Nuytsia The journal of the Western Australian Herbarium 28:229-246 Published online 8 June 2017 Stylidium miscellany 3: a synopsis of Robert Brown’s Stylidiaceae types and occasional notes on associated names Juliet A. Wege Western Australian Herbarium, Department of Parks and Wildlife, Locked Bag 104, Bentley Delivery Centre, Western Australia 6983 Email: [email protected]. gov. au Abstract Wege, J.A. Stylidium miscellany 3: a synopsis of Robert Brown’s Stylidiaceae types and occasional notes on associated names. Nuytsia 28: 229-246 (2017). A synopsis of the 39 species of Stylidiaceae named by Robert Brown is presented and a full synonymy provided. Lectotypes are selected for the following Brown names: Levenhookiapusilla R.Br., Stylidium alsinoides R.Br., S. articulatum R.Br., S. assimile R.Br., S. caespitosum R.Br., S. calcaratum R.Br., S. corymbosum R.Br., S. crassifolium R. Br., S. diversifolium R.Br., S. eriorhizum R.Br., S. fasciculatum R.Br., S. floribundum R.Br., S. guttatum R.Br., S. tenuifolium R.Br. (= S. laricifolium Rich.), S. luteum R.Br., S. pedunculatum R. Br., S. reduplicatum R.Br. (= S. pilosum (Labill.) Labill.), S. spinulosum R.Br., S. tenellum R.Br. (= S. tenerum Spreng.) and S. violaceum R.Br. Stylidium robustum Sond. is newly synonymised under S. diversifolium and lectotypes are selected for two additional synonyms of this species, S. marginatum Sond. and S.pruinosum Sond. Stylidium pilosum is lectotypified and a neotype designated for A pilosum var. brevius E.Pritz. Comments on the uncertain circumscription of S. armeria (Labill.) Labill. (of which S. melastachys R.Br. is a synonym) are provided. Introduction As the naturalist on board the HMS Investigator voyage under the command of Matthew Flinders, Robert Brown’s contribution to Australian botany is legendary (Stearn 1960; Mabberley 1985; Vallance et al. 2001; Moore 2005) and his research on the triggerplant family Stylidiaceae is no exception. His description of 38 species of Stylidium Sw. as part of his Prodromus (Brown 1810) remains an extraordinary taxonomic contribution to the genus, which is now known to comprise more than 300 species. He also named and circumscribed the family, which he insightfully defined on the basis of the reduced number of stamens, unique floral column and imbricate aestivation of the corolla lobes (Brown 1810, 1814), and described the genus Levenhookia R.Br. based on his collection of L. pusilla R.Br. Twenty-nine of the Stylidium species described by Brown occur in south-western Australia, a centre of diversity for the genus. Indeed, the Investigator anchorage at King George Sound (Albany) from 8 December 1801 to 5 January 1802 (Vallance etal. 2001) was auspicious in terms of Stylidium—not only is the greater Albany region an area of high triggerplant diversity, but many species are in flower in this region at this time. This is in stark contrast to many other genera, which are well past their peak spring-flowering period by then (Keighery & Gibson 2005). Brown had ample opportunity to explore and collect plants from a range of habitats and it is a testament to his skills that he managed to collect the majority of species known from the region; almost all of the species that he did not find © Department of Parks and Wildlife 2017 ISSN 2200-2790 (Online) https://florabase.dpaw.wa.gov.au/nuytsia/ ISSN 0085-4417 (Print) 230 Nuytsia Vol. 28 (2017) were either past flowering or require disturbance to stimulate flowering. However, Brown was not infallible. Examination of his Stylidium types has revealed three clear instances of mixed gatherings or mixed species concepts: S. scandens R.Br. comprised separate collections of S. scandens and S. nymphaeum Wege (Wege 2010), S.junceum R.Br. was based on a mixed gathering of S.junceum, S. squamosotuberosum Carlquist and S. thryonides Wege (Wege 2014), and S. fasciculatum R.Br. was based on a mixed gathering of S. fasciculatum and S. adnatum R.Br. (see under S. fasciculatum below). It is also possible that his type gathering of S. articulatum R.Br. includes a fragment of S. amoenum R.Br. (see under S. articulatum below). The following synopsis of Brown’s Stylidiaceae names and types is published in preparation of an account of the family for Flora of Australia and as an aid in the preparation of a forthcoming publication by David Mabberley and David Moore, which will provide a comprehensive register of Brown’s plant names and types (Mabberley & Moore 2007). This paper also provides an opportunity to provide additional taxonomic or typification notes on associated names. Methods For those species based solely on Brown’s own collections, the following approach has been adopted. Where