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| OUEENSLAND GOVERNMENT

Department of

Environment and Heritage

QUEENSLAND GOVERNMENT

epartment of

Environment and Fleritage

‘

Editorial Committee

L.W. Jessup (editor)

R.J.F. Henderson (technical advisor)

B.K. Simon (technical advisor)

Desktop Publishing

Amelia S. Cumberbatch

Austrobaileya

Vol. 1, No. 1 was published on | December 1977

Vol. 5, No. | was published on 16 December 1997

Austrobaileya is published once per year.

Exchange: This journal will be distributed on the basis of exchange.

Subscriptions: Orders for single issues and subscriptions may be placed. The price is (1999)

A$25 per issue for individuals, AS40 for institutions, including postage.

All correspondence relating to exchange, subscriptions or contributions to this journal should be

addressed to The Editor, Austrobaileya, Queensland Herbarium, Brisbane Botanic Gardens Mt.

Coot-tha, Mt. Coot-tha Road, Toowong Qld 4066, Australia.

ISSN 0155-4131

Austrobaileya ts the journal of the Queensland Herbarium and 1s devoted to publication of results

of sound research and of informed discussion on plant systematics, with special emphasis on

Queensland plants.

Opinions expressed by authors are their own and do not necessarily represent the policies or views

of the Queensland Herbarium.

Austrobaileya 5(2): 157-366 (1999)

Contents

A revision of the Babingtonia virgata (J.R.Forst. & G.Forst.) F.Muell.

complex (Myrtaceae) in Australia

PAGS SESCALD Stops te: 55,845 BygrE dh BR rsd, HE Gee AR Sad en ile. Feelin delve bad Wy bi Sek Wee RLS ate aS

Systematic and cladistic studies of Myrtella F. Muell. and Lithomyrtus F.

Muell. (Myrtaceae)

Neil Snow and Gordon P. Guymer ..... 0.000 ccc eee eee eee ees

Desmodium Desv. ( Fabaceae) and related genera in Australia: a taxonomic

revision

PoESAPCGISV Sh. lS ehis enycteryrtt ar0e3) + gol eaten VE toda een ke dean thae ytatn Weenie ar iba Geena n tyne 3

Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland, Australia

NlarCO Ar: ASUTSUG.. 6.2 a)51s:2,5 wae dee tie wns. a Bi EH ER Oe nd eM Pan qed eb ee

Taxonomic status and Australian distribution of the weedy neotropical grass

Leptochloa fusca subsp. uwninervia, with an updated key to Australian

Leptochloa (Poaceae, Chloridoideae)

Neth snow. atid: Bryair K Simi 2. aoe 4 eer wee iae Noa pa on Soe Seeley Rew ate BOR a tae Bee Bn

Notes on Acacia (Leguminosae: Mimosoideae) chiefly from northern

Australia

Les Pedley

* @ &» © @ &© © &© © &© #&© € &© #@ &© © #& © © © F&F © FE & & £€ SF *F 8 8 8 FF © 8 F 8 FF © ££ #£ ££ £ €£ F&F £& & £F F 2 2 HO ££ ££ BB © + # & F

Two new species of Stylidium Willd. (Stylidiaceae) from north Queensland

A.R. Bean

* @ © © © BS &€& € © © & © &€& © #€& © &€ FF & & #& © #& FF & & &€£ & #& #8 #8 © #8 8 ££ BF #8 8 #& BB #£ BS & £ &£ ££ FF 8 # #8 # SB 2 2 Bb FE FF &F £F

A new species of Habenaria Willd. (Orchidaceae) from North Queensland

Petet Ss Lavarack: and Altick: We DOCK 0 a). dnee ace 0a font Mn celina koe wake dee ad

Oldenlandia gibsonii (Rubiaceae: Hedyotideae), a new species from south

east Queensland

D.A. Halford

a 5 5 + a 5 s & 4 * 5 a a + + a a | * a 5 + + ' | ¥ a a a + a a a a I ¢ +s &#® & &£© #& £ £ €& 8 8 8 FF FF EF 2 FF F&F FF FE F&F Ff

Jasminum jenniae (Oleaceae), a new species from south eastern Queensland

wayne harris 2 Crlonn HONG 300 5- josey wv rabaoa) Cle bits was eee en We dee et Re wre

The seedling of Cassytha glabella R.Br.

H. Trevor Clifford

= = © © &© © © &© &€© &© &§ & #8 & &€& © §&§ & &€§ FF BF * 8 8 #8 #£# # #£ 8 £ FF & #£ & ££ FB FF F&F ££ £ © 2 + # FF £ FF FF F HF F 2 F

A checklist of bryophytes of the wallum habitat of south-eastern Queensland

and north-eastern New South Wales

J. Windolf

= §« &© &© © & &© © & &© &© €& & & & © FS © € © FF © © F&F F&F BE F&F &€& FF F&F FF FF FF S&F OF FF FF FF © FE F FE ££ FF *# *#£ FF 2 FF £F FF BF FF HE FF F&F BS &

Vanguerieae A. Rich. ex Dum. (Rubiaceae) in Australia, |. Everistia

S.T.Reynolds & R.J.F.Hend.

S. Li Reynolds & Ri Jub. Henderson... cen eh eas oboe eee eee kbar Lites.

Note

Contribution of isozymic analysis in differentiating Macrozamia moorei

D.L.Jones and K.D.Hill from Mjohnsoniti F.Muell igual

Eis. Sharma, O.L. Jones &-Pil. FOrster’ jsc.ccs-.isie oa ete ie Kae eee ao ae ends

2 ATE SL Dnt ES Bane Sen at ee ee ae

A revision of the Babingtontia virgata (J.R.Forst. & G.Forst.)

F.Muell. complex (Myrtaceae) in Australia

A.R. Bean

Summary

Bean, A.R. (1999), A revision of the Babingtonia virgata (J.R.Forst. & G.Forst.) F.Muell. complex

(Myrtaceae) in Australia, Austrobaileya 5(2): 157-171. Seven Australian species related to Babingtonia

_yirgata are described as new viz. B. angusta, B. collina, B. crassa, B. brachypoda, B. bidwillii, B.

papillosa and B. similis, and one new combination is made, B. pluriflora. All species are described and

illustrated and notes are provided on their distribution, habitat and conservation status. Babingtonia

virgata is considered to be endemic to New Caledonia. A description of it is provided for comparative

purposes, and its relationship to Australian taxa is discussed. A revised key is provided to ali species of

Babingtonia from eastern Australia, and for the Australian members of the B. virgata complex.

Keywords: Myrtaceae, Baeckea, Babingtonia, Baeckea virgata, Babingtonia virgata, Babingtonia

angusta, Babingtonia collina, Babingtonia crassa, Babingtonia brachypoda, Babingtonia bidwilli,

Babingtonia papillosa, Babingtonia similis, taxonomy, keys, Australia, New Caledonia

A.R. Bean, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,

Toowong, Queensland 4066, Australia.

Introduction

The shrub now known as Babingtonia virgata

was first collected by the Forsters from the

island of New Caledonia in 1774, and later

named by them as Lepfospermum virgatum.

The species was transferred by the younger

Linnaeus to the genus Me/aleuca in 1781, and

then to Baeckea by Andrews (1810). In his

discussion, Andrews refers to the presence of

Baeckea species in New Holland (soon after

to become known as Australia), but did not

suggest the presence of B. virgata there.

However, specimen determinations at the

Herbarium, Kew, from that era suggest that that

was the popular opinion. Schauer (1843)

erected a new genus, Harmogia, to

accommodate B. virgata and several other

species, but did not comment on the

distribution of any of the species. Both Mueller

(1864) and Bentham (1867) ascribed B. virgata

to eastern Australia as well as to New

Caledonia. Mueller (loc. cit.) reduced his own

species Camphoromyrtus pluriflora F.Muell.

to Baeckea virgata, then later in the same

publication made the combination Babingtonia

virgata, which 1s accepted here. Bentham (loc.

cit.) established a broad generic and species

concept, reducing Harmogia and Babingtonia

Accepted for publication 25 August 1998

to sectional status under Baeckea, and included

a wide range of Australian and New Caledonian

forms under the name Baeckea virgata. Bailey

(1900) described a new variety, B. virgata var.

parvula (described here as Babingtonia

bidwillii), but no other taxa in the group have

since been described from Australia. Dawson

(1992) revised the New Caledonian members

of the Baeckea virgata group, and recognised

4 species in it. Babingtonia was reinstated by

Bean (1997) and revised for eastern Australia

and New Caledonia, with the exception of the

B. virgata complex which ts here treated.

B. virgata and its allies form a

taxonomically very difficult group as the

species involved are still apparently evolving,

and there is continuing exchange of genetic

material between populations/taxa in some

areas. This results in the blurring of species

boundaries, with some species pairs

intergrading over a distance of 50-100

kilometres. As a result, it is sometimes very

difficult to allocate some specimens to a

particular taxon.

Taxonomic and ecological characteristics

Within the genus Babingtonia, the Australian

members of the B. virgata group of species are

distinguished by the following combination of

158

characters: trunk fluted on large specimens;

leaves entire, thin-textured, with length more

than 2.5 times the width: inflorescences

usually 3 or 7 (occasionally 9-, rarely 1-)

flowered; ovules more than 15 per loculus,

radially arranged around the placenta.

Within the numbering

presented by Bean (1997), all of the

Australian taxa presented in this paper can

be placed between No. 5 (2. virgata) and No.

6 (B. tozerensis).

In Australia, species in the B. virgata

group may occupy mesic habitats on

creekbanks or as understorey in eucalypt

forests, or paradoxically they may occupy

xeric sites on rocky outcrops. In all cases the

soils are sandy or skeletal and the available

nutrients are comparatively low.

scheme

Austrobaileya 5(2): 157-171 (1999)

Methods

This study 1s based upon the examination of

herbarium material from A, BM, BRI, G, GH,

K, MEL, NE, NSW, HO, P, and WELTU. Stipe

and peduncle lengths were measured on

flowering material, as the peduncle (and

possibly the stipe) elongate after anthesis, and

measurements taken on fruiting material can

be quite different.

Measurements of leaves and branchlets

were taken from dried herbarium material:

branchlet descriptions are based on young

material (within 50 mm of shoot apices).

Measurements of floral parts are based either

on material preserved in spirit or material which

has been reconstituted by boiling.

All species have been examined in the field.

Key to the Babingtonia species of eastern Australia

1. Leafmargins crenulate or conspicuously ttregular 2.0... cee ee tees 2

Leaf margins entire or minutely denticulate.. 0.0... ee eee eee 3

2. Leaves 4.26.1 mm long, crenulate. Mt Buffalo, Victoria................. B. crenulata

Leaves 1.2—2.4 mm long, margins conspicuously irregular. Central-northern

INS Wee rae a) alta dive Ati 's Mouionon etal btea Cash "he ol a-feraled oats, shld ete wow td EN oil what A 3 B. cunninghamii

3, “Leaves nore than 15 mi Widens os a x0 te bacy ote oe Sie ee Ee Gan nee ees es ERE ES eh eT PY 4

LOAVES [GSS idan | Neti WAGE. Fe Ce iar ay cen aps tires faa ne gate CO arte orth, agra aatn oe Rae acing 12

4. Inflorescences 3- or 7-(9)-floweted.. 0... cc ee eee e eee e ees 5

INT OFESCEHGeS. [-FIGWETER a icc. sox Soto a/4 bv eh eonivtieg'ges (A pnece dail, Sow Oe a! eth aod Sony eb Maas I]

5. Leaf midrib impressed on upper surface; quadrangular branchlets with

undulating surface and crenate margins ...... 0... eee ees B. pluriflora

Leaf midrib not tmpressed on either surface; quadrangular branchlets smooth ..........

or papillose witht entire-marginss, 64 akon ce Ee ee ba ls Eee ole RR oe es 6

6. Oil glands on branchlets raised, papillose .... 0... eens 7

Oil glands on branchlets not raised or impressed. ..... 0... ee ee eee ee ee 8

7. Flower stipes 2—-3.5 mm long; petals 2.5-3 mmm across .......... 2.000 eae B. papillosa

Flower stipes 3.5—5 mm long; petals 3—3.5 mm acroOSS .......... 0.0.0 0e. B. tozerensis

8 Mostinflofescences S=1lowered tic i ek Roles wae oa ll ne ates le ctonhe BET Neneh es 9

' Mostantlorescences 7 SHOWEretl ... 5 yes bow cicy Asnctcace ht base 0 Gedun Srdte Ue grease 4: nh eee reeeansae acy os 10

9. Leaves obovate; flower stipes 1.2—-2.5 mm long ......... 0.0... eee eee. B. brachypoda

Leaves elliptical; flower stipes 2.5-4.5 mm long........ 0.0... cc eee B. bidwillii

10. Leaves 1.7—2.5 mm wide; bracts up to 1.4mm long ............ 2.00 cee ees B. collina

Leaves 2.5—3.5 mm wide; bracts 1-2.5mmlong........ 0... ccc cee ee B. crassa

Bean, A.R., A Revision of the Babingtonia virgata in Australia 159

11. Petals 2.6-3.0 mm wide; hypanthium muricate; leaf length/breadth ratio

Lira yu ace EE i gH BS eA TS Be TE pa A WECM SE AE AS co OE ee WENO 8 B. prominens

Petals 1.5-2.3 mm wide; hypanthium smooth; leaf length/breadth ratio |

Deals coca aot gt patra tare Se ae antes, ya ee tyaraner a. ttcnayoane Man oe yar teaeh | kee was cee, RU he ere B. silvestris

PZ SBA AD ER MNCTIT AUC 5 Se 5— ene weld See vei Ml eo cisabiecWecace ells dsopeney ebay jew Ay howled inate poe en B. behrii

Pear ADS x ACUIC OE ODTUSE yf cise wees e Weecl ake Vas Sov aN apse dk MBI s, EeoeAt i ume, 48, 5 ge 13

TS. dnflorescences (ls}5—7-TlOWered oo psc gee pa ace outage ba Rea Me dele aS eda pane 14

Inflorescences ‘consistently 1-TlOWereds ocd ere ne ne oe ke wae ea Wa ae ld be 1S

14. Leaves plano-convex to concavo-convex, 5.5—10 x 0.5-1.0 mm ............ B. angusta

Leaves ciate false 1 EAI go ake}, sack GA ed oo eva ead a ed rece: denies cages B. similis

15. Outer sepals acute or acuminate, 0.8—-1.8 mmlong......... ccc ce eee ee 16

Outer sepals obtuse, up to 0.3 mm long, or absent ........ 0. ce ee eee 18

16. Leaves broadly ovate, 0.9-1.5 mm long; hypanthium smooth............ B. squarrulosa

Leaves linear to lanceolate, 2.5—6.5 mm long; hypanthium smooth or muricate ....... 17

17. Stamens 5~9; ovules 8 or 9 per loculus; leaves 0.7—-1.4 mm wide .......... B. granitica

Stamens 11-13; ovules 12—14 per loculus; leaves 0.5—0.8 mm wide ...... B. odontocalyx

18. Leaves obovate; pedicels 2.0—4.2 mm long; outer sepals present;

Hy AAS PID SUL ores osu oe bere Snot Sours tener enact Writblen gap 'clalt deuphele git leas B. jucunda

Leaves linear; pedicels 0.4-1.0 mm long; outer sepals absent; hypanthium

SETEOON LUT: 3 gt oe pal se oo ty phe pao roe SST SO Wie lane Ut AYMAN Wig Meas tert Yee NG hoc B. densifolia

Isey to the Australian species allied to Babingtonia virgata

{. Leaf midrib tmpressed on upper surface; quadrangular branchlets with

undulating surface and crenate margins ....... 0.0.0 cee eee eee ees 4. B. pluriflora

Leaf midrib not impressed on either surface; quadrangular branchlets

smooth or papillose, with entire margins 2.0... 0.0... eee tees 2

2. Oil glands on branchlets raised, papillose ......... 000.0. cee eee eee 8. B. papillosa

Oil glands on branchlets not raised or impressed... 0.0... eee eee ees 3

3:,, - WioatranTO@resceneds Si Owvene | salen w teu Leliences whe Rcicannean Powvenae dopants eb acksy beaded s 4

lost nifloreseences*7= Brow ered. i: 4. <on-< senseless Since Vac te 54d wales arab bateen: Sacer eg Aa 7

4. Leaves obovate; flower stipes 1.2—-2.5 mm long ................004. 7. B. brachypoda

Leaves elliptical to linear; flower stipes 2.5-4.5 mmlong ..... 0.0... cc ee 5

5. Leaves plano-convex to concavo-convex, 0.5—-1.0 mm wide ............. 3. B. angusta

DSGAVCS Bat, Het TT ELE & aid ge 5k dts oh eh wav eace in be Bib WE oted gohed Wk Hee oe Baa beh BE od key Ble sal 6

6. Leaves 9-17 mm long; bracts up to 1.3mm long ........ 0... eee ee 2. B. similis

Leaves 4.5—6.5 mm long; bracts 1.2-2 mm long .......... 0.0.00. 0 cee 1. B. bidwillii

7. Leaves 1.7—2.5 mm wide; bracts up to 1.4mm long ............ 00000 ee. 6. B. collina

Leaves 2.5—3.5 mm wide; bracts 1-2.5 mm long.......... 000... 0.0 eee 5. B. crassa

160

Babingtonia virgata (J.R.Forst. & G.Forst.)

F.Muell., Fragm. 4: 74 (1864);

Leptospermum virgatum J.R.Forst. &

G.Forst., Char. Gen. Pl. 48 (1776);

Melaleuca virgata (J.R.Forst. & G.Forst.)

L.f., Supp. Pl. 343 (1781); Baeckea

virgata (J.R.Forst. & G.Forst.) Andrews,

Bot. Repos. 9: t. 598 (1810); Harmogia

virgata (J.R.Forst. & G.Forst.) Schauer,

Linnaea 17: 238 (1843). Type: New

Caledonia, in 1774, J.R. Forster & G.

Forster (holo: i).

Leptospermum parvulum Labill., Sert.

Austro-Caledon. 62, t. 61 (1825);

Baeckea parvula (Labill.) DC., Prodr. 3:

229 (1828). Type: New Caledonia, in

1793, A.J. Labillardiere (holo: FI).

Baeckea parvula var. latifolia Brongn. & Gris,

Bull. Soc. Bot. France 11: 184 (1864).

Type: Gatope, New Caledonia, Vieillard

514 (ecto: P, fide Dawson (1992)).

Baeckea obtusifolia Brongn. & Gris, Bull.

Soc. Bot. France 11: 185 (1864). Type:

Balade, New Caledonia, Vieillard 445 (P

n.v., photo BRI).

Shrub to 3 m high. Bark unknown. Branchlets

quadrangular, with convex surfaces, not

flanged, white or grey, margins entire or

sinuate; oil glands absent. Leaves petiolate;

petiole 0.9-1.6 mm long; lamina lanceolate,

oblanceolate or elliptical, 5-11 mm long, 1.0-

2.0 mm wide, concolorous, straight, flat or

longitudinally striate, not keeled, oil glands

obscure on both surfaces, 0.25—0.5 mm apart,

midrib rarely faintly visible on abaxial surface,

invisible on adaxial surface, intramarginal vein

not visible, apex obtuse or acute. Inflorescence

axillary, 3-(4-7)-flowered; peduncles 2.5—9.0

min long; stipes 1.5—5.0 mm long; bracts 2,

caducous, narrowly deltate, to 1.5 mm long,

acute; bracteoles stmilar but somewhat smaller.

Hypanthrum smooth, glandular, obconical, 1 .2—

1.5 mm long, fused to the ovary except at top;

calyx lobes simple; inner lobe sem1-orbicular,

c. 0.8 x 1.5 mm, thin or rather thick, margins

entire; outer lobe absent. Corolla up to 7 mm

across; petals broadly ovate to orbicular, 2.0—

3.0 x 1.9-2.2 mm, white, oil glands present;

margins entire. Stamens 7—11, in groups of 1-

Austrobaileya 5(2): 157-171 (1999)

3 opposite the calyx lobes, stamen opposite to

centre of calyx lobe often shorter than

remainder; filaments terete, 0.5—0.8 mm long,

geniculate, with brown connective gland fused

to upper part of filament at the bend; anthers

adnate, dehiscing by small divergent slits, with

loculi free. Style terete, up to 1.0 mm long after

anthesis, set into a pit; stigma broadly capitate.

Ovary 3-locular; floral disc flat; ovules 12—16

per loculus, arranged radially around placenta.

Fruit hemispherical, 1.5-1.8 <x 2.1-2.8 mm,

valves broadly deltate, not woody, exserted.

Seeds semi-discoid, c. 0.75 mm long, brown,

with flat sides and rounded backs, finely

reticulate (Fig. 1 T—V).

Selected specimens: New Caledonia. 29 km E of

Noumea on the road to Yate, Jan 1978, Armstrong 1175

(BRI, NSW); Barrage de la Dumbea, Nov 1977, Bamps

5727 US); Mt Tchingou, Aug 1965, Bernardi 10402 CX);

Col dAmoss, Dec 1977, Dawson s.n. (WELTU); Col de

Boghen, Dec 1952, Everist sn. (BRI); SW base of Mt

Dore, Sep 1963, Green 1129 (K); Mt Boulinda, Nov 1977,

Jaffre 2005 (K, P); Quen Toro hill, Noumea, Dec 1991,

Kelch 1666 (NSW); Isle of Pines, Oct 1853, AfacGillivray

(K); Sommet est de la Roche Ouaieme, Jul 1968, A¢cKee

19191 (BRI, k, P); Col de Tiebo, Sep 1973, AdeKee 27418

(K, P); Haute Koealagoguamba, Aug 1974, AdcKee 29057

(BRI, P); Mont Ouin, Aug 1974, AdeKee 29095 (K, NSW,

P); Balabio, Tiaodmoin, Sep 1974, A¢dcK ee 29294 (K, P);

Paagoumene, Feb 1980, AdcKee 37807 (WELTU); Mt

Koniambo, south of Voh, Oct 1982, AdePherson 4993

(MO, NSW); lower reaches of Dumbea Valley, Nov 1982,

McPherson 5207 (MO, NSW); Aut den Bergen bei

Oubatche, Nov 1902, Schlechter 15519 CX); base of Mont

Mou, Oct 1923, IVhite 2077 (KB).

Distribution and habitat: Endemic to New

Caledonia and a few adjacent islands. It grows

mostly on soils derived from schist or peridotite

but also on serpentine alluvium (Dawson 1992).

Phenology: Flowers are recorded between

August and February, while fruits are recorded

from December to August.

Notes: Babingtonia virgata 1s most closely

related to B. leratii (Schitr.) A.R.Bean and B.

procera (J.W.Dawson) A.R.Bean from New

Caledonia. It is less closely related to Australian

species and can be distinguished from them by

its thick and often longitudinally wrinkled

leaves (when dried) with obscure oil glands,

its convex branchlet mternodes lacking oil

glands, its shorter hypanthia, its simple calyx

lobes and its 12—16 ovules per loculus (16-23

for Australian taxa).

Bean, A.R., A Revision of the Babingtonia virgata in Australia 161

1, Babingtonia bidwillii A.R.Bean sp. nov.

affinis B. simili A.R.Bean a qua foliis

brevioribus et 2.5—4plo longioribus quam

latioribus, bracteis 1.2—-2.0 mm longis, et

petalis saepe fimbriatis differt. Typus:

Queensland. Wipe Bay District: Yurol

State Forest, 3 km north-west of Cooroy,

26 October 1993, A.R. Bean 6803 (holo:

BRI iso: L, MEL).

Baeckea virgata var. parvula F.M.Bailey,

Queensl. Fl. 2: 585 (1900), nom. inval.,

nom. nud. ?

Babingtonia sp. (Yurol A.R. Bean 6803) in

Henderson (1997).

Shrub or tree to 5 m high. Bark grey, persistent,

scaly to fibrous. Branchlets quadrangular with

flat sides, slightly flanged, brown, margins

entire; oil glands present, not papillose. Leaves

petiolate; petiole 0.6—1.0 mm long; lamina

elliptical to obovate, 4.5-7.0 mm long, 1.3—

2.0 mm wide, discolorous, straight, flat, not

keeled, oil glands prominent, especially on

lower surface, c. 0.25 mm apart, midrib faintly

visible on abaxial surface, not visible on adaxial

surface, intramarginal vein not visible, apex

obtuse or acute and abruptly narrowed at apex,

with a tiny caducous mucro. Inflorescence

axillary, 3-flowered, rarely 4—7-flowered;

peduncles 4.5—8 mm long; stipes 3.0-4.5 mm

long; bracts 2, caducous, linear, 1.2—2.0 mm

long, acute; bracteoles similar but somewhat

smaller. Hypanthium smooth, glandular,

obconical, 1.5—2.0 mm long, fused to the ovary

throughout; calyx lobes compound; inner lobe

seml-elliptic, c. 0.5 x 1.2 mm, thin, margins

entire or fimbriate; outer lobe rudimentary or

occasionally conspicuous, 0.4—2 mm long,

thick, erect, obtuse or acute. Corolla up to 8

mm across; petals broadly ovate to orbicular,

2.22.6 x 2,0—-2.5 mm, white, oil glands

present, margins entire or fimbriate. Stamens

(6)7-—-10, in groups of 1-3 opposite the calyx

lobes, stamen opposite to centre of calyx lobe

shorter than remainder; filaments terete, 0.6—

1.0 mm long, geniculate, with brown

connective gland fused to upper part of filament

at the bend; anthers adnate, dehiscing by pores,

with loculi fused. Style terete, up to 1.0 mm

long after anthesis, set into a pit; stigma broadly

capitate. Ovary 3-locular; floral disc concave;

ovules 16-18 per loculus, arranged radially

around placenta. Fruit hemispherical, 1.7—2.0

x 2.6-3.5 mm, valves broadly deltate,

chartaceous, at rim level or slightly exserted.

Seeds discoid, c. 0.6 mm long, brown, with flat

sides and rounded backs, minutely reticulate

(Fig. 1 C, D).

Specimens examined: Queensland. Porr Curtis

District: Shoalwater Bay Training Area, Site SW06,

E tributary of Werribee Ck, c. 1.5 km WSW of Mt

Carrol, Sep 1993, AfdeDonald 5732 & Scriffignano

(BRI); c. 22 km from Agnes Waters, S of Gladstone,

Nov 1976, Stanley 78183 & Ross (BRI). Wipe Bay

District: Yalanga station, Bates Road, NE of Kin Kin,

Dec 1994, Bean 8152 & Grimshaw (BRI); N.E.

Australia, 1848-53, Bidwill 102 (&); Wide Bay,

tropical New South Wales, 1848-53, Bidwil/ (GH);

Maryborough, Nov 1948, Clemens (BRI, GH, K);

Cooloola, near Noosa, Teewah Ck, Dec 1971, Harrold

C204 (BRI); Cooloola N.P., north-east of Banyan

Creek, Oct 1982, McDonald 3764 & Williams (BRI);

c. 1.5 km SW of Toogoom, Oct 1996, Sparshott

KMS1020 & Baumgartner (BRI); Cooloola N.P.,

Noosa River at Cooloola Way bridge, Noy 1993,

Telford 11981 & Nightingale (BISH, BRI, CANB,

NSW); Burrum River, undated, Watson s.n. (A); Noosa

River near Lake Como, Nov 1977, Williams 77272

(BRI). Moreton District: Lefoes Road, Bli Bli, Dec

1996, Bean 11544 (BRI, NSW),

Distribution and habitat: B. bidwillii 1s

found in coastal areas of Queensland from

Shoalwater Bay to just north of Brisbane (Map

1). It grows in deep sandy soil in eucalypt forest

of the coastal lowlands, often adjacent to areas

of heathland. Associated species include

Syncarpia glomulifera (Sm.) Nied. subsp.

glomulifera, Melaleuca sieberi Schauer ,

Eucalyptus resinifera sm., E. intermedia

R.T.Baker and Lophostemon suaveolens

(Gaertn.) Peter G. Wilson & J.T. Waterh.

Phenology: B. bidwillii flowers from October

to December, and fruits from December to

April.

Notes: B. bidwillii differs from B. collina

A.R.Bean by its leaves 4.5—7 mm long (6.5—

12.5 mm for B. collina) often with obtuse apex,

its bracts 1.2—2 mm long (up to 1.4 mm for B.

collina) and its mostly 3-flowered

inflorescences. It differs from B. similis

A.R.Bean by its shorter leaves which are 2.5—

4 times longer than wide (8—10 times for B.

similis), bracts 1.2—-2 mm long (up to 1.3 mm

long for B. similis) and the often fimbriate

162

fo |

cH 4 Hy THAT

OER ETT TH

145 i350 [55

Map 1. Distribution of Babingtonia pluriflora ©, B. similis

fs, B. papillosa ®.

petals (entire for B. similis). Intergrades with

B. collina occur north of Brisbane.

Conservation status: This taxon is not

considered rare or threatened.

Etymology: The species epithet honours John

Carne Bidwill (1815-1853), who collected

the first known specimen of this species.

2. Babingtonia similis A.R.Bean sp. nov.

affinis B. angustae A.R.Bean a qua foltis

latioribus planisque, et stipitibus 2.5—4.0

Austrobaileya 5(2): 157-171 (1999)

mm longis differt. Typus: Queensland.

Moreton District: Springbrook, south-

west of Mudgeeraba, 16 January 1994,

A.R. Bean 7314 (holo: BRI; iso: DNA,

NSW).

Babingtonia sp. (Yatala P. Grimshaw+

G525) in Henderson (1997).

Shrub to 2 m high. Bark grey, persistent, scaly

to fibrous. Branchlets quadrangular with flat

sides, not flanged, grey, margins entire; oil

glands present, not papillose. Leaves petiolate;

petiole 0.6—1.0 mm long; lamina narrowly

lanceolate, 9-15 mm long, I.1-1.5 mm wide,

discolorous, straight, flat, not keeled, oil glands

prominent, especially on lower surface, c. 0.25

mm apart, midrib faintly visible on abaxial

surface, invisible on adaxial surface,

intramarginal vein not visible, apex acute.

Inflorescence axillary, 3- flowered; peduncles

5.0-9.0 mm long; stipes 2.5—4.0 mm long;

bracts 2, caducous, narrowly deltate, to 1.3 mm

long, acute; bracteoles similar but somewhat

smaller. Hypanthium smooth, glandular,

obconical, 1.5—2.0 mm long, fused to the ovary

except at top; calyx lobes compound; inner

lobe semi-elliptic or deltate, c. 0.7 1.5 mm,

thin, margins mostly fimbriate; outer lobe

rudimentary, c. 0.6 mm long, thick, obtuse.

Corolla up to 7 mm across; petals broadly ovate

to orbicular, 2.0—2.5 x 1.8-2.5 mm, white, oil

glands present, margins entire. Stamens 8—

LO, in groups of I1—3 opposite the calyx lobes,

stamen opposite to centre of calyx lobe

shorter than remainder; filaments terete, 0.7—

1.0 mm long, geniculate, with brown

connective gland fused to upper part of

filament at the bend; anthers adnate,

dehiscing by small divergent slits, with locul1

free. Style terete, up to 1.0 mm long after

anthesis, set into a pit; stigma broadly

capitate. Ovary 3-locular; floral disc

concave; ovules 16-18 per loculus, arranged

radially around placenta. Fruit hemispherical,

c. 1.7 x 3.0 mm, valves broadly deltate,

somewhat woody, exserted. Seeds sem1-

discoid, c. 0.75 mm long, brown, with flat sides

and rounded backs, minutely reticulate (Fig. 1

R, S).

Specimens exantined: Queensland. Moreton District:

Springbrook-Mudgeeraba road, Jan 1994, Bean 7319 (BRD;

Bean, A.R., A Revision of the Babingtonia virgata in Australia 163

145 {50

eT PaaS

sereesaesen)

BRE CT * 40

KERR

HEEEPCTEHE

145 150 {55

Map 2. Distribution of Babingtonia bidwillii A, B. crassa

©, B. brachypoda *.

near defunct Lion Park, Pacific Highway, Yatala, Mar 1994,

Grimshaw G525 & Gibbs (BRD; Oxenford, S of Brisbane,

Aug 1930, Hubbard 3690 (6); Beechmont road, 12 km S

of Nerang, Feb 1979, Olsen 826 & Lebler (BRI; 3 km S of

Nerang along road to Beechmont, Aug 1985, Reynolds &

Cahvay sn. (BRD; along Little Nerang Creek on road to

Springbrook, Apr 1959, Thorne 20481 (BRI). New South

Wales. Nortu Coast: Laurieton, Mar 1917, Baker s.n.

(NSW); New Italy, Nov 1895, Bauerlen s.n. (NSW);

Tooloom Falls, c. 5 km SSW of Urbenville, Nov 1987,

Coveny 12806 et al. (BRI, MEL, NSW); Blandford Ck,

Boundary Creek S.F., Feb 1979, Floyd AGF1210 (NSW);

Beechwood, Hastings River, May 1915, Adaiden s.n. (NSW);

Black Hull, between Maitland and Newcastle, Jun 1979,

Martin s.n. (NSW); banks of Tooloom Ck, Urbenville, May

1945, White 12775 (A).

Distribution and habitat: B. similis extends

from the Brisbane area in south-eastern

Queensland to near Newcastle in New South

Wales (Map 2). It occurs in a wide range of

habitats including Me/laleuca-dominated open

forest and eucalypt forest, and on rainforest

margins with Callicoma serratifolia Andrews

and Acrotriche sp.

Phenology: Flowers are recorded between

January and March, while fruits are recorded

from January to August.

Notes: B. similis is most closely related

to B. angusta but differs from that by its

discolorous leaves which are flat in cross-

section (concolorous and plano-convex for B.

angusta), and 1.0~-1.5 mm wide (0.5—1.0 mm

wide for B. angusta), and the flower stipes 2.5—

4.0 mm long (1-3 mm for B. angusta).

Intergrades may occur with B. angusta in some

parts of northern New South Wales, with 2.

collina south of Brisbane, and with B. pluriflora

around the Port Stephens—Newcastle area.

Conservation status: This taxon is not

considered rare or threatened.

Etymology: The specific epithet is from the

Latin similis, meaning like, resembling, similar;

in reference to the similarity between this

species and B. angusta.

3. Babingtonia angusta A.R.Bean sp. nov.

affinis B. simili A.R.Bean a qua foltis

concoloris planoconvexis 0.5—1.0 mm latis

et stipitibus |~3 mm longis differt. Typus:

New South Wales. NorTH Coast: 7 km SE

of Coutts Crossing, 2 February 1995, A.R.

Bean 8321 (holo: BRI; iso: A, CANB, K,

MEL, NSW, P, PERTH, distribuend)).

Babingtonia sp. (Atherton A.R. Bean 5707)

in Henderson (1997).

Baeckea sp. “Clarence River” in Elliot and

Jones (1982).

Shrub to 2.5 m high. Bark grey, persistent, scaly

to fibrous. Branchlets quadrangular with

slightly convex surfaces, not flanged, grey,

margins entire; oil glands present, not papillose.

Leaves petiolate; petiole 0.6—1.2 mm long;

lamina narrowly-oblanceolate to linear, 5.5—10,0

164

mm long, 0.5—-1.0 mm wide, concolorous,

straight, plano-convex to concavo-convex, not

keeled, oil glands visible on both surfaces, c.

0.5 mm apart, midrib not visible on either

surface, mtramarginal vein not visible, apex

acute, acuminate or uncinate. Inflorescence

axillary, 1—3 flowered; peduncles 2.5~7.5 mm

long; stipes 1.0—3.0 mm long; bracts 2,

caducous, narrowly deltate, 0.75—1.0 mm long,

acute; bracteoles similar but somewhat smaller.

Hypanthium smooth, glandular, obconical, |.5—

2.0 mm long, fused to the ovary except at top;

calyx lobes compound; inner lobe oblong to semi-

elliptic, c. 0.7 x 1.5 mm, thin, margins entire or

denticulate; outer lobe rudimentary, 0.4-0.7 mm

long, thick, erect, obtuse. Corolla up to 8 mm

across; petals orbicular, 2.0-2.8 x 2.0-2.7 mm,

white, oil glands present, margins entire. Stamens

8—13, in groups of 1-3 opposite the calyx lobes,

stamen opposite to centre of calyx lobe shorter

than remainder; filaments terete, 0.7—1.1 mm

long, geniculate, with brown connective gland

fused to upper part of filament at the bend; anthers

adnate, dehiscing by pores, with loculi fused. Style

terete, up to 1.0 mm long after anthesis, set into a

pit; stigma broadly capitate. Ovary 3-locular;

floral disc concave; ovules 17-20 per loculus,

arranged radially around placenta. Fruit

hemisherical, 1.8—2.2 x 3.0-3.5 mm, valves

broadly deltate, not woody, atrim level or slightly

exserted. Seeds D-shaped, c. 0.6 mm long, brown,

with flat sides and rounded backs, minutely

papillose (Fig. 1 E, F).

Speciinens examined: Queensland. Cook DIstRIcT:

Carrington Falls, SSW of Atherton, Jan 1993, Bean 5707

(BRI, CANB, K, L, MEL); powerline access road near

Herberton, Nov 1995, Ford 1671 (QRS); Herberton Wet,

Dec 1993, Forster PIF14481 (BRI, MEL, QRS). Norrr

KENNEDY District: Stony Creek, c. 2 km upstream from

Wallaman Falls, W of Ingham, Jan 1997, Bean 11595

(BRI, NSW, QRS); Blencoe Falls, 30 miles [48 km] W of

Cardwell, Nov 1967, Boyland & Gillieatt 583 (BRI, K);

Koombooloomba area, Dec 1964, Brooks s.n. (BRD);

Bluewater Creek near Bluewater, c. 25 km N of

Townsville, 6 km from coast, Dec 1983, Cattle s.n. (BRI);

Cameron Creek, on track east from Glen Ruth homestead,

between Cardwell and Mt Garnet, Dec 1993, Cumming

12565 (BRI, MEL); 17 km past Paluma on road to Hidden

Valley, Jan 1992, Forster PIF9476 (BRI, DNA, MEL,

PERTH); Nitchaga Creek, 6 km S of Tully Fails, Dec

1993, Forster PIF14476 (BRI, MEL, QRS); “Taravale”

near Hellhole Creek, 0.5-1.5 km E of homestead, Mar

1987, Jackes 8754 (BRI, CANB); Blencoe Ck, Nov 1975,

Travers C17 (A, BRI, K). Moreton District: Rocky

Creek, Mt Barney N.P., Jul 1994, Bean 7762 & Halford

(BRD; Mt Alford-Moogerah Dam road, just N of Mt

Austrobaileya 5(2): 157-171 (1999)

Alford, Mar 1994, Grimshaw G534 (BRI, NSW). DaRLING

Downs District: Portion 90, Wyberba, near Girraween

N.P., Sep 1993, Bean 6398 & Forster (BRI, NSW);

Dalveen, Dec 1962, Ped/ey 1168 (BRI, NSW). New South

Wales. Nortu Coast: Bean Creek Falis, 15 km S of

Urbenville, Dec 1993, Bean 7235 (BRI, NSW); Hortons

Creek, on Grafton-Armidale road, Apr 1994, Bean 7664

(AD, BRI, MEL, NSW); Nymboida River crossing, 5 km

S of Nymboida, Apr 1994, Bean 7671 (BRI, NSW); near

Sherwood, Oct 1981, Coveny & Armitage (NSW).

Distribution and habitat: B. angusta occurs m

north Queensland between Atherton and

Townsville, and also in south-eastern

Queensland e.g. near Boonah and Stanthorpe,

145 150

ree ae =I

Sel

RAE

CEHEA

145

Map 3. Distribution of Babingtonia angusta G.

Bean, A.R., A Revision of the Babingtonia virgata in Australia 165

and in north-eastern New South Wales as far

south as Sherwood (near Kempsey) (Map 3).

B. angusta inhabits rocky sites on forested

hillsides, or near (but not on) watercourses.

Altitudes range between 200 and 1050 metres.

Phenology: Flowers have been recorded from

November to March (with most from December

to February) and fruits are recorded from

March to September.

Notes: B. angusta is distinguished by its very

narrow leaves, which are less than 10 mm long.

A selected form of it has been cultivated in eastern

Australia for several years as Baeckea sp.

“Clarence River’. B. angusta is most closely

related to B. similis (see notes under that species),

and intermediates may occur in some areas.

Conservation Status: This taxon 1s not

considered rare or threatened.

Etymology: The species epithet is from the

Latin angustus Meaning narrow, in reference

to the leaves of this species.

4. Babingtonia pluriflora (F.Muell.)

A.R.Bean comb. nov.

Camphoromyrtus pluriflora F.Muell.,

Trans. & Proc. Victorian Inst. Advancem.

Sci. (1855). Type: Victoria. Tambo River,

February 1855, # Mueller s.n. (ecto:

MEL [MEL 73108] (here chosen);

isolecto: BM, G, K).

Shrub to 4m high. Bark grey, persistent, scaly

to fibrous. Branchlets quadrangular with

undulate surfaces, slightly flanged, white to

pink, margins crenate; oil glands present, not

paptilose. Leaves petiolate; petiole 0.8—1.5 mm

long; lamina lanceolate or elliptical, 10-29 mm

long, 2.5—-6.0 mm wide, discolorous, straight,

flat, not keeled, oil glands prominent, especially

on lower surface, c. 0.25 mm apart, equally

numerous but less prominent on upper surface,

midrib visible on abaxial surface, impressed on

adaxial surface, intramarginal vein sometimes

visible, apex acute or obtuse. Inflorescence

axillary, 3—7(9)- flowered; peduncles 5.0—13.0

mm long; stipes 3.0-7.0 mm long; bracts 2,

caducous, linear, c. 1.5 mm long, acute;

bracteoles similar but somewhat smaller.

Hypanthium smooth, glandular, obconical, 1.7—

2.2 mm long, fused to the ovary except at top;

calyx lobes compound; inner lobe sem1-elliptic,

c, 0.6 x 1.5 mm, thin, margins entire; outer lobe

rudimentary, rarely conspicuous, 0.3—1.5 mm

long, thick, erect, acuminate or obtuse, usually

not exceeding inner lobe. Corolla up to 10 mm

across; petals broadly ovate to orbicular, 2.5—

3.7 x 2.0-3.5 mm, white, oil glands present,

margins entire. Stamens 8—15, in groups of |-

4 opposite the calyx lobes, stamen opposite to

centre of calyx lobe shorter than remainder;

filaments terete, 0.6—1.0 mm long, geniculate,

with pale brown connective gland fused to

upper part of filament at the bend; anthers

adnate, dehiscing by short slits, with loculi free.

Style terete, up to 1.0 mm long after anthesis,

set into a pit; stigma broadly capitate. Ovary

3~-locular; floral disc concave; ovules 16—23 per

loculus, arranged radially around placenta.

Fruit hemispherical, 1.72.0 x 2.5~3.5 mm,

valves broadly deltate, not woody, enclosed or

at rim level. Seeds D-shaped, c. 0.75 mm long,

brown, with flat sides and rounded backs,

minutely papillose (Fig. | K—O).

Selected specimens: New South Wales. Norru Coast:

Port Stephens, Aug 1911, Boorman s.n. (BRI, NSW).

CENTRAL TABLELANDS: Moorara Boss Hiil, Mt Dunn road,

5.1 km NE of junction with Kananera road, Nov 1985,

Benson 2413 & Keith (NSW); Blue Mtns, undated,

Cunningham s.n, CX). CENTRAL Coast: The Peaks,

Burragorang, Aug 1905, Cambage 1290 (NSW); Grose

River, Sep 1906, Maiden & Cambage s.n. (NSW);

causeway on Glenbrook Ck, Jun 1952, Whaite 1162

(NSW). SouTHERN TABLELANDS: Correa Creek, Bolero

Creek, Sep 1898, Baeuerien (NSW). SoutH Coast:

Nadgee Nature Reserve, Newtons Beach, Jan 1985,

Albrecht 1507 (MEL, NSW); Araluen Valley, 8 mties [13

km] NW of Moruya, Dec 1961, Briggs s.n. (NSW);

Bodalla S.F., Sep 1953, Constable 26515 (K, NSW);

Woodburn S.F. on road to Pigeon House Mtn, WSW of

Burrill Lake, Oct 1981, Coveny 11012 & James (BRI,

NSW, PERTH); Araluen Valley, 7 mies [11 km] west of

Moruya on the Moruya-Araluen road, Dec 1967, Evans

2770 (A, K, NSW); Narrabarba Creek crossing on

Wonboyn road, c. 20 km SSW of Eden, Feb 1979, Haegi

1704 (BRI, K, MEL, NSW); Twofold Bay, Jan 1953,

Melville 2726 & Wakefield (A, K, MEL, NSW). Victoria.

Mt Dawson, north of Buchan, Mar 1984, Albrecht 364

(MEL, NSW); 15 miles [24 km] WNW of Bairnsdale,

Jan 1960, Aston 508 (A); 19.2 km S of Gelantipy, on

road to Buchan, Dec 1995, Bean 9432 & Jobson (BRI,

MEL); Errinundra Plateau, near mntersection of Helmers

Rd with Errinundra Rd, Jan 1993, Fletcher 135

(MEL);Double Creek nature walk, 6.7 km NW of

Mallacoota, Dec 1983, Parkes EG60b (CANB, MEL,

PERTH); Little River Gorge lookout, NE of Butchers

Ridge, Apr 1984, Parkes 214 (MEL).

166

Distribution and habitat: Babingtonia

pluriflora is common along the coast and

adjacent ranges from around Port Stephens

in New South Wales to the Mitchell River

(near Bairnsdale) in eastern Victoria (Map

1). It most commonly grows in eucalypt

forests close to watercourses in deep sandy

soils. However, it can sometimes grow on

rocky outcrops where tree cover is Sparse or

absent, and where there is little or no soil. It

occurs mostly at altitudes of 5 to 200 metres,

but occasionally occurs as high as 800

metres. Some recorded associated species

include Eucalyptus sieberi L.A.S. Johnson,

LE. gummifera (Gaertn.) Hochr., Aunzea

ericoides (A.Rich.) Joy Thomps., Kunzea

ambigua (Sm.) Druce and Banksia

integrifolia L.Ff.

Phenology: The main flowering period is

from October to January, but some flowering

also takes place in the April to July period.

Fruits are recorded from January to April.

Notes: B. pluriflora varies greatly in leaf size

and shape, and grows in a wide range of

habitats. Despite this, it has not been possible

to recognise more than one taxon as no other

characters consistently correlate with leaf

dimension or habitat. Itis closest to B. similis

and apparently intergrades with that in the

Port Stephens-Newcastle area. B. pluriflora

differs trom #. similis by its broader leaves

with the midrib impressed on the adaxial

surface, its crenate inflated branchlet

internodes, its longer petals, longer

hypanthium and fruiting valves enclosed or

at rim level.

Specimens from Little River gorge and Mt

Dawson in eastern Victoria have long

acuminate outer calyx lobes, but in other

respects are representative of B. pluriflora.

Specimens from the Budawang Range in

southern New South Wales (e.g. Telford

8833, 8905) have very short leaves and

pedicels, and may prove to be taxonomically

distinct.

Conservation Status: This taxon is not

considered rare or threatened.

5. Babingtonia crassa A.R.Bean sp. nov.

affinis B. collinae A.R.Bean a qua foltis

Austrobaileya 5(2): 157-171 (1999)

2.8—3.6 mm latis, hypanthio longiore,

lobis exterioribus calycis lobos interiores

excedentibus, et fructibus majoribus

differt. Typus: New South Wales.

NoRTHERN TABLELANDS: Dangar’s Falls,

20 km S of Armidale, 31 January 1995,

A.R. Bean 8289 (holo: BRI; iso: K, MEL,

NE, NSW).

Shrub to 2.5 m high. Bark grey, persistent, scaly.

Branchlets quadrangular with flat sides, not

flanged, grey, margins entire; o1l glands present,

not papillose. Leaves petiolate; petiole 1.0—1.5

mm long; lamina elliptical to lanceolate, 7-13

mm long, 2.5—3.5 mm wide, concolorous,

straight, with internodes flat, not keeled, oil

glands prominent on both surfaces, c. 0.25 mm

apart, midrib visible on abaxial surface, not

visible on adaxial surface, intramarginal vein

not visible, apex acute. Inflorescence axillary,

(3)7—9 flowered; peduncles 7-11 mm long;

stipes 2—4 mm long; bracts 2, caducous,

lanceolate, 1.0—2.5 mm long, acute; bracteoles

similar but somewhat smaller. Hypanthium

smooth, densely glandular, obconical, 2.0-2.5

mim long, fused to the ovary except at top; calyx

lobes compound; inner lobe semi-elliptic, c. 0.4

x 1.5 mm, thin, margins entire; outer lobe

rudimentary, c. | mm long, thick, erect, acute.

Corolla up to 7 mm across; petals orbicular,

2.0—2.8 x 2.0—2.8 mm, white, oil glands

present, margins entire. Stamens 8-11, in

groups of 1-3 opposite the calyx lobes, stamen

opposite to centre of calyx lobe shorter than

remainder; filaments terete, 0.5—1.0 mm long,

geniculate, with brown connective gland fused

to upper part of filament at the bend; anthers

adnate, dehiscing by short slits, with locul free.

Style terete, up to | mm long after anthesis, set

into a pit; stigma broadly capitate. Ovary 3-

ocular; floral disc concave; ovules 18—20 per

loculus, arranged radially around placenta.

Fruit campanulate to hemispherical, 2.53.0 x

3.54.0 mm, valves broadly deltate, not woody,

at rum level. Seeds not seen (Fig. 1 P, Q).

Specimens examined: New South Wales. NorTHERN

TABLELANDS: Church Point, Woilomombi Falls, Feb 1979,

Floyd 1207 (BRI, NE, NSW); Carrai Plateau edge, Sep

1984, King 353 (NSW); Dangar’s Falls, Jan 1883, Statter

s.n. (BM). Norrn Coast: McLeay River, undated, Beck/ler

s.n. (K); Upper Carrow Brook, on Mount Royal road, c.

2 km below Cassel’s road turnoff, Feb 1985, Faullding

31 (NSW).

Bean, A.R., A Revision of the Babinetonia virgata in Australia 167

Distribution and habitat: B. crassa is confined

to a small area on the eastern edge of the

northern tablelands of New South Wales, from

east of Armidale to the Barrington Tops area

(Map 2). It grows on rocky sites with little or

no soil, sometimes on very steep slopes.

Associated species include Leptospermum

petersonii F.M.Bailey, Callistemon sieberi DC.

and Acacia ingramii Tindale.

Phenology: Flowers have been recorded in

January and February; fruits in February.

Notes: B. crassa 1s related to B. collina, but

differs by its leaves 2.5-3.5 mm wide (1.7-2.5

mim for B. collina), outer calyx lobes exceeding

inner lobes, hypanthium 2~—2.5 mm long (1.5-

2mm for B. collina) and fruits 2.5—-3 x 3,5—4

mm (c. 1.5 x 3 mm for B. collina). No

intergrades with any other species are known.

Conservation status: The risk category of

Babingtonia crassa according to the criteria

of Chalson & Keith (1995) is “Priority for

Investigation” (criterion a).

Etymology: The specific epithet is from the

Latin crassus, meaning thick, 1n reference to

the leaves of this species.

6. Babingtonia collina A.R.Bean sp. nov.

affinis B. bidwillii A.R.Bean a qua foltis

longioribus, inflorescentiis plerumque 7-

floris, et bracteis plerumque longioribus

differt. Typus: Queensland. Moreton

District: Marstaeller Rd, Karana Downs,

1 km 8S of Mt Crosby, 26 April 1995, 1.77.

Bird & C. Hays |AQ 635743] (holo: BRI;

iso: AD, DNA, MEL, NSW).

Babingtonia sp. (Mt Crosby L.H. Bird+

AQ635744) in Henderson (1997).

shrub to 3 m high. Bark grey, persistent, scaly to

fibrous. Branchlets quadrangular with flat sides,

not flanged, grey, margins entire; oil glands

present, not papillose. Leaves petiolate; petiole

Q.6—-1.3 mm long; lamina lanceolate, 6.5—12.5 mm

long, |.7—2.5 mm wide, slightly discolorous or

concolorous, straight, flat, not keeled, oil glands

prominent, especially on lower surface, c. 0.25

mim apart, midrib not visible on either surface,

intramarginal vein not visible, apex acute, not

abruptly narrowed at apex. Inflorescence axillary,

(3)-7 flowered; peduncles 5.0—9.5 mm long;

stipes 2.5—4.0 mm long; bracts 2, caducous,

deltate, to 1.4 mm long, acute; bracteoles similar

but somewhat smaller. Hypanthium smooth,

glandular, obconical, 1.5—2.0 mm long, fused to

the ovary except at top; calyx lobes simple; inner

lobe serm1-elliptic, c. 0.6 x 1.5 mm, thick or thin,

margins entire; outer lobe absent. Corolla up to 8

mm across; petals orbicular, 2.0-2.8 x 2.0-2.8

mm, white, oil glands present, margins entire.

Stamens (7)8—11, in groups of 1—3 opposite the

calyx lobes, stamen opposite to centre of calyx

lobe shorter than remainder; filaments terete, 0.6—

1.0mm long, geniculate, with brown connective

gland fused to upper part of filament at the bend;

anthers adnate, dehiscing by short divergent slits,

with loculi fused. Style terete, up to 1.0 mm long

after anthesis, set into a pit; stigma broadly

capitate. Ovary 3-locular; floral disc concave;

ovules 16-18 per loculus, arranged radially

around placenta. Fruit hemispherical, c. 1.5 x 3.0

mm, valves broadly deltate, not woody, exserted

or at rim level. Seeds semi-discoid, c. 0.75 mm

long, brown, with flat sides and rounded backs,

minutely reticulate (Fig. 1 G, H).

Selected specimens: Queensland. NortH KENNeby District:

Frederick Peak, 25 km SW of Townsville, May 1991, Bean

3206 (BRI, PERTH); summit of Mt Aberdeen, Aug 1993,

Bean 6316 (BRI, MEL). Souru KENNepy District: Mount

Jukes N.P., c. 30 km NW of Mackay, May 1991, Bean 3186

(BRI). Port Curtis District: Mount Wheeler, 12 km SW of

Yeppoon, Aug 1991, Batianoff WH9108022 & Robins (BRI,

NSW); Mt Marta, c. 65 km NW of Bundaberg, Nov 1993,

Bean 6987 (BRI). Burnetr District: summit of Coongara

Rock, S.F. 1344, Gayndah shire, Mar 1994, Thomas COO4

(BRI). Wipe Bay District: Burnett River, 20 miles [32 kn]

W of Bundaberg, May 1983, Jansen 69 (BRI); Kenilworth

Bluff, c. 8 km N of Kenilworth, Apr 1987, Sharpe 4662 &

Bean (BRI). DARLING Downs District: Severn River gorge

between the falls and Low’s Waterholes, Apr 1988, Thomas

283 (BRD). Moreton District: Kureelpa Falls, 8 km W of

Nambour, Jun 1995, Bean 8728 (BRI, MEL); 3.7 km E of

Canungra, Jan 1995, Bean 8188 (BRI, MEL, NSW, QRS);

Stumers road 1 km W of Mt Crosby, Apr 1995, Bird &

Hays s.n. (BRI, DNA, NSW); Mt Blaine, 25 km S of Ipswich,

Peak Crossing area, Apr 1993, Birds.n. (BRI, NSW); Diana’s

Bath, D’Aguilar Range near Mt Byron, Apr 1995, Forster

PIF16403 (BRI, MEL, NSW); between Brisbane and

Redcliffe, Dec 1930, Hubbard 5510 (K, P); Aspley near

Brisbane, Dec 1930, White 7147 (A, BRI, K). New South

Wales. Nortu WEsT Stopes: Severn river, c. 4 km upstream

of confluence with Macintyre River, Jul 1991, Coveny 1441]

& Makinson (BRI, CANB, MEL, NSW, PERTH). Norra

Coast: Rocky Creek, 28 km from Grafton towards Coaldale,

Feb 1995, Bean 8340 (BRI, MEL, NSW); Copmanhurst,

Nov 1917, Cheel s.n. (BRI, NSW); Codhole, Nymboida R.,

23 km N of Dorrigo, Dec 1977, Floyd AGF826 (NSW).

168

Distribution and habitat: The main distribution

of B. coflina is from around Yandina in southern

Queensland to Dorrigo in New South Wales;

however it extends in isolated populations to as

far north as Frederick Peak near Townsville (Map

4), Altitudes are mostly 50 to 500 metres, but in

north Queensland it reaches 900 metres. It may

grow in shrubland on shallow soils derived from

sandstone, rhyolite or granite, in eucalypt forest

with shallow sandy soils, or in riparian

communities, especially where there is

outcropping rock.

Map 4. Distribution of Babingtonia collina @.

Austrobaileya 5(2): 157-171 (1999)

Phenology: Flowers may be found between

November and March, and sometimes as late

as May. Fruits occur from January to August.

Notes: B. collina \s related to both B. similis and

B. bidwillii (see notes under those species).

Collections of B. collina from Redcliffe, Aspley

and Petrie in south-eastern Queensland tend

towards B. bid\villii mn leaf dimensions and flower

number. B. colling and B. similis apparently

intergrade in the area south of Brisbane.

Conservation Status: This taxon is not

considered rare or threatened.

Etymology: The species epithet 1s from the

Latin col/inus, relating to hills, in reference to

the usual habitat of the species.

7. Babingtonia brachypoda A.R.Bean sp. nov.

affinis B. collinge A.R.Bean a qua folts

oblanceolatis apice obtusis, inflorescenttis

3-floris, strpitibus 1.2-2.5 mm longis et

petalis fimbriatis differt. Typus:

Queensland. LEICHHARDT DISTRICT:

“Humboldt”, 45 km north-east of

Rolleston, 26 January 1996, A.R. Bean

9541 (holo: BRI; iso: CANB, K, L, MEL,

NSW, distribuend)).

Babingtonia sp. (Comet P. Rowland

AQ634382) in Henderson (1997).

Shrub to 4 m high. Bark grey, persistent, scaly to

fibrous. Branchlets quadrangular with flat sides,

not flanged, grey, margins entire; oil glands

present, not papillose. Leaves petiolate; petiole

0.71.3 mm long; lamina obovate, 5.5~9.0 mm

long, 1.8—2.8 mm wide, concolorous, straight, flat,

not keeled, oi] glands prominent, especially on

lower surface, c. 0.25 mm apart, midrib not visible

on either surface, intramarginal vein not visible,

apex obtuse or with a tiny caducous mucro.

Inflorescence axillary, 3-flowered; peduncles 3—

6 mm long; stipes 1.2—2.5 mm long; bracts 2,

caducous, narrowly deltate, |.1—1.4 mm long,

acute; bracteoles similar but somewhat smaller.

Hypanthium smooth, glandular, obconical, 1.5—

1.8 mm long, fused to the ovary except at top;

calyx lobes simple; inner lobe sem1-elliptic, c. 0.7

x 1.5 mm, thin, margins entire or fimbriate; outer

lobe absent. Corolla up to 8 mm across; petals

orbicular, 2.2—2.5 x 1.8-2.5 mm, white, oil

¢ Chen ha eR td Ra edit eaaiedaie ie TALE iv iastanantttl i. be tok pute

Bean, A.R., A Revision of the Babingtonia virgata in Australia 169

glands present, margins fimbriate. Stamens 9—

12, in groups of 1-3 opposite the calyx lobes,

stamen opposite to centre of calyx lobe shorter

than remainder; filaments terete, 0.4-0.8 mm

long, geniculate, with brown connective gland

fused to upper part of filament at the bend;

anthers adnate, dehiscing by short divergent

slits, with loculi fused. Style terete, up to 1 mm

long after anthesis, set into a pit; stigma broadly

capitate. Ovary 3-locular; floral disc concave;

ovules 16-18 per loculus, arranged radially

around placenta. Fruit hemispherical, c. 2.0 x

3.5 mm, valves broadly deltate, somewhat

woody, exserted. Seeds D-shaped, c. 0.8 mm

long, brown, with flat sides and rounded backs,

minutely reticulate (Fig. 1 A, B).

Specimens examined: Queensland. LEICHHARDT

Disrrict: “Humboidt’, 45 km north-east of Rolleston,

Jan 1996, Bean 9543 (BRI, MEL); c. 30 km NW of

Woorabinda, Henry Creek, May 1996, Brushe JB206

(BRI); “Humboldt” S of Blackwater, Jan 1997,

Fensham 3002 (BRI); Precipice NP, Cables Ck

catchment, Sep 1996, Forster 19788 (BRD; “Apis

Creek”, W of Marlborough, Mar 1993, Fensham 1137

(BRI); “Humboldt” via Comet, Feb 1995, Rowland s.n. (BRD.

Distribution and habitat: B. brachypoda is

known from a few sites near the towns of

Rolleston, Woorabinda and Theodore (Map 2),

on sandstone gullies or on the sandy alluvials

adjacent to sandstone ranges.

Phenology: Flowers have been collected in

January and March, and fruits in January.

Notes: B. brachypoda is closely related to B.

collina, but differs from that by the obovate

leaves with obtuse apex, 3-flowered

inflorescence (mostly 7-flowered for B.

collina), fimbriate petal margins and stipes 1.2—

2.5 mm long (2.5—4 mm long for B. collina).

No intergrades with any other species are

known.

Conservation status: This taxon is not

considered rare or threatened.

Etymology: The species epithet is derived from

the Greek “brachy-” short and “-podus” footed,

based; in reference to the short flower stipes of

this species.

8. Babingtonia papillosa A.R.Bean sp. nov.

affinis B. tozerensi A.R.Bean a qua

foluis plerumque angustioribus,

pedicellis brevioribus, et petalis

fructibusque minoribus differt. Typus:

Queensland. NortH KENNEDY DISTRICT:

Ridge near headwaters of Cockatoo

Creek, Mt Elliot National Park, south

of Townsville, 4 January 1997, A.R.

Bean 11563 & P.G. Bean (holo: BRI;

iso: A, K, MEL, NSW, QRS).

Babingtonia sp. (Townsville A.R. Bean

3424) in Henderson (1997),

Shrub to 1.5 m high. Bark grey, persistent, scaly

to fibrous. Branchlets quadrangular with flat

sides, not flanged, white to grey, margins entire;

oil glands present, papillose. Leaves petiolate;

petiole c. 1.0 mm long; lamina obovate, 8.0—

11.5 mm long, 2.2—-3.8 mm wide, concolorous,

straight, flat, not keeled, oil glands prominent,

especially on lower surface, c. 0.5 mm apart,

midrib faintly visible on abaxial surface,

invisible on adaxial surface, intramarginal vein

not visible, apex obtuse or acute. Inflorescence

axillary, 7(--9)-flowered; peduncles 10—12.5

mm long; stipes 2.0-3.5 mm long; bracts 2,

caducous, deltate, 0.6-0.8 mm long, acute;

bracteoles similar but somewhat smaller.

Hypanthium smooth, glandular, broadly

campanulate, 1.8—-2.2 mm long, fused to the

ovary except at top; calyx lobes simple, semi-

elliptic, c. 0.6 x 1.8 mm, thin, margins entire.

Corolla up to 9 mm across; petals orbicular,

2.5-3.0 x 2.5-3.0 mm, white, oil glands

present, margins entire. Stamens 11-14, in

groups of |—4 opposite the calyx lobes, stamen

opposite to centre of calyx lobe shorter than

remainder; filaments terete, 0.7—1.0 mm long,

geniculate, with brown connective gland fused

to upper part of filament at the bend; anthers

adnate, dehiscing by short divergent slits, with

loculi free. Style terete, up to 1.2 mm long after

anthesis, set into a pit; stigma broadly capitate.

Ovary 3-locular; floral disc shallowly concave;

ovules 18-20 per loculus, arranged radially

around placenta. Fruit hemispherical, c. 1.5 x

3.0 mm, valves broad, somewhat woody,

slightly exserted. Seeds discoid, c. 0.8 mm long,

brown, with flat sides and rounded backs, finely

reticulate (Fig. 1 I, J).

Specimens examined: Queensland, NortH KENNEDY

District: Cockatoo Creek area, Mount Elliot, S of

170

Townsville, Aug 1991, Bean 3589 (BRI, NSW, PERTH);

Cape Cleveland section, Bowling Green Bay N.P., 8S of

Townsville, Jui 1991, Bean 3424 (BRI).

Distribution and habitat: B. papillosa occurs

only in the Bowling Green Bay National Park

south of Townsville. This National Park includes

Mt Elhot and Cape Cleveland, where the two

known populations are located (Map 1). The

species 1s confined to shrubland on outcrops of

geranite-like rocks, on skeletal soil, and is

associated with shrubs such as Lepfospermum

brachyandrum (F.Muell.) Druce.

Phenology: Flowers have been recorded in

January; fruits from February to July.

Affinities: B. papillosa ts very closely related

to B. tozerensis, but differs from that by its

flower stipes 2—3.5 mm long (3.5—5 mm for B.

fozerensis), petals 2.5—3 mm in diameter (3—

3.5 mm for B. fozerensis), fruits 1.5 mm long

and 3 mm in diameter (2 mm long, 4 mm

diameter for B. tozerensis) and the mostly

narrower leaves (2.2—3.8 mm wide, compared

to 3.0-6.5 mm for &. tozerensis). No

intergrades with any other species are known.

Conservation status: The risk category for

Babingtonia papillosa, according to the criteria

of Chalson and Keith (1995) is “critical”

(criterion a). The species is known from just two

populations. There are 20 known plants at the

type locality, and 3 known plants at Cape

Cleveland. The species is under threat due to its

small population size and specialised habitat.

The recommended conservation status for this

species as defined by the Queensland Nature

Conservation Act 1992 is ‘endangered’.

Etymology: The specific epithet is from the

Latin papillosus, meaning “covered with

papillae or small wart-like structures”, and

refers to the raised oil glands present on the

young branchlets.

Dubious name

Baeckea virgata var. polyandra Maiden &

E.Betche, Proc. Linn. Soc. New South Wales 23:

12 (1898). Type citation: “Forms dense bushes

6-8 ft. high on the banks of the Snowy River at

Jindabyne (J.H. Maiden, January 1898)’,

Austrobatieya 5(2): 157-171 (1999)

No type has been located for this name,

but from the description given in the protologue,

this taxon is not related to Babingtonia virgata,

but may be referable to Kunzea ericoides.

Acknowledgements

I thank the Directors of A, BM, G, GH, K, MEL,

NE, NSW, HO, P, and WELTU for the loan of

specimens and/or access to their herbaria; Lloyd

Bird and Kym Sparshott for making collections

of Babingtonia; Lyn Craven (CANB) for the

Latin diagnoses; Mr John Williams and Mrs

Peggy Rowland for showing me B. crassa and

B. brachypoda respectively. John Dawson

(WELTV) kindly sent me his English-language

manuscript of New Caledonian Baeckea.

Thanks are due to Paul Williams and Col Adams

of N.P.W.S. Townsville for visiting the type

population of B. papillosa, and to my brother

Peter for accompanying me to collect flowers

of that species.

References

ANDREWS, H.C. (1810). Baeckia virgata, Twiggy Baeckia.

The Botanist’s Repository 9: t. 598.

BalLey, F.M. (1900). Baeckea. In The Queensland Flora

2: 583-6. Brisbane: J.H. Diddams & Co.

Bean, A.R. (1997). Reinstatement of the genus Babingtonia

Lindl. (Myrtaceae, Leptospermoideae). Austrobaileya

Ad): 627-45.

BENTHAM, G. (1867). Baeckea. In Flora Australiensis 3:

71-89, London: Lovell Reeve & Co.

CHALSON, J.M. & Keitu, D.A. (1995), A Risk Assessment

scheme for Vascular Plants: Pilot Application to

the Flora of New South Wales. Hurstville: National

Parks and Wildlife Service.

Dawson, J.W. (1992). Flora de la Nouvelle-Caledonie et

Dependances, 18. Myrtaceae-Leptospermoideae.

Paris: Museum National d’Histoire Naturelle.

EvuioT, W.R. & Jongs, D.L. (1982). Encyclopaedia of

Australian Plants suitable for Cultivation, Vol. 2.

Melbourne: Lothian Publishing Company.

HeEnbeERSON, R.J.F. (ed.) (1997). Queensland Plants, Names

and Distribution. Brisbane: Department of

Environment

MUELLER, F. (1864). Fragmenta Phytographiae Australiae

4: 68-74. Melbourne: Government Printer.

SCHAUER, J.C. (1843). Genera Myrtacearum Nova vel

denuo recognita. Linnaea 17: 235-44.

Bean, A.R., A Revision of the Babingtonia virgata in Australia 171

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Fig. 1. Babingtonia brachypoda A. leaf x 5. B. transverse section of leaf x 10. Babingtonia bidwillii C. leat x 5. D.

transverse section of leaf x 10. Babingtonia angusta E. leat x 5. F. transverse section of leaf x 10. Babingtonia

collina G. leaf x 5, H. transverse section of leaf x 10. Babingtonia papillosa |. leaf x 5. J. transverse section of leaf x

10, Babingtonia pluriflora K, M. leaves x 5. L, N. transverse section of leaves x 10. O. branchlet x 5, Babingtonia

crassa P, leaf x 5. Q. transverse section of leaf x 10. Babingtonia similis R. leaf x 5. S. transverse section of leaf x 10.

Babingtonia virgata T. leaf x 5, U. transverse section of leaf x 10. V. branchlet x 5, A,B from Bean 9543; C,D from

Bean 6803; E,F from Bean 8321; G,H from Bird (AQ566671); LJ from Bean 3589; K,L,O from Me/ville 2726; M,N

from Pullen 4618; P,Q from Bean 8289; R,S from Thorne 20481; T-V from A/cKee 37807.

Systematic and cladistic studies of Myrtella F. Muell. and

Lithomyrtus F. Muell. (Myrtaceae)

Neil Snow and Gordon P. Guymer

Summary

snow, N, & Guymer, G.P. (1999). Systematic and cladistic studies of Afrtel/a F. Muell. and Lithomyrtus

F, Muell, (Myrtaceae). Austrobaileya 5(2): 173-207. A systematic revision of species formerly placed in

Myrtella F.Muell. (Myrtaceae) is presented. Species primarily from New Guinea remain in Adyrtella

sensu stricto (AZ. beccarii, M. bennigseniana) whereas all species from Australia are transferred to the

genus Lithomyrtus F.Muell. Lithomyrtus includes seven new species viz L. densifolia N.Snow & Guymer,

L. dunlopii N.Snow & Guymer, L. grandifolia N.Snow & Guymer, L. ypoleuca N.Snow & Guymer, L.

kakaduensis N.Snow & Guymer, L. linariifolia N.\Snow & Guymer and L. repens N.Snow & Guymer,

Four new combinations viz_L. cordata (A.J. Scott) N.Snow & Guymer, L. microphylla (Benth.) N.Snow

& Guymer, L. obtusa (Endl.) N.Snow & Guymer and ZL. refisa (Endl.) N.Snow & Guymer are made for

other species belonging to Lithomyrtus, Myrtella phebalioides (W. Fitzg.) A.J.Scott is reduced to

synonymy under ZL. retusa, and Af. rostrata Lauterb. is transferred to Uromyrtus as U. rostrata (Lauterb.)

N.Snow & Guymer. Character data and preliminary cladistic studies strongly support the generic separation

of Lithonryrtus from Myrtella, but all clades within Lithonryrtus in the strict consensus tree are highly

unstable. Iterative studies with additional outgroups are needed to more confidently infer generic and

specific relationships in Lithomyrtus, Mfyrtella and related genera.

Keywords: Afyrtella, Lithomyrtus, Fenzlia, Uromyrtus, Myrtaceae, Myrtoideae, systematics, cladistics,

Australia, New Guinea.

Gordon P, Guymer, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha

Road, Toowong, Queensland 4066, Australia; e-mail: Gordon.Guymer@env.qld.gov.au

Neil Snow, Department of Biological Sciences, University of Northern Colorado, Greeley, 80639,

USA; e-mail: nsnow@bentley.unco.edu

Introduction

The family Myrtaceae Juss. is particularly rich

in South America and Australasia (Cronquist

1981; Johnson & Briggs 1985; Wilson 1996;

Mabberley 1997), with new taxa being

described yearly (Sobral 1994; Bean & Brooker

1995; Lyne & Crisp 1996). Although some

genera have been revised recently (e.g.,

Landrum 1981, 1988; Ladiges & Humphries

1983; Thiele & Ladiges 1988; Grifo 1992; Hill

& Johnson 1995; Toelken 1996; Chappill &

Ladiges 1996; Bean 1997a, b), many remain

poorly studied. Other biological aspects of

Myrtaceae have recently been investigated from

various perspectives (Beardsell et al. 1993;

Krauss 1994; Costa et al. 1995; Nic Lughadha

& Proenca 1996; Byrne et al. 1996; Gadek et

al. 1996; Landrum & Bonilla 1996; Haron &

Moore 1996; Orlovich et al. 1996; Shapcott &

Playford 1996; Sale et al. 1996).

Accepted for publication 8 October 1998

Taxa previously placed in Myrtella

F,Muell. sensu lato were reviewed most

recently by Scott (1978). For some time,

Australian botanists have been aware of

undescribed species in the Northern Territory,

particularly in the sandstone escarpment

regions of Kakadu National Park and

neighbouring areas that relate to this genus. In

this paper, we cladistically test the

hypothesised monophyly of the predominantly

New Guinean taxa (included in Myrtella s.s.

herein) and the exclusively Australian taxa

Gneluded in Lithomyrtus F.Muell. herein),

describe seven new species in Lithomyrtus,

provide the first comprehensive revisions of

both genera and transfer Myrtella rostrata

Lauterb. to Uromyrtus Burret.

Taxonomic history

Species herein assigned to Myrtella and

Lithomyrtus traditionally have been treated as

congeneric under Myrtella (Scott 1978). This

174

account 1s the first to accept Lithomyrtus for

Australian species traditionally assigned to

Myrtella. As detailed below, this generic

segregation is strongly supported by cladistic

studies.

The name Myrtella has been associated

with Fenzlia which Endlicher (1834) published

with the detailed description and illustration of

two Australian species (F obtusa and F: retusa).

As correctly noted by Scott (1978), Fenzlia

Endl. is an illegitimate later homonym of

Fenzlia Benth. (Polemoniaceae). Mueller

(1877) published Myrte//a for two new species

from New Guinea based primarily on their

“nearly valvular preflorescence of the calyx”,

although he was unable to suggest tribal

affinities of this genus due to the absence of

fruits. One of Mueller’s two initially described

species, Myrtella hirsutula F.Muell. was

recently transferred to Kania Schlechter by

scott (1990).

Bentham (1866) described Fenzlia

obtusa var. microphylla as new based on its

leaf size, and placed Fenzlia in his tribe

Myrteae which he characterized as having an

ovary divided into two or more cells (rarely

|-celled) with parietal placentation and the

fruit having a berry or drupe (but see

discussion of characters, Appendix 1).

Bentham and Hooker (1867) largely followed

the more detailed account of Bentham

(1866). Niedenzu (1893) was the first to

correctly identify the testa layer as the hard

portion of the fruit, which earlier had been

designated incorrectly as endocarp.

Burret (1941) merged Myrtella with

Fenzlia, and partitioned the species then known

into sections M. sect. Myrtel/a F.Muell. and M.

sect. Eufenzlia Burret. He contradicted himself

by saying initially the genera always look very

different (“*. .. sehen habituell recht verschieden

aus’) but later indicated he was unable to find

any substantial differences between them

(“Wesentliche Unterschiede zwischen beiden

Gattungen finde ich night”). The latter

statement is also at odds with the characters he

used to recognise the sections.

Lithomyrtus was described by Mueller

(1857: 228) as a “new genus, Lithomyrtus,

Austrobaileya 5(2): 173-207 (1999)

MSS.,” which he distinguished from Psidiua

L. by its “dry berry and a circinate embryo”.

Although no particular species was included

in it at that time, Mueller’s diagnosis of

Lithomyrtus constitutes valid publication of the

name (see ICBN Articles 32.1 and 41.2). Later,

Mueller (1858: 156) again made reference to

Lithomyrtus, but although labels on some of

his specimens bear the name L. hypoleuca

(DBL and K, photos at BRD), this combination

was never published (Muir 1978). Later still,

Mueller (1882) apparently abandoned

Lithomyrtus, since he excluded it from the

synonymy for native genera of Australian

plants. It is not clear to which genus Mueller

then intended to assign specimens he annotated

as Lithomyrtus hypoleuca (see above). He may

have considered L. fhypoleuca conspecific with

L. obtusa.

In the most recent synopsis of the genus

Myrtella Scott (1978) recognized sections M.

sect. Myrtella (4 spp.) and M. sect. Fenzlia (5

spp.), and described as new the Australian

species M. cordata. Briggs & Johnson (1979)

and Johnson & Briggs (1985), however,

maintained Myrte//la as generically distinct

from Lithomyrtus (as Fenzlia).

Materials and methods

Taxonomic descriptions: Taxon boundaries

(Snow 1997) and character data for

Lithomyrtus were determined from herbarium

specimens and fresh and preserved material

collected in the field; for Myrtella only

herbarium specimens were used. The list of

characters (Appendix 1) and natural language

descriptions were generated using the DELTA

program of Dallwitz et al. (1993). Embryos

were absent in all fruits of Lithomyrtus

linariifolia and L. hypoleuca examined.

Terminology for tissue layers of fruits and seeds

follows Esau (1960). Conservations status

evaluations are made according to IUCN

categories (Species Survival Commission

1994),

Cladistic analysis: Phylogenetic relationships

of Myrtella and Lithomyrtus were inferred

using PAUP Vers. 3.1.1 (Swofford 1993) with

the branch and bound algorithm (Hendy &

Penny, 1982) with the following options —

A ee a ee a an err

Peed Get eed cot iocre Stee cd her eed PPE ee CEE ae EAE CORE EO

AI BT BL ST ce ge nan ee a et

Snow & Guymer, Systematic and cladistic studies of Afyrtella and Lithomyrtus (Myrtaceae) 175

keep minimal trees only, collapse zero length

branches, additional sequence furthest, and

MULPARS. All characters were unordered

and unweighted (Appendix 2; Table 1). A test

for phylogenetic signal was assessed using

the gl statistic with 10,000 random trees

(Huelsenbeck.1991). A successive weighting

procedure based on the rescaled consistency

index (Naylor & Kraus 1995) was

implemented following the unweighted

analysis. Candidate outgroups were not

obvious since many genera of Australasian

Myrtoideae have been poorly studied and

generic relationships among them untested.

We thus designated Myrtel/a as the outgroup

and rooted the trees to make Lithomyrtus a

sister clade to Myrtella, The stability of

clades in the strict consensus tree was

evaluated using minimal character support

values (Davis 1993).

Results and discussion of cladistic analysis

Unweighted analysis: The gl value of -2.82

indicates considerable left-hand skewness

suggesting greater phylogenetic signal than

expected at random (Huelsenbeck 1991). The

unweighted analysis resulted in four most-

parsimonious trees (MPTs) of 70 steps (after

rooting: ci = 0.857; 11 = 0.744, re = 0.637).

The one fully-resolved MPT and the strict

consensus tree are illustrated in Figures 1& 2.

In all MPT’s, the two species of Myrtella

s.s. formed a clade basal to Lithomyrtus giving

cladistic support to the hypothesis that these

genera are distinct (Fig. 1). Thirteen

uncontested synapomorphies support the

distinctness of Lithomyrtus from Myrtella (2,

3,8, 10, 13, 14, 16, 19, 22—26). Although some

of these characters may break down in

Myrtaceae when viewed collectively this

analysis strongly supports segregation of

Lithomyrtus from Myrtella.,

The exceptionally strong support for the

separation of Myrtella from Lithomyrtus

contrasts markedly with the weak support for

separation of clades within Lithomyrtus. Since

each clade within Lithomyrtus in the strict

consensus tree can be collapsed by the removal

of a single character (Davis 1993), little weight

should be accorded to those clades until

congruency testing with additional characters

suggests better support (Patterson 1982).

Successively weighted analysis: Successive

weighting based on the rescaled consistency

index converged on two MPT’s after only one

iteration. The strict consensus tree based on

successive weighting (Fig. 3) had only two

additional nodes resolved relative to the

unweighted strict tree, so this procedure was

of minimal help in inferring uncontested

patterns of descent. However, the tree very

closely resembled the one fully-resolved tree

from the unweighted analysis.

The choice of root can strongly influence

character polarity and the recognition of

monophyletic groups (e.g., Scotland 19972).

Since generic relationships in Myrtaceae remain

poorly known and most untested cladistically,

further analyses with additional outgroup genera

will be necessary before reliable inferences can

be made regarding relationships between

Lithomyrtus, Myrtella and related genera. In

particular, the monotypic Myrtastrum Burret

(New Caledonia) and Neomyrtus Burret (New

Zealand), which resemble Myrtel/a in gross

morphology, may be closely related and thus

are candidate genera. However, given the

seographic proximity of Myrtel/a (mostly New

Guinea) and Lithomyrtus (exclusively

Australian) and the fact that many myrtaceous

genera (e.g., Austromyrtus s.1., Melaleuca,

Asteromyrtus, Eucalyptus) range across these

areas, it was not unreasonable to use Myrtella

to root a cladistic analysis of Lithomyrtus for a

first approximation of species relationships.

176

~ rr Ey Pr

Austrobaileya 5(2): 173-207 (1999)

Mi. beccaril

M. bennigseniana

L. cordata

L. kakaduensis

L. grandifolia

. microphylla

. Aunlopii

. hypoleuca

. retusa

. linartifolia

. repens

- L. densifolia

- L, obtusa

Figure 1. One of four most parsimonious trees resulting from branch and bound search using unweighted and

unordered characters, with Afyrte//a designated as outgroup a priori.

Figure 2. Strict consensus tree.

or er a iii

Fak Pec raee tee PSCC Pet err

- M.beccarii

- M. bennigseniana

. cordata

. kakaduensis

L. grandifolia

- |, densifolia

- L. dunlopii

. hypoleuca

L, retusa

[. linariifolia

. repens

- L, microphylla

- 1, obtusa

snow & Guymer, Systematic and cladistic studies of Adprtella and Lithomyrtus (Myrtaceae) 177

M. beccaril

M. bennigseniana

L. cordata

L, Kakaduensis

L. grandifolia

L. microphylla

. dunlopii

. hypoleuca

. fetusa

- L, linariifolia

- L, repens

- L. densifolia

L. obtusa

Figure 3. Strict consensus tree based on successive weighting with the rescaled consistency index,

Taxonomic treatment

Key to Genera

Branchlets distinctly 4-angled, edges of younger stems winged, the wing apices

flaring outward; leaves mostly glabrous above and below (but often

sericeous on lower portions of midvein); stipules very short, thick, dark

red, scale-like, often fused laterally; apex of youngest leaves distinctly

comose; bracteoles glabrous (or minutely hairy on edges), leaf-like and

usually with a midvein, somewhat falcate and flexuous, usually persistent

in fruit; sepals valvate in bud; stamens unt- or multiseriate; fruits globular,

less than 5 mm long, glabrous, red to nearly black; stamina] disk glabrous;

style glabrous; ovary 2 or 3 (or 4)-locular; ovules 2—4 per locule; embryo

relatively thick, slightly curved or crescent-shaped; embryo not spirally

contorted, thickened towards tip; cotyledons straight, thick, very short.

Indonesia UIrian Jaya), Papua New Guinea, Guam; reported also from

Mariana and Caroline Islands... 00... cc eee eee eens Myrtella

Branchlets rounded, younger twigs rounded; leaves mostly somewhat hairy

above, often villous to tomentose below; stipules consisting of two to several

free ferrugineous setose hairs; apex of youngest leaves not distinctly

comose; bracteoles hairy (or at least glabrescent), scale-like and lacking a

midvein, straight or slightly curved, usually absent in fruit; sepals imbricate

in bud; stamens multiseriate; fruits globular to cylindrical or fusiform,

mostly greater than 5 mm long, glabrescent to densely tomentose, yellow-

to olive-green or whitish by virtue of dense hairs; staminal disk hairy; style

178

Austrobaileya 5(2): 173-207 (1999)

often sericeous, especially near base; ovary 1-locular; ovules (1—) 2 (or

3) per locule; embryo relatively thin; circinate; embryo sometimes spirally

contorted, slightly thickened towards tip; cotyledons circinate, thin,

relatively long. Tropical Australia ........

Myrtella F.Muell., Descr. Notes on Papuan

Plants 1: 105 (1877). Type: Myrtella

beccarii F.Muell. (lecto: fide Scott 1978).

Saffordiella Merrill, Philipp J. Sc. 9: 124

(1914). Type: Saffordiella bennigseniana

(Volk.) Merrill.

Shrubs or small trees, 0.3—-6.0 m tall. Bark

smooth or stringy, brown or grey. Branchlets

distinctly four-angled, the edges distinctly

winged, glabrous or sparsely sericeous to villous

neat apex of internodal wings, oil glands visible

and prominent, or lacking. Leaves decussate,

opposite, evenly distributed or mostly

concentrated near branch tips, coriaceous.

Stipules consisting of 2—several short, thick,

dark red, often laterally fused scale-like

structures. Petioles 0.5-1.0 mm long, not

channelled, eglandular. Leaf blades narrowly

elliptic to narrowly ovate, margins usually

slightly revolute, bases truncate, not clasping

the stem, apex obtuse, upper surface mostly

glabrous but sericeous on lower portion of

midvein (young leaves apically comose),

adaxial midvein impressed or not, lower surface

glabrous or sericeous on lower portion of

midvein, glands of lower surface visible and

about the same size, marginal veins of lower

surface invisible or indistinct. Inflorescence a

solitary axillary flower. Peduncles rigid,

glabrous. Bracteoles foliaceous, midrib usually

present, irregularly flexuous, the tips (in flower)

exceeding base of sepal lobes, glabrous, mostly

persistent in fruit. Hypanthium obconic,

ee yet oe ee wee ed ee Lithomyrtus

glabrous. Sepals 5, fused below. Sepal lobes

valvate in bud, apex acute, glabrous to sparsely

short tomentose along edges, persistent in fruit,

more or less ascending and mostly held above

fruit. Petals 5, white to pink, elliptic to ovate,

upper surface glabrous, lower surface glabrous

to minutely tomentose on edges. Stamens 15—

50, uni- or multiseriate, included, folded

centrewards in bud. Stamina] disk glabrous.

Anthers subglobose, dorsifixed, versatile, less

than '/, length of filaments, dehiscence by

longitudinal slits, connective glandular or

eglandular. Ovary 2 or 3 (or 4)-locular, ovules

2—4 per locule, placentation parietal. Style 1,

flexuous, or mostly straight, glabrous. Stigma

1, terete or scarcely (if at all) capitate. Fruit

indehiscent, a hard berry with bony, more or

less fused seeds, subglobose to globose, the base

somewhat tapered, glabrous, red or dark blue

to brown or nearly black (when dried). Seed

coat hard, bony. Adjacent seeds slightly fused,

their boundaries in longitudinal section

generally distinct and perpendicular to long axis

of fruit. Embryo slightly curved to crescent-

shaped. Radicle relatively short, relatively thin,

swollen near tip, the tip not spirally contorted

(held at same horizontal plane as embryo).

Cotyledons straight, relatively short; shorter

than hypocotyl, relatively thick, not folded back

towards hypocotyl.

Papua New Guinea, Irian Jaya, Guam;

reported but not verified from Caroline Islands

and Mariana Islands (Diels 1921).

Key to species of Myrftella

Leaves greater than 9 mm long; internodes of branchlets mostly longer

than 0.4 cm; wings of young twigs hairy only near apex; stamens

multiseriate ...... 030 ceccrececaetduvue

FL ES ts Ue WP eas We 1. M. becearn

Leaves less than 6 mm long; internodes of branchlets mostly shorter

than 0.3 cm; wings of young twigs hairy throughout; stamens uniseriate

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Deities ty Mesesenecer om pam yt ee Mae tanree rence pihscca dear ahs

Snow & Guymer, Systematic and cladistic studies of Adprtel/a and Lithomyrtus (Myrtaceae) 179

1. Myrtella beccarii F.Muell., Descriptive

Notes on Papuan Plants 106 (1877).

Type: Indonesia. Irian Jaya. Baia de

Humboldt, XIJ/1875, Beccarii 37777

(lecto: here designated FI).

Shrubs or small trees, 0.5—5 m tall. Bark smooth

or stringy, brown. Branchlets glabrous to

sparsely sericeous near apex of internodal

wings, oil glands prominent. Leaves more or

less evenly distributed. Petioles 0.5—1 mm long,

rounded, eglandular. Leaf blades narrowly

elliptic to elliptic, 7-11(—13) mm long, 2~-3.5(-

5) mm wide, base truncate, apex obtuse, margins

revolute, upper surface glabrous except for

sericeous lower portion of midvein and young

leaves, adaxial midvein flush above, lower

surface glabrous or sericeous on lower portion

of midvein, marginal veins of lower surface

absent or indistinct. Peduncles 3-4 mm long in

flower. Bracteoles 3~3.5 mm long, 0.6—0.8 mm

wide, glabrous. Sepal lobes 2.5—3 mm long,

sparsely short-tomentose along edges. Petals

white, ovate, 4 mm long; the lower surface

minutely tomentose on edges. Stamens

uniseriate or biseriate, c. 2.5 mm long; anther

connectives eglandular. Style 2.5-4 mm long,

flexuous or mostly straight. Fruit subglobose,

sometimes abruptly tapered to base, 2.5-4.5 mm

long, 2.5-3.5 mm wide, green, turning red to

dark brown or nearly black at maturity. Fig. 4 A-H.

Specimens examined: Papua New Guinea. MorosBe

District: LAE Supbistrict. Lasagna Island, 7°25’S,

L47°1S°E, Nov 1969, Streimann NGF44305 (BRI, L);

Buso Village, 7°25°S, 147°10’E, Nov 1969, Streimann

NGF44151 (BRI, L); Buso River, Foreman LAES2292

(.}; Buso River, SE of Lae on the coast, opposite Lasagna

Island, 7°25’S, 147°10’E, Nov 1973, Jacobs 9650 (L);

Buso River in lower land beside the river, Jun 1980, Rau

603 (L); 8 km S of Salamaua, Sep 1973, Paijmans 1564

(L.); Buso, 7°25’S, 147°10’E, Nov 1979, Henty LAE72469

(CANB, L, QRS); Koneipa Village, Kipu, 7°50’S,

147°10°B, Jan 1966, Streimann & Kairo NGF26129 (BRI,

L); Buso, S of Lal, Jun 1978, Bellanry B6 (L). Morospe

SUBDISTRICT: Paiawa Valley, along lower reaches of Paiawa

River, 7°35’°S, 147°15’E, May 1970, Jo/ins NGF47319

(BRI); Buso (Morobe), § of Lae, Jun 1970, Bellamy s.n.

(CANB, L); Creek at end of logging road near Paiawa

River, SPT, 7°30’, 147°15’E, Oct 1965, Gillison

NGF25026 (CANB, L); Payawa Village, 7°35’S,

147°10’E, Jun 1981, Katik LAE74928 (L). NorTHERN

District: Ca 2 miles S$ of Toma, Bariji-Managalese area,

Sep 1964, Pullen 5919 (BRI, L). West New Britain

District: Hoskins Suppistrict: NNE slope Mt Ulawon,

$°02’S, 15°22’E, Feb, 1971, Stevens LAE51251 (BRI,

CANB, L). District UNKNOWN: Korepa, 7°55’S,

147°05°E, Kairo 47 (CANB). Indonesia. [RIAN Jaya:

District HOLLANDIA: Cycloop Mts, along path to Neta,

Jun 1938, Mayer Dress 138 (L); Cycloop Mts, foothtils

W of Koejaboe River and E of Hoebal River, Royen 4149

(L). Districr RapJAH Ampatr: Weigo Island, E bank of

Majalibit Bay, Jan 1955, van Royer 5217 (CANB, L);

Town of Sukarnapura, Koatermanns & Soegong 560 (L);

Waigeo Island, Kabare Bay, Jan 1955, var Royer 5361 (L).

Distribution and habitat: Papua New Guinea,

Indonesia (Irian Jaya); reported but not verified

from Solomon Islands and Micronesia (Scott

1990). Growing in a wide range of habitats,

predominantly in moist rocky and disturbed

areas but also near beaches, in forests of

Weinmannia or Casuarina, and in xerophytic

vegetation on ultrabasic soils (van Royen 5361),

at altitudes from near sea level to 2500 metres.

Phenology: Flowering January to November;

fruiting mostly April to November.

Notes: The flowers of M. beccarii are white,

and the stamens are reported to be pale green

or cream coloured. The bright red fruit dries to

a much darker red colour. Due to the wide

elevational range, additional or cryptic taxa may

exist within it as currently accepted.

Conservation status: Herbarium labels often

report the species as locally common or

abundant,

Local names: zagicbara (Middle Waria,

Streimann & Kairo NGF26129 [BRI L]).

2. Myrtella bennigseniana (Volk.) Diels in

Engl. Bot. Jahrb. 56: 529 (1921);

Leptospermum bennigsenianum Volk. in

Engl. Bot. Jarhb. 31: 407 (1902);

Saffordiella bennigseniana (Volk.)

Merrill, Philipp. J. Sci. 9: 124 (1914);

Fenzlia bennigseniana (Volk.) Burret,

Notizb. Bot. Gart. Berlin 15: 500 (1941).

Type: Caroline Islands, Volkens 277

(lecto: CAL, here designated) (type

material at B destroyed, fide Hiepko

1987). |

Shrubs or small trees, 0.3-6.0 m tall. Bark

smooth, brown or grey. Branchlets sparsely

sericeous to villous, sometimes densely so, oil

glands absent. Leaves concentrated near branch

tips or mostly evenly distributed. Petioles 0.5—

1 mm long, rounded, eglandular. Leaf blades

180 Austrobaileya 5(2): 173—207 (1999)

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Figure 4. A-H: Myrtella beccarii. A. flowering branchlet x 3. B. section of branchlet x 8. C, adaxial leaf profile x 4.

D, adaxial leaf apex x 8. E. flower x 4. F. bracteole x 8. G. fruit x 4. H. flower bud * 3. A,C,D,H: Henty LAE72969 (L).

B: Streimann NGF44151 (BRI). E,F,G: Bellanry B6 (L). -O: M. bennigseniana., |. flowering branchlets x 3. J. section of

branchiet x 8. K. abaxial ieaf profile x 4. L. adaxial leaf apex x 8. M. flower = 6. N. bracteole x 12. QO. fruit x 12. [-O:

Rinehart 12019 (BRI). P—-W: Lithomyprtus cordata. P. flowering branchlet x 1.5. Q. section of branciet x 16. R.

abaxial leaf profile x 1. S. adaxial leaf apex (hairs not shown) x 2. T. flower with front petals removed x 3. U. bracteole

x 8, V. fruit x 2. W. flower bud x 3. P—W: Snow 7420 et al. (BRD).

Snow & Guymer, Systematic and cladistic studies of Adyrfella and Lithomyrtus (Myrtaceae) 18]

elliptic to narrowly ovate, 3.5—5(-7) mm long,

2—2.5 mm wide, base truncate, free from the

stem, apex obtuse, margins revolute, upper

surface glabrous except for sericeous lower

portion of midvein and young leaves, midvein

impressed above, lower surface glabrous or

sericeous on lower portion of midvein.

Peduncles 2—5 mm long in flower. Bracteoles

23.5 mm long, 0.4—-0.6 mm wide, glabrous or

minutely and sparsely sericeous along edges.

Sepal lobes 1.5—2 mm long, glabrous or sparsely

short tomentose along edges. Petals white to

pink, elliptic to ovate, 2-3 mm long; lower

surface glabrous. Stamens uniseriate, c. 1.5 mm

long; anther connectives glandular. Style

1.5—-3 mm long, flexuous. Fruit globose,

tapered at base, 1.5—3 mm long, 2.5—-3 mm

wide, brown to dark bluish-black when

mature. Fig. 4I—O.

Specimens examined: Guam. Savanna in Siqua Falls

area, between Alutom & Tenjo ridges, Jan 1986, Rinehart

LR12019 (BRD; Savanna along road (and pipeline) to

OMNI on north spur from Fonte Plateau (Nimitz Hill),

Sep 1985, Rinehart LR11544 (BRI); Savanna in Siqua

Falls area, between Alutom and Tenjo ridges, Jan 1986,

Rinehart LR12019 (BRI). Indonesia. [RIAN Jaya:

Tablasoefoe, E of Village, Aug 1961, var Royen &

Sleumer 6449 (BRI); Cycloop Mountains, southern slope

of Makanoi Range, N of Kotanica, Jul 1961, van Royen

& Sleumer 6211 (BRI); Cycloop Mountains, Arthur Hill,

E slope, Jul 1961, van Royen & Sleumer 6161 (BRD;

Hollandia [= Jayapura] and vicinity, Jun—Ju! 1938, Brass

8887 (BRI); Jayapura, Boschung der Ktistenstrasse im

nordéstlichen Teil der Stadt, Feb 1976, Hiepko & Shultze-

Motel 579 (L); Kota Nica, Hollandia, Oct 1956, Schram

BW2844 (1); Town of Sukarnapura (— Hollandia),

Kostermanns & Soegeng 560 (L); Cape Tanah Merah,

Aug 1961, van Royen & Sleumer 6501 (L); “Base G’,

Hollandia, Dec 1956, Sidje BW4126 (CANB, L); Cycloop

Mts, along path to Near, Aveyer Dress 138 (L); Cycloop

Mts, foothills W of Koejaboe River and E of Hoebal River,

van Royen 4149 (L); Jayapura, , Béschung der

_ Ktistenstrasse im nordéstlichen Teil der Stadt, Feb 1976,

Hiepko & Shultze-Motel 509 (L). Papua New Guinea.

