© 2016 Board of the Botanic Gardens & State Herbarium (South Australia) Journal of the Adelaide Botanic Gardens 29 (2016) 147–220 © 2016 Department of Environment, Water & Natural Resources, Govt of South Australia Notes concerning the classification of species included in Calocephalus R.Br. s.lat. and Gnephosis Cass. s.lat. (Asteraceae: Gnaphalieae), with descriptions of new genera and species P.S. Short 5 Chemnitz Court, Driver, Palmerston, Northern Territory 0830, Australia Previously: National Herbarium of Victoria (MEL) & Northern Territory Herbarium (DNA) Abstract Descriptions and keys to 40 species here included in, or which have often been considered to have affinities with the broadly-delimited genera Calocephalus R.Br. and Gnephosis Cass., are provided. Five new species are referred to Calocephalus, i.e. C. badmanii P.S.Short, C. beardii P.S.Short, C. birchii P.S.Short, C. glabratus P.S.Short, C. pilbarensis P.S.Short, and one to Gnephosis, i.e. G. newbeyi P.S.Short. The new genus Balladonia P.S.Short is described and the new combinations made for B. aervoides (F.Muell.) P.S.Short and B. multiceps (J.H.Willis) P.S.Short, species previously placed in Calocephalus and Chthonocephalus Steetz, respectively. The genus Notisia P.S.Short is described, the only species, N. intonsa (S.Moore) P.S.Short being transferred from Gnephosis. Lectotype specimens are designated for the names of 14 species and two varieties. Keywords: Asteraceae, Gnaphalieae, Balladonia, Blennospora, Calocephalus, Gilruthia, Gnephosis, Leucophyta, Notisia, review, taxonomy, lectotypifications, pollen:ovule ratios, Australia. Introduction unmanageable genus. Those here adopted are the best that suggest themselves [...] although it must be admitted that in George Bentham (1867a), in his treatment of the gna- some instances they are not altogether satisfactory. phalioid genera of Australian Compositae (essentially Bentham 1867a, p. 612, regarding the his tribes “Gnaphalieae” and “Eugnaphalieae”) in Flora limits of Helichrysum in Australia Australiensis, recognised 34 genera. Excluding several It is very near to Gnephosis, differing chiefly in the pappus. orthographic variants and nomenclatural synonyms he The general receptacle is also more broken up or slightly listed 77 generic synonyms. The previous year, in a branched, the partial heads are distinct and having more letter dated 22 July 1866 and sent to Ferdinand Mueller, florets connect the genus through Cephalipterum and Gna­ Bentham had indicated his progress with his treatment phalodes with Helipterum. Bentham 1867a, p. 573, regarding Calocephalus of the family for the Flora and noted that So far as I have gone, I have been able (pretty well to my It is closely allied to Angianthus, but the general involucre own satisfaction, although very likely not to that of others) and receptacle are less developed than in that genus and the very much to reduce the small genera. partial involucres much more so, consisting of much more Cited in Daly 1927, p. 158 numerous bracts and not flattened. Bentham 1867a, p. 569, discussing Gnephosis Mueller was to subsequently accept many of the However, non-specialist taxonomists found it con- genera recognised by Bentham. However, he noted, venient to simply follow Bentham’s concepts and prag- for example, that of the many names — often coined matically placed new species in these genera without, by Nicolai Turczaninow (1851) and Asa Gray (1851a, surely, giving much thought to their relationships. The b, 1852) — reduced to synonymy under Angianthus work of earlier taxonomists such as Gray and Turczani- Wendl., Gnephosis and Myriocephalus Benth by Ben- now were presumably overlooked or simply ignored, tham, “some were readily to be restituted” (Mueller with Bentham’s opinions holding sway for more than a 1882, pp. 82, 83). That Mueller found Bentham’s century. work unacceptable is un surprising given the following As part of a study of the genus Angianthus (sensu statements concerning generic limits in Flora Austra­ Bentham) in the late 1970s, I familiarised myself with liensis: many other compound-headed species of Australian The limits to be assigned to the group are very uncertain, as it is connected with so many others by almost insensible Gnaphalieae and, to a lesser extent, Australian species gradations [...] We are obliged [...] to make arbitrary which at that time were still, following Bentham demarcations, in order not to unite the whole tribe into one (1867a), included in Helipterum DC. and Helichrysum Published online: 15 Dec. 2016 • flora.sa.gov.au/jabg 147 ISSN 0313-4083 (Print) • ISSN 2201-9855 (Online) P.S. Short J. Adelaide Bot. Gard. 29 (2016) Mill. As with Mueller, I came to the conclusion that skirrophorum) out of the 40 species treated here, being Bentham’s generic concepts left much to be desired. included in their study. In neither case were species I have noted this to be the case for Calocephalus and of what I treat as Gnephosis s.str. and Calocephalus Gnephosis in several