent years (Jones 2009a, 
2009b, 2010a, 2010b; Jones & French 2012). 
Nine species are currently known from the 
Wet Tropics of north Queensland (Jones 2006, 
2010a; Bostock & Holland 2010). 

During a visit to the Hann Tableland, 
west of Mareeba in north Queensland in May 
2010, the author discovered a population 
of Pterostylis R.Br. that did not match any 
named species. However, one specimen at 
BRI corresponded with this species. Further 
material was collected in March 2012 and 
April 2013, enabling its distinctiveness to be 
confirmed and its formal description herein. 
I follow the classification of Janes & Duretto 
(2010) under which this species keys to 
Pterostylis subg. Pterostylis sect. Foliosae. 

Materials and methods 

This research is based on a study of herbarium 
specimens and associated spirit collection at 
BRI and CNS, and field studies at the type 


Accepted for publication 5 September 2013 


and other noted localities. All measurements 
have been made from live material or material 
preserved in spirit. 

Abbreviations in the specimen citations 
include NP (National Park). RE refers to 
Regional Ecosystem, descriptions of which 
can be viewed at (in this case): http://www. 
ehp.qld.gov.au/ecosystems/biodiversity/ 
regional-ecosystems/index.php 

Taxonomy 

Pterostylis caligna M.T.Mathieson sp. nov. 
ab omnibus aliis speciebus Pterostylis sepalis 
lateralibus clavatis, galea decurva apice sepali 
dorsalis filiformis clavati, labello semper 
abscondito et apice suo rotundato vel obtuse 
acuto distat. Typus: Queensland. Cook 
District: Hann Tableland National Park, NW 
of Mareeba, 8 April 2013, M.T. Mathieson 
MTM1438 & P.I.Forster (holo: BRI [1 sheet + 
spirit]; iso: CNS, distribuendi ). 

Tuberous, terrestrial herb growing in clonal 
colonies. Tubers globose to ovoid, 6-9 mm 
long, 5-9 mm wide when fertile. Leaves 3-5 
in a rosette encircling the base of the scape, 
mid-green; lamina linear-ovate to oblong- 
ovate, 14-34 mm long, 9-19 mm wide, 
margins entire or occasionally crisped, apex 
obtuse to bluntly acute; petioles 3—13 mm 
long, slender, not winged. Scape 42-100 mm 


Mathieson, Pterostylis caligna 

tall, 0.9-1.2 mm wide, glabrous; stem bracts, 
sterile 1 (rarely 2), fertile 1, 8-15 mm long, 
2.5-5 mm wide, lanceolate, acute, sheathing 
at base, margins entire. Flower solitary 17-23 
mm long, erect, translucent white with pale 
to bright green stripes on the galea and sinus. 
Galea slightly gibbous at the base, then erect 
before curving forwards in a semicircular 
fashion with the tip decurved. Dorsal sepal 
translucent white with bright green stripes, 
three narrow inner stripes and two broad 
marginal stripes which coalesce in the distal 
quarter, 15-20 mm long, 6-9 mm wide, 
inflated at the base and tapered beyond to the 
apex with a slight constriction in the proximal 
third, apex filiform, clubbed, 2.5-3 mm long. 
Lateral sepals erect closely embracing the 
galea; sinus protruding slightly to moderately, 
shallowly curving inwards, the upper margin 
shallowly notched at the centre then curving 
upwards; conjoined part 7-10 mm long, 6-8 
mm wide, narrowed to c. 2.5 mm at the base, 
bright green fading somewhat towards the 
base, the upper margins inrolled, gradually 
tapered into the free points; free points 
filiform, minutely clubbed, 8-13 mm long, 
erect, held upright above the galea. Petals 
asymmetrically lanceolate, falcate, 13-16 mm 
long, 3.5-4.5 mm wide, tapered in the distal 
half, apex acute to subattenuate, largely bright 
green with a small area of the central proximal 
section translucent pale green to white; flange 
1-1.2 mm wide, broadly deflate, obtuse. 
Labellum attached to a short basal claw c. 1 
mm long, erect, slightly curved in the distal 
quarter, not visible through the sinus in the 
set position, green in the proximal half and 
suffused tan-brown distally; lamina narrowly 
oblong to narrowly oblong-lanceolate, 5-7 mm 
long, 1.5-2 mm wide, slightly constricted in 
the distal half; apex bluntly acute to rounded; 
central callus slightly raised, running the 
length of the labellum ending just short of 
apex, c. 0.25 mm wide tapered to c. 0.1 mm in 
the centre, green in proximal half, tan-brown 
distally; basal appendage linear 2-3 mm 
long, curved, apex irregularly lobed. Column 
green, brown and white, 6-7 mm long, erect, 
angled away from ovary at about 40° in the 
proximal quarter, then erect. Column wings 
3-4 mm long; basal lobe 1.5-2 mm long, c. 


103 

1.2 mm wide, green and brown, at an angle 
of 50°, apex obtuse, inner margins incurved, 
sparsely adorned with short, translucent white 
cilia; mid section 0.8-1 mm long, brown 
and white; apical lobe, white, linear, 0.6-0.7 
mm long, obtuse. Stigma elliptic, 1.5-1.8 
mm long, c. 0.75 mm wide, raised, situated 
centrally on the column. Anther c. 0.7 mm, 
shortly rostrate. Pollinia linear-clavate, 0.6- 
0.7 mm long, yellow, mealy. Ovary cylindrical 
to slenderly obovoid, 6-11 mm long, 1.5-2 
mm wide, green, irregularly ribbed. Capsule 
narrowly obovoid, 13-18 mm long, 4-5 mm 
wide, ribbed, smooth. Fig. 1-3. 

Additional specimens examined : Queensland. Cook 
District: Hann Tableland NP, NW of Mareeba, May 
2010, Mathieson MTM828 & Forster (BRI); Windsor 
Tableland NP, N of Mt Carbine, Apr 2013, Mathieson 
MTM1523, 1555 & Forster (BRI); Windsor Tableland, 
Jan 2009, Sankowsky 3084 & Sankowsky (BRI). 

Distribution and habitat : Pterostylis caligna 
is endemic to northern Queensland and has 
been found only in the Hann Tableland NP, 
about 20 km north-west of Mareeba and Mt 
Windsor NP approximately 90 km north- 
north-west of Mareeba. It grows in simple 
notophyll vine forest with emergent Agathis 
robusta (C.Moore ex F.Muell.) F.M.Bailey on 
granite (RE 7.12.16a) on the Hann Tableland 
and in simple to complex notophyll vine forest 
with emergent Agathis robusta (RE 7.12.7a) 
on the Windsor Tableland between 880 m 
and 1000 m. At both localities, this species 
appears to be associated with vine forest- 
open forest ecotones. 

Phenology : Flowers and fruits are recorded 
for January through to April; also in May, 
when very few flowering plants were observed 
and most fruit had dehisced. 

Affinities : Pterostylis caligna stands apart 
from all other members of the genus by 
the following characters: clubbed lateral 
sepals, decurved galea with apex of dorsal 
sepal filiform and clubbed, labellum always 
concealed when in the set position and 
labellum apex rounded or bluntly acute. Sterile 
rosettes can resemble those of Pterostylis 
taurus M.A.Clem. & D.L.Jones. 


104 


A ustrobaileya 9(1): 102-106 (2013) 


Fig. 1. Pterostylis caligna. A. whole plant *2. B. rosette *2. C. root stock and tuber x4. D. flower from the side x3. 
E. flower from the front x3. F. undehisced capsule x3. All from Mathieson MTM1438 & Forster (BRI). Del. W.Smith. 


Mathieson, Pterostylis caligna 


105 


Fig. 2. Pterostylis caligna. A. lateral sepals (synsepalum) M. B. petal *4. C. column and labellum from the side *8. 
D. column from the front xl2. E. labellum from above xl2. F. labellum apex x32. G. basal appendage of labellum from 
above x32. All from Mathieson MTM1438 & Forster (BRI). Del. W.Smith. 


Conservation status : This species is known 
from three locations; one within the Hann 
Tableland NP and two in the Windsor 
Tableland NP. It was not encountered 
elsewhere on the Hann Tableland despite 
extensive searches in similar habitats during 
three expeditions between 2010 and 2013. It 
is likely to occur elsewhere on the Windsor 
Tableland and adjacent ranges in the Wet 
Tropics biogeographic region given the extent 
of appropriate habitat that exists. 


The population at the type locality 
is estimated to be between 100 and 200 
plants occupying an area of less than one 
hectare. The two populations on the Windsor 
Tableland were estimated at between 300 
and 400 plants, both occupying an area of 
less than two hectares. Repeated fire events 
may threaten this species by encouraging the 
encroachment of introduced, hyperinvasive 
grasses (viz. giant rat’s tail grass Sporobolus 
pyramidalis (Lam.) Hitchc., grader grass 
Theme da quadrivalvis (L.) Kuntze and 


106 


gamba grass Andropogon gayanus Kunth), 
the rampant herbaceous weed Praxelis 
clematidea R.M.King & H.Rob. and lantana 
Lantana camara L. into its’ ecotone habitats. 
Applying criteria of the IUCN (IUCN 2001), 
the recommended conservation status is 
Vulnerable (Dl, D2). 

Etymology : From the Greek calignis - of the 
mists, in reference to the upland cloud forest 
this species inhabits. 


Fig. 3. Pterostylis caligna , side view of the flower, 
photographed in habitat ( Mathieson MTM828). 

Acknowledgements 

Collections of this species were made on a Bush 
Blitz nature discovery expedition in 2010 and 
in 2012 and 2013 using a Bush Blitz Tactical 
Taxonomy Grant and Applied Taxonomy 
Grant respectively, awarded to MTM; all 
funded by the Commonwealth Government in 
part. I thank Will Smith for the illustrations, 
and Peter Bostock for the Latin diagnosis. 
The following people and organisations are 
thanked for their assistance that made the 
survey possible: Keith McDonald (formerly of 


Austrobaileya 9(1): 102-106 (2013) 

Dept, of Environment & Heritage Protection 
(DEHP) - Atherton) for logistical support 
and co-ordination; Traditional Owners for 
the Hann Tableland, John and Troy Grainer; 
Cairns/Mareeba/Mossman National Parks & 
Wildlife Service staff (Jonathon Roth, Robert 
Miller, Brendan Malone, Andrew Hedges and 
Alastair Freeman) and Queensland Herbarium 
staff (Paul Forster, Megan Thomas, Gordon 
Guymer, Peter Bostock, Ashley Field, Eda 
Addicott and Mark Newton) for logistical 
and field support; Cape York Tenure Unit 
Georgianna Fien, Eric Wason); Cape York 
Helicopters and Pat Gorman for safe and 
efficient helicopter provision. 

References 

Bostock, P.D. & Holland, A.E. (eds) (2010). Census 
of the Queensland Flora 2010. Queensland 
Herbarium, Department of Environment and 
Resource Management: Brisbane. 

Iucn (2001). IUCN Red List of Categories and 
Criteria: Version 3.1. IUCN Species Survival 
Commission: Gland (Switzerland)/Cambridge 
(United Kingdom). 

Janes, J. & Duretto, M. (2010). A new classification 
for subtribe Pterostylidinae (Orchidaceae), 
reaffirming Pterostylis in the broad sense. 
Australian Systematic Botany 23: 260-269. 

Jones, D.L. (2006). A complete guide to native orchids of 
Australia, including the island territories. New 
Holland: Sydney. 

- (2009a). Five threatened new species of 

Hymenochilus (Orchidaceae) from southern 
Australia. The Orchadian 16: 176-187. 

- (2009b). Oligochaetochilus pedinus, a new 

species of Orchidaceae from south-western 
New South Wales. The Orchadian 16: 214-221. 

- (2010a). Four new species of Oligochaetochilus 

(Orchidaceae) from tropical Queensland. The 
Orchadian 16: 302-311. 

- (2010b). A new species of Oligochaetochilus 

(Orchidaceae) from Queensland with affinities 
to O. woollsii. The Orchadian 16: 370-375. 

Jones, D.L. & Clements, M.A. (2002). A review of 
Pterostylis (Orchidaceae). Australian Orchid 
Research 4: 1-168. 

Jones, D.L. & French, C.J. (2012). Two new species of 
Pterostylis from Western Australia. Australian 
Orchid Review 77(4): 39-44. 


Two new subspecies of Eucalyptus tereticornis Sm. (Myrtaceae) 


A.R. Bean 
Summary 

Bean, A.R. (2013). Two new subspecies of Eucalyptus tereticornis Sm. (Myrtaceae). Austrobaileya 
9(1): 107-113 Two new subspecies of Eucalyptus tereticornis Sm., E. tereticornis subsp. rotunda 
A.R.Bean and E. tereticornis subsp. basaltica A.R.Bean, are described and illustrated. A key to 
the four subspecies of E. tereticornis is provided, together with a distribution map for the new taxa. 

Key Words: Myrtaceae, Eucalyptus , Eucalyptus tereticornis subsp. basaltica , Eucalyptus tereticornis 
subsp. rotunda , new subspecies, Australia flora, Queensland flora, identification key, taxonomy 

A.R. Bean, Queensland Herbarium, Department of Science, Information Technology, Innovation 
and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. 
E-mail: Tony.Bean@science.dsitia.qld.gov.au 


Introduction 

Eucalyptus tereticornis Sm. is one of the 
best known members of the largely eastern 
Australian ‘Red Gum’ group (E. sect. 
Exsertaria L.D.Pryor & L.A.S.Johnson ex 
Brooker), characterised by the reniform 
cotyledons, concolorous adult leaves, ovules 
in 6-8 longitudinal rows, and the toothed seed 
coat. Brooker (2000) placed E. tereticornis 
in E. ser. Erythroxylon (Blakely) Brooker, 
along with 14 other species, distinguished 
by the patchily deciduous bark, leaves with 
moderately dense reticulation, erect stamens, 
pyramidal or obliquely pyramidal seeds and 
axillary inflorescences. 

Eucalyptus tereticornis is a widespread 
and variable species; it extends from eastern 
Victoria to north Queensland, and to 
Papua New Guinea. Brooker & Slee (2000) 
documented a number of variants of E. 
tereticornis : E. tereticornis subsp. tereticornis 
(long operculum swollen at the base), the 
‘Hunter Valley’ form (with short obese 
opercula, ± parallel-sided), the ‘Tenterfield- 
Cunningham’s Gap’ form (operculum shorter 
than typical form and lacking the swollen 
base), E. tereticornis subsp. mediana Brooker 
& Slee (from eastern Victoria), and E. blakelyi 
-E. tereticornis intergrades. 


Accepted for publication 10 October 2013 


Eucalyptus tereticornis subsp. tereticornis 
is the most widespread subspecies, extending 
from the south coast of New South Wales 
to north of Cooktown in Queensland, and 
Central and Oro (Northern) Districts in 
Papua New Guinea. In southern Queensland, 
it extends west to near Roma (Queensland 
Herbarium 2013). 

Eucalyptus tereticornis subsp. basaltica 
A.R.Bean, described here, is a widespread 
tree on hills and ridges (predominantly 
basaltic) of southern Queensland and northern 
New South Wales. The ‘Hunter Valley’ form 
and ‘Tenterfield-Cunningham’s Gap’ form of 
Brooker & Slee (2000) are both attributable to 
this subspecies. 

An additional taxon, described here 
as Eucalyptus tereticornis subsp. rotunda 
A.R.Bean, is a riparian tree of restricted 
distribution from the upper reaches of the 
Maranoa River of southern Queensland with 
large orbicular juvenile leaves. 

Eucalyptus tereticornis subsp. mediana 
Brooker & Slee differs from E. tereticornis 
subsp. tereticornis by the glaucous juvenile 
leaves, shorter obese operculum, mixed 
black single-coated and yellow double-coated 
seeds, and the later flowering period (Brooker 
& Kleinig 1999). It inhabits stream banks and 
river floodplains, but also low hills and plains 
not subject to flooding (Nicolle 2006). 


108 


Eucalyptus tereticornis subsp. mediana 
and E. tereticornis subsp. rotunda are 
geographically isolated from the other 
subspecies, although the latter occurs in 
close proximity to an outlying stand of E. 
tereticornis subsp. basaltica. E. tereticornis 
subsp. tereticornis and E. tereticornis 
subsp. basaltica are parapatric in southern 
Queensland and northern New South Wales, 
and some hybridisation or intergradation is 
suspected between the two taxa. 

Subspecies rank is considered appropriate 
for the new taxa, as there are consistent 
characters to separate them from E. 
tereticornis subsp. tereticornis. 

Materials and methods 

This study is based on a combination of 
field studies in Queensland and northern 
New South Wales, measurements made 
during an earlier seedling trial that included 
Eucalyptus tereticornis subsp. tereticornis 
and E. tereticornis subsp. basaltica , and 
an examination of specimens at BRI. E. 
tereticornis subsp. mediana has not been 
observed in the field, and knowledge of the 
taxon has been gleaned from the literature and 
from herbarium specimens. Measurements 
have been made from dried herbarium 
material. 


Austrobaileya 9(1): 107-113 (2013) 

Common abbreviations used in the text 
include Mt (Mountain), S.F. (State Forest). 

Taxonomy 

Eucalyptus tereticornis Sm., Spec. Bot. New 
Holland , part 4, 41 (1795). Type: New South 
Wales. Port Jackson, 1793, J. White s.n. (holo: 
LINN; iso: BM ,fide Chippendale [1988: 324]). 

Tree to 50 m high. Bark smooth throughout 
or with persistent grey, scaly-fibrous bark 
at base, forming a stocking less than one- 
tenth tree height; smooth bark deciduous, 
somewhat shiny, dappled white, grey, brown 
or pale blue, shedding in irregular strips 
less than a metre long. Adult branchlets not 
glaucous. Seedling leaves broadly ovate 
to orbicular, stems quadrangular. Juvenile 
leaves petiolate, alternate, broadly-lanceolate 
to orbicular, concolorous or slightly 
discolorous. Adult leaves alternate, narrow- 
lanceolate to lanceolate or falcate, 8-21 x 
1-3.5 cm, concolorous, green, petiolate, 
lateral veins at 35-55° to the midrib, vein 
reticulation moderately dense, oil glands 
abundant, several per areole; intramarginal 
vein less than 20% of distance from margin 
to midrib, margins entire, base cuneate. 
Inflorescence with umbels borne singly in 
leaf axils or along leafless stems. Individual 
umbels 7(-9)-flowered; peduncle terete or 


Key to subspecies of Eucalyptus tereticornis 

1 Opercula of mature buds expanded at base; juvenile leaves broadly 

lanceolate to ovate, 2-3.2 times longer than wide . . . E. tereticornis subsp. tereticornis 

1. Opercula of mature buds not expanded at base; juvenile leaves orbicular 

to ovate, 0.85-2 times longer than wide.2 

2 Juvenile leaves orbicular, 0.85-1.2 times longer than wide, 5-16 cm 

across, green, apex obtuse, apiculate or emarginate . . . . E. tereticornis subsp. rotunda 

2. Juvenile leaves broadly ovate to ovate, 1.2-2.0 times longer than wide, 

dull bluish, apex acute.3 

3 Pedicels 1-3 mm long; opercula 1.3-1.6 times longer than broad; new 

growth of juvenile plants pruinose; margins of juvenile leaves usually 

undulate; seeds coats a mixture of black and yellow. . . E. tereticornis subsp. mediana 

3. Pedicels 2.5-6 mm long; opercula 1.5-2 times longer than broad; new 

growth of juvenile plants dull green; margins of juvenile leaves not 

undulate; seed coats all black.E. tereticornis subsp. basaltica 


109 


Bean, Eucalyptus tereticomis 

angular, 7-20 mm long. Buds with operculum 
scar present, stamens erect to slightly sinuate, 
staminodes absent, filaments white or rarely 
pink; opercula longer than hypanthium. 
Stigma blunt or slightly dilated. Ovules 
in six or more longitudinal rows. Fruits 
cupular, oblate or hemispherical, 4-7(-8) 
mm long, 5—8(—10) mm wide, pedicellate. 
Disc convex, obliquely ascending; valves 
3-5, strongly exserted. Seeds pyramidal or 
cuboid, larger and darker than chaff, 0.8-1.5 
mm long, usually black with toothed edges, 
but sometimes yellow and with smooth edges; 
hilum terminal. 

