The cover of Nuytsia Volume 30 showcases a golden display of Cephaiipterum drummondii and Waitzia acuminata at Karara Rangeland Park in 2016. Photograph by Steve Dillon Nuytsia WESTERN AUSTRAEIAN HERBARIUM VOLUME 30 2019 DEPARTMENT OF BIODIVERSITY, CONSERVATION AND ATTRACTIONS WESTERN AUSTRAEIA Nutysia Editorial Committee Nuytsia is an open access, peer-reviewed journal that publishes original research on the systematics, taxonomy and nomenclature of Australian (particularly Western Australian) plants, algae and fungi. Descriptions of taxa, revisions, identification guides, nomenclatural and taxonomic issues, systematic analyses and classifications, censuses, and information on invasive species are all considered. Published papers and information for authors can be downloaded from the Nuytsia website at https:// fiorabase.dpaw.wa.gov.au/nuytsia. Managing Editor Scientific & Copy Editor Production Editor Associate Editors Curation assessment Conservation assessment Kelly Shepherd John Huisman Kelly Shepherd Juliet Wege Terry Macfarlane Barbara Rye Meriel Falconer Anthea Jones Melanie Smith Published by the Department of Biodiversity, Conservation and Attractions, Eocked Bag 104, Bentley Delivery Centre, Western Australia 6983. Enquiries regarding subscriptions and submissions should be addressed to: The Managing Editor, Nuytsia Email: nuytsia@dbca.wa.gov.au Postal address; Western Australian Herbarium Biodiversity and Conservation Science Department of Biodiversity, Conservation and Attractions Eocked Bag 104, Bentley Delivery Centre Western Australia 6983 © Copyright of the Department of Biodiversity, Conservation and Attractions 2019. All material in this journal is copyright and may not be reproduced except with the written permission of the publishers. ISSN 0085-4417 (print) ISSN 2200-2790 (online) GOVERNMENT OF WESTERN AUSTRALIA Department of Biodiversity, Conservation and Attractions CONTENTS Updates to Western Australia’s vascular plant census for 2018. J.M. Percy-Bower and C.M. Parker.1 Acacia corusca (Fabaceae: Mimosoideae), a new species from the Pilbara bioregion in north-western Australia. J.P Bull, S.J. Dillon and D.R. Brearley.19 Gastrolobium sp. Harvey (G.J. Keighery 16821) (Fabaceae) is not distinct from G. capitatum. R. Butcher and M. Hislop.23 Eremophila oldfieldii subsp. papula, E. sericea and E. xantholaema (Scrophulariaceae), three new taxa from Western Australia. A.P Brown and R.W Davis.25 An update to the taxonomy of the Western Australian genera of the Myrtaceae tribe Chamelaucieae. 6. Scholtzia. B.L. Rye.33 A revision of the tiumdins {Tribonanthes, Haemodoraceae). E. J. Hickman and S.D. Hopper.87 A taxonomic review of the Styphelia xerophylla group (Ericaceae: Epacridoideae: Styphelieae). M. Hislop and C. Puente-Eelievre.155 Elionurus tylophorus (Poaceae: Paniceae: Andropogoneae), a new species from the Kimberley region of Western Australia. M.D. Barrett and T. Handasyde.177 A key to the species of Thomasia (Malvaceae: Byttnerioideae). C.F. Wilkins and K.A. Shepherd.195 Two new Western Australian species segregated from Banksia densa (Proteaceae). K.R. Thiele.203 Additions to Eremophila (Scrophulariaceae). R.J. Chinnock.215 Triodia veniciae (Poaceae), a new species from the Pilbara region. Western Australia. M.D. Barrett.221 Styphelia quartzitica and S. rectiloba (Ericaceae: Epacridoideae: Styphelieae), two new, morphologically anomalous species of restricted distribution. M. Hislop.229 Calandrinia monosperma and C. uncinella (Montiaceae), two new indehiscent species from Western Australia. F. J. Obbens.237 Elevation of Eucalyptus ravensthorpensis, and notes on relationships between obligate-seeder and resprouter members of subseries Levispermae (Myrtaceae). C.R. Gosper, T. Hopley, M. Byrne, S.D. Hopper, S.M. Prober and C.J. Yates.247 Nomenclatural updates and a new species of annual Hydrocotyle (Araliaceae) from Western Australia. A.J. Perkins.253 Update on generic and specific nomenclature in Paracaleana (Drakaeinae), Caladeniinae and a new name in Caladenia (Orchidaceae). S.D. Hopper and A.R Brown.279 Paracaleana ferricola (Orchidaceae), a rare new species from south-west Western Australia. A.R Brown and G. Brockman.287 The Hibbertia polystachya-H. spicata (Dilleniaceae) species group in Western Australia. K.R. Thiele.291 A new name, clarification of synonymy, and a new subspecies for Isopogon (Rroteaceae) in Western Australia. B.L. Rye and T.D. Macfarlane.309 Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae), with validation of six species names and two new combinations. N.G. Marchant.317 Hibbertia tuberculata (Dilleniaceae), a new, rare species from Western Australia K.R.Thiele.335 Updates to Western Australia’s vascular plant census for 2019. C.M. Barker and J.M. Rercy-Bower.339 Other content Referees for Volume 30.353 Conservation Codes for Western Australian Flora and Fauna.354 Nuytsia The journal of the Western Australian Herbarium 30: 1-18 Published online 28 March 2019 SHORT COMMUNICATION Updates to Western Australia’s vascular plant census for 2018 The census database at the Western Australian Herbarium (PERTH), which provides the nomenclature for the website FloraBase (Western Australian Herbarium 1998-), lists current names and recent synonymy for Western Australia’s native and naturalised vascular plants, as well as algae, bryophytes, lichens, slime moulds and some fungi. The names represented in the census are either sourced from published research or denote as yet unpublished names based on herbarium voucher specimens. We herein summarise the changes made to vascular plant names in this database during 2018. One hundred and twenty-nine taxa were newly recorded for the State, of which 24 are naturalised and 41 have been added to the Threatened and Priority Flora list for Western Australia (Smith & Jones 2018; Western Australian Herbarium 1998-) (Table 1). A total of 185 name changes were made, including the formal publication of 29 phrase-named taxa (Table 2). Plant groups for which a number of name changes were made include Hydrocotyle L. (Perkins 2018a, 2018b), Drosera L. (Lowrie 2013a, 2013b, 2014), Lepilaena Harv. (Ito et al. 2016; Macfarlane et al. 2017) and Zygophyllum L. (transferred to Roepera A. Juss. following Beier et al. 2003). Numerous phrased-named taxa in the genus Baeckea L. were formally published under an expanded circumscription of Hysterobaeckea (Nied.) Rye (Rye 2018). Table 2 also includes cases where there has been a change of taxonomic concept, misapplication, exclusion or rank change. Table 1. New records added to Western Australia’s vascular plant census during 2018. in litt. = in correspondence; in sched. = on herbarium sheet/label; * = naturalised; T, P1-P4 = Conservation Codes for Western Australian flora (Smith & Jones 2018; Western Australian Herbarium 1998-)- New Name Status Comments Acacia prominens G.Don * New naturalised record for WA. G.J. Keighery in litt. (11/01/2018). Acacia sp. Binningup (G. Cockerton et al. WB 37784) PI K.R. Thiele in litt. (02/07/2018). Acacia sp. Southern Cross (G. Cockerton et al. WB 38518) PI G. Cockerton in litt. (14/05/2018). Aglaia brownii Pannell PI New record for WA. M. Hislop pers. comm. (23/10/2018). Angianthus sp. Salmon Gums (G.F. Craig 3074) PI M. Hislop in litt. (23/05/2018). Arthropodium sp. Ironstone (J. Bull & J. Waters ONS PJ 36.01) P2 S. Dillon in litt. (08/04/2018). Beyeria sp. Lake King (PR. Jefferies 680514) P2 M. Hislop in litt. (27/03/2018). Bolboschoenusfluviatilis (Torr.) Sojak PI New record for WA. M. Hislop in litt. (26/04/2018). Bonnaya tenuifolia (Colsm.) Sprang. New naturalised record for WA. W.R. Barker in sched. (06/04/2018). © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 2 Nuytsia Vol. 30 (2019) New Name Status Comments Bossiaea moylei Keighery P2 See Keighery (2018). Byblis pilbarana Lowrie & Conran See Lowrie (2014). Caladenia lateritica K.W.Dixon & Christenh. See Dixon & Christenhusz (2018b). Caladenia rosea K.W.Dixon & Christenh. See Dixon & Christenhusz (2018b). Calandrinia sp. Edel Land (F. Obbens FO 01/17) P2 F. Obbens in litt. (26/02/2018). Calandrinia sp. Gypsum (F. Obbens & L. Hancock FO 10/14) F. Obbens in litt. (21/02/2018). Callistemon citrinus (Curtis) Skeels New naturalised record for WA. G.J. Keighery in litt. (10/10/2018). Callitris endlicheri (Pari.) F.M.Bailey * New naturalised record for WA. G.J. Keighery in litt. (10/10/2018). Comesperma sabulosum A. J.Ford & Halford P3 See VoxAetal. (2017). Comesperma secundum subsp. oligotrichum A. J.Ford & Halford See Ford et al. (2017). Cucumis myriocarpus Naudin subsp. myriocarpus * New naturalised record for WA. See Shaiketa/. (2016). Cuscuta suaveolens Ser. New naturalised record for WA. G.J. Keighery in litt. (04/10/2017). Desmanthus virgatus (L.) Willd. * New naturalised record for WA. G.J. Keighery in litt. (12/03/2018). Diuris sp. South Coast (G. Brockman GBB 3041) G.B. Brockman & C.J. French in litt. (09/06/2018). Drosera albonotata A.SKdb., A.T.Cross, Meisterl & A.Fleischm. P2 See Robinson et al. (2018). Drosera aquatica Lowrie See Lowrie (2014). Drosera aurantiaca Lowrie See Lowrie (2014). Drosera x badgingarra Lowrie & Conran See Lowrie (2014). Drosera barrettorum Lowrie See Lowrie (2014). Drosera bindoon Lowrie See Lowrie (2014). Drosera calycina Planch. New record for WA. See Lowrie (2014). Drosera x carbarup Lowrie & Conran See Lowrie (2014). Drosera coalara Lowrie & Conran See Lowrie (2014). Drosera coomallo Lowrie & Conran See Lowrie (2014). Drosera cucullata Lowrie See Lowrie (2014). J.M. Percy-Bower & C.M. Parker, Updates to Western Australia’s vascular plant census for 2018 3 New Name Status Comments Drosera depauperata Lowrie & Conran See Eowrie (2014). Drosera esperensis Lowrie See Eowrie (2014). Drosera fragrans Lowrie See Eowrie (2014). Drosera glabriscapa Lowrie See Eowrie (2014). Drosera hirsuta Lowrie & Conran See Eowrie (2014). Drosera huegelii Endl. var. huegelii New record for WA. See Utz & Gibson (2017). Drosera huegelii var. phillmanniana Y.-A.Utz & R.P.Gibson P2 See Utz & Gibson (2017). Drosera indumenta Lowrie & Conran See Eowrie (2014). Drosera x legrandii Lowrie & Conran See Eowrie (2014). Drosera micra Lowrie & Conran See Eowrie (2014). Drosera micrantha Lehm. New record for WA. See Eowrie (2014). Drosera minutiflora Planch. New record for WA. See Eowrie (2014). Drosera x pingellyensis Lowrie & Conran See Eowrie (2014). Drosera planchonii Planch. New record for WA. See Eowrie (2014). Drosera verrucata Lowrie & Conran See Eowrie (2014). Eremophila compacta subsp. Kennedy Range (B. Buirchell BB 107) P2 B. Buirchell in litt. (22/11/2017). Eremophilaplatycalyx subsp. Granites (D.J. Edinger & G. Marsh DJE 4782) B. Buirchell in litt. (27/11/2017). Eremophila platycalyx subsp. Ilgarari (B. Buirchell BB 326) B. Buirchell in litt. (22/11/2017). Eremophila platycalyx subsp. Eeonora (J. Morrisey 252) B. Buirchell in litt. (22/11/2017). Eremophila platycalyx subsp. Neds Creek (N.H. Speck 1228) B. Buirchell in litt. (27/11/2017). Eremophila platycalyx subsp. Wongawol (N.H. Speck 1291) PI B. Buirchell in litt. (22/11/2017). Eremophila platycalyx subsp. Woolgorong (F. Hort et al. FH 3253) B. Buirchell in litt. (22/11/2017). Eremophila platycalyx subsp. Yalgoo (A. Markey & S. Dillon 3337) P3 B. Buirchell in litt. (22/11/2017). Eremophila sp. Coodardo Gap (B. Buirchell BB 330) PI B. Buirchell in litt. (22/11/2017). Eremophila sp. Mt Channar Range (C. Keating & M.E. Trudgen CK 408) PI B. Buirchell in litt. (08/05/2018). 4 Nuytsia Vol. 30 (2019) New Name Status Comments Eremophila sp. Pingandy dentate (B. Buirchell BB 331) PI B. Buirchell in litt. (22/11/2017). Eremophila sp. Thundelarra (B. Buirchell BB 324) PI B. Buirchell in litt. (22/11/2017). Eremophila sp. Tropicana (B. Buirchell BB 323) PI B. Buirchell in litt. (01/11/2017). Eremophila sp. Wubin North (R. Wait 8305) PI B. Buirchell in litt. (22/11/2017). Erythrina crista-galli L. New naturalised record for WA. G.J. Keighery in litt. (15/02/2018). Eucalyptus revelata D.Nicolle & R.L.Barrett P2 See Nicolle & Barrett (2018). Grevillea sp. Victoria Desert (R. Davis et al. RD 11611) PI R. Davis in litt. (23/10/2018). Halodule tridentata (Steinh.) Unger New record for WA. See Kuo (2011). Hibbertia pubens K.R.Thiele See Thiele (2018). Hysterobaeckea setifera Rye See Rye (2018). Isotropis sp. Arid zone (G. Byrne 2775) R. Davis & J.A. Wege in litt. (28/03/2018). Isotropis sp. Yalgoo (S. Patrick 2375) R. Davis & J.A. Wege in litt. (28/03/2018). Josephinia sp. Northern (T.E.H. Aplin 6360) W.R. Barker in litt. (11/11/2013). Leucopogon sp. Kirup (M. Hislop 3919) P2 M. Hislop in litt. (02/11/2018). Limnobium laevigatum (Willd.) Heine New naturalised record for WA. G.J. Keighery in litt. (11/01/2018). Lindernia cyanoplectra W.R.Barker See Barker (2018). Lindernia dunlopii W.R.Barker See Barker (2018). Lindernia porphyrodinea W.R.Barker & M.D.Barrett See Barker (2018). Lindernia sp. Minute-flowered (A.S. George 12433) P2 W.R. Barker in litt. (18/07/2018). Lindernia sp. Pilbara (M.N. Lyons & L. Lewis LV 1069) PI W.R. Barker & M. Lyons in litt. (18/07/2018). Melaleuca quinquenervia (Cav.) S.T.Blake * New naturalised record for WA. G.J. Keighery in litt. (10/10/2018). Microcorys sp. Mt Holland (D. Angus DA 2397) PI M. Hislop in litt. (01/12/2017). Nymphaeapubescens Willd. * New naturalised record for WA. M.D. Barrett & E. Dalziell in litt. (04/12/2017). Olearia sp. Gregory (M. Hislop 3784) PI M. Hislop in litt. (11/07/2018). Paracaleana sp. Laterite (G. Brockman GBB 3571) P2 G. Brockman & A. Brown in litt. (18/04/2018). J.M. Percy-Bower & C.M. Parker, Updates to Western Australia’s vascular plant census for 2018 5 New Name Status Comments Pelargonium x hortorum L.H.Bailey * New naturalised record for WA. G.J. Keighery in litt. (10/10/2018). Pentameris patula (Nees) Steud. New naturalised record for WA. T.D. Macfarlane in litt. (22/08/2018). Petrophile brevifolia Lindl. subsp. brevifolia New record for WA. See Keighery & Keighery (2017). Petrophile brevifolia subsp. rosea B. J.Keighery & Keighery See Keighery & Keighery (2017). Polymeria mollis (Benth.) Domin New record for WA. D A. Halford in litt. (17/10/2018). Pterostylis arbuscula (D.L. Jones & C. J.French) D.L.Jones & C.J.French See Jones & French (2018). Pterostylis atrosanguinea (D.L. Jones & C. J.French) D.L.Jones & C.J.French See Jones & French (2018). Pterostylis ectypha (D.L.Jones & C. J.French) D.L.Jones & C. J.French See Jones & French (2018). Pterostylis faceta (D.L. Jones, C. J.French & M. A.Clem.) D.L. Jones & C.J.French See Jones & French (2018). Pterostylis galgula (D.L.Jones & C.J.French) D.L.Jones & C.J.French See Jones & French (2018). Pterostylis longicornis (D.L.Jones & C.J.French) D.L.Jones & C.J.French See Jones & French (2018). Pterostylis orbiculata (D.L.Jones & C.J.French) D.L.Jones & C.J.French See Jones & French (2018). Pterostylisprecatoria (D.L.Jones, C.J.French & M.A.Clem.) D.L.Jones & C.J.French See Jones & French (2018). Pterostylis saxosa (D.L.Jones & C.J.French) D.L.Jones & C.J.French See Jones & French (2018). Pterostylis serotina (D.L.Jones, C.J.French & M.A.Clem.) D.L.Jones & C.J.French See Jones & French (2018). Pterostylis sigmoidea (D.L.Jones & C.J.French) D.L.Jones & C.J.French See Jones & French (2018). Ptilotusyapukaratja R.W.Davis & T.Hammer PI See Davis & Hammer (2018). Quercus palustris Mlinchh. * New naturalised record for WA. See Lohr& Keighery (2016). Quercus robur L. * New naturalised record for WA. See Lohr& Keighery (2016). Ranunculus repens L. New naturalised record for WA. G.J. Keighery in litt. (10/10/2018). Rhizanthella johnstonii K.W.Dixon & Christenh. T See Dixon & Christenhusz (2018a). Salvia misella Kunth * New naturalised record for WA. M. Hislop pers. comm. (23/10/2018). Scaevola sp. Mt Ragged (M. Goods GG 004) P2 B. Maslin & R. Davis in litt. (23/01/2012). Scholtzia sp. Coburn (N. Murdock NM 031) P2 B.L. Rye in litt. (03/04/2018). Scholtzia sp. Overlander (M.E. Trudgen 12138) PI B.L. Rye in litt. (08/05/2018). Nuytsia Vol. 30 (2019) New Name Status Comments Sida fallax Walp. * New naturalised record for WA. G.J. Keighery in litt. (10/10/2018). Solarium nitidibaccatum Bitter New naturalised record for WA. See Sarkinen etal. (2018). Solarium sp. Red Hill (S. van Leeuwen et al. PBS 5415) P3 A.A. Mitchell in litt. (01/05/2018). Stemodia sp. Carnarvon (W.R. Barker 2154) W.R. Barker in litt. (06/04/2018). Stemodia sp. Doongan (R.L. Barrett RLB 4704) W.R. Barker in litt. (26/09/2013). Stemodia sp. Shay Gap (B. Cook 7) W.R. Barker in litt. (26/09/2013). Styphelia sp. Stirling Range (R.D. Royce 6087) M. Hislop in litt. (30/11/2017). Styphelia sp. Tarin Rock (W.E. Blackall 1315) PI M. Hislop in litt. (12/07/2018). Thinopyrum ponticum (Podp.) Barkworth & D.R.Dewey New naturalised record for WA. G.J. Keighery in litt. (28/01/2016). Thymophylla tenuiloba (DC.) Small New naturalised record for WA. V. Stajsic in litt. (19/01/2018). Thymophylla tenuiloba (DC.) Small var. tenuiloba * New naturalised record for WA. V. Stajsic in litt. (19/01/2018). Ursinia nana DC. New naturalised record for WA. See Keighery & Keighery (2018). Ursinia nana DC. subsp. nana New naturalised record for WA. G.J. Keighery in litt. (21/07/2018). Utricularia albertiana R.W. Jobson & Baleeiro PI See Jobson et al. (2018). Utricularia bidentata R.W.Jobson & Baleeiro P3 See Jobson et al. (2018). Utricularia foveolata Edgew. New record for WA. See Eowrie (2014). Utricularia magna R.W.Jobson & M.D.Barrett P2 See Jobson et al. (2018). Utricularia nivea Vahl New record for WA. M.D. Barrett inlitt. (12/12/2017). Utriculariapapilliscapa R.W.Jobson & M.D.Barrett P2 See Jobson et al. (2018). Verticordia sp. Koolyanobbing (B.H. Smith 1457) PI B.E. Rye in litt. (03/04/2018). J.M. Percy-Bower & C.M. Parker, Updates to Western Australia’s vascular plant census for 2018 7 Table 2. Changes to existing entries in Western Australia’s vascular plant census during 2018. Excluded taxon = a name used in the botanical literature that refers to a taxon never occurring in WA; misapplied name = a name used in the botanical literature but now considered to refer to one or more different WA taxa; nomenclatural synonym = a superseded name based on the same type specimen as the accepted name—^the epithet is usually transferred to a different genus name or rank; taxonomic synonym = a superseded name based on a different type specimen to the accepted name; orthographic variant = mis-spelling of a name in original publication; in litt. = in correspondence; in sched. = on herbarium sheet/label. Status: * = naturalised; T,P1-P4 = Conservation Codes forWesternAustralian flora (Smith & Jones 2018; Western Australian Herbarium 1998-). Old Name New Name Status Comments Aristida personata Henrard n/a Excluded taxon. S.J. Dillon in sched (30/10/2018). Baeckea sp. Barbalin (B.L. Rye & M.E. Trudgen BLR 241022) Hysterobaeckea petraea Rye Name synonymised. See Rye (2018). Baeckea sp. Bencubbin-Koorda (M.E. Trudgen 5421) Hysterobaeckea setifera subsp. meridionaJis Rye Name synonymised. See Rye (2018). Baeckea sp. Boorabbin (J.H. Willis s.n. 4/10/1961) Hysterobaeckea ochropetala subsp. reliqua Rye Name synonymised. See Rye (2018). Baeckea sp. Bulla Bulling (D. J.E. Whibley 4648) Hysterobaeckea ochropetala subsp. reliqua Rye Name synonymised. See Rye (2018). Baeckea sp. Bungalbin Hill (B.J. Lepschi & L.A. Craven 4586) Hysterobaeckea cornuta Rye P3 Name synonymised. See Rye (2018). Baeckea sp. Comet Vale (A.S. George 8078) Hysterobaeckea ochropetala subsp. cometes Rye P3 Taxon formally published. See Rye (2018). Baeckea sp. Coolgardie (A. Strid 21320) Hysterobaeckea ochropetala subsp. reliqua Rye Name synonymised. See Rye (2018). Baeckea sp. Die Hardy Range (E. Mattiske J91) Hysterobaeckea cornuta Rye P3 Name synonymised. See Rye (2018). Baeckea sp. Exclamation Lake (M.E. Trudgen 1524) Hysterobaeckea sp. Exclamation Lake (M.E. Trudgen 1524) PI Name synonymised. See Rye (2018). Baeckea sp. Fitzgerald Peaks (P.J. Poll 53) Hysterobaeckea graniticola Rye P2 Taxon formally published. See Rye (2018). Baeckea sp. Flying Fox Mine (A.O’Connor & V. Longman FF532) Hysterobaeckea pterocera Rye PI Taxon formally published. See Rye (2018). Baeckea sp. Gnarlbine Rocks (G. Barrett GRH469) Hysterobaeckea ochropetala subsp. reliqua Rye Name synonymised. See Rye (2018). Baeckea sp. Kalgarin Hill Road (A M. Lyne, L. Craven & F.ZichAML1018) Hysterobaeckea glandulosa Rye PI Taxon formally published. See Rye (2018). Baeckea sp. Lake Brown (E. Merrall s.n. 1889) Hysterobaeckea ochropetala (F.Muell.) Rye subsp. ochropetala PI Name synonymised. See Rye (2018). Baeckea sp. Lake Campion (A. Coates AC 2285) Hysterobaeckea setifera subsp. meridionalis Rye Name synonymised. See Rye (2018). Baeckea sp. Melita Station (H. Pringle 2738) Hysterobaeckea occlusa Rye Name synonymised. See Rye (2018). Baeckea sp. Mt Clara (R.J. Cranfield 11693) Hysterobaeckea ochropetala subsp. reliqua Rye Name synonymised. See Rye (2018). Baeckea sp. Mt Clifford (B. Seveme 74002) Hysterobaeckea occlusa Rye Name synonymised. See Rye (2018). Nuytsia Vol. 30 (2019) Old Name New Name Status Comments Baeckea sp. Mt Jackson (G. J. Keighery 4362) Hysterobaeckea cornuta Rye P3 Name synonymised. See Rye (2018). Baeckea sp. Pigeon Rocks (D. Grace DJP281) Hysterobaeckea petraea Rye Name synonymised. See Rye (2018). Baeckea sp. Queen Victoria Rock (K.R. Newbey6103) Hysterobaeckea ochropetala subsp. reliqua Rye Name synonymised. See Rye (2018). Baeckea sp. Roundtop Hill (P. Armstrong 05/843) Hysterobaeckea ochropetala subsp. reliqua Rye Name synonymised. See Rye (2018). Baeckea sp. Ubini (R. Pullen 9610) Hysterobaeckea ochropetala subsp. reliqua Rye Name synonymised. See Rye (2018). Baeckea sp. Wanarra (M.E. Trudgen MET 5376) Hysterobaeckea setifera Rye subsp. setifera Name synonymised. See Rye (2018). Baeckea sp. Wubin (M.E. Trudgen 5404) Hysterobaeckea longipes Rye Taxon formally published. See Rye (2018). Bartsia trixago E. Bellardia trixago (F.) All. Nomenclatural synonym. See Uribe-Convers & Tank (2016). Borya jabirabela Churchill nJdL Excluded taxon. M.D. Barrett in litt. (04/09/2018). Borya sp. Synnot Range (M.D. Barrett MDB 444) Borya stenophylla M.D.Barrett PI Taxon formally published. See Barrett (2018). Bossiaea sp. Bremer (K. Newbey 2980) Bossiaea lalagoides F.Muell. P3 Name synonymised. See Keighery (2018). Bossiaea sp. Cairn Hill (M. Henson MJH 2-28) Bossiaea moylei Keighery P2 Taxon formally published. See Keighery (2018). Bossiaea sp. Waroona (B. J. Keighery & N. Gibson 229) Bossiaea angustifolia (Meisn.) Keighery Name synonymised. See Keighery (2018). Caesalpinia gilliesii (Hook.) D.Dietr. Erythrostemon gilliesii (Hook.) Klotzsch Nomenclatural synonym. See Gagnon et al. (2016). Caladenia lateritica K.W.Dixon & Christenh. Caladeniaflava subsp. sylvestris Hopper &A.P.Br. Taxonomic synonym. A.P Brown in sched. (23/05/2018). Caladenia rosea K.W.Dixon & Christenh. Caladenia x spectabilis Hopper & A.P.Br. Taxonomic synonym. A.P. Brown in sched. (23/05/2018). Calandrinia sp. Goongarrie (F. Obbens, F. Hort& J. Hort FO 18/13) Calandrinia quartzitica Obbens PI Taxon formally published. See Obbens (2018). Calandrinia sp. Meekering (F. Obbens 42/02) Calandrinia wilsonii Obbens P2 Taxon formally published. See Obbens (2018). Calandrinia sp. Widgiemooltha (F. Obbens & E. Reid FO 9/05) Calandrinia lefroyensis Obbens PI Taxon formally published. See Obbens (2018). Callitris huegelii J.W. Green Callitris glaucophylla Joy Thomps. & F.A.S.Johnson Misapplied name. A name of uncertain application. See Thompson & Johnson (1986). Cayratia trifolia (F.) Domin Causonis trifolia (F.) Mabb. & J.Wen Nomenclatural synonym. See Mabberley (2017). Celtis philippensis Blanco Celtis strychnoides Planch. Misapplied name. See Guymer (2013). Chionachne cyathopoda (F.Muell.) Benth. Polytoca cyathopoda (F.Muell.) F.M. Bailey Nomenclatural synonym. See Mabberley (2017). J.M. Percy-Bower & C.M. Parker, Updates to Western Australia’s vascular plant census for 2018 9 Old Name New Name Status Comments Chionachne hubbardiana Henrard Citrullus lanatus (Thunb.) Matsum. & Nakai Comesperma pallidum Pedley Commersonia cygnorum Steud. Corunastylis tepperi (Tapper) D.L Jones & M. A.Clem. Cynanchum carnosum (R.Br.) Schltr. Davenportia davenportii (F.Muell.) R.W. Johnson Drosera brevicornis Lowrie Drosera bulbosa subsp. major (Diels) N.G.Marchant & Lowrie Drosera erythrorhiza subsp. collina N.G.Marchant & Lowrie Drosera erythrorhiza Lindl. subsp. erythrorhiza Drosera erythrorhiza subsp. magna N.G.Marchant & Lowrie Drosera erythrorhiza subsp. squamosa (Benth.) N.G.Marchant & Lowrie Drosera gigantea subsp. geniculata N.G.Marchant & Lowrie Drosera gigantea Lindl. subsp. gigantea Drosera lanata K.Kondo Drosera macrantha subsp. eremaea N.G.Marchant & Lowrie Drosera macrantha Endl. subsp. macrantha Drosera macrophylla Lindl. subsp. macrophylla Drosera macrophylla subsp. monantha Lowrie & Carlquist Drosera marchantii DeBuhr subsp. marchantii Drosera marchantii subsp. prophylla N.G.Marchant & Lowrie Polytoca hubbardiana (Henrard) Mabb. Citrullus amarus Schrad. Comesperma sabulosum A. J.Ford & Halford Commersonia corniculata (Sm.) K.A.Sheph. & C.F.Wilkins Corunastylis fuscoviridis (Reader) D.L.Jones & M.A.Clem. Vincetoxicum carnosum (R.Br.) Benth. Distimake davenportii (F.Muell.) A.R.Simoes & Staples n/a Drosera major (Diels) Lowrie Drosera collina (N.G.Marchant & Lowrie) Lowrie Drosera erythrorhiza Lindl. Drosera magna (N.G.Marchant & Lowrie) Lowrie Drosera squamosa Benth. Drosera geniculata N.G.Marchant & Lowrie) Lowrie Drosera gigantea Lindl. nJdL Drosera eremaea (N.G.Marchant & Lowrie) Lowrie & Conran Drosera macrantha Endl. Drosera macrophylla Eindl. Drosera monantha (Eowrie & Carlquist) Eowrie Drosera marchantii DeBuhr Drosera prophylla (N.G.Marchant P3 & Eowrie) Eowrie Nomenclatural synonym. See Mabberley (2017). * Misapplied name. See Chomicki & Renner (2015). P3 Misapplied name. See Voxdetal. (2017). Taxonomic synonym. See Shepherd & Wilkins (2018a). Misapplied name. A.P Brown insched (18/04/2018). Nomenclatural synonym. See Eiede-Schumann & Meve (2018). Nomenclatural synonym. See Simoes & Staples (2017). Excluded taxon. See Eowrie (2013a). Nomenclatural synonym. See Eowrie (2014). Nomenclatural synonym. See Eowrie (2014). Nomenclatural synonym. No subspecies recognised. See Eowrie (2013a). Nomenclatural synonym. See Eowrie (2014). Nomenclatural synonym. See Eowrie (2014). Nomenclatural synonym. See Eowrie (2014). Nomenclatural synonym. No subspecies recognised. See Eowrie (2013b). Excluded taxon. See Eowrie (2013b). PI Nomenclatural synonym. See Eowrie (2014). Nomenclatural synonym. No subspecies recognised. See Eowrie (2013b). Nomenclatural synonym. No subspecies recognised. See Eowrie (2013b). Nomenclatural synonym. See Eowrie (2014). Nomenclatural synonym. No subspecies recognised. See Eowrie (2013b). Nomenclatural synonym. See Eowrie (2014). 10 Nuytsia Vol. 30 (2019) Old Name New Name Status Comments Drosera menziesii subsp. basifolia N.G.Marchant & Lowrie Drosera basifolia (N.G.Marchant & Lowrie) Lowrie Nomenclatural synonym. See Lowrie (2014). Drosera menziesii DC. subsp. menziesii Drosera menziesii DC. Nomenclatural synonym. No subspecies recognised. See Lowrie (2013b). Drosera menziesii subsp. penicillaris (Benth.) N.G.Marchant & Lowrie Drosera drummondii Planch. Taxonomic synonym. See Lowrie (2014). Drosera menziesii subsp. thysanosepala (Diels) N.G.Marchant Drosera thysanosepala Diels Nomenclatural synonym. See Lowrie (2014). Drosera neesii subsp. borealis N.G.Marchant Drosera neesii Lehm. Taxonomic synonym. No subspecies recognised. See Lowrie (2014). Drosera neesii Lehm. subsp. neesii Drosera neesii Lehm. Nomenclatural synonym. No subspecies recognised. See Lowrie (2014). Drosera occidentalis subsp. australis N.G.Marchant & Lowrie Drosera australis (N.G.Marchant & Lowrie) Lowrie & Conran Nomenclatural synonym. See Lowrie (2014). Drosera occidentalis Morrison subsp. occidentalis Drosera occidentalis Morrison P4 Nomenclatural synonym. No subspecies recognised. See Lowrie (2013b). Droserapaleacea DC. subsp. paleacea Drosera paleacea DC. PI Nomenclatural synonym. No subspecies recognised. See Lowrie (2013b). Drosera paleacea subsp. trichocaulis (Diels) N.G.Marchant & Lowrie Drosera trichocaulis (Diels) Lowrie & Conran Nomenclatural synonym. See Lowrie (2014). Drosera parvula Planch. Drosera paleacea DC. PI Taxonomic synonym. See Lowrie (2014). Droserapetiolaris DC. nJdi Excluded taxon. See Eowrie (2013b). Elythranthera x intermedia (Fitzg.) M.A.Clem. Caladenia x intermedia (Fitzg.) M.A.Clem. & D.L.Jones PI Nomenclatural synonym. See Jones & Clements (2017). Eremophila glabra subsp. green flowers (E.A. Griffin 5347) Eremophila glabra subsp. chlorella (Gand.) Chinnock T Name synonymised. A.P Brown in sched. (02/08/2013). Eremophila sp. Narrow leaves (J.D. Start D12-150) Eremophila subangustifolia A.P.Br. & T.M.Llorens T Taxon formally published. See Brown et al. (2018). Eucalyptus brachycorys Blakely Eucalyptus comitae-vallis Maiden Taxonomic synonym. See French (2012). Eucalyptus capillosa Brooker & Hopper subsp. capillosa Eucalyptus capillosa Brooker & Hopper Nomenclatural synonym. No subspecies recognised. See French (2012). Eucalyptus capillosa subsp. polyclada Brooker & Hopper Eucalyptus capillosa Brooker & Hopper Taxonomic synonym. See French (2012). Eucalyptus sp. Wagerup (L. Johnson 9127 & B. Briggs) Eucalyptus x graniticola Hopper P4 Taxon formally published. See Hopper (2018). Fioria vitifolia (L.) Mattei Hibiscus vitifolius L. Nomenclatural synonym. See Mabberley (2008). Flemingia involucrata Benth. Flemingia trifoliata (Jungh.) C.Y.Wu Nomenclatural synonym. See van der Maesen (2012). J.M. Percy-Bower & C.M. Parker, Updates to Western Australia’s vascular plant census for 2018 11 Old Name New Name Status Comments Glochidion perakense Hook.f Glochidion sumatranum Miq. Misapplied name. M.D. Barrett in litt (14/12/2017). Glochidion perakense var. supra-axillare (Benth.) Airy Shaw Glochidion sumatranum Miq. Misapplied name. M.D. Barrett in litt. (14/12/2017). Hemiarrhena plantaginea (F.Muell.) Benth. Linderniaplantaginea (F.Muell.) F.Muell. Nomenclatural synonym. See Barker (2018). Hibbertia sp. Geraldton Sandplains (R. Edmiston E 421) Hibbertia squarrosa K.R.Thiele Taxon formally published. See Thiele (2018). Hydrocotyle sp. Coorowensis (PG. Wilson 12580) Hydrocotyle spinulifera A.J.Perkins P3 Taxon formally published. See Perkins (2018a). Hydrocotyle sp. Coraginaensis (K.R. Newbey 7477) Hydrocotyle perforata A.H?Qxkms, P2 Taxon formally published. See Perkins (2018b). Hydrocotyle sp. Hexaptera (T. Erickson TEE 173) Hydrocotyle decorata A.H?Qxkms, P2 Taxon formally published. See Perkins (2018b). Hydrocotyle sp. Truslove (M.A. Burgman 4419) Hydrocotyle asterocarpa A.J.Perkins P2 Taxon formally published. See Perkins (2018b). Hydrocotyle sp. Warriedar (PG. Wilson 12267) Hydrocotyle dimorphocarpa A.J.Perkins PI Taxon formally published. See Perkins (2018a). Hypoxis sp. Prince Regent River (M.D. Barrett 924 B) Hypoxis cavernicola M.D.Barrett P2 Taxon formally published. See Barrett (2018). Jacquemontia sp. Keep River (J.E. Egan 5051) Jacquemontia sp. Keep River (J.E. Egan 5015) PI Name synonymised. See CHAH (2006). Josephinia sp. Marandoo (M.E. Trudgen 1554) Josephinia eugeniae F.Muell. Name synonymised. W.R. Barker in sched. (06/04/2018). Lasiopetalum sp. Badgingarra (E.A. Griffin 5278) Lasiopetalum decoratum K.A.Sheph. & C.F.Wilkins P2 Taxon formally published. See Shepherd & Wilkins (2018b). Lasiopetalum sp. Hill River (T.N. Stoate 5) Lasiopetalum rupicola K.A.Sheph. & C.F.Wilkins PI Taxon formally published. See Shepherd & Wilkins (2018b). Lasiopetalum sp. Kukerin (C.A. Gardner 13646) Lasiopetalum rosmarinifolium (Turcz.) Benth. Name synonymised. See Shepherd & Wilkins (2018c). Lepilaena australis Harv. Althenia australis (Harv.) F.Muell. Nomenclatural synonym. See Ito etal. (2016). Lepilaena bilocularis Kirk Althenia bilocularis (Kirk) Cockayne Nomenclatural synonym. See Ito etal. (2016). Lepilaena cylindrocarpa (MlilLBerol.) Benth. Althenia cylindrocarpa (Mull.Berol.) Asch. Nomenclatural synonym. See Ito etal. (2016). Lepilaena marina E.E.Robertson Althenia marina (E.E.Robertson) Yulto Nomenclatural synonym. See Ito etal. (2016). Lepilaenapatentifolia E.E.Robertson Althenia patentifolia (E.E.Robertson) T.Macfarlane & D.D.Sokoloff Nomenclatural synonym. See Macfarlane et al. (2017). Lepilaenapreissii (Eehm.) F.Muell. Althenia preissii (Eehm.) F.Muell. Nomenclatural synonym. See Ito etal. (2016). Lepilaena sp. Wheatbelt (M.T. Collins et al. 265) Althenia hearnii T.Macfarlane & D.D.Sokoloff Taxon formally published. See Macfarlane et al. (2017). Leucopogon brevicuspis Benth. Styphelia sp. Cascades (R. Davis 11037) Misapplied name. M. Hislop in litt. (29/03/2018). 12 Nuytsia Vol. 30 (2019) Old Name New Name Status Comments Leucopogon crassifolius Sond. Styphelia sp. South Coast (J.M. Powell 3374) Misapplied name. M. Hislop in litt. (29/03/2018). Leucopogon ovalifolius Sond. Styphelia sp. Albany (M. Hislop 2218) Misapplied name. M. Hislop in litt. (29/03/2018). Lindernia Crustacea (L.) F.Muell. Torenia Crustacea (L.) Cham. & Schltdl. Nomenclatural synonym. W.R. Barker in sched. (06/04/2018). Lindernia subulata R.Br. nJdL Excluded taxon. M.D. Barrett in litt. (03/09/2018). Lipocarpha chinensis (Osbeck) J.Kern Cyperus albescens (Steud.) Larridon & Govaerts P3 Taxonomic synonym. See Larridon cr a/. (2016). Meiogyne cylindrocarpa (Burck) Heusden subsp. cylindrocarpa Meiogyme cylindrocarpa (Burck) Heusden Nomenclatural synonym. No subspecies recognised. See Xue etal (2014). Merremia aegyptia (L.) Urb. Distimake aegyptius (L.) A.R.Simoes & Staples * Nomenclatural synonym. See Simoes & Staples (2017). Merremia dissecta (Jacq.) Hallier f. Distimake dissectus (Jacq.) A.R.Simoes & Staples Nomenclatural synonym. See Simoes & Staples (2017). Merremia dissecta (Jacq.) Hallier f. var. dissecta Distimake dissectus (Jacq.) A.R.Simoes & Staples var. dissectus * Nomenclatural synonym. See Petrongari etal. (2018). Merremia kimberleyensis R.W. Johnson Distimake kimberleyensis (R.W.Johnson) A.R.Simoes & Staples Nomenclatural synonym. See Simoes & Staples (2017). Merremia quinata (R.Br.) Ooststr. Distimake quinatus (R.Br.) A.R.Simoes & Staples Nomenclatural synonym. See Simoes & Staples (2017). Merremia umbellata (L.) Hallier f. nJdi Excluded taxon. See Simoes & Staples (2017). Merremia umbellata subsp. orientalis (Hallier f.) Ooststr. Camonea pilosa (Houtt.) A.RSimoes & Staples PI Misapplied name. See Simoes & Staples (2017). Mnesithea granularis (L.) de Koning & Sosef Hackelochloa granularis (L.) Kuntze Nomenclatural synonym. See Arthan et al. (2016). Monochoria cyanea (F.Muell.) F.Muell. Pontederia cyanea (F.Muell.) M.Pell. & C.N.Horn Nomenclatural synonym. See Pellegrini etal. (2018). Monochoria vaginalis (Burm.f.) Kunth Pontederia vaginalis Burm.f. Nomenclatural synonym. See Pellegrini etal. (2018). Monodia stipoides S.W.L. Jacobs Triodia stipoides (S.W.L.Jacobs) Crisp & Mant Nomenclatural synonym. See Crisp etal. (2015). Opercularia rubioides Juss. Opercularia nubicola A.S.Markey P2 Misapplied name. See Markey (2018). Opercularia sp. Stirling Range (M. Hislop 2839) Opercularia nubicola A.S.Markey P2 Taxon formally published. See Markey (2018). Ozothamnus sp. Filifolius (A.S. George 8292) Ozothamnus filifolius Puttock Name synonymised. See Schmidt-Lebuhn ct fl/. (2018). Ozothamnus sp. Northampton (J. Brooker &M. Weir 913) Ozothamnus vespertinus R.W.Davis, Wege & Schmidt-Leb. PI Taxon formally published. See Schmidt-Lebuhn ct a/. (2018). Paracaleana sp. Israelite Bay (W. Jackson 8/11/1995) Paracaleana parvula Hopper & A.P.Br. P2 Name synonymised. See Hopper & Brown (2006). J.M. Percy-Bower & C.M. Parker, Updates to Western Australia’s vascular plant census for 2018 13 Old Name New Name Status Comments ParentuceUia viscosa (L.) Caruel Bellardia viscosa (L.) Fisch. & C.A.Mey. * Nomenclatural synonym. See Uribe-Convers & Tank (2016). Polymeria sp. A Kimberley Flora (T.E.H. Aplinetal. 418) Polymeria quadrivalvis R.Br. Name synonymised. D A. Halford in litt. (17/10/2018). Pomax sp. desert (A.S. George 11968) Pomax sp. Sand dunes (PG. Wilson 752) Name synonymised. G.J. Keighery in litt. (08/06/2015). Pontederia cordata L. nJdL Excluded taxon. G.J. Keighery in litt. (02/10/2018). Pterostylis sp. Cape Arid (G. Brockman GBB 263) Pterostylis meridionalis (D.L.Jones & C.J.French) D.L.Jones & C.J.French Name synonymised. G.B. Brockman & C.J. French insched. (18/04/2018). Pterostylis sp. cauline leaves (N. Gibson &M.N. Lyons 1490) Pterostylispyramidalis Lindl. Name synonymised. G.B. Brockman & C.J. French insched. (18/04/2018). Pterostylis sp. granite (W. Jackson BJ351) Pterostylis microphylla D.L.Jones & C.J.French Name synonymised. A.P Brown in sched. (23/05/2018). Pterostylis sp. Helena River (G. Brockman GBB 340) Pterostylis angulata (D.L.Jones & C.J.French) D.L.Jones & C.J.French Name synonymised. G.B. Brockman & C.J. French insched. (18/04/2018). Ptilotus gaudichaudii subsp. eremita (S.Moore) Lally Ptilotus eremita (S.Moore) T.Hammer & R.W.Davis Nomenclatural synonym. See Hammer et al. (2018a). Ptilotus gaudichaudii (Steud.) J.M.Black subsp. gaudichaudii Ptilotus gaudichaudii (Steud.) J.M.Black Nomenclatural synonym. No subspecies recognised. See Hammer et al. (2018 a). Ptilotus nobilis (Lindl.) F.Muell. subsp. nobilis Ptilotus nobilis (Lindl.) F.Muell. Nomenclatural synonym. No subspecies recognised. See Hammer et al. (2018b). Ptilotus petiolatus Farmar Ptilotus murrayi F.Muell. Taxonomic synonym. See Hammer (2018). Ptilotus sp. Doolgunna (D. Edinger 4419) Ptilotus actinocladus T.Hammer & R.W.Davis PI Taxon formally published. See Hammer & Davis (2018). Ptilotus sp. Edaggee Station (T.E.H. Aplin 3208) Ptilotus unguiculatus T.Hammer PI Taxon formally published. See Hammer (2018). Ptilotus sp. Goldfields (R. Davis 10796) Ptilotus exaltatus Nees Name synonymised. See Hammer et al. (2018b). Rhyncharrhena linearis (Decne.) K.L.Wilson Vincetoxicum lineare (Decne.) Meve & Liede Nomenclatural synonym. See Liede-Schumann & Meve (2018). Sansevieria trifasciata Prain Dracaena trifasciata (Prain) Mabb. Nomenclatural synonym. See Mabberley (2017). Schinus terebinthifolius Raddi Schinus terebinthifolia Raddi * Orthographic variant. See Zona (2015). Secamone timoriensis Decne. Secamone timorensis Decne. Orthographic variant. See Decaisne (1844). Sida sp. verrucose glands (F.H. Mollemans 2423) Sida sp. L (A.M. Ashby 4202) Name synonymised. R.M. Barker pers. comm. (2017). 14 Nuytsia Vol. 30 (2019) Old Name New Name Status Comments Sisyrinchium exile E.P.Bicknell Sisyrinchium rosulatum E.P.Bicknell * Taxonomic synonym. See USDA(2018). Spartothamnella teucriiflora (F.Muell.) Moldenke Teucrium teucriiflorum (F.Muell.) Kattari & Salmaki Nomenclatural synonym. See Salmaki et al. (2016). Tecticornia sp. Little Sandy Desert (K.A. Shepherd & C. Wilkins KS 830) Tecticornia willisii K.A.Sheph. PI Taxon formally published. See Shepherd (2018). Tephrosia sp. C Kimberley Flora (K.F. Kenneally 5599) Tephrosia pedleyi R. Butcher P3 Taxon formally published. See Butcher (2018). Thespesia thespesioides (Benth.) Fryxell Azanza thespesioides (Benth.) F.Areces Nomenclatural synonym. See Areces-Berazain & Ackerman (2016). Thomasia sp. Merredin (M.B. Mills 11) Thomasia sarotes Turcz. Name synonymised. K.A. Shepherd & C.F. Wilkins inlitt. (28/11/2018). Thomasia sp. Salmon Gums (C.A. Gardner s.n. PERTH 02708639) Thomasia sarotes Turcz. Name synonymised. K.A. Shepherd & C.F. Wilkins inlitt. (28/11/2018). Thomasia sp. Toolbrunup (G.J. Keighery 9895) Thomasia brachystachys Turcz. P2 Name synonymised. C. Wilkins in sched. (26/07/2016). Tylophora benthamii Tsiang Vincetoxicum polyanthum Kuntze PI Nomenclatural synonym. See Liede-Schumann & Meve (2018). Tylophora cinerascens (R.Br.) P.I.Forst. Vincetoxicum cinerascens (R.Br.) Meve & Liede Nomenclatural synonym. See Liede-Schumann & Meve (2018). Tylophora flexuosa R.Br. Vincetoxicum flexuosum (R.Br.) Kuntze Nomenclatural synonym. See Liede-Schumann & Meve (2018). Utricularia sp. Theda (M.D. Barrett MDB 2056) Utricularia hamata R.W. Jobson & M.D.Barrett PI Taxon formally published. See Jobson et al. (2018). Vaccaria hispanica (Mill.) Rauschert Gypsophila vaccaria (L.) Sm. * Taxonomic synonym. See Madhani et al. (2017). Wahlenbergia communis Carolin Wahlenbergia capillaris (G.Lodd.) G.Don Taxonomic synonym. See Turner (2016). Zygophyllum ammophilum F.Muell. Roepera ammophila (F.Muell.) Beier & Thulin Nomenclatural synonym. See Beier eta/. (2003). Zygophyllum angustifolium H.Eichler Roepera angustifolia (H.Eichler) Beier & Thulin Nomenclatural synonym. See Beier eta/. (2003). Zygophyllum apiculatum F.Muell. Roepera apiculata (F.Muell.) Beier & Thulin Nomenclatural synonym. See Beier eta/. (2003). Zygophyllum aurantiacum (Lindl.) F.Muell. Roepera aurantiaca Lindl. Nomenclatural synonym. See Beier eta/. (2003). Zygophyllum aurantiacum (Lindl.) F.Muell. subsp. aurantiacum Roepera aurantiaca Lindl. subsp. aurantiaca Nomenclatural synonym. See Beier eta/. (2003). Zygophyllum billardierei DC. Roepera billardierei (DC.) G.Don Nomenclatural synonym. See Beier eta/. (2003). Zygophyllum compressum J.M. Black Roepera compressa (J.M.Black) Beier & Thulin Nomenclatural synonym. See Beier eta/. (2003). Zygophyllum eichleri R.M.Barker Roepera eichleri (R.M.Barker) Beier & Thulin Nomenclatural synonym. See Beier et al. (2003). J.M. Percy-Bower & C.M. Parker, Updates to Western Australia’s vascular plant census for 2018 15 Old Name New Name Status Comments Zygophyllum eremaeum (Diels) Ostenf Roepera eremaea (Diels) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum fruticulosum DC. Roepera fruticulosa (DC.) G.Don Nomenclatural synonym. Beier et al. (2003). See Zygophyllum glaucum F.Muell. Roepera glauca (F.Muell.) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum halophilum R.M.Barker Roepera halophila (R.M.Barker) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum iodocarpum F.Muell. Roepera iodocarpa (F.Muell.) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum kochii Tate Roepera kochii (Tate) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum lobulatum (Benth.) R.M.Barker Roepera lobulata (Benth.) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum ovatum Ewart & Jean White Roepera ovata (Ewart & Jean White) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum reticulatum R.M.Barker Roepera reticulata (R.M.Barker) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum retivalve Domin Roepera retivalvis (Domin) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum simile H.Eichler Roepera similis (H.Eichler) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum tesquorum J.M.Black Roepera tesquorum (J.M.Black) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Zygophyllum tetrapterum R.M.Barker Roepera tetraptera (R.M.Barker) Beier & Thulin Nomenclatural synonym. Beier et al. (2003). See Acknowledgements Curation staff at PERTH and the Department of Biodiversity, Conservation and Attraction’s ‘ Taxonomic Review Committee’ are acknowledged for their contributions to the information presented herein. References Areces-Berazain, F. & Ackerman, J.D. (2016). Phylogenetics, delimitation and historical biogeography of the pantropical tree genus Thespesia (Malvaceae, Gossypieae). Botanic alJournal of the Linnean Society 181: 171-198. Arthan, W., Traiperm, P, Gale, S.W., Norsaengsri, M. & Kethirun, L. (2016). Re-evaluation of the taxonomic status of Hackelochloa (Poaceae) based on anatomical and phenetic analyses. BotanicalJournal ofthe Linnean Society 181:224-245. Barker, W.R. (2018). Notes on the taxonomy ofAmimlmnLindernia subg. Didymadenia (Linderniaceae). Swainsona 31:59-80. Barrett, M.D. (2018). Three new species of Asparagales from the Kimberley region of Western Australia. Telopea 21; 25-37. Beier, B.-A., Chase, M.W. & Thulin, M. (2003). Phylogenetic relationships and taxonomy of subfamily Zygophylloideae (Zygophyllaceae) based on molecular and morphological data. Plant Systematics and Evolution 240: 11-39. Brown, A.P, Llorens, T.M., Coates, D.J. & Byrne, M. (2018). Eremophila subangustifolia (Scrophulariaceae), a rare new species from the Mid West Region of Western Australia, with notes on E. microtheca. Nuytsia 29: 17-20. Butcher, R. (2018). Tephrosia pedleyi (Fabaceae: Millettiae), a new species from the west Kimberley of Western Australia. Nuytsia 29: 69-73. CHAH (Council of Heads of Australasian Herbaria) (2006). National Species List, https://id.biodiversity.org.au/name/ apni/206590 [accessed 25 October 2018]. Chomicki, G. & Renner, S.S. (2015). Watermelon origin solved with molecular phylogenetics including Linnaean material: another example of museomics. New Phytologist 205: 526-532. 16 Nuytsia Vol. 30 (2019) Crisp, M.D., Mant, J., Toon, A. & Cook, L.G. (2015). Australian spinifex grasses: new names in Triodia for Monodia and Symplectrodia. Phytotaxa 230: 293-296. Davis, R.W. & Hammer, T.A. (2018). Ptilotus yapukaratja (Amaranthaceae), a new species from the Gascoyne bioregion of Western Australia. Nuytsia 29: 157-160. Decaisne, J. (1844). Asclepiadaceae. In. Candolle, A.L.P.P. de (ed.). Prodromus Systematis Naturalis Regni Vegetabilis. Vol. 8. (Fortin, Masson et Sociorum: Parisiis.) Dixon, K.W. & Christenhusz, M.J.M. (2018a). Flowering in darkness: a new species of subterranean orchid Rhizanthella (Orchidaceae; Orchidoideae; Diurideae) from Western Australia. Phytotaxa 334: 75-79. Dixon, K.W. & Christenhusz, M.J.M. (2018b). Endangered fairies: two new species of Caladenia (Orchidaceae; Orchidoideae; Diurideae), from the bauxite plateaux of southwestern Western Australia. Phytotaxa 334: 87-90. Ford, A.J., Halford, D.A., Van Der Merwe, M. & Mathieson, M.T. (2017). A revision of the tropical white-flowered species of Comesperma (Polygalaceae) in Australia. Australian Systematic Botany 30: 159-182. French, M. (2012). Eiicalypts of Western Australia’s Wheatbelt. (Malcolm French: Perth.) Gagnon, E., Bruneau, A., Hughes, C.E., De Queiroz, E.P & Eewis, G.P (2016). A new generic system for the pantropical Caesalpinia group (Eeguminosae). PhytoKeys 71: 1-160. Guymer, G.P. (2013). Celtis strychnoides Planch, an earlier name for Celtis australiensis Sattarian (Ulmaceae). Austrobaileya 9: 146-147. Hammer, T.A. (2018). The Ptilotus murrayi species group: synonymisation of P. petiolatus under P. murrayi and description of the new Western Australian species P. unguiculatus (Amaranthaceae). Swainsona 31: 93-100. Hammer, T.A. & Davis, R.W. (2018). Ptilotus actinocladus (Amaranthaceae), a new and rare species from the Gascoyne bioregion. Western Australia. Nuytsia 29: 145-149. Hammer, T.A., Davis, R.W. & Thiele, K.R. (2018a). The showy and the shy: reinstatement of two species from Ptilotus gaudichaudii (Amaranthaceae). Australian Systematic Botany 31: 1-7. Hammer, T.A., Macintyre, P.D., Nge, F.J., Davis, R.W., Mucina, E. & Thiele, K.R. (2018b). The noble and the exalted: a multidisciplinary approach to resolving a taxonomic controversy within Ptilotus (Amaranthaceae). Australian Systematic Botany 3\ \ 262-280. Hopper, S.D. (2018). Natural hybridization in the context of Ocbil theory. South African Journal of Botany 118: 284-289, Appendix A. Hopper, S.D. & Brown, A.P (2006). Australia’s wasp-pollinated flying duck orchids revised (Paracaleana: Orchidaceae). Australian Systematic Botany 19: 211-244. Ito, Y, Tanaka, N., Garcia-Murillo, P. & Muasya, A.M. (2016). A new delimitation of the Afro-Eurasian plant genus Althenia to include its Australasian relative, Lepilaena (Potamogetonaceae) - Evidence from DNAand morphological data. Molecular Phylogenetics and Evolution 98: 261-270. Jobson, R.W., Baleeiro, PC. & Barrett, M.D. (2018). Six new species of Utricularia (Eentibulariaceae) from Northern Australia. TelopealV. 57-77. Jones, D.E. & Clements, M.A. (2017). New combinations in Australian Orchidaceae. Australian Orchid Review 82(2): 63. Jones, D.E. & French, C.J. (2018). New combinations in the Pterostylidinae. Australian Orchid Review 83(4): 55. Keighery, G.J. (2018). Swan Coastal Plain studies: taxonomic notes on Bossiaea eriocarpa Benth. and Bossiaea ornata (Eindl.) Benth. (Bossiaeeae: Fabaceae). Western Australian Naturalist 3\ ’. 95-104. Keighery, B.J. & Keighery, G.J. (2017). A limestone subspecies of Petrophile brevifoliaLindl. (Proteaceae). Western Australian Naturalist 3\ : 7-10. Keighery, G.J. & Keighery, B.J. (2018). Ursinia nana DC. (Asteraceae) is naturalised in Western Australia. We stern Australian Naturalist 31:7 8-81. Kuo, J. (2011). Cymodoceaceae. In. Wilson, A.J.G. (ed.). Flora of Australia. Vol. 39: Alismatales to Arales. pp. 120-134. (Australian Biological Resources Study: Canberra / CSIRO Publishing: Melbourne.) Earridon, E, Govaerts, R., Banters, K. & Goetghebeur, P. (2016). Cyperus albescens, a new combination in Cyperus (Cyperaceae) for the common (sub)tropical African and Asian species Lipocarpha chinensis. Kew Bulletin 71: 30. Eiede-Schumann, S. & Meve, U. (2018). Vincetoxicum (Apocynaceae—Asclepiadoideae) expanded to include Tylophora and ed\\QS. Phytotaxa 369'. 129-184. Eohr, M.T. & Keighery, G. (2016). Quercus (Fagaceae) in Western Australia. Western Australian Naturalist 39'. 172-175. Eowrie, A. (2013a). Carnivorous Plants of Australia Magnum Opus. Vol. 1. (Redfern Natural History Productions: Poole.) Eowrie, A. (2013b). Carnivorous Plants of Australia Magnum Opus. Vol. 2. (Redfern Natural History Productions: Poole.) Eowrie, A. (2014). Carnivorous Plants of Australia Magnum Opus. Vol. 3. (Redfern Natural History Productions: Poole.) J.M. Percy-Bower & C.M. Parker, Updates to Western Australia’s vascular plant census for 2018 17 Mabberley,D.J. (2008). Mabberley’s plant-book: aportable dictionary of plants, their classification anduses. 3'^‘'edn. (Cambridge University Press: Cambridge.) Mabberley, D. J. (2017). Mabberley’s plant-book: aportable dictionary of plants, their classification and uses. 4* edn. (Cambridge University Press: Cambridge.) Macfarlane, T.D., Sokolotf, D.D. & Remizowa, M.V. (2017). Filling a morphological gap between Australasian and Eurasian/ African members of Althenia Potamogetonaceae, Alismatales): A. hearnii sp. nov. from SW Western Australia. Phytotaxa 317: 53-60. Madhani, H., Rabeler, R., Pirani, A., Oxelman, B., Heubl, G. & Zarre, S. (2017). Untangling phylogenetic patterns and taxonomic confusion in tribe Caryophylleae (Caryophyllaceae) with special focus on generic boundaries. Taxon 67: 83-112. Maesen, L.J.G. van der (2012). The correct name for Flemingia involucrata^dW. ex Benth. (Leguminosae-Papilionoideae). Webbia; raccolta di scritti botanici 67: 33-36. Markey, A. (2018). By their fruit you will recognise them: species notes and typifications in Western Australian species of Opercidaria (Rubiaceae: Anthosperminae). Nuytsia 29: 119-140. Nicolle, D.L. & Barrett, R.L. (2018). Eucalyptus revelata, a rare new species related to E. mooreana (Myrtaceae) from the Kimberley region of Western Australia. Nuytsia 29: 109-118. Obbens, F.J. (2018). Three new perennial species of Calandrinia (Montiaceae) from southern Western Australia. Nuytsia 29: 193-204. Pellegrini, M.O.O., Flom, C.N. & Almeida, R.F (2018). Total evidence phylogeny of Pontederiaceae (Commelinales) sheds light on the necessity of its recircumscription and synopsis of Pontederia L. PhytoKeys 108: 25-83. Perkins, A. J. (2018a). Hydrocotyle spinulifera and H. dimorphocarpa (Araliaceae), two new Western Australian species with dimorphic mericarps. Nuytsia 29: 57-65. Perkins, A.J. (2018b). Hydrocotyle asterocarpa, H. decorata and H. perforata (Araliaceae), three new Western Australian species with spicate inflorescences. Nuytsia 29: 205-216. Petrongari, F.S., Simoes, A.R. & Simao-Bianchini, R. (2018). New combinations and lectotypiflcations in Distimake Raf (Convolvulaceae). Phytotaxa 340: 297-300. Robinson, A. S., Cross, A.T., Meisterl, M.E. &Fleischmann, A. (2018). A new pygmy sundew, Z)ro5erafl//jo«ototo(Droseraceae), from the western Wheatbelt and an updated diagnostic key to the orange-flowered pygmy Drosera of Western Australia. Phytotaxa 346: 221-236. Rye, B.E. (2018). An update to the taxonomy of some Western Australian genera of Myrtaceae tribe Chamelaucieae: 5. Hysterobaeckea. Nuytsia 29: 75-107. Salmaki, Y, Kattari, S., Fleubl, G. & Brauchler, C. (2016). Phylogeny of non-monophyletic Teucrium (Eamiaceae: Ajugoideae): Implications for character evolution and taxonomy. Taxon 65: 805-822. Sarkinen, T., Poczai, P, Barboza, G.E., van der Weerden, G.M., Baden, M. & Knapp, S. (2018). A revision of the Old World Black Nightshades (Morelloid clade of Solanum E., Solanaceae). PhytoKeys 106: 1-223. Schmidt-Eebuhn, A.N., Puttock, C.F, Davis, R. W. & Wege, J. (2018). Two new species of Ozothamnus (AstemcQdLe: Gnaphalieae) from Australia. Phytotaxa 336: 190-196. Shaik, R.S., Eepschi, B. J., Gopurenko, D., Urwin, N.A.R., Burrows, G.E. & Weston, E.A. (2016). An integrative morphological and molecular approach to identiflcation ofthree Australian cucurbitaceous invasive weeds: Citrullus colocynthis, C. lanatus and Cucumis myriocarpus. Australian Systematic Botany 29: 247-264. Shepherd, K.A. (2018). Tecticornia willisii (Chenopodiaceae), a new samphire from the Tittle Sandy Desert in Western Australia. Nuytsia 29: 141-144. Shepherd, K.A. & Wilkins, C.F. (2018a). Commersonia corniculata (Malvaceae), a newname for C. cygnorum. Nuytsia 29: 67-68. Shepherd, K.A. & Wilkins, C.F. (2018b). A taxonomic revision of species with a petaloid epicalyx bract allied to Lasiopetalum bracteatum (Malvaceae: Byttnerioideae). Nuytsia 29: 161-179. Shepherd, K.A. & Wilkins, C.F. (2018c). Typiflcation of Lasiopetalum and an interim key to the Western Australian species of the genus (Malvaceae: Byttnerioideae). Nuytsia 29: 181-192. Sim6es,A.R. & Staples, G.W. (2017). DissolutionofConvolvulaceaetribeMerremieaeandanewclassiflcationoftheconstituent genera. BotanicalJournal of the Linnean Society 183: 561-586. Smith, M.G. & Jones, A. (2018). Threatened and Priority Flora list 5 December 2018. Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 23 January 2019]. Thiele, K.R. (2018). Two new species of Hibbertia (Dilleniaceae) from Western Australia. Nuytsia 29: 151-155. Thompson, J. & Johnson, E.A.S. (1986). Callitris glaucophylla, Australia’s ‘White Cypress Pine’ - a new name for an old 18 Nuytsia Vol. 30 (2019) species. Telopeal: 731-736. Turner, I.M. (2016). Rather for the nomenclaturist than for the scientific botanist: The Botanical Cabinet of Conrad Loddiges & Sons. Taxon 65'. 1107-1149. Uribe-Convers, S. & Tank, D.C. (2016). Phylogenetic Revision of the Genus Bartsia (Orobanchaceae): Disjunct Distributions Correlate to Independent Lineages. Systematic Botany 41: 672-684. USDA, Agricultural Research Service, National Plant Germplasm System (2018). Germplasm Resources Information Network (GRIN-Taxonomy). https;//npgsweb.ars-grin.gov/gringlobal/taxonomydetail.aspx?id=455012 [accessed 23 July 2018]. Utz, Y.-A. & Gibson, R.P (2017). Drosera huegelii Endl. var. phillmanniana from the Stirling Range, south western Australia. Carnivorous Plant Newsletter 46: 132-138. Western Australian Herbarium (1998-). FloraBase—the Western Australian Flora. Department of Biodiversity, Conservation and Attractions, https://florabase.dpaw.wa.gov.au/ [accessed 23 January 2019]. Xue, B., Thomas, D.C., Chaowasku, T., Johnson, D.M. & Saunders, R.M.K. (2014). Molecular phylogenetic support for the taxonomic merger of Fitzalania and Meiogyne (Annonaceae): new nomenclatural combinations under the conserved name Meiogyne. Systematic Botany 39: 396-404. Zona, S. (2015). The correct gender of Schinus (Anacardiaceae). Phytotaxa 222'. 15-11. J.M. Percy-Bower^ and C.M. Parker Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 ‘Corresponding author, email: Julia.Percy-Bower@dbca.wa.gov.au Nuytsia The journal of the Western Australian Herbarium 30: 19-22 Published online 28 March 2019 SHORT COMMUNICATION Acacia corusca (Fabaceae: Mimosoideae), a new species from the Pilbara bioregion in north-western Australia Acacia corusca J.RBull, S.J.Dillon & Brearley, sp. nov. Type: c. 35 km east-northeast of Newman, Western Australia [precise locality withheld for conservation reasons], 26 April 2014, J. Bull & D. Roberts ONS A 27.01 {holo\ PERTH 08612277; iso\ CANB). Acacia sp. East Fortescue (J. Bull & D. Roberts ONS A 27.01), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au [accessed 13 February 2019]. Rounded to broadly rounded, robust, multi-stemmed shrub or small tree 1.5^.0(-5.0) m high, 1.5- 5.0(-6.0) m wide. Bark grey to dark grey, smooth except longitudinally Assured and fibrous towards the base of mature stems. Branchlets terete, slightly angular at extremities, youngest branchlets ribbed, with a dense indumentum of appressed, flattened, pale yellow to white, simple hairs and scattered, red-brown, glandular hairs between the ribs, and a dense indumentum of red-brown, glandular hairs on the ribs, a moderately thick layer of yellow, translucent resin often obscuring the indumentum, both the resin and indumentum becoming absent with age. New shoots resinous, the young phyllodes with a conspicuous marginal nerve invested with dense, red-brown, glandular hairs and the faces with a dense indumentum of pale yellow to white, appressed, flattened, simple hairs and scattered, red- brown, glandular hairs, the indumentum somewhat obscured by resin. Stipules triangular, red-brown, (0.4-)0.5-0.65 mm long. Phyllodes narrowly elliptic, narrowed at both ends, slightly kinked at the gland, (36-)38-72 mm long (occasionally interspersed with a few less than 35 mm long), 4.1-8.7 mm wide, l:w = 5.0-13.4, ascending to erect, straight to shallowly incurved or shallowly recurved, green to dark green, with a moderately dense indumentum of flattened, appressed, simple hairs and scattered red-brown glandular hairs, indumentum becoming sparser with age, slightly resinous; midrib (and sometimes 2 additional, imperfectly developed, longitudinal nerves either side of the midrib) straighter and slightly more pronounced than the minor nerves, often becoming obscure towards the phyllode apex, minor nerves numerous and anastomosing to form a dense reticulum with longitudinally elongated nerve-islands; marginal nerve pale yellow with moderately dense red-brown glandular hairs embedded in resin; apex acute to acuminate, innocuous, often shallowly recurved; pulvinus2.0-5.3 mm long; gland on upper margin of phyllode (2.9-)3.4-13.4 mm above the pulvinus, not prominent. Inflorescences simple or vestigial racemes 0.5-0.8 mm long, initiated in the axils of young phyllodes; peduncles 3.0-6.2 mm long, with a sparse to moderately dense indumentum of short, appressed hairs, resinous; basal peduncular bract persistent, single, ovate, 1.0-1.4(-l .65) mm long, yellow to red-brown, with a moderately dense indumentum of appressed, simple hairs; spikes (lO-)l 1.5-25(-27) mm long, flowers densely arranged. Bracteoles 0.9-1.1 mm long; claws narrowly oblong to linear, glabrous; lamina ovate, thickened proximally, ciliolate and with scattered, glandular hairs. Flowers 5-merous; sepals united for up to V 4 (-almost V 2 ) of their length, 0.5-1.1 mm long, narrowly ovate, slightly expanded at the apex, sparsely papillose on margins becoming denser at apex together with simple hairs, abaxial face at apex papillose; petals 1.3-1.8(-1.9) mm long, glabrous, 1-nerved; ovary densely sericeous. Pods narrowly oblong, flat, scarcely raised over seeds, 17-58 mm long, 2.7-5.1 mm wide, coriaceous- crustaceous, straight, resinous but not sticky, yellow-brown to brown, with a sparse indumentum of © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 20 Nuytsia Vol. 30 (2019) appressed, white hairs, with numerous anastomosing longitudinal nerves; marginal nerve discrete, yellow. longitudinal in pods, obloid-ellipsoid, (3.3-)3.5-5.5(-6.0)mm long, (1.5-)1.8-3.1 mm wide, brown; areole ‘u’-shaped, 0.2-0.3 mm long, 0.15-0.3 mm wide; funicle expanded into a poorly defined, once-folded, terminal aril. (Figure 1) Diagnosticfeatures. Acacia corusca can be distinguished from other Western Australian^c^zczh! species by the following combination of characters: phyllodes with 1 (-3) longitudinal nerves that are slightly more prominent than the minor nerves and with numerous anastomosing minor nerves that form a dense reticulum with longitudinally elongated nerve islands; a gland that is distinctly distant from the pulvinus resulting in a slight though distinct kink in the phyllode (Figure IB); the presence of red- brown glandular hairs on new growth and along the margins of the phyllodes; cylindrical inflorescences (Figure 1C); sepals that are united for %(-14) the length of the calyx; narrow, firm-textured pods with anastomosing longitudinal nerves (Figure ID). Selected specimens examined. WESTERN AUSTRALIA: [localities withheld for conservation reasons] 16 July 2015, J.5W//0NSAEF 1 (PERTH); 16 July 2015, J. 5 m//ONSAEF 3 (PERTH); 3 Aug. 2015, J. Bull & D. Brearley ONS 2.4 (PERTH); 3 Aug. 2015, J. Bull & D. Brearley ONS 2.5 (PERTH); 25 Apr. 2014, J. Bull & D. Roberts ONS A 25 (PERTH); 25 Apr. 2014, J. Bull & D. Roberts ONS A 57 (PERTH); 27 Apr. 2014, J. Bull & D. Roberts ONS A100 (PERTH); 8 July 2014, J. Bull & D. Roberts ONS S 71 (PERTH); 10 July 2014, J. Bull & D. Roberts ONS S 107 (PERTH); 13 Sep. 2014, J. Bull & D. Roberts ONS R 25 (PERTH); 13 Sep. 2014, J. Bull & D. Roberts ONS R 28 (PERTH). Phenology. Flowering has been observed from mid-autumn to late winter (April to August), and fruiting from late winter to mid-spring (August to October). Like many arid dwelling Acacia species, A. corusca appears to set fruit in response to adequate winter rainfall (Preece 1971; Mangadas & Fox 2002) but further study is required to confirm this. Distribution and habitat. Acacia corusca occurs east-northeast of the town of Newman in the southeast Pilbara bioregion of Western Australia. A total of 567 plants have been recorded from three populations covering approximately 8.1 hectares (Onshore Environmental 2014, 2015a). Extensive targeted surveys of the surrounding region were undertaken but no further populations were located (Onshore Environmental 2015a, 2015b). Acacia corusca grows in red-brown sandy-loam soils on hill crests, ridges, slopes and minor drainage lines upon low, subdued and undulating stony hills. It rarely grows on hill summits and the largest populations appear to prefer exposed hill ridges, outcrops and rocky hill slopes composed of Boolgeeda Iron Formation overlying Woongarra Rhyolite (Figure 1 A). Associated vegetation is typically spinifex hummock grassland of Triodiapungens and Triodia vanleeuwenii with scattered low trees of Eucalyptus leucophloia subsp. leucophloia. Acacia aptaneura and Grevillea berry ana, open shrublands of Grevillea wickhamii subsp. hispidula. Senna glutinosa and Acacia marramamba and scattered tussock grasses ofEriachne mucronata and Cymbopogon ambiguus. Conservation status. Acacia corusca is listed as Priority One under Conservation Codes for Western Australian Flora (Smith & Jones 2018), as Acacia sp. East Fortescue (J. Bull & D. Roberts ONS A 27.01). Etymology. The epithet is from the Latin corusco (to flash, glitter or shimmer), in reference to the shimmering effect of the dark green foliage on windy days. This character was valuable in locating populations of Acacia corusca from a distance, especially on typically exposed hill slopes. J.P. Bull, S.J. Dillon & D.R. Brearley, Acacia corusca (Fabaceae), a new species from the Pilbara 21 Figure 1. Acacia corusca. A - habit and habitat; B - branches showing ascending phyllodes with longitudinal reticulation and gland on adaxial edge; C - upper branch with both immature and mature cylindrical flowering spikes; D - immature pod with two seeds. Image from J. Bull & D. Roberts ONS R 28 (PERTH) (D). Photographs by Jerome Bull. Affinities. Acacia corusca appears to have affinities with A. melleodora Pedley and A. dictyophleba F.Muell, both of which also occur in the Pilbara. These taxa are similar to A. corusca in that they are rounded shrubs, have aphyllode nervature pattern comprising a dense reticulum of anastomosing nerves and have funicles with a poorly defined aril. However, bothH. melleodora and A. dictyophleba differ significantly from A. corusca in that they are glabrous; have phyllodes with a rounded apex, a gland positioned adjacent to the apical mucro and an open and net-like reticulum (rather than longitudinally elongated); globular inflorescences; sepals that are united almost to the apex; flat, papery pods; and have seeds with a larger areole (1.1-1.5 mm wide) that are transversely arranged in the pods. Common name. Shimmer Wattle. Acknowledgements The authors would like to thank Bruce Maslin for his advice, expertise and highlighting the significance of this taxon, and BHP Western Australia Iron Ore for providing research support. Jerome would like to thank Mr Daniel Roberts, Ms. Jessica Waters and Mrs Ellen Whitten (nee Palmer) for their work ethic and humour in the field whilst researching this taxon. 22 Nuytsia Vol. 30 (2019) References Mangadas, L.G. & Fox, J.E.D. (2002). Reproductive potential of Acacia species in the central wheatbelt: variation between years. Conservation Science of Western Australia A{3y. 147-157. Onshore Environmental Consultants Pty Etd. (2014). Orebody 31 Eevel 2 Flora and Vegetation Survey. March 2014. Internal report prepared for BHP Billiton Iron Ore Pty Etd. pp. 38-41. Onshore Environmental Consultants Pty Etd. (2015a). Targeted Survey for Acacia sp. East Fortescue. March 2015. Internal report prepared for BHP Billiton Iron Ore Pty Etd. Onshore Environmental Consultants Pty Etd. (2015b). Targeted Survey for Acacia sp. East Fortescue. Phase2. August 2015. Internal report prepared for BHP Billiton Iron Ore Pty Etd. Preece, PB. (1971). Contributions to the biology of Mulga. I. fXosNer'mg. Australian Journal of Botany 19: 21-38. Smith, M.G. & Jones, A. (2018). ThreatenedandPriorityFloralist 5 December 2018. Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 13 February 2019]. Western Australian Herbarium (1998-). FloraBase—the Western Australian Flora. Department of Biodiversity, Conservation and Attractions, https://florabase.dpaw.wa.gov.au/ [accessed 13 February 2019]. Jerome P. BulP, Steven J. Dillon^ and Darren R. Brearley^ ‘Onshore Environmental Consultants Pty Etd, PO Box 227, Yallingup, Western Australia 6282 ^Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Eocked Bag 104, Bentley Delivery Centre, Western Australia 6983 ‘Corresponding author, email: info@onshoreenvironmental.com.au Nuytsia The journal of the Western Australian Herbarium 30:23-24 Published online 28 March 2019 SHORT COMMUNICATION Gastrolobium sp. Harvey (G.J. Keighery 16821) (Fabaceae) is not distinct from G, capitatum The phrase-name Gastrolobium sp. Harvey (G.J. Keighery 16821) was placed on the Western Australian plant census in 2006 following the collection of three specimens (G.J. Keighery 16715,16788 & 16821) in 2005 from two small wetland reserves in the Pinjarra-Harvey area. It was given a Priority Two conservation listing in 2008 based on its restricted distribution and potential threats to habitat posed by the construction of the Perth-Bunbury Highway, a conservation status it still retains (Smith & Jones 2018). In the absence of contemporary documentation detailing the morphological basis for its recognition, it is not known with what species G. sp. Harvey was compared in order to determine that it was distinct. We suspect a comparison was made with G ebracteolatum G. Chandler & Crisp, under which all three specimens were originally placed, presumably because of a shared wetland habitat, similar long, narrow, often alternate leaves and a tall habit (variously recorded in the three specimens as ‘sprawling shrub 2 m tall X 2 m wide’, ‘twining erect shrub 2 m tall x 2 m wide’ and ‘slender erect scrambling shrub 1-2 m tall X 2 m wide’). There are, however, some major differences between the two, with G sp. Harvey having condensed axillary or terminal racemes (inflorescences 2-6-flowered; rachis 1-3 mm long) rather than elongate terminal racemes (inflorescences at least 20-flowered; rachis 60-180 mm long), and four to eight ovules rather than at least 15 (Chandler et al. 2002). Examination of the Gastrolobium R.Br. collection at the Western Australian Herbarium (PERTH) has since found that G sp. Harvey is comparable in its floral and foliar morphology to a large number of G capitatum (Benth.) G.Chandler & Crisp specimens. In particular, specimens referred to as a ‘narrow leafed form, generally recumbent, typically on/around swamps of the Swan Coastal Plain’ (E.A. Griflhn 1993, in sched.) match G sp. Harvey in all features other than the size of the plants (G. capitatum is typically a sprawling sub-shrub to 1 m tall). Two ‘Swan Coastal Plain Survey’ (Gibson et al. 1994) voucher specimens collected from seasonally wet sites and identified as Gastrolobium aff. capitatum (originally asNemcia aff. capitata; B.J. Keighery & N. Gibson 120 & 132) also match Griflin’s concept of this swamp form. Chandler et al. (2002: 679) noted that G. capitatum ‘grows in a variety of habitats, from wet to quite dry, on sandy to loamy soils in woodland or open forest’ but did not recognise any ecotypes as warranting formal taxonomic status. It is noteworthy that their specimen citations under G capitatum included B.J. Keighery & N. Gibson 120. Relative to plants collected from drier habitats there is a tendency for wetland specimens of G capitatum to have leaves that are narrower and for there to be a higher ratio of alternate to opposite leaves (note that a mixed phyllotaxis is the norm for this species). However, these are tendencies only and neither constitutes a consistent difference. The following specimens exemplify the overlapping nature of these characters: S. Turner 257 and S. Turner 310, from dry Banksia woodland in metropolitan Kiara and Noranda, respectively, have the narrow leaves (to 5 mm) typical of the wetland variant but with opposite leaves predominating; M.D.Crisp & W.M. Keys MDC 8943, from a wetland in the Busselton area, has mostly alternate leaves but their width (to 8 mm) is more commonly found in plants growing in dry habitats; M.J. Kealley 1304, from the Darling Range north-west of Bannister, has narrow, mostly alternate leaves but is from a dry, lateritic site. © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 24 Nuytsia Vol. 30 (2019) Examination of specimens of G. sp. Harvey against the G capitatum collection at PERTH cannot identify any morphological characters by which this entity might be considered distinct. The narrow leaf form and predominantly alternate phyllotaxis characteristic of G. sp. Harvey is shared with most G. capitatum specimens collected from seasonally wet areas (as well as some collected from drier habitats). Both phyllotaxis and leaf size must be regarded as variable characters in this species. The only feature of G sp. Harvey that falls outside the known parameters of G. capitatum is plant size. The recorded dimensions of up to 2 m high and 2 m wide are certainly well beyond the proportions otherwise recorded for G. capitatum. Perhaps the habitat at these sites is particularly favourable to the species or the areas are long-unbumt, or both, but even if the reasons for the large growth habit are genetic, a single difference of this kind does not provide a strong basis for the recognition of a distinct taxon. In summary therefore, we believe that G sp. Harvey is best regarded as a variant or ecotype of G capitatum. Taxonomic investigation of this phrase-name has also identified that the key to species of Gastrolobium (Chandler et al. 2002) contains contradictory information to that presented in species descriptions, such that good representatives of a species may not key out correctly. For example, lead 114 (Chandler et al. 2002: 627) provides the alternatives ‘rachis >5 mm long’ and ‘rachis <1 mm long’, but G. capitatum, which is recorded as having ‘rachis 1-2 mm long’ in its description (Chandler et al. 2002: 678), falls into neither category and can only be reached in the key if the first alternative is taken. Problems with the key may have contributed to the original misidentification of the three specimens that later became the basis for the phrase name G. sp. Harvey, which highlights the importance of a well-curated research herbarium collection for accurate species identification. Acknowledgements This taxonomic research was funded under the Western Australian Government’s Saving our Species biodiversity conservation initiative (2006-2008) and by a grant to the (then) Department of Environment and Conservation’s Science Division from Ministerial funds for Swan Bioplan: a new conservation plan for the Swan Coastal Plain and adjacent escarpments. References Chandler, G.T., Crisp, M.D., Cayzer, L. W. & Bayer, R.J. (2002). Monograph of Gastrolobium {YdbdiCQdLQ’. MfxhQMQdiQ). Australian Systematic Botany 15; 619-739. Gibson, N., Keighery, B.J., Keighery, G.J., Burbidge, A.H. & Lyons, M.N. (1994). A floristic survey of the southern Swan Coastal Plain. Unpublished report for the Australian Heritage Commission prepared by Department of Conservation and Land Management and the Conservation Council of Western Australia (Inc.) (Department of Conservation and Land Management and the Conservation Council of Western Australia: Perth.) Smith,M.G.& Jones, A. (2018). ThreatenedandPriority Floralist 05 December 2018.TtQ^wcimQoioffModwexsxfy^ComexNdiixon and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 21 January 2019]. Ryonen ButcheH and Michael Hislop Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 'Corresponding author, email: Ryonen.Butcher@dbca.wa.gov.au Nuytsia The journal of the Western Australian Herbarium 30:25-31 Published online 28 March 2019 SHORT COMMUNICATION Eremophila oldfieldii subsp./7«pw/«, E. sericea and E. xantholaema (Scrophulariaceae), three new taxa from Western Australia Eremophila oldfieldii F.Muell. subsp. papula A.RBr., subsp. nov. Type: Karara, Western Australia [precise locality withheld for conservation reasons], 22 September 2017, A.R Brown & R.W Davis AVB 4307 {holo: PERTH 09033971; iso\ AD, CANB, MEL). Eremophila oldfieldii subsp. Karara (D. Coultas s.n. PERTH 07341717), WestemAustralian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed October 2018]. Illustration. A.P Brown & B.J. Buirchell, A Field Guide to the Eremophilas ofW. Austral, p. 197 (2011), as E. oldfieldii subsp. Karara. An erect to spreading much branched shrub 1.5-3.0 m high, 2-A m wide. Branches grey, terete, tuberculate, young parts with sparse grey-white branched hairs, old parts glabrescent. Leaves green to grey-green, sessile, although gradually tapering to a short petiole-like base, alternate, erect or spreading, scattered along branches, oblanceolate, (20-)40-55(-65) mm long, (6-)7-10(-12) mm wide, the upper and lower surfaces glandular-pustulate; apex acute to subobtuse; margins entire. Flowers 1 per axil; pedicel terete basally, becoming dilated distally, straight or slightly curved, 10- 15 mm long, with scattered, grey-white branched hairs. Sepals 5, oblanceolate, attenuate, imbricate towards the base, equal or subequal, 12-18 mm long, 4-8 mm wide, not enlarging after flowering; outer and inner surfaces green, sometimes tinged with brown, with sparse short simple hairs. Corolla zygomorphic, 22-30 mm long, 8-10 mm wide; outer and inner surfaces orange-red to brick red, unspotted, glabrous with the exception of scattered simple hairs along the margins of lobes; lowermost lobe dilated, truncate to emarginate; upper 4 lobes acute. Stamens 4, exserted; filaments 25-30 mm long, glabrous; anthers glabrous. Ovary ovoid, 4-locular with 2 ovules per locule, 3-4 mm long, 2.0- 2.5 mm wide, glabrous or with sparse short simple hairs. Style 25-30 mm long, glabrous. Fruit dry, subglobular, 7-9 mm long, 4-5 mm wide, rugose, glabrous. Seed ovoid, 2.2-2.4 mm long, 1-1.2 mm wide. (Figure 1) Diagnostic features. Eremophila oldfieldii papula may be distinguished from other subspecies of E. oldfieldii F.Muell. by its glandular-pustulate leaves and the following combination of characters: branches tuberculate; leaves oblanceolate, 40-55 x 7-io mm. Other specimens examined. WESTERN AUSTRALIA: [localities withheld for conservation reasons] 16Nov. 2005,D. Coultas s.n. (PERTH); 22 Sep. 2010,D. Coultas & A. SaligariDCA5) Opp9(PERTH); 14 Nov. 2015, A. Crawford ADC 2686 (PERTH); 29 Sep. 2010, K. Greenacre & B. Stratton, KIOP 145-01 (PERTH). Phenology. Predominantly flowers from August to November with rare flowering following rainfall at other times of the year. Fruiting throughout the year following flowering. © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 26 Nuytsia Vol. 30 (2019) Figure 1. Eremophila oldfieldii subsp. papiz/tz. A-flowering plant in situ showing a much branched, spreading habit; B - flower, showing the large oblanceolate, attenuate sepals and unspotted, subglabrous corolla; C - leaves, showing the characteristic glandular-pustulate upper and lower surfaces of the lamina. Images from A.P. Brown & R.W. Davis APB 4307. Photographs by A.P. Brown. Distribution and habitat. Found over a small geographic range north-east of Perenjori in the Yalgoo bioregion (sensu Department of the Environment 2017), growing in red brown clay loam on the lower slopes of rocky hills. Associated species include Acacia tetragonophylla. Eucalyptus loxophleba subsp. supralaevis, Exocarpos aphyllus, Tecticornia pterygosperma subsp. pterygosperma and Mesembryanthemum nodiflorum. Conservation status. Currently listed as Priority One under Conservation Codes for Western Australian Flora (Smith & Jones 2018), under the name E. oldfieldii subsp. Karara (D. Coultas s.n. PERTH 07341717). The subspecies is known from rare, scattered populations in an area that may be subject to future mining. Etymology. From the Eatin papula (pimple or pustule), in reference to the glandular-pustulate leaves of this subspecies. Notes. Eremophila oldfieldii subsp. papula may be distinguished from the other two subspecies by its glandular-pustulate leaves and tuberculate branches. It also differs from subsp. oldfieldii in its shorter stature 1.5-3 m high when mature {cf. 3-6 m in subsp. oldfieldii) and usually shorter, broader leaves 40-55 X 7-10 mm {cf. 50-100 x 3-8 mm) and from subsp. angustifolia (S.Moore) Chinnock in its broader leaves 7-10 mm wide {cf. 1.5-2.5 mm in subsp. angustifolia). A.P. Brown & R.W. Davis, Three new Eremophila (Scrophulariaceae) from Western Australia 27 Although Eremophila oldfieldii subsp. papula is found near E. oldfieldii subsp. oldfieldii and occurs in similar habitat, they are not known to grow together. Eremophila sericea A.RBr., sp. nov. Type'. Karara, Western Australia [precise locality withheld for conservation reasons], 22 September 2017, A.P. Brown &R.W Davis APB 4308 {holo\ PERTH 09033963; iso\ CANB, MEL). Eremophila sp. Rothsay (D. Coultas & J. Kelt s.n. PERTH 08200440), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed October 2018]. Illustration. A.P Brown & B.J. Buirchell, A Field Guide to the Eremophilas ofW. Austral, p. 316 (2011), as E. sp. Rothsay. A small much branched shrub, 0.2-1.2 m high, 0.8-1.6 m wide. Branches grey, terete, non-tuberculate, young parts with a dense indumentum of grey-white, branched hairs, old parts glabrescent. Leaves grey-white, more rarely grey-green, shortly petiolate, opposite, erect or spreading, scattered along branches; pedicel terete, straight or slightly curved, 2-3 mm long, with crowded, grey-white branched hairs; lamina elliptic, 15-25 mm long, 6-14 mm wide, the upper and lower surfaces with crowded, grey-white branched hairs; apex obtuse to subobtuse, margins entire. Flowers 2 per axil. Sepals 5, lanceolate, attenuate, imbricate, equal, appressed against the corolla, 5-7 mm long, 1.5-2.0 mm wide, not enlarging after flowering; outer surface grey-white, more rarely grey-green, with a dense indumentum of short, grey-white branched hairs; inner surface grey-green with short, grey-white branched hairs in distal third, more sparsely hairy below except along margins. Corolla zygomorphic, 17-22 mm long, 8-10 mm wide, abruptly constricting above ovary; outer surface purple to pale lilac, unspotted, with crowded grey-white branched hairs; inner surface white, with scattered villous-arachnoid hairs extending down from below medial lobe of lower lip; lobes subequal, spreading, obtuse. Stamens 4, included; filaments 5-8 mm long, glabrous; anthers glabrous. Ovary ovoid-obloid, 4-locular with 1 ovule per locule, 3-5 mm long, 2-4 mm wide, with crowded branched hairs. Style 6-8 mm long, glabrous. Fruit dry, ovoid-conical, 6-8 mm long, 5-7 mm wide, rugose, with scattered branched hairs when young. Seed unknown. (Figure 2) Diagnostic features. Eremophila sericea may be distinguished from all other members of the genus by the following combination of characters: small much branched shrub up to 1.2 m high; branches, leaves and sepals with a dense indumentum of grey-white branched hairs; leaves shortly petiolate, opposite, lamina elliptic; flowers 2 per axil; sepals lanceolate, attenuate, imbricate, equal, appressed against the corolla; corolla purple to pale lilac, unspotted, abruptly constricting above the ovary; outer surface with crowded, grey-white branched hairs; inner surface white with scattered villous-arachnoid hairs extending down from below the medial lobe of lower lip. Other specimens examined. WESTERN AUSTRALIA: [localities withheld for conservation reasons] 31 Mar. 2015, J. Borger PJ 04 (PERTH); 26 Oct. 2009, D. Coultas & J. Kelt Opp 1 (PERTH); 16 Nov. 2009, D. Coultas & J. Kelt s.n. (PERTH); 15 Sep. 2010, D. Coultas & K. Greenacre DCKG Opp 1 (PERTH); 6 Jan. 2016, A. Crawford ADC 2727 (PERTH). Phenology. Predominantly flowers from September to November with rare flowering following rainfall at other times of the year. Fruiting throughout the year following flowering. 28 Nuytsia Vol. 30 (2019) Figure 2. Eremophila A - flowering plant in situ showing the much branched habit; B - flowering stem, showing the grey-white indumentum on the stems, leaves and sepals, the opposite leaves and the purple to pale lilac, unspotted corolla; C - close up of flower. Images from A.P. Brown & R.W. Davis APB 4308. Photographs by A.P Brown. A.P. Brown & R.W. Davis, Three new Eremophila (Scrophulariaceae) from Western Australia 29 Distribution and habitat. Found over a small geographic range north-east Perenjori in the Avon Wheatbelt and Yalgoo bioregions (sensu Department of the Environment 2017), growing in red brown clay loam on the lower slopes of rocky hills. Associated species 'mcXudQ Acacia andrewsii. Eucalyptus loxophleba subsp. supralaevis, E. salubris, Eremophila oldfieldii subsp. oldfieldii, E. scoparia and Senna charlesiana. Conservation status. Currently listed as Priority One under Conservation Codes for Western Australian Flora (Smith & Jones 2018), under the name E. sp. Rothsay (D. Coultas&J. Kelts.n. PERTH 08200440). The species is known from several mostly small populations. One population is on a weedy road verge and all others are in an area that may be subject to future mining. Etymology. From the Eatin sericeus (silky), in reference to the densely silky-hairy branches, leaves and sepals of this species. Affinities. Eremophila sericea appears closest in morphology to E. malacoides Chinnock and like that species is a densely hairy, much branched shrub up to 1.2 m high with opposite, elliptic leaves, lanceolate, attenuate sepals and a tubular, lilac to purple corolla that is constricted just above the ovary. Eremophila sericea may be distinguished from E. malacoides by its larger leaves, 15-25 x 6-14 mm {cf. 5-15 X 2.5-6.0 mm in E. malacoides), two flowers per axil (cf. one flower per axil) and smaller corolla 17-22 mm long (cf. 20-32 mm long). Notes. Eremophila sericea occasionally hybridises with E. scoparia (R.Br.) F.MuelL, producing offspring that are intermediate in morphology, i.e. R. Meissner & R. Coppen 4312. However, the parent species do not appear to be closely related as they differ in their stature, leaf width, sepal length and corolla indumentum. Eremophila xantholaema R.W.Davis, sp. nov. Type-. Bulong, Western Australia [precise locality withheld for conservation reasons], 16 September 2018, R. Davis & A. Brown RD 12904 (holo'. PERTH 08171033; iso'. CANB, MEE). Eremophila sp. Kalgoorlie (V. Clarke & A. Brown VTC 590), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed October 2018]. Illustration. A.P. Brown & B.J. Buirchell, A Field Guide to the Eremophilas ofW. Austral, p. 298 (2011), as E. sp. Kalgoorlie. Erect shrub to 1.2-3.0 m high, 0.9-1.3 m wide. Branches grey, compressed to terete, with a dense indumentum of short, appressed, flattened hairs. Leaves grey-green, opposite to sub-opposite; lamina linear, 25-40 mm long, 1.5-2.0 mm wide, terete to sub-terete and channelled, with crowded, appressed, flattened hairs intermixed with scattered, short glandular hairs; apex uncinate; margins entire. Flowers 1 per axil; pedicel straight or slightly curved, 2-3 mm long; with appressed, flattened hairs. Sepals 5, separated at base, sub-equal, spathulateto oblanceolate, 5-6 mm long, 1.2-2.5 mm wide, not enlarging after flowering (splaying slightly towards the apex in fruit); outer and inner surfaces mauve to pale pink, with crowded, flattened, appressed hairs, intermixed with scattered glandular hairs. Corolla zygomorphic, 9-12 mm long, 6.0-7.5 mm wide, outer surface pale pink and mauve, more rarely pale yellow and mauve, glabrous, inner surface below the lobes predominantly yellow with darker brown or orange markings, with densely tangled hairs in the throat; lobes mauve to pale pink, subequal. 30 Nuytsia Vol. 30 (2019) spreading, obtuse with sparse, short hairs. Stamens 4, included; filaments 4.5-5.5 mm long, with crowded, simple hairs on the lower half; anthers glabrous. Ovary ovoid, 4-locular with one ovule per locule, 1.3-1.5 mm long, 0.8-0.9 mm wide, with crowded simple hairs. Style 6-6.5 mm long, with scattered, long, simple hairs on the lower two thirds. Fruit dry, ovoid, 2.8-3.0 mm long, 1.5-1.7 mm wide, with crowded, simple hairs. Seed unknown. (Figure 3) Diagnostic features. Eremophila xantholaema may be distinguished from all other members of the genus by the following combination of characters: tall erect shrub up to 3 m high; sepals oblanceolate, free or fused only at the base; vegetative parts clothed in a fine pubescence of fiattened, appressed hairs; corolla mauve to pale pink, 9-12 mm long, the inner surface below the lobes predominantly yellow with brown or orange markings. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 16 Oct. 2005, V. Clarke & A. Brown VTC 590 (PERTH); 16 Oct. 2005, M.J. Grieve & J.D. Start D7 122 (PERTH). Phenology. Predominantly Powers from September to October with rare Powering following rainfall at other times of the year. Fruiting throughout the year following Powering. Distribution and habitat. Found near Bulong in the Coolgardie Bioregion (sensu Department of the Environment 2017), growing in stony, brown loam soils in Eucalyptus-Casuarina woodland on the upper slopes of low rocky hills. Associated species include Casuarina pauper, Eremophila glabra subsp. glabra, E. parvifolia subsp. auricampa, E. pustulata. Senna artemisioides s\xhs^.filifolia and Westringia rigida. Figure 3. Eremophila xantholaema. A - plant in situ showing the erect habit; B - leaves and flowers showing the terete or sub-terete grey-green leaves and flowers with sub-equal sepals and glabrous corolla with markings in the throat. Images from R. W. Davis & A.P. Brown RD 12904 (A) and V. Clarke & A. Brown VTC 590 (B). Photographs by A.P. Brown. A.P. Brown & R.W. Davis, Three new Eremophila (Scrophulariaceae) from Western Australia 31 Conservation status. Currently listed as Priority One under Conservation Codes for Western Australian Flora (Smith & Jones 2018) under the name E. sp. Kalgoorlie (V. Clarke & A. Brown VTC 590). The species is known from just three populations near Bulong. Etymology. The epithet is from the Greek xantho- (yellow-) and laimos (throat), in reference to the inner surface of the corolla which is predominantly yellow with brown or orange markings. Affinities. Eremophila xantholaema appears closest in morphology to E. oppositifolia R.Br. subsp. angustifolia (S.Moore) Chinnock and co-occurs with that taxon east of Kalgoorlie. Eremophila xantholaema may be distinguished from E. oppositifolia subsp. angustifolia by its usually shorter, sub-opposite leaves 25^0 mm long {cf 29-120 mm long and opposite in E. oppositifolia subsp. angustifolia), shorter corolla 9-12 mm long {cf 15-30 mm long) and the prominent yellow mottled markings in the throat of the corolla {cf no markings). Notes. Eremophila xantholaema was first recognised as distinct in October 2005, when plants were found near Bulong. The species has since been surveyed for over a wide area, with just two additional populations found, both within 10 km of the type location. Acknowledgements We would like to thank David Coultas and Greg Woodman who were the first to collect Eremophila oldfieldii subsp. papula and E. sericea and recognise them as distinct. We also thank Bevan Buirchell, Phil and Marlene James, and Jotf and Joan Start who have often accompanied us in the field and who have made valuable comments during discussions about these taxa. In addition, we thank Melanie Smith and Anthea Jones for assessing the conservation status of these taxa, the Curator and staff at the Western Australian Herbarium for access to specimens and Barbara Rye for useful comments and edits on the manuscript. References Department of the Environment (2017). Australia’s bioregions (IBRA), IBRA7, Commonwealth of Australia. https://www. environment, gov. au/land/nrs/science/ibra#ibra [accessed 2 July 2018]. Smith, M.G. & Jones, A. (2018). Threatened and Priority Flora list 5 December 2018. Department of Biodiversity, Conservation and Attractions. https;//www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 7 March 2019]. Andrew P. Brown^’^ and Robert W. Davis^ 'Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 ^Corresponding author, email: andrewbrown3@optusnet.com.au 32 Nuytsia Vol. 30 (2019) Nuytsia The journal of the Western Australian Herbarium 30:33-86 Published online 28 March 2019 An update to the taxonomy of some Western Australian genera of Myrtaceae tribe Chamelaucieae. 6. Scholtzia Barbara L. Rye Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 Abstract Rye, B.L. An update to the taxonomy of the Western Australian genera of the Myrtaceae tribe Chamelaucieae. 6. Scholtzia. Nuytsia 30: 33-86 (2019). Twenty five new species and five new subspecies of Scholtzia are described: S. bellairsiorum Rye, S. brevistylis Rye, S. brevistylis subsp. prow aka Rye, S. calcicola Rye, S. chapmanii Trudgen ex Rye, S. cordata Trudgen ex Rye, S. corrugata Rye, S. denticulata Rye, S. halophila Rye, S. halophila subsp. meridionalis Rye, S. halophila subsp. mortlockensis Rye, S. inaequalis Rye, S. laciniata Rye, S. longipedata Rye, S. longipedata subsp. procera Rye, S. multiflora Rye, S. oleosa Rye, S. peltigera Rye, S. pentamera Rye, S. pentamera subsp. collina Rye, S. prostrata Rye, S. quindecim Rye, S. recurva Rye, S. subsessilis Rye, S. tenuissima Rye, S. thinicola Rye, S. trilocularis Rye, S. truncata Rye, S. uniflora Rye and S. uniovulata Rye. Scholtzia sp. Shark Bay (M.E. Trudgen 7429) is reduced to synonymy under S. capitata Benth. and most of the other phrase names in the genus are treated as synonyms of the above new taxa. A key is given to aid the identification of approximately 40 species and subspecies. Most taxa have conservation priority. Introduction This paper on Scholtzia Schauer is one of a series designed to reduce the large backlog of unnamed members of Myrtaceae tribe Chamelaucieae and draw attention to any problematic taxa. Scholtzia is endemic to Western Australia, extending from near Carnarvon south to near Harvey. It is the largest genus of a proposed new subtribe (Rye et al. in prep.) that includes Babingtonia Lindl., Hysterobaeckea (Nied.) Rye, Malleostemon J.W.Green and Sannantha Peter G.Wilson. Apart from a recent paper establishing lectotypes (Rye 2017), there has been very little published regarding the taxonomy of Scholtzia since Bentham’s (1867) treatment of it in Flora Australiensis. Background When Schauer (1843, 1844) named Scholtzia, only two species, now known as S. obovata (DC.) Schauer and S. involucrata (Endl.) Druce, had been described. Schauer regarded Scholtzia as having axillary, dichotomously cymose infiorescences, anthers dehiscent by pores, and a 2-locular ovary with two ovules in each loculus, but was apparently unaware of the indehiscent fruit, which is one of the important characters that define the genus. He recorded 20 stamens, a common number in5. involucrata (see Figure lA), although stamen numbers range from about 15 to 30 in this species. © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 34 Nuytsia Vol. 30 (2019) A third species, now known as S. spatulata (Turcz.) Benth., was described by Turczaninow (1862), who treated it as a new genus, Piptandra Turcz. This species differed from the two previously named species in having a 3-locular ovary. Ten more names were published by Mueller (1864) and Bentham (1867), although not all of them are still considered to represent distinct species. Based on this larger number of species, Bentham (1867: 66) described the genus as having a 2- or 3-locular ovary, usually with two superposed ovules per loculus, but rarely one or three ovules per loculus. Actually, there is sometimes only one loculus and the ovule number is either one or two per loculus. Bentham also described the genus as having three kinds of fruits, valvate, indehiscent and schizocarpic, when in fact the fruits are uniformly indehiscent. Since 1867, only one species, S. eatoniana (Ewart & J.R.White) C.A.Gardner, has been named; altogether 12 of the named taxa in Scholtzia are considered here to be good species. A flora treatment for the Perth region (Rye 1987) treated only three species. Trudgen established manuscript names for two very distinctive new species during the 1990s, one of which was listed by Paczkowska and Chapman (2000), but both were later replaced by phrase names. A total of 44 phrase names have been applied to members of the genus (Table 1). Rye (2017) selected lectotypes for the genus and several species, and relegated two phrase names and one published name to synonymy. This left about 40 phrase names still in use prior to the current paper. Need for further work One of the unnamed species, S. sp. Geraldton (F. Lullfltz L 3216), does not match Scholtzia as circumscribed here, but does not appear to match any of the other named genera either (see Rye 2016: 105), so is retained under Scholtzia pending molecular studies to help establish its afflnities. Four of the phrase names that remain unaltered by the current study are of taxa that are considered to be too poorly collected to adequately assess and describe. These are S. sp. Bickley (W.H. Foaring s.n. PERTH 06165184), 5. sp. Nolba (E. Place s.n. Jan. 1964), 5. sp. Walebing (S. McNee 4) and 5. sp. Whelarra (M.E. Trudgen 12018). Several others belong to three complexes that need further study to determine how many taxa should be recognised: 1. Scholtzia involucrata complex. This has both low-growing plants and erect ones up to 3 m tall. 2. Scholtzia laxiflora Benth. complex. This is variable in characters such as leaf shape and peduncle length. Mike Hislop (pers. comm.) has observed two variants that sometimes co-occur but differ in habit, flowering time and microhabitat. 