Volume 9 Number 1 2013 AUSTROBAIIEYA A Journal of Plant Systematics and Conservation Biology Queensland Herbarium Department of Science, Information Technology, Innovation and the Arts Queensland Government Volume 9 Number 1 2013 A Journal of Plant Systematics and Conservation Biology Queensland Herbarium Department of Science, Information Technology, Innovation and the Arts iSh QUEENSLAND ffilr GOVERNMENT Editorial Committee P.I.Forster (editor) P.D.Bostock (technical advisor) G.P.Guymer (technical advisor) Graphic Design Will Smith Desktop Publishing Aniceta Cardoza Austrobaileya Vol. 1, No. 1 was published on 1 December 1977 Vol. 8, No. 4 was published on 21 December 2012 Austrobaileya is published once per year. Exchange: This journal will be distributed on the basis of exchange. Australian Subscribers: Orders for single issues and subscriptions may be placed. The price is (GST included): AUD$44.00 per issue for individuals, AUD$71.50 for institutions, including postage. Overseas Subscribers: Orders for single issues and subscriptions may be placed. The price is AUD POA per issue for individuals, AUD$95.00 for institutions, including postage. All correspondence relating to exchange, subscriptions or contributions to this journal should be addressed to: The Editor, Austrobaileya, Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts (DSITIA), Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: paul.forster@science.dsitia.qld. gov.au ISSN 0155-4131 © Queensland Herbarium 2013 Web site: http://www.qld.gov.au/environment/plants-animals/plants/herbarium/publications/ Austrobaileya is the journal of the Queensland Herbarium and publishes peer-reviewed research on plants, algae, fungi and lichens (systematics, morphology, geography, anatomy, karyology, conservation biology and botanical history), with special emphasis on taxa from Queensland. Opinions expressed by authors are their own and do not necessarily represent the policies or view of the Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts. Austrobaileya 9 ( 1 ): 1-154 ( 2013 ) Contents A taxonomic revision of Garcinia L. (Clusiaceae) in Australia, including four new species from tropical Queensland W.E. Cooper .1-29 Reinstatement and revision of Sphaeromorphaea DC. and Ethuliopsis F.Muell. (Asteraceae: Plucheinae ) A.R.Bean .30-59 Reinstatement of intraspecific taxa for Bosistoa pentacocca (F.Muell.) Baill. (Rutaceae) with a new combination B. pentacocca subsp. connaricarpa (Domin) P.I.Forst. P. I. Forster .60-65 Three new species of Pluchea Cass. (Asteraceae: Inuleae-Plucheinae) from northern Australia A.R.Bean .66-74 Cryptocarya cercophylla W.E.Cooper (Lauraceae), a new species from Queensland’s Wet Tropics W.E.Cooper .75-79 Violaperreniformis (L.G.Adams) R. J.Little & G.Leiper, stat. nov., with notes on Australian species in Viola section Erpetion (Violaceae) R. J.Little & G.Leiper .80-101 Pterostylis caligna M.T.Mathieson (Orchidaceae), a new species from northern Queensland M.T.Mathieson .102-106 Two new subspecies of Eucalyptus tereticornis Sm. (Myrtaceae) A. R.Bean .107-113 A new fern species for Queensland: Diplazium squamuligerum (Rosenst.) Parris (Woodsiaceae) D.J.Ohlsen & A.R.Field .114-125 New records for Queensland in Lindernia All. (Linderniaceae) B. S.Wannan .126-129 Status and lectotypification of Craspedia gracilis Hook.f. (Asteraceae: Gnaphalieae) A.C.Rozefelds .130-135 Craspedia uniflora G.Forst. (Asteraceae) is a New Zealand endemic and not part of the Australian flora A.N.Schmidt-Lebuhn .136-139 Cullen spicigerum (Domin) A.E.Holland (Fabaceae), a new combination and reinstatement of a north Queensland species A. E. Holland .140-145 Celtis strychnoides Planch, an earlier name for Celtis australiensis Sattarian (Ulmaceae) G.P.Guymer .146-147 Reinstatement of Rapanea leucobrachya P.Royen (Myrsinaceae) from New Guinea, with a new combination as Myrsine leucobrachya (P.Royen) PI.Forst. P.I.Forster. .148-149 Emmenosperma pancherianum Baill. (Rhamnaceae) newly recorded for Australia A. R. Be an .150-152 Euphorbia operta Halford & W.K.Harris (Euphorbiaceae), a new name for Euphorbia occulta Halford & W.K.Harris, nom. illeg. and lectotypification of the name Euphorbia australis var. subtomentosa Domin D.A.Halford & W.K.Harris .153-154 A taxonomic revision of Garcinia L. (Clusiaceae) in Australia, including four new species from tropical Queensland W.E. Cooper Summary Cooper, W.E. (2013). A taxonomic revision of Garcinia L. (Clusiaceae) in Australia, including four new species from tropical Queensland. Austrobaileya 9(1): 1-29. The genus Garcinia is revised for Australia with nine native and three naturalized species included. Garcinia jensenii W.E.Cooper, G. leggeae W.E.Cooper, G. russellii W.E.Cooper and G. zichii W.E.Cooper are newly described and illustrated. All species are fully described, together with notes on habitat and distribution. A key to Garcinia species in Australia is provided. Key Words: Clusiaceae, Garcinia , Garcinia brassii , Garcinia dnlcis , Garcinia gibbsiae, Garcinia jensenii , Garcinia leggeae , Garcinia livingstonei, Garcinia mangostana , Garcinia mestonii , Garcinia riparia , Garcinia russellii , Garcinia zichii , Garcinia warrenii, Garcinia xanthochymus , Australia flora, Queensland flora. Northern Territory flora, new species, taxonomy, identification keys, rainforest, naturalised species W.E. Cooper, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor Road, Smithfield, Queensland 4878, Australia. Introduction Garcinia L. (Clusiaceae) is a pantropical genus of about 260 species (Stevens 2007) of mostly dioecious shrubs and medium sized trees found in Africa, Madagascar, Comoros, the Indian subcontinent (including Bangladesh, Bhutan, India & Sri Lanka), SE Asia (including Cambodia, China, India, Laos, Myanmar, Nepal, Thailand and Vietnam), Malesia (including Malaysia, Indonesia, the Philippines and New Guinea), Solomon Islands, Australia, New Caledonia, Fiji and South America. Species are restricted to various forms of rainforest between the lowlands and the highlands, the exception being Garciniaamplexicaulis Vieil 1. exPierre, which occurs in exposed areas and secondary regrowth thickets on degraded and mined areas in New Caledonia. Within Australia, 12 species (nine native, three naturalised) of Garcinia occur in Queensland’s rainforests and gallery forests from the Torres Strait islands south to the Kirrama Range in the Wet Tropics bioregion, with one species (G. warrenii) on Melville Island in the Northern Territory where it occurs in monsoon Accepted for publication 9 September 2013 vine forest. Seven species are endemic to Queensland. Garcinia plants have been recorded as having white, cream or yellow exudate (Nazre 2006); whereas, some Australian species also have clear or slightly milky exudate. Bracts and glands are a feature of this genus and there are three different types on most species, all are insignificant in species delimitation (Nazre 2006). Fruits are baccate with a single ovule per carpel, a unique combination in Clusioideae and a single ovule per carpel is a synapomorphy for the genus (Sweeney 2008: 1298). However, two Australian species, Garcinia gibbsiae S.Moore and G. russellii W.E.Cooper produce dry berries and commonly contain up to 4 ovules per locule and therefore the presence of a single ovule per carpel is no longer a diagnostic feature for the genus. The first record of Garcinia from Australia was of G. mestonii F.M.Bailey (as ‘mestonf ) (Bailey 1889), followed quickly by G. warrenii F.Muell. (Mueller 1891). The last overall flora account of Australian Garcinia enumerated only three species (G. cherryi F.M.Bailey [Bailey 1902; subsequently transferred to Ternstroemia (Theaceae) as T. 2 cherryi (F.M.Bailey) J.F.Bailey & C.T.White], G. mestonii and G. warrenii ) (Bailey 1899). Additional species were added by Moore (1917) and White (1933, 1936) and names for Malesian taxa have been variously misapplied to some species; however, the genus has been little studied taxonomically in the intervening years to the present. Consequently, putatively undescribed species known only by phrase names have been recognised in the last twenty years and nine native species were listed for Australia by Cooper & Cooper (2004) and CHAH (2007). Examination of both live populations of plants and herbarium material, coupled with preliminary phylogenetic analyses of a limited DNA sequence dataset indicate that four undescribed species occur in Australia. Previously, Garcinia species endemic to Australia have been named in honour of worthy recipients and this revision continues the tradition by honouring four people who have made valuable contributions to Queensland botany. G. jensenii W.E.Cooper is described for material from Cape York Peninsula previously assigned to G. riparia A.C.Sm. at CNS. G. leggeae W.E.Cooper is described for all of the CNS and some of the BRI material of G. sp. (Claudie River) (the remaining BRI material is G. jensenii). G. zichii W.E.Cooper is described for specimens known as G. sp. (Davies Creek) and G. russellii had not been recognised by any herbaria but was listed in Cooper & Cooper (2004) as G. sp. (Mossman). Lineages, clades and associated morphology A phylogenetic analysis of DNA sequence data resolved two major lineages within Garcinia (Sweeney 2008). Australia is represented by 12 species (nine native and three naturalised) distributed among 6 clades. Two species occur in Clades 1 & 2 within Lineage A, and ten species occur in Clades 4, 5, 6 & 9 within Lineage B. Lineage A is distinguished by the presence of floral nectaries. Lineage B is distinguished in part, by the lack of disks, appendages and rings (nectaries) (Sweeney 2010: 158). Austrobaileya 9(1): 1-29(2013) Sweeney (2008: 1299) in reference to Lineage B, stated that Clades 2 and 4 usually lack a well-developed pistillode. However, using Sweeney’s evidence (2008: 1292 and in lift .), Clade 2 is part of Lineage A and Clade 4 does have a well-developed pistillode. It is Clade 7 and some species within Clades 5 and 9, which lack a well-developed pistillode. Clade 1 (Garcinia dulcis (Roxb.) Kurz and G. xanthochymus Hoolc.f. [the latter naturalised in Queensland]) is distinguished by 5-merous flowers. Clade 2 (G. Iivingstonei [naturalised in Queensland]) is distinguished by free stamens. Clade 4 (G. jensenii and G. zichii ) is distinguished by fleshy bilocular flowers and fruit, which are capped with a disk-like stigma, unlike the woody pericarp described by Sweeney (2008). Clade 5 (G.mestonii and G. leggeae ) is distinguished by an androecium of fused filaments creating a solitary fleshy receptacle that is not adnate to the petals. Sweeney (2008) stated an additional possible synapomorphy for Clade 5 was the fruit, which usually have furrows along the septal radii, and for Clade 9, the presence of a distinctive adaxial, exudate- containing canal pattern in the leaves, consisting of highly branched, anastomosing canals. In Australian material; however, these characters poorly predict clade membership because furrows (or at least grooves) along the septal radii (putatively synapomorphic for Clade 5) occur in two species from Clade 9 but not in the Clade 5 species. Furthermore, intricately branched exudate-containing canals (putatively synapomorphic for Clade 9: G. brassii C.T.White, G. gibbsiae, G. rnsselli and G. warrenii) are found in G. dulcis and G. Iivingstonei (Clades 1 & 2). Clade 6 (G. mangostana L.) has no clear morphological synamorphies but is usually distinguished by a fungiform pistillode surrounded by an undivided 4-lobed androecium (Sweeney 2008). Clade 9 in Australian plants is distinguished by an androecium that is adnate to the petals. Materials and methods The study is based upon the examination of 521 specimens or photos from BRI, CANB, CNS, K, MEL, NSW and P. All specimens Cooper, Garcinia in Australia cited have been seen by the author and all native species have been observed in the field. Naturalised species have been seen in the field and/or cultivation. Measurements of the floral parts and fruits are based on material preserved in 70% ethanol as well as fresh material from the field; d.b.h. refers to the forestry logging term of diameter at breast height. Abbreviations in the specimen citations are: EP (Experimental Plot), FR (Forest Reserve), LA (Logging Area), NP (National Park), SF/SFR (State Forest/State Forest Reserve) and TR (Timber Reserve). The superscript 1 refers to online images of specimens. Species are arranged sequentially following the disposition of taxa outlined by Sweeney (2008). Taxonomy Garcinia L. Sp. PI. 1: 443 (1753). Type species: G. mangostanaV. Rheedia L., Sp. PI. 1193 (1753). Type species: Rheedia laterifolia L. [=Garcinia humilis (Vahl) C.D.Adams], Cambogia L., Gen. PI. , ed. 5:225 (1754). Type species: Cambogia gummi-gutta L. [=Garcinia gummi-gutta (L.) N.Robson], Oxycarpus Lour., FI. Cochinch. 647 (1790). Type species: Oxycarpus cochinchinensis Lour. [=Garcinia cochinchinensis (Lour) Choisy], Verticillaria Ruiz & Pav., FI. Peruv. Prodr. 81: t. 15. (1794). Type species: Verticillaria acuminata Ruiz & Pav. Xanthochymus Roxb., PI. Coromandel 2: 51 (1798). Type species: Xanthochymuspictorius Roxb. [=Garcinia xanthochymus Hook.fi], Brindonia Thouars, Diet. Sci. Nat. 5: 339 (1804). Type species: Brindonia indica Thouars. [=Garcinia indica (Thouars) Choisy], Stalagmitis Murray, Commentat. Soc. Regiae Sci. Gott. 9: 173 (1789). Type species: Stalagmitis cambogioides Murray \=Garcinia ovalifolia 01 iv.]. 3 Hebradendron Graham, Companion Bot. Mag. 2: 199 (1837). Type species: Hebradendron cambogioides Graham [=Garcinia morella Desr.]. Discostigma Hassk., Flora 25(2, Beibl .): 33 (1842). Type species: Discostigma rostrata Hassk. [=Garcinia rostratum (Hassk.) Hook.fi], Rhinostigma Miq., FI. Ned. Ind., Eerste. Bijv. 3: 495 (1861). Type species: Rhinostigma parvifolia Miq. [=Garciniaparvifolia Miq.]. Pentaphalangium Warb., Bot. Jahrb. Syst. 13: 382 (1891). Type species: Pentaphlangium crassinerve Warb. \=Garcinia crassinervis (Warb.) Kosterm.]. Garcinia in Australia: Dioecious or monoecious (agamospermous in the naturalised G. mangostana ), mostly subcanopy trees but also shrubs or canopy trees to 30 m; exudate, yellow, white or colourless, opaque or translucent. Branchlets square, ribbed or terete; petioles mostly channelled and ligulate. Interpetiolar gland-like bracts at nodes on stems and inflorescences, deltoid, peltate, tiny, caducous, brown. Leaves simple, opposite or rarely whorled, coriaceous or chartaceous, shape variable, glabrous; margin entire (or crenate in the naturalised G. livingstonei), mostly recurved; venation brochidodromous or paxillate; primary vein raised or flush; exudate-containing canals as parallel or random lines, branched or unbranched, continuous or broken dashes and dots. Inflorescences terminal, axillary or ramiflorous; bracts opposite, leaf-like with a primary vein, occurring at peduncle apex on racemose or paniculate species. Bracteoles mostly opposite, convex or cupular, occurring at pedicel base on most species. Flowers solitary or in cymes, compact panicles, racemes or fascicles, up to 66-flowered; actinomorphic, hypogynous, unisexual, may be fragrant, sessile or pedicellate; sepals 4 or 5, free or joined; petals 4 or 5, free, convex, cupular, erect to almost explanate. Male flowers: androecium of fused stamens as phalanges, free stamens or fused filaments as a receptacle, pistillode may be present; anthers mostly bithecate, sessile or with a 4 short filament. Female flowers: staminodal phalanges present or absent; ovary superior, 2-13-locular, 1-4 ovules per locule; stigma sessile or with a short style, discoid or lobed. Fruit a fleshy or dry berry, sometimes with seams or furrows along septal radii, Austrobaileya 9(1): 1-29 (2013) 2-13-locular; seeds 1-13, 1-4 per locule; aril present in most species. Germination is hypogeal or epigeal. Etymology : The generic epithet Garcinia is named for the Dutch army doctor and naturalist Laurentius Garcin (1683-1751). Key to Garcinia species in Australia 1 Stigma on flowers and fruit disk-like and entire.2 1. Stigma on flowers and fruit rayed, lobed or crenate. 4 2 Leaves always opposite; flowers with stamens united as phalanges and surrounding a fungiform pistillode; fruit red or blackish. 3 2. Leaves mostly 3-whorled (occasionally opposite or 4-whorled); flowers with free stamens, pistillode absent; fruit yellow to orange. 3 . *G. livingstonei 3 Fruit much longer than wide, occurs in the Wet Tropics area.5. G. zichii 3 . Fruit equidimensional, occurs north from the Mcllwraith Range. 4 . G. jensenii 4 Seeds and ovules usually more than one per locule.5 4 . Seeds and ovules only one per locule.6 5 Leaves peltate.11. G. russellii 5. Leaves never peltate.10. G. gibbsiae 6 Exudate in stems and leaves white; petals 5.7 6. Exudate in stems and leaves yellow; petals 4.8 7 Secondary veins 20-25 pairs.2. *G. xanthochymus 7. Secondary veins 9-14 pairs.1. G. dulcis 8 Leaves with 10 pairs of secondary veins; exudate-containing canals as short streaks, dashes and dots, unbranched.7. G. mestonii 8. Leaves with 12 or more pairs of secondary veins; exudate- containing canals as continuous long lines.9 9 Leaves with 2 distinct intramarginal veins.8. *G. mangostana 9. Leaves with 1 intramarginal vein or intramarginal vein absent or not obvious 10 10 Leaves almost membranous; petals up to 3 mm long.6. G. leggeae 10. Leaves coriaceous; petals 5 mm or more long.11 11 Leaves with tertiary veins absent.9. G. brassii 11. Leaves with tertiary veins present.12 12 At least some or most tertiary veins diminishing and not anastomosing . . . . 9. G. brassii 12. All tertiary veins anastomosing.12. G. warrenii 1. Garcinia dulcis (Roxb.) Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 43: 88 (1874); Xanthochymus dulcis Roxb., PI. Coromandel 3: 66 (1820). Type: “ex Molucca Islands, cult. In Indian Botanic Garden, Calcutta” n.v.,fide Maheshwari (1964: 116). Illustrations : Hooker (1831: t. 3088); Wight (1840: 1: t. 192); van Nooten (1880: t. 15); Christophel & Hyland (1993: 81); Cooper & Cooper (2004: 123); CANBR (2010); Hyland etal. (2010); Sompong (2007: 248-250). 5 Cooper, Garcinia in Australia Tree to 20 m, dioecious; exudate white in stems and petioles, yellow in fruit; bark with fine vertical fissures, becoming thin flakey on older trees; branchlets square, ribbed, pendulous, glabrous. Leaves discolorous, coriaceous, glabrous; petioles 10-30 mm long, channelled, ligulate; lamina elliptic, ovate or oblong-ovate, 75-280 mm long, 32-180 mm wide; base attenuate, cuneate, obtuse, cordate or asymmetrical; apex shortly acuminate, acute or obtuse; margin recurved; venation brochidodromous, primary vein ± flush on upper surface and protruding on abaxial surface; secondary veins 9-14 pairs, raised on both sides in dried specimens, angle of divergence from primary vein 30-60°, forming loops within 1.5-5 mm from margin; exudate-containing canals on upper side are numerous many-branched wriggly lines running more or less parallel to secondary veins, on the underside they are numerous parallel branched or unbranched lines which run at angles between being parallel to primary vein and margin, sometimes running parallel to secondary veins. Male and female inflorescences similar, axillary or ramiflorous, a raceme or a fascicle on a pulvinus, 1-many-flowered; bracteoles persistent, clustered at base of pedicels, ovate, convex, keeled, 1-2.25 mm long, 1-2 mm wide, base truncate, apex obtuse or acute, entire, minutely hairy especially around margin, sparsely hairy abaxially. Male flowers: fragrant, diameter 11-12 mm, glabrous; pedicels 7-14 mm long; sepals 5, free, orbicular, 2-4 mm long, 3-4 mm long, green, white or cream, margin minutely ciliate; petals 5, free, erect, imbricate, orbicular, convex, 5-7 mm long, 5-7 mm wide, membranous, green, lime-yellow, cream or yellow, glabrous, margin thin and minutely ciliate; androecium 5-phalangiate, glabrous; phalanges antepetalous, strap-shaped at base and flabellate towards apex, introrse, 8-9 mm long, 0.8-1.4 mm wide, apically branched into 5-10 filaments, filaments 0.3- 0.7 mm long, 1 anther per filament; anthers bithecate, introrse, broadly-globose, c. 0.2 mm long and 0.3 mm wide; disk diameter c. 4 mm, pitted or honeycombed; pistillode rudimentary, conical-attenuate c. 0.5 mm long. Female flowers: fragrant, diameter 8-9 mm; pedicels 9-17 mm long; sepals 5, free, imbricate, orbicular, convex, 3-4.5 mm long, 4.5-6.5 mm wide, green or yellowish, glabrous, margin minutely ciliate; petals 5, free, imbricate, orbicular, convex, 7-8 mm long, 7-8 mm wide, green, cream or yellow, glabrous, margin minutely ciliate; staminodal phalanges 5, antepetalous, solitary and filament-like or 2 or 3-fused, distally indexed, 2-3.5 mm long, 2 or 3-branched at apex, filaments 0.9-1.4 mm; antherodes 1-3 per phalange, introrse, bilobed, flattened, 0.2-0.3 mm long, c. 0.3 mm wide; nectaries alternating between phalanges, c. 2 mm long and 2 mm wide, pitted; disk c. 4 mm wide; ovary globular, c. 5 mm long and 4 mm wide, sessile, 5-locular, locules uniovulate; stigma 5-branched, style c. 1.8 mm long; stylar arms 5, c. 2.5 mm long. Fruiting pedicel c. 15 mm long, 5 sepals persistent at base; fruit a fleshy berry, ovoid, globular or depressed globular, apex often beaked, 35-50 mm long, 45-53 mm wide, yellow, glabrous; stigma 5-rayed, diameter 2-3 mm; seeds usually 5, c. 30 mm long and 20 mm wide; testa brown, smooth; aril fleshy, cream or yellow. Claudie mangosteen, gourka, mundu tree. Fig. 1A & B, 2B. Additional selected specimens (from 47 examined): Malaysia. Kelantan: Pasir Putih, Semarak Bt., Sep 1992, FRI37081 (CNS). Papua New Guinea. West New Britain District: 32 km SW of Linga Linga Plantation, May 1973, Isles & Katik NGF32248 (CNS). Milne Bay Province: Esirava Creek, Tagula Island, Mar 1979, Damas LAE74527 (CNS). Australia. Queensland. Cook District: Banks Peak, Moa Island, Torres Strait, Jul 2008, Fell 9722 & Stanton (BRI); Moa (Banks) Peak, Moa Island, Torres Strait, Feb 1989, Gray 4993 (CNS); Escarpment of Great Dividing Range, 14.6 km NE of Heathlands Ranger Base, Heathlands D & O Reserve, Oct 1993, Fell 3746 & Stanton (BRI); CSIRO Plot, West Claudie River, Dec 2011, Cooper 2216 & Venables (CNS); Gordon Creek, Oct 1972, Hyland 6416 (CNS); Claudie River, Oct 1981, Hyland 11181 (CNS); Iron Range, base of Lamond Hill, Sep 1988, Sankowsky 861 (CNS); Nesbit River, Silver Plains Holding, Oct 1999, Hyland 16253 (CNS); Nesbit River, Silver Plains Holding, Oct 1999, Hyland 16254 (CNS); Junction of Leo Creek and Nesbit River, Aug 1993, Fell 3385, Jensen & Barnes (BRI); Chester River Scrub, Jun 1992, Forster PIF10440 & Tucker (BRI); TR 14, Leo Creek Road, Dec 1979, Hyland 10132 (CNS); TR 14, Leo Creek Mine area, Mcllwraith Range, Jun 1992, Forster P1F10236, Tucker & Kenning (BRI); Rocky River crossing, Oct 1996, Cooper 1016 & Cooper (CNS); Rocky River catchment. A ustrobaileya 9(1): 1-29(2013) Fig. 1. Flowers. A. Garcinia dulcis showing rayed stigma at ovary apex {Hyland 16253 [CNS]). B. G. dulcis S, showing apically branched phalanges {Hyland 16254 [CNS]). C. G. mestonii S, showing small opening and anthers {Sankowsky 2326 [BRI]). D. G. jensenii showing 3-flowered inflorescence and yellow stigma at apex of fungiform gynoecium {Cooper 2225, Zdenek, Venables & Pritchard [CNS]). E. G. leggeae S showing anthers {Cooper 2175, Jensen & Kemp [CNS]). F. G. mestonii showing small opening and stigma {Forster PIF24028 [CNS]). G. G. zichii % showing viscous fungiform gynoecium {Sankowsky 2547 [BRI]). H. G. zichii 3 , showing viscous fungiform pistillode and anthers at apices of 4-phalangiate androecium {Dowe 1159 [CNS]). Photos: A, B, F, CSIRO; C, G, G.Sankowsky, D, B. Venables, E, R. Jensen, H, J.Dowe. Cooper, Garcinia in Australia 7 Fig. 2. Exudate-containing canals on leaf undersides shown crossing secondary venation. A. Garcinia brassii showing as pale lines {Cooper 2163 & Ford [CNS]). B. G. dulcis showing as pale lines and reticulations {Cooper 2216 & Venables [CNS]). C. G.jensenii showing as dark lines {Cooper 2194, Jensen & Kemp [CNS]). D. G. mestonii showing as pale wriggly dashes and dots {Cooper 2203 & Jensen [CNS]). E. G. gibbsiae showing as pale lines {Cooper 2212 [CNS]). F. G. leggeae showing as dark lines {Cooper 2187, Jensen & Kemp [CNS]). G. G. warrenii showing as dark lines {Cooper 474 [CNS]). H. G. livingstonei showing as dark lines {Cooper 2218 [CNS]). Photos: A, B, C, D, E, G, H W.Cooper ; F, R. Jensen. 8 Silver Plains, Aug 1997, Cooper & Jensen 44 (CNS); Upper Massey (Massy) Creek, Oct 1962, Smith 11719 (BRI); Melville Range 4 km N of Abbey Peak, Nov 1981, Irvine 2195 (CNS). Distribution and habitat : Garcinia dulcis is widespread from Australia and New Guinea to South East Asia and India. Within Australia, it is restricted to north Queensland where it occurs in rainforest (complex, semideciduous notophyll and mesophyll vine forest) from the Melville Range on Cape York Peninsula to the Torres Strait islands, altitude 10-500 m (Map 1 ) Phenology : Flowers have been recorded from September to February; fruit has been recorded from August to October and January to March. Etymology : The specific epithet is derived from the Latin dulcis (sweet or pleasant) referring to the sour but palatable aril. 2. *Garcinia xanthochymus Hook.fi ex T.Anders., FI. Brit. India 1: 269 (1874); Xanthochymus pictorius Roxb., PI. Coromandel 2: 51, t. 196 (1805); Stalagmitis pictoria (Roxb.) G.Don., Gen. Syst. 1: 620 (1831); Xanthochymus tinctorius DC., Prodr. 1: 562 (1824) [sphalm. for pictorius] ; Garcinia tinctoria (DC.) W.F.Wight, Bull. Bur. PI. Industr. U.S.D.A. 137: 50 (1909). Type: India: Circars, s.dat., Roxburgh s.n. (holo: BR, herb. Mart.). Illustrations : Roxburgh (1805, 2: t. 196) as Xanthochymus pictorius Roxb.; Kirtikar & Basu (1918, 1: t. 104); Sweeney (2008: 1289, K; 2010: 158, 160, 162); Sharma et al. (2012: 8,9). Shrub or tree to 20 m, dioecious; exudate yellow or milky; bark dark brown, with short vertical fissures, flaky on older trees; branchlets square, mostly pendulous, twigs distinctly 6-8-ribbed. Leaves discolorous, coriaceous, glabrous, shiny; petioles 10-25 mm long, ligulate; lamina oblong, linear- oblong, oblong-lanceolate, oblong-ovate or ovate, 120-410 mm long, 38-120 mm wide, base cuneate or rounded; apex acute, acuminate or apiculate; margin recurved; venation brochidodromous, primary vein slightly raised on upper side and distinctly Austrobaileya%\). 1-29(2013) raised on underside; secondary veins 20-25 pairs raised on both sides, angle of divergence from primary vein 50-60°, forming loops within margin, intramarginal vein 0.5-5 mm from margin; exudate-containing canals sparse on the underside as faint long lines running at angles between parallel to the margin and the primary vein. Male flowers: not seen for Australian material, recorded as: diameter 15 mm, fascicles 4-8-flowered; sepals 5; petals 5, diameter c. 8 mm; androecium 5-phalangiate; phalanges strap-shaped at base and flabellate towards apex, indexed, apically branched into 3-6 filaments; anthers 2-celled, introrse; nectaries in between phalanges. Female inflorescences axillary or ramiflorous, a reduced raceme, fascicle-like, 2-14-flowered. Female flowers: fragrance not recorded; diameter 10—12(— 20) mm; pedicels (18-)24-36 mm long; sepals 5, free, suborbicular, convex, 3.5-4 mm long, 6-7 mm wide, green or whitish, glabrous, margin minutely ciliate; petals 5, free, orbicular, convex, erect, 7.5-8 mm long, 10-11 mm wide, white, cream or pale yellow, margin minutely ciliate; staminodal phalanges 5, antepetalous, solitary and slender or 3 or 4-fused and strap-shaped at base and flabellate towards apex, indexed, c. 3 mm long, branched into 3 or 4 filaments at apex, filaments 0.5-0.7 mm long, antherodes solitary or 3 or 4 per phalange, introrse, bilobed, flattened, c. 0.3 mm long and 0.5 mm wide; nectaries alternating between phalanges, c. 2 mm long and 2.5 mm wide, pitted; disk c. 5 mm wide; ovary globular, c. 4 mm long and 6 mm wide, sessile, 5-locular, locules uniovulate; stigma 5-branched, style c. 1.7 mm long; stylar arms 5, 2.5-3 mm long. Fruiting pedicel to 40 mm long, sepals persistent at base; fruit a fleshy berry, often asymmetrical with beak not centred, ovoid or globose, beaked, 38-65 mm long, 35-50 mm wide, yellow or orange; stigma 5-rayed, diameter 4-5 mm; seeds 1-5 (usually 1 or 2), 25-35 mm long, 15-18 mm wide, testa brown, veined; aril fleshy, yellow or orange. sour mangosteen, false mangosteen , yellow mangosteen, gamboge tree , eggtree. Additional selected specimens (from 6 examined): Australia. Queensland. Cook District: Tributary of Cooper, Garcinia in Australia Bushy Creek, Rumula, Oct 1998, Holmes 66 (CNS); Cultivated, Cairns Botanic Gardens, Feb 2013, Cooper 2217, Cooper & Venables (CNS); Wadda Creek, East Palmerston, Sep 1941, Wilson s.n. (CNS); Patrick Road off Dunne Road, East Palmerston, Dec 2005, Jensen 1486 & Cauley (BRI); Tully, Dec 1945, Shearer s.n. (CNS). North Kennedy District: Delta, 3 miles [5 km] W of Bowen, Nov 1971, Wright s.n. (BRI [AQ313]). Moreton District: Rosemount Road, Nambour, Nov 1989, Brufords.n. (BRI [AQ458316]). Distribution and habitat : Garcinia xanthochymus is a native of Bangladesh, Bhutan, Cambodia, China, India, Laos, Myanmar, Nepal, Thailand and Vietnam. It is cultivated for its edible sour fruit and has become naturalised in a few locations in Queensland from the Rumula area south to Nambour (Map 6), altitude 20-500 m. Phenology : Flowers have been recorded from February to May. Fruit has been recorded from February to May, November and December. Typification : Garcinia xanthochymus was named by Anderson in Hooker’s Flora of British India (1874); however, nomenclature for this species commenced nearly 70 years earlier when Roxburgh (1805) described Xanthochymus pictorius Roxb. (non Garcinia pictoria Roxb.). De Candolle (1824) did not help the situation by totally misprinting (or perhaps ‘correcting’) the epithet as X. tinctorius DC. (Maheshwari 1964; Singh 1993). Anderson (1874) interpreted de Candolle’s (1824) account as a misspelling, so coined the new name G. xanthochymus with Roxburgh’s X. pictorius and de Candolle’s X. tinctorius both in synonymy. Much later, D’Arcy (1980) overlooked the name Garcinia pictoria Roxb. and published the superfluous new combination G. pictorius (Roxb.) D’Arcy based on Xanthochymus pictorius Roxb. Anderson (1874) did not unambiguously specify a type collection for Garcinia xanthochymus , but did list X. pictorius Roxb. and the misprinted X. tinctorius DC. in synonymy, together with references to other literature and one specimen in the Wallich Catalogue (no. 4837). Under the distribution notes for this species Anderson (1874) cited a single specimen collected by Roxburgh from the Circars. Maheshwari 9 (1964) listed the type for G. xanthochymus as this Roxburgh collection (deposited in BR) and this interpretation appears to have been universally accepted by later authors (Kostermans 1980; Singh 1993). Notes : Sharma et al. (2012) suggested that G. xanthochymus produces bisexual flowers; however they concluded that functional bisexuality had not been confirmed. Agamospermy is known or suspected for ten Garcinia species (not including G. xanthochymus) (Richards 1990a). Separate male and female flowers have been observed and illustrated by Sweeney (2010). Etymology : The specific epithet is derived from the Greek xanthos (yellow) and chymos (sap); referring to the exudate. 3. *Garcinia livingstonei T.Anders., Bot. J. Linn. Soc. 9: 263 (1866). Type: cult. In hort. bot. Calcutta, ex Mozambique, “in rupibus schistosis prope flumen Zambesi”, s.dat., Kirk s.n. (holo: CAL);, fide Robson (1960: 400). Illustrations : Sweeney (2008: 1289, Q). Shrub or tree to 18 m, monoecious; exudate yellow or reddish; bark grey and rough with numerous vertical fissures; branches terete, arching. Leaves discolorous, opposite or 3-4-whorled, coriaceous, glabrous; petioles 2-7 mm long, channelled, ligulate; lamina oblong, elliptical or oblanceolate, 35-110 mm long, 20-55 mm wide; base cuneate, rounded or cordate; apex acute, rounded, emarginate or apiculate; margin crenate or entire, recurved; venation brochidodromous, primary vein raised in a groove adaxially and distinctly raised abaxially; secondary veins 9-15 pairs, pale yellow, raised on both surfaces, angle of divergence from primary vein 60-70°; intramarginal vein merges with margin; exudate-containing canals on upperside are numerous branched lines ± parallel to secondary veins, on the lower side they are branched lines running at angles between the primary vein and margin. Male and female inflorescences: similar, axillary or ramiflorous, a fascicle, 2-7-flowered; bracteoles ± quadrate or rhombic, slightly convex, 1-1.5 mm long, 1-1.5 mm wide, base truncate, apex acute, margin 3-toothed, 10 glabrous. Male flowers: fragrant, diameter 8-10 mm; pedicels 6.5-8 mm long, softly muricate; sepals 4,2-whorled, free, D-shaped, c. 1 mm long and 1.8 mm wide, green, margin entire; inner pair orbicular, convex, c. 2 mm long and 2.5 mm wide, green or yellowish- green, margin entire; petals 5, free, orbicular, reflexed, 4.5-5.5 mm long, 3.5-4.5 mm wide, membranous, green or greenish-yellow, glabrous; androecium a globose yellow disk with c. 30 free stamens radiating like needles in a pin cushion, disk sunken directly below each stamen, filaments 1.5-1.7 mm long; anthers bithecate, globose, diameter c. 0.3 mm; pistillode absent. Female flowers: fragrant, diameter c. 7 mm; pedicels 16-20 mm long, softly muricate; sepals 4, free, green; outer pair D-shaped, c. 1.2 mm long and 1.4 mm wide; inner pair ovate, c. 2.5 mm long and 2 mm wide, glabrous; petals 5, free, orbicular or obovoid, c. 8 mm long and 6 mm wide, green or greenish-yellow; disk oblate, yellow, with c. 26 free staminodes, diameter c. 3.5 mm, sunken directly below each staminode; staminodes c. 2 mm long; antherodes 1 per staminode, ± globose, bilobed, diameter c. 0.5 mm; ovary obovoid, c. 3 mm long and 3 mm wide, 2-locular, locules uniovulate; stigma domed, not rayed. Fruiting pedicel: not seen for Australian material, recorded as: at least as long as fruit, staminodes usually persistent at base, sepals absent at base; fruit (not seen for Australian material) a fleshy berry, globular or obovoid, 10-30 mm long, 10-30 mm wide, orange or reddish-orange; seeds 1 or 2, cylindrical, 8-20 mm long; aril fleshy, orange. African mangosteen , lowveld mangosteen, imbe. Fig. 2H. Additional selected specimens (from 54 examined): Senegal. Ouassadou, Jan 1959, Berhaut 4302 (P 1 ). Mali. Bankoumana, Nov 2005, Birnbaum 920 (P 1 ); Tienfala (25 km E Bamako), Dec 1978 (P 1 ). Cameroon. Southwest Province: Mt Kupe, Feb 1986, Thomas & McLeod 5480 (P 1 ). Comoros. Mayotte, Sohoa, May 1997, Pascal 934 (P 1 ); Itsamia, Nov 1999, Labat 3223 & Yahaya, Dloubeiri & Mindhiri (P 1 ). Kenya. Sep 1957, Verdcourt 1936 (P 1 ). Tanzania. 40 km west of Lindi, Sep 1934, Schlieben 5402 (P 1 ); Zanzibar, Nov 1889, Sacleux 886 (P 1 ); Pwani Coast Region, Matumbi Hills, Oct 1997, Phillipson 4960, Sitoni & Kibure (P 1 ). Zimbabwe. Urungwe District, Oct 1964, Bingham 1370 (P 1 ). Mozambique. Gaza, Nov 1970, Correia 2015 (P 1 ); Lugela-Mucuba, Namagoa, s.dat., Faulkner Kew 77 (P 1 ). Australia. Queensland. Cook District: Cooktown, Austrobaileya 9(1): 1-29 (2013) Anzac Memorial Park, Sep 2008, McKenna 267 (CNS); Helen St, Cooktown, May 2013, Cooper 2218 & Cooper (CNS); Trevethan Creek, Cooktown Development Road, May 1998, Jago s.n. & Wannan (CNS). Distribution and habitat: Garcinia livingstonei is a native of Africa with a widespread distribution, occurring on the mainland from West Africa (including Senegal, Mali, Nigeria and Cameroon) through to Somalia and East Africa (including Kenya and Tanzania) and south to Zimbabwe, Angola and South Africa; as well as the Comoros islands. In Australia it has been cultivated and a recent record ( Jago s.n. & Wannan CNS) from Trevethan Creek near Cooktown indicates possible naturalization (Map 1) Phenology: Flowers from Australian material have been recorded in May and no fruiting records exist. Notes: Cultivated plants of Garcinia livingstonei in Cooktown, which had been poisoned, were sprouting vigorous suckers from the base of otherwise dead trees. One tree at Cooktown {Cooper 2218 ) had both female and male flowers confirming that this species is monoecious. Etymology: The specific epithet is in honour of the explorer David Livingstone (1813— 1873), who was amongst the first to collect this species. 4. Garcinia jensenii W.E.Cooper sp. nov. Similar to Garcinia hunsteinii Lauterb. but differs in the leaf apex (bluntly acute versus acuminate), secondary veins (7-10 pairs versus c. 50 pairs), and secondary vein angle (c. 45° versus c. 80°). Typus: Australia: Queensland. Cook District: about 3 km N of 3-ways, Iron Range National Park, 29 July 2013, W.Cooper 2225, C.Zdenek, B.Venables & J.Pritchard (holo: CNS [2 sheets + spirit], iso: 13 sheets to be distributed to A, BRI, CANB, FRI, K, KEP, KL, MEL, MO, NSW, NY, SING, US). Garcinia riparia auct. non A.C.Sm.; Hyland etal. (2003, 2010); CHAH (2007). Garcinia sp. (Claudie River L.J.Brass 19658) in part; Jessup (2007: 54, 2010: 44). Cooper, Garcinia in Australia Illustrations : Christophel & Hyland (1993: 81) as G. riparia vel aff; Hyland et al. (2003, 2010), CHAH (2007) as G. riparia; Zodiac Publications (undated), as Garcinia sp. Claudie River. Tree to 15 m, dioecious; exudate white or colourless; dbh to 150 mm; bark slightly flakey with fine vertical fissures, lenticels horizontal and more prevalent on younger trees; branchlets square, young twigs narrowly 4-winged. Leaves discolorous, coriaceous, glabrous; petioles 5-7 mm long, channelled, ligulate; lamina narrowly to broadly elliptic, ovate-elliptic or obovate, 52-160 mm long, 20-72 mm wide; base cuneate, attenuate or decurrent; apex acute, retuse-acute or rounded-acute; margin entire; venation brochidodromous, primary vein on upperside flush or slightly raised, distinctly raised on underside; secondary veins 7-10 pairs, slightly raised on both surfaces, angle of divergence with primary vein c. 45°, forming intramarginal loops 0.4-1.4 mm from margin; exudate-containing canals numerous and conspicuous on the underside as unbranched lines ± parallel to each other and running at angles between primary vein and margin. Male inflorescences axillary, a fascicle or raceme, 5-many-flowered, rachis up to 3.5 mm long; bracts at pedicel bases broadly ovate, apex acute, base truncate, convex, c. 1.6 mm long and 1 mm wide, glabrous; bracteoles present or absent, part way along pedicels, solitary or paired, broadly ovate, convex, c. 1 mm long and 1 mm wide, glabrous. Male flowers: fragrance not recorded, diameter 7-8 mm; pedicels 4.5-12 mm long; sepals 4, 2-whorled, colour not recorded but probably white or green white like females, glabrous; outer pair joined, D-shaped, convex, may be keeled, c. 1.6 mm long and 1.8 mm wide; inner pair free, petal-like, orbicular, convex, c. 4 mm long and 3 mm wide; petals 4, free, imbricate, orbicular, convex, c. 4 mm long and 3-4 mm wide, membranous, colour not recorded but probably white like females, glabrous; androecium 4-phalangiate, glabrous; phalanges antepetalous, strap¬ shaped, slightly ribbed, erect or distally indexed, c. 3.2 mm long and 2-3 mm wide, apex divided into 2-7 short filaments; ll anthers 15-30 per phalange, completely enveloping phalange apex, D-shaped, circular or irregular, margin lobed, erect or introrse, diameter 0.5-6 mm; pistillode fungiform, c. 3 mm long and 2 mm wide at apex, apex colour not recorded but probably yellow like females. Female inflorescences terminal or axillary, supra-axillary or ramiflorous, a solitary flower, fascicle or condensed raceme, 3-9-flowered; peduncles 1.2-2 mm long; bracteoles triangular, keeled, c. 1 mm long and 1 mm wide, entire, persistent. Female flowers: fragrance not detected, diameter 4.5-6.5 mm; pedicels 3.5-4.5 mm long; sepals 4, 2-whorled, D-shaped, convex, white or palest green, glabrous; outer pair joined, c. 1 mm long and 2 mm wide, coriaceous; inner pair free, petal-like, orbicular, convex, 1.2-2 mm long and 2-3 mm wide, membranous; petals 4, free, imbricate, orbicular, convex, c. 2 mm long, 2-3 mm wide, margin thin, white, glabrous; gynoecium fungiform, unlobed, glabrous; ovary obovoid and laterally compressed, c. 1.2 mm long and 1.8 mm wide, 2-locular, uniovulate; stigma disc¬ like, entire, diameter c. 2.5 mm, yellow, not viscous. Fruiting pedicel c. 5 mm long; 2 joined sepals persistent at base; fruit a fleshy berry, globose or ellipsoid-globose, 12.5-15.5 mm long, 11-16 mm wide, pink or reddish; stigma at apex, diameter 3-3.8 mm; seeds 1 or 2, c. 10 mm long and 5 mm wide; testa and aril not recorded. Fig. ID, 2C. Additional selected specimens (from 23 examined): Australia. Queensland. Cook District: Widul Island, 4 km west of Mabuyang Island Airfield, 73 km NNW of Horn Island Airfield, Apr 2009, Fell 10037 (CNS); Ulu (Saddle Islet), 62 km NE of Horn Island Airfield, Torres Strait, Feb 2012, Fell 10712 (CNS); Eet Hill Track, Moa Island, Torres Strait, May 1987, Budworth 1185 (BRI); c. 2 km N of St Paul’s community, Moa Island, 46 km NNE of Horn Island Airfield, Torres Strait, Nov 2007, Fell 8955 (CNS); Mew River, Muddy Bay, Cape York, Jun 1994, Forster PIF15300 & Tucker (BRI); Lake Boronto - Newcastle Bay, Cape York Peninsula, Sep 1974, Tracey 14349 (BRI); Between Lockerbie & Somerset, Feb 1980, Hyland RFK3984 (CNS); Lockerbie Scrub, Sep 1974, Tracey 14306 (BRI); 5.5 km NE of Moreton, Jun 1994, Fell 4457 & fiMdt(BRI); Olive River, N bank, Sep 1974, Traceyl4373 (BRI); Nelson Creek, Aug 2011, Cooper 2145, Venables & Murphy (CNS); Iron Range NP, southside of Pascoe River, Sep 2004, Sankowsky 2517 & Sankowsky (BRI); about 3 km N of 3-ways, Iron Range NP, Dec 2012, Cooper 2194, Jensen & Kemp (CNS); Iron Range, Jul 1923, Brass 19658 (BRI); about 2.8 km 12 N of 3-ways Iron Range NP, Jun 2013, Cooper 2219, Cooper & Venables (CNS); Chester River, Jun 1977, Hyland RFK3555 (CNS); TR 14, Leo Creek Mine area, Mcllwraith Range, Jun 1992, Forster PIF10160, Tucker & Kenning (BRI); Mcllwraith Range, E foothills. Rocky River, Oct 1969, Webb 9345 & Tracey (BRI). Distribution and habitat : Garcinia jensenii grows on red soils derived from a mixture of basic igneous rocks and ferrugineous sandstone. It is as an understory tree in rainforest (semideciduous notophyll/ mesophyll vine forest) from the Torres Strait islands and down the eastern side of Cape York Peninsula to the Mcllwraith Range at altitudes between sea level and 500 m (Map 2 ). Garcinia jensenii occurs in rainforest dominated by Archidendron hirsutum I. C.Nielsen, Buchanania arborescens (Blume) Blume, Calamus caryotoides A. Cunn. ex Mart., Dillenia alata (R.Br. ex DC.) Martelli, Euonymus australiana F.Muell., Hydriastele costata F.M.Bailey, Maranthes corymbosa Blume, Planchonella xylocarpa C.T.White, Ptychosperma elegans (R.Br.) Blume, Syzygium bamagense B. Hyland & Syzygium pseudofastigiatum B. Hyland. Phenology : Flowers have been recorded in July and August; fruits have been recorded in December and February. Affinities : Within Australia, Garcinia jensenii is most similar to G. zichii W.E.Cooper. Both species have similarly shaped leaves and exudate canal pattern, a 2-locular ovary and 1 or 2-seeded fruit. G. jensenii has leaves with 7-10 pairs of secondary veins, flowers with white petals and fruit that are globose or ellipsoid-globose. G. zichii has leaves with 12—18 pairs of secondary veins, flowers with green or yellow petals and fruit that are distinctly ellipsoid. Notes: In Queensland, Garcinia jensenii has been confused with G. riparia that is considered to be endemic to New Guinea; however, the two species are significantly different morphologically. G. jensenii has white or colourless exudate, flowers pedicellate, ovary 1 or 2-locular, fruit not ribbed and 1 or 2-seeded. G. riparia has yellow Austrobaileya 9(1): 1-29 (2013) exudate, flowers sessile with a 7-10-locular ovary, fruit ribbed and many-seeded. Etymology: The specific epithet honours botanist Rigel Jensen (1959-) acknowledging his contribution to botany in Australian and New Guinea, as well as his enthusiastic assistance with all of my botanical projects. 5. Garcinia zichii W.E.Cooper sp. nov. Similar to Garcinia hunsteinii but differs in the leaf apex (acute versus acuminate), lateral veins (7-10 pairs versus c. 50 pairs), lateral vein angle (c. 45° versus c. 80°) and the fruit shape (ellipsoid versus globose). Typus: Australia: Queensland. North Kennedy District: Tully Falls National Park, 2.2 km along Tully Falls Road from Charmillan Creek bridge, 8 November 2011, A. Ford 5908 & M. Torello-Raventos (holo: CNS [2 sheets + spirit], iso: 10 sheets to be distributed to BRI, CANB, K, KL, L, MEL, MO, NSW, SING). Garcinia sp. aff. G. hunsteinii ; Hyland (1982); http://www.anbg.gov.au/cgi-bin/apni Accessed January 2013. Garcinia sp. (Davies Creek JGT 14745); Hyland etal. (2003). Garcinia sp. (Davies Creek); Cooper & Cooper (2004: 124). Garcinia sp. (Davies Creek J.G.Tracey 14745); Jessup (2007: 49, 2010: 44). Garcinia sp. Davies Creek (J.G.Tracey 14745); CHAH (2007); Hyland et al. (2010). Garcinia sp. (=AFO/1268); http://www.anbg. gov.au/cgi-bin/apni Accessed January 2013. Illustrations: Christophel & Hyland (1993: 81) as G. sp. aff. G. hunsteinii ; Hyland et al. (2003) as G. sp. (Davies Creek JGT 14745); Cooper & Cooper (2004: 124) as G. sp. (Davies Creek); Hyland et al. (2010), as G. sp. Davies Creek (J.G.Tracey 14745). Small tree or canopy tree to 30 m, dioecious; exudate colourless, white or slightly milky, meager; dbh to 20 cm, may have small buttresses; bark rough, fissured, slightly flaky, lenticels pale orange; branchlets square, deeply grooved between last 2 pairs of leaves, becoming terete. Leaves discolorous, Cooper, Garcinia in Australia coriaceous, glabrous; petioles 5-10 mm long, channelled, ligulate; lamina elliptical or rarely obovate, 35-110 mm long, 15-35 mm wide; base attenuate, cuneate or decurrent; apex bluntly acute; margin flat or may be recurved; venation brochidodromous, primary vein distinctly raised on upperside, slightly raised near base on underside and flush or slightly depressed towards apex; secondary veins obscure, 12-18 pairs, flush on both sides, angle of divergence with primary vein c. 60°, forming intramarginal loops c. 0.5 mm from margin; exudate-containing canals numerous and conspicuous on the underside as unbranched lines ± parallel to each other and running at angles between primary vein and margin. Male inflorescences axillary or rarely terminal, an umbel or rarely a cyme, 2-11-flowered (mostly 3-5-flowered), peduncles 0.8-2.5 mm long; bracts at peduncle bases and bracteoles at pedicel bases similar, triangular or broadly rhomboid, convex, keeled, 0.8-2.6 mm long, muricate, apex serrate with 3-7 teeth, caducous. Male flowers: not fragrant, diameter 7.5-8.5 mm, glabrous; pedicels 3.5-5 mm long; sepals 4, outer pair joined, inner pair free, imbricate, D-shaped, convex but not keeled, 2-4 mm long, 2-4 mm wide, coriaceous-membranous, yellow or green, glabrous, margin thin; petals 4, free, orbicular, convex, not keeled, becoming reflexed but remaining incurved at apex, 3-4.5 mm long, 3-4.5 mm wide, membranous, green; androecium 4-phalangiate; phalanges antepetalous, strap-shaped, ribbed, erect and becoming slightly reflexed, 1-4 mm long, 1-2 mm wide, apex undivided or divided into two short arms; anthers c. 20 per phalange, completely enveloping phalange apex, oblong, 0.8-1 mm long, c. 0.3 mm wide; pistillode fungiform, 2-2.5 mm long, 2-2.8 mm wide at apex, apex dark orange-red or reddish-brown and quickly becoming viscous once flowers are open. Female inflorescences axillary, an umbel, 3-5-flowered, peduncles 1-2 mm long; bracteoles triangular, c. 1.2 mm long and 1 mm wide, muricate, margin 3-5-toothed, caducous. Female flowers: occasionally fragrant, diameter 6-8 mm; pedicels 2-5 mm long; sepals 4, D-shaped or deltoid, convex, not keeled or slightly keeled, 13 coriaceous-membranous, yellow or green, margin thin; outer pair joined, c. 2 mm long, 2-3 mm wide; inner pair free, 2-4 mm long, 2.5-4 mm wide; petals 4, free, imbricate, becoming reflexed, orbicular, convex, not keeled, 2.2-4.2 mm long, 2.8-4.2 mm wide, coriaceous-membranous, yellow or green, margin thin; gynoecium fungiform, glabrous; ovary ellipsoid, 0.6-1.5 mm long, 4-7 mm wide, 2-locular, uniovulate; stigma disc-like, entire, diameter 4-5 mm, yellow, orange, rusty red or brownish-red, quickly becomes viscous once flowers are open. Fruiting pedicel 2.5-5 mm long; sepals absent from base; fruit a fleshy berry, ellipsoid, 25-37 mm long, 10-18 mm wide, red to dark red or blackish; stigma at apex, diameter 4-6 mm; seeds one or two, surface slightly undulating, 22-32 mm long, 11-12 mm wide; testa brown; aril meagre, clear, gelatinous, baconwood , marblewood. Fig. 1G & H. Additional selected specimens (from 118 examined): Australia. Queensland. Cook District: Cedar Bay NP, western slopes of Mt Finnigan, Oct 1999, Forster PIF25016 & Booth (BRI); Mt Hemmant just N of Noah Creek in Cape Tribulation area, Jul 1973, Webb 11967 & Tracey (BRI); Mt Windsor NP, Nov 2008, Forster PIF34675, Jensen & McDonald (BRI); Mt Misery, Jun 1992, Forster PIF10759, Sankowsky & Tucker (BRI); Pinnacle Rock Track, 4.5 km W of Karnak, Jun 1992, Forster PIF10700, Sankowsky & Tucker (BRI); TR 66, Mt Lewis, Jun 1997, Forster PIF21239, Jensen & Tucker (BRI); Mt Lewis Road, Aug 1957, Smith 10094 (CANB); Davies Creek, SF 607, Aug/Sep 1971, Tracey 14745 (BRI); SFR 607, Emerald LA, Aug 1982, Gray 2701 (CNS); Mt Haig, Oct 2012, Cooper 2147 & Ford (CNS); Mt Haig, CSIRO plot E27, Oct 2012, Cooper 2148 & Ford (CNS); Lake Morris Road at start of track to tower 20, Sep 2012, Dowe 1159 (CNS); Copperlode Dam, Cairns, Oct 1970, Gittins 2196 (NSW); Palmerston Ridge Track, Wooroonooran NP, Feb 1995, Hunter 1109 (BRI); Breeden property, Hughes Road, Topaz, Aug 1997, Cooper 1152 & Cooper (CNS); Track to Phoenix Battery, Towalla, Jul 1997, Cooper 1132 & Cooper (CNS); Elinjaa LA, Oct 1974, Irvine 961 (CNS); Tully Falls NP, 2.2 km along Tully Falls road from Charmillan Creek bridge, Nov 2011, Ford5909 & Torello-Raventos (CNS); SFR 344, Douglas Creek, 0.7 km along Carron Creek Road from Kirrama Road, Jun 2002, Ford 3510 & Holmes (CNS). Distribution and habitat : Garcinia zichii is endemic to the Wet Tropics bioregion of north-east Queensland from Mt Finnigan to the Kirrama Range (Map 2) at altitudes between 150-1300 m. It occurs in rainforest (evergreen simple to complex notophyll/ 14 mesophyll vine forest) on soils derived from granite and rhyolite. Garcinia zichii grows often in association with Beilschmiedia oligandra L.S.Sm., Cryptocarya angulata C.T.White, Elaeocarpus elliffii B.Hyland & Coode, Flindersia pimenteliana F. Muell., Planchonella euphlebia (F.Muell.) W.D.Francis, Syzygium endophloium B.Hyland & S. wesa B.Hyland. Phenology : Flowers have been recorded in October and November; fruits have been recorded from June to November. Affinities : Within Australia, Garcinia zichii is most similar to G. jensenii. Both species have similarly shaped leaves and exudate canal pattern, a 2-locular ovary and 1 or 2-seeded fruit. G. zichii has leaves with 12- 18 pairs of secondary veins, flowers with green or yellow petals and fruit that are distinctly ellipsoid. G. jensenii has leaves with 7—10 pairs of secondary veins, flowers with white petals and fruit that are globose or ellipsoid-globose. G. zichii is also similar to G. hunsteinii Lauterb. from New Guinea. G. zichii has colourless, slightly milky or white latex, leaf apex bluntly acute and fruit much longer than wide, whereas G. hunsteinii has white or yellow latex, leaf apex acuminate and globose fruit. Notes : There is one specimen (Cooper 1152) as well as an observation by Tim Hawkes pers. comm., of fruit dehiscing, which was presumably caused by recent abundant rainfall. This phenomenon is well documented within horticulture for other plant groups, occurring after superfluous moisture is applied to orchard plantings. The abundant water causes osmotic absorption through the epicarp and the disintegration of the cuticle (Opara et al. 1997: 227). The exudate-containing canals within the leaves (a useful distinguishing character in Australian Garcinia) of this species are indistinguishable from G. jensenii (Fig. 2C). Etymology : The specific epithet honours Frank Zich (1968-), a most competent and Austrobaileya 9( 1): 1-29(2013) obliging curator of the collection at the Australian Tropical Herbarium (CNS). 6. Garcinia leggeae W.E.Cooper sp. nov. Similar to Garcinia riparia but differs by flower diameter (5 mm versus 15 mm), stamen number (5-9 versus 40-50), fruit shape (ovoid versus obovoid, oblate or patelliform), fruit size (20-27 mm long and 29-45 wide versus c. 70 mm long and c. 50 mm wide), sepal size on fruit (c. 1.5 mm long versus c. 5 mm in G. riparia). Typus: Australia: Queensland. Cook District: Gordon Creek, Iron Range National Park, 5 December 2012. W.Cooper 2187, R. Jensen & J.Kemp (holo: CNS [2 sheets + spirit], iso: 7 sheets to be distributed to BRI, CANB, K, KEP, L, MO, SING). Garcinia sp. (Claudie River LJB 19658) in part; Hyland et al. (2003). Garcinia sp. (Claudie River); Cooper & Cooper (2004: 123). Garcinia riparia auct. non A.C.Sm.; Cooper & Cooper (2004: 123). Garcinia sp. (Claudie River L.J.Brass 19658) in part; Jessup (2007: 54, 2010: 44). Garcinia sp. Claudie River (L.J.Brass 19658) in part; CHAH (2007); Hyland et al. (2010). Illustrations : Christophel & Hyland (1993: 81) as G. riparia vel aff.; Cooper & Cooper (1994: 146-147) as G. riparia vel. aff: y Hyland et al. (2003) as G. sp. (Claudie River LJB 19658); Cooper & Cooper (2004: 123) as G. sp. (Claudie River) & as G. riparia. Shrub or small tree to 6 m, dioecious; exudate yellow; dbh to 100 mm, often multi-stemmed; bark grey or greyish-brown, fissured and flakey, lenticels numerous; branchlets square, glabrous. Leaves discolorous, almost membranous, shiny on upperside and less so on underside, glabrous; petioles 5-8 mm long, channelled, not ligulate; lamina elliptical, 30-145 mm long, 25-57 mm wide, base cuneate; apex acuminate or caudate, rarely acute or obtuse; margin narrowly recurved; venation brochidodromous, primary vein flush on upperside and raised on underside; secondary veins 12-16 pairs, slightly raised on both surfaces, becoming puckered in dry Cooper, Garcinia in Australia specimens, angle of divergence from primary vein c. 45°, intramarginal vein 0.6-1.2 mm from margin; exudate-containing canals numerous, continuous parallel or intertwined lines running at angles between + parallel to primary vein and diverging up to c. 45° Male inflorescences axillary or ramiflorous, a fascicle seated on a pulvinus, 1-3-flowered; bracteoles persistent, triangular, c. 0.7 mm long and 0.8 mm wide, base truncated, apex acute, glabrous, margin entire. Male flowers: fragrant, diameter 5-6.5 mm, glabrous, sessile; sepals 4, 2-whorled, free, ovate, convex, entire, c. 1.5 mm long and 1 mm wide, membranous, green or yellowish, glabrous, margin thin; petals 4 (rarely 5), free, explanate, oblong-elliptic, sometimes slightly convex, 2.5-3 mm long, c. 1.5 mm wide, membranous, cream or yellow, margin entire or minutely ciliate; androecium a fleshy receptacle not adnate to petals, c. 0.5 mm long; anthers 5-9, sessile or almost sessile, erect or extrorse, quadrate, c. 0.4 mm long and 0.4 mm wide, bithecate; pistillode absent. Female inflorescences axillary or ramiflorus, a solitary flower, sessile; bracteoles persistent, triangular, convex, c. 0.8 mm long and 0.8 mm wide, base truncated, apex acute, glabrous, margin entire. Female fiowers: fragrant, diameter c. 5 mm, sessile; sepals 4, 2-whorled, free, ovate, convex, 2-3 mm long, 1.5-2 mm wide, membranous, green or yellowish, glabrous, margin thin, entire; petals 4, free, spreading widely, oblong- elliptic, sometimes slightly convex, c. 3 mm long and 1.5 mm wide, membranous, cream or yellow, glabrous, margin entire and thin; staminodal phalanges 4, adnate to petals, filament-like with a globose apex, distally inflexed, c. 1.4 mm long; antherodes 1 per phalange, c. 0.8 mm long; ovary urceolate, c. 2.5 mm long and 1.5 mm wide, sessile, 6-13-locular, uniovulate; stigma sessile, diameter c. 1.5 mm. Fruiting pedicel 0.7-1 mm long, sepals persistent at base c. 1.5 mm long, fruit a fleshy berry, ovoid (oblate, patelliform or asymmetrical when some ovules infertile), 20-27 mm long, 29-45 mm wide, pink-red to dark red, 6-13 septum lines, 6-13 segments; seeds 6-13, hairy, c. 11 mm long and 7-8.5 mm wide (not including hairs); 15 testa brown; aril fleshy, translucent whitish or pinkish. Figs. IE, 2F. Additional selected specimens (from 22 examined): Queensland. Cook District: Claudie River, Nov 1977, Hyland RFK3623 (CNS); ditto , Jan 1982, Hyland 11515 (CNS); ditto , Dec 1982, Hyland 12412 (CNS); ditto , Dec 1982, Hyland 12413 (CNS); West Claudie River, Iron Range, Aug 1997, Jensen 874 (CNS); Iron Range, Sep 1962, Volck AF02600 (CNS); Iron Range NP, CSIRO EP/42, Sep 2008, Ford 5422, Metcalfe, Murphy & Bradford (CNS); Claudie River EP/42, July 1978, Unwin 625 (CNS); Claudie River, Iron Range, May 1992, Cooper 279 & Cooper (CNS); Claudie River area, Oct 1968, Webb 8335 & Tracey (BRI); Claudie River, Iron Range NP, Dec 2012, Cooper 2168, Jensen & Kemp (CNS); Gordon Creek, Iron Range NP, Dec 2012, Cooper 2171 , 2175, 2177, 2186, 2188, Jensen & Kemp (CNS); ditto , Jun 2014, Cooper 2220 & Cooper (CNS); Gordon Creek, 10 km ENE of Mt Tozer, Iron Range NP, May 1992, Fell 2543 (BRI). Distribution and habitat : Garcinia leggeae is endemic to rainforest (evergreen mesophyll vine forest) in the Iron Range area, mostly on the Claudie River and Gordon Creek floodplains but occasionally on hillslopes to an altitude of 75 m (Map 3). Garcinia leggeae grows as an understory shrub or small tree on alluvial soils derived from basalt, frequently under a canopy of Blepharocarya involucrigera F.Muell., Calamus australis Mart., Garcinia dulcis , G. warrenii, Lasjia claudiensis C.L.Gross & B. Hyland, Myristica insipida R.Br., Nauclea orientalis (L.) L., Pandanus zea H.St.John, Syzygium bamagense B.Hyland & Tetrameles nudiflora R.Br. Phenology: Flowers have been recorded in June and December; fruits have been recorded in May, August and September. Affinities: Garcinia leggeae is similar to G. riparia from New Guinea. Compared with G. riparia, G. leggeae is a more diminutive plant, having much smaller flowers (5 mm versus 15 mm) with fewer stamens (5-9 versus 40-50), smaller fruit (20-27 mm long and 29-45 wide versus c. 70 mm long and c. 50 mm wide) and smaller sepals on ripe fruit ( c . 