Eucalyptus tereticomis subsp. basaltica 
A.R.Bean subsp. nov. a subspecie typica 
operculis quae sunt e basi non expansa, foliis 
juvenilibus late ovatis (late lanceolatis usque 
ovatis in subspecie typica) et habitatione 
clivorum et summorum jugorum differens. 
Typus: Queensland. Darling downs district: 
Near Bottle Tree Lookout, Bunya Mountains 
National Park, 21 May 1996, P. Grimshaw 
PG2468 & L.M. Grimshaw (holo: BRI [3 
sheets]; iso: NSW). 

Eucalyptus tereticomis ‘Hunter Valley form’ 
(Brooker & Slee 2000) 

Eucalyptus tereticomis ‘Tenterfield- 
Cunningham’s Gap form’ (Brooker & Slee 
2000) 

Eucalyptus tereticomis subsp. (Bunya 
Mountains P.V. Holzworth AQ397993), 
Bostock & Holland (2010: 121). 

Eucalyptus tereticomis subsp. (Consuelo 
Tableland M.I. Brooker B4880), Bostock & 
Holland (2010: 121). 

Tree to 20 m high. Base of trunk sometimes 
with slabs of persistent grey, scaly-fibrous 
bark, extending up to 1 metre; bark smooth 
above, deciduous, somewhat shiny and 
dappled with white, creamy-white, yellow 
and brown. Juvenile leaves ovate to almost 
orbicular, bluish green, 6.5-12.5 x 4-7 cm, 
1.2-2 times longer than broad, concolorous; 
apex acute, base cuneate; margins entire or 
crenulate, petioles 11-22 mm long. Adult 
leaves lanceolate to falcate, 90-160 x 12-32 
mm, green; apex attenuate; petioles 11-21 


mm long. Inflorescences 7-flowered or rarely 
9-flowered; peduncles ± terete or obviously 
flattened, 7-13 mm long; pedicels 2.5-6 mm 
long; mature buds ovoid to ellipsoid, 9-14 x 
5-6 mm; outer operculum shed early, inner 
operculum cylindrical-oblong to conical, 7.5- 
11 x 5-6 mm, 1.5-2 times longer than broad, 
not expanded at base, apex obtuse or acute; 
stamens white; outer stamens erect in bud, up 
to 9.5 mm long. Fruits cupular to oblate, 5-7 
mm long (excluding valves), 6-8 mm wide, 
valves 3-5, pedicels 2-7 mm long. Fig. 1A-E. 

Additional selected specimens examined : Queensland. 
Port curtis district: Kroombit Tableland, c. 60 km SW 
of Gladstone, Amy’s Peak, Jun 1977, Crisp 2824 (BRI). 
Burnett district: Foothills of Bunya Mountains S.F., Jul 
1985, Holzworth s.n. (BRI [AQ397993]); MtMowbullan, 
Bunya Mountains, Jul 1994, Fensham 1781 (BRI); Bunya 
Mountains, Mt Kiangarow, Dec 1954, Smith 6261 (BRI). 
Wide bay district: Woowoonga S.F. 287, 2 km from Mt 
Goonaneman, c. 12 km NE of Biggenden, Apr 1989, Last 
s.n. (BRI [AQ455525]); Widgee Mt, Apr 1996, Forster 
PIF19137 & Leiper (AD, BRI, CANB, MEL, NSW). 
Maranoa district: 3 km W of the ‘Stock Yard’ on the 
Consuelo Tableland, Dec 1980, Gunn BG227 (BRI); 
Consuelo Tableland, May 1982, Neldner & Thomas 683 
(BRI). Darling downs district: c. 6.6 miles [10.6 km] 
N of Yamsion on Bunya Mountains road. May 1961, 
Smith 11365 (BRI); western slope of Cunningham’s Gap, 
Jun 1957, Blake 20134 (BRI); c. 4 km along Spicer’s 
Gap road from the west, at S.F. gate, Oct 1995, Brooker 
12417 & Slee (BRI, CANB, MEL, NSW); Spicer’s Gap, 
Jun 1963, Everist 7285 (BRI); Allora Council Reserve, 
Sep 2002, Menkins 1LM0122 (BRI). Moreton district: 
Main Range, between Spring Bluff & Murphy’s Creek, 
Aug 1930, Hubbard 3530 (BRI); near Kooralbyn Resort, 
Sep 1998, Larson s.n. (BRI [AQ662937]); Mt Cotton 
ridge, Oct 2008, Bean 28106 (BRI); WNW ridge of Lost 
World, Jan 1989, Weston 1368 & Richards (BRI, NSW); 
Mt Cainbabel, Macpherson Range, May 1930, White 
6766 (BRI). New South Wales. Upper Acacia Creek, Jun 
1951, Everist 2550 & Webb (BRI); 3.8 km along Middle 
Creek Road from Cressfield Road, c. 2 km up ridge to 
west, Sep 1985, Hill 1319 (BRI, NSW); 8.6 km SW of 
Scone towards Merriwa, Oct 1995, Brooker 12350 & 
Slee (BRI, CANB); 8.3 km N of Wiseman’s Ferry, W 
edge of MacDonald River, Oct 1995, Brooker 12340 & 
Slee (BRI, CANB). 

Distribution and habitat : Eucalyptus 
tereticomis subsp. basaltica extends from 
north-west of Sydney to Kroombit Tops, south¬ 
west of Gladstone, and there is an outlier on 
the Carnarvon Range north of Mitchell (Map 
1). It is a component of eucalypt woodland, 
commonly associated with Eucalyptus 
eugenioides Sieber ex Spreng., E. melliodora 
A.Cunn. ex Schauer, and E. biturbinata 


110 


A ustrobaileya 9(1): 107-113(2013) 


m 


1 

i 

t h) 

j 

> 


Fig. 1 . Eucalyptus tereticornis subsp. basaltica A. juvenile leaf x0.5. B. adult leaf x0.6. C. cluster of mature buds x2. 
D. cluster of mature buds x2. E. cluster of mature fruits x2. E. tereticornis subsp. rotunda F. juvenile leaf x0.5. G. 
adult leaf x0.6. H. cluster of mature buds x2.1. cluster of mature buds x2. J. cluster of mature fruits x3. A,B,E from 
Grimshaw 2468 & Grimshaw (BRI); C from Bean 28106 (BRI); D from Larson s.n. (BRI [AQ662937]); F,G,I from 
Bean 31323 (BRI); H from Bean 31321 (BRI); J from Bean 25804 (BRI). Del. W.Smith. 


Bean, Eucalyptus tereticomis 

L.A.S. Johnson & K.D.Hill. It most commonly 
grows on basalt substrates, in brown to black 
clayey soil, but may sometimes grow on soils 
derived from rhyolite or granite, and rarely, 
serpentinite. Altitudes vary from 100 metres 
to 1150 metres. 

Phenology : Flowers have been collected from 
May to December; fruits may be found all 
year round. 

Notes : Eucalyptus tereticornis subsp. 

basaltica differs from E. tereticornis subsp. 
tereticornis by the opercula that are not 
expanded at the base, the broadly ovate 
juvenile leaves (broadly lanceolate to ovate 
for E. tereticornis subsp. tereticornis ), and 
the hillside or ridge top habitat. The buds 
and leaves of E. tereticornis subsp. basaltica 
are often smaller than those of the typical 
subspecies, but there is some overlap. 

This subspecies was recognised as a 
distinct taxon by Clifford (1972), who referred 
to differences in the size of the buds and 
fruits, and the shape of the juvenile leaves. 

Conservation status : A widespread and 
common subspecies. 

Etymology : The epithet refers to the basaltic 
substrate upon which this subspecies usually 
grows. 

Eucalyptus tereticornis subsp. rotunda 
A.R.Bean subsp. nov. a subspecie typica 
foliis juvenilibus orbicularibus apice obtuso 
emarginatove praeditis (lanceolato usque 
ovato apice acuto in subspecie typica) et 
operculis brevioribus quae sunt e basi non 
expansa differens. Typus: Queensland. 
Maranoa district: Western Creek, ‘Mt 
Owen’, c. 140 kmN of Mitchell, 28 November 
2006, A.R. Bean 25804 (holo: BRI; iso: 
CANB, MEL, NSW, distribuendi). 

A tree to 28 m high. Base of trunk with 
slabs of persistent grey, scaly-fibrous bark, 
extending up to 2 metres; bark smooth above, 
deciduous, somewhat shiny and dappled with 
white, creamy-white and brown. Juvenile 
leaves orbicular, 60-180 x 60-160 mm, 0.85- 
1.2 times longer than broad, concolorous or 
slightly discolorous, bright green; apex obtuse 
or emarginate, rarely apiculate, base cuneate 


ill 

or obtuse; margins entire or crenulate, petioles 
12-32 mm long. Adult leaves lanceolate to 
falcate, 90-190 x 18-32 mm, concolorous, 
green; apex attenuate; petioles 18-30 mm 
long. Inflorescences 7-flowered; peduncles 
± terete, 7-14 mm long; pedicels 2-3.5 mm 
long; mature buds ovoid to ellipsoid, 7.5-10 
x 4.5-5.5 mm; outer operculum shed early, 
inner operculum cylindrical-oblong to 
conical, 6-7 x 4.5-5.5 mm, 1.3-1.7 times 
longer than broad, apex obtuse, acute or 
umbonate; stamens white; outer stamens erect 
in bud, though slightly sinuate, up to 5.5 mm 
long. Fruits cupular to oblate, 4-5 mm long 
(excluding valves), 5.5-7 mm wide, valves 4 
or 5, pedicels 1.5-3.5 mm long. Fig. 1F-J. 

Additional specimens examined : Queensland. 

Maranoa district: Western Creek, c. 12 km N of ‘Mt 
Owen’ Homestead, N of Mitchell, Nov 2011, Bean 31321 
& Halford (BRI, CANB, MEL, NSW); ditto loc., Nov 
2011, Bean 31323 & Halford (BRI, NSW); 16.2 km from 
Consuelo Tableland towards Mt Moffatt, Dec 1982, 
Brooker 7891 (BRI, CANB). 

Distribution and habitat : Eucalyptus 
tereticornis subsp. rotunda is endemic to 
Queensland. It is known from a limited area 
north of Mitchell, on ‘Mt Owen’ Station, 
and in the Mt Moffatt section of Carnarvon 
National Park (Map 1). It grows along creek 
banks, at altitudes between 550 and 700 
metres. 

Phenology : Flowers have been recorded in 
November; fruits in November and December. 

Affinities : Eucalyptus tereticornis subsp. 
rotunda differs from E. tereticornis subsp. 
tereticornis by the orbicular juvenile leaves 
with obtuse or emarginate apex (broadly 
lanceolate to ovate juveniles with acute apex 
for E. tereticornis subsp. tereticornis ); the 
opercula of mature buds not expanded at base, 
7.5-10 mm long with obtuse apex (opercula 
of mature buds expanded at base, 11-17 mm 
long, with acute apex for E. tereticornis 
subsp. tereticornis ). 

Eucalyptus tereticornis subsp. rotunda 
differs from E. tereticornis subsp. basaltica 
by the orbicular juvenile leaves with obtuse 
or emarginate apex (broadly ovate juveniles 
with acute apex for E. tereticornis subsp. 
basaltica ), and the riverine habitat. 


112 


The juvenile leaves of Eucalyptus 
tereticornis subsp. rotunda are similar in size, 
shape and colour to those of E. amplifolia 
subsp. amplifolia , but E. tereticornis subsp. 
rotunda differs by the mature buds 7.5-10 
mm long (10-12 mm for E. amplifolia subsp. 
amplifolia ); the inflorescences 7-flowered 
(9-11-flowered for E. amplifolia subsp. 
amplifolia)., the obtuse opercula (acute for E. 
amplifolia subsp. amplifolia) and the narrower 
adult leaves. 

Notes : Eucalyptus tereticornis subsp. 

rotunda is found on the upper tributaries of 
the Maranoa River. The type specimen of 

E. camaldulensis subsp. acuta Brooker & 
M.W.McDonald was collected at ‘Forest Vale’, 
a property between Mitchell and ‘Mt Owen’, 
also on the Maranoa River. However, E. 
tereticornis subsp. rotunda and Eucalyptus, 
camaldulensis subsp. acuta are thought 
to be allopatric, with occurrences of the 
latter starting 30-40 km downstream from 
occurrences of the former. E. camaldulensis 
subsp. acuta can be distinguished by its 
bluish, lanceolate juvenile leaves, the fruiting 
valves set at a considerable angle to the disc, 
and the seeds with a yellow coat. 

Eucalyptus tereticornis subsp. rotunda is 
conspecific with the type of E. populifolia 
Hook., which is the basionym for E. populnea 

F. Muell., a name currently applied to the 
Poplar box. The typification of E. populnea is 
the subject of a current conservation proposal 
(Bean 2012). 

Conservation status : A conservation status of 
Near Threatened is proposed under Criterion 
D, i.e. area of occupancy less than 40 square 
kilometres, and fewer than 10 localities known 
(IUCN 2001). A minimum of 2000 trees have 
been observed at Mt Owen. The population 
size at Mt Moffatt is not known. 

Etymology : The subspecific epithet is from 
the Latin rotundus , meaning round. This 
refers to the shape of the juvenile leaves. 


A ustrobaileya 9(1): 107-113(2013) 

Acknowledgements 

I am grateful to Will Smith for the illustrations 
and distribution map, Peter Bostock for the 
Latin diagnoses, and an anonymous referee 
for helpful comments and suggestions. 

References 

Bean, A.R. (2012). Proposal to conserve the name 
Eucalyptus populnea (Myrtaceae) with a 
conserved type. Taxon 6\ . 1332-1333. 

Bostock, P.D. & Holland, A.E. (eds) (2010). Census 
of the Queensland Flora 2010. Queensland 
Herbarium, Department of Environment and 
Resource Management: Brisbane. 

Brooker, M.I.H. (2000). A new classification of 
the genus Eucalyptus L’Her. (Myrtaceae). 
Australian Systematic Botany 13: 79-148. 

Brooker, M.I.H. & Kleinig, D.A. (1999). Field Guide to 
Eucalypts , Volume 1 - South-eastern Australia , 
2 nd edition. Bloomings Books: Hawthorn, 
Victoria. 

Brooker, M.I.H. & Slee, A.V. (2000). Studies in the Red 
Gums of south-eastern Australia with particular 
emphasis on Eucalyptus subser. Erythroxyla. 
Australian Forestry 63: 86-106. 

Chippendale, G.M. (1988). Eucalyptus, Angophora 
(Myrtaceae). Flora of Australia Volume 19. 
Australian Government Publishing Service: 
Canberra. 

Clifford, H.T. (1972). Eucalypts of the Brisbane Region. 
Queensland Museum: Brisbane. 

Iucn (2001). IUCN Red List Categories: version 3.1. 
IUCN Species Survival Commission. IUCN: 
Gland, Switzerland. 

Nicolle, D. (2006). Eucalypts of Victoria and Tasmania. 
Bloomings Books: Melbourne. 

Queensland Herbarium (2013). Specimen label 
information (HERBRECS), viewed 22 Feb 
2013. 


Bean, Eucalyptus tereticomis 113 


Map 1 . Distribution of Eucalyptus tereticomis subsp. basaltica • and E. tereticomis subsp. rotunda A. 


A new fern species for Queensland: Diplazium 
squamuligerum (Rosenst.) Parris (Woodsiaceae) 

Daniel J. Ohlsen 1 & Ashley R. Field 2 

Summary 

D.J.Ohlsen & A.R.Field (2013). A new fern species for Queensland: Diplazium squamuligerum 
(Rosenst.) Parris (Woodsiaceae). Austrobaileya 9 ( 1 ): 114 - 125 . The taxonomic status of an unknown 
fern species from the Atherton Tableland, north-east Queensland, hitherto attributed to Asplenium 
L., was investigated. Phylogenetic analysis of trnL-F and rbcL chloroplast DNA sequences supported 
classification in Diplazium Sw., a finding supported by closer examination of scale features. 
Inspection of Diplazium type material determined that the Australian material belongs to Diplazium 
squamuligerum (Rosenst.) Parris, a species previously described from Papua New Guinea. A thorough 
description of this species and an amended key to the Diplazium species of Australia are provided. 
This study highlights the value of molecular study and close inspection of scale features for fern 
identification. Taxonomic revision in Diplazium is also discussed in light of the findings presented. 

Key Words: fern, Athyriaceae, Woodsiaceae, Diplazium , Diplazium squamuligerum , Australia flora, 
Queensland flora. Wet Tropics bioregion, new species record, taxonomy 

'D.J.Ohlsen, School of Botany, The University of Melbourne, Parkville, Victoria 3010, Australia. 
Email: d.ohlsen@student.unimelb.edu.au 

2 A.R.Field, Queensland Herbarium, Department of Science, Information Technology, Innovation and 
the Arts, Brisbane Botanic Gardens, Toowong, Queensland 4066, Australia &Australian Tropical 
Herbarium, James Cook University, Cairns Campus, Smithfield, Queensland 4878, Australia. Email: 
A shley. F ield@science. dsitia. qld. gov. au 


Introduction 

The Wet Tropics of north-east Queensland, 
from Ingham north to Cooktown, contains 
the highest fern species diversity in Australia. 
257 leptosporangiate fern species occur in 
the region, which is 66% of the total for 
Australia (McCarthy 1998). While 37 of these 
are considered to be endemic to this region, 
the vast majority are widespread being 
shared with neighbouring tropical islands, 
particularly New Guinea which shares 56% 
of tropical Australian leptosporangiate fern 
species (McCarthy 1998). Several such species 
are extremely rare in Australia, known only 
from one or a few small populations. Notable 
examples ar q AspleniumpeUucidiim Lam. and 
Hymenasplenium unilaterale (Lam.) Hayata, 
both restricted to a single known population 
in Australia (Brownsey 1998). 

The rarity of some taxa has delayed their 
discovery until relatively recently in Australia. 
Lecanopteris sinuosa (Wall, ex Hook.) Copel., 


Accepted for publication 29 July 2013 


known from only three sites in Australia, was 
described (as Polypodium sinuosa Wall, ex 
Hook.) from Indonesia in 1863, but was not 
collected in Australia until 1985 when it was 
found on Moa Island, Torres Strait. It was 
later collected on Cape York in 1987, and was 
formally recognised as an Australian species 
(Andrews 1990). Entirely new taxa, endemic 
to north-east Queensland have also gone 
unnoticed until relatively recently. These 
include Antrophyum jagoanum D.L.Jones 
& Bostock, Diplazium bostockii D.L.Jones, 
and Lastreopsis windsorense D.L. Jones 
(McCarthy 1998). However, the most recent 
new fern species confirmed for Australia is 
Oleandra musifolium Blume. In contrast to 
the previous cases, populations of this species 
have been known in Australia for over 60 
years, but under the misapplied name of O. 
neriiformis Cav. (Hovenkamp & Ho 2012). 