3. Scholtzia obovata complex. This complex has the longest geographic range and includes specimens currently identified as S. umbellifera F.Muell. Methods Measurements were taken from dried material using the largest leaves and bracts, and using fully mature floral parts that were well pressed. The stigma is recorded as peltate if it is more than twice as wide as its attachment point at maturity and as capitate if it is less than twice as wide. B.L. Rye, Update to the taxonomy of Scholtzia 35 Table 1. Phrase names and manuscript names for taxa in Scholtzia, with the year each was established and its new name or a comment. Phrase name or manuscript name Year Published name or comment Scholtzia chapmanii Trudgen ms 1993 Scholtzia chapmanii Trudgen ex Rye Scholtzia cordata Trudgen ms 1994 Scholtzia cordata Tmdgen ex Rye Scholtzia sp. Ajana (TA. Halliday 137) 1994 Scholtzia bellairsiorim Rye Scholtzia sp. Ajana East Road (M.E. Trudgen 21734 A) 2006 Scholtzia truncata Rye Scholtzia sp. Bickley (W.El. Eoaring s.n. PERTH 06165184) 2002 known from a single collection Scholtzia sp. Billeranga Hills (B.J. Conn 2159) 2010 Scholtzia siibsessilis Rye Scholtzia sp. Binnu (M.E. Trudgen 2218) 1994 Scholtzia uniflora Rye Scholtzia sp. Binnu East Road (M.E. Trudgen 12013) 1995 Scholtzia thinicola Rye Scholtzia sp. Binnu-Yuna (M.E. Trudgen 12016) 2003 Scholtzia longipedata subsp. procera Rye Scholtzia sp. Bungabandi Creek (M. Quick EURA 48) 2004 Scholtzia peltigera Rye Scholtzia sp. Burma Road (A.C. Burns 138) 2002 Scholtzia prostrata Rye Scholtzia sp. Coburn (N. Murdock NM 031) 2018 Scholtzia corrugata Rye Scholtzia sp. Coomberdale (M.E. & M.E. Trudgen MET 1724) 2006 Scholtzia halophila Rye subsp. halophila Scholtzia sp. Dongara (R. Hart 8401) 2003 Scholtzia calcicola Rye Scholtzia sp. Duck Pool (M.E. Trudgen MET 5427) 2006 Scholtzia halophila subsp. mortlockensis Rye Scholtzia sp. East Yuna (A.C. Bums 6) 1994 Scholtzia inaequalis Rye Scholtzia sp. Eneabba (S. Maley 8) 1996 Scholtzia trilocularis Rye Scholtzia sp. Eradu (R.D. Royce 8016) 1994 Scholtzia longipedata subsp. procera Rye Scholtzia sp. Eurardy (J.S. Beard 6886) 1994 Scholtzia oleosa Rye Scholtzia sp. Folly Hill (M.E. Trudgen 12097) 1995 belongs to Scholtzia obovata (DC.) Schauer complex Scholtzia sp. Galena (W.E. Blackall 4728) 1994 Scholtzia truncata Rye Scholtzia sp. Geraldton (F. Eullfitz E 3216) 1994 atypical of the genus in ovule number Scholtzia sp. Gunyidi (J.D. Briggs 1721) 1994 Scholtzia quindecim Rye Scholtzia sp. Jurien (R.J. Cranfield & PJ. Spencer RJC 8443) 2003 belongs to Scholtzia involucrata (Endl.) Druce complex Scholtzia sp. Kalbarri (N. Hoyle 623) 1996 Scholtzia oligandra F.Muell. ex Benth. Scholtzia sp. Kojarina (A.M. Ashby 1904) 1994 Scholtzia multiflora Rye Scholtzia sp. Eancelin (M.E. Trudgen 1516) 2003 Scholtzia laciniata Rye Scholtzia sp. Murchison (M.E. Trudgen 1685) 2002 belongs to Scholtzia laxflora Benth. complex Scholtzia sp. Murchison River (A.S. George 7908) 71994 Scholtzia denticulata Rye Scholtzia sp. Nolba (E. Place s.n. Jan. 1964) 1994 known from a single collection Scholtzia sp. Northampton (A Strid 20714) 1995 Scholtzia pentamera Rye subsp. pentamera Scholtzia sp. Overlander (M.E. Trudgen 12138) 2018 Scholtzia recurva Rye Scholtzia sp. Prowaka Springs (R.J. Cranfield & P. Spencer 8083) 1995 Scholtzia brevisiylis subsp. prowaka Rye Scholtzia sp. Red Bluff (A. Gunness 2373) 1996 Scholtzia longipedata Rye subsp. longipedata Scholtzia sp. Ross Graham Eookout (S Maley 6) 1994 Scholtzia denticulata Rye Scholtzia sp. Shark Bay (M.E. Trudgen 7429) 1996 Scholtzia capitata F.Muell. ex Benth. 36 Nuytsia Vol. 30 (2019) Phrase name or manuscript name Year Published name or comment Scholtzia sp. Valentine Road (S. Patrick 2142) 1995 Scholtzia thinicola Rye Scholtzia sp. Walebing (S. McNee 4) 2006 known from a single collection (see S. halophila) Scholtzia sp. Whelarra (M.E. Trudgen 12018) 1994 known from a single collection (see S. tenuissima) Scholtzia sp. Winchester (C. Chapman s.n. PERTH 05625386) 2010 Scholtzia chapmanii Trudgen ex Rye Scholtzia sp. Wonganderrah (M.E. & M.R. Trudgen MET 12000) 2003 belongs to Scholtzia involucrata (Endl.) Druce complex Scholtzia sp. Yandanooka (R. Soullier 646) 1999 Scholtzia brevistylis Rye Scholtzia sp. Yenyening Takes (A. Gunness 2824) 2003 Scholtzia halophila subsp. meridionalis Rye Scholtzia sp. Yerina Springs (N. Hoyle 517) 2002 Scholtzia capitata F.Muell. ex Benth. Scholtzia sp. Yuna (C.A. Gardner 14286) 2010 Scholtzia cordata Trudgen ex Rye Scholtzia sp. Z-Bend (Bellairs-Kalflora 912a) 1994 Scholtzia tenuissima Rye Fruit measurements were only taken from fertile fruits as far as possible. Fully mature fruits that are sterile contain hard structures formed from unfertilised ovules or inviable seeds; they tend to be somewhat larger than fertile fruits with a fully formed viable seed enclosed. A similar phenomenon was observed in Thryptomene Endl. (see Rye 2014: 274). Taxonomy Scholtzia Schauer, Linnaea 17: 241 (1843). Baeckea sect. Scholtzia (Schauer) Bail!., Hist. PL 6: 358 (1876). Type: Baeckea involucrata Endl, lecto,fide B.E. Rye, Nuytsia 27: 160 (2017) [= Scholtzia involucrata (Endl.) Druce]. [Pritzelia Schauer ms, Flora 27: 407 (1843), nom. inval. & nom. nud] Piptandra Turcz., Bull. Soc. Imp. Naturalistes Moscou 35: 323 (1862). Type'. Piptandra spatulata Turcz. [= Scholtzia spatulata (Turcz.) Benth.]. Shrubs prostrate to tall, up to 3(^) m high, mostly without a lignotuber, glabrous. Young stems developing a loose, whitish epidermis, which is smooth or fairly smooth in nearly all species. Leaves opposite. Petioles usually well defined, but sometimes very reduced or absent. Leaf blades dorsiventrally compressed, obtuse. Peduncles up to 25 mm long, often many-fiowered and with secondary axes, rarely consistently 1 -fiowered. Sepals often of varied shape within a flower, usually much shorter than, but up to about as long as, the petals, persistent in fruit, often with a longitudinal ridge on the base. Petals broadly ovate to circular, 1.0-4.5 mm long, white or pink, abruptly narrowed to a short claw at base, deciduous or (in some species) persistent (closed erect) in fruit. Stamens 3-30, antisepalous (often in irregular antisepalous groups) in most species, in a continuous circle in some species. Anthers (including basal connective gland) bent more or less at right angles to the filament to face the centre of the fiower, somewhat 2-lobed, dehiscent by 2 terminal pores or short, ± vertical slits. Ovary 1-3-locular (rarely 4-locular in one taxon); ovules 1 or 2 per loculus, if 2 then superposed. Style 0.3-3.5 mm long; base inset in a cylindrical depression. Fruits indehiscent, inferior to c. 1/2 inferior, usually 1-seeded. Seeds unfacetted, usually ± obovoid, 0.8-1.9 mm long; testa thin, membranous. Size and distribution. A Western Australian genus of more than 40 species, extending from near B.L. Rye, Update to the taxonomy of Scholtzia 37 Carnarvon south to near Harvey and inland to Anderson Rocks, north of Hyden, with a concentration of species in the northern sandplains of the Southwest Botanical Province. Kalbarri National Park is particularly rich, including more than a quarter of the species. Etymology. The genus is dedicated to Johann Eduard Heinrich Scholtz (1812-1859), a physician and naturalist from Bratislava, who studied the flora and fauna of one of the provinces of Poland. Gender: feminine. Chromosome numbers. Several species have been recorded with n = \\ and a single species, S. drummondii Benth., with the tetraploid number of w = 22 (Rye 1979). Horticultural potential and fire tolerance. Several species are in cultivation as garden ornamentals. They have the advantage of being relatively drought-tolerant plants for Perth gardens, since most come from drier habitats. The local species S. involucrata is one of the best known species as it is widespread in the Perth metropolitan area. It is a small, single-stemmed shrub, often low-growing, and is readily killed by fires. Seedlings are commonly produced in the two years after fires, but rarely, if ever, during fire-free periods. Some single-stemmed species of Scholtzia are large, sturdy shrubs that may be able to survive low-intensity fires. Species groups. Although Scholtzia is one of the largest genera in the tribe Chamelaucieae, no formal or informal infra-generic classification has been proposed for it. Stamen number and arrangement are of importance in distinguishing individual species or groups of species in the genus. When stamens are 15 or more, they are arranged in a complete circle (Figure lA), and when of moderate number or fewer they are usually in obvious antisepalous groups (Figure IB, C), although sometimes the stamens of an antisepalous group are quite widely separated. When there are very few stamens per flower, some of the antisepalous groups may be absent (see Figure ID). Scholtzia species can be divided into two groups based on whether they have two superposed ovules, or a solitary ovule, in each loculus of the ovary. Both situations are unknown in related genera, which have 3-25 radially arranged ovules per loculus or rarely two collateral ovules. Most of the Scholtzia species with only one ovule per loculus have three ovary loculi, whereas species with two ovules per loculus are much more likely to have two loculi. However the number of loculi ranges from one to three in both categories, with just one member of each having a one-locular ovary. Past reliance on keys such as Blackall and Grieve (1980) has led to one-locular specimens often having been misidentifled as members of consistently one-locular genera, such as Thryptomene and Malleostemon J.WGreen. A much less complete difference between the two main groups is in stamen arrangement, but this is only evident when stamen numbers are low, as most members of the 2-ovulate group appear to always have at least one stamen opposite each sepal, so that the minimum number of stamens per flower is five, whereas members of the 1 -ovulate group have up to two sepals without a stamen and a minimum number of three stamens per flower. However, S. brevistylis of the former group has antisepalous groups of 0-2 stamens, with a minimum of four stamens per flower. In this species, as in the 1-ovulate group, the most common arrangement of the stamens when there are five per flower is 2,0,1,2,0 around the circumference such that two sepals lack stamens but two others have two stamens and one has a single stamen. 38 Nuytsia Vol. 30 (2019) Figure 1. Images of Scholtzia species showing varied stamen arrangements. A- numerous stamens in a continuous circle in S. involucrata; B - ten antisepalous stamens in the arrangement 3,1,3,1,2 in a flower of S. drummondii; C - nine or ten stamens per flower, arranged mostly in close pairs opposite each sepal, in S. halophila, D - three or four, widely spaced stamens per flower in S. uniovulata, showing one enlarged flower with four antisepalous stamens in the arrangement 2,0,1,1,0. Taken by Kevin Thiele at Inglewood, voucher K.R. Thiele 3159 (A) and near Kellerberrin, voucher K.R. Thiele 3859 (D); by Rob Davis from near Irwin River, voucher R. Davis 11272 (B); and by Jean Hort from near Cunderdin (C). B.L. Rye, Update to the taxonomy of Scholtzia 39 Key to species and subspecies of Scholtzia *taxa that appear in more than one section of the key 1 . Stamens (10-) 13-30, in a continuous circle, with antipetalous stamens as well as antisepalous ones 2 . Ovary 3-locular; ovules 1 per loculus. Erect shrubs up to 2.5 m high, occurring north of Geraldton 3. Peduncles absent or up to 0.5 mm long, 1-flowered. Sepals 0.7-1.3 mm long (Binnu area). S. uniflora 3: Peduncles 7-13 mm long, 5-15-flowered. Sepals 0.4-0.8 mm long (Nolba area). S. sp. Nolba 2 . Ovary 2-locular; ovules 2 per loculus, superposed. Mostly prostrate or low- growing shrubs but one variant of S. involucrata up to 3 m high, occurring south of Geraldton 4 . Leaves markedly laciniate-ciliate; longest cilia 0.4-0.8 mm long (near Lancelin-N of Seabird-Moore River NP). S. laciniata 4 : Leaves entire to moderately laciniate-ciliate; longest cilia (when present) up to 0.4 mm long 5. Peduncles 3-7-flowered. Outer (and inner) sepals scarious throughout, not ridged (Eneabba-near Harvey). S. involucrata complex 5: Peduncles 1-3-flowered, all or mostly 1-flowered in most taxa. Outer sepals longitudinally ridged at the base 6. Leaves about as thick as wide, with margins poorly deflned (rounded) and entire except for a few apical teeth. Llowers borne usually at 2-7 consecutive nodes (Meckering-Youndegin area). S. eatoniana 6: Leaves usually wider than thick, angled on each edge to form distinct margins, often denticulate along most or distal parts of the margins. Llowers borne at up to 35 consecutive nodes, rarely all at fewer than 10 nodes 7. Leaves ovate to linear, 3.5-10 mm long. Peduncles 1-3-flowered 8. Leaves flat, with the longest lacinia or cilia 0.2-0.4 mm long. Petals 3.5-4.5 mm long. Antipetalous stamens with a filament c. 3.3 mm long (Bickley area). S. sp. Bickley 8: Leaves usually somewhat thickened on a fold, entire or denticulate- ciliolate. Petals 2.5-3 mm long. Antipetalous stamens with a filament 1.8-2.4 mm long (near Eneabba-near Moore River). S. teretifolia 7: Leaves obovate to broadly elliptic or narrowly obovate to linear, 1.2-5 mm long. Peduncles all or nearly all 1-flowered 9. Leaves ± entire. Peduncles 3-7 mm long. Mature style (including embedded part) commonly 2.5-3.4 mm long (Gunyidi-Dalwallinu). S. quindedm 9: Leaves toothed along the margins, sometimes becoming entire with age. Peduncles 1.5-3 mm long. Mature style (including embedded part) 1.5-2.3 mm long 10 . Leaves narrowly obovate to linear, 1.2-3 mm long. Hypanthium distinctly 5-ribbed, each rib connecting to a marked ridge on the base of a sepal. Stamens 15-20 (W of Eneabba-near Winchester) S. chapmanii 40 Nuytsia Vol. 30 (2019) 10 : Leaves obovate to broadly elliptic, 2.5-4.5 mm long. Hypanthium not appearing 5-ribbed, the sepals only moderately ridged at the base. Stamens 11-17 (E of Walkaway).S. prostrata 1 : Stamens 3-12(-14), grouped opposite the sepals, with no antipetalous stamens 11 . Ovules 1 per loculus 12 . Leaves rather densely covered by fairly uniform oil glands on abaxial surface; foliar colleters up to 0.6 mm long on each side of the petiole of young leaves. Sepals fairly erect in flower and fruit 13 . Leaves broadly to depressed obovate, 0.5-2.5 mm long, usually with midvein not noticeably raised or only at the apex. Hypanthium about as broad as long. commonly 1.5-1.6 mm long in fruit Peduncles 3-14-flowered (Coburn Stn-near Binnu).S. oleosa 13 : Leaves narrowly to broadly obovate, 2.5-5 mm long, usually with midvein ridged for about half or more of the lamina. Hypanthium longer than broad, commonly 2-2.5 mm long in fruit. Peduncles 9-23-flowered (Tamala Stn-Yerina Springs) .S. capitata 12 : Leaves with few or more scattered oil glands on abaxial surface, usually with central glands much larger than lateral ones; foliar colleters minute or apparently absent. Sepals somewhat to markedly incurved in flower and fruit in most species but spreading in fruit in S. uniovulata 14 . Hypanthium wrinkled-rugose or reticulate, with lowered areas not very deep or not restricted to oil glands 15 . Llowers borne at 5-42 consecutive nodes on most branchlets and with many or all of them occurring well below apex of flowering branchlets (i.e. not all in terminal clusters). Most flowers with 4-6 stamens and a 2-locular ovary, and all with sepals of fairly uniform length 16 . Leaf blades 2.3-3 mm wide. Peduncles 5-7 mm long, 3-15-flowered (near Kojarena-E of Walkaway).S. multiflora 16 : Leaf blades 1.8-2.3 mm wide. Peduncles 2-2.5 mm long, 1-3-flowered (near Winchester).S. atf. multiflora 15 : Llowers borne at l-5(-8) consecutive nodes on most branchlets, either all in ± terminal clusters or at very few nodes. Llowers not matching above choice in all characters, most commonly by having 6-11 stamens or a 3-locular ovary 17 . Leaf blades 2-7 x 2-4 mm, scarcely thickened (Eurardy Stn area- Canning River).S. laxiflora complex 17 : Leaf blades 1.0-2.5 x 0.8-2.1 mm, usually distinctly thickened towards the centre 18 . Peduncles 3-5 mm long, 3-7-flowered. Llowers most commonly with a 3-locular ovary and 5 stamens 19 . Leaf blades obovate, 0.8-1.4 mm wide. Hypanthium 0.7-0.9 mm long. Petals 1.0-1.2 mm long (Dongara-Drovers Cave NP).S. calcicola 19 : Leaf blades broadly obovate, 1.3-2.1 mm wide. Hypanthium 1.2-1.4 mm long. Petals 1.3-1.6 mm long (Coburn-southern Zuytdorp Clilfs).S. corrugata 18 : Peduncles 0.4-2.3 mm long, 1-3-flowered. Llowers either with a 2-locular ovary or mostly with more than 5 stamens B.L. Rye, Update to the taxonomy of Scholtzia 41 20. Inner sepals much larger than outer ones. Stamens 3-6 (East Yuna NR-Wicherina area). S. inaequalis 20: Inner sepals not markedly dilferent from outer ones. Stamens mostly 6-11 (N of Eurardy Stn-Indarra). S. truncata* 14: Hypanthium rugose-pitted, the shallow to very deep pits containing a sunken oil gland 21. Peduncles 3.5-14 mm long, 3-12-flowered 22. Petioles absent or up to 0.25 mm long. Occurring in non-saline habitats (Morawa area) .S. subsessilis* 22: Petioles 0.3-0.7 mm long. Occurring in subsaline habitats 23. Stamens 5-8. Ovary 2-locular in all or most flowers (Yenyening Eakes NR).S. halophila subsp. meridionalis 23: Stamens 7-12. Ovary 2- or 3(4)-locular, most specimens with about half or more of the ovaries 3-locular 24. Inner sepals 1.0-1.4 mm long (Mortlock River branches).. S. halophila subsp. mortlockensis 24: Inner sepals 0.5-0.8 mm long (near Coorow-Tammin area).S. halophila subsp. halophila 21. Peduncles 0.4-2.5(-3.5) mm long, 1-6-flowered 25. Ovary 3-locular. Peduncles at least 2.5 mm long and 3-6- flowered (Morawa area).S. subsessilis* 25: Ovary 1- or 2-locular (possibly rarely 3-locular in S. truncata). Peduncles 0.5-2.5 mm long and/or 1-3-flowered 26. Stamens (5-)7-12, always more than 6 in some flowers. Mature style (including embedded part) 0.9-1.6 mm long (N of Eurardy Stn-Indarra).S. truncata* 26: Stamens 3-6, always 5 or fewer in most flowers. Mature style (including embedded part) 0.45-0.8 mm long 27. Sepals mostly spreading in fruit; innermost one 0.8-1.2 mm long. Hypanthium pitted but usually not as deeply as in S. parviflora. Ovary 1-locular or much less commonly 2-locular (Northampton-Corrigin- Anderson Rocks). S. uniovulata 27: Sepals more erect or incurved in fruit; innermost one 0.3-0.6 mm long. Hypanthium deeply rugose-pitted. Ovary 2-locular (Hill River area-Moore River-Watheroo NP).S. parviflora 11: Ovules 2 per loculus, superposed 28. Ovary 3-locular in all or most flowers or (in S. oligandra) sometimes 2-locular in up to half of the flowers 29. Stamens 5-8, commonly 5 with 1 opposite each sepal. Ovary 2-locular in up to half of the flowers. Mature style (including embedded part) 0.6-1 mm long (Kalbarri-near Eucky Bay). S. oligandra 42 Nuytsia Vol. 30 (2019) 29: Stamens usually 8-14, rarely 6 or 7 (but then with mature style 1-1.6 mm long), with 0-4 opposite each sepal. Ovary 3-locular in all or nearly all of the flowers. Mature style (including embedded part) 0.7-1.6 mm long 30. Peduncles 0.4-6(-10) mm long, 1-3-flowered 31. Leaves ciliate to laciniate; longest cilia 0.2-0.4 mm long. Peduncles 0.4-2.1 mm long. Stamens 6-10 (Howatharra Hills area-N of Irwin River).S. dliata 31: Leaves entire, denticulate or ciliolate; longest cilia less than 0.2 mm long. Peduncles 2-6(-10) mm long. Stamens 10-14 (Maya-Tammin area). S. drummondii 30: Peduncles 4-25 mm long, mostly 3-15-flowered 32. Bracts 2-5 mm long. Petals 2.5-3.5 mm long. Largest stamens usually with a filament 1-1.3 mm long (Kalbarri NP-Binnu area -Howatharra).S. spatulata 32: Bracts 0.9-1.8 mm long. Petals 1.5-2.5 mm long. Largest stamens with a filament 0.4-0.7 mm long (E of Geraldton- Watheroo NP).S. trilocularis 28: Ovary 1- or 2-locular in all or most flowers 33. Leaves broadly to depressed orbicular-cordate or broadly ovate to depressed obovate, consistently broader than long 34. Leaves sessile, somewhat stem-clasping at base, 1.3-1.8 mm long, 1.5-2.5 mm wide, 0.5-1.1 mm thick (Eurardy Stn).S. peltigera 34: Eeaves distinctly petiolate at base, 2.2-7 mm long, 2.5-9 mm wide, not very thick 35. Flowers borne at 1-4 consecutive nodes. Mature style 0.8-1.4 mm long (Kalbarri NP-N of Yuna).S. cordata 35: Flowers mostly borne at 8-23 consecutive nodes. Mature style 0.3-1.0 mm long 36. Stamens always or mostly 6-10 per flower, 1-3 opposite each sepal. Bracts 0.6-1.6 mm long. Mature style 0.45-1.0 mm long 37. Eeaf blades broadly or very broadly obovate, with lateral veins usually obscure. Peduncles 2-4 mm long. Petals persistent in fruit (Kalbarri NP area).S. bellairsiorum 37: Eeaf blades broadly to depressed orbicular-cordate, with lateral veins clearly visible. Peduncles 5-15 mm long. Petals deciduous in fruit (Murchison River-near Balline Homestead).S. uberiflora 36: Stamens always or mostly 5 per flower, 1 opposite each sepal. Bracts 1.3-3 mm long. Mature style 0.3-0.4 mm long 38. Bracts 1.3-1.8 mm long. Ovary 1-locular (Ajana- Greenough).S. pentamera subsp. pentamera 38: Bracts 2-3 mm long. Ovary 2-locular (Moresby Range).S. pentamera subsp. collina 33: Eeaves of varied shape, all or mostly longer than broad 39. Petioles 0.1-0.35 mm long B.L. Rye, Update to the taxonomy of Scholtzia 43 40 . Leaves with a recurved apex produced into a minute point (Meadow Stn area). S. recurva 40 : Leaves with a straight apex and no point (except sometimes on the youngest leaves of S. thinicola) 41 : Peduncles 0.7-4 mm long, 0.4-0.7 mm wide. Sepals with a herbaceous base that is longitudinally ridged (near Yandi Stn -W of Mullewa).S. thinicola 41 . Peduncles 5.5-13 mm long 0.15-0.35 mm wide. Sepals scarious throughout, not ridged 42 . Leaves ± obovate, 0.6-1.0 mm wide. Secondary axes up to 2.5 mm long (Kalbarri NP). S. tenuissima 42 : Leaves obovate to broadly elliptic, 1.0-1.5 mm wide. Secondary axes up to 0.7 mm long (E of Binnu). S. sp. Whelarra 39: Petioles 0.4-1.1 mm long 43 . Stamens 4-6, usually 5, with no stamens opposite one or two sepals of each flower 44 . Leaves 1.3-1.6 mm wide, with fairly flat margins. Inflorescences extending for 3-8 nodes, tending to be concentrated into a small zone towards the ends of the branchlets; peduncles of the lowest node 2-4 mm long (Carnamah-Coorow area).S. brevistylis subsp. prowaka 44 : Leaves either wider than 1.6 mm or with recurved margins (at least towards the base). Inflorescences extending for 5-14 nodes but usually spike-like, usually well below the apex of each branchlet; peduncles of the lowest node l-2(-3) mm long 45 . Leaves with margins recurved, at least at base, the largest ones 1.0-1.5 mm wide. (Mingenew area-Yandanooka area)....S. brevistylis subsp. brevistylis 45 : Leaves with margins fairly flat, the largest ones 2.0-2.4 mm wide. (Nebroo Hill area).S. brevistylis unnamed variant 43 : Stamens 5-11, mostly more than 5, with at least one stamen opposite each sepal 46 : Leaves entire. Flowers with a hypanthium 1.0-1.6 mm long and sepals up to 0.75 mm long 47. Outer sepals 0.3-0.5 mm long, smooth or shortly ridged. Stigma peltate or capitate (Kalbarri NP-E of Balline).S. longipedata subsp. longipedata 47 : Outer sepals (0.4-)0.5-0.75 mm long, with base ridged for most of length. Stigma capitate (East Yuna NR - Eradu).S. longipedata subsp. procera 46 . Eeaves initially denticulate-ciliolate, sometimes becoming entire as they age. Flowers either with a long hypanthium (more than 1.6 mm long) or long sepals (more than 1 mm long) 48 . Hypanthium 1.0-1.6 mm long, rugose, not ribbed. Inner sepals 1.3-2.5 mm long (Kalbarri NP).S. denticulata 48 : Hypanthium 1.8-2.5 mm long, 5-ribbed at least in distal part. Inner sepals 0.3-0.6 mm long (N of Carnarvon-Eancelin).S. obovata complex 44 Nuytsia Vol. 30 (2019) Scholtzia bellairsiorum Rye, sp. nov. Typus\ Kalbarri National Park, Western Australia [precise locality withheld for conservation reasons], 24 October 1995, M.N. Lyons 2411 {holo\ PERTH 05002257; iso\ K, MEL). Scholtzia sp. Ajana (T. A. Halliday 137), in G. Paczkowska & A.R. Chapman, West. Austral. FI.: Descr. Cat. p. 401 (2000); Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Shrub 0.9-3 m high, width not recorded; flowering branchlets with peduncles borne at up to 20 consecutive nodes in a raceme-like arrangement, but usually at 5-10 nodes and more clustered, in both cases usually well below apex of branchlet. Leaves antrorse to patent. Petioles 0.5-\ . 1 mm long. Leaf blades broadly or very broadly obovate, 3.5-5.5 mm long, 3-5 mm wide, with a translucent margin up to c. 0.1 mm deep, entire; abaxial surface flat except for a slight narrow furrow or ridge along midvein, with lateral veins usually obscure, the oil glands in more than 3 rows on each side of midvein but inconspicuous. Peduncles 2-4 mm long, 0.4-0.6 mm wide, mostly 7-12-flowered; secondary axes up to 1 mm long. Bracts 0.6-1.6 mm long, caducous. Pedicels 0-1.2 mm long. Flowers 4-5.3 mm diam. Hypanthium 1.5-1.8 mm long, rugose. Sepals mostly broadly to depressed ovate, 0.5-1.0 mm long, the outer ones with base somewhat ridged; petaline margin 0.4-0.6 mm deep, entire. Petals 1.5-2.0 mm long, white or pale pink. Stamens 7-10, with 1-3 opposite each sepal. Longest filaments 0.2-0.4 mm long. Anthers 0.35-0.4 mm long. Ovary inferior, 2-locular; ovules 2 per loculus. Style 0.5-1.0 mm long; stigma capitate. Fruits largely inferior, 2.0-2.5 mm long, 1.6-1.8 mm wide; sepals erect; petals persistent. Seeds 1.3-1.8 mm long, 0.6-0.8 mm wide. Diagnostic features. Among species that have a 2-locular ovary with 2 ovules per loculus, S. bellairsiorum is distinguished by the following combination of characters: leaves broadly or very broadly obovate, with lateral veins usually obscure; peduncles 2-4 mm long; stamens 7-10; petals persistent in fruit. Selected specimens examined. [localities withheld for conservation reasons] 21 Oct. 1974, J.S Beard 7137 (PERTH); 10 Oct. 1995, D.R. Bellairs s.n. (PERTH); 12 Jan. 2005, A. Crawford 867 (PERTH); 19 Dec. 1968, H. Demarz 925 (PERTH) Distribution and habitat. Occurs in the eastern part of Kalbarri National Park (F igure 2A), on sandplain, often dominated by Banksia. Phenology. Flowers from September to December. Mature fruits recorded from November to January. Etymology. Named in honour of Don and Barbara Bellairs, who collected extensively in the Kalbarri area over their many years of residence there, and kept a local herbarium. They were the first to collect a number of rare species from Kalbarri National Park and nearby, including S. tenuissima Rye, and are the only collectors of Thryptomene pinfolia Rye & Trudgen. Conservation status. Priority Three under Conservation Codes for Western Australian Flora. Listed with this conservation status by Smith and Jones (2018) under the name S. sp. Ajana (T.A. Halliday 137). Affinities. Scholtzia bellairsiorum is closest in morphology to S. uberiflora F.Muell. and S. pentamera Rye, but those species have more obviously veined leaves, longer peduncles (4-15 mm long) and less persistent petals, and S. pentamera also differs in having only five stamens. B.L. Rye, Update to the taxonomy of Scholtzia 45 Figure 2. Distribution of Scholtzia species and subspecies. A- S. bellairsiorum (■), S. chapmanii (o), S. prostrata (•) and S. quindecim ( a); B - 5. brevistylis subsp. brevistylis (a), 5. brevistylis subsp. prowaka (a), unnamed variant of S. brevistylis (a), S. calcicola (•), S. cordata (o) and S. corrugata (■). Scholtzia brevistylis Rye, sp. nov. Typus: Yandanooka, Western Australia [precise locality withheld for conservation reasons], November 1998, R. Soullier 646 {holo\ PERTH 05447860; iso\ K, MEL). Shrub 0.6-2.5 m high, commonly 0.9-2 m wide, rather spindly; flowering branchlets commonly with peduncles borne at 4-15 consecutive nodes in a raceme-like arrangement. Leaves antrorse to patent. Petioles 0.4-0.8 mm long. Leaf blades obovate, 2^.5 mm long, 1.0-2.4 mm wide, with a clear-translucent margin up to c. 0.1 mm deep, entire; abaxial surface gradually raised towards centre, sometimes grooved along midvein, with lateral veins obscure, the oil glands usually in 2 main rows on each side of midvein, with 4-7 glands in innermost rows. Peduncles \-A mm long, 0.25- 0.3 mm wide, 3-7-flowered; secondary axes up to 0.5 mm long. Bracts 0.7-2.0 mm long, deciduous or caducous. Pedicels 0.2-0.6 mm long. Flowers 3-3.5 mm diam. Hypanthium 0.6-1.0 mm long, smooth to somewhat glandular-rugose. Sepals almost deflate to depressed ovate, 0.3-0.5 mm long, the outer ones often with a green ridge; petaline margin 0.25-0.4 mm deep, entire. Petals 1.2-1.5 mm long, white or pale pink. Stamens 4-6, usually 5, with 0-2 opposite each sepal. Longest filaments 0.2- 0.4 mm long. Anthers 0.2-0.25 mm long. Ovary inferior, 2-locular; ovules 2 per loculus. Style 0.3- 0.6 mm long; stigma capitate. Fruits 2/3-3/4 inferior, 1.3-1.5 mm long, 1.0-1.2 mm wide; sepals erect or incurved; petals deciduous. Seeds 1.0-1.3 mm long, 0.5-0.6 mm wide. Diagnostic features. Distinguished from all other named species of Scholtzia by the combination of a short style (0.3-0.6 mm long) and 4-6 stamens with 0-2 opposite each sepal. Other important characters: 46 Nuytsia Vol. 30 (2019) 3-7-flowered peduncles, the smooth to somewhat glandular-rugose (i.e. not pitted) hypanthium and 2 ovules per loculus. Distribution and habitat. Occurs from the Mingenew area south to the Nebroo Hill area and south¬ east to the Coorow area (Figure 2B). Etymology. From the Latin brevis (short) and -stylis (styled), as this species has a shorter style than any other member of the genus except for S. pentamera. Affinities. Scholtzia brevistylis is very distinctive, its aflinities uncertain. It is readily distinguished from the other particularly short-styled species, S. pentamera, for example in its narrower leaves, which are longer than broad, shorter peduncles and smaller flowers. Although S. brevistylis has two ovules per loculus, it is similar to a number of species with a single ovule per loculus in having one or two sepals with no stamen opposite them. Most flowers have five stamens in the sequence 2,0,1,2,0 around the circumference of the flower, i.e. with a maximum of two stamens opposite each sepal. Notes. Three groups of specimens are included here, from the northern, southern and western parts of the range respectively. The poorly known western variant is described informally below, while the other two variants are treated as subspecies brevistyla and prowaka. a. Scholtzia brevistylis Rye subsp. brevistylis Scholtzia sp. Yandanooka (R. Soullier 646), in G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. p. 402 (2000); Western Australian Herbarium, inF/oraR(3!5e, https://florabase.dpaw.wa.gov. au/ [accessed 2 March 2018]. Leaf blades 1-1.5 mm wide, with margins usually markedly recurved, at least at base. Flowering branchlets with peduncles borne at 5-11 consecutive nodes in a raceme-like arrangement 10-60 mm long, always with some over 20 mm long, usually well below the end of the branchlet; peduncles of the lowest node l-2(-3) mm long. Selected specimens examined. WES TERN AU S TRAEIA: [localities withheld for conservation reasons] 16 Nov. 1996, A. Carr 383 (PERTH); 10 Dec. 2015, A. Crawford ADC 2646 (PERTH); 2 Dec. 1999, S.J. Patrick 3358A (NSW, PERTH); Oct. 1997, R. Soullier 431 (PERTH). Distribution and habitat. Occurs in the Mingenew to Yandanooka area (Figure 2B), often on sandplain, sometimes on the margins of granite outcrops or associated with damp depressions. Phenology. Flowers from October to early December. Mature fruits recorded in November and December. Conservation status. Priority One under Conservation Codes for Western Australian Flora. Eisted with this conservation status by Smith and Jones (2018) under the name S. sp. Yandanooka (R. Soullier 646). b. Scholtzia brevistylis subsp. prowaka Rye Typus\ east of Carnamah, Western Australia [precise locality withheld for conservation reasons], 7 September 2006, M. Hislop, P. Aynsley & J. Borger MH 3650 {holo\ PERTH 07403267; iso\ K, MEE). B.L. Rye, Update to the taxonomy of Scholtzia 47 Scholtzia sp. Prowaka Springs (R. J. Cranfield & R Spencer 8083), in G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. p. 402 (2000); Western Australian Herbarium, in FloraBase, https:// florabase. dpaw.wa.gov.au/ [accessed 2 March 2018]. Leaf blades 1.3-1.6 mm wide, with margins fairly flat. Flowering branchlets with peduncles borne at 3-8 consecutive nodes in a short raceme-like arrangement up to 15 mm long, tending to be concentrated into a small zone towards the end of the branchlet; peduncles of the lowest node 2-A mm long. Diagnostic characters. Differs from subsp. brevistylis in usually having longer peduncles and leaves with fairly flat margins. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 16 Oct. 2006, J. Borger AK 1610-12 (CANB, NSW, PERTH); 24 June 2008, J. Borger IB 246-08 (CANB, NSW, PERTH); 19 Sep. 1991, R.J. Cranfield 8083 & P.J. Spencer (PERTH). Phenology. Flowers from late June to September, with mature fruits recorded in October. Distribution and habitat. Extends from north of Carnamah to south-east of Coorow (Figure 2B), on granite outcrops, also recorded on laterite. Etymology. Named after Prowaka Springs, as this subspecies was first collected near there. Conservation status. Priority Two under Conservation Codes for Western Australian Flora. Previously listed (Smith & Jones 2018) as Priority One under the name S. sp. Prowaka Springs (R.J. Cranfield 8083 & PJ. Spencer). Only four collections have been made of this taxon, but it is known from a nature reserve. Notes. Flowering branchlets often have peduncles borne at, or fairly close to, the apex of the branchlet, whereas in subsp. brevistylis and the unnamed eastern variant, the uppermost peduncles are usually well below the apex of each branchlet. c. Scholtzia brevistylis unnamed eastern variant Leaf blades 2.0-2.4 mm wide, with margins fairly flat. Flowering branchlets with peduncles borne at 5-14 consecutive nodes in a raceme-like arrangement 15-70 mm long, always with some over 20 mm long, usually well below the end of the branchlet; peduncles of the lowest node 1-2 mm long. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 17 Oct. 2007, J. Borger BR 1710-14 (PERTH); 17 Nov. 2009, J. Borger NR21 5 (PERTH); 10 Oct. 2002, S. Patrick (CANB, PERTH). Phenology. Flowers from October to December. Distribution and habitat. Restricted to a small area near Nebroo Nature Reserve (F igure 2B), apparently associated with damp habitats or on hillsides, recorded in sandstone, siltstone or ironstone gravel. Notes. The three specimens placed here have broader leaves than either of the named subspecies. They 48 Nuytsia Vol. 30 (2019) match subsp. brevistylis in their inflorescences but have fairly flat leaf margins as in subsp. prowaka. More collections are needed to assess the taxonomic status of this variant. Scholtzia calcicola Rye, sp. nov. Typus\ south-west of Dongara, Western Australia [precise locality withheld for conservation reasons], 9 December 2002, R.P. Hart 8401 {holo\ PERTH 06256449; iso\ CANB, K, MEL, NSW). Scholtzia sp. Dongara (R. Hart 8401), Western Australian Herbarium, in FloraBase, https ://florabase. dpaw.wa.gov.au/ [accessed 2 March 2018]. Shrub erect, dense, 0.2-2 m high, 0.3-1.2 m wide; flowering branchlets mostly with peduncles borne at 2-8 consecutive nodes, in a raceme-like arrangement or sometimes with flowers in a globular cluster. Leaves appressed or antrorse. Petioles 0.2-0.4 mm long. Leaf blades obovate, 1.8-2.5 mm long, (0.8-) 1.2-1.4 mm wide, entire, obtuse, not apiculate; abaxial surface flattened at the centre in basal 1/2-3/4 of its length and often with midrib ridged distally, with lateral veins obscure, the oil glands in 1 or 2 main rows on each side of midvein, with 3-6 glands in innermost rows. Peduncles 3-5 mm long, 0.2-0.35 mm wide, mostly 3-7-flowered; secondary axes up to 0.6 mm long. Bracts 0.6-0.8 mm long, deciduous. Pedicels absent or up to 0.3 mm long. Flowers 2.5-3.5 mm diam. Hypanthium 0.7- 0.9 mm long, wrinkled or reticulate-rugose. Sepals broadly to depressed ovate, 0.35-0.5 mm long, the base ridged; petaline margin 0.15-0.3 mm deep, entire. Petals 1.0-1.2 mm long, usually pale pink. Stamens commonly 5, with 0-2 opposite each sepal. Longest filaments 0.25-0.35 mm long. Anthers 0.25-0.3 mm long. inferior, 3-locular; ovules 1 per loculus. Style c. 0.6 mm long; stigma capitate. Fruits globular, 2/3-3/4 inferior, 1.3-1.35 mm long, 1.1-1.2 mm wide; sepals strongly incurved; petals deciduous. Seeds 1.0-1.1 mm long, c. 0.8 mm wide. Diagnostic features. Among species that have 1 ovule per loculus, S. calcicola is distinguished by the following combination of characters: leaves 0.8-1.4 mm wide; hypanthium wrinkled or reticulate- rugose, 0.7-0.9 mm long; petals 1.0-1.2 mm long. Other specimens examined. WESTERN AEISTRALIA: [localities withheld for conservation reasons] 11 Nov. 2014, S Ruoss SBR 020 (PERTH); 16 Oct. 2014,7: Stehbens & D. Panickar TS 003 (PERTH); 16 Oct. 2014, T Stehbens & D. Panickar TS 005 (PERTH); 26 Sep. 1984, R.T. Wills s.n. (PERTH^ 14 Oct. 2011, G. Zemunik25 (PERTH). Distribution and habitat. Recorded from near Dongara south to Drovers Cave National Park (Figure 2B), in heath on shallow sand over limestone. Phenology. Flowers from October to December. Etymology. From the Latin calcis (lime) and -cola (inhabitant), as the species is found in limestone habitats. Vernacular name. Tiny-flowered Scholtzia. Conservation status. Priority Two under Conservation Codes for Western Australian Flora. Previously listed (Smith & Jones 2018) as Priority One under the name S. sp. Dongara (R. Hart 8401), this species has recently been collected from a National Park and its known range extended to about 100 km. B.L. Rye, Update to the taxonomy of Scholtzia 49 Affinities. This belongs to a species group that has the sepals strongly incurved, both in flower and fruit. It shows greatest similarity to S. corrugata but that species occurs much further north and differs in having broader leaves and larger flowers with a longer hypanthium (see key). Scholtzia multiflora is closer in distribution to S. calcicola but has more numerous flowers and a 2-locular ovary. Co-occurring species. Recorded growing with a member of the S. obovata complex (G. Zemunik 24). Notes. Scholtzia calcicola has the smallest flowers in the genus. Its fruits are particularly delicate and dotted with somewhat glittering oil glands. ScholtziacapitataF.Muell exBenth., FI. Austral. 3: 69 (1867). Baeckeacapitata (F.Muell. exBenth.) F.MuelL, Census Austral. PI. 54(1883). Type: Murchison River, Western Australia, 1859-1863, A.F. Oldfields.n. (lecto: MEL2278625,^(ieB.L. Rye, Nuytsia 28: 161 (2017); isolecto: K 000357124, MEL 2278624). Scholtzia sp. Shark Bay (M.E. Trudgen 7429), in G. Paczkowska & A.R. Chapman, West. Austral. FI : Descr. Cat. p. 402 (2000); Western Australian Herbarium, in FloraBase, https://florabase.dpaw. wa.gov.au/ [accessed 2 March 2018]. Scholtzia sp. Yerina Springs (N. Hoyle 517), Western Australian Herbarium, in FloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Shrub 0.4-3 m high, 0.5^ m wide; flowering branchlets with peduncles mostly borne at 1-3 consecutive nodes, often in an umbel-like arrangement. Leaves widely antrorse or patent; foliar colleters (one on each side of the petiole of young leaves) Aliform, 0.35-0.6 mm long, white. Petioles 0.5-1 mm long. Leaf blades narrowly to broadly obovate, 2.5-5 mm long, 1.3-3.5 mm wide, usually not very thick; abaxial surface usually ridged along midvein for at least half the blade length, sometimes deep bluish green, entire; abaxial surface with lateral veins obscure, the oil glands densely covering the surface rather than in rows. Peduncles 8-18 mm long, 0.5-0.8 mm wide, usually 9-23-flowered; secondary axes up to 1.3 mm long. Basal bracts 1.8-2.3 mm long. Pedicels 0.5-1.0 mm long. Flowers 4.3- 4.8 mm diam. Hypanthium 1.1-1.4 mm long, densely glandular and pitted, each gland depressed below the surface and forming the full area of the pit. Sepals ovate to oblong or semicircular, commonly broadly ovate, (0.5-)0.8-1.3 mm long, thickened or ribbed and reddish at base; petaline margin 0.4- 0.8 mm deep, entire. Petals 1.5-1.8 mm long, pale to medium pink. Stamens 6-10, with 0-3 opposite each sepal. Longest filaments 0.2-0.35 mm long. Anthers c. 0.35 mm long. Ovary inferior, 2-locular; ovules 1 per loculus. Style usually 0.6-1 mm long; stigma capitate or peltate. Fruits 1/2-2/3 inferior, 2-2.5 mm long, c. 1.6 mm wide; sepals erect; petals deciduous or persistent. Seeds 1.2-1.4 mm long, c. 0.6 mm wide. Diagnostic features. Among species that have 1 ovule per loculus, S. capitata is distinguished by the following combination of characters: leaves densely glandular; peduncles 8-18 mm long, usually 9-23-flowered; hypanthium densely glandular and pitted, longer than wide; stigma capitate. Selected specimens examined. WESTERN AUSTRALIA: between Hamelin and Tamala, 16 Oct. 1974, J.S. Beard 7067 (PERTH); near the Murchison river on the River Rd track, 2.5 km N of Kalbarri, 5 Oct. 1990, S. Maley 3 (AD, CANB, PERTH); 6.7 km from Tamala-Useless Loop fork on road to Hamelin Pool, 18 Sep. 1974, B.L. Powell 74090 (PERTH); Meanarra Hill, Kalbarri National Park, 29 Oct. 1996, R. Schuh & G. Cassis 96-33 (PERTH). 50 Nuytsia Vol. 30 (2019) Distribution and habitat. Extends fromXamala Station south-south-east to near Yerina Springs (Figure 3A), in yellowish to reddish sandy soils. See below for a discussion of an outlying collection {F. Hort & J. Hort 3797) from the Perth suburb of Stratton. Conservation status. Not listed by Smith and Jones (2018); this species has most of its known populations protected within Kalbarri National Park. Chromosome number. n = \\,fide B.L. Rye, Austral. J. Bot. 27: 572 (1979) [as Scholtzia sp.]. Voucher: B.L. Powell 74090. Phenology. Flowers mainly from August to October. Fruits recorded from October and January. Etymology. From the Fatin capitatus (capitate, head-like), referring to the head-like arrangement of the flowers, which are actually in a condensed dichasial cyme. Vernacular name. Pom-pom Scholtzia. Affinities. This species and S. oleosa Rye are referred to here as the S. capitata complex. They have crowded oil glands on the leaves and also on the hypanthium, with the glands sunken into small pits on the hypanthium. See S. oleosa for the differences between the two species. Figure 3. Distribution of Scholtzia species. A - 5. capitata, with outlying southern collection (•), S. multiflora (a) and S. off. multflora ( a); B - S. denticulata (o), S. inaequalis (a) and S. peltigera (•). B.L. Rye, Update to the taxonomy of Scholtzia 51 Notes. When this species was lectotypified in Rye (2017), it comprised only the specimens that had been housed under the phrase name S. sp. Yerina Springs (N. Hoyle 517). Now the circumscription of S. capitata is enlarged to include specimens previously identified as S. sp. Shark Bay (M.E. Trudgen 7429). Scholtzia sp. Shark Bay (M.E. Trudgen 7429) is a variant known from only a few, mostly poor specimens from the north-western part of the range of the complex. It tends to have narrower leaves than the other variants, often with the blade narrowly obovate, but broader leaves can be present on the same specimens. Eeaf width varies considerably in the remainder of the complex, often with considerable variation on individual specimens that may occasionally include narrowly obovate leaves. Foliar colleters are more conspicuous in this species than in most other species that have one ovule per loculus, but are usually shed before the leaves mature. Similar filiform colleters are associated with the bracts and bracteoles; these are commonly noticeable, especially if they are left protruding from the top of peduncles after the bracts and fruits have been shed. The very isolated Stratton collection {F. Hort & J. Hort 3797: PERTH) occurs more than 350 km south of the known natural range of S. capitata. It also differs from the rest in having only 4-6 stamens per fiower. It may represent a naturalised occurrence since over 500 plants have been recorded in a dense population, in which case it could be a cultivated hybrid. Alternatively the specimen could be the sole representative of a new taxon. Scholtzia chapmanii Trudgen ex Rye, sp. nov. Typus\ 0.8 km E of Eake Indoon entrance along Eeeman-Eneabba road. Western Australia, 9 December 1992, R.J. Cranfield&P.J. Spencer 8705 {holo\ PERTH 02934256; iso\ AD, CANB, K, MEE, NSW). Scholtzia chapmanii Trudgen ms. Western Australian Herbarium, in FloraBase, https: //fiorabase. dpaw. wa.gov.au/ [accessed 2 March 2018]. Scholtzia sp. Winchester (C. Chapman s.n. PERTH 05625386), Western Australian Herbarium, in FloraBase, https://fiorabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Shrub ± prostrate, 0.1-0.3 m high, commonly 0.8-1.8 m wide, at least sometimes with multiple, close branches arising at ground level from a woody base; fiowering branchlets with peduncles borne at up to 20 consecutive nodes but usually at 3-12 nodes, often in a one-sided raceme-like arrangement. Leaves appressed or antrorse. Petioles absent or up to 0.4 mm long. Leaf blades narrowly obovate or oblong to linear in outline, 1.2-3 mm long, 0.5-1.0 mm wide, 0.4-0.7 mm thick, ciliolate or denticulate at first, the cilia up to 0.2 mm long; abaxial surface raised and somewhat fiattened at centre, with lateral veins obscure, the oil glands in 1 or sometimes 2 main rows on each side of midrib, with 3-6 glands in innermost rows. Peduncles 1-3.5 mm long, 0.2-0.35 mm wide, l(2)-fiowered; secondary axis (when present) c. 0.8 mm long. Bracteoles 0.7-2 mm long, deciduous. Pedicels 0.3-1.5 mm long. Flowers 5-8 mm diam. Hypanthium 1.0-1.3 mm long, 5-ribbed. Sepals broadly ovate to almost semicircular, usually very broadly ovate, 1.1-1.9 mm long, ridged on the herbaceous base, the ridge usually very prominent; scarious or petaline margin 0.5-0.7 mm deep, ± entire. Petals 2-3.5 mm long, white or pale pink, very erect in fruit. Stamens 15-20, in a continuous circle. Longest filaments 1.5-2 mm long. Anthers 0.3-0.5 mm long. Ovary c. 1/2 inferior, 2-locular; ovules 2 per loculus. Style 1.5-2.3 mm long; stigma capitate (possibly sometimes slightly peltate). Fruits c. 1/2 inferior, 1.6-2.3 mm long, 1.8- 2.2 mm wide, 5-ribbed; sepals erect; petals persistent. &^2\ (AD,BRI,K,NSW,PERTH); 17 Sep. 2002, E fff/Aow279(PERTH). Distribution and habitat. Associated with salt lakes in Yenyening Lakes Nature Reserve (Figure 4A), on white or yellow sand. Phenology. Flowers mainly from September to November. Fruits recorded in November and July. Etymology. From the Latin meridionalis (southern), as this subspecies occurs in the far south of the distribution of the genus. Conservation status. Recently listed as Priority Two under Conservation Codes for Western Australian Flora. This species is known from a conservation reserve. Notes. The sepals of this subspecies tend to be intermediate in length between those of subspecies halophila and mortlockensis but show some overlap with both of them. Subsp. meridionalis also B.L. Rye, Update to the taxonomy of Scholtzia 59 usually has fewer stamens (5-8 cf 7-12), and its mature style tends to be shorter (0.7-1.1 cf. 0.8- 1.4 mm long). However, good fruiting material is absent in subsp. meridionalis and this is needed to determine whether the mature style is sometimes close to the maximum length recorded for the other two subspecies. c. Scholtzia halophila subsp. mortlockensis Rye, subsp. nov. Typus\ Mortlock River, Western Australia, 18 November 1986, M.E. Trudgen 5427 {holo\ PERTH 06171877; iso\ CANB, K, MEL, NSW). Scholtzia sp. Duck Pool (M.E. Trudgen MET 5A21), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Leaf blades 2.5^.0 mm long, 2.0-3.6 mm wide. Peduncles 4-8 mm long. Sepals broadly or very broadly ovate, petal-like, 1.0-1.4 mm long. Petals 2.3-2.5 mm long, white to medium pink. Stamens (6-)8-l 1, with 0-3 opposite each sepal. Longestfilaments 0.4-0.6 mm long. Ovary usually 3-locular (rarely with a proportion of them 4-locular). Style 1.1-1.3 mm long. Diagnostic features. Subspecies mortlockensis is distinguished from the other two subspecies by the following combination of characters: sepals 1-1.4 mm long; stamens (6-)8-ll; ovary 3-locular in most flowers. Selected specimens examined.WESTERN A\J^TPJslAA\ [localities withheld for conservation reasons] 24 Jan. 1946, C.A. Gardner s.n. (BRI, CANB, NSW, PERTH); 26 Nov. 2003, B. Oversby BO 142 (PERTH); 23 Oct. 1967, PG. Wilson 6394 (AD, PERTH). Distribution. Occurs along the Mortlock River North, Mortlock River South and Mortlock River East, extending from north of Northam inland to near Cunderdin (Figure 4B). Etymology. Refers to its occurrence along all three branches of the Mortlock River, combining Mortlock with the Latin -ensis (native of). Conservation status. Recently listed as Priority Three under Conservation Codes for Western Australian Flora. Only known from 15 collections, some of which are historical records, in habitat that is susceptible to hydrological changes and salinity. Further survey is needed to check the conservation status of this taxon. Notes. This subspecies is intermediate in its average latitude and peduncle length between the other two subspecies. While it is readily separated in both morphology and distribution from subsp. meridionalis, it shows a greater approach in both these aspects to subsp. halophila. The length of its largest sepal is the character that best separates it from subsp. halophila (1-1.4 mm cf 0.5-0.8 mm long), but there may be some intermediate specimens occurring in the region where the ranges of the two taxa more or less meet. Subsp. mortlockensis is distinguished from subsp. meridionalis in having the ovary 3-locular rather than 2-locular in a majority of the flowers, and is the only subspecies known to rarely have a 4-locular ovary. At least one PERTH specimen, C. Howell 494, has a number of 4-locular ovaries, a character state not recorded elsewhere in the genus. 60 Nuytsia Vol. 30 (2019) Figure4. Distribution of5'c/?ofe/a species and subspecies. A-^. halophilasubs^. halophila{p)^S. halophilasvibs^. meridionalis (•) and S. uniflora (a); B - 5. halophila subsp. mortlockensis (a), S. longipedata subsp. longipedata (•) and S. longipedata subsp. procera (o). Scholtzia inaequalis Rye, sp. nov. Typus\ East Yuna Nature Reserve, Western Australia [precise locality withheld for conservation reasons], August 1985, T.F. Houston 617-2 {holo\ PERTH 03627721; iso\ CANB). Scholtzia sp. East Yuna (A.C. Bums 6), in G. Paczkowska & A.R. Chapman, West. Austral. FI : Descr. Cat. p. 402 (2000); Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 2 March 2018], Shrub 1-3 m high, 0.5-3 m wide; flowering branchlets with l-4(-6) pairs of peduncles, often with flowers in a dense cluster or short raceme-like arrangement. Leaves appressed or antrorse. Petioles 0.2-0.5 mm long. Leaf blades obovate or broadly obovate, 1.0-2.5 mm long, 1.0-1.7 mm wide, entire, often apiculate; adaxial surface flattened at the centre in basal 1/2-3/4 of its length and ridged distally, with lateral veins obscure, the oil glands in 1 main row of 3 or 4 large glands and 1 or 2 rows of smaller glands on each side of midvein. Peduncles 0.4-1.2(-1.7) mm long, 0.3-0.5 mm wide, 1-3-flowered; secondary axes ± absent. Bracts 0.8-2.0 mm long. Pedicels 0.2-0.8 mm long. Flowers 4-5 mm diam. Hypanthium 0.7-1.2 mm long, wrinkled-rugose. Sepals greatly differing in size and sometimes orientation, the outermost one broadly or very broadly ovate and 0.35-0.5 mm long and the innermost one broadly to depressed ovate and 0.8-1.4 mm long, the base ridged; petaline margin up to 0.7 mm deep, but much smaller on outermost sepals, entire. Petals 1.5-2.5 mm long, pale pink. Stamens 3-6, 0-2 opposite each sepal. Longest filaments 0.5-0.7 mm long. Anthers 0.25-0.3 mm B.L. Rye, Update to the taxonomy of Scholtzia 61 long. Ovary inferior, 2-locular in all or most flowers, rarely 3-locular; ovules 1 per loculus. Style 0.8- 1.5 mm long; stigma peltate or capitate. Fruits c. 2/3-inferior, c. 1.5 mm long, c. 1.6 mm wide; sepals fairly erect or (especially outer ones) tending to be incurved; petals deciduous. Seeds c. 1.3 mm long, c. 1.1 mm wide. Diagnostic features. Among species that have 1 ovule per loculus, S. inaequalis is distinguished by the following combination of characters: 1-3-flowered peduncles 0.4-1.2 mm long; petals 1.5- 2.5 mm long; stamens 3-6; outer sepals (incurved), usually much shorter than inner sepals (erect); hypanthium wrinkled-rugose. Selected specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons], 12-16 Oct. 19768, B.G. Muir 38 6.2 (PERTH); 15 Aug. 2006, A. Gunness AG 2988 (MEE, PERTH); 28 Aug. 2008, A. Gunness AG 3004 (CANB, NSW, PERTH); 31 July 2003, M.E. Trudgen MET 22021 (PERTH). Distribution and habitat. Extends from East Yuna Nature Reserve south to the Wicherina area (Figure 3B), recorded from varied habitats including the base of a low granite breakaway, lateritic and yellow sandplain areas, sometimes in open Eucalyptus woodlands. Phenology. Flowers mainly from June to October. Mature fruits recorded in August. Etymology. From the Eatin inaequalis (unequal), referring to the usually unequally sized sepals, with the innermost sepal tending to be more erect, as well as much larger than, the outermost sepal. Conservation status. Priority Two under Conservation Codes for Western Australian Flora. Fisted with this status by Smith and Jones (2018) under the name S. sp. East Yuna (A.C. Burns 6). This taxon occurs in a large nature reserve and one other reserve. Affinities. This species is very similar to S. truncata, which differs in having less difference in size between the inner and outer sepals, more numerous stamens and possibly a shorter style. It is also very similar to S. uniovulata, which differs in having a pitted hypanthium and often only a 1-locular ovary. Both of those species overlap in distribution with S. inequalis. Scholtzia inaequalis could also be confused with S. corrugata but that species differs in having longer, mostly 3-7-flowered peduncles and occurs further north. Notes. Two PERTH specimens that may belong to this species but which have more uniform-sized sepals than usual are P. Fairall 754 and S. Patrick SP 1760, both from the large East Yuna Flora Reserve (PERTH). Scholtzia lacmiata Rye, sp. nov. Typus\ near Eancelin, Western Australia, 1975, M.E. Trudgen 1516 {holo\ PERTH 06362168; iso\ AD, CANB, K, MEE, NSW). Scholtzia sp. Eancelin (M.E. Trudgen 1516), Western Australian Herbarium, in FloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. 62 Nuytsia Vol. 30 (2019) Prostrate or widely spreading shrub up to 0.4 m high, width not recorded, with young stems often markedly tuberculate; flowering branchlets with peduncles borne at 3-15 consecutive nodes, in a raceme-like arrangement. Leaves widely antrorse to retrorse. Petioles 0.3-0.7 mm long. Leaf blades elliptic to broadly ovate or cordate, 2.5^ mm long, 2-4 mm wide, markedly ciliate-laciniate, the longest cilia 0.4-0.8 mm long; abaxial surface raised only at the centre (near the midvein) and tending to be grooved along the midvein, with lateral veins obscure, the oil glands in several main rows on each side of midvein but inconspicuous. Peduncles 1.5-2.5 mm long, 0.4-0.7 mm wide, 3-7-flowered; secondary axes up to 0.6 mm long. Bracts 1.3-2 mm long, deciduous. Longest pedicels 0.4-0.8 mm long. Flowers 5-8 mm diam. Hypanthium 1.5-1.8 mm long, smooth or 5-ribbed. Sepals mostly very broadly to depressed ovate, 0.8-1.3 mm long, largely scarious, the herbaceous base somewhat ridged or almost smooth; petaline margin 0.5-0.8 mm deep, denticulate. Petals 2.5-3.5 mm long, pale pink. Stamens (10-)12-22, in a continuous circle. Longest filaments 1.7-2.5 mm long. Anthers 0.4-0.5 mm long. Ovary largely inferior, 2-locular; ovules 2 per loculus. Style 2.4-3.1 mm long; stigma capitate or peltate. Fruits 1/2-2/3 inferior, only sterile ones seen; sepals erect; petals deciduous. Diagnostic features. Distinguished by the following combination of characters: young stems often markedly tuberculate; leaf blades elliptic to broadly ovate or cordate, markedly ciliate-laciniate; stamens up to 22, in a continuous circle. Selected specimens examined. WESTERN AUSTRALIA: [localities withheld for conservation reasons] 8 Dec. 1992, E.A. Griffin 8404 (PERTH); 25 Jan. 1966, J.J. Havel H 251 (PERTH); 5 Oct. 1999, M.A. Langley & PM. Smith MAE 2173 (PERTH); 23 Nov. 2017, B. Morgan IOMH-12 (PERTH); 1 Dec. 1974, A.E. Orchard A261 (PERTH). Distribution and habitat. Extends from near Lancelin south to near Seabird and inland to Moore River National Park (Figure 5A), in shallow sand over limestone near the coast and in deep yellow sand further inland. Phenology. Flowers from November to January. Fruits recorded from December to January. Etymology. From the Latin laciniatus (laciniate), in reference to the irregularly incised leaf margins. Vernacular name. Ragged-leaved Scholtzia. Conservation status. Recently listed as Priority Two under Conservation Codes for Western Australian Flora, this species has a restricted range only c. 30 km long. Affinities. Similar to S. involucrata in many respects, including having flat leaves and peduncles usually 3-7-flowered, but differing in having markedly laciniate-ciliate margins to the leaves and in having the base of the sepals somewhat ridged. Notes. Scholtzia laciniata seems to be the only member of the genus to commonly have markedly tuberculate young stems. Judging from the most mature fruits examined, seeds probably tend to be 1.7-2 mm long. One PERTH specimen, B. Morgan IOMH-12, appeared to have most flowers with only 10-12 stamens. All other specimens examined had mostly 14-22 stamens per flower. B.L. Rye, Update to the taxonomy of Scholtzia 63 Figure 5. Distribution of Scholtzia species and subspecies. K-S. laciniata(A), S. pentamera suhsp.pentamera(*), S. pentamera subsp. collina (o) and S. thinicola (a); B - 5. oleosa (a) and S. uniovulata (o). Scholtzia longipedata Rye, sp. nov. Typus\ 4.3 km along firebreak, c. 8.3 km E of Kalbarri along Ajana-Kalbarri road. Western Australia, 30 August 2001, R. Davis 10004 {holo\ PERTH 05823854; iso\ MEE). Shrub 0.3-2.5 m high, 1-2.5 m wide; fiowering branchlets with peduncles borne at 1-5 consecutive nodes, varying in arrangement, sometimes combined into an umbel-like or short raceme-like arrangement but often discrete or more densely arranged. Leaves antrorse to patent, usually widely antrorse. Petioles 0.4-1.0 mm long. Leaf blades obovate or broadly obovate, 2.5^.5 mm long, 1.4-3.5 mm wide, entire, with incurved margins; abaxial surface gradually raised towards the centre, often indented or fiattened along the midvein but narrowed to a ridge distally, with thin translucent margins less than 0.1 mm deep, with lateral veins obscure, the oil glands in more than 2 rows on each side of midvein but inconspicuous. Peduncles 6-16 mm long, 0.4-0.8 mm wide, mostly 7-21-flowered; secondary axes up to 1.5 mm long. Basal bracts 1.4-2.5 mm long, caducous or deciduous. Pedicels 0.3-1.0 mm long. Flowers 3.5-5.5 mm diam. Hypanthium 1.0-1.6 mm long, somewhat rugose. Sepals broadly to depressed ovate, 0.3-0.75 mm long, smooth or with a basal ridge; petaline margin 0.2-0.3 mm deep, entire. Petals 1.3-2 mm long, white or pink. Stamens 6-9, with 1-3 opposite each sepal (commonly 8 in the arrangement 2,1,2,2,1). Longest filaments 0.4-0.6 mm long. Anthers 0.25-0.35 mm long. Ovary inferior, 2-locular; ovules 2 per loculus. Style 0.7-1.4 mm long. Fruits c. 3/4 inferior, commonly 1.9-2.3 mm long, 1.6-1.7 mm wide; sepals erect; petals persistent or deciduous. Seeds 1.2-1.6 mm long, 0.5-0.7 mm wide. 64 Nuytsia Vol. 30 (2019) Diagnostic features. Among species that have a 2-locular ovary with 2 ovules per loculus, S. longipedata is distinguished by the following combination of characters: leaf blades entire, obovate or broadly obovate; petioles 0.4-1 mm long; peduncles 7-16 mm long, mostly 7-21 -flowered; sepals 0.3-0.75 mm long; stamens 6-9, with 1-3 opposite each sepal. Distribution and habitat. Extends from the coast of the northern part of Kalbarri National Park south¬ east to East Yuna Reserve and Eradu (Figure 4B). Phenology. Flowers mainly from July to November but with one record from December. Mature fruits have been recorded from September to November. Etymology. From the Eatin longus (\ong),pes {-pedis', foot) and -atus (indicating possession or likeness), referring to the long peduncles in this species. Scholtzia spatulata has the longest peduncles in the genus, but also tends to have longer leaves, so the two taxa are similar in the degree to which the flowers are separated from the foliage. Another particularly long-pedunculate species is S. cordata. Co-occurring species. Recorded {A.G. Gunness AG 2382) growing upslope from, but apparently not intermixed with, S. denticulata (A.G. Gunness AG 2379). Affinities. All other species of Scholtzia differ in several characters from this species, making it diflicult to suggest which might be its closest relative. Notes. This species was first collected in 1958 ‘near the Murchison River’ by Charles Gardner, who recorded the plant height as ‘30 cm’ and flowering in early December. These details, if accurate, give the lowest plant height and latest flowering time recorded for the species. Two subspecies are recognised, occurring in the north and south of the range respectively. a. Scholtzia longipedata subsp. longipedata Scholtzia sp. Red Bluff (A. Gunness 2373), in G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. p. 402 (2000); Western Australian Herbarium, inE/oraR(3!5e, https://florabase.dpaw.wa.gov. au/ [accessed 2 March 2018]. Shrub 0.3-2.4 m high, 1.0-2.5 m wide. Leaf blades 2.5-4.5 mm long, 1.4-3.5 mm wide, not very thick. Peduncles 7-16 mm long. Sepals usually depressed ovate, 0.3-0.5 mm long, smooth or with a basal ridge; petaline margin 0.2-0.3 mm deep. Style 0.7-1.4 mm long; stigma peltate or capitate. Selected specimens examined. WESTERN AUSTRAEIA: Transect RB, Red Bluff, Kalbarri National Park, 9 Sep. 2003, D. & B. Bellairs 6336 (PERTH); Erriary Rd, 8.4 km N of the intersection with Binnu West Rd, 9 July 1997, R. Davis 3674 (PERTH); between Eookout and parking lot, Z Bend, Murchison River gorge, Kalbarri National Park, 2 Oct. 1991, W. Greuter 22482 (PERTH); Pot Alley Gorge, Kalbarri National Park, 26 Sep. 1974, G. Perry 322 (CANB, MEE, PERTH); 33.5 km E of the Meanarra Hill turnoff on the road to North West Coastal Hwy, Kalbarri National Park, 25 Sep. 2002, M.E. Trudgen 21708 (AD, BRI, CANB, DNA, NSW). Distribution and habitat. Occurs in Kalbarri National Park and just south of the park (Figure 4B), on the coast in sand over sandstone and in near-coastal areas with sand. B.L. Rye, Update to the taxonomy of Scholtzia 65 Conservation status. Not listed by Smith and Jones (2018); most of the range of this species is within a large national park. Notes. Specimens collected from along the coast are mostly 0.3-1.5 m high, apparently stunted from their exposed position, whereas those collected further inland are 1.2-2.5 m high. There is ample flowering material but better fruiting material is needed for this subspecies. The stigma is often peltate in subsp. longipedata but is always capitate in subsp. procera Rye. See under the latter for other differences. b. Scholtzia longipedata subsp. procera Rye, subsp. nov. Typus\ Binnu East Road, Western Australia [precise locality withheld for conservation reasons], 6 December 1993, M.E. Trudgen & M.R. Trudgen MET 12016 {holo\ PERTH 06171869; iso\ AD, BRI, CANB, K, NSW, MEE). Scholtzia sp. Binnu-Yuna (M.E. Trudgen 12016), Western Australian Herbarium, inFloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Scholtzia sp. Eradu (R.D. Royce 8016), in G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. p. 402 (2000); Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Shrub 1.2-3.5 m high, commonly 1-1.7 m wide. Leaf blades 2.0-3.3 mm long, 1.5-3.0 mm wide, up to c. 0.4 mm thick. Peduncles 6-11 mm long. Sepals broadly or very broadly ovate, (0.4-)0.5- 0.75 mm long, with herbaceous base distinctly ridged; petaline margin 0.25-0.5 mm deep. Style 0.5- 1.0 mm long; stigma capitate. Diagnostic features. Differs from subsp. longipedata in its usually thicker leaves and usually longer sepals that are more markedly ridged, and tends to be a taller plant with more persistent petals and a shorter style. Selected specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 19 Nov. 1968, A.C Burns 99N (PERTH); 24 Oct. 1992, F.A. Griffin 7516 (PERTH); 15 Aug. 2008, A. AG 2987 (PERTH); 24 0ct. 2001, YJ. 4082 (PERTH); 31 Oct. 1963,P.D. Royce 8016 (PERTH); 31 July 2003, M. Trudgen MET 22017 (PERTH); 28 Sep. 2007, M.F. Trudgen & M. Guest MET 22544 (PERTH). Distribution and habitat. Extends from East Yuna Reserve south-west to Eradu (Figure 4B), in varied habitats that are usually sandy, often with malices, Melaleuca or Allocasuarina. This subspecies may form a tall shrubland 2-3 m high. Etymology. From the Eatin procerus (tall, long), as this is one of the tallest members of the genus, up to 3.5 m high. Conservation status. Priority Three under Conservation Codes for Western Australian Flora. Previously listed as Priority Two by Smith and Jones (2018) under the name Scholtzia sp. Eradu (R.D. Royce 66 Nuytsia Vol. 30 (2019) 8016) but not listed under the name Scholtzia sp. Binnu-Yuna (M.E. Trudgen 12016). This species is known from two nature reserves. Notes. The phrase names S. sp. Eradu and S. sp. Binnu-Yuna have been applied to western and eastern specimens respectively, with all of the former in fruit and the latter mainly in bud or flower. Western specimens tend to have shorter sepals and styles than the eastern ones, but there is considerable overlap in the measurements of these characters. Scholtzia multiflora Rye, sp. nov. Typus\ Burma Road, Western Australia [precise locality withheld for conservation reasons], 1 August 2003, M. Trudgen MET 22022 {holo\ PERTH 08209022; iso\ CANB, K, MEE, NSW). Scholtzia sp. Kojarena (A.M. Ashby 1904), Western Australian Herbarium, in FloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Shrub commonly 1.6-2.0 m high, recorded as 1.5-2.7 m wide; flowering branchlets mostly with peduncles at 5^2 consecutive nodes in a raceme-like arrangement. Leaves widely antrorse or patent. Petioles 0.2-0.4 mm long. Leaf blades ± very broadly obovate, 1.7-3.3 mm long, 2.3-3.0 mm wide, entire, obtuse, not apiculate; abaxial surface flattened at the centre in basal 1/2-3/4 of its length and with midrib ridged distally, with lateral veins obscure, the oil glands usually in 3 or more main rows on each side of midvein, with 4 or 5 glands in innermost rows. Peduncles 5-7 mm long, 0.2-0.3 mm wide, 3-15-flowered; secondary axes 0.4-0.7 mm long. BractsO.A-\.5 mm long. Pedicels absent or up to 0.3 mm long. Flowers 3.5^.5 mm diam. Hypanthium 0.7-1.1 mm long, reticulate-rugose. Sepals mostly broadly or very broadly ovate, 0.4-0.5 mm long, smooth or ridged at extreme base, largely petaline, entire. Petals 1.4-1.6 mm long, white or pale pink. Stamens 4-7, with 0-2 opposite each sepal. Longest filaments c. 0.5 mm long. Anthers c. 0.2 mm long. Ovary inferior, 2-locular; ovules 1 per loculus. Style 0.6-0.7 mm long; stigma capitate or peltate. Fruits c. 2/3 inferior, c. 1.3 mm long, c. 1.3 mm wide; sepals incurved; petals deciduous. Seeds 0.65-1.1 mm long, 0.6-0.7 mm wide. Diagnostic features. Among species that have 1 ovule per loculus, S. multifiora is distinguished by the following combination of characters: leaves wider than long; peduncles many-flowered 5-7 mm long, borne at usually 5-42 consecutive nodes; hypanthium reticulate-rugose; stamens 4-7. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 14Aug. \966,A.M. Ashby 1904 (PERTH); 15Aug. 2008,A. GunnessAG299\ (PERTH); 8 Sep. 1998, S. Patrick 3007 (PERTH); Aug. 1924. G.L. Sutton s.n. (PERTH). Distribution and habitat. Recorded from Kojarena south-east to east of Walkaway (Figure 3 A), one record from a lateritic ridge but also recorded from a sandy swale. An outlying specimen from about 100 km to the south-east near Winchester, may also belong to this species (see Notes section below). Phenology. Flowers recorded from July to September and mature fruits in August. Etymology. From the Eatin multi- (many-) and -fiorus (-flowered). This species is given the epithet multiflora because it has the largest number of consecutive nodes bearing flowers so far recorded in the genus, and often has numerous flowers on each peduncle. B.L. Rye, Update to the taxonomy of Scholtzia 67 Conservation status. Priority One under Conservation Codes for Western Australian Flora. Listed with this status by Smith and Jones (2018) under the name S. sp. Kojarena (A.M. Ashby 1904). Affinities. Of the currently named species, S. calcicola and S. corrugata are perhaps the closest in their morphology, but S. multijiora differs from both in having widely spreading, depressed-obovate leaves with oil glands in 3 or more main rows on each side of midvein, more numerous peduncles on the flowering branchlets and a 2-locular ovary. Its fruit also appears to differ in being wider than broad and less glandular, but this requires confirmation with better fruiting material. All three species have sepals that are usually strongly incurved in flower as well as in fruit. Notes. A single, atypical specimen {E. Wittwer 813: PERTH) from the Winchester area (see Figure 3A) is currently determined as S. off. multijiora. This specimen might prove to be sufliciently distinctive to be treated as a subspecies of S. multijiora or a separate species but is too poorly known to assess its taxonomic status. The Winchester collection differs in having shorter peduncles (2.0- 2.5 mm cf. 5-7 mm long) with only 1-3 flowers {cf. up to 15 flowers), but as it is in bud with only a few flowers opened, the possibility that some peduncles produce more than three flowers in more mature inflorescences cannot be completely ruled out. Flower colour is recorded as pink, but that may be because the inflorescences are still in bud. E. Wittwer 813 also differs in having smaller leaves, commonly only three stamens per flower and larger petals (c. 1.7 mm long cf. 1.4-1.6 mm long). More material is needed to determine whether these differences are sufliciently reliable to warrant formal recognition of more than one entity. Scholtzia oleosa Rye, sp. nov. Typus\ north of Northampton, Western Australia [precise locality withheld for conservation reasons], 23 September 2002, M.E. Trudgen MET 21676 {holo\ PERTH 06361005; iso\ CANB, K, MEE, NSW). Scholtzia sp. Eurardy (J.S. Beard 6886), in G. Paczkowska & A.R. Chapman, West. Austral. El: Descr. Cat. p. 402 (2000); Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Shrub 0.4-3 m high, 0.5-4 m wide; flowering branchlets with peduncles borne at 1^ consecutive nodes, in umbel-like or other kinds of arrangements. Leaves widely antrorse or patent. Petioles 0.3- 0.6 mm long. Leaf blades obovate to depressed obovate (usually broadly or very broadly obovate), 0.5- 2.5 long, 1.2-1.7 mm wide, often 0.5-0.7 mm thick at first, entire; abaxial surface commonly raised and rounded over the full surface, but sometimes distinctly thinner on each side of the midvein, which is often grooved in the basal half but may be ridged close to the apex, with lateral veins obscure, the oil glands densely covering the surface rather than in rows. Peduncles 4-14.5 mm long, 0.5-0.8 mm wide, usually 3-14-flowered; secondary axes up to 1.5 mm long. Basal bracts 1.5-1.8 mm long. Pedicels 0.3-0.8 mm long. Flowers 4.5-5.5 mm diam. Hypanthium 1.1-1.3 mm long, rugose-pitted (the numerous pits each corresponding to a sunken gland). Sepals ovate to oblong or semicircular, commonly broadly ovate, (0.5-)0.8-1.3 mm long, thickened or ribbed and reddish at base; petaline margin 0.4-0.8 mm deep, entire. Petals 1.6-2.3 mm long, pale to medium pink. Stamens 7-10, with 0-3 opposite each sepal. Longestfilaments c. 0.2 mm long. Anthers 0.35-0.4 mm long. Ovary inferior, 2- or 3-locular; ovules 1 per loculus. Style 0.6-0.8 mm long; stigma capitate. Fruits 1/2-2/3 inferior, 1.5-1.6 mm long, 1.5-1.6 mm wide; sepals erect; petals deciduous or persistent. Seeds 1.0-1.4 mm long, c. 0.55 mm wide. Nuytsia Vol. 30 (2019) Diagnostic features. Among species that have 1 ovule per loculus, S. oleosa is distinguished by the following combination of characters: leaves densely glandular; hypanthium pitted, ± as wide as long; stigma capitate. Selected specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 21 Oct. 1973, J.5. Beard (PERTH, AD, CANB); 3 Dec. 2004, A. Crawford ADC 809 (PERTH); 13 Sep. 2001, R. Davis 10044 (PERTH): 2 Oct. 1998, N. Gibson 4143 (PERTH); 26 Oct. 2006, S. Robinson SR 29 (PERTH); 25 Sep. 2002, M.E. Trudgen MET 21713 (PERTH); 29 Aug. 2003, Wildflower Society ofWA EURA 39 (AD, BRI, PERTH); 3 Oct. 2003, Wildflower Society of WA EURA 38 (PERTH). Distribution and habitat. Extends from Cobum Station south to near Binnu (Figure 5B), in yellowish to reddish sandy soils. Phenology. Flowers from August to October. Fruits recorded from October and December. Etymology. From the Eatin oleum (oil) and -osus (abounding in), referring to the numerous oil glands on the leaves and hypanthium. Conservation status. Fisted by Smith and Jones (2018) as Priority Two under the name S. sp. Eurardy (J.S. Beard 6886). This species has many more collections than when it was previously assessed for conservation status so should probably be reduced in priority. Co-occurring species. Scholtzia oleosa {M.E. Trudgen MET 22151) has been recorded growing with a member of the S. obovata complex {M.E. Trudgen MET 22153), with both species adjacent to a patch of vegetation where S. tenuissima {M.E. Trudgen MET 22154) occurred. Affinities. This species belongs to the S. capitata complex, in which there are crowded oil glands on the leaves and also on the hypanthium, where the glands are sunken into pits. Scholtzia oleosa differs from S. capitata in its less elongated hypanthium, usually shorter leaves, and usually shorter (4-14.5 mm cf. 8-18 mm) and fewer-flowered peduncles (3-14-flowered cf. usually 9-23-flowered). It also differs in having the ovary commonly 3-locular {cf. consistently 2-locular). As the two taxa overlap in range and flowering time, they may well have the opportunity to hybridise, which could account for one or two specimens that are diflicult to identify. Notes. Specimens from north of Eurardy (e.g. F. Lullfitz E 3197) have a consistently 2-locular ovary and tend to have small sepals whereas most specimens from Eurardy southwards have a 3-locular ovary. Scholtzia peltigera Rye, sp. nov. Typus: Eurardy Station, Western Australia [precise locality withheld for conservation reasons], 1 November 2016, B. Parkhurst BP 1 {holo\ PERTH 08986533; iso\ CANB, K, MEE, NSW). Scholtzia sp. Bungabandi (M. Quicke EURA48), Western Australian Herbarium, inFloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Shrub 0.5-2.0 m high, width not recorded, single-stemmed at base; flowering branchlets with peduncles borne at 1-5 consecutive nodes, sometimes with their flowers combined into a dense cluster. Leaves B.L. Rye, Update to the taxonomy of Scholtzia 69 widely antrorse to patent, sessile, with a broad, stem-clasping base. Leaves broadly to depressed ovate in outline, 1.3-1.8 mm long, 1.5-2.5 mm wide, 0.5-1.1 mm thick, with a clear-translucent margin up to 0.15 mm deep or rarely slightly deeper, entire; abaxial surface raised throughout, rounded, with lateral veins obscure, the oil glands in 3 or more rows on each side of midvein, usually with 3-5 large glands in innermost rows. Peduncles 2-8 mm long, 0.3-0.5 mm wide, mostly 3-12-flowered; secondary axes up to 0.4 mm long. Basal bracts caducous or deciduous, 1.2-1.5 mm long. Pedicels 0.2-0.7 mm long. Flowers 4.0-5.5 mm diam. Hypanthium 0.8-1.3 mm long, with prominent glands, becoming smooth in fruit. Sepals entire; outer ones almost triangular to depressed ovate, short, slightly ridged at base, with a scarious margin; inner ones mostly broadly ovate, 1.0-1.3 mm long, largely or fully scarious but with base tending to be glandular and reddish. Petals 1.7-2.0 mm long, white or pale pink. Stamens 7-10, with 1-3 opposite each sepal. Longest filaments 0.5-0.6 mm long. Anthers 0.25- 0.3 mm long. Ovary inferior, 2-locular; ovules 2 per loculus. Style 0.5-0.7 mm long; stigma capitate. Fruits c. 1/2 inferior, 1.6-1.7 mm long, c. 1.4 mm wide; sepals spreading or recurved; petals persistent. Seeds c. 1.3 mm long, 0.5-0.6 mm wide. Diagnosticfeatures. Unique in having stem-clasping, sessile leaves. Other important characters: leaves broader than long; peduncles 2-8 mm long, mostly 3-12-flowered; ovary 2-locular, with 2 ovules per loculus. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 24 Nov. 2004, M. Quiche EURA 48 (BRI, NSW, PERTH); 8 Sep. 2016, B.L. Rye, J. Norman & B. Parkhurst BLR 290166 (PERTH); 23 Aug. 2015 Wildfiower Society ofWA CREEK NORTH 2/22 (PERTH). Distribution and habitat. Known only from Eurardy Station (Figure 3B), mostly collected from a site on yellow sand, in a slight hollow well above a usually dry creek, in low open Eucalyptus woodland OYQY Acacia-Scholtzia tall shrubs. Phenology. Flowers recorded from September to November, and mature fruits in late November. Etymology. From the Eatin pelta (small, crescent-shaped shield) and -ger (bearing), referring to the presence of sessile, somewhat stem-clasping, leaves along the stems. Each leaf has three surfaces, the broad base that shallowly clasps the stem and is attached near its centre to the stem, the broadly crescent-shaped abaxial surface, and the smaller, incurved adaxial surface. Vernacular name. Eurardy Scholtzia. Conservation status. Priority One under Conservation Codes for Western Australian Flora. Fisted with this priority by Smith and Jones (2018) under the name S. sp. Bungabandi (M. Quicke EURA 48), S. peltigera has a very restricted distribution. Co-occurring species. There is one record {B.L. Rye, J. Norman & B. Parkhurst BER 290166) of S. peltigera growing intermixed with S. truncata {B.L. Rye, J. Norman & B. Parkhurst BER 290165), but only the latter was in flower. Affinities. Scholtzia peltigera is readily distinguished from all other members of the genus by its sessile, stem-clasping leaves, and it is not clear from its morphology which of the other species is likely to be its closest relative. 70 Nuytsia Vol. 30 (2019) Notes. Although narrow translucent margins are found on the leaves of a number of Scholtzia species, they tend to be more noticeable in S. peltigera, perhaps due to the contrast with the remainder of its very short, thick leaves. Scholtzia pentamera Rye, sp. nov. Typus\ 7 km N along Yerina Springs Road from Port Gregory Road, Western Australia, 15 August 1996, E. Holland & K. Kershaw EH 1539 {holo\ PERTH 04506111; iso\ K, MEE, NSW). Shrub 0.6-2.5(-3) m high, commonly 1-2.5 m wide; flowering branchlets mostly with peduncles borne at 8-23 consecutive nodes. Leaves widely antrorse to patent. Petioles 0.1-0.7 mm long. Leaf blades orbicular-cordate, 2.5-5 mm long, 3-6 mm wide, with a translucent margin less than 0.1 mm deep, entire; abaxial surface level except for a narrow raised midvein, usually with 4-6 lateral veins clearly visible on each side of midvein, the oil glands in 4 or more rows on each side of midvein, mostly with 5-7 glands in innermost rows. Peduncles 4-13 mm long, 0.4-0.6 mm wide, mostly 3-15-flowered; secondary axes up to 1.5 mm long. Bracts 1.3-3 mm long, deciduous. Pedicels 0-1.2 mm long. Flowers 3.5-5.0 mm diam. Hypanthium 1.5-2.0 mm long, wrinkled-ribbed or fairly smooth. Sepals usually broadly ovate to depressed elliptic, 0.5-0.9 mm long, almost fully scarious but outer ones often narrowly ridged or with a somewhat ridged green base; petaline margin 0.5-0.8 mm deep, entire. Petals 1.3-2 mm long, white or pink, deciduous in fruit. Stamens 5, with 1 opposite each sepal. Longest filaments 0.25-0.4 mm long. Anthers 0.25-0.35 mm long. Ovary inferior, 1- or 2-locular; ovules 2 per loculus. Style 0.3-0.4 mm long; stigma capitate. Fruits inferior, 1.5-1.8 mm long, c. 0.8 mm wide; sepals erect; petals deciduous. Seeds c. 1.2 mm long, c. 0.65 mm wide. Diagnosticfeatures. Differs fromY bellairsiorum andY uberiflora in having smaller flowers, including a shorter style, and only five stamens. Distribution and habitat. Extends from Aj ana south to and near Greenough (Figure 5A). Phenology. Flowers from July to November. Etymology. From the Greek penta- (five-) and -merus (refers to parts or their numbers) as the constant stamen number of five per flower in this species matches the number of sepals and petals. Vernacular name. Constant Scholtzia. This is the only species of Scholtzia known to have a constant stamen number, rather than having the number vary between flowers. Affinities. Very similartoY uberiflora oyerall, for example in having leaves often with multiple spreading veins visible, and both taxa are unusual in having long peduncles borne at many consecutive nodes along the stems, as other species with long peduncles (sometimes exceeding 10 mm) have them at a maximum of 6 nodes and mostly at 1^ nodes. Scholtzia uberiflora differs in having smaller bracts but usually larger flowers, 6-10 stamens per flower, with 1-3 opposite each sepal, a style 0.45-0.8 mm long, and in always having a 2-locular ovary. Notes. Two allopatric subspecies are recognised; these could be treated as separate species but are certainly more similar to one another than either is to S. uberiflora. B.L. Rye, Update to the taxonomy of Scholtzia 71 a. Scholtzia pentamera subsp. collina Rye, subsp. nov. Typus\ Howatharra,WestemAustralia [precise locality withheld for conservation reasons], 13 September 1977, A.S. George 14878 {holo\ PERTH 04062647; iso\ CANB, K). Bracts 2-3 mm long. Ovary markedly expanded distally, 2-locular. Diagnostic features. Distinguished from the other subspecies by its longer bracts and 2-locular ovary. Selected specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 3 July 1966, A.C. Burns 8 (PERTH); 3 July 1966, A.C. Burns 9 (PERTH); 26 Aug. 1983, R.J. Cranfield 297 \ (PERTH). Distribution and habitat. Occurs in Moresby Range (Figure 5A), the habitat recorded as ‘gravelly soil’ or ‘lateritic soil’. Etymology. From the Eatin collinus (pertaining to low hills), as this subspecies is restricted to a range of hills. Conservation status. To be listed as Priority Two under Conservation Codes for Western Australian Flora (M. Smith, pers. comm.). Probably restricted to a small area within a nature reserve. Notes. Subsp. collina is like the related species S. uberiflora in its ovary shape and loculus number, but is otherwise much more like S. pentamera subsp. pentamera. All three taxa differ in bract size. b. Scholtzia pentamera Rye subsp. pentamera Scholtzia sp. Northampton (A. Strid 20714), in G. Paczkowska & A.R. Chapman, West. Austral. FI : Descr. Cat. p. 402 (2000); Western Australian Herbarium, inFloraBase, https://florabase.dpaw.wa.gov. au/ [accessed 2 March 2018]. Bracts 1.3-1.8 mm long. tubular, 1-locular. Selected specimens examined. WESTERN AUSTRAEIA: S of Ogilvie, 3 Sep. 1947, N.T. Burbidge 2158 (PERTH); Bishops Gully Rd, 4.3 km from Port Gregory Rd, 8 July 1997, R. Davis 3624 (PERTH); private property, Rob Rd, slope above Yarda Gully Creek, W of homestead, 30 May 2008, A. Gunness AG 3022 (PERTH); W side of Ogilvie Rd W, close to Hutt River crossing, W of Binnu, 25 July 2008, M. Hislop 3783 (PERTH); 7 km N along Yerina Springs Rd from Port Gregory Rd, 15 Aug. 1996, E. Holland 1539 & K. Kershaw (MEE, NSW, PERTH); Northwest Coastal Hwy, 8 km S of junction with Ogilvie East Rd and 9.5 km S of crossing of the Hutt River, 11 Sep. 1999, J. W. Horn 2400 (AD, PERTH); on dunes near Flat Rocks, SE of Greenough, 4 Oct. 1972, S. Paust 1222 (PERTH); 9.5 km N of Northampton along North West Coastal Hwy, 7 Oct. 1982, A. Strid 20714 (PERTH). Diagnostic features. Distinguished from the other subspecies in having a 1-locular, tubular ovary. Distribution and habitat. Mostly collected between Ajana and Northampton, but with one outlying specimen from Flat Rocks Beach, near Greenough (Figure 5A), in varied habitats including yellow sand, the outlier recorded from sand dunes. 72 Nuytsia Vol. 30 (2019) Conservation status. Not listed by Smith and Jones (2018) but does not appear to be known from any nature reserves so may need to be surveyed. Notes. The uni-locular ovary is a rarity in Scholtzia, otherwise being known only in S. uniovulata, Scholtzia pentamera subsp. pentamera is unique in having a tubular ovary. Scholtzia prostrata Rye, sp. nov. Typus\ Burma Rd, Western Australia [precise locality withheld for conservation reasons], 13 October 1969, A.C. Burns 138 {holo\ PERTH 06165192; iso\ CANB, MEL). Scholtzia sp. Burma Road (A.C. Burns 138), Western Australian Herbarium, in FloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Prostrate shrub c. 0.1 m high, commonly 0.3-1.0 m wide, at least sometimes with multiple, close branches arising at ground level from a woody base; flowering branchlets with peduncles borne at up to 35 consecutive nodes, usually in a one-sided raceme-like arrangement. Leaves antrorse to patent. Petioles 0.3-0.5 mm long. Leafhlades narrowly obovate to elliptic, 2.5^.5 mm long, 0.8-2 mm wide, with a midrib commonly 0.3-0.5 mm thick but remainder thin, ciliolate or ciliate-laciniate, the longest cilia 0.2-0.3 mm long; abaxial surface raised along central part, often with a groove along midvein, with lateral veins obscure, the oil glands inconspicuous or in usually 2 or 3 main rows on each side of midvein, with 3-6 glands in main rows. Peduncles 1.5-3 mm long, 0.25-0.4 mm wide, 1-flowered. Bracteoles 1.1-1.5 mm long, deciduous or persistent. Pedicels 0.2-0.7 mm long. Flowers 5-8 mm diam. Hypanthium 0.8-1.0 mm long, finely rugose. Sepals broadly ovate to almost semicircular, usually very broadly ovate, 0.7-1.2 mm long, the herbaceous base ridged; petaline margin 0.6-0.8 mm deep, denticulate-ciliolate or dentate. Petals 2.5-3.5 mm long, white or pale pink. Stamens 11-17, in a continuous circle. Longest filaments 1.5-2.0 mm long. Anthers 0.3-0.35 mm long. Ovary c. 1/2 inferior, 2-locular; ovules 2 per loculus. Style 1.5-2.3 mm long; stigma capitate. Fruits c. 1/2 inferior, probably 1.5- 1.8 mm long, not seen at maturity; sepals erect; petals persistent. Diagnostic features. Among the species that have 2 ovules per loculus and mostly to consistently 1 -flowered peduncles, S. prostrata is distinguished by the following combination of characters: peduncles 1.5- 3 mm long; hypanthium not ribbed; sepals ridged at base; stamens 11-17, in a continuous circle; style 1.5-2.3 mm long. Selected specimens examined. WESTERN AUSTRALIA: [localities withheld for conservation reasons] 21 Oct. 1963, A.C. Burns 3 (NSW, PERTH); 3 Oct. 2007, A. Crawford ADC 1392 (PERTH); 23 Oct. \9n,G.J.Keighery&N. Gibson3Q)\\ (PERTH); 22Mm.2m\,S.J. Patrick3im &A. Chant (PERTH); 24 Oct. 2001, S.J. Patrick AOll (PERTH). Distribution and habitat. Occurs between Ambania and Strawberry, east of Walkaway (Figure 2A), in sand, often over laterite, in heath, often with emergent banksias or malices. Phenology. Flowers from October to December. Conservation status. Recently listed as Priority Three under Conservation Codes for Western Australian Flora. The species is known from at least one nature reserve. B.L. Rye, Update to the taxonomy of Scholtzia 73 Etymology. From the Latin prostratus (prostrate, lying flat along the ground), in reference to the habit of this species. Several apparently closely related species also tend to be prostrate but the habit is possibly most extreme in S. prostrata, with the species reported to be largely hidden {S. Patrick 4097) by other heath species. Vernacular name. Creeping Scholtzia. Co-occurring species. A possible hybrid {S. Patrick 4079 PERTH 08989702) was collected together with S. prostrata (S. Patrick 4079, PERTH 05947561). The putative hybrid is similar to S. prostrata but shows two significant differences, as it has up to three flowers per peduncle and fewer stamens arranged in antisepalous groups. These differences would be consistent with the other parent species being either S. ciliata or S. obovata, both of which are common in the general area and may well co¬ occur with S. pro strata. Affinities. This is the northernmost member of the S. chapmanii complex and has the fewest stamens on average. Scholtzia prostrata is most similar to S. chapmanii but differs in that its usually shorter hypanthium lacks the antisepalous ribs that are usually obvious in S. chapmanii. Also, its leaves tend to be broader and more spreading, and its sepals tend to be less prominently ridged at the base. Notes. Possibly this taxon should be treated as a subspecies of S. chapmanii, but since there are multiple minor differences between them it seems better to treat them as distinct species. The two taxa appear to be geographically separated by a distance of at least 50 km. As in S. chapmanii, there is just one specimen of S. prostrata showing that the base of the plant may have multiple stems arising from it at about ground level. In this case, two plants mounted on A.C. Burns 138, are complete except for their roots. There are no specimens with mature fertile fruits. Scholtzia quindecim Rye, sp. nov. Typus\ north ofWatheroo, Western Australia [precise locality withheld for conservation reasons], 5 October 1984, J.D. Briggs 1721 {holo\ PERTH 03628191; iso\ AD, CANB). Scholtzia sp. Gunyidi (J.D. Briggs 1721), in G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. p. 402 (2000); Western Australian Herbarium, inFloraBase, https://florabase.dpaw.wa.gov. au/ [accessed 2 March 2018]. Prostrate shrub usually 0.05-0.15m high, 0.6-2.5 m wide; flowering branchlets mostly with peduncles borne at 4-17 consecutive nodes, usually in a one-sided raceme-like arrangement. Leaves antrorse to patent. Petioles 0.3-0.5 mm long. Leaf blades narrowly obovate to narrowly oblong, 3.5-5 mm long, 0.5-1.1 mm wide, up to 0.5 mm thick, entire or denticulate-ciliolate; abaxial surface raised along the midvein, the raised part rounded, with lateral veins obscure, the oil glands usually in 2 rows on each side of midvein, with 4-8 glands in innermost rows but not obvious. Peduncles 3-7 mm long, 0.25-0.5 mm wide, nearly always 1-flowered. Bracteoles 1.1-2.2 mm long, deciduous or persistent. Pedicels 0.4-0.8 mm long. Flowers 6-8 mm diam. Hypanthium 0.9-1.1 mm long, finely rugose. Sepals broadly ovate to almost semicircular, usually very broadly ovate, 1.1-1.9 mm long, the herbaceous base thickened or ridged; petaline margin 0.7-1.2 mm deep, entire. Petals 3)-4 mm long, white or pale pink. Stamens 13-18, in a continuous circle. Longest filaments usually 2-2.5 mm long. Anthers 74 Nuytsia Vol. 30 (2019) 0.4-0.5 mm long. Ovary c. 1/2 inferior, 2-locular; ovules 2 per loculus. Style 2.5-3.4 mm long; stigma ± capitate. Fruits c. 1/2 inferior, only sterile ones seen; sepals erect; petals probably persistent. Diagnostic features. Among the species with two ovules per loculus and mostly to consistently 1-flowered peduncles, S. quindecim is distinguished by having peduncles 3-7 mm long, an unribbed hypanthium and a style 2.5-3.4 mm long. Selected specimens examined. [localities withheld for conservation reasons] 27 Oct. 1992, E.A. Griffin 7716 (PERTH); Reserve 21788,20 Nov. 1992, S.J. Patrick 1450 (PERTH); 9 Nov. 1961, Mrs Strickland s.n. (PERTH). Distribution and habitat. Occurs from Gunyidi to Dalwallinu (Figure 2A), in yellow or grey sand, recorded in sites with low woodlands or malices. Phenology. Flowers from September to November. Etymology. From the Eatin quindecim (fifteen), referring to the number of stamens, since this species has an average of about 15 stamens per flower. Vernacular name. Gunyidi Scholtzia. Conservation status. Priority Two under Conservation Codes for Western Australian Flora. Fisted with this priority by Smith and Jones (2018) under the name S. sp. Gunyidi (J.D. Briggs 1721). Affinities. This is the most inland member of the S. chapmanii complex and the only one with more or less entire leaves. It has the longest peduncles within the complex. It may also have longer styles than the other species, as measurements of 2.8 and 3.4 mm have been recorded for styles on the single fruiting specimen. No other reliable measurements of mature styles could be made, but specimens in the early stages of flowering have shorter styles to c. 2.5 mm long. Notes. A single 2-flowered peduncle has been observed, indicating that the peduncles are not always 1-flowered. Material with mature fertile fruits is needed. Scholtzia recurva Rye, sp. nov. Typus\ south of Overlander Roadhouse, Western Australia [precise locality withheld for conservation reasons], 26 September 2002, M.E. Trudgen 21720 {holo\ PERTH 08986525; iso\ CANB, K, MET). Scholtzia sp. Overlander (M.E. Trudgen 12138), Western Australian Herbarium, inFloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 20 July 2018]. Shrub 1-2 m high, 1.5-3.0 m wide, rather spindly; flowering branchlets commonly with peduncles borne at 1-4 consecutive nodes in a raceme-like arrangement. Leaves antrorse. Petioles 0.1-0.35 mm long. Leaf blades broadly obovate, 1.3-1.7 mm long, 1.1-1.5 mm wide, with a clear-translucent margin less than 0.1 mm deep, denticulate/ciliolate, the apex recurved and with a point up to 0.15 mm long; abaxial surface gradually raised towards centre, sometimes grooved along midvein, with lateral veins obscure, the oil glands in 1 or 2 main rows on each side of midvein, with 3-5 glands in innermost rows. B.L. Rye, Update to the taxonomy of Scholtzia 75 Peduncles 4-6.5 mm long, 0.3-0.35 mm wide, mostly 6-9-flowered; secondary axes up to 1.3 mm long. Bracts 1.3-1.5 mm long, deciduous. Pedicels 0-0.5 mm long. Flowers c. 4 mm diam. Hypanthium 1.5-2 mm long, dotted with oil glands but otherwise rather smooth. Sepals ± debate to very broadly ovate, 0.35-0.45 mm long, the outer ones with a green ridge; petaline margin c. 0.15 mm deep, entire. Petals 1.5-1.7 mm long, white or pale pink. Stamens commonly 10, with 1-3 opposite each sepal. Longest filaments c. 0.35 mm long. Anthers c. 0.3 mm long. Ovary inferior, 2-locular; ovules 2 per loculus. Style c. 0.5 mm long; stigma capitate. Fruits c. 3/4 inferior, sterile ones 1.6-1.75 mm long, 1.3-1.4 mm wide; sepals erect or incurved; petal persistence unknown. Seeds probably c. 1.2 mm long. Diagnostic features. Among the species that have a 2-locular ovary with 2 ovules per loculus, S. recurva is distinguished by the following combination of characters: petioles mostly 0.1-0.3 mm long; leaf blades 1.3-1.7 mm long, longer than wide, with apexrecurved and shortly pointed; peduncles 4-6.5 mm long, mostly 6-9-flowered; hypanthium smooth apart from its glands. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 22 Oct. 1994, M.E. Trudgen 12138 (NSW, PERTH); 26 Sep. 2002, M.E. Trudgen 21721 (PERTH). Distribution and habitat. Occurs in the Meadow Station area (Figure 6A), in red or orange-brown soil on the slopes of dunes or in the swale, with Acacia high open shrubland. Phenology. Flowers recorded in late October. A few old fruits were present on the specimens collected in late September. Etymology. From the Eatin recurvus (curved backwards), referring to the recurved apex of the leaves, which is of importance in distinguishing this species. Conservation status. Recently listed as Priority One under Conservation Codes for Western Australian Flora. Known from three collections that were possibly all made from a single population. Affinities. Closest in morphology to S. thinicola Rye but differing in having leaves with a recurved and shortly pointed apex. Scholtzia recurva also has shorter sepals with a narrower petaline margin, and tends to have longer peduncles but shorter pedicels. The two species occupy a similar habitat but with S. recurva occurring more than 100 km north of the range of S. thinicola. Notes. Scholtzia recurva appears to have the smallest leaves of all the species with two ovules per loculus. The description of its flowers is based entirely on one specimen. Further collections of both flowering and fruiting material are needed. All fruits dissected were sterile, although one had a cavity that appeared to have contained an incompletely formed seed c. 1.2 mm long. Scholtzia subsessilis Rye, sp. nov. Typus\ South-west of Morawa, Western Australia [precise locality withheld for conservation reasons], 15 September 1985, B.J. Conn2\59 {holo\ PERTH 03414515; iso\ B, CHR, MEE, MO, NSW all n.vf Scholtzia sp. Billeranga Hills (B.J. Conn 2159), Western Australian Herbarium, in FloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. 76 Nuytsia Vol. 30 (2019) Figure 6. Distribution of Scholtzia species. A-S. recurva (a) and S. truncata (•); B-S. subsessilis (o), S. tenuissima (a), S. trilocularis (•) and S. sp. Whelarra (a). Shrub 0.8-2.5 m high, erect, with one record of 1.5 m wide; flowering branchlets with peduncles borne at 1-5 consecutive nodes, often with the flowers combined into a dense cluster. Leaves antrorse or widely antrorse. Petioles absent or not exceeding 0.25 mm long. Leaf blades mostly broadly elliptic or broadly obovate, 1.5-2.3 mm long, 1.7-2.2 mm wide; abaxial surface often raised and grooved in basal 1/2-2/3 and ridged along midvein above, or ridged for full length, with lateral veins obscure, the oil glands in 2 or 3 main rows on each side of midvein, with 3-5 glands in innermost rows. Peduncles 2.5-5.5 mm long, 0.4-0.6 mm wide, 3-6-flowered; secondary axes absent or up to 0.3 mm long. Bracts 0.8-1.4 mm long, caducous or deciduous. Pedicels 0.2-0.8 mm long. Flowers 5-6 mm diam. Hypanthium 1.1-1.5 mm long, rugose-pitted. Sepals broadly to depressed ovate, 0.5- 0.6 mm long, the outer sepals ridged and dark reddish at base; petaline margin c. 0.3 mm deep, entire. Petals 2.3-2.6 mm long, white or pink. Stamens usually 8-10, with 1-3 opposite each sepal. Longest filaments 0.7-0.8 mm long. Anthers c. 0.3 mm long. Ovary inferior, 3-locular; ovules 1 per loculus. Style 1-1.2 mm long; stigma tending to be peltate. Fruits largely inferior, not seen at maturity; sepals incurved; petals deciduous. Diagnostic features. Among species that have a 3-locular ovary with 1 ovule per loculus, S. subsessilis is distinguished by having sessile or subsessile leaves. Other important characters: peduncles 2.5-5.5 mm long, 3-6-flowered; hypanthium pitted-rugose; stamens usually 8-10, with 1-3 opposite each sepal. Selected specimens examined. [localities withheld for conservation reasons] 12 Sep. 1996, A. Carr 368 (PERTH); 25 Sep. 1990, R.J. Cranfield 7847 & P.J. Spencer (CANB, PERTH); 19 Aug. 1997, F Keast M6A 067 (PERTH). B.L. Rye, Update to the taxonomy of Scholtzia 11 Distribution andhabitat. Occurs in the Billeranga Hills area, west of Morawa (Figure 6B), in sandy soils. Phenology. Flowers from August to December. Fruits recorded from September to December. Etymology. From the Latin sub- (somewhat, less than) and sessilis (sessile, unstalked), as the leaves are almost sessile. Vernacular name. Billeranga Scholtzia. Conservation status. Recently listed as Priority One under Conservation Codes for Western Australian Flora. Scholtzia sub sessilis does not appear to have been collected from any nature reserves. Affinities. The affinities of S. subsessilis are unclear, but it shows some similarities to S. halophila, which has longer petioles and peduncles. Notes. Good fruiting material is needed for this species. Scholtzia tenuissima Rye, sp. nov. Typus: Kalbarri National Park, Western Australia [precise locality withheld for conservation reasons], 30 November 2003, M.E. Trudgen MET 22155 {holo\ PERTH 08238162; iso\ CANB, K, MEE). Scholtzia sp. Z-Bend (Bellairs-Kalflora 912a), in G. Paczkowska & A.R. Chapman, West. Austral. FI : Descr. Cat. p. 402 (2000); Western Australian Herbarium, in FloraBase, https://florabase.dpaw. wa.gov.au/ [accessed 2 March 2018]. Shrub spindly, 0.4-1.0 m high, 0.3-1.0 m wide; flowering branchlets with peduncles borne at \-4 (-7) consecutive nodes, in a very open raceme-like arrangement or with flowers more clustered. Leaves appressed or antrorse. Petioles 0.1-0.3 mm long. Leaf blades narrowly obovate or obovate, 1.4- 2.7 mm long, 0.6-1 mm wide, 0.3-0.5 mm thick; abaxial surface highly raised throughout, rounded or flattened, with lateral veins obscure, the oil glands in 2 or 3 main rows on each side of midvein, with 4-6 glands in innermost rows. Peduncles 5.5-13 mm long, 0.15-0.25 mm wide, 1-3-flowered; secondary axes up to 2.5 mm long. Bracts 0.8-1.3 mm long, deciduous. Pedicels 0-0.5 mm long. Flowers 4-5.5 mm diam. Hypanthium 1.2-1.4 mm long, rugose. Sepals mostly ± broadly ovate, 1.0- 1.4 mm long, scarious throughout, not ridged, tending to be recurved, entire. Petals 1.5-2.5 mm long, white. Stamens 12 or 13, with 1-3 opposite each sepal. Longest filaments 0.8-1.2 mm long. Anthers c. 0.35 mm long. Ovary inferior, 2-locular; ovules 2 per loculus. Style 0.8-1.0 mm long; stigma capitate. Fruits c. 1/2 inferior, 1.8-2.2 mm long, 1.3-1.4 mm wide; sepals reflexed; petals persistent. Seeds 1.3-1.4 mm long, 0.6-0.7 mm wide. Diagnostic features. Among the species that have a 2-locular ovary with 2 ovules per loculus, S. tenuissima is distinguished by the following combination of characters: petioles 0.1-0.3 mm long; peduncles extremely slender (0.15-0.25 mm wide), 1-3-flowered, 5.5-13.0 mm long, with secondary axes up to 2.5 mm long; hypanthium mgose; sepals entirely scarious, reflexed in fruit. 78 Nuytsia Vol. 30 (2019) Selected specimens examined. WES TERN AUSTRALIA: [localities withheld for conservation reasons] 17NOV. \9m,Bellairs-Kalflora9\2A(?mJYL)-6m\. 1995,7). &B. 3008 (PERTH); 16Sep. 1999, D. & B. Bellairs 6041 (PERTH^ Distribution and habitat. Occurs in Kalbarri National Park (Figure 6B), in sandy soil, sometimes over sandstone or laterite, one record from a winter-wet flat, where S. tenuissima formed a low shrubland over Lepidobolus open sedgeland. Phenology. Flowers from September to December. Mature fruits recorded in November Etymology. From the Latin tenuis (thin) and -issimus (very), i.e. the thinnest or most delicate, in reference to the peduncles, which are long (up to 13 mm) but extremely slender (0.15-0.25 mm wide). Vernacular name. Delicate Scholtzia. Conservation status Priority Two under Conservation Codes for Western Australian Flora. Listed with this priority Smith and Jones (2018) under the name S. sp. Z-Bend (Bellairs-Kalflora 912a). Known from an area c. 30 km long within a large national park. Co-occurring species. Scholtzia tenuissima (M.E. Trudgen MET 22154) has been recorded adjacent to, and downhill from, an area where S. oleosa {M.E. Trudgen MET 22151) and a member of the S. obovata complex (M.E. Trudgen MET 22153) co-occurred. Affinities. Very similar to S. sp. Whelarra (M.E. Trudgen 12018), both taxa having a somewhat spindly appearance and very slender peduncles. Scholtzia sp. Whelarra is very poorly known but occurs to the south-east (Figure 6B) and has broader, smoother, less thickened leaves, somewhat wider peduncles, much shorter secondary axes, and possibly fewer stamens. Good flowering material is needed to determine stamen numbers and other flowering characters for S. sp. Whelarra, but the differences in its leaves indicate that it should probably be treated as a distinct species rather than as a subspecies of S. tenuissima. Notes. The sepals are about 2/3 the length of the petals. Only S. denticulata sometimes has a greater sepal/petal length ratio. Scholtzia thinicola Rye, sp. nov. Typus: Binnu East Road, Western Australia [precise locality withheld for conservation reasons], 5 December \993, M.E. &M.R. Trudgen MET 12013 {holo\ PERTH 03979318; iso\ AD, BRI, CANB, K, NSW, MEL). Scholtzia sp. Binnu East Road (M.E. Trudgen 12013), in G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. p. 401 (2000); Western Australian Herbarium, mFloraBase, https ://florabase. dpaw.wa.gov.au/ [accessed 2 March 2018], Scholtzia sp. Valentine Road (S. Patrick 2142), G. Paczkowska & A.R. Chapman, West. Austral. FI : Descr. Cat. p. 402 (2000); Western Australian Herbarium, inE/ora^tz^e, https://florabase.dpaw.wa.gov. au/ [accessed 2 March 2018], B.L. Rye, Update to the taxonomy of Scholtzia 79 Shrub commonly 1.0-1.5 m high, with one record of 2 m wide; flowering branchlets with peduncles borne at 1-3 consecutive nodes with the flowers combined into a dense cluster. Leaves mostly widely antrorse. Petioles 0.1-0.3 mm long. Leaf blades obovate to almost circular in outline, 1.4-3.7 mm long, 1.0-1.6 mm wide, 0.35-0.6 mm thick, sometimes with a minute subterminal point at flrst, entire; abaxial surface somewhat angled, with the central part flattened and with sloping sides, with lateral veins obscure, the oil glands usually in 2 or 3 main rows on each side of midvein but inconspicuous. Peduncles 0.7-4 mm long, 0.5-0.7 mm wide, mostly 3-9-flowered; secondary axes scarcely developed or up to 0.6 long. Bracts 0.1-\ .4 mm long, deciduous or persistent. Pedicels 0.3-2.0 mm long. Flowers c. 4 mm diam. Hypanthium 0.8-1.2 mm long, rugose. Sepals transversely oblong to depressed ovate, mostly broadly ovate, 0.7-1.2 mm long, the herbaceous base ridged; petaline margin 0.4-0.6 mm deep, entire. Petals 1.5-2.3 mm long, white or pale pink. Stamens 10-12, with 1-3 opposite each sepal. Longest filaments 0.25-0.4 mm long. Anthers 0.4-0.5 mm long. Ovary inferior, 2-locular; ovules 2 per loculus. Style 0.5-0.8 mm long; stigma capitate. Fruits 1/2-2/3 inferior, 1.3-2.0 mm long, 1.3- 1.5 mm wide; sepals erect; petals persistent for some time. Seeds 0.9-1.6 mm long, 0.6-0.8 mm wide. Diagnostic features. Among the species that have a 2-locular ovary with 2 ovules per loculus, S. thinicola is distinguished by the following combination of characters: petioles 0.1-0.3 mm long; leaf blades 1.4-3.7 mm long, longer than wide, with apex straight and not pointed; peduncles 0.7^.0 mm long, mostly 3-9-flowered; hypanthium rugose. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 21 Dec. 2005, A.D. Crawford 841 (PERTH); 12 Jan. 2005, A.D. Crawford 871 (K, PERTH); 29 Oct. 1991, G.J. Keighery 14734 (PERTH); 1 Nov. 1994, SJ. Patrick 2142 (PERTH); 12 Dec. 2002, S.J. Patrick & G. Paczkowska SP 4676 (PERTH). Distribution and habitat. Occurs from near Yandi Station south to east of Mullewa (Figure 5A), on yellow sand dunes, with several records from the crest of the dunes. Phenology. Flowers from October to December, with mature fruits recorded from November to January. Etymology. From the Eatin thinium (dune) and -cola (dweller) since the species is known only from sand dunes. Vernacular name. Dune Scholtzia. Conservation status. Priority One under Conservation Codes for Western Australian Flora. Fisted with this priority by Smith and Jones (2018), under the names S. sp. Binnu East Road (M.E. Trudgen 12013) and Scholtzia sp. Valentine Road (S.J. Patrick 2142). This taxon is currently known from five locations, two of which are very close and probably represent a single population. Co-occurring species. The type specimen {M.E. & M.R. Trudgen MET 12013) was collected from a site where it co-occurred with S. uniflora {M.E. & M.R. Trudgen MET 12011). Affinities. Very similar to S. recurva in morphology and habitat; see affinities section under that species for details. 80 Nuytsia Vol. 30 (2019) Notes. Like the inflorescences, the infmctescences are globular and dense, and located well below the apex of each branchlet. The name S. sp. Valentine Road (SJ. Patrick 2142) was applied to southern specimens that tend to have longer peduncles than the other specimens. Scholtzia trilocularis Rye, sp. nov. Typus'. 20 km north of Eneabba on Brand Highway, Western Australia, 8 October 1990, S. Maley 8 (holo\ PERTH 04279115; iso\ AD, MEE, NSW). Scholtzia sp. Eneabba (S. Maley 8), in G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. p. 402 (2000); Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 2 March 2018]. Shrub erect, 0.4-2(-3?) m high, 1-2.3 m wide; flowering branchlets with peduncles borne at 1^ consecutive nodes, often in an umbel-like arrangement. Leaves antrorse or widely antrorse. Petioles 0.2-0.8 mm long. Leaf blades obovate to almost circular, 1.5-3.5 mm long, 1.1-2.1 mm wide, entire; abaxial surface usually raised and flattened at centre in basal 1/2-3/4 and distally ridged along midvein or sometimes ridged along full length, with lateral veins obscure, the oil glands in 1-3 main rows on each side of midvein but inconspicuous. Peduncles 4-15 mm long, 0.4-0.6 mm wide, mostly 3-13-flowered; secondary axes up to 1.3 mm long. Bracts 0.9-1.8 mm long, deciduous. Pedicels mostly 0.6-1.0 mm long. Flowers 3.5-7 mm diam. Hypanthium 1.0-1.4 mm long, wrinkled- rugose or sometimes reticulate-rugose. Sepals usually broadly to depressed ovate, 0.5-1.0 mm long, the outer ones slightly to markedly ridged at base; petaline margin 0.3-0.6 mm deep, entire. Petals 1.5-2.5 mm long, pale or medium pink. Stamens 8-12, with 1-3 opposite each sepal. Longestfilaments 0.4-0.7 mm long. Anthers c. 0.35 mm long. Ovary inferior, 3-locular; ovules 2 per loculus. Style 0.7- 1.0 mm long; stigma capitate. Fruits c. 2/3 inferior, 1.6-1.8 mm long, c. 1.4 mm wide; sepals erect or spreading; petals persistent in early or mature fruit. Seeds 1.3-1.6 mm long, 0.6-0.7 mm wide. Diagnostic features. Among species that have a 3-locular ovary with 2 ovules per loculus, S. trilocularis is distinguished by the following combination of characters: leaves entire; peduncles 4-15 mm long, mostly 3-13-flowered; stamens 8-12, with 1-3 opposite each sepal; petals 1.5-2.5 mm long. Selected specimens examined. WES TERN AU S TRAEIA: Eoc. No. 144, Arrowsmith River Valley, near Blue Water Rd. Shallow siltstone (Mound Springs 16-19), E side, Arrino, 17 Oct. 2005, J. Borger CH 1710-1 (PERTH); Nebroo Nature Reserve, Bunney Rd, Western side Tag 21 ferricrete site, 17 Nov. 2009, J. BorgerNKll 1 (PERTH); track between Ambania and Casuarina Rds,10 km S off Geraldton road, 13 Nov. 2005, J. Docherty 414 (PERTH); Moore Rd, 9 km S of Mullewa-Geraldton Rd, Indarra Nature Reserve, 23 Oct. 1998, G.J. Keighery & N. Gibson 5025 (BRI, CANB, PERTH); Mingenew, Oct. 1909, J.H. Maiden s.n. (PERTH); Brand Mudge Rd, 2.9 km N of Hughes Rd, W of Coorow, 22 Oct. 2001, S.J. Patrick 4051 (NSW, PERTH); on E boundary of park, Watheroo National Park, W of Watheroo, 7 Oct. 1971, R.D. Royce 9697 (PERTH). Distribution and habitat. Extends from near Ambania (east of Geraldton) south to Watheroo National Park (Figure 6B), usually in sandy habitats, commonly in yellow sand, the dominant species often eucalypts (such as E. todtiana),Allocasuarina campestris, Banksia species orXylomelum angustifolium. Phenology. Flowers from August to November. Mature fruits recorded in November and December. B.L. Rye, Update to the taxonomy of Scholtzia 81 Conservation status. Not listed by Smith and Jones (2018); this species is known from a national park and from several nature reserves. Etymology. From the Latin tri- (three) and locularis (having compartments or cells). Most of the species with two ovules per loculus have regularly 2-locular ovaries or have both 2- and 3-locular ovaries common, whereas this species is one of four that are regularly 3-locular. Co-occurring species. Recorded (M Hislop WF 3362) growing with two variants of S. laxiflora (M. Hislop WF 3363 & WF 3367) near Eneabba. Affinities. Keys out with S. spatulata, which tends to have larger flowers with the sepals less ridged at the base. Another possible close relative is S. oligandra F.Muell. ex Benth., which differs in having denticulate to ciliate leaves and only 5-8 stamens. Notes. In S. trilocularis all of the flowers examined had at least one stamen opposite each sepal, whereas in S. spatulata there may not be any stamens opposite one of the sepals of a flower. Scholtzia truncata Rye, sp. nov. Typus\ Eurardy Station, Western Australia, 2 September 2004, Wildflower Society ofWA EURA 171 {holo\ PERTH 06922856; iso\ K, MEE). Scholtzia sp. Galena (WE. Blackall 4728), in G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. p. 402 (2000); Western Australian Herbarium, mFloraBase, https://florabase.dpaw.wa.gov. au/ [accessed 2 March 2018]. Scholtzia sp. Ajana East Road (M.E. Trudgen 21734 A), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 30 May 2016]. Shrub 0.5-3 m high, 0.6-3.5 m wide; flowering branchlets with peduncles borne at 1-5 consecutive nodes, often with their flowers combined into a terminal globular cluster. Leaves appressed to widely antrorse, but mostly antrorse. Petioles 0.3-0.5 mm long. Leaf blades broadly obovate to very broadly elliptic, 1.3-2.2 mm long, 1.4-2.0 mm wide, entire, with a very short mucro present on at least some of the leaves of each specimen; abaxial surface flattened at the centre for 1/2-2/3 of its length and ridged distally, with lateral veins obscure; oil glands in 1 or 2 main rows on each side of midvein, with 2-5 glands in innermost rows. Peduncles 0.5-2.0(-2.3) mm long, 0.4-0.6 mm wide, 1-3-flowered; secondary axes ± absent or up to 0.4 mm long. Bracts 0.6-1.8 mm long. Pedicels 0.3-1.0 mm long. Flowers 3.5-6.0 mm diam. Hypanthium 1.0-1.3 mm long, wrinkled or reticulate-rugose to pitted, sometimes becoming fairly smooth in mature fruit. Sepals broadly or very broadly ovate, 0.5-0.7 mm long, the base ribbed (but appearing scarious in fruit); petaline margin often hidden from side view, 0.3-0.5 mm deep, entire. Petals 1.5-2.5 mm long, white or pale pink. Stamens (5-)7-10(-12), with 0-3 opposite each sepal. Longest filaments 0.5-0.^ mm long. Anthers 0.25-0.3 mm long. Ovary inferior, 2- or 3-locular; ovules 1 per loculus. Style 0.9-1.6 mm long; stigma peltate. Fruits c. 1/2 inferior, 1.4-1.5 mm long, 1.8-1.9 mm wide; sepals strongly incurved; petals deciduous. Seeds 0.9-1.3 mm long, 0.6-0.8 mm wide. 82 Nuytsia Vol. 30 (2019) Diagnostic features. Among species that have 1 ovule per loculus, S. truncata is distinguished by the following combination of characters: petioles 0.3-0.5 mm long; peduncles 0.5-2.0(-2.3) mm long, 1-3-flowered; sepals strongly incurved; petals 1.5-2.5 mm long; stamens usually 7-11; style 0.9- 1.6 mm long, stigma peltate. Selected specimens examined. [localities withheld for conservation reasons] 14 Sep. 1990, A.H. Burbidge 4349 (PERTH); 26 Aug. 1991, A.H. Burbidge 4365 (PERTH); 6 Aug. 1976, R.J. Hnatiuk 760431 (PERTH); 8 Sep. 2016, B.L. Rye 290165, J. Norman & B. Parkhurst (PERTH); 29 Aug. 1985, C.I. Staceyl6^ (PERTH); 30 Aug. 2003, Wildflower Society of W. A. EURA26 (PERTH); 11 Oct. 2004, C. Wilkins & J. Wilkins CW 1984 (PERTH). Distribution and habitat. Extends from Eurardy Station south-east to near Indarra (Figure 6A), in yellow to brown sand in varied vegetation dominated by malices or high shrubs. Phenology. Flowers from June to September. Fruits recorded in September and October. Etymology. From the Eatin truncatus (cut off, blunt-ended) after the more or less truncate young flower buds with strongly incurved sepals that lie almost flat across the top. Conservation status. Priority Two under Conservation Codes for Western Australian Flora. Fisted with this priority by Smith and Jones (2018) under the name S. sp. Galena (W.E. Blackall 4728), this species is known from nature reserves and a national Park . Co-occurring species. Recorded at one location {Wildflower Society ofWA EURA 1) growing with S. obovata and at another (B.L. Rye 290165, J. Norman & B. Parkhurst) growing with S. peltigera. Affinities. This is similar to several other species that have a peltate stigma, but most of those species have longer peduncles (2.5-15 mm cf. 0.5-2(-2.3) mm long). Three species with peduncles shorter than 2.5 mm long differ from S. truncata in having only 3-6 stamens per flower. Notes. Currently S. truncata includes some specimens with somewhat pitted hypanthia (e.g. reference specimen of S. sp. Galena) as well as those with wrinkled ones (reference specimen of S. sp. Ajana East Road) but a few specimens are hard to classify into either category because they are somewhat intermediate in hypanthium patterning. This kind of difference in hypanthium patterning has proved useful in distinguishing some members of the genus and is used in the key, but in this case does not appear to be correlated with any other characters that could be used to recognise more than one species. Consequently the two phrase names are considered to be conspeciflc. Scholtzia truncata keys out at two positions (see key to species above) because of this variability in hypanthium ornamentation. Sepals are strongly incurved at all stages of flowering and fruiting. The fruits can be 2-seeded but mostly contain only one seed. Scholtzia uniflora Rye, sp. nov. Typus\ east of Binnu, Western Australia [precise locality withheld for conservation reasons], 13 September 1978, M.E. Trudgen 2218 {holo\ PERTH 03628167; iso\ CANB, MEE). B.L. Rye, Update to the taxonomy of Scholtzia 83 Scholtzia sp. Binnu (M.E. Trudgen 2218), in G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. p.401 (2000); Western Australian Herbarium, inE/oraRtjr^e, https://florabase.dpaw.wa.gov. au/ [accessed 2 March 2018]. Shrub commonly 1.2-2.5 m high and 1.4-2 m wide; flowering branchlets with peduncles borne at \-A consecutive nodes, often with their flowers combined into a dense cluster. Leaves antrorse. Petioles 0.5-0.8 mm long. Leaf blades broadly or very broadly obovate, 2-4 mm long, 2.0-3.5 mm wide, entire; abaxial surface raised only near the middle, often with a flattened section along the midvein that contracts into a ridge distally, with lateral veins usually obscure, the oil glands in more than 3 rows on each side of midvein, with 5-7 glands in innermost rows. Peduncles 0-0.5 mm long, 0.4- 0.6 mm wide, 1-flowered. Bracteoles 1.5-1.8 mm long. Pedicels 0.4-0.7 mm long. Flowers 5-7 mm diam. Hypanthium 1.8-2.0 mm long, wrinkled-rugose. Sepals transversely oblong to semicircular or deflate to depressed ovate, 0.7-1.3 mm long, entire, the outer ones smooth or slightly keeled; petaline margin 0.3-0.5 mm deep, entire. Petals 2.5-3.5 mm long, pale to medium pink. Stamens 15-17, in a continuous circle. Longest filaments c. 1 mm long. Anthers 0.4-0.5 mm long. Ovary inferior, 3-locular; ovules 1 per loculus. Style c. 1.6 mm long; stigma ± peltate. Fruits largely inferior, 2.2-2.3 mm long, 2.3-2.5 mm wide; sepals erect, but scarious margins incurved; petals deciduous. Seeds 1.3-1.4 mm long, 0.8-1.0 mm wide. Diagnostic features. Among species that have a 3-locular ovary with 1 ovule per loculus, S. unifiora is distinguished by the following combination of characters: peduncles absent or up to 0.5 mm long, 1-flowered; stamens 15-17, in a continuous circle. Selected specimens examined. [localities withheld for conservation reasons] 2 Dec. 2004, A.D. Crawford 807 (MEL, PERTH); 16 Aug. 2001, R. Davis 9935 (CANB, PERTH); \ My 2001, J.Docherty 62 (PERTH); 5 Dec. 1993,ME. Trudgen&M.R. Trudgen MET UOU (AD, BRI, K, NSW, PERTH). Distribution and habitat. Occurs east of Binnu (Figure 4A), and has been recorded in yellow sand on the top of a dune, in Actinostrobus tall open shrubland. Phenology. Flowers from July to September. Etymology. From the Latin unus (one) and -fiorus (-flowered). Refers to the presence of only one flower on each peduncle. Of the species with one ovule per loculus, this is the only one that has consistently 1-flowered peduncles. Conservation status. Priority Two under Conservation Codes for Western Australian Flora. Previously listed (Smith & Jones 2018) as Priority One under the name S. sp. Binnu (M.E. Trudgen 2218). Co-occurring species. At one locality, S. unifiora was recorded growing with S. thinicola. Affinities. Similar to S. sp. Nolba in having 15-17 stamens and a 3-locular ovary with one ovule per loculus. The fruits in both species are largely inferior and have a smooth, convex summit. Scholtzia sp. Nolba has larger leaves, multi-flowered peduncles and usually shorter sepals. All other species with 15 or more stamens have a 2-locular ovary with two ovules per loculus. 84 Nuytsia Vol. 30 (2019) Scholtzia uniovulata Rye, sp. nov. Typus'. Midlands Road, 2.5 km north of Coorow, Western Australia, 28 August 2003, B. Moyle & N. Gibson BM 005 (holo\ PERTH 08126291; iso\ 7CANB, K, MEL). Shrub 0.5-3(-4) m high, 0.3-3 m wide; flowering branchlets with peduncles borne at 1-6 consecutive nodes, often with their flowers combined into a dense cluster. Leaves antrorse to patent. Petioles 0.1- O. 3 mm long. Leaf blades obovate or broadly obovate, (0.8-)l .3-2.8(^) mm long, 1.2-1.6 mm wide, ± entire; abaxial surface raised towards the middle, often with a flattened section along the midvein that contracts into a ridge distally, with lateral veins obscure or sometimes visible, the oil glands in 1 or 2 main rows on each side of midvein, with 2-A glands in innermost rows. Peduncles 0.8-2.5(- 3.5) mm long, 0.3-0.45 mm wide, mostly 1-3-flowered. Bracts 0.8-1.5 mm long. Pedicels 0.5-0.8 mm long. Flowers 3.5^.5 mm diam. Hypanthium 1.0-1.4 mm long, rugose-pitted. Sepals broadly or very broadly obovate to transversely elliptic, of varied length, entirely petaline, the innermost one 0.8-1.2 mm long. Petals 1.2-1.6 mm long, white or pale pink. Stamens 3-6, 0-2 opposite each sepal (commonly 5 in the arrangement 2,0,1,2,0 or sometimes with mostly fewer than 5). Longestfilaments 0.2-0.4 mm long. Anthers 0.2-0.3 mm long. Ovary inferior, 1 (2)-locular (rarely 2-locular in a majority of the flowers); ovules 1 per loculus. Style 0.5-0.8 mm long; stigma ± peltate. Fruits 2/3-3/4-inferior, 1.0-1.2 mm long, c. 1.3 mm wide; sepals erect or spreading; petals deciduous. Seeds 0.7-0.8 mm long, 0.8-1.0 mm wide. (Figure ID) Diagnostic features. Among the species with 1 ovule per loculus, S. uniovulata is distinguished by its usually 1-locular ovary. Other important characters: hypanthium pitted; stamens 3-6. Selected specimens examined. WESTERN AUSTRALIA: Yuna, Sep. 1930, E. Ashby s.n. (ADW, NSW); in a dip in Cunderdin-Minnivale Rd, 0.8 kmN of Berry Rd, Minnivale Nature Reserve, 15 Oct. 2013, R. Davis & B.L. Rye DR 012 (PERTH); Wilroy, 17 Aug. 1964, J. Galbraith WA383 (MEL); Cowcowing, Sep.-Oct. 1904, M. Koch 1091 (AD, NSW); 7.1 km E of Konnongorring Siding on Dowerin-Konnongorring Rd, c. 27 km SW ofWongan Hills, 22 June 1996, T.R. Lally & B.J Lepschi TRL 1029 (CANB, PERTH); 6.1 km on road to Ellendale Pool from Walkaway-Nangetty Rd, 26 Oct. 2001, S.J. Patrick 40M (PERTH); Corrigin to Quairading, 28 Sep. 2002, M.E. Trudgen 21783 (AD, BRI, MEL, NSW, PERTH). Distribution and habitat. Extends from the Greenough River area south-east to near Corrigin (Figure 5B), usually in sandy soils, occasionally with laterite or gravel. Occasionally the species occurs along watercourses, where it has been recorded reaching heights of up to 4 m or possibly more. Phenology. Flowers from June to November. Etymology. From the Latin unus (one) and ovulatus (having ovules). Refers to the presence of only one ovule in each loculus of the ovary and frequently only one ovule per ovary as the ovary is usually unilocular. Vernacular name. Common Scholtzia. Scholtzia uniovulata appears to be distributed over the greatest area in the genus, although the coastal species S. obovata has a longer range if S. umbellifera is treated as its synonym. B.L. Rye, Update to the taxonomy of Scholtzia 85 Conservation status. Being one of the most common species in the genus, S. uniovulata is not considered to be at risk. Chromosome number, n = \ \,fide B.L. Austral. J. Bot. 27: 571 (1979) [as Scholtziaparviflora]. Voucher: B.L. Powell 73128. Affinities. Many herbarium specimens have previously been identified as either S. parviflora or S. oligandra, both of which have an excluded syntype that matches this new species (see Rye 2017). Scholtzia oligandra is not closely related, differing in having a 2- or 3 -locular ovary with two superposed ovules in each loculus. Scholtzia parviflora is very similar in morphology to S. uniovulata, but has more uniform sepals, with the innermost one 0.3-0.6 mm long (cf 0.8-1.2 mm), a consistently 2-locular ovary, and usually larger leaves. Most specimens of S. uniovulata are distinguished from those of all other species of Scholtzia by having a 1-locular ovary with a single ovule, but the much rarer specimens with a 2-locular ovary in all or most fiowers are fairly widely dispersed. Co-occurring species. No confirmed cases of co-occurrence are recorded on PERTH herbarium specimens but S. uniovulata is a widespread species that overlaps in range with many others. Notes. Many specimens have tiny leaves less than 1.3 mm long on most of their branchlets, although all specimens have at least a few leaves greater than 1.3 mm long. Leaves 3-A mm long are atypical and have only been observed on very fast-growing shoots. Despite the large number of specimens available for study at PERTH, only one could be found with fully mature, fertile fruits. Acknowledgements I am grateful for the loan of specimens from AD, BRI, CANB, MEL and NSW. Images of type specimens were checked on GlobalPlants (2018). I would also like to thank Alex George for nomenclatural advice, Ben Parkhurst for collecting type material for one of the species, Rob Davis, Kevin Thiele and Jean Hort for their images, Steve Dillon for help with the mapping, Lisa Rye for arranging the figures and Julia Percy-Bower for assistance with specimen processing. Part of this work was supported by ABRS funding. References Bentham, G. (1867). Flora Australiensis. Vol. 3. (Reeve & Co.: London.) Blackall, W.E. & Grieve, B.J. (1980). Flow to know Western Australian wildflowers. Part 3A. Revised 2"‘' edn by B.J. Grieve. (University of Western Australia Press: Nedlands, Western Australia.) Global Plants (2018). JSTOR https://plants.jstoer.org/ [accessed 30 January 2018], Mueller, F. (1864). Myrtaceae. In: Fragmentaphytographiae Australiae. Vol. 4. pp. 51-77. (Government Printer: Melbourne.) Paczkowska, G. & Chapman, A.R. (2000). The WesternAustralianflora: a descriptive catalogue. (WesternAustralian Herbarium: Kensington, Western Australia.) Rye, B.L. (1979). Chromosome number variation in the Myrtaceae and its taxonomic implications. Australian Journal of Botany 21 ■. 547-573. Rye, B.L. (1987). Myrtaceae. In: Marchant, N.G., Wheeler, J.R., Rye, B.L., Bennett, E.M., Lander, N.S. & Macfarlane, T.D. Flora of the Perth region. Vol. 1. pp. 377-423. (WesternAustralian Herbarium, Perth.) Rye, B.L. (2014). An update to the taxonomy of some Western Australian genera of Myrtaceae tribe Chamelaucieae. 3. Thryptomene. Nuytsia 24: 269-306. 86 Nuytsia Vol. 30 (2019) Rye, B.L. (2016). An update to the taxonomy of some Western Australian genera of Myrtaceae tribe Chamelaucieae. 4. Malleostemon. Nuytsia 27: 103-120. Rye, B.L. (2017). New lectotypes and synonyms in the Western Australian genus Scholtzia (Myrtaceae: Chamelaucieae). Nuytsia!^-. 159-167. Schauer, J.C. (1843). Genera Myrtacearum nova vel denuo recognisa. Linnaea 17: 235-244. Schauer, J.C. (1844). Myrtaceae R.Br. In'. Lehmann, C. Plantae Preissianae. Vol. 1. pp. 98-158 (Meissneri: Hamburg.) Smith, M.G. & Jones, A. (2018). Threatened and Priority Flora list 16 January 2018. Department of Parks and Wildlife. https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 30 January 2018]. Turczaninow, N. (1862). Decus octava. Generum plantarum hucusque non descriptorum. Bulletin de la Societe /raperiale des Naturalistes de Moscou 35: 321-325. Western Australian Herbarium (1998-). FloraBase-the Western Australian flora. Department of Biodiversity, Conservation and Attractions. http/Zwww.dbca.wa.gov.au/ [accessed 30 January 2018] Nuytsia The journal of the Western Australian Herbarium 30: 87-154 Published online 16 April 2019 A revision of the tiurndins (Tribonanthes, Haemodoraceae) Ellen J. Hickman^’^ and Stephen D. Hopper^ 'Centre of Excellence in Natural Resource Management and School of Biological Sciences, University of Western Australia, Albany 6330, Western Australia, Australia ^Corresponding author, email: ellen.hickman@uwa.edu.au Abstract Hickman, E.J. & Hopper, S.D. A revision of the tiurndins (Tribonanthes, Haemodoraceae). Nuytsia 30: 87-154 (2019). The tiurndins {Tribonanthes Endl.) comprise a small taxonomically challenging genus of geophytes endemic to the Southwest Australian Floristic Region. Six named and one undescribed species of Tribonanthes are recognised in recent literature. Tribonanthes poses the most difficult taxonomic problems remaining for the Haemodoraceae in Western Australia. It is a genus where herbarium studies alone are insufficient to delineate taxa. A knowledge of variation in wild living material has proved essential to resolve taxonomic difficulties in the tiurndins. This revision recognises and comprehensively illustrates living plants of the 12 recognised species, four of which are new {T elongata E.J.Hickman & Hopper, T keigheryi E.J.Hickman & Hopper, T monantha E.J.Hickman & Hopper and T porphyrea E.J.Hickman & Hopper). Tribonanthes variabilis Eindl. and T uniflora Eindl. are reinstated. Morphological and molecular data also support the recognition of three new subgenera: subg. Tribonanthes, subg. Salina E.J.Hickman & Hopper and subg. Boya E.J.Hickman & Hopper. Botanical illustration, especially of internal floral morphology, has revealed several previously unrecorded characters that help diagnose the species of Tribonanthes. Introduction Tribonanthes Endl. is a small, taxonomically challenging, genus of geophytes comprising some 12 species (this study) placed in Haemodoraceae subfamily Conostylidoideae (Eindl.) T.Macfarlane & Hopper. The genus is endemic to the Southwest Australian Floristic Region (SWAFR, sensu Gioia & Hopper 2017). Tribonanthes is currently placed in the monotypic Tribe Tribonantheae T.Macfarlane & Hopper, sister to all other members of the Conostylidoideae (Macfarlane & Hopper 1987; Simpson 1990; Hopper etal 1999,2009; Smith etal. 2011). Divergence of Tribonanthes has been estimated as occurring in the Eocene about 40 MA, with contemporary species diversifying since 5 MA (Hopper etal 2009). No common name has been adopted for the genus, except occasionally ‘flannel flowers’, which is confusing as it has been widely taken up for Actinotus Eabill. in the Apiaceae. There is also no definitive Aboriginal name for members of Tribonanthes recorded in the literature or among extant Noongars SDH has interviewed over the past four decades. Colonial botanist James Drummond (1842a: 4), in a discussion of the SWAFR’s rich Haemodoraceae flora, mentioned a possible name, but left its application in doubt: ‘There are two sorts of round white roots called Jitta Cara by the natives, which they sometimes eat; pigs also are fond of them. I cannot © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) Nuytsia Vol. 30 (2019) refer these plants with certainty to any described.’ Grey (1840: 54) listQdJeeta as ‘the root of a species of rush; this root is something like a grain of Indian corn in appearance and resembles it in taste. It is in season in June.’ Moore (1884: 37) gave jitta as meaning ‘the bulbous root of an orchis, eaten by the natives, about the size of a hazel nut’, and kara was for ‘a spider’. Some modem Noongars use Kara for Burchardia congesta Lindl. (Daw et al. 1997), with clusters of long slender white corms, but Drummond (1842b: 5) was clear that 'carra" was applied to the much sought after orchid tuber of Lyperanthus suaveolens (= L. serratus Lindl.) in Western Australia. These alternative applications render the inferred application of Kara to Tribonanthes problematic. Another similarly problematic name is djoobak(djubak, dubakjoobakjubak, tubac, tuboc). Historically, this was applied to an edible small yam, usually of an orchid ‘... in season in .. October. The flower is a pretty white blossom, scented like the heliotrope.’ (Moore 1884: 22). Resemblance to a hyacinth was suggested by Grey (1840: 33). Botanist James Drummond (1842b: 5) stated that ' Prasophyllum giganteum produces the roots called dubad by the natives’. Modern Noongars apply the name to the orchid Pyrorchis nigricans (R.Br.) D.L. Jones & M.A.Clem. (Daw et al. 1997) but Yued elder Ned Mippy identified as djoobak specimens of Tribonanthes longipetala near Moora and New Norcia (Rooney 2002: 286). However, subsequently Rooney (2011: 125) gave a more general meaning for djoobak as a native potato, an ‘edible kidney-shaped tuber species’, and Wheatbelt Natural Resource Management (n.d.) similarly listed djubak as a general term for bush potato. Brandenstein (1988: 115) conveyed that some Esperance Noongars applied tyuubaq to the tubers of Platysace ejfusa (Turcz.) C.Norman (Apiaceae). It remains difflcult to unequivocally apply this name to tubers of Tribonanthes species, with the weight of evidence pointing more towards traditionally being a name for a scented, multiflowered (like a hyacinth) edible orchid tuber. Here, as a mark of respect to the indigenous people of the SWAFR, we propose instead that diurndin ’ is an apposite common name for the genus Tribonanthes. Tiurndin (pronounced churn-din) means ‘fair, white (flowers)’ according to Noongar informants from the Esperance region as recorded by von Brandenstein (1988: 101). Introduced sheep sometimes were also bestowed the same name, indicating that white wool may be regarded as meriting the label tiurndin. However, the first use of tiurndin was for white flowers, perhaps with a woolly appearance, which commends the name for most species of Tribonanthes. Tiurndin is derived from Tyiurtt, ‘the ancestral hero involved in the creation of the southern coastline’ (von Brandenstein 1988: 113), which is renowned for its brilliant white sand beaches. Indeed, Tyiurtt’s name recurs in many forms where the colour white appears on the south coast, as in chuditch (white-spotted native cat), Mt Chudalup (a granite inselberg bedecked by white lichen), maartyiurt (white-pointer shark = hands white-tipped), tyiurndur (bright stars), and Tyiurtigellup (Eake Seppings at Albany, brilliantly reflecting white light on a still, cloudy day). At present, six named and one undescribed species of Tribonanthes are recognised in contemporary literature (e.g. Parsons & Hopper 2003; Smith et al. 2011) and data bases maintained by the Western Australian Herbarium (1998-continuously updated). While some of these taxa are clear-cut, others are polymorphic and difihcult to key out with the latest taxonomic treatment, now three decades old (Macfarlane 1987). Indeed, these taxonomic problems were evident to Bentham (1873: 426) who wrote from studies of specimens at the herbarium in Kew that: ‘the species appear to be very variable and diflicult to distinguish’. We are currently involved in a broad study of the Haemodoraceae commenced in the 1970s (see Hopper et al. 2009 and references therein). This research soon identified the need to better understand taxonomic variation in Tribonanthes. Arguably, following recent resolution of tropical Kimberley Haemodorum Sm. species (Barrett et al. 2015), Tribonanthes poses the most diflicult taxonomic E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 89 problems remaining for the family in Western Australia, although there remain undescribed taxa in other genera still to be dealt with (Smith et al. 2011). Until the alpha-taxonomy of Tribonanthes is resolved, molecular phylogenetic and other biological studies will be, and have been, significantly hampered (e.g. Hopper et al. 2009), with the need for collecting voucher herbarium specimens for such studies remaining critical. Indeed, the few biological investigations conducted on Tribonanthes have been accompanied by the lodgement of few or no vouchers (e.g. Pate & Dixon 1982), rendering their results difficult to ascribe to species today. We have also been interested in exploring the use of modern techniques of botanical illustration as a means of discovering novel variation (Hickman et al. 2017). The contributions of botanical artists in discovery through illustration are rarely explicit in the literature. Here, we set out with the aim of having a botanical illustrator lead the research towards a taxonomic revision, and we make clear below what traits have been discovered through that process. Consequently, this paper focuses on a comprehensively illustrated revision of the genus Tribonanthes, with our combined efforts revealing the presence of 12 species, four of which are new and two reinstated. Taxonomic history If they were collected, no surviving specimens of Tribonanthes persist from the earliest maritime explorations of the SWAFR coastline, despite the genus being common enough around Perth, King George Sound and the Esperance region granites (Hopper 2003,2004). The earliest scientific specimens extant were collected by Baron Karl A.A. von Huegel during the first 12 days of 1834 when at King George Sound. We know from his field journal that he ascended Mt Clarence on January 5* (Hopper 2004), and that T australis Endl. occurs there today and on other nearby inselbergs such as Mt Melville. The precise location of HuegeTs collection was not documented. Endlicher (in Endlicher & Fenzl 1839) based comprehensive descriptions of the new genus and species on HuegeFs collection. No comparison with other genera was made, other than the sequence in which the description appeared, but the diagnostic characters of Tribonanthes cited below were well-described. An illustration of T australis was published by Endlicher in 1841 (Endlicher 1837^1) (Figure 1). James Drummond, who arrived in the Swan River Colony in 1829, undoubtedly was aware of species of Tribonanthes well before Huegel. However, Drummond’s earliest extant collections were made in 1839, some of which were used by Eindley (1839^0) as type material for T brachypetala Eindl. (CGE 06823) and T longipetala Eindl. (CGE 06824). Eindley (1839^0) named two other new species - T variabilis Eindl. based on Georgiana Molloy collections from the Vasse River (CGE 23342), and T uniflora Eindl. (CGE 06825) from Captain Mangles’ collections (or acquisitions from Drummond and others more likely - Hopper 2004). Eindley offered the rather disparaging comment that the tiumdins ‘are plants of no beauty, as far as can be ascertained by their appearance in the form of dried specimens’ (Eindley 1839^0: 44). Had he seen living plants, Eindley may have been more impressed by Tribonanthes. Eindley’s (1839^0) contribution was the first to reveal diversity amid seeming uniformity among the few herbarium specimens of Tribonanthes then available for study in Britain and Europe. This is a genus where herbarium studies alone are insufficient to delineate taxa. Field studies and dissection of living material are essential, we have found, to develop sound taxonomic insights for the tiumdins. Consequently, many herbarium collections have mixed material mounted on the same sheets. 90 Nuytsia Vol. 30 (2019) Figure 1. Uncoloured engraving of Tribonanthes australis from a drawing of Hugel’s King George Sound collection of this species by Putterlick, published by Endlicher in Iconographia Generum Plantarum (Endlicher, 1841) (Image from the Biodiversity HeritageEibrary. Digitisedby [FieldMuseuniNatural History Eibrary] https://www.biodiversityhbrary.Org/item/130502#page/235/ mode/lup [accessed 5 May 2016]). This illustration, in its original publication, is the neotype of T. australis. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 91 The type sheet of T. variabilis (CGE 23342), for example, includes one specimen of T. longipetala (second from left). All other specimens on the sheet are T. variabilis. Also, sketches of the stamen connective appendages at the bottom of the sheet show a T. variabilis stamen connective appendage on the left and a T. longipetala stamen connective appendage on the right. Lindley’s original description of T. variabilis, based on this collection, mentions two length conditions; ‘filamentis antherae subaequalibus dentatis v. petaloideis longioribus’ (Lindley 1839^0: 44) hence the name variabilis, alluding to what Lindley perceived as the variable nature of the stamen connective appendages in this species. Even more confusing is the type sheet of T. uniflora (CGE 08625) which includes three taxa. The single lectotype from Mangles on the left side of the sheet agrees well with Lindley’s protologue, notably in having testicular (cormous) roots, the bract and upper leaf with noticeably membranaceous margins, and the anthers and filaments much shortened. On the right hand side, the Molloy collection from ‘swamps. The Vasse’ (River near Busselton) has the earliest extant representatives of one of the new species we name below as T. porphyrea E.J.Hickman & Hopper (if stamen connective appendages are cream), or T. elongata E.J.Hickman & Hopper (if stamen connective appendages are yellow). A solitary plant embedded in this collection, second from the right, labelled ‘A and determined in 1984 as T. violacea Endl. by T.D. Macfarlane, is another new species collected by Mrs Molloy that we name below as T. keigheryi E.J.Hickman & Hopper. Endlicher (1846) published a new taxon {T. violacea, MEL 104289) based on collections of Ludwig Preiss made in October 1840 between Mounts Elphinstone and Melville at King George Sound (Albany). Endlicher erected new names for species already described by Lindley (1839^0) using Preiss specimens from Perth as types, thus creating the synonyms T. odora Endl. (= T. brachypetala) and T. lindley ana Endl. (= T. longipetala). George Maxwell, during or before 1873, on one of his expeditions eastwards from Albany discovered at Cape le Grand more Tribonanthes violacea (e.g. K 000356594, incorrectly determined as T. uniflora). On this same sheet are two specimens collected by Georgiana Molloy in 1841 at the Vasse River (K 000356593). The left-hand specimen is more of the earliest extant collection of one of the new species we name below as T. porphyrea or T. elongata while the right-hand specimen is of the slender swamp-dwelling T. keigheryi. Additional complexity on this sheet is evident in a single specimen (K 000356595) collected by Augustus Oldfield in 1873 at the Kalgan River north of Albany. This is correctly determined as T. unflora. Several such mixed collections were available to Bentham (1873), who correctly used Lindley’s (1839-40) four earlier names and effected the relevant synonymies for Endlicher’s (1846) later names. Moreover, Bentham, without the benefit of field studies of living material, regarded T. violacea as a synonym of T. uniflora, and suggested that T. variabilis was ‘perhaps a variety only of T. australis'. Understandably, given the limited and mixed material available to him, Bentham obfuscated as much as clarified the taxonomy of Tribonanthes. Mueller (1872-1874), perhaps at the time unaware of Bentham’s (1873) synonymies, supported Endlicher’s (1846) Tribonanthes species rather than Lindley’s (1839-40) which had priority. However, Bentham’s (1873) treatment prevailed for close to a century until Geerinck (1969: 65) suggested that Tribonanthes had only three, problematic species (i.e. the type T australis and two others not specified). No workers before or since Geerinck (1969) have agreed to such a conservative view of the number of species of tiumdins. 92 Nuytsia Vol. 30 (2019) Macfarlane’s (1987) Flora of Australia treatment, the first by a resident Western Australian botanist, achieved some clarification, although based on decisions informed by limited field work (Macfarlane pers. comm. 2014). He recognised four species among the six that Bentham (1873) had listed ( T. australis, T brachypetala, T longipetala, T. violacea), and synonymised two - T. uniflora under T. longipetala, and T. variabilis under T. australis. A year earlier, in 1986, one of us (SDH) with colleagues Sue Patrick and Andrew Brown from the then Department of Conservation and Land Management, had discovered an almost glabrous and stemless pink-fiowered new species on a granite outcrop in the central wheatbelt. This was named T. purpurea T. Macfarlane & Hopper in the appendix of the Flora of Australia volume in which Macfarlane’s (1987) treatment of Tribonanthes was published. Subsequently, a second striking new species was discovered by SDH (Hopper 6931, PERTH 07438761), accompanied by Stephen van Leeuwen, near Meckering in October 1988. This was a diminutive species, growing barely more than 3-5 cm tall, the smallest of any Haemodoraceae known. Four years later, in 2002, Mike Lyons collected this same undescribed species from near Pingrup and Lake King (Lyons & Keighery 2006), and it was named T. minor M.Lyons & Keighery. Lastly, in 1997 a small-fiowered and slender-stemmed taxon in swamps at Lake Muir was recognised as an undescribed species by G.J. Keighery and given the phrase name Tribonanthes sp. Lake Muir (G.J. Keighery & N. Gibson 2134). In recent times, then, Tribonanthes was regarded as comprising this unnamed species and six described species - T. australis, T. brachypetala, T. longipetala, T. minor, T. purpurea and T. violacea. Methods We have examined herbarium specimens of Tribonanthes, including types lodged at the Western Australian Herbarium (PERTH), and AD, BM, CANB, CBG, K NSW and MEL. Also images of types and specimens were examined through Global Plants (http://plants.jstor.org/) at B, E, L, LD, P, S and W. Opportunistic collections, photography and field notes of Tribonanthes were made by SDH since the 1970s during field work in the SWAFR. For this intensive revision, at least five geographically representative populations for each of the six described and one undescribed species were identified by EJH for field visits and collection of fresh material for illustration so that discernible variation within known species was covered. Not all populations were able to be relocated due to limited or incorrect GPS co-ordinate information (often due to retrospective or automated estimation) associated with the herbarium specimens, or historical habitat alteration or deterioration. During searches for target populations, if other Tribonanthes species were found, opportunistic collections were taken. A voucher herbarium collection was made for each taxon at each population and lodged at PERTH. Leaf material from specimens at each of the populations visited was collected for subsequent molecular analysis. The results (to be published elsewhere) were taken into account in making our taxonomic decisions. Colour botanical illustrations of each species we recognised, including variants within species, were made from fresh specimens with or without the aid of a dissecting microscope. Whole plant, details of above ground and below ground parts, including dissections of Lowers, capsules, leaves, corms and roots, were illustrated to discover and elucidate morphological character traits. Measurements of quantitative character traits for each species were taken from dried pressed material (Figure 2). Flowers, inflorescence and floral bracts were removed and rehydrated prior to dissecting and measuring. Inflorescence and floral bracts were splayed out to measure width as shown in Figure E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 93 2. Measurements are presented as ranges compiled from specimens of several populations across the distribution of each species. Descriptive terminology used herein follows that adopted by Simpson (1990,1998,2006). Specimens cited are a selection seen. For types, we use an exclamation mark (!) for those seen as specimens, ‘image! ’ for those seen as images online and ‘w.v. ’ for those not seen. The species concept applied is both morphological and biological, following that articulated by Hopper and Brown (2001), and first used by Darwin (1868). New species are recognised if they grow in populations (i.e. are not aberrant individuals within populations displaying normal variation), if they have traits or a combination of traits not seen in any previously named species, and especially if they grow with previously named species and produce few or no natural hybrids, the latter exhibiting evidence of partial or full sterility. Ecological data, including habitats occupied, were also determined through field survey and summary of the information provided from herbarium specimens examined. We were unable to conduct experimental tests of reproductive interactions of putative species. However, as an independent test of reproductive isolation, we sequenced DNA from each study population to search for genetic divergence or uniformity, as well as explore molecular phylogenetics. To highlight discoveries made through illustration, examples are listed within figure captions below. Distribution maps were compiled on a base phytogeographic map from Gioia and Hopper (2017) showing fioristic districts and provinces for the SWAFR. Locations came from label details of specimens held at PERTH and determined by the senior author in line with this taxonomic revision. Life Cycle and Comparative Morphology Seedlings (Figure 3). Little has been published on seed biology and germination requirements. However, T. australis seed are recorded as germinating on average within 65 days of sowing, ranging from 55 to 103 days (Fryer 2006). Contractile roots of the germinating seedling pull the apex down into the soil soon after germination Corms and roots. Colourless roots lacking rhizosheaths (Smith et al. 2011) are produced in the autumn just above the corms at a depth of 2-5 cm below soil level. Roots are simple (T. variabilis) or may have a covering of fine short or long colourless root trichomes, with the long trichomes having some sand-binding ability (T. minor and T. purpurea). Aerne-Hains and Simpson (2017) described the root anatomy of T. australis (? T. variabilis). They found the endodermal cell wall of the roots is not thickened. The ground tissue of the central vascular cylinder consists of small isodiametric cells, with thin cell walls. These isodiametric cells have peripheral globular tanniniferous deposits present. The roots have two xylem poles (diarch) with a single large vessel fianked by 10-12 small vessels on either side and only two groups of phloem alternating between the xylem archs. Roots wither during the onset of dry soil in late spring (Pate & Dixon 1982; Figure 3). Corms provide the sole organ of perennation over the dry summers. Parent corms are annually replaced, globoid to ovate, white, fully enlarged to 0.5-1.5 cm diameter by the end of spring as above ground parts wither and dry. In winter a replacement daughter corm is produced alongside the parent corm, usually at the end of a shortly descending dropper stem. Protection is afforded by a pale brown scarious covering remaining from the previous-year’s corms and leaf bases. 94 Nuytsia Vol. 30 (2019) PLANT HEIGHT ^6 Figure 2. Caption opposite. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 95 Flowering. Flowering has been claimed to be obligately fire-stimulated for all Tribonanthes species (Lament & Downes 2011). However, several species found on granite outcrops fiower in the absence of recent fire (e.g. T. australis, T brachypetala, T monantha, T purpurea, T uniflora, T variabilis, T. violacea), as does the salt-lake endemic T. minor and species of ephemeral ponds such as T. keigheryi. Facultative flowering after fire is seen in species when they grow in other damplands such as T. brachypetala and T. violacea on paperbark {Melaleuca spp.) fiats. Aerial stem (Figure 4). The aerial stem is terete in cross section. It is glabrous from the basal leaf to the infiorescence bract, although sometimes there are scattered trichomes near the base of the infiorescence bract attachment, and then it is covered in woolly white trichomes above the infiorescence bract attachment to the base of the infiorescence, except T. purpurea which is glabrous throughout. Tribonanthes brachypetala and T. variabilis also possess a cauline leaf on the stem between the basal leaf and the infiorescence bract. Aeme-Hains and Simpson (2017) found in T. australis (? T. variabilis) that the aerial stem has a cortex of two to several cells thick, consisting of irregularly shaped cells. A sclerenchyma cylinder is present consisting of two to several cell layers, distinct from the adjacent cortical cells at the outer edge and grading into the parenchyma cells at the inner edge. Vascular bundles are inside the sclerenchyma cylinder, randomly arranged throughout, extending into the parenchyma but not found in the centre of it. Vascular bundles are often surrounded by sclerenchyma. Basal leaves (Figure 4). These are 1.5^6 cm long (1.5-2.5 cm in T. minor and 2.5-5.5 cm in T. purpurea), basal (sub-basal to 7.5 cm above ground in T. porphyrea), narrowly linear to filiform (e.g. T. keigheryi), distinctly flat (e.g. T. longipetala), otherwise terete, with a long open basal sheath. The basal sheath has a narrow membranous margin, colourless or tinged pink to deep purple-red and no fringing trichomes. In Aeme-Hains & Simpson’s (2017) study of the vegetative anatomy of Haemodoraceae, the epidermis of the leaf of Tribonanthes australis (? T variabilis), T brachypetala and T longipetala (? T uniflora) consisted of a single layer of cells. The epidermal cells are axially elongated, being longer than wide, with the cell body raised relative to the Junction with the adjacent cells. Epidermal papillae are absent. Stomata are scattered across the leaf surface with each stomate having two paracytic subsidiary cells. The vascular bundles are arranged in a ring at the junction of the palisade and spongy mesophyll. Within the vascular bundles the xylem consists of two to 10 layers of vessels and phloem consists of two to eight layers of sieve members. Tribonanthes lack fibres enveloping the vascular bundles, which are usually present in the Haemodoraceae (Aerne-Hains & Simpson 2017). The cells of the bundle sheath are irregular and not surrounded by sclerenchyma. The palisade region contains scattered tannin cells. The leaves are fistulose with the spongy mesophyll containing aerenchymous cells, with the exception of T minor and T purpurea. The lack of fibres and presence of the aerenchyma cells in the leaves of Tribonanthes are consistent with features present in species that dwell in habitats that are at least periodically wet (Aerne-Hains & Simpson 2017). Cauline leaves (Figure 4). Tribonanthes brachypetala and T variabilis (not illustrated in Figure 2) often possess one or two cauline leaves between the basal leaf and inflorescence bract. It is shorter than the basal leaf, with a broad sheathing base and becoming terete above. The basal sheath has a narrow membranous margin, colourless or tinged pink to deep purple-red and no fringing trichomes. The apex is sometimes mucronate. Figure 2 (opposite). Diagram of a Tribonanthes plant indicating quantitative characters measured (not to scale). A- whole plants including above and below ground parts; B - flower viewed from top; C - longitudinal section through flower; D - stamen showing anther and stamen connective appendage; E - inflorescence bract (splayed out) showing transparent margins; F - floral bract (splayed out) showing transparent margins; G - longitudinal section through underground corms showing this seasons corm, last seasons shrivelled corm and downward extension of stem forming next seasons corm. Illustration by E.J. Hickman. 96 Nuytsia Vol. 30 (2019) Figure 3. Seedling development and seasonal phenology of Tribonanthes. A - germinating seed of T. keigheryi (CT - cotyledon, P - plumule, SC - seed coat, R - radicle, RH - root hairs); B - seedling morphology of T. variabilis, showing mode of formation and burial of first-season’s storage organs. Stages in seedling development are arranged in numbered sequence, the last stage showing the likely condition of the seedling on entering dormancy at the end of the first season of growth (CT - cotyledon; P- plumule; R-radicle; LI - seedling leaf; SA - stem apex; SC - seed coat; D - ‘dropper’ shoot; ST - stem tuber). Ground level marked by a dotted line. (From Pate & Dixon 1982: 22-23); C - phenology of growth, dry matter production and reproduction in the geophyte T. variabilis. Morphological changes accompanying growth are recorded and provide a key to the symbols for plant organs used in the cumulative plots of change in dry matter of plant organs. A single parent stem tuber (stippled and marked PT) is replaced by a single replacement tuber (RT) (R - root; L - leaf; IS - inflorescence stem; F - flowers; S - seed). Total rainfall for the year of harvest was 923 mm. Long-term rainfall average for harvest site was 873 mm. (From Pate & Dixon 1982: 168). Image by R.J. Smith, Royal Botanic Gardens, Kew (A). E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 97 Inflorescence bract (Figure 4). The inflorescence bract is solitary. It is shorter than the basal (and cauline) leaf (0.6-13.8 cm). The bract is usually well-separated from basal leaves to a third or more up the stem, with a broad sheathing base and becoming shortly terete above. The apex is sometimes mucronate. The basal sheath has a broad membranous margin, and is colourless or tinged pink, lilac, or deep purple-red. In T. australis, T keigheryi, T porphyrea and T. uniflora the bract has trichomes along the margin. Peduncle. The peduncle is the structure between the base of the inflorescence bract and the base of the floral bract (or lowest floral bract), with the exception of T. longipetala, with its conspicuous branching, where the upper limit of the peduncle is the lowest branching point (F igure 2). T. longipetala is anomalous in its open inflorescence with wide branching, which usually lacks a bract at the lower branch points. The base of the peduncle is the node at the base of the inflorescence bract. Pedicel (Figure 4). Pedicel from the insertion of the floral bract to the base of the ovary. Pedicels are absent (e.g. T. minor) or vary in length up to 30 mm (e.g. T. longipetala). Floral bracts (Figure 4). Inflorescences are either a solitary flower or a compact or loosely arranged cyme. A floral bract subtends each flower. The floral bracts are elliptic to orbicular. They have a terete tip which is sometimes mucronate. There are three to five prominent veins, broad membranous margins that are pale green or tinged pink or lilac, and fringed with colourless trichomes, except T. purpurea which is glabrous. Flowers andperianth (Figure 5). Flowers are actinomorphic, slightly zygomorphic in T. minor, tubular, and with six perianth lobes. The lobes vary from narrow linear to broadly obovate. The perianth lobes can be longer than, equal or shorter than the perianth tube. They can be erect, spreading or strongly reflexed. The perianth tube is covered in silky or woolly white trichomes, sometimes tinged purple on the outer surface but the inner surface is glabrous and ranges in colour from white, pale green to a deep yellowish green. The perianth lobes are covered in silky or woolly white trichomes, sometimes tinged purple on both outer and inner surfaces, sometimes the centre of the outer surface has less trichomes and can have a central broad stripe that is either green or purple in colour. The exception is T. purpurea which has pink, glabrous flowers. Stamen connective appendages (Figure 6). A distinctive feature of Tribonanthes is the presence of stamen connective appendages (Simpson 1990). Detailed developmental studies and homology issues require further studies to determine the anatomical origins of these structures. Their function is completely unknown. The stamen connective appendage includes the filament and the appendage, which are not easily distinguishable. The appendages consist of either two small lateral points at the top of the filament (e.g. T. purpurea, T. keigheryi), or a spreading apex of either short dentate points or more petal-like structures, sometimes with longitudinal ridges on the back (abaxial) surface, or a cluster of fleshy finger-like structures {T. minor, T. monantha), or a broad fleshy structure with dentate top and longitudinal ridges on the back surface (T. brachypetala). The stamen connective appendages are either white to cream or bright yellow in colour under natural light, aging to deep orange. The stamen connective appendages can be shorter than, equal to or project above the anthers, collectively forming a distinct corona-like structure. Anthers (Figure 6). The anthers are yellow, except in T. minor in which they are cream. In terms of location of the anther on the stamen connective appendage, they appear centrally placed, or less frequently occupy the apical half (i.e. T. keigheryi, T. purpurea, T. violacea), or the basal half (i.e. 98 Nuytsia Vol. 30 (2019) T. minor, T monantha). The locules are free below the attachment and fused above with a sterile apex, whose function is unknown. They dehisce introrsely through longitudinal slits. Pollen. Tribonanthes has globose-spherical pollen grains usually with 5 flattened apertures (sometimes 6-7) and scattered exinous elements (Simpson 1983; Pierce & Simpson 2009). Styles and stigmas (Figure 6). The style is usually sub-sessile or short, stout and green, or with purple longitudinal striations, except T. purpurea where it is slender-flliform and elongated. The stigma is terminal, simple or has a prominent tuft of silky white trichomes atthe apex(i.e. T. elongata, T longipetala, T variabilis, less so in T. porphyrea), or three protuberances at the apex (i.e. T. brachypetala, less so in T. porphyrea). Nectaries. Simpson (1993) described T. variabilis as having three supralocular septal nectaries primarily traversing the exposed portion of its half-inferior ovary. They open to the outside at the extreme distal end of their extent, near the base of the style. They are composed of a single layer of radially elongated nectar-secreting epithelial cells. Ovary position influences the type and extent of nectaries, so other species of Tribonanthes may have different nectaries depending on their ovary position (Simpson 1993). Ovary (Figure 7). The ovary is superior (i.e. T. minor and T. purpurea), inferior (i.e. T. elongata and T. longipetala) or otherwise half-inferior, except T. variabilis which can be half-inferior to inferior. The ovary consists of three locules. All species have axile placentation. The number of ovules per locule illustrated in Figure 7 was counted from available botanical drawings herein. It varied between different species: T. brachypetala (6 ovules); T. purpurea (7); T. longipetala (9); T. minor (10-12); T. keigheryi (14); T. uniflora (15); T. monantha (25); T. elongata (26); T. variabilis (26); T. australis (48); T. violacea (55); and T. porphyrea (78). Tribonanthes brachypetala displays early abortion of some ovules resulting in only a few seeds per capsule. Capsule (Figure 7). All species have 3-valved capsules. The ovary swells as the seeds enlarge. The perianth is persistent, with the perianth lobes remaining reflexed or becoming erect, except T variabilis where the perianth lobes remain spreading, prior to dehiscence. The perianth lobes then become reflexed post-dehiscence, except in T purpurea where they are spread slightly from the erect position. When mature the capsule splits loculicidally from the apex to a third or half way down the central ridge of each carpel. Seeds (Figure 7). Seeds of all species except T minor are 1.0-2.2 mm in length, irregularly ovoid or cuboid with angular folds and a few ridges. The seed shape is affected by compression between neighbouring seeds during development. The seeds are brown tending to purple-grey, glabrous, with a starchy endosperm and a minute embryo near the micropyle. Those of T minor are noticeably diminutive, measuring 0.4-0.5 mm in maximum length. The seeds have no specialised means of dispersal and appear to be released as capsules are shaken by wind and rain. Presumably the seeds do not fall far from the parent plant. Chromosomes. Simpson (1990) reportedN=7 for an unknown Tribonanthes species. This chromosome count was attributed to “G. J. Keighery pers. comm.” However, Hopper and Stone (cited in Hopper etal. 2006) obtained N = 11 for Z variabilis from Mt Benia and Toodyay Road. Moreover, G.J. Keighery (pers. comm. 2005) has no record of a count of N = 7 for Tribonanthes. The report of N = 7 for the genus was an error in Simpson (1990) (Simpson pers. comm. 2005). Subsequently, cytological collections by EJH of most species in the genus failed to yield squash preparations of pollen mother cells and root tips suitable for chromosome counts. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 99 Figure 4. Caption on page 103. 100 Nuytsia Vol. 30 (2019) Figure 5. Caption on page 103. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 101 Figure 6. Caption on page 103. 102 Nuytsia Vol. 30 (2019) Figure 7. Caption opposite. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 103 Figure 4 (page 99). Whole plant specimens of the twelve Tribonanthes species to scale (dashed line indicates ground level). A - T brachypetala (no voucher, from Millinup Rd, south side of Porongurup Range after a fire in 2010); B - E australis {EJ. Hickman 2067); C-T. violacea {E.J. Hickman 2090); D - E purpurea {EJ. Hickman 2083); E-T. uniflora {E.J. Hickman 2084); E -T. minor {E.J. Hickman 2079); G-T. longipetala (E.J. Hickman 2021); H - E monantha (holotype E.J. Hickman 2048; \-T. porphyrea (holotype E.J. Hickman 2025); J - E elongata (holotype E.J. Hickman 2073); K - 7^ keigheryi (holotype E.J. Hickman 2065); E-T. variabilis (E.J. Hickman 2053). Scale bar = 20 mm. Discoveries through illustrations for particular species include the variation in peduncle length (species C, D, I and K have little to no peduncle, while all other species have relatively long peduncles); the sub-basal leaf up to 7.5 mm above ground level (species I). Illustrations by E.J. Hickman. Figure 5 (page 100). Flowers of Tribonanthes species viewed from top or face-on (A-F) and with half perianth removed to show ovary position, stamen, stamen connective appendages and style (M-X). A -T. australis', E-T. monantha, C-T. purpurea, E)-T. elongata, E-T. unflora, E-T. minor, G-T. longipetala, Ei-T. brachypetala, \-T. variabilis', i -T. porphyrea, K - T. violacea, E-T. keigheryi. M - 7! australis', N - F monantha, 0-T. purpurea, E-T. elongata, Q-T un flora', R-T. minor, S - T. longipetala, E-T. brachypetala, G-T. variabilis', Y - T. porphyrea, W - T. violacea, X- T. keigheryi. Scale bar = 2.5 mm. Discoveries through illustrations for individual species include: difference in ratio of length to width of perianth lobes (particularly A>I and G>J distinguishing these species pairs); position of ovary (inferior- P, S, half-inferior - M, N, Q, T, U, V, X, superior - O, R, half- inferior to inferior - W). Voucher specimens as cited in caption for Figure 4. Illustrations by E. J. Hickman. Figure 6 (page 101). Stamen and styles of Tribonanthes species. A - stamen viewed from front; B - stamen viewed from back; C - stamen viewed from side; D - style and stigma; aus - T. australis', bra - T. brachypetala, elo - T. elongata, kei - T. keigheryi. Ion - T. longipetala, min - T. minor, mon - T. monantha, por - T. porphyrea, pur - T. purpurea, uni - T. unflora, var - T. variabilis', vio - T. violacea. Scale bar = 1 mm. Discoveries through illustration for particular species include; colour of stamen connective appendages (cream - aus, uni, mon, por, pur, kei, var, vio; yellow - bra. Ion, min, elo); colour of anthers (yellow - aus, bra, uni. Ion, mon, por, pur, elo, kei, var, vio; cream - min); position of attachment of anther on stamen connective appendage (central - aus, bra, uni. Ion, por, elo, var; lower - min, mon; upper - pur, kei, vio); trichomes on inner surface of filament (elo); relative length of style (sub-sessile - aus, bra. Ion, min,mon, por, elo, var; short - uni, kei, vio; long - pur); style colour (green - bra, uni, min, por, pur, var; purple striations - aus. Ion, mon, elo, kei, vio); stigma colour (yellow - aus, bra, uni, min, mon, por, pur, kei, vio; green - Ion, elo, var); stigma vestiture (none - aus, uni, min, mon, pur, kei, vio; tuft of silky white trichomes - Ion, por, elo, var; three protuberances - bra, por). Voucher specimens as cited in caption for Figure 4. Illustrations by E.J. Hickman. Figure 7 (opposite). Capsules and seeds of Tribonanthes species. A - immature capsule; B - T.S. capsule; C - F.S. capsule; D - dehisced capsule; E - seeds from side top and sectioned to show seed volume; aus - T. australis', bra - T. brachypetala, elo - T. elongata, kei - T. keigheryi. Ion - T. longipetala, min - T. minor, mon - T. monantha, por - T. porphyrea, pur - T. purpurea, uni - T. unflora, var - T. variabilis', vio - T. violacea. Scale bars = 5 mm (A, B, C, D); 1 mm (E). Discoveries through illustration for particular species include persistent perianth position before dehiscence (erect - aus, uni, mon, por, kei, vio, pur; splayed- Ion, elo, var; refiexed - bra, mon); persistent perianth position after dehiscence (refiexed - aus, uni. Ion, mon, por, elo, kei, var, vio, bra, min; upwardly splayed- pur); ovules per locule (<10 - bra, pur. Ion; 10-25 - min, kei, uni, mon; 25-50 - elo, var, aus; >50 - vio, por); the relative size of seeds particularly the diminutive Tminor seeds. Voucher specimens as cited in caption for Figure 4. Illustrations by E.J. Hickman. Molecular Phylogeny Early analysis of the molecular phylogeny of the Haemodoraceae included a few species of Tribonanthes (Hopper et al. 1999, 2009). Some of the identifications in these studies need updating in light of this present revision. This update is being done, and a more comprehensive study of all twelve species at up to eight populations each is underway. Current evidence unequivocally establishes the monophyly of Tribonanthes as a genus (Hopper et al. 2009). Morphological, ecological and current unpublished molecular studies indicate strong support for recognition of T minor and T purpurea as monotypic subgenera. It seems clear that further species-level resolution of the phylogeny of Tribonanthes and a clearer understanding of the role of natural hybridization in the genus will require next generation sequence data. 104 Nuytsia Vol. 30 (2019) Taxonomy Tribonanthes Endl. in S. Endlicher & E. Fenzl, Nov. Stirp. Dec. 4: 27 (1839). Type: Tribonanthes australis Endl. Herbs, perennial terrestrial geophytes, erect (usually) but stemless or nearly so in T. purpurea, to 41 cm tall. Corms annually replaced at the end of a short, vertically descending dropper emergent near the top of the parent corm, globoid to ovate, white, to 0.5-1.5 cm diameter when fully enlarged, ensheathed with brown scarious papery multi-layered tunic. Roots annually replaced, emergent from the top of the corm, colourless, sinuous, lacking a rhizosheath, or sometimes with a loose sand-binding rhizosheath {T. longipetala), c. 1 mm diameter. Eeaves l(-3), usually 1 basal, occasionally additional leaves on aerial stem (E brachypetala, T. variabilisy, sheath cylindrical, closely (or loosely) surrounding the stem, dilated and funnel-like in upper part, with margins fused (or free); lamina terete, sometimes slightly grooved or flat, narrowly linear. Inflorescence terminal, a single flower, a compact or loose cyme, or bifurcate pair of helicoid cymes; axes lanate, bracts usually glabrous with hairy margins. Flowers actinomorphic, slightly zygomorphic in T. minor, white, sometimes tinged purple, rarely purplish pink (in T. purpurea). Perianth shortly tubular, white-lanate, the 6 lobes imbricate, divergent or reflexed at anthesis. Stamens 6, inserted at base of the perianth lobes; filament connective extended upwards as an apically lobed, fleshy, yellow or cream flange often irregularly serrate at top; anthers dorsiflxed with sterile apex, dehiscence longitudinal. Ovary inferior, half-inferior or superior, globose, with few to numerous ovules (c. 6-80 per locule), anatropous or heterotropous, the roof green. Style broadly conical or slender-filiform; stigma capitate, yellow or green, obscurely 3-lobed with elongate apex sometimes tufted with short trichomes or topped with 3 protuberances. Capsules enlarged, ellipsoidal, loculicidally dehiscent. Seeds globose or cuboidal angular, often irregular and variable in the one capsule, purplish grey-brown. Etymology. From the Greek tribon (threadbare cloak) and anthos (flower), which could refer to the cobwebby-woolly indumentum of the flowers of most species (Macfarlane 1987) or to the stamen connective appendages ‘which rise above the anthers’ (Sharr 1996: 69). Notes. Tribonanthes is ataxonomically isolated genus within Haemodoraceae subfamily Conostylidoideae (Eindl.) T.Macfarlane & Hopper, placed in its ownmonotypic tribe Tribonantheae (Macfarlane & Hopper 1987; Simpson 1990; Hopper et al. 1999, 2009; Smith et al. 2011). The 12 species share a common ecology as inhabitants of seasonally water-logged or shallowly inundated clay-based wetlands or seeps and moss mats on granite outcrops. These are often old, climatically-buffered infertile landscapes (Hopper 2009; Hopper et al. 2016), refugial habitats from today’s strongly seasonal Mediterranean climate, implying that ecological niche conservatism is a hallmark of Tribonanthes. The genus is most diverse in higher rainfall provinces of the SWAFR (Gioia & Hopper 2017), with a few species penetrating further inland in locally mesic habitats afforded by granite outcrops or seasonal damplands. Occupying such habitats are the two smallest species, the granite pink (E purpurea) and the diminutive salt-lake tiurndin (E minor). These are the only taxa of conservation concern. Their population structures are quite insular, and their habitats at some risk to disturbance processes affecting granite outcrops and freshwater rises on the margins of salt lakes respectively. In particular, threats include grazing by native mammals and feral rabbits and pigs, mechanical disturbance of granite moss mats by recreational or mining vehicles, the risk of salinization due to land clearing and climate change, and invasion of habitats by weeds. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 105 Based on our field, morphological and molecular studies it is reasonable to recognise three subgenera in Tribonanthes, two of which are monotypic; they are keyed out below. Key to subgenera of Tribonanthes 1. Perianth white with spreading or reflexed lobes 2. Aerial stem up to 3 cm tall.subg. Salina 2: Aerial stem longer than 5 cm tall.subg. Tribonanthes 1 : Perianth purple-pink with erect lobes.subg. Boya Key to species of Tribonanthes Note. Species are listed by relatedness, based on our morphological studies and initial results from continuing molecular studies, and numbered consecutively below for the entire genus. 1. Inflorescence multi-flowered 2. Stamen connective appendages white to cream 3. Stigma glabrous or few scattered trichomes. 1. T. anstralis 3: Stigma with tuft of trichomes. 8. T. variabilis 2: Stamen connective appendages yellow 4. Perianth lobes spreading. 3. T. longipetala 4: Perianth lobes strongly reflexed. 10. T. brachypetala 1: Inflorescence 1 (-2) flowered 5. Stamen connective appendages yellow 6. Perianth lobes spreading. 6. T. elongata 6: Perianth lobes strongly reflexed. 11. T. minor 5: Stamen connective appendages white to cream 7. Stamen connective appendages residual 8. Perianth lobes erect, glabrous. 12. T. purpurea 8: Perianth lobes spreading, hairy 9. Inflorescence bract sheath margin hairy, fine slender plant, swamp dweller.7. T. keigheryi 9: Inflorescence bract sheath margin glabrous, robust plant, granite dweller.9. T. violacea 7: Stamen connective appendages prominent 10. Stigma with tuft of trichomes. 5. T. porphyrea 10: Stigma glabrous or few scattered trichomes 11. Inflorescence bract sheath margin glabrous. 4. T. mouautha 11: Inflorescence bract sheath margin hairy. 2. T. uuiflora 106 Nuytsia Vol. 30 (2019) Tribonanthes Endl. subg. Tribonanthes Aerial stem more than 5 cm tall. Flowers erect, solitary or few per inflorescence. Perianth with white spreading lobes, or with green reflexed lobes. Style 0.4-3.2 mm long. Notes. A subgenus of ten species regarded as the typical tiurndins with white lanate erect (rarely nodding, i.e. T. brachypetala) star-like flowers. Found on granite outcrops or seasonal damplands or in shallow pools. 1. Tribonanthes anstralis Endl. in S. Endlicher & E. Fenzl, Nov. Stirp. Dec. 4: 27 (1839). Type: King George Sound, Western Australia, C. Hugel s.n. (holo: W, n.v. - destroyed in Are during World War II [E. Vitek pers. comm. 6 May 2016]). Neotype - illustration in S. Endlicher, Icon. Gen. PI. t. 109 (1841), here chosen (Figure 1). Illustrations. S. Endlicher, Icon. Gen. PI. t. 109(1837^ 1) - Fig. 1.; A. Schnizlein Iconogr. i. t. 62 Fig. 4 (1843); WE. Blackall & B.J. Grieve, How to Know W. Austral. Wildfl. Part I, p. 75 (1954), reprinted as Parts I, II, III p. 75 (1974); J. Scott & P. Negus Wildfl. Aust. S. W. p. 44 Fig. 5 (2002). Herb annually renewed from a white ovoid corm, 17-41 cm tall, consisting of a single stem and single basal leaf Corms 10-12.5 mm diameter, covered by numerous old brown, papery sheaths, 3.2-8.9 cm below soil surface, white, simple with no visible roottrichomes and no sand-binding rhizosheath. Aerial stem (from basal leaf to inflorescence bract) 2.6-29.4 cm, glabrous. Basal leaf ^-23 cm long, erect, stem-clasping to terete, red-purple base grading to deep green apically, with dark mucronate tip, glabrous. Inflorescence bract 2.7-10.8 cm long, erect, funnel-like sheath with broad dilated mouth, 6-10 mm circumference at widest point, green with noticeable longitudinal veins and thin scarious margins fringed with white trichomes; lamina 10-61 mm long, terete, green, with darker mucronate tip, glabrous. Peduncle (from inflorescence bract to floral bract) 7.5-26 cm long, covered in white woolly trichomes, sparser towards inflorescence bract. Inflorescence a compact cyme of 2-5 sessile or shortly pedicellate flowers, each subtended by a single floral bract. Floral bract ovate, 5-11 mm long, 2-7 mm wide, greenish-purple with prominent longitudinal veins, hairy on margins and at base outside, glabrous inside, with a short terete mucronate apex, 1-2 mm long, ending in a darker tip, not exceeding perianth lobes. Perianth lobes 5-9.1 mm long, 2^. 8 mm wide, spreading, obovate to elliptic, inner surface white hairy, outer surface purple with white trichomes sparser along centre of lobe, tips dark mucronate. Perianth tube 3-7 mm long, outer surface covered in long silky white trichomes, inner surface pale green, glabrous. Stamen connective appendages 2-4 mm long, 1.3-1.5 mm wide, creamy white, topped with 4 or 5 flnger-like projections with rounded tips, equal to slightly exceeding anther tips. Anthers 1.5-3.3 mm long, yellow, with cream sterile apex that projects towards centre of flower, attached centrally on stamen connective appendages. Ovary green, half-inferior, locules 3, with many (48) ovules per locule. Style 0.6-2 mm long, with green and purple longitudinal striations. Stigma yellow, with few scattered trichomes on upper surface. Capsule with perianth persistent, erect before dehiscence, reflexed after dehiscence. Seeds purplish grey-brown, angular. (Figures 1, 8, 9) Diagnostic features. Tall plant, terete leaf, long peduncles, multi-flowered inflorescence, white hairy flowers with faint tinge of purple, large creamy white stamen connective appendages, short to sub- sessile style with simple stigma bearing few scattered trichomes on upper surface, half-inferior ovary. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 107 Selected specimens examined. WESTERN AETSTRAEIA: 7.2 km W of Denmark along South Coast Hwy, 18 Sep. 1991, A.R. Annels 1668 (PERTH); Kangaroo Rd, 4.1 km E of Collis Rd, Hazelvale, 14 Sep. \99AA.R. Annels &R.W. Hearn 4416A (PERTH); banks of the Gordon River, N of Frankland, 13 Sep. 1973, A.M. Ashby 4899 (PERTH); Holland Rocks, 4.4 km from Newdegate-Pingrup Rd along Holland Tank Rd, 17 Sep. 2014, A.M Coates 6735 (PERTH); road to water monitor. Waychinicup, E of Albany, 30 Oct. 1992, E.J. Croxford 6659 (PERTH); Table Hill Forest Block, Bevan Rd Denbarker, 8 Oct. 2007, C.P. Dornan 387 (PERTH); Nollajup Nature Reserve, 8 km SW Boyup Brook, 20 Sep. 2014, E.J. Hickman 2057 (PERTH); Mettabinup Nature Reserve, 3.3 km W on Flora and Fauna Rd from Tone Rd, then 500 m into reserve to SW boundary of reserve, 20 Sep. 2014, E.J. Hickman 2060 (PERTH); Wamballup Fake Nature Reserve, 500 m W on Wamballup Rd from Boyup Rd, S side of road in broad drainage line, 30 Sep. 2014, E.J. Hickman 2066 (PERTH); Mt Melville, 350 m SE from top carpark along Mt Melville Circuit Trail, both sides of trail, 31 Oct. 2014, E.J. Hickman 2067 (PERTH); 20 km S of Bridgetown towards Manjimup, 8 Oct. 1982, G.J. Keighery 5346 (PERTH); Porongurup Range, SW margin, below Halls Rock, 3 Nov. 1986, G.J.Keighery 8481 (PERTH); Moingup Spring, Chester Pass, Stirling Range, 20 Sep. 1987, G.J. Keighery 11382 (PERTH); Symmonds Block, Tuart Forest, W of Eudlow, 15 Sep. 1994, G.J. Keighery 13595 (PERTH); Cowerup Nature Reserve, 23 Oct. 1997, G.J. Keighery & N. Gibson 2749 (PERTH); 3 km E of Windy Harbour, 13 Nov. 1986, M. Prest s.n. (PERTH); ScottNational Park camping reserve west, 17 Sep. 1990, G.J. Robinson 123 (PERTH). Proposed vernacular name. Southern Tiumdin. Phenology. Flowers from August to December. Distribution and habitat. The species ranges from Augusta across the south-west comer of Western Australia to east of Albany, with an out-lying population recently recorded near Hollands Rock. It is mainly concentrated in the Jarrah and Muir Districts of the Bibbulmun Province of south-west Western Australia (Gioia & Hopper 2017), but there are collections from the Fitzgerald-Stirling and Maalak Districts of the Southeast Coastal Province. Tribonanthes australis grows on brown loams around granite outcrops associated with herbfields, shrublands and open woodlands, and on grey sandy clay in winter wet flats associated with shrublands and heathlands adjacent to open woodlands. Associated species include Corymbia calophylla. Eucalyptus rudis. Eucalyptus wandoo, Melaleuca preissiana, Melaleuca viminea, Hakeaprostrata, Taxandria marginata, Verticordiaplumosa, Borya nitida, Drosera menziesii, and Stypandra glauca (Figure 9). Conservation status. Widespread with no special conservation needs. Etymology. Named from the Eatin for southern, alluding to its discovery at King George Sound near the southern-most part of the Swan River Colony. Notes. Macfarlane (1987) synonymised T. variabilis under T. australis. However, while the stigma of T. australis can sometimes have scattered trichomes on the top, the prominent appendage covered in silky white trichomes, present on the stigma of T. variabilis, is absent. Also the perianth of T. australis is more noticeably tinged purple than the perianth of T. variabilis, and the perianth lobes of T. australis are generally narrower than T. variabilis, relative to their length. Tribonanthes australis is usually more south and east in its distribution (Figure 9C) while T. variabilis is found on the west coast (Figure 23C). Young plants of T. australis may appear to have a single-flowered inflorescence, and hence are confused with their smaller relatives, T. monantha and T. uniflora. Tribonanthes australis is identified as a herbaceous species in the floristic summary of the Melaleuca cuticularis/M.preissiana Open 108 Nuytsia Vol. 30 (2019) Figure 8. Tribonanthes australis. A - whole plant; B - bud; C - flower from top; D - flower from side; E - flower with part of perianth removed to show ovary and style; F - stamens from front, side and back view showing creamy white stamen connective appendages exceeding anthers and yellow anthers with apex apiculate and projecting forwards; G - style sub-sessile with simple stigma; H - capsule; I - cross section of capsule; J - longitudinal section of capsule; K - dehisced capsule; L - seeds; M - leaf section near apex; N - leaf section near base; O - inflorescence bract; P - floral bract; Q - longitudinal section of corm; R - root; S - cross section of root; T - detail of perianth trichomes. Scale bars = 1 cm (A); 2.5 mm (B, C, D, E, H, I, J, K, P, R); 1 mm (F, G, M, N); 4 mm (O, Q); 0.5 mm (E, S); 0.65 mm (T). Drawn from fresh material of E.J. Hickman 2067 (PERTH 08989354). Illustrations by E.J. Hickman. E. J. Hickman & S .D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 109 Figure 9. Tribonanthes australis. A - plants showing multi-flowered inflorescences {E.J. Hickman 2067); B - habitat {E.J. Hickman 2067), shrubland of Taxandria marginata and Verticordia phimosa associated with herbfleld of Stypandra glauca and Drosera menzesii over granite on south-west slopes of Mt Melville, Albany, Western Australia; C - distribution in south-western Australia with SWAFR provinces and districts shown in grey (Kalbarri Province: SB - Shark Bay District, Nan - Nanda District; Bibbulmun Province; Les - Lesueur District, Jar - Jarrah District, Nar - Narrogin District, Mui - Muir District; Southeast Coastal Province: Maa - Maalak District, F-S - Fitzgerald-Stirling District, Boy - Boylya District, Esp - Esperance District; Transitional Rainfall Province: Won - Wongan District, Mer - Merredin District, Hyd - Hyden District (Gioia & Hopper 2017). Photos by E.J. Hickman. 110 Nuytsia Vol. 30 (2019) Woodland vegetation association, of the Albany Regional Vegetation Survey (Sandiford & Barrett 2010). 2. Tribonanthes uniflora LindL, Sketch Veg. Swan R. xliv (1840). Type: Swan River, Western Australia, J Mangles s.n. (lectotype: CGE 06825! fide T.D. Macfarlane, FI. Australia 45: 465 (1987)). Illustrations. WE. Blackall & B.J. Grieve, How to Know W. Austral. Wildfl. Part I, p. 75 (1954), reprinted as Parts I, II, III p. 75 (1974); J.R. Wheeler in J. Wheeler, N. Marchant & M. Eewington, Flora 5. IE Vol. 1: 314 (2002). Herb annually renewed from a white ovoid corm, small to medium plant, 6.5-21.4 cm tall above ground, consisting of a single stem and single leaf Corms 4-8 mm diameter, covered by numerous old brown, papery sheaths, 1.2-5.3 cm below soil surface. Roots white, no root trichomes, no sand¬ binding rhizosheath. Aerial stem (from basal leaf to inflorescence bract) 1.5-15 cm, glabrous. Basal leaf 4-16.5 cm long, erect, stem-clasping to terete, red-purple base grading to green apically, with mucronatetip, glabrous. Inflorescence bract 1.4-6.4 cm long, erect, funnel-like sheath with broad dilated mouth, 4-8 mm circumference at widest point, red-purple, broad membranous margin tinged red and fringed with white trichomes, lamina 5-35 mm long, terete, green, with mucronate tip. Peduncle (from inflorescence bract to floral bract) 0.4-7.7 cm long, covered in white woolly trichomes. Inflorescence a solitary flower, subtended by one or two floral bracts. Floral bract obovate, 7-13 mm long, 3-8 mm wide, green with purple margin fringed with white trichomes and scattered trichomes on the base, with a short terete mucronate apex, 1-5 mm long, ending in a darker tip, exceeding perianth lobes. Perianth lobes 5-9.4 mm long, 1.5-3.2 mm wide, spreading, obovate, with white hairy inner and outer lobe surfaces, dark mucronate tips. Perianth tube 2.8-7.4 mm long, outer surface covered in long white woolly trichomes forming a distinctly fluffy skirt, inner surface white, glabrous. Stamen connective appendages 0.5^ mm long, 1.2 mm wide, creamy white topped with 3 flnger-like projections with rounded tips, equal to slightly exceeding anther tips. Anthers 1.5-3.4 mm long, yellow, with cream sterile apex that projects towards centre of flower, attached centrally on stamen connective appendages. Ovary green, half-inferior, locules 3 with several (15) ovules per locule. Style 0.7-2.5 mm long, green. Stigma simple, capitate, yellow, glandular, glabrous. Capsule greyish brown; perianth persistent; lobes erect before dehiscence, spreading to reflexed after dehiscence. Seeds purplish grey-brown, angular. (Figures 10, 11) Diagnostic features. Small to medium plant, terete leaf, long woolly peduncles, solitary flowered inflorescence, woolly white flowers, obvious cream stamen connective appendages, medium style with simple stigma, half-inferior ovary. Selected specimens examined. WESTERN AUSTRAEIA: Guildford, Sep. 1901, C.Andrews s.n. (PERTH); Coomalbidgup, 4 Aug. 1965, J. Bowen s.n. (PERTH); Wandoo Conservation Park, Goonapping Swamp, 17 Sep. 2014, K.L. Brown & G. Paczkowska KEB 1153 (PERTH); N side of Eake Powell, W of Albany, 25 Sep. 1983, D. Davidson s.n. (PERTH); Cowalellup Rd, 1.2 km W of intersection with Boxwood Hill-Ongerup Rd, S side of road, 29 Oct. 2013, E.J. Hickman 2010 (PERTH); McNeil Rd, 2.6 km E of Kellerberrin-Voting Rd, 100 m S of road side, 12 Sep. 2014, E.J. Hickman 2049 (PERTH); Moses Rock, 150 m S of T-junction at end of Moses Rock Rd, 50 m W to granite rock, 20 Sep. 2014, E.J. Hickman 2055 (PERTH); Mettabinup Nature Reserve, 3.3 km W along Flora and Fauna Rd from Tone Rd, then 500 m W into reserve to SW boundary, 20 Sep. 2014, E.J. Hickman 2061 (PERTH); Jingalup Nature Reserve, 1.1 km W of Kojonup-Frankland Rd on Settlers Rd, S side E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 111 of road in claypan, 20 Sep. 2014, EJ. Hickman 2062 (PERTH); Break Rd, 2.6 km W of turn-off to Mt Lindesay, N side of road, 3 Nov. 2014, EJ. Hickman 2071 (PERTH); Moses Rock, 150 m S of T-junction at end of Moses Rock Rd, 50 m W to granite rock, 16 Aug. 2016, E.J. Hickman 2084 (PERTH); 8 km S of Eneabba, 27 Sep. 1977, R. HnatiukllU96 (PERTH); Kemerton, 2 Sep. 2003, G.J & B.J. Keighery 261 (PERTH); Pinjarra Nature Reserve, 30 Aug. 2007, G.J. Keighery 17134 (PERTH); 17.2 km W of Northampton on Port Gregory Rd, 25 July 2009, G.J. & B.J. Keighery 1569 (PERTH); Warrenup Nature Reserve, W of Kendenup, 19 Sep. 2009, G.J. Keighery 17561 (PERTH); Plain Rd, NW of Walpole, 21 Sep. 1983, R.Orndujf 9329 (PERTH); Mt Chudalup, 5 Sep. 1985, J.R. ffTzee/er 2389 (PERTH). Proposed vernacular name. Woolly Tiurndin. Phenology. Flowers from August to October. Distribution and habitat. Tribonanthes uniflora is known from scattered populations both coastal and inland, from north of Geraldton to west of Esperance. Most collections are from the Jarrah, Muir and Eesueur Districts of the Bibbulmun Province of south-west Western Australia (Gioia & Hopper 2017), but there are collections from the Nanda District of the Kalbarri Province, Wongan District of the Transitional Rainfall Province and Maalak District of the Southeast Coastal Province. It grows on seasonally wet brown to grey sandy clays in moss swards and shallow soil pockets of granite sheets and outcrops or on winter wet flats associated with Melaleuca shrublands, open woodlands and forest fringes around granite. Associated species include Eucalyptus occidentalism E. rudis, Melaleuca preissiana, M. hamulosa, M. lateritia, M. uncinata, M. vimineam Hakeaprostrata, H. varia, Verticordia plumosa, Dodonaea ceratocarpa,Xanthorrhoea preissii and a variety of herbs and sedges (Figure 11). Conservation status. Widespread with no special conservation needs. Etymology. From the Eatin unus (one) and florus (flowered), referring to the single-flowered inflorescence of this species. Notes. Tribonanthes uniflora collections have previously been identified as T. australis, T. longipetala or T. violacea. Tribonanthes uniflora differs from T. australis in having a smaller habit and a smaller, single-flowered inflorescence. It differs from T. longipetala in having terete leaves and shorter perianth lobes and obvious cream stamen connective appendages, rather than bright yellow stamen connective appendages. It differs from T. violacea in having an obvious peduncle, which is covered in woolly trichomes from the inflorescence bract to the first floral bract. This species is similar to T. monantha in its small stature and single-flowered habit but differs in having an inflorescence bract with a narrower transparent margin fringed with white simple trichomes. Also the flowers of T. monantha tend to turn purple at senescence while T. uniflora flowers remain white. 