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
© Queensland Herbarium 2022. ISSN 2653-0139 (Online)
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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
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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.
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| 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.
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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)
al 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.#
yf - = a 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:
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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 © CSIRO
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
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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
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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.
1] References ANON. (7 Oct 1886). Departures: 2. Cairns Post.
Cairns, Queensland. https://trove.nla.gov. au/newspaper/article/39423230, accessed 20 August 2021.
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BACHMAN, S., Moar, J., Hitt, A.W., DE LA TORRE, J.. & ScoTt., B. (2011). Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. In V. Smith & L. Penev (eds.), e-/nfrastructures for data publishing in biodiversity science. ZooKeys 150: 117-126. (Version BETA).
BAILEY, F.M. (1899). Kayea larnachiana [Guttiferae]. The Queensland Flora |: 105. H.J. Diddams & Co.: Brisbane.
BEENTJIE, H. (2010). The Kew Plant Glossary. Kew Publishing: Kew, Surrey.
BENTHAM, G. & Hooker, J.D. (1862). Hypericineae, Guttiferae. Genera Plantarum 1(1): 163-177. L. Reeve & Co. London.
CABRAL, F.N., TRAD, R.J., AMORIM, B.S., MACIEL, J.R., ESTANISLAU DO AMARAL, M.C. & STEVENS, P. (2021). Phylogeny, divergence times, and diversification in Calophyllaceae: Linking key characters and habitat changes to the evolution of Neotropical Calophylleae. Molecular Phylogenetics and Evolution 157: 107041.
CHRISTOPHEL, D.C. & HYLAND, B.P.M. (1993). Leaf Atlas of Australian Tropical Rain Forest Trees. CSIRO Publications: Melbourne.
CHAH |COUNCIL OF HEADS OF AUSTRALASIAN HERBARIA] (2005). Australian Plant Census. https://biodiversity.org.au/nsl/services/search/
names, accessed 30 July 2021.
— (2021). Australian Plant Census. https:// biodiversity.org.au/nsl/services/search/names, accessed 30 July 2021.
COOPER, W. & COOPER, W.I. (2004). Fruits of the Australian Tropical Rainforest. Nokomis Editions: Melbourne.
DEH [DEPARTMENT OF THE ENVIRONMENT AND HERITAGE] (2004). Species Profile and Threats Database. http://www.environment.gov.au/cgi- bin/sprat/public/sprat.pl, accessed 30 July 2021.
12
FROGGATT, W.W. (1887). Travels in Northern Queensland. The Bendigo Advertiser, 9 March 1887, p. 1. Sandhurst, Victoria: R.R. Haverfield. https:// trove.nla.gov.au/newspaper/article/88926416, accessed 20 August 2021.
HYLAND, B.P.M. (1982). Mesua sp. aff. M. elmeri. A revised card key to rainforest trees of North Queensland. CSIRO: Melbourne.
HYLAND, B.P.M., & WHIFFIN, T.P. (1993). Mesua sp. (=RFK/3128). Australian tropical rainforest trees: an interactive identification system 1. CSIRO: Melbourne.
HYLAND, B.P.M., Gray, B. & ELiIck, R.W. (1994). Appendix I: Provisional Species List. In W.E. Cooper, W.E. & W.T. Cooper, Fruits of the Rain Forest, pp. 300-313. Geo: Sydney.
HyYLAND, B.P.M., WHIFFIN, T.P., CHRISTOPHEL, D.C., GRAY, B., ELIcK, R.W. & ForD, A.J. (1999). Australian tropical rainforest trees and
shrubs:60. CSIRO: Collingwood.
HYLAND, B.P.M., WHIFFIN, T.P. & ZICH, F.A. (2010). Mesua_ sp. Boonjee (A.K.Irvine 1218). Australian Tropical Rainforest Plants, Edition 6. [No longer online].
IucNn (2012). International Union for the Conservation of Nature. JUCN Red List categories and Criteria: Version 3.1, 2nd edition. https:// www.iucnredlist.org/resources/categories-and- criteria, accessed 30 July 2021.
Jessup, L.W. (1994). Clusiaceae. In R.J.F. Henderson (ed.), Queensland Vascular Plants: Names and Distribution, p. 74. Queensland Herbarium: Indooroopilly.
—— (1997). Clusiaceae. In R.J.F. Henderson (ed.), Queensland Vascular Plants: Names and Distribution, p. 50. Queensland Herbarium, Department of Environment: Indooroopilly.
— (2002). Clusiaceae. In R.J.F. Henderson (ed.), Names and Distribution of Queensland Plants, Algae and Lichens, p. 49. Queensland Herbarium, Environmental Protection Agency: Brisbane.