Four fern specimens collected from two 
areas on the Atherton tableland, in north-east 
Queensland, hitherto identified as Asplenium 
sp. indet. (A. sp. ‘RE.I. Road’ in BRI) 


115 


Ohlsen & Field, Diplazium squamuligerum 

(Aspleniaceae) on herbarium specimens, also 
appear to represent a new taxon for Australia 
(Fig. 1). This taxon is clearly morphologically 
distinct from the 30 Australian Asplenium 
species treated by Brownsey (1998). It was 
first collected in 1983: once from the North 
Johnstone Logging Area ( Lockyer s.n.), which 
is thought to be from a population on the west 
side of Topaz National Park (N.P.) (B.Gray 
pers. comm.), and twice from Maalan, 20 
km further south. The fourth and most recent 
collection, in 2005 ( Sankowsky & Sankowsky 
2637), was from near the end of P.E.I. Road 
in Topaz N.P, within one km of the suspected 
location of the Lockyer collection. 

Determining the taxonomic status of 
taxa in Aspleniaceae is often fraught with 
difficulty. Aspleniaceae is the largest fern 
family with over 700 species occurring 
worldwide (Kramer & Viane 1990; Smith 
et al. 2006). This makes verification of 
identity or novelty challenging for unresolved 
Australian collections; the putative taxon 
must be compared with a large number of 
type specimens from overseas. In addition to 
the large species diversity in Aspleniaceae, 
hybridisation and polyploidy are frequently 
encountered (see Lovis 1977). Hybrids and 
polyploids are often morphologically distinct 
from parental lineages (Kramer & Viane 
1990), obscuring their origins and giving the 
impression of completely new lineages. 

While identification within Aspleniaceae 
is often difficult, members of the family 
are generally easy to distinguish from 
other families. Those in Aspleniaceae are 
distinguished by linear sori, clathrate scales 
on the stipes, and stipes which have two 
C-shaped vascular bundles at the base which 
unite apically up the stipe to form a ‘butterfly- 
like’ shape (Kato & Kramer 1990). Species 
of Aspleniaceae are morphologically most 
similar to some members of the Woodsiaceae 
sensu Smith et al. (2006) (=Athyriaceae 
sensu McCarthy (1998)), which also possess 
elongate indusiate sori. The majority of 
these Woodsiaceae species are most easily 
distinguished from Aspleniaceae by being 
large terrestrial plants; however, they are 
consistently distinguished by more subtle 


features such as differences in the arrangement 
of vascular bundles in the stipe, and by the 
possession of non-clathrate scales (Kato & 
Kramer 1990), which are often only obscurely 
differentiated from the clathrate scales 
observed in Aspleniaceae. Two Woodsiaceae 
genera occur in north-east Queensland: 
Deparia Hook. & Grev. and Diplazium Sw. 
sensu lato (including Callipteris Bory). 

This study sequenced two chloroplast 
regions, rbcL and trnL-L, of a fresh collection 
of Asplenium sp. ‘RE.I. Road’, in order to 
determine this taxon’s affinity amongst the 
ferns, and to resolve its taxonomic status. 

Materials and methods 

Collections of Asplenium and Diplazium 
in Australian herbaria, and selected type 
material from extra-Australian herbaria were 
examined. One specimen of Asplenium sp. 
‘P.E.I. Road’ was collected (i Ohlsen 461, BRI, 
MELU) in the vicinity of the most recent P.E.I. 
Road collection {Sankowsky & Sankowsky 
2637, BRI). 

DNA isolation, amplification and 
sequencing. 

DNA was extracted from 20 mg of silica gel- 
dried leaf tissue. Leaf tissue was ground using 
a mortar and pestle with the aid of acid washed 
grinding sand (Ajax Finechem, Australia). 
DNA was isolated from ground samples 
using a DNeasy Plant Mini Kit (QIAGEN, 
Germany), following the manufacturer’s 
instructions. DNA was eluted in 100 pL of the 
supplied elution buffer. 

The chloroplast DNA markers rbcL and 
trnL-L were sequenced. These regions were 
chosen because 1) both are routinely used 
in fern systematics, enabling comparison 
of sequences obtained from this species to 
those of many other species, and 2) effective 
phylogenetic placement (at least to genus 
level) can be achieved with both regions. 
rbcL is a gene enabling comparison across 
multiple plant lineages, and the trnL-L region 
contains an intron and intergenic spacer, 
which have faster mutation rates, enabling 
differentiation between closely related species 
and populations. 


116 


Austrobaileya 9(1): 114-125(2013) 


Fig. 1. Diplazium squamuligerum (syn. Asplenium sp. RE.I. Road) from Topaz National Park, Atherton Tableland, 
Queensland (Ohlsen 461 et al. [BRI, MELU]). (a) a mature frond with elongate sori, (b) a fully developed scale from 
the base of the stipe, (c) D. squamuligerum in habitat (two plants are shown: top left, and bottom beside the creek 
pool), (d) detail of the rachis and stipe wing, and the protuberances, some with scales developing upon them. 


Chloroplast DNA markers were 
amplified by Polymerase Chain Reaction 
(PCR), performed on a MyCycler thermal 
cycler (Bio-Rad, USA). Reaction mixtures 
comprised 5 pL of 5x MyTaq Reaction Buffer 
containing 5mM of each dNTP and 15 mM 
MgCl 2 (Bioline, Australia), 50 pg BSA 
(Thermo Fisher Scientific, Australia), 0.125 
pL (0.625 Units) MyTaq DNA Polymerase 
(Bioline, Australia), 10 pmol of each primer, 
2.0 pL of extracted DNA, and distilled water 
added to make a total volume of 25 pL. No 
amplification occurred without the addition 
of BSA. The rbcL gene was amplified using 
the primers ESRBCL1F (5’-ATG TCA CCA 
CAA ACG GAG ACT AAA GC-3’) and 
ESRBCL1361R (5’-TCA GGA CTC CAC 


TTA CTA GCT TCA CG-3’) (Schuettpelz & 
Pryer 2007). The trnL intron, trnL 3’-exon 
and trnL-F intergenic spacer were amplified 
using the primers F (5’-ATT TGA ACT GGT 
GAC ACG AG-3’) (Taberlet et al. 1991) and 
Fernl (5’-GGC AGC CCC CAR ATT CAG 
GGR AAC C-3’) (Trewick et al. 2002). PCR 
thermocycling conditions involved an initial 
denaturation step of 95°C for 1 minute, 
followed by 33 cycles of 95°C for 1 minute, 
55°C for 1 minute, and 65°C for 4 minutes, 
and a final extension time of 65°C for 5 
minutes. DNA concentrations were quantified 
by electrophoresis against Hyperladder I 
and EasyLadder I (Bioline, Australia) and 
PCR products were purified using illustra 
ExoSTAR 1-step enzymatic purification (GE 


117 


Ohlsen & Field, Diplazium squamuligerum 

Healthcare Life Sciences, UK). Purified PCR 
products were then sent to the Australian 
Genome Research Facility (AGRF), 
Melbourne Branch, where sequencing 
reactions, and capillary separation, using the 
96-capillary analyser AB 3730x/ sequencing 
platform, were performed. 

Sequence editing, alignment, and analysis. 
Sequences were edited using Sequencher 
v. 3.0 (Gene Codes Corporation, Ann 
Arbor, MI, USA) and the NCBI nucleotide- 
nucleotide BLAST (blastn) tool (Altschul 
et al. 1997) was then used to evaluate the 
similarity of both rbcL and trnL-L sequences 
to existing sequences in GenBank. After the 
search using the BLAST tool was performed, 
available rbcL and trnL-L sequences of 
Diplazium species from GenBank were 
gathered (Appendix 1) and aligned manually 
in Se-Al Sequence Alignment Editor v. 2.0all 
(Rambaut 2002). Outgroup sequences were 
also gathered from GenBank (Appendix 1). 

Parsimony analyses were run in PAUP* 
v 4.0(310 (Swofford 2000). Gap characters 
in the alignment were treated as a fifth 
character state. For multiple base indels, 
characters were excluded from analyses so 
that indels were represented only by a single 
gap character, when variability did not occur 
within indels. Question marks, the character 
recognised in PAUP for missing data, were 
used to fill gaps, where needed, when indels 
fell across otherwise variable regions of the 
alignment. A heuristic tree search was used, 
with delayed character-state optimization 
(DELTRAN) and starting trees obtained 
by a closest addition sequence followed by 
tree bisection-reconnection (TBR) branch 
swapping. Bootstrap support for tree nodes 
was determined for each analysis, with 100 
replicates. Separate parsimony analyses were 
performed for each chloroplast DNA region. 
Athyrium felix-femina (L.) Roth was chosen as 
an outgroup, based on the sister relationship 
of Athyrium to Diplazium in past molecular 
phylogenies (Sano et al. 2000; Wang et al. 
2003; Schuettpelz & Pryer 2007) and for 
the rbcL analysis, in which it is possible 
to align distantly related taxa, the more 
distantly related Blechnum occidental L. and 


Asplenium marinum L. were also included as 
outgroups. 

Species identification. 

Online images of Diplazium types (B, 
MICH) were examined, to determine whether 
this taxon could be assigned to an existing 
Diplazium species, or if it was an undescribed 
species. 

Common Abbreviations used in Specimen 
Citations 

LA (Logging Area); NP (National Park). 

Results 

Searches using the BLAST tool of both 
trnL-L and rbcL sequences, determined that 
sequences from the P.E.I. Road specimen had 
the greatest similarity to species of Diplazium 
in the family Woodsiaceae. 37 Diplazium 
rbcL sequences were used to produce an 
alignment 1188 base pairs long, which 
contained 110 parsimonious characters. 
Two most parsimonious trees were obtained 
(length=398 steps, CI= 0.626) one of which is 
shown (Fig. 2a). This had a topology identical 
to the strict consensus tree, except for the 
placement of D. doderleinii (Luerss.) Mak., 
which in the consensus tree is placed in a 
polytomy with the clade of D. amamianum 
Tagawa + D. hachijoense Nakai and the clade 
of D. taiwanense Tagawa + D. virescens 
Kunze. 10 Diplazium trnL-L sequences were 
used to produce an alignment of 753 included 
base pairs, which contained 139 parsimonious 
characters. A single most parsimonious tree 
was obtained (length=302 steps, 0=0.831) 
(Fig. 2b). Maximum parsimony analyses of 
rbcL (Fig. 2a) and trnL-L (Fig. 2b) shows that 
the P.E.I. Road taxon is nested well within 
Diplazium sensu lato. Of all Diplazium 
species currently sequenced for rbcL, it is 
most closely related to D. proliferum (Lam.) 
Thouars. ( =Callipteris prolifera (Lam.) Bory 
sensu Jones [1998]) with high bootstrap 
support (98%). It differs from D. proliferum 
by 12 base pairs in rbcL. 

Inspection of Diplazium type material 
revealed that the P.E.I. Road taxon matches 
Asplenium varians var. squamuligerum 
Rosenst., from New Guinea, now treated as 


118 


15 


70 


73 


6 


1 


93 


99 

2 


Austrobaileya 9(1): 114—125(2013) 

I Diplazium amamianum 
' Diplazium hachijoense 


— Diplazium doederleinii 
Diplazium taiwanense 

- Diplazium virescens var. conterminum 

- Diplazium virescens 
2 

|— Diplazium crassiusculum 
Diplazium donianum var. aphanoneuron 


99 

1 


1 


a 


87 


1 


91 


^ Diplazium lobatum 
Diplazium dilatatum 
Diplazium hayatamae 
Diplazium deciduum 
Diplazium fauriei 
U- Diplazium mettenianum 
Diplazium griffithii 
— Diplazium pullingeri 


L-=— Diplazium kawakamii 


8 


76 


- Diplazium centripetale 

—— Diplazium incomptum 
4 

9 |- Diplazium longicarpum 

- 12 

97 1 --- Diplazium subserratum 


Diplazium proliferum 


98 

p 

5 Diplazium chinense 

L_6 


Diplazium squamuligerum 


87 

4 


8 


10 


Diplazium subtripinnatum 
■ Diplazium esculentum 
— Diplazium nipponicum 


■ Diplazium sibiricum var. glabrum 


Diplazium squamigerum 

9 Diplazium bombanasae 


95 


93 


— Diplazium cristatum 
Diplazium lonchophyllum 
- Diplazium plantaginifolium 
13 | Diplazium okudairae 

' Diplazium wichurae 


18 


43 


—— Diplazium pinfaense 
- Athyrium felix-femina 

- Blechnum occidentale 

- Asplenium marinum 


(a) 


Fig.2a. Parsimony analyses of rbcL showing the phylogenetic position of Australian Diplazium squamuligerum. 
Branch lengths are given above braches and bootstrap support values greater than 70% are given below branches. 


Ohlsen & Field, Diplazium squamuligerum 


119 


24 

100 


12 


■ Diplazium pinfaense 


r^- Diplazium wichurae 


11 


100 


L^- Diplazium heterocarpum 


7 

99 


Diplazium hainanense 


Diplazium hachijoense 


7 

89 


16 


15 


Diplazium esculentum 


Diplazium nipponicum 


46 


15. 

90 


17 


Diplazium mettenianum 


Diplazium squamuligerum 


59 


Diplazium ovatum 


Athyrium filix-femina 

(b) 


Fig.2b. Parsimony analyses of trnL-F. Branch lengths are given above braches and bootstrap support values greater 
than 70% are given below branches. 


120 

Diplazium squamuligerum (Rosenst.) Parris. 
The original diagnosis treated it as a variety 
of the unrelated African Asplenium varians 
Wall, ex Hook. & Grev. (Rosenstock 1913). 
The diagnosis is very brief, outlining only 
some immediately obvious features which 
distinguish D. squamuligerum from A. 
varians. Many of the morphological features 
that are quite distinctive are not mentioned. 
The following description describes 
Diplazium squamuligerum in greater 
detail, including several distinctive features 
overlooked by the earlier diagnosis. 

Taxonomy 

Diplazium squamuligerum (Rosenst.) 
Parris, Kew Bull. 41 (1): 69 (1986); Asplenium 
varians var. squamuligerum Rosenst., Repert. 
Spec. Nov. Regni Veg. 12: 528 (1913); A. 
squamuligerum (Rosenst.) Hieron, Hedwigia 
61:5 (1919).Type: PapuaNew Guinea: Morobe 
Province: Sattelberg Hinterland, 1400-1500 
m [I.C.]Keysser 228, April 1913 (holo: S, S-P- 
7625, online image!; iso: UC 378428, online 
image!; MICH 1190092, online image!). 

Rhizome erect, to 1.5 cm tall, scaly; scales to 
1 mm long, dark brown with toothed margins. 
Fronds arcuate, 5-15 cm long, 1.5-4 cm wide. 
Stipe 0.5-5 cm long, dark green, enveloped by 
green tissue which elongates to produce wings 


Austrobaileya 9(1): 114—125(2013) 

and protuberances; protuberances eventually 
bearing light brown scales to 1 mm long, 
with darker brown veins and bifid marginal 
teeth. Lamina bipinnate-tripinnatifid, 3-10 
cm long, 1.5-4 cm wide, dark green above, 
paler below, occasionally proliferous from the 
rachis. Primary pinnae 5-25 mm long, 2-10 
mm wide, most with an acroscopic secondary 
pinna to 5 mm long and an elliptic apical 
segment 5-10 mm long; margins serrate; 
veins free. Sori to 4 mm, elongate along most 
veins; indusium entire, narrow, light brown 
(Fig. la, b, d) 

Specimens examined: Australia: Queensland. Cook 
district: North Johnstone LA, July 1983, Lockyer s.n. 
(CANB); P.E.I. Road, Topaz NP, July 2005, Sankowsky 
& Sankowsky 2637 (BRI); North Johnstone LA, Jan 
2013, Ohlsen 461 etal. (BRI, MELU); Portion 545 Parish 
of Dirran, Malian Road, July 1983, Gray 3133 (CNS). 

Distribution and habitat : In Australia 
Diplazium squamuligerum is known only 
from two areas on the Atherton Tableland, 
north-east Queensland; however, it also 
occurs in Papua New Guinea. It grows in 
mixed mesophyll rainforest, on metamorphic 
rocks or between tree roots in volcanic soil, 
in steep embankments of small, slow-flowing 
creeks (Fig. lc). 

Notes : Wings of the axes often become 
obscure when dried, but are obvious in fresh 
material. 


Key to Australian species of Diplazium (naturalised taxa indicated *) 


1 Veins anastomosing.2 

1 . Veins free. 5 

2 Lamina simple.D. cordifolium 

2. Lamina pinnate to tripinnate.3 

3 Lamina pinnate, rachis proliferous.D. proliferum (syn. Callipteris prolifera) 

3. Lamina bipinnate or tripinnate, rachis not proliferous. 4 

4 Secondary pinnae 1-1.5 cm wide, margins incised more than half-way to 

the pinnule midvein.D. dietrichianum 

4 . Secondary pinnae 1.5-2.5 cm wide, very shallowly lobed.*D. esculentum 

5 Lamina pinnate.6 

5. Lamina bipinnate or more divided. 7 

6 Lamina apex a single pinna similar to lateral pinnae.D. pallidum 

6. Lamina apex formed by reduction of lateral pinnae.D. dameriae 


121 


Ohlsen & Field, Diplazium squamuligerum 

7 Fertile laminae < 250 mm long, stipe and rachises winged, wing extending 
to form scale bearing protuberances, pinnae with a basal acroscopic 


secondary pinna or lobe. D. squamuligerum 

7. Fertile Laminae > 250 mm, stipe, rachis and pinnae not as above.8 

8 Lamina bipinnate. 9 

8. Lamina tripinnate or more divided. 10 

9 Basal lobes of secondary pinnae longest, abaxial surface of pinnae 

glabrous. D. dietrichianum 

9. Basal lobes of secondary pinnae similar to or smaller than the rest, abaxial 

surface bearing red glandular hairs. D. dilatatum 


10 Lamina pale green, pinnules less than 5 mm long and 3 mm wide. D. assimile 

10. Lamina dark green, pinnules greater than 10 mm long and 4 mm wide. 11 

11 Apex of pinnules acute to caudate. D. bostockii 

11. Apex of pinnules obtuse. 12 

12 Pinnules of fertile lamina dissected < one third of the distance to the 

midvein, pinnules decurrent on basiscopic margin. D. australe 

12. Pinnules of fertile lamina dissected > half-way to the midvein, pinnules 

not decurrent. D. queenslandicum 


Discussion 

The identification of the PE.I. Road taxon as 
a species of Diplazium by chloroplast DNA 
sequence is also supported by morphology. 
Populations of D. squamuligerum in Australia 
have elongate sori and scales that are borne 
on small protuberances: both features typical 
of Diplazium (Jones 1998). The scales are 
characterised by bifid teeth on a darkened 
margin and conform to the ‘Callipteris type’ 
(Fig. lb). This scale type is known only from 
the segregate genus Callipteris and a few 
other Diplazium species (Pachebo & Moran 
1999; Pachebo & Moran 2003). 

These specimens were repeatedly 
misidentified as a species of Asplenium, 
rather than being correctly identified as a 
Diplazium. Hence this study is another case 
which highlights the value of DNA sequences 
for phylogenetic placement and identification 
when morphology may be misleading 
(Gastony et al. 2001). It also begs caution in 
identifying species by overall morphology 
without close inspection of more subtle but 
taxonomically informative features such as 
scale characters. 


Prior to this study, 12 species of Diplazium 
were known in Australia (one species, D. 
esculentum is naturalised). All of these 
Diplazium species are large terrestrials, with 
fronds over 50 cm long. The small size of D. 
squamuligerum is therefore unique amongst 
Australian Diplazium , and unusual amongst 
all Diplazium (Kato & Kramer 1990). Its 
position in the chloroplast phylogeny, as sister 
to D. proliferum is supported by its possession 
of the ‘Callipteris type’ scales, that are also 
possessed by D. proliferum (Jones 1998). 
Both D. squamuligerum and D. proliferum 
are also proliferous from the rachis, albeit 
rarely so in D. squamuligerum. Otherwise 
these species are quite dissimilar in general 
appearance. D. proliferum is a very large fern 
with pinnate fronds to over 2 metres in length 
whereas the fronds of D. squamuligerum are 
at least 2-pinnate and less than 15 cm long. 
D. squamuligerum also has free venation and 
winged axes (Fig. Id); the latter feature is not 
shared with any other Australian Diplazium 
species. 