3. Tribonanthes longipetala Eindl, Sketch Veg. SwanR. xliv (1840). Type'. Swan River [Colony], Western Australia, 1839, J. Drummond s.n. (lecto here chosen: CGE 06824! left specimen; isolecto: K 000846201 image!); Swan River, Western Australia, J. Mangles s.n. {syn\ CGE 06824! right specimen). TribonantheslindleyanaEndl. inEehm.,P/. Preiss. 2: 27 (1846), nom. illeg., nom. superfl. Type: Swan River [Colony], Western Australia, 1839, J.A.L. Preiss 1561 {holo\ ED 1811505 image!; iso\ MEE 104294 image!, P 01698194 image!, S 06-11224 n.v). 112 Nuytsia Vol. 30 (2019) Figure 10. Tribonanthes uniflora. A - whole plant; B - bud; C - flower from top; D - flower from side; E - flower with part of perianth removed to show ovary and style; F - stamens from front, side and back view showing stamen connective appendages small, just shorter than anthers and yellow anthers with apex apiculate and projecting forwards; G - style slightly elongated with simple stigma; H - capsule; I - cross section of capsule; J - longitudinal section of capsule; K - dehisced capsule; L - seeds; M - leaf section near apex; N - leaf section near base; O - inflorescence bract; P - floral bract; Q - longitudinal section of corm; R - root; S - cross section of root (scale bar = 0.5 mm); T - detail of perianth trichomes (scale bar = 0.65 mm). Scale bars = 1 cm (A); 2.5 mm (B, C, D, E, O, P, Q, R); 1 mm (F, G, M, N); 2 mm (H, I, J, K); 0.5 mm (E, S); 0.65 mm (T). Drawn from fresh material from E.J. Hickman 2084 (PERTH 08988579). Illustrations by E.J. Hickman. E. J. Hickman & S .D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 113 Figure 11. Tribonanthes uniflora. A - group of plants each with a solitary-flowered inflorescence, {E.J. Hickman 2084); B - habitat {E.J. Hickman 2084), granite pavement with wind pruned shrubs Dodonaea ceratocarpa, Verticordia plumosa var. plumosa and Leucopogon parvflorus over herbs, sedges and grasses at Moses Rock, south of Yallinup, Western Australia; C - distribution (abbreviations for floristic provinces and districts as in Figure 9). Photos by E.J. Hickman. 114 Nuytsia Vol. 30 (2019) Illustrations. W.E. Blackall & BJ. Grieve, How to Know W Austral. Wildfl. Part I, p. 75 (1954), reprinted as Parts I, II, III p. 75 (1974); T.D. Macfarlane zwN.G. Marchant, J.R. Wheeler, B.L. Rye, E.M. Bennett, N.S. Eander & T.D. Macfarlane, FI. Perth Region p. 859, Figure 310 (1987); T.D. Macfarlane, FI. Australians-. 135,Figure52A-E(1987); J.R. Wheeler zwJ.WheelepN. Marchant&M. Eewington, Flora 5. IE Vol. 1: 313 (2002). Herb annually renewed from a white ovoid corm, 7.3^0 cm tall, consisting of a single branched aerial stem and single leaf Roots white, with visible white root trichomes, that sometimes have an associated loose sand-binding rhizosheath. Corms 12.5-15 mm diameter, covered by numerous old brown, papery sheaths, 2-10 cm below soil surface. Aerial stem (from basal leaf to inflorescence bract) 2-13.3 cm, lower 1/2 glabrous, the remainder with scattered trichomes. Basal leaf 4.5-23 cm long, erect, sheath stem-clasping, lamina flattened, green, with dark mucronate tip, glabrous. Inflorescence bract 2.5-\3 cm long, erect, stem-clasping below, 6-10 mm circumference at widest point, green with noticeable longitudinal veins and thin scarious margins; lamina 2-100 mm long, flattened, green with darker mucronate tip, glabrous. Peduncle (from inflorescence bract to lowest branching point) 3.3- 28 cm long, covered in white woolly trichomes, sparser on lower 1/2. Inflorescence a loose cyme of 2-5 pedicellate flowers, each subtended by a single floral bract. Pedicels 30-60 mm long, covered in white woolly trichomes. Floral bract 11-22 mm long, 2-5 mm wide, lanceolate, green with prominent longitudinal veins, outside surface hairy at base, inside surface glabrous, margins membranous, hairy, apex shortly terete, 0.5-5 mm long, ending in a darker tip, not exceeding perianth lobes. Perianth lobes 10-16 mm long, l-A mm wide, spreading to recurved, linear to narrowly obovate, inner and outer surfaces white woolly hairy, with purple tinge along midline of outer surface, apex with a dark mucronate tip. Perianth tube A-1 mm long, outer surface covered in silky white trichomes, with longer silky trichomes at base forming a distinct skirt, inner surface pale green, glabrous. Stamen connective appendages small, 2.0-3.0 mm long, 1.1-1.2 mm wide, yellow, topped with 5-7 short rounded tips, their apices below anther tips. Anthers 3^ mm long, yellow, with deep yellow sterile apex that projects upwards, attached centrally on stamen connective appendage. Ovary green, inferior, locules 3 with few (9) ovules per locule. Style short to sub-sessile, 0.7-2 mm long, green and purple longitudinal striations. Stigma capitate, green, with prominent tuft of white silky trichomes on top. Capsule greyish brown; perianth persistent; lobes spreading before dehiscence, reflexed after dehiscence. Seeds purplish grey-brown, angular. (Figures 12, 13) Diagnostic features. Tall plant, flattened leaves, long peduncles, multi-flowered inflorescence with long pedicels, woolly white flowers with long narrow perianth lobes, small yellow stamen connective appendages, short to sub-sessile style, stigma with prominent tuft of silky white trichomes on top, inferior ovary. Selected specimens examined. WESTERN AUSTRAEIA: 4.8 km NW of Mount Yetar, 5 Nov. 1996, MG. Allen 136 (PERTH); Ellis Brook Valley Reserve, 25 Aug. 1999, H. Bowler 411 (PERTH); Waroona,29Oct. 1991,R.J. Craz 2 /ze/'/io«a«r/?es'e/oog'c/to.A-single-floweredinflorescencewithwhitewoollyhairyspreadingperianthlobes,andyellow stamen connective appendages (E.J. Hickman 2073); B - habitat {E.J. Hickman 2073), burnt, seasonal wet flat, dominated by Melaleuca spp. shrubland, surrounded by Jarrah/Marri woodland, on Watershed Road, Mt Romance area of the state forest, north-west of Denmark, Western Australia; C - distribution (abbreviations for floristic provinces and districts as in Figure 9). Photos by E.J. Hickman. 128 Nuytsia Vol. 30 (2019) (PERTH); Moorialup Rd, E of Porongumps, 30 Aug. 1983, D. Davidson s.n. (PERTH); Jingaring Reserve, ESE ofBrookton, 1 Sep. 1998, R. Davis 6521 (PERTH); Hillman Forest Block, 27 Sep. 2006, S. Fisher BNC 1070 (PERTH); Beaufort River Bridge Nature Reserve, N of bridge, E of Albany Hwy, 18 Aug. 2014, EJ. Hickman 2012 (PERTH); Northam-Pithara Rd, 400 m S of Konnongorring West Rd, E side of road at base ofW sloping sheet of granite, 11 Sep. 2014, EJ. Hickman 2046 (PERTH); Nollajup Nature Reserve, Jayes Rd, 11.5 km E from Bridgetown-Boyup Brook Rd, N side of road around granite outcrop, 20 Sep. 2014, E.J. Hickman 2058 (PERTH); Mettabinup Nature Reserve, 3.3 km W on Flora and Fauna Rd from Tone Rd, then 500 m W into reserve to the SW boundary, 20 Sep. 2014, E.J. Hickman 2059 (PERTH); reserve on Frankland-Cranbrook Rd, 100 m N on Addis Rd from Frankland-Cranbrook Rd, 20 Sep. 2014, E.J. Hickman 2063 (PERTH); Wamballup Fake Nature Reserve, 500 m W on Wamballup Rd from Boyup Rd, S side of road in broad drainage line, 30 Sep. 2014, E.J. Hickman 2064 (PERTH); Mokine Nature Reserve, WSW of Northam, 2 Aug. 1985, G.J. Keighery 7794 (PERTH); Fake Muir Nature Reserve, NE side of Fake Muir, 27 Aug. 1998, G.J Keighery 15411 (PERTH); Poison Paddock, New Norcia, 5 Aug. 2004, K. Macey 594 (PERTH); 14 km N of Moora from Moore River Bridge, along road to Geraldton, 27 July 1982, T.D. Macfarlane 856 (PERTH); Mount Hardey Reserve, 19 Aug. 1998, J. Monks JM 108 (PERTH); Beverley Airfield Reserve, 27 Sep. 2000, M. Ochtman BAR 109 (PERTH); Dryandra State Forest, 30 Aug. 1988, D. Rose 669 (PERTH); Sullivan Rock S of Armadale, 30 Aug. 1992, R.J Rudall 20 (PERTH); 21 km from Collie towards Darkan, 22 Sep. 1983, J. Taylor & P. Ollerenshaw JT2101 (PERTH); Reserve 424A, Talbot West Road, York, 9 Aug. 2003, C. cS: A. Warburton AW 27 (PERTH); Highbury Block State Forest, 15 Oct. 1999, G. Warren, C. Taylor & P. Rose 315 (PERTH). Proposed vernacular name. Brilliant-fiowered Tiurndin. Phenology. Flowers from August to October. Distribution and habitat. Tribonanthes elongata is known from Watheroo in the north to the Porongurup Range in the south and from Perth east of the Darling Scarp to Pingelly. Most populations occur within the Jarrah, Narrogin and Muir Districts of the Bibbulmun Province and Wongan District of the Transitional Rainfall Province of south-west Western Australia, but one population is on the border of the Jarrah District of the Bibbulmun Province and the Fitzgerald-Stirling District of the Southeast Coastal Province (Gioia & Hopper 2017). Tribonanthes elongata grows on brown to grey sandy loam to sandy clay, on winter wet fiats and at the base of granite outcrops that support open woodlands, shrublands and herbfields. Associated species include Eucalyptus rudis, E. wandoo, E. loxophleba, E. marginata, E. occidentalis, Corymbia calophylla, Allocasuarina spp., Melaleuca preissiana, M. viminea, Hakea spp.. Acacia acuminata and various herbs and sedges (Figure 19). Conservation status. Widespread with no special conservation needs. Etymology. From the Eatin elongatus (elongated), referring to the long perianth lobes of this species. Notes. Based on determinations on herbarium specimens, Macfarlane (1987) included our Tribonanthes elongata in his concept of T. longipetala. However, T. elongata differs from T. longipetala in having terete leaves, a smaller stature and a single-fiowered inflorescence. Tribonanthes elongata also has a more widespread distribution, in winter wet areas throughout the SWAFR, while T. longipetala is generally found in seasonally wet areas on the slopes of the Darling Scarp. The other significant difference between these two species is that T. elongata has fine transparent trichomes on the inside of the filament below the anther, while T. longipetala has glabrous filaments. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 129 7. Tribonanthes keigheryi EJ.Hickman & Hopper, sp. nov. Type-. Wamballup Lake Nature Reserve, 500 m west along Wamballup Road from Boyup Road, south side of road in broad drainage line. Western Australia, 30 September 2014, EJ. Hickman 2065 {holo'. PERTH 08986940; iso-. CANB). Tribonanthes sp. Lake Muir (G.J. Keighery & N. Gibson 2134), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 1 Feb. 2019]. Illustration. Nil other than those herein. Herb annually renewed from a white ovoid corm, 3.0-26.5 cm tall above ground, consisting of a single stem and single leaf Corms 6.4-6.8 mm diameter, covered by numerous old brown, papery sheaths, 1.5^.6 cm below soil surface. Roots white, with no root trichomes, no sand-binding rhizosheath. Aerial stem (from basal leaf to inflorescence bract) 2-23 cm, glabrous. Basal leaf 4.4-17.5 cm long, erect, stem-clasping to Aliform, deep red-purple base grading to green, glabrous, with mucronate tip. Inflorescence bract 0.8-6.1 cm long, erect, funnel-like sheath with broad dilated mouth, 4-9 mm circumference at widest point, green with broad transparent membranous margins, sometimes tinged pink, fringed at top with fine white trichomes; lamina 5^4 mm long. Aliform, green, glabrous, with a dark mucronate tip. Peduncle (from inflorescence bract to floral bract) absent, or if present to 0.7 cm long, with very few scattered trichomes. Inflorescence a solitary flower, subtended by a single floral bract. Floral bract 4-11 mm long, 1.5-6 mm wide, orbicular to ovate, green with broad membranous margin tinged pink and fringed with white trichomes, otherwise glabrous, apex 0.5-3 mm long, terete, ending in a darker tip, not exceeding perianth lobes. Perianth lobes 3-6 mm long, 1.5-3 mm wide, spreading to erect, obovate, inner surface white woolly hairy, on outer surface sparsely hairy with broad central purple or green stripe. Perianth tube 2.5-5 mm long, outer surface green with few scattered white trichomes, inner surface white and glabrous. Stamen connective appendages small to residual 0-1.5 mm long, 0.7 mm wide, white with purple tinged spot where anther attaches, two pointed tips, their apices below anther tip. Anthers 0.7-2 mm long, yellow, with small paler yellow sterile apex that projects upwards, attached at top of the stamen connective appendage. Ovary green, half-inferior, locules 3 with several (14) ovules per locule. Style relatively long, 0.7-2.5 mm, green. Stigma capitate, yellow, with few short trichomes on top. Capsule greyish brown; perianth persistent; lobes erect before dehiscence, reflexed after dehiscence. Seeds purplish grey-brown, angular. (Figures 20, 21) Diagnostic features. Medium plant. Aliform leaf, short peduncles, solitary-flowered inflorescence, woolly white flowers with purple sparsely hairy outer surfaces, small to residual white stamen connective appendages, long style with simple stigma, half-inferior ovary. Selected specimens examined. WESTERN AUSTRALIA: Kangaroo Rd, 4.1 km E of Collis Rd, 14 Sep. 1994, A.R. Annels & P.W. Hearn ARA 4416 (PERTH); Wandoo Conservation Park, Goonapping Swamp, 17 Sep. 2014, K.L. Brown & G. Paczkowska KLB 1154 (PERTH); Brockman Highway, 6 Oct. 2000, R. Cranfield 15475 (PERTH); Coalfields Rd, 4 km NE of Bowelling, 25 Sep. 1995, V. Crowley DKN 70 (PERTH); King George Sound, 1 Dec. 1900, Col. Goadby B2063 (PERTH); Lake Muir Nature Reserve, 5.9 km W of Unicup Rd on Muirs Hwy, 60 m S of road side, 17 Oct. 2013, E.J. Hickman 2001 (PERTH); Kulunilup Nature Reserve, 2.3 km S ofWingebellup Rd on Unicup Rd, E side of road, 17 Oct. 2013, E.J. Hickman 2002 (PERTH); Cobertup Nature Reserve, 2.5 km N on Noobijup Rd from Muirs Hwy, 400 m W along N boundary firebreak, 17 Oct. 2013, E.J. Hickman 2004 (PERTH); Wimballup Swamp, NW side, 3.8 km W of Frankland River bridge, 2.9 km S on track to 130 Nuytsia Vol. 30 (2019) Figure 20. Tribonanthes keigheryi. A - whole plant; B - bud; C - flower from top; D - flower from side; E - flower with part of perianth removed to show ovary and style; F - stamens from front, back and side view showing stamen connective appendages scarcely developed, much shorter than anthers and yellow anthers without prominent apiculate apex; G - style elongated with simple stigma; H - capsule; I - cross section of capsule; J - longitudinal section of capsule; K - dehisced capsule; L - seeds; M - leaf section near apex (hollow); N - leaf section near base (hollow); O - inflorescence bract; P - floral bract; Q - longitudinal section of corm; R - root; S - cross section of root; T - detail of perianth trichomes. Scale bars = 1 cm (A); 2.5 mm (B, C, D, E, O, P, Q); 1 mm (F, G, E); 2 mm (H, I, J, K); 0.65 mm (M, N); 1.25 mm (R); 0.5 mm (S); 0.4 mm (T). Drawn from fresh material E.J. Hickman 2065 (PERTH 08986940). Illustrations by E.J. Hickman. E. J. Hickman & S .D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 131 Figure 21. Tribonanthes keigheryi. A - smdiW single-flowered inflorescence with white woolly hairy spreading perianth lobes, and minute stamen connective appendages {E.J. Hickman 2065); B - habitat {E.J. Hickman 2065), seasonally wet flat, in open Eucalyptus occidentalis and E. rudis woodland, Wamballup Lake Nature Reserve, west of Kendenup; C - distribution (abbreviations for floristic provinces and districts as in Figure 9). Photos by E.J. Hickman. 132 Nuytsia Vol. 30 (2019) Neerabup Rd, 5.5 km W to site, 23 Oct. 2013, EJ. Hickman 2005 (PERTH); Cannington, 28 July 1973 GJ. Keighery 366 (PERTH); 1 km S of Tambellup to Cranbrook, 20 Oct. 1983, GJ. Keighery 6715 (PERTH); Beaufort River Water Reserve, 7 Oct. 2003, GJ. Keighery 16357 (PERTH); Yoongarillup Townsite Reserve, SE ofBusselton, 15 Sep. 2006, G.J. Keighery 17045 (PERTH); South West Hwy, 6.5 km N of Palgarup, 12 Oct. 1983, T.D. Macfarlane 1244 (PERTH); Tooregullup Swamp, 15 km NE of Bremer Bay, 25 Sep. 1976, KR. Newbey 4987 (PERTH); Wandoo National Park, 4 Oct. 2007, M. Wheeler, P. Armstrong & students 480 (PERTH). Proposed vernacular name. Keighery’s Tiurndin. Phenology. Flowers from August to October. Distribution and habitat. The distribution of T. keigheryi is broadly scattered from Perth to north-east of Bremer Bay, but the majority of populations are located around the wetlands and swamps between Manjimup and Mt Barker, where the plants are often in standing water. Populations are concentrated in the Jarrah and Muir Districts of the Bibbulmun Province of the south-west of Western Australia (Gioia & Hopper 2017), but also occur in the Narrogin District of the Bibbulmun Province, and Fitzgerald- Stirling District of the Southeast Coastal Province. Tribonanthes keigheryi grows on grey to brown sand, sandy clay or clay, in the winter wet flats, valley floors and wetlands that support open woodlands, and sedgelands. Associated species include Eucalyptus rudis, E. wandoo, E. occidentalis, Corymbia calophylla, Melaleuca rhaphiophylla, M. cuticularis, M. viminea, M. lateritia, Hakea prostrata, H. varia Callistemon phoeniceus, Hypocalymma angustifolium, Chorizandra enodis, Leptocarpus spp., Orndujfia submersa and Pauridia spp. (Figure 21). Conservation status. Widespread with no special conservation needs. When first recognised, T. keigheryi was listed as Priority Three (Smith & Jones 2018). However, in light of the present study, this species has been shown to be widespread with no special need for conservation and it has consequently been de-listed. Etymology. Named for Gregory J. Keighery, survey botanist with the Department of Conservation and Eand Management and its successors, for his many contributions to the study of Western Australian flora and vegetation. Greg has an encyclopaedic knowledge of plant names and has become one of the most prolific collectors of Western Australian plants, as well as naming more than 100 new species. The specific epithet is pronounced ‘key-er-ee-eye’. Notes. Tribonanthes keigheryi differs from T. violacea in having a slenderer habit and smaller flowers. This difference was noted by Greg Keighery with specimens he first collected in 1997 and lodged at the Western Australian Herbarium with the designation Tribonanthes sp. Fake Muir. 8. Tribonanthes variabilis Find!., Sketch Veg. Swan R. xliv (1840). Type: Damp places. The Vasse, Western Australia, Mrs Molloy p.p. (lecto: CGE23342! all plants except second from the left, fide T.D. Macfarlane, El. Australia 45: 465 (1987); possible isolecto: K000356597 image!). Illustrations. WE. Blackall & B.J. Grieve, How to Know W. Austral. Wildfl. Part I, p. 75 (1954), reprinted as Parts I, II, III p. 75 (1974); T.D. Macfarlane El. Australia 45: 135 Figure 52 F-I (1987); E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 133 M. Simpson, Haemodoraceae, in K. Kubitzki, Fam. Gen. vascular plants IV, p. 213, Figure 54 A-B (1998); J.R. Wheeler in J. Wheeler, N. Marchant & M. Lewington, Flora S.W.V.m (2002). Herb annually renewed from a white ovoid corm, 13.6-39 cm tall above ground, consisting of a single stem and l(-2) leaves, the basal leaf and sometimes a cauline leaf Corms 7-10 mm diameter, covered by numerous old brown, papery sheaths, 2.7-6.0 cm below soil surface. Roots white, with rootlets but no visible root trichomes, no sand-binding rhizosheath. Aerial stem (from basal leaf to inflorescence bract) 3.2-18 cm, glabrous. Basal leaf 6.8-21.7 cm long, erect, stem-clasping to terete, red-purple below grading to deep green, with dark mucronate tip, glabrous, some plants with a cauline leaf on the aerial stem between the basal leaf and the inflorescence bract. Cauline leaf similar to basal leaf only shorter. Inflorescence bract2.1-\A.5 cm long, erect funnel-like sheath, 4-11 mm circumference at widest point; lamina 5-80 mm long, terete, green, with a narrow membranous margin and darker mucronate tip, glabrous. Peduncle (from inflorescence bract to floral bract) 6.8-28 cm long, lower 1/2 becoming sparsely hairy, the remainder covered in white woolly trichomes. Inflorescence a compact cyme of 2-7 shortly to moderately long-pedicellate flowers, each subtended by a single floral bract. Floral bract 7-18 mm long, 3.5-8.5 mm wide, lanceolate, green with one to several darker longitudinal striations, membranous margin fringed with white trichomes, outside surface hairy at base, inside surface glabrous, apical part 1-6 mm long, terete, ending in a darker tip, not exceeding perianth lobes. Perianth lobes 5-12 mm long, 2-6 mm wide, spreading, dilated toward the apex, inner and outer surfaces creamy white hairy, with a darker mucronate tip. Perianth tube 3.5-9.0 mm long, outer surface covered in long silky creamy white trichomes, inner surface green, glabrous. Stamen connective appendages 3-6 mm long, 1.5-1.7 mm wide, creamy white, topped with 4 or 5 flnger- like projections with pointed tips, exceeding anther tips. Anthers 2-A mm long, yellow, with cream to pink sterile apex that projects upwards, attached centrally on stamen connective appendages. Ovary green, half-inferior to inferior, locules 3 with many (26) ovules per locule. Style short to almost absent, 0.4-2 mm long, dark greenish purple. Stigma capitate, green, with elongated tip topped with three glandular protuberances and covered in white trichomes. Capsule greyish brown; perianth persistent; lobes spreading before dehiscence, reflexed after dehiscence. Seeds purplish grey-brown, angular. (Figures 22, 23) Diagnostic features. Tall plant, terete leaf, long peduncles, multi-flowered inflorescence, woolly creamy white flowers with perianth lobes dilated toward the apex, cream stamen connective appendages, short to almost absent style with stigma bearing an elongated tip topped with three glandular protuberances and covered in white trichomes, half-inferior to inferior ovary. Selected specimens examined. WESTERN AUSTRALIA: Brand Hwy, 2.2 km W of Eneabba, 26 Aug. 1985, M.E. Ballingall 1955 (PERTH); Lake Wannamal Nature Reserve, 1 km E of Bindoon-Moora Rd, 9 Sep. 2014, K.L. Brown & G. Paczkowska KLB 1150 (PERTH); Surprise Forest Block, 2.8 km W of Nornalup Rd on Mountain Rd, 22 Nov. 2004, R.J. Cranfield & B.G. WardWYM 298 (PERTH); Kooljerrenup Nature Reserve, 11 Sep. 2003, P. Foreman & J. Kelly KO 275 (PERTH); Leda Nature Reserve, 1.2 km S of Beekeepers Rd, 6 km W of Brand Hwy, 20 Aug. 2014, E.J. Hickman 2014 (PERTH); Marchagee Track, 2 km W of intersection with Dewars Rd, S side of road and S side of Dewars Creek, 20 Aug. 2014, E.J. Hickman 2017 (PERTH); Bashford Nature Reserve, SE comer of reserve, Bootoo Rd, E of Indian Ocean Dve, 28 Aug. 2014, E.J. Hickman 2026 (PERTH); Munbinea Rd, 10 km S of Bibby Rd, W side of road, 28 Aug. 2014, E.J. Hickman 2028 (PERTH); Cantabilling Rd, 3.5 km E of Munbinea Road, S side of road, 29 Aug. 2014, E.J. Hickman 2035 (PERTH); Ambergate Regional Park, Queen Elizabeth Ave., 150 mN of Doyle Rd, and 100 mW of road side, 19 Sep. 2014, E.J. Hickman 2053 (PERTH); Albany Hwy, 8.6 km S of Mt Cooke turn-off, 9 Nov. 1975, S.D. Hopper 842 (PERTH); Blackwood River bridge, 11 km E of Boyup Brook, 13 Nov. 1975, S.D. Hopper 882 134 Nuytsia Vol. 30 (2019) Figure 22. Tribonanthes variabilis. A - whole plant; B - bud; C - flower from top; D - flower from side; E - flower with part of perianth removed to show ovary and style; F - stamens from front, side and back view showing creamy white stamen connective appendages exceeding anthers and yellow anthers with apex apiculate and projecting upwards; G - style sub-sessile with distinct tuft of white silky trichomes on stigma; FI - capsule; I - cross section of capsule; J - longitudinal section of capsule; K - dehisced capsule; L - seeds; M - leaf section near apex; N - leaf section near base; O - inflorescence bract; P - floral bract; Q - longitudinal section of corm; R - root; S - cross section of root; T - detail of perianth trichomes. Scale bars = 1 cm (A); 2.5 mm (B, C, D, E, H, I, J, K, P); 1 mm (F, G, M, N); 0.5 mm (E, S); 5 mm (O); 4 mm (Q); 2 mm (R); 0.65 mm (T). Drawn from fresh material E.J. Hickman 2053 (PERTH 08986975). Illustrations by E.J. Hickman. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 135 Figure 23. Tribonanthes variabilis. A- multi-flowered inflorescence with white woolly hairy spreading perianth lobes, and corona of cream stamen connective appendages {E.J. Hickman 2035); B - habitat {E.J. Hickman 2053), seasonally wet flat, in open Corymbia calophylla woodland, with dominant species Kingia australis, Xanthorrhoeapreissii, Hypocalymma angustifolium, Adenanthos obovatus, Stirlingia latifolia, Synaphea sp., Nuytsia floribunda, Ambergate Regional Park, south of Busselton, Western Australia; C - distribution (abbreviations for floristic provinces and districts as in Figure 9). Photos by E.J. Hickman. 136 Nuytsia Vol. 30 (2019) (PERTH); Ring Rd, Bunbury, 19 Sep. 1983, GJ. Keighery 6436 (PERTH); Nullilla Nature Reserve, Gingin, 14 Sep. 2005 GJ. Keighery 16580 (PERTH); Harvey River District, Nov. 1901, Miss Lambert 814 (PERTH); Wongonderrah Rd, 9 km E of turn-off from Nambung Homestead, 30 Oct. 1999, C. MacPherson s.n. (PERTH); Manjimup, 28 Sep. 1948, R.D. Royce 2710 (PERTH); Wongan Hills, 9 Aug. 1949, E. Salisbury s.n. (PERTH); Augusta, 28 Aug. 2001, J. Scott A2\ (PERTH); Brixton Street Wetlands, Kenwick, E of Brixton St and N of Wanaping Rd, 17 Aug. 2009, K.R. Thiele 3766 (PERTH). Proposed vernacular name. Hairy-stigma Tiurndin. Phenology. Flowers from August to November. Distribution and habitat. Tribonanthes variabilis is found from near Geraldton along the west coast to Bunbury and inland to the Darling Scarp. Populations occur mainly within the Eesueur, Jarrah and Muir Districts of the Bibbulmun Province of the south-west of Western Australia (Gioia & Hopper 2017), but a few populations are located in the Wongan District of the Transitional Rainfall Province and the Nanda District of the Kalbarri Province. Tribonanthes variabilis grows on grey, brown, red-brown or black, sand, sandy clay, clay, sandy loam or peat in the winter wet flats, creek beds, along riversides and surroundings of granite outcrops, that support open woodlands, shrublands, heaths, sedgelands and herbflelds. Associated species include Corymbia calophylla. Eucalyptus rudis, E. loxophleba, Melaleuca rhaphiophylla, M. cuticularis, M. viminea, M. lateritia, M. uncinata. Acacia acuminata, A. lasiocalyx, Hakea ceratophylla, Viminariajuncea, Hypocalymma angustifolium mdXanthorrhoea preissii (Figure 23). Conservation status. Widespread with no special conservation needs. Etymology. Eatin for variable, originally referring to the variable nature of the stamen connective appendages within this species, of which two length conditions are mentioned in the original description of this species (Eindley 1839^0: 44), based on Georgiana Molloy collections of this species from the Vasse River (CBG 23342). As stated above, this specimen sheet actually contains two species; T. variabilis and T. longipetala, leading Eindley to describe two conditions for a single species. Notes. Macfarlane (1987) synonymised T. variabilis with T. australis. However, T. variabilis and T. australis specimens can be distinguished by their stigmas. Tribonanthes variabilis has a prominent appendage covered in silky white trichomes, present on the top of the stigma, while T. australis only has a few scattered trichomes on the top of the stigma. In addition, the perianth of T. variabilis is creamy white, while T. australis has a purple tinge, and the perianth lobes of T. variabilis are dilated at the ends making their width broader relative to their length than in T. australis. 9. Tribonanthes violacea Endl. in Eehm., PI. Preiss. 2: 28 (1846). Type'. Between mounts Elphinstone and Melville, Albany, Western Australia, 11 October 1840, J.A.L. Preiss 1562 {holo\ ED 1811441 image!; iso\ MEE 104289 image!, P 00753522 image!, E 1470274 image!). Illustration. T.D. Macfarlane, El. Australia 45: 135, Figure 52 J-M (1987). Herb annually renewed from a white ovoid corm, 4.5-12.5 cm tall above ground, consisting of a single stem and single leaf Corms 6.9-9.2 mm diameter, covered by numerous old brown, papery sheaths. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 137 1-5 cm below soil surface. Roots white, with a few white rootlets, no sand-binding rhizosheath. Aerial stem (from basal leaf to inflorescence bract) 2.5-8 cm, glabrous. Basal leaf 4.5-11 cm long, spreading, stem-clasping to terete, deep red-purple below grading to green with deep red margins, with mucronate tip, glabrous. Inflorescence bract 1.5-6.9 cm long, funnel-like sheath with broad dilated mouth, 5-11 mm circumference at widest point; lamina 5^3 mm long, terete, green with broad transparent membranous margin tinged pink, mucronate tip, glabrous. Peduncle (from inflorescence bract to floral bract) absent, or if present to 1.5 cm long, with a few scattered trichomes. Inflorescence a solitary flower, subtended by a single floral bract. Floral bract 5-19 mm long, 5-11 mm wide, orbicular to ovate, green with one to several darker longitudinal striations, broad membranous margin tinged pink and fringed with white trichomes, otherwise glabrous, apex 0.5-5 mm long, terete, ending in a darker tip, exceeding perianth lobes. Perianth lobes 4-9 mm long, 2-5 mm wide, spreading to erect, obtuse, white woolly hairy inner surface and sparsely hairy with broad central purple stripe on outer surface. Perianth tube 3-7.5 mm long, outer surface covered in white trichomes, inner surface white, glabrous. Stamen connective appendages small to residual, 1-2.5 mm long, 1.3-1.5 mm wide, white with purple tinged spot where anther attaches, with two pointed tips, shorter than anthers. Anthers 1.2- 3.1 mm long, yellow, with pale yellow sterile apex that projects upwards, attached at top of stamen connective appendage. Ovary green, half-inferior, locules 3 with many (55) ovules per locule. Style relatively long, 1-3.2 mm, green. Stigma capitate, simple, yellow, with few short trichomes on top. Capsule greyish brown; perianth persistent; lobes erect before dehiscence, spreading to reflexed after dehiscence. Seeds purplish grey-brown, angular. (Figures 24, 25) Diagnostic features. Medium plant, leaf lamina terete, short peduncles, solitary flowered inflorescence, woolly white flowers with more sparsely hairy purple outer surface, small to residual white stamen connective appendages, relatively long style with simple stigma, half-inferior ovary. Selected specimens examined. WESTERN AUSTRAEIA: Mingenew, 12 July 1970, A.M. Ashby 3275 (PERTH); Carnaby Close, Barrens Beach Estate, Hopetoun, 22 Aug. 2004, M. Bennett 1121 (PERTH); Sandalwood Rd 1.5 km from Cape Riche camping area, 1 Oct. 2013, G. Byrne 4813 (PERTH); Wilyung Hill, Albany, 30 Aug. 2004, R.J. Cranfield 20469 (PERTH); Gravel pit 14 km N of Mullewa on Gascoyne Junction Rd, 14 Sep. 2005, J. Docherty 380 (PERTH); Cape Arid National Park, 2.5 km NW from Seal Creek Campsite on firebreak track, 27 Oct. 2013, E.J. Hickman 2006 (PERTH); N boundary of reserve S of Esperance Airport, 2.2 km W on Crawford Rd from Esperance- Coolgardie Rd, 28 Oct. 2013, E.J. Hickman 2008 (PERTH); Eort River, E bank 100 m N of bridge on South Coast Hwy, 28 Oct. 2013, E.J. Hickman 2009 (PERTH); Merivale, intersection of Merivale Rd and Cape le Grand Rd, on two granite hills, 1 Aug. 2016, E.J. Hickman 2090 (PERTH); parking bay 1.8 km N of Gibson on Coolgardie-Esperance Hwy, 6 Sep. 1982, S.D. Hopper 2507 (PERTH); Moore River National Park, Beermullah, 26 July 2008, F. Hort & J. Hort FH 3203 (PERTH); 9 km SE ofYornup, 11 Sep. 1981, G.J Keighery 3995 (PERTH); 15.4 km S of Northampton along North West Coastal Hwy, 21 Aug. 1983, C.M. Lynch 24 (PERTH). Proposed vernacular name. Violet Tiurndin. Phenology. Flowers July to September. Distribution and habitat. Tribonanthes violacea is known from scattered populations from Northampton, north of Geraldton, along the coast to Cape Arid National Park east of Esperance. Populations occur in the Nanda District of the Kalbarri Province, Merredin District of the Transitional Rainfall Province, Eesueur, Jarrah and Muir Districts of the Bibbulmun Province, and Fitzgerald-Stirling, Maalak, Boylya and Esperance Districts of the Southeast Coastal Province of the south-west of Western 138 Nuytsia Vol. 30 (2019) Figure 24. Tribonanthes violacea. A - whole plant; B - bud; C - flower from top; D - flower from side; E - flower with part of perianth removed to show ovary and style; F - stamens from front, side and back view showing stamen connective appendages small, much shorter than anthers and yellow anthers with apex apiculate and projecting upwards; G - style elongated with simple stigma; Ft - capsule; I - cross section of capsule; J - longitudinal section of capsule; K - dehisced capsule; L - seeds; M - leaf section near apex; N - leaf section near base; O - inflorescence bract; P - floral bract; Q - longitudinal section of corm; R- root; S - cross section of root; T - detail of perianth trichomes. Scale bars = 1 cm (A); 2.5 mm (B, C, D, E, I, J, K, O, P, Q); 1 mm (F, G, M, N, T); 4 mm (H); 0.5 mm (L, S); 2 mm (R). Drawn from fresh material E.J. Hickman 2090 (PERTH 08989524). Illustrations by E.J. Hickman. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 139 Figure 25. Tribonanthes v/o/acea. A - single-flowered inflorescence with purple tinged, white woolly hairy spreading perianth lobes, on a short peduncle {E.J. Hickman 2090); B - habitat {E.J. Hickman 2090), mosaic of shrubland and moss swards on two granite hills at the intersection of Merivale Rd and Cape Le Grand Rd, Merivale, east of Esperance; C - distribution (abbreviations for floristic provinces and districts as in Figure 9). Photos by E.J. Hickman. 140 Nuytsia Vol. 30 (2019) Australia (Gioia & Hopper 2017). Tribonanthes violacea grows on grey sand over clay associated with sandstone along creek lines and in winter wet depressions in the north and on grey sand over clay around granite outcrops or on wet winter flats in the south. These habitats support shrublands, open scrub, and heaths. Associated species include Hakea spp., Melaleuca spp., Borya nitida, and various herbs, sedges and grasses (Figure 25). Conservation status. Widespread with no special conservation needs. Etymology. From the Latin for violet colour, referring to the purple tinge of the flowers of this species. A number of species have been misidentifled as T. violacea in the past including T. uniflora and the newly described T. keigheryi, T. monantha and T. porphyrea. Tribonanthes violacea is similar to T. keigheryi and T. porphyrea in having a short to indistinguishable peduncle, but differs from T. monantha and T. uniflora, which have a relatively long peduncle covered in white woolly trichomes. Tribonanthes violacea is similar to T keigheryi in having short to residual stamen connective appendages, which are often reduced to two small points on the tip of the filament, while T monantha, T porphyrea and T uniflora have prominent stamen connective appendages forming a distinct corona-like structure. Tribonanthes violacea is similar to T monantha, T keigheryi and T uniflora in having small flowers with the perianth lobes equal or shorter than the perianth tube but differs from T porphyrea which has large flowers with perianth lobes longer than the perianth tube. Tribonanthes violacea is similar to T monantha in having inflorescence bracts with a glabrous margin, by which it differs from T keigheryi, T porphyrea and T uniflora which have a fringe of white simple trichomes along their inflorescence bract margins. Tribonanthes violacea is probably most similar to T keigheryi but they are distinguished by T violacea being a more robust plant that dwells in damp environments associated with granite outcrops, while T keigheryi is a fine slender plant that prefers swampy habitats. 10. Tribonanthes brachypetala Lindl, Sketch Veg. SwanR. xliv (1840). Type'. Swan River, Western Australia, 1839, J. Drummond s.n. {holo: CGE 06823!; iso\ K 000846203 image!, K 000846202 image!). Tribonanthes odora End!., in Eehm., PI. Preiss. 2: 28 (1846), nom. illeg., nom. superfl. Type: Canning River, Western Australia, 1840, J.A.L. Preiss 2394 {holo\ ED 1811569 image!; iso\ MEE 104290 image!, MEE 104291 image!, MEE 104292 image!, S 06-11227 image!). Illustrations. WE. Blackall & B.J. Grieve, How to Know W Austral. Wildfl., Part I, p. 75 (1954), reprinted as Parts I, II, III p. 75 (1974); T.D. Macfarlane, FI. Australia 45: 135, Figure 52 N-Q (1987); J.R. Wheeler in J. Wheeler, N. Marchant & M. Eewington, Flora S.W. 1: 313 (2002). Herb annually renewed from a white ovoid corm, 21^1 cm tall above ground, consisting of a single stem and 2(-3) leaves, 1 basal and l(-2) cauline. Corms 10-11 mm diameter, covered by numerous old brown, papery sheaths, 3.2-5.8 cm below soil surface. Roots white, with no visible root trichomes, no sand-binding rhizosheath. Aerial stem (from basal leaf to inflorescence bract) 3.5-20 cm, mostly glabrous except upper 1/4 sparsely hairy. Basal leaf3).A-22 cm long, erect, stem-clasping to terete, deep green with dark mucronate tip, glabrous. Cauline leaf similar to basal leaf only shorter; some plants with l(-2) cauline leaves positioned 1/3-1/2 way up aerial stem (Figure 26A1). Inflorescence bract 1.8-8.2 cm long, sheath funnel-like, with broad dilated mouth 6-9 mm circumference at widest point, green with thin membranous margins, glabrous; lamina 14-55 mm long, terete, green with E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 141 darker mucronate tip, glabrous. Peduncle (from inflorescence bract to first floral bract) 13-26.5 cm long, covered in white woolly trichomes, with the trichomes becoming sparser in the lower 1/2. Inflorescence a nodding compact cyme of 3-7 sessile or shortly pedicellate flowers, each subtended by a single floral bract. Floral bracts 6-24 mm long, 2-10 mm wide, lanceolate, asymmetric, green with one to several darker longitudinal striations, hairy margins, hairy at base on outside surface, glabrous inside surface, apex 1-15 mm long, terete, green ending in a darker tip. Perianth lobes 3.5-5 mm long, 2-3 mm wide, strongly reflexed, ovate to triangular, green with white hairy outer surface, glabrous inner surface, dark red mucronate tips. Perianth tube length 5.2-8 mm, outer surface covered in white woolly trichomes, inner surface mostly glabrous except for scattered trichomes near the base. Stamen connective appendages 2.5-4 mm long, 2.2 mm wide and 1.8 mm deep, fleshy, deeply grooved on back, well exceeding anther tips, forming a conspicuous crown-like structure, yellow. Anthers 2-3 mm long, yellow, with cream sterile apex that projects upwards, attached on lower part of stamen connective appendages. Ovary green, half-inferior, locules 3 with few (6) ovules per locule. Style short, 0.8-2 mm long, green. Stigma capitate, greenish yellow, topped with three short glandular protuberances and covered with short trichomes. Capsule greyish brown; perianth persistent; lobes reflexed before and spreading to reflexed after dehiscence. Seeds purplish grey-brown, angular. (Figures 26, 27) Diagnostic features. Tall plant, terete leaf, long peduncles, multi-flowered nodding inflorescence, perianth lobes strongly reflexed, very obvious yellow stamen connective appendages, short style, stigma with three glandular protuberances, half-inferior ovary. Selected specimens examined. WESTERN AUSTRAEIA: Gooseberry Hill, 170 m E of Watsonia Rd and 210 m N of Gooseberry Hill Rd, 27 Aug. 2014, EJ Hickman 2022 (PERTH); Watershed Rd, 4.5 km N of intersection with Basin Rd, SE side of road on wet flat, 3 Nov. 2014, E.J. Hickman 2074 (PERTH); Tindale Rd, Kentdale, W of Denmark, 3 Aug. 1992, S.D. Hopper 7286 (PERTH); 18 km S of Bridgetown, near Yornup, 8 Oct. 1982, G.J. Keighery 5321 (PERTH); airport, SE Bunbury, 18 Sep. 1983, G.J. Keighery 6426 (PERTH); Ambergate Regional Park, S of Busselton, 13 Sep. 1994, G.J. Keighery 13398 (PERTH); Yoongarillup Townsite Reserve, SE of Busselton, 15 Sep. 2006, G.J. Keighery 17030 (PERTH); Serpentine townsite, 19 Oct. 1983, T.D. Macfarlane 1291 (PERTH); W of Harvey, 21 Sep. 1948, R.D. Royce 2654 (PERTH); Unicup Rd off Muir Hv^, 19 Sep. 1987, M Sherwood ^9 (PERTH). Proposed vernacular name. Nodding Tiurndin. Phenology. Flowers from July to October. Distribution and habitat. Tribonanthes brachypetala has been collected from north east of Perth on and west of the Darling Scarp, to Dunsborough, and south-east through the state forest to Mt Barker. Its most easterly population is at the base of the southern slopes of the Porongurup Range. Its distribution is concentrated along the western edge of the Bibbulmun Province of south-west Western Australia (Gioia & Hopper 2017), mainly in the Jarrah and Muir Districts. Its easterly population is on the border of the Muir District and Fitzgerald-Stirling District of the Southeast Coastal Province. It grows on seasonally wet brown loams around granite outcrops associated with open woodlands, and on black to grey peaty or sandy clays in winter wet flats associated with open woodlands. Associated species include Corymbia calophylla. Eucalyptus marginata, E. wandoo, Melaleuca rhaphiophylla md Xanthorrhoea preissii (Figure 27). Conservation status. Widespread with no special conservation needs. 142 Nuytsia Vol. 30 (2019) Figure 26. Tribonanthes brachypetala. A1 - whole plant; A2 - inflorescence; B - bud; C - flower from top; D - flower from side; E - flower with part of perianth removed to show ovary and style; F - stamens from front, side and back view showing yellowfleshy stamen connective appendages exceeding anthers and yellow anthers with apexapiculate and projecting upwards; G - style short with three protuberances on stigma; H - capsule; I - cross section of capsule; J - longitudinal section of capsule; K - dehisced capsule; L - seeds; M - leaf section near apex; N - leaf section near base; O - inflorescence bract; P - floral bract; Q - longitudinal section of corm; R - root; S - cross section of root; T - detail of perianth trichomes. Scale bars = 1 cm (Al); 5 mm (A2); 2.5 mm (B, C, D, E, H, I, J, K, P, R); 1 mm (F, G); 0.5 mm (E, S, T); 0.65 mm (M, N); 4 mm (O, Q). A- G & M- T drawn from fresh material at Millinup Road, Porongurup Range after a Are in 2010 (no voucher) and H- E drawn from EJ. Hickman 2022 (PERTH 08989370). Illustrations by E.J. Hickman. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 143 Figure 27. Tribonanthes brachypetaJa. A- plant showing multi-flowered inflorescence, with prominentyellow stamen connective appendages and reflexed perianth lobes (no voucher, Millinup Rd, Porongurup Range after a fire in 2010) B - habitat (no voucher, Millinup Rd, Porongurup Range after a fire in 2010), recently burnt open woodland of Corymbia calophylla and Eucalyptus marginata over Taxandria parviceps, Astartea sp. and Cyathochaeta equitans on grey brown loam over laterite at base of the southern slopes of the Porongurup Range, Western Australia; C - distribution (abbreviations for fioristic provinces and districts as in Figure 9). Photos by E.J. Hickman. 144 Nuytsia Vol. 30 (2019) Etymology. From the Greek brachys (short) and petalon (leaf, but in modern botany referring to petals). Notes. Tribonanthes brachypetala is more readily observed in the flowering period following a burn of its habitat and is then found in diminishing numbers of plants in subsequent seasons for a few years after the bum. The exception is when the plants’ habitat is open woodland of Eucalyptus wandoo, where the understorey is sparse, or on granite outcrops, when plants may flower in the absence of Are. The fruiting specimens examined show T. brachypetala displays early abortion of many ovules resulting in only a few seeds per capsule. Tribonanthes Endl. subg. Salina E. J.Hickman & Hopper, subg. nov. Type'. Tribonanthes minor M.Eyons & Keighery Aerial stem 1.3-4 cm tall. Flowers laterally horizontal, solitary. Perianth white to pale green with reflexed lobes. Style 0.5-1.5 mm long. Etymology. From the Eatin salina for salty, alluding to the general habitat occupied on the margins of salt lakes. Notes. This monotypic subgenus is distinctive in its diminutive size, the smallest species known of Haemodoraceae. 11. Tribonanthes minor M.Eyons & Keighery, Nuytsia 16: 78 (2006). Type'. Chinocup Nature Reserve [precise locality withheld for conservation reasons]. Western Australia, 18 October 2000, M.N Lyons 2734 Qiolo'. PERTH 07245890!). Illustration. M. Eyons & G. Keighery, Nuytsia 16: 79, Figure 1 (2006). Herb annually renewed from a white ovoid corm, 1.3-3.9 cm tall above ground, consisting of a single stem and single leaf Corms 4-6 mm diameter, covered by numerous old brown, papery sheaths, 1.3- 3.8 cm below soil surface. Roots white, with long fine root trichomes, some sand-binding rhizosheaths. Aerial stem (from basal leaf to inflorescence bract) 0.3-0.8 cm, glabrous. Basal leaf \.3-3 cm long, spreading, stem-clasping to terete, deep red-purple, with a mucronate tip, glabrous. Inflorescence bract 0.5-0.9 cm long, funnel-like sheath with broad dilated mouth, red-purple, 2-4 mm circumference at widest point, with glabrous margin, lamina 0.2-1.5 mm long, terete, red-purple to green, with mucronate tip. Peduncle (from inflorescence bract to floral bract) 4.5-22 mm long, covered in white woolly trichomes. Inflorescence a solitary flower, subtended by a single floral bract. Floral bract 4-5.5 mm long, 3-5 mm wide, ovate, pinkish-purple, broad membranous margins, with a few fringing trichomes, otherwise glabrous inside and out, apex 0.5-1 mm long terete, ending in a darker tip, not exceeding perianth lobes. Perianth lobes 1.1-2 mm long, 0.8-1.2 mm wide, strongly reflexed, acute, outside surface white woolly hairy, inside surface pale green to purplish, glabrous. Perianth tube 2.6-3.5 mm long, outer surface covered in white woolly trichomes, inner surface pale green, glabrous. Stamen connective appendages relatively large, 0.7-1 mm long, 0.8-1 mm wide, fleshy, yellow, topped with 4 broad, round-tipped flnger-like projections, extending above anther tips. Anthers 1-1.3 mm long, cream, with cream sterile apex that projects away from centre of flower, attached at base of stamen connective appendage. Ovary green, superior, 3 locules with several (10-12) ovules per locule. Style short to almost absent, 0.5-1.5 mm long, green. Stigma capitate, simple, yellow, covered with short E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 145 trichomes. Capsule greyish brown; perianth persistent; lobes reflexed before and after dehiscence. Seeds purplish grey-brown, angular. (Figures 28, 29) Diagnostic features. Small plant, terete leaf, long peduncles, solitary flowered inflorescence, strongly reflexed perianth lobes, obvious yellow stamen connective appendages, short to almost absent style with simple stigma, superior ovary. Selected specimens examined. WESTERN AUSTRAEIA [localities withheld for conservation reasons]: 4 Oct. 2007, T Erickson 116 (PERTH); 28 July 2015, E.J Hickman 2079 (PERTH); 28 July 2015, E.J. Hickman 2081 (PERTH); 12 Oct. 1988, S.D. Hopper 6931 (PERTH); 19 Oct. 1999, M.N. Lyons 2929 (PERTH); 24 Oct. 2000, M.N. Lyons 2735 (PERTH); 28 Aug. 2007, R.J. Smith, S.D. Hopper & L. Sweedman 059 (K, PERTH). Proposed vernacular name. Salt-lake Tiurndin. Phenology. Flowers July to August. Distribution and habitat. Tribonanthes minor is known from scattered populations from the Meckering area south to the Take King area. Populations are located in the Wongan District of the Transitional Rainfall Province and Maalak District of the Southeast Coastal Province of south-west Western Australia (Gioia & Hopper 2017). It grows on seasonally wet, low sandy rises on the the margins of naturally saline lakes. The vegetation is heath, scrub and samphire flats, associated species include, Melaleuca thy aides, Atriplex hymenotheca, Tecticornia indica, T. halocnemoides, T. leptoclada, Poycea spinescens, Samolus caespitosus, Tecticornia moniliformis, Gnephosis acicularis, G. tridens, Hyalochlamys globifera and a variety of annual herbs (Figure 29). Conservation status. Currently listed as Priority Three under Conservation Codes for Western Australian Flora (Smith & Jones 2018). Due to its preference for low elevations on the margins of naturally saline lakes T. minor is under threat from increased risk of flooding associated with changes in catchment hydrology following agricultural clearing (Eyons & Keighery 2006; Smith & Jones 2018). Etymology. From the Eatin minor (smaller), referring to the very small leaves, stems, flowers and perianth lobes of this species. Notes. Tribonanthes minor is presently known from three or four locations, with herbarium specimens collected from all but the Eake Goorly location. Its presence at Eake Goorly was recorded in quadrats for the Salinity Action Plan survey (Keighery 2004) across the state of Western Australia in 1999 (M. Eyons pers. comm., 13 September 2013), but is thought to be dubious. Field visits to Eake Goorly by N. Gibson in October 2013 (N. Gibson pers. comm. 22 October 2013), EJH in July 2015 and SDH in September 2017 failed to locate any specimens. Eake Goorly is not shown on the map (Figure 29C). Tribonanthes Endl. subg. Boya E.J.Hickman & Hopper, subg. nov. Type '. Tribonanthes purpurea T.Macfarlane & Hopper. Aerial stem 1.1^.3 cm tall. Flowers erect, solitary. Perianth pink with erect lobes. Style 4.0-7.0 mm long. 146 Nuytsia Vol. 30 (2019) Figure 28. Tribonanthes minor. A - whole plant; B - bud; C - flower from top; D - flower from side; E - flower with part of perianth removed to show ovary and style; F - stamens from front, back and side view showing yellow fleshy, stamen connective appendages exceeding anthers and cream anthers with apexapiculate and projecting upwards; G- style sub-sessile with simple stigma; FI - capsule; I - cross section of capsule; J - longitudinal section of capsule; K - dehisced capsule; L - seeds; M - leaf section near apex; N - leaf section near base; O - inflorescence bract; P - floral bract; Q - longitudinal section of corm; R - root; S - cross section of root; T - detail of perianth trichomes. Scale bars = 2.5 mm (A); 1 mm (B, C, D, E, H, I, J, K, O, P); 0.5 mm (F, G, M, N, R, T); 0.4 mm (E); 2 mm (Q); 0.2 mm (S). Drawn from fresh material E.J. Hickman 2079 (PERTH 08989427). Illustrations by E.J. Hickman. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 147 Figure 29. Tribonanthes minor. A - group of plants each with a single-flowered inflorescence with obvious yellow stamen connective appendages and strongly reflexed perianth lobes {E.J. Hickman 2079); B - habitat of T. minor {E.J. Hickman 2081), samphire on low sandy rises; C - distribution (abbreviations for floristic provinces and districts as in Figure 9). Photos by E.J. Hickman. 148 Nuytsia Vol. 30 (2019) Etymology. From the Noongar word boya for granite rock or stone, as a noun in apposition, alluding to the habitat occupied by the sole species. Notes. A monotypic subgenus distinctive in its almost stemless pink erect tubular flowers. 12. Tribonanthes purpurea T.Macfarlane & Hopper, FI. Australia 45: 465 (1987). Type -, near Kuender [precise locality withheld for conservation reasons]. Western Australia, 27 August 1986, S.D. Hopper 5237 {holo'. PERTH 01008625!; iso'. CANB. 00394484 image!). Illustrations. T.D. Macfarlane, FI. Australia 45: 135, Figure 52 R-U (1987); S.D. Hopper, S. van Leeuwen, A.R Brown & S.J. Patrick, WA Endemic Flora p. 79 (1990); A. Brown, C. Thomson-Dans &N. Marchant, WA Threatened Flora 167 (1998). Herb annually renewed from a white ovoid corm, 1.1-4.3 cm tall, consisting of a single stem and single leaf Corms 5-6 mm diameter, covered by numerous old brown, papery sheaths, 1 -4.6 cm below surface. Roots white, with long fine root trichomes, lacking a sand-binding rhizosheath. Aerial stem (from basal leaf to inflorescence bract) 0.1-2.2 cm, glabrous. Basal leaf 2-5. A cm long, spreading, stem- clasping base, becoming terete, the base white grading to green, with darker mucronate tip, glabrous. Inflorescence bract 9-20 mm long, erect, funnel-like sheath with broad dilated mouth, 6-12 mm circumference at widest point, purplish green with broad pinkish purple membranous margins, glabrous; lamina 2-10 mm long, terete, green with a mucronate tip. Peduncle (from inflorescence bract to floral bract) absent, or if present to 0.2 cm long, glabrous. Inflorescence a solitary flower, often appearing geoflorous, subtended by the inflorescence bract and a floral bract. Floral bract 6-13 mm long,4- 10 mm wide, ovate, green with broad pinkish purple membranous margins, inside and outside surfaces glabrous, apex 1^ mm long, terete, ending in a darker tip, not exceeding perianth lobes. Perianth lobes 4-6.1 mm long, 1.6-3 mm wide, erect, oblong, pink, glabrous inside and outside, outer lobes with reddish-purple gland-like protuberances at apex. Perianth tube A.6-1 mm long, white, glabrous inside and outside. Stamen connective appendages scarcely developed, 2-3 white projections, much shorter than anther tips. Anthers 1-2 mm long, yellow, with cream sterile apex that projects upwards, attached towards top of stamen connective appendage. Ovary green, superior, 3 locules with few (7) ovules per locule. Style long, 4.0-7.0 mm, green. Stigma capitate, simple, yellow, with a few short trichomes on top. Capsule greyish brown; perianth persistent; lobes erect before dehiscence, upwardly spreading after dehiscence. Seeds purplish grey-brown, angular. (Figures 30, 31) Diagnostic features. Small plant, terete leaf, peduncles absent or rudimentary, solitary flowered inflorescence, pink glabrous flowers often appearing geoflorus, stamen connective appendages scarcely developed, superior ovary, long style with simple stigma. Selected specimens examined. WESTERN AUSTRALIA [localities withheld for conservation reasons]: 29 Aug. 1994, A.M. Coates 4312 (PERTH); 31 July 1999, V. Crowley s.n. (PERTH); 12 Sep. 1995, R. Cugley 1 (PERTH); 23 Aug. 2004, P. Cugley 159 (PERTH); 22 Aug. 1990, D. Davidson s.n. (PERTH); 28 July 2015, E.J. Hickman 2080 (PERTH); 29 July 2015, E.J. Hickman 2082 (PERTH); 1 Aug. 2015, E.J. Hickman 2083 (PERTH); 1 Aug. 2004, G.J. & B.J. Keighery 270 (PERTH); 2 Sep. 2011, D. Sandow et al. DSA07 (PERTH). Proposed vernacular name. Granite Pink (proposed by Hopper et al. 1990). E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 149 Phenology. Flowers from July to September. Distribution and habitat. Tribonanthespurpurea is known from the Mt Dale area in the Darling Range to the Porongurup Range and east to Varley. Populations are located within the Jarrah and Narrogin Districts of the Bibbulmun Province, the Hyden District of the Transitional Rainfall Province and the Maalak and Fitzgerald-Stirling Districts of the Southeast Coastal Province of south-west Western Australia (Gioia & Hopper 2017). Tribonanthes purpurea mostly grows on brown sandy loam associated with granite outcrops but has also been found growing on grey sandy clays of winter wet flats. These habitats support moss swards, herbflelds, heath and shrublands with associated species including Thryptomene australis, Borya constricta, Dodonaea viscosa, Melaleuca elliptica, M. viminea, M. uncinata. Acacia lasiocalyx, Spartochloa scirpoidea, Stypandra glauca, Drosera spp., Stylidium spp., Caladenia spp. and Wurmbea spp. (Figure 31). Conservation status. Threatened Flora protected under the Biodiversity Conservation Act 2016 with a ranking of Vulnerable. Due to its small stature, succulent leaves and occurrence on open herbflelds, it is under threat from grazing by rabbits (Brown et al. 1998; Smith & Jones 2018). Its ability to regenerate after grazing is unknown. Etymology. Specific name from the Latin purpureus meaning purple, or red with a touch of blue, referring to the pink perianth lobes of this species. Notes. Tribonanthes purpurea is the most distinctive species of the genus with its glabrous pink flowers. Presently it is known from 14 populations, several of which have sub-populations within the same granite outcrop complex. However, the senior author has revisited several of these populations over the last five years and not relocated the plants at three locations - Beaufort River, Porongurup Range and north of the Stirling Range. Neither author has visited the Hillman location. Natural hybrids A small number of natural hybrid combinations of Tribonanthes have been recorded in herbarium collections at PERTH. Apart from morphological intermediacy and comparative rarity in populations of abundant likely parents, evidence in favour of the occurrence of natural hybridization is cited on the label of one specimen where pollen fertility was reduced to 62% in the putative hybrid of T australis X ? violacea (S.D. Hopper 5776). Tribonanthes australis Endl. x brachypetala Eindl. Specimens examined. Brixton Road, Beckenham, 13Aug. 1983, G.J. Keighery 6249 (PERTH); airport, 6 km SE Bunbury to Boyanup, 19 Sep. 1983, G.J. Keighery 6427 (PERTH); 9 km along Railway Rd, Boyanup to Capel, 29 Aug. 1984, G.J. Keighery 6828 (PERTH); Ambergate Regional Park, S of Busselton, 13 Sep. 1994, G.J. Keighery 13399 (PERTH); Yoongarillup Townsite Reserve, SE of Busselton, 15 Sep. 2006, G.J. Keighery 17029 (PERTH). Tribonanthes australis Endl. x longipetala Eindl. Specimens examined. WESTERN AUSTRAEIA: Brixton Rd, Beckenham, Perth, 21 Aug. 1983, G.J. Keighery 6260 (PERTH); Pinjarra Nature Reserve, Pinjarra, 30 Aug. 2007, G.J. Keighery 17133 (PERTH). 150 Nuytsia Vol. 30 (2019) Figure 30. Tribonanthes purpurea. A - whole plant; B - bud; C - flower from top; D - flower from side; E - flower with part of perianth removed to show ovary and elongated style; F - stamens from front, side and back view showing stamen connective appendages scarcely developed, translucent, surface punctate, much shorter than anthers and yellow anthers with apex apiculate and projecting upwards; G - style tip with capitate stigma; Ft - capsule, glabrous; I - cross section of capsule; J - longitudinal section of capsule; K - dehisced capsule; L - seeds; M - leaf section near apex; N - leaf section near base; O - inflorescence bract; P - floral bract; Q - longitudinal section of corm; R - root; S - cross section of root; T - detail of perianth tip. Scale bars = 4 mm (A); 1.5 mm (B, C, D, E, H, I, J, K); 0.5 mm (F, G, E, M, N); 2 mm (O, P, Q); 1 mm (R); 0.4 mm (S, T). Drawn from fresh material E.J. Hickman 2083 (PERTH 08989443). Illustrations by E. J. Hickman. E. J. Hickman & S .D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 151 Figure 31. Tribonanthes purpurea. A - single-flowered inflorescence with pink, glabrous, erect perianth lobes, {E.J. Hickman 2080); B - habitat of T. purpurea {E.J. Hickman 2080), Borya herbfleld and moss swards in shallow soil pockets between granite sheets; C - distribution (abbreviations for floristic provinces and districts as in Figure 9). Photos by E.J. Hickman. 152 Nuytsia Vol. 30 (2019) Tribonanthes australis Endl. x ?violacea Endl. Specimen examined. WESTERN AETSTRAEIA: 12.6 km off Unicup Rd along Wingebellup Rd, ESE to Unicup Eake, 16 Oct. 1986, S.D. Hopper 5776 (PERTH). Tribonanthes variabilis Eindl. x brachypetala Eindl. Specimen examined. WESTERN AUSTRAEIA: Brixton Street Wetlands, Kenwick, 24 Aug. 2010, K.L. Brown & G. Paczkowska KEB 854 (PERTH). Acknowledgements Thanks to Charmaine Hickman, Kate Brown, Carol Wilkins, Harriet Paterson, Anne Coates, Rosemary Cugley, Gillian & Bay Collison, Sarah Barrett, Ross Burnett and Philip Kinsey for assistance in the field and accommodation during fieldwork. Collections were made under scientific licenses issued by the Department of Parks and Wildlife and its predecessors. We are grateful to the Directors of PERTH, K, MEE, NSW, CANB, AD, CBG and BM for approval to work on specimens under their care. Travel by EJH to examine collections at K and the BM was supported by a Churchill Fellowship and by grants awarded by Kings Park and Botanic Garden and the Royal Botanic Gardens Kew while SDH worked as Director/CEO at these organisations. Mike Eyons and Neil Gibson provided information on Tribonanthes minor populations. Terry Macfarlane and Greg Keighery helpfully discussed species concepts with EJH. Matthew Barrett and Benjamin Anderson provided able training and assistance with molecular analyses. Aspects of seed biology were profitably discussed with Anne Cochrane, Andrew Crawford and David Coates. Karina Knight of the Western Australian Herbarium facilitated access to the specimen collection and processed incoming specimens from our research. Christine Bartram from Cambridge University Herbarium was most helpful with access to the specimen collection and the supply of high-resolution images of type specimens. Pina Milne from the Royal Botanic Gardens, Victoria supplied a high-resolution image of a type specimen of Tribonanthes violacea. Ernst Vitek from the Naturhistorisches Museum Wien provided information on the type specimen of Tribonanthes australis. Alan Paton and Elizabeth Howard enabled access to the Herbarium, Royal Botanic Gardens Kew. Ellen J. Hickman undertook this research while studying for her Doctor of Philosophy degree at the University of Western Australia, supported by an Australian Postgraduate Award with a University of Western Australia Top-Up Award. Stephen D. Hopper was supported by an Australian Research Council Discovery Outstanding Researcher Award as part of a Discovery Project (DP140103357), as well as by grants from the Great Southern Development Commission and the Jack Family Trust. References Aerne-Hains, L. & Simpson, M.G. (2017). Vegetative anatomy of the Haemodoraceae and its phylogenetic significance. International Journal of Plant Science 178(2): 117-156. Barrett, R.L., Hopper, S.D., Macfarlane, T.D. & Barrett, M.D. (2015). Seven new species of Haemodorum (Haemodoraceae) from the Kimberley region of Western Australia. Nuytsia 26; 111-125. Bentham, G. (1873). Flora Australiensis: A description ofplants of the Australian territory. Vol. 6. pp. 426-428. (L. Reeve & Co.: London, UK). Brown, A., Thomson-Dans, C. & Marchant, N. (1998). Western Australia’s Threatened Flora. (Department of Conservation and Land Management; Como, Western Australia). Darwin, C.R. (1868). On the specific difference between Primula veris, Brit. FI. (var. officinalis, of Linn.), P. vulgaris, Brit. FI. (var. acaulis, Linn.) and P elatior, Jacq.; and on the hybrid nature of the common Oxlip. With supplementary remarks on naturally-produced Hybrids in the genus Verhascum. Journal of the Linnean Society of London (Botany) 10: 437-454. E. J. Hickman & S.D. Hopper, A revision of the tiumdins (Tribonanthes, Haemodoraceae) 153 Daw, B., Walley, T. & Keighery, G. (1997). Bush Tucker Plants of the South-West. (Department of Conservation and Land Management: Kensington, Western Australia). Drummond, J. (1842a). On the botany ofWestem Australia. Inquirer (Perth, Western Australia) Wednesday 10 August 1842, p. 4-5. Drummond, J. (1842b). On the botany of Western Australia. Inquirer (Perth, Western Australia) Wednesday 17 August 1842, p. 5. Endlicher, S. (1837-41). Iconographia generumplantarum. (F. Beck: Vindobonae). Endlicher, S. & Fenzl, E. (1839). Novarum stirpium decas I-X. (Typis Sollingerianis: Vindobonae). Endlicher, S. (1846). Haemodoraceae. In: Lehmann, J.G.C. (Qd.) Plantae Pressianae, Vol. 2. pp. 14-29. (Sumptibus Meissneri: Hamburg). Fryer, R. (2006). Appendix 1: seed germination records. In: Sweedman, L. & Merrit, D. {Qds) Australian seeds: a guide to their collection identification and biology, pp. 199-219. (CSIRO Publishing: Melbourne; Botanic Gardens and Parks Authority: Perth & Millenium Seed Bank: Royal Botanic Gardens, Kew, London). Geerinck, D. (1969). Genera des Haemodoraceae et des Hypoxidaceae. Bulletin du Jardin Botanique National de Belgique 39: 47-82. Gioia, P. & Hopper, S.D. (2017). A new phytogeographic map for the Southwest Australian Floristic Region after an exceptional decade of collection and discovery. BotanicalJournal of the Linnean Society 184: 1-15. Grey, G. (1840). A vocabulary of the dialects of south-western Australia. (T. & W. Boone: London, UK). Hickman, E. J., Yates, C. J. & Hopper, S.D. (2017). Botanical illustration and photography - a Southern Hemisphere perspective. Australian Systematic Botany 30(4): 291-325. Hopper, S.D. (2003). South-western Australia, Cinderella of the world’s temperate floristic regions 1. Curtis’s Botanical Magazine 20(2)'. 101-126. Hopper, S.D. (2004). South-western Australia, Cinderella of the world’s temperate floristic regions 2. Curtis’s Botanical Magazine 2\(2): 132-180. Hopper, S.D. (2009). OCBIL theory: towards an integrated understanding of the evolution, ecology and conservation of biodiversity on old, climatically-buffered, infertile landscapes. Plant and Soil 322: 49-86. Hopper, S.D. & Brown, A.P (2001). Contributions to Western Australian orchidology: 1. History of early collections, taxonomic concepts and key to genera. Nuytsia 14(1/2): 1-26. Hopper, S.D., Silveira, F.A.O. & Fiedler, RE. (2016). Biodiversity hotspots and Ocbil theory. Marschner Review. Plant Soil 403: 167-216. Hopper, S.D., Fay, M.F., Rossetto, M. & Chase, M.W. (1999). A molecular phylogenetic analysis of the bloodroot and kangaroo paw family, Haemodoraceae: taxonomic, biogeographic and conservation implications. BotanicalJournal of the Linnean Society 131: 285-299. Hopper, S.D., van Leeuwen, S., Brown, A.P. & Patrick, S.J. (1990). Western Australia’s Endangered Flora and other plants under consideration for declaration. (Department of Conservation and Land Management: Perth.) Hopper, S.D., Chase, M.W. & Fay, M.F. (2006). A molecular phylogenetic study of generic and subgeneric relationships in the south-west Australian endemics Conostylis and Blancoa (Haemodoraceae). Aliso: A Journal ofSystematic and Evolutionary Botany 22{\y. 527-538. Hopper, S.D., Smith, R.J., Fay, M.F., Manning, J.C. & Chase, M.W. (2009). Molecular phylogenetics of Haemodoraceae in the Greater Cape and Southwest Australian Floristic Regions. Molecidar Phylogenetics and Evolution 51: 19-30. Keighery, G.J. (2004). State Salinity Strategy biological survey of the Western Australian wheatbelt: background. Records of the Western Australian Museum Supplement 61’. 1-6. Lamont, B.B. & Downes, K.S. (2011). Fire-stimulated flowering among resprouters and geophytes in Australia and South Africa. Plant Ecology 2\2{\2y. 2111-2125. Findley, J. (1839-40). A Sketch of the vegetation of the Swan River Colony. (James Ridley: Piccadilly, London). Lyons, M.N. & Keighery, G.J. (2006). A new species of Tribonanthes (Haemodoraceae) from saline wetland margins in Western Australia. Nuytsia 16(1): 77-80. Macfarlane,T.D. (1987). Tribonanthes. In: George,A.S. {Qd.) Flora of Australia. Vol. 45. pp. 131-134. (Australian Government Publishing Service: Canberra). Macfarlane,T.D. & Hopper, S.D. (1987). Haemodoraceae. Appendix: Newtaxa, combinations and lectotypiflcations, 7;^: George, A.S. {Qd.) Flora of Australia. Vol. 45. pp. 454-455. (Australian Government Publishing Service: Canberra). Moore, G.F. {\'&M). Diary of Ten Years Eventful Life of an Early Settler in Western Australia and also A Descriptive Vocabulary of the Language of the Aborigines. Facsimile edition (1978). (University of Western Australia Press: Nedlands, Western Australia). Mueller, F. (1872-1874). Fragmenta Phytographiae Australiae, Vol. 8. pp. 23. (Government Printer: Melbourne). 154 Nuytsia Vol. 30 (2019) Parsons, R.F. & Hopper, S.D. (2003). Monocotyledonous geophytes: comparison of south-western Australia with other areas of Mediterranean climate. Australian Journal of Botany 51(2): 129-133. Pate, J.S. & Dixon, K.W. (1982). Tuberous, cormous and bulbous plants. (University of Western Australia Press: Nedlands, Western Australia). Pierce, N. B., & Simpson, M. G. (2009). Polyaperturate pollen types and ratios of heteromorphism in the monocot genus Conostylis (Haemodoraceae). Australian Systematic Botany, 22(1): 16-30. Rooney, B. (2002). The Legacy of the Late Edward Mippy: An Ethnographic Biography. PhD Thesis. (Curtin University of Technology: Perth). Rooney,B. (2011). The Nyoongar Legacy: The naming ofland and the language of its people. (Batchelor Press: Batchelor, NT). Sandiford, E.M. & Barrett, S. (2010). Albany Regional Vegetation Survey Extent, Type and Status, A project funded by the Western Australian Planning Commission (EnviroPlanning ‘Integrating NRM into land Use Planning’ and State NRM Program), South Coast Natural Resource Management Inc. and City of Albany for the Department of Environment and Conservation. Unpublished report. Department of Environment and Conservation, Western Australia. Sharr, F.A. (1996). Western Australian Plant Names and their Meanings: A Glossary. (University of Western Australia Press: Nedlands, Western Australia). Simpson, M.G. (1983). Pollen ultrastructure of the Haemodoraceae and its taxonomic significance. Grana 22: 79-103. Simpson, M.G. (1990). Phylogeny and classification of the Haemodoraceae. Annuals Missouri Botanic Garden 77(4): 722-784. Simpson, M.G. (1993). Septal nectary anatomy and phylogeny in the Haemodoraceae. Systematic Botany 18: 593-613. Simpson, M.G. (1998). Haemodoraceae. In: Kubitzki, K. (ed). The families and genera of vascular plants. Flowering plants. Monocotyledons: Alismatanae and Commelinanae (except Gramineae). Vol. 4. pp. 212-222. (Springer: Berlin). Simpson, M.G. (2006). Plant systematics. (Elsevier Academic Press, Amsterdam). Smith, M.G. & Jones, A. (2018). Threatenedand Priority Floralist 5 December 201 8 . Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 5 February 2019]. Smith, R.J., Hopper, S.D. & Shane, M.W. (2011). Sand-binding roots in Haemodoraceae: global survey and morphology in a phylogenetic context. Plant and Soil 348: 453-470. von Brandenstein, C.G. (1988). NyungarAnewphonology, text samples and etymological and historical 1500-word vocabulary of an artificially re-created aboriginal language in the south-west of Australia. Pacific Linguistic Series C - No. 99. (Department of Linguistics, Research School of Pacific Studies, The Australian National University: Canberra, Australia). Western Australian Herbarium (1998- continuously updated). FloraBase—the Western Australian Flora. Department of Biodiversity, Conservation and Attractions, https://fiorabase.dpaw.wa.gov.au/ [accessed 10 January 2018]. Wheatbelt Natural Resource Management (n.d.). Nyungar Budjara Wangany Nyungar NRM wordlist and language collection booklet of the Avon Catchment Region. (Wheatbelt NRM, Northam). Nuytsia The journal of the Western Australian Herbarium 30: 155-175 Published online 30 July 2019 A taxonomic review of the Styphelia xerophylla group (Ericaceae: Epacridoideae: Styphelieae) Michael Hislop^ and Caroline Pnente-Lelievre^ ‘Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 ^United States Food and Drug Administration, 5001 College Park, Maryland 20740, USA Abstract Hislop, M. & Puente-Lelievre, C. A taxonomic review of the Styphelia xerophylla group (Ericaceae: Epacridoideae: Styphelieae). AwyAzh! 30:155-175(2019). Species belonging to the Western Australian S. xerophylla (DC.) F.Muell. group (or Group IX) are described and illustrated. The treatment includes two species published in the nineteenth century, S. xerophylla and S. stomarrhena (Sond.) Sleumer, together with four new species, S. angustiflora Hislop & Puente-Eef, S. cernua Hislop & Puente- Eef, S. disjuncta Hislop & Puente-Eel. and S. sulcata Hislop & Puente-Eel. A seventh member of the group, S. longissima Hislop & Puente-Eel., was described in a recent paper and is not treated again here. The morphological attributes of the group are outlined and a key to species provided. Introduction The Styphelia xerophylla (DC.) F.Muell. group (or Group IX) is one of 12 phylogenetic groups resolved within the large Styphelia ^m.-Astroloma R.Br. clade of tribe Styphelieae Barth, as a result of recent molecular research (Puente-Eelievre et al. 2016). Although these lineages were generally well-supported by the molecular data, not all of them could be diagnosed by unique morphological characters or character combinations. This led the researchers to adopt a conservative approach and accept a broad circumscription for Styphelia, inclusive of all 12 groups. As a result, the genera Astroloma, Coleanthera Stschegl. and Croninia J.M.Powell, together with a large number of the taxa previously included in Leucopogon R.Br., have now been subsumed into Styphelia (Crayn et al. 2019). The Styphelia xerophylla group comprises seven species and is thought to be exclusively Western Australian. Morphological assessment suggests that it is unlikely that other western or eastern Australian taxa, not included in previous molecular analyses, belong to this group. The earliest named species in the group, S. xerophylla and S. stomarrhena (Sond.) Sleumer, are also the most widespread, both having geographical ranges encompassing more than one bioregion. The other five have very restricted, sometimes disjunct, distributions in the Geraldton Sandplains, Jarrah Forest and Mallee bioregions. All five are of conservation concern, including two that have already been assessed as Threatened Flora (Smith & Jones 2018). One of these threatened species, Styphelia longissima Hislop & Puente- Eel., has only recently been deseribed (Hislop & Puente-Eelievre 2017) and is not treated again here. © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 156 Nuytsia Vol. 30 (2019) The phylogenetic tree topology obtained by Puente-Lelievre et al. (2016) indicates that the closest relative of the S. xerophylla group is the S. conostephioides (DC.) F.Muell. group (or Group VIII). Together the two groups are sister to Groups I-VII, which collectively encompass a large proportion of all Styphelia species. This paper is the first in a series detailing the morphological attributes of these groups and providing descriptions of their member species. Some of the groups comprise at least as many undescribed as described taxa, and a significant number of these are of conservation concern. A formal infrageneric classification will be published once the relationships of a number of mostly eastern Australian species are resolved. Further consideration of the best way to classify the morphologically heterogeneous Group X {sensu Puente-Lelievre et al. 2016) is also required. Methods This study was based on an examination of dried specimens housed at PERTH, together with field observations of the species described and their relatives in Western Australia. Foliar measurements and observations were taken from dried specimens in natural posture. Care was taken to confine observations to mature leaves. Inflorescence length was measured from the point of attachment at the axil to the tip of the bud-rudiment. Floral measurements were taken from rehydrated flowers in natural posture, with the exception of the corolla lobes which were uncurled to their fullest length before measuring. Fruit length is inclusive of a gynophore, if present. Bioregions referred to in the text and shown on distribution maps follow Interim Biogeographic Regionalisation for Australia (IBRA) v. 7 (Department of the Environment 2013). The Styphelia xerophylla group Morphological synopsis Leaves helically arranged; apex long-mucronate, pungent, although sometimes the mucro rather delicate and brittle; lamina adaxially concave; abaxial surface smooth to variously grooved, glabrous or variously hairy. Inflorescence 1- or occasionally 2-flowered; axis erect (widely spreading to ± pendulous in S. cernua Hislop & Puente-Eef), extending above floral node, or upper node if more than one flower is present, and terminating in a bud-rudiment; distal portion of axis, above the floral node, distinctly flattened, usually bract-like and often lobed on either side of the bud-rudiment; basal portion of axis, below the floral node with 4-5 sterile bracts, the lowest two opposite. Bracteoles conspicuously striate when dry, not keeled. Sepals usually conspicuously striate when dry (sometimes only faintly so in S. xerophylla), usually equal to, or longer than, the corolla tubes (always shorter in S. stomarrhena, occasionally shorter in S. xerophylla). Corolla usually white (red in S. stomarrhena). Corolla tube internal surface hairy in the upper part below the lobes, basal hair tufts usually absent (present only in S. stomarrhena), external surface usually glabrous (hairy in S. sulcata Hislop & Puente-Eef). Corolla lobes mostly erect in the basal 1/4-1/2 and then spreading and recurved above; outer surface glabrous or hairy (in S. stomarrhena and S. sulcata), inner surface usually densely hairy (sparsely and unevenly hairy in S. stomarrhena) with twisted, distinctly ornamented hairs. Anthers usually partially exserted from corolla tube and free from each other (fully exserted and cohering in S. stomarrhena). Filaments usually terete (plano-convex in S. stomarrhena), attached 1/2-3/4 above anther base (attached just below anther apex in S. stomarrhena), adnate to the corolla tube Just below the sinuses. Ovary usually M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 157 glabrous (hairy in S. longissima), 2-5-locular, pale to mid green in dried material. Nectary annular, truncate to variously lobed. Style usually scabrous or very shortly hairy, at least in the upper half (may be glabrous in S. stomarrhena), well exserted from the corolla tube in most species, with the stigma held at about the same level as the erect bases of the corolla lobes (greatly exserted in S. stomarrhena with the stigma held well beyond the erect corolla lobe bases); style base arising directly from ovary apex, not inset in a cylindrical depression and enveloped by ovarian tissue. Drupe variously shaped, longer or shorter than the calyx, circular in section, apex obtuse; mesocarp well-developed in most species (poorly developed and a gynophore present in S. sulcata). Important distinguishing features The relatively large, striate bracteoles and sepals found in the species of Group IX are an important diagnostic feature. Among the western groups only Group I (thQ Astroloma s. str species group) and the species-pair, S. crassiflora F.Muell. and Leucopogon sp. Badgingarra (R. Davis 421) from Group X, have similarly large and striate bracteoles and sepals. Members of Group IX can be distinguished from those of Group I by their white flowers (cf. red, orange, green or cream-coloured), in having corolla tubes that lack basal hair tufts {cf. basal hair tufts usually present), terete filaments {cf. usually distinctly flattened, sometimes compressed only) and corolla lobes of uniform texture (rather than bitextured). Styphelia stomarrhena is aberrant in regard to all except the last of these features. It is however easily recognised by its long-exserted anthers and hairy filaments. Its remarkable morphology is discussed below under the description of that species. From S. crassiflora and L. sp. Badgingarra, members of Group IX differ in having internal corolla tubes that are hairy in the upper part below the lobes {cf glabrous in this area), corolla lobe hairs twisted and strongly ornamented {cf straight and not or barely ornamented), and flat (rather than undulate) sepal margins. The sepal length alone is useful in recognising members of Group IX. Very few western Styphelia outside of Group I have sepals longer than those of S. disjuncta Hislop & Puente-Lel. (i.e. 4.2-5.3 mm long), the smallest flowered member of Group IX. Species other than those from Group I that may occasionally have longer sepals belong to the S. conostephioides group. Members of the latter group however can be readily distinguished from those of Group IX in always having partite nectaries {cf annular in Group IX), usually pendulous inflorescences {cf erect apart from S. cernua) and usually hairy ovaries {cf glabrous apart from S. longissima). Distribution The seven species that constitute Group IX are mainly distributed in near west coastal districts (within about 60 km of the coast) from the Kalbarri area to a little south of Perth and then with disjunct occurrences south and east as far as the Condingup area, east of Esperance. The Geraldton Sandplains bioregion is the main centre of distribution with four species. Key to species 1. Corolla red; anthers cohering, long-exserted from corolla tube on hairy filaments (Eneabba-eastern Perth suburbs). S. stomarrhena 1: Corolla white; anthers free, partially included within corolla tube, filaments glabrous 158 Nuytsia Vol. 30 (2019) 2. Inflorescence axis widely spreading to ± pendulous, 3.4-5.2 mm long; drupe ellipsoid, at least as long as the sepals (Kalbarri area).S. cernua 2: Inflorescence axis erect, 1.5-3.6 mm long (usually < 3 mm); drupe variously shaped but not ellipsoid, always much shorter than the sepals 3. Ovary densely hairy; at least some leaf margins densely long-ciliate with hairs 0.5-1.0 mm long (Eneabba area).S. longissima 3: Ovary glabrous; leaf margins glabrous, minutely ciliolate or irregularly ciliate with hairs <0.5 mm long 4. Leaf abaxial surfaces narrowly and deeply grooved, densely hairy in the grooves; ovary 2-locular; drupe cylindrical, gynophore present (Cascades area and NE of Condingup).S. sulcata 4: Leaf abaxial surfaces broadly and shallowly grooved, glabrous or sometimes sparsely and evenly hairy; ovary 3-5-locular; drupe obovoid, depressed-obovoid or globose, gynophore absent 5. Sepals 4.2-5.3 mm long; corolla tube 3.4-4.2 mm long; drupe obovoid (Dumbleyung and Ongerup areas).S. disjuncta 5: Sepals 6.5-9.2 mm long; corolla tube 6.0-8.2 mm long; drupe depressed- obovoid or ± globose 6. Style base abruptly differentiated from ovary apex; drupe depressed- obovoid (eastern Darling Range).S. angustiflora 6: Style base smoothly attenuated from ovary apex, such that it is difficult to tell where ovary ends and style begins; drupe ± globose or occasionally ovoid (Mount Adams-Watheroo-southern Perth suburbs.S. xerophylla Taxonomy Styphelia angustiflora Hislop & Puente-Lel, sp. nov. Typus\ south-west of York [precise locality withheld for conservation reasons], Western Australia, September 1997, J.L. Robson 657 {holo\ PERTH 04832353; iso\ CANB, K, MEL, NSW). Astroloma sp. sessile leaf (J.L. Robson 657), in G. Paczkowska & A.R. Chapman, West. Austral. Fl. \ Descr. Cat. p. 235 (2000); Western Australian Herbarium, inF/ora^t/^e, https://florabase.dpaw.wa.gov. au [accessed 21 February 2019]. Erect, compact shrubs to c. 80 cm high and 100 cm wide; fire-tolerance of rootstock unknown. Young branchlets with a moderately dense to dense indumentum of retrorse hairs, 0.05-0.20 mm long. Leaves spirally arranged, steeply antrorse to antrorse-appressed; apex long-mucronate, pungent, the mucro fine and rather brittle, 1.0-2.0 mm long; base obtuse to rounded; petiole well-defined, 0.3-0.6 mm long; lamina narrowly ovate to narrowly ovate-elliptic, 5.0-12 mm long, 1.3-3.0 mm wide, strongly concave adaxially, longitudinal axis gently incurved; surfaces discolorous, shiny; adaxial surface glabrous or with a few hairs towards the base, venation not evident; abaxial surface glabrous, paler, smooth, or shallowly grooved with 7-9 slightly raised primary veins and broad grooves between; the margins narrowly hyaline, at least towards the base, mostly glabrous or occasionally irregularly ciliate. Inflorescence axillary, erect; axis 2.5-3.5 mm long, 1 (2)-flowered, ± terete in lower portion, fiat and bract-like above the fertile node and terminating in a bud rudiment, sparsely hairy on the terete portion with a moderately dense indumentum on all surfaces of the fiat portion; flowers sessile. M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 159 Fertile bracts broadly ovate to ± orbicular, 1.4-2.0 mm long, 1.4-1.7 mm wide, subtended by 4 or 5 smaller, sterile bracts. Bracteoles ovate to elliptic, 2.6-3.7 mm long, 1.7-2.5 mm wide, obtuse with a very short sub-terminal mucro; abaxial surface striate, not keeled, shortly hairy, straw-coloured to pale brown when dry; margins ciliolate. Sepals narrowly elliptic or narrowly elliptic-obovate, 6.5- 9.2 mm long, 1.8-2.5 mm wide, acute and mucronate; abaxial surface striate, glabrous or with a very short, sparse indumentum towards the apex, straw-coloured when dry; adaxial surface with a well- defined patch of hairs towards the base and scattered, shorter hairs above; margins ciliate, the hairs to 0.2 mm long, the longer ones concentrated towards the base and apex. Corolla tube white, narrowly obovoid to ± cylindrical, shorter than the sepals, 62-1 mm long, 2.8-3.1 mm wide, glabrous externally, internal surface with hairs extending to a point well below the anther bases. Corolla lobes white, shorter than the tube, 5.0-5.7 mm long, 1.4-1.7 mm wide at base, erect for 1/3-1/2 of their length and then spreading and recurved; external surface glabrous, internal surface with a dense indumentum of twisted and ornamented hairs. Filaments terete, glabrous, 0.3-0.5 mm long, attached to anther c. 2/3 above the base, adnate to tube just below the sinuses. Anthers partially exserted from the tube (by 1/3-1/2 of their length), 2.2-3.4 mm long, apex shallowly emarginate. Nectary annular, 0.4-0.7 mm long, shallowly lobed, glabrous. Ovary mid-green, globose or depressed-globose, 1.2- 1.4 mm long, 1.2-1.4 mm wide, glabrous, deeply rugose, 3-5-locular. Style scabrous throughout, 6.5-9.1 mm long, abruptly differentiated from ovary apex, exserted from corolla tube with the stigma held at c. the level of the erect bases of the corolla lobes; stigma slightly expanded. Fruit depressed- obovoid, 2.3-2.5 mm long, 3.2-3.5 mm wide, much shorter than the calyx, shallowly rugose towards the apex, with 3-5 broad longitudinal ribs demarcating the suture lines on the endocarp; the apical surface ± fiat, scarcely descending towards the style base; gynophore absent. (Figure 1) Diagnostic characters. Within the S. xerophylla group, S. angustiflora is distinguished by the following character combination: leaf abaxial surfaces ± smooth to shallowly and broadly grooved; ovary glabrous, 3-5-locular; style abruptly differentiated from the ovary apex; fruit depressed-obovoid. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] I Oct. 1997, R. Davis 4220 (PERTH); 22 Aug. 1999, F Hort 543 (PERTH); 20 Sep. 1999, F&J. Hort 697 (CANB, NSW, PERTH); 16 Dec. 1999, F Hort 862 (PERTH); 7 Aug. 2000, F Hort 1105 (PERTH); 7 Sep. 2000, F Hort 1106 (CANB, NSW, PERTH); 16 Oct. 1987, J.L. Robson JER 12 (PERTH^ II Sep. 1999, L. W Sage, F Hort & J. Hort EWS 1565 (CANB, MEE, NSW, PERTH). Distribution and habitat. Known only from a small area of the eastern Darling Range, south-west of York, in the Jarrah Forest bioregion, where it occurs on sandy fiats in the understorey of Jarrah and Banksiav^JoodXdind. Commonly associated species includQBanksiaattenuata, B. menziesii,Adenanthos cygnorum, Allocasuarina humilis and Leptospermum erubescens. Phenology. Peak fiowering is between August and early October. The only specimen with mature fruit present was collected in mid-December. Etymology. From the Eatin angustus (narrow) and -florus (-fiowered), a reference to the noticeably narrower fiowers of this species relative to those of S. xerophylla, to which it otherwise bears a similarity in gross morphology. Conservation status. Eisted as Priority Two (Smith & Jones 2018) under Conservation Codes for Western Australian Flora under the name Astroloma sp. sessile leaf (J.E. Robson 657). It is known from very few populations, all of which are within the boundaries of Wandoo National Park. On the 160 Nuytsia Vol. 30 (2019) Figure 1. Styphelia angustiflora. A - habit; B - flowering branchlet in situ\ C - scanned image of flowering branchlet; D - fruit. Scale bars = 1 cm (C), 1 mm (D). Vouchers R. Davis 4220 (C), F. Hort 862 (D). Photographs by Fred and Jean Hort (A, B). Drawing by Skye Cofley (D). basis of current knowledge its geographic range is less than 12 km on a north-south axis and less than 5 km from east to west. Affinities. The topology of Group IX obtained by Puente-Lelievre et al. (2016) indicates that the closest relative of S. angustiflora is S. disjuncta. The two can be readily distinguished by the distinctly M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 161 larger leaves and floral parts of S. angustiflora (measurements for S. disjuncta given in parenthesis): leaves 5-12 mm long with a mucro 1-2 mm long {cf. 3.0-6.8 mm long with a mucro 0.4-1.0 mm long); sepals 6.5-9.2 mm long {cf. 4.2-5.3 mm); corolla tube 6.2-7.8 mm long {cf. 3.4^.2 mm); style 6.5-9.7 mm long {cf 3.9-5.2 mm). The fruits of the two species are also quite dissimilar. Those of S. angustiflora are wider than long with a more or less flat apical surface, while in S. disjuncta the drupe is significantly longer than wide and the apical surface ascends markedly towards the style base. Styphelia angustiflora also resembles S. xerophylla. Indeed when the first collection of the new species reached the Western Australian Herbarium in 1987, it was regarded as an outlier of that species. The most significant morphological difference between the two relates to the gynoecium. Whereas in S. angustiflora the style base is abruptly differentiated from the ovary apex, in S. xerophylla it is smoothly attenuated such that, in flowering specimens, it is not clear where ovary becomes style. The flowers of S. angustiflora are also noticeably narrower, with narrower sepals (1.8-2.5 mm wide, cf. 3.0^.3 mm in S. xerophylla), corolla tubes (2.8-3.1 mm wide, cf. 3.7-5.3) and corolla lobes (1.4-1.7 mm at base cf. 2.0-2.5 mm). Notes. The phrase name that had been in use for this species, A. sp. sessile leaf (J.L. Robson 657), was not an apt one. It was coined to highlight a perceived difference between the new species and S. xerophylla. Although the petioles of S. angustiflora are usually shorter and less conspicuous than those of S. xerophylla, the leaves are never truly sessile. Styphelia cernua Hislop & Puente-Lef, sp. nov. Typus\ north-east of Port Gregory [precise locality withheld for conservation reasons]. Western Australia, 7 July 2010, C. Puente-Lelitvre, M. Hislop & E.A. Brown CPL 63 {holo\ PERTH 08281998; iso\ CANB, K, MEL, NSW). Astroloma sp. Kalbarri (D. & B. Bellairs 1368), Western Australian Herbarium, in FloraBase, https:// florabase.dpaw.wa.gov.au [accessed 21 February 2019]. Erect shrubs to c. 1.7 m high and 1.5 m wide, from a fire-sensitive rootstock. Young branchlets with a sparse or moderately dense indumentum of ± patent hairs, to c. 0.05 mm long. Leaves antrorse, usually steeply so; apex long-mucronate, pungent, the mucro 0.8-1.5 mm long; base cuneate to attenuate; petiole well-defined, 1.0-1.7 mm long; lamina narrowly obovate to narrowly obovate-elliptic, 9.0-18.0 mm long, 2.0-3.2 mm wide, concave adaxially, longitudinal axis straight to gently incurved; surfaces distinctly discolorous; adaxial surface ± shiny, glabrous, except for a few hairs towards the base, venation not evident; abaxial surface paler, matt, grooved, with 7-9 raised, primary veins and narrow, prominent grooves between, shortly hairy in the grooves; margins narrowly hyaline, usually irregularly ciliate when young, the hairs soon abrading, leaving only the thickened bases, or occasionally the margins glabrous. Inflorescence axillary, widely spreading to more or less pendulous; axis 3.4-5.2 mm long, 1- or less often 2-flowered, ± terete in lower portion, flat and bract-like above the upper fertile node and terminating in a bud rudiment, with a moderately dense indumentum throughout; flowers sessile. Fertile bracts broadly elliptic, ± orbicular to transversely elliptic, 1.3-2.1 mm long, 1.5-2.0 mm wide, subtended by 4-5 smaller, sterile bracts. Bracteoles ovate, elliptic or ± orbicular, 2.6-3.5 mm long, 2.0-2.7 mm wide, obtuse to subacute, mucronate; abaxial surface striate, not keeled, shortly hairy, pale green in the basal half and straw-coloured above or straw-coloured throughout when dry; margins 162 Nuytsia Vol. 30 (2019) ciliate. Sepals narrowly ovate, 5.6-6.5 mm long, 2.5-2.8 mm wide, acute, mucronate; abaxial surface striate, glabrous or hairy with a short antrorse indumentum, straw-coloured when dry, sometimes with pink tinges; adaxial surface glabrous except for a discrete patch of hairs close to the base; margins ciliate with hairs to 0.4 mm long. Corolla tube white, ellipsoid or obovoid, about equal to, or a little shorter than the sepals, 4.5-5.8 mm long, 3.4-3.8 mm wide, glabrous externally, internal surface with hairs extending into the top of the tube below the lobes. Corolla lobes white, shorter than the tube, 2.6-3.2 mm long, 1.6-2.0 mm wide at base, erect for c.1/4 of their length and then spreading and recurved; external surface mostly glabrous but becoming papillate towards the apex, internal surface with a dense indumentum of twisted, strongly ornamented hairs. Filaments terete, glabrous, 0.4-0.5 mm long, attached to anther 2/3-3/4 above the base, adnate to tube just below the sinuses. Anthers partially exserted from the tube (by 1/4-1/3 of their length), 3.0-3.8 mm long, apex distinctly emarginate to ± truncate. Nectary annular, 0.6-0.9 mm long, shallowly lobed, glabrous or minutely papillate. Ovary pale green, globose or ellipsoid, c. 1.0-1.5 mm long (but refer comment under notes below), 1.0-1.5 mm wide, glabrous, 5-locular, with indistinct longitudinal ribs. Style scabrous or very shortly hairy in the upper half, c. 4.0-5.0 mm long (but refer comment under notes below), tapering gradually from the ovary apex, slightly exserted from the tube with the stigma held at c. the level of the erect bases of the corolla lobes; stigma slightly expanded. Fruit ellipsoid, 62-1.0 mm long and 4.0^.5 mm wide, slightly longer than calyx, the surface rugose with a markedly raised reticulum; gynophore absent. (Figure 2) Figure 2. Styphelia cemua. A - scanned image of flowering branchlet; B - fruit. Scale bars = 1 cm (A), 1 mm (B). Vouchers C. Puente-Lelievre, M. Hislop & E.A. Brown CPL 63 (A), A. Franks, S. Branigan & B. Smith BS 35 (B). Drawing by Skye Coffey (B). M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 163 Diagnostic characters. Within the S. xerophylla group, S. cernua is distinguished by the following character combination: leaf abaxial surface narrowly and prominently grooved with short hairs in the grooves; inflorescences widely spreading to ± pendulous; ovary glabrous, 5-locular; style tapering smoothly from ovary apex; fruit ellipsoid. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 31 July 1993, D. & B. Bellairs 1368 (PERTH); 17 May 1995, D. & B. Bellairs 1372 (PERTH); 9 July 1997, R. Davis 3669 (PERTH); 1 Nov. 2005, A. Franks, S. Branigan & B. Smith BS 35 (PERTH); 24 July 2008, M. Hislop 3779 (CANB, NSW, PERTH); 8 July 2010, C. Puente-Lelievre, M. Hislop & E.A. Brown CPE 63A (NSW, PERTH); 30 Aug. 2006, M Weir 100 (CANB, PERTH). Distribution and habitat. Styphelia cernua has a restricted distribution in and around Kalbarri National Park in the northern part of the Geraldton Sandplain bioregion (F igure 3). In this area it occurs on white or yellow sandplain in low, open woodland or heath. Commonly associated species include Banksia prionotes. Acacia scirpifolia, Daviesia divaricata, Jacksonia rigida and Scholtzia spp. Phenology. The main flowering period is between June and August. Although the only specimen with mature fruit was collected in early November, it could be expected that fruit would be present between September and November. 164 Nuytsia Vol. 30 (2019) Etymology. From the Latin cernuus (slightly drooping), a reference to the orientation of the flowers. This species being the only one in the group not to have an erect inflorescence. Conservation status. Recently listed as Priority Two under Conservation Codes for Western Australian Flora under the xmnQ Astroloma sp. Kalbarri (D. & B. Bellairs 1368) (Western Australian Herbarium 1998-). Styphelia cernua is currently known from less than 10 collections. Most of these are from a relatively small area south of Kalbarri National Park, with only two having been made within the park boundaries. Its distribution and frequency inside the park needs to be investigated. Affinities. This species is most similar to, and had previously been included in, Styphelia xerophylla. It differs from that species in its widely spreading to more or less pendulous inflorescences and smaller floral parts (measurements for S. xerophylla given in parenthesis): sepals 5.6-6.5 mm long {cf. 7.5-9.0 mm); corolla tube 4.5-5.8 mm long {cf. 6.0-8.2 mm); corolla lobes 2.6-3.2 mm long {cf. 4.0-5.2 mm). There is also a foliar difference with the abaxial leaf surfaces of S. cernua being deeply and narrowly grooved and shortly hairy within the grooves, whereas in S. xerophylla they are more shallowly grooved and either glabrous or with scattered hairs across the entire surface. The two species typically also have a different fruit shape: ellipsoid in S. cernua and usually ± globose in S. xerophylla (but refer to the notes heading under the latter for discussion of some variation that has been observed in regard to this character). The two species are allopatric with the southernmost population of S. cernua about a 160 km to the north-east of the most northerly known population of S. xerophylla. Notes. Because the style tapers so gradually from the ovary apex in this species the given measurements for the ovary and style lengths are necessarily imprecise. Styphelia disjuncta Hislop & Puente-Lel, sp. nov. Typus\ west of Lake Grace [precise locality withheld for conservation reasons]. Western Australia, 28 August 2001, M. Hislop 2272 {holo\ PERTH 05826616; iso\ CANB, MEL, NSW). Leucopogon sp. Ongerup (A.S. George 16682), Western Australian Herbarium, in FloraBase, https:// florabase.dpaw.wa.gov.au [accessed 21 February 2019]. Erect, compact shrubs to c. 60 cm high and 60 cm wide, branching from close to the base but with a Are-sensitive rootstock. Young branchlets with a dense indumentum of mostly retrorse hairs, 0.05- 0.15 mm long. Leaves steeply antrorse to antrorse-appressed; apex long-mucronate, with a rather delicate, scarcely pungent mucro, 0.4-1.0 mm long; base obtuse to rounded; petiole well-defined, broad, 0.3-0.5 mm long; lamina ovate to narrowly ovate, 3.0-6.8 mm long, 1.2-2.4 mm wide, strongly concave adaxially, longitudinal axis gently incurved; surfaces slightly discolorous, shiny; adaxial surface glabrous, or with a few scattered hairs towards the base and apex, venation not evident; abaxial surface glabrous, paler, shallowly grooved, with 7-9 raised primary veins and broad, shallow grooves between; margins of most leaves conspicuously hyaline (only those produced towards the end of a growth flush without hyaline margins), variably ciliate to ± glabrous. Inflorescence axillary, erect; axis 1.9-2.6 mm long, 1-flowered, ± terete in lower portion, flat and bract-like above the fertile node and terminating in a bud rudiment, sparsely hairy on the terete portion with a moderately dense indumentum on all surfaces of the flat portion; flowers sessile. Fertile bracts ovate, 1.3-1.8 mm long, 1.0-1.3 mm wide, subtended by 4 or 5 smaller, sterile bracts. Bracteoles broadly ovate, ovate or elliptic. M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 165 2.0-2.6 mm long, 1.7-1.8 mm wide, obtuse, with a very short sub-terminal mucro; abaxial surface striate, not keeled, shortly hairy, straw-coloured when dry; margins ciliolate. Sepals narrowly ovate- elliptic, 4.2-5.3 mm long, 1.6-2.0 mm wide, obtuse to acute, indistinctly mucronate; abaxial surface striate, with a sparse to moderately dense indumentum of short, antrorse hairs, straw-coloured when dry; adaxial surface with a well-defined patch of hairs towards the base and scattered hairs in the upper half; margins ciliate with hair to 0.2 mm long. Corolla tube white, narrowly obovoid or narrowly ellipsoid, shorter than the sepals, 3.3^.2 mm long, 1.8-2.0 mm wide, glabrous externally, internal surface with hairs extending below the lobes. Corolla lobes white, from shorter than to occasionally equal to the tube, 3.2^.0 mm long, 1.0-1.2 mm wide at base, erect for 1/3-1/2 of their length and then spreading and recurved; glabrous externally, internal surface with a dense indumentum of twisted, ornamented hairs. Filaments terete, glabrous, 0.4-0.5 mm long, attached to anther 1/2-2/3 above the base, adnate to tube just below the sinuses. Anthers partially exserted from the tube (by 1/2-2/3 of their length), 1.7-2.2 mm long, apex emarginate. Nectary annular, 0.4-0.5 mm long, shallowly lobed, glabrous. Ovary pale green, globose, 0.9-1.2 mm long, 0.9-1.1 mm wide, glabrous, deeply rugose, 3-locular. Style scabrous in the upper half, 3.9-5.2 mm long, well-differentiated from the ovary apex, exserted from the corolla tube with the stigma held at c. the level of the erect bases of the corolla lobes; stigma distinctly expanded. Fruit obovoid, 3.0-3.5 mm long, 1.8-2.2 mm wide, much shorter than the calyx, shallowly rugose with indistinct longitudinal grooves also evident; the apical surface distinctly raised towards the style base; gynophore absent. (Figure 4) Diagnostic characters. Within the S. xerophylla group, S. disjuncta is distinguished by the following character combination: short leaves (the longest < 7 mm long); leaf abaxial surfaces shallowly and broadly grooved; ovary glabrous, 3-locular; style abruptly differentiated from the ovary apex; fruit obovoid. Figure 4. Styphelia disjuncta. A - flowering branchlet in situ; B - scanned image of flowering branchlet; C - fruit. Scale bars = 1 mm (C), 1 cm (B). Vouchers M. Hislop 2272 (B), M. Hislop 3063 (C). Photograph by Jolanda Keeble (A). Drawing by Skye Coffey (C). 166 Nuytsia Vol. 30 (2019) Other specimens examined. WESTERN AETSTRAEIA: [localities withheld for conservation reasons] 27 June 2003, S. Barrett 1109 (PERTH); 1 Sep. 2005, J.A. Cochrane & S. Barrett JAC 5386 (K, PERTH); 16 Oct. 2007, J.A. Cochrane JAC 6871 (PERTH); 29 July 1986, A.S. George 16682 (CANB, PERTH); 12 Oct. 2003, M. Hislop 3063 (PERTH); 19 July 2016, M. Hislop 4610 (PERTH); 77 July 1965, K. Newbey 1808 (PERTH); 4 Aug. 1969, K. Newbey 2^33 (PERTH); 31 May 1970, K. Newbey 3184 (PERTH); 26 July 2002, L. Polomka & S. Patrick SP 4208 (PERTH). Distribution and habitat. Styphelia disjuncta has a restricted, disjunct distribution in the Dumbleyung and Ongerup areas at the western end of the Mallee bioregion. Grows in sand or sandy loam soils over laterite, and in association with species-rich heath or open mallee woodland. Phenology. The main flowering period appears to be between June and early September. Collections with mature fruit present have been made in September and October. Etymology. The epithet is derived from the Eatin disjunctus (separate, distinct), a reference to the significant geographic disjunction between the northern and southern populations. Conservation status. Eisted as Threatened Flora in Western Australia with a ranking of Vulnerable (Smith & Jones 2018) under the name L. sp. Ongerup (A.S. George 16682). Currently known only from three small populations, the two northern ones being around 100 kilometres distant from the southern population. Affinities. Styphelia disjuncta is morphologically most similar to, and only likely to be confused with, S. angustiflora. The differences between the two are given under S. angustiflora. Notes. Despite the significantly disjunct distribution, there is little morphological difference between the populations. Relative to those from the northern populations there is a tendency for plants from the southern population to have somewhat longer inflorescence axes and very slightly larger floral parts. Styphelia stomarrhena (Sond.) Sleumer, Blumea 12: 154 (1964); Astroloma stomarrhena Sond. in Eehm., PI. Preiss. 1(2): 301 (1845). Type'. In regionibus interioribus Australasiae occidentalis [Western Australia], dat.ffi. Preiss 410 {sym ED 1075918, MEE 1549325); In Australia occidentali [Western Australia], 1843, J. Drummond 467 (n.v.). Styphelia lasionema F.MuelL, Fragm. 6: 40 (1867). Type'. Swan River [Western Australia], dat., J. Drummond 475 {sym K 000277501, MEE 1549327); Ibidem ad oppidulum Hamdeni [Western Australia], dat., W. Clarke s.n. {sym. MEE 1549326). Eow, spreading, compact shrubs to c. 30 cm high and 40 cm wide, multi-stemmed from a fire- tolerant rootstock. Young branchlets with a moderately dense to dense indumentum of ± patent hairs, < 0.05-1.50 mm long. Leaves mostly steeply antrorse; apex long-mucronate, pungent, the mucro 1.3-2.0 mm long; base attenuate to cuneate; petiole well-defined, 0.7-2.0 mm long; lamina narrowly elliptic to narrowly obovate-elliptic, 10-23 mm long, 2.7^.5 mm wide, strongly concave adaxially, longitudinal axis gently incurved; surfaces discolorous; adaxial surface shiny, with a sparse or moderately dense indumentum, or occasionally glabrous, venation not or barely evident; abaxial surface paler, usually with a dimorphic indumentum consisting of a layer of sparse or moderately dense short hairs overtopped by a sparse layer of much longer, coarse hairs, occasionally one or both (i.e. surface glabrous) of the layers absent, openly grooved between 7-9 raised primary veins; M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 167 margins usually variably ciliate, sometimes ± glabrous. Inflorescence axillary, erect, 3.4-5.0 mm long, l(2)-flowered, ± terete and obscured by bracts in the lower portion, flat and bract-like above the fertile node and terminating in a bud-rudiment, flat portion hairy on all surfaces; flowers sessile. Fertile bracts elliptic, 2.8-3.8 mm long, 2.0-2.8 mm wide, subtended by 4 smaller sterile bracts. Bracteoles elliptic, 3.7-5.0 mm long, 2.4-3.3 mm wide, acute, mucronate; abaxial surface striate, not keeled, hairy with longer spreading hairs in the upper half, straw-coloured when dry; margins ciliate. Sepals narrowly elliptic or narrowly obovate-elliptic, 7.0-8.5 mm long, 2.6-3.8 mm wide, acute andmucronate; abaxial surface striate, with a moderately dense to dense indumentum of shallowly antrorse hairs, mostly in the upper half, straw-coloured when dry; adaxial surface glabrous throughout; margins ciliate at least in the upper half, with hairs 0.1-0.5 mm long. Corolla tube red, narrowly obovoid, much longer than the sepals, 10.2-16.5 mm long, 4.0-5.2 mm wide, glabrous externally, internal surface with 5 ± well- defined hair tufts close to the base and scattered hairs between the tufts and tube apex. Corolla lobes red, much shorter than the tube, 6.0-7.4 mm long, 2.2-3.0 mm wide at base, erect for up to c. 1/4 of their length and then spreading and recurved to revolute; external surface hairy in the upper half (the hairs with smooth surfaces); internal surface with a sparse indumentum of twisted and ornamented hairs, becoming denser towards the base and apex. Filaments plano-convex, stout, densely hairy with twisted, ornamented hairs, 4.4-5.5 mm long, adnate to the tube just below the sinuses, attached just below anther apex. Anthers well exserted from the tube, 2.0-2.8 mm long, cohering for much of their length, apex entire. Nectary annular, 0.6-1.0 mm long, shallowly lobed, glabrous. Ovary mid-green, globose to ovoid, 1.1-1.5 mm long, 1.1-1.3 mm wide, glabrous, 4- or 5-locular. Style glabrous or minutely scabrous immediately below the stigma, 14-24 mm long, well-differentiated from ovary apex, well-exserted from corolla tube; stigma lobed, much expanded. Fruit depressed-obovoid, with the upper surface ± flat except for a small raised area at the style base, 3.5-3.8 mm long, 4.2^.8 mm wide, much shorter than the calyx, the surface rugose; gynophore absent. (Figure 5) Selected specimens examined. WESTERN AUSTRALIA: Ellis Brook Valley Reserve [Orange Grove], 19 May 1999, H. Bowler 560 (PERTH); Brand Hv^, 6.9 km N of Greenhead Rd, 27 May 1997, R. Davis 3232 (PERTH); Cadda Rd, c. 11.5 km WSW of Badgingarra, 29 July 2012, R. Davis 12101 (PERTH); Lesueur National Park, S break of buffer to Mt Peron, 200 m E of Cockleshell Gully Rd, 20 June 1993, B. Evans WE 627 (PERTH); depressions near Cannington, June 1935, C.A. Gardner s.n. (PERTH); 8 km SW of Eneabba, 9 July 1977, E.A. Griffin 912 (PERTH); Irwin, 9 July 1986, R. Gueho 6 (PERTH); remnant bushland, adjacent ACTIV Industries, High Wycombe, 6 June 1998, M. Hislop 1060 (PERTH); Hi Vallee property (D. & J. Williams) Warradarge, above NW head of main valley, 15 July 2001, M Hislop, F&J. Hort MH 2256 (PERTH); Reserve No. 29801 E of Warradarge, along S internal firebreak adjacent to Greenhead-CoorowRd, 18 July 2004, M. Hislop 3281 (PERTH); Boonanarring Brook, 25 kmNNE of Gingin, 25 May 1988, GJ. Keighery 10050 (PERTH); 24.7 km E of Jurien Bay on road to Brand Hwy, 15 Aug. 1986, J.M. Powell 2522 (NSW, PERTH); White Rd Plot 1, c. 120 m from Kelvin Rd, and 80 m from White Rd, Orange Grove, 19 July 2006, J. Pryde & M. Hoskins MM 2 (PERTH); walking track of Badgingarra National Park on Brand Hwy, front of roadhouse petrol station, 6 July 2010, C. Puente-Lelievre & E.A. Brown CPE 50 (NSW, PERTH); Forrestfleld-Welshpool Rd, 0.5 km from Lewis Rd, in bushland patch N of road, 13 Aug. 2009, K.R. Thiele 3761 (PERTH); c. 13 km N of Jurien Rd junction on Cockleshell Gully Rd, 10 Aug. 1988, A.J.G. Wilson 128 (CBG, PERTH). Distribution and habitat. Styphelia stomarrhena has a scattered distribution in the southern Geraldton Sandplains and northern Swan Coastal Plain bioregions, with limited occurrence in the adjacent parts of the Avon Wheatbelt and Jarrah Forest bioregions; from a little south of Eneabba to the eastern suburbs of of Perth and east as far as the Chittering area (Figure 6). Grows in deep sandy soils or sand over laterite, mostly in Banksia woodland or various heathland communities. 168 Nuytsia Vol. 30 (2019) Figure 5. Styphelia stomarrhena flowering branchlet. Voucher K. Thiele 3761. Photograph by Kevin Thiele. An old, undated specimen (JJ. Havel s.n.) with no information other than the collection locality, ‘Coolup-Dwellingup Rd’, ostensibly represents a disjunct southerly occurrence for the species. However with such scant collecting information and no other collections of the species having been made south of Perth there would seem to be the possibility that this is a locality-error. Phenology. The main flowering period is between May and July. The only fruiting collections of the species were made in August, but presumably fruit is likely to be present at least until October. Conservation status. Although quite widespread the species is rarely common locally; its occurrence at any locality is mostly described as occasional. It is however well represented on the conservation estate and no conservation coding is recommended here. Notes. In the context of the morphology of Group IX as a whole S. stomarrhena is clearly anomalous in regard to a number of characters. Within the group it is the only species to have the following features (the usual character states given in parenthesis): red flowers (white); hair tufts present near the base of the internal corolla tube (hair tufts absent); anthers cohering, long-exserted from the corolla tube on hairy, stout, plano-convex filaments (anthers free, partially included within the tube on glabrous, slender, terete filaments); corolla lobes recurved to frequently revolute, hairs on internal surfaces unevenly distributed, mostly sparse but becoming denser towards the base and apex (corolla lobes recurved, never revolute, hairs on internal surfaces evenly distributed, dense); style much longer than the tube with the stigma ultimately presented well beyond the corolla lobes (style a little longer than M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 169 the tube with the stigma held at about the same level as the erect bases of the corolla lobes); rootstock fire-tolerant (fire-sensitive). The red fiower colour, long corollas (effectively made longer by the robust, densely hairy, exserted filaments), as well as differences in floral hair distribution, anther and stigma presentation strongly suggests that a particular pollination strategy is at work and has led to the morphological divergence exhibited by this species. Johnson (2013) has suggested that birds play a significant role in the pollination of epacrids with these features. This has been corroborated by field observations recorded by Michael Whitehead (pers. comm.) of the University of Melbourne during the course of recent research into the pollination systems of certain Styphelia species. During flowering the stigma is held in two different positions. When the flowers first open it is presented at the top of the cohering anthers, which dehisce introrsely against the upper style. At this stage the style is bent into angles within the corolla tube. Later, the style straightens so as to be finally well- exserted beyond the anthers. It seems likely that this sequence is protandrous in effect. There is an interesting difference in the shape of the ovary apex within the Western Australian Herbarium’s holding of this species. Some specimens have the ovary tapering quite smoothly to the style base, while in others the style arises from a well-defined flat surface at the ovary apex, so that the style base is conspicuously narrower than the ovary apex. There is no obvious geographical pattern to this variation, neither does it appear to correlate with any other potentially significant taxonomic 170 Nuytsia Vol. 30 (2019) difference. In order to evaluate whether this variation in ovary shape translates to a consistent difference in fruit shape it would be necessary to compare mature fruit of the two morphotypes. Currently there are only two fruiting collections of the species at the Western Australian Herbarium and they have very similar fruit, as given in the description above. Styphelia sulcata Hislop & Puente-Lel, sp. nov. Typus\ north of Cascade [precise locality withheld for conservation reasons], Western Australia, 16 May 2002, M. Hislop & F Hort 2598 {holo\ PERTH 06132804; iso\ CANB, NSW). Leucopogon sp. Bonnie Hill (K.R. Newbey 9831), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au [accessed 21 February 2019]. Low, compact shrubs to c. 40 cm high and 40 cm wide, from a Arc-sensitive rootstock. Young branchlets with a moderately dense to dense indumentum of patent to shallowly antrorse hairs, 0.05-0.5 mm long. Leaves steeply antrorse; apex long-mucronate, the mucro very fine and brittle, scarcely pungent, 1.0-2.4 mm long; base attenuate to ± cuneate; petiole well-defined, 0.3-0.6 mm long; lamina narrowly ovate, 4.0-8.2 mm long, 0.8-1.6 mm long, adaxially concave, longitudinal axis incurved; surfaces slightly discolorous, shiny; adaxial suface glabrous, the venation not evident; abaxial surface slightly paler, strongly grooved, with 5-7 raised primary veins and deep, narrow grooves between, shortly and densely hairy in the grooves, the raised veins glabrous; margins usually coarsely ciliolate with very short, antrorse hairs, occasionally ± glabrous. Inflorescence axillary, erect; axis 1.5-2.7 mm long, 1 -flowered, ± terete in lower portion, distinctly flattened above the fertile node and terminating in a bud rudiment, with a moderately dense indumentum throughout; flowers sessile. Fertile bracts narrowly ovate, 2.0-2.5 mm long, 0.9-1.3 mm wide, subtended by 4 smaller, sterile bracts. Bracteoles ovate, 2.8-3.5 mm long, 1.3-1.5 mm wide, acuminate, long-mucronate; abaxial surface striate, not keeled, shortly hairy, straw-coloured when dry, sometimes with pink tinges; margins ciliate. Sepals narrowly ovate, 4.8-5.5 mm long, 1.2-1.4 mm wide, acute and long-mucronate; abaxial surface striate, variably antrorse-hairy, at least in the upper half, straw-coloured, sometimes with pink tinges; adaxial surface hairy towards the apex and with a well-defined patch of hairs towards the base; margins ciliate with hairs to 0.3 mm long. Corolla tube white, narrowly obovoid to ± cylindrical, shorter than the sepals, 3.2^.2 mm long, 1.6-2.1 mm wide, external surface with a sparse to moderately dense indumentum in the upper half, glabrous below, internal surface with hairs extending to a point below the anther bases. Corolla lobes white, shorter than, to longer than, the tube, 3.2^.5 mm long, 0.9-1.1 mm wide at the base, erect for 1/4-1/3 of their length and then spreading and recurved; external surface with a sparse to moderately dense indumentum of antrorse hairs, internal surface with a dense indumentum of twisted and ornamented hairs. Filaments terete, glabrous, 0.2-0.3 mm long, attached to anther 1/2-2/3 above the base, adnate to tube just below sinuses. Anthers partially exserted from the tube (by 1/2-2/3 of their length), 1.7-2.3 mm long, apex emarginate. Nectary annular, 0.4-0.6 mm long, glabrous. Ovary pale to mid green, ellipsoid to obovoid, 0.8-1.0 mm long, 0.5-0.7 mm wide, glabrous, rugose, 2-locular. Style minutely scabrous throughout or at least in upper half, 3.2^.2 mm long, well- differentiated from ovary apex, slightly exserted from corolla tube, with the stigma held at a point c. level with the recurved bases of the corolla lobes; stigma slightly expanded. Fruit ± cylindrical, 3.2^.0 mm long, 1.2-1.6 mm wide, much shorter than the calyx, not rugose, with faint longitudinal striations; the apical surface ascending to the style base; a well-defined gynophore present. (Figure 7) Diagnostic characters. Within the S. xerophylla group, S. sulcata is easily distinguished by the deep, narrow grooves on the abaxial leaf surfaces, the hairy external surfaces of the corolla tube and lobes and a 2-locular ovary. M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 171 Figure 7. Styphelia sulcata. A - scanned image of flowering branchlet; B - flower, external view; C - fruit. Scale bars = 1 cm (A), 1 mm (C). Vouchers M. Hislop & F Hort 2598 (A), M. Hislop & F Hort 2598 (B), M. Hislop 4231 (C). Drawings by Skye Coffey (B, C). Other specimens examined. WESTERN AETSTRAEIA: [localities withheld for conservation reasons] 27 Sep. 1984, M.A. Burgman 3708 (PERTH); 24 May 1983, M.A. Burgman & S. McNee 1476 (PERTH); 13 Sep. 1992, G.F. Craig 2X12 (PERTH); 19 Sep. 1993, C.F. Craig2912 (PERTH); 21 Aug. 2012, M Hislop 4231 (PERTH); 16 May 2002, M. Hislop & F Hort MH 2600 (MEE, PERTH); 13 Oct. 2000, GJ. Keighery & N. Gibson 5597 (PERTH); 15 Nov. 1980, K. Newbey 8148 (PERTH); 10 May 1982, K.R. Newbey 9831 (PERTH). 172 Nuytsia Vol. 30 (2019) Distribution and habitat. Has a disjunct distribution, north of Cascades and north-east of Condingup, in the Mallee bioregion. Grows in sandy soils in open mallee woodland. Associated species include Eucalyptus leptocalyx, E. pleurocarpa, Calothamnus gracilis, Banksia media and Grevillea aneura. Phenology. Peak flowering is likely to be in May and June. Mature fruit is present on specimens collected in August and September. Etymology. The epithet is derived from the Latin sulcatus (furrowed), a reference to the narrowly and deeply grooved abaxial leaf surfaces. Conservation status. Priority One (Smith & Jones 2018) under Conservation Codes for Western Australian Flora under the name Leucopogon sp. Bonnie Hill (K.R. Newbey 9831). Styphelia sulcata is very poorly known and needs to be surveyed as a matter of priority. Because of imprecise label details it is not clear whether the collections from the western distribution node are from one or more populations. When the type collection was made in 2002 the species was very common locally, with at least many hundreds, and probably several thousand plants. However in 2012 when the first author revisited the site, there had been a wildfire in the intervening years and only occasional plants were seen. The eastern distribution node is currently represented by a single collection made in 1982; its precise locality is unknown. Although on the strength of our current knowledge there is cause for concern that this is a rare plant, it must also be acknowledged that both population nodes occur in remote areas of Western Australia containing large tracts of natural vegetation. There must therefore still be a good chance that future survey will bring to light new populations. Affinities. This species was not included in the molecular phylogenetic analysis of Puente-Lelievre et al. (2016), and its position as the seventh member of Group IX is based on morphology alone. Although possessing the core morphological attributes of the group, as described above, there is a significant difference in its fruiting character. Rather than having the usual fleshy drupe this species has a very-reduced mesocarp (more or less dry) and with a well-defined gynophore. None of the other species in the group has a fruiting gynophore although a very-reduced mesocarp is the norm in Group VIII, the sister group to Group IX. It is worth noting in this regard that Styphelia erubescens F.Muell. (until recently treated as Leucopogon oxycedrus Sond.) of Group V is a confirmed example of another species with a ± dry drupe and a fruiting gynophore that is nested within a group in which all of its closest relatives have a fleshy mesocarp and no gynophore. Styphelia sulcata is also the only species in the group in which the external surface of the corolla tube is hairy. Styphelia xerophylla (DC.) YM\xQ\\.,Fragm. 6: 38 (1867); StomarrhenaxerophyllaDC.,Prodr. 7(2): 738 (1839); Astrolomaxerophyllum (DC.) Sond. inLehm., PI. Preiss. 1(2): 301 (1845). Type: InNova Hollandiaad Swan River [Western Australia], 1835-1838, J. Drummonds.n. (holo: GDC G00464096). Erect, compact shrubs, to c. 60 cm high and 60 cm wide, branching from close to base but with a Are-sensitive rootstock. Young branchlets with a moderately dense to dense indumentum of patent to shallowly antrorse hairs, 0.05-0.60 mm long. Leaves variably antrorse or occasionally some leaves patent to shallowly retrorse; apex long-mucronate, pungent, the mucro 0.6-1.2 long; base attenuate, cuneate or sometimes rounded; petiole well-defined, 1.0-1.8 mm long; lamina narrowly ovate to ovate, narrowly elliptic to elliptic, or sometimes obovate, 5-20 mm long, 1.6-3.8 mm wide, strongly concave M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 173 adaxially, sometimes ± stem-clasping, longitudinal axis gently incurved, ± straight or occasionally gently recurved; surfaces discolorous shiny; adaxial surface glabrous or sparsely hairy, venation not evident; abaxial surface paler, glabrous or sparsely hairy with 5-9 raised primary veins, broadly grooved, or sometimes ± flat between the veins; margins glabrous or irregularly ciliate. Inflorescence axillary, erect; axis 2.5-3.6 mm long, 1 -flowered, ± terete in lower portion, flat and bract-like above the fertile node and terminating in a bud rudiment, sparsely hairy on the terete portion, with a moderately dense indumentum on the flat portion; flowers sessile. Fertile bracts orbicular, depressed-ovate or broadly ovate, 1.8-2.8 mm long, 2.1 -3.0 mm wide, subtended by 4-5 smaller, sterile bracts. Bracteoles broadly ovate to broadly elliptic to ± orbicular, 3.0-4.2 mm long, 2.4-3.6 mm wide, obtuse with a very short, sub-terminal mucro; abaxial surface striate, not keeled, glabrous or with spreading hairs, straw-coloured when dry, sometimes with a pinkish tinge; margins ciliolate. Sepals narrowly elliptic or narrowly ovate-elliptic, 7.5-9.0 mm long, 3.0^.3 mm wide, obtuse to subacute, shortly mucronate; abaxial surface finely striate, glabrous or sparsely hairy, straw-coloured when dry, sometimes with a pinkish tinge; adaxial surface with a well-defined patch of hairs towards the base; margins ciliate with hairs <0.05-0.20 mm long. Corolla tube white, obloid or narrowly obovoid, usually shorter than, but sometimes longer than, the sepals, 6.0-8.2 mm long, 3.7-5.3 mm wide, glabrous externally, internal surface with hairs extending to a point well below the anther bases. Corolla lobes white, shorter than the tube, 4.0-5.2 mm long, 2.0-2.5 mm wide at base; erect for basal 1/3-1/2 of their length and then spreading and recurved; external surface glabrous, internal surface with a dense indumentum of twisted, ornamented hairs. Filaments terete, glabrous, 0.6-1.0 mm long, attached to anther 2/3-3/4 above the base, adnate to tube just below the sinuses. Anthers partially exserted from the tube (by 1/3-2/3 of their length), 2.2-3.6 mm long, apex emarginate and often slightly recurved. Nectary annular, 0.4-0.8 mm long, truncate or lobed, glabrous or minutely papillate, longitudinally grooved below the sinuses. Ovary pale green, globose, c. 1.2-1.5 mm long (but refer comment under notes below), 1.2-1.4 mm wide, glabrous, strongly rugose, 5-locular. Style scabrous in the upper half, c. 6.2-8.0 mm long (but refer comments under notes below), tapering gradually from the ovary apex, slightly exserted from the corolla tube with the stigma held at c. the level of the erect bases of the corolla lobes; stigma slightly expanded. Fruit ± globose (but refer exception under notes below), 3.7^.2 mm long and 3.8-4.2 mm wide, much shorter than the calyx, the surface rugose, at least in the upper half; gynophore absent. (Figure 8) Selected specimens examined. WESTERN AUSTRAEIA: Gillingarra West Rd [NW of Mogumber], 4 Oct. 2006, C. Danese & D. Rayner B 1006-43 (PERTH); Brand Hwy (truck stop), 32 km N of Eneabba turnoff, 19 June 1997, R. Davis 3392 (PERTH); AMG 50JEE811721, W of S end of Moochamulla Rd, N of Moore River, 12 Aug. 1988, E.A. Griffin 4929 (MEE, PERTH); Alexander Morrison National Park, S side of Green Head-Coorow Rd, 18.3 km E of Brand Hwy, 13 Nov. 2004, M Hislop 3349 (CANB, NSW, PERTH); High Hill corner of Badgingarra National Park, on internal firebreak c. 150 m Wof corner, 26 July 2008, M Hislop 3788 (NSW, PERTH); Boothendarra Nature Reserve, off Boothendarra Rd 5.2 km E of Dewar Rd, NE of Badgingarra, 16 Aug. 2008, M. Hislop 3801 (CANB, NSW, PERTH); intersection of Mt Adams Rd and the gas pipe line [SE of Dongara], 3 Aug. 1994, E.D. Kabay 297 (PERTH); Chandala Nature Reserve, loppolo Rd, S of Gingin, 27 July 2005, G.J. Keighery 16680 (PERTH); site 23,2.25 km E along Wongonderrah Rd from the turn off to Nambung Homestead, then due S along track for 2 km, 29 Oct. 1999, C. MacPherson s.n. (PERTH); Gnangara-Moore River State Forest, Melaleuca block, 50 m N of a point 1360 m W along Quicke Rd from intersection of Quicke Rd and St Patrick Rd, Bullsbrook, 12.8 km ENE of Wanneroo, 25 Sep. 2008, D.A. Mickle & M.L. Swinburn 567 (PERTH); Reserve 28685 at Junction of Sundalara and Tomkins Rd, W of Arrino, 10 July 1991,5. Patrick 6\8(VEKT}1); Cooljarloo, S end of gravel reserve in Conservation Park C41986, 8 Oct. 1991, 5. Patrick 878 (PERTH); Jandakot area, 25 Aug. 1979, J.M. Powell 1320 (AD, CANB, CHR, NSW, NY, PERTH); Watheroo Rd, 28.7 km E of Brand Hwy, 174 Nuytsia Vol. 30 (2019) Figure 8. Styphelia xerophylla. A - flowering branchlet. B - fruit. Scale bar = 1 mm (B). Voucher M.E. Trudgen 20474 (B). Photograph by Rob Davis (A). Drawing by Skye Coffey (B). 14 Aug. 1986, J.M. Powell 2497 (HO, K, NSW, PERTH); Hi Vallee property along E track in the main valley, locality of Warradarge, 7 July 2010, C. Puente-Lelievre, M. Hislop & E.A. Brown CPE 57 (NSW, PERTH); 300 m into bush on E side, 1.1 km S on Amherst Rd from Warton Rd, Gosnells, 17 Aug. 1996, L. Sage 694 (PERTH); N of Gnangara Rd, towards SE part of Eot 47 Eexia Ave., locality of Ellenbrook, 30 July 1999, M. Trudgen & M. Trudgen MET 20006 (K, MEE, PERTH). Distribution and habitat. Widespread in the southern Geraldton Sandplains and northern Swan Coastal Plain bioregions, with limited occurrence in the adjacent parts of the Avon Wheatbelt and Jarrah Forest bioregions; from the Mount Adams area south to the southern suburbs of Perth and east as far as the Arrino area and Wannamal (Figure 3). Styphelia xerophylla grows in deep sandy soils or sand over laterite, mostly in Banksia woodland or various heathland communities. Phenology. The main flowering period is between June and September. Mature fruit is present on specimens collected between September and November. Conservation status. A common species, well represented on the conservation estate. Notes. On the uplands of the Dandaragan Plateau in the north-east of the species’ range there occurs a variant (e.g. M. Hislop 3001 & 3801) that differs from the typical form in having relatively shorter and broader leaves. The flower size of this variant is also consistently at the upper end of the range seen in the species as a whole, but no other qualitative floral or fruiting differences were identified during this study. M. Hislop & C. Puente-Lelievre, Taxonomic review of Styphelia xerophylla group (Ericaceae) 175 Styphelia xerophylla is a well-collected species with more than 160 specimens currently housed at the Western Australian Herbarium. These provide ample evidence that a globose fruit shape is the norm for the species across most of its range. However there are a few specimens that are aberrant in having ovoid drupes, distinctly longer than wide. One of these (J. D’alonzo 15) is from Karragullen in the Darling Range close to Perth. This collection is also noteworthy because it is the only one from laterite on the Darling Range, all other collections from the Perth region being from the deep sands of the coastal plain. Two other collections with ovoid fruit {E.A. Brown 97/129 & G. Taajfe., J.M. Powell 1363) have been made in the Eneabba area. No collections with mature fruit have been made from areas north of this area, which raises the possibility that ovoid fruit are the norm in the far north of the species’ range. Fruit shape is very often a diagnostic character for species of Styphelia and the presence of two fruit shapes in S. xerophylla is potentially of taxonomic significance, even in the apparent absence of correlating characters. Further research and targeted fieldwork is required to clarify the morphological and geographical pattern of variation in fruit shape before proper consideration can be given to whether segregate taxa should be recognised. Acknowledgements We would like to thank the following people for their assistance in the preparation of this paper: Skye Coffey for the line drawings, Steve Dillon for generating the distribution maps, Julia Percy-Bower for assisting with the specimen photographs and Kelly Shepherd for formatting the images to best effect. Fred and Jean Hort, Rob Davis, Jolanda Keeble and Kevin Thiele are also thanked for their images of live material. References Crayn, D.M., Hislop, M.& Puente-Lelievre, C. (2019). Aphylogenetic recircumscription (Ericaceae: Epacridoideae: Styphelieae). Australian Systematic Botany accepted (4 June 2019). Department of the Environment (2013). Australia’s bioregions (IBRA). IBRA 7, Commonwealth of Australia. http://www. environment.gov.au/land/nrs/science/ibra#ibra [accessed 21 February 2019]. Hislop, M., & Puente-Lelievre, C. (2017). Five new species of Styphelia (Ericaceae: Epacridoideae: Styphelieae) from the Geraldton Sandplains, including notes on a new, expanded circumscription for the genus. Nuytsia 28: 95-116. Johnson, K. A. (2013). Are there pollination syndromes in the Australian epacrids (Ericaceae: Styphelioideae)? Anovel statistical method to identify key floral traits per syndrome. Annals of Botany 112: 141-149. Puente-Lelievre, C., Hislop, M., Harrington, M., Brown, E.A., Kuzmina, M. & Crayn, D.M. (2016). A flve-marker molecular phylogeny of the Styphelieae (Epacridoideae, Ericaceae) supports a broad concept of Styphelia. Australian Systematic Botany 28: 368-387. Smith, M.G. & Jones, A. (2018). Threatened and Priority Floralist 5 December 2018. Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 21 February 2019]. Western Australian Herbarium (1998-)- FloraBase—the Western Australian Flora. Department of Biodiversity, Conservation and Attractions, https://florabase.dpaw.wa.gov.au [accessed 7 February 2018], 176 Nuytsia Vol. 30 (2019) Nuytsia The journal of the Western Australian Herbarium 30: 177-194 Published online 30 July 2019 Elionurus tylophorus (Poaceae: Paniceae: Andropogoneae), a new species from the Kimberley region of Western Australia Matthew D. Barrett^’^’^’^ and Tricia Handasyde'^ ‘Kings Park and Botanic Garden, Department of Biodiversity, Conservation and Attractions, 1 Kattidj Close, Kings Park, Western Australia 6005 ^Western Australian Herbarium, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 ^School of Biological Sciences, The University of Western Australia, 35 Stirling Highway, Crawley, Western Australia 6009 ''Department of Biodiversity, Conservation and Attractions, PO Box 942, Kununurra, Western Australia 6743 ^Corresponding author, email: matthew.barrett@dbca.wa.gov.au Abstract Barrett, M.D. & Handasyde, T. Elionurus tylophorus (Poaceae: Paniceae: Andropogoneae), a new species from the Kimberley region of Western Australia. Nuytsia 30: 177-194 (2019). A new annual species, Elionurus tylophorus M.D.Barrett & T.Handasyde, is described in the genus Elionurus W\\\d. of the grass tribe Andropogoneae. It differs from all previously described Elionurus species in having keels of the lower fertile glumes ornamented with projections, these often terminated by minute solitary bristles, rather than regular cilia lacking projections, or with projections terminated by hair-tufts in other species. It differs further from E. citreus (R.Br.) Benth. and E. purpureus E.J.Thomps., the only other Australian Elionurus, in having strongly developed fertile glume nerves, rachilla longer and lacking a wing, and presence of lemmas and anthers in the pedicellate spikelet. Elionurus tylophorus is currently known from a single location in the King Leopold Ranges in the central Kimberley region, and has conservation significance. A primary division of Elionurus into two clades is hypothesised based on a preliminary chloroplast gene phylogeny and two morphological synapomorphies, presence of tufted glume hairs and oil streaks. Introduction The purpose of this paper is to describe a previously unknown species of Elionurus Willd. that was recently discovered in tropical Australia. The genus Elionurus belongs to the largely tropical Poaceae tribe Andropogoneae Dumort. and contains 16 species (Renvoise 1978; Kellogg 2015; Thompson 2017) distributed through Africa, the Arabian Peninsula, India and the Americas, with two disjunct species, E. citreus (R.Br.) Benth. and E. purpureus E.J.Thomps., occurring in tropical Australia and Papua New Guinea (Thompson 2017). Elionurus is absent from south-east Asia, the closest records to Australia being in the Middle East and India. Elionurus belongs to the distinctive tribe Andropogoneae of subfamily Panicoideae Link, and shares the dominant tribal characters of paired spikelets (one fertile and sessile, the other pedicellate and usually sterile or staminate), disarticulating rachis and C4 photosynthesis (Kellogg 2015). Most species of Andropogoneae have twisted and hygroscopic lemma awns (possibly an apomorphy for the core Andropogoneae), hui Elionurus is one of the few Andropogoneae genera to lack lemma awns. Although © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 178 Nuytsia Vol. 30 (2019) molecular phylogenies(e.g. Skendzic^rr//. 2007; Teerawatananon err//. 2011) strongly support as a member of the Andropogoneae, the relationship of Elionurus within the tribe is uncertain and it is unplaced to subtribe in Kellogg (2015) and treated as Andropogoneae incertae sedis. Soreng et al. (2017), in their phylogenetic classification of Poaceae, place Elionurus in subtribe Tripsacinae Dumort., but many aberrant species of basal Andropogoneae have not yet been sequenced. It should be noted that a significant proportion of species and genera of tribe Andropogoneae are of allopolyploid origin and are difficult to treat in a hierarchical classification based on the principal of monophyly (Estep et al. 2014), but Elionurus species have not been investigated for allopolyploid origins. Thompson (2017) provided a detailed discussion of morphological relationships with other genera and identified the presence of a proximal beak on the sessile spikelet as a key distinguishing character for Elionurus, which amongst the candidate close relatives is shared only with Urelytrum Hack. Most Elionurus species can also be identified to genus by the presence of one or two oil streaks on the lower glumes, but this is lacking in three species (Thompson 2017). Important morphological characters delimiting from other Andropogoneae genera (following Kellogg 2015; Thompson 2017) are: infiorescences unbranched above final leaf blade, rachis fragmenting at maturity, spikelets paired with one sessile and fertile, the other pedicellate and sterile or male, (lower) glume coriaceous to indurated and thicker than the lemmas, fertile spikelets dorsiventrally compressed, lower glume of sessile spikelet with a bifid apex and two distant keels adjacent to an oil streak (streak absent in three species) and with cilia in lines or tufts along at least part of the keel, lemmas lacking awns, and both paleas absent. Most but not all species of Elionurus have two distinctive lobes or awns at the apex of the lower glume. Elionurus citreus is a perennial (or sometimes annual?) species that occurs in tropical and subtropical Australia from the Kimberley to Queensland and south to New South Wales, and also in southern Papua New Guinea (Simon 1992; Simon & Alfonso 2011-). Elionurus citreus was considered the only Australian representative of the genus until Thompson (2017) described E. purpureus, an annual species known only from Cape York in Queensland. Some Elionurus species (e.g. E. elegans Kunth) have been characterised for the scent-producing volatiles. The Australian species E. citreus and E. purpureus have lemon-scented foliage and are known as Lemon Grasses or Lemon-scented Grasses. In2004, during a Department of Conservation and Land Management and Tropical Savannas Cooperative Research Centre survey in the King Leopold Ranges area of the Kimberley region of Western Australia (Start et al 2012), an unusual grass was collected at one of the survey sites by T. Handasyde (TH). The specimen could not be confidently placed to genus, so it was sent to M. Lazarides in Canberra who suggested further material would be required to determine the genus. Subsequently, TH organised collection of further material from the same site in 2015 and 2017. Examination of both collections by M. Barrett (MB) confirmed that the material belonged to the genus Elionurus, possessing all the diagnostic characters of the genus as described above; however, the King Leopold specimens could not be accommodated in any known species (reviewed by Renvoise 1978 and Thompson 2017). It is therefore described below as a new species, E. tylophorus M.D.Barrett & T.Handasyde. Methods Morphology Gross morphology of four accessions of E. tylophorus held at PERTH, including the proposed type, were examined in detail. Images of the types of E. citreus and E. purpureus were examined, as were all E. citreus collections held at PERTH and DNA. M.D. Barrett & T. Handasyde, Elionurus tylophorus (Poaceae), a new species from WA 179 DNA sequencing In order to place the new species phylogenetically, DNA sequences were generated for two chloroplast regions, ndhY and frwK intron (including the matK gene), and one nuclear rRNA region, the internal transcribed spacer (ITS), from the holotype specimen of E. tylophorus. All available Elionurus sequences from these regions were downloaded from GenBank, along with a range of outgroups from the tribe Andropogoneae (Table 1). Extraction of genomic DNA was performed using a method modified from Doyle and Dickson (1987). The Carlson extraction buffer (Csaikl et al. 1998) contained 2% cetyltrimethylammonium bromide (CTAB), 1 % polyethylene glycol (PEG), 1.4 M NaCl, 100 mM Tris-HCl and 20 mM EDTA (diluted from 0.5M pH 9.5 stock). Approximately 0.1 g of tissue was ground in 1 mE of buffer with 3 pE of 100 mg/mE RNase (Qiagen, Melbourne, Victoria, Australia) added before being incubated at 65°C for 15 mins. Samples were then centrifuged at 11,500 rpm for 10 mins, and the supernatant transferred to a new tube, with 1 vol. of chloroform: isoamyl alcohol (24:1) added before being shaken for 30 mins. After centrifuging at 9,000 rpm for 10 mins, DNA was precipitated from the supernatant by adding 1 vol. of cold (-20°C) isopropanol to each sample and stored at -20°C in a freezer for 20 mins before pelleting the DNA in a centrifuge at 13,000 for 10 mins. The pellet was then washed in 500 pE of 70% ethanol, re-centrifuged at 13,000 for 10 mins, and air-dried after removal of the ethanol. Polymerase chain reactions (PCR) were performed on an Applied Biosystems Veriti® thermocycler, under the following conditions: 95°C for 3 mins, 37 cycles of ((95°C for 30 s, 48°C for 30 s, 72°C for 1.5 mins), 72°C for 10 mins). PCR amplifications were performed in 20 pE volumes, and consisted of a final concentration of 67 mM Tris-HCl (pH 8.8 at 25°C), 16.6 mM (NH 4 ) 2 S 04 ,0.45% Triton X-100, 0.2 mg/mE gelatin, 0.2 mMof each dNTP (all diluted 1:4 from 5x polymerisation buffer, Fischer Biotec, Subiaco, Western Australia), 2.5 mM MgCl 2 , 0.12% v/v DMSO, 0.06% v/v BSA, and 0.2 units Taq DNA polymerase (Fischer Biotec, Subiaco, Western Australia), 0.5 pM final concentration of each primer per reaction and c. 10-500 ng of template DNA. Primers used for DNA sequences were: ITS - 18SF / 26SR (Prince 2010); matY. - trnKmatKF / 2545R (Aliscioni et al. 2012); ndhY - 1311F / 209IR (Peterson et al. 2010). Cycle sequence reactions were done using BigDye® Terminator v. 3.1 chemistry (Applied Biosystems) with the same primers as used for the PCR reaction, in both the forward and reverse directions, using the following PCR protocol: 96°C for 1 min, 25 cycles of (96°C for 10 s, 50°C for 5 s, 60°C for 4 min). Fragment visualisation was performed using an Applied Biosystems 3500 Genetic Analyser. Chromatograms were manually edited using CodonCode Aligner v.7.0.1 (CodonCode Corporation, Dedham, MA, USA, http://www.codoncode.com/). Comparisons to other sequences were made following alignment with sequences downloaded from GenBank (http://www.ncbi.nlm.nih.gov/) in the sequence database software Geneious® v.6.1.7 (http://www.geneious.com/; Kearse etal. 2012). Phylogenetic analyses were performed separately for (1) a concatenation of ndhY and rrwK intron (including the matK gene) and (2) ITS, since the corresponding regions were only available for the same Elionurus sample in a few cases (Table 1). Outgroups were chosen to represent several different groups of Andropogoneae, since no close relatives of Elionurus have been identified in phylogenies. Phylogenetic trees were reconstructed using RAxME (Stamatakis 2014), using a rapid bootstrapping and search algorithm, with 100 bootstrap replicates and a GTR+GAMMA nucleotide substitution model, as implemented in Geneious® v.6.1.7 (http://www.geneious.com/; Kearse et al. 2012). 180 Nuytsia Vol. 30 (2019) Table 1. Sequences used for phylogenetic analyses and GenBank numbers. Samples in bold were generated for this study. Species Voucher ndhF truK / matK ITS Reference Chrysopogon gryllus RI. Kellogg 250984 (A) (Rep. of Macedonia) KY596161 KY59616E Arthan et al. (2017) 0. Neamsuvan 165 (BCU) GQ856347 Neamsuvan et al. (unpublished) Elionurus citreus SJ. & T.R. Hodkinson 9561 (TCD) (Australia) HE573560 HE574449 GQ870207 Aliscioni etal. (2012); Teerawatananon etal. 2011 K.R. McDonald KRM 14437 (BRI AQ914411) (Australia) MK396908 This paper K.R. McDonald KRM 15855 (BRI AQ915512) (Australia) MK396910 This paper Elionurus muticus J.M. Kimeu et al JMK 145 (EA, K) (Kenya) MF998637 MF998984 Hackel et al. (2018) Not specified AF190758 Spies & Kellogg (unpublished) Elionurus purpureus P.l. Forster PIF 45151 (BRI AQ941416) (Australia) MK396909 This paper Elionurus royleanus MX Vorontsova et al. MSV 826 (EA, K) (Kenya) MF998646 MF998985 Hackel et al. (2018) Elionurus tripsacoides Manrique 1895 (COCA) (Mexico) DQ005046 Skendzic et al. (2007) Elionurus tripsacoides Manrique 1904 (COCA) (Mexico) DQ005047 Skendzic et al. (2007) M.D. Barrett & T. Handasyde, Elionurus tylophorus (Poaceae), a new species from WA 181 Species Voucher ndhF truK / matK ITS Reference Elionurus tristis M.S. Vorontsova et al MSV 589 (K, TAN) (Madagascar) LN908091 LN906693 Hackel et al. (2018) Elionurus tylophorus T. Sonneman TH 8073 (PERTH, holotype) (Australia) MK097174 MK097175 MK095935 This paper Hemisorghum mekongense Traiperm 569 (BKF) (Thailand) KY596132 KY596132® Arthan et al. (2017) Miscanthus transmorrisonensis Not specified Hodkinson 20 (K) LN869229 LN869229® AYl16271 Lloyd & Joshi (unpublished) Hodkinson et al. (2002) Themeda villosa Arthan 065, (BKF) (Thailand) E00697528 (China) KY596131 KY59613F KY991079 Arthan et al. (2017) Dunning et al. (2017) Tripsacum dactyloides Not specified (China) Sanchez-Ken 607 (ISC) (Mexico) NC_037087 NC_037087® DQ005086 Wang et al. (2017) Skendzic et al. (2007) ® Extracted from a plastid genome, hence ndhY and frwK / matK have the same accession number. Results The chloroplast RAxML tree (Figure 1 A) supported the monophyly of all sampled Elionurus species (100% bootstrap (BS) support). Elionurus royleanus AKioh. was recovered as sister to all remaining species, but with low support (70% bootstrap, hereafter BS) support. Within the core Elionurus clade resolution was poor, and only a sister relationship between E. muticus (Spreng.) Kunth and E. tristis Hack, was supported (82%). In the ITS RAxML tree (F igure 1B), Elionurus was not recovered as monophyletic due to the unresolved position of the single E. muticus sample. The E. muticus sample is very divergent from all other ITS sequences, and is likely a misidentified or contaminated sample, given that a dilferent specimen of E. muticus is placed within the core Elionurus clade on chloroplast data. A clade containing all remaining Elionurus samples was strongly supported (100% BS), in the ITS tree, and divided into two sub-clades, one containing the two E. tripsacoides Kunth samples (100% BS), the other clade containing E. tylophorus sister to E. citreus (77% BS). 182 Nuytsia Vol. 30 (2019) Figure 1. Phylogenies of Elionurus based on chloroplast and nuclear sequences. A - phylogenetic tree of ten chloroplast ndhV and trwK-intron sequences showing the monophyly of Elionurus and sister relationship of E. royleanus and all other Elionurus species; the position of two potential apomorphies for basal sister clades of Elionurus are indicated; B - phylogenetic tree of twelve ITS sequences showing the monophyly of Elionurus, a well-supported E. citreus / E. purpureus clade, and variation between E. citreus samples. See text for further discussion. M.D. Barrett & T. Handasyde, Elionurus tylophorus (Poaceae), a new species from WA 183 Discussion Morphological comparison and evidence for a new species A detailed summary of key features of the Elionurus collections are listed in Table 2. From this data, there are numerous discrete differences to both E. citreus and E. purpureus, spread across all parts of the plant. Thompson (2017) identified several apparently apomorphic features that unite the Australian species E. citreus and E. purpureus: pedicellate spikelets neuter (lacking anthers), and rachillas chartaceous, inflated and usually with a distal, bilobed fiange. In comparison, all other Elionurus species, including E. tylophorus have male pedicellate spikelets and cartilaginous, slender rachillas lacking a fiange. Elionurus tylophorus therefore is expected to lie outside of a presumed E. citreus+E. purpureus clade, but with unknown affinities to extra-Australian species. Elionurus tylophorus can be accommodated in the Appendix 2 table of Thompson (2017) comparing all species of Elionurus, by listing the following character states: paired spikelets dissimilar; sessile spikelet: lower glume margin hairs pectinate (but the lower ones tubercle-based, resembling the tufted condition but with only a single apical hair); lower glume submargins with an oil streak; lower glume apex attenuate and 2-lobed; pedicellate spikelet: fioret male; lower glume asymmetrical; lower glume margins with an oil streak on both margins; lower glume margins 2-keeled; lower glume apex long- attenuate, 1-lobed/awned. This combination of characters is unique, but most similar to E. hensii K.Schum. and E. muticus. Elionurus hensii shares with E. tylophorus the annual habit (according to Renvoise (1978), although Clayton et al. (2018) describe it as perennial), but differs in having smaller spikelets 3-4 mm long. Elionurus muticus is a perennial with short rhizomes and mostly basal leaves, and has spikelets in a pair similar to each other, while E. tylophorus is annual with cauline leaves and spikelets in a pair dissimilar. The projections on the margins of the lower glume of the sessile spikelet of E. tylophorus are found in three other species, E. elegans, E. hirtifolius Hack, and E. royleanus (Renvoise 1978; Thompson 2017 Figure A3.1, A3.2), which might possibly indicate a relationship. However, these species all have tufts of hair on the submarginal projections of the sessile spikelet lower glumes, lack oil streaks on the glumes and have 1-keeled lower glume of the pedicellate spikelet. In contrast, E. tylophorus has submarginal projections of the sessile spikelet lower glumes with at most a single hair, lower glumes of both sessile and pedicellate spikelets with two oil streaks, and lower glume of the pedicellate spikelet 2-keeled. E. tylophorus therefore differs in multiple characters from all described species of the genus, indicating that species rank is appropriate for the new taxon. Keys are provided below to Elionurus in Australia, and to place E. tylophorus within the global key of Renvoise (1987). Different characteristics of E. tylophorus weakly suggest relationships with various other species in the genus, and true affinities will require molecular data. Key to Australian species of Elionurus 1. Keels (submarginal) of lower glume of sessile spikelet with a row of prominent projections often terminated by a single minute bristle, the projections grading into long cilia near glume apex; rachilla not winged at apex; pedicellate spikelet with lemmas and anthers.E. tylophorus 1: Keels (submarginal) of lower glumes of sessile spikelet pectinate (regularly ciliate), lacking projections; rachilla usually winged at apex; pedicellate spikelet lacking lemmas and anthers.2 184 Nuytsia Vol. 30 (2019) 2. Annual; sessile spikelet 8.1-9.2 mm long; lower glume of sessile spikelet with lobes shorter than body, purple at maturity; lower and upper glumes of pedicellate spikelet 5-veined.E. purpureus 2: Perennial; sessile spikelet 10.6-13.1 mm long; lower glume of sessile spikelet with lobes longer than body, pallid to pale pink, rarely purplish; lower and upper glumes of pedicellate spikelet 3-veined.E. citreus Elionurus tylophorus can be included in the global key to Elionurus species of Renvoise (1978) by adding the following couplet at the start of the key (see Thompson (2017) for the addition of E. purpureus to the Renvoise (1978) key): 1. Keels (submarginal) of lower glume of fertile spikelet with a row of prominent projections, glabrous or terminated by a single minute bristle, the projections grading into long cilia near glume apex.E. tylophorus 1: Keels (submarginal) of lower glumes of fertile spikelet with longer cilia either regularly arranged and lacking projections, or if with projections these terminated by hairs in tufts.Go to step 1 in Renvoise (1978) Table 2. Distinguishing morphological characters between the three Australian Elionurus species (after Thompson 2017). Character Elionurus citreus Elionurus purpureus Elionurus tylophorus Growth habit perennial (or occasionally annual annual annual according to collector notes and herbarium specimens) Inflorescence type Culms (shape in T.S.) Leaf blades and sheath Distribution Margin prickle hairs Margins and mid-vein indumentum Mid-vein Sessile spikelet (mid-raceme) Total length (beak, body and lobes) (mm) Colour at maturity single racemes at nodes on cultivated plant and topotypes narrowly convexo- concave mostly basal medium (60-70 um) usually scabrous acutely keeled 10 . 6 - 12.1 pallid to pale pink, rarely purplish multiple racemes at nodes at least on cultivated plants broadly convexo-concave cauline absent smooth obtusely keeled 8.1-9.2 purple multiple racemes at nodes on wild-collected plants broadly convexo-concave cauline present, to 1.5 mm long smooth iobtusely keeled 9.0-13.2 green turning straw with purplish margins or pale purplish all over Lower glume lobe length (mm) 5.5-7 2.6-3.2 2.0-4.5 M.D. Barrett & T. Handasyde, Elionurus tylophorus (Poaceae), a new species from WA 185 Character Elionurus citreus Elionuruspurpureus Elionurus tylophorus 1.5-1.7 X 3.6-4.3 1.7-2.0 X 3.5-4.2 1.4-2.0 X 7.0-12.0 body width x length (mm) texture nerve prominence indumentum type orientation hair length (mm) keeled margins Upper lemma margin texture marginal indumentum apex Callus length (mm) Pedicellate spikelet Lower glume width X length (mm) colour at maturity body Upper glume veins shape in section Lemmas Anthers Anther length Chasmogamous (mm) Cleistogamous (mm) Pedicellate floret Rachilla Width X length (mm) Shape cartilaginous not raised glabrous to pilose ascending c. 2 narrowly winged hyaline ciliate apically or glabrous attenuate 1.8-2.7 c. 0.9 X 7.2-8.5 pallid to pale pink, rarely purplish linear to narrow lanceolate asymmetrical 3-veined 1-keeled one margin with oil streak unequal to lower glume 3-veined laterally compressed absent absent 1.3-2.7 0.7-1.8 absent 0.9-1.3 X 3.3-5 clavate chartaceous not raised pubescent appressed c. 0.2 not winged hyaline pilose acute 1.0-2.3 0.8-L3 X5.2-5.7 purple lanceolate asymmetrical 5-veined 1-keeled both margins with oil streak subequal to lower glume 5-veined rounded absent absent 0.9-2.0 c. 0.8 absent L2-L4 X4.3-4.4 distinctly clavate cartilaginous strongly raised pubescent in rows between nerves ascending (appressed) 0.05-0.25 very narrowly winged pale straw isparely ciliate attenuate 1.5-2.5 0.8-1.2x9-11.7 green turning straw with purplish margins or pale purplish all over lanceolate to narrowly lanceolate asymmetrical 7-veined 2-keeled both margins with oil- streak subequal to lower glume obscurely 3-5-nerved rounded to laterally compressed present present 2.5-3.0 not seen 2.2-3.8 0.9-L2 X 7.5-16, parallel-sided 186 Nuytsia Vol. 30 (2019) Character Elionurus citreus Elionurus purpureus Elionurus tylophorus Apex rim usually narrowly winged and unequally bilobed, or not winged distinctly winged, unequally bilobed entire, not winged, not bilobed Subapical beard length of longest hairs (mm) Pedicel 3.6-4.9 2.8-2.9 2.5-4.5 Width X length (mm) 0.5-0.6 X 2.9-4.0 c. 0.8 X 2.5-3.5 0.6-0.8 X 4.1-6.2 Phytogeny o/Elionums In the chloroplast ndhY + tree (Figure 1 A), E. royleanus was found sister (70% BS) to all other Elionurus species, including E. tylophorus. Elionurus royleanus was the only sampled representative of three similar species with tufted hairs on the submargins of the lower glume of the sessile spikelet (the others being F. elegans mdE. hirtifolius, hereafter the ‘tufted group’). These unique hair-tufts, along with other similarities (see Appendix 2 in Thompson 2017) suggest that the ‘tufted group’ might be a monophyletic clade sister to the remaining ‘core group’ of Elionurus; if so the presence of 1 or 2 oil streaks on the lower glumes would then be an apomorphy for the ‘core group’ (oil streaks are lacking in the ‘tufted group’; Thompson 2017). The presence of glume hair tufts and oil streaks might then be reciprocal apomorphies for a primary dichotomy within Elionurus. The position of E. tylophorus remote from E. royleanus suggests that the tuberculate projections on lateral submargins of the lower glume of the sessile spikelet found in the ‘tufted group’ and E. tylophorus are not indicative of a close relationship, but instead arose through convergence or homoplasy. The relationship between E. tylophorus and E. citreus was insufficiently resolved, and too many potentially intermediate species remain unsampled, to draw any conclusions about the monophyly or otherwise of Australian from the chloroplast data. In the ITS tree (Figure 1B), the available samples of Elionurus are supported (90% B S) as a monophyletic group, however no species of the ‘tufted group’ discussed above have ITS sequences available. A clade composed of the three representatives of E. citreus and one E. purpureus was strongly supported (90% BS), to the exclusion of E. tylophorus. This E. citreus clade is in agreement with the morphological evidence presented in Thompson (2017) for a close relationship between E. citreus and E. purpureus, and also corroborates the distinctiveness of E. tylophorus from the other Australian species. Two of the E. citreus specimens {K.R. McDonald 15855 and S.J. & T.R. Hodkinson 9561) formed a strongly supported clade (95% BS) and were divergent from the third specimen of E. citreus {K.R. McDonald 14437), which formed a polytomy with the sample of E. purpureus. This result suggests that E. citreus might not be monophyletic with the separation of E. purpureus. The specimens K.R. McDonald 15855 mvdK.R. McDonald 14437 had been annotated as belonging to ‘ small spikelet’ and ‘ long spikelet’ forms respectively, which could indicate a morphological difference between them. The ITS data therefore argue for further taxonomic revision of E. citreus across its range. Although a sister relationship was recovered between the E. citreus - E. purpureus clade and E. tylophorus, it was poorly supported (5 8% BS). Since E. tripsacoides is the only non-Australian species for which ITS sequences are available, it is notyet possible to draw any conclusions about the monophyly or otherwise of the Australian species. The ITS sequence generated for F. tylophorus differs in 18-21 substitutions plus several indels (95% similarity) from the sequences of E. citreus, which is considerably greater than is typically observed between closely related grass species (only 3 or 4 substitutions between E. purpureus and the various E. citreus sequences). The number of substitutions between F. tylophorus andF. citreus is comparable M.D. Barrett & T. Handasyde, Elionurus tylophorus (Poaceae), a new species from WA 187 to the difference between E. tylophorus and a Mexican accession of E. tripsacoides (DQ005047, Skendzic etal. 2007; 20 substitutions plus several indels, 94% similarity). The Australian citreus - E. purpureus clade and E. tylophorus therefore seem to be relatively distantly related to each other. Greater sampling of ITS and other loci within Elionurus will be required to determine whether the two Australian lineages arose following independent dispersals into Australia or represent a single more ancient lineage that speciated within Australia. The apparent restricted distribution of E. tylophorus is puzzling, given that its habitat is unassuming, appears representative of a widespread soil type and plant community, and is not obviously linked to any localised geological or hydrological feature (Figure 2). Many grass species in the Australian Monsoon Tropics have wide distributions, often extending to south-west Asia or beyond; however, occasional species with apparently restricted distributions do exist (e.g. Clausospicula extensa, a restricted monotypic genus in the Top End, Lazarides et al. 1991). It is possible that E. tylophorus has simply been overlooked amongst the array of other co-occurring grasses and is in fact more widespread than current data suggests. Taxonomy Elionurus tylophorus M.D.Barrett & T.Handasyde, sp. nov. Type'. King Leopold Ranges, Western Australia [precise locality withheld for conservation reasons], 13 April 2015, T Sonneman TH 8073 {holo\ PERTH 08662908; iso\ BRI, K, PERTH 09033955 (smutted specimen)). Erect annual grass 45-60 cm tall; culms <4 mm wide, unbranched, green to reddish; nodes 4-8, glabrous, internodes purplish, ±terete near base, broadly convexo-concave (±D-shaped) above, mostly glabrous but just below inflorescence-subtending node with appressed hairs along the angles, hairs becoming longer (up to 4.5 mm long) just below the node. Ligule a dense fringe of hairs 0.7-2.5 mm long. Leaves cauline; leaf sheath 26-52 mm long, 9-23-nerved, glabrous or sparsely hispid near apex, leaf blades linear, flat to canaliculate or involute, 65-175 mm long, 0.7-2.5 mm wide, 9-11-nerved, irounded on the abaxial surface but with raised nerves that are sometimes ±acute, adaxial surface glabrous to sparsely hispid near the base, abaxial surface glabrous except for the submargins with sparse tubercles often terminated by a stiff hair up to 1.5 mm long. Inflorescence a spatheolate raceme but appearance spiciform, racemes in upper 1-3 nodes. Spatheoles 45-69 mm long, mostly tightly sheathing and obscure, with a small, erect blade enclosing only the base or sometimes to apex of the lowest spikelet at maturity. Racemes 1-7 per node, 35-75 mm long, 3-8 mm wide, of 4-12 spikelet- pairs; rachis fragmenting at maturity, rachillas free from the internode, dissimilar to pedicel, shorter than or longer than pedicels, 7.5-16.0 mm long (tip to tip), 0.9-1.2 mm thick at widest point, ±parallel- sided, hollow, oblique at joints leaving sharply acute apices, somewhat flattened in section, mostly glabrous to very sparsely minutely to shortly hairy, but abaxial edge densely shortly hairy in a broad band with hairs ascending, 0.7-1.0 mm long, with longer stiff white hairs at apex, the longest hairs 3.0^.5 mm long; apex oblique, not winged or bilobed; disarticulation scar flattened, rim thickened, glabrous. Spikelets paired, one sessile and fertile, the other pedicellate and sterile, dimorphic within the pair, not gradational (similar length along the raceme), appressed and closely overlapping in two opposed staggered rows along a jointed disarticulating axis; diaspore composed of a pair of spikelets remaining attached to the rachilla. Sessile spikelet containing 2 florets, the lower sterile, the upper bisexual, apparently always chasmogamous, 9.0-13.2 mm long (including callus and proximal beak), 1.4-2.0 mm wide, dorsiventrally compressed; proximal beak c. 1.4-1.8 mm long, glabrous or hairy along margins and gradually merging with callus; callus elliptic, c. 1.0-1.5 mm long, gradually 188 Nuytsia Vol. 30 (2019) merging with beak, apex moderately acute to obtuse in face view, sharply acute in side view, glabrous along broad midline, pubescent with white hairs to 0.8-1.2 mm long on lateral faces, green turning straw sometimes with purplish margins or pale purplish all over; glumes dissimilar; lower glume of sessile spikelet 7-12 mm long (excluding callus), 1.4-2.0 mm wide, oblong to ±elliptic, broadest near the middle, apex bifid with 2 long beak-like teeth each 2.0^.5 mm long (much shorter than glume body), body thickly indurated, prominently 6-8-veined with thickened raised veins, strongly 2-keeled (keels thickened, sub-marginal and highly divergent, with nerves between them), body with sparse ±appressed to pubescent white hairs 0.05-0.25 mm long in lines in the narrow grooves between nerves, the grooves narrower than the nerves, a narrow, pale reddish brown oil streak present just inside each of the keels in the basal half (extending onto the inner face); keel not winged, with a row of strong tuberculate projections 0.15-0.3 mm tall in the lower l/2^/5 of the body, projections rounded to acute containing a single oblique apical hair up to 0.5 mm long, gradually merging into a dense fringe of stiff white hairs on keel margin (the longest hairs 0.5-0.8 mm long); glume margins very narrow, ±membranous, entire, slightly in-rolled beyond the keels and difficult to observe; upper glume of sessile spikelet 7-8 mm long, 1.7-2.0 mm wide, narrowly 1-keeled (carinate), 3-veined, shortly acuminate, chartaceous to coriaceous in the inner part between the nerves, membranous to chartaceous marginally outside the lateral nerves, shortly hairy (0.05-0.4 mm, irregular orientation, ascending) in a band either side of the keel and including on the keel, glabrous on marginal areas, narrow membranous margins with sparse to moderately sparse cilia (0.2-0.4 mm, wavy) in upper half; lower lemma of sessile spikelet 4.2-4.7 mm long, c. 1.4 mm wide, lanceolate, acute, lacking nerves or obscurely 2-4-nerved near base, membranous, pale straw to pale brown near base, sub-hyaline above and on margins, glabrous or minutely ciliate on upper margins; upper lemma of sessile spikelet elliptic, 4.2^.6 mm long, c. 1.3 mm wide, lanceolate, acute to acuminate, membranous to chartaceous, pale straw to sub-hyaline, 0-nerved or obscurely to moderately 3-nerved, glabrous or with a few cilia c. 0.25 mm long in upper part; lower and upper paleas of sessile spikelet absent; anthers 3, c. 2.5- 3.0 mm long, purple; styles 2, exserted at anthesis, purple; caryopsis 3.6^.9 mm long, 1.0-1.1 mm wide, moderately to distinctly dorsiventrally compressed, hilum half as long as caryopsis; lodicules 2, c. 0.4 mm long, membranous, broadly rounded at apex, entire. Pedicellate spikelet 8.0-11.7 mm long (excluding pedicels), 2-8 mm wide, rounded to weakly dorsiventrally compressed, with two fiorets, the lower sterile and the upper functionally male (anther may be difficult to demonstrate in older specimens with dropped anthers) with 2 glumes and 2 lemmas, initially lanceolate but gaping at maturity; pedicels 4.1-6.2 mm long, 0.6-0.8 mm wide, glabrous on one face, the opposite ‘external’ face with a longitudinal curving arc of short hairs that terminates in a sub-apical tuft of hairs 2.5- 4.5 mm long; glumes dissimilar; lower glume of pedicellate spikelet asymmetric most noticeable in keel indumentum, 9.0-11.7 mm long, 0.8-1.2 mm wide, dorsiventrally compressed, lanceolate to narrowly lanceolate, apex long-acuminate into an awn (2.0-)4.0-6.0 mm long, coriaceous, green to pale purplish all over, obscurely to moderately strongly 7-veined (not or scarcely raised, visible in transmitted light), 2-keeled close to margins, oil-streak present on both sides, with the keel glabrous or with stiff white hairs c. 0.2 mm long (indumentum usually asymmetric, more prominent on one keel than the other), margins membranous, 0.2-0.4 mm wide and wider on one side than the other; upper glume of pedicellate spikelet rounded to laterally compressed, shorter, subequal to or slightly shorter or longer than lower glume, very narrowly lanceolate, long-attenuate into an awn c. 3-5 mm long, indurated in central zone, chartaceous to membranous on sides, 1 -keeled ±in centre, obscurely to moderately 3-5-nerved, scabrous along upper keel and margin, glabrous; lower lemmas of pedicellate spikelet 3.0-4.0 mm long, 0.8-0.9 mm wide, lanceolate, acute to muticous, chartaceous to membranous, not nerved, glabrous or with marginal cilia in upper part; upper lemma of pedicellate spikelet shorter than lower, 2.6-3.0 mm long, glabrous, otherwise similar to lower lemma; paleas absent; anthers present in upper fioret, 3, apparently dimorphic, 2.2-3.8 mm long and exserted at maturity (a few 0.4 mm long are probably merely immature), purple; ovary and styles rudimentary, c. 0.2 mm long, not developing further. (Figure 3) M.D. Barrett & T. Handasyde, Elionurus tylophorus (Poaceae), a new species from WA 189 Diagnostic features. Elionurus tylophorus differs from all known Elionurus species in having keels (that are submarginal) of the lower fertile glumes ornamented with projections, which are often terminated by minute solitary bristles. Other notable features include: annual habit, prominently thickened lower glume nerves in the fertile (sessile) floret, pedicellate spikelets male with well-developed lemmas and anthers, lower glume body ±oblong with parallel-sided lateral outline, and glumes of the pedicellate spikelet subequal in length and awned. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] same locality as type, 29 May 2017, K. Carnes TH 8193 (DNA, KNR, PERTH); same locality as type, 26 May 2004, T. Handasyde TH 2148 (BRI, CANB, DNA, KNR, PERTH); same locality as type, 13 April 2015, T Sonneman TH 8072 (PERTH). Phenology. Fertile collections have been made between April and May (the latter senescing). Distribution and habitat. The single known E. tylophorus population was growing on a rocky ridge/slope of colluvium and alluvium in a sparse low woodland of a mix of species including Eucalyptus tectifica, E. brevifolia, Corymbia cadophora and Erythrophleum chlorostachys over scattered Cochlospermum fraseri and Calytrix achaeta (Figure 2). Ground cover included sparse to moderately dQnsQHeteropogon contortus, Eriachne sulcata, Mnesithea rottboellioides and Cymbopogon bombycinus. Elionurus tylophorus is currently known from a single location in a geologically diverse section of the King Eeopold Ranges in the central Kimberley. F igure 2. Habitat of Elionurus tylophorus from type location. Arrow indicates E. tylophorus, drying straw-coloured at time of photo. 190 Nuytsia Vol. 30 (2019) Figure 3. Elioniiriis tylophorus {T. Sonneman TH8073). A - splayed spikelet pair, showing pedicellate spikelet, fertile sessile spikelet and rachis internode; B -abaxial surface of base of lower glume of sessile spikelet showing strongly raised nerves, short hairs in rows between nerves, and marginal projections; C - adaxial surface of base of lower glume of sessile spikelet showing projections, and upper glume showing keel and surface hairs; D - dissected spikelet pair: a - rachilla, b - male pedicellate spikelet with lemmas visible, c - lower glume (with callus cut away), d - upper glume, e - lower lemma, f - upper lemma, g - caryopsis, h (insert) - apex of rachilla, which lacks a wing and is not bilobed. All scale bars 1 mm. M.D. Barrett & T. Handasyde, Elionurus tylophorus (Poaceae), a new species from WA 191 Conservation status. To be listed as Priority Two under Conservation Codes for Western Australian Flora (M. Smith, pers. comm.). It is conserved in the King Leopold Ranges Conservation Park (previously Mt Hart Station). Etymology. The species epithet is Greek from tylo- (with knobs, lumps or projections), and -phorus (-bearing). Notes. Cleistogamous flowers have been reported in both of the other Australian species, E. citreus and E. purpureus (Thompson 2017), but were not detected in any of the three specimens of E. tylophorus. However, plants were not cultivated, and some of the available E. tylophorus material is over-mature. In the K. Carnes TH 8193 collection, the best racemes have styles that are exserted but have already exserted and dropped their anthers from all sessile florets, while the sterile florets all have exserted or dropped anthers. In this collection at least, it appears that cleistogamous flowers are lacking, the sessile flowers are protandrous (as in most grasses), while the styles of sessile spikelets are ±synchronous within a raceme and with anthers of the pedicellate spikelets. Racemes in Elionurus fragment from the apex downward; although the lowermost spikelet is the last to fall in E. tylophorus, it does not appear tardily dehiscent as reported for E. purpureus (and at least one E. citreus plant) by Thompson (2017). Much of the available material is in the stage of dehiscing, and even prior to the secondmost-basal spikelet dehiscing the basal joint is already weak, suggesting that there is no significant delay of dehiscence of the basal florets. The characteristic marginal projections on the lower fertile glumes of E. tylophorus are possibly developmentally homologous with tuberculate bases of hairs, as found on leaves in the same species, since the projection is frequently terminated by a single short hair. Alternatively, since the projections are tightly fused with the keel, they may be outgrowths of the keel itself, centred on hair-cells. Elionurus tylophorus differs from all known Elionurus species (Renvoise 1978) in having submarginal projections on the keels of the lower fertile glumes, which are often terminated by minute solitary bristles, compared to cilia or hair-tufts that are not swollen at the base in other species. Elionurus tylophorus is also unusual within the genus in its annual habit. Aside from E. purpureus discussed above, the only other annuals or short-lived perennials in the genus are the African and Indian species E. elegans, E. hensii and.fi’. royleanus (Renvoise 1978). Elionurus tylophorus differs further from both of the only other Australian Elionurus, E. citreus (F igure 4) and^". purpureus (Thompson 2017), in its annual habit (usually tufted perennial in^". citreus), prominent thickened lower glume nerves in the fertile (sessile) floret (lower glume smooth or with thin scarcely raised nerves in E. citreus), lemmas and anthers present in pedicellate spikelets (absent in E. citreus and E. purpureus), lower glume body ±oblong with parallel-sided lateral outline (ovate and rounded lateral outline in E. citreus and E. purpureus), and glumes of the sterile spikelet roughly subequal in length (upper glume usually 1/2-2/3 as long as the lower glume in E. citreus and E. purpureus). Further differences between all three Australian species are contrasted in Table 2, and in the keys. The leaves of E. citreus have a lemon scent when crushed, but only a weak scent from crushed inflorescences is reported for E. tylophorus {K. Carnes TH 8193). 192 Nuytsia Vol. 30 (2019) Figure 4. Elionurus citreus (G.N. BatianoJflQW 8, PERTH 01780565, QLD, Lizard Is.), which is a good match for the holotype of E. citreus [Andropogon citreus R.Br.; R. Brown 6176, 1802; images of holotype at K and isotypes at E, W seen], except in being less obviously perennial than the type. A - splayed spikelet pair, showing sterile pedicellate spikelet, sessile spikelet and rachilla with expanded apex; B - base of lower glume showing obscure nerves, and marginal cilia (rachilla and pedicel cut away from callus); D - dissected spikelet pair: a - rachilla showing slightly winged and bilobed apex, b - sterile pedicellate spikelet, c - lower glume (with callus cut away), d - upper glume, e - lower lemma, f- upper lemma. All scale bars 1 mm. M.D. Barrett & T. Handasyde, Elionurus tylophorus (Poaceae), a new species from WA 193 Elionurus tylophorus superficially resembles some slender species of Ischaemum L. or Sehima Forssk., but is immediately distinguished by lacking an awn on the fertile lemma (twisted and geniculate awn present in Ischaemum and Sehima). Two of the collections are smutted [K. Carnes TH 8193 (PERTH 08654972) and TH 8073 (PERTH 09033955)]. No smut has previously been reported onE/zowwrw^ from Australia (Shivas etal. 2014), but three species, Anthracocystis elionuri (Henn. & A.Evans) McTaggart & R. G. Shivas, Macalpinomyces elionuri-tripsacoidis Vanl15 mm long 3. Calyx inner surface stellate hairy (NE Northclitfe-Fitzgerald River NP). T. stelligera 3: Calyx inner surface glabrous (New Norcia). T. sp. GreenHill* 2: Leaves ovate, oblong to orbicular, < 15 mm long 4. Style 5-winged; petals absent; ovary with peltate scale-like stellate hairs over glandular hairs (Albany E-Esperance W) . T. pygmaea 4: Style Aliform; petals present; ovary with peltate scale-like stellate hairs or flat stellate hairs 5. Ovary with 4-5 locules; petals with dense marginal peltate scale-like stellate hairs or multiangulate stellate hairs; calyx > 8 mm long; peduncles < 6 mm long (Mount Holland). T. gardneri^ 5: Ovary with 3 locules; petals glabrous or occasional scale-like stellate hairs on abaxial surface; calyx < 8 mm long; peduncles > 7 mm long 6. Leaves usually > 7 mm long, lower leaf surface with flat scale-like stellate hairs, sometimes with occasional multiangulate stellate hairs; peduncles (10-)30-50 mm long with 3-7 flowers 7. Calyx outer surface with flat scale-like stellate hairs alone, inner surface glabrous; leaves with midvein only indented (New Norcia). T. sp. GreenHill* 7: Calyx outer surface with flat scale-like and scattered multiangulate stellate hairs, inner surface with scattered white stellate hairs; leaves with midvein and minor veins indented (W of Woodanilling). T. sp. Arthur River 6: Leaves < 7 mm long, lower leaf surface with peltate scale-like stellate hairs; peduncles 10-23(-30) mm long with 1-2(3) flowers (Ongerup-Esperance east). T. microphylla 1: Stipules present, leaf-like and conspicuous or minute and inconspicuous; flowering stems with only multiangulate stellate hairs C.F. Wilkins & K.A. Shepherd, A key to the species of Thomasia (Malvaceae) 197 Figure 2. Floral features of Thomasia. A - ^owqx showing the petaloid calyx and stellate hairs with long arms on the peduncle and epicalyx bracts; B - flat scale-like hair; C - multiangulate stellate hair; D - anther with poricidal dehiscence, glabrous scale-like petal, ovary with stellate hairs, and a glabrous style; E - anthers with early lateral dehiscence and ovary with flat, scale-like stellate hairs and a glabrous style; F - anthers with poricidal dehiscence, scale-like petals with dense stellate hairs, ovary with stellate hairs, and base of style prominently tomentose to 1/3 of length; G - papillose ovary with a glabrous style, transverse section showing 5 carpels. Scale bars = 2.5 mm (A); 0.25 mm (B); 1 mm (C); 0.5 mm (D); 0.5 mm (E); 1 mm (F); 0.5 mm (G). Vouchers; Thomasia sp. Vasse (C. Wilkins & K. Shepherd CW 581) {K.A. Shepherd & J.A. Wege KS 345) (A-C, E); T. sp. Hopetoun (K.R. Newbey 4896) {K.A. Shepherd & J.A. Wege KS 277) (D); T. paniculata Lindl. {P.M. Olde 678, NSW 270044) (F); T. rulingioides Steud. {KA. Shepherd & J.A. Wege KS 201) (G). Illustration by Lorraine Cobb. 198 Nuytsia Vol. 30 (2019) Figure 3. Informative floral characters in Thomasia. A - style > 3 times longer than ovary and well exserted from the calyx; B - ovary with pink stellate hairs with short arms to 0.15 mm long; C - scale-like petals (black arrow); D - calyx thickened along the central area with thin, undulating margins; E - poricidally dehiscent anthers; F - perigynous cup with staminodes (white arrow). Vouchers: T. off. foUosa (K.A. Shepherd & C.F Wilkins KS 1614) (A); T. pauciflora Lindl. {K.A. Shepherd & C.F. Wilkins 1643) (B); Thomasia sarotes Tnrcz. {K.A. Shepherd et al. KS 1462) (C); T. grandiflora Lindl. (in cultivation) (D); T. montana Steud. {D. Rasmussen 2013/003) (E, F). Images by K.A. Shepherd. C.F. Wilkins & K.A. Shepherd, A key to the species of Thomasia (Malvaceae) 199 8. Stipules conspicuous; anthers poricidally dehiscent or with late lateral splitting; style c. 2 times longer than ovary and not well exserted from the calyx 9. Leaves and stipules linear, with margins tightly revolute 10. Inflorescence with 2 flowers (rarely 3); calyx <4.5 mm long; ovules 2 per locule (Hopetoun-Ravensthorpe). T. sp. Hopetoun 10: Inflorescence with 3-6 flowers; calyx usually >4.5 mm long; ovules 6-8 per locule (Mukinbudin-Quaalup). T. sarotes 9: Leaves and stipules not linear, margins flat or scarcely recurved to recurved 11. Mature leaves lobed 12. Mature leaves divided almost to midvein, appearing ‘oak-like’ 13. Leaves coriaceous, lower surface with a close tomentum of short stellate hairs each with arms to 0.2 mm long (Denmark-Esperance). T. querdfolia 13: Leaves soft, lower surface with scattered to medium dense long stellate hairs each with arms to 0.8 mm long (Augusta-Denmark). T. sp. Vasse 12: Mature leaves lobed or trilobed, not divided almost to the midvein 14. Leaves discolorous, lower surface with a close tomentum of stellate hairs; calyx 13-20 mm long with a strongly dissected tube c. 20% of the total length (Albany). T. discolor 14: Leaves almost concolorous, lower surface with scattered to dense stellate hairs; calyx 6-10 mm long with a moderately dissected tube c. 40% of the total length 15. Leaves (20)30-83 mm long, soft, lower surface with moderately dense stellate hairs 16. Petals absent or minute c. 0.4 mm long; epicalyx bracts narrowly obovate or narrowly elliptic (Denmark-Bremer Bay). T. solanacea 16: Petals > 1 mm long; epicalyx bracts ovate (Stirling Range National Park).... T. brachystachys 15: Leaves 12-44 mm long, coriaceous, lower surface glabrous or with scattered stellate hairs 17. Leaf margins strongly multi-lobed with a ±trilobed base, lamina usually longer than wide; anther filaments >2.5 mm long (Cervantes to Esperance in coastal heath). T. triphylla 17: Eeaf margins distinctly trilobed, lamina as long as wide; anther filaments < 2 mm long (Unknown). T. triloba 11: Mature leaves entire to scarcely sinuate, not distinctly lobed 18. Stamens and staminodes united in a glabrous or hairy perigynous cup c. 2 mm long (often longer than the ovary) 19. Perigynous cup hairy (Perth-Bridgetown). T. macrocarpa 19: Perigynous cup glabrous 20. Petals with scattered to dense stellate hairs (Brookton-Tutanning NR). T. montana 20: Petals glabrous (York area). T. glabripetala 18: Stamens free or scarcely united at base, or with a glabrous staminal tube <0.8 mm long 21. Calyx thickened along the central area either side of the midvein and with thin, undulating margins; leaves usually coriaceous with very few scattered 200 Nuytsia Vol. 30 (2019) Stellate hairs (Geraldton-Augusta-Esperance). T. grandiflora 21: Calyx thin except for midvein; leaves thin and soft, indumentum various 22. Leaves usually discolorous with a tomentose indumentum of dense stellate hairs below (Dryandra-Stirling Range NP). T. dielsii 22: Leaves almost concolorous with an indumentum of scattered to moderately dense stellate hairs below 23. Leaves < 10 mm long; pedicels with long glandular hairs (Morawa-Bencubbin). T. tremandroides 23: Leaves > 10 mm long; pedicel with glandular hairs absent or inconspicuous 24. Ovary with ±sessile papillose glands sometimes mixed with stellate hairs, without stalked glandular hairs 25. Petioles 0.2-1.5 mm long; leaf margins strongly recurved; upper surface of epicalyx bracts glabrous or rarely with 1-2 hairs (S of Dongara-N of Cervantes). T. rulingioides 25: Petioles (1.6-)2-8 mm long; leaf margins scarcely recurved; upper surface of epicalyx bracts with hairs throughout 26. Calyx usually <6.5 mm long, ovules 2 per locule 27. Ovary with red papillose glands only; flowers (3-)5-6 per inflorescence (Albany area). T. purpurea 27: Ovary with red papillose glands and scattered to dense white stellate hairs at the apex; flowers (4)5-8(-12) per inflorescence (Albany). T. multiflora^ 26: Calyx usually >6.5 mm long; ovules 4-8 per locule 28. Calyx lobes broadly ovate, c. 6 mm wide, tube almost half of total length; ovary with papillose glands only (Three Springs-Israelite Bay) ...T. macrocalyx 28: Calyx lobes ovate to narrowly ovate, c. 3.5 mm wide, tube < 20 % of total length; ovary with papillose glands only or a mix of papillose glands and white stellate hairs (Lrankland River-W of Cape Arid NP; SA, Vic). T. petalocalyx 24: Ovary with stellate hairs, sometimes with white, red- or pink-tipped stalked glandular hairs, or with stalked glandular hairs only 29. Base of style prominently tomentose to 1/3 of length; leaf margin recurved but entire to scarcely crenulate 30. Ovary with short stellate hairs only 31. Epicalyx bracts ovate, (1.7-)2-3 mm wide; flowers 1-2(3) per inflorescence; ovules (2-)4-7 per locule ; erect spreading shrub (Gracetown-Albany-Stirling Range National Park). T. rhyuchocarpa 31: Epicalyx bracts narrowly ovate, 0.8-1 mm wide; flowers (2)3(4) per inflorescence; ovules 2 per locule; low mounded shrub (Perth-Augusta-E of Albany). T. paudflora 30: Ovary with short stellate hairs and conspicuous glandular hairs 0.5-0.6 mm long C.F. Wilkins & K.A. Shepherd, A key to the species of Thomasia (Malvaceae) 201 32. Inflorescence a monochasium of < 3-7(-9) flowers; calyx > 7 mm long; leaves usually narrowly ovate (L;W c. 6:1) (Perth-Albany).T. paniculata 32: Inflorescence a dichasium or monochasium of 7-20 flowers; calyx < 7 mm long; leaves usually ovate (L;W c. 2.5:1) (N of Collie-W of Pemberton-E to the Porongurups).T. sp. Big Brook 29: Base of style completely glabrous or with a few stellate hairs; leaf margin recurved and somewhat crisped 33. Leaves ovate, 8-27 mm wide 34. Leaf upper surface prominently rugose; epicalyx bracts 2.1-3.3 mm wide; petals absent (Wongan Hills-E of Ravensthorpe).T. rugosa 34: Leaf upper surface not prominently rugose; epicalyx bracts 0.8-1.2 mm wide; petals stellate hairy (Carnamah to Bendering).T. tenuivestita 33: Leaves narrowly ovate, elliptic or oblong, 1.2-8 mm wide 35. Petals with stellate hairs on the margin; anthers <1.5 mm long; ovary hairs 0.7-1.25 mm long; peduncle stellate hairs with arms to 1.3 mm long (Cervantes-Esperance, in coastal heath). T. cognata 35: Petals glabrous; anthers > 1.7 mm long; ovary hairs 0.15-0.3 mm long; peduncle stellate hairs with arms 0.2-0.4 mm long 36. Leaves with veins scarcely impressed; inflorescence < 60 mm long; epicalyx bracts c. 1.5 mm wide; flowers 5-merous; anthers 1.8-1.9 mm long; ovules 2 per locule (Kojonup-Israelite Bay). T. angustifolia 36: Leaves with veins strongly impressed; inflorescence > 60 mm long; epicalyx bracts 2.1-4 mm wide; flowers 5- or 6-merous; anthers c. 3 mm long; ovules 6-8 per locule (Three Springs). T. formosa** 8: Stipules minute and inconspicuous; anthers showing early lateral dehiscence; style > 3 times longer than ovary and well exserted from the calyx* 37. Leaf lower surface and calyx outer surface with a dense indumentum of small stellate hairs, each with c. 12 arms (c. 0.2 mm long) arms, beneath moderately dense large stellate hairs (c. 0.5 mm long); moderately dense red-tipped glandular hairs on new growth and outer calyx of flowers in bud (Lesueur NP). T. sp. Lesueur 37: Leaf lower surface and base of calyx outer surface with an indumentum of very few small stellate hairs, each with c. 6 arms (0.2 mm long), beneath moderately dense large stellate hairs (0.5 mm long); scattered red-tipped glandular hairs on new growth and outer calyx of flowers in bud (Eneabba-Esperance E). T. foliosa ^Presumed extinct (see Smith & Jones 2019). ^Thomasia multiflora is currently only known from two collections (Z. Diels 5528, K. Newbey 4869). This species is poorly dilferentiated from T. purpurea and may not be supported as distinct. ''Possible hybrid between Lysiosepalum rugosum x T. macrocalyx (see Wilkins & Chappill 2001: 593). “Further work is required to resolve this complex. Thomasia foliosa and T. sp. Lesueur (M. Hislop 4217) usually have dark green leaves with an indumentum of scattered stellate hairs on the upper leaf surface; however, a variant currently determined as T. off. foliosa {K.A. Shepherd & C.F. Wilkins KS 1614, K.A. Shepherd & C.F. Wilkins KS 1617) with pale green leaves and an indumentum of dense stellate hairs on the upper leaf surface (Figure 3A) was collected east of Bindoon, co-occurring with typical T. foliosa {K.A. Shepherd & C.F. Wilkins KS 1616, K.A. Shepherd & C.F. Wilkins KS 1616b). Furthermore, another unusual specimen (C W. Parker 1628.08) with affinity to the closely allied T. sp. Lesueur (M. Hislop 4217) was collected east of Hopetoun, which is more than 800 kms to the south of known populations. 202 Nuytsia Vol. 30 (2019) Acknowledgements We sincerely thank Wendy Thompson for her helpful review, John Huisman for editorial assistance, and our late friend Lorraine Cobb for her wonderful illustrations. CFW was partially funded by a Science Project Support Grant from Biodiversity and Conservation Science (DBCA). The Australian Biological Resources Study is also gratefully acknowledged for financing early aspects of this research including the illustrations. References AVH(2019). The Australasian Virtual Herbarium. Council of Heads of Australasian Herbaria, http://avh.chah.org.au [accessed 19 March 2019], Bentham, G. (1863). FloraAustraliensis. Vol. 1. (Reeve and Co.: London.) Shepherd, K.A. & Wilkins, C.F. (2018). Typification of Lasiopetalum and an interim key to the Western Australian species of the genus (Malvaceae: Byttnerioideae). Nuytsia 29: 181-192. Smith,M.G.