— (2007). Clusiaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2007, p. 49. Queensland Herbarium, Environmental Protection Agency: Brisbane.
— (2010). Clusiaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2010, p. 44. Queensland Herbarium, Department of Environment & Resource Management: Brisbane.
Austrobaileya 12: 1-13 (2022)
— (2019). Clusiaceae. In G.K. Brown & P.D. Bostock (eds.), Census of the Queensland Flora 2019. Queensland Department of Environment and Science, Queensland Government: Brisbane. https://data.qld.gov.au/dataset/census-of-the- queensland-flora-2019, accessed 30 July 2021.
— (2021). Clusiaceae. In G.K. Brown & P.D. Bostock (eds.), Census of the Queensland Flora 2021. Queensland Department of Environment and Science, Queensland Government: Brisbane. https://www.data.qld.gov.au/dataset/census-of- the-queensland-flora-2021, accessed | February 2022.
JsToR (2021). Sayer, W.A. (fl. 1886-1897). Global Plants — Plant Collectors. https://plants.jstor. org/stable/10.5555/al.ap.person.bm000153859, accessed 30 July 2021.
KOSTERMANS, A.J.G.H. (1969). Kayea Wall. and Mesua L. (Guttiferae). Reinwardtia 7: 425—431.
MUELLER, F.J.H. (1887). Descriptions of new Australian plants (continued). The Victorian Naturalist 3(9): 126-127.
Notis, C. (2004). Phylogeny and character evolution of Kielmeyeroideae (Clusiaceae) based on molecular and morphological data. M.Sc. Thesis, University of Florida: Gainesville, Florida, USA.
QPNS [Queensland Place Names Search] (2022). https:// www.qld.gov.au/environment/land/title/place- names/search, accessed 28 March 2022.
RUHFEL, B.R., Bove, C.P., PHILBRICK, C.I. & DAVIS, C.C. (2016). Dispersal largely explains the Gondwanan distribution of the ancient tropical clusioid plant clade. American Journal of Botany 103: 1117-1128.
STEVENS, P.F. (1993). A new species and new combination in Clusiaceae—Calophylloideae from New Guinea. 7Jelopea 5: 359-361.
— (2001 onwards). Calophyllaceae. Angiosperm Phylogeny Website, Version 14. http://www. mobot.org/MOBOT/research/A Pweb/, accessed 30 July 2021.
— (2007). Clusiaceae-Guttiferae. In K. Kubitzki (ed.),
Flowering plants: Eudicots; Berberidopsidales, Buxales, Crossosomatales, Fabales p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales, Vitales, Zygophyllales, Clusiaceae Alliance, Passifloraceae Alliance, Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae. Series Title: The Families and Genera of Vascular Plants LX., pp. 48—66. Springer Verlag: Berlin/New York.
Cooper & Zich, Kayea in Australia
THOMAS, M.B. & MCDONALD, W.J.F. (1987). Rare and threatened plants of Queensland: a checklist of geographically restricted, poorly collected and/or threatened vascular plant species, p. 21. Queensland Herbarium: Indooroopilly.
— (1989). Rare and threatened plants of Queensland: a checklist of geographically — restricted, poorly collected and/or threatened vascular plant species, Edition 2, p. 22. Queensland Herbarium: Indooroopilly.
Tscs |THREATENED SPECIES AND COMMUNITIES SECTION, BIODIVERSITY GROUP, ENVIRONMENT AUSTRALIA (8 JULY 1998)|. Endangered Species Protection Act 1992. Schedules |, 2 & 3: p. 23. Environment Australia, Canberra.
TURLAND, N.J., WIERSEMA, J.H., BARRIE, F.R., GREUTER, W., HAWKSWoRTH, D.L., HERENDEEN, P.S., KNAPP, S., KUSBER, W.-H., Li, D.-Z., MARHOLD, K., May, T.W., McNEILL, J.. MONRO, A.M., PRADO, J., PRicE, M.J. & SmitTH, G. F. (eds.) (2018). International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashititten: Koeltz Botanical Books. https://www.iapt-taxon.org/ nomen/main.php, accessed 20 August 2021.
WALLICH, N. (1831), Kayvea floribunda. Plantae Asiaticae Rariores 3: 5, t. 210. Treuttel & Wiirtz: London.
ZAKARIA, R. & CHOONG, C.Y. (2007). Systematic study on Guttiferae Juss. of Peninsular Malaysia based on Plastid Sequences. 7ropics 16: 141-— 150.
