CSIRO PUBLISHING Australian Systematic Botany, 2021, 34, 1–205 https://doi.org/10.1071/SB19049 A taxonomic revision of the Australasian genera Dracophyllum and Richea (Richeeae: Styphelioideae: Ericaceae) Stephanus Venter Australian Tropical Herbarium, James Cook University, PO Box 6811, Cairns, Qld 4870, Australia. Email: fanie.venter@viewnaturalhistory.com Abstract. The genus Dracophyllum Labill. is revised, with a total of 61 species being recognised in four subgenera and two species (Dracophyllum minimum F.Muell. and D. strictum Hook.f.) are listed as incertae sedis. The genus Richea R.Br. is reduced to synonymy under Dracophyllum where it is divided into two new subgenera, namely, Dracophyllum subgenus Cystanthe (R.Br.) S.Venter and D. subgenus Richea (R.Br.) S.Venter. Replacement names published here are Dracophyllum laciniatum S.Venter, D. persistentifolium S.Venter and D. tasmanicum S.Venter, and new combinations published here are Dracophyllum alpinum (Menadue) S.Venter, D. continentis (B.L.Burtt) S.Venter, D. desgrazii (Hombr. ex Decne.) S.Venter, D. gunnii (Hook.f.) S.Venter, D. pandanifolia (Hook.f.) S.Venter, D. procerum (F.Muell.) S.Venter, D. sprengelioides (R.Br.) S.Venter and D. victorianum (Menadue) S.Venter. Nomenclature, descriptions, illustrations, photographs and distribution maps are provided for each species and lectotypes are designated where necessary. A key to the subgenera and keys to species within these are provided. Keywords: Australasia, Dracophyllum, Richea, epacrid, Ericaceae, Richeeae, taxonomy. Received 18 December 2019, accepted 25 August 2020, published online 4 January 2021 Introduction Dracophyllum Labill. is one of three currently recognised genera (Dracophyllum, Richea R.Br. and Sphenotoma (R.Br.) Sweet) in tribe Richeeae Crayn & Quinn, subfamily Epacridoideae Arn. family Ericaceae Durande (Crayn et al. 1998; Kron et al. 2002). Recent family-level classification has listed 38 genera in Epacridoideae on the basis of phylogenetic analyses (Quinn et al. 2005; Albrecht et al. 2010; PuenteLelièvre et al. 2015). The subfamily is predominantly Australasian (Powell 1983), with some representatives in south-eastern Asia (Leucopogon R.Br.), Hawaii (Styphelia Sm.), Tierra del Fuego and Patagonia in South America (Lebetanthus Endl.). Species of Dracophyllum vary in habit from small cushion plants a mere 5 mm tall to trees in excess of 13 m tall. Dracophyllum is distributed from mainland Australia (e.g. Powell 1992; Brown and Streiber 1999; Streiber et al. 1999), Tasmania (e.g. Rodway 1903; Curtis 1963; de Salas and Baker 2017), Lord Howe Island (Oliver 1917), New Caledonia (Virot 1975; Venter 2004a; S. Venter, unpubl. data), to New Zealand (Allan 1961; Venter 2002, 2004b; S. Venter, unpubl. data), with subgenus Dracophyllum being found across the full geographic range of the genus (Wagstaff et al. 2010). Dracophyllum has traditionally been divided into the following three subgenera: subgenus Dracophyllum (New Zealand 7 spp., Australia 4 spp., Tasmania 1 species, Lord Journal compilation
CSIRO 2021 Howe Island 1 species and New Caledonia 7 spp.), subgenus Cordophyllum W.R.B.Oliver, with a single species in New Caledonia, and subgenus Oreothamnus (F.Muell.) W.R.B. Oliv. (New Zealand 28 spp. and Tasmania 1 species; Oliver 1952). All species are endemic at the national level. Dracophyllum species in Australia are the most phylogenetically diverse (Wagstaff et al. 2010), whereas the greatest morphological diversity in the genus is found in New Zealand and its offshore islands, and also in New Caledonia (Oliver 1928, 1952; S. Venter, unpubl. data). The New Caledonian and New Zealand species have recently radiated following at least two instances of long-distance dispersal from eastern Australia (Wagstaff et al. 2010; S. Venter, unpubl. data). The other two genera in the Richeeae are Richea, which is endemic to south-eastern Australia (10 species restricted to Tasmania and two to mainland south–eastern Australia; Australian Plant Census, see https://biodiversity.org.au/nsl/ services/apc), and Sphenotoma (seven recognised species; Australian Plant Census, see https://biodiversity.org.au/nsl/ services/apc), which is restricted to south-western Western Australia. Richea appears somewhat similar to Dracophyllum; however, its ovoid or conical corolla with united lobes that form an operculum easily distinguishes it. The operculum splits transversely and falls off as an entire unit, leaving a persistent basal ring-like structure (Menadue and Crowden 2000). Sphenotoma lacks a leaf sheath and has a narrow corolla www.publish.csiro.au/journals/asb 2 Australian Systematic Botany S. Venter tube with the throat almost closed by the presence of prominent longitudinal ridges that extend from the base of the corolla lobes down into the corolla tube (Bentham 1869). Taxonomic history of Dracophyllum and Richea The descriptive history of the genus Dracophyllum began with two plant specimens collected in March 1773 by J. R. Forster and his son J. G. A. Forster at Dusky Bay in Fiordland, New Zealand (Table 1). These specimens were described as two different species, one by father and son (Forster and Forster 1776) as Epacris longifolia J.R.Forst. & G.Forst. (Dracophyllum longifolium (J.R.Forst & G.Forst.) R.Br. ex Roem. & Schult.) and the other by J. G. A. Forster (1786), as Epacris rosmarinifolia G.Forst. (Dracophyllum rosmarinifolium (G.Forst.) R.Br. ex Roem. & Schult.). The genus name Epacris Cav. [1797], which is now attached to a genus of 46 species in Australia and New Zealand (Quinn et al. 2015), is conserved against Epacris J.R.Forst. & G.Forst. [1776] and contains neither of the above species published by the Forsters. Labillardière (1800) established the genus Dracophyllum for a plant that he collected in New Caledonia during April 1793. He named this plant D. verticillatum as he observed the flowers to be arranged in rings or verticels on the inflorescence. Ten years later, Brown (1810) divided Dracophyllum into sections Dracophyllum and Sphenotoma on the basis of corolla shape, corolla-lobe apex shape, position of the stamens, inflorescence shape and the inflorescence bracts being persistent or deciduous. In the same publication, under the genus Dracophyllum, he mentioned (in a note) that the two Epacris species of the Forsters must now be included in Dracophyllum. According to Article 35.2 of the International Code of Nomenclature for algae, fungi, and plants (Turland et al. 2018), Roemer and Schultes (1819) were the first to validly publish these two combinations. Poiret (1811) extended the descriptions of the Forsters’ two species, maintaining them as E. longifolium and E. rosmarinifolium in his Epacris section Dracophyllum (Labill.) Poir., a grouping that was based on the number of bracts, shape of the corolla tube and position of the stamens. Sprengel (1825) also did not accept Dracophyllum as a separate genus and retained D. longifolium, D. rosmarinifolium and species of Cosmelia R.Br. in the genus Epacris, transferring D. secundum R.Br. to the genus Prionotes R.Br. Richard (1832) was the first to revise Dracophyllum, which he based on the New Zealand collections in the Muséum Table 1. Taxonomic history of the genus Dracophyllum Labill. sens. lat. Author Publication date Publication Forster, J.R. & G. Forster, G. Labillardière, J.J.H. Brown, R. Poiret, J.L.M. Roemer, J.J. & Schultes, J.A. Sprengel, C. Sweet, R. Richard, M.A. De Candolle, A.P. Cunningham, A. Hooker, J.D. Raoul, E. Hooker, J.D. Hooker, J.D. Brongniart, A.D. & Gris, A. Hooker, J.D. Bentham & Mueller Mueller, F. von Colenso, W. Colenso, W. Colenso, W. Cheeseman, T.F. Cheeseman, T.F. Oliver, W.R.B. Oliver, W.R.B. Virot, R. Brown, E.A. & Streiber, N. Venter, S. Venter, S. Venter, S. 1776 1786 1800 1810 1811 1819 1825 1830 1832 1839 1838 1844 1846 1853 1860 1864 1867 1869 1870 1887 1888 1890 1906 1925 1929 1952 1975 2000 2002 2004 2004 Characteres generum plantarum Florulae Insularum Australium Prodromus Rélation du voyage à la recherche de la Pérouse, vol. 2. Prodromus Florae Novae Hollandiae, vol. 1. Encyclopédie Méthodique, Botanique, Supplement, vol. 2. Systema Vegetabilium, vol. 4. Systema Vegetabilium, vol. 1. Hortus Britannicus Essai d’une flore de la Nouvelle Zélande Prodromus Systematis Naturalis Regni Vegetabilis, vol. 7. Annals of Natural History, vol. 2 Flora Antarctica, vol. 1. Choix de plantes de la Nouvelle-Zélande Flora Novae-Zelandiae, vol. 2. Flora Tasmaniae, vol. 2 Bulletin Société Botanique de France, vol. 2. Handbook of the New Zealand Flora, vol. 1. Flora Australiensis, vol. 4. Fragmenta Phytographiae Australiae, vol. 7. Transactions and Proceedings of the New Zealand Institute, vol. 20. Transactions and Proceedings of the New Zealand Institute, vol. 21. Transactions and Proceedings of the New Zealand Institute, vol. 22. Manual of the New Zealand Flora Manual of the New Zealand Flora, 2nd edn Transactions of the New Zealand Institute, vol. 59. Transactions of the Royal Society of New Zealand, vol. 80. Flore de la Nouvelle Calédonie et dépendances, vol. 6. Telopea, vol. 8(3). New Zealand Journal of Botany, vol. 40(1). New Zealand Journal of Botany, vol. 41(4). New Zealand Journal of Botany, vol. 42(4). Number of species covered 1 5 1 3 1 3 2 2 4 7 5 12 11 14 2 5 11 3 1 1 1 1 18 20 45 48 7 2 2 1 1 Taxonomic revision of Dracophyllum and Richea Nationale d’Histoire Naturelle in Paris (P). The notes in Richard’s publication give the impression that he had a modern concept of the circumscription and characteristics of Dracophyllum. He included four species (D. lessonianum A.Rich., D. longifolium, D. rosmarinifolium and D. urvilleanum A.Rich.; the first and last were described as new) and gave a lengthy discussion as to how the species differed from each other, with a particular reference to floral characters. De Candolle (1839) placed Dracophyllum in his family Epacrideae, recircumscribing it and providing the first full description of the genus. He included only members of Brown’s (1810) section Dracophyllum, considering the species in the second section as belonging to a different genus, namely Sphenotoma. De Candolle listed seven species of Dracophyllum, five from New Zealand, one from Australia (D. secundum) and D. verticillatum from New Caledonia. These were also the first comprehensive descriptions of the species and he was the first author to mention species with a deciduous corolla, a characteristic of some species of Dracophyllum. Endlicher (1836) described the following two sections: the new section Dacryanthus Endl. characterised by free stamens in order to accommodate D. secundum, and a remodelled section Dracophyllum sensu Endl. (as Eudracophyllum), characterised by stamens adnate to the corolla to accommodate all the other known species. However, later authors have not upheld Endlicher’s sections. J. D. Hooker (1844) was the first author to give full descriptions of the species and to discuss species relationships: ‘. . .but none of these authors proposed any sectional characters for these species, which differ most materially from those of Mr. Brown’s first section of the genus, in many of them having epipetalous stamens, as also in the flowers being spiked and the bracteae persistent, and which equally differ from Sphenotoma in habit and the form of the corolla’ (p. 45). In 1853, he published the second revision of Dracophyllum (Hooker 1853) wherein he discussed 14 species, of which nine were described as new. He placed the species in two unnamed sections on the basis of inflorescence shape, inflorescence bracts being deciduous or persistent, and the length of the calyx lobes in relation to tube length. This was published as part of an account of the plants collected on the Antarctic voyage during 1839–1843. Hooker was unsure about the differences between D. urvilleanum, D. lessonianum and D. filifolium Hook.f., thus starting the 72-year period of confusion pertaining to these three species. Mueller (1858) proposed the section Oreothamnus F.Muell. in Dracophyllum, encompassing species with the inflorescence in a raceme, spike or with a solitary flower, and stamens attached to the corolla tube for most of their length. Bentham and Hooker (1876) divided the genus into three unnamed groups on the basis of leaf characters and the structure of the inflorescence. Hooker’s concept of D. urvilleanum differed markedly from that of Richard’s (1832) and he included D. filifolium, D. lessonianum and D. scoparium in synonymy of D. urvilleanum. This publication was also noteworthy for containing the first published key to Dracophyllum species. Australian Systematic Botany 3 Brongniart and Gris (1864) were the first to treat New Caledonian Dracophyllum species. They formally published the name D. ramosum Pancher ex Brongn. & Gris. and mentioned D. cosmelioides, both names written by Pancher onto the specimen sheets. Dracophyllum cosmelioides was merely mentioned in a note as a manuscript name, but was later validly published by Oliver (1952). The Australian Dracophyllum species were fully revised by Mueller (1867); however, the descriptions provide little in the way of discussion. Bentham and Hooker (1876) gave a full generic description of Dracophyllum and divided the genus into two sections, Dracophyllum (as Eudracophyllum) and Sphenotoma, on the basis of floral characters. This is in contrast to Hooker (1844) mentioning that section Sphenotoma had been raised to genus level by Sweet (1827) as Sphenotoma (R.Br.) Sweet. Bentham and Hooker (1876) made no mention of Mueller’s section Oreothamnus. Fifty-three years after Hooker’s second revision, Cheeseman (1906) revised the genus for New Zealand, wherein he recognised 18 species. He classified the species in two unnamed sections that correspond with Oliver’s (1928) later division into subgenera (Eu)dracophyllum and Oreothamnus. Cheeseman brought some order to Dracophyllum taxonomy when he placed many of Colenso’s species in synonymy (Cheeseman 1906). Sadly though, he was unable to clear up the D. filifolium–D. urvilleanum–D. lessonianum confusion that had begun with Bentham and Hooker (1876). Cheeseman’s concepts of D. rosmarinifolium, D. scoparium and D. uniflorum also differed from those of Hooker. The next significant addition to Dracophyllum taxonomy was the establishment of subgenera. Oliver (1928) raised the sections Dracophyllum (as Eudracophyllum) and Oreothamnus to subgenus level, described a third subgenus Cordophyllum to accommodate D. involucratum from New Caledonia, and removed the section Sphenotoma from Dracophyllum. This was also the first major monograph of Dracophyllum and, in it, Oliver accepted 45 species. This publication gave the first comprehensive introduction to the genus, with discussions on the macro-morphology and grouping of the species. He brought order to the old D. filifolium–D. lessonianum–D. urvilleanum confusion, but, in the process, divided D. filifolium into three varieties, none of which is upheld in the present study. Oliver’s grouping of the species is not natural and differs markedly from that presented here, but was, nonetheless, a commendable effort to resolve the taxonomy of Dracophyllum. Oliver (1952) later published a ‘Supplement’ to the revision, wherein he regrouped and cited types for the species, with many lectotypes being chosen by him. In many cases, he did not write the word ‘TYPE’ on the specimen sheets. Subsequent research (Allan 1961; Virot 1975; Brown and Streiber 1999; Venter 2002) has shown serious shortcomings in the nomenclature, and Oliver’s groups have been shown to poorly reflect the phylogeny inferred by Wagstaff et al. (2010). The latest publication covering all Dracophyllum for New Zealand is that of Allan (1961) wherein he stated that he followed the findings of Oliver (1928; 1952) in a very large part, and was unable to compile a satisfactory key with the 4 Australian Systematic Botany S. Venter knowledge available. Allan included 35 species and agreed that the D. rosmarinifolium–D. uniflorum, the D. filifolium– D. urvilleanum and the D. longifolium groups remained unresolved. Allan seems to have been unsure about the status of D. filifolium, D. longifolium, D. recurvum Hook.f. and D. strictum. He also published lengthy comments on Colenso’s and Oliver’s species and varieties. These comments are in small type, a procedure Allan used to report names, but to make no judgment on their distinctness. He provided considerable discussion of hybridism in the genus, saying that it is a common phenomenon and that: ‘. . .field evidence abundantly supports the view that hybrids occur freely between several members of the subgenus Oreothamnus’ (Allan 1961, p. 538). Virot (1975) revised the Dracophyllum species of New Caledonia mostly on the basis of material collected by the surgeons Vieillard and Déplanche, and also MacKee specimens from Missouri Botanical Gardens (MO). Virot accepted seven endemic Dracophyllum species for New Caledonia, six belonging to the subgenus Dracophyllum and D. involucratum to subgenus Cordophyllum. The species descriptions and the material studied are precise, but notes on the distribution, ecology and observations are scanty, with hardly any mention being made of related species or how the species differ from each other. The most recent publication dealing with the taxonomy of Dracophyllum for Australia is by Brown and Streiber (1999), wherein they describe two new species (D. macranthum E.A.Br. & N.Streiber and D. oceanicum E.A.Br. & N.Streiber) on the basis of a morphometric study. The latest publications on Dracophyllum for New Zealand and New Caledonia were by Venter (2002, 2004a, 2004b), wherein three new species (D. marmoricola S.Venter, D. ophioliticum S.Venter and D. elegantissimum S.Venter) were described from the northern part of the South Island of New Zealand and D. mackeeanum S.Venter from New Caledonia. The descriptive history of Richea R.Br. began with a specimen collected in April 1804 by Robert Brown at Mount Wellington near the Derwent River, Tasmania. Brown (1810) described two genera, namely Richea and Cystanthe R.Br., and described the Mount Wellington specimen as Cystanthe sprengelioides R.Br. (1810); however, Mueller (1867) transferred the species into Richea as R. sprengelioides (R.Br.) F.Muell. In the same publication, Brown (1810) described R. dracophylla R.Br., which was also collected by him on Mount Wellington. Lindley (1836) described a third genus, Pilitis Lindl., on the basis of his newly described Pilitis acerosa Lindl. Mueller (1867) sank both these taxa into Richea, but recognised two sections, Dracophylloides and Cystanthe, the latter incorporating Cystanthe and Pilitis. Hooker (1844) described Richea pandanifolia Hook.f. from a specimen collected by Gunn (without locality) and, in 1847, he described R. gunnii Hook.f. and R. scoparia Hook.f., both from collections made by Gunn on Mount Wellington. The first revision of Richea was by Mueller (1858). He treated the species as belonging to the genus Cystanthe and described the new sections C. section Atomanthera F.Muell. (C. sprengelioides), C. section Lobanthera F.Muell. (C. procera F.Muell.) and C. section Pilitis (Lindl.) F. Muell. (C. acerosa, C. milliganii (Hook.f.) F.Muell.) (Table 2). The next revision was that of Hooker in Flora Tasmaniae (Hooker 1860), where he recognised three separate genera of Cystanthe (C. sprengelioides with three forms), Pilitis (P. acerosa, P. milliganii Hook.f.) and Richea (R. pandanifolia, R. dracophylla, R. gunnii and R. scoparia). Mueller (1867) published a revision of the genera Cystanthe and Pilitis, which are now included in Richea. The last and most comprehensive revision of Richea was by Menadue and Crowden (2000), where 11 species were accepted. Richea pandanifolia subsp. ramulosa Menadue and R. alpina Menadue were described for the first time in this revision. Phylogenetic relationships Since Bentham (1869), Dracophyllum has generally been regarded as being closely related to Richea and Sphenotoma. Indeed, these three genera have consistently formed a clade in phylogenetic analyses (Powell et al. 1996; Crayn et al. 1998; Kron et al. 2002; Wagstaff et al. 2010; Johnson et al. 2012). This clade corresponds to the tribe Richeeae of Powell (1983) and Powell et al. (1996), which was formalised in Kron et al. (2002). The molecular study of Wagstaff et al. (2010), by far the most comprehensive that has been published on the group, Table 2. Taxonomic history of the genus Richea R.Br. Author Publication date Publication Brown, R. Lindley, J. Hooker, W.J. Hooker, W.J. Hombron & Jacquinot 1810 1836 1844 1847 1852 Mueller von, F. Hooker J.D. Mueller von, F. Menadue, Y. 1858 1859 1868 2000 Prodromus florae Novae Hollandiae, vol. 1. Introduction to the Natural System Flora Antarctica, vol. 1. London Journal of Botany, vol. 6. Voyage au pole sud et dans l’océanie sur les corvettes l’Astrolobe et la Zélée pendent les anné 1837–1840 Fragmenta Phytographiae Australiae, vol. 6 Flora Tasmaniae, vol. 1. Fragmenta Phytographiae Australiae, vol. 6 Australian Systematic Botany Number of species covered 2 1 1 2 1 3 7 6 11 Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 52 86 A 100 53 B 96 70 56 88 95 C G 91 68 D 100 53 100 100 82 E 99 96 I 95 H F 100 100 99 100 82 100 61 J 96 100 100 100 100 100 98 100 66 100 68 53 100 Oreothamnus). The two subgenera of Richea newly published here (subgenus Cystanthe and subgenus Dracophylloides) were monophyletic, but not sisters. The northern Queensland endemic Dracophyllum sayeri F.Muell. was robustly placed sister to the remaining species of Dracophyllum and Richea, with the exception of a Dracophyllum acerosum Dracophyllum densum Dracophyllum filifolium Dracophyllum kirkii Dracophyllum ophioliticum Dracophyllum patens Dracophyllum rosmarinifolium Dracophyllum trimorphum Dracophyllum arboreum Dracophyllum longifolium Dracophyllum muscoides Dracophyllum pronum Dracophyllum scoparium05.30 Dracophyllum scoparium05.77 Dracophyllum strictum Dracophyllum fiordense Dracophyllum menziesii Dracophyllum latifolium Dracophyllum townsonii Dracophyllum traversii Dracophyllum alticola Dracophyllum balansae Dracophyllum cosmelioides Dracophyllum mackeeanum Dracophyllum ramosum Dracophyllum involucratum Dracophyllum verticillatum Dracophyllum ouaiemense Dracophyllum thiebautii Richea acerosa Richea milliganii Richea procera Richea sprengelioides Dracophyllum oceanicum Dracophyllum secundum Dracophyllum macranthum Dracophyllum filzgeraldii Richea alpina Richea continentis Richea pandanifolia Richea scoparia Richea gunnii Richea victoriana Dracophyllum minimum Dracophyllum sayeri Dracophyllum milliganii Sphenotoma capitata Sphenotoma dracophylloides Sphenotoma drummondii Sphenotoma gracilist Acrotriche divaricata Cyathodes glauca Astroloma humifusum Styphelia viridis Brachyloma daphnoides Melichrus procumbens Leucopogon microphyllus Monotoca scoparia Pentachondra pumila Trochocarpa sp. Cosmelia rubra Sprengelia incamata Andersonia sprengelioides Lysinema ciliatum Rupicola sprengelioides Needhamiella pumilio Oligarrhena micrantha Archeria comberi Prionotes cerinthoides Leucothoe racemosa Vaccinium uliginosum Harrimanella hypnoides Cassiope mertensiana Rhododendron kaempferi Arbutus canariensis Pyrola rotundifolia Enkianthus campanulatus Oreothamnus Dracophyllum Dracophyllum Dracophyllum Cordophyllum Dracophyllum Cystanthe Dracophyllum Tribe Richeeae grouped species of Dracophyllum and Richea in a moderately supported clade (Bremer support 6, bootstrap 76%, jack knife 78%; Fig. 1) on the basis of sequences of the plastid genes rbcL and matK. Neither Dracophyllum nor Richea were monophyletic, nor were two of the three subgenera of Dracophyllum (subgenus Dracophyllum and subgenus 5 Dracophylloides Oreothamnus Dracophyllum Fig. 1. The strict consensus tree from parsimony analysis of the combined rbcL and matK datasets (CI = 0.512, RI = 0.775). Bootstrap values >50% are presented above the branches (Wagstaff et al. 2010). Reproduced with permission of Annals of the Missouri Botanical Garden. 6 Australian Systematic Botany Tasmanian species, D. milliganii Hook., which was placed in a tritomy with Sphenotoma and the Dracophyllum + Richea clade. The relationships of D. milliganii remain unresolved. Johnson et al. (2012) sought to resolve the relationships among Epacridoideae genera using supertree and supermatrix analyses of sequence data from five gene regions. Although their taxon sampling of Dracophyllum and Richea species was much lower (5 and 3 species respectively) than that of Wagstaff et al. (2010), their analyses also resolved a Dracophyllum + Richea clade (including D. milliganii as sister to the remainder, albeit weakly supported) but with neither genus being retrieved as monophyletic. Tribe Richeeae was strongly supported but its relationships to other major epacrid lineages were not unambiguously resolved; in different analyses, either Richeeae or Cosmelieae was sister to Epacrideae + Styphelieae. Historical biogeography and evolution The historical biogeography and evolution of Richeeae were investigated in detail by Wagstaff et al. (2010). Their analyses dated the split between Sphenotoma and Dracophyllum + Richea to the early–mid-Miocene, c. 16 million years ago. The early radiations within the clade took place in Australia, and the occurrence of Dracophyllum in New Caledonia and New Zealand was attributed to independent long-distance dispersal from Australia 5.6 and 6.2 million years ago respectively. The presence of a much older (20–25 million years) New Zealand fossil attributed to Richeeae was explained as representing an earlier, now extinct lineage (Jordan et al. 2010; Mildenhall et al. 2014). A similar explanation has been made for fossil Styphelieae (Jordan et al. 2010) that predates the inferred ages of New Zealand members of tribe Styphelioideae (Puente-Lelièvre et al. 2013). This is not unreasonable because of the submergence of New Caledonia and most of New Zealand in the Oligocene (Mildenhall et al. 2014), which presumably caused widespread extinction of terrestrial biota. Since their arrival in New Caledonia and New Zealand, the extra-Australian lineages underwent significant evolutionary radiations, most of which occurred within the past two million years on the basis of a lineage through time plot (Wagstaff et al. 2010). In New Zealand, in particular, this rapid radiation produced from a single ancestor a great degree of morphological diversity (plants range from cushion plants to trees), which rivals that found among the much older Australian lineages. This morphological diversity is not reflected in the chloroplast genes rbcL and matK, the sequences of which are nearly identical among the New Zealand species (Wagstaff et al. 2010). This radiation was presumably promoted by the geological uplift of the New Zealand Alps and the accompanying climatic changes during the Pliocene, which generated new ecological niches (Heenan and McGlone 2013; Schwery et al. 2015; Wood et al. 2017). However, in Australia, the Miocene was a period of increasing aridity, which caused fragmentation and contraction of the mesic flora and the proliferation of sclerophyll and arid plant communities (Hill 2004). Indeed, a slowing of the diversification rate (due to extinction, reduced S. Venter speciation, or both) during this time is evident in the lineage through time plots (Wagstaff et al. 2010). Materials and methods Herbaria and libraries This study followed the standard methods for taxonomic revisions (Davis and Haywood 1973). Relevant literature was consulted in libraries at Auckland Museum, University of Auckland, and Landcare Research at Lincoln, Lincoln University, Museum of New Zealand, Victoria University of Wellington, Nelson Public Library and Wellington Public Library. Herbarium material was received on loan from the following herbaria: AK, AKU, BM, BRIU, CANU, FI, GZU, HO, K, L, LINC, MEL, MPN, MW, NSW, NZFRI, O, OTA, P, S, TCD, UNSW, UPS, W, WAIK, WELT, WELTU and Z. Terminology used is mainly in accordance with Stearn (1996). Field studies Field studies were conducted on populations of all the New Zealand, New Caledonian and northern Australian species of Dracophyllum in their natural habitat. Many different populations per species were surveyed to ascertain variability within species. Flowering and fruiting material were collected and preserved in specimen jars filled with preservative (FAG) made up as follows: formaldehyde (1 mL), ethanol (10 mL), glycerol (1 mL) and water (8 mL). Glycerol is added to the mixture to keep the material soft and pliable, so that it can be easily handled under the microscope without fear of it breaking up. Herbarium material Vouchers were taken to verify all observations reported, and these are housed in CHR as well as NOU and annotated accordingly. Other specimens used are cited with the necessary reference to the herbaria, which sent them as loan material. Measurements Measurements were taken from live material in the field as well as from dried specimens. Distribution maps Distribution maps for the New Caledonia and New Zealand Dracophyllum species were generated with Distribution Plotter (ver. 2.0.2, Department of Conservation proprietary software, Auckland, New Zealand) and, for Australian species, maps generated from records in the Atlas of Living Australia (ALA, see www.ala.org.au) were used. Taxonomic treatment An amplified genus description for Dracophyllum is given followed by keys to the subgenera and the species in the different subgenera. For each species, listed in alphabetical order, complete synonymy is given. All types examined for this study are followed by a ‘!’. Excluded names are listed in Appendix 1. Taxonomic revision of Dracophyllum and Richea Typification Type material of all accepted species and varieties of Dracophyllum and their synonyms was obtained from AK, BM, FI, GZU, HO, K, L, MEL, NSW, O, P, S, W, WELT and Z. All holotypes were checked with the respective protologues. Where lectotypification was necessary, Articles 7–10 of the International Code of Nomenclature for algae, fungi, and plants (Turland et al. 2018) were followed. Results In this study, Richea is included within the circumscription of Dracophyllum, which has nomenclatural priority. In this case, the two sections of Richea that are indicated as the clades ‘Richea D’ and ‘Richea F’ and nested within Dracophyllum (Fig. 1) become Dracophyllum subgenus Cystanthe and subgenus Dracophylloides. The tribe Richeeae then consists of the genera Dracophyllum and Sphenotoma. With the inclusion of Richea in Dracophyllum, eight new combinations are required. New names must be given to three species (Richea acerosa, R. milliganii and R. scoparia), because these specific epithets are already preoccupied in Dracophyllum. Richea is morphologically similar to Dracophyllum. Both share prominent annular scars from fallen leaves, leaves spirally arranged, leaves with parallel venation, leaf bases sheathing, persistent sepals, anthers lacking appendages, five-locular ovary with numerous ovules per loculus, axile placentation, style inserted in a depression in the ovary and the fruit a loculicidal capsule. The major difference between Richea and Dracophyllum is that, in Richea, the stamens are enclosed in the corolla with its lobes fused until the anthers mature and they push the corolla off to be shed as an entire unit (an operculum). An operculum is also present in Dracophyllum (D. elegantissimum, D. latifolium A.Cunn. and D. traversii Hook.f.), but here the stamens are epipetalous and shed with the corolla after pollination. One molecular phylogenetic study provided some evidence that the traditional subgenera are not monophyletic (Wagstaff et al. 2010). However, that study was based on plastid sequence data only and did not resolve the placement of many of the species with strong support. For the purposes of this paper, the subgeneric relationships are not considered adequately tested in a molecular phylogenetic framework and, therefore, the proposed subgenera in Dracophyllum reflect morphological affinity. The classification of tribe Dracophylleeae (=Richeeae) is, therefore, as follows: Sphenotoma (11+ species), Dracophyllum subgenus Dracophyllum (21 species), D. subgenus Oreothamnus (29 species), D. subgenus Cystanthe (4 species) and D. subgenus Richea (7 species). Australian Systematic Botany 7 D. scoparium to form abundant fertile hybrids. These are polymorphic, connecting the parent species by a wide spectrum of transitional forms and have been discussed in detail by Cockayne (1904), Du Rietz (1930) and Wardle (1987). On the Chatham Islands, Dracophyllum longifolium commonly hybridises with D. scoparium (D.
insulare W.R.B.Oliv.) and with D. rosmarinifolium (D.
acicularifolium (Cheeseman) W.R.B.Oliv.) in Fiordland (Oliver 1928). Dracophyllum filifolium hybridises with D. recurvum (D.
arcuatum W.R.B.Oliv.) and with D. subulatum Hook.f. (D.
vulcanicum W.R.B.Oliv.) on the Volcanic Plateau and also freely with D. rosmarinifolium on Mount Arthur in the Kahurangi National Park. Dracophyllum kirkii Berggr. hybridises with D. pronum W.R.B.Oliv. (D.
saxicola W.R.B.Oliv.) at Arthur’s Pass and D. lessonianum sometimes hybridises with D. subulatum (D.
marginatum W.R.B.Oliv.) and with D. sinclairii Cheeseman (D.
densiflorum W.R.B.Oliv.) on the Volcanic Plateau. (Oliver 1928). Du Rietz (1930) described hybrids between D. oliveri Du Rietz and D. prostratum Kirk from west of Lake Wakatipu, but no specimen could be found to verify this and searches in the named area proved fruitless in locating hybrid plants. On the lower slopes of Mount Ruapehu, D. recurvum commonly hybridises with D. filifolium (D.
arcuatum of Oliver 1928) and Colenso (1896) described this hybrid as D.
varium Colenso. Oliver (1928) regarded this as a putative hybrid between D. longifolium and D. recurvum, but only D. filifolium occurs naturally with D. recurvum, D. longifolium being absent from the North Island. On Campbell Island, D. scoparium commonly hybridises with D. cockayneanum (D.
insulare; Oliver 1928). Hybrids are known to occur between D. persistentifolium S.Venter and D. pandanifolium (Hook.f.) S.Venter (Richea
curtisiae) in disturbed areas at Mount Field National Park, Tasmania, that result from fire or clearance (Menadue and Crowden 2000) Fire has dramatically affected New Zealand’s landscapes and ecosystems in the post-settlement era and, in many places, permanently altered the composition of the vegetation (Perry et al. 2014). Many Dracophyllum species are highly flammable; hence, the common name ‘turpentine scrub’. Dracophyllum has the potential to rapidly colonise disturbed sites. According to Wardle (1987), vegetation modified by sheep grazing and burning are factors that encourage the development of hybrid populations. Taxonomic treatment Dracophyllum Labill., Voy. Rech. Pérouse 2: 210–211 (1800) Hybrids No hybrids were recorded for D. subgenus Dracophyllum. A single hybrid is known in D. subgenus Richea and has been described as
Richea curtisiae A.M.Gray (R. scoparia
R. pandanifolia (Gray 1971; Menadue and Crowden 2000). A multitude of hybrids have been recorded and named in Dracophyllum subgenus Oreothamnus. On Campbell Island, D. cockayneanum Du Rietz hybridises freely with Type: D. verticillatum Labill. Richea R.Br., Prodr. 555 (1810). Type: R. dracophylla R.Br. Perennial cushion plants, subshrubs, shrubs or trees up to 14 m tall. Leaves crowded at the ends of branches spirally arranged 8 Australian Systematic Botany along the branches or imbricate; juvenile leaves present in some species; spirally arranged along branches or crowded at ends of branches; lamina sheath tapering to auricled and margin smooth to ciliate; juvenile lamina longer and wider than adult lamina, coriaceous, linear–triangular, glabrous, rugose, scabrid or pubescent, sometimes with a patch of scabrid hairs at the base of the lamina on the adaxial surface, striated in some species; adult leaves with base of lamina sheathing and leaving ringed scars on branches when falling away forming a sheath; with shoulders tapering, rounded, truncate or auricled, smooth to ciliate; lamina coriaceous to rigid and hard, linear, linear–triangular to triangular, 1–1000
0.3–50.0 mm, with parallel veins, glabrous, rugose, scabrid, pubescent to tomentose, sometimes with a tuft of hairs at base on adaxial surface, sometimes striated; lamina margin entire, serrate, serrulate or thickly covered in hairs, sometimes cartilaginous; lamina apex thickened, obtuse, lobed, acicular, triquetrous to acuminate. Inflorescence a solitary flower, terminal or lateral raceme, spike or terminal or lateral panicle, erect to drooping; lax to dense, flowers opening acropetally, rarely basipetally, 5–460 mm long, linear–oblong to pyramidal in outline; inflorescence bracts persistent or caducous, 1 to numerous, shorter than flowers or overtopping, light green to red, subulate to ovate–triangular, 1.5–67.0
0.5–45.0 mm, surfaces glabrous to sericeous, margins entire, serrulate to ciliate, apex obtuse to mucronate. Flowers 1–5000+ per inflorescence, sessile or pedicellate. Flower bracts persistent or caducous, shorter than or overtopping the flower, membranous to rigid and hard, linear to triangular, 2–20
0.4–8.0 mm, surfaces glabrous to sericeous, sometimes striate, sometimes with a tuft of scabrid hairs either at the apex or the base, margins entire or ciliate, apices obtuse to subulate. Bracteoles 2–6, persistent or caducous, shorter or longer than flower, linear to triangular, 0.5–8.0
0.1–1.7 mm, glabrous to pubescent; pedicels straight to curved, 0.2–20.0 mm long, glabrous, tomentose to pubescent. Calyx 5-lobed, persistent, polysepalous; sepals green to red, lanceolate to broadly ovate–triangular, 0.7–17.0
0.6–5.5 mm, shorter to longer than the corolla tube, surfaces glabrous to pubescent, sometimes with the top half pubescent or with scabrid hairs at the base, sometimes striate; margins entire, denticulate to ciliate or the upper half toothed or ciliate; apex obtuse to acuminate. Corolla white or yellowish or light pink to dark red; corolla tube cylindrical to conical, slightly urceolate to broadly campanulate, mouth of tube closed, narrowed to widened, 1.0–22.0
0.8–5.0 mm, exterior surface glabrous to pubescent; corolla lobes 0–5, free or fused to form an operculum that splits transversely, shed as an entire unit leaving a persistent basal ring; operculum may sometimes persist and the stigma project between the lobes; imbricate in bud, spreading to strongly recurved, broadly lanceolate to triangular, white to dark red, shorter to longer than corolla tube, 0.8–5.0
0.8–4.0 mm, apical ridge sometimes present, sometimes with the margin inflexed, apex acute to obtuse, adaxial surface glabrous or papillate to slightly rugose–verrucate, abaxial surface glabrous or pubescent. Stamens 5, hypogynous or adnate to the corolla tube or inserted on the receptacle; filaments 0.1–20.0 mm long; anthers included or exserted, dorsifixed, oblong to rectangular S. Venter in outline, light yellow to purple, 0.3 – 3.0 mm long. Ovary 5-locular with pendulous placentae; cylindrical to oblong, 0.5–4.5
0.5–3.0 mm, apex tapering to truncate, glabrous to pubescent; nectary scales 5, free, rarely fused to form a ring (D. verticillatum), rectangular to round, 0.2–2.0