only one sheet has been located (i.e. for S. adnatum, S. amoenum, S. breviscapum R.Br., S.falcatum R.Br., S. propinquum R.Br. and S. rotundifolium R.Br.), it is treated as the holotype. These specimens have a blue Iter Australiense label (prepared according to the instructions of J. J. Bennett), indicating that they originated from Brown’s own herbarium (Mabberley 1985), and in each case they bear Brown’s original field labels as well as an annotation by Brown indicating the page number for that taxon in his Prodromus. Where more than one sheet has been found, the material at BM from Brown’s own herbarium (as described above) is designated as an appropriate lectotype; invariably these specimens are the best and most complete of the available material. Of these, there are several species (Levenhookia pusilla, S. assimile R.Br., S. diversifolium R.Br., S. inundatum R.Br. and S. reduplicatum R.Br.) for which the only material known is the lectotype and one or more ‘ Dryander duplicates’ (see explanation in Vallance et al. 2001: 14). The Dryander duplicates were unlikely to have used by Brown for his Prodromus descriptions, in which case they could be interpreted as isotypes (D.J. Mabberley pers. comm.). However, all material was used to infonn Brown’s descriptive slips (his plant descriptions made on the expedition and now held at BM) and it remains a possibility that he consulted these slips when compiling his Stylidiaceae account (e.g. for detail concerning the presence or absence of appendages on the corolla lobes and labellum, which can be difficult to see in the dried state). While Brown’s descriptions were based primarily on his own material, three species of Stylidium (S. alsinoides R.Br., S. capillare R.Br. and S. pedunculatum R.Br.) were based solely, or in part, on material collected by Joseph Banks and Daniel Solander from Endeavour River (Queensland) in 1770, while S. despectum R.Br. was based on a collection made by William Paterson from Tasmania in 1805 (Wege 2011). Only one sheet has been located for both S. capillare and S. despectum and accordingly they are treated as holotypes. For both S. alsinoides and S. pedunculatum, the primary specimen in Banks’ Herbarium (annotated by Brown) has been designated as the lectotype since it is of better quality than the material that formed part of Brown’s ‘study set’ that he took to Australia, although the latter may also have been used in descriptions made on board Investigator. Notes on lectotypifications of taxa named by authors other than Brown are included under each species. J.A. Wege, Stylidium miscellany 3 (Stylidiaceae) 231 Typifications and miscellany Levenhookia pusilla R.Br., Prodr. FI. Nov. Holland. 573 (1810). Type citation: ‘(M.) v. v.’ Type specimen', near the observatory, Princess Royal Harbour, King George’s Sound [Western Australia], 21 December 1801, R. Brown s.n. [Bennett No. 2613] (lectotype, here designated: BM 001041273!; isolectotype: BM 000948765!). Stylidium adnatum R.Br., Prodr. FI. Nov. Holland. 572 (1810). Candollea adnata (R.Br.) F.Muell., Syst. Census Austral. PI.: 86 (1882). Type citation. ‘(M.) v. v. ’ Type specimen. Goose Island Bay larger island [Middle Island, Western Australia], January 1802, R. Brown s.n. [Bennett No. 2611] (holotype: BM 000812588!). Stylidium propinquum R.Br., Prodr. FI. Nov. Holland. 572 (1810). Stylidium adnatum R.Br. var. propinquum (R.Br.) R.Br. in Sims, Bot. Mag. 52, t. 2598 (1825). Stylidium adnatum var. abbreviatum Benth., FI. Austral. 4: 33 (1868), nom. illeg., nom. superfl. Candollea adnata var. abbreviata (Benth.) De Wild., Icon. Horti Then. 2: 9, t. 43 (1901), nom. illeg. Type citation: ‘(M.) v. v.’ Type specimen: King George’s Sound [Western Australia], December 1801-January 1802, R. Brown s.n. [Bennett No. 2612] {holotype. BM 000812587!). Notes. The holotype of S. adnatum is a single, fairly robust individual with numerous stems that arise from a lignotuberous stock. This species, which is widespread in southern Western Australia (Western Australian Herbarium 1998-), is usually geophytic, with plants dying back to a lignotuber over the dry, summer months and resprouting following autumn rains. Occasionally, the above-ground stems persist (presumably if there is sufficient water available over summer), with the apical stem node giving rise to new shoots and occasionally adventitious roots. Apical shoots are evident in the left hand individual on the holotype of S. propinquum; material that differs further from the holotype of S. adnatum in having shorter and narrower stems and, for the most part, shorter scapes with fewer flowers. Brown later treated this taxon as variety of S. adnatum, noting the two forms share an ovary that is sterile in one loculus (Sims 1825). Although Brown’s variety was recognised by Erickson (1958), the illegitimate name S. adnatum var. abbreviatum has been inexplicably used in Western Australia for many years (Paczkowska & Chapman 2000; Western Australian Herbarium 1998-). Stylidium adnatum is a variable species in terms of its habit and overall robustness (e.g. stem length and width, inflorescence length and flower number) and var. propinquum is not worth recognising. Stylidium alsinoides R.Br., Prodr. FI. Nov. Holland. 