The current phylogeny of Diplazium is 
limited in the number of species sampled. 
Chloroplast DNA sequences of only 38 
species are publicly available, all of which 


122 

are incorporated in the phylogenies presented 
here. This is a low proportion of the 400 species 
of Diplazium in total (Kato & Kramer 1990). 
So it is highly likely that another unsampled 
species of Diplazium is more closely related 
to D. squamuligerum than D. proliferum, 
especially given the large number of base 
pair differences between the two species. 
Several Diplazium species, which occur in 
New Guinea, or possess the ‘Callipteris type’ 
scales, are yet to be sequenced, and may be 
more closely related. 

Diplazium has received little attention 
from molecular study, as is demonstrated by 
the low number of sequences available on 
GenBank. Further molecular work, involving 
a more comprehensive sampling of species, 
is required in Diplazium not only to gain 
a better understanding of the systematics 
and biogeography of this genus, but also to 
aid taxonomic revision, particularly at the 
generic level. It has been suggested, based on 
a recent molecular study using a rather small 
number of species (Wang et al. 2003), that 
several genera which had been segregated 
from Diplazium be placed back into that 
genus. This included Callipteris , a genus still 
accepted in Australia (Jones 1998). However, 
revision of Diplazium cannot be completed 
until more species sampling is undertaken. 
The close relationship of D. squamuligerum 
to D. proliferum suggests that the ‘Callipteris 
type’ scale may define a monophyletic group. 
It has been suggested that this feature may 
be used to recircumscribe Callipteris , to 
include the few Diplazium species, such as 
D. squamuligerum , that have this scale type, 
and also have free venation (Pachebo & 
Moran 2003). However, the monophyly of 
all species possessing this scale type needs 
to be demonstrated before any revision is 
undertaken (Pachebo & Moran 2003). 

Identification of this species as D. 
squamuligerum provides an additional 
example of a fern species that is localised 
in a few sites on the Atherton Tableland. 
Populations at Topaz NP are within close 
proximity to other localised fern species. They 
are within 3 km of the only known locality of 
Hymenasplenium unilateral in Australia, and 


Austrobaileya 9(1): 114—125(2013) 

adjacent to one of approximately ten locations 
of Dryopteris wattsii M.McKeown, Sundue & 
Barrington. In addition to these exceptionally 
rare species, other uncommon fern species 
such as Dicksonia herbertii W.Hill, Oleandra 
musifolia , and Pteridoblechnum acuminatum 
(C.T.White & Goy) Hennipman and P. 
neglectum (F.M.Bailey) Hennipman also occur 
at or near the D. squamuligerum population 
localities. Likewise, the Maalan population is 
within 7 km of one of only three populations 
of Asplenium normale D.Don in Australia. 
The occurrences of multiple, very localised 
fern species near sites of D. squamuligerum 
highlights the high conservation priority that 
should be given to these areas, especially 
considering the high diversity of seed plant 
species also present. It also suggests that a 
number of equally localised species may have 
been lost due to vegetation clearing around 
these populations. 

Diplazium squamuligerum is currently 
one of the most threatened fern species in 
Australia. While it is reserved in Topaz NP, 
all populations of this species are small in 
area and size and are particularly vulnerable 
to stochastic events. Periods of particularly 
high rainfall, with increased runoff due to 
surrounding cleared land, can swell the small 
creeks along which it occurs. Consequently 
plants face the danger of being uprooted from 
the creek embankments. This species would 
also be extremely susceptible to desiccation. 
It occurs in very high rainfall areas, and along 
small slow flowing creeks, that provide the 
constant soil moisture and high humidity that 
is probably required by the plant. Extended 
dry periods may reduce such small creeks and 
limit water access to plants. Such stochastic 
weather events are expected to become more 
extreme in the future with ongoing climate 
change (Abbs et al. 2006; Mpelasoka et al. 
2008; Walsh et al. 2004). 

The one collection from Papua New 
Guinea that exists in Australian herbaria 
(Brass 25731 [CANB]) was collected from 
Mt Pabinama, Normanby Island, which is 
550 km south-east from the type locality, 
and suggests that D. squamuligerum is 
more widespread in Papua New Guinea. 


123 


Ohlsen & Field, Diplazium squamuligerum 

A conservation status of Vulnerable is 
recommended for D. squamuligerum 
in accordance with the distribution and 
abundance characteristics of other taxa listed 
as Vulnerable under the Nature Conservation 
Act 1992 (Queensland), the Environment 
Protection and Biodiversity Conservation Act 
1999 (Australian Commonwealth) and also 
in line with the criteria of the International 
Union for Conservation of Nature (IUCN). A 
conservation advice report is currently being 
prepared for D. squamuligerum in Australia. 

Acknowledgements 

We would like to express our gratitude 
to Bruce Gray, who assisted the recent 
collection of Diplazium squamuligerum. All 
sequencing was performed in the Cookson 
Laboratory, School of Botany, The University 
of Melbourne, and was funded by a BushBlitz 
Research Grant. Much appreciated advice in 
sequencing D. squamuligerum was provided 
by Lara Shepherd and Leon Perrie. We are also 
thankful to Leon Perrie, Nada Sankowsky, 
Darren Crayn, and to the anonymous 
reviewers for their helpful comments on the 
manuscript. 

References 

Abbs, D., Aryal, S., Campbell, E., McGregor, 
J., Nguyen, K., Palmer, M., Rafter, T., 
Watterson, I. & Bates, B. (2006). Projections 
of extreme rainfall and cyclones. CSIRO, Final 
report to the Australian Greenhouse Office. 

Andrews, S.B. (1990). Ferns of Queensland. A 
handbook to the ferns and fern allies , 402-404. 
Queensland Department of Primary Industries: 
Brisbane, Australia. 

Altschul, S.F., Madden, T.L., Schaffer, A.A., Zhang, 
J., Zhang, Z., Miller, W. & Lipman, D. J. (1997). 
Gapped blast and psi-blast: a new generation 
of protein database search programs. Nucleic 
Acids Research 25: 3389-3402. 

Bell, G.H. (1998). Davalliaceae. In PM. McCarthy 
(ed.), Flora of Australia Ferns, Gymnosperms 
and Allied Groups 48: 434-450. ABRS/CSIRO: 
Melbourne. 

Brownsey, P.J. (1998). Aspleniaceae. In PM. McCarthy 
(ed.). Flora of Australia Ferns, Gymnosperms 
and Allied Groups 48: 295-327. ABRS/CSIRO: 
Melbourne. 


Gastony, G.J. & Johnson, W.P (2001). Phylogenetic 
placements of Loxoscaphe thecifera 
(Aspleniaceae) and Actiniopteris radiata 
(Pteridaceae) based on analysis of rbcL 
nucleotide sequences. American Fern Journal 
91: 197-213. 

Hovenkamp, P.H. & Ho, B-C. (2012). A revision of the 
fern genus Oleandra (Oleandraceae) in Asia. 
PhytoKeys 11: 1-37. 

Jones, D.L. (1998). Athyriaceae. In P.M. McCarthy 
(ed.). Flora of Australia Ferns, Gymnosperms 
and Allied Groups 48: 418-429. ABRS/CSIRO: 
Melbourne. 

Kato, M. & Kramer, K.U. (1990). Subfamily 
Athyrioideae. In K.U. Kramer & PS. Green 
(eds.). The families and genera of vascular 
plants Pteridophytes and gymnosperms 1: 130- 
144. Springer-Verlag: Berlin. 

Kramer, K.U. & Viane, R. (1990). Aspleniaceae. In 
K.U. Kramer & PS. Green (eds.). The families 
and genera of vascular plants Pteridophytes 
and gymnosperms 1: 52-56. Springer-Verlag: 
Berlin. 

Lovis, J.D. (1977). Evolutionary patterns and processes 
in ferns. Advances in Botanical Research 4: 
223-415. 

McCarthy, PM. (1998). Flora of Australia Ferns, 
Gymnosperms and Allied Groups 48. ABRS/ 
CSIRO: Melbourne. 

Mpelasoka, F., Hennessy, K., Jones, R. & Bates, B. 

(2008). Comparison of suitable drought indices 
for climate change impacts assessment over 
Australia towards resource management. 
International Journal of Climatology 28: 1283— 
1292. 

Pachebo, L. & Moran, R.C. (1999). Monograph of 
the neotropical species of Callipteris with 
anastomosing veins (Woodsiaceae). Brittonia 
51: 343-388. 

- (2003). Lectotypification of several names 

currently placed in Diplazium (Woodsiaceae). 
American Fern Journal 93: 90-92. 

Parris, B.S. (1986). New combinations in filices. Kew 
Bulletin 41: 69-70. 

Rambaut, A. (2002). ‘Se-Al: sequence alignment editor’. 
Available at: http://tree.bio.ed.ac.uk/ 

Rosenstock, E. (1913). Filices novoguineenses 
Keysseranae, III. Repertorium Specierum 
Nov arum Regni Vegetabilis 12: 524-530. 

Sano, R., Takamiya, M., Ito, M., Kurita, S. & Hasebe, 

M. (2000). Phylogeny of the lady fern group, 
tribe Physematieae (Dryopteridaceae), based 
on chloroplast gene sequences. Molecular 
Phylogenetics & Evolution 15: 403-413. 


124 

Schuettpelz, E. & Pryer, K.M. (2007). Fern phylogeny 
inferred from 400 leptosporangiate species and 
three plastid genes. Taxon 56: 1037-1050. 

Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, 
P, Schneider, H. & Wolf, PG. (2006). A 
classification for extant ferns. Taxon 55: 705- 
731. 

Swofford, D.L. (2000). PAUP* 4.0: phylogenetic 
analysis using parsimony (*and other methods). 
Sinauer Associates: Sunderland, MA. 

Taberlet, P, Gielly, L., Pautou, G. & Bouvet, J. (1991). 
Universal primers for amplification of three 
non-coding regions of chloroplast DNA. Plant 
Molecular Biology 17: 1105-1109. 


Austrobaileya 9(1): 114—125(2013) 

Trewick, S.A., Morgan-Richards, M., Russell, 
S.J., Henderson, S., Rumsey, F.J., Pinter, I., 
Barrett, J.A., Gibby, M. & Vogel, J.C. (2002). 
Polyploidy, phylogeography and Pleistocene 
refugia of the rockfern Asplenium ceterach: 
evidence from chloroplast DNA. Molecular 
Ecology 11: 2003-2012. 

Walsh, K. J.E., Nguyen, K.C. & McGregor, J.L. (2004). 
Finer-resolution regional climate model 
simulations of the impact of climate change 
on tropical cyclones near Australia. Climate 
Dynamics 22: 47-56. 

Wang, M-L., Chen, Z-D., Zhang, X-C., Lu, S-G. & Zhao, 
G-F. (2003). Phylogeny of the Athyriaceae: 
evidence from chloroplast trnL-F region 
sequences. Acta Phytotaxonomica Sinica 41: 
416-426. 


Appendix 1. Chloroplast DNA sequences included in analyses, identified by GenBank 
accession numbers 


Taxon 

rbcL 

trnL 

Asplenium marinum L. 

AF240647 

- 

Athyrium felix-femina (L.) Roth 

FJ821348 

HQ676519 

Blechnum occidentals L. 

U05909 

- 

Diplazium amamianum Tagawa 

AB574961 

- 

D. bombanasae Rosenst. 

EF463308 

- 

D. centripetale (Bak.) Maxon 

EF463309 

- 

D. chinense (Bak.) C.Chr. 

AB574962 

- 

D. crassiusculum Ching 

AB574963 

- 

D. cristatum (Desr.) Alston 

EF463310 

- 

D. deciduum N.Ohta & M.Takamiya 

AB574964 

- 

D. dilatatum Blume 

EF463311 

- 

D. doederleinii (Luerss.) Mak. 

AB574966 

- 

D. donianum (Mett.) Tard var. aphanoneuron (Ohwi) Tagawa 

AB574968 

- 

D. esculentum (Retz.) Sw. 

U05619 

AF514838 

D. fauriei Christ 

AB574970 

- 

D. griffithii (T.Moore) Diels 

AB574971 

- 

D. hachijoense Nakai 

EF463312 

EU329111 

D. hainanense Ching 

- 

AF515257 


Ohlsen & Field, Diplazium squamuligerum 


125 


Taxon 

rbcL 

trnL 

D. hayatamae N.Ohta & M.Takamiya 

AB574973 

- 

D. heterocarpum Ching 

- 

- 

D. incomptum Tagawa 

AB574974 

- 

D. kawakamii Hayata 

AB574975 

- 

D. lobatum (Tagawa) Tagawa 

AB574976 

- 

D. lonchophyUum Kunze 

U05920 

- 

D. longicarpum Kodama 

AB574977 

- 

D. mettenianum (Miq.) C.Chr. 

AB574978 

AF515233 

D. nipponicum Tagawa 

AB574979 

EU329110 

D. okudairae Mak. 

AB574980 

- 

D. ovatum Wall. 

- 

AF515232 

D. pinfaense Ching 

AB574981 

AF515250 

D. plantaginifolium (L.) Urb. 

EF463314 

- 

D. proliferum (Lam.) Thouars 

EF463918 

- 

D. pullingeri (Bak.) J.Sm. 

AB574982 

- 

D. sibiricum (Turcz ex Kunze) Kurata in Namegata 
var. glabrum (Tagawa) Sa.Kurata 

AB574983 

- 

D. squamigerum (Mett.) Christ 

AB574984 

- 

D. squamuligerum (Rosenst.) Parris 

KC900228 

KC900229 

D. subserratum (Blume) T.Moore 

AB021721 

- 

D. subtripinnatum Nakai 

AB574985 

- 

D. taiwanense Tagawa 

AB574986 

- 

D. virescens Kunze var. conterminum (Christ.) Sa.Kurata 

JN168017 

- 

D. virescens Kunze 

EF463316 

- 

D. wichurae (Mett.) Diels 

JN168018 

AF515245 


Austrobaileya 9(1): 126-129 (2013) 


126 


SHORT COMMUNICATION 

New records for Queensland in Lindernia All. (Linderniaceae) 

B.S. Waiman 

Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor Road Smithfield, 
Queensland 4878, Australia. Email: bswannan@bigpond.com 


The genus Lindernia All. (Linderniaceae 
Borsch, K.Mull. & Eb.Fisch.) is a pan- 
tropical genus represented in Australia by 
approximately 50 species. In Australia, many 
of the species are yet to be formally named, 
especially those from the Northern Territory. 

During the current taxonomic revision of 
Lindernia in Queensland (Wannan in prep.) 
two new species have been recorded, namely: 
Lindernia pusilla (Willd.) Bold., which is a 
new record for Australia and L. tectanthera 
W.R.Barker, which is a new record for 
Queensland. 

Lindernia pusilla (Willd.) Bold., Zakfl. Java 
165 (1916); Gratiolapusilla Willd., Sp. Pl. ed. 
4 , 1(1): 105 (1797); Pyxidaria pusilla (Willd.) 
Kuntze, Rev. Gen. PI. 2: 468 (1891); Vandellia 
pusilla (Willd.) Merr., Philipp. J. Sci. Bot. 7: 
246 (1912). Type: India, s.dat., s.coll. (holo: 
B-W 377). 

Weakly erect herb to 10 cm. Stems decumbent, 
densely pubescent with filamentous 
trichomes to 1 mm. Leaves simple opposite, 
very shortly petiolate or subsessile, petioles 
up to 1 mm. long, hirsute; lamina 5-15(20) 
mm long, 3-15 mm. wide, broadly ovate 
to subcircular to cordate, crenate-dentate, 
filamentous trichomes on both sides but more 
densely below; 5-nerved from base of leaf 
Flowers axillary, in clusters of 1-3 per axil; 
pedicels slender, to 20 mm in flowers, 30 mm 
in fruit, with stalked glandular trichomes. 
Calyx deeply 5-lobed, almost to the base, 
lobes 3-4 mm long, single nerved and with 
stalked glandular and filamentous trichomes; 
margin finely serrate, scarious. Corolla 5-8.5 


Accepted for publication 29 July 2013 


mm. long, white to pale violet to purple; tube 
externally glabrous or with sparse stalked 
glandular trichomes, internally with stalked 
glandular trichomes near antero-lateral 
ridges, otherwise glabrous; upper lip narrow 
emarginate; lower lip 3-lobed. Stamens 4, 
fertile, cohering in pairs; upper filaments 
1-1.5 mm. long, glabrous; lower filaments 
3-4 mm long, broadly geniculate just above 
the base, forming a ‘webbed’ bend. Ovary 
c. 1 mm; style 5-7 mm. Capsule subglobose 
to ovoid, more or less equal in length to the 
calyx. Seeds light brown, cylindric, 0.5 x 0.3 
mm, with up to 8 longitudinal ribs. Fig. 1. 

Selected specimens examined : Malaysia. Lawn at 
Forestry Department Kuching Sarawak, Nov 1988, 
Wannan UNSW16578 (CANB, NSW, UNSW). Australia: 
Northern Territory. 2 km NE of Port Bradshaw, Gove 
Peninsula, Sep 1987, Russell-Smith 3380 & Lucas , (AD, 
DNA). Queensland. Cook District: North of Coen, Jul 
1999, Wannan 1379 (BRI, CANB); North of Coen, Sep 
2007, Wannan 4855 & Graham (BRI); ditto loc., Jun 
2009, Wannan 5696 & Thompson (CNS). 

Notes : Lindernia pusilla was first recognised 
in Australia from a collection made in 1999 
just north of Coen beside the Peninsula 
Developmental Road, Queensland ( Wannan 
BSW1379). Despite searches, no further 
localities have been recorded on Cape York 
Peninsula. During subsequent examination 
of Lindernia collections at AD a further 
collection was identified from the Northern 
Territory ( Russell-Smith 3380 & Lucas). 
These sites remain the only known Australian 
localities. 

According to Philcox (1968), Lindernia 
pusilla is widespread in the Indian 
subcontinent, China, Myanmar, Thailand 
and Malesia (Borneo, Celebes, Java, Malay 
Peninsula, Moluccas, Philippines, Sumatra 
and mainland New Guinea) up to an altitude 
of 1600 metres. It has also subsequently been 


Wannan, Lindemia 


127 


Fig. 1 . Lindemia pusilla (Wannan 1379\ BRI). Photo: B. Wannan. 


recorded from the Bismarck Archipelago 
of Papua New Guinea (Peekel 1984). The 
Australian material is a good match with 
the type and Malesian material (cf Wannan 
UNSW16578 ) based on its morphology and, 
for the latter, seed anatomy. 

Although a key to Queensland species will 
be provided in the revision (Wannan in prep.), 
L. pusilla can be distinguished from other 
Australian Lindemia by its multiple (pink) 
flowers per axil, long (to 30 mm) pedicels, 
and its stems pubescent with filamentous 
trichomes. 

Lindemia tectanthera W.R. Barker, J. 
Adelaide Bot. Gard. 13: 86-87 (1990). Type: 
Western Australia. Kimberley Region: 23.3 
km along King River Road, 25 April 1983, 
R.M. Barker 284 (holo: AD). 