0.3–2.0 mm, apices acute to obtuse, bifid to variously toothed; style inserted in a depression at the apex of the ovary, 0.5–19.0 mm long, glabrous, papillose distally or pubescent, sometimes lengthening in fruit; stigma included or exserted, obscurely to prominently 5-lobed. Fruit a dry loculicidal capsule, 5-valved, mostly included in persistent calyx, sessile or pedicellate, light brown to purplish-brown, 1–5 mm long and wide, depressed-globose to oblong, apex pointed to truncate, glabrous or pubescent. Seeds are numerous, filiform, ovoid to trigonous, 0.2–1.3 mm long, testa variously reticulate, cells pitted. Distribution The genus comprises 61 species: restricted to the western Pacific, extending from Australia, Lord Howe Island, New Caledonia to New Zealand and its off-shore islands (Appendix 2). Key to the subgenera 1. Corolla not forming an operculum, corolla lobes well developed........2 Operculum present with corolla lobes not to slightly developed..........3 2. Inflorescence a panicle; inflorescence bracts variously coloured and caducous .....................................................subgenus Dracophyllum Inflorescence a solitary flower, spike or raceme; inflorescence bracts persistent......................................................subgenus Oreothamnus 3. Inflorescence of simple clusters; flowers solitary, crowded in terminal heads; bracts persistent...................................... subgenus Cystanthe Inflorescence a panicle; bracts caducous..................... subgenus Richea Dracophyllum subgenus Dracophyllum Gen. Pl. 2(2): 618 (1876). (As Eudracophyllum). Type: Dracophyllum verticillatum Labill., designated by Virot (1975). Key to the species of subgenus Dracophyllum 1. Inflorescences lateral (below the leaves).............................................2 Inflorescences apical ............................................................................4 2. Pedicel glabrous; stamens hypogynous; fruit not enclosed in persistent sepals; lamina 40–50 mm wide..................................... D. fiordense Pedicel pubescent to tomentose; stamens epipetalous; fruit enclosed in persistent sepals; lamina 4–18 mm wide .........................................3 3. Plants erect-stemmed and sparingly branched; flowers arranged in groups of 3 on the lower inflorescence branches; stamens and style included ......................................................................... D. menziesii Plants with branches forming an open candelabrum; flowers arranged in groups of 5–10 on lower inflorescence branches; stamens and style exserted......................................................................... D. townsonii 4. Flower maturation basipetal; style 14 mm or longer... D. macranthum Flower maturation acropetal; style shorter than 5 mm .......................5 5. Branchlets glabrous .............................................................................9 Branchlets pubescent or tomentose .....................................................6 6. Inflorescence axis prominently ribbed; stamens inserted in the lower half of the corolla tube ........................................... D. mackeeanum Inflorescence axis smooth; stamens either inserted in the upper-third of the corolla tube or hypogynous ...................................................7 Taxonomic revision of Dracophyllum and Richea 7. Stamens epipetalous.............................................................................8 Stamens hypogynous .........................................................................10 8. Lamina 2–5 mm wide; sepals longer than the corolla tube; flowers in groups of three on lower inflorescence branches...........D. balansae Lamina 5–8 mm wide; sepals shorter than the corolla tube; flowers in groups of 5–10 on lower inflorescence branches............ D. strictum 9. Lamina margin smooth; leaves below inflorescence imbricating; corolla 9–17 mm long; outer surface of corolla tube densely pubescent .......................................................................... D. alticola Lamina margin serrulate; leaves below inflorescence not imbricating; corolla (2–)4–6 mm long; outer surface of corolla tube glabrous ....................................................................D. cosmelioides 10. Inflorescence axis prominently ribbed...............................................11 Inflorescence axis smooth..................................................................12 11. Pedicel covered in dense overlapping bracts............. D. involucratum Pedicel without bracts...................................................... D. milliganii 12. Flowers arranged in groups of 3 on the lower inflorescence branches ..................................................................... D. ouaiemense Flowers arranged in groups of 5 or more on the lower inflorescence branches ..........................................................................................13 13. Flowers arranged in groups of 5–10..................................................14 Flowers arranged in groups of >10 ...................................................17 14. Flowers borne in clusters (verticils) at close intervals..D. verticillatum Flowers not in clusters (verticils) ......................................................15 15. Stamens hypogynous, >3.7 mm long; inflorescence rachis glabrous ........................................................................D. secundum Stamens epipetalous, 1.75 mm long; inflorescence rachis pubescent to tomentose ...................................................................................16 16. Stamens exserted; leaves below inflorescence not different from the other leaves; corolla lobes longer than the corolla tube; corolla campanulate .................................................................. D. latifolium Stamens included; leaves below inflorescence different and smaller than the sterile leaves; corolla lobes shorter than the corolla tube; corolla tube narrowed at mouth .................................... D. ramosum 17. Flowers cylindrical; abaxial surface of inflorescence bract scabrid; apex of corolla lobe subacute...................................... D. fitzgeraldii Flowers narrowly to broadly campanulate; abaxial surface of inflorescence bract glabrous; apex of corolla tube obtuse .............18 18. Lamina margin entire; bracteoles equalling flower in length; nectary scales connate at the base....................................................D. sayeri Lamina margin serrulate; bracteoles shorter than flower; nectary scales separate ...........................................................................................19 19. Corolla tube 4–7 mm long; inflorescence bract 10–65 mm long; pedicel glabrous.......................................................... D. oceanicum Corolla tube shorter than 3.5 mm; inflorescence bract 129–615 mm long; pedicel pubescent to tomentose ............................................20 20. Corolla lobe shorter than corolla tube; margin of inflorescence bract entire; sepal shorter than corolla tube; apex of ovary tapering ................................................................D. elegantissimum Corolla lobe longer than corolla tube; margin of inflorescence bract ciliate; sepal equalling corolla tube; apex of ovary round............... D. traversii Dracophyllum alticola Däniker, Vierteljahrsschr. Naturf. Ges. Zürich. 78 (19): 339 (1933) – as ‘alticolum’ Type: New Caledonia. Mount Humboldt. 5 Nov. 1924. A.U. Däniker D509 (lecto: Z!), designated by Oliver (1952). Mount Humboldt. A.U. Däniker 509a (syn: Z!), designated by Virot (1975). Australian Systematic Botany 9 Multi-stemmed shrub, 0.2–1.0 m tall. Branches: bark on old branches grey, smooth, on young stems pubescent and yellowish-brown. Leaves crowded at tips of branches, spreading, light green to grey, sometimes reddish at the base, decreasing in size below inflorescence; lamina sheath 10–15
10–11 mm, coriaceous, striate, tapering, and margin smooth; lamina triangular, 30–100
10–30 mm, slightly concave, surfaces glabrous, prominently striated; margin entire; apex thickened, lobed. Inflorescence a panicle subtended by 5–9 short, imbricating leaves; overtopping the leaves, erect, lax, 80–250 mm long, sparsely branched; rachis and pedicels densely white pubescent to tomentose; inflorescence axis 3–5 mm in diameter; basal inflorescence branch 1–5 mm long, suberect; inflorescence bracts overtopping flowers, subcoriaceous, light green with the tips pink, ovate–triangular, 12–30
9–10 mm, surfaces glabrous, striate, margins entire. Flowers 45–50, in groups of three at the base of the inflorescence, pedicellate; bracteoles caducous, recaulescent, with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, 3–5
0.2–0.3 mm, pubescent, straight and reddish-brown; pedicels reddish-brown, (2–)5–20 mm long, pubescent. Sepals lanceolate to ovate–lanceolate, 9–17
2.5–3.5 mm, longer than the corolla tube, striate, adaxial surfaces glabrous, abaxial surfaces pubescent; margins toothed. Corolla light to dark pink; corolla tube cylindrical, narrowed at mouth, (7–)9–17
2.0–3.5 mm, exterior pubescent; corolla lobes spreading, ovate–triangular, shorter than corolla tube, (3–)4–5
(1.5–) 2.0–4.0 mm, apices subacute; adaxial surface papillate, abaxial surface pubescent. Stamens hypogynous, filaments 5.0–6.5 mm long; anthers included, oblong, purple, and 1.0 – 1.3 mm long. Ovary ovoid, (1.5–)2–4 mm
(1.5–)2–3 mm, densely pubescent, apex truncate; nectary scales rectangular, 0.5–0.8
0.5–1.0 mm, apex obtuse to irregularly toothed; style included, 3–4 mm long, pubescent; stigma capitate. Fruit 3–4 mm long and wide, broadly obovoid, apex round, variously pubescent. Seeds ovoid, 0.5–0.6 mm long, testa slightly reticulate (Fig. 2, 3). Distribution and ecology A New Caledonia endemic restricted to the upper slopes of Mount Humboldt and Massif de Kouakoue (Fig. 4). Dracophyllum alticola appears to be uncommon and restricted in distribution. It occurs on gentle (5–20) southern and southeastern mountain slopes at 1000–1634-m elevation in open maquis vegetation (Jaffré 1991). It sometimes grows in the forest margin, but mostly in open grassy areas among low shrubs. The soil is brown to reddish-brown loam or clay loam derived from peridotite and serpentinite. Phenology Flowering September–November(–May). Illustrations Etymology W. R. B. Oliver, Trans. Roy. Soc. N.Z. 80 (1): t. 5 (1952); R. Virot, Fl. Nouv. Calédonie et Dépend. 6: t. 18 (1975). The specific epithet describes the habitat, meaning ‘dweller of high places’. 10 Australian Systematic Botany S. Venter G Selected specimens I NEW CALEDONIA. Province Sud: Mount Humboldt, 23 Nov. 1951, Baumann-Bodenheim 15505 (P, Z); ibid., 17 May 2005, Venter 13855 (CHR, NOU); ibid., 17 May 2005, Venter 13856 (NOU); ibid., 13 Oct. 1956, MacKee 5412 (L); ibid., 19 Sep. 1980, Hoff 2645 (NOU, P); ibid., 1 Aug. 1993, Cherry 344 (NSW); ibid., 18 Sep. 1980, McPherson 3098 (NOU); ibid., 26 Apr. 1973, Veillon 2849 (NOU); ibid., 17 May 2005, Venter 13855 (CHR, NOU); Massif de Kouakoue, July 1955, Chevalier 11, (NOU). H A Dracophyllum balansae Virot, Fl. Nouv.-Calédonie & Dépend. 6: 138 (1975) Type: New Caledonia. Port-Bouquet, Nov. 1869. B. Balansa 2194 (holo: P!, iso: P!). J F Illustration E B D C Fig. 2. Dracophyllum alticola. A. Flowering branch (1
). B. Sepal abaxial surface (3
). C. Sepal cross-section (3
). D. Lamina apex (10
). E. Laid-out flower (3
). F. Lower inflorescence branch (1.5
). G. Leaf (3
). H. Nectary scale (10
). I. Ovary (5
). J. Flower (3
). Drawn from Däniker 509. Del. S. Venter. Diagnostic features and discussion Dracophyllum alticola is characterised by its low (20–100 cm tall) shrubby habit, light green–grey triangular and striated leaves with prominently lobed apices, 5–10 imbricating bract-like leaves below the inflorescence, large pink flowers (9–17 mm long) with sepals longer than the corolla tube and their abaxial surfaces densely pubescent, corolla tube pubescent on the outside, pedicels 5–20 mm long and pubescent, the ovary, style and fruit densely pubescent. Dracophyllum alticola shows little variation, the young leaves are pubescent, but they soon lose this and become glabrous. R. Virot, Fl. Nouv.-Calédonie et Dépend. 6: t. 23 (1975). Multi-stemmed shrub, 0.6–1.0 m tall. Branches: bark on old branches grey to dark brown, deeply fissured, on young stems pubescent and yellowish-brown. Leaves erect–spreading; lamina sheath 6.7–7.0(–22)
3–5(–9) mm, subcoriaceous, tapering with the top half ciliate, and margin membranous; lamina coriaceous, linear, (30–)40–80(–180)
2–3(–6) mm, flat, surfaces glabrous, sometimes with a tuft of scabrid hairs at the base of the adaxial surface, slightly striated; margin serrulate with 60–100 teeth per 10 mm; apex rarely obtuse. Inflorescence slightly overtopping the leaves, erect, lax, 20–81 mm long, linear–oblong and sparsely branched; rachis and pedicels mostly tomentose; inflorescence axis smooth, 1.0–1.5 mm in diameter; basal inflorescence branch 0.5–1.0 mm long, widely spreading; inflorescence bracts caducous, overtopping flowers, red, ovate–lanceolate to ovate, 8.0–16.4
3.0–6.8 mm, surfaces glabrous, margins ciliate. Flowers hidden by leaves, 25–50, in groups of three at the base of the inflorescence, pedicellate; bracteoles caducous, recaulescent, with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, 2–3
0.2–0.25 mm, glabrous to pubescent; pedicels 0.5–2.0 (–3.0) mm long, glabrous to pubescent. Sepals rose-coloured to red, lanceolate–triangular, 3.0–5.2
1.0–1.7 mm, equal to or longer than the corolla tube, striate, surfaces glabrous; margins ciliate. Corolla white-tipped pink to red; corolla tube narrowly campanulate, 3–4
0.8–1.0 mm; corolla lobes spreading, ovate–triangular to triangular, shorter than corolla tube, 1.2–1.8
1.0–1.2 mm, apices subacute to obtuse; adaxial surface papillate. Stamens inserted in upper-third of corolla tube, filaments 0.3–0.32 mm long; anthers included, oblong, light yellow and 0.5 mm long. Ovary subglobose, 1.0–1.5
0.5–0.8 mm, glabrous, apex round; nectary scales oblong, 0.5–1.2
0.4–0.5 mm, apex retuse; style included, 0.7–1.5 mm long, glabrous; stigma capitate. Fruit reddish-brown, 1.5–1.7
1.5–2.0 mm, obovoid, apex round and glabrous. Seeds yellowish-brown, filiform, 0.2–0.3 mm long, testa slightly reticulate (Fig. 5, 6). Distribution and ecology New Caledonia endemic, restricted to the southern part of Grande Terre as far north as the To De River, with most known Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A B D C E Fig. 3. Dracophyllum alticola. A. Habitat on Mount Humboldt. B. Plant showing the leaf shape and silvery wax on the lamina surfaces. C. Young inflorescences showing the overlapping inflorescence bracts. D. Young plant showing the characteristic triangular leaf shape. E. Flowering plant at the type locality (Venter 13856). Photographs: S. Venter (A–E). 11 12 Australian Systematic Botany S. Venter 165ºE 167ºE Capitale/capital city 20ºS Mer de Corail Coral Sea Ouvéa Lifou Nouvelle Calédonie New Caledonia Île sL oy au té s/L oy alt yI sla nd s Maré 22ºS 0 100 km Nouméa Île des Pins © Australian National University CartoGIS CAP 00-032 Fig. 4. Known distribution of Dracophyllum alticola in New Caledonia. localities east of Nouméa (Fig. 7). Dracophyllum balansae occurs at elevations of 50–700 m and grows along small streams and rivers, sometimes among rocks in streambeds. The vegetation consists of maquis growing on ferrous soils derived from serpentinite. Most plants of D. balansae grow in full sun, but plants in shaded conditions have slightly larger, thinner, lighter green and more pubescent leaves. Phenology Flowering May–December. Etymology Named after Benedict Balansa (1825–1891), French explorer and botanist from Narbonne, France. Diagnostic features and discussion Dracophyllum balansae is characterised by its shrubby habit (0.6–1.0 m tall) with erect branches; erect narrow and long (40–80
2–3 mm) leaves with the abaxial surfaces finely pubescent; inflorescence included in the foliage, flowers arranged in groups of three at the base of the inflorescence, short peduncles (0.5–1.0 mm), pedicels 0.5–2.0 mm long, calyx red with the white corolla tipped pink, corolla tube hypocrateriform and 4–5 mm long with spreading corolla lobes, stamens included and the capsule 1.5–2.0 mm in diameter. Dracophyllum balansae is similar to D. cosmelioides in the long narrow leaves, the inflorescence being shorter than the leaves and the sepals longer than the corolla tube, but differs in lamina, inflorescence and flower characters (Table 3). The lamina is glabrous, but a patch of scabrid hairs is sometimes present at the base on the adaxial surface. Some variation occurs in the length (20–81 mm) and hairiness of the inflorescence axis, which is mostly glabrous, but can be shortly tomentose in some populations. The bracteole and pedicel can be either glabrous or pubescent. Sepals vary from lanceolate to triangular on a single plant. Selected specimens NEW CALEDONIA. Province Sud: Hill at Ngoye, 1 July 1965, Bernardi 9331 (L); Ouinné, 29 Nov. 1984, Jaffré 2668 (NOU); Rivière Bleu, 1 July 1965, Bernardi 9331 (L); Route to Yaté, 25 Dec. 1967, MacKee 18158 (K, L); north of Val des Pins, 30 Mar. 1951, Guillamin & Baumann-Bodenheim 11722 (Z); Bleue River, 7 Dec. 1966, Schmid 824 (NOU, P); along the Bleue River, 5 Aug. 1951, Baumann-Bodenheim 15053 (Z); ibid., 21 May1997, Taxonomic revision of Dracophyllum and Richea I B Australian Systematic Botany 13 Illustration C D A H E F G Fig. 5. Dracophyllum balansae. A. Flowering branch (1
). B. Nectary scale (10
). C. Ovary (10
). D. Flower (6
). E. Inflorescence-bract abaxial surface (3
). F. Sepal abaxial surface (5
). G. Laid-out flower (5
). H. Leaf (1
). I. Lower inflorescence branch (6
). Drawn from MacKee 18158. Del. S. Venter. Musselman, Delzell & Rich 5305 (BM); Creek Pernod at bridge, 12 May 2005, Venter 13845 (NOU); 4 km from Prony to Mount Kouré, 14 May 2005, Venter 13848 (NOU); Creek Pernod, 4 Nov. 2003, Brown 2003/153, Crayn & Quinn (CHR, NOU, NSW). Dracophyllum cosmelioides Pancher ex W.R.B.Oliv., Trans. & Proc. Roy. Soc. NZ. 80 (1): 15 (1952) Type: New Caledonia. Lac Arnaud, 1860. E. Vieillard 828 (lecto: P!), designated by Oliver (1952). Dracophyllum gracile Brongn. & Gris, Ann. Sci. Nat. Bot. 2: 156 (1864). nom. illeg. non R. Br. (1810). R. Virot, Fl. Nouv. Calédonie. & Dépend. 6: t. 24 (1975). The nectary scales are incorrectly illustrated as being fused at the base. Single to multi-stemmed shrub 0.2–1.0 m tall. Branches with the bark on old branches grey to blackish-brown, finely to deeply fissured, young stems tomentose to pubescent and dark brown. Leaves erect; lamina sheaths 4–8
(1.5–)2.5–3.6 mm, membranous, tapering and margin ciliate; lamina linear, rarely linear–triangular, 15–70
0.8–1.5 mm, slightly concave, surfaces glabrous with a tuft of scabrid hairs at the base of the adaxial surface; slightly striated; margin serrulate with 50–70 teeth per 10 mm; apex obtuse, sometimes semi-acute. Inflorescence overtopping the leaves, erect, lax, 10–50 mm long, oblong and sparsely branched; rachis and pedicels tomentose; inflorescence axis 0.5–1.0 mm in diameter; basal inflorescence branch 0.5–1.0 mm long, suberect; inflorescence bracts caducous, overtopping flowers, pink to red, ovate–lanceolate, 4.5–6.0
1.5–2.0 mm, adaxial surface pubescent at the apex, margins ciliate. Flowers hidden by leaves, 5–13, in groups of three at the base of the inflorescence, pedicellate; bracteoles caducous, recaulescent, with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, 2–3
0.2 mm, glabrous; pedicels straight to curved, reddish-brown, 0.5–2.5 mm long, tomentose. Sepals rose-coloured to red, lanceolate–ovate, 2.5–4.0(–8.0)
1.5–2.0 mm, equalling corolla tube, striate, glabrous; margins ciliate. Corolla light pink to red; corolla tube narrowly campanulate, narrowed at the mouth, (2–)4–6
1.0–1.5 mm; corolla lobes spreading, ovate triangular, shorter than corolla tube, 0.8–1.0 mm long and wide, apices obtuse; glabrous. Stamens hypogynous, filaments 2.5–3.0 mm long; anthers included, oblong, light yellow and 0.40–0.45 mm long. Ovary subglobose, 0.5–1.0 mm long and wide, glabrous, apex round; nectary scales rectangular, 0.5–0.7
0.3–0.5 mm, apex retuse; style included, 0.5–0.7 mm long, glabrous; stigma clavate. Fruit pedicellate, reddish-brown, 1.5–3.0 mm long and wide, broadly obovoid, apex round, glabrous. Seeds brown, filiform, 0.20–0.21 mm long, testa slightly reticulate (Fig. 8, 9). Distribution and ecology A New Caledonian endemic restricted to the mountains in the south-east, mostly in the Plain des Lacs area (Fig. 10). Dracophyllum cosmelioides occurs on flat areas or on gentle (0–10) slopes at elevations of 160–200 m. It grows in open areas along streams and rivers that flow through maquis vegetation. Soils are lithosols derived from serpentinite or laterite. In certain populations, plants grow between rocks in streambeds, with their roots permanently wet. Phenology Dracophyllum thiebautii Brongn. & Gris, Ann. Sci. Nat., Bot. 3: 238 (1865). Flowering May–December. Type: New Caledonia: montagnes d’Arama, 1865. C. Thiebaut 339 (holo: P!). Etymology Resembling the genus Cosmelia R.Br. 14 Australian Systematic Botany S. Venter A B C D E F Fig. 6. Dracophyllum balansae. A. Habitat along the Creek Pernod. B. Plant with fruiting branches. C. Fruiting branch showing the spreading leaves. D. Adult plant growing in minimum soil between rocks in creek bed. E. Adult plant (Venter 13845). F. Branch showing the many-flowered inflorescence and the red leaf sheaths. Photographs: S. Venter (A–F). Diagnostic features and discussion Dracophyllum cosmelioides is characterised by the low shrubby habit, linear leaves having blunt apices, inflorescence equal or slightly longer than the leaves, sepals and rachis shortly pubescent, sepals pink to red and striate equalling the corolla tube in length, corolla pink to red and the filaments hypogenous. Virot (1975) placed D. cosmelioides close to D. alticola and D. ouaiemense on the basis of the general morphology of the inflorescence; however, D. cosmelioides is similar to D. balansae in general appearance. For differences between D. balansae and Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 165ºE 15 167ºE Capitale/capital city 20ºS Mer de Corail Coral Sea Ouvéa Lifou Nouvelle Calédonie New Caledonia Île sL oy au té s/L oy alt yI sla nd s Maré 22ºS 0 Nouméa 100 km Île des Pins © Australian National University CartoGIS CAP 00-032 Fig. 7. Known distribution of Dracophyllum balansae in New Caledonia. D. balansae D. cosmelioides (K, L, MEL, NOU, Z); Rivière des Lacs, 14 May 2005, Venter 13849 (NOU); Prony, Oct. 1913, Franc 1825 (L, NBI, Z); l’Odjijoni, 19 Aug. 1958, Hürliman 3346 (Z); 100 m from turnoff to Chute de la Madeleine along Creek Pernod, 12 May 2005, Venter 13844 (NOU). 2–3 8.0–16.4