1.5 mm long versus c. 5 mm). Notes: Plants are frequently multi-stemmed, presumably caused by the forces of floodwater and floating debris during the wet season. 16 Some leaves on most herbarium collections, and all plants in the field seen by the author, are marked with black spots (diameter c. 5 mm), which appear to be caused by insects rather than fungi. Etymology : The specific epithet honours ornithologist and ecologist Sarah Legge (1969-) who collected this species as well as many other valuable specimens for illustration in the fruit book (Cooper & Cooper 2004), while working in the Iron Range area. 7. Garcinia mestonii F.M.Bailey, Report on New Plants, Preliminary to General Report on Botanical Results on Meston’s Expedition to the Bellenden-Ker Range 2 (1889), as ‘mestoni ’■ Type: Australia: Queensland. Cook District: Bellenden-Ker Range at an altitude of 2000 feet, [June 1889], [A.Meston & D.Whelan s.n .] (holo: BRI [AQ340247] 1 ). Illustrations : Christophel & Hyland (1993: 81); Cooper & Cooper (2004: 123), Hyland et al. (2010). Shrub or small tree to 6 m, dioecious (rarely monoecious); exudate yellow, meagre; bark nondescript with irregular-shaped lenticels; branchlets square when young, becoming terete. Leaves discolorous, coriaceous, glabrous; petioles 6-11 mm long, channelled, not ligulate; lamina elliptic, ovate, lanceolate, obovate, oblanceolate, 35-80 mm long, 12-34 mm wide; base attenuate, cuneate or oblique; apex caudate or acuminate; margin may be recurved in dry specimens; venation brochidodromous, primary vein + flush on upperside and raised on underside; secondary veins 10 pairs, angle c. 70° to primary vein, forming loops c. 1 mm from margin; exudate-containing canals numerous as short streaks, dashes and dots (more conspicuous on upperside), running at angles between 45° and parallel to primary vein or margin. Male inflorescences axillary, terminal or ramiflorous, a 2-5-flowered umbel or fascicle; bracts oblong, convex, 1.5-5.5 mm long, 0.5- 0.8 mm wide, glabrous, apex obtuse, base truncate, caducous; bracteoles triangular, c. 0.8 mm long and 0.6 mm wide, caducous; peduncles absent or up to 4 mm long. Male flowers: not fragrant, diameter c. 4 mm; Austrobaileya%\). 1-29(2013) pedicels 1.5-2.5 mm long; sepals 4,2-whorled, outer pair joined, D-shaped, cupular, c. 4 mm long and 4.5 mm wide, green, glabrous, margin thin; petals 4, orbicular, cupular with apex recurved, c. 4.8 mm long and 4.8 mm wide, membranous, green or cream, glabrous, margin thin; androecium a fleshy receptacle not adnate to petals, c. 2.5 mm long and 3.2 mm wide; anthers c. 16, triangular, sessile or almost sessile, 0.6-1 mm long, 0.6-1 mm wide at base, bithecate; pistillode absent. Female inflorescences axillary or terminal, solitary flowers or 2-3-flowered umbels or fascicles; bracts 4-whorled, oblong, 1.5-3.2 mm long, c. 1 mm wide, caducous; bracteoles triangular, c. 1 mm long and 0.6 mm wide; peduncle c. 1 mm long. Female flowers: not fragrant, pedicels c. 2.5 mm long, diameter 6-7.5 mm; sepals 4, 2-whorled, outer pair joined, inner pair free, D-shaped, cupular, 3.8-4.4 mm long, 4.3-5.5 mm wide, membranous, green or cream, glabrous; petals 4, free, orbicular, cupular with apex recurved, c. 5 mm long and 4 mm wide; staminodal phalanges adnate to petals, narrowly or broadly triangular, apically 2 or 3-branched, distally indexed, c. 2 mm long, antherodes 2 per branch; ovary oblate, 2.2-2.8 mm long, 4-4.5 mm wide, sessile, 3-9-locular, uniovulate; style c. 0.5 mm long, stigma diameter c. 2.5 mm, 3-9-rayed. Fruiting pedicel 1-3 mm long; 4 sepals persistent at base; fruit a fleshy berry, depressed globular, 31-50 mm long, 40-60 mm wide, cream or greenish-cream; stigma 3-9-rayed, diameter 3.5-4 mm; seeds 13-25 mm long, c.10 mm wide, testa brown; aril fleshy, cream. Meston’s mangosteen. Fig. 1C & F, 2D. Additional selected specimens (from 26 examined): Australia. Queensland. Cook District: Little Cooper Creek, Cow Bay, Cape Tribulation area, Oct 1994, Small s.n. (CNS [QRS104526]); Behana Creek, S of Gordonvale, Jul 2005, Halford Q8446 & Jensen (BRI); Divide between Babinda Creek and East Mulgrave, May 2002, Jago 6211 (CNS); Mt Harold, Aug 1985, Irvine 2322 , 2323 (BRI, CNS); Goldsborough Road, Mulgrave River Valley, Jul 1992, Tucker s.n. (CNS [QRS105577]); Goldsborough Road, Jul 1992, Sankowsky 1332 (CNS); East Mulgrave River, Nov 1995, Jensen 483 (CNS); Ridgeline NW of airplane wreck, Bellenden Ker, Jun 1995, Hunter 5308 (BRI); Mt Bellenden Ker, c. 7/8 mile SE of centre peak, Jun 1969, Smith 14691 (BRI); Harvey Creek, N of Bellenden Ker, Dec 1998, Forster PIF24028 (CNS); Harvey Creek, Dec 2012, Cooper 2203, 2204 & Cooper, Garcinia in Australia Jensen (CNS); Cultivated at Tolga, ex Harvey Creek, Dec 2003, Sankowsky 2326 & Sankowsky (BRI); Sankowsky’s garden, Tolga [ex Harvey Creek], Jan 2013, Cooper 2209, Sankowsky & Jensen (CNS); NPR 226, Parish of Bellenden Ker, Dec 1983, Hyland 25062 RFK (BRI, CNS); Ridgetop S of Tower 6, Bellenden Ker, Jun 1995, Hunter 5292 (BRI); E slopes of Bellenden Ker near Station 3 on the Bellenden Ker Cable Car Track, Jan 1981, Irvine 2056 (BRI, CNS); Bellenden Ker, s.dat., Meston 1 (BRI [AQ166293]); Bellenden Ker Range, Mar 1922, White s.n. (BRI [AQ166291]); Bellenden Ker, below summit, Jun 1949, Smith 4236 (BRI); Wooroonooran NP, headwaters of East Mulgrave River, 5 km W of Bellenden Ker township, Nov 2000, Forster PIF26412, Booth & Jensen (BRI). Distribution and habitat: Garcinia mestonii is endemic to the Wet Tropics bioregion of north-east Queensland between Cape Tribulation and the Mt Bellenden Ker Range (Map 3). It occurs in rainforest (evergreen, simple/complex notophyll/mesophyll vine forest) on granitic and metamorphic soils at altitudes between 30 and 1340 m. Phenology: Flowers have been recorded in December and January; fruit has been recorded from August to February. Typification: The type collection at BRI has minimal original labelling attached to the sheet, apart from a tattered scrap of paper with the number 18 and the words ‘Meston’s Apple’. No specific collectors or date of collection are given; although a copy of the protologue is attached. In the protologue Bailey (1889) noted that it was collected by Messrs. Meston and Whelan on the first ascent of Bellenden Ker. As noted by Dowe & Broughton (2007) this was in June 1889. Archibald Meston is well known as the leader and organiser on the 1899 and 1904 Bellenden Ker expeditions, but was also a significant collector of plant specimens, particularly on the latter expedition when F.M.Bailey was absent (Dowe & Broughton 2007, 2008). The co-collector of the type of Garcinia mestonii was Daniel Whelan, a Sergeant in the infamous [Queensland] Native Mounted Police (Richards 2008) and who led the force of troopers allocated as protection on the 1889 expedition. Whelan was honoured by Bailey with the Proteaceous tree Lasjia whelanii (originally as Helicia whelanii F.M.Bailey) and the fern Blechnum whelanii 17 F.M.Bailey. The ‘Whelanian Pools Camp’ on Bellenden Ker was also named for him (Dowe & Broughton 2007). Notes: Garcinia mestonii has been recorded as dioecious in the field; however, one plant cultivated by G & N Sankowsky at Tolga is distinctly monoecious with about 50% of flowers being male and 50% female. Individual branches on this tree appear to bear flowers of one sex only. Etymology: The specific epithet honours Archibald Meston (1851-1924), a botanical collector, journalist, civil servant, explorer and member for Rosewood in the Queensland Legislative Assembly from 1878-1882. 8. * Garcinia mangostana L ., Sp. PI. 443 (1753). Type: ‘Mangostan’ in L.Garcin, Philos. Trans. 38: t. 1 (1734); fide Hammel (1993: 28). Illustrations: Fleming ( c. 1795-1805); Hooker (1855: t. 4847); van Nooten (1863); Marsden (2005: plate 3); Nazre (2006: 41, 45, 49, 52, 54, 57, 63, 181); Te-chato (2007: 248, 249, 250). Tree to 20 m, agamospermous; exudate yellow; bark fissured and flaky; branchlets ± terete with some ribbing, glabrous. Leaves discolorous, coriaceous, glabrous, drying reddish-brown; petioles 10-30 mm long, not channelled on fresh specimens and slightly channelled on dried specimens, ligulate; lamina elliptic, broadly elliptic, ovate or oblanceolate, 100-250 mm long, 45-105 mm wide, base cuneate or obtuse; apex acute, shortly acuminate or obtuse; margin narrowly recurved; venation brochidodromous, primary vein distinctly raised on both sides; secondary veins 15-17 pairs, prominent, angle of divergence 50-70° from primary vein; 2 intramarginal veins, outermost 0.5-2 mm from margin, innermost 3-7 mm from margin; exudate-containing canals on underside as numerous branched lines running at angles between parallel to primary vein and margin, on upperside as numerous branched lines mostly running parallel to secondary venation. Male flowers unknown. Female inflorescences terminal, a solitary flower, pairs or rarely in 3’s. Female flowers: 18 diameter 25-60 mm; pedicels 1.5-20 mm long; sepals, 4, 2-whorled, c. 20 m long, green, cream, yellowish, pinkish or reddish, cupular, coriaceous; petals 4, free, erect to spreading, broadly obovate or broadly orbicular, c. 25 mm long, cream, pink or reddish, glabrous, thickly fleshy; staminodes up to 20 shorter than ovary, adnate to petals, free, filament-like, c. 5 mm long, caducous; antherodes 1 per filament; ovary broadly ovoid or ellipsoid or globose, 12-16 mm, sessile, 4-8-locular, locules uniovulate; stigma mass sessile, domed, diameter 8-12 mm, 4-8-lobed. Fruiting pedicel with 4 green to reddish sepals persistent at base; fruit a dry berry with a thick corky-leathery pericarp c. 9 mm wide, globose or depressed globose, diameter 35-70 mm, purple to blackish, glabrous; sepals persistent; stigma persistent, 4-8-rayed; seeds 4-8, 15-25 mm, testa brown; aril thickly fleshy and white. mangosteen, Queen of fruit. Additional selected specimens (from 5 examined): Queensland. Cook District: Cultivated, Stanton property. Intake Road, Redlynch, Cairns, Jul 2013, Cooper 2222 (CNS); North Johnstone River, off the Cooroo Lands Road, W of Innisfail, Jul 2007, Bean 26749 (BRI); Dunk Island, Aug 1963, Scanlan AFO 2812 (CNS); Dunk Island, Aug 1964, Scanlan AFO 3490 (CNS); Tully, Sep 1941, Shearer s.n. (CNS). Distribution and habitat: Garcinia mangostana is unknown in the wild but is likely to have originated on Peninsula Malaysia (Richards 1990b). In Australia it is cultivated in the tropical lowlands of Queensland, and a recent collection ( Bean 26749), in the Innisfail area, confirms naturalisation within Queensland rainforest. Three old collections from Dunk Island and Tully also indicate possible naturalisation (Map 3). Phenology : Flowers and fruit have not been collected from naturalised plants; however, Peter Stanton and Allen Sheather have cultivated plants in the Cairns and Daintree areas and record flowers from September to March; fruit December to July. Notes: Garcinia mangostana is thought to be a hybrid of multiple origins (Richards 1990b). Within tropical Asia, Mangosteens have been widely cultivated over hundreds of years for Austrobaileya 9(1): 1-29(2013) their delicious arils and are referred to as the Queen of Fruit. Several authors including Richards (1990a), suggest that male flowers are probably always absent and the species is an obligate agamosperm, forming fruit without fertilisation. Etymology: The specific epithet is derived from Latinising the Malay name manggis. 9. Garcinia brassii C.T.White, Proc. Roy. Soc. Queensland 47: 52 (1936). Type: Australia: Queensland. Cook District: Thornton Peak, in low scrub at the summit, 14 March 1932, L.J. Brass 2277 (holo: BRI). Illustrations: Christophel & Hyland (1993: 81); Cooper & Cooper (2004: 123); Hyland et al. (2010). Tree to 18 m, dioecious; exudate yellow; bark with fine vertical fissures; branchlets square becoming terete, new twigs mostly red, glabrous. Leaves discolorous, coriaceous, glabrous; petioles 5-12 mm long, shallowly channelled on younger growth and on dried specimens, ligulate; lamina obovate or elliptical, 25-125 mm long, 16-63 mm wide, base attenuate or cuneate; apex acute, obtuse or emarginate; margin recurved; venation brochidodromous, primary vein ± flush on upperside but may be raised near base on dry specimens; secondary veins raised on both sides, 12-35 pairs, angle of divergence from primary vein 65-80°, forming loops 0.3-1.2 mm (rarely 4 mm) from margin, intramarginal vein often hidden in the recurved margin; some tertiary veins or veins forking off secondary veins may be present, if so, most diminish and do not reach the intramarginal vein; rarely are reticulations of similar strength to secondary veins present and if they do they are few and near the margin; exudate-containing canals on underside as numerous parallel lines, branched, straight, wriggly or curled, running at angles radiating between primary vein and margin. Male inflorescences terminal, a panicle, 3-18-flowered, 19-28 mm long; peduncles 10-21 mm long; bracts obovate or elliptical, keeled, 5-7 mm long, 2-3 mm wide (these may expand into small leaves up to 55 mm long and 21 mm wide), caducous; bracteoles 2-2.5 mm long and c. 2 mm wide. Cooper, Garcinia in Australia Male flowers: fragrant, diameter 15-28 mm; pedicels absent or up to 6 mm long; sepals 4, 2-whorled, dark red, glabrous, margin thin; outer pair joined, D-shaped or ovate- orbicular, convex, 4-6.5 mm long, 4-6.5 mm wide, coriaceous; inner pair free, orbicular or oblong-orbicular, convex, 7-8 mm long, 6-7 mm wide; petals 4, free, erect or spreading, imbricate, orbicular, convex, 9-14 mm long, 8-14 mm wide, waxy-coriaceous, cream, yellow, pink or reddish, unmarked or with dark red markings, glabrous, margin thin; androecium 4-phalangiate, glabrous; phalanges adnate to petals, obovate or elliptical at anthesis, 5-8 mm long, 5-6 mm wide; anthers several to numerous per phalange, sparsely to densely clustered over phalange, bilobed, c. 0.5 mm long and 0.5 mm wide; pistillode rudimentary, attenuate-conical, 1- 3.5 mm long. Female inflorescences terminal, a solitary flower 10.5-12 mm long; peduncle 1.5-3 mm long; bracts at apex of peduncle triangular, keeled, 3-7 mm long, 2- 3 mm wide, caducous; bracteoles 1.5-2 mm long and 2 mm wide. Female flowers: fragrant, diameter 13-18 mm; pedicels absent or up to 1.5-3.5 mm long; sepals 4,2-whorled, coriaceous, green and mostly blushed dark- red towards apex, glabrous, margin thin; outer pair joined, oblong, convex, c. 5 mm long and 4 mm wide; inner pair free, broadly ovate or orbicular, convex, 6-8 mm long, 5-7 mm wide; petals 4, free, erect to spreading, orbicular, convex, 8-10 mm long, 8—10 mm wide, waxy-coriaceous, cream or yellowish, often with red markings and often ageing to entirely reddish, glabrous, margin thin; staminodal phalanges 4, erect, adnate to petals, strap-shaped with flabellate apex, c. 3.5 mm long and 4 mm wide, apically branched into 5-8 individual filaments c. 1 mm long; antherodes 1 or 2 per filament, 0.5- 0.7 mm long; gynoecium globose, 6-7.5 mm long, 4-5 mm wide, glabrous; ovary ovoid, 5-6 mm long, 4-5 mm wide, sessile, 4-locular, locules uniovulate; stigma mass sessile, domed, diameter 4.5-5.5 mm, cream becoming reddish with age or exposure. Fruiting pedicels 2-3.5 mm long, 4 sepals persistent at base, fruit a fleshy berry, depressed globular, 22-45 mm long, 27-50 mm wide, dark red 19 to blackish, glabrous; stigma shallowly 3 or 4-lobed, diameter 5.5-6.5 mm; seeds 1-4, 12-22 mm long, 10-18 mm wide; testa brown, smooth; aril fleshy, dark red. Fig. 2A, 3A-C. Additional selected specimens (from 36 examined): Queensland. Cook District: Near peak of walking track to Mt Sorrow, 3 km W of Cape Tribulation, Daintree NP, Sep 2009, Worboys 825 (CNS); NPR 133, Daintree, summit of Mt Sorrow track, Dec 1998, Ford 2147 (CNS); VCL Noah, Mt Hemmant, Oct 1975, Hyland 8490 (CNS); Thornton Peak, Jul 2008, Jago 7152, Ford & Keith (BRI); Thornton Peak, Sep 1937, Brass & White 280 (BRI, K 1 ); Thornton Peak, in valley between N and S peaks near campsite on Hilda Creek, Sep 1984, Clarkson 5588 (CNS); Pinnacle Rock Track, 4.5 km W of Karnak, Jun 1992, Forster PIF10708, Sankowsky & Tucker (CNS); Along track to Devils Thumb, Sep 1997, Jago 4496 (BRI); Mt Misery, E of Mt Spurgeon, 15.7 km NE of Mt Carbine, Nov 1988, Jessup GJM861, Guymer & McDonald (BRI); Mossman Bluff c. 10 km W of Mossman, Dec 1998, Fell 1683 (BRI); Near Schiller’s hut, Mt Spurgeon, Sep 1972, Webb 13438 & Tracey (CNS & BRI); 27.3 km from Rex Highway on Mt Lewis Road, Nov 1990, Holland 31 & Hind (NSW); Mt Lewis Road S[South] Mary LA, Nov 1988, Jessup GJM1526, Guymer & McDonald (BBS), Mt Lewis FR, 28 km along Mt Lewis Road, Jan 2002, Forster PIF28164, Booth & Jensen (BRI); Mt Lewis, on track to Rhododendrons, Sep 2012, Cooper 2163 & Ford (CNS); ditto , Dec 2012, Cooper 2207 & Ford (CNS); Mt Lewis, CSIRO plot EP18, Sep 2012, Cooper 2162, 2164 & Ford (CHS). Distribution and habitat : Garcinia brassii is endemic to the Wet Tropics bioregion of north-east Queensland from Mt Sorrow to Mossman Bluff and west to the Mt Lewis -Mt Spurgeon areas (Map 1), at altitudes between 500-1300 m. It occurs in rainforest (simple notophyll vine forest, complex notophyll vine forest and simple microphyll vine-fern thicket) on granitic soils. Phenology : Flowers have been recorded in June, August to November; fruit has been recorded in December and January. Etymology : The specific epithet is in honour of botanist Leonard J. Brass (1900-1971), collector of the type specimen. 10. Garcinia gibbsiae S.Moore, J. Bot. 55: 302 (1917). Type: Australia: Queensland. Cook District: Bellenden Ker, forest on long ridge, March 1914, L.S.Gibbs 6306 (holo: BM 1 ; iso: K 1 ). 20 Austrobaileya 9(1): 1-29 (2013) Fig. 3. Flowers. A. Garcinia brassii showing stigma and apically branched staminodal phalanges around the ovary (i Cooper 2163 & Ford [CNS]). B. G. brassii $, showing 4-phalangiate androecium prior to anthesis ( Cooper 2164 & Ford [CNS]). C. G. brassii S, showing 4-phalangiate androecium post anthesis ( Cooper 2164 & Ford [CNS]). D. G. gibbsiae showing stigma and apically branched staminodal phalanges around ovary {Cooper 2213 & Cooper [CNS]). E. G. gibbsiae S, showing androecium of 4 fused phalanges {Cooper 2212 & Cooper [CNS]). F. G. warrenii & showing anthers on 4-phalangiate androecium post anthesis {Cooper 2158 & Hawkes [CNS]). G. G. russellii showing stigma and apically branched staminodal phalanges around ovary {Cooper 2150, Russell & Hawkes [CNS]). H. G. russellii 3 , showing androecium of 4 fused phalanges {Cooper 2149, Russell & Hawkes [CNS]). I. G. warrenii showing stigma and apically branched staminodal phalanges around the ovary {Cooper 2159 & Hawkes [CNS]). Photos: A, D, E, W.Cooper, B, C, F, G, H, I, T.Hawkes. Cooper, Garcinia in Australia Illustrations : Christophel & Hyland (1993: 81); Cooper & Cooper (1994: 288); Cooper & Cooper (2004: 123); Hyland et al. (2010). Tree to 12 m, dioecious; exudate bright lime-yellow; bark with fine vertical fissures and irregular lenticels; branchlets shallowly ribbed, horizontal, erect or pendulous, glabrous. Leaves discolorous, coriaceous, glabrous; petioles 8-20 mm long, channelled, ligulate; lamina oblong-elliptic, ovate, oblanceolate, elliptical or elliptic-obovate, 65- 200 mm long, 30-93 mm wide, base cuneate or obtuse, apex acute or obtuse, margin thickened and sometimes remotely recurved; venation paxillate, primary vein raised on both sides; secondary veins numerous (more than 35), + flush, parallel, angle of divergence with primary vein 70-90°, intramarginal vein on the margin; exudate-containing canals similar on both sides as sparse lines, branched or unbranched, mostly running at a slightly narrower angle than the secondary veins and sometimes crossing over them. Male inflorescences terminal or rarely axillary, a solitary flower, pairs, or a 2-8-flowered raceme, up to 38 mm long; peduncle 3-22 mm long; bracts narrowly triangular, convex, 5-10 mm long, 3-4 mm wide, glabrous; bracteoles triangular, convex, c. 2 mm long, glabrous. Male flowers: not fragrant, diameter 17-46 mm; pedicel absent or up to 2.7 mm long; sepals 4, 2-whorled, convex, may be slightly keeled, green, margin thin; outer pair joined, D-shaped, 5-8 mm long, 7-9 mm wide; inner pair free, orbicular, 5.5-7 mm long, 8-10.5 mm wide; petals 4 (rarely 3), free, orbicular, convex or flat, 11-20 mm long, 11-20 mm wide, thick, waxy, yellow (sometimes blushed with pink), glabrous, margins thin and mostly incurved; androecium of 4 fused phalanges creating a solitary receptacle, rhombic, adnate to petals, divided into 4 shallow lobes at apex, diameter 10-13 mm; anthers numerous, densely clustered over receptacle, subsessile, strap-shaped or wedge-shaped, bithecate, 0.9-1.3 mm long, 0.5-1 mm wide, glabrous; pistillode absent. Female inflorescences terminal, solitary or 2-flowered; peduncles 1-2.5 mm long; bracts triangular, keeled, 3-5 mm long, 1.5-2 mm wide, caducous; bracteoles triangular, orbicular or ovate, 21 slightly keeled, c. 3 mm long and 2 mm wide, caducous. Female flowers: faintly fragrant, diameter 15-22 mm, pedicels 1.5- 2.5 mm long; sepals 4, convex, green, glabrous, margin entire and membranous; outer pair narrowly joined, D-shaped, 5-6.5 mm long, 8-10 mm wide; inner pair free, orbicular, 6.5-8 mm long, 7.5-12 mm wide; petals 4, free, orbicular, convex, 14-17 mm long, 11-15 mm wide, waxy-coriaceous, yellow, glabrous, margin thin; staminodal phalanges 4, free or adjacent pairs fused, adnate to petals, strap-shaped, erect, 2.5-5 mm long, 3-3.5 mm wide, apically branched into 6-9 individual filaments c. 1 mm long; antherodes 1 or 2 per filament, erect or introrse, oblate, c. 1 mm long and 0.6 mm wide; gynoecium urn-shaped, 7-9-locular, surface irregularly lobed, glabrous; ovary globular, 4-5 mm long, 5-6 mm wide, sessile, 7-9-locular, 1-4 ovules per locule; stigma sessile, domed, 7-9-rayed, diameter 5-6 mm. Fruiting pedicel 2-4 mm long, 4 sepals persistent at base; fruit a semi-dry berry, top-shaped, broadly-ovoid or truncated-globose, base truncated, apex acute, beaked or rarely truncated, 30-62 mm long, 30-67 mm wide, green, septal radii conspicuous as 7-9 shallow ribs or lines; stigma persistent at apex, 7-9-rayed; locules 7-9, septa thick, coriaceous and entire; seeds 1-4 per locule, ± globular, 10-18 mm long, 7-11 mm wide, testa brown, aril absent. mountain mangosteen. Figs. 2E, 3D & E. Additional selected specimens (from 29 examined): Australia. Queensland. Cook District: Mt Bellenden Ker, east slopes, April 1948, Brass 18330 (BRI); Mt Bellenden Ker, c. 7/8 mile SE of the centre peak, June 1969, Smith 14690 (BRI); Near junction of E and W Mulgrave River in SFR 310, Upper Goldsborough LA, 19.1 km SSE Little Mulgrave town, Nov 1988, Jessup GJM1746, Guymer & McDonald (BRI); Old Mailman’s Track, W of Windin Creek, Feb 1995, Hunter 849 (BRI); SFR 755, Gosschalk LA, EP/34, Nov 1976, Unwin 117 (CNS); SFR 310, Windin LA, Apr 1973, Hyland 6702 (CNS); Lamonds [Lamins] Hill, Atherton Tableland, Oct 1949, Stephens NQ Nat. Club No. 13312 (BRI); SFR 310, Swipers LA, Oct 1963, Hyland 278RFK (CNS); Headwaters of Jimpee Creek, s.dat ., Hunter 4522 (CNS); TR 1230, Boonjee LA, Oct 1979, Moriarty 2678 (CNS); ditto , May 1982, Gray 2550 (CNS); TR 1230, Boonjee LA, Boonjee, walking track to Stockwellia site, June 1992, Tucker & Sankowsky s.n. (BRI [AQ546523]); Wooroonooran NP, track to Chuck Lunga Creek, E of Boonjee, Forster PIF19286 & Tucker (BRI, CNS); Mt Bartle Frere, 1.8 km WSW Bobbin Bobbin Falls, 4.4 km 22 NNE Boonjee, Nov 1988, Jessup GJM1049, Guymer & McDonald (BRI); Boonjie LA near Bartle Frere track, 0.9 km SE of Bobbin Bobbin Falls, Nov 1988, Jessup GJM269, Guymer & McDonald (BRI); Westcott Road, Topaz, Jul 1992, Cooper 320 & Cooper 320 (CNS); ditto , Mar 2013, Cooper 2212, 2213 & Cooper (CNS); Topaz, Dec 1970, Kitchener s.n. (CNS); Downey Creek SA 34, 24.6 km SE of Millaa Milla, Oct 1988, Jessup GJM2217, Guymer & McDonald (BRI). Distribution and habitat : Garcinia gibbsiae is endemic to the Wet Tropics bioregion of north-east Queensland where it occurs in rainforest (complex mesophyll vine forest and simple notophyll vine forest) on basalt soil from the eastern slopes of the Mt Bellenden Ker Range to Downey Creek near Innisfail and west to the Boonjie-Topaz area (Map 4), at altitudes between 120 and 1160 m. Phenology : Flowers have been recorded from February to June; fruit has been recorded from July to January. Affinities : Garcinia gibbsiae appears to be most similar to G. russellii. Both species have tough leathery leaves with numerous parallel secondary veins and green fruit with up to three or four seeds in each locule. G. gibbsiae is a moderately to densely branched shrub or small tree, with bright lime-yellow exudate, leaf bases which are either cuneate or obtuse and never peltate or cordate, male inflorescences solitary or a few-branched raceme and petals yellow, fruit with spacious locules not completely filled by seeds; locular septa thick, fleshy and entire. G. russellii is a slender and sparsely branched shrub or small slender tree with colourless, milky or white exudate, peltate leaves usually with cordate bases, male inflorescences a 9- or more- flowered panicle and petals cream or white, fruit with seeds tightly packed within each locule which mostly cause the exocarp to bulge; locular septa thin, pithy and incomplete. Etymology : The specific epithet honours British botanist Lilian Suzette Gibbs (1870— 1925). 11. Garcinia russellii W.E.Cooper sp. nov. Similar to Garcinia gibbsiae but differs in the exudate colour (clear or slightly milky versus bright lime-yellow), leaves (peltate versus not peltate), male inflorescences (nine or more Austrobaileya%\). 1-29(2013) flowers versus up to 8-flowered but usually less); fruit septa (thin, pithy and incomplete versus thick, fleshy and entire). Typus: Australia: Queensland. Cook District: North of Mossman River, Daintree National Park, 9 July 2011, W.Cooper 2137, R.Russell, R. Jensen & R.Jago (holo: CNS [1 sheet + spirit]; iso: 3 sheets to be distributed to BRI, CANB, MO). Garcinia sp. (Mossman); Cooper & Cooper (2004: 124). Illustrations : Cooper & Cooper (2004: 124) as Garcinia sp. (Mossman). Shrub or small tree to 6 m, sparsely branched with branches mostly pendulous, dioecious; exudate opaque white, colourless or slightly milky; bark with a few horizontal raised rings at old interpetiolar scars, dotted with small pustules; branchlets slightly 4-sided on new growth and becoming terete on older growth, glabrous. Leaves discolorous, coriaceous, glabrous, peltate; petioles 10-25 mm long, attached on underside of lamina; lamina oblong or ovate-oblong, 110-295 mm long, 45-100 mm wide, base cordate or rounded, apex acute and often almost aristulate, margin entire and strongly recurved; venation paxillate, primary vein distinctly raised on both sides, green on upperside and yellow on underside; secondary veins c. 100, raised on both sides, angle of divergence from primary vein 70-90°, intramarginal vein on the margin; exudate-containing canals sparse on both sides as unbranched or branched lines, mostly running at a slightly narrower angle than the secondary veins and sometimes crossing over them. Male inflorescences terminal, a panicle, 9-39-flowered, 30-62 mm long; peduncle 5-11 mm long; bracts narrowly triangular, convex, 8-9 mm long, c. 2.5 mm wide, glabrous; bracteoles triangular, convex, 2-3 mm long, 1.5-2 mm wide. Male flowers: not fragrant, diameter 13-27.5 mm, pedicels 2.5-5 mm long; sepals 4, 2-whorled, orbicular, convex, may be slightly keeled; outer pair joined, with margin entire or irregularly lobed, 4-6 mm long, 7-8 m wide, green, glabrous; inner pair free, with margin recurved, entire or irregularly lobed, 4-8 mm long, 4-10 mm wide, pale green, glabrous; petals 4, free, orbicular or oblong-orbicular, Cooper, Garcinia in Australia convex, 8.5-15 mm long, 7.5-15 mm wide, white or cream on the inner surface and cream or pale yellow on the outer surface, glabrous; androecium of 4 fused phalanges creating a solitary receptacle, rhombic, adnate to petals and flower base, undivided or divided into 4 shallow lobes at apex, diameter 5.5-11 mm; anthers numerous, densely clustered, subsessile, strap-shaped or broadly wedge- shaped, bithecate, c. 2 mm long and 0.5 mm wide, glabrous; fungiform pistillode present in some flowers, c. 4.5 mm long, diameter at apex c. 2 mm. Female inflorescences terminal, a solitary flower (rarely pairs); bracteoles narrowly triangular, convex, c. 8.5 mm long and 2.5 mm wide at base, glabrous, caducous. Females flowers: not fragrant, diameter 16-25 mm; pedicel c. 2 mm long; sepals 4, free, D-shaped, base truncate, apex obtuse, cream; outer pair narrowly joined, c. 6.3 mm long and 7.2 mm wide; inner pair free, c. 8.3 mm long and 10 mm wide; petals 4, oblong-orbicular, convex, c. 13.5 mm long and 10.5 mm wide, cream; staminodal phalanges 4, adnate to petals, somewhat square, apex rounded, antherodes 5-8 per phalange, c. 1 mm diameter; gynoecium urn-shaped, c. 6 mm long; ovary ovoid or top-shaped, 6-8-locular; stigma domed, 6-8-rayed, diameter c. 3.5 mm; ovules 1-4 per locule. Fruiting pedicels c. 2 mm long; 4 sepals persistent at base; fruit a semi-dry berry, top-shaped or broadly ovoid, mostly beaked, beak up to 20 mm long, 6-8 shallow septum-lobes or septum-lines, 60-72 mm long, 50-58 mm wide, green, dull (shiny when unripe), stigma remnants 6-8-rayed, diameter 5-6 mm; locules 6-8; septa thin, transparent, pithy and incomplete; seeds 1-3 per locule, wedge-shaped, 20-22.5 mm long, 18-23 mm wide; testa cream to brown; aril cream and netted, not fleshy. Fig. 3G & H, 4. Additional selected specimens (from 12 examined): Queensland. Cook District: Finlayvale, NW of Mossman, May 2012, Hawkes s.n. (CNS); Mossman Gorge NP on circuit track, Dec 1989, Jessup GJD3330, Guymer & Dillewaard (BRI); above the north bank of the Mossman River west of Mossman, Jan 2003, Jago 6382, Russell & Darwin (BRI); N of Mossman River in Daintree NP, Dec 2002, Russell 11 (CNS); North of Mossman River, Daintree NP, Jul 2011, Cooper 2138, Russell, Jensen & Jago (CNS); ditto , May 2012, Cooper 2149, 2150, 2151, Russell & Hawkes (CNS); ditto , Nov 23 2012, Cooper 2167, Russell, Ford & Sheather (CNS); ditto , Dec 2012, Cooper 2205, 2206, Russell & Sheather (CNS). Distribution and habitat : Garcinia russellii is endemic to the Wet Tropics bioregion of north-east Queensland where it is currently known to occur in rainforest (evergreen mesophyll vine forest) on granitic soils in the Mossman Gorge section of Daintree NP and at Finlayvale near Mossman (Map 4) at altitudes between 150 and 300 m. At the type locality Garcinia russellii grows as a slender understory tree in rainforest under a canopy of Backhousia bancroftii F.M.Bailey & F.Muell. ex F. M.Bailey, Flindersia bourjotiana F.Muell., Linds ay omyrtus racemoides (Greves) Craven, Hicksbeachia pilosa P.H.Weston & Calamus caryotoides A.Cunn. ex Mart. Phenology : Flowers have been recorded in May and July; fruits have been recorded in December. Notes : Male flowers within the same inflorescence can be inconsistent with some flowers having a fungiform pistillode and others not having any indication of a pistillode. The exudate-containing canals within the leaves of this species are indistinguishable from G. gibbsiae (Fig. 2E). Affinities : Garcinia russellii appears to be most similar to G. gibbsiae. Both species have tough coriaceous leaves with numerous parallel secondary veins and green fruit with up to three or four seeds within each locule. G. russellii is a slender and sparsely branched small slender tree with colourless, milky or white exudate, peltate or almost peltate leaves with distinctly cordate bases, male inflorescences are a 9- or more-flowered panicle and petals cream or white, fruit with seeds tightly packed within each locule which mostly cause the exocarp to bulge; locular septa thin, pithy and incomplete. Whereas, G. gibbsiae is a moderately to densely branched small to medium-sized tree, bright lime-yellow exudate, leaf bases which are either cuneate or obtuse and never peltate or cordate, male inflorescences solitary or a few-branched raceme and petals yellow; fruit 24 Austrobaileya 9(1): 1-29(2013) Fig. 4. Garcinia russellii fruit A. showing longitudinal cross-section and multi-carpelled locules, thin transparent septa and netted aril. B. showing beaked fruit, bracts and terminal attachment to branch (Cooper 2206, Russell & Sheather [CNS]). Photos: A, W.Cooper, B, T.Hawkes. with seeds usually in spacious locules; locular septa thick, fleshy and entire. Etymology : The specific epithet is in honour of naturalist and conservationist Rupert Russell (1939-). 12. Garcinia warrenii F.Muell., Viet. Nat. 8: 109 (1891). Type: Australia: Queensland. Cook District: Coen River, in 1891, S. Johnson s.n. (holo: MEL n.v.\ iso: BRI [AQ0340253 1 ], K 000677819 1 ). Garcinia kajewskii C.T.White, Contr. Arnold Arbor. 4: 72 (1933). Type: Queensland. Cook District: Daintree River, 30 November 1929, S.F.Kajewski 1418 (holo: BRI [AQ0340243 1 ]; iso: S 11-34563 1 ). Illustrations: Christophel & Hyland (1993: 82); Cooper & Cooper (1994:288); Hyland et al. (2003); Cooper & Cooper (2004: 123). Tree to 25 m, dioecious; exudate yellow; dbh to 35 cm; bark tessellated and flakey in large trees, vertically fissured and with vertical lenticels; branchlets square orterete andmostly horizontal, glabrous. Leaves discolorous, coriaceous, glabrous; petioles 8-30 mm long, not channelled or becoming shallowly channelled in dry specimens, ligulate; lamina elliptic, oblong-elliptic, ovate, obovate, orbicular or oblanceolate, 40-230 mm long, 28-123 mm wide; base attenuate, cuneate or obtuse; apex acuminate, acute, obtuse, retuse or emarginate; margin not recurved or rarely slightly recurved; venation brochidodromous, often with a second set of loops near margin, primary vein on upperside raised towards base and more or less flush towards apex; secondary veins 24-50, angle of divergence from primary vein 60-90°, forming an intramarginal vein 0.5-4 mm from margin; exudate-containing canals difficult to see on fresh leaves, distinct on underside of dried leaves as numerous unbranched or branched straight or wriggly + parallel lines, sometimes a second set of canals is visible as more widely spaced long straight lines running at angles 45-80° to primary vein and across the other canal set. Male inflorescences terminal or rarely axillary, a panicle, 8-66-flowered, 15-120 mm long; peduncle 5-38 mm long; bracts (which sometimes become full- sized leaves) at peduncle apex, triangular, becoming incurved, glabrous, 8.5-10 mm long, 2-3 mm wide, green with a yellowish primary vein, caducous; bracteoles triangular, 25 Cooper, Garcinia in Australia incurved, 2.5-3 mm long, 2.5-3 mm wide, green, caducous. Male flowers: fragrant with a clove-like smell, diameter 10-15 mm; pedicels absent or up to 4 mm long; sepals 4, 2-whorled, outer pair joined, D-shaped, convex, 4-5 mm long, 4-5mm wide, green; petals 4, free, erect, imbricate, orbicular or oblong-obovate, may be convex, 5-7 mm long, 4.5-6 mm wide, cream or yellow, glabrous, margin usually recurved and thin; androecium 4-phalangiate; phalanges adnate to petals, obovate-elliptic at anthesis, c. 3 mm long, 2-2.5 mm wide; anthers several, most are clustered around phalange margin, bilobed, bithecate, c. 0.4 mm long and 0.2 mm wide, glabrous; pistillode rudimentary, attenuate-conical, c. 2.5 mm long. Female inflorescences terminal, a solitary flower or a 2 or 3(rarely 4)-flowered raceme, 17-25 mm long; peduncles 4-12 mm long; bracts at apex of peduncle triangular, keeled, 6-11 mm long, 3-5 mm wide, green with a yellowish primary vein, glabrous, caducous; bracteoles triangular, 3-4 mm long, 3-4 mm wide, green, caducous. Female flowers: fragrant with a clove-like smell, diameter 14-16 mm; pedicels absent or 3-5 mm long; sepals 4, 2-whorled, outer pair joined, inner pair free, D-shaped, convex, 5-6.5 mm long, 8-9 mm wide, green, glabrous, margin thin; petals 4, free, orbicular, convex, 9-11 mm long, 10-12 mm wide, cream or yellow, sometimes blushed with pink, glabrous, margin thin; staminodal phalanges 4, erect, adnate to petals, strap-shaped, c. 5 mm long and 3.5 mm wide, apex divided into 4-6 filaments with 1 or 2 antherodes on each side; filaments 1-1.5 mm long; antherodes obovate or rhomboid, c. 1 mm long; gynoecium oblong- globose, c. 8.5 mm long and 6-8 mm wide; ovary globose or 4-sided, 7-8 mm long and 6-8.5 mm wide, sessile, 4-locular, uniovulate; stigma mass sessile, domed, diameter 7-8 mm, cream becoming pink or reddish with age or exposure. Fruiting pedicel 3-5 mm long, 4 sepals persistent at base; fruit a fleshy berry, globular or depressed globular, often shortly beaked, 40-58 mm long, 50-75 mm wide, yellow or dark red to blackish; stigma shallowly 3 or 4-lobed or rayed, diameter 8-9.5 mm; seeds 1-4, 32-41 mm long, 23-35 mm wide; testa brown, smooth; aril fleshy, dark red, purple or cream, native mangosteen, Warren’s mangosteen. Fig. 2G, 3F & I. Additional selected specimens (from 144 examined): Papua New Guinea. Western District: Tarara, Wasi Kussa River, Dec 1936, Brass 8430 (BRI). Australia. Northern Territory. Jessie River, Melville Island, Aug 2000, Harwood907 (CNS). Queensland. Cook District: Gabba Island, Torres Strait, Mar 2003, Waterhouse 6592 (BRI); lama (Yam) Island, Torres Strait, Nov 2007, Fell 8855 & Stanton (BRI); Pajinka Lodge, Feb 2001, Cooper 1453 & Jensen (CNS); Escarpment of the Great Dividing Range, 18.2 km northeast of Heathlands Ranger Base, Heathlands D and O Reserve, Oct 1993, Fell 3759, Stanton & Dibella (BRI); Wattle Hills, Sep 2002, Sankowsky 1836 & Sankowsky (BRI); Stone Crossing, Wenlock River, Oct 1980, Hyland 1075 (CNS); Hey Point, Aug 1985, Gunness 1899 (BRI); Aurukun, banks of Watson River, Mar 2001, Smith 4609 (BRI); Ling Creek Spring, Steve Irwin Wildlife Reserve, c. 16km due NE of Stone Crossing, Wenlock River, Sep 2009, McDonald 2009- 09-034 & Addicott (BRI); 12.6 km WNW of Lockhart River Community, Mar 1994, Fell 4194 & Stanton (BRI); Nesbit River, Aug 1948, Brass 19915 (BRI); Rocky River Crossing, Oct 1996, Cooper 1011 & Cooper (CNS); Gap Creek, c. 38 km S by E of Cooktown, Aug 1959, Smith 10744 (BRI); Pilgrim Sands, Cape Tribulation, Jan 1993, Cooper 474 & Cooper (CNS); Clacherty Road, Julatten, Aug 2012, Cooper 2158 & Hawkes (CNS); Euluma Creek Road, Julatten, Aug 2012, Cooper 2159 & Hawkes (CNS); SFR 843, Parish of Bellenden Ker, Frenchman Creek, Sep 1986, Gray 4333 (CNS); Babinda, Aug 2003, Jago 119 (CNS). Distribution and habitat: Garcinia warrenii occurs in Australia and New Guinea. Within Australia it is recorded from north Queensland where it occurs in rainforest (complex mesophyll vine forest, mesophyll vine forest, complex notophyll vine forest, notophyll vine forest and simple microphyll vine-fern forest) and vine thickets on granitic, basaltic and rarely metamorphic soils from the Torres Strait islands to Babinda, altitude 0-1050 m. There are also two records from Melville Island, in the Northern Territory (Map 5) Phenology : Flowers have been recorded in July, August and September; fruit has been recorded in October, November, December, January and February. Etymology : The specific epithet is in honour of Dr William Henry Warren (1852-1926), Professor of Engineering at the University of Sydney. 26 Excluded Names Garcinia cherryi F.M.Bailey, Queensl. FI. 6: 103 (1902). Basionym of Ternstroemia cherryi (F.M.Bailey) Merr. ex J.F.Bailey & C.T.White (Theaceae). Acknowledgements Christina Zdenek, Brian Venables and John Pritchard are sincerely thanked for making several trips to check for flowers from the female tree of Garcinia jensenii. I am especially grateful to Rupert Russell for showing me G. russellii in the field, where he initially discovered them and for assisting on numerous trips across the Mossman River during varying water levels to finally collect enough fertile material for a formal description. Andrew Ford, Rigel Jensen and Jeanette Kemp have been invaluable in the field for their good company and sources of expertise. I also thank Frank Zich and Darren Crayn for support and access to the Australian Tropical Herbarium (CNS); Paul Forster for the complicated typification for G. xanthochymus and notes on the typification of G. mestonii, as well as other assistance; Ben Trupperbaumer for old German translations and Peter Bostock for translation of Latin descriptions as well as creating the maps. Bill Cooper, David Fell, Mark Harrington, Mel Harrison, Tim Hawkes, Bob Jago, Sarah Legge, Stephen McKenna, Steve Murphy, Garry Sankowsky, Allen Sheather, Peter Stanton & Gary Wilson are thanked for much appreciated assistance in various forms. Darren Crayn made valuable suggestions to an earlier manuscript. The curators of BRI, CANB, MEL & NSW are thanked for loans and/or photos of relevant material. Permits to collect were issued by the Queensland Department of Environment and Heritage Protection. References Anderson, T. (1874). Guttiferae. In J.D. Hooker, Flora of British India 1: 258-278. L.Reeve & Co.: London. Bailey, F.M. (1889). Report on new plants, preliminary to general report on botanical results on Meston’s expedition to the Bellenden-Ker Range, by F.M. Bailey, F.L.S., Colonial Botanist , p. 2. Department of Agriculture: Brisbane. Austrobaileya 9(1): 1-29 (2013) - (1899). Guttiferae. In The Queensland Flora , Part 1, pp. 102-105. H.J.Diddams & Co: Brisbane. - (1902). The Queensland Flora , Part 6, p. 103. H.J.Diddams & Co: Brisbane. Canbr (Centre for Australian National Biodiversity Research) (2010). Australian Tropical Rain Forest Plants Edition 6 (online version), viewed 20 June 2012. http://www.anbg.gov.au/cpbr/cd- key s/rfk/index. html Chah (2007). Australian Plant Census. http://www. anbg.gov.au/cgi-bin/apclist; Australian Plant Names Index, http://www.anbg.gov.au/cgi-bin/ apni Accessed November 2011. Christophel, D.C. & Hyland, B.P.M (1993). Leaf Atlas of Australian Rain Forest Trees. CSIRO Publishing: Melbourne. Cooper, W. & Cooper, W.T. (1994). Fruits of the Rain Forest. Geo: Sydney. - (2004). Fruits of the Australian Tropical Rainforest. Nokomis Editions: Melbourne. D’arcy, W.G. (1980). Flora of Panama. Family 123. Guttiferae. Annals of the Missouri Botanical Garden 67: 969-1043. De Candolle, A.P (1824). Guttiferae. In Prodromus Systematis Naturalis Regni Vegetabilis 1: 557- 564. Treuttel & Wtirtz: Paris. Dowe, J.L. & Broughton, A.D. (2007). F.M.Bailey’s ascent of Mt Bellenden-Ker in 1889, and notes on the publication priority of new vascular plant species from the Expedition. Austrobaileya 7: 555-566. - (2008). F.M.Bailey’s taxonomy of A.Meston’s collections from the Bellenden Ker Expedition of 1904. Austrobaileya 7: 677-679. Fleming, (circa 1795-1805). Fleming Indian Drawings Collection circa 1795-1805, Natural History Museum, London/Science Photo Library http:// www.sciencephoto.com/media/428689/enlarge Accessed July 2013 Hammel, B. (1993). In C.E. Jarvis, F.R. Barrie, DM. Allan & J.L. Reveal. A list of Linnean generic names and their types. Regnum Vegetabile 127: 28. Hooker, W.J. (1831). Xanthochymus dulcis. Sweet- fruited Xanthochymus. Curtis’s Botanical Magazine 58: t. 3088. - (1855). Garcinia mangostana. Mangosteen, or Mangostan. Curtis’s Botanical Magazine 81: t. 4847. Hyland, B.P.M. (1982). A revised card key to rainforest trees of North Queensland. CSIRO: Melbourne. 27 Cooper, Garcinia in Australia Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray, B. & Elick, R.W. (2003). Australian Tropical Rain Forest Plants. Trees, Shrubs and Vines. CD-ROM. CSIRO Publishing: Melbourne. Hyland, B.P.M., Whiffin, T., Zich, F. Edition 6. (2010). Australian Tropical Rain Forest Plants, CSIRO. http://www. anbg.gov.au/cpbr/cd-keys/rfk/ index.html Accessed October 2011. Jessup, L.W. (2007). Clusiaceae. In P.D. Bostock & A.E. Holland (eds.). Census of the Queensland Flora 2007, p. 54. Environmental Protection Agency, Brisbane. - (2010). Clusiaceae. In P.D. Bostock & A.E. Holland (eds.). Census of the Queensland Flora 2010, p. 44. Environmental Protection Agency: Brisbane. Kirtikar, K.R. & Basu, B.D. (1918). Indian Medicinal Plants. Sudhinra Nathbasu: Bahadurganj. Kostermans, A.J.G.H. (1980). Clusiaceae (Guttiferae). In M.D. Dassanayake (ed.), Flora of Ceylon 1: 72-110. Amerind Publishing Co. Pvt. Ltd.: New Delhi. Maheshwari, J.K. (1964). Taxonomic studies on Indian Guttiferae III. The genus Garcinia L. s.l. Bulletin of the Botanical Survey of India 6: 107-135. Marsden, W. (2005). The History of Sumatra containing an Account of the Government, Laws, Customs and Manners of the Native Inhabitants. ebook #16768 http://www.gutenberg.org/ files/16768/16768-h/16768-h.htm#sumatra-03 Accessed July 2013. Moore, S. (1917). A contribution to the phytogeography of Bellenden-Ker. II. Systematic account. Journal of Botany 55: 302-310. Mueller, F.J.H. (1891). Descriptions of new Australian plants with occasional other annotations (continued). The Victorian Naturalist 8(7): 109. Nazre, M. (2006). Taxonomic Revision and Molecular Studies of Garcinia Section Garcinia (Guttiferae ), unpublished PhD thesis. The University of Edinburgh. Opara, L.U., Studman, C.J. & Banks, N.H. (1997). Fruit Skin Splitting and Cracking. In J. Janick (ed.), Horticultural Reviews 19(4): 227. Richards, A.J. (1990a). Studies in Garcinia , dioecious tropical forest trees: agamospermy. Botanical Journal of the Linnean Society 103: 233-250. - (1990b). Studies in Garcinia , dioecious tropical forest trees: the origin of mangosteen. Botanical Journal of the Linnean Society 103: 301-308. Richards, J. (2008). The secret war: a true history of Queensland’s Native Police. University of Queensland Press: St Lucia. Robson, N.K.B. (1960). Guttiferae. In A.W. Excell & H. Wild (eds.). Flora Zambesiaca 1(2): 378-404. Crown Agents for Overseas Governments & Administrations: London. Roxburgh, W. (1805). Plants of the Coast of Coromandel 2: 51, t. 196. W. Bulmer and Co.: London. - (1820). Plants of the Coast of Coromandel 3: 66. W. Bulmer and Co.: London. Sharma, B.P., Singh, T.D., Handique, P.J., Devi, H.S. (2012). Unambiguous identification of Garcinia xanthochymus (Clusiaceae) in Manipur, India. Nebio 3(3): 7-10. Singh, N.P (1993). Clusiaceae. In B.D. Sharma & M. Sanjappa (eds.). Flora of India 86-159. Botanical Survey of India: Calcutta. Sompong, T. (2007). Floral and fruit morphology of some species in Garcinia spp. Songklanakarin Journal of Science and Technology 29: 245- 252. Stevens, P.F. (2007). Clusiaceae - Guttiferae. In K. Kubitzki (ed.). The Families and Genera of Vascular Plants. 9: 48-66. Springer-Verlag: Berlin/Heidelberg. Sweeney, P.W. (2008). Phylogeny and floral diversity in the genus Garcinia (Clusiaceae) and relatives. International Journal of Plant Sciences 169:1288-1303. - (2010). Floral anatomy in Garcinia nervosa and G. xanthochymus (Clusiaceae): A first step toward understanding the nature of nectaries in Garcinia. Bulletin of the Peabody Museum of Natural'History 51: 157-168. Te-chato, S. (2007). Floral and fruit morphology of some species in Garcinia spp. Songklanakarin Journal of Science Technology 29: 245-252. van Nooten, B.H. (1880). Fleurs, fruits et feuillages choisis de l ’ille de Java: peintes d ’apres nature. C.Muquardt: Bruxelles. White, C.T. (1933). Ligneous plants collected for the Arnold Arboretum in north Queensland by S.F. Kajewski in 1929. Contributions from the Arnold Arboretum of Harvard University 4: 1-113. - (1936). Contribution to the Queensland flora. No. 5. Proceedings of the Royal Society of Queensland 47: 51-84. Wight, R. (1840). Xanthochymus dulcis. leones Plantarum Indiae Orientalis. 1: t. 192. J.B.Pharoah: Madras. Zodiac Publications (undated). Australian Tropical Plants version 4.1 DVD ROM. Zodiac Publications. 