Erect, sometimes decumbent, glabrous 
annual herb, 8-40 cm tall, with slender 
scapiform stems arising from a basal cluster 


of leaves; sometimes with stem branched in 
upper parts below the terminal inflorescence. 
Leaves at the base spathulate, 6-22 mm long, 
with the petiole 1-5 mm long, the blade ovate 
to obovate, sometimes narrowly so, 3-15 
x 1.5-8 mm, entire, with midvein. Cauline 
leaves/ bracts subulate, 1-4 mm x 1-2 mm. 
Inflorescence terminal racemes or panicles; 
pedicels 5-32 mm at anthesis and erect, 
15-40 mm in fruit and sometimes deflexed. 
Calyx 2-4.5 mm long; sepals 5, equal, divided 
almost to base, triangular, 3-nerved, often 
reddened. Corolla 2-lipped, with internal 
longitudinal flaps enclosing stamens and 
style; stalked glandular trichomes externally; 
white, with purple or mauve; tube 6-7 mm; 
upper lip recurved, 1.5-2 mm long, shallowly 
emarginate; lower lip widely spreading, 4-6.5 
mm long, the lobes shallowly emarginate, the 
mid lobe wider than the laterals. Stamens 
with upper 2 stamens functional, filaments 
c. 1 mm; anthers 0.5-0.8 mm long, yellow, 


128 


Austrobaileya 9(1): 126-129 (2013) 


Fig. 2. Lindernia tectanthera ( Wannan 6291 & Trenerry. BRI). Photo: B. Wannan. 


with 2 locules end to end, approximately 
180° divergent; lower stamens sterile, anthers 
represented by 2 appendages geniculate at 
summit, proximally thick, near the apex 
often bearing a minute anther vestige. Ovary 
1-1.5 mm, glabrous, style c. 5 mm, glabrous. 
Capsule ovoid, 4-5 x 2.5 mm, thin-walled, 
yellow-brown; seeds broad obovoid to 
oblong-ellipsoid, 0.3-0.35 x 0.2-0.25 mm, 
light brown, with 4 longitudinal angles and 
4-6 transverse ridges. Fig. 2. 

Selected specimens examined : Western Australia. 3 
km S of Pitta Creek near Prince Regent River, Jul 2009, 
Wannan 5740 (BRI, CNS, NSW, PERTH); Drysdale 
River NP, Jul 2012, Wannan 6563 & Wardrop (CNS, 
PERTH). Queensland. Cook District: Sand Ridge, 
Staaten River NP, Jan 2003, Fox IDF2002 (BRI); 
Wrotham Park, Jul 2011, Wannan 6114 (BRI); Gamboola, 
Aug 2011, Wannan 6291 & Trenerry (NSW), Wannan 
6298 & Trenerry (BRI). 

Notes: Lindernia tectanthera was first 
described from the Kimberley region of 
Western Australia (Barker 1990). This 
species has been recently confirmed from 
west of Chillagoe in Queensland and is a 


good match with material from the Kimberley 
including the holotype, based on morphology 
and seed anatomy. Examination of Fox 
IDF2002 (Staaten River NP) has shown it to 
be also L. tectanthera. Queensland specimens 
of this taxon have been previously incorrectly 
determined as Lindernia lobelioides (F.Muell.) 
F.Muell., a Northern Territory species. 

Lindernia tectanthera has been recorded 
widely from the Top End and Kimberley 
regions. The species typically occurs in moist 
areas often on the margin of sandy rises. It is 
distinguished from other Queensland species 
by its usual basal rosette of leaves and its 
upper corolla with a longitudinal flap which 
encloses the anthers and style. 

Acknowledgements 

Thanks to Darren Crayn (CNS) for his support 
of the project and the directors of AD, BRI, 
CANB, DNA, MEL, NSW and UNSW for 
loans and/or access to collections. This work 
has been supported by Australian Biological 
Resources Study (ABRS) National Taxonomy 


Wannan, Lindemia 


129 


Research Grant Program CN211-26 which has 

been undertaken at the Australian Tropical 

Herbarium. 

References 

Barker, W.R. (1990). Newtaxa, names and combinations 
in Lindemia , Peplidium , Stemodia and Striga 
(Scrophularlaceae) mainly of the Kimberley 
Region, Western Australia. Journal of the 
Adelaide Botanic Gardens 13: 79-93. 

Peekel, P.G. (1884). Flora of Bismarck Archipelago for 
Naturalists. Translated by E.E. Henty. Office of 
Forests, Division of Botany: Lae, Papua New 
Guinea. 

Philcox, D. (1968). Revision of the Malesian species of 
Lindemia All. (Scrophulariaceae). Kew Bulletin 
22: 1-72. 


Status and lectotypification of Craspedia gracilis 
Hook.f. (Asteraceae: Gnaphalieae) 

Andrew C. Rozefelds 

Summary 

Rozefelds, A.C. (2013). Status and lectotypification of Craspedia gracilis Hook.f. (Asteraceae: 
Gnaphalieae). Austrobaileya 9(1): 130-135. The identity of Craspedia gracilis Hook.f. is 
determined and it is shown to be conspecific with C. coolaminica J.Everett & Joy Thomps. which, 
being the younger name, is therefore placed in synonymy. The identity of Craspedia richea var. 
linearis Hook.f. is also resolved and it is shown to be a synonym of C. gracilis. Lectotypes of C. 
gracilis and C. richea var. linearis are designated and illustrated. 

Key Words: Asteraceae, Gnaphalieae, Craspedia, Craspedia coolaminica, Craspedia gracilis, 
Craspedia richea var. linearis, Australia flora, Tasmania flora, taxonomy, billy buttons 

A.C.Rozefelds, Queensland Museum, GPO BOX 3300, South Brisbane, Queensland 4101, Australia. 
Email: andrew. rozefelds@qm. qld.gov. au 


Introduction 

Previous studies of Craspedia G.Forst. by 
Rozefelds (2002) and Rozefelds et al. (2011), 
have resolved the nomenclature of species 
of Craspedia described from Tasmania; 
however, the status of two taxa described 
by Hooker (1847), C. gracilis Hook.f and C. 
richea Cass. var. linearis Hook.f, has been 
unclear. 

Cassini (1818) recognised that Richea 
glauca , described by Labillardiere (1800) 
from Tasmania, should be transferred to 
Craspedia. His new combination, Craspedia 
richea Cass.; however, is a nomen illegitimum, 
as recognized by Chapman (1991), since it 
was based on Richea glauca Labill. whose 
epithet should have been adopted. Sprengel 
(1826) successfully transferred this species to 
Craspedia , i.e. C. glauca (Labill.) Sprengel. 

Hooker (1847) described Craspedia 
gracilis Hook.f from the Middlesex Plains 
[Tasmania], although he later, in ‘ Flora 
Tasmaniae’, reduced C. gracilis to a variety, 
C. richea var. gracilis Hook.f Curtis (1963: 
346) recognised that ‘C. glauca included 
several further spp. and varieties [that] have 
been named’. She recognised four varieties 


including the new combination C. glauca 
var. gracilis , but as the original reference and 
dates of publication for this taxon were not 
cited, it is invalid (Chapman 1991). 

Hooker (1847) also recognised two new 
varieties of Craspedia richea. The status of C. 
richea var. glabrata Hook.f from the western 
mountains of Tasmania, was examined by 
Rozefelds (2002) and a new combination C. 
glabrata (Hook.f.) Rozefelds was proposed. 
The status of C. richea var. linearis Hook.f 
from the western mountains of Tasmania is 
reviewed herein. Curtis (1963) did not discuss 
the status of this variety. 

Craspedia coolaminica J.Everett & 
Joy Thomps. was described from material 
collected on mainland Australia and 
Tasmania, but the description of this species 
overlaps with that of both C. richea var. 
linearis and C. gracilis. To resolve the status 
of these taxa described by Hooker (1847), 
cibachromes of early Tasmanian collections 
of Craspedia gracilis and C. richea var. 
linearis, held at Kew, were obtained; and from 
the study of these images it is proposed that 
C. coolaminica and C. richea var. linearis are 
placed in synonymy under C. gracilis. 


Accepted for publication 25 September 2013 


Rozefelds, Craspedia gracilis 

Taxonomy 

Craspedia gracilis Hook.f. 

Craspedia gracilis Hook.f., Lond. J. Bot. 6: 
118 (1847); C. richea Cass. [nom. illeg .] var. 
gracilis (Hook.f.) Hook.f., Flora Tasmaniae 3: 
197 (1857); C. glaiica var. gracilis (Hook.f.) 
W.M.Curtis comb, invalid., The Student’s 
Flora of Tasmania 2: 247 (1963). Type 
citation: “Middlesex plains; Gunn”. Type: 
Tasmania. Middlesex PL, 15 February 1837, 
[R.C.]Gunn 835/1837 (lecto [here designated]: 
K n.v., cibachrome at HO!). The specimen 
associated with the label on the right hand 
side of the plate is the lectotype (Fig. 1). 

Craspedia richea Hook.f. [nom. illeg .] 
var. linearis Hook.f., Lond. J. Bot. 6: 118 
(1847); C. richea Cass. [nom. illeg] var. |3 
linearis Hook.f., Flora Tasmaniae 3: 197 
(1857). synon. nov. Type citation: “Western 
Mountains; Gunn”. Type: Tasmania. Western 
Mountains, 17 February either 1842 or 1843 (as 
the year is not consistent on the label), [R.C.] 
Gunn 1160/1842 (lecto [here designated]: K 
n.v., cibachrome at HO!). The more complete 
specimen on the left hand side of the plate is 
the lectotype (Fig. 2). 

Craspedia coolaminica J.Everett & Joy 
Thomps., synon. nov., Telopea 5: 45 

(1992). Type: New South Wales. Southern 
Tablelands: Happy Jacks, Snowy Mountains, 
near Happy Jacks Road, above sheep yard, 27 
January 1964, M.E. Phillips 20419 (holo: NSW 
n.v/, iso: CBG n.v) 

Typification : In describing Craspedia gracilis. 
Hooker (1847) did not specify a numbered 
collection of Gunn’s. The specimen Gunn 
835/1837 from Middlesex Plains comprises 
three individual plants and was examined 
by Hooker; hence, it is an appropriate choice 
for lectotypification of the name. A lectotype 
is chosen based on the right-hand specimen 
closest to the collector’s label. 

Craspedia richea var. linearis was 
described by Hooker (1847) as having narrowly 
linear leaves with arachnose hairs and known 
from the “Western Mountains”, and although 
no specific locality, nor collector number were 
recorded a specimen in K, Gunn 1160/1842, 


131 

dated 18 February 1843, is consistent with the 
limited published information. Notes on this 
herbarium sheet, in Ronald Campbell Gunn’s 
handwriting, record it as Craspedia, but the 
note has been annotated by a later author with 
‘richea var. linearis’. The narrowly linear 
leaves and silvery grey foliage are consistent 
with C. gracilis. Gunn’s notes on the type of 
C. richea var. linearis records the florets as 
orange (Fig. 2), and while no orange-floreted 
species are known from Tasmania, the bright 
yellow flowers of C. gracilis approach an 
orange colour (Fig. 3). The combination, 
C. richea var. linearis, was only used by 
Hooker, so it seems very likely that this 
collection was examined by Hooker, and as 
it comes from the “Western Mountains” and 
was collected by Gunn, it is an appropriate 
choice for lectotypification of this name. As 
the herbarium sheet includes two similar¬ 
sized specimens and the right-hand specimen 
is more complete, it is designated as the 
lectotype. 

Hooker (1857) in Flora Tasmaniae, 
considered C. richea var. |3 linearis as being 
restricted to the Western Mountains and 
referred a second collection ( Gunn 1160 ) from 
Arthurs Lake to this variety. This collection 
{Gunn 1160 ) comprises four individual plants; 
all have long narrow silvery grey leaves with 
finely appressed hairs, and with arachnose 
hairs on the margin and are consistent with C. 
gracilis as defined herein. 

Mainland Australian and Tasmanian 
populations of Craspedia gracilis 

Craspedia gracilis as defined here occurs in 
subalpine communities in New South Wales, 
Victoria and Tasmania (Everett & Thompson 
1992; Schmidt-Lebuhn & Milner 2013; 
Rozefelds pers. obs). Like many species of 
Craspedia, plants are often morphologically 
variable, with the size of the plants and the 
various organs (leaves, scapes, bracts on 
scape and compound head) phenotypically 
plastic. Material from Tasmania is variable in 
size from small herbs (like the type), to large, 
robust specimens 60 cm high (Rozefelds pers. 
obs). The mainland material show a similar 
variation in size (Everett & Thompson 1992) 
and no morphological differences were noted 


Fig. 1 . Lectotype of Craspedia gracilis Hook.f. (arrowed), Gunn 835/1837 (K). The lectotype shows the typical narrow 
linear silvery grey leaves, fine hairs and secondary veins are clearly evident on the underside of the leaves. Reproduced 
by kind permission of the Board of Trustees of the Royal Botanic Gardens, Kew. 


Rozefelds, Craspedia gracilis 


133 


Fig. 2. Lectotype of Craspedia richea Cass. var. p linearis Hook.f. (arrowed), Gunn 1160 (K). Reproduced by kind 
permission of the Board of Trustees of the Royal Botanic Gardens, Kew. 


134 


Austrobaileya 9(1): 130-135 (2013) 


Fig. 3. Floret colour in Craspedia gracilis Hook.f (left hand inflorescence) compared with C. rosulata Rozefelds & 
A.M.Buchanan (right hand inflorescence). 


between these mainland Australian and 
Tasmania populations by these authors in 
their original description. 

Sequence data for mainland Australian 
and Tasmanian samples of C. gracilis (i.e. 
C. coolaminica in Ford et al. 2007) showed 
no variation in ITS, seven base differences 
in ETS, and in psbA-trnH there were two 
base differences and also two deletions in 
the Tasmanian specimen (Kerry Ford, pers. 
comm. 2013). There seems little justification 
based upon the current molecular data, and 
the overall similarities in morphology to 
support separation of mainland Australian 
and Tasmanian populations. As the two 
samples of C. gracilis were sister to each other 
in Ford et al. (2007) analysis, it also becomes 
an arbitrary decision, in a phylogenetic sense, 
as to whether they are considered conspecific 
or sister species. 

Discussion 

The lectotype of Craspedia gracilis Hook.f 
{Gunn 835/1837 ) has long linear, narrowly 
lanceolate leaves, fine appressed hairs that 
impart the silver grey foliage, secondary 
veins clearly visible on the underside of the 
leaves due in part to the appressed hairs on 
the veins, and arachnose hairs along the leaf 
margins. 


The distinguishing features of Craspedia 
coolaminica J.Everett & Joy Thomps. (Everett 
& Thompson 1992; Rozefelds et al. 2011) 
easily distinguish this species from others 
in Tasmania, viz. distinctive basal rosette of 
long linear to narrowly attenuate silvery grey 
leaves that have a dense covering of long, fine 
appressed hairs, conspicuous secondary veins 
on the abaxial leaf surface, yellow corolla 
and mid-summer flowering (December- 
February). As no characters have been 
recognised that separate these taxa apart, I 
conclude that C. coolaminica and C. richea 
var. linearis are both synonyms of C. gracilis. 

Dawson et al. (1999) attributed a 
chromosome count of 2n=44 for Craspedia 
gracilis (as C. glauca ; Kerry Ford pers. 
comm. 2010). Putative hybrids have been 
noted between C. gracilis and the typically 
earlier flowering C. rosulata Rozefelds & 
A.M.Buchanan which is sympatric with this 
species at Miena, Central Plateau and the two 
species overlap slightly in flowering time. 
Putative hybrids are robust plants and have 
bright yellow florets like C. gracilis , rather 
than the light yellow of C. rosulata , but lack 
the conspicuous silver grey indumentum of C. 
gracilis. 


Rozefelds, Craspedia gracilis 

Acknowledgements 

I thank Alex Chapman in obtaining 
cibachromes of the relevant collections, 
Kerry Ford for advice on molecular data, and 
Alex Buchanan for his help and interest in the 
project. Fig. 1. and Fig. 2 are reproduced by 
kind permission of the Board of Trustees of 
the Royal Botanic Gardens, Kew. 

References 

Cassini, A.H.G. de (1818). Craspedie, Craspedia. In 
Dictionnaire des sciences naturelles, dans 
lequel on traite methodiquement des differens 
etres de la nature. Second Edition. F.G. 
Levrault: Strasbourg/ Le Normant: Paris. 

Chapman, A.D. (1991). Australian Plant Name Index 
A-C. Australian Biological Resources Study: 
Canberra. 

Curtis, W.M. (1963). The Student’s Flora of Tasmania, 
Part 2. Government Printer: Hobart. 

Dawson, M.I., Breitwieser, I. & Ward, J.M. (1999). 
Chromosome numbers in Craspedia , Ewartia 
and Pterygopappus (Compositae: Gnaphalieae). 
Australian Systematic Botany 12: 671-674. 

Everett, J. & Thompson, J. (1992). New alpine and 
subalpine species in Craspedia sens, strict. 
(Asteraceae: Gnaphalieae). Telopea 5: 45-51. 

Ford, K.A., Ward, J.M., Smissen, R.D., Wagstaff, 
S.J., Breitwieser, 1. (2007) Phylogeny and 
biogeography of Craspedia (Asteraceae: 
Gnaphalieae) based on ITS, ETS and psbA-trnH 
sequence data. Taxon 56: 783-794. 


135 

Hooker, J.D. (1847). Florae Tasmaniae Spicilegium or 
Contributions towards a Flora of Van Diemens 
Fand. London Journal of Botany 6: 106-125. 

Hooker, J.D. (1857). Botany of the Antarctic Voyage 
of H.M. Discovery ships Erebus and Terror in 
the years 1839-1843. Vol. 3. Flora Tasmaniae. 
Fovell Reeve: Fondon. 

Fabillardiere, J.J.H. de (1800). Relation du voyage a 
la recherche de La Perouse, fait par ordre de 
L’assemblee Constituante, pendant les annies 
1791, 1792, et pendants l ere la 2 de annee de la 
Republique Francoise. 2 Vols. H.J. Jansen: 
Paris. 

Rozefelds, A.C. (2002). A new species and new 
combination in Craspedia (Asteraceae) from 
Tasmania. Telopea 9: 813-820. 

Rozefelds, A.C., Buchanan, A.M. & Ford, K.A. 
(2011). New species of Craspedia (Asteraceae: 
Gnaphalieae) from Tasmania, and determination 
of the identity of C. macrocephala Hook. 
Kanunnah 4: 93-116. 

Schmidt-Febuhn, A.N. & Milner, K.V. (2013). A 
quantitative study of morphology in Australian 
Craspedia (Asteraceae: Gnaphalieae). 

Australian Systematic Botany 26: 238-254. 

Sprengel, C.P.J. (1826). Systema Vegetabilium 3. 
Gottingae sumtibus librariae: Dieterichlanae. 


Craspedia uniflora G.Forst. (Asteraceae) is a New 
Zealand endemic and not part of the Australian flora 

Alexander N. Schmidt-Lebuhn 

Summary 

Schmidt-Lebuhn, A.N. (2013). Craspedia uniflora G.Forst. (Asteraceae) is a New Zealand endemic 
and not part of the Australian flora. Austrobaileya 9(1): 136-139. Specimens identified as Craspedia 
uniflora from Australia in the Queensland Herbarium are shown to have been misidentified and this 
species should therefore be deleted from both the Census of the Queensland Flora and the Australian 
Plant Census. The absence of any Australian material of this species in Australian herbaria also 
indicates that C. uniflora is a New Zealand endemic. Four species of Craspedia are identified 
as occurring in Queensland, C. canens J.Everett & Doust, C. haplorrhiza J.Everett & Doust, C. 
paludicola J.Everett & Doust (syn. C. sp. (GirraweenNP S.T.Blake 23643)) and C. variabilis J.Everett 
& Doust, and a key to these species is provided. 

Key Words: Asteraceae, Compositae, Gnaphalieae, Australian Plant Census, Census of the 
Queensland Flora, Craspedia , Craspedia canens , Craspedia haplorrhiza , Craspedia paludicola , 
Craspedia uniflora , Craspedia variabilis , identification key 

A N. Schmidt-Lebuhn, CSIRO Plant Industry, Centre for Australian National Biodiversity Research, 
GPO Box 1600, Canberra, ACT 2601, Australia. Email: alexander.schmidt-lebuhn@csiro.au 


Introduction 

Craspedia G.Forst. (Asteraceae, Gnaphalieae) 
is a genus of c. 20 Australian and at least 6 New 
Zealand species of mostly perennial rosette 
plants bearing yellow, white or orange flowers 
in solitary compound heads on leafy scapes. 
Although found in all states of Australia 
except the Northern Territory, Craspedia is 
most diverse in subalpine to alpine regions of 
the south-eastern mainland and Tasmania. 

