Lomandra grayi Jian Wang ter (Laxmanniaceae), a new Species from north-east Queensland, Australia

Jian Wang!

Summary

Wang, J. (2023). Lomandra grayi Jian Wang ter (Laxmanniaceae), a new species from north-east Queensland, Australia. Austrobaileya 13: 1-6. Lomandra grayi Jian Wang ter from tropical north- east Queensland is described, illustrated and compared to the morphologically similar species L. filiformis. Notes are provided on its distribution (including a map), habitat, phenology and affinities. A conservation status of Least Concern is recommended.

Key Words: Laxmanniaceae; Lomandra; Lomandra filiformis; Lomandra grayi; flora of Australia;

flora of Queensland; taxonomy; new species

‘Queensland Herbarium and Biodiversity Science, Department of Environment and Science, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: Jian.

Wang(@des.qld.gov.au

Introduction

The genus Lomandra Labill. (Laxmanniaceae) includes 59 species to date (Wang 2021; CHAH 2022). There are 56 species endemic to Australia, two endemic to Australia and New Guinea and one endemic to New Caledonia (Lee & Macfarlane 1986; Wang 2021). Lomandra has had various family placements over the past decades, including Xanthorrhoeaceae in ‘Flora of Australia’ (Lee 1966; Lee & Mactarlane 1986), Dasypogonaceae (Briggs 1986; Chase et al. 1995) and Laxmanniaceae (Chase & Stevens 1998; Wang 2021). The genus 1S more often placed in a broadly circumscribed Asparagaceae under subfamily Lomandroideae Thorne & Reveal (Chase ef al. 2009; Gunn ef al. 2020; Govaerts ef al. 2022). The family Laxmanniaceae 1s retained here following the systematics applied at BRI where a narrower circumscription of monocot families 1s preferred.

The genus Lomandra was revised by Lee & Macfarlane (1986) for ‘Flora of Australia’, and they recognised 15 species and four subspecies from Queensland. Currently, there are 19 recognised species 1n Queensland, as well as three non-autonymic subspecies (Wang 2021). A taxonomic review of Lomanadra in Queensland is underway.

Lomandra_ filiformis (Yhunb.) Britten belongs to Lomandra section Lomandra, series Sparsiflorae (Benth.) A.T.Lee (Lee & Macfarlane 1986). It includes three subspecies, L. filiformis subsp. filiformis, L. filiformis subsp. coriacea A.T.Lee, and L. filiformis subsp. flavior A.T.Lee. The first 1s distributed throughout eastern Queensland and New South Wales, the second in south- east Queensland, New South Wales and Victoria, and the third is restricted to north- east New South Wales.

Examination of herbarium material of Lomandra, especially from north-east tropical Queensland, has revealed the existence of a distinctive species that has been misidentified, mostly as Lomandra filiformis subsp. filiformis. The new species Lomandra grayi 1s notably different from L. filiformis s.1. in the dissimilar inflorescences of the male and female plants and the male flowers being arranged perpendicularly on the rachis.

Materials and methods

This study is based on morphological examination of Lomandra accessions from the Queensland Herbarium (BRI) and the National Herbarium of New South Wales (NSW). Only the BRI and CNS

Accepted for publication 17 January 2023, published online | March 2023

© Queensland Herbarium 2023. ISSN 2653-0139 (Online)

2

duplicates of the type collection have been examined, with the remaining duplicates to be distributed from CNS after publication. All measurements were undertaken using a binocular microscope and are based on dried material, except the dimensions of florets, which were reconstituted with boiling water. High resolution images of the type specimen of L. filiformis stored at UPS were viewed in detail. Dimensions of measurements are inclusive, 1.e. 1.0—1.8 is given as I—1.8.

Herbarium acronyms follow Thiers (updated continuously). Common abbreviations in the specimen citations are CYP (Cape York Peninsula), Mt (Mount or Mountain, except where part of a National Park or State Forest name) and NP (National Park).

Taxonomy Lomandra grayi Jian Wang ter, sp. nov.

Resembling Lomandra filiformis and _ its subspecies, differing in the dissimilar appearance of male and female inflorescences, longer male inflorescence, longer female scapes, the longer pedicel of male flowers, and male flowers arranged perpendicularly to rachis. Type: Australia. Queensland. Cook District: Portland Roads, 2 km E of Brown Creek, 10 June 2004, B. Gray 8931 (holo: BRI [AQ830232]; isotypes: CNS; BO, CANB, L, MO, NOU, NSW n.v., distribuendi ex CNS).

Plants are tussocks with condensed ascending rhizomes, each tussock usually comprising few to numerous tufts. Each tuft is 5-15 mm in diameter at the base with leaves arranged distichously or irregularly. Leaves glabrous and usually upright. Leaf sheath margins at first membranous or cartilaginous, fraying into strips or fibres up to 10 cm long, white to light brown. Leaf blades flattened or curved adaxially, slightly to strongly convex abaxially, or inrolled especially for the lower part; with up to 25 parallel veins on both sides; the margins smooth or scabrid; leaf apex obtuse, near rounded, minutely toothed, or occasionally acuminate. Leaves of male plants 28—70 cm long, (1-)2—4(—-5) mm wide. Leaves of female plants 33—52 cm long, (2—)3—

Austrobaileya 13: 1—6 (2023)

5 mm wide. Male and female inflorescences dissimilar in appearance. Male inflorescence a raceme, unbranched or bearing a few slender branches, each branch 3—9 cm long at the base or near the base of rachis; the scape flattened, smooth, several vertical parallel veins on both sides obvious, especially on dried specimens, 9-32 cm long, (0.6—)1.5—2.5 mm broad, white to light brown; the rachis slightly flat to terete, smooth or rarely slightly verrucose, irregularly vertically angled or channelled, (6—)11-23 cm long, 0.5—2 mm broad, white to light brown; inflorescence and its branches usually subtended by a bract, long- to short-deltoid, 2.5—10 mm long, 0.5—1 mm wide at the base (that 1s usually the widest point), white to light brown, membranous except the mid vein. Male flowers (20—)40- 80(—120), solitary, alternate or sometimes appearing paired or a few grouped together, rarely whorled; each flower subtended by bract, 1.5—2(—4) mm long, 0.5—1 mm wide, usually completely encircling the pedicel; flowers borne perpendicularly to rachis, pedicels terete, (2—)3—4(—7) mm long, 0.1—0.2 mm wide, pale, flowers usually of similar age within each inflorescence; buds ellipsoid to globular, becoming hemispherical or flattened-globose at anthesis, 1.5—2 mm long, 2.5—3 mm wide at open end. Male perianth segments 6 with distinct outer and inner whorls; outer tepals (Sepals) 3, long- to short- deltoid, free except on the very base, uniform in size, texture thick with thin whitish margins, 1.6—1.8 mm long, 0.6—0.8 mm wide, ereen to light green; inner tepals (petals) 3, broadly elliptical, free except on the basal c. 1/5, uniform in size, texture thick with slightly thin margins, 1.5—2 mm long, I-1.5 mm wide, light yellow to light brown except for whitish colour on the margins. Stamens 6, 3 adnate basally to swollen inner surfaces of inner tepals, 2 adnate basally to swollen base of inner tepals, | adnate basally to swollen base of outer tepal; filaments lacking; anthers all similar, versatile, 0.4—0.7 mm long and 0.25—0.5 mm wide, bright yellow to occasionally creamy yellow; pistillode in the centre very rarely developed, mostly whitish; stigma and style hardly formed. Female inflorescence simple, (6—)14—-30 cm _ long,

Wang, Lomandra grayi

rarely with a few branches, each up to | cm long at the base or near the base of rachis; the scape flattened, smooth, often with several vertical parallel veins on both sides, obvious on dried specimens, creamy to light brown, 10—22 cm long, 1.4—2 mm broad; the rachis irregularly angled, smooth or rarely minutely verrucate, 1.5-3.5 cm long; inflorescence bracts 0—2, long- to short-deltoid, up to 5 mm long and 4 mm wide at the base (the widest point); each flower has a bract resembling the general bracts, but can also be broadly cucullate, 1.5—4 mm long and 1-3 mm wide, white to light brown, membranous, 1/2—4/5 encircling the pedicel. Female flowers 10— 25, alternate, single, appearing crowded; borne in clusters on rachis; pedicels terete, or irregularly angled and channelled, 0.5—0.6 mm long, 0.4—0.5 mm wide, creamy to pale brown, flowers usually of similar age within each inflorescence; buds globular, becoming cup-shaped with age, pale cream to greenish. Female perianth segments 6 with distinct outer and inner whorls; outer tepals (sepals) 3, greenish, ovate, 2.2—2.8 mm long, 1.4—1.8 wide, connate at the base; inner tepals (petals) 3, pale cream, broadly ovate, 2.6-3.6 mm long, 1.6—2.6 mm wide, connate near base. Staminodes inconspicuous or sometimes absent, when present lacking filaments and bearing vestigial anthers, 3 inserted on the middle part of inner tepals, 3 alternating with them on the middle of lower part of each outer tepal; pistil conspicuous, styles very short and fused with 3 stigmatic lobes; ovary sessile, flattened globular, 0.6—1 mm long, 1.5-18 mm diameter, with 3 locules; ovules | per loculus. Fruiting pedicels |—3 mm long, single, occasionally in groups of 2 or 3 in the lower section. Capsules obovoid, usually 6—7 mm long, 5—7 mm diameter, light brown to brown, each carpel with 1—5 transverse wrinkles; carpel light to dark brown inside; the carpel margins smooth; fruiting perianths with usually 6 tepals, leathery, persistent, outer whorl 2.1-2.3 mm long, 1.8—2 mm wide; inner whorl 2.9-3.2 mm long, 2.6—2.8 mm wide, bracts occasionally present. Seeds 1 per locule, usually two seeds in each fully developed fruit, narrowly ovoid to ovoid, 4.1-4.5 mm long, 2.4—2.6 mm wide, usually

3

2-angled on inner face, rounded on outer face, smooth to slightly wrinkled, translucent in appearance, light brown. Fig. 1.

Additional specimens examined: Queensland. Cook District: Lockerbie, 10 miles [16.6 km] WSW of Somerset, Apr 1948, Brass 18469 (BRI); ibid, Apr 1948, Brass 18502 (BRI); 18 km north of Captain Billy turn, on Cape York Road, Jun 2004, Gray 8924 (BRI, CNS); Mine lease near Skardon River, Apr 2011, Wannan 6065 et al. (BRI); Unigan Reserve, Head of Swamp, Weipa, Feb 1989, O’Reilly 166 (BRI); 8 km from the beach turnoff from Bolt Head on the track to Bromley Station, Jul 1990, Clarkson 8849 (BRI); Moreton Telegraph Station campground western side, May 2013, Thompson ST13231 (BRI); W side of Wenlock crossing on Frenchman’s Track, May 2013, Thompson ST13299, Nobel & Nelson (BRI); 7.3 km (by road) W of Tozer Gap, 92 km (by road) NE of Peninsula Development Road, Jun 1993, Conn 3795 & Doust (BRI, NSW); 12 km N of the Pennefather Road turnoff on the Weipa — Mapoon Road and 50 m to the E of the road, CYP, Jun 2017, Addicott EPA3824 (BRI); Olive River Environmental Reserve, 8 km E by road of ‘Bromley’ Homestead, 48 km E of Moreton Telegraph Station, CYP, Jun 2007, Forster PIF32827 & McDonald (BRI, NSW); Browns Creek area, 18 km E of Pascoe River crossing of Portland Road toward Lockhart River, Jun 2017, Fell DGFBCIS/14 & Pritchard (BRI); 24.2 km S of Batavia Downs on the Peninsula Development Road, Apr 1990, Clarkson 8322 & Neldner (BRI); MclIlwraith Range, 10.8 km from Peach Creek Crossing, past Forest Pocket, Aug 2002, Gray 8206 & Jones (BRI); NNE of Strathburn airstrip on road to Yarraden, Jun 2005, Wannan 4001 & Beasley (BRI); 11.5 km W of the Laura to Musgrave Road on the track to Dixie, Jun 1989, Clarkson 8127 (BRI); Laura Quinkan country, Sandy Creek about 200 m of junction with Laura River, Jun 2018, McDonald KRM20476 & Gueu Yalangi Rangers (BRI); Midslope of southern fall of Carbine Tableland on Brooklyn Wildlife Sanctuary, c. 5.4 km W of Mt Molloy, Feb 2013, Jensen 2777 & Kemp (BRI); c. 700 m N of high transmission powerline, Bridle Creek area, N of Bare Hill, 18.5 km ENE of Mareeba, Feb 2018, Kemp JEK20062 & Jensen (BRI); 1 km SW of Stones Hill, east of Mareeba, Jun 1993, Bean 5695 & Forster (BRI); Springmount Road, c. 10 km W of Walkamin, May 1990, van der Werff 11631 (BRI). NORTH KENNEDY DISTRICT: Department of Primary Industry Experimental Plots, Boomerang Station via Mt Garnet, Mar 1975, Staples 2034 (BRI).

Distribution and habitat: Lomandra grayi 1s endemic to tropical north-east Queensland, primarily on Cape York Peninsula, from the tip of Cape York to Boomerang Station near Mt Garnet (Map 1). It has been recorded in the Cape York Peninsula (CYP), Wet Tropics (WET) and Einasleigh Uplands (EIU) Bioregions (REF 2023). The altitudes of locations range from 18 m to 840 m.

1-6 (2023)

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Fig. 1. Lomandra grayi.

Wang, Lomandra grayi

Lomandra grayi grows with a diverse suite of grass and herbaceous species. Its habitat 1s mainly open forest or woodland but also occurs on rainforest margins on a variety of substrates including sandy loam, clay loam and ironstone gravel. The dominant tree species may include, in varying compositions: Eucalyptus crebra F.Muell., £. drepanophylla F.Muell. ex Benth., £. platyphylla F.Muell., E. portuensis K.D.Hill, £&. tetrodonta F.Muell., Corymbia_ citriodora (Hook.) K.D.Hill & L.A.S.Johnson subsp. citriodora, C. clarksoniana (D.J.Carr & S.G.M.Carr) K.D.Hill & L.A.S.Johnson, C. confertiflora (F.Muell.) K.D.Hill & L.A.S.Johnson, C. hylandii (D.J.Carr & S.G.M.Carr) K.D.Huill & L.A.S.Johnson, C. nesophila (Blakely) K.D.Hul & L.A.S.Johnson, Lophostemon confertus (R.Br.) Peter G.Wilson & J.T.Waterh., Erythrophleum — chlorostachys — (¥.Muell.) Baill., Canarium australianum F.Muell. var. australianum, Melaleuca viridiflora Sol. ex Gaertn. var. viridiflora and Banksia dentata Lf.

Phenology: Based on herbarium collections of Lomandra grayi, male flowering was recorded mainly from April to July. However, its flowering can be as early as in February. Female flowering was recorded in June and July. Mature fruits were collected mostly during June to August with one record in March.

Affinities: Lomandra grayi 1s _ readily distinguished by its male flowers arranged perpendicularly to the inflorescence rachis. Although no species can be easily confused with Lomandra grayi, it is morphologically similar to L. filiformis which 1s widespread in eastern Australia, from northern Queensland, through New South Wales to Victoria. The two species share similar characteristics: such as tussock forming from condensed ascending rhizomes, the size of leaves and shape of leaf tips. Lomandra grayi can be easily distinguished from L. filiformis by the dissimilar appearance of male and female inflorescences (male and female inflorescences are similar for L. filiformis), the longer male inflorescence (20—55 cm for L. grayi, 2-15 cm for L. filiformis), longer

%

female scapes (10-22 cm for L. grayi, 1-5 cm for L. filiformis), longer pedicel of male flowers (3-4 mm for L. grayi, 0.5—2 mm for L. filiformis), and male flowers vertically arranged along rachis.

Conservation status: Lomandra — grayi is widespread in north-east Queensland. Currently there are many locations known stretching along its geographic range. It can be a very common species where it occurs. It is recorded from Batavia NP (CYPAL), Kulla (MclIlwraith Range) NP (CY PAL) and Dinden NP. The species is not considered to be threatened in the wild and a Least Concern conservation status is recommended using the IUCN (2019) criteria.

Etymology: This specific epithet honours Bruce Gray. Bruce has been a keen botanical collector for many years, being previously employed as a technical officer with CSIRO at the Australian National Herbarium (QRS) in Atherton. Bruce has collected more than 10,000 plant specimens for Australian herbaria.

Acknowledgements

I am grateful to the following staff at the Queensland Herbartum who helped in the preparation of this manuscript: Nicole Crosswell for the illustrations; Jiaorong Li for the distribution map; Peter Bostock for his Latin expertise; Dr Paul Forster for constructive comments. I also wish to thank Dr Mats Hjertson, Curator (vascular plants), Uppsala University, Sweden for the type specimen photos of Lomandra_filiformis at UPS. Sincere thanks to the Directors of CANB, CNS, DNA, JCT, MEL, NSW and NT for providing loan specimens.

References

Briccs, B.G. (1986). Chromosome numbers in Lomandra (Dasypogonaceae). Telopea 2: 741— 744,

CHAH [COUNCIL OF HEADS OF AUSTRALASIAN HERBARIA| (2022). Australian Plant Census. http:// biodiversity.org.au/nsl/servicers/apc, accessed 10 February 2022.

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CHASE, M.W. & STEVENS, P.F. (1998). An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Garden 85: 531-553.

CHASE, M., DUVAL, M.H., HILLs, H.G., CONRAN, J.G., Cox, A.V., EGUIARTE, L.E., HARTWELL, J., Fay, M.F., CADDICK, L., CAMERON, K. & Hoot, S. (1995). Molecular phylogenetics of Lillanae. In P.J. Rudall et al. (eds.), Monocotyledons: Systematics and Evolution, pp. 109-137. Royal Botanic Gardens, Kew: London.

CHASE, M.W., REVEAL, J.L. & Fay, M.F. (2009). A subfamilial classification for the expanded Asparagaceae families |§ Amaryllidaceae, Asparagaceae and Xanthorrhoeaceae. Botanical Journal of the Linnean Society 161: 132-136.

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, J.D. & BircuH, J.L. (2020). Evolution of Lomandroideae: Multiple origins of polyploidy and biome occupancy

in Australia. Molecular Phylogenetics and Evolution 149 (106836): 1—16.

IucN [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.

Leg, A.T. (1966). Xanthorrhoeaceae. Contributions from the New South Wales National Herbarium, Flora Series 34: 16—42.

Leg, A.T. & MACFARLANE, T.D. (1986). Lomandra. In A.S. George (ed.), Flora of Australia 46: 100-141. Australian Government Publishing Service: Canberra.

REF [REGIONAL ECOSYSTEM FRAMEWORK] (2023). https:// www.qld.gov.au/environment/plants-animals/ plants/ecosystems/descriptions/framework, accessed 17 January 2023.

THIERS, B.M. (updated continuously). /ndex Herbariorum. http://sweetgum.nybg.org/ science/ih/, accessed 26 November 2022.

WANG, J. (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 10: 266— Dien

Austrobaileya 13: 1—6 (2023)

140°E 142°E 144°E 146°E

140°E 142°E 144°E 146°E

Map 1. Distribution of Lomandra grayi based on collections in Australian herbaria. Abbreviations for Queensland Bioregions: CYP, Cape York Peninsula; EIU, Einasleigh Uplands; GUP, Gulf Plains; WET, Wet Tropics (REF 2023).

Scleria psammitica R.Booth (Cyperaceae), a new Species from Cape York Peninsula, Queensland

R. Booth!

Summary

Booth, R. (2023). Scleria psammitica R.Booth (Cyperaceae), a new species from Cape York Peninsula, Queensland. Austrobaileya 13: 7-13. A new species, Scleria psammitica R.Booth, is described and illustrated. Notes are provided on its distribution and habitat. A key to Queensland species of Diplacrum R.Br. and Scleria P.J.Bergius 1s provided.

Key Words: Cyperaceae; Diplacrum; Scleria; Scleria psammitica; flora of Australia; flora of Queensland; new species; taxonomy; identification key

‘Queensland Herbarium and Biodiversity Science, Department of Environment and Science, Brisbane Botanic Gardens Mt Coot-tha, Mt Coot-tha Rd, Toowong, Queensland 4066, Australia.

Email: Ronald.Booth@des.qld.gov.au

Introduction

Cyperaceae is a cosmopolitan family which includes 95 genera (Larridon ef al. 2021). Scleria P.J.Bergius (Bergius 1765: 142) is the sixth largest genus in the family, with about 260 species (Govaerts et al. 2015; Bauters ef al, 2016). The genus 1s pantropical, consisting of 23 species in Australia, with c. four being endemic. The genus has received relatively little attention in Australia, with all formally recognised species having been named over 100 years ago.

The genus Diplacrum R.Br. was described to accommodate D. caricinum R.Br., a species where the glumes enclose the achene (resembling a sort of perigynium) (Brown 1810: 241). Brown compared Diplacrum to both Scleria and Carex, the latter based on an assumption that the glumes of D. caricinum were homologous’ with _ the perigynium or utricle in Carex species. Kunth (1835) subsequently argued that the glumes in Diplacrum and the utricle in Carex L. are not homologous and Bentham (1878) subsequently merged Diplacrum with Scleria. However, Simpson & Koyama (1998) and Govaerts & Simpson (2007) reinstated Robert Brown’s Diplacrum as a genus. This has been corroborated based on

chloroplast (ndhF, rps16) and nuclear (ITS) DNA markers (Bauters ef a/. 2016). In the current classification of Cyperaceae based on an extensive nuclear genomic dataset the

two genera are now placed in separate tribes (Larridon et al. 2021).

Amongst the Australian species of Scleria, only S. sphacelata F.Muell. is dioecious. After studying the collections of S. sphacelata at the Queensland Herbarium (BRI), several of the specimens, although superficially resembling S. sphacelata, were noted to be monoecious. All these specimens were collected in the sandstone hills surrounding Laura on Cape York Peninsula, north Queensland. After further study of collections of Scleria from the Same area, several were also found identified as S. brownii Kunth. Preliminary descriptions for this species as Scleria sp. (Laura N.Byrnes 3285), were included in a DELTA key (Jessup et al. 2005 onwards) and the phrase name has been listed 1n the ‘Census of the Queensland Flora (Booth 2007, 2010, 2021).

Since then, more collections of this plant have become available for detailed taxonomic study. Critical examination of these by the author has now enabled this new species to be formally described and named as

Accepted for publication 17 January 2023, published online | March 2023

© Queensland Herbarium 2023. ISSN 2653-0139 (Online)

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Scleria psammitica R.Booth. A key to the Queensland species of Scleria 1s provided. It includes the two species of Diplacrum that occur 1n Queensland due to their superficial morphological similarity and likelihood of confusion with species of Scleria.

Materials and methods

All herbartum specimens of Sc/eria held at BRI have been examined. Measurements were made from dried material. The description 1s modelled on those in Jessup ef al. (2005 onwards). Specimens were viewed with a Nikon SMZ25 microscope and images were captured using NIS Elements Software with Z-stacking features (https://www. microscope.healthcare.nikon.com/products/ software/nis-elements).

Common abbreviations in the specimen citations are Mt (Mountain, except where part of a National Park or State Forest name) and NP (National Park).

Taxonomy Scleria psammitica R.Booth, sp. nov.

Similar to Scleria sphacelata F.Muell. but differs in being monoecious versus dioecious; the glumes being appressed pubescent to scaberulous versus scabrid to glabrous, and yellow-brown to pale brown versus dark brown with reddish margins. Type: Queensland. Cook DIsTRIcT: Normanby Holding, Trap Creek catchment, S of Battle Camp Road, 21 June 2007, PJ. Forster PIF32856 & K.R. McDonald (holo: BRI [AQ0737233 comprising | sheet]; iso: CANB, NSW).

Scleria sp. (Laura N.Byrnes 3285): Booth (2007, 2010, 2021).

Scleria sp. Laura (N.Byrnes 3285) Qld Herbarium: CHAH (2012).

Robust perennial with a stout, woody rhizome; monoecious. Culms trigonous to terete, bases swollen, 30—65 cm tall, 1.5—2.5 mm wide, scabrous or papillose. Leaves: sheath scabrous, not winged; contra-ligule truncate,

Austrobaileya 13: 7-13 (2023)

ciliate; lamina 25—65 cm long, 1.75-3.5 mm wide, glabrous, or scabrous, apex acute. Inflorescence simple or once compound, panicle-like, 3—7-branched, 15-25 cm long, 2.5—4 cm wide. Involucral bracts leaf-like, lowest bract up to 25 cm long. Flowers unisexual. Spikelets sessile, or sub-sessile, 4—6 mm long, 2—2.8 mm wide. Glumes 3-5 per spikelet, broadly ovate, coriaceous, 2.5—4 mm long, apex acute to mucronate, surface appressed pubescent to scaberulous, yellow- brown to pale brown. Male flowers: stamens 3, anthers linear, 2.7-3 mm long. Female flowers: stigmas 3. Achene 3-sided or terete, wrinkled to rugulose, 2.5—3 mm long, 2—2.5 mm wide, white, with ferruginous hairs, rarely glabrous, apex rounded or slightly apiculate; disc brown, 3-lobed, shallow, lobes obtuse, undulate. Figs. 1-3.

Additional selected specimens examined: Queensland. Cook District: NE corner of Piccaninny Wildlife Sanctuary, c. 35.6 km from the homestead, Apr 2014, Jensen RJ31IS2 (BRI); Chuula Outstation, Kaanju Nation, Central Cape York, May 2005, Smith 4866 & Claudie (BRI); Orchid Creek Station, Wallaby Range; SW of Lockhart River, Cape York Peninsula, Apr 2014, Forster PIF40910 et al. (BRI); Bacon Creek, 15 km past Archer River, Apr 1988, Forster PIF4039 (BRI); 18 km S of Coen on the Peninsula Development Road and just E of a borrow pit east of the road, Cape York Peninsula, Jun 2017, Addicott EPA3S809 & Newton (BRI); Crystal Vale Station, 16 km S of homestead, 38.7 km SW of Coen, Jun 2018, Leitch & Starkey ODAOISI61 (BRI); Mary Valley, S of the Peninsula Developmental Road, Apr 2009, Wannan 5642 (BRI); Melsonby NP, N of Battle Camp Road; NW of Cooktown, May 2010, Forster PIF36773 & Thomas (BRI); Mt Jack Station, Cape York Peninsula, Oct 2009, Wannan 5756 & Thompson (BRI); 5 km ENE of disused airstrip, Mt Jack Station, Cape York, Mar 1995, Garnett STG/1228 (BRI); Bridge Creek Holding (proposed NP), N of Battle Camp Road; NW of Cooktown, May 2010, Forster PIF36730 & Thomas (BRI); Escort Creek Holding, ESE of Laura, Mar 2017, Forster PIF44649 et al. (BRI); Cormay Creek catchment, East Quinkan, SE of Laura, Mar 2017, McDonald KRMI18997 et al. (BRI); Laura River on Crocodile Station, May 1975, Byrnes 3285 (BRI); On track to Giant Horse art site, SE of Laura, Jul 1990, Bean 1890 (BRI); adjacent to Split Rock Gallery, 14 km S of Laura, Apr 1992, Neldner 3696 & Clarkson (BRI); Turtle Rock area, Laura sandstone escarpment, Jan 1993, Forster PIF 12860 & Bean (BRI); Amphitheatre on the NE side of Ngarrabullgan, May 2002, Fox s.n. (BRI [AQ764125]); Mt Mulligan, c. 40 km NW of Dimbulah, Apr 1985, Clarkson 5766 (BRI).

Booth, Scleria psammitica 9

Fig. 1. Scleria psammitica. A. habit showing the base of the plant and rhizome. B. inflorescence. C. spikelet with achene in situ. D. achene. All from Forster PIF32856 & McDonald (holotype, BRI). Scales as indicated. Del. N. Crosswell.

10

Fig. 2. Scleria psammitica. achene. From Forster PIF 32856 & McDonald (BRI, holotype).

Fig. 3. Scleria psammitica. spikelet with in situ achene. From Forster PIF32856 & McDonald (BRI, holotype).

Austrobaileya 13: 7-13 (2023)

Distribution and habitat: Scleria psammitica is endemic to Cape York Peninsula in far northern Queensland between Mt Mulligan in the south and as far north as the Lockhart River (Map 1). The species invariably occurs in eucalypt-dominated woodland on sandstone hills, often restricted to crevices in outcropping rocks.

Notes: Scleria psammitica is morphologically similar to S. sphacelata, differing most obviously in being monoecious’ versus dioecious, the glumes being appressed pubescent versus scabrid to glabrous and brown to pale brown versus dark brown with reddish margins. It 1s also somewhat similar in appearance to S. brownii but differs in its larger more robust form, 30—65 cm tall versus 20—60 cm tall, culms 1.5-2.5 mm wide versus 1—1.5 mm wide and the pubescent or scaberulous glumes, versus glabrous glumes.

Conservation status: Although Scleria psammitica 1s apparently limited to sandstone hills, it 1s quite widespread within its geographic range and is not considered threatened.

Etymology: The epithet is derived from the Greek psammités (made from sand) and refers to the geological strata where the plant is commonly found.

Key to the Queensland species of Diplacrum and Scleria (nomenclature as per Booth 2021).

1. Glumes with long patent hairs, c. | mm long; inflorescence consisting of globose clusters of 1 to many spikelets ona long peduncle ........ S. carphiformis 1. Glumes glabrous or with short hispid hairs < 0.5 mm long, without

LOA ANGIE MAINS ES Oo thatch oad fy Be, Ge all a oetivel: cad by By See Gh oes, Sates o ce Me oe 2 2 Small annuals (or short-lived perennials); rhizomes absent or not well developed .... . 3 2, Perennials; rhizome well-developed: -o4.4 64 oa 4 bed ame ee bee ee ed 10 Dwarf plants usually 1—10(—20) cm high; mature achene hidden by glumes and not readily visible (Diplacrum) .............. 0.2.0.0 0 eee eees él 3. Plants (S—)15—60 cm high; mature achene not completely hidden by CUES SAOSIN PV ISTOLO™ 5, wc Pa de sae. Fact ad wm ne Be he tte tt ae a one tt Sa’ © Ss ds een 5 4 Glume enclosing the achene prominently 3 or more nerved, apex 3-lobed . . D. caricinum

4. Glume enclosing the achene | or 2-nerved, apex acuminate, not lobed . . . .D. pygmaeum

Booth, Scleria psammitica 1]

5 Inflorescence spike-like, not branched; spikelet clusters sessile, subtended

by bracts: notexceedino the spikelets... . 2: c.f yes ar be we ce yk SES S. pergracilis 5. Inflorescence paniculate, subtended by bracts usually far exceeding the HAMIOPESCOMCGr 5 cic. piel Ge WSs opldn SHOE RG Fle oho ode PES BORE MSL ced pee 6

6 Plants frequently very hairy; disc of achene with many small bumps

and protrusions; peduncles of the inflorescence clusters stout, often

WHTSCG Fe peach, eet eee eee ae Wel ered iveta SRAGEN Me We alfe MOF athe Wt nae SG ee GN tee S. rugosa 6. Plants glabrous or only slightly hairy; disc of achene smooth, with no

protrusions; peduncles of the inflorescence clusters slender, not

WAT OCS By ce cx wen was amass areca BE pyneds nic mde prepay ot ay RE AS Sled, a teens san oo ones EM cg, ee ese wen span oe 22 os © 7 7 ~chene pubescent, beak purplish 5 aq ecud xe eo lls ce Style ee Web® gens S. biflora 7m. Achene clabrons,bealk not-purplishy. A occ. He De Whee eteaee ste Hoe Ee © ae seat ee EG 8 Disc lobes with short, erect, subulate tips .................6. S. tricuspidata S$; (DiISGlObeS TORSUDUIATS TPC! RE igs: wc aw ge Se a id eee Vi Ye yk coe Figs G 9 9. Achene globose, with irregular elongated pits, a smooth triangular area

immediately Above each CiselObe: © .. «ional & SB: de sew we ee he Ee ae S. laxa 9. Achene oblong-ellipsoid, smooth. ..................4 S. novae-hollandiae 10 Middle leaves of the fertile culms clustered in pseudo-whorls of 3-5... . 2... 2... 11 10. Middle leaves not in pseudo-whorls of 3—5, evenly spaced along the stem ........ 13 11 Disc of achene cup-like, covering at least the lower half of the achene . . . S. Sumatrensis 11. Disc of achene shallowly 3-lobed, covering < 1/3 of the achene.............. 12

12 Panicle branches at a narrow angle to the main axis; achene slightly rugulose to smooth, 2—2.5 mm long, hardly beaked, often tinged

POLIT IS 2A Pye x Sea ewdelt liere 9 el Bed eee eal nye WSs Ge Gaertn aA Pte ce S. polycarpa 12. Panicle branches widely spreading; achene pitted, with many shallow,

rounded depressions, 2.5—3 mm long, beaked, white ........... S. scrobiculata PS Aceair shed ie-wiieeds 5) 5. h gee 8 tae cody BY seh: so ois cee Peed eee oe ee ee BS es geen ce 14 TS ear sheatissnotsvinn@ede = SoS Fe os ase yo ie BARBS SS ace ae ne oe tet Be RAR AS oe es ge ten 18

14 Inflorescence a terminal, compact, much branched panicle; lowest

involucral bract < 3 cm long, much shorter than the inflorescence ...... S. poiformis 14. Inflorescence terminal and with 1-5 lateral panicles some distance from

each other; bracts leafy, > 3.5 cm long, as long or longer than the

INMOTESCENCE “. ice en dw en ep Pace bk wath ee a £ See ate hs 15

15 Plant stoloniferous, stolons covered with purplish, lanceolate scales;

achene Simin long nor Peaked, +, +, at + ao aw sore ow Oe ek S. psilorrhiza 15. Plant shortly rhizomatous; achene <3 mm long, beaked.................. 16 16 Disc deeply 3-lobed, lobes very acute, frequently bidentate at the tip;

ACCME -SIMOOCM. 5, Sey SPIE, LE UE ae Be se oe te Be ee Be, wen Fe we os ats SP S. levis 16. Disc lobes rounded, obtuse; achene frequently cancellate (pitted)... ........2.. 17

17 Contraligule 1-2 cm long, a scarious, oblong, purplish strap-like appendage; prominent, stiff bracteoles below each spikelet cluster,....... S. ciliaris 17. Contraligule <1 cm long, with brown scarious margin; bracteoles indistinct . .S. terrestris

12 Austrobaileya 13: 7-13 (2023)

18 Inflorescence a_ terminal, compact much _ branched panicle,

subtended by small, glume like bracts < 12 mm _ long,

MUCH-SHOTbert Than TheAniorescenee “sas Abs ve wa we EN ow oe ee 8 eS S. poiformis 18. Inflorescence terminal and with 1-5 lateral panicles some distance from

each other: bracts leafy, > 35 mm long, as long or longer than the

INTIOTESCENCE: .. 6... .: «oe ee. Sev SSE a G: eeke bee Sey ASE, DPE wee ete ee eS 19 19 Disc obsolete, or reduced to a narrow band at the base of the achene ............ 20 19, Dise Swell: aeveloped. 5. -% fei See & Pas wi Se twee en b Behe tw 2h 21 20 Culms 60—200 cm tall, 3—7 mm wide; leaf lamina 7—25 mm wide ...... S. corymbosa 20.Culms 30-60 cm tall, 1.42 mm wide; leaf lamina 1-4 mm

WIC ae yon Medic tin ge 2 tals apeeeaan a 4 Wake Rye eeaed S. lithosperma var. linearis 21 Inflorescence solely with stamens, or stigmas with fruit, but not both... . . S. sphacelata 21. Inflorescence with both stamens and stigmas ............ 0.00002 2 eee 22

22 Plant with the culms separated along a thick, robust rhizome; achene with curly ferruginous hairs, glumes tomentose; found on Cape York

Penimnsilacin Saivdstoie His: ob a. a a ee et be end me S. psammitica 22.Culms clumped, stoloniferous; achene glabrous or slightly hispid; not LOUIE Th SANCSTOMENIUS 66. yew cesta a 8 fa Joke eet wn eM ween CAME 9 ta ite 23

23 Large plants 1—2 m tall, culms 2—10 mm wide; inflorescence paniculate . . . .S. terrestris 23. Smaller plants 5—60 cm tall, culms 0.8—1.5 mm wide; inflorescence of

2S Taseicles. on Alone peduncle: 8552 Abas wat te SM oe ee ee ANE oe. 24

24 Achene with a long cylindrical beak; contraligule tongue shaped up U7 TE CIO? oy. oP oe gy BR a ge Ns oe eS aa A Se a ene 24. Achene with a brown tip, no beak; contraligule absent or truncate and less chat O, Sai Oe: =. So ec cet Bae en & Ree a a BS ee Bn hte thee S. brownil

S. mackaviensis

The last two species are very closely allied and in North Queensland are not easy to differentiate. Scleria brownii occurs as far south as Port Curtis and only S. mackaviensis Boeckeler occurs 1n both south-east and north Queensland.

Acknowledgements

I thank Nicole Crosswell for the illustrations and Tony Bean for his support and helpful Suggestions 1n compiling this manuscript.

References

BAUTERS, K., ASSELMAN, P., SIMPSON, D.A., MUASYA, A.M., GOETGHEBEUR, P. & LARRIDON, I. (2016). Phylogenetics, ancestral state reconstruction, and a new infrageneric classification of Scleria (Cyperaceae) based on three DNA markers. Taxon 65: 444—466.

BENTHAM, G. (1878). Flora Australiensis: A Description of the Plants of the Australian Territory. 7: 246— 449. L. Reeve & Co.: London.

Beratius, P.J. (1765). Scleria, et nytt Orte-genus ifran America, framlagt och beskrivet. Kongliga Vetenskaps Academiens Handlingar 26: 142— 148.

BooTtH, R. (2007). Cyperaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2007, p. 63. Queensland Herbarium, Environmental Protection Agency: Brisbane.

—— (2010). Cyperaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2010, p. 58. Queensland Herbarium, Department of Environment & Resource Management: Brisbane.

— (2021). Cyperaceae. In G.K. Brown (ed.), Census of the Queensland Flora 2020. Queensland Department of Environment and Science, Queensland Government. https://www.data.

qld.gov.au/ dataset/census-of-the-queensland- flora-2020, accessed 19 May 2021.

Brown, R. (1810). Prodromus Florae Novae Hollandiae et Insulae Van-Diemen. R. Taylor et soc.: London.

Booth, Scleria psammitica

CHAH [COUNCIL OF HEADS OF AUSTRALIAN HERBARIA]. (2012). Australian Plant Census. https:// id.biodiversity.org.au/instance/apni/771113

GOVAERTS, R. & SIMPSON, D.A. (2007). World Checklist of Cyperaceae. Sedges. Kew Publishing: Royal Botanic Gardens, Kew.

GOVAERTS, R., SIMPSON, D.A., GOETGHEBEUR, P., WILSON, K.L., EGorova, T. & BRUHL, J.J. (2015). World checklist of Selected Plant Families. Cyperaceae. The Board of Trustees of the Royal Botanic Gardens: Kew.

JESSUP, L.W., SHARPE, P.R. & BOOTH. R. (2005 onwards). Cyperaceae in Queensland: Descriptions, Illustrations, Identification, and Information Retrieval. Version: 31 March 2015. Queensland Herbarium: Brisbane.

Kuntu, K.S. (1835). Uber die natur des schlauchartigen organs utriculus, welches in der gattung Carex das pistill und spater die frucht einhiillt. Wiegmann’s Archiv fiir Naturgeschichte 1(2): 349-353, t. 6.

1135

he ee 7 ee ee ~ > = * * “* *%

en "Wis i ee ee ee a i ee Oe

Map 1. Distribution of Scleria psammitica.

. ‘=

13

LARRIDON, I[., ZUNTINI, A.R., LEVEILLE-BOURRET, E., BARRETT, R.L., STARR, J.R., Muasya, A.M., VILLAVERDE, T., BAUTERS, K., BREWER, G.E., BRuUHL, J.J.. Costa, S.M., ELtiottT, T.L., EPITAWALAGE, N., ESCUDERO, M., FAIRLIE, I., GOETGHEBEUR, P., Hipp, A.L., JIMENEZ- Mesias, P., SABINO Kikucul, I.A.B., LUCENO, M., MARQUEZ-CorRRO, J.I., MARTIN-BRAVO, S., MAURIN, O., POKORNY, L., ROALSON, E.H., SEMMOURI, I., SIMPSON, D.A., SPALINK, D., THOMAS, W.W., WILSON, K.L., XANTHOS, M., Forest, F. & BAKER, W.J. (2021). A new classification of Cyperaceae (Poales) supported by phylogenomic data. Journal of Systematics and Evolution 59: 852—895.

SIMPSON D. & KoyAMA, T. (1998). Cyperaceae. In Flora of Thailand 6(4): 426—447. Forest Herbarium, Royal Forest Dept.: Bangkok.

=

Austrobaileya 13: 14—15 (2023) 14 SHORT COMMUNICATION Reinstatement and lectotypification of the name Cissus muelleri Planch. (Vitaceae) Betsy R. Jackes

College of Science and Engineering, James Cook University, Townsville, Queensland, Australia.

Email: betsy.jackes@jcu.edu.au

Key Words: Vitaceae; Cissus cardiophylla; Cissus muelleri; flora of Australia; flora of Queensland;

taxonomy; lectotype

The specific epithet ‘cardiophylla’ has been applied to two different species of Cissus L. (PNI 2023). Standley (1929) described C. biformifolia Standl., from Panama, and C. cardiophylla Standl. from Costa Rica. In the Flora of Costa Rica, Standley (1937) considered these two species to be synonymous and placed C. cardiophylla Standl. into synonymy under C. biformifolia Standl. Jackes (1988) transferred the Australian species Vitis cardiophylla F.Muell. to Cissus as C. cardiophylla (F.Muell.) Jackes, but this name 1s an illegitimate later homonym of C. cardiophylla Standl. According to the ICN rules (Turland ef al. 2018), another name is required for the Australian species. Originally the intention was to rename the species with a new epithet. However, the name Cissus muelleri Planch. 1s available for this species, having been previously included in the synonymy of C. cardiophylla (F.Muell.) Jackes (Jackes 1988).

Taxonomy

Cissus muelleri Planch., Monogr. Phan. [A.DC. & C.DC.] 5(2): 516 (1887). Type citation: “Australie tropicale: Queensland, district de Kennedy (Mueller, herb. Melb. sous le nom de Vitis cordata). Hodjkinson river (Gulliv., n 204, in herb. Melb.). Rockingham bay (Dallachy in herb. Melburn).’ Type: Queensland. NorTH KENNEDY DISTRICT: Basalt Wall Fletchers C[ree]k, s.dat., R. Daintree s.n. (ecto [here designated]: MEL 565994).

Vitis cardiophylla F.Muell., Fragm. 2: 73 (1861); Cissus cardiophylla (F.Muell.) Jackes, Austrobaileya 2: 491 (1988), nom. illeg. non C’. cardiophylla Standl., syn. nov. Type: Jn nemoribus ad oppidum, Rockhampton, s.dat., Thozet [94] (holo: MEL 540180), labelled as Vitis cordata.

Vitis cordata Benth., Fil. Austral. 1: 447 (1863), pro parte taxonomic synonym.

Cissus repens auct., non Lam., Domain, Biblioth. Bot. 22(89): 921 (1927).

Typification: Of the three original syntypes for Cissus muelleri, only the collection by R. Daintree has been located. This collection was referred to by Planchon as “Mueller, herb. Melb.”, but was actually collected by R. Daintree as indicated by the label “Vitis, Kennedy District, R. Daintree” on MEL 565994. There is an additional blue label attached to the specimen which indicates it was collected on the “Basalt Wall, Fletcher’s Creek, North Kennedy”. A second sheet (MEL 565993) of what is apparently the same collection has had some of this information added with later labels. Although I have not seen a specimen labelled as Gulliver 204 from the Hodjkinson River, Gulliver 804 from the Hodjkinson (Hodgkinson) River (MEL 565093) 1s C. muelleri; 1t 1s possible that the hand-written number was incorrectly transcribed by Planchon. The third syntype (a collection by Dallachy) has not been located to date.

Accepted for publication 14 March 2023, published online 5 April 2023

© Queensland Herbarium 2023. ISSN 2653-0139 (Online)

15

Acknowledgments

My sincere thanks to Alison Vaughan (MEL) and Frank Zich (CNS) for assistance in locating some of the Cissus muelleri collections, and Paul Forster (BRI) for nomenclatural advice.

References

IPNI (2023). International Plant Names Index. The Royal Botanic Gardens, Kew, Harvard University Herbaria & Libraries and Australian National Herbarium. http://www.ipni.org, accessed 6 January 2023.

JACKES, B.R. (1988). Revision of the Australian Vitaceae, 3. Cissus. L. Austrobaileya 2: 481-505.

STANDLEY, P.C. (1929). Studies of American Plants I. Field Museum of Natural History, Botanical Series 4(8): 225-226.

— (1937). Flora of Costa Rica. Field Museum of Natural History, Botanical Series 18(2): 653.

TURLAND, N.J., WIERSEMA, J.H., BARRIE, F.R., GREUTER, W., HAwWkKSworRTH, D.L., HERENDEEM, P.S., KNAPP, S., KUSBER, W.-H., Lt, D.-Z., MARHOLD, K., May, T.W., McNeILy, J., Monro, A.M., PRADO, J., PRicE, M.J. & SmiITH, 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, Glashiitten: Koeltz Botanical Books.

Cissus montana (Lauterb.) Jackes & Trias-Blasi, a newly recognised species of Vitaceae from New Guinea

Betsy R. Jackes'> & Anna Trias-Blasi’

Summary

Jackes, B.R. & Trias-Blasi, A. (2023). Cissus montana (Lauterb.) Jackes & Trias-Blasi, a newly recognised species of Vitaceae from New Guinea. Austrobaileya 13: 16-19. Cissus montana (Lauterb.) Jackes & Trias-Blasi comb. & stat. nov. from New Guinea was initially described by Lauterbach in 1925 as C. adnata var. montana Lauterb. A comparison of herbarium material indicated that this variety was distinct from C. adnata Roxb. and should be recognised at species rank. The newly recognised species is fully described with notes provided on distribution, habitat, phenology and typification. It is easily distinguished by a carpet-like indumentum of rusty-brown 2-armed hairs on the abaxial leaf surface. A lectotype is selected for C. adnata var. montana.

Key Words: Vitaceae; Cissus; Cissus adnata var. montana; Cissus montana; flora of Indonesia; flora of Papua; flora of Papua New Guinea; new species; new combination and status

'College of Science and Engineering, James Cook University, Townsville, Queensland 4811. Australia; “Royal Botanic gardens, Kew, Richmond, TW9 9AE, United Kingdom; °Corresponding

author. Email: betsy.jackes@jcu.edu.au

Introduction

Cissus L. 1s the largest genus 1n the Vitaceae, with over 350 species occurring throughout tropical and subtropical regions of the world (POWO 2022). Most of the species are in the core Cissus group with several in Cissus I] (Lu et al. 2018). Species referred to Cissus II by Lu et al. (2018) have a neotropical-Australian distribution (Lui et a/. 2013), although the more recent molecular phylogenetic study sampled only a few taxa.

The last overall account of Cissus in New Guinea was nearly 100 years ago (Lauterbach 1925). Revisionary work in adjacent regions such as Australia (Jackes 1988) has meant that many of the New Guinea taxa are relatively well known, although several remain known only from few collections or just type material with continuing uncertainty to their taxonomic status.

An examination of species with similar morphological features indicated that Cissus adnata var. montana Lauterb., should be treated as a separate species, thus requiring

a new combination and change in status to be made. The importance of hair morphology in differentiating between species in the Vitaceae, has been shown by Jackes (1987a, b) and has been critical in the recognition of Cissus montana.

Eighteen species of Cissus are now recognised for New Guinea (up from POWO 2022).

Materials and methods

Herbarium specimens were examined at A, BISH, BRI, E, K and L, plus additional duplicates available on the JSTOR Global Plants website; many of these specimens were duplicates of relatively few collections. Herbarium acronyms follow those of Thiers (continuously updated).

The foliage hairs were examined by Scanning Electron Microscopy (SEM) in the Advanced Analytical Centre at James Cook University. Images of the lectotype and isolectotype specimens are available on

Accepted for publication 14 March 2023, published online 5 April 2023

© Queensland Herbarium 2023. ISSN 2653-0139 (Online)

Jackes & Trias-Blasi, Cissus montana

the JSTOR Global Plants website (https:// plants.jstor.org/stable/viewer/10.5555/al.ap. specimen.k000736439; —__ https://plants.jstor. org/stable/viewer/10.5555/al.ap.specimen. s11-22390). Images of cited specimens at the Herbarium of the Netherlands are available online via their website (https://www. naturalis.nl/en/collection) and are accessible via the given accession numbers.

Taxonomy

Cissus montana (Lauterb.) Jackes & Trias- Blasi, comb. et stat. nov.

Cissus adnata Roxb. var. montana Lauterb., Bot. Jahrb. Syst. 59: 522 (1925). Type citation: [Papua New Guinea.| “Nordostl. Neu-Guinea: Walder des Kani-Gebirges, 1000m (SCHLECHTER n. 16676!, in Knospe 9, Okt. 1907)”. Type: Papua New Guinea. in der WAaldern des Kani-Gebirges [Kani Range], 9 October 1907, R. Schlechter 16676 (lecto [here designated]: S 11-22390; isolecto: K 000736439).

Vine; stems striate, reddish 2-armed hairs common; tendrils bifid, angular, pubescent. Stipules prominent to 4 x 3 mm, pubescent. Leaves simple, firm, petiole 2—6 cm long; lamina ovate to broadly ovate, 6—10 x 3—8 cm, apex acute, base truncate to weakly cordate, margins bristly-serrate; at maturity upper surface sparsely pubescent, usually drying ereyish, lower surface densely tomentose, the rusty-brown 2-armed hairs appearing as though brushed (similar to a shag pile carpet), rather than randomly arranged, arms relatively short, stalk munute; veins raised above the surface. Inflorescence to 2.5 cm long when flowering, 3 cm long when fruiting. Flowers to 2.5 mm long, pedicel to 3 mm long; calyx and corolla densely rusty- pubescent on the outside. Corolla 2 mm long, ovary glabrous. Berry globular, 4-6 mm diameter, black. Seed 1 per fruit; endosperm in transverse section almost twice divided by the thin endotesta. Figs. 1, 2.

Additional specimens examined: Indonesia. PAPUA: Rouffaer [Tariku] River, Aug 1926, Docters van Leeuwen 10084 (A, K, L [L.2333595]); ibid, Nov 1926, Docters van Leeuwen 11095 (A, K, L [L.2334484)]); Nassau Mountains, Oct 1926, Docters van Leewen 10645 (kK, L [L.2345252, L.2345253]). Papua New

17

Guinea. CENTRAL PROVINCE: Track from Mt Kumme, E of Woitape, Jan 1965, van Royen NGF 20358 (BRI, K); Near landing paddock Mt Ehuhu, Mt Lamington area, Nov 1984, Kuduk & Naoni 6198 (L [L.2334433]). New BRITAIN PROVINCE: Gasmata subdivision, Torlu River, Mar 1965, Sayers NGF 24206 (A, BISH, BRI, CANB, E, L [L.2334487]).

Distribution and habitat: Cissus montana is endemic to New Guinea in Indonesian Papua and Papua New Guinea, occurring in rainforest communities along the northern coastline from Jayapura to New Britain, usually above 300 m altitude.

Phenology: Only four of the seven collections of Cissus montana were flowering, the dates of collection indicated that flowering occurs between August and March. The specimen Docters van Leeuwen 10845 was fruiting when collected in October.

Typification: Collections made by Rudolf Schlechter were deposited at the Botanic Garden and Botanical Museum Berlin-Dahlem (B) with duplicates distributed elsewhere. At the time of Lauterbach’s account, it was not specifically stated where the cited specimens were deposited; however, the inference was that they were at that institution. Most type specimens at B were destroyed during an Allied bombing raid during World War Two. Duplicates of the type collection are present at K and S. The collection at S is the more complete and 1s selected here as the lectotype.

Notes: The importance of indumentum for differentiating species of Vitaceae has been previously emphasised (Jackes 1987a,_ b). Cissus montana 1s distinguished from other New Guinea species by the greyish, glaucous adaxial leaf surface and the dense carpet-like indumentum of rusty-red 2-armed hairs on the abaxial surface. These hairs appear as though they are brushed rather than randomly arranged, as found in C. aristata Blume and C.. conchigera Ridley (Fig. 2). The veins on the abaxial surface are raised above the surface and are also covered with rusty- brown 2-armed hairs. This species is readily distinguished from C. adnata Roxb. by the dense covering of hairs, particularly on the abaxial leaf surface, the pubescent stipules, and the calyx and corolla being densely pubescent on the outside.

18 Austrobaileya 13: 16-19 (2023)

100 wu

Fig. 1. Cissus montana. Single 2-armed or T-branched hair. From Sayers NGF 24206 (BRI).

Fig. 2. Cissus montana. Dense carpet-like indumentum of 2-armed or T-branched hairs. From Sayers NGF 24206 (BRI).

Jackes & Trias-Blasi, Cissus montana

Acknowledgments

The authors thank the directors of A, BISH, BRI, E, K and L for access to specimens.

References

JACKES, B.R. (1987a). Revision of the Australian Vitaceae, 2. Cayratia Juss. Austrobaileya 2: 365-379.

— (1987b.) A study of the trichomes of several frequently confused species of Cissus L. (Vitaceae). Blumea 32: 143-147.

— (1988). Revision of the Australian Vitaceae, 3. Cissus L. Austrobaileya 2: 481—505.

LAUTERBACH, C. (1925). Die Vitaceen Papuasiens. Botanischer Jahrbticher fiir Systematik, Pflanzengeschichte und Pflanzengeographie 59: 505-534.

Liu, X.Q., ICKERT-BOND, S.M., CHEN, L.Q. & WEN, J. 2013. Molecular phylogeny of Cissus L. of Vitaceae (the grape family) and evolution of its pantropical intercontinental disjunctions. Molecular Phylogenetics and Evolution 66: 862-878.

Lu, L.M., Cox, C.J., MATHEWS, S., WANG, W., WEN J. & CHEN, Z.D. (2018). Optimal data partitioning, multispecies coalescent and Bayesian concordance analysis resolve early divergences of the grape family (Vitaceae). Cladistics 34: 57-77.

Powo (2022). Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. http://www. plantsoftheworldonline.org/, accessed 11 July 2022.

THIERS, B. (continuously updated). Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden. http://sweetgum.nybg.org/science/ih/, accessed 29 August 2022.

19

Plantago nupera Menkins (Plantaginaceae), a new species from the basaltic uplands of the southern Darling Downs, Queensland

Ian L. Menkins! & Paul I. Forster?

Summary

Menkins, I.L. & Forster, P.J. (2023). Plantago nupera Menkins (Plantaginaceae), a new species from the basaltic uplands of the southern Darling Downs, Queensland. Austrobaileya 13: 20—33. Plantago nupera Menkins is described as a new species. It is a grassland or grassy woodland specialist from the basaltic uplands on the Darling Downs of Queensland where it has a restricted distribution in the area encompassing Allora, Clifton and Maryvale. The new species is illustrated, with notes provided on its distribution and habitat, a suggested conservation status, and etymology. Plantago L. comprises 12 species in Queensland with seven of these being native. An identification key to these species 1S provided.

Key Words: Plantaginaceae; Plantago; Plantago nupera; flora of Australia; flora of Queensland; Darling Downs basalt flora; new species; taxonomy; identification key; conservation status

‘Oakey, Queensland 4401, Australia. Email: imenkins@optusnet.com.au; “Queensland Herbarium and Biodiversity Science, Department of Environment and Science, Brisbane Botanic Gardens, Mt

Coot-tha Road, Toowong, Queensland 4066, Australia. Email: paul.forster@des.qld.gov.au

Introduction

Plantago L. with c. 250 species (Rahn 1996; Rensted et al. 2002; Hassemer ef al. 2019; Shipunov et al. 2021) comprises both annual and perennial herbs. The genus has a cosmopolitan distribution, although with a concentration of species in temperate regions or at higher altitudes in the subtropics and tropics (Rahn 1996; Hassemer ef al. 2019; Shipunov ef al. 2021). Many of the most widespread species occur as naturalisations in Australia, hence the genus can be falsely construed as comprising predominantly alien species (WOA 2016; Blake 2020). Despite this, 24 of the 33 species that occur in Australia are considered native (Briggs 2022). Many of the Australian species have limited distributions in restricted habitats (Briggs ef al. 1977). Plantago has been divided into five subgenera with further division into sections (Shipunov ef al. 2021). The native Queensland species are all classified in Plantago section Mesembrynia Decne., whereas the naturalised species belong in other subgenera and sections (Table 1).

Speciation in Plantago is complex, involving polyploidy (Ishikawa ef al. 2009; Meudt 2011, 2012), inbreeding (cleistogamy) and outbreeding (chasmogamy), often with admixtures of the two (Hassemer ef al. 2020). Genomic evolution is inferred to have been quite rapid, based on plastome data that indicates considerable upheaval resulting in rearrangements driving hypermutations (Mower et al. 2021).

The seeds of Plantago species are coated with mucilage and this has been postulated to aid long range dispersal events by birds resulting in subsequent radiation (Rensted et al. 2002; Tay et al. 2010; Ahlstrand et al. 2018). Local dispersal events are, however, more likely from ground dwelling animals (especially insects), and wind or water. Pollination, where it occurs, can be by wind, insects, or a combination of the two (Abrahamczyk ef al. 2020), although the majority of species are probably wind pollinated (Hammer 1978). This feature, together with widespread self-compatibility is thought to have enabled colonisation of remote land masses (Ahlstrand ef al. 2018).

Accepted for publication 13 March 2023, published online 10 May 2023

© Queensland Herbarium 2023. ISSN 2653-0139 (Online)

Menkins & Forster, Plantago nupera

Herbarium collections of Plantago were first made in Australia by Joseph Banks and Daniel Solander in 1770, then by Robert Brown in 1802-1803; however, surprisingly, none seem to have collected them in what is now Queensland. Brown (1810) named four new species based on his collections: P. carnosa R.Br. nom. illeg., P. debilis R.Br., P. hispida R.Br. and P. varia R.Br. Within the geographical jurisdiction of the Queensland flora, Plantago was first recorded by Bentham (1870) with P. debilis and P. varia \isted. Bentham’s (and later Bailey’s 1900) inclusion of P. varia in the Queensland flora was based on T.L. Mitchell’s collection “In the interior”. This collection is most likely represented by two sheets in herbaria at the Royal Botanic Gardens, Kew (K 000779672) and _ the Muséum National d’Histoire Naturelle, Paris (P 00609916) respectively. The K specimen was determined on the 31 May 1977 as P. gaudichaudii Barnéoud by L. Johnson; however, it appears to be P. cunninghamii Decne. based on our examination of the digitised specimen images. The K specimen is undated and labelled as ‘Lieuten. Col... T.L. Mitchell’, ‘Subtropical N. Holl’ and ‘aff. varia’. The P specimen was determined on 8 May 1972 as the holotype of P. mitchellii Decne. by B.G. Briggs, who later included this name in the synonymy of P. cunninghamii (Briggs et al. (1977). The P specimen is also undated and labelled as ‘Cpn Mitchell’, ‘Subtrop. N. Holl. with another label stating it is from the Decaisne herbartum and with Plantago mitchellii on that label. Decaisne (1852), when naming P. mitchellii stated “In Nov. Holl. subtropica (Cap. Mitchell)” and “(v.s. in herb. Hooker)’. Decaisne probably saw the material while it was still in the personal possession of Hooker, with subsequent splitting of the collection between the two herbaria. The designation of the P collection as the holotype for the name P. mitchellii by Briggs et al. (1977) should now be considered as a lectotypification in retrospect (Art. 7.11 and 9.10) (Turland ez al. 2018) with duplicates of this collection to be designated as isolectotypes.

21

As noted by Gilbert (1971), Bentham “often had trouble with localities’, and the Mitchell collection may have been from either New South Wales or Queensland given the lack of a date. Further investigation of Mitchell collections at BM, CGE and K is required to determine whether any other collections of Plantago were made by him or whether the cited collection can be dated and further localised.

The last overall taxonomic account of Plantago for Queensland remains that of Bailey (1900), who listed the native species P. debilis and P. varia in a direct lift from Bentham (1870), the latter once again based on Mitchell’s collection from “In the Interior”’. Bailey (1900) also included the alien species P. lanceolata L. and P. major L. Bailey (1913) listed the same species, with the addition of an illustration of “P. varia’. This illustration may be based on an unaccessioned BRI specimen of P. cunninghamii which bears a Bailey handwritten label “Plantago varia R.Br.”’, but with no other information. Domin (1929) listed only P. debilis as native, with P. varia 1n synonymy. By the time of the Flora of South-eastern Queensland, four native species (P. cunninghamii, P. debilis, P. gaudichaudii, P. hispida) and five alien species were recorded (Ross 1986). Recent Census of the Queensland Flora accounts (Bean 2021, 2022) enumerate 11 species, with six of these considered native, including P. varia once again.

To some extent, the application of names to Plantago species in Queensland has been haphazard over time, given that there has never been an overall revision of the Australian species. This, together with misapplications and misidentifications, has resulted in some incorrect names (e.g. P. varia) being applied to species locally (Bentham 1870; Bailey 1900, 1913; Bean 2019, 2021, 2022). Superficial similarity between species is largely to blame for these errors and this has not been restricted to Australian material (cf. Shipunov ef al. 2021). Some species are obligate annuals, whereas others (e.g. P. cunninghamii, P. debilis), including some of the naturalised species (e.g. P. lanceolata)

can be annual or short-lived perennials, exhibiting marked developmental changes in gross morphology. Investigation of the flora of the Darling Downs in southern Queensland resulted in several identification problems with Plantago, and the realisation that a unique taxonomic entity was present in a small area around Allora. This entity is described as the new species Plantago nupera Menkins in this paper.

Plantago nupera was initially suspected of being an alien species, partly because the first collections were only made in the 1990s. Species such as Plantago media L., P. tomentosa Lam. and P. virginica L. from Plantago subgenus Plantago were among the list of suspects based on superficially similar morphology. However, DNA analysis using the nuclear internal transcribed spacer

Austrobaileya 13: 20—33 (2023)

(ITS) region conducted by the University of Adelaide RABLAB showed that the species eroups with the other native Australian species sequenced. The phylogeny suggests that P. nupera is most closely allied to P. debilis and P. hispida (J. Cowley, pers. comm., 2021), which are classified in P. section Mesembrynia (Shipunov 2021). By contrast, Plantago media is in P. section Lamprosantha Decne. and P. tomentosa and P. virginica are in P. section Virginica Decne. & Steinh. ex Barneoud (Hassemer et al. 2019; Shipunov 2021; Shipunov ef al. 2021). Given the complexity of the genomics in Plantago (Mower et al. 2021), together with relatively limited overall taxon sampling, it 1s rather likely that allocation of species to the morphology-based sections may change in the future.

Table 1. Subgeneric and sectional placement of Queensland species of Plantago from

Shipunov (2021, pers. comm., 2022)

cunninghamii Decne. Native debilis R.Br Native drummondii Decne. Native

gaudichaudii Barnéoud hispida R.Br.

lanceolata L.

Plantago section Mesembrynia Native

website by PIF. Results from phylogenetic analysis using ITS sequence data were kindly supplied by J. Cowley and R. Burton of the University of Adelaide, based on samples of Plantago nupera from the type locality.

major L. myosuros Lam. nupera Menkins

turrifera B.G.Briggs, Carolin & Pulley

Materials and methods

Measurements and descriptions are based on observations of plants in habitat and in cultivation in a common garden situation at Oakey, Queensland by ILM, herbarium collections at the Queensland Herbarium (BRI) and type collections of Plantago available online via the JSTOR Global Plant

A binocular microscope at magnification levels 20X and 40X was used to measure and describe the smaller details.

Menkins & Forster, Plantago nupera

Descriptive measurements are inclusive, e.g. 0.5—3.0 would be given as 0.5—3. In the description of hairs, stout is defined as rigid or coarse to the touch with the hairs generally relatively short and thick, while slender is defined as soft and flexible to the touch with the hairs generally long and thin.

The species key has been developed by ILM based on observation of the species and character states and data included in Kafri (1974), Feinbrun-Dothan (1978), Briggs ef al. (1977), Briggs (1980, 2022), Meikle (1985), Jeanes (1999), Brown (1991), Hassemer et. al. (2019), Jager (2011), Meudt (2012), Bean (2019) and Shipunov (2021). A key to all Australian species of Plantago 1s also available (Menkins 2022).

Taxonomy Plantago nupera Menkins, sp. nov.

Differs from P. debilis by the longer, more elliptic leaves, the spreading to near patent hairs on the lower portion of the scapes, the more pubescent inflorescence, the larger fruits, and the very long and persistent styles 5—9 mm long which often remain intact on pressed and dried specimens. The styles + straight or slightly arcuate in fresh material, becoming irregularly curled and _ twisted on drying. Type: Queensland. DARLING Downs DistTrRIcT: Forest Plain Road, 1.1 km E of Allora, 21 September 2022, PI. Forster PIF48254, G. Leiper & ILL. Menkins (holo: BRI [2 sheets comprising 3 individual plants + spirit]; iso: CANB, MEL, NSW).

Rosulate perennial, either prostrate or upright (tufted); taproot stout and very well developed. Indumentum comprising simple, eglandular, uniseriate (comprising 3-8 cells) hairs. Leaves thickened but not fleshy, slightly coriaceous, in juvenile stage ovate to obovate-oblanceolate, by fruiting stage lanceolate to narrow elliptic (rarely broadly falcate), broadest slightly above middle of lamina, both surfaces moderately hairy, lime green or yellowish-green, rarely darker green, dull to the naked eye, glistening and minutely colliculate on magnification; veins (including mid-vein) numbering 3-—S(-7), obvious on both surfaces, outermost obscure

23

if close to margins, depressed with scattered hairs on adaxial surface, prominently raised and ciliate on abaxial surface; hairs to c. | mm long, translucent to white, spreading, mostly patent, a few inclined, stout but soft to touch, straight or flexuous, apex aristate and often slightly deflexed; leaf margins entire or shallowly scalloped with 3—5 opposing pairs of small symmetrical teeth, the largest measuring 1—1.5 mm long X c. 2 mm wide, each ending on the adaxial surface in a small, ill-defined tubercle measuring 1-3.5 mm diameter X c. | mm high and surmounted by a small, unilateral tuft of short, antrorse hairs, the corresponding abaxial position forming a nearly glabrous dimple; inner and juvenile leaves spreading, imbricate, 3-6 X 2-4 cm, both surfaces covered in moderately dense, short, stout, erect hairs, apex subobtuse, petiole indistinct; outer and mature leaves 8-22 X 2-4 cm (including petiole), spreading, petiolate, not imbricate, each covered in moderately dense, short, stout hairs on both surfaces, apex sub-obtuse or broadly acute, leaf base attenuate, tapering very gradually to a slender petiole measuring 0-50 X c. 4.5 mm, axillary hairs tufted, 1.1- 6 mm long, dark golden brown, obscured by the broadened leaf base. Inflorescence erect or ascending, often somewhat arcuate about the middle (in rare cases coiling), narrow- cylindrical, 14—56 cm long, of two types with no obvious intermediates that often appear on the same plant, one with a short and compact spike, the other long with flowers irregularly spaced on an interrupted spike, surface dull ereen or dark yellowish-purple when fresh, pale brown or dull grey when dry, moderately pilose, hairs c. 1 mm long, white, appressed in upper part, antrorse to near patent in lower third to half; scapes stout, 10—30.5(-35) xX 0.2—0.3 cm, spike 8—26 X 0.6—1 cm wide, typically one third to approximately equal the length of the scape, of two types, the first compact-cylindrical with close, evenly spaced flowers, the other narrow-cylindrical and elongated with irregularly spaced flowers and fruits, in both cases with flowers and fruits more widely spaced in the lowest portion of the spike; bracts narrow elliptic- deltoid, c. 2/3 length of sepals, 1.5-3.2 xX

24

l1-1.6 mm, dull green in fresh state, dull brown in dry state, coarsely hairy, margins glabrous and narrowly membranous or ciliate, apex acute and slightly incurved-mucronate, keel somewhat thickened, either angled and distinct or broad and indistinct. Flowers sessile, chasmogamous, numerous; style straight or arcuate, sometimes kinked or twisted apically, 5—7(—9) mm long, exserted beyond corolla lobes, dark grey, rusty brown or purplish, pilose, hairs dense, short, white, patent; sepals elliptic, subequal, 2.2—3.5(—4) x 1.5—1.8mm, lime green in fresh state, brown in dry state, glabrous except on keel, apex broadly acute to subobtuse, membranous margins about the same width as the keel, glabrous or with a few ciliate hairs in apical region, white with purplish highlights in fresh state, pale brown and translucent in dry state, keel somewhat thickened, either angled and distinct or broad and rather flat, darker than membranous margins, covered in short, coarse, antrorse hairs; corolla tube 1.5—2.1 mm long, inner and outer surfaces membranous, glabrous, pale brown to brown; corolla lobes ovate-deltoid with an acute to slightly attenuate apex, 1.3-2.1 xX 1.5—1.7 mm, minutely rugose, pale brown or purplish, translucent, often darker brown or purplish black and + opaque towards base, spreading to reflexed, rarely remaining erect, midvein distinct; filaments straight or slightly arcuate, 5—7 mm long, white; anthers apicifixed, obovate or narrowly obcordate, vertically symmetrical, 1-2 X< 1-1.5 mm, pale creamy brown or yellow with purplish margins; anther connective 1/20" to 1/3" the total width of the anther at its widest point, both surfaces greenish-yellow or creamy white, abaxial surfaces of the thecae cream to yellow with purplish margins, adaxial surfaces purplish throughout or cream to yellow with purplish margins, inner thecal margins vertical, incurved, adpressed upon the connective, outer thecal margins slightly wavy, narrowly rounded in outline, rupturing along the stomium, apex obtuse and purplish, base mucronate and deep purple. Fruita pyxis, bilocular, ovoid-conical to subglobular, 3—4.8 xX 2.5-2.6 mm, asymmetrically and partially

Austrobaileya 13: 20—33 (2023)

contracted in upper portion, tapering to apex; lid 2.5-3 X 2-2.1 mm, creamy brown or golden-brown, surface minutely rugose with 5 or 6 obscure but evenly-spaced vertical impressions, apex tapering acutely to a shrivelled, acuminate-attenuate mucro or cusp that is 1-1.5 mm long, of similar shape to the spike on a pickelhaube; seeds very obscurely rugose, with a small superficial hilum and a thin mucilaginous layer, ellipsoidal, flattened adaxially, numbering 1—4(—5) per capsule, 1.5—2.5 X 1-1.1 mm, yellowish brown. Figs. 1-5.

Additional specimens examined: Queensland. DARLING Downs District: 24 km ENE of Clifton, Feb 1995, Fensham 2070 (BRI); 13 km ENE of Allora, May 1994, Fensham 1533 (BRI); Mount Pleasant Road, off Forest Springs — Goomburra Road, Forest Springs, Sep 2022, Forster PIF48270, Leiper & Menkins (BRI); 2.2 km E of Allora along Forest Plain Road, Oct 1996, Bean 10853 (BRI); 7.7 km along North Branch Road, near Maryvale, Oct 1996, Bean 10830 (BRI). CULTIVATED:

Oakey (ex Forest Plain Road, 1.1 km E of Allora), Sep 2022, Menkins s.n. (BRI, MEL).

Distribution and habitat: Plantago nupera is endemic to Queensland on the eastern Darling Downs in a small area from near Maryvale in the south, to just east of Hirstglen in the north. It occurs in ungrazed, or seldom erazed situations, in relatively intact remnant erassland comprising predominantly an admixture of Cymbopogon refractus (R.Br.) A.Camus, Dichanthium sericeum (R.Br.) A.Camus subsp. sericeum and Themeda triandra Forssk. (Fig. 6), in relict or partly cleared grassy woodland dominated by Eucalyptus orgadophila Maiden & Blakely and/or &. tereticornis subsp. basaltica A.R.Bean or comprising variously E. albens Benth., &. melliodora A.Cunn. ex Schauer, £. molucanna Roxb., FE. nobilis L.A.S.Johnson & K.D.Hill and Angophora floribunda (Sm.) Sweet. It appears to show a preference for south-facing slopes. Collectively these locations are all part of the basalt uplands of the Eastern Darling Downs; they largely equate to the eastern portion of the ‘hill woodland’ of Fensham (1998) and are part of the western foothills of the Great Dividing Range.

Menkins & Forster, Plantago nupera 25

Fig. 2. Rosette of flowering plant of Plantago nupera in situ at type location. Photo: P.I. Forster.

Fig. 1. Flowering plant of Plantago nupera in situ at type location. Photo: P.I. Forster.

Fig. 4. Flowers mainly in staminate stage for Plantago nupera at type location. Photo: G. Leiper.

Within this general landscape, several Plantago species occur, including P. cunninghamii, P. debilis, P. nupera and the alien species P. lanceolata, although we have not observed the other species co-occurring with P. nupera. Plantago lanceolata 1s often in proximity, although it is invariably Fig. 3. Flowers in pistillate stage for Plantago nupera at associated with disturbed habitats such as type location. Photo: G. Leiper. irregularly mown roadside verges.

26

Fig. 5. Flowers predominantly bisexual for Plantago nupera at type location. Photo: G. Leiper.

At the type locality, Plantago nupera 1s also associated with a large stand of Discaria pubescens (Brong.) Druce on a steep hillside that is deeply eroded and channelled (Fig. 7). Plants occur on the highest point, growing amongst tussocks of Cymbopogon refractus and Themeda triandra where the soil is crumbling and falling away (Fig. 8). The situation 1s quite exposed, despite being a south-facing slope and is immediately adjacent to a grassy woodland of mainly Eucalyptus orgadophila with occasional E. tereticornis subsp. basaltica. At a second location (Forster PIF48270 et al., BRI), the plants occur in grassy woodland dominated mainly by £. moluccana, mainly towards the base of a low basalt rise.

The soil where Plantago nupera occurs is invariably a black vertisol containing nodules of basalt and calcined clay. The pH is alkaline. These soils are basalt derived and often characterized by self-mulching,

Austrobaileya 13: 20—33 (2023)

cracking clays (black earths) (Beckmann et al. 1974). Plantago nupera occurs on the rather more skeletal examples of these soils that occur 1n the foothills of the Great Dividing Range, which typically show little or no profile development. The soils are very dark grey or brown to black in colour and often stony or possessing a granular texture. They are generally well-drained and friable when dry, but able to retain some moisture in the clay particles for extended periods after rain events.

Notes: Plantago nupera_ has — gross morphological similarities to P. exilis Decne. from Western Australia and P. varia from southern temperate Australia (Table 2). It differs most noticeably from P. exilis in the axillary hair colour, the shorter petioles, the longer capsules and broader seeds and from P. varia in the axillary hair colour, hair orientation on the lower scape, the stout hairs on the adaxial leaf surface, bract shape and seed colour.

Based on ITS sequence data, Plantago nupera forms a monophyletic group with P. debilis and P. hispida (J. Cowley, pers. comm. 2021). Plantago nupera differs from P. debilis in the axillary hair colour, shape of the leaf laminae, form of the indumentum on the lower scape and the lower lamina surface, bract shape and longer sepals. Plantago hispida is disjunct in occurrence from P. nupera, being further south on the Granite Belt near the Queensland — New South Wales border. Plantago nupera differs from P. hispida in the axillary hair colour, form of the indumentum on the lower scape and the lower leaf lamina surface, bract shape, capsule shape and seed colour. ITS data is presently not available for Plantago exilis to enable a_ genetic comparison. Plantago exilis grows almost exclusively on sandy loam overlying granite formations where the pH is generally acidic, while P. nupera occurs on rather deeper, more clayey loams of basaltic origin and the pH 1s invariably alkaline.

Menkins & Forster, Plantago nupera 27

Table 2. Morphological characters of Plantago nupera and allied Australia species

Stout

Leaf lamina shape narrow-elliptic to | narrow-elliptic | narrow-elliptic oblanceolate | narrow-elliptic to oblanceolate or obovate to oblong- oblancolate

Leaf length (cm) 85-23 (2-)4-9(-16) | 3-15(-25)

Axillary hair colour dark golden pale brown white or pale pale brown reddish brown brown yellow-brown

Hairs on lower 2 of spreading to near | spreading spreading to + appressed + appressed patent appressed

scape

Leaf hairs on adaxial stout, but soft stout, but soft | slender Slender Slender

lamina surface

Bract shape narrow-elliptic narrow-elliptic | ovate ovate-elliptic | ovate deltoid deltoid

Sepal length (mm) 2.2-3.5(-4) 2.2-2.8(-3) 1.5-2.2(-3) | 2.8-3.5(-4)

Corolla lobe length 1.3-2.1 1.6—2.5 1.3-1.5 0.8—] 1.5—2.3 (mm)

Leaf lamina rib count | 3-5(-7) 3(-5) 1-3(-5)

Pyxis shape and length | ovoid-conical ovoid-conical, | ovoid-ellipsoid, | ovoid to ovoid-ellipsoid (mm) to subglobose, 2.8—3.5 2.8—3.5 subglobose, | to subglobose, 3.4-8 1.7-3 2.8—3

1-5, 1.52.5, 1-5, 1-1.5, 1-5, c. 1, 1-5, 1.5—2.5, yellow-brown yellow-brown yellow to brown

elliptic

Seed number in capsule, length (mm), colour

Conservation Status: The relict, grassland and grassy woodland communities in the eastern foothills of the southern Darling Downs are characterised by soil types much valued for agriculture. These habitats are collectively becoming known as a minor regional centre for local speciation or species distribution rarity, and this is reflected in an increasing number of taxa being allocated conservation status listing. It has been largely cleared for agriculture, or greatly altered by land management practices or the invasion of alien species (Fensham 1998; Fensham & Fairfax 1997), particularly introduced pasture grasses and legumes (Silcock & Scattini 2007).

The handful of locations where Plantago nupera has been found are ungrazed by domestic stock. They are either on road reserves or land portions that are subject to indifferent management, and decreasing

brown

in natural biodiversity and concomitant conservation values every year. This decrease in habitat quality and composition is indirectly impacted by management of adjacent agricultural land (e.g. herbicide drift, changes in moisture run off patterns, alien weed invasion) or directly by herbicide application, slashing, burning and overall clearing. Fensham (1998) found that 10% of species from his ‘hill woodland’ community were sensitive to grazing pressure and this appears to be the case for P. nupera based on its absence from grazed areas adjacent to known locations. Plantago nupera 1s yet to be found in the Allora Mountain Fauna and Flora Reserve that is in close proximity to the type location for the species; however, that reserve has been heavily grazed by private leaseholders in recent times with heavy

infestations of alien weed species (PIF, pers. obs.).

: 20-33 (2023)

Austrobaileya 13

28

ee

_—a

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P.I. Forster.

(arrowed). Photo

10n

tu at type locat

lh St

Fig. 6. Flowering plants of Plantago nupera scattered

ia pubescens, local

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D

ing

bundant flower!

ing a

show P.I. Forster.

9

tat for Plantago nupera at type location

1

ty to aroad and agr

Fig. 7. Grassland hab

Itural land. Photo:

1cu

m1

d prox

Crosilon an

Menkins & Forster, Plantago nupera

29

Fig. 8. Grassland habitat for Plantago nupera at type location, a single plant indicated by arrow. Photo: P.I. Forster.

An assessment of the Extent of Occurrence (EOO) and Area of Occupancy (AOO) for Plantago nupera using the GeoCat analysis tool (Bachmann ef al. 2011), found an EOO of 298 km* and an AOO of 16 km°. The species is currently known from four locations (one with two sublocations) in a highly fragmented and predominantly cleared landscape. The habitat at these locations continues to be impacted by herbicide drift from adjacent farmland, alien weed invasion and occasional grazing by domestic stock. By applying the categories of the IUCN (2012), a conservation status of Endangered (E) is suggested, based on criteria Bla,b(i—v),c(i1i,1v);_ 2a,b(i—v),c(111,1V); however, a formal assessment 1s yet to be completed.

The likelihood of returning this landscape to a pre-European state in remaining remnants is likely difficult to achieve, with management to maintain a predominantly native composition thought to be dependent on the right combination of seasonal conditions, fire and grazing pressure (Silcock & Scattini

2007). Environmental factors such as soil pH, silt, water logging and seasonal conditions in these communities were found to be more significant than fire regimes, slashing or erazing by Fensham ef al. (2017); however, this is likely to be more relevant to larger remnants rather than in narrow road reserves with large edge effects and recurring impacts.

Plantago nupera_ co-occurs with at least one other threatened plant (Discaria pubescens) at one of its known localities. All the known locations are very small in extent, with the estimated AOO being much greater than the actual area on the ground. The Plantago 1s rather cryptic within the broader landscape and may well prove to be more widespread 1f all land tenures within its EOO could be adequately surveyed.

Etymology: The specific epithet comes from the Latin adjective niiperus (feminine nupera, neuter nuperum) meaning “late”, “fresh”, or “recent”. The name alludes to its relatively recent recognition as a species.

30 Austrobaileya 13: 20—33 (2023)

Key to Plantago taxa in Queensland (* indicates naturalised taxon)

1 Plants caulescent; leaves opposite on erect, elongated, branching stems;

MAT RS ATION oh or 5-75-08 ea aes se. or sh BR A ale ae ta ee Plantago arenaria* 1. Plants acaulescent; leaves basal, rosulate, prostrate or tufted, occasionally ascending; hairs, if present,eglandular ................... 2

2 Leaves pinnately or bipinnately lobed; spikes compact with

flowers appressed to axis; corolla tube sparsely hairy or hairy

ii LOG Mal OTy 8 2 gs og tintcves dae fp SSeS dh a toe ae Wy Ee Plantago coronopus* 2a Most scapes longer than leaves; bracts mostly longer than

sepals; keel of anterior sepals about equal to width of

MemMmbranous Mares: 4 4 6 4 bark a ae wes Plantago coronopus subsp. coronopus* 2b Most scapes shorter or equal to leaves; bracts mostly shorter

or equal to sepals; keel of anterior sepals much wider than

INEMBTANOUS INMATOINS oa. yb be a Ee Plantago coronopus subsp. commutata* 2. Leaves entire or toothed; spikes loose, or 1f compact then with

spreading flowers; corolla tube glabrous throughout. ................... 3 3 Flowers cleistogamous; corolla lobes erect; anthers not exserted . . . Plantago myosuros* 3. Flowers chasmogamous; corolla lobes spreading or reflexed; anthers exserted....... al 4 Capsules 8—16-seeded; leaf bases abruptly contracted or truncate ... . Plantago major* 4. Capsules up to 5-seeded; leaf bases attenuate or rounded, never truncate ......... 5 5 Peduncles with a series of longitudinal, parallel grooves and ridges;

spikes broad, subconical, ovate, or almost cylindrical; anterior sepals

fused tor at least other length, os a a Yee ee ho ee we Plantago lanceolata* 5. Peduncles without grooves and ridges; spikes cylindrical; anterior

SONA SHMCCLOE TOQEIV 2S O58 ag x, ce Bs Se as cle Mae eS ad ws cee eee ee tt ah see Dos Bt Arata Ms de Bt 6

6 Spikes narrowly cylindrical, elongating and lax at maturity; small capsules to 3 mm long, often well interspaced, particularly in lower

BYU FOE VOA UIT oS yy ez ie hea chee) Eh at OE cere newt Ae Fe Hot) AA Ws hat of? oe ake Snes Ss P. debilis 6. Spikes cylindrical, upright at maturity; capsules larger, 3.3-6 mm long,

IV PiealIN Close TOCETNE 2 an et te Sa Be MERE Soe oe at et te Nae be a a eet ot tt SE TS era 8 7 7 Leaves more than 15 times longer than wide. ............ Plantago gaudichaudii 7. Leaves tarely more than 3 Times as longas Wade .. & ksi bawle bh ne wee ee ee 8 8 Plants annual; bracts and sepals glabrous or with few hairs along the keels ........ 9 8. Plants perennial; bracts and sepals with numerous coarse or soft

HaIVS-AlOMe Ie KCCIS:s othe ote doe, i foe a hbesee Th doen ei teach Gee ele Ales se PB up lie 11

9 Mature pyxis ovoid, obovate, or ellipsoid, 2.5—5 mm long, dark purple or

dark brown; apex acute to subobtuse; plant of arid and semi-arid

floodplains but generally not of swampy ground. ......... Plantago drummondil 9, Mature pyxis obovate-pyriform to pyriform, 1.5-2.5 mm

long, pale brown or tinged a purplish red; constriction and taper

symmetrical or asymmetrical in outline; apex acute ortruncate ............ 10

Menkins & Forster, Plantago nupera

10 Pyxis obovate-pyriform, broadly constricted in upper third to half and tapering to apex; constriction and taper symmetrical or asymmetrical in outline; apex acute, not quadrangular or lobed; plant mostly of inland colluvial and alluvial plains, preferring more fertile soils in

31

HATUTANOEISSIATIGS: fox. cpl) AN tas lat wk eget Sp oY oe br eteaee atest Plantago cunninghamii

10.Pyxis prominently pyriform with the narrowest part forming a cylindrical beak above mid-point; beak generally parallel in outline or with an additional slight compression around the middle; apex truncate, quadrangular, and minutely lobed; small plant of mostly impoverished soils in inland regions, e.g. on hard, stony or

sandy ridges and plains, and on claypans and “scalds” ........ Plantago turrifera

11 Small plants; spikes 1-6 cm long; bracts ovate, minute, 1-1.9 mm long, less than half the length of the sepals; hairs in leaf axils short, to c. 2 mm long, whitish, pale brown or pale yellowish-brown; pyxis ovoid-globular to ovoid-ellipsoid. Confined to the extreme SE of the Granite Belt east of Wallangarra where it has been collected from crevices in rock pavement, but possibly to be found elsewhere on the

CTPA IGE ESCM wtnetcugs stag) hy Se, fet a Gh ES, Sa eaeee 5 cp ROE ele, cn te oe oh Ng Plantago hispida

11. Robust plants, spikes 8—26 cm long; bracts narrowly elliptic-deltoid, 1.5—3.2 mm long, c. 2/3 the length of the sepals; hairs in leaf axils long, 2—6 mm long, dark golden brown; pyxis ovoid-conical to subglobular. In ungrazed or seldom grazed grassland on the basaltic slopes of the extreme south-east of the Darling Downs, but possibly found elsewhere

1, SUT Lak NAIA Dee <1 qn ea: pro ake ee we Fae ere oe OE Plantago nupera

Acknowledgements

ILM wishes to thank Dr B.G. Briggs (NSW), Dr J. Cowley and Professor R. Burton (University of Adelaide), Dr R. Parsons (Latrobe University), Professor A. Shipunov (USA) and the very helpful staff of the Western Australian Herbarium (PERTH). Images of type specimens were viewed on the JSTOR Global Plants website. Glenn Leiper kindly shared numerous photographic images for the species.

References

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AHLSTRAND, N.I., VERSTRAETE, B., HASSEMER, G., DuNCAR-Co, S., HOGGARD, R., MEUDT, H.M. & R@NSTED, N. (2018). Ancestral range reconstruction of remote oceanic island species of Plantago (Plantaginaceae) reveals differing scales and modes of dispersal. Journal of Biogeography 46: 706-722.

BACHMAN, S., Moat, J. Hitt, A.W., DE LA TORRE, J.. Scott, B. (2011). Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. ZooKeys 150: 117-126.

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FENSHAM, R.J. & FAIRFAX, R. (1997). The use of the land survey record to reconstruct pre- European vegetation patterns in the Darling Downs, Queensland, Australia. Journal of Biogeography 24: 827-836.

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FENSHAM, R.J., BUTLER, D.W., LAFFINEUR, B., MacDernmott, H.J., MORGAN, J.W. & SILCOCK, J.L. (2017). Subtropical native grasslands may not require fire, mowing or grazing to maintain native-plant diversity. Australian Journal of Botany 05: 95-102.

GILBERT, L.A. (1971). Botanical Investigation of New South Wales 1811-1880. Volume 3. https://rune. une.edu.au/web/bitstream/1959.11/18449/31/ open/SOURCE24.pdf, accessed 15 February 2022.

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HASSEMER, G., BRUUN-LUND, S., SHIPUNOV, A.B., BricGs, B.G., MEUDT, H.M. & R@NSTED, N.A.H. (2019). The application of high-throughput sequencing for taxonomy: the case of Plantago subg. Plantago (Plantaginaceae). Molecular Phylogenetics and Evolution 138: 156-173.

HASSEMER, G., DOS SANTOS, A.P., SHIPUNOV, A.B. & FUNEZ, L.A. (2020). Plantago australis (Plantaginaceae) produces both chasmogamous and cleistogamous' flowers: Field work, herbarium and literature-based evidence. Flora 273. https//doi.org/10.1016/j.flora.2020.151724, accessed 31 January 2022.

ISHIKAWA, N., YOKOYAMA, J. & TSUKAYA, H. (2009). Molecular evidence of reticulate evolution in the subgenus Plantago (Plantaginaceae). American Journal of Botany 96: 1627-1635.

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JEANES, J.A. (1999). Plantaginaceae. In N.G. Walsh & T.J. Entwisle (eds.), Flora of Victoria 4: 463- 474. Inkata Press: Melbourne.

KAFRI, S.M.A. (1974). Plantago species and subspecies. In E. Nasir & S.J. Ali (eds.), Flora of West Pakistan 62: 1-21. Department of Botany, University of Karachi: Karachi.

MEIKLE, R.D. (1985). Flora of Cyprus 2. Bentham- Moxon Trust, Royal Botanic Gardens, Kew: London.

MeENKINS, I. (2022). Flora of Australia: vascular plants: Species of Plantago. https://keybase. rbg.vic.gov.au/keys/show/12277, accessed 25 October 2022.

Menkins & Forster, Plantago nupera

Meupt, H.M. (2011). Amplified fragment length polymorphism data reveal a history of auto and allopolyploidy in New Zealand endemic species of Plantago (Plantaginaceae): new perspectives on a taxonomically challenging group. [International Journal of Plant Sciences

172(220): 237.

— (2012). A taxonomic revision of native New Zealand Plantago (Plantaginaceae). New Zealand Journal of Botany 50: 101-178.

Mower, J.P., Guo, W., PARTHA, R., FAN, W., LEVSEN, N., WoLFr, K., NUGENT, J.M., PABON-Mora, N. & GONZALEZ, F. (2021). Plastomes from tribe Plantagineae (Plantaginaceae) reveal infrageneric structural synapormorphies and localized hypermutation for Plantago and functional loss of ndh genes from Littorella. Molecular Phylogenetics and Evolution 162. https//doi.org/10.1016/).ympev.2021.107217, accessed 31 January 2022.

RAHN, K. (1996). A phylogenetic study of the Plantaginaceae. Botanical Journal of the Linnean Society 120: 145-198.

R@NSTED, N., CHASE, M.W., ALBACH, D.C. & BELLO, M.A. (2002). Phylogenetic relationships within Plantago (Plantaginaceae): evidence from nuclear ribosomal ITS and plastid trnLl-F sequence data. Botanical Journal of the Linnean Society 139: 323-338.

Ross, E.M. (1986). Plantaginaceae. In T.D. Stanley & E.M. Ross (eds.), Flora of South-eastern Queensland 2: 469-472. Queensland Department of Primary Industries: Brisbane.

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SHIPUNOV, A. (2021). Plantagineae of the World. Identification keys. http://ashipunov.me/ shipunov/plantago/plantagineae of the world. pdf, accessed 8 February 2022.

SHIPUNOV, A., ALONSO, J.L.F., HASSEMER, G., ALP, S., Lee, H.J. & Pay, K. (2021). Molecular and morphological data improve the classification of Plantagineae (Lamiales). Plants 10: 2299. https://doi.org.10.3390/plants1l0112299, accessed 2 February 2022.

SILCOcCK, R.G. & SCATTINI, W.J. (2007). The original native pasture ecosystems of the eastern and western Darling Downs — can they be restored? Tropical Grasslands 41: 154-163.

Tay, M.L., Megupt, H.M., GARNOCK-JONES, P.J. & Ritcuig, P.A. (2010). DNA sequences from three genomes reveal multiple long-distance dispersals and non-monophyly of sections in Australasian Plantago (Plantaginaceae). Australian Systematic Botany 23: 47-68.

TURLAND, N.J., WIERSEMA, J.H., BARRIE, F.R., GREUTER, W., HAWKSwWorRTH, D.L., HERENDEEN, P.S., KNAPP, S., KUSBER, W.-H., L1, D.-Z., MARHOLD, K., May, T.W., McNEILL, J.. MONRO, A.M., PraAbDo, J., Price, M.J. & SMITH, 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. Koeltz Botanical Books: Glashiitten.

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Calandrinia halophila Albr. & J.G.West (Montiaceae), a new Species from estuarine environments in northern Australia

David E. Albrecht' & Judy G. West'

Summary

Albrecht, D.E. & West, J.G. (2023). Calandrinia halophila Albr. & J.G.West (Montiaceae), a new species from estuarine environments in northern Australia. Austrobaileya 13: 34—40. Calandrinia halophila Albr. & J.G.West sp. nov. is described and illustrated, and differences between it and similar species of Calandrinia discussed. Notes on distribution, habitat and conservation status are also provided.

Key Words: Montiaceae; Portulacaceae; Calandrinia; Calandrinia halophila; Parakeelya; Rumicastrum; flora of Australia; flora of Western Australia; flora of Northern Territory; flora of Queensland; taxonomy; new species

‘Australian National Herbarium, Centre for Australian National Biodiversity Research, (a joint

venture between Parks Australia and CSIRO), GPO Box 1700, Canberra, ACT 2601, Australia.

Introduction

In the early 1970s, initial collections of a small rosette-forming species of Calandrinia Kunth were made from a few estuarine sites in northern Australia. Since that time an additional 20 or so collections of the same taxon have been made from similar sites scattered across northern Australia. The species has been independently recognised as distinct by botanists, and assigned the phrase names Calandrinia sp. Berry Springs (M.O.Parker 855) and Calandrinia sp. Olive River (J.R.Clarkson+ 10012) at the Northern Territory Herbarium (DNA) and Queensland Herbarium (BRI) respectively. We here formally describe this entity as the new species Calandrinia halophila Albr. & J.G.West, acknowledging that the generic name is likely to change in the near future once the proposal to conserve the name Parakeelya Hershk. against Rumicastrum Ulbr. 1s voted on at the International Botanical Congress scheduled for 2024 (Thiele et al. 2018; West & Albrecht 2022).

Methods and materials

The species description is based on pressed herbarium specimens and spirit material housed at BRI, CANB, DNA and NSW. Representative flowers and _ fruits’ on dried specimens were rehydrated prior to assessment.

Measurements of plant parts are inclusive (e.g. 4.0—6.7 is given as 4—6.7). Following West & Albrecht (2022) the flower—bearing shoots were measured in two parts. The proximal portion was measured from the base to the closest bract/pedicel junction above. The distal flowering portion (1.e. inflorescence axis sensu Obbens 2011) was measured as the distance from the lowermost bract/pedicel junction to the apex of the distal-most bud or flower. Where inflorescences were reduced to a single flower the distal flowering portion equalled the combined length of pedicel and flower. Pedicel length was measured as the distance from its apex (at the junction with the sepals) to the insertion point of the closest bract below. For the purposes of consistency with descriptions of recently

Accepted for publication 29 March 2023, published online 10 May 2023

© Queensland Herbarium 2023. ISSN 2653-0139 (Online)

Albrecht & West, Calandrinia halophila

described species we here treat the two structures below the petals as sepals although it is acknowledged that they are considered to correspond to bracteoles or involucral bracts by some authors (see Nyfteler & Eggli 2010 for discussion). These structures were detached and flattened to measure. Fruit measurements are based on unflattened mature fruits prior to dehiscence.

Taxonomy

Calandrinia halophila Albr. & J.G.West, sp. nov.

Distinguished from other species’ of Calandrinia by the combination of the following characters: annual herb with leaves narrow and _ dorsi-ventrally compressed, strictly in a basal rosette; flower-bearing shoots not or only slightly exceeding the length of the longest leaves; bracts alternate; sepals non-persistent; petals (S—)6—7(—8), pale pink or rarely white; stamens 5—10; stigmata 5—7; capsules 2—3.2 mm long with an obtuse to subacute apex and splitting into 5—7 non- persistent valves; and shiny seeds mostly c. 0.35 mm long and 0.25 mm wide, with a very indistinct colliculate surface pattern. Type: Northern Territory. Berry Springs, 18 May 1977, M.O. Parker 855 (holo: CANB 366252; iso: DNA [D0011526]).

Calandrinia sp. Berry Springs (M.O.Parker 855) Dixon, R.A.Kerrigan & Cowie: CHAH (2006); Kerrigan & Albrecht (2007); Short et al. (2011); Cowie et al. (2017); Parker & Percy-Bower (2022: 2).

Calandrinia sp. (Olive River J.R.Clarkson+ 10012): Thomas (2007, 2010, 2013, 2016); Brown (2021).

Illustrations: Northern Territory Herbarium (2015) as Calandrinia sp. Berry Springs (M.O.Parker 855) Dixon, R.A.Kerrigan & Cowie.

Glabrous annual herb to c. 7.5 cm high with lateral roots arising from a non- or scarcely thickened taproot. Leaves all basal, mostly spreading, sometimes ascending or flat on substrate surface, succulent, narrowly oblanceolate, 3-60 mm long (the longer outermost leaves rarely < 15 mm long), 0.7—

35

3 mm wide, dorsi-ventrally compressed to oval in cross-section, dull green or tinged red/pink, orange or brown; base attenuate and lacking clearly differentiated petiole; margins rounded in cross-section for most of leaf length but becoming more angular proximally; apex narrowly rounded to subacute. Flower-bearing shoots arising from leaf axils, spreading or prostrate on substrate surface, equal to or slightly shorter or slightly longer than the longest leaves; proximal leafless portion (1.5—)4-38 mm long; distal flowering portion (2.5—)6—37 mm long, flowers arranged in |—3(—4)-flowered monochasial cymes or rarely dichasial cymes with monochasial branches, pedicels (2.5—)4—28 mm long, broader distally, fine, becoming somewhat wiry with age, narrowly longitudinally winged in dried specimens, persistent long after fruit abscission, not obviously elongating in fruit; bracts alternate, mostly appressed or ascending, triangular- ovate to elliptic, deeply concave, 1—2.5(—3) mm long, with a herbaceous midvein and broad hyaline margins, or rarely almost entirely hyaline, sub-acute to acuminate, tip sometimes recurved. Flowers 5—8 mm diameter. Sepals ovate, broadly ovate or broadly elliptic, 2-4 mm long and 2-3 mm wide in flower, enlarging to 2.5—5 mm long and 2.5—4 mm wide in fruit, at least one shedding before or with the capsule valves, free to base, venation not evident; margins narrow hyaline to c. 0.4 mm wide; apex broadly acute to obtuse. Petals (S—)6—7/(—8), oblong-elliptic to narrowly obovate, (2—)2.4—3.5(—4) mm long, Q0.6—-1.6 mm wide, free or virtually so, pale pink or rarely white, apex obtuse. Stamens 5—10; filaments linear, appearing free, 1.3—2.2 mm long, pale pink to white, smooth; anthers versatile, 2-celled, each cell oblong in outline, 0.3-0.7 mm long, 0.15-0.25 mm _ wide, latrorse, dehiscing longitudinally, violet pre- dehiscence. Ovary ovoid to broadly ellipsoid or almost spherical, 1.2—2.2 mm long, 0.8—1.5 mm diameter, reddish-pink to orangish-pink. Stigmata 5-7, linear, 0.4—0.8 mm long, free to the base, pale pink to white. Capsule ovoid to broadly ellipsoid, 2—3.2 mm long, 1.3—2.5 mm wide, not exceeding the sepals, pale brown, apex obtuse to broadly acute, initially

36

the distal third separating into valves and sometimes the proximal third also splitting longitudinally, finally some or all valves separating for the whole length; valves 5-7, the tips sometimes recurved, not persisting and thus exposing placentas and funicles, thin-textured, when dry the external surface with numerous close minutely roughened striations. Seeds numerous, obovoid, (0.3— )0.35(—0.4) mm long, (0.2—)0.25(—0.3) mm wide, mid brown, evenly shiny, appearing smooth at X10 magnification but very weakly colliculate, with a rudimentary strophiole. Fig. 1.

Additional specimens examined: Western Australia. Mary Island, Vansittart Bay, Kimberley Region, Jun 2010, Keighery KIBSI190 (PERTH); Northern bay of Coronation Island, Bonaparte Archipelago, May 1972, Wilson 10940 (PERTH). Northern Territory. Middle Arm area, adjacent Inpex site, May 2020, Brennan 11733 (DNA); Limmen National Park, Apr 2008, Dixon 1506 (DNA); Bathurst Island, track NE of sewerage ponds Neguiu, Feb 2001, Kerrigan & Risler 325 (DNA); Cooper Creek catchment, c. 47 km NW of Gunbalanya, Jul 1972, Lazarides 7510 (CANB, DNA, K, L); Limmen National Park, St Vidgeons block, south of Limmen River adjacent to salt flats, Apr 2009, Lewis 1157 (CANB, DNA); Middle Arm — Kittyhawk, Apr 2020, Lewis 3099 (CANB, DNA); Bathurst Island, Delawa Long Arm, May 1998, Michell & Harwood 1467 (DNA); N of Black Rock Landing, McArthur River, Feb 1976, Rice 2387 (CANB); 3 km S of Bing Bong Homestead, May 1984, Thomson 516 (DNA); Crab Claw Island, Bynoe Harbour, May 2010, Westaway 3236 (DNA). Queensland. Cook District: Saibai, Jul 1973, Stocker 1336 (CANB); c. 24 km SSE of the mouth of the Olive River, c. 3 km S of Mosquito Point, Apr 1993, Clarkson 10012 & Neldner (BRI, CANB, DNA, K); Lakefield National Park, Knifehole, 2 km from Saltwater Creek crossing on the Musgrave to Lakefield Road, May 1987, Clarkson 7060 & Simon (BRI, CNS, DNA, L, NSW, PERTH). BuRKE District: 54 km NW of Burketown on Escott Station (Gulf site 378), Apr 2007, Thompson & Wilson WES1371 (BRI); 7 km NE of Normanton (Gulf site 57), Jun 2001, Thompson & Turpin NOR294 (BRI); c. 10 km along the Normanton to ‘Mogoura’ Station road, SW of Normanton, Apr 1974, Pullen 8858 (CANB); 15 km c. SW of Normanton on Burketown Road, Apr 1974, Jacobs 1194 (NSW). NORTH KENNEDY DiIstRIcT: Bohle River estuary, c. 13 km WNW of Townsville, Feb 1998, Cumming 16799 (BRI); Townsville, at the east end of Marina Drive, Bushland Beach, Jul 2021, Horsfall s.n. (BRI, CANB 954477); Bushland Beach, Stony Creek, Bohle River boat ramp, Sep 2022, Horsfall s.n. (CANB 955859); Wunjunga Road, off Bruce Highway, S of Inkerman, S of Ayr, Jun 1997, Cumming 16082 (BRI); Gloucester Island, S end, site 38, Apr 1994, Batianoff & Figg 940430G (BRI).

Austrobaileya 13: 34—40 (2023)

Distribution and habitat: Calandrinia halophila is currently known from scattered locations along the northern Australian coast, including the Kimberley, greater Darwin region and Tiwi Islands, Gulf of Carpentaria and north-eastern Queensland north of Mackay (Map 1). There is also an occurrence on Saibai Island in the Torres Strait, approximately 5 km south of Papua New Guinea. It is anticipated that the species probably grows on the southern coast of mainland New Guinea.

Calandrinia halophila occurs within, or adjacent to estuarine environments in northern Australia. Plants are often found in areas lacking or with a very sparse overstorey canopy of Melaleuca spp. (predominantly M. acacioides F.Muell.) or mangrove species (Fig. 2). Associated ground layer species include Sporobolus virginicus (L.) Kunth, Xerochloa imberbis R.Br., Fimbristylis spp., Calandrinia gracilis Benth. sens. lat. and Tecticornia spp. Many collectors explicitly state or imply that the substrate is saline or subsaline, with sandy to gravelly surface soil over sandy saline clay being a typical description of the substrate. A small number of collections have been made on islands where the landform type may not be strictly estuarine but edaphic conditions are comparable. A collection from south-west of Normanton (Jacobs 1194) would appear to have been made from an atypical habitat described as ‘ironstone pavement partially laterised, low open woodland with open shrub layer’. Typical habitat for C. halophila also occurs in the vicinity and although it is possible that the species extends into the habitat described, further field observations would provide clarification.

Phenology: Flowering and fruiting specimens have been collected in the field between February and September.

Affinities: Most specimens of Calandrinia halophila in Australian herbaria have hitherto been filed under Calandrinia indeterminate or an undescribed species; however, a few were determined as C. pumila (Benth.) F.Muell. Calandrinia pumila has a similar habit to C.. halophila but the leaf is more clearly differentiated into a lamina (1-10 mm wide) and long petiole, the lamina being lanceolate,

Albrecht & West, Calandrinia halophila 37

— ; “ i rs ¢

atte i | : - we ~~ ee - = ee , =

y ** o ae) S. : _

Fig. 1. Calandrinia halophila: A. habit. B. flower. C. seeds. D. capsule valves separating with subtending sepal (LHS); exposed placentas, funicles and seed with abscising sepal (RHS). Scale bar: B = 0.5 mm; C= 0.1 mm; D=0.5 mm. A, B & D from Brennan 11733 (DNA); C from Horsfall s.n. (CANB 954477). Photos C & D: J. FitzGerald.

38 Austrobaileya 13: 34—40 (2023)

» ae ie oy

=4 a" :. 7 .

-

Fig. 2. Typical habitat of Calandrinia halophila. Bushland Beach, Townsville (Horsfall s.n., CANB 955859). Photo: P. Horsfall.

Albrecht & West, Calandrinia halophila

elliptic, ovate, triangular or cordate, flower- bearing shoots usually distinctly exceeding the leaves and with more numerous (2—40+) flowers, opposite or rarely subopposite bracts, 5 petals, 3 stigmata, fruits that exceed the sepals and are basally circumscissile (lower part may split longitudinally but apex does not separate into valves), and slightly larger seeds (0.3—0.35 mm wide) with the swollen (but tiny) apex of the funicle persisting as a distinct strophiole. Calandrinia pumila occurs in more inland sites and is usually found in, or on the margins of temporary freshwater swamps, claypans and gilgais, and along intermittent non-saline watercourses and in run-on areas.

Dried herbarium specimens of Calandrinia halophila also resemble a species of rock pavements in eastern New South Wales and south-eastern Queensland recently described as C. petrophila J.G.West & Albr. However, that species differs from C. halophila in a range of features including being a perennial, having opposite bracts, fruiting pedicels |1.5—7 mm long, persistent sepals, 4 or 5 white petals (2.2—)3—6.5 mm long, longer (4—5.2 mm long) truncate capsules that split no deeper than halfway to the base into 4 persistent valves, and larger darker seeds (0.4—) 0.45—0.6 mm long, that are unevenly shiny and more prominently patterned.

Notes: Although molecular sequence data are currently unavailable for Calandrinia halophila, stigmata and petal number would suggest placement in clade 6 recognised by Hancock ef al. (2018). This clade aligns well with Calandrinia section Basales Poelln. and includes species with 4—6 stigmata and six or more petals. Calandrinia halophila difters from other species in clade 6 in having small petals, relatively few (5—10) stamens, and a unique combination of 5—7 stigmata and (5— )6—7(—8) petals.

Conservation status: Very little 1s known about the size and health of populations and threatening processes operating regarding Calandrinia halophila. This uncertainty 1s reflected in the current risk category assigned for the NT (data deficient) and WA (priority one poorly known). In the NT the species

39

occurs in Limmen National Park and in Queensland it occurs in Rinyirru (Lakefield) National Park. Significant disturbance caused by recreational activities is evident at one known site (Bushland Beach near Townsville) and appears to be impacting the currently extant population of C. halophila (P. Horsfall pers. comm.).

Etymology: The specific epithet is derived from Greek halo-, salt, and phileo, I love, in reference to the species preference for saline or semi-saline environments.

Acknowledgements

We thank Frank Obbens for providing information on specimens held at PERTH and comments on distinctiveness and affinities of Calandrinia halophila, Peter Horsfall for providing specimens, field observations and photographs, Kym Brennan for photographs and field observations, John FitzGerald for the seed photograph, Sally Mumford for preparing Fig. 1, Margarita Goumas for preparing the map, and the directors of BRI, DNA and NSW for loans and/or access to specimens.

References

Brown, G.K. (ed.) (2021). Portulacaceae. Census of the Queensland Flora 2021. Queensland Department of Environment and Science, Queensland Government. https://www.data. qld.gov.au/dataset/census-of-the-queensland- flora-2020, accessed 17 October 2022.

CHAH (2006). Australian Plant Census. https:// biodiversity.org.au/nsl/services/rest/reference/ apni/42942, accessed 20 December 2022.

Cowl, I.D., CUFF, N.J., LEwis, D.L. & JOBSON, P. (2017). Checklist of the vascular plants of the Northern Territory. Northern Territory Herbarium, Department of Environment and Natural Resources. http://eflora.nt.gov.au, accessed 20 December 2022.

Hancock, L.P., OBBENS, F., Moore, A.J., THIELE, K., DE Vos, J.M., West, J., HoLttum, J.A.M. & Epwarps, E.J. (2018). Phylogeny, evolution, and biogeographic history of Calandrinia (Montiaceae). American Journal of Botany 105: 1021-1034.

KERRIGAN, R.A. & ALBRECHT, D.E. (2007). Checklist of NT vascular plant species. Northern Territory Herbarium, Department of Natural Resources, Environment and the Arts: Darwin.

40

NORTHERN TERRITORY HERBARIUM (2015). FloraNT Northern Territory Flora Online. http://eflora. nt.gov.au, accessed 17 October 2022.

NYFFELER, R. & EGGLI, U. (2010). Disintegrating Portulacaceae: A new familial classification of the suborder Portulacineae (Caryophyllales) based on molecular and morphological data. Taxon 59: 227-240.

OBBENS, F.J. (2011). Five new species of Calandrinia (Portulacaceae) from Western Australia with additional information on morphological observations. Nuytsia 21: 1-23.

PARKER, C.M. & PERCY-Bower, J.M. (2022). Updates to Western Australia’s vascular plant census for 2021. Nuytsia 33: 2.

SHoRT, P.S., ALBRECHT, D.E., Cowie, I.D., LEwis, D.L. & STUCKEY, B.M. (2011). Checklist of the vascular plants of the Northern Territory. Northern Territory Herbarium, Department of Natural Resources, Environment and the Arts: Darwin.

THIELE, K.R., OBBENS, F., HANCOCK, L., EDWARDS, E. & WEST, J.G. (2018). (2587) Proposal to conserve the name Parakeelya against Rumicastrum (Montiaceae). Zaxon 67: 214-215.

‘ * — babe , \ 4 Q °°? ih ' ; - . . “hee Kae Darwin _@ tn” oe ( I me " iw Ne } ~» . f

> |

' t ‘ + My —we f 4 "s é

0 Borroloola 1

Austrobaileya 13: 34—40 (2023)

THOMAS, M.B. (2007). Portulacaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2007, p. 168. Queensland Herbarium, Environmental Protection Agency: Brisbane.

— (2010). Portulacaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2010, p. 162. Queensland Herbarium, Department of Environment & Resource Management: Brisbane.

— (2013). Portulacaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2013. — https://id.biodiversity.org.au/reference/ apni/51389804, accessed 20 December 2022.

— (2016). Portulacaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2016. https://id.biodiversity.org.au/instance/ apni/51390859, accessed 20 December 2022.

WEST, J.G. & ALBRECHT, D.E. (2022). A new species of Calandrinia (Montiaceae) from rocky pavements in eastern New South Wales and south-eastern Queensland. 7elopea 25: 323-

-~ apua - c - S nao! G Ul fo | : ae | ' Ng ra Port Moresby . * | . 6 ) rn J \. Cairns Ke O Oo ri | Burketown 4 e : o n . "Mackay : 0 500 7 T\s a 4 vi Kilometres dy

Map I. Distribution of Calandrinia halophila: based on specimens lodged at BRI, CANB, DNA, NSW and PERTH.

XA bacopterella altifrons (Thelypteridaceae), a new intergeneric fern hybrid from Australia

Thais Elias Almeida’, Alan R. Smith’, Peter D. Bostock’, Zoé Bloesch* & Ashley Raymond Field>*

Summary

Almeida, T.E., Smith, A.R., Bostock, P.D., Bloesch, Z. & Field, A.R. (2023). xAbacopterella altifrons (Thelypteridaceae), a new intergeneric fern hybrid from Australia. Austrobaileya 13: 41-50. The new nothogenus, xAbacopterella T.E.Almeida & A.R.Field and a new nothospecies, <Abacopterella altifrons T.E.Almeida & A.R.Field (Abacopteris aspera (C.Presl) Ching x Christella queenslandica (Holttum) A.R.Field & Z.Bloesch, comb. nov.) are described, based on recent phylogenomic evidence identifying the Australian ‘Russell River Fern’ to be an intergeneric hybrid. A new combination, Christella queenslandica (Holttum) A.R.Field & Z.Bloesch, accommodates findings of recent investigations of this species’ phylogenetic placement based on phylogenomic data. A description, illustrations, notes, and a conservation status assessment for the nothospecies are presented, as well as an updated list of all Thelypteridaceae occurring in Australia.

Key Words: Thelypteridaceae; <Abacopterella; <Abacopterella altifrons; Abacopteris; Christella; Christella queenslandica; flora of Queensland; flora of Wet Tropics; checklist of species; hybridisation; nothogenus; nothospecies; conservation status

‘Universidade Federal de Pernambuco, Centro de Biociéncias, Departamento de Botanica, Avenida Professor Morais Rego 1235, CEP 50.670—420, Recife, PE, Brazil; *The University Herbarium, University of California, Berkeley, 1001 Valley Life Sciences Building 2465, Berkeley, CA 94720- 2465, USA; °Queensland Herbarium and Biodiversity Science, Department of Environment and Science, Brisbane Botanic Gardens, Mt Coot-tha Road, Queensland 4060, Australia; “Australian Tropical Herbarium, James Cook University, PO Box 6811, Cairns, Queensland 4878, Australia.

*corresponding author: thais.elias@ufpe.br

Introduction

Reticulation is known to be a relevant process in evolution, connecting and promoting genetic exchange in otherwise isolated lineages (Dunning & Christin 2020). Hybridisation (Mallet 2007) and horizontal gene transfer (Dunning & Christin 2020) are the main processes that promote reticulation. In hybridisation, several outcomes are possible, including the formation of single or recurrent Fl hybrids, hybrid swarms or zones that can promote introgression, and the emergence of a new lineage of hybrid origin (Mallet 2007; Schumer ef al. 2014). Traditional species description based upon alpha-taxonomic species delimitation (as defined by Mayo et al. 2008) and in-use hierarchical categorical classification systems both fail to accommodate this evolutionary reality that lineages can reticulate. This is particularly relevant among ferns.

Reproductive barriers in_ ferns are suggested to evolve more slowly than in flowering plants, with multiple interspecific and intergeneric hybrids recognized (Barrington ef al. 1989; Rothfels et al. 2015; Liu et al. 2020). There are currently 32 described nothogenera, of which 12 are validly published and required if adopting the current classification of ferns (Liu ef al. 2020). Two belong to Thelypteridaceae: xChrismatopteris Quansah & D.S.Edward (Christella H.Lév. x Pneumatopteris Nakat) and xChrinephrium Nakaike (Christella H.Lév. x Pronephrium C.Presl).

Recently, and for the first time, an intergeneric hybrid in Thelypteridaceae was tested using phasing of genomic data, providing evidence of a hybrid between species of Abacopteris Fée and Christella (Bloesch et al. 2022). Thelypteridaceae is a

Accepted for publication 4 January 2023, published online 10 May 2023

© Queensland Herbarium 2023. ISSN 2653-0139 (Online)

42

cosmopolitan family with an estimated 1190 species and 37 genera (Fawcett & Smith 2021). The family is monophyletic (Almeida et al. 2016; Fawcett et al. 2021), with two main lineages, subfamilies Thelypteridoideae C.F.Reed and Phegopteridoideae Salino, A.R.Sm. & T.E.Almeida (PPG I 2016). Eleven genera and 23 species were recorded from Australia (Bostock 1998; Field 2020) with two species, Amblovenatum queenslandicum (Holttum) T.E.Almeida & A.R.Field and Chingia australis Holttum, considered endemic (Field 2020).

We examined new material = of Thelypteridaceae collected since the family was revised for Flora of Australia (Bostock 1998) and found two putative hybrids from the Australian tropics. One, named by Bloesch et al. (2022) as the ‘Russell River Fern’ has three recorded populations (Map 1; Bloesch et al. 2022). The other, named the ‘Tully River Fern’ (Bloesch et al. 2022) 1s known from a single locality. Using genomic data from nuclear loci, Bloesch et al. (2022) were able to confirm the hybrid origin of the two ferns and indicate the putative parents. The “Russell River Fern’ was found to be an F2, or later hybrid involving Abacopteris aspera (C.Presl) Ching and Amblovenatum queenslandicum, and the “Tully River Fern’ was considered likely to be an F1 hybrid between Christella subpubescens (Blume) Holttum and C. parasitica (L.) H.Leév.

Evidence from allele ratios of the sampled population indicates that the ‘Russell River Fern’ 1s more likely an F2 hybrid or a lineage of hybrid origin than a primary hybrid (Bloesch et al. 2022). The ‘Russell River Fern’ has been collected multiple times and is known from multiple sites; hence, this evidence warrants formal taxonomic recognition. Conversely, further research is needed to resolve the parentage of the “Tully River Fern’ and to review its identification as Amblovenatum tildeniae (Holttum) T.E.Almeida & A.R.Field (Field 2020).

Named taxa _ enable _ stability of scientific communication and recognition in biodiversity legislation and, in the case of nothotaxa, better reflect the uncertainty

Austrobaileya 13: 41-50 (2023)

around parentage and whether a plant 1s an Fl hybrid, complex hybrid, or species of hybrid origin. Therefore, we here describe a new nothogenus and nothospecies for the ‘Russell River Fern’. Although a nothospecies is often considered to be a primary hybrid, we chose to describe the ‘Russell River Fern’ as a nothospecies because only one of three populations has thus far been sampled phylogenomically, and whether the three populations are a cohesive lineage or resultant from separate hybridisation events remains unknown.

In addition to providing insights about reticulating evolution, the phylogenetic tree presented by Bloesch ef al. (2022) provided evidence that Amblovenatum queenslandicum 1s more appropriately placed in Christella (Bloesch et al. 2022) and a new combination 1n that genus 1s also made in this paper. An updated checklist for Australian Thelypteridaceae is presented (Appendix 1) following the classification of Fawcett & Smith (2021).

Materials and methods

We examined herbarium materials from the following herbaria: BRI, CNS, UC and US (all specimens cited have been seen), and material collected from wild populations throughout the north-eastern coast and hinterland of the Wet Tropics of Queensland, Australia. Living material was cultivated for four years in a tropical shade house in Cairns, Queensland, Australia, under 80% shade cloth, irrigated on alternating days and liquid fertilized on alternating weeks. Plants were grown in 250 mm diameter black plastic pots in a 1-5 mm 1:1 bark:scoria growing medium and were repotted biennially. Living plants were also examined in the private collection of Nada and Garry Sankowsky, Tolga, Australia. The terminology used to describe general morphology was based on Radford (1986) and Lellinger (2002).

Conservation status was assessed using IUCN Red List Categories and Criteria (IUCN 2018) to calculate the Extent of Occurrence (EOO) and the Area of Occupancy (AOO), using the GeoCAT tool (Bachman ef al. 2011)

Almeida et al., x Abacopterella altifrons

based on the records from the three locations (Map 1). A 2 km width grid cell was used for the estimation of AOO.

Taxonomy

Christella queenslandica A.R.Field & Z.Bloesch, comb. nov.

(Holttum)

Amphineuron queenslandicum Holttum, Kew Bull. 41:518 (1986); Thelypteris queenslandica (Holttum) Christenh., Global Fl. 4: 36 (2018); Amblovenatum queenslandicum (Holttum) T.E.Almeida & A.R.Field, Aust. Syst. Bot. 33: 68 (2020). Type: Australia: Queensland. Cook DISTRICT: State Forest Reserve 756, Kaaru Logging Area, Queensland, 4 September 1981, D.L. Jones 114 (holo: CNS [QRS 65258]; iso: BRI [AQ0411774]).

A species description can be sourced in the Flora of Australia account (Bostock 1998: 350).

Distribution and habitat: Christella queenslandica 1s an occasional fern in well- lit rainforest understory, near creeks, and long-term canopy gaps in lowland to lower montane rainforest. It 1s endemic to the wet tropical rainforests of northeast Queensland. By contrast, sympatric Christella dentata (Forssk.) Brownsey & Jermy and C. parasitica (L.) Lev. are much more abundant and tend to be pioneers on disturbed rainforest edges. Both C. dentata and C. parasitica have widespread tropical distributions.

Notes: Christella queenslandica (Tully River fern) 1s placed phylogenetically within the genus Christella and not within the genus Amblovenatum (Bloesch et al. 2022). Holttum (1986) does not discuss why it was placed in the genus Amphineuron (= Amblovenatum). It lacks the spherical yellow glands on the laminae characteristic of Amblovenatum and keys to the genus Christella in the generic keys of Bostock (1998) and Fawcett & Smith (2021). Christella queenslandica has similar fronds and rhizome to those of C. dentata, being distinguished in the field by its foliage which has a consistently spicy smell and mildly irritant exudate when crushed, more deeply incised pinna lobes, and a single pair of reflexed basal pinnae (compared to 2—4

43

pairs of reduced basal pinnae in C. dentata). Christella queenslandica has similar frond shape and division, spicy smell, and basal pinnae to C. parasitica, being distinguished by its erect rhizome (compared to horizontal creeping rhizome in C. parasitica).

xAbacopterella T.E.Almeida & A.R.Field, nothogen. nov.

A taxon of hybrid origin with parentage originating from species of Abacopteris Fée and Christella H.Lev.

xAbacopterella altifrons T.E.Almeida & A.R.Field, nothosp. nov.

A taxon of hybrid origin originating from Abacopteris aspera (C.Presl) Ching x Christella queenslandica (Holttum) A.R.Field & Z.Bloesch). It is similar to Abacopteris aspera in having anastomosing venation and short-creeping rhizomes. It differs from Abacopteris aspera by the larger, 1-pinnate- pinnatifid fronds, with 10—20 pairs of pinnae. It is similar to Christella queenslandica in having | pinnate-pinnatifid fronds, but differs in having anastomosing veins, broader pinnae, and short creeping rhizomes. Type: Australia: Queensland. COOK DISTRICT: Wooroonooran National Park, Russell River pack Track, 14 February 2018, A.R. Field 4646 & R. Jago (holo: BRI [AQ629845, comprising | sheet]; iso: BHCB, CNS 146459.1, UC).

Plants terrestrial. Rhizomes short-creeping; scales lanceolate, (4)5-12 mm wide, brown, glabrous. Fronds monomorphic to slightly dimorphic (fertile pinnae slightly narrower and more incised), 102—140 cm long; stipes 37—69 cm long, stipe bases covered with scales similar to those on the rhizome, abaxially with sparse narrow linear scales, adaxially sulcate with one-celled acicular trichomes. Laminae elliptic to ovate, chartaceous, 53-— 72 x 28-50 cm, 1-pinnate-pinnatifid, apical pinnae pinnatifid, not pinna-like, always widened and lobed towards their base, lateral pinnae 10—20 pairs, 20-26 =< 2.6-3.6 cm, perpendicular to ascending, proximal pair slightly deflexed, sessile, narrowly triangular to triangular, apex caudate, base truncate, pinna margins lobed, 1/4—1/3 towards costae; buds absent, aerophores round and slightly

44

raised, acroscopically at axils of pinnae; rachises abaxially with short, setiform hairs, 0.05—0.2 mm long, and capitate, stalked glandular hairs, 0.01—0.03 mm long, adaxially with acicular hairs, 0.18—0.4 mm long and glandular hairs, 0.01—0.03 mm long; costae abaxially rounded, with setiform hairs to 0.2 mm long, some to 0.5 mm, adaxially sulcate and covered with acicular hairs; costules, veins, and blades abaxially bearing setiform hairs, adaxially glabrous; veins 10-14 pairs per segment, at 60° to costule, mostly anastomosing and forming parallel rows of areoles below (proximal to) the sinuses, the basal-most areoles joining to form excurrent veinlets that join the next row of areoles, veins free on the lobes; segments 4.1—5.1 mm wide, apices rounded. Sori round, medial, indusiate; indusia conspicuous, round-reniform, entire, with setiform hairs to 0.2 mm long; sporangia setose. Spores variable, with three observed types: 50-60 um _ diameter, spherical, unreduced spores (rare), 30—40 um long, monolete, winged spores (rare); and 30—40 um collapsed monolete unwinged spores (predominant type). Figs. 1 & 2.

Additional specimens examined: Queensland. Cook District: National Park 226, Bartle Frere, first Combo Russell River pack trail, Jun 1993, Bostock 1414 & Jago (BRI, NSW); ibid, Sep 1993, Bostock et al. 1457 (BRI, NSW); ibid, ex plant cultivated at Sankowsky living plant collection, Tolga, ex Bostock 1414 & Jago, Nov 2015, Field 3918 (BRI, CNS, UFP). NorTH KENNEDY District: Porter’s Creek, off Bruce Highway, NW of Ingham, May 1992, Cumming 11952 & Thomas (BRI).

Distribution and habitat: *Abacopterella altifrons (Russell River Fern) is endemic to Australia in the Wet Tropics of Queensland. It has been documented from two locations, near the Russell River pack track in the Innisfail region, and Porter’s Creek in the Hinchinbrook region and also reported from Melele Creek (7ng s.n., Daintree Research Observatory Field Herbarium) and a tributary of Bailey Creek (Nada Sankowsky, pers. comm.) in the Daintree—Bloomfield region.

xAbacopterella altifrons 1s an extremely rare, terrestrial fern that forms small colonies in brightly lit areas on forest edges, tracks, or in long-term canopy gaps in lowland mesic complex mesophyll vineforests. This

Austrobaileya 13: 41-50 (2023)

species tends to form localized multi-plant colonies with each plant bearing three to nine fronds; the plants generally long-lived. New plant establishment has been observed both from rhizome division and apparently from spores, evidenced by plants appearing at sites not connected to established plants. The population near the Russell River pack track has been monitored for over 30 years (R.L. Jago pers. comm.; A.R. Field pers. obs.) and has persisted in the same general area through severe canopy disturbance caused by Tropical Cyclones Winifred (1986), Larry (2006) and Yasi (2011).

Notes: *Abacopterella altifrons occurs sympatrically with Abacopteris aspera, from which it differs by the higher number of pinna pairs (10—20 pairs in <Abacopterella altifrons vs. up to eight pairs in Abacopteris aspera), the incised lobulate pinna margins (vs. entire to crenulate in Abacopteris aspera), and the pinnatifid apical pinna (vs. conform in Abacopteris aspera). Mature plants of <Abacopterella altifrons generally bear numerous fronds and form a larger and localized colony, differing from mature Abacopteris aspera which bear few fronds and usually are isolated plants scattered throughout the forest floor. Fronds of xAbacopterella altifrons are more erect and paler green than Abacopteris aspera, which has fronds that are more spreading and usually a darker green. These traits were maintained when plants were cultivated side- by-side under uniform conditions.

xAbacopterella altifrons also occurs sympatrically with Christella queenslandica, from which it differs by its short-creeping rhizomes with a loose cluster of fronds (vs. erect with a vase-shaped rosette of fronds in C. queenslandica) and anastomosing veins (vs. free veins in C. qgueenslandica, except for costal areoles formed by basal veins from adjacent segments uniting and forming an excurrent vein toward the sinus). xAbacopterella altifrons 1s a brighter yellow- ereen than C. gueenslandica and also lacks the spicy aroma and irritating exudate sometimes recorded in C. queenslandica.

Almeida et al., xAbacopterella altifrons

Fig. 1. xAbacopterella altifrons. A. habit. B. abaxial view of a fertile frond, showing non-conform apex. C. adaxial side of pinnae. D. abaxial side of a pinna, showing soral pattern. E. abaxial side of sterile (left) and fertile (right) pinnae, showing venation pattern. A—E taken in habitat at location of Field 4646 (CNS).

46 Austrobaileya 13: 41—50 (2023)

Fig. 2. xAbacopterella altifrons. A & B. abaxial view of a fertile frond, showing overall aspect and details of indument, sori disposition, and venation. C & D. adaxial view of a fertile frond, showing overall aspect and details of indument. E. sori, showing acicular hairs on the indusium. Scale bar: A—D: 0.5 cm. E: 0.5 mm. All from Field 4646 (BHCB). Del.

Keven S. Lima.

Almeida et al., xAbacopterella altifrons

Spore production was observed in both wild specimens and cultivated plants. Although we did not test for viability, most spores are malformed and appear inviable. New, apparently spore-grown individuals, have been observed in the wild and once in cultivation (A.R. Field, pers. obs.), but we do not know if these are the result of sexual or apomictic reproduction or are de novo hybrids with the same parentage. Both Abacopteris and Christella have x=36 (n=72) (Tindale & Roy 2002), and future studies should focus on investigating the ploidy level to clarify if any of the populations are allopolyploid (Bloesch et al. 2022). A survey on the genetic structure of known populations of xAbacopterella altifrons 1s also recommended to assess the genetic divergence and understand its sexual and asexual reproductive processes.

Etymology: *Abacopterella_ altifrons 1s named for having tall fronds, in comparison with the species from the parental genera (Abacopteris and Christella).

Conservation Status: Hybrids are not included in IUCN Red Lists (UCN 2022a) with their presence considered a species stressor (IUCN 2022b); however, this policy does not explicitly cover nothospecies. This policy of not applying IUCN conservation status to hybrid individuals has been criticised for potential conservation decision bias (Bauer ef al, 2021).

For the purposes of conservation assessment, we here consider _ that xAbacopterella altifrons 1s not a primary hybrid (Bloesch eft al. 2022); hence it 1s eligible for conservation status assessment.

The species has an EOO of 29,673 km? and an AOO of 12.000 km’, and which respectively meet the CR and EN categories under IUCN criteria IUCN 2019). Although the species is known from protected areas, populations observed in the field have a very small number of individuals. <Abacopterella altifrons 18 maintained in four ex situ cultivated collections in Australia.

47 Acknowledgments

We thank Nada and Garry Sankowsky of Tolga for providing access to their superb living collection of Thelypteridaceae, Robert Jago of Cairns for providing specimens, advice on field localities, and support in fieldwork, and two anonymous reviewers for their comments that helped improve this manuscript. TEA thanks CNPq for the research grant (317091/2021-2) and NSF for support through erant DEB1456232 to Benjamin Torke (the New York Botanical Garden). ARF was supported by a Queensland-Smithsonian Fellowship 2017 to US and UC herbaria. TEA and ARF’s fieldwork in Australia was supported by Queensland Department of Environment and Science’s core-funding. Illustrations were drawn by K.S. Lima.

References

ALMEIDA, I.E., HENNEQUIN, S., SCHNEIDER, H., SMITH, A.R., BATISTA, J.A.N., RAMAHO, A.J., PROITE, K. & SALINO, A. (2016). Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alterations of neotropical taxa and further study of paleotropical genera. Molecular Phylogenetics and Evolution 94: 688-700. DOI: 10.1016/j. ympev.2015.09.009

BACHMAN, S., Moat, J., DE LA TORRE, J. & SCOTT, B. (2011). Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. ZooKeys 150: 117-126. DOL: 10.3897/zookeys.150.2109

BARRINGTON, D.S., HAUFLER, C.H., WERTH, C.R. (1989). Hybridization, reticulation, and species concepts in the ferns. American Fern Journal 79: 55—64. DOI: 10.2307/1547160

BAUER, H., TEHOU, A.C., GUEYE, M., GARBA, H., DOAMBA, B., DIouCK, D. & SILLERO-ZUBIRI, C. (2021). Ignoring species hybrids in the IUCN Red List assessments for African elephants may bias conservation policy. Nature Ecology & Evolution 5: 1050-1051.

BLOESCH Z., NAUHEIMER, L., ALMEIDA, I.E., CRAYN, D. & FreL_p, A.R. (2022). HybPhaser identifies hybrid evolution in Australian thelypterid ferns. Molecular Phylogenetics and Evolution 173: 107526. DOI: 10.1016/j.ympev.2022.107526

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Bostock, P.D. (1998). Thelypteridaceae. In P.M. McCarthy (ed.), Flora of Australia 48: 327-358. ABRS/ CSIRO: Melbourne.

DUNNING, L.T. & CHRISTIN, P.-A. (2020). Reticulate evolution, lateral gene transfer, and innovation in plants. American Journal of Botany 107(4): 1—4. DOI: 10.1002/ajb2.1452

FAWCETT, S., SMITH, A.R., SUNDUE, M., BURLEIGH, J.G., SESSA, E.B., Kuo, L.-Y., CHEN, C.-W. TESTO, W.L., KESSLER, M., GOFLAG CONSORTIUM & BARRINGTON. D.S. (2021). A global phylogenomic study of the Thelypteridaceae. Systematic Botany 46: 891-915. DOI: 10.1600/ 036364421 X16370109698650

FawceTT, S. & SMITH, A.R. (2021). A Generic Classification of the Thelypteridaceae. Sida, Botanical Miscellany 59. BRIT Press, Botanic Garden | Botanical Research Institute of Texas, U.S.A.: Fort Worth.

Fretp, A.R. (2020). Classification and _ typification of Australian lycophytes and ferns based on Pteridophyte Phylogeny Group classification PPG I. Australian Systematic Botany 33: 1-102. DOI: 10.1071/SB18011

HoLttum, R.E. (1986). The genus Christella Léveilleé, sect. Christella. Studies in the Family Thelypteridaceae, XI. Kew Bulletin 31: 293-339.

IucN (2018). Guidelines for Using the IUCN Red List Categories and Criteria, version 2; http://jr.tucnredlist.org/documents/ RedListGuidelines.pdf, accessed 19 Sep 2020.

—— (2022a). Taxonomic Sources. https://www. lucnredlist.org/resources/tax-sources, accessed 23 November 2022.

— (2022b). Stresses Classification Scheme (Version 1.0). https://www.iucnredlist.org/resources/

stresses-classification-scheme, accessed 23 November 2022.

LELLINGER, D.B. (2002). A modern multilingual glossary for taxonomic pteridology. Pteridologia 3: 1-263. DOI: 10.5962/bh1.title.124209

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Liu, H.-M., SCHUETTPELZ, E. & SCHNEIDER, H. (2020). Evaluating the status of fern and lycophyte nothotaxa in the context of the Pteridophyte Phylogeny Group classification (PPG 1). Journal of Systematics and Evolution 58(6): 988-1002.

MALLET, J. (2007). Hybrid speciation. Nature 446: 279-— 283. DOI: 10.1038/nature05706

Mayo, S., ALLKIN, R., BAKER, W., BLAGODEROV, V., BRAKE I., CLARK, B., GOVAERTS R., GODFRAY C., HAIGH A., HAND R., HARMAN K., JACKSON M., KILIAN N., Kirkup D.W., KITCHING L., KNAPP S., LEwis G.P., MALCOLM P., RAAB- STRAUBE E. VON, ROBERTS D. M., SCOBLE M., SIMPSON D.A., SMITH C., SMITH V., VILLALBA S., WALLEY L. & WILKIN P. (2008). Alpha E-Taxonomy : Responses from the Systematics Community to the Biodiversity Crisis. Kew Bulletin 63: 1-16. DOI: 10.1007/s12225-008- 9014-1

Pec I (2016) A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54(6): 563-603. DOI: 10.1111/jse.12229

RADFORD, A.E. (1986). Fundamentals of Plant Systematics. Harper & Row: New York.

ROTHFELS, C.J., JOHNSON, A.K, HOVENKAMpP, P.H., SWOFFORD, D.L., ROSKAM, H.C., FRASER- JENKINS, C.R., WINDHAM, M.D. & PRYER, K.M. (2015). Natural hybridization between genera that diverged from each _ other approximately 60 million years ago. American Naturalist 185: 433-42. https://www.jstor.org/ stable/10.1086/679662

SCHUMER, M., ROSENTHAL, G.G., ANDOLFATTO, P. (2014). How common is homoploid hybrid speciation? Evolution 68: 1553-1560. DOL: 10.11 11/evo.12399

TINDALE, M. & Roy, S. K. (2002). A cytotaxonomic survey of the Pteridophyta of Australia. Australian Systematic Botany 15: 839-937.

Almeida et al., x Abacopterella altifrons 49

J ; | 144.00°E

L eich

’ i,

= 18.00°S :

~147.00°E

oi XY L | | — io ‘ -

Map 1. Distribution of xAbacopterella altifrons. The Extent of Occurrence (EOO) and the Area of Occupation (AOO) were calculated based on these records.

50 Austrobaileya 13: 41—50 (2023) Appendix 1. An updated list of Australian Thelypteridaceae according to the classification of Fawcett & Smith

(2021). Superseded names listed in Field (2020) are presented in square brackets. * putative hybrid identity based on Bloesch ef al. (2022).

Subfamily Phegopteridoideae Salino, A.R.Sm. & T.E.Almeida Macrothelypteris polypodioides (Hook.) Holttum Macrothelypteris torresiana (Gaudich.) Ching

Subfamily Thelypteridoideae C.F.Reed xAbacopterella altifrons T.E.Almeida & A.R.Field Abacopteris aspera (C.Presl) Ching [= Pronephrium asperum (C.Presl) Holttum] Amblovenatum immersum (Blume) Parris Amblovenatum opulentum (Kaulf.) J.P.Roux Amblovenatum terminans (Hook.) J.P.Roux * Amblovenatum tildeniae (Holttum) T.E.Almeida & A.R.Field [? Christella subpubescens (Blume) Holttum =< Christella parasitica (L.) H.Lév] Ampelopteris prolifera (Retz.) Copel. Chingia australis Holttum Christella arida (D.Don) Holttum Christella dentata (Forssk.) Brownsey & Jermy Christella hispidula (Decne.) Holttum Christella parasitica (L.) H.Lev. Christella queenslandica (Holttum) A.R.Field & Z.Bloesch |= Amblovenatum queenslandicum (Holttum) T.E.Almeida & A.R.Field] Christella subpubescens (Blume) Holttum Cyclosorus interruptus (Willd.) H.Ito Grypothrix triphylla (Sw.) S.E.Fawc. & A.R.Sm. [= Pronephrium triphyllum (Sw.) Holttum | Pakau pennigera (G.Forst.) S.E.Fawc. & A.R.Sm. [= Pneumatopteris pennigera (G.Forst. | Holttum) Plesioneuron tuberculatum (Ces.) Holttum Reholttumia costata (Brack.) S.E.Fawc. & A.R.Sm. [= Pneumatopteris costata (Brack.) Holttum] Reholttumia sogerensis (Gepp) S.E.Fawc. & A.R.Sm. [= Pneumatopteris sogerensis (A.Gepp) Holttum] Reholttumia truncata (Poir.) S.E.Fawc. & A.R.Sm. [= Pneumatopteris truncata (Pott.) Holttum] Sphaerostephanos heterocarpos (Blume) Holttum Strophocaulon unitum (L.) S.E.Fawc. & A.R.Sm. [= Sphaerostephanos unitus (L.) Holttum] Thelypteris confluens (Thunb.) C.V.Morton

Two new species of Enteropogon Nees (Poaceae: Chloridoideae: Cynodonteae: Eleusininae) for northern Australia

E.J. Thompson

Summary

Thompson, E.J. (2023). Two new species of Enteropogon Nees (Poaceae: Chloridoideae: Cynodonteae: Eleusininae) for northern Australia. Austrobaileya 13: 51—93. Two new species, Enteropogon pubifolius E.JVhomps. and &£. scabrilemma E.J.Thomps., with morphological similarities to E. paucispiceus (Lazarides) B.K.Simon and E. macrostachyus (Hochst. ex A.Rich.) Munro ex Benth., respectively, are described and illustrated. Enteropogon pubifolius differs from other Australian species by a combination of characteristics including the pubescent, broader basal leaves, and E. scabrilemma by the scabrid broader lemmas and inflorescences with usually one or two rigid racemes. Some morphological characters used to identify Australian species of Enteropogon were found to be variable and consequently, a new modified key was developed to reduce potential ambiguity. Results from phenetic analyses of 43 morphological characters were used to explore hypotheses for the taxonomic relationships of the species.

Key Words: Poaceae; Chloridoideae; Cynodonteae; Eleusininae; Chloris; Enteropogon; Enteropogon dolichostachyus; Enteropogon macrostachyus; Enteropogon paucispiceus; Enteropogon pubifolius; Enteropogon scabrilemma; flora of Australia; flora of Queensland; taxonomy; new species; identification key; anatomy; micromorphology; cork cells; paraligule; stipe

E.J. Thompson, c/o 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 chloridoid genus Enteropogon Nees 1s represented by 17 species, commonly known as windmill grasses, distributed across Africa, Asia and Australia (Stapf 1934; Watson & Dallwitz 1992; Nightingale eft al. 2005; Peterson ef al. 2015; Simon & Alfonso 2022; Tropicos 2022). Six species have been recorded for Australia, four of which are endemic. Two of the Australian species are exclusively tropical, including Enteropogon dolichostachyus (Lag.) Keng ex Lazarides, a species that is also recorded across parts of Asia. Two species occur along the Australian central east coast including Enteropogon unispiceus (F.Muell.) Clayton that 1s also found in Taiwan and the Cook Islands. The other two species are widespread across the Australian arid and semi-arid zones (Lazarides 1972; Nightingale et al. 2005; Simon & Alfonso 2022; Map 1).

The taxonomic composition of Enteropogon, formerly included in Chloris Sw., has changed somewhat since its initial description. New species or combinations in Enteropogon (E. coimbatorensis K.K.Nair, S.K.Jain & M.P.Nayar, E. paucispiceus (Lazarides) B.K.Simon and FE. ramosus B.K.Simon) have occurred (Nair et al. 1977; Simon 1984), and several species were segregated into two genera 7etrapogon Dest. and Enteropogonopsis Wipft & R.B.Shaw, (Lazarides 1972; Anderson 1974; Tothill & Hacker 1983; Jacobs & Highet 1988; Peterson ef al. 2015). Characters used to distinguish Enteropogon and related taxa include leaf anatomy and micromorphology, and morphology of spikelets, inflorescences and leaf sheaths (Lazarides 1972; Carolin & Jacobs 1973; Anderson 1974; Tothill & Hacker 1983; Watson & Dallwitz 1992; Prendergast & Hattersley 1987; Wipft & Shaw 2018).

Accepted for publication 29 March 2023, published online 28 June 2023

© Queensland Herbarium 2023. ISSN 2653-0139 (Online)

52

Enteropogon, like most of the chloridoid erasses, exhibits Kranz anatomy but differs from the C, type known in Chloris (Prendergast & Hattersley 1987; Watson & Dallwitz 1992). Comparative micromorphology of chloridoid grasses that includes £. dolichostachyus have been presented by Anderson (1974) and Liu ef al. (2010). Anderson (1974) provided illustrations of bicellular micro- hairs and silica bodies of several genera with morphological similarities to Chloris. Lemmatal micromorphology and scanning electron micrographs (SEMs) of chloridoid cork cells have been provided by several authors including Valdes-Reyna & Hatch (1991), Snow (1996) and Liu ef al. (2010).

The breeding system of Enteropogon involves _cleistogamy _—_ (Self-fertilisation within a closed flower) (Campbell ef al. 1983; Culley & Klooster 2007; Thompson 2021). Enteropogonopsis chloridea (J.Presl) Wipfl & R.B.Shaw (syn. Enteropogon chlorideus (J.Presl.) Clayton) exhibits a very rare syndrome that involves subterranean spikelets borne on rhizomes (rhizanthogenes); these are morphologically differentiated from the spikelets in the terminal inflorescences that are chasmogamous as commonly found in the stereotypical grasses (Campbell ef al. 1983). All Australian species of Enteropogon have terminal inflorescences with both cleistogamous and chasmogamous spikelets of similar morphology (Thompson 2021). The spikelets of Enteropogon disarticulate above the glumes and the diaspore consists of united florets (one or two fertile proximal and one sterile distal), with passive awns (Jurado et al. 1991; Watson & Dallwitz 1992; Cavanagh et al. 2019).

Recent curation of Enteropogon specimens held at the Queensland Herbarium (BRI) revealed two new species, wiz. E. scabrilemma E.J.Thomps. and E. pubifolius E.JThomps., that are described here. The specimens of E. scabrilemma were stored under EF. dolichostachyus, but the morphology of the spikelets more closely resembles £. macrostachyus (Hochst. ex A.Rich.) Munro ex Benth from tropical Africa. Enteropogon scabrilemma and E. macrostachyus ditter

Austrobaileya 13: 51—93 (2023)

from LE. dolichostachyus by several characters including broader scabrid lemmas (Table 1). Enteropogon pubifolius was stored as an undetermined species differing most conspicuously by the hairy leaves, an unusual occurrence in the Australian species (Simon & Alfonso 2022). Enteropogon pubifolius is considered to have close morphological similarity to £. paucispiceus as they share some characters including basal leaves, a character used 1n the identification key to the Australian species of Enteropogon by Simon & Alfonso (2022) (Table 1).

Materials and methods

This study explores multiple hypotheses about the delimitation of species in Enteropogon, including some generated from phenetic analyses for the purpose of aiding accountability, repeatability and objectivity in taxonomic decision-making. The robustness of the hypotheses generated as a result of the analyses was tested using variations in data inputs. This study also reviews the morphological affinities and discriminating characters of the species and examines some distinctive characters of Enteropogon in the context of Poaceae.

Nomenclature, terminology and circumscription

Botanical nomenclature complies’ with

Thompson (2022a) and Tropicos (2022).

General taxonomic concepts conform with Stuessy (2009), and Enteropogon species concepts follow those of Lazarides (1972) and Simon (1984).

Etymology follows Clifford & Bostock (2007).

General botanical terminology follows Harris & Harris (1994), Henslow (2009) and Beentje (2010). Terminology relating to inflorescences and spikelets follows Tothill & Hacker (1983), Jacobs et al. (2008), Gibson (2009) and Thompson (2021). The spikelet is a reduced inflorescence with florets subtended by bracts, viz. glumes, lemmas and paleas (Kellogg 2006; Endress 2010).

Thompson, New species of Enteropogon

Most of the terminology relating to grass anatomy and micromorphology used here follows Ellis (1976, 1979) and Dengler ef al. (1994). Bicellular micro-hairs have sometimes been referred to as salt glands as in the case of Enteropogon macrostachyus (Liphschitz & Waisel 1974; Jacobs 1986; Ceccoli et al. 2015). Diaspore stipe, lemma, rachilla and callus are depicted in Fig. 1 for Enteropogon species. Diaspore stipe is defined as the portion of the diaspore from the distal tip of the callus to the junction of the rachilla and first lemma (Fig. 1). The chloridoid stipe has broad similarities to that found in some panicoid grasses but is distinct from a diaspore that is stipitate (Hitchcock 1950; Zuloaga 1986; Freckmann & Lelong 2003; Aliscioni ef al. 2016; Zuloaga et al. 2018; Thompson & Fabillo 2021; Thompson 2022b,c). Due to a tendency for immature or aborted spikelets to have higher variability in size, shrinkage or distortion, spikelet measurements presented in the diagnoses were made from specimens with mature spikelets bearing caryopses.

Silica bodies (also referred to as phytoliths by some authors), stomata, trichomes (such as bicellular micro-hairs), prickles and hooks were Classified using information provided by Ellis (1979), Amarasinghe & Watson (1990), Watson & Dallwitz (1992), Renvoize (2002), Rugolo de Agrasar & Vega (2004) and Neumann eft al. (2017).

Cork cells, comprising a silica body and lid, on first lemmas were evaluated using four criteria viz., spacing, separation, size and shape (Metcalfe 1960; Ellis 1979; Valdes- Reyna & Hatch 1991; Acedo & Llamas 2001; Fig. 2). Metcalfe (1960) and Ellis (1979) reported cork cells on leaves and Valdes- Reyna & Hatch (1991) and Acedo & Llamas (2001) studied cork cells on lemmas, with the latter authors using characters including shape, density and length.

Classification of epicuticular wax follows Barthlott et al. (1998) and Ortunez & de la Fuente (2010). The paraligule is defined here as arow of hairs behind the ligule (Fig. 3).

53 Taxon sampling

Sampling included the eight Australian native species and one African species of Enteropogon and two species of Chloris. Species data was obtained from numerous accessions across the geographic distribution for each species. Forty-five specimens of E. dolichostachyus trom Australian herbaria, collected from Australia, Sri Lanka, Papua New Guinea, Philippines and Thailand, were examined and specimens cited by other authors are listed in Appendix 1. The specimens of EF. macrostachyus examined are also listed in Appendix 1. Two specimens of E. scabrilemma (Blake 23488 and Pullen 6529), have been cited by other authors including Lazarides (1972), Nightingale ef al, (2005) and Peterson ef al. (2015) in their synopses of E. dolichostachyus.

Chloris divaricata R.Br. and Chloris ventricosa R.Br. were selected as_ the outgroup with the former having resemblance to Enteropogon at least to the naked eye, with the consequence that occasionally some specimens at BRI have been misidentified.

Cultivation of plants

All species, except Enteropogon macrostachyus, were cultivated in pots under the same nursery conditions in full sun at Brisbane, Australia during the period 2009-2022. Plants were propagated from five or SIX caryopses per species acquired from herbarium specimens, except E. macrostachyus for which no viable seeds were available (Appendix 2).

Specimens of Enteropogon acicularis (Lindl.) Lazarides held at BRI display some variation, particularly in_ inflorescence structure. To explore plasticity, plants were propagated from caryopses taken from accessions of three provenances. The three accessions were chosen because they provided viable caryopses.

Austrobaileya 13: 51—93 (2023)

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56 Austrobaileya 13: 51—93 (2023)

jad floret

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‘\, rachilla broader

Fig. 1. Diaspores of Enteropogon dolichostachyus, E. macrostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma from ventral and lateral views. The diaspore comprises the united florets that have disarticulated leaving the glumes attached to the pedicel. A, C, E, G & J lateral views; B, D, F, H & I ventral views. A & B. E. paucispiceus. C & D. E. dolichostachyus. E & F. E. pubifolius. G & H. E. scabrilemma. | & J. E. macrostachyus. A & B from Thompson MORS42 (BRI); C & D from Thompson MORS37 (BRI); E & F from Thompson MORS4/ (BRI); G & H from Thompson MORS44 (BRI); 1 & J from Davidse 5886 (BRI). Photos: E.J. Thompson.

Thompson, New species of Enteropogon

The cultivated plants provided fresh material for transverse sectioning, and studies of inflorescence morphology, breeding systems, flowering and fruiting phenology, and growth habit.

Imagery

Photographs were taken using _ light microscopes, viz. Nikon SMZ25 binocular microscope with Nikon DS-Ril camera and images viewed using NIS-Elements BR (2020) and Leica DMLB compound binocular microscope with an industrial digital camera and images viewed using ToupView (2020). Potted plants were photographed using a Fuji FinePix SL280 digital camera.

Scanning electron micrographs (SEMs) were used to examine the surface micromorphology of lemmas, paleas, leaves, culms and caryopses. SEMs were obtained without sputter coating the specimens, using a Phenom G2 Skev scanning electron microscope with backscatter detector.

Type specimens were examined using digital images obtained from JSTOR Plants (2022).

Acquisition of data and classification of morphological characters

Three main groups of morphological characters were examined and classified for this study. These include spikelets (macro- and micromorphology of glumes, lemmas, paleas, style and caryopsis, leaves (including ligule, paraligule, anatomy of transverse sections, micromorphology of the abaxial surface) and culms (anatomy and surface micromorphology).

Characters and character states used by other authors were reviewed and further information was gathered from observation of herbarium specimens and cultivated plants (Appendix 3). A broad spectrum of characters was examined, and information relating to character states and plasticity was obtained from character lists, diagnostic descriptions, illustrations and keys used by other authors including Lazarides (1972), Anderson (1974) Nair et al. (1977), Phillips (1982), Davis (1983), Tothill & Hacker (1983), Jacobs

57

(1986), Jacobs & Highet (1988), Lazarides et al. (1992), Watson & Dallwitz (1992), Van den Borre & Watson (1997), Nightingale ef al. (2005), Liu et al. (2010), Giraldo-Cafias et al, (2012), Fahey et al. (2019), and Simon & Alfonso (2022).

Characters and character states were evaluated in terms of similarity, ambiguity, reliability, plasticity, practicality, repeatability and standardisation of assessment or measurement using considerations by other authors (Hillis 1987; Wagner 1989; Smith 1990; Lipscomb 1992; Hillis & Wiens 2000; Poe & Wiens 2000; Rieppel & Kearney 2002; Scotland et al. 2003; Wiens 2004; Smith & Turner 2005; Thompson & Fabillo 2021; Thompson 20225, c).

Spikelet morphology

Micromorphology of first lemmas was studied using SEMs to classify silica bodies, cork cells, stomata, and macro- and micro-hairs using information provided by Hsu (1965), Anderson (1974), Ellis (1979), Valdes-Reyna & Hatch (1991), Snow (1996), Columbus (1999), Acedo & Llamas (2001), Liu ef al. (2010), Mashau ef al. (2015) and Olonova ef al. (2016) (Appendix 3). SEMs were captured near the mid-point of mature first lemmas (bearing a caryopsis).

The morphology of caryopses included shape and size, length of scutellum, shape of hilum, embryo spermaderm, and stylopodium, as used by Reeder (1957), Jacobs & Highet (1988), Watson & Dallwitz (1992), Terrell & Peterson (1993), Soreng & Davis (1998), Snow (1998), Liu et al. (2005), Gandhi et al. (2013) and Zhang et al. (2014).

Leaf and inflorescence culm anatomy and surface micromorphology

Leaf and culm materials from cultivated plants and herbarium specimens were used to obtain transverse sections with the method described by Thompson (2017) as modified from Frohlich (1984). Mature leaves were selected from the upper parts of culms and sections were taken from near the middle of the leaves. Culm sections were taken from a portion just below the first inflorescence branch. Leaf and

58

culm transverse sections were classified by characters from the sclerenchyma, vascular bundles and parenchyma building on the information (including illustrations) provided by authors including De Wet (1960), Ellis (1976), Clifford & Watson 1977, Dengler et al. (1994), Columbus (1999), Renvoize (1983, 2002), Siqueiros-Delgado (2007) and Ahmad et al. (2012).

Leaf surface micromorphology was also studied from replicas of the abaxial surface of fresh leaves acquired using the method described by Hilu & Randall (1984). To improve ease of application, acrylic nail varnish was diluted to about 60% using acetone.

Datasets and phenetic analyses

The robustness of results from _ cluster analyses was explored by varying inputs, viz. sample composition, algorithm (ordination, classification, association measure), as well as the format of characters. This approach was adopted following findings made by other authors (Clifford & Goodall 1967; Mannetje 1967; Clifford & Williams 1973; Austin & Belbin 1982; Hilu & Wright 1982; Johnson 1982; Stevens 1991; Thiele 1993; Wills ef al. 2000; Scotland et al. 2003; Wortley et al. 2005; Pereira et al. 2007; Newmaster et al. 2008; Zuloaga et al. 2014; Peichoto et al. 2015; Aliscioni et al. 2016; Thompson 2022b,c).

The baseline dataset was composed of 11 samples of Enteropogon species and 43 morphological characters (Appendix 4). The characters comprised nine vegetative characters, 41 characters relating to spikelets and inflorescences and seven relating to leaf anatomy (Appendix 3). The dataset consisted of 32 binary and |] multistate characters.

Cluster analyses were conducted using PATN 4.00 (Belbin & Collins 2013). Classification analyses used the unweighted pair method with arithmetic mean (UPGMA), and dendrograms were generated using agglomerative hierarchical fusion with unweighted pair group method with a Beta value of -0.1. Two association measures were used to examine the potential impact of assumptions about the nature of the states

Austrobaileya 13: 51-93 (2023)

of polymorphic characters. Of the several association measures available in the PATN package, Gower metric and Czekanowsk1 (Bray-Curtis) (Somerfield 2008) were applied where differences between states of polymorphic characters are considered equal and unequal, respectively (L. Belbin pers. comm.). Three-dimensional ordination plots were generated using semi-strong hybrid multidimensional scaling (SSH). Ordination stress value (OSV) was used as the measure of closeness of fit: stress values of <0.05 = excellent, <0.1 = good, <0.1—0.15 may be OK, <0.2 = not good (Belbin https://patn.org/ PATN - Finding patterns 1n data/, accessed 17 February 2022). Discriminating characters were generated for analysis of the baseline dataset from Kruskal-Wallis (KW) values (Appendix 3).

Three tests of impact, based on the methods used by Thompson (2022b, c), were conducted for comparison with results from analyses of the baseline dataset. In test 1, all multistate characters were transformed to binary format resulting in 72 characters. In test 2, four datasets in addition to the baseline were established from a list of the most discriminating characters in descending order based on KW values generated from analysis of the baseline dataset with the number of eroups set at 10, corresponding to the nine putative species used in the sample. Datasets were created by successively removing batches of 5 characters with the lowest KW values starting from 40 characters, followed by 35, 30, and so on. During test 3, the effect of an outgroup was assessed by adding the two species of Chloris to the baseline dataset resulting in a data matrix of 13 samples and 58 characters, of which 13 were multistate (Appendix 4). This dataset was analysed and the test on discriminating characters (test 2) was carried out with the number of groups set at nine and datasets with batches of five characters removed. The multistate characters were transformed to binary resulting in 87 characters.

Thompson, New species of Enteropogon Results

Topologies generated from analyses were affected by the data format and the algorithm, but the addition of Chloris to the sample as an outgroup had little impact (Figs. 4 & 5, Table 3). Four clusters were consistently recovered from all topologies, viz. Chloris spp., Enteropogon minutus with E. unispiceus, E. macrostachyus with E. scabriblemma, and E. paucispiceus with E. ramosus (Fig. 4). In ordination topologies, EF. paucispiceus, E. pubifolius and E. ramosus were commonly erouped. Enteropogon dolichostachyus was typically grouped with E. macrostachyus and E. scabriblemma in ordination topologies, but only as such in dendrogram topologies generated from analyses using Czekanowsk1 association measure (Figs. 4 & 5).

The top 20 most discriminating characters generated from cluster analysis of the baseline dataset covered a broad range of character types (Table 1; Appendix 5). Several of these characters are new to studies of Enteropogon and are useful diagnostic characters in the identification key below, while others are micromorphological characters, of which some relate to cork cells (Fig. 2).

Observation of the cultivated plants of Enteropogon revealed some _ character differences not always readily visible from herbarium specimens. Character differences including growth habit, number of inflorescence culms per plant and colour of spikelets, were more evident in cultivated plants of some species. For example, E. scabrilemma differs from E. dolichostachyus by having more numerous inflorescence culms per plant with leaves more obviously cauline; leaf-bearing portions of the culms elongated and occasional decumbent culms rooting at the nodes but lacking stolons; inflorescences mostly 1 or 2 racemes, and when fresh the inflorescence culms and rachis green; and racemes appearing tinted purple (Table 1).

Enteropogon scabrilemma and E. macrostachyus are morphologically simular in growth habit, inflorescence structure, and spikelet morphology (Table 1). The two species differ by several characters including

59

narrower, shorter spikelets (1-1.5 mm X 5.2- 6.4 mm vs 1.9-2 mm X 7-10 mm; the latter measurements for EL. macrostachyus from Clayton et al. 1974) in E. scabrilemma.

The micromorphology of the _ first lemma, abaxial surface of leaves and culms of SEnteropogon dolichostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma differ by micro-hairs, silica bodies and cork cells (Figs. 2, 6—8). The four species can be placed in two groups based on micromorphology. Cork cells across the four species vary 1n spacing, distribution, size and shape (Fig. 2). The bicellular micro-hairs of E. dolichostachyus are slightly larger than the other three species (Fig. 2). Enteropogon dolichostachyus and E. scabrilemma_ share slightly larger stomata, cork cells and silica bodies. The latter pair of species also share slightly wider long cells on the surface of inflorescence culms (Fig. 8).

Caryopses of Enteropogon species vary in shape, length and width (Fig. 9). There was also some variation in the relative length of the scutellum, size of the stylopodium, development of the dorsal ridge and cell width on the spermaderm (Figs. 9 & 10).

Leaf blades, sheaths and growth habits differed between species of Enteropogon (Figs. 1 & 11, Appendix 3). Indumentum of young leaves is a useful differentiating character (Fig. 3) but on some species the hairs are readily shed with age and mature leaves and the paraligule can be hairless.

There was a relatively small degree of variation in the anatomy of transverse sections of leaves and culms of Enteropogon dolichostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma (Figs. 12 & 13). All four species have clear parenchyma 8—20 cells wide by 3 or 4 cells thick above the mid-vein vascular bundle (Fig. 12). The strip of cells tends to give the mid-vein a silvery appearance on fresh leaves. The other four Australian species, viz. FE. acicularis, E. minutus, E. ramosus and E. unispiceus, were found to differ from the former four species by having only a few clear parenchyma cells above the mid-vein vascular bundle.

60 Austrobaileya 13: 51—93 (2023)

Fig. 2. Scanning electron micrographs of surface of lemmas (callus to RHS) of Enteropogon dolichostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma with classification of cork cells. The body of cork cells comprises a silica body (dark portion) and lid. A. FE. dolichostachyus (r, 1, L, a,c). B. E. paucispiceus (1, 2, s, b,c). C. E. pubifolius (i, 2,8, a,c). D. E. scabrilemma (r, 1, L, a, r). A from Thompson MORS37 (BRI); B from Thompson MORS42 (BRI); C from Thompson MOR841 (BRI); D from Thompson MOR844 (BRI). Micrographs (captured at *1000): E.J. Thompson.

Abbreviations: bmh bicellular micro-hair; CC cork cell; mh macro-hair; pr prickle

Ratings for cork cells (for most frequent occurrences) - Spacing: r regular; 1 irregular; separation: 1 mostly < one diameter, 2 frequently two diameters; size: s small (<< 15 um), L large (> 20 um); size of silica body: a > half of area, b < half of area; shape of silica body: ¢c crescent, r rectangular.

Thompson, New species of Enteropogon 61

Fig. 3. Leaf, collar and sheath at the mid-point along the vegetative portion of the culm of Enteropogon dolichostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma. A. E. dolichostachyus. B. E. paucispiceus. C. E. pubifolius D. E. scabrilemma. A from Thompson MORS37 (BRI); B from Thompson MORS42 (BRI); C from Thompson MORS41 (BRI); D from Thompson MORS44 (BRI). Photos: E.J. Thompson.

Abbreviation: pl — paraligule.

62 Austrobaileya 13: 51—93 (2023)

Chloris divaricata

Chloris ventricosa Enteropogon acicularis 1 Enteropogon acicularis 2 Enteropogon acicularis 3 Enteropogon paucispiceus Enteropogon ramosus Enteropogon minimus Enteropogon unispiceus Enteropogon dolichostachyus Enteropgon pubifolius Enteropogon macrostachyus

Enteropogon scabrilemma

_ Oo _ a) oS — cl Ww) rm = ry \o is GO — — va) cn — co) oS oe, a — cm)

Chloris divaricata Chloris ventricosa Enteropogon acicularis | Enteropogon acicularis 2 Enteropogon acicularis 3 Enteropogon dolichostachyus Enteropgon pubifolius Enteropogon paucispiceus Enteropogon ramosus ——_—_——_ Enteropogon minimus Enteropogon untspiceus sO nteropogon macrostachyuss

Enteropogon scabrilemma

© ‘a | wy) tt —) c~ rr) cr) _ > Ca — — co a) ry ae) ~— = = => —) S oe)

Fig. 4A & B. Variation in topologies for dendrograms generated from PATN cluster analyses for four examples of variations in analysis inputs using two datasets and two association measures (AM). Two datasets: Tl, 13 samples (includes Chloris) and 58 characters (13 multistate); T2, all multistate characters transformed to binary format (87 characters). A. Dataset = T1, AM = Gower metric. B. Dataset = Tl, AM = Czekanowsk1i.

Thompson, New species of Enteropogon 63

Chloris divaricata

Chloris ventricosa Enteropogon acicularis 1 Enteropogon acicularis 2 Enteropogon acicularts 3 Enteropogon ramosus Enteropogon dolichostachyus Enteropogon macrostachyus Enteropogon scabrilemma Enteropogon paucispiceus Enteropgon pubifolius Enteropogon minimus

Enteropogon unispiceus

0.8314 0.6235 0.2078 0.0000

0.4157

D Chloris divaricata Chloris ventricosa Enteropogon acicularis 1 Enteropogon acicularts 2 Enteropogon acicularts 3 Enteropogon paucispiceus Enteropogon ramosus Enteropgon pubifolius Enteropogon dolichostachyus Enteropogon macrostachyus Enteropogon scabrilemma Enteropogon minimus

Enteropogon unispiceus

i. — st rm > =t o or oO S — a rn oO S fo Ne +t oO — cS S oS —)

Fig. 4C & D. Variation in topologies for dendrograms generated from PATN cluster analyses for four examples of variations in analysis inputs using two datasets and two association measures (AM). Two datasets: Tl, 13 samples (includes Chloris) and 58 characters (13 multistate); T2, all multistate characters transformed to binary format (87 characters). C. Dataset = T2, AM = Gower metric. D. Dataset = T2, AM = Czekanowski.

64 Other discoveries include:

e Plants of Enteropogon acicularis have fragile inflorescence culms that usually disarticulate at maturity, very likely as are- sult of the sparser peripheral layer of scler- enchyma, compared to plants of E. ramo- sus that retain the old inflorescence culms.

e Enteropogon acicularis and E. ramo- sus differ by the length of the diaspore stipe, c. 1 mm and 0.5 mm, respectively.

e The indumentum in the raceme axils dif- fers with Enteropogon acicularis having hairs > 2 mm long and E. ramosus < 0.7 mm long. Axil indument is a reliable character on young inflorescences, but axils are sometimes glabrescent with age.

e Enteropogon minutus difters from. all the other Australian species by hav- ing inflorescence culms with a longi- tudinal groove or flattened edge. The contraligule may not be evident at all leaf blade-sheath junctions on a plant.

e nteropogon paucispiceus can have cleistogamous second florets.

e Besides compression, another useful char- acter that can be used to distinguish spike- lets of Chloris and Enteropogon 1s the rel- tive width of the first and second florets with Chloris having the second florets about half the width of the first, but occa- sionally Chloris divaricata R.Br. was ob- served in herbarium specimens to be like Enteropogon with narrow second florets.

Discussion

This study used results from phenetic analyses of a comprehensive set of morphological characters to provide an objective and repeatable process to aid decision making for alpha and beta taxonomy (Stuessy 2009; Thompson 2022b, c). The additional new morphological characters observed for Enteropogon were useful in discriminating the Australian species and helped provide information on the plasticity of some of the characters used in identification keys. The study affirmed the significance of the role of cultivation of plants over several years as part of the process of gathering information

Austrobaileya 13: 51—93 (2023)

on flowering, fruiting and growth habit (Thompson 2017, 2019, 2022b, c). Also, the results from cluster analyses supported the initial intuitive sense of similarities and differences in the putative species and provided potentially useful discriminating morphological characters for phylogenetic reconstructions that include molecular sequence data for correlated genes.

Examination of herbarium specimens and observation of cultivated plants for this study revealed that some characters such as growth habit and number of racemes per inflorescence, used in keys to identify some Australian species of Enteropogon, can lead to ambiguous identifications. For example, Nightingale et al. (2005) and Simon & Alfonso (2022) used “leaves mostly basal” and “leaves cauline” as distinguishing characters. The distribution of leaves on live plants is usually evident as either basal or cauline in Enteropogon. However, some herbarium specimens comprise only one or two culms, thus presenting potentially ambiguous character evaluation, especially when both the specimen and the label notes (including sometimes misleading terminology) are inadequate. Growth habit characters can be more evident when observed on living plants. This was exemplified by the stoloniferous habit of cultivated plants of Australian provenances of E. dolichostachyus. Further ambiguity in the identity of EF. dolichostachyus was encountered due to variations in descriptions and illustrations in floras over its geographic range across northern Australia, New Guinea, Timor, China, India, and the Philippines, the latter location where the type specimen was collected (Bor 1960; Lazarides 1972; Hsu 1978; Lazarides et al. 1992; Noltie 2000; Barkworth 2003; Nightingale et al. 2005; Bixing & Phillips 2006). It 1s possible that more than one taxon currently identified as E. dolichostachyus 1s present across Asia.

The inflorescences of &Enteropogon acicularis also exhibit degrees of plasticity in number and orientation of racemes, indumentum of the axils and culm diameter. Inflorescences of the cultivated plants of E. acicularis occasionally consisted of only five

Thompson, New species of Enteropogon

OSV = 0,0820 Enteropogon unispiceus Enteropegon minutus * 0.933 Enteropogon PaMOSUS Enteropogon paucispiceus nteropagon acicularis Enteropogoen macyostachyus Enteropegon acicularis | Enteropogon™acicularis 2 © Enteropogon pubifolius Enteropogon dolichostachyus -1,056 \ Pm ea, ee 0.833 “haha | 0,722 Enteropogon scabrilemma 0.917

OSV = 0.1073

Enteropogon unispiceus

, Enteropogon minutus Enteropogon ramosus

0.9

Enteropogon paucispiveus

>

Enteropogon pubifolius Enteropogon macyostackfus wai aad se 5 Enteropogon acicularis 2

Enteropogon acicularis 1

Enterepogon dolichostachyus

-1.038

-0.936 0.767

Enteropogon scabrilemma

0.845

65

Fig. 5A & B. Variation in topologies for 3-D ordinations generated from PATN principal component analyses using semi-strong, hybrid multidimensional scaling for four examples of variations in analysis inputs using four datasets and two association measures (AM). Four datasets: BI, baseline dataset with 11 samples and 43 characters (11 multistate); B2, as for Bl but with multistate characters transformed to binary (72 characters); Tl, 13 samples (includes Chloris) and 58 characters (13 multistate); T2, all multistate characters transformed to binary format (87 characters). A. Dataset

= Bl, AM = Gower metric. B. Dataset = B2, AM = Czekanowsk1.

66 Austrobaileya 13: 51—93 (2023)

OSV = 0.0863 Chloris divaricata 0.996 Chloris ventricosa Ir, Enteropogon minutus R Enteropogon unispiceus _ Enteropogon paucispiceus ; Enteropogon pubifolius Enteropogen ramosus Enteropogok aciculgris 1,2 & 3 Enteropogon macrostach a Zl Enteropogon dolichostachyus 0,842 x -0.924 -0.692 1.677 Enteropogon scabrilemma 0.548 D | Chloris ventricos OSV an 0. | 723 Moris Venmrircosa 1.252 Chloris divaricata Enteropogon unispiceus Enteropogon ramost Enteropogon paucispigeus Entcropegon minutus Enterapoyon acicularis 1 Enteropoguna acicularis 2 . ia wi ee cee Enteropogon pubifolius Enteropogon Enteropogon dolichostachyus -0,912 1.5395 Enteropogon scabrilemma 0.806

Fig. 5C & D. Variation in topologies for 3-D ordinations generated from PATN principal component analyses using semi-strong, hybrid multidimensional scaling for four examples of variations in analysis inputs using four datasets and two association measures (AM). Four datasets: BI, baseline dataset with 11 samples and 43 characters (11 multistate); B2, as for BI but with multistate characters transformed to binary (72 characters); T1, 13 samples (includes Chloris) and 58 characters (13 multistate); T2, all multistate characters transformed to binary format (87 characters). C. Dataset = Tl, AM = Gower metric. D. Dataset = T2, AM = Gower metric.

Thompson, New species of Enteropogon 67

‘iPapepres mF AS eee ‘ rT " . :

L., s wt re * \* a9 io ete :

Pe era Tees : ths ys,

fas! Le ga “A

» ™e”

*

i? yA let. . - ew

Fig. 6. SEM micrographs of abaxial leaf surface of Enteropogon dolichostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma (apex to RHS). A. FE. dolichostachyus. B. E. paucispiceus. C. E. pubifolius. D. E. scabrilemma. A from Thompson MORS37 (BRI); B Thompson MORS&42 (BRI); C from Thompson MORS4/ (BRI); D from Thompson MORS44 (BRI). Micrographs (captured at *1000): E.J. Thompson,

Abbreviations: bmh bicellular micro-hair; ew epicuticular wax (variable thickness layer); h hook; mh macro-hair; p papillae; pr prickle; sb silica body; S stoma.