Sewa Bay, Normanby Island, April 1956, Womersley

NGF8687 (BRI); Normanby Island, Mt Pabinama, May

1956, Brass 25703 (L); Province Miine Bay, Subprovince

Esa’ala, NE of Bwasiaiai, Dec 1977, Croft & Marsh

LAE71203 (L); Along Arumu River S of Botue village,

Sep 1953, Hoogland 3963 (BRI, L).

Distribution and habitat: Indonesia, Papua

New Guinea, Guam; also reported but not

verified from Caroline and Mariana Islands. In

open grasslands, occasionally in understorey of

Dacrydium forests, on lateritic soils, between

20 and 2500 metres altitude.

Phenology: Flowering March to September;

fruiting March to October.

Notes: The flowers of M. bennigseniana are

white or pinkish. The reddish fruit dries to

nearly black. he uniseriate, glandular stamens

are reliable diagnostic attributes in flower as

are the densely pubescent young twigs in sterile

material. Reports of the species from the

Caroline Islands (Merrill 1914; Diels 1922) and

Mariana Islands (Diels 1921, 1922) have not

been confirmed. The wide elevational range

suggests the possibility of more than one taxon

within this species as currently accepted.

Conservation status: The relative few

collections ofthis species and dearth of detailed

information on specimen labels indicate the

species is data deficient.

Local name: ététi (Orokaiva language,

Mumuni; see Hoogland 3963 [BRI, L]).

Excluded species

Uromyrtus rostrata (Lauterb.) N.Snow &

Guymer, comb. nov.

Myrtella rostrata Lauterb,, Nova Guinea 8:

855(1912). Type: New Guinea. Hellwig

Mountains, Roemer 1050 (holo: L, photo

BRI).

The type specimen of Myrtella rostrata has the

inflorescence, fruit and seed characters of

Uromyrtus and is subsequently transferred to

this genus.

Kanta hirsutula (F.Muell.) A. J. Scott, Kew

Bull. 45: 206 (1990); Myrtella hirsutula

F.Muell., Descr. Notes Papuan Pl. 106

(1877); Type: Indonesia. [rian Jaya.

Mount Arfak, Jul 1875, Beccari 3776

(lectotype, here designated: FI).

According to Article 37.3 of the International

Code of Botanical Nomenclature (Greuter et al.,

1994), sufficient information was provided by

Mueller (1877) to indicate a holotype. However,

given that Mueller loc. cit. did not cite a

collection number of Beccari’s in the protologue

and that Scott (1990) indicated no specimen can

be found in the National Herbarium of Victoria,

Melbourne on which Mueller might have based

182

his description, lectotypification here eliminates

ambiguity surrounding the possible existence

of a putative holotype implied by Scott’s (1990)

citation of an isotype. Scott loc. cit. provides

itrefutable evidence for transferring Myrtella

hirsutula to the genus Kania.

Lithomyrtus F.Muell., J. of Bot. and Kew Gard.

Misc. 9: 228 (1857). Type: Lithomyrtus

hypoleuca F.Muell. ex N.Snow &

Guymer, lecto: here designated.

Fenzlia Endl., Atakta Bot. 19. 1834, nom.

illegit. non Fenzlia Benth., Bot. Reg. sub

t. 1622. 1833. Type: Fenzlia obtusa Endl.

(ecto:fide Scott 1978)

Frutices prostrati usque plerumque erecti vel

raro arbusculae, folia juvenia et rami villosi

usque tomentosi, saepe minus hirsuti

vetustatibus. Stipulae parvae at prominentes,

pilorum 2 aliquot ferrugineorum setosorum

consistentes. Bracteolae non foliosae, absque

costis distinctis. Flores solitari1 in axillis

foliorum. Sepala petalaque pentamera, lobi

sepalorum in alabastro seorsi, stamina

multiseriata, numerosa. Fructus baccam vel

baccatus. Ovarium maturitate plerumque

solitarrum. Semina plerumque 2 vel 3, oblique

conferruminata, testa lignea. Embryo circinatus,

cotyledones tenues, circinatae, apex radiculae

incrassatus, ultra embryonem extendens.

Plants suffrutescent, shrubs or small trees,

().2—5 m tall, stems prostrate, ascending or erect.

Bark smooth or stringy, brown, grey to

somewhat orange-coloured. Branchlets

rounded, glabrous to tomentose, oil glands

present and prominent or absent. Leaves

decussate, opposite or rarely whorled at some

nodes, evenly distributed or concentrated near

erowing tips, coriaceous to velvety. Stipules of

2~several free, ferruginous, setose hairs.

Petioles channelled or rounded, eglandular or

rarely glandular. Leaf blades linear to elliptic,

ovate or sometimes obovate; margins flat or

revolute; bases truncate, cordate or cuneate,

occasionally clasping the stem; apex obtuse,

retuse or acute, adaxial surface glabrous to

tomentose with midvein impressed; lower

surface glabrous to tomentose, marginal veins

of lower surface prominent, indistinct or absent.

Inflorescence a solitary axillary flower.

Austrobaileya 5(2): 173-207 (1999)

Peduncles rigid or sometimes flexuous, sparsely

antrorsely sericeous to villous or tomentose.

Bracteoles lacking a midrib, ascending, scale-

like, slightly curved to straight, exceeding or

shorter than base of sepal lobes, sparsely

sericeous to densely villous or tomentose,

persistent or caducous in fruit. Hypanthium

obconic, densely sericeous to tomentose. Sepals

5, fused at the base. Sepal lobes imbricate in

bud, apex acute to acuminate, sparsely sericeous

to tomentose, persistent in fruit, held above or

reflexed towards fruit. Petals 5, white, pink or

magenta, elliptic to ovate or obovate. Stamens

numerous, multiseriate, included, folded

centripetally in bud. Staminal disk hairy.

Anthers subglobose, dorsifixed, versatile, less

than ‘4 length of filaments, dehiscing by

longitudinal slits, connectives glandular or

eglandular. Ovary 1-locular, ovules (1 or) 2 (or

3) per locule, placentation parietal. Style 1,

flexuous or mostly straight, glabrous or sparsely

sericeous, Stigma |, terete, rarely capitate. Fruit

a hard berry with bony, tightly fused seeds,

subglobose to cylindrical or sometimes

fusiform, base rounded or tapered, glabrous to

tomentose, light yellow or olive-green, or

whitish by dense pubescence. Seed coat hard,

bony. Adjacent seeds tightly fused, their

boundaries mm longitudinal section generally

indistinct and often somewhat oblique to the

long axis of fruit. Embryo circinate. Radicle

relatively long, relatively thin, swollen near tip,

the tip held at same horizontal plane as embryo

or spirally contorted above or below.

Cotyledons circinate, relatively long, shorter

than hypocotyl, relatively thin, not folded back

towards hypocotyl. Tropical Australia (Western

Australia, Northern Territory, Queensland).

All Australian species formerly in

Myrtella are now assigned to Lithomyrtus,

which is confined to the North Australian

Province of the Northeast Australian Region

(Takhtajan 1986).

Notes: Leaves of L. retusa seedlings under a

metre tall are generally longer than those of

mature plants (Snow, pers. obs.) and the

possibility of heteromorphy between juvenile

and adult leaves needs further investigation. The

black hair-like structures occasionally occurring

on the foliage of some species (particularly LZ.

Snow & Guymer, Systematic and cladistic studies of Myrtella and Lithomyrtus (Myrtaceae) 183

obtusa) are actually an ophiostimoid fungus (J.

Simpson, pers. comm.).

The outer layers (perisperm) of the mature fruit

of this genus typically disintegrate, leaving as

a single diaspore the hard, fused seeds. The

individual seed coats often have openings

(opercula) from which the radicles probably

emerge during germination of the seed.

Most plants with mature fruit of all Lithomyrtius

species show evidence of some insect predation.

A preliminary identification of insect casings

and pupae found on their leaves and fruits

shows these insects represent two families (P.

Gullan, pers. comim.). The first of these families,

Diaspididae (armoured scale insects), consist

of insects phytophagous on a variety of plant

parts, including fruits. The second, family

Tachinidae (tachinid flies), comprises mostly

parasites of other insects. Whether either insect

is actually attacking the frutts of this genus

however, is unknown. A single moth pupa was

found embedded in a fruit of LZ. hypoleuca but

it was insufficiently mature for identification to

family level (J. Lawrence, pers. comm.).

Investigation of fruit predation therefore would

appear to be a worthwhile field study,

particularly mn the Kakadu region of the Northern

Territory where several Lithomyrtus species

coexist,

The ornamental potential of the genus is well

known (Hubbard 1978) and evaluations of this

aspect is currently in progress at the Darwin

Botanical Garden (G. Leach, pers. comm.).

Potential horticultural disadvantages observed

in the field include insect and fungal attack on

L. grandifolia and ophiostemoid fungal infection

occasionally noted on other species, most

notably L. obtusa. Attempts to germinate mature

seeds of seven species representing several

accessions for each were unsuccessful.

Key to species of Lithomyrtus

1. Leaves linear, 10-51 mm long and |—3 mm wide; Northern Territory .... 7. L. linartifolia

Leaves narrowly elliptic to ovate or obovate . 0... eect eens 2

2. Leaves 26-61 mm long, 10-46 mm wide; Northern Territory ......... 4, L. grandifolia

Leaves mostly less than 25 mm long and less than 20 mm wide... ..... 0... cee eee 3

Hey (GSAT DOSES COR alee al. cit i stake wt «Teles fe 4 las os AS OPS toy WRU Abe eas a BNE SS Wa aly Slat # a 4

Peal DASES-THCaLS OMCUNECALS ar. pese oe Ch a ea wt pets oe alee pa a neg ane) Sa 6

4. Plants prostrate; peduncles in fruit often longer than subtending leaves;

NOMMEI LEMITOIy (ir sts eo ate et eae oe, Boal d Wee pen beg oo ee el CN a a 10. L. repens

Plants erect; peduncles in fruit generally shorter than subtending leaves .............. 5

5. Base of leaf blades clasping stem; petiole up to 0.5 mm; leaves mostly

eglandular abaxially; Northern Territory

= 8s © #8 #& # *F © 8 © © & © © & © £ * B&B BF B 82 8

6. L. kakaduensis

Base of leaf blades free from stem; petiole mostly greater than 1 mm

long; leaves always glandular abaxially; Northern Territory ............

6. Plants prostrate; Northern Territory ...... 0.00.0. eens 10. L. repens

PUQIES GRE CTs nis. auc aks, Bie eure enetad aL es 8 Wiens a AE GLP Whey a emma! wag HY Ma eon ake bet ete sie 7

7. Leaves less than 30 mm long and less than 2 mm wide; Queensland .... 8. L. microphylla

Leaves mostly longer than 30 mm and greater than 2mm wide ............ 200s

8. Leaf blades eglandular abaxially (rarely glandular in L. hypoleuca and L. obtusa) ...... 9

Leal Diades 2 lati has a ba mtely 6405 bee o's tale. vad Be eet 4 Boyind K, veke Gegie Sarnalahied Sealla baled ba a 9 1]

9. Leaf blades narrowly elliptic; tips of sepal lobes mostly held above fruit;

Northern Territory ......... 0000. eee

184

Austrobaileya 5(2): 173-207 (1999)

Leaf blades broadly elliptic to ovate or obovate; tips of sepal lobes

tefiexed and held adjacent to fruit ......

10. Bracteoles not exceeding base of sepal lobes in flower; anther connectives

eglandular; lateral abaxial leaf veins invisible; Northern Territory,

Oucensland’ . oc. 4s sy bs «dean + oa o

= & & © # #€ # © @#@ #8 *#© @# # @&@ &© © @#@ &# 8 © &£€£ & & @

5. L. hypoleuca

Bracteoles exceeding base of sepal lobes m flower; anther connectives

glandular; lateral abaxial leaf veins clearly visible; Northern Territory,

ROUTES] ANGE, pose oot aha, a a 8 Seca ce Fe eg od

> + © © © &© FF B 8 8 #*# @8 © © 8 ££ © 8B & & & © £ F&F HF FF Ff

9, L. obtusa

11. Oil glands often inconspicuous (use magnification); plants usually less

than | m tall; foliage relatively dense and mostly evenly distributed on

stems; petioles not channelled adaxially; tips of sepal lobes mostly erect;

Northern Territory ..............0000.

Oil glands visible without magnification; plants usually greater than |

metre tall; foliage not dense and often concentrated near branch tips;

petioles channelled adaxially; tips of sepal lobes reflexed..................0005. 12

12. Foliage oil glands ferruginous, unequal in size; leaf blades narrowly elliptic

to elliptic or obovate, cuneate at base; petals 3-6 mm long; anther

connectives usually with several glands; fruit globular to subglobular,

3-6 mm jong, glabrous to sparsely villous; Western Australia, Northern

Territory, Queensland .............0..

= AP lag Suk Os "ot US YE oben cd eyes LE 11. L. retusa

Fohage oil glands yellow-green, equal in size; leaf blades narrowly ovate

to ovate, cordate or truncate at base; petals 7-12 mm long; anther

connectives usually with a single gland; fruit subcylindrical, 7-12 mm

long, sparsely tomentose to tomentose; Northern Territory .............

1. Lithomyrtus cordata (A.J. Scott) N.Snow

& Guymer comb. nov.

Myrtella cordata A.J. Scott, Kew Bull: 33:

301 (1978). Type: Australia. Northern

Territory. DARWIN AND GULF REGION: Deaf

Adder Basin, 10 Jun 1972, R. Schodde

AES 1 (holo: K; iso: CANB, DNA, L, NT).

Erect shrubs to small trees, 1-5 m tall. Bark

smooth or stringy, brown, grey or orangish.

Branchliets tomentose, oil glands present and

prominent or absent. Leaves opposite, mostly

concentrated near branch tips, soft, hairy,

petioles 1.5—6 mm long, channelled, adaxially

eglandular or glandular. Leaf blades narrowly

ovate to ovate, 12-48 mm long, 5—26 mm wide,

base often cordate or occasionally truncate,

apex obtuse or acute, margins revolute, adaxial

surface sparsely short villous, glabrescent,

midvein impressed, adaxial surface shortly and

densely tomentose, glands of lower surface

visible and about the same size, marginal veins

of lower surface prominent (usually), or

invisible or indistinct (occasionally). Peduncles

3. L. cordata

rigid, 3-14 mm long, tomentose. Bracteoles 4—

5 mm long, c. 0.5 mm wide, the tips (in flower)

not exceeding base of sepal lobes, or rarely

exceeding base of sepal lobes, tomentose,

persistent in fruit. Hypanthium tomentose. Sepal

lobes 2.5—4.5 mm long, apex mostly acuminate

or rarely acute, villous or tomentose, persistent

in fruit, mostly reflexed towards body of fruit.

Petals white to magenta, elliptic to ovate or

obovate, 8-13 mm long, upper surface glabrous,

glabrescent or sparsely sericeous, lower surface

sericeous to villous. Stamens 5—9 mm long,

anther connectives glandular. Style 9-14 mm

long, flexuous, sparsely sericeous. Fruit

subcylindrical, base rounded, 7-12 mm long,

6—9 mm wide, sparsely tomentose to tomentose,

yellow-green or olive-green to whitish.

Peduncle 3-14 mm long in fruit. Radicle tip

somewhat spirally contorted above or below

plane of embryo. Fig. 4P—W.

Selected specitinens: Northern Territory. DARWIN AND

GuLF District: Headwaters East Alligator River, 12°47’S,

133°21°E, May 1997, Guymer 2503 et al. (BRI, DNA,

CANB); Arnhem Land, edge or Wellington Range, c. 40

Snow & Guymer, Systematic and cladistic studies of Myrtella and Lithomyrtus (Myrtaceae) 185

km SSE of Murganella settlement, 11°49’°34.7"S,

133°03715.3"E, May 1997, Snow & Mangion 7420 (AD,

BRI, CANB, DNA, K, NSW, MEL, MO, PERTH);

Kakadu N.P., ca 25 km NE of Jabiru, 12°30’30.4"S;

132°57’°08.1°E, May 1997, Srow 7437 et al. (BRI,

CANB, DNA, L, MO, US); Head of Koolpin Creek,

13°26’°06”"S, 132°39°13”’E, Apr 1995, Leach 4382 &

Greshke (DNA); Kakadu N.P., Mt Brockman, c. 16 km

SE of Jabiru, 12°47°37.8°S, 132°55°42.9"E, May i997,

Snow 7447 et al. (BRI, CANB, DNA, L. NSW, PERTH);

i—2 miles S [of] Cannon Hill, Aug 1972, Maritenz AE261

(BRI, DNA); East Alligator River near Cahills Crossing

on road to Oenpelli, 12°27’S, 132°58’E, Jun 1974, Pullen

9446 (BRI, CANB, DNA, MEL); 10 km W [of] Island

Lagoon, Mudginberry Stn, May 1976, Brow s.n. (BRD);

Arnhem Land, Magela Creek upper catchment,

12°49°19"S, 133°00°25”"E, Apr 1995, Cowie 5608 &

Brennan (BRI, DNA); Deaf Adder Gorge, 13°07’S,

132°56’E, Apr 1980, Dunlop 5466 (AD, BRI, CANB,

DNA); Little Nourlangie Rock, 12°52’S, 132°48’E, Aug

1978, Dunlop 5005 (BRI, CANB, DNA, MEL); Kakadu

N.P., Twin Falis, 13°19°S, 132°50’E, Sep 1980, Dunlop

5539 (BRI CANB, MEL); c. 4 miles NNE of Mudgtinberry

HS, Jul 1972, Lazarides 7526 (BRI, CANB); 3 km S of

Naradge Ck crossing, 500 m W of Oenpelli Road,

12°28’S, 132°53’E, May 1988, Bishop 840 (DNA);

Jabiluka Lease, sandstone outcrop at north end, 12°29’S,

132°54’E, Jun 1992, Taylor 103 (DNA); 11.5 km NE of

Jabiru E, 12°35’S, 132°58’E, May 1980, Craven 5939

(CANB, DNA); Arnhem Land, sources of Goomadeer

River, 12°34’S, 133°23°E, Jun 1978, Henshali 1987

(DNA); 25 km from Jabiru towards Oenpelli, 12°32’S,

132°55’E, May 1988, Adunir 5771 (DNA); 16 km SW of

East Alligator River crossing, 12°28’S, 132°55’E, Jun

1972, Maconochie 2273 (DNA); Koongarra Saddie,

Kakadu N.P., 12°51°S, 132°51’E, Aug 1985, Wightnian

2032 (DNA); Nabarlek, 12°19°S, 133°19’E, Jun 1988,

Hinz 34 (DNA); Kakadu N.P., Upper Gimbat Creek,

13°34°S, 133°00’E, Apr 1990, Cowie 1151 & Leach(BRL,

DNA, MEL); Koongarra Jump-up, [2°51°S, 132°50°E,

May 1978, Rice 2617 (CANB); 17,5 km NE of Jim Jim

Falls, 13°08.5’S, 132°56’E, May 1980, Craven 6108 (CANB).

Distribution and habitat: Darwin and Gulf

Region, Northern Territory. Growing in rocky

areas amongst the escarpments and slopes of

Kakadu National Park and in Arnhem Land, in

sandy or rocky soils.

Phenology: Flowering mostly April to July;

fruiting May to November.

Notes: Plants of Lithomyrtus cordata are the

tallest-growing in the genus. Their leaf shape

varies from broadly ovate to narrowly ovate or

almost elliptic with bases truncate or cordate.

This is the only species of Lithomyrtus typically

having both abaxial leaf glands and prominent

abaxial lateral veins, although the latter are not

always conspicuous. This combination also

occurs rarely in L. obtusa but these species are

allopatric. The ferruginous foliage glands on

Gittins 2886 (BRI, CANB) from near the East

Alligator River suggests hybridisation with L.

retusa. Several of the specimens cited by Scott

(1978) as paratypes of Myrtella cordata have

been found to belong to other species of

Lithomyrtus.

Conservation status: The species is widespread

and can be locally common.

2. Lithomyrtus densifolia N.Snow & Guymer,

sp. nov. Plantae erectae, plerumque

minores quam | metrum altae, in ambitu

rotundatae, dense foliosae ramosaeque.

Folia anguste elliptica, subtus glandes

inconspicuas et venas secundarias

laterales prominentes gerentia. Fructus

plerumque cylindrici, glabri usque sparse

tomentosi. Typus: Australia. Northern

Territory. DARWIN AND GULF REGION: Up-

per East Alligator River, Arnhem Land,

12°49’°30"S, 133°21°59”E, 13 May 1997;

C. Mangion 400 & PS. Short (holo: DNA;

iso: AD, BRI, CANB, CHR, K, HO, L,

NSW, MEL, GREE, MO, PERTH, UPS,

US [Note: type material represents either

of the two genotypes collected, indicated

on labels as A collection or B collection).

Mostly erect, compact, densely branching

shrubs, 0.3—-1 (—1.5) m tall. Bark stringy,

orangish or brown. Branchlets tomentose, oil

glands lacking. Leaves opposite, evenly

distributed, soft (younger) or coriaceous (older).

Petioles 1-1.5 mm long, rounded, eglandular

or glandular. Leaf blades narrowly elliptic, 6—

15(-20) mm long, 1.5—3.5 mm wide, base

cuneate, free from the stem, apex obtuse,

margins revolute or flat, upper surface

glabrescent or sparsely short- villous, adaxial

midvein impressed, lower surface villous to

densely villous, oil glands of lower surface

visible and about the same size (but usually

faint), marginal veins of lower surface invisible

or indistinct. Peduncles rigid, 5-12 mm long,

villous. Bracteoles 3-5 mm long 0.5—0.7 mm

wide, the tips (in flower) not exceeding base of

sepal lobes, villous to tomentose, caducous in

fruit. Hypanthium tomentose. Sepal lobes 2.5

mim long; apex acute, tomentose, persistent in

186

fruit, mostly ascending and above body of fruit.

Petals pink to magenta, ovate to obovate, 5—

8.5 mm long, upper surface mostly glabrous,

lower surface sericeous to villous. Stamens 2—

7 mm long, anther connectives glandular. Style

5—8 mm long, flexuous, glabrous. Fruit

cylindrical or rarely fusiform, base rounded or

tapered, 6-10 mm long, 3—5.5 mm wide,

glabrescent to sparsely tomentose, yellow-green

or olive-green. Radicle tip spirally contorted

above or below plane of embryo, or held at same

horizontal plane as embryo. Fig. SA-G.

Selected specinens: Northern Territory. DARWIN AND

GULF ReGcion: Arnhem Land, 15 km SE of Jabiru,

12°47°S, 132°56’E, Apr 1989, Johnson 4770 (BRI);

Kakadu N.P., 14.9 km (by road) from Pine Creek/Jabiru

road (Kakadu Hwy), on road to Koongarra Saddle, c. 16.3

km S of Jabiru, 12°50’S, 132°51’E, May 1992, West 5311

(CANB, DNA); Mt. Gilruth area, in sand amongst rocks

on sandstone escarpment, 12°58’S, 133°10’E, Jun 1978,

Dunlop 4877 (DNA, PERTH); Koongarra area, | km S

of jumpup, 12°51°S, 132°S1’E, Apr 1979, Rankin 1981

(BRI, DNA); Kakadu N.P., 6 km SW of Mt. Brockman,

12°46’°S, 132°54’E, Apr 1980, Zelford 8086 & Wrigley

(CANB); Headwaters of the Liverpool River, Arnhem

Land, 12°46’S, 133°44’E, Apr 1984, Wightman 1444 &

Craven (BRI, CANB, DNA); Headwaters East Alligator

River, 12°47,76S, 133°21.61S8, May 1997, Guymer 2505

etal. (BRI, DNA, CANB); Kakadu N.P., Little Nourlangie

Rock, 12°52’S, 132°47°E, Aug. 1980, Telford 8459 &

Wrigley (CANB); Vicinity of Nourlangie Rock, Jul 1972,

Martensz AE114 (DNA); Giddy River area, Gove, Oct

1971, Hinz 71-1610B (DNA); Arnhem Land, 12°54’S,

135°27°E, Jun 1972, Latz 2957 (CANB, DNA); Doyndji

area, 12°S5’S, 135°24’E, Oct 1976, Scarlett 30 (DNA);

1.5 km NE of Koongarra, 12°52’S, 132°51’E, Sep 1978,

Rankin 1390 (CANB, DNA); Mt. Brockman, 12°46’S,

132°54’E, Apr 1980, Dunlop 5490 (CANB, DNA),

Koongarra Saddle, NE of Nourlangie Rock, 12°S1’S,

132°52’E, May 1988, Munir 5750 (DNA); Upper East

Alligator River, Arnhem Land, 12°49’S, 133°21°E, Apr

1988, Russell-Smith 5226 & Lucas (DNA); c. 17 km SE

of Jabiru, 12°47.5’°S, 132°57,5’E, Craven 6595 (CANB,

DNA, MEL); Buffalo Springs, Mt. Brockman, 5 km NE

of Koongarra, 12°50’S, 132°53’E, May 1980, Lazarides

8910 (CANB, DNA); Arnhem Land, Magela Creek upper

catchment, 12°49°02”S, 133°00719”E, Apr 1995, Cowie

5672 & Brennan (CANB, DNA); Kakadu N.P., Mt.

Brockman, 12°50’00”S, 132°55’17°E, Mar 1995, Kean

4624 (DNA); Kakadu N.P., 12°50’S, 132°S51’E, May

1988, Adunir 5717 (DNA); Kakadu N.P., Mt. Brockman

area, 12°48’S, 132°54°E, Mar 1995, Russell-Smith 10239

(DNA); Kakadu N.P., Koongarra Saddle, 12.51S,

132,51E, Nov 1991, Leach 2904 & Dunlop (CANB,

DNA); Little Nourlangie Rock, 12°52’S, 132°48’E, Mar

1979, Dunlop 4948 (CANB, DNA); Near Koongarra

Saddie, 1.5 km N of Koongarra, 12°51.5’S, 132°51.5”E,

May 1980, Craven 5717 (CANB, DNA); Mt. Gilruth,

Feb 1977, Dunlop 4431 (CANB); Kakadu N.P., Little

| Austrobaileya 5(2): 173-207 (1999)

Nourlangie Rock, 12°52’S, 132°47°E, Apr 1980, Telford

7789 & Wrigley (CANB); Koongarra, 12°51°S; 132°52’E,

May 1978, Rice 2637 (CANB).

Distribution and habitat: Northern Territory;

Darwin and Gulf Region. Sandstone

escarpments, scree slopes and ledges, in open

shrublands; soils skeletal or sandy.

Phenology: Flowering February to June;

fruiting April to November.

Notes: Lithomyrtus densifolia and L. dunlopii

are distinct in the field but herbarium material

can be very similar in appearance. In the field,

L. densifolia is a compact, spreading, densely

branching shrub usually less than a metre tall,

with abundant grey-green leaves occurring

densely throughout whereas L. dunlopii is a

stragely, openly branching shrub to 1.5 m. The

peduncles of mature fruits of L. dunlopii

generally exceed 10 mm in length, whereas

those of L. densifolia are usually shorter than

10 mm. The frutts of L. densifolia are highly

aromatic when crushed.

Conservation status: Although the species Is

not locally common, its wide distribution does

not suggest any vulnerability at this time.

Etymology: The specific epithet is derived from

Latin and refers to the close spacing of the

leaves.

3. Lithomyrtus dunlopii N.snow & Guymer

sp. hov. Plantae erectae. Folia anguste

elliptica, venis secundartis lateralibus

prominentibus plerumque carentia. Pe-

dunculi fructum folia plerumque paulo

longiores. Fructus cylindrici, raro

fusiformes, sparse villosi. Typus: Aus-

tralia. Northern Territory: Arnhem Land,

along mostly abandoned track c. 7.1 km

NE of Murganella Settlement,

11°32°08.7°S, 132°56’13.7°E, May

1997, N.Snow 7419 & C.Mangion

(holo: DNA; itso: BRI, CANB, K, L,

MEL, MO, NSW, US).

Erect shrubs 0.7~-1.5 (—2) m tall. Bark stringy,

brown or orangish. Branchlets villous or

tomentose, oil glands lacking. Leaves opposite,

mostly evenly distributed, soft but becoming

coriaceous. Petioles 1.5—4 mm long, rounded,

Snow & Guymer, Systematic and cladistic studies of Myrtella and Lithontyrtus (Myrtaceae) 187

eglandular. Leaf blades narrowly elliptic, 10-

40(—55) mm long, (3—)6-10 mm wide, base

narrowly cuneate, free from the stem, apex

obtuse, margins revolute or flat, upper surface

glabrescent to sparsely short villous, adaxial

midvein impressed, lower surface villous to

tomentose, oil glands of lower surface not

visible, marginal veins of lower surface invisible

or indistinct. Peduncles rigid, sparsely villous

to villous, 10-30 mm long. Bracteoles 3—3.6

mm long, c. 0.5 mm wide, tips (in flower)

exceeding base of sepal lobes or not, tomentose,

caducous in fruit. Hypanthium tomentose. Sepal

lobes 1.5—-3.3 mm long; apex acute or

acuminate, sparsely tomentose to tomentose,

persistent in fruit, mostly ascending and above

body of fruit. Petals white to magenta, ovate to

broadly ovate, 5.56.8 mm long, upper surface

sparsely sericeous, lower surface tomentose.

stamens 4—5 mm long; anther connectives

glandular. Style 4-5 mm long, flexuous or

mostly straight, glabrous or sparsely sericeous.

Fruit cylindrical or rarely fusiform, base tapered,

7-10 mm long, 4-5.5 mm wide, sparsely villous,

yellow-green or olive-green. Radicle tip

somewhat spirally contorted above or below

plane of embryo. Fig. SH—N.

Selected specimens: Northern Territory. DARWIN AND

GuLF Recion: Arnhem Land, along track bearing east-

west towards Gningarg Point, from Murganella

settlement, 11°287°30.8°S; 132°56°31.6"E, May 1997,

Snow 7418 & Mangion (BRI, CANB, DNA, K, L, NSW,

MO, PERTH, US); Kakadu N.P., c. 25 km NE of Jabiru,

12°29°58.3"8; 132°56’52.0"E, May 1997, Snow 7441

et al, (BRI, CANB, DNA, NSW, K, MO); Arnhem Land,

edge of Wellington Range, c. 40 km SSE of Murganella

settlement, 11°49°34.7°S; 133°03’15.3”, May 1997,

Snow 7421 & Mangion (BRI, UPS); Kakadu N.P., 6.5

km SSW of Mt. Brockman, 12°48’S, 132°56°E, Telford

8040 & Wrigley (CANB); Arnhem Land, Flinders

Peninsula, 12°06’S, 135°59°E, May 1992, Cowie 2768

(DNA,MEL); c. 22 km NE of Jabiru, 12°31’S,

132°58.5°E, Mar 1981, Craven & Whitbread 6604

(CANB,DNA,MEL); Upper Liverpool River, 12°26’S,

134°04’E, Sep 1976, Duncan s.n. (DNA); Kakadu N.P.,

near Cannon Hill, 12.298, 132.55E, Mar 1983, Russell-

Smith 522 (DNA); 15 km NNE of Jabiru East, 12°32’S,

132°57°E, Jun 1980, Craven 6458 (DNA); Headwaters

of East Alligator River, Arnhem Land, 12°48’S,

133°21°E, Mar 1984, Wightman 1376 & Craven (BRI,

CANB, DNA); Nabariek, 12°17’S, 133°19’E, Apr 1979,

Rankin 2093 (BRI, CANB, DNA); Mt. Brockman,

12°45°S, 132°56’E, Apr 1980, Dunlop 5501 (CANB,

DNA); Upper East Alligator River, Arnhem Land, 12.508,

133,.20E, Apr 1988, Russell-Smith 5268 & Lucas (BRI,

DNA); c. 8 km NNE of Jabiru, 12°35753”S, 132°59’34”E,

Apr 1995, Egan 4826 (BRI, CANB, DNA); 11.5 km NE

of Jabiru East, 12°35’S, 132°58’E, May 1980, Craven

5938 (CANB, DNA); Arnhem Land, Murganella

Settlement, vicinity and west of Workshop Road,

11°32748.3"S, 132°55’25.3°E, May 1997, Snow 7414 &

Mangion (BRI, DNA, L, UPS); Kakadu N.P., c. 25 km

NE of Jabiru, 12°30’°24.9"S, 132°57°05,5”E, Snow 7432

et al. (BRI, DNA); Giddy River area, Gove, Oct 1971,

Hinz 71-1610B (CANB, DNA); Nabariek, 12.2058,

133.16E, Hinz 46 (DNA); North Caledon Bay, Oct 1968,

Byrnes NB964 (DNA); Gimbat River Station, source of

South Alligator River, 13°45’S, 132°38°E, Jul 1983,

Russell-Smith 748 (DNA); 15 km NNE of Jabiru East,

12°32’S, 132°57’E, Jun 1980, Craven 6458 (CANB);

Cato River Rd, 6.5 km N Dahlinbury turnoff, 12°21’S,

136°25’E, Sep 1987, Clark 1694 (DNA); Marchinbar

Istand South, Hopeful Bay, 11°25°38”S, 136°29°60”E,

Sep 1994, Brennan 2900 (DNA); Wessel Islands, 11°26’S,

136°31°E, Oct 1972, Latz 3480 (DNA).

Distribution and habitat: Northern Territory;

Darwin and Gulf Region. Occurs in heaths,

woodlands or A/losyncarpia ternata S.J.Blake

forests, along sandstone escarpments and

outcrops; soils sandy or lateritic.

Phenology: Flowering April to August; fruiting

April to November.

Notes: L. dunlopii most resembles L. densifolia.

The tips of the bracteoles of L. dunlopii

occasionally exceed the base of the sepal lobes

(e.g., Russell-Smith 5381 & Lucas (BRI); kgan

4826 (BRI)), an attribute which otherwise

usually occurs only inl. obtusa. However, the

known distributional ranges of L. .dunlopii and

L. obtusa do not overlap. A few specimens of

L. dunlopii with narrowly elliptic leaves

approach L. finariifolia (Egan 4826 [BRI}), but

the dense pubescence on the abaxial leaf surface

distinguishes them from the latter. The three

specimens (Clark 1694, Brennan 2900, Latz

3480) need further study; they resemble

specimens of L. densifolia, but are tentatively

placed under L. dunlopii here because oil glands

are not clearly visible on the abaxial leaf

surfaces.

Conservation status: The frequent occurrence

of subpopulations in excess of fifty individuals,

recruitment of seedlings in extant subpopulations

(Snow, pers. obs.) and the known number of

collections indicate this species 1s not of

conservation concern at this time.

Etymology: The specific epithet honours Clyde

Dunlop of the Northern Territory Herbarium,

Austrobaileya 5(2): 173-207 (1999)

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Figure 5. A-G: Lithomyrtus densifolia, A. fruiting branchlet x 2. B. section of branchlet x 16. C. abaxial leaf profile

x 4, D. adaxial leaf apex = &. E. flower x 2. F. bracteole x 8. G. fruit x 2. A-D, G: Mangion 400B & Sharp. E-F: Rutssell-

Smith 10239, H-N: L. duntopit. H. fruiting branchlet x 1. I. section of branchlet x 16. J. abaxial leaf profile x 2. K. adaxial

leaf apex x 6. L. flower and peduncle x 2. M. bracteole x 8. N. fruit x 2. H-N: Snow 7419 ef al. O-Y: L. grandifolia, O.

fruiting branchlet x 0.5. P. section of branchlet x 8. Q. abaxial leaf profile x 0.5, R. adaxial leaf apex x 1. S. flower = 2. T.

bracteole x 8, U. fruit x 1.5. V. detail of sepal lobe and fruit x 4. O-P: Snow 7444—B ef al. Q—-R, U—V: Snow 7443 et al. S—

T: Wightman 1377 & Craven, All DNA.

snow & Guymer, Systematic and cladistic studies of Myrtella and Lithomyrtus (Myrtaceae) 189

who has made many important collections of

this taxon and other species of Lithomyrtus.

4. Lithomyrtus grandifolia N.Snow &

Guymer sp. nov. Plantae adscendentes

usque erectae. Folia magna, late elliptica

usque ovata, basibus cordatis autem haud

amplexicaulibus; pagina inferior foliorum

dense hirsuta, venas secundarias laterales

prominentes gerens sed glandibus carens,

Fructus subglobosi usque globosi, dense

tomentosi. Typus: Australia. Northern

Territory. DARWIN AND GULF REGION:

Kakadu National Park, c. 25 km NE of

Jabiru, underneath overhang of large sand-

stone outcrop, 12°29°12.2”S, 132°56’°37.8°E,

15 May 1997, N.Snow 7444 et al. (holo:

DNA; iso: BRI, CANB, K, L, MO).

Shrubs 0.62 m tall, stems erect (or merely

ascending). Bark stringy, orangish. Branchlets

tomentose, oil glands lacking. Leaves opposite,

mostly evenly distributed, soft, Petioles 1.84

mm long, channelled, eglandular. Leaf blades

ovate, (17—)25—61 mm long, 10-46 mm wide,

base cordate, clasping (occasionally, especially

the youngest leaves) or free from the stem, apex

obtuse to acute (rarely), the margins flat, upper

surface glabrescent to sparsely tomentose,

adaxial midvein impressed, lower surface

tomentose, glands of lower surface not visible,

marginal veins of lower surface prominent.

Peduncles rigid, 9-51 mm long in flower, villous

to tomentose. Bracteoles 4-6 mm long, c.1 mm

wide, tips (in flower) not exceeding base of

sepal lobes, densely villous to tomentose,

persistent in fruit. Hypanthium tomentose. Sepal

lobes 4—7 mm long, apex acuminate, villous to

tomentose, persistent in fruit, mostly reflexed

towards body of fruit. Petals pink, elliptic to

ovate, or obovate, 9-11 mm long, upper surface

glabrous, lower surface villous to tomentose.

Stamens 6—8 mm long; anther connectives

eglandular. Style 5—7 mm long, flexuous,

glabrous. Fruit mostly globose, base rounded,

11-13 mm long, 8-10 mm wide, tomentose,

whitish. Radicle tip not spirally contorted, held

at same plane as embryo. Fig. 5O—Y.

Specimens examined: Northern Territory, DARWIN AND

GULF REGION: Kakadu N.P., c. 25 km NE of Jabiru,

12°30°15.2"S, 132°57°08.6"E, May 1997, Snow 7430 et

al. (BRI, DNA); Kakadu N.P., c. 25 km NE of Jabiru,

near top of sandstone cliff arising some 20 metres,

12°30738.7°S, 132°57°06.47°E, Snow 7439 et al. (BRI,

DNA, MEL, PERTH, UPS); Kakadu N.P., c. 25 km NE

of Jabiru, deeply dissected sandstone crevices,

12°30°30.4"S, 132°57°08.1°E, Snow 7438 et al. (BRI,

DNA, GREE, NSW, US); Kakadu N.P., c. 25 km NE of

Jabiru, 12°30°24.9"S, 132°57°05.5”"E, Snow 7433 et al.

(BRI); Kakadu N.P., remote area c. 25 km NE of Jabiru,

12°30°15.2”S, 132°57°08,6”"E (BRI, DNA); Kakadu N.P.,

c, 25 km NE of Jabiru, very base of vertical sandstone

outcropping in sandy soil, 12°29°28,5"S, 132°56’40.8”E,

May 1997, Srow 7442 et al. (BRI, DNA, CANB, K, EL,

MO); Kakadu N.P., c. 25 km NE of Jabiru, underneath

overhang at very base of sandstone cliff in complete

shade, 12°29°16.2”S, 132°56’40.1"E, May 1997, Snow

7443 et al. (BRI, DNA, K, MO); Northern Kakadu,

12°32°S, 132°39”"E, Apr 1983, King 226 (DNA);

Headwaters of the East Alligator River, Arnhem Land,

{2°48’S, 133°21°E, Mar 1984, Wightman 1377 & Craven

(AD, BRI, CANB, DNA, PERTH); East Alligator River,

13°01’S, 133°25°E, Nov 1987, Dunlop 7308 (BRI, DNA);

ESE Mudginberry, 12°36’S, 132°58’°E, Feb 1973, Dunlop

3297 (BRI, DNA); East Alligator River, 38 miles SSE of

Oenpelli, 12°47’°S, 133°21’E, Jul 1972, Adams 2866

(BRI, DNA); Magela Creek, Sept 1970, Byrnes 1973

(DNA); Nabarlek, 12°30’S, 133°21”E, Feb 1989, Hinz

418 (CANB, DNA); 19 km E of Jabiru, 12°37’S,

133°03°E, Russeil-Smith 8032 & Lucas (DNA); 18 km

NNE of Jabiru East, 12°30.5S, 132°57°E, Jun 1980,

Craven 6360 (CANB, DNA); 14.5 km NE of Jabiru East,

12°33.5’S, 132°59’E, May 1980, Craven 5967 (CANB,

DNA); Kakadu N.P., 7 km NNE of Jabiru, 12°35’52”S,

132°59°27°E, Apr 1995, Egan 4782 & Knox (DNA); 20

km S of Nabarlek, 12°30’S, 133°21’E, Jul 1989, Hinz

551 (DNA); Upper East Alligator River area, Arnhem

Land, 12°39’S, 133°29°E, Feb 1991, Russell-Sniith 8424

& Brock (DNA); 44 km SE Oenpelli, 12°34’S, 133°23’E,

Jun 1978, Dunlop 4916 (DNA); Arnhem Land, 19 km E

of Jabiru, 12°37°S, 133°03’E, Apr 1989, Jo/mson 4514

(BRI).

Distribution and habitat: Northern Territory,

Darwin and Gulf Region. Often growing in the

semi-shaded overhangs at base of sandstone

escarpments; in sandy soils.

Phenology: Flowering February to May;

fruiting April to November.

Notes: The species is unmistakable in the genus

because of its leaf size. It is prone to mealy

bug and ophiostomoid fungal infestations (Snow,

pers. obs.).

Conservation status: Given the relatively few

collections, subpopulation sizes of usually less

than fifteen individuals, and the observed lack

of seedling recruitment (Snow and Guymer,

pers. obs.), the species 1s considered vulnerable

(criterion D1).

190 Austrobaileya 5(2): 173-207 (1999)

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Figure 6. A-G: Lithomyprtus hypoleuca. A. fruiting branchlet x 1, B. section of branchlet x 8. C. abaxial leaf profile

x 2. D, abaxial leaf apex x 4 E. bracteole <x 16 F. flower x 3 G, fruit x 2, A-D, G. Zatz 7335 (DNA). E-F. Egan

4861(DNA). H-N: LZ. kakaduensis. H. flowering branchlet x 1. 1. section of branchlet x 8. J. abaxial leaf profile x 2.

K. adaxial leaf apex x 4. L. flower x 2. M. bracteole x 8. N. fruit x 1.5. H—-N. Syow 7450 ef al. (BRI) O-U: Z£,

linariifolia. ©. flowering branchiet x 1. P. section of branchlet x 16. Q. abaxial leaf profile x 2. R. abaxial leaf apex x

[6.S. flower x 3. T. bracteole x 16. U. fruit x 2, O-T: Craven 8248 & Wightman (DNA), U: Cowie 1109 & Leach (BRD.

Snow & Guymer, Systematic and cladistic studies of Myrtella and Lithomyrtus (Myrtaceae) 19]

Etymology: The specific epithet is derived

from Latin and refers to the large leaves of

the species.

5. Lithomyrtus hypoleuca F.Muell, ex N.Snow

& Guymer sp. nov. Plantae erectae. Folia

plerumque elliptica; pagina inferior

foliorum dense hirsuta, glandibus nullis

vel raro praesentibus et venis secundariis

lateralibus prominentibus carens. Fructus

plerumque cylindrici, glabrescentes usque

villosi. Typus: Australia. Northern Terri-

tory. DARWIN AND GULF REGIon: McArthur

River station, Abner Ranges, 16°44’45’S,

13S°S7°31°E, 21 May 1997, CL. R.

Michell & D. S. Calliss 869 (holo: DNA;

iso: BRI, CANB, K, MEL, MO, NSW),

Shrubs 0.5—1.5 m tall, stems erect or ascending.

Bark smooth or stringy, brown, grey or orangish.

Branchlets tomentose, oi] glands lacking.

Leaves opposite, mostly concentrated near

branch tips or evenly distributed, soft but

becoming coriaceous. Petioles |—2.5 mm long,

channelled (slightly), glandular. Leaf blades

narrowly elliptic to ovate, 10-33 mm long, 6—

12 mm wide, base truncate or cuneate, apex

obtuse, margins revolute, upper surface

glabrescent to sparsely short villous, adaxial

midvein impressed, lower surface tomentose,

glands of lower surface not visible, marginal

veins of lower surface invisible or absent.

Peduncles rigid, 6.5-19 mm long, sparsely

villous to villous. Bracteoles 3-4.7 mm long,

0.40.7 mm wide, tips (in flower) not exceeding

base of sepal lobes, densely villous, caducous

in fruit. Hypanthium densely villous. Sepal

lobes 3.2—-4 mm long, apex acute or acuminate,

villous, persistent or deciduous in fruit, mostly

reflexed towards body of fruit. Petals pink,

ovate to obovate, 5.5—-9 mm long, upper surface

glabrous, lower surface villous. Stamens 2.5—6

mm long; anthers connectives eglandular. Style

3—9 mm long, flexuous, sparsely sericeous. Fruit

subcylindrical, base rounded, 6-9 mm long,

4.5—7 mm wide, glabrescent to villous, whitish.

(Embryos not seen.) Fig. 6A—G.

Specimens examined: Northern Territory. DARWIN AND

GULF ReGion: 14 km NE of Mann River Gorge, 12°39°S,

134°08’E, Nov 1987, Leach & Dunlop 1594 (BRI, DNA);

Near Magela Creek, 13.5 km SE of Jabiru East, 12°45’S,

132°59°E, Jun 1980, Craven 6337 (CANB); Headwaters

E. Alligator River, 12°47°17"S, 133°21’61°E, May 1997,

Guymer 2501 et al. (BRI, CANB, DNA, K, MO); Mt.

Brockman, 12°46’S, 132°54’E, Apr 1980, Dunlop 5489

(BRI, CANB); Kakadu N.P., Birdie Creek, 13°57’S,

132°52’K, Apr. 1990, Cowie 1110 & Leach (BRI); Cox

River Station, 16°O1°S, 134°46’E, Jul 1977, Zatz 7335

(DNA); 10 km NNE of Jabiru, Kakadu N.P., 12°34’54"S,

132°59°10”"E, Apr 1995, Egan 4861 (CANB, DNA);

Kakadu N.P., 5 km § of Cahill’s Crossing, East Alligator

River, 12°28’S, 132°57’°E, Apr 1983, Thompson 306

(CANB); 6km SW of Mt. Brockman, 12°46’S, 132°54’E,

Apr 1980, Telford 8085 & Wrigley (CANB); Beetle

Spring, 20 km SE of McArthur River Stn Hmstd, 16°47’S,

135°S9’E, Jun 1990, Menkhorst 967 (DNA, MEL); Upper

East Alligator River, Arnhem Land, 12°49’S, 133°22’E,

Russell-Smith 5914 & Lucas (DNA); 10 miles SSW of

Borroloola, Jun 1971, Dunlop 2217 (BRI, CANB),

Queensland. Burke District: Hells Gate, 17°28’S,

138°22°E, May 1974, Carolin 9171 (BRI); Hells Gate

between Doomadgee and Westmoreland Station, May

1974, Pullen 9177 (BRI, CANB); Westmoreland, Big

Amphitheatre, 17°24°08”S, 138°1S5’25°E, May 1997,

Forster PIF21046 (BRI); Westmoreland Station, 70 km

from Wollogorang towards Burktown, 17°20’S,

138°15’E, Jul 1972, Gittins 2492 (BRI). Coox District:

Gilbert River Holding site EU 553, 19°04’54"S,

143°31°32”E, Apr 1994, Godwin s.n. (BRI [AQ

630612]).

Distribution and habitat: Northern Territory,

Darwin and Gulf Region; Queensland, Burke

and Cook Districts. Occurring on rocky

slopes and sandstone escarpments in open

woodlands.

Notes: Lithomyrtus hypoleuca is the most

difficult species of this genus to identify because

it can resemble several other species. It most

resembles LZ. densifolia, but can be distinguished

from this species by almost always lacking

abaxial leaf oil glands, generally having larger

and fewer leaves, and having eglandular anther

connectives. Embryo characters were not seen

in the nine fruits sampled. The species was not

seen in the field.

Phenology: Flowering April to June; fruiting

June to November.

Conservation status: Aithough a recent

collection (Forster PIF21046) records the

species as locally common in the Westmoreland

area, the few collections in herbaria suggest

the species should be considered vulnerable

(criterion D1) pending further information.

Etymology: The specific epithet is from Greek

hypo-, below or under, and /euca, white, a

reference to the white underside of the leaves.

192

6. Lithomyrtus kakaduensis N.Snow &

Guymer sp. nov. Plantae plerumque

erectae. Folia ovata, apetiolata, basibus

cordatis ac amplexicaulibus; pagina infe-

rior foliorum dense hirusta, venas

secundarias laterales prominentes gerens

autem glandibus carens. Fructus

subglobosi usque subcylindrici, villosi.

Typus: Australia: Northern Territory.

DARWIN AND GULF District: Kakadu Na-

tional Park, 5.7 km off Kakadu highway

along road to Gunlom,13°32’20.3"S,

132°20°06.8”"E, 18 May 1997, N.Snow

7450 etal. (holo: DNA; 1so: BRI, CANB,

Kk, L, MEL, MO, NSW, PERTH, US).

Erect shrubs 0.5—2 m tall. Bark smooth, or

stringy, orangish. Branchlets tomentose, o11

glands lacking. Leaves opposite, concentrated

near branch tips or mostly distributed, soft.

Petioles 0.2—0.5(—1) mm long, eglandular. Leaf

blades ovate to broadly ovate, 9-34 mm long,

6—22 mm wide, base cordate, clasping the stem,

apex obtuse, margins revolute, upper surface

sparsely short villous to sparsely tomentose,

adaxial midvein impressed, lower surface

tomentose, oil glands of lower surface not

visible, marginal veins of lower surface

prominent. Peduncles rigid, 10-20 mm long,

sparsely villous to villous. Bracteoles 4-6 mm

long, 0.5—0.8 mm wide, tips Gn flower) not

exceeding base of sepal lobes, villous, persistent

or (usually) caducous in fruit. Hypanthium

villous to densely villous. Sepal lobes 3—5 mm

long, apex acuminate, villous to tomentose,

persistent in fruit, mostly ascending and above

body of fruit. Petals pink, obovate, 7-9 mm

long, upper surface glabrescent to sparsely

sericeous, lower surface sericeous, Stamens 4—

6 mm long; anther connectives glandular. Style

5—6 mm long, flexuous or mostly straight,

glabrous. Fruit subglobose to subcylindrical,

base rounded, 7-14 mm long, 69.5 mm wide,

villous, yellow-green or olive-green or whitish.

Radicle tip somewhat spirally contorted above

or below plane of embryo. Fig. 6H-N.

Specimens examined: Northern Territory. DARWIN AND

GULF ReGIoN: Kakadu N.P., 5.7 km along dirt road to

Guniom from Kakadu Highway, c. 50 m S of road,

13°32°18.5°S, 132°28712.9"E, May 1997, Snow 7448 et

al. (BRI, CANB, DNA, K, L, MO, NSW, PERTH);

Kakadu N.P., Twin Fails, 13°18’S, 132°51°E, Jul 1983,

Dunlop 6723 & Wightnan (BRI, DNA, MEL, PERTH);

Austrobatieya 5(2): 173-207 (1999)

Without precise locality, 13°44’S, 132°40’E, Jul 1972,

Lazarides 7684 (BRI, CANB); Kakadu N.P., 8 km NNE

of Mt. Evelyn, 13°32’S, 132°56’E, Apr 1989, Menkhorst

343 (BRI, DNA, MEL); Headwaters Katherine River,

Arnhem Land, 13°46’15°S, 133°09°03”S, Jul 1996,

Mangion 237 & Dunlop (BRI); Gradys Creek, Arnhem

Land, 13°29’S, 133°O01’E, Jul 1995, Mlerrotsy 2474A

(BRI); Gimbat Station, source of South Alligator River,

13°45°S, 132°38’E, Jul 1983, Russell-Smith 743 (DNA,

MEL); Jim Jim Creek, 3.5 km ESE of Jim Jim Falls,

13°17’S, 132°52’E, May 1980, Craven 5819 (CANB,

DNA); Kakadu N.P., adjacent to Round Jungle, 13°18’S,

132°38’E, Apr 1987, Russell-Smith 2176 & Lucas

(DNA); 75 km NE of Pine Creek along road to El Sharana,

13°32’S, 132°21°E, May 1983, Briggs 925 (BRI, CANB,

DNA); Kakadu Hwy at Gunlom turnoff, 13°33’S,

132°17°E, Dec 1995, Brennan 3191 (DNA); UDP Fails

road, c. 5.5 km E from Kakadu Hwy, 12°32’S, 132°20°E,

Apr 1987, Purdie 3228 (CANB, DNA); Headwaters of

Twin Creek, 13°26’43”S, 133°51°04”E, Apr 1995, Leach

4337 & Greschke (DNA); 31 km WSW of Twin Falls,

13°20’S, 132°29.5’E, Jun 1980, Craven 6404 (CANB,

DNA); 6 km ESE of Twin Falls, 13°22S, 132°48’E, May

1980, Craven 5845 (CANB); Kakadu N.P., Upper Birdie

Creek, 13°53’S, 132°57’E, Apr 1990, Slee & Craven

2526 (CANB) & 2527 (AD, CANB, MEL); Kakadu N.P.,

Kakadu Hwy 1.2 km S of entrance to Mary River Ranger

Station, 13°33’S, 132°16’E, Apr 1990, Slee & Craven

2959 (CANB, MEL).

Distribution and habitat: Northern Territory,

Darwin and Gulf Region. On gentle slopes or

sandstone ridges with Eucalyptus phoenicea F.

Muell., 2. miniata A. Cunn. ex Schauer, £.

tetrodonta or Acacia spp. 1n sandy souls.

Phenology: Flowering December to May;

fruiting May to December.

Notes: Specimens of L. kakaduensisi with oil

glands on the abaxial leaf surface may be

hybrids of this species with L. cordata or L.

retusa (viz Hearne 1740; Craven 6404 [CANB

307316]|). The second accession of Craven

6404 (CANB 307317) is an intergrade of this

species with L. densifolia, as evidenced by its

dense foliage and abundant fruit on short

peduncles.

Conservation status: Although occasionally

occutring in subpopulations greater than 10

individuals (Snow, pers. obs.), the absence of

seedlings and narrow geographic range suggest

the species should be considered vulnerable

(criterion D1).

Etymology: The epithet is a contraction of

‘Kakadu’ and Latin -ensis indicating the

engi tenner nenetentn tare ble meee ne

Snow & Guymer, Systematic and cladistic studies of Myrtella and Lithonryrtus (Myrtaceae) 193

occurrence of the species largely within Kakadu

National Park.

7. Lithomyrtus linariifolia N.Snow & Guymer

sp. nov. Plantae plerumque decumbentes.

Folia linearia, glandibus subtus carentia.

Pedunculi sparse sericei, folia subtenta

usque duplo longiores. Fructus

fusiformes, glabri usque sparse villosi.

Typus: Australia, Northern Territory, Up-

per East Alligator River, Arnhem Land,

12°39°S, 133°23’°E, 20 Feb 1991,

J.Russell-Smith 8435 & Brock (holo:

DNA).

Plants suffrutescent, rarely shrubby, 0.1—0.2 (—

1.0) m tall, stems prostrate or rarely erect. Bark

stringy, brown to orangish. Branchlets sparsely

sericeous (with antrorse hairs bending sharply

at base) or villous, glands lacking. Leaves

opposite, mostly evenly distributed, coriaceous.

Petioles 0,5—1.3 mm long, rounded, eglandular.

Leaf blades linear, mostly 10-51 mm long, 1-3

mm wide, base cuneate, free from the stem, apex

acute, margins revolute, upper surface glabrous

to sparsely sericeous, adaxial midvein

impressed, lower surface somewhat sericeous

to densely villous, glands of lower surface not

visible (usually), or visible and about the same

size (very rarely), marginal veins of lower

surface invisible or indistinct. Peduncles

flexible, 27-58 mm long, sparsely antrorsely

sericeous to sparsely villous. Bracteoles 22.5

mm long, c. | mm wide, tips (in flower) not

exceeding base of sepal lobes, sparsely

sericeous to glabrous, mostly caducous in fruit.

Hypanthium densely sericeous. Sepal lobes 1.1—

2 mm long, sparsely sericeous, persistent in

fruit, mostly ascending and above body of fruit.

Petals pink, elliptic to ovate, 5-7 mm long,

upper surface mostly glabrous, lower surface

sericeous to sparsely villous. Stamens 3—3.5 mm

long; anther connectives glandular. Style 2.5—3

mm long, flexuous or mostly straight, sparsely

sericeous. Fruit fusiform, base tapered, 8.5—11]

mm long, 3.5—4 mm wide, sparsely sericeous

to sparsely tomentose, glabrescent, yellow-

green or olive-green. Embryos not seen. Fig.

60-U.

Selected specinens: Northern Territory. DARWIN AND

GULF REGION: Kakadu N.P., Birdie Creek, 13°57’S,

132°52°E, Apr 1990, Cowie 1109 & Leach (BRI, DNA,

MEL); Head of gorge between Twin Falls and Jim Jim

Falis, 13°19’S, 132°59’E, Mar 1984, Craven 8248 &

Wightman (AD, CANB, DNA, MEL); c. 10 km NNE of

Jabiru, 12°34°23”S, 132°59°16"E, Hean 4849 (CANB,

DNA); Kakadu N.P., 10 km NE Jabiru, 12°34’02”S,

132°59’°02”E, Apr 1995, Russell-Smith 10481 CDNA); 6

km ESE of Twin Falls, 13°22’S, 132°48’E, May 1980,

Craven 5837 (CANB, DNA); | km upstream from Twin

Falls, 13°20’S, 132°47°E, Mar 1988, Fenshan: 775

(DNA); 14.5 km NE of Jabiru East, 12°33.5’S, 132°59’E,

May 1980, Craven 5966 (CANB, DNA).

Distribution and habitat: Northern Territory,

Darwin and Gulf Region. In heaths or eucalypt

woodlands of sandstone escarpments, in sandy

or skeletal souls.

Phenology: Flowering March to April; fruiting

known only from April.

Notes: The linear leaves and fusiform fruits are

distinctive for the species; the sparsely sericeous

midvein on the undersurface of the leaf and the

densely sericeous peduncles are also reliable

diagnostic characters. The fruits are often

irregularly swollen near the midpoint, but this

feature can also occur in L. dunlopii. Embryo

characters have not been seen in the fruits

sampled. The species was not seen in the field.

Conservation status; Few populations of this

species are known and it is considered

vulnerable (criterion D1).

Etymology: The specific epithet is dertved from

Latin and refers to the linear leaves of the

species.

8, Lithomyrtus microphylla (Benth.) N.Snow

& Guymer comb. nov.

_ Fenzlia obtusa var. microphylla Benth., FI.

Austr. 3: 279 (1866); Fenzlia microphylla

(Benth.) Domin, Biblioth. Bot. 89: 476

(1928); Myrtella microphylla (Benth.) A.

J. Scott, Kew Bull. 33: 301 (1978). Type:

Australia. QUEENSLAND: Dividing range

between the rivers Thomson and

Burdekin, J. Sutherland s.n. (holo: K).

Erect or prostrate shrubs 0.3—1.5 m tall. Bark

smooth, or stringy, brown, grey or orangish.

Branchlets villous to tomentose, oil glands

lacking. Leaves opposite or occasionally

whorled, mostly evenly distributed, coriaceous.

Petioles 0.8—1.6 mm long, rounded, eglandular.

194

Leaf blades narrowly elliptic to ovate, 8—30(-—

42) mm long, 1.7—-2(—-4.2) mm wide, the base

truncate or cuneate, free from the stem, apex

obtuse to acute, margins revolute or flat, upper

surface glabrescent to sparsely tomentose,

adaxial midvein impressed, lower surface

tomentose, oil glands of lower surface usually

not visible, marginal veins of lower surface

invisible or indistinct. Peduncles rigid, 3—10 mm

long, villous to tomentose. Bracteoles 2.5-3.2

mim long, 0.4—0.6 mm wide, tips (in flower) not

exceeding base of sepal lobes, villous to

tomentose, persistent or caducous in fruit.

Hypanthium densely villous to tomentose. Sepal

lobes 1.4-—2.4 mm long, apex acute to

acuminate, glabrous to villous (above), or

villous to tomentose (below), persistent in fruit,

mostly appressed towards body of frutt. Petals

white to magenta, elliptic to obovate, 4~4.7 mm

long, upper surface glabrous to sparsely

sericeous, lower surface sparsely villous to

villous. Stamens 2.5—4 mm long; anther

connectives eglandular or glandular. Style 3-4

mm long, flexuous, sparsely sericeous. Stigma

apex terete or occasionally capitate. Fruit

subcylindrical, base tapered, 5—8.4 mm long,

2.5-4.5 mm wide, glabrescent to sparsely

tomentose, yellow-green or olive-green. Radicle

tip not spirally contorted, held at same

horizontal plane as embryo. Fig. 7A—G.

Selected specimens: Queensland. BuRKE District: Burra

Range, 2.5 km from the Great Divide Crest along Flinders

Hay towards Torrens Creek, 20°44’S, 145°13’E, May

1991, Telford 11451 (CANB); 102 km N of Hughenden,

on road to Lynd Junction, Poison Creek, 19°56’S,

144°16’E, May 1990, Halford Q248 (AD, BRI, DNA);

Torrens Creek, Mar 1933, White 8723 (BRI); “Warang”

Holding, White Mountains, c. 37 km NNW of Torrens

Creek township, 20°29S, 144°48’E, Jul 1988, Fel/

DF1292 & Swain (BRI); Coox District: 74 km from the

Chillagoe-Wrotham Park road towards Bulimba, 16°58’S,

143°43°E, Jan 1993, Bean 5656 & Forster (BRD; 87 km

along road to Bulimba Station, off Chillagoe to Wrotham

Park road, 16°54’S, 143°36’E, Feb 1994, Forster

PIF14743 (BRI, DNA, MEL); Mopata H-W boundary on

SWER Line to Robinhood, 18°498S, 143°48’E, Sep 1994,

Godwin MG4i78C (BRD; Poison Creek, 87 miles N of

Hughenden, Sep 1937, Brass & White 68 (BRI). MircHELL

District: Barcaldine Shire; Busthinia Quarry, c, 43 km

E of Barcaldine, 23°34’S, 145°43’E, Anderson 3730

(BRI); 17 km E of Torrens Creek, North Queensiand,

Mitchell District, 20°44’S, 145°11’°E, Jul 1975, Chapman

1359 (BRI, CANB, PERTH); N side of Flinders Highway,

140 km of Charters Towers, 20°44’S, 145°11’E, Jobson

1804 (BRI, MEL,); 41 km NW of Torrens Creek,

20°26°32”S, 144°50’°05”E, Apr 1993, Thonipson

Austrobaileya 5(2): 173-207 (1999)

HUG413 et al. (BRI); “Fleetwood” Station, Aramac, Aug.

1962, Cockburn s.n. (BRI). NorTH KENNEDY District:

Burra Range, 4 km from Great Divide Crest towards

Pentland along Flinders Hwy, 20°41’°S, 145°14’E, May

1991, Telford 11430 & Rudd (CANB); Beside Greenvale-

Charters Towers road, 32.3 km from Greenvale, 19°07’S,

145°20’E, Jun 1989, Bean 1069, (BRD: 5 km SW of

“Milray”’, 20°43’S, 145°41’E, Nov 1991, Thompson 392

and Robins (BRI); 7.5 km SSE of Mt. Cooper HS, 20°35S,

146°48’°E, Jun 1992, Thompson CHAI8 & Sharpe (BRD);

8 miles W of Pentland, Jun 1953, Perry 3553 (BRI, DNA,

MEL, PERTH). Souru KENNepy District: 19,7 km W of

Oxenhope outstation, at Charlies Creek crossing, 21°07’S,

145°30°E, May 1991, Neldner 3190 & Thompson (AD,

BRI); c. 21 km NNW of Yarrowmere Station homestead

on Great Dividing Range, 21°17’°S, 145°48°E, Oct 1983,

Henderson H2856 et al. (BRI, CANB); c. 22.5 km NNW

of Yarrowmere Station homestead on the Great Dividing

Range, 21°16°S,145°49’°E, Oct 1983, Henderson H2846

et al. (BRI, CANB); c. 4 km N of the homestead, on

Moonoomoo Station, 21°46S, 145°058’E, Oct 1983,

Henderson H2776 et al. (BRI); c. 22 km NE of Mirtna

Station homestead on S side-road to the Charters Towers

to Clermont road, 21°12’S, 146°12’E, Oct 1983, Henderson

H2903 et al. (BRI).

Distribution and habitat: Queensland: Cook,

Burke, North Kennedy, South Kennedy, and

Mitchell Pastoral Districts. In open forests,

shrublands and heaths, on ridgetops, in sandy

or gravelly soils; elevation from 170 to 500

metres.

Phenology: Flowering February to November;

fruiting March to November.

Notes: A few specimens of L. microphylla with

longer than average leaves occasionally have a

few oil glands faintly visible on the lower leaf

surface. Some specimens (Bean 5656; Forster

PIF14743) have three leaves per node.

Conservation status: Often common to

abundant locally.

9, Lithomyrtus obtusa (Endl.) N.Snow &

Guymer comb. nov.

Fenzlia obtusa Endl., Ataka Bot.: 19, t. 17

(1834); Myrtella obtusa (Endl.) A. J.

Scott, Kew Bull. 33: 300 (1978). Type:

Australia: QUEENSLAND: Port Curtis Pas-

toral District: Between Curtis and Facing

Islands, & L. Bauer s.n. (holo: W; iso: K

[photo BRI]).

Erect shrubs 0.3—2.5 m tall. Bark smooth or

stringy, brown or grey. Branchlets tomentose,

andasnysrsterrslnncines.

nininddicindkd Maiviesh baat

J =

STUSSY

Snow & Guymer, Systematic and cladistic studies of Myrteila and Lithomyrtus (Myrtaceae) 195

‘i

) *

& a

ans SEG tte q

ee Fi He a

| Fi ata la ati

ee Ue

eee &

eg ee

‘yt

yo aes ae

dit

ioe

*2*

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wi,

1» *

Pad

™

a rf

int rf i!

i eda

hs, —

Figure 7. A-G: Lithomyrtus microphylla, A. fruiting branchlet x 2. B. section of branchlet x 16. C, adaxial leaf

profile x 4. D. adaxial leaf apex x 8. E. flower x 4. F. bracteole x 16. G. fruit x 3. A—-D, G: Forster PIF 16665 (BRI). E—

F: Bean 5758 (BRI). H-N: LZ. obtusa. H. flowering branchlet x 1.5.1. section of branchlet (hairs and fungal bodies) x

16. J. adaxial leaf profile x 1. K. adaxial leaf apex x 2. L. flower x 3. M. bracteole x 8. N. fruit x 3. H—-M: Bailey s.n.

(BRI AQ 040624). N: White 10126 (BRI). O-U: L. repens. O. branchlet x 0.5. P. section of branchlet x 12. Q. adaxial

leaf profile x 1. R. adaxial leaf apex x 2. S. flower x 4. T. bracteole x 16. U. fruit x 3. O: Mangion 218 (DNA). P-R,

U: Snow 7425 et al.(BRI). S, T: Dimlop & Byrnes 2109 (DNA).

196

oil glands lacking. Leaves opposite,

concentrated near branch tips or evenly

distributed, coriaceous. Petioles 1—3 mm long,

channelled, eglandular. Leaf blades narrowly

elliptic to ovate, or obovate, 15-42 mm long,

(4—)7—13 mm wide, base cuneate, free from the

stem, apex obtuse or retuse, margins revolute

or flat, upper surface glabrous, glabrescent or

sparsely short-villous, adaxial midvein

impressed, lower surface tomentose, oil glands

of lower surface not visible, marginal veins of

lower surface mostly invisible or indistinct.

Peduncles rigid, 3-8 mm long in flower, 5—14

mm long in fruit, villous to tomentose.

Bracteoles 4~9 mm long, 0.40.6 mm wide, tips

(in flower) exceeding base of sepa] lobes,

sometimes considerably so, villous to densely

villous, caducous in fruit. Hypanthium

tomentose. Sepal lobes 2.5—4 mm long, apex

acuminate, glabrous (above) or tomentose

(below), persistent in fruit, mostly reflexed

towards body of fruit. Petals pink, ovate to

obovate, 6-9 mm long, upper surface

glabrescent, lower surface glabrous to sparsely

villous. Stamens 4—7 mm long; anther

connectives glandular. Style 5-6 mm long,

mostly straight, sparsely sericeous. Fruit

globose to subcylindrical, sometimes slightly

tapered above, base slightly tapered, 5-9 mm

long, 3.5—5 mm wide, glabrescent to sparsely

villous, yellow-green or olive-green to dark

bluish-black (rarely reddish). Radicle tip not

spirally contorted above or below horizontal —

plane of embryo. Fig. 7H-N.

Selected specimens: Queensland. Coox DISTRICT:

Endeavor River, near Marton, 15°28’S, 145°11°E, May

1991, Dunlop 7932 & Clarkson (BRI, CANB, DNA); E

of Escape River, Old Comalco campsite near lake, W of

Sadd Point, c. 11°03’S, 142°45°E, Aug 1978, Kanis 2073

(BRI, CANB); Between Half Moon Creek and Earl Hill

near Cairns, 16°48’S, 145°42’E, Oct 1979, Batianoff

1306 & McDonald (BRI); Mt. Saunders, near Cooktown,

15°25°S, 145°14°E, Mar 1984, Scarth—Johnson 1439A

(BRI); Wangetti Beach, Cook Hwy, 16°40’S, 145°34”E,

Sep 1992, Gray 5556 (QRS); Fitzroy Island, near Cairns,

16°56’S, 146°00’E, May 1968, Berry 14942 (QRS); Road

to Cape Flattery, c. 0.5 km W of beach, 15°07’S,

145°13°E, Jun 1992, LeCussan 98 (QRS); 5 miles N of

crossing on Massey Creek on road between Silver Plains

Station and Rocky River, c. 13°50’S, 143°29’E, Oct 1969,

Webb & Tracey 9732 (BRI); Cape York, 100 metres from

the tip, 10°40’S, 142°35’°E, Aug 1986, Archer s.n. (DNA);

Headland between Narau and Nanthau beaches,

Newcastle Bay, 10°47’S, 142°35’E, Feb 1990, Forster

PIF6359 (BRI, DNA); Mt. Tozer summit area, Iron Range

Austrobaileya 5(2): 173-207 (1999)

N.P., 12°45°S, 143°12°E, Jul 1994, Forster PIF15428

(BRI); Coastal Cliffs above Captain Billy Beach, 11°38°S,

142°51’E, May 1980, Morton 636 (BRI, MEL); 8.5 km

N of the Lockhart River Road on the track to Wattle Hill,

12°41°S, 143°05’S, Aug 1991, Clarkson 9077 & Neldner

(BRI, DNA); 3.8 km E of Garraway Creek on road to

Portland Roads, 12°44’S, 143°12’E, Jul 1991, Neldner

3533 & Clarkson (BRI, DNA); Finch Bay, Mar 1966,

Smith 13102 (BRI, CANB); Cooktown, mouth of

Endeavour River, north bank, 15°28’S, 145°15’E, May

1970, Blake 23297 (MEL, PERTH); Moa Island, Torres

Strait, LO°11’S, 142°16’E, May 1953, Marks &

Mackerras s.n. (BRT); 15.1 km S of beachfront at Bathurst

Bay on track to “Wakooka”, Jul 1984, Puttock & King

UNSW16751 (BRI, MEL); Cairns, semi-shade on edge

of swamp, Apr 1967, Brass 33539 (BRI); 2.5 km N of

the mouth of the MclIvor River, 15°07’S, 145°14’°E, Feb

1984, Clarkson 5163 (BRI, MEL); Sharp Point, 10°57’S,

142°43°E, Jun 1978, Clarkson 2104 (BRI); Norru

KENNEDY District: Forrest Beach, c. 12 miles E of

Ingham, May 1970, Fagg 721 (AD, CANB);

Hinchinbrook Island, N end of Ramsay Bay, 18°46’S,

146°18°E, Jul 1988, Thompson 915 (CANB); Sout

KENNEDY District: Shaw Island, 20°29’S, 149°09’E, Nov

1985, Batianoff 3212 & Dalliston (BRI, CANB). Port

Curtis District: Clinton Lowland, Shoalwater Bay area,

22°35’S, [50°42°E, Feb 1992, Thompson 73 (BRI);

Upper reaches of Leeks Creek, Great Keppel Island,

23°10’S, 150°S5S7’E, Nov 1987, Batianoff 9467 &

Dillewaard (BRI); Port 1, between Curtis and Facing

Islands, Aug 1802, R. Brown “no. 14 spec.” (BRD); South

Percy Island, NW of Onslow Point, 21°45’E, 150°20’E,

Oct 1989, Batianoff 11470 et al. (BRI); Nine Mile Beach,

2—2.5 km NW of Waterpark Point, 22°55’S, 150°47°E,

Jul 1977, Batianoff 485 & McDonald (BRI); Middle

Percy Island, 87 miles SE of Mackay, Apr 1956, Lazarides

5625 (AD, BRI, DNA, MEL, PERTH); Port Curtis (Port

1), between Curtis and Facing Island, Aug 1802, Rk. Brown

s.n. (CANB); “Green Horizons” property of A. Geckeler,

8.4 km from Rules Beach Rd along Fingerfield Rd, c. 60

km direct line NW of Bundaberg, 24°26’S, 151°59°E,

Oct 1992, Geckeler et al. 9 (CANB).

Distribution and habitat: Queensland: Cook,

North Kennedy, Port Curtis, and South Kennedy

Pastoral Disticts. In heaths, woodlands,

shrublands, open forests and vineforest, in sandy

soils; altitude from sea level to c. 120 metres.

Phenology: Flowering January to September;

fruiting throughout the year.

Notes: The leaves and fruits of this species are

sometimes reported to be reddish. A few

specimens have oil glands slightly visible on

the petioles and abaxial leaf surface (Marks &

Mackerras s.n., BRI; Pedley 2730, BRI; Forster

PIF5337, DNA). Possible hybrids with LZ. retusa

include Pedley 2730 and Gray 6126 (QRS). The

black hair-like structures often visible on the

Snow & Guymer, Systematic and cladistic studies of Myrtella and Lithonyrtus (Myrtaceae) 197

leaves and stems represent an ophiostomoid

fungal infection (J. Simpson, pers. comm., the

longer, straight projections in Fig. 71).

Details from Scott (1978) regarding the

type appear to be correct, except for our

replacement of “east coast” with more detailed

locality information (Burbidge, 1955; see also

Burbidge manuscript “An index to the microfilm

of Robert Brown’s botanical descriptions

(manuscript) of Australian plants held at the

British Museum (Natural History) at the British

Museum” [copies at BRI, CANB]).

Conservation status: Common throughout

much of its range.

Local name: “arings” (Brass 18650, BRI).

10. Lithomyrtus repens N.Snow & Guymer

sp. nov. Plantae procumbentes. Folia

anguste ovata usque ovata, basi truncata

vel cordata, non amplexicaulia; pagina in-

ferior foliorum dense hirsuta, glandibus

nullis autem venis secundarils lateralibus

saepe praesentibus. Pedunculi folia

plerumque longiores. Fructus subcylindrici

usque cylindrici, glabrescentes usque sparse

villosi. Type: Australia. Northern Terri-

tory. DARWIN AND GULF REGIon: Arnhem

Land, Upper Catchment of East Alligator

River, 12°47°31.3’S, 133°21°51.8”E, 12

May 1997, N.Snow 7425 et al. (holo:

DNA; iso: BRI, CANB, K, L, MEL, MO,

NSW, US).

Plants suffrutescent, 0.2—0.4 m tall, stems

prostrate. Bark stringy, orangish. Branchlets

villous to tomentose, oil glands lacking. Leaves

opposite, mostly evenly distributed, coriaceous.

Petioles 1.3—1.7 mm long, rounded or

channelled, eglandular. Leaf biades narrowly

ovate to ovate, 14-45 mm long, 7—13(—22) mm

wide, base truncate or cordate, free from or

occasionally clasping the stem, apex obtuse or

acute, margins revolute or flat, upper surface

glabrous, glabrescent or sparsely short villous,

adaxial midvein impressed, lower surface

tomentose, oil glands of lower surface visible,

marginal veins of lower surface prominent, or

invisible or indistinct. Peduncles rigid or

flexible, 10-55 mm long, sparsely villous to

villous. Bracteoles 1.8—3.1 mm long, 0,.2—0.5

mm wide, tips (in flower) not exceeding base

of sepal lobes, villous, persistent or caducous

in fruit. Hypanthium densely villous to

tomentose. Sepal lobes 1—2 mm long, apex

acute, villous to tomentose, persistent in fruit,

mostly reflexed towards body of fruit. Petals

pink to magenta, elliptic to obovate, 5—7 mm

long, upper surface glabrous, lower surface

villous. Stamens 3.5—5 mm long anther

connectives eglandular. Style 4—-5.5 mm long,

mostly straight, glabrous. Fruit subcylindrical

to cylindrical, base tapered, 7.5—-10 mm long,

4—5 mm wide, glabrescent to sparsely villous,

or villous, yellow-green or olive-green. Radicle

tip somewhat spirally contorted above or below

plane of embryo. Fig. 7O-U.

Selected specitinens: Northern Territory. DARWIN AND

GuLF REGION: East Alligator River, 12°40°S, 133°13’E,

Mar 1973, Dunlop 3409 (MEL); Arnhem Land, Upper

East Alligator River catchment, 12°49°40.1"S;

133°21°47.2"E, May 1997, Snow 7427-a (BRI, DNA,

CANB); Headwaters E Alligator River, 12°47°17"S,

133°21’61°E, May 1997, Guymer 2504 et al. (BRI, DNA,

L, MO, NSW); Arnhem Land, Magela Creek upper

catchment, 12°49°19”S, 133°00’25”E, Apr 1995, Cowie

5606 & Brennan (DNA); Kakadu N.P., Upper Koolpin

Creek, 13°27°S, 132°39’E, Jun 1988, Russell-Sinith 5496

& Lucas (BRI, DNA); Kakadu N.P., 8 km NNE of Mt.

Evelyn, 13°32’S, 132°56’°E, Menkhorst341 (DNA, MEL);

13 km SSW of Twin Falls, 13°25.5’S, 132°44’°E, May

1980, Craven 5897 (DNA); Near Mt. Gilruth, 13°10°S,

133°06°E, Mar 1984, Craven & Wightman 8290 (CANB,

MEL); Kakadu N.P., Deaf Adder Creek Gorge, 13°07’S,

132°56’E, Apr 1980, Telford 7994 & Wrigley (CANB);13

km SSW of Twin Falls, 13°25.5’S, 132°44’E, May 1980,

Craven 5897 (CANB); 41 miles NE of Pine Creek, Mar

1971, Dunlop & Byrnes 2109 (DNA); Above U.D.P. Falls,

May 1975, Gittins 2863 (BRI, DNA); Katherine Gorge

N.P., May 1968, Byrnes 870 (BRI, DNA, MEL, PERTH);

Kakadu N.P., c. 25 km NE of Jabiru, 12°30’°24.9"S,

132°57’°0,5.5"E, May 1997, Snow 7435 et al. (BRI,

DNA); Magela Creek, 12°40’S, 133°03’E, Feb 1973,

Dunlop 3377 (DNA); Upper Magela Creek Valley,

Arnhem Land, 12°45’S, 133°05’E, May 1991, Russe/l-

Smith 8470 & Brock (DNA); Kakadu N.P., Mt. Brockman

area, 12°48’S, 132°54’E, Mar 1995, Russel/-Smith 10221

(DNA); Kakadu Nat. Park, Dinner Creek, 13°38°18"S,

132°36’18”E, Apr 1995, Leach 4396 & Greschke (DNA);

Headwaters Katherine River, 13°46°06"S, 133°097°03”E,

Jul 1996, Mangion 218 (DNA); Upper East Alligator,

Arnhem Escarpment, 12°46’°45”S, 133°22’04”E, Jun

1996, Michell 299 & Knox (DNA); Kakadu N.P., 10.5

km NE of Mt. Evelyn, 13°317S, 132°57°E, Apr 1990,

Slee & Craven 2592 (CANB, MEL).

Distribution and habitat: Northern Territory,

Darwin and Gulf Region. Amongst rocks on

sandstone escarpments in open heaths; soils

sandy or skeletal.

198

Phenology: Flowering February to May;

fruiting March to June.

Notes: Lithomyrtus repens can resemble ZL.

hypoleuca in overall morphology (e.g., Slee &

Craven 2592), but its prostrate habit and more

sparsely pubescent and usually longer peduncles

distinguish it from the latter.

Conservation status: The relatively small

geographical range and small size of

subpopulations (Snow, pers. obs.) suggest L.

repens 1s a vulnerable species (criterion D1).

Etymology: The specific epithet 1s from Latin

repens, creeping or prostrate, a reference to the

habit of the species.

11. Lithomyrtus retusa (Endl.) N.Snow &

Guymer comb. nov.

Fenzlia retusa Endl., Ataka Bot. 20, t.

18(1834); Myrtella retusa (Endl.) A. J.

scott, Kew Bull. 33: 300 (1978). Type:

Australia, F’ L. Bauer (ecto: here desig-

nated, W, illustration) (fide also Scott

1978),

Fenzlia phebalioides W. Fitzg., J. & Proc.

Roy. Soc. Western Australia 3: 189

(1918); Myrtella phebalioides (W. Fitzg.)

A.J. Scott, Kew Bull. 33: 301 (1978).

Type: Australia. Western Australia:

Tabletop Mountain, Synnot Range, Aug

1905, WV. Fitzgerald 1350 (holo:

PERTH).

Shrubs or rarely small trees, 1—3(—5) m tall.

Bark smooth or stringy, brown or grey.

Branchlets tomentose, oil glands visible and

prominent. Leaves opposite, concentrated near

branch tips or evenly distributed, coriaceous.

Petioles 2—3 mm long, channelled, glandular.

Leaf blades narrowly elliptic to elliptic, or

obovate (narrowly so), 10-41 mm long, 1-9 mm

wide, base cuneate, free from the stem, apex

obtuse retuse or rarely acute, margins revolute

or flat, upper surface sparsely short- villous to

glabrescent, adaxial midvein impressed, lower

surface densely tomentose, oil glands of lower

surface visible and of differing sizes, marginal

veins of lower surface invisible or indistinct.

Peduncles rigid, 4-7 mm long, sparsely villous

Austrobaileya 5(2): 173-207 (1999)

to tomentose. Bracteoles (0.5—)]—2(—3) mm

long, 0.2—0.5 mm wide, ascending but not

flexuous, the tips (in flower) not exceeding base

of sepal lobes, villous to tomentose, caducous

in fruit. Hypanthium tomentose. Sepal lobes 1—

1.5 mm long, apex acute, glabrous to sparsely

villous, persistent in fruit, mostly reflexed

towards body of fruit. Petals pink, elliptic to

obovate, 3-6 mm long, upper surface glabrous

to glabrescent, lower surface sericeous to

sparsely villous. Stamens 3—4 mm long; anther

connectives glandular with usually 2 or more

oil glands. Style 4-5 mm long, flexuous,

sparsely sericeous. Fruit globose or subglobose,

base rounded, 3-6 mm long, 44.5 mm wide,

glabrous to sparsely villous, yellow-green or

olive-green to dark bluish-black. Radicle tip

somewhat spirally contorted above or below

plane of embryo. Fig. 8A-—G.

Selected specimens: Western Australia. 54.5 km N of

Broome on Pt Columbo track, Dampier Peninsula, SW

Kimberley, 17°36’S, 122°11’E, Jun 1981, Kenneally

7584 (BRI, MEL); Forest Creek, near Drysdale River,

14°39’°S, 126°S57’E, Aug 1975, George 14114 (CANB,

PERTH); King Edward River, c. 50 km NE of Mitchell

River H.S,, Aug 1978, Beauglehole 59127 & Errey 2853

(CANB, PERTH). Northern Territory. DARWIN AND GULF

REGION: c. 6 miles N of Pine Creek Township, 13°48’S,

131°SO°E, Mar 1965, Lazarides & Adams 148 (DNA,

MEL); Arnhem Land, near Murganella Settlement, west

of Workshop Road, 11°32’48.3"S, 132°55°25,3"E, May

1997, Snow 7413 & Mangion (BRI, CANB, DNA, K, L,

MO, NSW); Near Raft Point, 20 km from Mandorah

Road, 12°38°S, 130°34’E, Feb 1991, Cowie 1491 &

Dunlop (BRI, CANB, DNA, MEL, PERTH); Gulf of

Carpenteria, Maria Island, 14°54’S, 135°41’S, Jul 1972,

Dunlop 2946 (BRI, DNA, MEL); Wigram Island,

11°45°S, [36°37°E, Jul 1992, Leach 3058 (BRI, CANB,

DNA); Darwin River, near bridge, 12°42’°S, 130°54’E,

Feb 1977, Parker 705 (CANB, DNA); Headwaters of

Waterfall Creek, 13°24’44°S, 132°29°23”E, Apr 1995,

Leach 4387 & Greshke (CANB, DNA); Katherine Gorge

N.P., 14°14’S, 132°29°E, Jun 1982, Siversten 667 (DNA);

Goyder River, [3°01’S, 134°59’E, Dunlop 8687 & White

(CANB, DNA, MEL); Bathurst Island, 8 km E of Negutu,

11°48°S, 130°30°R, Apr 1987, Wightman 3550 & Smith

(CANB, DNA); Astell Island, N end, 11°51°52”S,

136°24°31"E, Apr 1996, Cowie 6585 (DNA); Groote

Eylandt, headwaters of Amakula River, 14°06’S,

136°33’E, Sep 1991, Cowie 2036 & Brocklehurst

(CANB, DNA, MEL); Black Islet, Sir Edward Pellew

Group, 15°36’S, 136°40’E, Jul 1988, Thomson 2573

(DNA); Melville Island, 11°42’S, 130°50’E, Jan 1992,

Leach 2955 & Cowie (CANB, DNA, MEL); Near

Doyndgji, E Arnhem Land, 12°55’00”S, 135°24’00”E,

Oct 1976, Scarlett 45 (DNA). Queensland. Cook

District: Iron Range Road, c. 2 km upstream [from] creek

crossing Cape Weymouth, 12°46’S, 143°07°E, Apr 1988,

RS eb daca etd dhnucd in iogt pee LL

“APRA SSUES Se se nero bie On eee ce oa

Face al een en en evn ev nn EU eee ee EEE PE EE renter eee ee tn ae :

Snow & Guymer, Systematic and cladistic studies of Myrtel/a and Lithomyrtus (Myrtaceae) 199

Forster PIF4177 & Liddle (BRI, CANB); Agate Creek,

Robinhood Station, SW of Forsayth, 18°50’S, 143°25’E,

Apr 1996, Forster PIF 19080 et al. (BRI); Moa Island,

LO° LVS, 142° E, Budworth 319 (BRI); McDonald

Creek area, 42 km from Mt Surprise township, Mt

Surprise gemfields, 18°03°S, 144°07°E, Apr 1985,

Champion 132 (BRD; 19.8 km from Oroners on the track

to New Dixie, 15°21°S, 143°09’E, Jun 1981, Clarkson

3762 (BRI); c. 24 km SSE of the mouth of the Olive

River, c. 3 km S of Mosquito Point, 12°22’S, 143°10°E,

Apr 1993, Clarkson 9997 & Neldner (BRI); 74 km from

the Chillagoe — ““Wrotham Park” road towards “Bulimba”,

16°58’S, 143°44°E, Jan 1993, Bean 5658 & Forster

(BRI, DNA); 20 km NW of Mount Garnet, on road to

Lappa, 17°36’S, 144°59°E, Jan 1993, Bean 5488 &

Forster (BRI); 5 km N of Fairlight on the Palmerville

road, 15°42°S, 144°03’E, Jun 1992, Clarkson 9610 &

Neldner (BRI).

Distribution and habitat: Western Australia;

Northern Territory, Darwin and Gulf Region;

Queensland, Cook District. In Eucalyptus

woodlands; on sandstone escarpments,

occasionally along streambanks, in sandy and

skeletal soils.

Phenology: Flowering December to June;

fruiting December to September.

Notes: Lithomyrtus retusa has the most

variation in leaf characters of all species of

Lithomyrtus. Plants are covered with oil glands

nearly throughout, including the upper leaf

surfaces where the glands are sometimes dark

and prominent. The foliage hairs can appear

ferruginous on close inspection (Svow et al.

7413; Bean 5065 & Forster, BRI). A few

specimens have the bracteoles exceeding the

base of the sepal lobes in length (Bean 5658 &

Forster, BRI; Bean 5665 & Forster, BRI).

Conservation: Widespread and locally

common.

Acknowledgments

Our thanks to the curators of AD, CANB, DNA,

FI, K, L, MEL, PERTH, and QRS for loans of

and permission to view specimens; the curators

of B, BM, C, E, H, Kk, MO, NY, WRSL for

checking for specimens of Volkens; Dr Greg

Leach for assistance in planning a field trip to

the Northern Territory; Chris Mangion for

considerable assistance during fieldwork; Dr

Don Foreman when Australian Botanical

Liaison Officer, Kew, England, for information

= eu te

=» ‘i

* eg

a Ok oo * = =u *

- -” 7 = .

me ee ie ee te Pp guees terl

al = r ae Pl

Are D ead eeT Pe

re el ee ee

7 Ct a 7 *. .*

+ + . a* +

ad - ae Ft * ae

= 5

Sis

o oky *

ail

Figure 8. A-G: Lithomyrtus retusa, A. flowering

branchlet x 3, B. section of branchlet x 2. C, adaxial leaf

profile x 2. D, adaxial leaf apex x 4. E. flower x 6. F.

bracteole x 24. G. fruit x 4. A-G: Snow 7413 & Mangion

(BRI).

200

on type material in London; Drs Penny Gullan

and John Lawrence for identification of msects;

Will Smith for illustrations; Peter Bostock for

the Latin descriptions; the Northern Land

Council for permission to collect in the Wauk

(Murganella) region; and Philip Sharpe for

several translations from German. This

research was supported by grants from the

Australian Biological Resources Study

during 1997-1999,

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Appendix I. Characters for natural language

descriptions for Lithomyrtus and Myrtella as coded

in DELTA (Dallwitz et al., 1993). Some characters

invariant for this study are listed, given their likely

value in later studies of Myrtaceae.

*Character List

#1. including <synonyms: genera’ included

in the future description>/

#2. plants <growth form>/

1. suffrutescent/

2. shrubby/

3, tree-like shrubs/

4. small trees/

#3. plants <main> stems <habit>/

1. erect/

2. prostrate/

#4, plant height/

m tall/

#5, bark <texture>/

1. smooth/

2. stringy/

#6. bark <coloutr>/

1. brown/

2. grey/

3. orangish/

#7. branchlets <whether four-angled>/

i. distinctly four-angled/

2. rounded/

#8. branchlets <whether winged>/

1. with distinctly winged edges/

2. not winged/

Austrobaileya 5(2): 173-207 (1999)

#9. branchlets <vestiture >/

1. glabrous/

2. Sparsely sericeous/

3. villous/

4. tomentose/

#10. branchlets oif glands <whether visible and prominent

or not visibie>/

1. visible and prominent/

2. absent/

#11. leaves <arrangement between nodes>/

1. distichous/

2. decussate/

#12, leaves <arrangement at nodes>/

1. opposite/

2. disjunct opposite/

3. whorled/

4, alternate/

#13. leaves <distribution: whether concentrated near

branch tips or more or less evenly distributed>/

1. mostiy concentrated near branch tips/

2. mostly evenly distributed/

#14. apex acuminate, leaves <texture: whether coriaceous

or soft>/

|. coriaceous/

2. soft/

#15. stipules <type>/

1. consisting of 2—severai short, thick,

deep red, often laterally

fused scaie-like structures/

2. consisting of 2—severa! free

ferrugineous setose hairs/

#16, petioles <length>/

mm long/ —

#17, petioles <whether channelled>/

1. rounded/

2, channeiled/

#18. petioles <glandularity>/

1. eglandular/

2. glandular/

#19. leaf blades <shape>/

. Hinear/

. narrowly elliptic/

. elliptic/

. lanceolate/

. narrowly ovate/

. ovate/

. broadly ovate/

. obovate/

CoO™! ON Un fe Ge hoe

#20. leaf blades <length>/

mm fong/

#21. leaf blades <width>/

mm wide/

BME EITS SA eS en

Snow & Guymer, Systematic and cladistic studies of Myrtella and Lithomyrtus (Myrtaceae)

#22. leaf blades at base <shape>/

1. truncate/

2. cordate/

3. cuneate/

#23. leaf blades at base <whether clasping stem>/

1. free from the stem/

2. Clasping the stem/

#24. leaf blades at apex <shape>/

L. obtuse/

2. acute/

3, retuse/

#25. leaf blade margins <whether folded>/

1. revolute/

2. flat/

#26. leaf blades adaxial surface <indumentum>/

. glabrous/

. glabrescent/

. sparsely sericeous/

. Sericeous on lower portion of midvein/

. sparsely tomentose along lower edges/

. Sparsely short villous/

. sparsely tomentose/

. tomentose/

CO~] oO On & Whe

#27, <leaf blades> adaxial midvein/

i. impressed/

2. flush/

#28. leaf blades lower surface <indumentum>/

. glabrous/

. glabrescent/

. Somewhat sericeous/

, sericeous on lower portion of midvein/

. villous/

. densely villous/

. tomentose/

. densely tomentose/

OO —] ON On . ta bBo ee

#29. leaf blade oil glands of lower surface/

1. absent/

2. visible and about equal in size/

3, visible and of differing sizes/

#30. leaf blade marginal veins of lower surface/

1. prominent/

2. absent or indistinct/

#31. inflorescence <type>/

l. a solitary flower/

2. other/

#32. peduncle <arrangement>/

|. solitary in leaf axils/

2. other/

#33. peduncle <stifiness>/

1. rigid/

2, flexible/

#34. peduncle (in flower) <length>/

mm long/

#35. peduncle <pubescence in flower>/

. glabrous/

, sparsely antrorsely sericeous/

. sparsely villous/

. Villous/

. tomentose/

1 ff Go bo ke

#36. bracteoles subtending flowers <number>/

1. two/

2. other/

#37. bracteoles <appearance>/

1. foliaceous and usually with midrib/

2, scale-like and lacking midrib/

#38. bracteoles <length>/

mm long/

#39. bracteoles <width>/

mm wide/

#40, bracteoles <flexuosity>/

1. ascending and rigid/

2. trregularly flexuose/

—

. bracteole tips <length> Cin flower)/

1. exceeding base of sepal lobes/

2. shorter than base of sepal lobes/

#42. bracteoles <indumentum>/

. glabrous/

. sparsely sericeous/

. villous/

. densely villous/

, tomentose/

Nin be WNM re >

#43, bracteoles <persistence>/

1. persistent in fruit/

2. caducous in fruit/

#44, hypanthium <shape in flower>/

1. obconic/

2. other/

#45. hypanthium <indumentum>/

1. glabrous/

2. densely sericeous/

3, vilious/

4. densely villous/

5. tomentose/

6, densely tomentose/

#46.sepals <number>/

1. five/

2. four/

#47.sepals <fusion at base>/

1. fused proximally/

2. free/

#48. sepal lobes <fusion>/

203

. minutely and sparsely sericeous along edges/

204

1. coherent in mature bud/

2. separate in mature bud/

#49.sepal lobes apex <shape>/

1, acute/

2. acuminate/

#50. sepal lobes <length>/

mm long/

#51.sepal lobes <indumentum>/

. glabrous/

. Sparsely sericeous/

, sparsely villous/

. villous/

. sparsely short tomentose along edges/

. sparsely tomentose/

. tomentose/

~J oN On fe GQ hw

#52.sepal lobes <duration>/

1, persistent in fruit/

2. deciduous in fruit/

#53.sepal lobes <habit in fruit>/

1, mostly reflexed towards body of fruit/

2. mostly ascending and above body of fruit/

#54. petals <number>/

1. ftve/

2. four/

#55. petals <color>/

1. white/

2, pink/

3. magenta/

#56. petals <shape>/

l.elliptic/

2. ovate/

3. broadly ovate/

4, obovate/

#57. petals <iength>/

mm long/

#58. petals — adaxial surface <indumentum>/

lL. glabrous/

2. glabrescent/

3. sparsely sericeous/

4, villous/

#59. petals — abaxial surface <indumentum>/

. glabrous/

. sericeous/

. sparsely villous/

. Vilious/

. tomentose/

. minutely tomentose on edges/

Ov tn 4m Go BO Re

#60.stamens <number>/

1. numerous/

2. five/

#61. stamens <number of whorls>/

#62,

#63,

#64,

#65,

#66.

#67,

#68,

#69,

#70.

fed

#72.

#73.

#74,

#75,

#76.

#77.

Austrobaileya 5(2): 173-207 (1999)

i. uniseriate/

2. multiseriate/

stamens <length relative to petals>/

1. included/

2. exserted/

stamens <position in bud>/

|. folded centripetally in bud/

2. other/

stamens <length>/

mm long/

staminal disk <indumentum>/

1. glabrous/

2. hairy/

anthers <shape>/

1. rounded/

2. sagittate/

anthers <attachment to filaments>/

1. dorsifixed/

2. basifixed/

anthers <length reijative to filaments>/

1. less than one fourth length of

filaments/

2. other/

anthers <dehiscence>/

1. dehiscing via longitudinal slits/

2. other/

anthers connectives <glandularity>/

{. eglandular/

2. glandular/

styles <length>/

mun long/

styles <habit>/

1. flexuous/

2. mostly straight/

styles <indumentum>/

1. glabrous/

2. sparsely sericeous/

stigma apex <type>/

1. terete/

2. capitate/

fruit <dehiscence>/

1. indehiscent/

2. dehiscent/

fruit <type>/

1. a hard berry with bony, more or less

fused seeds/

2. a soft berry/

fruit <shape>/

Oe Sr nn ee ee ee ee ee ee eee te tee eee eeenn eee ee BOE Bg ee

SOMES HARM one nmin te alae db oe ers em torte oan mien

nn tree Merete need AA it ADIT

EE TEE SS ST er

Snow & Guymer, Systematic and cladistic studies of Myrfella and Lithomyrtus (Myrtaceae)

, subglobose/

. globose/

, subcylindrical/

. eylindrical/

. fusiform/

Lh - Go Bho

#78. fruit base <shape>/

1. rounded/

2. tapered/

#79. fruit <length>/

mm long/

#80. fruit <width>/

mm wide/

#81, fruit <pubescence>/

. glabrous/

, glabrescent/

. sparsely sericeous/

. sparsely villous/

. Villous/

. sparsely tomentose/

, tomentose/

, densely tomentose/

Com) MH OA & GW hoe

#82. fruit <color>/

1, dark bluish-black/

2. yellow-green or olive-green/

3, whitish by virtue of dense hairs/

4, red/

5. brown/

#83. peduncle <length>/

mm long in fruit/

#84. ovaries <nmumber in frutt>/

in fruit/

#85. placentation/

1. parietal/

2. axtle/

#86. locules/

per ovary/

#87,seed coat <texture>/

1. hard, bony/

2. soft, somewhat pulpy/

#88. adjacent seeds <relative fusion>/

1. slightly fused, the boundaries in

longitudinal section generally

distinct and perpendicular to

long axis/

2. tightly fused, the boundaries in

longitudinal section generally

indistinct and often somewhat oblique

to long axis/

#89, embryo <shape>/

|, falcate to crescent/

2. circinate/

#90. embryos <oiliness>/

l. oily/

2. starchy/

#91. hypocotyl <length>/

!. relatively short/

2. relatively long/

#92. hypocotyl <thickness>/

{, relatively thick/

2. relatively thin/

#93, hypocoty! <swelling>/

{. swollen near tip/

2. not or slightly swollen near tip/

#94, hypocoty! tip <position reiative to plane

of embryo>/

1. somewhat spirally contorted above or

below plane of embryo/

2. held at same horizontal plane as

embryo/

#95, cotyledons <curvature>/

1, circinate/

2. mostly stratght/

#96.cotyledons <length>/

1. relatively short/

2. relatively long/

#97, cotyledons <length relative to

hypocoty|>/

1. shorter than hypocotyl/

2. about equalling hypocotyl/

#98, cotyledons <thickness>/

1. relatively thick/

2. relatively thin/

#99. cotyledons <position>/

1. not folded back towards hypocotyl/

2. folded back towards hypocoty!/

#100. <geographical range>/

1. Queensland/

2. Northern Territory/

3. Western Australia/

4, Papua New Guinea/

5. Irian Jaya/

6, Guam/

#101. <geographical range in Queensland>/

L. Cook/

2, Burke/

3. North Kennedy/

4, South Kennedy/

5. Mitchell/

6, Gregory North/

7. Port Curtis/

#102. habitat/

#103. elevation/

#104. <taxon name>/

205

206

Appendix 2. Enumeration and discussion of characters

used in cladistic studies. Numbers in parenthesis

following character description correspond to DELTA

numbering system (Appendix 1),

1, Stem habit (3): prostrate (0); erect (1). Z. repens is

prostrate and herbarium specimens indicate a prostrate

habit for L. linartifolia.

2. Young branch cross section (7): quadrangular (0); round

(1). The character is most easily observable in young

twigs.

3. Young branch wings (8): present (0); absent(1). This

character is also most easily observable in young

twigs.

4, Oil glands on branchlets (10): absent (0); present (1).

Since the presence of oil glands on branchlets does

not correlate invariably with the presence of oil glands

abaxially on leaves, their expression must have a

different genetic basis.

5. Leaf distribution (13): concentrated near branch tips

(0); more or less evenly distributed (1). This character

is not always easily determined on herbarium

specimens. It is pronounced for L. densifolia, which

has very dense foliage.

6. Leaf texture (14): coriaceous (0); soft (1). This character

partially reflects patterns of pubescence on the leaves,

and may need modification.

7, Mature leaf bases (23): free from the stem (0); clasping

the stem (1). Stem-clasping leaves occur in JL.

kakaduensis and are occasionally present inZ. repens.

Youngest leaves of L. grandifolia can be stem-

clasping.

8, Stipules (15): short squamose structures that are often

fused laterally (0); free short setose hairs (1). The

differences in character states are readily apparent on

herbarium specimens. The stipules of Lithoniyrtius

resemble those in Fig. 3b of Dahigren & Thorne

(1985); those of Afsrtella somewhat resemble the

stipules in Fig. 3f (op. cit.), but are shorter, broader

and fused at the base.

9, Petiolar oi! glands (18): absent (0); present (1). Since

petiolar oil gland presence Is not correlated with glands

elsewhere, it is assumed to have a distinct genetic

basis.

10. Leaf blade base shape (21): truncate (0); cordate (1);

cuneate (2).

1]. Abaxial leaf oil glands (29): indistinct (0); visible (1).

12. Marginal adaxial leaf veins (30): absent or indistinct

(0); prominent (1)

13. Bracteole type (37): foliaceous with a distinct midrib

(0); seale-like and lacking a midrib (1).

Austrobaileya 5(2): 173-207 (1999)

14. Bracteole flexuosity (40): ascending and straight (0);

flexuose (1).

15, Bracteole length (41): longer than base of sepal lobes

(0); shorter than base of sepal lobes (1).

16. Sepal lobe arrangement In bud (48): coherent in mature

bud (0); separate in mature bud (1)

17. Sepa! lobe apex (49): acute (0); acuminate (1).

18. Sepal lobe position in fruit (53): mostly reflexed against

body of fruit (0); mostly above body of fruit (1).

19, Staminal disk vestiture (65): glabrous (0); hairy (1).

The staminal disk is the apex of the hypanthium on

which stamens are inserted.

20, Anther connectives (70): eglandular (0); glandular (1).

Lithomyrtus retusa usually has two or more oil glands

per anther, which can arise from the connective or

anther wall. No quantification of glands was attempted

(e.g., Landrum & Boniila 1996).

jem,

. Fruit base shape (78): rounded (0); tapered (1).

22. Relative fusion of seeds (88): slightly fused, the

boundaries in longitudinal section distinct and

perpendicular to long axis of fruit (0); tightly fused,

the boundaries in |.s. indistinct and often oblique to

long axis of fruit (1).

23. Embryo shape (89): slightly curved (0); circinate (1).

24. Hypocoty! length (91): relatively short (0); relatively

long (1). The alternative states are very apparent

between Lithomyrtus and Myrtel/a, although they may

be of limited use in Myrtaceae as a whole (see

Landrum & Stevenson 1986).

25. Cotyledon length (96): relatively short (0); relatively

long (1). The comments for character 24 apply equally

here.

26. Cotyledon thickness (92): relattvely thick (0); relatively

thin (1). The comments for character 24 apply equally

here.

27. Hypocotyl tip (94): somewhat spirally contorted above

or below plane of embryo (0); held in same plane as

embryo (1).

207

snow & Guymer, Systematic and cladistic studies of Myrtella and Lithomyrtus (Myrtaceae)

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LITLE ERATE Ell lg pd eee es pq ee epee eg tee rte ttt eee at 2

Desmodium Desvy. ( Fabaceae) and related genera in Australia:

a taxonomic revision

Les Pedley

Summary

Pediey, Les (1999). Desmodium Desv. (Fabaceae) and related genera in Australia: a taxonomic revision.

Austrobaileya.5(2):209-261, The concepts of Ohashi (1973) are adopted and five genera are recognised

for Australia: Aphyllodium (DC.) Gagnep. (Dicerma DC. sensu Benth.), Dendrolobium Benth.,

Desmodium Desv., Phyllodium Desy. and Tadehagi Ohashi. A classification of Desmodium in Australia,

closely following Ohashi’s for Asia, is presented, Taxa described as new are: Desmodium subg.

Acanthocladum (type: D. acanthocladum F. Muell.), Desmodium ser. Arillata (type: D. microphyllum

(Thunb.) DC.), Desmodium ser Sagotia (type: D. triflorum (L.) DC.), Aphyllodium glossocarpum, A.

latifolium, A. parvifolium, A. schindleri, A. stylosanthoides, Dendrolobium multiflorum, Desmodium

glareosum, D. pullenii, D. pycnotrichum, D. tiwiense, Phyllodium pulchellum vay. glabrius, P. hackeri

and Tadehagi robustum. New combinations are Desmodium sect. Desmodiopsis (based on Alysicarpus

sect. Desmodiopsis Schindl.) Desmodium ser. Stenostachys (based on Desmodium sect. Stenostachys

Schindl., lectotype D. filiforme Zoll. & Moritzi), Dendrolobium cheelii (based on Alysicarpus cheelti

C.A. Gardner), Dendrolobium polyneurum (based on Desmodium polyneurum S.T.Blake),

Dendrolobium umbellatum var. hirsutum (based on Desmodium umbellatum var. hirsutimn DC.) and

Desmodium whitfordii based on Desmodium nemorosum subvar. whitfordii Schindl., the last two

from the Mascarenes and the Phillipines respectively. Keys to species are provided; new and poorly

understood taxa are described and references to descriptions of all others provided; brief notes on

distribution and habitat of all species are included; and typification of some species is discussed.

Forty-eight Australian species are treated: six Apiyllodium, six Dendrolobium, 32 Desmodium, three

Phyllodium and one Tadehagi.

Key words: Aphyllodium, Dendrolobium, Desmodium, Dicerma, Phyllodium, Tadehagi

Les Pedley c/- Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,

Toowong, Queensland 4066, Australia.

Introduction

This account relies heavily on the work of

Ohashi (1973), and may be considered a

supplement to it. I have adopted Ohashi’s

circumscription of genera and, in most cases,

of species. Where his descriptions of taxa are

satisfactory I have merely referred to them.

For species widely spread in the Old World

or recently introduced into Australia I have

referred to other accounts, especially those

of Schubert (1971) and Verdcourt (1979).

Ohashi (1973) discussed at some length

the various treatments of Desmodium sensu

lato. Candolle (1825) recognised a large genus

Desmodium Desv. and much smaller

Nicolsonia DC. and Dicerma DC. (which

included Phyllodium Desy.). Dicerma 1s an

illegitimate name and has to be replaced by

Aphyllodium (DC.) Gagnep.; this matter 1s

Accepted for publication 25 August 1998

discussed below (p.211 ). Bentham (1852), taking

a rather natrow view of genera, described several

as new and recognised Candolle’s two sections

of Dicerma as distinct genera, namely

Phyllodium Desv. and Dicerma DC, (for sect.

Aphyllodium). Later, however, Bentham

(1865a) tergiversated; he wrote “In the first

general account of the group which I made for

the ‘Plantae Junghunianae’ it appeared to me

that ifthe universally accepted genera Dicerma

and Nicolsonia of De Candolle are maintained,

it would be necessary to keep up several others,

and even to establish new ones, for which I gave

the characters in that work. I have however,

since then, had occasion to examine in detail

above a hundred species, ... and the characters

relied on for separation of the smaller genera

have proved too uncertain or artificial to be

available for any but sectional divisions; that

in our ‘Genera Planarum’ I have found it

necessary not only to restore the genus to the

210

extent originally contemplated by De Candolle,

but to add to it his Dicerma and Nicolsonia...”.

He further stated that it was with much

hesitation that he refrained from adding to them

the small or monotypic genera Ougeinia

Benth., Adecopus Benn. and Pseudarthria

Wight & Arn. So it was that Bentham (1864,

1865b) and Baker (1879) adopted a broad

concept of Desmodium. Schindler had different

concepts and recognised not only the segregate

genera proposed by Candolle (1825) and

Bentham (1852) but described several new

ones; see Schindler (1928) for details of his

numerous papers. Schindler’s views were not

genetally accepted; Fosberg (1966), for

example, was forthright in his opposition,

noting of Desmodium that ‘some authors have

dismembered this very natural and distinctive

genus into a number of ill-distinguished

segregate genera. Nothing seems to be gained

by accepting these, so they are ignored here’.

Knaap-van Meeuwen (1962) and Verdcourt

(1979), both of whom dealt with floras of

regions where segregate genera occur, also

‘kept to the more extended view of

Desmodium’ (Verdcourt 1979). Ohashi

(1973) adopted a position between that of

Bentham and his followers and Schindler. On

the whole Ohashi et al. adopted Ohashi’s

(1973) treatment, though they discussed the

difficulties in arriving at a ‘sensible

classification’ of Desmodieae.

Adopting a ‘more extended view of

Desmodium’ has some attraction. Having dealt

with Australian and Sri Lankan species which

has necessitated a more-than-cursory

examination of most of the Old World

Desmoditinae I would favour recognition of four

genera in Australia, namely Tadehagi,

Desmodium, Phyllodium, Aphyllodium, with

Dendrolobium included in Phyllodium.,

However, until further scientific

investigation, rather than intuition, informed

as 1t might be, clarifies the situation I am

following the classification proposed by

Ohashi (1973). Though recorded by Ohashi,

Codariocalyx does not occur in Australia.

Austrobaileya 5(2): 209-261 (1999)

Aphyllodium (DC.) Gagnep., Not. Syst. 3:254

(1916); Pedley, Rev. Handb. FI.

Ceylon10:106(1996); Ohashi, Taiwania

42: 142(1997); Dicerma DC., Prodr.

2:339 (1825), Mém. Leg. 326 (1826),

incl. sect. Aphyllodium DC. but excl. sect.

Phyllodium (Desv.) DC.; Wight & Arn.,

Prodr. FIL. Ind. orient. 238 (1834); Benth.

in Migq., Pl. Jungh. 217 & 219 (1852);

Schindl., Rep. sp. nov. reg. veg. 20:267

(1924); Hutch., Gen, FI. Pl. 1:483 (1964);

Ohashi, Ginkgoana 1:251 (1973), nom.

ileg; Desmodium sect. Dicerma Benth.

in Benth. & J.D. Hook., Gen. Pl. 1:519

(1865); Desmodium subg. Dicerma

Baker in J.D. Hook., FI. Brit. India 2:163

(1876); Type: Aphylodium biarticulatum

(L.) Gagnep.

Aphyllodium is the most distinctive and well

defined of the segregates of the Linnean

Hedysarum though Bentham (1865b),

followed by later authors, referred it (as

Dicerma) to Desmodium. It has united

amplexicaul stipules that are mostly divided

to the middle, secondary bracts similar in size

and shape to the primary bracts, and deeply

constricted I- or 2- jointed pods with

orbicular to transversely elliptic (rarely, in

A, glossocarpum, oblong) articles.

Ohashi (1973) considered Aphyllodium

(as Dicerma) to be related to Phyllodium

because of the similarity of their

inflorescences. The primary bracts are,

however, not homologous with the foliar

bracts of Phyllodium (q.v.) and the genera

are not particularly closely related.

Hitherto Aphyllodium has been

considered to consist of either three species,

one widespread and polymorphic (Schindler

1924, Ohashi 1997b), or one species (Ohashi

1973). This study indicates that there are seven

species in Australia and New Guinea , one (A.

biarticulatum) wide ranging and one (A.

novoguineense (Schindl.)Ohashi) poorly

known and probably misunderstood. The

misapplication of the name is discussed briefly

below.

SEGRE SS oe

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 211

Nomenclature

The genus Phyllodium was described by

Desvaux who included in it two species, P

pulchellum, based on Hedysarum pulchellum

L., and P /utescens, based on H. lutescens Pott.

His description and illustration were derived

solely from P pulchellum, and P. lutescens has

since been transferred to the monotypic genus

Pycnospora,

Candolle described the genus Dicerma,

with two sections: Aphyllodium, with one

species Dicerma biarticulatum (Hedysarum

biarticulatum L.), and Phyllodium, Desvaux’s

genus Phyllodium. He explained (1826) that

he wished to retain Phy/lodium in its original

sense and had therefore, in adding another

section, coined another generic name. Under

the current Code of Botanical Nomenclature,

Candolle should have used the existing generic

name Phyllodium for his enlarged genus and

because he did not do so, his generic name

Dicerma was superfluous and therefore

illegitimate.

Bentham (1852) treated the two sections

of Dicerma DC. as genera for which he used

the names Phyllodium Desv. and, for sect.

Aphyllodium, the illegitimate name Dicerma.

In the next 60 or so years the name Dicerma

was used at generic, subgeneric and sectional

rank. Except for its use as a generic name these

usages do not conflict with the International

Code of Botanical Nomenclature.

Gagnepain, without any discussion at all

of the standing of the name Dicerma, raised

Dicerma sect. Aphyllodium to generic rank as

Aphyllodium (DC.) Gagnep. and in doing so

provided a legitimate name for Dicerma as

circumscribed by Bentham (1852).

Unfortunately Schindler, who sometimes

adopted names which are at variance with the

present International Code of Botanica!

Nomenclature, quibbled about Gagnepain’s

generic distinctions, but agreed using other

criteria that Aphyllodium (DC.) Gagnep. was

indeed distinct and again applied to it the

illegitimate name Dicerma. Such has been the

prestige of Bentham and Schindler (rightly so

as taxonomists) that Aphyllodium has not been

taken up as a generic name. Conservation of

the name Dicerma tor Dicerma sect.

Aphyllodium is not warranted as it has never

been applied to more than three species, two

of them little known with narrow geographic

ranges. I have therefore used the correct name

Aphyllodium for the Australian species.

A number of infra-specific taxa of 4.

biarticulatum. has been described and a

knowledge of the circumscription of these taxa

and the rather loose application of their names

is essential to an understanding of the taxonomy

of the genus in Australia. These matters are

dealt with in some detail under A. biarticulatum.

Key to species

1. Terminal leaflet more than 12 mm wide, more than 30 mm long and 2.2— __

3.2 MUMS LONG! ANAL WAGE. Sic tess aos costae tan dave w Sige ic beabiestas oe akatlat samen 1. A. latifolium

Terminal leaflet less than 12 mm wide, if more than 30 mm long then either

more than 3 times longer than wide or less than 1.5 times longer than

WAG Creo Sno dbx Wren dated eke utara tee att oh chee es ib orld eta lpn hoe 6 ti apse Mee Ree 2

2. Leaflets to 4mm long, 1—2 times longer than wide.................. 2. A. parvifolium

Leaflets more than 10 mm long, more than twice as long as wide ........ 0.00.0 eee 3

3. Bracteoles 5—9 mm long, more than 1.5 times longer than calyx ........ 3. A. schindleri

Bracteoles to 5 mm long, only slightly longer than calyx ....... 0.0.00. 0c ec ee ee eee 4

4. Distal (or single) article of pod 7-9 mm long .......... 0.0000 eae 4. A, glossocarpum

Articles of pod less than 5.5 mm long

ss &§ &©§ *€* *§ #&# 8 FEF F&F BB *# ££ 8 @ BF © B&B #£ BF #F ££ FF 8B 8 BF 8 BH HF FF FF FEF © 2 FF FF SF FE FF F F F

. Leaflets + acute; stipules 10-20 mm long; petioles 10-16 mm long . 5. A. stylosanthoides

Leaflets obtuse; stipules 6-15 mm long; petioles 3-11 mmlong ..... 6. A. biarticulatum

212

Il. Aphyllodium latifolium Pedley, sp. nov.

habitu simile A. schindleri Pedley sp.

nov. autem foliolis latioribus minus

elongatis, stipulis brevioribus,

inflorescentiits brevioribus bracteis

brevioribus praeditis, ramulorum pilis

patentibus differt. Typus: Queensland.

Cook District: Dead Horse Creek, 8 km

S of Morehead River, 15°04’S 143°43’E,

May 1987, JR. Clarkson 7193 & B.K.

Simon (holo: BRI; iso: BRI, K, L, MBA,

NSW, PERTH, QRS).

Aphyllodium sp. (Edward River J.R.

Clarkson 3544) Pedley in Henderson

(ed.): Queensland plants: names and

distribution: 76 (1997),

Erect shrub to 1.5 m tall, branching from the

base; branchlets with indumentum of dense

spreading fulvous hairs to | mm long; stipules

glabrous, 6-10 mm long. Leaflets oblong,

elliptic or obovate, rounded at the base, obtuse,

minutely apiculate at the apex, margins slightly

recutved, appressed pubescent on upper

surface, prominently reticulate and appressed

pubescent on lower surface; terminal ones 30—

50 mm long, 12—16 mm wide, 2—3 times as long

as wide, the laterals somewhat smaller; stipels,

often hidden in indumentum, c. 0.6 mm long;

petioles 10—20 mm long. Inflorescences to 18

cm long; primary and secondary bracts c. 5 mm

long; pedicels 1.4—-2.4 mm long. Flowers: calyx

with indumentum of uncinate hairs c. 0.5 mm

long, 3-4 mm long, the tube cylindrical 1.8—

2.2 mm long, lobes 1.2—1.8 mm long;

bracteoles as long as tube; corolla mauve or

purple; standard obovate or orbicular with short

claw, 6—7.3 mm long, 3.4—5 mm wide; wings

rectangular, auriculate, 4.2-5 mm long, 1—1.7

mm wide on claw 1.2-2 mm long; keel about

as long as wings; ovary densely pubescent, style

slender, stigma minute, surrounded by hairs.

Pods of two articles, densely appressed hairy,

each 4—5 mm long, 3.3—4 mm wide; seeds c. 2

mm long, 1.5 mm wide.

Selected specimens: Queensland. Cook DISTRICT:

Edward River Aboriginal Reserve, 15°10’S 141°40’E,

Oct 1980, Clarkson 3544 (BRI, MBA); Edward River,

Apr 1980, Garnett 10 (BRI); c. 80 km NW of Laura,

15°12’S 143°53’E, Jul 1990, Pedley 5520 (BRI); east

bank of Little Laura River, S of Laura, |15°32’S

144°20°E], May 1975, Byrnes 3419 (BRI); 33 miles [53

Austrobaileya 5(2): 209-261 (1999)

km] NE of ‘Dunbar’ H.S., 87 miles [139 km] ESE of

Mitchell River mouth [15°45’S 142°46’E}, Aug 1966,

Story 8029 (CANB).

Distribution and habitat: Aphyllodium

latifolium occurs on sandy soils on roadsides

and in eucalypt woodland tn the southern part

of Cape York Peninsula. Map 1.

Affinities: The species resembles A. schindleri,

with which it is sympatric, in habit and general

appearance but differs 1n its wider, less elongate

leaflets, shorter stipules, shorter inflorescences

with shorter bracts, and the spreading hairs of

the branchlets.

Etymology: The specific epithet is a compound

of Latin, /atus, wide, and foliun7, leaf: a

reference to the wide leaflets of the species.

2. Aphyllodium parvifolium Pedley, sp. nov.

ab omnibus aliis speciebus Aphyllodii

foliis parvioribus usque 4 mm longis, 1—

1.2 plo longioribus quam latis et floribus

magnis vexillo 7.5—8.5 mm longo ornatis

distinguenda. Typus: Western Australia.

Near Barred Creek, 33 km N of Broome,

17°40’°S 122°12’E, 3 April 1988, KF

Kenneally 10612 (holo: PERTH; iso:

CANB, DNA).

Prostrate subshrub; branchlets with

indumentum of dense, spreading to ascending

hairs c. 1 mm long; stipules 4-5.5 mm long

with long marginal hairs. Leaflets oblong or

orbicular, obtuse at base and apex, moderately

dense, long, ascending hairs on both surfaces,

terminal and lateral leaflets about equal in size,

3.2—7.5 mm long, 3.2—4 min wide, c. 1-2 times

longer than wide; stipels minute; petioles 2.5—

4.5 mm long. Inflorescences to c. 6 cm long;

primary and secondary bracts 2-3 mm long;

pedicels 1.5—2 mm long. Flower: calyx with

sparse indumentum of short uncinate hairs and

longer straight marginal ones, 44.5 mm long,

the tube c, 2.6 mm long, the lobes 1.3—1.9 mm

long; corolla pink; standard obovate with short

claw, 7.5-8.5 mm long, c. 4 mm wide; wings

rectangular, auriculate, c. 5—5.5 mm long, c.

1.2 mm wide on claw 2—2.5 mm long; keel

slightly longer than wings; ovary densely

pubescent; style flattened towards the tip,

stigma minute, fringed with hairs. Pods of 2

articles, appressed hairy on veins, each c. 6 mm

long and 4 mm wide; seeds not seen.

ESSE EEA as ne gn EL BB

eee

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 213

Other Speciinen: Western Australia. McLarty Huls O14

Camp [c. 19°30’S 123°30°E] JS. Beard (PERTH).

Distribution and habitat: The species is known

only from the collections cited. Map 2.

Affinities: Aphyllodium parvifolium 1s

probably most closely related to A.

biarticulatum but is easily distinguished by its

small leaflets less than 1.5 times longer than

wide and its large flowers.

Etymology: The epithet is from Latin parvus,

small and fo/ium, leaf, a reference to the small

leaves of the species.

3. Aphyllodium schindleri Pedley, sp. nov. ab

allis speciebus Aphyllodii bracteolis

calyce 1.5—3 plo longioribus, 5-9 mm

longis, ab A. /atifolio Pedley foliolis

angustioribus, ab A. glossocarpo Pedley

lomentis brevioribus, speciebus

Australiensibus ceteris inflorescentis

densioribus longioribusque distinguenda.

Typus: Queensland. Cook District: 8 km

SW of Beagle North Camp, 13°04’S

141°45’E, June 1982,/7.R2. Clarkson 4479

(holo: BRI; iso: BRI, DNA, K, NSW,

PERTH, QRS).

Dicerma biarticulatum {. longibracteatum

ochindl., Rep. sp. nov. reg. veg. 20:268

(1924). Type: Carpentaria. Mainland

opposite Groote Eylandt, Brown ‘4183’

(isosyn: BM, BRI, K).

Aphyllodium novoguineense auct. non

(Schindl.) Ohashi; Ohashi, Taiwania 42:

146(1997) p. p.

Aphyllodium sp. (Lockerbie L. J. Brass

18464) Pedley in Henderson (ed.):

Queensland plants: names and

distribution: 76 (1997).

Shrub to 1.5 m tall; branchlets glabrous or with

sparse indumentum of appressed hairs to 2 mm

long, often ascending hairs at nodes, or (in

Western Australia and Papua New Guinea) of

sparse to moderately dense ascending hairs;

stipules (10—)13-—25 mm long, glabrous, often

with a few marginal hairs towards the tips.

Leaflets lanceolate, narrowly elliptic, narrowly

oblong or oblanceolate, attenuate at the base,

acute or occasionally obtuse, mucronulate at

the apex, upper surface glabrous or (especially

in northern Cape York Peninsula and Papua

New Guinea) with sparse appressed hairs,

lower surface with sparse appressed hairs to 1

mm long, densest on margins and midrib, often

with sparse erect minute (0.05 mm long)

glandular hairs as well; terminal leaflets (1 5—)25—

60(—70) mm long, 4.5—11(-22) mm wide, (3—-)4—

7(-9) times longer than wide, the laterals

slightly smaller; stipels glabrous to 2.4 mm

long, often less and sometimes hidden in

indumentum; petioles 6—-18(-—24 in New

Guinea) mm long; pulvinuses 1—2(—2.7) mm

long, laterals slightly smaller than terminal.

Inflorescences dense, 6-16 mm long; striate

primary bracts (6—)8—12(—16) mm long,

secondary bracts 6—-14(—-16) mm long

(specimens from New Guinea at upper ends of

ranges), both with hyaline marginal hairs;

pedicels 0.7—2.6 mm long. Flower: calyx with

sparse indumentum of hooked hairs c. 0.05 mm

long, 2.6-3.7 mm long, tube 1.5—2.3 mm, lobes

1.1—1.5 mm long; bracteoles similar in shape

and texture to bracts, shorter or longer than the

flower, 5—9,3 mm long; corolla white to mauve;

standard obovate, 5.6—9.3 mm long, (2.5—)3—

6 mm wide; wings narrowly rectangular,

sometimes auriculate, 3.5—5.7 mm long, 0.8—

1.4 mm wide, on claw |.4—1.7 mm long; keel

about as long as wings; ovary densely appressed

pubescent; style flattened in upper half; stigma

minute, surrounded by hairs. Pods of 2 (rarely

1) articles, each 3.5—5 mm long, 2.8-4.2 mm

wide, appressed hairy; seeds 2.2—2.8 mm long,

1.6—2 mm wide.

Selected specimens: Western Australia. Carson River,

in 1891, Bradshaw & Allen (MEL, NSW 213630); Gorge

of Glenelg River, Jul 1950, Gardner 9663 (PERTH).

Northern Territory. Arnhem Land, in 1928, Basedow

157 (AD, K); Katherine Gorge National Park, 14°19’S

132°28’E, Jun 1983, King 252 (DNA); Groote Eylandt,

May 1972, Levitt (DNA); South Bay, Bickerton Is.,

13°45’S 136°06’E, Jun 1948, *Specht 508 (AD, BRI,

CANB, K, MEL, PERTH); Gove, Jun 1972, Maconochie

1532 (DNA, K, MEL, PERTH); c. 39 miles [62 km] NE

of Maranboy Police Station, Mar 1965, Lazarides &

Adams 113 (BRI, CANB); near Knuckey Lagoon,

Nicholson’s Place, NE of Katherine, Apr 1956, Burbidge

5107 (BRI, CANB); 7 km from ‘Mudginberri’ along road

to Oenpelli, 12°33’S 132°54’E, Apr 1980, Telford 7752

& Wrigley (CANB, CBG); East Alligator River, 11°59’S

133°41°E, Feb 1973, Dunlop 326 (DNA); Angularli

Creek, Murganella-Oenpelli Road, 11°46’S 133°10°E,

214

Feb 1984, Whiteman 1093 (DNA). Queensland. Cook

DistricT: Cape York, 10°42’S 142°32’E, Jul 1985, Thiele

919 (BRI, CANB); Lockerbie, 10 miles [16 km] WSW

of Somerset, Apr 1948, *Brass 18464 (BRI, CANB, K);

Captain Billy Landing, 11°35’S 142°43’E, May 1971,

Norris [AQ 230277] (BRI); Mapoon, 11°58’S 141°52’E,

Jun 1985, Gunness AG 2024 (BRI); 17 km from Merapah

homestead, 13°49’S 142°22’E, May 1987, Clarkson 7143

& Simon (BRI, K, MBA, QRS). Papua New Guinea.

Between Laloki & Hiwick R., 9°25’S 147°17’E, Feb

1964, *Womersley 19094 (BRI, K, L, NSW); Port

Moresby, Feb 1937, *Brass 8781 (BRI, BM).

* These and other specimens seen, determined

by Ohashi (1973) as Dicerma biarticulatum

subsp. australiense.

Distribution and habitat: The species occurs

in sandy soils, sometimes on sandstone, usually

in open eucalypt communities, though also

reported from the edges of Melaleuca swamps

and on coastal dunes. It occurs in the Kimberley

region of Western Australia, the Northern

Territory (north of about 15°S latitude), eastern

Queensland from Cape York south to about

19°S latitude, and near Port Moresby in Papua

New Guinea. There are notable disjunctions

between the Kimberley and Northern Territory

occurrences and across the southern shore of

the Gulf of Carpentaria. Map 3.

Affinities: Aphyllodium schindleri is

distinguished from other species by its dense

inflorescences and remarkably long bracts and

bracteoles; its closest relative among Australian

species is probably A. latifolium. From

specimens determined by Ohashi at BRI and

other herbaria, his key to subspecies and the

distribution map published by him (Ohashi

1973 ), it appears that he considered this to be

Dicerma biarticulatum subsp. australiense

Schindl. Schindler (1924) recognised three

formae of D. biarticulatum var. australiense,

each of which is treated as a distinct species

here. ‘Typical’ subsp. australiense (that is, f.

australiense, f. ‘genuinum’ of Schindler) is

included in Aphyllodium biarticulatum. The

identity of Dicerma biarticulatum var.

australiense, its typification, and the

misapplication of the name Aphyllodium

novoguineense are discussed under A.

biarticulatum.

Austrobaileya 5(2): 209-261 (1999)

Etymology: The _— specific epithet

commemorates Anton Karl Schindler (1879-—

1964) who, in a remarkably brief but fruitful

botanical career in the first quarter of this

century, laid solid foundations for later critical

work on the Desmodiineae. Regrettably a

quantity of his work remained unpublished on

his retirement from plant taxonomy (see

Schubert 1964).

4. Aphyllodium glossocarpum Pedley, sp.

nov. ab aliis speciebus Aphyllodii

fructibus in lomenta distincta aegre

separatis, lomento solitario vel distali 7—

9 mm longo purpurascenti distinguenda.

Typus: Western Australia. 19.5 km from

Beagle Bay Mission turn-off on the road

to Cape Leveque, June 1981, B.R. Maslin

4928 (holo: PERTH; iso: BRI, K).

Dicerma biarticulatum f. plumosum

Schindl., Rep. sp. nov. reg. veg. 20:268

(1924). Type: Western Australia.

Brunswick Bay, Oct 1820, _ A.

Cunningham 267 (holo: K; 1so: BM).

Desmodium novae-hollandiae_ var.

caudatum Domin, Biblioth. Bot. 89:212

(1926). Type: Western Australia.

Brunswick Bay, Oct 1820, _ A.

Cunningham 267 (holo: K; iso: BM).

Shrub to 2 m tall; branches with sparse

indumentum of appressed hairs c. 1 mm long;

stipules + glabrous, 5—12 mm long. Leaflets

linear, narrowly oblong or narrowly obovate,

obtuse at the base, obtuse or acutish at the tip,

glabrous on the upper surface, sparsely

appressed pubescent on the lower; terminal

leaflets 15—40 mm long, 3.5—5.5 mm wide, 3.5—

8 times longer than wide, the laterals slightly

smaller; stipels c. 0.5 mm long; petioles 5—10

mm long. Inflorescences to c. 25 cm long;

primary bracts 3.5—5.5 mm long, secondary

bracts slightly smaller, both with a few marginal

hairs; pedicels 1.4—2.5 mm long. Flower: calyx

with sparse indumentum of short (c. 0.05 mm

long) uncinate hairs, 3.5—4.5 mm long, the tube

c. 2 mm long, the lobes 1.5—2.5 mm long;

bracteoles about as long as the tube; corolla

mauve to pale purple; standard obovate 6.5—

7.5 mm long, 3.5—4.5 mm wide; wings narrowly

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Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 215

rectangular, not auriculate, 4—5 mm long, 1.2—

1.4 mm wide, on claw 2—2.4 mm long; keel

about as long as wings; ovary densely

pubescent; stigma surrounded by hairs. Pods 1

or 2 seeded, scarcely divided into distinct

articles, the proximal one 5.5—7.5 mm long, the

distal or single one longer, 7—9 mm long, all

3.2—4.5 mm wide, appressed pubescent; seeds

c. 3 mm long and 2 mm wide.

Specimens: Western Australia. 1 km NW of Beagle Bay

Mission, 16°59’S 122°40’E, Apr 1985, Martin 31

(CANB, PERTH); Wonganut Spring, 19 km ESE of

Coulomb Point, Dampier Penin., 17°15’S 122°19’E, Jun

1984, Kenneally 9039 (PERTH).

Distribution and habitat: The species is known

only from the four collections cited, all from

coastal areas of the Kimberley region, Western

Australia. Cunningham’s specimen probably

came from Marigui Promontory (c. 15°15’S

125°00’E) where he ‘obtained a large addition

to his collection’ (King, 1827:439) on this, the

last anchorage of the ‘Mermaid’ on the north-

west coast. It is reported to occur on sand

(Kenneally, in sched.) Map 4.

Affinities: Aphyllodium glossocarpum bears a

general resemblance to A. biarticulatum but

differs from it and other species in having

purplish fruits which are scarcely divided into

distinct articles. The single or distal loment

(when two are found) is 7—9 mm long.

Etymology: The epithet is from Greek g/ossa,

tongue and karpos, fruit, an allusion to the long

purplish tongue-like articles of the fruit.

5. Aphyllodium stylosanthoides Pedley, sp.

nov. aspectu distinctivo sed simul A.

biarticulato (L.) Gagnep. arcte affinis a

quo differt stipulis plerumque

longioribus, 10—20 mm longis, petiolis

longioribus, 10—16 mm longis, foliolis

plus minusve acutis longioribus, (15—)20—

35(—55) mm longis, elongatioribus, 4—7(—

12)-plo longioribus quam latis; ab A.

schindleri Pedley foliolis plerumque

angustioribus, 3—6 (raro 12) mm latis,

bracteis plerumque brevioribus, 4.5—10.5

mm longis, bracteolis brevioribus, 3.5—

4.2 mm longis; ab A. latifolio Pedley

stipulis longioribus, foliolis angustioribus

elongatioribus. Typus: Western

Australia. Near Wonjarring Gorge in

Carson Escarpment, 10 km E of ‘ Theda’

H.S., 25 March 1978, M. Lazarides 8712

(holo: CANB; iso: BRI, K, NSW,

PERTH).

Erect or ascending annual stems to c. | m from

perennial woody rootstock; branchlets with

indumentum of sparse to moderately dense

appressed hairs up to 2 mm long; stipules 10—

20 mm long, glabrous or with a fringe of hairs.

Leaflets linear, narrowly oblong, narrowly

elliptic or occasionally oblanceolate or

narrowly obovate, + acute, glabrous on upper

surface, with sparse appressed hairs on lower

surface, dense on margins and midrib, as well

as minute (< 20 needed) hairs on lower surface,

rarely also on upper; terminal leaflets 15—35(—

55) mm long, 3—6(—12) mm wide, 4—7 times

longer than wide, laterals a little smaller; stipels

glabrous, c. 1—-1.5 mm long; pulvini densely

pubescent, 1—2 mm long; petioles 10-16 mm

long. Inflorescences to 12.5 cm long, but often

shorter; primary bracts 4.5—10.5 mm long,

secondary bracts slightly shorter; pedicels 1.5—

2 mm long. Flower: calyx with sparse short

hooked hairs, 3.3—4 mm long; corolla pink;

standard narrowly ovate c. 6.5 mm long, 3.3—4

mm wide; wings narrowly rectangular, not

auriculate, 4—-5.2 mm long, 0.5—1.2 mm wide

on claw 1.5—2 mm long; keel about as long as

the wings; ovary densely pubescent, style

flattened towards the tip. Pods of 1 or usually

2 articles, each 3.8—5.3 mm long, 2.8—4 mm

wide, rather sparsely pubescent at maturity with

short appressed hairs; seeds c. 3 mm long, |.5—

2 mm wide.

Selected specimens: Western Australia. Carson

Escarpment, S of Coucal Gorge, Drysdale River National

Park, + 150°2’S 126°49’E, Aug 1975, George 13883

(PERTH); Bioga Falls, Drysdale River National Park,

15°08’S 127°06’E, Aug 1975, Kenneally 3061 (PERTH).

Northern Territory. 10 km SE of Nourlangie Ranger

Station, 12°50’S 132°42’E, May 1980, Lazarides 8890

(CANB); 5 km NNW of Koongarra, 12°50’S 132°50°E,

May 1980, Craven 5580 (CANB, MEL); 21.5 miles [34.5

km] from Adelaide River camp, 13°02’S 131°03’E, Aug

1946, Blake 16715 (BRI, DNA); 6 miles [10 km] from

Litchfield, 13°27’S 130°307’E, Jul 1946, Blake 16644

(BRI, DNA, K, MEXU, MO); Noonamah [12°38’S

131°04’E], Apr 1968, Byrnes NB602 (DNA, NSW).

Distribution and habitat: The species has been

recorded from habitats similar to those of A.

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216

schindleri: sandy soil, sometimes shallow or

rocky, in eucalypt communities (Lucalyptus

tetrodonta F. Muell and &£. miniata A. Cunn.

ex Schauer particularly), but also in damp

situations where it is associated with Melaleuca

spp. It occurs in the eastern part of the

Kimberley region of Western Australia and the

north-western part of the Northern Territory.

Map 5.

Affinities: Aphyllodium stylosanthoides has a

distinctive facies but appears to be close to A.

biarticulatum. It has generally longer stipules,

petioles and leaflets; the leaflets are more or

less acute and have a greater length/width ratio.

It is less closely related to A. schindleri and A.

latifolium; the former has longer bracteoles and

usually longer bracts and wider leaflets; and

the latter shorter stipules and wider leaflets.

Etymology: The specific epithet is derived

from the generic name Stylosanthes. In habit

and in its rather slender reddish stems, A.

stylosanthoides resembles some species of

Stylosanthes introduced into tropical Australia

as fodder plants.

6. Aphyllodium biarticulatum (L.) Gagnep.,

Not. Syst. 3(8):254 (1916); Pedley, Rev.

Hand. Fl. Ceylon 10:167 (1996);

Dicerma biarticulatum (L.) DC., Prodr.

2:339 (1825), nom. illeg.; Echinolobium

biarticulatum (L.) Desv., Mem. Soc.

Linn. Paris 4:310 (1825); Desmodium

biarticulatum (L.) F. Muell., Fragm. Phyt.

Austr. 2:121 (1861); Hedysarum

biarticulatum L., Sp. Pl. 747 (1753).

Type: Herb. Hermann vol. 3 fol. 15, No.

296 (lecto: BM, fide Pedley in Turland

& Jarvis (1997)).

Dicerma biarticulatum var. australiense

Schindl., Rep. sp. nov. reg. veg. 20:268

(1924) (excl. f. /ongibracteatum Schind..

& f{. plumosum Schindl.); Desmodium

biarticulatum var. australiense (Schindl.)

Meeuwen, Reinwardtia 6:247 (1962);

Dicerma biarticulatum — subsp.

australiense (Schindl.) Ohashi,

Ginkgoana 1:255 (1973); Aphyllodium

australiense (Schindl.) Ohashi, Taiwania

42: 143 (1997). Type: Bowen River,

Austrobaileya 5(2): 209-261 (1999)

Bowman 269 (lecto: MEL, chosen

here).

Desmodium novae-hollandiae Domin,

Biblioth. Bot. 89:212 (1926) var. novae-

hollandiae (typicun’ Domin). Type:

Sandige Mischwalder in der Nahe der

Russell-Mundung, Feb 1910, Domin

‘4758’ (lecto: PR 527353, chosen here).

Desmodium novae-hollandiae f. latifoliuim

Domin, loc. cit. Type: Savannenwalder

zwischen Chillagoe und Walsh River, Feb

1910, Domin ‘4755’ (holo: PR 527350).

Prostrate, ascending or erect subshrub to c. 0.5

m high usually with dense indumentum of

appressed or somewhat spreading white hairs

to c. | mm long; stipules 4-13(—15) mm long

with a few hairs on margin or tip. Leaflets

linear, narrowly elliptic, narrowly obovate,

elliptic or obovate, attenuate at the base, obtuse,

mucronulate at the apex, glabrous or with

Sparse appressed or weak ascending hairs on

upper surface, sparsely appressed pubescent or

with rather dense ascending hairs on lower

surface; terminal ones 7—25(—38) mm long, 2

6.5(-10) mm wide, (1.5—)2.5—6.5(—9) times

longer than wide, the laterals usually a little

smaller, (6—)7.5—22 mm long, 2—6(--8) mm

wide, 1.5—-5.5(—9) times longer than wide;

stipels 0.3—0.7(—1) mm long often hidden in

the hairs of the pulvinuses which are 0.6—1(—

1.5) mm long; petioles 3—9(-12) mm long.

Inflorescences terminal and axillary, rather

dense, up to 20 cm long, the rachis with

indumentum of moderately dense short

uncinate hairs, often with some straight

appressed hairs towards the base; primary

bracts 3.5—-10 mm long, secondary bracts 2.5-

7 mm long, pedicels with indumentum of short

uncinate hairs, 0.9-—-1.8(—2.7) mm long at

anthesis. Flower: calyx with sparse indumentum

of short uncinate and a few long straight white

hairs, 2.5-—-3.5(—4) mm long, the tube 1.2—1.8(-

2.2) mm long, the lobes about equal in length,

1.2—2.2 mm long; bracteoles (1.7—-)2—3.6(-4.8)

mm long; corolla described by most collectors

as mauve or pink, but also bluish, purple or

red, standard obovate, (5—)5.5—7.5 mm long,

(2.5—)3—4.4 mm wide; wings narrowly oblong,

+ auriculate, obtuse (2.8—)3—-4.7 mm long on

aelaseset ete

adeee ecto ee een

PRMRITMPA eee

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 217

claw (0.6—)0.8—1.3 mm long; keel petals about

as long as the wings; ovary with dense

appressed or rarely ascending hairs. Pods with

1 or 2 articles, each 3—5 mm long, 2.5-4 mm

wide with indumentum of appressed hairs,

glabrescent, or dense + spreading and often

persistent hairs; seeds 2—3.8 mm long, |.2—2.2

mm wide.

Selected specimens: Western Australia. Near junction

of Drysdale River & Mogurnda Creek, 15°02’S 126°5S’E,

Aug 1975, George 1353 (PERTH); 60 km from

Kununurra, 16°O1’S 128°46°E, Apr 1977, Pullen 10686

(CANB, NSW). Northern Territory. 30 miles [48 km]

E of ‘O.T. Station, Aug 1948, Perry 1487 (BRI, CANB,

NSW); 11 miles [18 km] NE of ‘Alexandria’ Station, Jun

1948, Perry 1472 (AD, BRI, CANB, K, MEL, NSW,

PERTH). Queensland, Burke District: Woolgar, c.

19°40’S 143°10’°E, Aug 1915, Bick (BRI, NSW);

Between ‘Corinda’ & ‘Westmoreland, 17°40’S 138°28’E,

Pullen 10686 (CANB, NSW); Albert River, Mueller (K,

MEL); Sweers Is., Nov 1802, Brown (BRI, BM); same

place, Henne (MEL) & same place, Jun 1901, JE Bailey

[AQ 97444] (BRI); ‘Chudleigh Park’ Station, 110 miles

[176 km] N of Hughenden, Feb 1931, Hubbard &

Winders 7606 (BRI, K); Coox District: Yorkey’s Knob

beach, Apr 1962, McKee 9034 (BRI, CANB, K, NSW);

3 miles [5 km] E of Mareeba, May 1953, Thorne 21116

& Jones (BRI, RSA); NortH KENNEDY District: Magnetic

Is., Jan 1931, Hubbard & Winders 6665 (BRI, K); 6 km

N of Elliot River, c. 19°50’S 147°S0’E, Apr 1975,

McDonald & Batianoff 1386 (BRI, K); MircHELy

District: c. 12 km SW of Barcaldine 23°29’S 145°12’E,

Jun 1977, McDonald 2692 (BRD; ad opp. Jericho, Donzin

‘4760’? (PR).

Distribution and habitat: Aphyllodium

biarticulatum is restricted to sandy soils in open

situations 1n eucalypt communities, on coastal

dunes, on roadsides and occasionally in

cultivation. In Australia it extends from the

northern part of the Kimberley region of

Western Australia, across the north of the

Northern Territory to Queensland south to the

Tropic of Capricorn. It appears to be more

common in the east of its range than it is in the

west. Its extreme variability within Australia

is discussed below. Outside Australia it extends

through Malesia to southern China, Indo-China,

India and Sri Lanka. Map 6.

Affinities: The species is closely related to A.

parvifolium, A, stylosanthoides and A.

hispidum (Schindl.) Ohashi which have rather

restricted ranges, the first two in north-western

Australia, the last in south-eastern Asia. A.

latifolium, A. schindleri, A. glossocarpum and

probably A. novoguineense are less closely

related to 1t and may represent a more remote

evolutionary line.

Variation and typification: Beside

nomenclatural problems discussed (p. 211) the

treatment of Hedysarum biarticulatum L. has

been confused by different classifications

resulting from differing concepts of genera and

infra-generic taxa.

Mueller (1861), in transferring H. biarticulatum

to Desmodium noted that Wight’s figure (Icon.

Plant. Ind. Or. 2 (1841) t. 419) did not at all

clearly represent (‘haud clare exprimit’) the

Australian species, but invoked the authority

of J.D. Hooker in treating Indian and Australian

species as the same. Bentham (1864) noted that

some Australian specimens differed from the

Indian ones but considered all belonged to one

species; despite Bentham’s remark to the

contrary, Wight’s figure is not a good

representation of the species, either in India or

Australia.

Schindler (1924) and Domin (1926) used

much the same set of specimens, but

independently produced significantly

different infra-specific classifications of A.

biarticulatum. Schindler considered the whole

genus Dicerma (sensu Bentham) while Domin

only the Australian occurrences of Desmodium

biarticulatum sensu Mueller ( and Bentham).

Schindler described Dicerma hispidum and D.

novoguineense from single specimens from

Burma and New Guinea respectively and

divided Dicerma biarticulatum into six taxa:

three varieties, one consisting of three formae.

They are:

Dicerma biarticulatum QL.) DC.

var. biarticulatuim

var. collettii Schindl.

var. australiense Schindl.

f. qustraliense (£. genuinum Schindl., nom.

inval.)

f. longibracteatum Schindl.,

f. plumosum Schindl.

218

He considered D. biarticulatum var.

biarticulatum to be widely spread on all coasts

of the Indian Ocean and to be very uniform

everywhere. Evidently, however, he did not

believe it to occur in Australia as he cited no

Australian specimens and considered

Desmodium biarticulatum sensu Mueller and

Bentham to be Dicerma biarticulatum var.

australiense which he described as ‘varietas

valde variabilis’. I have seen no material of

Dicerma biarticulatum vay. collettii which was

described from a single collection from

“Upper Burma”. Dicerma biarticulatum Tf.

longibracteatum and f. pliumosum are treated

here as distinct species Aphy/lodium schindleri

and A. glossocarpum respectively. With the

removal of these, var. australiense still exhibits

a wide range of variation..

Domin placed Hedysarum biarticulatum

in Desmodium and described D. novae-

hollandiae as new, referring to it most, though

not all, Australian specimens previously

considered D. biarticulatum. He subdivided D.

novae-hollandiae into four taxa: two varieties,

one made up of two formae:

Desmodium novae-hollandiae Domin

var. novae-hollandiae (var. typicum Domin,

nom. inval.)

{. novae-hollandiae - automatically

{. fatifolium Domin

var. caqudatum Domin

Desmodium biarticulatum var. caudatum 1s

based on Cunningham 267 which is also the

type of Dicerma biarticulatum f. plumosum

Schindl. and consequently is treated here as

Aphyllodium glossocarpum. Here the

correspondence between Domin’s and

Schindler’s classifications ends. Domin was

aware of the diffuseness of his classification

and remarked that in f. /atifolium one could see

a weak approach to var. caudatum and that

Brown ‘4183’ was almost intermediate between

the type (that is, f. novae-hollandiae) and var.

caudatum. Brown 4183 is, in fact, one of the

two syntypes of Dicerma biarticulatum f.

longibracteatum Schindl. Furthermore,

Austrobaileya 5(2): 209-261 (1999)

Domin cited one specimen as Desmodium

biarticulatum sensu stricto, Schultz 534,

which is a syntype of Dicerma biarticulatum

f. australiense Schindl.

Later workers have modified Schindler’s

scheme of classification but have paid scant

attention to Domin’s. Regrettably these later

workers have either not consulted Schindler’s

and Domin’s type specimens or, if they have,

they have not cited them. Knaap-van Meeuwen

(1962) included Dicerma in Desmodium, She

suggested that Desmodium novae-hollandiae

to be not specifically distinct from D.

biarticulatum (to which she also referred

Dicerma biarticulatum var. collettii, Dicerma

hispidum and Dicerma novoguineense) and

transferred Dicerma biarticulatum var.

australiense to Desmodium biarticulatum.

Study of her key shows that her concept of var.

australiense was based entirely on f.

longibracteatum and not at all on either f.

australiense or f. plumosum. Verdcourt (1979),

whose work was not intended to be a critical

treatment, followed Knaap-van Meeuwen

exactly.

To a large extent Ohashi (1973) adopted

schindler’s generic concepts but, with a change

of rank, Knaap-van Meeuwen’s treatment of

infrageneric taxa. He accepted Dicerma

biarticulatum subsp. biarticulatum, subsp.

australiense (Schindl.) Ohashi and subsp.

hispidum (Schindl.) Ohashi. From the key to

subspecies, the distribution map of Dicerma

biarticulatum and the photograph of Brass

18464, it is evident that Ohashi also based his

subsp. australiense solely on Dicerma

biarticulatum f. longibracteatum. Ohashi cast

doubt on Knaap-van Meeuwen’s treatment of

Dicerma novoguineense but took no action,

probably because he had seen no specimen.

Ohashi (1997b) accepted Aphyllodium as the

correct name for the genus previously treated

by Bentham as Dicerma (see p. 211 ). He effected

the transfer of some names from Dicerma to

Aphyllodium and modified his previous

treatment. In doing so he further complicated

an already tangled situation. He cited no

specimens, type or otherwise, and seems not

to have appreciated the desirability of

lectotypifying names to establish their

Pediey, L., Desmodium and related genera in Australia: a taxonomic revision 219

application. He changed the status of Dicerma

biarticulatum f. australiense which he raised

to specific rank as Aphyllodium australiense.

He transferred D. novoguineense to

Aphyllodium, placing D. biarticulatum f.

longibracteatum, f. plumosum and f.

australiense (p. p.) in its synonymy. He gave

its distribution as Australia and New Guinea.

Perhaps, appreciating that D. novoguineense,

D. biarticulatum {. longibracteatum and D.

biarticulatum { plumosum differ from D.

biarticulatum {. australiense most conspicuously

in having larger leaflets, stipules and bracts,

he referred them to A. novoguineense, since

the epithet novoguineense must be used at

specific rank,

Application of the name A,

novoguineense presents a major difficulty. I

have not seen the type but have seen one poor

specimen of the species ( J.H. Barrett 4270,

BRI ex LAE) collected from near the type

locality, Astrolabe Range, Papua New Guinea.

Examination of it and a reading of the

protologue have persuaded me that it represents

a species distinct from A. schindleri (D.

biarticulatum {. longibracteatum), A.

glossocarpum (D. biarticulatum f. plumosum)

and A. australiense, the last of which as

lectotypified (p. 216), is the same as A.

biarticulatum sensu stricto. A. novoguineense

has stems with long ascending fulvous hairs and

usually longer stipules and bracts than have

other taxa. It appears to be rare and restricted

to Papua New Guinea.

In habit, size and shape of vegetative parts

and in the indumentum of stems, leaves and

pod-articles A. biarticulatum is extremely

variable, with the degree of variability within

Australia probably greater than it 1s outside the

continent. This variability is reflected in the

different treatment of species discussed above.

study of specimens from Sri Lanka (Pedley

1996), the type locaity of Hedysarum

biarticulatum, indicated that though the plants

there tend to have somewhat smaller stipules

and primary and secondary bracts, they fall

within the range of variability of Australian

plants and recognition of a subspecies or variety

for Australian plants is not warranted. |

After specifically distinct entities have

been removed A. biarticulatum remains a

disconcertingly variable species. Preliminary

studies suggested that two variants could be

recognised: (a) a prostrate plant with narrow

(2—3 mm wide) leaflets found usually on coastal

sands on the east coast, and (b) subshrub with

short obovate leaflets 1.5—3 times longer than

wide with large pod articles with dense

somewhat spreading hairs which occurs in

semi-arid areas inland around the south-western

part of the Gulf of Carpentaria. Though

extremes of these variants are distinctive the

variants merge into the more ‘typical’

populations of the species and forma!

recognition is not warranted.

Dendrolobium (Wight & Arn.) Benth. in

Miquel, Pl. Junghuniana: 215,216

(1852); Schindl., Rep. sp. nov. reg. veg.

20:278 (1924); Hutchinson, Gen. PI.

1:483 (1964); Ohashi, Ginkgoana 1:50

(1973), Taiwania 42: 135 (1997);

Desmodium subg. Dendrolobium Wight

& Arn., Prodr. 1:223 (1834); Baker in

J.D. Hooker, Fl. Brit. India 2: 161 (1876);

Desmodium sect. Dendrolobium (Wright

& Arn.) Benth. in Bentham & J.D.

Hooker, Gen. Pl. 1:519 (1868). Type:

Dendrolobium umbellatum (L.) Benth.

Holtzea Schindl., Rep. sp. nov. reg. veg.

22:285 (1926). Type: H. umbellata

schindl. (= Dendrolobium stipatum 8.1.

Blake). |

Ohashi (1973) characterised Dendrolobium by

its axillary subumbellate inflorescences,

uniparous flowers, monadelphous stamens,

conspicuously long styles, thick corky or

coriaceous pericarps and conspicuously rim-

arillate seeds. The inflorescences of D.

polyneurum are elongate racemes, but the

combination of the other characters listed

distinguished Dendrolobium sharply from all

other genera except Phyllodium, The emphasis

placed on the foliaceous bracts of Phyllodium

obscures the relationship of it with

Dendrolobium. This relationship 1s discussed

under Phyllodium.

Ohashi (1973) recognised 11 species in

the genus with the Indo-Chinese region the

220

centre of diversity with eight species occurring

there. The recognition of five species confined

to the Australian region in the present study

places northern Australia as another area of

Austrobaileya 5(2): 209~261 (1999)

speciation. Some of the Australian species are

adapted to a strongly seasonal climate with

wildfires in the dry season.

Key to species

1. Lower calyx lobe markedly (1 mm or more) longer than other lobes ................. 2

Lower calyx lobes almost the same length as other lobes ........ 0.0... cece ees 3

2. Plant prostrate; branchlets with spreading or ascending hatrs; pedicels

4—10 mm long; bracteoles c. | mm long. .

* € © &© @&© © FF EF &#& FF & #© FF BP & €£ BF FF & HF HF £

1. D. multiflorum

Plant a shrub or tree to 6.5 m (? 12.5 m); branchlets with dense white

appressed hairs; pedicels c. 2 mm long; bracteoles 2.5—3.6 mm long ...

6. D. arbuscula

3. Flowers in elongate racemes, axis up to 70 mm long; leaflets with dense

silvery appressed hairs on underside. ....

Wsted ihe BERR niaedes gy Stead og. Meteo 2. D. polyneurum

Flowers in umbellate racemes, sessile or axis to 10 mm long; undersides of

leaflets with appressed hairs, sometimes dense but not silvery..............000 eee 4

4. Wing petals shorter and much narrower than the keel petals; terminal leaflets

50-140(-170) mm x (25—)35—70(-85) mm

yA or Sy ee eee Laat Ley 5. D. umbellatum

Wing petals as long as or longer than keel petals and not markedly narrower

than them; terminal leaflets to 65mm xX 30mm...... ee ee ee ee eens 5

5. Veins conspicuously depressed on upper surface of leaflets; terminal leaflets

8—I4(—-17) mm wide; pedicels (2—)5—7 mm long; ...... 0... 0... eee eae

Veins not conspicuously depressed on upper surface of leaflets; terminal

leaflets 17-30 mm wide; pedicels 6-16 mm long ...............008.

1. Dendrolobium multiflorum Pedley, sp.

nov. affinis D. stipato S.T. Blake; habitu

prostrato, plerumque petiolis brevioribus,

foliolatis latioribus instructis, calycis

lobis (praesertim inferioribus)

longioribus petalis parvioribus differt.

Typus: Northern Territory: 4 miles [6

km] SE of Katherine [14°29’S 132°19’E],

December 1963, M Lazarides 6995 (holo:

CANB; iso: BRI, DNA, MEL, PERTH).

Annual stems up to 1.2 m long trailing from

perennial rootstock; branchlets angular with

indumentum of crisp matted hairs to 0.5 mm

long; stipules striate, pubescent on the outside,

47 mm long. Leaflets ovate or broadly ovate,

obtuse or slightly mucronulate, discolorous,

sparse slightly ascending hairs on both surfaces,

mainly on veins beneath, sometimes becoming

glabrous on upper surface, 8—11 lateral veins

on each side of midrib; terminal leaflet 50-90

mm long, 30-55 mm wide, laterals distinctly

smaller 30-45 mm long, 17-35 mm wide, both

1.3—2 times longer than wide; rachis 8-17 mm

long, petiolules 1.3-—2.3 mm long, stipels 1-3

mm long, petioles 8-11 mm long. Flowers in

fascicles of 10-20, sessile in the upper axils;

bracts 2—2.5 mm long, bracteoles sometimes a

little below the base of the calyx, c. 1 mm long;

pedicels 4-10 mm long, with spreading white

hairs. Flowers: calyx with spreading hairs, 5.3—

6 mm long, tube c. 1.7 mm long, upper lobe

2.3—3.5 mm long, laterals the same size or

shorter, 2.3-3 mm long, the lower setulose,

incurved, distinctly longer, 3.6—4.4 mm long;

corolla white or yellowish tinged with pink:

keel oblong, rounded at the top, narrowed but

not clawed at the base, 6.5—8 mm long, 4.5—6

mim wide; wings 4.5—6 mm long on claw 1—1.5

mim long, slightly shorter (by 0.5 mm) than keel

petals; ovary pubescent with 1 ovule. Pod not seen.

Other specimens: Northern Territory. 22.4 miles [35.8

km] E of Adelaide River [c. 13°25’°S 131°25’E], Mar

1961, Chippendale NT 7716 (BRI, DNA, K MEL,

PERTH); Bull Creek, 13°54’S 131°17°E, Jul 1946, Blake

16294 (BRI).

SASSAN SS EM SES SSCS ES nd By VE vic nak

"TUSHAR et a I

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 221

Distribution and habitat: The species is

restricted to a small area in the extreme north

of the Northern Territory where it occurs on

sandy soils. Map 7.

Affinities: Dendrolobium multiflorum appears

to be related to D. stipatum but differs in its

prostrate habit, wider and usually less elongate

leaflets, fascicles with more flowers and the

lower lobe of the calyx longer than the others.

Etymology: The specific epithet is derived

from Latin, mu/tus many, and flos, floris,

flowers, an allusion to the plant’s many

flowered fascicles.

2. Dendrolobium polyneurum (S.1T. Blake)

Pedley, comb. nov.

Desmodium polyneurum S.T, Blake, Austr.

J. Bot. 2:119 t.3 (1954). Type: Northern

Territory. Near Station Creek, 13°42'4’S

131°04’E, July 1946, 8.7) Blake 16474

(holo:BRI; tso:K).

Ascending annual stems to 1 m long arising

from perennial rootstock; branches + terete with

indumentum of dense appressed matted hairs

c. | mm long; stipules deltoid, connate nearly

to the middle, pubescent 3—5.5 mm long, early

deciduous. Leaves: leaflets obovate or oblong,

obtuse or retuse, sometimes mucronulate,

lamina with dense short appressed hairs on both

surfaces, becoming rather sparse on upper

surface, 10-13 lateral veins on each side of

midrib; terminal leaflet 40-90 mm long, 23—

70 mm wide, 1.6—2 times longer than wide,

laterals smaller, usually oblong, 35-70 mm

long, 20-40 mm wide; stipels setaceous, 1-2

mm long; petiolules 2—3.5 mm long, rachis 4—

20 mim long; petioles 3-20 mm long. Flowers

in axillary racemes up to 70 mm long, the young

unopened flowers crowded at the apex of the

rachis; bracts ovate-acuminate, 2~-3 mm long,

deciduous; pedicels 2-7 mm long; bracteoles

1.5—2 mm long. Flowers: calyx 5.5-7.5 mm

long, tube 3 mm long, obtuse at base, gradually

widening upwards; lobes lanceolate, acute, the

two upper ones at first connate nearly to the

tip, finally separating; corolla white: standard

obovate, c. 10 mm long and 7 mm wide, +

clawed; wings 10 mm long, spurred near the

base; keel petals as long as the wings,

subobtuse, slightly incurved; ovary hirsute.

Pods densely pubescent becoming only

moderately so when old, both margins

constricted between the seeds; articles (1—-)2-

4, longitudinally oval, nearly symmetrical, 6—

9 mm long, 5-6 mm wide, woody and

indehiscent; seeds not seen mature.

Selected specimens: Northern Territory. 2.5 miles [4

km] SW of ‘Tipperary’ homestead, Jul 1961, Lazarides

6677 (CANB, K, MEL PERTH); 39 miles [50 km| SW

of ‘Dorisvale’ Station [14°46’S 131°06’E] May 1952,

Perry 2796 (CANB, K, MEL); 6 miles [10 km] N of

Wilton River Crossing, Jun 1972, Byrnes 2617 (DNA, K).

Distribution and habitat; Dendrolobium

polyneurum occurs only in the northern part of

the Northern Territory on sandy soils in

eucalypt communities. It has not often been

collected and seems to be rare, Like D. cheelii

and D. stipatum the species appears to be a

pyrophyte with annual stems arising from a

woody perennial rootstock. Map 8.

Affinities: In the protologue of Desmodium

polyneurum, Blake referred it to Desmodium

sect. Dollinera (Endl.) Schindl. and it does bear

a general resemblance to some species of that

section, particularly D. hayatae Ohashi. Its

monadelphous stamens and bracteoles support

such a placement, but the structure of its

inflorescence is quite different from that of sect.

Dollinera. Its flowers are arranged in long

axillary racemes, each pedicel subtended by a

bract, whereas in sect. Do/linera flowers are

in 6-8 flowered fascicles; that is, each bract

subtends 6-8 pedicellate flowers. Though long

racemes are unusual in Dendrolobium, Ohashi

(1973:14) noted some specimens of D.

umbellatum and D. triangulare (Retz.) Schindl.

with ‘a few axillary racemes mixed with short

racemes or subumbels’. The pods of D.

polyneurum are similar to those of other species

of Dendrolobium such as D. arbuscula and D.

umbellatum. Though its axillary racemes sets

it apart from all other species the affinities of

D. polyneurum are with D. stipatum and D.

cheellti.

3. Dendrolobium cheelii (C.A. Gardner)

Pedley, comb. nov.

Alysicarpus cheelii C.A. Gardner, West.

Austr. For. Dept. Bull. 32:57 (1923).

222

Type: Western Australia. Between

Vansittart and Napier Broome Bays,

August 1921, C_A. Gardner 1040 (holo:

PERTH).

Erect annual stems to 0.5 m tall from perennial

woody rootstock; branches with indumentum

of moderate to dense appressed, often matted

hatrs 0.5—0.7 mm long; stipules 3-6 mm long,

striate, sparsely to densely hairy outside,

glabrous inside. Leaves (1-) or 3- foliolate;

leaflets elliptic or obovate, obtuse or rarely

slightly acuminate, mucronulate, sparsely

pubescent with appressed or weak, slightly

ascending hairs on both surfaces, mainly on

veins beneath, 4—10 lateral veins on each side

of midrib, deeply tmpressed on upper surface;

terminal leaflet (13—)20—60 mm long, (4—)8-

14(-18) mm wide, 2.54.5 times longer than

wide, laterals somewhat smaller; stipels to 1.8

mm long, slightly longer than the petiolules;

petiolules 0.5—1.5 mm long, rachis 1-6 mm

long; petioles 4-10 mm long. Flowers in

fascicles of 3-8 in upper axils; bracts and

bracteoles c. 1 mm long, pedicels (2—)5—7 mm

long. Flowers: calyx 3.5—4 mm long with

sparse, longish, + appressed hairs and short, +

erect, uncinate hairs, tube 1.7—-2 mm long, lobes

of+ equal length, 1.6-2 mm long, or the lower

to 2.5 mm long, longer than the others; corolla

pink or yellow with a pink keel; standard +

orbicular 4.8—5.5 mm long, 3.5—4.5 mm wide,

on claw c 1.5 mm long; wings 4.5—5.5 mm

long, c 1.8 mm wide, on claw 1.7 mm long;

keel petals slightly shorter than the wings;

ovary densely pubescent, ovules up to 3. Pod

of 1 or 2 articles, each about 8 mm long and 5

mm wide, reticulately veined, densely

pubescent; seeds not seen.

Other speciinens: Western Australia. Near Carlia

Creek, near Carson River, Drysdale River National Park,

15°02’S 126°49’E, Aug 1975, George 13939, sterile

(PERTH); Little Falls, 14°49°S 125°42’E, July 1976,

Lewis 44 (&, PERTH); 4 miles [6 km] N of mining

campsite, Mitchell Plateau, 14°47°S 125°48’E, Dec

1982, Kenneally 8666 (PERTH); Gibb River, Hann

District, Jan 1951, Gardner 9893 (PERTH).

Distribution and habitat: Dendrolobium

cheelii is confined to the Kimberley region of

Western Australia where it occurs on sandy or

loamy soils in eucalypt woodland. It is a

pyrophyte with annual stems arising from a

Austrobaileya 5(2): 209-261 (1999)

perennial rootstock. One specimen (George

13939) is sterile and has somewhat larger leaves

than other specimens and is possibly from a

plant surviving from the previous wet season.

Map 9.

Affinities: The species is related to.D. stipatum

which has a similar life-form, but is certainly

distinct. D. cheelii has leaflets with veins

prominently depressed on the upper surface of

the leaflets, usually short leaf-rachises, usually

narrower leaflets, shorter pedicels, bracts and

bracteoles, and pods with fewer articles. The

geographic ranges of the two species do not

overlap.

4, Dendrolobium stipatum S.T. Blake, Austr.

J. Bot. 2:117 t.2 (1954). Type: E mouth

of Daly River, 13°23’S 130°32’E, July

1946, S.T. Blake 16670 (holo:BRI;

iso:BRI, K).

Holtzea umbellata Schindl., Rep. sp. nov.

reg. veg. 22:285 (1926). Type: N.

Australia, in 1891, Af Holtze (iso:K).

Subshrub to c. | m high, tufted stems from

perennial rootstock; branches with indumentum

of moderately dense to dense appressed hairs

0.20.4 mm long; stipules triangular, 2—5.5 mm

long, 2 mm wide at base, striate, pubescent

outside. Leaves I- or 3- foliolate; leaflets

oblong, elliptic or ovate, obtuse or retuse,

sparsely appressed pubescent on both surfaces,

the hairs c. 0.2 mm long, 6-10 prominent lateral

veins on each side of midrib; terminal leaflet

38-65 mm long, 17-45 mm wide, 1.8~2.3 times

longer than wide, laterals smaller, 23-45 mm

long, 11-20 mm wide, 1.7—2.3(-2.8) times

longer than wide; rachis 1-12 mm long,

petiolules 1.5—-2.5 mm long, stipels about as

long as the petiolules; petioles 4-14 mm long.

Flowers in fascicules of up to c. 12 in the upper

axils, sessile or on peduncles to 6 mm long;

bracts 1-3 mm long; pedicels 6-16 mm long,

bracteoles at base of calyx, 1—1.5 mm long.

Flowers: calyx c. 4mm long with + appressed

hairs c. 0.3 mm long and short erect uncinate

hairs, tube 2—2.2 mm long; corolla pink, the

standard whitish in the upper half, concave,

oblong c. 10 mm long, 6 mm wide on claw c. 1

mim long; keel petals about as long as the wings;

ovary appressed pubescent. Pods curved with

pe ee aan, =

SURREY RAE EAs BSS

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 223

4-6 articles, each oblong, 6-8 mm long, 3.55

- mm wide, appressed pubescent. Seeds not seen.

Other specinens: Northern Territory. Old Litchfield

Road, 13°32’S 130°54’E, Sep 1975, Dunlop 3588 (DNA,

K); Woodcutters track, 50 miles [80 km] S of Darwin,

Sep 1970, Byrnes 1991 (DNA, K); Batchelor, Sep 1913,

Hill 7023/13 (NSW).

Distribution and habitat: Dendrolobium

stipatum is restricted to a small area in the north

of the Northern Territory where it occurs on

sandy soils in eucalypt woodland. Map 10.

Affinities: The species is closely related to D.

cheelii; differences between them are discussed

under the latter. Blake compared D. stipatum

with D. umbellatum but acknowledged that it

differed sharply from that species.

5. Dendrolobium umbellatum (L.) Benth. in

Miquel, Pl. Jungh. 216 & 218 (1852);

Miquel, Fl. Ind. Bat. 1:262 (1855);

Ohashi, Ginkgoana 1:82 (1973); Pedley,

Rev. Handb. FI. Ceylon 10: 162 (1996);

Desmodium umbellatum (L.) DC., Prodr.

2:325 (1825); Benth., Fl. Austr. 2:230

(1864); Prain J. Asiatic Soc. Bengal

66:387 (1897); Schubert, Fl. Trop. East

Africa Legum.: Papilion.: 455 (1971);

Lui & Chuang, Taiwania 8:98 (1962);

Hedysarum umbellatum L., Sp. Pl.: 747

(1753). Type: Herb. Hermann vol. 2 fol.

26, No.293 (lecto: BM, fide Pedley in

Turland & Jarvis (1997)).

Aeschynomene arborea L.,Sp. Pl.: 713

(1753). Type: Sri Lanka: N. of

Trincomalie, 1 September 1931, N.D.

Simpson 8516 (BM, neotype, fide Pedley

in Turland & Jarvis (1997)).

Two varieties are recognised.

5a.Dendrolobium umbellatum (L.) Benth.

var. umbellatum

Branches and pedicels with indumentum of

silky appressed hairs, becoming sparse; leaflets

ovate or elliptic, sparsely appressed pubescent

beneath, the hairs minute; pods sparsely

appressed pubescent.

Selected specimens: Queensland. Cook District: Boigu

Is., 9°14°S 142°13’E, Oct 1981, Clarkson 3870 (BRI, K,

MBA); Endeavour River, Aug 1820, Cunningham 88

(BM, K); Barnard Is. (no 2), Jun 1848, McGillivray. Voy.

of ‘Rattlesnake’ Bot. 284 (K); NorTH KENNEDY DISTRICT:

Cardwell, c. 18°10°S 146°00’E, Aug 1963, Blake 22070

(BRI, K, MEL); Daydream (West Molle) Is, 20°15’S

148°49°, Jun 1984, Pedley 5220 (BRI, MEL).

Sb. Dendrolobium umbellatum var. hirsutum

(DC.) Pedley, comb. nov. Desmodium

umbellatum var hirsutum DC., Prodt. 2:325

(1925); Dendrolobium umbellatum f.

hirsutum (DC.) Ohashi Ginkoana 1:84

(1973). Type: Calcutta Botanic Gardens,

Wallich 5687D (iso:K)

Branches and pedicels densely hirsute; leaflets

almost orbicular, densely hairy beneath; pods

with + persistent long yellowish hairs.

This variety does not occur in Australia.

Distribution and habitat: In Australia D.

umbellatum var. umbellatum occurs along the

east coast, north of about Mackay (22°S),

usually on sand, often in dense stands above

high-water mark, though sometimes on rocks

- or forming a fringe behind mangroves. It does

not occur on the western shore of Cape York

Peninsula nor farther west in the Northern

Territory or Western Australia. Outside

Australia, the variety extends into the Pacific

(Melanesia as far east as Tonga, Samoa and

Micronesia) and through Malesia to Taiwan,

southern China, Indo-China, the Andaman Is.,

Peninsular India, Sri Lanka to east Africa. Map 11.

Dendrolobium umbellatum var.

hirsutum is found in Madagascar and the

Mascarene Islands. Ohashi in making the

combination Dendrolobium umbellatum f.

hirsutum cited only cultivated specimens and

stated that its native habitat was unknown.

Affinities: Dendrolobium umbellatum, D.

arbuscula, D. quinquepetalum and D.

triangulare are closely related but their

relationship to other species is uncertain. The

wide geographic range of D. umbellatum could

indicate it to be less advanced than the other

species all of which (except for D. triangulare

and possibly D. quinquepetalum) have

restricted ranges and which may have been

derived from it. Its occurrence on sea-shores

and its thick-walled pod articles seemingly

adapted to dispersal by water could, however,

224

point to the wide distribution of the species

regardless of its level of advancement.

Ohashi (1973:85) noted the great

variability of D. umbellatum but could not find

any distinctive local assemblages. However,

he did distinguish D. uwmbellatum f. hirsutum

(DC.) Ohashi, based on Desmodium

umbellatum var. hirsutum DC., which was

described from a plant grown in the Calcutta

Botanic Gardens. D. umbellatum from

Madagascar, Mauritius and Reunion is the same

as D. umbellatum f. hirsutum. It has densely

hirsute branches and pedicels, almost orbicular

leaflets densely hairy and prominently veined

on the lower surface, and pods with more or

less persistent long yellowish hairs, and is

distinct enough to warrant varietal rank.

6. Dendrolobium arbuscula (Domin) Ohashi,

Tarwania 42:137 (1997).

Desmodium arbuscula Domin, Biblioth.

Bot. 89:211 (1926), Type: Queensland.

In xerodrymio ad Chillagoe, Domin

‘4764’ (lecto: PR 527346, chosen here).

Dendrolobium sp. (Mt Scatterbrain J.R.

Clarkson+ 6708) Pedley in Henderson

(ed.): Queensland Plants: Names and

Distribution: 79 (1997).

Shrub or small tree, in Australia up to 6 m tall

but reported to reach 12.5 m in New Guinea

(Pullen 6992; BRI, CANB, K); bark soft, grey;

branchlets covered with rather dense appressed

white hairs, occasionally somewhat retrorse,

Q.3-0.5 mm long; stipules to 10 mm long,

deciduous. Leaves 3-foliolate; leaflets oblong,

elliptic or obovate, rounded or sometimes

attenuate at the base, obtuse or subacute at the

apex, margins tending to be slightly recurved,

glabrous or with sparse appressed hairs on

upper surface, sparse to moderately dense,

appressed or slightly ascending hairs on the

lower, c. 10 lateral veins prominent on each

side of midrib; terminal leaflet 30-65 mm long,

16-32 mm wide, laterals smaller, 20-50 mm

long, 9-17(—24) mm wide, both (1.4-)1.8-2.6

times longer than wide; stipules 1.5—4 mm long,

usually about as long as petiolules, 1.5—3.5 mm

long; rachis S—9 mm long; petioles 9-16 mm

long. Flowers mn condensed, umbellate, axillary

Austrobaileya 5(2): 209-261 (1999)

racemes, sessile or peduncle up to 8 mm long;

bracts 3—4 mm long, pedicels c. 2 mm long,

bracteoles 2.5—3.5 mm long. Flowers: calyx

5,5—8 mm long, tube c. 3 mm long, upper lobe

shortly bifid at apex, 2.5—3.5 mm long, c. 1.5

mm wide at base, lateral lobes 3—3.2 mm long,

lower lobe acuminate, 4.5—5 mm long, allc. 1

mm wide; corolla white, occasionally with a

green area at the base of the standard; standard

orbicular, 9-10 mm long (7—)8—11 mm wide

on claw 1.5—2 mm long; wings (5.5—)9—11 mm

long, (2—)3.3—mm wide, auriculate at the base,

ona claw 1.5—2 mm long; keel slightly shorter

than wing petals. Pods with 2 or 3 articles;

articles oblong, 4-6 mm long, 3.5—4 mm wide

with sparse appressed hairs 0.1—0.2 mm long.

Seeds not seen.

Selected specimens: Papua New Guinea. Between

Dabora and Cape Vogel Peninsula, Apr 1953, Brass

21875 (K); vicinity of Rigo, Central District, Aug 1962,

Schodde 2699 (CANB, K, L); Mori R., c. 15 km NE of

Rodney, Central District, Aug 1969, Pullen 8132 (CANB,

K, L). Queensland. Coox District: 17 miles [27 km]

SW of Coen, Aug 1966, Story 7927 (CANB, K);

Chiliagoe, alt. 360m, Jan 1931, Hubbard & Winders 6794

(BRI, K); 4-mile Creek, Alma-den, alt. 450m, 17°208S

144°35°E, Jan 1972, Hyland 5818 (BRI, K, QRS).

Distribution and habitat: The species occurs

on limestone outcrops, in dry rainforest and in

eucalypt woodland often on margin of

rainforest from c. 18°S lat. through Cape York

Peninsula to Papua New Guinea in the vicinity

of Port Moresby and eastward. It has also been

recorded from Fiji (Ohashi 1997a). Map 12.

Affinities: Despite its position close to D.

multiflorum in the key to species, D. arbuscula

is related not to D. multiflorum but to D.

umbellatum and D, quinguepetalum (Blanco)

Merr. Knaap-van Meeuwen (1962) referred

four specimens (Brass 8789, Carr 11845, NGF

4225 from New Guinea and Aubbard &

Winders 6794 from Queensland) with some

doubt to Desmodium quinquepetalum.

Verdcourt (1979: 411) accepted the New

Guinean specimens cited by Knaap-van

Meeuwen as mere variants of Desmodium

umbellatum. Ohashi (1997a) pointed out

differences between D. arbuscula and

Dendrolobium quinquepetalum and referred all

four specimens to D. arbuscula. NGF 4228 ts

not a good specimen and difficult to identify

PPh ee Ce cert cert Cer tere ee aod ea ee a ce eo ae Pee eae ee heh ee ee ee eh ret nd

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 225

but I agree with Ohash1’s placement of the other

specimens.

D. arbuscula difters from D. guinguepetalum

in leaflets not acuminate with veins not

prominent beneath, both calyx and corolla

somewhat smaller, the lower lobe of the calyx

longer than the others by 1 mm or more, and

small, less pubescent articles of the pod. From

D. umbellatum it differs in its leaflets smaller,

more coriaceous and tendency to shine on the

upper surface, and, contrary to its protologue,

wing petals longer than the keel and about as

wide.

ThoughD, guinquepetalum has apparently

been excluded from the flora of New Guinea,

one specimen from the island (Laloki River,

June 1926, Brass 1654 (BRI, K)) probably

represents an undescribed species closely

related to it.

Desmodium Desy., J. Bot. Agric, 1:122 t.5

(1813), nom. conserv.; Benth. in Miq., Pl.

Jungh. 220 (1852); Hutchinson, Gen. FI.

Pl. 1:481 (1964); Ohashi, Ginkoana 1:87

(1973) — with synonyms listed, except

Holtzea Schindler which is

Dendrolobium, Smith, Fl. Viti. Nova 3:

188 (1985); Pedley, Rev. Handb. FI.

Ceylon 10:196 (1996). Type: Desmodium

scorpiurus (Sw.) Desv.

Note: Most other published descriptions of

Desmodium (for example, Schubert (1971),

Verdcourt (1979)) are quite wide and include

the genera Aphyllodium, Dendrolobium,

Phyllodium and Tadehagi which are treated

here as distinct.

A genus of some 300 species distributed mainly

through the subtropics and tropics with perhaps

half the species in Mesoamerica.

Bentham (1852, 1865b) is the only person

to have published an infrageneric classification.

He probably saw specimens of fewer than half

the currently accepted species and these,

because of a preponderance of specimens from

British colonies, mainly from the Old World.

Consequently his treatment of New World

species is rather sketchy. Schindler, whose

annotations of specimens and his publications

from 1911 to 1928 suggest that his knowledge

of the Desmodieae was unsurpassed, and who

had a narrow concept of Desmodium,

recognised the segregate genera of Bentham

(1852), and described others. Some were

recognised by Hutchinson (1964), Ohashi

(1973) and Ohashi et al. (1981). Ohashi (1973)

compared the different treatments.

Schindler never proposed a complete

infrageneric classification of Desmodium and

Meibomia which he treated as distinct genera,

though he did recognise several subgenera and

sections. Ohashi (1973) proposed a new

classification of Desmodium and allied genera.

His study was ‘intended to delimit all the

Asiatic species of the genus Desmodium and

its allied genera as well as these genera

themselves’. Since he considered less than a

fifth of the total number of species of

Desmodium over only part of its range, 1t might

be expected that his classification would have

only limited application. However, because

south-eastern Asia is the centre of generic

diversity (but not of speciation) of the

Desmodiineae and has few species in common

with the New World, Ohashi’s classification can

stand alone. Consideration of New World

species would probably lead to the recognition

of additional subgenera and sections rather than

the modification of those already proposed; that

is, a classification of American species could

be added to Ohashi’s classification with minor

disturbance.

Ohashi treated only 14 Australian

species, those that occur in Asia or Malesia as

well, and his classification requires minor

modification to accommodate all the Australian

species, The classification 1s set out below, Four

recently introduced American species, D.

incanum, D. intortum, D. tortuosum and D.

uncinatum have not been classified. Alterations

to Ohashi’s classification are:

1. Description of a new subgenus to

accommodate D. acanthocladunt.

Desmodium subg. Acanthocladum Pedley,

subg. nov. Frutex; ramuli in spinis

brevibus immutati; inflorescentia brevia

usque 4 nodis constructa; bracteoleae et

bractae primariae secundariaeque

226

praesentae; legumen usque 7 articulis,

aliquot plerumque abortis constructus,

unusquisque c. 10 mm longis, 4-4.5 mm

latis. Typus: Desmodium acanthocladum

F. Muell.

Shrub; branchlets modified into short spines;

inflorescence short with up to 4 nodes; primary

and secondary bracts and bracteoles present;

pod with up to 7 articles, some usually aborted,

each c. 10 mm long, 44.5 mm wide.

2. Transfer of Alpsicarpus sect.

Desmodiopsis Schindl. to Desmodium

(subg. Sagotia), with Desmodium

campylocauton F. Muell. ex Benth. the

only species

Desmodium sect. Desmodiopsis (Schindl.)

Pedley, comb. nov.

Alysicarpus sect. Desmodiopsis Schindl.,

Rep. sp. nov. reg. veg. 21:14 (1925).

Type: Desmodium campylocauton F.

Muell. ex Benth. (Ahysicarpus campylocaulos

(F. Muell. ex Benth.) Schindl.).

3. Recognition of (a) ser. Stenostachys and

(b) ser. Arillata, both sufficiently

distinct from the type of sect. Sagotia,

D. triflorum, to warrant acceptance

a. Desmodium ser. Stenostachys (Schindl.)

Pedley, stat. nov.

Desmodium sect. Stenostachys SchindL.,

Rep. sp. nov. reg. veg. 22:255 (1926).

Lectotype: Desmodium filiforme Zoll. &

Moritzi

Annual herbs; leaves 1- or 3-foliolate, often on

same plant; leaflets dimorphic; inflorescences

long, sparsiflorous, terminal and/or axillary;

flowers single, or rarely in pairs along the

rachis; secondary bracts and bracteoles absent;

pods with upper suture continuous or notched

between articles; seed with small rim aril only.

b. Desmodium ser. Arillata Pedley, ser. nov.

Herbae perennes; folia 1- et 3- foliolata,

rhachide saepe petiolo longiore;

inflorescentia terminalia, plerumque 3—

10-flora, compacta, saepe in frondibus

tecta; legumen suture supera undulata,

Austrobaileya 5(2): 209-261 (1999)

inferna crenata praeditum, ita inter

articulos constrictum; semen arillo bene

evoluto simili elei Codariocalycis

motorii (Houtt.) Ohashi ornatum. Typus:

Desmodium microphyllum (Thunb.) DC,

Perennial herbs; leaves |- and 3-foliolate,

rachis often longer than the petiole;

inflorescence terminal, usually 3—10-flowered,

compact, often hidden in foliage; upper suture

of pod undulate, lower crenate, hence pod

constricted between articles; seeds with well

developed aril stmilar to that of Codariocalyx

motorius (Houtt.) Ohashi.

4. Recognition of Sagotia at the rank of

series

Desmodium ser. Sagotia (Duchass. &

Walpers) Pedley, comb. et stat. nov.

Sagotia Duchass. & Walpers, Linnaea

23:737 (1850).

Perennial herb; leaves 3-foliolate; inflorescence

an axillary or usually a leaf-opposed 2—5

flowered fascicle, sometimes fascicles arranged

in short racemes; upper suture of pod

continuous; seed with small rim aril only. Type:

Desmodium triflorum (L.) DC. (Sagotia triflora

(L.) Duchass. & Walpers).

Systematic arrangement of Desmodiunt

Desv. in Australia.

subg. Hanslia (Schindl.) Ohashi

1. D. ormocarpoides DC

subg. Acanthocladum Pedley

2. D. acanthocladum F. Muell.

subg. Desmodium

3. D, scorpiurus (Sw.) Desv.

subg, Sagotia (Duchass. & Walpers) Baker

sect. Oxytes (Schindl.) Ohashi

4. D. brachypodum A, Gray

sect. Desmodiopsis (Schindl.) Pedley

5. D. campylocaulon F. Muell. ex Benth.

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 227

sect. Sagotia (Duchass. & Walpers.) Benth. 19. D. strigillosum Schindl.

ser. Arillata Pedley 20. D. nemorosum F. Muell. ex Benth.

6. D. microphyllum (Thunb. ) DC. 21. D. tiwiense Pedley

ser. Sagotia (Duchass. & Walpers) Pedley sect. Heteroloma Benth

7. D., triflorum (L.) DC. 22. D. velutinum (Willd.) DC.

8. D. heterophyllum (Willd.) DC. 23. D. gangeticum (L.) DC.

ser. Stenostachys (Schindl.) Pedley 24. D. rhytidophyllum F.Muell. ex Benth.

9. D. pycnotrichum Pedley 25. D. tenax Schindl.

10. D. trichostachyum Benth. 26. D. macrocarpum Domin

11. D. filiforme Zoll. & Moritzi 27. D. gunnii Benth. ex J.D. Hook.

12. D. brownti Schindl. 28. D. varians (Labill.). D. Don

13. D. flagellare Benth.

14. D. hannii Schindl. defined:

15. D. mueller? Benth. 29. D. tortuosum (Sw.) DC.

16. D. glareosum Pedley 30. D. uncinatum (Jacq.) DC.

17. D. pullenii Pedley 31. D. intortum (Mill.) Urban

sect. Nicolsonia (DC.) Benth. 32, D. incanum DC.