28 Austrobaileya 9(1): 1-29(2013) Map 1. Distribution of Garcinia dulcis • (within Australia), G. brassii A and *G. livingstonei M. Map 3. Distribution of Garcinia leggeae A, G. mestonii ■ and *G. mangostana •. Map 2. Distribution of Garcinia jensenii A and G. zichii •. Map 4. Distribution of Garcinia gibbsiae A and G. russellii ■. Cooper, Garcinia in Australia 29 Map 5. Distribution of Garcinia warrenii • (within Australia). Map 6. Distribution of *Garcinia xanthochymus •. Reinstatement and revision of Sphaeromorphaea DC. and Ethuliopsis F.Muell. (Asteraceae: Plucheinae) A.R. Bean Summary Bean, A.R. (2013). Reinstatement and revision of Sphaeromorphaea DC. and Ethuliopsis F.Muell. (Asteraceae: Plucheinae). Austrobaileya 9(1): 30-59. Sphaeromorphaea DC. and Ethuliopsis F.Muell., historically included in Epaltes Cass., are reinstated and taxonomically revised. Sphaeromorphaea occurs in Australia, Indian Subcontinent, Indochina, China, eastern Asia, Malesia, Papuasia, northwestern Pacific and southwestern Pacific, and comprises six species. Three new species; S. ephemera A.R.Bean, S. major A.R.Bean and S. subintegra A.R.Bean are described, and two new combinations; S. harrisii (F.Muell.) A.R.Bean and S. littoralis (Retz.) A.R.Bean are made. Ethuliopsis occurs only in Australia and comprises the single species E. cunninghamii (Hook.) F.Muell. A neotype is chosen for Epaltes australis Less., and lectotypes are chosen for Artemisia littoralis Retz., Ethulia cunninghamii Hook, and Epaltes harrisii F.Muell. Descriptions, illustrations and distribution maps are provided for all taxa. Diagnostic morphological characters and ecological aspects are discussed, and identification keys are provided to the species and to the Australian genera of Subtribe Plucheinae. Key Words: Asteraceae, Plucheinae, Epaltes, Ethuliopsis, Sphaeromorphaea, Sphaeromorphaea ephemera, Sphaeromorphaea harrisii, Sphaeromorphaea littoralis, Sphaeromorphaea major, Sphaeromorphaea subintegra, Australia flora, India flora, China flora, taxonomy, new species, identification key, distribution maps A.R. Bean, Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: Tony.Bean@science.dsitia.qld.gov.au Introduction Sphaeromorphaea DC. and Ethuliopsis F.Muell. are often regarded as synonyms of Epaltes Cass. (Cooke 1986; Bremer 1994; Anderberg & Eldenas 2007; GRIN 2012) and their histories are intertwined. Cassini (1818) named the genus Epaltes with a single species Epaltes divaricata (L.) Cass., native to the Indian subcontinent, China and Indochina. C.F. Lessing added Epaltes australis Less., E. mexicana Less, and E. hirsuta Less, in 1830 and 1832. In the ensuing century, nearly 20 Epaltes species were described from all tropical parts of the world. Sphaeromorphaea was named by De Candolle (1838) with three species at that time, two originating from southern Asia and one from Australia. No other species were subsequently described under that generic Accepted for publication 16 July 2013 name; however, the combination S. australis (Less.) Kitam., based on Epaltes australis Less., was made in 1936. In Australia, Mueller (1861) proposed a new genus, Ethuliopsis , to accommodate Ethulia cunninghamii Hook. Bentham (1867) relegated Ethuliopsis to synonymy with Epaltes , while maintaining Epaltes australis ; the two species he placed in Epaltes (E. australis , E. cunninghamii (Hook.) Benth.) were to remain there for well over a century. Bentham (1873) dealt incompletely with Sphaeromorphaea , saying correctly only that it is, in part, synonymous with Centipeda Lour. One of the three named species, S. centipeda DC., is a nomenclatural synonym of Centipeda minima (L.) A.Braun & Asch. Hooker (1882) accepted Sphaeromorphaea with a single species (for India), S. russeliana DC., and placed the genus next to Centipeda in his systematic arrangement. Bean, Sphaeromorphaea and Ethuliopsis Mueller described three further species under Epaltes (E. harrisii F.Muell., E. tatei F.Muell. and E. pleiochaeta F.Muell.) from Australia; Moore & Betche (1893) listed E. pleiochaeta as an accepted species, and Bailey (1900) listed E. harrisii as an accepted species, but neither name ever came into general usage. Epaltes tatei is an accepted species that was transferred to a new genus, Haegiela P.S.Short & Paul G.Wilson (Short & Wilson 1990), belonging to the Tribe Gnaphalieae. Contemporary Asian flora treatments have used either Epaltes australis or Sphaeromorphaea australis for the taxa found there. For Taiwan, Li (1978) accepted S. australis , but Peng (2004) reversed this, with Epaltes australis then the accepted name; Naithani (1995) accepted S. australis , placing the genus near Centipeda in the Tribe Anthemideae\ Hu & Wong (2009) used E. australis as the accepted name, with S. australis given as a synonym. In recent decades Epaltes has been regarded as a heterogenous genus incorporating any Plucheoid taxa whose achenes lack pappus bristles (Leins 1971; Merxmueller et al. 1978; Cooke 1986; Anderberg 1991). The present author agrees that Epaltes is paraphyletic, and that the Australian species currently referred to that genus belong in Sphaeromorphaea DC. or Ethuliopsis F.Muell. Dunlop (2000) was the first author to accept the genus name Sphaeromorphaea (and the species S. australis ) in an Australian flora treatment and some major Australian herbaria subsequently adopted Sphaeromorphaea for their state in plant checklists or censuses (Walsh & Stajsic 2007; Short et al. 2011). Short & Wilson (1990) foreshadowed the acceptance of the genus name Ethuliopsis , but the Australian Plant Census (APC 2012) continued to list it as a synonym of Epaltes. Sphaeromorphaea and Ethuliopsis are revised in this paper. Six species of Sphaeromorphaea are enumerated, including three new species and two new combinations; all of these species have previously been referred to a broadly circumscribed Epaltes 31 australis. They occur in Australia, Indian subcontinent, Indochina, China, eastern Asia, Malesia, Papuasia, northwestern Pacific and southwestern Pacific. Ethuliopsis is monotypic, and is endemic to Australia. Materials and methods This revision is based on a morphological study of herbarium specimens from A, AD, AMES, BM, BRI, C, CANB, GH, K, LD, MEL, PE, PERTH and TAI. Images of specimens, mainly types, from BM, C, CAL, G-DC, K, M and P have been examined. In addition, the author has collected and studied plants in the field in Queensland, New South Wales and the Northern Territory. All measurements were made from dried material, using a stereo microscope equipped with a graticule. Commonly used abbreviations in the specimen citations are HS for Homestead, NP (National Park), SF (State Forest). The distribution maps were compiled using DIVA-GIS Version 7.5.0, using label data of specimens from the herbaria listed above. Geographical regions are listed according to the scheme of Brummitt (2001); for the generic distributions, the areas cited are Regional Names; for the specimen citations, the areas cited are Level 3 Names. Specimen citations within Queensland are further subdivided into Pastoral districts. Relationships Anderberg (1991) provided a cladistic analysis of the subtribe Plucheinae based on morphological characters. He reinstated the African genus Litogyne Harv., and transferred Epaltes gariepina (DC.) Steetz to it, based mainly on the very different shape of the receptacle. Litogyne has since gained some acceptance as a distinct genus (Retief & Herman 1997; Beentje 2002). Anderberg (1991) studied material of the following Epaltes species: E. australis , E. brasiliensis (Link) DC., E. cunninghamii and E. mattfeldii. Interpretation of his data is fraught because E. australis is here 32 regarded as belonging to Sphaeromorphaea, E. cunninghamii is regarded as belonging to Ethuliopsis , and the generic status of E. brasiliensis and E. mattfeldii is uncertain. The type species of Epaltes (E. divaricata ) was not included in the analysis. Anderberg (1991) placed Thespidium F. Muell. ex Benth. and Coleocoma F.Muell. in the ‘Coleocoma group’, while Epaltes was placed in the ‘Pluchea group’. The present author is of the opinion that while Epaltes sens. str. and Ethuliopsis probably do belong in the ‘Pluchea group’, Sphaeromorphaea belongs to the ‘Coleocoma group’ and is most closely related to the monotypic genus Thespidium. The persistent ebarbellate bristles formed on the achenes of some Sphaeromorphaea spp. are very similar to those of Thespidium basiflorum (F.Muell.) Benth., and the cylindrical longitudinally- ribbed achenes are a feature of both genera. Austrobaileya 9(1): 30-59 (2013) Leins (1971) had previously proposed a close relationship between Epaltes australis and Thespidium based on a study of the pollen grains and style morphology. The molecular study of Anderberg et al. (2005) placed Coleocoma , Streptoglossa Steetz and Epaltes cunninghamii (i.e. Ethuliopsis) as part of a largely unresolved clade dominated by species of Pluchea Cass. The presence of a coronal pappus is a uniting character for all species of Sphaeromorphaea and Ethidiopsis. Coronal pappus is not common in Asteraceae, being confined to some genera in Tribe Anthemideae and Tribe Inuleae (Heywood & Humphries 1977; Mukherjee & Sarkar 2001). The species of Sphaeromorphaea and Ethuliopsis differ from Epaltes sens. str. (E. divaricata) in a number of significant ways (Tables 1 & 2): Table 1. Morphological character differences between Sphaeromorphaea and Epaltes divaricata Characters Sph aeromorph aea Epaltes divaricata stems not winged winged capitula borne usually in pairs borne singly involucral bracts mostly obtuse, cartilaginous, incurved all acute, herbaceous, straight or excurved marginal florets conical or lageniform, or filiform with an expanded base filiform, not significantly expanded at the base style of disc florets rather slender, not divided style arms short and broad, distinctly divided pappus a stiff persistent corona, bristles sometimes present no corona, no bristles achenes of disc florets full sized, often indistinguishable from marginal-floret achenes not formed; ovary vestigial achene shape cylindrical ellipsoidal to obovoid carpophore prominent white ring small, obscure Bean, Sphaeromorphaea and Ethuliopsis 33 Table 2. Morphological character differences between Ethuliopsis and Epaltes divaricata Characters Ethuliopsis Epaltes divaricata stems not winged winged sexuality subdioecious bisexual receptacle hemispherical (in heterogamous capitula) flat style of disc florets rather slender, not divided style arms short and broad, distinctly divided pappus a persistent cylindrical corona, bristles present on disc florets no corona, no bristles achene shape and ribbing lunate; 1-ribbed ellipsoidal to obovoid; several-ribbed carpophore prominent white ring small, obscure In Ethuliopsis , some plants bear homogamous capitula (with disc florets only), while other plants bear heterogamous capitula (nearly all marginal florets, but with a few disc florets). In Epaltes divaricata , all capitula on all plants are heterogamous. Ecology and distribution Ethuliopsis cunninghamii is an erect annual herb reaching one metre in height. It is found on alluvial flats with heavy clay soils, in swamps, or beside river channels and lakes. It is endemic to arid and semi-arid eastern Australia. All Sphaeromorphaea species are herbs or small shrubs 15-60 cm high or wide. Some species are seemingly annual (e.g. S. ephemera ), while other species can develop a substantial taproot and probably live for two or three years, with older stems dying, replaced by new stems shooting from near the base of the plant. They prefer sunny areas with bare soil in grasslands, sedgelands, woodlands or forests. They appear to have no edaphic preference, occurring on soils as diverse as pure beach sand and heavy cracking clays. All Sphaeromorphaea species inhabit places where the soil is intermittently or permanently moist or wet. Poor drainage is readily tolerated, but they will not persist where there is standing water. They can most often be found on alluvial flats, in swamps, or beside rivers and lakes; however, some species can also occur in hilly terrain, where they occupy the wetter microhabitats, such as depressions, drains and gullies. Most species display a considerable tolerance for high levels of salt in the soil or in the air. One species (Sphaeromorphaea major A.R.Bean) is confined to marine couch grasslands, and most other species sometimes inhabit salt affected areas. S. australis can grow on coastal headlands, exposed to salty winds. Label data suggest that S. subintegra A.R.Bean sometimes grows on serpentinite, a substrate well known to be toxic to many plant species. Waterbirds are well known as dispersal agents for the seeds of wetland plants (e.g. Figuerola & Green 2002; Soons et al. 2008), and although there is no literature documenting the dispersal of Sphaeromorphaea or Ethuliopsis , it seems reasonable to assume that waterbirds play a major role. This would help explain the distribution of S. littoralis , which is widespread from southern Australia to southern Asia. Its apparent absence from Indonesia may be partly due to the 34 predominance of rainforest communities that are not suitable habitat for the light-loving Sphaeromorphaea. It is interesting to note that Sphaeromorphaea and the ecologically comparable genus Centipeda (taxonomy revised by Walsh [2001]) have similar distribution patterns, to the extent of possessing a single widespread species shared by southern Asia and Australia, and a centre of diversity in Australia. Some diagnostic morphological characters 1. Resin globules Ethuliopsis and most species of Sphaeromorphaea have resin globules on the lower and often on the upper leaf surfaces, and on the corollas and achenes. They are shining, transparent and globose, but may be difficult to see in very old herbarium specimens, as with age they degrade and deflate. Their presence, distribution and sometimes their numbers are diagnostic. S. major does not have resin globules. 2. Leaf and stem indumentum Sphaeromorphaea major has glabrous leaves and stems. All other species bear crisped, multicellular, eglandular hairs, at least on young developing leaves and stems. For S. ephemera and Ethuliopsis cunninghamii, leaves are glabrous at maturity; S. australis may be glabrous or sparsely hairy. For S. littoralis, S. harrisii and S. subintegra, at least some hairs persist on mature leaves. Leaf indumentum is particularly variable in S. littoralis and S. harrisii, sometimes sparsely hairy and sometimes densely hairy on fully expanded leaves. 3. Leaf shape and margins Sphaeromorphaea species have leaves that at first glance appear petiolate, but they are in fact sessile, as the lamina does extend to the stem in all taxa. In most species, the leaf shape is more or less obovate, but S. major is distinguishable by being especially narrow¬ leaved. No species has entire margins; in S. subintegra the margins are denticulate with teeth less than 0.5 mm long, and hence may appear entire without close inspection. In Austrobaileya 9(1): 30-59 (2013) S. australis, some large teeth or lobes (>1 mm long) are usually present, often on the basal part of the lamina (the pseudo-petiole). Sphaeromorphaea littoralis and S. harrisii similarly are often conspicuously toothed on some leaves, with the teeth extending to the basal half of the lamina. Ethuliopsis cunninghamii leaves are not pseudo-petiolate, and the lamina is broad almost to the base. The leaf margins are denticulate to dentate. 4. Marginal floret shape The shape of the marginal florets is an important diagnostic character for Sphaeromorphaea. In S. littoralis, S. subintegra and S. ephemera, the corolla is consistently conical, tapering more or less evenly from the base to the very narrow apex (e.g. Fig. 3E); in S. australis and S. harrisii, the corolla is consistently lageniform i.e. swollen in appearance, broadest above the base, then tapering rapidly towards the apex (Figs. 1G, 2C); in S. major and Ethuliopsis cunninghamii, the corolla of the marginal florets is filiform (Figs. 1C, 4H). 5. Achene features All Sphaeromorphaea achenes are cylindrical, providing an instant distinction from Ethuliopsis whose achenes are lunate. Ethuliopsis achenes have a single adaxial longitudinal rib, although several anastomosing veins on the surface are often also visible. The number of longitudinal ribs for Sphaeromorphaea species varies from 5-14, and is moderately diagnostic for each species. In S. littoralis and S. harrisii, there is a whorl of antrorse to appressed twin hairs at the base of the achene, with a few twin hairs sometimes occurring also on the achene body. In other species, the twin hairs are nearly always absent, although S. australis and S. subintegra may occasionally have 1-4 twin hairs at the achene base. 6. Pappus corona All species of Sphaeromorphaea and Ethuliopsis have a stiff persistent coronal pappus. In Ethuliopsis cunninghamii and in Sphaeromorphaea major, the corona is Bean, Sphaeromorphaea and Ethuliopsis erect and shortly cylindrical with an entire or fimbriate margin, while in the other species of Sphaeromorphaea , it is annular or disc¬ like, transverse or nearly so, with an entire, pentagonal or erose margin (Fig. 3G). The width (and therefore the visibility) of this corona varies with the species; in S. littoralis , S. harrisii and S. subintegra it is relatively broad and easy to see, while in S. australis and S. ephemera , it is very narrow and difficult to detect. 7. Pappus bristles In some species of Sphaeromorphaea pappus bristles emerge from the margin of the corona. They are consistently ascending, terete, persistent, and shorter than the corolla. With the exception of S. major , they lack barbellae. Sometimes only a single bristle is produced, but there may be as many as five. The length and number of bristles produced may differ according to the type of floret; S. major can produce up to five bristles on the disc achenes, but not more than three on the marginal achenes. Pappus bristles occur frequently in S. harrisii and S. major , but only rarely in S. littoralis and S. subintegra ; they have not been observed for S. australis and S. ephemera. Ethuliopsis consistently produces slightly flattened barbellate bristles on the disc florets, but never any bristles on the achenes. In summary, Sphaeromorphaea is characterised by the hemispherical to oblate capitula borne singly or in pairs opposite a leaf; the cartilaginous incurved glabrous involucral bracts; the presence of a coronal pappus (sometimes produced into slender persistent bristles); the cylindrical achenes, formed from all florets (though achenes produced from disc florets may not be fertile), glabrous or with straight twin hairs mainly at the base. Ethuliopsis is characterised by the subdioecious habit; the vestigial ovary of the disc florets; white corollas; barbellate pappus bristles on the disc florets; and the glabrous one-ribbed lunate achenes with a cylindrical coronal pappus, obliquely placed on the achene. 35 Taxonomy Sphaeromorphaea DC., Prodr. 6: 140 (1838). Type: S. russeliana, fide J.R. Drummond in Merrill & Rolfe (1908: 126) [=S. littoralis ]. Annual or perennial herbs with a well developed rootstock. Latex absent. Indumentum of crisped, multicellular, eglandular hairs. Sessile to very shortly stalked resin globules frequently present on stems, leaves, involucral bracts, corolla and achenes, initially shiny and transparent, becoming opaque and deflated with age. Stems terete, not winged. Leaves alternate, sessile, spreading; margins denticulate, dentate or lyrate; base attenuate, not decurrent. Capitula axillary or leaf-opposed, solitary or in pairs on often short peduncles, hemispherical to depressed-globose, disciform, heterogamous. Involucral bracts subequal in length, cartilaginous, incurved, in 3-4 rows, innermost row conspicuously narrower. Receptacle flat, glabrous, without paleae. Marginal florets fertile, female, without rays; corolla without multicellular hairs, resin globules present or absent, white or pink to purple at anthesis; lobes 3, tiny; style with bulbous base, branches divergent. Disc florets bisexual, usually producing achenes, but in some species infertile; corolla cylindrical to narrowly campanulate, without multicellular hairs, resin globules present or absent, pink to maroon; lobes 4 or 5; anthers tailed, apically obtuse; style bulbous at the base, divided or undivided at apex, sweeping hairs obtuse and extending well down the shaft. Achenes cylindrical, longitudinally ribbed; carpopodium ring-like, prominent, white. Pappus comprising a stiff, transverse or erect annular corona, sometimes produced into short slender persistent bristles. Twin hairs (when present) straight, antrorse to appressed, in a basal whorl, and occasionally on body of achene. Six species, occurring in Australia, Indian subcontinent, Indochina, China, eastern Asia, Malesia, Papuasia, northwestern Pacific and southwestern Pacific. 36 Austrobaileya 9(1): 30-59 (2013) Key to the species of Sphaeromorphaea using microscopic characters 1 Corolla of marginal florets lageniform.2 1. Corolla of marginal florets conical or filiform.3 2 Coronal pappus 0.02-0.05 mm wide, obscure; leaves glabrescent; marginal florets with several resin globules; pappus bristles always absent; achene hairs usually absent, or rarely 1-4 present at base.2. S. australis 2. Coronal pappus 0.05-0.08 mm wide, obvious; leaves persistently hairy; marginal florets with few or no resin globules; pappus bristles frequently present; hairs 5-20 at base of achene.3. S. harrisii 3 Fully expanded leaves sparsely to densely hairy; achenes with several resin globules ... 4 3. Fully expanded leaves glabrous; achenes without resin globules.5 4 Leaves dentate or denticulate, teeth present on basal half of leaf; peduncles 2-14 mm long; achenes with 10-40 hairs, mainly at the base.5. S. littoralis 4. Leaves denticulate, teeth absent from basal half of leaf; peduncles 1.5-3.5 mm long; achene hairs usually absent, or rarely 1-4 present at base . . . 6. S. subintegra 5 Pappus bristles absent; young stems and leaves sparsely hairy; fertile achenes 0.45-0.7 mm long.4. S. ephemera 5. Pappus bristles present on some florets; all stems and leaves glabrous; fertile achenes 1.0-1.3 mm long.1. S. major Field key to the species of Sphaeromorphaea 1 Leaves narrow, 5.5-14 times longer than broad, without resin globules.1. S. major 1. Leaves broader, 2.3-5.3 times longer than broad, resin globules present.2 2 Older leaves glabrous, bright green.3 2. Older leaves sparsely to densely hairy, green to grey-green.4 3 Annual; achenes without resin globules, marginal florets conical.4. S. ephemera 3. Perennial; achenes with resin globules, marginal florets lageniform.2. S. australis 4 Teeth on leaf margin very small on all leaves and absent from basal half. . 6. S. subintegra 4. Teeth on leaf margin > 1 mm long on some leaves, teeth present on basal half.5 5 Most stems erect; capitula almost sessile; pappus bristles frequently present; leaves deeply and irregularly lobed.3. S. harrisii 5. Most stems sprawling; capitula conspicuously pedunculate; pappus bristles rarely present; leaves shortly lobed, lobes fairly regular.6 6 Leaves grey-green; achenes with numerous twin hairs at base and with conspicuous corona.5. S. littoralis 6. Leaves bright green; achenes lacking twin hairs at base (or just a few present) and with very narrow corona.2. S. australis Bean, Sphaeromorphaea and Ethuliopsis 37 Diagnostic characters for Sphaeromorphaea species Achenes with 5^10 twin hairs, mainly at base S. harrisii, S. littoralis Achenes with 1-4 twin hairs at base, or twin hairs absent S. australis, S. ephemera, S. subintegra, S. major Achene ribs 5-10 S. australis, S. harrisii, S. littoralis, S. major Achene ribs 10-14 S. australis, S. ephemera, S. littoralis, S. subintegra Corona transverse, prominent, >0.05 mm wide S. harrisii, S. littoralis, S. subintegra Corona transverse, obscure, < 0.05 mm wide S. australis, S. ephemera Corona erect, entire or fringed S. major Marginal floret lageniform S. australis, S. harrisii Marginal floret conical S. ephemera, S. littoralis, S. subintegra Marginal floret filiform S. major Pappus bristles frequently present S. harrisii, S. major Pappus bristles rarely present S. littoralis, S. subintegra Pappus bristles absent S. australis, S. ephemera 1. Sphaeromorphaea major A.R.Bean sp. nov. affinis S. anstrali sed acheniis longioribus costis longitudinalibus paucioribus, flosculorum discoideorum corolla 2-2.5 mm longa (in S. australi 1-1.3 mm) et foliis acheniis corollisque globulis resinae carentibus differens. Typus: Queensland. Port Curtis District: 100 metres from Huttonvale track, SE of Sabina Point, Shoalwater Bay Training Area, 8 April 2011, A.R. Bean 30830 & D. Halford (holo: BRI [1 sheet + spirit]; iso: BM, CANB, MEL, MO, distribuendi). Prostrate or procumbent perennial shrub to 30 cm high and 60 cm across. Stems glabrous. Leaves linear to narrowly oblanceolate, 11-85 x 2-15 mm, 5.5-14 times longer than broad, glabrous, concolorous, resin globules absent; margins dentate to denticulate, with 3-6 pairs of teeth 0.1-07 mm long; apex acute or obtuse. Capitula solitary or in pairs, leaf opposed, hemispherical, 3-4.5 mm long, 4.5- 8 mm wide; peduncles 2-28 mm long. Involucral bracts incurved, glabrous; median bracts ovate, 27-3.2 x 1.3-1.8 mm, apex acute or obtuse. Marginal florets 75-250, in several rows, female; corolla cylindrical, 1.5- 2.4 mm long, apex very narrow, white, without resin globules. Disc florets 16-68, functionally male; corolla cylindrical to narrowly campanulate, 2-2.5 mm long, 0.25- 0.35 mm wide near base, 0.4-0.5 mm wide near apex, pink, resin globules absent; lobes 4 or 5, triangular, each 0.4-0.5 mm long; style undivided. Achenes formed from all florets, but those from disc florets infertile. Marginal achenes fertile, cylindrical, 1-1.3 mm long, 0.25-0.3 mm wide, brown; ribs 5-10, white; twin hairs absent or 1-4, resin globules absent. Pappus corona almost erect, 0.02-0.05 mm high, margin entire or fimbriate; pappus bristles absent or with 1-3 bristles up to 1.9 mm long, and (in inland populations) with numerous short bristles. Disc achenes infertile, cylindrical, 0.8-17 mm long, 0.3-0.4 mm wide, white; ribs absent; pappus corona erect, white, margin entire or fimbriate; pappus bristles absent or with 1-5 barbellate bristles arising from the corona, up to 1.9 mm long. Fig. 1A-D. Additional specimens examined : Queensland. Port Curtis District: Broad Sound, Sep 1802, Brown s.n. (CANB, MEL); Bay of Inlets, Jun 1770, Banks & Solander s.n. (BM); 2 km NW of Seahound Hard ramp, Shoalwater Bay Training Area, N of Yeppoon, Feb 2012, Bean 31610 & Mathieson (AD, B, BRI, US, W, to be distributed); Ramsay Crossing, squatters 38 Austrobaileya 9(1): 30-59 (2013) Fig. 1. A-D. Sphaeromorphaea major. A. Flowering branchlet x0.5. B. disc floret and infertile achene with pappus bristles xl6. C. marginal floret x32. D. achene x32. E-I, S. australis. E. flowering branchlet xl. F. disc floret x32. G. marginal floret x32. H. achene, lateral view x32.1. achene, plan view x32. A-D from Bean 30830 & Halford (BRI); E from Bean 9548 (BRI); F-I from Kemp 3362H& Kutt (BRI). Del. W.Smith. Bean, Sphaeromorphaea and Ethuliopsis camp, 40 km NW of Gladstone, near Curtis Island, Dec 2004, Batianoff 0412502 & Halford (BRI); Targinnie district, 18.5 km NW of Gladstone, Nov 2011, Fell DGF QEQ79/1 (BRI). Mitchell District: Bowen Downs, in 1873, Birch s.n. (MEL); Second Spring, Edgbaston Reserve, NE of Aramac, Apr 2012, Bean 31643 (A, BRI, DNA, G, HO, MEL, NSW, to be distributed); Measuring Spring, Edgbaston Reserve, NE of Aramac, Apr 2012, Bean 31646 (BRI, CANB); Edgbaston, Mar 1998, Fensham 3464 (BRI). South Kennedy District: Little Moses Spring, Doongmabulla Station, 165 km NW of Clermont, May 2012, Danielsen 1555 (BRI). Distribution and habitat: Sphaeromorphaea major is endemic to Queensland. It is known from some coastal locations near Rockhampton, and three inland locations at Edgbaston Reserve, Bowen Downs and Doongmabulla station (Map 1). On the coast, it is confined to those salt marsh areas (inundated by spring tides and dominated by grassland of Sporobolus virginicus (L.) Kunth) that apparently receive freshwater seepage from adjacent eucalypt or melaleuca forest. At the inland locations, it grows on the margins of artesian springs in open salt-encrusted areas dominated by Cyperus laevigatus L., but not where there is surface water. Phenology: Flowers and fruits have been recorded from September to April. Notes: Banks and Solander were the first to collect this species, from ‘Bay of Inlets’ (near Stanage Bay) in 1770, and Solander gave it the manuscript name Cotula glabrata. Robert Brown then collected it in 1802; the label on his specimen at BM [Bennett 2131] states that the plant was seen “in paludosis salsis Port 1, Keppel Bay, Shoalwater Bay [&] Broad Sound”. ‘Port 1’ and ‘Keppel Bay’ are near the southern and northern ends of Curtis Island respectively (Vallance et al. 2001), places where there are no modern collections. The area covered by Brown’s excursions at ‘Broad Sound’ (between the present-day towns of Ogmore and Carmila) is also without modern collections. For this species, there was a time lag of 243 years between the first herbarium collection and formal description. This lag figure is surely the highest on record for an Australian plant taxon, and it may well be a record not easily broken. Fontaine et al. (2012) found that, for species described in 2007, the 39 average time lag (or ‘shelf life’) was around 21 years for all species, and around 32 years for plant species. Sphaeromorphaea major differs from the sympatric S. australis by the achenes 1-1.3 mm long (0.65-0.9 mm long in S. australis ); marginal florets 1.7-2.2 mm long (0.5-0.9 mm long in S. australis)-, leaves without resin globules (always present in S. australis, though sometimes hard to see); no resin globules on achenes or corolla (always present in S. australis ). The Edgbaston populations have a coronal pappus with a strongly fimbriate margin, while the coastal populations have an entire corona margin. The Doongmabulla specimen is intermediate with regard to this character, with a weakly fimbriate coronal pappus. There seem to be no other consistent morphological differences between the coastal and inland populations. Sphaeromorphaea major can be distinguished by its narrow leaves and often very long peduncles. It is the only species in which the leaves lack eglandular multicellular hairs and resin globules. Furthermore, its disc florets are functionally male and the achenes produced are infertile, while in the other species the “disc achenes” are usually fully fertile. Conservation status: Sphaeromorphaea major is known from about 400 plants at Edgbaston Reserve near Aramac, on the margins of three springs. The population size at Doongmabulla station is unknown. Its continued existence at Bowen Downs is unknown. The author has observed several hundred plants at Shoalwater Bay Training Area, and Brown’s recordings of it suggest that there is a strong possibility of extending the known distribution. However, in view of the very specialised habitat, the total area covered is unlikely to exceed 40 square km. Applying the Red Fist criteria (IUCN 2001), a status of ‘Near Threatened’ is recommended. Etymology: The specific epithet refers to the disc florets, which are larger in this species than in any other species of Sphaeromorphaea. 40 2. Sphaeromorphaea australis (Less.) Kitam., Acta Phytotax. Geobot. 5(4): 276 (1936), 6(2): 80 (1937); Epaltes australis Less., Linnaea 5: 148 (1830), Syn. Gen. Compos. 206 (1832); Erigerodes australe (Less.) O.Kuntze, Revis. Gen. PL 1: 335 (1891). Type: not cited (B, t). Neotype (here designated): Australia: New South Wales. Myall Creek road, Bungawalbin National Park, c. 22 km SSE of Casino, 30 December 2011, A.R. Bean 31450 (neo: BRI [1 sheet + spirit]; iso: CANB, E, K, MEL, NSW, RSA, US, W, to be distributed). Sphaeromorphaea petiolaris DC., Prodr. 6: 140 (1838). Type: [Australia: New South Wales.] Port Jackson, undated [in 1819], C. Gaudichaud (syn: G-DC, image!; P, image!); Port Jackson, undated [in 1823], F.W. Sieber 697 (syn: G-DC, image!). Illustrations : Logan River Branch SGAP (2005: 130); Stephens & Sharp (2009: 44), both as Epaltes australis. Prostrate to ascending perennial herb, to 30 cm high. Stems with sparse multicellular hairs when young, glabrescent. Leaves spathulate, obovate or oblanceolate, 26-70 x 7-21 mm, 2.6-5.3 times longer than broad, discolorous, resin globules present on both surfaces; young leaves with sparse multicellular hairs, often absent at maturity; margins dentate to denticulate, with 4-8 pairs of teeth up to 5 mm long, the longer lobes frequently on the attenuate basal part of the lamina; apex obtuse or acute. Capitula solitary or in pairs, leaf opposed, hemispherical to oblate, 2-3 mm long, 4-5.5 mm wide; peduncles 1-6 mm long. Involucral bracts incurved; median bracts ovate, 1.5-1.9 x 0.8-1.2 mm, glabrous or with a few resin globules on outer surface, apex acute. Marginal florets 150-300, in several rows, female; corolla lageniform, 0.5-0.9 mm long, pink to purple, resin globules few to numerous, apex very narrow. Disc florets 13-34, bisexual or functionally male; corolla narrowly campanulate, 1-1.3 mm long, 0.25-0.3 mm wide near base, 0.4- 0.45 mm wide near apex, pink to maroon, resin globules present throughout; lobes 4, triangular, each c. 0.2 mm long; style undivided. Achenes formed from all florets, but those from disc florets sometimes infertile, Austrobaileya 9(1): 30-59 (2013) white to pale yellow. Marginal achenes fertile, cylindrical, 0.65-0.9 mm long, 0.2-0.25 mm wide, brown, ribs 8-12, white, twin hairs absent or 1-4 at base; resin globules few to numerous, shining. Pappus corona transverse, obscure, translucent, 0.02-0.05 mm wide, with numerous radial striations, margin entire to erose; pappus bristles absent. Fig. 1E-I. Additional selected specimens examined : Taiwan. Tamsuy [Tan-shui], 1864, Oldham s.n. (GH); Houlung, Miaoli County, Aug 1968, Kao 7364 (TAI). New Caledonia. Koutio-Koudta, Dumbea, Sep 1971, McKee 24355 (NOU). Australia: Western Australia. 1 km N of Serpentine, Feb 1985, Keighery 7182 (PERTH); Lambkin Reserve, Serpentine, Apr 1996, Keighery 14346 (PERTH). Queensland. Cook District: Rocky Isles, Great Barrier Reef, c. 18 km S of Cooktown, Jun 1969, Done s.n. (BRI [AQ7513]). North Kennedy District: Alva Beach, Jun 1949, Smith 4400 (BRI). South Kennedy district: Western edge of Lake Buchanan, Yarrowmere Station, Mar 1998, Kemp 3362H & Kutt (BRI). Mitchell District: Myross, east of Aramac, May 2000, Fensham 3885 (BRI). Leichhardt District: Baralaba - Woorabinda Road, 1.8 km W of Dawson Range, Mar 2005, Bean 23551 (BRI); Humboldt, 45 km NE of Rolleston, Jan 1996, Bean 9548 (BRI). Port Curtis District: Livingstone Shire Vegetation Survey, site 227 behind Lammermoon Beach, Sep 1977, Batianoff 806 & McDonald (BRI, CANB). Burnett District: Allies Creek SF, Compartment 146, W of Middle Creek Road, Apr 2004, Halford Q8271 & Jessup (BRI). Wide Bay District: Old Hollett Road, c. 5 km SW of Noosaville, Dec 2003, Bean 21577 (BRI). Maranoa District: Around barracks, Carellen, c. 90 km WNW of Bollon, Mar 2008, Bean 27456 (BRI). Darling Downs District: Girraween NP, 22 km S of Stanthorpe, Apr 2001, Batianoff 210534 & Collyer (BRI). Moreton District: Hay’s Landing, Wivenhoe Dam, N of Fernvale, Jan 2001, Bean 17303 (BRI). New South Wales. Tenterfield, May 1921, Cheel s.n. (NSW); Pilliga, Nov 1932, Rupp 15 (NSW); Trial Bay to Laggers Point, NE of Kempsey, Jan 1953, Constable s.n. (NSW, US); c. 4 km by road W then N of Crowdy Head on road to Diamond Head, Mar 1981, Haegi 2026 (AD, MEL, NSW); 3 miles [5 km] along Currambean Creek Road, 5 miles [8 km] S of Nowra, Mar 1951, Ford s.n. (NSW); Huskisson, Jervis Bay, Apr 1916, Rodway 4509 (NSW). Distribution and habitat : Sphaeromorphaea australis occurs naturally in Australia and in New Caledonia. In Australia it extends from Ulladulla, New South Wales to Townsville, Queensland (with an outlier further north near Cooktown), and up to 600 km inland e.g. Lake Buchanan, Bollon, Pilliga (Map 2). It is naturalised in Taiwan (first collection in 1864) and near Perth, Western Australia (first collection in 1985) (Map 3). In New South Bean, Sphaeromorphaea and Ethuliopsis Wales and southern Queensland, it occupies a range of habitats in forest dominated by Eucalyptus spp., Acacia harpophylla F.Muell exBenth. ox Melaleuca spp., and sometimes in marine couch grassland adjacent to mangrove trees. Towards the northern end of its native range, the species appears to be increasingly confined to saline habitats, such as coastal salt marsh, artesian springs or the edges of saline inland lakes. Soil ranges from white sands to dark heavy clays. Phenology : Flowers and fruits can be found in any month of the year. Typification: Lessing (1830) catalogued all Asteraceae specimens held at the Berlin Botanical Museum (B), wherein he provided a three word description of Epaltes australis as a footnote, without mention of the specimen(s) upon which the name is based. Soon after, in his Synopsis Generum Compositarum , he (Lessing 1832) expanded the description of E. australis slightly, again without referring to the specimen(s) used. He did not explicitly give the country of origin, but did state that the genus is found in “Asiae vel Novae- Hollandiae vel Americae aequinoctialis” The other three species treated ( E. divaricata, E. mexicana, E. hirsuta) were known to him from India, Mexico and India respectively, so it follows that E. australis is the Australian species. While it is unclear from reading A.P. de Candolle’s treatment of E. australis (de Candolle 1836) whether he viewed the specimen that Lessing used to describe it, in hindsight it is obvious that he did not. Candolle cited a specimen for E. australis , collected by Cunningham from the Lachlan River in New South Wales and thanks to a paper by O.W. Sonder (1853), we know this is not the correct application of the name Epaltes australis. Sonder examined both Lessing’s specimen at B and the Cunningham specimen cited by de Candolle, and he indicated unambiguously (Sonder 1853: 482) that E. australis sensu de Candolle was not that of Lessing, and that Ethulia cunninghamii Hook, is synonymous with E. australis sensu de Candolle. He further stated (p. 485) that the cited specimens of Sphaeromorphaea petiolaris DC. belong to 41 the same taxon as Lessing’s (type) specimen of Epaltes australis. However, Sonder did not impart any information about the collector or provenance of Lessing’s type specimen. Nearly all material at B was destroyed during World War II (Hiepko 1987), including the type of E. australis , and the absence of information about the collector, date of collection or provenance of the type (except ‘Novae-Hollandiae’) means that there is no starting point to search for potential duplicate specimens. As no known type material exists for E. australis , a neotype is here chosen from material collected in “Novae-Hollandiae”, i.e. Australia, and from the taxon that occurs around Sydney, the most likely collection site of the original specimen. Notes : Sphaeromorphaea australis is distinguished from other species in the genus by its dark green leaves, the frequent presence of conspicuous lobes on the basal attenuate part of the lamina; the lageniform marginal florets; the narrow and inconspicuous coronal pappus; the lack of pappus bristles; and the absence (usually) of hairs at the base of the achene. Leaves are often glabrous at maturity, but specimens from the more westerly areas have leaves that often retain some hairs at maturity. In Queensland, its distribution overlaps extensively with Sphaeromorphaea subintegra, but both species maintain their identities without any sign of intergradation or hybridization. S. subintegra may be differentiated by its pale green leaves with persistent (though often sparse) hairs and denticulate margins (teeth <0.5 mm long), the conical marginal florets, and the relatively broad and conspicuous coronal pappus. Conservation status : A common and widespread species. 3. Sphaeromorphaea harrisii (F.Muell.) A.R.Bean comb, nov.; Epaltes harrisii F.Muell., Fragm. 11: 101 (1880); Erigerodes harrisii (F.Muell.) O.Kuntze, Revis. Gen. PL 1: 335 (1891). Type: Australia: Queensland. Cook District: Possession Island, in 1880, C.C. Harris s.n. (lecto [here designated]: K 42 000373331; isolecto: MEL 2159948, NSW 582206). Illustrations : Britten (1901: t. 158), as Epaltes australis ; Dunlop (2000: 185), as Sphaeromorphaea australis. Erect annual or perennial herb to 35 cm high. Stems with dense multicellular hairs when young, hairs persistent. Leaves spathulate to oblanceolate, 22-61 x 7-21 mm, 2.5-3.8 times longer than broad, discolorous or concolorous, resin globules present on both surfaces; young leaves with dense multicellular hairs, fully expanded leaves sparsely to densely hairy; margins dentate to lyrate, rarely denticulate, with 6-10 pairs of teeth up to 5 mm long; apex acute. Capitula solitary or in pairs, hemispherical to cupular, 3-3.5 mm long, 4.5-6 mm wide; peduncles 1-3 mm long. Involucral bracts incurved; median bracts ovate, 2-2.5 x 0.7-1.1 mm, glabrous, apex acute. Marginal florets 200-300, in several rows, female; corolla lageniform, 0.9-1.3 mm long, maroon to purple, resin globules few or none, apex very narrow. Disc florets 15-31, bisexual; corolla cylindrical, 1.25-1.4 mm long, c. 0.3 mm wide near base, c. 0.35 mm wide near apex, pink to maroon, resin globules scattered throughout; lobes 4, triangular, 0.2-0.25 mm long; style undivided; pappus bristles sometimes present. Achenes formed from all florets, but those from the disc florets sometimes infertile, white to pale yellow, not ribbed. Marginal achenes fertile, cylindrical, 0.7-1.1 mm long, 0.2-0.3 mm wide, brown, ribs 6-10, white, twin hairs 5-20 at base; resin globules numerous. Pappus corona transverse, pale and translucent, 0.05-0.08 mm wide, margin entire to erose; pappus bristles absent or present, slender, without barbellae, 0-6 per floret, 0.1-1 mm long. Fig. 2A-E. Additional selected specimens examined : Papua New Guinea. Western Province: Arufi, Wassi Kussa River, Jul 1968, Henty & Katik NGF38641 (A, BRI, CANB, K, L); Bula Plains, Morehead subdistrict, Nov 1972, Henty & Foreman NGF49355 (A, BRI, CANB, K, L, LAE); Garmari on the Bula Plains, c. 30 km S of Bensbach Lodge, Apr 1997, Mitchell 4696 (BISH, BRI, LAE); Balamuk, Bensbach River, c. 9 km S of Bensbach Wildlife Lodge, May 1992, Waterhouse BMW2503 & Obedi (BRI). Australia: Northern Territory. Nhulunbuy Lagoon, Oct 1993, Egan 2804 (DNA). Queensland. Cook District: Saibai Island, Sep 1994, Austrobaileya 9(1): 30-59 (2013) Wannan 130 (BRI); [North] Possession Island [Iem Islet], Sep 1792, Smith & Wiles s.n. (BM); Possession Island, in 1880, Harris s.n. (MEL2162771); Thursday Island, June 1897, Bailey s.n. (BRI [AQ247412]); 1 km S of Seisia, 250 m from beach, Nov 1999, Wannan 1412 (BRI, CANB, NSW); New Holland, in 1770, Banks & Solander s.n. (BRI [AQ450752]; MEL); Bolt Head, Temple Bay, Jul 1991, Forster PIF8985 (BRI, DNA, MEL); Coen River [Pennefather River], Gulf of Carpentaria, Nov 1802, Brown s.n. (CANB); 8.5 km NW of Weipa Mission, Dec 1974, Specht W243 & Salt (BRI); Evans Landing rubbish dump, Weipa, Aug 1996, Waterhouse BMW3918 (BRI); 11 km N of the Chester River Crossing, W of the Embley Range, Silver Plains Station, Jun 1992, Forster PIF10539 et al. (BRI); 18.6 km from Running Creek on the track to Old Port Stewart, Jun 1993, Clarkson 10103 & Neldner (BRI, DNA, L, MEL); Lakefield NP, 29.4 km by road from Hann River Crossing, Aug 2011, McDonald KRM11779 (BRI); East of mouth of Muck River, Cape Melville NP, Jul 1998, Bean 13666 (BRI); Lakefield NP, 22.8 km by road N of Ranger base, Aug 2011, McDonald KRM11776 (BRI); Lizard Island, Nov 1997, Muir s.n. (BRI [AQ667812]); Lizard Island, Dec 1974, Specht LI420 & Specht (BRI). North Kennedy District: Edmund Kennedy NP, near Cardwell, Jan 1992, Bean 3912 (BRI, DNA, MEL). Distribution and habitat : In Australia Sphaeromorphaea harrisii occurs along the east coast of Queensland from Cardwell to Cape York and on several of the Torres Strait islands as well as in the north-eastern Northern Territory. It is also found in southern Papua New Guinea (Map 4). It grows on a wide range of soil types, from sand to black clay, on floodplains and riverbanks where Melaleuca spp. or Lophostemon suaveolens (Sol. ex Gaertn.) Peter G.Wilson & J.T.Waterh. are dominant. It also occurs at the base of sand dunes, and in Sporobolus virginicus grassland adjacent to mangrove communities. Phenology : Flowers and fruits can be found in any month of the year. Typification: The original material of Epaltes harrisii (now distributed in K, MEL and NSW) comprises numerous very young plants that bear only one or two capitula. Most capitula are immature; only the material at K has florets at anthesis, hence the K sheet has been selected as the lectotype. There is a second gathering of S. harrisii at MEL (MEL2162771) with the same label details as the type. This sheet comprises three mature plants bearing many flowers and mature achenes. Judging by the description Bean, Sphaeromorphaea and Ethuliopsis 43 Fig. 2. A-E, Sphaeromorphaea harrisii. A. Flowering branchlet *1. B. disc floret and achene with pappus bristle x32. C. marginal floret x32. D. achene, lateral view x 32. E. achene, plan view *32. F-J, S. ephemera. F. flowering branchlet xl. G. disc floret and achene x32. H. marginal floret x32.1. achene, lateral view x32.1. achene, plan view x32. A, from Forster PIF10539 et al. (BRI); B-E from McDonald KRM11779 (BRI); F-J from Forster PIF22217 & Booth (BRI). Del. W.Smith. 44 in the protologue, Mueller did not use this collection when naming the species and hence it cannot be considered a type. Notes : Sphaeromorphaea harrisii is distinctive in the erect stems, the lageniform marginal florets with few or no resin globules; capitula not as broad as in other species; the very short peduncles (often c. 1 mm long); the frequent presence of pappus bristles; and the achenes with few (6-10) ribs. It is perhaps closest to S. australis , but differs from that species by the more upright habit, persistently hairy leaves, the longer corolla of the marginal florets, the broader coronal pappus and the frequent presence of pappus bristles. Pappus bristles are present on most S. harrisii collections, but completely lacking in others (including the type). There is no geographical basis to this variation. For example, Specht LI420 & Specht, from Lizard Island, is without pappus bristles; while Muir s.n., also from Lizard Island, has numerous pappus bristles. Conservation status : A common and widespread species. Etymology : This species was named for Charles Christopher Harris (1844-1887), an amateur naturalist. He was borne in Maine, U.S.A., and immigrated to Australia in the 1860s. He arrived at Cooktown in 1873 or 1874, and resided there until his death. He was accidently killed while on board the sailing vessel ‘Spey’ on 30 th April 1887, when the boom struck him on the head (M.Scully pers. comm.; National Library of Australia (2009- onwards)). 4. Sphaeromorphaea ephemera A.R.Bean sp. nov. affinis S. australi sed habitu annuo, flosculis foemineis conicis et acheniis brevioribus globulis resinae carentibus differens. Typus: Australia: Queensland. Gregory North District: Bladensburg National Park, S of Winton, Opalton Road, 19 March 1998, P.I. Forster PIF22217 & R. Booth (holo: BRI). Prostrate to ascending annual herb, to 15 cm high. Stems with sparse multicellular hairs when young, glabrescent. Leaves spathulate Austrobaileya 9(1): 30-59 (2013) to oblanceolate, 14-53 x 5-17 mm, 2.8-4.8 times longer than broad, discolorous, resin globules present on both surfaces; young leaves with sparse multicellular hairs, usually absent at maturity; margins dentate to denticulate, with 3-12 pairs of teeth mostly < 0.5 mm long, but with some up to 2 mm long; apex obtuse. Capitula solitary or in pairs, leaf opposed, hemispherical, 1.5-2 mm long, 4-5 mm wide; peduncles 1-7 mm long. Involucral bracts incurved; median bracts ovate, 1.4- 2.3 x 0.5-0.9 mm, glabrous or with a few resin globules on outer surface, apex acute. Marginal florets 100-200, in several rows, female; corolla conical, 0.6-0.7 mm long, pink to purple, broadest at base, few to several resin globules present, apex narrow. Disc florets 13-24, bisexual; corolla cylindrical to narrowly campanulate, 1-1.1 mm long, 0.25- 0.3 mm wide near base, 0.25-0.35 mm wide near apex, pink to maroon, resin globules present throughout; lobes 4, each c. 0.2 mm long; style undivided. Achenes formed from all florets, cylindrical, tapered at both ends, 0.45-0.7 mm long, 0.15-0.2 mm wide, brown, ribs 10-14, white, twin hairs absent; resin globules absent, or rarely 1 or 2. Pappus corona transverse, obscure, translucent, 0.01-0.04 mm wide, with numerous radial striations, margin entire; pappus bristles absent. Fig. 2F-J. Additional specimens examined'. Queensland. South Kennedy District: Cudmore, Apr 1997, Fensham 3130 (BRI). Mitchell District: Lake Inveresk, 70 km NNE of Muttaburra, Mar 2004, Cumming 22432 & Thompson (BRI). Warrego District: Mariala NP, 52.5 km east by road from Adavale, Aug 2009, Forster PIF35727 & Thomas (BRI); Road reserve, south of Biloola HS, 2 km off Adavale Road, west of Charleville, May 2010, Wang JW0180 (BRI); 60 km NW [actually WSW] of Charleville, off the Diamantina Developmental Road, Jun 1976, Purdie 373D (BRI). Distribution and habitat : Endemic to western Queensland, between Adavale and Winton (Map 1). Most occurrences are along drainage lines under shrubland of Acacia aneura F.Muell. ex Benth., often with emergent Eucalyptus populnea F.Muell., and in red clayey soils. One site is on the margins of a salt lake, with fine sandy soil. Phenology : Flowers and fruits have been recorded from March to August. Bean, Sphaeromorphaea and Ethuliopsis Affinities : Sphaeromorphaea ephemera is apparently related to S. australis, but differs by the conical marginal florets, the annual habit, and the shorter achenes that lack resin globules. S. ephemera differs from S. littoralis by the glabrescent leaves, the shorter achenes that lack resin globules, the absence of twin hairs on the achenes, and the narrower pappus corona. Conservation status : Data deficient. While the species is known from just a few specimens, it is potentially common. Etymology : From the Greek ephemeros, meaning ‘short-lived’. 5. Sphaeromorphaea littoralis (Retz.) A.R.Bean comb, nov.; Artemisia littoralis Retz., Observ. Bot. 5, p. 28 (1789); Epaltes littoralis (Retz.) Less., Linnaea 6: 151 (1831). Type: seashores of eastern India, undated [1768-1785], J.G. Koenig s.n. (lecto, here designated: C 10007263, image!). Epaltes hirsuta Less., Syn. Gen. Compos. 206 (1832). Type: ‘In India orientali lecta in herbario Thunbergiano’, J.P. Rottler s.n., n.v. Cotula quinqueloba Blanco, FI. Filip. [F.M. Blanco] 626 (1837), nom. illeg. non L.f. (1782). Sphaeromorphaea russeliana DC., Prodr. 6: 140 (early Jan 1838); in Deless., Icon. Sel. 4, t. 49 (Feb 1838); Sphaeromorphaea russeliana DC. var. russeliana. Prodr. 6: 140 (early Jan 1838); in Deless., Icon. Sel. 4, t. 49 (Feb 1838); Centipeda russeliana (DC.) F.Muell. as ‘ russellianaFragm. 8: 142 (1874). Type: India. The Circars, undated [1781-1790], P. Russell s.n. (syn: G-DC, image!; syn: K-W, image!). Cotula russeliana Wal 1., Ntimer. List[Wallich] 3240 (1831), nomen nudum Illustrations : Gagnepain (1924: 557), as Sphaeromorphaea russeliana, Cunningham et al. (2011: 685), as Epaltes australis. Prostrate to sprawling perennial herb, to 25 cm high. Stems with dense multicellular hairs when young, hairs persisting at maturity. Leaves spathulate to oblanceolate or obovate, 9-60 x 4-15 mm, 2.3-4.8 times longer than broad, concolorous, resin globules present on 45 both surfaces; young leaves densely hairy, hairs sparse to dense at maturity; margins dentate to denticulate, with 4-14 pairs of teeth usually less than 1 mm long, but occasionally up to 2 mm long; apex acute. Capitula solitary or in pairs, hemispherical, 2.5-3.5 mm long, 3.5-7 mm wide; peduncles 2-14 mm long. Involucral bracts incurved, glabrous; median bracts ovate, 1.6-2.3 x 0.8-1.2 mm, glabrous or with few to many resin globules on outer surface, apex acute. Marginal florets c. 200, in several rows, female; corolla conical, 0.7- 1.2 mm long, pink to purple, resin globules numerous, apex very narrow. Disc florets lb- 40, bisexual; corolla cylindrical, 1.1-1.4 mm long, 0.25-0.4 mm wide, pink, resin globules present throughout; lobes 4, triangular, each 0.2-0.3 mm long; style undivided, arms parallel. Achenes formed from all florets, 0.7-1.3 mm long, 0.25-0.35 mm wide, brown, ribs 8-14; twin hairs 10-40, mainly at base but with some scattered above; resin globules numerous, shining. Pappus corona transverse, circular or somewhat pentagonal, 0.05-0.09 mm wide, with numerous radial striations, margin entire, erose or rarely toothed; pappus bristles usually absent, rarely present. Fig. 3. Additional selected specimens examined : India. Hazaribagh, Apr 1884, Clarke 34806 (K); Eastern India, s.dat ., Koenig s.n. (LD); Visakhapatnam, Waltan Beach near university, Sep 1969, Rao 7101 (CAL). China Southeast. Chung Chi college, Hong Kong, Apr 1972, Hu 11888 (A, K, US); Nanning, Guangxi province, Nov 1935, Liang 67192 (PE); Yongning, Guangzi province, Jul 1958, ZhongA62026 (PE). Hainan: Yaichow, Hainan Island, Sep 1933, Liang 62870 (GH, US); Danzhou county, Hainan, May 1928, Tsang 16813 (PE, US). Thailand. Nawng Kai, Chaiyaburi, Feb 1924, Kerr 8543 (BM); Klong Hoy Kong, Songkla province. May 1986, Maxwell 86-305 (A); Kao Sabab, Chantaburi, Feb 1935, Seidenfaden 2735 (C). Vietnam. Near beach, Tourane, May-Jul 1927, Clemens & Clemens 3079 (BM, K, US); Phu Yen province, Jan 1936, Petelot 6285 (GH, US); Nha Trang, s.dat., Poilane 6046 (K, US). Malaysia. Near Kampong Padangnegeri, Kuala Trengganu, Jul 1953, Sinclair 39806 & Salleh (BM); Tumpat, Kota Bahru, Feb 1917, Ridley 535 (K). Philippines. Manila, Jan 1915, Merrill Species Blancoanae No 773 (AMES, BM, CAL, GH, NSW, US); Caloocan, province of Rizal, Luzon, Nov 1903, Merrill 3655 (US). Marianas, southern end of Cabras Island, Guam, Mar 1966, Evans 1673 (US); Waste ground just inland from Asan Point, Guam, Jun 1983, Raulerson 4561 (US). Australia: Western Australia. Piccaninny Creek Gorge, 15 km SE of Bungle Bungle Outcamp, Bungle Bungle Range, Apr 1985, Blackwell BB297 (PERTH); King Edward River, old CRA 46 Austrobaileya 9(1): 30-59 (2013) Fig. 3. Sphaeromorphaea littoralis. A. Flowering branchlet xl.5. B. leaf surface with resin globules and eglandular multicellular hairs x24. C. flowering capitulum x8. D. disc floret and achene x32. E. marginal floret x32. F. achene, lateral view x32. G. achene, plan view x32. A-B from Bean 23777 (BRI); C from Brass 8806 (BRI); D-G from Badman 1473 (AD). Del. W. Smith. campsite, 1 km S of track to old Mitchell River Station, Jun 1988, Edinger 541 (DNA, PERTH); Drysdale River above Mogurnda Creek, Drysdale River NP, Aug 1975, George 13570 (CANB, PERTH). Northern Territory. 1.5 km NE of Fish River Gorge in China Wall, Barkly Tableland, Jun 1974, Kanis 1823 (CANB, DNA, L, US); SE corner of entry to Lake Surprise, Mar 1973, Maconochie 1672 (DNA, MO, NSW); Mary Ann Dam, c. 5km NE of Tennant Creek, May 2005, Bean 23777 (BRI); c. 12 miles [c.19 km] NE Finniss River HS, Oct 1971, Must 842 (CANB, DNA, NSW); Near mouth of Foelsche River, Jul 1948, Perry 1829 (AD, BRI, CANB, MEL, NSW, US). South Australia. Carruranna Waterhole, Cooper Creek, 32 km WNW of Etadunna HS, Feb 1984, Badman 1473 (AD, BRI, CANB, MEL, NSW); c.lkmN of Big Bend S of Swan Reach, May 1973, Donner 4113 (AD, BRI); Sunnyside opposite Mypolonga on riverbank, Dec 1961, Symon 1926 (AD, Bean, Sphaeromorphaea and Ethuliopsis BRI). Queensland. Cook District: Cumberland, Gilbert River, May 1937, Brass 8806 (BRI). Burke District: eastern shore of Lake Moondarra, 16 km NNE of Mt Isa, Oct 1997, Fraser 245 (A, CANB). Mitchell District: Yalleroi, Mar 1946, Clemens s.n. (BRI [AQ247452]; GH). Gregory North District: SE edge of Lake Machattie, c. 70 km SE of Bedourie, May 2007, Bean 26406 (BRI). Warrego District: Currawinya NP, Lake Numalla, Mar 1996, Forster PIF18688 (BRI). New South Wales. Cobar, Sep 1910, Abrahams 387 (NSW); 4 miles [6 km] S of Gwabegar, between Baradine and Pilliga, Nov 1967, McGillivray 2746 (NSW); Cobham Lake, c. 32 km SSE of Milparinka, Mar 1972, Milthorpe 711 (NSW). Victoria. Dimboola, Feb 1901, D’Alton s.n. (NSW); Post No 7 on Hattah Nature Drive, Hattah NP, Jan 1989, DAubert 483 (MEL, NSW); On SW side of Lake Lockie, 465 m NNE (at 188 degrees) from the E end of Lockie Track, Mar 2007, Stajsic 4364 (MEL, NSW). Distribution and habitat : Sphaeromorphaea littoralis is widely distributed in southern Asia, with native occurrences in India, Thailand, Vietnam, Malaya, Philippines and southeastern China. It is naturalised on the island of Guam (Map 6). In Australia, it is indigenous in the Kimberley region of Western Australia, South Australia, Northern Territory, western Queensland, western New South Wales and north-western Victoria (Map 5). The species inhabits sunny places on the margins of creeks and dams, saline coastal flats and swamps, and in Australia extends into areas receiving very low annual rainfall. Soils are variable. Phenology : Flowers and fruits may be found at any time of the year. Typification: I am aware of just two specimens of Artemisia littoralis that were collected by J.G. Koenig - one at C and one at LD. The LD specimen is notable for the presence of 1 or 2 pappus bristles, up to 0.6 mm long, on many of its achenes. The specimen at C lacks these bristles, and it is therefore deduced that these two specimens were from separate gatherings. The C specimen is superior in quality, and pappus bristles are not mentioned in the protologue; hence it is chosen as the lectotype. Type material of Epaltes hirsuta was sought from all herbaria that are listed as having collections of C.P. Thunberg or J.P Rottler (HUH 2012), but without success. However, I consider it a synonym of S. littoralis ; the words “capitulis solitariis vel geminis” given 47 in the protologue strongly suggests that E. hirsuta is a Sphaeromorphaea , and the “tota hirsuta” accords with the hairy Indian form of S. littoralis. Notes: Sphaeromorphaea littoralis differs from S. australis by the grey-green hairy leaves (green glabrous leaves in S. australis)', the conical-shaped marginal florets that have numerous resin globules (lageniform florets with few resin globules for S. australis), the prominent corona on the achenes; and the numerous hairs at the base of the achene. Sphaeromorphaea littoralis differs from S. subintegra by the longer teeth on the leaf margins, presence of teeth on the basal half of the leaves, fully expanded leaves moderately to densely hairy, mostly longer peduncles (to 14 mm, versus 1.5-3.5 for S. subintegra), and the presence of numerous antrorse hairs at the base of the achene. Australian specimens of S. littoralis tend to have larger leaves than is usual for Asian specimens. Indian and northern/central Australian specimens are generally densely hairy, while specimens from south-east Asia (e.g. Thailand, China) and around the Murray River in southern Australia are generally only sparsely hairy. According to Cunningham et al. (2011), it is completely ignored by domestic livestock. The collector of the type of S. russeliana was Patrick Russell, usually spelt with two “l’s” (Hawgood 1994). However, early references (Wallich 1831; Wight 1831; De Candolle 1838) cited his surname as ‘Russel’, while Lowndes (1834) cited his surname as ‘Russel’ or ‘Russell’. The spelling of the epithet ‘russeliana’ by de Candolle (1838) was clearly intentional and that spelling must be maintained (Art. 60). Wallich (1831) included a Russell manuscript name (Artemisia chrysanthemum) as a synonym of his Cotula russeliana [Cat. 3240], The latter, also a nomen nudum, was later validated as Sphaeromorphaea russelliana DC. The epithet chrysanthemum means ‘golden-yellow flowers’, and so some later accounts (Hooker 1882; Li 1978; Naithani 1995) and some specimen labels have stated 48 that Sphaeromorphaea has yellow flowers. All taxa and populations of Sphaeromorphaea known to the present author have pink, purple, maroon or (rarely) white florets, and the occurrence of yellow flowered populations is considered highly unlikely. It is probable that Russell’s manuscript name referred to another species e.g. Grange a maderaspatana (L.) Poir., and Wallich merely misapplied it. Sphaeromorphaea littoralis is naturalised at Guam, being first recorded in 1966 (Fosberg & Sachet 1980), adjacent to the Naval Base. Conservation status : A common and widespread species. 6. Sphaeromorphaea subintegra A.R.Bean sp. nov. affinis S. australi sed infoliis dentibus aequabiliter parvis, pappo coronali comparate lato et conspicuo, pilis persistentibus in foliis maturis et corolla foeminea conica (in S. australi lageniformi) differens. Typus: Australia: Queensland. South Kennedy District: Lambert Beach, Slade Point, Mackay, 16 July 1992, G.N. Batianoff9207103 & H.A. Dillewaard (holo: BRI; iso: MEL). Prostrate to ascending perennial herb, to 25 cm high. Stems with sparse to dense multicellular hairs when young, hairs ± persistent. Leaves spathulate to obovate, 14-50 x 4-19 mm, 2.4-37 times longer than broad, discolorous, resin globules present on both surfaces; young leaves with moderately dense multicellular hairs, some hairs persisting at maturity; margins denticulate, with 3-12 pairs of small teeth <0.5 mm long; apex obtuse. Capitula solitary or in pairs, leaf-opposed, hemispherical, 2-3 mm long, 5-6 mm wide; peduncles 1.5-3.5 mm long. Involucral bracts incurved; median bracts 1.5-2.5 x 0.8-0.9 mm, glabrous or with a few resin globules on outer surface, apex acute. Marginal florets 150-300, in several rows, female; corolla conical, 0.6-0.9(-l) mm long, mauve to purple, broadest at base, with few to numerous resin globules; apex narrow. Disc florets 22-44, bisexual; corolla cylindrical to narrowly campanulate, 0.9-1.3 mm long, 0.25-0.35 mm wide near base, 0.25-0.4 mm wide near apex, pink to maroon, resin globules numerous; lobes 4, triangular, each 0.15-0.25 Austrobaileya 9(1): 30-59 (2013) mm long; style undivided. Achenes formed from all florets, cylindrical, 0.7-0.9 mm long, 0.25-0.3 mm wide, brown, ribs 10-14, twin hairs absent or 1-4 at base; resin globules numerous, shining. Pappus corona transverse, obvious, white to grey, 0.05-0.08 mm wide, margin entire to erose; pappus bristles absent or very rarely present. Fig. 4A-E. Additional selected specimens examined : Papua New Guinea. Milne Bay Province: Narian, Misima Island, Aug 1956, Brass 27606 (A); Joe’s Landing, Sudest Island, Aug 1956, Brass 27780 (A, US). Western Province: Daru, near airstrip, Jun 1973, Henty NGF49517 (A, BRI); Daru Island, Sep 1963, Womersley NGF17786 (BRI). New Caledonia. Presqu’ile de Ouano, Sep 1971, Raynal 16504 (NOU). Australia: Queensland. Cook District: Booby Island, Great Barrier Reef, c. 34 km from Thursday Island, Jul 1969, Heatwole s.n. (BRI [AQ8142]); East of the Mossman to Daintree Road north of Wonga Beach, Sep 2002, Jago 6297 (BRI); Mareeba, Jan 1918, White s.n. (BRI [AQ247418]). North Kennedy District: E of Mount Garnet, Feb 2008, Wannan 4999 (BRI, NSW); Little Ramsay Bay, Hinchinbrook Island, Apr 2012, Mathieson MTM1327 (BRI, NSW); Snake Creek, c. 95 km NW of Charters Towers on Gregory Developmental Road, Apr 1991, Batianoff SC9104010 (BRI). South Kennedy District: Brampton Island near airstrip, Sep 1986, Batianoff 5123 (AD, BRI); Black Wattle Creek Crossing, on Gregory Development Road, May 1991, McFadyen 453 (BRI); Collinsville Coal Pty Ltd mine lease area, near Collinsville and Scottville, Apr 1984, Thompson 587 (BRI). Leichhardt District: 8.8 km from Mt Dillingen peak and c. 87 km NNE of Clermont, Feb 1999, Bailey LB65 (BRI); 45.8 km from Taroom, on road to Bauhinia Downs, May 2010, Bean 29617 (BRI). Port Curtis District: 200m down road to St Lawrence from Bruce Highway, Jul 1998, Thompson 1317 (BRI); Facing Island, Oct 2004, c. 0.7 km NW of Catcombe, Halford Q8670 (BRI). Burnett District: About 3.3 km SE of Culcraigie along Culcraigie - Eidsvold Road, Apr 1997, Pollock ABP391 (BRI); Toondahra HS yards, Jun 1989, Forster PIF5076 (BRI, US). Wide Bay District: Near Maroondan, E of Gin Gin, Nov 1996, Bean 11229 (BRI). Distribution and habitat : Sphaeromorphaea subintegra is recorded from southern lowland parts of Papua New Guinea, New Caledonia, and in Australia is widespread in eastern Queensland from the Torres Strait islands to Mundubbera (Map 7). It grows on wet or moist microhabitats in open forest and woodland dominated by Eucalyptus or Melaleuca , often on alluvium, but also on hillsides and plateaux. Soils vary from sand to reddish basaltic clay, and including soils derived from serpentinite. It has not been recorded from salt marsh communities. Bean, Sphaeromorphaea and Ethuliopsis 49 Fig. 4. A-E, Sphaeromorphaea subintegra A. Flowering plant *0.6. B. disc floret and achene *32. C. marginal floret x32. D. achene, lateral view x32. E. achene, plan view x32. F-J, Ethuliopsis cunninghamii. F. flowering branchlet xl. G. disc floret xl6. H. marginal floret x32. L achene, abaxial view x32. J. achene, adaxial view x32. A, from Bean 29617 (BRI); B-E from McFadyen 453 (BRI); F, from Cowan 81 (BRI); G, from Mitchell 824 (BRI); H-J from Emerson s.n. (BRI [AQ501325]). Del. W.Smith. 50 Phenology : Flowers and fruits can be found in any month of the year. Notes : Sphaeromorphaea subintegra differs from S. australis by the consistently small teeth on the leaves (no more than 0.5 mm long), the relatively broad and conspicuous coronal pappus, persistent hairs on mature leaves, and the marginal corolla conical in shape. Field observations and herbarium specimen labels suggest that where the distributions of this species and S. australis overlap, S. subintegra inhabits the more mesic sites (where fresh water is more readily available), whereas S. australis inhabits salt-affected sites, including salt marsh or headlands near the ocean, or sites that may dry out more quickly. No intergradation between these two species has been observed in the field or in the herbarium, even though they sometimes occur less than a kilometre from each other. S. subintegra differs from S. littoralis by the lobing of the leaves (more conspicuously lobed in S. littoralis ) and the very few twin hairs on the achene, if any (numerous twin hairs in S. littoralis). The oldest herbarium specimen of S. subintegra that I have seen is one collected by Ernst Betche at Cooktown in 1881. Conservation status : A common and widespread species. Ethuliopsis F.Muell., Fragm. 2: 154 (1861); Epaltes sect. Ethuliopsis (F.Muell.) F.Muell., Fragm. 10: 100 (1877). Type: E. dioica F.Muell. (= E. cunninghamii). Gynaphanes Steetz in Peters, Naturw. Reise Mossambique [Peters] 6(2): 457 (1864). Type: Gynaphanes australis (= Ethuliopsis cunninghamii). Perennial herbs. Latex absent. Stems terete, not winged. Leaves alternate, sessile, toothed, spreading, not decurrent. Capitula pedunculate, in terminal panicles; on some plants disciform and heterogamous; on other plants homogamous, with disc florets only. Involucral bracts variable in length, cartilaginous, in 3-6 rows. Receptacle glabrous, without paleae. Marginal florets Austrobaileya 9(1): 30-59 (2013) fertile, female, without rays; corolla lobes 3, tiny; style base not bulbous, branches divergent. Disc florets bisexual, functionally male, ovary vestigial, not producing achenes. Anthers tailed, apically obtuse. Style base not bulbous, style undivided at apex; sweeping hairs obtuse, extending well down the shaft. Achenes lunate, with a single longitudinal rib on adaxial side; carpopodium annular, white, prominent. Pappus on the marginal florets comprising a persistent erect cylindrical corona; on the disc florets comprising 3-6 barbellate bristles. A monotypic genus, endemic to Australia. Ethuliopsis cunninghamii (Hook.) F.Muell., Key to the System of Victorian Plants 1: 313 (1888); Ethulia cunninghamii Hook, in T.Mitch., J. Exped. Trop. Australia 62 (1848); Ethuliopsis dioica F.Muell, Fragm. 2: 155 (1861), nom. illeg .; Gynaphanes australis Steetz in Peters, Naturw. Reise Mossambique [Peters] 6(2): 458 (1864), nom. illeg .; Epaltes cunninghamii (Hook.) Benth., FI. Austral. 3: 530 (1867); Erigerodes cunninghamii (Hook.) O.Kuntze, Revis. Gen. PI. 1: 335 (1891). Type citation: “We saw... Also an ETHULIA, which may, on further examination, constitute a new genus; it was found by Allan Cunningham on the Lachlan” Type: Australia: New South Wales. Swampy banks of the Lachlan River, 26 June 1817, A. Cunningham 290/1817 (lecto [here designated]: K 000373334; isolecto: G-DC). Illustrations : Cunningham et al. (2011: 685), as Epaltes cunninghamii. Erect herb, to 100 cm high. Stems glabrous. Leaves oblong to oblanceolate, 20-69 x 4.5- 17 mm, 3.6-8.5 times longer than broad, bright green, concolorous, resin globules sometimes present on lower surface; glabrous or with sparse multicellular hairs; margins denticulate to dentate with 5-13 pairs of teeth each 0.3-3 mm long; apex obtuse or acute; base cuneate to obtuse. Capitula in small terminal panicles. Peduncles 0-4.5 mm long, glabrous or sparsely glandular-hairy. Homogamous capitula. Capitula cylindrical to campanulate at anthesis, 3.5-5 mm long, 2.5- 5.5 mm wide. Involucral bracts flat, Bean, Sphaeromorphaea and Ethuliopsis glabrous, outer ones much shorter than inner; median involucral bracts 3.1-3.5 x 0.6-1.1 mm, apex obtuse, margin entire. Receptacle flat. Marginal florets absent; disc florets 20- 35, bisexual; corolla narrowly campanulate, 2-3.1 mm long, 0.2-0.3 mm wide near base, 0.5-1 mm wide near apex, white, resin globules present throughout; lobes 5, 0.25-0.3 mm long; style undivided; ovary vestigial; pappus bristles 2-6, 1.9-2.6 mm long, barbellate, slightly flattened, often bent or twisted near apex. Heterogamous capitula: Capitula globose at anthesis, 2.5-3 mm long, 2.5-3 mm wide. Involucral bracts incurved, glabrous; outer ones much shorter than inner; median bracts 2-2.7 x 1-1.2 mm, apex obtuse, glabrous, margin entire or denticulate. Receptacle hemispherical. Marginal florets 100-200, in several rows, female; corolla filiform, 0.8-1.3 mm long, white, cylindrical, multicellular hairs absent, few to numerous resin globules present; lobes 3, tiny; styles slender, branched. Disc florets 2-16, bisexual; corolla narrowly campanulate, 1.2-1.5 mm long, 0.2-0.3 mm wide near base, c. 0.5 mm wide near apex, white, resin globules present throughout; lobes 5, 0.25-0.3 mm long; style undivided; ovary vestigial; pappus bristles 2-6, 0.8-1.1 mm long, barbellate, slightly flattened, often bent or twisted near apex. Achenes formed only from marginal florets, lunate, 0.6-0.7 mm long, c. 0.2 mm wide, with 1 conspicuous longitudinal rib, and several raised anastomosing veins sometimes visible; twin hairs absent; resin globules absent. Pappus corona obliquely placed, erect, cylindrical, white, 0.03-0.05 mm high, margin entire; pappus bristles absent. Fig. 4F-J. Additional selected specimens examined: Australia: South Australia. Katarapko Reserve, N of Loxton, Jul 1985, Alcock 10249 (AD, MEL, NSW, NY); 7 km W of Mundondna HS, May 1984, Badman 1022 (AD, CANB, MEL, NSW); Waukatanna waterhole. Cooper Creek, Apr 1985, Badman 1646 (AD, BRI, MEL); 50 km SE of Coongie, Lake Eyre basin, Oct 1986, Conrick 1939 (AD, BRI, NSW); Pandiburra Bore, Aug 1975, Donner 5177 (AD, US); Neals River, 56 km S of Oodnadatta on track to William Creek, Nov 1989, Nordenstam & Anderberg 972 (AD, MEL, NSW, PERTH); Engenina Creek, on the Coober Pedy - William Creek Road, Sep 1990, Wilson 789 & Rowe (AD, MEL, NSW). Queensland. Gregory North District: Cuttaburra Channels, Eyre Development Road, S of Bedourie, May 2007, Bean 51 26385 (BRI, MEL); 116 km N of Birdsville, Sep 1989, Cowan 81 & Bushel1 (BRI). Warrego District: c. 15 km W of Quilpie, Nov 1975, Henderson H2384 (BRI). Gregory South District: Thylungra, c. 75 miles [c. 121 km] NW of Quilpie, Oct 1955, Everist 5768 (BRI); 1.3 km S of Plevna Downs HS gate, W of Eromanga, Aug 2010, Bean 30103 (BRI); Lake Yamma Yamma, Sep 1991, Mitchell 824 (BRI); Windorah, ‘Tanbar’, Cooper Creek, Nov 1990, Emerson s.n. (BRI [AQ 501325]). New South Wales. Yantara Lake, Oct 1887, Baeuerlen s.n. (NSW); 15.1 km SW of Goodooga, Dec 1998, Bean 14508 (BRI, NSW); Near Barrier Range, Feb 1861, Beckler s.n. (MEL); Delalah Downs, Aug 1972, Cunningham 476 (BRI, NSW); Willandra Creek, c. 28 miles [c. 43 km] SE of Ivanhoe, Nov 1973, Salasoo 5427 (NSW). Victoria. Kings Billabong Wildlife Reserve, near Red Cliffs, Aug 1975, Beauglehole 50256 (AD, MEL); Kings Billabong, Red Cliffs area. May 1969, Henshall s.n. (MEL); W of Neds Corner HS, Mullaroo Creek, 5.5 km due N of Lake Wallawalla, Lindsay Island, Dec 1981, Browne 73 (MEL); Lake Lalbert, Dec 1853, Mueller s.n. (MEL). Distribution and habitat : Ethuliopsis cunninghamii is widely distributed in south-western Queensland, western New South Wales, eastern South Australia and north-western Victoria (Map 8), in areas that receive less than about 600 mm annual rainfall. It grows in heavy clay soil on alluvial flats, and around ephemeral swamps and lakes. Associated species include lignum {Muehlenbeckia flondenta Meisn.), gidgee (Acacia cambagei R.T.Baker), coolabah {Eucalyptus coolabah Blakely & Jacobs), and bluebush ( Chenopodium auricomum Lindl.). Phenology : Flowers and fruits have been recorded from March to December, but flowering and fruiting is most likely synchronised to rainfall events rather than season. Notes : Ethuliopsis is distinct from both Epaltes sens. str. (E. divaricata) and from Sphaeromorphaea , and would appear to be more closely related to some members of the large genus Pluchea. The latter is a very complex and possibly paraphyletic assemblage (Anderberg et al. 2005), and none of the Pluchea species in Australia or south¬ east Asia have characters similar to those exhibited by Ethuliopsis. Ethuliopsis cunninghamii is sub- dioecious. Some plants have homogamous capitula that bear only disc florets. These florets are bisexual, but do not produce 52 achenes and the ovary is vestigial, hence they are effectively male. The capitula on these plants are campanulate or cylindrical, the median involucral bracts are 3.1-3.5 mm long, and the corolla of the disc florets is 2-3.1 mm long. Other plants have heterogamous capitula that bear predominantly marginal florets, with a few disc florets in the centre. The capitula on those plants are globose, the median involucral bracts are 2-2.7 mm long, and the corolla of the disc florets is 1.2-1.5 mm long. Hence, specimens of the “male” and “female” plants can be readily distinguished by capitulum shape and size, or by floret size. The corolla of both marginal and disc florets is white, unlike Sphaeromorphaea Austrobaileya 9(1): 30-59 (2013) spp. (except marginal florets of S. major) and Epaltes divaricata which are pink to purple. There is one herbarium specimen reportedly collected from the Northern Territory. It is at MEL, and has a handwritten label written in ink stating “118. Epaltes cunninghamii B., 1885/6, Lieut. Dittrich, Lindsay’s Exped.” Someone else has later written, in pencil, “finke river”. As this species has never otherwise been recorded from the Northern Territory, it seems likely that the locality is in error, and that the specimen was in fact collected in South Australia. Conservation status : A common and widespread species. Key to the Australian genera of Subtribe Plucheinae 1 Capitula grouped into secondary heads or glomerules.2 1. Capitula all separate.3 2 Pappus absent.Sphaeranthus 2. Pappus present.Pterocaulon 3 Pappus of the marginal florets consisting of a small corona, sometimes with 1-5 bristles arising from it.4 3. Pappus of all florets with 10 or more bristles, corona absent.5 4 Achenes cylindrical, with 5-14 longitudinal ribs; capitula all heterogamous, hemispherical to oblate; corolla pink to purple, rarely white.Sphaeromorphaea 4. Achenes lunate, with 1 longitudinal rib; capitula heterogamous or homogamous, campanulate or globose; corolla white.Ethuliopsis 5 Pappus bristles fused into a tube at their base.Coleocoma 5. Pappus bristles all free.6 6 Pappus bristles broad-based, persistent, ebarbellate.Thespidium 6. Pappus bristles slender throughout, somewhat caducous, barbellate.7 7 Disc florets 3-merous; anthers not tailed.Allopterigeron 7. Disc florets 4-merous or 5-merous; anthers tailed.8 8 Corolla of marginal florets irregularly lobed to distinctly ligulate; achenes densely sericeous.Streptoglossa 8. Corolla lobes of marginal florets equal; achenes glabrous or sparsely hairy.Pluchea 53 Bean, Sphaeromorphaea and Ethuliopsis Excluded names in Epaltes and Sph aeromorph aea Epaltes alata (Sond.) Steetz in Peters, Naturw. Reise Mossambique [Peters] 6(Bot., 2): 452 (1864) = Litogyne gariepina (DC.) Anderb. Epaltes gariepina (DC.) Steetz in Peters, Naturw. Reise Mossambique [Peters] 6(Bot., 2): 451 (1864) = Litogyne gariepina (DC.) Anderb. Epaltespleiochaeta F.Muell., Fragm. 10: 100 (1877) = Pluchea baccharoides (F.Muell.) Benth. Epaltes tatei F.Muell., Trans. & Proc. Roy. Soc. South Australia 6: 31 (1883) = Haegiela tatei (F.Muell.) P.S.Short & Paul G.Wilson. Sphaeromorphaea centipeda DC., Prodr. 6: 140 (1838). An illegitimate name based on Artemisia minima L. (= Centipeda minima). Centipeda sect. Sphaeromorphaea C.B.Clarke, Compos. Ind. 151 (1876) is typified by Centipeda orbicularis nom. illeg. (= Centipeda minima). This sectional name is therefore a synonym of Centipeda. Sphaeromorphaea russeliana var. glabrata DC., Prodr. 6: 140 (1838). Type: NW India, J.F. Royle s.n. (syn: K 000373336, image!) = Centipeda minima subsp. minima. A Royle specimen at G-DC [G00318749] identified as var. glabrata is evidently from a different gathering. It bears the number 260, and the locality Cachemire [?], rather than the ‘prov. bor.-occid.’ as stated in the protologue. This latter specimen is not considered a type. It represents Sphaeromorphaea littoralis. Acknowledgements I thank the Directors of A, AD, AMES, BM, C, CANB, GH, K, LD, MEL, NOU, PE, PERTH and TAI for the loan of specimens; the Directors of CAL and M for sending images of specimens, V. Fonjallaz (G) for sending images of type specimens, J. Hunnex (BM) for sending images of historical specimens; and lb Friis and Hans Hansen (C) for assistance with the type of Artemisia littoralis. I am grateful to Philip Sharpe for translating some German text, and Jian Wang (BRI) for interpreting Chinese specimen labels; to Keith McDonald and Mike Mathieson for collecting Sphaeromorphaea material from Queensland. I also thank Melinda Peters (HUH) and Nicola Biggs (K) for friendly and efficient assistance; Murray Haseler and Steve Heggie for enabling access to Edgbaston Reserve; Marge Scully of the Cooktown Historical Society for providing information about C.C. Harris; and Dr Barbara Hawgood for kindly sending a copy of her article on Patrick Russell. Peter Bostock (BRI) kindly provided the Latin diagnoses, and Will Smith (BRI) provided the illustrations and put the finishing touches on the distribution maps. References Anderberg, A. A. (1991). Taxonomy and phylogeny of the tribe Plucheeae (Asteraceae). Plant Systematics and Evolution 176: 145-177. Anderberg, A.A., Eldenas, P., Bayer, R.J. & Englund, M. (2005). Evolutionary relationships in the Asteraceae tribe Inuleae (incl. Plucheeae) evidenced by DNA sequences of ndhF; with notes on the systematic positions of some aberrant genera. Organisms, Diversity & Evolution 5: 135-146. Anderberg, A.A. & Eldenas, P. (2007). Tribe Inuleae. In: K. 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Recent Researches in Plant Anatomy and Morphology , pp. 275-294. Scientific Publishers: Jodhpur, India. Naithani, B.D. (1995). Tribe Anthemideae. In Flora of India 12: 3-75. Botanical Survey of India: Calcutta. National Library Of Australia (2009-onwards). TROVE, http://trove.nla.gov.au/ Accessed November 2012. Peng, C. (2004). Digital Flora of Taiwan. http://www. efloras.org/flora_page.aspx?flora_id=100 Accessed 29 January 2012. Retief, E. & Herman, PP J. (1997). Plants of the northern provinces of South Africa: keys and diagnostic characters. Strelitzia 6: 1-681. Short, P.S. & Wilson, P.G. (1990). Haegiela , a new genus of Australian Asteraceae (Inuleae: Gnaphaliinae), with notes on the genus Epaltes Cass. Muelleria 7: 259-265. 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, May 2011. Northern Territory Herbarium: Department of Natural Resources, Environment, The Arts and Sport, http://www.nt.gov.au/nreta/wildlife/ plants_herbarium/pdf/200701nt_checklist.pdf Accessed 3 March 2012. Sonder, O.W. (1853). Compositae. In Plantae Muellerianae, Beitrag zur Flora Sudaustraliens, aus den Sammlungen des Dr. Ferd. Muller. Linnaea 25: 450-656. Soons, M.B., Vlugt, C. Van Der, Lith, B. Van, Heil, G.W. & Klaassen, M. (2008). Small seed size increases the potential for dispersal of wetland plants by ducks. Journal of Ecology 96: 619— 627. Stephens, K. & Sharp, D. (2009). Flora of North Stradbroke Island. Queensland Herbarium: Brisbane. Vallance, T.G., Moore, D.T. & Groves, E.W. (2001). Nature’s Investigator, The Diary of Robert Brown in Australia. Australian Biological Resources Study: Canberra. Wallich, N. (1831). Numerical List of Dried Specimens of plants in the Muse um of the Honl. East India Company , numbers 2604-4877. East India Company: London. Walsh, N.G. (2001). A revision of Centipeda (Asteraceae). Muelleria 15: 33-64. Walsh, N.G. & Stajsic, V. (2007). A Census of the Vascular Plants of Victoria , 8 th edition. Royal Botanic Gardens, Melbourne, http://www.rbg. vic.gov.au/viclist/ Accessed 3 March 2012. 56 Austrobaileya 9(1): 30-59 (2013) Map 2. Australian distribution (native and naturalised) of Sphaeromorphaea australis. Bean, Sphaeromorphaea and Ethuliopsis 57 Map 3. World distribution (native and naturalised) of Sphaeromorphaea australis. Map 4. World distribution of Sphaeromorphaea harrisii. 58 Austrobaileya 9(1): 30-59 (2013) Map 6. World distribution (native and naturalised) of Sphaeromorphaea littoraJis. Bean, Sphaeromorphaea and Ethuliopsis 59 160°E Map 7. World distribution of Sphaeromorphaea subintegra. 25°S 30°S 35°S 150°E Map 8. Distribution of Ethuliopsis cunninghamii. Reinstatement of intraspecific taxa for Bosistoapentacocca (F.Muell.) Baill. (Rutaceae) with a new combination B. pentacocca subsp. connaricarpa (Domin) P.I.Forst. Paul I. Forster Summary Forster, P.I. (2013). Reinstatement of intraspecific taxa for Bosistoa pentacocca (F.Muell.) Baill. (Rutaceae) with a new combination B. pentacocca subsp. connaricarpa (Domin) RI.Forst. Austrobaileya 9(1): 60-65. Geographic variation in morphology of Bosistoa pentacocca is discussed and the recognition of subspecies is proposed with the new combination B. pentacocca subsp. connaricarpa (Domin) RI.Forst. Typification of the name Bosistoa connaricarpa Domin is reviewed with the holotype designated as being at K. Key Words: Rutaceae, Bosistoa , Bosistoa pentacocca subsp. connaricarpa , Australia flora, Queensland flora, identification key, taxonomy, subspecies, John Dallachy P.I.Forster, Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: Paul. Forster@science. dsitia.qld.gov. au Introduction Bosistoa pentacocca (F.Muell.) Baill. as defined by Hartley (2013), is a variable species from eastern Australia with numerous geographically discrete population centres (Map 1). This revised his initial analysis of these same populations as comprising one species with three varieties, with each variety based on discrete population centres and defined on leaf morphology (Hartley 1977). In this instance Hartley’s use of variety was equivalent to that of subspecies (sensu Forster 2005; Duretto & Forster 2007) wherein the relevant taxon is confined to a discrete geographic region and is morphologically distinct. Hartley’s (2013) more conservative taxonomic conclusion was based on the greater number of collections now available and that the previously perceived differences between taxa were no longer valid. Diverse chemical evidence also tends to support a single species, albeit with some variation between populations (Auzi et al. 1997; Brophy et al. 2007), although both these studies sampled only a small number of populations. Accepted for publication 26 August 2013 Notwithstanding Hartley’s (2013) conclusion, there remain clear morphological differences between the most southern and the other population centres, principally the number of leaflets in each leaf and the number of secondary (lateral) veins per side of the primary (midrib) vein in each leaflet. These differences are easily discernible both in the field and on herbarium material and persist in material of known origin cultivated in a common garden. Although occasional collections exhibit divergence towards the other morphotype, the overall differences are consistent within a given geographic area (Map 1). My conclusion is that two intraspecific taxa can still be recognised within Bosistoa pentacocca (maintaining the status quo of Forster 1997, 2002, 2007, 2010; Harden et al. 2006); however, I prefer the rank of subspecies rather than variety as the latter taxonomic category is more suited to within population variation. The existence of some populations (see below) exhibiting intermediate combinations of characters are to be expected and demonstrate continuity as required by Stebbins (1950), viz. “subspecies....connected....by a series of intergrading forms”. Forster, Bosistoa pentacocca If a metapopulation concept (Hanski & Gilpin 1991; Freckleton & Watkinson 2002) is applied to Bosistoa pentacocca then it can be hypothesised that the species consists of at least six, geographically discrete, metapopulations with B. pentacocca subsp. pentacocca existing as a single metapopulation (1 on Map 1). Each of the metapopulations comprise few to many populations or subpopulations in ‘core’, possibly refugial areas, with occasional outlying populations/subpopulations. Applying this hypothesis, the ‘core’ area for B. pentacocca subsp. pentacocca exists within the Border Ranges (McPherson/Macleay overlap) centre of plant species richness and endemism of southeast Queensland and northeast New South with a concentration of populations on the basalt flows associated with the Mt Warning caldera and an ‘outlier’ population at Mt French, Boonah. Five metapopulations can be hypothesised for B. pentacocca subsp. connaricarpa (viz. 2. Mt Dryander - Conway Range area; 3. Eungella - Kuttabul area; 4: Rockhampton area; 5. Bulburin - Miriam Vale area; 6. Mt Urah - Cold Creek area (Map 1). These proposed metapopulations remain to be tested using molecular data, although it is perhaps likely that most are refugial in a genetic sense (i.e. with little existence of active gene flow apart from within catchments). 61 Materials and methods Populations of Bosistoa pentacocca throughout its known range in Queensland were examined in the field. Herbarium collections at BRI and CANB were reviewed, plus online or scanned images of selected specimens at K and MEL. Common abbreviations in the specimen citation text include NP (National Park), SF (State Forest) and TR (Timber Reserve). Taxonomy Bosistoa pentacocca (F.Muell.) Baill., Hist. PI. 4: 470 (1873); Euodiapentacocca F.Muell., Fragm. 3: 41 (1862); B. sapindiformis F.Muell. ex Benth., FI. Austral. 1: 359 (1863), nom. illeg.', Acradenia bosistoi F.Muell., Pap. & Proc. Roy. Soc. Tasmania 8: 8 (1868) & Fragm. 6 167 (1868), nom. illeg. Type: New South Wales. Clarence River, s.dat., H. Beckler s.n. (holo: MEL 67499, online image!; iso: K 000717477, online image!, P 04355028, online image!). The species description is as per Hartley (2013) and two subspecies are recognised here. The specimens cited are representative of the six proposed metapopulations (numbered as above). Key to the subspecies of Bosistoa pentacocca 1 Leaves with 3-5(-6) leaflet pairs; leaflets with 16-30 secondary (lateral) veins per side of primary (midrib) vein.B. pentacocca subsp. pentacocca 1. Leaves with 1 or 2 (rarely 3) leaflet pairs; leaflets with 13—16(—18) secondary (lateral) veins per side of primary (midrib) vein. Bosistoa pentacocca subsp. pentacocca B. pentacocca var. pentacocca Hartley, J. Arnold Arbor. 58: 423 (1977). Leaves imparipinnate, 14-45 cm long, with 3-5 (rarely 6) leaflet pairs; terminal leaflet usually oblanceolate, more rarely elliptic- oblong, subterminal leaflets oblanceolate, elliptic-oblong to narrowly elliptic, margins of all leaflets crenate to coarsely serrate, rarely entire, with 16-30 secondary (lateral) veins per side of primary (midrib) vein. .B. pentacocca subsp. connaricarpa Selected specimens examined'. Queensland. Moreton District: Ormeau, corner of Barrenjoey & Upper Ormeau Roads, Jan 1995, Forster PIF16086 & Leiper (BRI); NW slopes of Mt French, Apr 1982, Bird s.n. (BRI [AQ339215]); Mt French, NW slopes, 7.8 km WNW of Boonah, Jan 1983, Guymer 1847A & Jessup (BRI, CANB, MEL, NSW); Tambourine Mt, in Guanaba Gorge, Mar 1947, Blake 17387 (BRI); Numinbah Forest Reserve, c. 15 km SW of Nerang, southern slope of Pages Pinnacle, Jan 2006, Halford Q8958 (BRI); Upper Currumbin Creek, Tomewin Range, Oct 1981, Williams 81250 (BRI); Nicholl Scrub NP, Currumbin Valley, Feb 2011, Forster PIF37846 & Fernando (BISH, BRI, MEL); Egg Rock/Prison Farm track Nixons Creek, Lamington 62 NP, Nov 1993, Grimshaw G168 (BRI); Cave Creek, Natural Bridge [Springbrook] NP, Numinbah Valley, Mar 1978, Field s.n. (BRI [AQ264150], New South Wales. North Coast: Booyong Recreation Reserve, Big Scrub Remnant, Feb 1987, Floyd s.n. (BRI [AQ495572]). Distribution and habitat : Bosistoa pentacocca subsp. pentacocca is endemic to southeast Queensland (northern limit at Ormeau) and northeast New South Wales (south to the Clarence River) (Map 1). A western, outlying population occurs at Mt French, Boonah. It is found in rainforest (araucarian notophyll to microphyll vineforest) on diverse substrates (basalts, metasediments) predominantly on alluvium and colluvium along ephemeral watercourses. Notes : The most commonly encountered morphological state for this subspecies has plants with leaves with 4 or 5 leaflet pairs and predominantly oblanceolate leaflets that are always toothed. Occasionally the leaflets are more or less entire, but they are usually mixed in with toothed leaflets on the same branch. Fertile branchlets generally have leaflets at the larger end of the recorded dimensions whereas in Bosistoa pentacocca subsp. connaricarpa , the leaflets are usually much smaller and tend to be elliptic in shape. Bosistoa pentacocca subsp. connaricarpa (Domin) P.I.Forst., comb, et stat. nov.; B. connaricarpa Domin, Feddes Repert. Spec. Nov. Regni Veg. 12: 390 (1913); B. pentacocca var. connaricarpa (Domin) T.G.Hartley, J. Arnold Arbor. 58: 424 (1977). Type citation: “Queensland: Rockhampton, leg. Dallachy”. Type: Queensland. Port Curtis District: Rockhampton, 12 December [1862], [J. Dallachy s.n.] (holo: K, image!). Bosistoa pentacocca var. dryanderensis T.G.Hartley, J Arnold Arbor. 58: 425 (1977). Type: Queensland. North Kennedy District: Mt Dryander, 11 August 1972, V.K. Moriarty 968 (holo: CANB; iso: BRI). Leaves imparipinnate or pinnately trifoliate, 9-30 cm long, with 1 or 2, rarely 3 leaflet pairs; terminal leaflet usually elliptic, subterminal leaflets oblanceolate to elliptic or narrowly elliptic, margins of all leaflets entire, or crenate to coarsely serrate, with 13—16(—18) secondary (lateral) veins per side Austrobaileya 9(1): 60-65 (2013) of primary (midrib) vein. Additional selected specimens examined: Queensland. North Kennedy District: 2 *Portello’s Crossing, Dryander Creek, base of Mt Dryander, Feb 1994, Forster PIF14859 & Bean (A, BRI, CNS, MEL); 2 Sawmill Beach, Whitsunday Island [Whitsunday Islands NP], Sep 1990, Batianoff9009838 & Batianoff (BRI; BISH, LAE o.v.); impulse Creek, SF299 Conway, May 1991, Forster PIF8300 & McDonald (BRI, CNS, K, MEL). South Kennedy District: 3 Mt Blackwood, S side c. 5 km from Kuttabul, Jun 1984, Champion 68 (BRI); 3 Mt Adder, southern base. Mount Jukes NP, Apr 1991, Forster PIF8115 & McDonald (BRI, CANB, CNS, K, L); 3 Camerons Pocket, 12 km W of Calen [Mia Mia SF], Jan 2003, Hambleton 4 (BRI); 3 23 km S of Eungella Dam Road on Cockies Creek Road, Crediton SF, Sep 2002, Gumming 21133 (BRI). Port Curtis District: 4 Black Creek, SW base of Mt Sleipner, Mar 1989, Forster P1F4999 & Hoy (BRI, CANB, CNS, K, L, MEL, MO, NSW, US); 5 TR115 [now Castle Tower NP], Mt Coulston, 9 km NNW of Bororen, Sep 1989, Gibson TOI828 (BRI); 5 Bindaree, NE of Miriam Vale & just S of Arthurs Seat, Jun 1995, Meter RM911 (BRI); 5 TR122, Wietalaba [now Wietalaba NP], 32 km S of Calliope, Oct 1989, Gibson TO1740 (BRI, CANB); 5 TR202 [now Bulburin NP], Feb 1995, Forster P1F16176 (BRI, CNS, MEL); 5 2.5 km N of Glassford Vale, TR353 [now Dawes NP], Sep 1989, Forster PIF5674 & Bean (BRI, CANB); 5 Bulburin NP, Pine Creek headwaters, Jan 2009, Forster PIF34931 6 Brushe (BRI, MEL); 5 Bulburin NP, Cassilus Creek, Jan 2009, Forster P1F34944 et al. (BRI, CNS). Burnett District: 5 SF 695 Kalpowar, Mt Fort William, Mar 2000, Forster PIF25439 & Booth (BRI, CNS). Wide Bay District: 6 SF50 Glenbar, 1 km WSW of Mt Urah summit [now Glenbar NP], Feb 1993, Forster PIF 13130 & Machin (BRI); 6 Glenbar NP, Cold Creek catchment, 8 km NW of Miva, Dec 2008, Forster PIF34782 et al. (BRI). * superscript number refers to the metapopulation in Map 1 Distribution and habitat : Bosistoa pentacocca subsp. connaricarpa occurs in eastern Queensland in five metapopulations from Mt Dryander to northeast of Gympie (Map 1). It invariably occurs in rainforest (microphyll to notophyll vineforest, often with Araucaria cunninghamii Aiton ex A.Cunn.) on diverse substrates (andesites, basalts, granites, granodiorites), often on alluvium or colluvium along ephemeral watercourses. The type of B. pentacocca subsp. connaricarpa is from ‘Rockhampton’; and the subspecies has only been recollected from the Mt Sleipner locality in this general area, thus this proposed metapopulation (4 on Map 1) consists of a single population. By contrast the other metapopulations are more diverse (2 has at least seven populations, 3 has at least Forster, Bosistoa pentacocca four populations, 5 has at least 13 populations and 6 has at least two populations). Typification : Domin (1913) cited a single collection in the protologue for Bosistoa connaricarpa, viz. “Queensland: Rockhampton, leg. Dallachy”. Earlier he had stated “I give descriptions of new forms according to specimens preserved in the herbarium of the Royal Botanic Gardens at Kew” when basing names on collections other than his own from Australia (Domin 1911). It can be concluded from this that a holotype for this name should be at Kew. Hartley (2013) gives the type of this taxon as “Rockhampton, Qld, 5 Feb. 1863, J.Dallachy, iso: MEL”. Earlier he stated “Type: Dallachy, February 5,1863, Queensland, Rockhampton” with an isotype at MEL (Hartley 1977). There are two sheets at MEL (67497 & 67503) of this collection, both labelled as an isotype by Hartley in January 1977. Both of these sheets have original collection labels from Dallachy with his collection number #297 and the locality of “Devils Creek”, but no mention of Rockhampton. There exists atK a sheet with a ‘ Phytologic Museum of Melbourne’ label stating “Bosistoa sapindiformis F.M. Rockhampton” (although the specimen can now be clearly identified as Bosistoa pentacocca subsp. connaricarpa). The handwriting on this label is Mueller’s. Whilst there is no mention of a collector or specific date on this label, further evidence can be found in a field label attached to the bottom left corner of the sheet. This field label is written in Dallachy’s hand using blue crayon and amongst various points on plant habit, states “12 Dec”. Dallachy is known to have made collections in the Rockhampton area “at least from the 5 th [of] January 1863” to “at least April 16 th ” (Blake 1954) and this field label indicates that he was active there three weeks earlier. Prior to the Rockhampton area, Dallachy collected about Brisbane (Blake 1954). It is unlikely that Domin saw any of the MEL sheets discussed and although labelled as isotypes by Hartley they are not duplicates of the K material as they were collected on 63 a different date. Mueller distributed material sent to him by collectors to many herbaria, often with inadequate or different labels to that of those that were retained in MEL. Collectors such as Dallachy also often recollected material and it can be difficult to ascertain the specific collections subsequently cited by Mueller and others as they were generally unmounted in the 1800s and often with multiple sheets. Precisely where Dallachy made his “Rockhampton” collection can probably never be resolved; however, it is likely that he returned to the same vicinity on subsequent occasions and so the collections by him of this plant in K and MEL may be from the same spot. His locality “Devils Creek” is possibly equivalent to Frenchmans Creek in a modern context for the Rockhampton area as this ephemeral watercourse flows into the Fitzroy River at Devils Elbow. The known locality for Bosistoa pentacocca at the southwestern base of Mt Sleipner is on the eastern side of Black Creek that is east of Mt Berserker and only about 3 km in a direct line from where Dallachy may have collected his specimens. Suitable habitat still exists in the upper reaches of Frenchmans Creek and should be explored to determine if B. pentacocca populations persist. Notes : Several populations of this subspecies occasionally exhibit characteristics of B. pentacocca subsp. pentacocca. These are at Kalpowar, Bulburin and Mt Urah (all at the southern end of the northern subspecies range) where the leaves can occasionally have three leaflet pairs; however, these are mixed in with leaves predominantly of 1 or 2 leaflet pairs. Acknowledgements I am very grateful to David Goyder (Royal Botanic Gardens Kew) for locating the Dallachy material in the general herbarium and Lucia Lopez Poveda for photographing material. Elucidation of Dallachy’s handwriting (scrawl) was assisted by Peter Bostock, Gordon Guymer and Will Smith (BRI). The population of Bosistoa pentacocca at Black Creek was located with 64 the assistance of Dianne & Neil Hoy in 1989. Useful discussions of possible localities for Dallachy’s ‘Devils Creek’ were held with Neil Hoy and Susan Cunningham. Map 1 was prepared by Will Smith. References Auzi, A.A., Forster, P.I., Gray, A.I., Hartley, T.G., Waigh, R.D. & Waterman, P.G. (1997). Phloroglucinol derivatives from leaves of Bosistoapentacocca. Phytochemistry 45: 1673— 1678. Blake, S.T. (1954). Some pioneers in plant exploration and classification. Proceedings of the Royal Society of Queensland. 66: 1-19. Brophy, J.J., Goldsack, R.J. & Forster, PI. (2007). Essential oils from the leaves of Bosistoa F.Muell. ex Benth. (Rutaceae). Journal of Essential Oil Research 19: 249-254. Domin, K. (1911). First contribution to the Flora of Australia. Feddes Repertorium Specierum Novarum Regni Vegetabilis 9: 550-553. - (1913). Eighth contribution to the Flora of Australia. Feddes Repertorium Specierum Novarum Regni Vegetabilis 12: 388-390. Duretto, M.F. & Forster, PI. (2007). A taxonomic revision of the genus Zieria Sm. (Rutaceae) in Queensland. Austrobaileya 7: 473-544. Forster, PI. (1997). Rutaceae. In R. J.F. Henderson (ed.), Queensland Plants - Names and Distribution , 184-188. State of Queensland, Department of Environment: Brisbane. - (2002). Rutaceae. In R.J.F. Henderson (ed). Names and Distribution of Queensland Plants, Algae and Lichens , pp. 177-181. Queensland Government, Environmental Protection Agency: Brisbane. - (2005). New species of Philotheca Rudge (Rutaceae) from Queensland. Austrobaileya 7: 175-181. Austrobaileya 9(1): 60-65 (2013) Freckleton, R.P & Watkinson, A.R. (2002). Large-scale spatial dynamics of plants: metapopulations, regional ensembles and patchy populations. Journal of Ecology 90: 419-434. Hanski, I. & Gilpin, M. (1991). Metapopulation dynamics: brief history and conceptual domain. Biological Journal of the Linnean Society 42: 3-16. Harden, G., McDonald, W.J.F. & Williams, J. (2006). Rainforest Trees and Shrubs. A field guide to their identification. Gwen Harden Publishing: Nambucca Heads. Hartley, T.G. (1977). A revision of the genus Bosistoa (Rutaceae). Journal of the Arnold Arboretum 58: 416-436. - (2013). Bosistoa. In A.J.G. Wilson (ed.). Flora of Australia 26: 48-52. ABRS/CSIRO Publishing: Melbourne. Stebbins, G.L. (1950). Variation and Evolution in Plants. Columbia University Press: New York. - (2007). Rutaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2007 ; pp. 180-183. Queensland Herbarium, Environmental Protection Agency: Brisbane. - (2010). Rutaceae. In P.D. Bostock & A.E. Holland (eds.), Census of the Queensland Flora 2010 , pp. 174-178. Queensland Herbarium, Department of Environment and Resource Management: Brisbane. Forster, Bosistoa pentacocca 65 Map 1 . Distribution of Bosistoa pentacocca in eastern Australia. Numbered areas refer to proposed metapopulations (1 -B.pentacocca subsp. pentacocca A, outlier population at Mt French arrowed; 2-6 -B. pentacocca subsp. connaricarpa •. Three new species of Pluchea Cass. (Asteraceae: Inuleae-Plucheinae) from northern Australia A.R. Bean Summary Bean, A.R. (2013). Three new species of Pluchea Cass. (Asteraceae: Inuleae-Plucheinae) from northern Australia. Austrobaileya 9(1): 66-74. Three new species of Pluchea Cass, are described; P. longiseta A.R.Bean from northern Western Australia, P. mesotes A.R.Bean from the Northern Territory, and P. alata A.R.Bean from central Queensland. All species are illustrated and distribution maps are provided. A revised key to the Australian of Pluchea species is provided. Key Words: Asteraceae, Pluchea , Pluchea alata , Pluchea longiseta , Pluchea mesotes , Australia flora, taxonomy, identification key A.R. Bean, Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: Tony.Bean@science.dsitia.qld.gov.au Introduction Pluchea Cass, was revised for Australia by Hunger (1997), with seven species. Bean (2011) named two additional species of Pluchea from Queensland. Further investigations have revealed three additional species, which are named here: P. alata is endemic to the Aramac area of central Queensland, P. longiseta occurs in coastal and near-coastal parts of north-western Western Australia, and P. mesotes occurs in coastal parts of north¬ eastern Northern Territory. Materials and methods This study is based on herbarium specimens from BRI, DNA, MEL and PERTH, and (for P. alata) field investigations. Measurements of florets were made from material preserved in spirit or reconstituted in boiling water, while leaves, bracts and achenes were measured from dried specimens. Common abbreviations in the text include NT (Northern Territory), Qld (Queensland), SA (South Australia), WA (Western Australia). Accepted for publication 25 July 2013 Taxonomy Pluchea alata A.R.Bean sp. nov. affinis P. rubelliflorae (F.Muell.) B.L.Rob. sed capitulo cylindrico sessili vel subsessili, flosculis marginalibus prope horum apices expansis, setis pappi in flosculis disci paucioribus et alis caulis usque ad 1.2 mm latis differens. Typus: Queensland. Mitchell District: First Spring, Edgbaston Reserve, NE of Aramac, 6 April 2012, A.R. Bean 31641 (holo: BRI; iso: CANB, MEL). Woody shrub to 70 cm high and 120 cm across. Stems erect to spreading, well-branched, glabrous throughout, conspicuously winged, wings 0.5-1.2 mm wide. Leaves sessile, elliptical, spreading, slightly fleshy, decurrent, 15-30 x 4.2-7 mm, 3.5-4.5 times longer than broad, eglandular, glabrous; margins denticulate; apex acute. Capitula cylindrical, 5-6.5 x 2.5-3 mm, in terminal clusters of 2-4; peduncles 0-1 mm long. Involucral bracts in several rows; outer bracts ovate, 1.2-2 x 0.4-0.6 mm, with apex acute, outer surface glabrous, margins entire, not ciliate and glabrous; median bracts lanceolate, 2-3.5 x 0.3-0.6 mm, apex acute, outer surface glabrous, margins glabrous; inner bracts linear, 3.7-4 x 0.8- 0.9 mm, apex acute, outer surface glabrous, upper margins not ciliate, entire. Receptacle epaleate, verrucose, glabrous, flat, 1-1.2 Bean, Pluchea 67 Fig. 1. Pluchea alata. A. flowering branchlet xl.5. B. leaf showing decurrent base and winged stem *4. C. flowering capitulum xl2. D. disc floret xl6. E. outer floret x24. F. achene x48. A & B from Bean 31648 (BRI); C-E from Bean 31641 (BRI); F from Fairfax s.n. (BRI [AQ875934]). 68 mm diameter. Outer florets 55-75, female; corolla filiform, but expanded (0.2-0.25 mm wide) towards apex, 2.8—3.1 mm long, white; lobes 3 or 4, equal, triangular, 0.15-0.25 mm long, glabrous; style bifid, exserted, glabrous; pappus corona present, erect, cylindrical, c. 0.03 mm long; pappus bristles 4- 12, 2.4-2.8 mm long, caducous, in a single whorl, not coherent at base, all of similar length, barbellate throughout;. Disc florets 5- 13, ovary fully developed but infertile, pink; corolla cylindrical, 3-3.4 mm long, pink; lobes 5, triangular, 0.4-0.6 mm long, glabrous; anthers c. 1.5 mm long, including tail c. 0.15 mm long; style undivided, with obtuse sweeping hairs extending below the furcation;. Achenes developing only from outer florets; narrowly-ellipsoidal, 0.7-0.85 mm long, brown, glabrous; surface obscurely longitudinally striate; carpopodium prominent, white. Fig. 1. Additional specimens examined: Queensland. Mitchell District: Bowen Downs, in 1874, Birch s.n. (MEL); Big Spring, Edgbaston Station, N of Aramac, Apr 2005, Fairfax s.n. (BRI [AQ875934]); Edgbaston, NE of Aramac, Sep 2006, Fensham 5549 (BRI); Edgbaston Station 33 km NNE of Aramac, Apr 2010, Thompson MUT410 et al. (BRI, PE); Measuring Spring, Edgbaston Reserve, NE of Aramac, Apr 2012, Bean 31648 (BRI, NSW, US). Distribution and habitat: Pluchea alata is endemic to Queensland and is known from Edgbaston Reserve, NE of Aramac, and an old record from nearby Bowen Downs station (Map 1). It inhabits the margins of artesian springs, on saline soils. Phenology : Flowers and fruits have been recorded for April and September. Affinities: This species is allied to Pluchea rubelliflora , but differs by the woody stems, the relatively broader leaves (length 3.5-47 times width versus 4.5-12 times for P. rubelliflora ), the more or less cylindrical capitula that are sessile or with peduncles up to 1 mm long (hemispherical capitula on peduncles (1—)2—10 mm long for P. rubelliflora ); the outer florets expanded towards apex, with the lobes 0.15— 0.25 mm long (florets not expanded and with lobes 0.08-0.12 mm long for P. rubelliflora ); and the disc florets with 3-6 pappus bristles (6-18 bristles for P. rubelliflora). The stem Austrobaileya 9(1): 66-74 (2013) wings are up to 1.2 mm wide, whereas for P. rubelliflora in the Aramac - Barcaldine area, the stem wings are 0.2-0.4 mm wide. Note: ‘Bowen Downs’ was a much larger station when this species was collected there in 1874. In view of the habitat occupied by this species, Birch’s collecting locality was most likely in the Lake Huffer area (R. Fensham, pers. comm. Feb 2013). Conservation status: The species is known from fewer than 100 plants at the type locality. Applying the Red List criteria (IUCN 2012), a category of Endangered is recommended (Criterion D). Etymology: The epithet is from the Latin alatus meaning winged, in reference to the prominently winged stems of this species. Pluchea longiseta A.R.Bean sp. nov. ab omnibus aliis speciebus Plucheae foliis carnosis, flosculis achenisque longioribus et pappi setis in verticillis duobus tribusve dispositis differens. Typus: Western Australia. North shore of Willie Creek inlet, Dampierland Peninsula, 6 September 1985, K.F. Kenneally 9526A (holo: BRI; iso: CANB, DNA, PERTH). Pluchea sp. A; Wilson (1992: 950). Pluchea sp. B; Wilson (1992: 950). Pluchea sp. B Kimberley Flora (K.F. Kenneally 9526A); Western Australian Herbarium (1998-). Woody shrub to 60 cm high and 80 cm across. Stems terete, erect to spreading, well-branched, glabrous or with very sparse multicellular eglandular hairs near growing point. Leaves sessile, narrowly-oblanceolate to linear, spreading, fleshy, not decurrent, 28-52 x 2-5.5 mm, 9-14 times longer than broad, eglandular, multicellular hairs sparse or absent; margins entire; apex acuminate or apiculate. Capitula in small terminal clusters, narrowly campanulate to cylindrical, 12-15 x 8-12 mm; peduncles 2-27 mm long, with 1-3 lanceolate bracts near the apex, each 2-4 mm long. Involucral bracts in several rows; outer bracts ovate, 2.5-5.5 x 1.5-3 mm, apex acuminate, outer surface glabrous, margins Bean, Pluchea 69 entire and ciliate; median bracts lanceolate, 6.5-9.5 x 2.3-3 mm, apex acute to acuminate, outer surface glabrous, margins glabrous or with sparse eglandular hairs; inner bracts linear, 10.5-12 x 0.7-1.1 mm, apex laciniate, outer surface glabrous, upper margins glabrous, not ciliate. Receptacle epaleate, verrucose, glabrous, flat to concave, 1.7-2 mm diameter. Outer florets 100-200, female; corolla filiform, 8-9.5 mm long, pink; lobes 3, equal, triangular, 0.2-0.3 mm long, glabrous; style bifid, exserted, glabrous; pappus 8.2-9.2 mm long, pappus bristles 25-40, in 2 or 3 whorls, coherent at base, all of similar length. Disc florets 12-22, ovary fully developed but infertile, pale yellow; corolla cylindrical, 7.9- 9.3 mm long, pink; lobes 4(-5), triangular, 0.5-0.6 mm long, glandular; anthers c. 3 mm long, including tail c. 0.2 mm long; style bifid, with obtuse sweeping hairs extending below the furcation. Achenes narrowly-ellipsoidal, 1.3-1.5 mm long, brown, with numerous appressed twin hairs; longitudinal ribs 4-6; carpopodium prominent, white. Fig. 2. Additional specimens examined : Western Australia. Montgomery Island, E of Koolan Island, May 1998, Mitchell 5381 (DNA, PERTH); Cygnet Bay, Nov 1906, Fitzgerald 1706 (PERTH); Buckley Plain, north of town boundary, Broome, Feb 2005, Byrne 1374 (PERTH); Buckley Plain, 6 km NE of Broome, Nov 1987, Foulkes 442 (DNA); c. 9 km NE of Mandora Homestead, between Broome and Port Hedland, Aug 1997, Mitchell PRP1787 (PERTH); Loam 2 site, Mandora Marsh area, Oct 1999, Duero WEL2-10 et al (DNA, PERTH); NW Island, Montebello Islands, Oct 2000, Kenneally 11545 (PERTH); Hermite Island, Montebello Islands, Oct 2000, Kenneally 11573 (PERTH); below tip at side of DSI lease, Dampier Island, Sep 1987, Glennon 397K (PERTH); Boodie Island, S of Barrow and Middle Islands, Sep 2004, Long VL360-3 (PERTH); Duffers Bore, 24 km N of Dampier Downs, Aug 1976, Kenneally 5782 (DNA, PERTH); between T-bone Bay and hill towards lighthouse, Oct 1975, Weber 4949 (PERTH); Willie Creek marsh, Jan 1985, Foidkes 97 (PERTH); just N of Dragon Tree Soak, Great Sandy Desert, Aug 1977, George 14747 (BRI, PERTH); 80 Mile Beach, N of Wooroo Creek, Jul 1941, Burbidge 1300 (PERTH). Distribution and habitat : This species is endemic to Western Australia, mainly in coastal areas from the Montgomery Islands in the western Kimberley region, to the Cape Range, but also in a few scattered places away from the coast (Map 1). It inhabits samphire communities behind frontal dunes, saline marshy plains, and the margins of salty lagoons. Soils are usually pale and clayey but can have a sandy surface layer. Phenology : Flowers and fruits are recorded from May to February, with most collections made in September and October. Affinities : Because of the relatively large floral parts and the broad involucral bracts, this species resembles members of the genus Streptoglossa Steetz, and some specimens have previously been identified as such. But Streptoglossa species have ligulate outer florets and achenes with very dense twin hairs that conceal the surface. Pluchea longiseta has the tubular outer florets and almost glabrous achenes that are typical of this genus. Pluchea longiseta does differ from the usual characteristics of Pluchea in that the pappus bristles are arranged in two or three whorls. Pluchea usually has pappus bristles in a single whorl. P. longiseta can be distinguished from other Australian species of Pluchea also by the fleshy leaves, and the longer achenes, pappus and florets. Conservation status : This species is relatively widespread, but no surveys have been carried out to determine population sizes. Its habitat is potentially under threat from port development and infrastructure in some areas. A category of Data Deficient is appropriate under the IUCN guidelines (IUCN 2012). Etymology: The epithet longiseta refers to the pappus bristles that are longer in this species than in any other Australian Pluchea. Note: According to Western Australian Herbarium (1998-), the taxon “Pluchea sp. A” listed in Wilson (1992) is synonymous with “Pluchea sp. B”. 70 Austrobaileya 9(1): 66-74 (2013) Fig. 2. Pluchea longiseta. A. flowering branchlet xl. B. flowering capitulum x4. C. disc floret x8. D. outer floret x8. E. achene x32. A from Byrne 1374 (PERTH); B-D from Kenneally 9526A (PERTH); F from Kenneally 11573 (PERTH). Bean, Pluchea 71 Pluchea mesotes A.R.Bean sp. nov. affinis P. indicae (L.) Less, sed foliis angustioribus dentibus marginalibus paucis vel nullis, pedunculis capitulisque longioribus et pappi setis in quoque flosculo paucioribus differens. Typus: Northern Territory. Connexion Island, off Groote Eylandt, 29 September 1988, P.K. Latz 10901 (holo: BRI; iso: DNA). Multi-stemmed annual shrub to 100 cm high. Stems terete, erect to spreading, well- branched, with sparse to moderately dense multicellular eglandular hairs. Leaves sessile, narrowly-elliptical to spathulate, spreading, chartaceous, not resinous, glands absent or sparse, multicellular eglandular hairs sparse, bases not decurrent, 24-55 x 4-16 mm, 3.4- 6.2 times longer than broad, margins entire or occasionally with 1-3 pairs of teeth, apex acute. Capitula in small terminal clusters, narrowly campanulate, 5.5-7 x 5-5.5 mm; peduncles 0-7 mm long, with 1-3 lanceolate bracts near the apex, each 1-2.5 mm long. Involucral bracts in several rows; outer bracts broadly ovate, 1.6-2.7 x 1—1.4 mm, with apex acute, outer surface glabrous, upper margins ciliate; median bracts lanceolate, 3.2-3.8 x 0.8-1 mm, apex acute, outer surface glabrous, upper margins ciliate; inner bracts linear, 4.8-5.2 x 0.2-0.3 mm, upper margins ciliate (cilia 0.05-0.1 mm long), outer surface glabrous. Receptacle epaleate, verrucose, densely hairy, flat, 1.4-1.7 mm diameter. Outer florets 60-120, female; corolla filiform, 4-4.5 mm long, colour unknown; lobes 3, equal, triangular, 0.1-0.15 mm long, glabrous; style bifid, exserted, glabrous; pappus 3.8-4.1 mm long, pappus bristles 12-15, in a single whorl, coherent at base, all of similar length; Disc florets 2-5, ovary fully developed but infertile; corolla cylindrical, 4-4.3 mm long, white to pale yellow; lobes 5, triangular, 0.4- 0.7 mm long, glandular; anthers c. 1.9 mm long, including tail c. 0.3 mm long; style bifid, with obtuse sweeping hairs extending below the furcation. Achenes narrowly-ellipsoidal, 0.9-1 mm long, brown, with scattered appressed twin hairs; longitudinal ribs 2-5; carpopodium prominent, white. Fig. 3. Additional specimens examined : Northern Territory. Bennett Bay, Eastern Arnhem Land, Nov 1987, Russell- Smith 4219 & Lucas (BRI, DNA); Roper River, Sep 1867, Gulliver s.n. (MEL). Distribution and habitat: This species is apparently confined to the coastal parts of north-eastern Northern Territory (Map 1). It is recorded from “dune thicket”, and from “coral-shell gravel above mangroves”. Phenology: Flowers and fruits have been recorded for September and November. Affinities: Pluchea mesotes is related to P. indica but differs by the leaves 3.4-6.2 times longer than broad, that are without teeth or sometimes with a few small teeth (leaves 1.7-3.3 times longer than broad and with numerous prominent teeth for P. indica ); the peduncles 0-7 mm long (versus 0-1 mm long for P. indica}, the longer capitula 5.5-7 mm long (versus 4-5.5 mm forP. indica)', the cilia on the upper margins of the inner involucral bracts 0.05-0.1 mm long (versus 0.2-0.3 mm long for P. indica)', and the 12-15 pappus bristles (versus 17-25 for P. indica). P. indica occurs in the Darwin - Kakadu area, and on Cape York Peninsula, but apparently not in the region occupied by P. mesotes. Conservation status: A category of Data Deficient is appropriate under the IUCN guidelines (IUCN 2012). Etymology: From the Greek mesotes, meaning ‘in the middle position’. This is reference to the geographical position of this species in Australia compared with P. indica. The epithet is treated as a noun in apposition. Acknowledgements I am grateful to the Directors of DNA, MEF and PERTH for the loan of specimens, to Peter Bostock for the Fatin diagnoses and to Wifi Smith for the illustrations and distribution map. Thanks to Murray Haseler for organising access to Edgbaston Reserve, and Rod Fensham for elucidating Birch’s probable collecting locality for P. alata. 72 Austrobaileya 9(1): 66-74 (2013) Key to the Pluchea species in Australia 1 Leaves and stems glabrous (disregarding sessile glands) or with very few scattered hairs . 2 1. Leaves and stems with frequent to abundant eglandular hairs or stalked glandular hairs. . 8 2 Pappus bristles 8.2-9.2 mm long, in 2 or 3 whorls; leaves fleshy WA .P. longiseta 2. Pappus bristles 1.8-5 mm long, in a single whorl; leaves not fleshy.3 3 Leaves narrow, 4.5-23 times longer than wide.4 3. Leaves broad, 1.5-47 times longer than wide.6 4 Leaves with dark punctate glands; stems not winged.5 4. Leaves without punctate glands; stems narrowly winged NT, Qld, SA, WA. . P. rubelliflora 5 Capitula 6-7 mm long; disc florets 5-lobed; leaves 7-24 mm long; margins of involucral bracts fimbriate, eglandular NSW, Qld .P. baccharoides 5. Capitula 7-10 mm long; disc florets 4-lobed; leaves 18-54 mm long; margins of involucral bracts with stalked glands Qld .P. punctata 6 Stems winged; leaf glands absent Qld .P. alata 6. Stems not winged; leaf glands present.7 7 Leaves not resinous; capitula 4-5.5 mm long, clustered into sessile groups of (1—)2—4 NT, Qld .P. indica 7. Leaves resinous; capitula 6-8.5 mm long, usually solitary; some peduncles on any given plant more than 3 mm long WA .P. tetranthera 8 Hairs on leaves extremely dense, obscuring leaf surfaces NT, Qld, WA .P. ferdinandi-muelleri 8. Leaf surfaces readily visible.9 9 Hairs on leaves and stems predominantly gland tipped NSW, NT, Qld, SA, WA . . P. dentex 9. Hairs on leaves and stems all or mostly eglandular.10 10 Leaves narrowly-elliptic or oblanceolate, 3.4-6.4 times longer than wide.11 10. Leaves obovate, 1.5-3.3 times longer than wide.12 11 Leaf margins denticulate throughout; leaf surfaces with abundant, yellow glands; receptacle glabrous Qld .P. xanthina 11. Leaf margins entire or with 1 or 2 pairs of teeth; leaf surface glands absent or sparse, not yellow; receptacle hairy NT .P. mesotes 12 Capitula 5.5-75 mm long, not clustered, at least some peduncles more than 2 mm long; leaves often resinous NSW, NT, Qld, SA .P. dunlopii 12. Capitula 4-5.5 mm long, clustered into sessile groups of (l-)2-4; leaves not resinous NT, Qld .P. indica Bean, Pluchea 73 Fig. 3. Pluchea mesotes. A. flowering branchlet x0.5. B. leaf *2. C. flowering capitulum x6. D. disc floret xl6. E. achene x32. A, C-E from Latz 10901 (BRI); B from Russell-Smith 4219 & Lucas (DNA). 74 References Bean, A.R. (2011). Two new species of Pluchea Cass. (Asteraceae: Plucheinae ) from Queensland, Australia. Austrobaileya 8: 340-346. Hunger, S. (1997). A survey of the genus Pluchea (Compositae, Plucheeae) in Australia. Willdenowia 27: 207-223. Iucn (2012). IUCN Red List Categories and Criteria , Version 3.1, 2 nd edn. IUCN - The World Conservation Union: Gland. Austrobaileya 9(1): 66-74 (2013) Western Australian Herbarium (1998-). FloraBase — the Western Australian Flora. Department of Environment and Conservation, http:// florabase.dec.wa.gov.au/Accessed 17 Feb 2013. Wilson, A.J.G. (1992). Pluchea. In J.R. Wheeler (ed.). Flora of the Kimberley Region. Department of Conservation and Land Management: Como, Western Australia. Map 1 . Distribution of Pluchea longiseta •, P.mesotes A and P. alata M. Cryptocarya cercophylla W.E.Cooper (Lauraceae), a new species from Queensland’s Wet Tropics W.E. Cooper Summary Cooper, W.E. (2013). Cryptocarya cercophylla W.E.Cooper (Lauraceae), a new species from Queensland’s Wet Tropics. Austrobaileya 9(1): 75-79. Cryptocarya cercophylla W.E.Cooper is described and illustrated. Notes on habitat, habit and distribution are provided. The new species is restricted to a small area near Boonjee in the Wet Tropics of north-eastern Queensland. Key Words: Lauraceae, Cryptocarya , Cryptocarya cercophylla , rainforest, Australia flora, Queensland flora. Wet Tropics bioregion, new species W.E. Cooper, Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor Road, Smithfield, Queensland 4878, Australia Introduction Cryptocarya R.Br. (Lauraceae) is a pantropical genus of200-250 species of small to large trees, distributed from Africa, Asia, Malesia, Australia, the Pacific and South America (Le Cussan & Hyland 2007). In Australia, 48 species are now recorded, 39 or 40 are endemic and 36 of those occur in the rainforests of tropical Queensland. Many Cryptocarya species are very restricted in distribution. Most species in Australia occur in various forms of rainforest, ranging from wet evergreen rainforest to gallery forest and other more seasonal rainforests, with a few also occurring in wet sclerophyll forest. They occur in a variety of substrates from sea level to 1600 m. In February 1981, Laurence Jessup with J. Geoff Tracey and Tony Irvine collected the first specimens {Jessup 319, Tracey & Irvine and Tracey 14993 ) of the species described below as Cryptocarya cercophylla W.E. Cooper within the Boonjee Logging Area in the Russell River headwaters (now part of Wooroonooran National Park). The plant was originally thought to be a species of Icacinaceae. Flowering specimens were subsequently collected by Jessup, Gordon Guymer and Bill McDonald in October 1988, Bernie Hyland in July 1990, Bob Jago in August 1994 and Rigel Jensen in November 1994. By this time the plant was recognised as a species of Cryptocarya. Visiting Nigerian researcher, Olusegan Osunkoya with direction from Jensen, collected the first specimen with one fruit, in February 1994. Hyland et al. (2003) listed this entity under the phrase name Cryptocarya sp. (Boonjee BH 25794RFK), whereas the Queensland Herbarium listed it as Cryptocarya sp. (Boonjie L.W. Jessup+ 319) (Jessup 2007). However, it was not treated in either Hyland (1989) or the Flora of Australia (Le Cussan & Hyland 2007). With advice from Andrew Ford, a tree was located in January 2013 with enough fruiting material to enable a complete description of this species as Cryptocarya cercophylla W.E.Cooper. Materials and methods The study is based upon the examination of herbarium material from CNS together with relevant field observations. All specimens cited have been seen by the author. Measurements of the floral parts and fruits are based on material preserved in 70% ethanol and fresh specimens in the field. Abbreviations in the specimen citations are: LA (Logging Area), NP (National Park), SFR (State Forest Reserve) and TR (Timber Reserve). Accepted for publication 16 July 2013 76 Taxonomy Cryptocarya cercophylla W.E.Cooper sp. nov. Similar to Cryptocarya meissneriana Frodin but differs in the leaf apex (distinctly caudate, with acumen 8-20 mm long and 1- 2 mm wide, versus shortly acuminate or caudate with acumen up to 10 mm long and 2- 4 mm wide), lateral veins (3-6 versus 7-10), tepals (erect and not opening widely versus opening widely at anthesis), and anther apex (acute versus truncate). Typus: Australia: Queensland. Cook District: Timber Reserve 1230, Bartle Frere, Boonjee Logging Area [east of Malanda], 17 November 1994, R. Jensen 72 (holo: CNS [2 sheets + spirit], iso: BRI, CANB, MO distribuendi). Cryptocarya sp. (Boonjee BH 25794RFK); Hyland etal. (2003). Cryptocarya sp. (Boonjee); Cooper & Cooper (2004: 144). Cryptocarya sp. (Boonjie L.W.Jessup+ 319); Jessup (2007: 92, 2010: 97). Cryptocarya sp. Boonjee (B.Hyland 25794RFK); Hyland et al. (2010). Illustrations : Hyland et al. (2003), as Cryptocarya sp. (Boonjee BH 25794RFK); Cooper & Cooper (2004: 144) as Cryptocarya sp. (Boonjee). Hyland et al. (2010), as Cryptocarya sp. (Boonjee B.Hyland 25794RFK). Small tree to 10 m, poorly formed and often with coppice shoots, stems to 12 cm diameter at breast height, not buttressed. Bark pale with sparse vertical elongated lenticels and fine fissures; twigs terete, green. Leaves simple, alternate, petiolate, discolorous, glabrous; petioles 7-13 mm long, grooved and yellowish; lamina ovate or elliptical, 53-100 mm long, 22-45 mm wide, thinly leathery, upper side glossy dark green, lower side shiny (less so than the upper surface), paler green, base attenuate or cuneate, apex caudate, acumen 8-20 mm long, 1.3-2 mm wide; margins entire and slightly recurved, mostly sinuous; venation brochidodromous, midrib depressed on upper side; secondary lateral veins indistinct, 3-6 pairs at 45-90° to the primary midrib, oil dots numerous (visible Austrobaileya 9(1): 75-79 (2013) with a lens). Inflorescences axillary or rarely terminal, racemose or rarely paniculate, 4-7-flowered, 10-28 mm long, not exceeding leaves, rachis ± glabrous with occasional minute simple hairs; bracts triangular, c. 0.5 mm long and 0.2 mm wide, sparsely and minutely hairy, caducous. Flowers bisexual, actinomorphic, not fragrant, 3-merous, c. 2.5 mm long and 1.75 mm diameter; pedicels 0.6-2 mm long, sparsely and minutely hairy; perianth tube 2.5-2.8 mm long, c. 1.5 mm wide at apex, inner surface hairy, outer surface glabrous; tepals 6, in 2 whorls of 3, glabrous outside, sparsely hairy inside, creamy-green or cream, erect at anthesis; outer tepals 1.2- 1.7 mm long, 1.2-1.5 mm wide; inner tepals c. 1.7 mm long and 1.6 mm wide; stamens with filaments sparsely hairy, c. 5 mm long; gland heads conical, glabrous, c. 1 mm long and 0.4 mm wide, stalks sparsely hairy or glabrous c. 0.2 mm long; anthers 6 introrse and 3 extrorse, 2-locular, inner and outer anthers c. 0.5 mm long and 0.5 mm wide, glabrous; staminodes 3, head cordate, sessile, c. 0.7 mm long; ovary glabrous to sparsely hairy, 0.8-1 mm long, c. 0.5 mm wide; style glabrous, 1.2-1.6 mm long. Fruit a nut, appearing as an inferior drupe, elliptical, rarely ovoid or obovoid, mostly asymmetric at base with one side slightly lobed, 16.5-20 mm long, 10-14.5 mm wide, ribbed, glabrous, red to shiny black at maturity, epicarp 0.4-0.6 mm thick, mesocarp + epicarp c. 1.8 mm thick, mesocarp green, stigma persistent at apex with diameter c. 1.5 mm, pedicel 1-2.8 mm long; seed solitary, 16-19.5 mm long, 8-12.5 mm wide; endocarp beaked at base, cream, ribbed; cotyledons white, uniform in texture (not ruminate), radicle apical. Figs. 1 & 2. Additional selected specimens (from 25 examined): Queensland. Cook District: Tolga, cultivated, Jan 1995, Sankowsky 1437 (CNS); Atherton, cultivated, Jul 1990, Gray 5210 (CNS); Butchers Creek, c. 20 km ESE of Yungaburra, Stockwellia site, Jan 1985, Morawetz, Waha & Weber 114-30185 (CNS); TR 1230 Boonjie LA, Feb 1981, Jessup 319, Tracey & Irvine (BRI); Wooroonooran NP, Stockwellia site, July 2000, Forster PIF25913, Booth & Jensen (BRI); TR 1230, Bartle Frere, Boonjee LA, July 1990, Hyland 14052 (CNS), 14053 (CNS), 25793RFK (CNS), 25794RFK (CNS), 25795RFK (CNS); TR 1230, Boonjee LA, the headwaters of the Russell River, Feb 1981, Tracey 14993 (CNS); Wooroonooran NP, Stockwellia area south of road to Mt Bartle Frere, Site 216, Dec 2009, Ford 5636 & Cooper (BRI, CNS); Cooper, Cryptocarya cercophylla Stockwellia site, Boonjee, Wooroonooran NP, Jul 2011, Cooper 2136 & Ford (CNS), Oct 2012, Cooper 2165 (CNS); SFR 755, Bartle Frere, Gosschalk LA, Sep 1994, Gray 5774 (CNS); TR 1230, Boonjee LA, Feb 1994, Osunkoya 1 (CNS); Wooroonooran NP, along Russell 77 River track between lookout and Chucklunga track, Sep 2012, Ford 6082 (CNS); Wooroonooran NP, Donkey Track near Zig Zag, Mar 2003, Forster PIF29259 & Cooper (BRI); Wooroonooran NP, along Chucklunga track, Jan 2013, Cooper 2210 & Ford (CNS); Palmerston Ridge Track, Feb 1995, Hunter 1154 (BRI). Fig. 1. Cryptocarya cercophylla. A. stem with leaves showing main lateral veins *0.5. B. leaf showing primary, secondary and some tertiary venation xi. C. inflorescence showing open flower, buds and bracts on pedicels x4. D. flower (side view) x8. E. flower (side view with 3 tepals removed) *12. F. staminode x32. G. anther (inner whorl) x32. H. anther (outer whorl) x32.1. gland showing head and stalk xl. A & B from Forster PIF29259 & Cooper (BRI); C-I from Jensen 72 (CNS). Del. W. Smith. 78 Austrobaileya 9(1): 75-79 (2013) Fig. 2. Cryptocarya cercophylla fruiting stem (i Cooper 2210 & Ford [CNS]). Photo: A. Ford. Distribution and habitat : Cryptocarya cercophylla is endemic to Wooroonooran National Park within the Wet Tropics bioregion of north-eastern Queensland, where it is known to occur in a small area between what is locally referred to as the Stockwellia site at Boonjie and the nearby Russell River, at altitudes 580-680 m. The species may well occur in a broader area of the western Russell River catchment (Map 1). Cryptocarya cercophylla grows as an understory tree in a high rainfall area of notophyll rainforest on clay soils derived from metasediments. Tree species that it co-occurs with are Stockwellia quadrifida D.J.Carr, S.G.M.Carr & B.Hyland, Beilschmiedia oligandra L.S.Sm., Ceratopetalum virchowii F.Muell., Elaeocarpus johnsonii F.Muell Endiandra dichrophylla F.Muell, Flindersia bourjotiana F.Muell, Franciscodendron laurifolium (F.Muell) B.Hyland & Steenis and Peripentadenia mearsii (C.T.White) L.S.Sm., Bobea myrtoides (F.Muell.) Valeton and Aceratium doggrellii C.T.White. It appears to favour areas disturbed by landslips. Phenology : Flowers have been recorded in July, October, November and December: fruits have been recorded in January and February. Notes : In cultivation, Cryptocarya cercophylla has flowered as a shrub at 1.5 m tall; however, flowering collections in the wild are only known from trees 6 m and above. Fallen stems produce adventitious roots, which appear to eventually grow into individual trees following decomposition of the fallen stem. Evidence of such a phenomenon can be seen as a number of ‘individuals’ occurring close together (3-5 m) and in a straight line. Affinities: Cryptocarya cercophylla appears to be most similar to C. meissneriana from southeast Queensland and northeast New South Wales but differs from this species in the leaf apex (distinctly caudate, with acumen 8-20 mm long and 1-2 mm wide, versus shortly acuminate or caudate with acumen up to 10 mm long and 2-4 mm wide), leaf venation (3-6 secondary lateral veins versus Cooper, Cryptocarya cercophylla 7-10 secondary lateral) and the tepals erect and not opening widely at anthesis (versus tepals opening widely at anthesis). Conservation status : Cryptocarya cercophylla is a geographically restricted species and can be classified as Vulnerable (Criterion D2) using the criteria of the IUCN ( 2001 ). Etymology : The specific epithet is derived from the Greek, kerkos (tail), and phyllon (leaf), in reference to the distinctive caudate or tail-like leaf apices. Acknowledgements Frank Zich and Darren Crayn are thanked for support and access to CNS herbarium, Andrew Ford and Cath Moran for assistance and good company in the field and especially Andrew for finding the elusive fruiting tree, Peter Bostock and Will Smith for the map and illustrations. Laurie Jessup enlightened me regarding the discovery of the plant. Darren Crayn and Andrew Ford made worthwhile suggestions to an earlier manuscript. Permits to collect in National Parks and State Forests were issued by the Queensland Department of Environment and Resource Management. References Cooper, W. & Cooper, W.T. (2004). Fruits of the Australian Tropical Rainforest. Nokomis Editions: Melbourne. Hyland, B.P.M. (1989). A revision of Lauraceae in Australia (excluding Cassytha ). Australian Systematic Botany 2: 135-367. Hyland, B.P.M., Whiffin, T., Christophel, D.C., Gray, B. & Elick, R.W. (2003). Australian Tropical Rain Forest Plants. Trees, Shrubs and Vines. CD-ROM. CSIRO Publishing: Melbourne. Hyland, B.P.M., Whiffin, T. & Zich, F. (2010). Australian Tropical Rain Forest Plants Edition 6 (online version). CSIRO. http://www.anbg.gov.au/cpbr/ cd-keys/rfk/index.html Accessed October 2011. Iucn (2001). IUCN Red List of Categories and Criteria: Version 3.1. IUCN Species Survival Commission: Gland (Switzerland)/Cambridge (United Kingdom). 79 Jessup, L.W. (2007). Lauraceae. In P.D. Bostock & A.E. Holland (eds.). Census of the Queensland Flora 2007, p. 54. Environmental Protection Agency, Brisbane. - (2010). Lauraceae. In P.D. Bostock & A.E. Holland (eds.). Census of the Queensland Flora 2010, p. 92. Environmental Protection Agency, Brisbane. http://www.derm. qld.gov. au/wildlife- ecosystems/plants/pdf/qld-flora-census.pdf Accessed October 2011. Le Cuss an, J. & Hyland, B.P.M. (2007). Lauraceae. In A.S. George (ed.). Flora of Australia. 2: 106- 223. Australian Biological Resources Study: Canberra. Map 1. Distribution of Cryptocarya cercophylla in north-east Queensland. Shaded area on map indicates Wooroonooran National Park. Violaperreniformis (L.G.Adams) R.J.Little & G.Leiper, stat. nov with notes on Australian species in Viola section Erpetion (Violaceae) R. John Little 1 & Glenn Leiper 2 Summary Little, J. & Leiper, G. (2013). Viola perreniformis (L.G.Adams) R.J.Little & G.Leiper, stat. nov., with notes on Australian species in Viola section Erpetion (Violaceae). Austrobaileya 9(1): 80-101. Viola perreniformis (L.G.Adams) R.J.Little & G.Leiper is recognized as a distinct species in section Erpetion in Australia. A description of the species is presented. Leaf and floral characters of four closely related species in section Erpetion, V. banksii Thiele & Prober, V. hederacea Labill. subsp. hederacea, V. perreniformis stat. nov., and V. silicestris Thiele & Prober, are compared and discussed. Various aspects of the biology of these species are discussed and provisional keys are presented based on floral and leaf characters. Comments are also provided for four other species in section Erpetion. V. cleistogamoides (L.G.Adams) Seppelt, V eminens Thiele & Prober, V. fnscoviolacea (L.G.Adams) T.A. James, and V. sieberiana Sprengel, and a putative hybrid, V x zophodes Thiele & Prober. Key Words: Violaceae, Viola , Viola section Erpetion, Viola banksii, Viola betonicifolia, Viola caleyana, Viola cleistogamoides, Viola eminens, Viola fuscoviolacea, Viola hederacea subsp. hederacea, Viola perreniformis, Viola sieberiana, Viola silicestris, Viola x zophodes, anther gland, cleistogamy, nectary 'R. John Little, MagnaFlora, 16 Pebble River Circle, Sacramento, California 95831, U S A. Email: bugsandbunnies4u@sbcglobal. net 2 Glenn Leiper, 30 Tweedvale Street, Beenleigh, Queensland 4207, Australia. Email: leiper30@ bigpond.com Introduction Viola hederacea Labill. subsp. perreniformis L.G.Adams was one of eight subspecies recognized by Adams in the V hederacea complex in Australia (Adams 1982a,b). This was the first attempt to classify the diversity of forms that had been assigned to V hederacea sensu lato in section Erpetion (Sweet) Benth. & Hook.fi Later, James (1990) recognized seven forms (A through G) of V hederacea in New South Wales based on leaf shape and pubescence, and in addition, elevated three of Adams’ subspecies to species rank. Thiele & Prober (2003) summarized the taxonomic status of each subspecies of V hederacea described by Adams with all subspecies except subsp. perreniformis and subsp. curtisae L.G.Adams having been elevated to species rank. In the original description, Viola hederacea subsp. perreniformis was distinguished from V hederacea subsp. hederacea by the erect, soft, short pubescence, by the corolla being more or less concolorous, pale (lazuline) blue- violet, with the anterior petal white at the base with violet-coloured veins, and distinguished from other V. hederacea subspecies by suborbicular leaves with a deeply cordate base (Adams 1982a). The epithet ‘perreniformis’ means ‘very reniform.’ Interestingly, the adjective ‘reniform’ with respect to leaves of subsp. perreniformis was not mentioned in the Latin diagnosis and appears only in the description of subspecies (Adams 1982b). The leaves of subsp. hederacea were stated by Adams (1982b) to be variable but never suborbicular (authors’ emphasis). Adams reported that subsp. perreniformis occurred in coastal regions between Cairns, Queensland (Qld) and Sydney, New South Wales (NSW). Accepted for publication 26 September 2013 81 Little & Lei per, Viola perreniformis Some Australian botanists have considered Viola hederacea subsp. perreniformis as warranting specific status (e.g. Thiele & Prober 2003), but the nomenclatural change has not been made. Based on a study of the holotype and isotype as well as other collections of the V hederacea complex at BRI, CANB, and NSW, and recent field observations by the authors, we believe subsp. perreniformis differs sufficiently from subsp. hederacea to warrant recognition at species rank, as V perreniformis. The close relationship among the four taxa, V banksii, V hederacea subsp. hederacea, V perreniformis and V silicestris, can be seen in the morphology of their leaves and flowers. Long regarded as a difficult taxon because of its many leaf forms, recent studies of the Viola hederacea complex in Australia in section Erpetion have begun to clarify relationships among taxa in this group. In 2003, Thiele & Prober presented the most comprehensive study and summary to date of species in section Erpetion (Thiele & Prober 2003), which included a circumscription of species in the V hederacea complex, descriptions of several new species including V banksii , and a key to most members of the complex. Their circumscription of V hederacea did not include V hederacea subsp. perreniformis , which they believed to be a taxon distinct from V hederacea. Thiele & Prober (2006) described V silicestris as a new species and discussed similarities and differences of it compared to four other species in section Erpetion. V. cleistogamoides (L.G.Adams) Seppelt, V fuscoviolacea (L.G.Adams) T.A.James, V. hederacea and V sieberiana Sprengel. During our study of Viola perreniformis, we found that V banksii, V. hederacea subsp. hederacea , and V silicestris were the taxa most often confused with Vperreniformis stat. nov. In this paper we examine and compare these four species (collectively referred to herein as the “four taxa”) with new and existing data, document previously unreported observations of various morphological features, and suggest topics for future study. The taxonomy of section Erpetion is challenging and far from complete. The four taxa and other species in the section appear to have many unique features not known to occur in other Viola species. Materials and methods In November and December 2011 and January 2012, the first author examined herbarium specimens in the Viola hederacea complex at the herbaria BRI and NSW, including specimens identified as subsp. perreniformis. Both authors conducted numerous field studies in Qld and NSW during this timeframe. In 2013, the first author continued study of species in the V. hederacea complex at the herbaria CANB and NSW, and both authors examined specimens at BRI identified as subsp. perreniformis and subsp. hederacea. Scanned images were examined of slides of the type of V hederacea at CANB. At BRI, all specimens previously determined as subsp. perreniformis were compared with the isotype housed at BRI and an image of the holotype housed at CANB. Specimens were then sorted into three groups: 1) specimens that matched or closely resembled the holotype and isotype (these specimens are listed below under Additional specimens examined)', 2) specimens generally similar to the holotype and isotype, but with leaves too immature or in too poor a condition to make a definitive determination; and 3) specimens which, in the authors’ opinion, did not fit the concept of the perreniformis taxon, most of which are referable to subsp. hederacea. Over 1000 high-resolution, digital photographs were taken of herbarium specimens to facilitate further study. Field observations of Viola perreniformis were made by both authors in February 2013 at Nightcap National Park (NP) in NSW. The second author made additional observations and collections of this taxon in March, April, and May 2013 at Crows Nest NP and in March 2013 at Moogerah Peaks NP, both in Qld. In June 2013, the second author collected fresh flowers of V banksii from Springbrook NP (possibly a horticultural escape) and Venman Bushland NP and V hederacea from Springbrook NP, all sites in Qld; the anther 82 glands on these flowers were then observed with a stereozoom microscope. Using a stereozoom microscope with a camera attachment, the first author obtained micro¬ photographs of flowers and anther glands of V banksii and seeds of V perreniformis and V betonicifolia. The species description is based on a composite of observations and measurements of leaves, petioles, peduncles, and flowers from a) the holotype and isotype; b) from herbarium specimens at BRI, and c) from plants collected by the authors in 2012 and 2013 and photographs taken of fresh material. The number of ‘crenations’ (rounded lobes) on leaf margins were recorded only from one side of a leaf because on herbarium specimens, overlapping leaves often prevented observation of the entire margin. As used herein in reference to pubescence, the term pilose means soft, thin, straight, and clearly separated hairs; the type of hairs on leaf surfaces of Viola perreniformis. Villous means ± densely covered with soft, thin, long, straight or sometimes shaggy hairs, not clearly separated; the type of hairs often found on petioles and peduncles of V perreniformis. Common abbreviations used in specimen citation include CP (Conservation Park), NP (National Park) and SF (State Forest). Taxonomy Viola perreniformis (L.G.Adams) R.J.Little & G.Leiper, stat. nov.; Viola hederacea Labill. subsp. perreniformis L.G.Adams, FI. Aust. 8: 386 (1982). Type: Queensland. Port Curtis District: c. 40 km WSW of Gladstone, Many Peaks Range, 2 June 1977, M.D. Crisp 2751 & R.K. Ellyard (holo: CANB, image!; iso: BRI). Plants perennial, often appearing acaulescent, 3-6.5 cm high, stoloniferous, rooted sections of the stolon forming rosettes of leaves on short, fleshy rhizomes. Stems sometimes present, 1-8 cm long. Leaves: stipules narrow-ovate or lanceolate, margin irregularly fimbriate, apex acute, long-aristate; petiole 9-23 cm long, sparsely or densely pilose or villous, occasionally glabrous; blade reniform, sub- Austrobaileya 9(1): 80-101 (2013) reniform, or ovate (outline ovate, orbicular, suborbicular, or semicircular), 5.5-23 mm long, 7-27 mm wide, base cordate or deeply cordate, occasionally subtruncate or truncate, often with distinct sinus between lobes; margins ciliate or eciliate, uniformly or irregularly crenate from basal lobes to apex, 4-9(-10) lobes per side, sometimes poorly defined, occasionally some lobes ± serrate, apex acute, obtuse, or rounded, occasionally ± truncate, the midvein terminating in a blunt mucro; abaxial surfaces of mature and immature leaves sparsely or densely pilose, with hairs scattered on 1° and 2° veins or over entire surface, occasionally glabrous, small or immature leaves sometimes tomentose; adaxial surfaces of mature and immature leaves sparsely or densely pilose, with hairs scattered over entire surface, occasionally glabrous; blades bi-colored, darker green adaxially, lighter abaxially. Flowers solitary, peduncles arising from a fleshy, usually vertical rhizome, extending above the leaves, 25-77 mm long, sparsely or densely pilose or villous, occasionally glabrous; bracteoles narrowly linear or aciculate, 2.5-5 mm long, opposite or alternate, often above the middle during capsule development; sepals lanceolate, margin entire, scarious, eciliate, apex acute, usually gland-tipped, auricles not prominent, c. 0.5 mm long; upper petals erect in young and reflexed in mature flowers, pale blue-violet on both surfaces, white basally, occasionally with a faint or prominent violet- coloured blotch distal to the proximal white area; lateral petals spreading in new and twisted in mature flowers, pale blue-violet on both surfaces, occasionally appearing white distally, white basally with a green and white, ± 1-2 mm long, ± 1-2 mm wide indurate area at the base, often with a faint or prominent violet-coloured blotch distal to the proximal white area, with faint or prominent violet-coloured veins, usually with a dense patch of white clavate and cylindrical beards on proximal third of petal, often with some tips violet-coloured, distal beards sometimes entirely violet-coloured; lowest petal obovate to oblong, usually white on proximal half with a prominent green and white, 1-2 mm long, 1-2 mm wide indurate area at the base, 83 Little & Lei per, Viola perreniformis the distal half of petal pale blue-violet, the veins violet-coloured, prominent on adaxial surface, faint or prominent on abaxial surface, usually not extending to distal margin of petal, the midvein and the two adjacent veins distinct, prominent, parallel, ± same length, scarcely or not anastomosing (without evident, interconnecting 2° veins), 5-9 mm long, apex retuse, truncate, rounded, occasionally with a short, sharp or rounded point between two lobes, spur not present; style geniculate near attachment to ovary, ± terete, style head glabrous, not differentiated from style shaft; anthers cream-coloured, anther glands pale green, smooth, c. 14-2/3 as long as anther, prominent, margins of terminal anther appendages not hairy; pollen cream- coloured. Capsules 5-6 mm long, 3-valved, often purple-spotted, glabrous. Seeds ovoid, surface uneven to irregularly rugose, 1—1.2 mm long, shiny or dull dark brown mottled with patches of light brown or tan, elaiosome not present. Additional selected specimens examined : Queensland. Cook District: Intersection of road to radio tower and Longlands Gap Road, Longlands Gap SF, Aug 2007, Zich 617 & Quinn (BRI, CANB). North Kennedy District: Mt Fox, Dec 1949, Clemens s.n. (BRI [AQ115414]). South Kennedy District: Dalrymple Heights & vicinity, Jul 1947, Clemens s.n. (BRI [AQ115415]). Leichhardt District: Carnarvon Creek Gorge, 70 miles [112 km] NW of Injune, May 1962, Johnson 2411 (BRI); Blackdown Tableland, c. 1 km E of main N-S road crossing, Jan 1973, Henderson H1392 & Sharpe (BRI). Port Curtis District: c. 60 km SW of Gladstone, 8 km SE of Amys Peak, Jun 1977, Crisp 2793 (BRI); Kroombit SF 316, Aug 1984, Gibson 634 (BRI); Castletower Mt, Oct 1988, Randall 591 (BRI); Portion 10 V, Oaky Creek, Castletower, Apr 1989, Gibson TOI482 (BRI). Wide Bay District: Mt Woowonga, Woowonga Range SF, Jan 1982, Forster PIF1153 (BRI); Tinana Creek at Missings Crossing, May 2001, Stephens 138903A(11) (BRI); Mt Boulder, SE of Gympie, Oct 1993, Bean 6831 (BRI); Mapleton Logging Area, Mapleton SF, W of Mapleton, Mar 1998, Bean 13121 (BRI, CANB, HO, MEL, NSW). Moreton District: Mt Mee SF 893,2 km SE of Forestry office, Sep 1997, Halford Q3357 & Boyle (BRI); Alice Creek, 7.5 km ESE of Murphys Creek Rail Siding, Aug 1990, Forster PIF7100 & Bird (BRI); Mt Edwards NP, 14 km W of Boonah, Sep 1992, Forster P1F11483 & Reilly (BRI); Mt Edwards, Moogerah Peaks NP, c. 14 km W of Boonah, Feb 2010, Halford Q9754 (BRI); Mt May, Sep 1992, Forster PIF11794 et al. (BRI); Southern slopes of Mt Greville, SW of Lake Moogerah, Jan 2007, Phillips 1679 & Phillips (BRI); Between N and S peaks of Mt Maroon, W of Rathdowney, Apr 2009, Phillips 2006 & Phillips (BRI); Cronins Creek between Mt Barney & Mt Ernest, Dec 1932, White 8757 (BRI); Near summit of Mt. Lindsay [Lindesay], Sep 1973, Dowling 109 (BRI). New South Wales. Terania Creek, 10 km NNE of The Shannon [Channon], Feb 1980, Coveny 10628 (BRI); Summit of Mt Carnham, W of Grafton, in the Baryugil area, Jun 1993, Bryant s.n. (BRI [AQ566756]); Summit of Munningyundo Mt, Nymboida NP, between Grafton and Glen Innes, Mar 2003, Bean 20105 (BRI); North Rocks Road, Nightcap Range, Little 13697 & Leiper (BRI). Distribution and habitat : The species occurs in a series of disjunct localities from west of Cairns, north Qld, to Sydney in NSW. Recorded elevations of specimens examined in this study range from 75 m at Oaky Creek, Castletower to 1160 m at Mt Lindesay, both in Qld. Of the 14 specimens that provide elevation data nearly 60% were collected at 500 m or higher. Microhabitat situations reported by collectors include: moist situations in wet sclerophyll forests, in woodlands, in open eucalypt forests, on steep slopes, in crevices of rocky cliffs, cascading over rock faces with or without water flowing, on rhyolite or on rocky brown loam, granite, or sandstone; east, south, and west-facing slopes (Fig. 1). Phenology : Flowering and fruiting occurs throughout the year. Conservation status : Viola perreniformis is widespread, occurs in several National Parks, and is not known to be at risk. Notes on Viola section Erpetion Observations on aspects of morphological variation and reproductive biology focus primarily on the four taxa Viola banksii, V hederacea subsp. hederacea , V. perreniformis and V silicestris. Information is also presented for four other species in the section, V. cleistogamoides, V. eminens Thiele & Prober, V fuscoviolacea, and V sieberiana and a putative hybrid, V. x zophodes Thiele & Prober. Other taxa recognized in section Erpetion that are not discussed include V hederacea subsp. curtisiae and V improcera L.G.Adams. Stolons and Rhizomes. All species in section Erpetion produce stolons and two small bracts are always present between stolon nodes. Thiele & Prober (2003) reported that 84 Fig. 1. V. perreniformis. Habitat, Crows Nest NP, Qld, Mar 2013. Photo: G. Leiper. rootstocks of Viola banksii and V hederacea were sometimes somewhat swollen and bulbous at the stem bases, but they were not or scarcely swollen in V silicestris (Thiele & Prober 2006). The authors made similar observations for these species. Stolons in section Erpetion are initially produced on the surface. Overtime depending on their location, they can become buried with eroded soil, leaf litter, etc. Numerous herbarium specimens and the author’s recent collections show that buried, subterranean stolons can eventually become lignified in some species. This phenomenon is not known to have been previously reported for Australian species of Viola in the section. Although a buried stolon is technically a ‘rhizome,’ buried stolons of these species do not become thick and fleshy, but retain their narrow shape. A stolon node situated in a suitable site has the capacity to produce a vertically oriented rhizome and a rosette of leaves. As a rhizome matures it elongates vertically, its diameter increases, and it becomes fleshy, similar to many other Viola species. New leaves are produced distally as older leaves die. Leaf abscission scars and the remains of stipules are visible on the surface of the rhizome. If a vertically oriented rhizome is not buried, it can attain the appearance of a miniature tree; a large clone can attain the appearance of a miniature forest. A similar situation is sometimes observed in other Viola species e.g. V odorata L. and V riviniana Rchb. If a Austrobaileya 9(1): 80-101 (2013) vertically oriented rhizome is slowly buried with eroded soil, it can survive by elongating vertically, a useful strategy in a geologically unstable environment. All that may be visible at the surface is a rosette of leaves. Although empirical data are lacking, it is possible that fleshy rhizomes function as water storage organs. Enlarged, subterranean rhizomes were found by the second author in a population of Viola perreniformis plants at Crows Nest NP, Qld (Fig. 2). Its gourd-like shape suggests it may function as a water and/or nutrient storage organ. Perhaps this rhizome was initially above ground and then subsequently buried; perhaps it was then able Fig. 2. V perreniformis. An enlarged, subterranean rhizome; length between arrows is c. 22 mm. Crows Nest NP, Qld, Mar 2013. Photo: G. Leiper. to retain water more efficiently and gradually assumed a gourd-like shape. Clones of some species can occupy up to 1 m or more in diameter. Although detailed studies of how these species spread via stolons have not been made, the first author recorded preliminary observations of the stages that a single ‘mother’ plant of Viola hederacea went through as it matured and spread over time; a brief description follows. After several leaves in the rosette of the mother plant matured and enlarged, five stolons developed that spread out in all directions. A rosette of miniature leaves formed at the distal end of each elongating stolon. If the stolon tip contacted suitably moist soil, a white, root-like structure developed and grew into the soil. After several days, the leaves in the newly rooted rosette increased in size and simultaneously 85 Little & Lei per, Viola perreniformis several new stolons formed and spread out in all directions and as they elongated, rosettes of miniature leaves formed at the tip of each new stolon. This process was repeated over and over in all directions from the initial mother plant until there was a mass of stolons and leaves c. 1 m in diameter. If a node didn’t contact moist soil, the stolon continued to elongate, developing rosettes of leaves at each node. Because the available evidence suggests seeds of the four taxa may not be ant dispersed (see below) and they may not produce cleistogamous flowers (see below), vegetative reproduction through vigorous stolon growth may be an important strategy for survival in section Erpetion. Stems. A valuable taxonomic character commonly used to separate Viola species in European, North American, and Asian floras, is whether the plant has stems (caulescent) or does not (acaulescent). Stems in caulescent Viola species are usually upright structures that originate from a fleshy or somewhat woody rhizome and possess alternating leaves subtended by stipules. Peduncles that produce chasmogamous and in many species, cleistogamous flowers, arise from axillary positions on the stem. In caulescent species in North America, the internode distance is usually short early in the growing season, elongating later in the season. Peduncles in caulescent species are normally produced only from stems and not directly from a rhizome. Caulescent Viola species in North America have both basal and cauline leaves, although a few species have only cauline leaves and no basal leaves. When present, basal leaves arise directly from the rhizome and are taxonomically useful in some species. In all acaulescent species known outside of Australia, the leaves and peduncles arise directly from a rhizome. Of interest relative to the foregoing is that all Australian species in section Erpetion produce stolons. By analogy with the acaulescent, stoloniferous species in Europe and North America, species in section Erpetion would be assumed to be acaulescent. However, the Viola treatments in most Australian floras e.g. Flora of Australia (Adams 1982b), Flora of South-East Queensland (Stanley & Ross 1983), Flora of New South Wales (James 1990), Flora of Victoria (Entwisle 1996), Electronic Flora of South Australia (2007), Flora of Tasmania Online (Duretto 2009), and New South Wales Flora Online (James 2013), state that stems are present for all species in the section. “Stems short” and “stems short, erect” are common descriptions in these floras for stems of species in section Erpetion. Also of interest is that the peduncles of all Australian species in section Erpetion , with two known exceptions, appear to develop only from rhizomes produced at a stolon node, similar to acaulescent, stoloniferous species elsewhere in the world. Thus, at present it appears that stems do not normally occur in section Erpetion , such as those typically found in caulescent species, e.g. V caleyana Don in Australia. A remarkable exception was documented by Thiele & Prober (2006) when they reported that Viola silicestris often develops “a caulescent growth habit when growing in sheltered sites amongst other vegetation, with elongate, weakly erect, scrambling aerial stems to 30 cm high and scattered leaves. Plants in exposed sites have contracted stems and fasciculate leaves”. In northern NSW and Qld, we found stems on V silicestris plants that had developed at stolon nodes and peduncles that developed from axillary nodes on the upright stems. Peduncles may also develop directly from rhizomes at stolon nodes, but this has not been verified. Stems can be short, e.g. 1.5 cm, or greatly elongated, 30-40 cm (Thiele & Prober 2006). The authors found a population in Springbrook NP, Qld, where clearly discernable stems occurred at 6 of 8 nodes on a stolon 75 cm long. At another location in Springbrook NP an 85 cm long stolon had discernable stems at 3 of 10 nodes. It may be that stem development in this species is more common, perhaps overlooked when stems are short and hidden in a rosette of leaves. The development of a stem from a stolon node has not previously been documented in other Viola species known to the authors and may be unique to section Erpetion. Acaulescent, stoloniferous species such as V odorata typically root at a node and form a rosette 86 Austrobaileya 9(1): 80-101 (2013) of leaves similar to the four taxa, but erect stems are never produced by such species at a stolon node. The growth habit of V silicestris may represent a transitional stage from an acaulescent to a caulescent habit. Viola perreniformis also produces stems. In 2013 the authors found plants with stems in Nightcap NP in NSW (Little 13697 & Leiper [BRI]) (Fig. 3). In addition, possible evidence of stems have been observed on several herbarium specimens at BRI that were collected in Qld and identified by the authors as V perreniformis. Stems were not observed in populations of V perreniformis at Crows Nest NP or on Mt Edwards (Moogerah Peaks NP) in Qld. Although phenotypically similar, it is possible these populations are genetically different and never produce stems, or stems were produced but overlooked, or perhaps microhabitat conditions precluded stem development. Thiele & Prober (2006) noted that Viola hederacea and V sieberiana, in contrast with V silicestris , never (author’s emphasis) develop a caulescent growth habit, even when growing amongst dense vegetation. Our observations of herbarium specimens and of living populations of V banksii in Qld have not revealed evidence of stems. In V banksii , V hederacea and V sieberiana , leaves and peduncles appear to arise only from a rhizome and never from a caulescent stem. These observations suggest these taxa could be acaulescent, although additional observations are needed. No data are available for V cleistogamoides or V fuscoviolacea regarding presence/absence of stems. Leaves. The size and shape of leaves of most Viola species can vary greatly due to many factors, such as time of year (spring versus summer), the amount of soil moisture and other edaphic factors, and whether in sun or shade (Little & McKinney in press). Thiele & Prober (2006) appear to have been the first to document similar observations in reference to Australian violets, stating “This is a problem throughout the section, with reduced leaves of all species converging in shape [when immature] while fully developed leaves from sheltered sites are usually highly Fig. 3. V perreniformis. Pressed specimen showing stem (arrow) arising from a stolon node; 3 peduncles from the stem. Nightcap NP, NSW. Feb 2013. Photo: J. Little. distinctive”. They noted that most herbarium specimens fail to adequately represent the local variation present in leaves of species in section Erpetion and that the location of the plant, e.g. sun or shade, can have a profound effect on leaf size. We concur and emphasize that mature, fully developed leaves can facilitate identification of the four taxa. However, plants may have mature leaves that are not fully expressed, i.e. not typical, if they are growing in difficult sites (pers. comm., K. Thiele and observations by the authors). Thus, having fully developed, mature leaves may be essential for identification. In addition, it is necessary to consider all the leaves in a population and not focus on a few variants because in most populations of the four taxa, a few leaves can often be found at different stages of growth that are more typical of one of the other species. Also important to note is that various combinations of the four taxa are sympatric and it is possible that the stolons of two or more species could become intertwined in a given population resulting in some confusion if the leaves of one species Little & Lei per, Viola perreniformis are associated with the flowers of another. The leaves of all four taxa are darker green adaxially compared to lighter coloured abaxial surfaces (Fig. 4). The overall outline of Viola banksii and V perreniformis leaves remain ± proportionally constant during development from immature to fully developed, whereas the leaves of V hederacea and V silicestris expand laterally as the blade matures, with the blade becoming much wider than long. The information in Table 1 is an attempt to qualitatively summarize and contrast morphological leaf characters in the four taxa typically found in fully developed leaves. Some characters for a given species 87 appear genetically fixed and do not vary, e.g. fully developed Viola silicestris leaves never have an orbicular outline; the outline of V banksii and V. perreniformis leaves are never ± rectangular and their shapes are never falcate; a distinct sinus never occurs in fully developed V hederacea or V silicestris leaves, etc. As noted in the table, other morphological characters are only ‘sometimes’ present in a given leaf or population. ‘Usually’ in the table indicates the norm for the species, recognizing that variation occurs in that character. Immature leaves of some species have characters associated with fully developed leaves of other species (see discussions below). Table 1. Qualitative summary of some leaf characters typically found in fully developed leaves of some Viola species Leaf characters V banksii V perreniformis V hederacea subsp. hederacea V silicestris outline orbicular usually sometimes no no outline ovate no sometimes no no outline ± rectangular no no sometimes sometimes outline semi-circular no sometimes usually no shape distinctly reniform sometimes usually no no shape sub-reniform (i.e. cordate lobes not well developed) no sometimes sometimes sometimes shape falcate no no no sometimes leaf margins crenate from base near netiole to apex no usually no no 1 leaf base cordate yes usually no no leaf base truncate no sometimes usually usually distinct basal lobes present yes usually no no distinct sinus present formed by lobes yes usually no no leaves much wider than long no no yes yes The distal leaf margins of V silicestris are occasionally ± crenate, but the basal part of the blade is never crenate. Austrobaileya 9(1): 80-101 (2013) Viola banksii: Mature leaves ultimately attain an ± orbicular outline and shape or sometimes a reniform shape. The apex of mature leaves is usually truncate or rounded; a distinct mucro at the apex of the midvein is sometimes present; the base is cordate, never truncate. The two distinct basal lobes often overlap in fresh material, but overlapping lobes can be obscured when pressed. As the leaf matures and expands, the marginal teeth become further apart and often become small and less obvious on the upper half of the leaf. The margin is occasionally ± dentate. Occasionally small teeth and/or 3 or 4 small lobes (crenations) are present on the lower half of the leaf, but never around the entire margin. Mature, orbicular-shaped leaves are about as wide as long; reniform-shaped leaves are slightly wider than long. In small, immature leaves a distinct sinus is often present. A distinguishing characteristic of this species is the tendency for the apex of fully developed leaves to be truncate or somewhat rounded while also having cordate basal lobes and a distinct sinus (Fig. 4). Viola hederacea subsp. hederacea: Mature leaves have been described as broad-reniform or semi-circular, usually truncate at the base and a margin with 8—16 teeth, the base occasionally with a broad sinus or broadly cuneate (Thiele & Prober 2003). Images of Fig. 4. V. banksii. Sequence of development from new (1) to fully developed leaf (8). The smallest leaves often have a distinct sinus. Adaxial surfaces are shown; abaxial in 7. Pilose pubescence present on leaves and petioles. Photo: J. Little. the type specimen show many small and large leaves with a distinct cuneate base. We found blade outlines of fully developed leaves to be ± rectangular (Fig. 5.8). Leaf apices are variable, rounded, obtuse, truncate, or occasionally convex. When the apex is truncate or convex, the outline is arguably not semi-circular, but closer to rectangular. The terminal midvein is usually just a small point or mucro. The distal (upper) leaf margin is ± entire with only a few short points where the major veins terminate. The base of fully developed leaves is usually Fig. 5. V hederacea. Sequence of development from new/immature (1) to fully developed leaf (8). Basal angle increases with age until becoming truncate in fully developed leaves. Adaxial surfaces shown. Photo: J. Little. 89 Little & Lei per, Viola perreniformis truncate and does not form distinct lobes as in V banksii or V perreniformis. Occasionally, the basal blade margin is curved or arched downward between the petiole attachment and the outermost edge of the blade, similar to V silicestris. Such leaves are sometimes described as having a ‘wide’ sinus. The basal margin of young and fully developed leaves is entire or may have a small point or two and often forms a ± straight line between the petiole attachment and the outer edge of the blade. The angle formed by the leaf base in fully developed leaves is obtuse to usually truncate (180°). Distinguishing features in fully developed leaves separating Viola hederacea (Fig. 5.8) from V banksii and V perreniformis are a truncate leaf base and lack of a sinus. Fully developed leaves of V hederacea are somewhat similar to V silicestris in that both can be ± rectangular and are much wider than long. Immature V hederacea leaves usually have small lobes, but as the leaf matures and enlarges, the angle between the lobes gradually increases and the lobes ‘disappear’ as the base becomes truncate. Occasionally, small, immature leaves of V hederacea have very distinct lobes and a distinct sinus and are very similar to immature leaves of other species, e.g. compare Figs. 4.5, 5.3, 6.1. Our observations of herbarium and living specimens of Viola hederacea suggest that leaves that match the description of “broad- reniform” are not fully developed or perhaps were growing in difficult sites. The lobes in broad-reniform leaves are small (=sub- reniform) and the width of such blades are smaller compared to a fully developed leaf. The typical leaf shape of V eminens was described as “broad-reniform” (Thiele & Prober 2003); however, we have not studied this species and our observations that broad- reniform leaves are not fully developed pertain only to V hederacea. Although V hederacea subsp. hederacea was reported to be replaced in Qld by V. hederacea subsp. perreniformis (Thiele & Prober 2003), many populations have been found in Qld that morphologically match the description of V hederacea and are considered by the authors and others to be V hederacea. The second author has found several populations in Qld where the flowers are somewhat similar to V hederacea, but the leaves are cordate or reniform with distinct basal lobes. Viola perreniformis : Outlines of fully developed leaves are ovate, orbicular, suborbicular, or semicircular; their shapes are reniform or ovate. The apex is acute, obtuse, or rounded, occasionally ± truncate. The base of mature and immature leaves is usually cordate with a distinct sinus formed by the lobes of the blade. The sinus is usually distinct, but wider than in V banksii. Most populations of V perreniformis in Qld occasionally have some truncate and/or subtruncate leaf bases in addition to typical leaves with cordate bases, but these are usually associated with immature leaves. The marginal crenations (lobes) are usually quite uniform, but can be somewhat ‘ragged’ and irregularly shaped in some populations. Although the number of crenations per blade can vary in the same clone and among different populations, distinct crenations are present on most leaves in a clone. The number of crenations on one side varies from four in small/immature leaves to nine (seldom ten) in the largest/fully developed leaves. Fully developed leaves are usually slightly wider than long. Distinguishing features are leaves with crenate margins, usually around the entire blade in fully developed and usually also in immature leaves (Fig. 6). Fig. 6. V perreniformis. Sequence of development from immature (1) to fully developed leaf (5). Leaves at all growth stages are lobed. Adaxial surfaces shown. Photo: G. Leiper. 90 Austrobaileya 9(1): 80-101 (2013) Viola silicestris : Fully developed leaves have been described as broad, occasionally sub-reniform, the margin with (5—)7-12(— 14) obscure teeth, the apical tooth not prominently larger than the rest (Thiele & Prober 2006). Here, we describe the overall outline of a fully developed blade as broad or ± rectangular, often with a falcate shape and occasionally sub-reniform. When falcate, the upper and lower margins are parallel. Occasionally a leaf is found with a shape resembling the tail of a diving whale. The apex is often truncate, sometimes rounded; the terminal midvein usually not much longer than the two adjacent veins, but sometimes shorter. The terminal midvein is usually just a small point or mucro. The blades are not distinctly lobed and a distinct sinus is not present. The base of fully developed leaves is truncate, although sometimes slightly curved or arched downward between the petiole attachment and the outermost edge of the blade. Occasionally, the blade bases curve upward from the petiole. The basal margin is entire or may have a small point or two. The distal leaf margin is ± serrate, subserrate, sometimes denticulate, or occasionally ± crenate. The outline of immature leaves is semi-circular or sub-reniform, smaller leaves are sometimes ± ovate; the apex obtuse or acute; the upper margin serrate, subserrate, or ± denticulate, the leaf base attenuate (cuneate) in some populations and truncate in others. Distinguishing features of this species are leaves that are usually much wider than long through all stages of growth from new/ immature to fully developed. Fully developed leaves have a ± rectangular outline or falcate shape and never have distinct lobes or a sinus (Fig. 7). Indumentum Viola banksii: This species was described as glabrous (Thiele & Prober 2003). However, several specimens at BRI collected in Qld have pilose hairs on adaxial, or on adaxial and abaxial leaf surfaces (e.g., BRI [AQ11278], [AQ115406]). A number of other populations in Qld appear to be V banksii based on leaf shape, but the plants were variously pubescent on one or both surfaces, or leaves and petioles Fig. 7. V silicestris. Sequence of development from immature (1) to fully developed leaf (5). Leaves are much wider than long at all growth stages. Adaxial surfaces shown. Photo: G. Leiper. were densely pubescent (Table 2). V. banksii plants are sold commercially worldwide under the name “ Viola hederacea ” (Thiele & Prober 2003; Elliot & Jones 2010). The first author has maintained a clone of V banksii for over 10 years purchased as “ Viola hederacea ” from a nursery in Sacramento, California, U.S.A. The leaves, petioles, and peduncles on this clone are usually pubescent (Fig. 4), similar to some of the populations noted in Table 2. When leaves of V banksii are pubescent, the margins are usually ciliate, as in Vperreniformis. Based on our observations of pubescence on V banksii , we believe this character state needs to be recognized in descriptions of this taxon. Further study is needed of the populations noted in Table 2. Viola hederacea subsp. hederacea : Leaves were described as glabrous or with scattered unicellular hairs on the upper surface (Thiele & Prober 2003). Our observations of populations in southeast Qld and northern NSW are similar, although we have observed scattered hairs are sometimes present on both leaf surfaces. In addition, this species occasionally has hairs proximally on petioles and peduncles. In comparing the leaves of this taxon with V silicestris , a general tenet is that V hederacea leaves are often short- hairy, at least on adaxial surfaces, whereas V silicestris leaves are usually glabrous. Viola perreniformis : Adams (1982b) stated that leaves of this species were softly pubescent, rarely almost glabrous. The authors Little & Lei per, Viola perreniformis 91 Table 2. Populations of pubescent Viola banksii cf. in Queensland Location in Queensland Comment Pine Ridge CP and Jacobs Well (Gold Coast coastal areas) Plants appear to be V banksii. At both locations leaves have scattered hairs; at Jacobs Well, petioles are hairy. Point Cartwright (Sunshine Coast) Plants appear to be V banksii. Leaves are sparsely hairy. Alexandra Headland (Sunshine Coast) Plants appear to be V banksii. Leaves and petioles are very hairy. Maroochy River CP (Sunshine Coast) Plants appear to be V banksii. Leaves with scattered hairs on both surfaces. Ningi (Abbey Museum at the southern end of the Sunshine Coast) Plants appear to be V banksii. Leaves with scattered hairs on both surfaces. conducted a brief evaluation of pubescence on leaf surfaces, petioles, and peduncles among V perreniformis populations in Qld and NSW. The evaluation revealed substantial variation in pubescence on the same plant, between plants in the same population, and among different populations. Most populations were sparsely to densely pubescent on both leaf surfaces although the density of hairs usually varied between the abaxial and adaxial surfaces on the same leaf. Some populations had both glabrous and pubescent leaves on the same plant. Leaf margins were ciliate or eciliate. The petioles and peduncles of most populations were sparsely or densely villous, although some populations had both glabrous and pubescent petioles and peduncles on the same plant. No population was completely glabrous, although a population at Crows Nest NP in Qld was almost completely glabrous, with scattered pilose hairs only on the cordate lobes of the adaxial leaf surfaces. The pubescence on the abaxial surface of smaller leaves on the type specimens is considered tomentose, but larger leaves are considered densely pubescent. Based on our observations, we conclude the absence of hairs on leaves, petioles, and/or peduncles is not taxonomically useful. However, the presence of dense pubescence on leaves, petioles, and/ or peduncles can be helpful for identifying V perreniformis if appropriate leaf shapes and flower colours are also present. When leaves of V perreniformis are pubescent, the margins are normally ciliate, as in V banksii. Viola silicestris : Leaves of this species were described as glabrous or occasionally sparsely short-pubescent when young (Thiele & Prober 2006). Our observations of populations in southeast Qld and northern NSW are similar. The basal margins of young leaves on plants in Nightcap NP, NSW, occasionally had marginal cilia, but were otherwise glabrous. Many leaves on the isotype at BRI are ciliate C Thiele 2568 [BRI]). Flowers. Petal movements and petal twisting : The two upper petals of the four taxa are sometimes described as “petals erect or reflexed”. Reflexed upper petals and twisted lateral petals are common traits among the four taxa. Both authors have independently observed that petals of Viola banksii reorient themselves into a ‘sleep’ mode at dusk, then ‘reopen’ the next day. For example, as dusk approaches, the two completely reflexed upper petals of V banksii flowers fold forward over the front of the flower (a movement of c. 180°) and the lateral petals ‘untwist’ and extend straight out in front of the flower. In the morning of the next day, the upper petals reflex backward (another c. 180°) from their forward position and the lateral petals spread back laterally to their normal daytime position and become twisted. Thus, the upper petals are ‘erect’ only for a brief period twice a day 92 Fig. 8. V. perreniformis. Erect upper petals; all petals pale blue-violet with violet-coloured blotches on upper and lateral petals. Moogerah Peaks NP, Qld. 2013. Photo: G. Leiper. (Fig. 8), at least for the first few days after the flower first opens. Initial observations indicate that after several days, the petals no longer exhibit these nyctinastic movements. The first author and W. Cherry (pers. comm.) have independently observed nyctinastic petal movements in flowers of V silicestris. This phenomenon is not known to have been previously documented in section Erpetion and its occurrence elsewhere in the section is unknown. Flower colours : Flowers of Viola perreniformis and V silicestris have been described as “concolorous” whereas flowers of V banksii and V hederacea have been described as “discolorous” (=bicoloured). Concolorous is also applied to the flowers of V cleistogamoides, V. fuscoviolacea and V sieberiana. The term concolorous in reference to the overall appearance of a flower is useful as an initial assessment to determine the species at hand, e.g., V Austrobaileya 9(1): 80-101 (2013) perreniformis versus V silicestris. However, because the face of V perreniformis flowers often has darker coloured blotches on the upper and lateral petals, darker veins on the lowest and sometimes all petals, and because the petal bases are white, the description as ‘concolorous’ is sometimes not so clear cut. The description of flower colours on collection labels can assist with taxon identification, but can be problematic for future researchers if ambiguous. For example, flower colours recorded for specimens we would consider Viola perreniformis based on leaf shape include the following descriptions: flowers pale blue; flowers purple; flowers sky blue; flowers violet with darker purple marks; flowers white and purple streaked; lilac-white flowers; mauve and white petals; violet flowers. None state whether the petal tips are white or not. Because the patterns of colouration on the face of flowers in the four taxa can be complex and because colours facilitate identification, we suggest 1) that names for flower colours, veins, and blotches be based on a readily available standard, and 2) that separate descriptions be recorded by the collector for the colour, presence or absence of darker coloured blotches, and presence or absence of darker coloured veins, for the upper and lateral petals, and for the lower (anterior) petal. For all petals, whether the tips are white or not should also be recorded. The colours on petals of Viola flowers fade rather quickly on herbarium sheets and if not quickly pressed flat, the petals shrink and become distorted. These factors can make it difficult or impossible to determine the original colour scheme and/or petal shape on herbarium specimens. For example, within a month after collecting V perreniformis and V silicestris flowers, the distal portions of the petals, which had been pale blue when fresh, had faded in the press to a whitish colour. Viola banksii and V hederacea subsp. hederacea : Flowers of V banksii were described as strongly discolorous violet- and-white (Thiele & Prober 2003). Flowers of V hederacea were described as usually discolorous violet-and-white (occasionally concolorous pale violet or almost white), 93 Little & Lei per, Viola perreniformis the colours usually not strongly demarcated {ibid). In general we concur with these descriptions, although some populations in Qld that appear to be V banksii based on leaf shape have flowers that are not strongly demarcated and other populations that appear to be V hederacea have flowers that appear to strongly demarcated. Among the possible explanations is that a greater range of variation needs to be recognized, or perhaps new taxa are involved. Viola perreniformis: Adams noted that petal colours of V perreniformis were pale (lazuline) blue-violet. We concur and have observed the abaxial surfaces of the five petals (Fig. 9) and portions of the adaxial surfaces (i.e., the ‘face’) are usually a pale blue-violet colour (Figs. 8, 10). The adaxial surfaces often have distinct purple blotches of various sizes on the lateral petals and occasionally basally on the upper petals (Fig. 8). In addition, faint to very distinct violet- coloured veins are often present on both surfaces of the lateral petals and lowest petal. The proximal portions on the adaxial surface of the upper and lateral petals are usually white, although it may be difficult to detect in some flowers without manually spreading the petals. The lowest petal is white proximally, blue-violet distally, and always has distinct, darker, violet-coloured veins (nectar guides). We observed substantial variation in the amount of pigmentation present on the ‘face’ of V perreniformis flowers at Crows Nest NP, Moogerah Peaks NP and Nightcap NP. Viola silicestris : Flowers were described as concolorous pale mauve-blue (rarely somewhat discolorous) (Thiele & Prober 2006), similar to our observations. At Springbrook NP, the authors found populations growing on rhyolite which had completely white flowers. These flowers did not have any darker coloured veins (i.e., no nectar guides) abaxially or adaxially on the upper, lateral, or lower petals. Venation (nectar guides) on lowest petal: Venation of the lowest petal was determined to be a useful taxonomic character for distinguishing V banksii , V hederacea , and V silicestris (Thiele & Prober 2003, 2006). We found general congruence with the venation Fig. 9. Abaxial surface of V perreniformis flowers wtih concolorous petals. Note the green and white indurate area at base of lowest petal (arrows). Crows Nest NP, Mar 2013. Photo: G. Leiper. Fig. 10. V perreniformis. Upper petals fully reflexed, lateral petals twisted. Crows Nest NP, May 2013. Photo: G. Leiper. pattern described for these species in some populations, but not in others. Green and white indurate area: A small, 1-2 mm long, 1-2 mm wide, green and white area is located at the proximal end of the lowest petal of the four taxa (Figs. 9, 11). A narrow, white, ere scent-shaped zone usually occurs between the coloured part of the petal and the green area. A similar green and white area has been reported in V eminens and is referred to as a “small green blotch” or “V-shaped blotch” in these species (Thiele & Prober 2003,2006). V cleistogamoides also has a green and white area on the lowest petal (Thiele & Prober 2006), but V fuscoviolacea does not (Thiele & Prober 2003). No information is available for V. sieberiana. In addition to the colour being distinctly different from the rest of the petal, 94 Fig. 11. V banksii, face view, lateral and upper petals removed. Yellow arrows point to anther glands. Green and white indurate area (not in focus) in lowest petal indicated by white arrow. Photo. J. Little. we found the tissue in the green and white area in V banksii and V perreniformis flowers to be slightly ‘harder’ (indurate) compared to the rest of the petal, a feature not previously reported for flowers in section Erpetion. In flowers of V banksii , V hederacea, and V perreniformis , we also observed a small green area at the base of the two lateral petals similar in colour to the indurate area on the lowest petal, a feature not previously reported in other species in section Erpetion. It has not been determined if the green area on the lateral petals is indurate compared to the rest of the petal. Although it may be presumed that the green and white indurate area in some way facilitates pollination and/or pollinator attraction, its purpose or function is unknown. When a flower is held up to a light source, the centre of the indurate area is translucent, perhaps functioning as a ‘window’ for the benefit of a pollinator. The lowest petal of most Viola flowers has the same texture throughout their length. For example, an indurate area at the base of the lowest petal was not reported for any of the 96 Viola species in China (Chen et al. 2008), nor any of the 73 Viola species in North America (Little & McKinney in press), and may be Austrobaileya 9(1): 80-101 (2013) unique to species in section Erpetion. Style : The style of the four taxa is similar: a thin, straight, ± terete structure, except angled downward near the distal end toward the lowest petal; the style head is glabrous. The base of the style in the four taxa is geniculate near the attachment to the ovary, similar to many Viola species. The geniculate base facilitates the style shaft being easily moved when ‘pushed aside’ by insects probing for nectar or pollen. The geniculate portion of the style functions as a spring, allowing the style to be easily moved by the pollinator. After the pollinator withdraws, the style returns to its original position. In some Viola species in Europe, when the style is lifted up by a probing insect the anthers are spread apart and pollen is released (Beattie 1974). Anther glands: The term ‘anther gland’, apparently first used by Thiele & Prober (2003), is an accurate and appropriate name for the glands found in the four taxa. Compared to the appendage and nectary found in many Viola species, the morphology and presentation of the anther glands in the four taxa are totally different. A few differences are highlighted. First, because the flowers have no spur, the anther glands are not protected within one; second, the anther glands develop laterally from and do not extend beyond the base of the anther; third, the nectaries in most Viola species are situated adjacent to each other whereas in the four taxa the anther glands are widely separated. In his description of the genus Viola , Adams (1982) stated that the anterior two anthers have “basal appendages” i.e., outgrowths from each anther, the distal portion of which secretes nectar. Although correct for species in which the two nectaries extend into a spur, e.g. V betonicifolia and V caleyana , it is inaccurate for the four taxa in section Erpetion because the anther glands are never longer than the anther and the nectaries are not enclosed in a spur. When a flower is viewed face on with a hand lens, the two anther glands in flowers of the four taxa are seen ‘hanging down’ like landing gear on a plane (Fig. 11, 12). Remarkably, because the anther glands are not protected within the confines of a 95 Little & Lei per, Viola perreniformis spur, they are ‘exposed’ to the elements and potentially, it would seem, to physical damage from pollinators. Because the anther glands are situated above and immediately adjacent to the indurate portion of the lowest petal, one might assume that nectar would pool in the concavity formed by indurate area. However, we have not observed nectar in fresh flowers of the four taxa and are not aware of data demonstrating that nectar is produced in these species. In photographs of anther glands from freshly collected flowers of Viola banksii, V hederacea and V perreniformis , the glands were often ‘glistening,’ but obvious liquid nectar was not present. The lack of observations of nectar is not, of course, evidence that none is produced. Perhaps because they are not protected within a spur, the anther glands in section Erpetion have a different function other than to produce nectar. Pollen has been observed adhering to the anther glands in the four taxa presumably due to secretion of a viscid exudate. The shape, colour, and surface texture of the anther gland is a useful diagnostic feature in section Erpetion (Thiele & Prober 2003). Table 3 summarizes information for anther glands reported in the literature. Table 4 summarizes preliminary observations made by the authors. The green colour of glands in some species is very similar to the colour of the indurate area of the lowest petal. The glands of some species are reported as pale or dull green or whitish. However, it is unknown if the colour is initially green and transitions to white or whitish over time, or vice-versa, or if some flowers typically produce only one colour, or if the gland is sometimes or always ± bi-coloured (e.g. white or pale distally, green proximally). A similar situation exists for gland surface texture. In a brief comparison of newer versus older flowers of Viola banksii , the first author found that glands of both newer and older flowers were white or whitish and both were rugose. However, the glands of this species have also been reported as pale green and smooth (Tables 3 & 4; Fig. 12). The anther glands of Viola silicestris were described as “irregular, narrow” and an illustration shows them to be sinuous (Fig. Fig. 12. V. banksii. Lateral view of one of the two anther glands (arrow). Lateral and upper petals removed. Photo: J. Little. 1 in Thiele & Prober 2006). However, in a population we sampled in Qld, the glands were prominent and curved, not narrow and sinuous. We believe that differences reported for shape, surface texture, and colour among the four taxa suggests that additional observations are needed. V hederacea sometimes has purplish glands (Thiele & Prober 2003). A pure white flower of V silicestris from Springbrook N.P was found without an anther gland on either of the two lower anthers. A critical survey is needed of the morphology and development of anther glands, the role they play in reproductive biology, and a standard terminology developed for species in section Erpetion. Pollination and cleistogamy. With certain exceptions, most species in the section have relatively large, showy flowers. Comparably sized flowers of Viola species in the northern hemisphere attract a wide range of insect pollinators such as solitary bees, bee- flies, syrphid flies and various Lepidoptera including butterflies and skippers. Beattie (1971) documented 43 different insect species in a Viola pollination study in England. The authors have not observed potential pollinators visiting flowers of any of the four taxa, although pollination of chasmogamous flowers is assumed to occur based on the many developing and dehisced capsules present in most populations and the fact that seeds are produced. 96 The fully reflexed upper petals of the four taxa could be an adaptation that facilitates sternotribic pollination by providing a platform on which insects can land. Sternotribic pollination occurs when an insect enters a flower ‘upside down/ in contrast with nototribic pollination where the insect enters the flower in a supine position (i.e. ‘right side up’) (Beattie 1974). Typical Viola pollinators seek nectar and/ or pollen, although some are only after pollen. The authors have observed copious amounts of pollen in fresh flowers of each of the four taxa. A pollination study conducted with two Viola species in Brazil provided evidence that both species may be shifting from flowers that typically produce nectar, to flowers that provide primarily pollen (Freitas & Sazima 2003). The first author has observed yellow-coloured thrips (Thysanoptera) with pollen adhering to their bodies in V banksii flowers, randomly and rapidly moving over and within dehisced anther sacs and around Austrobaileya 9 ( 1 ): 80-101 ( 2013 ) the ovary. Thrips were reported in Viola flowers (Baker 1935) and the first author has also observed them on flowers of several Viola species in California. Although thrips are often observed with pollen attached to them, there is currently no evidence they play a role in pollination of Viola. However, due to numerous observations of thrips in Viola flowers, it seems highly probable they could periodically affect pollination. Cleistogamy occurs in many species of Viola throughout the world and is interpreted as a mechanism to help ensure survival when pollinators are unavailable. In North America, c. 85% of all species are known to produce cleistogamous (CL) flowers; only nine species are known to never produce CL flowers (Little & McKinney in press). In Asia, CL flowers are very common in most Viola species (J. Leng, pers. comm.). The presence or absence of CL flowers is not mentioned in Australian floras for any species of Viola. The authors have not observed CL flowers in the field or Table 3. Summary of anther gland characters reported in the literature for Viola Species/Source Description Surface Texture Colour V banksii 1 long, narrow and high, scarcely flattened smooth pale green to whitish V hederacea subsp. hederacea 1 shorter than the anther cells, broad at the base and each distinctly flattened or depressed towards the other; short, rugose, often purple irregularly rugose purplish or dull green V silicestris 2 slightly shorter than the anther cells, irregular, narrow (Not reported) pale 'Thiele & Prober 2003. 2 Thiele & Prober 2006. Little & Lei per, Viola perreniformis Table 4. Summary of anther gland characters recorded by authors 97 Species/Source Description Surface Texture Colour V banksii Springbrook NP, Qld 1 anther gland prominent, ‘talk or ‘high’, almost as long but not as wide as anther, wider proximally, slightly curved, ‘top’ of gland ± flat smooth pale green basally, ± greenish-white distally Venman Bushland, NP, Qld anther gland not prominent, short, c. !4 length of and not as wide as anther, gland height ‘short’, not ‘tall’ or ‘high’, appears ± ‘flat’ entire length of gland smooth white or whitish with a greenish tinge Plant sold in U.S.A. as “V. hederacecT anther gland prominent, ‘tall’ or ‘high’ almost as long but not as wide as anther, much wider proximally, curve, ‘top’ of gland ± flat rugose white or whitish or pale green entire length V hederacea subsp. hederacea Springbrook NP, Qld anther gland prominent, almost as long and about as wide as the anther, gland is wider proximally, arched (curved), taller distally at lower magnifications, the gland appears irregularly rugose; at higher magnifications the surface appears smooth; the uneven, rugose appearance appears to be caused by clumps of pollen grains adhering to the glandular exudate; the normally white pollen grains then take on a greenish colour pale green basally, ± whitish distally; sometimes glistening when fresh V perreniformis anther gland prominent, c. Vz-% as long as anther smooth pale green V silicestris Springbrook NP, Qld anther gland prominent (on a violet-tinged anther), almost as long as, but narrower than the anther, proximal portion curved toward other gland not determined yellowish to pale ‘Possibly a horticultural escape. 98 on herbarium specimens of the four taxa, nor on V sieberiana. Cleistogamous flowers have not been observed on herbarium specimens of V cleistogamoides or V fuscoviolacea (the authors have not observed these species in the field). However, detecting CL flowers on herbarium specimens is difficult because CL flowers can appear similar to undeveloped or aborted chasmogamous flowers. Viola species exhibit numerous strategies that promote reproductive fitness, one of which is hybridization that results in the formation of amphiploids, diploids, and tetraploids. As the authors have found and others have reported (Thiele & Prober 2006), some of the four taxa and other species in section Erpetion are sympatric. For example, in NSW sympatric populations occur with V perreniformis and V silicestris ; V hederacea and V silicestris ; and V fuscoviolacea , V sieberiana and V. silicestris ; and in Qld, V hederacea and V silicestris. However, among the five species mentioned, no hybrids have yet been found (Thiele & Prober 2003, 2006). An example demonstrating that hybridization can potentially occur in section Erpetion was reported in Victoria between V eminens and V fuscoviolacea when an apparently sterile, but vegetatively vigorous putative hybrid, V x zophodes , was found (Thiele & Prober 2003). Adams (1982a) noted in the key to subspecies of Viola hederacea that subsp. fuscoviolacea L.G.Adams (=V fuscoviolacea) was faintly scented. No other information on floral odours of species in section Erpetion is known to the authors. Seeds and seed dispersal. Seeds of the four taxa, like most Viola species, have an ovoid outline, a shape that is assumed to facilitate ballistic travel after ejection from a capsule valve. The seeds of many Viola species possess an outgrowth, called an elaiosome or caruncle that develops around and extends out from the micropyle. Elaiosomes are fleshy food bodies of variable size that can be attractive to ants. Although the number of seeds examined in this study was limited, an obvious elaiosome was not detected on mature, recently collected seeds of three of the four taxa nor on V sieberiana (seeds not Austrobail'eya 9(1): 80-101 (2013) available for V banksii ). Seed characters reported in the literature and observed by the authors for the four taxa and V sieberiana are summarized in Table 5. The seeds are small, 1-2 mm long, although V banksii seeds are sometimes up to 2.5 mm long (Thiele & Prober 2003). While the first author was studying seeds of V sieberiana , initial observations suggested that mature seeds were reddish-brown, until shiny black seeds were found. At least for this species, it appears that a reddish-brown colour transitions into black. The apparent lack of an elaiosome in three of the four taxa and Viola sieberiana might lead one to suspect their seeds are not dispersed by ants. However, seeds of a Viola species in Europe with almost no elaiosome were reported to be removed by ants (Beattie & Lyons 1975). Lengyel etal. (2010) estimated that over 70% of Violaceae species throughout the world are myrmecochorous. Based on numerous observations of dehisced capsules split into three valves on herbarium specimens and in the field, the dispersal mechanism of seeds from capsules in the four taxa and other species in section Erpetion is assumed to be similar to V betonicifolia (Little & Leiper 2012). However, studies are needed to determine ballistic dispersal distances and to determine if ants play any role in dispersing seeds. Taxonomic challenges There is considerable evidence that Viola populations exhibit character divergence among local colonies of single species (Beattie 1976). It should therefore not be too surprising to find morphological variation among isolated populations of species in section Erpetion. Thiele & Prober (2003) noted there are several undescribed species in the V hederacea complex in NSW. Recognition of new taxa may help resolve situations where observed character states in a population conflict with the current taxonomy of the complex. Additional study of the four taxa may reveal other morphological characters or relationships between characters that provide additional or better approaches for taxonomic Little & Lei per, Viola perreniformis Table 5. Seed characters of five species in Viola section Erpetion 99 Species/ Location & Source Mature Seed Colour Surface Texture Seed Length V banksii 1 * * * V glossy purplish-black ± smooth to irregularly rugose 1.8-2.5 mm V hederacea subsp. hederacea 1 dull, mottled cream and brown (occasionally uniformly reddish-brown) ± smooth 1.2-2.0 mm Springbrook NP 2 reddish-brown warty-rugose 1.7-1.8 mm V perreniformis 2 Nightcap NP, NSW shiny or dull dark brown mottled with patches of light brown or tan uneven to irregularly rugose 1-1.2 mm V sieberiana 2 shiny black warty-rugose 1.5-1.8 mm V. silicestris 3 glossy purplish-black not reported 1.5-1.7 mm Springbrook NP 2 shiny black warty-rugose 1.6-1.7 mm ‘Thiele & Prober 2003. 2 Current authors. 3 Thiele & Prober 2006. recognition. Relationships in section Erpetion are largely unknown and will probably require DNA sequencing and chromosome counts to elucidate taxonomic relationships (Thiele & Prober 2006). The provisional keys provided herein are based on the authors’ observations and information in Thiele & Prober (2003, 2006). Among the four taxa, the flowers of Viola perreniformis are most similar to V silicestris ; flowers of V banksii are most similar to V hederacea. Fully mature leaves of V banksii are in general more similar to Vperreniformis , whereas leaves of V hederacea are in general more similar to V silicestris. Provisional Key to selected species in Viola section Erpetion (based on flowers and seeds) 1 Abaxial surfaces of all petals usually of one colour (concolorous), entire surface pale violet, pale blue, or pale lavender, whitish or pale coloured proximally; on adaxial surface (face), c. half of upper and lateral petals pale violet or pale blue distally; (abaxial and adaxial surfaces of V silicestris rarely mostly or entirely white); lowest petal obovate to oblong.2 1. Abaxial surface of all petals usually of two colours (bi-coloured, or discolorous), petals white distally and pale violet proximally (c. 2/3 of upper petals white distally, pale violet proximally) on adaxial surface (face) c. 2/3 of upper petals white distally, violet proximally; c. half of lateral petals white distally, violet proximally; lowest petal obovate, elliptic, or circular.3 100 Austrobaileya 9 ( 1 ): 80-101 ( 2013 ) 2 Anthers cream-coloured; fully mature seeds shiny or dull dark brown, 1-1.2 mm long.V. perreniformis 2. Anthers violet-tinged; fully mature seeds shiny black, 1.5-17 mm long ... V. silicestris 3 Anther gland smooth or rugose, white or whitish to pale green; the midvein and the two adjacent veins on lowest petal usually prominent and distinct, often without interconnecting 2° veins; the violet and white areas on petals usually strongly demarcated.V. banksii 3. Anther gland usually rugose, dull green or purplish; the midvein on lowest petal often less distinct than the two adjacent veins, often with several to many 2° veins connecting them; the violet and white areas on petals usually not strongly demarcated.V. hederacea Provisional Key to selected species in Viola section Erpetion (based on mature, fully developed leaves) 1 Leaf base usually cordate, sinus and lobes usually present, sometimes overlapping, blades about as wide as long (< 2x wider than long), blade outline ovate, orbicular, suborbicular, or semi-circular, blade shape orbicular, reniform, or ovate.2 1. Leaf base usually truncate, sinus and lobes not present, blades usually much wider than long (> 2x wider than long), blade outline semi-circular or ± rectangular, blade shape broad-reniform or sub-reniform (but lobes not prominent), or falcate (with the upper and lower margins parallel).3 2 Leaf margin not crenate from basal lobes to apex, margin nearly entire, sometimes ± scalloped or sinuous, outline usually orbicular, blade shape orbicular or reniform, base cordate, lobes often overlapping in fresh material, apex ± truncate, rounded or obtuse, occasionally ± dentate, the midvein usually not extending much beyond the length of adjacent veins, both leaf surfaces glabrous or with scattered, short hairs, never tomentose, leaf blades about as wide as long.V. banksii 2. Leaf margin regularly or irregularly crenate from basal lobes to apex, outline ovate, orbicular, suborbicular, or semicircular, blade shape reniform, sub-reniform, or ovate, base cordate, lobes seldom overlapping in fresh material, apex usually acute, not truncate, midvein usually extending beyond length of adjacent veins, both leaf surfaces often softly hairy, occasionally tomentose, rarely sub-glabrous; leaf blades usually slightly longer than wide.V. perreniformis 3 The distal (upper) blade margin ± entire or with only a few short points; stems not known to occur at stolon nodes.V. hederacea 3. The distal (upper) blade margin ± serrate or subserrate, sometimes denticulate, occasionally ± crenate; apex often truncate, sometimes rounded; stems (1.5-40 cm) sometimes present at stolon nodes .V. silicestris 101 Little & Leiper, Viola perreniformis Acknowledgements The authors thank the curators at BRI, CANB, and NSW for access to specimens and assistance. We thank RI. Forster for encouragement to prepare this paper and K.R Thiele for constructive comments on an earlier draft. The assistance of B. Lepschi, Curator, Australian National Herbarium, is greatly appreciated for providing images of the holotype of Viola hederacea subsp. perreniformis and scans of photographs of the type of V hederacea. Thanks to P. Bostock for translating the original Latin description and for scanning herbarium specimens. Thanks to J. Brock (Toowoomba, QLD), B. Neilly (Hurstville, NSW), and R. Ollerenshaw (Canberra, ACT) for timely processing of collection/export permits in early 2013 for the first author. References Adams, L.G. (1982a). Appendix. Violaceae. In A.S. George (ed.). Flora of Australia 8: 386. Australian Government Publishing Service: Canberra. Adams, L.G. (1982b). Viola. In A.S. George (ed.). Flora of Australia 8: 91-100. Australian Government Publishing Service: Canberra. Baker, M.S. (1935). Studies in western violets. I. Madrono 3: 51-57. Beattie, A.J. (1971). Pollination mechanisms in Viola. New Phytologist 70: 343-360. - (1974). Floral evolution in Viola. Annals of the Missouri Botanical Garden 61: 781-783. - (1976). Plant dispersion, pollination and gene flow in Viola. Oecologia 25: 291-300. Beattie, A.J. & Lyons, N. (1975). Seed dispersal in Viola (Violaceae): adaptations and strategies. American Journal of Botany 62: 714-722. Chen, Y., Yang, Q., Ohba, H. & Nikitin, V. (2008). Violaceae. In Flora of China 13: 72—111. eFloras 2008. Published on the Internet http:// www.efloras.org Missouri Botanical Garden: St. Louis, MO & Harvard University Herbaria: Cambridge, MA. Accessed 28 May 2013. Duretto, M.F. (2009). Violaceae. In M.F. Duretto (ed.). Flora of Tasmania Online. Tasmanian Herbarium, Tasmanian Museum & Art Gallery: Hobart, www.tmag.tas.gov.au/floratasmania. Accessed 28 May 2013. Electronic Flora of South Australia (2007). http:// www.flora.sa.gov.au/cgi-bin/speciesfacts_ display. cgi?form=speciesfacts&family=&gen us=viola&species=&iname=&submit=Display Accessed 28 May 2013. Elliot, R. & Jones, D. (2010). Encyclopaedia of Australian plants suitable for cultivation 9: 441. Lothian: Sydney. Entwisle, T.J. (1996). Violaceae. In N.G. Walsh & T.J. Entwisle (eds.), Flora of Victoria 3: 361-370. Inkata Press: Melbourne. Freitas, L. & Sazima, M. (2003). Floral biology and pollination mechanisms in two Viola species - from nectar to pollen flowers? Annals of Botany 91: 311-317. James, T.A. (1990). Violaceae. In G. J. Harden (ed.), Flora of New South Wales 1: 435-441. Royal Botanic Garden: Sydney. - (2013). Family Violaceae, In PlantNET - The Plant Information Network System of The Royal Botanic Gardens and Domain Trust, Sydney, Australia (version 2.0). http://plantnet. rbgsyd.nsw.gov.au Accessed 28 May 2013. Lengyel S., Gove, A.D., Latimer, A.M., Majer, J.D. & Dunn, R.R. (2010). Convergent evolution of seed dispersal by ants, and phylogeny and biogeography in flowering plants: A global survey. Perspectives in Plant Ecology, Evolution and Systematics 12: 43-55. Little, R.J. & Leiper, G. (2012). Capsule dehiscence in Viola betonicifolia Sm. (Violaceae). Austrobaileya 8: 624-633. Little, R.J. & McKinney, L.E. In press. Violaceae. In Flora of North America Editorial Committee (eds.). Flora of North America North of Mexico , Vol. 6. Flora of North America Association: New York & Oxford. Stanley, T.D. & Ross, E.M. (1983). Flora of South¬ eastern Queensland 2: 99-101. Queensland Department of Primary Industries: Brisbane. Thiele, K.R. & Prober, S.M. (2003). New species and a new hybrid in the Viola hederacea species complex, with notes on Viola hederacea Labill. Muelleria 18: 7-26. - (2006). Viola silicestris , a new species in Viola section Erpetion from Australia. Telopea 11: 99-104. Pterostylis caligna M.T.Mathieson (Orchidaceae), a new species from northern Queensland M.T. Mathieson Summary Mathieson, M.T. (2013). Pterostylis caligna M.T.Mathieson (Orchidaceae), a new species from northern Queensland. Austrobaileya 9(1): 102-106. Pterostylis caligna is described as new. It is a tuberous, terrestrial herb known only from the Hann and Windsor Tablelands in northern Queensland and is readily distinguished from any other members of the genus by its unique combination of characters. The species is illustrated and diagnosed herein. Its conservation status is assessed and a status of Vulnerable is recommended. Key Words: Orchidaceae, Pterostylis , Pterostylis caligna , Australia flora, Queensland flora, new species, taxonomy, conservation status. M.T.Mathieson, Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4006, Australia. Email: Michael.Mathieson@science.dsitia.qld.gov.au Introduction The greenhood orchids of Australia are diverse and occur in many habitats throughout eastern and southern Australia (Janes & Duretto 2010; Jones & Clements 2002) with a number of new Pterostylis species having been described in recent years (Jones 2009a, 2009b, 2010a, 2010b; Jones & French 2012). Nine species are currently known from the Wet Tropics of north Queensland (Jones 2006, 2010a; Bostock & Holland 2010). During a visit to the Hann Tableland, west of Mareeba in north Queensland in May 2010, the author discovered a population of Pterostylis R.Br. that did not match any named species. However, one specimen at BRI corresponded with this species. Further material was collected in March 2012 and April 2013, enabling its distinctiveness to be confirmed and its formal description herein. I follow the classification of Janes & Duretto (2010) under which this species keys to Pterostylis subg. Pterostylis sect. Foliosae. Materials and methods This research is based on a study of herbarium specimens and associated spirit collection at BRI and CNS, and field studies at the type Accepted for publication 5 September 2013 and other noted localities. All measurements have been made from live material or material preserved in spirit. Abbreviations in the specimen citations include NP (National Park). RE refers to Regional Ecosystem, descriptions of which can be viewed at (in this case): http://www. ehp.qld.gov.au/ecosystems/biodiversity/ regional-ecosystems/index.php Taxonomy Pterostylis caligna M.T.Mathieson sp. nov. ab omnibus aliis speciebus Pterostylis sepalis lateralibus clavatis, galea decurva apice sepali dorsalis filiformis clavati, labello semper abscondito et apice suo rotundato vel obtuse acuto distat. Typus: Queensland. Cook District: Hann Tableland National Park, NW of Mareeba, 8 April 2013, M.T. Mathieson MTM1438 & P.I.Forster (holo: BRI [1 sheet + spirit]; iso: CNS, distribuendi ). Tuberous, terrestrial herb growing in clonal colonies. Tubers globose to ovoid, 6-9 mm long, 5-9 mm wide when fertile. Leaves 3-5 in a rosette encircling the base of the scape, mid-green; lamina linear-ovate to oblong- ovate, 14-34 mm long, 9-19 mm wide, margins entire or occasionally crisped, apex obtuse to bluntly acute; petioles 3—13 mm long, slender, not winged. Scape 42-100 mm Mathieson, Pterostylis caligna tall, 0.9-1.2 mm wide, glabrous; stem bracts, sterile 1 (rarely 2), fertile 1, 8-15 mm long, 2.5-5 mm wide, lanceolate, acute, sheathing at base, margins entire. Flower solitary 17-23 mm long, erect, translucent white with pale to bright green stripes on the galea and sinus. Galea slightly gibbous at the base, then erect before curving forwards in a semicircular fashion with the tip decurved. Dorsal sepal translucent white with bright green stripes, three narrow inner stripes and two broad marginal stripes which coalesce in the distal quarter, 15-20 mm long, 6-9 mm wide, inflated at the base and tapered beyond to the apex with a slight constriction in the proximal third, apex filiform, clubbed, 2.5-3 mm long. Lateral sepals erect closely embracing the galea; sinus protruding slightly to moderately, shallowly curving inwards, the upper margin shallowly notched at the centre then curving upwards; conjoined part 7-10 mm long, 6-8 mm wide, narrowed to c. 2.5 mm at the base, bright green fading somewhat towards the base, the upper margins inrolled, gradually tapered into the free points; free points filiform, minutely clubbed, 8-13 mm long, erect, held upright above the galea. Petals asymmetrically lanceolate, falcate, 13-16 mm long, 3.5-4.5 mm wide, tapered in the distal half, apex acute to subattenuate, largely bright green with a small area of the central proximal section translucent pale green to white; flange 1-1.2 mm wide, broadly deflate, obtuse. Labellum attached to a short basal claw c. 1 mm long, erect, slightly curved in the distal quarter, not visible through the sinus in the set position, green in the proximal half and suffused tan-brown distally; lamina narrowly oblong to narrowly oblong-lanceolate, 5-7 mm long, 1.5-2 mm wide, slightly constricted in the distal half; apex bluntly acute to rounded; central callus slightly raised, running the length of the labellum ending just short of apex, c. 0.25 mm wide tapered to c. 0.1 mm in the centre, green in proximal half, tan-brown distally; basal appendage linear 2-3 mm long, curved, apex irregularly lobed. Column green, brown and white, 6-7 mm long, erect, angled away from ovary at about 40° in the proximal quarter, then erect. Column wings 3-4 mm long; basal lobe 1.5-2 mm long, c. 103 1.2 mm wide, green and brown, at an angle of 50°, apex obtuse, inner margins incurved, sparsely adorned with short, translucent white cilia; mid section 0.8-1 mm long, brown and white; apical lobe, white, linear, 0.6-0.7 mm long, obtuse. Stigma elliptic, 1.5-1.8 mm long, c. 0.75 mm wide, raised, situated centrally on the column. Anther c. 0.7 mm, shortly rostrate. Pollinia linear-clavate, 0.6- 0.7 mm long, yellow, mealy. Ovary cylindrical to slenderly obovoid, 6-11 mm long, 1.5-2 mm wide, green, irregularly ribbed. Capsule narrowly obovoid, 13-18 mm long, 4-5 mm wide, ribbed, smooth. Fig. 1-3. Additional specimens examined : Queensland. Cook District: Hann Tableland NP, NW of Mareeba, May 2010, Mathieson MTM828 & Forster (BRI); Windsor Tableland NP, N of Mt Carbine, Apr 2013, Mathieson MTM1523, 1555 & Forster (BRI); Windsor Tableland, Jan 2009, Sankowsky 3084 & Sankowsky (BRI). Distribution and habitat : Pterostylis caligna is endemic to northern Queensland and has been found only in the Hann Tableland NP, about 20 km north-west of Mareeba and Mt Windsor NP approximately 90 km north- north-west of Mareeba. It grows in simple notophyll vine forest with emergent Agathis robusta (C.Moore ex F.Muell.) F.M.Bailey on granite (RE 7.12.16a) on the Hann Tableland and in simple to complex notophyll vine forest with emergent Agathis robusta (RE 7.12.7a) on the Windsor Tableland between 880 m and 1000 m. At both localities, this species appears to be associated with vine forest- open forest ecotones. Phenology : Flowers and fruits are recorded for January through to April; also in May, when very few flowering plants were observed and most fruit had dehisced. Affinities : Pterostylis caligna stands apart from all other members of the genus by the following characters: clubbed lateral sepals, decurved galea with apex of dorsal sepal filiform and clubbed, labellum always concealed when in the set position and labellum apex rounded or bluntly acute. Sterile rosettes can resemble those of Pterostylis taurus M.A.Clem. & D.L.Jones. 104 A ustrobaileya 9(1): 102-106 (2013) Fig. 1. Pterostylis caligna. A. whole plant *2. B. rosette *2. C. root stock and tuber x4. D. flower from the side x3. E. flower from the front x3. F. undehisced capsule x3. All from Mathieson MTM1438 & Forster (BRI). Del. W.Smith. Mathieson, Pterostylis caligna 105 Fig. 2. Pterostylis caligna. A. lateral sepals (synsepalum) M. B. petal *4. C. column and labellum from the side *8. D. column from the front xl2. E. labellum from above xl2. F. labellum apex x32. G. basal appendage of labellum from above x32. All from Mathieson MTM1438 & Forster (BRI). Del. W.Smith. Conservation status : This species is known from three locations; one within the Hann Tableland NP and two in the Windsor Tableland NP. It was not encountered elsewhere on the Hann Tableland despite extensive searches in similar habitats during three expeditions between 2010 and 2013. It is likely to occur elsewhere on the Windsor Tableland and adjacent ranges in the Wet Tropics biogeographic region given the extent of appropriate habitat that exists. The population at the type locality is estimated to be between 100 and 200 plants occupying an area of less than one hectare. The two populations on the Windsor Tableland were estimated at between 300 and 400 plants, both occupying an area of less than two hectares. Repeated fire events may threaten this species by encouraging the encroachment of introduced, hyperinvasive grasses (viz. giant rat’s tail grass Sporobolus pyramidalis (Lam.) Hitchc., grader grass Theme da quadrivalvis (L.) Kuntze and 106 gamba grass Andropogon gayanus Kunth), the rampant herbaceous weed Praxelis clematidea R.M.King & H.Rob. and lantana Lantana camara L. into its’ ecotone habitats. Applying criteria of the IUCN (IUCN 2001), the recommended conservation status is Vulnerable (Dl, D2). Etymology : From the Greek calignis - of the mists, in reference to the upland cloud forest this species inhabits. Fig. 3. Pterostylis caligna , side view of the flower, photographed in habitat ( Mathieson MTM828). Acknowledgements Collections of this species were made on a Bush Blitz nature discovery expedition in 2010 and in 2012 and 2013 using a Bush Blitz Tactical Taxonomy Grant and Applied Taxonomy Grant respectively, awarded to MTM; all funded by the Commonwealth Government in part. I thank Will Smith for the illustrations, and Peter Bostock for the Latin diagnosis. The following people and organisations are thanked for their assistance that made the survey possible: Keith McDonald (formerly of Austrobaileya 9(1): 102-106 (2013) Dept, of Environment & Heritage Protection (DEHP) - Atherton) for logistical support and co-ordination; Traditional Owners for the Hann Tableland, John and Troy Grainer; Cairns/Mareeba/Mossman National Parks & Wildlife Service staff (Jonathon Roth, Robert Miller, Brendan Malone, Andrew Hedges and Alastair Freeman) and Queensland Herbarium staff (Paul Forster, Megan Thomas, Gordon Guymer, Peter Bostock, Ashley Field, Eda Addicott and Mark Newton) for logistical and field support; Cape York Tenure Unit Georgianna Fien, Eric Wason); Cape York Helicopters and Pat Gorman for safe and efficient helicopter provision. References Bostock, P.D. & Holland, A.E. (eds) (2010). Census of the Queensland Flora 2010. Queensland Herbarium, Department of Environment and Resource Management: Brisbane. Iucn (2001). IUCN Red List of Categories and Criteria: Version 3.1. IUCN Species Survival Commission: Gland (Switzerland)/Cambridge (United Kingdom). Janes, J. & Duretto, M. (2010). A new classification for subtribe Pterostylidinae (Orchidaceae), reaffirming Pterostylis in the broad sense. Australian Systematic Botany 23: 260-269. Jones, D.L. (2006). A complete guide to native orchids of Australia, including the island territories. New Holland: Sydney. - (2009a). Five threatened new species of Hymenochilus (Orchidaceae) from southern Australia. The Orchadian 16: 176-187. - (2009b). Oligochaetochilus pedinus, a new species of Orchidaceae from south-western New South Wales. The Orchadian 16: 214-221. - (2010a). Four new species of Oligochaetochilus (Orchidaceae) from tropical Queensland. The Orchadian 16: 302-311. - (2010b). A new species of Oligochaetochilus (Orchidaceae) from Queensland with affinities to O. woollsii. The Orchadian 16: 370-375. Jones, D.L. & Clements, M.A. (2002). A review of Pterostylis (Orchidaceae). Australian Orchid Research 4: 1-168. Jones, D.L. & French, C.J. (2012). Two new species of Pterostylis from Western Australia. Australian Orchid Review 77(4): 39-44. Two new subspecies of Eucalyptus tereticornis Sm. (Myrtaceae) A.R. Bean Summary Bean, A.R. (2013). Two new subspecies of Eucalyptus tereticornis Sm. (Myrtaceae). Austrobaileya 9(1): 107-113 Two new subspecies of Eucalyptus tereticornis Sm., E. tereticornis subsp. rotunda A.R.Bean and E. tereticornis subsp. basaltica A.R.Bean, are described and illustrated. A key to the four subspecies of E. tereticornis is provided, together with a distribution map for the new taxa. Key Words: Myrtaceae, Eucalyptus , Eucalyptus tereticornis subsp. basaltica , Eucalyptus tereticornis subsp. rotunda , new subspecies, Australia flora, Queensland flora, identification key, taxonomy A.R. Bean, Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: Tony.Bean@science.dsitia.qld.gov.au Introduction Eucalyptus tereticornis Sm. is one of the best known members of the largely eastern Australian ‘Red Gum’ group (E. sect. Exsertaria L.D.Pryor & L.A.S.Johnson ex Brooker), characterised by the reniform cotyledons, concolorous adult leaves, ovules in 6-8 longitudinal rows, and the toothed seed coat. Brooker (2000) placed E. tereticornis in E. ser. Erythroxylon (Blakely) Brooker, along with 14 other species, distinguished by the patchily deciduous bark, leaves with moderately dense reticulation, erect stamens, pyramidal or obliquely pyramidal seeds and axillary inflorescences. Eucalyptus tereticornis is a widespread and variable species; it extends from eastern Victoria to north Queensland, and to Papua New Guinea. Brooker & Slee (2000) documented a number of variants of E. tereticornis : E. tereticornis subsp. tereticornis (long operculum swollen at the base), the ‘Hunter Valley’ form (with short obese opercula, ± parallel-sided), the ‘Tenterfield- Cunningham’s Gap’ form (operculum shorter than typical form and lacking the swollen base), E. tereticornis subsp. mediana Brooker & Slee (from eastern Victoria), and E. blakelyi -E. tereticornis intergrades. Accepted for publication 10 October 2013 Eucalyptus tereticornis subsp. tereticornis is the most widespread subspecies, extending from the south coast of New South Wales to north of Cooktown in Queensland, and Central and Oro (Northern) Districts in Papua New Guinea. In southern Queensland, it extends west to near Roma (Queensland Herbarium 2013). Eucalyptus tereticornis subsp. basaltica A.R.Bean, described here, is a widespread tree on hills and ridges (predominantly basaltic) of southern Queensland and northern New South Wales. The ‘Hunter Valley’ form and ‘Tenterfield-Cunningham’s Gap’ form of Brooker & Slee (2000) are both attributable to this subspecies. An additional taxon, described here as Eucalyptus tereticornis subsp. rotunda A.R.Bean, is a riparian tree of restricted distribution from the upper reaches of the Maranoa River of southern Queensland with large orbicular juvenile leaves. Eucalyptus tereticornis subsp. mediana Brooker & Slee differs from E. tereticornis subsp. tereticornis by the glaucous juvenile leaves, shorter obese operculum, mixed black single-coated and yellow double-coated seeds, and the later flowering period (Brooker & Kleinig 1999). It inhabits stream banks and river floodplains, but also low hills and plains not subject to flooding (Nicolle 2006). 108 Eucalyptus tereticornis subsp. mediana and E. tereticornis subsp. rotunda are geographically isolated from the other subspecies, although the latter occurs in close proximity to an outlying stand of E. tereticornis subsp. basaltica. E. tereticornis subsp. tereticornis and E. tereticornis subsp. basaltica are parapatric in southern Queensland and northern New South Wales, and some hybridisation or intergradation is suspected between the two taxa. Subspecies rank is considered appropriate for the new taxa, as there are consistent characters to separate them from E. tereticornis subsp. tereticornis. Materials and methods This study is based on a combination of field studies in Queensland and northern New South Wales, measurements made during an earlier seedling trial that included Eucalyptus tereticornis subsp. tereticornis and E. tereticornis subsp. basaltica , and an examination of specimens at BRI. E. tereticornis subsp. mediana has not been observed in the field, and knowledge of the taxon has been gleaned from the literature and from herbarium specimens. Measurements have been made from dried herbarium material. Austrobaileya 9(1): 107-113 (2013) Common abbreviations used in the text include Mt (Mountain), S.F. (State Forest). Taxonomy Eucalyptus tereticornis Sm., Spec. Bot. New Holland , part 4, 41 (1795). Type: New South Wales. Port Jackson, 1793, J. White s.n. (holo: LINN; iso: BM ,fide Chippendale [1988: 324]). Tree to 50 m high. Bark smooth throughout or with persistent grey, scaly-fibrous bark at base, forming a stocking less than one- tenth tree height; smooth bark deciduous, somewhat shiny, dappled white, grey, brown or pale blue, shedding in irregular strips less than a metre long. Adult branchlets not glaucous. Seedling leaves broadly ovate to orbicular, stems quadrangular. Juvenile leaves petiolate, alternate, broadly-lanceolate to orbicular, concolorous or slightly discolorous. Adult leaves alternate, narrow- lanceolate to lanceolate or falcate, 8-21 x 1-3.5 cm, concolorous, green, petiolate, lateral veins at 35-55° to the midrib, vein reticulation moderately dense, oil glands abundant, several per areole; intramarginal vein less than 20% of distance from margin to midrib, margins entire, base cuneate. Inflorescence with umbels borne singly in leaf axils or along leafless stems. Individual umbels 7(-9)-flowered; peduncle terete or Key to subspecies of Eucalyptus tereticornis 1 Opercula of mature buds expanded at base; juvenile leaves broadly lanceolate to ovate, 2-3.2 times longer than wide . . . E. tereticornis subsp. tereticornis 1. Opercula of mature buds not expanded at base; juvenile leaves orbicular to ovate, 0.85-2 times longer than wide.2 2 Juvenile leaves orbicular, 0.85-1.2 times longer than wide, 5-16 cm across, green, apex obtuse, apiculate or emarginate . . . . E. tereticornis subsp. rotunda 2. Juvenile leaves broadly ovate to ovate, 1.2-2.0 times longer than wide, dull bluish, apex acute.3 3 Pedicels 1-3 mm long; opercula 1.3-1.6 times longer than broad; new growth of juvenile plants pruinose; margins of juvenile leaves usually undulate; seeds coats a mixture of black and yellow. . . E. tereticornis subsp. mediana 3. Pedicels 2.5-6 mm long; opercula 1.5-2 times longer than broad; new growth of juvenile plants dull green; margins of juvenile leaves not undulate; seed coats all black.E. tereticornis subsp. basaltica 109 Bean, Eucalyptus tereticomis angular, 7-20 mm long. Buds with operculum scar present, stamens erect to slightly sinuate, staminodes absent, filaments white or rarely pink; opercula longer than hypanthium. Stigma blunt or slightly dilated. Ovules in six or more longitudinal rows. Fruits cupular, oblate or hemispherical, 4-7(-8) mm long, 5—8(—10) mm wide, pedicellate. Disc convex, obliquely ascending; valves 3-5, strongly exserted. Seeds pyramidal or cuboid, larger and darker than chaff, 0.8-1.5 mm long, usually black with toothed edges, but sometimes yellow and with smooth edges; hilum terminal. Eucalyptus tereticomis subsp. basaltica A.R.Bean subsp. nov. a subspecie typica operculis quae sunt e basi non expansa, foliis juvenilibus late ovatis (late lanceolatis usque ovatis in subspecie typica) et habitatione clivorum et summorum jugorum differens. Typus: Queensland. Darling downs district: Near Bottle Tree Lookout, Bunya Mountains National Park, 21 May 1996, P. Grimshaw PG2468 & L.M. Grimshaw (holo: BRI [3 sheets]; iso: NSW). Eucalyptus tereticomis ‘Hunter Valley form’ (Brooker & Slee 2000) Eucalyptus tereticomis ‘Tenterfield- Cunningham’s Gap form’ (Brooker & Slee 2000) Eucalyptus tereticomis subsp. (Bunya Mountains P.V. Holzworth AQ397993), Bostock & Holland (2010: 121). Eucalyptus tereticomis subsp. (Consuelo Tableland M.I. Brooker B4880), Bostock & Holland (2010: 121). Tree to 20 m high. Base of trunk sometimes with slabs of persistent grey, scaly-fibrous bark, extending up to 1 metre; bark smooth above, deciduous, somewhat shiny and dappled with white, creamy-white, yellow and brown. Juvenile leaves ovate to almost orbicular, bluish green, 6.5-12.5 x 4-7 cm, 1.2-2 times longer than broad, concolorous; apex acute, base cuneate; margins entire or crenulate, petioles 11-22 mm long. Adult leaves lanceolate to falcate, 90-160 x 12-32 mm, green; apex attenuate; petioles 11-21 mm long. Inflorescences 7-flowered or rarely 9-flowered; peduncles ± terete or obviously flattened, 7-13 mm long; pedicels 2.5-6 mm long; mature buds ovoid to ellipsoid, 9-14 x 5-6 mm; outer operculum shed early, inner operculum cylindrical-oblong to conical, 7.5- 11 x 5-6 mm, 1.5-2 times longer than broad, not expanded at base, apex obtuse or acute; stamens white; outer stamens erect in bud, up to 9.5 mm long. Fruits cupular to oblate, 5-7 mm long (excluding valves), 6-8 mm wide, valves 3-5, pedicels 2-7 mm long. Fig. 1A-E. Additional selected specimens examined : Queensland. Port curtis district: Kroombit Tableland, c. 60 km SW of Gladstone, Amy’s Peak, Jun 1977, Crisp 2824 (BRI). Burnett district: Foothills of Bunya Mountains S.F., Jul 1985, Holzworth s.n. (BRI [AQ397993]); MtMowbullan, Bunya Mountains, Jul 1994, Fensham 1781 (BRI); Bunya Mountains, Mt Kiangarow, Dec 1954, Smith 6261 (BRI). Wide bay district: Woowoonga S.F. 287, 2 km from Mt Goonaneman, c. 12 km NE of Biggenden, Apr 1989, Last s.n. (BRI [AQ455525]); Widgee Mt, Apr 1996, Forster PIF19137 & Leiper (AD, BRI, CANB, MEL, NSW). Maranoa district: 3 km W of the ‘Stock Yard’ on the Consuelo Tableland, Dec 1980, Gunn BG227 (BRI); Consuelo Tableland, May 1982, Neldner & Thomas 683 (BRI). Darling downs district: c. 6.6 miles [10.6 km] N of Yamsion on Bunya Mountains road. May 1961, Smith 11365 (BRI); western slope of Cunningham’s Gap, Jun 1957, Blake 20134 (BRI); c. 4 km along Spicer’s Gap road from the west, at S.F. gate, Oct 1995, Brooker 12417 & Slee (BRI, CANB, MEL, NSW); Spicer’s Gap, Jun 1963, Everist 7285 (BRI); Allora Council Reserve, Sep 2002, Menkins 1LM0122 (BRI). Moreton district: Main Range, between Spring Bluff & Murphy’s Creek, Aug 1930, Hubbard 3530 (BRI); near Kooralbyn Resort, Sep 1998, Larson s.n. (BRI [AQ662937]); Mt Cotton ridge, Oct 2008, Bean 28106 (BRI); WNW ridge of Lost World, Jan 1989, Weston 1368 & Richards (BRI, NSW); Mt Cainbabel, Macpherson Range, May 1930, White 6766 (BRI). New South Wales. Upper Acacia Creek, Jun 1951, Everist 2550 & Webb (BRI); 3.8 km along Middle Creek Road from Cressfield Road, c. 2 km up ridge to west, Sep 1985, Hill 1319 (BRI, NSW); 8.6 km SW of Scone towards Merriwa, Oct 1995, Brooker 12350 & Slee (BRI, CANB); 8.3 km N of Wiseman’s Ferry, W edge of MacDonald River, Oct 1995, Brooker 12340 & Slee (BRI, CANB). Distribution and habitat : Eucalyptus tereticomis subsp. basaltica extends from north-west of Sydney to Kroombit Tops, south¬ west of Gladstone, and there is an outlier on the Carnarvon Range north of Mitchell (Map 1). It is a component of eucalypt woodland, commonly associated with Eucalyptus eugenioides Sieber ex Spreng., E. melliodora A.Cunn. ex Schauer, and E. biturbinata 110 A ustrobaileya 9(1): 107-113(2013) m 1 i t h) j > Fig. 1 . Eucalyptus tereticornis subsp. basaltica A. juvenile leaf x0.5. B. adult leaf x0.6. C. cluster of mature buds x2. D. cluster of mature buds x2. E. cluster of mature fruits x2. E. tereticornis subsp. rotunda F. juvenile leaf x0.5. G. adult leaf x0.6. H. cluster of mature buds x2.1. cluster of mature buds x2. J. cluster of mature fruits x3. A,B,E from Grimshaw 2468 & Grimshaw (BRI); C from Bean 28106 (BRI); D from Larson s.n. (BRI [AQ662937]); F,G,I from Bean 31323 (BRI); H from Bean 31321 (BRI); J from Bean 25804 (BRI). Del. W.Smith. Bean, Eucalyptus tereticomis L.A.S. Johnson & K.D.Hill. It most commonly grows on basalt substrates, in brown to black clayey soil, but may sometimes grow on soils derived from rhyolite or granite, and rarely, serpentinite. Altitudes vary from 100 metres to 1150 metres. Phenology : Flowers have been collected from May to December; fruits may be found all year round. Notes : Eucalyptus tereticornis subsp. basaltica differs from E. tereticornis subsp. tereticornis by the opercula that are not expanded at the base, the broadly ovate juvenile leaves (broadly lanceolate to ovate for E. tereticornis subsp. tereticornis ), and the hillside or ridge top habitat. The buds and leaves of E. tereticornis subsp. basaltica are often smaller than those of the typical subspecies, but there is some overlap. This subspecies was recognised as a distinct taxon by Clifford (1972), who referred to differences in the size of the buds and fruits, and the shape of the juvenile leaves. Conservation status : A widespread and common subspecies. Etymology : The epithet refers to the basaltic substrate upon which this subspecies usually grows. Eucalyptus tereticornis subsp. rotunda A.R.Bean subsp. nov. a subspecie typica foliis juvenilibus orbicularibus apice obtuso emarginatove praeditis (lanceolato usque ovato apice acuto in subspecie typica) et operculis brevioribus quae sunt e basi non expansa differens. Typus: Queensland. Maranoa district: Western Creek, ‘Mt Owen’, c. 140 kmN of Mitchell, 28 November 2006, A.R. Bean 25804 (holo: BRI; iso: CANB, MEL, NSW, distribuendi). A tree to 28 m high. Base of trunk with slabs of persistent grey, scaly-fibrous bark, extending up to 2 metres; bark smooth above, deciduous, somewhat shiny and dappled with white, creamy-white and brown. Juvenile leaves orbicular, 60-180 x 60-160 mm, 0.85- 1.2 times longer than broad, concolorous or slightly discolorous, bright green; apex obtuse or emarginate, rarely apiculate, base cuneate ill or obtuse; margins entire or crenulate, petioles 12-32 mm long. Adult leaves lanceolate to falcate, 90-190 x 18-32 mm, concolorous, green; apex attenuate; petioles 18-30 mm long. Inflorescences 7-flowered; peduncles ± terete, 7-14 mm long; pedicels 2-3.5 mm long; mature buds ovoid to ellipsoid, 7.5-10 x 4.5-5.5 mm; outer operculum shed early, inner operculum cylindrical-oblong to conical, 6-7 x 4.5-5.5 mm, 1.3-1.7 times longer than broad, apex obtuse, acute or umbonate; stamens white; outer stamens erect in bud, though slightly sinuate, up to 5.5 mm long. Fruits cupular to oblate, 4-5 mm long (excluding valves), 5.5-7 mm wide, valves 4 or 5, pedicels 1.5-3.5 mm long. Fig. 1F-J. Additional specimens examined : Queensland. Maranoa district: Western Creek, c. 12 km N of ‘Mt Owen’ Homestead, N of Mitchell, Nov 2011, Bean 31321 & Halford (BRI, CANB, MEL, NSW); ditto loc., Nov 2011, Bean 31323 & Halford (BRI, NSW); 16.2 km from Consuelo Tableland towards Mt Moffatt, Dec 1982, Brooker 7891 (BRI, CANB). Distribution and habitat : Eucalyptus tereticornis subsp. rotunda is endemic to Queensland. It is known from a limited area north of Mitchell, on ‘Mt Owen’ Station, and in the Mt Moffatt section of Carnarvon National Park (Map 1). It grows along creek banks, at altitudes between 550 and 700 metres. Phenology : Flowers have been recorded in November; fruits in November and December. Affinities : Eucalyptus tereticornis subsp. rotunda differs from E. tereticornis subsp. tereticornis by the orbicular juvenile leaves with obtuse or emarginate apex (broadly lanceolate to ovate juveniles with acute apex for E. tereticornis subsp. tereticornis ); the opercula of mature buds not expanded at base, 7.5-10 mm long with obtuse apex (opercula of mature buds expanded at base, 11-17 mm long, with acute apex for E. tereticornis subsp. tereticornis ). Eucalyptus tereticornis subsp. rotunda differs from E. tereticornis subsp. basaltica by the orbicular juvenile leaves with obtuse or emarginate apex (broadly ovate juveniles with acute apex for E. tereticornis subsp. basaltica ), and the riverine habitat. 