3.0–6.8 25–50 0.8–1.5 4.5–6.0
1.5–2.0 5–13 Dracophyllum elegantissimum S.Venter, New Zealand J. Bot. 42(1): 37–43 (2004) 3–4
0.8–1.0 Epitepalous 1.0–1.5 4–6
1.0–1.5 Hypogynous 0.5–1.0 Type: New Zealand. Abel Tasman National Park, Rameka Track, 28 Jan. 2001. S. Venter 13827 (holo: CHR!; iso: AK!, K!, NSW!, P!, WELT!). Table 3. Morphological differences between Dracophyllum balansae and D. cosmelioides Character Lamina width (mm) Inflorescence-bract size (mm) Number of flowers per inflorescence Corolla-tube size (mm) Filament habit Ovary length (mm) Illustration D. cosmelioides see comments under D. balansae. Pedicel length is variable, ranging from 0.5 to 2.5 mm on a single plant. Selected specimens NEW CALEDONIA. Province Sud, Goro-nickel, Rivière Kwé, 18 May 2002, Dagostini & Rigault 616 (NOU); River Yaté, 6 Oct. 1924, Däniker 223 (Z). Plaine des Lacs, La Chute River, 25 June 1963, Blanchon 207 (NOU, P); Valley of River des Lacs, 5 Oct. 1950, Guillaumin & Baumann 6542 (Z); near junction of River des Lacs and Ruisseau Pernod, 24 Nov. 1963, Green 1192 S. Venter, New Zealand J. Bot. 42(1): 38, t. 1 (2004). Single-stemmed tree 5–14 m tall. Branches form a closed candelabrum-shaped crown. Bark on old branches light brown, flaky, on young stems yellowish-brown. Leaves at tips of branches in a bromelioid manner, old leaves present; lamina sheath 22–58
13–43 mm, light brown, coriaceous, striate, tapering, margin smooth; lamina coriaceous, light to mid-green, linear to linear–triangular, 330–1000
10–20(–32) mm, surfaces glabrous, prominently striated; margin cartilaginous, serrulate with 15–24 teeth per 10 mm; apex acute and 16 Australian Systematic Botany D S. Venter E F G A H 1–2
1.0–1.5 mm, shorter than corolla tube, striate, surfaces glabrous; margins ciliate; apices subacute to obtuse. Corolla light to dark pink; corolla tube campanulate, widened at mouth, 1–2
1.3–2.0 mm, sometimes white; corolla lobes spreading horizontally to reflexed, ovate–triangular, shorter than corolla tube, 1.2–1.4
1.0–1.3 mm, apices obtuse; surfaces glabrous. Stamens inserted at top of corolla tube, filaments 0.3–0.5 mm long; anthers exserted, rectangular, young anthers pink, deep yellow when mature and 0.9–1.3 mm long. Ovary globose, 1.0–1.5
1.3–1.5 mm, glabrous, apex tapering; nectary scales rectangular, 0.6–1.0
0.5–1.0 mm, apex subacute to irregularly toothed; style exserted, 1.5–1.7 mm long, glabrous, lengthening in fruit; stigma clavate. Fruit not included in persistent calyx, reddish-brown, 1.2–1.5
1.5–1.8 mm, depressed-globose, apex round, glabrous. Seeds yellowishbrown, filiform, 0.7–0.8 mm long, testa slightly reticulate (Fig. 11, 12). Distribution and ecology C I B Endemic to the north-western Nelson area on the South Island of New Zealand, with most localities in the Abel Tasman and Kahurangi National Parks and a few scattered localities southward to Charleston (Fig. 13). Dracophyllum elegantissimum occurs from 101- to 164-m elevation in light to deep shade (rarely in full sun) in forest communities. It is common in the small tree tier of the high-altitude conifer–broad-leaved forest. All the known populations occur on gentle to steep (5–45) south-west to north-west facing mountain slopes. Soils are dark brown humus-rich loam to gritty brown sandy loam derived from granidiorite, calcareous sandstone and conglomerate (Mount Rochfort) or from limestone (Gouland Downs). Phenology Flowering December–February. Etymology Fig. 8. Dracophyllum cosmelioides. A. Habit (1
). B. Laid-out corolla (5
). C. Leaf (1
). D. Flower arrangement on inflorescence (5
). E. Nectary scale (20
). F. Ovary (10
). G. Flower (5
). H. Inflorescence-bract adaxial surface (3
). I. Sepal abaxial surface (5
). Drawn from Vieillard 828. Del. S. Venter. prominently curled. Inflorescence shorter than the leaves, erect, dense, 190–320 mm long, pyramidal and densely branched; rachis and pedicels tomentose, sometimes hirsute to pubescent, light green; inflorescence axis 7.6–13.5(–14.0) mm in diameter; basal inflorescence branch 38–42(–55) mm long, widely spreading (50–80) to right angles with the inflorescence axis; inflorescence bracts caducous, overtopping flowers, light green, ovate–triangular at the base, 270–610
30–45 mm, surfaces glabrous, margins entire. Flowers 600–1000+, in groups of more than 10 at the base of the inflorescence, pedicellate; bracteoles caducous, both bracteoles shorter than the perianth and situated in the middle of the pedicel, 2.0–6.5(–8.0)
0.5–1.0(–1.5) mm, glabrous; pedicels straight, 0.5–2.0 mm long, tomentose. Sepals broadly ovate, The epithet elegantissimum refers to the graceful and slender habit of the canopy and the long slender leaves, which is a distinctive feature of the species. Diagnostic features and discussion Dracophyllum elegantissimum is characterised by the columnar and closed candelabra-shaped crown, bark flaking in large pieces, smooth glossy leaves that are narrow and thinly textured with prominently curled apices, stout and hirsute panicle with small dark pink flowers, sepals and corolla lobes shorter than the corolla tube, nectary scales very small and the capsule 1.5–1.8 mm in diameter. Dracophyllum elegantissimum is separated from similar species, with which it sometimes co-occurs, by a suite of vegetative inflorescence, floral and seed characters, which constitute evidence that it is not conspecific with them (Table 4). Superficially, D. elegantissimum resembles D. traversii but it lacks the glaucous waxy bloom so characteristic of D. traversii (S. Venter, pers. obs.). The narrow leaves of D. elegantissimum are somewhat similar to those of D. latifolium. This similarity resulted in Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A 17 B C D E F Fig. 9. Dracophyllum cosmelioides. A. Habitat along Creek Pernod. B. Flowering branch showing the horizontally spreading flowers. C. Mature plant growing in rock cracks devoid of soil. D. Flowering branch showing the erect, linear leaves and the few-flowered inflorescence. E. Plant showing the tufts of leaves at the branch apices. F. The erect leaves with blunt apices. (B–C, D, Venter 13844). Photographs: S. Venter (A–F). Cockayne (1928) confusing the two species, and incorrectly stating that D. latifolium occurs on the South Island. Mature plants display crown shapes that vary from columnar to closed-candelabra-shaped, but plants growing in forest openings can sometimes have a more open crown resembling that of D. traversii. Leaf length varies in a single population from 330 to 1000 mm, with the lamina width being 10–20(–32) mm, but most plants have leaves 500–700
15–20 mm. The inflorescence bracts vary in shape and length (270–610 mm), even on an individual plant. Selected specimens NEW ZEALAND. South Island: north-western Nelson, along the Anatori River, 26 Feb. 2002, Courtney s.n. (CHR); Kahurangi National Park, 18 Australian Systematic Botany S. Venter 165ºE 167ºE Capitale/capital city 20ºS Mer de Corail Coral Sea Ouvéa Lifou Nouvelle Calédonie New Caledonia Île sL oy au té s/L oy alt yI sla nd s Maré 22ºS 0 100 km Nouméa Île des Pins © Australian National University CartoGIS CAP 00-032 Fig. 10. Known distribution of Dracophyllum cosmelioides in New Caledonia. Gouland Downs, 14 Feb. 2002, Courtney s.n. (CHR); Knuckle Hill, 28 Mar. 1999, Venter 13778 (CHR); Abel Tasman National Park, near Moa Park, Druce s.n. (CHR); Wainui Falls, 22 Dec. 2002, Venter 13837 (CHR), Charleston, Ananui Caves, along the Nile River, 7 Jan. 1999, Venter 13749 (CHR); Westport, Mount Rochfort, 25 Nov. 1998, Venter 13732 (CHR). Dracophyllum fiordense W.R.B.Oliv., Trans. & Proc. NZ. Inst. 59: 705 (1928) Type: New Zealand, Wilmot Pass on Wilmot Saddle, Mar. 1927. W.R.B. Oliver s.n. (lecto: WELT55115!), designated by Oliver (1952). Illustrations W. R. B. Oliver, Trans. & Proc. NZ. Inst. 59: t. 15 (1928); A. Eagle, Trees & Shrubs of N.Z. 2nd edn: t. 130 (1982); J. T. Salmon, Native Trees of N.Z.: t. 72 (1989); D. Norton, New Zealand Journal of Botany (2018). Tree 1.5–5.0 m tall. Branches erect and sparsely branched. Bark on old branches greyish-brown, deeply fissured to flaky at the base on old stems and branches, young stems yellowishbrown. Leaves crowded on tips of branches in a bromelioid manner; lamina sheath 60–87
30–43 mm, coriaceous, striate, tapering and margin smooth; lamina coriaceous, linear–triangular to lanceolate, 400–700
40–50 mm, surfaces glabrous, prominently striated; margin denticulate with 10–15 teeth per 10 mm; apex acute and often spiralling. Inflorescence an axillary panicle some distance below the leaves; much shorter than the leaves, drooping, dense, 100–120(–150) mm long, pyramidal, densely branched; rachis and pedicels glabrous; inflorescence axis 5.6–7.7 mm in diameter; basal inflorescence branch 20–25(–50) mm long, widely spreading; inflorescence bracts caducous, overtopping flowers, light green, ovate–triangular at the base, 40–51
18–21 mm, surfaces glabrous, margins ciliate. Flowers hidden by leaves, 113–120, in groups of more than 10 at the base of the inflorescence; bracteoles caducous, longer than the perianth and situated in the middle of the pedicel, 4.5 – 5.0
0.8 – 1.0 mm, glabrous; pedicels straight, 0.8 – 1.5 mm long, glabrous. Sepals ovate, 2.0–2.5
2.0–3.0 mm, shorter than the corolla tube, striate, surfaces glabrous; margins ciliate. Corolla light to dark pink; corolla tube broadly campanulate, widened at mouth, 2.0–2.5 mm long and wide; corolla lobes reflexed, oblong, equalling the corolla tube, 1.5–2.0
1.3–1.5 mm, apices Taxonomic revision of Dracophyllum and Richea A Australian Systematic Botany 19 B G C F E D Fig. 11. Dracophyllum elegantissimum. A. Habit (0.12
). B. Flowering branch (0.25
). C. Ovary (10
). D. Flower (5
). E. Laid-out corolla (5
). F. Inflorescence branch (1
). G. Inflorescence bract (0.3
). Drawn from Venter 13827. Del. S. Venter. obtuse; surfaces glabrous. Stamens hypogynous, filaments 2.3–2.5 mm long; anthers exserted, oblong, light yellow and 1.5–2.0 mm long. Ovary subglobose, 0.9–1.0
1.3–1.5 mm, glabrous, apex round; nectary scales rectangular, 0.6–0.7 mm long and wide, apex retuse; style exserted, 1.8–2.0 mm long, glabrous; stigma 5-lobed. Fruit reddish-brown, 2.0–2.8
2.5–4.0 mm, depressed-globose, apex round, glabrous. Seeds brown, ovoid, 0.55–0.6 mm long, testa slightly reticulate (Fig. 14, 15). Distribution and ecology Endemic to the Western Otago and Fiordland areas in New Zealand. There are two main distribution areas, the one 20 Australian Systematic Botany S. Venter C B D E Fig. 12. Dracophyllum elegantissimum. A. In forest habitat, Karamea, north-western Nelson. B. Young plant showing the characteristic long and narrow leaves. C. Characteristic peeling bark on old stem and young branch. D. Plant in fruit showing the characteristic long, narrow leaves with coiled leaf tips. E. Inflorescence showing the small flowers, Mount Rochfort. Photographs: Phil Bendle (A, B and D) and S. Venter (C and E). surrounding the Mount Cook and Westland Tai Poutini National Park, and the other in Fiordland National Park (Fig. 16). Recent northern range extension (75 km) as far as the Hokitika River catchment of 75 km was reported by Norton (2018). Dracophyllum fiordense is common on steep (50–80) northern, north-western and north-eastern slopes from near Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 21 180º –35º −40º −45º Fig. 13. Known distribution of Dracophyllum elegantissimum, South Island, New Zealand. sea level (50 m) to areas of high elevation (1280 m). The vegetation consists of permanently moist lowland, subalpine forest, subalpine woodland and shrubland or tussock grassland in mountain gullies, ravines, on ridges or on bluffs. The soil is humus-rich brown clay loam derived from schist and gneiss. Most populations grow in full sun, but occasionally some individuals grow in light shade, although this does not alter their external appearance. Plants occur in high rainfall areas and receive additional moisture from mist. Phenology Flowering January–March. Etymology Named after the Fiordland area in New Zealand. Diagnostic features and discussion Dracophyllum fiordense is characterised by the unbranched (occasionally only once) stems, very large and long leaves with narrowed lamina bases and prominently spiralled apices, the panicle situated below the leaves, sepals shorter than the corolla tube with spaced cilia on the margin, bracteoles longer than the flower, with a few teeth at the apex and the anthers exserted with the capsule short and very broad. Dracophyllum fiordense is similar to D. menziesii and D. townsonii (Oliver 22 Australian Systematic Botany S. Venter Table 4. Diagnostic characters of Dracophyllum elegantissimum, D. traversii and D. latifolium Character Crown habit Leaf size (mm) Rachis and pedicel Inflorescence-axis diam. (mm) Inflorescence-bract length (mm) Bracteole size (mm) Sepal size (mm) Sepal length (mm) Corolla-tube size (mm) Corolla-lobe size (mm) Corolla-lobe length Filament length (mm) Anther length (mm) Nectary-scale size (mm) Nectary-scale apex Ovary size (mm) Style length (mm) Fruit size (mm) Seed shape Seed length (mm) D. elegantissimum D. traversii D. latifolium Closed candalabrum 330–1000
10–20(–32) Tomentose 7.6–13.5(14.0) 270–610 2.0–6.5
0.5–1.0 1.0–2.0
1.0–1.5 <Corolla tube 1.0–2.0
1.3–2.0 1.2–1.4
1.0–1.3 <Corolla tube 0.3–0.5 0.9–1.3 0.6–1.0
0.5–1.0 Subacute to irregularly toothed 1.0–1.5
1.3–1.5 1.5–1.7 1.2–1.5
1.5–1.8 Filiform 0.7–0.8 Open candalabrum 90–300
40–50 Pubescent 13.0–16.5 130–240 4.0–4.8
0.5–0.7 2.0–3.0
2.0–2.5 Equalling corolla tube 2.7–3.0
4.0–5.1 2.5–2.8
2.0–2.5 >Corolla tube 1.0–1.2 1.8–2.0 1.0–1.5
1.0–1.5 Retuse 1.4–1.5
1.8–2.0 2.0–2.2 1.9–2.0
2.8–3.0 Ovoid 1.0–1.2 Open candalabrum 45–500
12–30 Pubescent or tomentose 15–20 105–180 2.0–5.5
0.5–1.7 0.7–1.5
1.0–1.7 <Corolla tube 1.5–2.0
1.5–2.5 1.5–2.0
1.5–2.0 >Corolla tube 1.0–1.2 1.3–1.5 0.6–1.2
0.8–1.0 Irregularly toothed 0.8–1.0
1.0–1.5 1.0–1.7 1.0–2.0
2.0–3.0 Ovoid 1.2–1.3 E F A G H B D C Fig. 14. Dracophyllum fiordense. A. Habit (0.25
). B. Inflorescence (0.5
). C. Flower (5
). D. Laid-out corolla (5
). E. Leaf (0.5
). F. Ovary (10
). G. Inflorescence bract (1
). H. Sepal (5
). Drawn from Venter 13801. Del. S. Venter. 1928) in the axillary panicles situated below the leaves. It resembles D. traversii, but differs in the unbranched to sparsely branched stems, fissured to flaky grey bark (Norton 2018), the large leaves with spiralled apices and the inflorescences situated below the leaves. The panicles are more branched than those of D. menziesii but closely resemble those of D. townsonii in size and shape. Leaves collected from populations of D. fiordense in the Franz Josef area are nearly 50 % shorter, with the main stems more branched than the plants from further south (Fig. 16, 17). This can possibly be attributed to the higher elevation, lower rainfall and higher temperatures of the northern populations. Plants from protected valleys on Mount Alexander attain a height of 5 m, compared with plants from exposed areas at high altitude that are not taller than 1.5 m. Fruit size varies (2.0–2.8
2.5–4.0 mm), with the largest fruit being found in the southern populations. Selected specimens NEW ZEALAND. South Island: Alex Knob, gully between Alex Knob and Louisa Peak, 19 Mar. 2000, Venter 13801 (CHR); ibid., 17 Jan. 1951, Oliver s.n. (WELT 13767); Westland National Park, head of Architect Creek, 7 Feb. 1971, Wardle s.n. (CHR); Westland National Park, between Twain Col and Douglas River, 3 Feb. 1971, Wardle s.n. (CHR); Westland National Park, between Mount Moltke and Cape Defiance, Castle Rocks, 9 Feb. 1971, Wardle s.n. (CHR); Whataroa Valley, Rocky Creek, May 1998, Giller s.n. (CHR); Westland National Park, Copland Valley head, near Copland Glacier Lake, 18 Nov. 1975, Wilson & Verhoef s.n. (CHR); Cleddau Valley, 20 Dec. 1944, Oliver s.n. (WELT 55113); Karangarua River, between Top Flat and Twain Saddle, 29 Mar. 1969, Wardle & Fryer s.n. (CHR); Fiordland, Sinbad Gulley, 27 Feb. 1975, Johnson s.n. (CHR); Thompson Sound, mountains above Lyall Bay, Jan. 1958, Metcalf s.n. (CHR); Caswell Sound, Mount Alexander, 29 Mar. 1949, Zotov s.n. (CHR); Wilmot Pass, Wilmot Saddle, Mar. 1927, Oliver s.n. (WELT). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 23 A C B D Fig. 15. Dracophyllum fiordense. A. Mature plants, Dark Cloud Range, Fiordland. B. Inflorescence below the leaves (Venter13801). C. Mature plants showing the typical bromelioid growth habit of the leaves. D. Nothofagus forest with D. fiordense, Fiordland. Photographs: Grant Dixon (A) and S. Venter (B, C). Dracophyllum fitzgeraldii Moore & F. Muell., Frag. Phyto. Austr. 7: 27 (1869). (As ‘Fitzgeraldi’) Type: Australia. Insula Lord Howe’s Island. 1869. Rob D. Fitzgerald 43 (holo: MEL!). Illustrations W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 21 (1928); H. F. Recher and S. S. Clark, Environmental Survey of Lord Howe Is.: 26 (1974); I. Hutton, Lord Howe Is.: 123 (1986); P. S. Green, Oceanic Islands. Fl. of Austr.: 49 (1): t. 42 (1994). 24 Australian Systematic Botany S. Venter 170º 175º 180º −35º −30º −35º −45º Fig. 16. Known distribution of Dracophyllum fiordense, South Island, New Zealand. A shrub to tree (1–)3–13 m tall. Branches: bark on old branches dark brown to blackish- brown, rough to deeply fissured, young stems reddish-brown. Leaves crowded on tips of branches in a bromelioid manner; lamina sheath light brown, 23–29
15–21 mm, coriaceous, striate, membranous, tapering, top half minutely ciliate; lamina coriaceous, light to mid-green, abaxial surface lighter coloured, linear–triangular, rarely lanceolate (150–)205–335 (–350)
(7–)9–11(–25) mm, surfaces glabrous, prominently striated; margin cartilaginous, serrate to serrulate with 16–38 teeth per 10 mm; apex acute. Inflorescence shorter than the leaves, erect, dense, 100–150(–280) mm long, densely branched; rachis and pedicels pubescent; inflorescence axis 3.0–5.5 mm in diameter; basal inflorescence branch 10–12 mm long, widely spreading; inflorescence bracts caducous, overtopping flowers, light green, ovate–lanceolate at the base, 29–49
11.8–15.2 mm, adaxial surfaces glabrous; abaxial surfaces scabrid, margins serrulate. Flowers 100–600+, in groups of more than 10 at the base of the inflorescence, pedicellate; bracteoles persistent, recaulescent with 1 Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 25 A B Fig. 17. Variation in Dracophyllum fiordense. Plants from northern populations (closed squares) tend to have branched main stems, small leaves and fruit (A. Westland National Park, Alex Knob, Venter 13801), compared with the southern populations (closed circles) with unbranched main stems, much longer leaves and larger fruit (B. Fiordland National Park, Sinbad Gulley, Johnson s.n.). bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, 4.5–6.0
0.5–0.7 mm, glabrous; pedicels straight, 1–2 mm long, pubescent. Sepals lanceolate to ovate–lanceolate, (3.0–) 4.4–5.0(–6.0)
1.5–2.5 mm, shorter than the corolla tube, white, striate, surfaces glabrous; margins ciliate. Corolla white, occasionally pale pink; corolla tube cylindrical, widened at mouth, (4,0–)5.3–6.0
2.0–2.3 mm; corolla lobes reflexed, triangular, shorter than corolla tube, 2.0–2.5
1.3–1.6 mm, apices subacute; surfaces glabrous. Stamens adnated to top of corolla tube, filaments 0.3–0.5 mm long; anthers slightly exserted, oblong, light yellow and 0.8–0.9 mm long. Ovary subglobose, 1.0–1.5
1.2–1.25 mm, glabrous, apex round; nectary scales rectangular, 0.6–0.7
0.3–0.5 mm, apex retuse to irregularly toothed; style included, 0.9–1.0 mm long, glabrous; stigma 5-lobed. Fruit pedicellate, dark brown, (3.9–)4.4–5.0