18. D. heterocarpon (L.) DC.

Key to species of Desmodium in Australia

1. Disk around base of ovary; stigma lateral; articles of pod 17-20 mm long;

SCAT S IGS ETT s eee oe uelatve, areas mvnslobeic reheat wari se tacts |dteerSl ee-g: Masgns tie nes 1. D. ormocarpoides

Disk absent; stigma terminal; articles of pod less than 12 mm long; herbs

GP SHU DS MOL SCATICEINE: tenes. 055-30 Phu tecaret yoo ye Reab tee Done a ERS fy oad adn isek aadu tery cneh ice

2. Woody shrub with spinescent branches; inflorescences axillary, up to 12

mm long with up to 4 fascicles of flowers; bracteoles present; pods of

4—6 articles, each 9-11 x 4-4.5 mm, some usually aborted....... 2. D. acanthocladum

Herbs or shrubs, not spinescent; inflorescences terminal or axillary,

usually longer; bracteoles absent; articles usually smaller, not

1H EIN ADOPT sony fasts boon 2c nce oy hs paren Shanes ae bck: ne ages aeek Es Siecseutcn doa ya hes WRinaen ose ae ees

3. Stipules persistent, distinctly auriculate at the base on the side further away

PEQTUE UNS HOCLTO LS: soya voters ebioas oct-as> eta Lo Re sree tra ota phan aay adbeast lone nese ss Nah ese: eh ote Ae

Stipules persistent or not, mostly symmetrical at the base, or if asymmetric

PNSINGTAUIICULALE, 66) dae aaa Kee Bed boa ao eM APE ice Bie ead Aba eed wae Ee gree ea

4, Erect annual; young pods twisted; mature articles with margins alternately

involute and revolute......... 0.0. ec ec cee eee eee teen enn 29, D. tortuosum

Introduced American species, position not

228 Austrobaileya 5(2): 209-26] (1999)

Sprawling or prostrate perennial herb; young pods not twisted,

mature articles with plane Marans: 5 caw eA GV ae Ee Moke Vad OV Na Soke aa woh aS 5

5. Leaves drying blue-black, sometimes with 5 leaflets; pod-articles

SEM Ciremlats Ae SANT eS Seemrce, ecdnbaeges Op & been aye Ge a-eceaie Hale lve woes 3.D. brachypodum

Leaves not drying blue-black, 3-foliolate; pod articles narrowly oblong

SLES A STITT, ay ot ov ode ae Ae, LF SVG, a Mca TE sey Ml elend Betoca 4. D. scorpiurus

6. Flowers large: calyx 6 mm or more long, lowest lobe c. 5 mm long; standard

c. lem long; pod shortly stipitate; stamens monadeiphous, the upper

stainen-Séparating as: POdS GEVElOP: 4. ss ck pe eke BEN Soc e Yea ew ne BP RG Ea ONE ES 7

Flowers smaller: calyx less than 6 mm long, lobes less than 5 mm long;

standard to c. 8 mm long; pod sessile; stamens diadelphous ............. 0.002 eee 8

7. Pod articles 5 x 3 mm; leaflets often with pale central zone along midrib

Lit etalk Moar ie hiy tus! + ack ty “eteiens tec ak yea 25d eS Oy a eT tod Sp SORE EE the Ms 30. D. uncinatum

Pod articles 4 x 2 mm; leaflets uniformly green above ...............4.. 31. D intortum

8. Pod articles inflated; leaves lanceolate to narrow ovate, 4-7 (—11) cm long,

4—7,5(—-10) times longer than wide ........... 00 cee eee ene §. D. campylocaulon

Pod articles flat; leaves smaller and/or less elongate... . 0.0... 0.0 ee eee 9

9. Upper calyx lobes connate at their base, united up to about half

their length, hence calyx subequally 5-lobed; inflorescences open;

pedicels often more than 5 mm long, filiform; secondary bracts absent;

PPATES SHIDO COOUES sy, oe tcecntez rb stc ots tne etd deme arian. afavuals Lanan Boar Misecetsarhe creak gate ae osels 10

Upper calyx lobe entire or bifid but not divided beyond the middle, hence,

calyx 4-lobed; inflorescence often dense; pedicels often less than 5 mm

long; secondary bracts often present; plants herbaceous or woody .............04. 21

10. Flowers in few-flowered axillary or leaf-opposed fascicles or 1n short

3—10-flowered terminal inflorescences, the inflorescences obscured by

TOMAS racic ce te dese Soha gd slat Mia Mcnae rcs Mea Sete ark Nowy wthelbre Meee aoree Realy Ave a Gleersecenlae tate wate ea Neale [1

Flowers single (rarely in pairs) in open terminal inflorescence projecting

PPO TO HAO 5 reget s-ch sake Whee Map Belcerh Le Sela Malar ta Paro aete betes ghd Uae w wendy 13

11. Inflorescence terminal; lateral leaflets 2.5—6 x 1.2—2.2 mm, usually 2—3

times longer than wide; both upper and lower sutures of pod incised;

seed-with well.Gevelopediarils so s.ce% 0 a6 $y ycale eae bmn eld a Bh 6. D. microphyllum

Flowers in axillary or leaf-opposed fascicles; lateral leaflets more than

3 mm wide and less than 3 times longer than wide; upper suture of pod

SrA PE BESG Wy WGlr MO AH OTT co ee oF eect ee neice eng bee ge nese eee Lae eset del ashen 12

12. Terminal leaflet cuneiform, 4—7.5 mm long; pedicels 5-8 mm long....... 7. D. triflorum

Terminal leaflet elliptic or obovate, 10-27 mm long; pedicels 10-20 mm

PONS Spt ar Ear a cade ann nek Ene toisyour ds hal aleve year ae 8. D. heterophyllum

13. Upper suture of pod distinctly notched between articles; nerves of articles

anastomosing, radiating from raised central area where anastomoses are

small and + equilateral; nerves often purplish ....... 0.0... ccc eee ee tees 14

Upper suture of pod continuous, not distinctly notched between articles;

nerves of articles + transverse, anastomoses + elongate not radiating

from central raised area; nerves not purplish ... 0... 0... ccc eee ees 17

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision | 229

14. Leaves |- and/or 3-foliolate; leaflets depressed ovate, orbicular or cuneiform ....... 15

Leaves usually 3-foliolate, terminal leaflet obovate, oblong or

PRS AE Pe ce ou be apo eliy 2% ved Ula Sek BIS Titi © UN ge os eRe wi et Aattad aADOT yb! erg; nod are la otalen? Bey dials certo? & 16

15. Leaves 3-foliolate; leaflets cuneiform; ovary and articles (at least when

young) with clavate trichomes 1... ccc eect eee ees 9, D. pycnotrichum

Leaves |- and/or 3-foliolate; leaflets cuneiform, depressed ovate or

orbicular; ovary and pod glabrous or with sparse uncinate hairs. 10. D. trichostachyum

16. Terminal leaflet oblong, 1O—35(—45) mm long, usually more than 3.5 mm

wide, 1—5(—7) times longer than wide; rachis of inflorescence usually

with long spreading hairs; plant prostrate... 2... eee 11. D. filiforme

Terminal leaflet linear, narrow lanceolate or narrow oblong, 10—50(—65)

mm long, usually less than 4 mm wide, 3.5—13 times longer than wide;

rachis (at least in upper part) without long spreading hairs; plant erect... 12. D. brownii

17. Pods with 2 (or 3) articles; articles 4.2—5 mm long; leaves usually 3-foliolate;

terminal leaflet oblong or obovate, 12-40 x 10-24 mm, 1|.1—1.7 times

LONG STRAWS os cna Be dyn nodes +: & oritebogho tie aunt Seedeet Ge ahve eller hunts Bhs ort 13. D. flagellare

Pods with 4—6 articles; articles 2.5-4.2(rarely —5 mm) long; leaves 1- or

3-foliolate; largest leaflet shorter, narrow or more elongate ........ 0.0.00. eee 18

18. Pod distinctly curved; articles (2.5—)2.8-4.2(—5) mm long; leaflets orbicular

(when 1-foliolate) or obovate or cuneiform (when 3-foliolate), usually

less than 1.5 times longer than wide ........0.. 0.0 ce eee eae ees 14. D. hannii

Pods straight; articles sometimes shorter; terminal (or single) leaflet oblong,

obovate-oblong, usually 1.5-4.5 or more times longer than wide ..............4.. 19

19. Articles (3.5—-)4—5 x 3-4 mm; largest leaflet of leaf widest below the middle,

glabrous, or almost so, on upper side 2.0... ee et eee 15. D. muelleri

Articles 2.5~3.5 x 2.2—3.3 mm; largest leaflet oblong or obovate,

not widest below the middle, usually pubescent on upper surface..............065 20

20. Hairs of upper surface of leaf usually weak, ascending; of stems usually

weak and ascending, white, up to 1 mm long; largest leaflet usually less

AAAS EE WEISS a oS acore tees ae er fon aie ayhdet S eca.ohle nor oh aeeee | 16. D. glareosum

Hairs of upper surface of leaflets appressed, occasionally somewhat

ascending; of stem stiff, yellow, curved ascending; largest leaflet usually

MMOLS THA SMT WIE Salas cea en Ae eee eh ee ee els che a Re re nee 17. D. pullenii

21. Leaves on individual plants predominantly 1-foliolate, occasionally with

afew S-1OlOlAIe 1EAV ES +n och pad le Ail does ad Be bs heath a Aten eta rene 22

Leaves on individual plants predominantly 3-foliolate, sometimes,

particularly on young plants, mixed with 1-foliolate leaves.............0 0.0.00 uae 25

22. Leaflet broadly ovate, 6—-12.5 x 4—-8.5(—10.5) cm, lower surface velutinous,

UPperSurlace PUBESCSNE oo ane ee tees Eke oh aw eh sees ee ow eee 22. D. velutinum

Leaflet oblong, elliptic, ovate or orbicular, 1.7—-8.5 (rarely —16) x

1-6(—8.5) cm, glabrous or almost so on upper surface, appressed

pubesconton. lower Surtace ot. Lore ans anata vista de own coe ate pea auicewle ane HE 23

23. Leaflet elliptic to ovate, acutish; secondary bracts present; pod deeply

incised; articles 2.5-3 x 2—2.5 mm; pedicels to4mmlong........ 23. D. gangeticum

Leaflet oblong, elliptic or orbicular, obtuse; secondary bracts absent; pods

230 Austrobaileya 5(2): 209-261 (1999)

shallowly incised, articles 3 mm or more long and wide; pedicels 2-3

PATS HODGE Padre Sao hed Gbcks feb ly seb lvash red Matec dosed prac Adeeb atria ine ol eee Lark lasers 24

24. Leaflet more or less orbicular, subcordate at base; pod-articles 3—3.5 x

aS OT EPVITDS cg ierhcinnatoene pF dice, aeioela hin b des arsn ele gs lag eee OA dla a eaeeeseeng ae 21. D. tiwiense

Leaflet oblong-elliptic,rounded at base; pod articles c.5 x 3.5mm .... 20. D. nemorosum

25. Primary bracts dense and entirely covering flower buds, the young

inflorescence therefore like coniferous strobiles; upper suture of pod

straight or slightly undulate, lower shallowly constricted, hence isthmus

usually more than half width of pod... . 0.0... ete tee eee eens 26

Primary bracts usually triangular or narrowly ovate, not entirely covering

flower buds; upper suture of pod undulate to distinctly incised, lower

suture deeply constricted, hence isthmus usually less than half width of

DOE od ha Rice taped aA arann Seddon rcshygiary yntaa ime Se dseee Nalin a as See eT EA ieee maT Sona Ask 28

26. Inflorescence open when mature; pedicels spreading or decurved in fruit;

podearticles: C,H Oe 2. dae eedioyepkcy a case yp lyse, acne aeace ace, busvcaes 20. D. nemorosum

Inflorescence dense when mature; pods erect, ascending or deflexed, often

obscuring rachis; pod articles 2.5—3 kK 1.8-3 mM........ 0. eee eee 27

27. Pods erect or ascending; terminal leaflet obovate, sometimes narrow; axis

of inflorescence either with rather spreading uncinate hairs or with stiff

yellowish appressed hairs ...... 0... 0c ccc eee eee eens 18. D. heterocarpon

Pods deflexed; terminal leaflet lanceolate to oblong; axis of inflorescence

shortly appressed pilose 0.0.0.0... ccc ccc ee eee teens 19. D. strigillosum

28.Plants rather densely pubescent; branchlets and at least undersides of

FSAVES WIEN BPEECAAINS AUS, 1 Lise te alge nay bee, 2 kG ARH en ln Wb EG BS 0 AGA PER Beet aR 29

Plants not densely pubescent; branchlets and undersides of leaves glabrous

OIMWIEDESPATSS AMPeESSEC IAT Sic fos a5, sce dil Sedots eS LY poe ach le cated once bore HL ic as 32

29. Stipules 3—11mm long, united at base, at least when young............. 32. D. incanum

SELES: TOGO. aT LOTS Ie. oot aca let ecevipn Salk enka wie ug eile DAE AMG departs 30

30. Terminal leaflet to 60 mm long; articles of pod 3.5—4 x 2.5—3 mm; trailing

OP SALLIE Ae a a an a 8 setecry ein Wyse 'e ows x ca I oe a (SP ae etka 24, D. rhytidophyllum

Terminal leaflet more than 60 mm long; articles of fruit more than 5 mm

LOVE AGRO CCIDPATIS 4, Bek hte uc kySete it San rinrceg dee gla eis. son teats ston eh casary ser ecent-aateMargeed verse 31

31.Leaflets ovate with long acute apex; upper surface with sparse

appressed to ascending hairs; articles 5.5—8 x 3.4-4mm ............... 25. D. tenax

Leaflets ovate or rhombic, obtuse, mucronulate, with moderately dense

stiff hairs on both surfaces; articles 9-10 x 5-6 mm ............ 26. D. macrocarpum

32, Leaves subdigitate; leaflets cuneiform and truncate at the apex or rarely

broadly obovate and rounded; pod articles 4—5 x 2-2.5mm............ 27. D. gunnii

Leaves usually with distinct rachis 24.5 mm long; leaflets lanceolate,

oblong or rarely oblanceolate; pod articles 3.5-4.2 x 2.7-3.8mm...... 28. D. varians

OPE A NE Sor et Eee eon Tee ETE a LEE UCE TS tte ete DES ty dng dec hhcvd alive antdece chee aa

* epee ph Te Lar Loe a nnn ean eta ng rae A RR na CE PII Be SOE PP EN een Ef PEE ERE SEEN FET SS STS DEE RESETS PRE PL LY,

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 231

1. Desmodium ormocarpoides DC., Prodr.

2:327 (1825); Desvaux, Mém. Soc. Linn.,

Paris 4:307 (1825); Ohashi, Ginkgoana

1:113 (1973); Verdcourt, Man. New

Guinea Leg: 403 (1979). Based on

Hedysarum adhaerens Poir., in Lamarck,

Encycl. Meth. Suppl 5:15 (1817), nom

illeg., non Vahl (1791); Hanslia

adhaerens (Poir.) Schindl., Rep. sp. nov.

reg. veg. 20:277 (1924) Type: ‘Cette

plante a été découverte a |’fle de Java par

M. de Labillardiere’ (P?, n.v.)

Desmodium dependens Blume ex Migq,, FI.

Ind, Bat. 1:248 (1855). Type:

Molukscheeilanden. Nieuw Guinea’ (n.v.)

Selected specitinens: Queensland. Cook DIsTRIcT:

Alligator Creek Catchment, 12°35’°S 143°20°E, Oct 1972,

Hyland 6440 (BRI, K); Shiptons Flat, 15°48’°S 145°14’E,

Oct 1973, Webb & Tracey 13643 (BRI, CANB). Norts

KENNEDY District: Rockingham Bay, Dallachy (MEL);

Port Denison, in 1874, Fitzalan (MEL).

Distribution and habitat: The species occurs

in rainforest, usually in disturbed situations

such as along logging tracks. It ranges from

the central Queensland coast through New

Guinea to central Malesia. It is not common in

Australia. Map 13.

Affinities: D. ormocarpoides ts a well defined

species isolated in the genus. Schindler referred

it to his monospecific genus Hanslia which

Ohashi reduced to subgeneric rank within

Desmodium. It is characterised by its stipitate

pods composed of slightly turgid, narrowly

obovate articles, the minute tubular disk around

the base of the ovary, the lateral stigma and the

bracteoles at the base of the calyx.

2. Desmodium acanthocladum F. Muell.,

Fragm. Phytog. Austr. 2:122 (1861);

Benth., Fl. Austr. 2:23 (1864); C. Muell.

in Walpers, Ann. Bot. 7:764 (1870);

Steen. & Veldk., Reinwardtia 10:24

(1982); Dicerma acanthocladum F.

Muell., loc. cit., pro. syn; Meibomia

acanthoclada (F. Muell.) Kuntze; Rev.

Gen. 1:197 (1891). Type: Clarence River,

Beckler (holo: MEL; iso: K).

Woody shrub to 2 m tall; branchlets Gnany

modified into short thorns) angular with

yellowish ribs running from the bases of

stipules, glabrous; stipules deltoid, 1-2 mm

long, veined. Leaves trifoliolate, coriaceous;

petioles sulcate, 1-4 mm long; rachis 0.6—2.3

min long; petiolules 0.3—0.5 mm long; stipels

setaceous, about as long as the petiolules;

terminal leaflet oblanceolate and obtuse or

narrowly elliptic and acute, the laterals oblong

or obovate, acute or obtuse, all glabrous and

slightly shining above or with scattered short

hairs and paler beneath, the terminal ones 12—

28 mm long, 3—-5.5 mm wide, 3—6 times longer

than wide; laterals smaller, 7—-18.5 mm long,

3—5 mm wide, 23.6 times longer than wide.

Flowers pink or mauve in fascicles of 2 or 3

along an axillary axis, up to 4-noded and 12

mm long; primary bracts c. | mm long,

secondary bracts similar, shorter; pedicels 2—3

mm long with bracteoles 0.6—1 mm long at top

of pedicel; calyx campanulate, 4-lobed, 3-4

mm long with scattered appressed hairs, the

upper lip notched at the tip, 1.8~2 mm long,

the lateral and lower ones apiculate with

thickened tips, 1~1.3 mm long. Corolla longer

than the calyx: keel orbicular, shortly clawed,

6.5—7 mm long, 6-7 mm wide; wings oblong,

obtuse 7-8 mm long, 2.3-3 mm wide,

auriculate at the base, on a claw 1.5—2 mm long,

keel petals 6.5~8 mm long, 2.3—-3 mm wide,

auriculate on claw 2—2.5 mm long; stamens

diadelphous; ovary minutely uncinate-hairy.

Pods with 2-6 (7) articles, some usually aborted

and occasionally appearing stipitate when the

proximal article is aborted, upper suture

_ Shallowly indented, the lower margin deeply

indented, the isthmus c. '/; width of pod. Articles

+ reticulately longitudinally veined, with

moderate indumentum of hooked hairs, 9-11

mm long, 4-4.5 mm wide; seeds obliquely

oblong, c. 6 mm long, 3.5 mm wide, rim aril small.

Selected specimens: New South Wales, NortH Coast:

Boat Harbour Flora Reserve, 17 km NE of Lismore

railway station, 28°47’°S 153°30’E, Feb 1989, Coveny

10651 (K, NSW); Wilsons River at Lismore, 28°48’S

153°17'%4’E, Feb 1987, Pullen 11139 (BRI, K, NSW).

Distribution and habitat: The species is

confined to the North Coast division of New

South Wales where it occurs on the fringes of

riverine rainforest. It was probably more widely

distributed before extensive clearing of the

land, but apparently viable populations occur

in flora reserves. Map 14.

232

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Austrobaileya 5(2):; 209-261 (1999)

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14, Desmodium ancanthocladum 15. Desmodium scorpiurus

CONTR ey ri

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 233

Affinities: The relationships of the species are

uncertain. Bentham (1864) dismissed Mueller’s

association of 1t with Desmodium biarticulatum

(L.) F. Muell. (= Aphyllodium biarticulatum)

and stated ‘itis otherwise more nearly allied to

the section Heteroloma subsect. Podocarpa’ .

Ohashi (1973) referred most species of subsect.

Podocarpa to subg. Podocarpium (Benth.)

Ohashi. D. acanthocladum differs from

species of subg. Podocarpium in its woody

habit, thorny branchlets, short axillary

inflorescences, bracteoles and diadelphous

stamens. This combination of characters

excludes it from other subgenera treated by

Ohashi, and it has therefore been placed in the

newly described, monotypic. subg.

Acanthocladum (see p. 225).

Its resemblance to Trifidocanthus

unifoliolatus Merr.( Desmodium unifoliolatum

(Merr.) Steen.) is due mainly to its ‘eco-

taxonomical character’ (Steenis & Veldkamp,

1982), its thorns. Otherwise it is not related to

the other species.

3. Desmodium scorpiurus (Sw.) Desv., J. Bot.

Agric. 1:122 (1813); Schubert, in

McBride, Field Mus. Nat. Hist. Publ. Ser.

13 (3,1):433 (1943); Verdcourt, Man.

New Guinea Leg: 407 1.94 (g) (1979);

Howard, Fl. Lesser Antill., Leeward &

Windward Is. 4(1):482 (1988); Smith FI.

Vitiensis Nova 3:190 (1985); Hedysarum

scorpiurus Swartz, Nov. Gen. & Sp.

Prodr.:107 (1788). Type: Jamaica,

Swartz (S, fide Howard, n.v.)

Selected specimens: Western Australia. Port Hedland,

Runich s.n. (PERTH). Northern Territory, Darwin, Oct

1971, Pickering sn. (DNA, K). Queensland. Cook

District: Thursday Is., Oct 1981, Clarkson 3817 (BRI,

DNA, CANB, K); 8 of Port Douglas, 16°30’°S 145°27’E,

Feb 1974, Gibbs sn. (CANB). Hyde Park, Townsville,

Feb 1990, Jobson 1084, (AD, BRI, CANB, HO, MEL).

Distribution and habitat: The species 1s native

to tropical America, the West Indies and

Mexico southward to Peru. It was introduced

into Australia as a potential fodder plant and is

now naturalised in a few places in the tropical

part of the country. It favours open situations

on well drained soils, particularly behind

coastal dunes. Map 15.

4. Desmodium brachypodum A. Gray in U.S.

Expl. Exped. 434 (1854); Benth., FL.

Austr, 2:232 (1864); Ohashi, Ginkgoana

1:227 (1973); Verdcourt, Man. New

Guinea Leg.:394 (1979). Type: Hunter

River, U.S. Explor. Exped. (n.v.)

D. indigotinum Harms. & K. Schum. in K.

Schum. & Laut, Nachr. Fl. Deutsch,

Schutz geb. Stidsee Nachtr.: 276 (1905).

Type: Kaiser Wilhelmsland: Finschhafen

Wieland s.n. (n.v.)

Selected specimens: Northern Territory. Nimbuwa

Rock, i2°11’S 133°2i°E, Jun 1974. Pullen 9502

(CANB). Queensland. LEICHHARDT District: West of

Moura, 24°34’8 149°39E, Apr 1961, Jones (CANB);

Burnett District: Monto, 24°52’S 151°O7’E, Jun 1937,

Regan (BRI); DarLinc Downs District: Canal Creek, in

1880, Hartmann 712 (MEL). New South Wales. 3 miles

[5 km] from Tent Hill on Torrington road, Nov 1973,

Rodd 2540 (NSW); Albion Park, Apr 1968, McBarron

4617 (NSW); Mt Kaputar National Park, 31 km ENE of

Narrabri, Nov 1967, Coveny 8916 & Roy (K, NSW).

Victoria. East Gippsland, Little River track, Mar 1971,

Beauglehole ACB 37153 (MEL).

Distribution and habitat: Vhe species is widely

spread in eastern Australia from Gippsland,

Victoria through coastal and subcoastal parts

of New South Wales and Queensland to Cape

York Peninsula, often in hilly country on sandy

or loamy soils in eucalypt communities. It

extends to the Northern Territory (two

specimens seen) and eastern New Guinea.

Desmodium brachypodon,D. rhytidophyllum

and D. varians are the most frequently

encountered species of Desmodium in eastern

Australia. Map 16.

Affinities: Schindler (1926) described

Desmodium subg. Oxytes to accommodate D.

brachypodum and two New Caledonian

species, D. deplanchei Harms and D.

pycnostachyum Benth. Ohashi (1973) reduced

this to sectional rank in subg. Sagotia with D.

brachypodum as lectotype.

5. Desmodium campylocaulon F. Muelil. ex

Benth., Fl. Austr. 2:233 (1864);

Meibomia campylocauton (F. Muell. ex

Benth.) Kuntze, Rev. Gen. 1:197 (1891)

(campylocaulis’); Afysicarpus campylocaulos

(F. Muell.) Schindl., Rep. sp. nov. reg. veg.

21:14 (1925). Type: Sturts Creek, March

1856, & Mueller (holo:s),

234

Prostrate or somewhat ascending perennial

herb; branches with indumentum of short erect

uncinate hairs when young, becoming sparse

or absent when old; stipules rather stout,

triangular, striate, 3—8 mm long, 1.5~-3 mm wide

at base. Leaves 3-foliolate; petioles (20—)30—

70 mm long, rachis 3-11 mm long; leaflets

lanceolate to (rarely) ovate, acute or obtuse and

mucronulate at the apex, attenuate at the base,

glabrous above with sparse hairs usually on

veins beneath; terminal leaflet 40-—-70(—110)

mm long, 6.5—15 mm wide, 3—7(—10.5) times

longer than wide, the laterals shorter, 30—60(—

100) mm long. Inflorescences terminal or

rarely leaf-opposed, to 20 cm long, rachis with

indumentum of spreading uncinate hairs c. 0.2

mm long, dense, becoming sparse as

inflorescence elongates; flowers in pairs

subtended by persistent oval-acuminate

glabrous bract, 3-4 mm long; secondary bract

and bracteoles absent; pedicels 2—4(rarely 5)

mim long at anthesis, doubling in length in fruit.

Flowers: calyx 4-merous, 2.5-3.5 mm long,

with scattered straight hairs; tube 1-1.5 mm

long, lobes 1.5—2 mm long, the upper bifid;

corolla pink to pale purple; standard obovate,

3.5 mm long, 2~3 mm wide; wings 3.8—-4 mm

long, 1 mm wide; keel petals 4.3—4.8 mm long,

1-1.5 mm wide; stamens diadelphous; ovary

pubescent. Pod straight, with upper suture _

thickened, the lower deeply indented; with 5—

7 articles, each inflated, indehiscent, (2.5—)3—

4 mm long, 2.2—-3.3 mm wide, reticulately

veined, with spreading uncinate hairs. Seeds

oblongoid, 2.2—2.8 mm long, 1.5—2 mm wide,

c. 1 mm thick, the hilum ecentrically placed on

the long side, rim aril small.

Selected specimens: Western Australia. 40 km SE of

Fitzroy Crossing, 18°28’S 125°45°E, May 1988, Pullen

11197 (CANB). Northern Territory. 30 miles [48 km]

E of ‘Alexandria’ Station, Jun 1948, Perry 1507 (BRI,

CANB, K); 25% [41 km] W of ‘Rockhampton Downs’,

Jun 1960, Chippendale NT 7118 (DNA, K, MEL)

Queensland. SouTH KENNEDY District: 2 miles [3 km]

NE of ‘Natal Downs’ Station, 21°04’S 146°08’E, Jul

1964, Adams 1147 (BRI, CANB, K); WarreGo DisTRIcT:

Gilruth Plains, Cunnamuila, 28°04’S 145°41° E, Apr

1963, AdcKee 10311 (BRI, CANB, K).

Distribution and habitat: Desmodiunt

campylocaulon extends from the northern part

of Western Australia through the north-central

part of the Northern Territory into inland

southern Queensland. It is confined to clay

Austrobaileya 5(2): 209-261 (1999)

soils, usually in grassland with Astrebla spp.

and Iseilema spp. Map 17.

Affinities: As sole member of sect.

Desmodiopsis (see above, p.226) D.

campylocaulon has no close relative, though

in habit, general appearance and ecology it

resembles D. muelleri (sect. Sagofia). In his

description of D. campylocaulon, Bentham

described the pod-articles as ‘membranous

turgid or almost inflated when ripe’. Such a

description is misleading. The pods are, in fact,

markedly inflated and unlike the pods of any

other Desmodieae. Presumably because of

these inflated articles, Schindler transferred the

species to Alysicarpus, describing a new

section Desmodiopsis to accommodate it.

Alysicarpus has turgid pods which are

symmetrical: that is, their upper and lower

margins are equally indented. They are also less

membranous. The calyx of Alysicarpus 1s

scarious with complex venation, unlike that of

D. campylocaulon, As generally understood

Alysicarpus is an easily recognised, nicely

circumscribed, homogeneous genus. The

addition of D. campylocaulon to it, even tn a

distinct section, obscures the present clear

generic limits without doing anything to clarify

the already rather indistinct limits of

Desmodium, Prain (1897), commenting on

Baker’s (1876) transfer of Alysicarpus

parviflorus Dalzell and other species to

Desmodium, and concerned that it would not

fit well in that genus, stated: ‘Compromises in

taxonomy are necessary, indeed the systematic

arrangement of the species is essentially the art

of happy compromise, but an arrangement

which places one half of a natural group of

forms in one genus, the other half in a second,

strains unduly the privileges that the art of

compromise allows’. Prain’s principle still

applies. Alysicarpus parviflorus and related

trifoliolate species are now referred to

Desmondiastrum A, Pramanik & Thoth.

(Pramanik & Thothathri 1986). Rather than to

complicate Alysicarpus which, despite

comments to the contrary by Prain (op. cit.)

and Meeuwen (1962), is quite distinct from

Desmodium, I have treated D. campylocautlon

as the only member of a section of Desmodium.

No other genus of Desmodieae has inflated pod

articles. The pods of Pycnospora, which 1s

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 235

conventionally, though possibly wrongly,

placed in the tribe are inflated but are 7-10

seeded, not segmented, and are not analogous

to those of D. campylocauton.

6. Desmodium microphylium (Thunb. ex

Murray) DC., Prodr. 2:337 (1825); van

Mecuwen, Reinwardtia 6:254 (1962);

Ohashi, Ginkgoana 1:241 (1973) & FI.

Camb. Laos Vietn. 27:105 (1994);

Verdcourt, Man. New Guinea Leg. 402

(1979); Pedley, Rev. Handb. Fl. Ceylon

10:176 (1996); Meibomia microphylla

(Thunb.) Kunze, Rev. Gen. 1:198 (1891);

Hedysarum microphyllum Thunb. ex

Murray, Syst. Veg. ed. 14:675 (1785), FI.

Jap. 284 (1784); Poir. in Lam, Encycl.

Method. Bot. 6:417 (1804). Type: from

Japan (n.v.)

Desmodium parvifolium DC., Ann. Sct.

Nat. 4:100 (1825) & Prodr. 2:234 (1825);

Wright & Arn., Prodr. FI. Ind. Orient. 229

(1834); Benth., Fl. Austral. 2: (1864).

Type: Nepal, in 1821, Wallich 5700

(iso:K).

Selected specittnens: Queensland. NORTH KENNEDY

District: Rockingham Bay, Dallachy (MEL, NSW); Porr

Curtis District: Port Clinton, 22°38’S 150°42’°E,

Clarkson & Stanley 855 (BRI, CANB); Moreton

District: Mor(e}ton Bay, Mueller Us, MEL). New South

Wales. Richmond River, Moore (K).

Distribution and habitat: Desmodium

microphyllum 1s not a common plant in

Australia. It has been collected in coastal areas

from about Cairns south to the Richmond River,

but little 1s known about its ecological

requirements. It does occur in eucalypt open-

forests and is evidently native. The species has

a huge latitudinal range, extending through

Malesia to south-eastern Asia and China and

Japan. Specimens collected near the type

locality, Nagasaki (Maximowicz in 1863,

Oldham in 1862, K) leave no doubt that the

Australian and the east Asian plants represent

the same species. Map 18.

Affinities: Bentham (1852) placed D.

microphyllum (as D. parvifolium) in sect.

Sagotia withD. triflorum and D. heterophyllum,

and it does have affinities with these species.

However, its pod incised along the upper suture

(not notched between articles) and, more

importantly, the well developed aril set it apart

from D. triflorum, D. heterophyllum and the

other species I have referred to sect. Sagotia. I

have therefore treated it as the single species

of ser. Arillata. The well developed aril

suggests a relationship with Codariocalyx, but

other attributes do not support such a

relationship.

7. Desmodium triflorum (L.) DC., Prodr.

2:334 (1825); Liu & Chang, Taiwania

8:96 t.22 (1962); Schubert, J. Arnold Arb.

44; 293 (1963) & FI. Trop. East Africa:

Legum. Papilion. 459 (1971); Ohashi,

Ginkgoana 1:245 (1973) & Fl. Camb.

Laos Vietn. 27:11 (1994); Verdcourt,

Kirkia 9:512 (1974) & Man. New Guinea

Leg. 409 (1979); Du Puy, Fl. Austral.

50:314 (1993); Pediey, Rev. Handb. FI.

Ceylon 10:178 (1996); Sagotia

triflora(L.) Duchass. & Walpers, Linnaea

23:738 (1850); Meibomia triflora (L.)

Kuntze, Rev. Gen. 1:197 (1891) (incl.

vars.); Hedysarum triflorum L., Sp. PL.

749 (1753). Type: Herb. Hermann vol. |

fol.21, No.297 (BM, lectotype fide

Pedley in Turland & Jarvis (1997).

Selected specimens: Northern’ Territory.

Palmerston,12°29’°8 130°59’E, May 1992, Cowie 3017

(BRI, DNA). Queensland. Cook District: Mareeba, Apr

1983, Clarkson 4587 (BRI, K); NorTH KENNEDY DIsTRICT:

Tully & vicinity, Jan 1950, Clemens (ex herb. Univ.

Michigan) (BRI, K); Wine Bay District: Bundaberg, Apr

1936, Blake 11253 (BRI, CANB, K, MEXU); MoreTon

District: Bribie Is., May 1930, Hubbard 2667 (BRI,K),

Distribution and habitat: This species is

distributed through the tropics and subtropics.

It occurs in open spaces, on roadsides and is

often a weed of lawns, usually on well drained

soils. In Australia it occurs in the extreme north

of the Northern Territory and along the east

coast north from about Brisbane. It is so well

established in south-eastern Queensland that it

is hardly conceivable that it does not extend

into New South Wales, but I have seen no

specimens. Map 19.

Affinities: Desmodium triflorum and D.

heterophyllum are closely related. The latter

has larger, more elongate leaflets and the upper

calyx lobes are united to the middle. The two

species constitute ser. Sagotia of sect. Sagotia

and therefore stand somewhat apart from other

species of the section.

236

8. Desmodium heterophylium (Willd.) DC.,

Prodr. 2:334 (1825); White & Francis,

Qld Dept. Ag. & Stock Bot. Bull. 22:14

(1920); Liu & Chang, Taiwania 8:81 t.10

(1962); Ohashi, Ginkgoana 1:239 (1973)

& Fl. Camb. Laos Vietn. 27:110 (1994);

Verdcourt, Man. New Guinea, Leg. 400

(1979); Pediey, Rev. Handb. FI. Ceylon

10:179 (1996); Meibomia heterophylla

(Willd.) 10:179 Kuntze, Rev. Gen. 1:196

(1891); Hedysarum heterophyllum

Willd., Sp. Pl. 3:1201 (1802); Type:

Herb. Willdenow 13832 (B, n.v,;

microfiche: BRI)

Hedysarum triflorum vay. B &y L., Sp. PL.

749 (1753).

Selected specittnens: Queensland. Coox District: Bolt

Head, Temple Bay, 12°15’S 143°05’E, Jul 1991, Forster

PIF 8988 (BRI); Cairns, Feb 1918, White [AQ 19947]

(BRI); South Johnstone, Apr 1952, Bureau of Tropical

Agriculture [AQ 97399], possibly cultivated (BRI); Sour

KENNEDY District: Slade Point, 21°05’S 149°13’E, Aug

1992, Batianoff 920853 & Champion (BRI).

Distribution and habitat: Desmodium

heterophyllum is a comparatively recent

introduction to Australia and is only sparingly

naturalised and poorly collected. Little is

known of its ecology though it 1s found on sand

behind dunes on beaches immediately north of

Cairns. Map 20.

Affinities: The species is closely related to D.

triflorum though not likely to be confused with

it. Ohashi (1973) described the upper two lobes

of the calyx to be ‘deeply incised near base’;

they are in fact united to about the middle as

can be seen in his illustration (t. 53:3).

9, Desmodium pycnotrichum Pedley, sp. nov.

leguminibus sutura supera inter articulos

acute Incisa praeditis similis eis D.

filiformis Zoll. et Moritzi et D.

trichostachyi Benth. autem ab ambabus

ovario et plerumque aliquis leguminis

trichomatibus clavatis ornatis differt. Id

foliolum terminale longum D. filiformis

caret. Typus: Northern Territory: Coastal

Plains Research Station [12°34’S

131°19’E], Mar 1963, M Lazarides 6828

(holo: CANB; iso: BRI, K, MEL).

Annual herb often flowering and fruiting when

+ erect, but becoming prostrate; branches with

Austrobaileya 5(2): 209-261 (1999)

moderate to dense, stiff, yellowish, spreading

hairs 0.8—1.2 mm long. Leaves 3-foltolate,

occasionally 1-foliolate; stipules 3—7 mm long,

hairy at the base or with marginal hairs; petioles

3.5—21(—24) mm long; rachis 2.5—-6.5(—-10) mm

long; leaflets cuneiform, obovate or sometimes

oblong, attenuate at the base, retuse or truncate

at the apex, usually glabrous, or occasionally

with sparse appressed hairs on the upper

surface, sparse appressed or occasionally

loosely ascending hairs 0.6 mm long on lower

surface; terminal leaflet 8.5—30 mm long, (8—)

10-—20(-22) mm wide, 0.8—-1.5 times longer

than wide, laterals smaller, 7—19(—24) mm long,

5—14(—17) mm wide, 0.9—1.6 times longer than

wide; pulvinuses 0.8—1.6 mm long; stipels | .5—

2 mm long. Inflorescences terminal, open, to

25 cm long, rachis with sparse to rather dense

spreading hairs, rarely appressed, to 1 mm long;

flowers single on the rachis; primary bract

acuminate, 2.5~5 mm long, pubescent;

secondary bract and bracteoles absent; pedicels

slender, 5—9 mm long. Flowers: calyx 2—3 mm

long with linear lobes, the upper bifid to the

middle, the tube %<—'/; total length, long hairs

on lobes and short sparse uncinate hairs on

tube; corolla pink, occasionally red or rarely

bluish with standard obovate, 3—3.5 mm long,

1.5—-2.5 mm wide, wings c. 2.5 mm long, 0.7—

I mm wide, on short claw, keel petals slightly

longer, tapered to the base, not clawed; stamens

diadelphous; ovary with dense covering of

clavate hairs. Pods + straight, the upper suture

notched between the lobes, the lower deeply

indented between them, the isthmus about half

width of pod, with 3 or 4 articles, each 2—3.5(—

4) mm long, 2—3(-4) mm wide, with

indumentum of clavate hairs, rarely some

glabrous, nervature similar to that of D.

trichostachyum; seeds reniform, black when

mature, 1.7—3(—3.6) mm long, (1.2—)1.4-2 mm

wide with small rim aril.

Selected specimens: Western Australia. Manning Creek

Gorge, near ‘Mt Barnett’ Station, 16°39’S 125°55’E, May

1988, Pullen 11211 (BRI, CANB, DNA, PERTH).

Northern Territory. Tortilla Flats, 13°06’S 131°14’E,

Apr 1974, Parker 411 (DNA, CANB); Woolaning,

13°07°S 130°40°E, Apr 1981, Dunlop 5879 & Craven

(BRI, CANB, DNA, MEL); 12 km S of Hayes Creek,

13°35’S 131°30’E, Apr 1988, Pullen & Spottswood

11165 (BRI, CANB, DNA); Douglas Hot Springs

National Park, 13°45’S 13°126’E, Apr 1988, Pullen &

Spottswood 11159 (BRI, CANB, DNA, K, L, PERTH);

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 237

Smith Creek, Coburg Penin., 11°07°S 132°08°E, Apr

1977, Pullen 10619 (CANB, DNA),

Distribution and habitat: The species usually

occurs on sandy soils, sometimes in disturbed

situation, sometimes associated withD. pullenti

(Pullen & Spottswood 11158 is a mixed

collection). It is confined to the Dieman and

Gulf Region of the Northern Territory and the

Kimberley Region of Western Australia. Pullen

(in sched., Pullen 11211) suggests that it may

be a recent introduction to Western Australia,

but it was collected by Bradshaw and Allen on

the Prince Regent River in 1891 (specimens,

MEL). Map 21.

Affinities: Desmodium pycnotrichum has pods

like those of D. filiforme and D. trichostachyum

and is probably closely related to the latter, It

differs from both in the unusual clavate

trichomes on the ovary which usually persist

on the pod. Its foliage resembles that of D.

trichostachyum and lacks the long terminal

leaflet of D. filiforie. However, one specimen

(Reeve & Watson 228, DNA) with elongated

terminal leaflets (34.5 times longer than wide}

possibly represents a D. filiforme x D. pycnotrichum

hybrid.

Etymology: The epithet is a Latinisation of

Greek: pychno, thick, and thrix, trichos, hair,

a reference to the peculiar indumentum of ovary

and, usually, pods. Specimens have been

distributed with the name D. ‘clavitricha’, a

name coined by Mr R. Pullen who recognised

the species as distinct. Regrettably the epithet

is neither grammatically nor etymologically

acceptable and has therefore been replaced.

10. Desmodium trichostachyum Benth., FI.

Austr. 2:234 (1864); Ohashi, Ginkgoana

1:244 (1973). Meibomia trichostachya

(Benth.) Kuntze, Rev. Gen. 1:198 (1891).

Type: Port Essington, Armstrong (lecto:

Kk, chosen here),

Prostrate or slightly ascending annual herb;

stems slender, terete, glabrous or with sparse

appressed or occasionally spreading hairs (0.6

min long). Leaves |- or 3-foliolate, sometimes

on separate plants, sometimes leaves of both

types on one plant, in which case the |-foliolate

produced before the 3-foliolate ones; stipules

ovate or deltoid, acute, 2.5—4 mm long, 1—-1.5

mm wide; petiole 3-12 mm long, rachis 1-4

mm long; leaflets depressed ovate, orbicular

or occasionally broadly oblong, rounded or

subcordate at the base, retuse at the apex when

1-foliolate, cuneiform, rounded or subcordate

at the base, truncate or retuse at the apex when

3-foliolate, all + glabrous or with sparse

appressed hairs on lower surface; when 1-

foliolate, leaflet 8-20 mm long, 9-20 mm wide,

Q.7—1.2 (-1.4) times wider than long; when 3-

foliolate terminal leaflet 5-15 mm long, 6-16

mim wide, i—1.4 times wider than long, lateral

leaflets smaller, 4-12 mm long and wide,

(0.7)1—1.2 times longer than wide; pulvinuses

Q.5-1.5 mm long; stipels 0.21 mm long.

Inflorescences terminal, very open, to 25 cm

long, the rachis with a few appressed or rarely

spreading hairs towards the apex, flowers single

(rarely in pairs) subtending by an ovate

acuminate bract 1~2 mm long, secondary bracts

and bracteoles absent; pedicels slender, 2-6

mim long, doubling in length in fruit. Flowers:

calyx 1.4—2.]1 mm long, uncinately pubescent

or occasionally lobes strigose with hairs to |

mm long; tube 0.5—0.7 mm long, upper lobe

0.8—1.4 mm long, incised to about the middle,

other lobes shorter or about equal in length,

0.6-1.3 mm long; corolla purplish: standard

obovate, obtuse or retuse at apex, narrowed to

base but not distinctly clawed, 2.5—3.3 mm long

(1.6-)2—2.5 mm wide, wings broad oblong with

short claw, 1.8—-2.5 mm long, 0.7—1.2 mm wide,

keel petals somewhat longer and narrower than

wings, 2.2—-2.9 mm long, 0.7-1 mm wide,

tapered to base, not distinctly clawed. Pods with

upper suture thickened, notched between

articles, lower indented, isthmus “2—/ width of

pod, with 3—5 articles, each 2—2.8 mm long and

wide, glabrous or with uncinate hairs,

transversely reticulately veined; seed 1.3~1.7

mim long, 0.9—1.5 mm wide, with a rim aril.

Selected specinens: New Guinea. MERAUKE: Koerike

Camp, c.15 km NE of Koembe village on Koembe river,

Sep 1954, P. van Royen 4910 (L). Western Australia.

King Edward River Crossing, May 1975, Symon 10250

(NSW); Augustus Is., 15°25’°S 124°33’E, May 1972,

Wilson 10719b & 10877 (PERTH). Northern Territory.

Humpty Doo, Feb 1961, AdcKee 8356 (CANB, K, L);

Arnhem Land, 12°55’S 135°17’E, Jun 1972, Maconochie

1498 (DNA, K); Elcho Is., Jul 1975, 11°47’S 135°53’E,

Maconochie 2223 (DNA). Queensland. Cook District:

Newcastle Bay [10°49’S 142°36’°E], May 1948, Brass

238

18740 (BRI, CANB, K, L); 2 km S of Moreton Telegraph

Station, 12°28’S 142°38’E, Kanis 2037 (CANB, L);

Endeavour River, Jun 1819, Cunningham 243 (K); Gorge

Creek, 10 miles [16 km] W of Mareeba, Apr 1962, McKee

9265 (BRI, CANB, K, L).

Distribution and habitat: Desmodium

frichostachyum occurs on sandy soils,

sometimes seasonally waterlogged, in eucalypt

and Melaleuca communities in coastal areas

from the Kimberley region of Western

Australia, through the Northern Territory to

northern Queensland south to about the

Atherton Tableland. It also occurs in southern

New Guinea. Map 22.

Affinities: Desmodium trichostachyum, D.

pycnotrichum, D. filiforme and D. brownti

constitute a complex of species of ser.

Stenostachys characterised among other

attributes by small pods with the upper suture

distinctly notched between the articles. This

attribute distinguishes them from another

complex of species, D. glareosum, D. hannii,

D. muelleri and D. pullenii, where the upper

suture 1s continuous, not notched between the

articles.

The dimorphic foliage of OD.

trichostachyum distinguishes it from all other

species. Living plants have not been studied

but a sequence of development of leaves can

be deduced from the large number of herbartum

specimens examined. The plants which are

annual or short-lived perennials first develop

leaves with a single depressed ovate leaflet. If

environmental conditions favour rapid growth

plants may flower and fruit at this stage, or, if

not, may develop leaves with three cuneiform

leaflets before flowering. Consequently plants

with unifoliolate leaves only, trifoliolate leaves

only, or commonly a mixture of both are

represented in herbarium collections. The

lectotype specimen has both uni- and

trifoliolate leaves.

11. Desmodium filiforme Zollinger & Moritzi,

Natuur-en Geneeskunding Archief voor

Nedeéland’s Indie 3:58, 77 (1846);

Ohashi, Ginkgoana 1:237 (1973);

Meibomia filiformis (Zoll. & Moritzi)

Kuntze 1:198 (1891). Type: ‘Zol/. & Mor

Herb. 2738’. Java in arenosis maritimis

prope Poeger (iso:BM)}

Desmodium neurocarpum Benth., Fl. Austr.

2:234 (1864); Meibomia neurocarpa

Austrobaileya 5(2): 209-261 (1999)

(Benth.) Kuntze, Rev. Gen. 1:198 (1891).

Type: Upper Victoria River, Mueller

(holo: K).

Desmodium neurocarpun var.

queenslandicum Domin, Biblioth. Bot.

89:214 (1926). Type: In fl. Flinders River

ad opp. Hughenden, Feb 1910, Domin

‘4733’ (lecto: PR 527380, chosen here).

Sprawling annual, stems terete with

indumentum of long (to 1 mm) spreading hairs,

occasionally subglabrous. Leaves 3-foliolate

or, usually on young plants, 1-foliolate or both

1- and 3-foliolate; stipules triangular, 2~5 mm

long; petioles 4-18 mm long; rachis 1.5-6 mm

long; leaflets chartaceous, linear-oblong,

oblong or obovate-oblong, rounded or

subcordate at the base, emarginate at the apex,

usually with scattered appressed hairs on the

upper surface, sparse appressed hairs on the

lower surface, sometimes glabrous; terminal

leaflet 10—35(-45) mm long, 3.5—12.5(-14)

mm wide, I—5(—8) trmes longer than wide,

lateral leaflets smaller, (S—)10—22(-26) mm

long, 3—7(—10) mm wide, 1.4-4.5(—6.5) times

longer than wide. Inflorescences terminal,

open, to 20 cm long, rachis with long hairs,

usually spreading, sometimes appressed, to |

mim long; flowers single along rachis, primary

bract 2-3 mm long, deciduous, secondary

bracts and bracteoles absent; pedicels 3—4 mm

long. Flowers: calyx 2.3—2.5 mm long,

indumentum of minute uncinate hairs and long

(0.6—-1 mm) straight hairs on lobes; tube 0.7

mm long, all lobes about equal length, tending

sometimes to incurve, the upper divided to 4~/,

length, the lower sometimes narrower than the

others; corolla white to pink or mauve; standard

obovate, obtuse at apex, 2.5—2.7 mm long, 1.6—

2.2 mm wide, wings and keel petals about equal

in length, 2—2.6 mm long; stamens diadelphous;

ovary usually with rather sparse uncinate hairs

or occasionally glabrous. Pods straight, the

upper suture sharply notched between articles,

the lower undulate, isthmus about 4% width of

pod, with 3, 4 or rarely 5 articles, each (2.5)3—5

min long, 2.5-4(-4.5) mm wide, one exceptional

specimen with only one article per pod

developed, 5.5—6.2 mm long, 4 mm wide,

glabrous or sparsely uncinate pubescent,

reticulated veined (often purplish), raised in

centre over seed, usually the anastomoses small,

replete bev

AIS Ra nae ey RI ee E

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 239

+ equilateral in central area and larger, radially

rectangular to the periphery; seeds 2-3 mm

long, 1.5—2 mm wide.

Selected specimens: Java. Berachi, Poegar, Apr 1929,

Backer 3650 (L). New Guinea. Merauke, Aug 1904,

Kock (L); Koitaki, Jun 1935, Carr 12673 (CANB, K, L).

Western Australia. 16 km S of Port Hedland, 20°23’S

118°40°E, Apr 1977, George 14565 (K, PERTH); 19.6

km 8 of Derby, 17°27’S 123°44’E, Apr 1985, Aplin et

al. 38 (PERTH); 5 miles [8 km] W of Kununurra, East

Kimberleys, 14°46’S 128°41’°E, Mar 1963, Lazarides

6733 (CANB); NW of Deception Range, 15°53’S

128°36’E, Mar 1978, Hartley 14784 (CANB, PERTH);

Piccaninny Creek Gorge, 17°27’S 128°25’E, Blackwell

BB 187, 180b & 377 (PERTH). Northern Territory.

Daly River, 13°52’S 131°1L1’E, Apr 1988, Pullen &

Spottswood 11162 (CANB); McArthur River area,

16°07’S 136°07°E, Jun 1976, Craven 4088 (CANB);

South Bickerton Is., 13°45’S 136°06’E, Jun 1948, Specht

505 (BRI, CANB, K, MEL); Maude Creek, 12 miles [19

km] NE of Katherine, 14°23’S 132°24’E, Mar 1965,

Wilson 370 (CANB, K). Queensland. Burke District:

E of ‘Westmoreland’ Station, 17°30’S 138°20’E, May

1974, Pullen 9131 (CANB, K); Coox DISTRIcT:

Cooktown, mouth of Endeavour River, 15°28’S

145°1S’°E, May 1970, Blake 23296 (BRI, K, MEL);

Mareeba, Apr 1967, Pedley 2270 (BRI, K); Nort

KENNEDY District: 70 km § of Charters Towers, 20°40’S

146°17°E, May 1988, Pullen 11255 (CANB); Port Curtis

District: Rockhampton, Apr 1867, O’Shanesy 116

(MEL).

Distribution and habitat: Desmodium

filiforme, the most widespread species of ser.

Stenostachys in tropical Australia, occurs over

most of the northern half of Queensland and

extends across the Northern Territory to the

northern part of the Kimberley region of

Western Australia with isolated occurrences

near Port Hedland (George 14565) and Lake

Surprise, Lander River (Maconochie 1695). It

is also found in southern New Guinea and Java.

The species is confined to sandy soils,

sometimes seasonally waterlogged, in eucalypt

communities, dry beds of streams, flood-outs

and coastal dunes; in coastal areas it is

sometimes associated withD. trichostachyum.

Map 23.

Affinities: Because of its long terminal leaflet

D. filiforme most closely resembles D. pullenti

but differs from it in having the upper suture of

the pod notched between articles and the central

portion of each article raised over the seed with

the veins closely anastomosing and radiating

to the margins. It and D. brownii which has

narrower leaflets are probably more closely

allied toD. trichostachyum andD, pycnotrichum

which have similar pods.

Knaap-van Meeuwen (1962) included D.

muelleri, and judging from her remarks,

possibly alsoD. pulleniiinD. filiforme. Ohashi

(1973) pointed out that such a treatment was

unacceptable; he also suggested that D.

filiforme can be separated from D.

trichostachyum only with considerable

difficulty except for the shape of the leaflets

and pods. The shape of the leaflets and the

indumentum of the branchlets and leaves, but

not the shape of the pods, are sufficient to

differentiate the two species. Ohashi may also

have had material of more than one species.

12. Desmodium brownii Schindl., Notes Roy.

Bot. Gard. Edinburgh 25:13 (1926) &

Rep. sp. nov. reg, veg. 22:256 (1926).

Type: (syntypes): North Coast, Brown

sn. (E?, not seen); Brown ‘4188’ (iso:

BM, K).

Desmodium neurocarpum var. gracile

Benth., Fl. Austr. 2:234 (1864). Type:

North Coast, Brown (holo:K).

Erect annual to 75 cm tall; stems + terete with

indumentum of weak erect uncinate hairs, often

with stiff ascending or spreading hairs 0.6—1

mm long. Leaves 3-foliolate or 1-foliolate

towards the base of the plant; stipules rather

attenuate, 26.5 mm long; usually with long

hairs on the margin; petioles 2—9(-15) mm

long; rachis (1-)2—4(—5) mm long; leaflets

linear or oblong-linear, rounded at the base,

obtuse at the apex, glabrous or with sparse

appressed, rarely ascending hairs on upper

surface, sparse appressed hairs on lower;

terminal leaflet 10-50(—65) mm long, 1.8-4(--5.5)

mm wide, 3.5—13 times longer than wide, laterals

smaller 5.5—36 mm long, 1.5-4.5 mm wide 3-

12(—14) times longer than wide. Inflorescence

terminal and in the upper axils, to 20 cm long,

open; rachis with indumentum of uncinate hairs,

occasionally with long hairs near the base;

flowers single or rarely in pairs; primary bracts

1.5-—3 mm long, acuminate, pubescent on

margin in upper half, deciduous; secondary

bracts and bracteoles absent; pedicels filiform,

240

5—15 mm long. Flowers: calyx tube c. 0.5 mm

long, the lobes all 0.6-0.7 mm long, the upper

divided to about the middle, the lower

acuminate; corolla pink or mauve with standard

obovate c. 3 mm long, 3 mm wide; wings

oblong, c. 2 mm long; keel petals about as long

as the wings. Pod flat, the upper suture notched

at the junction of the articles, the lower more

gently incised, the isthmus /—’4 width of pod,

with 3 or 4 articles, each oblong or orbicular

2.9—-3.5 mm long, 2.2-3 mm wide, uncinate

hairs on faces and fringe of uncinate hairs on

margins, sometimes becoming glabrous,

reticulately nerved, raised in centre over the

seed, usually the anastomoses small, +

equilateral in central area, larger and radially

rectangular to the periphery; seeds 1.7—-2.5 mm

long, 1.3—-1.7 mm wide, with a small rim aril.

Selected speciinens: Western Australia. King Leopold

Range, 17°02’S 125°14°E, May 1988, Pullen 11206

(CANB); Carr Boyd Range, 20 km ENE of ‘Dunham

River’ H.S., 16°15’S 128°30’E, Mar 1978, Lazarides

2814 (CANB, K, PERTH); Weaber Range, 15°20’S

128°48’°E, Mar 1978, Hartley 14472 (CANB, PERTH).

Northern Territory. Woolaning, 13°07’S 130°40°E., Apr

1981, Dunlop 5880 (CANB, DNA, MEL NSW); 8 miles

[13 km] NNE of Edith River siding, Mar 1965, Lazarides

& Adams 117 (BRI, CANB, K, MEL); 16 miles [26 km]

SE of El Sharana mine, 13°41°S 132°41’E, Feb 1973,

Lazarides 7873 (AD, CANB, K, PERTH); Victoria River,

Gregory National Park, 15°28’S 131°IS°E, Wightman

2767 & Clark (DNA). Queensland. Cook District: 19

km S of Palmer River crossing 16°15’S 144°43’E, Mar

1987, Clarkson 6639 & McDonald (BRI, K, MBA).

Distribution and habitat: Desmodium brownii

is confined to the southern part of Cape York

Peninsula, Queensland, the northern part of the

Northern Territory and the Kimberley region

of Western Australia, on sandy soil, often

shallow or gravelly on hillsides. Map 24.

Affinities: The species is closely related to D.

filiforme to which Bentham referred it as a

variety (of D. neurocarpum), but differs in its

erect habit, narrower leaflets, rachis of the

inflorescence lacking spreading hairs except at

the base, and longer pedicels.

13. Desmodium flagellare Benth., Fl. Austr.

2:233 (1864); Schindl., Rep. sp. nov. reg.

veg. 22:255 (1926); Meibomia flagellaris

(Benth.) Kuntze 1:198 (1891). Type:

Beagle Valley |[approx. 15°35’S

130°SSE’, in 1855, Mueller (holo:K).

Austrobaileya 5(2): 209-261 (1999)

Prostrate annual spreading to a metre or more;

branches terete with sparse to moderately dense

ascending to + spreading hairs to 0.5 mm long

with shorter dense uncinate hairs. Leaves 3-

foliolate or [-foliolate towards base of plant;

stipules 3-6 mm long, somewhat acuminate,

with a few long hairs; petiole 5-12 mm long;

rachis 1.5-4 mm long; leaflets cuneiform,

obovate or oblong, rounded or slightly cordate

at the base, slightly retuse at the apex, glabrous

on upper surface, sparse appressed pubescent

on lower surface or hairs confined to midrib

and margins, hairs rather loose; terminal leaflet

15-25 mm long, 14-20 mm wide, the lateral]

ones smaller, 10-20 mm long, c. 12 mm wide,

all (1—)1.2-1.4 times longer than wide;

pulvinuses and stipels 1—-1.5 mm long.

Inflorescences terminal and in upper axils, to

20 cm long, rachis with sparse appressed hairs,

fruiting pedicels c. 20 mm apart on rachis,

rather closer than in related species; flowers

single on the rachis; primary bract concave,

ovate, 2-5 mm long, rather persistent,

secondary bracts and bracteoles absent;

pedicels 3—7 mm long. Flower: calyx 3-4 mm

long, the tube c. 1 mm long, upper lobe divided

to about the middle, lobes with long white

spreading hairs; corolla not examined; stamens

diadelphous; ovary densely uncinate pubescent.

Pods with upper suture continuous, not notched

between articles, the lower indented, isthmus

c. “/; width of pod, with 1-3, rarely 4 articles,

each 3.5-4.5 mm long, 3-3.5 mm wide,

uniformly sparsely to densely pubescent on

faces and margins with uncinate hairs; seeds

not seen.

Specimens examined: Western Australia. Kimberley

Research Station [15°39’S 129°43’E], Mar 1963,

Lazarides 6784 (CANB, K, PERTH). Northern

Territory. Victoria River, Gregory National Park,

15°35’S 131°21’E, Feb 1986, Wightnan 2777 & Clark

(DNA).

Distribution and habitat: Desmodium

flagellare is restricted to heavy soils and has

an extremely limited geographic range in the

lower part of the Ord and Victoria River basins.

Map 25.

Affinities: The type material of D. flagellare

is so poor as to make interpretation of the name

of the species difficult; not only is the type

specimen fragmentary but, as Schindler pointed

Lan RC Sa a en Bn COSTE Ee i on het on eet Lac betia en htm aramid to E

1 MISTY SS BU rns tnt dag nated

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 241

out, it is intertwined with another legume

(Cajanineae, probably Rhynchosia sp.). The

pods with the continuous upper suture and with

rather large articles, suggest a relationship with

D. hannii and D. muelleri, species also common

on heavy soils. Further specimens are needed

for the species to be described in detail and its

relationships clarified.

14, Desmodium hannii Schindl., Rep. sp. nov.

reg, veg, 21:4 (1925). Type: Carpentaria,

F.C. Hann (Br). Queensland. Cook

District: 4.8 km N of Little Laura River

on Peninsula Development Road,

(15°30’S 144°17’W), 27 April 1983, 7B.

Clarkson 4780(neo: BRI; isoneo. CANB,

K, MBA, QRS; chosen here).

Decumbent or prostrate annual herb, stems up

to 1 m long; branches with sparse spreading or

+ appressed white hairs and sometimes shorter

uncinate hairs. Leaves 1- or 3-foliolate (usually

on same plant); stipules 2.5-4.5 mm long, 1-

1.8 mm wide at base; petioles (1.5—)4-9(—16)

mm long, rachis 1—3.5 mm long; leaflets

chartaceous, cuneiform or broadly obovate

when trifoliolate, or obicular when 1|-foliolate,

rounded or slightly emarginate at the base,

rounded at the apex, glabrous or with scattered

appressed hairs above, with sparse to moderate

appressed or somewhat ascending hairs

especially on midrib beneath, 3 or 4 prominent

lateral veins on each side of midrib; terminal

leaflet 6—25(-30) mm long, 6-20 mm wide,

Q.9-—-1.5(—1.8) times longer than wide; lateral

leaflets smaller, 6-15 mm long, 6-12 mm wide,

0,9—1.3 times longer than wide. Inflorescences

terminal and/or axillary, open, to 12 cm long;

rachis with appressed hairs c. 1 mm long;

flowers solitary; primary bracts ovate, 3.2—3.5

mm long, early deciduous; secondary bracts

and bracteoles absent; pedicels 3—4.5 mm long.

Flowers: calyx 2.7-3.2 mm long, long hairs on

margins and midlines of lobes; tube 0.7-0.9 mm

long, upper lobe 2—2.5 mm long, narrowly

triangular, wider than the others, divided to

about the middle; lateral lobes 1.8-—2.5 mm

long, lower lobes c. 2 mm long, slightly wider

than laterals; corolla pink or mauve; standard

obovate, obtuse at the apex, 3-4 mm long, |.6—

3 mm wide; wings 2.3—2.8 mm long, shortly

clawed, c. 1 mm wide; keel petals longer and

narrower than wings, 2.6—-3.5 mm long, 0.5—1

mm wide; stamens diadelphous; ovary

pubescent. Pod lightly but distinctly recurved,

the upper margin continuous, not notched

between articles, the lower incised, the isthmus

'4,—Y2 or more width of pod, with 2-4, rarely 5

articles, each (2.5—)2.8-4.2(-5) mm long, 2.5—

3.5 mm wide, pubescent with uncinate hairs,

particularly on margins; seeds c. 2.5 mm long,

1.6 mm wide.

Selected specimens: Western Australia. King Leopoid

Ranges, 17°07’S 125°20’E, May 1988, Pullen 11205

(CANB). Northern Territory. 3 km S of Katherine,

14°29°S 133°46’E, May 1988, Pullen & Spottswood

14176 (CANB); Beswick Aboriginal Reserve, 14°33’S

133°00°E, May 1974, Pullen 9329 (BRI, CANB, K); 38

miles [61 km} ESE of ‘Limbunya’ Station, Jun 1949,

Perry 2294 (CANB, PERTH). Queensland. Coox

District: 4.8 km N of Little Laura River, 15°30’S

144°|7°E, Apr 1983, Clarkson 4780 (BRI, CANB, K,

MBA, QRS).

Distribution and habitat: Desmodium hannii

has a discontinuous distribution in tropical

Australia. It appears to be most common and

has been most frequently collected 1n the north-

central part of the Northern Territory with two

collections from the Kimberley region of

Western Australia. It also occurs in north-

eastern Queensland. It favours heavy soils and

sometimes occurs withD. muelleri (collector’s

note: Pullen 9329) and its absence from

grassland dominated by Astrebla spp. and

Dichanthium spp. in north-western Queensland

is unexpected. It is reported to be favoured by

stock. Map 26.

Affinities: The species is a member of ser.

Stenostachys closest to D. muelleri from which

it differs in its curved pods, leaves often

unifoliolate with less elongate leaflets. It may

be aneotonic derivative of D. muelleri bearing

the same relationship to it asD. trichostachyum

does to D. pycnotrichum.

15. Desmodium muelleri Benth., Fl. Austr.

2:235 (1864); van Meeuwen, Reinwardtia

6:249 (1962), pro syn. Meibomia

muelleri (Benth.) Kuntze, Rev. Gen.

1:198 (1992), Type: Upper Victoria

River, Mueller (holo:Ks).

Decumbent or prostrate annual; stems with

indumentum of sparse stiff, usually appressed,

242

occasionally ascending, rarely spreading, white

hairs to c. 1 mm long. Leaves 1- or 3-foliolate,

both on same or occasionally on different

plants; stipule 3-6 mm long glabrous or with

marginal hairs; petiole 3-12 mm long, rachis

1.5—5 mm long; leaflets chartaceous, narrowly

ovate, elliptic or oblong, occasionally broadly

oblong when 1-foliolate, rounded at the base,

obtuse or retuse at the apex, glabrous or

occasionally with sparse appressed hairs on

upper surface, sparse or moderately dense

appressed hairs on lower surface; terminal

leaflet 12-45(-60) mm long, 5-15(-20) mm

wide, 1.2—6 times longer than wide, the laterals

smaller, 10—30(-35) mm long, 2.5-12 mm

wide, 1.5—6 times longer than wide; pulvinuses

0.5-1.5 mm long, stipels 0.7—2.5 mm long.

Inflorescences terminal, usually rather sparsely

flowered, to 30 cm long, rachis glabrous or with

uncinate and/or scattered to rather dense

straight spreading hairs to 1 mm long; flowers

single or in pairs, subtended by primary bract

(2.5—)3—4.5 mm long, pubescent or hairs

confined to margin; secondary bracts and

bracteoles absent; pedicel 7-10 mm at anthesis,

15 mm long in fruit. Flowers: calyx 2.8—3.3

mm long, long straight hairs on lobes, the tube

c. | mm, upper lobe divided to about the

middie, 1.5—1.8 mm long, the others setaceous,

1.6—2.3 mm long, the lower slightly longer than

the laterals; corolla pink to mauve with standard

obovate, obtuse at the apex, 3.5—5 mm long,

2.5—3 mm wide, keel 3.5—4.5 mm long, slightly

longer than the wings; stamens diadelphous;

ovary densely pubescent. Pods straight, the

upper suture continuous, not notched between

articles, the lower slightly incised, the isthmus

‘/;or more width of pod; with 4-6 articles, each

(3.6—)4—5 mm long, 3—4 mm wide, transversely

reticulately veined, moderately uncinate

pubescent on faces with distinct marginal hairs;

seed (2.3—)2.8-3.2 mm long, (1.6—)1.8-2.3 mm

wide with distinct rim aril.

Selected specimens: Western Australia. Walsh Pt, Port

Warender, Keighery 4806 (PERTH); Mitchell Plateau

mining camp, 14°26’S 125°47’E, Apr 1977, George

14480 (kK, PERTH); Kimberley Research Station, Jul

1952, Perry 3036 (CANB, K, MEL, PERTH) & Mar

1963, Lazarides 6786 (CANB, PERTH). Northern

Territory. 4 miles [6 km] N of ‘Rankin’, Mar 1956,

Chippendale NT 1869 (DNA, MEL, PERTH); 12 km S

of Katherine, 14°31°S 132°23’E, Apr 1988, Pullen &

Spottswood 11172 (BRI, CANB, K); c. 26 miles [42 km]

Austrobaileya 5(2): 209-261 (1999)

NNW of ‘Brunette Downs’, May 1947, Blake 17813

(BRI, CANB, K, MEL). Queensland. Burk& District:

30 miles [48 km] NNE of Camooweal, May 1948, Perry

982 (CANB, K); SW of Normanton, 17°45’S 140°S7°E,

Apr 1974, Pullen 8887 (CANB, K); NortH KENNEDY

District: between Powelathunga and Charters Towers,

20°10’S 146°00’°E, May 1988, Pullen 11250 (BRI,

CANB).

Distribution and habitat: Desmodium muelleri

ranges across northern Australia from the

Kimberley region of Western Australia to

south-western and north-central Queensland

(near Charters Towers). It extends into semi-

arid areas, and is virtually confined to heavy

soils, often in grassland dominated by Astrebla

spp. or Dichanthium spp. The western part of

its range is rather fragmented, probably because

of lack of suitable habitat. Map 27.

Affinities: Desmodium muelleri and D. hanniti

are closely related and are distinguished from

other species of ser. Stenostachys (except

perhaps from the poorly known D. flagellare)

by pods not notched between the rather large

articles. As Ohashi (1973) and Verdcourt

(1979) pointed out, Knaap-van Meeuwen’s

referral of D. muelleri to D. filiforme is

unacceptable.

16. Desmodium glareosum Pedley, sp. nov.,

D. muelleri Benth. leguminis sutura

supera continua non inter articulos acute

incisa similis autem leguminis articulis et

foliolis parvioribus differt; a D. pullenii

Pedley pilis ramulorum foliolorumque

albis incohaerentibus, foliolo terminali

angustiore differt; a D. filiformi Zoll. et

Moritzi leguminis sutura supera continua

non inter articulos acute incisa, pilis

ramulorum albis differt. Typus:

NORTHERN TERRITORY: Cc. 8 miles [13 km]

NNE of Edith River Siding, March 1965,

M. Lazarides & L.G. Adams 115 (holo:

CANB; iso: BRI, DNA, K, L, NSW).

Annual, erect or becoming prostrate, foliage

described as greyish; stems with indumentum

of usually weak moderately dense ascending

(rarely appressed) white hairs 0.6—1 mm long.

Leaves 3-foliolate; stipules narrow lanceolate,

1.5—4 mm long, 0.5—1.2 mm wide, with a few

long hairs on the margins; petiole (0.5—)2.5—8

mm long, rachis 1.5—3.5 mm long; leaflets

SEM CS Phere nnn Ta a er DN hg BE Ee Sd oh oo me PSOE TSS

EMSS Ree eT ci tea

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 243

oblong, obovate or almost orbicular, rounded

or subcordate at the base, obtuse or retuse at

apex, indumentum of weak, sparse to moderate

loose ascending hairs c. 0.6 mm long on both

surfaces; terminal leaflet 5—-19(—26) mm long,

2.5-6.5(-11.5) mm wide, 1.4-4.5(—7) times

longer than wide, lateral leaflets smaller, 3.5—

13(-20) mm long, 1.8—6(—-8.5) mm wide, 1.1—

3.5(-4.5) times longer than wide; pulvinuses

Q.5-1 mm long, stipels 0.3-0.6 mm long.

Inflorescences terminal, rather open, to c. 10

cm long, the rachis with moderately dense,

rather spreading hairs; flowers single, pedicels

slender, 5-15 mm long, subtended by

deciduous ovate pubescent bract 1.5—3 mm

long; secondary bract and bracteoles absent.

Flower: calyx 1.5—1.7 mm long, with

indumentum of sparse spreading hairs to 0.8

mm long, tube 0.5—0.7 mm long, lobes c. 1 mm

long, the upper divided to about the middle;

corolla pink or mauve; standard c. 3 mm long

and 2 mm wide; wings c. 2 mm long, | mm

wide; keel petals slightly longer than the wings,

2—-2.8 mm long; stamens diadelphous; ovary |

moderately pubescent with ascending hairs.

Pods + straight, the upper suture + continuous

(as in D. muelleri, D. hannti and D. pulleniti),

not notched between articles; isthmus “4—/

width of pod; lower suture deeply indented;

with 2—4(—5) articles, each 2.5~3.4 mm long,

2.2—3(—3.3) mm wide, sparsely to moderately

pubescent on faces, fringed on sutures,

transversely reticulately veined; seeds 1.7—2.1

mm long, 1.4—1.8 mm wide, shining black when

mature.

Selected specimens: Western Australia. Mitchell

Plateau, 14°49’°S 125°45’E, May 1988, Pullen 11219

(CANB). Northern Territory. Manton River, 12°50’S

131°LQ’E, May 1973, Dunlop 3146 (BRI, CANB, DNA,

Kk, NSW); 11 km E of Katherine, 14°26’S 132°15’E, Apr

1977, Pullen 10592 (CANB, K, L); 16 km NE of

Katherine, 14°24’S 132°22’E, Apr 1988, Pullen 11171

(AD, BRI, CANB, K, MEL, NSW, PERTH); 22 miles

[35 km] N of Katherine, Feb 1965, Wilson 301 (BRI,

CANB, DNA, K, L).

Distribution and habitat: The species has

usually been recorded from gravelly soils,

sometimes in disturbed situations such as

roadsides. It is evidently sometimes associated

withD. pullenii, It occurs in Western Australia

(Mitchell Plateau and Weaber Range), and in

the north-west of the Northern Territory. Map 28.

Affinities: Desmodium glareosum is a member

of ser. Stenostachys, one of a group of species

that includes D. muelleri, D. hannii and D.

pullenii with the upper suture of the pod +

continuous (only slightly undulate) not sharply

notched between articles. It is most closely

related to D. pullenii.

Etymology: The specific epithet is Latin

glareosum, gravelly, a reference to the usual

substrate of the species.

17. Desmodium pullenii Pedley, sp. nov.

similis D. muelleri Benth. leguminis

sutura supera continua inter articulos non

incisa autem leguminis articulis

seminibusque parvioribus, foliolis

interdum parvioribus pilis ramulorum

aliquantum densis flavidis curvatis

differt. Typus: Northern Territory. 12 km

S of Hayes Creek, 13°35’S 131°30’E,

April 1988, R. Pullen & Spottswood

11169 (holo: CANB; iso: BRI, DNA,

NSW, PERTH).

Desmodium sp. (Laura V.J.Neldner + 3836)

Pedley in Henderson: Queensland Plants:

Names & Distribution: 79 (1997).

Weak erect annual herb to c. 30 cm high; stems

with indumentum of sparse to moderately dense

stiffly spreading or curved hairs 0.5—1 mm long.

Leaves trifoliolate; stipules deltoid to

lanceolate, 2.5-6 mm long, glabrous or with a

few marginal hairs; petioles 3-13 mm long,

rachis 1.5-6.5 mm long; leaflets obovate or

oblong, rounded at the base, obtuse or slightly

retuse at the apex, upper surface with sparse

appressed or rarely ascending hairs, rarely

glabrous, lower surface appressed pubescent,

hairs 0.5—1 mm long; terminal leaflet 7—22(—

27) mm long, 4.5~-11 mm wide, 1.4—2.5(-4)

times longer than wide, lateral leaflets

somewhat smaller, 5-17(-23) mm long, 3.2—

8(—9.5) mm wide, 1.2~2.5 very rarely to 4.5

times longer than wide; pulvinuses and stipels

0.5—1.3 mm long. Inflorescence open, to 20 cm

long, rachis with weak long spreading (rarely

appressed) usually yellowish hairs; flowers

single along the rachis; primary bract 2-3 mm

long, deciduous; secondary bract and

bracteoles absent; pedicels filiform 5-14 mm |

long. Flowers: calyx 1,5—2 mm long wth short

244

(0.1—0.3 mm) spreading hairs, tube 0.4—0.6 mm

long, lobes about of equal length, 1.2—-1.5 mm

long, the upper bifid to 0.3—0.5 mm; corolla

ranging in colour from white through pink to

pale purple, usually described as pink: standard

obovate occasionally somewhat truncate at

apex, 3-4 mm long, 2.4~2.7 mm wide, wings

2—2.5 mm long, c. 1 mm wide, shortly clawed;

keel petals longer than wings, 3-4 mm long,

about as wide; stamens diadelphous; ovary

pubescent. Pods straight, upper suture

continuous, not sharply notched between

articles, lower tncised, isthmus “4—% width of

pod; with usually 4 or 5 articles, each 2.7—3.5

mm long, 2.5—3.2 mm wide, moderately

densely uncinate pubescent with fringe of hairs,

occasionally becoming glabrous, transversely

reticulately veined; seeds 1.8—2.2 mm long,

1.5—1.7 mm wide, shining black at maturity,

with a small rim aril.

Selected specimens: Lesser Sunda Islands. Alor:

Landschaft Kei, Moroe-Gendok, May 1938, O. Jaag (L).

Western Australia. 5 km N of Port Warrender road,

Mitchell Plateau, 14°45’S 125°46’E, May 1988, Pullez

11222 (CANB, PERTH); 2.9 km SE of Mitchell Plateau

mining camp 14°50’S 125°51’E, Apr 1977, George

14504 (AD, MEL, PERTH). Northern Territory.

Beatrice Hill [12°39’S 131°19°E], Mar 1961,

Chippendale NT 7963 (CANB, DNA); 25 km W of Mary

River, 12°52’S 131°48’E, Mar 1978, Adaconoclhie 2332

(AD, BRI, CANB, K, MEL); Katherine Gorge National

Park, 14°20°S 132°25’E, Apr 1981, Craven 6708

(CANB). Queensland. Cook District: Endeavour River,

in 1878, Persietz 232 (MEL); Yorkey’s Knob Beach

116°49°S 145°43’E], Apr 1962, AdceKee 9011 (BRI,

CANB, K); NortH KeNNeEDyY District: Castle Hill,

Townsville, [19°16’S 146°45’E], Nov 1947, Shay 5507

(CANB),

Distribution and habitat: In Australia D.

pullenii ranges across northern parts from the

north of Western Australia to about Gladstone

in central Queensland. It occurs on sand dunes

and in eucalypt communities usually on sandy

soils, but it has also been recorded from clay.

It has also been collected in the Lesser Sunda

Is. Map 29.

Affinities: The species belongs to ser.

Stenostachys and is closely related to D.

glareosum which has long loose white hairs on

the stems and leaves, and usually narrower

terminal leaflets. The Malesian specimen cited

was determined by Knaap-van Meeuwen as D.

Austrobaileya 5(2): 209-261 (1999)

filiforme and she probably included D. pullenii

in her wide circumscription of that species.

Schindler annotated some specimens in herb.

Kew as D. muelleri var. minus and the name

has been used in other herbaria though never

validated.

Etymology: The epithet honours Mr Roy

Pullen, formerly of CSIRO, Canberra, whose

field, herbarium and other studies of

Desmodium in Australia have made this

account of the genus possible.

18. Desmodium heterocarpon (L.) DC.,

Prodr. 2:337 (‘heterocarpum’); Liu &

Chuang, Taiwania 8:80 t. 9 (1962);

Fosberg, Micronesiaca 2:145 (1966);

Schubert, FI. Trop. East Africa. Legum.

Papilion. 462 (1971); Ohashi, Ginkgoana

1:120 (1973), Man. Flow. Pl. Hawai’s

1:667 (1990), J. Jap. Bot. 66:14 (1991)

& (in part) Fl. Camb. Laos Vietn. 27:136

(1994); Verdcourt, Man. New Guinea

Leg. 399 (1979); Pedley, Rev. Handb. FI.

Ceylon 10: 187 (1996); Meibomia

heterocarpa(L.) Kuntze, Rev. Gen. 1:198

(1891); Hedysarum heterocarpon L., Sp.

Pl. 747 (1753). Type: Herb. Hermann

vol. 2 fol. 32, No. 94, left-hand specimen

lecto: BM, fide Pedley in Turland &

Jarvis (1997).

Desmodium polycarpon (Poir.) DC., Prodr.

2:334 (1825) (as ‘polycarpum’); Benth.,

Fl. Austr. 2:235 (1864); Hedysarum

polycarpon Poir. in Lam., Encycl. Meth.

6:413 (1805). Type: ‘Indes orientales in

herb. Lam’ (P, n.v.).

Hedysarum tuberculosum Labill., Sertum

Austro-Cal. 71 t. 72 (1824). Type: New

Caledonia, La Billadiére (n.v.).

Desmodium trichocaulon DC., Prodr. 2:335

(1825); Benth., Fl. Austr. 2:235 (1864).

Type: Nepal, Wallich (G-DC, n.v.;

microfiche, BRI).

Two varieties occur in Australia.

18a. Desmodium heterocarpon (L.) DC. var.

heterocarpon

Rachis of inflorescence with sparse short erect

uncinate hairs.

Fen MAT Ee SE

Het RNA NES MS Mae ed a

RGR Pees

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision

16. Desmodium brachypodum

19. Desmodium triflorum

22. Desmodium trichostachyum

25. Desmodium flagellare

28. Desmodium glareosuni

m2 a ae loot re

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att Rae re \ act MS 2

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17. Desmodium campylocauion 18. Desmodium microphyllum

20, Desmodium heterophyllum 21. Desmodium pycnotrichum

23. Desmodium filiforme 24. Desmodium brownii

26. Desmodium hannii 27, Desmodium muelleri

29, Desmodium pullenii 30. Desmodium heterocarpon vat.

heterocarpon

245

246

Selected specimens: Queensland, Coox District:

Cooktown, Jan 1958, Blake 21218 (BRI); Mt Molloy,

Apr 1962, AfcKee 9127 (BRI); SouTH KENNEDY DISTRICT:

Mt Fox, Sep-Dec 1949, Clemens s.n. (ex Univ. Michigan)

(BRI, K); Moreton District: Petrie, in 1931, Blake s.n.

(K}, New South Wales. Lismore, Mar 1893, Bauerlen

985 (NSW).

18b.Desmodium heterocarpon = var.

strigosum Meeuwen, Reinwardtia 6:95

(1962). Type: New Guinea, Kalkman

B.W. 3596 (holo: L, n.v.).

Rachis of inflorescence with dense long straight

white or yellowish hairs.

Selected specimens: Northern Territory. Port Essington,

Armstrong 373 (K); Bamboo Pass, Marrakai road, Mar

1967, Byrnes 211 (AD, DNA); Holmes Jungle, 8 miles

[13 km] E of Darwin, Mar 1961, Chippendale NT 7902

(DNA, MEL). Queensland. Cook District: Lockerbie,

10°48°S 142°28’E, Apr 1948, Brass 18437 (BRI, CANB);

Endeavour River, Jun 1819, Cunningham 244 (kK, MEL);

Moreton District: Buderim, in 1912, Longman s.n. (K).

New South Wales. Along Bonalbo road, north of Bottle

Tree village, 28°48’S 152°39’E, May 1973, Tindale 2094

(MEL, NSW).

Distribution and habitat: The species is widely

distributed in the Old World: Africa, Sri Lanka,

India, south-eastern Asia, Malesia and

throughout the Pacific. In Australia it ranges

from north-eastern New South Wales, through

eastern Queensland to the northern part of the

Northern Territory, usually in eucalypt

communities on well drained soils, occasionally

a weed in disturbed situations. Maps 30,31.

Affinities: Desmodium heterocarpon is one of

the four species of sect. Nicolsonia (subg.

Sagotia) that occur in Australia, but is not

closely related to the others. Ohashi (1991)

recognised eight infraspecific taxa, including

one based principally on flower colour. Both

Australian varieties are referred to subsp.

heterocarpon. D, heterocarpon vat. strigosum

is much the commoner of the two, as it is in

Africa, New Guinea and the Pacific.

Typification: In the protologue of Hedysarum

heterocarpon, Linnaeus quoted specific

differential characters from his own Flora

Zeylanica and J. Burman’s Thesaurus Zeylanicus.

The specific differential characters in both these

works emphasised that pods on the lower part

of the plant are one-seeded (Flora Zeylanica:

Austrobaileya 5(2): 209-261 (1999)

‘leguminibus articulatis: infimo monospermo’;

Thesaurus Zeylanicus: ‘siculis inferioribus

solitariis: superioribus articulatis’). Burman’s

plate cited by Linnaeus does show pods with,

in the lower part of the plant, one article, but

none of the five specimens in herb. Hermann,

only one of which has pods, has a one-seeded

pod. It could be argued that since none of

Hermann’s specimens agrees with the

protologue and Burman’s plate does, then, if

the species is to be lectotypified, Burman’s

plate should be chosen as the type.

Unfortunately the plate is not a good depiction

of the species currently accepted as

H. heterocarpon: the shape of the leaflets is not

at all typical and more importantly the fruiting

inflorescences are shown as being rather open

with the pods not appressed to and hiding the

rachis as they usually do. Since other species

(for example, Hedysarum diphyllum, H.

pulchellum and H. umbellatum) are so well

illustrated in Thesaurus Zeylanicus as to be

instantly recognisable, one is left with some

doubt as to whether the plant illustrated is the

Hf, heterocarpon of herb. Hermann. This doubt

is deepened by the fact that though I have

examined scores of specimens of UH.

heterocarpon collected throughout its range I

have not seen a specimen with pods with only

one article. In view of this doubt, my general

reluctance to typify a name with a plate when a

specimen Is available, and a desire to preserve

nomenclatural stability, I lectotypified

Hedysarum heterocarpon L. on herb. Hermann

vol. 2 fol. 32, the left-hand specimen. The

specimen bears flowers, but not pods, and

shows both upper and lower surface of leaflets

and the characteristically dense inflorescence

with the rachis uncinately pubescent.

19. Desmodium strigillosum Schindl., Bot.

Jahrb. 54:57 (1916); Gagnepain, FI. Gen.

Indochin. 2: 583 (1920); Ohashi,

Ginkgoana 1:233 t.32a, fig. 63, 64

(1973); Dy Phon et al., Fl. Camboge,

Laos, Vietnam 27: 129 t.27 (1994). Type:

Vietnam. Saigon, November 1864,

Lefevre s.n. (ecto: P, n.v., fide Ohashi).

Selected specimens: Vietnam. Onorlei, dans les lieux

incults, Nov 1885, Balavsa Pl. du Tonkin 1250 (K,

paralectotype). Queensland. Coox Districr: Andoom Top

Camp, 20 km N of Lorim Point,12°29’S 141°S0’E, Mar.

1981, Aforton AM 1153 (BRI); Cooktown, Jan 1958,

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 247

Blake 20218 (BRI); 2.5 km S of Mt Molloy on road to

Mareeba, 16°42’S 145°20’S, Jan 1982, Clarkson 6756

& McDonald (BRI).

Distribution and habitat; The species has

previously been recorded only from south-east

Asia (Burma, Cambodia, Laos and Vietnam,

fide Dy Phon et al.) at altitudes of 700-800m

(though the lectotype is evidently from

Saigon!). In Queensland it was first collected

at Cooktown in 1958 and seems to be a recent

introduction. It now occurs at rather low

elevations in the Mareeba-Cooktown area and

at the extreme north of the west coast of Cape

York Peninsula. Map 32.

Affinities: It is probably most closely allied to

D. heterocarpon from which it differs, most

obviously, in its lanceolate leaflets, large

flowers and reflexed pods.

20. Desmodium nemorosum F. Muell. ex

Benth., Fl. Austr. 2:234 (1864); Ohashi,

Ginkgoana 1:218 (1973), excl. var.

whitfordit; Verdcourt, Man. New Guinea

Leg. 403 (1979); Meibomia nemorosa

(F. Muell. ex Benth.) Kuntze, Rev. Gen.

1:198 (1891). Type: Brisbane River,

Mueller (syn: K); Pine River, Fitzalan

(syn: K); sine loc., Leichhardt (syn: K;

iso?: NSW).

Desmodium nemorosum var. simplex

Schindl., Rep. sp. nov. reg. veg. 21:10

(1925). Type: none designated.

Desmodium nemorosum — subvar.

eboracense Schindl., Rep. sp. nov. reg.

veg. 21:10 (1925). Type: Cape York,

Daemel (iso: BM).

Desmodium nemorosum var.

novoguineense Kaneh. & Hatus., Bot.

Mag. Tokyo 56:366 (1942). Type: New

Guinea. Vogelkop, Waren, 60 miles [96

km] S of Manokwari, Mar 1940,

Kanehira & Hatusima 12930 (holo: FU,

n.v.; photo: K),

Desmodium archboldianum E.G. Baker,

Brittonia 2:318 (1937). Type: Papua New

Guinea. Western District: Wuroi, Oriomo

River, Jan 1934, Brass 5732 (holo: BM;

iso: BRI, L).

Selected specimens: Queensland, Coox DISTRICT:

Newcastle Bay [10°49’S 142°26’] May 1948, Brass

18763 (BRI, CANB); NortH KENNEpy District: Mt Fox

[18°S1’°S 145°48’E], Dec 1949, Clemens (CANB);

MoRETON DISTRICT: Bellthorpe—Jimna road, 26°44’S

152°35’H, alt. 600m, Jan 1990, Weston 1469 & Richards

(BRI, NSW); Tamborine Mtn, Jan 1916, White s.n. [AQ

097953] (BRI). New South Wales. 5 miles [8 km] N of

Ttuka, Jun 1966, Boyd & McGillivray 2008 (NSW); Port

Macquarie, May 1819, Cunningham 41 (K) & Feb 1898,

Boorman (NSW); Camden Haven, Jan 1882, Befche

(NSW 105278).

Distribution and habitat: Desmodium

nemorosum extends from New Guinea through

coastal parts of eastern Queensland to north-

eastern New South Wales, in higher rainfall

areas in eucalypt open-forest and on margins

of rainforest. Though widespread, it is nowhere

common, and appears to be absent from central

Queensland between about 20°S and 25°S.

Map 33.

Affinities: The species differs from D.

heterocarpon the other species of sect.

Nicolsonia mainly in its elongate articles 5—7

mm long and its more elongate leaflets. Like

D. heterocarpon, D. nemorosum shows a wide

variation in size in both foliar and floral parts,

but unlike D. heterocarpon it cannot be divided

into infraspecific taxa. D. nemorosum subvar.

whitfordii Schindl. (D. nemorosum var.

whitfordii (Schindl.) Ohashi), syntypes of

which I have seen (herb. Kew), is best

considered a distinct species with a restricted

range in the Philippines some 2500 km from

Vogelkop the nearest known occurrence of D.

nemorosum in New Guinea. It has distinctly

ovate leaflets with dense long appressed silky

hairs on their undersides. Specimens from

Australia referred to D. nemorosum var.

whitfordii by Ohashi are well within the normal

range of variation of D. nemorosum var.

nemorosum.

Desmodium whitfordii (Schindl.) Pedley,

comb. nov.

Desmodium nemorosum subvar. whitfordii

Schindl., Rep. sp. nov. reg. veg. 21:10

(1925). Types: Philippines. Luzon:

Bataan Prov., Lamao River, Mt Marweles

(Herb. Bur. Gov. Lab. No. 227 leg. H.N.

Whitford, Herb. Bur. Sci. No. 7611 leg.

ELD. Merrill.) Gsosyn: K).

248

21. Desmodium tiwiense Pedley, sp. nov.,

affinis D. nemoroso F. Muell. ex Benth.

a qua foliis semper unifoliolatis, foliolis

orbicularibus (non ovatis ellipticisve)

infra minor dense pubescentibus et venis

lateralibus paucioribus ornatis et

leguminibus margine inferiore minor

valde inciso et articulis paucioribus

differt. Typus: Northern Territory:

Bathurst Island, Runku, 11°36°59”S

130°16°35”E, 27 March 1995, 7D, Cowie

5424 (holo: DNA; iso BRI, K, & (7. v)

CANB, L, MEL, NSW, PERTH).

Perennial trailing herb; stems terete with

indumentum of dense spreading or ascending

white hairs (to 0.7 mm long). Leaves 1-

foliolate; sttpules brown scarious, long-pointed,

somewhat asymmetrical at the base, 7—10 mm

long,1.2—2 mm wide; petiole 12-16 mm long;

leaflets more or less orbicular, cordate at the

base, obtuse at the apex, glabrous above,

moderately appressed pubescent below, with

c. 7 pairs of lateral veins not reaching the

margin, (25—)30—36 mm long, (25—)28-—32 mm

wide; pulvinuses 1—1.5 mm long; stipels

filiform 3-4 mm long. Inflorescences terminal,

open, to 12 cm long, rachis pubescent with

moderately dense, slightly retrorse, uncinate

hairs; flowers in pairs subtended by a deciduous

ovate acuminate bract c. 3 mm long, secondary

bracts and bracteoles absent; pedicels 2-3 mm

long. Flowers: calyx c. 3 mm long, very shortly

uncinately pubescent and with a few long hairs

at the base, tube c.1 mm long, all lobes

triangular c. 2 mm long, the upper shortly bifid

at the tip; corolla white or yellowish, standard

orbicular, emarginate at the top, 5.5 mm long,

6.5 mm wide, wings oblong c. 5 mm long

Gncluding claw c.1 mm), keel petals about as

long as wings; stamens rather stout; ovary

appressed pubescent. Pod to 25 mm long with

5 or 6 articles, upper suture straight or slightly

sinuate, not notched between articles, lower

shallowly indented (less than % width of pod)

between articles; articles quadrate 3—3.5 mm

long and wide, with sparse uncinate hairs,

denser on margins, transversely reticulately

nerved; seed c. 2 mm long and 1.5 mm wide

with a small rim aril.

Other specimens (all DNA): Northern Territory.

Melville Island, 11°28’S 130°30°E, Feb 1987, Fensham

Austrobaileya 5(2): 209-261 (1999)

445; Cape Gambier, Melville Is., 11°52S 130°40’E, May

1988, Fensham 828; Melville Is., 11°42’S [30°42E, Apr

1987, Fensham 508; Melville Is., Penell Beach to

MacClear Creek Rd, Jun., 11°49’S 130°54’E, Jan 1992,

Leach 2938 & Cowie.

Distribution and habitat: The species has a

limited geographic range, restricted to Bathurst

and Melville Islands in the extreme north-west

of the Northern Territory where it occurs In

eucalypt communities on sandy or gravelly

soils. Map 34.

Affinities: Desmodium tiwiense is most closely

related to D. nemorosum which usually has

trifoliolate leaves, narrower leaflets more

densely pubescent beneath with more lateral

veins, the lower margin of the pod more deeply

incised, and larger pod articles.

Etymology: Bathurst and Melville Islands are

the traditional home of the Tiwi people and are

often known as the Tiwi Islands. The specific

epithet is a combinationof tiwi, and the Latin

suffix (neuter) -ense, place of origin.

22. Desmodium velutinum (Willd.) DC.,

Prodr. 2:328 (1825); Schubert, Fl. Congo

Belge 5:194 (1954) & FI. Trop. East

Africa. Legum.: Papilion.: 466 (1971);

Ohashi, Ginkgoana 1:192 (1973) & FI.

Camb. Laos Vietn. 27:117 (1994) ;

Verdcourt, Kirkia 9:518 (1974), Manual

New Guinea Leg. 411 (1979); Pedley,

Rev. Handb. Fl. Ceylon 10:183 (1996);

Meibomia velutina Willd., Sp. PI.

3(2):1174 (1802). Type: herb. Willdenow

13763 (holo: B; microfiche, BRI).

Selected specimens: Northern Territory. Mt Bundy,

12°52’S 131°38’E, Apr 1969, Byrnes 1486 (DNA, K); 9

km N of Cannon Hill, 12°17’°S 132°5S’E, Jun 1983,

Russell-Smith 699 (DNA); Bamboo Pass, 12°55’S

131°1S’E, Mar 1968, Byrnes 213 (AD, DNA),

Distribution and habitat: In Australia confined

to the extreme north-west of the Northern

Territory where it occurs in eucalypt

communities and on margins of closed

communities. It may have been introduced into

Australia in recent years as all collections have

been made since 1965. It is widespread in

Africa, south-east Asia and Malesia. Map 35.

Affinities: With D. gangeticum it is the only

representative of sect. Heferoloma, but it is not

TSE EV Ss Ei wan cade oa teen eye

Pediey, L., Desmodium and related genera in Australia: a taxonomic revision 249

particularly closely related to D. gangeticum

or any other species of the section. Ohashi

recognised two subspecies, one with two

varieties, Australian specimens are referable to

D. velutinum vay. velutinum.

23. Desmodium gangeticum (L.) DC., Prodr.

2:327 (1825); Schubert, Fl. Congo Belge

5:196 (1954), J. Arn. Arb. 44:294 (1963)

& Fl. Trop. East Africa. Legum. Papilion.

467. t. 65 (1971); Ohashi, Ginkgoana

1:184 (1973) & Fl. Camb. Laos Vietn.

27:121 (1994); Verdcourt, Man. New

Guinea Leg. 397 (1979); Pedley, Rev.

Handb. Fl. Ceylon 10:182 (1996);

Meibomia gangetica (L.) Kuntze, Rev.

Gen. 1:196 (1891); Hedysarum

gangeticum L., Sp. Pl. 746 (1753). Type:

Hedysarum no. 13 (LINN).

Selected specimens: Western Australia. Junction of

Broc kman R. & Calder R., 16°17’S 125°00’E, May

1983, Milewski (K, PERTH). Northern Territory.

Victoria River, Mueller (KK); 43 miles [69 km] ENE of

Pine Creek, Mar 1965, Lazarides & Adams 193 (CANB);

5 miles [8 km} 8 of Batchelor, Mar 1965, Muspratt 112

(AD, DNA, MEL, NSW), Queensland. NortH KENNEDY

District: Ayr, 19°34’S 147°24’E, Jun 1951, Kleinschmidt

K.145 (CANB); Port Curtis District: Gladstone, Dietrich

41 (MEL); Moreton District: near Sidling Creek, Petrie 18

miles [29 km] N of Brisbane, Mar 1931, Blake 2349 (BRI, K).

Distribution and habitat: Desmodium

gangeticum has a wide geographic range in the

Old World: Africa, Sri Lanka, India, Indo-

China and Malesia. In Australia it is found from

north-eastern New South Wales, through

eastern Queensland to the northern part of the

Northern Territory and the Kimberley region

of Western Australia. It 1s nowhere common

and little is known ofits ecology except that it

occurs in eucalypt communities; it may have

been introduced in some places. Map 36.

Affinities: Ohashi referred D. gangeticum with

D, velutinum to sect. Heteroloma Benth. though

the relationship between the two is not close.

The nearest relative of D. gangeticum is

possibly D. pryonii DC. which is restricted to

Sri Lanka and the extreme south of Peninsular India.

24, Desmodium rhytidophyllum F. Muell. ex

Benth., Fl. Austr. 2:233 (1864);

Meibomia rhytidophylla (F. Muell. ex

Benth.) Kuntze, Rev. Gen. 1:198 (1891).

Types: Port Jackson, Brown (syn:K);

Parramatta, Woolls (syn:K, MEL);

between Burnett and Dawson, Mueller

(syn:K_); Queensland (cited by Bentham

as “near Rockhampton’), Dallachry (syn: K),

Trailing perennial herb; branchlets with

indumentum of brown spreading hairs c. 0.5

mim long; stipules deltoid or ovate acuminate,

3-5 mm long, 1-2 mm wide, densely hairy

outside, glabrous, striate inside. Leaves 3-

foliolate; petiole with indumentum of branches,

8—30 mm long, rachis 3—9 mm long, petiolules

1—2 mm long, stipels 0.7—1.7 mm long; terminal

leaflets rhomboid (rarely obovate or orbicular),

obtuse or rarely emarginate, discolorous with

+ dense spreading hairs on both surfaces,

becoming sparser on upper surface, 17—45(—

55) mm long, (10--)15—30 mm wide, (1—)1.5-

1.8(2.1) times longer than wide, lateral ones

smaller and often oblique 14-35(—40) mm long,

10-25 mm wide, 1.31.8 times longer than

wide. Inflorescence terminal, to 25 cm long,

rather open, axis with dense indumentum of

hooked hairs; flowers 1n fascicles of 2 or rarely

3; primary bracts 2-3 mm long, secondary

bracts c. | mm long; pedicels 2-4 mm long.

Flowers pink or purplish: calyx 4-lobed, 3-3.3

mm long, the tube hirsute 1-1.3 mm long, the

lobes c. 2 mm long, the lower and lateral ones

subulate, the upper wider, minutely bifid at the

tip; corolla standard obovate, sometimes

shortly clawed, 4.3-5.3 mm long, 44.5 mm

wide, wings 3.7 mm long, 1.5 mm wide,

somewhat auriculate, ona claw c. 1.5 mm long,

keel petals 3.5—4 mm long, 1.5—2 mm wide on

a claw 1.5—2 mm long; stamens diadelphous;

ovary with indumentum of hooked hairs. Pod

straight with up to 6 articles, upper suture and

lower margin incised, the lower slightly more

than the upper, the isthmus c. % width of

articles; articles transversely ovate, 3.5-4 mm

long, 2.5—3 mm wide, reticulately veined with

moderately dense indumentum of uncinate

hairs. Mature seeds not seen.

Selected specinens: Papua New Guinea, CENTRAL

District: Port Moresby subdistrict, Lake Myola swamp

grassland, 9°9’S 147° 43°E, Sep 1973, Croft & Lelean

NGF 34642 (BRI). Northern Territory. Gimbat, source

of South Alligator River, 13°45’E 132°48’E, Russelj-

Smith 734 (DNA). Queensland. Coox District: Tolga,

17°13’S 145°29°E, Apr 1962, McKee 9405 (CANB, K,

NSW); LeicHHARDT District: Blackdown Tableland, c.

23°50'S 149°00’E, Apr 1971, Henderson et al. 671 (BRI,

250

K, MEL); Burnetr District: Eidsvold, 25°22’S 151°07’E,

Apr 1963, McKee 10219 (CANB, NSW). New South

Wales. Brunswick Heads, Feb 1971, O’Hara & Coveny

3476 (K, NSW); Colo River Bridge, c. 12 miles [19 km]

N of Windsor, Mar 1967, Constable 7347 (K, NSW);

Huskisson, Jervis Bay, Feb 1941, £.C. AfcDonaild 143 (Kk).

Distribution and habitat. The species ranges

from the Atherton Tablelands of north

Queensland through coastal and subcoastal

areas to the South Coast District of New South

Wales, with isolated occurrences in the

Northern Territory and in Papua New Guinea.

It is often a prominent member of the ground

layer vegetation of open eucalypt communities,

usually on well drained soils. It also occurs in

New Caledonia. Map 37.

Affinities: AS Bentham noted in his protologue

D. rhytidophyllum has some affinity with D.

varians, but the relationship is not close. It is

however, close to D. tenax (which Verdcourt

(1979) treated as a subspecies of D.

rhytidophyllum) and D. macrocarpum but all

are rather isolated among species of sect.

Heteroloma Benth.

25. Desmodium tenax Schindl., Rep. sp. nov.

reg. veg. 21:10 (1925). Type: Rockingham

Bay, Dallachy (n.v.).

D. rhytidophyllum subsp. acutifoliolum

Verdc., Kew Bull. 32:250 (1977) & Man.

New Guinea Leg. 406 (1979). Type:

Papua New Guinea. WESTERN DISTRICT:

Moreshead Subdistrict: Wassi Kussi

River, Arufi, Jul 1968, Henty & Katik

NGF 38667 (holo: K).

Sprawling shrub to c. 1 m tall; branchlets with

indumentum of brown spreading hairs similar

to those of D. rhytidopyhyllum; stipules

lanceolate acute, to c. 6 mm long, densely hairy

outside, glabrous inside. Leaves trifoliolate;

petiole 10-40 mm long, rachis 8—20 mm long;

leaflets ovate with a long acute apex, upper

surface with sparse, appressed to ascending

hairs, lower surface with moderately dense

usually ascending hairs; terminal leaflet 65-105

mm long, 40-55 mm wide, 1.5—2.3 times longer

than wide, laterals less attenuate, smaller, 42—

60 mm long, 22—35 mm wide, 1.3-2 times

longer than wide; pulvinuses 22.5 mm long,

stipels 1.5-2.5 mm long. Inflorescences

Austrobaileya 5(2): 209-261 (1999)

terminal and in upper axils, to 200 mm long,

rachis with indumentum of hooked hairs;

fascicles of usually 2 flowers, primary bract 2—

3 mm long, secondary c. 1.5 mm, both

deciduous, bracteoles absent; pedicels to c. 3

mm long. Flowers pink or purple, similar to

those of D. rhytidophyllum though perhaps

standard smaller. Pods indented on both upper

and lower sutures, the lower more deeply,

isthmus about '/, width of pod, of up to 6

articles, each triangular, 5.5—8 mm long, 3.44

mm wide, reticulately veined with sparse

uncinate hairs; seed (only 1 measured) 3.5 mm

long, 2.2 mm wide.