Craspedia was described by Forster 
(1786) with one species, C. uniflora G.Forst. 
from New Zealand. Over the following two 
hundred years, several other species were 
described from New Zealand and Tasmania 
but only one from the Australian mainland 
(see Schmidt-Lebuhn & Milner 2013 for an 
overview), so that mainland specimens were 
often haphazardly referred to species from 
New Zealand and Tasmania, especially C. 
uniflora and C. glauca (Labill.) Spreng. Curtis 
(1963) did not consider C. uniflora to occur 
in Tasmania, while Stanley & Ross (1986) 
used the name for all specimens of Craspedia 


Accepted for publication 24 July 2013 


s.str. in south-eastern Queensland (the other 
two species they treated under Craspedia are 
now assigned to Pycnosorus Benth. which 
is characterized by sessile partial heads, 
yellow phyllaries and pappus, and a different 
base chromosome number; Everett & Doust 
1992a). 

Recent taxonomic or floristic treatments 
of Craspedia in Tasmania and the Australian 
mainland did not mention the species (Everett 
& Doust 1992b, c; Everett & Thompson 1992; 
Everett 1999; Costin et al. 2000; Rozefelds 
2002; McDougall & Walsh 2008; Rozefelds et 
al. 2011). Most Australian specimens formerly 
identified as C. uniflora , including all those at 
CANB/CBG, have been assigned to species 
described in these recent studies. However, C. 
uniflora is still listed in the latest Census of the 
Queensland Flora (Bostock & Holland 2010) 
and in the Australian Plant Census (CHAH 
2011) as occurring in all states except the 
Northern Territory. No nationwide taxonomic 
treatment of Craspedia is currently available. 

Ford et al. (2007) demonstrated that the 
New Zealand species of Craspedia form a 
monophyletic group nested in the Australian 
representatives of the genus and thus are likely 
to represent the descendants of a single long 


Schmidt-Lebuhn, Craspedia uniflora 

distance dispersal event with no subsequent 
re-colonization of Australia. Because the New 
Zealand lineage has diversified into at least 
six species since its arrival (Allan 1961), it 
appears a priori unlikely that a contemporary 
species such as C. uniflora would be found on 
both sides of the Tasman Sea. 

In this paper the status of Craspedia 
uniflora in Australia is examined to clarify 
whether Australian specimens identified as 
this species were merely overlooked when 
the taxonomy of Australian Craspedia was 
updated or whether at least some of them are 
conspecific with those from New Zealand. A 
recent quantitative study of morphological 
characters in the genus aimed to include 
specimens from all formally described native 
Australian species (Schmidt-Lebuhn & 
Milner 2013), and it was therefore important 
to determine whether C. uniflora needed to be 
included. 

Materials and methods 

All specimens labelled as Craspedia uniflora 
were requested as a loan from BRI, and a 
photograph of the type of the species was 
obtained from K. The morphology of the 
type as discernible from the photograph, 
the morphology of selected New Zealand 
specimens and the description of C. uniflora 
by Allan (1961) were compared against 
the morphology of Australian specimens 
previously identified as C. uniflora and 
the descriptions of and keys to Australian 
species (in particular Everett & Doust 1992a; 
Everett & Thompson 1992). The results of the 
quantitative analyses by Schmidt-Lebuhn & 
Milner (2013) were also taken into account. 

Common abbreviations in the specimen 
citation include National Park (NP). All 
specimens have been seen unless indicated 
n.v. 

Results 

As circumscribed by Allan (1961), Craspedia 
uniflora of New Zealand is a heterogeneous 
species of four varieties occurring from 
coastal rock crevices to subalpine herb-fields. 
It is characterized by a very distinctive margin 
of tangled long white hairs around the leaves 


137 

but overall plant size, glomerule size and 
flower colour are variable (yellow or white 
in the latter case). Under the typical variety, 
the type specimen at K is described as having 
leaves obovate- to lanceolate-spathulate, c. 7 
x 1.5 cm, scapes c. 12 cm long and glomerules 
of c. 1.5 cm diameter, values that are rather 
small for a Craspedia. Visual examination of 
the photograph of the type showed it to match 
this description, with the single glomerule 
c. 1.4 cm in diameter and the longest intact 
leaf c. 6 cm long. A very distinctive margin 
of white hairs around the rosette and scape 
leaves is clearly visible. 

No Australian specimens of Craspedia 
are consistent with the above morphology 
and descriptions. The only native species of 
similar stature as the type of C. uniflora are 
C. alba J.Everett & Joy Thomps., C. glabrata 
(Hook.f.) Rozefelds and C. rosulata Rozefelds 
& A.M.Buchanan, but all of them differ 
from this species especially in regard to the 
indumentum. Whereas Craspedia lamicola 
J.Everett & Joy Thomps. has a margin of hairs 
superficially similar to that of C. uniflora 
around otherwise glabrous to subglabrous 
leaves it is a much more robust plant restricted 
to wet alpine habitats with significantly larger 
scapes, leaves and glomerules. 

Craspedia specimens from Queensland 
previously identified as C. uniflora were 
referable to C. canens J.Everett & Doust, C. 
haplorrhiza J.Everett & Doust, C. paludicola 
J.Everett & Doust and C. variabilis J.Everett 
& Doust. The species of Craspedia in 
Queensland can be determined with the 
following key, with the caveats that more work 
is needed to understand the circumscription 
and taxonomy of what is currently called C. 
canens and that C. variabilis may ultimately 
turn out to be a synonym of Tasmanian C. 
glauca. (Schmidt-Lebuhn & Milner 2013). 


138 


Austrobaileya 9(1): 136-139 (2013) 

Key for the identification of Craspedia species that occur in Queensland 

1 Plants short-lived, with a single tap-root.2. C. haplorrhiza 

1. Plants perennial, with numerous secondary roots.2 

2 Tall plants up to 75 cm, with subglabrous to glabrous leaves, growing in 

swamps or drainage lines.3. C. paludicola 

2. Smaller plants usually only to 50 cm, with hairy leaves, growing in drier habitats.3 

3 Leaves with woolly indumentum.1. C. canens 

3. Leaves with robust septate indumentum and often with glands but not 

woolly.4. C. variabilis 


Taxonomy 

1. Craspedia canens J.Everett & Doust, 
Telopea 5: 37 (1992). Type: New South 
Wales. Central Tablelands: Wingecarribee 
[Wingicarribee] Swamp, 26 October 1971, 
L.A. Craven 1981 (holo: NSW 221409 n.v.-, 
iso: CANB; L n.v .). 

Additional specimens examined: Queensland. Darling 
Downs District: Stanthorpe, Feb 1916, Wright s.n. (BRI 
[AQ247074]); Stanthorpe, s.dat., Davidson s.n. (BRI 
[AQ247073, 247077]); “The Junction track”, Girraween 
NP, Oct 1974, McDonald 467 (BRI); Victoria. Port 
Phillip, s.dat., Mossman 321 (BRI). 

Note: Mossman 321 is very small and 
glabrescent, showing few of the diagnostic 
woolly hairs. 

2. Craspedia haplorrhiza J.Everett & 
Doust, Telopea 5: 37 (1992). Type: New 
South Wales. Far South Western Plains: 
southern crossroad, north of Ranger’s Station, 
Kinchega National Park, 12 September 1989, 
A. Denham 39 (holo: NSW n.v .; iso: AD n.v). 

Specimens examined : Queensland. Darling Downs 
District: 10 miles [16.6 km] SE of St George, Aug 1947, 
Everist 3137 (BRI); New South Wales, c. 56 km S of 
Walgett, Lightning Ridge/Angledoon Road, Sep 1989, 
White s.n. (BRI [AQ504366]). 

3. Craspedia paludicola J.Everett & Doust, 
Telopea 5: 35 (1992). Type: Victoria. Top End 
Reference Area Barmah State Park, Murray 
Valley Study Area Sector: F, sub-block: 41B, 
28 September 1985, A.C. Beauglehole 81321 
(holo: MEL 681491 n.v., iso: CANB; HO n.v., 
NSW n.v). 

Craspedia sp. (Girraween NP S.T.Blake 
23643), synon. nov. (Bostock & Holland 
2007, 2010). 


Additional specimens examined: Queensland. 

Darling Downs District: About 7 miles [11 km] E of 
Wyberba, Oct 1958, Pedley 297 (BRI); Girraween NP 
near Wyberba and Wallangarra, Nov 1971, Blake 23643 
(BRI); Racecourse Creek, Dingo Swamp, Girraween 
NP, Aug 1995, Forster PIF17626 (BRI, CANB). 

Notes: The loan made available by BRI also 
included several additional specimens not 
previously identified as C. uniflora, among 
them three sheets of the undescribed phrase 
name taxon Craspedia sp. (Girraween NP 
S.T.Blake 23643) (Bostock & Holland 2007, 
2010; CHAH 2011). They do not differ in 
any significant way from C. paludicola 
J.Everett & Doust, especially considering the 
morphological variability of the latter species 
over its range on both sides of the Bass Strait. 
Although Pedley 297 is an unusually small 
plant, its glabrous leaves and the habitat are 
typical of C. paludicola. 

4. Craspedia variabilis J.Everett & Doust, 
Telopea 5: 36 (1992). Type: South Australia. 
South Eastern Region: c. 15 km south-east 
of Mt Burr Township, 7 September 1966, 1.B. 
Wilson 511 (holo: AD n.v.-, iso: CANB). 

Additional specimens examined: Western Australia. 
Mundaring Weir, Aug 1926, Bicks.n. (BRI [AQ247068]); 
Dalyup, 30 miles [50 km] W of Esperance, Sep 1971, 
Scarth-Johnson 820 (BRI). Queensland. Darling 
Downs District: Cons Plain, NE of Killarney, Oct 
2000, Bean 16897 (BRI); Killarney, Sep 1937, Schindler 
5 (BRI). Moreton District: turnstile gate above 
Grace’s Hut, near Mt Barney, s.dat., Leiper s.n. (BRI 
[AQ584399]). New South Wales. 7.4 km along Cochrane 
Road, Carrai NP, W of Kempsey, Dec 2007, Bean 27170 
(BRI, CANB). Victoria. Melbourne, sine coll. s.n. (BRI 
[AQ247063]). South Australia. Near Kapunda, in 1888, 
Bailey s.n. (BRI [AQ247062]); Eyre Peninsula Region, 
Marble Range, slopes of North Block, Sep 1979, Weber 
6018 (BRI). 


Schmidt-Lebuhn, Craspedia uniflora 

Discussion 

No Australian specimen previously labelled 
as C. uniflora was found to be consistent with 
the type or with descriptions (Allan 1961) 
and herbarium vouchers from New Zealand. 
Craspedia uniflora therefore is endemic 
to New Zealand, and the name should be 
removed from the Census of the Queensland 
Flora and the Australian Plant Census. 

This study shows that four species of 
Craspedia s.str. occur in Queensland: C. 
canens, C. haplorrhiza, C. paludicola and 
C. variabilis. Craspedia haplorrhiza is a 
widespread species of the western slopes 
and the margin of the arid zone, occurring in 
Queensland south from St George to around 
Bourke, Brewarrina and Walgett in New 
South Wales. The distribution of the other 
three species in the state is restricted to small 
areas in its south-eastern corner. Craspedia 
paludicola is known only from Girraween 
National Park, and C. canens from the same 
National Park and from some historical 
collections labelled as ‘Stanthorpe’. 

Acknowledgements 

I thank the Director of the Queensland 
Herbarium for the loan of material, Nicholas 
Hind and Elizabeth Woodgyer (Kew 
Royal Botanic Gardens) for moving the 
photographing of the type of Craspedia 
uniflora ahead of the schedule of the Global 
Plants Initiative to make it available for 
examination, and Paul I Forster, Brendan 
Lepschi, Joseph T. Miller and one anonymous 
reviewer for helpful comments on a previous 
version of this manuscript. 

References 

Allan, H.H. (1961). Flora of New Zealand. Volume 
1. Indigenous Tracheophyta: Psilopsida, 
Lycopsida, Filicopsida, Gymnospermae, 
Dicotyledones. New Zealand Government 
Printer: Wellington. 

Bostock, P.D. & Holland, A.E. (2007). Census of 
the Queensland Flora 2007. Queensland 
Herbarium, Environmental Protection Agency: 
Brisbane. 

- (2010). Census of the Queensland Flora 

2010. Queensland Herbarium, Department 
of Environment and Resource Management: 
Brisbane. 


139 

CHAH [Council of Heads of Australasian Herbaria] 
(2011). Australian Plant Census. http://www. 
anbg. gov. au/chah/apc/ 

Costin, A., Gray, M., Totterdell, C. & Wimbush, D. 
(2000). Kosciuszko Alpine Flora , 2 nd edition. 
CSIRO Publishing: Melbourne. 

Curtis, W.M. (1963). The Student’s Flora of Tasmania 
Part 2 - Angiospermae: Lythraceae to 
Epacridaceae. St. David’s Park Publishing: 
Hobart. 

Everett, J. (1999). Craspedia. In N.G. Walsh & T.J. 
Entwisle (eds.). Flora of Victoria 4: 758-764. 
Inkata Press: Melbourne. 

Everett, J., Doust, A.N.L. (1992a). New species and a 
new combination in Pycnosorus (Asteraceae: 
Gnaphalieae). Telopea 5: 39-43. 

- (1992b). Four new Australian species 

of Craspedia sens, strict. (Asteraceae: 
Gnaphalieae). Telopea 5: 35-38. 

- (1992c). Craspedia. In G.J. Harden (ed), 

Flora of New South Wales 3: 221-226. NSW 
University Press: Sydney. 

Everett, J. & Thompson, J. (1992). New alpine and 
subalpine species in Craspedia sens, strict. 
(Asteraceae: Gnaphalieae). Telopea 5: 45-51. 

Ford, K.A., Ward, J.M., Smissen, R.D., Wagstaff, 
S.J. & Breitwieser, I. (2007). Phylogeny and 
biogeography of Craspedia (Asteraceae: 
Gnaphalieae) based on ITS, ETS and psbA-trnH 
sequence data. Taxon 56: 783-794. 

Forster, G. (1786). Florulae Insularum Australium 
Prodromus. J.C. Dieterich: Gottingen. 

McDougall, K.L. & Walsh, N.G. (2008). A conspectus 
of high-country Craspedia (Asteraceae: 
Gnaphalieae) of mainland south-eastern 
Australia. Muelleria 26: 3-10. 

Rozefelds, A.C. (2002). A new species and new 
combination in Craspedia (Asteraceae) from 
Tasmania. Telopea 9: 813-820. 

Rozefelds, A.C., Buchanan, A.M. & Ford, K.A. 
(2011). New species of Craspedia (Asteraceae: 
Gnaphalieae) from Tasmania and determination 
of the identity of C. macrocephala Hook. 
Kanunnah 4: 93-116. 

Schmidt-Lebuhn, A.N. & Milner, K.V. (2013). A 
quantitative study of morphology in Australian 
Craspedia (Asteraceae, Gnaphalieae). 
Australian Systematic Botany 26: 238-254. 

Stanley, T.D. & Ross, E.M. (1986). Flora of south¬ 
eastern Queensland. Volume 2. Queensland 
Department of Primary Industries: Brisbane. 


Cullen spicigerum (Domin) A.E.Holland (Fabaceae), a new 
combination and reinstatement of a north Queensland species 

Ailsa E. Holland 

Summary 

Holland, A.E. (2013). Cullen spicigerum (Domin) A.E.Holland (Fabaceae), a new combination 
and reinstatement of a north Queensland species. Austrobaileya 9(1): 140-145. Psoralea spicigera 
Domin has been relegated to synonymy under the Western Australian Cullen lachnostachys 
(F.Muell.) J.W.Grimes. This species is here reinstated and a new combination, Cullen spicigerum 
(Domin) A.E.Holland, is made. A revised description of this species is provided, along with notes on 
distribution and habitat and a key to the species of Cullen in Queensland. 

Key Words: Fabaceae, Cullen, Cullen spicigerum, Psoralea, Psoralea spicigera, Australia flora, 
Queensland flora, identification key, taxonomy, new combination 

Ailsa E. Holland, Queensland Herbarium, Department of Science, Information Technology, 
Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, 
Australia. E-mail: Ailsa.Holland@science.dsitia.qld.gov.au 


Introduction 

Cullen Medik., tribe Psoraleeae , is a 
widespread genus of 32 species of which 
26 occur in Australia (Grimes 1997; AVH 
2013). The genus was revised by Grimes 
(1997) who placed Psoralea spicigera Domin 
in synonymy under Cullen lachnostachys 
(F.Muell.) J.W.Grimes, albeit without any 
explanatory notes as to why this decision was 
made. However, in his introduction, Grimes 
commented that “a couple of problems 
remain which will best be addressed through 
detailed field-work.. .There are several 
forms included in C. badocanum that might 
merit some taxonomic status, but which are 
incompletely known from few collections”. It 
is possible that this taxon is referred to here, 
as Grimes probably saw only a few specimens 
and only one of the specimens cited below 
was mentioned anywhere in his revision 
{Bolton 475 was determined by Grimes as C. 
pustulatum (F.Muell.) J.W.Grimes). Numerous 
new collections are all consistent with 
Domin’s description of Psoralea spicigera 
except for the stipules which are generally 
shorter than the 1 cm described by him. The 
species is however quite distinct and the 
main differences are listed in Table 1. This 


Accepted for publication 9 September 2013 


north Queensland species is therefore here 
reinstated, and a new combination, Cullen 
spicigerum (Domin) Holland, is made. 

Materials and methods 

This paper is based on herbarium collections 
in the Queensland Herbarium (BRI). The 
species was examined in the field in 2012. 
A common abbreviation in the specimen 
citations is: NP (National Park). 

Taxonomy 

Cullen spicigerum (Domin) A.E.Holland, 
comb, nov.; Psoralea spicigera Domin, 
Biblioth. Bot. 89(2): 186-187 (1926). Type: 
Queensland. Cook District: Elizabeth Ck 
[Creek], 16 July [1872], W.Hann, Cape York 
Peninsula Expedition No. 40 (lecto: PR; 
isolecto: K, online image!;^^ Grimes [1997: 
615]). 

Cullen sp. (Westmoreland C.H.Gittins 2491); 
Holland & Pedley (2013). 

