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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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.

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.].

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

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).

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

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.

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

(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,

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;

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

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,

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/

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);

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.

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

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:

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

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 ).

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,

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

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

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

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,

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.

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.

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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.

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

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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

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

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

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.

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.

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

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

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.

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

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

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

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.

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

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

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

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

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

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.

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

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

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

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.

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

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

Bean, Sphaeromorphaea and Ethuliopsis

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

Map 4. World distribution of Sphaeromorphaea harrisii.

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

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

Bean, Sphaeromorphaea and Ethuliopsis

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).

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

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

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

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

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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

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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

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

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]).

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

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”.

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

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.

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

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).

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

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

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.

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).

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

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

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,

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

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

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

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

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

outline ovate

sometimes

outline ± rectangular

sometimes

outline semi-circular

sometimes

usually

shape distinctly

reniform

sometimes

usually

shape sub-reniform

(i.e. cordate lobes not

well developed)

sometimes

shape falcate

sometimes

leaf margins crenate

from base near netiole

to apex

usually

no 1

leaf base cordate

yes

usually

leaf base truncate

sometimes

usually

distinct basal lobes

present

yes

usually

distinct sinus present

formed by lobes

yes

usually

leaves much wider

than long

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.

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.

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

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),

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,

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

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.

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

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.

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

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)

t h)

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

Austrobaileya 9(1): 114—125(2013)

I Diplazium amamianum

' Diplazium hachijoense

— Diplazium doederleinii

Diplazium taiwanense

- Diplazium virescens var. conterminum

- Diplazium virescens

|— Diplazium crassiusculum

Diplazium donianum var. aphanoneuron

^ Diplazium lobatum

Diplazium dilatatum

Diplazium hayatamae

Diplazium deciduum

Diplazium fauriei

U- Diplazium mettenianum

Diplazium griffithii

— Diplazium pullingeri

L-=— Diplazium kawakamii

- Diplazium centripetale

—— Diplazium incomptum

9 |- Diplazium longicarpum

- 12

97 1 --- Diplazium subserratum

Diplazium proliferum

5 Diplazium chinense

L_6

Diplazium squamuligerum

Diplazium subtripinnatum

■ Diplazium esculentum

— Diplazium nipponicum

■ Diplazium sibiricum var. glabrum

Diplazium squamigerum

9 Diplazium bombanasae

— Diplazium cristatum

Diplazium lonchophyllum

- Diplazium plantaginifolium

13 | Diplazium okudairae

' Diplazium wichurae

—— 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

100

■ Diplazium pinfaense

r^- Diplazium wichurae

100

L^- Diplazium heterocarpum

Diplazium hainanense

Diplazium hachijoense

Diplazium esculentum

Diplazium nipponicum

15.

Diplazium mettenianum

Diplazium squamuligerum

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.

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Mpelasoka, F., Hennessy, K., Jones, R. & Bates, B.

(2008). Comparison of suitable drought indices

for climate change impacts assessment over

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1292.

Pachebo, L. & Moran, R.C. (1999). Monograph of

the neotropical species of Callipteris with

anastomosing veins (Woodsiaceae). Brittonia

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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

5-8.5

inflorescence

shape

cylindrical

subglobose

cylindrical

inflorescence

bract persistence

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

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.

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