1.8–2.0 mm, broadly obovoid, apex round, glabrous. Seeds yellowish-brown, ovoid, 0.6–0.65 mm long, testa slightly reticulate (Fig. 18, 19). Distribution and ecology Endemic to Lord Howe Island restricted to the upper slopes of Mount Gower and that of Mount Lidgebird (Fig. 20). Dracophyllum fitzgeraldii occurs on gentle to steep (10–70) northern to north-western slopes at 200–1000-m elevation, which are covered in dense oceanic montane cloud forest and shrubland. The soil is clay loam derived from basalt. 26 Australian Systematic Botany B A S. Venter C D On the slopes of Mount Gower and in the Erskine Valley, D. fitzgeraldii grows as a tree (up to 13 m tall). Where the habitat is more exposed, the vegetation changes from forest to closed shrubland where Dracophyllum plants rarely exceed 2 m in height, especially on the high plateau. Inflorescence length varies from plant to plant (100–150 mm) and most corolla lobes have subacute apices but there are some specimens with obtuse apices. The apices of the nectary scales vary from retuse to irregularly toothed. Selected specimens E G F H Fig. 18. Dracophyllum fitzgeraldii. A. Flowering branch (0.25
). B. Leaf (0.5
). C. Ovary (10
). D. Sepal abaxial surface (5
). E. Flower (5
). F. Inflorescence-bract adaxial surface (1
). G. Laid-out corolla (5
). H. Bottom inflorescence branch (2
). Drawn from Green 1656. Del. S. Venter. Phenology Flowering October–February(–April). Etymology Named after Robert D. Fitzgerald F.L.S. (1830–1892), Deputy Surveyor General of New South Wales, who collected the type. Diagnostic features and discussion Dracophyllum fitzgeraldii is characterised by the spreading crowns, deeply fissured and rough bark, lamina sheath 23–29
15–21 mm, lamina 205–335
9–11 mm with prominent striation and drawn-out lamina apices, panicle shorter than the leaves with flowers arranged in groups of >10 at the base of the inflorescence, sepals shorter (4.4–5.0
1.5–2.5 mm) than the 5.3–6.0-mm long corolla tube, anthers slightly exserted with the ovary subglobose and the style included. Dracophyllum fitzgeraldii is related to species in northern Queensland (D. sayeri) and New Caledonia (D. verticillatum; Oliver 1952; Green 1994; Streiber et al. 1999). Dracophyllum fitzgeraldii is similar to D. milliganii, D. sayeri, D. oceanicum and D. verticillatum (Table 5). AUSTRALIA. Lord Howe Island: eastern side of Mount Lidgebird, near the Goat House, 28 Aug. 1969, Game 69/248 (K); beyond Goat House around Mount Lidgebird, 2 Dec. 1993, Beard s.n. (HO, WAIK); near Goat house Cave, 1 Sep. 1981, Rodd 3714 (NSW); along footpath south of Goat House cave, 7 Apr. 2006, Venter 13863 (CHR, NSW); northern spur of Mount Gower, Johnson & Rodd 1366 (K, NSW); summit of Mount Gower, Green 1656 (K, MEL); Mount Gower, Get Up Place, 17 Sep. 1998, Parkes s.n. (CHR); Mount Gower, 6 June 1882, Duff s.n. (MEL); Erskine Valley, Mount Gower Track, 27 Sep. 1980, Rodd 3591 (NSW); The Saddle, 14 Apr. 1999, Hutton s.n. (NSW); along Erskine Creek on The Saddle, 9 Apr. 2006, Venter 13862 (CHR, NSW); Mount Lidgbird, 22 July 1965, Van Balgooy 1076 (L); Erskine Valley, Von Mueller 17 (BM); Erskine Valley to summit of Mount Gower, 17 May 1920, Boorman s.n. (NSW). Dracophyllum involucratum Brongn. & Gris, Ann. Sci. Nat. Bot. sér(5) 2: 157 (1864) Type: New Caledonia. Montagne d’Yate. Vieillard 832 (lecto: P!, isolecto: NSW!, P!) designated by Oliver (1952); New Caledonia. s. loc., Pancher 367 (remaining syn: P!.) designated by Virot (1975). Dracophyllum compactum S. Moore, J. Linn. Soc. Bot. 45: 349 (1921). Type: New Caledonia. Plaine des Lacs, 23 Feb. 1914. R. H. Compton 371 (holo: BM!). Illustrations W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 25 (1928); R. Virot, Fl. Nov. Calédonie et Dépend. 6: t. 17 (1975). Trees 2–4 m tall. Branches spreading sparsely branched. Bark on old branches grey to brown, smooth, young stems yellowish-brown. Leaves spreading to recurved; lamina sheath light green to light brown, 40–48
25–30 mm, coriaceous, striate, tapering and margin smooth or with the top half minutely ciliate; lamina with lower surface lighter green, linear–triangular to sometimes lanceolate, (150–) 250–300(–500)
15–25(–30) mm, surfaces glabrous, prominently striated; margins serrulate with 6–8 teeth per 10 mm; apex acute to rarely obtuse. Inflorescence overtopping the leaves, erect, dense, 220–400 mm long, linear–oblong and densely branched; rachis and pedicels tomentose; inflorescence axis prominently ribbed, 4–7 mm in diameter; basal inflorescence branch 1–2 mm long, widely spreading; inflorescence bracts caducous, overtopping flowers, light green, ovate–lanceolate at the base, 25–30(–40)
10–12 mm, surfaces glabrous, margins ciliate. Flowers 400–600+, in groups of more than 10 at the base of the inflorescence, pedicellate; flower bracts persistent, shorter than flowers, coriaceous, ovate Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A 27 B C D E Fig. 19. Dracophyllum fitzgeraldii. A. Habitat on slopes of Mount Lidgebird. B. Flowering branch. C. Deeply fissured and rough bark. D. Flowers. E. Mature plant. B–E from Venter 13863. Photographs: S. Venter (A–E). to broadly ovate, 2.5–4.0
1.5–2.5 mm, surfaces glabrous, striate, margins ciliate; apices acute; bracteoles caducous, situated at the base of the pedicel and the pedicel above the bracteoles covered in persistent bracts, shorter than the flower, 0.5–1.0
0.1–0.2 mm, glabrous; pedicels straight, (1.5–) 4–12 mm long, tomentose. Sepals lanceolate to occasionally ovate–lanceolate, 2.5–5.0
2.0–2.5 mm, shorter than the corolla tube, striate, surfaces glabrous; margins ciliate. Corolla white to greenish-white; corolla tube narrowly campanulate; (2–)4–5
1.4–1.5 mm; corolla lobes reflexed, ovate–triangular, shorter than corolla tube, 1–2
1.2–1.5 mm, apices obtuse; surfaces glabrous. Stamens adnate to top of corolla tube, filaments 1.0–1.5 mm long; anthers exserted, oblong and purple when young turning light yellow when mature and 1.2–1.5 mm long. Ovary subglobose to mainly obovoid, 1.0–1.2
1.0–1.3 mm, glabrous, apex round; nectary scales Australian Systematic Botany S. Venter contour interval 150 m Malabar 209 m Malabar Ridge Mount Ellza 147 m Dawsone Ridge Transit Hill 121 m ing Tre e Ridg r Intermediate Hill 205 m Smok 28 Goat House 0 1 Mount Lidgbird 777 m 2 km Erskine Valley The Saddle 150 300 450 Get up Place Mount Gower 875 m 159º05' Fig. 20. Known distribution of Dracophyllum fitzgeraldii, Lord Howe Island. Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 29 Table 5. Morphological differences among Dracophyllum fitzgeraldii, D. milliganii, D. oceanicum, D. sayeri and D. verticillatum Parameter Lamina sheath length (mm) Lamina margin texture Number of teeth per cm Inflorescence length (mm) Inflorescence axis texture Inflorescence-bract length (mm) Inflorescence-bract margin Sepal to corolla-tube length Corolla-tube length (mm) Stamen fusion Filament length (mm) Nectary-scale fusion D. fitzgeraldii D. milliganii D. oceanicum D. sayeri D. verticillatum 23–29 Serrulate 16–20 100–150 Smooth 24–49 Serrulate Shorter 5–6 Top 0.3–0.5 Free 30–40 Serrulate 15–70 130–460 Ribbed 40–180 Serrulate Equalling 2.5–3.0 Free 2.5–5.0 Free 12–17 Serrulate 10–20 70–100 Smooth 10–65 Entire/serrulate Shorter/equalling 4–7 Middle 3.0–4.5 Free 20–27 Smooth Absent 160–210 Smooth 85–110 Entire Shorter 3–5 Top 1.0–1.3 Fused at the base 17.3–25.2 Serrate or serrulate 20–32 190–400 Smooth 42–43 Ciliate Shorter 2–3 Upper third 1.0–1.5 Fused at base rectangular, 0.5–0.6 mm, apices retuse to irregularly toothed; style included, 1.5–2.0 mm long, glabrous; stigma 5-lobed. Fruit dark to reddish-brown, 2–3 mm long and wide, obovoid, apex round, glabrous. Seeds brown, filiform, 0.9–1.0 mm long, testa slightly reticulate (Fig. 21, 22). E A F Distribution and ecology A New Caledonia endemic restricted to the montane areas east of Tontouta as far north as Thio (Fig. 23). Dracophyllum involucratum grows on mountain slopes, plateaus and in valley floors at altitudes of 150–1400 m. The vegetation consists of maquis and forest. Soils are ferruginous, mostly rocky loam to clay loam and derived from peridotite or serpentinite. Plants grow mostly in full sun but some populations occur in light shade inside the forest. Phenology Flowering October–April. Etymology The specific epithet describes the prominent involucres of flowers. G B Diagnostic features and discussion Dracophyllum involucratum is characterised by leaves having long drawn apices, remote minute teeth on the lamina margin, ribbed and densely tomentose rachis with flowers arranged in dense contracted clusters surrounding the rachis at regular intervals, peduncles clothed in persistent imbricating bracts and the narrow corolla tube. Dracophyllum involucratum is similar to D. sayeri, D. elegantissimum and D. fitzgeraldii, but can be distinguished by the flowers being in involucres and the pedicel of each individual flower covered with tightly overlaid persistent bracts. These characters prompted Oliver (1952) to describe the subgenus Cordophyllum to accommodate D. involucratum. The results of cladistic and phenetic studies (S. Venter, unpubl. data) and of a molecular study (Wagstaff et al. 2010) have shown that it is nested within Dracophyllum and is included as such to avoid paraphyly. Dracophyllum involucratum is a species showing limited variation. Some variation occurs in inflorescence length C D Fig. 21. Dracophyllum involucratum. A. Flowering branch (0.25
). B. Flower (5
). C. Laid-out corolla (5
). D. Sepal adaxial surface (5
). E. Leaf (0.5
). F. Ovary (10
). G. Inflorescence bract (5
). Drawn from MacKee 21621. Del. S. Venter. (220–400 mm), flower-bract size (2.5–4.0
1.5–2.5 mm), sepal length (2.5–5.0 mm) and nectary scale apices that are either retuse or irregularly toothed. 30 Australian Systematic Botany S. Venter A B C D Fig. 22. Dracophyllum involucratum. A. Habitat at Pic du Pin. B. Adult plant in habitat, Pic du Pin. C. The characteristic involucres with young fruit. D. Fruiting plant, Pic du Pin (Venter 13850). Photos: S. Venter (A–D). Selected specimens NEW CALEDONIA. Province Sud: Mount Kouakoué, 20 Nov. 1981, MacKee 39946 (NOU); ibid., 1 Dec. 2002, Tronchet 609 (NOU, MO); Montagne des Sources, 1 Apr. 1968, Bernardi 12464 (K); Yate, 18 Mar. 1969, Jaffré 175 (NOU, P); 4 km south-east of S2 TER on road to Carenage, 19 July 1977, Whaite 3697-11 (NSW); track, ~0.3 km off Carenage Road, 2 km directly east-north-east of Pic du Pin, Weston 1685 (NOU, NSW); Creek Pernod, 22 Mar. 1967, Nothis 333 (NOU); Vallée du Pernod, 9 Mar. 1967, Schmid 1956 (NOU); Chute de la Madeleine, 28 June 1966, Schmid 1358 (NOU); Prony, Dec. 1913, Franc 1518 (Z); Plaine des Lacs, 6 Feb. 1926, Däniker 3074 (Z); ibid., 23 Feb. 1914, Compton 371 (BM); Plaine des Lacs, Ruisseau Pernod, 2 Nov. 1967, MacKee 17841 (L, NOU); ibid., 25 Dec. 1967, MacKee 18161 (K, L, NOU); Vallée des Lacs, 7 Oct. 1950, Guillaumin & BaumannBodenheim 6747 (Z); Ouénarou, 22 Feb. 1970, MacKee 21621 (K, L, NOU); Ouenghi, east de la Dent de Saint-Vincent, 22 Dec. 1970, MacKee 23100 (L); Pic du Pin, 14 May 2005, Venter 13850 (NOU). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 165ºE 31 167ºE Capitale/capital city 20ºS Mer de Corail Coral Sea Ouvéa Lifou Nouvelle Calédonie New Caledonia Île sL oy au té s/L oy alt yI sla nd s Maré 22ºS 0 100 km Nouméa Île des Pins © Australian National University CartoGIS CAP 00-032 Fig. 23. Known distribution of Dracophyllum involucratum, New Caledonia. Dracophyllum latifolium A.Cunn., Ann. Nat. Hist. 2: 48 (1837) Type: New Zealand. Bay of Islands, dry woods on the Kana Kana [Kawakawa] River, 1826. A. Cunningham s.n. (lecto: BM!; isolecto: MEL!; WELT 79400!), designated by Oliver (1952). Dracophyllum recurvatum Colenso, Trans. & Proc. NZ. Inst. 21: 92 (1889). Type: New Zealand. Hills around Lake Waikare [Lake Waikaremoana], on high grounds, 1888. H.T. Hill & T. Kirk s.n. (holo: WELT 23606!, iso: AK 6904!; K!; WELT!). Dracophyllum latifolium var. matthewsii Carse, Trans. & Proc. NZ. Inst. 48: 238 (1916). Dracophyllum matthewsii Carse, Trans. Proc. NZ. Inst. 56: 86 (1926); Oliver, Trans. Roy. Soc. N.Z. 80 (1): 16 (1952). Type: New Zealand. North Island, near Kaitaia, Taumatamahoe, 1900 feet [~579 m], Oct. 1913. H. Carse & H.B. Matthews s.n. (holo: CHR 332571!; iso: AK 6910!, AK6911!), designated by Oliver (1952). Illustrations T. Kirk, Forest Fl. N.Z.: t. 123 (1889); T. F. Cheeseman, Illustr. N.Z. Fl., t. 129 (1914); A. Eagle, Trees & Shrubs of N.Z.: tt. 129A, B (1982); J. T. Salmon, Native Trees of N.Z.: 271 (1989) Tree 3–10 m tall. Branches form an open candelabrumshaped crown. Bark on old branches greyish-brown to brown, rough or flaky, young stems yellowish-brown. Leaves crowded at tips of branches in a bromelioid manner; lamina sheath 30–65
18–55 mm, striate, membranous, tapering and margin smooth; lamina linear–triangular to rarely lanceolate, (100–)145 –700(–800)
12–30 mm, surfaces glabrous, prominently striated; margins serrate to denticulate with 2–4 teeth per 10 mm; apex thickened. Inflorescence shorter than leaves, erect to drooping, dense, 100–340(–400) mm long, oblong to pyramidal and densely branched; rachis and pedicels pubescent to tomentose; inflorescence axis yellowish to light green, 15–20 mm in diameter; basal inflorescence branch 30–60 mm long, suberect to at right angles with inflorescence axis; inflorescence bracts caducous, overtopping flowers, whitish at the base and pink tipped, broadly ovate to ovate–triangular 32 Australian Systematic Botany at the base, 105–180(–210)
20–35 mm, surfaces glabrous, margins ciliate, apices acute. Flowers 600–2000+, in groups of 5–10 at the base of the inflorescence, pedicellate; bracteoles caducous, recaulescent, with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, (1.5–)2.0–5.5
0.5–1.7 mm, glabrous; pedicels straight, 1.0–2.5 mm long, pubescent to tomentose. Sepals broadly ovate to triangular, 0.7–1.5
1.0–1.7 mm, shorter than the corolla tube, striate, adaxial surfaces glabrous; abaxial surfaces pubescent; margins with the upper-third toothed. Corolla dark pink to dark red; corolla tube campanulate to broadly campanulate, widened at mouth, 1.5–2.0
1.5–2.5 mm; corolla lobes reflexed, oblong to ovate–triangular, longer than corolla tube, 1.5–2.0 long and wide, apices obtuse, rarely subacute; surfaces glabrous. Stamens inserted at top of corolla tube, filaments (0.5–) 1.0–1.2 mm long; anthers exserted, rectangular, pink turning light yellow with age and 1.3–1.5 mm long. Ovary ovoid, 0.8–1.0
1.0–1.5 mm, glabrous, apex round; nectary scales rectangular to oblong, 0.6–1.2
0.8–1.0 mm, apex retuse to irregularly toothed; style exserted, 1.0–1.7 mm long, glabrous; stigma clavate to 5-lobed. Fruit not included in persistent calyx, reddish to purplish-brown, 1–2
2–3(–4) mm, depressed-globose, apex round and glabrous. Seeds yellowish-brown, ovoid, 1.2–1.3 mm long, testa slightly reticulate (Fig. 24, 25). S. Venter E A F B Distribution and ecology Endemic to the North Island of New Zealand (Fig. 26). The known localities are north of a line from Mount Egmont to Hawke’s Bay. Dracophyllum latifolium occurs on gentle to steep (0–70) slopes in river valleys, along stream banks and on mountain slopes from sea level to 1100-m elevation. Dracophyllum latifolium is an important element of the small tree and shrub tier of the low-altitude conifer–broadleaved forest (Wardle 1964, 2002). Soils are brown sandy loam derived from calcareous sandstone and greywacke or dark brown clay loam or clay derived from andesite, basalt, rhyolite, siltstone and mudstone. Dracophyllum latifolium sometimes grows in full sun, but is primarily a shade lover of forest communities. It is not known to occur in areas that are exposed to salt spray. Phenology Flowering September–May. Etymology: Describes the narrow leaves, a prominent feature of this species. Diagnostic features and discussion Dracophyllum latifolium is characterised by the rough to flaky bark; leaves recurved in a bromelioid manner, glabrous, thinly textured; panicle slender, erect or drooping with the pubescent branches at acute angles, flowers purplish-red, capsules 2.0–2.5 mm in diameter with the pedicels 1.5–2.5 mm long. Young plants form erect, unbranched stems with a tuft of leaves at the top, a character shared with D. fiordense, D. fitzgeraldii, D. elegantissimum, D. involucratum, D C Fig. 24. Dracophyllum latifolium. A. Flowering branch (0.25
). B. Inflorescence bract (1
). C. Flower (5
). D. Lower inflorescence branch (1
). E. Leaf adaxial surface (0.5
). F. Ovary (10
). Drawn from Venter 13764. Del. S. Venter. D. ramosum, D. townsonii, D. traversii and D. verticillatum. Dracophyllum latifolium is similar to D. traversii, but differs in having narrower leaves (12–30 mm compared with 40–50 mm), lamina margin serrate to denticulate (not serrulate) and having fewer teeth on the lamina margin (2–4 compared with 18–20 per 10 mm). The flowers are in groups of 5–10 (compared with >10), sepals shorter than the corolla tube, with the upper half distinctly toothed not ciliate, corolla tube shorter and narrower, and ovary ovate and much smaller, with the seeds larger than those of D. traversii. Oliver (1928, 1952) and Allan (1961) kept D. matthewsii separate from D. latifolium on the basis of the drooping inflorescence. In many D. latifolium populations, drooping inflorescences occur together with erect inflorescences (S. Venter, pers. obs.), making this an unreliable character to separate species. Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 33 A B C D E Fig. 25. Dracophyllum latifolium. A. Montane habitat, Coromandel Peninsula. B. Bark of mature specimen. C. Mature plant. D. Flowering plant from Mount Rowe (Venter 13764). E. Plant with drooping inflorescence (=D. matthewsii) from Mount Kaitarakihi on the Coromandel Peninsula. Photos: Kathrin Marks (A), S. Venter (B, D–E) and Phil Bendle (C). Inflorescence bracts vary from broadly ovate to ovate–triangular and can sometimes be short and narrow, especially in populations growing in full sun. Sepal size varies (0.7–1.5
1.0–1.7 mm), with corolla lobes being oblong to ovate–triangular and this does not seem to be associated with specific populations. The nectary scales show variation in shape (rectangular to oblong), size (0.6–1.2
0.8–1.0 mm) and shape of the apices (retuse to irregularly toothed). Selected specimens NEW ZEALAND. North Island: near Kaitaia, Jan. 1913, Matthews & Carse s.n. (CHR); Manganui County, Kaiaka, 3 Jan. 1904, Carse s.n. (CHR); Pukepoto, 1 Oct. 1915, Carse & Matthews s.n. (CHR); Maungataniwha, Nov. 1920, Carse 7573 (K); Puketi State Forest, Pukatea Ridge Track, 9 Oct. 1973, Orchards 4075 (AK); north of Whangaroa Harbour, 27 Sep.1990, Wright 10520 (AK, HO); Bay of Islands, Kerikeri, Sep. 1977, Bartlett & O’ Brien s.n. (CHR); Waima Forest, Sep. 1991, Druce 62 (CHR); Waipoua State Forest, 2 miles [~3.2 km] south of Headquarters, 12 Apr. 1972, Rawlings, Esler, Smith & Astridge 3830 (CHR); Otaua, Nov. 1874, Berggren s.n. (W); Bay of Islands, Cape Brett, Collett s.n., Sep 1964 (CHR); Little Barrier Island, July 1949, Parkin s.n. (CHR); Wayby, 31 Jan. 1944, McKenzie s.n. (AK); Great Barrier Island, T. Kirk s.n. (CHR); Mount Kohukohunui summit, Thames, Mount Kaitarakihi, 12 Feb. 1999, Venter 13763 (CHR); Mount Te Aroha, near top, 2 July 1939, Molesworth s.n. (AK); Kaimai Range, 34 Australian Systematic Botany S. Venter 170º 175º 180º –35º –40º –45º Fig. 26. Known distribution of Dracophyllum latifolium, North Island, New Zealand. Ngatamahinerua, July 1977, Bartlett s.n. (CHR); Huia, Rickard’s Bush, 6 Aug. 1949, Wood s.n. (AK); Tauranga, McLaren’s Falls, Simpson 1960 (CHR); Raukumara Range, Mount Honokawa, Feb. 1969, Druce s.n. (CHR); Raukumara Range, Mount Aorangi, Jan. 1983, Druce s.n. (CHR); Herangi Range, Mangatoa Road near Saddle, Mar. 1972, Druce s.n. (CHR); Waitaanga Plateau, Feb. 1967, Druce s.n. (CHR); Mount Pirongia peak, 31 Mar. 1940, Molesworth s.n. (AK); Urewera National Park, Lake Waikareiti, Sandy Bay, Apr. 1973, Druce s.n. (CHR); Mount Whakapunake, Dec. 1967, Druce s.n. (CHR); Mount Hikurangi, 14 Feb. 1966, Burke s.n. (WELTU). Dracophyllum mackeeanum S.Venter, New Zealand J. Bot. 42: 747–751 (2004) Virot, Fl. Nov. Calédonie et Dépend. 6: 147–151 p.p. (1975). Type: New Caledonia. Province Sud, Upper Rivière Tontouta, near Mine Gallieni, 25 Dec. 1960. H.S. MacKee 7745 (holo: NSW 461862!; iso: CANB!, L!). Illustrations R. Virot, Fl. Nov. Calédonie et Depend. 6: t. 26 (1975) (as D. ramosum); S. Venter, New Zealand. J. Bot. 42: 748, t. 1 (2004). A fairly compact and multi-stemmed shrub, up to 1.5 m tall. Branches: bark on old stems greyish-brown with spaced deep furrows, young stems yellowish to reddish-brown, smooth and covered with fine whitish antrorse, multicellular hairs. Leaves erect–spreading; lamina sheath slightly wider than the lamina, light brown to reddish-brown, 10–16