572 (1810). Candollea alsinoides (R.Br.) F.Muell., Syst. Census Austral. PL . 86 (1882). Type citation. ‘(T.) B. v. sT Type specimen Endeavour River [Queensland, June-August], 1770, J. Banks & D. Solander s.n. {lectotype, here designated: BM 000645715!; isolectotypes. BM 001041266! [study set], NSW 133570 n.v.). Typification. Prior to 1 January 2001, if an author stated that a particular specimen was the holotype of a previously published name but other specimens of the cited gathering existed, then this was an error to be corrected to lectotype (see McNeill 2014); however, Bean’s (2000: 635) citation ‘holo: ?BM n.vT cannot be corrected in this manner since there are two Banks and Solander duplicates housed at BM (neither of which were seen or annotated by Bean). StylidiumamoenumR.Br.,Prodr. FI. Nov. Holland. 570(1810). Candolleaamoena(R.Br.)F.Muell., Syst. Census Austral. PL : 86 (1882). Type citation: ‘(M.) v. v.’ Type specimen: King George’s Sound [Western Australia], December 1801, R. Brown s.n. [Bennett No. 2592] {holotype: BM 000903578!). 232 Nuytsia Vol. 28 (2017) Notes. Refer to the notes under S. articulatum regarding material of S. amoenum on the CANB isolectotype of that species. Stylidium armeria (Labill.) Labill., Nov. Holl. PI. Spec. 2: 66, t. 216 (1806). Candollea armeria Labill., Ann. Mus. Hist. Nat. 6: 455 (1805). Type: ‘Habitat in capite van-Diemen’ [May 1792, J.J.H. de Labillardiere s.n.] (syntypes. BM 000894111! [individuals labelled ‘B’], B-W 17048-010 image!, FI 006811!, G 00354007!, G-DC G 00493487 image!, ?MPU 016343 image!). Stylidium melastachys R.Br., Prodr. FI. Nov. Holland. 568 (1810). Type citation. ‘(D.) v. v.’ Type specimen Kent’s Group, Bass Straight, December 1803, R. Brown s.n. [Bennett No. 2577] (lectotype fide E. J. Raulings & P.Y. Ladiges, A wVra/. Syst. Bot. 14(6): 929 (2001): BM 000603724!; isolectotypes: BM 000603725!; DBN n.v., E 00279234!, E 00279235!, E 00279236!, K 000060478!, MEL 2257703!, MEL 2257704!, MEL 2257705!, P 00712398 image!). Stylidium dilatatum W.D.Jacks. & R.J.E.Wiltshire, Austral. Syst. Bot. 14(6): 961, 964, 965 (2001). Type ', ridge west of Proctors Road, opposite The Lea, Tasmania (holotype: HO 22793!; isotypes'. BM n.v., HO 22790!, MEL 2330860 image!). Typification. Lectotypification of S. armeria should only be considered once the taxonomy of the S. graminifolium Sw. complex is resolved (see notes below). All duplicates of Labillardiere’s gathering are therefore treated here as syntypes regardless of how they are annotated or databased. Note that FI 006811, which I have mistakenly annotated as the holotype, bears extensive annotations by Labillardiere. My annotation slip on this specimen should read ‘flowering specimens possess two broad channels of stomata on lower [not upper] surface and lack stomata on upper [not lower] surface’. I also observed stomata towards the apex of the upper leaf surface in the fruiting individual that is second from the left. Notes. Stylidium armeria—a species from the widespread and morphologically variable south-eastern Australian S. graminifolium complex—was simultaneously reinstated by Jackson and Wiltshire (2001) and Raulings and Ladiges (2001), albeit with differing circumscriptions (both studies treated Brown’s S. melastachys as a synonym). It is not my aim here to provide a comprehensive commentary on the discrepancies within and differences between these two publications, nor a solution to the taxonomic confusion that has been created by their publication. This group of triggerplants (which, in addition to the taxa referred to herein, includes the New South Wales endemic S. productum Hindm. & Blaxell) exhibits complex patterns of morphological and ecological variation and I have limited field knowledge of them so far. Moreover, I have made only a cursory examination of a fraction of the specimens available for study in Australian herbaria. The following brief summary simply serves to highlight some of the taxonomic issues that need resolving and the scope of research that will need to be conducted in order to do so. The study by Jackson and Wiltshire (2001), which had a