& Jones, A. (2018). awr/Pnon'/v C/ora/At 05 2075. Department ofBiodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 19 March 2019]. Western Australian Herbarium (1998-). FloraBase—the Western Australian Flora. Department of Biodiversity, Conservation and Attractions, https://florabase.dpaw.wa.gov.au [accessed 7 February 2018], Wilkins, C.F. & Chappill, J.A. (2001). A taxonomic revision of the Western Australian genus Lysiosepalum (Malvaceae: Lasiopetaleae). Nuytsia 13 (3): 571-594. Carolyn F. Wilkins^’^ and Kelly A. Shepherd^’^’^ 'Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 ^School of Biological Sciences, Faculty of Science, The University of Western Australia, 35 Stirling Highway, Crawley, Western Australia 6009 ''Corresponding author, email: Kelly.Shepherd@dbca.wa.gov.au Nuytsia The journal of the Western Australian Herbarium 30:203-214 Published online 30 July 2019 Two new Western Australian species segregated from Banksia densa (Proteaceae) Kevin R. Thiele^ 'Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 ''School of Biological Sciences, University of Western Australia 35 Stirling Hwy, Crawley WA 6009 Email: kevin.thiele@uwa.edu.au Abstract Thiele, K.R. Two new Western Australian species segregated from Banksia densa (Proteaceae). Nuytsia30: 203-214 (2019). Banksia densa A.R.Mast ScK.R.ThiQle has until now been regarded as a widespread species in the Western Australian wheatbelt, comprising a widespread nominate variety and a geographically more restricted variety in the southern part of its range (var. parva (A.S.George) A.R.Mast & K.R.Thiele). An informally recognised morphological variant, which matches the type of the species, is geographically restricted to an area on the edge of the species range in the immediate vicinity of Corrigin. Morphological assessment shows that the Corrigin form differs significantly in leaf morphology from the remainder of the species and comprises the true B. densa. The widespread taxon previously referred to as B. densa var. densa, and the southern var. parva, are here raised to species rank as B. zygocephala K.R.Thiele sp. nov. and B. parva (A.S.George) K.R.Thiele comb, et stat. nov. respectively. Introduction Banksia densa A.R.Mast & K.R.Thiele was described, as Dryandra conferta Benth., by Bentham (1870) based on a specimen collected by James Drummond from south-west Western Australia and distributed in his third collection as n. 295. Drummond’s third collection comprised specimens collected between Toodyay and Albany, and in the vicinity of Bolgart (Erickson 1969; George 2009). All species ofDryandraRBx. were transferred to by Mast and Thiele (2007). The replacement name B. densa was used for D. conferta due to the preoccupation of the original epithet in Banksia by the eastern Australian R. cow/erfa A.S.George. George (1996) placed B. densa (as D. conferta) in Dryandra ser. Obvallatae Benth., which he recircumscribed to include five (later seven; see George 1999) species: B.fasciculata (A.S.George) A.R.Mast & K.R.Thiele, B. densa, B. columnaris (A. S.George) A.R.Mast& K.R.Thiele, B.platycarpa (A.S.George) A.R.Mast & K.R.Thiele, B. seneciifolia (R.Br.) A.R.Mast & K.R.Thiele, B. rufistylis (A.S.George) A.R.Mast & K.R.Thiele and B. insulanemorecincta (A.S.George) A.R.Mast & K.R.Thiele. All these have an unusual infiorescence morphology in Banksia, with the fiowers all © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 204 Nuytsia Vol. 30 (2019) ‘ downcurved’ (George, 1996) giving the inflorescences a distinctive zygomorphic, bilateral symmetry (inflorescences in most other species in the genus are radially symmetric). Six of the seven species of ser. Obvallatae were included in a molecular phylogeny of Banksia by Cardillo and Pratt (2013). Five of these (B. fasciculata, B. densa, B. platycarpa, B. rufistylis and B. insulanemorecincta) form a clade, albeit with weak support. Sister to this clade is B. pallida (A.S.George) A.R.Mast & K.R.Thiele, which George (1999) placed in D. ser. Armatae Benth. The sixth included species in ser. Obvallatae, B. columnaris, was resolved close to but separate from the clade comprising the other ser. Obvallatae species. Support values on nodes in the dryandra clade on the Cardillo and Pratt (2013) phylogeny are extremely low, substantially reducing confldence that the clades recovered there are meaningful. However, the fact that most of the included ser. Obvallatae species were recovered as a clade indicates that the series is probably monophyletic or largely so. Banksia pallida, the species placed sister to the ser. Obvallatae species by Cardillo and Pratt (2013), also has a zygomorphic, bilaterally symmetric inflorescence with down-curved flowers, indicating that it may be misplaced in George’s ser. Armatae (all other species in that series have radially symmetric inflorescences) and that this feature may be a synapomorphy for the clade. Only six other species in the dryandra clade (the species in George’s ser. Concinnae Benth. and Plumosae A.S.George) have zygomorphic inflorescences, but in these cases the styles curve upwards whereas in ser. Obvallatae and B. pallida they curve downwards. George (1996) segregated plants from the southern part of the range of D. conferta, having smaller flowers and velvety rather than villous involucral bracts, as D. conferta var. parvaA.^.GQOxgQ. Even with var. parva excluded, he commented (George 1996: 387) that var. conferta (i.e. B. densa var. densa) was a variable taxon, noting that ‘typical var. conferta has linear leaf teeth and is of spreading habit’ while ‘the more common form has broad teeth and is columnar’. Despite this comment, he circumscribed var. conferta to include both these forms. Cavanagh and Pieroni (2006) recognised George’s ‘typical’ morphotype (that is, the form with linear leaf teeth that includes the type of D. conferta) as a ‘Corrigin form’, distinct from the widespread form which they called D. conferta var. conferta. They noted that the ‘Corrigin form’ is restricted to a small area of roadside between Corrigin and Quairading and has bluish leaves in the held. Mast and Thiele (2007), when transferring all species of Dryandra to Banksia, provided the new combinations Banksia densa var. densa and B. densa var. parva, without an assessment of the status of these two taxa or of the variability described by George (1996) and Cavanagh and Pieroni (2006). The present study was instigated when Margaret Pieroni brought to my attention the vulnerable state of the only population known to her of the ‘Corrigin form’. Weed invasion at the site and roadside clearing had reduced this population to a small number of individuals, raising the possibility, if the ‘Corrigin form’ and the more widespread morphotype discussed by George (1996) are distinct taxa, that urgent conservation assessments and action may be needed for the former. An initial assessment of all specimens of B. densa held at PERTH showed that the two morphotypes included under var. densa are in fact readily separable on the basis of leaf morphology. Given this, and to trigger a desktop assessment of conservation status of the ‘Corrigin form’, the phrase name Banksia densa var. Wheatbelt (M. Pieroni s.n. PERTH 04083407) was raised at PERTH for the widespread morphotype. With this change, B. densa var. densa includes only specimens of the ‘Corrigin form’; this was given a conservation ranking of Priority Two on the Western Australian K.R. Thiele, Two new species segregated from Banksia densa (Proteaceae) 205 Department of Biodiversity, Conservation and Attractions ’ Threatened and Priority Flora /Ar (Smith & Jones 2018). Using an informal varietal name for ‘var. Wheatbelf was preferred at that time, pending assessment of appropriate rank. This paper provides a morphological assessment and taxonomic reappraisal of all three taxa currently recognised within Banksia densa, following a more complete study of specimens held at PERTH and detailed observations in the field. Materials and Methods All specimens at PERTH were assessed, both to determine the distinction between B. densa var. densa and the widespread B. densa var. Wheatbelt, and to assess the most appropriate status for these and for B. densa var. parva. Measurements were made of style length for all specimens with mature flowers, as this feature was used by George (1996) to segregate var. parva from var. densa sens. lat. (style length 16-26 mm in var. parva cf 25-30 mm in var. densa). Given that var. Wheatbelt and var. parva are latitudinally separated (with the latter at the southern end of the range of the former), style lengths were plotted against latitude and a regression analysis performed in Genstat v. 19 (VSN International 2017) to assess whether the variation in style length between the two taxa is clinal. All putative taxa were visited and compared in the field during spring 2017. Type specimens were viewed using JSTOR Global Plants (https://plants.jstor.org/). Maps are based on all specimens held at PERTH and are drawn using IBRA v. 7 (Department of the Environment 2013) bioregion and subregion boundaries. Results Both in the herbarium and in the field, B. densa var. densa and B. densa var. Wheatbelt are clearly separable on the basis of leaf morphology alone (Figure 1). As first noted by George (1996), leaf lobes of B. densa var. densa are ±linear (parallel-sided) to narrowly triangular, while those of var. Wheatbelt are broadly triangular. While these differences in shape do not lend themselves to simple measurements, in practice all available specimens can be adequately and unambiguously assigned to the two taxa. Banksia densa var. Wheatbelt and var. parva are distributed more or less along a north-south axis in the Western Australian wheatbelt (Figure 2), with both taxa narrowly sympatric in the southern part of the range of the former, approximately between Nyabing and Ongerup. A plot of style length against latitude (Figure 2) shows that (1) the two taxa have non-overlapping style lengths, and (2) style length invar. Wheatbelt is not correlated with latitude (R^=0.065, p=0.079 n.s.). Neither of these would be expected if var. parva and var. Wheatbelt comprised a single, clinal taxon. The clear morphological separation between B. densa var. densa, var. Wheatbelt and var. parva, lack of intermediates, lack of any indication of a north-south cline in style length in var. Wheatbelt, and partial sympatry of var. Wheatbelt and var. parva, all provide morphological evidence that the three taxa comprise independently evolving lineages. Given the lack of intermediates and clear morphological separation, I regard that species rank is appropriate for all three; accordingly, they are treated below as B. densa, B. zygocephala K.R.Thiele sp. nov. and B. parva (A.S.George) K.R.Thiele comb, et stat. nov., respectively. 206 Nuytsia Vol. 30 (2019) Figure 1. Leaves of the type form of Banksia densa (A); the widespread var. Wheatbelt (B); and B. densa var. parva (C). Top row, from left: A. Mast 558; K.R. Newbey 2488; A. Mast 505; A.S. George 16754; G.S. Durell 55; C.A. Gardner 13621; K.Alcock5Al,A. Cochrane 552', A.S. George 14361;7?. Davis 12819. Bottom row, from left: 7?. J. Cranfield d,066; G.J. Keighery & N. Gibson 5801; R. Meissner 5654; G.J. Keighery & N. Gibson 3470; A.S. George 16704; S. Barrett 1461; S. Barrett 842; S. Barrett 1249; A.S. George 16691; ^.5'. George 16662. Key to species previously referred to Banksia densa 1. Styles 14-23 mm long. B. parva 1: Styles 26-34 mm long 2. Leaf lobes broadly triangular, the sides ±straight; plants ±columnar and erect to 3 m. B. zygocephala 2: Leaf lobes narrowly triangular to ±linear, the sides nearly parallel before curving to the apex; plants spreading, to 1 m. B. densa K.R. Thiele, Two new species segregated from Banksia densa (Proteaceae) 207 40 35 30 ^ 25 E M 20 _a) _a) >■ CO 15 10 5 0 29 30 31 32 33 34 35 Latiitude (degrees South) Figure 2. Variation in style length (mm) with latitude for B. densa var. Wheatbelt (circles) and B. densa var. parva (diamonds). Open circles are specimens that were not determined to variety but are within the range of var. Wheatbelt. The regressions (dotted lines) are not significantly correlated with latitude (var. Wheatbelt R^=0.065, p=0.079 n.s.; var. parva R^=0.185, p=0.091 n.s.). Taxonomy •> % ° • • • A • • • • cfe A AA o z ■■■ 0 ^ Banksia densa A.R.Mast & K.R.Thiele, Austral. Syst. Bot. 20(1): 65 (2007). Dryandra conferta Benth., FI. Austral. 5: 578 (1870). Type citation: ‘W. Australia, Drummond, 3rd. coll. n. 295.’ (lecto, fide George, Nuytsia 10(3): 387 (1996): K 796238 image!; isolecto: BM fide George, n.v., CGE fide George, w.v., G389422 image!, K 796239 image!, K 796237 image!, MEL 602358!, P 750880 image!, PERTH 1590650 [fragments]!). Non-lignotuberous, rounded to spreading or sprawling shrubs to c. 1 m high. Young stems densely tomentose with short, curled, whitish hairs overtopped by sparse longer, straighter, reddish-brown hairs. Leaves pinnatipartite, mostly densely crowded on short-shoots lateral to the main stems, often dull bluish-green, 90-160 mm long, 8-16 mm wide; lamina linear, sparsely to moderately tomentose above with minute, tightly curled, whitish hairs persisting or deciduous above when mature, closely and densely tomentose beneath with pale fawn to whitish, tightly curled hairs obscuring the surface; lobes 6-20 each side, at c. 70°-90° to the axis, 3-10 mm long, linear to narrowly triangular, acute, shortly pungent, the margins narrowly revolute. Inflorescences lateral on short-shoots; involucral bracts very narrowly ovate to linear, acute, the innermost to 17 mm long, sparsely to moderately pale to dark brown-villous with spreading hairs that are denser, darker and less spreading towards the apex. Flowers 56-70 per head, creamy yellow; perianth 18-23 mm long, woolly above base, spreading brown-villous above; limb 3^ mm long, sparsely brown-villous often with an apical tuft of denser hairs. Pistil 27-30 mm long, downcurved, glabrous except hirsute immediately above ovary; pollen 208 Nuytsia Vol. 30 (2019) presenter 1.5-2.5 mm long, narrowly cylindrical, not distinctly thickened, scarcely contracted to the pistil, faintly ribbed. Follicles obliquely obovate, 9-10 mm long, sparsely hirsute. Diagnostic features. May be distinguished from the other taxa previously included in B. densa by the leaf lobes that are narrowly triangular to linear, often bluish-green leaves, and spreading, non- columnar habit. Specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 12 Aug. 2018, R. Davis 12819 (PERTH); 19 Oct. 1961, C.A. Gardner 13621 (PERTH); 6 Sep. 1976, A.S. George 14361 (PERTH); 2 Aug. 1986, A.5. George 16754 (PERTH); 22 Oct. 2004, A. Mast 558 (PERTH); 16 Oct. 1966, K.R. Newbey 2488 (PERTH). Phenology. Flowers between April and October with a probable peak in August. Distribution and habitat. Restricted to an area within a radius of approx. 30 km from Corrigin in the Western Australian wheatbelt (Figure 3). Grows in white to yellow sand over laterite, in open, proteaceous mallee-heath withEucalyptus macrocarpa,Allocasuarina huegeliana, Isopogon divergens, Banksia sphaerocarpa, B. vestita and B. cirsioides. Conservation status. Currently listed as Priority Two under the Conservation Codes for Western Australia Flora (Smith & Jones 2018), under the name Banksia densa var. densa. Historical collections have been made at approx, six localities; however, only two are known to be extant. One 1993 collection {G.S. Durell 55) from private land has not yet been revisited. Figure 3. Distribution of Banksia densa. K.R. Thiele, Two new species segregated from Banksia densa (Proteaceae) 209 James Drummond is not known to have collected near Corrigin. His collection comprised specimens collected between Toodyay and Albany, and in the vicinity of Bolgart (Erickson 1969; George 2009). It is possible that Banksia densa occurred further west than it does now, before extensive clearing of the wheatbelt. Notes. Banksia densa (formerly Dryandra confertd) is here recircumscribed to include only the ‘type’ or ‘Corrigin’ form as discussed by George (1996) and Cavanagh and Pieroni (2006), with linear to narrowly triangular leaf lobes and a low, sprawling habit. The foliage of living plants tends to be bluish due to the persistence of an indumentum; however, this is not universal, especially in larger populations. Banksia zygocephala K.R.Thiele, sp. nov. Type: Nyabing-Kukerin Road, 3.7 km N of Nyabing, Western Australia, 19 August 2018, K.R. Thiele 5524 {holo\ PERTH 09083502; iso\ CANB, MEE, K). Banksia densa var. Wheatbelt (M. Pieroni s.n. PERTH 04083407), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 15 March 2018]. Non-lignotuberous, erect, multi-stemmed, usually columnar to 3 m high. Young stems densely tomentose with short, curled, fawn to brown hairs overtopped by sparse longer, straighter, reddish- brown hairs. Leaves pinnatipartite, mostly densely crowded on short-shoots lateral to the main stems, dull green, 60-160 mm long, 15-20 mm wide; lamina linear, sparsely to moderately tomentose above when young with minute, tightly curled, whitish hairs, glabrous above when mature, closely and densely tomentose beneath with pale fawn to whitish, tightly curled hairs obscuring the surface; lobes 10-18 each side, at c. 60°-80° to the axis, 5-10 mm long, broadly triangular, acute, pungent, the margins narrowly revolute. Inflorescences lateral on short-shoots; involucral bracts very narrowly ovate to linear, sometimes narrowly spathulate at apex, acute, the innermost 15-27 mm long, sparsely to moderately pale- to brown-villous with spreading hairs sometimes restricted to the margins, the hairs denser, darker and less spreading towards the apex. Flowers 55-80 per head, creamy yellow; perianth 22-25 mm long, woolly above base, moderately to densely spreading fawn- to brown-villous above; limb 3.5^ mm long, sparsely brown-villous often with an apical tuft of denser hairs. Pistil 26-34 mm long, downcurved, glabrous except hirsute immediately above ovary; pollen presenter c. 2 mm long, narrowly cylindrical, not distinctly thickened, scarcely contracted to the pistil, faintly ribbed. Follicles obliquely obovate, 9-14 mm long, sparsely to moderately hirsute. (Figure 4.) Diagnostic features. May be distinguished from the other taxa previously included in B. densa by the leaf lobes that are broadly triangular, dull green leaves, usually columnar habit, and pistils 26- 34 mm long. Specimens examined. WESTERN AUSTRAEIA: 4 km E of Cadoux, 5 Apr. 1987, K. Alcock 548 (PERTH); 26 km due SW of Bodallin, 16Sep. 1982,i?.J. Craw>/^/2315 (PERTH); Martin] inni Nature Reserve, 18 Sep. \99\,R.J. Cranfield & P.J. Spencer 8066 (PERTH); [10 km W of Bodallin on Great Eastern Hwy], 18 Sep. 1962, A.S. George 4162 (PERTH); c. 22 km SE of Nyabing along Rabbit Proof Fence, 26 Jun. 1976, A.S. George 14289 (PERTH); Paterson Rd, 0.7 km W of Junction with Balls Rd, NE of Woodanilling, 26 July 1986, A.S. George 16633 (PERTH); 26.3 km N of Nyabing on Kukerin Rd, 30 July 1986, A.S. George 16704 (PERTH); 4 km E of Cadoux on Johnson Rd, 3 Aug. 1986, A.S. George 16762 (PERTH); Kojonup, E side of Merilup Rd, 22 Dec. 1998, M.S Graham 1038 (PERTH); Along E boundary of Harrismith Reserve, 18 Oct. 2001, K. Kershaw 2452 (PERTH); 3.9 km 210 Nuytsia Vol. 30 (2019) Figure 4. Banksia zygocephala: top—habit; bottom left—inflorescence, front view; bottom right—inflorescence longitudinal section, natural orientation. All photos: R. Davis (plants from the type population, Nyabing-Kukerin Rd 3.7 km N of Nyabing, 19 Aug. 2018). N of Wickepin-Harrismith Rd, c. 4 km NE of Harrismith, 28 Aug. 1997, G.J. Keighery & N. Gibson 5801 (PERTH); 14.5 km SE of Dumbleyung, 6 Nov. 1992, S.A. McNee DY 904 (PERTH); Mount Holland, 14 Oct. 1963, K.R. Newbey 1113 (PERTH); Gibb Rock, 1972, L. Sedgwick s.n. (PERTH), Miling, Aug. 1912, Seymour s.n. (PERTH). K.R. Thiele, Two new species segregated from Banksia densa (Proteaceae) 211 Phenology. Flowers between April and October with a distinct peak in August. Distribution and habitat. Occurs in two disjunct areas, a northern one between Gunyidi and Lake Cronin and a southern one from around Toolibin to near Ongerup and as far west as Dardadine (Figure 5). An outlying collection from the southern boundary of Stirling Range National Park {S. Barrett 843) is well within the range of B. parva, but clearly has a style length within the range of B. zygocephala and is typical in all respects. In the northern area of distribution, grows in white to yellow clayey sand over laterite, in heathy woodlands with Eucalyptus oldfieldii, E. burracoppinensis, E. macrocarpa. Acacia yorkrakinensis, Banksia armata, B. purdieana, Beaufortia interstans, Melaleuca conothamnoides and M cordata. In the southern area, grows in white, yellow or brown sands and loams in proteaceous and myrtaceous heath, mallee-heath and open wandoo woodlands with Eucalyptuspluricaulis, Melaleuca platycalyx, Allocasuarina thymoides, Hakea prostrata, H. cygnorum, Grevillea cagiana, Banksia sphaerocarpa, B. baueri, B. cynaroides, B. nivea, B. drummondii and Isopogon buxifolius. Conservation status. Widespread and not considered to be at risk. Etymology. From the Greek zygos (a yoke, pair) and kephale (a head), in reference to the zygomorphic inflorescences with downcurved flowers. Notes. With the removal of B. zygocephala from B. densa sens, lat., a distribution that appeared more or less continuous is now shown to be discontinuous, with a gap of around 125 km between the northern and southern areas of the range of the former. Suitable habitats occur within the gap, indicating that B. zygocephala as currently circumscribed may comprise two taxa. Plants from the Figure 5. Distribution of Banksia zygocephala. 212 Nuytsia Vol. 30 (2019) northern area have larger, coarser leaves with more widely spaced and larger leaf lobes than plants in the southern area; however, the differences are subtle and individual specimens are difficult to place. It is currently premature to recognise two taxa, but further work may indicate that this is the case. The type specimen is from the southern area of distribution. Banksia parva (A.S.George) K.R.Thiele, comb, et stat. nov. Dryandra conferta var. parva A.S.George, Nuytsia 10(3): 388 (1996). Banksia densa var. parva (A.S.George) A.R.Mast & K.R.Thiele, Austral. Syst. Bot. 20(1): 67 (2007); Type: South Fence Rd, 7 kmNW of Albany-Lake Grace Rd, SE ofNyabing, Western Australia, 30 July 1986, A.S. George 16694 {holo\ PERTH 03462595!; iso\ CANB 4853261, PERTH 034626091). Non-lignotuberous, erect, multi-stemmed, usually columnar shrubs to 1.5 m high. Young stems densely tomentose with short, curled, fawn to brown hairs overtopped by sparse longer, straighter, reddish-brown hairs. Leaves pinnatipartite, mostly densely crowded on short-shoots lateral to the main stems, dull green, 50-150 mm long, 8-18 mm wide; lamina linear, sparsely to moderately tomentose above when young with minute, tightly curled, whitish hairs, glabrous above when mature, closely and densely tomentose beneath with pale fawn to whitish, tightly curled hairs obscuring the surface; lobes 8-20 each side, at c. 60°-80° to the axis, 3-9 mm long, triangular, acute, pungent, the margins narrowly revolute. Inflorescences lateral on short-shoots; involucral bracts very narrowly ovate to linear, sometimes narrowly spathulate at apex, acute, the innermost to 18 mm long, sparsely to moderately pale- to brown-villous with spreading hairs sometimes restricted to the margins, the hairs denser, darker and less spreading towards the apex. Flowers 40-65 per head, creamy yellow; perianth 15-19 mm long, woolly above base, moderately to sparsely to densely spreading fawn- to brown- villous above; limb 3-4 mm long, sparsely to moderately pale brown-villous often with an apical tuft of denser hairs. Pistil 14-23 mm long, downcurved, glabrous except hirsute immediately above ovary; pollen presenter c. 2 mm long, narrowly cylindrical, not distinctly thickened, scarcely contracted to the pistil, faintly ribbed. Follicles obliquely obovate, 6-10 mm long, sparsely to moderately hirsute. Diagnostic features. May be distinguished from the other taxa previously included in B. densa by the leaf lobes that are broadly triangular, dull green leaves, usually columnar habit, and pistils 14- 23 mm long. Specimens examined. WESTERN AUSTRAEIA: [precise localities withheld for conservation reasons] North Woogenilup, 3 May 2000, S. Barrett 843 (PERTH); Nyabing, 26 June 1976, A.S. George 14290 (PERTH); Stirling Range, 27 July 1986, A.S. George 16662 (PERTH); Take Grace, 30 July 1986, A. S. George 16691 (PERTH); Porongurup Range, 18 Aug. 1971, ilT. 3414 (PERTH); Boxwood Hill-Ongerup, 8 July 1987, M. Pieroni 33; North Woogenilup, 11 July 1998, M. Pieroni 98/3 (PERTH). Phenology. Flowering specimens have been collected between May and July, with probable peak flowering in the latter month. Banksia parva flowers significantly earlier than either B. densa or B. zygocephala, despite its more southerly range. During fieldwork in mid-August 201 8 , R. zygocephala was in full flower while B. parva had consistently finished flowering. Distribution and habitat. Occurs from Badgebup to the Porongurup Range and east to near Ongerup (Figure 6). Grows in sandy and gravelly loams and sands over laterite, in mallee-heath and shrublands with Eucalyptuspleurocarpa, E. incrassata, Allocasuarina huegeliana, Hakea varia, H. lissocarpha, H. marginata, Banksia nervosa, B. arctotidis, B. gardneri, B. armata, B. sphaerocarpa, B. drummondii. K.R. Thiele, Two new species segregated from Banksia densa (Proteaceae) 213 Regelia inops, and Xanthorrhoea platyphylla. Conservation status. Currently listed as Priority Four under the Conservation Codes for Western Australian Flora (Smith & Jones 2018), under the name Banksia densa var. parva. Notes. Banksia parva is morphologically very close to B. zygocephala, differing mainly in its shorter styles and perianths. Leaves in B. parva also tend to be shorter than in B. zygocephala, but the ranges overlap and leaf length and lobe shape is not diagnostic. George (1996) discriminated the two varieties of B. densa partly on the indumentum of their involucral bracts, with B. parva (as Dryandra conferta war. parva) described as having velvety bracts and B. zygocephala (as D. conferta var. conferta) having villous bracts. While the indumentum on the bracts of B. parva is on average slightly less dense and more spreading than 'mB. zygocephala, the differences are slight, are not consistent, and are difficult to use for diagnosis. Likewise, George(1996) described the follicles ofvar. parva as ‘more oblique (almost transversely obovate)’ {l.c. p. 388). Comparison of a range of follicles of B. parva and B. zygocephala shows that follicle shape in both species is variable and is not diagnostic. George (1996) used varietal rank for Dryandra conferta var. parva based on a ranking scheme used throughout his work at the time. Under this scheme, taxa were given species rank if ‘morphological characters and states are considered significant in the context of the genus’, subspecies rank was used when ‘diflference(s) are less significant and there is a geographical and/or ecological discontinuity’, and variety was used when ‘diflference(s) are less significant than at specific rank and there is no geographical or ecological separation’ (l.c. p. 314). Figure 6. Distribution of Banksia parva. 214 Nuytsia Vol. 30 (2019) Under this scheme, D. confertayar.parvawas recognised at varietal rank because it comprises southern populations of D. conferta that are not ecologically or geographically disjunct from var. conferta. I regard, by contrast, that the discontinuity in style lengths between B. zygocephala and B. parva, despite at least partial sympatry, differences in flowering times, and lack of any indication of clinal variation, indicates that the two taxa are reproductively isolated lineages, and hence are appropriately recognised at species rank. Summary of taxon circumscriptions Banksia densa A.R.Mast & K.R.Thiele in this paper has a narrower circumscription than B. densa A.R.Mast & K.R. Thiele var. densa as previously accepted at PERTH due to the removal of B. zygocephala. Banksia zygocephala K.R.Thiele in this paper is a new taxon and is removed from B. densa A.R.Mast & K.R.Thiele var. densa as previously accepted at PERTH. Banksia zygocephala K.R.Thiele in this paper has the same circumscription as B. densa A.R.Mast & K.R.Thiele var. Wheatbelt (M. Pieroni s.n. PERTH 04083407) as previously accepted at PERTH. Banksia parva (A.S.George) K.R.Thiele in this paper has the same circumscription as B. densa var. parva (A.S.George) A.R.Mast & K.R.Thiele as previously accepted at PERTH. Acknowledgements I thank Margaret Pieroni for bringing to my attention the taxonomic and conservation issues around Banksia densa, Rob Davis and Tim Hammer for assistance in the held, the Curator and staff of PERTH for access to the collections, and Kelly Shepherd and an anonymous reviewer for helpful comments on the ms. References Erickson, R. (1969). The Drummonds of Hawthornden. (Lamb Patterson: Perth.) Bentham, G. (1870). FloraAustraliensis. Vol. 5. (Reeve and Co.: London.) Cardillo, M. & Pratt, R. (2013). Evolution of a hotspot genus: geographic variation in speciation and extinction rates in Banksia (Proteaceae). BMC Evolutionary Biology 13(155): 1-11. Cavanagh, A.K. & Pieroni, M. (2006). The Dryandras. (Wildflower Society of Western Australia: Perth, Australia.) Department of the Environment (2013). Australia’s bioregions (IBRA). IBRA7, Commonwealth of Australia. http://www. environment.gov.au/land/nrs/science/ibra#ibra [accessed 5 January 2016], George, A.S. (1996). New taxa and a new infrageneric classification in Dryandra R.Br. (Proteaceae: Grevilleoideae). Nuytsia 10(3): 313-408. George, A.S. (1999). Dryandra. In: Wilson, A. {ed.) Flora of Australia. Vol. 17B. pp. 251-363. (Australian Biological Resources Study: Canberra.) George, A.S. (2009). The Australian Botanist’s Companion. (Four Gables Press: Kardinya, Western Australia.) Mast, A.R. & Thiele, K. (2007). The transfer of Dryandra R.Br. to Banksia L.f (Proteaceae). Australian Systematic Botany 20(1): 63-71. Smith, M.G. & Jones, A. (2018). Threatened and Priority Floralist December 2018. Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 6 February 2019], VSN International (2017). Genstat for Windows 19th Edition (VSN International, Hemel Hempstead, UK.) Nuytsia The journal of the Western Australian Herbarium 30:215-219 Published online 15 October 2019 Additions to Eremophila (Scrophulariaceae) Robert J. Chinnock State Herbarium of South Australia, Department of Environment, Water and Natural Resources, GPO Box 1047, Adelaide, South Australia 5001 Email: bob.chinnock@sa.gov.au Abstract Chinnock, R.J. Additions to Eremophila (Scrophulariaceae). Nuytsia 30: 215-219 (2019). One new species of Eremophila R.Br.,E. waitii Chinnock, is described and one subspecies of E. glabra (R.Br.) Ostenf, E.glabra subsp. verrucosa Chinnock, is raised to species level and recognised herein as E. viridissima Chinnock. Introduction Since the treatment of 215 species of Eremophila R.Br. in the Myoporaceae monograph by Chinnock in 2007, an additional 19 species have been added to the genus (See Brown & Buirchell 2007; Chinnock & Doley 2011; Edginton 2015; Buirchell & Brown 2016; Brown & Davis 2016; Brown etal. 2018). The addition of the two species here brings the current number of recognised species in Eremophila to 238. A recently discovered endangered species of Eremophila from the northern Wheat Belt of Western Australia is described and a subspecies of Eremophila glabra (R.Br.) Ostenf, described by Chinnock (2007), is raised here to species level. Taxonomy Eremophila waitii Chinnock, sp. nov. Type-. Mullewa, Western Australia [precise locality withheld for conservation reasons], 21 September 2018, RJ. Chinnock 10600 {holo'. PERTH 09083618; iso'. AD, CANB, MEE). Eremophila sp. Mullewa (R. Wait 7311), Western Australian Herbarium, mFloraBase, https ://florabase. dpaw.wa.gov.au/ [accessed 20 January 2019]. Erect, densely branched aromatic shrub 1-1.5(-2.5) m tall, bark pale grey, rugose. Branches distinctly ribbed, ribs extending down from raised leaf bases, obscurely tuberculate, resin extruded in dried specimens as small translucent bubbles, densely finely stellate-tomentose, glabrescent in older parts. Leaves very densely clustered along branches, ± alternate-sub-whorled or in whorls of 3, straight or slightly curved, linear-subterete, tapering towards tip, broadly acute to obtuse, (6-)10-18(-26) mm © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 216 Nuytsia Vol. 30 (2019) X 0.5-0.9(-l.l) mm, dendritic pubescent to dendritic-tomentose, glabrescent. Flowers 1 per axil, subsessile (pedicel <1 mm), densely clustered into strobiloid structures 30^0 mm long at branch tips, and flowers developing from base upwards or opening at various positions along strobilus at same time. Sepals 5, subequal, valvate, lanceolate, acute, 5-8 x 0.7-1.2 mm; outer surface densely clothed in large dendritic hairs partially or completely obscuring sepals except at tip, inner surface with long dendritic hairs in distal half, glandular pubescent below; green or dark purple. Corolla 9-15 mm long, outer surface and inside of lobes and upper inside of tube deep lilac, inside of tube below lower lip white with purple spots; outside surface and inner surface of lobes glabrous, tube with white woolly hairs especially in lower part; lobes obtuse. Stamens 4, enclosed, filaments glabrous. Ovary oblong, 4-locularwith 1 ovule per loculus, glabrous. Fruit dry, obconical, 3^.5 x 2.5-3.0 mm, exocarp papery, bifid at apex, rugulose, very pale brown; endocarp, brown, split into 4 at apex, rugulose, granular with numerous minute resinous spots. Seed oblong, c. 1.5 x 0.5 mm, pale cream. (Figure 1) Diagnostic features. Eremophila waitii is readily distinguished from other species in Eremophila sect. Australophilae Chinnock in having sub-whorledto 3-whorled, densely pubescent to densely tomentose, linear-subterete leaves that are clustered along the branches and flowers aggregated together into woolly strobili 30-40 mm long at branch tips. Figure 1. Eremophila waitii. A - type population; B - habit, note the much taller senescent plant in background (centre); C - inflorescence showing the well-deflned strobili and flowers emerging from various places; D - comparison between E. waitii (left) and the larger flowered E. nivea (right). Vouchers; R.J. Chinnock 10599 (C, D left (right not vouchered). Photographs by R.J. Chinnock. R. J. Chinnock, Additions to Eremophila (Scrophulariaceae) 217 Other specimens examined. WESTERN AETSTRAEIA. [localities withheld for conservation reasons] 14Sep. 2018,5. 5wz>cMBB332 (PERTH, image seen); 19Aug. 2018,5. Waitl?>\\ (PERTH, image seen). Phenology. Flowers appear from late winter to at least mid-spring with fruits maturing from early spring onwards. Distribution and habitat. Eremophila waitii is known only from one population consisting of about 55 plants (Figure 1 A) in a very small area around low sandstone rock outcrops. It was growing with Melaleuca dind Acacia species, Solanum sp., various chenopods and a mixed annual ground cover of composites and grasses in an open area in remnant low mallee woodland. It is possible that because many of the plants were very close together they may have resulted vegetatively from root-suckering, but this could not be confirmed. Most plants in the population were between 1-1.5 m tall and wide (Figure IB) with a rounded crown but a few senescent plants, usually with a few unbranched stems, were up to 2.5 m tall. It was noted that leaves on new vigorous basal branches of actively growing plants are longer (>18 mm) with a more sparse indumentum and more obviously resinous. Conservation status. Eremophila waitii, is known only from the one population and has recently been listed as a Priority One species under the Conservation Codes for Western Australian fiora under the name Eremophila sp. Mullewa (R. Wait 7311) (Western Australian Herbarium 1998-). Etymology. Named after Russell Wait, who over many years has trekked to various parts of Australia, but especially Western Australia, to study and collect Eremophila. I am delighted to name this species after him in recognition of the many new or rare species that he has discovered and introduced into cultivation and the significant contribution he has made to our understanding of this genus. Affinities. Eremophila waitii is most closely related to Eremophila subangustifolia A.P.Br. & Elorens (see Notes) and E. microtheca (F.Muell. ex Benth.) F. MuelL, sharing with these species the dendritic- pubescent or dendritic-tomentose leaves and branches. It is readily distinguished from these species by having a well-defined strobiloid infiorescences (Figure 3) at the branch tips rather than having scattered fiowers along the upper parts of the branches. Eremophila nivea Chinnock is also closely related to E. waitii sharing a similar dense white to grey-white indumentum on the branches and leaves. Eremophila nivea can have a ± loose cluster of flowers near the apex (Figure 4) but it is readily distinguished from E. waitii by the larger flowers 15-23 mm long {cf. 9-15 mm long), and the distinctly flattened, broader, leaves 1.5-3.5 mm wide {cf. 0.5-0.9(-l.l) mm wide). The four species E. waitii, E. subangustifolia, E. microtheca and E. nivea form a well-defined group within Eremophila sect. Australophilae. All four species share a strong and distinctive musky odour, although it is not initially as strong in E. waitii the smell becomes more obvious in dried herbarium specimens. Notes. Chinnock (2007) noted that northern populations of E. microtheca in the Kalbarri National Park differed from more southern ones in the Fake Indoon-Fake Fogue depression area, in having distinctly flattened leaves. Brown et al. (2018) undertook a molecular study of the two forms and found significant divergences between plants found in the two areas. Subsequent field work and herbarium studies on the populations by one of the authors (A.P Brown) resulted in additional morphological 218 Nuytsia Vol. 30 (2019) features being identified and as a consequence, the southern form was described as a new species, Eremophila subangustifoUa (Brown et.al. 2018). Eremophila viridissima Chinnock, nom. et stat. nov. Eremophila glabra subsp. verrucosa Chinnock, Eremophila 592 (2007). Type: Lake Raeside, Western Australia, 22 September 1986, R.J.Chinnock 7273 {holo\ AD 98647195; iso\ K 000961395, MEL 2147389, PERTH 07538677, US 01050399). Etymology. From the Eatin viridissima, very green; referring to the deep green leaves. Figure 2. EremophilaviridissimafiowQnng stem showing the distinctive ascending, deep green leaves. Photograph by R.J. Chinnock. R. J. Chinnock, Additions to Eremophila (Scrophulariaceae) 219 Notes. Eremophila viridissima is well defined (see description in Chinnock 2007, p. 592) and is now considered as distinct from typical E. glabra as are satellite species like E. subteretifolia Chinnock, E. subfloccosa^Qnt\\. mdE. denticulataYMxxQW. (Figure2; see also Chinnock 2007, unnumbered Figure on p. 592 and Figure 297 p. 593, as E. glabra subsp. verrucosa). The species is readily distinguished from Eremophila glabra by its ascending, resinous deep green leaves, branches that are usually not divided in the upper parts and its distinctly verrucose endocarp of the fruit. When stressed, leaves of E. viridissima may turn yellowish green. When I published the account of Eremophila glabra in 2007,1 recognised nine subspecies but noted that the treatment was a tentative subdivision of this large polymorphic super species that includes a number of polyploid states (n=l 8,36,54). A number of these subspecies were known to be complexes in themselves like subsp. elegans Chinnock, subsp. albicans (Bartling) Chinnock and subsp. tomentosa Chinnock. Furthermore, these often showed intergradation e.g. subsp. elegans and subsp. tomentosa. To further complicate matters, I was also aware that some of the subspecies were essentially ‘buckets’ consisting of a number of taxa that required a more detailed study before segregation could occur. Subsp. tomentosa, for example, consists of at least three taxa: one in inland WA that forms large intricate plants 1.5 m tall and broad; the type form found on coastal cliffs around Kalbarri that is often a spindly shrub up to 1.8 m tall with one or a few main stems branched in the upper parts; and a third common in coastal areas around Geraldton and northwards, including off-shore islands, which is a prostrate plant or an erect shrub. These forms were beyond the scope of the monograph and required a more detailed study to make sense of these complexes. I also acknowledged thsit Eremophila glabra was part of a larger complex, which at that time included seven very closely related species that were actively in the process of speciation. Eremophila calcicola R.W.Davis has since been added to this complex (Brown & Davis 2016) and I am also aware of a number of other un-named taxa belonging to this group. Hopefully, the current molecular studies being undertaken by Dr R. Fowler at the University of Melbourne will resolve many of the problems found within the Glabra group. Acknowledgement I thank Karina Knight for providing scans of specimens of Eremophila waitii held at the Western Australian Herbarium. References Brown, A.P & Buirchell, B.J. (2007). Eremophila densifolia subsp. erecta and E. grandiflora (Myoporaceae), two new taxa from south west Western Australia. Nuytsia 17(1); 81-86. Brown, A.P. & Davis, R.W. (2016). Eremophila buirchellii and E. calcicola (Scrophulariaceae), two new species from Western Australia. Nuytsia 27; 211-216. Brown, A.P Llorens, T.M., Coates, D.J. & Byrne, M. (2018). Eremophila subangustifolia (Scrophulariaceae), a rare new species from the Mid West Region of Western Australia. Nuytsia 29; 17-20. Buirchell, B.J. & Brown, A.P. (2016). New species of(Scrophulariaceae); thirteen geographically restricted species from Western Australia. Nuytsia 27; 253-283. Chinnock, R.J. (2007). Eremophila and allied genera. A monograph of the plant family Myoporaceae. (Rosenberg Publications; Dural, New South Wales.) Chinnock, R.J. & Doley, A.B. (2011). Eremophila koobabbiensis (Scrophulariaceae), a new, rare species from the Wheatbelt of Western Australia. Nuytsia2\{A): 158-161. Edginton, M.A. (2015). Eremophila woodiae Edginton (Scrophulariaceae), a new species from Queensland. Austrobaileya 9(3); 408 415. Western Australian Herbarium (1998-). FloraBase—the Western Australian Flora. Department of Biodiversity, Conservation and Attractions, https;// florabase.dpaw.wa.gov.au/ [accessed 20 January 2019] 220 Nuytsia Vol. 30 (2019) Nuytsia The journal of the Western Australian Herbarium 30:221-228 Published online 15 October 2019 Triodia veniciae (Poaceae), a new species from the Pilbara region, Western Australia Matthew D. Barrett Kings Park and Botanic Garden, Dept, of Biodiversity, Conservation and Attractions, 1 Kattidj Close, Kings Park, Western Australia 6005 Western Australian Herbarium, Dept, of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 School of Biological Sciences, The University of Western Australia, Crawley, Western Australia 6009 Australian Tropical Herbarium, Sir Robert Norman Building (E2), James Cook University, McGregor Road, Smithfield, Queensland 4878 Corresponding author, email: matt.barrett@jcu.edu.au Abstract Barrett, M.D. Triodia veniciae (Poaceae), a new species from the Pilbara region. Western Australia. Nuytsia30\ 221-228 (2019). TheAustralian hummock grass genus TriodiaR.Br. is currently undergoing taxonomic revision following increased botanical survey in remote Australia. This paper describes a recently discovered species of ‘soft’ Triodia from the Pilbara region, T veniciae M.D.Barrett. The lemma lobes of the new species are narrow and produced into a short awnlet, intermediate between the old, and now united, generic concepts of Plectrachne Henrard with awned lemmas and Triodia with lobed lemmas. Triodia veniciae M.D.Barrett is only known from rocky shale slopes of the eastern Chichester Range, and has priority conservation status. Diagnostic character combinations for T veniciae are given to allow discrimination from all other Triodia. Introduction Triodia R.Br. hummock grasses are restricted to mainland Australia, where they are characteristic components of many arid and semi-arid communities, and the dominant elements of hummock grasslands that cover 18% of Australia (Department of the Environment and Heritage 2007). As at late 2018 the genus Triodia consists of 82 described and accepted species (Lazarides 1997; Barrett et al. 2005; Armstrong 2008; Barrett & Barrett 2011, 2015; Bmry et al. 2012; Crisp etal. 2015; Anderson et al. 2017a; Barrett & Trudgen 2018) but is currently undergoing phylogenetic investigation and taxonomic revision (e.g. Toon et al. 2015; Anderson et al. 2016, 2017b). This paper describes a new species of Triodia, in the group of less pungent species colloquially known as ‘soft’ spinifex. In 2016, during a survey of ploidy variation in Triodia in the eastern Pilbara, a morphologically anomalous variant was located on shale near Roy Hill Mine, referred to in an unpublished report under the informal name ''Triodia sp. Roy Hill (M.D. Barrett MDB 5412)’. Although this taxon had been previously overlooked amongst the similarly resinous T. epactia S.W.L.Jacobs, which is abundant in the same area, subtle features of its glumes, lemmas and leaf sheaths were found to be consistently discrete from all known Triodia species. Its morphology suggested a relationship with T. sp. Karijini (S. van Leeuwen 411 \),T. melvillei (C.E.Hubb.) Eazarides or T. basitricha M.D.Barrett. Despite these © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 222 Nuytsia Vol. 30 (2019) apparent similarities, DNA sequences from the ITS and ETS regions suggest that it is phylogenetically closer to T. bynoei (C.E.Hubb.) Eazarides of the eastern Kimberley and adjacent Northern Territory than to the aforementioned species. Previous reports of T. bynoei from the Pilbara region (Eazarides 1997; Eazarides et aJ. 2005) were based on misidentified specimens that are morphologically and genetically indistinguishable from T. melvillei. An intensive further survey for the new species located additional populations over a range of c. 140 km, between Roy Hill and Mulga Downs, and confirmed its exclusive occurrence on shale substrates. The Roy Hill Triodia is described below as a new species, T. veniciae. The lemma lobes of T. veniciae could be interpreted as either long lobes or short awns at the extremes of variation, and are here termed ‘sub-awned’. Along with T. hubbardii and T. triaristata, the sub- awned lemma lobes of T. veniciae provide an additional example of an intermediate between the old concepts of Plectrachne Henrard and Triodia, which were delimited on the presence or absence of lemma awns. As noted by Eazarides (1997), lemma lobing is untenable as a generic character, and the presence of an awn is homoplastic across the whole of Triodia s. lat. (Toon et al. 2015). Methods and terminology The term epistomatous (Toon etal. 2015; c/ amphistomatous) is taken up for the soft-type leaf anatomy in which stomata, stomatal grooves, and green chlorenchyma tissues are lacking adjacent to the lateral faces of the abaxial side of the leaf blade, with a corresponding lateral increase in colourless mesophyll tissue to fill the same space. Stomata are also extremely difficult to visualise, being protected within longitudinal grooves, so this condition is usually inferred from associated structures. Epistomatous leaf blades are diagnosed externally by a lack of stomatal grooves on the adaxial surface other than in a central band, and in section by the lack of chlorenchyma tissues adjacent to the lateral adaxial surfaces. Epistomatous leaf anatomy is synonymous with the terms ‘soft-type leaf anatomy’ or ‘centro- abaxial stomata’ that are sometimes used (e.g. Mant et al 2000). Use of the term ‘epistomatous’ for Triodia leaf blades is, however, slightly misleading, since stomatal grooves (and presumably also stomata) are still present on the abaxial surface but are confined to a median band. The conclusions made here are based on morphological data, but some notes are made on relationships inferred from phylogenetic analyses of sequence alignments from ITS and ETS (Internal Transcribed Spacer and External Transcribed Spacer respectively of the nuclear ribosomal DNA region). Extraction and sequencing protocols followed those described in Anderson etal (2016). Results mentioned here are based on preliminary phylogenetic trees generated using both Maximum Parsimony and Maximum Eikelihood tree generation methods. Full results of these analyses will be presented elsewhere. Taxonomy Triodia veniciae M.D.Barrett, sp. nov. Type'. North of Roy Hill Station, Western Australia [precise locality withheld for conservation reasons], 25 February 2017, M.D. Barrett 54U(holo\ PERTH 08776245; iso\ BRI, CANB, DNA, K,NSW, MEE). Hummock-forming perennial, very resinous, not obviously stoloniferous; hummocks loose, 30-70 cm high, 40-60 cm widQ,flowering culms 1.1-1.5 m high. Culm internodes 0. 1-10 cm long and obscured by subtending foliage, red-brown, glabrous; aerial roots absent. Fastigiate branching absQnt. Leaf sheaths 2.8-5.4 mm wide near apex, surface glabrous or sparsely to moderately pilose with hairs 1.5-3.2 mm M.D. Barrett, Triodia veniciae (Poaceae), a new species from the Pilbara 223 long, resinous or not resinous, pale green to straw coloured, prominently nerved; margins (excluding orifice) glabrous; margins of orifice truncate to oblique, with a dense fringe of hairs, the longest hairs 3.5- 5.5 mm long. Ligule a dense fringe of hairs 0.5-0.9 mm long; pseudopetiole obscure. Leaf blades fiattened-V-shaped when fresh, conduplicate and tightly in-rolled when dry, initially straight but becoming somewhat curled in older and dead leaves, 20-35 cm long, 0.8-1.5 mm wide when rolled, 1.5- 3 mm wide when unrolled, relatively soft when fresh, weakly pungent, glabrous abaxially and densely papillose adaxially, resinous or not over abaxial surface, bright green to dark green when fresh, dull green when dry; abaxial stomatal grooves confined to central part, 4 (2 either side of midrib), unequally spaced, absent on marginal c. 1/3 but the surface finely obscurely ribbed; abaxial stomatal grooves 7-8 each side of midrib; margins minutely scaberulous with prickle hairs c. 0.05 mm long. Panicle linear to ovate, 14-37 cm long, 1.0-15 cm wide, with 61-192 spikelets; primary axis minutely antrorsely scabrous, angular and ribbed; branches openly racemose with some basal branches ternate, loose, glabrous including the branch axils, non-resinous; longest basal panicle branches 5.6-15.8 cm long, sub-terete to angular, with 6-15 loosely-arranged ± uniform-sized spikelets, which are 7-16 mm apart (the distance between point of insertion of adjacent pedicels along the branch) with adjacent glumes not or slightly overlapping; longest basal pedicels (on longest basal panicle branches) 4.3- 14 mm long, 0.1-0.2 mm wide, ± filiform, becoming slightly thicker just below spikelet, angular to slightly fiattened, minutely scabrous; terminal pedicel 7-14 mm long; Spikelet 8-15 mm long, 1.9- 3.9 mm wide, loosely 4-8-fiowered with 3-7 fertile florets, sometimes with 1 apparently sterile (but possibly immature) fioret at apex, narrowly elliptic to narrowly oblanceolate, compressed at maturity; lowest rachilla internode c. 0.8-1.7 mm long, c. 0.15-0.2 mm diam., glabrous. Lower glume narrowly lanceolate, 5.4-8.5 mm long, 1.0-1.6 mm wide, L:W 5.0-6.8, apex acute, shorter than the combined fiorets (including lobes), scarious, sub-membranous to chartaceous, with minute scabrosities less than 0.1 mm long over whole surface or only on margins and apex, 3-5-nerved, the midnerve distinctly raised, lateral nerves slightly raised; margins narrowly membranous, glabrous. Upper glume inserted c. 0.3-0.6 mm above lower glume, 5.0-8.5 mm long, similar to and subequal to lower glume, shorter than the combined fiorets (including lobes). Lowest lemma narrowly elliptic to narrowly lanceolate, 6.6- 8.5 mm long including lobes, uniformly chartaceous, deeply 3-lobed, the midlobe somewhat drawn out into an awnlet; body 3.8-5.2 mm long including callus, 0.8-1.1 mm wide, the lower 1/2-3/4 with dense indumentum of appressed to lifting hairs 0.15-0.35(0.5) mm long arranged in 6-10 rows (or hair lines sometimes missing from the central zone), with nerves mostly obscure, but the upper part with 3 groups of 3 faint nerves visible and radiating into the lobes; midlobe 1.8-3.2 mm long, narrowly triangular at base narrowing to apex and sometimes produced into a short awnlet, similar in width to the lateral lobes; lateral lobes 1.0-2.3 mm long, 1/3-3/4 as long as midlobe, narrowly triangular at base, long-acuminate at apex but lacking a distinct awnlet, margins with a narrow membranous wing; callus 0.2-0.6 mm long, straight to distinctly curved in profile, attached obliquely, blunt to acute with an abruptly blunt apex in face view, acute in profile, white-bearded except in midline, the longest hairs 0.3-0.6 mm long. Upper lemmas similar to but smaller than lowest lemma, the midlobe becoming more awn-like toward the apex of the spikelet (i.e. the awnlet becomes a more prominent fraction of the length as lemmas decrease in size). Palea of basal lemma very slightly longer than lemma body, oblanceolate, 4.0-5.6 mm long, 0.6-0.8 mm wide, 2-keeled, not bitextured, membranous, glabrous or with a tuft of hairs between 1/4 and 1/2 of the length, apex acute, entire or erose; keels with a very narrow wing c. 0.1 mm wide, keel margins scabrous; flaps c. 0.15 mm wide, broadest in central part, narrower than 1/2 width of the palea body and not overlapping, entire. Lodicules c. 0.4 mm long, apex ± truncate, undulate, glabrous. Anthers 3,1.9-2.7 mm long, exserted at maturity. Styles 2, c. 0.5- 0.9 mm long. Caryopsis lenticular to narrowly obovate, 2.0-2.2 mm long, c. 0.7 mm wide, L:W 3.2- 3.5:1, abruptly obtuse at apex, ± acute at base, slightly dorsiventrally flattened in section, with a broad shallow furrow on ventral side, pale reddish straw-coloured, base of styles only slightly thickened; hilum 0.55-0.65 mm long, c. 0.27-0.30 times as long as caryopsis. (Figure 1) 224 Nuytsia Vol. 30 (2019) Figure 1. Triodia veniciae. A- flowering plants in situ showing habit and broken shale substrate; B - spikelet, showing the awned lemmas, variation in florets and relative midlobe (awnlet) length within a spikelet, and glumes shorter than combined florets; C - glumes, showing shape in side view; D - lemma of basal florets in side and face view showing relative proportions of body and lobes; the intense purple pigmentation in this specimen is atypical; E - base of lemma of basal floret in side and face view, showing hairs in longitudinal rows; F - palea in face view showing zone hairs in central area (paleas can also be glabrous), and very shortly winged lemma keels. Scale bars = 1 mm. Images from M.D. Barrett MDB 5414. Photographs by M. Barrett. Diagnostic features. Foliage copiously resinous. Leaf sheaths glabrous or hairy on surfaces, glabrous on margins. Leaf blades epistomatous (soft-type), lacking stomatal grooves on the lateral margins of the abaxial surface. Panicles open, prominently ternately branched, lacking hairs in panicle axils. Glumes 5.4-8.5 mm long, narrowly lanceolate. Lemmas with short callus 0.2-0.3 mm long, body uniformly textured, lobes long-acute to shortly awned, the midlobe 1.8-3.2 mm long. Palea uniformly textured. Habitat restricted to shale slopes of the Chichester Range, Pilbara. M.D. Barrett, Triodia veniciae (Poaceae), a new species from the Pilbara 225 Other specimens examined. WESTERN AUSTRALIA: Chichester Range, [precise localities withheld for conservation reasons] 6 Aug. 2016, M.D. BarrettMDB 5408 (PERTH); 8 Aug. 2016, M.D. Barrett MDB 5412 (PERTH); 22 Feb. 2016, M.D. Barrett MDB 5415 (PERTH); 19 Mar. 2016, M.D. Barrett MDB 5416 (PERTH); 22 June 2017, M.D. Barrett & N. Bezemer MDB 5421 (PERTH); 23 June 2017, M. D. Barrett & N. Bezemer MDB 5424 (PERTH); 24 June 2017, M.D. Barrett & N. Bezemer MDB 5434 (PERTH); 25 June 2017, M.D. Barrett & N. Bezemer MDB 5440 (PERTH); M.D. Barrett & N. Bezemer MDB 5441 (PERTH); M.D. Barrett & N. Bezemer MDB 5442 (PERTH); 27 June 2017, M.D. Barrett & N. Bezemer MDB 5455 (PERTH); 27 June 2017, M.D. Barrett & N. Bezemer MDB 5458 (PERTH). Phenology. Observed flowering February to March. Aseasonal fertile collections (very rare plants flowering among thousands of non-flowering plants) have been made in June and August following significant winter rainfall. Distribution and habitat. Occurs on slopes of low hills of broken shale on the northern/eastern flanks of Marra Mamba mesa formations, along a 140 km transect of the eastern Chichester Ranges, between Roy Hill Station in the east and Mulga Downs Station in the west (Figure 2). Often occurs intermixed with or immediately adjacent to T. epactia. Replaced by other species such as T. brizoides, T. scintillans and T. wiseana on the more widespread ironstone hills in the area. Conservation status. Triodia veniciae is to be listed as Priority One under Conservation Codes for Western Australian Flora (A. Jones pers. comm.). It is currently known from 27 patches, but aggregating patches of close proximity (0.1-2 km apart) consolidates the known distribution to 14 ‘populations’, spread over a 140 km distance along an almost linear zone of the eastern Chichester Range system. These 14 populations were estimated to have a total of more than 144,500 clumps. Some of the populations are potentially under threat due to the existence of mines in close proximity to known patches. Further evidence of substantial genetic division within T. veniciae might also increase the protection required for the eastern genetic lineage. Etymology. The epithet vcw/czhfc is after VeniciaDe San Miguel, fromRoy Hill Environment Department, who was instrumental in the discovery, collection of type material and surveys for this species. Affinities. Morphologically, T. veniciae is similar to a number of other Triodia species from the Pilbara. It co-occurs with the similarly resinous T. epactia, with which it is easily confused in the held. Triodia epactia (and T. pungens R.Br.) can be separated from T. veniciae by the broader ovate to elliptic glumes (L:W 1.7—4.0), glabrous sheath surfaces in T. epactia / T. pungens, and lemma hairs present only along the midline and margins, compared to lanceolate to narrowly lanceolate glumes (L:W > 5.0-6.8), frequently hairy sheath surfaces, and lemma hairs concentrated in many longitudinal rows in T. veniciae. Triodia veniciae is similar to T. sp. Karijini (S. van Leeuwen 4111), in having narrow lemma lobes 2.0-3.2 mm long that could be interpreted as short awns, but differs in being copiously resinous (non- resinous to weakly resinous in T. sp. Karijini), in having paleas usually with some hairs (palea always glabrous in T. sp. Karijini) and 6-15 spikelets on its longest basal panicle branches (3-6 spikelets in T. sp. Karijini). Triodia sp. Karijini is only known from above 900 m on ironstone ridges on mountains in the Hamersley Range, while T. veniciae is only known from shale slopes in the Chichester range north of the Fortescue River (Figure 2). 226 Nuytsia Vol. 30 (2019) Figure 2. Distribution map of Triodia veniciae, showing the location of known populations along the eastern Chichester Range just to the north of the Fortescue Valley sub-region of the Pilbara bioregion. Black circles - eastern diploid populations. Black triangle - eastern tetraploid population. Black square - western population that is genetically somewhat divergent and is inferred to have a reduced-tetraploid DNA content, see text for details. Grey circles - eastern populations not assessed for ploidy. Triodia veniciae is also similar to T. melvillei (to which it would key in Lazarides et al. 2005), sharing a very resinous habit, narrow glumes and lemma hairs concentrated in longitudinal rows. However, the lemmas are 3-awned in T. melvillei (lowest lemma midlobe 5.5-6.8 mm long in Pilbara specimens), compared to lobed to sub-awned with lowest lemma midlobe 2.1-3.2 mm long in T. veniciae. The leaf sheaths are most frequently hairy on their surfaces in T. veniciae, whereas they are always glabrous in T. melvillei (at least in hundreds of specimens seen by me, including all Pilbara specimens at CANB and PERTH, however Lazarides (1997) reports lower sheaths can be partially hirsute, without reporting specimen details nor their origin). The panicle axis and branches are strongly scabrous in T. veniciae but only weakly scabrous in T. melvillei. The distributions of T. melvillei and T. veniciae do not overlap, with T. veniciae restricted to the Chichester Range north of the Fortescue River, while T. melvillei occurs in the Pilbara south of the Fortescue River, south and east into central Australia. The Pilbara species T. basitricha also has hairy leaf sheaths, but has awned lemma lobes, and lemmas bitextured with a sharp transverse demarcation line (uniformly textured in T. veniciae). Although both species occur in the eastern Pilbara, they are not known to occur in close proximity. Specimens of T. veniciae lacking florets can be distinguished by having shorter lower glumes 5.4-8.5 mm long (10- 12.5 mm long in T. basitricha) and more numerous spikelets on longest basal branches (7-14 compared to 4-5 in T. basitricha). Despite the morphological similarities of the above Pilbara species, ITS and ETS sequences (data not shown) suggest T. veniciae has a closer relationship to the Kimberley and Top End species T. bynoei. Triodia bynoei has distinct lemma awns, and lemma hairs scattered over the surface, not clearly concentrated in rows. M.D. Barrett, Triodia veniciae (Poaceae), a new species from the Pilbara 227 Notes. Preliminary unpublished genetic investigation suggests that there may be two genetic units within T. veniciae, as plants from the Mulga Downs patches (Western group, n=2) consistently differ from samples from more eastern populations (Eastern group, n=6) in three and five substitutions in ITS and ETS regions respectively (data not shown). The Mulga Downs patches also appear to have a ‘reduced tetraploid’ DNA content based on flow cytometric measurements of DNA content, consistent with a long-divergent tetraploid population that is in the process of becoming diploidised (Wendel 2015). Flow cytometric measurements of DNA content from populations in the Eastern group are predominantly diploid, or in one population tetraploid with a typical 2:1 ratio relative to diploid populations (M. Barrett unpublished data). Field investigations have shown that plants in the western-most populations on Mulga Downs always have glabrous leaf sheaths, while in those around Roy Hill Station and Christmas Creek Mine always have sparsely to moderately densely hairy sheaths. In geographically intermediate areas near Cloudbreak Mine (genetically belonging to the ITS/ETS Eastern group), plants can have either glabrous or hairy sheaths, and these populations often have a mixture of the two morphotypes. As a result, the glabrous sheath character is not completely diagnostic for the Western group lineage. No other characters differing between the two groups are known, and characteristics from Mulga Downs specimens {M.D. Barrett & N. Bezemer MDB 5440 to MDB 5442) were always within the variation found amongst other collections made from further east. The vernacular name Roy Hill Spinifex is suggested. Acknowledgements The Roy Hill Environment Department, in particular Venicia De San Miguel, Todd Bell and Susanna Beech, are thanked for providing fieldwork and logistical assistance. Roy Hill Holdings Pty Etd provided funding for Triodia surveys and for the formal description of T. veniciae. References Anderson, B.M., Barrett, M.D., Krauss, S.L. & Thiele, K. (2016). Untangling a species complex of arid zone grasses {Triodia) reveals patterns congruent with co-occurring animals. Molecular Phylogenetics and Evolution 101:142-162. doi: 10.1016/j. ympev.2016.05.014. Anderson, B.M., Thiele, K.R. & Barrett, M.D. (2017a). A revision of the Triodia basedowii species complex and close relatives (Poaceae: Chloridoideae). Australian Systematic Botany 30; 117-229. Anderson, B.M., Thiele, K.R., Krauss, S.L. & Barrett, M.D. (2017b). Genotyping-by-Sequencing in a Species Complex of Australian Hummock Grasses {Triodia) : Methodological Insights and Phylogenetic Resolution. PLOSOne 12( 1): eO 171053. doi:10.1371/journal.pone.0171053. Armstrong, G. (2008). Triodia caelestialis (Triodieae: Chloridoideae: Poaceae), a new species from the central Kimberley, Western Australia. Journal of the Royal Society of Western Australia 91:313-317. Barrett, R.L., Wells, G.B. & Dixon, K.W. (2005). New taxa and combinations: Subfam. Chloridoideae, Trib. Triodieae, Triodia. In: K Mallett, (ed.). Flora of Australia 44B pp. 458-459. (CSIRO Publishing: Melbourne) Barrett, R.L. & Barrett, M.D. (2011). Two new species of Triodia (Poaceae: Triodieae) from the Kimberley region of Western Australia. Telopea 13: 57-67. Barrett, R.L. & Barrett, M.D. (2015). Twenty-seven new species of vascular plants from Western Australia. Nuytsia 26'. 21-87. Barrett, M.D. & Trudgen, M.E. (2018). Triodia pisoliticola (Poaceae), a new species from the Pilbara region. Western Australia, and a description for T. sp. Mt Ella (M.E. Trudgen MET 12739). Nuytsia 29: 271-281. Crisp, M.D., Mant, J., Toon, A. & Cook, E.G. (2015). Australian spinifex grasses: new names in Triodia for Monodia and Symplectrodia. Phytotaxa 230: 293-296. 228 Nuytsia Vol. 30 (2019) Department of the Environment and Heritage (2007). Australia’s native vegetation: a summary of Australia’s major vegetation. Canberra, ACT. http://www.environment.gov.au/resource/australias-native-vegetation-summary-australias-major-vegetation- groups [accessed 5 May 2017]. Hurry, C.R., Walsh, N.G. &Murphy, D.J. (2012). Ataxonomicreviewof rnW/a/>M?7/co/aandT5Cflno5a(Poaceae: Chloridoideae), based on morphological and molecular data. Australian Systematic Botany 25: 304-312. doi:10.1071/SB10044. Lazarides, M. (1997). A revision of Triodia including Plectrachne (Poaceae, Eragrostideae, Triodiinae). Australian Systematic Botany 10: 381-489. doi:10.1071/SB96012. Eazarides, M., Weiller, C. & McCusker, A. (2005). Triodia. In. K Mallett, (ed.). Flora of Australia 44B, pp. 203-256. Mant, J.G., Bayer, R.J., Crisp, M.D. & Trueman, J.W.H. (2000). A phylogeny ofTriodieae (Poaceae: Chloridoideae) based on the ITS region of nrDNA: Testing conflict between anatomical and inflorescence characters. In. Jacobs, S.W.E. & Everett, J. (eds.). Grasses: Systematics and Evolution, pp. 213-217. (CSIRO Publishing: Melbourne.) Toon, A., Crisp, M.D., GamageH., Mant, J., Morris, D.C., Schmidt, S., Cook, E.G. (2015). Key innovation or adaptive change? A test of leaf traits using Triodiinae in Australia. Scientific Reports 5: 12398. Wendel, J.F. (2015). The wondrous cycles of polyploidy in plants. American Journal of Botany 102:1753-1756, Nuytsia The journal of the Western Australian Herbarium 30:229-235 Published online 15 October 2019 SHORT COMMUNICATION Styphelia quartzitica and S, rectiloba (Ericaceae: Epacridoideae: Styphelieae), two new, morphologically anomalous species of restricted distribution Recently published research by Puente-Lelievre et al. (2016) dealing with the molecular phylogeny of the large Styphelia Sm.-Astroloma R.Br. clade has led to the adoption of a greatly enlarged circumscription for Styphelia (Crayn et al. 2019). The sampling of the Western Australian taxa for that project was both relatively dense (96 taxa, or about two-thirds of the western members of the clade were sampled) and strategic, with all major morphological groups included, in addition to most of those taxa that showed no clear morphological athnities. However two uncommon species, which did not fit well with any morphological group, were not sampled. While their closer affinities are uncertain and will require further molecular investigation to resolve, there is little reason to doubt that the two are Styphelia in the newly expanded sense. They are described below. Styphelia quartzitica Hislop, sp. nov. Typus\ Fitzgerald River National Park, Western Australia [precise locality withheld for conservation reasons], 17 May 2016, M. Hislop 4596 {holo\ PERTH 08836639; iso\ CANB, K, MEL, NSW). Leucopogon sp. Barren Range (A.S. George 10092), Western Australian Herbarium, in FloraBase, https://fiorabase.dpaw.wa.gov.au/ [accessed 21 June 2019] Erect shrubs to 60 cm high and 40 cm wide, single-stemmed at ground level from a fire-sensitive rootstock. Young branchlets with a dense indumentum of mostly retrorse hairs, 0.05-0.20 mm long. Leaves spirally arranged, crowded, variably antrorse, or occasionally some leaves ± patent; apex long- mucronate, pungent, mucro 0.4-1.0 mm long; base attenuate; petiole rather obscure, 0.3-0.8 mm long, shortly hairy on adaxial surface and margins; lamina linear or very narrowly ovate, 7-14 mm long, 0.8-1.5 mm wide, fiat or adaxially concave, the longitudinal axis gently incurved, usually distinctly twisted; surfaces markedly discolorous, shiny; adaxial surface shortly and sparsely hairy, ± transversely rugose, the venation not evident; abaxial surface paler, glabrous, with 5 fiat or slightly raised primary veins; margins glabrous or minutely scabrid. Inflorescence axillary, widely spreading; axis 1.4-2.6 mm long, 1- or less often 2-fiowered, ± terete in lower portion, plano-convex above the fertile node, terminating in a bud-rudiment; axis indumentum moderately dense, c. 0.1 mm long; fiowers widely spreading, sessile. Fertile bracts broadly ovate to depressed-ovate, 0.4-0.6 mm long, 0.4-0.6 mm wide, subtended by 4 or 5 sterile bracts. Bracteoles broadly ovate to depressed-ovate, 1.0-1.3 mm long, 1.0-1.4 mm wide, obtuse; abaxial surface not keeled, glabrous, greenish, ± striate; margins minutely ciliolate. Sepals narrowly ovate, 2.8-3.6 mm long, 1.2-1.3 mm wide, acute or subacute, shortly mucronate; abaxial surface glabrous, pale greenish to straw-coloured, venation obscure; adaxial surface with a discrete patch of very short, sparse hairs close to the base; margins minutely ciliolate, with hairs < 0.05 mm long. Corolla tube cream, ± cylindrical or narrowly obovoid, longer than the sepals, 3.0^.5 mm long, 1.7-2.2 mm wide, glabrous externally; internal surface densely hairy in a medial © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 230 Nuytsia Vol. 30 (2019) band, glabrous at base. Corolla lobes cream, shorter than the tube, 2.0-2.5 mm long, 1.0-1.2 mm wide at base, erect in lower 1/2-2/3 of their length and then spreading and recurved; glabrous externally, internal surface with a sparse indumentum of twisted and ornamented hairs, concentrated towards the margins. Filaments terete, 0.3-0.4 mm long, attached to anther 2/3-3/4 above base, adnate to tube just below sinuses. Anthers partially exserted from the tube (by 1/3-1/2 of their length), 1.2-1.3 mm long, apex rounded to shallowly emarginate. Nectary annular, shallowly lobed, 0.3-0.4 mm long, glabrous, usually grooved longitudinally below the sinuses. Ovary pale green, narrowly ovoid, c. 1.3-1.6 mm long (but refer comments under notes below), 0.6-0.8 mm wide, glabrous, 5-locular. Style glabrous or minutely papillose in the upper half, c. 2.3-3.6 mm long (but refer comment under notes below), tapering smoothly from ovary apex, with stigma presented at the top of the corolla tube; stigma not or barely expanded. Frwzf narrowly ellipsoid or obloid, 3.2^.0 mm long, 1.6-1.8 mm wide, much longer than the calyx; surface glabrous, with longitudinal grooves, mesocarp poorly developed; gynophore absent; style persistent. (Figure 1) Figure 1. Styphelia quartzitica. A - scanned image of flowering branchlet; B - flower, slit open longitudinally; C - fruit. Scale bars = 1 cm (A), 1 mm (B & C). Vouchers: M. Hislop 4596 (A & B), M. Hislop 3040 (C). Drawings by Skye Coffey. M. Hislop, Styphelia quartzitica and S. rectiloba (Ericaceae), two new species 231 Diagnostic characters. Readily distinguished from all other western Styphelia by the following combination of characters: leaves linear, longitudinally twisted; internal surfaces of corolla lobes with a sparse indumentum concentrated towards the margins; internal corolla tubes with a medial band of dense hairs; fruit ± dry, longitudinally grooved, narrowly ellipsoid or obloid, lacking a gynophore. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 4Aug. 1995, S Barrett323. \ (PERTH); 26Nov. 2002, S. Barrett 1054 (PERTH); 9 Oct. 2007, S. Barrett 1648 (PERTH); 26 Mar. 2012, S. Barrett2094 (PERTH); 16 July 1970, A.5. George 10092 (PERTH); 10 Oct. 2003, M. Hislop 3034 (CANB, NSW, PERTH); 10 Oct. 2003, M. Hislop 3040 (MEE, PERTH^ 28 Nov. 2002, M Hislop, S. Barrett & J.A. Cochrane MH 2876 (PERTH); 30 May 1970, K.R. Newbey 3 164 (PERTH); 17 Sep. 2011, E>. A. Rathbone 997 (PERTH); 22 Sep. 2011, D.A. Rathbone 998 (PERTH). Distribution and habitat. Restricted to the upper slopes of quartzite hills and mountains in remote parts of the Fitzgerald River National Park within the Esperance Plains bioregion (Department of Environment 2013). Grows in shallow sandy soils over quartzite in the understory of dense heath. Commonly associated species include Regelia velutina, Melaleuca lutea, Agonis baxteri, Taxandria conspicua and Banksia oreophila. Phenology. Apparently has a prolonged flowering period with a peak most likely between March and May, although flowering collections have been made as late as October. Specimens with mature fruit have been collected between July and November. Etymology. The epithet is Eatinised from quartzite and refers to the rock type on which the species is found. Conservation status. Currently listed by Smith and Jones (2018) as Priority Two under Conservation Codes for Western Australian Flora under the name Leucopogon sp. Barren Range (A.S. George 10092). All populations of this species are conserved in the Fitzgerald River National Park. However its occurrence there is very localised and habitat-speciflc. Affinities. Styphelia quartzitica is a very distinctive species and no close morphological aflhnities are apparent with any of the species-groups resolved in the recent molecular phylogenetic study (Puente- Eelievre et al. 2016). The unevenly distributed corolla lobe hairs and dense medial band of hairs in the internal corolla tube are both unusual features for the genus. The conspicuously twisted leaves are another uncommon attribute. Notes. Because the style tapers so gradually from the ovary apex in this species the given measurements for the ovary and style lengths are necessarily imprecise. Styphelia rectiloba Hislop, sp. nov. Typus: west of Kambalda, Western Australia [precise locality withheld for conservation reasons], 23 May 2013, M. Hislop 4249 {holo\ PERTH 08514054; iso\ CANB, MEE, NSW). Leucopogon sp. Kambalda (J. Williams s.n. PERTH 07305028), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 21 June 2019] 232 Nuytsia Vol. 30 (2019) Spreading, compact shrubs to c. 1.5m high and 1.5m wide, branching from close to base but probably with a fire-sensitive rootstock. Young branchlets with a sparse to moderately dense indumentum of very short, patent hairs, < 0.05 mm long. Leaves spirally arranged, variably antrorse; apex long- mucronate, pungent, the mucro 0.8-2.0 mm long; base rounded to cuneate; petiole well-defined, 0.5-1.0 mm long, usually with a few hairs about the adaxial surface and margins or ± glabrous; lamina ovate to narrowly ovate, 6-11 mm long, 2.3-3.5 mm wide, strongly concave adaxially, the longitudinal axis variable, from distinctly incurved to distinctly recurved; surfaces discolorous; adaxial surface glabrous, ± glaucous on younger leaves, becoming shiny, the venation not or barely evident, often with transverse striations; abaxial surface paler, matt, with 5 usually slightly raised primary veins, openly grooved or flat between veins, glabrous or very shortly and sparsely hairy; margins glabrous, irregularly denticulate. Inflorescence axillary, erect; axis 2.4-3.5 mm long, l-3(4)-flowered, ± terete basally, becoming sharply angular in flowering portion, terminating in a bud-rudiment; axis indumentum moderately dense to c. 0.2 mm long; flowers erect, sessile. Fertile bracts ovate or broadly ovate, 0.7-1.2 mm long, 0.7-1.0 mm wide, subtended by 4 or 5 sterile bracts. Bracteoles ovate- elliptic 1.8-2.5 mm long, 1.4-1.6 mm wide, obtuse to ± retuse; abaxial surface not keeled, shortly and sparsely hairy, greenish but often tinged brown at least in the upper half; margins glabrous or minutely ciliolate. Sepals narrowly ovate, 3.8^.5 mm long, 1.4-1.7 mm wide, obtuse with a short, subapical mucro; abaxial surface shortly hairy towards apex, glabrous below, initially greenish, but soon becoming brown, at least in the distal half, venation obscure; adaxial surface glabrous; margins ciliate with hairs to 0.2 mm long, mostly towards the apex. Corolla tube cream, cylindrical, usually slightly shorter than, to occasionally distinctly longer than, the sepals, 3.4^.2 mm long, 1.3-1.6 mm wide; external surface usually with a few long hairs in lines; internal surface hairy in the upper 2/3, glabrous below. Corolla lobes cream, shorter than the tube, erect for up to c. 1/4 of their length and then spreading and recurved (but refer to comment under Affinities heading below), 2.4-3.2 mm long, 0.8-1.0 mm wide at base, glabrous externally; internal surface with a rather sparse indumentum of twisted and ornamented hairs. Filaments terete, 0.7-0.9 mm long, attached to anther 2/3-3/4 above anther base, adnate to tube just below sinuses. Anthers fully exserted from the tube and held at right angles to the floral axis post-anthesis, 1.2-1.4 mm long, apex emarginate. Nectary partite, the scales 0.45-0.55 mm long, 0.35-0.45 mm wide, glabrous. Ovary pale yellow-green, globose to ellipsoid, 0.6-0.8 mm long, 0.5-0.6 mm wide, hairy, 3-locular. Style sparsely hairy in lower 2/3-3/4, scabrous above, 4.2-5.3 mm long, tapering smoothly from ovary apex, well-exserted from the corolla with the stigma presented well beyond the corolla lobe bases and anthers; stigma expanded. Fruit narrowly ellipsoid, c. 3.0^.0 mm long, 1.5-2.0 mm wide (but see comment under below), c. equal to the sepals, surface sparsely hairy, with irregular, raised, ribs representing the dried mesocarp; gynophore absent; style shed at or close to maturity. (Figure 2) Diagnostic characters. Readily distinguished from all other western Styphelia by the following combination of characters: leaves ovate, strongly concave adaxially with irregularly denticulate margins and pungent, long-mucronate apices; corolla lobes straight and sharply reflexed on dried specimens; anthers exserted from the tube and held at right angles to the floral axis post-anthesis; ovary hairy, 3-locular, style hairy at least in the lower half; fruit narrowly ellipsoid, with a thin mesocarp, lacking a gynophore. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 13 Jan. 2006, V. Clarke VTC 647 (NSW, PERTH); 10 May 2012, M Hislop 4194 (PERTH); 10 May 2012, M.HislopAl91 (K,PERTH); 14Dec. 2011, J. Warden33627 (PERTH); May 2005, J. Williams s.n. (PERTH); 10 June 2013, V. Yeomans & A. Sleep VY 840-05 (PERTH). M. Hislop, Styphelia quartzitica and S. rectiloba (Ericaceae), two new species 233 Figure 2. Styphelia rectiloba. A - scanned image of flowering branchlet; B - flower, slit open longitudinally; C - fruit. Scale bars = 1 cm (A), 1 mm (B & C). Vouchers: M. Hislop 4249 (A), M. Hislop A\91 (B & C). Drawings by Skye Coffey. 234 Nuytsia Vol. 30 (2019) Distribution and habitat. Currently only known from a few populations in the Kambalda area of the Coolgardie bioregion (Department of Environment 2013) where the species is restricted to rocky, skeletal soils, on and in close proximity to, decomposed granitic breakaways. The associated vegetation is very open woodland or heath with Eucalyptus stricklandii, Melaleuca leiocarpa, Alyxia buxifolia and Ptilotus helichrysoides among the more frequently encountered species. Phenology. Flowering specimens have been collected between December and June. The only collection to include mature fruit was made in May. Peak flowering is probably dependent on the extent and pattern of rainfall through the summer months but in average seasonal conditions it is likely to be between March and May. Fruit are likely to be present at least between May and September. Etymology. From the Fatin recti- (straight) and lobus (lobe), in reference to the noticeably straight axis of the dried corolla lobes (see Affinities below). Conservation status. Fisted by Smith and Jones (2018) as Priority Three under Conservation Codes for Western Australian Flora under the name Leucopogon sp. Kambalda (J. Williams s.n. PERTH 07305028). This species is localised, habitat-specific and known from very few populations. However it does occur in a remote part of the state where there is considerable potential for the discovery of new populations. Surveys are needed to better understand the extent of its distribution. Affinities. As with S. quartzitica described above, S. rectiloba has no obvious morphological aflhnities with any of the species groups resolved in the recent molecular phylogenetic study (Puente-Felievre etal. 2016). The epithet highlights an interesting feature of the species. Although there is nothing unusual in the orientation of the corolla lobes in live flowers, these being erect towards the base and then spreading and recurved above, when dried the lobes become straight and reflexed. This characteristic does not occur elsewhere in the western members of the genus. Another unusual feature relates to the disposition of the anthers. While the anthers are slightly exserted from the corolla tube at anthesis, once the pollen is shed they are held at right angles to the floral axis, and in so doing appear more prominent. The species is also notable for the presence of stylar hairs, an uncommon character in the western Styphelia. Notes. The fruit measurements are based on a single fruiting collection and therefore cannot be regarded as definitive. Acknowledgements I would like to thank Skye Coffey for the fine illustrations, Julia Percy-Bower for help with specimen photographs and Kelly Shepherd for formatting the images to best effect. References Crayn, D.M.,Hislop,M.& Puente-Lel ievre, C. (2019). A phylogenetic recircumscription of Styphelia (Ericaceae: Epacr ido ideae: Styphelieae). Australian Systematic Botany (accepted 4 June 2019) Department of the Environment (2013). Australia’s bioregions (IBRA). IBRA 7, Commonwealth of Australia. http://www. environment.gov.au/land/nrs/science/ibra#ibra [accessed 19 December 2018] Puente-Eelievre, C., Hislop, M., Harrington, M., Brown, E.A., Kuzmina, M. & Crayn, D.M. (2016). A five-marker molecular phylogeny of the Styphelieae (Epacridoideae, Ericaceae) supports a broad concept of Styphelia. Australian Systematic Botany 2^'. 368-387. M. Hislop, Styphelia quartzitica and S. rectiloba (Ericaceae), two new species 235 Smith, M.G. & Jones, A. (2018). ThreatenedandPriority Floralist 5 December 2018. Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 21 June 2019]. Michael Hislop Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 Author, email: michael.hislop@dbca.wa.gov.au 236 Nuytsia Vol. 30 (2019) Nuytsia The journal of the Western Australian Herbarium 30:237-245 Published online 15 October 2019 Calandrinia monosperma and C. uncinella (Montiaceae), two new indehiscent species from Western Australia Frank J. Obbens c/o Western Australian Herbarium, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 Corresponding author, email; frank.obbens@aapt.net.au Abstract Obbens, F.J. Calandrinia monosperma and C. uncinella (Montiaceae), two new indehiscent species from Western Australia. Nuytsia 30: 237-245 (2019). Two unique indehiscent species of Calandrinia Kunth. from Western Australia are described and mapped and their affinities are discussed with regards to a recent phylogeny for Australian Calandrinia. Introduction This paper describes two new annual species of Calandrinia Kunth. from Western Australia, both having capsules that are indehiscent and deciduous. The newly described C. monosperma Syeda ex Obbens is a prostrate, small to medium-sized species with a widespread distribution in the IBRA Eremaean bioregion, while C. uncinella Obbens is a decumbent, usually smaller-sized species of patchy distribution within the IBRA Southwest bioregion. The distinctive capsules of both these species make them readily identifiable when in fruit, however, their seeds are also quite distinctive. This is somewhat unusual, as seed shape and pattern are generally considered important diagnostic characters within the genus. Hancock et a/.’s (2018) recent molecular phylogeny for Australian Calandrinia, based on a targeted gene enrichment approach, placed C. monosperma (as C. sp. The Pink Hills (F. Obbens FO 19/06)) within their clade 3 and C. uncinella (as C. sp. Piawaning (A.C. Beauglehole 12257)) within their clade 5. Two other indehiscent species, C. disperma J.M. Black and Rumicastrum chamaecladum (Diels) Ulbr. were also placed in clade 3. Both these species also have unusual and distinctive fruits; however their fruits and seeds are quite different to the above, allowing for all four indehiscent species to be quite easily recognised. Methods Methods used are the same as those described in Obbens (2011). In both C. monosperma and C. uncinella it is impossible to distinguish between stems and scapes. Therefore, in this paper, the term ‘flowering shoot’ is used to refer to the length of the stem, scape and inflorescence axis measured as one unit along the main axis of the shoot. The length of the inflorescence axis is also recorded separately. The bioregions referred to in describing species distributions and indicated on the map are from Australia’s bioregions (IBRA) (Department of the Environment 2013). © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 238 Nuytsia Vol. 30 (2019) Taxonomy Calandrinia monosperma Syeda ex Obbens, sp. nov. Type'. Site is c. 3.8 km E along Talawana Track from the turnoff into Balfour Downs homestead, Pilbara region, Western Australia, 1 June 2004, F. Obbens 8 l B. Bromilow FO 32/04 {holo: PERTH 06609740; iso: CANB). Calandrinia sp. The Pink Hills (F. Obbens FO 19/06), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed June 2019]. Prostrate, sometimes slightly decumbent annual herbs, 10-110 mm tall, 60-385 mm wide, glabrous, the root system comprising a small taproot (occasionally larger) with several fibrous roots. Basal leaves succulent, narrowly linear, sometimes slightly curved inwards, 7.5-80 mm long, 1.2-3.8 mm wide, terete to subterete in mid cross-section, green to brownish-green, often turning blackish when aging. Flowering shoots usually 4-11 (-20), 25-270 mm long, radiating from base, the proximal 10^8 mm leafless, thereafter very leafy including proximal portions of inflorescence axes, branched several times with secondary branching relatively common, each branch terminating in either a singular or branched inflorescence. Stem leaves fleshy, narrowly linear to narrowly elliptic, 2.5-15 mm long, 1.2-3.2 mm wide, shortly mucronate, alternate to somewhat scattered, ascending, often curved inwards, usually terete to compressed in mid cross-section, colour as above. Inflorescences 4-12 flowered; axis 8-140 mm long, generally fomiing a loose cyme or panicle, the inflorescence tip unfurling with maturity, bare except for several ± scarious bracts. Inflorescence axis bracts appressed, narrowly triangular, occasionally broader, 0.75-1.1 mmlong, 0.4-0.55 mm wide, apex acuminate, scattered, but often located near bases of individual flower pedicels, withering when flowers reach mature fruit stage. Pedicels 0.1-0.2 mm long and erect, to 0.4 mm long in fruit. Flowers 1.5-3.0 mm diam. Sepals thick, broadly ovate to orbicular, 0.9-1.3 mm long, 1-1.2 mm wide, fused over a short distance to the apex of the pedicel, with a moderately prominent midvein and two lateral veins and extensive reticulation, upper edge distinctly hyaline. Petals 5, usually pale pink or white, narrowly elliptic to ovate, occasionally broader, 1.1-1.6 mm long, 0.6-1 mm wide, free to base. Stamens 3-5 in one series; filaments flattened, very broad at the base, 0.4-0.6 mm long, shortly adnate to the base of adjoining petals and attached to the top of basal ring beneath ovary; anthers orbicular to broadly elliptic in outline, 0.25-0.35 mm long, 0.25-0.35 mm wide, versatile, extrorse, dehiscing longitudinally. Ovary obovoid, 0.55-0.7 mm diam., brown. Stigmas 3, squat, linear to narrowly triangular, lengthening and spreading with maturity, 0.15-0.3 mm long, free to base, with a dense covering of stigma trichomes. Capsule ellipsoid to globular, coriaceous to crustose, surface distinctly verrucose-colliculate, 1.3-1.7 mm long, 1.1- 1.3 mm wide, the apex relatively obtuse, usually much longer than the sepals, capsules indehiscent and when mature also deciduous with sepals attached; valves 3, sometimes very shortly split at the apex, probably splitting fully with age. Seeds one per capsule, greyish to black, obovoid to globular with a relatively large strophiole, 1-1.05 mm long, 0.8-0.9 mm wide, 0.65-0.7 mm thick, surface minutely and parallelly rugose. (Figures 1, 2). Diagnostic features. Calandrinia monosperma may be uniquely diagnosed within the genus by its distinctive hard, one-seeded, indehiscent capsules with a verrucate-colliculate surface that are unlike those of any other species of Calandrinia. Other specimens examined.WF'FIFRISAIJ'FIBJ kLIA: Mine Flats, Paraburdoo, 23 Sep. \919, K.J. Atkins 584 (PERTH); F ish holes, 36.5 km from Doolgunna Homestead on the northern boundary of the station, 9 Aug. 2006, G. Byrne 2308 (PERTH); Site FMS 18, 17 km SW of Fake Mason Homestead, 21 Sep. F. J. Obbens, Calandrinia monosperma and C. uncinella (Montiaceae), two new species from WA 239 Figure 1. Calandrinia monosperma capsules with sepals and portion of pedicel attached (i.e. dispersal unit) (A) and capsules with sepals removed (B). Scale bars = 1 mm. Voucher: F. Obbens & B. Bromilow FO 32/04. 2004, DJ. Edinger & G. Marsh DJE 4989 (PERTH); Mount Narryer, SW of main peak, 10 Aug. 1997, A.S. George 17374b (PERTH); Booylgoo Range, survey site BOOY07, on Booylgoo Spring Station approximately 4.1 km SE of Number 1 Bore and 2.8 km north of Phils Bore. Approximately 62.6 km east north-east of Sandstone, 3 Sep. 2006, A. Markey & S. Dillon 4778 (PERTH); Jack Hills, survey site JACK35, located at the end of the main range approximately 3 km S of spot-height 482 m, and approximately 30 km from the junction of Berringarra - Cue Rd and the main track running adjacent and parallel N of the Jack Hills Range, 29 Aug. 2005, R. Meissner & Y. Caruso 705 (PERTH); MtNairn, Milly Milly Station, survey site MTNN04, c. 2.7 km NE of spot-height 371 m and 4.1 km E of Tardy Well, 15 Aug. 2007, R. Meissner & G. Owen 1690 (PERTH); Mount Barloweerie survey site MTBW04, located on Mount Barloweerie (BIA Aboriginal Reserve), c. 3.8 km SE of Burra Burra Well and 7.5 km SW of Pia Well. Eocated c. 134 km NNW of Yalgoo, 25 Aug. 2008, R. Meissner & J. Wright 2416 (PERTH); Site 702, Weld Range, 19 July 2006, J. Naaykens JN 702 07 (PERTH); Area c. 6 to 7 km E of Channar East Mine (ESE of Paraburdoo) and c. 0.5 km downslope from Pilbara Biological Survey site TCMBC05, 7 June 2006, F. Obbens FO 19/06 (PERTH); Area along track c. 50 km directly ESE of Paraburdoo and c. 4 km S of Turee Creek East Branch. Also located nearby Pilbara Biological Survey site TCMBC12, 8 June 2006, F. Obbens FO 22/06 (PERTH); 5.3 km N along the Cobra-Dairy Creek Rd from junction with the Carnarvon - Mullewa Rd (W side), 9 Aug. 2015, A Obbens 18/15 (PERTH); 6.4 km S along Carnarvon - Mullewa Rd from junction with the Byro - Beringarra Rd. On W side of road c. 250 to 300 m away, 10 Aug. 2015, A Obbens 19/15 (PERTH); c. 40-45 km N of Murchison settlement on Carnarvon - Mullewa Rd (before Curbur Homestead turnoff), 30 Aug. 2018, A Obbens FO 01/18 (PERTH); Site number: 648, 6.8 km NW of Mount Hilditch, Hamersley Ranges, Fortescue Botanical District, 2 June 1997, M.E. Trudgen 18309 (PERTH); Foot of The Pink Hills, 17 Aug. 1973, A. fP/fOverW 1050 (PERTH). Phenology. The main flowering and fruiting period for C. monosperma is from early June through to late September. This extensive period relates to the species’ wide distribution, hence flowering and fruiting occurs first in northern populations and later at more southern locations. 240 Nuytsia Vol. 30 (2019) Figure 2. Calandrinia monosperma seeds with dorsal and plan views. Scale bar = 0.2 mm. Voucher: F. Obbens & B. Bromilow FO 32/04. Distribution and habitat. Calandrinia monosperma occurs over a relatively wide area, from southern parts of the IBRA Pilbara subregion to the Murchison subregion, and from Gascoyne Junction- Murchison settlement areas in the west and eastwards to Wiluna-Leonora areas (Figure 3). This species is found growing in soils described as mainly red to orange-brown, clayey or sandy loams, often pebbly or rocky and frequently ironstone on various landforms that include gibber or rocl6{20Q2)\Jonesiopsisincensa(Ro^^QX &A.P.Br.)D.L.Jones& M.A.Clem., Orchadian 14(4): 182(2002). Typus: Chiddarcooping Hill Nature Reserve, near southern boundary, 30°54’S, 11 SATE, Western Australia, 17 August 1988, A. Brown 829 {holo\ PERTH 01708112; iso\ AD 156477, CANB 556721, K, MEE 2279957, NSW 520141). CaladeniavulgataiAo^^QxScA.VBx.^Nuytsia 14(1/2): 280(2001),/?ro/?(3!rreastotype. Typus: Kalbarri National Park, 17 km south of Eurardy Homestead, 13.1 km north of Murchison River Bridge, 500 m south-west of Highway, 27°43’S, 114°40E, Western Australia, 23 August 1983, S.D. Hopper 3330 (holo: PERTH 00273600; iso: AD 168970, CANB 598564.1, K). Caladenia varians Hopper & A.PBr., sp. nov. Typus: Coorow Greenhead Road, 25 km west of Midlands Road, south-west of Coorow, Western Australia, 4 September 2010, G. Brockman GBB 2621 {holo: PERTH 08420068; iso: CBG). Caladenia vulgata Hopper & A.P.Br., Nuytsia 14(1/2): 280 (2001),pro parte not as to type. Caladenia variansdo^^QX ScAiBBx. subsp. varians, inN. Hoffman&A. Brown, Orchids of South-West Australia, 2nd edn. p. 33 (1992), nom. nud. Illustrations. N. Hoffman & A. Brown, Orchids of South-West Australia, 2nd edn, p. 33 [as Caladenia varians subsp. varians] (1992); N. Hoffman & A. Brown, Orchids ofS-W. Austral., rev. 2nd edn with suppL, p. 33 (1998) and 3rd edn, p. 37 (2011); A. Brown, P. Dundas, K. Dixon & S. Hopper, Orchids of W. Austral., p. 36, Figure A (2008); G. Backhouse, Spider-orchids - the Genus Caladenia and its Relatives in Austral., p. 258-259 (2011) and rev. 2nd edn, p. 276-277 (2018); A. Brown, K. Dixon, C. French & G. Brockman, Field Guide to the Orchids ofW. Austral, p. 72 (2013); N. Hoffman, A. Brown & J. Brown, Orchids ofS-W. Austral, 4th edn. p. 38 (2019) [all as C. vulgata]. Plants solitary or in small clumps. Leaf 50-220 mm long, 3-5 mm wide, linear, erect, incurved in cross section, pale green, the basal 1/3 usually irregularly blotched with red-purple. Scape 150- 300 mm tall. Flowers 1 to 3, 70-100 mm across, cream, with pale maroon lines, spots and blotches; floral odour faintly sweet to putrid. Petals and sepals linear-lanceolate, scarcely glandular-hirsute on the outside and glabrous on the inside in the basal 1/5 to 1/4 then abruptly narrowing to a red-black, densely glandular-hairy, long-acuminate filamentous apex lacking a swollen osmophore. Dorsal sepal 45-100 mm long, 2-3 mm wide, erect and slightly incurved. Lateral sepals 50-110 mm long, 2-A mm wide, spreading obliquely downwards near the base and pendulous towards the apex. Petals 45-100 mm long, 2-3 mm wide, spreading horizontally near the base and pendulous towards the apex. Labellum obscurely 3-lobed, cream with pale to deep red stripes, spots and blotches, stiflly articulated on a claw 1-1.5 mm wide; lamina 10-15 mm long, 7-11 mm wide, narrowly triangular to triangular (rarely rhomboidal) in outline, erect with entire margins in the basal 1/3, nearly horizontal in middle 1/3 and apical 1/3 prominently recurved; lateral lobes with broadly truncate to triangular forwardly uncinate cream to red-marked marginal calli which are decrescent towards the mid lobe; lamina calli cream, often with pale red markings, dull on top, broadly to narrowly anvil-shaped, the longest c. 1 mm tall, in 8-12 pairs in two rows extending over about 1/2 the length of the labellum and slightly decrescent towards the apex. Column 8-12 mm long, 4-6 mm wide, narrowly-winged, opaque cream with pale red stripes and blotches, sparsely hirsute with short glandular hairs on outer surface. Anther 2-3 mm long, 2-3 mm wide, greenish yellow. Pollinia >2 mm long, kidney-shaped. S.D. Hopper & A.P. Brown, Update on Paracaleana and a name in Caladenia (Orchidaceae) 283 flat, yellow, mealy. Stigma 2-2.5 mm long, 2-2.5 mm wide. Capsule not seen. (Figure 1) Diagnostic features. Caladenia varians may be distinguished from all other members of the genus by the following combination of characters: Flowers cream, with pale maroon lines, spots and blotches; Petals and sepals with a red-black, densely glandular-hairy, long-acuminate filamentous apex lacking a swollen osmophore: Labellum lateral lobes with broadly truncate to triangular forwardly uncinate marginal calli; lamina calli broadly to narrowly anvil-shaped in 8-12 pairs in two rows extending over about 1/2 the length of the labellum. Selected specimens examined. WESTERN AUSTRALIA: Coolinup Rd, 3 km S of Kettles Road. Track follow power line to W, 23 Sep. 2004, G. Brockman 1417 (PERTH); 500m SE of Beaumont Wheat facility; Beaumont Nature Reserve, 23 Sep. 2004, G. Brockman GBB 1477 (PERTH); Reserve 10147, Robinson Rd,Woodanifling, 8 Oct. 2014, G. Byrne 5274 (PERTH); 16 miles NW ofBadgingarra, 1 Sep. 1966, A.S George 7805 (PERTH); 22 km SSE of Kalbarri, 45 km WNW of Binnu, 24 Aug. 1983, S.D. Hopper 3342 (PERTH); 26 km ENE of Wilga siding, 6 Oct. 1983, S.D. Hopper 3477 (PERTH); 300 metres E of Mandurah-Fremantle Rd on Paganoni Rd, 12 km NNE of Mandurah, 12 Sep. 1984, S.D. Hopper 4136 (AD, CANB, PERTH); Yalgorup National Park, 4.1 km W of Mandurah - Bunbury Rd on Preston Beach Rd, 12 Sep. 1984, YD. Hopper 4145 (AD, CANB, K, PERTH); Yalgorup National Park, 1.2 km W of Mandurah - Bunbury Rd on Preston Beach Rd, 12 Sep. 1984, S.D. Hopper 4149 (CANB, PERTH); 26 kmNWof Kojonup, intersection of Collie - Changerup and Stirlings Rd, 11 Sep. 1985, S.D. Hopper 4543 (PERTH); Dragon Rock Nature Reserve (36128), 31 km N of Newdegate, Figure 1. Caladenia varians. K-lowers showing their cream and pale maroon colouration and pendulous petals and lateral sepals with densely glandular-hairy, long-acuminate filamentous apexes; B - labellum, showing the broadly truncate to triangular forwardly uncinate marginal calli and broadly to narrowly anvil-shaped lamina calli. Photographs by A.P. Brown. 