Taxonomic revision of Dracophyllum and Richea 7.0–9.6(–13.35) mm, coriaceous, striate, shoulder tapering with the top half shortly ciliate; lamina coriaceous, light green to dark green above and lighter beneath, elongatetriangular, 27.8–58.6(–128)
3.8–7.8(–11.53) mm, flat, adaxial surface glabrous to pubescent but abaxial surface tomentose, prominently striated; margins serrulate with 22–40 teeth per 10 mm; apex subacute to obtuse. Inflorescence exceeding the leaves, erect, dense, 72.0–81.5 mm long, linear–oblong and sparingly branched; rachis and pedicels white pubescent; inflorescence axis prominently ribbed, 2.0–2.7 mm in diameter at the base; basal inflorescence branch 5–12 mm long, sub erect; inflorescence bracts caducous, longer than flowers, straw yellow-tinged pink to light red, broadly ovate at the base with apices drawn-out, 15–23
10–15 mm; adaxial surfaces glabrous, apices pubescent, abaxial surfaces white tomentose, margins ciliate. Flowers 26–32, arranged in groups of three on the basal inflorescence branches, pedicellate; bracteoles persistent, recaulescent with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, weakly keeled, 3–4
0.2–0.5 mm, ciliate; pedicels fused at the base and flattened, middle pedicel longest, 1.5–3.0
0.5 mm, tomentose, lateral pedicels 1.5–2.0
0.5 mm. Sepals rose to light pink, ovate–lanceolate, 4–6
1.5–1.7 mm, equal to or longer than corolla tube, striate, adaxial surface glabrous, abaxial surface pubescent; margin ciliate. Corolla white; corolla tube cylindrical, 4.5–5.0
1.2–1.3 mm, exterior pubescent in distal part, slightly verrucose inside; corolla lobes spreading to horizontal, broadly lanceolate, shorter than corolla tube, 2.0–2.7
1.2–1.5 mm, margin laciniate; adaxial surface papillate, abaxial surface pubescent, apex obtuse. Stamens inserted at mid-point of corolla tube; filaments 1.0–1.5 mm long; anthers included, oblong, light yellow, 0.5–0.7 mm long. Ovary ovoid, 0.9–1.0 mm long and wide, glabrous, apex round; nectary scales rectangular, 0.6–0.7
0.4–0.5 mm, apices retuse; style included, 0.95–1.0 mm long, glabrous; stigma included, 5-lobed. Fruit light to dark brown, ovoid, 1.5–2.0 mm long and wide, apex round, glabrous. Seed yellowish-brown, ovoid, 0.75–0.8 mm long, with slightly reticulate testa (Fig. 27, 28). Distribution and ecology New Caledonian endemic restricted to the valleys of the Thio, Tontouta and Dombea rivers (Fig. 29). Dracophyllum mackeeanum occurs on gentle (5–15) to steep (40–50) hill slopes at 200–600-m elevation. It is restricted to low maquis vegetation in exposed open areas. Soils are rocky clay loam derived from serpentinite. Phenology Flowering October–December. Etymology Named after Dr H. S. MacKee, University of Sydney, who made extensive collections in New Caledonia. Australian Systematic Botany H 35 A H G C F D E Fig. 27. Dracophyllum mackeeanum. A. Flowering branch (1
). B. Ovary (10
). C. Flower (5
). D. Inflorescence-bract adaxial surface (2
). E. Laidout corolla (5
). F. Calyx (5
). G. Cross-section through inflorescence axis (10
). H. Leaf (1
). Drawn from MacKee 7745. Del. S. Venter. Diagnostic features and discussion Dracophyllum mackeeanum is characterised by the young stems covered in fine whitish hairs lying flat on the surface, leaves glabrous to pubescent adaxially and tomentose abaxially, striate and light green; prominently ribbed and white pubescent inflorescence axis, abaxially whitish tomentose inflorescence bracts with flowers arranged in groups of three on the basal inflorescence branches, the base of pedicels fused and flattened, cylindrical corolla tube being 4.5–5.0 mm long, with the distal part of the outer surface pubescent, broadly lanceolate corolla lobes that are abaxially pubescent with laciniate margins and stamens inserted in the 36 Australian Systematic Botany S. Venter A B C D Fig. 28. Dracophyllum mackeeanum. A. Habitat along the To De River near Thio. B. Young inflorescence showing the shape of the inflorescence bracts. C. Mature plant. D. Flowering branch (Venter 13847). Photos: S. Venter (A–D). mid-point of the corolla tube. Dracophyllum mackeeanum is similar to D. ramosum, but lacks the marked difference between the leaves of the vegetative branch and those of the flowering branch (Table 6). Virot (1975) confused D. mackeeanum with D. ramosum and thought it to be just another variation in branching, leaf and inflorescence characters, not realising the stability of these characters. Dracophyllum mackeeanum is a species having some variation in the leaf size (27.8–58.6
3.8–7.8 mm) and with 22–40 teeth per 10 mm on the lamina margin. Basal Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 165ºE 37 167ºE Capitale/capital city 20ºS Mer de Corail Coral Sea Ouvéa Lifou Nouvelle Calédonie New Caledonia Île sL oy au té s/L oy alt yI sla nd s Maré 22ºS 0 Nouméa 100 km Île des Pins © Australian National University CartoGIS CAP 00-032 Fig. 29. Known distribution of Dracophyllum mackeeanum, New Caledonia. Table 6. Major morphological differences between Dracophyllum mackeeanum and D. ramosum Character D. mackeeanum one or two plants in a population have flowers with sepals equalling the corolla tube in length (MacKee 3480). D. ramosum Lamina size (mm) Inflorescence axis Basal inflorescence branches (mm) Inflorescence-bract length (mm) Inflorescence-bract adaxial surface 28–59
4–8 65–125
6–11 Ribbed Smooth 5–12 0.8–1.0 15–23 (10–)22–70 Pubescent Glabrous at apex Inflorescence-bract length 14.5–23.0 22–70 Number of flowers or inflorescence 25–33 65–350 Abaxial surface of sepal Pubescent Glabrous Corolla-tube diameter (mm) 1.2–1.3 2.3–2.5 Corolla-lobe size (mm) 2.0–2.7
1.2–1.5 1.5–1.7
1.0–1.1 Stamen fusion Lower third Upper third of tube of tube Filament length 1.0–1.5 1.5–1.7 Nectary-scale apices Retuse Irregularly toothed Seed length (mm) 0.75–0.8 0.48–0.5 inflorescence branches can vary from 5 to 12 mm long and the inflorescence-bract size is 15–23
10–15 mm. Occasionally, Selected specimens NEW CALEDONIA. Province Sud: Montagnes de la Dombea, Vieillard 2831 (NSW); 5 miles [~8 km] north-west of Thio on road to Canala, 25 July 1952, McMillan 5159 (L); Combui Concession SLN, 18 Dec. 2002, Dagostini & Rigault 575 (NOU); Thio-Canala, Vallée de la Nekuitourou, 29 Dec. 1977, Bamps 6099 (NOU); valley of the Dothio River, ~12 km north-west of Thio, 17 Dec. 1973, Webster 19349 (NOU); route de PortBouquet, 10 km from Thio, 6 Dec. 1965, MacKee 14017 (NOU); Thio area, ~2 km upstream along the To De River, 13 May 2005, Venter 13847 (NOU); upper Tontouta Valley, hillside, 20 Nov. 1955, MacKee 3480 (L); Col de Vulcain, 24 Sep. 1951, Baumann-Bodenheim 15540 (Z); Bord de la Tontouta, mine Liliane, 15 Dec. 1964, Blanchon 1297 (NOU); Mount Humboldt, 17 May 2005, Venter 13854 (NOU). Col de Ouirange, 14 May 2005, Venter 13852 (NOU). Dracophyllum macranthum E.A.Br. & N.Streiber, Telopea 8 (3): 394 (1999) Type: Australia. New South Wales, North Coast, Coorabakh National Park, track to Newbeys Cave ~100 m from Newbeys 38 Australian Systematic Botany S. Venter Creek Road, on Newbeys Creek, 270 m, 12 Aug. 1997. E.A. Brown 97/51, N. Streiber & D.M. Crayn (holo: NSW 411514!; iso: CHR!, MEL!, NY). Etymology Illustration Diagnostic features and discussion E. A. Brown & N. Streiber Telopea 8 (3): t. 1 (1999). Shrub to small tree 0.6–2.0(–3.0) m tall. Branches frequently pendant. Bark on old branches grey, deeply fissured, young stems reddish-brown. Leaves spirally arranged along young branches but crowded at tips of old branches, spreading; lamina sheath light brown, 7–8
10–12 mm, membranous, striate, tapering and the top half ciliate; lamina coriaceous, abaxial side lighter green, linear–triangular, (85–)140–200
6–9 mm, surfaces glabrous, prominently striated; margins serrulate with 8–13 teeth per 10 mm; apex acute. Inflorescence overtopping the leaves, erect, dense, 100–160 mm long, oblong and sparsely branched; flowers maturing basipetally; rachis and pedicels glabrous; inflorescence axis reddish-brown, 1.0–1.5 mm in diameter; basal inflorescence branch 0.5–1.0 mm long, suberect; inflorescence bracts caducous, overtopping flowers, pink-tipped to pink, broadly ovate at the base, 40–50
8–11 mm, surfaces glabrous, margins ciliate. Flowers 15–38, in groups of 3 at the base of the inflorescence, pedicellate; bracteoles persistent, recaulescent with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, 5–8
0.45–0.5 mm, glabrous; pedicels straight, 1–3 mm long, glabrous. Sepals rose-coloured, ovate–lanceolate, (6–)9–10 (–11)
(1.2–)2.0–2.5 mm, shorter than corolla tube, striate, surfaces glabrous, margins ciliate in upper half. Corolla light to dark pink, becoming red with age; corolla tube cylindrical, widened at mouth, (10–)18–22(–25)
(2–)3.0–3.5(–4) mm; corolla lobes spreading, ovate to ovate–triangular, shorter than corolla tube, 2.2–4.0
1.2–3.0 mm, apices obtuse; adaxial surface mostly papillate, sometimes minutely rugose–verrucate. Stamens hypogynous, sometimes loosely adnated to the wall of the corolla tube, filaments (12–) 18–20 mm long; anthers included, oblong, pink and 2–3 mm long. Ovary cylindrical, 2–3
1–2 mm, glabrous, apex round; nectary scales rectangular to oblong, 1.0–1.5
0.5–0.6 mm, apices irregularly toothed, sometimes retuse; style enclosed, (14–)17–19 mm long, sometimes distally papillose; stigma 5-lobed. Fruit light to reddish-brown, 2.5–4.5
2–3 mm, oblong, apex round, glabrous. Seeds light brown, ovoid, 0.8–1.0 mm long, testa prominently reticulate (Fig. 30, 31). Dracophyllum macranthum has the autapomorphy of flowers maturing basipetally. The corolla tube is pink to dark pink with white corolla lobes. The corolla tube is 18–22 mm long and it is also longer than the sepals. In the past, D. macranthum has been confused with D. secundum and is similar in general appearance. It differs in the leaves being wider (6–9 mm v. 4–6) with fewer teeth (8–13 per 10 mm v. 18–27 per 10 mm) on the lamina margin. The sepals are ovate–lanceolate not broadly ovate triangular and are larger (9–10
2.0–2.5 mm v. 4–6
1.1–1.8 mm in D. secundum). The mouth of the corolla tube is widened and the corolla tube (18–20 mm compared with 4–8 mm in D. secundum) and the corolla lobes (2.2–4.0 mm) are longer. The adaxial surfaces of the corolla lobes are glabrous and the filaments are longer. The nectary scales are larger (1.0–1.5
0.5–0.6 mm). The fruit is also wider (2–3 mm) and the seed longer (0.8–1.0 mm). There is some variation in leaf size (140–200
6–9 mm) and in inflorescence length (100–160 mm). Flower number per inflorescence can vary from 15–38 and minor variation in flower size (18–22
3.0–3.5 mm), but variation in corolla lobe size is more prominent (2.2–4.0
1.2–3.0 mm). The corolla lobes can sometimes be white. Distribution and ecology Endemic to the Coorabakh National Park in New South Wales, Australia (Fig. 32). Dracophyllum macranthum occurs along forested stream gullies, low cliffs and on rock outcrops and faces at an elevation of 800–900 m. The annual rainfall varies between 1200 and 1600 mm. The soil is a sandy loam or clay loam derived from conglomerate and sandstone. Phenology Flowering August–October. The specific epithet describes the large flowers of this species, the largest in the genus. Selected specimens AUSTRALIA. New South Wales: North Coast, Starrs Creek Catchment, south of Big Nellie, 13 Aug. 1997, Brown 97/59, Streiber & Crayn (NSW). Dracophyllum menziesii Hook.f., Fl. Nov. Zel. 2(1): 168 (1853) Type: New Zealand. Dusky Sound, 1791. A. Menzies s.n. (lecto: K!; isolecto: BM 577674!), designated by Oliver (1952). Illustrations W. R. B. Oliver, Trans. Proc. N.Z. Inst. 59: t. 14 (1928); A. Eagle, Trees & Shrubs of N.Z. 2nd series.: t. 132 (1982) – the habit specimen drawn is atypical; J. C. Smith–Dodsworth, N.Z. Native Shrubs & Climbers: t. 60, Pl. 24C & 24D (1991). Shrub 0.5–1.0 m tall. Branches: bark on old branches grey, smooth, rarely deeply fissured at the base, young stems brown. Leaves crowded at tips of branches in a bromelioid manner; lamina sheath light brown, 10–20
(7–)17.2–20.4 mm, coriaceous, striate; tapering, margins membranous and smooth; lamina coriaceous, linear–triangular to occasionally triangular, 90–220
9–17 mm, surfaces glabrous, prominently striated; margins cartilaginous, serrate, thickened, with 20–32 teeth per 10 mm; apex thickened, acute. Inflorescence an axillary panicle situated below the leaves; shorter than the leaves, drooping, dense, 50–150 mm long, oblong and sparsely branched; rachis and pedicels pubescent; inflorescence axis mid-green to reddishbrown, 1.5–2.4 mm in diameter; basal inflorescence branch 11.0–12.4 mm long, widely spreading; inflorescence bracts Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 39 F C D A E B Fig. 30. Dracophyllum macranthum. A. Flowering branch (1
). B. Inflorescence bract (1
). C. Leaf (1
). D. Lower branch of inflorescence (1
). E. Flower (5
). F. Ovary (10
). Drawn from Brown 97/50. Del. S. Venter. caducous, overtopping the flowers, dark green to red, broadly ovate at the base, 12.0–19.2
6.0–8.8 mm, adaxial surfaces glabrous to sericeous in basal half; abaxial surfaces glabrous, margins ciliate. Flowers 8–38, in groups of 3 at the base of the inflorescence, pedicellate; bracteoles caducous, recaulescent, with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, 4–5
0.8–1.0 mm, glabrous; pedicels straight, 1.5–5.5 mm long, tomentose. Sepals ovate to broadly ovate, (2.5–)3.0–3.5
(1.5–)2.0–2.5 mm, shorter than the corolla tube, striate, surfaces glabrous; margins ciliate. Corolla white to red; corolla tube campanulate, widened at mouth, (4–)6–7
3–5 mm; corolla lobes reflexed, ovate–triangular, shorter than corolla tube, 2.0–4.5
1.1–1.6 mm, apices obtuse; surfaces glabrous. Stamens inserted at top of corolla tube, filaments (0.5–)1.0–1.2 mm long; anthers included, oblong, light yellow and 1.3–1.5 mm long. Ovary obovoid, 1.3–1.5
1.7–2.0 mm, glabrous, apex round; nectary scales rectangular, 40 Australian Systematic Botany S. Venter B A C D E E Fig. 31. Dracophyllum macranthum. A. Fruiting plant in habitat. B. Seedlings on a road cutting. C. Mature plant in habitat, Newby’s Creek. D. Flowering branch showing the long corolla tubes and light-coloured corolla lobes. E. Inflorescences showing the flowers maturing from the apices of the inflorescences. F. Fruits showing the persistent long styles. Photos: Barry Ralley (A, B, D–F) and S. Wagstaff (C). 0.7–0.8 mm long and wide, apices retuse; style included, 2.5–3.5 mm long, glabrous, lengthening in fruit; stigma 5-lobed. Fruit reddish-brown, 1.5–2.5
(2.5–)4.0–5.0 mm, depressed-globose, apex round, glabrous. Seeds dark brown, ovoid, (0.55–)0.8–0.6(–1.3) mm long, testa slightly reticulate (Fig. 33, 34). Distribution and ecology Endemic to the South Island of New Zealand and occurs west of a line from Haast to Invercargill and Halfmoon Bay on Stewart Island (Fig. 35). Dracophyllum menziesii grows from close to sea level up to 1500-m elevation on mountain slopes of Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 41 Fig. 32. Known distribution of Dracophyllum macranthum. 10–50 and on Stewart Island occasionally on cliffs, also on bluffs and rock outcrops. Vegetation consists of montane to subalpine woodland, montane and subalpine shrubland, grassland and herb field or snow-tussock herb field. Dracophyllum menziesii is not recorded from forest communities and is the only true grassland species in the genus. The soil is brownish-grey gritty sandy loam derived from granidiorite, schist, argillite, diorite gneiss or occasionally peridotite. All populations visited were recorded as growing fully exposed and covered in a thick layer of snow during the winter. Phenology Flowering November–February. 42 Australian Systematic Botany S. Venter D A E B F C H G Fig. 33. Dracophyllum menziesii. A. Flowering branch (0.5
). B. Leaf (1
). C. Bottom branch of inflorescence (2
). D. Ovary (10
). E. Sepal (5
). F. Flower (5
). G. Inflorescence bract (2
). H. Laid-out corolla (5
). Drawn from Venter 13788. Del. S. Venter. Etymology Named after Archibald Menzies (1754–1842), Scottish surgeon and naturalist. Diagnostic features and discussion Dracophyllum menziesii is characterised by its shrubby habit, mostly unbranched stems, leaves crowded in a bromelioid fashion, lamina short and broad (90–220
9–17 mm). Dracophyllum menziesii is a well defined species, readily distinguished from other Dracophyllum species. It is similar to D. townsonii, but is easily distinguished by the unbranched stems and the shorter leaves. The distribution of the two species does not overlap. Bark on mature stems can vary from smooth to deeply fissured and the lamina shape is very variable, from Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 43 A B C D E Fig. 34. Dracophyllum menziesii. A. Habitat Milford Sound, Fiordland. B. Plant showing the red-coloured winter leaves (Venter 13788). C. Mature plant showing the multi-stemmed growth habit, Fiordland. D. Mature plant showing the inflorescences situated below the leaves, E. Fruiting branch (Venter 13788). Photos: S. Venter (B, E), David Noble (C) and L. and A. Stridvall (D). linear–triangular to triangular in a single population. Leaf size is also variable (90–220
9–17 mm). Plants in subalpine habitats display short inflorescences (50–78 mm), compared with inflorescences from plants growing at lower altitudes (80–150 mm long). The inflorescence bracts vary in size (12.0–19.2
6.0–8.8 mm), with several populations on Stewart Island having the inflorescence bracts red, not the normal green. The corolla can either be white with pink lobes, light pink or red depending on the population. Corolla lobes of subalpine plants are larger than those from lower altitudes. 44 Australian Systematic Botany S. Venter 170º 175º 180º –35º –40º –45º Fig. 35. Known distribution of Dracophyllum menziesii, South Island, New Zealand. Selected specimens NEW ZEALAND. South Island: South Westland, Okuru District, Woolsack, 1 Apr. 1979, Williams & Wardle s.n. (CHR); South Westland, Mataketake Range, Mount Clarke, 7 Mar. 1980, Wardle s.n. (CHR); head of Clarke River, Kea Cliffs, 9 Mar. 1978, Wardle s.n. (CHR); Copland Range, Ngataus Knob, 8 Dec. 1967, Wardle & Fryer s.n. (CHR); Karangarua River, between Cassel and Lame Duck Flat, 31 Mar. 1969, Wardle & Fryer s.n. (CHR); Westland National Park, Fox Range, McKenna Creek, 22 Mar. 1970, Wardle s.n. (CHR); Ben Ohau Range, head of Fred Stream, 8 June 1969, Wilson 456 (CHR); Fiordland, Tutoko Valley, Tutoko high bench, Leader Creek, 13 Jan. 1976, Morris s.n. (CHR); Fiordland, Homer Saddle, 6 Mar. 1962, Simpson 3874 (CHR); South Westland, Gorge Plateau area, Red Hills, Fiordland, north end of Olivine Range, 5 Jan. 1977, Webb 77016 (CHR); Fiordland National Park, Lake Harris, 21 Mar. 2000, Venter 13809 (CHR); Lake Harris, 27 Feb. 1927 Petrie s.n. (CHR); south Canterbury, Hopkins River, head of North Temple Stream, 7 Jan. 1986, Metcalf s.n. (CHR); Fiordland, Doubtful Sound, Secretary Island, Blanket Bay, 13 Dec. 1962, Morice s.n. (CHR); Dusky Sound, Woodhen Cove, 8 Dec. 1946, Allan s.n. (CHR); Fiordland National Park, near West Cape, 5 Feb. 1972, Mark s.n. (CHR); Fiordland, valley south of Mount Anderson, Feb. 1972, Given 72650 (CHR): Fiordland, Dusky Sound, Cascade Valley, 2 Jan. 1969, Dorizac s.n. (CHR); Caswell Sound, above Leslie Clearing, Expectation Stream, 8 Mar. 1949, Murie s.n. (CHR); Fiordland, Mount Gorge, above hut at head of Elizabeth Burn, Mar. 1977, Garnock-Jones, Lee, Anderson & Given 10308 (CHR); mountains head of Lake Hauroko between Hay River and Hauroko Burn, 11 Jan. 1926, Thomson s.n. (CHR); Takitimu Range, head of Aparima River, 31 Jan. 1962, Wardle s.n. (CHR); Preservation Inlet, Useless Bay, 27 Jan. 1946, Allan s.n. (CHR). Stewart Island; Mount Anglem, near top, 11 Jan. 2000, Venter 13788 (CHR). Taxonomic revision of Dracophyllum and Richea Dracophyllum milliganii Hook., Icon. Pl. 9: t. 845 (1852), as ‘Milligani’ Australian Systematic Botany A 45 F D Type: Tasmania, Mount Sorel, Macquarie Harbour, 15 Jan. 1846. J. M. Milligan 747 (holo: MELB 2064389!; iso: BM!; HO!; K!; NSW!; W!). Illustrations J. D. Hooker, Icon. Plant. 9: t. 845 (1851); W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 20 (1928). A shrub 0.2–3 m tall. Branches: bark on old stems brown, deeply fissured, young stems reddish-brown. Leaves spreading to recurved; lamina sheath light green to light brown, (24–) 30–40
(9–)15–20 mm, coriaceous, striate, tapering, top half of margin ciliate; lamina coriaceous, linear–triangular to lanceolate, 140–300(–900)