Tasmanian focus, redefined S. graminifolium as a diploid species (n = 15,2n = 30) widespread on infertile soils or sites of high evapotranspirational stress in summer and autumn, and characterised by narrow (1 to almost 3 mm wide), bifurrowed leaves. They describe its frequent parapatric distribution in Tasmania with broader-leaved, tetraploid cytotypes (2n = 60) that favour fertile soils or sites of low evapotranspiration. The study by Raulings and Ladiges (2001), which had a Victorian focus and did not investigate ploidy levels, similarly delineated S. graminifolium as a species from sandy habitats with narrow (1-2.5 mm wide), bifurrowed leaves. The leaves of the broader-leaved forms (variously described as S. armeria, S. dilatatum or S. montanum Raulings & Ladiges) were described in both studies as >3 mm wide and flat in section. J.A. Wege, Stylidium miscellany 3 (Stylidiaceae) 233 While at first glance these studies appear to be in broad agreement with respect to the circumscription of S. graminifolium, there are some significant discrepancies, not limited to the following three examples. Firstly, Raulings and Ladiges (2001) consider the regular serrations on the leaf margins to be taxonomically informative, with the broad-leaved forms either having smooth margins or occasional serrations that are mostly confined to near the tip; however, Jackson and Wiltshire (2001), state that this feature is taxonomically unreliable, describing S. graminifolium as variably serrulate, with the margins sometimes smooth or serrulate near the apex. Secondly, scape width is described by Raulings and Ladiges as >2 mm and diagnostic with respect to S. montanum (<2 mm wide) but not S. armeria (>2 mm), whereas Jackson and Wiltshire describe it as narrower (mean 1.5 mm; upper scape c. 1 mm) and diagnostic with respect to S. dilatatum (mean 3.2 mm; upper scape c. 2 mm) and S. armeria (mean 4.4 mm; upper scape c. 4 mm wide). Thirdly, Raulings and Ladiges consider throat appendage morphology (specifically the reduced size of the anterior-most appendages) as a useful character upon which to separate S. graminifolium from S. armeria (but not S. montanum), but Jackson and Wiltshire state that throat appendage morphology is uninformative. The two studies differ even further with respect to the broader-leaved forms (leaves >3mm wide). Jackson and Wiltshire (2001) recognised two tetraploid species, S. armeria s. str. and S. dilatatum, the former restricted to littoral habitats of highly exposed coasts in Tasmania and on islands of the Bass Strait (and ‘probably extending to about Twofold Bay in NSW’), and the latter widespread in more inland areas of south-eastern Australia and Tasmania. They were differentiated by leaf shape (spathulate in S. armeria s. str. and linear in S. dilatatum) and stomata distribution (confined to the lower leaf surface in S. armeria s. str. and present on both surfaces in S. dilatatum), with the flowers of S. armeria s. str. noted to be c. 20% larger (although measurement ranges were not provided for either species). Additional diagnostic information for S. dilatatum was mostly provided with respect to S. graminifolium and consequently taxonomic placement of broad-leaved herbarium specimens has proven difficult (Gray 2011). Of note is the contradictory information given by Jackson and Wiltshire with respect to the distribution of stomata in S. armeria s. str. (e.g. the description on p. 960 indicates stomata can in fact be present on the upper leaf surface) and the existence of collections from well outside the cited geographic range of S. armeria s. str. which appear to lack stomata on the upper leaf surface (e.g. Hj. Eichler 14634, Bogong High Plains, CANB). The broad-leaved form was broadly circumscribed by Raulings and Ladiges (2001) under S. armeria, with the exception of S. montanum, a species they recognised from subalpine and alpine habitats in Victoria, New South Wales and Tasmania that grows in close proximity to S. armeria on the Bogong High Plains and at Mt Buffalo. Stylidium montanum was said to be characterised by its relatively short, pale to mid-green leaves (4-15 cm long cf. 15-40 cm and dark green in S. armeria), shorter, narrower scapes (10—45 cm high and 2 mm wide cf. 50-100 cm high and >2 mm wide in S. armeria) with fewer flowers (10-30 cf. (25-)30-100 in S. armeria) and a sparser indumentum (glabrous or glabrescent below the lowest flower cf. glandular-hairy below the lowest flower in S', armeria). Jackson and Wiltshire (2001), who were aware of the taxonomic studies being conducted by Raulings as part of her PhD dissertation, refer to S. montanum on p. 953 (as Stylidium sp. nov.) and suggest that it is a tetraploid (2n = 60) and does not occur in Tasmania. Since no specimens of S. montanum from Tasmania were cited by Raulings and Ladiges, it is not recognised in that state (Council of Heads of Australasian Herbaria 2011a). In Tasmania, S. armeria and S. dilatatum are considered insufficiently distinct to warrant recognition of the latter species (Gray 2011). A broad concept of S. armeria is also followed in Victoria, where Best et al. (2009) described a Critically Endangered subspecies, S. armeria subsp. pilosifolium R. J.Best, D.E.Francis & N.G. Walsh, with eglandular and glandular hairs on both leaf surfaces. In preparation for 234 Nuytsia Vol. 28 (2017) a treatment of Stylidiaceae for Flora of South Australia, I have applied Raulings and Ladiges’ broad concept of S. armeria to South Australian material at AD; however, I did not have time to examine their holdings of this complex from other states. Indeed, there are a significant number of collections at AD, CANB and NSW that have not been formally assessed. I recently conducted a very preliminary sort of material at CANB which indicated that there is considerably more variation in this species complex than has been accounted for in either study. The reliability of leaf width to separate S. graminifolium (<3 mm) from S. armeria s. lat. and S. montanum (>3 mm) must be questioned in view of the significant number of specimens that intergrade. These include specimens with leaves <3 mm that otherwise have features considered by one or both studies as characteristic of S. armeria or S. montanum (e.g. leaves flat in T.S., leaf margin entire, corolla lobes large, anterior-most throat appendages conspicuous and column very long). Jackson and Wiltshire (2001) describe a higher density of smaller stomata in S. graminifolium s. str. as compared with the polyploidy species S. armeria and S. dilatatum, a feature that could perhaps inform taxonomic work if S. armeria s. lat. is indeed polyploid throughout its range, but this must also be questioned in view of recent ecological studies of S. armeria s. lat. that have indicated populations in alpine Victoria are diploid (Hoffman et al. 2009). While both Raulings and Ladiges (2001) and Jackson and Wiltshire (2001) make insightful observations at a local scale, resolution of the taxonomy of the S. graminifolium complex demands a geographically comprehensive, modem and integrative approach that uses multiple lines of evidence (molecular, morphological, cytological and ecological) and includes examination and curation of the more than 2,500 specimens currently available in Australian herbaria. This is not something that I will be able to pursue from Western Australia. StylidiumarticulatumR.Br.,Prodr. FI.Nov. Holland. 570(1810). Candolleaarticulata(R.Br.)F.Muell., Syst. Census Austral. PL . 86 (1882). Type citation. ‘(M.) v. v.’ Type specimens. King George’s Sound [Western Australia], 3 January 1801, R. Brown s.n. [Bennett No. 2593] (lectotype, here designated: BM 000903579!; isolectotypes: BM 001041332!, CANB 279057! [right hand individual], E 00279199!, E 00279200!, K 000060586!, K 000060587!). ?Paralectotype [syntype]: King George’s Sound [Western Australia], 3 January 1801, R. Brown s.n. (CANB 279057! [left hand individual]), =S. amoenum R.Br. Typification. CANB 279057 consists of a complete individual of S. articulatum on the right hand side together with a fragment of S. amoenum. It is unclear whether the latter is type material of S. amoenum (due to an error that has occurred post-Prodromus) or indeed whether Brown made a mixed gathering. I have observed both species growing in close proximity east of Albany suggesting the latter is not beyond the realms of possibility. Stylidium assimile R.Br., Prodr. FI. Nov. Holland. 569 (1810). Candollea assimilis (R.Br.) F.Muell., Syst. Census Austral. PI.: 86 (1882), nom. illeg. non Steud. (1845 = a name of uncertain application in Dilleniaceae). Type citation ‘(M.) v. v. ’ Type specimen Princess Royal Harbour, King George’s Sound [WesternAustralia], December 1801-January 1802, R. Browns.n. [BennettNo. 2584] {lectotype, here designated: BM 000797560!; isolectotype: BM 000797561!). Stylidium breviscapum R.Br., Prodr. FI. Nov. Holland. 572 (1810). Candollea breviscapa (R.Br.) F.Muell., Syst. Census Austral. PL . 86 (1882). Type citation ‘(M.) v.v.’ Type specimen. Bay I South Coast [Lucky Bay, Cape Le Grand, Western Australia, January 1802], R. Brown s.n. [Bennett No. 2608] (iholotype: BM 001041346!). J.A. Wege, Stylidium miscellany 3 (Stylidiaceae) 235 Stylidium breviscapum var. erythrocalyx Benth., FI. Austral. 