Erect herbaceous shrubs to 3 m tall; bark 
grey. Branches striate, densely pubescent with 
short crisped white or grey hairs, moderately 
glandular with large raised light brown or 
blond glands that are generally sunken or 
flush with surface or pustulate but generally 
not stipitate (see below). Leaves heteroblastic; 
the proximal leaves pinnately trifoliolate, the 


Holland , Cullen spicigerum 141 

Table 1. Comparison of morphological characters for Cullen spicigerum and putatively 
related taxa 


Character state 

C. spicigerum 

C. badocanum 

C. pustulatum 

C. lachnostachys 

indumentum 

dense, short 
crisped 

villose to 
lanate-villose 

glabrate to 
villulose 

velutinous or 
short-villose 

stipule length 
(mm) 

3-5 

4-12 

4-12 

5-8.5 

inflorescence 

shape 

cylindrical 

subglobose 

cylindrical 

cylindrical 

inflorescence 
bract persistence 

persistent 

persistent 

deciduous or 
persistent 

deciduous 

flower length 
(mm) 

6-10 

8.5-15 

7-10 

10.5-14.5 

lower calyx lobe 
length (mm) 

5-8 

7.7-14 

2.2-3.5 

3.5-6.5 

pedicel length 
(mm) 

0.5-2 

0.5-3 

0.5-3 

4-7 


distal leaves unifoliolate; petioles 5-20 mm 
long; rachis 3-15 mm long; both petiole and 
rachis striate, pubescent and with scattered 
glands. Leaflets lanceolate, rounded to 
cuneate at base, acute or obtuse and apiculate 
at apex; the upper surface sparsely pubescent 
or nearly glabrous, with scattered glands; 
lower surface moderately densely pubescent, 
glandular with larger stipitate and crateriform 
glands congregated on the veins. Terminal 
leaflets 25-80 mm long; 8-35 mm wide, 
length:width ratio 1.8-4.4. Lateral leaflets 
similar, 2/3 of the length of the terminal 
leaflet. Stipules triangular, reflexed, 3-5 
mm long, 1-1.5 mm wide, glandular and 
pubescent on outer surface, glabrous on inner 
surface, with several obscure veins. Stipels 
absent. Pseudoracemes indeterminate, one 
per axil, compact, cylindrical, occasionally 
branched, 1-10 cm long, 1.5-2 cm wide, 
initially shorter than the subtending leaf, 
longer at maturity; peduncle 20-80 mm long; 
rachis with 9—16 flowering nodes, initially 
shorter than the peduncle, elongating through 
the growing season. Flowers usually in pairs, 
rarely 3 together, overlapping on the rachis; 
internodes 2- 8 mm long. Bracts persistent, 


initially erect, becoming relaxed, broadly 
ovate to suborbicular, 3-6 mm long, 2.5-4.5 
mm wide, the apex abruptly narrowed into a 
slightly reflexed acuminate tip; outer surface 
sparsely hairy mainly on the margins and 
densely covered in large, light brown pustular 
glands; inner surface glabrous. Bracteoles 
absent. Pedicels 0.5-2 mm long, pubescent. 
Flowers 6-10 mm long, corolla shorter or 
slightly longer than the calyx. Calyx tubular- 
campanulate, somewhat flattened, 7-10 mm 
long, 5-lobed, purplish-green, pubescent with 
short crisped white or grey hairs (sometimes 
tending to villous but not obscuring veins 
or glands); glands scattered, large, pustular; 
calyx tube 2-3 mm long, rounded at base, 
with 10 raised veins of which 5 end in the lobe 
sinuses and 5 continue to lobe apices; adaxial 
4 lobes triangular, acute, 2-4 mm long, the 
uppermost two slightly longer and partly 
joined; abaxial lobe 5-8 mm long, carinate, 
curving upwards over the keel to a long 
attenuate point. Corolla glabrous, purplish at 
the apex and fading to white at base; standard 
obovate, 6-9 mm long, 6-7 mm wide, slightly 
longer than wings and keel, emarginate, 
tapered to an indistinct claw at base; wings 


142 


oblong, 8-9 mm long including a 2-3 mm 
claw, c. 2.5 mm wide, rounded at apex; keel 
upturned and enclosed in the other petals, c. 8 
mm long including a 3-4 mm claw and 2 mm 
wide, rounded at apex and curved along the 
lower edge, the two halves joined in the distal 
half. Ovary glabrous, glandular, stipitate. 
Fruit and flower remnant falling together 
(excluding bract). Fruit stipitate; stipe to 3 mm 
long; pod broadly elliptic, flattened, 3-3.5 
mm long, 1.7-2.3 mm wide, glabrous or with 
a few marginal hairs, very densely glandular 
with pustular glands, the fruit wall strongly 
adhering to the seed. Seed reniform, 2.3-2.5 
mm long, 3.2-3.8 mm wide, smooth, green to 
light brown, hilum minute. Fig. 1. 

Additional specimens examined : Queensland. Cook 
District: 1 km N of Old Laura Homestead, Lakefield 
NP, Aug 2012, Thompson ST12628 (BRI, MEL); near 
Old Laura Homestead, Lakefield NP, Jul 1993, Stanton 
2 (BRI); Lakefield NP, 3.5 km along 12 Mile Waterhole 
Road, Jul 2010, McDonald KRM9701 (BRI, PE); c. 0.5 
km from Kowanyama, Oct 1981, Birchley 127 (BRI); 
Cooktown Development Road, c. 0.5 km E of Boggy 
Creek, Apr 2001, Jago 5899 & Wannan (BRI, MEL, 
NSW); Wrotham Park, Trimbles Crossing, Walsh River, 
May 2012, McDonald KRM 12753 & Holland (BRI); 
Murphy’s Farm, Gilbert River, Aug 2001, Smith AS2038 
(BRI); Near Gamboola Homestead, N of Chillagoe, 
Aug 1985, Bolton 475 (BRI); Van Lee Homestead, 65 
km NNW of Mt Surprise, Jul 2001, Addicott EPA873 
(BRI); Gilbert River, May 1954, Ever is t 5477 (BRI). 
Burke District: 14.5 km SE of Lotus Vale, on Lotus 
Vale to Stirling Road (Site 106), Jun 2001, Turpin 
GPT592 & Thompson (BRI); Westmoreland, 56 km 
from Wollogorang towards Burketown, Jul 1972, Gittins 
2491 (BRI). North Kennedy District: Cape Cleveland 
section of Bowling Green Bay NP, S of Townsville, Jul 


Austrobail'ey a 9(1): 140-145 (2013) 
1991, Bean 3459 (BRI, MEL, MEXU). 

Distribution and habitat: Cullen spicigerum 
occurs in open woodland of Corymbia, 
Eucalyptus or Melaleuca usually with a 
grassy understorey and generally on alluvial 
silty clay or loam soils, often on river banks, 
flood plains or disturbed areas. The species 
is endemic to north Queensland where it is 
widespread on southern Cape York Peninsula 
and west nearly to the Northern Territory 
border (Map 1). 

Phenology: This species flowers and fruits 
continuously through the cooler months from 
April through to August, occasionally as late 
as October. 

Notes: Cullen spicigerum is related to the 
widespread C. badocanum (Blanco) Verde, 
and C. pustulatum , and the Western Australian 
endemic C. lachnostachys. It differs from C. 
badocanum by the cylindrical pseudoracemes 
and the shorter stipules, flowers and abaxial 
calyx lobe, and from C. pustidatum by the 
dense indumentum (especially on the calyx), 
shorter pseudoracemes, shorter stipules and 
much longer abaxial calyx lobe. It differs 
from C. lachnostachys by the persistent and 
smaller bracts, the shorter pedicels, flowers 
and stipules, and the shorter pubescence 
(especially on the calyx) and the longer 
abaxial calyx lobe. 

Conservation status: Cullen spicigerum 
is Least Concern using the IUCN (2001) 
criteria. 


Key to the species of Cullen in Queensland 


1 Leaflets 3 or 5, palmately arranged. C. tenax 

1. Leaflets 3, pinnately arranged, or leaves unifoliolate.2 

2 Plants covered in long spreading white hairs > 1 mm long; bracts 

lanceolate and more than 6 mm long.C. plumosum 

2. Plants glabrous or if hairs present then < 1 mm long; bracts ovate to 

orbicular, less than 6 mm long.3 

3 Leaves all unifoliolate.4 

3. Leaves trifoliolate, or both unifoliolate and trifoliolate leaves present.6 

4 Inflorescence an elongate spike; calyx shorter than the fruit.C. leucanthum 

4. Inflorescence subglobular on a shortened rachis; calyx longer than the fruit.5 


Holland , Cullen spicigerum 143 

5 Whole plant densely covered in black pustular glands; leaves distinctly 

denticulate; flowers to 7 mm long.C. balsamicum 

5. Sparsely to moderately glandular with brown glands and flush with surface; 

leaves entire or weakly toothed; flowers 9-15 mm long.C. badocanum 

6 Glands on stems and petioles pustular or stipitate.7 

6. Glands on stems and petioles flush with surface, or sunken.8 

7 Calyx much shorter than corolla, the hairs confined to veins; abaxial 

calyx lobe scarcely longer than the others.C. pustulatum 

7. Calyx longer or shorter than corolla, moderately hairy; abaxial calyx lobe 

c. 3 mm longer than the others.C. spicigerum 

8 Leaflets mostly more than 5 cm long, lanceolate, entire.9 

8. Leaflets mostly less than 5 cm long, ovate to lanceolate, denticulate, 

dentate, erose or serrate.10 

9 Plants glabrous or nearly so; corolla much longer than calyx, predominantly 

white, cream or yellowish with a purple blotch.C. leucanthum 

9. Plants hairy to some degree; corolla as long or slightly shorter than calyx, 

purple to white.C. spicigerum 

10 Calyx 2-4 mm long, shorter than the fruit.C. cinereum 

10. Calyx 3-8 mm long, longer than the fruit.11 

11 Calyx sparsely hairy with appressed white hairs in rows, and much 

longer than corolla.C. graveolens 

11. Calyx densely hairy with spreading or appressed white or black hairs, and 

shorter than or equal to corolla.12 

12 Inflorescence condensed, usually shorter than the subtending leaf; 

peduncles less than 4 cm long; corolla usually white.C. discolor 

12. Inflorescences elongated and interrupted, much longer than the subtending 

leaf; peduncles longer than 4 cm; corolla usually purple, pink or blue.13 

13 Plants strictly prostrate.C. patens 

13. Plants erect or ascending, bushy.14 

14 Entire plants densely white velvety hairy, the glands of the leaves obscured . . C. pallidum 

14. Only calyces with dense white hair; the glands on the leaves apparent . . C. australasicum 


Acknowledgements 

I would like to thank Les Pedley for drawing 
this species to my attention and Keith 
McDonald for his excellent specimens, 
information and on ground guidance. I also 
acknowledge Ruth Clark who located the 
isolectotype at Kew and arranged for an 
image to be provided. Will Smith prepared 
the illustrations and map. 

References 

AVH (Australia’s Virtual Herbarium) (2013). Council 
of Heads of Australasian Herbaria, viewed 16 
August 2013, <http://avh.ala.org.au/> 


Domin, K. (1926). Psoralea L. Bibliotheca Botanica 
89(2): 186-187. 

Grimes, J.W. (1997). A revision of Cullen (Leguminosae: 
Papilionoideae). Australian Systematic Botany 
10: 565-648. 

Holland, A.E. & Pedley, L. (2013). Fabaceae. In 
P.D. Bostock & A.E. Holland (eds). Census 
of the Queensland Flora 2013. Queensland 
Herbarium, Department of Science, Information 
Technology, Innovation and the Arts: Brisbane. 
https://data.qld.gov.au/dataset/census-of-the- 
queensland-flora-2013 

Iucn (2001). Red List Categories and Criteria: Version 
3.1. IUCN Species Survival Commission. 
IUCN: Gland, Switzerland/Cambridge, U.K. 


144 


Austrobaileya 9(1): 140-145 (2013) 


Fig. 1 . Cullen spicigerum. A. branch xl. B. leaflet lower surface xl6. leaflet upper surface xl6. D. stipule x8. E. bract 
x8. F. calyx x6. G. standard petal x6. H. keel petal x6.1. wing petal x6, J. staminal column x6. K. ovary and style 
enclosed in staminal column x6. L. fruit x8. K. seed x8. All from McDonaldKRM9701 (BRI). Del. W.Smith. 


Holland, Cullen spicigerum 


Map. 1 . Distribution of Cullen spicigerum in Queensland. 


Austrobaileya 9(1): 146-147 (2013) 


146 


SHORT COMMUNICATION 

Celtis strychnoides Planch, an earlier name for 
Celtis australiensis Sattarian (Ulmaceae) 

Gordon P. Guymer 

Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic 
Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: Gordon.Guymer@science.dsitia.qld.gov. 
au 


Celtis australiensis Sattarian was described 
by Sattarian (2005) as occurring in north west 
Western Australia, the Northern Territory and 
north west Queensland and was distinguished 
from Celtisphilippensis Blanco by its elliptic, 
cordate, pilose leaves (2-7 x 1.5-5 cm), 
spinose leaf apices and margins. This name 
has subsequently been incorrectly applied to 
all Australian specimens previously known as 
Celtis philippensis (CHAH 2007). Sattarian 
(2005) referred to the Flora of Australia 
treatment of Celtis by Hewson (1989) but 
overlooked the name Celtis strychnoides 
Planch, given by Hewson as a synonym of 
Celtis philippensis. 

Planchon (1848) described Celtis 
strychnoides from a collection of Allan 
Cunningham’s from Careening Bay, 
Kimberley, northern Western Australia. He 
distinguished it from Celtis philippensis by 
its small, pubescent, ovate or rarely broadly 
ovate leaves that were barely 1.5 inches (4 cm) 
long. 

Examination of the type collections 
and images of Celtis strychnoides , C. 
australiensis and specimens of Celtis at BRI, 
MEL and NSW confirm that C. strychnoides 
is conspecific with C. australiensis and as 
such is the earlier name. 

Celtis strychnoides Planch., Annales des 
Sciences Naturelles, Botanique ser. 3, 10: 
306-307 (1848). Type: [Western Australia] 
Careening Bay, north west Australia, January 
1820, A. Cunningham 193 in herb. Hook, 
(holo: K [852750, 852752 images !] , iso: BM 
[838618, 838619, 799835 images !], BRI p.p , 
MEL p.p ). 


Accepted for publication 2 September 2013 


Celtis australiensis Sattarian, Blumea 50: 499 
(2005), synon. nov. Type: Western Australia. 
Northern Province: Geikie Gorge, near 
gorge mouth, 14 May 1992, I. Telford 11722 
(holo: CANB n.v., iso: BISH n.v., BRI, L n.v., 
NSW, PERTH n.v.). 

Illustration : Sattarian & van der Maesen 
(2005: 500, Fig. 1). 

Additional specimens examined : Western Australia. 
N end of Bougainville Peninsula, Apr 1982, Kenneally 
8076 (BRI); Walsh Point, Port Warrender, May 1981, 
Tracey 15170, 15075 (BRI); Port Warrender, Oct 1974, 
Rodd 2900 (NSW); Surveyor’s Creek Road, May 1981, 
Tracey 15122 (BRI); Lone Dingo, May 1981, Tracey 1500 
(BRI); Merten’s Falls, May 1981, Tracey 15062 (BRI); 
Near Crusher Pool, May 1981, Tracey 15233 (BRI); 
Cambridge Gulf, in 1886, Ranford & Nyulasy 18 & 36 
(MEL); Wyndham, in 1903 & Nov 1906, Fitzgeralds.n. 
(NSW); Cockburn Range, 13 km W of King River, July 
1974, Carr 3316 & Beaiiglehole 47094 (NSW); 4 km S 
of junction of Neville Creek & Calder River, May 1983, 
Fell 9 (BRI); Cape Leveque, Jul 1973, Webb & Tracey 
13141 (BRI); King’s [King] Sound, in 1869, Hughan 
s.n. (MEL 2210491, 2210489, 2210481); ditto, in 1888, 
Froggart s.n. (NSW); Napier Range, Jul 1973, Tracey 
14807 (BRI); Mornington Wildlife Sanctuary, NE of 
Fitzroy Crossing, Apr 2005, Murphy MULE453 & Legge 
(BRI); Geikie Gorge, 16 km NE of Fitzroy Crossing, 
Jun 1970, Briggs 3671 (NSW); ditto, May 1981, Tracey 
15143 (BRI); 10 miles [16 km] SW of Lissadell Station, 
Aug 1949, Perry 2672 (MEL); S of Ord River Station, 
Aug 1973, Tracey 14100 (BRI). Northern Territory. 
Victoria River, Dec 1855, Mueller s.n. (K 852751, MEL 
2210482, 2210475); Limestone Gorge, Gregory NP, Apr 
1990, Evans 3091 (MEL); Gregory NP, Apr 1996, Booth 
1696 (MEL); ditto , Apr 1996, Duretto 1187 (MEL); ditto , 
Apr 1996, Foreman 2249 (MEL); ditto , Apr 1996, Jones 
83 (MEL); 5 km S of Timber Creek, May 1983, Briggs 
904 (MEL); 15 miles [24 km] SSE of Timber Creek, Jun 
1952, Perry 2869 (BRI); 41 km SSE of Turkey Creek, 
Jul 1984, Scarlett 84-334 (MEL); 6 miles [10 km] NW 
of Katherine, Adams 806 (NSW); 11 miles [18.3 km] S 
of Katherine, Feb 1961, McKee 8487 (NSW); 16-Mile 
Cave Reserve, S of Katherine, May 1978, Webb & 
Tracey 12588 (BRI); 4 miles [6 km] E of Elsey, Oct 1958, 
Chippendale 5065 (BRI, NSW); 15 miles [24 km] S of 
MacArthur River Station, Jul 1949, Perry 1732 (BRI, 


147 


MEL); near head of Kilgour River, Sep 1911, Hill 556 
(MEL); Settlement Creek, Jun 1922, Brass 161 (BRI). 
Queensland. Burke District: 39 km SW of Hells Gate, 
Apr 2006, Thompson WES384 & Edginton (BRI); 
17 km W of Adel’s Grove, Lawn Hill NP, Nov 1983, 
Buttock UNSW14474 & St George (MEL); 20.2 km S of 
Musselbrook, Apr 1995, Thomas & Johnson MRS469 
(BRI); Lawn Hill NP, May 1988, O’Keefe 11 (BRI); 1 
km N of Gregory River, Riversleigh, May 1970, Webb 
& Tracey 1349 (BRI); 29 km NW of Norfolk Station, 
Apr 2005, Kelman DTK1021 (BRI); Seymour River, 
vicinity of Nicholson River, Jul 1886, Dittrich s.n. (BRI 
[AQ88147]); 7 km NE of Camooweal, Jul 2006, Booth 
CAM29-1 & Kelman (BRI). 

Distribution : Celtis strychnoides occurs in 
the Kimberley region of northern Western 
Australia, central northern Northern Territory 
and north western Queensland. 

Habitat : The species occurs in semi-deciduous 
notophyll vine thickets or woodlands usually 


in rocky situations on slopes or hills in fire 
refugia, on limestone, basalt or sandstone. 

Notes: TheNorthern Territory and Queensland 
collections from higher rainfall areas (>1100 
mm mean annual rainfall) that have glabrous, 
ovate or oblong leaves 6-18 x 3.5-8.5 cm 
and that were included in C. australiensis 
by CHAH (2007) are referrable to Celtis 
philippensis. These collections may represent 
a species distinct from C. philippensis but a 
revision of this species complex is required to 
resolve the taxonomy 

The Cunningham sheets at BRI and MEL 
include specimens of both C. strychnoides 
and C. philippensis. 

Celtis strychnoides can be distinguished 
from Australian material of Celtis philippensis 
by the following key: 


1 Leaf blades ovate or broadly ovate, acute, 2-7 x 1.5-4 cm, young leaves 
sparsely pubescent, glabrescent except veins and leaf base below; base 
rounded or cordate; petioles and branchlets pubescent with white erect 
hairs 0.1-0.3 mm long, petioles 3-6 mm long; juvenile leaves with 
spinose margins, occasionally adult leaves with short marginal spines; 

small tree to 4-8 (-10) m. WA, NT & NW Qld.Celtis strychnoides 

1. Leaf blades oblong to ovate, acuminate, 6-18 x 3.5-8.5 cm, glabrous; 
base cuneate to truncate; petioles glabrous, occasionally sparsely 
pubescent, 5-8 mm long; juvenile leaves with serrate or crenate 
margins; tree 4-25 m. Coastal NT & N & E Qld.Celtis philippensis 


Acknowledgements 

I thank the Directors and staff of MEL and 

NSW for allowing access and examination of 

their collections. 

References 

CHAH (2007). Australian Plant Census. Council of 
Heads of Australasian Herbarium, http://www. 
anbg. gov. au/chah/apc/ 

Hewson, H.J. (1989). Ulmaceae. In AS. George 
(ed.). Flora of Australia 3: 10-13. Australian 
Government Publishing Service: Canberra. 