5.1–7.0 mm, with scabrid hairs restricted to veins on abaxial surface, slightly striated; margins serrulate with 50–70 teeth per 10 mm; apex acute, coils often. Inflorescence overtopping the leaves, erect, dense, 130–460 mm long, oblong and densely branched; rachis and pedicels pubescent; inflorescence axis mid-green, reddishbrown to reddish, prominently ribbed, 3.8–8.2 mm in diameter; basal inflorescence branch 14–25(–101) mm long, widely spreading; inflorescence bracts persistent for a long period, overtopping flowers, light green, pink or with only the apices pink, ovate to broadly ovate at the base, 40–180(–320)
6–19; adaxial surface scabrid at apex; abaxial surfaces scabrid on veins to pubescent, margins serrulate, ciliate. Flowers 150–500+, in groups of more than 10 at the base of the inflorescence, pedicellate; bracteoles persistent, recaulescent with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, 2.0–2.5
0.1–0.2 mm, glabrous; pedicels straight, 0.5–1.5 mm long, tomentose. Sepals reddish-brown to rose coloured, lanceolate to oblong, (2–)3–4
0.8–1.5 mm, equalling corolla tube, striate, surfaces glabrous; margins ciliate in upper half. Corolla white to light pink; corolla tube narrowly campanulate, narrowed at mouth, (2–)2.5–3.0
1.5–2.0 mm; corolla lobes spreading horizontally to reflexed, oblong to ovate–triangular, shorter than corolla tube, 1.0–1.5
(0.9–)1.0–1.2 mm, apices obtuse; surfaces glabrous. Stamens hypogynous, filaments 2.5–5.0 mm long; anthers exserted, oblong, papillate, pink turning deep yellow with age and (0.5–)0.8–1.0 mm long. Ovary subglobose, 1.3–1.5
1.5–2.0 mm, glabrous, apex round; nectary scales appear to be connate, rectangular, 0.2–0.25
0.6–0.8 mm, apices retuse; style included, 0.9–1.0(–1.1) mm long, glabrous; stigma 5-lobed. Fruit reddish-brown, 2.2–2.5
1.5–2.0 mm, depressed-globose, apex round, glabrous. Seeds yellowish-brown, ovoid, 0.47–0.5 mm long, testa slightly reticulated (Fig. 36, 37). Distribution and ecology Endemic to Tasmania with most known localities in the Cradle Mountain–Lake Saint Clair National Park in the north, Walls of Jerusalem National Park, Franklin–Gordon Wild Rivers National Park, Mount Field National Park and the Southwest National Park in the south (Fig. 38). Most common in G B H E C Fig. 36. Dracophyllum milliganii. A. Flowering branch (0.5
). B. Inflorescence-bract adaxial surface (1
). C. Laid-out corolla (5
). D. Leaf (1
). E. Lower inflorescence branch (1
). F. Ovary (10
). G. Sepal (5
). H. Flower (5
). Drawn from Croft 10070. Del. S. Venter. the quartzite mountains of the west and south-west. Dracophyllum milliganii occupies a wide altitudinal range, from almost near sea level in the humid western cold lowlands to 1050-m elevation on Mount Field West. It grows in mountain gullies or on mountain slopes and ridges that are covered in shrubland, moorland, heathland or sedge lands, with the shrub form on the edges of open montane temperate rainforest. The soil is usually shallow yellowish-brown to brown stony clay loam, low in nutrients, and derived from Precambrian or Ordovician quartzite and quartzitic conglomerates. It occasionally occurs on granite and dolerite. Lithosols and shallow fibrous peats cover the glacially and periglacially eroded areas occupied by the alpine vegetation (Kirkpatrick 1983). Phenology Flowering January–May. 46 Australian Systematic Botany S. Venter A B C D Fig. 37. Dracophyllum milliganii. A. Flowering specimens in a grassland habitat on the Sentinel Range. B. Flowering specimen from the quartzite of the Sentinel Range. C. Inflorescence showing clearly the persistent inflorescence bracts, Mount Eliza. D. Plant in fruit, Derwent Valley. Photos: James Wood (A, B), J. Harrison (C) and Natalie Tapson (D). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 47 Fig. 38. Known distribution of Dracophyllum milliganii. Etymology Named after J. S. Milligan (1835–1911) surgeon-superintendent and naturalist who collected plant specimens for W. J. Hooker. Diagnostic features and discussion Dracophyllum milliganii is characterised by its unbranched stems, leaves 140–300 mm long with thin drawn-out apices and finely crenulated lamina margins, panicle 130–460 mm long, having the rachis prominently ribbed and hairy, inflorescence bracts broadly sheathing and abruptly contracted into long and attenuated apices, sepals and corolla tube the same length, anthers exserted and the bilobed apices of the nectary glands. Dracophyllum milliganii is, to some extent, similar to D. involucratum, but easily distinguished from the latter on the account of the flowers not in involucres and the inflorescence bracts that are persistent till the flowers are fully open. Dracophyllum milliganii differs from other Dracophyllum species in the inflorescence bracts that persist deep into the flowering period. There are two different forms. The common form is a small plant, with subterranean stem growing in exposed sites and the other, a tall plant with a stem up to 48 Australian Systematic Botany 3 m tall, long leaves and a more robust inflorescence. This large form is found growing in more protected areas in dense shrub or in forest. There are specimens of intermediates linking the two forms and there are no differences between the two forms in the flowers and inflorescence except that the inflorescence is larger in the shrub form. Dracophyllum milliganii has leaves that vary in size, being 140–300
5–10 mm, depending on the locality and habitat. The colour of the inflorescence axis varies from green, reddish-brown to red. Plants growing in open subalpine areas have short inflorescence bracts (40–90 mm compared with 98–180 mm). Flower colour varies from white to pink with the white forms being sometimes intermingled with the pink. The sepals are variable in shape and size, ranging from lanceolate to oblong and 3–4
0.8–1.5 mm. Corolla lobes are oblong to ovate–triangular, with the filaments 2.5–5.0 mm long. Selected specimens AUSTRALIA. Tasmania: north of Lake Huntley, 23 Sep. 1985, Brown 1261 (HO); Strahan, Jan. 1913, Rodway s.n. (HO); Queenstown, Mount Sedgewick, 28 Feb. 1930, Comber 2162 (K); Macquarie Harbour, Mount Sorell, Von Mueller s.n. (W); Frenchman’s Cap, Barron Pass, 29 Dec. 1966, Olsen 37 (NSW); Twelve Trees Range, 1 km north-east of Strathgordon, 19 Feb. 1989, Croft 10224 (HO, NSW); Clear Hill, 3 Dec. 1997, Buchanan 14990 (HO); Mount Cullen, Summit area, 13 Apr. 1986, Moscal 12874 (HO, NSW); Mount Humboldt, Mount Field West, 25 Dec. 1896, Rodway s.n. (HO); Western Arthur Range, Moraine A, 7 Dec. 1986, Collier 1999 (HO); South West National Park, Schnells Ridge, 11 Jan. 1998, Rozefelds 658 (HO); Port Davy, Cox Bight, near Kings Mine, 15 Dec. 1939, King s.n. (HO); between Mount La Perouse and Maxwell Ridge, 21 Mar. 1984, Buchanan 3025 (HO, NSW); Mount La Perouse, Dec. 1926, Rodway s.n. (HO); ridge between Mount Brady and Mount La Perouse, Dec. 1898, Rodway s.n. (NSW). Dracophyllum oceanicum E.A.Br. & N.Streiber, Telopea. 8 (3): 397 (1999) S. Venter spreading; inflorescence bracts caducous, overtopping flowers, whitish at the base and pink-tipped to pink, ovate–triangular at the base, 10–65
1.5–15.0 mm, surfaces glabrous, margins entire to serrulate. Flowers 300–700+, in groups of more than 10 at the base of the inflorescence, pedicellate; bracteoles persistent, recaulescent with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, linear to triangular, 4–8
0.2–0.6 mm, glabrous; pedicels straight, 1.5–2.0 mm, glabrous. Sepals green to rose-coloured, lanceolate to ovate–lanceolate, 4.5–7.0
1–2 mm, shorter to equalling corolla tube, striate, surfaces glabrous; margins ciliate in upper half. Corolla white; corolla tube narrowly campanulate, widened at mouth, 4–7
2.5–3.5 mm; corolla lobes spreading horizontally to reflexed, ovate, shorter than corolla tube, 1.5–2.5
(1–)1.5–2.0 mm; apices obtuse; adaxial surface papillate and rugose. Stamens inserted in the middle of the corolla tube, filaments 3.0–4.5 mm long; anthers included, rectangular, pink and 1.5–2.0 mm long. Ovary obovoid, 1.5–3.0
1–2 mm, glabrous, apex round; nectary scales oblong, 1.8–2.0
0.5–1.5 (–2.0) mm, apex subacute to obtuse; style included, 2–3 mm long, glabrous; stigma 5-lobed. Fruit dark brown, 1.5–3.0
1.5–2.5 mm, obovoid, apex round, glabrous. Seeds yellowish-brown, trigonous, 0.5–0.6 mm long, testa prominently reticulate (Fig. 39, 40). Distribution and ecology Australian endemic restricted to the Jervis Bay area in New South Wales (Fig. 41). Dracophyllum oceanicum grows in full sun to light shade at a low elevation (10–50 m), on or at the bases of coastal cliffs and along stream margins in sheltered bays. It rarely grows more than 50 m from the sea. The surrounding vegetation is open heathland to dry sclerophyll woodland. The soil is sand or sandy loam derived from sandstone. Type: Australia. New South Wales, South Coast, cliffs ~300 m north of Point Perpendicular, Beecroft Peninsula, Jervis Bay, 9 Nov. 1997. E.A. Brown 97/80, N. Streiber & C.C. Simpson (holo: NSW 412483!; iso: BRI; CANB!; CHR!; NY!). Phenology Illustration The specific epithet describes the habitat as plants grow on seaward cliffs and slopes. E. A. Brown and N. Schreiber, Telopea. 8 (3): t. 2 (1999). A shrub 0.4–2.0 (–2.5) m tall with a scrambling habit. Branches erect–spreading and many branched. Bark on old branches grey, deeply fissured, young stems reddish-brown. Leaves crowded at tips of branches, spirally arranged along stem in young plants, spreading to recurved; lamina sheath olive-green to light brown, 12–17
14–21 mm, coriaceous, striate, membranous, tapering to rounded and the top half minutely ciliate; lamina coriaceous, adaxial surface lightercoloured, linear–triangular, (90–)150–200(–230)
(9–)10–15 (–18) mm, surfaces glabrous, prominently striated; margins serrulate with 10–20 teeth per 10 mm; apex thickened, obtuse. Inflorescence shorter than the leaves, erect, dense, 70–100 mm long, oblong and densely branched; rachis and pedicels glabrous, reddish-brown; inflorescence axis 3.0–3.6 mm in diameter; basal inflorescence branch 0.5–1.0 mm long, widely Flowers from August–December. Etymology Diagnostic features and discussion Dracophyllum oceanicum is characterised by the scrambling growth habit, roughly fissured bark, leaves mostly bronzed green and 150–200
10–15 mm, primary-axis of the inflorescence reddish-brown with flowers on the basal inflorescence branches arranged in groups of 20–30, pedicels 1.5–2.0 mm long, sepals shorter or equalling the cylindrical corolla tube, obtuse corolla lobes, hypogynous filaments and the trigonous seed. Dracophyllum oceanicum shows affinities with D. fitzgeraldii and D. sayeri in the many flowers per inflorescence node, wide coriaceous leaves and floral measurements showing overlap (Streiber et al. 1999). The stigma is capitate and the corolla lobes are of a similar shape in both D. oceanicum and D. fitzgeraldii. Dracophyllum Taxonomic revision of Dracophyllum and Richea C A Australian Systematic Botany D E 49 84, Streiber & Simpson (AK, CANB, NSW); ibid., 22 Oct. 1994, Buchanan 13818 (HO); ibid., 7 Sep. 1997, Brown 97/87, Streiber & Simpson (HO, NSW); ibid., 13 Sep. 1930, Rodway 49 (HO); Beecroft Peninsula, ~500 m W of Beecroft Head, at Eves Ravine, 27 Aug. 1991, Lyne 374 (BISH, NSW, P); 1/4 mile N of Point Perpendicular, (first gully), northern head of Jervis Bay, 7 Oct. 1960, Constable s.n. (HO, K, NSW); Beecroft Peninsula, Gumgetters Inlet, 9 Oct. 1997, Brown 97/83, Streiber & Simpson (NSW); eastern end of Steamers Bush, Brown 97/ 90, Streiber & Simpson, 7 Oct. 1997 (NSW); Drum and Drumsticks, near Point Perpendicular, Jervis Bay, 23 Oct. 1932, Rodway 917 (HO, K, NSW). Dracophyllum ouaiemense Virot, Fl. Nouv.-Calédonie et Dépend. 6: 156 (1975) Type: New Caledonia. Roche Ouaième (massif de Ton-Non), 900 m, 7 Jan. 1968. H.S. MacKee 18230 (P, holo.!; iso: NOU,! P!.). Illustration F B Fig. 39. Dracophyllum oceanicum. A. Flowering branch (0.25
). B. Laidout corolla (5
). C. Leaf (1
). D. Ovary (10
). E. Flower (5
). F. Inflorescence bract (1
). Drawn from Brown 97/82. Del. S. Venter. oceanicum and D. sayeri have glabrous peduncles and pedicels, whereas they are hairy in D. fitzgeraldii (Brown and Streiber 1999). Leaf size varies, being 150–200
10–15 mm, and the lamina might be either flat or channelled in cross-section. The inflorescence varies in length from 70 to 100 mm, with the green inflorescence bracts varying in size (10–65
1.5–15 mm), having a whitish base or it can be pink-tipped or wholly pink. Plants in a population occasionally have the margin of the inflorescence bracts entire, not serrulate. Bracteoles may vary in shape from linear to triangular on the same inflorescence. Sepal size is 4.5–7.0
1–2 mm and the corolla tube 4–7
2.5–3.5 mm, with the corolla lobes being 1.5–2.5
1.5–2.0 mm. The adaxial surface of the corolla lobes can either be papillate or slightly rugose–verrucate. Ovary size varies, being 1.5–3.0
1–2 mm, and fruit size is 1.5–3.0
1.5–2.5 mm. Selected specimens AUSTRALIA. New South Wales: South Coast, Beecroft Peninsula, 9 Sep. 1997, Brown 97/82, Streiber & Simpson (AK, HO, NSW), ibid. Brown 97/ R. Virot, Fl. Nouv.-Calédonie et Dépend. 6: t. (1975). A shrub 30–50 cm tall. Branches erect-stemmed and sparsely branched. Bark on old branches grey to brown, deeply fissured, young stems reddish-brown. Leaves erect to spreading; lamina sheath light green, 5–6
6–8 mm, coriaceous, striate, membranous, tapering and margin ciliate; lamina coriaceous, linear–triangular to triangular, 15–60
4–12 mm, surfaces glabrous, slightly striated; margins denticulate with 20–30 teeth per 10 mm; apex thickened, acute. Inflorescence overtopping the leaves, erect, lax, 20–100 mm long, oblong and sparsely branched; rachis and pedicels pubescent; inflorescence axis reddish, 0.8–2.0 mm in diameter at the base; basal inflorescence branch 2–3 mm long, widely spreading; inflorescence bracts caducous, longer than flowers, light green to reddish, ovate–lanceolate to ovate, 6–12
4–8 mm, surfaces glabrous, margins ciliate, apices obtuse. Flowers 20–35, in groups of 3 at the base of the inflorescence, pedicellate; bracteoles caducous, both bracteoles shorter than the perianth and situated in the middle of the pedicel, 1.8–2.0
1.0–1.3, abaxial surface pubescent; pedicels curved, (2–) 3–11, pubescent. Sepals rose-coloured to red, ovate to broadly ovate, (3.5–)4–6
2.0–2.5 mm, shorter than the corolla tube, striate, adaxial surfaces glabrous, abaxial surfaces with minute scattered papillae; margins ciliate. Corolla red; corolla tube narrowly campanulate to suburceolate, narrowed at mouth, 7–9
0.8–1.0 mm, exterior apically pubescent; corolla lobes spreading, ovate, shorter than corolla tube, 2.3–2.5
1.2–1.5 mm, apices obtuse; adaxial surface papillate; abaxial surface pubescent. Stamens implanted at the middle of the corolla tube, filaments 1–2 mm long; anthers included, rectangular, 0.7–1.5 mm long. Ovary globose, 0.8–1.0 mm long and wide, glabrous, apex round; nectary scales rectangular, 0.4–0.5 mm long and wide, apex retuse to irregularly toothed; style included, 0.7–1.0 mm long, glabrous; stigma capitate. Fruit reddish-brown, 1.2–1.5
2–4 mm, obovoid to broadly obovoid, apex round, glabrous. Seeds light brown, ovoid, 0.55–0.6 mm long testa slightly reticulate (Fig. 42, 43). 50 Australian Systematic Botany S. Venter A B C D E Fig. 40. Dracophyllum oceanicum. A. Plants growing on the cliffs at Jervis Bay. B. Mature plant showing the scrambling habit, Picnic Point. C. Old inflorescence showing the dropped inflorescence bracts, Picnic Point. D. Plant with mature fruit, Jervis Bay. E. Young inflorescence showing the inflorescence bracts. Photos: S. Wagstaff (A, D), Cas Liber (B, C) and M. Fagg, ANBG (E). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 51 Fig. 41. Known distribution of Dracophyllum oceanicum. Distribution and ecology New Caledonia endemic restricted to the Roche Ouaième area of Grande Terre in the north (Fig. 44). Dracophyllum ouaiemense occurs in full sun on mountain tops and gentle to steep (10–40) slopes at 850–900-m elevation. The vegetation is typical maquis and D. ouaiemense grows in close association with Xeronema moorei on clay loam derived from schist. This habitat is usually moist during the rainy season and receives mist throughout the year, especially over the summer months. Phenology Flowering January–July. Etymology Named after the Roche Ouaième area where the species occurs. Diagnostic features and discussion Dracophyllum ouaiemense is characterised by its low (up to 50 cm tall) growth habit, thick stems with the leaves 15–60
4–12 mm and glabrous, short inflorescence (20–100 mm long) normally carried above the leaves, 7–9-mm-long flowers arranged in groups of 3 at the base of the inflorescence, 3–11 mm long filiform pedicels, flexuose and caducous filiform bracteoles, glabrous sepals 4–6 mm long, red suburceolate 7–9-mm-long corolla tube, obtuse oval-shaped corolla lobes and the included stamens. Virot (1975) placed D. ouaiemense close to D. alticola and D. cosmelioides, but it differs from them mainly in the inflorescence characters. Dracophyllum ouaiemense is superficially similar to D. balansae and D. cosmelioides, but differs from them in the glabrous branchlets, wider lamina (4–12 mm compared with 0.8–3 mm) and denticulate lamina margin with 20–30 52 Australian Systematic Botany S. Venter E A D inflorescence branches. The apices of the nectary scales are normally retuse, but occasionally they are irregularly toothed. Selected specimens NEW CALEDONIA. Province Nord: Roche Ouaième (massif de Ton-Non), 13 July 1968, MacKee 19135 (P); ibid., 7 Jan. 2003, Tronchet 714 (MO, NOU, P). F Dracophyllum ramosum Pancher ex Brongn. & Gris., Ann. Sci. Nat., Bot. sér 5(2): 156 (1864) Type: New Caledonia. Montagne de M’bée, 1861. E. Vieillard 830 (lecto: L!; isolecto: BR, G, GH, GH, NSW, W!, Z!). New Caledonia. Kanala, J.A.I. Pancher s.n. (syn: L!). Dracophyllum amabile Brongn. & Gris, Bull. Soc. Bot. France 11: 68 (1864). C Type: New Caledonia. Montagnes de Kanala, 1855, E. Vieillard 829 (lecto: P!; iso: K!, L!, LY!, W!, Z!.), designated by Oliver (1952). Dracophyllum vieillardii Lenorm. ex Guillaumin, Ann. Mus. Colon. Marseille 9 (2): 181 (1911). nom. nud. Type: New Caledonia. Dombéa, E. Vieillard s.n. (holo: P!; iso: P!, W!, Z!). B H Illustrations G Fig. 42. Dracophyllum ouaiemense. A. Flowering branch (1
). B. Laidout corolla (5
). C. Bottom inflorescence branch (2
). D. Leaf (1
). E. Ovary (10
). F. Flower (5
). G. Sepal abaxial surface (5
). H. Inflorescence bract (5
). Drawn from MacKee 18230. teeth per 10 mm (compared with 50–100 teeth per 10 mm). The rachis is pubescent, with the basal branches of the inflorescence being longer (2–3 mm compared with 0.5–1 mm). The inflorescence bracts have obtuse apices and the pedicel is much longer (3–11 mm compared with 0.5–2.5 mm). The sepals are shorter than the corolla tube and the corolla tube is longer (7–9 mm compared with 3–6 mm) with the outer surface pubescent compared to glabrous. The corolla lobes are longer (2.3–2.5 mm compared with 0.8–1.8 mm) with pubescent abaxial surfaces, and the stamens are inserted in the middle of the corolla tube, with the ovoid seed being longer (0.5–0.6 mm compared with 0.2–0.3 mm). The leaves of D. ouaiemense show variation in shape from linear–triangular to triangular and in size (15–60
4–12 mm). The inflorescence bracts vary from ovate–lanceolate to ovate at the base and also in size (6–12
4–8 mm). The flowers at the base of the inflorescence are sometimes arranged in groups of two, a rare reduction from the normal groups of three on the basal W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 17 & 18 (1928); W. R. B. Oliver, Trans. & Proc. Roy. Soc. N.Z. 80 (1): t. 17, 18 (1952); R. Virot, Fl. Nov. Calédonie et Dépend. 6: t. 25 (1975) p.p. A shrub or a small tree, 0.5–5.0 m tall. Branches spreading and sparsely branched, with side branches growing from the same point. Bark on old branches greyish to blackish-brown, finely to deeply fissured, young stems reddish-brown. Leaves spirally arranged on young plants, spreading to recurved, decreasing in size below inflorescence; lamina sheath light green to light brown, 19–28
10–14 mm, coriaceous, striate, membranous, tapering and margin smooth; lamina coriaceous, linear–triangular to lanceolate, 65–125
6–11 mm, surfaces glabrous, prominently striated; margins serrulate with 40–60 teeth per 10 mm; apex obtuse to acute. Inflorescence overtopping the leaves, erect, dense, 40–200 mm long, linear–oblong and sparsely branched; rachis and pedicels pubescent; inflorescence axis light to mid-green, 1–2 mm in diameter; basal inflorescence branch 0.8–1.0 mm long, widely spreading; inflorescence bracts caducous, overtopping flowers, coriaceous, pink-tipped, pink or red, narrowly ovate to ovate–triangular at the base, (10–)22–70
10–15 mm, surfaces glabrous, margins ciliate. Flowers 70–250+, in groups of 5–10 at the base of the inflorescence, pedicellate; bracteoles persistent, both bracteoles situated at a basal position on the pedicel, with the top bracteole longer than the perianth and the lower shorter than or equalling the flower, linear, 3–8
0.6–1.0 mm, glabrous; pedicels straight, reddishbrown, 0.5–1.5 mm, tomentose. Sepals green to rose-coloured, ovate–lanceolate to ovate, (3.0–)3.5–4.0
(1.3–)1.5–1.8, Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A B C E 53 D E Fig. 43. Dracophyllum ouaiemense. A. Habitat on Roche Ouaième at the type locality. B. Flowering branch showing the urceolate pink flowers and the pink sepals. C. Adult flowering plant along the Ouaième River. D. Inflorescence showing the flower bracts. E. Plants growing among the shrub vegetation. F. Vegetation on Roche Ouaième. Photos: T. Whitaker (A, E, F), Gildas Gateblé (B) and J. Munzinger (C, D). shorter than or equalling the corolla tube, striate, surfaces glabrous; margins ciliate; apices acute. Corolla white to dark pink; corolla tube cylindrical, narrowed at mouth (2.5–) 4.0–6.0
2.3–2.5 mm, exterior pubescent in upper part; corolla lobes spreading to horizontal, oblong to ovate– triangular, shorter than corolla tube, 1.5–1.7
1.0–1.1 mm, apices obtuse with an erose margin, inflexed at apices; adaxial surface papillate, abaxial surface glabrous to pubescent. Stamens inserted in the upper-third of the corolla tube, filaments 1.5–1.7 mm long; anthers included, oblong, light yellow and 0.5–0.7(–1.0) mm long. Ovary subglobose to globose, 0.8–1.0
1.1–1.3 mm, glabrous, apex round; nectary scales rectangular, 0.9–1.0
0.4–0.5 mm, apex irregularly toothed; style included, 0.7–0.8 mm long, glabrous; stigma 5-lobed. Fruit reddish brown, 1.4–1.5
1.5–3.0 mm, obovoid to round, apex round, glabrous. Seeds light brown, filiform, 0.48–0.5 mm long, testa slightly reticulate (Fig. 45, 46). Distribution and ecology New Caledonia endemic, growing on the main island Grande Terre, and on Ile des Pins in the south (Fig. 47). Dracophyllum ramosum is the most common Dracophyllum species in New 54 Australian Systematic Botany S. Venter 165ºE 167ºE Capitale/capital city 20ºS Mer de Corail Coral Sea Ouvéa Lifou Nouvelle Calédonie New Caledonia Île sL oy au té s/L oy alt yI sla nd s Maré 22ºS 0 100 km Nouméa Île des Pins © Australian National University CartoGIS CAP 00-032 Fig. 44. Known distribution of Dracophyllum ouaiemense, New Caledonia. Caledonia. Dracophyllum ramosum grows on mountain summits and slopes, in river valleys, on plateaux and commonly along streams from 100- to 1200-m elevation. The vegetation varies from forest, maquis to shrub mosaic. Soils are rocky loam or clay loam, mostly ferruginous and derived from peridotite or serpentinite. Phenology Flowering September–February. Etymology Named for the many branches developing in the same place, a diagnostic character of the species. Diagnostic features and discussion Dracophyllum ramosum is characterised by the side branches that arise from the same point, leaves that decrease in size below the inflorescence, inflorescence being longer than the leaves, flowers arranged in groups of 5–10 on the basal inflorescence branches, pubescent outer surface of the corolla and the inflexed apices of the corolla lobes. When Brongniart and Gris (1864) published the Pancher manuscript name D. ramosum, they mentioned two specimens, namely, Vieillard 830 collected at M’bée and Pancher s.n. collected at Kanala. Oliver (1928) gave M’bée as the type locality, but he did not designate a specimen, and in a later publication (1952) he selected the Vieillard 830 (L) specimen as lectotype. However, Virot (1975) rejected the Vieillard 830 specimen as lectotype, and correctly chose the Pancher specimen from Kanala, because the name ‘Dracophyllum ramosum’ occurs on the label in Pancher’s writing. Dracophyllum ramosum is similar to D. fitzgeraldii but differs in leaf, inflorescence and flower characters. Dracophyllum ramosum is a polymorphic species, with a great deal of variation in the leaves and inflorescence (Virot 1975). It grows as a small shrub scarcely 1 m tall at Mount Tchingou to a forest element of 5 m at Mount Dore. Branches are spreading and mostly sparsely branched or might be many branched, or it may have many branches all from one point. These variations appear to be the result of environmental conditions. Leaves vary in size (65–125
6–11 mm), sometimes having obtuse lamina apices. Inflorescences vary, being 40–200 mm long, with the inflorescence bracts being narrowly ovate to ovate–triangular at the base and 22–70
Taxonomic revision of Dracophyllum and Richea B Australian Systematic Botany C D A E 55 Mé Maoya, 28 Dec. 1962, MacKee 9894 (K, L, NOU, NSW); Bogota, Sarasin 283 (Z). Province Sud: Dothio, 21 Oct. 1965, MacKee 13670 (L, NOU); Mé Ori, 26 May 1965, MacKee 12695 (L, NOU); Canala, Mondi, 24 Nov. 1970, MacKee 22571 (K, L, NOU, Z); Baie des Pirogues, 4 Feb. 1926, Däniker 3073 (Z); Prony, 15 Oct. 1914, Franc 1868 (Z); Thio, Jan. 1872, Balansa 3671 (K); Baie d’Uié, base du Pic Ia, 17 Sep. 1868, Balansa 277 (Z); Ngoye, 26 Nov. 1902, Schlechter 15093 (BM, W, Z); northern Dumbea Valley, 5 Feb. 1956, MacKee 3974 (K, L); Montagne des Sources, 29 Sep. 1979, McPherson 1921 (NOU, NSW); Col de Ouirange, 14 May 2005, Venter 13851 (NOU); De Touaourou ad Mamié, 26 July 1965, Bernardi 9812 (K, L); Vallee N’Go, Baie du Sud, Jan. 1903, Cribs 1267 (K); Mount Dore area, along the Lembi River, 12 May 2005, Venter 13843 (NOU); Vallée de la Rivière des Lacs, Pont, 20 Nov. 1969, Guillamin & Baumann 6619 (K); Ile des Pins, Germain s.n. (Z). Dracophyllum sayeri F.Muell., Australas. J. Pharm. 2: 85 (1887) F Dracophyllum sayeri var. sayeri F.M.Bailey, Compr. Cat. Queensl. Plants: 296 (1913). Type: Australia. Northern Queensland, on Mount BellendenKer, at elevations between 4500 and 5200 feet [~1371–1584 m]; W. Sayer s.n. (holo: MEL 2064424!). G Dracophyllum sayeri var. reflexum F.M.Bailey, Compr. Cat. Queensl. Plants: 296 (1913). Mentioned in a note. Dracophyllum sayeri var. normale F.M.Bailey, Compr. Cat. Queensl. Plants: 296 (1913). [Intended to be type variety]. Illustrations Fig. 45. Dracophyllum ramosum. A. Flowering branch (1
). B. Leaf (1
). C. Inflorescence bract (1
). D. Ovary (10
). E. Flower (5
). F. Sepal (5
). G. Bottom branch of inflorescence (1
). Drawn from MacKee 22571. Del. S. Venter. 10–15 mm. The sepals vary from ovate–lanceolate to ovate, with the corolla being white to dark pink, the corolla lobes oblong to ovate–triangular, sometimes glabrous on the abaxial surface in certain populations, but these are not geographically separated. Because most of the above-mentioned variation could occur in a small area, it is not practical to depict this on a pictorial map. Selected specimens NEW CALEDONIA. Province Nord: Mount Tiebaghi, 21 Oct. 1959, Thorne 28058 (L, Z); Pouembout Valley, 3 July 1967 MacKee 17024 (L, NOU, Z); Taom, Mount Homédéboa, 16 Oct. 1969, MacKee 20999 (K, L, NOU); Mount Kaala, southern slopes, 20 Oct. 1956, MacKee 5546 (K, L); Mount Tchingou, 21 Aug. 1965, Bernardi 10426 (L, S, Z); Kopéto, east of Mount Vert, 8 Aug. 1970, MacKee 22213 (L, NOU); Moneo, 8 Oct. 1973, MacKee 27565 (K, L); Mount Mé Maoyo, Mine Emma, 11 July 1965, Bernardi 9557 (L, Z); Haute Houailou and Haute Baraoua, Spur of F. M. Bailey, Compr. Cat. Queensl. Plants. t. 269 (1913); K. Domin, Bibliotheca Botanica. 89(4): t. 173 (1928); W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 19 (1928). A shrub or tree 2–8 m tall. Branches sparsely to many branched, sometimes forming a tangled mass. Bark on old branches light brown, flaky, young stems yellowish-brown. Leaves recurved in a bromelioid manner; lamina sheath light brown to pinkish-red, 20–27
14–16 mm, coriaceous, striate, tapering and margin ciliate; lamina coriaceous, adaxial surface lighter green; linear–triangular, (240–)310–440
9.0–11.5 (–20) mm, surfaces glabrous, prominently striated; margins entire; apex acute. Inflorescence shorter than leaves, erect, dense, 160–210 mm long, linear–oblong and densely branched; rachis and pedicels glabrous to minutely hirsute; inflorescence axis light to mid-green, 2.8–6.8 mm in diameter; basal inflorescence branch 15–28 mm long, widely spreading; inflorescence bracts caducous, overtopping flowers, whitish at the base to pink-tipped or wholly pink, ovate–lanceolate at the base, 85–110
15–25 mm, surfaces glabrous, margins entire. Flowers 300–500+, in groups up to 20 at the base of the inflorescence, pedicellate; bracteoles caducous, recaulescent, with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, equalling the flower, 4–5
0.5–1.0 mm, glabrous; pedicels straight, 1.5–2.5 mm long, glabrous to pubescent. Sepals green to rose-coloured, lanceolate to ovate–lanceolate, 2.3–3.2
1.0–1.5 mm, shorter than the corolla tube, striate, surfaces glabrous; margins ciliate. Corolla white turning cream- 56 Australian Systematic Botany S. Venter B A C D E Fig. 46. Dracophyllum ramosum. A. Mature plant growing, Lac Pernod. B. Maquis habitat near Mount Dore. C. Flowering branch, Dumbea (Venter 13851). D. Mature plant showing the characteristic multiple branching from a single point, Mont Moné. E. Young inflorescences showing the broad and early deciduous flower bracts, Tiebaghi. Photos: S. Venter (A, B), S. Wagstaff (C), Gildas Gateblé (D) and T. Whitaker (E). coloured with age, to light pink, turning deeper pink with age or white with pink lobes; corolla tube narrowly campanulate, widened at mouth, 3–5
2–3 mm; corolla lobes reflexed, oblong to ovate, shorter than corolla tube, 2.5–3.8
1.5–2.0 mm, apices obtuse; adaxial surface papillate, abaxial surface glabrous. Stamens inserted at the top of the corolla tube, filaments 1.0–1.3 mm long; anthers exserted, rectangular, light yellow and 1.0–1.5 mm long. Ovary Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 165ºE 57 167ºE Capitale/capital city 20ºS Mer de Corail Coral Sea Ouvéa Lifou Nouvelle Calédonie New Caledonia Île s Lo ya uté s/L oy alt yI sla nd s Maré 22ºS 0 100 km Nouméa Île des Pins © Australian National University CartoGIS CAP 00-032 Fig. 47. Known distribution of Dracophyllum ramosum, New Caledonia. subglobose, 0.7–1.0
1.0–1.2 mm, glabrous, apex round; nectary scales connate at the base, rectangular, 0.4–0.5 mm long and wide, apices retuse to irregularly lobed; style included, 1.0–2.2 mm long; stigma 5-lobed. Fruit pedicellate, included in persistent calyx, reddish-brown, 1.7–2.0
2.5–3.0 mm, depressed-obovoid; apex round, glabrous. Seeds light brown, ovoid, 0.95–1.0 mm long, testa slightly reticulate (Fig. 48, 49). Phenology Flowering sometimes starts during April; however, the main flowering occurs from June to August. Etymology Named after W. A. Sayer, plant collector for Baron von Mueller who collected the type material. Diagnostic features and discussion Distribution and ecology Australian endemic, restricted to the Bellenden-Ker and Bartle Frere Mountain Ranges in the Wooroonooran National Park, with an isolated record from the eastern slopes of Mount Spurgeon near Mossman, north-eastern Queensland (Fig. 50). Dracophyllum sayeri is common in the understorey of well developed low microphyll vine–fern rainforest and less so in stunted rain forest on high ridges and slopes at elevations of 1371–1730 m. The soil is skeletal and derived from granite. Plants of D. sayeri grow mainly in light shade, but there are populations in which all the plants are in full sun, having shorter and thicker textured leaves. Dracophyllum sayeri is characterised by the panicles that are shorter than the leaves and having up to 20 flowers per basal inflorescence branch, inflorescence bracts tapered at both ends and 85–110 mm long, sepals much shorter than the corolla, corolla lobes 2.5–3.8 mm long with obtuse apices, exserted anthers and club-shaped stigmas. The Sayer specimen (MEL 2064424) is the only specimen mentioned in the protologue and Oliver (1952) incorrectly stated that the Johnson s.n. (MEL) specimen collected on Mount Bartle Frere is the type. Dracophyllum sayeri is similar to D. fitzgeraldii and D. verticillatum, but differs in leaf, inflorescence and flower characters. The only record of birds feeding on D. sayeri nectar 58 C Australian Systematic Botany S. Venter 1.5–2.0 mm. The nectary-scale apices vary from retuse to irregularly toothed, with the latter being most common and the style measures 1.0–2.2 mm long. These variants are present in all the known populations. D A E Selected specimens F AUSTRALIA. Queensland: Cairns, summit of Mount Bellenden-Ker, 16 Sep. 1977, Powell 817 (BRI, CBG, HO, NE, Z); ibid. 27 July 1990, Baylis & Wardle s.n. (CHR); Mount Bellenden-Ker, South Peak, 2 Aug. 1971, Van Balgooy 1455 (L); ibid., summit, 6 Apr. 1984, Brass 18313 (L); ibid., 1892, Podanza s.n. (BM); Mount Bellenden-Ker, Centre Peak, 7 June 1972, Wrigley & Telford 837 (L); ibid., south-west of Centre Peak, 7 June 1969, Smith 14615 (L); ibid., Centre Peak, near TV Tower, 9 Nov. 1972, Webb & Tracey 10805 (L); Mount Bellenden-Ker, on ridge between cableway and North Peak, 3 Sep. 1986, Clarkson 6566 (MBA, L, NSW); ibid., 8 Apr. 2006, Venter 13860 (CHR, BRI); Queensland, Mount Bartle Frere, 1892, Johnson s.n. (MEL); ibid., 11 June 1949, Smith 4212 (L); 3 km north of Mossman Gorge, plane-crash site, 27 Nov. 1990, Monteith s.n. (BRI); runction of Roots Creek and Mossman Falls, Sep. 1936, White 10604 (BRI). Dracophyllum secundum R.Br. ex Roem. & Schultes., Syst. Veg. 4: 385 (1819) Type: Australia. Eastern New South Wales, Port Jackson. 18 July 1802. R. Brown s.n. [Bennett 2,805] (lecto: MEL 2064414!; isolecto: BM 577, 617!, BM 577,619!, K000349838, P00760781, P00760783), designated by Oliver (1952). B G Epacris secunda Poiret, nom. illeg. In: Lamarck, Encycl. Méth. Bot. Suppl. 2: 556 (1811). Prionotes secunda (R.Br.) Sprengel, Syst. Veg. 1: 631 (1825). Dracophyllum secundum R.Br. forma secundum Domin, Bibliotheca Botanica. 89: 501 (1928). Autonym. Dracophyllum secundum R.Br. forma rubreo–rosea Domin, Bibliotheca Botanica. 89: 501 (1928). Type: Australia. New South Wales, Sandsteinfelsen der Blue Mountains. Apr. 1897. Domin IV (holo: PR?, n.v.). Fig. 48. Dracophyllum sayeri. A. Flowering branch (1
). B. Laid-out flower (5
). C. Leaf (0.5
) with enlargement of lamina margin (5
). D. Flower (5
). E. Ovary (10
). F. Inflorescence bract (1
). G. Sepal (5
). Drawn from Powell 817. Del. S. Venter. is that recorded by Van Balgooy (CANB 330631) on Mount Bellenden-Ker. Two varieties recognised by Bailey (1913) cannot be upheld, because the erect-stemmed plants with white corolla tubes and those with pink corolla lobes (=D. sayeri var. normale) grade into plants with more scandent stems, allwhite flowers and strongly reflexed corolla lobes (=D. sayeri var. reflexum). Some variation occurs in the size of the lamina (310–440
9.0–11.5 mm) and in inflorescence length (160–210 mm). The inflorescence axis varies from 2.8 to 6.8 mm in diameter at the base and the inflorescence bracts can vary from subulate to ovate–lanceolate and measure 85–110
15–25 mm in a single population (Clarkson 6566). Pedicels are either glabrous or pubescent and the corolla lobes are oblong to ovate, measuring 2.5–3.8
Illustrations Hooker, Curtis’s Bot Mag. 7: t. 3264 (1833); Engler and Prantl, Die Natürlichen Pflanzenfamilien 4(1): t. 38. A subshrub to shrub, (0.1–)0.2–1.0(–2.0) m tall. Branches spreading. Bark on old branches grey to brown, deep and broadly fissured, young stems reddish-brown. Leaves crowded at tips of branches on mature stems, spirally arranged on young branches; spreading to recurved, decreasing in size below inflorescence; lamina sheath light green to light brown, 5–17
5–18 mm, subcoriaceous, striate, membranous, tapering to rounded and the top half ciliate; lamina coriaceous, glaucous to dark green and glossy, abaxial surface lighter coloured, linear–triangular, (30–)50–160 (–180)
(1.3–)4–6(–18) mm, glabrous, prominently striated; margins serrulate with 18–27(–40) teeth per 10 mm; apex acute. Inflorescence overtopping the leaves, erect, dense, 80–140 mm long, linear–oblong and sparsely branched; rachis and pedicels glabrous; inflorescence axis light green to reddish, 1.2–1.8 mm in diameter; basal Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A B C D E 59 F Fig. 49. Dracophyllum sayeri. A. Plant in habitat. B. Rooting of stems where they touch the ground. C. Large inflorescence bracts. D. Flowers and young fruit. E. Mature inflorescence. F. Adult plant on Mount Bellenden-Ker. Photos: S. Venter (A–F). inflorescence branch 0.5–1.0 mm long, widely spreading; inflorescence bracts caducous, overtopping flowers, light green, pink-tipped to wholly pink, ovate to broadly ovate at the base, 35–105
10–20 mm, surfaces glabrous, margins ciliate. Flowers 24–67, in groups of 5–10 at the base of the inflorescence, pedicellate; bracteoles persistent, shorter than the perianth, both bracteoles situated at a basal position on the pedicel, 2.0–4.7
0.4–1.1 mm, glabrous, a few teeth at apices; pedicels straight, (0.5–)2.0–3.5 mm long, glabrous. Sepals green to rose-coloured, broadly ovate to triangular, (3–)4–6 (–7)
(0.9–)1.1–1.8(–2.2) mm, shorter than the corolla tube, striate, surfaces glabrous; margins ciliate in the upper half. Corolla white to deep pink, becoming light red with age; corolla tube cylindrical to narrowly campanulate, narrowed at 60 Australian Systematic Botany S. Venter Fig. 50. Known distribution of Dracophyllum sayeri. mouth, 4–8(–10)
(1.5–)2–3(–3.5) mm; corolla lobes reflexed, ovate–triangular, shorter than corolla tube, 1–2 (–3) mm long and wide, apices obtuse; adaxial surface papillate; abaxial surface glabrous. Stamens hypogynous, filaments 4–8 mm long with the filaments sometimes adhering to the corolla-tube wall with the free part 1–2 mm long; anthers included, rectangular, purple turning light yellow with age and (0.5–)1.2–2.0 mm long. Ovary pale pink, obovoid, 1.5–3.0
1–2 mm, glabrous, apex round; nectary scales rectangular, 0.8–1.0
0.3–0.5 mm, apex obtuse to irregularly toothed; style pale pink, included, 3–4 mm long and papillose distally; stigma 5-lobed. Fruit light brown, 3.5–5.0 (0.5–)1.2–2.0
1.3–1.6 mm, round to oblong, apex round, glabrous. Seeds light brown, ovoid 0.7–0.75 mm long with a prominently reticulated testa (Fig. 51, 52). Distribution and habitat Australian endemic restricted to the Central and Southern Tablelands in eastern New South Wales (Fig. 53). Dracophyllum secundum occurs at elevations of 165–960 m on cliffs, in gorges, valleys, and occasionally along riverbeds. The vegetation consists of open forest, dry woodland or heathland. The soils are sandy to loamy lithosols derived from conglomerate, sandstone or, occasionally, shale. Dracophyllum secundum grows mostly in full sun but some plants grow in light shade and have larger and darker green leaves. Phenology Flowering June–December. Etymology The specific epithet describes the flowers borne in a ‘secund’ or single-sided inflorescence. Diagnostic features and notes Dracophyllum secundum is characterised by having prominently striated leaves, flowers moderately large and on long pedicels, frequently arranged to one side of the inflorescence (hence, the specific epithet), flowers arranged in groups of 5–10 on the lower inflorescence branches, sepals much shorter than the corolla tube, which is narrowed at the mouth, spreading long Taxonomic revision of Dracophyllum and Richea E Australian Systematic Botany A 61 F D G H C B I Fig. 51. Dracophyllum secundum. A. Flowering branch (0.5
). B. Lower inflorescence branch with only three of the five flowers shown (2
). C. Laidout corolla (2
). D. Leaf (1
). E. Lamina margin (40
). F. Ovary (10
). G. Flower (2
). H. Sepal (2
). I. Inflorescence bract (1
). Drawn from Powell 341. corolla lobes and the distally papillose styles. Roemer and Schultes (1819), Hooker (1844) and Mueller (1864) gave the type locality as the Port Jackson area but without any collector details. Oliver (1952) was the first to designate a type with collector data and he chose the R. Brown [Bennett 2,805] (MELB 2064414) specimen. A polymorphic species with no main stem and variously branched from the base. The main variation is in the lamina size (50–160
4–6 mm), inflorescence length (80–140 mm) and inflorescence bract size (34–105
10–20 mm). