Reinstatement and revision of Kayea Wall.

(Calophyllaceae) in Australia, including two new

species from Queensland’s Wet Tropics bioregion

W.E. Cooper’? and F.A. Zich'”

Summary

Cooper, W.E. & Zich, F.A. (2022). Reinstatement and revision of Kayea Wall. (Calophyllaceae) in

Australia, including two new species from Queensland’s Wet Tropics bioregion. Austrobaileya 12:

1-13. The genus Kayea Wall. is taxonomically revised for Australia and comprises three species:

Kayea larnachiana F.Muell. (syn. Mesua larnachiana F.Muell.), and two new species described

here: K. concinna W.E.Cooper & Zich and K. meridionalis W.E.Cooper & Zich. All species are

described and illustrated, with notes on habitat, distribution and a suggested conservation status. A

comprehensive synonymy for the new species, listing published phrase names and a discussion of

lectotypification in Kayea larnachiana are also provided. An identification key is included for the

three species of Kayea in Australia.

Key Words: Calophyllaceae; Clusiaceae; Kayea concinna; Kayea larnachiana; Kayea meridionalis;

Mesua; Australia flora; Queensland flora; Wet Tropics bioregion; taxonomy; new species;

identification key

‘Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor Road,

Smithfield, Queensland 4878, Australia.

“National Research Collections Australia, Commonwealth Industrial and Scientific Research

Organisation (CSIRO), GPO Box 1700, Canberra, ACT 2601, Australia.

>Corresponding Author. Email: wendy@williamtcooper.com.au.

Introduction

Calophyllaceae belongs to the _ clusioid

clade (Malpighiales) which comprises

five monophyletic families: Bonnetiaceae,

Calophyllaceae, Clusiaceae Ss

Hypericaceae, and Podostemaceae (Ruhtfel ef

al. 2016; Cabral et al. 2021). Calophyllaceae

is a pantropical family comprised of 14

genera and about 460 species (Stevens 2001

onwards). It includes the genera Calophyllum

L., Caraipa Aubl., Clusiella Planch. & Triana,

Endodesmia Benth., Haploclathra Benth.,

Kayea Wall., Kielmeyera Mart. & Zucc.,

Lebrunia Staner, Mahurea Aubl., Mammea

L., Marila Sw., Mesua L., Neotatea Maguire,

and Poeciloneuron Bedd. (Stevens 2001

onwards; Ruhfel ef al. 2016; Cabral ef al.

2021). Calophyllum (six species), Mammea

(two species, including Christmas Island) and

Kayea Wall. (three species) occur in Australia.

The Calophyllaceae is divided into two

tribes, Endodesmieae Engl. and Calophylleae

Choisy. In their reconstruction of phylogeny

for the tribe, Cabral et al. (2021) placed the

genera into four main clades: a Calophyllum

clade [Calophyllum + Mesua|, a Mammea

clade [Mammea]|, a Kayea clade [Kayeal,

and a Neotropical clade. Kayea and Mesua

have until quite recently been considered

congeneric, but are in separate clades with

other genera (Zakaria & Choong 2007; Ruhfel

et al. 2016; Cabral et al. 2021), thus supporting

the reinstatement of Kayea by Stevens (1993).

Early authors recognised Kayea and

Mesua as distinct genera based on differences

in the ovaries and stigma lobes (Wallich

1831; Bentham & Hooker 1862; Mueller 1887;

Bailey 1899). Kostermans (1969) reduced

Kayea to synonymy under Mesua and

Accepted for publication | April 2022, published online 29 June 2022

recognised 40 species in the combined genus.

Stevens (1993) distinguished the two genera

morphologically and, stating that there was

no evidence for them forming a monophyletic

eroup, reinstated Kayea but this was not

adopted, or was overlooked in the Australian

context. Kayea 1s distinguished from Mesua

by the presence of papillate stems, a four-

carpelled ovary (Mesua with two) with 4-12

ovules (Mesua with two), a long style which

is Shortly branched, four narrow stigma lobes

(Mesua with two that are broadly peltate),

c-Shaped staminal thecae with a terminal

gland and accrescent sepals which continue

to expand after anthesis enclosing the ripe

indehiscent fruit (vs. not accrescent and not

enclosing the fruit in Mesua) (Stevens 2007).

The growing terminal bud in most Kayea

species aborts as new growth emerges from

the upper axils and this feature was considered

a synapomorphy by Stevens (2007). However,

Australian Kayea have terminal bud scales

which are stipule-like, and often persistent.

Kayea 1s a genus of more than 70 species

(Stevens 2001 onwards, 2007; Cabral ef al.

2021) of monoecious trees and shrubs from

the Old World tropics (Notis 2004; Ruhfel

et al. 2016), occurring in India, Thailand,

Vietnam, Cambodia, Malesia including

New Guinea and Australia. The three Kayea

species that occur in Australia are endemic to

the rainforests of the Wet Tropics bioregion

in north Queensland: Kayea larnachiana

F.Muell., K. concinna W.E.Cooper & Zich sp.

nov. and K. meridionalis W.E.Cooper & Zich

Sp. nov.

Kayea meridionalis was first collected at

Boonjie in 1974 by A.K. Irvine (AKI 772, CNS)

and the first collection of K. concinna was

made near the head of Noah Creek in 1977 by

B.P.M. Hyland (B. Hyland 9355, CNS). These

two taxa have, until recently, been identified

under one phrase name (Mesua sp. (Boonjee

A.K.Irvine 1218), or numerous variants

thereof; however, with additional specimens

collected in recent years, two morphologically

distinct species have been identified. In 1984

a manuscript describing one new taxon and

recognising one putatively distinct but poorly

collected taxon (then known from one sterile

Austrobaileya 12: 1-13 (2022)

Specimen), was submitted for publication

but never published (P.F. Stevens & B.P.M.

Hyland, pers. comm.).

All Australian Kayea species are restricted

in occurrence within the Wet Tropics

bioregion with allopatric distributions. Kayea

meridionalis is also altitudinally disjunct,

occurring above 600 m at Boonyie and at

the head of the East Mulgrave valley within

Wooroonooran National Park, while K.

concinna and K. larnachiana occur below

500 m in the Daintree National Park.

Materials and methods

The study is based upon the examination

of herbarium material from BRI and CNS

combined with field observations. Images of

types and other specimens held at BRI, K

and MEL have also been examined and are

indicated as i.d.v. (imago digitalis visa).

Measurements of the floral parts and fruits

are based on freshly collected specimens and

material preserved in 70% ethanol. Common

abbreviations in the specimen citations are:

LA (Logging Area), NPR/NP (National Park

Reserve/National Park) and TR (Timber

Reserve). Boonyjie has sometimes been spelt

as ‘Boonjee’ on specimen labels; however, the

accepted spelling has the ‘ie’ (QPNS 2022).

Botanical terminology is based on

Beentjie (2010).

Taxonomy

Kayea Wall., Plantae Asiaticae Rariores

3(10): 4 (1832).

Type: Kayea floribunda Wall.

Kayea in Australia: Monoecious subcanopy

trees, glabrous; bark smooth or slightly

flaky, exudate meagre or absent; twigs

flattened at nodes, papillate; new growth

pink; stipules absent. Interpetiolar bud scales

stipule-like, mostly persistent, colleters at

base. Leaves petiolate, simple, opposite,

glabrous, apex often mucronulate; venation

brochidodromous, tertiary venation reticulate

with a latex cavity within each reticulum.

Inflorescence terminal or rarely axillary, a

1—6-flowered racemose cyme; bracts and

bracteoles present; sepals 4, 2-whorled, outer

Cooper & Zich, Kayea in Australia

pair initially connate, concave, inner pair

free; petals 4; stamens numerous with slender

filaments; anthers with a gland at base; ovary

superior, 4-carpellate with 4-12 ovules,

placentas intrude at the base; style solitary;

stigma 4-fid. Fruit indehiscent, a woody berry

surrounded by large, accrescent, furfuraceous

sepals.

Etymology: Kayea is named in honour of

English botanist, mycologist, bryologist and

artist Dr Robert Kaye Greville (1794-1866).

Key to Australian Kayea species

1 Mature leaves greater than 150 mm long; leaf base cordate or rounded. .3. K. larnachiana

1. Mature leaves less than 80 mm long; leaf bases cuneate. .................. 2

2 Stipule-like bud scales up to 1.5 mm long; leaf lamina broadly ovate,

width at widest point greater than one third of the length; sepals

ORES pote lychee! caard ath Site eee dss Wien as

ai Ate wy, a lw ee 2. K. meridionalis

2. Stipule-like bud scales 3—5 mm long; leaf lamina lanceolate, width at

widest point less than one third of the length; sepals pinkish-fawn... .

1. Kayea concinna W.E.Cooper & Zich sp.

nov.

Similar to Kayea stylosa Thwaites, but differs

from that species by having leaf bases cuneate

(vs. obtuse or rounded); inflorescence length

up to 18 mm (vs. 30—35 mm); flower diameter

8—l11 mm (vs. c. 5mm); style extending beyond

anthers by up to 2 mm (vs. 5 mm). Typus:

Australia. Queensland. Cook DIsTRIcT: Noah

Creek, 20 February 2020, W. Cooper 2635

& R. Russell (holo: CNS 151003.1 [1 sheet +

spirit|), iso: BRI, CANB, MO distribuendi).

Mesua sp. (Boonjee A.K.Irvine 1218) (in

part): Jessup (1997: 50, 2021).

Mesua sp. Boonjee (A.K.Irvine 1218) (in

part): TSCS (1998); Hyland ef al. (2010); Zich

et al. (2020).

Mesua sp. (Boonjee AKI 1218) (in part):

Hyland et al. (1999: 60).

Mesua sp. (Boonyjie A.K.Irvine 1218) (in part):

Jessup (2002: 49, 2007: 49, 2010: 44, 2019).

Mesua sp. (Boonjee) (in part): Cooper &

Cooper (2004: 126).

Mesua sp. 1 Boonjee (in part): DEH (2004).

Mesua sp. (A.K.Irvine 1218; Boonjee) (in

part): DEH (2004).

Mesua sp. Boonyie (A.K.Irvine 1218) Qld

Herbarium (in part): CHAH (2005, 2021).

1. K. concinna

Illustrations: Cooper & Cooper (2004: 126,

as Mesua sp. (Boonjee)).

Small tree to 7 m, dbh to 90 cm; buttresses

absent; exudate meagre and clear, or absent;

bark with numerous vertical fissures and

occasional + horizontal creases, thin, papery,

smooth but seasonally flaky, reddish-brown;

twigs + terete, flattened at nodes with

several minutely shallow longitudinal ribs

becoming slightly flaky, minutely papillate

becoming less pronounced in older growth,

internodes 13-38 mm long; new growth

pink; interpetiolar bud scales often persistent,

initially connate but soon dehiscing into a

pair, narrowly triangular, 3—5 mm long, c. 0.5

mm wide at base, glabrous. Leaves: petioles

flat along upper side and becoming shallowly

erooved in dried specimens, 4—8 mm long;

lamina lanceolate, coriaceous, 53-103 mm

long, 13-30 mm wide, discolorous, glabrous,

base cuneate, apex acuminate and usually

mucronulate, margins entire; venation:

primary vein slightly raised on both surfaces;

secondary veins 10—14, 3—5.5 mm apart, angle

of divergence from primary vein 10—20",

forming loops 1—2 mm from margin, flush

with adaxial surface, slightly raised abaxially;

tertiary veins reticulate with a solitary round

or slightly elongated latex cavity in the centre

of each reticulum. Inflorescence a terminal

(rarely axillary) 3—5-flowered racemose cyme,

13-18 mm long; bracts several, triangular, c.

| mm long and wide, clustered at peduncle

bases and persistent well after inflorescences

have dehisced; colleters several, coppery-

coloured, clustered amongst and behind

bracts; peduncle 4.5—11 mm long; bracteoles

paired at pedicel junctions, caducous,

narrowly triangular, c. 2.5 mm long and 0.2

mm wide at base, glabrous, entire; lateral

pedicels 1.5—4 mm long; terminal pedicels

up to 3. mm long. Flowers not recorded as

fragrant, diameter 8—11 mm; sepals in outer

whorl ovate-orbicular, apex obtuse, inner

pair orbicular, deeply concave, 3—4 mm long

and wide, pale pinkish-fawn coloured; petals

broadly ovate, becoming convex, c. 7 mm long

x 4.5 mm wide, white, apex acute, margin

entire but fimbriate near apex and folding to a

point, glabrous; stamens numerous, filaments

c. 4.5 mm long; anthers c-shaped, c. 0.8 X 0.6

mm; ovary broadly ellipsoid, c. 2 X1.5 mm;

ovules usually 4 per locule, c. 1 mm long;

style c. 5 mm long; stigma rays 0.25—0.6 mm

long. Fruit unknown. Fig. 1.

Additional selected specimens (from 7 examined):

Queensland. Cook District: VCL Noah, head of Noah

Creek, Mar 1977, Hyland 9355 (CNS); Noah Creek,

Cape Tribulation, Mar 1996, Jensen 643 (BRI, CNS);

Noah Creek, Feb 1998, Cooper & Jensen 73 (BRI, CNS,

NSW); Noah Creek, Daintree NP, Feb 2019, Cooper

2584, Jensen & Hawkes (CNS); ibid, Feb 2020, Cooper

2629, Hawkes & Carmichael (CNS); ibid, Jun 2020,

Cooper 2639 & Ford (CNS).

Distribution and habitat: Kayea concinna

is known from mesophyll and notophyll

rainforest on riparian granite silt in the

braided channel of the Noah Creek valley

within the Daintree National Park (Map 1) at

altitudes from near sea level to 500 m.

Kayea concinna has’ morphological

similarities to K. stylosa from Sri Lanka

rather than to other species from Australia,

Asia or Malesia.

Notes: Several visits were made to Kayea

concinna trees that sparsely flowered over a

season of more than 6 months but failed to

set fruit over 2 years. Other plant features are

enough to confirm that this species belongs in

Kayea: papillate twigs, colleters behind bud

scales and bracts, a small latex gland on the

Austrobaileya 12: 1-13 (2022)

anther connective, 4-fid stigma and ovary

l-locular with accrescent sepals around the

fruit.

Specimens from Noah Creek were first

collected in 1977 and were maintained as a

distinct taxon from specimens collected at

Boonyjie by Bernie Hyland at the Australian

National Herbarium — Atherton (QRS; now

Australian Tropical Herbarium, CNS) until

1996 when they were incorporated into Mesua

sp. (Boonjee AKI 1218).

Phenology: Flowers have been recorded from

February, March, May, June and September.

Fruiting has not yet been observed.

Conservation status: Kayea concinna 1s

known from only a few collections along

Noah Creek in Daintree National Park. The

population is very restricted in area and

only recorded within the National Park and

while no immediate threats were observed,

plants have not been found in fruit despite

monitoring of flowering individuals over

two years. Using GeoCat (Bachman ef al.

2011) based on known locations, Extent

of Occurrence is estimated at 10 km’, and

Area of Occupancy is estimated at 24 km7’.

A suggested conservation status for Kayea

concinna 1S Endangered [EN BlabQ, 1,

1v)+B2ab(, 101, 1v)| (UCN 2012).

Etymology: The specific epithet 1s derived

from the Latin concinna (neat, pretty, elegant);

referring to the elegant leaves and growth

habit of this tree.

2. Kayea meridionalis W.E.Cooper & Zich

Sp. nov.

Similar to Kayea stylosa Thwaites but differs

from that species in the length of the stipule-

like bud scales up to 1.5 mm (vs. 2—3 mm); leaf

shape broadly ovate (vs. lanceolate); flower

diameter greater than 7.5 mm (vs. 5 mm);

inflorescence length up to 23 mm (vs. c. 40

mim); style extending beyond anthers by | mm

(vs. c. 5 mm). Typus: Australia. Queensland.

Cook Dzstrict: Timber Reserve 1230,

Boonjee Logging Area, 4 March 1975, A.K.

Irvine 1218 (holo: CNS [comprising 3 sheets

Cooper & Zich, Kayea in Australia

ORS 20579.1, QRS 20580.3, QRS 20581.4 and

spirit QRS 20579.2]), iso: A, BRI, CANB, K,

L, MEL, MO, SING distribuendi).

Mesua sp. aft. M. elmeri: Hyland (1982: 107,

133, 143, Code 745).

Mesua sp. “‘Boonjee’ (A.K.Irvine 1218):

Thomas & McDonald (1987: 21, 1989: 22).

Mesua sp. (=RFK/3128): Hyland & Whiffin

(1993: 106, 116, 174, Code 745); Christophel

& Hyland (1993: 4, 35, 83, pl. 21(b), Code

745); Hyland & Whiffin (1993: 74, Code 745).

Mesua sp. (Boonjee AKI 1218): Hyland ef al.

(1994: 302); (in part) Hyland ef al. (1999: 60).

Mesua sp. (Boonjee A.K. Irvine 1218): Jessup

(1994: 74); (in part) Jessup (1997: 50, 2021).

Mesua sp. Boonjee (A.K.Irvine 1218): (an

part) TSCS (1998); Hyland et al. (2010); Zich

et al. (2020).

Mesua sp. (Boonyjie A.K.Irvine 1218): (in part)

Jessup (2002: 49, 2007, 2010: 44, 2019).

Mesua sp. (Boonjee): (in part) Cooper &

Cooper (2004: 126).

Mesua sp. 1 Boonjee: (in part) DEH (2004).

Mesua sp. (A.K.Irvine 1218; Boonjee): (in

part) DEH (2004).

Mesua sp. Boonyjie (A.K.Irvine 1218) Qld

Herbarium: (in part) CHAH (2005, 2021).

Illustrations: Christophel & Hyland (1993:

83) as Mesua sp. (=RFK/3128); Zich ef al.

(2020) as Mesua sp. Boonjee (A.K.Irvine

1218).

Small tree to 20 m, dbh 22 cm; buttresses

absent; exudate meagre and clear, or absent;

bark mostly smooth with horizontal creases

and sections of circular impressions which

resemble beaten copper, reddish-brown;

twigs + terete, flattened at nodes, with several

minutely shallow longitudinal ribs becoming

slightly flaky, minutely and sparsely papillate,

internodes 14-38 mm; new growth pink;

interpetiolar bud scales, persistent or

caducous, initially connate but soon dehiscing

into a pair, ovate or triangular, 0.65—1.5 mm

long, 0.6—0.8 mm wide; colleters coppery-

coloured, glabrous, often becoming dehiscent.

Leaves: petioles flat along upper side and

not grooved in fresh specimens but grooved

in dried material, 6-11 mm long; lamina

broadly ovate, 45-73 mm long, 16-30 mm

wide, discolorous, glabrous, base cuneate,

apex acuminate and mucronulate, margins

entire; venation: primary vein slightly raised

on upper side and distinctly raised below;

secondary veins 10—14, 2-3 mm apart, angle

of divergence from primary vein 30—40°,

forming loops 1.5-3 mm from margin,

slightly raised on both surfaces; tertiary veins

reticulate with a solitary round or slightly

elongated latex cavity in the centre of each

reticulum. Inflorescences a terminal (rarely

axillary) 1—6-flowered (rarely 4 or more)

cyme or racemose cyme 20—23 mm long or

a solitary flower; bracts triangular, several,

clustered at peduncle bases and persistent,

c. 0.75 X 0.75 mm, several colleters at base

between bract and peduncle; peduncle 4—20

mm long; bracteoles in opposing pairs part

way along peduncle or apical, caducous,

narrowly ovate, c. 1.8 < 0.5 mm, glabrous,

entire; solitary flowers with pedicels 2.5—

7 mm long; cymose flowers with lateral

pedicels 5—11.5 mm long, terminal pedicels

3—5 mm long. Flowers not noted as fragrant,

diameter 7.5—-12 mm; sepals in outer whorl

oblate, apex rounded or minutely acute;

inner pair + orbicular, concave, c. 4.5 X 4.5

mm, green, entire; petals broadly elliptical,

concave, boat-shaped, or obovate, c. 7 X

4 mm, white, apex acute, margin entire;

stamens numerous, filaments c. 5 mm long;

anthers c-shaped, c. 0.5 X 0.7 mm, gland near

base; ovary broadly ovoid or turbinate, 2—2.25

mm long and wide; ovules usually 4, c. 0.7

mm long; style 5-6 mm long; stigma rays

0.5—0.7 mm long. Fruiting peduncle 8-12

mm long. Fruit a dry and somewhat woody

berry surrounded by accrescent furfuraceous

sepals, apex of the fruit exposed, stigma lobes

persistent, obovoid, 25—35 mm long and wide,

rusty brown; seeds | or 2, c. 14mm long X 20

mm wide X 18 mm deep. Figs. 2 & 3.

Additional selected specimens (from 21 examined):

Queensland. Cook District: East Mulgrave River, Nov

1995, Jensen 484 (CNS); Stockwellia Track, Boonjee,

Dec 1998, Jensen 944 (CNS); TR 1230, Boonjee, Nov

1974, Hyland 3129RFK (BRI, CANB, CNS); ibid, Nov

1974, Hyland 3I130RFK (BRI, CANB, CNS); ibid, Nov

6 Austrobaileya 12: 1-13 (2022)

stipule-like bud scal

dehisced

bracteoles

bud scale

Fig. 1. Kayea concinna. A. showing terminal connate stipule-like bud scales and dehisced stipule-like bud scales

(Cooper 2629, CNS). Photo: W. Cooper B. showing open-flowered inflorescence, stipule-like bud scale at inflorescence

base, bracteoles and flowers at anthesis with 4-fid stigma (Cooper 2584, CNS). Photo: R. Jensen.

Cooper & Zich, Kayea in Australia

_— =

bracts.#

- =

4 a

a . ™

y aj :

ate * Yh

ia Ry

A 3 =.

Fig. 2. Kayea meridionalis. A. flower at anthesis with 4-fid stigma and bracts (Cooper 2741A, CNS). B. fruit (Cooper

2734, CNS). Photos: T. Hawkes.

1974, Hyland 313IRFK (CNS); ibid, Jan 1975, Irvine

IIII (CNS); ibid, Jan 1975, Irvine 1136 (CNS); ibid, Jan

1975, Irvine 1137 (CNS); ibid, Dec 1975, Irvine 1682

(CNS); ibid, Jan 1976, Irvine 1735 (CNS); TR 1230,

Bartle Frere, Boonjee LA, Aug 1992, Gray 5470 (CNS);

Stockwellia Track, Boonjee, Oct 2020, Cooper 2734 &

Hawkes (CNS); ibid, Jan 2021, Cooper 2741A (CNS).

Distribution and habitat: Kayea meridionalis

occurs in complex notophyll rainforest on

the contact zone of basalt and metamorphic

geology in the Boonjie area on the Atherton

Tableland and in the upper East Mulgrave

River valley (Map 1) at altitudes between 600

m and 760 m.

Kayea meridionalis has morphological

similarities to K. stylosa from Sri Lanka

rather than to other species from Australia,

Asia or Malesia.

Phenology: Flowers have been recorded in

January and February and fruit in August,

October, December, January and February.

Conservation status: Kayea_ meridionalis

is known from two disjunct areas in

Wooroonooran National Park, in the upper

East Mulgrave River and the Boonjie area

near Mt Bartle Frere. Remote areas of the

National Park remain poorly explored and the

size of known subpopulations has not been

assessed extensively, but the subpopulations

are restricted 1n area and only recorded within

the National Park. There are no immediate

threats evident for the species. Using GeoCat

(Bachman ef al. 2011) based on known

locations, Extent of Occurrence is estimated at

46 km’, and Area of Occupancy is estimated

at 32 km’. A suggested conservation status

for Kayea meridionalis is Endangered |EN

Blab@, 111)+B2ab(, 111)] (UCN 2012).

Etymology: The specific epithet is derived

from the Latin meridionalis (southern);

referring to the distribution of this species

being the southernmost of all known Kayea

species.

3. Kayea larnachiana F.Muell., The Victorian

Naturalist 3(9): 126 (1887).

Mesua_ larnachiana (F.Muell.) Kosterm.,

Reinwardtia 7: 427 (1969). Type citation:

“On the Mossman-River; W. Sayer”. Type:

Australia. Queensland. Cook DISTRICT:

8 Austrobaileya 12: 1-13 (2022)

55 mm

Fig. 3. Kayea meridionalis. A. habit of branchlet. B. seedling. C. fruit, lateral and apical view. D. embryo with 3

cotyledons. E. flower, longitudinal section. A & E from /rvine 1215 (CNS); B—D from /rvine 1735 (CNS). Del. T. Nolan

Cooper & Zich, Kayea in Australia

Mossman’s River, Trinity Bay, s.dat., W.

Sayer s.n. (lecto: BM 000611321 i.dv. [buds

present], fide Kostermans (1969: 427);

possible isolecto: s.dat., W. Sayer 147A (BRI

[AQ0166459| id.v. [fruit present], MEL

2227332 i.d.v., [fruit present], MEL 2227333

i.d.v. [flower present]; MEL 2227334 i.d.v.

[buds present]; s.dat., W. Sayer 143 (MEL

2227335 i.d.v. |buds present]).

Kaya larnachiana’ F.Muell., Systematic

Census of Australian Plants Suppl. 4: 3

(1889), orth. var.

Illustrations: Christophel & Hyland (1993:

83); Cooper & Cooper (2004: 125), Zich et al.

(2020).

Small tree to 15 m, dbh 30 cm; buttresses 1f

present are small; exudate meagre and clear,

or absent; bark flaky; twigs + terete, flattened

at nodes, bark flaky, papillate, internodes

25-70 mm; new growth pink; interpetiolar

bud scales persistent or caducous, initially

connate soon dehiscing into a pair, narrowly

triangular with a midrib, 3.5—6.5 mm long,

Q.75—1 mm wide at base, glabrous, colleters

coppery-coloured, adaxially persistent.

Leaves: petioles terete, flaky, 2-10 mm long,

2-3 mm thick; lamina oblong-ovate, 150-—

240 mm long, 40—100 mm wide, discolorous,

glabrous; base cordate, obtuse or rounded;

apex acuminate often mucronulate, entire;

venation: primary vein slightly raised on

upper side and distinctly raised below;

secondary veins 15—25 mm apart, angle of

divergence from primary vein 5—20°, forming

loops 2—3 mm from margin, slightly raised

on both surfaces; tertiary veins reticulate

with latex cavities within each reticulum.

Inflorescence a terminal (rarely axillary)

3—10-flowered cyme or racemose cyme, 20—

23 mm long, or a solitary flower; bracts and

bracteoles similar, caducous or persistent at

peduncle and pedicel bases, triangular, 1-2 x

to 1.5 mm, several colleters at base between

bract and peduncle; peduncle 4—20 mm long;

pedicels 6—7.5 mm long, papillate. Flowers

fragrant or not fragrant, diameter 11—13.5

mm; sepals in outer whorl orbicular, concave,

glabrous, sometimes papillate, apex rounded

c. 7 X 8 mm; inner pair oblong-orbicular,

concave, c. 7 X 6 mm, white, entire; petals

concave or boat-shaped, broadly-elliptical or

orbicular, c.7 X 6mm, white, apex recurved,

entire; stamens numerous, filaments 5—6 mm

long; anthers c-shaped, c. 1 X 0.9 mm, gland

near base; ovary broadly ovoid, c. 3 mm long

and wide; ovules usually 4-12, 1.5—2 mm

long; style 4—5 mm long, stigmarays c. 0.4mm

long. Fruiting peduncle 5—12 mm long. Fruit

a dry somewhat woody berry surrounded

by accrescent furfuraceous sepals, only

the apex of the true fruit 1s exposed and the

stigma lobes are persistent, oblate-orbiculate,

20-38 mm long, 26—51 mm wide and c. 46

mm breadth, rusty brown, endocarp c. 3 mm

thick; seeds 1-3, 20—26 mm long, c. 16 mm

wide and 20 mm deep. Fig. 4.

Additional selected specimens (from 27 examined):

Queensland. Cook DISTRICT: Cooper Creek, Jun 1969,

Mazlin 4346 (BRI i.d.v.); Parish of Alexandra, Cooper

Creek, Dec 1984, Gray 3817 (BRI, CNS); Portion 49V,

Cooper Creek, Hyland 13878 (BRI, CNS); Nr Portion

54V, Cooper Creek, Parish of Alexandra, Nov 1983,

Hyland 12859 (BRI, CNS, DNA, NSW); Portion 49,

Alexandra, Cooper Creek, Oct 1975, Hyland 3335RFK

(BRI, CNS); Cooper Creek between Daintree River

and Noah Creek, Nov 1986, Zucker MCT 129 (BRI

i.d.v.); Cooper Creek, Turpentine Road, Nov 1989,

Jessup, Guymer & Dillewaard GJD2808 (BRI i.d.v.);

W. of Cooper Creek between Daintree River and Cape

Tribulation, Oct 1973, Webb 11013 & Tracey (CNS);

Mossman Gorge NP, Jan 2000, Gray 7754 (BRI, CNS);

Mossman Gorge NP, Mossman River, Dec 1984, Jessup

701 (BRI 7.d.v.); Mossman River Gorge, Nov 2004, Jago

6778 & Keith (BRI, MEL, NSW i.d.v.); Mossman Gorge

National Park, Dec 1997, Forster PIF 21951, Jensen

& Booth (BRI i.d.v.); ibid, Nov 2018, Cooper 2572 &

Hawkes (CNS); NPR 133, Mossman Gorge, Dec 1976,

Hyland 9243 (BRI, CNS).

Typification: The type citation in the

protologue of Kayea larnachiana 1s given

as “On the Mossman-River; W. Sayer.” and

does not provide a collector’s number or date.

Following the taxon description, Mueller

(1887: 127) writes that “The descriptive notes

have been elaborated from specimens with

young flower-buds and with over-ripe fruit”.

Six herbarium sheets have been located in

BM, BRI and MEL that appear to be original

material for Kayea larnachiana. They are all

labelled as having been collected by Sayer

and all lack a collection date. The collection

locality is given as either ““Mossman|’s] River”

or “Trinity Bay” or both localities are given.

Label locality information varies slightly

Austrobaileya 12: 1-13 (2022)

Fig. 4. Kayea larnachiana. A. tightly clustered inflorescence, flowers at anthesis. B. fruit. Both from Cooper 2572

and Hawkes (CNS). Photos: T. Hawkes.

between the sheets, and the specimens vary in

phenology, having either flower buds or fruit

(except for MEL 2227333 which has a single

flower). Most of the sheets can be linked to

Mueller through handwriting on original

labels, except for the BM sheet which has no

original label. Its label was clearly added much

later and is typewritten, with no collector

number, and the sheet is not annotated by

Kostermans despite having been cited by him

(Kostermans 1969: 427). Four of the sheets at

MEL and BRI (MEL 2227332, MEL 2227333,

MEL 2227334, BRI [AQ0166459]) have the

same collector number ‘147A’. One sheet in

MEL has the collector number *143’ (MEL

2221339).

Despite these factors, all these specimens

are likely to be part of the one collection,

as Sayer appears to have only collected in

Queensland in 1886-1887 (JSTOR 2021;

ANBG 2021) and made one visit to the

Mossman area after leaving Cairns on 30

Sept. 1886 with W.W. Froggatt (Anon. 1886:

2). They set up camp for about four weeks

on the Mossman River where they collected

specimens (Froggatt 1887: 1). Froggatt

departed in late October leaving Sayer who

had the intention of “staying a few weeks

longer” (Froggatt 1887: 1), which suggests that

the collection were made during October and

November 1886. It is not possible (although

rather unlikely) to rule out the possibility that

the specimens were collected from different

plants or on different days.

The citation by Kostermans (1969: 427)

of “typus: W. Sayers [as Sayer] (BM), Trinity

Bay, Queensland” appears to be an effective,

though possibly inadvertent, lectotypification

under Art. 7.11 of the International Code of

Nomenclature for algae, fungi, and plants

(Turland et al. 2018). The lectotype sheet (BM

000611321) is a rather unfortunate choice due

to the absence of handwritten and original

labels, but we can see no grounds to supersede

it. The remaining original specimens are

herein recognised as possible 1solectotypes.

Distribution and habitat: Kayea larnachiana

is known from mesophyll and notophyll

rainforest on granite derived substrates 1n the

Daintree National Park in the Mossman River

and Cooper Creek areas (Map 1) at altitudes

from near sea level to 100 m.

Phenology: Flowers have been recorded in

November and December; fruit has been

recorded in from Novermber to January.

Cooper & Zich, Kayea in Australia

Conservation status: Kayea larnachiana 1s

known from two main areas that are disjunct

in the Daintree National Park, along the

Mossman River and along Cooper Creek.

Much of the lowland vegetation outside of

the National Park in the Mossman area has

been cleared, and most of the remaining

potential habitat in remote areas of the park

remain poorly explored. The size of the

known subpopulations has not been assessed

extensively, but they are restricted in area

and only recorded within the National Park.

There are no immediate threats evident for

the species. Using GeoCat (Bachman ef al.

2011) based on known locations, Extent of

Occurrence is estimated at 131 km’, and

Area of Occupancy is estimated at 40 km’.

A suggested conservation status for Kayea

larnachiana is Vulnerable [VU Blabd,

111)+B2abQ, 111)]| (IUCN 2012) and the species

is currently listed as this under the Queensland

Nature Conservation Act 1992.

Etymology: This species was named in

honour of James McDonald Larnach (1837-—

1887), sometimes given as ‘James Macdonald

Larnach’ (Anon. 1887: 6), one of the founders

of the Historical Society of Australasia

and member of the council of the Royal

Geographical Society (Victoria).

Acknowledgments

Tim Hawkes, Rigel Jensen, Rupert Russell,

Terry Carmichael, Andrew Ford, and Trixie

Tuck are thanked for able assistance with

collections; Peter Stevens for advice on

Kayea morphology; Tim Hawkes and Rigel

Jensen for photos; Terry Nolan (© CSIRO)

for the illustration and Steve Murphy for

creating the map. We thank Anna Monro

at CANB for advice on nomenclature and

typification. Collection Managers at CANB,

MEL and BRI are thanked for access to

specimens on loan and to images on the

Australasian Virtual Herbarium and Global

Plants JSTOR. Permits to collect were issued

to the Australian Tropical Herbarium by the

Queensland Department of Environment and

Science or its precursors.

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»145_5° 145.75°

ine

TRIBULATION

CAIRNS 4.~- »

MALANDA

Map 1. Distribution of Kayea species. A K. concinna,

® K. larnachiana and @ K. meridionalis.

Thysanotus admirabilis Jian Wang ter (Laxmanniaceae), a

remarkable new species from western Queensland, Australia

Jian Wang & Jenny Silcock

Summary

Wang J. & Silcock J. (2022). Thysanotus admirabilis Jian Wang ter (Laxmanniaceae), a remarkable

new species from western Queensland, Australia. Austrobaileya 12: 14-18. Thysanotus admirabilis

Jian Wang ter is described and is distinguished by the lanate flower buds and perianth, and sometimes

by the lanate bracts. The new species is known only from Mariala National Park and adjacent areas of

the neighbouring grazing property ‘Varna’, western Queensland. Notes are provided on distribution,

habitat, phenology and affinities of the newly described species and its conservation status discussed.

A key to Thysanotus species in Queensland is provided.

Key Words: Laxmanniaceae; Thysanotus; Thysanotus admirabilis; Australia flora; Queensland flora;

taxonomy; new species; identification key; conservtion status

J. Wang, Queensland Herbarium, Department of Environment and Science, Brisbane Botanic

Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: jian.wang@des.qld.gov.

J. Silcock, School of Biological Sciences, University of Queensland, St Lucia, 4072, Australia. Email:

j.silcock@uq.edu.au

Introduction

The genus 7hysanotus R.Br., commonly

known as fringe lilies, includes 54 species

and one subspecies to date (CHAH 2020;

Govaerts ef al. 2022). Thysanotus has had

various family placements over the past

decades, including Liliaceae (Brittan 1981,

1987), Anthericaceae (Chase eft al. 1995)

and Laxmanniaceae (Chase & Stevens 1998;

Wang 2021). The genus is more often placed

in the family Asparagaceae under subfamily

Lomandroideae as recently circumscribed

(Chase eft al. 2009; Gunn ef al. 2020;

Macfarlane et al. 2020; Govaerts et al. 2022).

The family Laxmanniaceae 1s retained here

following the family systematics applied at

BRI.

Thysanotus is mainly found in Australia,

with two species (1. banksii R.Br. and T.

chinensis Benth.) extending to New Guinea.

The latter of these is also distributed in China,

Thailand, Vietnam, the Malay Peninsula, the

Philippines, Lesser Sunda Islands, Celebes

and Aru Island (Brittan 1981, 1987).

Currently, there are three species and

a subspecies recognised as occurring in

Queensland, namely: 7Thysanotus banksii, T.

chinensis, T. tuberosus R.Br. subsp. tuberosus

and 7! tuberosus subsp. parviflorus (Benth.)

Brittan (Wang 2019). A taxonomic review

of Thysanotus in Queensland north-eastern

Australia by the senior author and P. Forster

is underway.

The first collection of the taxon described

here, with flower buds and near mature

flowers, was made by C. Pennay and S.

Hudson who were conducting flora surveys

for the Queensland Spatial BioCondition

project in June 2019. Their specimen was

brought to the senior author’s attention.