ZICH, F.A., HYLAND, B.P.M., WHIFFIN, T. & KERRIGAN, R.A. (2020). Mesua sp. Boonjee (A.K.Irvine 1218). Australian Tropical Rainforest Plants, Edition 8. https://apps.lucidcentral. org/rainforest/text/entities/mesua sp _ boonjee a k.irvine 1218.htm, accessed 30 July 2021.
13 \
»145_5° 145.75° ine
TRIBULATION
\,
Pa CAIRNS 4.~- » p
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. au
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
© Queensland Herbarium 2022. ISSN 2653-0139 (Online)
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
[5
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
16
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;
18
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.
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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
© Queensland Herbarium 2022. ISSN 2653-0139 (Online)
20
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
21
(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.
References
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BULTITUDE, R.J., GARRAD, P.D., ROBERTS, C.W. (1998). Maytown Region, Queensland, 1:100 000 Sheet 7765 and Part 7865. 1* edition. Queensland Department of Mines and Energy: Brisbane.
COOPER, W. & COOPER, W.I. (2004). Fruits of the Australian Tropical Rainforest. Nokomis: Melbourne.
Cowl, I.D., KERRIGAN, R.A. & Drxon, D.J. (2013). Hibiscus. In P.S. Short & I.D. Cowie (eds.), Flora of the Darwin Region |: 12—19. Northern Territory Herbarium, Department of Land Resource Management. http://eflora.nt.gov.au/ viewfile?file id=4721, accessed 16 February 2021.
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HYLAND, B., WHIFFIN T., CHRISTOPHEL D.C., GRAY, B. & Etick R.W. (2020). Australian Tropical Rainforest Plants 8" Ed. CSIRO, CANBR, ATH, ABRS: Canberra. https://apps.lucidcentral.org/ rainforest/text/entities/hibiscus meraukensis. htm, accessed 16 February 2021.
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WHEELER, J.R. (1992). Hibiscus. In J.R. Wheeler (ed.) Flora of the Kimberley Region, pp. 213-223. Department of Conservation and Land Management: Como.
WILson, F.D. (1974). Aibiscus section Furcaria (Malvaceae) in Australia. Australian Journal of Botany 22: 157-82.
— (2006). A distributional and cytological survey of the presently recognized taxa of Hibiscus section Furcaria (Malvaceae). Bonplandia 15: 53-62.
WILSON, F.D. & CRAVEN, L.A. (1995). Two new species of Hibiscus section Furcaria DC. (Malvaceae) from northern Queensland. Austrobaileya 4: 439-447,
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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
© Queensland Herbarium 2022. ISSN 2653-0139 (Online)
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
Ze
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.
30
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
31
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
32
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
33
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).
34
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).
36
. . = 7 a? ' ; > —w . ; j ' 7 eS . » . . ~_—. i ee : a. + F = ft ae ee, x tae eo tee Pe 1 tare *. - <2 —— er > wae ‘ “ - * UT 7 - “* “prig __ » i oe y . - h* 7 a Ate = ee le ne ‘py ok. 7 _ - e - > - sO ) — ~ : > os wes a - - — wa : % ws ae a ey ie 4 tiny , Ty - - ’ o~ : = =
Austrobaileya 12(1):; 26-58 (2022)
intercostal costal zone
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.
a]
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
s 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 1 100 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.
40
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
:;
SR _&
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 i : :
£ -_——!1 eee
6
pesieseas jyHuAdoOS 8 =
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.
v
G == eee || bill
9
Z =
paniesel 1u6iuAdoS 8 =
Det.: Thompson, E.J., Mar 2013 Dup.: 359.0 Poaceae
~NN
RI- 0219 | Prep at BRI: Sheet
(ey)
co
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
s
Fig. 13B. Holotype of Cryptachne columboola (BRI [AQ1021906], sheet 2 of 2).
46 Austrobaileya 12(1): 26-58 (2022)
5mm
oe : : 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
NE
RI- 02 Prep at BRI: Sheet
2 3 ) < =. o > + = ® ” © < @ a.
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
TM
rn fe) .- > =e a > re re | © a © P D 2.
NN
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.
52
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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35
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56
Austrobaileya 12(1): 26-58 (2022)
Legend
C. coolumboola
— «<= boundary of climatic zones Ps *C duaringa BC
T equatorial to tropical S subtropical
D desert
Ds semidesert
M temperate
Map 1. Distribution of species of Cryptachne based on BRI collection records. Climatic zones of Australia modified
from BOM (2020) following Thompson (2021c).
57
Thompson, Cryptachne
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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
© Queensland Herbarium 2022. ISSN 2653-0139 (Online)
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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
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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
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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
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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.
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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
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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
67
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]).
68
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
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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.
70
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).
TI
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.
72
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
73
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.
74 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
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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.
76
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.
78
(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.
T9
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).
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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:
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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.
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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