112 The juvenile leaves of Eucalyptus tereticornis subsp. rotunda are similar in size, shape and colour to those of E. amplifolia subsp. amplifolia , but E. tereticornis subsp. rotunda differs by the mature buds 7.5-10 mm long (10-12 mm for E. amplifolia subsp. amplifolia ); the inflorescences 7-flowered (9-11-flowered for E. amplifolia subsp. amplifolia)., the obtuse opercula (acute for E. amplifolia subsp. amplifolia) and the narrower adult leaves. Notes : Eucalyptus tereticornis subsp. rotunda is found on the upper tributaries of the Maranoa River. The type specimen of E. camaldulensis subsp. acuta Brooker & M.W.McDonald was collected at ‘Forest Vale’, a property between Mitchell and ‘Mt Owen’, also on the Maranoa River. However, E. tereticornis subsp. rotunda and Eucalyptus, camaldulensis subsp. acuta are thought to be allopatric, with occurrences of the latter starting 30-40 km downstream from occurrences of the former. E. camaldulensis subsp. acuta can be distinguished by its bluish, lanceolate juvenile leaves, the fruiting valves set at a considerable angle to the disc, and the seeds with a yellow coat. Eucalyptus tereticornis subsp. rotunda is conspecific with the type of E. populifolia Hook., which is the basionym for E. populnea F. Muell., a name currently applied to the Poplar box. The typification of E. populnea is the subject of a current conservation proposal (Bean 2012). Conservation status : A conservation status of Near Threatened is proposed under Criterion D, i.e. area of occupancy less than 40 square kilometres, and fewer than 10 localities known (IUCN 2001). A minimum of 2000 trees have been observed at Mt Owen. The population size at Mt Moffatt is not known. Etymology : The subspecific epithet is from the Latin rotundus , meaning round. This refers to the shape of the juvenile leaves. A ustrobaileya 9(1): 107-113(2013) Acknowledgements I am grateful to Will Smith for the illustrations and distribution map, Peter Bostock for the Latin diagnoses, and an anonymous referee for helpful comments and suggestions. References Bean, A.R. (2012). Proposal to conserve the name Eucalyptus populnea (Myrtaceae) with a conserved type. Taxon 6\ . 1332-1333. Bostock, P.D. & Holland, A.E. (eds) (2010). Census of the Queensland Flora 2010. Queensland Herbarium, Department of Environment and Resource Management: Brisbane. Brooker, M.I.H. (2000). A new classification of the genus Eucalyptus L’Her. (Myrtaceae). Australian Systematic Botany 13: 79-148. Brooker, M.I.H. & Kleinig, D.A. (1999). Field Guide to Eucalypts , Volume 1 - South-eastern Australia , 2 nd edition. Bloomings Books: Hawthorn, Victoria. Brooker, M.I.H. & Slee, A.V. (2000). Studies in the Red Gums of south-eastern Australia with particular emphasis on Eucalyptus subser. Erythroxyla. Australian Forestry 63: 86-106. Chippendale, G.M. (1988). Eucalyptus, Angophora (Myrtaceae). Flora of Australia Volume 19. Australian Government Publishing Service: Canberra. Clifford, H.T. (1972). Eucalypts of the Brisbane Region. Queensland Museum: Brisbane. Iucn (2001). IUCN Red List Categories: version 3.1. IUCN Species Survival Commission. IUCN: Gland, Switzerland. Nicolle, D. (2006). Eucalypts of Victoria and Tasmania. Bloomings Books: Melbourne. Queensland Herbarium (2013). Specimen label information (HERBRECS), viewed 22 Feb 2013. Bean, Eucalyptus tereticomis 113 Map 1 . Distribution of Eucalyptus tereticomis subsp. basaltica • and E. tereticomis subsp. rotunda A. A new fern species for Queensland: Diplazium squamuligerum (Rosenst.) Parris (Woodsiaceae) Daniel J. Ohlsen 1 & Ashley R. Field 2 Summary D.J.Ohlsen & A.R.Field (2013). A new fern species for Queensland: Diplazium squamuligerum (Rosenst.) Parris (Woodsiaceae). Austrobaileya 9 ( 1 ): 114 - 125 . The taxonomic status of an unknown fern species from the Atherton Tableland, north-east Queensland, hitherto attributed to Asplenium L., was investigated. Phylogenetic analysis of trnL-F and rbcL chloroplast DNA sequences supported classification in Diplazium Sw., a finding supported by closer examination of scale features. Inspection of Diplazium type material determined that the Australian material belongs to Diplazium squamuligerum (Rosenst.) Parris, a species previously described from Papua New Guinea. A thorough description of this species and an amended key to the Diplazium species of Australia are provided. This study highlights the value of molecular study and close inspection of scale features for fern identification. Taxonomic revision in Diplazium is also discussed in light of the findings presented. Key Words: fern, Athyriaceae, Woodsiaceae, Diplazium , Diplazium squamuligerum , Australia flora, Queensland flora. Wet Tropics bioregion, new species record, taxonomy 'D.J.Ohlsen, School of Botany, The University of Melbourne, Parkville, Victoria 3010, Australia. Email: d.ohlsen@student.unimelb.edu.au 2 A.R.Field, Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Toowong, Queensland 4066, Australia &Australian Tropical Herbarium, James Cook University, Cairns Campus, Smithfield, Queensland 4878, Australia. Email: A shley. F ield@science. dsitia. qld. gov. au Introduction The Wet Tropics of north-east Queensland, from Ingham north to Cooktown, contains the highest fern species diversity in Australia. 257 leptosporangiate fern species occur in the region, which is 66% of the total for Australia (McCarthy 1998). While 37 of these are considered to be endemic to this region, the vast majority are widespread being shared with neighbouring tropical islands, particularly New Guinea which shares 56% of tropical Australian leptosporangiate fern species (McCarthy 1998). Several such species are extremely rare in Australia, known only from one or a few small populations. Notable examples ar q AspleniumpeUucidiim Lam. and Hymenasplenium unilaterale (Lam.) Hayata, both restricted to a single known population in Australia (Brownsey 1998). The rarity of some taxa has delayed their discovery until relatively recently in Australia. Lecanopteris sinuosa (Wall, ex Hook.) Copel., Accepted for publication 29 July 2013 known from only three sites in Australia, was described (as Polypodium sinuosa Wall, ex Hook.) from Indonesia in 1863, but was not collected in Australia until 1985 when it was found on Moa Island, Torres Strait. It was later collected on Cape York in 1987, and was formally recognised as an Australian species (Andrews 1990). Entirely new taxa, endemic to north-east Queensland have also gone unnoticed until relatively recently. These include Antrophyum jagoanum D.L.Jones & Bostock, Diplazium bostockii D.L.Jones, and Lastreopsis windsorense D.L. Jones (McCarthy 1998). However, the most recent new fern species confirmed for Australia is Oleandra musifolium Blume. In contrast to the previous cases, populations of this species have been known in Australia for over 60 years, but under the misapplied name of O. neriiformis Cav. (Hovenkamp & Ho 2012). Four fern specimens collected from two areas on the Atherton tableland, in north-east Queensland, hitherto identified as Asplenium sp. indet. (A. sp. ‘RE.I. Road’ in BRI) 115 Ohlsen & Field, Diplazium squamuligerum (Aspleniaceae) on herbarium specimens, also appear to represent a new taxon for Australia (Fig. 1). This taxon is clearly morphologically distinct from the 30 Australian Asplenium species treated by Brownsey (1998). It was first collected in 1983: once from the North Johnstone Logging Area ( Lockyer s.n.), which is thought to be from a population on the west side of Topaz National Park (N.P.) (B.Gray pers. comm.), and twice from Maalan, 20 km further south. The fourth and most recent collection, in 2005 ( Sankowsky & Sankowsky 2637), was from near the end of P.E.I. Road in Topaz N.P, within one km of the suspected location of the Lockyer collection. Determining the taxonomic status of taxa in Aspleniaceae is often fraught with difficulty. Aspleniaceae is the largest fern family with over 700 species occurring worldwide (Kramer & Viane 1990; Smith et al. 2006). This makes verification of identity or novelty challenging for unresolved Australian collections; the putative taxon must be compared with a large number of type specimens from overseas. In addition to the large species diversity in Aspleniaceae, hybridisation and polyploidy are frequently encountered (see Lovis 1977). Hybrids and polyploids are often morphologically distinct from parental lineages (Kramer & Viane 1990), obscuring their origins and giving the impression of completely new lineages. While identification within Aspleniaceae is often difficult, members of the family are generally easy to distinguish from other families. Those in Aspleniaceae are distinguished by linear sori, clathrate scales on the stipes, and stipes which have two C-shaped vascular bundles at the base which unite apically up the stipe to form a ‘butterfly- like’ shape (Kato & Kramer 1990). Species of Aspleniaceae are morphologically most similar to some members of the Woodsiaceae sensu Smith et al. (2006) (=Athyriaceae sensu McCarthy (1998)), which also possess elongate indusiate sori. The majority of these Woodsiaceae species are most easily distinguished from Aspleniaceae by being large terrestrial plants; however, they are consistently distinguished by more subtle features such as differences in the arrangement of vascular bundles in the stipe, and by the possession of non-clathrate scales (Kato & Kramer 1990), which are often only obscurely differentiated from the clathrate scales observed in Aspleniaceae. Two Woodsiaceae genera occur in north-east Queensland: Deparia Hook. & Grev. and Diplazium Sw. sensu lato (including Callipteris Bory). This study sequenced two chloroplast regions, rbcL and trnL-L, of a fresh collection of Asplenium sp. ‘RE.I. Road’, in order to determine this taxon’s affinity amongst the ferns, and to resolve its taxonomic status. Materials and methods Collections of Asplenium and Diplazium in Australian herbaria, and selected type material from extra-Australian herbaria were examined. One specimen of Asplenium sp. ‘P.E.I. Road’ was collected (i Ohlsen 461, BRI, MELU) in the vicinity of the most recent P.E.I. Road collection {Sankowsky & Sankowsky 2637, BRI). DNA isolation, amplification and sequencing. DNA was extracted from 20 mg of silica gel- dried leaf tissue. Leaf tissue was ground using a mortar and pestle with the aid of acid washed grinding sand (Ajax Finechem, Australia). DNA was isolated from ground samples using a DNeasy Plant Mini Kit (QIAGEN, Germany), following the manufacturer’s instructions. DNA was eluted in 100 pL of the supplied elution buffer. The chloroplast DNA markers rbcL and trnL-L were sequenced. These regions were chosen because 1) both are routinely used in fern systematics, enabling comparison of sequences obtained from this species to those of many other species, and 2) effective phylogenetic placement (at least to genus level) can be achieved with both regions. rbcL is a gene enabling comparison across multiple plant lineages, and the trnL-L region contains an intron and intergenic spacer, which have faster mutation rates, enabling differentiation between closely related species and populations. 116 Austrobaileya 9(1): 114-125(2013) Fig. 1. Diplazium squamuligerum (syn. Asplenium sp. RE.I. Road) from Topaz National Park, Atherton Tableland, Queensland (Ohlsen 461 et al. [BRI, MELU]). (a) a mature frond with elongate sori, (b) a fully developed scale from the base of the stipe, (c) D. squamuligerum in habitat (two plants are shown: top left, and bottom beside the creek pool), (d) detail of the rachis and stipe wing, and the protuberances, some with scales developing upon them. Chloroplast DNA markers were amplified by Polymerase Chain Reaction (PCR), performed on a MyCycler thermal cycler (Bio-Rad, USA). Reaction mixtures comprised 5 pL of 5x MyTaq Reaction Buffer containing 5mM of each dNTP and 15 mM MgCl 2 (Bioline, Australia), 50 pg BSA (Thermo Fisher Scientific, Australia), 0.125 pL (0.625 Units) MyTaq DNA Polymerase (Bioline, Australia), 10 pmol of each primer, 2.0 pL of extracted DNA, and distilled water added to make a total volume of 25 pL. No amplification occurred without the addition of BSA. The rbcL gene was amplified using the primers ESRBCL1F (5’-ATG TCA CCA CAA ACG GAG ACT AAA GC-3’) and ESRBCL1361R (5’-TCA GGA CTC CAC TTA CTA GCT TCA CG-3’) (Schuettpelz & Pryer 2007). The trnL intron, trnL 3’-exon and trnL-F intergenic spacer were amplified using the primers F (5’-ATT TGA ACT GGT GAC ACG AG-3’) (Taberlet et al. 1991) and Fernl (5’-GGC AGC CCC CAR ATT CAG GGR AAC C-3’) (Trewick et al. 2002). PCR thermocycling conditions involved an initial denaturation step of 95°C for 1 minute, followed by 33 cycles of 95°C for 1 minute, 55°C for 1 minute, and 65°C for 4 minutes, and a final extension time of 65°C for 5 minutes. DNA concentrations were quantified by electrophoresis against Hyperladder I and EasyLadder I (Bioline, Australia) and PCR products were purified using illustra ExoSTAR 1-step enzymatic purification (GE 117 Ohlsen & Field, Diplazium squamuligerum Healthcare Life Sciences, UK). Purified PCR products were then sent to the Australian Genome Research Facility (AGRF), Melbourne Branch, where sequencing reactions, and capillary separation, using the 96-capillary analyser AB 3730x/ sequencing platform, were performed. Sequence editing, alignment, and analysis. Sequences were edited using Sequencher v. 3.0 (Gene Codes Corporation, Ann Arbor, MI, USA) and the NCBI nucleotide- nucleotide BLAST (blastn) tool (Altschul et al. 1997) was then used to evaluate the similarity of both rbcL and trnL-L sequences to existing sequences in GenBank. After the search using the BLAST tool was performed, available rbcL and trnL-L sequences of Diplazium species from GenBank were gathered (Appendix 1) and aligned manually in Se-Al Sequence Alignment Editor v. 2.0all (Rambaut 2002). Outgroup sequences were also gathered from GenBank (Appendix 1). Parsimony analyses were run in PAUP* v 4.0(310 (Swofford 2000). Gap characters in the alignment were treated as a fifth character state. For multiple base indels, characters were excluded from analyses so that indels were represented only by a single gap character, when variability did not occur within indels. Question marks, the character recognised in PAUP for missing data, were used to fill gaps, where needed, when indels fell across otherwise variable regions of the alignment. A heuristic tree search was used, with delayed character-state optimization (DELTRAN) and starting trees obtained by a closest addition sequence followed by tree bisection-reconnection (TBR) branch swapping. Bootstrap support for tree nodes was determined for each analysis, with 100 replicates. Separate parsimony analyses were performed for each chloroplast DNA region. Athyrium felix-femina (L.) Roth was chosen as an outgroup, based on the sister relationship of Athyrium to Diplazium in past molecular phylogenies (Sano et al. 2000; Wang et al. 2003; Schuettpelz & Pryer 2007) and for the rbcL analysis, in which it is possible to align distantly related taxa, the more distantly related Blechnum occidental L. and Asplenium marinum L. were also included as outgroups. Species identification. Online images of Diplazium types (B, MICH) were examined, to determine whether this taxon could be assigned to an existing Diplazium species, or if it was an undescribed species. Common Abbreviations used in Specimen Citations LA (Logging Area); NP (National Park). Results Searches using the BLAST tool of both trnL-L and rbcL sequences, determined that sequences from the P.E.I. Road specimen had the greatest similarity to species of Diplazium in the family Woodsiaceae. 37 Diplazium rbcL sequences were used to produce an alignment 1188 base pairs long, which contained 110 parsimonious characters. Two most parsimonious trees were obtained (length=398 steps, CI= 0.626) one of which is shown (Fig. 2a). This had a topology identical to the strict consensus tree, except for the placement of D. doderleinii (Luerss.) Mak., which in the consensus tree is placed in a polytomy with the clade of D. amamianum Tagawa + D. hachijoense Nakai and the clade of D. taiwanense Tagawa + D. virescens Kunze. 10 Diplazium trnL-L sequences were used to produce an alignment of 753 included base pairs, which contained 139 parsimonious characters. A single most parsimonious tree was obtained (length=302 steps, 0=0.831) (Fig. 2b). Maximum parsimony analyses of rbcL (Fig. 2a) and trnL-L (Fig. 2b) shows that the P.E.I. Road taxon is nested well within Diplazium sensu lato. Of all Diplazium species currently sequenced for rbcL, it is most closely related to D. proliferum (Lam.) Thouars. ( =Callipteris prolifera (Lam.) Bory sensu Jones [1998]) with high bootstrap support (98%). It differs from D. proliferum by 12 base pairs in rbcL. Inspection of Diplazium type material revealed that the P.E.I. Road taxon matches Asplenium varians var. squamuligerum Rosenst., from New Guinea, now treated as 118 15 70 73 6 1 93 99 2 Austrobaileya 9(1): 114—125(2013) I Diplazium amamianum ' Diplazium hachijoense — Diplazium doederleinii Diplazium taiwanense - Diplazium virescens var. conterminum - Diplazium virescens 2 |— Diplazium crassiusculum Diplazium donianum var. aphanoneuron 99 1 1 a 87 1 91 ^ Diplazium lobatum Diplazium dilatatum Diplazium hayatamae Diplazium deciduum Diplazium fauriei U- Diplazium mettenianum Diplazium griffithii — Diplazium pullingeri L-=— Diplazium kawakamii 8 76 - Diplazium centripetale —— Diplazium incomptum 4 9 |- Diplazium longicarpum - 12 97 1 --- Diplazium subserratum Diplazium proliferum 98 p 5 Diplazium chinense L_6 Diplazium squamuligerum 87 4 8 10 Diplazium subtripinnatum ■ Diplazium esculentum — Diplazium nipponicum ■ Diplazium sibiricum var. glabrum Diplazium squamigerum 9 Diplazium bombanasae 95 93 — Diplazium cristatum Diplazium lonchophyllum - Diplazium plantaginifolium 13 | Diplazium okudairae ' Diplazium wichurae 18 43 —— Diplazium pinfaense - Athyrium felix-femina - Blechnum occidentale - Asplenium marinum (a) Fig.2a. Parsimony analyses of rbcL showing the phylogenetic position of Australian Diplazium squamuligerum. Branch lengths are given above braches and bootstrap support values greater than 70% are given below branches. Ohlsen & Field, Diplazium squamuligerum 119 24 100 12 ■ Diplazium pinfaense r^- Diplazium wichurae 11 100 L^- Diplazium heterocarpum 7 99 Diplazium hainanense Diplazium hachijoense 7 89 16 15 Diplazium esculentum Diplazium nipponicum 46 15. 90 17 Diplazium mettenianum Diplazium squamuligerum 59 Diplazium ovatum Athyrium filix-femina (b) Fig.2b. Parsimony analyses of trnL-F. Branch lengths are given above braches and bootstrap support values greater than 70% are given below branches. 120 Diplazium squamuligerum (Rosenst.) Parris. The original diagnosis treated it as a variety of the unrelated African Asplenium varians Wall, ex Hook. & Grev. (Rosenstock 1913). The diagnosis is very brief, outlining only some immediately obvious features which distinguish D. squamuligerum from A. varians. Many of the morphological features that are quite distinctive are not mentioned. The following description describes Diplazium squamuligerum in greater detail, including several distinctive features overlooked by the earlier diagnosis. Taxonomy Diplazium squamuligerum (Rosenst.) Parris, Kew Bull. 41 (1): 69 (1986); Asplenium varians var. squamuligerum Rosenst., Repert. Spec. Nov. Regni Veg. 12: 528 (1913); A. squamuligerum (Rosenst.) Hieron, Hedwigia 61:5 (1919).Type: PapuaNew Guinea: Morobe Province: Sattelberg Hinterland, 1400-1500 m [I.C.]Keysser 228, April 1913 (holo: S, S-P- 7625, online image!; iso: UC 378428, online image!; MICH 1190092, online image!). Rhizome erect, to 1.5 cm tall, scaly; scales to 1 mm long, dark brown with toothed margins. Fronds arcuate, 5-15 cm long, 1.5-4 cm wide. Stipe 0.5-5 cm long, dark green, enveloped by green tissue which elongates to produce wings Austrobaileya 9(1): 114—125(2013) and protuberances; protuberances eventually bearing light brown scales to 1 mm long, with darker brown veins and bifid marginal teeth. Lamina bipinnate-tripinnatifid, 3-10 cm long, 1.5-4 cm wide, dark green above, paler below, occasionally proliferous from the rachis. Primary pinnae 5-25 mm long, 2-10 mm wide, most with an acroscopic secondary pinna to 5 mm long and an elliptic apical segment 5-10 mm long; margins serrate; veins free. Sori to 4 mm, elongate along most veins; indusium entire, narrow, light brown (Fig. la, b, d) Specimens examined: Australia: Queensland. Cook district: North Johnstone LA, July 1983, Lockyer s.n. (CANB); P.E.I. Road, Topaz NP, July 2005, Sankowsky & Sankowsky 2637 (BRI); North Johnstone LA, Jan 2013, Ohlsen 461 etal. (BRI, MELU); Portion 545 Parish of Dirran, Malian Road, July 1983, Gray 3133 (CNS). Distribution and habitat : In Australia Diplazium squamuligerum is known only from two areas on the Atherton Tableland, north-east Queensland; however, it also occurs in Papua New Guinea. It grows in mixed mesophyll rainforest, on metamorphic rocks or between tree roots in volcanic soil, in steep embankments of small, slow-flowing creeks (Fig. lc). Notes : Wings of the axes often become obscure when dried, but are obvious in fresh material. Key to Australian species of Diplazium (naturalised taxa indicated *) 1 Veins anastomosing.2 1 . Veins free. 5 2 Lamina simple.D. cordifolium 2. Lamina pinnate to tripinnate.3 3 Lamina pinnate, rachis proliferous.D. proliferum (syn. Callipteris prolifera) 3. Lamina bipinnate or tripinnate, rachis not proliferous. 4 4 Secondary pinnae 1-1.5 cm wide, margins incised more than half-way to the pinnule midvein.D. dietrichianum 4 . Secondary pinnae 1.5-2.5 cm wide, very shallowly lobed.*D. esculentum 5 Lamina pinnate.6 5. Lamina bipinnate or more divided. 7 6 Lamina apex a single pinna similar to lateral pinnae.D. pallidum 6. Lamina apex formed by reduction of lateral pinnae.D. dameriae 121 Ohlsen & Field, Diplazium squamuligerum 7 Fertile laminae < 250 mm long, stipe and rachises winged, wing extending to form scale bearing protuberances, pinnae with a basal acroscopic secondary pinna or lobe. D. squamuligerum 7. Fertile Laminae > 250 mm, stipe, rachis and pinnae not as above.8 8 Lamina bipinnate. 9 8. Lamina tripinnate or more divided. 10 9 Basal lobes of secondary pinnae longest, abaxial surface of pinnae glabrous. D. dietrichianum 9. Basal lobes of secondary pinnae similar to or smaller than the rest, abaxial surface bearing red glandular hairs. D. dilatatum 10 Lamina pale green, pinnules less than 5 mm long and 3 mm wide. D. assimile 10. Lamina dark green, pinnules greater than 10 mm long and 4 mm wide. 11 11 Apex of pinnules acute to caudate. D. bostockii 11. Apex of pinnules obtuse. 12 12 Pinnules of fertile lamina dissected < one third of the distance to the midvein, pinnules decurrent on basiscopic margin. D. australe 12. Pinnules of fertile lamina dissected > half-way to the midvein, pinnules not decurrent. D. queenslandicum Discussion The identification of the PE.I. Road taxon as a species of Diplazium by chloroplast DNA sequence is also supported by morphology. Populations of D. squamuligerum in Australia have elongate sori and scales that are borne on small protuberances: both features typical of Diplazium (Jones 1998). The scales are characterised by bifid teeth on a darkened margin and conform to the ‘Callipteris type’ (Fig. lb). This scale type is known only from the segregate genus Callipteris and a few other Diplazium species (Pachebo & Moran 1999; Pachebo & Moran 2003). These specimens were repeatedly misidentified as a species of Asplenium, rather than being correctly identified as a Diplazium. Hence this study is another case which highlights the value of DNA sequences for phylogenetic placement and identification when morphology may be misleading (Gastony et al. 2001). It also begs caution in identifying species by overall morphology without close inspection of more subtle but taxonomically informative features such as scale characters. Prior to this study, 12 species of Diplazium were known in Australia (one species, D. esculentum is naturalised). All of these Diplazium species are large terrestrials, with fronds over 50 cm long. The small size of D. squamuligerum is therefore unique amongst Australian Diplazium , and unusual amongst all Diplazium (Kato & Kramer 1990). Its position in the chloroplast phylogeny, as sister to D. proliferum is supported by its possession of the ‘Callipteris type’ scales, that are also possessed by D. proliferum (Jones 1998). Both D. squamuligerum and D. proliferum are also proliferous from the rachis, albeit rarely so in D. squamuligerum. Otherwise these species are quite dissimilar in general appearance. D. proliferum is a very large fern with pinnate fronds to over 2 metres in length whereas the fronds of D. squamuligerum are at least 2-pinnate and less than 15 cm long. D. squamuligerum also has free venation and winged axes (Fig. Id); the latter feature is not shared with any other Australian Diplazium species. The current phylogeny of Diplazium is limited in the number of species sampled. Chloroplast DNA sequences of only 38 species are publicly available, all of which 122 are incorporated in the phylogenies presented here. This is a low proportion of the 400 species of Diplazium in total (Kato & Kramer 1990). So it is highly likely that another unsampled species of Diplazium is more closely related to D. squamuligerum than D. proliferum, especially given the large number of base pair differences between the two species. Several Diplazium species, which occur in New Guinea, or possess the ‘Callipteris type’ scales, are yet to be sequenced, and may be more closely related. Diplazium has received little attention from molecular study, as is demonstrated by the low number of sequences available on GenBank. Further molecular work, involving a more comprehensive sampling of species, is required in Diplazium not only to gain a better understanding of the systematics and biogeography of this genus, but also to aid taxonomic revision, particularly at the generic level. It has been suggested, based on a recent molecular study using a rather small number of species (Wang et al. 2003), that several genera which had been segregated from Diplazium be placed back into that genus. This included Callipteris , a genus still accepted in Australia (Jones 1998). However, revision of Diplazium cannot be completed until more species sampling is undertaken. The close relationship of D. squamuligerum to D. proliferum suggests that the ‘Callipteris type’ scale may define a monophyletic group. It has been suggested that this feature may be used to recircumscribe Callipteris , to include the few Diplazium species, such as D. squamuligerum , that have this scale type, and also have free venation (Pachebo & Moran 2003). However, the monophyly of all species possessing this scale type needs to be demonstrated before any revision is undertaken (Pachebo & Moran 2003). Identification of this species as D. squamuligerum provides an additional example of a fern species that is localised in a few sites on the Atherton Tableland. Populations at Topaz NP are within close proximity to other localised fern species. They are within 3 km of the only known locality of Hymenasplenium unilateral in Australia, and Austrobaileya 9(1): 114—125(2013) adjacent to one of approximately ten locations of Dryopteris wattsii M.McKeown, Sundue & Barrington. In addition to these exceptionally rare species, other uncommon fern species such as Dicksonia herbertii W.Hill, Oleandra musifolia , and Pteridoblechnum acuminatum (C.T.White & Goy) Hennipman and P. neglectum (F.M.Bailey) Hennipman also occur at or near the D. squamuligerum population localities. Likewise, the Maalan population is within 7 km of one of only three populations of Asplenium normale D.Don in Australia. The occurrences of multiple, very localised fern species near sites of D. squamuligerum highlights the high conservation priority that should be given to these areas, especially considering the high diversity of seed plant species also present. It also suggests that a number of equally localised species may have been lost due to vegetation clearing around these populations. Diplazium squamuligerum is currently one of the most threatened fern species in Australia. While it is reserved in Topaz NP, all populations of this species are small in area and size and are particularly vulnerable to stochastic events. Periods of particularly high rainfall, with increased runoff due to surrounding cleared land, can swell the small creeks along which it occurs. Consequently plants face the danger of being uprooted from the creek embankments. This species would also be extremely susceptible to desiccation. It occurs in very high rainfall areas, and along small slow flowing creeks, that provide the constant soil moisture and high humidity that is probably required by the plant. Extended dry periods may reduce such small creeks and limit water access to plants. Such stochastic weather events are expected to become more extreme in the future with ongoing climate change (Abbs et al. 2006; Mpelasoka et al. 2008; Walsh et al. 2004). The one collection from Papua New Guinea that exists in Australian herbaria (Brass 25731 [CANB]) was collected from Mt Pabinama, Normanby Island, which is 550 km south-east from the type locality, and suggests that D. squamuligerum is more widespread in Papua New Guinea. 123 Ohlsen & Field, Diplazium squamuligerum A conservation status of Vulnerable is recommended for D. squamuligerum in accordance with the distribution and abundance characteristics of other taxa listed as Vulnerable under the Nature Conservation Act 1992 (Queensland), the Environment Protection and Biodiversity Conservation Act 1999 (Australian Commonwealth) and also in line with the criteria of the International Union for Conservation of Nature (IUCN). A conservation advice report is currently being prepared for D. squamuligerum in Australia. Acknowledgements We would like to express our gratitude to Bruce Gray, who assisted the recent collection of Diplazium squamuligerum. All sequencing was performed in the Cookson Laboratory, School of Botany, The University of Melbourne, and was funded by a BushBlitz Research Grant. Much appreciated advice in sequencing D. squamuligerum was provided by Lara Shepherd and Leon Perrie. We are also thankful to Leon Perrie, Nada Sankowsky, Darren Crayn, and to the anonymous reviewers for their helpful comments on the manuscript. References Abbs, D., Aryal, S., Campbell, E., McGregor, J., Nguyen, K., Palmer, M., Rafter, T., Watterson, I. & Bates, B. (2006). Projections of extreme rainfall and cyclones. CSIRO, Final report to the Australian Greenhouse Office. Andrews, S.B. (1990). Ferns of Queensland. A handbook to the ferns and fern allies , 402-404. Queensland Department of Primary Industries: Brisbane, Australia. Altschul, S.F., Madden, T.L., Schaffer, A.A., Zhang, J., Zhang, Z., Miller, W. & Lipman, D. J. (1997). Gapped blast and psi-blast: a new generation of protein database search programs. Nucleic Acids Research 25: 3389-3402. Bell, G.H. (1998). Davalliaceae. In PM. McCarthy (ed.), Flora of Australia Ferns, Gymnosperms and Allied Groups 48: 434-450. ABRS/CSIRO: Melbourne. Brownsey, P.J. (1998). Aspleniaceae. In PM. McCarthy (ed.). Flora of Australia Ferns, Gymnosperms and Allied Groups 48: 295-327. ABRS/CSIRO: Melbourne. Gastony, G.J. & Johnson, W.P (2001). Phylogenetic placements of Loxoscaphe thecifera (Aspleniaceae) and Actiniopteris radiata (Pteridaceae) based on analysis of rbcL nucleotide sequences. American Fern Journal 91: 197-213. Hovenkamp, P.H. & Ho, B-C. (2012). A revision of the fern genus Oleandra (Oleandraceae) in Asia. PhytoKeys 11: 1-37. Jones, D.L. (1998). Athyriaceae. In P.M. McCarthy (ed.). Flora of Australia Ferns, Gymnosperms and Allied Groups 48: 418-429. ABRS/CSIRO: Melbourne. Kato, M. & Kramer, K.U. (1990). Subfamily Athyrioideae. In K.U. Kramer & PS. Green (eds.). The families and genera of vascular plants Pteridophytes and gymnosperms 1: 130- 144. Springer-Verlag: Berlin. Kramer, K.U. & Viane, R. (1990). Aspleniaceae. In K.U. Kramer & PS. Green (eds.). The families and genera of vascular plants Pteridophytes and gymnosperms 1: 52-56. Springer-Verlag: Berlin. Lovis, J.D. (1977). Evolutionary patterns and processes in ferns. Advances in Botanical Research 4: 223-415. McCarthy, PM. (1998). Flora of Australia Ferns, Gymnosperms and Allied Groups 48. ABRS/ CSIRO: Melbourne. Mpelasoka, F., Hennessy, K., Jones, R. & Bates, B. (2008). Comparison of suitable drought indices for climate change impacts assessment over Australia towards resource management. International Journal of Climatology 28: 1283— 1292. Pachebo, L. & Moran, R.C. (1999). Monograph of the neotropical species of Callipteris with anastomosing veins (Woodsiaceae). Brittonia 51: 343-388. - (2003). Lectotypification of several names currently placed in Diplazium (Woodsiaceae). American Fern Journal 93: 90-92. Parris, B.S. (1986). New combinations in filices. Kew Bulletin 41: 69-70. Rambaut, A. (2002). ‘Se-Al: sequence alignment editor’. Available at: http://tree.bio.ed.ac.uk/ Rosenstock, E. (1913). Filices novoguineenses Keysseranae, III. Repertorium Specierum Nov arum Regni Vegetabilis 12: 524-530. Sano, R., Takamiya, M., Ito, M., Kurita, S. & Hasebe, M. (2000). Phylogeny of the lady fern group, tribe Physematieae (Dryopteridaceae), based on chloroplast gene sequences. Molecular Phylogenetics & Evolution 15: 403-413. 124 Schuettpelz, E. & Pryer, K.M. (2007). Fern phylogeny inferred from 400 leptosporangiate species and three plastid genes. Taxon 56: 1037-1050. Smith, A.R., Pryer, K.M., Schuettpelz, E., Korall, P, Schneider, H. & Wolf, PG. (2006). A classification for extant ferns. Taxon 55: 705- 731. Swofford, D.L. (2000). PAUP* 4.0: phylogenetic analysis using parsimony (*and other methods). Sinauer Associates: Sunderland, MA. Taberlet, P, Gielly, L., Pautou, G. & Bouvet, J. (1991). Universal primers for amplification of three non-coding regions of chloroplast DNA. Plant Molecular Biology 17: 1105-1109. Austrobaileya 9(1): 114—125(2013) Trewick, S.A., Morgan-Richards, M., Russell, S.J., Henderson, S., Rumsey, F.J., Pinter, I., Barrett, J.A., Gibby, M. & Vogel, J.C. (2002). Polyploidy, phylogeography and Pleistocene refugia of the rockfern Asplenium ceterach: evidence from chloroplast DNA. Molecular Ecology 11: 2003-2012. Walsh, K. J.E., Nguyen, K.C. & McGregor, J.L. (2004). Finer-resolution regional climate model simulations of the impact of climate change on tropical cyclones near Australia. Climate Dynamics 22: 47-56. Wang, M-L., Chen, Z-D., Zhang, X-C., Lu, S-G. & Zhao, G-F. (2003). Phylogeny of the Athyriaceae: evidence from chloroplast trnL-F region sequences. Acta Phytotaxonomica Sinica 41: 416-426. Appendix 1. Chloroplast DNA sequences included in analyses, identified by GenBank accession numbers Taxon rbcL trnL Asplenium marinum L. AF240647 - Athyrium felix-femina (L.) Roth FJ821348 HQ676519 Blechnum occidentals L. U05909 - Diplazium amamianum Tagawa AB574961 - D. bombanasae Rosenst. EF463308 - D. centripetale (Bak.) Maxon EF463309 - D. chinense (Bak.) C.Chr. AB574962 - D. crassiusculum Ching AB574963 - D. cristatum (Desr.) Alston EF463310 - D. deciduum N.Ohta & M.Takamiya AB574964 - D. dilatatum Blume EF463311 - D. doederleinii (Luerss.) Mak. AB574966 - D. donianum (Mett.) Tard var. aphanoneuron (Ohwi) Tagawa AB574968 - D. esculentum (Retz.) Sw. U05619 AF514838 D. fauriei Christ AB574970 - D. griffithii (T.Moore) Diels AB574971 - D. hachijoense Nakai EF463312 EU329111 D. hainanense Ching - AF515257 Ohlsen & Field, Diplazium squamuligerum 125 Taxon rbcL trnL D. hayatamae N.Ohta & M.Takamiya AB574973 - D. heterocarpum Ching - - D. incomptum Tagawa AB574974 - D. kawakamii Hayata AB574975 - D. lobatum (Tagawa) Tagawa AB574976 - D. lonchophyUum Kunze U05920 - D. longicarpum Kodama AB574977 - D. mettenianum (Miq.) C.Chr. AB574978 AF515233 D. nipponicum Tagawa AB574979 EU329110 D. okudairae Mak. AB574980 - D. ovatum Wall. - AF515232 D. pinfaense Ching AB574981 AF515250 D. plantaginifolium (L.) Urb. EF463314 - D. proliferum (Lam.) Thouars EF463918 - D. pullingeri (Bak.) J.Sm. AB574982 - D. sibiricum (Turcz ex Kunze) Kurata in Namegata var. glabrum (Tagawa) Sa.Kurata AB574983 - D. squamigerum (Mett.) Christ AB574984 - D. squamuligerum (Rosenst.) Parris KC900228 KC900229 D. subserratum (Blume) T.Moore AB021721 - D. subtripinnatum Nakai AB574985 - D. taiwanense Tagawa AB574986 - D. virescens Kunze var. conterminum (Christ.) Sa.Kurata JN168017 - D. virescens Kunze EF463316 - D. wichurae (Mett.) Diels JN168018 AF515245 Austrobaileya 9(1): 126-129 (2013) 126 SHORT COMMUNICATION New records for Queensland in Lindernia All. (Linderniaceae) B.S. Waiman Australian Tropical Herbarium, James Cook University, Cairns Campus, McGregor Road Smithfield, Queensland 4878, Australia. Email: bswannan@bigpond.com The genus Lindernia All. (Linderniaceae Borsch, K.Mull. & Eb.Fisch.) is a pan- tropical genus represented in Australia by approximately 50 species. In Australia, many of the species are yet to be formally named, especially those from the Northern Territory. During the current taxonomic revision of Lindernia in Queensland (Wannan in prep.) two new species have been recorded, namely: Lindernia pusilla (Willd.) Bold., which is a new record for Australia and L. tectanthera W.R.Barker, which is a new record for Queensland. Lindernia pusilla (Willd.) Bold., Zakfl. Java 165 (1916); Gratiolapusilla Willd., Sp. Pl. ed. 4 , 1(1): 105 (1797); Pyxidaria pusilla (Willd.) Kuntze, Rev. Gen. PI. 2: 468 (1891); Vandellia pusilla (Willd.) Merr., Philipp. J. Sci. Bot. 7: 246 (1912). Type: India, s.dat., s.coll. (holo: B-W 377). Weakly erect herb to 10 cm. Stems decumbent, densely pubescent with filamentous trichomes to 1 mm. Leaves simple opposite, very shortly petiolate or subsessile, petioles up to 1 mm. long, hirsute; lamina 5-15(20) mm long, 3-15 mm. wide, broadly ovate to subcircular to cordate, crenate-dentate, filamentous trichomes on both sides but more densely below; 5-nerved from base of leaf Flowers axillary, in clusters of 1-3 per axil; pedicels slender, to 20 mm in flowers, 30 mm in fruit, with stalked glandular trichomes. Calyx deeply 5-lobed, almost to the base, lobes 3-4 mm long, single nerved and with stalked glandular and filamentous trichomes; margin finely serrate, scarious. Corolla 5-8.5 Accepted for publication 29 July 2013 mm. long, white to pale violet to purple; tube externally glabrous or with sparse stalked glandular trichomes, internally with stalked glandular trichomes near antero-lateral ridges, otherwise glabrous; upper lip narrow emarginate; lower lip 3-lobed. Stamens 4, fertile, cohering in pairs; upper filaments 1-1.5 mm. long, glabrous; lower filaments 3-4 mm long, broadly geniculate just above the base, forming a ‘webbed’ bend. Ovary c. 1 mm; style 5-7 mm. Capsule subglobose to ovoid, more or less equal in length to the calyx. Seeds light brown, cylindric, 0.5 x 0.3 mm, with up to 8 longitudinal ribs. Fig. 1. Selected specimens examined : Malaysia. Lawn at Forestry Department Kuching Sarawak, Nov 1988, Wannan UNSW16578 (CANB, NSW, UNSW). Australia: Northern Territory. 2 km NE of Port Bradshaw, Gove Peninsula, Sep 1987, Russell-Smith 3380 & Lucas , (AD, DNA). Queensland. Cook District: North of Coen, Jul 1999, Wannan 1379 (BRI, CANB); North of Coen, Sep 2007, Wannan 4855 & Graham (BRI); ditto loc., Jun 2009, Wannan 5696 & Thompson (CNS). Notes : Lindernia pusilla was first recognised in Australia from a collection made in 1999 just north of Coen beside the Peninsula Developmental Road, Queensland ( Wannan BSW1379). Despite searches, no further localities have been recorded on Cape York Peninsula. During subsequent examination of Lindernia collections at AD a further collection was identified from the Northern Territory ( Russell-Smith 3380 & Lucas). These sites remain the only known Australian localities. According to Philcox (1968), Lindernia pusilla is widespread in the Indian subcontinent, China, Myanmar, Thailand and Malesia (Borneo, Celebes, Java, Malay Peninsula, Moluccas, Philippines, Sumatra and mainland New Guinea) up to an altitude of 1600 metres. It has also subsequently been Wannan, Lindemia 127 Fig. 1 . Lindemia pusilla (Wannan 1379\ BRI). Photo: B. Wannan. recorded from the Bismarck Archipelago of Papua New Guinea (Peekel 1984). The Australian material is a good match with the type and Malesian material (cf Wannan UNSW16578 ) based on its morphology and, for the latter, seed anatomy. Although a key to Queensland species will be provided in the revision (Wannan in prep.), L. pusilla can be distinguished from other Australian Lindemia by its multiple (pink) flowers per axil, long (to 30 mm) pedicels, and its stems pubescent with filamentous trichomes. Lindemia tectanthera W.R. Barker, J. Adelaide Bot. Gard. 13: 86-87 (1990). Type: Western Australia. Kimberley Region: 23.3 km along King River Road, 25 April 1983, R.M. Barker 284 (holo: AD). Erect, sometimes decumbent, glabrous annual herb, 8-40 cm tall, with slender scapiform stems arising from a basal cluster of leaves; sometimes with stem branched in upper parts below the terminal inflorescence. Leaves at the base spathulate, 6-22 mm long, with the petiole 1-5 mm long, the blade ovate to obovate, sometimes narrowly so, 3-15 x 1.5-8 mm, entire, with midvein. Cauline leaves/ bracts subulate, 1-4 mm x 1-2 mm. Inflorescence terminal racemes or panicles; pedicels 5-32 mm at anthesis and erect, 15-40 mm in fruit and sometimes deflexed. Calyx 2-4.5 mm long; sepals 5, equal, divided almost to base, triangular, 3-nerved, often reddened. Corolla 2-lipped, with internal longitudinal flaps enclosing stamens and style; stalked glandular trichomes externally; white, with purple or mauve; tube 6-7 mm; upper lip recurved, 1.5-2 mm long, shallowly emarginate; lower lip widely spreading, 4-6.5 mm long, the lobes shallowly emarginate, the mid lobe wider than the laterals. Stamens with upper 2 stamens functional, filaments c. 1 mm; anthers 0.5-0.8 mm long, yellow, 128 Austrobaileya 9(1): 126-129 (2013) Fig. 2. Lindernia tectanthera ( Wannan 6291 & Trenerry. BRI). Photo: B. Wannan. with 2 locules end to end, approximately 180° divergent; lower stamens sterile, anthers represented by 2 appendages geniculate at summit, proximally thick, near the apex often bearing a minute anther vestige. Ovary 1-1.5 mm, glabrous, style c. 5 mm, glabrous. Capsule ovoid, 4-5 x 2.5 mm, thin-walled, yellow-brown; seeds broad obovoid to oblong-ellipsoid, 0.3-0.35 x 0.2-0.25 mm, light brown, with 4 longitudinal angles and 4-6 transverse ridges. Fig. 2. Selected specimens examined : Western Australia. 3 km S of Pitta Creek near Prince Regent River, Jul 2009, Wannan 5740 (BRI, CNS, NSW, PERTH); Drysdale River NP, Jul 2012, Wannan 6563 & Wardrop (CNS, PERTH). Queensland. Cook District: Sand Ridge, Staaten River NP, Jan 2003, Fox IDF2002 (BRI); Wrotham Park, Jul 2011, Wannan 6114 (BRI); Gamboola, Aug 2011, Wannan 6291 & Trenerry (NSW), Wannan 6298 & Trenerry (BRI). Notes: Lindernia tectanthera was first described from the Kimberley region of Western Australia (Barker 1990). This species has been recently confirmed from west of Chillagoe in Queensland and is a good match with material from the Kimberley including the holotype, based on morphology and seed anatomy. Examination of Fox IDF2002 (Staaten River NP) has shown it to be also L. tectanthera. Queensland specimens of this taxon have been previously incorrectly determined as Lindernia lobelioides (F.Muell.) F.Muell., a Northern Territory species. Lindernia tectanthera has been recorded widely from the Top End and Kimberley regions. The species typically occurs in moist areas often on the margin of sandy rises. It is distinguished from other Queensland species by its usual basal rosette of leaves and its upper corolla with a longitudinal flap which encloses the anthers and style. Acknowledgements Thanks to Darren Crayn (CNS) for his support of the project and the directors of AD, BRI, CANB, DNA, MEL, NSW and UNSW for loans and/or access to collections. This work has been supported by Australian Biological Resources Study (ABRS) National Taxonomy Wannan, Lindemia 129 Research Grant Program CN211-26 which has been undertaken at the Australian Tropical Herbarium. References Barker, W.R. (1990). Newtaxa, names and combinations in Lindemia , Peplidium , Stemodia and Striga (Scrophularlaceae) mainly of the Kimberley Region, Western Australia. Journal of the Adelaide Botanic Gardens 13: 79-93. Peekel, P.G. (1884). Flora of Bismarck Archipelago for Naturalists. Translated by E.E. Henty. Office of Forests, Division of Botany: Lae, Papua New Guinea. Philcox, D. (1968). Revision of the Malesian species of Lindemia All. (Scrophulariaceae). Kew Bulletin 22: 1-72. Status and lectotypification of Craspedia gracilis Hook.f. (Asteraceae: Gnaphalieae) Andrew C. Rozefelds Summary Rozefelds, A.C. (2013). Status and lectotypification of Craspedia gracilis Hook.f. (Asteraceae: Gnaphalieae). Austrobaileya 9(1): 130-135. The identity of Craspedia gracilis Hook.f. is determined and it is shown to be conspecific with C. coolaminica J.Everett & Joy Thomps. which, being the younger name, is therefore placed in synonymy. The identity of Craspedia richea var. linearis Hook.f. is also resolved and it is shown to be a synonym of C. gracilis. Lectotypes of C. gracilis and C. richea var. linearis are designated and illustrated. Key Words: Asteraceae, Gnaphalieae, Craspedia, Craspedia coolaminica, Craspedia gracilis, Craspedia richea var. linearis, Australia flora, Tasmania flora, taxonomy, billy buttons A.C.Rozefelds, Queensland Museum, GPO BOX 3300, South Brisbane, Queensland 4101, Australia. Email: andrew. rozefelds@qm. qld.gov. au Introduction Previous studies of Craspedia G.Forst. by Rozefelds (2002) and Rozefelds et al. (2011), have resolved the nomenclature of species of Craspedia described from Tasmania; however, the status of two taxa described by Hooker (1847), C. gracilis Hook.f and C. richea Cass. var. linearis Hook.f, has been unclear. Cassini (1818) recognised that Richea glauca , described by Labillardiere (1800) from Tasmania, should be transferred to Craspedia. His new combination, Craspedia richea Cass.; however, is a nomen illegitimum, as recognized by Chapman (1991), since it was based on Richea glauca Labill. whose epithet should have been adopted. Sprengel (1826) successfully transferred this species to Craspedia , i.e. C. glauca (Labill.) Sprengel. Hooker (1847) described Craspedia gracilis Hook.f from the Middlesex Plains [Tasmania], although he later, in ‘ Flora Tasmaniae’, reduced C. gracilis to a variety, C. richea var. gracilis Hook.f Curtis (1963: 346) recognised that ‘C. glauca included several further spp. and varieties [that] have been named’. She recognised four varieties including the new combination C. glauca var. gracilis , but as the original reference and dates of publication for this taxon were not cited, it is invalid (Chapman 1991). Hooker (1847) also recognised two new varieties of Craspedia richea. The status of C. richea var. glabrata Hook.f from the western mountains of Tasmania, was examined by Rozefelds (2002) and a new combination C. glabrata (Hook.f.) Rozefelds was proposed. The status of C. richea var. linearis Hook.f from the western mountains of Tasmania is reviewed herein. Curtis (1963) did not discuss the status of this variety. Craspedia coolaminica J.Everett & Joy Thomps. was described from material collected on mainland Australia and Tasmania, but the description of this species overlaps with that of both C. richea var. linearis and C. gracilis. To resolve the status of these taxa described by Hooker (1847), cibachromes of early Tasmanian collections of Craspedia gracilis and C. richea var. linearis, held at Kew, were obtained; and from the study of these images it is proposed that C. coolaminica and C. richea var. linearis are placed in synonymy under C. gracilis. Accepted for publication 25 September 2013 Rozefelds, Craspedia gracilis Taxonomy Craspedia gracilis Hook.f. Craspedia gracilis Hook.f., Lond. J. Bot. 6: 118 (1847); C. richea Cass. [nom. illeg .] var. gracilis (Hook.f.) Hook.f., Flora Tasmaniae 3: 197 (1857); C. glaiica var. gracilis (Hook.f.) W.M.Curtis comb, invalid., The Student’s Flora of Tasmania 2: 247 (1963). Type citation: “Middlesex plains; Gunn”. Type: Tasmania. Middlesex PL, 15 February 1837, [R.C.]Gunn 835/1837 (lecto [here designated]: K n.v., cibachrome at HO!). The specimen associated with the label on the right hand side of the plate is the lectotype (Fig. 1). Craspedia richea Hook.f. [nom. illeg .] var. linearis Hook.f., Lond. J. Bot. 6: 118 (1847); C. richea Cass. [nom. illeg] var. |3 linearis Hook.f., Flora Tasmaniae 3: 197 (1857). synon. nov. Type citation: “Western Mountains; Gunn”. Type: Tasmania. Western Mountains, 17 February either 1842 or 1843 (as the year is not consistent on the label), [R.C.] Gunn 1160/1842 (lecto [here designated]: K n.v., cibachrome at HO!). The more complete specimen on the left hand side of the plate is the lectotype (Fig. 2). Craspedia coolaminica J.Everett & Joy Thomps., synon. nov., Telopea 5: 45 (1992). Type: New South Wales. Southern Tablelands: Happy Jacks, Snowy Mountains, near Happy Jacks Road, above sheep yard, 27 January 1964, M.E. Phillips 20419 (holo: NSW n.v/, iso: CBG n.v) Typification : In describing Craspedia gracilis. Hooker (1847) did not specify a numbered collection of Gunn’s. The specimen Gunn 835/1837 from Middlesex Plains comprises three individual plants and was examined by Hooker; hence, it is an appropriate choice for lectotypification of the name. A lectotype is chosen based on the right-hand specimen closest to the collector’s label. Craspedia richea var. linearis was described by Hooker (1847) as having narrowly linear leaves with arachnose hairs and known from the “Western Mountains”, and although no specific locality, nor collector number were recorded a specimen in K, Gunn 1160/1842, 131 dated 18 February 1843, is consistent with the limited published information. Notes on this herbarium sheet, in Ronald Campbell Gunn’s handwriting, record it as Craspedia, but the note has been annotated by a later author with ‘richea var. linearis’. The narrowly linear leaves and silvery grey foliage are consistent with C. gracilis. Gunn’s notes on the type of C. richea var. linearis records the florets as orange (Fig. 2), and while no orange-floreted species are known from Tasmania, the bright yellow flowers of C. gracilis approach an orange colour (Fig. 3). The combination, C. richea var. linearis, was only used by Hooker, so it seems very likely that this collection was examined by Hooker, and as it comes from the “Western Mountains” and was collected by Gunn, it is an appropriate choice for lectotypification of this name. As the herbarium sheet includes two similar¬ sized specimens and the right-hand specimen is more complete, it is designated as the lectotype. Hooker (1857) in Flora Tasmaniae, considered C. richea var. |3 linearis as being restricted to the Western Mountains and referred a second collection ( Gunn 1160 ) from Arthurs Lake to this variety. This collection {Gunn 1160 ) comprises four individual plants; all have long narrow silvery grey leaves with finely appressed hairs, and with arachnose hairs on the margin and are consistent with C. gracilis as defined herein. Mainland Australian and Tasmanian populations of Craspedia gracilis Craspedia gracilis as defined here occurs in subalpine communities in New South Wales, Victoria and Tasmania (Everett & Thompson 1992; Schmidt-Lebuhn & Milner 2013; Rozefelds pers. obs). Like many species of Craspedia, plants are often morphologically variable, with the size of the plants and the various organs (leaves, scapes, bracts on scape and compound head) phenotypically plastic. Material from Tasmania is variable in size from small herbs (like the type), to large, robust specimens 60 cm high (Rozefelds pers. obs). The mainland material show a similar variation in size (Everett & Thompson 1992) and no morphological differences were noted Fig. 1 . Lectotype of Craspedia gracilis Hook.f. (arrowed), Gunn 835/1837 (K). The lectotype shows the typical narrow linear silvery grey leaves, fine hairs and secondary veins are clearly evident on the underside of the leaves. Reproduced by kind permission of the Board of Trustees of the Royal Botanic Gardens, Kew. Rozefelds, Craspedia gracilis 133 Fig. 2. Lectotype of Craspedia richea Cass. var. p linearis Hook.f. (arrowed), Gunn 1160 (K). Reproduced by kind permission of the Board of Trustees of the Royal Botanic Gardens, Kew. 134 Austrobaileya 9(1): 130-135 (2013) Fig. 3. Floret colour in Craspedia gracilis Hook.f (left hand inflorescence) compared with C. rosulata Rozefelds & A.M.Buchanan (right hand inflorescence). between these mainland Australian and Tasmania populations by these authors in their original description. Sequence data for mainland Australian and Tasmanian samples of C. gracilis (i.e. C. coolaminica in Ford et al. 2007) showed no variation in ITS, seven base differences in ETS, and in psbA-trnH there were two base differences and also two deletions in the Tasmanian specimen (Kerry Ford, pers. comm. 2013). There seems little justification based upon the current molecular data, and the overall similarities in morphology to support separation of mainland Australian and Tasmanian populations. As the two samples of C. gracilis were sister to each other in Ford et al. (2007) analysis, it also becomes an arbitrary decision, in a phylogenetic sense, as to whether they are considered conspecific or sister species. Discussion The lectotype of Craspedia gracilis Hook.f {Gunn 835/1837 ) has long linear, narrowly lanceolate leaves, fine appressed hairs that impart the silver grey foliage, secondary veins clearly visible on the underside of the leaves due in part to the appressed hairs on the veins, and arachnose hairs along the leaf margins. The distinguishing features of Craspedia coolaminica J.Everett & Joy Thomps. (Everett & Thompson 1992; Rozefelds et al. 2011) easily distinguish this species from others in Tasmania, viz. distinctive basal rosette of long linear to narrowly attenuate silvery grey leaves that have a dense covering of long, fine appressed hairs, conspicuous secondary veins on the abaxial leaf surface, yellow corolla and mid-summer flowering (December- February). As no characters have been recognised that separate these taxa apart, I conclude that C. coolaminica and C. richea var. linearis are both synonyms of C. gracilis. Dawson et al. (1999) attributed a chromosome count of 2n=44 for Craspedia gracilis (as C. glauca ; Kerry Ford pers. comm. 2010). Putative hybrids have been noted between C. gracilis and the typically earlier flowering C. rosulata Rozefelds & A.M.Buchanan which is sympatric with this species at Miena, Central Plateau and the two species overlap slightly in flowering time. Putative hybrids are robust plants and have bright yellow florets like C. gracilis , rather than the light yellow of C. rosulata , but lack the conspicuous silver grey indumentum of C. gracilis. Rozefelds, Craspedia gracilis Acknowledgements I thank Alex Chapman in obtaining cibachromes of the relevant collections, Kerry Ford for advice on molecular data, and Alex Buchanan for his help and interest in the project. Fig. 1. and Fig. 2 are reproduced by kind permission of the Board of Trustees of the Royal Botanic Gardens, Kew. References Cassini, A.H.G. de (1818). Craspedie, Craspedia. In Dictionnaire des sciences naturelles, dans lequel on traite methodiquement des differens etres de la nature. Second Edition. F.G. Levrault: Strasbourg/ Le Normant: Paris. Chapman, A.D. (1991). Australian Plant Name Index A-C. Australian Biological Resources Study: Canberra. Curtis, W.M. (1963). The Student’s Flora of Tasmania, Part 2. Government Printer: Hobart. Dawson, M.I., Breitwieser, I. & Ward, J.M. (1999). Chromosome numbers in Craspedia , Ewartia and Pterygopappus (Compositae: Gnaphalieae). Australian Systematic Botany 12: 671-674. Everett, J. & Thompson, J. (1992). New alpine and subalpine species in Craspedia sens, strict. (Asteraceae: Gnaphalieae). Telopea 5: 45-51. Ford, K.A., Ward, J.M., Smissen, R.D., Wagstaff, S.J., Breitwieser, 1. (2007) Phylogeny and biogeography of Craspedia (Asteraceae: Gnaphalieae) based on ITS, ETS and psbA-trnH sequence data. Taxon 56: 783-794. 