284 Nuytsia Vol. 30 (2019) 13 Sep. 1985, S.D. Hopper 4567 (CANB, PERTH); Kalbarri Road, c. 9 km WSW of Murchison House Station turnoff, 8 Aug. 1986, S.D. Hopper 5X1% (PERTH); Great Northern Hwy, 1 kmNNE ofWubin, 23 Aug. 1988, S.D. Hopper 6504 C (PERTH); SE foot and W foot of Totadgin Rock, c. 47 km NE of Kellerberrin, 14 Sep. 1988, S.D. Hopper 6610 (PERTH); c. 12 km W of Z Bend Gorge, Kalbarri National Park, and 10 km SSW of T- junction on road to Z Bend Gorge, 8 Aug. 1990, S.D. Hopper 7815 (PERTH); Bullsbrook Nature Reserve, 1 kmN of Pearce, 9 Aug. 1992, G.J. Keighery 13514 (PERTH); Pikaring West Nature Reserve, Old Beverley Rd; ca 25 km E Quairading, 18 Aug. 1998, G.J. Keighery 16388 (PERTH); Bridge crosses salty creekline on the Coorow-Greenhead Rd 7.6 km E of the junction of Carger Rd and the Coorow-Greenhead Rd, 29 Aug. 2014, R. Phillips 0269 (PERTH); Peak Charles car park area, 6 Aug. 1978, D.R. Voigt 46pp (PERTH). Phenology. Flowering from August to October. Fruiting from October to November. Distribution and habitat. Widespread throughout the south-west from north of Kalbarri to Augusta and eastwards to Condingup. Favours sandy soils in a range of communities including coastal heath, inland mallee heath, woodlands and forests. Conservation status. Not considered rare or under immediate threat. Etymology. From the Eatin varians (varying), in reference to the variable morphology of the species over its very large geographic range. Affinities. Caladenia varians is most similar to C. pendens Hopper & A.P.Br. from which it may be distinguished by its shorter petals 45-100 x 2-3 mm {cf. 60-120 x 3-5 mm in C. pendens), generally smaller labellum 10-15 x 7-11 mm {cf. 11-19 x 9-12 mm in C. pendens) and smaller column 8-12 x 4-6 mm {cf. 10-15 x 5-7 mm in C. pendens). It is also similar to C. exilis Hopper & A.P.Br. from which it may be distinguished by its larger labellum 10-15 x 7-11 mm cf (8-12 x 5-7 mm in C. exilis). Caladenia varians may be found growing with C. nobilis Hopper & A.P.Br., which has much larger flowers with a broader labellum 12-25 x 12-16 mm and larger column 13-18 x 5-10 mm. It may also be found with C. hiemalis Hopper & A.P.Br., which generally has shorter petals 35-60 mm long and an earlier flowering season between June and August, and with C. meridionalis Hopper & A.P.Br., which also flowers earlier (June-August) and has stiflly held petals and sepals with dark glandular apices. Caladenia meridionalis is also confined to the south coast whereas C. varians is widespread from Kalbarri to Condingup. Northern populations of C. varians may grow near to C. incensum but the latter species differs in its broader leaf 4-15 mm across, broader tepals 3-6 mm across, short squat calli and its preference for heavier (clay-loam) soils. To the north-west of Northampton, C. varians grows with C. elegans Hopper & A.P.Br. which differs in its pale yellow colouration, thicker more squat calli and its preference for winter-wet clay soils. Notes. Caladenia varians occasionally hybridises with C. nobilis on the Swan Coastal Plain and with C. wanosaA. S .George in Kalbarri National Park. Rare hybrids of C. varians and C. drakeoides Hopper & A.P.Br. found near Pithara are formally named Caladenia xhopperi J.M.H.Shaw (Shaw 2014). Acknowledgements S.D. Hopper & A.P. Brown, Update on Paracaleana and a name in Caladenia (Orchidaceae) 285 We are grateful to many colleagues who have shared their time in conducting fieldwork and have assisted us by making additional collections. We would like to thank the Curator and staff of PERTH for access to specimens. Kelly Shepherd provided preliminary editorial guidance that materially improved the paper. References Brown, A.P. & Brockman, G. (2007). Caladeniapetrensis and C. saxicola (Orchidaceae), two new ironstone endemics from south-west Western Australia. Nuytsia 17: 73-79. Brown, A.P. & Brockman, G. (2015). New taxa of Caladenia (Orchidaceae) from south-west Western Australia. Nuytsia 25: 45-123. Brown, A.P. & Brockman, G. (2019). Paracaleana ferricola (Orchidaceae), a rare new species from south-west Western Australia. Nuytsia 30: 287-289. Brundrett, M. (2014). Identification and ecology ofSouthwestAustralian Orchids. (Western Australian Naturalists’ Club Inc.: Perth.) Clements, M.A., Howard, C.G. and Miller, J.T. (2015). Caladenia revisited: results of molecular phylogenetic analyses of Caladeniinae plastid and nuclear loci. American Journal of Botany 102: 581-597. Hopper, S.D. (2009). Taxonomic turmoil down-under: recent developments in Australian orchid systematics. Annals of Botany 104:447-455. Hopper, S.D. & Brown, A.P. (2001). Contributions to Western Australian orchidology: 2. New taxa and circumscriptions in Caladenia (Spider, Fairy and Dragon Orchids of Western Australia). Nuytsia 14: 21-301. Hopper, S.D. & Brown, A.P. (2004). Robert Brown’s Caladenia revisited, including a revision of its sister genera Cyanicula, Ericksonella and Pheladenia (Caladeniinae: Orchidaceae). Australian Systematic Botany 17: 171-240. Hopper, S.D. & Brown, A.P. (2006). Australia’s wasp-pollinated flying duck orchids revised {Paracaleana. Orchidaceae). Australian Systematic Botany 19: 211-244. Jones, D.L. & Clements, M.A. (2001). Subtribe Drakaeinae. In. Pridgeon, A.M., Cribb, P.J., Chase, M.W. & Rasmussen, F.N. (ed.) Genera Orchidacearum, 2. Orchidoideae (Part 1). pp. 134 - 155. (Oxford University Press: Oxford, UK.) Kores, P.J., Molvray, M., Weston, P.W., Hopper, S.D., Brown, A.P, Cameron, K.M. & Chase, M.W. (2001). A phylogenetic analysis of Diurideae (Orchidaceae) based on plastid DNA sequence data. American Journal of Botany 88: 1903-1914. JMcNeil, J., Barrie, F.R., Buck, W.R., Demoulin, V., Greuter, W., Hawksworth, D.L., Herendeen, PS., Knapp, S., Marhold, K., Prado, J., Prud’homme van Reine, W.F, Smith, G.F, Wiersema, J.H. & Turland, N.J. (2012). International Code of Nomenclature for Algae, Fungi, and Plants (Melbourne Code). Regnum Vegetabile 154. (Koeltz Scientific Books.) Miller, J.T., & Clements, M.A. (2014). Molecular phylogenetic analyses of Drakaeinae: Diurideae (Orchidaceae) based on DNA sequences of the internal transcribed spacer region. Australian Systematic Botany 27: 3-22. Shaw, J.M.H. (2014). The Orchid Review 122 (1305, Suppl.): 16. 286 Nuytsia Vol. 30 (2019) Nuytsia The journal of the Western Australian Herbarium 30:287-289 Published online 10 December 2019 SHORT COMMUNICATION Paracaleana ferricola (Orchidaceae), a new, rare species from the south¬ west of Western Australia Paracaleana ferricola A.P.Br. & G.Brockman, sp. nov. Type : Canning Mills, Western Australia [precise locality withheld for conservation reasons], 18 November 2017, G. Brockman GBB 3571 {holo\ PERTH 08961646; iso\ CANB). Paracaleana sp. Laterite (G. Brockman GBB 3571), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 23 July 2018]. Illustration. A.P Brown, K.W Dixon, C.J. French & G. Brockman, Field Guide to the Orchids of W Austral, p. 305 (2013), as Paracaleana sp. ‘Darling Range’. An erect perennial geophytic herb to 10 cm high. Plants solitary, more rarely in small groups. Tuber oblong, 6-11 mm long, 3-6 mm wide, white, annually replaced with a new tuber produced at the end of an elongate descending side-dropper (root-like stolon). Leaf solitary, basal, erect to spreading, withered or more rarely fresh at anthesis; lamina when fresh 10-20 mm long, 5-8 mm wide, glabrous, thin, narrowly elliptic to ovate, flattened in TS, dark maroon above and below, apex acute to obtuse. Scape 6-10 cm tall, wiry, dull brownish maroon and greenish yellow, with floral bract(s) erect, sheathing, foliaceous, acuminate; pedicels 5-15 mm long. Flowers solitary, rarely paired, nonresupinate, inverted, glabrous, greenish yellow and pale maroon with darker maroon blotches and markings; floral odour undetectable to humans but known to attract male thynnid wasps. Sepals and petals similar in shape, narrowly linear, margins revolute, apices narrowly clavate to acute. Dorsal sepal 7-10 mm long, 1-1.5 mm wide, splayed downwards, appressed against the column and distinctly elbowed in terminal 1/4 to 1/3, apex narrowly clavate. Lateral sepals 7-10 mm long, 1 mm wide, splayed downwards, usually away from the column wings, distinctly elbowed in terminal 1/4 to 1/3, apex not clavate. Petals narrower than sepals, 6-9 mm long, 0.5 mm wide, splayed downwards, appressed to column wings, shallowly curved, apex not clavate. Labellum motile, stiffly sprung on a long broadly rhomboidal claw 3-5 mm long, 1.5-2 mm wide, attached to the front of the extended column foot, the claw upcurved when set, recurved when sprung, terminating in a triangular apex extending backwards off the base of the labellum lamina; lamina 8-10 mm long, 2.5-3 mm wide, undivided, narrowly lageniform, entire, flattened or sometimes with a slight hump at 2/3 its length with a shortly projecting narrowly triangular neck-like narrowly obtuse apex; lamina calli sessile, irregularly hemispheric, smooth, black, glossy, confined to the apical 1/3 to 1/2. Column 6-10 mm long, downcurved, 2-winged; the wings 3^ mm wide, broadly rectangular, incurved, somewhat opaque, greenish to greenish maroon with darker maroon blotches and markings. Anther 1.5-2 mm long, 1.5-2 mm wide, yellowish green, obtuse, not terminating in a definite point. Pollinia 1.5-2 mm long, four, lamellate, granular, yellow. Stigma ovate, 1.5-2 mm long, 1.5-2 mm wide, dull yellow. Ovary narrowly pyramidal, greenish yellow with brownish maroon longitudinal stripes. Capsule not seen. (Figure 1) Diagnostic features. Paracaleana ferricola may be distinguished from all other members of the genus by the following combination of characters: leaf elliptic to ovate, dark maroon above and below, apex © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 288 Nuytsia Vol. 30 (2019) F igure 1. Paracaleanaferricola. A - flowering plants in situ at the type locality showing the sometimes paired flowers, thin wiry stems and basal leaves which are often withered at anthesis; B - flower, showing the narrow sepals and petals, broadly winged column and motile labellum with calli confined to the apical 1/3 to 1/2; C - habitat showing the iron-rich lateritic soils occupied by the species and associated vegetation including Banksia sessilis and Xanthorrhoea preissii. Photographs by A.P. Brown. acute to obtuse, withered or rarely fresh at anthesis; labellum lamina entire, flattened, sometimes with a slight hump at 2/3 its length, with calli conflned to the apical 1/3 to 1/2. Other specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 6 Dec. 2005, G. Brockman 1694 (PERTH); 30 Oct. 2006, G Brockman 1986 (PERTH); 5 Nov. 2002, F.&J. Hort 1892 (PERTH); 14 Nov. 2002, F. & J. Hort 1908 & G Brockman (PERTH). Phenology. Flowering from late October to early December. Fruiting December. Distribution and habitat. Found over a small geographic range east of Perth, growing in lateritic soils containing an unusually high amount of Haematite. The habitat is open Allocasuarina fraseriana, Corymbia calophylla. Eucalyptus marginata woodland over Banksia sessilis and Xanthorrhoea preissii (Figure 1). Conservation status. Currently listed as Priority Two under Conservation Codes for Western Australian Flora (Smith & Jones 2018), under the name P. sp. Eaterite (G. Brockman GBB 3571). The species is known from two small populations 1.8 km apart in Korung National Park, these together comprising just 30 mature plants in an area threatened by altered Are regimes, dieback and habitat damage by off-road vehicles. A.P. Brown & G. Brockman, Paracaleana ferricola (Orchidaceae), a new species from WA 289 Etymology. The epithet ferricola is derived from the Latin ferrum (iron) and the suffix -cola (dweller or inhabitant) in reference to the iron-rich lateritic soils occupied by this species. Affinities. Paracaleana ferricola is most similar to P. disjuncta Hopper & A.P. Br. and like that species has a late flowering period, a leaf that is often withered at anthesis and a small greenish yellow and maroon flower. It is distinguished, however, by its smaller flower with petals and sepals 6-10 mm long {cf 8-12 mm long for P. disjuncta), sometimes slightly humped, narrower labellum 2.5-3 mm wide {cf labellum lacking a hump and 3-5 mm wide for P. disjuncta), labellum lamina calli confined to the apical 1/3 to 1/2 {cf labellum lamina calli confined to the apical 1/3 for P. disjuncta) and narrower column wings, 3-A mm wide {cf column wings 4-6 mm wide forP. disjuncta). FurthermoreP ferricola occurs some 250 km to the north of the nearest known population of P. disjuncta and occupies lateritic soils the sandy soils favoured by that species. At the type location P ferricola grows with P. nigrita (Lindl.) Blaxell and P. brockmanii Hopper & A.P.Br., but flowers over a month after P. nigrita has finished flowering and is in late flower when P brockmanii begins flowering. It is distinguished from P. nigrita by its labellum either lacking a hump or with a slight hump at 2/3 its length {cf labellum prominently humped at 1/2 its length for P. nigrita) and its lamina calli confined to the apical 1/3 to 1/2 the length of the labellum {cf lamina calli confined to the apical 2/3 the length of the labellum for P. nigrita). From P. brockmanii, P. ferricola is distinguished by its smaller flowers with petals and sepals 6-10 mm long {cf 10-15 mm long for P. brockmanii), smaller, narrower labellum 8-10 x 2.5-3 mm {cf 9-12 x 4-7 mm for P. brockmanii) and shorter column 6-10 mm long {cf 10-12 mm long forP. brockmanii). Acknowledgements The authors would like to thank Fred and Jean Hort and Joflf and Joan Start who have helped in locating plants in the held, the Curator and staff at the Western Australian Herbarium for access to specimens and Terry Macfarlane for useful comments and edits on the manuscript. References Department of the Environment (2017). Australia’s bioregions (IBRA), IBRA7, Commonwealth of Australia. https://www. environment.gov.au/lancl/nrs/science/ibra#ibra [accessed 11 December 2018], Smith, M.G. & Jones, A. (2018). Threatenedand Priority Floralist 5 December 2018. Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 11 December 2018], Andrew P. Brown^ and Garry Brockman Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 ^Corresponding author, email: andrewbrown3@optusnet.com.au 290 Nuytsia Vol. 30 (2019) Nuytsia The journal of the Western Australian Herbarium 30:291-308 Published online 10 December 2019 The Hibbertia polystachya-H, spicata (Dilleniaceae) species group in Western Australia Kevin R. Thiele School of Biological Sciences, University of Western Australia 35 Stirling Hwy, Crawley WA 6009 Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 Email; kevin.thiele@uwa.edu.au Abstract Thiele, K.R. The Hibbertiapolystachya-H. spicata (Dilleniaceae) species group in Western Australia. Nuytsia 30: 291-308 (2019). Hibbertia spicata F.Muell. and H. polystachya Benth. are widespread in south-western Western Australia, where they are unique in Hibbertia Andrews in having flowers in a cincinnus rather than solitary. The boundary between these two species has been problematic, as they mostly dififer in the composition and density of indumentum on their sepals and leaves. Close study has revealed that//; polystachya is widespread andtaxonomically relatively straightforward, while//, spicata sens. lat. comprises six distinctive, readily resolvable species (including H. leptotheca (J.R.Wheeler) K.R.Thiele comb, etstat. nov., previously H. spicata subsp. leptotheca J.R.Wheeler) differing in leaf and sepal indumentum, number of flowers in the cincinnus, floral bract shapes, stamen number, and number and arrangement of staminodes. New species described here are H. capensis K.R.Thiele sp. nov., H. asterella K.R.Thiele sp. nov. and H. subglabra K.R.Thiele sp. nov. The new name H. prolata K.R. Thiele is provided for a widespread taxon based on Hemistephus linearis J.Drumm. ex Harv. With these species removed, H. spicata is recircumscribed and is now more restricted in distribution. Introduction Hibbertia spicata F.Muell. was described by Mueller (1860) based on a specimen collected by Walcott and A.F. Oldfield, probably in 1859, at Port Gregory (between Geraldton and Kalbarri). In the protologue, Mueller noted its close aflinity with Hemistephus linearis J.Drumm. ex Harv., described by Harvey (1855) based on a Drummond specimen from ‘Northern Districts’. Bentham (1863) synonymised the latter name under the former (the combination in Hibbertia Andxovts being unavailable due to the epithet being preoccupied by H. linearis R.Br. ex DC.). Bentham (1863) also described Hibbertia polystachya Benth. based on two specimens, a Drummond collection from ‘Swan River’ and an Oldfield collection from the ‘Blackwood River’. He included both species in his sect. Hemipleurandra Benth., noting that unlike other species in the section, H. spicata and H. polystachya have flowers borne in a ‘ 1-sided spike’ rather than singly. He gave as the key difference glabrous leaves in H. spicata compared with hirsute leaves (and sepals) in H. polystachya, noting also that the former has staminodes completing the ring of stamens while the latter has no or few staminodes (sometimes with only a single one behind the stamens). © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 292 Nuytsia Vol. 30 (2019) Wheeler (1984,1987,2004) commented on the close relationship between these two species and the difficulty of separating them at times. Wheeler (2004) gave the key difference as the sepals, bracts and young leaves being glabrous or with minute stellate and hooked hairs in H. spicata c.f having fairly long, coarse, simple hairs (sometimes also with underlying minute stellate hairs) in H. polystachya. She considered (Wheeler 1984) that the difference in staminodes provided by Bentham (1863) was unreliable, with some specimens from each species having identical staminode arrangements. She further noted that (1) some specimens appear intermediate between the two species, speculating that closer study might reveal that H. polystachya and H. spicata are conspecific, and conversely that (2) H. spicata is morphologically variable within its range, with the implication that it may contain further taxa. Wheeler (1984) added a third taxon to the group when she described H. spicata subsp. leptotheca J.R. Wheeler for specimens occurring along the coastal plain from Lancelin, north of Perth, southwards to Yalgorup National Park. These differ from typical subsp. spicata in having longer, more slender anthers, fewer (or no) staminodes, and outer sepals that are glabrous or almost so. She noted that subsp. leptotheca is geographically and ecologically separated in the Perth region from subsp. spicata, the former occurring on coastal limestones and the latter being absent from the coastal plain in that region (although elsewhere occurring on coastal limestones e.g. at North West Cape). Hibbertiapolystachya and if. spicata are striking amongst Hibbertia species in south-west Western Australia in having flowers borne in cincinni (often described as ‘one-sided spikes’; Figure 1), all other species in the area having sessile or pedicellate flowers borne singly and terminating shoots (sometimes appearing axillary to leaves when terminating axillary short-shoots or following overgrowth of an axillary shoot arising below the flower). Two other groups of Hibbertia also have cincinnate inflorescences: a northern Australian species group including H. muelleri Benth. and H. candicans (Hook.f) Benth., and a New Caledonian radiation including H. pancheri (Brongn. & Gris) Briq. and 77 vieillardii (Brongn. & Gris) Gilg. Despite striking similarities in their inflorescences, the molecular phylogeny of Horn (2005) resolves these as separate clades, with H. spicata in a clade with other (single-flowered) species from south-western Australia such as H. acerosa (R.Br. ex DC.) Benth. {H. polystachya was not included in the phylogeny). Horn (2005) analysed inflorescence structure and floral development in//, polystachya and concluded that the cincinnus is a highly sylleptic sympodium; that is, each flower terminates its axis, with subsequent flowers ‘higher up’ in the inflorescence derived from axes produced from the axil of the subtending bract of the terminal flower and with the bract carried sylleptically up the axis so that it appears to subtend the next-distal flower. Usually only one flower is open in each cincinnus at any given time. This study assessed morphological variation in H. polystachya and H. spicata to determine whether the two taxa should be retained as separate or whether the variation noted by Wheeler (1984) indicates that further taxa should be recognised. Significant variation was found, in the indumentum of young stems, leaves and sepals, in the number of flowers and shape and arrangement of bracts in the cincinnus, in flower size, in the number of stamens, and in the number, shape and arrangement of staminodes. The morphological variation is geographically and ecologically correlated and consistent with the recognition of four new species and the raising of H. spicata subsp. leptotheca to species rank. The group thus comprises seven distinct and readily recognisable species. K.R. Thiele, Hibbertiapolystachya-H. spicata (Dilleniaceae) species group in WA 293 Figure 1. Flowering branches of two species in the Hibbertia spicata-H. polystachya group. A- H. prolata (Zig Zag Road, Gooseberry Hill, 28 Sep. 2019); B -Ff. asterella {K.R. Thiele 5590, 2 Oct. 2019). Photos by K.Thiele. Methods All specimens at PERTH were examined. Types of Hibbertia spicata, H polystachya and Hemistephus linearis were viewed at MEL; other type specimens were viewed on JSTOR Global Plants (https:// plants.jstor.org/). Leaf and sepal measurements are based on dried specimens; petal, stamen and style measurements are based on herbarium material rehydrated in hot water with a little detergent. Maps are based on all specimens held at PERTH and are drawn using IBRA v. 7 (Department of the Environment 2013) bioregion and subregion boundaries. Key to species in the Hibbertia spicata-H. polystachya species group 1. Outer sepals moderately to densely stellate-hairy, with no or few simple hairs 2. Whole plant (young stems, leaves and sepals) densely stellate-pubescent; stamens 10-12; most staminodes opposite the stamens. 1. H. asterella 2: Young stems and leaves ±glabrous; stamens 12-16; most staminodes behind the stamens. 2. H. capensis 1: Outer sepals glabrous to pilose, mostly with simple or hooked hairs (stellate hairs, when present, a minor component of the indumentum) 2. Abaxial leaf lamina and midrib glabrous. 3. H. subglabra 2: Abaxial leaf lamina where exposed (and where hidden by the recurved margins) densely stellate-pubescent, the midrib often glabrous 294 Nuytsia Vol. 30 (2019) 3. Flowers (6-)8-14 per cincinnus, overlapping; cincinnus bracts distinctly heteromorphic, each flower (except the lowermost) subtended by a ±linear and a broadly ovate to triangular bract. 4. H. spicata 3: Flowers (l)2-6(7) per cincinnus, the lowermost at least well-separated; cincinnus bracts not distinctly heteromorphic, each flower (except the lowermost) subtended by two ±linear to ovate bracts dilFering mainly in length 4. Adaxial leaf laminas and outer sepals quite glabrous.5. H. leptotheca 4: Adaxial leaf laminas and/or outer sepals hairy 5. Adaxial leaf laminas ±glabrous; sepals mostly with hooked hairs. 6. H. prolata 5: Adaxial leaf laminas pubescent to pilose with simple hairs; sepals with a mix of simple, stellate and hooked hairs. 7. H. polystachya Taxonomy 1. Hibbertia asterella K.R.Thiele, sp. nov. Type : Bokal Road North, 1.4 km from Boyup Brook-Arthur River Road, Western Australia, 2 October 2019, K.R. Thiele 5590 (holo\ PERTH 09085122; iso\ AD, CANB, MEL) Prostrate, compact or spreading shrubs to 0.3(-0.7) m high, multi-stemmed at base and resprouting from the rootstock after fire; young stems moderately to densely stellate-pubescent; older stems with smooth to scabrid, papery, dull or silvery bark decorticating in flakes and strips. Leaves spreading- erect, scattered, linear, (10-) 12-25 mm long, 1-2 mm wide; margins revolute but the abaxial lamina usually exposed; adaxial lamina coarsely stellate-pubescent with tubercle-based hairs (rarely with scattered simple hairs amongst the stellate ones); abaxial lamina finely and densely stellate- pubescent, the midrib with indumentum as for the adaxial surface; apex obtuse and recurved-apiculate. Inflorescences comprising 2^-flowered cincinni, the lowermost flowers usually well-separated; cincinna axis with indumentum as for the stems; bracts ±homomorphic, narrowly ovate to linear, the lowermost 3-6 mm long, 0.8-2 mm wide, often leaf-like in having recurved margins, a prominent midrib, and indumentum as for the leaves. Sepals ovate, 5-6 mm long, abaxially densely tuberculate- stellate; midribs prominent; outer sepals acute; inner sepals similar to the outer but smaller. Petals yellow, broadly obovate, 7-10 mm long, shallowly to deeply emarginate. Stamens 10-12, all on one side of the gynoecium, shortly fused by their filaments; anthers oblong, 1.5-2 mm long, dehiscing by introrse, longitudinal slits. Staminodes 10-16, a few lateral to but most opposite the stamens. Carpels 2; ovaries rather rectangular-cuboid, densely pubescent; styles excentric from near the carpel apex, istraight then slightly curved at the apex, 1-1.5 mm long. Ovules 2 per carpel. Fruiting carpels and seeds not seen. (Figure lA) Diagnostic features. May be distinguished from all other species in the Hibbertia spicata species group by the densely and coarsely stellate-pubescent leaves, cincinnus bracts and sepals. Other specimens examined (all PERTH). WESTERN AUSTRALIA: Wingebellup Rd, E of Unicup Rd and N of Kulanilup Reserve, 15 Nov. 2017, G. Byrne 6738; Dardadine Rd South between Bunce King Rd and O’Connor Rd, ESE of Dardadine, 8 Nov. 1993, V. Crowley DKN392A; NE of Dinninup, 28 Oct. 1998, R. Davis 8228; c. 23 km NW from Darkan Rock, 25 Aug. 1997, A. Gundry 583; Yarnup Nature Reserve, 25 Oct. 1997, GJ. Keighery & N. Gibson 2631; Kululinup [Kulunilup] Nature Reserve, 26 Oct. 1997, GJ. Keighery & N. Gibson 2746; Albion Rd, c. 32 km SW of Kojonup, 29 Oct. 1997, K.R. Thiele, Hibbertiapolystachya-H. spicata (Dilleniaceae) species group in WA 295 C.M. Lewis 311; Graham Rd, SW Narrogin, 8 Oct. 1998, G. Warren 43; North Kulikup, Darkan, 14 Sep. 2005, A. Webb BNC970; Between Albany Hwy and Boddington, c. 2 km from Albany Hwy, 26 Sep. 1983, J.R. Wheeler 2202; Just W of Cuballing, c. 1.5 km along Cuballing West Rd, 10 Oct. 2001, J.i?. Wheeler AU5. Phenology. Flowers in September and October, most records being from the latter month. Distribution and habitat. Scattered in south-west Western Australia from the vicinity of Boddington to Yarnup Nature Reserve west of Frankland River (Figure 2), growing in grey to red-brown loamy, clay and sandy soils over granite and laterite, in open marri, jarrah and wandoo woodlands. Conservation status. Widespread and scattered, including in several nature reserves, and not considered to be at risk. Etymology. From the Latin aster (a star) with the diminutive -ella, in reference to the leaves and sepals that are densely stellate-pubescent. Notes. Hibbertia asterella is distinctively grey-hoary due to the abundant tubercle-based stellate Figure 2. Distribution of Hibbertia asterella. 296 Nuytsia Vol. 30 (2019) hairs on all parts. While minute stellate hairs are variably found in some other taxa in this group (e.g. H. prolata, H. polystachya) they are a minor component of the indumentum (except in H. capensis which has minutely stellate-hairy sepals). Most specimens of H. asterella lack simple hairs amongst the stellate ones. 2. Hibbertia capensis K.R.Thiele, sp. nov. Type -. Track north of Charles Knife Road; summit of Cape Range, Cape Range National Park, Western Australia, 17 June 2019, GJ. Keighery & K. Lilburn s.n {holo-. PERTH 09085165!; iso-. AD, CANB). Much-branched, erect, compact shrubs to 0.8 m high, multi-stemmed at base and resprouting from the rootstock after fire; young stems sparsely stellate-pubescent with minute hairs; older stems with dull, grey, fissured bark. Leaves spreading to erect, scattered, linear, 15-35 mm long, c. 1 mm wide; margins tightly revolute to the midrib and hiding the abaxial lamina; adaxial lamina glabrous except for fine hairs at the very base, scattered-tuberculate; abaxial lamina obscured by the recurved margins but when visible densely stellate-pubescent; midrib glabrous; apex acute. Inflorescences comprising 2^(-6)-fiowered cincinni, the lowermost fiowers well-separated to overlapping; cincinnaaxis minutely and finely stellate-pubescent; bracts ±heteromorphic with one broader and one narrower and with a iconstricted base, the broader one ovate to broadly ovate or triangular, the lowermost 3^ mm long, 1-2.5 mm wide, finely stellate-pubescent. Sepals very broadly ovate to almost orbicular, 6.5-8 mm long, abaxially very finely and minutely stellate-pubescent; midribs prominent; outer sepals obtuse to subacute; inner sepals similar to the outer but smaller and more obtuse. Petals yellow, broadly obovate, 10-13 mm long, shallowly to deeply emarginate. Stamens 12-16, all on one side of the gynoecium, shortly fused by their filaments; anthers oblong, 1.8-2 mm long, dehiscing by introrse, longitudinal slits. Staminodes 10-12, all behind and lateral to the stamens, filiform. Carpels 2; ovaries rather rectangular-cuboid, densely pubescent; styles excentric from near the carpel apex, ±straight then slightly curved at the apex, 1.8-2 mm long. Ovules 2 per carpel. Fruiting carpels and seeds not seen. Diagnostic features. May be distinguished from all other species in the Hibbertia spicata species group by the combination of very finely and minutely stellate-pubescent sepals and leaves that are glabrous above and densely stellate-pubescent on the lamina below. Selected specimens (all PERTH). WESTERN AUSTRAEIA: Eastern end ofYardie Creek, Cape Range National Park, 2 July 2006, J. English 58; Cape Range - Charles Knife Rd, 30 Aug. 1960,^.5. George 1328; NW Cape - Eighthouse Hill, 31 Aug. 1960, A.S. George 1365; Cape Range Peninsula, small creek 5 km S of Exmouth, 2 Oct. 1995, S. Hunger & N. Kilian 4195; c. 4 miles [6 km] E of Ningaloo Station Homestead, 4 Sep. 1970, A.S. George 10222; Pilgonaman Creek, 67 km from Exmouth, on Yardie Creek Rd, Cape Range National Park, 26 July 1980, K.F. Kenneally 7308; Minilya-Exmouth Rd near Gales Bay, 6 July 2008, K.R. Thiele 3558. Phenology. Flowers from May to November, with a peak in July. Distribution and habitat. Restricted to Cape Range between North West Cape and Ningaloo Station (Figure 3), on red loams and sands over limestone, often in watercourses, in Acacia shrublands, low open mallee and Triodia grasslands. Conservation status. Common in its restricted area of distribution and not considered to be at risk in K.R. Thiele, Hibbertiapolystachya-H. spicata (Dilleniaceae) species group in WA 297 the short term, although its occurrence in an arid environment indicates that it is likely to be adversely affected by climate change. Etymology. From the Latin ‘of the Cape’ in reference to the disjunct distribution on North West Cape. Notes. Hibbertia capensis is the only Hibbertia species found in the semi-arid Cape Range and is widely disjunct from all other species. It shares a limestone habitat with H. leptotheca and H. spicata, the former differs in having glabrous outer sepals and longer, narrower anthers, while the latter has more flowers in the cincinnus and long, simple hairs on the cincinnus bracts and sepals. Leaves of H. capensis have very strongly recurved margins tightly abutting the midrib and entirely obscuring the abaxial leaf lamina. It is often necessary to break leaves to see the stellate lamina pubescence. A specimen from Charles Gardner Flora Reserve near Tammin {R.D. Royce 8377) closely matches H. capensis in leaves and in its sepal indumentum, and does not match H. polystachya, which occurs in the area, or the other widespread species H. prolata. Unless matching material from the locality is found and assigned to a taxon, it is most likely that the sheet is mis-labelled. 298 Nuytsia Vol. 30 (2019) 3. Hibbertia subglabra K.R.Thiele, sp. nov. Type : Coorow- Green Head Road, Western Australia [Precise locality withheld for conservation reasons], 7 September 2019, K.R. Thiele 5565 {holo\ PERTH 09085130; iso\ AD, CANB). Erect, dwarf shrubs to 0.3(-0.5) m high, multi-stemmed at base and resprouting from the rootstock after fire; young stems glabrous except for prominent hair-tufts in the axils; older stems with smooth, papery, silvery bark decorticating in flakes and strips. Leaves erect to spreading, scattered, linear, 12-30(^0) mm long, (0.6-)l-2(-3) mm wide; margins loosely revolute, the abaxial lamina usually exposed; adaxial lamina glabrous except at the very base above the axil, smooth to finely scattered- tuberculate; abaxial lamina and midrib glabrous; apex acute, straight to slightly upturned. Inflorescences comprising 2-6-flowered cincinni, the lowermost flowers usually well-separated; cincinna axis glabrous or sparsely simple-hairy; bracts ±homomorphic, ovate to triangular, the lowermost 2-3.5 mm long, 0.8-1 mm wide, sparsely hairy as for the sepals. Sepals broadly ovate, 3-6 mm long, abaxially with sparse, long, sub-appressed to spreading hairs that are simple or sub-stellate and straight or hooked at the apex; midribs not prominent; outer sepals obtuse; inner sepals similar to the outer but smaller, broader and with fewer hairs, minutely stellate-hairy where covered by the outer. Petals yellow, broadly obovate, 8-10 mm long, shallowly to deeply emarginate. Stamens 9 or 10, all on one side of the gynoecium, shortly fused by their filaments; anthers oblong, 1.5-1.8 mm long, dehiscing by introrse, longitudinal slits. Staminodes 5-7, linear-flattened, behind and lateral to the stamens. Carpels 2; ovaries rather rectangular-cuboid, densely pubescent; styles excentric from near the carpel apex, istraight then slightly curved at the apex, 1.4-1.8 mm long. Ovules 2 per carpel. Fruiting carpels and seeds not seen. Diagnostic features. May be distinguished from all other species in the Hibbertia spicata species group by the glabrous abaxial leaf lamina. Other specimens examined (all PERTH). WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 19 Sep. 1983,i?.J. Cranfleld4266; 15 Oct. 1946, C.A. Gardner M30; 5 Sep. 1976,E.A. Griffin 1006; 18 Sep. \919,E.A. Griffln2214; 10Sept. \999,J.W.Horn2366;3 0a. \912,B.R. Maslin3Q4Q; 4 Sep. 1984, J.R. Wheeler 2343. Distribution and habitat. Restricted to a relatively small area between Cockleshell Gully and Eneabba (Figure 4). A disjunct occurrence near Mogumber {R.J. Cranfleld 4266) may be an erroneous location; searches at the locality have failed to reveal any plants of H. subglabra. Occurs in sand over laterite on slopes and in gullies, in kwongan heath with scattered eucalypts. Conservation status. Range-restricted and known from only seven specimens, some of which occur in conservation reserves. To be listed as Priority Three under Conservation Codes for Western Australian Flora (Smith & Jones 2018). Etymology. From the Eatin glaber (hairless) with the prefix sub- (under, beneath), in reference to the leaves that are abaxially glabrous. Notes. Hibbertia subglabra is distinctive in its glabrous abaxial leaf lamina, all other species in the group being densely stellate-pubescent (except on the midrib). Specimens of all species often have the leaf margins strongly recurved and abutting the midrib, thus obscuring the abaxial lamina; in these cases, care must be taken to find leaves with less strongly recurved margins or that were folded on pressing to expose the abaxial lamina. Occasionally it is necessary to break a leaf to expose the lamina. K.R. Thiele, Hibbertiapolystachya-H. spicata (Dilleniaceae) species group in WA 299 This species often has unusually long, slender leaves compared with e.g. H. prolata. One specimen of H. prolata {J. Liddelow 228) also has a glabrous abaxial lamina; however, in all other respects it is typical of its species and cannot be confused with specimens of H. subglabra. 4. Hibbertia spicata F.Muell, Fragm. (Mueller) 2(11): 1 (1860). Type citation: ‘Ad portum Gregorii [Port Gregory], Walcott et Oldfield.’ (syn: K 687450 image!; MEL 666907 (fragments)!). Erect to spreading, compact to open, sometimes dwarfed shrubs to 0.7 m high, multi-stemmed at base and resprouting from the rootstock after fire; young stems minutely and sparsely stellate-pubescent at first, soon glabrous; older stems with grey, fissured, papery bark. Leaves erect to spreading, scattered, linear, 12-25(-35) mm long, 1.2-2(^) mm wide; margins revolute, the abaxial lamina hidden or exposed; adaxial lamina glabrous, scattered-tuberculate; abaxial lamina densely stellate-pubescent, the midrib glabrous; apex obtuse to sub-acute. Inflorescences comprising (6-)8-14-fiowered cincinni, all fiowers close and overlapping; cincinna axis sparsely stellate-pubescent; bracts heteromorphic, each flower (except the lowermost) subtended by a narrowly obovate-spathulate bract 3-4.5 mm long, 0.8-2 mm wide and a broadly ovate bract 3.5-7 mm long, 2.5-3 mm wide, the bracts sparsely hispid with hairs often restricted to the margins. Sepals ovate, 5-7 mm long, abaxially sparsely pubescent to 300 Nuytsia Vol. 30 (2019) hispid with stellate or simple hairs especially along the midline; midribs not to moderately prominent; outer sepals obtuse; inner sepals similar to the outer but broader and with fewer or no simple hairs. Petals yellow, broadly obovate, 7.5-8 mm long, shallowly to deeply emarginate. Stamens 6(7), all on one side of the gynoecium, shortly fused by their filaments; anthers oblong, 1.5-1.7 mm long, dehiscing by introrse, longitudinal slits. Staminodes 10-14, short and broadly fiattened, all around the gynoecium. Carpels 2; ovaries rather rectangular-cuboid, densely pubescent; styles excentric from near the carpel apex, ±straight then slightly curved at the apex, 2-2.5 mm long. Ovules 2 per carpel. Fruiting carpels and seeds not seen. Diagnostic features. May be distinguished from all other species in the Hibbertia spicata species group by the cincinnus comprising 6-14 fiowers, each fiower (except the lowermost) subtended by two distinctly heteromorphic bracts, one broad and one narrow. Selected specimens (all PERTH). WESTERN AUSTRAEIA: Greenough Rd near Greenough River, 18 Aug. 2008, L. Atkins Sp 317; Spalding Park, 3 miles [5 km] N of Geraldton, 30 Aug. 1965, A.C. Burns 17; Horrocks Rd, 500 m from beach, 7 July 1997, R. Davis 3587; 20 km S of Kalbarri National Park boundary on Grey Rd, 7 Sep. 1997, S. Donaldson & G. Flowers SD1496; 2 miles [3 km] S of Red Bluff, 4 Sep. 1963, A.R. Fairall 1226; 60.4 km W along State Barrier Fence Access track from NW Coastal Hwy, 26 Aug. 1994, G.J. Keighery & N. Gibson 1349; Horrocks, 27 Aug. 1983, C.M. Lynch 154; Pot Alley Gorge, Kalbarri National Park, 26 Sep. 1974, G. Perry 301; Shark Bay, 29 Sep. 1989, M.E. Trudgen 7400; Chapman River Regional Park, Geraldton, 2 June 1999, S. Vigilante 61; Red Bluff, Kalbarri, 6 Sep. 1984, J.R. Wheeler 2375. Phenology. Flowers from April to October, with a peak in early September. Distribution and habitat. Occurs on coastal limestones from Geraldton to the southern end of Shark Bay (Figure 5). A few specimens have been recorded from coastal sandstone bluffs (e.g. Red Bluff) close to limestone. Grows in sandy and clay soils, in coastal heathlands, shrublands and low woodlands. Conservation status. Relatively widely distributed including in several national parks and nature reserves, and not considered to be at risk. Notes. Specimens of H. spicata are immediately recognisable by the many-fiowered cincinni with the fiowers and bract overlapping, giving a distinctively crowded appearance. In all other species the fiowers are fewer and more widely-spaced {H. capensis sometimes also has overlapping fiowers in the cincinnus). The cincinnus bracts in H. spicata are also distinctively heteromorphic. One bract is obliquely inserted on the cincinnus axis and is large and broad, while the other is transversely inserted and is much narrower. 5. Hibbertia leptotheca (J.R.Wheeler) K.R.Thiele, comb, et stat. nov. Hibbertia spicata Benth. subsp. leptotheca J.R.Wheeler, Nuytsia 5(1): 35-37 (1984). Type. Yalgorup National Park, between the north end of Take Preston and Martin’s Tank Take, Western Australia, 17 September 1981, NG. Marchant 81/76 {holo\ PERTH 1627198!; iso\ CANB 363019 n.v, MEE 6775771). Small, spreading shrubs to 0.3 m high; young stems glabrous, multi-stemmed at base and resprouting from the rootstock after fire; older stems with grey, papery-fissured bark. Leaves spreading, scattered. K.R. Thiele, Hibbertiapolystachya-H. spicata (Dilleniaceae) species group in WA 301 Figure 5. Distribution of Hibbertia spicata. linear, (8-)12-25(-35) mm long, 0.8-1 mm wide; margins revolute and hiding the abaxial lamina or nearly so; adaxial lamina glabrous, smooth to rarely sparsely scattered-tuberculate; abaxial lamina where visible densely stellate-pubescent, the midrib glabrous; apex acute. Inflorescences comprising 2-5-flowered cincinni, the flowers well-separated; cincinna axis glabrous; bracts ±homomorphic, linear to narrowly obovate, 2-A mm long, 0.8-1.5 mm wide, glabrous. Sepals ovate, 3.5-5 mm long, the outermost abaxially glabrous; midribs not prominent; outer sepals broadly acute; inner sepals similar to the outer but broader, more obtuse, glabrous or finely and sparsely minutely stellate-pubescent. Petals yellow, broadly obovate, 5-6 mm long, shallowly to deeply emarginate. Stamens (8-)10-12(- 15), all on one side of the gynoecium, shortly fused by their filaments; anthers linear, 1.8-2.8 mm long, dehiscing by introrse, longitudinal slits. Staminodes 0(-3), when present lateral to or behind the stamens. Carpels 2; ovaries rather rectangular-cuboid, densely pubescent; styles excentric from near the carpel apex, ±straight then slightly curved at the apex, 2-2.5 mm long. Ovules 2 per carpel. Fruiting carpels and seeds not seen. Diagnostic features. May be distinguished from all other species in the Hibbertia spicata species group by the glabrous outer sepals and long, narrow anthers. Selected specimens (all PERTH). WESTERN AEISTRAEIA: [localities withheld for conservation reasons] 20 Sep. 2003, P. Foreman 391; 8 Dec. 1992, E.A. Grijfln 8401; 1 Sep. 1897, R. Helms s.n. 302 Nuytsia Vol. 30 (2019) (PERTH 2625482); 21 Sep. 1990, GJ. Keighery 11492; 14 Sep. 1995, GJ. Keighery 14093; 25 Oct. 1982, J.R. Wheeler 27 Aug. 1957, C.L. Wilson 870. Phenology. Flowers between August and October, with a peak in September. Distribution and habitat. Occurs in coastal and near-coastal sites from west of Cataby south to Lake Preston (Figure 6), growing in sand over limestone in coastal heaths and thickets usually dominated by species of Melaleuca and Acacia. Conservation status. Listed as Priority Three under Conservation Codes for Western Australian Flora (Smith & Jones 2018), under the name H. spicata subsp. leptotheca. Notes. Wheeler (1984) described H. leptotheca as a subspecies of H. spicata s.l. She presumably used subspecies rank due to the wide variation she perceived in the latter, and because elsewhere in its range H. spicata s.l. shares a limestone habitat with H. leptotheca (i.e. the species segregated here as H. capensis and H. spicata s.s.). However, it is clearly separate both geographically and morphologically from other taxa in the group, with no intermediates, and constitutes a separately evolving lineage. I regard that species rank is appropriate for this taxon. 6. Hibbertia prolata K.R.Thiele, nom. nov. Hemistephus linearis J.Drumm. ex Harv., in Hooker, W. J. (ed.). Hooker’s Journal of Botany and Kew Garden Miscellany 7:52(1855); Hemistemma line are (J. Drumm. ex Harv.) F. Muell., Fragm. (Mueller) 1(7): 162 (1859). Type citation'. ‘Northern Districts.’ {sym TCD 9690 image!, MEL 6666641). Dense to open, often rounded, erect to spreading shrubs to 0.8 m high, multi-stemmed at base and resprouting from the rootstock after fire; young stems glabrous to sparsely stellate-pubescent with minute hairs, soon glabrous; older stems with grey, papery, fissured bark. Leaves erect to spreading, scattered, linear to very narrowly obovate, (10-) 12-25(-30) mm long, 0.8-2 mm wide; margins revolute and usually hiding the abaxial lamina or nearly so; adaxial lamina glabrous, scattered-tuberculate; abaxial lamina densely stellate-pubescent, the midrib glabrous; apex acute, sometimes bluntly so. Inflorescences comprising (l-)2-5(-7)-flowered cincinni, the flowers well-separated; cincinna axis glabrous to sparsely stellate-pubescent; bracts ±homomorphic, linear to narrowly obovate, 2-3.5(-5) mm long, 0.5-1.5 mm wide. Sepals ovate, 5-6 mm long, abaxially with few to many, spreading to retrorse, hooked hairs mostly towards the base, sometimes underlain by sparse stellate hairs; midribs prominent; outer sepals acute; inner sepals broader and thinner-textured than the outer, with sparser hooked hairs. Petals yellow, broadly obovate, 6-8.5 mm long, shallowly to deeply emarginate. Stamens 10(-12), all on one side of the gynoecium, shortly fused by their filaments; anthers oblong, 1.2-1.8 mm long, dehiscing by introrse, longitudinal slits. Staminodes present, 6-20, behind, lateral to and/or opposite the stamens. Carpels 2; ovaries rather rectangular-cuboid, densely pubescent; styles excentric from near the carpel apex, ±straight then slightly curved at the apex, c. 1.5 mm long. Ovules 2 per carpel. Fruiting carpels and seeds not seen. (Figure IB) Diagnostic features. May be distinguished from all other species in the Hibbertia spicata species group by the leaf lamina that is adaxially glabrous and abaxially densely stellate-pubescent, and the outer sepals with few to many hooked hairs. K.R. Thiele, Hibbertiapolystachya-H. spicata (Dilleniaceae) species group in WA 303 Selected specimens (all PERTH). WESTERN AUSTRAEIA: Dingo Hill, 5 km along First North Rd from Eneabba-Three Springs Rd, 11 Oct. 2003, J. Borger BB212; junction of Cape Naturaliste Rd and Bunker Bay Rd, Dunsborough, 13 Oct. 2008, H. Cole 744; 3 km E of Woodanilling, 3 Nov. 1978, R.J. Cranfield s.n. \ behind Gillingarra School, 6 Oct. 2006, C Danese & D. Rayner BK 1006- 06; Greenough River, 1 km W of the first railway crossing on Geraldton-Mullewa Rd, 55 km W of Mullewa, 20 Oct. 1983, SJ. Forbes 1714; Wagin, 26 Oct. 1920, C.A. Gardner 1003; Avondale Research Station, 6 km W of Beverley, 22 Oct. 1979, RJ. Hnatiuk 790181; near The Casuarinas (c. 36 miles [58 km] E of Geraldton), 18 Sep. 1971, R.D. Hoogland 11982; along Northwest Coastal Hwy S of Ajana-Kalbarri turn-off, near mile peg 369, 20 Sep. 1971, R.D. Hoogland 11996; S side of Eneabba- Three Springs Rd, 6.4 km W of its Junction with Reserve Rd, 25 Aug. 2001, J. W. Horn 4008; Benn Reserve, 1 km NW of Kojonup, 14 Nov. 1999, C.M. Lewis 464; Serpentine National Park, about 750 m ESE of Chatfield Rd, 31 Oct. 1996, A. Markey 492; Red Hill, 22 Sep. 1944, R. D. Royce s.n. (PERTH 3107299); Helena Valley, 26 Sep. 1977, J. Seabrook 3\2; 2 miles [3.5 km] W of Eradu on Mullewa-Geraldton Rd, 1 Oct. 1966, E.M. Scrymgeour 1449; Helena Valley, Ridge Rd, 25 Sep. 1981, J.R. Wheeler 2013; Jarrahdale Scenic Drive, 2.5 km along Barge Rd from South West Hwy, 5 Oct. 1983, J.R. Wheeler 2231; 4.5 km N of Binnu, 6 Sep. 1984, J.R. Wheeler 2372; Eagle Bay, Meelup, 7 Sep. 1985, J.i?. Wheeler 2AQ2. Phenology. Flowers between June and November, with a peak in September. 304 Nuytsia Vol. 30 (2019) Distribution and habitat. Widely distributed from Binnu to Katanning, in the southern part of its range occurring in the far western Wheatbelt and along the Darling Range, with a disjunct population around Cape Naturaliste (Figure 7). A few collections from Jurien Bay and Green Head are close to the coast, but these are not usually on limestone. Mostly found in sandy soils over granite or laterite, in woodlands, mallee, shrublands and heath. Conservation status. Widespread and common and not considered to be at risk. Etymology. From the Latin prolatus (extended, elongated) in reference to the cincinnus having rather well-separated flowers compared with H. spicata s.s., from which it has been segregated. Notes. Hibbertia prolata is a widespread species. It tends to occur westwards of the range of the other widespread species, H. poJystachya, but there is substantial overlap. Most specimens of H. prolata have a distinctive sepal indumentum, comprising few to many spreading to retrorse, hooked hairs mostly concentrated towards the sepal base. The only other species in the group with a similar indumentum is H. subglabra, which differs in having the abaxial leaf lamina glabrous. The disjunct specimens from near Cape Naturaliste are typical in all other respects. A few specimens (e.g. R.D. Royce s.n. PERTH 3107299; R.J. Hnatiuk790\8\) lack these hairs and appear superficially similar to H. leptotheca but can be distinguished from that species by the shorter, less slender anthers. Seven specimens collected between Gillingarra, Jurien Bay and Arrino lack the hooked hairs and have a denser sepal indumentum of simple, ±appressed hairs. These are provisionally included in H. prolata pending further held work. 7. Hibbertia polystachya Benth, FI Austral. 1: 22-23 (1863). Type citation'. ‘W. Australia. Swan River, Drummond; Blackwood river, Oldfield. ’ {sym [Oldfield] MEL 666681!; [Drummond] K 687446 image!, K 687447 image!, K 687448 image!, MEL 666680!). Compactto straggling, often sprawling^/2rwfeto0.3(-0.7)mhigh, multi-stemmed at base andresprouting from the rootstock after Are; young stems sparsely to moderately stellate-pubescent, sometimes also with scattered simple hairs; older stems with grey, papery bark decorticating in strips. Leaves erect to spreading, scattered, narrowly elliptic to linear, 5-20 mm long, 0.8-3 mm wide, the margins revolute with the abaxial lamina usually exposed; adaxial lamina with sparse to abundant, short to long, soft, simple, often tubercle-based hairs, sometimes also with scattered, tubercle-based stellate hairs; abaxial lamina densely stellate-pubescent, the midrib with indumentum as for the adaxial lamina; apex obtuse to subacute. Inflorescences comprising (l-)2-5-flowered cincinni, the flowers well-separated; cincinna axis sparsely pubescent with usually long simple hairs, sometimes also with intermixed stellate hairs; bracts ±homomorphic, narrowly ovate to linear, 3-6 mm long, 1-2 mm wide, with indumentum as for the leaves. Sepals narrowly ovate to ovate, 7-10 mm long, abaxially pubescent to pilose with long, simple hairs sometimes intermixed with hooked and/or stellate hairs; midribs prominent; outer sepals often leaf-like with recurved margins, acute; inner sepals thinner, broader and less hairy than the outer, with broader (to obtuse) apex. Petals yellow, broadly obovate, 7.5-12 mm long, shallowly to deeply emarginate. Stamens (6-) 10(11), all on one side of the gynoecium, shortly fused by their filaments; anthers oblong, 1.6-2 mm long, dehiscing by introrse, longitudinal slits. Staminodes 5-10, mostly lateral to and opposite the stamens (a few sometimes behind the stamens), those opposite sometimes broad and petaloid. Carpels 2; ovaries rather rectangular-cuboid, densely pubescent; styles excentric from near the carpel apex, ±straight then slightly curved at the apex, 1.2-1.8 mm long. Ovules 2 per carpel. Fruiting carpels and seeds not seen. K.R. Thiele, Hibbertiapolystachya-H. spicata (Dilleniaceae) species group in WA 305 Diagnostic features. May be distinguished from all other species in the Hibbertia spicata species group by the adaxial leaf laminas and sepals with sparse to abundant simple hairs. Selected specimens WESTERN AUSTRALIA: 11.5 km S ofCachionalgoHill, offBindoon- Dewars Pool Rd, 7 Nov. 1996, M.G. Allen 303; 5 km NE of Cataby, 28 Oct. 1993, P. Armstrongs.n.; 8.5 km NW of Calingiri, 20 Sep. 1983, R.J. Cranfield 4339; Dobaderry Rd, 4.6 km N of Dale West Rd intersection, 8 Oct. 1997, R. Davis 4256; Bowelling-Duranillin Rd, 2.9 km from Bowelling, 17 Nov. 1997, R. Davis 4471; Jingaring Reserve, c. 33 km ESE of Brookton, 1 Sep. 1998, R. Davis 6526; Wambyn Nature Reserve, 9 Oct. 1998, R. Davis 7148; N end of Cony Rd, W of Corrigin, 24 Sep. 2007, M Hislop & H. Mills WW 209-38; Along main road from Gingin to Dongara at crossing with Mullering Brook, 15 Sep. 1971, R.D. Hoogland 11969; Brand Hwy, 2.3 km S of its junction with Yandin Rd, 22 Aug. 2001, J. W. Horn 4000; Kondinin-Narembeen Rd, 220 m S of Billericay-East Rd, 24 Sep. 1997, G.J. Keighery & N. Gibson 3715; Narlingup Reserve, 28 km W of Kojonup, 17 Oct. 1997, C.M. Lewis 300; Stirling Rd, 25 km WNW of Kojonup, 5 Nov. 2000, C.M. Lewis 486; Airstrip Rd, 0.3 km from intersection of Jilakin Rd, 21 Oct. 2001, S. Murray 525; 73 km from Perth along main road to Brookton, by Christmas Tree Well, 27 Oct. 1982, A. Strid 21108; Chittering Valley Rd 35.8 km from Bullsbrook towards Bindoon, 5 Sep. 1982, J.R. Wheeler 2034; Dumbleyung-Lake Grace Rd, 17 km E of Dumbleyung, 21 Sep. 1986, J.R. Wheeler 2405; Junction [of] Talbot West Rd and Talbot Rd, 9 Oct. 2001, J.R. Wheeler 4129; Beekeepers Reserve, 18 Sep. 1985, R.T. Wills 110. 306 Nuytsia Vol. 30 (2019) Phenology. Flowers between August and December, with a peak in October. Distribution and habitat. Widely distributed in the Western Australian wheatbelt and Darling Range from east of Mingenew to Katanning and west to near Koj onup, generally inland from the coast (Figure 8). Occurs on sandy, loamy and clay soils generally over granite or laterite, in wandoo woodlands. Acacia md Allocasuarina shrublands, mallee, and heathlands. Conservation status. Hibbertia polystachya is widely distributed and common and is not considered to be at risk. Notes. Hibbertia polystachya is more widespread and variable in leaf, cincinnus and sepal indumentum than any other species in the H. spicata-H. polystachya group. It tends to occur inland (east) of the other widespread taxon, H. prolata, although there is substantial overlap. Indumentum varies from sparsely pubescent with short, straight simple hairs arising from prominent tubercles to villous with long, soft, silky hairs without distincttubercles. Theremay also be varying development of short stellate hairs interspersed with the simple ones. The variation, however, is continuous and, although there is a tendency for villous specimens to be at the western edge of the range, not geographically partitioned. Figures. Distributionof///'Zj6ert/a/7o/>’stoc/?3^a. K.R. Thiele, Hibbertiapolystachya-H. spicata (Dilleniaceae) species group in WA 307 In almost every case, specimens of H. polystachya may be distinguished from H. prolata by the presence of simple hairs on the adaxial leaf surface at least on young leaves. Leaves in H. prolata are adaxially glabrous. One specimen (J. Liddelow 228) from near Lake Muir is unusual in having a glabrous abaxial leaf lamina (as for H. subglabra). However, it is typical for H. polystachya in all other respects, with simple-hairy adaxial leaf lamina and abaxial midrib, and typical sepal indumentum. There are four Drummond specimens of H. polystachya, three at K and one at MEL, annotated variously as Drummond 15, Drummond 12, Drummond 17 and Drummond s.n.. All appear to be separate gatherings based on differences in leaf and sepal indumentum, and all fall within the range of variation of H. polystachya in these characters. Given that Bentham did not specify a Drummond number in the protologue for H. polystachya, all are here regarded as syntypes. I choose not to lectotypify as no benefit is to be gained by doing so at this time and I prefer to leave lectotypification as an option for future workers if the need arises. Summary of taxon circumscriptions Hibbertia polystachya Benth. in this paper has the same circumscription as H. polystachya Benth. as previously accepted at PERTH. Hibbertia spicataY MwqW. in this paper has a narrower circumscription than 77. spicataYMwQll. subsp. spicata as previously accepted at PERTH due to the removal of 77. asterella K.R.Thiele, 77. subglabra K.R.Thiele, 77. capensis K.R.Thiele and 77. prolata K.R.Thiele. Hibbertia asterella K.R.Thiele sp. nov. in this paper is removed from 77. spicata F. Muell. subsp. spicata as previously accepted at PERTH. Hibbertia subglabra K.R.Thiele sp. nov. in this paper is a new taxon and is removed from 77. spicata F. Muell. subsp. spicata as previously accepted at PERTH. Hibbertia capensis K.R.Thiele sp. nov. in this paper is a new taxon and is removed from 77. spicata F. Muell. subsp. spicata as previously accepted at PERTH. Hibbertia prolata K.R.Thiele nom. nov. in this paper is removed from 77. spicata F.Muell. subsp. spicata as previously accepted at PERTH. Hibbertia leptotheca (J.R.Wheeler) K.R.Thiele comb, et stat. nov. in this paper has the same circumscription as 77. spicata subsp. leptotheca as previously accepted at PERTH. Acknowledgements I thank the staff of the Western Australian Herbarium (PERTH) and National Herbarium of Victoria (MEE) for access to their collections. 308 Nuytsia Vol. 30 (2019) References Bentham, G. (1863). FloraAustraliensis. Vol. 1. (Reeve and Co.: London, United Kingdom.) Harvey, W.H. (1855). Characters of some new genera of plants recently discovered by Mr. James Drummond in Western Australia. In: Hooker, W.J. (ed.). Hooker’s Journal of Botany and Kew Garden Miscellany 1: 51-52. Horn, J. W. (2005). The phylogenetics and structural botany of Dilleniaceae and HibbertiaAndxQws. PhD thesis, Duke University, Durham, NC, USA. Mueller, F.J.H. von (1860). Fragmenta Phytographiae Australiae 2(11); 1-2. Smith, M.G. & Jones, A. (2018). Threatened and Priority Flora list December 2018. Department of Biodiversity, Conservation and Attractions. https;//www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 20 June 2019]. Wheeler, J.R. (1984). Taxonomic notes on some Western Australian species of Hibbertia (Dilleniaceae). Nuytsia 5(1); 31-42. Wheeler, J.R. (1987). Hibbertia. In: Marchant, N.G., Wheeler, J.R., Rye, B.L., Bennett, E.M., Lander, N.S. & Macfarlane, T.D. (eds) Flora of the Perth Region. Part 1. pp. 119-133. (Western Australian Herbarium: Perth, Western Australia.) Wheeler, J.R. (2004). An interim key to the Western Australian species of Hibbertia (Dilleniaceae). Nuytsia 15(2): 311-320. Nuytsia The journal of the Western Australian Herbarium 30:309-316 Published online 10 December 2019 A new name, clarification of synonymy, and a new subspecies for Isopogon (Proteaceae) in Western Australia Barbara L. Rye and Terry D. Macfarlane Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 Abstract Rye, B.L. & Macfarlane, T.D. A new name, clarification of synonymy, and a new subspecies for Isopogon (Proteaceae) in Western Australia. Nuytsia30\ 309-316 (2019). Isopogon drummondii Benth. nom. illeg. is lectotypified and replaced by the new name I autumnalis Rye & T.Macfarlane while I drummondii Hiigel ex Jacques is recognised as a probable synonym of I sphaerocephalus Lindl. The new subspecies I sphaerocephalus subsp. lesueurensis Rye is described. Introduction This paper undertakes to settle the uncertainty around the name of a Western Australian species of Isopogon R.Br. (Proteaceae), the solution to which involves a second species, in which a new subspecies is recognised. In his treatment of Isopogon for the Flora of Australia, Foreman (1995) described one species as I sp. A because he was uncertain whether the name I drummondii Hiigel ex Jacques (Jacques 1843) applied to it. A later-published name with the same epithet, I drummondii Benth. (Bentham 1870), does apply to Foreman’s species A but cannot be used because it is an illegitimate later homonym. The name I drummondii Hiigel ex Jacques was not mentioned by Bentham (1870), possibly because it was published in a horticultural context. It was based on cultivated plants with no precise locality of origin, has no illustration and evidently lacks a type, problems commonly encountered among Jacques’s names. The protologue does not give any information regarding the flowers and fruits of the species but does provide a moderately detailed description of its leaves. Whereas Jacques’s species was described as having linear leaves 6-15 cm (i.e. 60-150 mm) long, Bentham’s species has terete leaves 20-65 mm long. This difference alone is sufficient to show that they are distinct entities, so I drummondii Benth. needs to be replaced by a new, legitimate name. The new name chosen here is I autumnalis Rye & T.Macfarlane. A far more likely match for I drummondii Hiigel ex Jacques in leaf morphology is I sphaerocephalus Lindl., an attractive species that was collected by James Drummond during the early stages of settlement of the Swan River area of Western Australia. It has a moderately large range from near Gidgegannup to the south coast, with an isolated, distinctive variant occurring much further north near Mt Lesueur. This northern variant is named here as subsp. lesueurensis Rye. © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 310 Nuytsia Vol. 30 (2019) Taxonomy Isopogon autumnalis Rye & T.Macfarlane, nom. nov. Isopogon drummondiiBmth., FI. Austral. 5: 344—345 (1870), nom. illeg., Atylus drummondii Kuntz, Revis. Gen. 2: 577 (1891). Type citation. ‘Swan River, Drummond, \st coll., Preiss\ Type specimens: Swan River [Western Australia], 1839, J. Drummond 1st coll, s.n. (lecto, here designated: K 000736646 image!; isolecto: BM 000991907 image!, K 000736645 image!); Swan River [Western Australia], loc.,J.A.L. Preiss s.n. {syn: MEL 1531863 image!). Isopogon sp. A; D.B. Foreman, FI. Australia 16: 204 (1995). [Isopogon drummondii auct. wowHiigel ex Jacques: Council ofHeads ofAustralasianHerbaria, Plant Census, https://biodiversity.org.au/nsl/services/APC (2006-2019); FloraBase, https://florabase. dpaw.wa.gov.au/ (2006-2019).] [Isopogonpetrophiloides auct. non R.Br.: C.F. Meissner in J.G.C. Lehmann, PI. Preiss 1: 503 (1845).] Illustrations.^E. Blackall &B.J. Gx'xqwq, How Know W. Austral. Wildflowers 1:146 (1988) [dcslsopogon drummondii^, D.B. Foreman, FI. Australia 16: 206, fig. 100 (1995) [as Isopogon sp. A]. Shrubs 0.3-1 m high, commonly 0.5-1 m wide, lignotuberous. Young stems with a very dense, short indumentum of whitish, coiled hairs and also with patent hairs 0.4-0.7 mm long. Leaves mostly curved, simple, (20-)30-50(-65) mm long, 1.3-2.1 mm wide, broadest towards the apex, terete and glabrous except for the base, mucronate; petiole or flattened base c. 4 mm long, densely coiled-hairy on abaxial surface and sometimes on basal part of adaxial surface; mucro erect, stout, c. 1 mm long, pale at first but usually with a dark tip, becoming dark. Flower heads solitary, terminal, erect, very broadly or depressed ovoid and 20-35 mm diam. when in full fiower (often smaller and broadly ovoid in bud and fruit), with numerous, densely packed involucral and floral bracts and closely surrounded by leaves; bracts very densely hairy outside with long whitish hairs, sometimes with a short, dark, glabrous tip. Tepals usually 10-12 mm long, pale yellow; claw covered throughout by widely spreading wavy (curved or loosely curled), silky hairs, separating to just below the base of the pollen presenter, which extends slightly below the base of the limb; limb 3.5^ mm long, usually ciliate in lower part, with a terminal tuft of hairs 0.3-0.6 mm long, the four parts of the limb becoming curved after separating from one another. Anthers 2.3-2.5 mm long including terminal appendage; connective appendage 0.45-0.5 mm long, becoming dark. Pollen presenter 3-3.5 mm long, densely and minutely papillose-hairy on the pedestal and swelling, with a slight to obvious constriction separating the pedestal from the much shorter swelling; pedestal slightly to distinctly shorter than the receptor; swelling c. 0.3 mm wide; receptor 1.5-2.3 mm long, lacking papillae or with papillae restricted to the base. Cones very broadly ovoid; involucral bracts similar to cone scales. Cone scales spathulate, up to c. 10 mm long, with margin incurved; outer surface very densely villous; inner surface glabrous or sparsely hairy. Diaspores ovoid, 5-5.5 mm long, 2.3-2.5 mm diam.; largest hairs widely spreading (with some directed downwards), c. 5 mm long; mature seed not seen. Diagnostic characters. Distinguished from all other members of the genus in having the following combination of characters: simple, terete leaves and pale yellow fiowers. Selected specimens examined. WESTERN AUSTRALIA [localities withheld for conservation reasons]: B.L. Rye & T.D. Macfarlane, A new name and a new subspecies for Isopogon (Proteaceae) 311 23 July 2001, F. Hort 1322 (PERTH); 26 Feb. 2003, GJ. Keighery 16418 (PERTH); 8 June 2006, M Morley & V. English JP 1 (PERTH); 27 Apr. 2013, K.R. Thiele 4767 (PERTH). Distribution and habitat. Occurs in sandy soils, often in Banksia woodlands, near the west coast of southern Western Australia in the Geraldton Sandplains, Jarrah Forest and Swan Coastal Plain bioregions. The distribution extends from Cockleshell Gully (north-east of Jurien Bay) south to Serpentine (Figure lA). Phenology. Flowers recorded primarily in the first half of the year, especially from February to May, and mature fruits recorded from March to May. Conservation status. Currently listed as Priority Three under Conservation Codes for Western Australian Flora (Smith & Jones 2018). This species appears to be susceptible to dieback. Etymology. From the Eatin adjective autumnalis (autumnal) to refiect the species’ unusual fiowering period, which peaks in autumn. Most other species of Isopogon have their main fiowering time in spring. Vernacular name. Autumn Isopogon. Figure 1. Distribution maps. A - Isopogon autumnalis (•); B - 7. sphaerocephalus subsp. lesueurensis (o) and subsp. sphaerocephalus (•). 312 Nuytsia Vol. 30 (2019) Affinities. This species seems very distinctive. It was placed in sequence between I. longifolius R.Br. and I. villosus Meisn. by Foreman (1995) but is unlikely to be confused with either of those species and its closest affinities are uncertain. Typification. Foreman (1995: 204) listed Kew material of the Drummond syntype, which he had examined and confirmed as type material of I. drummondii on 12 January 1994, but did not list any material of the Preiss syntype. Three specimens of the J. Drummond s.n. syntype currently have images on Global Plants, two from the same Kew sheet annotated by Bentham and Foreman, and the other from the Natural History Museum (BM), but none of the Preiss syntype. A Preiss specimen that bears the “B” annotation indicating that it was seen by Bentham has been located at MEL (1531863); this specimen, is consistent with the syntype citation in lacking collection details. We have selected one of the Drummond specimens, K 000736646, as the lectotype because it has two inflorescences and the date 1839, which accords well with the attribution to Drummond’s first collection. The other specimen on the same sheet, K 000736645, is from Hooker’s herbarium but has only one inflorescence and no date indicated, while the BM 000991907 specimen is not annotated by Bentham. The Preiss syntype at MEL was not chosen because it is in a poorer physical condition. In the protologue for/, drummondii Benth., Bentham (1870:345) lists the synonymy as ‘/ petrophiloides, Meissn. in PI. Preiss. i. 503, and in DC. Prod. xiv. 276, partly, but not of Br.’, and notes ‘The foliage of the species is nearly that of the undivided states of I. teretifolia and 1. scabriuscula, with the former of which (the / petraphiloides, Br.) it may have been confounded by Meissner, as he quotes Baxter’s specimens as well as Drummond’s and Preiss’s.’ This note refers to a partial misapplication by Meisner (1845,1856)ofthe name/R.Br. to Bentham’s new species/ drummondiiBmth. inregard to Meisner’s citation of two numbered Preiss specimens (679, 680) and an un-numbered Drummond first collection specimen. These are all represented in the de Candolle Prodromus herbarium at Geneva (G-DC; International Documentation Centre 1962). It is possible that the un-numbered Preiss syntype at MEL is a duplicate of one of the numbered specimens cited by Meisner under / petrophiloides, but evidently its number had been lost before Bentham saw it. Notes. A description of / autumnalis [as / drummondii Benth.] was given in the Flora of the Perth Region (Rye 1987). Later, the species was treated for some time as / drummondii Hiigel ex Jacques. See the discussion above and below outlining why the latter name is now considered likely to be a synonym of / sphaerocephalus. Isopogon sphaerocephalus Lindl, Sketch Veg. Swan R. 34 (1840); Atylus sphaerocephalus (Lindl.) ¥amtzQ,Revis. Gen. PI. 2:577 (1891). Type\ Swan River district [Western Australia], 1839, J. Drummond s.n. (lecto: CGE [image PERTH 015539761], fide D.B. Foreman, Flora of Australia 16: 480 (1995); isolecto: GH 00035491 image!, MEL 672802 image!). Shrubs 0.3-2 m high, 0.3-1.5 m wide. Young stems with long patent hairs and more numerous very short coiled hairs. Leaves sessile, erect, simple, linear to narrowly obovate, 50-160 mm long, 4-18 mm wide, with a somewhat obtuse, mucronate apex; margins tending to be recurved throughout their length, entire; mucro erect, triangular, 1.2-1.6 mm long, dark-coloured but the tip sometimes pale or the whole mucro covered by appressed, whitish hairs. Flower heads solitary or crowded with one or two others at end of a stem, erect, ± globular, commonly 25-30 mm diam., with numerous, densely packed involucral and floral bracts, and subtended by very short leaves that grade into the much larger leaves below; bracts hidden when head is in full flower, very densely hairy outside with long whitish hairs. Tepals 10-15 mm long, cream to moderately deep yellow, usually pale yellow, separating down to the base of the pollen B.L. Rye & T.D. Macfarlane, A new name and a new subspecies for Isopogon (Proteaceae) 313 presenter, which extends slightly below the base of the limb; claw glabrous throughout or with few hairs at the summit; limb 3.5^ mm long, densely hairy with widely spreading, white hairs 0.7-2 mm long, the four parts of the limb becoming strongly curved after separating from one another. Anthers 2.2-2.8 mm long including terminal appendage; connective appendage 0.4-0.6 mm long, becoming dark. Pollen presenter 2.3-5 mm long, with an obvious constriction between the pedestal and swelling; pedestal 0.4-2 mm long, densely covered by hair-like or elongated papillae up to 0.25 mm long, which may protrude along four main longitudinal ridges to form four thick rows; swelling 0.35-0.55 mm wide; receptor 1.4-2.2 mm long, usually without papillae. Cones ovoid to cylindrical at first but often becoming broader when diaspores are shed; involucral bracts narrowly ovate, 7-8 mm long, attenuate or with a long apical point, with similar indumentum to the cone scales. Cone scales ovate or broadly ovate, as long as, or somewhat shorter than, involucral bracts; outer surface very densely hairy, with spreading hairs up to 2.2 mm long; inner surface glabrous; apical point hairy or becoming glabrous and dark. Diaspores ovoid, 3.5-5.3 mm long, 1.8-2.4 mm diam.; largest hairs widely spreading (with some directed downwards), 5-7 mm long; seed c. 3 mm long, c. 2 mm wide. Diagnostic features. Distinguished from all other members of the genus by the following combination of characters: leaves linear to narrowly obovate, entire, with an erect mucro but with margins tending be recurved throughout their length; tepals usually pale yellow, with a glabrous claw and hairy limb. Notes. An unusual characteristic of I. sphaerocephalus is its tendency to have slightly to markedly recurved leaf margins from the base to the apex. The closest approach to the fully recurved leaf margins of I. sphaerocephalus is in I. cuneatus, which has the base more or less flat or slightly incurved to slightly recurved. Other species of Isopogon have the leaf margins flat or incurved to meeting on the upper surface, or have the lamina recurved only distally. Two geographically separated subspecies are recognised here. a. Isopogon sphaerocephalus Lindl. subsp. sphaerocephalus llsopogon drummondii Hiigel ex Jacques, Ann. FI. Pomone ser. 2,1:216 (1843) [as Drumundii]. Type citation'. ‘Lieu originaire, laNouvelle-Hollande’. Isopogon eriocladus Gand., Bull. Soc. Bot. France 66: 228 (1919). Type', near Serpentine, Darling Range, Western Australia, September 1900, W.V. Fitzgerald s.n. {syn'. LY n.v.,fide D.J. McGillivray, Contr. New South Wales Natl. Herb. 4: 344 (1937)). Isopogon ovoideus Gand., Bull. Soc. Bot. France 66: 228 (1919). 7)pe: Smith’s Mill, Darling Range, Western Australia, September 1900, A. Morrison s.n. Qiolo: LY (two pieces mounted on same sheet) n.v.,fide D.J. McGillivray, Contr. New South Wales Natl. Herb. 4: 344 (1937)). Illustrations. A.S. GQOXgQfntr. Proteaceae W. Australia 83, Plate 123 (1984); N.G. Marchant, J.R. Wheeler, B.L. Rye, E.M. Bennett, N.S. Lander & T.D. Macfarlane, FI. Perth Region 1: 347, Figure 125 (1987); WE. Blackall & B.J. Grieve, How Know W. Austral. Wildflowers 1: 144 (1988); D.B. Foreman, FI. Australia 16: frontispiece & 220, Figure 1031 (1995); J.R. Wheeler, N.G. Marchant & M. Eewington, FI. South West 2: 838 (2002) [all as I sphaerocephalus]. Shrubs 0.3-2 m high, lignotuberous. Flowering branches moderately to densely leafy, moderately to densely hairy. Leaves 50-160 mm long, 4-12 mm wide, glabrous or sparsely hairy. Tepals 10-15 mm long. Pollen presenter 2.3-3.3 mm long, with papillae 0.15-0.25 mm long on approximately the basal 314 Nuytsia Vol. 30 (2019) third of the presenter; pedestal 0.4-1 mm long; swelling usually glabrous, sometimes with some very short papillae on the base, rarely with long papillae on one side only. Diaspores 3.5^ mm long, c. 22 mm wide; largest hairs 5-6 mm long; seed c. 2.8 x l .8 mm. Selected specimens examined. WESTERN AUSTRAEIA: McCorkill Forest Block, 22 km WNW of Nannup, 11 Nov. 2009, RJ. Cranfield 2441 (PERTH); reserve near comer of Hardey Rd and Strettle Rd, Glen Forrest, 15 Sep. 2008, K.R. Thiele 3688 (PERTH). Distribution and habitat. This subspecies extends from near Gidgegannup (north-east of Perth) south to the Scott River area and south-east to near Kent River (Figure IB), in lateritic areas with Jarrah forest, sometimes associated with watercourses. This distribution extends through near-coastal parts of the far south-west of Western Australia, in the Jarrah Forest, Swan Coastal Plain and Warren bioregions. Phenology. Flowers and fruits recorded more or less throughout the year but with the main flowering season apparently from July to December. Common name. Drumstick Isopogon. Conservation status. Not considered to be at risk. Synonyms. McGillivray (1973: 344) identifled/. eriocladus Gand. and/, ovoideus Gand. as synonyms of / sphaerocephalus after examining type material, which in both cases he considered to be holotypes. In the case of / eriocladus, it appears likely that Gandoger saw only material of one kind, and only on a single sheet now housed at FY, which can therefore be assumed to be the holotype. However, for / ovoideus McGillivray stated ‘The type sheet bears two specimens, one in flower (on the right), the other in fmit. ’ In this case the type material possibly should be regarded as two syntypes rather than a holotype. The protologue of / drummondii Hiigel ex Jacques (Jacques 1843) describes the leaves as sparsely distributed, undivided, linear, sessile, 60-150 mm long [as 6-15 cm] and somewhat obtuse with a rather long, white point at the apex. This description of the leaves matches that given above for / sphaerocephalus although the colour of the leaf point is usually dark. However, the tip is sometimes pale and sometimes covered by appressed whitish hairs. It is evident from the original text that the description of / drummondii Hiigel ex Jacques was based on glasshouse-grown specimens. In addition to giving the native country for each as ‘laNouvelle-Hollande’, Jacques (1843: 217) stated that this was one of six species he had described that were cultivated by Monsieur Martine’s establishment. Apparently the plants of / drummondii were derived from a plant catalogue from HiigeTs garden and nursery business at Hietzing near Vienna, Austria (‘EUGEF cat. a.’, interpreted here as HiigeTs annual catalogue) under the manuscript name / "Drumundif (Jacques 1843: 216). In view of the epithet, it may be assumed that the original source of seed was collected by James Drummond from south-western Australia. As is typical for Jacques’s names for Australian plants, there is no known herbarium specimen that can be connected with his / drummondii. We are aware of only one case of a specimen associated, albeit indirectly, with a Jacques name, that of the doubtful name Acacia semperflorens Jacques (see Maslin 2001; O’Feary 2007); that specimen is housed at the Florence Herbarium (FI) and came from the same garden (Jacques 1837) that was cited in the description of / drummondii Hiigel ex Jacques. In response to our enquiries, we were informed that there are no relevant specimens of Isopogon at FI (C. Nepi, pers. comm.). Despite the lack of any original material, we are confident that the protologue B.L. Rye & T.D. Macfarlane, A new name and a new subspecies for Isopogon (Proteaceae) 315 is sufficient to show that it is not the same species as I. drummondii Benth.. Notes. The diaspore and seed measurements given above are based on diaspores from only a few specimens. From the available samples it seems likely that subsp. lesueurensis tends to have larger diaspores than subsp. sphaerocephalus but further fruiting material is needed to check this. b. Isopogon sphaerocephalus subsp. lesueurensis Rye, subsp. nov. Typus\ Mount Lesueur area, Western Australia [precise locality withheld for conservation reasons], June 1931, C.A. Gardner s.n. {holo\ PERTH 03439380). Shrubs 0.5-1.5 m high, probably lignotuberous. Flowering branches densely leafy, densely or very densely hairy. Leaves 60-100 mm long, 7-18 mm wide, densely or moderately densely hairy. Tepals 13-15 mm long. Pollen presenter 3.5-5 mm long, with finger-like papillae 0.2-0.25 mm long on the basal half of presenter; pedestal 1.5-2 mm long; swelling with long papillae on both sides or rarely just on one side and with short papillae on the other side, also often with papillae extending upwards for a short distance onto the base of the receptor. Diaspores 4-5.3 mm long, 1.8-2.4 mm wide; largest hairs 6-7 mm long; seed c. 3.2 x 2.2 mm. Diagnostic features. Distinguished from subsp. sphaerocephalus mainly by its longer pollen presenter (3.5-5 mm cf. 2.3-3.3 mm long), including a pedestal 1.5-2 mm long {cf. 0.4-1 mm long) and with long papillae on the lower half including the swelling above the pedestal {cf. long papillae extending for less than half the length of the pollen presenter, with the swelling usually glabrous or just with a few short papillae at the base). Plants of this subspecies also tend to be more hairy on their vegetative parts and to have more densely arranged, broader leaves (7-18 mm cf. 4-12 mm wide). Selected specimens examined. WESTERN AUSTRALIA [localities withheld for conservation reasons]: 25 Aug. 1938, W.E. Blackaineil (PERTH); June 1931, C.A. Gardner s.n. (PERTH); 27 Aug. 1979, E.A. Griffin 1980 (PERTH), 26 Sep. 1976, R. W. Johnson 3285 (BRI n.v., PERTH). Distribution and habitat. Occurs in lateritic soils, recorded on the top of one hill and from the slopes and base of mesas, in the Mt Lesueur area (Figure IB), north-east of Jurien Bay in the far south of the Geraldton Sandplains bioregion of Western Australia. No details of the vegetation are given but presumably this subspecies occurs in the shrublands with a high species diversity that characterise the lateritic uplands of the Mt Lesueur area. Subspecies lesueurensis occurs in a drier habitat and is separated by a disjunction of about 180 km from the northernmost locality known for subsp. sphaerocephalus. Phenology. Flowers recorded from June to early November and fruits continuing to February. Conservation status. To be listed under Conservation Codes for Western Australian Flora as Priority Two (A. Jones pers. comm). This subspecies has a very restricted range, being known only from three hills over a distance of a few kilometres. Etymology. Named after the Mt Lesueur area where the taxon occurs. This area has a high biodiversity and includes several other endemic taxa, such as Grevillea batrachioides McGill, and Hypocalymma tenuatum Strid & Keighery. Common name. Lesueur Isopogon. 316 Nuytsia Vol. 30 (2019) Notes. The earliest collection of this subspecies was made by Charles Gardner in June 1931. One of Gardner’s specimens (PERTH 01904957) bears the manuscript name 1. longifolius var. Nillosd’ and another gathering was recorded in his collecting book as I. sp. off. sphaerocephalus but he made no mention of the taxon in his unpublished flora treatment*. Foreman (1995: 218) described this variant of I. sphaerocephalus as ‘a broad, hairy leaf form’. One flowering specimen (PERTH 03418626) labelled as having been collected by Gardner from Mt Eesueur in January 1940 is of the typical subspecies and does not match a second specimen labelled with the same details (PERTH 01904957), which is in fruit. There appears to have been a mislabelling of the former specimen. Perhaps the confusion in the labelling of specimens contributed to Foreman’s (1995: 218) note that the Mt Eesueur variant seemed to intergrade with the typical variant. Although some of the individual characters may show some overlap between the two subspecies, possibly mainly because of seasonal variation in most cases, there are no known specimens that cannot be conclusively identifled to the subspecies level. Acknowledgements Digital images of types not available at PERTH were examined using JSTOR’s Global Plants database (https://plants.jstor.org/). We are grateful to Dr Chiara Nepi (University of Florence) for her advice regarding specimens at FI, the Collections staff at MEE for information and specimen scans, Anna Monro and Kelly Shepherd for their helpful comments on the manuscript, and Steve Dillon for assistance with the distribution maps. References Bentham, G. (1870). Proteaceae. Flora Anstraliensis. Vol. 5, pp. 315-584. Foreman, D.B. (1995). Isopogon. Flora of Australia. Vol. 16, pp. 194-223,451-453,479-481. (Australian Biological Resources Study: Canberra.) Gardner, C.A. (1952). Flora of Western Australia. Vol. 1, part 1. Gramineae. (Government Printer: Perth.) International Documentation Centre. (1962). Candolle Prodromi Herbarium. Herbariumgenevense. IDC microform collection number 800/2. (International Documentation Centre: Tumba, Sweden.) Jacques, H.A. (1837). Suite de la revue des genres de vegetaux cultives en France. Suite des Acacies. 2° Division. Annales de Flore et de Pomone; ou Journal des Jardins et des Champs 1837-38: 33-48. Jacques, H.A. (1843). Plantes rare ou peu connu. Annales de Flore et de Pomone; ou Journal des Jardins et des Champs 1842-1843, ser. 2, 1:215-221. Maslin, B.R. (2001). Mimosaceae, AcaciaPart2. Flora of Australia. Vol. 1 IB. (Australian Biological Resources Study: Canberra.) McGillivray, D.J. (1973). Michel Gandoger’s names of Australian plants. Contributions from the New South Wales National Herbarium 4(6): 319-365, plates XXVIII-XXXI. Meisner, C.D.F. (1845). Proteaceae. In’. Lehmann, C. (ed.) Plantae Preissianae. Vol. 1. pp. 491-601. (Meissneri: Hamburg.) Meisner, C.D.F. (1856). Proteaceae. In’. Candolle, A.P de (ed.) Prodromus systematis naturalis regni vegetabilis. Vol. 14: 209-492. (Victoris Masson: Parishs [Paris].) O’Leary, M.C. (2007). Review of Acacia retinodes and closely related species, A. uncifolia and A. provincialis (Leguminosae: Mimosoideae: sect. Phyllodineae). Journal of the Adelaide Botanical Gardens 21: 95-109. Rye, B.L. (1987). Proteaceae. Flora of the Perth Region. Vol. 1, pp. 309-357. (Western Australian Herbarium, Department of Agriculture: Western Australia.) Smith, M.G. & Jones, A. (2018). Threatenedand Priority Floralist 5 December 2018. Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 24 June 2019]. *This was prepared during the 1930s for his proposed Flora of Western Australia. However, only the first part of Volume One, on grasses (Gardner 1952), was ever published. Nuytsia The journal of the Western Australian Herbarium 30:317-334 Published online 10 December 2019 Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae), with validation of six species names and two new combinations Neville G. Marchant Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 Abstract Marchant, N.G. Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae), with validation of six species names and two new combinations. Nuytsia 30: 317-334 (2019). An updated description of the genus Chamelaucium Desf compiled from all currently recognised formal and informal taxa is presented to establish the status of Chamelaucium at generic level within the Myrtaceae, tribe Chamelaucieae. Descriptions are provided for C. erythrochlorum N.G.Marchant, C. floriferum N.G.Marchant, and its two subspecies, C. /w///zrrzzN.G.Marchant, C. orar/ww N.G.Marchant, C. roycei N.G.Marchant, and C. xanthocladum N.G.Marchant. New combinations are made and descriptions provided for C. forrestii (F.Muell.) N.G.Marchant, and C. repens (A.S.George) N.G.Marchant. Precise localities for all taxa have been withheld for conservation reasons. Introduction Chamelaucium Desf is one of a group of 37 currently recognised genera placed in the Myrtaceae tribe Chamelaucieae DC. (Rye et al. in press). Phylogenetic and taxonomic studies on member genera are ongoing by various authors, especially in Actinodium Schauer, Chamelaucium, Darwinia Rudge, and Verticordia DC. Current phylogenetic studies by others recognise that of these four genera only Chamelaucium is best treated as a discrete genus as currently circumscribed (B.L. Rye, pers. com. 5 March 2019). The present paper provides a generic description of this genus recognising it as a distinct entity in the tribe Chamelaucieae, endemic to south-western Australia, and currently comprised of around 40 described and undescribed species. Six species are formally validated here and two new combinations are made. All taxa have conservation priority, including two being currently assigned ‘vulnerable’ status. Methods Descriptions of new taxa have been developed from PERTH herbarium specimens, and from field observations. Measurements were made using rehydrated fiowers and leaves. Relevant type material has been studied at CGE, K, MEE and PERTH, and other specimens have been viewed on Global Plants (https://plants.)stor.org/). © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 318 Nuytsia Vol. 30 (2019) Circumscription of Chamelaucium Desf. Taxonomic history The generic name Chamelaucium dates from the publication of two species by Desfontaines (1819a). Sprengel (1825) adopted the variant spelling Chamaelaucium which was taken up by Candolle (1827, 1828) and used in later literature, even after the original spelling had been pointed out by Bibby (1944). Desfontaines (1819a) based the genus on C. ciliatum Desf and C. plumosum Desf; subsequently, in the same volume, he added C. brownii Desf (Desfontaines 1819b). The latter two species were later transferred to Verticordia. Chamelaucium sensu Desfontaines was characterised by membranous, caducous bracteoles, and either prominently ciliate calyx lobes (in C. ciliatum) or by each calyx lobe being deeply divided into ciliate segments (C. brownii and C. plumosum). In 1828 A.P. de Candolle described Verticordia in which he placed C. plumosum (as V. fontanesii DC.) and C. brownii. He retained C. ciliatum in Chamelaucium which he distinguished from Verticordia by the lower degree of division of the calyx lobes, the included style, the form of the stigma, and absence of substigmatic hairs. Chamelaucium ciliatum Desf can thus be regarded as the type species of Chamelaucium. This species is extremely morphologically variable and unlike almost all other members of the genus it has prominently ciliate calyx lobes and a glabrous style, both characters resembling those of some species of Verticordia. Endlicher (1838) and Schauer (1844) each added another species of Chamelaucium (C. virgatum Endl. and C. uncinatum Schauer), both characterised by large flowers with entire, short, broad calyx lobes and entire petals. Turczaninov, working on collections of James Drummond, described the genus Decalophium Turcz. (Turczaninov 1847), listing one species, D.pauciflorum Turcz. In 1852, he published a detailed description of this species as well as another five species of Decalophium (Turczaninov 1852). Bentham (1867) transferred D. pauciflorum to Chamelaucium, and added four more species to those already published; he was, however, unaware of Turczaninov’s 1852 paper, and it was not until Domin (1923) that the names contained in that paper were brought to the attention of botanists with the publication of several new combinations, including C. micranthum (Turcz.) Domin, based onDecalophium micranthum Turcz. Taxonomic confusion between the gmoraChamelaucium rndDarwiniapartly dates from Turczaninov’s description of GenetyllispaucifloraJmcz. (Turczaninov 1849). Turczaninov later transferred this species to his Decalophium (Turczaninov 1852); however, because he had already used the same epithet under that genus name in 1847, he provided the new name Decalophium rugulosum Turcz. Genetyllis had been established by Candolle in 1828, and was later regarded by Bentham (1865,1867) as a section of Darwinia Rudge, referring only to south-western Australian species. Further nomenclatural confusion resulted from the publication of the Systematic Census of Australian Plants (Mueller 1882), in which Mueller transferred ten species of Chamelaucium to Darwinia, and because the same epithets had been used for different taxa in the two genera, new species names were created. Although Mueller’s inclusion of Chamelaucium in Darwinia was not recognised by Gardner (1931), nor subsequently by Blackall (1954) or Blackall and Grieve (1980), the difference between the two genera was not clear. This is exemplified by the number of misidentiflcations on herbarium N.G. Marchant, Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae) 319 specimens, and the publication of two species of Chamelaucium as Darwinia, most likely on the basis of their exserted styles; the recombinations are made below. Generic description Chamelancinm Desf, Mem. Mus. Hist. Nat. 5: 34^4 (1819); A.R de Candolle, Prodr. 3: 209 (1828). Type species'. C. ciliatum Desf Decalophium Tmcz., Bull. Soc. Imp. Naturalistes Moscou 20: 153 (1847). Type species'. D. pauciflorum Turcz. Shrubs up to 4 m high, widely spreading, much branched or often slender with a single lower stem, sometimes prostrate or nearly so. Leaves 2-30 mm long, opposite decussate, or alternate, sometimes aggregated at ends of branchlets, often borne in fascicles on short lateral shoots; linear-triquetrous, narrowly obovate to lanceolate, or ovate to elliptic, adaxial surface frequently grooved, abaxial surface flat or ± convex; margins entire or serrulate, or densely ciliate with simple or plumose hairs, rarely hyaline; apex truncate, acuminate or mucronate, sometimes minutely bifid; leaf scars persistent, usually very prominent; floral leaves undifferentiated, or shorter and broader, sometimes with a hyaline margin, or rarely ciliate with simple or plumose hairs. Flowers solitary or in lax or dense axillary or terminal racemes, or dichasiaf Bracteoles 0.5-7 mm long, brown or reddish, caducous or persistent, rarely connate at the base and retained to fruiting, cucullate or deeply concave and spathulate, usually umbonate on the outside at the highest part, the umbo of the inner bracteole often less prominent; umbo (when present) very prominent and curved, hom-like or barely developed. Floral tube 2-8 mm long, turbinate, campanulate or obconic to cylindric; lower part adnate to ovary, 1.5-5 mm long, 5 to 10-ribbed, rugose, or smooth, frequently 10-foveolate, upper part free from ovary, 0.5^ mm long, smooth or with 10 or more ribs, or rugose, glands less prominent. Calyx lobes 5,0.5-3.5 mm long, usually under 2 mm long, green, white, or reddish, semicircular, oblong, ovate, spathulate or triangular, entire, ciliate, ciliolate, crenate or irregularly dentate; sinuses narrow to wide, rarely absent. Petals 5,2-7 mm long, white, often aging red, pink, purple or yellow-green, mostly obovate, ovate, or orbicular, concave, entire, ciliolate or irregularly dentate. Stamens 10, rarely up to 13, anthers opening by longitudinal slits; staminodes 10, rarely 12, less than 0.5 or up to 3 mm long, linear or spathulate, often divided at the tip into two uneven segments. Style 2-24 mm long, included, shortly exserted, or long exserted, usually dilated in lower part, most species with substigmatic hairs, which are most commonly in a band subtending the stigma. Ovules 4-10. Fruit a one-seeded nut, encased by the dried, lower part of the floral tube. 