Selected Specimens: Queensland. Cook DISTRICT:

Lockerbie Scrub, 10°47’S 142°29°E, Feb 1990, Forster

PIF 6325 (BRD); Iron Range [12°44’S 143°17’E] Jun

1948, Brass 19320 (BRI, CANB, K); Mcllwraith Ra.,

13°42’S 143°18’E, Jun 1992, Forster PIF 10247 (BRD;

NorTH KENNEDY District: Mt Fox [18°S1’S 145°48’E],

Nov 1949, Clemens s.n. [AQ 417008] (BRI).

Distribution and habitat: The species ranges

from southern New Guinea to Mt Fox, a little

north of Townsville, Queensland. It has been

collected in eucalypt-Melaleuca communities,

but is evidently rare. Map 38.

Affinities: As Schindler noted in the protologue

of the species, it differs from D. rhytidophyllum

on its larger acute leaflets and larger pods, and

they are obviously closely related.

26. Desmodium macrocarpum Domin,

Biblioth. Bot. 89:214 (1926). Type: in

xerodrymio ad opp. Jericho, Mar 1910,

Domin 4744 (holo: PR 527366).

Subshrub woody at the base, with ascending

stems to c. 0.5 m tall; branchlets with rather

dense indumentum of erect, slightly uncinate,

stiff hairs to 0.5 mm long; stipules ovate,

acuminate, 5-6 mm long, c. 3 mm wide,

longitudinally veined, pubescent. Leaves 1- or

rarely 3-foliolate; petiole 25-50 mm long;

leaflets ovate or rhombic (or laterals oblong-

orbicular), obtuse, mucronulate at the apex,

truncate or subcordate at the base, with

moderately dense short stiff hats on both upper

and lower surfaces; terminal leaflets 25-30 mm

long, 20-40 mm wide, 1.2—1.6 times longer

than wide, laterals about half as large;

pulvinuses 2—3 mm long, stipels 2.5-4 mm

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 251

long, setaceous. Inflorescences terminal and in

upper axils, to c. 200 mm long, rachis with

indumentum similar to that of branchlets;

fascicles of 2 flowers; primary bracts similar

to stipules, 3-5 mm long, 1.5 mm wide,

secondary bracts absent or early deciduous;

bracteoles absent; pedicels 2—3 mm long.

Flowers white or pale mauve; calyx 4-lobed,

c.4mm long, pubescent with short stiff hairs,

lobed to about the middle, the upper wider than

the others, notched at the apex; standard

orbicular, tapered to the base but not distinctly

clawed, 6—6.5 mm long, 6.5—7 mm wide; wing

6-6.5 mm long, 2—3 mm wide including a

distinct claw; keel petals about as long as the

wings, slightly wider; stamens diadelphous;

ovary with dense indumentum of short uncinate

hairs. Pod straight with up to 4 articles, upper

suture straight, lower margin incised to c. “/s

width of pod; articles + triangular, 9-10 mm

long, 5-6 mm wide, with dense indumentum

of short stiff uncinate hairs. Seeds brown, +

oblong, 5—6 mm long, 3~—3.5 mm wide, with a

distinct rim aril.

Selected specimens: Queensland. NoRTH KENNEDY

District: 40-mile scrub, 18°15’S 144°45’E, Feb 1972,

Irvine 165 (BRI, QRS); LeicHHarpt District: ‘Redlands’

5 km W of Balfes Creek, [approx 20°15’S 145°55’E],

Apr 1977, Rebgetz 355 (BRI); ‘Kooralbyn’ c. 24 km SSW

of Duaringa, 23°57’°S 149°37’°E, Mar 1984, Anderson

3660 & 3661 (BRD; Burnetr District: ‘Narayen’ near

Mundubbera, May 1988, Hacker JBH 870 (BRD.

Distribution and habitat: Desmodium

macrocarpum 1S an uncommon plant known

from several widely separated localities in

eastern Queensland. It is found on sandy or

loamy soil, sometimes shallow, usually in

communities of Eucalyptus crebra F. Muell,

sometimes with Acacia shirleyi Maiden. Map 39.

Affinities: As Domin stated the species is

related to D. rhytidophyllum, but has usually

unifoliolate leaves, stiffer hairs and much larger

pod articles,

27. Desmodium gunnii Benth. ex J.D. Hook.,

Fl. Tasmanicae 1:101 (1856); .D. varians

var. gunnii (Benth. ex J.D. Hook.) Benth.,

Fl, Austr. 2:233 (1864). Type: Tasmania,

Gunn 243 (lecto: K, see below; isolecto: BM).

Sprawling or somewhat ascending perennial,

sometimes rooting at the nodes; branchlets

terete, glabrous or with a few appressed hairs;

stipules triangular, striate, 2—4 mm long, 1—2.5

mm wide at base. Leaves trifoliolate, petioles

10-30 mm long, rachis minute or to 1 mm long,

petiolules (i.e. pulvinuses) 0.6—-1 mm long,

stipels setaceous, 0.5—2 mm long, those of

lateral leaflets longer than those of terminal

one; leaflets cuneate and truncate at the apex,

or rarely obovate and obtuse, the terminal 6—

20 mm long, 7-11 mm wide, 0.8—1.5 times

longer than wide, the laterals usually smaller,

6-18 mm long, 6.5—13 mm wide, 0.8—1.6 times

longer than wide, subglabrous, a few hairs on

veins on lower surface. Inflorescences termina]

or in the upper axils, to 16 cm long, open,

flowers solitary, bracts persistent, stipule-like

to 1.7 mm long, pedicels with indumentum of

hooked hairs, 1.5~7 mm long, bracteoles none.

Flowers: calyx tube campanulate 1~1.2 mm

long, the lobes broad 0.7—1 mm long; corolla

pink or lavender, standard 3—3.3 mm long, the

other petals longer, c. 3.5 mm long; stamens

diadelphous; ovary appressed pubescent. Pods

with up to 6 articles, the upper suture indented

between the articles, the lower more deeply so;

articles oblong, 4—5 mm long, 2—2.5 mm wide,

with dense indumentum of spreading hooked

hairs. Mature seeds not seen.

Selected specimens: Queensland. Port Curtis DISTRICT:

Many Peaks Range, 24°10’S IS1°L7°E, Jun 1977, Crisp

2752 (CANB); Gladstone, Dietrich 215 (MEL); MoreToN

District: Tamborine Mtn, Mar 1947, Clemens (K). New

South Wales. Brindabella Range, 35°19’S 148°50’E, Jan

1977, Crisp et al. 2471 (CANB); Shoalwater River Gorge,

34°48’°S 150°12’E, Feb 1974, Pullen 8794 (CANB).

Victoria. Mallacoota Inlet, Dec 1969, Beauglehole 32372

& Finck (MEL); Sherbrooke, 37°54’°S8 145°21E, Mar

1977, Gullan 62 & Opie (MEL).

Distribution and habitat: The species ranges

from Papua New Guinea (one collection)

through coastal parts of mainland eastern

Australia to Tasmania, commonly in eucalypt

open forest, usually in shady situations. It also

occurs in New Caledonia. Map 40.

Affinities: Desmodium gunnii is close to D.

varians to which Bentham referred it as a

variety. It differs in the structure of its pod and

its extremely short leaf rachis. Leaflets of D.

varians, when extremely short, approach those

of D. gunnii in size and shape, but whereas the

leaves of D. gunnii are subdigitate with the leaf

252

rachis never exceeding | mm, those of D.

varians are distinctly pinnate with the leaf

rachis at least 1 mm longer and usually

considerably more.

Typification: Hooker cited Gunn 243 as the

type. In herb. K two sheets have been

segregated as type material, one from

Herbarium Hookerianum, the other from

Herbarium Benthamianum. Thirty-two

fragments are mounted on the sheets.

Annotations and labels indicate that, in part,

they represent Gunn 243, but more than one

collection (including some by Milligan) is

represented. Unfortunately five of the

fragments are of D. varians. Despite the mixed

material, Hooker’s description applies to D.

gunnii. | have therefore lectotypified the name

by the fragments mounted on the sheet

‘Herbarium Hookerianum 1867’, excepting two

sizeable fragments of D. varians on the sheet.

This sheet has pinned to it a note and a field

label both in Gunn’s hand identifying it as his

n. 243. The isolectotype at BM consists of six

fragments all from Circular Head; none is D.

varians.

28. Desmodium varians (Labill.) G. Don, Gen.

Hist. Dich. Plants 2:298 (1832); Benth.,

Fl. Austr. 2:233 (1861); Meibomia

varians (Labill.) Kuntze, Rev. Gen. 1:198

(1891); Hedysarum varians Labill., Sert.

Austro-cal. 71 t. 71 (1824). Type: none

designated.

Desmodium varians var. angustifolium

Benth., Fl. Austr. 2:233 (1864).

Desmodium spartioides DC., Prodr.

2:337 (1825). Type: ex herb. Lambert

(G—DC, n.v., microfiche: BRI).

Desmodium dietrichiae Dormin, Biblioth.

Bot. 89(3): 213 (1926). Type: prope

Brisbane River, Australia or. Dietrich

1450 (lecto: PR 527357, chosen here).

Sprawling perennial or tufted annual with stems

arising from perennial rootstock; branchlets

terete, glabrous or with a few appressed hairs;

stipules deltoid or ovate acuminate, 2—5.5 mm

long. Leaves trifoliolate, petiole S—10 mm long,

rachis 2—4.5 mm long; leaflets lanceolate,

oblong or occasionally oblanceolate, rounded

Austrobaileya 5(2): 209-261 (1999)

or cuneate at the base, obtuse or slightly retuse

at the apex, usually a few appressed hairs on

the veins beneath; terminal leaflet 5-35 mm

long, 4-5.5(—7) mm wide, 1—8.5 times longer

than wide, lateral leaflets smaller, 3.5—26 mm

long, 3—4.5(—6) mm wide, |.2—8 times longer

than wide; pulvinuses 0.51.5 mm long, stipels

0.4—1 mm long. Inflorescence terminal to c.

[5cm long in flower, elongating in fruit, the

rachis glabrous except for some uncinate hairs

near the tip, fascicles of 1—3 flowers, subtended

by primary bract 1.5—3.5 mm long, deciduous;

secondary bract usually 0.7—1 mm long,

deciduous; bracteoles absent; pedicels 3—9 mm

long with indumentum of uncinate hairs.

Flowers: calyx 4-lobed, covered with short

uncinate and longer straight hairs, tube 1.2—

1.5 mm long, the lobes all of about equal length,

1,2—-1.6 mm long, the upper + entire or divided

to about the middle; corolla pink or purplish;

standard orbicular, about 6 mm long and wide,

wings about 5 mm long, 2.5 mm wide, with

claw c. 2 mm long, keel petals somewhat

shorter than the wings. Pods with 4—6 articles,

the upper suture thickened, somewhat undulate,

the lower deeply indented, the isthmus '4A—%

width of pod; articles 3.5-4.2 mm long, 2.7-

3.8 mm wide, with moderate to dense

indumentum of uncinate hairs; seeds about 2.5

mm long, 1.6 mm wide with a distinct rim aril.

Selected specimens: New Caledonia. Wagap, Miellard

366 (K). Queensland, BuRKE District: 13 km N of turn-

off to ‘Gregory Springs’, 19°40’S 144°14’°EB, May 1988,

Pullen 11248 (CANB); LeicuHarpr District: 8 miles [13

km] S of Springsure, Oct 1964, Addams 1391 (BRI,

CANB, K); Moreton District: Blunder Creek near

Brisbane, Oct 1930, Hubbard 4713 (BRI, K). New South

Wales. Yamba, 29°27’S 153°22’E, Feb 1973, Tindale

2038 (NSW); 30 miles [48 km] S of Tamworth, 31°31’S

150°51’E, Feb 1941, Roe R61 (CANB); Canberra, Nov

1960, AfcKee 7672 (CANB, K). Victoria. Dandenong

Ranges, Belgrave, Dec 1932, Willis (MEL); Upper

Turnback Creek, 37°06’S 148°22’E, Feb 1980, MHa/s/ 352

(MEL). Tasmania. South Esk, in 1887, Oakden (MEL);

without definite locality, Adve/ler (MEL),

Distribution and habitat: The species occurs

over a wide area of eastern Australia, ranging

from Tasmania through coastal districts of

Victoria and subcoastal and coastal districts of

New South Wales to north-central Queensland,

usually on well drained soils in eucalypt

communities. It also occurs in New Caledonia.

Map 41.

baletada eee aya al a py le ee ae a ea iy one Sy PraEre DEPRTY, parE, “lint aitlieduan wind intuniidsteta

Sibert Be Cee red Ses tre a eae and bol Pee Dra oe cake a ee eT ee ee ea

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 253

Affinities: Desmodium varians is closely

related to D. gunnii which Bentham treated as

a variety of D. varians. The two are sympatric

in many coastal areas but D. gunnii usually

occurs in moister shaded situations. The species

is variable and there are two different growth

forms. In coastal areas the plant usually sprawls

among grass but in inland areas which often

have a marked dry season it often forms

multistemmed tufts from a woody base. The

latter may be merely first growth in response

to rain, but possibly Bentham should be

followed and two taxa recognised .

29. Desmodium tortuosum (Sw.) DC., Prodr.

2:332 (1825); Schubert, Flora Trop. East

Africa, Legum. Papilion. 474 (1971);

Verdcourt, Kirkia 9:256 (1974) & Man.

New Guinea Leg. 408 (1979); Long &

Lakela, Fl. Trop. Florida 487 (1971);

Correll & Correll, Fl. Baham. Arch. 64

(1982); Liogier, Fl. Espafiola 3:197

(1985); Howard, Fl. Lesser Antilles

4(1):482 (1988); Green, Fl. Austral.

49:179 (1994); Pedley, Rev. Handb. F.

Ceylon 10:191 (1996); Meibomia

tortuosa (Sw.) Kuntze, Rev. Gen. 1:198

(1891); Hedysarum tortuosum Sw.,

Prodr, Veg. Ind. Occ. 107 (1788). Type:

Jamaica, Swartz (holo: 8S, fide Schubert,

Howard, n.v.).

Desmodium purpureum (Miller) Fawcett &

Rendle, Fl. Jamaica 4:36 (1920), non

Hook. & Arn. (1832); Hedysarum

purpureum Miller, Gard, Dict. ed. 8 no.

6 (1768). Type: Mexico, Vera Cruz, in

1730, Houston (holo: BM).

Selected specimens: Western Australia, Mitchell

Piateau, 14°49’S 125°51°E, May 1978, Kenneally 6801

(K, PERTH). Northern Territory. Darwin, Oct 1946,

Blake 17311 (BRI, K); Katherine, 14°28°S 132°16’E, Apr

1977, Pullen 10607 (CANB) & Dunlop 4182 (DNA).

Queensland. Cook District: Yorke {[s., 9°45’S 143°24’E,

Oct 1981, Clarkson 3972 (BRI, K, MBA); Moreton

District: Gatton, Apr 1947, Everist 2965 (BRI, K). New

South Wales. Braidwood District, Jan 1885, Bauerlen

393 (MEL).

Distribution and habitat: Desmodium

fortuosum 1s a native of subtropical and tropical

America now naturalised in Africa and Asia. It

was probably introduced to Australia as a

green-manure crop but is widely (though not

commonly) naturalised in high rainfall areas

in eastern coastal areas and in the north-west,

usually on roadsides. Map 42.

Affinities: Because of its twisted pods D.

tortuosum is unlikely to be confused with any

other species known from Australia, but it 1s

closely related to the tropical American D.

procumbens (Mill.) Hitche. and the African D.

ospriostreblum Chiov. which is probably only

a variant of D. procumbens. The terminal article

of the pods of Clarkson 3972 1s about twice as

large as the other articles, an attribute which

allegedly distinguishes D. ospriostreblum from

D. procumbens (Schubert 1971).

30. Desmodium uncinatum (Jacq.) DC.,

Prodr. 2:331 (1825); Schubert, Field

Mus. Nat. Hist. Publ. Bot. 13(3):412

(1943); Fosberg, Micronesica 4:257

(1968), pro syn.; Verdcourt, Man. New

Guinea Leg. 411 (1979); Meibomia

uncinata (Jacg.) Kuntze, Rev. Gen. 1:197

(1891); Hedpsarum uncinatum Jacq., PI.

Hort. Schoenbr. 3:27 t.298 (1798). Type:

‘Crescit in Caracas’ (W?,n.v.). |

Selected specimens: Queensland, WipE Bay District: 3

km NE of Kin Kin, 26°15’S 152°54°E, May 1976, Jacobs

2520 & Rodd (K, NSW); Moreton District: cultivated

CSIRO Experimental Farm, Samford, Apr 1960, (BRI,

Kk). New South Wales. 4.6 miles [7.4 km] from Sheep

Station sign on Lynch’s Creek road, Wiangaree State

Forest, 28°23°S 153°06’E, May 1973, Tindale 2114

(NSW).

Distribution and habitat; Desmodium

uncinatum (“silver-leaf Desmodium’) was

deliberately introduced into Australia as a

forage crop and has become naturalised in high

rainfall areas of south-eastern Queensland and

north-eastern New South Wales, usually on

loamy volcanic soils. Map 43.

Affinities: Schubert, Fosberg and and Ohashi

(1990) commented on the confusion of the

identities of D. wnctnatum, D. intortum and D.

limense Hook. Contrary to Fosberg’s treatment

of D. uncinatum as a synonym of D. tntortum,

Schubert’s and Verdcourt’s recognition of them

as distinct species is reasonable. The

differences given in the key hold for South

American as well as Australian and New

Guinea specimens. Ohashi (1990) applied the

: 209-261 (1999)

Austrobaileya 5(2)

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33. Desmodium neniorosum

32. Desmodium strigillosum

31. Desmodium heterocarpon

Var. Si1gOSuHI

34, Desmodium tiviense

36. Desmodium gangeticum

35, Desniodium velutiniuni

38. Desmodium tenax

39, Desmodium macrocarpum

42, Desmodium torttosiup

37. Desmodium rhytidophyllum

40, Desmodium gunnii

41, Desmodium varians

45, Desmodium tneanii

44, Desmodium intortuni

43, Desmodium uncinatuii

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 255

name D. sandwicense to plants growing in

Hawai’1. The correct name for the species (if it

is different from D. uncinatum) 1s the earlier

D. pilosiusculum DC.

31. Desmodium intortum (Mill.) Urban,

symbol. Antill. 8:292 (1920); Schubert,

Field Mus. Nat. Hist. Publ. Bot.

13(3):427 (1943); Verdcourt, Man. New

Guinea Leg. 400 (1979); McVaugh, FI.

Novo-Galiciana 5:470 (1987); Ohashi,

Man. Fl. Pl. Hawai’i 667 (1990);

Meibomia intorta (Mill.) Blake, Bot.

Gaz. 78:286 (1924); Hedysarum intortum

Mill., Gard. Dict. ed. 8. no. 11 (1788).

Type: Jamaica, Houston (2BM, n.v.)

Selected specimens: Queensland. Moreton District:

Nambour, Jan 1972, Everist [AQ 4279] (BRI, K);

Brisbane Botanic Gardens, Dec 1975, Verdcourt s.n. (K).

New South Wales. Wiangaree State Forest, 28°23’S

152°06’°E, May 1973, Tindale 2116 (NSW).

Distribution and habitat: Like D. uncinatium,

D. intortum (green-leaf Desmodium’) was

introduced into Australia as a forage plant and

has become naturalised, though not as

frequently as D. uncinatum. Map 44.

Affinities: The affinities of the species are

commented on under D. uncinatum.

32. Desmodium incanum DC., Prodr. 2:332

(1825); Benth. in Martius, Fl. Bras. 15:98

(1859) & synonyms cited; Nicolson,

Taxon 27:365—370 (1978); McVaugh, FI.

Novo-Galiciana 5:468 (1987); Green, FI.

Austral. 49: 179 (1994); Hedysarum

incanum Sw., Nov. gen. 107 (1788), nom.

illeg. Type: Plumier (ed. Burman), Pl.

Amer. 140. 149. f£.1. (1757) “Hedysarum

foliis ternatis, ovatis, floribus specatis”’.

Desmodium frutescens Schindl., Rep. sp.

nov. reg. veg. 21:9 (1925) & synonyms

cited. Type: Jacq., Hort. Vindob. 3:47. t.

89 (1776) “Hedysarum frutescens Jacq.

non L.”

Desmodium canum Schinz & Thellung,

Mem. Soc. Neuchatel. Sci. Nat. 5:371

(1913); Schubert, Fl. Trop. East Africa. are subtended by foliage leaves, whereas in

Legum. Papilion. 456 (1974); Verdcourt,

Man. New Guinea Leg. 394 (1979);

Hedysarum canum Gmel., Syst. Veg.

2:1125 (1792). Type: as for D. incanum

DC,

Specimens; Queensland. MoORETON DISTRICT:

Indooroopilly, Brisbane 27°31°S 152°59’E, Mar 1994,

Pedley 5678 (BRI); Scarborough, Redcitffe City, 27°12S

153°07°E, May 1994, Galagher s.n. (BRI).

Distribution and habitat: This species is a

native of tropical America, often used as a cover

crop and naturalised in many countries. It

occurs as a weed in a lawn at Indooroopilly

and on a walk path at Redcliffe, both in the

extreme south-east of Queensland. Prior to its

collection in the field it had been grown

experimentally under glass in the vicinity of

Brisbane at least as early as 1964.

Affinities: The plant resembles D. heterocarpon,

but has an open inflorescence and its leaves

have more primary lateral veins. Its fused stipules

distinguish it from other species. Map 45.

Note: Nicholson, in his detailed discussion of

the complex nomenclature of D. incanum and

D. canum, pointed out that both Bentham and

Schindler considered D. ancistrocarpum

(Ledebour) DC. (based on Hedysaruin

ancistrocarpum Ledebour) to belong to this

species. In the interest of stability however, he

refrained from reviving the earlier name. IfD.

ancistrocarpum is indeed conspecific with D.

incanum it is to be expected in the interest of

continuing stability that the name D.

ancistrocarpum be rejected in favour of D.

Incanum.,

Phyllodium Desv,, J. Bot. Agric. 1:123.1.5.f.

24 (1819); Benth. in Miq., Pl. Jungh. 217

(1852); Ohashi, Ginkgoana 1:260 (1973);

Dicerma sect. Phyllodium (Desv.) Benth.

in Benth. & J.D. Hook., Genera PI. 1:519

(1865); Desmodium subg. Phylodium

(Desv.) Baker in J.D. Hook., Fl. Brit.

India 2:162 (1876). Type: Phyllodium

pulchellum (L.) Desv.

Phyllodium is a genus allied to Dendrolobium.

In Dendrolobium flowers are arranged in

fascicles or short (rarely long) racemes which

Phyllodium fascicles of flowers are usually

subtended and hidden by ‘foliar bracts’. These

256

foliar bracts are modified foliage leaves: the

terminal leaflet is lost or reduced to a terminal

seta and the lateral leaflets are much reduced

in size; stipules and stipels are retained. In other

Desmodieae flowers are subtended by stipule-

like bracts not at all like foliage leaves in colour,

texture or shape.

Ohashi (1973) described subg. Prainea

to accommodate P. insigne (Prain) Schindl. in

which foliar bracts are only sporadically

developed. In the field, plants of P hackeri sp.

nov. produce foliar bracts at the bases of

inflorescences only, but plants grown under

Austrobaileya 5(2); 209-261 (1999)

glass (Hacker 389A, BRI) have foliar bracts at

most nodes of the inflorescence. Thus, since

the production of foliar bracts is largely

controlled by environmental factors, their

presence or absence is not a sound basis for

the recognition of a subgenus. Consequently,

subg. Prainea is ignored.

In Australia there are three species of

Phyllodium; namely, P. pulchellum with an ill-

defined variant described here as a variety, a

well-defined species confined to northern

Queensland (P. hackeri) and a third, known

only from one specimen.

Key to species

1. Pedicel 7-12 mm long; flowers large: calyx c. 5 mm long, standard 10-

10.5 mm long and 5—6 mm wide pods with up to 5 articles, each 5.5—7 x

SP di pek sell Reaee are Boxcar on 2 be wee Date dc oo oe

Pedicel to 3 mm long; flowers small: calyx to 3 mm long, standard to 7 mm

long and 4 mm wide; pods with up to 3 articles, each4 x 3.5mm ..............44. 2.

2. Foliar bracts subtending fascicles of flowers at anthesis; calyx with upper

lobe narrowly ovate, lower lobe 1.5—2 mm long, longer than others.... 2. P. pulchellum

Foliar bracts a few at base of inflorescence at anthesis; calyx with upper

lobe broadly ovate, obtuse, lower lobe to c. | mm long, about as long as

EPPELIOWE=<.. 4. ue patels Cork od ede Gecko

1. Phyllodium hackeri Pedley, sp. nov. forsan

affinis P insigne (Prain) Schindler

bracteis foliatis + caducis praedito sed a

quo foliolis infra non dense lanatis,

interdum parvioribus differt, sed similis

P. pulchello (L.) Desv. a quo bracteis

foliolatis + caducis, pedicellis longioribus

(7-12 mm), floribus grandioribus,

articulis leguminis multo grandioribus

differt: Typus: Queensland. Cook

DistricT: between Irvinebank and

Petford, c. 17°20’S 145°00’E, February

1980, JR. Clarkson 2832A (holo:BRI;

iso:K, MBA, PERTH, QRS).

Phyllodium sp. (Montalbion H.S. McKee

9430) Pedley in Henderson, Queensland

Plants: Names & Distribution: 84 (1997).

Desmodium megaphyllum auct. non Zoll.;

Knapp-van Meeuwen, Blumea 12:16 (1964).

Shrub to 1.5 m tall; branchlets angular with

moderately dense appressed hairs 0.4 mm long.

Leaves 3-foliolate; stipules subulate, broad at

the base, c. 5 mm long; petioles 12-30 mm

long, rachis 15—25 mm long; leaflets ovate or

elliptic, rounded at the base, obtuse or slightly

retuse, mucronulate, sparse minute uncinate

hairs and appressed hairs on veins above,

minute (c. 0.2 mm long) appressed hairs on

lower surface; terminal leaflet 75—105 mm

long, 30-50 mm wide, 2—2.5 times longer than

wide; lateral leaflets smaller, 60-80 mm long,

30—40 mm wide; pulvinuses 2.3—2.8 mm long,

stipels 1.5—2.2 mm long; foliar bracts (often

absent) bifoliolate, the terminal leaflet reduced

to a bristle, leaflets with indumentum of foliage

leaves, petiole 3-6 mm long, petiolules c. 2.5

mm long, stipels c. 1.5 mm long; leaflets ovate,

attaining 25 mm long, 15—20 mm wide. Flowers

in fascicles of up to 16, in axillary or terminal

racemes, each fascicle subtended by a foliage

leaf or a foliar bract or without subtending leaf

or foliar bract; bract c. 1 mm long; pedicels

slender, appressed pubescent 7-12 mm long.

Flowers: calyx 4.7-5.1 mm long, with minute

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision Zor

uncinate hairs and appressed white hairs c. 0.3

mm long , tube 2.2 mm long, upper and lower

lobes 2.52.9 mm long, laterals about as long

as the tube; corolla cream; standard oblong,

obtuse or slightly retuse at apex, 10—10.5 mm

long, 5-6 mm wide; wings 7,3-8.5 mm long,

auriculate at base, with claw c. 2 mm long; Keel

petals auriculate, 85-9 mm long, 4-5 mm wide

on claw 2.5-3 mm long; ovary sparsely

pubescent. Pod with upper suture straight,

lower somewhat indented, with up to 5 articles

each 5.5—7 mm long, c. 5 mm wide, reticulately

veined, glabrous except for appressed hairs on

each suture; seed 3—3.7 mm x 2.5—3.2 mm with

a small rim aril.

Specimens: Queensland, Cook District: Lankelly Creek,

Mcilwraith Ra., 13°53’°S 143°14’E, Jun 1992, Forster

PIF 10365 & Tucker (BRI); Montalbion on Petford-

Herberton road [17°24’S 145°09’E}, Apr 1962, McKee

9430 (BRI, CANB); Irvinebank-Emuford, 17°25’S

145°00’E, Jan 1972, Hyland 5814 (BRI, QRS); cultivated

from seeds collected on Chillagoe road, 17°25’S

145°04’°E, Apr 1987, Hacker BH389A (BRI).

Distribution and habitat: Phyllodium hackeri

has a limited distribution in northern

Queensland where it occurs on sand and

shallow rocky soil. Map 46.

Affinities: On account of its frequent lack of

foliar bracts P hackeri might be considered

related to P. insigne or P. pulchellum var.

glabrius. The relationship to other species ts,

however, not close and P. hackeri 1s rather

isolated in the genus. Fascicles of flowers

sometimes subtended by reduced foliage leaves

in P. hackeri indicate that Phyllodium and

Dendrolobium are closely related.

Etymology: Vhe species is named to honour

Dr J.B. Hacker, CSIRO, Brisbane who first

brought the species to my attention. Plants

stown in the glasshouse by Dr Hacker

demonstrated that the production of foliar

bracts by individual plants varied during its

srowing period. |

2. Phyllodium pulchellum (L.) Desv., J. Bot.

Agric. 1:124t. 5. £24 (1813), Mem. Soc.

Linn. Paris 4:324 (1825); Benth. in Miq.,

Pl. Jungh. 217 (1852); Schindl., Rep. sp.

nov. reg. veg. 20:270 (1924): Ohashi,

Ginkgoana 1:276 (1973); Pedley, Rev.

Handb. Fl. Ceylon 10:165 (1996);

Dicerma pulchellum (L.) DC., Ann. Sci.

Nat. 4:236 (1825), Prody. 2:239 (1825),

Mem. Leg. 328 (1826); Desmodium

pulchellum (L.) Benth., Fl. Hongkong 83

(1961), Fl. Austr. 2:231 (1864); van

Meeuwen, Reinwardtia 6:256 (1962); Liu

& Chuang, Taiwania 8:89 t. 16 (1962);

Verdcourt, Man. New Guinea Leg. 404

(1979); Meibomia pulchella (L). Kuntze,

Rev. Gen. 1:197 (1891); Hedysarum

pulchellum L., Sp. Pl. 747 (1753). Type:

Herb. Linn. No 921.24 (LINN, lectotype

fide Pedley in Turland & Jarvis (1997).

Two varieties are recognised in Australia.

2a. Phyllodium pulchellum (L.) Desv. var.

pulchellum.

Foliar bracts always present, persisting until

pods are produced; articles of pod glabrous or

appressed pubescent on faces, appressed

pubescent on sutures.

Specimens: Northern Territory. Keep River National

Park, 15°48°S 129°05’E, Apr 1982, King 72 (DNA);

Elcho Is., 11°57’S 135°43’E, Jul 1975, Maconochie 2115

(BRI, DNA, K). Queensland, Cook District: Iron Range,

12°43’°S 143°17°E, Jul 1988, Hacker JBH 892 (BRI);

Norts Kennepy District: ‘Maidavale’, E of Mingela, Apr

1991, Bean 2950 (BRI).

Distribution and habitat: Phyllodium

pulchellum var. pulchellum occurs only

sporadically in Australia and is nowhere

common. In the Northern Territory it occurs

on edge of coastal vine forests and in similar

situtations in Cape York Peninsula, but usually

among boulders on hillslopes in eucalypt

woodland in north-central Queensland. Map 47.

2b. Phyllodium pulchellum var. glabrius

Pedley var. nov., a P. pulchello vat.

pulchello bracteis foliatis in

inflorescentiis tantum sporadice

productis, articulis leguminis glabris vel

pilis aliquot in suturis ornatis differt.

Typus: Northern Territory. Muldiva

Creek, 14°01’S 130°43’E, February

1989, CR. Dunlop 976 & G. Leach (holo:

DNA; iso: BRI, K, MEL).

Foliar bracts only sporadically producted, often

absent on fruiting inflorescences; pod articles

glabrous with a few hairs on sutures.

258

Specimens: NORTHERN TERRITORY: Upper East Alligator

Region, 12°45’S 133°15’E, Apr 1988, Russell-Smith

5137 & Lucas (BRI, DNA, K, MEL); Leader Creek, Gunn

Point, 12°12’S 131°06’°E, Mar 1982, Wightman 226 (DNA).

Distribution and habitat: Phyllodium

pulchellum var. glabrius occurs in the northern

part of the Northern Territory in eucalypt

communities, usually on sandy soils. The plant

srows in a harsher environment than P

pulchellum var. pulchellum and the differences

between the two may be due to the differences

in habitat. Map 48.

3. Phyllodium sp. A

Shrub to 1 m; branchlets ribbed with

moderately dense indumentum of matted white

hairs 0.3 mm long. Leaves 3-foliolate; stipules

subulate, to 6 mm long, c. 3 mm wide at the

base; petioles 17 mm long, axis 20 mm long;

leaflets ovate, rounded at the base, obtuse

mucronulate, discolorous, sparse appressed

hairs on upper surface, sparse curved hairs on

lower, hairs 0.3 mm long; terminal leaflet 95—

105 mm long, 50—55 mm wide, lateral leaflets

smaller 55—70 mm long, 27-37 mm wide, about

10 lateral veins on each side of midrib all about

twice as long as wide; foliar bracts present only

at base of inflorescence, 2-foliolate, leaflets

broadly ovate, truncate at apex, 7—9 mm long,

6—7 mm wide, sparsely hairy on both surfaces.

Flowers in fascicles of up to 7 in axillary and

terminal racemes up to 20 cm long; the rachis

with indumentum similar to that of branchlets;

primary bracts stipule-like to 2 mm long,

pubescent at base and on margins; pedicels c.

3 mm long; bracteoles 0.6—0.8 mm long, 0.4

mm wide. Flowers: calyx c. 2.5 mm long with

moderatly dense appressed hairs; tube c. 1.5

min long, upper and lower lobes c, 1 mm long,

laterals c. 0.7 mm long; corolla cream; standard

oblong or obovate, slightly retuse at apex, 6—7

mm long, c. 4mm wide; wings 5.8—6 mm long,

including claw 1.7—2 mm long, 1—1.2 mm wide,

deeply auriculate; keel petals longer and wider

than the wings, c. 7 mm long, including claw c.

2 mm long, c. 2.5 mm wide; ovary laterally

compressed, glabrous on faces with fringe of

hairs on margins, with 1-3, usually 2, ovules.

Specimen: Northern Territory. Adelaide River, c. | km

downstream from Daly River road, Feb 1979, Rankin

1754 (BRI, DNA, K).

Austrobaileya 5(2): 209-261 (1999)

Distribution and habitat: The species is known

from the single specimen cited. There is no

information on its habitat. Map 49.

Affinities: Phyllodium pulchellum appears to

be the nearest relative of the species. Because

I know of the taxon from one specimen only, I

have not formally described it.

Tadehagi Ohashi, Ginkgoana 1:280 (1973) and

synonyms cited; Pedley, Rev. Hand. Fl. Ceylon

10:168 (1996). Type: Zadehagi triquetrum (L.)

Ohashi.

Tadehagi appears to be somewhat removed

from Desmodium and immediately related

genera but its relationship to Droogsmansia 1s

close and requires further investigation.

Ohashi included three species tn the genus each

of which he referred to a different subgenus.

He recorded 7! triguetrum subsp. triguetrum

from northern Australia and his distribution

map indicates that it extends into south-eastern

Queensland. He did not cite any specimen from

Australia and the only one I have seen bears

the label ‘Tropical East Australia, Evans’

(MEL). The record is a doubtful one. A species

related to 7. triqguetrum does however occur.

Tadehagi robustum Pedley, sp. nov.a 7

triquetro (L.) Ohashi bracteolis brevibus

et leguminibus plerumque articulis

paucioribus instructis; a 7. triquetro

subsp. triguetro foliolis minus quam 3-

flo longioribus quam latis, leguminis

articulis grandioribus in suturis non nisi

parce pubescentibus, aT triguetro subsp.

pseudotriguetro (DC.) Ohashi

leguminibus grandioribus in suturis non

nisi parce pubescentibus, a T. triguetro

subsp. aduriculato (DC.) Ohashi

leguminibus maturis neque suberosis

neque glabris, a T. triguetro subsp. alato

(DC.) Ohashi leguminibus glabris differt.

Typus: Western Australia. Mitchell

Plateau (c. 1 km N of CRA mining camp),

14°49°S 125°40°R, January 1982, BR.

Maslin 5085 (holo: PERTH; iso: MEL, K).

Shrub to 1.2 m tall, sometimes branched from

the base; branchlets strongly triquetrous, the

angles ribbed, glabrous or with appressed hairs

Pedley, L., Desmodium and related genera in Australia: a taxonomic revision 259

an £

46 eY Z & S . 47 3 5 g

at T e ps

oi * | as

( nue (

: a: ~ ros \

: j tae j —

) ate aa vb eo f ate za

2 ge at

49 90

46. Phyllodium hackeri

49, Phyllodium sp. A

to 0.7 mm long on the ribs; stipules 20-25 mm

long, 6-10 mm wide at the base. Leaves and

petiolar wing discolorous, coriaceous, glabrous

or with a few appressed hairs on major veins

beneath; petiole 17-40 mm long, wing 2.5—6

mm wide on each side of petiole, widest in

distal quarter; leaf blades oblong or ovate, acute

at apex, rounded or auriculate at base, 55—135

mm long, 25-60 mm wide, 2—2.5 times longer

than wide. Inflorescences terminal or in upper

axils, to 35 cm long; fascicles usually 2-

flowered; primary bracts 7-12 mm long

decreasing in size along rachis; pedicels to 3.5

mm long; secondary bracts and bracteoles c. |

mm long. Flowers: calyx with scattered stiff +

appressed white hairs and denser shorter

hooked erect ones; tube 1.7 mm long, upper

and lateral lobes c. 2.2 mm long, lower 2.7 mm

long. Corolla dark pink or red; standard

orbicular, 5 mm long, 6 mm wide, on stout claw

1.5 mm long; wings widest towards the tip, 4.5

mm long, 2.5 mm wide, auriculate at the base,

on a claw 1.8 mm long; keel petals slightly

shorter than the wings, 3.7 mm long, 2 mm

wide, auriculate at the base, on a claw 2.3 mm

long; ovary sparsely pubescent, moderately so

on the margins. Pods sessile, of 4—5 articles,

slightly indented on both sutures, the middle

47. Phyllodium pulchellum

var, pulchelluim

50. Tadehagi robustum

48, Phyllodium pulchellum var.

glabrius

articles c. 5 mm long, 8 mm wide, the proximal

and distal ones longer. Seeds oblongoid, with

a well developed rim aril, 3.6—3.7 mm long,

2.6—2,.7 mm wide.

Other specitnens (all Mitchell Plateau, all PERTH):

May 1978, Kenneally 6814; Jun 1976, Kenneally 4946;

Feb 1979, Beard 8327; Mar 1982, Keighery 4740.

Distribution and habitat: The plant is

apparently restricted to the Mitchell Plateau of

the Kimberley region of Western Australia

where it occurs in woodland, sometimes in

wetter areas, some seasonally flooded, and was

reported by one collector to be common. Map 50.

Affinities: Though the taxon could be referred

to T. triguetrum sensu lato it does not accord

well with the subspecies recognised by Ohashi.

It differs from subsp. triguetrum in having

leaflets less than 3 times longer than wide and

larger pod articles only sparsely hairy on

sutures; from subsp. pseudotriquetrum in its

larger pods only sparsely hairy on sutures; from

subsp. auriculatum in its mature pods being

neither corky nor glabrous; from subsp. a/atui

in its pods being not glabrous; and from all in

its short bracteoles and pods usually with fewer

articles.

260

Acknowledgements

This study could not have been carried out

without a great deal of support. I am specially

erateful for financial assistance from the

Australian Biological Resources Study, to the

officers-in-charge of the Australian herbaria,

AD, CANB, DNA, MEL, NSW and PERTH

for the extended loan of specimens, and to those

of CAL, NOU, PR, and T for lending special

material. Much of the work was carried out at

the Royal Botanic Gardens, Kew, and I am

indebted to the Director, Keeper of the

Herbarium and individuals of the Legume

Section for their assistance and for making my

stay at Kew pleasant as well as profitable. Dr

Charles Jarvis generously gave his time in

suiding me through the intricacies of the

Hermann Herbarium at the Natural History

Museum (BM): I wish to thank him and also

the Linnean Society for allowing me to consult

the Linnean Herbarium (LINN). I am grateful

to the Royal Geographical Society of

Queensland whose 1992 expedition to Cape

York Peninsula gave me the opportunity to

study some species of Desmodium ser.

Stenostachys in the field.

The work would have been sketchy

indeed without the extensive collections of Mr

Roy Pullen and J thank him for them and also

for his annotations and notes on herbarium

specimens which have been of great value. Ms

Judy Wheeler (PERTH) drew my attention to

Alysicarpus cheelii C.A. Gardner which I

would surely have missed otherwise.

Finally I must thank the Chief Botanist

of the Queensland Herbarium, Dr G.P. Guymer,

for allowing me the continuing use of the

facilities and collections of the institution, and

Mr David Halford who patiently taught me the

rudiments of word-processing.

Austrobaileya 5(2): 209-261 (1999)

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SCHINDLER, A.K, (1924), Uber eingige kleine Gattungen

aus der Verwandtschatt von Desmodium Desv.

Repertorium specierum novarum regni vegetabilis

20:266—-286.

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combinationes novae II. Repertorium specierun

novarum regni vegetabilis 22:250—-288.

——— (1928). Die Desmodiinen tn der botanischen

Literatur nach Linné. Repertorium specterum

novarum regni vegetabilis, Beihefte 49:1-371.

SCHUBERT, B.G, (1964), Anton K. Schindler, Zaxon 13:7-

10,

—— (1971). Desmodium. In E. Milne-Redhead & R.M.

Polhill (ed.):; Leguminosae subf. Papilionoideae.

Flora of Tropical East Africa. Kew: Crown

Agents for Overseas Governments &

Administrations.

STEENIS, C.G.G.J. vAN & VELDKaAmpP, J.F, (1982).

Miscellaneous Botanical Notes XXV 156.

Reduction of Trifidacanthus Merr. to Desmodium.

Reinwardtia 10:23-24.

TURLAND, N.J. & JArvis, C.E. (1997). Typification of some

Linnaean specific and varietal names in the

Leguminosae (Fabaceae). 7axon 46: 457-485.

VERDCOURT, B. (1997), Desmodium. In A manual of New

Guinea Legumes; 385-413. Office of Forests,

Papua New Guinea. Botany-Bulletin No. 1

ADSL Ae ALPE PO AU a aR St ta ae aan VeVi VIN ape ee ieee ee ee ett tet te ener eee eeeeeeeeeeee foe ne at bt

ARAM papi oite ahve Rie en,

Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland,

Australia

Marco F. Duretto

Summary

Duretto, Marco F, (1999), Boronia sect. Valvatae (Benth.) Engi. (Rutaceae) in Queensland, Australia.

Austrobaileya 5(2): 263-298. A numerical analysis, using phenetic methods, was undertaken on the

Boronia rosmarinifolia species complex. Four taxa were identified, three of which are new and are

described here as B. forsteri, B. splendida and B. palasepala. Nine other new taxa belonging to Boronia

sect. Valvatae (Benth.) Engl. (8. bella, B. duiganiae, B. excelsa, B. foetida, B. hoipolloi, B. jensziae, B.

odorata, B. quinkanensis and B. squamipetala) are also described. All new taxa are confined to

Queensland. A key to Boronia sect. Valvatae in Queensland is provided.

Keywords: Boronia sect. Valvatae, Boronia, Rutaceae, Boronia bella, Boronia duiganiae, Boronia

excelsa, Boronia foetida, Boronia forsteri, B. hoipolloi, Boronia jensziae, Boronia odorata, Boronia

palasepala, Boronia quinkanensis, Boronia rosmarinifolia, Boronia splendida, Boronia squamipetala.

Marco F, Duretto, School of Botany, The University of Melbourne, Parkville, Vic. 3052, Australia.

Present address: National Herbarium of Victoria, Royal Botanic Gardens Melbourne, Birdwood Avenue,

South Yarra, Vic. 3141, Australia. ph: 03 9252 2300 fax: 03 9252 2350 e-mail: duretto@rbgmelb.org.au

Introduction

As the ‘Flora of Australia’ account of

Boronia Sm. sect. Valvatae (Benth.) Engl.

was being prepared it became apparent that

B. rosmarinifolia A.Cunn. ex Endl., as currently

circumscribed, included a number of taxa.

Endlicher (1837) described B. rosmarinifolia

from material that was collected by Alan

Cunningham from Peel’s Island, Moreton Bay,

Queensland (Fig. 1). Boronia rosmarinifolia 1s

characterised by simple, sessile leaves that are

hirsute abaxially, glabrous adaxially and have

recurved to revolute margins. Specimens with

these features occur in coastal areas from

Bundaberg (Queensland) to Grafton (New South

Wales) and inland to the Carnarvon Range area

of Queensland (Fig. 1). In comparison with B.

rosmarinifolia sensu stricto (coastal

populations), plants previously included in

B. rosmarinifolia from the Carnarvon Range and

Robinson Gorge have smaller leaves and

flowers; and those from the Monto and Dalby-

Chinchilla—Haldon areas are taller and have

larger leaves and flowers as noted by Lebler

(1972). To ascertain whether these forms

watrant taxonomic recognition a numerical

Accepted for publication 29 May 1998

analysis, using phenetic methods, was

undertaken.

An apparently undescribed taxon from

the Pilliga Scrub (western slopes, New South

Wales) has been known in the literature as B.

sp. aff. rosmarinifolia B (Jacobs & Pickard

1981; Weston 1990; Weston & Porteners 1991).

Though superficially similar to B. rosmarinifolia,

it appears to be more closely related to B. glabra

(Maiden & Betche) Cheel which is also found

in the Pilliga Scrub (Duretto 1995, submitted),

though it occupies a different local habitat (D.

Mackay, NE, pers. comm.) The status of this

form is being assessed by D. Mackay and will

not be dealt with further in this paper.

There are a number of other Queensland

Boroniataxa with simple leaves that have been

assigned previously to B. rosmarinifolia or B.

sp. aff. B. rosmarinifolia (e.g. in Stanley & Ross

1983). Most of these taxa have broad, flat,

simple leaves (at least on older plants) and are

quite distinct from B. rosmarinifolia and other

Boronia species. These are described here as

B. bella Duretto, B. excelsa Duretto, B. foetida

Duretto, B. jensziae Duretto and B. odorata

Duretto. To complete the revision of Boronia

sect. Valvatae in Queensland, four pinnate leaved

264

147° E

6 Carnarvon

Ranges Robinson

> % Gorge

0 200 km

Austrobaileya 5(2): 263-298 (1999)

24°Ss

-_ Bundaberg™

Monto "O

laeaie' pg

O38

‘Manar

2 e

we Dalby \)

Brisbane 5 .

Toowoomba s ne . as

Peel Island

QUEENSLAND Moreton Bay

NEW SOUTH WALES eo’ S

Grafton a

Fig. 1. Distribution of collection localities of specimens used in analysis of Boronia rosmarinifolia species complex (1—

45); Boronia rosmarinifolia Group A («); B. forsteri Group B (¢); B. splendida pro parte Group C (A); B. palasepala

Group D (@); B. splendida pro parte Group E (0),

species (B. duiganiae Duretto, B. hoipolloi

Duretto, B. guinkanensis Duretto and B.

squamipetala Duretto) are also recognised and

described here.

A specimen assignable to B. ledifolia

(Vent.) DC. from the Pioneer River area of

central Queensland (Pioneer River, Queensland,

Dr Griffith, 1889 [MEL]) is the only collection

of a Boronia species that has been made in the

Mackay region seen by the author. Boronia

ledifolia, belonging in Boronia sect. Valvatae,

is found in New South Wales and Victoria. The

above specimen is therefore presumed to be

mislabelled and may have been collected in

Victoria where Dr Griffith had travelled

(Duretto 1995, submitted), and this species is

hence not dealt with further here. The taxon in

southern Queensland called B. ledifolia by

Neldner (1992), Ross (1994) and Forster

(1997) is probably either B. duiganiae or B.

odorata; the taxon in north-eastern Queensland

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 265

called by that name by Tennison-Woods (1882)

is probably a species of Zieria Sm. as it was

ascribed very small flowers and trifoliolate

leaves.

Materials and Methods

Material

Herbarium specimens from AD, BRI, CANB,

DNA, MBA, MEL, MELU, NSW, PERTH,

QRS and TCD were made available to the

author. Herbarium abbreviations follow

Holmgren et al. (1990). These specimens were

augmented by material collected 1n the field. A

list of all specimens examined is available from

the author.

Anatomy

The central portion of the leaves of all taxa was

sectioned transversely. Material to be sectioned

was fixed in Mirsky’s fixative (MAA) or 70%

ethanol. If fresh material was not available,

herbarium samples were re-hydrated by being

placed in water with a small amount of

detergent, brought to the brink of boiling, left

simmering for one hour and soaked overnight

before fixing in MAA. All fixed material was

then placed in 70% ethanol overnight,

dehydrated through a graded ethanol series up

to 100% ethanol, infiltrated with 100% LR-

White (London Resin) through a resin/ethanol

series, and polymerised at 60°C. Sections 2 pm

in thickness were cut on a Reichert Ultracut

ultra-microtome, stained with 0.05% toluidine

blue solution (pH 4.4) and observed and

photographed using an Olympus BHS

compound microscope. Anatomical features

observed are described in the taxonomic

descriptions below. Voucher specimens for leaf

anatomy are listed in Appendix 1.

Scanning electron microscopy

Trichomes on leaves and petals, and seed

surfaces were surveyed for all taxa (where

material was available) using a scanning

electron microscope. Dry leaves, petals and

seeds were mounted on stubs using double sided

or carbon tape with conductive carbon paint,

coated with gold using an Edwards Sputter

Coater $150B and examined and photographed

at SK V using a JEOL 840 Scanning Electron

Microscope equipped with a lanthanum

hexaboride filament. All photographs of seeds

were of central areas on a lateral side. Trichome

and seed characters are described in the

taxonomic descriptions below.

Phenetic Analysis of the B. rosmarinifolia

species Complex

Characters

Eleven characters (Table 1) were scored for 45

herbarium specimens (Table 2) covering the

geographic range of the B. rosmarinifolia

species complex. Scores are an average of 10

measurements (where 10 organs were available)

and ratios are the average of the individual ratios

of the 10 organs measured. There are some

problems associated with the use of ratios in

phenetic analyses (see Duretto & Ladiges 1997

and references therein for discussion): here

ratios are used as a means of quantifying and

standardizing leaf and sepal shape (characters

4 and 7).

Table 1. Morphological characters used in

the phenetic analysis of the Boronia

rosmarinifolia species complex.

I Style glabrous/hirsute, 0/1

ps Terminal leaf length (TLL) (mm)

3 Terminal leaf width (TLW) Gnm)

4 TLW/TLL

5 sepal length (SL) (mm)

6 Sepal width (SW) (mm)

7 SW/SL

8 Petal length (mm)

9 Petal width Gmm)

10 Stellate hair rays on sepals <0.25 mm

long/+0.5 mm long, 0/1

1] Anther appendage recurved/erect, 0/1

266

Austrobaileya 5(2): 263-298 (1999)

Table 2. Data used in the phenetic analysis of the Boronia rosmarinifolia species complex.

Principal collector only given. For quantitative characters, mean values are given (see Table 1).

Specimen

Number

Gollector

& number (or date)

Herbarium

& sheat number

smith 7 MEL (MEL259152)

Ross 3196 MEL {(MEL1552623)}

Jobson 930 MEL (MEL221851)

Parish, 6.x.1982 MEL (MEL626024)

Walsh 1399 MEL {MEL1545124)

Baxter 1132

Bird, 7.vii.1990

NSW (NSW243819)

BRI (AQ472316)

a PONaanrana

Willis, 10.viil.1982 MEL (MEL628666)

Olsen 330 NSW (NSW243826)

McDonald 476 BRI (AQ117773)}

14 White, 12.vili.1930 NSW (NSW243829)

12 Moriarty 415 CANB (CANB253236}

13 Duretto 259 MEL ({(MEL2036432)

14 Duretto 256 MEL (MEL2036427)

15 Duretio 254 MEL (MEL2036425)

16 Duretio 257 MEL (MEL2036428)

17 Duretto 258 MEL (MEL2036430)

18 Grieves, 22.vii.1979 NSW (NSW243816)

19 Foraman 907 MEL (MEL1539697)

20 Duretto 276 MEL (MEL2036609)

2 | Duretto 277 MEL (MEL2036610)

22 Duretio 275 MEL (MEL2036608)

23 Duretto 279 MEL (MEL2036674)

24 Forster 6906 BRi (AQ472561)

25 Forster 6961 BRI (AQ472512)}

26 Martensz 1014

27 Forster 4647

28 Forster 2243

2g Forster 4762

30 Shoobridge, 29.ix.1964

31 Williams 84159

32 omith 14462

33 Duretto 337

34 Duretto 339

35 Duretto 338

36 Duretto 342

37 shoobridge, 30.ix

38 Forster 41202

39 Forster 411235

40 Forster 11453

4] Forster 11429

42 Forster 11244

43 Gittens 2745

44 Thomas 138

45 Wiliams 86097

CANB (CANB284160)

BRI (AQ408650)

BRI {AQ441712)

MEL (MEL1575271)

BRI (AQ15118)

1 (AQ416779)

BRI (AQ403268)

MEL (MEL2036656)

MEL (MEL2036657)

MEL (MEL2044555 )

MEL (MEL2036660)

CANB (CBG15711)

MEL (MEL 2049143)

MEL (MEL 2049140)

MEL (MEL 2049118)

MEL (MEL 2049141)

MEL (MEL 2049142)

BRI (AQ264152)

CANB (CBG8900796)

BRI (AQ406813)

.1964

Most characters are self explanatory but

a few require clarification.

For characters 5 to 9, lengths, widths and

ratios of perianth members proved difficult to

measure accurately due to shrinkage of organs

while drying, and their haphazard orientation

on the herbartum sheet. Usually only a small

number of these organs could be measured with

any confidence on any herbarium specimen so

measurements cited here should be treated as

minimum values. Sepals and petals were

measured on flowers without fruit as these

organs enlarge during fruit development in most

are gsart

2 : { 4

oe oo eo ooec Osama OOO KH Ocoee oc eo Oe eco co mMO moo ce oOo oe Oe Or

3 5 6 7 8 9 10 41

19.3 2.7 0.1 2.5 1.5 0.60 7.0 3.5 0 0O

15.7. 1.9 0.1 2.5 2.0 0.80 6.5 3.5 0 O

18.6 1.8 mee: 3.0 2.5 0.83 6.0 3.0 0 O

24.3 4.1 0.17 2.5 1.5 0.60 7.0 3.5 0 O

22.3 2.6 0.12 3.0 2.5 0.83 7.0 3.8 9O Oo

19.8 2.1 0.10 3.0 2.0 0.67 6.0 3.3 0 0

18.7 2.5 0.13 4.0 2.5 0.63 6.0 3.5 90 Oo

19.5 1.7 0.09 4.0 2.5 0.63 7.0 3.3 0 Oo

16.1 1.7 0.10 2.5 1.5 0.60 6.0 3.0 0 0

19.0 2.0 0.11 3.0 2.0 0.67 6.5 2.7 O OO

16.6 1.8 0.12 3.0 2.0 0.67 6.5 3.0 0 9

24.6 2.0 0.08 4.0 2.5 0.63 8.0 3.38 90 0

12.8 2.2 0.17 2.0 1.5 0.75 6.0 4.0 0 90

12.8 2.1 0.17 2.5 2.0 0.80 5.5 4.60 0 O

14.8 2.9 0.20 2.5 2.0 0.80 6.0 4.0 06 0O

12.7 2.5 0.20 2.5 14.5 0.60 6.0 4.0 0 @Q

21.6 4.1 0.19 3.0 2.0 0.67 6.5 4.0 0 90

19.8 1.7 0.08 3.5 2.5 0.71 7.0 3.8 0 O

19.5 2.4 0.12 4.0 2.0 0.50 8.0 4.0 0 06

30.0 3.7 0.12 4.5 3.5 0.78 8.0 5.0 1. O

26.0 2.9 0.11 4.0 2.5 0.63 8.56 4.5 1 ?

28.2 4.0 0.14 4.5 3.5 0.78 8.0 5.0 1 ?

30.0 3.5 0.12 3.5 2.5 0.71 8.0 5.0 1 ?

30.9 4.7 0.16 5.0 4.0 0.80 10.0 6.0 1 ?

$0.2 4.3 0.15 5.0 4.0 0.80 9.0 5.5 1 9

24.0 2.7 0.12 4.0 3.0 0.75 9.0 5.5 1 9

22.5 2.4 0.11 6.0 4.0 0.67 10.0 6.0 1 9

24.0 1.0 0.04 3.5 2.0 0.66 ? ? Oo 9

35.6 2.0 0.06 3.0 2.0 0.67 10.0 6.0 06 9Q

19.3 1.3 0.07 4.0 2.5 0.63 86 5.0 0 0

18.3 1.2 0.07 4.0 2.5 0.63 10.0 5.5 90 O

17.9 1.1 0.06 4.0 2.0 0.50 9.0 5.0 0 0

18.1 4.9 0.11 4.0 3.0 0.75 8.5 5.0 0 Oo

19.3 1.4 0.07 3.5 2.0 0.57 8.0 5.0 0 0O

19.8 1.3 0.07 3.5 2.0 0.57 8.5 5.0 0 O

18.8 1.2 0.06 3.5 2.0 0.57 8.0 4.5 0 0

28.2 1.9 0.07 4.5 3.0 0.67 10.0 65 0 O

25.9 2.1 0.08 4.5 3.0 0.67 11.0 6.0 0 0

15.1 2.6 0.17 3.0 1.5 0.50 5.5 2.0 0 1

19.1 383.41 6.16 2.5 1.5 0.60 5.0 2.0 0 1

15.7 2.7 0.18 2.5 1.5 0.60 5.0 2.5 a 1

17.2 2.7 0.16 2.5 1.5 0.60 4.5 2.0 0 17

19.0 1.9 0.10 2.5 1.5 0.60 5.0 2.5 0 1

12.4 2.6 0.21 2.0 1.0 0.50 5.5 2.5 0 1

12.0 2.1 0.18 2.0 14.0 0.50 4.0 141.8 oO 1

members of Boronia sect. Valvatae.

For character 10, most specimens had

multiangular stellate haits with rays that were

too small to measure confidently as they were

much less than 0.25 mm long. Specimens 20 to

27, though, have hairs with rays that reach 0.5

mm in length. As there was no gradation

between these states, this numerical character

was scored as a binary character (Table 1).

Data Analysis

All data sets were analysed using PATN (Belbin

1987) following the methodology outlined in

NRSREA RMS TEAR a ee ee

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 267

Duretto & Ladiges (1997). Data were range

standardised before Manhattan dissimilarity

measures were calculated. For cluster analysis,

both flexible UPGMA (unweighted pair group

arithmetic averages) and flexible WPGMA

(weighted pair group arithmetic averages) were

utilised as fusion strategies. Data were ordinated

in three dimensions using the multidimensional

scaling, MDS, KYSP algorithm (Kruskal et al.

1973). The Hybrid option of Faith et al. (1987)

was chosen. Twenty different random starting

points were used for each analysis and the run

with the lowest stress value is shown. Character

correlations with the ordination vectors were

calculated using the PCC function of PATN.

Minimum spanning trees (MST) were also

calculated.

Taxon descriptions

Descriptive terminology follows Theobald et ai.

(1979) and Hewson (1988) for hairs, Briggs &

Johnson (1979) and Weston (1990) for

inflorescence structure, and Murley (1951),

Powell & Armstrong (1980) and Barthlott

(1984) for seed surfaces. Conservation codes

follow the format of Briggs & Leigh (1996).

Results

Analysis 1 (all specimens)

Analysis one was based on the entire data set

(45 specimens x 10 characters; Table 2). Five

groups, A to E, are recognisable in both the

UPGMA (Fig. 2) and WPGMA (Fig. 3)

classifications, in the ordination (Fig. 4 & 5)

and in the MST (Fig. 6). Group A includes all

coastal collections (specimens 1-19); Group B

includes all specimens collected from Robinson

Gorge and Carnarvon Ranges (specimens 39—

45); Group C includes collections from the

Dalby and Haldon areas (specimens 29-37),

specimen 28 from near the ‘Mimosa’

Homestead and specimen 38 from near the

‘Beeron’ homestead; Group D includes all

collections from Coominglah State Forest

(specimens 20-26); Group E is comprised of

the single specimen 27 from near the ‘Manar’

Homestead (Fig. 1).

In the UPGMA classification (Fig. 2),

Group A fuses first with Group C and then with

Group B, while in the WPGMA classification

(Fig. 3), Group A fuses with Group B. In the

MST (Fig. 6), Groups B, C and D connect to

Group A at different places. Group E fuses with

Group D in both classifications (Fig. 2 & 3).

This larger group of D with E is the most

dissimilar in the UPGMA (Fig. 2) but fuses with

Group C in the WPGMA (Fig. 3). Group E is

isolated but closer to Group D in the ordination

(Fig. 4 & 5), but joins members of Group C in

the MST (Fig. 6). Characters highly correlated

with the vectors are 2, 5 and 6 for vector 1, 1, 3

and 4 for vector 2, and 7—9 for vector 3.

Analysis 2 (Specimens 20-38, characters

1-7, 10)

For the five groups (A—E) recognised in

Analysis 1, the relationships between Groups

C, D and E (specimens 20-38) were ambiguous

and so a data set containing specimens 20 to 38

and characters 1-7 and 10 was reanalysed.

(Invariant characters in the data set were

excluded from this analysis.)

Analysis 2 confirmed that Groups C, D

and E of Analysis 1 (Fig. 2—6) are distinct.

Group E fuses with Group D in both

classifications (not shown). Though isolated in

the ordination (Fig. 7, 8), Group E 1s closer to

Group D (Fig. 8). In the MST (not shown)

Group E is well within Group C, as was the case

in Analysis 1 (Fig. 6). Characters highly

correlated with the vectors are 3 and4 for vector

1,5 for vector 2, and 7 and !0 for vector 3.

Taxonomic interpretation

On the basis of the above analyses (Fig. 2-8),

four taxa, corresponding to Groups A

(specimens 1-19), B (specimens 39-45), C

(specimens 28-38) and D (specimens 20-26), |

are recognised at the specific level. Results on

the position of Group E (specimen 27) are

conflicting. Geographically, the closest

specimens to Group E are specimens 28 and

38 of Group C (Fig. 1) and in the MST (Fig. 6)

specimen 27 links with specimen 38. Both these

specimens have hirsute styles. As Groups C, D

and E do not chain in the MST there is no

evidence of a cline. Specimen 27 differs from

members of Group C in having wider leaves

and larger sepals, and from members of Group

268

D by having a hirsute style; a feature that is

variable in Group C. Given this pattern of

variation Group E is here considered to be

conspecific with Group C.

The members of Group A are

characterised by short hairs, recurved anther

appendage and floral parts that are larger than

those in members of Group B but smaller than

those in members of Groups C/E and D (Table

3). The members of Group B are characterised

by small floral parts, an erect anther appendage

and short hairs. The members of Group C

(including Group E) are characterised by the

extremely narrow (usually recurved) leaves,

large floral parts and short hairs. The members

of Group D are characterised by wide leaves,

comparatively large floral parts and long hairs.

Austrobaileya 5(2): 263-298 (1999)

Characters not used in these analyses that

confirm these results include: members of Group

B have hirsute fruit unlike the other groups (except

for two specimens of Group A); and members of

Group D have minute anther appendages unlike

members of Groups B and C, this character being

variable in members of Group A.

Coastal specimens (Group A) retain the

name 8&8. rosmarinifolia as the distribution of

this group of specimens includes the type

locality (Peel Island, Moreton Bay, Queensland)

of that name and these specimens match the

diagnosis given by Endlicher (1837) for this

species. Groups B, C (including E) and D are

here described as B. forsteri Duretto, B.

splendida Duretto and B. palasepala Duretto

respectively.

0.5

0.4

0.3

Group |A Group ;C Group |B Group|D Group|E

0.2

0.7

1 17 14 163 46 141 7 442 19 30 37 32 35 33 39 42 43 45 22 25 23 27

41315 2 5 10 9 8 i8 28 29 31 34 36 38 40 44 44 20 24 21 26

Fig. 2. Unweighted pair group arithmetic averages (UPGMA) classification, analysis one, all specimens.

sa . REC SARS

(panier te antec

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 269

0.5

Group A Group B Group C Group D Group E

Fig. 3. Weighted pair group arithmetic averages (WPGMA) classification, analysis one, all specimens.

A A A

C A

A 28

A A

es . EO

o ©

‘4 © ©

B A

‘ Oo O° re

> $

@ @

® es)

Fig. 4. Ordination (KYSP), vector | verses vector 2, analysis one, all specimens. Boronia rosmarinifolia, Group A(o);

B, forsterii, Group B (@); B. splendida (pro parte), Group C (A); B. palasepala, Group D (e); B. splendida (pro parte),

Group E (0). Specimen 28, Group C numbered.

270 Austrobaileya 5(2): 263-298 (1999)

% oF

$ B ¢ Ad, p ‘

° A

* g 4 ep

¥ > ih

res o °? res

mn A? A

“ ° 28

Fig. 5. Ordination (K YSP), vector | verses vector 3, analysis one, all specimens. Symbols are: Boronia rosmarinifolia,

Group A (<); B. forsterii, Group B (@); B. splendida (pro parte), Group C (A); B. palasepala, Group D (e); B. splendida,

(pro parte) Group E (©). Specimen 28, Group C (numbered).

Group B Group A Group C

45 17 5 16 37 36

44 —_— 39 4 3 13 19 29 34

Py] oF Tye tal LA

Os i 1 2 —— 14 7 28 35

| | AE EE LA pee

40 —— 43 9 ——~—11—— 10 ——6 — 8 30 —— 31 ——-_ 38 27

12 Group D 24

26 ——— 21 —— 23 —— 20 ——._ 22 ———. 25

Fig. 6. Minimum spanning tree (MST), analysis one, all specimens.

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 27)

AA ee |

28 A

Fig. 7. Ordination (K YSP), vector 1 verses 2, analysis two, specimens 20-38. Boronia splendida (pro parte), Group C

(A); B. palasepala, Group D (¢); B. splendida (pro parte), Group E (©). Specimen 28 Group C (numbered).

A A 28 D

Fig. 8. Ordination (K YSP), vector | verses vector 3, analysis two, specimens 20-38. Boronia splendida (pro parte),

Group C (A); B. palasepala, Group D (¢); B. splendida (pro parte), Group E (0), Specimen 28 Group C (numbered).

2ye

Austrobaileya 5(2): 263-298 (1999)

Table 3. Character ranges for Groups A, B, C, D and E, with means values given in brackets.

Character

fond,

. Style glabrous /hirsute

2. Leaf length (LL) (mm)

. Leaf width (LW) (mm)

4, LW/LL

5. Sepal length (SL) (mm)

6. Sepal width (SW) Gnm)

7. SW/SL

8. Petal length (mm)

9, Petal width (mm)

10. Stellate hair rays on

sepals < 0.25 mm long/c.

0.5mm long, 0/1

ll. Anther appendage

recurved/erect, 0/1

Group A

12,7-24.6

(18.3)

1.7-4.]

(2.3)

0.08-0.20

(0.13)

2,.0-4.0

(3.0)

1,5-2.5

(2.03)

0.50-0.83

(0.68)

5.5-8.0

(6.6)

2.67-4.00

(3.52)

Group B

0-1

(0.64)

17.9-35.6

(22.3)

— 1.0-2.1

(1.5)

0.04-0.11

(0.07)

3.0-4.5

(3.8)

2.0-3.0

(2.36)

0.50-0.75

(0.63)

8.0-11.0

(9.2)

4.5-6.5

(5.4)

Group C

1.6-3.1

(2.5)

1.9-3.]

(2.5)

0.10-0.21

(0.17)

2.0-3.0

(2.43)

1.0-1.5

(1.36)

0.50-0.60

(0.56)

4.0-5.5

(4,9)

1.8-2.5

(2.2)

Group D

24.0-4.7/

(3,7)

2.7-4.7

(3.7)

0.11-0.16

(0.13)

3,5-5.0

(4.4)

2.9-4.0

(3.3)

0.63-0.80

(0.75)

8 .0-10.0

(8.6)

4.5-6.0

(5.2)

22.5

2.4

0.11

6.0

4.0

0.67

Taxonomy

Key to Boronia sect. Valvatae in Queensland

F. ‘Pitriate leaves Presents ca cesic x lo) ace latenlla! wcncteon bo doen eed g eke wlaNacde aie bo we aU) Ble alere pel? Z

AU TO AVES RIMES O5 chelsea lsc hes cose ee trriccd oo tect arete legs, syadeawhcte ane asda 0h ore ato see analtete aces 16

2. Stellate hairs, especially on petals, with fused rays and often appearing

peltate; abaxial surface of sepals glabrous (N Qld)... 2... . cc ee ee ee 3

Stellate hairs with distinct rays; abaxial surface of sepals glabrous or with

ASpParse To MENSe INGUMEMEUNT 0.2.2 Lace pata te Ate ete Rata tA eon od ote Hae Beeson wae 4

PEAY EVES eS ee ence y IG ey eS

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 273

3. Pinnae linear; branches obviously glandular; petals with a sparse

PLUCEUITS TCU ATA as 5 se, wae esting fa be a SA lena staal a cle ee A ed ww! Fae RS, KIDS . B. bowmanii*

Pinnae elliptic; branches not distinctly glandular; petals with a dense

indumentum abaxially, scaly in appearance. .......... 0c eee eee 13. B. squamipetala

4. Adaxial and abaxial leaf surfaces with a dense indumentum (no epidermis

SIS EOE Vous ett ada th el 4 Peake ss al a tenaaiaens. DAC iainl ge aalliod one heen tile fem Won HA hee etks ain ecireh dee nd’ §

Adaxial surface of leaves without a dense indumentum (epidermis clearly

visible); abaxial leaf surface glabrous or with a sparse to dense

TELOUIETE SLICED Fe aes sates ere Bhan eck koe owe We Ti GaP heer a ed eT acl 9 Pa ghtestah Odea Nou ghee“ eee heel pe §

5, Sepals much longer and wider than petals ......... 00. cece eee B. lanuginosa’®

Sepals much shorter than petals, or as long as but then much narrower

VATA PIS GANS var Bos ayont 7 toad tat ctae, Peyeacane S8e. ean ce ANote se yneueatahceshard deen hearin atest eee pak tetra kan 6

6. Sepals ovate, 1.5 to 2 times as long as wide, with acuminate tip (central

COTELIES Ss, oo Base csuy. 9 va tebieseait eaLdEN. | al bodes episee fhe? a) lan alot abas Gackie! eh epee b-g FI MUR 11. B. duiganiae

Sepals narrowly deltoid, at least 2.5 times as long as wide, with acute tip

CVE, GEN WY Qh sa sare nat Espana Macwe Meet pean deat athlete Dg lta iw & hace ote eke Eiht wedges weds 7

7. Pinnae linear to narrowly elliptic, c. 1 mm wide; sepals 2—3.5 mm long

ENEWS) GLY Wi a3 asain atid s fal is alt fa wan Seda be Goghe al Acts ty wRaamzdhOe bet aue Geese ap Momma 9, B. hoipolloi

Pinnae elliptic to oblanceolate, (1—)3—7 mm wide; sepals 3—5 mm long

CIS REET: Bs seus. eure nceiar ele duced Mote cteck. a? ati baa GFE saath we, anal Ata UA A 10. B. quinkanensis

8. Leaves strongly discolourous with a dense indumentum on the abaxial

surface Lepiderinis OL MISIDIG), ys ¢ Wweew hee te eos o's Oe Beeler n GLUES > liletie aay'd Bethe. 9

Leaves slightly discolourous or concolourous, abaxial surface glabrous

or with a sparse to moderate indumentum (epidermis clearly visible) ............. 14

9, Sepals narrowly deltoid, 2.5 times as long as wide, with tip acute...............006. 10

Sepals ovate, 1.5 to 2 times as long as wide, with tip acute or acuminate ............. 12

10, Leaves trifoliolate (Blackdown Tbld, Central Qld) .............0. 0.0000. B. obovata*

Leaves 5—17-fohtolate GN or SE QI). 2. ce ek ha Pb ea ben eae ew ba ee 1]

11. Leaflets < 5 mm wide; petals 3—7 mm long, the adaxial surface with a

dense indumentum; perianth often glabrous abaxially (N Qid)............. B. alulata’®

Widest leaflets > 5 mm wide; petals (6—)8—12 mm long, the adaxial surface

with a sparse indumentum; perianth never glabrous abaxially (SE Qld)..... B. amabilis”

12, Leaves sometimes trifoliolate when juvenile but simple when mature,

margins flat to slightly recurved; peduncle < 2 mm long; anthopodium |-

Sm long: petals 327 MAMHONG ec ppc pete aed Fy ee oe lee 2 12. B. odorata

Leaves imparipinnate, sometimes becoming simple with age, margins flat

to revolute; peduncle (1—)2—10 mm long; anthopodium 7-11 mm long;

petals (oS, 512 TONG fh eva lia Serpe a atid dames boeatare o 4aarg ars 4, 8 ae es 13

13. Sepals with tip acuminate, > 3.5 mm long, > 2 mm wide; adaxial surface

of leaves with a sparse to dense indumentum .............0000 eee 11. B. duiganiae

Sepals with tip acute, < 3 mm long, < 2 mm wide; adaxial surface of

leaves glabrous or with a sparse indumentum ......... 0.0000 c eee een B. ledifolia®

14, Midrib raised on abaxial surface of leaves (Cooloola sand mass, SE Qld) ...... B. keysii*

Midrib not raised on abaxial surface of leaves (inland Qld). .... 0.0... 0. cee eee ee eee 15

274 Austrobaileya 5(2): 263-298 (1999)

15. Branchlets not conspicuously glandular; leaves with sparse to moderate

indumentum of hairs with flexuous rays, the hairs sometimes stalked

(Granite Bet SE OG) 25 say, ere oes Ga eRe wae ER Lk pe on ey REG eee B. granitica’

Branchlets with large hemispherical glands; leaves glabrous or with a sparse

indumentum of sessile hairs with straight rays (Central Highlands &

DV AL ARVO CS TUNIC) pact, obs fark Bosrana res > eerosectcat enh or carey pom cme y ox hee aise wens SOLA B. eriantha*

16. Mature leaves only slightly discolourous, glabrous or with a sparse to

moderate indumentum on abaxial surface ...... 0. cc eee eee ees 17

Mature leaves markedly discolourous with a dense indumentum

(epidermis not visible) on abaxial surface (juvenile leaves notso).............04. 19

LF (LGRAV ES PCTO LAGS. 1. aU yet wyhaedd BES re oct M yeuan'e 2 av astios ti ste on ee ae owe eae ie Ne B. keysii"

EAVES SESSEIE Fo perry sistas ate Sle Ed a finte bu al we eeeslnn th wily ghratias ae Wha Ooh qcleccitens og! al ws fe SLAG iE aiey Bees 3 18

18.Leaf margin glandular punctate; leaves with a sparse to moderate

indumentum of stalked hairs with flexuous rays ........ 0... cece eee B. repanda*

Leaf margin smooth, leaves glabrous or glabrescent; hairs sessile, rays

SHEEP OITE oe eras xcs a Westen cecteteciie e,-ahta 1 acts lle ip eoascdtaBiaa oe feragl LSE a Slag atin: aaa wth alla Goin B. glabra*

19. Leaves sessile, base not strongly attenuate... 0... ee eee ee eee 20

Leaves petiolate or leaf base strongly attenuate ..... 0... eee eee 24

20. Petals (6—)8—13 mm long; sepals 2.5—6 mm long, (2—)3—-4 mm wide ................ 21

Petals 4~7.5 mm long; sepals 2-4 mm long, 1-2 mm wide ............. 0.0. c ce eee 22

21. Leaves strictly revolute, 1-2(—4 mm) wide; anther apiculum large and

reflexed; stellate hairs with rays to0.25mmlong................... 2. B. splendida

Leaves flat to recurved, sometimes revolute on drying, 2-6 mm wide; anther

apiculum absent or minute; stellate hairs with rays to 0.5 mmlong .... 3.B. palasepala

22, Abaxial surface of petals glabrous or glabrescent; largest leaves greater

ENE SOAPS ere ang dev ak po ee on nba eee Sree A aria eng bed ng 6. B. excelsa

Abaxial surface of petals with a sparse to moderate simple indumentum;

largest leaves usually less than 35 mm long ..... 0.0... ccc eee ee eee 23

23. Fruit glabrous or with a sparse indumentum, very rarely densely hirsute;

anther apiculum reflexed; stems terete to slightly quadrangular; sepals

2—4.5 mm long; petals 5~7.5 mm long (coastal and near coastal SE Qld

ATA DNS Yeo cca, 9: ee iv te, SOE eo harp Lge in he eva es er eee detoregerec +... 1. B. rosmarinifolia

Fruit densely hirsute; anther apiculum erect; stems quadrangular; sepals

2—2.5 mm long; petals 4—-6 mm long ( central and inland Qld)............... 4. B. forsteri

24. Stamen filaments glabrous or with 1 to 3 simple hairs; petals < 5.5 mm

HOTT WV SOU, IND vs ern nese ecstacy, ¢ 4 ARR fem aay Te Bote Selene: Be of eedcocylldl Wie B. lanceolata *

Stamen filaments densely hirsute; petals (4.5—)5.5-12 mm long (NE &

EWM ann oz. 3 Lteewleroe wie & piaunanetegeltias See fF otc Seteccva Se fac Bl ales HM af EOP ps at Be gee Oia SH ee SLED 25

25, Leaf adaxial surface with a sparse to moderate indumentum .............00 0c cee 26

Leaf adaxial surface glabrous or with few hairs along midrib ............ 2.0.00 ues 27

26. Leaves elliptic, (2—)4—8 mm wide; peduncle 1-2 mm long; iin erat

1—5 mm long (central inland Old) goa SL SSN ek dear. 389, GSM a asene ae seems fetes te a 12. B. odorata

Duretto, Boronia sect. Valvatae (Benth.) Engl, (Rutaceae) in Queensland 275

Leaves narrowly elliptic, 1.5-—5 mm wide; peduncle 3-5 mm long;

anthopodium 4—8 mm long (central coastal Qld?, NSW, Vic.)

a eee B. ledifolia*

27. Adaxial surface of petals with sparse to moderate indumentum of simple

hairs (Hinchinbrook Is. of N Qid).......

ae ea Bhp tee tar niyo vl SUE eta meee a . 5. B. jensziae

Adaxial surface of petals glabrous or with very few simple hairs (Mt

Windsor Tblds of N Qid or SE Qld) .....

28. Leaves narrowly elliptic, < 6 mm wide (Mt Windsor Tblds of N Qld)

Leaves elliptic, to 14 mm wide (SE Qld) oo... cc ee eee eens 29

29. Sepals 2—3.5 mm long (before fruit development); petals 6-8 mm long;

peduncles 2~3 mm long (Mt Walsh)... ... 00.0.0... ccc eee ee eens 7. B. foetida

Sepals 4.5—5 mm long (before fruit development); petals 9-10 mm long;

peduncles to 0.5 mm long (Many Peaks Ra)... ok. ees 8. B. bella

# Currently accepted Queensland species that are not dealt with further here but discussed in detail in Duretto (1997,

and/or submitted). Boronia lanuginosa Endl. has recently been collected from NW Qld (P.L. Forster pers. comm.)

* Species not found in Queensland but included in Key as it is found very close to Queensland-New South Wales border

(see Duretto submitted),

Boronia sect. Valvatae (Benth.) Enel., Nat.

Pflanzen. 3(4), 135 (1896); Boronia ser.

Valvatae Benth., Fl. Austral. 1: 308, 311

(1863). Type: type not cited (see below).

Boronia sect. Valvatae has recently been revised

(Duretto submitted) and is lectotypified therein.

To avoid confusion and duplication in the

species descriptions below a short description

of this section is given here.

Inflorescence cymose, axillary. Sepals

valvate, persistent with mature fruit. Petals

valvate, with tip not inflexed, persistent with

mature fruit. Stamens 8, all fertile; anthers

glabrous. Stigma rounded, not or scarcely wider

than style. Seed elliptical in outline with adaxial

surface flattened.

1. Boronia rosmarinifolia A, Cunn. ex Endl.,

Enum. Plant., Htigel: 16 (1837). Type:

Queensland. Moreton District: Peel’s Is-

land, Moreton Bay, in 1824, A.

Cunningham (holo: W?, 1.¥.)

Boronia ledifolia var. rosmarinifolia (A.

Cunn. ex Endl.) Benth., Fl. Austral. 1: 314

(1863).

Illustrations: B.A. Lebler, Qld Ag. J. 98:

196 (1972); K.A.W. Williams, Native PI.

Qid 1: 37 (1979); L. Cronin, Concise

Aust. Fl. 80 (1989); PH. Weston & M.

Porteners, Fl. NSW 2: 232 (1991); Fig.

9A-F,

Erect or weakly ascending, much branched

shrub to | m tall. Multiangular stellate hairs

sessile, with 5-10 rays; rays unicellular, free,

firm, straight, (0.05—)0.1 mm long, glossy,

smooth, white to yellow. Branches terete to

slightly quadrangular in TS, not glandular, with

little or no cork development, with a moderate

to dense stellate indumentum, becoming

glabrous with age, will regrow from a rootstock;

decurrent leaf bases absent. Leaves simple, not

conspicuously glandular, sessile, elliptic to

obovate, 6-30 mm long, 1-4.5 mm wide, with

tip obtuse, strongly discolourous, paler

beneath, lamina with palisade and spongy

mesophyll; margins entire, recurved or flat;

midrib raised slightly to prominent abaxially,

with tightly packed parenchyma without

secondary thickening between midvein and

abaxial epidermis, impressed adaxially; adaxial

surface glabrous or with few hairs along midrib;

abaxial surface with a dense indumentum of two

hair types, a moderate layer of multiangular

stellate hairs over a dense layer (sometimes

lacking from midrib) of peltate stellate hairs;

276 Austrobaileya 5(2): 263-298 (1999)

oe ae lata eae

ee a en

on | mE us image *"*

a Mes elt

Fig. 9, A-F, Boronia rosmarinifolia. A, flowering branchlet; B, flower; C, sepal; D, abaxial view of antesepalous

stamen; E, abaxial view of antepetalous stamen; F, disc and gynoecium. A-F, Duretfto 257 (MEL). G—L, B. splendida. G,

flowering branchlet; H, flower; I, sepal; J, abaxial view of antesepalous stamen; K, abaxial view of antepetalous stamen;

L, disc and gynoecium. G—L, Duretto 337 (MEL). M-R, B. palasepala. M, flowering branchlet; N, flower; O, sepal; P, —

abaxial view of antesepalous stamen; Q, abaxial view of antepetalous stamen; R, disc and gynoecitum. M-R, Duretto 279

et al. (MEL). S—X, B. forsteri. S, flowering branchlet; T, flower; U, sepal; V, abaxiai view of antesepalous stamen; W,

abaxial view of antepetalous stamen; X, disc and gynoecium, S—X, Forster 11235 (MEL). Scale bar: A, G, M, S = 24

mm; B, H, N, T= 10 mm; C, [, O, U =6 mm; D-F, IF-L, P-R, V—X =4 mm. Del. Peter Neish.

SRE a A SB VA Bg 2

PERSE SERIA ASS We

Duretto, Boronia sect. Valvatae (Benth.) Enel. (Rutaceae) in Queensland 277

juvenile leaves to 48 mm long and 10 mm wide,

glabrous but becoming progressively more

hirsute along shoot. Inflorescence 1(-—3)-

flowered, with a moderate to dense stellate

indumentum; peduncle to 0.5 mm long,

deciduous with flower; prophylls unifoliolate,

1.5—2 mm long, to 0.5 mm wide; metaxyphylls

to 0.5 mm long; anthopodium 1—6 mm long.

sepals (Fig. 9C) ovate-deltoid, 2~4 mm long,

1.5—2.5 mm wide, enlarging slightly with mature

fruit, with tip acute; adaxial surface densely and

minutely pubescent, becoming glabrous towards

base; abaxial surface with a dense stellate

indumentum. Petals pink to white, 5—7.5 mm

long, 3—4 mm wide, enlarging to 8-10 mm long

and 6 mm wide with mature fruit, with midvein

raised abaxially; adaxial surface sparsely to

moderately simple pubescent; abaxial surface

with a moderate to dense stellate indumentum.

stamen filaments bearing stiff simple hairs

abaxially and on margins below glandular tip;

antesepalous filaments clavate, tapering to anther

connective, c. 2 mm long, the distal 0.5—1 mm

prominently glandular (Fig. 9D); antepetalous

filaments c. 1.5 mm long, the distal end

glandular (Fig. 9E). Anthers monomorphic or

antepetalous anthers slightly larger before

dehiscence; anther appendage large, reflexed,

glabrous. Disc entire, not surrounding base of

filaments, glabrous (Fig. 9F). Gynoecium

glabrous (Fig. 9F). Coccus 4—5.5 mm long, 2.5—

3.5 mm wide, glabrous or very rarely densely

hirsute. Seeds black, shiny, 44.5 mm long, 2—

2.5 mm wide, adaxial side without a ridge;

elaiosome (placental portion of endocarp)

yellow-white; surface at magnification as with

5. odorata, Fig. 10A,B). Rosemary Boronia,

Forest Boronia or Possum Boronia.

Additional selected specimens (c. 50 collections

examined): Queensland. Burnett District: Curtis Rd,

Kingaroy, 26°31°S 151°52’E, Sep 1996, Bean 10650

(MEL); Wipe Bay District: W side of highway, Sunshine

Beach, 2 miles S of Noosa, 26°26’S 153°04’E, Oct 1968,

Baxter & Lebler 1132 (CANB, MEL, NSW); Rainbow

Beach Rd towards Rainbow Beach, c, 300 m inside

Cooloola NP opposite sandstone hill, 26°01’S 153°00’E,

Sep 1992, Duretfto 258-60, Bayly & Marsh (258 - BRI,

MEL, NSW; 259, 260 - MEL); Wide Bay, E side of

Cooloola Coast Rd, 49 km S of Maryborough, 25°56’S

152°S1’E, Sep 1989, Jobson 930 & Lum (MEL); Cooloola

NP between Camp Milo & Freshwater Ck, 26°0-’S 153°0—

"B, Jun 1970, McDonaid476 (BRI, CANB); Eiliot R., near

Bundaberg, May 1967, O/se 330 (NSW); Fraser Is., Lake

Boemingen, | km § of lake along Dilt Village walking

tract, Oct 1982, Parish s.n. (WEL); 2.8 km S of Rainbow

Beach, Cooloola NP, 25°58’S 153°09’E, Sep 1986, Ross 3196

(AD, MEL); Fraser Is., between Lake Birrabeen & Lake

Boemingen, 25°32’S 153°04’E, Aug 1971, Smith'7 (MEL);

Fraser Is., southern half, 1.5 km W of Lake Boemingen

camping area, 25°33’°S 153°04’E, Aug 1984, Walsh i399

(MEL); Moreton District: Collingwood Park near

Ipswich, 27°37’°S 152°52’E, Jul 1990, Bird s.n. (BRI,

CANB, MEL); Mt Tamborine, May 1930, Cheel s.n.

(NSW); 4 km S of Sunshine Beach turnoff along coast Rd

S of Noosa Heads, 100m along track heading W opposite

car park, 26°28’S 153°06’E, Sep 1992, Duretto 253-7,

Bayly & Marsh (253, 255 - MEL; 254, 257 - BRI, MEL;

256 - BRI, MEL, NSW); Miami, south coast, Sep 1965,

Jones 3060 (CANB); North Stradbroke Is., ¢. 27°28’S

153°30°E, Aug 1970, Moriarty 415 (CANB); Moreton Is.,

Aug 1855, Muellers.n. (MEL, TCD); Near Dunwich, North

Stradbroke, Sep 1941, Perry s.n. (BRI); Sunnybank, 8

miles E of Brisbane, Aug 1930, White & McKie s.n.

(NSW); Karawatha bushland, 1-1.5 km WNW of Trinder

Park Railway Station, Woodridge, Jui 1982, Willis s.n.

(MEL); New South Wales, NortH Coast: Fortis Ck, 24

km N of Grafton on the road to Coaldale, Aug 1985,

Foreman 907 (CANB, MEL); Property of Mr A. Ford at

Whiteman Ck near Copmanhurst, Jul 1979, Grieves s.n. (NSW).

Typification: The type of B. rosmarinifolia has

not been seen by the author but it should be tn

W where Endlicher worked. There is no

confusion regarding application of this name

however as B. rosmarinifolia 1s the only member

of Boronia sect. Valvatae occurring in the

Moreton Bay area of Queensland. The only

other member of this section found close to the

Moreton Bay area is B. keysii Domin (Cooloola

sand mass) which has pinnate or rarely simple,

broad, flat, petiolate leaves with a sparse

indumentum.

Taxonomy: Bentham (1863) reduced B.

rosmarinifolia to varietal rank under

B. ledifolia which was followed by Bailey

(1899) in his Queensland Flora and later in

his various catalogues of Queensland plants

(e.g. Bailey 1913). Cheel (1928) reinstated

B. rosmarinifoliato specific rank, which is the

status accepted in this paper. Cheel (1928) also

described B. rosmarinifoliavar. albiflora Cheel.

This variety was based on material of

B, ledifolia s. str. and so 1s not discussed

further here (see Duretto submitted).

Notes: Normally this species has glabrous fruit

but two collections (Baxter & Lebler 1132 [BRI

NSW]; Perry s.n., Sep 1941 [BRI]) have

278

densely hirsute fruit as with fruits of B. forsteri

and B. glabra, The presence of hirsute fruit in

these two specimens of B. rosmarinifoliais not

considered to be of any significant taxonomic

importance. Boronia rosmarinifolia 1s

distinguished from 8B. forsteri by its larger

flowers and leaves, reflexed anther apiculum

and usually glabrous fruits, from B. splendida

and B. palasepala by its smaller flowers and

leaves, and from #8. chartacea P.H.Weston

(North Coast, NSW) by its sessile leaves.

Distribution and ecology: Found in coastal and

near coastal areas from Bundaberg, Wide Bay

District, Queensland, to Grafton, North Coast,

New South Wales (Fig. 1). Common in coastal

heath (wallum) and woodland communities on

well drained sand and sandstone derived soils.

Flowering and fruiting material collected from

May to December.

Conservation status: Common, widespread and

found in several conservation reserves. Under no

immediate threat except local extinction in and around

Brisbane and the Gold Coast of Queensland.

Etymology: The specific epithet refers to the

leaves that are similar to those of species of

Rosmarinus L. (Lamiaceae).

ine

Sees Hames a

Sort eer ees

Austrobaileya 5(2): 263-298 (1999)

2. Boronia splendida Duretto, sp. nov. a

oronia rosmarinifolia A.Cunn. ex Endl.

oliis angustissimis revolutis, et floribus

grandioribus (petalis 8-13 non 5—7.5 mm

ongis) differt. Typus: Queensland.

Moreton District: Falls Ck, 4 km NW of

Haldon, Helidon 9342—084285, 27°45’S

152°04’E, 2 October 1988, PI. Forster

4762 & LH. Bird (holo: MEL [MEL

1575271]; iso: AD [AD 99120272], BISH

(n.v.), BRI [AQ429500], CANB [CBG

8908090], K (n.v.), MO (n.v.) Fig. 9G-L).

Erect, much branched shrub to 2.5 m tall.

Multiangular stellate hairs sessile, with 5-10

rays; rays unicellular, free, firm, straight, to

0.05(—0.1) mm long, glossy, smooth, white to

yellow. Branches slightly quadrangular in TS,

not glandular, with little or no cork

evelopment, with a dense stellate indumentum,

ecoming glabrous with age; decurrent leaf

ases absent. Leaves simple, not conspicuously

landular, sessile, linear to narrowly elliptic, 9—

0 mm long, 1~2(—4) mm wide, with tip obtuse,

ase attenuate, strongly discolourous, paler

eneath, lamina with palisade and spongy

mesophyll; margins entire, strongly revolute;

so ung oO Oo

ale

pire

peek

aes

Rae

ieee

Sethe cin

sere en te

perenne

Senet

soientee.

Fig. 10. Scanning electron micrographs of Boronia seed surfaces. A—B, Boronia odorata. Duretto 285 et al. (MEL); A

x 14, B x 250). C-D B. hoipolloi. Clarkson 10473 (BRI); C x 19, D x 300.

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 279

midrib raised abaxiaily, with tightly packed

parenchyma without secondary thickening

between midvein and abaxial epidermis,

impressed adaxially; adaxial surface glabrous

or with few hairs along midrib; abaxial surface

with a dense indumentum of two hair types, a

moderate layer of multiangular stellate hairs

over a dense layer (lacking or sparse cover on

midrib) of peltate stellate hairs. Inflorescence

1(—3)-flowered, with a dense stellate

indumentum; peduncle 0—0.5 mm long,

deciduous with flower; prophylls unifoliolate,

0.5—3 mm long, to 0.5 mm wide, with a dense

stellate indumentum, or as leaves; metaxyphylls

to 0.5 mm long; anthopodium 2—6 mm long.

Sepals (Fig. 91) ovate-deltoid, 2.5—4(-6) mm

long, 2-4 mm wide, enlarging slightly with

mature fruit, with tip acute; adaxial surface

densely and minutely pubescent, becoming

glabrous towards base; abaxial surface with a

dense stellate indumentum. Petals pink to white,

(6—)8—-13 mm long, 4.5-6 mm wide, enlarging

to 12-14 mm long and 6-7 mm wide with

mature fruit, with midvein raised abaxially;

adaxial surface moderately simple pubescent;

abaxial surface with a moderate stellate

indumentum. Stamen filaments clavate, tapering

to anther connective, densely covered with stiff

simple hairs abaxially and on margins below

glandular tip; antesepalous filaments c. 1.5 mm

long, the distal c. 0.5 mm prominently glandular

(Fig. 9J); antepetalous filaments slightly

tuberculate, c. | mm long (Fig. 9K). Anthers

monomorphic; anther appendage large,

reflexed, glabrous. Disc entire, not surrounding

base of filaments, glabrous (Fig. 9L). Ovary

glabrous (Fig. 9L). Style glabrous or hirsute.

Coccus 5-6 mm long, 2.5-3 mm wide, glabrous.

Seeds black, shiny, c. 4mm long, c. 2 mm wide,

with adaxial side without a ridge; elaiosome

yellow-white; surface at magnification as with

B. odorata (see Fig. 10A,B).

Additional specimens examined: Queensland. BURNETT

District: Stallworth Rd, north of Proston, 26°07’S

151°36°E, Sep 1996, Bean 10670 (MEL); Mundubbera,

9146—-374346, 1.5 km W of ‘Mimosa’ Homestead, 25°54’S

151°23°E, Sep 1985, Forster 2243 (BRI); 8 km W of

‘Manar’, Homestead, Boondooma, 9145—303219, 26°O1’S

1S1°18°E, Aug 1988, Forster 4647 (BRI, CANB); Beeron

Holding, 5 km W of Toondahra Homestead, 25°58’S

151°20°E, Sep 1992, Forster 11202 & Sharpe (BRI, MEL);

Beeron Holding, 25°59’S 151°20’E, Sep 1996, Forster

19603 & Ryan (MEL); Darina Downs District: 4.8 km

E of Tara turn off, & 5.3 km E of Kogan on Condamine

Hwy, near dog fence, c. 27°02’S 150°46’E, Sep 1992,

Duretto 337-344, Bayly & Marsh 337 - AD, BRL CANB,

MEL, NSW, PERTH; 338 - BRI, CANB, MEL; 339, 342—

344 - MEL; 340 - BRI, CANB, MEL, NSW; 341 - BRI,

MEL, NSW); Darling Downs, Lace s.n. (MEL); c. 2 miles

E of Kogan, on the Condamine Hwy, Aug 1961, Phillips

s.n. (CANB); Condamine Hwy, near dog fence, Sep 1964,

Shoobridge s.n. (BRI {AQ15118], CANB, DNA); Dalby-

Condamine, Sep 1964, Shoobridge s.n. (CANB

[CBG15711]); 3 mies c. SE of Kogan, 27°02’S 150°46’E,

Oct 1940, Smith & Everist 817 (MEL); c. 29 miles WNW

of Dalby, near grid on the Condamine Hwy, 27°0-’S

150°4-"E, Sep 1968, Smith 14102 (BRI, DNA);

Condamine, 26°56’S 150°08’E, Jul 1964, Ward s.n.

(PERTH); On Condamine Hwy near rabbit fence, Oct 1984,

Williams 84159 (BRD); Moreton District: East Egypt, 25

km SW of Gatton, 27°40’S 152°07’°E, Oct 1991, Bird s.n.