135 Hooker, J.D. (1847). Florae Tasmaniae Spicilegium or Contributions towards a Flora of Van Diemens Fand. London Journal of Botany 6: 106-125. Hooker, J.D. (1857). Botany of the Antarctic Voyage of H.M. Discovery ships Erebus and Terror in the years 1839-1843. Vol. 3. Flora Tasmaniae. Fovell Reeve: Fondon. Fabillardiere, J.J.H. de (1800). Relation du voyage a la recherche de La Perouse, fait par ordre de L’assemblee Constituante, pendant les annies 1791, 1792, et pendants l ere la 2 de annee de la Republique Francoise. 2 Vols. H.J. Jansen: Paris. Rozefelds, A.C. (2002). A new species and new combination in Craspedia (Asteraceae) from Tasmania. Telopea 9: 813-820. Rozefelds, A.C., Buchanan, A.M. & Ford, K.A. (2011). New species of Craspedia (Asteraceae: Gnaphalieae) from Tasmania, and determination of the identity of C. macrocephala Hook. Kanunnah 4: 93-116. Schmidt-Febuhn, A.N. & Milner, K.V. (2013). A quantitative study of morphology in Australian Craspedia (Asteraceae: Gnaphalieae). Australian Systematic Botany 26: 238-254. Sprengel, C.P.J. (1826). Systema Vegetabilium 3. Gottingae sumtibus librariae: Dieterichlanae. Craspedia uniflora G.Forst. (Asteraceae) is a New Zealand endemic and not part of the Australian flora Alexander N. Schmidt-Lebuhn Summary Schmidt-Lebuhn, A.N. (2013). Craspedia uniflora G.Forst. (Asteraceae) is a New Zealand endemic and not part of the Australian flora. Austrobaileya 9(1): 136-139. Specimens identified as Craspedia uniflora from Australia in the Queensland Herbarium are shown to have been misidentified and this species should therefore be deleted from both the Census of the Queensland Flora and the Australian Plant Census. The absence of any Australian material of this species in Australian herbaria also indicates that C. uniflora is a New Zealand endemic. Four species of Craspedia are identified as occurring in Queensland, C. canens J.Everett & Doust, C. haplorrhiza J.Everett & Doust, C. paludicola J.Everett & Doust (syn. C. sp. (GirraweenNP S.T.Blake 23643)) and C. variabilis J.Everett & Doust, and a key to these species is provided. Key Words: Asteraceae, Compositae, Gnaphalieae, Australian Plant Census, Census of the Queensland Flora, Craspedia , Craspedia canens , Craspedia haplorrhiza , Craspedia paludicola , Craspedia uniflora , Craspedia variabilis , identification key A N. Schmidt-Lebuhn, CSIRO Plant Industry, Centre for Australian National Biodiversity Research, GPO Box 1600, Canberra, ACT 2601, Australia. Email: alexander.schmidt-lebuhn@csiro.au Introduction Craspedia G.Forst. (Asteraceae, Gnaphalieae) is a genus of c. 20 Australian and at least 6 New Zealand species of mostly perennial rosette plants bearing yellow, white or orange flowers in solitary compound heads on leafy scapes. Although found in all states of Australia except the Northern Territory, Craspedia is most diverse in subalpine to alpine regions of the south-eastern mainland and Tasmania. Craspedia was described by Forster (1786) with one species, C. uniflora G.Forst. from New Zealand. Over the following two hundred years, several other species were described from New Zealand and Tasmania but only one from the Australian mainland (see Schmidt-Lebuhn & Milner 2013 for an overview), so that mainland specimens were often haphazardly referred to species from New Zealand and Tasmania, especially C. uniflora and C. glauca (Labill.) Spreng. Curtis (1963) did not consider C. uniflora to occur in Tasmania, while Stanley & Ross (1986) used the name for all specimens of Craspedia Accepted for publication 24 July 2013 s.str. in south-eastern Queensland (the other two species they treated under Craspedia are now assigned to Pycnosorus Benth. which is characterized by sessile partial heads, yellow phyllaries and pappus, and a different base chromosome number; Everett & Doust 1992a). Recent taxonomic or floristic treatments of Craspedia in Tasmania and the Australian mainland did not mention the species (Everett & Doust 1992b, c; Everett & Thompson 1992; Everett 1999; Costin et al. 2000; Rozefelds 2002; McDougall & Walsh 2008; Rozefelds et al. 2011). Most Australian specimens formerly identified as C. uniflora , including all those at CANB/CBG, have been assigned to species described in these recent studies. However, C. uniflora is still listed in the latest Census of the Queensland Flora (Bostock & Holland 2010) and in the Australian Plant Census (CHAH 2011) as occurring in all states except the Northern Territory. No nationwide taxonomic treatment of Craspedia is currently available. Ford et al. (2007) demonstrated that the New Zealand species of Craspedia form a monophyletic group nested in the Australian representatives of the genus and thus are likely to represent the descendants of a single long Schmidt-Lebuhn, Craspedia uniflora distance dispersal event with no subsequent re-colonization of Australia. Because the New Zealand lineage has diversified into at least six species since its arrival (Allan 1961), it appears a priori unlikely that a contemporary species such as C. uniflora would be found on both sides of the Tasman Sea. In this paper the status of Craspedia uniflora in Australia is examined to clarify whether Australian specimens identified as this species were merely overlooked when the taxonomy of Australian Craspedia was updated or whether at least some of them are conspecific with those from New Zealand. A recent quantitative study of morphological characters in the genus aimed to include specimens from all formally described native Australian species (Schmidt-Lebuhn & Milner 2013), and it was therefore important to determine whether C. uniflora needed to be included. Materials and methods All specimens labelled as Craspedia uniflora were requested as a loan from BRI, and a photograph of the type of the species was obtained from K. The morphology of the type as discernible from the photograph, the morphology of selected New Zealand specimens and the description of C. uniflora by Allan (1961) were compared against the morphology of Australian specimens previously identified as C. uniflora and the descriptions of and keys to Australian species (in particular Everett & Doust 1992a; Everett & Thompson 1992). The results of the quantitative analyses by Schmidt-Lebuhn & Milner (2013) were also taken into account. Common abbreviations in the specimen citation include National Park (NP). All specimens have been seen unless indicated n.v. Results As circumscribed by Allan (1961), Craspedia uniflora of New Zealand is a heterogeneous species of four varieties occurring from coastal rock crevices to subalpine herb-fields. It is characterized by a very distinctive margin of tangled long white hairs around the leaves 137 but overall plant size, glomerule size and flower colour are variable (yellow or white in the latter case). Under the typical variety, the type specimen at K is described as having leaves obovate- to lanceolate-spathulate, c. 7 x 1.5 cm, scapes c. 12 cm long and glomerules of c. 1.5 cm diameter, values that are rather small for a Craspedia. Visual examination of the photograph of the type showed it to match this description, with the single glomerule c. 1.4 cm in diameter and the longest intact leaf c. 6 cm long. A very distinctive margin of white hairs around the rosette and scape leaves is clearly visible. No Australian specimens of Craspedia are consistent with the above morphology and descriptions. The only native species of similar stature as the type of C. uniflora are C. alba J.Everett & Joy Thomps., C. glabrata (Hook.f.) Rozefelds and C. rosulata Rozefelds & A.M.Buchanan, but all of them differ from this species especially in regard to the indumentum. Whereas Craspedia lamicola J.Everett & Joy Thomps. has a margin of hairs superficially similar to that of C. uniflora around otherwise glabrous to subglabrous leaves it is a much more robust plant restricted to wet alpine habitats with significantly larger scapes, leaves and glomerules. Craspedia specimens from Queensland previously identified as C. uniflora were referable to C. canens J.Everett & Doust, C. haplorrhiza J.Everett & Doust, C. paludicola J.Everett & Doust and C. variabilis J.Everett & Doust. The species of Craspedia in Queensland can be determined with the following key, with the caveats that more work is needed to understand the circumscription and taxonomy of what is currently called C. canens and that C. variabilis may ultimately turn out to be a synonym of Tasmanian C. glauca. (Schmidt-Lebuhn & Milner 2013). 138 Austrobaileya 9(1): 136-139 (2013) Key for the identification of Craspedia species that occur in Queensland 1 Plants short-lived, with a single tap-root.2. C. haplorrhiza 1. Plants perennial, with numerous secondary roots.2 2 Tall plants up to 75 cm, with subglabrous to glabrous leaves, growing in swamps or drainage lines.3. C. paludicola 2. Smaller plants usually only to 50 cm, with hairy leaves, growing in drier habitats.3 3 Leaves with woolly indumentum.1. C. canens 3. Leaves with robust septate indumentum and often with glands but not woolly.4. C. variabilis Taxonomy 1. Craspedia canens J.Everett & Doust, Telopea 5: 37 (1992). Type: New South Wales. Central Tablelands: Wingecarribee [Wingicarribee] Swamp, 26 October 1971, L.A. Craven 1981 (holo: NSW 221409 n.v.-, iso: CANB; L n.v .). Additional specimens examined: Queensland. Darling Downs District: Stanthorpe, Feb 1916, Wright s.n. (BRI [AQ247074]); Stanthorpe, s.dat., Davidson s.n. (BRI [AQ247073, 247077]); “The Junction track”, Girraween NP, Oct 1974, McDonald 467 (BRI); Victoria. Port Phillip, s.dat., Mossman 321 (BRI). Note: Mossman 321 is very small and glabrescent, showing few of the diagnostic woolly hairs. 2. Craspedia haplorrhiza J.Everett & Doust, Telopea 5: 37 (1992). Type: New South Wales. Far South Western Plains: southern crossroad, north of Ranger’s Station, Kinchega National Park, 12 September 1989, A. Denham 39 (holo: NSW n.v .; iso: AD n.v). Specimens examined : Queensland. Darling Downs District: 10 miles [16.6 km] SE of St George, Aug 1947, Everist 3137 (BRI); New South Wales, c. 56 km S of Walgett, Lightning Ridge/Angledoon Road, Sep 1989, White s.n. (BRI [AQ504366]). 3. Craspedia paludicola J.Everett & Doust, Telopea 5: 35 (1992). Type: Victoria. Top End Reference Area Barmah State Park, Murray Valley Study Area Sector: F, sub-block: 41B, 28 September 1985, A.C. Beauglehole 81321 (holo: MEL 681491 n.v., iso: CANB; HO n.v., NSW n.v). Craspedia sp. (Girraween NP S.T.Blake 23643), synon. nov. (Bostock & Holland 2007, 2010). Additional specimens examined: Queensland. Darling Downs District: About 7 miles [11 km] E of Wyberba, Oct 1958, Pedley 297 (BRI); Girraween NP near Wyberba and Wallangarra, Nov 1971, Blake 23643 (BRI); Racecourse Creek, Dingo Swamp, Girraween NP, Aug 1995, Forster PIF17626 (BRI, CANB). Notes: The loan made available by BRI also included several additional specimens not previously identified as C. uniflora, among them three sheets of the undescribed phrase name taxon Craspedia sp. (Girraween NP S.T.Blake 23643) (Bostock & Holland 2007, 2010; CHAH 2011). They do not differ in any significant way from C. paludicola J.Everett & Doust, especially considering the morphological variability of the latter species over its range on both sides of the Bass Strait. Although Pedley 297 is an unusually small plant, its glabrous leaves and the habitat are typical of C. paludicola. 4. Craspedia variabilis J.Everett & Doust, Telopea 5: 36 (1992). Type: South Australia. South Eastern Region: c. 15 km south-east of Mt Burr Township, 7 September 1966, 1.B. Wilson 511 (holo: AD n.v.-, iso: CANB). Additional specimens examined: Western Australia. Mundaring Weir, Aug 1926, Bicks.n. (BRI [AQ247068]); Dalyup, 30 miles [50 km] W of Esperance, Sep 1971, Scarth-Johnson 820 (BRI). Queensland. Darling Downs District: Cons Plain, NE of Killarney, Oct 2000, Bean 16897 (BRI); Killarney, Sep 1937, Schindler 5 (BRI). Moreton District: turnstile gate above Grace’s Hut, near Mt Barney, s.dat., Leiper s.n. (BRI [AQ584399]). New South Wales. 7.4 km along Cochrane Road, Carrai NP, W of Kempsey, Dec 2007, Bean 27170 (BRI, CANB). Victoria. Melbourne, sine coll. s.n. (BRI [AQ247063]). South Australia. Near Kapunda, in 1888, Bailey s.n. (BRI [AQ247062]); Eyre Peninsula Region, Marble Range, slopes of North Block, Sep 1979, Weber 6018 (BRI). Schmidt-Lebuhn, Craspedia uniflora Discussion No Australian specimen previously labelled as C. uniflora was found to be consistent with the type or with descriptions (Allan 1961) and herbarium vouchers from New Zealand. Craspedia uniflora therefore is endemic to New Zealand, and the name should be removed from the Census of the Queensland Flora and the Australian Plant Census. This study shows that four species of Craspedia s.str. occur in Queensland: C. canens, C. haplorrhiza, C. paludicola and C. variabilis. Craspedia haplorrhiza is a widespread species of the western slopes and the margin of the arid zone, occurring in Queensland south from St George to around Bourke, Brewarrina and Walgett in New South Wales. The distribution of the other three species in the state is restricted to small areas in its south-eastern corner. Craspedia paludicola is known only from Girraween National Park, and C. canens from the same National Park and from some historical collections labelled as ‘Stanthorpe’. Acknowledgements I thank the Director of the Queensland Herbarium for the loan of material, Nicholas Hind and Elizabeth Woodgyer (Kew Royal Botanic Gardens) for moving the photographing of the type of Craspedia uniflora ahead of the schedule of the Global Plants Initiative to make it available for examination, and Paul I Forster, Brendan Lepschi, Joseph T. Miller and one anonymous reviewer for helpful comments on a previous version of this manuscript. References Allan, H.H. (1961). Flora of New Zealand. Volume 1. Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. New Zealand Government Printer: Wellington. Bostock, P.D. & Holland, A.E. (2007). Census of the Queensland Flora 2007. Queensland Herbarium, Environmental Protection Agency: Brisbane. - (2010). Census of the Queensland Flora 2010. Queensland Herbarium, Department of Environment and Resource Management: Brisbane. 139 CHAH [Council of Heads of Australasian Herbaria] (2011). Australian Plant Census. http://www. anbg. gov. au/chah/apc/ Costin, A., Gray, M., Totterdell, C. & Wimbush, D. (2000). Kosciuszko Alpine Flora , 2 nd edition. CSIRO Publishing: Melbourne. Curtis, W.M. (1963). The Student’s Flora of Tasmania Part 2 - Angiospermae: Lythraceae to Epacridaceae. St. David’s Park Publishing: Hobart. Everett, J. (1999). Craspedia. In N.G. Walsh & T.J. Entwisle (eds.). Flora of Victoria 4: 758-764. Inkata Press: Melbourne. Everett, J., Doust, A.N.L. (1992a). New species and a new combination in Pycnosorus (Asteraceae: Gnaphalieae). Telopea 5: 39-43. - (1992b). Four new Australian species of Craspedia sens, strict. (Asteraceae: Gnaphalieae). Telopea 5: 35-38. - (1992c). Craspedia. In G.J. Harden (ed), Flora of New South Wales 3: 221-226. NSW University Press: Sydney. Everett, J. & Thompson, J. (1992). New alpine and subalpine species in Craspedia sens, strict. (Asteraceae: Gnaphalieae). Telopea 5: 45-51. Ford, K.A., Ward, J.M., Smissen, R.D., Wagstaff, S.J. & Breitwieser, I. (2007). Phylogeny and biogeography of Craspedia (Asteraceae: Gnaphalieae) based on ITS, ETS and psbA-trnH sequence data. Taxon 56: 783-794. Forster, G. (1786). Florulae Insularum Australium Prodromus. J.C. Dieterich: Gottingen. McDougall, K.L. & Walsh, N.G. (2008). A conspectus of high-country Craspedia (Asteraceae: Gnaphalieae) of mainland south-eastern Australia. Muelleria 26: 3-10. Rozefelds, A.C. (2002). A new species and new combination in Craspedia (Asteraceae) from Tasmania. Telopea 9: 813-820. Rozefelds, A.C., Buchanan, A.M. & Ford, K.A. (2011). New species of Craspedia (Asteraceae: Gnaphalieae) from Tasmania and determination of the identity of C. macrocephala Hook. Kanunnah 4: 93-116. Schmidt-Lebuhn, A.N. & Milner, K.V. (2013). A quantitative study of morphology in Australian Craspedia (Asteraceae, Gnaphalieae). Australian Systematic Botany 26: 238-254. Stanley, T.D. & Ross, E.M. (1986). Flora of south¬ eastern Queensland. Volume 2. Queensland Department of Primary Industries: Brisbane. Cullen spicigerum (Domin) A.E.Holland (Fabaceae), a new combination and reinstatement of a north Queensland species Ailsa E. Holland Summary Holland, A.E. (2013). Cullen spicigerum (Domin) A.E.Holland (Fabaceae), a new combination and reinstatement of a north Queensland species. Austrobaileya 9(1): 140-145. Psoralea spicigera Domin has been relegated to synonymy under the Western Australian Cullen lachnostachys (F.Muell.) J.W.Grimes. This species is here reinstated and a new combination, Cullen spicigerum (Domin) A.E.Holland, is made. A revised description of this species is provided, along with notes on distribution and habitat and a key to the species of Cullen in Queensland. Key Words: Fabaceae, Cullen, Cullen spicigerum, Psoralea, Psoralea spicigera, Australia flora, Queensland flora, identification key, taxonomy, new combination Ailsa E. Holland, Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: Ailsa.Holland@science.dsitia.qld.gov.au Introduction Cullen Medik., tribe Psoraleeae , is a widespread genus of 32 species of which 26 occur in Australia (Grimes 1997; AVH 2013). The genus was revised by Grimes (1997) who placed Psoralea spicigera Domin in synonymy under Cullen lachnostachys (F.Muell.) J.W.Grimes, albeit without any explanatory notes as to why this decision was made. However, in his introduction, Grimes commented that “a couple of problems remain which will best be addressed through detailed field-work.. .There are several forms included in C. badocanum that might merit some taxonomic status, but which are incompletely known from few collections”. It is possible that this taxon is referred to here, as Grimes probably saw only a few specimens and only one of the specimens cited below was mentioned anywhere in his revision {Bolton 475 was determined by Grimes as C. pustulatum (F.Muell.) J.W.Grimes). Numerous new collections are all consistent with Domin’s description of Psoralea spicigera except for the stipules which are generally shorter than the 1 cm described by him. The species is however quite distinct and the main differences are listed in Table 1. This Accepted for publication 9 September 2013 north Queensland species is therefore here reinstated, and a new combination, Cullen spicigerum (Domin) Holland, is made. Materials and methods This paper is based on herbarium collections in the Queensland Herbarium (BRI). The species was examined in the field in 2012. A common abbreviation in the specimen citations is: NP (National Park). Taxonomy Cullen spicigerum (Domin) A.E.Holland, comb, nov.; Psoralea spicigera Domin, Biblioth. Bot. 89(2): 186-187 (1926). Type: Queensland. Cook District: Elizabeth Ck [Creek], 16 July [1872], W.Hann, Cape York Peninsula Expedition No. 40 (lecto: PR; isolecto: K, online image!;^^ Grimes [1997: 615]). Cullen sp. (Westmoreland C.H.Gittins 2491); Holland & Pedley (2013). Erect herbaceous shrubs to 3 m tall; bark grey. Branches striate, densely pubescent with short crisped white or grey hairs, moderately glandular with large raised light brown or blond glands that are generally sunken or flush with surface or pustulate but generally not stipitate (see below). Leaves heteroblastic; the proximal leaves pinnately trifoliolate, the Holland , Cullen spicigerum 141 Table 1. Comparison of morphological characters for Cullen spicigerum and putatively related taxa Character state C. spicigerum C. badocanum C. pustulatum C. lachnostachys indumentum dense, short crisped villose to lanate-villose glabrate to villulose velutinous or short-villose stipule length (mm) 3-5 4-12 4-12 5-8.5 inflorescence shape cylindrical subglobose cylindrical cylindrical inflorescence bract persistence persistent persistent deciduous or persistent deciduous flower length (mm) 6-10 8.5-15 7-10 10.5-14.5 lower calyx lobe length (mm) 5-8 7.7-14 2.2-3.5 3.5-6.5 pedicel length (mm) 0.5-2 0.5-3 0.5-3 4-7 distal leaves unifoliolate; petioles 5-20 mm long; rachis 3-15 mm long; both petiole and rachis striate, pubescent and with scattered glands. Leaflets lanceolate, rounded to cuneate at base, acute or obtuse and apiculate at apex; the upper surface sparsely pubescent or nearly glabrous, with scattered glands; lower surface moderately densely pubescent, glandular with larger stipitate and crateriform glands congregated on the veins. Terminal leaflets 25-80 mm long; 8-35 mm wide, length:width ratio 1.8-4.4. Lateral leaflets similar, 2/3 of the length of the terminal leaflet. Stipules triangular, reflexed, 3-5 mm long, 1-1.5 mm wide, glandular and pubescent on outer surface, glabrous on inner surface, with several obscure veins. Stipels absent. Pseudoracemes indeterminate, one per axil, compact, cylindrical, occasionally branched, 1-10 cm long, 1.5-2 cm wide, initially shorter than the subtending leaf, longer at maturity; peduncle 20-80 mm long; rachis with 9—16 flowering nodes, initially shorter than the peduncle, elongating through the growing season. Flowers usually in pairs, rarely 3 together, overlapping on the rachis; internodes 2- 8 mm long. Bracts persistent, initially erect, becoming relaxed, broadly ovate to suborbicular, 3-6 mm long, 2.5-4.5 mm wide, the apex abruptly narrowed into a slightly reflexed acuminate tip; outer surface sparsely hairy mainly on the margins and densely covered in large, light brown pustular glands; inner surface glabrous. Bracteoles absent. Pedicels 0.5-2 mm long, pubescent. Flowers 6-10 mm long, corolla shorter or slightly longer than the calyx. Calyx tubular- campanulate, somewhat flattened, 7-10 mm long, 5-lobed, purplish-green, pubescent with short crisped white or grey hairs (sometimes tending to villous but not obscuring veins or glands); glands scattered, large, pustular; calyx tube 2-3 mm long, rounded at base, with 10 raised veins of which 5 end in the lobe sinuses and 5 continue to lobe apices; adaxial 4 lobes triangular, acute, 2-4 mm long, the uppermost two slightly longer and partly joined; abaxial lobe 5-8 mm long, carinate, curving upwards over the keel to a long attenuate point. Corolla glabrous, purplish at the apex and fading to white at base; standard obovate, 6-9 mm long, 6-7 mm wide, slightly longer than wings and keel, emarginate, tapered to an indistinct claw at base; wings 142 oblong, 8-9 mm long including a 2-3 mm claw, c. 2.5 mm wide, rounded at apex; keel upturned and enclosed in the other petals, c. 8 mm long including a 3-4 mm claw and 2 mm wide, rounded at apex and curved along the lower edge, the two halves joined in the distal half. Ovary glabrous, glandular, stipitate. Fruit and flower remnant falling together (excluding bract). Fruit stipitate; stipe to 3 mm long; pod broadly elliptic, flattened, 3-3.5 mm long, 1.7-2.3 mm wide, glabrous or with a few marginal hairs, very densely glandular with pustular glands, the fruit wall strongly adhering to the seed. Seed reniform, 2.3-2.5 mm long, 3.2-3.8 mm wide, smooth, green to light brown, hilum minute. Fig. 1. Additional specimens examined : Queensland. Cook District: 1 km N of Old Laura Homestead, Lakefield NP, Aug 2012, Thompson ST12628 (BRI, MEL); near Old Laura Homestead, Lakefield NP, Jul 1993, Stanton 2 (BRI); Lakefield NP, 3.5 km along 12 Mile Waterhole Road, Jul 2010, McDonald KRM9701 (BRI, PE); c. 0.5 km from Kowanyama, Oct 1981, Birchley 127 (BRI); Cooktown Development Road, c. 0.5 km E of Boggy Creek, Apr 2001, Jago 5899 & Wannan (BRI, MEL, NSW); Wrotham Park, Trimbles Crossing, Walsh River, May 2012, McDonald KRM 12753 & Holland (BRI); Murphy’s Farm, Gilbert River, Aug 2001, Smith AS2038 (BRI); Near Gamboola Homestead, N of Chillagoe, Aug 1985, Bolton 475 (BRI); Van Lee Homestead, 65 km NNW of Mt Surprise, Jul 2001, Addicott EPA873 (BRI); Gilbert River, May 1954, Ever is t 5477 (BRI). Burke District: 14.5 km SE of Lotus Vale, on Lotus Vale to Stirling Road (Site 106), Jun 2001, Turpin GPT592 & Thompson (BRI); Westmoreland, 56 km from Wollogorang towards Burketown, Jul 1972, Gittins 2491 (BRI). North Kennedy District: Cape Cleveland section of Bowling Green Bay NP, S of Townsville, Jul Austrobail'ey a 9(1): 140-145 (2013) 1991, Bean 3459 (BRI, MEL, MEXU). Distribution and habitat: Cullen spicigerum occurs in open woodland of Corymbia, Eucalyptus or Melaleuca usually with a grassy understorey and generally on alluvial silty clay or loam soils, often on river banks, flood plains or disturbed areas. The species is endemic to north Queensland where it is widespread on southern Cape York Peninsula and west nearly to the Northern Territory border (Map 1). Phenology: This species flowers and fruits continuously through the cooler months from April through to August, occasionally as late as October. Notes: Cullen spicigerum is related to the widespread C. badocanum (Blanco) Verde, and C. pustulatum , and the Western Australian endemic C. lachnostachys. It differs from C. badocanum by the cylindrical pseudoracemes and the shorter stipules, flowers and abaxial calyx lobe, and from C. pustidatum by the dense indumentum (especially on the calyx), shorter pseudoracemes, shorter stipules and much longer abaxial calyx lobe. It differs from C. lachnostachys by the persistent and smaller bracts, the shorter pedicels, flowers and stipules, and the shorter pubescence (especially on the calyx) and the longer abaxial calyx lobe. Conservation status: Cullen spicigerum is Least Concern using the IUCN (2001) criteria. Key to the species of Cullen in Queensland 1 Leaflets 3 or 5, palmately arranged. C. tenax 1. Leaflets 3, pinnately arranged, or leaves unifoliolate.2 2 Plants covered in long spreading white hairs > 1 mm long; bracts lanceolate and more than 6 mm long.C. plumosum 2. Plants glabrous or if hairs present then < 1 mm long; bracts ovate to orbicular, less than 6 mm long.3 3 Leaves all unifoliolate.4 3. Leaves trifoliolate, or both unifoliolate and trifoliolate leaves present.6 4 Inflorescence an elongate spike; calyx shorter than the fruit.C. leucanthum 4. Inflorescence subglobular on a shortened rachis; calyx longer than the fruit.5 Holland , Cullen spicigerum 143 5 Whole plant densely covered in black pustular glands; leaves distinctly denticulate; flowers to 7 mm long.C. balsamicum 5. Sparsely to moderately glandular with brown glands and flush with surface; leaves entire or weakly toothed; flowers 9-15 mm long.C. badocanum 6 Glands on stems and petioles pustular or stipitate.7 6. Glands on stems and petioles flush with surface, or sunken.8 7 Calyx much shorter than corolla, the hairs confined to veins; abaxial calyx lobe scarcely longer than the others.C. pustulatum 7. Calyx longer or shorter than corolla, moderately hairy; abaxial calyx lobe c. 3 mm longer than the others.C. spicigerum 8 Leaflets mostly more than 5 cm long, lanceolate, entire.9 8. Leaflets mostly less than 5 cm long, ovate to lanceolate, denticulate, dentate, erose or serrate.10 9 Plants glabrous or nearly so; corolla much longer than calyx, predominantly white, cream or yellowish with a purple blotch.C. leucanthum 9. Plants hairy to some degree; corolla as long or slightly shorter than calyx, purple to white.C. spicigerum 10 Calyx 2-4 mm long, shorter than the fruit.C. cinereum 10. Calyx 3-8 mm long, longer than the fruit.11 11 Calyx sparsely hairy with appressed white hairs in rows, and much longer than corolla.C. graveolens 11. Calyx densely hairy with spreading or appressed white or black hairs, and shorter than or equal to corolla.12 12 Inflorescence condensed, usually shorter than the subtending leaf; peduncles less than 4 cm long; corolla usually white.C. discolor 12. Inflorescences elongated and interrupted, much longer than the subtending leaf; peduncles longer than 4 cm; corolla usually purple, pink or blue.13 13 Plants strictly prostrate.C. patens 13. Plants erect or ascending, bushy.14 14 Entire plants densely white velvety hairy, the glands of the leaves obscured . . C. pallidum 14. Only calyces with dense white hair; the glands on the leaves apparent . . C. australasicum Acknowledgements I would like to thank Les Pedley for drawing this species to my attention and Keith McDonald for his excellent specimens, information and on ground guidance. I also acknowledge Ruth Clark who located the isolectotype at Kew and arranged for an image to be provided. Will Smith prepared the illustrations and map. References AVH (Australia’s Virtual Herbarium) (2013). Council of Heads of Australasian Herbaria, viewed 16 August 2013, <http://avh.ala.org.au/> Domin, K. (1926). Psoralea L. Bibliotheca Botanica 89(2): 186-187. Grimes, J.W. (1997). A revision of Cullen (Leguminosae: Papilionoideae). Australian Systematic Botany 10: 565-648. Holland, A.E. & Pedley, L. (2013). Fabaceae. In P.D. Bostock & A.E. Holland (eds). Census of the Queensland Flora 2013. Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts: Brisbane. https://data.qld.gov.au/dataset/census-of-the- queensland-flora-2013 Iucn (2001). Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission. IUCN: Gland, Switzerland/Cambridge, U.K. 144 Austrobaileya 9(1): 140-145 (2013) Fig. 1 . Cullen spicigerum. A. branch xl. B. leaflet lower surface xl6. leaflet upper surface xl6. D. stipule x8. E. bract x8. F. calyx x6. G. standard petal x6. H. keel petal x6.1. wing petal x6, J. staminal column x6. K. ovary and style enclosed in staminal column x6. L. fruit x8. K. seed x8. All from McDonaldKRM9701 (BRI). Del. W.Smith. Holland, Cullen spicigerum Map. 1 . Distribution of Cullen spicigerum in Queensland. Austrobaileya 9(1): 146-147 (2013) 146 SHORT COMMUNICATION Celtis strychnoides Planch, an earlier name for Celtis australiensis Sattarian (Ulmaceae) Gordon P. Guymer Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: Gordon.Guymer@science.dsitia.qld.gov. au Celtis australiensis Sattarian was described by Sattarian (2005) as occurring in north west Western Australia, the Northern Territory and north west Queensland and was distinguished from Celtisphilippensis Blanco by its elliptic, cordate, pilose leaves (2-7 x 1.5-5 cm), spinose leaf apices and margins. This name has subsequently been incorrectly applied to all Australian specimens previously known as Celtis philippensis (CHAH 2007). Sattarian (2005) referred to the Flora of Australia treatment of Celtis by Hewson (1989) but overlooked the name Celtis strychnoides Planch, given by Hewson as a synonym of Celtis philippensis. Planchon (1848) described Celtis strychnoides from a collection of Allan Cunningham’s from Careening Bay, Kimberley, northern Western Australia. He distinguished it from Celtis philippensis by its small, pubescent, ovate or rarely broadly ovate leaves that were barely 1.5 inches (4 cm) long. Examination of the type collections and images of Celtis strychnoides , C. australiensis and specimens of Celtis at BRI, MEL and NSW confirm that C. strychnoides is conspecific with C. australiensis and as such is the earlier name. Celtis strychnoides Planch., Annales des Sciences Naturelles, Botanique ser. 3, 10: 306-307 (1848). Type: [Western Australia] Careening Bay, north west Australia, January 1820, A. Cunningham 193 in herb. Hook, (holo: K [852750, 852752 images !] , iso: BM [838618, 838619, 799835 images !], BRI p.p , MEL p.p ). Accepted for publication 2 September 2013 Celtis australiensis Sattarian, Blumea 50: 499 (2005), synon. nov. Type: Western Australia. Northern Province: Geikie Gorge, near gorge mouth, 14 May 1992, I. Telford 11722 (holo: CANB n.v., iso: BISH n.v., BRI, L n.v., NSW, PERTH n.v.). Illustration : Sattarian & van der Maesen (2005: 500, Fig. 1). Additional specimens examined : Western Australia. N end of Bougainville Peninsula, Apr 1982, Kenneally 8076 (BRI); Walsh Point, Port Warrender, May 1981, Tracey 15170, 15075 (BRI); Port Warrender, Oct 1974, Rodd 2900 (NSW); Surveyor’s Creek Road, May 1981, Tracey 15122 (BRI); Lone Dingo, May 1981, Tracey 1500 (BRI); Merten’s Falls, May 1981, Tracey 15062 (BRI); Near Crusher Pool, May 1981, Tracey 15233 (BRI); Cambridge Gulf, in 1886, Ranford & Nyulasy 18 & 36 (MEL); Wyndham, in 1903 & Nov 1906, Fitzgeralds.n. (NSW); Cockburn Range, 13 km W of King River, July 1974, Carr 3316 & Beaiiglehole 47094 (NSW); 4 km S of junction of Neville Creek & Calder River, May 1983, Fell 9 (BRI); Cape Leveque, Jul 1973, Webb & Tracey 13141 (BRI); King’s [King] Sound, in 1869, Hughan s.n. (MEL 2210491, 2210489, 2210481); ditto, in 1888, Froggart s.n. (NSW); Napier Range, Jul 1973, Tracey 14807 (BRI); Mornington Wildlife Sanctuary, NE of Fitzroy Crossing, Apr 2005, Murphy MULE453 & Legge (BRI); Geikie Gorge, 16 km NE of Fitzroy Crossing, Jun 1970, Briggs 3671 (NSW); ditto, May 1981, Tracey 15143 (BRI); 10 miles [16 km] SW of Lissadell Station, Aug 1949, Perry 2672 (MEL); S of Ord River Station, Aug 1973, Tracey 14100 (BRI). Northern Territory. Victoria River, Dec 1855, Mueller s.n. (K 852751, MEL 2210482, 2210475); Limestone Gorge, Gregory NP, Apr 1990, Evans 3091 (MEL); Gregory NP, Apr 1996, Booth 1696 (MEL); ditto , Apr 1996, Duretto 1187 (MEL); ditto , Apr 1996, Foreman 2249 (MEL); ditto , Apr 1996, Jones 83 (MEL); 5 km S of Timber Creek, May 1983, Briggs 904 (MEL); 15 miles [24 km] SSE of Timber Creek, Jun 1952, Perry 2869 (BRI); 41 km SSE of Turkey Creek, Jul 1984, Scarlett 84-334 (MEL); 6 miles [10 km] NW of Katherine, Adams 806 (NSW); 11 miles [18.3 km] S of Katherine, Feb 1961, McKee 8487 (NSW); 16-Mile Cave Reserve, S of Katherine, May 1978, Webb & Tracey 12588 (BRI); 4 miles [6 km] E of Elsey, Oct 1958, Chippendale 5065 (BRI, NSW); 15 miles [24 km] S of MacArthur River Station, Jul 1949, Perry 1732 (BRI, 147 MEL); near head of Kilgour River, Sep 1911, Hill 556 (MEL); Settlement Creek, Jun 1922, Brass 161 (BRI). Queensland. Burke District: 39 km SW of Hells Gate, Apr 2006, Thompson WES384 & Edginton (BRI); 17 km W of Adel’s Grove, Lawn Hill NP, Nov 1983, Buttock UNSW14474 & St George (MEL); 20.2 km S of Musselbrook, Apr 1995, Thomas & Johnson MRS469 (BRI); Lawn Hill NP, May 1988, O’Keefe 11 (BRI); 1 km N of Gregory River, Riversleigh, May 1970, Webb & Tracey 1349 (BRI); 29 km NW of Norfolk Station, Apr 2005, Kelman DTK1021 (BRI); Seymour River, vicinity of Nicholson River, Jul 1886, Dittrich s.n. (BRI [AQ88147]); 7 km NE of Camooweal, Jul 2006, Booth CAM29-1 & Kelman (BRI). Distribution : Celtis strychnoides occurs in the Kimberley region of northern Western Australia, central northern Northern Territory and north western Queensland. Habitat : The species occurs in semi-deciduous notophyll vine thickets or woodlands usually in rocky situations on slopes or hills in fire refugia, on limestone, basalt or sandstone. Notes: TheNorthern Territory and Queensland collections from higher rainfall areas (>1100 mm mean annual rainfall) that have glabrous, ovate or oblong leaves 6-18 x 3.5-8.5 cm and that were included in C. australiensis by CHAH (2007) are referrable to Celtis philippensis. These collections may represent a species distinct from C. philippensis but a revision of this species complex is required to resolve the taxonomy The Cunningham sheets at BRI and MEL include specimens of both C. strychnoides and C. philippensis. Celtis strychnoides can be distinguished from Australian material of Celtis philippensis by the following key: 1 Leaf blades ovate or broadly ovate, acute, 2-7 x 1.5-4 cm, young leaves sparsely pubescent, glabrescent except veins and leaf base below; base rounded or cordate; petioles and branchlets pubescent with white erect hairs 0.1-0.3 mm long, petioles 3-6 mm long; juvenile leaves with spinose margins, occasionally adult leaves with short marginal spines; small tree to 4-8 (-10) m. WA, NT & NW Qld.Celtis strychnoides 1. Leaf blades oblong to ovate, acuminate, 6-18 x 3.5-8.5 cm, glabrous; base cuneate to truncate; petioles glabrous, occasionally sparsely pubescent, 5-8 mm long; juvenile leaves with serrate or crenate margins; tree 4-25 m. Coastal NT & N & E Qld.Celtis philippensis Acknowledgements I thank the Directors and staff of MEL and NSW for allowing access and examination of their collections. References CHAH (2007). Australian Plant Census. Council of Heads of Australasian Herbarium, http://www. anbg. gov. au/chah/apc/ Hewson, H.J. (1989). Ulmaceae. In AS. George (ed.). Flora of Australia 3: 10-13. Australian Government Publishing Service: Canberra. Planchon, J.E. (1848). Sur les Ulmacees, Celtis. Annales des Sciences Naturelles, Botanique ser. 3, 10: 283-316. Sattarian, A. & van der Maesen, L. J.G. (2005). Two new species of Celtis (Celtidaceae) from Australia and Madagascar. Blumea 50: 499-503. Austrobaileya 9 ( 1 ): 148-149 ( 2013 ) 148 SHORT COMMUNICATION Reinstatement of Rapanea leucobrachya P.Royen (Myrsinaceae) from New Guinea, with a new combination as Myrsine leucobrachya (P.Royen) P.I.Forst. Paul I. Forster Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: Paul. For ster @science. dsitia. qld. gov. au Van Royen (1982) named a number of new species of Rapanea Aubl. from montane areas of New Guinea. Many of these were subsequently reduced to synonymy by Sleumer (1986) when he revised the New Guinea species. The New Guinea species recognised by Sleumer (1986) were uncritically transferred to Myrsine L. by Pipoly (Takeuchi & Pipoly 2009), together with the addition of one new species {Myrsine warrae W.N.Takeuchi & Pipoly). During curation of New Guinea Myrsinaceae at BRI it became obvious that Sleumer’s treatment of certain Myrsine species was overly conservative. This pertained particularly to his reduction to synonymy of Rapanea leucobrachya P.Royen and R. wapu P.Royen under R. rhombata P.Royen without any accompanying reasons. Whilst the recognition of three species for this group of montane populations from the Eastern and Western Highlands of Papua New Guinea may be too liberal, recognition of only one is not supported by easily discernible morphological differences such as leaf shape, venation and gland morphology. Hence, two species, M. rhombata (P.Royen) Pipoly and M. leucobrachya (P.Royen) P.I.Forst. are recognised herein. Myrsine rhombata (P.Royen) Pipoly, Edinburgh J. Bot. 66: 462 (2009); Rapanea rhombata P.Royen, Alpine FI. New Guinea 3: 1976 (1982). Type: Papua New Guinea. Accepted for publication 25 July 2013 Eastern Highlands Province: Mt Wilhelm, east slopes, 28 July 1959, L.J.Brass 30757 (holo: L, online image!; iso: US, online image!). Specimens examined : Papua New Guinea. Eastern Highlands Province: Kurumigl-bokwa, Kuski River above Toromambuno Mission, Upper Chimbu, Aug 1957, Pullen 324 (BRI); Mt Wilhelm, Lake Piunde, Aug 1957, Robbins 684 (BRI). Western Highlands Province: track to Mt Hagen, E of Mt Kigum, Apr 1977, Vinas & Veldkamp LAE59812 (BRI). Central Province: Murray Pass, Wharton Range, Jun-Sep 1933, Brass 4546 (BRI); Murray Pass, Aug 1970, Foreman & Lelean NGF48412A (BRI); Mt Victoria area, Iawan Swamp, c. 2 km SSW of Mt Service, May 1976, van Royen 11033 (BRI). Myrsine leucobrachya (P.Royen) P.I.Forst. comb, nov.; Rapanea leucobrachya P.Royen, Alpine FI. New Guinea 3: 1975 (1982). Type: Papua New Guinea. Southern Highlands Province: Mt Ialibu, slope to SW of Repeater Station, 10 August 1972, P.F. Stevens & D.B.Foreman LAE55853 (holo: K, online image!; iso: BRI; L, online image!). Rapanea wapu P.Royen, Alpine FI. New Guinea 3: 1971 (1982), synon. nov. Type: Papua New Guinea. Western Highlands Province: Northern slope of Sugarloaf complex (along Wapu River), 21 July 1960, R.D.Hoogland 7232 & R.Schodde (holo: L, online image!; BM, online image!). Additional specimens examined : Papua New Guinea. West Sepik Province: Bielga Range, W of Folongonom, second camp below Tamanagabip on Busilmin Track, May 1975, Vinas & Wiakabu LAE59529 (BRI). Eastern Highlands Province: track to Mt Michael, 5 km SW of Lufa, Apr 1982, Kairo 458 (BRI). Western Highlands Province: around the base camp, Mt Kigum, Apr 1977, Vinas & Veldkamp LAE59836 (BRI). Central Province: E slope to Lake Myola No 1, Sep 1973, Croft & Lelean NGF34965 (BRI). 149 The two species can be distinguished by the following key. Lamina rhombate-obovate, tip acuminate to retuse, below with numerous elongate glands, round glands along margins; 2° lateral veins obscure or very indistinct, 9-12 per side of 1° vein (midrib). Bracteoles broadly ovate to elliptic-ovate, 1-1.5 x 0.8-1.2 mm. Lamina elliptic, rhombate-elliptic to obovate, tip acuminate to obtuse, below with numerous nail-like glands on either side of 1° vein; 2° lateral veins prominent and easily discernible above and below, 8-14 per side of 1° vein (midrib), joined and forming an intramarginal vein, 3° veins prominent and reticulate below. Bracteoles ovate, 0.8-2.2 x 1-1.2 mm. References Sleumer, H. (1986). A revision of the genus Rapanea Aubl. (Myrsinaceae) in New Guinea. Blumea 31:245-269. Takeuchi, W. & Pipoly, J. (2009). Nomenclatural transfers in East Malesian Myrsine (Myrsinaceae), and the description of M. warrae , a distinctive new species from ultrabasic environments in Papua New Guinea. Edinburgh Journal of Botany 66: 459-467. Van Royen, P. (1982). The Alpine Flora of New Guinea. Volume 3: taxonomic part Winteraceae to Polygonaceae. J. Cramer: Vaduz. M. rhombata M. leucobrachya Austrobaileya 9 ( 1 ): 150-152 ( 2013 ) 150 SHORT COMMUNICATION Emmenospermapancherianum Baill. (Rhamnaceae) newly recorded for Australia A.R. Bean Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. E-mail: tony. bean@science.dsitia.qld.gov.au Emmenosperma F.Muell. is a genus of five species found in northern and eastern Australia, New Guinea, Sulawesi, New Caledonia and Fiji. It was described by Mueller (1862) based on Emmenosperma alphitonioides F.Muell. from Australia. Soon after, Bentham (1863) described another Australian species, E. cunninghamii Benth. Baillon described E. pancherianum Baill. from New Caledonia in 1874. In the 20 th century, two additional species, originally described in the genus Colubrina Rich, ex Brongn., were transferred to Emmenosperma , viz. E. micropetalum (A.C.Sm.) M.C.Johnst. and E. papuanum (Merr. & L.M.Perry) M.C.Johnst. In 2005, an unusual Emmenosperma specimen was submitted to the Queensland Herbarium, collected from Mt Zero-Taravale Sanctuary near Townsville. It was very unlike the two well-known Australian species, and an ensuing literature search suggested a strong affinity with E. pancherianum. Flowering and fruiting material was obtained from the Mt Zero-Taravale population in 2009, and this material has been compared with specimens of E. pancherianum collected from New Caledonia. The comparison revealed that there are no significant morphological differences between the New Caledonian material and the Mt Zero-Taravale material. Emmenosperma pancherianum Baill., Adansonia 11: 269 (1874). Type citation: ‘in Austro-Caledonia ubi in sylvis collium [hill- forests of southern Caledonia], leg. Pancher (herb.)’. Type: Nouvelle Caledonie, May 1865, J.A.I. Pancher s.n. (lecto: P 06886541 [here chosen]). Illustration : Sprague (1915: t. 3027). Shrub or small tree to 5 m. Young branches with dense indumentum of appressed simple hairs, older branches glabrous. Stipules caducous, hairy, triangular, c. 0.8 mm long. Leaves 16-40 x 8-22 mm, lamina obovate, with 2-4 obscure pairs of lateral veins at 45-70° to midrib, base cuneate, apex obtuse or emarginate; upper surface glabrous (except at base on midrib), very glossy; lower surface less glossy, glabrous; petioles 1.5-4 mm long, with appressed hairs. Inflorescences axillary, borne on short lateral branchlets 2-5 mm long with rusty hairs near apex; rachises absent or up to 2 mm long. Flowers 5-merous. Hypanthium and sepals with antrorse appressed brownish hairs; sepals deflate, 1-1.2 mm long, obtuse. Petals elliptical, 0.9- 1.1 mm long, white, glabrous, hooded. Ovary glabrous, disc inconspicuous at base of ovary; style bulbous, 0.6-0.8 mm long, 2-lobed at apex. Fruiting pedicels sparsely hairy, 2.5-5(-6.5) mm long; fruits ± globose, 5-6 mm long, 2-locular; endocarpids 2, splitting longitudinally from the apex for up to half their length. Seeds flattened ellipsoidal, 4-5.7 mm long, surface smooth and shiny, orange to red, remaining attached to fruiting pedicels after endocarpids have been shed. Accepted for publication 6 May 2013 Bean, Emmenosperma pancherianum Additional specimens examined : New Caledonia: Nekoro, Aug 2003, Dagostini & Rigault 703 (NOU); Gouaro Deva, Oct 2005, Dagostini 979 (NOU); Noumea, Jun 1909, Franc 1382 (BRI); Noumea, presqu’ile Ducos, Apr 1970, McKee 21778 (NOU); Tontouta, Jul 1971, McKee 24789 (NOU); Ngoue, Paita, Mar 1981, McKee 38781 (NOU); Plaine des gaiac, Jul 1987, Veillon 6411 (NOU); Mt Tereka, Noumea, Jul 1991, Veillon 7405 (NOU); Poya-Nord, Oct 1998, Veillon 8134 (NOU). Australia: Queensland. North Kennedy District: Near Return Creek falls, ‘Taravale’, NW of Townsville, May 2009, Bean 28865 & Jensen (BM, BRI, CANB, CNS, MO, NSW, P); Return Creek Falls, ‘Taravale’, 80 km WNW of Townsville, Jul 2005, Camming s.n. (BRI [AQ612648]). Cultivated. Botanic Gardens, Sydney, Jan 1921, White s.n. (BRI [AQ23692]). Distribution and habitat : This species is distributed along the west coast of New Caledonia, between Koumac and Noumea (Barrault 2006). In Australia, it is known from a single population at Mt Zero- Taravale Sanctuary north-west of Townsville, Queensland. In New Caledonia, it inhabits the “dry forests” of the western coast (Barrault 2006), a habitat that has been decimated for grazing and agriculture (Hequet 2010). In Queensland, it grows on the steep south¬ facing slope of a granite ridge, in open forest dominated by Araucaria cunninghamii Mudie and Eucalyptus granitica L.A.S.Johnson & K.D.Hill. Phenology: Flowers April to October; fruits September to March. Typification : The protologue specified ‘Pancher’ as the collector. Several Pancher specimens at P predate the publication date. 151 P 06886541 was chosen as the lectotype because it is a very good flowering specimen, and attached to the sheet is a handwritten description of the species, most likely by the author of the name. Notes: Emmenosperma pancherianum provides one more instance of the well known floristic connection between tropical eastern Australia and New Caledonia (Morat et al. 1986; Ladiges & Cantrill 2007). While a large proportion of New Caledonian plant species are endemic, there are numerous examples of species that occur in both New Caledonia and eastern Australia. These include not only littoral species that are dispersed by ocean currents e.g. Premna serratifolia L. and Cerbera manghas L., but others such as Carissa ovata R.Br., Croton insularis Baill., Dry petes deplanchei (Brongn. & Gris) Merr., Polyalthia nitidissima (Dunal) Benth., Psydrax odorata (G.Forst.) A.C.Sm. & S.P.Darwin, Pterocaulon serrulatum (Montrouz.) Guillaumin and Rhamnella vitiensis (Benth.) A.C.Sm. Conservation status: Emmenosperma pancherianum is listed as ‘Vulnerable’ under the criteria of the IUCN Red List, based on its New Caledonian occurrence (Hequet 2010). In Australia, it is currently known from a single population comprising between six and 10 individuals. Its recommended status, based on the Australian population only, is Critically Endangered, Criterion D (IUCN 2001 ). Key to the species of Emmenosperma in Australia 1 Inflorescences axillary, borne in dense ± spherical clusters, rachises lacking or < 2 mm long; petioles 1.5-4 mm long Qld, New Caledonia . E. pancherianum 1. Inflorescences predominantly terminal, some in upper leaf axils, paniculate to corymbose, rachises 13-70 mm long; petioles 5-27 mm long. 2 2 Bracts and bracteoles conspicuous at anthesis; style 2-fid; fruiting pedicels (3-)4-5.5 mm long; fruits 2-locular Qld, NSW .E. alphitonioides 2 . Bracts and bracteoles inconspicuous at anthesis; style 3-fid; fruiting pedicels 1.5-2.5(-4) mm long; fruits 3-locular Qld, NT, WA, New Guinea .E. cunninghamii 152 Austrobaileya 9 ( 1 ): 151-153 ( 2013 ) Acknowledgements I am grateful to Rigel Jensen, botanist with the Australian Wildlife Conservancy, for taking me to the ‘Taravale’ population. Thanks are also due to the Director of NOU for the loan of specimens and to Juergen Kellerman for helpful comments and suggestions. References Barrault, J. (2006). Plantes des Forets Seches de Nouvelle-Caledonie. Programme Forets Seches et Centre d’Initiation a FEnvironnement: Noumea. Bentham, G. (1863). Emmenosperma. In Flora Australiensis 1: 414-415. L.Reeve: London. Hequet, V. (2010). Emmenosperma pancherianum. In IUCN Red List of Threatened Species. Version 2012.2. <www.iucnredlist.org>. Downloaded on 20 October 2012. Iucn (2001). Red List Categories and Criteria: Version 3.1. IUCN Species Survival Commission. IUCN: Gland, Switzerland/Cambridge, U.K. Ladiges, P.Y. & Cantrill, D. (2007). New Caledonia - Australian connections: biogeographic patterns and geology. Australian Systematic Botany 20: 383-389. Morat, P, Veillon, J.-M. & Mckee, H.S. (1986). Floristic relationships of New Caledonian rainforest phanerogams. Telopea 2: 631-679. Mueller, F. (1862). Emmenosperma. Fragmenta Phytographiae Australiae 3: 62-63. Government Printer: Melbourne. Sprague, T.A. (1915). Emmenosperma pancherianum , Baill. Hooker’s leones Plantarum 31: t. 3027. Austrobaileya 9 ( 1 ): 153-154 ( 2013 ) 153 SHORT COMMUNICATION Euphorbia operta Halford & W.K.Harris (Euphorbiaceae), a new name for Euphorbia occulta Halford & W.K.Harris, nom. illeg. and lectotypification of the name Euphorbia australis var. subtomentosa Domin D.A. Halford & W.K. Harris d- Queensland Herbarium, Department of Science, Information Technology, Innovation and the Arts, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia In our recent taxonomic treatment of Euphorbia section Anisophyllum Roeper (Euphorbiaceae) in Australia (Halford & Harris 2012) we incorrectly published the name E. occulta Halford & W.K.Harris for an endemic Australian species from northwest Queensland. We were unaware that the name had already been applied by Klotzch (1856) to a North America species, so the use of the name by us was illegitimate, being a much later homonym. Therefore a new name Euphorbia operta Halford & W.K.Harris is provided here for the Australian species. The specific epithet is from Latin, opertus , hidden from sight, concealed, in reference to the concealment of the involucral glands by the erect hood-like gland appendages of this species. Euphorbia operta Halford & W.K.Harris, nom. nov.; Euphorbia occulta Halford & W.K.Harris, Austrobaileya 8: 529-530 (2012), nom. illeg., non Klotzsch, Bot. Voy. Herald [Seemann] 7-8: 277 (1856). Type: Queensland. Burke District: 12.3 km along Lake Moondara Road from Barkly Highway, N of Mt Isa, 1 July 2011, D. Halford QM524A (holo: BRI, iso: DNA, K, MEL, MICH, NT, distribuendi). We also overlooked the need to lectotypify the name Euphorbia australis var. subtomentosa Domin in order to fix the application of this name. Domin’s (1927) description of E. australis var. subtomentosa was based on material collected by Dr E. Clement from between the Ashburton and De Grey Rivers in Western Australia. There are four sheets that were seen by Domin, one in PR [528303] and three in K [186468, 186496, 186467], These comprise all the original material for the name that we have been able to locate. Careful examination of the four sheets clearly indicated that the specimens affixed to them belong to two different taxa, namely those currently known as E. australis var. subtomentosa (K 186468, PR 528303 (lower two fragments)) and E. vaccaria Baill. var. vaccaria (K 186496, 186467, PR 528303 (upper three fragments)). There is little doubt that all the material was used by Domin in his formulation of the original description of E. australis var. subtomentosa. The specimen numbered K 186468 is selected here as lectotype of E. australis var. subtomentosa. If either of the sheets (K 186496, 186467) or fragments (PR 528303 (upper three fragments)) were chosen as lectotype, E. australis var. subtomentosa would have to be placed in synonymy under E. vaccaria and a new name would be required for what is currently known as E. australis var. subtomentosa. Euphorbia australis var. subtomentosa Domin, Biblioth. Bot. 89(4): 310 (1927 ‘1926’). Type: Western Australia. Inter Ashburton et De Gray [Grey] River s.d., E. Clement s.n. (lectotype [here chosen]: K 186468; isolecto: PR 528303 (lower two fragments)). Accepted for publication 10 October 2013 Halford & Harris, Euphorbia names References Austrobaileya 9(1): 153-154 (2013) Domin, K. (1927). Euphorbia L. In Beitrage zur Flora und Pflanzengeographie Australiens, 1. Tiel, 3. Abt. Bibliotheca Botanica 89(4): 306-312. Halford, D.A. & Harris, W.K. (2012). A taxonomic revision of Euphorbia section Anisophyllum Roeper (Euphorbiaceae) in Australia. Austrobaileya 8: 441-600. Klotzsch, J.F. (1856). Euphorbiaceae. InB.C. Seemann, Botany of the Voyage of H.M.S. Herald, under the Command of Captain Henry Kellett, R.N., C.B., during the years 1845-1851 , pp. 276-278. L. Reeve: London.