'Briggs and Johnson (1979) regard the inflorescence of Chamelaucium as either a monad, or corymbiform, racemiform or a spiciform conflorescence, or a superconflorescence. Diagnosticfeatures. Bracteoles cucullate and deeply concave and spathulate, not keeled, almost always with an umbo arising from the highest part of the outer bracteole. Flowers are never subtended by an involucre. The 5 calyx lobes and 5 petals are never deeply divided. The 10 anthers dehisce by two longitudinal slits, and often have a large connective. There are 10 staminodia. The stigma is usually subtended by a narrow band of hairs, and the ovule numbers range from 4 to 10. Size and distribution. The genus comprises around 40 formally described and undescribed species, and is confined to the south-west of Western Australia where it occurs in various coastal and inland habitats from the lower Murchison to east of Esperance, with a few species associated with granitic monadnocks in the south-western forests. A number of species commonly occur in sandy and rocky soils in kwongan vegetation. 320 Nuytsia Vol. 30 (2019) Etymology. Desfontaines (1819a) coined the generic name from the strong resemblance of the bracteoles, especially the outer bracteole, to a cape or hood. A late antique Greek word for a type of cap worn by court officials, or the hood of a monastic habit, is a KapeJ^auKiov, i.e. kamelaukion (Sophocles et al. 1900). In the type species C. ciliatum Desf, the bracteoles are brown and closely resemble the hood or cape of a medieval monk. An additional monastic hood-like character is the erect or curved, cone- shaped nipple, referred to here as an ‘umbo’, borne on the highest part of the bracteoles of C. ciliatum Desf, and most other species of the genus. New taxa and new combinations Chamelancinm erythrochlornm N.G.Marchant, sp. nov. ^/>c:Yoongarillup [precise locality withheld for conservation reasons]. Western Australia, 19November 1980, G.J. Keighery 3635 {holo\ PERTH 04997158!; iso\ PERTH 012575601). Chamelaucium sp. Yoongarillup (G.J. Keighery 3635), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. Chamelauciumerythrochloruml^.G.Maxchmt&KQighQryms; G.Paczkowska&A.R. Chapman, West. Austral. FI : Descr. Cat 355 (2000); Western Australian Herbarium, in FloraBase, https://florabase. dpaw.wa.gov.au/ [accessed 6 June 2019]. Shrub, erect, 0.5-E5 m high, rarely more. Leaves opposite decussate, linear, 15-23 mm long, 0.7- 0.8 mm wide, 0.8-1 mm deep; adaxial surface deeply grooved; margins entire, rarely serrulate; apex acuminate-mucronate; petioles 0.3-1 mm long, appressedto stem; decurrences prominent, E5-3 mm long. Flowers solitary in leaf axils forming open racemes of 1-15 flowers on young branches, 3.5-5 mm diam.; disc 2.5^ mm diam.; pedicels 1.5-5.5 mm long, glandular, with two tiny auricles at the top, representing decurrences of the bracteoles. Bracteoles 2-2.5 mm long; umbo prominent, slender, erect to incurved to almost horizontal, to 1.5 mm long. Floral tube green, obconic-turbinate 5.5-8 mm long; lower part of floral tube 2.5^ mm long, strongly 10-ribbed, rugose, glandular, foveolae not obvious; upper tube 3^ mm long, sometimes clearly 10-ribbed, frequently ribs indistinct. Calyx lobes red-pink, triangular-ovate to oblong with a triangular-rounded apex, 1.5-2.5 mm long; margins ciliate to shallowly lacerate; sinuses narrow. Petals bright red, erect, obovate-broadly obovate, concave, ± hooded, with a narrowed base adnate to the staminal tube, 4.5-6.5 mm long; margins ciliate to shallowly lacerate. Stamens in a tube 0.2-1 mm long, the sepaline stamens set slightly lower than the petaline; filaments narrowly oblong, slightly broader at the base, 0.5-1 mm long; anthers 0.5-0.6 mm diam., connective darker, red-brown, swollen; staminodes narrowly oblong, sometimes slightly wider towards the base, 0.8-E8 mm long, adaxial surface frequently slightly concave so that the expanded apex is shortly hooded; apex shortly lobed, one lobe frequently longer (to 0.3 mm) and darker coloured. Style well exserted, 16-24 mm long; stigma barely broader than the upper part of the style, convex, 1 mm diam., subtended by a band of hairs; substigmatic hairs Aliform, 0.5-0.6 mm long. Ovules 6-10. Diagnostic features. Tall spindly shrub with a single stem at the base. The floral tube is bright green and the corolla red and the style is well exserted. Selected specimens examined.V\fFW^WBAAA\JW^BJkLlA\ [localities withheld for conservation reasons] 7 Jan. 1997, N. Casson & B. Evans SC137.9 (PERTH); 13 Dec. 2001, R.J. Cranfield 17674 (PERTH); 12 Nov. 1957, H.F. Dawson s.n. (PERTH); 26 Nov. 1975, A.S Ceorge 14219 (PERTH); 6 Feb. 2002, N.G. Marchant, Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae) 321 J Liddelow s.n. (PERTH); 20 Dec. 1978, N.G. Marchants.n. (PERTH); 7 Feb. 1989, G. S. McCutcheon 1994(PERTH); lODec. 1985,5. 19(PERTH);28Dec. 1976,RZ. 76040(PERTH);21 Jan. 1993, G. Wardell-Johnson Mil (PERTH). Phenology. Flowers from November to February. Distribution and habitat. Restricted to remnant localities on the foothills and more widespread in the Whicher Range scarp and nearby jarrah forest and jarrah marri woodlands. Grows on lateritic soil, pale sandy clay, or loam soils. Conservation status. Currently listed as Priority Four under Conservation Codes for Western Australian Flora. Fisted (Smith & Jones 2018) under the name C. sp. Yoongarillup (G.J. Keighery 3635). Etymology. From Greek words erythros for red, and chloros for green, referring to the bright red corolla and bright green floral tube. Chamelaucium floriferum N.G.Marchant, sp. nov. Type: Walpole Nornalup National Park [precise locality withheld for conservation reasons]. Western Australia, 10 November 1976, N.G. Marchant s.n. {holo\ PERTH 06763324). Chamelaucium floriferum N.G.Marchant & Keighery ms; G. Paczkowska & A.R. Chapman, West. Austral. FI : Descr. Cat. 355 (2000); Western Australian Herbarium, in FloraBase, https://florabase. dpaw.wa.gov.au/ [accessed 6 June 2019]. Shrub, erect, 0.6-3 m high. Leaves distinctly opposite decussate, vertically oriented or spreading, linear, ± triquetrous, 7-20 mm long, 0.5-0.9 mm wide, 0.6-1 mm deep; adaxial surface usually deeply grooved; apex usually acute, sometimes finely bifid, rarely obtuse; petioles 0.4-1 mm long, usually appressed to stems; decurrences usually paler coloured than stems, 0.6-6(-12) mm long Flowers solitary in upper axils forming racemes of 2-12(-16) flowers, 5.5-8 mm diam. at maturity; disc 2-2.4 mm diam.; pedicels 2-10 mm long, erect or spreading. Bracteoles 2-3.3 mm long, rarely longer; umbo usually prominent, incurved, narrowly to broadly conic, and laterally compressed, 0.5-0.9(-1.6) mm long, or only 0.1-0.3 mm long, or obscure. Floral tube narrowly campanulate to broadly campanulate, glandular, shallowly 10-ribbed or ribbing sometimes obscure, shallowly rugose, shallowly pitted or ± smooth, 2.5- 4.5 mm long; lower part of tube 1.3-3.4 mm long, with or without 10 shallow foveolae on the upper part; upper part of tube 0.8-1.4 mm long. Calyx lobes broadly triangular-deltate to shallowly triangular, 0.5-0.8 mm long; margins usually entire, sometimes irregularly serrulate-serrate near the apex; sinuses wide. Petals white, pink, or reddish ± erect-spreading; concave, broadly obovate-orbicular, 2.6-3.5(- 4.5) mm long; margins entire or finely uneven to shallowly crenate, sometimes with a few scattered serrations. Stamens in a tube 0.2-0.8 mm long, with filaments and staminodes borne at the same level; filaments usually narrowly triangular-triangular, or sometimes widening at the base and linear above, 0.5-1.5 mm long; anthers 0.4-0.5 mm diam.; connectives swollen; staminodes narrowly triangular or linear, sometimes slightly dilated in the subapical region, 0.5-1 mm long; apex acute, reddish. Styles shortly exserted, usually thickened at the base, 2.5-3.5(-4.5) mm long; stigma dilated, hemispherical- subglobose, 0.3-0.4 mm diam. and subtended by a band of hairs, but sometimes globose and papillose; substigmatic hairs (when present) reflexed. Aliform, 0.3-0.4 mm long. Ovules 7-9. Etymology. The name is derived from Fatin flos, a flower, and ferre, bear, an adjectival form implying many flowers. 322 Nuytsia Vol. 30 (2019) Notes. Two subspecies are recognised. Key to subspecies 1. Compact shrub; leaves vertically oriented, light green, rigid; pedicels 2-7.5 mm long, as short as flower, or slightly longer than the flower.C. floriferum subsp. floriferum 1: DiflUse-branched shrub; leaves mostly horizontally oriented, dark green, flaccid; pedicels 6-10 mm long, longer than the flower. C. floriferum subsp. diffusum Chamelaucium floriferum subsp. floriferum Shrub to 0.2 m high, with a distinctly trullate outline. Leaves vertically oriented, 7-15 mm long, rigid, light green. Flowers crowded in upper axils, as long as subtending leaves, slightly shorter or sometimes exceeding leaves; pedicels 2-7.5 mm long, as short as flower or slightly longer. Petals white, pink or reddish. Diagnostic features. Floriferous pine-like shrub with rigid, vertically oriented, light green leaves. Pedicels as short as the flower or slightly longer. Flowers white, pink or reddish, crowded in upper axils. Selected specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 18 Dec. 1990, A.R. Annels 1513 (PERTH); 6 Oct. 2007, M. Crowhurst 173 (PERTH); 26 Nov. 1997, E.D. Middleton 103 (PERTH); 9 Oct. 2005, E.D. Middleton 739 (PERTH); 7 Dec. 1986 A.5. Weston 15258 (PERTH); 22 Sep. 1992, J.R. Wheeler 3253 (PERTH). Phenology. Flowers between September and December. Distribution and habitat. Found in near-coastal areas from just west of Point Nuyts to Mt Hopkins, in open coastal heath, and thickets, on sandy or loam soils between granitic outcrops or on sandy soil over limestone above granitic country rock. Conservation status. Recently up-graded from Priority Three to Priority Two under Conservation Codes for Western Australian Flora. Eisted under the erroneous informal name C. sp. Walpole (PG. Wilson 6318); see note under C. floriferum subsp. dijfusum below [accessed 24 June 2019]. Notes. This subspecies is widely cultivated and is known as ‘Walpole Wax’. Artificial hybrids have been created with C. uncinatum. It is restricted to a few hundred metres from the coast and may be subjected to salt spray. It retains its form and rigid leaf characters in cultivation. Some specimens from the eastern part of the range, a short distance inland near Mt Hopkins, resemble C. floriferum subsp. dijfusum. Chamelaucium floriferum subsp. diffusum N.G.Marchant, subsp. nov. Type'. Walpole Nomalup National Park [precise locality withheld for conservation reasons]. Western Australia, 12 November 1976, N Marchant 76/125 {holo\ PERTH 06796443). Chamelaucium sp. Nornalup (N.G. Marchant 76/125), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. N.G. Marchant, Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae) 323 Chamelaucium sp. Walpole (RG. Wilson 6318), Western Australian Herbarium, in FloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. This specimen is not the typical subspecies. It is C. floriferum subsp. dijfusum., see Notes below. Chamelaucium floriferum subsp. dijfusum N.G.Marchant & Keighery ms; G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. 355 (2000); Western Australian Herbarium, inFloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. Shrub to 3 m high, with diffuse branching. Leaves usually at right angles to stem, 15-20 mm long, flaccid, dark green, widely spaced. Flowers in open racemes; pedicels up to 10 mm long, longer than flowers. Petals white or pale pink. (Figure 1) Diagnostic features. A many-flowered, open diffuse-branched shrub, not pine tree-like or trullate in outline. Leaves usually at right angles to stem and generally distantly spaced. Pedicels longer than flowers, up to 10 mm long. Selected specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 25 Nov. 1985, A.R. Annels 1870 (PERTH); 7 Jan. 2003, J.A. Cochrane 4530 (PERTH); 23 Oct. 2007, M. Crowhurst 136 (PERTH); 7 Nov. 2007, M. Crowhurst 180 (PERTH); 28 Nov. 2000, G. Freebury 35 (PERTH); 7 Aug. 1980, N.G. Marchant 80/65 (PERTH). Phenology. Peak flowering is between August and December; some populations may flower between January and May. Distribution and habitat. Sandy loam apron soils of granitic rocks, away from the coast. Populations are scattered and associated with granitic monadnocks from Kent River near Denmark, westwards to south east of Northcliffe. Conservation status. Currently listed as Priority Two under Conservation Codes for Western Australian Flora (Smith & Jones 2018), under the name Chamelaucium sp. Nomalup (N.G. Marchant 76/125), [accessed 24 June 2019]. Etymology. The subspecies name refers to the diffuse branching habit. Notes. The specimen used to represent the temporary name C. sp. Walpole (PG. Wilson 6318) is C. floriferum subsp. dijfusum. It was collected at the type locality for this subspecies, an inland locality, and not the near-coastal habitat of the typical subspecies. Chamelaucium forrestii (F.Muell.) N.G.Marchant, comb. nov. Darwinia forrestii F.Muell, Fragm. 11:9 (1878). Type\ In rupibus graniticis ad summitatem mentis. Burrobunup prope flumen Gordon’s River [Western Australia], December 1877, F. Mueller s.n. (lecto here designated: MEE2290893!; isolecto: CGEl, GH 000689811, MEE 7265091, MEE7265101, MEE 22908941, PERTH 06796427!). Chamelaucium sp. Mt Frankland (A.S. George 11117), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. 324 Nuytsia Vol. 30 (2019) Figure 1. Chamelauciumfloriferum subsp. A-flowering leafy shoot and close-up; B-leaf, side view and cross section; C - flower bud surrounded by overlapping bracteoles, each with an umbo; D - flower bud, bracteoles removed; E - flower at anthesis; F - flower from above; G - flower in longitudinal section; H - staminodia and stamens with front and side view of anthers; I - style showing band of substigmatic hairs. Scale bars = 1 mm. Drawn by Donna Terrington from N.G. Marchant 80/65 (PERTH 01170864). N.G. Marchant, Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae) 325 Chamelaucium forrestii (F.Muell.) N.G.Marchant & Keighery ms; G. Paczkowska & A.R. Chapman, West. Austral. FI.: Descr. Cat. 355 (2000); Western Australian Herbarium, inFloraBase, https ://florabase. dpaw.wa.gov.au/ [accessed 6 June 2019]. Shrub, erect, up to 2 m high, sometimes broadly spreading up to 2 m across. Leaves opposite decussate, erect, dark green, linear, 10-24 mm long, 1-1.5 mm diam., somewhat fleshy; adaxial surface deeply grooved; margins entire; apex truncate with two, minute, lateral mucrones; petioles to 1 mm long, appressed to stem; decurrences pale coloured, prominent. Flowers solitary in upper leaf axils, forming open racemes of 1-15 flowers on young branches, 3^ mm diam.; disc 2.5-3 mm diam.; pedicels 3-6 mm long, deflexed so that flowers are well clear of the erect leaves. Bracteoles 4-5 mm long, 3mm across, deeply cucullate; umbo acuminate 0.8-1 mm long. Floral tube narrowly obconic, 3.5-5 mm long; lower part of floral tube 2.5-3.5 mm long, ribbed, rugose, foveolae 10; upper part of floral tube 1-1.5 mm long, ribs not prominent, intervening surfaces finely pitted. Calyx lobes broadly triangular, 1-1.5 mm long, 1.5-1.7 mm across; margins minutely serrate, apex angular. Petals cream coloured, tinged green, erect, obovate-broadly obovate, deeply concave, 2.5-3.5 mm long; margins entire. Stamens in a tube 0.5 mm long, filaments narrowly triangular, 0.5-0.6 mm long; anthers 0.3-0.5 mm diam., connective thickened; staminodes narrowly oblong, slightly broader in upper part, 0.3-0.5 mm long, longer than the stamens; apex obtuse. Style exserted, sometimes well exserted, 4-6.5 mm long; stigma convex, 0.4— 0.6 mm across, subtended by a band of somewhat reflexed, straight hairs, 0.3-0.4 mm long. Ovules 4-6. Diagnostic features. Leaves dark green. Style excluded. Flowers on deflexed pedicels clear of the erect leaves. Floral tube foveolate. Distribution and habitat. The species grows on granitic monadnocks in high rainfall] arrah and karri forest. Etymology. The epithet commemorates John Forrest, who as an explorer collected specimens for von Mueller. Notes. The type locality, Burrobunup, is an early name for Granite Peak. Selected specimens examined. WESTERN AUSTRAEIA: [localities withheld for conservation reasons] 3 Oct. 1971, A.S George 11117 (PERTH); 21 July 2009, J. Liddelow 81 (PERTH); 17 June 2010, J. Liddelow 132 (PERTH); 3 Oct. 1971, A.G. Marchant 16I\U (PERTH); 9Nov. 1976, AG Marchant 76/114 (PERTH); 15 June 1997, E.D. Middleton 34 (PERTH); 30 Sep. 1997, E.D. Middleton 52 (PERTH); 7 Nov. 1998, E.D. Middleton 186 (PERTH); 13 Oct. 1976, B.L. Rye 76009 (PERTH); 10 Nov. 1990, LB. Wheeler AZ (PERTH). Phenology. Peak flowering is between July and November, also recorded as early as May and as late as January. Distribution and habitat. Associated with granitic monadnocks in humic sandy and loamy soils in high rainfall jarrah and karri forests in the Mount Frankland district. Conservation status. Recently up-graded from Priority Three to Priority Two under Conservation Codes for Western Australian Flora. Eisted under Chamelaucium sp. Mt Frankland (A.S. George 11117), [accessed 24 June 2019]. 326 Nuytsia Vol. 30 (2019) Chamelaucium lullfitzii N.G.Marchant, sp. nov. Type', between Muchea and Gingin [precise locality withheld for conservation reasons], Western Australia, 16 September 1987, N. G. Marchant 6 {holo\ PERTH 07576196; iso\ CANB, K, MEL, NSW). Chamelaucium sp. Gingin (N.G. Marchant 6), Western Australian Herbarium, in FloraBase, https:// florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. ChamelauciumlullfitzUl^.G.MarchantScKQighQryms; G. Paczkowska&A.R. Chapman, West.Austral. Fl.:Descr. Cat. 356(2000); WestemAustralianHerbarium, inF/oraRt/^e, https://florabase.dpaw.wa.gov. au/ [accessed 6 June 2019]. Shrub 100 to 200 cm high, with many erect branches bearing numerous, short axillary shoots 5-20(- 30) mm long. Leaves semi-spreading to spreading, mostly crowded on the numerous short axillary shoots, narrowly obovate-very narrowly obovate or linear-narrowly elliptic, or sometimes very narrowly elliptic, 5.5-11.5 mm long, 1.2-1.4 mm wide, 0.6-0.8 mm deep; adaxial surface flat; abaxial surface convex, rounded or angular, plano-convex to triquetrous; margins entire; apex mucronate; petiole 0.5- 1.5 mm long, appressed; decurrences prominent; floral leaves flattened, ovate-triangular-oblong, 1.3- 2.2 mm long, 0.9-1.3 mm wide; adaxial surface concave; margins rarely ciliate on the upper half; apex mucronate; decurrences strongly and abruptly raised but short, 0.2-0.6 mm long. Flowers 2-9 in small heads on axillary shoots and up to 20 in the clusters at the end of main branches, 6.5-9 mm diam.; disc 3^.2 mm diam.; pedicels 0.8-2 mm long. Bracteoles 3.5^.5 mm long; umbo incurved, narrowly conic-acicular, 0.3-1 mm long. Floral tube broadly or very broadly obconic to broadly or very broadly turbinate, 4.4-5.4 mm long; lower part of floral tube 2.8-3.4 mm long, not foveolate, shallowly 10-ribbed; upper part of tube 1.5-2.2 mm long, obscurely ribbed or smooth. Calyx lobes ± erect, ovate, glandular, 2-2.8 mm long; margins irregularly denticulate and ciliate, cilia 0.5-2 mm long; sinuses wide. Petals white, ± erect, broadly elliptic-broadly obovate, concave, 4.6-6 mm long; margins irregularly denticulate or very sparsely and finely ciliate. Stamens in a tube 0.6-1 mm long, erect or arching inwards; sepaline and petaline stamens similar; filaments narrowly triangular, 0.7- 1.1 mm long; anthers 0.4 mm long, connective not prominent; staminodes oblong-narrowly ovate, 0.9-1.3 mm long; apex obtuse or acute. Styles narrowly conic, 6.2-7.1 mm long, reaching to the top of petals; stigma strongly dilated resembling a swollen disc, finely papillate, 0.5-0.6 mm diam.; subtended by hairs from the base of the disc and directed downwards, hairs 0.4-1 mm long. Ovules 5-8. Diagnosticfeatures. Spindly erect, few-branched shrub with terminal leafy inflorescences of broad, white- petaled flowers when mature. Flowers on very short pedicels; floral tube broad; petals up to 6 mm long. Selected specimens examined. WESTERN AUSTRALIA: [localities withheld for conservation reasons] 22 Nov. 1995, J.A. Cochrane 1703 (PERTH); 9 Dec.1995, J.A. Cochrane 1813 (PERTH); 21 Nov. 2008, B. Fellows 1 (PERTH); 20 Oct. 2010, F & J. Hort 3700 (PERTH); 19 Oct. 2009, M. Hoskins 25 & F Felton (PERTH); 22 Sep. 1992, S. Patrick 1246 (PERTH^ Phenology. Flowers from September to December. Distribution and habitat. Restricted to a very small area associated with the Gingin scarp, south of Gingin. Plants grow on white, grey, or yellow sands in low open banksia woodland. Conservation status. Listed as Vulnerable in Western Australia (Smith & Jones 2018), under the name Chamelaucium sp. Gingin (N.G. Marchant 6) [accessed 24 June 2019]. N.G. Marchant, Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae) 327 Etymology. Named after George Lullfitz, native plant enthusiast who pioneered the widespread use of Western Australia native plants in landscaping and who first recognised this taxon as an undescribed species. Chamelaucium orarium N.G.Marchant, sp. nov. Type'. Cape Vancouver [precise locality withheld for conservation reasons], Western Australia, 18 September 1978, /. Abbotts.n. {holo\ PERTH 01259679!; iso\ CANBl, Kl, MEL!, NSW!). Chamelaucium sp. Cape Vancouver (B. Swainson & D. Davidson s.n. PERTH 1259660), Western Australian Herbarium, in FloraBase, https://fiorabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. Chamelauciumforrestii subsp. orar/MmN.G.Marchant&Keighery ms; G. Paczkowska&A.R. Chapman, West. Austral. FI.: Descr. Cat 355 (2000); Western Australian Herbarium, inFloraBase, https ://fiorabase. dpaw.wa.gov.au/ [accessed 6 June 2019]. Shrub, erect, 0.8-1.5 m high. Leaves opposite decussate, erect, linear, 10-27 mm long, mostly aggregated at ends of branches; adaxial surface grooved; margins entire; apex truncate with two, minute, lateral mucrones; petioles 1.5-2.5 mm long, appressedto stem; decurrences prominent, tinged yellow. Flowers solitary in upper leaf axils, erect, barely exceeding leaves, forming open racemes of 1-12 fiowers on young branches, 3.5-5 mm diam.; disc 2.5^ mm diam.; pedicels 1.5^ mm long, erect. Bracteoles 5 mm long, 3 mm across; umbo prominent, acuminate, up to 2.2 mm long, incurved. Floral tube narrowly campanulate, 5-7 mm long; lower part of floral tube 3^ mm long, 10-ribbed, ribs widely spaced, intervening surface distinctly rugose, foveoli absent; upper part of floral tube 1.5-2 mm long, ribs divided into three branches, intervening surfaces finely pitted. Calyx lobes triangular, 0.5-1 mm long and across; margins minutely serrate, apex rounded. Petals cream coloured, spreading at maturity, obovate-broadly obovate, shallowly concave, 3.5-5 mm long; margins entire. Stamens in a tube 0.3 mm long, filaments broader at the base, 0.5-1 mm long; anthers 0.5-0.6 mm diam., connective thickened; staminodes narrowly oblong, slightly wider atthe base, 0.6-1 mm long; apex obtuse, sometimes slightly widened. Style included 2-4 mm long; stigma convex, 0.4-0.7 mm diam., the base subtended by a band of crinkled hairs, 0.5-0.8 mm long. Ovules 6-8. Diagnostic features. Floral tube narrowly campanulate. Ribs widely-spaced, each dividing into three in upper part. Style included. Flowers shorter than leaves; pedicels erect, up to 4 mm long. Foveolae absent. Other specimens examined. WESTERN AUSTRALIA: [localities withheld for conservation reasons] Oct 1976, 1. Abbott s.n. (PERTH); 28 Dec 2003, J.A. Cochrane 4868, T. Friend Sc S. Hands (PERTH); 10Dec.l986,G.J.^^zg/zeO^8586(PERTH);28Oct. 1993,C.J.1189(PERTH); lOOct. 1984, B. Swainson & D. Davidson s.n. (PERTH). Phenology. Flowering recorded for September to December. Distribution and habitat. Only recorded for a single locality on sea cliffs east of Albany. Grows in low coastal heath, and tall shrub thickets on white or grey sand associated with granitic outcrops. Conservation status. Currently listed as Priority Two under Conservation Codes for Western Australian Flora (Smith & Jones 2018), under the name Chamelaucium sp. Cape Vancouver {B. Swainson & D. Davidson s.n. PERTH 1259660). 328 Nuytsia Vol. 30 (2019) Etymology. The subspecies name is derived from the Latin word ora, meaning of the coast. Chamelaucium repens (A.S.George) N.G.Marchant, comb. nov. DarwiniarepensA.S.GQorgQ,J Roy. Soc. WestemAustralia50\99(\967). Type. 15 miles east of Mingenew [WestemAustralia], August 1955, A.i?. Mains.n. {holo\ PERTH 01018671!; Ao: K000566466!, MEL 2290895!, PERTH 01018663!, PERTH 01018665!). Chamelaucium sp. Canna (G. Keighery s.n. PERTH 02236435), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. Chamelaucium repens (A.S.George) N.G.Marchant & Keighery ms; G. Paezkowska & A.R. Chapman, West Austral. Fl.iDescr. Cat 356(2000); WestemAustralianHerbarium, inF/oraR^z^e, https ://florabase. dpaw.wa.gov.au/ [accessed 6 June 2019]. Shrub, prostrate or almost so, branches to 0.1 m long, bearing erect branchlets 20^0 mm long, frequently rooting at the nodes. Leaves in terminal fascicles, opposite decussate, vertically oriented, sometimes tinged reddish, linear-narrowly clavate, subterete-terete, (3.5-)6-l 1 mm long, 0.4-0.6 mm wide; apex obtuse with two very fine lateral spine-like points on the adaxial surface; petiole slightly flattened and slightly concave adaxially 0.2-0.4 mm long, with 1-3 pairs of reddish brown, flattish, ovate-triangular persistent enations 0.1-0.25 mm long in the axils; floral leaves undifferentiated; decurrences completely covering younger parts of stems, 0.3-1.4(-3) mm long. Flowers solitary in upper leaf axils forming a loose cluster of 1-6 on erect branchlets, 2.6-4 mm diam.; disc 1.6-2.2 mm diam.; pedicels short and stout, 0.4-1 mm long. Bracteoles 2.2-3 mm long, margins entire; umbo incurved, narrowly conic, 0.2-0.8 mm long. Floral tube cylindric, shallowly to obscurely 10-ribbed, shallowly glandular-pitted, 3^.3 mm long; lower floral tube 1.5-2.3 mm long, without foveolae; upper tube 1.5-2.3 mm long. Calyx lobes erect, sparsely fine-glandular, broadly to very broadly ovate, 0.6-1.2 mm long; margins irregularly dentate and ciliate to shallowly incised; sinuses wide. Petals red, erect, concave, ovate-ovate elliptic, 2.6-3.6 mm long, 1.2-1.9 mm wide; margins entire to minutely undulate. Stamens in atube 0.1- 0.3 mm long; sepaline and petaline similar; filaments erect or sometimes bent inwards or ± geniculate, 0.3-0.6 mm long; anthers 3-lobed, with 2 larger lateral lobes and a smaller dorsal one, 0.35-0.55 mm diam.; connectives enlarged; staminodes erect or bent into floral tube, narrowly obtrullate-oblong, 0.3- 0.5 mm long. Style Aliform, long exserted, 12.5-22 mm long; stigma slightly dilated, subglobose, 0.2- 0.25 mm diam., papillose, subtended by a sparse whorl of hairs; substigmatic hairs 0.3-0.4 mm long, and mostly directed downwards, sometimes ± spreading. Ovules 6-8. Diagnostic features. The only recorded prostrate species of Chamelaucium ', leaves reddish, on short lateral branches. Flowers less than 5 mm long. Pedicels less than 1 mm long. Style up to 22 mm long, well exserted. Other specimens examined. WESTERN AUSTRALIA: [localities withheld for conservation reasons] 22Aug. 2005, J. Docherty 366 (PERTH); 23 Aug. 1990, G. J. Keighery s.n. (PERTH); no date,4.R. Main s.n. (PERTH). Phenology. Flowering only recorded for August. Distribution and habitat. Only recorded from a small area growing on lateritic breakaways. N.G. Marchant, Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae) 329 Conservation status. Currently listed as Priority One under Conservation Codes for Western Australian Flora (Smith & Jones 2018), under the name C. sp. Canna(G. Keighery w. PERTH 2236435), [accessed 24 June 2019], Etymology. The epithet is Latin for prostrate or creeping, referring to the habit. Chamelaucium roycei N.G.Marchant, sp. nov. Type: Busseltondistrict[preciselocahty withheld for conservation reasons]. Western Australia, 1 October 1954, R.D. Royce 4872 {holo\ PERTH 01224565!; iso\ PERTH 012245731). Chamelaucium sp. S coastal plain (R.D. Royce 4872), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. Chamelaucium roycei N.G.Marchant & Keighery ms; G. Paczkowska & A.R. Chapman, West. Austral. Fl.-.Descr. Cat. 356(2000); Western Australian Herbarium, inF/oraR^/^e, https://florabase.dpaw.wa.gov. au/ [accessed 6 June 2019]. Shrub to 120 cm high and 60 cm diam., branches tangled; brown to reddish. Leaves opposite decussate, widely spaced, usually spreading, narrowly oblong to linear, 4-10 mm long, 0.3-0.9 mm wide, sometimes triquetrous towards the apex, some leaves are ± flat-shallowly concave adaxially and are wider than deep; apex mucronate, sometimes acute-acuminate; floral leaves undifferentiated; petiole frequently appressed to stem, 0.2-0.5 mm long; decurrences 0.6-2(-6) mm long. Flowers usually in pairs in the upper leaf axils in a lax raceme with 1-12 flowers; 2.2-3 mm diam.; disc 1.4-2.4 mm diam.; pedicels (0.6-)l-2.8 mm long. Bracteoles 1.4-2.6 mm long; umbo prominent, ± erect, narrowly acicular, 0.6- 1.3 mm long. Floral tube narrowly obconic to narrowly turbinate, 3.9-5 mm long, sometimes slightly falcate, green, shallowly to obscurely 10-ribbed, glandular; lower tube 2.2-3.2 mm long, without foveolae; upper tube 1.6-2.2 mm long, frequently striate. Calyx lobes triangular-broadly triangular- ovate, sparsely glandular, 0.6-0.8 mm long; upper margins ciliate; sinuses wide. Petals white, obovate, shallowly concave, 2-3 mm long; margins ciliolate-serrate; apex rounded, sometimes ± truncate. Stamens in a tube, 0.2-0.7 mm long, with sepaline filaments borne slightly lower than the petaline; filaments narrowly triangular, 0.3-0.6 mm long; sepaline filaments leaning inwards, or curved outwards; in some flowers slightly larger than the petaline; petaline filaments ± erect; anthers 0.3-0.7 mm diam.; connectives enlarged, frequently with a concavity on the dorsal-abaxial surface; staminodes narrowly triangular, concave adaxially, 0.5-0.9 mm long; apex shallowly hooded, acute and curved slightly inwards or ± bilobed with the adaxial lobe longer than the abaxial. Style usually exserted, or sometimes barely so, 6-9 mm long; apical part curved; usually thickest at slightly below the middle, then narrowing slightly to the base, or sometimes not narrowing; stigma not dilated, ± flat, papillose, subtended by a band of hairs; substigmatic hairs Aliform, 0.4-0.5 mm long, usually tapering towards the tip, erect and slightly curved over the stigma. Ovules (4-)7-9. (Figure 2) Diagnostic features. Low-growing tangled shrub. Leaves often reddish with a waxy appearance. Umbo acicular. Floral tube narrow, falcate. Style up to 9 mm long, curved. Selected specimens examined. WESTERN AUSTRALIA: [localities withheld for conservation reasons] 23 Oct. 2007,E. Bennett\5IU{VFBJYiX 3 Oct. 2013, V. Clarkes.n. (PERTH);2Dec. 2009, J.A. Cochrane 7603 (PERTH); 3 Nov. 1993, R.J. Cranfield^9^\ &D. Afa/jqy (PERTH); 25 Aug. 1985, G.J. Keighery 8094 (CANB, PERTH); 9 Dec. 1976, G.S McCutcheon 859 (PERTH); 3 Nov. 1947, R.D. Royce 330 Nuytsia Vol. 30 (2019) Figure 2. Chamelaucium roycei. A - flowering leafy shoot and close-up; B - leaf, side view and cross section; C - flower bud surrounded by overlapping bracteoles, each with a long narrow acute umbo; D - flower bud, bracteoles removed; E - flower at anthesis; F - flower in longitudinal section; G - part of upper floral tube with petals, stamens, and staminodes; H - staminode and stamens with front and side view of anthers; I - style showing band of substigmatic hairs. Scale bars = 1 mm. Drawn by Donna Terrington from R.D. Royce 5765 (PERTH 01227718). N.G. Marchant, Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae) 331 2483 (PERTH); 17 Oct. 1949, R.D. Royce 3178 (PERTH, MEE); 8 Oct. 1957, R.D. Royce 5765 (AD, PERTH); 10 Sep. 1997, A. Webb 062 (PERTH); 23 Sep. 1997, A. Webb 066 & D. Hammer (PERTH). Phenology. Peak flowering is from August to November with records for July and December. Distribution and habitat. Restricted to a small area of winter wet low-lying habitats in open forest of the Donnybrook sunklands south of Busselton. Grows on red lateritic sandy or clayey soils. Conservation status. Eisted as Vulnerable in Western Australia (Smith & Jones 2018), under the name C. sp. S coastal plain (R.D. Royce 4872), Etymology. Named after Robert Dunlop Royce, a cadet of the Department of Agriculture from 1934, who joined the Botany Branch and Western Australian Herbarium in 1944 to become the Curator of the herbarium from 1960 to 1974. He made numerous plant and algal collections in the state, especially of plants in the Busselton-Margaret River area during his many annual trips certifying clover seed. He was the first to collect C. roycei in 1947 and made a further four collections of the species over the next ten years. Chamelaucium xanthocladum N.G.Marchant, sp. nov. Type-. Waychinicup [precise locality withheld for conservation reasons]. Western Australia, 28 October 1990, N.G. Marchant 90/10/7 {holo'. PERTH 06763308). Chamelaucium sp. Waychinicup (D. Davidson s.n. PERTH 01486527), Western Australian Herbarium, in FloraBase, https://florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. Chamelaucium aerocladus N.G.Marchant & Keighery ms; G. Paczkowska & A.R. Chapman, West. Austral. FI: Descr. Cat. 354 (2000); Western Australian Herbarium, in FloraBase, https://florabase. dpaw.wa.gov.au/ [accessed 6 June 2019]. Shrub 100-150 cm high or more in moist, sheltered habitats; branches weeping, with closely packed yellowish green young stems and leaves, decurrences, 2-5 mm long, widely spaced. Leaves opposite, linear, appressedto stem, 10-20 mmlong, 0.5-0.7 mm wide; abaxial surface rounded, adaxial surface with a shallow groove in lower part, apex minutely bifid; petiole 0.5-1 mm long. Flowers few, 1-2 in upper axils, 5-7 mm diam.; disc 2.3-3.5 mm diam.; pedicel 3.7-5.4 mm long. Bracteoles mostly comprised of a broad based, acuminate umbo. Floral tube turbinate to narrowly turbinate, 4-6 mm long; lower part 2.8^.4 mm long, surface shallowly rugose, without ribs; upper part 1.4-2 mm long, smooth; foveolae absent. Calyx lobes 0.2-0.6 mm long, very broadly triangular to hemispheric; margins entire or with small serrations, sinuses very broad. Petals creamy white to greenish cream, 2.5^.4 mm long, erect, obovate to ovate, deeply concave, margins entire or with minutely serrate margins. Stamens united in a tube 0.3-0.5 mm long; anthers 0.4-0.6 mm long, antisepalous stamens and staminodes geniculate and antipetalous stamens and staminodes erect or all stamens and staminodes geniculate; connective globose, reddish brown or cream; staminodes narrowly triangular, 0.4-0.6 mm long, apex obtuse. Style exserted, 4.7-5.5 mm long, very broad towards the base; stigma capitate, 0.7-0.9 mm diam., subtended by a band of hairs; substigmatic hairs white, 0.3-0.5 mm long. Ovules 4-6. (Figure 3) Diagnostic features. The distinctly weeping branches and the thin, yellow-green leaves and the often- greenish flowers distinguish this species from all other south coast species. 332 Nuytsia Vol. 30 (2019) Other specimens examined. WESTERN AETSTRAEIA: [localities withheld for conservation reasons] 1 Oct. 2010, S. Barrett & S. Comer 1978 (PERTH); 7 Oct. 2008, S. Barrett & G. Freebury 1783 (PERTH); 11 Dec. 2000, J.A. Cochrane 3792 (PERTH); 17 Nov. 1986, E.J Croxford (PERTH); 15 Oct. 1984, D. Davidson s.n. (PERTH); 27 Aug. 1986, D. Davidson s.n. (PERTH). Phenology. Flowers from late August through to January. Figure 3. Charnelaiicium xanthocladum. A - flowering leafy shoot; B - leaf, side view and cross section; C - flower bud surrounded by overlapping bracteoles, each with a broad-based umbo; D - flower bud, bracteoles removed; E - flower at anthesis; F - flower in longitudinal section; G - part of upper floral tube with petals, stamens, and staminodes; Ft - staminode and stamens with font and side view of anthers; I - style showing ring of substigmatic hairs. Scale bars = 1 mm. Drawn by Donna Terrington from D. Davidson s.n. (PERTH 01486527). N.G. Marchant, Circumscription of Chamelaucium (Myrtaceae: Chamelaucieae) 333 Distribution and habitat. Restricted to a limited range in heathland and coastal thickets between Albany and Cheyne Beach, Western Australia. Grows on brown sandy or loam soils associated with granitic outcrops. Conservation status. Currently listed as Priority Two under Conservation Codes for Western Australian Flora (Smith & Jones 2018), under the name C. sp. Waychinicup (D. Davidson s.n. PERTH 01486527). Etymology. The species name is derived from the Greek words xanthos, yellow, and dados, branches. Acknowledgements The taxonomic skills of Chin See Chung are acknowledged for his excellent support. Greg Keighery provided much valuable field knowledge and advice in the early stages of studies on Chamelaucium and Darwinia. Annette Wilson, Barbara Rye and other staff of the WA Herbarium, PERTH are sincerely thanked for their help and encouragement. Donna Terrington prepared the illustrations. References Bentham, G. (1865). Notes on the Genera Darwinia, Rudge, and Bartlingia, Ad. Brongn. The Journal of the Linnean Society. Botany 9: 176-181. Bentham, G. (1867). Flora AustraJiensis. Vol. 3. (Lovell Reeve & Co.: London.) Bibby, R (1944). Geraldton Wax-flower is a Chamelaucium. The Victorian Naturalist 61 (4): 66. Blackall, W.E. (1954). How to Know Western Australian Wildflowers, Part 1. B.J. Grieve (ed.) (University of Western Australia Press: Nedlands, Western Australia.) Blackall, W.E. & Grieve, B.J. (1980). How to Know Western Australian Wildflowers, Part 3A. Revised 2"‘' edn by B.J. Grieve. (University of Western Australia Press: Nedlands, Western Australia.) Briggs, B.G. & Johnson, L.A.S. (1979). Evolution in the Myrtaceae - Evidence from Inflorescence Structure. Proceedings of the Linnean Society of New South Wales 102: 157-256. Candolle, A.P de (1827). In: Bory de Saint-Vincent, J.B.G.N. Dictionnaire Classique d’HistoireNaturelle 11 (Rey et Gravier; Baudoinfreres: Paris.) Candolle, A.P. de (1828). Prodromus Systematis Naturalis. 3. (Treuttel and Wurtz: Paris.) Desfontaines, R.E. (1819a). Description de trois nouveaux genres. Memoires du Museum d’Histoire Naturelle 5: 34-44, t. 4. Desfontaines, R.E. (1819b). Supplement au memoire sur le genre Chamelaucium. Memoires du Museum d’Histoire Naturelle 5: 271-273,1 19. Domin, K. (1923). New additions to the flora of Western Australia. Verticordia plumosa (Desf) Domin. Vestnik Krdlovske Ceske Spolecnosti Nauk, Tfida Matematicko-pfirodovedecke 2: 79. Endlicher, S.E. (1838). Stirpium australasicarum herbaria hugeliani decades tres: 5. (J.P Sollinger: Wein.) Gardner, C.A. (1931). Enumeratio Plantarum Australiae Occidentalis (Government Printer: Perth.) George, A.S. (1967). Additions to the flora of Western Australia: ten miscellaneous new species. Journal of the Royal Society Western Australia 50(4): 97-104. Mueller, F. (1878). Myrtaceae. In: Fragmentaphytographiae Australiae. Vol. 11. (Government Printer: Melbourne.) Mueller, F. (1882). Systematic Census of Australian Plants 1. (M’Carron, Bird: Melbourne.) Rye, B.E.,Wilson, P.G., Heslewood, M.M., Perkins, A.J. & Theile, K.R. (in press). A new subtribal classification of Myrtaceae tribe Chamelaucieae. Australian Systematic Botany. Schauer, J.C. (1844). Myrtaceae R.Br. In’. Eehmann, C. (ed.) Plantae Preissianae 1: 96-160. (sumptibus Meissneri: Hamburg!.) Smith, M.G. & Jones, A. (2018). Threatened and Priority Flora list 5 December 2018. Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 24 June 2019], 334 Nuytsia Vol. 30 (2019) Sophocles, E.A., Thayer, J.H., & Drisler, H. (eds) (1900). Greek Lexicon of the Roman and Byzantine Periods (from B.C. 146 toA.D. 1100). (C. Scribner’s Sons: New York.) Sprengel, K.[P.J.] (1825). Linnaeus. C., Systema Vegetabilium Ed. 16, 2. (sumptibus Librariae Dieterichianae: Gottingen) Turczaninov, N. (1847). Decas tertia generum adhuc non-descriptorum adjectis descriptionibus nonnullarum specierum Myrtacearum xerocarpicarum atque umbelliferarum imperfectarum. Bulletin de la Societe Imperiale des Naturalistes de MoscoulQ-. 148-174. Turczaninov, N. (1849). (as Tourczaninov) Decas sexta generum plantarum hucesque non descriptorum Bulletin de la Societe Imperiale des Naturalistes de Moscou 22(2): 1-38. Turczaninov, N. (1852). Myrtaceae xerocarpicae, in Nova Hollandia. Bulletin de la Class Physico-Mathematique de L ’Academic Imperiale des Sciences de Saint-Petersbourg 10 (21, 22): 321-346. Western Australian Herbarium (1998-). FloraBase-the Western Australian flora. Department of Biodiversity, Conservation and Attractions, https://florabase.dpaw.wa.gov.au/ [accessed 6 June 2019]. Nuytsia The journal of the Western Australian Herbarium 30:335-338 Published online 10 December 2019 SHORT COMMUNICATION Hibbertia tuberculata (Dilleniaceae), a new, rare species from Western Australia During a 2009 flora and vegetation survey of the Forrestania Greenstone Belt (Thompson & Allen 2013), a specimen of Hibbertia AndxQv^s was collected from a rocky basalt area east of North Ironcap and north-west of Lake Cronin Nature Reserve. The specimen closely matched an earlier collection provisionally determined by J. Wheeler at the Western Australian Hebarium as H. atf. oligantha J.R.Wheeler, collected by Ken Newbey in 1981 from a nearby locality. Subsequently, further collections have been made from the Mount Holland area c. 20 km further north. The specimens are consistent, distinctive and not referable to any known species; accordingly, they are here described as H. tuberculata K.R. Thiele sp. nov. Hibbertia tuberculata K.R.Thiele, sp. nov. Type', c. 100 km S of Southern Cross, Western Australia [precise locality withheld for conservation reasons], 23 August 2019, B. Ellery BE 1415 (holo'. PERTH 09155317; iso: AD, CANB, K, MEE). Eow, spreading, woody shrubs to 45 cm high, with papery, exfoliating bark on all but the youngest stems, which are reddish and sparsely hairy with minute, spreading to retrorse hairs c. 0.02 mm long. Leaves crowded on new growth, erect to spreading, scattered, shortly petiolate and slightly stem- clasping at the base, green, elliptic to linear, (2-)3-5 mm long, 0.8-1.2(-2) mm wide, prominently tuberculate and sparsely hairy with short, simple hairs to 0.3 mm long, each hair arising antrorsely from a tubercle; margins revolute to the broad, tuberculate midrib, obscuring the minutely papillate true abaxial surface; apex obtuse to subacute, ± straight. Flowers solitary, terminating short lateral shoots, ± sessile; bracts 3-7, narrowly triangular to narrowly ovate, 0.8-1.2 mm long, 0.2-0.3 mm broad, acute, ± glabrous except for minutely ciliolate margins, sometimes scarious in the upper half Sepals 5; outer sepals broadly ovate, acute but not pungent, (4.5-)5-6 mm long, glabrous, without a prominent midrib; inner sepals similar to the outer ones but broader and less acute. Petals 5, yellow, obovate, 6-8 mm long, slightly emarginate. Stamens (9-)12-14, free, erect, arranged on one side of the carpels, varying in size but without true staminodes; filaments free to the base, 0.5-1.9 mm long; anthers linear-oblong, 0.5-2.0 mm long, dehiscing by longitudinal slits. Carpels 2; ovaries ± oblong, glabrous; styles lateral and strongly reflexed from ovary apex then spreading-erect, c. 3-A mm long. Ovules 2-A per carpel. Fruiting carpels and seeds not seen. (Figure 1) Diagnostic features. May be distinguished from all other species of Hibbertia in Western Australia by the combination of sessile flowers with 3-7, narrowly triangular to narrowly ovate bracts, erect stamens with free filaments on one side of the two glabrous carpels, and leaves (2-)3-5 mm long and prominently tuberculate. Other specimens examined. WESTERN AUSTRAEIA [localities withheld for conservation reasons]: 11 Sep. 2017, D. Angus 2916 (PERTH); 7 Oct. 1981, K.R. Newbey 9251 (PERTH); 5 Oct. 2009, W. Thompson & J. Allen 950 (PERTH). © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 336 Nuytsia Vol. 30 (2019) Figure 1. Flowering branch of Hibbertia tuberculata. Photo: W. Thompson {W. Thompson & J. Allen 950). Phenology. The known collections were flowering in September and October. Distribution and habitat. Occurs from Lake Cronin and North Ironcap to near Mount Holland, in the Southern Cross subregion of the Coolgardie IBRA Bioregion (Figure 2). The type locality is a gentle slope with rocky, red-brown sandy clay soils developed on greenstone. Associated vegetation is an open heathland of Melaleuca clijfortioides, Allocasuarina campestris and Dodonaea adenophora over low, sparse Grevillea lissopleura and Trymalium myrtillus subsp. myrtillus. The Newbey specimen was collected from a gentle slope on an undulating plain of well- drained, crumbly red clay supporting Melaleuca cardiophylla dwarf scrub. The site near North Ironcap is a rocky hill crest with exposed greenstone bedrock and shallow red-brown clay-loam soils, where H. tuberculata grows with Eucalyptusprotensa, Santalum acuminatum, A llocasuarina helmsii, Trymalium myrtillus subsp. myrtillus. Acacia kerryana, A. pachypoda, Grevillea acuaria, Eutaxia nanophylla and Hemigenia ajf. diplanthera. All known localities are on fine and medium-grained mafic amphibolite with metabasalt regolith adjacent to banded chert (Chin et al. 1984). Conservation status. The three known collection localities are all within c. 25 km of each other, and none is in a current conservation area. Accordingly, a status of Priority One is appropriate under the Conservation Codes for Western Australian Flora (M. Smith pers. comm.). Etymology. From the Latin tuberculatus (tuberculate, covered with wart-like projections), in reference to the leaves. K.R. Thiele, Hibbertia tuberculata (Dilleniaceae), a new species from Western Australia 337 Figure 2. Distribution of H. tuberculata. Shaded lines show the boundaries of IBRA6.1 bioregions (Department of the Environment, Water, Heritage and the Arts, 2008). Notes. Hibbertia tuberculata belongs in//, subg. Hemistemma (DC. in Thou.) J.W.Horn and is relatively unusual amongst Western Australian species of that subgenus in having erect, free stamens on one side of two glabrous carpels, and spreading-erect styles. In most other Western Australian species with stamens on one side of two carpels, the carpels are densely hairy, the staminal filaments are fused at their bases and the stamens are arranged like a hand of bananas curving forward over the carpels, with the short styles curved beneath them. Only three other Western Australian species, H. andrewsiana Diels, H. psilocarpa J.R.Wheeler and H. oligantha J.R.Wheeler, share this arrangement of stamens and carpels with H. tuberculata, the pattern is more commonly seen in eastern Australian species of subg. Hemistemma. Hibbertia tuberculata is morphologically most similar to H. oligantha. It is a woodier shrub than H. oligantha, with stout branchlets and very short seasonal growth units, whereas H. oligantha has a softer appearance with long, more or less straight seasonal shoots. The leaves of H. oligantha are longer (3.5-15 mm long), narrower (0.6-1 mm wide), and are typically smooth and glabrous (occasionally with obscure tubercles), unlike the short ((2-)3-5 mm long), broad (0.8-1.2(-2) mm wide), prominently tuberculate leaves of H. tuberculata. The distributions of the two species do not overlap, H. oligantha occurring from Esperance to Ravensthorpe and inland to the vicinity of Peak Charles. Two PERTH specimens currently included in H. oligantha from near Salmon Gums {R.D. Hoogland 12051 and W.E. Blackall 1038) have fine, trichome-tipped tubercles. These are likely to comprise another new species, which differs from H. tuberculata in the finer, denser tubercles and distinctly hairy leaves. Hibbertia psilocarpa has smooth, often glaucous leaves which, when hairy, have fine, straight, erect, non-tubercle-based hairs, 4-8(-10) stamens, and styles that are not strongly refiexed from the carpel apex. Hibbertia andrewsiana has slender-pedicellate fiowers with a single fioral bract at the apex of the often sigmoid pedicel. 338 Nuytsia Vol. 30 (2019) Summary of taxon circumscriptions Hibbertia tuberculata K.R.Thiele sp. nov. in this paper does not cause the recircumscription of any other taxon. Acknowledgements I thank Wendy Thompson for bringing this species to my attention, and Kelly Shepherd and Juliet Wege for helpful comments on the manuscript. References Chin, R.J., Hickman, A.H. & Thom, R. (1984). Hyden, Western Australia. Western Australian Geological Survey 1:250 000 Geological Series Map and Explanatory Notes Sheet SI/50-4. Geological Survey of Western Australia, Perth. Department of the Environment, Water, Heritage and the Arts (2008). Interim Biogeographic Regionalisation for Australia (IBRA), Version 6.1. http://www.environment.gov.au/parks/nrs/science/bioregion-framework/ibra/index.html [accessed October 2009]. Thompson, W.A. & Allen, J. (2013). Flora and vegetation of greenstone formations of the Yilgarn Craton: The northern Forrestania Greenstone Belt (Mount Holland area). Conservation Science Western Australia 8(3): 277-294. Kevin R. Thiele School of Biological Sciences, University of Western Australia 35 Stirling Hwy, Crawley WA 6009 Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions, Eocked Bag 104, Bentley Delivery Centre, Western Australia 6983 Email: kevin.thiele@uwa.edu.au Nuytsia The journal of the Western Australian Herbarium 30:339-351 Published online 10 December 2019 SHORT COMMUNICATION Updates to Western Australia’s vascular plant census for 2019 The census database at the Western Australian Herbarium (PERTH), which provides the nomenclature for the website FloraBase (Western Australian Herbarium 1998-), lists current names and recent synonymy for Western Australia’s native and naturalised vascular plants, as well as algae, bryophytes, lichens, slime moulds and some fungi. The names represented in the census are either sourced from published research or denote as yet unpublished names based on herbarium voucher specimens. We herein summarise the changes made to vascular plant names in this database during 2019. Ninety-five taxa were newly recorded for the State, of which 11 are naturalised and 33 have been added to the Threatened and Priority Flora list for Western Australia (Smith & Jones 2018; Western Australian Herbarium 1998-) (Table 1). A total of 119 name changes were made, including the formal publication of 50 phrase-named taxa (Table 2). Plant groups for which a number of name changes were made include Eremophila R.Br. (Brown & Davis 2019; Chinnock 2019), Hibbertia Andrews (Thiele 2019a), HydrocotyleL. (Perkins 2018,2019) and Styphelia Sm. (Hislop 2019; Hislop & Puente-Lelievre 2019). Numerous changes were made in Scholtzia Schauer in the latest of a series of papers aimed at reducing the large backlog of informally-named taxa in the Myrtaceae tribe Chamelaucieae (Rye 2019). A comprehensively illustrated revision of the tiurndins (Tribonanthes, Haemodoraceae) recognised 12 species, of which four were newly described and two were reinstated (Hickman & Hopper 2019). Table 2 also includes cases where there has been a change of taxonomic concept, misapplication, exclusion or rank change. Table I. New records added to Western Australia’s vascular plant census during 2019. in litt. = in correspondence; = on herbarium sheet/label. Status: * = naturalised; T,P1-P4=Conservation Codes for Western Australian Flora (Smith & Jones 2018; Western Australian Herbarium 1998-)- New Name Status Comments Acacia sp. Forrestania (D. Angus DA 3001) PI M. Hislop in litt. (20/09/2019). Actinotus sp. Whicher (G.J. Keighery 12394) Correction to voucher-specimen identifier. See Keighery (1999). Austrostipa echinata (Vickery, S.W.L.Jacobs & J.Everett) S.W.L.Jacobs & J.Everett PI New record for WA. T.D. Macfarlane in litt. (13/12/2018). Boronia clavata Paul G. Wilson subsp. clavata T See Duretto (2019). Boronia clavata subsp. grandis Duretto P2 See Duretto (2019). Boronia denticulata Sm. subsp. denticulata See Duretto (2019). Boronia denticulata subsp. whoogarupensis Duretto P2 See Duretto (2019). Boronia sp. Eeeuwin (J. Scott 235) P2 A. Webb in litt. (07/05/2019). © Department of Biodiversity, Conservation and Attractions 2019 https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 2200-2790 (Online) ISSN 0085-4417 (Print) 340 Nuytsia Vol. 30 (2019) New Name Status Comments Calandrinia sp. Cape Range (F. Obbens FO 10/18) P2 F. Obbens/>?//«. (16/11/2018). Calandrinia sp. Nanutarra (F. Obbens FO 08/18) PI F. Ohhens in litt. (16/11/2018). Calochilus sp. Theda (R.L. Barrett et al. RLB 6158) R E. Barrett & T.D. Macfarlane in litt. (01/07/2019). Calyptocarpus vialis Less. * New naturalised record for WA. G.J. Keighery in litt. (10/12/2018). Cassinia sifton Orchard See Orchard (2017). Comesperma sp. A Kimberley Flora (D.W. Rust 48) Correction to voucher-specimen identifier. Convolvulus sp. Cascades (W. Archer 1110161) PI M. Hislop in litt. (11/07/2019). Cor chorus drysdalensis R.L. Barrett See Barrett (2019b). Corybas sp. Peat (A.P. Brown & D. Edmonds APB 4314) A.P. Brown in litt. (21/05/2019). Danvinia sp. Corrigin (T. Erickson TEE 308) P2 B.E. Rye in litt. (03/09/2019). Dillwynia sp. Mallee (W.R. Archer 1709959) M. Hislop in litt. (10/10/2019). Diuris sp. Augusta (G. Brockman GBB 1469) C. French & G. Brockman in litt. (05/12/2018). Drosera sp. Branched styles (S.C. Coffey 193) S.C. Coffey in litt. (30/11/2018). Elionurus tylophorus M.D.Barrett & Handasyde P2 See Barrett & Handasyde (2019). Eremophila sp. Kalgoorlie (V. Clarke & A. Brown VTC 590) Correction to voucher-specimen identifier. A.P. Brown in litt. (30/01/2019). Eremophila sp. Mullewa (R. Wait 7311) R. Davis in litt. (09/01/2019). Eriachne sp. Dugald River (B.K. Simon+ 3007) M.D. Barrett in litt. (18/01/2019). Eucalyptus cladocalyx F.Muell. subsp. cladocalyx * New naturalised record for WA. G.J. Keighery in litt. (08/07/2019). Eucalyptus vokesensis D.Nicolle & E.A.S. Johnson New record for WA. M.E. French in litt. (12/06/2019). Exocarpos sp. Ardath (J. Buegge D 62) B.J. Eepschi in litt. (30/10/2018). Gloriosa superba E. New naturalised record for WA. G.J. Keighery in litt. (10/02/2019). Goodenia sp. Cunyu (C.J. Nicholson & PJ. Curry 1001) PI K.A. Shepherd & M. Hislop in litt. (14/02/2019). Grevillea sp. Yerilgee Hills (T. Easlett TE 025) PI J. Bull & D. Brearley in litt. (05/06/2019). Habenaria hymenophylla Schltr. New record for WA. R E. Barrett in litt. (02/01/2019). C.M. Parker & J.M. Percy-Bower, Updates to Western Australia’s vascular plant census for 2019 341 New Name Status Comments Haemodorum coccineum R.Br. New record for WA. See Barrett et al. (2015). Haemodorum subvirens F.Muell. New record for WA. See Barrett et al. (2015). Heteranthera reniformis Ruiz & Pav. New naturalised record for WA. M. Hislop in litt. (25/07/2019). Hibbertia acrotoma K.R.Thiele PI See Thiele (2019a). Hibbertia ambita K.R.Thiele PI See Thiele (2019a). Hibbertia davisii K.R.Thiele See Thiele (2019a). Hibbertia elegans K.R.Thiele PI See Thiele (2019a). Hibbertia hortiorum K.R.Thiele PI See Thiele (2019a). Hibbertia improna K.R.Thiele See Thiele (2019a). Hibbertia inopinata K.R.Thiele See Thiele (2019a). Hibbertia sandifordiae K.R.Thiele PI See Thiele (2019a). Hibbertia semipilosa K.R.Thiele See Thiele (2019a). Hibbertia spectabilis K.R.Thiele See Thiele (2019a). Hibbertia wheelerae K.R.Thiele P2 See Thiele (2019a). Hibiscus sp. Perrinvale Station (J. Warden & E. Ager WB 10581) P3 G. Cockerton in litt. (05/08/2019). Lagenophora gracilis Steetz New record for WA. See Wang & Bean (2019). Lagenophoraplatysperma Jian Wang ter & A.R.Bean See Wang & Bean (2019). Lagenophora stipitata (Labill.) Druce New record for WA. See Wang & Bean (2019). Lechenaultia sp. Cascade (W.R. Archer 212122) PI K.A. Shepherd & M. Hislop in litt. (14/02/2019). Lepidium sp. Stirling Range (S. Barrett 1359) G.J. Keighery in litt. (23/10/2019). Leucopogon sp. Cascades (M. Hislop 3693) PI M. Hislop in litt. (07/02/2019). Levenhookia sp. Whicher Range (J.A. Wege 2090) J.A. Wege in litt. (11/04/2019). Micrococca mercurialis (L.) Benth. * New naturalised record for WA. G.J. Keighery in litt. (10/02/2019). Microcorys sp. Mt Holland broad-leaf (G. Barrett s.n. PERTH 04104927) PI M. Hislop in litt. (12/09/2019). Nandina domestica Thunb. * New naturalised record for WA. G.J. Keighery in litt. (10/10/2018). 342 Nuytsia Vol. 30 (2019) New Name Status Comments Nicotiana gascoynica M.W. Chase & Christenh. See Chase & Christenhusz (2018b). Nicotiana karijini M.W.Chase & Christenh. See Chase & Christenhusz (2018a). Nicotiana stenocarpa H.-M.Wheeler New record for WA. See Chase & Christenhusz (2018c). Opuntia polyacantha Haw. * New naturalised record for WA. R.J. Chmnock in litt. (23/05/2019). Opimtiapolyacantha var. hystricina (Engelm. & J.M.Bigelow) B.D.Parfitt New naturalised record for WA. R.J. Chinnock in litt. (23/05/2019). Phehalium sp. Yerilgee Sandplain (J. Jackson 223) P2 J. Bull in litt. (24/07/2019). Phytolacca americana L. * New naturalised record for WA. G.J. Keighery in litt. (09/07/2019). Pterostylis sp. Bloated bird orchid (A.R. Annels et al. ARA 5478) C. French & G. Brockman in litt. (05/12/2018). Pterostylis sp. Bloated snail orchid (W. Jackson BJ 486) C. French & G. Brockman in litt. (05/12/2018). Pterostylis sp. Coastal clubbed sepals (G. Brockman GBB 255) C. French & G. Brockman in litt. (05/12/2018). Pterostylis sp. Paynes Find (G. Brockman GBB 526) C. French & G. Brockman in litt. (05/12/2018). Pterostylis sp. Robust snail orchid (W. Jackson BJ 358) C. French & G. Brockman in litt. (05/12/2018). Ptilotus xerophilus T. Hammer & R.W. Davis See Hammer et al. (2019). Rorippa sp. Fortescue Valley (M.N. Fyons & R.A. Coppen FV0760) PI M.T. Collins & M.N. Fyons in litt. (25/06/2019). Scholtzia brevistylis Rye See Rye (2019). Scholtzia halophila Rye See Rye (2019). Scholtzia longipedata Rye See Rye (2019). Scholtzia pentamera Rye See Rye (2019). Scholtzia pentamera subsp. collina Rye See Rye (2019). Scholtzia uniovulata Rye See Rye (2019). 5/^/asp.R(P. Copley 1390) PI New record for WA. R.M. Barker in sched. (May 2007). Siylidium sp. Dragon Rocks (J.A. Wege & K.A. Shepherd JAW 2054) P2 J.A. Wege in litt. (11/04/2019). Styphelia sp. Wandoo (F. & J. Hort 2441) P2 M. Hislop in litt. (22/01/2019). Syncarpia glomulifera (Sm.) Nied. New naturalised record for WA. G.J. Keighery in litt. (10/10/2018). C.M. Parker & J.M. Percy-Bower, Updates to Western Australia’s vascular plant census for 2019 343 New Name Status Comments Thysanotus sp. Ennuin (N. Gibson & M. Lyons 2665) PI T. Macfarlane in litt. (20/02/2019). Thysanotus sp. Mount Madden (G.F. Craig 10516) PI T. Macfarlane in litt. (27/05/2019). Thysanotus sp. Yellowdine (A.S. George 6040) P2 T. Macfarlane in litt. (11/03/2019). Tribonanthes australis Endl. x Tribonanthes brachypetala Lindl. See Hickman & Hopper (2019). Tribonanthes australis Endl. x Tribonanthes longipetala Lindl. See Hickman & Hopper (2019). Tribonanthes brachypetala Lindl. x Tribonanthes variabilis Lindl. See Hickman & Hopper (2019). Tribonanthes elongata E.J.Hickman & Hopper See Hickman & Hopper (2019). Tribonanthes monantha E. J.Hickman & Hopper See Hickman & Hopper (2019). Tribonanthesporphyrea E. J.Hickman & Hopper See Hickman & Hopper (2019). Triodia sp. Silvergrass (P.-L. de Kock BES 00808) PI M.D. Barrett & P.-L. de Kock in litt. (06/02/2018). Triodia veniciae M.D.Barrett See Barrett (2019a). Trithuriafitzgeraldii D.D.Sokolotf, LMarques, T.Macfarlane, Rudall & S.W.Graham See Sokoloffeta/. (2019). Utricularia oppositiflora R.Br. New record for WA. R.W. Jobson in sched. (06/09/2019). Vallisneria sp. Weelarrana (M.N. Lyons & S.D. Lyons 3050) PI T. Macfarlane & M. Lyons in litt. (19/06/2019). Table 2. Changes to existing entries in Western Australia’s vascular plant census during 2019. Excluded taxon = a name used in the botanical literature that refers to a taxon never occurring in WA; misapplied name = a name used in the botanical literature but now considered to refer to one or more different WA taxa; nomenclatural synonym = a superseded name based on the same type specimen as the accepted name—the epithet is usually transferred to a different genus name or rank; taxonomic synonym = a superseded name based on a different type specimen to the accepted name; orthographic variant = mis-spelling of a name in original publication; in litt. = in correspondence; in sched. = on herbarium sheet/label. Status: * = naturalised; T, P1-P4 = Conservation Codes for Western Australian Flora (Smith & Jones 2018; Western Australian Herbarium 1998-). Old Name New Name Status Comments Abutilon halophilum Schldl. n/a Excluded taxon. No records from WA. Acacia sp. East Fortescue (J. Bull & D. Roberts ONSA27.01) Acacia corusca J.P.Bull, S. J.Dillon & Brearley PI Taxon formally published. See Bull etal. (2019). Actinotus sp. Whicher (G.J. Keighery 12394) Actinotus whicheranus Keighery P2 Taxon formally published. See Keighery (1999). 344 Nuytsia Vol. 30 (2019) Old Name New Name Status Comments Ageratina adenophora (Sprang.) R.M.Kmg & H.Rob. n/a Name made current. Taxon reinstated. G.J. Keighery in litt. (10/10/2018). Arthrochilus byrnesii Blaxell Phoringopsis byrnesii (Blaxell) D.F. Jones & M.A.Clem. Nomenclatural synonym. A.P. Brown in litt. (13/06/2018). Astroloma sp. Dumbleyung (A.J.G. Wilson 146) Styphelia sp. Dumbleyung (A.J.G. Wilson 146) P3 Name synonymised. M. Hislop in litt. (15/05/2019). Astroloma sp. Eneabba (N. Marchant s.n. PERTH 01291777) Styphelia sp. Eneabba (N. Marchant s.n. PERTH 01291777) Name synonymised. M. Hislop in litt. (15/05/2019). Astroloma sp. Kalbarri (D. &B. Bellairs 1368) Styphelia cerniia Hislop & Puente-Eel. P2 Taxon formally published. See Hislop & Puente-Lelievre (2019). Astroloma sp. Nannup (R.D. Royce 3978) Styphelia sp. Nannup (R.D. Royce 3978) Name synonymised. M. Hislop in litt. (15/05/2019). Astroloma sp. Narrogin (R.D. Royce 8158) Styphelia sp. Narrogin (R.D. Royce 8158) Name synonymised. M. Hislop in litt. (15/05/2019). Astroloma sp. sessile leaf (J.L. Robson 657) Styphelia angustiflora Hislop & Puente-Eel. P2 Taxon formally published. See Hislop & Puente-Lelievre (2019). Astroloma stomarrhena Sond. Styphelia stomarrhena (Sond.) Sleumer Nomenclatural synonym. See Hislop & Puente-Lelievre (2019). Astroloma xerophyllum (DC.) Sond. Styphelia xerophylla (DC.) F.Muell. Nomenclatural synonym. See Hislop & Puente-Lelievre (2019). Banksia densa A.R.Mast & K.R.Thiele var. densa Banksia densa A.R.Mast & K.R.Thiele P2 Nomenclatural synonym. No varieties recognised. See Thiele (2019b). Banksia densa var. parva (A S.George) A.R.Mast & K.R.Thiele Banksia parva (A. S. George) K.R.Thiele P4 Nomenclatural synonym. See Thiele (2019b). Banksia densa var. Wheatbelt (M. Pieroni s.n. PERTH 04083407) Banksia zygocephala K.R.Thiele Taxon formally published. See Thiele (2019b). Caladenia x intermedia (¥itzg.) M. A.Clem. & D.L. Jones Elythranthera x intermedia (Fitzg.) M.A.Clem. Nomenclatural synonym. A.P. Brown in litt. (09/01/2018). Calandrinia sp. Piawaning (A.C. Beauglehole 12257) Calandrinia uncinella Obbens PI Taxon formally published. See Obbens (2019). Calandrinia sp. The Pink Hills (F. Obbens FO 19/06) Calandrinia monosperma Obbens Taxon formally published. See Obbens (2019). Clerodendrim heterophyllum (Poir.) R.Br. n/a Excluded taxon. PERTH collections represent cultivated material. Clerodendrum heterophyllum var. baueri Moldenke nJdL Excluded taxon. PERTH collections represent cultivated material. Comesperma sp. A Kimberley Flora (D.W. Rust 8) Comesperma sp. A Kimberley Flora (D.W. Rust 48) Error. Correction to voucher-specimen identifier. Comesperma sp. A Kimberley Flora (D.W. Rust 48) Comesperma minutum A. J.Ford & Halford PI Taxon formally published. See Ford etal. (2017). Corybas sp. Peat (A.P. Brown & D. Edmonds APB 4314) Corybas autumnalis A.P.Br. & D.Edmonds P2 Taxon formally published. See Brown & Edmonds (2019). Crosslandia setifolia W.Fitzg. Fimbristylis crosslandii Roalson, R E. Barrett & Larridon Nomenclatural synonym. See Roalson etal. (2019). Cyanthillium gracilis (Fander & P.J.H.Hurter) K.R.Thiele & E.E.Schill. Cyanthillium gracile (Lander & P.J.H.Hurter) K.R.Thiele & E.E.Schill. P3 Orthographic variant. A.S. George in litt. (13/12/2018). C.M. Parker & J.M. Percy-Bower, Updates to Western Australia’s vascular plant census for 2019 345 Old Name New Name Status Comments Cyathostemon sp. Mt Dimer (C. McChesney TRL 4/72) Cyathostemon verrucosus Trudgen & Rye P3 Taxon formally published. See Rye (2018). Delosperma vinaceum L.Bolus n/a Excluded taxon. See Keighery (2005). Dillwynia sp. Coolgardie (V.E. Sands 637.3.1) Dillwynia acerosa S.Moore Informal synonym. M. Hislop in sched (03/10/2019). Diuris sp. South Coast (G. Brockman GBB 3041) Diuris brockmanii D.E. Jones & C.J.French Taxon formally published. See Jones & French (2019). Drosera indica L. nJdL Excluded taxon. See Eowrie (2013). Enneapogon nigricans (R.Br.) PBeauv. nJdL Name made current. Taxon reinstated. T.D. Macfarlane & U. Bell in sched. (21/03/2019). Eremophila glabra subsp. Kokeby (R. Davis 5080) Eremophila glabra subsp. York (PG. Wilson 12172 B) PI Name synonymised. A.P Brown in sched (08/02/2017). Eremophila glabra subsp. verrucosa Chinnock Eremophila viridissima Chinnock Nomenclatural synonym. See Chinnock (2019). Eremophila oldfieldii subsp. Karara (D. Coultas s.n. PERTH 07341717) Eremophila oldfieldii subsp. papula A.VBr. PI Taxon formally published. See Brown & Davis (2019). Eremophila sp. Kalgoorlie (V. Clark & A. Brown VTC 590) Eremophila sp. Kalgoorlie (V. Clarke & A. Brown VTC 590) Error. Correction to voucher-specimen identifier. A.P. Brown in litt. (30/01/2019). Eremophila sp. Kalgoorlie (V. Clarke & A. Brown VTC 590) Eremophila xantholaema R.W. Davis PI Taxon formally published. See Brown & Davis (2019). Eremophila sp. Mullewa (R. Wait 7311) Eremophila waitii Chinnock PI Taxon formally published. See Chinnock (2019). Eremophila sp. Rothsay (D. Coultas & J. Kelt s.n. PERTH 08200440) Eremophila sericea A.P.Br. PI Taxon formally published. See Brown & Davis (2019). Eucalyptus gardneri subsp. ravensthorpensis Brooker & Hopper Eucalyptus ravensthorpensis (Brooker & Hopper) Gosper & Hopper P4 Nomenclatural synonym. See Gosper etal (2019). Eucalyptus obtusiflora subsp. cowcowensis E.A.S.Johnson & K.D.Hill Eucalyptus obtusifiora subsp. dongarraensis (Maiden & Blakely) E.A.S.Johnson & K.D.Hill Taxonomic synonym. See Nicolle etal (2019). Gastrolobium sp. Harvey (G.J. Keighery 16821) Gastrolobium capitatum (Benth.) G.Chandler & Crisp Name synonymised. See Butcher & Hislop (2019). Genus sp. Nillup (R.D. Royce 98) Hemigenia sp. Nillup (R.D. Royce 98) P2 Name synonymised. B E. Rye in litt. (25/02/2014). Geodorum neocaledonicum Kraenzl. Geodorum densifiorum (Earn.) Schltr. Taxonomic synonym. R E. Barrett in litt. (08/06/2018). Geodorum terrestre (E.) Garay Geodorum densifiorum (Earn.) Schltr. Misapplied name. R.E. Barrett in litt. (08/06/2018). Goodenia campestris Carolin nJdi Name made current. Taxon reinstated. M.D. Barrett in litt. (23/04/2019). Grewia glabra Blume Grewia guazumifolia Juss. Taxonomic synonym. See Barrett (2019b). 346 Nuytsia Vol. 30 (2019) Old Name New Name Status Comments Grewia retusifolia Kurz Grewia pindanica R.E.Barrett Misapplied name. See Barrett (2019b). Grewia retusifolia Kurz Grewia savannicola R.E.Barrett Misapplied name. See Barrett (2019b). Habenaria elongata R.Br. Pecteilis elongata (R.Br.) M.A.Clem. & D.E.Jones Nomenclatural synonym. See Clements & Jones (2018). Habenaria eurystoma Schltr. Pecteilis eurystoma (Schltr.) M.A.Clem. & D.E.Jones Nomenclatural synonym. See Clements & Jones (2018). Habenaria ochroleuca R.Br. Pecteilis ochroleuca (R.Br.) M.A.Clem. & D.E.Jones Nomenclatural synonym. See Clements & Jones (2018). Habenaria sp. Beverley Springs Station (M.D. Barrett MDB185) Pecteilis eurystoma (Schltr.) M.A.Clem. & D.E.Jones Name synonymised. R.E. Barrett in litt. (02/01/2019). Habenaria triplonema Schltr. Pecteilis elongata (R.Br.) M.A.Clem. & D.E.Jones Taxonomic synonym. R E. Barrett in litt. (02/01/2019). Hibbertia sp. Toolbrunup (J.R. Wheeler 2504) Hibbertia barrettiae K.R.Thiele T Taxon formally published. See Thiele (2019a). Hydrocotyle sp. Crassipes (K.R. Newbey 7567) Hydrocotyle e/c/z/en A.J.Perkins P3 Taxon formally published. See Perkins (2018). Hydrocotyle sp. Decipiens (G.J. Keighery 463) Hydrocotyle tuberculata A.J.Perkins P2 Taxon formally published. See Perkins (2018). Hydrocotyle sp. Hamelinensis (G.J. Keighery s.n. PERTH 02391325) Hydrocotyle tetragonocarpa Bunge Name synonymised. See Perkins (2019). Hydrocotyle sp. Puberula (H. Eichler 22058) Hydrocotyle scutellifera Benth. Name synonymised. See Perkins (2019). Hydrocotyle sp. Vigintimilia (PG. Wilson 7940) Hydrocotyle papilionella A. J. Perkins P2 Taxon formally published. See Perkins (2018). Lepidosperma leptophyllum Benth. n/a Name made current. Taxon reinstated. See CHAH (2012). Leptocarpus crassipes Pate & Meney Leptocarpus scariosus R.Br. Taxonomic synonym. B.G. Briggs in litt. (13/03/2019). Leucopogon sp. Barren Range (A.S. George 10092) Styphelia quartzitica Hislop P2 Taxon formally published. See Hislop (2019). Leucopogon sp. Bonnie Hill (K.R. Newbey 9831) Styphelia sulcata Hislop & Puente-Eel. PI Taxon formally published. See Hislop & Puente-Eelievre (2019). Leucopogon sp. Kambalda (J. Williams s.n. PERTH 07305028) Styphelia rectiloba Hislop P3 Taxon formally published. See Hislop (2019). Leucopogon sp. Ongerup (A.S. George 16682) Styphelia disjuncta Hislop & Puente-Eel. T Taxon formally published. See Hislop & Puente-Eelievre (2019). Livistona lorophylla Becc. Livistona leichhardtii F.Muell. Taxonomic synonym. See Dowe (2018). Neosciadium glochidiatum (Benth.) Domin Hydrocotyle glochidiata Benth. Nomenclatural synonym. See Perkins (2019). Planchonella pohlmaniana (F.Muell.) Dubard Planchonella arnhemica (Benth.) PRoyen Misapplied name. See Jessup (2019). Plectranthus congestus R.Br. Coleus congestus (R.Br.) A.J.Paton Nomenclatural synonym. See Paton etal. (2019). C.M. Parker & J.M. Percy-Bower, Updates to Western Australia’s vascular plant census for 2019 347 Old Name New Name Status Comments Plectranthus intraterraneus S.T.Blake Coleus intraterraneus (S.T.Blake) P.I.Forst. & T.C.Wilson Nomenclatural synonym. See Paton etal (2019). Plectranthus scutellarioides (L.) R.Br. Coleus scutellarioides (E.) Benth. Nomenclatural synonym. See Paton etal. (2019). Pleurocarpaea gracilis Lander & PJ.H.Hurter Cyanthillium gracilis (Eander & PJ.H.Hurter) K.R.Thiele & E.E.Schill. Nomenclatural synonym. See Thiele & Schilling (2018). Pterostylis sp. Karri forest (W. Jackson BJ270) Pterostylis sp. crinkled leaf (G.J. Keighery 13426) Name synonymised. G. Brockman in sched (08/02/2013). Pterostylis sp. short sepals (W. Jackson BJ259) Pterostylis ectypha (D.E. Jones & C.J.French) D.E.Jones & C.J.French Name synonymised. G. Brockman & C.J. French in sched (19/12/2018). Ptilotus macrocephalus (R.Br.) Poir. nJdL Excluded taxon. See Hammer et al. (2019). Rhagodia sp. Yeelirrie Station (K.A. Shepherd et al. KS 1396) Chenopodium nitrariaceum (F.Muell.) Benth. Name synonymised. K.A. Shepherd in litt. (27/02/2019). Sauropiis crassifolius (Mull.Arg.) Airy Shaw Synostemon crassifolius (Mull.Arg.) I.Telford & Pruesapan Nomenclatural synonym. See Telford etal. (2019). Sauropus rigidiilus (Mull.Arg.) Airy Shaw Synostemon rigidulus (Mull.Arg.) I.Telford & Pruesapan P3 Nomenclatural synonym. See Telford etal. (2019). Scholtzia sp. Ajana (T.A. Halliday 137) Scholtzia bellairsiorum Rye P3 Taxon formally published. See Rye (2019). Scholtzia sp. Ajana East Road (M.E. Trudgen 21734 A) Scholtzia truncata Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Billeranga Hills (B.J. Conn 2159) Scholtzia subsessilis Rye PI Taxon formally published. See Rye (2019). Scholtzia sp. Binnu (M.E. Trudgen 2218) Scholtzia uniflora Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Binnu East Road (M E. Trudgen 12013) Scholtzia thinicola Rye PI Name synonymised. See Rye (2019). Scholtzia sp. Binnu-Yuna (M.E. Trudgen 12016) Scholtzia longipedata subsp. procera Rye P3 Name synonymised. See Rye (2019). Scholtzia sp. Bungabandi Creek (M. QuickC EURA48) Scholtzia peltigera Rye PI Taxon formally published. See Rye (2019). Scholtzia sp. Burma Road (A C. Burns 138) Scholtzia prostrata Rye P3 Taxon formally published. See Rye (2019). Scholtzia sp. Coburn (N. Murdock NM 031) Scholtzia corrugata Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Coomberdale (M E. & M.E. Trudgen MET 1724) Scholtzia halophila Rye subsp. halophila Taxon formally published. See Rye (2019). Scholtzia sp. Dongara (R. Hart 8401) Scholtzia calcicola Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Duck Pool (M.E. Trudgen MET 5427) Scholtzia halophila subsp. mortlockensis Rye P3 Taxon formally published. See Rye (2019). Scholtzia sp. East Yuna (A.C. Burns 6) Scholtzia inaequalis Rye P2 Taxon formally published. See Rye (2019). 348 Nuytsia Vol. 30 (2019) Old Name New Name Status Comments Scholtzia sp. Eneabba (S. Maley 8) Scholtzia trilocularis Rye Taxon formally published. See Rye (2019). Scholtzia sp. Eradu (R.D. Royce 8016) Scholtzia longipedata subsp. procera Rye P3 Name synonymised. See Rye (2019). Scholtzia sp. Eurardy (J.S. Beard 6886) Scholtzia oleosa Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Galena (W.E. Blackall 4728) Scholtzia truncata Rye P2 Name synonymised. See Rye (2019). Scholtzia sp. Gunyidi (J.D. Briggs 1721) Scholtzia quindecim Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Kojarena (A.M. Ashby 1904) Scholtzia multiflora Rye PI Taxon formally published. See Rye (2019). Scholtzia sp. Eancelin (M E. Trudgen 1516) Scholtzia laciniata Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Murchison River (A.S. George 7908) Scholtzia denticulata Rye P2 Name synonymised. See Rye (2019). Scholtzia sp. Northampton (A. Strid 20714) Scholtzia pentamera Rye subsp. pentamera Taxon formally published. See Rye (2019). Scholtzia sp. Overlander (M E. Trudgen 12138) Scholtzia recurva Rye PI Taxon formally published. See Rye (2019). Scholtzia sp. Prowaka Springs (R.J. Cranfield & P. Spencer 8083) Scholtzia brevistylis subsp. prowaka Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Red Bluff (A. Gunness 2373) Scholtzia longipedata Rye subsp. longipedata Taxon formally published. See Rye (2019). Scholtzia sp. Ross Graham Eookout (S. Maley 6) Scholtzia denticulata Rye P2 Name synonymised. See Rye (2019). Scholtzia sp. Shark Bay (M.E. Trudgen 7429) Scholtzia capitata Benth. Name synonymised. See Rye (2019). Scholtzia sp. Valentine Road (S. Patrick 2142) Scholtzia thinicola Rye PI Name synonymised. See Rye (2019). Scholtzia sp. Winchester (C. Chapman s.n. PERTH 05625386) Scholtzia chapmanii Rye Taxon formally published. See Rye (2019). Scholtzia sp. Yandanooka (R. Soullier 646) Scholtzia brevistylis Rye subsp. brevistylis PI Taxon formally published. See Rye (2019). Scholtzia sp. Yenyening Eakes (A. Gunness 2824) Scholtzia halophila subsp. meridionalis Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Yuna (C.A. Gardner 14286) Scholtzia cordata Rye P2 Taxon formally published. See Rye (2019). Scholtzia sp. Z-Bend (Bellairs-Kalflora 912a) Scholtzia tenuissima Rye P2 Taxon formally published. See Rye (2019). Tephrosia sp. Carnarvon (J.H. Ross 2681) Tephrosia gardneri R.Butcher Name synonymised. See Butcher (2018). Tephrosia sp. Fortescue (A.A. Mitchell 606) Tephrosia densa (Benth.) R.Butcher Name synonymised. See Butcher (2018). Tephrosia sp. Meentheena (S. van Eeeuwen 4479) Tephrosia densa (Benth.) R.Butcher Name synonymised. See Butcher (2018). C.M. Parker & J.M. Percy-Bower, Updates to Western Australia’s vascular plant census for 2019 349 Old Name New Name Status Comments Tephrosia sp. Onslow (K.R. Newbey 10571) Tephrosia gardneri R.Butcher Name synonymised. See Butcher (2018). Tribonanthes sp. Lake Muir (G.J. Keighery & N. Gibson 2134) Tribonanthes keigheryi E. J.Hickman & Hopper Taxon formally published. See Hickman & Hopper (2019). Tribonanthes uniflora Lindl. n/a Name made current. Taxon reinstated. See Hickman & Hopper (2019). Tribonanthes variabilis Lindl. n/a Name made current. Taxon reinstated. See Hickman & Hopper (2019). Triodia sp. Robe River (M.E. Trudgen et al. MET 12367) Triodia pisoliticola Trudgen & M.D.Barrett P3 Taxon formally published. See Barrett & Trudgen (2018). Acknowledgements Curation staff at PERTH and the Department of Biodiversity, Conservation and Attraction’s ‘ Taxonomic Review Committee’ are acknowledged for their contributions to the information presented herein. References Barrett, M. D. (2019a). Triodiaveniciae (Poaceae), a new species from the Pilbara region. Western Australia. Niiytsia 30:221-228. Barrett, M.D. & Handasyde, T. (2019). Elionurus tylophoriis (Poaceae: Paniceae: Andropogoneae), a new species from the Kimberley region of Western Australia. Niiytsia 30: 177-194. Barrett, M.D. & Trudgen, M.E. (2018). Triodia pisoliticola (Poaceae), a new species from the Pilbara region. Western Australia, and a description for T. sp. Mt Ella (M.E. Trudgen MET 12739). Nuytsia 29: 271-281. Barrett, R.L. (2019b). Three new species of Corchoriis L. and GrewiaL. (Sparmanniaceae /Malvaceae subfamily Grewioideae) from northern Australia, an earlier name in Grewia, and recircumscription of Triumfetta kenneallyi Elalford. Austrobaileya 10: 458-472. Barrett, R.L., Hopper, S.D., Macfarlane, TD. & Barrett, M.D. (2015). Seven new species of Haemodorum (Haemodoraceae) from the Kimberley region of Western Australia. Nuytsia 26: 111-125. Brown, A.P & Davis, R.W. (2019). Eremophila oldfieldii subsp. papula, E. sericea and E. xantholaema (Scrophulariaceae), three new taxa from Western Australia. Nuytsia 30: 25-31. Brown, A.P. & Edmonds, D. (2019). Corybas autumnalis (Orchidaceae), a rare new species of helmet orchid from south-west Western Australia. Orchadian 19: 392-394. Bull, J.P, Dillon, S.J. & Brearley, D.R. (2019). Acacia corusca (Fabaceae: Mimosoideae), a new species from the Pilbara bioregion in north-western Australia. Nuytsia 30: 19-22. Butcher, R. (2018). Making it official—formal description of two orange-flowered Tephrosia (Fabaceae: Millettieae) species from north-west Western Australia. Nuytsia 29: 251-267. Butcher, R. & Hislop, M. (2019). Gastrolobium sp. Harvey (G.J. Keighery 16821) (Fabaceae) is not distinct from G. capitatum. Nuytsia 30: 23-24. CHAH (Council of Heads of Australasian Herbaria) (2012). National Species List, https ://id. biodiversity.org. au/name/apni/86176 [accessed 19/11/2019]. Chase, M.W. & Christenhusz, M.J.M. (2018a). Nicotiana karijini, Solanaceae. Curtis’s Botanical Magazine 3>5: 228-236. Chase, M.W. & Christenhusz, M.J.M. (2018b). Nicotianagascoynica, Solanaceae. Curtis’s Botanical Magazine 35: 245-252. Chase, M.W. & Christenhusz, M.J.M. (2018c). Nicotiana stenocarpa, Solanaceae. Curtis’s Botanical Magazine 35: 319-327. Chinnock, R.J. (2019). Additions to Eremophila (Scrophulariaceae). Nuytsia 30: 215-219. Clements, M.A. & Jones, D.L. (2018). Pecteilis Raf (Orchideae): a molecular phylogenetic based determination of their status in Australian Orchidaceae. Australian Orchid Review 83(6): 50-53. 350 Nuytsia Vol. 30 (2019) Dowe, J.L. (2018). Livistona leichhardtii is the correct name for Livistona lorophylla (Arecaceae). Nuytsia 29: 245-250. Duretto, M.F. (2019). New subspecies for the south-western Australian species Boronia clavata and B. denticulata (Rutaceae). Telopeall-. 31-39. Ford, A.J., Flalford, D.A., Van Der Merwe, M. & Mathieson, M.T. (2017). A revision of the tropical white-flowered species of Comesperma (Polygalaceae) in Australia. Australian Systematic Botany 30; 159-182. Gosper, C.R., Hopley, T., Byrne, M., Hopper, S.D., Prober, S.M. & Yates, C.J. (2019). Elevation of Eucalyptus gardneri subsp. ravensthorpensis, and notes on relationships between obligate-seeder and resprouter members of subseries Levispermae (Myrtaceae). Nuytsia 30; 247-252. Hammer, T.A., Davis, R.W. & Thiele, K.R. (2019). Of a dilferent feather; two new species of featherheads from the Ptilotus macrocephalus (Amaranthaceae) complex. Australian Systematic Botany 32; 61-70. Hickman, E.J. & Hopper, S.D. (2019). A revision of the tiumdins (Tribonanthes, Haemodoraceae). Nuytsia 30: 87-154. Hislop, M. (2019). Styphelia quartzitica and S. rectiloba (Ericaceae: Epacridoideae: Styphelieae), two new, morphologically anomalous species of restricted distribution. Nuytsia 30: 229-235. Hislop, M. & Puente-Eelievre, C. (2019). A taxonomic review of the Styphelia xerophylla group (Ericaceae: Epacridoideae: Styphelieae). VMyA/fl 30: 155-175. Jessup, E.W. (2019). A taxonomic revision of Sapotaceae for mainland Australia. Austrobaileya 10: 321-382. Jones, D.E. & French, C.J. (2019). a new species in the complex (Orchidaceae: Diurideae) from Western Australia with aflinities to Diuris brumalis. Australian Orchid Review 84(2): 32-34. Keighery, G.J. (1999). Two new species of Actinotus (Apiaceae) from Western Australia. Nuytsia 13: 23-27. Keighery, G.J. (2005). New and noteworthy plant species recognised as naturalised in Western Australia. Nuytsia 15: 523-528. Eowrie, A. (2013). Carnivorous plants of Australia magnum opus. Vol. 1. (Redfern Natural History Productions: Poole, United Kingdom.) Nicolle, D., George, A. & Wilson, PG. (2019). Application of the name Eucalyptus obtusiflora DC. (Myrtaceae). Telopea 22: 61-66. Obbens, F.J. (2019). Calandrinia monosperma and C. uncinella (Montiaceae), two new indehiscent species from Western Australia. Nuytsia 30: 237-245. Orchard, A.E. (2017). A revision of Cassinia (Asteraceae: Gnaphalieae) in Australia. 7. Cassinia subgenus Achromolaena. Australian Systematic Botany 30: 337-370. Paton, A.J., Mwanyambo, M., Govaerts, R.H.A., Smitha, K., Suddee, S., Phillipson, P.B., Wilson, T.C., Forster, PI. & Culham, A. (2019). Nomenclatural changes 'mColeusmdPlectranthus{Ldim\diCQdiQy. ataleofmore than two genera.P/?ytofey5129; 1-158. Perkins, A.J. (2018). Hydrocotyle eichleri, H. papilionella and H. tuberculata (Araliaceae), three new annual species from Western Australia. Nuytsia 29: 233-243. Perkins, A.J. (2019). Molecular phylogenetics and species delimitation in annual species of Hydrocotyle (Araliaceae) from South Western Australia. Molecular Phylogenetics and Evolution 134: 129-141. Roalson, E.H., Barrett, R.E., Wilson, K.E., Bruhl, J.J. & Earridon, I. (2019). Crosslandia setifolia is a partly monoecious species of Fimbristylis (Abildgaardieae: Cyperaceae). Phytotaxa 399: 163-166. Rye, B.E. (2018). Placement of three names in Cyathostemon, Hysterobaeckea and Oxymyrrhine (Myrtaceae: Chamelaucieae). Nuytsia 29: 269-270. Rye, B.E. (2019). An update to the taxonomy of some Western Australian genera of Myrtaceae tribe Chamelaucieae. 6. Scholtzia. Nuytsia 30: 33-86. Smith, M.G. & Jones, A. (2018). Threatenedand Priority Floralist 5 December 2018. Department of Biodiversity, Conservation and Attractions, https://www.dpaw.wa.gov.au/plants-and-animals/threatened-species-and-communities/threatened-plants [accessed 2 December 2019]. Sokoloff, D.D., Marques, I., Macfarlane,T.D., Remizowa, M. V, Earn, VK. Y, Pellicer, J , Hidalgo, O., Rudall, P. J. & Graham, S.W. (2019). Cryptic species in an ancient flowering-plant lineage (Hydatellaceae, Nymphaeales) revealed by molecular and micromorphological data. Taxon 6^: 1-19. Telford, I.R.H., Pruesapan, K., Welzen, PC. van & Bruhl, J.J. (2019). Morphological and molecular data show an enlarged tropical Australian radiation in Synostemon (Phyllanthaceae, Phyllantheae) previously concealed by heteromorphic species concepts. Australian Systematic Botany 32: 146-176. Thiele, K.R. (2019a). A revision of the Hibbertia commutata (Dilleniaceae) species group. Australian Systematic Botany 32; 71-109. C.M. Parker & J.M. Percy-Bower, Updates to Western Australia’s vascular plant census for 2019 351 Thiele, K.R. (2019b). Two new Western Australian species segregated from Banksia densa (Proteaceae). Nuytsia 30: 203-214. Thiele, K.R. & Schilling, E.E. (2018). Cyanthillium gracZ/A, a new combination forthe WestemAustralian endemic P/ewracarpaea gracilis (Asteraceae; Vemonieae). Nuytsia 29: 229-232. Wang, J. & Bean, A. R. (2019). Ataxonomic revision of Lagenophora Cass. (Asteraceae) in Australia. Austrobaileya 10:405-442. Western Australian Herbarium (1998-). FloraBase—the Western Australian Flora. Department of Biodiversity, Conservation and Attractions, https://florabase.dpaw.wa.gov.au/ [accessed 2 December 2019]. C.M. Parker^ and J.M. Percy-Bower Western Australian Herbarium, Biodiversity and Conservation Science, Department of Biodiversity, Conservation and Attractions Eocked Bag 104, Bentley Delivery Centre, Western Australia 6983 ‘Corresponding author, email: Cheryl.Parker@dbca.wa.gov.au 352 Nuytsia Vol. 30 (2019) Nuytsia Vol. 30 (2019) 353 Referees for Volume 30 The assistance of referees in providing expert review of papers submitted to Nuytsia is gratefully acknowledged. The referees consulted for Volume 30 include those listed below and six anonymous reviewers. Each paper was also refereed internally by Nuytsia Editorial Committee members. Benjamin Anderson Fred Barrie Anna Monro Marco Pellegrini Bevan Buirchell Barbara Rye Bob Chinnock Kelly Shepherd Mark Clements John Thompson Alex George Helen Kennedy Wendy Thompson Juliet Wege Bruce Maslin 354 Nuytsia Vol. 30 (2019) CONSERVATION CODES for Western Australian Flora and Fauna Threatened, Extinct and Specially Protected fauna or flora' are species^ which have been adequately searched for and are deemed to be, in the wild, threatened, extinct or in need of special protection, and have been gazetted as such. The Wildlife Conservation (Specially Protected Fauna) Notice 2018 and the Wildlife Conservation (Rare Flora) Notice 2018 have been transitioned nnder regnlations 170, 171 and 172 of the Biodiversity Conservation Regulations 2018 to be the lists of Threatened, Extinct and Specially Protected species nnder Part 2 of the Biodiversity Conservation Act 2016. Categories of Threatened, Extinct and Specially Protected fauna and flora are: T Threatened species Eisted by order of the Minister as Threatened in the category of critically endangered, endangered or vulnerable under section 19(1), or is a rediscovered species to be regarded as threatened species under section 26(2) of the Biodiversity Conservation Act 2016 (BC Act). Threatened fanna is that subset of ‘Specially Protected Fauna’ listed under schedules 1 to 3 of the Wildlife Conservation (Specially Protected Fauna) Notice 2018 for Threatened Fauna. Threatened flora is that subset of‘Rare Flora’listed under schedules 1 to 3 of the Wildlife Conservation (Rare Flora) Notice 2018 for Threatened Flora. The assessment of the conservation status of these species is based on their national extent and ranked according to their level of threat using lUCN Red Fist categories and criteria as detailed below. CR Critically endangered species Threatened species considered to be 'facing an extremely high risk of extinction in the wild in the immediate future, as determined in accordance with criteria set out in the ministerial guidelines'’'. Fisted as critically endangered under section 19(1 )(a) of the BC Act in accordance with the criteria set out in section 20 and the ministerial guidelines. Published under schedule 1 of the Wildlife Conservation (Specially Protected Fauna) Notice 2018 for critically endangered fauna or the Wildlife Conservation (Rare Flora) Notice 2018 for critically endangered flora. EN Endangered species Threatened species considered to be 'facing a very high risk of extinction in the wild in the nearfuture, as determined in accordance with criteria set out in the ministerial guidelines’’'. 'The definition of fiora includes algae, fungi and lichens. ^Species includes all taxa (plural of taxon - a classificatory group of any taxonomic rank, e.g. a family, genus, species or any infraspecific category i.e. subspecies or variety, or a distinct population). 355 Listed as endangered under section 19(l)(b) of the BC Act in accordance with the criteria set out in section 21 and the ministerial guidelines. Published under schedule 2 of the Wildlife Conservation (Specially Protected Fauna) Notice 2018 for endangered fauna or the Wildlife Conservation (Rare Flora) Notice 2018 for endangered flora. VU Vulnerable species Threatened species considered to be "facing a high risk of extinction in the wild in the medium-term future, as determined in accordance with criteria set out in the ministerial guidelines’’". Listed as vulnerable under section 19(l)(c) of the BC Act in accordance with the criteria set out in section 22 and the ministerial guidelines. Published under schedule 3 of the Wildlife Conservation (Specially Protected Fauna) Notice 2018 for vulnerable fauna or the Wildlife Conservation (Rare Flora) Notice 2018 for vulnerable flora. Extinct Species Listed by order of the Minister as extinct under section 23(1) of the BC Act as extinct or extinct in the wild. EX Extinct species Species where ""there is no reasonable doubt that the last member of the species has died”, and listing is otherwise in accordance with the ministerial guidelines (section 24 of the BC Act). Published as presumed extinct under schedule 4 of the Wildlife Conservation (Specially Protected Fauna) Notice 2018 for extinct fauna or the Wildlife Conservation (Rare Flora) Notice 2018 for extinct flora. EW Extinct in the wild species Species that ""is known only to survive in cultivation, in captivity or as a naturalised population well outside its past range; and it has not been recorded in its known habitat or expected habitat, at appropriate seasons, anywhere in its past range, despite surveys over a time frame appropriate to its life cycle and form”, and listing is otherwise in accordance with the ministerial guidelines (section 25 of the BCAct). Currently there are no threatened fauna or threatened flora species listed as extinct in the wild. If listing of a species as extinct in the wild occurs, then a schedule will be added to the applicable notice. Specially protected species Listed by order of the Minister as specially protected under section 13(1) of the BC Act. Meeting one or more of the following categories: species of special conservation interest; migratory species; cetaceans; species subject to international agreement; or species otherwise in need of special protection. Species that are listed as threatened species (critically endangered, endangered or vulnerable) or extinct species under the BC Act cannot also be listed as Specially Protected species. 356 Nuytsia Vol. 30 (2019) MI Migratory species F auna that periodically or occasionally visit Australia or an external Territory or the exclusive economic zone; or the species is subject of an international agreement that relates to the protection of migratory species and that binds the Commonwealth; and listing is otherwise in accordance with the ministerial guidelines (section 15 of the BC Act). Includes birds that are subj ect to an agreement between the government of Austral ia and the governments of Japan (JAMBA), China (CAMBA) and The Republic of Korea (ROKAMBA), and fauna subject to the Convention on the Conservation of Migratory Species of Wild Animals (Bonn Convention), an environmental treaty under the United Nations Environment Program. Migratory species listed under the BC Act are a subset of the migratory animals, that are known to visit Western Australia, protected under the international agreements or treaties, excluding species that are listed as Threatened species. Published as migratory birds protected under an international agreement under schedule 5 of the Wildlife Conservation (Specially Protected Fauna) Notice 2018. CD Species of special conservation interest (conservation dependent fanna) Fauna of special conservation need being species dependent on ongoing conservation intervention to prevent it becoming eligible for listing as threatened, and listing is otherwise in accordance with the ministerial guidelines (section 14 of the BC Act). Published as conservation dependent fauna under schedule 6 of the Wildlife Conservation (Specially Protected Fauna) Notice 2018. OS Other specially protected species Fauna otherwise in need of special protection to ensure their conservation, and listing is otherwise in accordance with the ministerial guidelines (section 18 of the BC Act). Published as other specially protected fauna under schedule 7 of the Wildlife Conservation (Specially Protected Fauna) Notice 2018. P Priority species Possibly threatened species that do not meet survey criteria, or are otherwise data deficient, are added to the Priority Fauna or Priority Flora Lists under Priorities 1,2 or 3. These three categories are ranked in order of priority for survey and evaluation of conservation status so that consideration can be given to their declaration as threatened fauna or fiora. Species that are adequately known, are rare but not threatened, or meet criteria for near threatened, or that have been recently removed from the threatened species or other specially protected fauna lists for other than taxonomic reasons, are placed in Priority 4. These species require regular monitoring. Assessment of Priority codes is based on the Western Australian distribution of the species, unless 357 the distribution in WA is part of a contiguous population extending into adjacent States, as defined by the known spread of locations. 1 Priority 1: Poorly-known species Species that are known from one or a few locations (generally five or less) which are potentially at risk. All occurrences are either: very small; or on lands not managed for conservation, e.g. agricultural or pastoral lands, urban areas, road and rail reserves, gravel reserves and active mineral leases; or otherwise under threat of habitat destruction or degradation. Species may be included if they are comparatively well known from one or more locations but do not meet adequacy of survey requirements and appear to be under immediate threat from known threatening processes. Such species are in urgent need of further survey. 2 Priority 2: Poorly-known species Species that are known from one or a few locations (generally five or less), some of which are on lands managed primarily for nature conservation, e.g. national parks, conservation parks, nature reserves and other lands with secure tenure being managed for conservation. Species may be included if they are comparatively well known from one or more locations but do not meet adequacy of survey requirements and appear to be under threat from known threatening processes. Such species are in urgent need of further survey. 3 Priority 3: Poorly-known species Species that are known from several locations, and the species does not appear to be under imminent threat, or from few but widespread locations with either large population size or significant remaining areas of apparently suitable habitat, much of it not under imminent threat. Species may be included if they are comparatively well known from several locations but do not meet adequacy of survey requirements and known threatening processes exist that could affect them. Such species are in need of further survey. 4 Priority 4: Rare, Near Threatened and other species in need of monitoring (a) Rare. Species that are considered to have been adequately surveyed, or for which sufficient knowledge is available, and that are considered not currently threatened or in need of special protection but could be if present circumstances change. These species are usually represented on conservation lands. (b) Near Threatened. Species that are considered to have been adequately surveyed and that are close to qualifying for vulnerable but are not listed as Conservation Dependent. (c) Species that have been removed from the list of threatened species during the past five years for reasons other than taxonomy.