68

Austrobaileya 13: 51—93 (2023)

yf ' Fi Send 7

ae ’ , re ° . -_— - i #), “y TaN _ : To VV. B+ - a ay =! he i -

4 y

hy 4 +) a - .

yy ,

e giret sae “BD.

= ~% o T' = . . - _* > ~ <a _ -* — _ £ “ . = < ee, | er - “a * eo ey — ea “ > — a ‘. ‘ 4 iw —_—" E ny a =F -« : “. . i es ee oe — * = ieee > ' —— | Pe

* —

_ . - Pr Se — -—- ——— al - +

a. oe -s “

° — Fa “a . e y ~ - - =. > a A —— A " - . - sn - ' tad _—_— > “ye ites ’ Ce 7” : — — ag ie _, —_—- ~—— ‘go eee * os — -~ «4 eo Team © .

_ = ~- “ean - , —Se* OS - ~ . - ~. 2 ae P—— © Fr © ioe. Ft ~—_— S - Tin Se eo ~, ae Lt = aie a " : -y~< - » wo fs > — + oe oe oofeaom ©) Te ohh . " : -_. —1- . o * a . = ” > - : . * °. - > 4 - . od ae Ce Fe "

Fig. 7. Replica of abaxial leaf surface of Enteropogon dolichostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma (apex to RHS). A. FE. dolichostachyus. B. E. paucispiceus. C. E. pubifolius. D. E. scabrilemma. A from Thompson MORS37 (BRI); B Thompson MORS&42 (BRI); C from Thompson MORS41/ (BRI); D from Thompson

MORS44 (BRI). Photos: E.J. Thompson.

Abbreviations: bmh bicellular micro-hair; mh macro-hair; p papillae; pr prickle; sb silica body; S stoma.

racemes overlapping with the number found in the closely related EF. ramosus (Simon 1984; Nightingale et a/. 2005). Culm width just below the raceme junction, a character not used in other studies of Enteropogon, was found to be positively correlated with number of racemes for E. acicularis ranging from 0.6 to 2.1 mm for 5 to 14 racemes, rarely 24, while for E. ramosus the corresponding ranges were 0.4 to 0.8 mm, occasionally 0.9, and 1 to 5, rarely 6.

The diaspore stipe found in species of Enteropogon and in some other chloridoid erasses has not been reported as a discriminating morphological character in other studies (Lazarides 1972; Watson &

Dallwitz 1992; Van Der Borre & Watson 1997). In Australian Enteropogon, the stipe is most pronounced, about | mm long, in E. paucispiceus and E. ramosus whereas in all the other Australian species it is < 0.6 mm long (Fig. 1). The stipe 1s uncommon in other Australian chloridoid genera with the diaspore composed of united florets such as in Astrebla F.Muell., Chloris, Oxychloris (F.Muell.) Lazarides, and also in subtribe Eleusininae. The stipe reaches its most conspicuous development in Australian chloridoid grasses in Oxychloris where it 1s c. 2 mm long. Furthermore, the stipe is absent in Eragrostis N.M.Wolf where the composition of the diaspore 1s highly variable with some African species having united florets and in

Thompson, New species of Enteropogon

the Australian species the diaspore ranging from the caryopsis alone to an individual floret with or without the rachilla or palea (Palmer et al, 2005). Another form of diaspore stipe occurs in andropogonoid grasses including Elionurus Willd., reported by Thompson (2017: 156) as a “proximal beak”. Although the stipe can have similarities in function for dispersal and burial, differences in the structure of the diaspore across grass taxa suggest ontogenetic differences, for example as found in many chloridoid grasses that have disarticulation above the glumes compared to andropogonoid grasses where disarticulation occurs below a spikelet pair (Peart 1979; Cheplick 1998; Cavanagh ef al. 2019).

ee rrr ~—a : hs il a - <a; : ee . =e -_—" _ — of oe . ~~ , >_> o&.

Ss , - hl ol

. a _ *

ss Ls

?

> a : - Ee saiterits

coca

. Soper. -.4 a + - rank ;

/

a -&

eee a: Ps ae nS teers ot *

x “yee

-

69

The paraligule found in some chloridoid erasses 1S also an uncommon occurrence in panicoid taxa but is rarely used as a discriminating character. Paraligules were recorded for species of Paspalum L. as “hairs behind the ligule” by Rua & Aliscion1 (2002) and have been found useful by the author (unpubl.) in aiding identification of the Australian species.

The frequency distribution of species of Enteropogon by climatic zones follows a broadly similar trend to that of Poaceae collectively in Australia (Map 1; Thompson 2021). The highest frequencies of species occur in tropical and subtropical zones across the Northern Territory and Queensland.

Fig. 8. SEM micrographs of surface of terminal inflorescence culm of Enteropogon dolichostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma (apex to RHS). A. E. dolichostachyus. B. E. paucispiceus. C. E. pubifolius. D. E. scabrilemma. A from Thompson MORS37 (BRI); B from Thompson MORS42 (BRI); C from Thompson MORS41 (BRI); D from Thompson MOR844 (BRI). Micrographs (captured at *1000): E.J. Thompson,

Abbreviations: ew epicuticular wax (variable thickness layer); h hook; sb silica body; S stoma.

70 Austrobaileya 13: 51—93 (2023)

Fig. 9. Caryopses of Enteropogon dolichostachyus, E. macrostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma from dorsal view and cross-section just above scutellum. A & E. E. dolichostachyus. B & F. E. macrostachyus. C & G. E. pubifolius. D & H. E. scabrilemma. A from Thompson MORS37 (BRI); B from Davidse 5886 (BRI); C from Thompson MORS844 (BRI); D from Thompson MORS&41 (BRI). Scale bar = 4 mm. Photos: E.J.

Thompson.

Abbreviations: Se scutellum; St stylopodium.

Thompson, New species of Enteropogon 71

Fig. 10. SEM micrographs of caryopsis endosperm of Enteropogon dolichostachyus, E. macrostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma from dorsal view. A. E. dolichostachyus. B. E. macrostachyus. C. E. scabrilemma. D. E. pubifolius. A from Thompson MORS837 (BRI); B from Davidse 5886 (BRI); C from Thompson MOR844 (BRI); D from Thompson MOR841 (BRI). Micrographs (captured at *1000): E.J. Thompson,

72 Austrobaileya 13: 51—93 (2023)

Fig. 11. Growth habit of cultivated plants of Enteropogon dolichostachyus, E. paucispiceus, E. pubifolius and E. scabrilemma. A. E. dolichostachyus. B. E. scabrilemma. C. E. paucispiceus. D. E. pubifolius. A. from Thompson MORS37 (BRI); B. from Thompson MORS44 (BRI); C. from Thompson MORS842 (BRI); D. from Thompson MORS841

(BRI). Photos: E.J. Thompson.

Thompson, New species of Enteropogon 73

sclerenchyma colourless bulliform cells mesophyll

papillae tertiary vascular bundle

prickle chlorenchyma

thick-walled cells

between xylem and : primary vascular phicem bundle

sclerenchyma

Fig. 12. Transverse section of fresh leaf at mid-vein of Enteropogon scabrilemma. Broad-leaved species of Enteropogon exhibit variation in thickness (up to 6 cells) and width (up to c. 16 cells) of parenchyma above the mid-vein and some narrow-leaved species have none or few of these cells. From Thompson MORS44 (BRI). Del. E.J. Thompson.

sclerenchyma

chlorenchyma

100 um

Fig. 13. Transverse section of portion of inflorescence culm of Enteropogon scabrilemma. Morphology and anatomy of the Australian species shows little variation except for E. minutus that has semi-circular cross-section and E. acicularis with lower density of primary vascular bundles. From Thompson MORS44 (BRI). Del. E.J. Thompson.

74

Taxonomy

Austrobaileya 13: 51—93 (2023)

Key to the Australian species of Enteropogon (modified from Simon & Alfonso (2011))

1 Diasporée with an elongated stipe;> | mm lone . 22 ea ee ee we ee eS 2

1. Diaspore witha stipe<0.7mmlong....

2 Culms robust; leaves cauline, drying flat, often coiled; hairs at raceme

junction ¢;-O STON, 2 os ie BSS

2. Culms slender; leaves basal, drying flat and straight; hairs at

raceme junction>2mmliong.......

3 Number of racemes > 7; culms > 0.9 mm wide just below junction of

Leaves with pilose abaxial surface, >2mmwide............... E. pubifolius

3 4 4. Leaves glabrous abaxially, width varies . . 5

Leaves < 1.5 mm wide; racemes usually | (—3); awn of basal lemma< 9mm

HOR mye fee ess opt: oo Beene la Gh ON cos ro ne Ree A one we A Sina E. unispiceus 5. Leaves > 1.5 mm wide; racemes | or more; awn of basal lemma > 9 mm ROTO cr ae os aa tam RS Balen hee anaes oe Rag Ra mae Ae ap an itl A ha Rag ee el ute ve atet its Ran Re ct somee 8 6

6 Lemma of proximal floret < 3.8 mm long; contraligule present; inflorescence culm furrowed or flattened on one side ............. E. minutus 6. Lemma of proximal floret > 3.5 mm long; contraligule usually absent;

inflorescence culm circtilar In-cross-section . . . 6 ee ee a ee eS 7 7 First floret > 1 mm wide; racemes usually 1-2 (-4) ..........0... E. scabrilemma 7. First lemma smooth or sparsely scabrid; racemes usually6................. 8 8 First lemma conspicuously scabrid; widest leaves<3mm........... E. acicularis 8. First lemma smooth or sparsely scabrid; widest leavesc.6mm.... . E. dolichostachyus

1. Enteropogon pubifolius E.J.Thomps., sp. nov.

Similar to Enteropogon paucispiceus (Lazarides) B.K.Simon differing by having pilose broader leaves, spikelets drying dark erey, short diaspore stipe and rachilla with a distal beard. Type: Queensland. Ex-sITu CULTIVATED: Ashgrove (ex Cape Melville National Park, Mookai Creek), 1 May 2019, E.J. Thompson MORS4I/ (holo: BRI [AQ1023144, comprising two sheets]).

Perennial grass, 30-90 cm high, tussock- forming, occasionally with decumbent culms rooting at the nodes. Fertile culms erect, rigid, 3—7 noded, < 0.9 mm wide, internodes short giving the appearance of basal leaves. Leaf blades 3-15 cm long, 3—4.5 mm wide, drying flat; abaxial leaf blade epidermis:

pilose with tuberculate-based simple hairs to 0.3 mm long and scabridulous with prickles; densely papillate in the intercostal zone; epicuticular wax present. Ligule a fringed membrane, 0.1—0.2 mm long; paraligule hairs to 3.2 mm long. Leaf sheaths usually shorter than the internodes, glabrous, bearded at the orifice with stiff hairs c. 2.7 mm long, round on the back; collar glabrous. Inflorescence a digitate to subdigitate panicle with 3-5 racemes, 40—80 mm long, c. 1.6 mm wide and 1.4 mm thick, divaricate, with or without a pseudo-node with a subtending reduced leaf and sheath to 9.8 mm long, or with a bract- like scale to 0.3 mm long; rachis c. 0.3 mm wide. Spikelets with 2 or 3 florets. Glumes narrow lanceolate, membranous, glabrous; lower glume 1.6—2 mm, entire, apex acute to awned, awn c. 0.5 mm long; upper glume

Thompson, New species of Enteropogon

TS

Fig. 14. Spikelet of Enteropogon pubifolius. A. lower glume. B. upper glume. C. dorsal view of first floret. D. ventral view of second floret and dorsal view of third floret (Second floret manipulated outward). E. dorsal view of first palea. F, transverse view of first palea. G. ventral view of caryopsis. H. dorsal view of caryopsis. I. Lateral view of caryopsis. J. transverse view of caryopsis. K. ventral view of first floret with dorsal view of second floret. All from Thompson

MORS41 (BRI). Del. E.J. Thompson.

4—4.3 mm long, apex attenuate to awned, awn to 0.9 mm long. Florets 2 or 3, | fertile, stipe c. 0.4mm long. Lowest lemma elliptical, 4.1— 5.1 mm long, 0.6—0.8 mm wide, membranous, smooth; apex acute, bilobed, lobes to 0.3 mm long; awn 7.5—8.5 mm long, straight, filiform. Palea subequal to lemma, membranous, minutely scabrid between the keels, apex acute. Anthers 3, chasmogamous 0.9-1 mm long; cleistogamous 0.7—0.9 mm long. Caryopsis of first floret 2.5—2.9 mm long, c. 0.7 mm wide. First rachilla c. 1.2 mm long,

sometimes bearded towards the apex with hairs c. 0.3 mm long. Second lemma 1.4—1.9 mm long, 0.2—0.3 mm wide; awn 3.7—6.5 mm long. Palea present or absent, subequal to lemma. Second rachilla 0.2 mm long. Third lemma to 1.8 mm long, c. 0.1 mm wide; awn 1.4-1.9 mm long. Micromorphology of the lemmas: surface densely covered in relatively large cork cells and sparsely scabridulous with prickles and simple micro-hairs to c. 50 um long; bicellular micro-hairs to 26 um long (Figs. 1, 6, 7, 14, 15).

76

Queensland Herbarium (BRI) Ex-situ cultivated

Australia; Queensland Cook Enteropogon

Coll.: Thompson, E.J. 1 May 2019 Coll, no.: MOR841

Lat,: 14° 15' 33.900"S Long.: 144° 27’ 32.300"E

Datum: GDA94 Alt.: 80m

Cape Melville National Park, Mookai Creek. Provenance: Ashgrove, Queensland

Pot cultivation in nursery. Grown in full sun. pave habut with mostly basal leaves. Fertile culms to 30

Propagated from caryopsis collected from BRI-AQ837374 (Forster et al. PIF41368).

Det.: Thompson, E.J., May 2019 Dup.: 359.0 Poaceae

M000

RI-AQ1023144 Prep at BRI: Sheet

Austrobaileya 13: 51—93 (2023)

QUEENSLAND HERBARIUM (BRI) Brisbane Australia

AQ /OL 3/ 4¢Y

Fig. 15. Holotype specimen of Enteropogon pubifolius. Sheet | of 2. Photo: Queensland Herbarium.

Thompson, New species of Enteropogon

Additional specimens examined: Queensland. Cook District: Cape Melville National Park, Mookai Creek, May 2014, Forster PIF41368, S.L. Thompson & Cape Melville Traditional Owners (BRI); Kings Plains Station, near Kings Plains Lake, Apr 2015, McDonald KRMI6891 & E.J. Thompson (BRI); Alkoomie Station, Melody Rocks, Apr 2016, McDonald KRMI8361 & Forster (BRI).

Distribution and habitat: Enteropogon pubifolius 1s endemic to the tropical east coast of Queensland, Australia from northwest of Cooktown up to Cape Melville (Map 2). Plants have been collected from tussock grassland dominated by 7riodia microstachya R.Br. intermingled with small copses of vinethicket on exposed granite bluffs and clifflines at Cape Melville; from small vinethicket patches on limestone karst on Alkoomie Station northwest of Cooktown and from a lake edge in woodland dominated by eucalypts and melaleucas at Kings Plains Station.

Phenology: Plants produce flowers and fruits mostly in April—May.

Notes: —Enteropogon — pubifolius has similarities to £. paucispiceus in growth habit and inflorescence typology. The spikelets dry dark grey (Fig. 1).

Etymology: The specific epithet 1s derived from Latin and refers to the pubescent leaves.

Conservation status: Enteropogon pubifolius is known from three herbarium specimens collected from three locations about 120 km apart. Populations at these sites are not accurately known so further field assessment is required. However, it is suggested that this species should be considered Vulnerable based on Criterion B2a (IUCN 2019) and a formal conservation status nomination should be made.

2. Enteropogon scabrilemma E.J.Thomps., Sp. nov.

Similar to £. macrostachyus (Hochst. ex A.Rich.) Munro ex Benth., differing by having smaller spikelets with longer and more slender rachillas. Type: Queensland. Ex-sITu CULTIVATED: Ashgrove (ex Riversleigh

T7

Station, on a remote mining exploration track), 1 May 2019, E./. Thompson MORS35 (holo: BRI [AQ1023137]).

Illustrations: Lazarides (1972: 26; although the specimen source was not cited, morphology including collar and ligule (Figs. 48E&F) and spikelet with bearded rachilla apex (Fig. 49K) match E. scabrilemma).

Perennial grass, 70-90 cm high, tussock- forming, occasional decumbent culms rooting at the nodes. Culms erect, wiry, 5—8 noded, < 0.9 mm wide. Leaf blades 4—45 cm long, 2-9 mm wide, drying flat. Ligule 0.3—0.4 mm long; paraligule to 4.5 mm long. Leaf sheaths usually longer than the internodes, bearded at the orifice with stiff hairs 2-4 mm long, rounded on the back; collar glabrous. Inflorescence a digitate panicle with 1-5 racemes, 90-120 mm long, c. 1.5 mm wide and 2.5 mm thick, divaricate; rachis 0.5—0.6 mm wide, relatively stiff. Spikelets with 2 or 3 florets. Glumes narrow lanceolate, membranous, acute to awned, glabrous; lower glume 1.7—2.7 mm, entire, awn to 0.9 mm long; upper glume 4.2—5.6 mm _ long, awn to 1.9 mm long. Florets 2(or 3), 1 or 2 fertile; stipe c. 0.6 mm long. Lowest lemma chartaceous, elliptical, 4.1-6.5 mm _ long, 1—1.5 mm wide, scabrous; apex acute, bilobed, lobes to 0.3 mm long; awn filiform, straight, 10.7-16.4 mm long. Palea subequal to lemma, chartaceous, scabrid, apex acute, bilobed. Anthers 3, chasmogamous 1|.5—1.7 mm long; cleistogamous c. 1.4 mm long. Caryopsis of first floret 3.5—4.9 mm long, 0.9-1.3 mm wide, keeled dorsally, ventrally concave. First rachilla 1.4—2.4 mm long, glabrous or bearded towards apex with hairs to 0.4 mm long. Second lemma, sterile or fertile, scabrid; fertile: cleistogamous, to 3.1 mm long and 0.7 mm wide; infertile: c. 1.1 mm long; awn 3.9-7.3 mm long. Palea present or absent, equal to lemma. Anthers 3, 1.2—1.8 mm long. Caryopsis of second floret c. 2.2 mm long, to 0.7 mm wide. Second rachilla 1—1.3 mm long. Third lemma sterile, vestigial to 1 mm long, awn c. 1.3 mm long. Figs. 16 & 17.

78 Austrobaileya 13: 51-93 (2023)

5 mm

Fig. 16. Spikelet of Enteropogon scabrilemma. A. lower glume. B. upper glume. C. dorsal view of first floret. D. ventral view of first floret and dorsal view of second floret. E. dorsal view of first palea. F. transverse view of first palea. G. ventral view of caryopsis. H. dorsal view of caryopsis. I. transverse view of caryopsis. J. dorsal view of fertile second floret showing third floret. K. ventral view of spikelet showing dorsal view of fertile second floret. L. ventral view of caryopsis from second floret. L. dorsal view of caryopsis from second floret. All from Thompson MORS44 (BRI). Del. E.J. Thompson.

Thompson, New species of Enteropogon T9

ee =

Queensland Herbarium (BRI)

Ex-situ cultivated QUEENSLAND HERBARIUM (BRI) Australia: Queensland Burke Brisbane Australia Enteropogon dolichostachyus

AQ /OLDI SDT

Coll.: Thompson, E.J. 1 May 2019 Coll. no.: MOR835

Lat.: 19° 38' 58,379"S Long.: 138° 56’ 22.109"E

Datum: WGS84

Riversleigh Stn, on a remote mining exploration track. Provenance: Ashgrove, Queensland

Pot cultivation in nursery. Grown in full sun.

Culms to 60 cm tall with leaves distributed evenly. Occasional decumbent culms rooting at the nodes. Propagated from caryopsis collected from BRI-AQ850308 (Booth and Kelman CAM 28-1).

Det.: Thompson, E.J., May 2019 359.0 Poaceae

Dup.: | ; BRI-AQ1023137

Prep at BRI: Sheet

Fig. 17. Holotype specimen of Enteropogon scabrilemma. Photo: Queensland Herbarium.

80

Additional specimens examined: Papua New Guinea. CENTRAL PROVINCE: c. | mile [1.6 km] N of Kapa Kapa, Rigo Sub-district, July 1972, Pullen 3278 (CANB); Tavai Creek area, c. 46 miles [74 km] SE of Port Moresby, Apr 1967, Pullen 6829 (BRI). Australia. Western Australia. 3 km E of Cape Leveque, N Dampier Peninsula, Carter 397 (PERTH); Gibb River — Kalumburu Mission Road; c. 2 km NE of Carson River crossing, 115 km (by road) NE of Mitchell River turnoff, c. 175 km NW of. Wyndham, Jun 1976, Beauglehole 51960 (CANB); Long Island, Buccaneer Archipelago, NE of Derby, Apr 1997, Martin CBIOI (CANB); 13.5 km NE of Crystal Head on SW Osbourne Island, Mar 1989, Keighery 10639 (PERTH); Martin CBI0I (PERTH); 3 km E of Cape Leveque, N Dampier Peninsula, May 1989, Carter 397 (PERTH); Long Island, Buccaneer Archipelago, NE of Derby, Apr 1997, Martin CBI0O] (PERTH); Wyndham, East Kimberley, Apr 1992, Mitchell 2320 & Willing (PERTH); Behind sand dune cemetery at One Arm Point, about 200 m N of Broome, Jul 1997, Mitchell 4809 (BRI). Northern Territory. S of Wollogorang Station. Plot 1421, Oct 1998, Harwood 535 (DNA). Queensland. BuRKE DIstTRIcT: Riversleigh Station, on a remote mining exploration track, Jun 2006, Booth CAM28-I & Kelman (BRI). Cook District: Princess Charlotte Bay, Aug 1980, Buckley 6203 (BRI); ibid, Aug 1980, Buckley 6330 (BRI); 2 km S of Bathurst Heads Camping Area, Kalpowar, Aug 2016, Thompson SLT16513.1, Ross, Ross & Wallace (BRI); Kings Plains Station, Kings Creek track, Apr 2017, McDonald KRMI1935I, Forster & Paradise (BRI); Cooktown, Quarantine Bay, May 1970, Blake 23488 (BRI). CULTIVATED. Ashgrove, Mar 2017, Thompson MORS36 (BRI); Ashgrove, Jun 2020, Thompson MORS44 (BRI).

Distribution and habitat: Enteropogon scabrilemma 1s widespread 1n coastal tropical Australia (Kimberley of Western Australia, top end of the Northern Territory, Cape York Peninsula, Queensland) and southern Papua New Guinea (Map 2). Plants occur in eucalypt dominated woodlands on sandy substrates such as dunes or alluvium along watercourses.

Phenology: Dates of herbarium collections reveal flowering and fruiting throughout the year. Cultivated plants bore flowers mostly in summer.

Affinities: Although specimens of Enteropogon scabrilemma were originally identified as EF. dolichostachyus, E. scrabilemma has more morphological similarities to the African species, £E. macrostachyus. Enteropogon scabrilemma

Austrobaileya 13: 51—93 (2023)

and E. macrostachyus are very similar in erowth habit, leaf size and distribution, and inflorescence characteristics including the number of racemes and spikelet imbrication. Both species have relatively broad scabrid spikelets and rachillas with a distal beard but E. macrostachyus has much larger lemmas

(Fig. 1).

Notes: Enteropogon scabrilemma _ has “monomorphic cleistogamous anthers” as defined by Thompson (2021) where the chasmogamous and cleistogamous anthers are of the same size. The second florets can be reduced to a lemma or sometimes present fertile with cleistogamous caryopses (Fig. 16). Spikelets dry straw-coloured (Fig. 1).

The additional specimens of Enteropogon scabrilemma cited above have mostly been stored in Australian herbaria as E. dolichostachyus.

Two specimens, Keighery 10125 (CANB) from North Kimberley and Lazarides 5635 (CANB), have morphological similarities to Enteropogon scabrilemma and E. minutus, respectively. The former specimen differs from £. scabrilemma by the smaller spikelets, slender growth habit and narrower leaves and the latter specimen differs from E. minutus by the larger spikelets and absence of contraligule.

Etymology: The specific epithet 1s derived from Latin and refers to the scabrid lemmas.

Conservation Status: Enteropogon scabrilemma has a wide distribution; however, its abundance at any of its known locations 1s unclear. Further field investigation 1s required before its conservation status can be assessed.

Acknowledgements

I greatly appreciate the loan of specimens from CANB, PERTH and NT. I am very erateful to Dr Gordon Guymer for the provision of resources at BRI. Thanks to Dr Melody Fabillo and Dr Paul Forster for reviewing drafts of the manuscript.

Thompson, New species of Enteropogon References

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