(BRI, CANB); East Egypt, 16 km SW of Gatton, 27°40’S

152°07°B, Mar 1992, Bird & Pahl s.n. (BRI, CANB).

Notes: The tall inland form of B. rosmarinifolia

referred to by Lebler (1972) probably is

B. splendida. Boronia splendida is closely

related to B. forsteri, B. palasepala and

B. rosmarinifolia from which it can be

distinguished by its tall stature, comparatively

long and narrow leaves with revolute margins,

and large flowers. A Proston specimen (Bean

10670) has smaller flowers and a smaller anther

appendage than other collections but its strictly

revolute, narrow leaves and small hairs

identifies it as B. splendida. Further research

and collections, preferably of several plants per

population, of B. spendida are required in the

northern part of its range to ascertain whether

or not the specific distinction between B.

splendidaand B. palasepala, as described here,

is warranted. |

Distribution and ecology: Occurs in the

Condamine-Kogan area, and north to ‘Mimosa’

homestead c. 50 km S of Mundubbera (Fig. 1).

Found on sandstone derived soils in eucalypt

and acacia woodland. Flowering material

collected from March to November; fruiting

material in November.

Conservation status: Though found over a wide

area, collections of 8B. splendida are

geographically isolated and populations at each

site are small. This species is not known to occur

in any reserves and a ROTAP code of 2R is

therefore appropriate.

Etymology: The specific epithet is derived from

Latin, splendidus (splendid, showy, striking),

and refers to the spectacular display of

comparatively large flowers by this species.

280

3. Boronia palasepala Duretto, sp. nov. a

Boronia rosmarinifolia A.Cunn. ex Endl.

sepalis majoribus ((3—-)4—6 non 2—4 mm

longis, 2—4 non 1.5—2.5 mm latis) ad api-

ces acuminatis, petalis longioribus (8—

10.5 non 5—7.5 mm longis), et antheris

non-apiculatis differt. Typus: Queens-

land. Burnett District: Coominglah State

Forest 28, c. 24°51’30"S 150°56’E, Grid

Ref. 9048—-916493, 6 September 1992,

MF. Duretto 277, M. Bayly & N. Marsh

(holo: MEL [MEL 2036610]; iso: AD,

BRI, CANB, HO, K, MEL [MEL

2036611, MEL 2036612], NSW, PERTH)

(Fig. 9M—R).

Erect, much branched, rounded shrub to 2 m

tall. Multiangular stellate hairs sessile, with 5—

10+ rays; rays unicellular, free, firm, straight,

to 0.25(-0.5) mm long, glossy, smooth, white

to yellow or red. Branches slightly quadrangular

in TS, not glandular, with little or no cork

development, with a moderate to dense stellate

indumentum, becoming glabrous with age,

branches will regrow from a rootstock;

decurrent leaf bases absent. Leaves simple, not

conspicuously glandular, sessile, elliptic to

obovate, 14-42 mm long, 2—6 mm wide, with

tip obtuse, base attenuate, strongly discolourous,

paler beneath, lamina with palisade and spongy

mesophyll; margins entire, flat to recurved

(revolute on drying); midrib raised abaxially,

with tightly packed parenchyma without

secondary thickening between midvein and

abaxial epidermis, impressed adaxially; adaxial

surface glabrous or with few hairs along midrib;

abaxial surface with a dense indumentum of two

hair types, a moderate layer of multiangular

stellate hairs over a dense layer of peltate stellate

hairs. Inflorescence 1(—3)-flowered, with a

moderate to dense stellate indumentum;

peduncle to 0.5 mm long, deciduous with

flower; prophylls unifoliolate, 1-3 mm long,

0.5-1 mm wide, with a dense stellate

indumentum, or as leaves; metaxyphylls minute,

to 1.5 mm long; anthopodium I—3(—5 mm in

Biloela specimens) mm long. Sepals (Fig. 90)

broadly ovate-deltoid, (3—-)4—6 mm long, (2—)3—

4 mm wide, with tip acuminate to acute; adaxial

surface densely and minutely pubescent,

becoming glabrous towards base; abaxial

surface with a dense stellate indumentum. Petals

pink to white, 8—10.5 mm long, 4.5—6 mm wide,

Austrobaileya 5(2): 263-298 (1999)

with midvein raised abaxially; adaxial surface

moderately simple pubescent; abaxial surface

with a moderate stellate indumentum. Stamen

filaments bearing stiff simple hairs abaxially and

on margins below glandular tip; antesepalous

filaments clavate, tapering to anther connective,

c. 2mm long, the distal 0.5—1 mm prominently

glandular (Fig. 9P); antepetalous filaments c.

1.5 mm long, the distal end glandular (Fig. 9Q).

Anthers monomorphic; anther appendage

absent or minute. Disc entire, not surrounding

base of filaments, glabrous (Fig. 9R).

Gynoecium glabrous (Fig. 9R). Fruit and seed

not seen.

Additional speciinens examined: Queensland. BURNETT

District: Coominglah State Forest 28, c. 24°51.5 §

150°56’E, 9048-9 16493, Sep 1992, Duretto 275, 276, 278,

279, Bayly & Marsh (275 - BRI, CANB, MEL, NSW; 276

- BRI, K, MEL, NSW; 278 - AD, BRI, HO, NSW, MEL,

PERTH; 279 - BRI, CANB, K, MEL, NSW); ibid, 24°S1’S

150°57°E, 9048-914493, Jul 1990, Forster 6961 (BRI);

ibid, 24°55’S 150°59’E, 9048971425, Jul 1990, Forster

6906 (BRI, CANB, MEL, NSW); Coominglah SF28,

boundary between compartments 18 & 33, 14 km SW of

Monto, 9048-KT982410, Aug 1976, Afartensz 1014

(CANB); 15 km NE of Biloela, 3 km N of Callide dam,

Jul 1992, Thompson BIL10 (AD, PERTH).

Notes: Boronia palasepalacan be distinguished

from the other members of the B. rosmarinifolia

species complex by its comparatively large

flowers, usually wide leaves with recurved

margins (which can become revolute on drying)

and spade-shaped sepals.

Distribution and ecology: Occurs near Biloela

and in Coominglah State Forest (SF28, near

Monto), Queensland (Fig. 1). Found growing

on sandstone in eucalypt open forest or

woodland where it can dominate the

understorey. Flowering material collected from

July to September.

Conservation status: Boronia palasepala is

known from few small populations outside

existing conservation reserves; a ROTAP code

of 2R is therefore appropriate.

Etymology: The specific epithet is derived from

Latin pala (spade) and sepala (sepal), and

alludes to the spade shaped (as of playing cards)

sepals (Fig. 9O).

4. Boronia forsteri Duretto, sp. nov. a Boronia

rosmarinifolia A.Cunn. ex Endl. petalis

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 281

et sepalis minoribus (2—2.5 non 2-4 mm

longis) et coccis hirsutis differt. Typus:

Queensland, LEICHHARDT District: 7 km

past Glenhaugton Homestead on Mapala

Rd, SF46, 25°21’°S 149°19’E, 10

September 1992, PI. Forster 11235 &

PR. Sharpe (holo: MEL [MEL 2049140];

iso: BRI [AQ561403], NSW) (Fig. 9S—X).

Boronia sp. (Robinson Gorge P.I. Forster+

PIF11235) (Forster 1997),

Erect, much branched shrub to 1(—2) m tall.

Multiangular stellate hairs sessile, with 5—10

rays; rays unicellular, free, firm, straight, to 0.1

mm long, glossy, smooth, white to yellow.

Branches terete to slightly quadrangular in TS,

not glandular, with little or no cork

development, with a moderate to dense stellate

indumentum, becoming glabrous with age;

decurrent leaf bases absent. Leaves simple, not

conspicuously glandular, sessile, elliptic to

obovate, 6—25 mm long, 0.5—5 mm wide, with

tip obtuse, base attenuate, strongly discolourous,

paler beneath, lamina with palisade and spongy

mesophyll (fresh material unavailable); margins

entire, flat or slightly recurved; midrib raised

slightly abaxially, with tightly packed

parenchyma without secondary thickening

between midvein and abaxial epidermis,

impressed adaxially; adaxial surface glabrous

or with few hairs along midrib; abaxial surface

with a dense indumentum of two hair types, a

moderate layer of multiangular stellate hairs

over a dense layer of peltate stellate hairs;

juvenile leaves to 35 mm long, abaxial surface

glabrous or with a sparse indumentum.

Inflorescence 1(—3)-flowered, with a

moderate to dense stellate indumentum;

peduncle to 0.5 mm long, deciduous with

flower; prophylis unifoliolate, 1.5—2.5 mm |

long, to 0.5 mm wide; metaxyphylls to 0.5

mm long; anthopodium 1.5-3 mm long.

Sepals (Fig. 9U) ovate-deltoid, 2—-2.5 mm

long, 1—1.5 mm wide, enlarging slightly to 3

mm long with mature fruit, with tip acute;

adaxial surface densely and minutely

pubescent near margins, becoming glabrous

towards base; abaxial surface with a dense

stellate indumentum. Petals pink, 4-6 mm

long, 2-3 mm wide, enlarging to 7-8 mm

long and 5 mm wide with mature fruit, with

midvein raised abaxially; adaxial surface

sparsely simple pubescent; abaxial surface

with a moderate to dense stellate

indumentum. Stamen filaments bearing stiff

simple hairs abaxially and on margins below

glandular tip; antesepalous filaments clavate,

tapering to anther connective, c. 1.5 mm long,

the distal c. 0.5 mm prominently glandular

(Fig. 9V); antepetalous filaments c. 1 mm

long, the distal end glandular (Fig. 9W).

Anthers monomorphic; anther appendage

large, erect, glabrous. Disc entire, not

surrounding base of filaments, glabrous (Fig.

9X). Gynoecium glabrous (Fig. 9X), Coccus

5—6 mim long, 2.5—-3 mm wide, with a

moderate to dense indumentum of erect,

simple hairs. Seeds black, shiny, 4.5—5 mm

long, 2—2.5 mm wide, with adaxial side

without a ridge; elaiosome yellow-white;

surface at magnification as with B. odorata

(see Fig. 10A,B).

Additional specimens examined: Queensland.

LEICHHARDT District: Marlong Arch-Thombs area,

25°0S’S 147°52’E, Sep 1978, Benyon s.n. (CANB);

Gwambagwine, Ruined Castle Ck catchment,

25°13’°08"S 149°27°02”E, Sep 1995, Forster 17836,

Fige & Carter (MEL); Gwambagwine, Ruined Castle

Ck catchment, 25°12’43”S 149°28°11”E, Sep 1995,

Forster 17851, Figg & Carter (MEL); 5 km past

Glenhaugton Homestead on Mapala Rd, SF46, 25°21°S

149°09°E, Apr 1992, Forster 9753 & Manchin (MEL);

Robinson Gorge NP, northern end in headwaters of

Gienhaugton Ck in Murphy Range, 25°12’S 149°07°E,

Sep 1992, Forster 11429 & Sharpe (BRI, MEL);

Robinson Gorge NP, near Starckvale Ck campsite,

25°18’S 149°11’E, Sep 1992, Forster 11244 & Sharpe

(MEL); Get Down section, Robinson Gorge,

Expedition NP, 25°18’08”"S 149°11’°23”E, Sep 1995,

Forster 17696 & Figg (MEL); Starckvale Creek,

Expedition NP, 25°18’34"S 149°10°53”K, Sep 1995,

Forster 17714 & Figg (MEL); 11 km past Glenhaugton

Homestead on Mapala Rd, 25°18’S 149°17°E, Sep

1992, Forster 11453 & Sharpe (BRI, MEL); 11.8 km

N of ‘Yoothapinna’, Injune District, 25°15’S 148°20’E,

Sep 1974, Gittins 2745 (BRI, NSW); 117.5 km S of

Bauhinia Downs on Glenhaughton Rd, 25°17°20"S

149°16°52”"E, Oct 1996, Hilf 4863 (MEL, NSW); 21

miles SE of Bedourte, Oct 1963, Speck 1854 (BRI);

500m N of Robinson Gorge, c. 25 km NW of

‘Glenhaughton’ Homestead, 25°11’S 149°12’E,

Telford 5635 (CANB); Mt Moffatt section of

Carnarvon NP, behind Tombs Bluff, Sep 1986, Thomas

138 (CANB); Mt Moffatt “The Tombs’, Sep 1986,

Williams 86097 (BRI).

Notes: Boronia forsterican be distinguished

from B. rosmarinifolia, B. splendida and B.

palasepala by its smaller floral parts, erect

anther apiculum and hirsute cocci. The

282

distributions of B. forsteri and B. glabra (a

simple leaved species) may overlap in the

Carnarvon Ranges (Duretto 1995,

submitted), These two species both have

hirsute cocci and the stamens and sepals of

each are similar in size and shape. Boronia

forsteri can be distinguished from B. glabra

by having a dense indumentum on the abaxial

surface of the leaves, as opposed to the

glabrous leaves of B. glabra (at least in

Queensland).

Distribution and ecology: Occurring on the

Chesterton, Carnarvon and Expedition

Ranges, and the Central Highlands of

Queensland (Fig. 1). Found in dissected

sandstone country in eucalypt open woodland

or forest. Flowering and fruiting material

collected in September and October.

Conservation status: Boronia forsteri

occurs in Expedition Range and Carnarvon

National Parks; a ROTAP conservation code

of 2RC- is therefore appropriate.

Etymology: This species is named in honour

of Paul Forster (BRI) whose prolific and

untiring work, including collection of an

impressive number of specimens for world-

wide herbaria (often from remote and poorly

collected areas), has increased our

knowledge of the flora of Queensland and

adjacent tropical areas considerably.

5. Boronia jensziae Duretto, sp. nov. a Boronia

rosmarinifolia A.Cunn. ex Endl. folltis

petiolatis, late ellipticis, et sepalis

acuminatis, et a B. bella Duretto, B.

excelsa Duretto et B. foetida Duretto

indumento adaxiali petalorum sparso

differt. Typus: Queensland. Cook

District: c. 300 mS of Banksia Bay turn

off along the East Coast Trail between

Little Ramsey & Zoe Bays, Hinchinbrook

Is., 18°21.73’S 146°18.65’E, 29 May

1993, M. Duretto 406 (holo: MEL [MEL

2037448]; iso: AD, BRI, CANB, DNA,

K, MEL [MEL 2037449], NSW) (Fig.

11 A—-F).

Boronia sp. ‘Hinchinbrook Is.’ (Thomas &

McDonald 1989).

Austrobaileya 5(2): 263-298 (1999)

Boronia sp.1 (Hinchinbrook Island; S.L.

Everist 7786) (Briggs & Leigh 1996),

Boronia sp. (Hinchinbrook Is. S.L. Everist

7786) (Forster 1997).

Illustration: K.A.W. Williams, Native Pl.

Qid 2, 58 (1984) (as Boronia sp.)

Erect, much branched shrub to 2 m tall.

Multiangular stellate hairs sessile, with 8—15

rays; rays unicellular, free, firm, straight, 0.05—

0.1(—0.25) mm long, glossy, smooth, white to

yellow. Branches terete to slightly quadrangular

in TS, not glandular, with little or no cork

development, with a dense stellate indumentum,

becoming glabrous with age, will regrow from

a rootstock; decurrent leaf bases absent. Leaves

simple, not conspicuously glandular, subsessile

to petiolate; petiole 2-4 mm long; lamina

elliptic, (10—)15—45 mm long, (4—)6—-11.5 mm

wide, strongly discolourous, paler beneath, with

palisade and spongy mesophyll, with tip acute

and +: mucronate, with base strongly attenuate;

margins entire, flat to slightly recurved; midrib

prominently raised abaxially, with tightly

packed parenchyma with secondary thickening

between midvein and abaxial epidermis,

impressed adaxially; adaxial surface glabrous

or with few hairs along midrib; abaxial surface

with a dense indumentum of two hair types, a

moderate layer of multiangular stellate hairs

over a dense layer of peltate stellate hairs.

Inflorescence |-flowered, with a dense stellate

indumentum; peduncle 0.5—1 mm long,

deciduous with flower; prophylls unifoliolate,

2—2.5 mm long, 0.5—1 mm wide, with a dense

stellate indumentum, or as leaves; metaxyphylls

0.5—1 mm long; anthopodium 2—5 mm long.

Sepals (Fig. 11C) broadly ovate-deltoid, c. 4

mm long, c. 2.5 mm wide, not enlarging

significantly with mature fruit, with tip

acuminate; adaxial surface densely and minutely

pubescent, becoming glabrous towards base;

abaxial surface with a dense stellate

indumentum. Petals pink to white, 5.5—-7 mm

long, 3-3.5 mm wide, enlarging to 7.5-8.5 mm

long with mature fruit, with midvein raised

abaxially; adaxial surface with a sparse simple

indumentum, becoming glabrous towards base;

abaxial surface with a moderate to dense stellate

indumentum, Stamen filaments bearing stiff

simple hairs abaxially and on margins below

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 283

i ge

Lf } \X

Fig. 11. A-F, Boronia jensziae. A, flowering branchlet; B, flower; C, sepal; D, abaxial view of antesepalous stamen; E,

abaxial view of antepetalous stamen; F, disc and gynoecium. A-F, Duretto 406 (MEL), G—-L, B. excelsa. G, flowering

| branchiet; H, flower; I, sepal; J, abaxial view of antesepalous stamen; K, abaxial view of antepetalous stamen; L, disc and

| gynoecium. G—L, Forster 17248 (MEL). M-R, B. feetida. M, flowering branchlet; N, flower; O, sepal; P, abaxial view of

antesepalous stamen; Q, abaxial view of antepetalous stamen; R, disc and gynoecium. M, Forster 7483 (MEL); N-R,

Duretto 263 (MEL). S—X, B. bella. S, flowering branchlet; T, flower; U, sepal; V, abaxial view of antesepalous stamen; W,

abaxial view of antepetalous stamen; X, disc and gynoecium; S—X, Duretto 269 (MEL). Scale bar: A, G, M, S= 16 mm;

B, H, N, T= 8 mm; C, I, O, U =4 mm; D-F, J-L, P-R, V—X = 2 mm. Figure 1! was prepared by Peter Neish for inclusion

in Flora of Australia vol. 26 (in prep.) and is reproduced here with the permission of the artist and ABRS.

284

glandular tip; antesepalous filaments clavate,

tapering to anther connective, c. 2 mm long, the

distal 0.5—I mm prominently glandular (Fig.

11D); antepetalous filaments, c. 1.5 mm long,

the distal end slightly glandular (Fig. 11E).

Anthers monomorphic; anther appendage

minute to large and reflexed, glabrous. Disc

entire, not surrounding base of filaments,

glabrous (Fig. 11F). Gynoecium glabrous (Fig.

LIF). Coccus 44.5 mm long, 2—3.5 mm wide,

glabrous. Seeds black, shiny, 2.5—3.5 mm long,

1.5-2 mm wide, with adaxial side without a

ridge; elaiosome yellow-white; surface at

magnification as with B. odorata (see Fig.

10A,B). Andy Jensz's Boronia, Hinchinbrook

Boronta.,

Additional spectnens examined: Queensland. NorTH

KENNEDY District: Zoe Bay, Hinchinbrook Is., Aug 1951,

Blake 18857 (BRI, CANB); Mt Diamantina, 18°26’°S

146°18°E, Jul 1991, Cumming 11273 (BRI); Mount

Bowen, Hinchinbrook Is., 18°41°S 146°16’E, Jun 1991,

Cumming 11217 (BRI); c. 300 m S of Banksia Bay turn

off along the East Coast Trail between Little Ramsey &

Zoe Bays, Hinchinbrook Is,, 18°21.73’S 146°18.65°E, May

1993, Duretto 405 & 407 (405 - AD, BRI, MEL, PERTH;

407 - BRI, CANB, MEL, NSW); On the East Coast Trail

between Banksia & Zoe Bays, Hinchinbrook Is.,

18°21.86’°S 146°18.74’°E, May 1993, Duretto 402 &

Vadala (BRI, CANB, MEL, NSW, PERTH); iéid,

18°22.17’S 146°18.86°E, May 1993, Duretto 404 &

Vadala (BRI, MEL); Southern end of Missionary Bay, N

end of Hinchinbrook Is., 18°27°S 146°12’E, Feb 1965,

Everist 7786 (BRI, CANB, MELU, NSW); Hinchinbrook

Is., southern end of Missionary Bay, 18°19’S 146°13’E,

Jun 1979, Thornsborne & Thornsborne 535 (BRD; Zoe

Bay, Hinchinbrook Is., Sep 1967, Thornsborne s.n. (BRI).

Notes: Boronia jensziae 1s closely related to

B. excelsa, B. bella and B. foetida from which

it can be distinguished by having a sparse

indumentum of simple hairs on the adaxial

surface of the petals rather than being glabrous

to glabrescent.

Distribution and ecology: Restricted to

Hinchinbrook Island, north-eastern Queensland

(Fig. 12). A poorly collected species found in

a variety of habitats including Syncarpia Ten.

or eucalypt open forest and montane heath,

from sea level to c. 840 m (summit of Mt

Bowman). Flowering material collected

between February and September; fruiting

material in August and September.

Conservation Status: Briggs & Leigh (1996)

gave a ROTAP code of 2KC- to this taxon but

a ROTAP code fo 2RC+ seems more

Austrobaileya 5(2): 263-298 (1999)

appropriate. Present collections and field

observations by the author indicate that though

B, jensziae does appear to be widespread on

the eastern half of Hinchinbrook Island the

populations are small and often near hiking

trails. Further field research is required to

ascertain the range of this species and to study

the effect of the tourism on the size of the

known populations.

Etymology: This species is named for Andrea

Suzan Jensz, for her support and invaluable

help to the author throughout the Boronia

section Valvatae project.

ahd

Haeis

Pees

Fig. 12. Distribution of Boronia bella (ma), B. ditiganiae

(A), B. excelsa (©), B. foetida (0), B. jensziae (e), B.

odorata (@), B. hoipalloi («), B. quinkanensis (+) and B.

squamipetala (A).

6. Boronia excelsa Duretto, sp. nov. a Boronia

rosmarinifolia A.Cunn. ex Endl. sepalis

acuminatis, et a B. bella Duretto, B.

foetida Duretto et B. jensziae Duretto

foliis sessilis anguste ellipticis differt.

Typus: Queensland. Cook DIstTrRIct:

State Forest 144, Mt Windsor Tableland,

16°15°52”S 145°02’28”E, 11 July 1995,

PI, Forster 17248 & S.J Fige (holo:

BRI; iso: AD, BRI [x2], CANB, DNA,

K, L, MEL [MEL 243038, MEL 249902,

Flieger tS VI RE ee nett -

1A RRSEBI AGS SSDS SES pg ee NE 0 A

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 285

MEL 249903, MEL 2025931], MO,

NSW, PERTH, ORS (Fig. 11G—L).

Boronia sp. (Mt Windsor Tableland P.I.

Forster+ PIF 15225) (Forster 1997).

Erect, much branched shrub to 3 m tall.

Multiangular stellate hairs sessile, with 8—20+

rays; rays unicellular, free, firm, straight,

0.05—-0.1(-0.25) mm long, glossy, smooth, white

to yellow. Branches terete, not glandular, with

little or no cork development, with a dense stellate

indumentum, becoming glabrous with age;

decurrent leaf bases absent. Leaves simple, not

conspicuously glandular, sessile, narrowly

elliptic, 14-60 mm long, 2-6 mm wide, with

tip acute, base attenuate, strongly discolourous,

paler beneath, lamina with palisade and spongy

mesophyll (fresh material not seen); margins

entire, flat to slightly recurved; midrib

prominently raised abaxially, with tightly

packed parenchyma with secondary thickening

between midvein and abaxial epidermis,

impressed adaxially; adaxial surface glabrous

or with few hairs along midrib; abaxial surface

with a dense indumentum of two hair types,

a moderate layer of multiangular stellate hairs

over a dense layer of peltate stellate hairs.

Inflorescence 1-flowered, with a dense stellate

indumentum; peduncle c. 0.5 mm long, deciduous

with flower; prophylls unifoholate, 1.5-2.5 mm

long, 0.5-1 mm wide, with a dense stellate

indumentum, or as leaves; metaxyphylls 0.5—1

mim long; anthopodium 2—4 mm long, Sepals (Fig.

11I) broadly ovate-deltoid, 3 mm long,

1.5 mm wide, with tip acuminate to acute;

adaxial surface densely and minutely pubescent,

becoming glabrous towards base; abaxial surface

with a dense stellate indumentum. Petals pink

to white, 4.5—5 mm long, 2-3 mm wide, with

midvein raised abaxially; adaxial surface glabrous

or glabrescent; abaxial surface with a moderate

stellate indumentum. Stamen filaments bearing

stiff simple hairs abaxially and on margins

below glandular tip; antesepalous filaments

clavate, tapering to anther connective, c. 1.5 mm

long, the distal c. 0.5 mm prominently glandular

(Fig. 11J); antepetalous filaments c. 1 mm long,

the distal end slightly glandular (Fig. 11K).

Anthers monomorphic, apiculum absent. Disc

entire, not surrounding base of filaments,

glabrous (Fig. 11L). Gynoecium glabrous (Fig.

11L). Coccus c. 4.5 mm long, c. 2 mm wide,

glabrous. Seeds black, shiny, 3~3.5 mm long,

c. 1.5 mm wide, adaxial side without a

ridge; elaiosome yellow-white; surface at

magnification as with B. odorata (see Fig, 10A,B).

Additional specimens examined: Queensland. Cook

District: State Forest i144 Mt Windsor Tableland,

16°15’52"S 145°02’28"E, Jul 1995, Forster 17253 & Figg

(BRI, MEL); Spencers Creek, downstream about 2 km from

Forestry Camp, Mt Windsor Tableland, Whypalla SF,

16°15’S 145°7’E, Aug 1988, Hind 56791 & D’Aubert

(NSW); SFR144 (Mt Windsor Tableland), 16°15’S

145°00’E, Jun 1969, Hyland 4784 (BRI, QRS).

Notes: Boronia excelsa 1s closely related to B.

jensziae, B. bella and B. foetida from which it

can be distinguished by its narrow, sessile leaves

and smaller flowers,

Distribution and ecology: Restricted to the

Mount Windsor Tableland, north-eastern

Queensland (Fig. 12). Found growing on

sranite-derived soils in wet sclerophyll and

Syncarpia forests and along rainforest margins.

All collections have been made above 1000 m

in altitude.

Conservation status: As the only known

collections of B. excelsa are from a limited area

within a logging reserve (SFR144) a ROTAP

conservation code of 2R is appropriate. The type

collection was made from a population of c. 40

plants (Forster pers. comm.)

Etymology: The specific epithet 1s derived from

the Latin, exce/sus (high or elevated), and refers

to the comparatively high altitudes where this

species occurs.

7. Boronia foetida Duretto, sp. nov. a Boronia

rosmarinifolia A.Cunn, ex Endl. foltis

petiolatis, late ellipticis, et sepalis

acuminatis, a B. bella Duretto floribus

minoribus (sepalis 2—3.5 non 4.5—3.5 mm

longis, petalis 7-8 non 7-12 mm longis)

et stylis glabris, a B. jensziae Duretto

petalis adaxialiter glabris et a B. excelsa

Duretto foliis petiolatis differt. Typus:

Queensland. Wipe Bay District: Mt

Walsh, 7 km south of Biggenden, Grid

Ref. 9347-046709, 25°34’S 152°03’E, 28

September 1990, PJ. Forster 7483 (holo:

MEL [MEL 1597019]; iso: AD [AD

99135181], BRI [AQ474340], CANB

|CANB 406384], K (n.v.), NSW, PERTH

(n.v.) (Fig, LLM-R).

Boronia sp. (Mt Walsh P.I. Forster+

PIF 17253) (Forster 1997).

Erect, much branched shrub to 2 m tall.

Multiangular stellate hairs sessile, with 8—20+

rays; rays unicellular, free, firm, straight, 0.05—

Q.1(—0.25) mm long, glossy, smooth, white to

yellow (Fig. 13A). Branches terete to slightly

quadrangular in TS, not glandular, with little or

no cork development, with a dense stellate

indumentum, becoming glabrous with age, will

regrow from a rootstock; decurrent leaf bases

absent. Leaves simple, subsessile to petiolate;

petiole 2—7 mm long; lamina not conspicuously

glandular, elliptic to slightly lanceolate, 20-52

Austrobaileya 5(2): 263-298 (1999)

mim long, 7—14 mm wide, strongly discolourous,

paler beneath, with palisade and spongy

mesophyll, with tip acute, with base attenuate;

margins entire, flat to slightly recurved; midrib

prominently raised abaxially, with tightly

packed parenchyma with secondary thickening

between midvein and abaxial epidermis,

impressed adaxially; adaxial surface glabrous

or with few hairs along midrib; abaxial surface

with a dense indumentum of two hair types, a

moderate layer of multiangular stellate hairs

over a dense layer of peltate stellate hairs (Fig.

13A). Inflorescence 1(-—3)-flowered, with a

dense stellate indumentum; peduncle 2—2.5 mm

long, deciduous with flower or rarely persistent;

‘Sena

Pear

Fig. 13. Multiangular stellate hairs of Boronia species; abaxial leaf surface (A, B, D, E), adaxial leaf surface (C), or

abaxial petal surface (F). A, Boronia foetida , x 180, Bean 28 (BRD. B, B. bella, x 55. Duretto 269 et al. (MEL). C—D, B.

quinkanensis, C x 55, D x 170. Clarkson 6914 (MEL). E, B. duiganiae, x 55, Duretto 315 et al. (MEL). F, B. sguamipetala,

x 200. Moreton 631 (BRI).

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 287

prophylls unifoliolate, 1-6 mm long, 0.5—2 mm

wide, with a dense stellate indumentum, or as

leaves; metaxyphylls 0.5—-I mm long;

anthopodium 7-13 mm long. Sepals (Fig. 110)

broadly ovate-deltoid, 2—3.5 mm long, 1.5—2.5

mm wide, enlarging to 4 mm long and 3 mm

wide with mature fruit, with tip acuminate;

adaxial surface densely and minutely pubescent,

becoming glabrous towards base; abaxial

surface with a dense stellate indumentum. Petals

pink to white, c. 7 mm long, c. 4 mm wide,

enlarging to 8 mm long with mature fruit, with

midvein raised abaxially; adaxial surface

glabrous or glabrescent; abaxial surface with a

moderate to dense stellate indumentum. Stamen

filaments bearing stiffsimple hairs abaxially and

on margins below glandular tip; antesepalous

filaments clavate, tapering to anther connective,

c. 2 mm long, the distal 0.5—1 mm prominently

glandular (Fig. 11P); antepetalous filaments c.

1.5 mm long, the distal end slightly glandular

(Fig. 11Q). Anthers monomorphic; anther

appendage large, reflexed, glabrous. Disc entire,

not surrounding base of filaments, glabrous (Fig.

11R). Gynoecium glabrous (Fig. 11R). Coccus

4—5 mm long, 2—3.5 mm wide, glabrous. Seeds

black, shiny, c. 4 mm long, c. 2 mm wide,

adaxial side without a ridge; ela1losome yellow-

white; surface at magnification as with B.

odorata (see Fig. 10A,B).

Additional specimens examined: Queensland. WIDE BAy

District: Mt Walsh near Biggenden, 25°3-’S 151°5—’E,

Jun 1983, Bean 28 (BRD); Gully just below saddie between

Mt Walsh & The Bluff, Mt Walsh NP, 25°34’S 152°03’E,

Sep 1992, Duretto 261-265, Bayly & Marsh (261-MEL;

262-MEL, NSW; 263-BRI, MEL; 264-HO, MEL; 265-

CANB, MEL); Mt Walsh NP, c. 15 km SW of Biggenden,

Sep 1973, Randell sn. (BRI); 13 km S of Biggenden,

25°38 152°0E, Jun 1979, Rayner s.n. (BRD); Mt Waish,

c. 6.5 km S of Biggenden, 25°34’°S 152°02’°E, May 1977,

Telford 5316 (BRI, CANB),

Notes: Boronia foetida was referred to as the

Mt Walsh form of B. rosmarinifolia by Stanley

and Ross (1983). Leaves of B. foetida show

some variation in size. Specimens collected in

montane heath communities have smaller leaves

than those of specimens collected in

the forest communities in gullies at lower

altitudes. This phenomenon is common in

Boronia species and is considered not to be of

any taxonomic significance. Boronia foetida is

closely related to B. be/la from which it can be

distinguished by its smaller flowers, smaller

hairs (Fig. 13A,B), and glabrous styles. It can

be distinguished from B. jenszide by its petals

being glabrous adaxially and from B. excelsa

by its much wider leaves.

Distribution and ecology: Restricted to Mount

Walsh, south of Biggenden (Fig. 12). Found in

a variety of habitats ranging from montane heath

to densely forested gullies. Flowering and

fruiting material collected from May to

September.

Conservation status: A ROTAP conservation

code of 2RC+ is appropriate as the species is

confined to Mt Walsh National Park.

Etymology: The specific epiphet is derived

from Latin foetidus (stinking), and alludes to

the foul smelling foliage of this species (much

more so than that of other members of Boronia

sect. Valvatae). Some collectors have noted the

smell as ‘reminiscent of dead possum’, but to

me the leaves smell like an unpleasant

combination of burnt styrofoam, tar and a very

mature cheese.

8. Boronia bella Duretto, sp. nov. a Boronia

rosmarinifolia A.Cunn. ex Endl. foltis

petiolatis, ellipticus late, et sepalis

acuminatis, et a B. jensziae Duretto,

B. excelsa Duretto et B. foetida Duretto

floribus grandioribus (sepalis 4.5-5.5 mm

longis, petalis 7-12 mm longis)

et stylis hirsutis differt. Typus:

Queensland. Porr Curtis District: Upper

Oaky Ck, Many Peaks Range, c.

24°11.5°S 151°17.5°E, 9149-—263238, 5

Sep 1992, MF Duretto 269, M. Bayly &

N. Marsh (holo: MEL [MEL 2036441];

iso: AD, BRI, CANB [CBG 9604106],

DNA, K, MEL [MEL 2036442], NSW,

PERTH). (Fig. 11S—X).

Boronia sp. Telford CBG7702560

(Batianoff & Dillewaard 1988).

Boronia sp. (Many Peaks Range LR. Telford

CBG7702560) (Forster 1997).

Erect, much branched shrub to 2 m tail.

Multiangular stellate hairs sessile, with 10—20+

rays; rays unicellular, free, firm, straight, 0.1-

0.25(-0.5) mm long, glossy, smooth, white to

yellow (Fig. 13B). Branches terete to slightly

288

quadrangular in TS, not glandular, with little or

no cork development, with a dense stellate

indumentum, becoming glabrous with age, will

regrow from a rootstock; decurrent leaf bases

absent. Leaves simple, subsessile to petiolate;

petiole 2-4 mm long; lamina not conspicuously

glandular, elliptic, 18—35 mm long, 3.5—10 mm

wide, strongly discolourous, paler beneath, with

palisade and spongy mesophyll, with tip acute,

with base attenuate; margins entire, flat to

slightly recurved; midrib prominently raised

abaxially, with tightly packed parenchyma with

secondary thickening between midvein and

abaxial epidermis, impressed adaxially; adaxial

surface glabrous or with few hairs along midrib;

abaxial surface with a dense indumentum of two

hair types, a moderate layer of multiangular

stellate hairs over a dense layer of peltate

stellate hairs (Fig. 13B). Inflorescence 1(-—3)-

flowered, with a dense stellate indumentum;

peduncle 0.5—2 mm long, deciduous with

flower or rarely persistent; prophylls

unifoliolate, 2—5.5 mm long, 0.5—2.5 mm wide,

with a dense stellate indumentum, or as leaves;

metaxyphylls 0.5—2.5 mm long; anthopodium

2—7 mm long. Sepals (Fig. 11U) broadly ovate-

deltoid, 4.5-5.5 mm long, 2—2.5 mm wide, not

enlarging significantly with mature fruit, with

tip acuminate; adaxial surface densely and

minutely pubescent, becoming glabrous

towards base; abaxial surface with a dense

stellate indumentum. Petals pink to white, 7—

8 mm long, 4~-5.5 mm wide, enlarging to 12

min long with mature fruit, with midvein raised

abaxially; adaxial surface glabrous or

glabrescent; abaxial surface with a moderate

to dense stellate indumentum. Stamen

filaments bearing stiff simple hairs abaxially

and on margins below glandular tip;

antesepalous filaments clavate, tapering to

anther connective, c. 2.5 mm long, the distal

0.5—-1 mm prominently glandular (Fig. 11V);

antepetalous filaments c. 2 mm long, the distal

end slightly glandular (Fig. 11W). Anthers

monomorphic; anther appendage large, erect

or reflexed, glabrous. Disc entire, not

surrounding base of filaments, glabrous (Fig.

11X). Ovary glabrous (Fig. 11X). Style hirsute.

Coccus 4.5—6 mm long, 2.5-—3.5 mm wide,

glabrous or with few hairs along suture. Seeds

black, shiny, 4-5 mm long, 2~—2.5 mm wide,

adaxial side without a ridge; elaiosome yellow-

Austrobaileya 5(2): 263—298 (1999)

white; surface at magnification as with B.

odorata (see Fig. 10A,B).

Additional specimens examined: Queensland. Port

Curtis District: Upper Oaky Ck, Many Peaks Range, c.

24°11.5’°S 151°17.5’E, Calliope 9149-263238, Sep 1992,

Duretto 270-273, Bayly & Marsh (270 - BRI, CANB,

MEL; 271 - BRI, CANB, DNA, K, MEL, NSW; 272 -

BRI, MEL, NSW; 273 - BRI, CANB, HO, MEL, NSW,

PERTH); Mt Castletower NP, eastern slopes of Many

Peaks Range, 24°07°417°S 151°18’25”E, Feb 1995,

Forster 16338 (MEL); SF521, Many Peaks Range,

24°12°42”8 151°20°31°E, Feb 1995, Forster 16255

(MEL); Many Peaks Range, Olsen 348 (NSW); Many

Peaks Range, Mt Castletower, 24°10’S 151°17’E, Telford

5479 (BRI, CANB).

Notes: Boronia bella is closely related to 8B.

foetida from which it can be distinguished by

its larger flowers, larger hairs (Fig. 13A,B), and

hirsute styles. Both these species can be

distinguished from #. jensziae by having petals

that are glabrous adaxially and from B. excelsa

by having much wider leaves.

Distribution and ecology: Known only from

the Many Peaks Range near Gladstone (Fig.

12). Found in eucalypt forest and woodland

on granite-derived soils. Flowering material

collected from May to September; fruiting

material in September.

Conservation status: Batianoff & Dillewaard

(1988) considered this species to be rare.

Collections have been made within the Mount

Castletower National Park so the species

does not appear to be threatened. A ROTAP

conservation code of 2RC-— is therefore

appropriate.

Etymology: Vhe specific epithet is derived

from Latin bel/us (beautiful), and refers to

the spectacular displays made by the species

large, deep-pink flowers. :

9, Boronia hoipolloi Duretto, sp. nov. a

Boronia alulata Sol. ex Benth. paginis

ubique dense hirsutis, et a 8B.

quinkanensis Duretto foliolis

angustioribus differt. Typus: Queensland.

BurKE District: Amphitheatre, a

sandstone escarpment c. 27 km north

of Musslebrook mining Camp, 18°21’S

138°09°S, 12 June 1995, JR. Clarkson

10473 (holo: BRI; iso: MEL [MEL

2032037, MEL 2032038]) (Fig. 14A—EB).

. PERE pT Ee Een Cet ea erect 2 nnediis se eiules e e lee Dee h wits

Duretto, Boronia sect. Valvatae (Benth.) Engi. (Rutaceae) in Queensland 289

Pendulous or erect, much branched shrub to

50 cm long, with a dense stellate indumentum

throughout. Multiangular stellate hairs

sessile, with 4—12 rays; rays unicellular, free,

firm, straight, to 0.2 mm long, glossy,

smooth, white. Branches terete to slightly

quadrangular in TS, not glandular, with little

or no cork development, becoming glabrous

with age; decurrent leaf bases absent. Leaves

imparipinnate, with 7-25 pinnae, gradually

increasing in number of pinnae along axillary

branches, not conspicuously glandular, entire

leaf 15—35 mm long, 5—13 mm wide; petiole

winged, 2-5 mm long; rhachis segments

winged, oval, 1.5-6 mm long, c. 0.5 mm

wide; pinnae opposite or sometimes

subopposite, narrowly-elliptic to linear,

subsessile, with tip obtuse, margins entire and

recurved, discolourous, slightly paler

beneath, lamina with palisade and spongy

mesophyll; midrib raised abaxially, with

tightly packed parenchyma between midvein

and abaxial epidermis with secondary

thickening in cells in the layer above the

epidermis only, impressed adaxially; adaxial

surface with a dense stellate indumentum;

abaxial surface with a dense indumentum

of two hair types, a moderate layer of

multiangular stellate hairs over a dense layer

of peltate stellate hairs; terminal pinnae

longer than the most distal lateral pinnae

but shorter than others, 1-8 mm long, 0.5—1

mm wide; lateral pinnae 1—7 mm long,

0.5—1 mm wide. Inflorescence |—5-flowered;

peduncle to 2 mm long, not deciduous with

flower; prophylls unifoliolate or pinnate,

to 2.5 mm long; metaxyphylls minute;

anthopodium 1—4 mm long. Sepals (Fig. 14C)

narrowly deltoid, 2—3.5 mm long, 0.75—1.25

mm wide, not enlarging significantly with

fruit, with tip acute to slightly acuminate;

adaxial surface densely and minutely

pubescent, becoming sparse to glabrous

towards base or hirsute at tip only; abaxial

surface with a moderate to dense stellate

indumentum. Petals pink, 3.5—5 mm long,

1.5—2 mm wide, not enlarging significantly

with mature fruit, with midvein raised

abaxially; adaxial surface with a moderate

simple indumentum, becoming glabrous

towards base; abaxial surface with a dense

stellate indumentum. Stamen filaments

capitate, tapering to anther connective, with

stiff simple hairs abaxially and on margins

below glandular tip; antesepalous filaments,

c. 2 mm long, the distal c. 0.5 mm

prominently glandular (Fig.14D);

antepetalous filaments 1-1.5 mm long, the

distal end slightly glandular or eglandular

(Fig.14E). Anthers more or _ less

monomorphic, appendage absent or minute.

Dise entire, not surrounding base of

filaments, glabrous. Ovary glabrous. Style

hirsute or glabrous. Coccus (fully mature

not seen) c. 3.5 mm long, c. 2 mm wide,

glabrous or glabrescent. Seeds (mature not

seen) grey, dull, 1.5~2 mm long, 1.5—2 mm

wide, adaxial side without a ridge; elaiosome

yellow-white; surface at magnification

composed of collapsed tubercle like units,

these units free and 10-30 um across (Fig.

10 C,D)

Other specitnen examined: Queensland. BURKE

District: Amphitheatre, 40 km (by road) north of

Musslebrook Mining Camp, 18°21’S 138°10°S, May

1995, Jofinson 779 & Thomas (BRI),

Notes: Boronia hoipolloi was reterred to as

‘Boronia aff. alulata (NW Qld, Clarkson

10473)’ by Duretto (1997). It can be

distinguished from B. alulataby having a dense

stellate indumentum on all its parts, from B.

guinkanensis by its narrower leaf pinnae, and

from B. lanuginosa, which is also found in NW

Queensland, by its sepals being shorter and

narrower than the petals, its petals having a

distinctly raised midrib abaxially, and its dull

seed lacking a conspicuous ridge on its adaxial

side.

Seeds of B. hoipolloi are dull and the

structures on its testa appear to be collapsed

tuburcles (Fig. 10C,D), quite unlike those of

most other members of Boronia sect. Valvatae

(cf. Fig. LOA,B, Duretto 1995, submitted,

Duretto & Ladiges 1997, in press).

Interestingly, B. viridiflora Duretto of the north-

western Arnhem Land plateau, which is also a

cliff dwelling species, also has dull seeds with

apparently collapsed tubercles on the testa

(Duretto & Ladiges 1997).

Distribution and ecology: Known only from

two recent collections from The Amphitheatre,

north of the Musslebrook Mining Camp in

290 Austrobaileya 5(2): 263-298 (1999)

Fig. 14. A-E, Boronia hoipolloi. A, flowering branchlet; B, flower; C, sepal; D, abaxial view of antesepalous stamen; E,

abaxial view of antepetalous stamen. A-E, Clarkson 10473 (BRI). F-K, B. quinkanensis. F, flowering branchlet; G,

flower; H, sepal; I, lateral view of antesepalous stamen; J, abaxial view of antepetalous stamen; K, lateral view of a

coccus. F, K, Clarkson 3712 (BRD; G—J, Clarkson 9619 (MEL). L-Q, B. duiganiae. L, flowering branchlet, M, flower;

N, sepal; O, abaxial view of antesepalous stamen; P, abaxial view of antepetalous stamen; Q, lateral view of a coccus. L,

Thomas 137 (BRD; M-P, Duretto 319 (MEL); Q, Storey & Yapp 211 (NSW), R-X, B. ederata. R, flowering branchlet;

S, flower; T, sepal; U, abaxial view of antesepalous stamen; V, abaxial view of antepetalous stamen; W, lateral view of a

coccus; X, seed. R, Bean 2194 (BRI); S—V, Duretto 280 (MEL); W—X, Everist 8033 (CANB). Scale bar: A, F, L, R = 16

mm; B, G, M, S = 8 mm; C, H, K, N, Q, T, W, X =4 mm; D-_E, I-J, O-P, U-V =2 mm. Figures 14F—X were prepared by

Peter Neish for inclusion in Flora of Australia vol. 26 (in prep.) and are reproduced here with the permission of the artist

and ABRS.

se eSATA Saree SU A

Duretto, Boronia sect. Valvatae (Benth.) Enel. (Rutaceae) in Queensland 29]

north-western Queensland (Fig. 12). Found in

crevices in vertical sandstone cliff faces and

on scree slopes (collectors’ notes). Flowering

material collected in May and June; fruiting

material in June.

Conservation status: A ROTAP conservation

code of 2R is appropriate for this species as

the species is apparently common where found

(J. R. Clarkson, pers. comm.; collectors’ notes).

Field research is required to ascertain the size

and extent of the known population, and if

indeed other populations exist elsewhere.

Etymology: The specific epithet, hoipolloi, is

derived from Greek for rabble (hoi polloi or of

polloi), and refers to individuals of the species

being found on the outer parts of an

amphitheatre, where one expects to find ‘the

rabble’ congregating.

10. Boronia quinkanensis Duretto, sp. nov. a

Boronia alulata Benth. paginis ubique

dense hirsutis et sepalis et petalis

subaequilibus vel aequalibus, et a

B. hoipolloi Duretto foliolis latoribus

differt. Typus: Queensland. Cook District:

22.4 km from Kennedy River on the Jedda

Creek Track to King River Station, 15°41°S

143°47°E, 24 June 1981, JAR. Clarkson

3712 (holo: BRI [AQ348406]; iso: CANB

[CANB 372104, CBG 8505343], DNA, K,

MO, NSW [NSW 244358]) (Fig. 14F-Is).

Boronia sp. “Jedda Creek” (J.R. Clarkson

3712); Boronia sp. “Mt Mulligan” (7.

Clarkson 5769) (Thomas & McDonald

1989),

B. sp. (Mt Mulligan, J.R. Clarkson 5301)

(Ross 1994; Forster 1997).

Boronia sp.4 (Mt Mulligan; J.R. Clarkson

5301 (Briggs & Leigh 1996).

Erect, much branched shrub to 2.5 m tall, with

a dense stellate indumentum throughout.

Multiangular stellate hairs sessile, with 7—15+

rays; rays unicellular, free, firm, straight, 0.1—

0.5 mm long, glossy, smooth, white (Fig. 13C,D).

Branches terete to slightly quadrangular in TS,

not glandular, with little or no cork

development, becoming glabrous with age;

decurrent leaf bases absent. Leaves

imparipinnate, (1—)3—11 pinnae, gradually

increasing in number of pinnae along axillary

branches, not becoming unifoliolate with age,

not conspicuously glandular, entire leaf 6-25

mm long, 4-15 mm wide; petiole winged, 1—5

mim long; rhachis segments winged, broader at

distal end, 1.5—-6 mm long, 0.5-2 mm wide;

pinnae elliptic to oblanceolate, subsessile, with

tip obtuse, discolourous, paler beneath, lamina

with palisade and spongy mesophyll; margins

entire, recurved; midrib raised abaxially, with

tightly packed parenchyma between midvein

and abaxial epidermis with secondary

thickening in cells in the layer above the

epidermis only, impressed adaxially; adaxial

surface with a sparse to moderate stellate

indumentum; abaxial surface with a dense

indumentum of two hair types, a moderate layer

of multiangular stellate hairs over a dense layer

of peltate stellate hairs; terminal pinnae longer

than the most distal lateral pinnae

but shorter than others, (2—)6—-15 mm long,

(1-)3—7 mm wide; lateral pinnae (2—)5—11 mm

long, (1-)3—5 mm wide. Inflorescence 1-3

(—9)-flowered; peduncle 1-23 mm long, not

deciduous with flower; prophylls unifoliolate

or pinnate, 2.5—5 mm long, 1.5—3 mm wide;

metaxyphylls to 0.5 mm long; anthopodium

1-10 mm long. Sepals (Fig. H) narrowly

deltoid, 3-5 mm long, 1—-1.5 mm wide, not

enlarging significantly with fruit, with tip acute

to slightly acuminate; adaxial surface densely

and minutely pubescent, becoming sparse to

glabrous towards base; abaxial surface with a

moderate to dense stellate indumentum. Petals

pink to white, 4—5.5 mm long, 2~3 mm wide,

enlarging to 6-7 mm long with mature fruit,

with midvein raised abaxially; adaxial surface

with a sparse simple indumentum, becoming

glabrous towards base; abaxial surface with a

dense stellate indumentum. Stamen filaments

capitate, tapering to anther connective, with

stiff simple hairs abaxially and on margins

below glandular tip; antesepalous filaments,

1.5—2 mm long, the distal 0.5 mm prominently

glandular (Fig. 141); antepetalous filaments

1—-1.5 mm long, the distal end slightly to

strongly glandular (Fig. 14J). Anthers more or

less monomorphic, apiculum present but

minute. Disc entire, not surrounding base of

filaments, glabrous. Gynoecium glabrous.

Coccus 3,5—4.5 mm long, 2—2.5 mm wide,

292

glabrous or glabrescent (Fig. 14K). Seeds

black, shiny, 3-4 mm long, 1.5—2 mm wide,

adaxial side without a ridge; elaiosome yellow-

white; surface at magnification as with

B. odorata (see Fig. 10A,B).

Additional specimens examined: Queensland. Cook

District: Sandy Ck area N of Jowalbinna, 15°43’S

144°18’E, Jul 1990, Bean 1710 (BRI, NSW); Near Laura

R,, 15°45’S 144°39’E, Aug 1974, Byrnes 3079 (BRI,

MEL, NSW); 4 km S of the crossing of Shepherd Ck on

the Maytown Track, 15°47°S 144°16’E, Jun 1992,

Clarkson 9619 & Nelder (BRI, DNA, K, L, M, MBA,

MEL, NSW, PERTH, QRS); 6 km south of Jowalbinna

turn off on the Maytown track, 15°48’S 144°16’E, Nov

1983, Clarkson 5050 (CANB); Mount Mulligan, c. 30

km NW of Dimbulah, 16°48’°S 144°49’E, Jun 1995,

Clarkson 10541 (BRI, MBA, MEL); Mt Mulligan, c. 40

km NW of Dimbulah, 16°52’S 144°S1’E, Apr 1985,

Clarkson 5769 (BRI, CANB, DNA, MBA, MEL, QRS);

ibid, Apr 1987, Clarkson 6914 (DNA, CANB, MBA,

MEL); Mt Mulligan, on the southern plateau of the

mountain, 16°54’S 144°S1’E, Apr 1984, Clarkson 5301

(BRI, CANB, DNA, MBA, NSW, PERTH, QRS); SSW

part of Mt Mulligan, c. 16°53’S 144°S1’E, May 1993,

Duretto 380, 385, 388, 389 & Vadala 380 - MEL; 385 -

BRI, CANB, DNA, K, MEL, NSW, PERTH; 388 - AD,

BRI, MEL; 389 - AD, BRI, CANB, DNA, MEL, NSW);

Foot of cliffs, Mt Mulligan, 16°52’S 144°52’E, Dec 1936,

Flecker s.n. (QRS); The Gorge, Mt Mulligan, Apr 1934,

Flecker s.n, (BRD); 35 km directly SW of Laura, just below

escarpment of Pine Tree Ck, 15°47’S 144°12’°E, Apr

1987, Parris 9198 (BRI, CANB); 35 km SW of Laura,

on plateau leading to escarpment above Brady Ck,

15°47°S 144°13°E, May 1987, Parris 9200 (BRI, CANB,

NSW); c. 42 km directly SSW of Laura, & c. 2 km W of

Maytown track just above escarpment of Mossman Ck,

15°SS5’S 144°18°E, May 1987, Parris 9190 (CANB);

Jowalbinna camp, c. 30 km SSW of Laura, 15°45’S

144°1S°E, Jun 1990, van der Werff 11716 (QRS),

Notes: The Flecker specimen from The Gorge,

Mt Mulligan, collected in April 1934 (BRI),

referred to as B. artemesiifolia F.Muell. (= B.

lanuginosa) by White (1942), is probably the

first collection of B. guinkanensis held in any

herbarium. Both Hnatiuk (1990) and Ross

(1994) were probably either referring to White

(1942) or to incorrectly determined specimens

of B. gquinkanensis when they stated that B.

lanuginosa (includes 8. artemesiifolia) had

been collected in the Cook district of

Queensland. Boronia lanuginosa has only

recently been collected from north-western

Queensland (P.I. Forster pers. comm.; Duretto

submitted).

Boronia quinkanensis 1s not easily

confused with any other species of Boronia in

north-eastern Queensland as it 1s the only

Austrobaileya 5(2): 263-298 (1999)

Species with a dense indumentum throughout.

It is distinguished from B. lanuginosa by its

more ovate leaflets, its sepals never being wider

and rarely longer than its petals, its petals

having a distinctly raised midrib abaxially, and

its seed lacking a conspicuous ridge on its

adaxial side. From 8B. hoipolloi it 1s

distinguished by its much wider leaflets.

Distribution and ecology: Occurs in the

‘Quinkan’ sandstone country south of Laura,

and also on Mt Mulligan (near Dimbulah) to

the south of that (Fig. 12). Found in woodland

and heath, on sandstones. These sandstones,

Mesozoic in origin, are extensive in the Laura

area with an isolated occurrence of the ‘pepper

pot’ type on Mt Mulligan (Keyser & Lucas

1968; Arnold & Fawckner 1980). Surrounding

these sandstones are the Hodgkinson

formations of greywacke, siltstones, shale,

slates etc. (Arnold & Fawckner 1980) on which

B. quinkanensis is not found. Flowering and

fruiting material collected from April to

December.

Conservation status: Briggs & Leigh (1996)

gave a ROTAP conservation code of 3K to this

taxon, but a conservation code of 3R is more

appropriate as the species does not appear to

be under any immediate threat.

Etymology: The specific epithet is derived

from the name of the area where this species is

commonly found, the so-called Quinkan

country.

11. Boronia duiganiae Duretto, sp. nov. a

Boronia lanceolata F. Muell. et B.

odorata Duretto foliis pinnatis cum

indumento adaxialis moderato ad densum

differt. Typus: Queensland. LEICHHARDT

District: Consuelo, 16 miles SW of

Rolleston Township, 1 September 1961,

Lazarides & Storey 116 (holo: CANB

[CANB 112028]; iso: AD [AD

96244143], BRI [AQ 121206], MEL

[MEL 250602], NSW [NSW 238032]}).

Erect, much branched shrub to 2 m tall.

Multiangular stellate hairs sessile, with 10-25+

rays; rays unicellular, free, firm, straight, c.

0.75(-1) mm long, glossy, smooth, becoming

weak, flexuous and dull with age, white to

ATARI OVS To

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 293

yellow (Fig. 13E). Branches terete, not

glandular, with little or no cork development,

with a dense stellate indumentum, becoming

glabrous with age; decurrent leaf bases absent.

Leaves imparipinnate, 1—5 pinnae, gradually

increasing in number of pinnae along axillary

branches, not conspicuously glandular, entire

leaf (6—)13—45 mm long, (3—)6—35 mm wide;

petiole winged, 2-8 mm long; rhachis segments

winged, oval shaped or triangular with distal

end wider, 4-10 mm long, 1—2 mm wide;

lamina slightly to strongly discolourous, paler

beneath, lamina with palisade and spongy

mesophyll; margins entire and flat to recurved;

midrib raised abaxially, with tightly packed

parenchyma with secondary thickening

between midvein and abaxial epidermis,

impressed adaxially; adaxial surface with a

sparse to moderate (rarely dense) stellate

indumentum; abaxial surface with a dense

indumentum of two hair types, a moderate layer

of multiangular stellate hairs over a

dense layer of peltate stellate hairs (Fig. 135);

pinnae elliptic to oblanceolate, sessile to

subsessile, petiolule to 1 mm long, with tip obtuse;

terminal pinnae longer than lateral pinnae, 6-31

mm long, 3—12 mm wide; lateral pinnae 5—17 mm

long, 2.5-8 mm wide. Inflorescence 1-3-

flowered, with a dense stellate indumentum;

peduncle 0.5—] mm long; prophylls unifoliolate

or pinnate, 1-5 mm long, to 1.5 mm wide;

metaxyphylls minute; anthopodium 1—2 mm long.

Sepals (Fig. 14N) ovate-deltoid, 3.5—5 mm long,

2—3 mm wide, not enlarging significantly with

fruit, with tip acuminate; adaxial surface

glabrescent; abaxial surface with a dense stellate

indumentum. Petals pink to white, 6-11 mm long,

3-6 mm wide, enlarging slightly with mature fruit,

with midvein raised abaxially; adaxial surface

with a sparse to moderate simple indumentum

becoming glabrous towards base; abaxial surface

with a moderate to dense stellate indumentum.

Stamen filaments bearing stiff simple hairs

abaxially and on margins below glandular tip;

antesepalous filaments clavate, tapering to anther

connective, 2—2.5 mm long, the distal 0.5—1 mm

prominently glandular (Fig. 140); antepetalous

filaments c. 1.5 mm long, the distal end glandular

(Fig. 14P). Anthers monomorphic; anther

apiculum minute or large and reflexed, glabrous.

Dise entire, not surrounding base of filaments,

glabrous. Gynoecium glabrous. Coccus 4-5.5 mm

long, 2-3 mm wide, with a sparse to moderate

indumentum (Fig. 14Q). Seeds black, shiny, 4—

4.5 mm long, 2—2.5 mm wide, adaxial side without

a ridge; elaiosome yellow-white; surface at

magnification as with B. odorata (see Fig.

10A,B). (Fig. 14 L-Q).

Additional selected specimens (c, 20 collections

examined): Queensland. LetcHHARDT District: Staircase

Range, 22 km SE of Springsure, 24°13’S 148°14’E, Sep

1993, Bean 6910 (MEL); 20 km from Springsure towards

Rolleston, 24°13’S 148°14’E, Sep 1992, Duretto 314-319

(314 - BRI, CANB, MEL, NSW, PERTH; 315 - AD, BRI,

MEL, NSW; 316 - BRI, CANB, MEL, NSW; 317-318 -

BRI, CANB, MEL, NSW; 319 - MEL); Hilltop, 9.35 km N

of Ist Carnarvon Gorge turnoff & 125,35 km N of Injune,

24°32°S 148°31’E, Sep 1992, Duretto 320-324 & Bayly

(320 - BRI, CANB, MEL, NSW, PERTH; 321 - BRI, MEL;

322-333 - BRI, CANB, MEL, NSW; 324 - AD, BRI,

CANB, MEL, NSW); Stonecroft Caves [c. 24°55’S

149°33’E|N of Taroom, Jul 1958, Gray, DMG4370 (BRI);

Ceres holding, 10.8 km (by road) W of Rolleston-Injune

Rd at Christmas Ck Crossing, Springsure 1:250000

(673917), 24°48’S 148°29?’E, Aug 1978, Martensz 1082A

(CANB); Carnarvon Gorge, 25°0—’S 148°1-’E, Aug 1989,

Morley s.n. (BRI); Near Dawson highway on Expedition

Ra., 24°4-’S8 149°0-"E, Aug 1988, Phillips s.n. (BRI);

Rolleston Rd, c. 13 miles from Springsure township, Sep

1962, Storey & Yapp 211 (AD, BRI, CANB, MEL, NSW);

Mt Moffatt section of Carnarvon National Park behind

Tambo Bluff, 25°02’S 147°27°E, Sep 1986, Thomas 137

(BRI); Orion Downs, Muth s.n. (MEL). MARANOA DISTRICT:

‘The Tombs’, Maranoa R, West Branch, Carnarvon NP,

Apr 1981, Blaxwell 1892 (BRI, NSW); Mt Moffatt NP,

25°0-’S 147°5—’E., Sep 1988, Hando 454 (BRI); Mt

Moffatt turnoff to Kenniffs Cave, 25°01’S 147°57°E, Sep

1986, Williams 86083 (BRI).

Notes: Boronia duiganiae1s not easily confused

with any other taxon except B. odorata from

which it can be distinguished by having pinnate

leaves that usually have a moderately dense

indumentum adaxially and hairs with longer

rays (to 1 mm long as opposed to 0.1 mm

long; Fig. 13E), Many specimens of it have

previously been determined as B. obovata

C.T White, which 1s endemic to the Blackdown

Tableland area. Boronia duiganiae has

ovate-deltoid sepals with a dense indumentum

on the abaxial surface (the epidermis is not

visible) which gives the sepa] the light cream

or tan colour, while B. obovata has narrowly

deltoid sepals with a moderate indumentum

on the abaxial surface (the abaxial surface 1s

visible) and are dark brown.

Distribution and ecology: Restricted to the

Great Dividing, Carnarvon and Expedition

294

Ranges, south and south-west of Springsure and

Rolieston (Fig. 12). Found growing in open

woodland or forest on sandstone. Flowering

material collected from February to November;

fruiting material from September to November.

Conservation status: As the species is found in

Carnarvon Gorge National Park and Mt Moffatt

National Park, a ROTAP conservation code of

2RC- 1s appropriate.

Etymology: The species is named in honour of

Dr Suzanne L. Duigan (1924-1993) in

recognition for her long and distinguished career

at the School of Botany, the University of

Melbourne.

12. Boronia odorata Duretto, sp. nov. a

Boronia lanceolata F.Muell. foliis

juvenalibus trifoliolatis, floribus

majoribus (petalis (4—-)6—11 non 2-5 .5(—

7) mm longis) et filamentis hirsutis differt.

Typus: Queensland. LEICHHARDT DISTRICT:

Bull Creek Gorge, 15 km W of ‘Castlevale’,

24°30’°S 146°52’E, 3 September 1990, 4.2.

Bean 2194 (holo: BRI [AQ474979]; iso:

NSW) (Fig. 14R-X).

Erect, much branched shrub to 2 m tall.

Multiangular stellate hairs sessile, with 5—25

rays; rays unicellular, free, firm, straight, 0.05(—

0.1) mm long, glossy, smooth, white to red-

brown. Branches terete to slightly quadrangular

in TS, not glandular, with little or no cork

development, with a dense stellate indumentum,

becoming glabrous with age; decurrent leaf

bases absent. Leaves simple at maturity but

juvenile leaves trifoliolate for several nodes, not

conspicuously glandular, subsessile to petiolate;

petiole winged, 1-8 mm long; pinnae or

unifoliolate leaf elliptic, with tip obtuse, strongly

discolourous, paler beneath, lamina with

palisade and spongy mesophyll; margins entire,

flat to recurved (becoming revolute on drying);

midrib raised abaxially, with tightly packed

parenchyma with secondary thickening between

midvein and abaxial epidermis, impressed

adaxially; adaxial surface with a sparse to

moderate stellate indumentum; abaxial surface

with a dense indumentum of two hair types, a

moderate layer of multiangular stellate hairs

over a dense layer of peltate stellate hairs;

juvenile leaves trifoliolate, initially glabrous,

Austrobaileya 5(2): 263-298 (1999)

becoming more hirsute with each node until as

hirsute as mature leaves; unifoliolate and

terminal pinnae longer than lateral pinnae, (S—)12—

40 mm long, (2-)4-8 mm wide; lateral pinnae 10—

15 mm long, 2-4 mm wide. Inflorescence 1—

3(—7)-flowered, with a dense stellate

indumentum; peduncle 1—2 mm long, not

deciduous with flower; prophylls unifoliolate,

1-4 mm long, 0.5—2 mm wide, with a dense

stellate indumentum or as leaves; metaxyphylls

minute; anthopodium I—7 mm long. Sepals (Fig.

141) ovate-deltoid, 2-4.5 mm long, 1—-2.5 mm

wide, not enlarging significantly with mature

fruit, with tip acute to shightly acuminate; adaxial

surface densely and minutely pubescent,

becoming glabrous towards base; abaxial surface

with a dense stellate indumentum. Petals pink to

white, (4—-)6-10 mm long, 4—6 mm wide,

enlarging to 8-11 mm long and 5—7 mm wide

with mature fruit, with midvein raised abaxially;

adaxial surface moderately simple pubescent;

abaxial surface with a moderate to dense stellate

indumentum. Stamen filaments bearing stiff

simple hairs abaxially and on margins below

glandular tip; antesepalous filaments clavate,

tapering to anther connective, 22.5 mm long,

the distal c. 1 mm prominently glandular (Fig.

14U); antepetalous filaments c. 1.5 mm long, the

distal end glandular (Fig. 14V). Antepetalous

anther slightly larger than antesepalous anthers

before dehiscence; anther apiculum large,

reflexed, glabrous. Disc entire, not surrounding

base of filaments, glabrous. Gynoecium glabrous.

Coccus (4-)5.5~7 mm long, (2-)3—3.5 mm wide,

glabrous or sparsely hirsute (Fig. 14W). Seeds

black, shiny, 3.5—5 mm long, 2.5~-3 mm wide,

with adaxial side without a ridge; elaiosome

yellow-white (Fig. 14X), surface at magnification

tuberculate; tubercles erect, unicellular, 10-44 um

across, free, with surface smooth and anticlinal

walls not visible (Fig. 10 A,B).

Additional selected specimens (c. 40 collections

examined): Queensland. LeIcHHARDT District: 6 miles

W of ‘Mt Playfair’ Station, 24°52’S 146°51’E, Oct 1964,

Adams 1356 (AD, BRI, CANB); 1.5 miles S of Ball Ck

& Robinson Ck junction, Glenhaughton holding, Oct

1974, Clarkson s.n. (BRI); 26 km WSW of Bauhinia

Downs on the Dawson Hwy towards Rolleston, 24°39’S

149°02’E, Sep 1992, Duretto 288-292, Bayly & Marsh

(288 - BRI, MEL, NSW; 289~—291- BRI, MEL; 292 - BRI,

CANB, MEL, NSW); E of car park & camping area, Isla

Gorge NP, 25°12’S 149°59’E, Sep 1992, Duretto 280-

285 (280 - BRI, MEL; 281 - BRI, MEL; 282 - BRI,

LE PE

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 295

CANB, MEL; 283 - MEL; 284-285 - BRI, MEL, NSW);

Isla Gorge, c. 18 miles SW of Theodore, 25°09’S

L49°S7°E, Sep 1968, Everist 8033 (AD, BRI, CANB,

NSW); Glenmore Gap, 13 km WSW of Theodore,

24°S58°S 149°S57°E, Mora Map 8848969354, Sep 1986,

Forster 2637 (BRI, CANB, MEL); 10 km S of Isla Gorge

lookout, 37 km S by road from Theodore, 25°05’S

150°00’E, Jun 1971, Jolmson 7203 & Briggs (BRI,

NSW); Bauhinia Downs, 24°34’S 149°17’E, Feb 1968,

Jones 3729 (CANB); W of Moura, Apr 1961, Jones 1814

(BRI); Watershed 23 miles ESE of Rolleston Township,

24°35’S 148°56’°E, Aug 1961, Lazarides & Storey 112

(BRI, CANB, MEL, NSW). Warreco District: SW

boundary of Chesterton NP, 26°13°S 147°20°E, Jul 1995,

Dollery 84 (BRI); Mt Mobil Holding, 15-20 km W of

Umberill Homestead, 26°14’S 147°25’E, Nov 1990,

Grimshaw CHR20 (BRI). MarAnoa District: SE of Surat,

Thomby Range, May 1960, Blake 21293 (BRI, CANB,

NSW, PERTH); Thomby Range, Glenmorgan-St. Georges

Rd, Aug 1948, Gordon 115 (BRI); Claravale, c. 37 miles

N of Mitchell on stony ridge, May 1962, Johnson 2434

(BRI, CANB).

Notes: Specimens ofa trifoliolate and glabrous

Boronia taxon that were thought to be a

form of B. glabra by Stanley & Ross (1983)

are probably juvenile specimens of B. odorata.

Plants of this species from Isla Gorge

and Thomby Range often have a more dense

stellate indumentum on the adaxial surfaces

of the leaves than do those of the typical

form, and may, with further collections and

research, be found to represent a distinct

taxon. The majority of herbarium specimens

of B. odorata seen have only simple leaves.

Trifololate leaves are produced on the

primary axis only, and then for only a few

nodes. In Boronia sect. Valvatae this

ontogenetic sequence also occurs in

B, paucifiora W.Fitzg. (NW WA) (Duretto

1997), while some normally pinnate leaved

species produce simple leaves as the plant ages,

e.g. B. keysil (SE Qld), B. ledifolia (NSW, Vic.),

B. rupptt Cheel (NSW), and B. ternata Endl.

(SW WA) (Duretto 1995, submitted).

Boronia odorata can be distinguished from

B. duiganiae by its simple mature leaves that

have a sparse to moderate indumentum on the

adaxial surface, and hairs with shorter rays (to

0.1 mm long rather than to | mm long). From

B. lanceolata F.Muell. (NW Qld, N.T.) it may be

distinguished by its larger flowers and pilose

rather than glabrous staminal filaments (Duretto

1997), and from B. jensziae, B. excelsa, B. foetida

and B. bellaby its trifoliolate juvenile leaves and

its sparse to moderate stellate indumentum, as

apposed to being glabrous, on the adaxial surface

of the leaves.

Distribution and ecology: Restricted to the

Central Highlands of Queensland in an area

approximately bounded by Springsure,

Theodore, Surat, Mitchell and Tambo (Fig. 12).

Found in open woodland on sandstone.

Flowering material collected from February to

October; fruiting material from April to

November.

Conservation status: As B. odorata is

widespread, though not evenly collected, and

found in some conservation reserves, e.g. Isla

Gorge and Expedition Range National Parks,

it is not considered to be under threat.

Etymology: The specific epithet is derived from

Latin, odoratus (smelling), and refers to the

unpleasant (to some) tat/coffee odour of the

leaves when crushed.

13. Boronia squamipetala Duretto, sp. nov. a

Boronia bowmanti F. Muell. petalis

majoribus (4-8 non 3—4 mm longis)

indumento denso abaxialiter differt.

Typus: Queensland, Cook District: 19

km from Peninsular development Rd on

a track to Wolverton via the Cook Tin

Mine, 13°21’S 143°3’E, 23 June 1993,

JR. Clarkson 10112 & VJ. Neldner (holo:

MEL [MEL 2036781]; iso: BRI [AQ

621834], K, L, MBA, MEL [MEL

2036782]). :

Boronia sp. “Massy Creek, Rocky River” (R.

Coveny 7174) (Thomas & McDonald

1989).

Boronia sp. 3 (Massy Creek, Rocky River;

R. Coveny 7174) (Briggs & Leigh 1996).

Boronia sp. (Massy Creek R.G. Coveny+

7174) (Forster 1997),

Erect, much branched shrub to 1 m tall.

Multiangular stellate hairs sessile, with 6—23+

rays; rays unicellular, fused and apressed,

appearing peltate at times, firm, straight, 0.1—0.3(—

0.5) mm long, glossy, smooth, white (Fig. 13F).

Branches quadrangular in TS, not conspicuously

glandular, with little or no cork development, with

296

a sparse to moderate stellate indumentum,

becoming glabrous with age; decurrent leaf bases

absent. Leaves imparipinnate, with 5—13 pinnae,

not conspicuously glandular, 33-55 mm long,

12—20 mm wide, glabrescent or with a sparse

indumentum, with hairs mainly on midrib; petiole

winged, 6-15 mm long; rhachis segments winged,

broader at distal end, 2-10 mm long, 1-3 mm

wide; pinnae sessile, elliptic, slightly

discolourous, paler beneath, lamina with palisade

and spongy mesophyll, with tip acute; margins

entire and flat to slightly recurved; midrib not or

slightly raised abaxially, with tightly packed

parenchyma between midvein and abaxial

epidermis with secondary thickening 1n cells in

the layer immediately above the epidermis only,

slightly impressed adaxially; terminal pinnae

longest, 8-20 mm long, 2-6 mm wide; lateral

pinnae 3-13 mm long, 1-3 mm wide.

Inflorescence (1-)3—7- flowered, with a sparse to

moderate stellate indumentum; peduncle 1-2 mm

long, woody, not deciduous with flower; prophylis

linear, unifoliolate or pinnate, 1~3 mm long, 0.5—

1 mm wide; metaxyphyl!s minute, to 0.5 mm long;

anthopodium 2—6 mm long, Sepals ovate-deltoid,

c. 2 mm long, c. 1 mm wide, not enlarging

significantly with fruit, with tip acute; adaxial

surface glabrous to glabrescent with few hairs

along margin at tip; abaxial surface glabrescent

or with a sparse to moderate stellate indumentum,

hairs concentrated at base. Petals white to green,

4—~7 mm long, 2.5-4 mm wide, enlarging to 6-8

mim long with mature fruit, with midvein not

raised abaxially; adaxial surface glabrous or with

a sparse simple indumentum, mainly at tip and

along margins; abaxial surface with a moderate

stellate indumentum with hairs concentrated along

midrib, Stamen filaments bearing stiff simple hattrs

abaxially and on margins below glandular tip;

antesepalous filaments clavate, tapering to anther

connective, c. 1.5 mm long, the distal 0.5—0.75

mm prominently glandular; antepetalous

filaments smooth, c. 1 mm long. Anthers

monomorphic, appendage absent or minute to

large and erect, glabrous. Disc entire, not

surrounding base of filaments, glabrous.

Gynhoecium glabrous. Coccus 4—5.5 mm long,

2.5-3 mm wide, glabrous. Seeds black, shiny, 3—

4 mim long, 1.5—2 mm wide, adaxial side without

a ridge; elaiosome yellow-white; surface at

magnification as with B. odorata (see Fig.

10A,B).

Austrobaileya 5(2): 263-298 (1999)

Additional specimens examined: Queensland. Cook

District: 4.2 km (2.6 miles) by road E of Wenlock R. towards

Pascoe river on Iron Range Rd, 124 km by road NNW of

Coen PO, 13°06 142°59’E, Sep 1975, Coveny7174 & Hind

(BRI, MELU, NSW, PERTH); 13 km along road to Leo Ck

mine, McIlwraith Range, 13°43’S 143°12’E, Jun 1992,

Forster 10098 (BRI, MEL); 3.5 km NNE Massy Ck

crossing, Silver Plains Station, eastern fall of McIfwraith

Range, 13°53’S 143°31’E, Jul 1993, Forster 13618 (CANB,

MEL, NSW); 8 miles from Kennedy Rd on Leo Ck Track,

13°3-’S 143°2-"E, Jul 1968, Gittens 1781 (BRI, CANB,

NSW); Bacon Ck, Archer R., 13°20’°S 142°50’E, Jul 1972,

Hyland 6239 (BRI, CANB, NSW, QRS); 10 miles N of

Archer R. on Kennedy Rd, 13°25’S 142°50’E, Oct 1973,

Hyland 7014 (BRI, QRS); Between Massy Ck & Rocky R.

on Cape York Rd, 13°55’S 143°30°E, Sep 1971, Hyland

5515 (BRI, MEL, QRS); T.R. 14, Leo Ck Rd, 13°40’S

143°20’E, Sep 1972, Jrvine 372 (QRS); Heathiands Pastoral

Station on road between the slaughter yard & the Telegraph

Line read, 11°47°S 142°30’E, May 1980, Morton 631 (BRD;

45 km N of Coen on Cape York Rd, Jun 1972, Wrigley &

Telford NQ1710 (BRI, CANB).

Notes: Boronia squamipetala\s closely related

to B. bowmanii (Duretto & Ladiges in press;

Duretto 1995, submitted) from which it can be

distinguished by its shorter and wider leaflets,

and its larger petals that have a dense, rather

than a sparse to moderate, peltate indumentum

abaxially.

Distribution and ecology: Occurs mainly in the

Iron and MclIllwraith Ranges tn Cape York

Penninsula (Fig. 12). Found in open woodland

or forest and heath on loams, sand, or rock

pavements. Flowering and fruiting material

collected from May to October.

Conservation status: Though this taxon was

given a ROTAP conservation code of 2K by

Briggs & Leigh (1996), because of its wider

geographical range, a code of 3RC- 1s more

appropriate. It is probably represented in Iron

Range and Mclllwraith Range National Parks.

Etymology: The specific epithet is derrved from

Latin, sguamosus (scaly) and petala (petals), and

refers to the scaly appearance of the petals when

viewed at low magnification. This scaly

appearance Is attributable to the fused rays of the

densely packed hairs.

Acknowledgments

I would like to thank Prof. P.Y. Ladiges for her

support and advice; Dr J. Ross for making

available space at MEL for loan material and

advice on nomenclature; the directors and curators

of the various herbaria for loaning material; A.

Duretto, Boronia sect. Valvatae (Benth.) Engl. (Rutaceae) in Queensland 297

Jensz, M. Bayly, N. Marsh, and A. Vadala for

assistance during field work; the Forestry Service

Queensland, Department of Primary Industries,

and the National Parks and Wildlife Service,

Queensland Department of Environment for

permission to collect in Queensland State Forest

Reserves and National Parks respectively; N.

Walsh for completing the Latin diagnoses; P.

Neish for completing figures 9, 12 and 14; and I.

Lambiris for useful comments on the manuscript.

This project was funded by the Australian

Biological Resources Study (grant Ref. N°. 91/

0228).

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WILLIAMS, K.A.W. (1984). Native Plants of Queensland

2. Ipswich: Keith Williams.

320); B. excelsa (Forster 17248); B. foetida

(Duretto 263); B. forsteri (Forster 11429); B.

hoipolloi (Clarkson 10473); B. jensziae

(Duretto 409); B. odorata (Duretto 282, 289);

B. palasepala (Duretto 279); B. quinkanensis

(Duretto 385, Clarkson 9619); B. rosmarinifolia

(Duretto 102, 257); B. splendida (Duretto 337);

B. squamipetala (Clarkson 10112).

Taxonomic status and Australian distribution of the weedy

neotropical grass Leptochloa fusca subsp. uninervia, with an

updated key to Australian Leptochloa (Poaceae, Chloridoideae)

Neil Snow‘ and Bryan K. Simon?

Summary

Snow, Neil & Simon, Bryan K, (1999), Austrobaileya 5(2): 299-305. The neotropical grass Leptochloa

jusca subsp. uninervia is reported for the first time in Queensland, Western Austrailia, Northern Territory,

South Australia and Tasmania, and its weedy tendencies are discussed. A brief overview is given

regarding the taxonomy of Lepftochioa, including why Diplachne P. Beauv. is no longer recognised,

and of the L. fusca species complex. An updated key is provided for the fourteen confirmed taxa of

Australian Leptochloa.

Keywords: Leptochloa, Poaceae, Chloridoideae, Australia, weediness, systematics.

Neil Snow!, Queensland Herbarium, Brisbane Botanic Gardens Mt. Coot-tha, Toowong, Queensland 4066,

Australia, e-mail: Neil. Snow@env.qid.gov.au. Current address: Department of Biological Sciences, University

of Northern Colorado, Greeley, Colorado, 80639 USA. e-mail: nsnow@bentley.unco.edu

Bryan K. Simon’, Queensland Herbarium, Brisbane Botanic Gardens Mt. Coot-tha, Toowong, Queensland

4066, Australia

Introduction

During fieldwork in April of 1996 in Bowen,

Queensland, we encountered a large population

of the neotropical grass Leptochloa fusca (L.)

Kunth subsp. uwninervia (J. Presl) N. Snow. Until

recently (Snow 1997a, 1998; Snow & Davidse

1998) this taxon was recognised as a distinct

species (Gould 1975; McVaugh 1983; Nicora

1995; Snow 1996). Prior to this report L. fusca

subsp. uninervia was only known in Australia

from a single collection in New South Wales

(Jacobs & McClay 1993). Recent monographic

work on the genus worldwide (Snow 1997a) has

confirmed its presence in the Australian states

of Queensland, Western Australia, Northern

Territory, south Australia and Tasmania. Before

discussing the weedy properties and distribution

of this taxon, a brief discussion is necessary

regarding the systematics of the genus, and of

the species complex to which L. fusca subsp.

uninervia belongs.

Accepted for publication 27 May 1998

Systematics of Leptochloa sensu lato

Leptochloa sensu lato frequently has been split

by Australian, African, and South American

workers into Leptochloa s.s. and Diplachne P.

Beauv. (Simon 1993; Gibbs-Russell et al. 1991;

Nicora 1995). However, since cladistic studies

consistently rejected the null hypothesis that

Diplachne represents a monophyletic clade

distinct from Leptochloa, Diplachne has been

reduced to synonymy under Leptochloa (Snow

1997a, 1998).

Systematics and nomenclature of the

Leptochloa fusca complex

Leptochloa fusca subsp. uninervia belongs to

the highly polymorphic species L. fusca, which

occurs worldwide in warm temperate and

tropical regions (Correll & Johnston 1970;

Phillips 1974; Lazarides 1980; Stanley and Ross

1989; Scholz and Boécker 1996; Snow

1997a). The nomenclature associated with

L, fusca has been tortuous and ambiguous. A

few authors have begun using the epithet

‘malabarica’ rather than the much more widely

300

known ‘fusca’, and for this reason

rejection of the epithet ‘malabarica’ has

been proposed (Snow & Davidse 1998).

Morphological forms of L. fusca s.\. that appear

relatively distinct regionally often have been

accorded formal taxonomic recognition at

various ranks. Consequently, dozens of names

have been given to local forms.

The recent revision of Leptochloa (Snow

1997a) reconsidered the systematics of this

species group. Included in the re-evaluation were

several thousand herbarium specimens from all

continents except Antarctica, and fieldwork on

three continents (North America, Africa,

Australia). Univariate statistical studies of

herbarium specimens and population samples

from the field (Snow in prep.) were unable to

find characters that could consistently diagnose

separate species in this group by ordinary

morphological means (Snow 1997b), with the

exception of the rare African species

Leptochloa gigantea (Launert) T. A. Cope & N.

Snow. However, multivariate statistical studies

(Snow in prep.) of population samples (7=20)

from the USA, Mexico, Botswana, Namibia, and

Australia, supported recognition of four

subspecies, given a general tendency of

populations to segregate into four entities. Of

the four recognised subspecies (Snow 1997a),

three occur in Australia, including L. fusca subsp.

uninervia, L. fusca subsp. fusca (nto which

Diplachne parviflora (R. Br.) Benth. and

Diplachne reptatrix (L.) Druce have been

synonymised), and L. fusca subsp. muelleri

(Benth.) N. Snow (Snow 1998).

Weediness and geographical distribution

of Leptochloa fusca subsp. untnervia

The occurrence of Leptochloa fusca subsp.

uninervia Should be of interest to the

agricultural community in Australia, Like several

other species in the genus, L. fusca subsp.

uninervia has pronounced weedy tendencies

(Hafliger & Scholz 1981: 98; McIntyre et al.

1988) and frequently can be seen growing in

mesic soils of agricultural crops (Snow pers.

Austrobaileya 5(2): 299-305 (1999)

obs.). Given the high vagility and high

germination rates of seed in the genus (Snow

unpubl.), Z. £ subsp. wninervia has the potential

to spread rapidly within Australia as a weed. This

potential is made even more likely by its

remarkable salinity tolerance (McVaugh 1983;

Jacobs & McClay 1993) and its ability to grow

in seasonally inundated habitats, properties

common to the species complex to which it

belongs (Snow 1997a). Since the seed can

overwinter in the soil under normal conditions

in a continental climate at ca. latitude 39°N in

St. Louis, Missouri (Snow pers. obs.), its weedy

potential in Australia probably covers the entire

continent at lower elevations. The related and

morphologically variable Leptochloa fusca

subsp. fusca has been amply documented as a

weed of rice crops in Australia (McIntyre 1985;

McIntyre et al. 1989). Leptochloa fusca subsp.

uninervia commonly grows in semi-disturbed,

seasonally inundated locations, and thus ts found

frequently in roadside ditches, along sandbars

of streams and smaller rivers, and in mesic

acricultural situations. Since it does not compete

well ecologically with other species, it often

occupies somewhat bare areas (Snow pers. obs.).

Contrary to the otherwise accurate presentation

in Hafliger & Scholz (1981), Leptochloa fusca

subsp. wninervia is often geniculate below and

frequently roots at the lower nodes (Snow

1997a). The fringed appearance of the ligule

(Hafliger & Scholz 1981) is an artefact of

mechanical damage, the undamaged condition

being apically attenuated.

The native range of the L. fusca subsp.

uninervia is from the southern third of the

United States, the West Indies, and south through

Argentina (Gould, 1975; Hafliger & Scholz,

1981; Nicora, 1995; Snow 1997a). However,

in addition to Australia, this taxon has become

introduced in a number of regions, including

Saudi Arabia, Egypt, the Canary Islands, and New

Zealand (Snow 1997a). Except for the lemma,

which is apically attenuate in its undamaged state

(Fig 1; shown erroneously as bi-lobed),an

excellent illustration of Leptochloa fusca subsp.

SE gS i in ee nin i

Snow & Simon, Taxonomic status Leptochloa fusca subsp. uninervia 301

uninervia from Mason (1957) is reproduced

here (Fig. 1.) by permission of the University of

by Herbert Mason). In the New World L. fusca

subsp. uninerva is commonly called ‘Mexican

Sprangletop’ (Gould, 1975: 229), ‘Zacate Salado

Mexicano’, or ‘Zacate Gigante Peruano’ (Beetle

et al., 1991: 291).

The only known previous Australian

collection (Jacobs & McClay, 1993) was in

August of 1992 from the Newington Naval Arms

Depot, Homebush Bay, on the central coast of

New South Wales (Jacobs 6546, NSW).

Additional Australian localities: Queensland. NortTH

KENNEDY District: Bowen, in ditches along roadside; locally

common in shallow water; GPS 20°00735”, 148°13'40" E; 13

April 1996, Snow 7387& Simon (BRI, MO, NE and duplicates

to be distributed). Porr Curtis District: Awonga Dam, Iveragh

Reach, 15 km SE of Calliope, GibsonTO1347 (BRI). Sout

Kennepy District: Munbura Road, Alligator Creek Mackay,

17 Nov 1994, Tilley s.12.(BRI). New South Wales, Newington

Naval Arms Depot, Homebush Bay, Jacobs 6546 (NSW).

Western Australia, Tank near Milbillillie H/S, Craver 5383

(CANB,MO); Kimberley Research Station, Kununurra,

Parker 471 (BRD; Carawine Gorge, ca 140 km SE of Shay

Gap, Newbey 10463 (CANB); Corong Creek, Woodstock

Station, S of Port Hedland, Burbidge 58454 (CANB);

Department of Agriculture Experimental Farm, Kununurra,

Gilbey s.n. (CANB), Northern Territory, Elparpa Swamp,

Latz 7607 (NSW); Palm Vailey, 12 miS W of Hermannsburg,

Mission, Lazarides 5290 (NSW). South Australia. Barker

Inlet South Wetland, Wingfield, Adelaide, 8 Apr 1997, Green

1988 (BRI), S.A. Water’s Bolivar Sewage Treatment Works,

Bolivar & St. Kilda, Adelaide, 8 Apr 1997, Green 1993 (BRI).

Tasmania. Woodbury, Black 1270.635 (CANB, MO).

Synopsis and key to Australian Leptochloa (sensu Snow 1997)

With the recent discovery of the new species Leptochloa southwoodii in Queensland (Snow &

Simon 1997), a first record of L. panicea subsp. panicea from Mt. Isa (Snow 1997a), and this report

of L. fusca subsp. wninervia, the number of taxa in Leptochloa for Australia stands at fourteen.

Details regarding nomenclature and synonymy will be published 1n the future (or see Snow 1997a).

Taxa in the following key with an asterisk are adventive in Australia.

1. Panicle branches digitate or subdigitate ...

Panicle branches not digitate or subdigitate

2. Culms ‘woody’; leaf blades deciduous at base

Culms not ‘woody’; leaf blades not deciduous at base... .... ee L. dubia*

3. Hidden inflorescences in axils of sheaths present......... 0.0.0 eee eee L. dubia*

Hidden inflorescences in axils of sheaths absent......... 0... eee ce ee eens 4

A OS DIKE ICISRONE =O WEICE Swale <n ass apie k bana Siac, Sorin 3 xtvne tolerant vas 3, eves PM in oo L. neesii

Spikelets TWO- OF ANOTETOWELER 05 ce accede ite eean oe Nok Acie eae we Fin Re en fete oe a 5

5. Ligule apex (undamaged) attenuate, mostly 5 mm or more long...............00005 6

Ligule apex (undamaged) truncate, mostly shorter than 5 mm (except

L: SouthiwoodiHeand £5. 1SU1GLAY conc occu sed .n 4 0 ehh Heedre b 40h sw LOCOS Ob oe 8

6. Lowermost panicle branches not exserted at maturity; uppermost leaf blade

often exceeding length of panicle; lower portion of leaf sheaths often

mottled purple; lemma often smoky white at maturity with a darker area

covering the caryopsis; marginal hairs of lemma often strongly divergent

RETA TNS cee ree eo in lg Meee ge aie ay oN metals EB, foe Syd Ose L. fusca subsp. muelleri

Lowermost panicle branches exserted at maturity; uppermost leaf blade

exceeding length of panicle; lower portion of leaf sheaths usually not

302 Austrobaileya 5(2): 299-305 (1999)

mottled purple; lemma colour various, but generally not smoky white

with a darker area; marginal hairs of lemma not strongly divergent at

PIPATIBIES 9 se ee a) ce Rs aD AL ak FF, HS TO Dav gk oe BADAE BLA GAP aBee |i eet BY ea PA a Teng CK BARA OO 7

7. Lemma apex obtuse to truncate, often notched and mucronate; lemma

dark green or lead coloured; panicles generally completely exserted

from sheaths, narrowly elliptic to elliptic in profile; panicle branches

held at greater than 45° angle, often greater than 30 branches in number;

anthers usually less than 0.5 mm long ................. L. fusea subsp. uninervia*

Lemma apex usually acute or acummate, notched or not, mucronate or

not; lemma generally not dark green or lead coloured; panicles

sometimes not completely exserted, often broader than elliptic in

profile; panicle branches sometimes held at less than 45° angle,

frequently fewer than 30 branches in number; anthers usually 0.5—2.5

LUAWD diye (3) 3 Sk ee ee | ROR 4 ee? PO eS te ae ee L. fusca subsp, fusca

8. Panicle apex erect; mature panicle branches (10-) 15-30 cm long and

divergent or reflexed; spikelets mostly distant................. L. divaricatissima

Panicle apex usually somewhat nodding; mature panicie branches mostly less

than 15 cm long, the branches only rarely divergent or reflexed; spikelets

seneraily sOmew hat-OVEtlan Pe avo neck pee ce dcbteode rah tdonige tose eiRrencens ate auace aneeRR wletal 9

9. Leaf sheaths with tubercule-based pilose hairs .. 0.0... 00 ccc ee ee eens 10

Leaf sheaths lacking turbercule-based pilose hairs ......... 0.000: cece eee eee 14

10. Leaf blades covered with sericeous or tomentose hairs; lower half of lateral

nerves of lemma densely sericeous, the hairs increasingly divergent at

Leaf blades glabrous; lower half of lateral nerves of lemma hairy or not, but

not with witely diFereet TaIES tyke Wee es bey ke an i ENN TY 1]

11, Plants over 100 cm tall; hairs on leaf sheaths more or less dense and occurring

throughout sheath; ligule apex sometimes notched at the middle .... L. southwoodii

Plants Gn Australia) mostly under 100 cm tall; hairs on leaf sheaths erratic,

often most dense near sheath apex; ligule apex not clearly notched at the )

(919 (6 (6 (see LA ewe As eae ROSE UG, 1 OO Oe OR RIOT LW ae PPE eee ee 12

12. Culms wiry; panicle branches generally naked along lowest 2 mm; leaf blades

— generally lacking a distinct midvein on upper surface (or, ifso, only occurring

at very base) ........... L. decipiens subsp. asthenes (formerly known as L. ciliolata)

Culms not wiry; panicle branches bearing one or more spikelets along lowest

2 mm; leaf blades with a distinct midvein on upper surface ............ 0.0000 ee 13

13. Caryopsis with a shallow groove and somewhat laterally compressed, the apex

often broadly acute; lower lemma mostly more than 1.3 mm long, mostly

glabrous between midnerve and lateral nerves............. 000000

ota Aneel ae a L. panicea subsp. brachiata* (formerly known as L. mucronata)

Caryopsis lacking shallow groove and mostly terete in cross section, the

Snow & Simon, Taxonomic status Leptochloa fusca subsp. uninervia 303

apex obtuse but.never acute; lower lemma mostly less than 1.2 mm

long, glabrous or hairy between midnerve and lateral nerves

(ot Perciatapetig SiAy Wa L. panicea subsp. panicea*

14, Plants perennial; ligules 0.9-1.7 mm long, apex erose but not notched near

Npmacat tema tnt L. decipiens subsp. decipiens

Plants annual; ligules 4.2—-7 mm long, sometimes with a central notch at the

ADEM. oie g etal e ley My ois tote wed Su PRU Sieyetale N34

Acknowledgments

The first author thanks Washington University (St.

Louis) and the Missouri Botanical Garden

(Andrew W. Mellon Foundation) for their

support. A generous grant from the National

Geographic Society (NGS 5594-95) provided

funding to collect in Africa and Australia. Thanks

to Dr. Surrey Jacobs for comments on the

manuscript, and to P, Green for details regarding

specimen localities.

References

BEETLE, A.A.,, MANRIQUE F,, E., MIRANDAS., JL.A., V JARAMILLO

L., V., CHIMAL H., A. & Ropricuez R., A. M. 1991.Zas

Gramineas de Mexico, Yomo Ill. COTECOCA, S.A.R.H.

Manzanillo, Mexico, D. F.

Corre t, D.S. & JoHNsTON, M. C. 1970, Manual of the Vascular

Plants of Texas. Texas Research Foundation, Renner,

Texas.

GIBBS RUSSELL, G. E., WATSON, L., KOEKEMOER, M., SMOOK, L.,

BARKER, N.P., ANDERSON, H.M. & Datiwirz, M.J. 1991.

Grasses of Southern Africa. Memoirs of the Botanical

Survey of South Africa No. 58.

Goutp, F.W. 1975. The Grasses of Texas. Texas A & M

University Press, College Station.

HAF ickr, E. & Scuouz, H. 1981.Grass Weeds 2: Weeds of the

Subfamilies Chloridoideae, Pooideae, Oryzoideae.

CIBA-GEIGY, Ltd., Basle, Switzerland.

JAcops, S. W. L. & McCray, K.L. 1993. Leptochioa and

Diplachne, In: Flora of New South Wales, Volame 4. G.

J. Harden (ed.). New South Wales University Press.

LAZARIDES, M. 1980. The genus Leptochloa Beauv.

(Poaceae, zur Grasflora (Poaceae) der Kanaren.

Willdenowita 25: 571-582.

Mason, H.L. 1957. A flora of the Marshes of California.

University of California Press, Berkeley.

Pets alee Mienea eke aeoek L. ligulata

McIntyre, 8. 1985. Seed reserves in temperate

Australian rice fields following pasture rotation

and continuous cropping, Journal of Applied

Ecology 22:875-884,

McInrvre, S., Lapicgs, P.Y. & ApAms, G. 1988. Plant

species-richness and invasion by exotics in

relation to disturbance of wetland comunities

on the Riverine Plain, NSW. Australian Journal

of Ecology 13:361-373.

McIntyre, 8., MircHeLL, D.S. & Lapicss, P.Y. 1989.

Germination and seedling emergence in

Diplachne fusca: a semi-aquatic weed of rice

fields. Journal of Applied Ecology 26:55 1-562.

McVauaeu, R. 1983. Flora Novo-Galiciana: A

Descriptive Account of the Vascular Plants of

Western Mexico, W.R. Anderson (editor). Vol.

14. Gramineae. The University of Michigan

Press, Ann Arbor.

Nicora, E.G, 1995. Los géneros Diplachne y

Leptochioa (Gramineae, Eragrosteae) de la

Argentina y paises limitrofes. Darwiniana

33:233-256. |

Puittips, 8,M. 1974. 67. Diplachne,. In: Flora of

Tropical East Africa. Gramineae (Patt 2), R.M.

Poihiil, Editor. Crown Agents for Oversea

Governments and Administrations, by

Whitefriars Press, Ltd., London.

ScHOLZ, H, & Bocxer, R. 1996. Ergénzungen und

Anmerkungen zur Grasflora (Poaceae) der

Kanaren, Wilidenowia 25:571-582.,

SIMON, B.K. 1993. A Key to the Australian Grasses.

Second Edition, Department of Primary

Industries, Brisbane.

Snow, N. 1996, The phylogenetic utility of lemmatal

micromorphology in Leptoch/oas.|. and related _

genera in subtribe Eleusininae (Poaceae,

Chloridoideae, Eragrostideae). Annals of the

Missouri Botanical Garden 83: 504-529.

304

—— 1997a. Phylogeny and Systematics of Leptochloa

P, Beauv. sensu lato (Poaceae, Chloridoideae).

Ph.D. dissertation, Washington University, St.

Louis, Missouri,

—- 1997b. Application of the phylogenetic species

concept: a botanical monographic perspective.

Austrobaileya 5: 1-8.

-— 1998. Nomenciatural changes in Lepfochioa P.

Beauv. sensu lato (Poaceae, Chloridoideae).

Novon 8: 77-80.

Snow, N., & DavipsE, G, 1998. Proposal to reject the

name Poa malabarica. Taxon 47; 157-159.

Snow, N., & Simon, B.K. 1997. Leptochloa southwoodii

(Poaceae: Chloridoideae), a new species from

south-east Queensland. Austrobaileya 5:137-

143.

STANLEY, T.D. & Ross, E.M. 1989. Flora of South-eastern

Queensland, Volume IH, Queensland Department

of Primary Industries Miscellaneous Publication

OM88001.

Austrobaileya 5(2): 299-305 (1999)

305

Snow & Simon, Taxonomic status Leptochloa fusca subsp. uninervia

showing palea and the

somewhat truncated lemma apex, the marginal nerves pubescent below; d, habit, e, grain, f£ leaf sheath and the ligule

(shown as bilobed; typically it is attenuate in undamaged condition). Reproduced with permission of the University of

California Press from Mason (1957)

floret,

™

a a ae a al

~ a

Fig. 1. Leptochioa fusca subsp. uninervia (J. Presl) N.Snow: a, spikelets; 6 and c,

Pee eee ee eee eee eee eee eee ees Seeeee eee Tete ee ge ee tT eee etn eee eee eet tent net dt ne Te tut fetal Sanne fe tat SSE OSEOEEEE eee ate eee fe ee ee atte fate oe att toes

SaGGh Gent behas an Aaa Saas ATS RAS eRR SAAS Enh Eh iG bGneTe eet ne ganic Een Raq eE tay tthe RGR ETRE eS

Notes on Acacia (Leguminosae: Mimosoideae) chiefly from

northern Australia

Les Pedley

Summary

Pedley, Les (1999), Notes on Acacia (Leguminosae: Mimosoideae) chiefly from northern Australia,

Austrobaileya 5(2): 307-321. Species are referred to sections recognised by Pedley (1978), Acacia

Abbatiana, A.arbiana, A. barakulensis, A. hendersonii, A. rubricola (sect. Phyllodineae), A.

argyrotricha, A. johannis, A. convallium (sect. Plurinerves), A. abbatiana, A. burdekensis, A. faucium,

A. filipes, A. fodinalis, A, lacertensis, A. proiantha, A, scopularum and A. solenota (sect. Juliflorae)

are described as new. A. tingoorensis nom, et stat. nov. is based on A. longispicata subsp. velutina

Pedley. Notes on affinities, distributions and habitats of all species are given. Variation within A.

leiocalyx (Domin) Pedley is discussed and a wide area of intergrade between A. fodinalis and A.

—_——————— Oo ee ee a a i ee ee ee ee SS eee tna tn lads nda in inthe dleain Se Panel senes Nec asin beats pen Eiht hes bt dies in ich eine en it

cretata Pedley is postulated.

Keywords: Acacia — Australia; Queensland; Northern Territory.

Les Pedley, c/- Queensland Herbartum Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,

Toowong, Queensland 4066, Australia.

Introduction

With the approaching treatment of Acacia in

Volumes LIA & 11B of the Flora of Australia

it has become necessary to validate names of

some species from Queensland and the

Northern Territory included there, and some

others that will not be included. Several of the

species are closely related to A. leiocalyx

(Domin) Pedley and variation within that

species is discussed, The species are referred

to sections according to the classification of

Pedley (1978). Other species described here

were recognised as distinct after editing of the

two volumes of the Flora began and therefore

cannot be included in it. These species are

indicated by an asterisk (*). In the Flora, for

the most part, they are included in rather wide

circumscriptions of other species.

Acacia sect. Phyllodineae DC., Prodr. 2: 448

(1825).

Acacia arbiana Pedley, sp. nov. ex affinitate

A.confertae A. Cunn. ex Benth. et

specierum affinium distinguenda

phyllodiis distincte mucronulatis,

angustioribus, elongatioribus, floribus

aliquantum grandioribus corollae

lobis longioribus praeditis. Typus:

Queensland. LeicHHARDT District: Ropers

Accepted for publication 3 September 1998

Peak, 22°52’S 148°13’E, alt. 780 m, 23

August 1990, PI. Forster PIF7209 (holo:

BRI iso: AD, BRI, CANB, DNA, K, MEL,

MO, NSW, PERTH, PRE, Z).

Acacia sp. (Ropers Peak P.I. Forster

PIF7209) (Pedley 1997),

Spreading shrub to 1.5 m tall with black stems;

branchlets, hidden by phyllodes, glabrous or

with scattered loosely appressed hairs, ribbed

by decurrent bases of phyllodes., Phyllodes

linear, straight or slightly curved, narrowed into

a longish curved mucro, crowded, spirally

arranged, sometimes in pseudo-whorls, 8-16

mm long, 0.6—-0.8 mm wide, thick without

obvious nerves, narrowed into a longish curved

mucro, with sparse, loose, + appressed hairs,

c. 0.6 mm long, particularly on abaxial margin

when young, glabrescent, apparently

exstipulate but with minute red-brown

trichomes in stipular position; pulvinus c. 0.5

mm long. Heads of 24-30 flowers on single

axillary peduncles 8-10 mm long, projecting

beyond the foliage, peduncle occasionally with

| or 2 bracts below the head. Flowers 5-merous,

subtended by bracteoles as long as the calyx

with thick oblique pubescent lamina as long as

the claw; calyx c. 1 mm long, lobes about one-

third of total length, ciliate, often with a stouter

brown trichome at the tip; petals obovate, c. 2

308

mm long, united to about the middle, glabrous;

stamens c. 3 mm long; ovary glabrous. Pods

similar to those of A. conferta, up to 4.5 cm

long, 10-14 mm wide, valves rather

membranous, finely reticulately veined,

somewhat glaucous, convex over the seeds;

seeds transverse, mature ones not seen.

Specimens (all BRI): Queensland, LEICHHARDT DISTRICT:

Ropers Peak, May 1987, Bean 569 & Aug 1987, Bean

630; Scotts Peak, 22°51’S 148°13’E, alt. 570 m, Jun

1951, Everist 4427; Peak Range, c. 30 km SW of Dysart,

Jan 1981, Podlich s.n. (AQ 348047),

Distribution and habitat: Vhe species is

confined to Ropers and Scotts Peak and perhaps

other peaks of the Peak Range, east of

Clermont. It is recorded from trachyte outcrops

where it is a component of heath-like vegetation

with Phebalium glandulosum Hook., Corymbia

frachyphloia (F.Muell.) K.D. Hill & L.A.S.

Johnson, Bertva pedicellata F.Muell., Acacia

gnidium Benth., Dodonaea filifolia Hook.,

Callistemon sp. and Zieria aspalathoides A.

Cunn. ex Benth.

Affinities: Acacia arbiana is close to

A. conferta, but its phyllodes are more crowded,

longer, narrower and distinctly mucronulate with

long loosely appressed hairs, and it also lacks

stipules. Its phyllodes resemble those of A.

gittinsii which has spreading hairs on the stems,

some heads in racemes and longitudinal seeds.

Etymology: The specific epithet is derived

from the initial letters of the names of Mr A.R.

(Tony) Bean whose flowering specimens

collected in 1987 clearly indicated the species

to be undescribed.

* Acacia barakulensis Pedley, sp. nov. affinis

A. burbidgeae Pedley a qua ramulosorum

pilis aliquantum — sparsioribus

brevioribusque, phyllodiis brevioribus

apice mucrone obliquiore pulvino 0.5—1

mm longo, plerumque brevioribus nervis

marginalibus minus prominentibus

ornatis, capitulis in pedunculis

longioribus differt. Typus: Queensland.

DARLING Downs District: Barakula State

Forest, 26°26’S 150°31’E, 18 August

1971, L.A.Nielsen 15 (holo: BRI).

Acacia barakulensis Pedley ex Lithgow, 60

Wattles of the Chinchilla and Murilla

Austrobaileya 5(2): 307-321 (1999)

Shires: 46 t.32 (1997), nom. invalid., no

Latin diagnosis or description.

Acacia sp. (Barakula L.A. Nielsen 15)

(Pedley 1997).

Shrub to c. 2 m high; branchlets brown,

resinous, ribbed, sparsely pubescent; stipules

setaceous, 0.3—0.6 mm long, persistent.

Phyllodes sometimes in pseuowhorls, borne on

short projections from ribs of branchlets, terete

or somewhat laterally compressed, linear, +

straight, 10-22(—28) mm long, 0.6—1 mm thick,

somewhat irregularly tuberculate with a few

hairs, an inconspicuous yellowish nerve on ab-

and adaxial surfaces with (when dry), a distinct

lateral furrow between them and usually some

obscure longitudinal folds; mucro oblique

(sometimes perpendicular), 0.1—0.3 mm long;

pulvinus 0.5—1 mm long; a single minute gland

behind the mucro. Heads c. 9 mm diam. of (20-—

)25—35 flowers on ebracteate peduncles 6-10

mm long, single in the upper axils. Flowers 5-

merous; calyx 0.7—0.8 mm long with wide

obtuse lobes 0.2—0.3 mm long, a few short hairs

at their tips; corolla 1.7—1.8 mm long, lobed to

the middle, glabrous; stamens c. 4 mm long;

ovary glabrous. Pods linear, + straight, slightly

contacted between the seeds and slightly

convex over them, to c. 40 mm long, 4 mm

wide, valves rather chartaceous, brown,

resinous, reticulately nerved, marginal nerves

prominent. Seeds longitudinal, 3.7-4.2 mm

long, 2~2.4 mm wide, brown; areole darker,

large; pleurogram constricted but open; funicle

filiform, creamy yellow, thickened into a

clavate aril. Waajie Wattle.

Selected specinens (all BRI): Queensland. DARLING

Downs Districr: 16-24 km N of Miles, Jul 1980, Hando

143; Woojie [sic] flower area, Panda Lane, c. 30 miles

[48 km] NW of Barakula Forestry Station, Aug 1981

(flowers) & Nov 1981 (pods), Lithgow 913; S.F.R. 16

Maicolm, N of Chinchilla, in 1958, Cameron QFDS9/84.

Distribution and habitat: The species occurs

only in the Barakula State Forest north of

Chinchilla on sandy soils in eucalypt

communities. It flowers August-September and

pods have been collected in November.

Affinities: The nearest relative of A.

barakulensis is A. burbidgeae but it is

distinguished by its shorter phyllodes with

Pedley, L., Acacia (Leguminosae: Mimosoideae) chiefly from northern Australia

usually more oblique mucro and its heads on

longer peduncles.

Etymology: The specific epithet is derived

from Barakula the only known locality of

occurrence of the species.

*Acacia hendersonii Pediey, sp. nov. ab 4.

jJohnsonii Pedley ramulis glabris

phyllodiis juventate midnervo

translucente ornatis in pulvinum brevem

abrupte contractis mucrone minori

praeditis, capitulis in pedunculis

longioribus portatis differt; ab A.

resinicostata Pedley phyllodtis pulvino

brevi praeditis (non sessilibus)

tenuioribus longioribusque differt.

Typus: Queensland. LEICHHARDT

District: Blackdown Tableland c. 32 km

SE of Blackwater, 23°50’S 149°0S’E, alt.

600 m, 11 September 1971, RJ.

Henderson, L. Durrington & PR. Sharpe

H1199 (holo: BRI).

Acacia sp. (Blackdown Tableland R.J.

Henderson+ H1199) (Pedley 1997).

Spreading, much-branched shrub to 3 m tall;

branchlets slender, resinous, glabrous,

prominently ribbed, the ribs tuberculate;

stipules subulate, 0.8—1 mm long, persistent.

Phyllodes flat but thick, linear, straight or

slightly decurved at apex or distinctly sigmoid,

(6—)12—22 mm long, 0.7—1.1 mm wide, (8—)15—

22 times longer than wide, glabrous, one vein

prominent on each face, yellow and translucent

on young (dark) phyllodes, raised and opaque

on older ones, margins thickened, a distinct

narrow longitudinal depression between midrib

and margins on dried phyllodes; a short oblique

mucro at apex; abruptly narrowed into pulvinus

0.2—-0.5 mm long; a small gland up to 1 mm

from base and a second smaller one at apex

behind mucro. Heads of 30-35 flowers on

ebracteate peduncles 9-15 mm long, single in

upper axils; bracteoles spathulate, c. 1.2 mm

long. Flowers 5-merous; calyx glabrous, c. 1.2

mm long, lobed to c. '/4 length; corolla glabrous,

1.72.2 mm long, lobed to middle; stamens c.

3 mm long; ovary glabrous or minutely

papillose towards top. Pod not seen.

Other specimens (all BRI): Queensland, LEICHHARDT

District: Blackdown Tableland, Aug 1966, Gittins 1184;

ditto, Aug 1973, Trapnell & Williams 40; ditto, Sep 1971,

309

Henderson et al. H1077; Blackdown Tableland, 23°48’S

149°O08’E, Sep 1988, J.G. Simmons & M._H.Simmons

2058.

Distribution and habitat: The species is

confined to Blackdown Tableland, an isolated

sandstone plateau with an interesting flora,

including the endemic, or near-endemic,

species Acacia gittinsii Pedley, A. storyi

Tindale, Corymbia bunites (Brooker &

A.R.Bean) K.D. Hill & L.A.S. Johnson,

Eucalyptus mensalis L.A.S. Johnson & K.D.

Hill and £. sphaerocarpa L.A.S. Johnson &

Blaxell.

Affinities: Acacia hendersonii is one of a group

of closely interrelated species that includes A.

barakulensis, A. burbidgeae Pedley, A.

Johnsonii, A. pilligaensis Maiden, A.

resinicostata and A. rubricola Pedley. It most

closely resembles A. johmsonii but differs in

having glabrous branchlets, phyllodes with

translucent midribs when young, abruptly

contracted into a short pulvinus, with a smaller

mucro and heads borne on longer peduncles. It

also has some affinity with A. resinicostata but

differs in its finer and longer phyllodes which

are not sessile but have pulvinuses 0.20.5 mm

long.

Etymology: The specific epithet honours my

friend and colleague Mr R.J.F. Henderson who

not only was joint collector of the type on the

first organised expedition to collect plants of

the Blackdown Tableland, but also a noted

systematist specialising in Liliaceae (sensu

latissimo) (for example, Henderson 1987).

* Acacia rubricola Pedley, sp. nov. affinis A.

johnsonii Pedley a qua phyllodiis vix

pubescentibus minus crassis plerumque

longioribus midnervo translucenti flavido

ornatis, ramulorum pilis aliquantum

brevioribus, leguminibus saepe

angustioribus differt. Typus: Queensland.

BuRNETT District: 6.7 km along

Gurgeena Plateau road, 25°29’S

151°23’E, 23 August 1989, P/.Forster

PIF5650 (holo:BRI; iso (n.v.): CANB,

MEL, PERTH).

Heavily foliaged, much branched shrub to 2

m high; branchlets ribbed, with phyllodes

borne on projections of ribs, resinous,

310

hispidulous with stiff straight hairs 0.1—0.2

mm long; stipules narrowly triangular, c. 0.5

mm long. Phyllodes flat, linear, straight or

slightly sigmoid, narrowed from about the

middle to base, 20-43 mm long, 1—1.8(-2.1)

mm wide, 13-30 times longer than wide, a

single translucent yellow longitudinal nerve

prominent on each face, with additional

obscure longitudinal folds when dry,

marginal nerves irregularly sparsely

tuberculate, covered with translucent resin

and with a few hairs at and near base when

young; gland at base small with thick

yellowish rim, a second smaller gland behind

mucro; mucro c. 0.5 mm long; pulvinus c. |

mm long. Heads c. 9 mm diam., of 20-35

flowers on single yellow resinous peduncles

6—8 mm long in the upper axils; peduncle

without basal bract but occasionally a bract

or flower at about middle of peduncle;

bracteoles spathulate, incurved, c. 0.8 mm

long. Flowers 5-merous; calyx 0.7—0.9 mm

long, divided to about the middle into wide,

obtuse lobes, glabrous except for a few minute

trichomes at apex of lobes; corolla 1.5—1.7

mm long glabrous; stamens c. 4 mm long;

ovary glabrous or with a few hairs at base of

style. Pods linear, dark to c. 40 mm long, 2.5

mim wide, dark brown; valves rather

chartaceous, raised over seeds. Seeds 3.7—

4.5 mm long, 1.6—2 mm wide, longitudinal;

areole large, pleurogram open; funicle fine,

folded and thickened into cream-coloured

clavate aril.

Other specimens (all BRI): Queensland. BURNETT

District: Binjour Plateau, near Gayndah, 25°28’S

151°23’°E, Oct 1987, Bean 673; Mundauran Pocket,

Gurgeena Plateau, 25°28’S 151°23’E, Jan 1990, Forster

PIF6174: State Forest Reserve 130, 2 km NW of

Nantglyn, 25°31’S 151°21’E, Sep 1989, Forster PIFS734

& Bean; Gurgeena Quarry, Aug 1988, 1.G.Simmonds &

MH Simmonds 2017.

Distribution and habitat: The species is

restricted to the Binjour Plateau near

Gayndah where it occurs on red loamy soil

in eucalypt open-forest or in heath. It flowers

from mid-August to September and a fruiting

specimen has been collected in January.

Affinities: Acacia rubricola is closely related

to A. johnsonii and duplicates have been

distributed under that name. It differs

Austrobaileya 5(2): 307-321 (1999)

however in having longer, less pubescent,

somewhat resinous phyllodes with quite

conspicuous yellow translucent midnerves

and branchlets with shorter hairs, and

narrower pods, though I have seen few

mature pods of either A. johnsonii or A.

rubricola.,

Etymology: The specific epithet is derived

from Latin rubrica, red earth, and -cola,

inhabitant, an allusion to the occurrence of

the species on red soils.

Acacia sect. Plurinerves (Benth.) Maiden

& Betche, Census Pl. NSW 93 (1916)

Acacia argyrotricha Pedley, sp. nov. affinis

A. rigenti A. Cunn. ex G. Don, a qua

ramulis angularibus non costatis,

phyllodiis saepe latioribus, capitulis

florum amplioribus tn pedunculis

parum longioribus portatis,

leguminibus plerumque angustioribus

et imprimis indumento pilorum 0.5 mm

longorum in ramulis, phyllodiis

juvenibus (remanenti in frustillis in

phyllodiis veteribus), pedunculis

leguminibusque differt. Typus:

Queensland. DARLING Downs DISTRICT:

Bracker State Forest, S of Inglewood,

28°36’S 151°02’E, 16 September 1989,

A.R. Bean 1115 (holo: BRI).

Acacia sp. (Inglewood A.R. Bean 1115)

(Pedley 1997).

Shrub to c. 2 m tall, much branched from the

base; bark smooth, grey to reddish brown;

branchlets angular with dense appressed

brown hairs, becoming grey with age, 0.5 mm

long; young foliage brownish. Phyllodes

linear, flattened, straight or slightly falcate,

rather thick, 10-15 cm long, I1—1.8 mm wide,

with indumentum of appressed brown hairs

when young, glabrescent or hairs becoming

white and retained in patches or along

midline; mucronulate with usually curved

brown mucro 1—2 mm long; gland at base of

phyliode; pulvinus stout, c. 1 mm long,

usually with appressed indumentum of stems.

Heads slightly elongate, deep yellow, of 30—

40 flowers in pairs in the upper axils; peduncles

2—2.5 mm long, pubescent, a short stout axis

"Lp SEE

Pedley, L., Acacia (Leguminosae: Mimosoideae) chiefly from northern Australia

between them. Flowers 5-merous, each

subtended by a concave curved bracteole c. 1.3

mm long; calyx rather stout, obconic c. 1 mm

long, lobed to about the middle, the tube with

dense hyaline hairs particularly in the upper

half, lobes ciliolate; corolla glabrous, c. 1.5 mm

long, c.'4 lobed, midrib in lower parts of lobes

only; stamens 3—4 mm long; ovary glabrous.

Pods linear, to 7.5 cm long, c. 2.5 mm wide,

somewhat apiculate, with c. 8 seeds; valves

coriaceous, brown, raised over the seeds,

sparingly veined, with patchy indumentum of

silvery appressed hairs c. 0.5 mm long. Seeds

longitudinal, oblongoid, 3-4 mm long, 1.7—2

mim across; pleurogram + distinct, open; areole

narrowly oblong; funicle folded and thickened

into cupular aril.

Specimens (all BRI, all from type locality): Apr 1989,

Bean 1023 & Ballingall 2542; Aug 1990, Ballingall

2579; Dec 1990, Bean 2735; s.d., Ballingall 2683.

Distribution and habitat: The species 1s known

only from the type locality where it occurs on

sandy soil in eucalypt woodland.

Affinities: Acacia argyrotricha resembles A.

rigens but differs notably in the appressed hairs

of the branchlets and young phyllodes; the

indumentum is often retained in patches on and

along the mid-line of old phyllodes. Its flower-

heads are larger and borne on longer peduncles.

Etymology: The epithet 1s fronr Greek,

argyros, silvery, and trichos, hair, a reference

to the silvery indumentum particularly of the

pods.

*Acacia johannis Pedley, sp. nov. ex affinitate

A. armillatae (Pedley) Pedley, A.

legnotae Pedley, A. ommatospermae

(Pedley) Pedley; a tribus omnibus

phyllodiis elongatioribus, pedunculis

filiformibus, leguminibus angustioribus,

ab A. armillata phyllodiis angustioribus

nervis prominentibus longitudinalibus

paucioribus pedunculis longioribus,

leguminium valvis super semina elevatis,

seminibus parvioribus, ab A. /egnota

phyllodiis elongatioribus angustioribus

nervis longitudinalibus paucioribus

leguminium valvis super semina elevatis,

ab A. ommatosperma phyllodiis apice

non latissimis, pedunculis longioribus,

311

pleurogrammate non prominenti differt.

Typus: Queensland. Cook District: Mt

Mulligan, 16°54’S 144°S1E, alt. 740 m,

13 April 1990, /.R. Clarkson 8217 (holo:

BRI; iso: K, MBA, MEL, MO, QRS).

Acacia sp. (Mt Mulligan J.R. Clarkson 8217)

(Pedley 1997).

Shrub to 2 m tall; bark smooth, grey; branchlets

glabrous, reddish brown, ribbed, lenticellate.

Phyllodes glabrous, narrowly elliptic, tapering

sradually to base and apex, markedly faicate,

14-18 cm long, 5-10 mm wide, 15-30 times

as long as wide; glabrous; 3 or occasionally 4

longitudinal nerves prominent, secondary

nerves forming open anastomoses between

them; acute with a callus point; gland +

prominent on margin at base; pulvinus 1.5-2

mm long. Heads ofc. 50 flowers, lemon-yellow,

on glabrous filiform peduncles 14—20 mm

long, arising from a brachyblastic axis c. 1

mm long at anthesis; axes usually two to

each axil occasionally growing out into leafy

shoots; bracteoles with a thick peltate shortly

pubescent lamina almost perpendicular to

the claw, about as long as the calyx. Flowers

5-merous; calyx divided almost to the base

into narrow ciliolate spathulate lobes c. 1.5 mm

long, the apex thickened; corolla deeply divided

into ovate lobes, 1.8—2 mm long, glabrous;

stamens 3—4 mm long; ovary glabrous. Pods

shortly stipitate, linear, straight, dehiscent,

the valves chartaceous, constricted between

the seeds and raised over them, alternately

on each valve, 7-10 cm long, 4 mm wide at

widest part, 2-3 mm at isthmuses, glabrous.

Seeds (up to 11 per pod), longitudinally

arranged, dark brown, depressed ellipsoidal,

c, 3.5 mm x 2.5 mm, pleurogram obscure,

closed; funicle thickened forming a small

clavate aril.

Specimens: Queensland. Cook Districr: Mt Pinnacle,

SSW of Dimbulah, 17°14’S 145°03’E, Jan 1993, Bean

5578 & Forster (BRI); 3.3 km, 8 of the crossing of

Shepherd Creek on Maytown track, 15°48’S 144°16’E,

Jun 1992, Clarkson 9615 & Neldner (BRI, MBA); Mt

Mulligan, 16°52’S 144°S1°E, alt. 700 m, Apr 1985,

Clarkson 5805 (BRI, K, MBA, MEL, NSW, PERTH) &

Apr 1987, Clarkson 6927 (BRI, K, MBA, MEL, NSW,

PERTH, QRS).

Distribution and habitat: The species is

restricted to north-eastern Queensland where

312

itis common on Mt Mulligan. It occurs on rock

outcrops and pavements and in shallow rocky

soils derived from sandstone, and is common

in places, forming thickets.

Affinities: Acacia johannis belongs to what has

been termed the Oligoneura group of

Racosperma (Pedley 1987), equivalent to the

Oligoneurae group of Acacia (Pedley 1978).

It is probably most closely related to Acacia

armillata, A. legnota and A. ommatosperma.

It differs from.A. armillata in its more elongate

phyllodes with fewer prominent longitudinal

nerves, longer peduncles and narrower pods

raised over the somewhat smaller seeds; from

A. legnota in its narrower, more elongate

phyllodes with fewer longitudinal nerves,

filiform peduncles and narrower pods raised

over the seeds; and from A. ommatosperma in

its less elongate phyllodes not widest near the

tip, shorter peduncles and narrower pods.

Etymology: The epithet is derived from the

Latin, Johannes, John. The species is named

in honour of John R. Clarkson (MBA) whose

work in far northern Queensland in the last 15

years has significantly increased scientific

knowledge of plant species and communities

of the region.

Acacia convallium Pedley, sp. nov. affinis

A. sericatae Cunn. ex Benth. et A.

platycarpae F. Muell.; a illa non denso

indumento pilorum brevium manifeste

stellatorum in phyllodiis ramulis

pedunculisque habenti; a hac areolis

nervorum anastomantium phyllodiorum

majoribus elongatioribusque, plerumque

ramulis pubescentibus non glaucis,

pluribus capitulis florum in quoque

fasciculo, ab utraque phyllodiis non

latissimis supra medium aliquantum

acutis differt. Typus: Northern Territory.

East Alligator River, 12°47’S 133°23’E,

18 July 1972, N.B. Byrnes 2750 (holo:

BRI; iso (n. v.): CANB, DNA, K, NSW).

Spindly shrub to 5 m tall; branchlets terete

with sparse to dense felty indumentum of

hairs to 0.2 mm long, sometimes becoming

glabrous, or rarely glabrous; stipules deltoid

c. | mm long. Phyllodes ovate, falcate, 9—

17.5 cm long, 1.2—4 cm wide, 3.5—-8.5 times

Austrobaileya 5(2): 307-321 (1999)

longer than wide; with sparse indumentum

of weak spreading hairs c. 0.15 mm long,

hairs sometimes confined to the base; upper

margin often undulate, with two, three or

rarely four longitudinal nerves prominent,

only one reaching the apex, all running

together into the lower margin at the base,

oblique secondary nerves forming a

reticulum; tapering to a callus point; glands

2~4(—5) on upper margin, small, sometimes

projecting, absent from the base; pulvinus 5—

10 mm long, with indumentum of branchlets.

Inflorescence of up to 7 fascicles of

pedunculate heads at anthesis at the one time,

on indeterminate axes in the upper axils, the

axis up to 10 cm long, with indumentum of

branchlets, with immature fascicles crowded

at apex; fascicles subtended by a deciduous

deltoid bract c. 1 mm long, of 4-6 slender

peduncles 12—17(—23) mm long, with

indumentum of spreading hairs c. 0.25 mm

long, the hairs sometimes confined to base.

Heads of c. 30 flowers each subtended by a

clawed bracteole, the claw as long as the

calyx, the lamina c. 0.5 mm long, obliquely

ovate, acute. Flowers 5-merous; calyx lobes

free, spathulate, c. 1 mm long, with spreading

hairs in upper half; corolla lobed to about

the middle, c. 1.6 mm long, the lobes with

spreading hairs; stamens c. 3.5 mm long, the

filaments cohering in the lower half

(especially in male flowers); ovary glabrous.

Pods 7-11 cm long, 28-36 mm wide, with

woody, glabrous, reticulately nerved valves,

the margins forming a ‘wing’ 2.5—3.5 mm

wide. Seeds 8—8.5 mm long, 5.3-6.2 mm

wide, 3—3.5 mm thick, transverse, arranged

in depressions in the material of the valves;

areole large; pleurogram open, well defined;

funicle c. 3 times folded and thickened into

a stout aril forming a cap over the seed.

Specimens: Northern Territory. East Alligator River

area, 12°41’°S 133°08’E, Dunlop 3421 (BRI, DNA, K);

44 km SE of Oenpelli, 12°34’S 133°23’E, Dunlop 4926

(BRI, DNA); 14 km NE of Mann River Gorge, 12°39’S

134°08’°E, Leach & Dunlop 1590 (BRI; Along road

to Smith Point, 73.5 km NW of Murgenella, 11°20’S

132°20’E, McDonald 421 (BRI).

Distribution and habitat: The species is

confined to a small area in the north of the

Northern Territory where it usually occurs on

sandstone, often in gorges.

Pediey, L., Acacia (Leguminosae: Mimosoideae) chiefly from northern Australia

Affinities: Acacia convallium 1s allied to A.

sericata but lacks the stellate hairs so prominent

on the phyllodes, branchlets and peduncles of

that species, and more closely to A. platycarpa

but the areoles among the anastomoses of the

nerves of its phyllodes are larger and more

elongate; its branchlets are usually pubescent,

not pruinose; and it has more flower heads in

each fascicle in the inflorescence. It differs from

both in having somewhat acute phyllodes not

widest above the middle. The cohesion of the

lower part of the staminal filaments is most

unusual in species of Acacia.

Etymology: The specific epithet is genitive

plural of Latin convallis, a valley shut in on all

sides, an allusion to the plant’s habitat.

Acacia. sect. Juliflorae (Benth.) Maiden &

Betche, Census Pl. NSW 95 (1916)

“Acacia abbatiana Pedley, sp. nov. A.

graniticae Maiden similis a qua cortice

lamellato, phyllodiis aliquantum

brevioribus, inflorescentiis multispicatis,

floribus parvioribus (staminibus tantum

c, 2.2 m longis) hinc spicis tenuioribus,

calyce pilis arachnoideis obtecto,

leguminibus brevioribus, seminibus

parvioribus pleurogrammate U-formi

ornatis differt. Typus: Queensland.

SOUTH IKENNEDY District: Mt Abbot, 50

km W of Bowen, 20°06’S 147°46’E, alt.

800 m, 2 August 1992, 4A.R Bean 4873

(holo: BRI; iso (n.v.): AD, BISH, HO,

K, L, MEL, MO, NSW).

Acacia sp., (Mt Abbot A.R.Bean 4873)

(Pedley 1997).

Shrub to c, 4 m tall, foliage confined to upper

part of plant, lower part of stems bare; bark

brown, lamellated (described by collector as

‘fibrous’); branchlets pale brown, often with

greyish overlay of dead epidermal! cells,

glabrous, angular. Phyllodes linear, straight,

(70—)95—150 mm long, 1.6~3 mm wide, 40—

65 times longer than wide, glabrous, with 13-—

19 parallel non-anastomosing longitudinal

nerves, the midnerve slightly more prominent,

marginal nerves somewhat thicker than rest,

a straight or slightly oblique brown callus

point at apex; small gland at base; pulvinus

313

not well defined. Spikes 8-9 mm long, c. 5

mm diam. at anthesis, rachis with scattered

curled hairs; peduncles |~2 mm long, borne

in 2s and 3s on a minute axillary shoot,

occasionally spikes becoming lateral as shoot

elongates; bracteole c. 0.5 mm long, with

oblique claw with long hairs on back. Flowers

5-merous; calyx 0.5-0.7 mm long, cupular,

membranous, sinuolately lobed, rather dense

white arachnoid hairs and a few brown scales

on lobes; corolla 1.3—1.5 mm long, glabrous,

lobed to middle, lobes uninerved, strongly

recurved; stamens 2—2.2 mm long; ovary with

a few long stiff hairs towards the apex. Pods

to 35 mm long, 2.5 mm wide with up to 8

seeds, pale brown with thick marginal nerves.

Seeds longitudinal 2.4—3.2 mm long, 1.3—1.4

mm wide, dark with paler areole; pleurogram

U-shaped; funicle pale, once folded,

thickened into a small aril capping the seed.

Other specinens (all BRD): Queensland, SoUTH KENNEDY

District: Mt Abbot, 20°06’°S 147°46’E, alt. 800m, Oct

1992, Bean 5196; Mt Abbot, 20°07’°S 147°46’E, alt.

400m, Jul 1992, Bean 4753.

Distribution and habitat: The species is

confined to Mt Abbot where it occurs in

heathland on rather steep slopes on skeletal

soils derived from granite. Flowering

material was collected in August and fruiting

in October.

Affinities: Acacia abbatiana is closely related

to A. granitica, and was recorded as such by

Bean (1994), but differs in having lamellated

bark, somewhat shorter phyllodes,

inflorescences of slender spikes on axillary

shoots, smaller flowers (stamens to only 2.2

mim long) and calyx with dense arachnoid hairs

and shorter pods with smaller seeds.

Etymology: The specific epithet is formed from

Latin abbas, -itis, abbot, and the suffix -ana,

indicating position: a reference to Mt Abbot,

the only known locality for the species.

*Acacia burdekensis Pedley sp. nov. affinis

A. leiocalyci (Domin) Pedley a qua habitu

arboris usque 9 m altae, phyllodiis forma

similis eis A. /eptostachyae Benth.

plerumque elongatioribus, plerumque

plus quam 6-plo longioribus quam latis,

saepe angustioribus nervis multis

314

tenuibus parallelis vix anastomantibus

ornatis differt. Typus: Queensland.

NorTH KENNEDY DISTRICT: 8 miles [c. [3

km] W of Pentland, 19 June 1953, M.

Lazarides 3543 (holo: BRI; iso: CANB

(n.v.), PERTH).

Acacia sp. (Torrens Creek C.T. White 8725)

(Pedley 1997),

Tree to 9 m tall; bark dark, rough, furrowed

or tending to form hard rectangular flakes;

branchlets reddish, angular, scurfy, resinous

(as are young phyllodes); stipules deltoid,

0.5-0.8 mm long, caducous. Phyllodes

somewhat crowded towards the ends of the

branches (internodes short), straight to

strongly falcate, widest below the middle,

(7)9.5—14.5 cm long, 7-18 mm wide, 6—11(—

15.5) times longer than wide; glabrous; 2 or

3 longitudinal nerves more prominent than

the rest, secondary nerves parallel, rather

crowded (30—40 per cm) with some oblique

nerves forming anastomoses; tapering to an

acute callus point at the apex; gland basal,

prominent with distinct swelling and

prominent orifice; abruptly contracted to a

reddish pulvinus 4-6 mm long. Spikes

moderately dense, 25-—40(—60) mm long,

rachis glabrous, on peduncles 7-12 mm long

in pairs in upper axils; bracteoles rather stout

0.4 mm long, caducous. Flowers 5-merous;

calyx cupular 0.5-0.75 mm long, lobes 0.1—

0.2 mm long, glabrous except for a few

minute trichomes on margins of lobes;

corolla (1.5—)1.7—2 mm long, glabrous, lobes

c. 1 mm long, strongly reflexed; stamens c.

4 mm long; ovary white-hirsute. Pods not

seen.

Spectnens (all BRI): Queensland. NortH KENNEDY

District: Greenvale Nickel Mine turn off, 18°53’S

144°53’E, Apr 1991, Batianoff GV9104106 & Franks;

10 km NE of ‘Valiey of Lagoons’ H.S., 18°38’S 145°12’8S

May 1971, Blaxvell 503 (ex NSW); Burra Range, Apr

1984, Brown 22; Charters Towers - Clermont Road 42

miles [67 km] from Charters Towers, May 1960, Johnson

1854; 50 km N of Cape River between Charters Towers

and Clermont, Jun 1994, Esser s.n. (AQ 627982); 3 km

W of Noname Hill, Valiey of Lagoons; 18°44’S 145°21’E,

Aug 1988, Mockett s.n. (AQ 476295); Torrens Creek,

Mar 1933, White 8725.

Distribution and habitat: The species is largely

confined to the northern part of the Burdekin

River basin between about 18°30’S and 21°S

Austrobatleya 5(2): 307-321 (1999)

where it occurs on stony and sandy soils on

hillsides and on creek banks.

Affinities: Acacia burdekensis has been

previously included in a rather heterogeneous

A, leiocalyx but differs in being a moderately

large tree with rather elongate phyllodes similar

in shape to those of A. /eptostachya with fine

crowded, only sparingly anastomosing,

secondary nerves.

Etymology: The epithet is a contraction of

‘burdekin’ and Latin -ensis indicating the

occurrence of the species largely within the

northern parts of the basin of the Burdekin

River.

*Acacia faucium Pedley, sp. nov. ramulis

valde angularibus, phyllodiis pulvino

brevi nervis longitudinalibus majoribus

basin versus confluentibus A. /eiocalyci

(Domin) Pedley similis autem

phyllodiis plerumque longioribus

chartaceioribus nervis secondariis

pluribus, spicis brevioribus, calyce

plerumque pilis hyalinis aliquot basi,

valvis leguminis cartilagineis differt.

Typus: Queensland. NortH KENNEDY

District: Bertya Creek, W of

“Warang”’, White Mountains National

Park, 20°27’°S 144°46’E, 21 June 1992,

A.R. Bean 4611 (holo:BRI).

Tree to 10 m tall; branchlets acutely angular,

somewhat scurfy, with sparse minute (c. 0.1

mm long), appressed hairs on young plants,

glabrescent; young growing tips brown;

stipules deltoid c. 0.5 mm long, deciduous.

Phyllodes straight or somewhat falcate,

widest above the middle, 13.5-18 cm long,

(16—)20-27 mm wide, 4.5—7.5(—9) times

longer than wide; rather chartaceous in

texture, glabrous or with sparse appressed

hairs c. 0.1 mm long on young plants; 2 or 3

longitudinal nerves more prominent than the

rest, confluent with each other near lower

margin towards the base, secondary nerves

rather crowded (20-—25/cm) anastomosing;

blunt at apex; gland prominent, basal;

tapering into pulvinus 0.5—0.8 mm long.

Spikes moderately dense, 35-55 mm long,

rachis pruinose, on reddish peduncles 5—8

mm long in pairs in upper axils; bracteoles

FOUR RTE ESS SS Ca oe a aa SS A

SSRs EES

Pedley, L., Acacia (Leguminosae: Mimosoideae) chiefly from northern Australia

c. 0.5 mm long, glabrous, the stipe somewhat

longer than the oblique lamina. Flowers 5-

merous; calyx rather broad, 0.5—0.8 mm long

with obtuse lobes 0.1—0.2 mm long, glabrous

or usually with a few spreading hyaline hairs

towards the base; corolla 1.7—2 mm long,

glabrous, lobed to about the middle, lobes

uninerved, strongly reflexed; stamens c. 2.5

mm long; ovary hirsute with white hairs. Pods

linear, + straight, to c. 8 cm long, 3 mm wide,

with up to 12 seeds; valves brown-black,

cartilaginous, glabrous, somewhat glaucous,

obscurely longitudinally wrinkled, raised

over seeds, marginal nerves thick, prominent,

yellowish; seeds longitudinal, pale brown,

3.5—4 mm long, c. 2 mm wide; areole large;

pleurogram not well defined, oblong, closed

or almost so; funicle yellow, folded about 3

times and thickened to form cupular aril.

Selected specimens (all BRI): Queensland. NortTH

KENNEDY District: Torrens Creek Gorge, White

Mountains, 20°28’°S 144°55’E, Oct 1991, Cumming

11422; ‘Warang’ Holding, White Mountains, 37 km

NNW of Torrens Creek, 20°29’S 144°48’E, Aug 1988,

Fell DF 1369; 88 km NE of Clermont, 22°09’S 148°08’E,

Jun 1972, AfdcDonald 556; 3 miles [c. 5 km] NE of ‘New

Twin Hills’ H.S., Aug 1964, Pedley 1738 (ex CANB).

Distribution and habitat: The species is

common in the headwaters of Torrens Creek

in White Mountains where it occurs in gorges

in sandstone. It also occurs in broken country

farther south, about 100 km north of

Clermont.

Affinities: Acacia faucium 1s closely allied to

A. leiocalyx but has usually longer, more

chartaceous phyllodes with more widely spaced

secondary nerves, shorter spikes, usually a few

hairs on the calyx and valves of the pod

cartilaginous.

Etymology: The specific epithet 1s Latin,

meaning ‘of gorges’ a reference to the habitat

of the species at the type locality.

Acacia filrpes Pedley, sp. nov. non prope

affinis ullae Australiae borealis specie,

phyllodiis teretibus, calycibus lobatis fere

ad basem, leguminibus angustis semina

parum obliqua pleurogrammatibus

inapertis gerentibus, et praecipue

pedunculis perlongis (35-45 mm longis)

315

notabilis. Typus: Northern Territory: 21

km N of Jim Jim Falls, near entrance to

Deaf Adder Gorge, 13°15’S 132°51’E,

29 May 1980, M. Lazarides 9075 cholo:

BRI; iso: (n.v.) CANB).

Spreading shrub to c. 1 m tall and 2 m wide;

stems smooth, grey; branchlets slender, angular,

ribbed, resinous. Phyllodes glabrous, terete (or

drying somewhat angular); 8.5—15 cm long,

0.4—0.8 mm diam. with 8 parallel longitudinal

nerves and a short oblique callus point; gland

basal or up to 5 mm from the base, the phyllode

usually slightly bent when gland not at the base;

pulvinus 1—1.5 mm long. Spikes in pairs at base

of rudimentary axillary shoot, 10-25 mm long

on slender peduncles 35-45 mm long;

bracteoles as long as the calyx, with a thickened

peltate lamina. Flowers 5-merous; calyx

glabrous, c. 0.7 mm long, divided almost to the

base into oblong obtuse lobes; corolla 1—1.2

mim long, divided to about the middle; stamens

c. 3mm long; ovary glabrous. Pods flat, linear,

44.5 cm long, 2.5—3 mm wide; valves slightly

resinous, rather woody with raised

anastomosing nerves. Seeds (up to 8 per pod),

longitudinally to slightly obliquely arranged,

dark grey, c. 3 mm long, 1.5 mm wide;

pleurogram closed; funicle straight, thickened

into a cupular aril similar to that of 4.

calyculata.

Other specimen: Northern Territory, Deaf Adder Gorge,

13°05’S 132°51’E, Feb 1977, Fox 2564 (BRI, DNA).

Distribution and habitat: This is evidently a

rare species confined to the vicinity of Deaf

Adder Gorge where it occurs on the top of the

sandstone escarpment.

Affinities: Acacia filipes 1s apparently not

closely related to any other species. A

combination of attributes (terete phyllodes,

long filiform peduncles, deeply lobed calyx,

pods with reticulately nerved valves and

slightly oblique seeds with a closed

pleurogram) sets it apart from all other species

known to me. The Fox specimen has finer

phyllodes than those of the type and came

from a smaller plant but otherwise the two

specimens are similar.

Etymology: The epithet is derived from Latin

fili, thread and pes, foot, an allusion to the

316

long threadlike peduncles of the

inflorescence.

*Acacia fodinalis Pedley, sp. nov. affinis A.

leiocalyci (Domin) Pedley a qua

phyllodiis plerumque angustioribus

elongatioribus (plus quam 7.5-plo

longioribus quam latis), spicis in

pedunculis longioribus, leguminibus

aneustioribus differt. Typus: Queensland.

LEICHHARDT District: Norwich Park

mine, c. 22°40’S 148°20’E, 8 June 1983,

J. Martin s.n. [AQ 349851] (holo: BRD).

Acacia sp. (Norwich Park J. Martin AQ

349851) (Pedley 1997).

Tree to 10 m tall with grey-brown coarsely

fibrous fissured bark; branchlets reddish,

sharply angular, resinous when young,

sometimes slightly scurfy, becoming smooth;

stipules deltoid, c. 1 mm long, caducous.

Phyllodes slightly falcate, tapering equally to

each end, 8.5—13 cm long, 8—15 mm wide, 7.5—

15 times longer than wide; glabrous; 2 or 3

longitudinal nerves more prominent than the

rest tending to run together near adaxial margin

near base, secondary longitudinal nerves

crowded (25-35 per cm), distinctly

anastomosing; apex acute with a callus point;

gland at, or within 1 mm of base, prominent, a

distinct swelling of margin with a marked

orifice. Spikes rather open, (25—)45~—70 mm

long, rachis glabrous and occasionally

glaucous, on peduncles (5~-)7—10(-13) mm

long in 2s or 4s in upper axils; bracteoles c. I

mm long, a narrow claw and oblique lamina

with a few hairs, caducous. Flowers 5-merous;

calyx cupular, glabrous or with a few hairs near

the base, 0.5—0.8 mm long, lobes c. 0.1 mm

long; corolla c. 1 mm long, glabrous, lobes

strongly reflexed; stamens c. 2.5 mm long;

ovary white hirsute. Pods linear, + straight, to

c. 45 mm long, 2.3—3 mm wide, dark brown,

with marginal nerves yellow; raised over seeds

and slightly constricted between them. Seeds

longitudinal, light brown, c. 3.5 mm long, 1.7-

2 mm wide; areole large, pale; pleurogram

rectangular, open; funicle folded c. 3 times,

thickened into yellow cupular aril.

Specimens (all BRI): Queensland. LEICHHARDT DISTRICT:

2 miles [c 3 km] N of ‘Logan Downs’ Stn, Aug 1964,

Adams 1263 (ex CANB); c. 8 km N of Dysart, Jun 1989,

Austrobaileya 5(2): 307-321 (1999)

Anderson 1011; Riverside coal project, 30 km NW of

Moranbah, Aug 1981, Anon.; Lake Elphinstone, 21°33’S

148°13°E, Jul 1985, Chanipion 146; 17 miles [27 km] E

of ‘Pasha’ homestead, Jul 1964, Pedley 1727A (ex

CANB). Sours KENNepy District: Peak Downs Highway

at top of Eton Range, 21°S 148°52’E, Champion 1341 &

Ritchie.

Distribution and habitat: The species occurs

in the upper part of the Isaacs River basin and

adjacent part of the Belyando River basin where

it occurs usually on sandy soils often associated

with Lucalytpus crebra.

Affinities: Acacia fodinalis is closely related

to A. leiocalyx but develops into a larger plant

with usually narrower and more elongate

phyllodes (more than 7.5 times longer than

wide), spikes on longer peduncles, and

narrower pods.

Etymology: The specific epithet is derived

from Latin fodina, a pit or mine, and the suffix

-alis, pertaining or belonging to, an allusion to

the large colleries in the geographic range of

the species.

Acacia lacertensis Pediecy, sp. nov. prope

affinis A. tropicae Tindale a qua

phyllodiis angustioribus minus crassis

plerumque 2-nervibus, pedunculis

spicarum brevioribus, spicis brevioribus,

probabiliter leguminibus longioribus

angustioribusque semina parviora

gerentibus differt. Typus: NORTHERN

TERRITORY. Narbalek, 12°19’S 133°19’E,

30 August 1988, R. Hinz 51 (holo: BRI

(2 sheets); iso (n.v.): AD, CANB, DNA,

MEL, NSW, PERTH).

Slender tree with sparse canopy to 8 m tall;

bark grey-brown; branchlets stout, angular,

glabrous, pruinose; young tips dark. Stipules

deltoid, c. 1 mm long, caducous. Phyllodes

straight or somewhat sigmoid, attenuate at the

base, 12.5-17(—20) cm long, 1—2(-—2.5) cm

wide, 6.5—12 times longer than wide, glabrous,

rather thin in texture with fine, widely spaced

(c. 0.5 mm apart in middle of phyllode)

longitudinal nerves, two or occasionally three

more prominent, these running together in

middle of phyllode at its base; obtuse at the

apex with a caducous callus; gland at base of

phyllode not prominent; pulvinus 5-14 mm

say ge ie tant y

Pedley, L., Acacia (Leguminosae: Mimosoideae) chiefly from northern Australia

long. Spikes golden-yellow (Chippendale NT

8094), in pairs at base of rudimentary shoot in

upper axils, 4-5 cm long on peduncles 5—8 mm

long. Flowers 5-merous; calyx 0.4—0.5 mm

long, broadly cupular, lobed to about the

middle, the lobes wide, obtuse, glabrous or

fimbriate, with a few hairs towards base of the

tube; corolla 1.4—1.6 mm long, lobed to about

the middle, glabrous; stamens 3-4 mm long;

ovary sparsely sericeous. Pods + straight, 8-9

cm long, 3.5 mm wide with up to [2 seeds;

valves convex over the seeds, with prominent

marginal nerves. Seeds obloid, 3.8-4.5 mm

long, 1.7-2.3 mm wide; pleurogram open;

areole oblong, 2.8—3.5 mm long, c. 1 mm wide;

funicle thickened and folded, forming cupular

aril beneath the seed.

Specimens: Northern Territory. East Alligator River,

12°47’S8 133°23’E, Jul 1972, Byrnes 2751 (BRI, DNA);

Narbalek, Cooper Creek, 12°19’S 133°19°E, Hinz 597

(BRI); Cooper Creek, 28 miles [45 km] N of Oenpelli,

Jul 1962, Chippendale NT 8094 (BRI, DNA); 17.8 km

along turn-off to Murgenella, Cooper Creek, 12°06’S

133°11’E, Sep 1987, McDonald MM 413. (BRI).

Distribution and habitat: The species has been

recorded only from sandy banks of the East

Alligator River and its tributary Cooper Creek.

Affinities: Acacia lacertensis and A. tropica

are closely related. The most obvious

differences between them are the texture of the

phyllodes and the prominence of the nerves.

The significance of the differences in pods and

seeds is difficult to assess as only one fruiting

specimen ofA. fropica has been examined and

the specimen is not a good representative of the

species. Itis George Creek, 18°15’S 137°16’E,

12 Sep 1967, A. Nicholls 733 (BRI; distributed

from DNA as Acacia sp. aff. A. gonoclada). It

has pods which are somewhat immature, about

60 mm long and 5 mm wide with dark valves

with lighter nerve-like margins and longitudinal

seeds. The single seed examined (not quite

mature) is 4.8 mm long, 2.5 mm wide with a

oblong areole; the pleurogram is pale and open,

and the funicle thickened and folded to form a

cupular aril beneath the seed.

Etymology: The specific epithet is derived

from Latin /acerta, lizard and the suffix -ensis

indicating place of origin: a rather indirect

reference to the Alligator River. English

317

alligator is a corruption of the Spanish e/.

lagerta which is derived from the Latin /acerta.

Acacia leiocalyx (Domin) Pedley, Contrib. Qld

Herb. 15:10 (1974), Austrobaileya 1:179

(1978); A. glaucescens var. leiocalyx

Domin, Bibliothec. Bot. 89:269 (1926).

Type: Queensland, prope Brisbane River,

Amalia Dietrich 568 (ex museo

Godeffroy Hamburgensi 5068) (lecto:

PR527897, fide Pedley, 1974).

Acacia leiocalyx 1s widespread in subtropical

Queensland from the coast west to about 147°E

longitude and in north-eastern New South

Wales. It is particularly common in south-

eastern Queensland where it is sympatric with

A. concurrens Pedley. It is conspicuous on

coastal dunes but away from the coast it is less

so. There it usually occurs on shallower soils

on steeper slopes than does A. concurrens. Prior

to 1974 the two were not formally distinguished

and were both referred to as A. cunninghamii

Hook.f., an illegitimate name. A. leiocalyx

differs from A. concurrens in having smooth

(not scurfy), sharply angular, often reddish

branchlets, shorter pulvinuses and completely

glabrous calyxes. It flowers earlier in the year

than A. concurrens, though in some areas it

may flower synchronously and hybrids between

the two are suspected to occur in north-eastern

New South Wales and on the northern outskirts

of Brisbane. Populations of A. leiocalyx west

of the Dividing Range often begin to flower

earlier than populations east of the range at the

same latitude and elevation, have longer

periods of flowering and deeper yellow

flowering spikes. Plants from the two regions are

morphologically similar. A. leiocalyx subsp.

herveyensis 1s distinguished from A. /eiocalyx

subsp. /eiocalyx by its narrow phyllodes widest

below the middle, long attenuate to the apex which

is slightly thickened. The subspecies form mixed

stands, particularly between Bundaberg and

Maryborough, but subsp. ferveyensis stands out,

not only on account of its narrower phyllodes but

also because it flowers later than subsp. leiocalyx.

Some workers, for example Tame (1992) do not

recognise subsp. Herveyensis.

Plants from subcoastal regions north of

the Tropic of Capricorn identified as A.

leiocalyx have been a puzzle. Pedley (1978)

318

referred to a variant from ‘a small area in central

Queensland’ with ‘remarkably consistent

narrower phyllodes’. Collections made since

that time revealed that the situation 1s more

complex than believed. Consideration of these

more recent collections and re-appraisal of

some old ones resulted in the recognition of

A. burdekensis, A. fodinalis and A. faucium.

The last is a distinctive species but the others

are close to A. leiocalyx, distinguished from it

in having more elongate, usually narrower

phyllodes, often with more crowded secondary

longitudinal nerves, longer pulvinuses, and at

least some flowers in the inflorescence with a

few erect hyaline hairs at the base of the calyx.

Mature plants of both species are larger than

those of A. leiocalyx. Though the distinctions

between the two species and between them and

A. leiocalyx are on occasions rather nebulous

they do warrant recognition, if only in providing

a basis for further studies, especially in the field.

Such studies should also include A. crassa

Pedley and A. /ongispicata Benth. which are

sympatric with some of these species.

Even with the recognition of the species

mentioned above, many collections, including

those of the variant with narrow phyllodes

originally noted, cannot be referred to any of

the species. Having studied populations of them

in the Emerald-Blackwater-Duaringa area, I

consider these specimens represent intergrades

between Acacia cretata Pedley and A. fodinalis.

A. cretata on the Blackdown Tableland has

stout angular pulverulent branchlets, large

phyllodes abruptly contracted into short

pulvinuses and long spikes. At lower altitudes

along the northern edge of the Tableland, plants

with more slender glaucous branchlets,

somewhat smaller phyllodes tapering into

longer pulvinuses, and shorter inflorescences

occur. These may conveniently be considered

A. cretata. Other plants, however, lack the

extreme characters ofA. crefata though angular

glaucous branchlets invariably occur; these are

considered to be the intergrades with A.

fodinalis. The zone of intergradation is an arc

some 80—100 km wide from the north-west to

the north-east of Blackdown Tableland.

Acacia proiantha Pedley, sp. nov. non arcte

speciebus ceteris Acaciae sectionis

Juliflorae affinis ; phyllodiis 11-17.5 cm

Austrobatieya 5(2): 307-321 (1999)

longis, 1—1.5 mm latis nervum medium

ceteris prominentiorem habientibus,

floribus parvis petala c. 1 mm longa

gerentibus, leguminibus aliquantum

moniliformis valvis reticulatim nervatis,

seminibus parvis c. 2.6 mm longis et 1.6

mm latis insignis. Typus: Northern

Territory. c. 2.5 miles [4 km] SW of Mt

Gilruth, 13°03’S 133°02’E, 28 February

1973, M. Lazarides 7938 (holo: BRI; iso

(n.v.): CANB).

Erect spindly shrub with sparse canopy to c. 3 m

tall; stems smooth, grey; branchlets angular,

glabrous, slightly resinous; young tips also

resinous. Phyllodes rather thick, linear, 11—-17.5

cm long, 1-1.5 mm wide; glabrous; one

longitudinal nerve forming a midrib, with 4-6 less

prominent non-anastomosing nerves on each side

of it; acute at the apex with a deciduous callus

point; gland basal, inconspicuous; pulvinus | .5—

3 mm long. Spikes single at the base of a

rudimentary axillary shoot, 15-20 mm long, on

peduncles 4-6 mm long, the peduncle and axis

somewhat resinous; bracteoles as long as the

calyx, with a narrow claw and lamina peltate,

thickened. Flowers 5-merous; calyx broadly

cupular, somewhat resinous, 0.4—0.6 mm long,

lobed to the middle, the lobes thick, obtuse,

sometimes with a few marginal hairs; corolla 0.9—

1.1 mm long, lobed to the middle, glabrous. Pods

straight, linear acute at the tip, the valves rather

thick with raised reticulate nerves, constricted

between the seeds and raised over them, to 6.5

cm long, 2.5 mm wide at widest part, 1.5 mm

wide at isthmuses, glabrous, slightly resinous.

Seeds (8 per pod) longitudinally arranged,

obloid, c. 2.6 mm long, 1.6 mm wide; areole

pale, narrow, oblong; pleurogram obscure,

open; funicle thickened into pale aril folded

twice beneath the seed.

Specimens: Northern Territory. 3 miles [S km] E of

Jim Jim Falls, Jul 1972, Byrnes 2724 (BRI, DNA, K);

Deaf Adder Gorge, 13°02’S 133°05’E, Feb 1977, Fox

2519 (BRI, DNA).

Distribution and habitat: The species is

restricted to the northern part of the Northern

Territory where it occurs on sandstone.

Affinities: Acacia proiantha is not closely related

to any other species of Acacia sect. Juliflorae. It

has long narrow phyllodes with one nerve more

Pedley, L., Acacia (Leguminosae: Mimosoideae) chiefly from northern Australia

prominent than the rest, forming a distinct midrib;

its flowers are small with petals only about 1 mm

long; and its pods are somewhat moniliform with

small longitudinal seeds.

Etymology: The epithet is derived from Greek

prois, early in the year, and anthos, a flower, an

allusion to the February flowering of the species.

Acacia scopularum Pedley, sp. nov. affinitatis

incertae A. spirorbi subsp. solandri (Benth. )

Pedley propter amplitudinem formamque

phyllodiorum, spicas interruptas in

pedunculo brevi primo adspectu simile, sed

nervis longitudinalibus phyllodiorum non

crebris, floribus 4-meris, petalis multo

brevioribus, ovario glabro, leguminibus non

torsivis differt. Arbor vel frutex usque 5 m

altus. Phyllodia glabra linearia, parum

falcata, 7—11 cm longa, 4.5—6 mm lata, 15—

20-plo latis longiora; nervis longitudin-

alibus 8-14 late separatis aliquando

anastomosantibus instructa; pulvinus 1.5—

2.5 mm longus. Flores 4-mert; calyx 0.5

mm longus, non nisi leviter lobatus;

corolla 1.5 mm longa; ovarium glabrum.

Legumina linearia, leviter constricta inter

semina et convexa super ea, usque 9 cm

longa, 2 mm lata, valvis reticulatim nervatis

et marginibus incrassatis praedita; seminia

parva in longitudinem disposita. Typus:

Northern Territory. ESE of Mudginbarry,

12°36’°S 132°58’E, 19 February 1973, CLR.

Dunlop 3313 (holo: BRI; iso (n.v.):

CANB, DNA, MEL).

Tree or large shrub, branches sometimes

sprawling, to 5 m high; bark rough, fissured;

branchlets dark red, angular, glabrous; stipules

minute or absent. Phyllodes narrow, lanceolate

falcate, widest above the middle, 7-11 cm long,

4.5—6 mm wide, 15—20 times longer than wide,

glabrous; a callus point, sometimes oblique at

the apex; with 8—14 widely spaced (0.3—0.5 mm

apart) longitudinal nerves, an occasional

anastomose; gland at the base; pulvinus 1.5—

2.5 mim long. Flowers tn interrupted spikes 3.5—

4.5 cm long in pairs, with a rudimentary axis

between them, in the upper axils; rachis

glabrous; peduncles 1.5~2 mm long. Flowers

4-merous; calyx 0.5 mm long, only shortly

lobed, glabrous except for a few short hairs on

5 oe

the lobes; corolla lobed to the level of the calyx,

c. 1.5 mm long, lobes with a distinct midrib,

strongly reflexed; stamens c. 2.5 mm long;

ovary glabrous. Pods glabrous, linear, straight

or slightly curved, with up to 15 seeds, 8-9 cm

long, 2 mm wide, valves reticulately veined

with thickened margins raised over the seeds.

Seeds longitudinal obloid, 2—2.5 mm long, 1.5—

1.7 mm wide; areole large, oblong; pleurogram

conspicuous, open; funicle folded, thickened

into small yellow cupular aril.

Specimens: Northern Territory. Radon Creek, Mt

Brockman, 12°45’S 132°56’°E, Dunlop 4679 (BRI,

DNA); Little Nourlangie Rock, 12°51°S 132°50’°E, Fox

2568 (BRI, DNA); Deaf Adder Gorge, 13°02’S 133°05’E,

Fox 2433 (BRI, DNA); ‘Common Rock’ Creek, Jabiluka

Outlier, Waterhouse 9620 (BRI).

Distribution and habitat: The species is

restricted to a small area in the northern part of

the Northern Territory where it is found in

shallow soils often on tops of cliffs.

Affinities: The relationships of A. scopulorum

are not at all clear. In general appearance it

resembles A. spirorbis and.A. leptostachya, and

has been distributed from DNA as the latter,

but differs from both in its small 4-merous

flowers. On the other hand, it does not appear

to be particularly close to the 4-merous

juliflorous species of south-eastern Queensland

(Pedley 1964).

Etymology: The specific epithet is genitive

plural of Latin scopulus, a rock or cliff, a

reference to the habitat of the species.

* Acacia solenota Pedley, sp. nov. quoad ramulos

complanatos, florium structuram

amplitudinemque, inflorescentias paene

albas, semina parva funiculo recto A.

calyculatae A.Cunn. ex Benth. similis

autem ab ea plantae statura majore (frutex

densus usque 5 m altus), cortice fibrosa,

phyllodiis saepe brevioribus, latioribus et

minus elongatis, praecipue leguminibus

angularibus valvis profunde canaliculatis

constructis distinguitur. Typus:

Queensland. Cook District: 11.9 km E of

Hopevale-Starcke road, on track to the

Mclvor River mouth, 15°06’S 145°12’E,

14 August 1984,/.R. Clarkson 5475 (holo:

BRI; iso: MBA, CANB, DNA, IK, MEL,

MO, NSW, PERTH, QRS).

320

Dense spreading shrub to c. 5 m tall, branching

from near the base; bark grey-brown,

longitudinally fibrous; branchlets glabrous,

complanate; young growth reddish brown and

scurfy. Phyllodes coriaceous, elliptic,

asymmetric, lower margin more or less straight,

upper curved, 8—-10.5(-12.5) cm long, 15-25(—

32) mm wide, 3.2—5.5 times longer than wide;

glabrous; with many fine longitudinal non-

anastomosing nerves, 2 or 3 more prominent than

the rest; obtuse with a small callus mucro; gland

at the base of the phyllode inconspicuous;

pulvinus 3-7 mm long. Flowers in dense spikes,

25-35 mm long, almost white, on peduncles 6-8

mm long, 1, 2 or 3 in the upper axils, each

peduncle subtended by a small ovate bract, rachis

glabrous; bracteoles curved, concave, hirsute on

the back, about as long as the calyx. Flowers 5-

merous; calyx broadly cupular, sinuolately lobed,

hirsute at base, c. 0.4 mm long; corolla deeply

lobed, glabrous, c. 1.8 mm long, midribs of lobes

prominent; stamens c. 3 mm long, ovary glabrous.

Pods straight, to 12 cm long, 8-10 mm wide, c. 8

mm thick, woody; valves widest near apex,

narrowed to the base, opening elastically from

the apex; each valve with a longitudinal dorsal

groove c. 3 mm deep. Seeds up to 12 in each

pod, longitudinally arranged; none seen but

impressions in valves quite plain.

Specimens: Queensland. Cook District: 12.5 km NW of

the beach on the track from Starcke Station to Mclvor River

mouth, 15°04’S 145°09°E, Feb 1984, Clarkson 5145 (BRI,

K, MBA, MEL, NSW, PERTH, QRS); Leprosy Creek,

Cooktown, Oct 1986, McLean (AQ441388) (BRI).

Distribution and habitat: The species is

confined to Quaternary sand-dunes between

Cooktown and Cape Flattery where it occurs,

sometimes in dense pure stands, in scrubs.

Affinities: Acacia solenota is closely related

to A. calyculata but is a much larger plant

with usually shorter and wider phyllodes and,

above all, by the deeply grooved valves of

the pod.

Etymology: The epithet is derived from

Greek solen, solenos, channel or pipe or

deeply grooved tile, a reference to the

characteristically grooved valves of the pod.

* Acacia tingoorensis Pedley, comb. et stat.

nov.

Austrobatleya 5(2): 307-321 (1999)

Acacia longispicata subsp. velutina

Pedley, Austrobaileya 1:176(1978).

Type: Queensland. BurRNETT DISTRICT:

12 km [sic] NW of Kingaroy, 26°23’S

151°41’E, 19 August 1973, L£. Pedley

4134 (holo: BRI; iso: A, B, BRI,

CANB, E, L, MO, NSW, PR).

Distribution and habitat: The species is

restricted to a low hill some 25 km NW of

Kingaroy. It occurs on shallow loamy and

sandy soils as understory in eucalypt

woodland and forms dense stands in disturbed

situations on roadsides.

Affinities: Acacia tingoorensis is one of a

taxonomically ‘difficult’ group of species that

includes A. concurrens Pedley, A. crassa

Pedley, A. leiocalyx and A. longispicata

Benth. It differs from the last, to which it was

referred as a subspecies, in having dense erect

velvety (not appressed) hairs on the

branchlets extending to the rachis of the

spikes. Pods and seeds may provide

additional differences but pods of A.

tingoorensis are unknown to me. Though

plants flower every year, all attempts at

collecting fruits in the last five or six years

have failed.

Etymology: The epithet is derived from

Tingoora, the name of the nearest centre of

population, some 10 km east of the type

locality.

SLY A etd tee de ec pe ee ee et gee EEE Se enn gtt tt eteeeeeeeee nee eeeee es Tee at

Pedley, L., Acacia (Leguminosae: Mimosoideae) chiefly from northern Australia

References

BEAN, A.R. (1994), An analysis of the vascular flora of

Mt Abbot near Bowen, Queensland. Proceedings

of the Royal Society of Queensland 104:43—66.

HENDERSON, R.J.F. (1987), Liliaceae (in part) in Flora of

Australia 45:175—232, 264-268, 281-299, 299-

306, 348-350.

PepLey, L. (1964). Notes on Acacia, chiefly from

Queensland, II. Proceedings of the Royal Society

of Queensland 75:29-35,

—— (1978). A revision of Acacia Mill. in Queensland.

Austrobaileya 1375-280.

-—— (1987). Notes on MRacosperma Martius

(Leguminosae: Mimosoideae), I. Austrobaileya

2:32 1—327.

—— (1997), Mimosaceae. In R.J.F, Henderson (ed.),

Oueensland Plants: Names and Distribution.

Brisbane: Queensland Herbarium, Queensland

Department of Environment and Heritage.

TAME, T. (1992). Acacias of southeast Australia.

Kangaroo Press: Kenthurst, NSW.

eldeieaee a 7 we ttnp dt etitadt feed ead adads pietedenapeeneenenedege eee 2 OP Oe aged ab ‘

Two new species of Stylidium Willd. (Stylidiaceae)

from north Queensland

A.R. Bean

Summary

Bean, A.R. (1999). Two new species of Stylidium Willd. (Stylidiaceae) from north Queensland.

Austrobaileya 5(2): 323-330. Two new Stylidium species, S. letophyllum and S. ramosissimum, and

their closest relative, S. eriorhizum R.Br. are described, illustrated and distinguished in a key.

Distribution maps, and notes on conservation status are provided.

Key words: Shlidium, Stylidiaceae, Australian flora, key, St-lidium eriorhizum, Stylidium leiophyllum,

Stylidium ramosissimum.

A.R. Bean, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha road,

Toowong, Queensland 4066, Australia.

Introduction

The genus Stylidium occurs throughout

Australia, especially in the south-west and

across the tropics. A few species also extend to

Malesia (Slooten 1954) and into south-east Asia

(Anon. 1972). A monograph of the genus was

provided by Mildbraed (1908), who provided

a subgeneric and sectional classification of the

genus, Mildbraed placed S. eriorhizum R.Br.

in S. sect. Debiles Milbraed.

S. eriorhizum and the two new species

described here all have a thick woolly base at

sround level, above which the leaves emerge.

The thick woolly plant base is rare in the genus,

being otherwise known in S. eglandulosum

F.Muell. (Qld, N.S.W.) and S. Aumphreysii

Carlquist (W.A.).

S. eriorhizum and the two newly described

species form a rather distinctive group within

the genus, most noticeably by virtue of the

woolly plant base and broad rosetted leaves with

a hair-like mucro, but also by the indeterminate

central rachis of the inflorescence, the mucronate

bracts and the more or less spherical seeds with

a colliculose surface.

Key to the species of the Stplidium eriorhizum group

1. Inflorescence branches and bracts glabrous, leaves (including petioles)

Oa VA AAPA LONE 08 a Gor cpt i ih Fels aeieswes

eee ke eel in edt S. ramosissimum

Inflorescence branches and bracts glandular-hairy, leaves (including

petioles) 20—-60(-100) mm long........

2. Leaf margins hairy; bracts 1.5—2.5 mm long; calyx lobes 1,0-1.7 mm long. ..S. eriorhizum

Leaf margins glabrous; bracts 3.5—6.5 mm long; calyx lobes 2.42.8 mm long. . .S. leiophyllum

Stylidium eriorhizum R.Br., Prodr. 569

(1810); Candollea eriorhiza (R.Br.)

F.Muell., Syst. Census Austral. pl. 86

(1883). Type: Queensland. Port Curtis

District: Thirsty Sound, Shoalwater Bay,

3 September 1802, R. Brown (holo: BM).

Accepted for publication 28 August 1998

Perennial herb, 12—25 cm high. Glandular hairs

present, 0.3-0.6 mm long, glands ellipsoidal.

Base of plant woolly, consisting of densely

packed eglandular multicellular trichomes up

to 9 mm long. Stems greatly reduced, appearing

absent, with leaves 1n basal rosette. Leaves 20—

100, spathulate to oblanceolate, grading into

indistinct petiole, 20-60(-100) mm long

324

(including petiole), 4-10 mm wide, glabrous

except for short eglandular hairs along margin;

apex obtuse, but with slender acumen 2—7 mm

long. Scapes |—3 per plant, 1—-1.2 mm in

diameter, glandular-hairy. Inflorescences

(including scape) 10—23 cm long, central rachis

indeterminate, branches monochasially (or

rarely dichasially) cymose, glandular-hairy;

peduncles 5-10 mm long; bracts ovate to

deltate, 1.5-2.5 mm long, glandular-hairy,

mucronate. Flowers solitary in axils of bracts.

Pedicels 3-6 mm long, glandular-hairy.

Hypanthium ellipsoidal, c. 2.5 mm long at

anthesis, glandular-hairy throughout. Sepals

lanceolate, all free, 1.0-1.7 <x 0.5—0.7 mm,

glandular-hairy, apex obtuse or acute. Corolla

pink to white, glandular-hairy on tube and

petals; tube 1.5—2.0 mm long, with sinus on

anterior side only; petals laterally paired,

anterior petals 2.5-3.3 x 1.0—-1.4 mm, entire,

obtuse; posterior petals 2.5-3.5 x 1.0-1.5 mm,

entire, obtuse. Throat appendages absent.

Labellum broadly ovate, 0.6—-0.9 mm long,

attached at base of anterior sinus, glabrous,

apex obtuse, basal appendages 2, minutely

papillose. Column of uniform width throughout,

7.5-8.5 mm long, glabrous; stigma cushion-

shaped; anthers fringed by short eglandular

corona, extending just beyond column. Capsule

obovoid to ellipsoidal, 5-6.5 mm long

(excluding sepals), 2.5—3 mm wide, faintly 5-

ribbed. Seeds spherical with small nipple, 0.4—

0.5 mm diameter, dark brown to black, surface

colliculose (Fig. | G—I).

Selected specimens: Queensland. Cook District: Portion

200, 6 km NW of Atherton, Jan 1966, Hyland 4077 (BRD.

NortH KeNNeDy District: Mt Abbot, 50 km W of Bowen,

Mar 1992, Bean 4207 (BRI); 1.5 km (by road) west of

Herberton, on Herberton-Petford road, May 1983, Conn

& Clarkson 1124 (BRI, CANB, HO, MBA, MEL, NSW);

Harold Island, Nov 1985, Batianoff 3396 & Dalliston

(BRI); west of Kaban, Nov 1989, Elick 85 (QRS); 17 km

past Paluma on road to Hidden Valley, Jan 1992, Forster

PIF9485 (BRI); Blencoe Creek, adjacent to the Cashmere-

Kirrama road, Dec 1995, Wannan 235 (BRI);

Hinchinbrook Island, Jun 1987, Warrian CW8217 (BRI).

SOUTH KENNEDY District: Cape Palmerston N.P., beach

hut headiand, Sep 1995, Champion 1247 & Pollock

(BRI); 9 miles [14 km] NE of “Glen Avon” H.S., Jul 1964,

Pedley 1727 (BRD; 12 km SW of “Mt Douglas” HLS.,

Jun 1992, Thompson BUC477 & Sharpe (AD, BRI).

MitTcHELL District: c. 15 km SSE of “Warang”, NNW of

Torrens Creek, Oct 1988, Crmming 8492 (BRI); 20 km

from Jericho on Blackall road, Feb 1994, Forster

PIFIS014 & Bean (BRI). LEICHHARDT DisTRICT:

Austrobaileya 5(2): 323-330 (1999)

“Wandobah”, c. 11 km NE of Dingo, Jul 1987, Anderson

4351 (BRI, CANB); Mount Flora-Dingo road, 6 km S of

May Downs turnoff, Jul 1992, Bean 4674 (BRD;

“Humboldt”, 45 km NE of Rolleston, Jan 1996, Bean

9584 (BRI); Carnarvon Range, between Roma and

Springsure, Oct 1933, White 9470 (BRI). Porr Curtis

District: west coast of Shoalwater Bay, near Mooly Ck,

Apr 1945, Blake 15587 & Webb (BRI). Burnett District:

on Eidsvold-Cracow road, 1 km N of Little Morrow Creek

crossing, Jul 1990, Forster PIF7006 (BRI, MEL).

DARLING Downs District: north of Waaje tower, Barakula

S.F., north of Chinchilla, Mar 1994, Bean 7566 (BRI).

Distribution and habitat: S. eriorhizum 1s

widespread in Queensland as far north as

Atherton, and as far south as the Barakula State

Forest near Chinchilla. It extends to the central

Queensland coast, and to some continental

islands, and inland as far as Jericho and Torrens

Creek (Map 1). It inhabits woodlands and

heathlands on sandy soils.

HME SERRE

EERE

L eT er CEE Hh

Teun ee

ae eeee

Grins.

PERE TENE

CCC RR

CCRC EN

CER CO BB

Map 1. Distribution of Shlidium eriorhizum @ and

S. PQMOSISSIMIUM A,

Phenology: Flowers and fruits can be found

throughout the year.

Note: The spelling of the species epithet

eriorhizum is a matter of some contention.

According to Stearn (1992: 261), the second

part of a compound Greek word beginning with

rh should have an additional r added to it when

preceded by a vowel, but he also points out

seat ithoeet pitch ecaiat tit tte te bi yeah ee eee ee een ey gene yee eee ee ee eeeee te toe te

RSS Sa ed ing tieg

Sar ere

SSSR REE ere BB ah Se NE Pe

ASSES EES PEE Ea

Bean, A.R., Two new species of Stvlidium Willd. (Stylidtaceae) from north Queensland

325

the

ree

“C8

600 F

ray

Ono 8

Ak jel

Vs600

UERO Gog

ce sg

nto'd x

Fig. 1. A-D: Séplidium ramosissimum, A. portion of inflorescence x 3, B. flower x 6. C. labellum x 20, D. seed x 24.

E-F: Stplidium leiophyllum. E. flower x 6. F. labellum x 20. G-I: Stylidium eriorhizum. G. fruit x 6. H. leaf x 3. 1.

close-up of leaf margin x 9. A-C, Forster 12850 & Bean; D, Bean 1740; E-F, Clarkson 7722 & Neldner; G, Johnson

2140; H-I, Forster 3740,

326

that many reputable authors have omitted this

additional 7, and so it is best regarded as

optional and an author’s original spelling

should be accepted. Brown spelt his epithet

eriorhizum, and hence this spelling is

maintained here. By contrast, Mueller named

Stylidium leptorrhizum with the additional r,

and this spelling should also be maintained.

Conservation status: S. ertorhizum 18 a

common and widespread species.

Stylidium leiophyllum A.R.Bean sp. nov.

affinis Stylidio eriorhizo sed foliis

apicibus acutis et marginibus glabris,

bracteis 3.5—6.5 mm longis, calycis lobis

2.42.8 mm longis et petalis aliquanto

longioribus differens. Typus: Queens-

land. Cook District: 0.9 km north of the

Big Coleman River on the Peninsula De-

velopment road, 14°34’S 143°25’E, 2]

December 1988, /.R. Clarkson 7722 &

VJ. Neldner (holo: BRI; iso: DNA, K,

MBA, PERTH, QRS).

Stylidium sp. (Big Coleman River

J.R.Clarkson+ 7722) in Henderson

(1997).

Perennial herb 8-17 cm high. Glandular hairs

0.25—-0.6 mm long, glands ellipsoidal. Base of

plant woolly, consisting of densely packed

eglandular multicellular trichomes up to 9 mm

long. Stems greatly reduced appearing absent,

with leaves in basal rosette. Leaves 20—100 per

plant, narrowly oblanceolate to narrowly

spathulate, grading into indistinct petiole, 28—

50 mm long (including petiole), 3-6 mm wide,

glabrous, including margins; apex acute, but

with slender acumen to 2.5 mm long. Scapes

1—6 per plant, 0.6—1.1 mm in diameter, hairs

erect and glandular or crisped and eglandular.

Inflorescences (including scape) 8-17 cm long,

central rachis indeterminate, branches

dichasially or monochasially cymose,

glandular-hairy; peduncles 5—7 mm long; bracts

deltate, glandular-hairy, 3.5-6.5 mm long, apex

acute to mucronate. Flowers solitary in axils

of bracts. Pedicels 3-4 mm long, glandular-

hairy. Hypanthium ellipsoidal, 2.5—2.8 mm long

at anthesis, glandular-hairy throughout. Sepals

deltate, all free, 2.4-2.8 x 0.8-1.0 mm,

glandular-hairy, apex obtuse to acute. Corolla

Austrobaileya 5(2): 323-330 (1999)

pink, glandular-hairy on petals only; tube 1.5—

1.8 mm long, with sinus on anterior side only;

petals laterally paired, anterior petals 3.0—3.5

x 1.4—1.6 mm, entire, obtuse; posterior petals

3,.5-4.0 x 1.8—2.0 mm, entire, obtuse. Throat

appendages absent. Labellum ovate, c. 1 mm

long, attached at base of anterior sinus, apex

obtuse; basal appendages 2, minutely papillose.

Column of uniform width throughout, glabrous,

7-7.5 mm long; stigma cushion-shaped; anthers

fringed by short eglandular corona, not

extending beyond column. Capsule ellipsoidal

to obovoid, 3.5~5.2 mm long (excluding

sepals), 1.7-3.1 mm wide, faintly 5-ribbed.

Seeds globose with small nipple, 0.5—0.6 mm

diameter, dark brown, surface colliculose (Fig.

1 E-F, Fig. 2).

Specimens examined: Queensland. Cook District:

upper reaches of Garden Creek, E of Jowalbinna-

Maytown road, Jui 1990, Bean 1760 (BRI); south-east

of Isabella Falls, towards Cooktown, Jul 1998, Bean

13636 (BRI, NSW); north-west of Cooktown near Isabella

Falls, May 1970, Blake 23418 (BRI); 19 km S of the

Palmer River crossing on the Peninsula Development

road, Mar 1987, Clarkson 6646 & McDonald (BRI, K,

MBA, PERTH, QRS); 13.3 km from the Mclvor River

crossing on the Hopevale to Starcke road towards

Battlecamp, May 1993, Clarkson 10070 & Ne/dner (BRI,

K, MBA, PERTH).

PTELELLLLLE LLL

CEPR

P| Tabet PL

abe

Pt td Ete TPR

SUSUSARBIRBISR Nn

Map 2. Distribution of Stvlidium leiophyllum *.

327

“a htt

a“

Merete :

fies

wm letophyllum x 1.

“rag

Bean, A.R., Two new species of Stlidiuim Willd. (Stylidiaceae) from north Queensland

328

Distribution and habitat: S. leiophyllum 1s

known from a few locations in south-eastern

Cape York Peninsula, between Coen and Mt

Carbine (Map 2). It grows in shallow to deep

sandy soils, on flat to very hilly sites, which

may be rocky. It is associated with species such

as Melaleuca citrolens Barlow, M. viridiflora

Sol. ex Gaertn., Eucalyptus nesophila Blakely

and Petalostigma banksii Britten & S.Moore.

Phenology: Flowers are recorded for

December and March; fruits are recorded for

May and July.

Affinities: S. leiophyllum is close to

S. eriorhizum, but differs by its leaves with

acute apices (obtuse for S. eriorhizum) and

glabrous margins (hairy for S. eriorhizun),

bracts 3.5—6.5 mm long (1.5-—2.5 mm for

S. erlorhizum), sepals 2.4—2.8 mm long (1.0—

1.7 mm for S. eriorhizum), and tts somewhat

larger petals.

Conservation status: S. leiophyllum has a

scattered and somewhat restricted distribution, but

itis not considered rare or threatened at this trme.

Etymology: The epithet is from the Greek, /eio-

smooth, and -phy//us- leaf. This is in reference

to the glabrous leaf margins, which

distinguishes it from S. eriorhizum.

Stylidium ramosissimum A.R.Bean sp. nov.

affinis Stylidio eriorhizo, sed foltis multo

minoribus, scapo pilis glandularibus

carente, inflorescentia semper

dichasialiter ramosa, bracteis longioribus

et seminibus majoribus differens. Typus:

Queensland. Cook District: Turtle Rock

area, Laura sandstone escarpment,

15°39°S 144°30’°E, 22 January 1993, PJ.

Forster PIF12850 & A.R. Bean (holo:

BRI; iso: DNA, MEL).

Stylidium sp. (Laura L.S.Smith 12050) in

Henderson (1997).

Perennial herb 11-17 cm high. Glandular hairs

0.2—0.4 mm long, glands ellipsoidal. Base of

plant woolly, consisting of densely packed

eglandular multicellular trichomes up to c. 3

mm long. Stems greatly reduced appearing

absent, with leaves in basal rosette. Leaves 7—

Austrobaileya 5(2): 323-330 (1999)

20 per plant, oblanceolate to broadly spathulate,

srading into indistinct petiole, 9-14 mm long

(including petiole), 1.8-4.5 mm wide, glabrous

except for short eglandular hairs along margin;

apex obtuse, but with slender acumen to 1.2

mim long. Scapes 1—5 per plant, with eglandular

hairs. Inflorescences (including scape) 11-17

x 1.5—3 cm long, central rachis indeterminate,

branches dichasially cymose, glabrous;

peduncles 2.5—3 mm long; bracts deltate,

glabrous, 2.8--5.0 mm long, mucronate. Flowers

solitary in the axils of the bracts. Pedicels 3.0—

3.5 mm long, glandular-hairy. Hypanthium

ellipsoidal, 3-4 mm long at anthesis, glandular-

hairy throughout. Sepals deltate, all free, 2.3—

2.8 x 0.9—[.0 mm, glandular-hairy, apex obtuse.

Corolla pink, glandular-hairy on petals only;

tube 1.6—2.2 mm long, with sinus on anterior

side only; petals laterally paired, anterior petals

3,0—3.5 x 1.4—1.7 mm, entire, obtuse; posterior

petals 4.0-4.5 x 1.5—1.8 mm, entire, obtuse.

Throat appendages absent. Labellum broadly-

ovate, c. 0.6 mm long, attached at base of

anterior sinus, apex obtuse or retuse, basal

appendages 2, minutely papillose. Column of

uniform width throughout, glabrous, 7.5—8 mm

long; stigma cushion-shaped; anthers fringed

by short eglandular corona, not extending

beyond column. Fruit ellipsoidal, 4.6-5.0 mm

long (excluding sepals), 1.8—2.5 mm wide,

faintly 5-ribbed. Seeds spherical except for

prominent nipple, 0.7—0.8 mm across, brown,

surface colliculose (Fig. 1. A—D, Fig. 3).

Specimens examined: Queensland, Cook District: 7 km

E of Jowalbinna, Jul 1990, Bean 1740 (BRI); W of

Branningham Bluff, c. 25 km NW of Cooktown, Jul 1990,

Bean 1974 (BRI); Split Rock gallery, 12.7 km south of

Laura, May 1982, Clarkson 4274 (BRI, QRS); c. 5-6

miles [8—10 km| SE of Laura, Oct 1962, Smith 12050

(BRI).

Distribution and habitat: S. ramosissimum

is confined to a relatively small area at the

southern end of Cape York Peninsula, from

Jowalbinna to near Cooktown (Map 1). It is

confined to sandstone hillsides and

escarpments in woodland dominated by

Eucalyptus stockeri D.J.Carr & S.G.M.Carr,

F.. tetrodonta F.Muell., FE. phoenicea F.Muell.

or E. nesophila Blakely.

Phenology: Unknown; it possibly flowers and

fruits sporadically throughout the year.

329

Bean, A.R., Two new species of Stlidium Willd. (Stylidiaceae) from north Queensland

Hake a

imum <x 1,

Hi FaTOSISS

Fig. 3. Whole plant of Styli

fe Ce Sa EDS It iten ects nt reer BO TLE Be re EIS UD be DE te Fe Bg 2g SA EE PE EM ESE EA ot

330

Affinities: S. ramosissimum differs from

S. eriorhizum by its leaves 9-14 mm long (20—

60(100) mm long for S. eriorhizum), scape

lacking glandular hairs, strictly dichasially

branched inflorescence (usually monochasial

for S. eriorhizum), bracts 2.8—5.0 mm long

(1.52.5 mm long for S. eriorhizum) and seeds

c. 0.9 x 0.75 x 0.75 mm (0.4—0.5 mm diameter

for S. eriorhizum).

S. ramosissimum differs from S.

leiophyllum by its labellum only 0.6 mm long

(obtuse and 1.0 mm long for S. /eiophyllum),

the glabrous peduncles and bracts, and the

leaves (including petioles) 9-14 mm long with

hairy margins (20-60(—100) mm long with

glabrous margins for S. leiophyllum),

Conservation status: S. ramosissimum has a

scattered and somewhat restricted distribution,

but it is not considered rare or threatened at

this time.

Etymology: The specific epithet is derived from

the Latin wordramosissimus, meaning very much

branched. This is in reference to the inflorescence.

Austrobaileya 5(2): 323-330 (1999)

Acknowledgements

| would like to thank Don Foreman (Australian

Botanical Liaison Officer 1996—97) for

photographing the type of S. eriorhizum, Peter

Bostock for the Latin diagnoses, the Director

of QRS for access to that Herbarium and Will

Smith for the illustrations.

References

ANON. (1972). Jeonographia Cormophytorum Sinicorum

4: 401, Beijing: Science Press.

HENDERSON, R.J. (1997). Strlidiaceae. In R.I.F,. Henderson

(ed.) Queensland Plants: Names and Distribution.

Brisbane: Queensland Department of

Environment.

MILDBRAED, G.W.J. (1908). Stylidiaceae. Das

Pflanzenreich Heft 35. Leipzig: Englemann.

SLOOTEN, D.F. van (1954). Stylidiaceae. In Steenis,

C.G.G.J. (ed.), Flora Malesiana Ser 1, 4: 529—

32. Djakarta: Noordhoff-Kol ff.

STEARN, W,T. (1992). Botanical Latin, 4th ed. Newton

Abbot: David & Charles.

ATS Ae ah od LOL Oe eee ee eye Bee

A new species of Habenaria Willd. (Orchidaceae)

from North Queensland |

Peter S. Lavarack and Alick W. Dockrill

Summary

Lavarack, Peter S. and Dockrill, Alick W. A new species of Habenaria Willd. (Orchidaceae) from

North Queensland. Austrobaileya 5(2): 331-335. Habenaria praecox Lavarack & Dockrill is described

and illustrated. Relationships of this species to other Australian species are discussed and a key to

Queensland species of Habenaria and Peristylus is provided.

Dr P.S. Lavarack, Queensland Department of Environment and Heritage, Box 5391, Townsville Mail

Centre, Queensland 4810.

A.W. Dockrill, c/- Box 58, Atherton, Queensiand 4883.

Introduction

For a number of years the species Habenaria

orchroleuca R.Br. was considered to occur in

north Queensland. A.W. Dockrill (1969)

illustrated material of a taxon in the genus

Habenaria as H. ochroleuca. Research by D.L.

Jones (pers. com.) in the Northern Territory,

from where Brown collected material he

originally described as H. ochroleuca,

established that the material previously thought

to belong to this taxon in north Queensland, is

significantly different from that from the

Northern Territory and probably represents an

undescribed species. Comparisons with the type

description of H. ochroleuca (Brown 1810),

also suggested that the taxa are separate. A

specimen was sent to M.A. Clements,while he

was at Kew, and he agreed that the North

Queensland taxon was distinct from H.

ochroleuca (M.A. Clements pers. com.). In

early 1997 one of the authors (P.S.L.) found

flowering plants of this taxon near Cardwell in

north Queensland. This fresh material allowed

further comparisons to be made with illustrations

of H. ochroleuca (Jones 1988), and these

confirmed that the north Queensland material

was distinct. Comparisons were made with

descriptions and illustrations of Habenaria

species from adjacent regions including New

Guinea, Java, Sulawesi, The Solomon Islands,

Vanuatu and New Caledonia and no identical

taxon was discovered.

Accepted for publication 25 August 1998

Habenaria praecox Lavarack et Dockrill, sp.

nov. affinis H. propinguiori Rchb.f. sed

tuberibus multum majoribus (35-50 non 7-15

min longis) et floribus labelli lobis lateralibus

latis et decurvis non tenuibus et sursum in semi-

circulo curvatis differt. Typus: Queensland.

NortH KENNEDY District: about 12 km south

of Cardwell. 18° 23'S, 146°05’E, 6 February

1997, PS. Lavarack PSL 4001 (holo: BRI; iso:

BRI).

Tubers 2, obloid or ellipsoid, up to 50 x

25 mm. Leaves 2-4 basal, or sometimes one

low on the inflorescence stem, more or less

erect and sometimes sheathing at the proximal

end, narrowly oblong to narrowly obovate, 50-

100 x 4-6 mm, canaliculate, acuminate or

acute. Inflorescence 20 to 50 cm tall, with axis

1—2 mm diameter; cauline bracts 2 to 6,

subulate, 6-20 mm long. Rachis 3- to 35-

flowered; flowers moderately dense or rather

sparse and often irregularly arranged; bracts

usually about half the length of the ovary;

pedicels about 1 mm long. Flowers 9-10 mm

across, white with the dorsal sepal often green;

dorsal sepal and petals galeate; lateral sepals

widely spreading; spur on the labellum curved

downwards or directed straight backwards.

Dorsal sepal cucullate-ovate, 3.54 x c, 2.5

mm, slightly constricted near the blunt apex.

Lateral sepals subfalcate, c. 2 mm longer and

1 mm narrower than the dorsal sepal,

constricted at the distal end. Petals broadly

subfalcate, about as long as or slightly longer

332

than, and about the same width as the dorsal

sepal, often with a small lobe on the posterior

side near the base. Labellum cuneate or semi-

circular at the proximal end and then deeply

trilobate, about twice as long as the dorsal

sepal; lateral lobes spreading and directed

downwards and forwards, about as long as the

mid-lobe, narrowly triangular to almost

narrowly oblong, the ends slightly curved

upwards, about the same length as the lateral

sepals and usually about 1 mm broad; mid-

lobe narrowly oblong, slightly tapered, obtuse

or rounded at the apex, 3.5—4 x c.]1 mm; spur

broad at the orifice, constricted at the middle,

dilating near the distal end, with the extreme

distal end constricted, 9-3 mm long. Column

auriculate, suberect, conical or subcylindrical

c.3x3x2mm. Anther emarginate distally;

thecal tubes widely separated at the proximal

end, somewhat converging distally, about half

as long as the stigmatophores and adnate to

them; bursicles not in evidence on any

specimens seen. Pollinia soft, granular and

difficult to remove from anther cells; caudicles

slender, tapered, reticulate, at about a 90° angle

to the pollinia and about two thirds their length.

Stigmatophores adnate only near the base to

the labellum, grooved below. Ovaries twisted

and curved to a varying extent and direction,

9—13 mm long (Fig. 1).

Austrobaileya 5(2): 331-335 (1999)

Additional specimen examined: Queensland. Cook

District: 5 km south of Bamaga, Dec 1976, PS. Lavarack

1070 |AQ193390] (BRD.

Distribution and habitat: Specimens of this

species from near Cardwell and from near

Bamaga on Cape York Peninsula have been

seen for this study. A. praecox (reported as H.

ochroleuca) has also been reported from near

Proserpine, Gordonvale, Julatten, Coen and the

Gulf of Carpentaria (Dockrill 1969).

This species occurs in lowland woodlands

dominated by tea trees (Melaleuca viridiflora)

and also in lowland woodlands comprising

bloodwoods (Corymbia spp) and Eucalyptus

spp. It is confined to areas where the drainage

is poor and the soil stays damp for long periods

during the wet season. The species is most

abundant in areas where the ground cover is

sparse and less than 0.5 m tail. All areas where

this species occurs are subject to a distinct dry

season from June to December. Dry season

fires, which occur about every five years, area

feature of this habitat.

Affinities: This species has been confused with

HT. ochroleuca R.Br. (eg Dockrill 1969 p

38,39), but is readily distinguished from that

species as shown in the following table:

Table 1. Morphological comparison of H. praecox and H. ochroleuca

Distinguishing Characters H. praecox

Lateral lobes of labellum

Labellum spur

Leaves

directed downwards,

about | mm broad

about the same length as the

mid-lobe of the labellum

2 to 4, basal

H. ochroleuca

prominently upturned,

about 3 mm wide

about twice the length of the

mid-lobe of the labellum

reduced to sheathing bracts

along the stem

Laverack, P.S. and Dockrill, A.W., Habenaria Willd. (Orchidaceae) 333

‘a

+ ‘a;

eed at ea

ro»

wT.

+. ie

ate’@ a

+ ok

iol pil

hae

tas ie

; eye i)

gy Pe

a: a

A Th Bid

a” yy Pad

was tna h a 3

' a” =

+S ert

wit, ee Es

HITS, on

o ed

{PLUG}

| gE ; L

= e

= *

4 ris

Fig. 1. Habenaria praecox. A. plant x 0.6. B. flower from the front x 3.6. C. flower from the side « 3.6. D. pollinarium

x 12. E. column from the front x 6. F.dorsal sepal x 3. G. lateral sepal x 3. H. petal (flattened out) x 3. I. longitudinal

section of flower x 6. Del. A.W. Dockrill.

334

While H. praecox is similar to H. ochroleuca

from the Northern Territory and Western

Australia, itis more likely to be confused with

Hf, propinquior Rchb.f. and H. xanthantha

F.Muell., both of which occur in the same

habitat in north Queensland. These three

species have a similar appearance and can

be confused when seen from a distance or when

not in flower. However, the details of the

labellum can be used to distinguish

them. In AH. propinquior, the lateral lobes

of the labellum are filiform and curved

upwards, while they are broader and trend

downwards in H. praecox. Both H. praecox

and H. propinquior have a well developed

spur on the labellum, at least as long as the mid

lobe of the labellum. In . xanthantha the spur

is either absent or, if present, is shorter than the

mid-lobe of the labellum and the lateral lobes

of the labellum are not distinct as they are in

H. praecox and H. propinquior, often being

reduced to very small bumps on the side of the

mid-lobe.

The species illustrated in colour plate 28 on

page 215 of Dockrill (1992) as H. ochroleuca, is

Hi, praecox.

Phenology: H. praecox flowers in December

and January, just as the wet season is

commencing. It flowers before both

H. xanthantha and H. propinquior, with little

overlap of flowering time. There is some

evidence that flowering is related to fire history

and to early season rains in November and

December. In some years, when conditions are

unfavourable few, if any, plants flower.