4: 31 (1868). Type'. ‘S West end Cape arid Range’ [Western Australia], 5. dat., G. Maxwell s.n. {lectotype fide J.A. Wege, Nuytsia 20: 80 (2010): MEL 2046601!; isolectotype: K 000060836!). Paralectotype [syntype]: Fitzgerald Ranges, 5. dat., G. Maxwell 150 (BM!, MEL 672624!), = S. involucratum F.Muell. Stylidium caespitosum R.Br., Prodr. FI. Nov. Holland. 569 (1810). Candollea caespitosa (R.Br.) F.Muell., Syst. Census Austral. PI.: 86 (1882). Type citation'. ‘(M.) v. v. ’ Type specimen. Princess Royal Harbour, King George’s Sound [Western Australia], December 1801, R. Brown s.n. [Bennett No. 2582] {lectotype, here designated: BM 000797556!; isolectotypes'. BM 000797555! CANB 279058!, E 00279194!, E 00279195!, E 00279196!, FI 006806, K 000060801!, K 000060857!, MEL 02593 83!, P 00712385!). Stylidium calcaratum R.Br., Prodr. FI. Nov. Holland. 570 (1810). Candollea calcarata (R.Br.) F.Muell., Syst. Census Austral. PL. 86 (1882). Type citation ‘(M.) v. v.’ Type specimen. Princess Royal Harbour, King George’s Sound [Western Australia], December 1801, R. Brown s.n. [Bennett No. 2598] {lectotype, here designated: BM 000645735!; isolectotypes'. BM 000645736!, E 00279218!, FI 006805!, K 000060137!, K 000060139!, P 00712386!). Stylidium mimeticum Lowrie & Carlquist, Phytologia 71(1): 16 (1991), syn. nov. Type: In sand, along Great Northern Highway north of Bullsbrook, 1 km south of Wandena Road (south end) on east side of the Highway, Western Australia, 3 December 1989, Allen Lowrie 243 {holotype. PERTH 01643126!; isotypes'. PERTH 01643118!, RSAn.v.). Notes. Examination of the floral column in type material of S. calcaratum at BM and K has revealed a minute, obtuse appendage on the bend identical to that described for the later-named S. mimeticum (see Lowrie & Carlquist 1991: 19, Figure 7F), leading me to regard the latter as a conspecific (Western Australian Herbarium 1998-). My revision of S. calcaratum and allies (subg. Centridium Lindl.) is in an advanced stage of preparation and seeks to resolve the taxonomy of this group across Australia. Stylidium capillare R.Br., Prodr. FI. Nov. Holland. 570 (1810). Candollea capillaris (R.Br.) F.Muell., Syst. Census Austral. PI.: 86 (1882). Type citation. ‘(T.) B v. s. ’ Type specimen. Endeavour River, New Holland [Queensland, June-August] 1770, J. Banks & D. Solander s.n. {holotype'. BM 000563895!). Stylidium quadrifurcatum F.L.Erickson & J.H.Willis, Viet. Naturalistic. 5 (1956), syn. fide A.R. Bean, Austrobaileya 5(4): 620 (2000). Type'. Pine Creek, Northern Territory, April 1904, J.H. Niemann s.n. {holotype: MEL 1061651!). Stylidium corymbosum R.Br., Prodr. FI. Nov. Holland. 571 (1810). Candollea corymbosa (R.Br.) F.Muell., Syst. Census Austral. PL . 86 (1882). Type citation. ‘(M.) v. v.’ Type specimen. South Coast Bay 1 [Lucky Bay, Western Australia], January 1802, R. Brown s.n. [Bennett No. 2602] {lectotype, here designated: BM 000603873!; isolectotypes'. BM 000603874!, E 00279182!, E 00279183!, K 000060777!, K 000060778!). Stylidium crassifolium R.Br., Prodr. FI. Nov. Holland. 571 (1810). Candollea crassifolia (R.Br.) F.Muell., Syst. Census Austral. Pl.: 86 (1882). Type citation. C(M.) v. v. ’ Type specimen. Princess Royal Harbour, King George’s Sound [Western Australia], December 1802 [sic, 1801 ], R. Brown s.n. [Bennett No. 2601] {lectotype, here designated: BM 001041312!; isolectotypes'. CANB 279059!, E 00279259!). 236 Nuytsia Vol. 28 (2017) Stylidium leptobotrys DC., Prodr. 7(2): 783 (1839). Stylidium leptobotrydium St.-Lag., Ann. Soc. Bot. Lyon 7: 135 (1880), nom. illeg., nom. superfl. Type: ‘in Nova-Hollandia ad Swan-river’ [Western Australia, 1835-1838], J. Drummonds.n. {holotype: G-DC G00458493!; isotypes. BM 000812571!, FI 006802, ?G 00358766!, ?K 000060346!, K 000060351!). Dampiera inundata de Vriese, in Lehm., PI. Preiss. 1(3): 404 (1845). Type. ‘In subturfosis hieme aqua inundatis planitiei prope urbiculam Albany, Plantagenet [Western Australia], d. 29 m. Jan. 1840. Herb. Preiss. No. 1523’ (,syn. L 0001767 image!, L 0843043 image!, LD 1043796!). Notes. L 0001767 is labelled as the lectotype of Dampiera inundata and L 0843043 as an isolectotype but to my knowledge no lectotypification has been published. StylidiumdespectumR.Br.,Prodr. FI. Nov. Holland. 571 (1810). Candolleadespecta(R.Br.) F.Muel 1., Syst. Census Austral. PL. 86(1882). Type citation. ‘(D.)v. sA Type specimen Port Dalrymple [George Town, Tasmania], 1805, W. Paterson s.n. {holotype: BM 000645719!). Stylidium brachyphyllum Sond., in Lehm., PL Preiss. 