publications (e.g. Short 1981a, s.str. included in the analyses. On the other hand, the b, c, 1986a), reinstating Blennospora A.Gray (Short results support my earlier beliefs (e.g. Short 1986a) 1987a) and subsequently Trichanthodium Sond. & that a number of genera should be segregated from F.Muell. ex Sond. (Short 1990a) and began (Short both Calocephalus s.lat. and Gnephosis s.lat. Thus, 1986b) to again use the name Leucophyta brownii Cass. of the four species of Calocephalus s.lat. included in for the species known for many years as Calocephalus analyses, it was no surprise to see Leucophyta and C. brownii (Cass.) F.Muell. Anderberg (1991), in his knappii in different clades while Blennospora drum­ world-wide review of the taxonomy and phylogeny of mondii and C. multiflora are in yet another clade. In the tribe Gnaphalieae, accepted many of the smaller regard to the four species of Gnephosis s.lat. used in genera which I had reinstated from Angianthus s.lat. the analyses, placement of the morphologically very as well as recognising Blennospora and Leucophyta. different Gnephosis arachnoidea and G. intonsa in He also recorded Calocephalus sensu Bentham as different clades is similarly unsurprising. However, the being “probably polyphyletic” and recognised that placement of Trichanthodium exilis and T. skirrophorum Calocephalus angianthoides is referable to Gnephosis in clades distantly removed from each other was totally s.str. (as defined by Short 1987b) but suggested that it unexpected. Ward et al. (2009) presented similar results would probably be paraphyletic without inclusion of in regard to Australasian Gnaphalieae members, using monotypic Hyalochlamys A.Gray. Anderberg (1991) many of the same species as Bayer et al., but in their also tentatively referred those species excluded from results both species of Trichanthodium are in the one Gnephosis s.str. to the genus Leptotriche Turcz. He clade. made no combinations to formally accommodate them A comparison of the generic concepts of Gray in that genus and, with the exclusion of those I have (1851a, b, 1852) and Turczaninow (1851), who often removed to Trichanthodium (Short 1990a) and the independently based their descriptions on duplicates tentative addition here of G. setifera P.S.Short (Short collections gathered by James Drummond, with the 1990c), they constitute what I treat here as species of treatment of the same taxa by Bentham (1867a) and Gnephosis s.lat. more recent workers, principally myself and Paul Following his morphology-based, cladistic analysis Wilson, is presented in Table 1. In view of his comments of the tribe Gnaphalieae, Anderberg (1991), placed the that many earlier names “were readily to be restituted” majority of Australian genera in the endemic subtribe one feels that Mueller (1882) would mostly approve Angianthinae, with most others in the subtribe Cassi- of a reversal to the recognition of at least some of the niinae. The subjects of this paper, Calocephalus s.lat. smaller genera. However, far more work pertaining and Gnephosis s.lat. were included in the Angianthinae. to the circumscription and status of most Australian Subsequent to Anderberg (1991), Bayer et al. (2002) gnaphalioid genera, particularly using molecular used a molecular approach to examine phylogenetic sequencing combined with morphological studies relationships in the Australian Gnaphalieae using non- (e.g. as in Weber & Schmidt-Lebuhn 2015), needs to coding chloroplast DNA sequences (the trnL intron and be carried out before there can be consensus on these trnL/trnF intergenic spacer), the maturase encoding matters. It is for this reason that, despite having been plastid enzyme matK, and the external transcribed critical of past workers in simply adopting Bentham’s spacer (ETS) of nulcear DNA. For their analyses, 69 generic concepts, I too largely adopt here what I of the 87 genera then recognised were sampled, but consider to be broad, non-monophyletic concepts of this equated to just 77 species out of a total of 475– Calocephalus and Gnephosis. However, I do erect 500 species. The results indicated that the subtribes two new genera, Balladonia and Notisia, and maintain Angianthinae and Cassiniinae as circumscribed by several which I have previously reinstated. Anderberg (1991) are not monophyletic. Furthermore, I have been aware of some of the new species for results generally support the recognition of many of the 30 years or more and, although they may have phrase- smaller genera which have been described or reinstated names, I believe it important that they be formally in recent years in papers by, for example, myself (e.g. named and their various features described. Short 1983, 1986c, 1987a, 1989b, 1990a) and Wilson I here provide descriptions of 40 species. Most (e.g. 1989a, b, 1992a, b, c). Bayer et al. (2002) also are detailed, being of species which are either first- indicated that some genera need to be more closely in- described here or of species which I have not previously vestigated.