Conservation status: 4. praecox is not known

to be conserved on any national parks, but

probably occurs on Lumholtz National Park

and Jardine River National Park. Coastal

habitats near Cardwell, north Queensland, have

been greatly reduced by clearing in recent times

and this must have resulted in a reduction in

Austrobaileya 5(2): 331-335 (1999)

population numbers. The exact conservation

status of this species is unclear and it may be

more common than it appears, but at present it

should be regarded as 3R (rare species with a

range greater than 100 km in Australia, but

occutring in small populations which are

mainly restricted to highly specific habitats)

according to the criteria of Thomas and

McDonald (1989).

Etymology:The specific epithet praecox

means “early” and refers to the comparatively

early flowering of this species. This is the first

of the four commonly-occurring species of

Habenaria in the coastal lowlands of the

Queensland’s wet tropics to flower each wet

season.

Acknowledgments

Thanks are due to Len and Kate Lawler of

Atherton for assistance on field trips and in

many other ways. Rod Henderson prepared the

Latin diagnosis. Both Dave Jones and Mark

Clements provided the authors with advice and

compared specimens on their behalf.

References:

Brown, R. (1810). Prodromus Flora Novae Hollandae:

313.

DockritL, A.W. (1969). Australian Indigenous Orchids

Volume 1:38,39. Sydney: Society for Growing Aus-

tralian Plants.

DockritL, A.W. (1992). Australian Indigenous Orchids

Volume 1 (Revised Ed.): 215-217, text & plate

28. Sydney: Surrey Beatty and Sons, in assoc.

with the Society for Growing Australian Plants.

Jones, D.L. (1988). Native Orchids of Australia: 338,339.

Sydney: Reed Books Pty Ltd.

THomas, M.B. & McDoNaLp, W.L.F. (1989). Rare and

Threatened Plants of Queensland. 2nd Edition.

Brisbane: Queensland Department of Primary

Industries.

Laverack, P.S. and Dockrill, A.W., Habenaria Willd. (Orchidaceae) 335

Key to the species of Habenaria Willd. and Peristylus Blume occurring in Queensland

L. (Leaves-arisiig at, of close-to, oround level, oo. ee ck ee ee new be ee oe ee gs 7

Leaves-not-arisine at proud level... x sce ea ace ole we aS bal aire Ace oh OL A 2

2. Lowest leaves several cm above ground level, in the middle of the stem

bceitet eh ditsh wy SPAR Fo pened woh ROP fa ca? So ar 9 EE P. tradescantifolius (Rchb.f.) Kores

Leaves SC IMETCRE PEO EEE SEETTY PoP E scales ave ig En ap ttre ceiaes Rae terepnlahetasahs) anton neikenteta hae eee ees 3

3. Labellum spur shorter than the sepals ............ P. banfieldii (F.M. Bailey) Lavarack

Labelhim-spur longer than the sepals... occ cuca ae ere ie oe Fad Wak Dave a we eye es 4

AEs PC Tea SOLS ts 8a 25 atiet ate wh tode 4 4, bu trey, Merctiniotenysttiga qe eee ial AU cadeaya niger: Muted, 4 pall dh eae ee 5

PELAIS OE ODOR 2s a 5,5 cacy sure ante ¥ Ceol CS b said QU Sek le a ak a PGA be aS Bee 6

Sp BetaSP AUC AS Aon ss yenecgreoceseca gp eae iy eats H. divaricata R.S.Rogers & C.T. White*

Petals not unguiculate (leaves sometimes basal in small plants)

sta Relan aSot boon GARTER oO aNd oa wicrfiaet WSS Withee bd webdles =i Diiedte AY Mh abacage SPS H. rumphii (Brongn.) Lindl.*

6. Anterior lobe of petal slightly longer than or equal to posterior lobe

si daa ic Se: hh ha beh Son, eB a cee wey aw Fore at stenlg Ao GANDA A ts Bah H. macraithii Lavarack

Anterior lobe of petal much shorter than posterior lobe ....... H. hymenophylla Schltr.

7. Flowers horizontal, tubular in proximal half .................006. P. candidus J.J.Sm.

Fhowets-crect- anid Opens a Avidely, genase caw dpe scales ohecenn se £4 og lychee ye eed do ates

8. Labellum spur less than 20 mm long, shorter than, equal to, or up to twice

as long asthe i bellumiobessy. +. «cick ess ene Cha be a eels a ple a ha ea eae eden 9

Labellum spur more than 20 mm long, at least three times as long as the

LADEN IG OES 17.5: ace nok ote ae fdas ann be Aden te vena Ae bea hate a Roah By aN tc nk Bo elem we Aes 12

9, Lateral lobes of labellum filiform, curving strongly upwards; labellum spur

almost twice as long as the mid lobe of the labellum ......... H. propinquior Rchb.f

Lateral lobes of labellum broadly or narrowly triangular, horizontal or

directed downwards oral most absent; labellum spur either almost absent,

or shorter than, or equal to, the mid lobe of the labellum ........ Red aes eee eee 1Q

10. Labellum spur always present and about as long as the mid lobe of the

labellum; lateral labellum lobes well developed and about the same length

ASME LONI Seca es SE ie oo, ey AER welt Ge eA Ig edt od H. praecox Lavarack & Dockrill

Labellum spur variable in size, even within flowers on a single

inflorescence, from absent to very short, or occasionally equal to the

mid lobe of the labellum; lateral lobes of labellum absent, or uneven

in size and shorter than the mid lobe .... 0... cee eee te eens I]

FL -Labeilim longer that! 740m. kd ee vee ee ea ae be H. harroldii D.L.Jones

Labellum shorter than 7mm ...... cee eee eee H. xanthantha F.Muell

12. Lateral lobes of labellum filiform; stigmatophores less than twice the

ISHetOr Te TReCal MESS e5 rae tencecy cane ote ee een avai oh eres abs H. triplonema Schitr.

Lateral lobes of labellum narrow near, often asymetrical; stigmatophores

more than twice the length of the thecal tubes.................00. H. elongata R.Br.

* H. divaricata and H. rumphii are very similar in both plant habit and flower form. No specimen referrable to H.

divaricata has been collected since the type and it is extremely likely that the two species will be shown to be conspecific.

Oldenlandia gibsonii (Rubiaceae: Hedyotideae), a new species from south

east Queensland

D.A. Halford

Summary

Halford, D.A. (1999). Oldenlandia gibsonii (Rubiaceae: Hedyotidieae), a new species from south east

Queensland. Austrobaileya 5(2): 337-339. Oldenlandia gibsonii is described and notes are provided on

its habitat, distribution and conservation status.

Key words: Queensland, Rubiaceae, Oldenlandia gibsonii

D.A, Halford, Queensland Herbarium, Brisbane Botanic Gardens, Mt Coot-tha, Mt-Coot-tha Road, Toowong,

Queensland 4066, Australia

Introduction

This new species of Oldenlandia was first

brought to my attention in 1992 when Norm

Gibson, a keen naturalist from Gladstone,

sent to the Queensland Herbarium for

identification a specimen of this species that

he had collected from Wietalaba State Forest

south west of Gladstone. This first collection

had only old fruit on it but was sufficient to

show that the plant represented either a disjunct

population of O. polyclada (F.Muell.) F.Muell.

from north Queensland or an undescribed

species closely related to O. polyclada. Since

then more material of it has become available

revealing that the plant warrants recognition as

a distinct species,

Oldenlandia gibsonii Halford sp. nov. arcte

affinis O. polycladae (F.Muell.) F.Muell.

autem ramificatione divaricata, foliis

parvioribus (3~7 x 1—2 mm non 7-40 x

2-6 mm), pedicellis brevioribus (1-3

mm non 3—6 mm longis), calycis lobis

parvioribus (0.1—-1.0 mm non 2.0—8.0

mm longis), floribus semper solitariis in

foliorum axillis (non solitariis vel in

fasciculis 2—8 floris) differt. Typus:

Queensland. Port Curtis DIstrRIcT:

State Forest 583 Wietalaba, 24°17’S

151°13’°E, 23 May 1996, PI. Forster

PIF19164 (holo: BRI: iso: BRI, CANB,

Kk, MEL, MO, NSW, QRS distribuendt).

Accepted for publication 12 October 1998

Oldenlandia sp. (Wietalaba N.Gibson

1344) in Henderson (1997).

Woody herbaceous perennial, densely

intricately branched, to 1 m high, with thin,

sreyish, papery bark on mature stems;

branchlets divaricate at c. 80 degrees,

quadrangular when young becoming rounded

with age, hispidulous; axillary branchlets short,

persistent after senescence giving the plant a

spinose appearance. Leaves opposite,

appearing fasciculate on older branches due the

short axillary branchlets, subsessile; lamina

narrowly oblong to oblong-elliptic, 3-7 mm

long, 1—2 mm wide, discolorous, glabrous or

with minute scabrous hairs above, glabrous or

sparsely hispidulous below, attenuate at base,

obtuse, subacute or sometimes mucronulate at

apex, with midrib prominent below and

margins recurved when dry. Stipule sheath c.

1 mm long, hispidulous, produced into

triangular lobe; margin fimbriate. Flowers

solitary in penultimate leaf axils, appearing

terminal by abortion or reduction of terminal

shoot; pedicels slender, 1—3 mm long, glabrous

or hispidulous. Hypanthium subglobose, c. 1.

mm diameter, glabrous or sparsely hispidulous;

calyx lobes broadly triangular, 0.6-1.0 mm

long, obtuse to subacute at apex, joined at the

base into free tube c. 1 mm long; margins

entire. Corolla cream with lilac tinge and faint

lilac blotches in throat, infundibular, glabrous;

tube 4—5 mm long; lobes reflexed, triangular,

4—-5 mm long. Stamens exserted; filaments.

0.5—1.0 mm long; anthers linear-oblong, c. 2

338

mm long. Ovules c. 25-35 per locule. Style

exserted, 6-7 mm long; stigma bifid; lobes

linear, c. 2 mm long, reflexed. Capsule

subglobose, c. 2 mm diameter, glabrous or

subglabrous, not markedly furrowed along

dissepiment; calyx lobes spreading; beak c. 1

mm long, rounded, not protruding above calyx

lobes. Seeds numerous, depressed obconic or

regularly polygonal, c. 0.5 mm long; testa

light brown, reticulate-areolate.

Selected specimens: Queensland. Port Curtis District:

State Forest 583 Wietalaba, 32 km S of Calliope, Jun 1992,

Gibson TOI1315 (BRD); ditto, Nov 1993, Gibson 1343 (BRI,

NSW); ditto, Apr 1994, Gibson 1344 (BRD; State Forest 583

Wietalaba, Dec 1995, Forster PIF 18265, Orford & Tucker

(BRI, MEL, QRS,); ditto, May 1996, Forster PIF19165 (BRD;

Wietalaba State Forest, c. 31 km S of Calliope, Nov 1997,

Halford Q3445 & Snow (BRI).

Distribution and habitat: O. gibsonii is

known only from State Forest 583, 35 km west

of Miriam Vale in central Queensland. It grows

on reddish brown loams itn Araucarian

microphyll vineforest dominated by

Choricarpia subargentea (C.T.White)

L.A.S.Johnson, Backhousia kingii Guymer and

Barklya syringifolia F. Muell.

Phenology: Flowers have been recorded in

May and November; fruits have been recorded

in April, November and December.

Affinities: O. gibsonii is closely related to

O,. polyclada but differs from that by its

divaricate branching, smaller leaves (3—7 x I-—

2 mm compared to 7-40 x 2—6 mm), shorter

pedicels (1-3 mm long compared to 3-6 mm

long), smaller calyx lobes (0.1—1.0 mm long

compared to 2.0-8.0 mm long) and flowers

always being solitary in leaf axils (compared

to flowers solitary or being in 2—8-flowered

fascicles in O. polyclada).

Conservation status: Oldenlandia gibsonii

is known only from the type locality in State

Forest 583 in central Queensland. The total

known population of this species is estimated

to consist of several thousand individuals.

Actions are currently being undertaken to

insure that the conservation of the site is

secured. The recommended conservation

status for this species as defined under the

Queensland Nature Conservation Act 1992 is

Endangered (E).

Austrobaileya 5(2): 337-339 (1999)

Etymology: This species is named in honour

of Norman Gibson, formerly of Gladstone,

who brought this species to my attention.

Acknowledgements

Iam grateful to N. Gibson for bringing this new

species to my attention, P.I. Forster for making

further collections of it, L. Pedley for the Latin

diagnosis and W. Smith for the illustrations.

References

HENDERSON, R.J.F. (ed.) (1997). Queensland Vascular Plants:

Names and Distribution. Brisbane: Queensland De-

partment of Environment.

Oldenlandia gibsonii (Rubiaceae: Hedyotideae), a new species from south east Queensland 339

Fig. 1. Oldenlandia gibsonii. A. branch x 1.B. detail of part ofa branch with short lateral branchlet and flower x 5. C. lateral

view of fruit x 10. A & B, Halford et al. Q3445 (BRD; C, Forster et al, PIF18265 (BRD),

Jasminum jenniae (Oleaceae), a new species from south eastern

Queensland

Wayne K. Harris & Glenn Holmes

Summary

Harris, Wayne K. & Holmes, Glenn (1999). Jasmintm jenniae (Oleaceae), a new species from south

eastern Queensland. Austrobaileya 5(2): 341-344. Jasminum jenniae sp. nov. is described, illustrated and

compared to related species. Notes are provided on its distribution, habitat and conservation status. A

key to the south eastern Queensland species of Jasminui is provided.

Key words: Oleaceae, Jasminiun jenniae

Wayne K. Harris, Department of Botany, The University of Queensland, St. Lucia Queensland, 4072, Australia

Glenn Holmes, P.O. Box 1246 Atherton, Queensland, 4883, Australia

Introduction

The genus Jasminum L. in Australia consists

of eight species and a number of subspecies

and was most recently revised by Green

(1984). The genus is widely distributed

throughout eastern and northern Australia and

two species extend further westwards. J.

calcareum F.Muell. extends from central

Australia through to Western Australia. J.

didymum subsp. lineare (R.Br.) P.S.Green is

the most widespread extending west from the

Great Dividing Range in eastern Australia

through central Australia and into the northern

parts of Western Australia south of the

Kimberley. The species described here has a

limited distribution in south-eastern

Queensland where four species have been

previously enumerated including J jenniae as

an undescribed species (Stanley and Ross

1986).

Jasminum jenniae W.K. Harris & G. Holmes,

sp. nov. affinis J. aennilo var. brassii B.S.

Green autem floribus parvioribus tubo 9—

12mm longo lobis 7—10mm longis;

calyce pedicelloque glabro; foltis

simplicibus oppositis venatione

acrodroma, petiolo versus laminae basem

articulato differt. Typus: Queensland.

Moreton District: Headwaters of south

branch of Kobble Ck, D’Aguilar Range,

27TAT’S 152°46°E; 22 Nov. 1997, S.

Phillips 143: (holo: BRI).

Accepted for publication 26 June 1998

Evergreen shrub to about 4 m, multistemmed,

straggling to prostrate; stem puberulent to

glabrous. Leaves opposite, simple; petioles 3—

5 mm long, sparsely puberulent to glabrous,

articulated near base of lamina, sometimes a

second articulation is present on the upper

third of the petiole; lamina coriaceous, ovate

to broadly ovate, glabrous, glossy, discolorous,

(4.0—-)4.5-6.5(—-8.5) cm long and (1.3—)2.5—

4,.2(-5.8) cm wide; margin entire not

thickened, slightly recurved; apex acute to

acuminate; base attenuate into the petiole;

venation raised above and below, acrodromous

with nerves clearly visible on both surfaces,

the first two or three starting at the base of

the lamina and extending to the apex; other

secondary venation (3—)5-—7 per side and

terminating at the veins originating from the

base of the lamina; base of leaf pairs often

forming a prominent ridge around the stem.

Inflorescence terminal or on axillary side

shoots, subumbellate, (1—) 3(—7)- flowered,

flowers perfumed, bracts linear subulate,

puberulent 3-6 mm long; pedicels 6-18 mm

long, glabrous. Calyx glabrous, tube 2-3 mm

long with 4 or 5 linear subulate teeth 4.5—10

mm long. Corolla hypocrateriform; pale pink

on the outside, paler pink internally, tube 9-

12 mm long with 4—6 lanceolate or narrowly

lanceolate, acute lobes, 7-10 mm long.

Stamens 2; anthers 3~—3.5 mm long, on

filaments 0.5—1 mm long, attached to the upper

third of the corolla tube, slightly exserted.

342

Ovary 0.8—-1.0 mm long; style about 10 mm

long, slightly exserted from the corolla tube,

heterostyly not observed; stigma slightly

bilobed about 1.5 mm long. Fruit spherical to

slightly ovoid, often paired (or single by

abortion), 8-10 mm long by 9—11 mm broad,

purple-black.

Phenology: Flowering period, November to

January, fruits appearing December to April.

Additional specimens examined: Queensland, Moreton

District: Mt Glorious, Jan 1945, Clemens s.n. [AQ47857]

(BRI; Brolga Park, 6km W of Woombye, Dec 1989, Forster

et al.,PIF6146 (BRI); Biack Shoot, Lower Beechmont, Nov

1993, Holmes & Holmes s.n. |AQ633577| (BRD; Tarimgton

Rd, Canungra, Sep 1994, Holines & Holmes s.n. [AQ637677 |

(BRI); South branch of Kobble Creek, Mt Glorious, 27°17’S

{52°46’E, Apr 1997, Phillips 12 (BRD

Distribution and habitat: Jasminum jernniae

has been recorded only from the Blackall,

D’ Aguilar and Beechmont Ranges. This

disjunct distribution extends over 160 km

between latitudes 26°38’ and 28°05’S. Most

records are from the Beechmont Range, in

the upper catchments of Black Shoot, Armitage

Creek, Back Creek (Killarney Falls) and

Clagiraba Creek, between 240m and 475m

altitude. Soil parent material 1s mainly

basaltic or metasediments. In the Beechmont

Range associated canopy species present

at each of four sites included Araucaria

cumninghamii Aiton ex D. Don, Lophostemon

confertus (R.Br.) Peter G. Wilson &

J.T. Waterh., Premna lignum-vitae (A. Cunn.

ex Schauer) W. Piep., and Olea paniculata

R.Br. Plants usually occur near the

boundary between araucarian notophyll

vineforest and eucalypt forest. Plants of

restricted distribution recorded at two of

four sites were Baloghia marmorata C.T.

White, Cryptocarya foetida R.T. Baker,

Rhodamnia dumicola Guymer & Jessup

and Alectryon reticulatus Radlk. (J. Holmes

data).

Diagnostic features: This species is

distinguished readily by its ovate, coriaceous,

glossy leaves with conspicuous and

characteristic leaf venation, its linear subulate

calyx teeth and its generally glabrous

appearance. The petiole is articulated near the

Austrobaileya 5(2): 341-344 (1999)

base of the lamina. Infertile material resembles

Strychnos psilosperma F.Muell. superficially.

Affinities: J. jenniae has close affinities with

J. aemulum var. brassii P.S.Green which

differs by its larger flowers (corolla tube

19-21 mm long, lobes 15-20 mm long),

penninerved leaves, distinct indumentum and

location of articulation in the middle of the

petiole. Green’s (1984) Jasminum sp. is also

very similar and Clemens s.n. [AQ47857]

(BRI) was included in this species by him. The

species is said to have a ‘ fine and minutely, if

thinly, velutinous indumentum on stem,

Woombye’

Canungra?

= Kilometres

Figure 1. Distribution of J. jenniae in south eastern

Queensland

petioles, lower surface of leaf, inflorescence,

and calyx’. Articulation of the petiole occurs

in the middie of or in the lower half. The other

specimens cited by Green are from north

Queensland and are a different species as yet

undescribed. The key in Stanley and Ross

(1986) also includes a Jasminum sp.1 which

is in patt the species described here and it is

clear that they have followed Green’s

treatment and confused the two species. J/.

Jenniae would key out at either J. aemulum

R.Br. or J. sp. in Green’s (1984) key. Several

other species in the Australasian region have

ing

343

” he oF

eet ee ey

mt ga

le leaf show

. sing

mith

. gynoecium x

stag

Se i |

from south eastern Queensland

, a hew species

woe bet ah ™

heneeee’ : ate aw

tx |. B. flower x 4. C. section of flower x 4. D

iae (Oleaceae)

venation x 1.5. F. fruit x 2, (Phillips 143).

A. hab

b td

Pilling,

75 at)

o_ =

bates sinned

wep iy

—_ ma

ne iy,

ssn

Wes

™ wna.

a ee Meal, oi

“eye a a ad ee A al i

et |

= Fl aa

Harris & Holmes, /asi

Figure 2. Jas

NAS ME REE SE Se SS Le eet ease eee pee IS Ss

344

simple leaves. In /. calcareum F. Muell. the

calyx lobes are blunt and the leaves are ovate

to linear- lanceolate and in/. simplicifolium

the leaves are less than 1 cm wide and the

calyx lobes are shorter, usually less than 5

mm long. J. kajewskii has corolla lobes

longer than the tube, in /. longipetalum the

calyx lobes are more or less triangular and

in J. molle the calyx lobes are equal to or

shorter than the tube. J. magnificum

Lingelsh. from Papua New Guinea differs

in having wider calyx segments and short

calyx lobes and much larger flowers and J.

papuasicum Lingelsh. has pinnately veined

leaves. New Caledonian species (Green

1962) with simple leaves are distinct and

differ in the details of the calyx lobes which

are generally shorter.

Austrobaileya 5(2): 341-344 (1999)

Conservation Status: By IUCN (1994) criteria

Bl and C, J. jenniae is Endangered (EN). The

population recorded in the Beechmont Range

comprises about 550 plants. Only a few have

been recorded elsewhere.

Etymology: The species is named after Jenny

Holmes (nee Todd, b. 1952) to recognise her

contribution to the knowledge of the flora of

south east Queensland and northern New South

Wales. During systematic geographic surveys

of vine forests she has collected numerous

significant specimens.

Key to the Jasminum species in S. E. Queensland

1. Leaves simple (1-foliolate) .............

Leaves 3-foliolate ............ cc eee

2. Stems, petioles and undersides of leaves glabrous to finely

puberulent; leaves 2—5 cm wide; calyx lobes 4 or 5, linear-subulate,

AD TTI VOM is ed cles ncaa set Oe egal nen ae

J. jenniae

Stems, petioles and undersides of leaves glabrous or with a few hairs,

leaves less than 1 cm wide; calyx lobes up to 4 mm long,

Pave ly DAN FOMO A Lae aneatdeenecnce le pte le 4"

J. simplicifolium

3. Stems, petioles and leaves pubescent-tomentose; inflorescences mostly

1—5-flowered.........0ccccceuseeeun

Penna ena eng tet No Rates Se Peete | J. dallachil

Stems, petioles and leaves glabrous or minutely puberulent;

inflorescences mostly 5—50-flowered...

Acknowledgements

We thank Dr. G. P. Guymer for provision of

facilities at BRI for one of us. Les Pedley

provided the Latin diagnosis. Sue Phillips made

collections of flowering material of this

species from the type locality for the authors.

Hees: tiralve WeKing x cote Rcd e tenhe he M4: BES as J. didymum

References

Anon (1994), JUCN Red List Categories. International Un-

ion for Conservation and Nature, Gland, Switzerland.

Z|p.

Green, P.S. (1962). Studies in the genus Jasminum, I the

species from New Caledonia and the Loyalty Islands.

Journal of the Arnold Arboretum 43 (2); 109-131.

——— (1984). A revision of Jasminum in Australia.

Allertonia 3(6): 403-438.

Stanley, T.D. and Ross, E.M. (1986). Flora of South-eastern

Queensland Volume 2, Brisbane: Queensland

Department of Primary Industries,.Miscellaneous

Publication QM 84007.

Se eos hha En sie he at ee Eee Eee a

The seedling of Cassytha glabella R.Br.

H. Trevor Clifford

Summary

Clifford, H. Trevor (1999), The seedling of Cassytha glabella R.Br. Austrobaileya 5(2) 345-347.

Cassytha glabella R.Br. is described with special reference to the cotyledons. Seedlings are

cryptocotylar and seed germination is epigeal with the remains of the diaspore covering the tip of the

plumular shoot.

Keywords: Cassytha glabella R.Br.

H. Trevor Clifford, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha

Road, Toowong, Queensland 4066, Australia

Introduction

The seedlings of Cassytha glabella are unusual

in several respects which may help account for

the inaccuracies in the published descriptions

of those of C. melantha R.Br. (Ewart 1919,

1930) and C. paniculata R.Br. (McLuckie &

McKee, as C. pubescens R.Br. 1954), As with

those of other species of Lauraceae, the

seedlings are cryptocotylar but differ from

them in that the remains of the diaspore with

its enclosed seed remnants is usually retained

on the stem apex of the young seedling rather

than being attached laterally to the shoot.

Such epigeal behaviour is characteristic

of many cryptocotylar species and was first

described by Miiller (1887) for Myristica

bicuhyba which he encountered in Brazil.

Furthermore, the cotyledons of Cassytha like

those of many other durian-type seedlings, as

they are often known, separate from the axis

at an early stage of seedling growth (Ng 1978).

The young shoot, with its apex capped by the

remains of the diaspore superficially resembles

a young seedling of A//ium but the presence of

scale leaves on the apparent cotyledon

confirms that it is a stem. Careful inspection

of the stem also reveals the presence of a pair

of opposite scars some distance below the first

scale-leaf. These scars mark the position of the

cotyledonary node.

Fruit and Seed

The diaspore is a superior drupe, embedded in

but free from, a fleshy hypanthtum which

Accepted for publication 14 July 1998

derives from the post anthesis expansion of the

floral receptacle. There is only one seed in each

drupe and this develops from a solitary

pendulous ovule (Sastri 1962). The embryo has

a very short axis and two massive, tightly

appressed plano-convex, peltate (Figs 1C, 1D),

pale-green cotyledons. As with other laurals the

mature seed lacks endosperm (Cronquist 1981).

Seedling

The initial stages of germination are marked

by a slight rupturing of the fruit wall followed

by the emergence and growth of the hypocotyl

to a length of several centimetres with little

concomitant growth of the radicle. From the

base of the hypocotyl, which is somewhat

swollen, there develop several adventitious

roots which anchor the plant in the soil (Figs

1A, 1B). Once the seedling has become

established the plumule expands and after a few

months the roots decay, at which time if the

seedling has not formed haustoria on a suitable

host it dies.

Extension of the plumular bud does not

result in its escape from the seed. Instead, as

the lowermost internodes of the shoot elongate,

its apex remains enclosed between the

cotyledons which are themselves retained

within the remnants of the diaspore.

Due to the abscission of the very narrow

cotyledon petioles at the site of their

attachment to the stem, the remnants of the

diaspore are often carried aloft on the tip of

the shoot (Fig. 1B).

346

The position of the cotyledonary node on

the axis is indicated by the pair of scars which

mark the junction of the hypocotyl and epicotyl

(Figs 1C, 1D).

Rarely, the diaspore wall ruptures

extensively in which circumstance its remnants

may be shed thereby exposing one or both

cotyledons still attached to the seedling (Figs

LA, IC, 1D).

Discussion

The failure of Ewart (1919,1930) to interpret

correctly the structure of Cassytha melantha

R.Br. seedlings is difficult to understand

because Bentham (1870), with whose work he

a — f

i? 9 eS

Icm

Austrobaileya 5(2):; 345-347 (1999)

was quite familiar, gave an excellent description

of the embryo.

Furthermore, Bentham summarised the

earlier literature in which the seed of this

species was initially described as endospermic

but later recognised to be non-endospermic

with massive fleshy cotyledons. In his

description of the family Brown (1810) not only

referred to the cotyledons as large and plano-

convex but noted that they were peltate, and

stressed that his description was the result of

direct observation.

It may be that Ewart (1919) was led into

the error of assuming the seed was endospermic

because he expected that since Cassytha was a

Imm

Fig. 1. Seedlings of Cassytha glabella A & B. Habit sketches; C & D. Details of cotyledonary node. ar, adventitious root;

sl, scale-leaf c, cotyledon; cn, cotyledonary node; f, remains of diaspore; e, epicotyl; h, hypocotyl.

SARI COC SOR cont a rel a one ale eM ees

a ee ee ee eee ee ee ee ee Rg QE, SERN NONEOES FS pce eee id ear Se .

Clifford, H.T, The seedling of Cassytha glabella R.Br.

twining leafless parasite, its embryo like,

those of many other parasitic groups would

lack cotyledons (Cronquist 1981). However,

this explanation does not absolve him from

failing to notice the scars marking the

position of the cotyledonary node on the

seedling axis. Furthermore, his assertion that

the ‘endosperm’ is absorbed by the stem tip

is not supported by his illustration in which

there is no indication that digestion has

occurred even though the seedling is well

developed.

Ewart’s description was soon

challenged by Hart (1925) who recognised

the presence of cotyledons in Cassytha seeds

and suggested that the pair of opposite scars

below the first scale leaf ‘may be the original

points of attachment of the cotyledons’. Ina

later paper (Hart 1946) he returned to the

subject of the morphology of Cassytha

seedlings but did little more than confirm his

original observations.

The scars marking the cotyledonary

node were overlooked by McLuckie and

McKee (1954) who failed to record them on

their otherwise excellent drawings of the

seedlings of Cassytha pubescens, Such an

oversight by two such otherwise careful

observers is difficult to understand especially

as they went on to follow Ewart (I.c.) and

described the embryo of the species as

acotyledonous.

The cotyledon scars were correctly

recognised by Clifford (1987) but as did

Kostermans (1957) and Weber (1981) he

referred to the remnants of the diaspore

covering the plumular axis as a seed. All

three writers thereby lapsed into the common

rather than the technical usage of the term.

The seedlings of all three Cassytha

species studied are similar and differ in only

minor respects from those of other

Lauraceae. Seedling morphology therefore

supports the view of Sastri (1962), based

largely on embryology, that there is no

justification for segregating the genus into a

separate family as proposed by Bartling Gin

Lindley 1833) or subfamily as proposed by

Kostermans (1957).

347

References

BENTHAM, G. (1870). Flora Australiensis 5:308-9.

London: Lovell Reeve & Co.

Brown, R. (1810). Prodromus Florae Novae

Hollandiae et Insulae van Diemen, London:

Johnson & Co.

Cuirrorp, H.T. (1987). Identification of seedlings in

the Australian Flora. In P.Langkamp (ed.),

Germination of Australian Native Plant Seed.

Melbourne: Inkata Press. xii, 236 pp.

Cronauist, A. (1981). An Integrated System of

Classification of Flowering Plants. New York:

Columbia University Press. xviii, 1262 pp.

Ewart, A.J. (14919). Contributions to the Flora of

Australia, No.27. Proceedings of the Royal

Society of Victoria 31(N.S.):367—78, Plate xviii.

Ewart, A.J. (1930). Flora of Victoria. Melbourne:

University Press. 1257 pp.

Hart, T.S. (1925). The Victorian species of Cassytha.

Victorian Naturalist 42:79—83.,

Hart. T.S. (1946). Notes on the tdentification and

erowth of certain dodder-laurels. Victorian

Naturalist 63:12-16.

KosTerMAnNS, A.J.G.H. (1957). Lauraceae. Reimvardtia

4:193-280,

LInDLeyY, J. (1833). Nixus plantarum, Londini: Ridgway

et filios. v, 28 pp.

McLuckig, J. & McKessg, H.S. (1954). Australian and

New Zealand Botany. Sydney: Horowitz

Publications Inc. xx, 758 pp.

MULuer, F, (1887). Keimung der Bicuiba. Berichte der

Deutschen Botanischen Gesellschaft 5:468-—

72.,Taf.23.

Ne, F.S.P. (1978). Strategies of establishment in

Malayan forest trees. In P.B. Tomlinson and

M.H. Zimmermann (eds), Tropical trees as

living systems. Cambridge: Cambridge

University Press,

SASTRI, R.L.N. (1962). Studies in Lauraceae: 3,

Embryology of Cassytha. Botanical Gazette

123:197-206.

Wesker, J.Z. (1981). A taxonomic revision of Cassytha

(Lauraceae) in Australia. Journal of the

Adelaide Botanic Garden 3:187—2672.

Perret arteries stiscisteceon ease not pita es sete tr eee Ce se al hn evade he hi rt er eres arth ted debe told siaeatend pect ah nt eht At

meget OM

* ASSO SE

A checklist of bryophytes of the wallum habitat of south-eastern

Queensland and north-eastern New South Wales

J. Windolf

Summary

Windolf, J (1999). A checklist of bryophytes of the wallum habitat of south-eastern Queensland and

north-eastern New South Wales (Austrobaileya 3(2) 349-352). A descriptive analysis of the

bryophytes occurring within the wallum environment, together with notes on their host/substratum,

micro-habitat and occurrence in the adjacent bryophyte community is provided. Six moss and five

liverwort species are recorded, and the most northerly known occurrence of the moss Sp/hagnut

australe Mitt. is noted.

Keywords: Bryophytes, Wallum

J. Windolf, 53 Pandanus Avenue, Coolum Beach, Queensland 4573, Australia

Introduction

The study reported on here was undertaken to

provide information on the bryophyte species

present in the wallum habitat of south-eastern

Queensland and north-eastern New South

Wales. Any previous studies in this specialist

area have apparently not been reported in the

literature, so it was considered to be a matter

of some urgency, given the rapidly diminishing

state of this unique habitat, that one be

undertaken. Because of its pleasant climate and

its proximity to large centres of population, the

area encompassing the wallum is in increasing

demand both as a tourist and recreational

destination, and as a source of agricultural land,

primarily for the cultivation of sugar cane. The

physical inroads of urban development as well

as of primary production, and their associated

infrastructures such as roads and drainage, have

contributed to severe modifications to the

landform of the wallum via reduction of intact

volume, altered watertables, the introduction

of foreign species and the unnatural frequency

of fire (Harold 1987, Windolf 1987a). Even

those areas which have until now escaped actual

destruction of their original habitats probably

have a limited time in their native state.

What is wallum?

The wallum habitat is widely accepted as a

narrow area of nutrient deficient wet heathland

occurring immediately inland of the coastal

Accepted for publication 22 December 1997

sand-dunes 1n south-eastern Queensland and

north-eastern New South Wales, from

Bundaberg in the north to Ballina in the south,

although limited outliers occur as far north

as Shoalwater Bay and southward to Port

Stevens. It is characterised by shallow, sandy,

acid (pH 3.54.0) soils, particularly deficient

in nitrogen, phosphorous and trace elements,

which, however, support a highly diversified

heathland type vegetation. These soils overlie

an impermeable organic sandstone which

restricts drainage. This results in the watertable,

except in extreme droughts, remaining relatively

high. Variations in altitude are generally less

than a metre, but even these create significant

zonation patterns ranging from the deepest

sections, which are either slow-moving drainage

channels or shallow, closed lagoons, to

relatively dry, open heathland. The surfaces of

the depressions, which are usually covered by

standing water, or at least remain moist

throughout the year, are occupied by the reed-

like sedges, Baumea rubiginosa and Lepironia

articulata, The margins of these areas are often

occupied by dense thickets of stunted (1-2 m

in height) Melaleuca quinquinervia, a species

which also occupies the occasional deeper

stream, but which then tends to adopt a more

normal growth pattern and form narrow belts

of closed forest up to 10 m high. On the sloping

sides of these depressions, there is a graduated

floristic change as the elevation increases,

passing through belts of Leptospermum,

Empodisma, Pultenaea, Xanthorrheoa and

350

Boronia species to the highest level where the

substratum is only seasonally waterlogged and

the vegetation is more characteristic of open

heathland. There it generally includes the

species Banksia aemula, or “wallum” to the

Aborigines, from which the habitat derives its

name (Campbell, Sharpe & Windolf 1995,

Windolf 1987a). This heathland then gradually

changes into sclerophyll forest and shrubland

which inhabits the adjoining sandy ridges.

Climate

The climate of the wallum is of the subtropical

humid, east-coast type characterised by hot

summers, mild winters and a clearly defined

wet season. Annual rainfall varies from 1400

min to 2000 mm, approximately half of which

falls, on average, between the beginning of

January and the end of April, but this is prone

to wide variation, both in timing and magnitude.

A limited number of frosts usually occur during

June, July and August (Windolf 1987a).

Methodology

The research was conducted over a period of

approximately five years and comprised the

collecting of specimens and the recording of

species, as well as the taking of notes on host/

substratum relationships and habitats at various

sites. Similar investigations were also carried

out at the same time in adjacent areas with a

view to determining whether the wallum

bryophyte community was in any way unique,

or exists merely as an extension of bryophyte

communities in adjoining areas.

Collecting was principally carried out in

the Wide Bay and Sunshine Coast regions in

Queensland and around Ballina and Evans

Head in New South Wales. Detailed studies

were conducted in the Noosa, Peregian,

Coolum region with general collecting being

carried out in other areas.

Species list

The following species of bryophytes have been

recorded for the wallum habitats of eastern

Australia. Voucher specimens are housed in the

Queensland Herbarium (BRI), the herbarium

of the Hattori Botanical Laboratory (NICH),

Austrobaileya 5(2): 349-352 (1999)

and the author’s private collection. The

nomenclature and classification systems used

are those published in The Mosses of southern

Australia (Scott, Stone & Rosser 1976) for

mosses, and Australian Liverworts

(Hepaticae): Annotated list of binominals and

checklist of published species with

bibliography (Scott & Bradshaw 1986) for

liverworts.

Mosses

Sphagnaceae

Sphagnum australe Mitt.

This species occurs along the margins of the

wallum depressions, generally mixed with

plants of Lepironia articulata which provide

it with a protected and partially shaded micro-

habitat. It is also recorded along the banks of

water courses and man-made drains

immediately adjacent to wallum areas. As it

appears to have been spread to these sites by

human intervention, it is assumed that the

wallum is its original habitat 1n this region. It

is unknown in Queensland outside of the

wallum and immediately adjacent areas,

although it may be more widespread than this

limited recording suggests.

Ditrichaceae

Ditrichum difficile (Dub.) Fleisch.

This species is found occasionally on bare earth

in the higher, drier areas of the wallum,

particularly on soil which has been compressed,

such as along vehicle tracks. This indicates

introduction by humans, so this species should

probably be seen as a randomly introduced

species in the wallum. It is common in a variety

of habitats throughout eastern Australia.

Dicranaceae

Leucobryum candidum (P.Beauv.) Wils.

This species occurs on dead, decaying timber

and occasionally on bare soil in the drier areas.

It is common in a variety of habitats throughout

eastern Australia.

Campylopus introflexus (Hedw.) Brid.

Windolf, A checklist of bryophytes of the wallum habitat

The habitat of this species is very similar to

that of Ditrichum difficile. It is relatively

common throughout eastern Australia.

Orthotrichaceae

Macromitrium aurescens Hainpe

This species is limited, occurring on the bark

of Casuarina sp. among the Melaleuca thickets.

These thickets, which are often quite dense,

provide a habitat similar to that on the inland

side of the coastal sand-dunes, which they often

adjoin, and where Macromitrium aurescens 1s

also present on the bark of both Casuarina and

Banksia species (Vitt & Ramsey 1985).

Bryaceae

Bryum billardieri Schwaegr.

This species is occasionally found on sand in

areas which are flooded periodically, but dry

out. Itis relatively common in both the wallum

and surrounding areas.

Liverworts

Ricclaceae

Riccia cartilaginosa Steph.

This species is rare, but is found occasionally

on bare soil patches which are periodically

flooded. These patches develop a heavier

texture than is usual in the sandy wallum soils

due to the accumulation of fine silts and organic

detritus in the lowest altitudes. This species has

not been recorded locally from other habitats

outside of the wallum.

Lepidoziaceae

Kurzia compacta (Steph.) Grolle

This species is found only, on peat at the base

and on paper-like absorbent bark on the lower

parts of Melaleuca quinquinervia trees. This

substratum remains damp for long periods

because of the sponge-like effect of drawing

up water from between the tree bases. As these

sites occupy the lowest elevations in the wallum

habitat they remain damp except in periods of

extreme drought. Outside of the wallum, this

species 1s relatively plentiful on dead decaying

351

timber tn dense riverine rainforests and also at

the margins of closed Melaleuca swamps in

adjacent areas (Windolf 1985).

Zoopsis argentea (J.D.Hook. & Tayl.)

J.D.Hook, & Tayl.

The habitat of this species is very similar to

that of Kurzia compacta, but it is a much rarer

species, both in the wallum and in adjacent

areas (Windolf 1985).

Frullaniaceae

Frullania rostrata (J.D.Hook. & Tayl.)

J.D.Hook. & Tayl.

This species occurs only very occasionally on

the bark of several Banksia species, usually in

open shrubland or on isolated plants in the

open. It is also found in a variety of similar

habitats throughout the region, but is never

common.

Lejeuneaceae

Lejeunea flava (Sw.) Nees subsp. orientalis

Schust.

This species is found on the bark of a variety

of hosts, and on dead timber in suitable micro-

habitats. It is very common in a wide range of

habitats in surrounding areas (Windolf 1985),

but is not particularly so in the wallum.

Discussion

It is obvious from the small number of species

recorded that bryophytes play only a minor role

in the overall biology ofthe wallum. However,

in view of the unusual nature of the habitat the

recording of the species and notes on their

occurrence produce an interesting extension of

knowledge in the field of bryophyte ecology,

as well as setting a baseline list of species

present in this threatened environment.

Although small the bryophyte flora

contributes to the overall biodiversity of the

region. Of particular significance is the

presence of Sphagnum australe which was

found only in the wallum and not in adjacent

areas. Although of limited extent, it appears

to be well established and native to this habitat.

352

Its discovery in the Coolum/Peregian area of

south-eastern Queensland is apparently the first

record of this species for Queensland and thus

of its most northerly occurrence in Australia,

normally being associated with the more

temperate regions of Australia, New Zealand,

South America and South Africa (Scott, Stone

& Rosser 1976). With this one exception the

taxa present all occur — albeit in varying degrees

— in adjacent habitats and their presence in the

wallum should be seen as nothing more than a

part of their natural distribution.

Acknowledgments

Acknowledgments are due to Dame Ella

Campbell, Dr Jessica Beever, Dr Helen Ramsey

and the late Dr Sinski Hattori who have all

assisted in the identification of specimens from

wallum habitats for me at some time. Also to

the Chief Botanist and staff of the Queensland

Herbarium for making their collection

available, and to my wife Frances and Philip

Sharpe for their ongoing encouragement and

company in the field.

Austrobaileya 5(2): 349-352 (1999)

References

CAMPBELL, E.O., SHARPE, P. & WINDOLF, J. (1995). The

significance of Kmpodisma minus (Restionaceae)

in mires of eastern Australia, with particular

reference to the coastal marshlands of S.E.

Queensland. New Zealand Botanical Society

Newsletter, Number 42,

Harrop, A. (1987). Heathland regeneration after fire

at Noosa. In L. Johnston, ed., Report of 1987

SGAP State Conference, 9-14.

SCOTT, GEORGE A.M. AND BRADSHAW, J.A. (1986).

Australian Liverworts (Hepaticae): Annotated list

of binomials and checklist of published species

with bibliography. Brunonia. 8(1).

scott, G.A.M., STONE, 1.G. & Rosser, C. (1976). The

mosses Of southern Australia. Academic Press.

56-59.

Vitt, DH. & Ramsey, H.P. (1985). The Macromitrium

complex in Australia (Orthotrichaceae:

Bryopsida). Part 1. J. Hattori Bot, Lab. 59: 325-

451.

Windolf, J. (1985). Survey of the Hepaticae and

Anthocerotae of the Sunshine Coast Region,

Queensland. J. Hattori Bot. Lab. 59: 171-176.

Windolf, J. (1987a). Climate and weather of the wallium

environment with particular emphasis on extreme

conditions and their effects. In L. Johnston, ed.,

Report of 1987 SGAP State Conference, 16-21.

Windolf, J. (1987b). Annotated checklist of Queensland

Hepaticae, Austrobaileya 2(4): 380-400.

oe EERSTE Be ei ee ed ei dS eg A ere bp

Vanguerieae A. Rich. ex Dum. (Rubiaceae) in Australia, 1.

Everistia S.T.Reynolds & R.J.F.Hend.

S. T. Reynolds & R. J. F. Henderson

Summary

Reynolds, S.T. & Henderson, R.J.F. (1999). Vanguerieae A.Rich, ex Dum, (Rubiaceae) in Australia, 1. Everistia

S.T.Reynolds & R.J.F.Hend. Austrobaileya 5(2): 353-361. The Australian representatives of tribe Vanguerieae

have been revised. Three genera, viz. Everistia S.T. Reynolds & R.J.F.Hend. gen. nov. (one species),

Cyclophyllum Hook.f. (9 species) and Psydrax Gaertn. (23 species) occur here. Australian plants formerly

included in Canthium Lam. belong to these genera. In this paper, the following new combinations are

made for some of them: Everistia vacciniifolia (F Muell.) S.T.Reynolds & R.J.F.Hend. (Canthiun

vacciniifoliitum F.Muell.), Everistia vacciniifolia var. nervosa S.T.Reynolds & R.J.F.Hend. and Everistia

vacciniifolia f. crassa S.T.Reynolds & R.J.F.Hend. All recognised taxa of Everistia are described, and

relevant keys, distributional maps and line drawings are provided.

Keywords: Hyverista, Vanguerteae, Rubiaceae, key, taxonomy, Everista vacciniifolia, Australian flora,

Queensland, New South Wales.

S.T. Reynolds, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road, Toowong

Queensland 4066, Australia.

R, J. FE. Henderson, Queensland Herbarium, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Road,

Toowong Queensland 4066, Australia,

Introduction

Recent studies evaluating the genus Canthium

Lam. in Africa have resulted in the splitting of

the genus and reinstatement of some of the

genera previously combined with Canthium.

Bridson (1985) reimstated the genus Psydrax

Gaertn. and included the African species and

one Australian species (P. lamprophylla

(F.Muell.) Bridson) in it. She later (Bridson

1987, p. 616) suggested transfer of several

other species to this genus. She characterised

the New Caledonian genus Cyclophyllum

Hook.f. and discussed it as one of several

groups of Vanguerieae allied to Pyrostria

Comm. ex Juss. noting that “the question of

whether recognition of Cyclophyllum should

be generic or at an infrageneric rank of

Pyrostria (or even Canthium) remains to be

settled”. Bridson (1992) noted that neither

Canthium sensu stricto nor any species

Accepted for publication 13 November 1998

approaching it occurs in Australia, Papuasia or

the Pacific Ocean basin, and that taxa from this

region previously included under the name

Canthium Lam. are referrable to either

Psydrax Gaertn. or Cyclophyllum Hook.f.

This study confirmed that the majority

of the Australian species of Vanguerieae are

referable to Psydrax or Cyclophyllum as they

agree quite well with others of those genera

from outside Australia. An exception is

Canthium vacciniifolium F.Muell. This species

differs from related Australian species by its

deeply 2-lobed, ovoid stigma, which has a

convex base, much-branched habit, its small

usually nerve-less leaves and its 1—3-flowered

umbelliform inflorescences with delicate

flowers. It is therefore treated as belonging to

a distinct new genus described as Everistia in

this account.

354

Taxonomy

Austrobaileya 5(2): 353-361 (1999)

Key to genera of tribe Vanguerieae in Australia

1. Stigma oblongoid, concave at the base; style (usually much) exceeding

the corolla tube; inflorescences usually of branched pedunculate cymes,

usually with secund flowers and a long-stalked central flower near branch

TOTES So Bao paced oSecty 18 OL toes a ks

convex at the

Stigma capitate or ovoid,

base; style as

long as or slightly exceeding the corolla tube; inflorescences of sessile

or short-peduncled umbelliform cymes

2. Stigma ovoid, deeply 2-lobed; cymes pedunculate, 1—3-flowered; corolla

tube delicate, with a band of reflexed filiform hairs at throat; anthers on

distinct filaments, lacking dark coloured connective tissue dorsally;

plants usually much branched (intricately branched); leaves nerveless

or obscurely Nerved.. o4 46 se eS ees

AEE eee PO ane yee eee ee 1. Everistia

Stigma capitate, obscurely 2-lobed; cymes sessile or pedunculate, 1—12-

flowered; corolla tube fleshy, usually with dense long moniliform hairs

projecting from mouth, but lacking reflexed hairs; anthers subsessile,

with dark coloured connective tissue dorsally; plants few-branched;

leaves conspicuously nerved...........

Notes: Only -veristia is treated in this paper;

Cyclophyllum and Psydrax will be dealt with

in forthcoming issues of Austrobaileya.

This study is based mostly on herbartum

material. Measurement ranges given for leaves,

inflorescences, flowers and fruit are based on

dried, fresh or spirit material.

In the citation of specimens, only

herbaria whose specimens have been seen are

listed; districts are provided only for

Queensland collections.

Everistia S.T.Reynolds & R.J.F.Hend. gen.

nov. a Psydrace Gaertn. et Cyclophyllo

Hook.f. stigmate profunde bilobo ad

stylum per basem convexam affixo,

habitu multo ramoso, foliis parvis

plerumque enervibus, floribus delicatis

differt. Typus: Everistia vacciniifolia

(F.Muell.) S.T.Reynolds & R.J.F.Hend.

(Canthium vacciniifolium F.Muell.)

Shrubs or small trees, erect, scandent or

prostrate, usually much branched; branchlets

(especially when young) usually spinose;

spines usually at tips of branchlets or supra-

axillary or leaf opposed, entire or sometimes

forked. Leaves nerveless or obscurely nerved.

Inflorescences umbelliform, with 1—3 flowers

3. Cyclophyllum

in fascicles on short slender peduncles;

pedicels exceedingly slender; flowers 4- or 5-

merous; corolla with short or long tube, with

reflexed hairs at throat; corolla lobes acute to

acuminate and sometimes long attenuated at

apex; stamens exserted; anthers oblongoid,

dorsifixed; style as long as or longer than

corolla tube; stigmatic knob ovoid, fleshy, with

two, broad, + flattened, ovate, recurved lobes,

convex at the base. Fruit ellipsoidal or obovoid,

slightly fleshy, blackish when ripe, 2-lobed;

pyrenes hemispherical, adaxially flattened,

woody, exceedingly rugose abaxially;

cotyledons borne more or less parallel to

ventral face of seed.

Distribution: Endemic in Australia;

represented by one very variable species.

Diagnostic characters: Everistiad 1s

characterised by its much-branched habit (the

divaricate intricately branched branches often

forming an entangled mass), slender branchlets

which are often spinose, obscurely nerved or

nerveless leaves, fragile 1—3-flowered

umbelliform inflorescences, delicate 4- or 5-

merous flowers and deeply 2-lobed stigma.

Affinities: Everistia resembles Psydrax

Gaertn. and Cyclophyllum Hook.f. in the

placement of the cotyledons and is closely

Vanguerteae A. Rich. ex Dum. (Rubiaceae) in Australia, 1. Everistia S.T.Reynolds & R.J.F.Hend., 355

related to these genera. It resembles Psydrax

in its corolla (with a band of reflexed hairs at

the throat of the corolla tube), exserted erect

stamens and umbelliform inflorescences. This

type of inflorescence although not known in

the species of Psydrax in Australia, is present

in some African species of that genus (Bridson

1985). It resembles Cyclophyllum in its

umbelliform inflorescence and stigma with

convex base.

Etymology: This genus is named in honour of

the late Dr Selwyn L. Everist, a former Director

of the Queensland Herbarium (1954-1976),

who gave us the opportunity to work in our

particular field of botanical interest and who

supported and encouraged many a botanist

during his directorship at BRI. Dr Gordon

Guymer, current Chief Botanist at BRI, is

thanked for suggesting this name.

1. Everistia vacciniifolia (F.Muell.)S.T.

Reynolds & R.J.F.Hend., comb. nov.

Canthium vaccintiifolium F.Muell., Trans.

Phil. Inst. Vict. 3: 47 (1859). Type:

Queensland. SouTH KENNEDY DISTRICT:

Suttor River, &’ Mueller s.n. (lecto, here des-

ignated: MEL [MEL 1538572}; isolecto: K).

Shrubs or small trees 1-7 m high, erect to

scandent to prostrate; bark grey or creamy

grey, smooth; trunk straight with spreading

uniplanar or divaricately branched branches;

branches and branchlets often intricately

branched and forming an entangled mass giving

the plant a rounded appearance; branchlets

greyish-brown and with white streaks, glabrous

or hairy with minute, spreading hairs, terete,

slender, stiff, straight or sometimes flexuose

or curved (sometimes recurved), spinose

distally especially in young growth, with few

scattered leaves or occasionally leaf-less and

spinescent; spines bifurcate or entire, straight,

often thickened at base, at tip of short leafless

branchlets, leaf-opposed or supra-axillary; leaf

lamina surfaces, stipules, petioles, peduncles,

pedicels and calyx with minute spreading hairs

or glabrous. Petioles 0.5-2.0 mm long;

stipules comparatively small, ovate, abruptly

acuminate with a short folded lobe at apex,

thick or thin, coriaceous; leaf laminas elliptic,

obovate or suborbicular, 3.5-17.0 x 3.5—10.0

mm, obtuse or retuse at apex, obtuse, + truncate

or acute at base, flat or recurved at margins,

flat or slightly convex or concave and broadly

channelled at midrib adaxially, thick or thin,

coriaceous; adaxial surface shiny; abaxial

surface dull: midrib slender, sometimes not

apparent; lateral nerves in | or 2 pairs, obscure,

very fine, arching, usually visible on abaxial

surface, or nerves absent. Peduncles 0.5—5.0

mm long; bracts minute, ovate. Flower buds

4,0-7.0 (-9.0) mm long, slender, sometimes

fleshy, with apex obtuse and apiculate or

acuminate; pedicels 0.5—5.0 mm long; calyx

1.0-1.25 mm long; limb 4 (or 5)-denticulate

with minute ovate lobes; corolla yellow, 5.0—

9.0 mm long; tube slender, dilated or inflated

at throat, 3.0-4.5 mm long, c. 2.5 mm wide at

mouth, sparsely hairy with a band of reflexed

filiform hairs at throat; lobes erect or recurved,

ovate or lanceolate, 2.5-5.0 x 1.0-2.0 mm,

with apex acute or subobtuse and apiculate or

subacuminate or acuminate and usually with a

long recurved acumen, hairy, often papillose

near apex; stamens exserted; filaments 0.75-—

2.5 mm long; anthers oblongoid, 1.0—2.5 mm

long; style with stigma 4—8 mm long, exserted

or included. Fruit 4.0—7.0 x 4.0—7.0 mm,

smooth, usually topped by remnants of calyx.

Notes: FE. vacciniifolia is readily

distinguishable by its much-branched habit,

stiff, slender, often spinose branchlets, straight

unbranched or bifurcate spines, small, elliptic

or obovate, faintly nerved or nerveless leaves,

small, fragile, shortly pedunculate 1-3-

flowered umbelliform inflorescences, and

obtuse- to acuminate-tipped flower buds.

The species as circumscribed here is extremely

variable in its aspect, branching, shape and

texture of leaf laminas and in the shape of its

flower buds. Two varieties are recognisable;

these are connected by intermediate forms.

356

Austrobaileya 5(2): 353-361 (1999)

Key to varieties of Everistia vaccinitfolia

1. Leaves 3.5—9.0 x 3.5—8.0 mm; lamina slightly concave or flat, coriaceous,

thick; lateral nerves not apparent; flower buds 4.0—7.0 mm long, obtuse

or abruptly narrowed and apiculate, or rarely subacuminate at apex;

peduncles and pedicels 0.5—2.0 mm long; usually much-branched

shrubs or small trees with divaricate branching .. (a) E. vacciniifolia var. vacciniifolia

Leaves 12.5-17.0 x 9.0-10.0 mm; lamina flat, coriaceous, thin (drying

very thin); lateral nerves distinct; flower buds 7.5—9.0 mm long, long

acuminate at apex; peduncles and pedicels 2.0-5.0 mm long; usually

small trees with planar layered branching

(a) E. vacciniifolia var. vacciniifolia

Shrubs, erect or prostrate, usually much

branched with divaricate branched branchlets;

branchlets densely leafy and straight or

recurved distally, or with a few scattered leaves

and spinose, or sometimes leafless when short

and spinescent. Leaf laminas flat or slightly

concave, with flat or recurved margins, thick,

coriaceous, nerveless or rarely obscurely

nerved, with surfaces minutely hairy or

glabrous. Peduncles slender, 0.5—2.0 mm long;

cP neacee Saar (b) E. vacciniifolia var. nervosa

flower buds 4.0—7.0 mm long, slightly fleshy,

obtuse and apiculate or abruptly narrowed and

crowned by minute lobes at apex; corolla tube

slender, slightly dilated or inflated at throat;

lobes obtuse, apiculate or sometimes

subacuminate.

This variety varies greatly in the shape

and texture of its leaf laminas. Two forms of it

are formally recognised here. These forms

occasionally overlap in their characters.

Key to forms of Everistia vaccinitfolia

1. Leaf laminas flat or slightly concave or deeply grooved along the middle

adaxially; margins flat or recurved; adaxial surface slightly shiny, hairy

or glabrous; nerves absent.............

... G) E. vacciniifolia forma vacciniifolia

Leaf laminas flat; margins flat; adaxial surface shiny, glabrous; nerves

obscure or absent ..........ccceuv vee

(i) E. vacciniifolia f. vacciniifolia

Shrubs with divaricate branching; branches

often much branched and forming an entangled

mass; branchlets distally straight or curved;

leaf laminas with nerves not apparent, minutely

hairy or glabrous. Flower buds 4.0—5.5 mm

long, usually obtuse and shortly apiculate at

apex; corolla tube 2.5—3.0 mm long, slightly

dilated at throat; lobes 2.5-3.0 x 1.0—-1.5 mm,

subobtuse or acute. (Fig. 1A)

Representative specimens: Queensland. LEICHHARDT

District: Junee Tableland, 80 km N of Dingo, Jun 1972, -

McDonald 554 (BRI); Katuroo, 23°05’S, 149°18’E, Feb 1993,

Fensham 713 (BRI); Murphy Range, 57 km from Taroom on

Glenhaughton road, 25°25’S, 149°30’E, Sep 1992, Forster

PIF 11226 & Sharpe (BRD; ditto, Oct 1993, Forster PIF 14111

& Holland (BRI). Port Curmis District: Fitzroy River, May

1863, Dallachy s.n. (MEL); Double Head Yeppoon, Emu

Puiad ehetee Gi) E. vacciniifolia forma crassa

Park, May 1970, 7elford 1700 (CANB). Burnetr District:

Monogorilby, Mundubbera Shire, 26°01’S, 151°O1°E, Dec

1984, Forster PIF 344B (BRI). DARLING Downs District:

Chinchiila, May 1946, White 1122 (BRI, CANB). MorEToN

District: Brisbane River, Hil/s.n. [MEL 15385711] (MEL).

Distribution and habitat: Central and

southern Queensland, between Suttor and

Brisbane Rivers; usually tn dry scrubs

especially in or at the edge of Brigalow (Acacia

harpophylla F.Muell. ex Benth.) scrubs, on dry

ridges, Jump-ups, slopes and gullies, on sandy

stony soils. Map 1.

Notes: This form varies greatly in its aspect,

branching, leaves and shape of flower buds.

Specimens from along the Fitzroy, Dawson,

Burnett and Brisbane Rivers usually have

robust stiff branchlets, which are mostly

arching especially towards their tips or are

ce tea ye petsbst Se es taba oa oa any T GLE tt A gh tae eee ie i ‘aeanlytelad fisted Meco ecag dee cb

Vanguerieae A, Rich. ex Dum. (Rubiaceae) in Australia, 1. EveristiaS.T.Reynolds & R.J.F.Hend. 357

short and sometimes spinescent, small elliptic

or orbicular, + concave leaf laminas with

usually recurved margins, shortly stalked

inflorescences and obtuse and apiculate flower

buds. Specimens from other areas, however,

have thicker, narrow or broad, flat leaf laminas

and usually more slender and longer flower

buds; some of these approach F. vacciniifolia

f, crassa, and probably represent intergrades

between the forms.

Typification: Mueller (1859) cited “Barren

scrubby localities near the Burdekin, Suttor,

McKenzie, Dawson and Burnett rivers” in his

protologue of Canthium vaccintifolium. Only

one of the collections on which Mueller

apparently based his description, viz. Suttor

River, leg, Mueller sn. UX, MEL), has been

seen in this study. The MEL specimen of this

collection is here chosen as lectotype of this

species’ name. The specimen, which is in

flower, agrees well with Mueller’s protologue

description.

(ii) E. vacciniifolia f. crassa S.T.Reynolds

& R.J.F. Hend., forma nov. a #.

vaccinilfolia (F. Muell.) S.T.Reynolds &

R.J.F.Hend. f. vacciniifolia laminis

foliorum crassis differt. Typus: Queens-

land. Port Curtis District: Neerkol

Creek, Bowman 21 (holo: MEL [MEL

1538080]).

Shrubs with divaricate or planar layered

branching; branchlets rarely arching distally;

leaf laminas with nerves and midrib not

apparent or rarely visible but obscure, glabrous.

Flower buds 5.5—7.0 mm long, abruptly

narrowed at apex and crowned by short narrow

lobes; corolla tube 3.5-4.5 mm long, usually

inflated at throat; lobes 2.5~3.0 x c.1.5 mm,

subacuminate.

Representative specimens: Queensland. Cook District: 40

Mile Scrub, 18°15’S, 141°45’E, Feb 1972, Hyland 5874 (BRD.

Burke District: Mt Walker, near Hughenden, 20°5-’S, 144°1-

'E, Apr 1935, Blake 8442 (AD). NortH KENNEDY DISTRICT:

Mingela Bluff, 19°53’S, 146°45’E, Jan 1992, Forster PIF 9416

& Bean (BRI). MircHe ty District: Cuttsy’s Springs, about

30 miles (48 km) ESE of Yalleroi, Feb 1940, Everist 1965 (BRD;

Jericho, Mar 1920, Francis 106 (BRI). SouTH KENNEDY

District: Near Glendon, Sep 1950, Smith 4625 (BRI); 7 km

NE of Belyando Crossing on Gregory Development Road,

21°30’S, 146°48°E, Jun 1992, Thompson 460 & Sharpe (BRI).

Port Curtis District: Gainsford, date unknown, Bowman

s.n. (MEL) (see below).

Distribution and habitat: Northern and

central western Queensland; in deciduous vine

thickets on rocky hillsides and on sandstone.

Map I.

Notes: This form is characterised by its flat,

thick leaf laminas with slightly shiny adaxial

surfaces, long flower buds and subacuminate

corolla lobes. It resembles &. vacciniifolia var.

nervosa in its planar layered branching, flat leaf

laminas and long flower buds and corolla, but

differs from that by its stout branchlets with

closely arranged leaves, thick, usually

nerveless leaves, and comparatively short

peduncles and pedicels. E. vacciniifolia var.

nervosa is a sparsely branched plant with

leaves usually widely spaced on the branchlet,

leaves thin and distinctly nerved, flower buds

slender and usually slightly fusiform, and

peduncles and pedicels comparatively long.

The label attached to Bowman’s

unnumbered specimen of this taxon in MEL

[MEL 1538079] is annotated ‘Gainsford’,

whereas the label attached to the specimen is

annotated ‘Herbert Creek’. This specimen

appears to be from the same collection as a

specimen in MEL labelled as collected at

Herbert Creek by Bowman [MEL 1538069].

Both Gainsford and Herbert Creek are close

to Neerkol Creek (Blake 1955), where the

holotype was collected.

Etymology: The epithet crassa, Latin for thick,

refers to the compartively thick leaf laminas

in this form.

(b). E. vacciniifolia var. nervosa S.T.Reynolds

& R.J.F.Hend., nom. & stat. nov.

Canthium microphyllum F.Muell., Fragm.

2: 134 (1861). Type: Queensland.

Moreton District: Brisbane River,

Moggill Scrub, # Mueller s.n. (lecto,

here designated: MEL [MEL 1538076];

isolecto: K).

- Small trees or large shrubs, usually with a

straight slender trunk and spreading layered

branching; branchlets more or less borne in the

same plane as the main branches or

occasionally suberect with divaricate

branches. Leaves usually widely spaced on

358

branchlets; laminas flat, glabrous, coriaceous,

thin (drying very thin), slightly shiny on adaxial

surface; midrib and nerves usually visible on

the abaxial surface; lateral nerves in 1 or 2 pairs,

very slender, slightly arching. Peduncles and

pedicels filiform; flower buds fragile, 7.5—9.0

mm long, with a slender tube and acuminate

lobes at apex; corolla tube slender, 3.5—4.5 mm

long; lobes lanceolate, 3.5—5.0 mm long, acuminate

with a long recurved acumen at apex. (Fig. 1B)

Representative specimens: Queensland, Port Curtis

District: Dan Dan Scrub, State Forest 53, Calliope Shire,

Dec 1984, Gibson 699 (BRI). WibE Bay District: NW base

of Boogooramunya, State Forest 648, 25°51’S, 152°08’E, Jan

1989, Forster PIF4913 (BRI); Mt Eerwah, about 4 km W of

Eurnoundi on Kenilworth road, 25°28’S, 152°55’E, Dec 1987,

Sharpe 4759 (BRD; 1 kmS ofbarracks, Oakview State Forest

220, 26°07’S, 152°20°E, Dec 1988, Forster PIF4861 & Orford

(BRI). Burnett District: Yarraman State Forest (State Forest

289), Neumgna, 26°56’S,151°49’E, King Logging Area, just

N of junction of Cooyar-Maidenweil road and New England

Highway, Dec 1987, McDonald 4135 & Williams (BRI).

Moreton District: McIntyre Scrub, 6 km W of Woombye,

26°39’S, 152°54’E, Jan 1990, Forster PIF6204, Bird & Bean

(BRI); Woongaroo Creek, S of Goodna, 27°40’S, 152°45’E,

Nov 1983, Williams 83021 (BRD; Moggill, Brisbane, 27°3-

’S, 152°5-’E, Feb 1984, Oakinan s.n. (BRI); Norman’s Creek,

Moreton Bay, Jui 1843, Leichhardt s.n. [NSW 193686]

(NSW); MtFrench, 6 km SW of Boonah, 28°02’S, 152°40’E,

Jan 1983, Telford9079 & Butler (CANB). New South Wales.

Totties Mount, Ramornie, Jul 1922, Blakely & Shiress s.n.

INSW 193685 ]{NSW); Pikapene State Forest, about 12 miles

(19 km) directly SE of Tabuian, Nov 1966, Hayes, Turner &

McGillivray 2649 (BRD); ditto, Apr 1969, Pickard & Blaxell

255 (NSW); MacLeay River, date unknown, Becker s.n.

[MEL 1538073 ](MEL); Broken Bago State Forest, about 9

km SW of Wauchope, Nov 1980, Covery 10922 (NSW).

Distribution and habitat: Central and

southeastern Queensland to Hunter River, New

South Wales; usually in Araucarian microphyll

vine forests, on reddish coloured soils; along

creeks and rivers, and on slopes. Map 1.

Notes: E. vacciniifolia var. nervosa is readily

recognisable by its usually layered branching

spreading out from a single stem, finely nerved

leaf laminas, leaves widely spaced on

branchlets, and by its long, fragile, acuminate

flower buds and acuminate corolla lobes.

The leaves of this variety are variable.

Specimens with thin, finely nerved leaves are

typical whereas specimens with comparatively

thicker leaves and obscure nerves approach £.

vacciniifolia f£. crassa. The latter taxon, however,

Austrobaileya 5(2): 353-361 (1999)

differs from them in its shorter peduncles and

pedicels which are 0.5—-2.0 mm long, and

obtuse- or acute-tipped flower buds.

Canthium microphyllum F.Muell. was

combined with C. vacciniifolium F.Muell. by

both Bentham (1867) and Mueller (1875)

without any formal infraspecific recognition.

It is recognised at varietal level here because

although the taxa intergrade, their extremes are

very distinctive.

Typification: Mueller (1861) cited three

collections, viz. Brisbane River, Hil/ s.n., ditto,

Mueller sn., and Rockhampton, Thozet s.n.,

with his diagnosis of Canthium microphyllum.

Of these, Mueller’s collection from Brisbane

River, annotated ‘Moggill Scrub, Brisbane

River’, at MEL [MEL 1538076] is here chosen

as lectotype of this name. The lectotype, which

is in flower, agrees well with the original

description.

The syntype from Brisbane River collected by

Hill [MEL 1538571] 1s &. vacciniifolia var.

vacciniifolia, whereas the syntype from

Rockhampton collected by Thozet s.n. [MEL

1538078] 1s also E. vaccintifolia var. nervosa.

Etymology: The new epithet nervosa, Latin for

nerved, referring to the usual obscurely nerved

leaf laminas, is proposed rather than making a

new combination from Mueller’s Canthium

microphyllum because the leaves of this

variety are usually much bigger than those of

E. vacciniifolia var. vacciniifolia.

Unplaced specimens of &. vaccinitfolta

Canthium sp. (Massy Creek P.J.Forster +PIF

10568), (Reynolds 1997, p. 180).

The collections listed below probably

represent another variety or form of this

species, but are insufficient to be sure. They

resemble £. vacciniifolia var. vacciniifolia in

their short peduncles and pedicels, and £.

vacciniifolia var. nervosa in their long flower

buds with pointed apices. However, they have

a different aspect from both these varieties in

attributes of their branchlets, their subobovate

or elliptic leaf laminas and the shape of their

flower buds. As only three of the collections

ee ee reste bede eae eee:

A ee ee

Baath at seg tl eile Loa deh a kg bee eke eile be ane EE Bot cea eek aa

UE LT A

nememeine.

Vanguerieae A. Rich, ex Dum. (Rubiaceae) in Australia, 1. Hveristia S.T. Reynolds & R.J.F.Hend. 359

Fig, 1. A-E. Everistia vacciniifolia vay. vacciniifolia., A. juventie branch x 1.5. B. portion of flowering branch * 2. C. flower

x 6. D. longitudinal section of flower x 6. E. fruit x 6. (A from Myers s.n. (BRI [AQ 124017]); B- D from Forster PIF 14111 (BRD;

E from Forster PIF9097 (BRD). F-J. &. vacciniifolia var. nervosa. F. flowering branch x 0.8. G. detail of leaves and

inflorescence x 3.H. flower x 6, I. longitudinal section of fruit showing embryos x 6. J, abaxial view of pyrene x 9. (F from

Lebler 1978, p.530; G from Forster PIF12062 (BRD; H from Forster PIFA919 (BRI); IJ from Forster PIF3750 & Bolton(BRD).

360

have a few depauperate flowers on them, and

their leaves are also variable, it is not possible

to be certain that they represent a taxon

constantly different from the named varieties.

If they do represent a distinct taxon, based on

specimens available, it may be described as

follows.

Shrubs 2—3 m high; branching spreading,

flattened; branchlets, peduncles and pedicels

finely hairy to glabrous; spines straight, leaf

opposed or at apex of small branchlets, usually

forked. Petioles 1-2 mm long; stipules small,

ovate, apiculate; leaf laminas elliptic to

subobovate, 5.59.0 (-18.0) x 3.5—9.0 (—16.0)

mim, obtuse or retuse at apex, subacute at base,

flat or slightly recurved at margins, + thin,

coriaceous; adaxial surface shiny or with a

slight sheen; midrib and 1 pair of lateral nerves

usually distinct. Inflorescences 1—3-flowered;

peduncles and pedicels 0.5—1.5 mm long;

flower buds narrowly ellipsoid to narrowly

obovoid, 5.5—7.5 mm long, with minute

slender, acuminate, hairy lobes at apex; calyx

indistinctly lobed, c.1 mm long; corolla 6.0—

7.0 mm long; tube cylindrical, 3.5-5.5 mm

long, c. 2.5 mm wide at mouth, densely reflexed

hairy at throat; lobes 2.5-3.5 x c.1.25 mm,

subacuminate; stamen filaments 0.5-1.5 mm

long; anthers 1.0—1.5 mm long; style with stigma c.

3 mm long. Fruit ellipsoid, c. 5.0 x c. 5.5 mm.

Specimens studied: Queensland. Cook DistricT: Goode

Island, in 1881, Powe// 26 [MEL 1538045] (MEL); Caimncross

Island, Aug 1855, collector unknown [MEL 1538072] (MEL);

Moa Island, Thomas Swamp, Aug 1985, Budworth 355

(BRI); Moa Island, Apr 1987, Budworth 973 (BRD; Galloways

Hill, 10°46’S, 142°28°E, Sep 1991, Sarkowsky & Sankowsky

1234 (BRD; Weipa Rd, 83 km W of Cape York Development

Rd, 13°14’S, 142°41°E, Jul 1984, Puttock & King UNSW

16975 (BRD; 5.5 km W of Lockhardt River turn off on

Portlands Rd, 12°44’S, 143°1S’E, Jul 1984, Puttock & King

16791 (BRI, UNSW); 6 km W of Rocky River mouth, 36.2 km

ENE of Coen, Silver Plains Holding, Cape York Peninsula,

13°46’S, 143°29°E, Aug 1993, Fe// 3480 (BRD); 5 miles (8 km)

N of Crossing on Massy Creek Rd, below Silver Plains

Station and Rocky River, Oct 1969, Webb & Tracey 9719

(BRD); 3 km N of Massy Creek Crossing, Silver Plains Station,

13°53’S, 143°31°B, Jun 1992, Forster PIF10574, Sankowsky

& Tucker (BRI); ditto, Forster PIF10568, Sankowsky &

Tucker (BRD).

Distribution and habitat: Cape York

Peninsula, Queensland, including the Torres

Strait islands; usually in evergreen vine

thickets, on sandy beach ridges on fine white

sand. Map 1.

Austrobaileya 5(2): 353-361 (1999)

Notes: The leaf laminas in the above specimens

are quite variable, being elliptic, thin,

coriaceous and with slightly recurved margins

in those from Massy Creek, subobovate, flat

and thin in the Torres Strait Islands specimens,

and flat, thick and coriaceous in the remainder.

However, in their overall aspect and colour of

branchlets and indumentum, the specimens

appear to be from the same taxon and are,

therefore, tentatively treated together.

Examination of more specimens from the

above areas 1s necessary before the variability

and affinities of these collections can be fully

assessed.

Acknowledgements

We thank colleagues at BRI especially Paul

Forster for collecting specimens of many

‘Canthium’ species and for their observations

on the habitat of the species, Will Smith for

the illustrations and maps, Directors/Keepers

of AD, BM, CANB, CGE, DNA, K, L, MEL,

NSW, P, PERTH, QRS, and UNSW for allowing

me (STR) full access to specimens in their

institutions and for the loan of herbarium

material, and The Australian Biological

Resource Study, Federal Department of Arts,

Science, Sports, The Environment, Tourism and

Territories for a grant (to STR) to undertake

research in the genus Canthium in Australia.

Vanguerieae A. Rich. ex Dum. (Rubiaceae) in Australia, 1. Everistia S.T.Reynolds & R.J.F.Hend. 36]

References: 145 150 145 150

aan wiz

THA OS Se

aie =Aan LETT

p fF qlg rae HEC Err 10

d A

BENTHAM, G., (1867). Canthium, in Flora Australiensis 3:420- 4 OE eRe

423, London: Lovell Reeve & Co.

BLAKE, S.T. (1955). Some Pioneers in Piant Exploration and

Classification. Proceedings of the Royal Society of

Queensland 66: 1-19.

Bripson, D.M. (1985). The reinstatement of Psydrax

(Rubiaceae subfamily Cinchonoideae tribe

Vanguierieae) and a revision of the African spe-

cies. Kew Bulletin 40(4):687-725.

———(1987). Studies in African Rubiaceae-Vangueriede: a

new circumscription of Pyrostria and a new

subgenus, Canthium subgenus Bullockia. Kew

Bulletin 42(3): 61 1-639.

— (1992). The genus Canthium (Rubiaceae-

Vanguerieae) in Tropical Africa. Kew Bulletin

47(3):353401.

LEBLER, B.A. (1978). The Canthiums of Southeastern

Queensland. Oueensland Agricultural Journal 104

(6):527- 532. Brisbane: Government Printer.

PRET RHE ET FP

MuELLer, F. (1859). Transactions of the Philosophical enim PH

Institute of Victoria 3:47.

145 150 155

—— (1861). Fragmenta phytographiae Australiae 2:134.

——— (1875). Fragmenta phytographiae Australiae 9:186.

REYNOLDS, S.T. (1997). Canthium, 1n R.J, Henderson (ed.),

Queensland Plants: Names and Distribution p. 180- Map 1. Everistia vacciniifolia f. vacciniifolia a,

181. Brisbane: Queensland Herbarium, Department Everistia vacciniifoliaf. crassa x, Everistia vacciniifolia

of Environment. Var. Nervosa @,

Note

Contribution of isozymic analysis in differentiating Macrozamia moorei

D.L.Jones and K.D.Hill from M.johnsonti F.Muell (Zamiaceae)

Macrozamia johnsonii D.L. Jones & K.D. Hill

was recently segregated from M. moorei

F.Muell. (Jones and Hill, 1992) based on

morphological characters, in particular,

differences in the seedlings and plant habit.

Populations of the two species are also disjunct,

being separated geographically by

a distance of about 800 km with M moorei

found in the central highlands of Queensland

around Carnarvon Gorge and Springsure and

M. johnsonii west of Grafton in north-eastern

NSW. While the two species are

morphologically distinct they are nevertheless

closely allied and may represent sister taxa. The

opportunity is taken here to compare enzyme

banding patterns of both species using isozyme

analysis.

Starch gel electrophoresis was

employed to determine the enzyme pattern

differences between two populations of

M. moorei (Mt Zamia Environmental Park,

Forster, PIF 14081, n=14; Staircase Range,

Forster and Machin, PIF 9766, n=14 and

one population of M. johnsonii (2.5km west

of Bobtail Road, P Machin s.n. (AQ540376),

n=14). Both species have a narrow geographic

range. Samples were collected from

population extremes of each species and

the voucher specimens housed at the

Queensland Herbarium (BRI). When an

extract of leaflet was assayed, following the

method of Wendel and Weeden (1989),

different but consistently reproducible

electrophoretic banding patterns were

produced (Fig. 1) for four of the eleven

anodal enzyme systems assayed namely:

Diaphorase (DIA, E.C 1.6.4.3), Menadione

Reductase (MR, E.C 1.6.9.92), Uridine

Diphosphogluconic Pyrophosphatase (UDP,

E.C 2.7.7.9) and Isocitrate Dehydrogenase

(IDH, E.C 1.1.1.42).

Accepted for publication 17 February 1998

DIA: This monomeric enzyme exhibited a

single band at locus one in all sampled

plants of M. moorei, whereas the locus

was absent in samples of M. johnsonii.

At locus two M. johnsonti samples were

monomorphic, but the locus was

polymorphic in M4 moorei with several

of the plants examined being

heterozygotes for the slower allelic

variant.

MR: The two loci for this enzyme in samples

of M. moorei and M. johnsonii revealed

totally different positions and are

presumed to be fixed for alternative

alleles.

UDP: There were only two heterozygotes

observed in samples of MZ. moorei with

this monomeric enzyme (frequency of

allele one =.07), the remaining samples

being homozygotes at position two,

whereas allele two was absent in samples

of M. johnsonii, all being fixed at position

one.

IDH: This dimeric enzyme produced only two

types of homozygote zymograms. [n all

the samples of 14. moorei examined, the

locus was monomorphic at position one

whereas in samples of M. johnsonii it was

also monomorphic but at position two.

The absence of locus 1 in enzyme DJA for

M. johnsonii and fixation of loci for alternate

alleles in MR and IDH 1s deserving of further

comment. There are three possibilities for these

results to occur namely: (a) the same locus is

present in both species but is fixed for

alternative alleles at a different location (IDH,

MR), (b) locus one for an enzyme DIA might

be present but is fixed for allele overlapping

locus 2 or a null allele or (c) there could have

been two loci at IDH and MR inM. moorei and

M. johnsonii, two loci at DIA in M. johnsonii,

but one of them is presumed to have been lost

during the course of evolution.

364

® M. moore

Eevee i —

Hee Teh ee ae a ae a ae

DIA

M. moore!

Austrobaileya 5(2): 363-365 (1999)

M, johnsonii

Loc. 1 absent

M. johnsonit

® M. moorei

20 +

M. moorei

IDH

M. johnsonii

M. johnson

Ey eee ee a PP LEA el Sry BL RPE LEP PB

Sharma et al, Differentiating Macrozantia moorei from M johnsonii 365

In assessing allelic variation at the same

loci, Nei’s genetic distance (Nei, 1978) was

calculated between the three populations

sampled. The genetic distance between two

congeneric species is generally considered to

be 0.4 (Gottlieb, 1977) and the genetic distance

between M. moorei and M. johnsonii was 0.48

(average of two populations of M. moorei)

which supports the previous conclusion of

Jones and Hill (1992) based on morphology,

that these are distinct species. The low levels

of genetic variability observance in M. moorei

and virtually no variability observance in M.

johnsonii may be due to the fact that the

populations have been derived from a small

number of plants and lack of gene flow from

other populations has resulted in independent

evolution of M moorei and M. johnsonii with

divergence caused mainly by random genetic

drift. The low number of polymorphic loci and

heterozygotes found in individuals may be

attributed to any of the following factors:

(a) the small and localised nature of the

population, (b) a genetic “bottleneck structure”

due to lack of gene flow from outside the

population, (c) inbreeding between individuals

within taxa, and (d) low sampling level.

1.K, Sharma, D.L. Jones

The results obtained here provide

supportive evidence for distinguishing

M. moorei and M. johnsonii as distinct species.

Acknowledgements

We thank Corinna Broers, Marion Garratt,

Ailsa Holland and Maggie Nightingale for

their technical support; Peter Machin for

collecting samples of M. johnsonii as permitted

by the Forestry Commission of New South

Wales; Ailsa Holland for assistance with the

sampling of M moore?; and Andrew Young for

his valuable input into the project.

References

GoTTLies, L.D. 1977. Electrophoretic evidence and

systematics. Avmals of Missourt Botanical Garden

64: 161-180.

Jones, D.L. and Hitt, K.D. 1992, Macrozamia johnsonii, a

new species of Macrozamia section Macrozamia

(Zamiaceae) from Northern New South Wales,

Telopea. 5: 31-34.

Nei, M. 1978. Estimation of average heterozygosity and

genetic distance from small numbers of individuals.

Genetics. 9: 583-590,

WENDEL, J.F. and WEEDEN, N.F. (1989), Visualisation and

interpretation of plant isozymes. In J/sozyimes in

Plant Biology. Eds. D.E. Soltis and PS. Soltis. pp.

5-45. Portland, Oregon: Discortdes Press.

Plant Biodiversity Research, GPO Box 1600, Canberra, Australian Capital Territory 2601,

Australia

P.l. Forster

Queensland Herbarium, Brisbane Botanic Gardens, Mt Coot-tha, Mt Coot-tha Road,

Toowong, Queensland 4066, Australia

SSE Be SEER SE ea i nora oa ro mane ab oc ab one dances batese asec Te oad aoe a ee ean enmnetmae em ntact ny deine d animate ht ae pee men siagee TLELLLD une pinignegeticcis sits gcienheris

Index of new names published in Austrobaileya 5(2)

Adacta abbatidnd Pedley Spin Vere obj esd cw sae ie ee ela hee altace eh te tells Bed oo vlaly nae bafele he biter 313

ACaACTE Gr bigtd PEGGY SPoDON: Hassle 5S aye dance: sce ber 88 ¥ 4, 4 # dos Sle cane ahaa: Babi elas Festal soe wee Lele ah ecdae ate 4 307

AGACIa GrevTOHICNG PEdey SP: DOVE jh fot oe eae sens Se RS OAS eat Wale Ef Hany ac HG grec ATEN os, wes 310

Acacia barakulensis Pedley SO. TOW: «<4 4 6 ue 4 uv 0d OF RW a a Res ee EOFS RS HEE deere HEE Ow 308

Avacta DUPdeKensts; PEGISY SPiN 4g soe ace ie tele vache ew ee Needs oe aoe Daan k Ane gis Ok Bap esetenc al acacia see Oy 313

ACHCTE LONVGL TION FECIEY-SO7 MOVs oe dts we eed pwd AG Ale, ara, ve Dean, Sande Bed & Fsod Iw A ATR gh See FR eal be 312

ACGCIG JONCITAEPEGICY SPIO 2 54 wreay ee o9 SS Wee BS WN BS BE TR ar EW Tye. eet VR Op A PT ten Eb fc we 6 314

AOGCIO-fil DES PEM SOs NOG cy hele ole tree ox w Wee te so ode oy aegnd Le chee bath 4 Boge bere beanas dead ROU a Sead poeta ON 315

ACAOIT TOMMUIS P CALEY SO. NOGA. Shes weed vaccine 8 9-u ehea gta an sivverdrs whut ace beet acd Say dakuant es whe at Page ooi 316

Acacia hendersonirPediey Sp. NOV... £4.04 % ee is od ee ee eee ee Ae Vea PES Hed thy ee EF meg 309

ALACIG JONAHTIS PEGICY SP UMIGV. oo os sre tet ae ceo 5 aye oe a Dat PRG wed Oe Ee ed etn ST ae 311

Acacia lacertensis PEAECY SPMOVa ¢ oe wt Sen celelics 6 00 hs xi 9 oh 9 ee Se aI rh ele A ene ee wae eh a 316

AGHCIG DP ROIGNING DL EAlLSy SPoOMOVE NL le oat sb end aah ee SR eo AES an hoe od ace Whee ce Rhee Ath bwsetlngen SEE ee EE 08 318

Atacia Fubricola Pedley SB: MOVs x 2i< 44 9 45 PERE e see VE we es ok OF eR ab ela 1 Share FT nae ea peel aa dale 309

ACaCirSCOpAGT I PedIGy SDATIOVE™ crac -e-g ete wee ol wor BRR EE EO eta Sone REL a ek AOE One adele p gk nie tal 319

Acacia solenoia PEdhesy Si HOV. 2 Lee aS, oh cline clare wale eae ot Wen eeee ath Wa Re ee SE ge pet ne eat ES pate Sate ee 319

Acacia tingoorensis Pedley nom. et stat. NOV. 2.0... cc ee eee eee eee ee eee eee ee heen ees Yaa ite Ca

Aphyliodiim glossocarpiin Pealey Spc NOV seg eh sett ye eas eat bd 2 ee ele hg ak bE EE ah hota beeen 214

Apnvilodiian (otfolinr PEGley SD. NOV nal osc 'e 6 aa. 8 294 Fike tes warn ace Malice tyler ee Pa eels eaten 212

Aphyllodium parvifolitum Pedley Sp. NOV... 6... cc ce cee ee ete eee nee eee eee eee eee eens 212

Aphyllodium schindleri Pedley Sp. nov. ... 0. eee eee ee eee eee ee eee eee eee eees 213

Aphyllodium stvlosanthoides Pedley sp. MOV... 0... cee eee eee eee ee eee ee eee eee tee 215

BapiiCiOnia Guensnes A. RABCAN-SD, TOV: 6 d65.4 35 vy kien ik sche Pian RO se GW ERE Dey ED LORE 4S pe aes 163

Babinetonia bidwillli AVR.Béati Sp. NOV. ce cc ce cei ee ee be ee dae deere eee ee eee eer oy Ban enes 161

Babineionia Drachvpoda A. R. Bean Sp. DOV ones ee een Ca 2 oe bee oa Ha Oo a AL Ppa pained eae 168

Babinetonia Govind AR Bean Se NOV 2 pads vcs seo oe OO ne Op de Eee PTE Ce pe we gle Keke 167

BaDiINGIONIG CLOSSAA REAL SP) TOV in: 4 plc5ei da eevee Vier Mie, ene areca, & be pualel al eavaneta ee! were ae nueg Wee ay ag leg aeRO eae 166

Babingtonia papillosa A.R.Bean 8p. MOV. 2. ce eee eee ee eee ne eet e ee eee ees 169

Babingtonia pluriflora (F.Muell.) A.R.Bean comb, nov... cc eee ene eee ett eee eens 165

BOBINGIONIO SHITE PIG BEA SOc TOM ans, oa ates, iar ysa ata able’ cath ope x DG, Lenerbat bare eae bd bP ge wes 162

Borenid Gala Duvet Sp, HOV s 63 ox. ccese da ef woth aA WS gl Sco. a cca ee doen bie APR GOR ade ayhh s Caeecaded Deety a eee abe Roe ke 287

Boronig-did ani ae: DUretta: SPs MOV. ace acne 4% wre eek 8 8 oC ah ewe ON Bea ban and, Ble qa ee omg What Sante eas pecans 292

Beronig- ex Celsg Duretto-Spe MOVs. 5 a ders a & eyk toe beh sh slege Oa ater abet E Gente e aoe gs eda aa cease arco gt hice pect ahatie hi 284

Boveria foctida Dutetto. Sp, MOVs x sep 6 Maric: bon BON bese 2 A ee tt Veto Ee ard & GER Sig LTS OES ORE Mego 285

Boronia forsterl PurettO-SpxWO Vs i x5 pete bee 4 5 ena ced Wate 5 & Sone andbece te AGE gel ped bee dcade eG AGI MLS denna doe eae 280

HOoroiia HOPOMOTDULETT SPs TAK aac ee So, eg Gk shee an Ste pol wok RPiep dal dasha te don A meats ls, eet ee acstereale edie | 288

POPOHIA: J CUSZICE TIUEETO Sh MOVE io. gon 8 eta se ew, ei ye ladle O Agia: 4p Abd wire Te A wlattrcend alti papacy A NON wre gb 282

Boronia OdOFatOUreito: SPs IOV, 4 rereany seg + bag Goa 0 acplsal 8 ek aces oe a ATE ae Al bein la dc ab antta Hae dale Ss Caedes See A Daete 294

Boronia palasepata. Durrett: Sp. DOWy cece hie ecg ween eee ee a ala a a Pg ee a ees waa, ele ang pon Haye eos 280

Boronia: GuINKANENSTS ADULTE: SP WOVIs Soc Sa HIS 5. Benge Fa Re lass phe nu, faded etn a Aa enact ta lets lo eiten Uplate 4a a ae 291

BOOM SPIendida-Duretto Spy DOW Gao vi wonce cn ek fae oy whe ad win Re Ee NLC R AR IE a pega BPR ance keke aa 278

Boronia squantipetala DarettO Sp. BOV:. ecko sce ob Mae % 4 DEW Ve Coe aware fine bp Red aoe & MLM Nels Rad te Ane bls 295

Dendrolobium cheelii (C.A. Gardner) Pedley comb. nov. .... 0 ccc eee eee ee beeen ees 221

Dendrolobium multiflorum Pedley sp.nov. 0... 0... ce ee ee eee eee eee eee beeen ee bea 220

Dendrolobium polyneurum (S.T.Blake) Pedley comb. nov... .... cece eee eee nee teenies 221

Dendrolobium umbellatum var. hirsutum (DC.) Pedley var. nov. . 0... 00.0 cc eee eee eee teas 223

Desmodium sabg. Acanthocladum Pedley subg. nov. 2... 0. cee eee eee eee eee eee ee tees 225

Desmodium sev. Ariligta Pedley- Ser DOV. vv e+ ak ste ewok aR oo ei Le Hig he Rhos HT hae A ob wzpiees Hoe Ee eS 226

Desmodium sect. Desmodiopsis (Schind].) Pedley comb. nov, ....... ce ee tenets 226

Hesiod eiareosiin Pedley SP. NOV. bc 5-.- pac ence ne lena, yn adie Hos ts nds old tw wht ahelecg plead tala ded 242

Desnodiwnpullent? Pedley Sp. NOV. 6. a6: 5 48 « ge arene hele wt os Cae By OF Welen ok Vane gle a, TMs eter EEG wea 242

Desmodium pycnotrichum Pedley sp. NOV... 0... een ee eee ee beeen eee e nes 236

Desmodium ser. Sagotia (Duchass.) & Walpers comb. & Stat. NOV... ee ee eee e ee eee nes 226

Desmodium ser, Stenostachys (Schindl.) Pedley stat. nov... 0... cc ce eee een eee eee 226

LIESTROGINUT THVIERSE PECIGY SPIO, eitrn veh tee a x ohh Se eel okde & oh ik yok oe AE A et aE hs. aed Rta w aA 248

Desmodium whitfordii (Schindl.) Pedley comb. nov. ...... 0... ccc ee ete eter eee eee enes 247

Everistia 8. F Reynolds & R.J,F. Hendenenove . oe ck eka ea ehcags Oye Meith ee hs be pha ar aa hk bee bbe ge 354

Everistia vacciniifolia (F, Muell.) 8.T.Reynolds & R.J.F.Hend, comb. nov, oo... ee eee etnies 355

Everistia vaccinitfolia f. crassa S.T.Reynolds & R.J.F.Hend. forma nov. ........0 0.0. cee ee ees 357

Eyeristia vacciniifolia var. nervosa S.T. Reynolds var. MOV... 0... ce eee teen e ee eee nn enans 357

Habenaria praecox Lavarack & Dockiill sp: Nowy ways es os a ee wa iv he a fee bb ee ga ne bea WO 331

Jasminum jenniae W.K.Aarris & G.Holmes sp. nov. . 6... ce eee tee ee eens 341

Lithomyrtus cordata (A.J, Scott) N.Snow & Guymer comb, nov... 0... cc ee tence ees 184

Lithomyrtus densifolia N.Snow & GUyMerPr SP. NOV... 6. eee eee eee eee eee eee eee ne ees 185

Lithomyrtus grandifolia N.Snow & GuymMer sp. NOV... 6.6.0... 00 eee dtd '4s soph glia erraabeeckhyes teks hse saute foeheeyts Leal 4 189

Lithomyrtus hypoleuca N. Snow & GUYMET SP. NOV. oc eee ee eee ee ete ee eee 19}

Lithomyrtus kakaduensis N.Snow & GUYMEL SP. NOV... eee eee tet tee eee eee eben i9]

Lithomyrtus linariifolia N.Snow & Guymer sp. nov... 6... ee bp dip acbiat hin sont a7ephe vende oad: oda 4g CS e's 193

Lithomyrtus repens N.Snow & Guymer sp. NOV... eee tees gf ET BEEN bd Eo ci a a . PY

Lithomyrtus retusa (Endi.) N.Snow & Guymer comb. NOV. 2... 2. eee tenner eee eens 198

Oldenlandia gibsonii, D.A.Halford sp, novi... oc eee eee nee eee eben ees 337

Piplladiint WaCkert. PEGley- SPU vie < os dase areca peicy 4k Ak gate 69 eee gn Sy cacy Bp a gears DOP Tee WA ATP Se Boece > 256

Phyllodium pulchellum vay. glabrius Pedley var. nov. 0... 00 cee eee eee een nes 257

Siplidnan Taiophipliran Fe, BeBeaty SBonOVe ene y eo depen eens len eer sen ee taeme gin eames dente pe he Aree firey ae acess 326

Stylidium ramosissimum A.R.Bean sp. DOV... 0. ee ee ee ee eee eee beens 328

Tadéhagi-rabtustrnt Pedley Spsnow. scr" asp cack ack SS a ed eB act ech aa EE gb ReaD Send pe 258

Uromyrtus rostrata (Lauterb.) N.Snow & Guymer comb. NOV... 66. ee ce eee eee e nna 18]

“AS saath loans as LALLA AL seen ie a atc pias arya ae nated 4 woh asian

Queensland Herbarium

Publications for sale

Floras

A Key to Australian Grasses by B.K. Simon (1990), 150 pp., card cover.

Ferns of Queensland by S.B. Andrews (1990), 427 pp., illustrated, hard cover.

Honey Flora of Queensland, 3rd edition by S.T. Blake and C. Roff (1988), 224 pp., illustrated,

card cover.

Keys to Cyperaceae, Restionaceae and Juncaceae of Queensland. Botany Bulletin No. 5 by

P.R. Sharpe (1986), 47 pp., illustrated, card cover.

Flora of South-eastern Queensland by T.D. Stanley and E.M. Ross.

Volume I (1983), 545 pp., illustrated, card cover.

Volume 2 (1986), 623 pp., illustrated, hard cover.

Volume 3 (1989), 532 pp., illustrated, hard cover.

Vegetation Surveys

Vegetation Survey of Queensland: South West Queensland. Queensland Botany Bulletin No. 4

by D.E. Boyland (1984), 151 pp., illustrated, including a map, card cover.

Vegetation Survey of Queensland: South Central Queensland. Queensland Botany Bulletin

No. 3 by V.J. Neldner (1984), 291 pp., illustrated, including a map, card cover.

Vegetation Survey of Queensland: Central Western Queensland. Queensland Botany Bulletin

No. 9 by V.J. Neldner (1991), 230 pp., illustrated, including a map, card cover.

Vascular Plants of Western Queensland. Queensland Botany Bulletin No. 11 by V.J. Neldner

(1992), 176 pp., illustrated, card cover.

Vegetation of the Sunshine Coast: Description and Management. Queensland Botany Bulletin

No. 7 by G.N. Batianoff and J.A. Elsol (1989), 107 pp., illustrated, including maps, card

cover.

Vegetation Map and Description, Warwick, South-eastern Queensland, Queensland Botany

Bulletin No. 8 by P.A.R. Young and T.J. McDonald (1989), 47 pp., illustrated, including a

map, card cover. |

Vegetation Description and Map: Ipswich, South-eastern Queensland. Queensland Botany

Bulletin No. 10 (1991), by James A. Elsol, 61 pp., maps, card cover.

Vegetation Survey and Mapping in Queensland: Its Relevance and Future, and the Contribution

of the Queensland Herbarium. Queensland Botany Bulletin No. 12 by V.J. Neldner (1993),

76 pp., card cover.