Planchon, J.E. (1848). Sur les Ulmacees, Celtis. Annales 
des Sciences Naturelles, Botanique ser. 3, 10: 
283-316. 

Sattarian, A. & van der Maesen, L. J.G. (2005). Two new 
species of Celtis (Celtidaceae) from Australia 
and Madagascar. Blumea 50: 499-503. 


Austrobaileya 9 ( 1 ): 148-149 ( 2013 ) 


148 


SHORT COMMUNICATION 

Reinstatement of Rapanea leucobrachya P.Royen 
(Myrsinaceae) from New Guinea, with a new combination 
as Myrsine leucobrachya (P.Royen) P.I.Forst. 

Paul I. Forster 

Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, 
Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: Paul. 
For ster @science. dsitia. qld. gov. au 


Van Royen (1982) named a number of new 
species of Rapanea Aubl. from montane 
areas of New Guinea. Many of these 
were subsequently reduced to synonymy 
by Sleumer (1986) when he revised the 
New Guinea species. The New Guinea 
species recognised by Sleumer (1986) were 
uncritically transferred to Myrsine L. by 
Pipoly (Takeuchi & Pipoly 2009), together 
with the addition of one new species {Myrsine 
warrae W.N.Takeuchi & Pipoly). 

During curation of New Guinea 
Myrsinaceae at BRI it became obvious that 
Sleumer’s treatment of certain Myrsine 
species was overly conservative. This 
pertained particularly to his reduction to 
synonymy of Rapanea leucobrachya P.Royen 
and R. wapu P.Royen under R. rhombata 
P.Royen without any accompanying reasons. 
Whilst the recognition of three species for 
this group of montane populations from the 
Eastern and Western Highlands of Papua New 
Guinea may be too liberal, recognition of only 
one is not supported by easily discernible 
morphological differences such as leaf shape, 
venation and gland morphology. Hence, two 
species, M. rhombata (P.Royen) Pipoly and 
M. leucobrachya (P.Royen) P.I.Forst. are 
recognised herein. 

Myrsine rhombata (P.Royen) Pipoly, 
Edinburgh J. Bot. 66: 462 (2009); Rapanea 
rhombata P.Royen, Alpine FI. New Guinea 
3: 1976 (1982). Type: Papua New Guinea. 


Accepted for publication 25 July 2013 


Eastern Highlands Province: Mt Wilhelm, 
east slopes, 28 July 1959, L.J.Brass 30757 
(holo: L, online image!; iso: US, online 
image!). 

Specimens examined : Papua New Guinea. Eastern 
Highlands Province: Kurumigl-bokwa, Kuski River 
above Toromambuno Mission, Upper Chimbu, Aug 
1957, Pullen 324 (BRI); Mt Wilhelm, Lake Piunde, Aug 
1957, Robbins 684 (BRI). Western Highlands Province: 
track to Mt Hagen, E of Mt Kigum, Apr 1977, Vinas & 
Veldkamp LAE59812 (BRI). Central Province: Murray 
Pass, Wharton Range, Jun-Sep 1933, Brass 4546 (BRI); 
Murray Pass, Aug 1970, Foreman & Lelean NGF48412A 
(BRI); Mt Victoria area, Iawan Swamp, c. 2 km SSW of 
Mt Service, May 1976, van Royen 11033 (BRI). 

Myrsine leucobrachya (P.Royen) P.I.Forst. 
comb, nov.; Rapanea leucobrachya P.Royen, 
Alpine FI. New Guinea 3: 1975 (1982). Type: 
Papua New Guinea. Southern Highlands 
Province: Mt Ialibu, slope to SW of Repeater 
Station, 10 August 1972, P.F. Stevens & 
D.B.Foreman LAE55853 (holo: K, online 
image!; iso: BRI; L, online image!). 

Rapanea wapu P.Royen, Alpine FI. New 
Guinea 3: 1971 (1982), synon. nov. Type: 
Papua New Guinea. Western Highlands 
Province: Northern slope of Sugarloaf 
complex (along Wapu River), 21 July 1960, 
R.D.Hoogland 7232 & R.Schodde (holo: L, 
online image!; BM, online image!). 

Additional specimens examined : Papua New Guinea. 

West Sepik Province: Bielga Range, W of Folongonom, 
second camp below Tamanagabip on Busilmin Track, 
May 1975, Vinas & Wiakabu LAE59529 (BRI). Eastern 
Highlands Province: track to Mt Michael, 5 km SW of 
Lufa, Apr 1982, Kairo 458 (BRI). Western Highlands 
Province: around the base camp, Mt Kigum, Apr 1977, 
Vinas & Veldkamp LAE59836 (BRI). Central Province: 
E slope to Lake Myola No 1, Sep 1973, Croft & Lelean 
NGF34965 (BRI). 


149 


The two species can be distinguished by the following key. 

Lamina rhombate-obovate, tip acuminate to retuse, below with numerous 
elongate glands, round glands along margins; 2° lateral veins obscure 
or very indistinct, 9-12 per side of 1° vein (midrib). Bracteoles 
broadly ovate to elliptic-ovate, 1-1.5 x 0.8-1.2 mm. 

Lamina elliptic, rhombate-elliptic to obovate, tip acuminate to obtuse, 
below with numerous nail-like glands on either side of 1° vein; 2° 
lateral veins prominent and easily discernible above and below, 8-14 
per side of 1° vein (midrib), joined and forming an intramarginal vein, 
3° veins prominent and reticulate below. Bracteoles ovate, 0.8-2.2 x 
1-1.2 mm. 

References 

Sleumer, H. (1986). A revision of the genus Rapanea 
Aubl. (Myrsinaceae) in New Guinea. Blumea 
31:245-269. 

Takeuchi, W. & Pipoly, J. (2009). Nomenclatural transfers 
in East Malesian Myrsine (Myrsinaceae), and 
the description of M. warrae , a distinctive new 
species from ultrabasic environments in Papua 
New Guinea. Edinburgh Journal of Botany 66: 

459-467. 

Van Royen, P. (1982). The Alpine Flora of New Guinea. 

Volume 3: taxonomic part Winteraceae to 
Polygonaceae. J. Cramer: Vaduz. 


M. rhombata 


M. leucobrachya 


Austrobaileya 9 ( 1 ): 150-152 ( 2013 ) 


150 


SHORT COMMUNICATION 

Emmenospermapancherianum Baill. (Rhamnaceae) 
newly recorded for Australia 

A.R. Bean 

Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, 
Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: tony. 
bean@science.dsitia.qld.gov.au 


Emmenosperma F.Muell. is a genus of 
five species found in northern and eastern 
Australia, New Guinea, Sulawesi, New 
Caledonia and Fiji. It was described by 
Mueller (1862) based on Emmenosperma 
alphitonioides F.Muell. from Australia. Soon 
after, Bentham (1863) described another 
Australian species, E. cunninghamii Benth. 
Baillon described E. pancherianum Baill. 
from New Caledonia in 1874. In the 20 th 
century, two additional species, originally 
described in the genus Colubrina Rich, ex 
Brongn., were transferred to Emmenosperma , 
viz. E. micropetalum (A.C.Sm.) M.C.Johnst. 
and E. papuanum (Merr. & L.M.Perry) 
M.C.Johnst. 

In 2005, an unusual Emmenosperma 
specimen was submitted to the Queensland 
Herbarium, collected from Mt Zero-Taravale 
Sanctuary near Townsville. It was very unlike 
the two well-known Australian species, and an 
ensuing literature search suggested a strong 
affinity with E. pancherianum. Flowering and 
fruiting material was obtained from the Mt 
Zero-Taravale population in 2009, and this 
material has been compared with specimens 
of E. pancherianum collected from New 
Caledonia. The comparison revealed that there 
are no significant morphological differences 
between the New Caledonian material and the 
Mt Zero-Taravale material. 


Emmenosperma pancherianum Baill., 
Adansonia 11: 269 (1874). Type citation: ‘in 
Austro-Caledonia ubi in sylvis collium [hill- 
forests of southern Caledonia], leg. Pancher 
(herb.)’. Type: Nouvelle Caledonie, May 
1865, J.A.I. Pancher s.n. (lecto: P 06886541 
[here chosen]). 

Illustration : Sprague (1915: t. 3027). 

Shrub or small tree to 5 m. Young branches 
with dense indumentum of appressed simple 
hairs, older branches glabrous. Stipules 
caducous, hairy, triangular, c. 0.8 mm long. 
Leaves 16-40 x 8-22 mm, lamina obovate, 
with 2-4 obscure pairs of lateral veins at 
45-70° to midrib, base cuneate, apex obtuse 
or emarginate; upper surface glabrous (except 
at base on midrib), very glossy; lower surface 
less glossy, glabrous; petioles 1.5-4 mm long, 
with appressed hairs. Inflorescences axillary, 
borne on short lateral branchlets 2-5 mm 
long with rusty hairs near apex; rachises 
absent or up to 2 mm long. Flowers 5-merous. 
Hypanthium and sepals with antrorse 
appressed brownish hairs; sepals deflate, 
1-1.2 mm long, obtuse. Petals elliptical, 0.9- 
1.1 mm long, white, glabrous, hooded. Ovary 
glabrous, disc inconspicuous at base of ovary; 
style bulbous, 0.6-0.8 mm long, 2-lobed 
at apex. Fruiting pedicels sparsely hairy, 
2.5-5(-6.5) mm long; fruits ± globose, 5-6 
mm long, 2-locular; endocarpids 2, splitting 
longitudinally from the apex for up to half 
their length. Seeds flattened ellipsoidal, 4-5.7 
mm long, surface smooth and shiny, orange 
to red, remaining attached to fruiting pedicels 
after endocarpids have been shed. 


Accepted for publication 6 May 2013 


Bean, Emmenosperma pancherianum 

Additional specimens examined : New Caledonia: 
Nekoro, Aug 2003, Dagostini & Rigault 703 (NOU); 
Gouaro Deva, Oct 2005, Dagostini 979 (NOU); Noumea, 
Jun 1909, Franc 1382 (BRI); Noumea, presqu’ile Ducos, 
Apr 1970, McKee 21778 (NOU); Tontouta, Jul 1971, 
McKee 24789 (NOU); Ngoue, Paita, Mar 1981, McKee 
38781 (NOU); Plaine des gaiac, Jul 1987, Veillon 6411 
(NOU); Mt Tereka, Noumea, Jul 1991, Veillon 7405 
(NOU); Poya-Nord, Oct 1998, Veillon 8134 (NOU). 
Australia: Queensland. North Kennedy District: Near 
Return Creek falls, ‘Taravale’, NW of Townsville, May 
2009, Bean 28865 & Jensen (BM, BRI, CANB, CNS, 
MO, NSW, P); Return Creek Falls, ‘Taravale’, 80 km 
WNW of Townsville, Jul 2005, Camming s.n. (BRI 
[AQ612648]). Cultivated. Botanic Gardens, Sydney, Jan 
1921, White s.n. (BRI [AQ23692]). 

Distribution and habitat : This species is 
distributed along the west coast of New 
Caledonia, between Koumac and Noumea 
(Barrault 2006). In Australia, it is known 
from a single population at Mt Zero- 
Taravale Sanctuary north-west of Townsville, 
Queensland. In New Caledonia, it inhabits the 
“dry forests” of the western coast (Barrault 
2006), a habitat that has been decimated 
for grazing and agriculture (Hequet 2010). 
In Queensland, it grows on the steep south¬ 
facing slope of a granite ridge, in open forest 
dominated by Araucaria cunninghamii Mudie 
and Eucalyptus granitica L.A.S.Johnson & 
K.D.Hill. 

Phenology: Flowers April to October; fruits 
September to March. 

Typification : The protologue specified 
‘Pancher’ as the collector. Several Pancher 
specimens at P predate the publication date. 


151 

P 06886541 was chosen as the lectotype 
because it is a very good flowering specimen, 
and attached to the sheet is a handwritten 
description of the species, most likely by the 
author of the name. 

Notes: Emmenosperma pancherianum 

provides one more instance of the well known 
floristic connection between tropical eastern 
Australia and New Caledonia (Morat et al. 
1986; Ladiges & Cantrill 2007). 

While a large proportion of New 
Caledonian plant species are endemic, there 
are numerous examples of species that occur 
in both New Caledonia and eastern Australia. 
These include not only littoral species that 
are dispersed by ocean currents e.g. Premna 
serratifolia L. and Cerbera manghas L., but 
others such as Carissa ovata R.Br., Croton 
insularis Baill., Dry petes deplanchei (Brongn. 
& Gris) Merr., Polyalthia nitidissima (Dunal) 
Benth., Psydrax odorata (G.Forst.) A.C.Sm. 
& S.P.Darwin, Pterocaulon serrulatum 
(Montrouz.) Guillaumin and Rhamnella 
vitiensis (Benth.) A.C.Sm. 

Conservation status: Emmenosperma 
pancherianum is listed as ‘Vulnerable’ under 
the criteria of the IUCN Red List, based on its 
New Caledonian occurrence (Hequet 2010). 
In Australia, it is currently known from a 
single population comprising between six 
and 10 individuals. Its recommended status, 
based on the Australian population only, is 
Critically Endangered, Criterion D (IUCN 
2001 ). 


Key to the species of Emmenosperma in Australia 

1 Inflorescences axillary, borne in dense ± spherical clusters, rachises 

lacking or < 2 mm long; petioles 1.5-4 mm long Qld, New Caledonia . E. pancherianum 

1. Inflorescences predominantly terminal, some in upper leaf axils, 

paniculate to corymbose, rachises 13-70 mm long; petioles 5-27 mm long. 2 

2 Bracts and bracteoles conspicuous at anthesis; style 2-fid; fruiting pedicels 

(3-)4-5.5 mm long; fruits 2-locular Qld, NSW .E. alphitonioides 

2 . Bracts and bracteoles inconspicuous at anthesis; style 3-fid; fruiting 

pedicels 1.5-2.5(-4) mm long; fruits 3-locular Qld, NT, WA, New 

Guinea .E. cunninghamii 


152 


Austrobaileya 9 ( 1 ): 151-153 ( 2013 ) 


Acknowledgements 

I am grateful to Rigel Jensen, botanist with the 
Australian Wildlife Conservancy, for taking 
me to the ‘Taravale’ population. Thanks are 
also due to the Director of NOU for the loan 
of specimens and to Juergen Kellerman for 
helpful comments and suggestions. 

References 

Barrault, J. (2006). Plantes des Forets Seches de 
Nouvelle-Caledonie. Programme Forets Seches 
et Centre d’Initiation a FEnvironnement: 
Noumea. 

Bentham, G. (1863). Emmenosperma. In Flora 
Australiensis 1: 414-415. L.Reeve: London. 

Hequet, V. (2010). Emmenosperma pancherianum. In 
IUCN Red List of Threatened Species. Version 
2012.2. <www.iucnredlist.org>. Downloaded 
on 20 October 2012. 

Iucn (2001). Red List Categories and Criteria: Version 
3.1. IUCN Species Survival Commission. 
IUCN: Gland, Switzerland/Cambridge, U.K. 

Ladiges, P.Y. & Cantrill, D. (2007). New Caledonia - 
Australian connections: biogeographic patterns 
and geology. Australian Systematic Botany 20: 
383-389. 

Morat, P, Veillon, J.-M. & Mckee, H.S. (1986). Floristic 
relationships of New Caledonian rainforest 
phanerogams. Telopea 2: 631-679. 

Mueller, F. (1862). Emmenosperma. Fragmenta 
Phytographiae Australiae 3: 62-63. 

Government Printer: Melbourne. 

Sprague, T.A. (1915). Emmenosperma pancherianum , 
Baill. Hooker’s leones Plantarum 31: t. 3027. 


Austrobaileya 9 ( 1 ): 153-154 ( 2013 ) 


153 


SHORT COMMUNICATION 

Euphorbia operta Halford & W.K.Harris (Euphorbiaceae), 
a new name for Euphorbia occulta Halford & 
W.K.Harris, nom. illeg. and lectotypification of the 
name Euphorbia australis var. subtomentosa Domin 

D.A. Halford & W.K. Harris 

d- Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, 
Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia 


In our recent taxonomic treatment of 
Euphorbia section Anisophyllum Roeper 
(Euphorbiaceae) in Australia (Halford & 
Harris 2012) we incorrectly published the 
name E. occulta Halford & W.K.Harris for an 
endemic Australian species from northwest 
Queensland. We were unaware that the name 
had already been applied by Klotzch (1856) 
to a North America species, so the use of 
the name by us was illegitimate, being a 
much later homonym. Therefore a new name 
Euphorbia operta Halford & W.K.Harris 
is provided here for the Australian species. 
The specific epithet is from Latin, opertus , 
hidden from sight, concealed, in reference to 
the concealment of the involucral glands by 
the erect hood-like gland appendages of this 
species. 

Euphorbia operta Halford & W.K.Harris, 
nom. nov.; Euphorbia occulta Halford 
& W.K.Harris, Austrobaileya 8: 529-530 
(2012), nom. illeg., non Klotzsch, Bot. Voy. 
Herald [Seemann] 7-8: 277 (1856). Type: 
Queensland. Burke District: 12.3 km along 
Lake Moondara Road from Barkly Highway, 
N of Mt Isa, 1 July 2011, D. Halford QM524A 
(holo: BRI, iso: DNA, K, MEL, MICH, NT, 
distribuendi). 

We also overlooked the need to lectotypify the 
name Euphorbia australis var. subtomentosa 
Domin in order to fix the application of this 
name. Domin’s (1927) description of E. 
australis var. subtomentosa was based on 


material collected by Dr E. Clement from 
between the Ashburton and De Grey Rivers 
in Western Australia. There are four sheets 
that were seen by Domin, one in PR [528303] 
and three in K [186468, 186496, 186467], 
These comprise all the original material for 
the name that we have been able to locate. 
Careful examination of the four sheets 
clearly indicated that the specimens affixed 
to them belong to two different taxa, namely 
those currently known as E. australis var. 
subtomentosa (K 186468, PR 528303 (lower 
two fragments)) and E. vaccaria Baill. var. 
vaccaria (K 186496, 186467, PR 528303 
(upper three fragments)). There is little doubt 
that all the material was used by Domin in his 
formulation of the original description of E. 
australis var. subtomentosa. The specimen 
numbered K 186468 is selected here as 
lectotype of E. australis var. subtomentosa. 
If either of the sheets (K 186496, 186467) 
or fragments (PR 528303 (upper three 
fragments)) were chosen as lectotype, E. 
australis var. subtomentosa would have to 
be placed in synonymy under E. vaccaria 
and a new name would be required for 
what is currently known as E. australis var. 
subtomentosa. 

Euphorbia australis var. subtomentosa 

Domin, Biblioth. Bot. 89(4): 310 (1927 ‘1926’). 
Type: Western Australia. Inter Ashburton et 
De Gray [Grey] River s.d., E. Clement s.n. 
(lectotype [here chosen]: K 186468; isolecto: 
PR 528303 (lower two fragments)). 


Accepted for publication 10 October 2013 


Halford & Harris, Euphorbia names 

References 


Austrobaileya 9(1): 153-154 (2013) 


Domin, K. (1927). Euphorbia L. In Beitrage zur Flora 
und Pflanzengeographie Australiens, 1. Tiel, 3. 
Abt. Bibliotheca Botanica 89(4): 306-312. 

Halford, D.A. & Harris, W.K. (2012). A taxonomic 
revision of Euphorbia section Anisophyllum 
Roeper (Euphorbiaceae) in Australia. 
Austrobaileya 8: 441-600. 

Klotzsch, J.F. (1856). Euphorbiaceae. InB.C. Seemann, 
Botany of the Voyage of H.M.S. Herald, under 
the Command of Captain Henry Kellett, R.N., 
C.B., during the years 1845-1851 , pp. 276-278. 
L. Reeve: London.