Subsequently, Tracy Wattz, the Queensland

Parks and Wildlife Service and Parnerships

ranger of Mariala National Park, collected

specimens in October 2021 on our request.

Two months later, J. Silcock and R. Fairfax

collected the type specimen and made a

preliminary conservation investigation of the

new species.

Accepted for publication 10 May 2022, published online 29 June 2022

Wang & Silcock, Thysanotus admirabilis

Materials and methods

This study is based on morphological

examination of 7hysanotus material from the

following herbaria: BRI, CANB, CNS, DNA,

MEL, NE, NT, PERTH. All measurements

are based on dried material, except the

dimensions of florets, which are based on

material reconstituted with boiling water.

Dimensions of measurements are inclusive,

1.e. 1.Q—1.8 1s given as I-1.8.

Taxonomy

Key to Thysanotus species in Queensland

1 Roots fibrous, rhizome small; inflorescence aterminal umbel. ......... T. chinensis

. Roots tuberous, tubers cylindrical to ellipsoidal; inflorescence simple to

paniculate-wath few-to more than:20 umbels... 2... aoa ee ee ee el See ee 2

Flower buds, perianth at anthesis and sometimes bracts lanate ....... T. admirabilis

. Flower buds, perianth at anthesis and bracts glabrous. ................04. 3

Inflorescence simple or with |I-—3 branches; antisepaline anthers bright

yellow with light purple or violet tips, 2—-2.8 mm long. ............. T. banksii

. Inflorescence paniculate and usually much branched; antisepaline anthers

light purple, purple or violet with white to pale yellow base, 4-5 mm long. .T. tuberosus

Thysanotus admirabilis Jian Wang ter sp.

nov.

With affinity to 7: tuberosus R.Br., but

differing in the lanate flower buds, perianth

and sometimes bracts; yellow anthers with

creamy to light purple or purple tips; obovoid

and longer seeds. Differing from all other

species of the genus in the lanate flower buds

and perianth and sometimes the lanate bracts.

Typus: Australia. Queensland. WARREGO

District: Mariala National Park, off eastern

boundary, 16 December 2021, J Silcock & R.

Fairfax JLS2332 (holo: BRI [AQ952631]).

Perennial geophytic herb, tufted and clumping;

rootstock tuberous, tubers cylindrical to

ellipsoidal, 1.6-2.5 cm long, 0.5—-l1 cm

diameter, 20-30 cm from stem base. Leaves

20—40, annual, basal, linear, + terete towards

apex, becoming channelled below with narrow

membranous wings, 250-370 mm long, 0.5—1

mm wide, longitudinally veined when dry,

glabrous; leaf sheath margins membranous,

entire, 50-80 mm from the base, 0.4—1 mm

wide; leaf apex obtuse. Each plant usually

has 2 or 3(—6) inflorescences. Inflorescence

paniculate with 6-10 branches, 26—41 cm

long; branched 7-25 cm from the base,

branches | or 2(—4) per node, 3.5—18 cm long,

ascending; axes (stems and branches) terete or

slightly flattened-terete in lower part, ridged

with the ridges becoming more prominent

distally. Umbels more than 20, terminal on

branches and stems; bracts of branches long-

to short-deltoid, (2—)5—33 mm long, 1—2.5 mm

wide at base, green with a purplish to whitish

membranous margin, glabrous to minutely

hairy; bracts of umbels short-deltoid, 3—4

mm long, 1.5—2 mm wide at base, purplish

with a wide and whitish membranous margin,

minutely to densely lanate hairy. Flowers 1-3,

of different ages for each umbel, occasionally

sessile ones below; umbels on old stems with

1 or 2 pedicel remnants; pedicels 7—16 cm long

at anthesis, articulate 1.5—9 mm (less than one

fifth to slightly more than halfway) from the

base, mostly erect in flower and fruit. Flower

buds lanate hairy. Perianth 13-15 mm long,

excluding fringes. Sepals lanceolate, 1.8-

2.2 mm wide, purple with creamy margins

on upper surface, light green, and covered

with lanate hairs on the under surface, not

obviously veined, the apex obtuse, apiculate,

mucro c. | mm long. Petals broad ovate to

elliptic, 1O—-14 mm wide including fringes, the

marginal fringe well-developed, 3.8—4.4 mm

long, dense, of even length throughout except

much shorter at base of petal, overall purple,

with a darker purplish central strip on upper

surface. Stamens 6, surrounding ovary, the

outer (antisepaline) and inner (antipetaline)

whorls similar in colour and shape, but

unequal in length; filaments 1.6—2 mm long;

outer anthers, upright, 5—5.5 mm long, 0.4—

0.5 mm wide, yellow with creamy to light

purple tips, the pores c. 0.5 mm long; inner

anthers, lanceolate, slightly curving inward,

8.5—9 mm long, 0.4—0.5 mm wide at the base,

yellow with creamy to light purple or purple

tips, the pores c. 0.4 mm long; both outer and

inner anthers more or less twisted. Ovary

spherical, 1.2—-1.3 mm diameter, pale with 2

ovules per locule; style terminal, 8.5—9 mm

long; light purple, straight, bent to one side

near tip, well-separated from the stamens,

the stigma facing the anther apices. Capsule

+ spherical or obovoid, 6—6.5 mm long, 5—6

mm wide, enclosed within persistent perianth

segments adhering above. Seeds obovoid with

a narrow end, 2.5—2.6 mm long, 1.4—1.6 mm

wide, black, partially covered with an orange

aril, c. 2 mm long, 1.5 mm wide. Fig. 1.

Additional specimens examined: Queensland.

WARREGO District: Mariala National Park,

approximately 100 m west from eastern side boundary

track, Jun 2019, Pennay & Hudson CPS&05 (BRI);

Mariala National Park, off eastern boundary, Oct 2021,

Wattz s.n. (BRI [AQ952630]).

Distribution and habitat: = Thysanotus

admirabilis is endemic to the Warrego

Pastoral District which lies in the Mulga Lands

biogeographic region of western Queensland

(DAWE 2012). Currently, itis known only from

Mariala National Park and adjacent areas of

the neighbouring grazing property ‘Varna’. It

grows in water run-on areas of mulga (Acacia

aneura F.Muell. ex Benth.) low woodland

with scattered poplar box (£ucalyptus

populnea F.Muell.) and shrubs including

silver turkey-bush (Eremophila bowmanii

F.Muell.) and emu bush (£. longifolia (R.Br.)

F.Muell.) on reddish brown earth (Fig. 1d).

Associated species include Aristida contorta

F.Muell., Brachyscome ciliaris (Labill.) Less.,

Calandrinia eremaea Ewart, Convolvulus

clementii Domin, Digitaria brownii (Roem.

& Schult.) Hughes, Dysphania glomulifera

(Nees) Paul G.Wilson, Erodium crinitum

Carolin, Fimbristylis dichotoma (1L.) Vahl,

Goodenia glabrata (Carolin) K.A.Sheph., G.

havilandii Maiden & Betche, Enneapogon

Austrobaileya 12: 14-18 (2022)

polyphyllus (Domin) N.T.Burb., Euphorbia

boophthona C.A.Gardner, Monachather

paradoxus Steud., Paspalidium rarum (R.Br.)

Hughes, Phyllanthus virgatus G.Forst.,

Polygala_ triflora L., Ptilotus modestus

T.Hammer, Stenopetalum nutans F.Muell.,

Trachymene cyanantha Boyland and Tripogon

loliiformis (F.Muell.) C.E.Hubb.

Phenology: Flowering was recorded in June

and October to December, and mature fruit 1n

December. The June 2019 collection followed

above-average autumn rainfall, while the

2021 collections followed average winter

rainfall and above-average spring rainfall

BOM (2022).

Notes: Thysanotus admirabilis 1s the most

outstanding species in this genus on account

of its lanate hairy flower buds and perianth,

together with sometimes lanate hairy

bracts. These characteristics makes it easily

distinguishable. Thysanotus tuberosus has

also been recorded from Mariala National

Park within 2 km of ZT. admirabilis, but it

was typically found on harder, stonier mulga

areas. Thysanotus admirabilis can be easily

distinguished from 7! tuberosus by, apart from

the lanate hairy flower buds and perianth and

sometimes the hairy lanate bracts, its leafy

habit and yellow anthers with creamy to light

purple or purple tips (anthers are light purple,

purple or violet with white to pale yellow base

for 7) tuberosus).

Conservation status: Thysanotus admirabilis

is currently only known from a small area of

Mariala National Park and a neighbouring

erazing property in western Queensland,

Australia. Extent of Occurrence (EOO) and

Area of Occupancy (AOO), as calculated in

the IUCN Geocat program using a 2 km grid

cell, are both 4 km’. The species population

comprises a single subpopulation and location

(IUCN 2019). It occurs patchily for at least |

km along the eastern boundary of Mariala

National Park on both sides of the fence.

Average density was conservatively estimated

as 50 plants/ha in December 2021, thus within

the 100 ha (1 km’) over which the species

is known to occur, total population size is

conservatively estimated as 5000 plants.

Wang & Silcock, Thysanotus admirabilis

‘4

| See

Hy ee Ce ‘a (ea ade me?

S| AS Bl aa ia ih Bale |

AIS ge Baa

Fig. 1. Thysanotus admirabilis. A. face view of flower, together with buds. B. side view of flower, together with buds.

C. plant habit. D. plants in habitat. All at the type locality. Photos: J. Silcock.

The habitat is in good condition and

has never been cleared. No weeds were

present at the site. Cattle were present on the

grazing property in December 2021, but no

Thysanotus admirabilis plants were grazed.

One plant on Mariala National Park had been

lightly grazed, apparently by a macropod.

Despite its very small known EOO, AOO

and population size, the species is not eligible

for listing under IUCN criteria B or C, as

there is no evidence of continuing decline in

any population or habitat parameters (IUCN

2019). Total population size is estimated to

be >3000 mature individuals, so the species

is not eligible for listing under criterion D1.

As there are no obvious threats that could

conceivably drive the species to CR or EX in

a very short time, which is needed to list as

Vulnerable under criterion D2, a conservation

listing of Least Concern is recommended

under the IUCN (2019) guidelines. Further

targeted surveys across Mariala National Park

and surrounding areas after above-average

rainfall are warranted. Mariala National

Park is relatively well-collected botanically;

however, the species 1s geophytic and cryptic

when not in flower. It is probably only present

above ground for short time periods following

rain.

Etymology: The specific epithet 1s derived

from the Latin for ‘admirable’, worthy

of admiration. It refers to the surprising

and astonishing appearance of the flower

perianth, which is covered with white hairs.

This characteristic distinguishes the species

from all others 1n the genus.

Acknowledgements

We are grateful to the following staff at

the Queensland Herbarium who helped in

the preparation of this manuscript: Chris

Pennay and Shannon Hudson first sighted

and collected a specimen of this little beauty,

Teresa Eyre for project management and

support, Tony Bean and Mark Edginton for

comments and discussion. Tracy Wattz from

Queensland Parks and Wildlife Service and

Partnership 1s thanked for her special trip and

collections on request. We also wish to thank

the Directors of BRI, CANB, CNS, DNA,

MEL, NE, NT and PERTH for providing loan

specimens.

References

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bom.gov.au/qld/charleville/, accessed 1 May

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BRITTAN, N.H. (1987). Thysanotus. In A.S. George (ed.),

Flora of Australia 45: 308-339. Australian

Government Publishing Service: Canberra.

(2020). Australian Plant Census. http://

biodiversity.org.au/nsl/servicers/apc, accessed

10 February 2022.

CHASE, M.W. & STEVENS, P.F. (1998). An _ ordinal

classification for families of flowering plants.

Annals of the Missouri Botanical Garden 835:

531-553.

CHASE, M., DUVAL, M.H., HILLs, H.G., CONRAN, J.G.,

Cox, A.V., EGUIARTE, L.E., HARTWELL, J., FAY,

M.F., CADDICK, L., CAMERON, K. & Hoot, S.

(1995). Molecular phylogenetics of Lilianae.

In P.J. Rudall et al. (eds), Monocotyledons:

Systematics and Evolution, pp. 109-137. Royal

Botanic Gardens, Kew: London.

CHAH

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CHASE, M.W.; REVEAL, J.L. & Fay, M.F. (2009). A

subfamilial classification for the expanded

Asparagaceae families | Amaryllidaceae,

Asparagaceae and Xanthorrhoeaceae, Botanical

Journal of the Linnean Society 161: 132—136.

DAwE (2012). Interim Biogeographic Regionalisation

for Australia (IBRA), Version 7 (Subregions)

- States and Territories. Commonwealth of

Australia: Canberra. https://www.environment.

gov.au/fed/catalog/search/resource/details.

page?uuid=%7B1273FBE2-F266-4F3F-895D-

CIE45D77CAF5%7D, accessed 3 May 2022.

GOVAERTS, R., ZONNEVELD, B.J.M. & ZONA, S.A. (2022).

World Checklist of Asparagaceae. Facilitated

by the Royal Botanic Gardens, Kew. http://

wcsp.science.kew.org/, accessed 10 February

2022.

GUNN, B.F., MURPHY, D.J., WALSH, N.G., CONRAN, J.G.,

Pires, J.C., MACFARLANE, I.D. & Bircu, J.L.

(2020). Evolution of Lomandroideae: Multiple

origins of polyploidy and biome occupancy

in Australia. Molecular Phylogenetics and

Evolution 149 (106836): 1-16. https://doi.

org/10.1016/j.ympev.2020.106836

IUCN STANDARDS AND PETITIONS COMMITTEE (2019).

Guidelines for Using the IUCN Red List

Categories and Criteria. Version 14. Prepared

by the Standards and Petitions Committee.

http://www.iucnredlist.org/documents/

RedListGuidelines.pdf.

MACFARLANE, T.D., FRENCH, C.J. & CONRAN, J.G. (2020).

A new Fringe Lily from Kalbarri National Park

(Thysanotus kalbarriensis, Asparagaceae).

Nuytsia 31: 23-27.

WANG, J. (2019). Laxmanniaceae. In G.K. Brown & P.D.

Bostock (eds.), Census of the Queensland Flora

2019. Queensland Department of Environment

and Science: Brisbane. https://www.data.qld.

gov.au/dataset/census-of-the-queensland-

flora-2019, accessed 1] February 2022.

— (2021). Lomandra altior Jian Wang ter and L.

breviscapa Jian Wang ter (Laxmanniaceae),

two new species from the Wet Tropics of north

Queensland, Australia. Austrobaileya 11: 266—

272:

Hibiscus graniticus Wannan (Malvaceae), a

new species from north-east Queensland

Bruce S. Wannan

Summary

Wannan, B.S. (2022). Hibiscus graniticus Wannan (Malvaceae), a new species from north-east

Queensland. Austrobaileya 12: 19-25. The new species Hibiscus graniticus is described and

illustrated. This species has affinities to the geographically widespread H. meraukensis Hochr., but

differs in features of habit, foliage, flowers, fruit and indumentum. Hibiscus graniticus is endemic to

granite habitats between Mareeba and Lakeland Downs in north-east Queensland. The new species is

illustrated with photographs and notes are provided on how it differs from related species.

Key Words: Malvaceae; Hibiscus; Hibiscus section Furcaria; Hibiscus graniticus; Hibiscus

meraukensis; flora of Australia; flora of Queensland; new species

B.S. Wannan, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor

Road, Smithfield, Queensland 4878, Australia. Email: bs;wannan@bigpond.com

Introduction

Hibiscus section Furcaria DC. 1s a speciose

eroup in Australia with more than 30 species

described (Wilson 1974; Wilson & Craven

1995; Wilson 2006; Craven ef al. 2003, 2016;

Badry et al. 2017). One of the most common

and widespread species of the group in

Australia is Hibiscus meraukensis Hochr.,

invariably an annual plant. It 1s also one of

the most morphologically diverse members of

Hibiscus section Furcaria with considerable

variation in foliage and floral characteristics

(Wilson 1974; Ross 1986; Brock 1988;

Wheeler 1992; Kenneally et al. 1996; Milson

2000; Cooper & Cooper 2004; Moore 2005;

Melzer & Plumb 2007; Cowie eft al. 2013;

Hyland et al. 2020). Hibiscus meraukensis was

described from Merauke in New Guinea and

considered as related to H. diversifolius Jacq.

and H. divaricatus Graham (Hochreutiner

1907). In areview of Malesian Malvaceae, the

distribution of H. meraukensis was described

as northern Australia (including the Torres

Strait Islands), southern New Guinea and the

southern Moluccas (Borssum Waalkes 1966).

Hibiscus graniticus Wannan sp. nov.

described below, was recognised during

fieldwork on southern Cape York Peninsula, as

a somewhat glaucous, upright, multistemmed,

mauve flowered perennial shrub, growing on

elevated granite areas. There are few previous

collections of 1. graniticus in herbaria. These

collections have been usually, previously

identified as H. meraukensis to which the new

species 1s superficially similar.

Materials and methods

Recognition of its morphological differences

and the taxonomic novelty of Hibiscus

graniticus have been confirmed from

comparative garden cultivation over the last

10 years at Speewah, alongside examples of

H.. meraukensis from north-east Queensland.

The differences have also been confirmed

by comparison with over 100 specimens in

herbaria of H. meraukensis from Western

Australia, the Northern Territory and northern

Queensland.

The species description and comparisons

with other species from Hibiscus section

Furcaria are based on herbarium specimens

(BRI, CANB, CNS, NT). Examination of

Accepted for publication 21 June 2022, published online 7 September 2022

spirit (author’s own collection) and fresh

material was used for detailed comparison

with H. meraukensis. Photographs of the type

specimens of H. meraukensis lodged at the

National Herbarium of the Netherlands (L

0012968, L 0012969, L 0012970, L 0012971,

L 0012972) were viewed online (https://web.

archive.org/web/20130125042043/http://

vstbol.leidenuniv.nl/).

Length by width measurements are

indicated as length x width mm. Dimensions

are inclusive, 1.e. 1.0—1.7 1s given as |1—1.7. The

terms climax leaves and distal leaves are used

here, sensu Craven ef al. (2003), to describe

the larger, mid-stem leaves and upper, smaller

leaves subtending flowers, respectively. The

term pedicel is used here to describe the stalk

of a flower; however, in some cases, this is

jointed and referred to as an articulation

in species other than that described below,

where it 1s inarticulate (see Table 1). This

does not include the frequent abscission layer

at the base of the pedicel near the subtending

leaf axil.

Taxonomy

Hibiscus graniticus Wannan sp. nov.

Similar to other Australian Hibiscus section

Furcaria but differing by the following

combination of characters: perennial

multistemmed shrub; glaucous foliage;

inarticulate pedicels; absence of stellate or

bifid hairs on foliage; presence of only sparse

aculei with simple hairs on leaves, epicalyx

and calyx and very rarely on stems, petioles

and pedicels; ovary with even covering of

simple hairs; and seeds hight brown with an

orange and white caruncle. Typus: Australia.

Queensland. COOK DISTRICT: Bonny Glen,

Cape York Peninsula, 27 October 2010, B.S.

Wannan 5990 & M. Trenerry (holo: BRI

[AQ880107 comprising a single sheet]; iso:

CNS, NSW distribuendi auctore).

Shrub to 3 m tall, evergreen, often

multistemmed from near ground level;

stems up to 4 cm diameter developing light

brown bark in older plants. Branchlets grey-

glaucous, very rarely with tubercle-based

aculei, glabrous. Stipules subulate, unlobed,

eventually deciduous, glabrous, 3-6 mm

Austrobaileya 12: 19-25 (2022)

long on climax leaves, up to 10 mm long on

distal leaves subtending flowers. Petioles

15-160 mm long on climax leaves, 0-20

mm on distal leaves, mostly glabrous, very

rarely with tubercle-based aculei. Climax

leaf lamina (sensu Craven et al. 2003) ovate

to broadly ovate, 60-150 mm X 60-150 mm,

mostly deeply 3—5-lobed with the lobes longer

than wide, discolorous, glabrous except for

scattered short aculei along midribs on both

surfaces; leaf laminae margins serrate, with

small aculei (<< 0.5 mm) at the apex of each

tooth; base mostly cordate. Distal leaves

simple, up to 75 X 20 mm, pinnately-veined;

discolorous, glabrous except for scattered

short aculei along midribs on both surfaces;

leaf laminae margins deeply serrate, with

small aculei (<< 0.5 mm) at the apex of each

tooth; base mostly cuneate. Foliar nectaries

2-4 mm long, present above the midvein

junction of climax leaves, one per lobe, or at

the base of each pherophyll. Flowers solitary

in axils, chasmogamous, lasting only one day.

Pedicels 13—22(—40) mm long, inarticulate,

glabrous. Epicalyx present, persistent, with

occasional tubercle-based aculei to 1.2 mm,

7—9-segments, 14—17 mm long at anthesis, less

than | mm wide; segments subulate, free to

the base, *4 the length of the calyx at anthesis

and in fruit, straight or incurved. Calyx at

anthesis not splitting, 5-lobed with the lower

Ya fused, not adnate to the corolla and not

falling with it after anthesis, green, abaxially

with tubercle-based aculei to 1.5 mm on

midrib and thickened margins, adaxially with

very fine simple hairs near lobe margins, lobes

triangular with prominent marginal ribs and

midrib; apex acute, (16—)19—22 mm long at

anthesis, nectary absent. Petals obovate, free

to base, 40—80 mm long, pink with maroon

basal spots and with simple hairs to 1.5 mm

at anthesis, with fine simple hairs to 0.5 mm

in bud. Staminal column straight, 5-toothed

at the apex, c. 17 mm long, with capitate

light maroon pollen clusters to 2 mm, up to

the apex of the staminal column. Style c. 27

mim long, extending 10 mm beyond the end of

the staminal column, 5-branched each with a

capitate stigma that 1s covered by fine maroon

hairs to 0.7 mm. Ovary 10—14 mm long, with

simple hairs to 1.5 mm distributed evenly over

Wannan, Hibiscus graniticus

ovary. Epicalyx in fruit up to 19 mm. Calyx in

fruit not distinctly inflated but slightly longer

than epicalyx, up to 28 mm. Fruits capsular,

dry-dehiscent, ovoid, 10-14 mm long, hairs

simple; dehisced capsules with attenuate

apices. Seeds trigonous reniform, up to 4.5 X

3.5 mm, light brown and with an orange and

white caruncle. Figs. 1-8, Table 1.

Additional selected specimens examined: Queensland.

Cook District: Near Mount Elephant, NW of Carbine,

Apr 2008, Wannan 5126 (BRI, CNS); Bobs Lookout

on the Desailly Range, Jun 1985, Clarkson 5950

(BRI, CANB, CNS, DNA); Ex horto Speewah (from

BSW35990), May 2012, Wannan 6426 (CNS); ibid, Jun

2017, Wannan 6932 (CNS); ibid, May 2020, Wannan

7043 (CANB, MEL); Footslope of Hann Tableland, Jul

2002, Fox IDF1640 (BRI); Ridge of Hann Tableland, Jul

2002, Fox IDF1653 (BRI).

Distribution and habitat: Hibiscus graniticus

is endemic to the north-east part of the

Einasleigh Uplands bioregion, on the southern

Cape York Peninsula in northern Queensland

where it grows in eucalypt dominated, open

woodland between Mareeba and Lakeland

Downs on Desailly, Kelly St George, Cannibal

Creek and Mareeba granites (Bultitude ef al.

1998).

Phenology: Hibiscus graniticus 1s a perennial

species that has been recorded to flower

mostly between April and August. However,

the type collection was made from a specimen

found flowering in late November 2010,

following 100 mm of rain in early November.

Individuals of H. graniticus survive for up to

5 years 1n cultivation, 1n contrast to the annual

erowth of H. meraukensis.

Notes: Hibiscus. graniticus differs from

examples of H. meraukensis from Australia

and New Guinea, by its inarticulated pedicels,

deeply serrate distal leaves, epicalyx less

than 80% of the calyx length at anthesis (vs

more or less equal, or slightly longer than

calyx at anthesis), absence of stellate hairs,

multistemmed perennial habit (vs annual) and

hairy ovary and capsule (vs mostly glabrous

or sparsely pubescent). Hibiscus graniticus

further differs from the type specimen and

descriptions of H. meraukensis from western

New Guinea (Borssum Waalkes 1966), by

its longer pedicels at anthesis (more than 10

mm long vs. less than 10 mm), longer corolla

(more than 45 mm vs. less than 45 mm),

and fawn brown seeds (rather than black-

brown). Comparison of H. graniticus with

other members of Hibiscus section Furcaria

from adjacent areas of northern Queensland

is Shown in Table 1. It differs from all other

species of this section in the region by its

absence of stellate hairs from all plant parts.

Etymology: The Latin species epithet refers

to the granite substrate on which it occurs.

Fig. 1. Hibiscus graniticus. Stem showing bark (Wannan

6426, CNS). Scale of 20 mm.

Fig. 2. Hibiscus graniticus. Adaxial surface of climax

leat (Wannan 7043, CNS). Scale of 20 mm.

22 Austrobaileya 12: 19-25 (2022)

Fig. 3. Hibiscus graniticus. Abaxial surface of climax

. Fig. 6. Hibiscus graniticus. Flower showing calyx,

wm WHEREAS LEM AMI US SAENO). BOURGES ye del edivel or konts (ania OLI6, CNS), Seale

of 20 mm.

Fig. 4. Hibiscus graniticus. Habit with lower climax Fig. 7. Hibiscus graniticus. Calyx in fruit (Wannan

leaves and flowering shoots with flowers and distal 7043, CNS).

leaves (unvouchered ex horto Speewah grown from

Wannan 5990, BRI, NSW).

one

~ ae ~” Te

_—

moe: nae fens

Fig. 5. Hibiscus graniticus. Flower showing stamens Fig. 8. Hibiscus graniticus. Seeds (Wannan 7043, CNS).

and ovary as well as stipules ex horto (Wannan 6426, Scale of 2 mm.

CNS).

Wannan, Hibiscus graniticus

Acknowledgements

I thank the curators of CNS, BRI, CANB and

NT for loans or access to specimens and for

their assistance 1n obtaining digital images of

specimens. The assistance of Todd McLay in

examining material from CANB 1s gratefully

acknowledged. Thanks also to Frank Zich

from CNS for helping access the online type

images of Hibiscus meraukensis. | would also

like to thank Garry Sankowsky and Bob Jago

for access to their images. Thanks also to Paul

Forster for his advice and encouragement

with this paper.

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Cryptachne E.J.Thomps. (Poaceae:

Panicoideae: Cleistochloinae), a new genus for

Queensland with three new species

E.J. Thompson

Summary

Thompson, E.J (2022). Cryptachne E.J.Thomps. (Poaceae: Panicoideae: Cleistochloinae), a new

genus for Queensland with three new species. Austrobaileya 12: 26—58. The new genus Cryptachne

E.J.Thomps., endemic to Queensland, is delimited and described. Three new species, all formerly

included with phrase names in Dimorphochloa, Cryptachne columboola E.J.Thomps., C. duaringa

E.J.Thomps. and C. trinerva E.J.Thomps., are described with illustrations of both macro and micro

morphology. Cryptachne is closely related to Calyptochloa differing in the tufted growth habit,

racemose terminal inflorescences with spikelets that have a lower lemma subequal to upper lemma,

and axillary spikelets with an elaiosome on the lower lemma. The three new species all have restricted

distributions in habitat types associated with lancewood dominated ironstone jump-ups; they are

suggested to warrant Endangered or Critically Endangered conservation status.

Key Words: Poaceae; Panicoideae; Cleistochloinae; Calyptochloa; Cryptachne; Cryptachne

columboola; Cryptachne duaringa; Cryptachne trinerva; flora of Australia; flora of Queensland;

amphigamy; chasmogamy; cleistogamy; anatomy; elaliosome; micromorphology; spikelet

dimorphism; new genus; new species; taxonomy; identification key

E.J. Thompson, c/ Queensland Herbarium and Biodiversity Science, Department of Environment

and Science, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Email: john.thompson@des.qld.gov.au

Introduction

The form of reproductive dimorphism found

in the Australian endemic grass subtribe

Cleistochloinae E.JThomps. (subfamily

Panicoideae), involves plants having two

types of inflorescences in separate places

(amphigamy), viz. terminal and axillary,

with spikelet dimorphism (Thompson 202Ic,

2022b). Spikelets in the terminal racemes

or reduced spike-like panicles are usually

wholly chasmogamous (flowers in terminal

inflorescences that open to release the stigmas

and anthers and thereby potentially cross-

fertilise) and morphologically differentiated

from spikelets in reduced racemes in the leaf

axils. The axillary spikelets are cleistogamous

where the flowers are obligately self-fertilized

and never open.

Cleistogamy is found in c. 3% of grass

species and c. 10% of the genera worldwide,

and 14% of the species and 30% of genera in

Australia (Campbell eft al. 1983; Thompson

2021c). Conversely, the typical grass has only

chasmogamous flowers. Anther dimorphism

is Shared by all species in Cleistochloinae with

the axillary cleistogamous spikelets having

much smaller anthers than the chasmogamous

ones 1n the terminal inflorescences.

The type of dimorphic reproductive

system found in the Cleistochloinae can also

be found in some North American grasses,

although spikelet morphology is distinctly

different across the genera. Chase (1918:

255) noted that “Often, if their source was

unknown, they would not be placed in the

same tribe’, in reference to the American

genera.

Calyptochloa C.E.Hubb., Cleistochloa

C.E.Hubb., Dimorphochloa S.Y.Blake and

Simonachne E.J.Thomps. comprise — the

Cleistochloinae (Thompson & Fabillo 2021;

Accepted for publication 4 April 2022, published online 9 November 2022

Thompson, Cryptachne

Thompson 2021b, 2021c, 2022b). Molecular

and morphological datasets generated

phylogenetic and phenetic analyses of this

eroup (Thompson & Fabillo 2021; Thompson

2021c). Dimorphochloa appears to be not

monophyletic based on these initial analyses

(Thompson & Fabillo 2021); however, more

extensive molecular sampling is still required

of extra-Australian taxa, before a final

classification can be achieved combining both

datasets. Dimorphochloa rigida S.1.Blake

and three undescribed species that have been

included in Dimorphochloa with phrase

names, represent two groups respectively,

on the basis of multiple morphological

characters.

The undescribed species previously

assigned to Dimorphochloa, have closest

affinity to Calyptochloa (Thompson &

Simon 2012; Thompson & Fabillo 2021;

Thompson 2021b). Calyptochloa differs in

several morphological characters including a

stoloniferous growth habit, relative length of

upper lemma of the chasmogamous spikelets

and the lower lemma of the cleistogamous

spikelets having an elaiosome (Fig. 1). They

also differ from Dimorphochloa rigida, as

that species has a rhizomatous habit and rigid

cane-like culms, absence of a contraligule,

cleistogamous and chasmogamous spikelets

similar, with the former apical on subordinate

leafy branches of culms separate from the

chasmogamous spikelets. By contrast, the

three undescribed species are distinct in

the combination of a tufted growth habit,

the indumentum composition on the upper

glumes and lemmas of the terminal spikelets,

and the lower lemmas of the axillary spikelets

having an elaiosome (Table 1).

The aim of this paper is to describe

a new genus Cryptachne E.J.Thomps. to

accommodate the undescribed species

currently known as Dimorphochloa sp.

(Charters Towers E.J.Thompson+ CHAS554),

D. sp. (Miles E.J Thompson EJT906) and

D. sp. (Mt Cooper R.J-Cumming 18623)

(Thompson 2021la, 2022a). The three species

of Cryptachne have been recorded from

relatively arid, subtropical habitats usually

composed of Acacia woodland (especially

Acacia shirleyi Maiden — lancewood), on hilly

terrain with shallow to skeletal well-drained

soils. They occur 1n eastern Australia over the

latitudinal range 20—26° (Fig. 2, Map 1). All

species occur on landscapes associated with

ironstone jump-ups (Cainozoic duricrusts)

(QRE 2020) and the allied grass Cleistochloa

subjuncea C.E.Hubb. is usually dominant in

the ground layer.

Materials and methods

Observations of leaf, culm and_ spikelet

morphology of the species of Cryptachne

and some superficially similar species of

Cleistochloa and Dimorphochloa were made

using imagery, herbarium specimens and

fresh samples taken from cultivated plants.

Plants were cultivated in pots under nursery

conditions in Brisbane, Australia from 2013—

2021. Nursery stock was initially grown

from ex situ plants and additional stock was

propagated from caryopses and stolons that

readily root at the nodes after rain and humid

weather in summer.

Data used in this paper were obtained

from the 16l-character list and data

matrix provided by Thompson (202Ic).

The categories of characters covered

include growth habit, culms and _ leaves,

inflorescence type, and terminal and

axillary spikelet macromorphology. Spikelet

micromorphology included glumes, paleas,

stigmas and caryopses. Micromorphological

characters also included those of leaves and

culms.

Habitat descriptions are provided 1n terms

of Regional Ecosystems as defined by the

Queensland Government (QRE 2020).

Nomenclature and terminology

Botanical nomenclature follows Brown

(2022).

General botanical terminology relevant

to grasses, follows Harris & Harris (1994),

Beentje (2010) and Thompson (2021b).

Terminology relating to inflorescences and

spikelets follows Tothill & Hacker (1983),

Jacobs et al. (2008), Gibson (2009) and

Thompson (2021c). The spikelet 1s viewed

here as a reduced inflorescence consisting of

28 Austrobaileya 12(1): 26-58 (2022)

5mm

Fig. 1. Elaiosomes on fresh and dry cleistogamous spikelets of taxa in subtribe Cleistochloinae. A—C. Lower lemma of

Cryptachne trinerva. A—-C from Thompson MOR796 (BRI). D—F. Lower lemma and upper glume of Dimorphochloa

rigida. D & E from Thompson MOR475 & Edginton (BRI). F from Thompson EJT958 & Simon (BRI). G—I. Lower

lemma of Cleistochloa subjuncea. G & H from Thompson MOR710 (BRI). | from Thompson HUGSI5 & Simon (BRI).

el fresh on lower lemma; e2 fresh on upper glume; e3 dry on lower lemma.

Thompson, Cryptachne 29

Fig. 2A. Growth habit and habitat of Cryptachne. A. C. trinerva in low woodland of Acacia shirleyi (lancewood)

on lateritic jump-up. B. C. columboola in woodland of Eucalyptus fibrosa on sandstone. Photos: E.J. Thompson.

Cryptachne indicated by black arrow, white arrow indicates the superficially similar Cleistochloa subjuncea.

Austrobaileya 12(1): 26-58 (2022)

Fig. 2C. Growth habit and habitat of Cryptachne duaringa in lancewood woodland on lateritic slope. Photo: E.J.

Thompson. Cryptachne indicated by black arrow.

bracts subtending one or more florets (Tothill

& Hacker 1983; Kellogg 2006; Endress 2010).

Terminology relating to grass anatomy and

micromorphology follows Ellis (1976, 1979),

Watson & Dallwitz (1992) and Dengler ef al.

(1994).

Imagery

Photographs were taken using _ light

microscopes to study anatomical and

micromorphological characters, viz. spikelets

using a Nikon SMZ25 binocular microscope

with Nikon DS-Ril camera. The resultant

Images were viewed using NIS-Elements

BR software (ver. BR 5.11.000 64-bit, USA;

Laboratory Imaging (http://www.lim.cz,

accessed 15 December 2019)). Stigmas and

sections of leaves and culms were examined

using a Leica DMLB compound binocular

microscope fitted with a digital camera

and images were viewed using ToupView

software (ver. x64 4.7.14326.20190401, China;

Touptek; http://wwwtouptek.com, accessed

20 September 2019).

Scanning electron micrographs (SEMs)

were obtained without sputter coating

(Phenom G2 5kev SEM with backscatter

detector).

Data acquisition and classification of

morphological characters

The freehand sectioning method described by

Thompson (2017), and modified from Frohlich

(1984), was used to obtain transverse sections

of leaves and inflorescence culms. For each

species, sections of leaves and culms were

obtained from fresh material of cultivated

plants.

Micromorphology of the abaxial leaf

surface was studied from replicas from fresh

leaves using the method described by Hilu &

Randall (1984) and SEMs.

Anatomy and micromorphology of leaves

and culms were studied with respect to

characters such as stomata, silica bodies and

microhairs (De Wet 1960; Metcalfe 1960;

Twiss et al. 1969; Ellis 1979; Renvoize 1987;

Watson & Dallwitz 1992; Siqueiros-Delgado

& Herrera-Arrieta 1996; Piperno & Pearsall

1998; Krishnan ef al. 2000; Siqueiros-Delgado

2007; Lu et al. 2009; Jattisha & Sabu 2015).

Observations of micromorphology were

made from SEMs of lower lemmas, upper

lemmas and upper paleas. Silica bodies,

stomata, epidermal long cell walls, microhairs

and macrohairs were classified using the

Thompson, Cryptachne

terminology of previous authors (Hsu 1965;

Jirasek & Jozifova 1968; Ellis 1979; Valdes-

Reyna & Hatch 1991; Snow 1996; Acedo &

Llamas 2001; Liu et al. 2010; Mashau et al.

2015; Olonova et al. 2016; Neumann ef al.

2017).

Images of fresh and dry _lodicules

and stigmas were obtained using light

microscopy. Lodicules were classified as

plicate or non-plicate (Hsu 1965; Jirasek

& Jozifova 1968; Guedes & Dupuy 1976).

Stigma macromorphology was classified by

the position of emergence from the spikelet,

Taxonomy

the outline shape of the branching and colour.

Micromorphology was examined _ using

characteristics of the lobes including shape

of apex, relative length and tilt (Thiele eft al.

1996).

Caryopsis and embryo morphology

(including characters relating to hilum,

scutellum and epiblast, embryo, spermaderm

and stylopodium) were observed and

classified with previously used terminology

(Kennedy 1899; Reeder 1957; Brown 1959,

1960; Watson & Dallwitz 1992; Klak 1994;

Kosina 1995; Snow 1998; Liu e¢ al. 2005; Liu

et al. 2015).

Key to the genera of subtribe Cleistochloinae (modified from Thompson (2022b))

1 Spikelets from the two types of inflorescences similar... ............2.0004 2

1. Spikelets from the two types of inflorescences strongly dimorphic. ............ 3

2 Plants stoloniferous; axillary racemes usually comprising four

cleistoOsamous-Spikelels- os 2 we 5 Gk gloee Re ASR Ge es ea Bes Simonachne

2. Plants rhizomatous with wiry, erect tufted culms; cleistogamous spikelets

apical on leafy subordinate branches, usually solitary. .......... Dimorphochloa

3 Upper lemmas awned; cleistogamous spikelets adaxial .................0.. ef

3. Upper lemmas mucronate; cleistogamous spikelets abaxial. .......... Cleistochloa

4 Plants stoloniferous; upper lemma much shorter than lowerlemma..... . Calyptochloa

4. Plants tufted; upper lemma subequal to lowerlemma.............. Cryptachne

Cryptachne E.J.Thomps., gen. nov.

Allied to Calyptochloa C.E.Hubb. and

differing in the tufted growth habit, the

indumentum on the upper glumes and

lemmas of the terminal spikelets, and the

lower lemmas of the axillary spikelets with

an elaiosome. Typus: Cryptachne trinerva

E.J.Thomps.

Tufted perennials sometimes with decumbent

stems rooting at the nodes. Culm with pith.

Internodes appressed pubescent. Leaf sheaths

pilose with simple tuberculate-based hairs,

one margin pilose. Ligule and contraligule a

fringe of hairs. Leaf blades lanceolate, base

truncate, pseudopetiolate, both surfaces

pilose; proximal portion of margins white,

ciliate with simple tuberculate-based hairs,

one margin undulate. Inflorescences of

two types in different parts of the plant

(amphigamous); terminal racemes and

axillary racemes with 1 to 2 spikelets

partly concealed by the leaf sheath. Axes

of inflorescence branches lacking pulvin1.

Pedicels scabrid, straight; apex a shallow cup

with thin walls. Spikelets dimorphic. Spikelets

from terminal inflorescences chasmogamous,

adaxial, slightly dorsally compressed,

elliptical in outline; apical ones longer than

laterals, disarticulating below the glumes.

Glumes unequal; lower glume much reduced;

upper glume elliptical, 5-veined, convex,

cartilaginous, villous in lower two-thirds with

long, straight, mostly appressed, tubercular-

based simple macrohairs, apex truncate;

margin flat hyaline. Lower floret sterile;

lemma 7-veined linear to lanceolate, convex,

cartilaginous, nerves smooth, indumentum

not exceeding the apex of the spikelet,

villous throughout, apex acute, similar in

shape, size and type of indumentum to upper

glume; palea absent. Upper floret bisexual;

upper lemma shorter than to subequal to

lower lemma, 3-veined, chartaceous, body

glabrous, minutely longitudinally ridged with

papillae; margins hyaline, apex with flattened

cilia; awned; germination lid a crescent-

shaped depression; palea obscurely 2-veined,

apex acute, similar texture and surface to

lemmas. Lodicules 2, free, triangular-flag

shaped. Anthers 3. Caryopsis slightly dorsi-

ventrally compressed, loosely clasped by

upper lemma and palea; hilum punctiform.

Spikelets in axillary inflorescences

cleistogamous; spikelets occurring 1n either

of three combinations: (a) sessile to subsessile

or pedicellate, sessile spikelets usually

shorter than the pedicellate ones; (b) paired,

with one sessile and the other pedicellate; (c)

sessile or paired spikelets at nodes, apical one

pedicellate. Pedicels mostly puberulous with

short simple hairs, straight; apex shallow cup-

shaped, scabrid. Spikelets elliptical, adaxial,

smooth, laterally compressed; concealed

within a coriaceous convolute leaf sheath at up

to several culm internodes in succession from

immediately below terminal inflorescence;

culm bowing and exserting from sheath as

spikelet matures with sheath partially gaping,

leaf and then leaf sheath disarticulating at

maturity with spikelet enclosed. Lower glume

absent or vestigial. Upper glume shorter

than the spikelet, elliptic, shallowly convex,

chartaceous, glabrous except at base, 3-veined,

apex obtuse. Lower floret sterile; boat shaped,

two-keeled, cartilaginous, glabrous, 5-veined,

apex truncate, basal elaiosome present;

palea absent. Upper floret bisexual; lemma

lanceolate, convolute, cartilaginous, base

hirsute, 5—7-veined, apex acute to attenuate,

awned; palea much shorter than lemmas,

convolute, cartilaginous, 4-veined, glabrous,

apex acute to attenuate. Anthers 3. Caryopsis

ellipsoid, slightly dorsi-ventrally compressed;

scutellum slightly less than half the length

of the caryopsis; hilum elliptical. Pedicellate

spikelet similar to sessile spikelet but

longer, adaxial; exposed at maturity; pedicel

puberulous. Caryopsis slightly dorsi-ventrally

Austrobaileya 12(1): 26-58 (2022)

compressed, loosely clasped by upper lemma

and palea, larger than chasmogamous; hilum

punctiform.

Three species; endemic to southern to central,

subcoastal areas of eastern Queensland.

Notes: Cryptachne differs from the other

genera in Cleistochloinae by a combination

of morphological characters (Table 1). The

erowth habit of all species of Cryptachne 1s

tufted and lacking rhizomes, but in summer

during humid, rainy periods, plants produce

decumbent stems that root at the nodes. In

the field, plants resemble the tufted habit of

Cleistochloa as opposed to the stoloniferous

form of Calyptochloa and Simonachne.

Caryopses have only been seen in

cultivated plants.

Fresh lodicules of Cryptachne are similar

to those in other taxa of Cleistochloinae with

asymmetrical lobes. Dry lodicules broadly

resemble the fresh ones, although reduced in

size (Fig. 3).

Etymology: The genus name is derived

from the Greek kryptos (hidden) and achne

(scale) in reference to the clandestine axillary

spikelets enveloped by the leaf sheaths.

Micromorphology

Stigma and anther morphology: Stigmas

emerge midway along the spikelet. Stigma

lobes are relatively long for species in

Cleistochloinae. The lobes are appressed to

the stigma branch axis with narrow rounded

apices. Cleistogamous anthers are about half

the length of the chasmogamous ones (Fig. 4).

Micromorphology of the lemmas and palea:

Upper lemmas and paleas of cleistogamous

spikelets with “beaked type” to truncate-

shaped papillae, irregularly spaced between

minute narrow ridges. Upper lemmas and

paleas of chasmogamous spikelets with

irregularly spaced dome-shaped papillae,

minute ridges absent (Fig. 5).

Abaxial leaf blade epidermis: Costal/

intercostal zonation conspicuous. Papillae

absent. Costal long cells rectangular;

anticlinal walls of intercostal long cells

()-shaped. Anticlinal walls of intercostal long

Thompson, Cryptachne

Fig. 3. Lodicules of taxa within Cleistochloinae demonstrating variation, Cleistochloa subjuncea: A & B. fresh,

in situ. Cryptachne duaringa: C. fresh, in situ. D. dry, ex situ. Calyptochloa johnsoniana: E. fresh, ex situ.

Dimorphochloa rigida: F. fresh, ex situ. A & B from Thompson MOR710 (BRI). C & D from Thompson MORS822

(BRI). E from Thompson MOR799 (BRI). F from Thompson MOR762 (BRI). Photos: E.J. Thompson. a anther; f

filament; o ovary; p palea.

cells moderately undulating, often irregular

with short wave-length (cf Ellis 1979).

Stomata 33-36 um long with parallel-sided

subsidiaries, in single rows separated by 6—8

files of long cells. Bicellular microhairs 50-—

70 um long, proximal cell longer than distal,

common. Silica bodies in 3—4 rows, bilobate,

14—20 um long, abundant. Hooks present.

(Figs. 6 & 7).

Transverse section of leaf blade: C,; XyMS+.

Mesophyll without radiate chlorenchyma;

adaxial palisade present. Mlidrib not

prominent; with a double bundle sheath; outer

complete ring of parenchyma cells and partial

inner ring of thick-walled cells with adaxial

arc of clear parenchyma cells. Bulliform cells

in discrete regular groups, in simple fans.

Sclerenchyma accompanying all vascular

bundles as adaxial strands and abaxial

cirders. (Fig. 8).

Austrobaileya 12(1): 26-58 (2022)

Fig. 4. Anthers and stigmas of Cryptachne columboola. A, B, D & F. from chasmogamous (CH) spikelet from terminal

raceme. C, E & G. from cleistogamous (CL) axillary spikelet. A. adaxial view showing stigmas emerging mid-way

along spikelet. B. upper floret showing CH anthers. C. CL anthers. D. CH stigma. E. CL stigma. F. CH stigma. G. CL

stigma. All from Thompson EJT906 et al. (BRI). Photos: E.J. Thompson.

Transverse section of culm: Culm examined

c. 0.4 mm in diameter. Vascular bundles

with a ring of clear parenchyma; three sizes

in separate circles, smallest to the periphery.

Outer smallest vascular bundles adjacent to

radial girder sclerenchyma and imbedded in

large-celled sclerenchyma. Chlorenchyma in

rectangular blocks, 2—4 cells deep by up to 12

cells wide; cells with regular size and shape,

slightly radially oblong. Inner ground tissue

consisting of large thin-walled cells (Fig. 9).

Surface of inflorescence culm: Hispid with

simple hairs to 0.2 mm long, muriculate with

hooks and scabridulous with prickles. Stomata

frequent, similar to those on the abaxial leaf

surface. Bicellar microhairs, 50—70 um long,

common. Silica bodies absent. (Fig. 10).

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Austrobaileya 12(1):; 26-58 (2022)

intercostal

costal zone

50 um

Fig. 6. Scanning electron micrograph of abaxial leaf surface of Cryptachne trinerva. From Thompson CHA555

& Turpin (BRI). Micrograph (captured at X1000): E.J. Thompson. le anticlinal walls of long cells; bm bicellular

microhair; sb silica body — bilobate type; S stomata; sh simple hair; tbsh tuberculate based simple hair.

1. Cryptachne trinerva E.J-Thomps., sp.

nov.

Culms to 80 cm tall. Apical spikelets in

terminal inflorescences mostly > 5 mm

long; axillary spikelets mostly single and

sessile, > 7 mm long. Spikelets in terminal

inflorescences with ascending hairs to 1.2 mm

long. Typus: Queensland. NORTH KENNEDY

District: 12 km W of Charters Towers near

edge of Flinders Highway, 30 March 2011,

E.J. Thompson CHA763, B.K. Simon & M.

Edginton (holo: BRI [AQ863142, comprising

1 sheet]).

Calyptochloa sp. (Charters ‘Towers

E.J.Thompson+ CHAS554): Simon ef al.

(2007); Thompson & Simon (2012: 636).

Dimorphochloa sp. (Charters ‘Towers

E.JThompson+ CHAS554): Simon &

Thompson (2013); Thompson (2021a, 2022a).

Perennial grass; culms 50—80 cm tall, clumps

spreading to 100 cm across, copiously

branched. Culm internodes with hairs to |

mm long. Sheaths with hairs to 4 mm long.

Ligule c. 0.3 mm long. Contraligule c. 0.2

mm. Leaf blades at mid-culm 20-50 mm

long, 1.5—3.5 mm wide, hairs to c. 4 mm long.

Terminal inflorescence: axes 20—50 cm long,

5—9-flowered. Apical spikelets 4.9-5.6 mm

long (without awn), 0.8—1 mm wide, lateral

spikelets 4.3—4.7 mm long; apical pedicels

3—7 mm long, lateral pedicels 0.2—1 mm long.

Lower glume to 0.2 mm long. Upper glume

Thompson, Cryptachne

4.1-5.6 mm long. Lower lemma 4.3—5.6 mm

long; pilose, hairs ascending to 1.2 mm long.

Upper lemma 3.5—3.7 mm long, awn 0.7—1.5

mm long. Lodicule c. 0.3 mm long. Anthers

2.5—-2.9 mm long. Axillary inflorescence:

most commonly occurring as single sessile

spikelet 7.3-—8.5 mm long (without awn),

Q.8—1.1 mm wide. Upper glume 5.6—6 mm

long. Lower lemma 7.3—8.5 mm long, 0.7-1.1

mm wide. Upper lemma 5.6—6.8 mm long,

awn |I—1.7 mm long; palea 5.3—5.7 mm long.

Single pedicellate spikelet: 6—6.8 mm long.

Paired spikelets: sessile, 5.9-6.7 mm long;

pedicellate, 6.9—7.5 mm long (without awn),

pedicels 9-22 mm long. Anthers c. 0.8 mm

long. Caryopsis not seen. Figs. 11 & 12.

Additional specimens examined: Queensland. NORTH

KENNEDY District: 11 km SW of Charters Towers

on edge of Flinders Highway, Mar 2002, Thompson

CHA)555 & Turpin (BRI); 11.6 km W of Charters Towers

near edge of Flinders Highway, Mar 2011, Thompson

CHA77I, Simon & Edginton (BRI); 12 km W of Charters

Towers, Apr 2000, Thompson CHA554 & Thomas (BRI);

ibid, Mar 2012, Thompson CHA792 & Simon (BRI);

ibid, May 2013, Thompson CHA8&27 & Simon (BRI).

Cultivated. Ashgrove, Feb 2016, Thompson MOR796

(BRI); ibid, Apr 2017, Thompson MORS/S (BRI).

Distribution and habitat: Cryptachne

trinerva 1s known from only a few localities

near Charters Towers (Map 1). It grows in

woodland of Acacia shirleyi or A. catenulata

C.T.White, on slopes of jump-ups or “stony

rises’, often dominating the ground cover

with Cleistochloa subjuncea usually present.

Associated shrubs include Erythroxylon

australe F.Muell., Beyeria viscosa Mig. and

Prostanthera leichhardtii Benth. Regional

Ecosystems represented include 10.7.3a and b,

and 11.7.2 (QRE 2020).

Phenology: Flowering December to June.

Etymology: Named after the lower lemma

commonly being 3-veined.

Conservation status: Cryptachne trinerva 1s

known from several, small and fragmented

subpopulations in a very restricted landscape

type. These are threatened by clearing for

pasture development, quarrying and road

works. A suggested conservation status for

the species is Endangered based on criterion

B2a (IUCN 2019) and a formal conservation

status nomination will be made elsewhere.

2. Cryptachne columboola E.J.Thomps., sp.

nov.

Culms to 80 cm tall. Apical spikelets in

terminal inflorescences mostly c. < 5 mm

long; axillary spikelets mostly pedicellate

only, < 7 mm long. Spikelets in terminal

inflorescences with appressed hairs to 0.8

mm long. Typus: Queensland. DARLING

Downs District: 23 km NE of Miles, 10 km

NNE of Columboola, 26 March 2013, £../.

Thompson EJT906 & B.K. Simon (holo: BRI

[AQ1021906, comprising 2 sheets]).

Dimorphochloa sp. (Miles E.J.Thompson

EJT906): Bostock & Holland (2014);

Thompson (2021a, 2021c, 2022a).

Perennial grass; culms 50-80 cm. tall,

copiously branched, clumps spreading to 50

cm across. Culm internodes with hairs to |

mm long. Sheaths with hairs to 3.5 mm long.

Ligule 0.3—0.4 mm long. Contraligule c. 0.1

mm long. Leaf blades at mid-culm 20—55 mm

long, 1.5—3.1 mm wide, hairs to 1.1 mm long;

proximal margins with hairs to 3.8 mm long.

Terminal inflorescence: axes 20-30 mm

long, 5—9-flowered. Spikelets 3.3-—4.8 mm

long (without awn), 1-1.2 mm wide; apical

pedicels 2.5—3.6 mm long, lateral pedicels 0.2—

0.6 mm long. Lower glume to 0.2 mm long.

Upper glume 3.5—4.8 mm long. Lower lemma

3.5—4.8 mm long; hairs sparse, appressed, to

0.8 mm long. Upper lemma 2.9—3.4 mm long,

awn c. 0.7 mm long. Upper palea 2.7-3.3 mm

long. Lodicule c. 0.3 mm long. Anthers 2-3

mm long. Axillary inflorescence: Spikelets

mostly present as pedicellate only; up to 5

per culm; 4.6—6.6 mm long, c. 0.8 mm wide.

Upper glume 4.2—4.6 mm long. Lower lemma

5—6.6 mm long. Upper lemma 5—6.6 mm

long, awn to 0.7 mm long; palea 4—4.7 mm

long. Solitary subsessile to sessile spikelet

rarely present, 1f present then 4.3—4.5 mm

long. Paired spikelets: sessile 5.4-5.9 mm

long; pedicellate 6.1-6.4 mm long, pedicels

1-10 mm long. Anthers 0.5—0.8 mm long.

Caryopsis not seen. Figs. 13 & 14.

Additional specimens examined: Queensland.

DARLING Downs District: 23 km NE of Miles, Jun

2012, Thompson EJTSS&8 (BRI). Cultivated. Ashgrove,

Apr 2017, Thompson MORS20 (BRI); ibid, Apr 2017,

Thompson MORS821 (BRI).

38 Austrobaileya 12(1): 26-58 (2022)

. [=e

SEN Sep sey vd SS PNET igins Tes 2 .

ae ot s Dey ; my Aes bast bk ademas

7 —-— - ~— : sn el — , — — - _ , a

oan intercostal

zone

MY ee Pe

<s*,. 950 ~~

costal zone

——— Pre ee

b—¢

~ -_ -—< ss

as?

Snes Seas

—~- 2 ; a™ has

: > rn ; 7

Fig. 7. Replica of abaxial leaf surface of Cryptachne trinerva. From Thompson MOR796 (BRI). Photo: E.J. Thompson.

bm bicellular microhair; h hook; sb silica body — bilobate type; S stomata; tbsh tuberculate-based simple hair.

gs thsh mid vein _‘'bs bs tertiary vein

100 um

Fig. 8. Transverse section of fresh leaf at mid vein of Cryptachne trinerva. From Thompson MOR796 (BRI). Photo

(captured at X20): E.J. Thompson. be bulliform cell; chl chlorenchyma; gs girder sclerenchyma; ibs inner bundle

sheath — inverted ‘horse-shoe’ of thick-walled cells with adaxial clear cells; obs outer bundle sheath — ‘horse-shoe’ of

clear cells with abaxial thick-walled cells; ss strand sclerenchyma; tbsh tuberculate-based simple hair.

Thompson, Cryptachne 39

Fig. 9. Transverse section of fresh fertile culm of Cryptachne trinerva. From Thompson MOR796 (BRI). Photo

(captured at X20): E.J. Thompson. ee clear cells; chl chlorenchyma; h hook; p prickle; sel radial sclerenchyma; sh

simple hair; vascular bundles: 1 primary; 2 secondary; 3 tertiary.

100 um

Fig. 10. SEM micrograph of surface of terminal inflorescence culm of Cryptachne trinerva. Thompson CHA555 &

Turpin (BRI). Micrograph (captured at X500): E.J. Thompson. bm bicellular microhair; h hook; p prickle; S stomata;

sh simple hair.

Queensland Herbarium (BRI)

Australia: Queensland North Kennedy

Dimorphochloa sp. (Charters Towers E.J.Thompson+

CHA554)

Coll.: Thompson, E.J.; Simon, B.K. 1 May 2012

Coll. no.: CHA79?

Lat.. 20° 7' 16.100"S Long.: 146° 9' 20.100"E

Woodland of Acacia shirleyi on lateritic jump-up. og

Dominant decumbent perennial grass ascending to 50 cm

tall,

Wet 33

Det.: Thompson, E.J., May 2012

Dup.: 359.0 Poaceae

HAN 0M

RI- 022169

Prep at BRI: Sheet

penseses JUBUAdOS

Fig. 11A. Holotype of Cryptachne trinerva (BRI [AQ1022169], sheet | of 2).

Datum: GDA34

11.6 km W of Charters Towers. eee

Austrobaileya 12(1): 26-58 (2022)

QUEENSLAND HERBARIUM (BRI)

Brisbane Australia

aa /02216q

‘ .

; fe :

Thompson, Cryptachne

ors E D HERBARIUM (BRI

= Queensland Herbarium (BRI) mes arsonist rv, ih (BRI)

| \

Australia: Queensland North Kennedy

N Dimorphochloa sp. (Charters Towers E.J.Thompson+ AQ / O a o / @ 9 '

} CHA554) f

to Coll. Thompson, E.J.; Simon. B.K. 1 May 2012 er ff of

Coll. no.: CHA792 | . J

> Lat,: 20" 7' 16.100"S Long,: 146° 9° 20.100"E }

| Datum: GDA94

oO 11.6 km W of Charters Towers.

Woodland of Acacia shirleyi on lateritic jump-up.

Dominant decumbent perennial grass ascending to 50 cm

tall,

Wot 33 \

. f

: :

-_——!1

eee

pesieseas jyHuAdoOS

Det.: Thompson, E.J., May 2012

Dup. 359.0 Poaceae

470

@ numa

Prep at BRI: Sheet

Fig. 11B. Holotype of Cryptachne trinerva (BRI [AQ1022169], sheet 2 of 2).

42 Austrobaileya 12(1): 26-58 (2022)

Fig. 12A. Terminal chasmogamous inflorescence and spikelets of Cryptachne trinerva. A—-F. A. adaxial view of

spikelet showing small lower glume and lower lemma. B. dorsal view of upper glume. C. dorsal view of upper lemma.

D. ventral view of upper lemma. E. ventral view of upper palea. F. terminal raceme. All from Thompson CHAS555 &

Turpin (BRI). Del. E.J. Thompson.

Thompson, Cryptachne 43

Fig. 12B. Axillary cleistogamous inflorescence and spikelets of Cryptachne trinerva. G. pedicellate spikelet in

situ. H. paired spikelets, sessile and pedicellate with leaf sheath removed. I. sessile spikelet in situ showing bowed

basal portion of culm internode. J—O. pedicellate spikelet. J. lateral view. K. adaxial view of spikelet showing

lower glume and lower lemma. L. dorsal view of upper glume. M. dorsal view of lower lemma. N. dorsal view of

upper lemma. O. ventral view of upper palea. All from Thompson CHA555 & Turpin (BRI). Del. E.J. Thompson.

44 Austrobaileya 12(1): 26-58 (2022)

a4 F QUEENSLAND HERBARIUM (BRI)

AA Queensland Herbarium (BRI) Brisbane Australia

Australia: Queensland Darling Downs

= | Dimorphochloa sp. (Miles E.J.Thompson EJT888) AQ /O2 / 9 0 6

6 Coll.: Thompson, E.J. 26 Mar 2013

Coll. no.: EJT 906

Lat.: 26° 36' 40.600"S Long.: 150° 23’ 3.000"E

Datum: GDA94

23 km NE of Miles, 10 km NNE of Columboola

Open woodland of Eucalyptus fibrosa with understorey of

Callitris glaucophylla and ground layer of Cleistochloa

subjuncea and Calyptochloa gracillima on sandy soil on

sandstone.

Occasional perennial grass to 40 cm tall with short stolons.

eee || bill

paniesel 1u6iuAdoS

Det.: Thompson, E.J., Mar 2013

Dup.: 359.0 Poaceae

~NN

RI- 0219

| Prep at BRI: Sheet

(ey)

Fig. 13A. Holotype of Cryptachne columboola (BRI [AQ1021906], sheet I of 2).

Thompson, Cryptachne

fy a QUEENSLAND HERBARIUM (BRI)

Queensland Herbarium (BRI) Vi Brisbane Australia

¢ f

Australia: Queensland Darling Downs

! Dimorphochloa sp. (Miles E.J. Thompson EJT888) AQ / 0 ol / g O C

Coll.: Thompson, E.J. 26 Mar 2013

| Coll. no.: EJT 906

| Lat.: 26° 36' 40.600"S Long.: 150° 23' 3.000"E

Datum: GDA94

23 km NE of Miles, 10 km NNE of Columboola

Open woodland of Eucalyptus fibrosa with understorey of

Callitris glaucophylla and ground layer of Cleistochloa

subjuncea and Calyptochloa gracillima on sandy soil on

sandstone.

Occasional perennial grass to 40 cm tall with short stolons.

— ————————— SS ———__—

Det.; Thompson, E.J., Mar 2013

MAT

Prep at BRI: Sheet

Fig. 13B. Holotype of Cryptachne columboola (BRI [AQ1021906], sheet 2 of 2).

46 Austrobaileya 12(1): 26-58 (2022)

5mm

: : a

mm die

ae ly

\ Ay

Es .T

Fig. 14A. Terminal chasmogamous inflorescence of Cryptachne columboola. A. adaxial view of spikelet showing

small lower glume and lower lemma. B. dorsal view of upper glume. C. dorsal view of upper lemma. D. ventral view

of upper lemma. E. ventral view of upper palea. F. terminal raceme. All from Thompson EJT906 (BRI). Del. E.J.

Thompson.

Thompson, Cryptachne 47

Fig. 14B. Pedicellate spikelet of axillary chasmogamous inflorescence of Cryptachne columboola. G. pedicellate

spikelet, in situ. H. lateral view. I. dorsal view. J. dorsal view of upper glume. K. dorsal view of upper lemma L. ventral

view of upper palea. All from Thompson EJT906 (BRI). Del. E.J. Thompson.

48 Austrobaileya 12(1): 26-58 (2022)

Queensland Herbarium (BRI)

Australia: Queensland Leichhardt QUEENSLAND HERBARIUM (BRI)

Dimorphochloa sp. (Mt Cooper R.J.Cumming 18623) Brisbane Australia

Coll.: Thompson, E.J.; Simon, B.K. 12 May 2013 AQ [0 a a [/ /

Coll. no.: EJT931

Lat.: 23° 42' 15.400"S Long.: 149° 34’ 45.900"E

Datum: WGS84

Duaringa SF, edge of Capricorn Highway, c. 10 km west of

Duaringa.

Woodland of Acacia shirleyi with A. rhodoxylon on laterite,

along creek line with ground layer dominated by

Cleistochloa subjuncea,.

Occasional perennial, tussock-forming grass to 50 cm tall.

Wot 14

Det.; Thompson, E.J., May 2013

Dup-: 359.0 Poaceae

RI- 02

Prep at BRI: Sheet

)

”

Fig. 15A. Holotype of Cryptachne duaringa (BRI [AQI022111], sheet 1 of 2).

Thompson, Cryptachne 49

Queensland Herbarium (BRI)

QUEENSLAND HERBARIUM (BRI)

Australia: Queensland Leichharal Brisbane Australia

Dimorphochloa sp. (Mt Cooper R.J.Cumming 18623)

Coll.: Thompson, E.J.: Simon, B.K, 12 May 2013 AQ LlOa o / ( [

Coll. no.: EJT931

Lat.: 23° 42' 15.400"5 Long.: 149° 34' 45.900"E

Datum: WGS84

Duaringa SF, edge of Capricorn Highway, c. 10 km west of (f

Duaringa. | f

Woodland of Acacia shirleyi with A. rhodoxylon on Jaterite, |

along creek line with ground layer dominated by

Cleistochioa subjuncea.

Occasional perennial, tussock-forming grass to 50 cm tall.

Wpt 14

Det.; Thompson, E.J., May 2013

Dup.:

RI-AQ1

359.0 Poaceae

fe)

re |

Prep at BRI: Sheet

Fig. 15B. Holotype of Cryptachne duaringa (BRI [AQI022111], sheet 2 of 2).

50 Austrobaileya 12(1): 26-58 (2022)

5 mm

Fig. 16A. Terminal inflorescence spikelets of Cryptachne duaringa. A. adaxial view of spikelet showing small lower

glume and lower lemma. B. dorsal view of upper glume. C. dorsal view of upper lemma. D. ventral view of upper

lemma. E. ventral view of upper palea. F. terminal raceme. All from Thompson EJT931 & Simon (BRI). Del. E.J.

Thompson.

Thompson, Cryptachne 5]

Smm

Fig. 16B. Cryptachne duaringa. G. sterile internode with leaf and sheath. J—P. Axillary inflorescence spikelets. J.

lower lemma. K. dorsal view of upper glume. L. cross-section of upper glume. M. dorsal view of upper lemma N.

ventral view of upper palea. O. ventral view of caryopsis showing scutellum. P. dorsal view of caryopsis showing

hilum. All from Thompson EJT931 & Simon (BRI). Del. E.J. Thompson.

Distribution and habitat: Cryptachne

columboola 1s known from a single location

near Miles (Map 1). Plants occur in a

woodland of Eucalyptus fibrosa F.Muell., on

undulating terrain with sandy soil derived

from duricrust. Regional Ecosystems

represented include 11.7.7. (QRE 2020).

Phenology: Flowering December to June.

Etymology: The specific epithet 1s derived

from the name of the location where the

species was first discovered, near the small

Queensland country town Columboola, in

Barunggam country. The town is named

after Columboola Creek, an Aboriginal

word, meaning plenty of white cockatoos

(http://en.wikipedia.org.wiki/Coolumboola,

accessed 13 September 2021). The epithet is

to be treated as a noun 1n apposition.

Conservation status: Cryptachne columboola

is known from a single location 1n a landscape

threatened by clearing for mining and pasture

development. It 1s suggested that this species

should be considered Critically Endangered

based on Criterion B2a (IUCN 2019) with a

formal conservation status nomination to be

made elsewhere.

3. Cryptachne duaringa E.J.Thomps., sp.

nov.

Culms to 60 cm tall. Apical spikelets in

terminal inflorescences mostly c. <5 mm long;

axillary spikelets mostly pedicellate only, < 7

mim long. Spikelets in terminal inflorescences

with ascending hairs to 2 mm long. Typus:

Queensland. LEICHHARDT DISTRICT:

Duaringa State Forest, edge of Capricorn

Highway, c. 10 km west of Duaringa, 12 May

2013, E.J. Thompson EJT931 & B.K. Simon

(holo: BRI [AQ1022111, comprising 2 sheets]).

Dimorphochloa sp. (Mt Cooper R.J.Cumming

18623): Simon & Thompson (2013); Thompson

(2021la, 2021c, 2022a).

Perennial grass; culms 40—60 cm tall, clumps

spreading to 100 cm across, copiously

branched. Culm internodes with hairs to 1.3

mm long. Leaf sheaths with hairs to 5 mm

long. Ligule c. 0.3 mm long. Contraligule c.

0.2 mm long. Leaf blades at mid-culm 20—55

mm long, 1.5—4 mm wide, hairs to 1 mm

Austrobaileya 12(1): 26-58 (2022)

long. Terminal inflorescence: axes 20-30

mm long, 5—9 flowered. Spikelets 3.2—5.2

mm long (without awn), 1.2—-1.4 mm wide;

lateral pedicels 0.2—0.6 mm long, terminal

pedicels 3.5—5.4 mm long. Lower glume to

0.2 mm long. Upper glume 3.2—4.7 mm long.

Lower lemma 3.2—5.2 mm long; pilose, hairs

ascending, to 1 mm long. Upper lemma 3.2-

5.2 mm long, awn 0.4—1 mm long. Lodicule

0.3-0.6 mm long. Anthers 2—3 mm long.

Caryopsis 2.9-3.3 mm long, 1.1—-1.3 mm

wide. Axillary inflorescence: Spikelets

mostly present as pedicellate only; up to 5 per

culm; 5.4—6.9 mm long, 0.9—1.1 mm wide.

Upper glume 4.9—6.2 mm long. Lower lemma

5.4—6.9 mm long. Upper lemma 5.4—6.9 mm

long, awn 0.4—1 mm long; palea 4.9—5.1 mm

long. Solitary subsessile to sessile spikelet

rarely present, if present then 4.8—7.2 mm

long. Paired spikelets: sessile 4.1-5.6 mm

long; pedicellate 6.1-—6.2 mm long, pedicels

1-10 mm long. Anthers 0.5—0.7 mm long.

Caryopsis 3.6—4.3 mm long, 1.2—-1.8 mm

wide. Figs. 15 & 16.

Additional specimens examined: Queensland. NORTH

KENNEDY District: Mt Cooper Station — 80 km SE of

Charters Towers, Mar 1999, Cumming 18623 (BRI);

20 km SW of Charters Towers, May 2012, Thompson

CHA790 & Simon (BRI); 81 km SE of Charters Towers

on Mt Cooper Station, May 2012, Thompson CHA8&22

& Simon (BRI); 16 km SW of Charters Towers, May

2013, Thompson CHAS31 & Simon (BRI). Cultivated.

Ashgrove, Apr 2017, Thompson MORS821 (BRI); ibid,

Apr 2017, Thompson MORSIY (BRI); ibid, Apr 2017,

Thompson MORS823 (BRI).

Distribution and habitat: Cryptachne

duaringa 1s known from a few localities

near Charters Towers and one location near

Duaringa (Map 1). Plants occur occasionally

in a woodland of Acacia shirleyi or A.

catenulata on slopes of jump-ups or “stony

rises’. Regional Ecosystems represented

include 10.7.3a and b, and 11.7.2 (QRE 2020).

Phenology: Flowering December to June.

Etymology: The specific epithet 1s derived

from the name of the location where the species

was first discovered near the Queensland

country town Duaringa, in Gangulu country.

The town’s name is popularly speculated to be

derived from the aboriginal words D’warra

D’nanjie (Duarininga) meaning a meeting

Thompson, Cryptachne

place in the swamp oaks (https://en.wikipedia.

org./wiki/Duaringa, accessed 13 September

2021). The epithet is to be treated as a noun

in apposition.

Conservation status: Cryptachne duaringa

is known from several small and fragmented

subpopulations at only a few locations. These

sites represent a very restricted landscape

type that is threatened by clearing for pasture

development. It is suggested that this species

should be considered Endangered based on

Criterion B2a (IUCN 2019) with a formal

conservation status nomination to be made

elsewhere. It has been recorded in a road

reserve adjacent to Duaringa State Forest, but

similar habitat occurs in the state forest, so it

is likely to be present there.

Acknowledgements

Iam indebted to B.K. Simon for his mentoring

in grass identification both in the field and

in the herbarium over many years. Dr G.P.

Guymer provided invaluable support at BRI.

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—— (1979). A procedure for standardizing comparative

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(GUEDES,

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Austrobaileya 12(1): 26-58 (2022)

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from BOM (2020) following Thompson (2021c).

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A revision of Neptunia Lour. (Leguminosae: subfamily

Caesalpinioideae, Mimosoid clade) in Australia and Malesia

A.R. Bean

Summary

Bean, A.R. (2022). A revision of Neptunia Lour. (Leguminosae: subfamily Caesalpinioideae,

Mimosoid clade) in Australia and Malesia. Austrobaileya 12: 59-106. Neptunia comprises 18

species for Australia and Malesia, including ten species newly named: N. heliophila A.R.Bean, N.

hispida A.R.Bean, N. insignis A.R.Bean, N. longipila A.R.Bean, N. paucijuga A.R.Bean, N. proxima

A.R.Bean, N. scutata A.R.Bean, N. tactilis A.R.Bean, N. valida A.R.Bean and N. xanthonema

A.R.Bean. One taxon below species rank (N. amplexicaulis f. richmondii Windler) is also accepted.

The distributions of all taxa are mapped, and all species are illustrated. Lectotypifications are

provided for N. amplexicaulis Domin, N. dimorphantha Domin, N. gracilis Benth., N. javanica Miq.

and N. monosperma F.Muell. ex Benth. Two identification keys are provided, one suited to flowering

material and the other for fruiting material. A conservation status is suggested for N. insignis.

Key Words: Leguminosae; Caesalpinioideae; Mimosoid clade; Neptunia; Neptunia heliophila;

Neptunia hispida; Neptunia insignis; Neptunia longipila; Neptunia paucijuga; Neptunia proxima;

Neptunia scutata; Neptunia tactilis; Neptunia valida; Neptunia xanthonema; flora of Australia; flora

of Malesia; flora of New Guinea; flora of Queensland; flora of Western Australia; flora of Northern

Territory; flora of New South Wales; flora of South Australia; new species; morphology; thigmonasty;

distribution maps; identification keys; conservation status

A.R. Bean, Queensland Herbarium and Biodiversity Science, Department of Environment and

Science, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia.

Email: tony.bean@des.qld.gov.au

Introduction

Neptunia Lour. was established by Loureiro

(1790) with the single species N. oleracea

Lour. The name is derived from Neptune,

the Roman god of freshwater and the sea,

and alludes to the aquatic habitat of the first

named species. Previously, Neptunia species

were described under the genus Mimosa L..,

e.g. M. plena L., M. natans L.f., M. prostrata

Lam. Willdenow (1806) named the genus

Desmanthus Willd. and included in it some

species now known as Neptunia, with that

usage followed by De Candolle (1825). It

was through the seminal work of Bentham

(1841) that the modern circumscription of

the genus Neptunia was established; therein

he distinguished Neptunia from the related

genus Desmanthus, but still likened it strongly

to his Piptadenia Benth. The morphological

definition of Neptunia was further elaborated

by Bentham (1864) in his treatment of the

three Australian species that he recognised.

Neptunia can be recognised by its

herbaceous habit, unarmed stems, bipinnate

stipulate leaves, and the globose to ellipsoidal

inflorescences solitary in the leaf axils,

usually comprising a mixture of bisexual

and neuter flowers that have yellow, petaloid

staminodia. It comprises about 22 species and

is distributed in the tropics and subtropics of

North America, South America, Australia,

mainland Asia, Malesia and Africa (Windler

1966). This taxonomic revision is restricted

to the species that occur in Australia and

Malesia.

The biogeographic region of Malesia

comprises Indonesia, Malaysia, Brunei, the

Philippines, Christmas Island (an Australian

external territory), East Timor and the whole

of New Guinea (van Steenis 1948). The two

species of Neptunia named from Malesia,

N. javanica Miq. and N. depauperata Merr.,

were described in 1855 and 1918 respectively.

Accepted for publication 29 August 2022, published online 16 November 2022

The first Australian Neptunia species to

be formally named was N. gracilis Benth.

(Bentham 1841). In the Flora Australiensis,

Bentham (1864) described N. gracilis var.

major Benth. (= N. major (Benth.) Windler)

and N. monosperma F.Muell. ex Benth., using

a manuscript name provided by F. Mueller for

the latter. Domin (1926) named two further

species, N. amplexicaulis Domin and N.

dimorphantha Domin, from specimens he

collected in 1910 while visiting Queensland.

In his revision of the genus, Windler (1966)

raised N. gracilis var. major to species

rank and named two new taxa at the rank

of forma. No Australian or Malesian taxa

have been described since that time. This

revision recognises 18 species for Australia

and Malesia, including ten that are newly

described.

Morphology

a) Habit. In most species, the stems are

quite prostrate, radiating from a central

rootstock. In others (e.g. Neptunia major, N.

monosperma, N. valida A.R.Bean) the stems

are upright or ascending. In some populations

of NV. heliophila A.R.Bean, the upright shrubby

plants occur in colonies, strongly suggesting

the presence of a network of rhizomes. Most

species are perennial, although the leaflets

abscise and the stems die back each year, a

few months after the wet season. Northern

Territory botanists have coined the term

“annual herb with perennial rootstock” for

local species of Neptunia, except for N. major

which they regard as an annual (Dunlop ef al.

1995).

b) Stipules. Stipules are present in all species,

but they vary in size (from 4—20 mm long)

and persistence. In Neptunia major and N.

monosperma, the stipules are narrow and

deciduous (often visible only on the newly

emerging vegetative growth). In all other

species, the stipules are broad and persistent

for many nodes.

c) Stipels. In Neptunia, the stipel 1s contiguous

with the raised margin of the petiole or rachis.

It 1s persistent, 1s sometimes deeply 2- to

4-lobed, and may have a gland-tipped apex.

Stipels seem to be present in all species, but

Austrobaileya 12: 59-106 (2022)

often they are extremely short and barely

noticeable. These structures were referred to

as “pseudostipels’ by Cowan (1998).

d) Leaves and nectaries. The leaves of

Neptunia are in all cases alternate and

bipinnate, with the leaflets oblong and

somewhat chartaceous. The number of pairs

of pinnae and the number of leaflets per pinna

vary considerably, but are to some degree

taxonomically diagnostic, particularly if

juvenile growth near the base of the plant

is ignored. The indumentum density and

pattern on the leaves and stems 1s somewhat

diagnostic but must be used with caution.

Neptunia major, N. javanica, N. oleracea,

N. plena (L.) Benth. and N. monosperma are

virtually glabrous on vegetative parts, while

most other species have at least sparse hairs on

the leaves or rachises. Neptunia amplexicaulis

has a glabrous and a pubescent form, and N.

gracilis has variable indumentum.

The petiole and sometimes the rachis,

can bear one or more circular or elliptical

nectaries on the upper (adaxial) side; the

presence, number, size and position of these

nectaries is usually diagnostic. However,

there 1s some variation within a species, e.g.

Neptunia scutata A.R.Bean. On a single plant

or specimen the nectaries can be present on

some leaves, but not on others, or may vary in

number. Therefore, more than one leaf should

be examined to properly assess the character.

With herbarium specimens, the difficulty 1s

that if a branch is mounted with the adaxial

side of all leaves down, this character will be

hidden from view.

e) Peduncles and bracts. The length of the

peduncle is diagnostic. From examination

of specimens, it is clear that 1n some species

the peduncle reaches its full length at, or

just before anthesis, as the peduncle does

not elongate between the flowering and

fruiting stages. In other species, the peduncle

elongates considerably between flowering and

fruiting. Neptunia monosperma, N. scutata

and N. xanthonema A.R.Bean have short

fruiting peduncles 10-20 mm long, while

in NV. gracilis, N. plena and N. heliophila,

the fruiting peduncle may reach 105-140

mm in length. The peduncle indumentum

Bean, Neptunia in Australia and Malesia

is also diagnostic, 1.e. the type (glandular or

eglandular), length and direction (antrorse or

patent) of hairs.

The peduncular bracts are highly

characteristic with regard to their number (0,

1, 2, or more), size and position (lower one-

third of peduncle, middle one-third, or distal

one-third). The bracts are usually prominently

veined, but the venation does not seem to vary

ereatly between species.

{) Inflorescence. One of the defining features

of Australian and Malesian Neptunia are

the dimorphic inflorescences, where the

spike has bisexual flowers at the apex, and

neuter flowers at the base. Some American

species (e.g. NV. pubescens) have trimorphic

inflorescences, with bisexual flowers at the

apex, male flowers below the middle, and

neuter or sterile flowers at the base (Tucker

1988).

The neuter flowers of Australian and

Malesian species comprise a calyx, corolla,

and 5-10, long flattened staminodes. Two

Australian species (Neptunia proxima

A.R.Bean and WN. tactilis A.R.Bean) have

wholly bisexual inflorescences. The

occurrence of dimorphic inflorescences

is often difficult to detect from herbartum

specimens. While some specimens appear to

have entirely bisexual inflorescences, this can

be because the neuter flowers have abscised.

Before anthesis, dimorphism is_ equally

difficult to detect because the neuter flower

buds are almost identical in appearance to

the bisexual flower buds. The flowers (both

bisexual and neuter) are nearly always yellow,

and the number of stamens is in most species

5, but in a few species (NV. oleracea, N. plena,

N. tactilis, N. hispida A.R.Bean, N. javanica),

there are between 5 and 10 stamens.

2) Pods. The pods for any given species of

Neptunia are relatively consistent in size,

shape, number of seeds, and indumentum.

Five species (NV. longipila, N. monosperma,

N. scutata, N. valida and N. xanthonema)

are consistently one-seeded and _ have

circular or broadly elliptical pods. Neptunia

dimorphantha and N. paucijuga have pods

with 1—3 seeds; pods of NV. javanica, N. plena

and N. tactilis typically have 7—14 seeds, and

the remainder (NV. amplexicaulis, N. gracilis,

N. heliophila, N. hispida, N. insignis, N.

major, N. proxima) usually have 3-7 seeds

per pod.

Pod indumentum is a useful delimiting

character: Neptunia dimorphantha has

curved antrorse hairs mainly on the pod

margins; N. longipila, N. scutata, N. valida

and N. xanthonema have patent glandular

hairs 0.2—0.5 mm long throughout; N. gracilis

and NV. monosperma pods are either glabrous

or have dense patent eglandular hairs c.

0.1 mm long; NV. insignis pods vary in the

density and distribution of hairs; N. proxima

has long patent hairs; N. amplexicaulis f.

richmondii has short pubescent eglandular

hairs; the remaining taxa (N. amplexicaulis

f. amplexicaulis, N. heliophila, N. hispida, N.

Javanica, N. major, N. paucijuga, N. plena, N.

tactilis) have pods that are strictly glabrous.

Neptunia pods are routinely described

as ‘dehiscent’ (Windler 1966; Murray 1991;

Nielsen 1992; Wheeler 1992), but for many

species that is not the case. For all species with

single-seeded pods, the pods are indehiscent

— they fall to the ground with the seed inside

and the seed is released only when the pod

decays. The pods of N. amplexicaulis, N.

dimorphantha, N. major and N. paucijuga are

also indehiscent. In N. hispida, N. javanica,

N. plena, and N. tactilis the pods open along

one suture only. In the remaining species,

dehiscence 1s along both sutures.

Chemical properties

Selenium hyperaccumulation

Selenium (Se) is a naturally occurring

metalloid element essential to human and

animal health in trace amounts, but harmful

in excess (Fordyce 2013). Some plant species

are known to accumulate high concentrations

of selenium. The Queensland species

Neptunia amplexicaulis is one of the strongest

selenium hyperaccumulators known (Harvey

et al. 2020) and it is very tolerant of soils

containing high levels of selenium. Irish ef

al. (2021) found that ‘N. gracilis’ (herein

recognised as N. heliophila), growing in the

same region as N. amplexicaulis, is not a

selenium accumulator and does not tolerate

high levels of this element in the soil. The

other Australian and Malesian species have

not yet been investigated in this regard.

Leaf sensitivity (thigmonasty)

Thigmonasty, leaf movement triggered

by touch, is found in numerous mimosoid

legumes. In some species, e.g. Mimosa pudica

L., the leaflets fold onto themselves within

a few seconds when touched. Some (and

possibly all) Australian species of Neptunia

have sensitive leaves, but the movement is

rather slow and the leaflets often do not fold

completely. Further examination 1s required

of live material of the species recognised in

this revision.

The response results from a loss of

turgor from extensor cells of the pulvini

(Braam 2005). It is unclear what benefit

species of Neptunia and other mimosoids

derive from the leaflet-folding response.

Eisner (1981) postulated that, for Schrankia

microphylla (Dryand.) J.F.Macbr., (now

Mimosa microphylla Dryand.) it is a defence

mechanism against grazing animals, as the

folding of the leaves exposes more of the

prickles on the rachises and stems. This was

disputed by Minorsky (2019), who considered

that this response would not deter most

herbivores.

Infrageneric classification and phylogeny

In his revision, Windler (1966) divided

Neptunia into two. sections, WN. ‘sect.

Neptunia’ and N. ‘sect. Pentanthera’. In his

key, these were distinguished (apart from

geography) solely on stamen number (10 for

sect. Neptunia and 5 for sect. Pentanthera).

As there is no Latin diagnosis for sect.

Pentanthera, tt is an invalid name, and hence

the autonym was not validly published either.

In any case, the benefit of having two sections

based on stamen number is debateable as some

apparently related species (e.g. N. gracilis

and N. javanica) would then be in separate

sections. Neptunia tactilis has 5—8 stamens

and would not fit easily into either section.

Austrobaileya 12: 59-106 (2022)

Previous authors have allied Neptunia

with Desmanthus (Bentham 1841, 1864, 1875;

Windler 1966) and Dichrostachys (Lewis

& Ehas 1981; Koenen 2020). Hughes ef al.

(2003) studied the generic relationships and

phylogeny of these and several other mimosoid

genera, establishing two monophyletic groups,

the informal ‘Dichrostachys group’ which

includes Dichrostachys, and the ‘Leucaena

eroup’ which includes Desmanthus. This

well-sampled study (including = seven

species of Neptunia) found that Neptunia

is monophyletic, but that it lies outside both

of these clades. The Australian species with

5 stamens formed a separate clade from

the American species with 10 stamens, but

problematic species (NV. hispida, N. javanica,

N. tactilis) were not sampled. Koenen ef al.

(2020) found that Neptunia belongs to a

more broadly circumscribed ‘Dichrostachys

clade’ along with 12 other genera, including

Desmanthus, Dichrostachys, Desmanthus,

Leucaena Benth. and Schleinitzia Warb. The

presence of heteromorphic inflorescences with

showy staminodes at the base is characteristic

of Neptunia and several other genera in this

clade, but the staminode character is perhaps

not homologous.

Hughes ef al. (2003) pointed out that the

yellow petaloid staminodes of Neptunia are

quite different from the filamentous, pink

or white staminodes of Desmanthus and

Dichrostachys.

Ecology

Koenen eft al. (2020) stated that Neptunia 1s

unlike any other mimosoid genus because ofits

“semi-aquatic lifestyle”, but the only aquatic

or semi-aquatic species are N. oleracea and N.

plena; all other species are strictly terrestrial.

Neptunia plena is naturalised in Australia and

N. oleracea is either indigenous in Malesia

or a long-established alien. The seemingly

uncommon WN. javanica inhabits terrestrial

open-forest habitats in Malesia, as does N.

gracilis.

Indigenous species of Neptunia in

Australia are strictly terrestrial, and they are

distributed in the northern half of Western

Australia, throughout the Northern Territory

Bean, Neptunia in Australia and Malesia

and Queensland, northern New South Wales,

and in northern South Australia. A majority

of species show a strong preference for heavy

clay vertosol soils, often growing in grassland

or open eucalypt woodland. Other species

grow on flat to somewhat hilly terrain, in

tropical woodland with loamy or sandy-loam

soils, often associated with laterite.

It 1s not unusual to find two, or even

three species of Neptunia growing at the

same location in Australia (pers. obs.). This

explains ‘mixed’ herbarium specimens, Le.

more than one taxon mounted on a herbarium

sheet. Despite this sympatry, there is no clear

evidence of hybridisation, from either field

or herbarium study. However, hybridisation

has played a part in Neptunia speciation, as

a recently described species from Brazil, WN.

windleriana J.Santos-Silva & V.F.Mansano is

a polyploid (Santos-Silva eft al. 2020).

Only one of the species of Neptunia

treated in this paper (NV. insignis) 1s considered

threatened under the criteria of the IUCN

(2012) with a suggested conservation status

provided. Most species are very widespread

and/or are represented in _ conservation

reserves. Neptunia javanica 1s represented

by relatively few collections at L and other

European herbaria; but was described by

Nielsen (1992) as “locally common”. The

Bogor herbarium (BO), the most likely

repository for specimens of N. javanica, could

not be consulted for this study.

Materials and methods

This revision is based on an examination

of specimens at BRI, and those received on

loan from AD, CANB, DNA, L, MEL, NT

and PERTH. Specimens at NSW were not

available for loan, but high resolution images

of all their specimens of Neptunia were

made available. Specimen images (mostly

types) from K, P, PR, NY, US and W have

also been examined and are indicated as

i.d.v. (imago digitalis visa) in the text. Most

species have been examined in the field, with

observations including habit, stipule and bract

persistence, extent of floral dimorphism and

fruit morphology. The species are treated in

alphabetical order.

Measurements of floral parts are based on

material reconstituted in hot water or preserved

in 70% alcohol; all other measurements were

taken from dried material. The range of

measurements 1s inclusive, 1.e. 7.5—8.0 1s given

as 7.5—8. Where there is apparent overlap in

some measurements given in the key, it 1s

important to examine a range of examples on

the specimen being identified (e.g. number

of pinnae) and use the average. The number

of pinnae and the number of leaflets per

pinna were assessed and measured adjacent

only to the flowering or fruiting peduncles

on the distal parts of the plant, because the

leaves from the basal part of the plant are not

fully developed, and consistently have fewer

pinnae and fewer leaflets per pinna. Only the

largest available leaflets were measured for

length and width on any given specimen. In

this paper, the portion of the leaf stalk below

the lowest pair of pinnae 1s termed the petiole;

the portion above this point is termed the

rachis; hence in leaves with a single pair of

pinnae, the rachis is absent.

Abbreviations used in the specimens cited

sections include ‘NP’ for National Park and

‘HS’ for Homestead. Herbarium acronyms

follow Thiers (updated continuously).

Distribution maps have been compiled with

DIVA-GIS Version 7.5.0 using localities or

geocodes given on the labels of specimens

from the herbaria listed above. Latitudes and

longitudes have been determined for some

locations in Indonesia (Neptunia javanica)

and are included in the specimen citation to

enable geocoding curation 1n herbaria.

Taxonomy

Neptunia Lour., Fl. Cochinch. 653 (1790).

Desmanthus sect. Neptunia (Lour.) DC.,

Prodr. 2: 444 (1825). Type: NV. oleracea Lour.

Hemidesmas Rat., Sylva Tellur. 119 (1838).

Type: not designated

Annual or perennial herbs or shrubs, prostrate

to upright, reaching 3 min height; most species

terrestrial, but some semi-aquatic; foliage

unarmed; stems terete, small branchlets

angular, glabrous or pubescent with simple

trichomes. Stipules lanceolate to broadly

ovate, free, in pairs; in most species persistent

for many nodes, but in some species caducous

and visible only near growing tip. Leaves

alternate, bipinnate, petiolate, with 1—6 pairs

of pinnae, frequently touch-sensitive, with the

leaflets folding in towards each other. Pinnae

lacking a terminal leaflet; rachis and petiole

canaliculate on upper side where | or more

extrafloral nectaries are often present; petiole

and rachis ridges extending at the nodes into

subulate, branched or unbranched persistent

stipels; rachis extending beyond last pair of

pinnae; rachillas without nectaries, extending

beyond distal pair of leaflets. Leaflets

opposite, obliquely attached to rachilla,

oblong, somewhat chartaceous, venation

usually visible on lower surface. Inflorescence

axillary, with a single peduncle per node,

bearing a congested terminal spike. Peduncle

usually with | or 2(—4) disjunct bracts (similar

in appearance to the stipules), but in some

species bracts are absent. Flowers 6—60 per

inflorescence, sessile, each subtended by a

bracteole. Inflorescences 1n some species

dimorphic, with the upper flowers bisexual,

Austrobaileya 12: 59-106 (2022)

and the lower flowers neuter. Bisexual flowers:

calyx fused, campanulate or obconical,

5-lobed, green; corolla with 5 free yellow

elliptical petals; stamens often 5 but up to 10

in some species, filaments glabrous, anthers

bilocular, with a small terminal gland (in

some species), ovary superior. Neuter flowers:

calyx campanulate; corolla with 5 free yellow

elliptical petals; stamens absent; staminodes

long-petaloid, yellow; gynoecium absent.

Pods orbicular, broadly elliptical, oblong or

falcate, flat, stipitate, dehiscent along one or

both margins, or indehiscent, 1—20-seeded,

brown at maturity; seeds flattened, orbicular

to broadly elliptical in outline, transversely

or obliquely arranged, pleurogram present.

Sensitive weed, Water Mimosa

Tropical and subtropical parts of North

America, South America, Asia, Africa,

Malesia and Australia. c. 22 species

worldwide; 18 species in Australia and

Malesia, 16 indigenous and 2 naturalised

(indicated * 1n text).

Key to the Australian and Malesian species of Neptunia (flowering material)

1 Plants aquatic, with inflated floating stems; staminal filaments 6—7.5 mm

[GTO ecw ee ty Ga nee ee fasts eed lie

1. Plants terrestrial, stems not inflated; staminal filaments 2-5.2mmlong .......... 3

2 Leaves with 2—4 pairs of pinnae, 12—24 pairs of leaflets per pinna; small

nectary usually present at distalend of petiole. ..............0... *N. plena

2. Leaves with 2 or 3 pairs of pinnae, 10—17 pairs of leaflets per pinna; petiole

nectary absent. < h 4a") ola o Soa RE:

2 ‘Petolencectary-absentiironiallicaves- 2c 68.0 a5 ox Ble Sm ei Roe bo Sealy Ele Bhar 4

3. Pétiole nectary present:on mostior all leaves «2 4 ace wa SS ce oe ee OY ee 14

4: Peduncles- with 2-2 bractsys. US Sos arhed tecesd, UL Sas gr pide pee be! Oo bege Arden. 4 5

4. Peduncles without bracts or with 1 bractonly ...................2.044 11

5 Peduncles mostly with 3 or 4 bracts, occasionally with2 ............. N. valida

Ss EALIMpCCUNCleS- With? WEACTS re wen wees ih beets Oy cn zd ath Dee ere dts Giese We Sia oe Hy Sy oes 6

6 Bracts lanceolate, both on the middle one-third of peduncle. ........... N. hispida

6. Bracts ovate to broadly ovate, with at least the lower one on proximal

OTS SANT GG Inf CCUG I hee 2h se wee Rab beatae es re Nee Sus ahecrlnee eal arene pore in Pv adc zagh atten ae. ee ee eS be 7

& Stipules 12-2 OmNy LON". 5 eke he be Mile ee ated dear bh sta die 5 N. amplexicaulis

Ts SOLUpPUles 4— (0: MTOM AL Breas SL ee 9, a: WE cage, GU Sp oe ho ee at ee BER a a Sle 8

8. Peduticles'/—35 mm long-atanthesis....4 vs +s BS RE dae we GREE eee 9

8. Peduncles 33-110 mm long atanthesis...........0.. 0.2... 002002 eee 10

Bean, Neptunia in Australia and Malesia 65

9 Peduncles glabrous except at apex; largest leaflets 5.8-11 mm _ long;

PINS Paves 2 Ot SAY. he a soe ws: ee BE Ree Se Gs ks Se SER EG GEES N. xanthonema

9, Peduncles hairy throughout; largest leaflets 3—7 mm long; pinna pairs

Da Ol Me ees cttw ta eine Sete its Greed sas See PB ce bee Sand a N. scutata

10 Leaves with 4—6 pairs of pinnae; largest leaflets 4.6-7.8 mm long ...... N. heliophila

10. Leaves with 2-3 pairs of pinnae; largest leaflets 8-13 mm long. ......... N. insignis

11 Neuter flowers absent; flowering peduncles 14-36 mm long. ................ 12

11. Neuter flowers present; flowering peduncles (25—)40-110 mm long. ........... 13

12 Pinnae with 16—21 pairs of leaflets; ovary hairy; stamens always5....... N,. proxima

12. Pinnae with 20—28 pairs of leaflets; ovary glabrous; stamens 5-8. ........ N. tactilis

13 Leaves predominantly with 2 or 3(rarely 4) pairs of pinnae; peduncles

OSU SAO Us SEN ee a Be eo oe ae Me as Senne een oe ie wor OR ys N. gracilis

13. Leaves with 4—6 pairs of pinnae; peduncles with sparse to dense antrorse

HANGS ais ote NAA oii ana oe Ba Se Se Gl eo eee ME ocey Sa a Ween N. heliophila

14 Peduncles without bracts; stamens 10 ..........0.....0....0200. N,. javanica

14, Peduncles-with-2 bracts; stamens 3... . 5 3 4b bee Gk we he he ee ee bees 15

15 Stipules deciduous, visible only on new growth; upright plants. ............. 16

15. Stipules persistent for many nodes; prostrate plants .................204. 17

16 Plant 1—2(-3) m high; petioles 10Q—18 mm long; peduncle 15—55 mm long

AU AMUNESISE. sk 8 YF oben BE nuda ow ee EE YE ete 8 BG N. major

16. Plant to 0.5 m high; petioles 5-11 mm long; peduncle 4-12 mm long

AIRATIENCSIS:.3 4 BAe re eye Ge ren oe be oh Be ca nt Gy eye ate Me ee ee tS N. monosperma

17 Peduncle glabrous except at apex; leaves with (1 or)2 or 3 pairs of pinnae ........ 18

17. Peduncles hairy throughout; leaves with 3—5 pairs of pinnae ............... 19

18 Ovary densely hairy; peduncles 7-32 mm long at anthesis; neuter flowers

PECSCIN Ve. a: US erage ay. Seg ee ty oie ae ot Ae oot Be BE ck ns wo AT cent ars ae al ea N. xanthonema

18. Ovary glabrous; peduncles 40-54 mm long at anthesis; neuter flowers

ASC Seeks ae By aces, Ba ww SSSA OM Ge theta: oleh BPR cg ieee eel “AOR asks N. paucijuga

19 Hairs on peduncle 0.5—0.9 mm long; inflorescences broadly-ellipsoidal ... .N. longipila

19. Hairs on peduncle 0.1—0.5 mm long; inflorescences globose. ...............

20 Staminodes 4—7.5 mm long; ovary hairy throughout. ............... N. scutata

20. Staminodes 8.5—14 mm long; ovary hairy along the margins only... . . N. dimorphantha

Key to the Australian and Malesian species of Neptunia (fruiting material,

thus excluding JN. oleracea)

Le <PcllPOOS SING IG=SECHCU om uc gg eee Flin g ORSi a WLS SSM. Gee Bile alum Wed eye pees 2

1. All pods, or a majority of pods, with more than one seed ................0.4 6

2 Peduncles glabrous throughout, or with hairs only in the upper one-fifth. ......... 3

De. PEAuncles: Waiey PNTOUeHOUTS, < a saon ve ae teas eld, OF ee ee EUROS Ge RM RST es “

3 Pods glabrous or with very dense eglandular hairs c. 0.1 mm long; leaflets

23-34 pairs per pinna; lower bract 1.5—2.3 mmlong............ N. monosperma

3. Pods with sparse glandular hairs 0.2—0.4 mm long; leaflets 11—24 pairs per

pinna; lower bract 5-14.5mm long..................... N. xanthonema

66 Austrobaileya 12: 59-106 (2022)

4 Peduncle hairs 0.5—0.9 mm long, eglandular................... N. longipila

4. Peduncle hairs-0.1—0;3 mnt long, elandular: -. 2... 2 26 ee eG ee ee 5

5 Erect shrub; some peduncles with 3 or 4 bracts; petiole nectary absent. .... . N. valida

5. Prostrate shrub; all peduncles with 2 bracts; petiole nectary usually present . . . N. scutata

6: Most-of allpetioles:satht AsISCtARY: & jo. Nos Ao oe we eee eh ae BOS Gh Hc Gs 7

G.. No: petibles WwitiadieClary se pd a Eo OE a a ea Doe ee sd 11

1 POUSssS=I5-=SGE0EG = woe 4 ala fat GH a Wom Bw AN He a OR fle OEM Re AAG 8

de (POUS: IS -6C6060)- <p ey etadale toe eee cette mm cle ten ree ~ 6 ee dnd 9

8 Semi-aquatic plant; leaflets 11-18 mmlong ..................0.. *N. plena

8. Terrestrial plant; leaflets 2.9-4.5mmlong.................... N,. javanica

9 Upright shrub; stipules deciduous; leaf pinnae with 23-39 pairs of leaflets... .. N. major

9. Prostrate shrubs; stipules persistent; leaf pinnae with 10—20 pairs of leaflets. ...... 10

10 Stems hairy; leaves with 3—5 pairs of pinnae; pods with short (0.1—0.2

mim) eglandular hairs along margin ................00.. N. dimorphantha

10. Stems glabrous; leaves with (1 or) 2 pairs of pinnae; pods glabrous. .... . N. paucijuga

I Darsestileaticts 6:21 lens: -.. en 8 Go a a ce ese ce ee Ew pe ae Se 12

Ph, Dareestleanets: 5-8-2 TR OMS me ares ek 4 Bea oe) BASALT ee we aS OR Ge CUE ww cnt 14

12 Plants semi-aquatic, with inflated floating stems; fruits 12-l5 seeded ...... *N. plena

12. Plants terrestrial, stems not inflated; fruits 2—7 seeded .................. 13

13 Fruiting peduncles 61-110 mm long; stems and peduncles with long

spreading hairs; lower peduncle bract 5—9 mm long. .............. N., insignis

13. Fruiting peduncles 36—52 mm long; stems and peduncles glabrous or with

short puberulous hairs; lower peduncle bract 11-19 mmlong...... N. amplexicaulis

14 Peduncle bracts lanceolate, both on the middle one-third of peduncle. ...... N. hispida

14. Peduncle bracts ovate to broadly ovate, on the proximal one-third of

PECUNCe ROMaOsent. = Shek, See et A Bek, See US eg OL ee See eB 15

15 Leaves with 4—6 pairs of pinnae; rachises with straight or curved antrorse

hairs:0;2 023m lone +. 4 whee Go Wa eke fae ee Ge boats & oS N. heliophila

15. Leaves with 1—3(—4) pairs of pinnae; rachises glabrous or with patent

PATS UCL TAINS gens GE Soca, na ae wee egdeim mo cen Se oa tee sgt ee BE oe, oj mg sew a ae ce A ep 16

16 Petioles 10-17 mm long; fruiting peduncles 50-105 mm long........... N. gracilis

16. Petioles 3—9 mm long; fruiting peduncles 20-59 mm long. ................ 17

17 Peduncle bracts consistently absent; pods 6—9-seeded, surface glabrous ..... N. tactilis

17. One bract present on some peduncles; pods 3—6-seeded, surface with

many patent eglandular hairs... .......0.. 0.0020. eee ene N. proxima

Bean, Neptunia in Australia and Malesia

1. Neptunia amplexicaulis Domin, Biblioth.

Bot. 22(89): 801, t. 24 (1926).

Type citation: “open grassland at Hughenden,

a robust, big form; at the Flinders River at

Hughenden, a small form.” Type: Australia.

Queensland. BURKE DISTRICT: in grassy

areas opposite Hughenden, February 1910,

K. Domin_ s.n. (lecto: PR 527689 [here

designated]; isolecto: PR 527688).

Shrub, perennial, terrestrial. Stems prostrate

or sprawling, to 0.35 m high, glabrous or

pubescent. Stipules persistent, broadly ovate,

12—20 mm long, 6—10 mm broad, coriaceous,

strongly nerved, glabrous or pubescent; apex

attenuate; base obliquely cordate. Leaves

with 2—4 pairs of pinnae; petioles 13—20 mm

long, glabrous or pubescent, nectary absent;

rachis 14-36 mm long, glabrous or pubescent,

nectary absent; rachis extension linear, 6—9.5

mm long; pinnae with 7—15 pairs of leaflets,

rachilla extension 2.6—5.3 mm long; stipels

usually linear, 0.3—0.6 mm long, but rarely (at

lowest node) expanded into a broadly elliptical

structure up to 2 mm long. Largest leaflets 10—

16 mm long, 3.5—6.7 mm wide, 2.2—4 times

longer than broad, glabrous or pubescent,

venation of 3 or 4 main veins, obvious on

lower surface. Inflorescences globose, all,

or most, with dimorphic flowers; peduncles

15—50 mm long at anthesis, 36—52 mm long at

fruiting stage, glabrous or pubescent; bracts 2,

at least the lower one positioned on proximal

one-third of peduncle, persistent, ovate to

broadly ovate, lower one 11-19 mm long,

glabrous or pubescent; flowers 25-50 per

spike; bracteole linear to lanceolate, 2.2—3.8

mm long, persistent. Bisexual flowers: calyx

campanulate, 1.3—2.4 mm long, glabrous or

pubescent; petals 2.8—3.5 mm long, glabrous

or sparsely hairy; stamens 5, filaments 3.6—4

mm long, anthers 0.6—0.9 mm long, terminal

gland not seen; style 2.4—-2.6 mm _ long,

glabrous. Neuter flowers: calyx 1.2—1.3 mm

long; petals 2.1-2.5 mm long, glabrous or

sparsely hairy; staminodes 5, 9-10 mm long,

Q.7—0.8 mm wide. Pods oblong, 2—6-seeded,

16—25 mm long, 9-12 mm broad, indehiscent,

stipe 1-2.5 mm long, apex obtuse; glabrous or

pubescent with hairs 0.15—0.3 mm long. Seeds

broadly elliptical to orbicular in outline, 5—5.3

mm long, 4.2—4.8 mm wide, brown to black.

Selenium weed.

Affinities: Neptunia amplexicaulis 1s

morphologically similar to WN. insignis.

Both have two or three pairs of pinnae, few

leaflets per pinna, no nectaries, and large

leaflets. Neptunia insignis differs by the long

spreading hairs on the stems and peduncles

(glabrous or with short puberulous hairs for

N. amplexicaulis); the stipules 4—7.5 mm

long (12-20 mm long for NV. amplexicaulis);

the bracteoles and calyx with a few hairs

(glabrous for N. amplexicaulis); the lower

bracts 5—9 mm long (11-19 mm long for N.

amplexicaulis); and the fruiting peduncles

61-110 mm long (36-52 mm long for N.

amplexicaulis).

Typification: Domin evidently made two

gatherings of Neptunia amplexicaulis in the

Hughenden area; a robust form from the

grasslands, and a small form from the Flinders

River (Domin 1926). A specimen from the

“orassland” gathering is here designated as

lectotype.

la. Neptunia amplexicaulis Domin f.

amplexicaulis, Windler, Aust. J. Bot. 14: 408

(1966).

Illustrations: Windler (1966: 406); Cowan

(1998: 22).

Stems, leaves, peduncles, ovaries and pods

glabrous. Fig. 1.

Additional specimens examined: Australia.

Queensland. BURKE DIsTRICcT: Kilterry, N of Richmond,

Mar 2009, Fensham 5856 (BRI); W of Swanson Street,

Hughenden, Jun 2021, Bean 34162 (BRI, CANB, MO,

NY); ‘Silver Hills’, Richmond, Mar 1960, McCray

s.n. (BRI [AQ235509]); near Ranmoor, 35 km N of

Richmond, May 1974, Byrnes 3064 (BRI); Gravel pit 11

km NW of Richmond, May 1974, Byrnes 3043 (BRI);

Flinders Highway, 19 km W of Hughenden, Jun 1998,

Bean 13356 (BRI); “Dunluce”, Hughenden, May 1963,

Johnston s.n. (BRI [AQ235495]); 60 miles [97 km] N W of

Maxwelton, Mar 1964, Entwistle s.n. (BRI [AQ235490]);

‘Sutherland’, 30 miles [48 km] N of Maxwelton, s.dat.,

Murray & Derrington s.n. (BRI [AQ235488]); ‘Yan

Yean’, c. 18 miles [29 km] NW of Richmond, Apr 1954,

Everist 5364 (BRI); ‘Lydia Downs’, c. 45 miles [72

km] NW of Maxwelton, Jan 1966, Pedley 1967 (BRI);

Telemon, 16 miles [26 km] NW of Hughenden, May

1958, Skerman s.n. (BRI [AQ235489]).

Austrobaileya 12: 59-106 (2022)

Fig. 1. Flowering branchlet of Neptunia amplexicaulis f. amplexicaulis (Bean 34162, BRI). Photo: A.R. Bean.

Distribution and habitat: Neptunia

amplexicaulis f. amplexicaulis 1s endemic to

northern Queensland, Australia, where it 1s

restricted to an area between Hughenden and

Cloncurry (Map 1). It inhabits dark cracking-

clay soils that often have a high concentration

of compounds with selenium content.

Phenology: Flowers are recorded from

December to June; fruits are recorded from

December to July.

Ib. Neptunia amplexicaulis f. richmondii

Windler, Aust. J. Bot. 14: 408 (1966).

Type: Australia. Queensland. BURKE

District: ‘Silver Hills’, Richmond, 23

March 1960, W. McCray s.n. (holo: BRI

[AQ0022907]).

Stems, leaves, peduncles, ovary and pods with

dense pubescent eglandular hairs 0.2—0.5 mm

long. Figs. 2D, 2E.

Additional specimens = examined: Australia.

Queensland. BURKE District: Flinders River, 20° 41’S

143° 08’E, Jul 1891, Plant 380 (BRI); Silver Hills Station,

Richmond, May 1974, Byrnes 30/11 (BRI); Gravel pit 11

km NW of Richmond, May 1974, Byrnes 3044 (BRI);

‘Silver Hills’, Richmond, Apr 1970, Murphy s.n. (BRI

[AQ235504]); c. 30 miles [48 km] WNW of Hughenden,

3 miles [5 km] SE of Telemon HS, Sep 1967, McCray s.n.

(BRI [AQ235511]); 5 miles [8 km] NW of Richmond, * Yan

Yean’ HS, Sep 1967, McCray s.n. (BRI [AQ235519]); c.

20 km NNW of Richmond on ‘Silver Hills’, Apr 1973,

Henderson H1927 (BRI); near Ranmoor, 35 km N of

Richmond, May 1974, Byrnes 3063 (BRI).

Distribution and habitat: Neptunia

amplexicaulis {. richmondii 1s endemic to

northern Queensland, Australia. Most records

are within 30 km of Richmond, while there is

a single record c. 45 km WNW of Hughenden

(Map 1). It grows in grassland communities

on flat or undulating terrain. The soils are

heavy clays that have a high selenium content

(Harvey ef al. 2020).

Bean, Neptunia in Australia and Malesia 69

Imm

a eh Te a

= Ped SAE telat)

Tis: 4

Vals

rus

=, 2

i= =

#500

feet

—

bo poe on be ae

a es oe er

Ls Ter

a ae se

Fig. 2. Neptunia dimorphantha. A. pods B. indumentum on pod margin. N. paucijuga. C. group of pods. N.

amplexicaulis f. richmondii. D. pods. E. pod indumentum. N. xanthonema. F. pods and peduncle bracts. G. pod

indumentum. N. heliophila. H. pods. N. major. |. pods. A & B from Hirst 49 (DNA); C from Michell & Risler 2338

(DNA); D & E from Byrnes 3063 (BRI); F & G from Bean 26282 (BRI); H from Bean 9866 (BRI); I from Clarkson

6977 & Simon (BRI). Del. N. Crosswell.

Phenology: Flowers are recorded for March,

May, September and October; fruits recorded

from March to December.

Affinities: Neptunia amplexicaulis ff.

richmondii differs consistently from WN.

amplexicaulis {. amplexicaulis only by the

indumentum. The two forms have been

recorded growing together at Silver Hills

Station (consecutive collections by McCray

and Byrnes), near Ranmoor (Byrnes 3063

& 3064) and at a gravel pit north-west of

Richmond (Byrnes 3043 & 3044), but no

intergrades are known.

Notes: Windler (1966) stated that Neptunia

amplexicaulis t. amplexicaulis has “stipels

large, leaf-like”, while in NV. amplexicaulis Tf.

richmondii they are absent. My study indicates

that leaf-like stipels are only rarely present in

N. amplexicaulis {. amplexicaulis, and hence

are of no practical value in distinguishing the

two formae.

2. Neptunia dimorphantha Domin, Biblioth.

Bot. 22(89): 802 (1926).

Neptunia dimorphantha var. dimorphantha,

Domin, Biblioth. Bot. 22(89): 802 (1926).

Type citation: “Queensland: grassige

Stellen auf den Karsthtigeln bei Chillagoe

(DOMIN II. 1910)” (translation: grassy spots

on the Karst hills opposite Chillagoe). Type:

Australia. Queensland. Cook DISTRICT:

Chillagoe, February 1910, K. Domin s.n.

(lecto: PR 527698 [here designated]; isolecto:

PR 527699).

Neptunia gracilis {. glandulosa Wiaindler,

Aust. J. Bot. 14: 416 (1966). Type: Australia.

Northern ‘Territory. Barkly Tableland, 26

miles [42 km] SW of Beetaloo, 10 March

1959, G.M. Chippendale NT5438 (syn: DNA

A0005438; NSW 404600; SIU, 7.¥.), syn. nov.

Shrub, perennial, terrestrial. Stems prostrate,

sparsely hairy, sessile glands present. Stipules

persistent, ovate, 5.2—6.3 mm long, 1.4—2.6

mm broad, coriaceous, strongly nerved,

glabrescent; apex attenuate; base obliquely

cordate. Leaves with 3-5 pairs of pinnae;

petioles 8—13 mm long, sparsely hairy, with a

small or large circular nectary just below first

pair of pinnae; rachis 15—28 mm long, with

Austrobaileya 12: 59-106 (2022)

sparse tubercle-based hairs, nectary absent;

rachis extension linear, 2—2.7 mm long; pinnae

with 14—20 pairs of leaflets, rachilla extension

0.8—1.3 mm long; stipels slender, 0.1—0.8 mm

long, bifurcated, glandular. Largest leaflets

4.2—8.5 mm long, 1.1-1.9 mm wide, 3-5.3

times longer than broad, with marginal cilia,

venation of 1-3 main veins, more obvious

on abaxial surface. Inflorescences globose,

some with dimorphic flowers; peduncles

25-90 mm long at anthesis, 42-90 mm

long at fruiting stage, with sparse, antrorse

eglandular hairs throughout, 0.25—0.5 mm

long; bracts 2, positioned on proximal one-

third of peduncle, persistent, broadly ovate,

amplexicaul, lower one 3.2—8.5 mm long,

glabrous or sparsely hairy, margin ciliate;

flowers 24—40 per inflorescence; bracteoles

oblanceolate, 1.4—1.6 mm long, caducous or

persisting until anthesis. Bisexual flowers:

calyx campanulate, 1—1.5 mm long, glabrous;

petals 1.7—2.2 mm long, glabrous; stamens

5, filaments 2.7-3 mm long, anthers 0.7—0.8

mim long, terminal gland present; style 2—2.5

mm long, glabrous; ovary hairy along the

margins only. Neuter flowers: calyx 0.5—0.8

mm long; petals 1.3—2 mm long, glabrous;

staminodes 5, 8.5—14 mm long, 0.7—0.9 mm

wide; gynoecium absent. Pods circular to

elliptical, 1-3-seeded, 8-16 mm long, 7—9.5

mm broad, indehiscent, stipe 1—2 mm long,

apex obtuse to mucronate; usually glabrous,

but with sparse to dense curved eglandular

hairs along pod margin (0.1—0.2 mm long),

most often near base of style. Seeds broadly

elliptical in outline, 3.5—3.7 mm long, 2.7—2.8

mm wide, brown. Figs. 2A—B, 4F.

Additional specimens examined: Australia. Western

Australia. At camp, Mornington Wildlife Sanctuary,

Jan 2006, Legge 73] (PERTH); Karunjie Station,

Kimberleys, Oct 1954, Rust 63K (CANB); Kimberley

Research Station, East Kimberley, Jul 1950, Langfield

217 (CANB). Northern Territory. S side of Lomarieum

Lagoon, Limmen NP, Apr 2009, Hirst 49 (DNA); 22 km

N of Connells Bore, Jun 1982, Latz 9270 (DNA); N of

Camerons Bore, Cattle Creek Station, May 2004, Cowie

10254 & Brocklehurst (DNA); Wyndham Road, 4 miles

[6 km] SW of Katherine, Jan 1964, Adams 835 (BRI,

CANB, DNA, L); Southern tributary of Broadarrow

Creek, c. 75 km SSW of Bullita Outstation, Gregory

NP, Apr 1996, Duretto 899 (DNA, MEL); 20 km W of

Kalkarindji, Mar 1992, Williams 213 (DNA); Mataranka

Reserve, May 1977, Must 1472 (DNA); Jilkminggan,

headwaters of Roper River, Feb 1990, Wightman &

Bean, Neptunia in Australia and Malesia

Jackson 4977 (DNA); c. 40 km W of Kalkarindji off the

Buntine Highway, near Burta, Mar 2012, Lewis 19/2

(CANB, DNA, NSW); 6 miles [10 km] W of Armchair

Bore, Brunette Downs, Mar 1956, Chippendale 1970

(AD, CANB, DNA); O.T. Station, May 1947, Blake

17697 (BRI, CANB). Queensland. Cook DISTRICT:

Brooklyn, 9.1 km WNW of Mt Carbine on Peninsula

Developmental Road, Feb 2008, Jensen 1607 (BRI);

21 km from Mt Surprise on O’Briens Gemfields Road,

Feb 1996, Forster PIFIS495 & Ryan (BRI); Chillagoe

Creek, Chillagoe, Jun 2021, Bean 34073 (BRI); Royal

Arch Cave track, Chillagoe — Mungana Caves NP, Mar

2008, McDonald KRM7336 & Little (BRI); Talaroo

Station, laneway to hay paddock, Mar 2017, McDonald

KRMI9III & Morrison (BRI). BURKE DISTRICT:

Finucane Island NP, 29 km NE of Burketown, May 2005,

Booth 4255 & Thompson (BRI); c. 30 km W of Burke

& Wills Roadhouse on Wills Development Road, Feb

2005, Fox IDF3438 et al. (BRI); Yelvertoft Station, c.

45 miles [72 km] SE of Camooweal, May 1963, Gittins

784A (BRI); ‘Woodlands’, on western side of Punchbowl

Road, c. 60 km NW of Julia Creek township, Apr 2001,

Johnson & Kelman s.n. (BRI [AQ733048]). NORTH

KENNEDY DISTRICT: corner Ingham Road & Ronald

Street, Garbutt, Townsville, Mar 2000, Gunther I81/2

(BRI); 1.5 km W of Bohle River on Hervey Range Road,

W of Townsville, Jan 1996, Cumming 13941 (BRI).

Distribution and habitat: Neptunia

dimorphantha is endemic to Australia

and known from a few collections in the

Kimberley region of Western Australia,

but widespread in Northern Territory and

Queensland (extending to the north-east coast

at Townsville) (Map 2). It grows in grassland

or open eucalypt woodland on grey to black

clay soils, or sometimes on loamy soil derived

from limestone, and prefers sites close to

creeks or waterholes.

Phenology: Flowers are recorded from

December to May; fruits from January to

June.

Affinities: Neptunia dimorphantha and N.

gracilis both have relatively long peduncles

and long staminodes. The former differs from

N. gracilis by the presence of a nectary on the

petiole (nectary absent for N. gracilis); the

1—3 seeds per pod (3—6 seeds for NV. gracilis);

pods glabrous except for hairs mainly on the

margins (pods either glabrous throughout or

puberulous throughout for N. gracilis); and

the peduncle with 2 bracts (no bracts or 1

bract for NV. gracilis).

Typification: In naming Neptunia gracilis

f. glandulosa, Windler (1966) cited two

specimens from one gathering as the type

(at DNA and SIU), but did not indicate either

specimen as the holotype. Those specimens,

therefore are syntypes (Turland et al. 2018,

Art. 40.2, Note 1), as is another duplicate at

NSW.

Notes: The name Neptunia dimorphantha

has for many years been misapplied to four

species with short peduncles and glandular

hairs covering the fruit surface (NV. longipila,

N. scutata, N. valida and N. xanthonema),

all newly named in this paper. The type

specimens of both N. dimorphantha and N.

gracilis {. glandulosa have relatively long

peduncles, a nectary at the top of the petiole,

and fruits with short curved eglandular hairs

mainly along the margins; they clearly belong

to the same taxon.

A specimen collected from Timor in

1962/63 by R. Cinatti (at L) was determined

by M. Lavaleye as Neptunia dimorphantha

and confirmed as that by I. Nielsen for the

Flora Malesiana (Nielsen 1992). I have

examined this flowering specimen and find

that it 1s not N. dimorphantha because of

its short peduncles and glabrous ovary. The

specimen appears to have much in common

with N. major (flowering peduncles 20—28

mm long, with two bracts; stems and leaves

virtually glabrous; large nectary at the apex

of the petiole). However, it has fewer leaflets

per pinna than NV. major and the stems on the

specimen are thin and curved, suggesting a

decumbent or prostrate habit. Neptunia major

is an erect shrub and specimens of it typically

display thick straight stems. It is not possible

at this stage to reliably classify the specimen

from Timor.

3. Neptunia gracilis Benth., /. Bot. (Hooker)

4: 355 (1841).

Neptunia gracilis var. typica Domin, Biblioth.

Bot. 22(89): 802 (1926), nom. illeg.; N.

gracilis f. gracilis, Windler, Aust. J. Bot. 14:

414 (1966). Type citation: “Australia, Bauer,

Mitchell”. Type: [Australia. New South

Wales] Interior of New Holland, s.dat. [1835

or 1836], 7. Mitchell s.n. (lecto: K 000791032

[here designated]; isolecto: P 02735407, W

0028233, W 0028231).

Neptunia depauperata Merr., Philipp. J.

Sci. Sect. C. Botany 13(3): 16 (1918). Type:

Philippines. LUZON: Ilocos Norte Province,

Burgos, 13 March 1917, M. Ramos 27169

(syn: K 000295941 i.d.v.; syn: BM 000946904

i.d.v.; syn: NY i.d.v.; syn: P 02436150 i.d.v.).

Illustrations: Windler (1966: 413); Pedley

(1983: 335); Murray (1991: 380).

Shrub, perennial, terrestrial. Stems prostrate,

glabrous, or with hispidulous hairs 0.1—0.2

mim long, or rarely with hispid hairs 0.2—0.5

mm long; sessile glands present or absent.

Stipules persistent, broadly ovate, 2.5—5.2

mm long, 1.8—2.3 mm broad, coriaceous,

strongly nerved, glabrous; apex attenuate;

base obliquely cordate. Leaves with 2 or 3(-

4) pairs of pinnae; petioles 10-17 mm long,

glabrous or hispidulous, nectary absent; rachis

9-21 mm long, with sparse tubercle-based

hairs or hispidulous hairs, nectary absent;

rachis extension linear, 1.8—2.4 mm _ long;

pinnae with 12-17 pairs of leaflets, rachilla

extension 1—].2 mm long; stipels slender, 0.4—

1.3 mm long, entire or divided, gland-tipped.

Largest leaflets 5.4—8.2 mm long, 1.1—2.2 mm

wide, 3.4—5 times longer than broad, glabrous

or with small marginal cilia, venation of 1-3

main veins, more obvious on lower surface.

Inflorescences globose, some with dimorphic

flowers; peduncles (25—)40—90 mm long at

anthesis, 50-105 mm long at fruiting stage,

glabrous throughout or with sparse tubercular

hairs; bracts absent or 1, positioned on

proximal one-third of peduncle, persistent,

ovate, amplexicaul, 2.5-9.5 mm long, with

hispid hairs on margin; flowers 12—25 per

inflorescence; bracteoles oblanceolate, 0.9—2

mm long, persisting until anthesis. Bisexual

Austrobaileya 12: 59-106 (2022)

flowers: calyx campanulate, 1.3—1.9 mm long,

glabrous; petals 2.5—2.8 mm long, glabrous;

stamens 5, filaments 2—2.5 mm long, anthers

0.7—0.9 mm long, terminal gland absent; style

2.3—3 mm long, glabrous; ovary glabrous or

hairy. Neuter flowers: calyx 0.6—0.8 mm long;

petals 1.8—2.1 mm long, glabrous; staminodes

5, 7-13 mm long, 0.5—0.9 mm wide. Pods

oblong, 3—6-seeded, 13—24 mm long, 6.5—8.5

mm broad, dehiscent along both margins,

stipe 1-2 mm long, apex obtuse; glabrous

or with dense patent hairs 0.1—0.3 mm long,

or rarely with hispid hairs 0.3—0.8 mm long.

Seeds broadly elliptical in outline, 3.5-3.9

mm long, 2.8—3 mm wide, dark brown.

Additional selected specimens examined: Philippines.

LUZON: Burgos, Ilicos Norte Province, Jul 1918, Ramos

32898 (US). Indonesia. LESSER SUNDA ISLANDS: Soenda,

Flores, Nov 1967, Schmutz 1888 (L); Manggarai, West

Flores, 8°34’S 116°377E, Mar 1982, Schmutz 5027 (L).

West Timor: Fatukopa, 9° 18S 124° 47E, Jan 1967,

Kooy 271 (L). Papua New Guinea. CENTRAL PROVINCE:

Moitaka, Dec 1964, Gillison NGF22061 (BRI, CANB,

L); Oval, UPNG Campus, Mar 1972, Stevens UPNG1371

(L). Australia. Queensland. Cook District: Mt

Molloy, Apr 1962, McKee 9/23 (BRI); Alma-den to

Mount Surprise Road, 8.8 km N of the Gulf Development

Road, Jun 2021, Bean 34105 (BRI; MEL, NSW, US,

distribuendi). BURKE DISTRICT: Bowthorn Station, 32.9

km NW of Bowthorn HS, Jun 2006, Thompson WES647

& Hogan (BRI). NORTH KENNEDY DistTrRIcT: Valley of

Lagoons, 1.8 km from the Ingham Road on the road to

Lake Lucy, Apr 1989, Clarkson 7936 & Henderson (BRI,

CNS, K). SourH KENNEDY District: Mt Blackjack,

‘Weetalaba’, Feb 1994, Bean 7322 & Forster (BRI).

LEICHHARDT District: c. 4 miles [6 km]| E of Moura,

Mar 1967, Henderson 226 (BRI). PorT CurRTIS DISTRICT:

Shoalwater Bay training area, Pine Mountain sector,

Stevens Road, Apr 2011, Halford OM271 & Bean (BRI);

State Forest 60, Rundle Range, Nov 1987, Gibson 936

(BRI). BURNETT District: 14 km from Murgon, towards

Nanango, Nov 1996, Bean 11377 (BRI, MEL). WIDE Bay

DistTRIcT: Booyal Hall, W of Childers, Nov 2001, Bean

18037 (BRI); Elliot Heads, Nov 1978, Stanley 78146 &

Ross (BRI). MARANOA District: Noondoo, Feb 1950,

Everist 4288 (BRI). DARLING Downs District: Lake

Broadwater, 25 km SW of Dalby, Nov 1984, Ballingall

NHA512 (BRI); 11 km N of Warra, Feb 1995, Fensham

2065 (BRI). Moreton District: Ipswich, Jan 1878,

Bailey s.n. (BRI [AQ235558]); Hays Landing, Wivenhoe

Dam, Feb 2020, Phillips 3022 & Phillips (BRI). New

South Wales. 10 km NNE of Ashley on the Boggabilla

Road, Mar 1987, Coveny 12531 et al. (MEL, NSW); 14

miles [23 km] N of Nyngan, Dec 1973, Cunningham 1659

(NSW); ‘Iolanthe’, 16 miles [26 km] W of Garah, Dec

1970, Solling s.n. (NSW 580902); Mitchell Highway, 6

miles [10 km] SSE of Dubbo towards Wellington, Nov

1969, Coveny 2506 (CANB, NSW).

Bean, Neptunia in Australia and Malesia

Distribution and habitat: In Malesia,

Neptunia gracilis is known from northern

Luzon in the Philippines, the islands of Flores

and Timor in Indonesia, and in the vicinity

of Port Moresby in Papua New Guinea. In

Australia, it 1s a common species in the

eastern half of Queensland south from the

Atherton Tableland, with two disjunct records

in the far north-west of the state and is also

in the north-east of New South Wales as far

south as Tuggerah Lake, near Gosford (Map

3). It has also been recorded from southern

Tarwan (Huang & Huang 1996). Away from

the coast, it typically inhabits heavy clay soils

in grassland or open eucalypt woodland, but

it also occurs in loamy or sandy-loam soils in

coastal areas.

Phenology: Flowers and fruits are mostly

recorded between October and April in

Australia, with few records from other months

of the year.

Affinities: The name Neptunia gracilis has

been broadly applied to any small-leaved

Neptunia without a nectary on the petiole, or

even with a nectary, in the case of N. gracilis

{. glandulosa. Here it is restricted to plants

having dimorphic inflorescences, leaves with

2 or 3 or sometimes 4 pairs of pinnae, long

peduncles bearing one bract or without bracts,

and bisexual flowers with 5 stamens. It is

perhaps most closely related to N. heliophila,

as both lack a petiole nectary, have neuter

flowers with five staminodes, and relatively

long peduncles. Neptunia heliophila differs

by the two bracts on the peduncle (except for

specimens from the Pilbara region of Western

Australia which frequently have only one);

stems, rachises and peduncles with antrorse

hairs (stems and peduncles either glabrous or

with patent hairs for N. gracilis), leat pinnae

4—6 pairs (2 or 3 pairs, sometimes 4 for N.

gracilis), and 25—40 flowers per inflorescence

(12—25 flowers for NV. gracilis).

Typification: Cowan (1996) published a

paper in which he typified many names

in the Mimosaceae. For Neptunia gracilis

(described by Bentham in 1841), he

designated as lectotype a specimen collected

by Thomas Mitchell in 1846. This specimen

was obviously not available to Bentham when

he was describing the species, hence it 1s

not original material and Cowan’s attempted

lectotypification 1s ineffective.

A specimen collected from Miuitchell’s

second or third expedition (which took

place in 1835 and 1836 respectively) is here

designated as the lectotype. The label of the

lectotype at K has printed on it “Interior of

New Holland/ Major Mitchell’s Expedition,

183 .. Handwritten notes have been added

saying “Neptunia gracilis Benth in Hook.

Journ. 4.355” and “Lindley 8 [1.e. 1838]”. This

indicates that the specimen was received by

Kew from John Lindley in 1838. Mitchell

had sent all his botanical specimens directly

to Lindley, who then described some of them

as new species in Mitchell’s journal (Mitchell

1838) and distributed specimens to major

herbaria.

The isolectotype at P has the identical

printed label, and the handwritten addition

says “Mlonsieur] Lindley 1838”. The

isolectotypes at W have the same printed

label, plus a handwritten “1839.V./ Lindley”

indicating that they received the specimens

from Lindley in (May’?) 1839.

Notes: Neptunia gracilis as circumscribed

here has three morphological forms; 1. the

most common form with glabrous stems,

glabrous peduncles and glabrous pods; 2.

a form with dense hairs 0.1—0.3 mm long

on stems, sparse tubercle-based hairs on

peduncles and dense puberulous hairs on

the pods; 3. an uncommon form with hispid

hairs 0.3—0.8 mm long on stems, peduncles

and pods. Forms | & 2 apparently grow in

mixed populations, as Fensham 2065 (BRI)

comprises one branchlet of each. Form 3 can

be confused with N. proxima, but the latter has

inflorescences comprised entirely of bisexual

flowers, shorter petioles and peduncles, and

smaller leaflets.

4. Neptunia heliophila A.R.Bean sp. nov.

Differing from Neptunia gracilis by the two

bracts on the peduncle; the stems, rachises

and peduncles with crisped antrorse hairs,

the greater number of flowers (25—40) per

inflorescence, and the leaves commonly

having a greater number (4—6 pairs) of

pinnae. Typus: Australia. Queensland.

MARANOA District: 7.2 km E of St George,

junction of Carnarvon Hwy [Highway] and

road to Dirranbandi, 8 December 2001, D.

Halford Q7680 & G.N. Batianoff (holo: BRI

[AQ783984 comprising | sheet]; iso: MEL

2196385A, NSW 672784).

Neptunia gracilis var. villosula Benth., FI.

Austral. 2: 300 (1864). Type: [Australia.

Western Australia/Northern Territory] Sturt’s

Creek, [in 1855], F! Mueller s.n. (lecto: K

000791029, lower right-hand specimen, fide

Cowan 1996: 14).

Shrub, perennial, terrestrial. Stems prostrate,

sprawling or erect, to | m high with sparse

to dense antrorse hairs, sessile glands present.

Stipules persistent, ovate to broadly ovate,

5.8—6.5 mm long, 2—2.8 mm broad, coriaceous,

strongly nerved, ciliate on margins; apex

attenuate; base obliquely cordate. Leaves

with 4—6 pairs of pinnae; petioles 9-13 mm

long, with sparse antrorse hairs, nectary

absent; rachis 16—37 mm long, with sparse

to dense antrorse hairs 0.2—0.5 mm _ long,

nectary absent; rachis extension linear, 2—4.5

mim long; pinnae with 15-19 pairs of leaflets,

rachilla extension 1.5—2.5 mm long; stipels

slender, 0.6—2.2 mm long, entire, gland-tipped

or not. Largest leaflets 4.6—7.8 mm long, 1.1—

2.6 mm wide, 3—5 times longer than broad,

glabrous except for marginal cilia, venation

of 1-3 main veins, more obvious on lower

surface. Inflorescences globose, some with

dimorphic flowers; peduncles 35-110 mm

long at anthesis, 45-130 mm long at fruiting

stage, with sparse to dense antrorse hairs

throughout; bracts 2 (rarely 1), positioned on

proximal one-third of peduncle, persistent,

ovate, amplexicaul, lower one 5.4—8 mm long,

with crisped hairs on margin; flowers 25—40

per inflorescence; bracteoles narrowly elliptic,

1.7-2.4 mm long, persisting until anthesis.

Bisexual flowers: calyx campanulate, 1.1—1.4

Austrobaileya 12: 59-106 (2022)

mm long, glabrous; petals 2—2.5 mm long,

glabrous; stamens 5, filaments 3.3-3.5 mm

long, anthers 0.6—0.8 mm long, terminal gland

present; style 1.8—2 mm long, glabrous; ovary

glabrous. Neuter flowers: calyx c. 0.7 mm long;

petals 1.4—1.5 mm long, glabrous; staminodes

5, 8.5-10 mm long, 0.5—0.8 mm wide. Pods

oblong to elliptical, 3—6(—8)-seeded, 13-22

mm long, 6—9 mm broad, tardily dehiscent

along both sutures, stipe 1-2 mm long, apex

obtuse; glabrous. Seeds broadly elliptical in

outline, 3.3—4.2 mm long, 3-3.5 mm wide,

dark brown to black. Figs. 2H, 3A—D.

Additional selected specimens examined: Australia.

Western Australia. Karunjie Station, Nov 1954, Rust

63 (PERTH); 30 km on Millewinde Road from Gibb

River Road, King Leopold Ranges, Apr 1988, Simon

4027 (BRI, PERTH); 2 km N of Beverley Springs

Station HS, May 1988, Cranfield 66856 (BRI, PERTH);

Mullalang Paddock, Flora Valley Station, Nov 2000,

Craig ABCI628 (PERTH); Munjina Claypan, Juna

Downs Station, 15.1 km S of Mt Lockyer, 24.8 km ENE

of Mt Windell, 37.4 km N of Packsaddle Hill, Hamersley

Range, Sep 1988, van Leeuwen 3928 (PERTH); c. 24

km S of the intersection of Nanutarra to Munjina Road

and Hamersley Road, 29.5 km NNE of Tom Price and

118.4 km SE of Silver Grass Peak, May 2011, Maier

& Adam CA006 (PERTH); Winning Pool, Oct 1941,

Gardner 6225 (PERTH). Northern Territory. c. 168

km SSW of ‘Calvert Hills’ on road to ‘Creswell Downs’,

May 1974, Pullen 9282 (CANB); Brunette Downs HS

waterhole, Nov 1999, Brock 50 (NT); 35 km SW of Eva

Downs HS, Nov 1999, Brock 37 (NT); Kidman Springs

Station fire plots, Victoria River, Apr 2016, Lebbink

GHL1I63 & Cowley (BRI); Pigeon Hole Station, May

2005, Risler & Fisher 2748 (DNA); Toplanes paddock,

Moolooloo Station, Victoria River district, May 1995,

Van Kerckhof 73 (DNA); Anthony Lagoon, Feb 1998,

Michell & Carrow 1307 (DNA). Queensland. BURKE

District: Calton Hills Station, c. 50 km N of Mt Isa,

May 2006, Booth CAM04-23 & Kelman (BRI); 12

km W of Richmond on Flinders Highway, May 2016,

McKenzie RAM16/12 (BRI). SOUTH KENNEDy DISTRICT:

2km N of Natal Downs HS, Jan 1987, Dorney 49

(BRI). GREGORY NortH District: Crawford Creek, c.

45 km SE of Winton on road to Longreach, Aug 2005,

Batianoff 0508448 & Butler (BRI, MEL). MITCHELL

District: Thomson River, 800 m SW of Waterloo HS,

Mar 1989, Emmott 268 (BRI); 30 km from Longreach

along Landsborough Highway towards Winton, May

1991, Telford 11469 (BRI, CANB). LEICHHARDT

District: Near ‘Yandina’ turn-off, Arcturus Road,

NW of Rolleston, Oct 1998, Bean /4/1/4 (BRI, NSW).

MARANOA DIstTRICT: Roma, Feb 1938, Blake /329/ (BRI,

CANB, K, MEXU, NSW); 1.7 km along Bindaroo Road,

E of Roma, Apr 2021, Bean 33909 (BRI, CANB, MEL,

NSW). DARLING Downs District: 15 km along Goodar

Road, NW of Goondiwindi, Feb 1996, Bean 9866 (BRI,

MEL); Barwon Highway, c. 35 km W of Goondiwindi,

Bean, Neptunia in Australia and Malesia

yur “Pas

Ae S,

Chae ye

ans me ay

out. C. anther. D. neuter flower

Fig. 3. Neptunia heliophila. A. bisexual flower, side view B. bisexual flower, opened

showing staminodes. N. proxima. E. pods. F. pod indumentum. N. tactilis. G. pods. N. javanica. H. pods. N. hispida.

I. pod. N. plena. J. pods. A—D from Bean 33909 (BRI); E & F from Forster PIF44127 & McDonald (BRI); G from

Brennan 6421 (DNA); H from Backer 19564 (L); I from Brennan 6913 (DNA); J from Mitchell 7041 (BRI). Del. N.

Crosswell.

Dec 2001, Halford O7SIS & Batianoff (BRI, CANB,

NSW). New South Wales. 3.6 km NW of Fort Grey

turnoff on Binerah Well — Tibooburra road, May 1977,

Donner 5704 (AD); c. 500 m upstream of South Werri

Tank, Sturt NP, Mar 1993, Logan s.n. (NSW 404883);

28 miles [45 km] N of Bourke on North Collerina Road,

Mar 1974, Millthorpe 1727 & Cunningham (NSW); S of

railway line c. 0.9 km E of the junction of the Quirindi

to Gunnedah Road and the Binnaway to Werris Creek

Railway line, Jan 2002, Hosking 2177 (CANB, MEL,

NSW). South Australia. Margaret Creek, 32 km W of

Coward Springs, Mar 1984, Badman 699 (AD).

Distribution and habitat: Neptunia

heliophila is endemic to Australia. It is

broadly distributed across northern Australia

(though sporadically) from the west coast of

Western Australia, through much of semi-

arid Northern Territory, the north of South

Australia, much of Queensland (as far east as

Collinsville) and extending south to Werris

Creek in New South Wales (Map 4). It grows

in heavy clay soils in grassland or open

eucalypt woodland.

Phenology: Flowers and fruits are recorded

from every month of the year except

September.

Notes: Neptunia heliophila has the most

conspicuous stipels of all the Australian and

Malesian taxa; they are frequently more than

1 mm long and can reach 2.2 mm 1n length.

Some populations from the Maranoa

district of Queensland and Werris Creek

area, N.S.W. (Hosking 2177) are notable

for their upright or ascending habit, and for

occurring in dense colonies, with adjacent

plants presumably connected by rhizomes. A

specimen from the Northern Territory (Pullen

9282) 1s likewise described as an “erect

annual to 80 cm high’.

Etymology: From the Greek helios (sun), and

philios (loving). This refers to the distribution

of this species in the part of Australia that

receives the most sunlight annually.

5. Neptunia hispida A.R.Bean sp. nov.

With affinity to Neptunia gracilis but differing

by the two lanceolate bracts positioned on the

middle one-third of the peduncle; the 9 or 10

stamens; the leaves with 3—5 pairs of pinnae,

and most vegetative parts with long hispid

hairs. Typus: Australia. Northern Territory.

Austrobaileya 12: 59-106 (2022)

Litchfield National Park, 16 March 1995,

[.D. Cowie 5319 & S. Taylor (holo: DNA

[D0122064 comprising | sheet]; 1so: MEL

0282099A).

Neptunia ‘Darwin’, pro parte: Dunlop et al.

(995-27);

Shrub, perennial, terrestrial. Stems prostrate,

with patent hispid hairs 0.4—0.8 mm _ long,

sessile glands absent. Stipules persistent,

ovate, 2.8—3.5 mm long, 1.3—2.1 mm broad,

chartaceous, faintly nerved, sparsely hairy;

apex acute; base cordate, symmetrical.

Leaves with 3—5(—6) pairs of pinnae; petioles

8—13 mm long, hispid, nectary absent; rachis

12-33 mm long, with sparse patent hispid

hairs, nectaries absent; rachis extension

linear, 1.9-2.6 mm long; pinnae with 16—25

pairs of leaflets, rachilla extension 1.4—2.3

mm long; stipels slender, 0.1—0.3 mm long,

entire, gland-tipped. Largest leaflets 4.1—6

mm long, 0.8-2 mm wide, 3-—5.]1 times

longer than broad, with a few marginal cilia,

venation of 1—3 main veins, more obvious on

lower surface. Inflorescences globose, all with

dimorphic flowers; peduncles 16—45 mm long

at anthesis, 25—47 mm long at fruiting stage,

with sparse hispid hairs; bracts 2, positioned

on middle one-third of peduncle, usually

persistent, lanceolate, + equal, base obtuse,

lower one 2.2—4.1 mm long, sparsely hairy

on margins; flowers 18—26 per inflorescence;

bracteoles oblanceolate, 0.9-1.6 mm _ long,

caducous, usually falling before anthesis.

Bisexual flowers: calyx campanulate, 1.5—2.2

mm long, ciliate; petals 2.5-3.1 mm long,

glabrous; stamens 9-10, filaments 5-—5.2

mm long, anthers 0.7-1 mm long, terminal

gland prominent; style 3.3-5.5 mm long,

glabrous; ovary glabrous. Neuter flowers:

calyx 1.8—2.2 mm long; petals 1.9—2 mm long,

glabrous; staminodes 7—10, 6.2—6.4 mm long,

0.6—0.8 mm wide. Pods oblong to elliptical,

3—7-seeded, 18—31 mm long, 7-8 mm broad,

dehiscent along one margin only, apex acute

to obtuse, stipe 2—6 mm long; glabrous.

Mature seeds not seen. Figs. 31, 4B, 4D.

Additional specimens examined: Australia. Northern

Territory. Melville Island, Nov 1986, Fensham 380

(DNA); [ibid, Melville Island, Apr 1986, Fensham 102

(DNA); ibid, Apr 1986, Johnson 4203 (BRI, DNA)|;

Miulikipiti Road, Melville Island, Feb 2018, Hirst 359

Bean, Neptunia in Australia and Malesia

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Fig. 4. Neptunia longipila. A. inflorescence, with peduncle and bracts. N. hispida. B. inflorescence, with peduncle and

bracts. N. valida. C. group of pods, with peduncle and bracts. N. hispida. D. indumentum on leaf rachis and rachilla. N.

monosperma. E. section of leaf rachis showing extra-floral nectaries and lack of stipule. NV. dimorphantha. F. section

of leaf rachis showing extra-floral nectary and persistent stipule. A from van Leeuwen et al. PBS2080 (PERTH); B

from Fensham 380 (DNA); C from Roberts APR1I079 (PERTH); D from Cowie 5319 & Taylor (DNA); E from Bean

15177 (BRI); F from McDonald KRM7336 & Little (BRI). Del. N. Crosswell.

(DNA); Maclear Creek Road, | km S of Notch Creek,

Melville Island, Jan 1992, Leach 294] & Cowie (BRI);

1 km E of Punelli, Melville Island, Feb 1992, Fensham

1207 (DNA); Emu Plains, Melville Island, Dec 1984,

Jones 1697 (DNA, PERTH); Port Darwin, in 1886,

Tenison-Woods 328 (MEL); Cobourg Peninsula, Apr

2006, Brennan 7141 (DNA); Gurig NP, Apr 2006,

Brennan 6913 (DNA); Charles Darwin NP, Feb 1998,

Harwood 315 (DNA); Channel Island Road, 1 km E

of Elizabeth River, Palmerston, Jan 1992, Cowie 2201

(DNA); power line track between Cox Peninsula Road

and Finn Road, Berry Springs, Jan 2016, Wirf 955 &

Holland (CANB, DNA); Berry Creek, Dec 1974, Must

1323 (DNA); Litchfield NP, 15 km WSW of Adelaide

River township, Mar 1996, Booth 1495 & Cowie (DNA);

Bulman Community near Telstra site, Nov 2009, Murfet

6651 (AD, DNA); 0.25 miles N of Ooloo crossing, Daly

River levee, Jun 1964, Robinson R599 (DNA); between

Douglas River and Block, Jan 1964, Robinson R244

(DNA); c. 21 miles [33.8km] SE of Oenpelli Mission, Feb

1973, Lazarides 7779 (BRI, CANB, DNA); Noonamah

area, Feb 1963, Muspratt SSO249 (DNA); Glenluckie

Creek, Stuart Highway, Dec 2009, Brennan 8230 (DNA).

Distribution and habitat: Neptunia hispida

is endemic to the “Top End’ of the Northern

Territory, Australia where it occurs from

Melville Island to Katherine and from Daly

River to Bulman (Map 5). It is most often

recorded from lateritic slopes and ridges

with gravelly or skeletal soil, although one

specimen from Melville Island was said to

be from a “cracking clay plain’. Dominant

eucalypts include Eucalyptus miniata

A.Cunn. ex Schauer, Corymbia_ bleeseri

(Blakely) K.D.Hiull & L.A.S.Johnson, £.

oligantha Schauer and E. tetrodonta F.Muell.

Phenology: Flowers are recorded from

November to April; fruits between December

and April.

Affinities: Neptunia hispida 1s closely related

to N. tactilis, these being the only two

Australian species with more than 5 stamens

per flower. Neptunia hispida differs by the

3—5(—6) pairs of leaf pinnae (1-3 pairs for N.

tactilis), the presence of bracts on the peduncle

(vs. absent for N. tactilis); the presence of long

spreading hispid hairs on the stems, rachises

and peduncles (glabrous or sparsely hairy for

N. tactilis); and the dimorphic inflorescences

(vs. all bisexual for N. tactilis).

Austrobaileya 12: 59-106 (2022)

Notes: Neptunia hispida is a prostrate plant.

The leaves have 3—5 pairs of pinnae and

no petiole nectaries. The stems, rachises,

leaves, peduncles, bracteoles and calyces

have conspicuous long spreading hispid hairs,

and there are two narrow bracts positioned

on the middle one-third of the peduncle; the

inflorescences are dimorphic with neuter

flowers at the base; anthers are 0.7—1 mm long

with a prominent terminal gland.

A specimen from Elsey Station (Menkhorst

1223, DNA) is similar to Neptunia hispida but

differs by the fruits that are very hairy and

have an obtuse apex. It could represent an

unnamed taxon.

Etymology: From the Latin hispidus, meaning

‘covered with stiff hairs’. The stems and

leaves of this species are densely covered in

stiff patent hairs.

6. Neptunia insignis A.R.Bean sp. nov.

With affinity to Neptunia amplexicaulis

but differing by the long spreading hairs

on the stems, peduncles and rachises, the

longer fruiting peduncles, the smaller bracts

and the smaller stipules. Typus: Australia.

Queensland. Cook DISTRICT: 3.7 km by road

from Elizabeth Creek towards Chillagoe, 6

March 2022, K.R. McDonald KRM22410 &

L. Little (holo: BRI [comprising | sheet]; iso:

AD, BM, CANB, DNA, L, MEL, NSW, US

distribuendi).

Shrub, perennial, terrestrial. Stems prostrate,

with dense patent hairs 0.4—0.8 mm long,

sessile glands absent. Stipules persistent,

broadly ovate, 4-7.5 mm long, 1.8—5 mm

broad, coriaceous, strongly nerved, hairy on

both surfaces; apex attenuate; base obliquely

cordate. Leaves with 2 or 3 pairs of pinnae;

petioles 15-24 mm long, hispid, nectary

absent; rachis 14-28 mm long, with hispid

hairs, nectary absent; rachis extension linear,

2.8-3.8 mm long; pinnae with 8-14 pairs of

leaflets, rachilla extension 2—2.6 mm long;

stipels slender, 0.9-—2 mm long, not divided,

not gland-tipped. Largest leaflets 8.2—-13 mm

long, 1.8-—4 mm wide, 2.9—4.9 times longer

Bean, Neptunia in Australia and Malesia

than broad, surfaces + glabrous, margin with

many cilia, venation of 1—3 main veins, more

obvious on lower surface. Inflorescences

globose, some with dimorphic flowers;

peduncles 33-95 mm long at anthesis, 61—

110 mm long at fruiting stage, with patent

hairs throughout, 0.4—0.8 mm long; bracts 2,

positioned on proximal one-third of peduncle,

persistent, broadly ovate, amplexicaul,

lower one 5—9 mm long, with hispid hairs

on lower surface and margin; flowers 25-35

per inflorescence; bracteoles oblanceolate,

1.4—1.7 mm long, persistent until anthesis.

Bisexual flowers: calyx campanulate, 1—1.3

mm long, glabrous or with a few hairs;

petals 2.3—2.5 mm long, glabrous; stamens

5, filaments c. 3 mm long, anthers c. 0.8

mim long, terminal gland absent; style c. 2.2

mim long, glabrous; ovary glabrous or hairy.

Neuter flowers: calyx c. | mm long; petals c.

2.1 mm long, glabrous; staminodes 5, 4.5—7

mm long, 0.3—0.4 mm wide. Pods oblong to

broadly elliptical, 3—/7-seeded, 13-22 mm

long, 9-11 mm broad, dehiscent along both

margins, stipe 1—-1.5 mm long, apex obtuse;

with dense patent eglandular hairs c. 0.4 mm

long, or short hairs c. 0.1 mm long, confined to

margins, or glabrous. Seeds broadly elliptical

in outline, 4—4.5 mm long, 3.5—3.7 mm wide,

dark brown. Figs. 5, 6.

Additional — specimens examined: Australia.

Queensland. Cook District: 4.9 km along Bellevue

Station access road, Mar 2022, McDonald KRM22415 &

Little (BRI; B, CANB, MEL, MO, P, PRE distribuendi);

Access road to Bellevue, NNW of Chillagoe, Apr 2002,

Bean 18740 & McDonald (BRI); 3 km E of Elizabeth

Creek on road to Chillagoe, Jan 1982, Clarkson 4256

(BRI, P); South of The Oasis on Greenvale Road, Jun

1999, Addicott 171 (BRI). NoRTH KENNEDY DISTRICT:

Lake Lucy, s.dat., Anon. 78 (MEL). SOUTH KENNEDY

District: Redcliffe Vale on Redcliffe Tableland, Apr

2012, Champion IGC2139 et al. (BRI).

Distribution and habitat: Neptunia insignis

is endemic to Queensland, Australia. It

extends from north-west of Chillagoe to

Redcliffe Tableland, north-west of Mackay

(Map 6). It grows on flat or gently undulating

terrain, in cracking clay soil, in open eucalypt

woodland.

Phenology: Flowers are recorded for January

and March; fruits for January, March, April

and June.

Fig. 5. Flowering branchlet of Neptunia insignis

(McDonald 22415 & Little, BRI). Photo: K.R. McDonald.

Affinities: Neptunia insignis 1s most similar

to N. amplexicaulis as they both have 2 or 3

pairs of pinnae, few leaflets per pinna (7—15

pairs), no nectaries on the petiole, and large

leaflets (8.2-16 mm long). Neptunia insignis

differs by the long spreading hairs on the

stems and peduncles (vs. glabrous or with

short puberulous hairs for NV. amplexicaulis);

the stipules 4—7.5 mm long (vs. 16-19 mm

long for N. amplexicaulis); the bracteoles

and calyx with a few hairs (vs. glabrous in N.

amplexicaulis); the lower bracts 5—9 mm long

(vs. 11-19 mm long for NV. amplexicaulis); and

the fruiting peduncles 61-110 mm long (vs.

36-52 mm long for NV. amplexicaulis). One

of the forms of N. gracilis has coarse hispid

hairs on the rachises and peduncles as in N.

insignis, but the latter differs consistently by

the larger leaflets and the presence of two

bracts on the peduncle.

80 Austrobaileya 12: 59-106 (2022)

Fig. 6. Fruiting branchlet of Neptunia insignis (McDonald 22410 & Little, BRI). Photo: K.R. McDonald.

Bean, Neptunia in Australia and Malesia

Note: The 3-—/-seeded pods of Neptunia

insignis may be sparsely hairy at the base

only (Bean 18740, McDonald KRM22415 &

Little), or on the margins (Clarkson 4256),

or hairy throughout (Addicott 171, Anon. 78,

Champion IGC2139 et al.).

Conservation status: Neptunia insignis 1s

suggested to qualify as ‘Endangered’ under

Red List Criterion B IUCN 2012). The area of

occupancy is far less than 500 km*, there are

five known locations, and there is a projected

continuing decline in area, extent and/or

quality of habitat, due to weed incursion,

especially by Grader grass (Themeda

quadrivalvis (L.) Kuntze), and heavy grazing

by cattle.

Etymology: From the Latin insignis meaning

‘distinguished’ or ‘conspicuous’. This species

is quite conspicuous by virtue of its relatively

large leaflets and bracts.

7. Neptunia javanica Mig., Fil. Ned. Ind. 1(1):

51 (1855).

Type citation: “Java, op heuvels van G.

Sahari, G. Soerabaja”. Type: Indonesia. G.

Saharie, near Surabaya, Java, 9 January

1847, H. Zollinger 3231 (lecto: P 02735395

[here designated]; isolecto: P 02735394, P

02735393).

[Neptunia acinaciformis auct. non (Span.)

Mig. (1841): Windler (1966: 393)]

Illustration: Windler (1966: 394), as WN.

acinaciformis.

Shrub, perennial, terrestrial. Stems prostrate,

glabrous, sessile glands absent. Stipules

persistent, ovate, 2-3.3 mm long, 0.8—1.2

mm broad, chartaceous, with a single nerve,

glabrous; apex attenuate; base obliquely

cordate. Leaves with (2—)3(—4) pairs of pinnae;

petioles 3—8 mm long, glabrous, with | small

nectary in the middle one-third, and rarely a

second one near the distal end; rachis 13-16

mm long, glabrous, nectary absent; rachis

extension linear, 0.8—1 mm long; pinnae with

9-15 pairs of leaflets, rachilla extension 1.3-—

2.5 mm long; stipels slender, 0.1—0.3 mm long,

entire, gland-tipped. Largest leaflets 2.9—4.5

mm long, 0.9-1.3 mm wide, 3.2—4.1 times

longer than broad, glabrous, venation of 1-3

8 |

main veins, more obvious on lower surface.

Inflorescences globose, some with dimorphic

flowers; peduncles 25—57 mm long at anthesis,

40—70 mm long at fruiting stage, glabrous;

bracts absent; flowers 8—12 per inflorescence;

bracteoles oblanceolate to obovate, c. 0.5

mm long, caducous, usually falling before

anthesis. Bisexual flowers: calyx obconical,

1.1-1.4 mm long, glabrous; petals 1.8—2.4 mm

long, glabrous; stamens 10, filaments c. 2.2

mm long, anthers 0.6—0.9 mm long, terminal

gland obscure; style c. 2.5 mm long, glabrous;

ovary glabrous. Neuter flowers: calyx 0.5—0.9

mm long; petals 0.9-1 mm long, glabrous;

staminodes 10, 1.8-—3 mm long, 0.2—0.4 mm

wide. Pods falcate, 8—ll-seeded, 26-33 mm

long, 7./—9 mm broad, dehiscing along one

suture only, stipe 5—7 mm long, apex acute;

glabrous. Seeds broadly elliptical in outline,

c. 4.5 mm long and 2.6 mm wide, brown. Fig.

3H.

Additional specimens examined: Indonesia. LESSER

SUNDA ISLANDS: Orong, Flores, 8°29’S 120° 39’E, Apr

1966, Schmutz 34 (L). JAvA: Soerbaja, Grisee, 7°09’S 112°

37°E, Feb 1924, Dorgelo 2292 (L); Koepang, Soerabaja,

May 1922, Dorgelo 377 (L); Sampang to Rapa, Madoera,

7°00’S 113°20’°E, Mar 1915, Backer 19773 (L); Sampang,

Madoera, 7° 00’S 113° 20’E, Mar 1915, Backer 19564

(BRI, L); Kesongo, Rembang, 7° 36’S 110° 50’E, s.dat.,

van Steenis 17436 (L); Mons Lengis, East Java, 7°09’S

112° 39°E, Apr 1932, Coert 1097 (L); road to Rapa,

Sampang, Madura, 7° 11’°S 113° 14E, Jan 1925, Rant s.n.

(L 2042660).

Distribution and habitat: In Malesia,

Neptunia javanica 1s known only from Java

and Flores (Map 3). Elsewhere, it is recorded

for Thailand, Cambodia and Vietnam. It

inhabits salty mud-plains and grassy fields

on heavy clay at altitudes below 650 metres

(Nielsen 1992).

Phenology: Flowers have been recorded in

March and April; fruits have been recorded in

January, February and March.

Affinities: Neptunia javanica 1s a terrestrial,

prostrate shrub with small leaflets and flowers

with 10 stamens. Itis similar to bothN. gracilis

and N. tactilis. From N. gracilis, it differs by

the presence of a petiole nectary (absent for

N. gracilis), flowers with 10 stamens (always

5 for N. gracilis); and the falcate acute pods

containing 8-11 seeds (pods oblong, apex

obtuse, pods with 3—6 seeds for N. gracilis).

From JN. tactilis, it differs by the leaf pinnae

with 9-15 pairs of leaflets (15-28 pairs

for N. tactilis); the presence of dimorphic

inflorescences (inflorescences all bisexual

in N. tactilis); and the presence of a petiole

nectary (absent for N. tactilis).

Typification: Nielsen (1981) cited a specimen

at Utrecht (U 164863) as the holotype of

Neptunia javanica, and he annotated it as

such. However, that specimen does not match

the protologue. In particular, the protologue

states that the fruits are 1.25 inches long,

while the sole fruit on U 164863 is 5 mm (0.2

inches) long. The “Index of Botanists” (HUH

2021) states that while Miquel’s herbartum and

types are at U, new names are “also based on

specimens at G, K, L (most), P”. A specimen

at P collected by Heinrich Zollinger (number

3231) matches the protologue with respect to

the dimensions of leaves, peduncles and fruit,

and its label includes text that is replicated in

the protologue. For example, Zollinger’s label

states “Fl. lute1 odorati” and the protologue

states “Flores odorati lutei”’. Zollinger’s

label includes “Daun littie Jav.” (perhaps an

indigenous name’), and the protologue states

“Daun litti jav.”. Furthermore, Zollinger’s

locality of ‘G. Saharie’ is cited in the

protologue. This specimen is designated as

the lectotype, while two other specimens

bearing the number 3231 are isolectotypes.

8. Neptunia longipila A.R.Bean sp. nov.

With affinity to Neptunia scutata but differing

by the non-glandular peduncle hairs 0.5—0.9

mm long, the bracts 7.5—10.4 mm long, and the

longer peduncles. Typus: Australia. Western

Australia. 20—25 km N of Fortescue Valley on

main Port Hedland Road from Wittenoom/

Newman, East Chichester Range, May 1988,

K.L. Tinley 3324 (holo: PERTH 1848410,

comprising | sheet).

Shrub, perennial, terrestrial. Stems prostrate,

with dense patent to antrorse hairs, sessile

glands absent. Stipules persistent, broadly

ovate, 5.9-8.2 mm long, 3—4.3 mm broad,

coriaceous, strongly nerved, sparsely hairy;

apex attenuate; base obliquely cordate. Leaves

with (3—)4 or 5 pairs of pinnae; petioles 9-12

mim long, with patent hairs, with a prominent

Austrobaileya 12: 59-106 (2022)

nectary around halfway along or just below

first pair of pinnae, rarely a second nectary

present; rachis 23-35 mm long, with dense

eglandular hairs, nectary absent; rachis

extension linear, 2.4—4.5 mm long; pinnae

with 17—22 pairs of leaflets, rachilla extension

1.2-1.8 mm long; stipels 0.1—0.2 mm long,

entire, gland-tipped. Largest leaflets 5—8.8

mm long, 1.2—2.3 mm wide, 3.8—4.6 times

longer than broad, glabrous on upper surface,

sparsely hairy on margins and lower surface,

venation of 1-3 main veins, more obvious

on lower surface. Inflorescences broadly

ellipsoidal, some with dimorphic flowers;

peduncles 27—52 mm long at anthesis, 28—69

mm long at fruiting stage, with dense patent

hairs throughout, 0.5—0.9 mm long; bracts 2,

positioned on proximal one-third of peduncle,

persistent, ovate, amplexicaul, lower one 7.5—

10.4 mm long, sparsely hairy, margin ciliate;

flowers 30—50 per inflorescence; bracteoles

oblanceolate, 1.5-1.8 mm long, persistent

until anthesis. Bisexual flowers: calyx

campanulate, 1.1-1.2 mm long, glabrous;

petals 1.7-2 mm long, glabrous; stamens 5,

filaments 2.5—3 mm long, anthers 0.6—0.7 mm

long, terminal gland absent; style 1.9—2.1 mm

long, glabrous; ovary hairy. Neuter flowers:

calyx 0.7—0.8 mm long; petals 1.3-1.5 mm

long, glabrous; staminodes 5, 5.2—6 mm long,

0.3—0.5 mm wide. Pods broadly elliptical,

l-seeded, 7.5—9 mm long, 5.5—6 mm broad,

indehiscent, stipe 0.5-1 mm long, apex

obtuse; with dense patent hairs 0.2—0.4 mm

long, at least some with glandular apex. Seeds

broadly elliptical in outline, 3.6—4 mm long,

2.7—3 mm wide, brown. Fig. 4A.

Additional specimens examined: Australia. Western

Australia. Fortescue River, in 1895, Cusack 211 (MEL);

11.1 km W of junction with main Millstream-Chichester

NP entry road on Millstream-Pannawonica Road, 14.6

km NNE of Mt Flora, 72.9 km E of Pannawonica,

Pilbara, Apr 2004, Leeuwen et al. PBS2081 (PERTH); 12

km SSW of Mt Herbert, Millstream-Chichester NP, May

1997, Trudgen METIS127 (PERTH); 10.5 km SSW of

Mount Herbert, Millstream-Chichester NP, May 1997,

Trudgen METI8/32 (PERTH); 16 km S of Barowina

Hill, Mar 1984, Newbey 9934 (PERTH); 7.2 km N

of Hamersley Iron Railway crossing on Roebourne-

Wittenoom Road, Millstream NP, Jul 2002, Wajon 444

(PERTH); c. 1 km W of Mt Florence HS on Roebourne

Road, Apr 1995, Mitchell PRP249 (PERTH); 11.6 km

ESE of Mt Sabine, Maiullstream-Chichester NP, Apr

1997, Trudgen 15196 (MEL, PERTH); 2.9 km N of Tom

Bean, Neptunia in Australia and Malesia

Price-Dampier Railway crossing on the Roebourne-

Munjina Road, 4 km N of Barowanna Hill, 69.4 km S

of Roebourne, Millstream-Chichester NP, Pilbara, Apr

2004, Leeuwen et al. PBS2080 (PERTH); 6.6 km SSE

of West Angela Hill, Hamersley Ranges, Jun 1997,

Trudgen 15805 (AD, PERTH); 11.7 km NE of Mt Regal,

15.9 km SE of White Peak, 3.7 km S of Karratha, Pilbara

IBRA, Aug 2005, Leeuwen et al. PBS5S87 (PERTH);

Millstream NP, Sep 2013, Davis & Dillon s.n. (PERTH

8562121); 8.7 km WSW of Mt Herbert, Millstream-

Chichester NP, Hamersley Ranges, Sep 1997, Trudgen

METISI31 (PERTH); 560 m N on Karratha Road from

the North West Coastal Highway, 1.6 km S of the

Coolawanyah Road turnoff, 2.8 km SE of Stove Hull,

2.5 km WSW of Tobacco Well, 3.9 km E of Tom Well,

Aug 2002, Leeuwen 5099 (CANB, PERTH); 2.9 km W

along track (from the main track) just south of the (old)

Robe camp. 4.7 km ESE of West Angela Hill, Hamersley

Ranges, Jun 1997, Trudgen MET1IS128 (PERTH).

Distribution and habitat: Neptunia longipila

is endemic to Western Australia in Australia

and confined to the Pilbara bioregion, from

Karratha to Newman (Map 7). It grows in

reddish-brown to orange-brown cracking

clay, or red to brown gravelly loam, on flat

or very gently sloping terrain, in grassland or

open shrubland.

Phenology: Flowers and fruits are recorded

from March to September.

Affinities: Neptunia longipila is_ clearly

allied to N. scutata, with which it shares the

glandular-hairy one-seeded fruits and the

leaves with 3—5 pairs of pinnae. It differs from

N. scutata by the generally longer peduncles

27-52 mm long at anthesis (8—35 mm long for

N. scutata), the hairs on the peduncle 0.5—0.9

mm long and eglandular (hairs 0.1—0.3 mm

long and glandular for V. scutata), the broadly

ellipsoidal inflorescences (globose for N.

scutata) and the peduncle bracts 7.5—10.4 mm

long (2.5—8 mm long for NV. scutata).

Etymology: From the Latin /ongus meaning

long and pilus meaning hair. This refers to

the long pilose hairs on the peduncles and

the leaflet margins, much longer than in the

related species N. scutata.

9, Neptunia major (Benth.) Windler, Aust. J.

Bot. 14: 408 (1966).

Neptunia gracilis var. major Benth., FI.

Austral. 2: 300 (1864). Type: Australia.

Queensland. [PorT CurRTIS DISTRICT:|

Rockhampton, s.dat., J. Dallachy s.n. (lecto:

K 000791035, fide Cowan (1996: 14); isolecto:

MEL 596304; P 02735402 i.d.v., right-hand

specimen only; P 02735403 i.d.v., right-hand

specimen only).

Illustrations: Windler (1966: 409); Wheeler

(1992: 342).

Shrub, annual, terrestrial. Stems erect, 1—2(—

3) m high, glabrous, thick, sessile glands

present. Stipules deciduous, visible only on

new growth, lanceolate to ovate, 3—4.8 mm

long, 0.8—1.4 mm broad, chartaceous, weakly

nerved, glabrous; apex acute; base obtuse

or slightly oblique. Leaves with 2—5 pairs of

pinnae; petioles 10-18 mm long, glabrous,

with a prominent nectary (Sometimes two)

in the upper half, often just below first pair

of pinnae; rachis 10-35 mm long, glabrous,

nectary sometimes present below some or all

pairs of pinnae; rachis extension linear, 2—2.8

mm long; pinnae with 23-39 pairs of leaflets,

rachilla extension I—1.7 mm _ long; stipels

slender, 0.1—0.5 mm long, entire or divided,

gland-tipped. Largest leaflets 5.4—7.1 mm

long, 1—1.6 mm wide, 4—5.5 times longer than

broad, glabrous or with a few marginal cilia,

venation of 1-3 main veins, more obvious

on lower surface. Inflorescences globose,

some with dimorphic flowers; peduncles

15—55 mm long at anthesis, 25—60 mm long

at fruiting stage, glabrous throughout; bracts

2, positioned on middle one-third of peduncle,

deciduous, ovate, amplexicaul, lower one

2.5—4 mm long, glabrous or with a few hairs

on margin; flowers 25-35 per inflorescence;

bracteoles oblanceolate, 1—-1.4 mm_ long,

caducous, usually falling before anthesis.

Bisexual flowers: calyx campanulate, 1.2—1.3

mm long, glabrous; petals 1.9—2 mm long,

glabrous; stamens 5, filaments 2.7-3.2 mm

long, anthers 0.7—0.8 mm long, terminal

gland absent; style 1.6—3 mm long, glabrous;

ovary glabrous. Neuter flowers: calyx 0.9-1

mm long; petals 1.9-2 mm long, glabrous;

staminodes 5, 6.5—11 mm long, 0.6—0.8 mm

wide. Pods oblong, (1—)2—5-seeded, 8—23 mm

long, 8—l11.5 mm broad, indehiscent, stipe

1—2.5 mm long, apex obtuse; glabrous. Seeds

broadly elliptical to broadly ovate in outline,

3.9-4.4 mm long, 3.4—-3.9 mm wide, brown.

Fig. 21.

Additional selected specimens examined: Australia.

Western Australia. Fitzroy River, 1881, Paterson s.n.

(MEL 596333); Kimberley Research Station, Ord River,

Apr 1956, Burbidge 5177 (CANB); Yanunara Gap, Napier

Range, May 1981, Demarz 8536 (CANB); Riverfarm

Road, Kununurra, May 1993, Mitchell 2853 (BRI); near

Mount House Station, Apr 1955, Lazarides 5166 (AD,

BRI, MEL, NSW, PERTH). Northern Territory. Port

Darwin, Oct 1888, Holtze 900 (MEL); Mouth of Daly

River, near Palmerston Island, Feb 1994, Leach 4007

(BRI, DNA); Wyndham Road, 4 miles [6 km] SW of

Katherine, Jan 1964, Adams 534 (CANB, NSW); 5 miles

[8.3 km] NE of Leguna Station, Jul 1949, Perry 2583

(CANB). Queensland. Cook District: 2.9 km N of the

Palmerville Road turnoff the Peninsula Development

Road, Fairview Station, Apr 2014, Thompson ST14019

(BRI, MEL, NSW); 18.5 km NNW of Bizant HS, 2 km

W of Bizant River, Apr 1992, Neldner 3796 & Clarkson

(BRI, DNA, NSW, PERTH); Lakefield NP, 12 km

from Jam Tin Creek on the track from Bizant to Jane

Tableland, May 1987, Clarkson 6977 & Simon (BRI);

Talaroo Station, Top Horse paddock boundary with

Eveleigh, Mar 2017, McDonald KRMI19147 & Morrison

(BRI). NORTH KENNEDY DistRIcT: Outskirts of Gumlu,

NW of Bowen, May 2000, Bean 166/4 (BRI, MEL,

PRE); Oonoonba, Feb 1980, Stanley 80204 (BRI); near

Guthalungra, Oct 1950, Blake 1861/7 (BRI, CANB, K);

Gregory Developmental Road, c. 85 km N of Charters

Towers, Jul 2017, Gardiner CPGT56I1 (BRI). Port

CuRTIS DIstTRIctT: 7.5 km from Bruce Highway at Bajool,

towards Port Alma, Apr 2012, Bean 3201] (BRI); Duck

Pond reserve, Rockhampton, Jan 2000, McCabe s.n.

(BRI [AQ550902]).

Distribution and habitat: Neptunia major

is endemic to Australia, where it is known

from near Derby in Western Australia to near

Borroloola in the Northern Territory, and

from Princess Charlotte Bay to Rockhampton

in Queensland (Map 8). It grows in grassland

or open eucalypt woodland on_black-soil

plains, usually on the margins of ephemeral

freshwater lagoons or gilgais, but sometimes

on the edge of marine saltpans 1n association

with Sporobolus virginicus (L.) Kunth.

Phenology: Flowers and fruits are most

commonly recorded from December to May,

but with a few records from other times of the

year.

Affinities: Neptunia major 1s closely related

to N. monosperma as both are upright

plants with glabrous stems and leaves, leaf

pinnae with numerous pairs of leaflets, and

deciduous stipules. Neptunia monosperma

differs by the peduncles at anthesis 4-12 mm

long (15-55 mm long for N. major); fruiting

peduncles 5—21 mm long (25—60 mm long for

Austrobaileya 12: 59-106 (2022)

N. major); 10—20 flowers per inflorescence

(25-35 flowers for NV. major); staminodes 2.5—

3.5 mm long (6.5—11 mm long for N. major);

and pods strictly l-seeded ((1 or)2—5-seeded

for NV. major).

Typification: Cowan (1996) selected as

the lectotype of Neptunia major, a fruiting

specimen at K collected at Rockhampton

by John Dallachy. Two specimens similar

in appearance, both bearing ripe fruits and

collected from Rockhampton are present at

P. The specimen P 02735402 is attributed to

“Ferd. Mueller” because it is Mueller’s name

that appears on the printed label, but Mueller

never visited Rockhampton, and the specimen

was almost certainly collected by Dallachy.

The sheet has two Neptunia branches mounted

on it. The one on the right 1s N. major; the

one on the left is N. gracilis. P 02735403

(right-hand branch) is without collector, but

it matches the other specimen very well and

looks like a duplicate of the other. The two

branches on the left, from “between the

Darling and Coopers Creek” appear to belong

to N. heliophila. The specimen at MEL has

a mounted branch bearing leaves and flower-

buds and there are several mature fruits in a

fragment packet. It bears a handwritten label

by Dallachy giving the collection date as

December 26, 1862.

Note: Neptunia major 1s the tallest species

of Neptunia, typically 1—2 metres high, but

reputed to attain 3 metres in height.

10. Neptunia monosperma F.Muell. ex

Benth., Fl. Austral. 2: 300 (1864).

Type: Australia. [Northern Territory.|

Victoria River, [1855-56], F.2 Mueller s.n.

(lecto [second-step, here designated]: K

000791027; isolecto: MEL 596309, branch ‘A’

only).

Illustrations: Windler (1966: 411); Wheeler

(1992: 342); Cowan (1998: 22, fig. 36 L—N).

Herb, perennial, terrestrial. Stems ascending

to erect, to 0.5 m high, glabrous, sessile glands

absent. Stipules deciduous, visible only on

new growth, narrowly ovate, 1.5—2.8 mm

long, 0.5—1.1 mm broad, chartaceous, without

nerves or with the midvein visible, glabrous;

Bean, Neptunia in Australia and Malesia

apex acute, base truncate or obliquely cordate.

Leaves with (1—)2 or 3(—4) pairs of pinnae;

petioles 5—1]1 mm long, glabrous, with one or

two prominent nectaries, one about halfway

along, the other (Sometimes absent) distal;

rachis 7-32 mm long or absent in 1-jugate

leaves, glabrous, nectaries present, 1 below

each pair of pinnae; rachis extension deltate,

0.83.5 mm long, glabrous; pinnae with

23-34 pairs of leaflets, rachilla extension

0.9—1.2 mm long; stipels slender, 0.1—0.3 mm

long, entire, sometimes faintly gland-tipped.

Largest leaflets 6.3-10 mm long, 1.5-—2.4

mm wide, 3.6—4.6 times longer than broad,

glabrous above and below, sometimes with

a few marginal cilia, venation of the midrib

only or sometimes | or 2 lateral veins that are

also visible on lower surface. Inflorescences

globose, some with dimorphic flowers;

peduncles 4-12 mm long at anthesis, 5—21

mim long at fruiting stage, glabrous; bracts 2,

positioned in middle one-third of peduncle,

deciduous, ovate, amplexicaul, glabrous;

lower bract 1.5—2.3 mm long; flowers 10—20

per spike; bracteole elliptical, 0.6—1 mm long,

caducous, usually falling before anthesis.

Bisexual flowers: calyx campanulate, 1—1.3

mm long, glabrous; petals 1.8—2.1 mm long,

glabrous; stamens 5, filaments 2.4—2.8 mm

long, anthers 0.5—0.6 mm long, terminal

gland absent; style 1.5—2 mm long, glabrous;

ovary glabrous or hairy. Neuter flowers: calyx

0.6—0.7 mm long, glabrous; petals 1.4—1.5

mm long, glabrous; staminodes 5, 2.5-3.5

mm long, 0.25—0.3 mm wide. Pods circular,

7—9 mm long, 7.5—8.5 mm broad, indehiscent,

glabrous or with dense patent eglandular hairs

0.05—0.2 mm long; stipe 1-2 mm long. Seeds

1 per pod, broadly elliptical in outline, 4.5—

4.8 mm long, 3.8—4.3 mm wide, brown. Figs.

4E, 7.

Additional selected specimens examined: Australia.

Western Australia. Kimberley Research Station,

Kununurra, Mar 1963, Lazarides 6729 (CANB, K,

PERTH); gravel extension of Packsaddle Road, Nov

2009, Byrne 3663 (MEL, PERTH); 44.3 km _ from

Karratha on North West Coastal Highway towards

Millstream, May 2008, Guerin GG1007 & Trudgen (AD,

PERTH); 3 km N of Nicholson Station, May 1973, Aplin

5363 (PERTH); Miles Bore area [near Derby], Feb 1992,

Mitchell 2061 (PERTH); Flora Valley Station, Nov 2000,

Craig ABC1627 (PERTH). Northern Territory. c. 20

km NW of Burta Wurta, Apr 2012, Lewis 2008 (BRI,

Ct] |

= .

: he

> *.

R Att

Vries cheese

Fig. 7. Shrubs of Neptunia monosperma (McDonald

22338, BRI). Photo: K.R. McDonald.

CANB, DNA); 7 miles [11 km] E of Stuart Highway,

Beetaloo Station Road, Feb 1969, Must 382 (CANB,

NT); K1 Bore, Brunette Downs Station, Jun 2016,

Schubert 811 (NT); 23 miles [37 km] SSW of Georgina

Downs Station, Mar 1953, Perry 3486 (BRI, CANB,

MEL, NSW); N of ‘Creswell Downs’ on the road to

‘Calvert Hills’, c. 148 km from ‘Calvert Hills’, Jun 1974,

Pullen 9270 (CANB, NSW); Daly Waters, Mar 1972,

Byrnes 2503 (CANB, DNA, NT); Bore 29, Wavehill

Station, Jun 1952, Perry 2908 (CANB, NSW [lower

specimen only|); 43 km E of Daly Waters, Carpentaria

Highway, May 1994, Albrecht 5893 (BRI, DNA, NT);

McDonnell trucking yards [Alice Springs], Jan 1975,

Mitchell 211 (CANB, DNA). Queensland. BURKE

District: 15 km along the road to Magoura Station, SW

of Normanton, Apr 1974, Pullen SSS9 (BRI, CANB);

Barkly Highway 74.2 km NW of Mt Isa (by air), Nov

2001, Fraser 363 (CANB). Cook District: 32.9 km §

from the Oasis Roadhouse on Kennedy Development

Road, Jan 2022, McDonald KRM22338 (BRI). NORTH

KENNEDY DISTRICT: 65 km ESE of Ayr, Jan 1982, Pedley

4802 (BRI, MEL). SouTH KENNEDy DiIstTRIcT: Suttor

River, [Feb—Mar 1864], Bowman 19 (MEL); 12 km

E of Avon Downs HS towards Bell Lagoon, Jan 1998,

Thompson 444 & Fox (BRI, MEL). GREGORY NORTH

District: Jessamine Creek, 8 km from Winton towards

Longreach, Jun 1999, Bean 1/5177 (BRI); Bladensburg

NP, S of Winton, near Mistake Hut Dam, Mar 1998,

Forster PIF22223 & Booth (BRI, MEL). MoRETON

District: 14 km WSW of Ipswich CBD off Coopers

Road, Willowbank, Feb 2017, Ryan CRO/ (BRI). New

South Wales. Gurley district, Feb 2016, Parsons s.n.

(NSW 991194).

Distribution and habitat: Neptunia

monosperma 1s endemic to _ northern

Australia where it 1s very widespread. The

main distribution is from Derby in Western

Australia, through the northern half of the

Northern Territory to Bowen in Queensland.

The highly disjunct specimen records from

Alice Springs, the Moreton district of Qld and

north-east N.S.W. (cited above) are presumed

to have resulted from inadvertent human-

mediated transportation. The habitat at the

Ipswich location is highly modified and right

beside a well-used road; the Alice Springs

location is a trucking yard — the specimen

label states “introduced in cattle fodder’; no

habitat data are available for the Gurley site.

Neptunia monosperma was first collected

from the Karratha area of Western Australia in

1997 — it is unclear whether these populations

are naturally occurring or not. (Map 9). It

inhabits grassland or open eucalypt woodland

on heavy clay soils.

Phenology: Flowers are recorded from

November to April; fruits from January to

June.

Affinities: Neptunia monosperma 1s an

upright plant to 50 cm high, and with strictly

l-seeded pods. Herbarium specimens of it

most resemble NV. major, as they both have

glabrous stems and leaves, leaf pinnae with

numerous pairs of leaflets, and deciduous

stipules. Neptunia monosperma differs by

the peduncles at anthesis being 4-12 mm

long (15-55 mm long for N. major); fruiting

peduncles being 5-21 mm long (25-60

mm long for N. major); 10—20 flowers per

inflorescence (25-35 flowers for N. major);

staminodes 2.5-3.5 mm long (6.5—-l11 mm

long for NV. major); and pods strictly 1-seeded

((1—)2—5-seeded for N. major).

Typification: Windler (1966) chose _ the

Victoria River syntype as lectotype for

Neptunia monosperma, although he did not

cite the herbarium 1n which it was held. Cowan

Austrobaileya 12: 59-106 (2022)

(1996) ignored this choice and selected instead

the specimen collected by Landsborough

from the Gulf of Carpentaria. Cowan argued

that Windler did not deliberately choose a

lectotype, and that he “simply repeated the

collection(s) cited in the protologue”. That 1s

not the case. In the protologue, Bentham cited

“N. Australia. Upper Victoria river, fF: Mueller;

Gulf of Carpentaria, Landsborough, Henne.

Queensland, Bowman.” while Windler (1966)

stated “Type—Upper Victoria River, F:-

Mueller”. Windler has clearly selected the

Victoria River gathering from amongst the

syntypes. As Windler did not designate which

herbarium holds the type, a second-step

lectotypification 1s needed, and that 1s effected

here. The sheet on which the lectotype is

mounted includes two other taxa, but the

barcode K 000791027 is associated only with

the lectotype, 1.e. the two branches on the left-

hand side of the sheet.

Notes: Neptunia monosperma has the shortest

peduncles of any of the Australian or Malesian

species. At anthesis, the peduncles are just

4—12 mm long, increasing to 5—21 mm at the

fruiting stage.

The specimen Perry 2908 (AD, CANB,

NSW) is a mixed gathering; the CANB

specimen 1s Neptunia monosperma, the AD

specimen is NV. scutata, the NSW specimen

comprises one branch of N. scutata and one of

N. monosperma.

*11. Neptunia oleracea Lour., Fl. Cochinch.

2: 654 (1790).

Type: Cochinchina, s.dat., J. Loureiro s.n.

(lecto [probably holo]: BM 000952356, fide

Merrill 1935: 188).

Mimosa prostrata Lam., Encycl. 1: 10 (1783),

nom. illeg. (M. natans L.f. (1782) cited in

synonymy); Neptunia prostrata (Lam.) Baill.,

Bull. Soc. Linn. Paris 1: 356 (1883), nom. illeg.

Mimosa lacustris Bonpl., Pl. Aequinoct. 1: 55,

t. 16 (1806); Desmanthus lacustris (Bonpl.)

Willd., Sp. Pl., ed.4, 4(2): 1044 (1806); Acacia

lacustris (Bonpl.) Desf., Tabl. Ecole Bot., ed.

3 (Cat. Pl. Horti Paris.) 301 (1829). Type:

[Colombia]. “Near Monpox’ [Santa Cruz de

Mompox], 77.¥.

Bean, Neptunia in Australia and Malesia

Desmanthus natans Willd., Sp. Pl. 4: 1044

(1806), nom. illeg. (Neptunia oleracea cited

in synonymy); Neptunia natans W.Theob.,

Burmah [Mason], ed. 3. 2: 541 (1883), nom.

illeg. nom. superfi.

Ilustration: Windler (1966: 402).

Shrub, perennial, aquatic. Stems prostrate

and trailing, glabrous, sessile glands absent,

frequently with inflated ellipsoidal “floats” up

to 8 cm long and 3 cm diameter and rooting

at the nodes. Stipules deciduous, broadly

ovate, 4.2—5.3 mm long, 2.8—3.5 mm broad,

chartaceous, faintly nerved, glabrous; apex

acute; base obliquely cordate. Leaves with

2 or 3 pairs of pinnae; petioles 35—47 mm

long, glabrous, nectary absent; rachis 10—28

mm long, glabrous, nectaries absent; rachis

extension linear, 1.2—-2.3 mm long; pinnae

with 10-17 pairs of leaflets, rachilla extension

0.8—0.9 mm long; stipels absent. Largest

leaflets 6.2-9.3 mm long, 1.8—2.5 mm wide,

3.4—4.4 times longer than broad, glabrous,

venation of 1-3 main veins, more obvious

on lower surface. Inflorescences ellipsoidal,

all with dimorphic flowers; peduncles 130-

175 mm long at anthesis, glabrous; bracts 2,

positioned on proximal one-third of peduncle,

caducous before anthesis, ovate, amplexicaul,

glabrous, lower one c. 3 mm long; flowers 40—

60 per inflorescence; bracteoles oblanceolate,

c. 2.2 mm long, caducous, usually falling

before anthesis. Bisexual flowers: calyx

campanulate, 2—2.5 mm long, glabrous;

petals 3.3-3.6 mm long, glabrous; stamens

10, filaments 6.5—7.5 mm long, anthers 0.8—1

mm long, terminal gland absent; style c. 6

mm long, glabrous; ovary glabrous. Neuter

flowers: calyx 1—1.3 mm long; petals 1.8—2.6

mim long, glabrous; staminodes 10, 6.5—9 mm

long, 0.7—0.9 mm wide. Pods and seeds not

seen.

Additional specimens examined: Philippines.

MINDANAO: Bunauan, in 1913, Taylor s.n. (US

02209018). Indonesia. JAvA: Sentiong, Aug 1904,

Backer s.n. (L 2042612); Batavia, s.dat., Blume s.n. (L

2042604); ibid, Aug 1907, Harreveld s.n. (L 2042613);

Rawa Bening bij Ambarawa, Jul 1940, Hoed & van der

Meer 600 (L 2042610); Rawah Gesian, Feb 1914, Backer

11723 (L 2042600); Rawah Bening, Jun 1927, Coert

440A (L 2042601).

8/7

Distribution and habitat: Neptunia oleracea

is distributed in tropical Asia, Africa, central

America and South America (Windler 1966).

From Malesia, I have seen specimens only

from Java and Philippines (Map 3), but ILDIS

(2022) records it also from Sumatra, Borneo,

Moluccas and Sulawesi. It inhabits warm,

slow-moving and often stagnant waters.

Phenology: It flowers year-round in the

tropics (Anon. 2022).

Affinities: Very closely allied to Neptunia

plena (see Notes under that species).

Notes: None of the available specimens of

Neptunia oleracea bears any fruits, and only

a few of them have flowers or flower buds.

Neptunia oleracea is anutritious vegetable,

containing Vitamins A, B,, B,, B, (niacin) and

C. In Thailand, the plant is cultivated for its

young shoots, which are eaten raw or as a

fried vegetable (Paisooksantivatana 1993).

12. Neptunia paucijuga A.R.Bean sp. nov.

With affinity to Neptunia dimorphantha, but

differing by the glabrous fruits, the leaves

with two pairs of pinnae (rarely one paizr),

and the nectary positioned away from the

distal end of the petiole. Typus: Australia.

Northern Territory. Nitmiluk National Park,

21 March 2000, K. Brennan 494] (holo: BRI

[AQ1024487 comprising | sheet]; iso: DNA

D0169730).

Shrub, perennial, terrestrial. Stems prostrate,

glabrous, sessile glands rare or absent. Stipules

persistent, ovate, 5.3-6.6 mm long, 1.2—1.9

mm broad, coriaceous, strongly nerved,

glabrous; apex attenuate; base obliquely

cordate. Leaves with (1 or)2 pairs of pinnae;

petioles 5-10 mm long, glabrous, with | or 2

nectaries on the lower half or about halfway

along or in upper half; rachis 7—9 mm long,

glabrous or with a few hairs, nectary absent;

rachis extension linear, 1.8—2.8 mm _ long;

pinnae with 10-18 pairs of leaflets, rachilla

extension 1.4—1.8 mm long; stipels slender,

0.4-1 mm long, divided, gland-tipped.

Largest leaflets 61-9 mm long, 1.3-2.4

mm wide, 3.7—4.7 times longer than broad,

glabrous or with a few marginal cilia, a single

vein visible on lower surface. Inflorescences

globose, all bisexual; peduncles 40-54 mm

long at anthesis, 44-65 mm long at fruiting

stage, glabrous; bracts 2, positioned on

proximal one-third of peduncle, persistent,

ovate, amplexicaul, lower one larger, 2.2—

6.6 mm long, glabrous; flowers 16—28 per

inflorescence; bracteoles oblanceolate, c. 1.3

mm long, caducous, usually falling before

anthesis. Bisexual flowers: calyx campanulate,

1.2-1.3 mm long, glabrous; petals 2—2.2 mm

long, glabrous; stamens 5, filaments c. 2.2 mm

long, anthers 0.9—1 mm long, terminal gland

prominent; style 3.5—3.8 mm long, glabrous;

ovary glabrous. Neuter flowers: absent. Pods

broadly elliptical, 1-—3-seeded, 11-19 mm

long, 9-13 mm broad, indehiscent, stipe

1—2 mm long, apex (excluding style) obtuse;

glabrous. Seeds broadly elliptical in outline,

c. 3.9 mm long and 3.2 mm wide, brown. Fig.

2C.

Additional specimens examined: Australia. Northern

Territory. Edith River area, Apr 1999, Michell & Risler

2338 (DNA); Fire plot 45, Nitmiluk NP, Mar 2000,

Brennan 4958 (DNA); 17 Mile Valley, Nitmiluk NP, Mar

2002, Michell & Kerrigan 3954 (DNA); Old Edith Falls

track, Mar 1987, Bowman 471 (DNA); Mt Todd mine

site, Mar 1995, Egan 4443 (DNA); Warlock [| Warloch|

Ponds, Apr 1968, Byrnes NB630 (AD, DNA); Bradshaw

Military Training Area, c. 60 km NW of Timber Creek,

May 2017, Cuff 662 (DNA).

Distribution and habitat: Neptunia paucijuga

is endemic to Australia in the Northern

Territory. It is known from the Katherine,

Timber Creek and Mataranka areas (Map

7). It inhabits rocky sandstone hills and

rises in grassy open woodland dominated

by eucalypts, except for the Warloch Ponds

location, which is a flat site with black

cracking clay.

Phenology: Flowers and fruits are recorded

from March and April.

Affinities: Neptunia paucijuga 1s related to

the widespread NV. dimorphantha, with which

it shares the 1—3-seeded pods, the relatively

long peduncle (42—90 mm long) bearing two

bracts on the proximal one-third, and the

presence of a nectary (or nectaries) on the

petiole. In N. paucijuga, the position of the

petiole nectary is variable, and sometimes

Austrobaileya 12: 59-106 (2022)

there are two adjacent nectaries. In N.

dimorphantha, the nectary is consistently

located at the distal end of the petiole

(between the lowest pair of pinnae), and there

is never a second nectary. Neptunia paucijuga

has leaves with 2 pairs (rarely 1 pair) of

pinnae, while N. dimorphantha has 3-5 pairs

of pinnae. The stems and leaflets of Neptunia

paucijuga are glabrous or almost glabrous

although the rachilla bears some long hairs; N.

dimorphantha stems and leaves are sparsely to

moderately hairy. An important characteristic

of N. dimorphantha is the presence of short

hairs along the margins of the pods. These

hairs are absent in N. paucijuga and its pods

are glabrous.

Note: The few flowering specimens of

Neptunia paucijuga bear only bisexual

flowers, and it is possible that neuter flowers

do not occur in this species.

Etymology: From the Latin paucus meaning

‘few’, and -jugus, meaning ‘paired’. This

refers to the fewer pairs of pinnae in this

species when compared to the related

Neptunia dimorphantha.

*13. Neptunia plena (L.) Benth., J Bot.

(Hooker) 4: 355 (1841).

Mimosa plena L., Sp. Pl. 1: 519 (1753);

Desmanthus plenus (L.) Willd., Sp. PL, ed.

4, 4(2): 1045 (1806); Hemidesmas plenus (L.)

Raf., Sylva Tellur. 119 (1838). Type citation:

“Habitat in Vera Cruce”. Type: Herb. Linn.

No. 1228.12 (lecto: LINN, fide Kostermans

1980: 462).

Mimosa punctata L., Syst. Nat. ed. 10, 2: 1311

(1759); Desmanthus punctatus (L.) Willd., Sp.

Pl., ed. 4, 4(2): 1047 (1806); Acacia punctata

(L.) Desf., Tabl. Ecole Bot., ed. 3 (Cat. PI.

Horti Paris.) 300 (1829). Type: “America”,

s.dat., P. Browne s.n., Herb. Linn. No. 1228.14

(lecto: LINN, fide Wiynands 1983: 152).

Desmanthus polyphyllus DC., Prodr. 2: 444

(1825); Neptunia polyphylla (DC.) Benth., J.

Bot. (Hooker) 2: 129 (1840). Type: locality

unknown, herb. Moricand ex hort. Napol.

(holo: G?, 72.v.).

Bean, Neptunia in Australia and Malesia

Mimosa adenanthera Roxb., Fl. Ind. 2: 554

(1832). Type: “East India’, s.dat., W. Roxburgh

s.n. (syn: K 000791019).

Illustration: Windler (1966: 399).

Shrub, perennial, aquatic. Stems sprawling

or erect, to | m high, glabrous, sessile glands

absent frequently with inflated ellipsoidal

“floats” up to 8 cm long and 3 cm diameter

and rooting at the nodes. Stipules deciduous,

broadly ovate, 9-12 mm long, 3.5—6 mm broad,

chartaceous, faintly nerved, glabrous; apex

acute; base obliquely cordate. Leaves with 2

or 3(—4) pairs of pinnae; petioles 35-55 mm

long, glabrous, nectary usually present, small,

near distal end, sometimes absent; rachis

20—48 mm long, glabrous, nectaries absent;

rachis extension linear, 3—6 mm long; pinnae

with 16—24 pairs of leaflets, rachilla extension

1.5—2 mm long; stipels absent. Largest leaflets

11-18 mm long, 2.2-3.3 mm wide, 4.7—5.8

times longer than broad, glabrous, venation

of 1-3 main veins, more obvious on lower

surface. Inflorescences ellipsoidal, all with

dimorphic flowers; peduncles 75-125 mm

long at anthesis, 80-140 mm long at fruiting

stage, glabrous; bracts 2, positioned on

proximal or middle one-third of peduncle,

caducous before anthesis, ovate, amplexicaul,

glabrous, lower one 5.5—8.2 mm long; flowers

30—50 per inflorescence; bracteoles elliptical,

1.2-1.5 mm long, caducous, usually falling

before anthesis. Bisexual flowers: calyx

campanulate, 2—2.4 mm long, glabrous;

petals 3.3-3.4 mm long, glabrous; stamens

10, filaments 6—6.5 mm long, anthers 0.9-1.2

mim long, terminal gland absent; style 4.3—4.7

mm long, glabrous; ovary glabrous. Neuter

flowers: calyx 0.8—1 mm long; petals 1.9—2.3

mm long, glabrous; staminodes 10, 8-13

mm long, 0.5—1.2 mm wide. Pods oblong,

12—15-seeded, 31-41 mm long, 8.5-10 mm

broad, dehiscent along one margin only, apex

obtuse, stipe 6—10 mm long; glabrous. Mature

seeds not seen. Figs. 3J, 8.

Additional selected specimens examined: Indonesia.

JAVA: S.loc., s.dat., Zollinger 3160 (MEL); Buitenzorg,

Jan 1950, Ooststroom 1264] (L); Bogor, Feb 1950,

Ooststroom 12732 (CANB). Papua New Guinea.

WESTERN (FLy): Old Mawatta, South Fly district, Feb

2018, Waterhouse BMW8301 (BRI, CANB, CNS).

Australia. Northern Territory. 80 Cunjevoi Road,

Virginia, Nov 2011, Mitchell 7041] (BRI, DNA);

Nhulunbuy town lagoon, NE Arnhem Land, Mar 2012,

Westaway JOW3786 (BRI, CANB, DNA). Queensland.

Cook District: Lake Placid, Cairns, May 2009, Vitelli

WMI19S (BRI). NORTH KENNEDY District: 15 km S of

Home Hill, May 2013, O’Bryan s.n. (BRI [AQ796758],

MEL). SOUTH KENNEDY DISTRICT: 6 Staceys Road,

Kuttabul, Jun 2020, Gunasekera s.n. (BRI [AQ952483]).

MoRETON' DISTRICT: 266 Ritchie Road, Pallara,

Brisbane, Mar 2006, Johnson s.n. (BRI [AQ751366]); c.

14 km W of Peak Crossing, Feb 2006, Wickman s.n. (BRI

[AQ751351)).

Distribution and habitat: Neptunia plena 1s

known from North and South America, almost

exclusively between the Tropic of Cancer and

the Tropic of Capricorn, including islands

of the Caribbean Sea (Windler 1966). In the

Old World, it is known from India, China,

Malaysia, Singapore, Java, Sri Lanka (ILDIS

2022), Papua New Guinea and Australia. It

is a recent introduction to Australia, with the

first non-cultivated record in 2001 (Map 10).

It inhabits dams, lagoons and swampy areas

where water movement is minimal.

Phenology: Flowers and fruits can be found

throughout the year.

Notes: Both Neptunia plena and N. oleracea

have been recorded as naturalised species for

Queensland (Brown 2020). These species have

always been poorly distinguished. Bentham

(1875) and Schery (1950) used the presence

or absence of spongy floating stem tissue as

the major determining character between

N. plena and N. prostrata (Lam.) Baill. (=

N. oleracea), and Schery (1950) introduced,

as a secondary distinguishing character, the

presence (N. plena) or absence (N. oleracea)

of a nectary gland on the leaf (at the apex

of the petiole near the basal pair of pinnae).

Windler (1966) gave increased importance to

the nectary character and did not mention the

spongy stem tissue in his key or discussion.

Other characters Windler (1966) used in his

key to distinguish them are 1. the number

of leaflets per pinna (but in his descriptions

there is considerable overlap (9-38 pairs for

N. plena; 8-20 pairs for N. oleracea); and

2. the number of seeds per pod (8—20 for N.

plena, 4-8 tor N. oleracea). These are of

little practical value as none of the south-east

Asian specimens at L determined by Windler

as N. oleracea (or its synonym WN. prostrata)

Austrobaileya 12: 59-106 (2022)

Fig. 8. Neptunia plena, flowering and fruiting (Waterhouse BMW8301/, BRI). Photo: B. Waterhouse.

bears any fruits. Subsequent authors (e.g.

Kostermans 1980; Nielsen 1992; Santos-Silva

et al. 2020) have not provided an alternative

key to these species — all have copied or

adapted Windler’s key.

Using Windler’s key, most Queensland

specimens can be assigned to Neptunia plena

because of the presence of the small nectary

on the petiole. However, in some specimens,

the nectary is developed on just one or two

leaves and is lacking on the remaining leaves.

There are also a few specimens where the

nectary is lacking altogether and these have

previously been identified as N. oleracea, but

their morphology in every other way matches

the NV. plena specimens and hence I regard

them as N. plena also.

Herbarium specimens of Neptunia

oleracea from Malesia do often appear

morphologically distinct from the N. plena

from Australia (fewer leaflets per pinna;

more abundant spongy stem tissue), but this

could be due to the growth stage that has

been selected, with N. oleracea specimens

representing younger plants growing across

the water (leaves in juvenile phase and

plants too young to bear fruits), and N. plena

representing older growth with ascending

mostly non-spongy stems, and often bearing

fruits.

Schery (1950) commented “Probably N.

prostrata |= N. oleracea| 1s not a valid species,

but merely a form of NV. plena growing floating

in water and assuming a prostrate position.”

Peck & Serviss (2011) accepted NV. oleracea

but said that “Neptunia oleracea 1s extremely

similar morphologically to NV. plena ...”.

It does seem doubtful that Neptunia

oleracea 1s taxonomically distinct from WN.

plena, but that determination would require

a more wide-ranging study than is possible

here.

14. Neptunia proxima A.R.Bean sp. nov.

With affinity to Neptunia gracilis but differing

by the long spreading hairs on_ leaves,

peduncles and pods; the lack of staminodes;

and the shorter peduncles. Typus: Australia.

Queensland. Cook District: Maytown, near

historical Jimmy Ah Fun stores site in Leslie

Street, 13 April 2016, PJ. Forster PIF 44127

& K.R. McDonald (holo: BRI [AQ940153

comprising | sheet}).

Bean, Neptunia in Australia and Malesia

Shrub, perennial, terrestrial. Stems prostrate,

with patent hispid hairs 0.2—0.5 mm long,

sessile glands absent. Stipules persistent,

ovate to broadly ovate, 2.8—-6.2 mm long,

22.3 mm _ broad, coriaceous, strongly

nerved, with hispid hairs along margin; apex

attenuate; base obliquely cordate. Leaves

with 2 or 3 pairs of pinnae; petioles 5—8 mm

long, hispid, nectary absent; rachis 6-15 mm

long, with patent hispid hairs 0.2—0.4 mm

long, nectary absent; rachis extension linear,

2.5—3.1 mm long; pinnae with 16—21 pairs of

leaflets, rachilla extension 1.2—1.6 mm long;

stipels slender, 0.1—0.4 mm long, divided,

gland-tipped. Largest leaflets 3—4.4 mm long,

0.9-1.1 mm wide, 3.3—4.4 times longer than

broad, surfaces glabrous but margins ciliate,

venation of 1-3 main veins, more obvious

on lower surface. Inflorescences globose, all

flowers bisexual; peduncles 17-24 mm long

at anthesis, 30—45 mm long at fruiting stage,

hispid hairy throughout; bracts absent or 1,

positioned on proximal one-third of peduncle,

persistent, broadly ovate, amplexicaul,

2.5-3.3 mm long, margin ciliate, surface

glabrous or sparsely hairy; flowers 12—25 per

inflorescence; bracteoles oblanceolate, 1—1.3

mm long, persistent until anthesis. Bisexual

flowers: calyx obconical, 1.3—1.5 mm long,

glabrous; petals 2—2.6 mm long, glabrous;

stamens 5, filaments 2.5—3 mm long, anthers

0.6—0.8 mm long, terminal gland absent;

style 2.1-3.4 mm long, glabrous; ovary

hairy. Neuter flowers absent. Pods oblong,

(2—)3—6-seeded, 15-27 mm long, 8-9 mm

broad, tardily dehiscent along both margins,

stipe 1.5—2 mm long, apex obtuse; with dense

eglandular patent hairs 0.2—0.4 mm long.

Seeds broadly elliptical in outline, 3.8—4 mm

long, 3.2—3.3 mm wide, brown. Fig. 3E—-F.

Additional = specimens examined: Australia.

Queensland. Cook DIstTRIcT: Cabbage Tree Creek, c.

1 mile [1.6 km] E of the Kennedy River, Laura — Coen

Road, Oct 1962, Smith 11691 (BRI); Kings Plains

Station, 10.3 km old coach road from homestead, Apr

2015, McDonald 17070 & Thompson (BRI); Littleton NP,

1.3 km by road N of ranger station, Mar 2016, McDonald

KRMI8/18S (BRI); Rockfields, Gilbert River, May 1954,

Everist 5473 (BRI).

Distribution and habitat: Neptunia proxima

is endemic to Queensland, Australia. It

extends from near Laura to Gilbert River, east

of Croydon (Map 7). It grows in hilly terrain

in eucalypt woodland with sandy-loam soil,

or in Melaleuca woodland on plains with

clayey soil.

Phenology: Flowers are recorded for March,

April and May; fruits are recorded for March,

April and October.

Affinities: Neptunia proxima difters from the

similar NV. gracilis by the petioles 5-8 mm

long (10-17 mm long for N. gracilis); 16-21

leaflet pairs per pinna (12-17 pairs for N.

gracilis); largest leaflets 3—4.4 mm long (5.4—

8.2 mm for N. gracilis); inflorescences all

bisexual (inflorescence dimorphic, with some

neuter flowers for N. gracilis); peduncles (at

anthesis) 17-24 mm long ((25—)40—90 mm

long for NV. gracilis) and peduncles (at fruiting

stage) 30—45 mm long (50—105 mm long for

N. gracilis).

Etymology: The epithet is from the Latin

proximus, meaning ‘next’ or ‘nearest’. This 1s

in reference to its relationship with Neptunia

gracilis.

15. Neptunia scutata A.R.Bean sp. nov.

With affinity to Neptunia xanthonema but

differing by the lack of nectaries on the leaf

rachis, the peduncles hairy throughout, the

smaller leaflets and the smaller bracts. Typus:

Australia. Northern Territory. Gregory

National Park, Bullita section, 14 February

1992, I. Cowie 2376 & P.S. Brocklehurst

(holo: CANB 00471857 comprising 1 sheet;

iso: DNA D0066573, MEL 0717866A).

[Neptunia dimorphantha auct. non Domin,

pro parte: Whibley (1981: 114); Wheeler

(1992: 341)].

Illustration: Cowan (1998: 22, fig. 36 J, K), as

N. monosperma

Shrub, perennial, terrestrial. Stems prostrate

or sprawling, glabrous, sparsely hairy or

densely hairy, glandular or not, sessile glands

rare or absent. Stipules persistent, broadly

ovate, 3.5—6.8 mm long, 1.2—2.9 mm broad,

coriaceous, usually with 1 strong nerve and

several faint nerves, apex attenuate, base

obliquely cordate, glabrous or with sparse

hairs. Leaves with 3-5, rarely 6 pairs of

pinnae; petioles 4-13 mm long, hairy, with

a prominent nectary about halfway along, or

in the upper half, occasionally absent; stipels

slender, 0.2—0.8 mm long, divided, gland-

tipped; rachis 7—28 mm long, with sparse to

dense simple hairs, nectaries absent; rachis

extension linear, 1—3.8 mm long, glabrous

or sparsely hairy; pinnae with 11—24 pairs of

leaflets, rachilla extension 0.8—1.5 mm long.

Largest leaflets 3—7 mm long, 1—1.8 mm wide,

2.8—4.3 times longer than broad, glabrous

above, glabrous or sparsely hairy below,

and with marginal cilia, venation of 1-3

main veins, more obvious on lower surface.

Inflorescences globose to broadly ellipsoidal,

some with dimorphic flowers; peduncles

8—35 mm long at anthesis, 18-35 mm long

at fruiting stage, with patent glandular hairs

throughout, 0.1-0.3 mm long; bracts 2,

positioned on proximal one-third of peduncle,

persistent, ovate, amplexicaul, the lower one

2.5—8 mm long, glabrous above, glabrous or

hairy below; flowers 25—50 per inflorescence;

bracteole oblanceolate, 0.9-1.2 mm _ long,

caducous, usually falling before anthesis.

Bisexual flowers: calyx campanulate, 0.8—1.6

mim long, glabrous or with sparse hairs; petals

1.4—2.4 mm long, glabrous or with sparse

hairs; stamens 5, filaments 2.1—3.2 mm long,

anthers 0.5—0.6 mm long, terminal gland

absent; style 1.3—-3 mm long, glabrous; ovary

hairy. Neuter flowers: calyx 0.5—0.8 mm long,

glabrous; petals 1.1-2.4 mm long, glabrous;

staminodes 5, 4—7.5 mm long, 0.3—0.6 mm

wide. Pods broadly elliptical to circular,

l-seeded, 7.5—9.5 mm long, 5—8.5 mm broad,

indehiscent, with patent glandular hairs 0.2—

0.5 mm long; stipe 1—2.5 mm long. Seeds

broadly elliptical in outline, 3.8—4.5 mm long,

2.8—3.4 mm wide, brown. Fig. 9.

Additional specimens examined: Australia. Western

Australia. Long Spring, 66 km NNE of Carlton Hill HS,

c. 10 km NE of Knob Peak in the Ningbing Range, Mar

1978, Lazarides 84851 (BRI, CANB); Langey Crossing on

west bank of Fitzroy River, on Jarrananga Plain in Yeeda

Station, Mar 1986, Foulkes 42] (CANB, PERTH); Great

Northern Highway, 84 km N from Halls Creek, May

1999, Purdie 4566 (CANB, PERTH); Revolver Creek,

upper slopes of southern Carr Boyd Ranges, Mar 1978,

Hartley 14516 (CANB); Kimberley Research Station,

Austrobaileya 12: 59-106 (2022)

Kununurra, Apr 1964, Richards 75 (CANB). Northern

Territory. Darwin wharf area, adjacent to duck pond,

Feb 2007, Roberts 137 (BRI, CANB); c. 27 km SW

of ‘Calvert Hills’ on the road to ‘Cresswell Downs’,

May 1974, Pullen 9254 (CANB); | mile [1.6 km] S of

Mt Sanford Outstation, Jun 1949, Perry 2183 (CANB);

Morphett Creek, Jan 1994, Egan 2995 (CANB, DNA).

Queensland. Cook District: 33 km along ‘Blackdown’

Road, from Chillagoe —*Wrotham Park’ Road, Jan 1993,

Bean 5637 & Forster (BRI, MEL). BURKE DISTRICT: 20

miles [32 km] NNE of Camooweal township, May 1948,

Perry 983 (BRI, CANB); | km along road to Herbert

Vale HS, off Lawn Hill to Riversleigh Road, Apr 1997,

Forster PIF20867 & Holland (BRI, DNA, MEL). NorTtH

KENNEDY District: 20 km from Charters Towers,

towards Clermont, Apr 2002, Bean 18966 (BRI); 7 km

from Homestead towards Pentland, Feb 1994, Bean

7485 & Forster (BRI). SOUTH KENNEDy DISTRICT: 3 km

W of Yarrowmere HS, Mar 1993, Thompson BUC987

& Henderson (BRI). GREGORY NORTH DISTRICT:

Diamantina NP, northern boundary of park, Apr 1997,

Forster PIF20754 & Holland (BRI, DNA); 20 miles

[32 km] W of Winton, May 1966, Pedley 1996 (BRI).

MITCHELL District: Stonehenge, on road to airstrip,

May 2004, Bean 22446 (BRI).

Distribution and habitat: Neptunia scutata

is widespread in northern Australia, from

Karratha in Western Australia to Charters

Towers in Queensland (Map 11). It inhabits

grassland often dominated by Mitchell grass

(Astrebla spp.) growing in brown to black clay

soul, and sometimes occurs in Melaleuca or

eucalypt woodland with sandy to clayey soils.

Phenology: Most flowering and _ fruiting

records are between December and June, with

occasional records in the other months.

Affinities: Neptunia xanthonema difters

from N. scutata by the frequent presence of

nectaries on the leaf rachis, one adjacent to

each pair of pinnae (nectaries absent from

rachis in JN. scutata); peduncles usually

glabrous except at the apex (hairy throughout

for N. scutata); seeds 2.7-3.6 x 2.4—2.8 mm

(3.8—4.5 x 2.8-3.4 mm for N. scutata); and

the mostly larger leaflets, 5.8-11 mm long

(3—7 mm long for N. scutata). In addition, N.

xanthonema has 2 or 3(—4) pairs of pinnae

(3—5(—6) pairs for N. scutata).

Note: The petiolar nectary that is a feature

of this species, is occasionally absent for all

petioles on a herbarium sheet.

Bean, Neptunia in Australia and Malesia

Fig. 9. Neptunia scutata, showing leaf, inflorescence,

peduncle, bracts and young fruits (Bean 18966, BRI).

Etymology: From the Latin scutatus, meaning

‘shield-shaped’. This is given in reference

to the flattened broadly elliptical fruits that

resemble a small shield.

16. Neptunia tactilis A.R.Bean sp. nov.

With affinity to Neptunia javanica but differing

by the lack of a nectary on the petiole, the lack

of staminodes, and the 15—28 pairs of leaflets

per pinna. Typus: Australia. Queensland.

Cook District: 4.6 km S of Batavia Downs

on the Peninsula Development Road, 19 April

1990, JR. Clarkson 8248 & V.J. Neldner

(holo: BRI [AQ512120 comprising 1 sheet];

iso: DNA D0061355, L 2042652).

Neptunia ‘Darwin’, pro parte: (Dunlop et al.

1993227),

Ilustration: Dunlop et al. (1995: 26).

Shrub, perennial, terrestrial, prostrate to

trailing. Stems glabrous, sessile glands absent.

Stipules persistent, narrowly ovate, 4.5—6.5

mm long, 1.1-1.8 mm broad, coriaceous,

with several strong nerves, apex attenuate,

filamentous, base obliquely cordate or obtuse,

glabrous or with a few hairs along margin.

Leaves with I-3 pairs of pinnae; petioles 3—9

mm long, glabrous or sparsely hairy, nectary

absent; rachis absent or 6-19 mm long, with

sparse simple hairs, nectaries absent; rachis

extension linear, 2.2—6 mm long; pinnae with

20-28 pairs of leaflets, rachilla extension

1—3.3 mm long; stipels slender, 0.1—0.9 mm

long, entire, sometimes gland-tipped. Largest

leaflets 4.5—6.7 mm long, 1.1—-1.5 mm wide,

3.6-5.2 times longer than broad, glabrous

above, glabrous or sparsely hairy below, and

with marginal cilia, midvein and a few lateral

veins visible on lower surface. Inflorescences

globose, all with bisexual flowers only;

peduncles 14-36 mm long at anthesis, 20—59

mm long at fruiting stage, glabrous or with

sparse eglandular patent hairs throughout;

bracts absent; flowers 6-12 per spike;

bracteoles oblanceolate, 0.8—1.9 mm _ long,

caducous or somewhat persistent. Bisexual

flowers: calyx campanulate to obconical,

1—2.2 mm long, glabrous; petals 1.9—2.8 mm

long, glabrous; stamens 5-8, filaments 2.5—4

mm long, anthers 0.6—1 mm long, terminal

gland very small or conspicuous; style 2.2—4

mm long, glabrous; ovary glabrous. Neuter

flowers absent. Pods oblong or elliptical,

16—42 mm long, 7.5—11 mm broad, dehiscent

along one margin only; stipe 2—6 mm long;

apex acute or obtuse; surface of pod glabrous.

Seeds 6-9 per pod, broadly elliptical in

outline, 3.8—4.5 mm long, 3.5—3.6 mm wide,

brown. Fig. 3G.

Additional selected specimens examined: Australia.

Western Australia. Boongaree Island, Prince Frederick

Harbour, Jul 1973, Wilson 11320 (PERTH); 2 km W

of camp, Mitchell Plateau, Apr 1982, Keighery 4998

(PERTH); Dunvert Island, Buccaneer Archipelago, NE

of Derby, Apr 1997, Martin CB132 (PERTH); Adjacent

to Mermaid Tree, behind beach at Careening Bay, North

Kimberley, May 1998, Mitchell 5410 (AD, PERTH);

Augustus Island, May 2008, Handasyde TH3541

(PERTH). Northern Territory. Elcho Island, Jul 1975,

Maconochie 2168 (CANB, DNA); Fossil Head, Feb

1994, Leach 4222 (DNA); 15 km WSW of Cape Arnhem,

Feb 1994, Brennan 2440 (DNA); Bickerton Island,

Milyakburra Community, Apr 1993, Leach 3450 &

Cowie (DNA); Arnhem Bay, mouth of Cato River, May

1992, Cowie 2882 (DNA); Copeland Island, Mountnorris

Bay, Apr 1883, Foelsche 82 (AD, MEL); Road to Matt

Wilson Lookout, Gregory NP, Jan 1998, Harwood 277

(DNA); Gunn Point, Mar 1993, Egan 617 (DNA); North

Goulburn Isle, May 1992, Dunlop 9042 (DNA); SE end of

Fisher airstrip, Gimbat, Nov 1991, Brennan 1593 (DNA);

Groote Eylandt, c. 25 km E of Angurugu, Mar 2005,

Risler & Brennan 2946 (DNA); Near Glenluckie Creek,

Stuart Highway, Mar 1991, Cowie 1566 & Munns (DNA,

MEL); on Katherine road, 10.2 km from Gove airport,

Nov 1989, Forster PIF6012 (BRI); Groote Eylandt, GE

115, Mar 2005, Brennan 6421 (DNA). Queensland.

Cook District: Beside the road to Pormpuraaw, Apr

2009, Wannan 5603 (BRI); 6.8 km N of the new road

turnoff to Weipa, on the Peninsula Development Road,

Jul 1985, Clarkson 6043 (BRI, CANB, MEL, NSW);

NW Cape York Peninsula, Skardon River landing

area, Mar 1995, Gunness 2478 (BRI); vicinity of Weipa

township, Jun 1997, Grimshaw JFG697E & Gollan

(BRI); South Cape Bedford, May 1990, Clarkson 8652 &

Neldner (BRI); East of Weipa on Peninsula Development

Road, Feb 2000, Wannan 1547 & Jago (BRI); S of

Strathburn boundary on Strathburn access road, Feb

2015, Thompson SLTI5006 (BRI); Alkoomie Station,

W of Cooktown, Mar 2015, McDonald KRM16695

(BRI); Brooklyn Nature Reserve, Mt Spurgeon Road,

19 February 2009, McDonald KRM8229 & Puschendorf

(BRI, MEL). BuRKE District: North Bountiful Island,

South Wellesley Group, Gulf of Carpentaria, Nov 2002,

Pedley BOI23 & Thomas (BRI).

Distribution and habitat: Neptunia tactilis

is endemic to Australia where it 1s found

in northern parts of the Kimberley region

of Western Australia, the “Top End’ of the

Northern Territory, and Cape York peninsula

in Queensland (Map 6). It grows in eucalypt

woodland on stony lateritic plains and slopes.

Phenology: Flowers are recorded from

November to April, although there is one

additional record from September. Fruits are

recorded from November to May, and one

record in September. The September record

was associated with regrowth after a fire.

Affinities: Neptunia tactilis 1s a very sparsely

hairy species with few pinnae, numerous

leaflets per pinna, short petioles, no bracts

on the peduncle, and bisexual inflorescences.

It is similar to N. javanica but differs by

the lack of a nectary on the petiole (present

in N. javanica), the 5—8 stamens (10 for N.

javanica), the 15-28 pairs of leaflets per

pinna (9-15 pairs for N. javanica), and the

lack of staminodes (present in NV. javanica).

N. tactilis difters from N. gracilis by the 5—8

stamens (always 5 for N. gracilis), the 15—28

pairs of leaflets per pinna (12-17 pairs for N.

gracilis), the petioles only 3—9 mm long (10-—

Austrobaileya 12: 59-106 (2022)

17 mm long for N. gracilis), the consistent

lack of bracts on the peduncle (1 bract present

on some peduncles for N. gracilis), and the

6—12 flowers/inflorescence (12—25 flowers for

N. gracilis).

Note: A specimen from 4 miles S of Katherine

U.B. Wilson 51, CANB, DNA) resembles

Neptunia tactilis, but has dense hispid hairs

on the stems and leaves. It may represent a

distinct taxon.

Etymology: The species epithet is from the

Latin tactilis, meaning ‘sensitive to touch’.

This refers to the leaflets which close in on

each other when touched, a feature of this and

other species of Neptunia.

17. Neptunia valida A.R.Bean sp. nov.

With affinity to Neptunia scutata but differing

by the upright habit, the lack of a nectary

on the petiole, and the frequent presence of

three or four bracts on the peduncle. Typus:

Australia. Western Australia. Near Wyndham

port hotel, 17 April 2000, A.A. Mitchell

6148 (holo: BRI [AQ642949]; iso: PERTH

6195342).

[Neptunia dimorphantha auct. non Domin,

pro parte: Wheeler (1992: 341)|.

Shrub, perennial, terrestrial. Stems erect,

1—1.5 m high, densely glandular-hairy, sessile

glands absent. Stipules persistent, broadly

ovate, 4.9-8.3 mm long, 2.2—4.1 mm broad,

coriaceous, strongly nerved, hairy; apex

acute to attenuate; base obliquely cordate.

Leaves with (2 or)3 pairs of pinnae; petioles

6—10 mm long, hairy, nectary absent; rachis

8—15 mm long, with dense glandular hairs,

nectary absent; rachis extension linear,

2.7/-2.9 mm long; pinnae with 9-15 pairs of

leaflets, rachilla extension 0.5—1.5 mm long;

stipels slender, 0.2—0.5 mm long, divided,

gland-tipped. Largest leaflets 4—7.4 mm long,

1.4—2.3 mm wide, 2.8—3.5 times longer than

broad, glabrous on upper surface, sparsely to

dense hairy on margins and lower surface,

venation of midvein only, visible only on

lower surface. Inflorescences globose, some

with dimorphic flowers; peduncles 14—27 mm

long at anthesis, 27-38 mm long at fruiting

stage, with glandular hairs throughout, 0.1—

Bean, Neptunia in Australia and Malesia

0.3 mm long; bracts 2—4, usually 3, positioned

on proximal or middle one-third of peduncle,

persistent, broadly ovate, amplexicaul, lower

one 5.4—7.5 mm long, glabrous above, hairy

below; flowers 25-40 per inflorescence;

bracteoles oblanceolate, 1.5—1.8 mm _ long,

persisting until anthesis. Bisexual flowers:

calyx obconical, 1—1.2 mm long, hairy on lobe

margins; petals 2.2—-2.5 mm long, sparsely

hairy on outer surface; stamens 5, filaments

3.4-3.6 mm long, anthers 0.7—0.9 mm long,

terminal gland present; style 3.5—4.5 mm

long, hairy on lower half; ovary hairy. Neuter

flowers: calyx 0.9-1 mm long; petals 1.8—2

mm long, sparsely hairy on outer surface;

staminodes 5, 8.5-9.5 mm long, 0.5—0.9

mm wide. Pods circular to broadly elliptical,

l-seeded, 7.5-9.5 mm long, 5.5—7.5 mm

broad, indehiscent, stipe c. 0.5 mm long,

apex obtuse; surface of pod with dense patent

hairs 0.1—0.25 mm long, at least some with

glandular apex. Seeds broadly elliptical in

outline, 4.3—4.6 mm long, 3.5—3.8 mm wide,

brown. Fig. 4C.

Additional specimens examined: Australia. Western

Australia. Cambridge Gulf, in 1887, Wright s.n. (MEL

596323); Gorge north of Devil Devil Spring, Smoke

Creek, SW of Lake Argyle, May 1980, Weston 12344

(PERTH); Shell garage grounds, Wyndham, Mar 1992,

Mitchell 214] (PERTH); along Victoria Highway, 15 km

S of highway, 40 km E of Kununurra, Apr 1989, Halford

H50 (BRI, PERTH); Wyndham Port, East Kimberley,

Jun 2008, Roberts APRIO79 (CANB, DNA, PERTH);

Port Wyndham, Jul 1974, Carr 3269 & Beauglehole

47047 (CANB, PERTH); Wyndham graveyard, Mar

1991, Petty s.n. (PERTH 1258222); Ord River station,

May 1985, Newland CN22 (PERTH); c. 4 km from

Wyndham along road to port, Apr 1987, Purdie 3309

(CANB).

Distribution and habitat: Neptunia

valida is endemic to Australia in Western

Australia, and apparently confined to the

eastern Kimberley region, mostly around

Wyndham and Kununurra (Map 5). Only

three specimen labels (all coastal) record

habitat information, viz. ‘disturbed roadside

near mangroves’, ‘next to tidal flat’ and ‘plain

just above supratidal plain’, but because the

species extends inland, it must have a broader

ecological range.

Phenology: Flowers and fruit are recorded

from March to July.

soe

Affinities: Neptunia valida and WN.

monosperma are the only upright species with

single-seeded fruits. Neptunia valida 1s very

different in the dense glandular indumentum

(vegetative parts glabrous in NV. monosperma)

and the (2—)3—4 hairy bracts (2 glabrous

bracts for NV. monosperma). Neptunia valida

is perhaps closest to N. scutata, but differs

by the upright habit, the lack of a nectary on

the petioles, the longer staminodes, and the

frequent presence of three or four bracts on

the peduncle.

Note: Some illustrations in the Flora of

Australia (Cowan 1998: 22, figs. E-—TI),

depict plant parts including mature fruits,

a nectary gland on the petiole, and 2 bracts

on the peduncles. They are labelled as

Neptunia dimorphantha, but all are in fact

N. xanthonema. Figs. E and F are said to be

based on Purdie 3309, but that specimen has

no petiole nectaries and all peduncles have

either 3 or 4 bracts. Clearly those illustrations

were not made using Purdie 3309 but the

actual voucher specimen is unknown. Purdie

3309 1s Neptunia valida.

Etymology: From the Latin validus, meaning

‘strong’. This is reference to the thick and

strong stems allowing an upright habit.

18. Neptunia xanthonema A.R.Bean sp. nov.

With affinity to V. monosperma, but differing

by prostrate habit, the conspicuous and

persistent stipules, the pods with glandular

hairs, the fewer leaflets per pinna, and

the larger bracts on the peduncle. Typus:

Australia. Northern Territory. Anthony

Lagoon, 19 February 1998, C.R. Michell

I31I & R. Carrow (holo: BRI [AQ1024483

comprising | sheet]; iso: DNA D0134434).

Neptunia dimorphantha var. clementii Domin,

Biblioth. Bot. 22(89): 802 (1926). Type:

Australia. Western Australia. “Between

Ashburton and Yule River’’, s.dat., E. Clement

S.n. (n.V.)

[Neptunia monosperma auct. non Domin:

Black (1948: 429)].

[Neptunia dimorphantha auct. non Domin,

pro parte: Whibley (1981: 114); Whibley

(1986: 513); Murray (1991: 380); Kenneally ef

al. (1996: 137); Wheeler (1992: 341)].

Illustrations: Cowan (1998: 22), as Neptunia

dimorphantha, but see note under N. valida;

Murray (1991: 380), as N. dimorphantha;

Kenneally et al. (1996: 137), as WN.

dimorphantha.

Shrub, perennial, terrestrial. Stems prostrate

or sprawling, to 0.25 m high, glabrous, sessile

glands present. Stipules persistent, broadly

ovate, 6—-9.7 mm long, 3—4.6 mm broad,

coriaceous, strongly nerved, glabrous; apex

attenuate; base obliquely cordate. Leaves

with 2 or 3(—4) pairs of pinnae; petioles 5—13

mm long, glabrous, with a prominent nectary

in the upper half, often just below first pair

of pinnae; rachis 8—28 mm long, with sparse

tubercle-based hairs, nectary present below

each pair of pinnae; rachis extension linear,

2.2—6 mm long; pinnae with 11-24 pairs of

leaflets, rachilla extension 0.9—1.8 mm long;

stipels slender, 0.2—0.8 mm long, divided,

gland-tipped. Largest leaflets 5.8-l11 mm

long, 1.6—2.4 mm wide, 3.7—4.6 times longer

than broad, glabrous or with a few marginal

cilia, venation of I-3 main veins, more

obvious on lower surface. Inflorescences

globose, some with dimorphic flowers;

peduncles 7-32 mm long at anthesis, 19—45

mm long at fruiting stage, usually glabrous

except at apex, sometimes sparsely hairy

throughout; bracts 2, positioned on proximal

one-third of peduncle, persistent, ovate,

amplexicaul, glabrous, lower one 5-—14.5

mm long; flowers 30—40 per inflorescence;

bracteoles oblanceolate, 0.9-1.1 mm _ long,

caducous, usually falling before anthesis.

Bisexual flowers: calyx campanulate, 0.9-1.3

mm long, glabrous; petals 1.5—2 mm long,

glabrous; stamens 5, filaments 2.2—2.8 mm

long, anthers 0.4—0.55 mm long, terminal

gland absent; style 1.3—2.1 mm long, glabrous;

ovary densely hairy. Neuter flowers: calyx

0.5—0.6 mm long; petals 1.2—1.4 mm long;

staminodes 5, 6—8.5 mm long, 0.5—-1 mm

wide. Pods broadly elliptical, l-seeded, 7.5—

9.5 mm long, 5.5—7 mm broad, indehiscent,

stipe 1-1.7 mm long, apex obtuse; surface

Austrobaileya 12: 59-106 (2022)

of pod with dense patent hairs 0.2—0.4 mm

long, at least some with glandular apex. Seeds

broadly elliptical in outline, 2.7—-3.6 mm long,

2.4—2.8 mm wide, brown. Figs. 2F, 2G, 10.

Additional selected specimens examined: Western

Australia. c. 5 km from Nanutarra on Mt Stuart Road,

Aug 1971, Ashby 4064 (AD); Jigalong Creek, near ‘Ethel

Creek’, NE of Newman, May 2006, Bean 25378 (BRI);

Mt Vernon Station, Nov 1976, Sarti 10/96 (PERTH);

Near Rudall River, May 1971, George 10776 (PERTH); 5

km W of Mulga Downs Station HS along Mulga Downs

Road, 91 km NE of Tom Price, Aug 2016, Lyons &

Coppen FV0589 (PERTH); Mardie HS, Mardie Station,

Aug 2002, Smith 136 (PERTH); Noonkanbah Station,

Fitzroy River area, May 1984, Stuart s.n. (PERTH

1669559); 4 miles [6 km] E of Derby, Jan 1971, Allan 559

(PERTH). Northern Territory. Track SW of Tobacco

Waterhole, Mittebah Station, Apr 2014, Jobson 11177

& Latz (DNA, NT); McArthur River, Feb 1976, Rice

2276 (CANB); 6.5 miles [10.5 km] E of Stuart Highway,

Beetaloo Station Road, Feb 1969, Must 384 (AD,

CANB, MEL); Burt Plain, 36 miles [58 km] N of Alice

Springs, May 1962, Chippendale 9014 (CANB, MEL,

NT); Narwietooma HS, Apr 1967, Maconochie 56 (AD,

CANB, DNA, NT); Charlotte Waters, c. 260 km SSE of

Alice Springs, May 1939, Crocker s.n. (AD 96434301).

Queensland. BURKE DISTRICT: Georgina River, just

SW of Camooweal, May 2006, Cumming 24151 (BRI).

SOUTH KENNEDY DISTRICT: SW of Mazeppa NP, Feb

1998, Fairfax 298 & Holman (BRI). GREGORY NORTH

DIsTRICT: Goomerchie Paddock, ‘Monkira’, c. 100 km E

of Bedourie, May 2007, Bean 26282 (BRI); Chartwage

Bore, Headingley, May 1985, Neldner 1776 & Stanley

(BRI). MITCHELL DtstrRIctT: Jireena Station, 90 km SSE

of Torrens Creek, Apr 2005, Booth 3830 & Thompson

(BRI). GREGORY SouTH DistRIcT: 10 miles [16 km] W

of Windorah, Jun 1967, Gittins 1252 (CANB); Windorah

to Yaraka Road, 24.4 km E from crossroads at Retreat,

Oct 1993, Slee 3332 & Lepschi (AD, BRI, CANB).

WARREGO District: Gilruth Plains, Feb 1941, Roe

RRID5 (CANB). New South Wales. Mount Murchison,

s.dat., Bonney 29 (MEL); Warri Gate, Sturt NP, May

1988, Marcom s.n. (NSW 213151). South Australia.

Eringa Waterhole, Lindsay Creek, Hamilton station, Apr

1985, Badman 1664 (MEL); 11 km S from Candradecka

Dam on Cordillo Downs — Innamincka road, May 1986,

Purdie 2931 (AD, CANB); c. 22 km NW of Stuart Creek

crossing (near Blower Waterhole) on track to Coward

Springs from Stuart Creek HS, Mar 1984, Jackson 5114

(AD); Welbourne Hill Station, 13 miles [21 km] S of

Hawks Nest Well, May 1980, Henshall 3023 (AD, NT);

Arckaringa Station, 21.5 km direct NNW of Arckaringa,

May 2000, Lang BSOP-450 (AD).

Distribution and habitat: Neptunia

xanthonema 1s endemic to Australia. It

is distributed in the Pilbara and western

Kimberley regions of Western Australia,

the northern part of South Australia, south-

western Queensland, southern Northern

Bean, Neptunia in Australia and Malesia

Territory and north-western New South Wales

(Map 12). It grows in red or brown clay soils

on alluvial flats or low rises in grassland or

Acacia woodland.

Phenology: Flowers and fruits are recorded

for every month of the year except December.

Affinities: Neptunia xanthonema difters

from N. scutata by the frequent presence of

nectaries on the leaf rachis, one adjacent to

each pair of pinnae (nectaries absent from

rachis in JN. scutata); peduncles usually

glabrous except at the apex (hairy throughout

for N. scutata); seeds 2.7-3.6 x 2.4—2.8 mm

(3.8—4.5 x 2.8-3.4 mm for N. scutata); and

the mostly larger leaflets, 5.8-11 mm long

(3-7 mm long for N. scutata). In addition, N.

xanthonema has 2 or 3(—4) pairs of pinnae (vs.

3—5(—6) pairs for NV. scutata).

Typification: Type material of Neptunia

dimorphantha var. clementii was sought

from PR, L, K and BM, without success. The

placement of this name as a synonym of N.

xanthonema is based on the description in the

protologue and assisted by the fact that there

are few species of Neptunia known to occur

in the area cited.

Notes: Specimens of Neptunia xanthonema

from the Derby area (western Kimberley

region) lack a nectary on the petiole or on the

rachis, but they are otherwise typical of the

species. This variant is accommodated in the

identification keys.

Etymology: The epithet is derived from

the Greek xanthos (yellow) and -nema

(thread) and refers to the conspicuous yellow

staminodes present on some inflorescences.

Excluded names

Neptunia spicata F.Muell. = Dichrostachys

spicata (F.Muell.) Domin

Neptunia cinerea (L.) FMuell. =

Dichrostachys cinerea (L.) Wight & Arn.

Dubious name

Neptunia acinaciformis (Span.) Miq., Fil. Ind.

Bat. 1: 51 (1855); Desmanthus acinaciformis

Span., Linnaea 15: 198 (1841).

Spanoghe (1841) described Desmanthus

acinaciformis in his posthumous treatise on

the flora of Timor, and it was transferred to

Neptunia some years later by Miquel. Windler

(1966) accepted this name, and applied it to

specimens that were formerly known as N.

javanica and collected from Java, India,

Burma and Thailand. Nielsen (1992) argued

that because no type material or iulustration

can be found for NV. acinaciformis, and because

the original description is non-diagnostic,

the species name should be regarded as

dubious. The present author agrees with that

conclusion.

Nielsen (1992) rejected a synonymy with

Neptunia javanica because that species has

more seeds per pod than the “4—5 seeds”

stated in the protologue for N. acinaciformis,

and because N. javanica has not been found

on Timor.

Acknowledgements

I thank the Directors of AD, CANB, DNA, L,

MEL, NT and PERTH for sending specimens

on loan, and Hannah McPherson (NSW) for

sending label data and hyperlinks to high-

quality specimen images of their Neptunia

holdings. Iam grateful to Otakar Sida, curator

of the herbarium at the National Museum in

Prague (PR), who sent high-quality images

of Domin’s Neptunia specimens, and John

McNeill (Royal Botanic Garden, Edinburgh)

for nomenclatural advice on the status of

N. prostrata and N. natans. Melody Fabillo

for providing Figures 9 and 10. Nicole

Crosswell provided the excellent illustrations.

I am_ grateful to Barbara Waterhouse

(Northern Australia Quarantine Strategy)

for the photograph of N. plena, and to Keith

McDonald and Lana Little for collecting N.

insignis for me from the far side of the Walsh

River causeway.

Austrobaileya 12: 59-106 (2022)

741280

Fig. 10. Neptunia xanthonema, showing leaves, inflorescences, peduncles and bracts (McKenzie 03/96, BRI).

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100

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(

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Fe _ a

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143 E 144 F |

Map 1. Distribution of Neptunia amplexicaulis f. amplexicaulis (@) and N. amplexicaulis {. richmondii (A).

Bean, Neptunia in Australia and Malesia 101

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102 Austrobaileya 12: 59-106 (2022)

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Map 4. Distribution of Neptunia heliophila.

130E 135E

Map 5. Distribution of Neptunia hispida (@) and N. valida (A).

Bean, Neptunia in Australia and Malesia 103

Map 6. Distribution of Neptunia insignis (~) and N. tactilis (A).

Map 7. Distribution of Neptunia longipila (®), N. paucijuga (A) and N. proxima (@).

104 Austrobaileya 12: 59-106 (2022)

Map 8. Distribution of Neptunia major.

Te hawt a

f , * ah F

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Map 9. Distribution of Neptunia monosperma.

Bean, Neptunia in Australia and Malesia 105

Map 11. Distribution of Neptunia scutata.

106 Austrobaileya 12: 59-106 (2022)

Map 12. Distribution of Neptunia xanthonema.

Pomax ammophila Ngugi (Rubiaceae), a new

species from arid, central Australia

Lorna B. Ngugi

Summary

Negugi, L.B. (2022). Pomax ammophila Ngugi (Rubiaceae), anew species from arid, central Australia.

Austrobaileya 12: 107-116. Pomax ammophila is described as new, distinguished from the related

P. rupestris F.Muell. by the recurved peduncles; the leaves, fruits and stems with abundant short

uncinate hairs and the larger fruit and seed. The new species is sporadically widespread on red sand

dunes in arid central Australia in the Northern Territory, Queensland, South Australia and Western

Australia. Notes are provided on distinctive features, habitat, phenology and conservation status,

along with detailed images and a distribution map.

Key Words: Rubiaceae; Pomax; Pomax ammophila; flora of Australia; flora of Northern Territory;

flora of Queensland; flora of South Australia; flora of Western Australia; new species; taxonomy

L.B. Ngugi, Queensland Herbarium and Biodiversity Science, Department of Environment and

Science, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066. Australia.

Email: lorna.ngugi@des.qld.gov.au

Introduction

Pomax Sol. ex DC. (Rubiaceae) is endemic

to Australia and is classified in the tribe

Anthospermeae Cham. & Schltdl. (Puff

1982). Puff described Anthospermeae as

comprising small trees or shrubs with

inconspicuously coloured, actinomorphic,

nectarless, odourless and wind pollinated

flowers. Together with Opercularia Gartn.,

Puff (1982) further classified Pomax into the

subtribe Operculariinae Benth. Molecular

data have provided some support for the

morphological classification outlined by Puff

(1982). Anderson ef al. (2001) constructed

a molecular phylogeny of Anthospermeae,

but with poor support for this subtribal

classification. Thureborn ef al. (2019)

concluded that Pomax formed a sister clade

to Leptostigma Arn. (subtribe Coprosminae

Fosberg) on the basis of plastid data; however,

when nuclear data was used, the relationship

was with Opercularia. This inconsistent

phylogenetic relationship was considered to

require further study, with Thureborn ef al.

(2019) continuing to support the placement

of both Opercularia and Pomax within

Operculariinae.

Pomax was described by the Swiss

taxonomist de Candolle (1830), with two

species P. glabra DC. and P. hirta DC. Richard

(1834) added a previously described species

from Opercularia as P. umbellata (Gaertn).

Sol. ex A.Rich. and Mueller (1853) named P.

rupestris F.Muell. Bentham (1867) in treating

Pomax for Flora Australiensis, reduced all

taxa to a single species, P. umbellata. Bailey

(1913) continued to recognise the genus as

consisting of a single species. Domin (1929)

then split this species into five varieties;

however, these names have not been applied in

Australian herbaria and remain of uncertain

application (CHAH 2011). Eichler (1965)

expressed doubts regarding the inclusion of all

material of Pomax under a single species and

included P. glabra, P. hirta and P. rupestris as

doubtful synonyms of P. umbellata.

Recent consensus has been to recognise

two described species in Pomax (CHAH

2022): P. umbellata and P. rupestris. Pomax

umbellata 1s a woody herb, prostrate or up to

40 cm tall, with stems pubescent or minutely

so; leaves with petioles 1-9 mm long, and

with leaf lamina ovate to obovate or narrowly

Accepted for publication 13 October 2022, published online 7 December 2022

108

so (Bentham 1867; Stanley & Ross 2002;

PlantNet 2022; VicFlora 2022). It is the

most widespread species of the genus and

erows on sandy soils derived from a broad

range of geologies in Eucalyptus dominated

woodlands in eastern Australia (Queensland

Herbarium 2021). Pomax rupestris 1s a

small shrub to about 50 cm tall with leaves

that are glabrous, sessile or shortly petiolate,

lanceolate-ovate or rarely elliptic (Flora NT

2022). It occurs from western to central and

southern parts of Australia (AVH 2022) in

rocky screes (WAH 2022).

A review was undertaken of the

morphological variation of Pomax umbellata

in Queensland, the only species recorded for

the state (Forster & Halford 2021). A single

collection from western Queensland did not

match this species and had been determined

by David Halford as Pomax sp. (Richter

DR282 & Turpin). After a second collection

was made from the same location, it could

be confirmed that this material represented

a new and distinctive species restricted to

sand dunes. Further investigation revealed

that this species extends into adjacent states

of Australia (AVH 2022). Botanists in

Western Australia and the Northern Territory

independently recognised this species within

their collections, applying different phrase

names. The Northern Territory herbartum

listed this first as Pomax A89438 Sand Dunes

(Dunlop 1995) then in 2011, listed it as

Pomax sp. Sand dunes (P.G.Wilson 752) NT

Herbarium, while Western Australia listed

it as Pomax sp. desert (A.S.George 11968)

WA Herbarium (CHAH 2011). Pomax sp.

Sand dunes (P.G.Wilson 752) NT Herbarium

became the accepted name for Pomax sp.

desert (A.S.George 11968) WA Herbarium

(CHAH 2018). In this paper a new species for

arid Australia is described based upon these

collections and of the author.

Materials and methods

This study is based on a morphological

examination of herbarium — collections

held at the Queensland Herbarium (BRI)

and specimens loaned from the Northern

Territory Herbarium (DNA, NT), the State

Herbarium of South Australia (AD) and the

Austrobaileya 12: 107-116 (2022)

Western Australian Herbartum (PERTH). The

measurements for corolla, stamens and style

are based on dried material reconstituted with

hot water, while all other measurements were

from dried material using aruler or microscope

eyepiece graticule. Stereomicroscopy and

Scanning Electron Microscopy (SEM) were

also used to clearly visualise and characterise

the stem indumentum, presence of colleters

in stipules, leaf indumentum, fruit dehiscence

and position of seeds, and idioblasts on the

seeds. The distribution map was compiled

from localities and/or geocode information

provided on the labels of specimens at AD,

BRI, DNA, NT and PERTH and was mapped

using DIVA-GIS Software.

Taxonomy

Pomax ammophila Neug1, sp. nov.

With affinity to Pomax rupestris, differing

by the presence of uncinate hairs on stems,

leaves and fruits; recurved peduncles; fruit

with larger, campanulate capitula; larger

seeds and a sand dune habitat. Typus:

Australia: Queensland. GREGORY SOUTH

District: 14 km from Carrinya Homestead,

about 60 km from Windorah, 28 September

2021, L.B. Negugi 112 & G.P. Turpin (holo:

BRI [AQ1026842, comprising 2 sheets]; 1so:

AD, CANB, MEL, NSW, NT, PERTH).

[Pomax umbellata auct. et pro parte, non

DC.: Marsden (1981: 283-284) (referring to

‘dunes’ ); Green (1985: 155)].

Pomax A89438 Sand Dunes: Dunlop (1995:

99); CHAH (2011).

Pomax sp. Sand dunes (P.G.Wilson 752) NT

Herbarium: CHAH (2011, 2018).

Pomax sp. desert (A.S.George 11968) WA

Herbarium: WAH (2011); Percy-Bower (2019:

13).

Perennial, woody subshrub, erect, 10—50 cm

high. Stems terete, indumentum comprising

mixed short uncinate or flexuose, or erect

hairs, 0.1—0.3 mm long. Mature stem with

outer surface layers decorticating in long

vertical plates. Stipules interpetiolar, narrowly

triangular or sometimes lobed, 0.9-5 mm

long, 0.4—1.4 mm wide; hairs uncinate 0.05—

Negugi, Pomax ammophila

0.3 mm long, mostly along margins; colleters

along margins. Leaves opposite, narrowly

elliptic or lanceolate, 7.5—26 mm long, 2.2—7

mm wide, light green, margins fimbriate; base

sessile or attenuate; apex acute or attenuate;

laminae indumentum predominantly 0.05—

0.4 mm long, but up to 0.5 mm long on veins

and along margins, abaxial indumentum

longer and denser compared to adaxial side.

Inflorescence terminal or axillary near the

top; 1-3 flowers fused at their bases to form

a capitulum; andromonoecious. Flowering

peduncles 0.5—5 mm long; indumentum

uncinate, reflexed or erect, 0.05—0.3 mm long,

elongating as it matures; calyx lobes variable

in shape and size. Pistillate flowers: corolla

2—3.7 mm long, lobes 5 or 6, 0.1—1.5 mm long,

green, indumentum external, uncinate, 0.05—

0.4 mm long, sparse; style filiform, 3—6.5

mim long, deeply bifid more than * its length,

stigma papillate. Staminate flowers: corolla

2—4 mm long, 5—6 lobed, 0.6—2.5 mm long,

indumentum 0.01—0.1 mm long, very sparse,

mostly around the outside edges; stamens

3—5, filament exserted for 1-5 mm, anthers

1.3-2.5 long. Fruiting peduncles 1.5—6.5

mm long, recurved; calyx lobes variable in

shape and size, 2.4—3.2 mm long. Fruits an

operculate campanulate capsule, non-fleshy,

3—5 mm long, 2—5 mm wide; locules 1—3, seed

per locule 1; indumentum uncinate, 0.01—0.2

mm long, very sparse; operculum deciduous

releasing the seed; empty capitulum persistent

for some time after seed dehiscence. Seeds

obdeltoid, 2.2—-3.8 mm long, 1.4—2 mm wide

and 0.3-0.9 mm broad, ridge on dorsal

surface 2 to % of seed length; covered with

white idioblasts containing calcium oxalate

crystals (raphides). Figs. 1—4.

Additional specimens examined: Western Australia.

80 km NNE of Bondya Homestead, c. 100 km N of

Laverton, Aug 1968, Wilson 7384 (PERTH); Hill N of

Good Camp Rockhole, South Carnarvon Range, Little

Sandy Desert, Aug 1998, Blackhouse et al. BEMJ 180

(PERTH); On S side of Lake Kerrilyn, c. 5.9 km on a

bearing of 25° from Mt Methwin, Birriliburu Indigenous

Protected Area, Aug 2012, Gibson et al. NG7030 (BRI,

PERTH). Northern Territory. Wild Eagle Plain,

Temple Downs Station, Sep 2014, Jobson 11515 & Latz

(DNA, K, MEL, PERTH); c. 5 km W [of] Redbank

Yard, Owen Springs Station, Sep 2000, Albrecht 9402

(DNA); 3 km NE of Mt Winter, 80 km WNW of Kings

Canyon, Aug 3003, Latz 18953 & Albrecht (DNA, MEL);

109

Norman Gully, Finke Gorge NP, 0.2 km E of fenceline,

Jun 2006, Schubert 137 (DNA). Queensland. GREGORY

SOUTH District: 14 km from Carrinya Homestead, c. 60

km from Windorah, Sep 2011, Richter DR282 & Turpin

(BRI). South Australia. 30 km W of Vokes Corner,

along track to Serpentine Lakes, Aug 1980, Weber 6408

(AD, KRA, PE); 11 km NE Mount Finke, Denton’s track,

Oct 1987, Simon 1083 (AD).

Distribution and habitat: Pomax ammophila

occurs in central inland Australia in Western

Australia, Northern ‘Territory, Queensland

and South Australia (Map 1). It 1s confined

to red sand dunes in open shrublands over

Triodia hummock-dominated grassland. In

the Northern Territory, typical associated

species include Calandrinia polyandra Benth.,

Calotis erinacea Steetz, Chrysocephalum

apiculatum (Labill.) Steetz, Eremophila willsii

F.Muell. and Grevillea stenobotrya F.Muell.

In Western Australia, typical associated

species include Acacia ligulata A.Cunn. ex

Benth., Corymbia_ chippendalei (D.J.Carr

& S.G.M.Carr) K.D.Hill & L.A.S.Johnson,

Eremophila_ platythamnos Diels and_ 4G.

stenobotrya. In Queensland, P. ammophila

has been recorded only near Windorah in

Sparse to open grassland dominated by

Trachymene cyanantha Boyland and Triodia

basedowii E.Pritz, with scattered Acacia

species, Corymbia aparrerinja K.D.Huill

& L.A.S.Johnson, C. terminalis (F.Muell.)

K.D.Hull & L.A.S.Johnson and Dodonaea

species (Fig. 5). The Queensland Regional

Ecosystem (RE) 1s 5.6.5/5.5.3b (Neldner ef al.

2020); this is described as a variable sparse

to open-herbland or shrubland on dune flanks,

crests and sandy interdunes.

Phenology: Flowers have been recorded from

May to October, and fruits recorded for every

month of the year.

Notes: The peduncles of Pomax ammophila

at the fruiting stage are recurved (Fig. 1A

& B), unlike P. umbellata which has erect

peduncles. The surface of the leaves, fruits

and stems are rough to the touch (at least when

dried), likely because of the abundant short

uncinate hairs (Fig. 2C), while P. rupestris

is glabrous and leathery to the touch, and P.

umbellata is hairy and feels smooth. The hairs

on the abaxial leaf surface of P. ammophila

are longer and denser compared with those on

110 Austrobaileya 12: 107-116 (2022)

Fig. 1. Flowers and fruit of Pomax ammophila. A. & B. mature fruits showing recurved peduncles. C. empty fruit

persistent on plant. D. bisexual flower showing pistillate and staminate parts. All from Neugi 1/2 & Turpin (BRI).

Negugi, Pomax ammophila 111

Fig. 2. Indumentum on vegetative parts of Pomax ammophila as seen under stereomicroscope. A. interpetiolar stipule

abaxial surface showing the colleters (indicated by blue arrow). B. interpetiolar stipule adaxial surface showing the

colleters (indicated by blue arrow). C. young stem showing uncinate hairs. A from Jobson 11515 & Latz (NT); B & C

from Richter 282 & Turpin (BRI).

112

Fig. 3. Fruit of Pomax ammophila. A. fruit showing

seed position and idioblasts on seeds. B. inside view of

fruit cap lid showing three locules. From Negugi //2 &

Turpin (BRI).

the adaxial leaf surface. The fruit size (3-5

mm long, 2—5 mm wide) (Fig. 1A—D) and the

seed size (up to 3.8 X 2 mm) (Fig. 4A & B)

are also much larger than the other species of

Pomax.

Colleters on the stipules of Pomax

ammophila are congruent with those described

for a large number of species of Rubiaceae

(Robbrecht 1988). A study of colleter types in

Rubiaceae found that colleters with uniform

appearance like those of Cinchona calisaya

Wedd., are widespread in the tribe in which

Pomax was included, although the genus was

not directly mentioned (Lersten 1975). In

recent years, colleters have only been noted

Austrobaileya 12: 107-116 (2022)

to be present or absent (WAH 2022). The

colleters of P. ammophila are cylindrical to

conical, amber in colour, tapering towards the

tip on rims of the interpetiolar stipules (Fig.

2A & B).

The occurrence of calcium oxalate

crystals in all organs of Pomax ammophila are

congruent with the description for Rubiaceae

(Robbrecht 1988). Crystals such as druses

and styloids are usually observable with the

naked eye or under low magnifications as

translucent points in leaf blades and so may

serve as practical identification or taxonomic

characters (Robbrect 1988; Teixeira ef ail.

2016). Specialised white cells called idioblasts

containing needle like raphide crystals are

found in the seeds of P. ammophila (Fig. 4A-

C). Druses found on leaf surfaces and hairs of

P. ammophila are noticeably shinier than in

other species of Pomax.

The fruit of Pomax (Fig. 3A & B) is a

pome and has two parts: the capitulum is the

lower part shaped like a cup and surrounded

by the calyx in variable shapes and sizes;

and the operculum is the fruit’s apical lid,

with an almost round shape. When the fruit

has matured, the operculum opens up along

a transverse dehiscence line. Seeds dehisce

through the operculum and empty fruits

persist for some time (Fig. 1C).

Conservation status: Pomax ammophila 1s

widespread with many populations scattered

over inland arid Australia. It occurs in a

number of conservation reserves and protected

areas in Western Australia, Northern Territory

and South Australia; however, 1n Queensland

it only occurs on private grazing land. The

recommended conservation status for the

species 1s Least Concern based on criterion

B UCN 2021).

Etymology: The specific epithet 1s derived

from the Greek ammophila, which means

sand-loving. This refers to the sand dune

habitat where Pomax ammophila grows and

also pays recognition to one of its phrase

names Pomax sp. Sand dunes (P.G.Wilson

752) NT Herbarium.

Negugi, Pomax ammophila 113

Fig. 4. Seeds of Pomax ammophila showing white idioblasts. A. dorsal view. B. ventral view. C. SEM of idioblast

showing multiple needle-like raphides crystals (indicated by blue arrow). A & B from Richter 282 & Turpin (BRI); C

from Neugi 112 & Turpin (BRI).

114 Austrobaileya 12: 107-116 (2022)

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Special thanks to Gerry Turpin for his help

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Sincere thanks to Melody Fabillo for her id=&name=Pomaxtumbellatatvar.+mit

; ; helliana+Domin&inc. scientific=&inc

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(2011).

116 Austrobaileya 12: 107-116 (2022)

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Map 1. Distribution of Pomax ammophila based on cited herbarium specimens.

Austrobaileya publication dates 1977-2022

Paul I. Forster

Queensland Herbarium and Biodiversity Science, Department of Environment and Science,

Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong 4066, Queensland, Australia. Email:

paul.forster@des.qld.gov.au

Individual numbered parts (comprising multiple papers, notes and book reviews)

in Austrobaileya, were generally published on an annual basis between 1977 and

2020. Each volume usually comprised four parts (initially five), published over

one to five years. The publication dates of previous parts were usually given in the

subsequent one and are of significance for documenting when new names were

officially published in relation to listing in online databases such as the Australian

Plant Name Index (APNI) and the /nternational Plant Names Index (1PN]).

Between 1977 and 2019, a single numbered part was generally published in any

one year as hard copy. Prior to 2020, commencing with 10(3) in 2015, papers were

subsequently placed online after hard copy publication. In 2020, papers were first

published online with the dates of publication indicated on the individual papers;

these were still within numbered parts, followed by a hard copy volume at the end

of that year.

As of 2020, volumes were compiled on an annual basis, 1.e. all papers published

in a calendar year comprise a single volume.

From 2021, publication of papers was moved entirely online, without numbered

parts and with the dates of publication indicated on the individual papers.

The publication dates of volumes, individual parts and papers therein are

indicated in Table 1. Journal parts with single, or a couple of missing pages are

mainly the consequence of commencing new parts on an odd numbered page.

The responsible Editor and Editorial Committee are also provided as they were

sometimes missing from a published part, or the information was lost when the

parts were bound as library volumes.

Accepted for publication 7 November 2022, published online 7 December 2022

118 Austrobaileya 12: 117-120 (2022)

Table 1. Austrobaileya publication dates 1977-2022, (* indicates missing pages due to

commencing journal parts on an odd page)

Volume Part Pagination Editor Editorial Committee Date of

Number | Number Publication

2 109-202 L. Pedley G.P. Guymer 29 August 1985

T.D. Stanley

2 3 203-306 L. Pedley G.P. Guymer 24 July 1986

T.D. Stanley

2 4 307-418 L. Pedley E.M. Ross 24 September 1987

T.D. Stanley

2 5 419-593 L. Pedley E.M. Ross 5 October 1988

T.D. Stanley

4 2 139-294" E.M. Ross R.J.F. Henderson 1 September 1994!

T.D. Stanley

4 3 297-453° L.W. Jessup R.J.F. Henderson 5 October 1995

B.K. Simon

4 4 455-696" L.W. Jessup R.J.F. Henderson 20 February 1997

B.K. Simon

5 l 1-153" L.W. Jessup R.J.F. Henderson 16 December 1997

B.K. Simon

5 2 157-365? L.W. Jessup R.J.F. Henderson 25 January 1999

B.K. Simon

5 3 405-5867 L.W. Jessup R.J.F. Henderson 29 December 1999

B.K. Simon

5 4 589-735" L.W. Jessup R.J.F. Henderson 15 December 2000

B.K. Simon

] 1-176 L.W. Jessup R.J.F. Henderson 11 December 2001

B.K. Simon

2 177-345" L.W. Jessup R.J.F. Henderson 30 September 2002

B.K. Simon

3 349-579" L.W. Jessup R.J.F. Henderson 3 December 2003

B.K. Simon

Forster, Austrobaileya publication dates lly

Volume Part Pagination Editor Editorial Committee Date of

Number | Number Publication

4 581-1006" L.W. Jessup P.D. Bostock 6 December 2004

B.K. Simon

7 l 1-252 P.I. Forster P.D. Bostock 21 December 2005

G.P. Guymer

7 2 253-385" P.I. Forster P.D. Bostock 4 December 2006

G.P. Guymer

7 3 387-576" P.I. Forster P.D. Bostock 30 November 2007

G.P. Guymer

7 4 5377-7137 P.I. Forster P.D. Bostock 18 December 2008

G.P. Guymer

1-105" P.I. Forster P.D. Bostock 23 November 2009

G.P. Guymer

107-223" P.I. Forster P.D. Bostock 7 December 2010

G.P. Guymer

225-440" P.I. Forster P.D. Bostock | December 2011

G.P. Guymer

441-723 P.I. Forster P.D. Bostock 21 December 2012

G.P. Guymer

1-154 PI. Forster P.D. Bostock 17 December 2013

G.P. Guymer

155-320 P.I. Forster P.D. Bostock 3 December 2014

G.P. Guymer

321-461" P.I. Forster P.D. Bostock 3 November 2015

G.P. Guymer

“4

463-600" P.I. Forster P.D. Bostock 8 December 2016

G.P. Guymer

1-206 P.I. Forster P.D. Bostock 18 December 2017

G.P. Guymer

207-289" P.I. Forster G.P. Guymer 14 November 2019

D.A. Halford

291-547" P.I. Forster G.P. Guymer 23 August 2019

D.A. Halford

549-664" P.I. Forster G.P. Guymer 20 April 2020

D.A. Halford

I—55 P.I. Forster G.P. Guymer 24 March 2021

D.A. Halford

56-123 P.I. Forster N. Fechner 15 September 2021

G.P. Guymer

D.A. Halford

124-169 P.I. Forster

n PI. Forster

—

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—

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N. Fechner 22 December 2021

G.P. Guymer

A.E. Wood

G.K. Brown 29 June 2022

N. Fechner

n/a

A.E. Wood