1(3): 386 (1845). Candollea brachyphylla (Sond.) F.Muell., Syst. Census Austral. Pl.: 86 (1882). Type'. In solo turfoso ad Princess Royal Harbour [Albany, Western Australia], December 1840, L. Preiss 2239 {lectotype fide J.A. Wege, Austral. Syst. Bot. 24: 384 (2011): MEL2069492!; isolectotypes'. G00354009!, LD 1751955!, P00712379!, TCD! [as L. Preiss 453]). Paralectotype [syntype]'. In depressis uliginosis silvae prope oppidulum, Perth [Western Australia], 26 September 1839, L. Preiss 2248 (BR 0000005422593 image!, FI 113175!, G 00354004!, G 00354006!, G 00354008!, L 0001764 image!, L 0001765 image!, LD 1097540!, M 0175802!, MEL 2069490!, MEL 2069491!, MO-797443 image!, P 00712380!, W!). Stylidium diffusum R.Br., Prodr. FI. Nov. Holland. 571 (1810). Candollea diffusa (R.Br.) F.Muell., Syst. Census Austral. PL . 86 (1882). Type citation ‘(T.) v. v.’ Type specimen Shoal water bay [Queensland], 1802-1805 [3 September 1802],R. Brown s.n. [Bennett No. 2605] {lectotypefide A.R. Bean, Austrobaileya 5(4): 627 (2000): BM 000563899!; isolectotypes'. E 00279231!, K 000060550!). Typification. Bean’s (2000: 627) type citation for S. diffusum (‘holo: BM’) is corrected herein to reflect his inadvertent lectotypification (see McNeill 2014). Stylidium diversifolium R.Br., Prodr. FI. Nov. Holland. 570 (1810). Candollea diversifolia (R.Br.) F.Muell., Syst. Census Austral. Pl.: 86 (1882). Type citation ‘(M.) v. v. ’ Type specimen King George’s Sound [Western Australia], December 1801, R. Brown s.n. [Bennett No. 2594] (,lectotype, here designated: BM 000903576!; isolectotypes'. BM 001041333!, BM 001041334!). Stylidium robustum Sond., in Lehm., Pl. Preiss. 1(3): 378 (1845), syn. nov. Type'. In arenosis districtus Sussex [Western Australia], 22 December 1839, L. Preiss 2235 {syn LD 1745623!, MEL 293341!, MEL 2235239!). Stylidium marginatum Sond., in Lehm., Pl. Preiss. 1(3): 379 (1845). Type specimens. In arenosis colliculosis sylvae prope oppidulum Guildford [Western Australia], 13 September 1839, L. Preiss 2232 (ilectotype, here designated: MEL 2258906! [Sheet 1 of2], MEL2258907! [Sheet2 of2]; isolectotypes'. G 00358783!, G 00358784!, L 0001773 image!, LD 1753748!, MO-797518 image!, P 00712392!, W! [2 sheets]); Swan River [Western Australia, 1841], J. Drummond 529 {paralectotypes [syntypes]: BM 000903577!, K 000060680!, K 000060684!, MEL 2258911! P 03025123!, W! [2 sheets]). J.A. Wege, Stylidium miscellany 3 (Stylidiaceae) 237 Stylidiumpruinosum Sond., inLehm., PI. Preiss. 1 (3): 379 (1845). Type. In subturfosis hieme inundatis haud longe a Kaudiup, Plantagenet [WesternAustralia], 24 November 1840, L. Preiss 2236 (lectotype, here designated: MEL 2258905!; isolectotypes: BR 0000005422555 image!, G-DC G00358785!, G 00358786!, G 00358787!, GOET 011204 image!, LD 1731539!, M 0175794!, MEL 2258903!, MEL 2258904!, MO-797526 image!, P 00712393! W! [2 sheets]). Typification. Mildbraed (1908) was correct to place both S. marginatum and S. pruinosum in the synonymy of S. diversifolium, although he failed to consider S. robustum, which is added to the synonymy presented herein. Sonder examined all of the known type material of S. robustum, of which LD 1745623 and MEL 293341 both comprise an individual in fruit while MEL 2235239 consists of three dissected corollas and columns. The two MEL sheets are from Sonder’s personal herbarium and it is of note that the tip of the inflorescence of the fruiting specimen is missing. It is therefore possible that flowers were once present on this specimen but were dissected by Sonder, the fragments of which were placed in a packet (subsequently mounted on MEL 2235239 together with a label bearing diagnostic information in Sonder’s hand). However, there remains an element of uncertainty with this interpretation and as such the three specimens are best regarded as syntypes. Of the syntypes listed above for S. marginatum, Sonder examined and annotated the Preiss material at MEL and LD as well as the Drummond material at BM. The MEL material (MEL 2258906 and MEL 2258907), which is from Sonder’s personal herbarium, is designated as an appropriate lectotype: the former sheet includes a single individual with an incomplete inflorescence, descriptive information in Sonder’s hand, and dissected flowers in a packet, and the latter sheet is a complete individual (albeit with a broken scape). I have seen several sheets of the type gathering of S. pruinosum but only three of these have been annotated by Sonder: MEL 2258904, MEL 2258905 and LD 1731539. MEL 2258905, which includes descriptive annotations by Sonder and dissected flowers fragments in a packet, is herein designated as the lectotype. MEL 2069493 is from Sonder’s personal herbarium and comprises a single, incomplete individual of S. diversifolium and two labels, one for Preiss 2232 (the type of S. marginatum) and the other for Preiss 2236 (the type of S. pruinosum). I am uncertain as to which gathering this specimen came from. Notes. Stylidium diversifolium exhibits variation in overall robustness, leaf size and shape, hyaline leaf margin morphology (whether erose, fimbriate or entire), the number of whorls of sterile scape bracts, the presence of labellum lateral appendages and glandular trichome density, which accounts for Sonder’s (1845) somewhat enthusiastic taxonomy. Stylidium eriorhizum R.Br., Prodr. FI. Nov. Holland. 569 (1810). Candollea eriorhiza (R.Br.) F.Muell., Syst. Census Austral. PI.. 86 (1882). Type citation: ‘(T.) v. v.’ Type specimen'. Shoal water Bay Thirsty Sound outer and inner entrance [Queensland], 3 September 1802, R. Brown s.n. [Bennett No. 2586] {lectotype, here designated: BM 000645731!; isolectotypes'. BM000645732!, E 00279213!, E 00279214!, FI 006824, K 000060209!, K 000060212!, MEL 2257317!, NSW 923303 image!, P 00712394 image!, P 00712395 image!). Typification. Bean’s (1999a) citation ‘holo: BM’ is not treated herein as an inadvertent lectotypification since he did not indicate which of the two duplicates at BM was the ‘holotype’ (neither duplicate has been annotated by Bean). 238 Nuytsia Vol. 28 (2017) Stylidium fa lea turn R.Br., Prodr. FI. Nov. Holland. 572 (1810). Candollea falcata (R.Br.) F.Muell., Syst. Census Austral. PI. . 86 (1882). Type citation. ‘(M.) v. v.’ Type specimen. King George’s Sound [Western Australia, December 1801-January 1802], R. Brown s.n. [Bennett No. 2610] (holotype: BM 000812589!). Stylidium lessonii DC., Prodr. 7(2): 337 (1839). Type, ‘in Nova Hollandia ad portum Regis Georgii legit cl. Lesson. ... (v.s. comm, a cl. Merat.)’ (holotype: G-DC G00458499!; isotype: P 00313107!). Stylidiumfalcatum f. robustaWawra, Itin. Princ. S. Coburgi 1:131(1883). Type. King George’s Sound [Western Australia], 1872-1873, H. Wawra Coll. I 854 [cited as 845 in protologue] (iholotype: W!). Stylidium fasciculatum R.Br., Prodr. FI. Nov. Holland. 572 (1810). Candollea fasciculata (R.Br.) F.Muell., Syst. Census Austral. PI.: 86 (1882), nom. illeg. non R.Br. ex DC .(1817 = Hibbertia depressa Steud., Dilleniaceae). Type citation. ‘(M.) v. v.’ Type specimen. Bald Head, King George’s Sound [Western Australia], December 1801, R. Brown s.n. [Bennett No. 2609] (lectotype, here designated: BM 000812593!; isolectotypes: BM 000812582!, BM 000812592!, CANB 279060!, E 00279240!, E 00279263!, E 00279264!, K 00060329!, MEL 2254211!). Paralectotype \syntype\. King George’s Sound [Western Australia], December 1801, R. Brown s.n. [Bennett No. 2609] (BM 000812591), = S. adnatum. Notes. Brown describes the capsules of S. fasciculatum as having seeds in both loculi. While this is evident in the bulk of the material cited above, only one fertile locule is evident in the three fragments which comprise BM 000812591 (mounted on the lower right hand side of the sheet that also bears BM 000812593 and BM 000812582). This anomalous material is labelled by Brown as ‘Stylidium fasciculatum (3’ and is referable to S. adnatum. Stylidium fasciculatum is a morphologically variable species that is the subject of ongoing taxonomic investigations that aim, in part, to resolve the status of S. fasciculatum subsp. gigantic (J.A. Wege JAW 1174) (Western Australian Herbarium 1998-) and S. fasciculatum var. elongatum Benth., the latter of which is a name of uncertain application (Council of Heads of Australasian Herbaria 2011b). Stylidium floribundum R.Br., Prodr. FI. Nov. Holland. 569 (1810). Candollea floribunda (R.Br.) F.Muell., Syst. Census Austral. PL . 86 (1882). Type citation. ‘(T.) v. v.’ Type specimen North Coast Carpentaria Island h [North Island, Northern Territory], 20 December 1802, R. Brown s.n. [Bennett No. 2588] (/lectotype, here designated: BM 000563896!; isolectotypes'. BM 000563900!, E 00279265!, E 00279266!, E 00279267!, K 000741770!, K 000741771!, MEL 1061579!, P 00712396!). Typifcation. Bean’s (1999b) citation ‘holo: BM (2 sheets)’ is not considered herein as an inadvertent lectotypification since the two sheets are duplicates and not a single specimen (BM 000563896 is the specimen from Brown’s own herbarium while BM 000563900 is the Dryander duplicate). Stylidium glandulosum Salisb., Parad. Lond. 2(1), t. 77 (1807). Stylidium fruticosum R.Br., Prodr. FI. Nov. Holland. 570 (1810), nom. illeg., nom. superfl. Candolleaglandulosa (Salisb.) F.Muell., Syst. Census Austral. PI.: 86 (1882). Type citation ‘Sponte nascentem in New Holland, oris occidentalibus, legit RtusBROWN. ... For this species, Mr. Hooker is indebted to E.J.A. WOODFORD, Esq. I saw it in flower at Kew, in July, 1804... ’ (lholotype\ K 000355286!). Notes. Peter Good collected seeds of S. glandulosum in January 1802 whilst Investigator was anchored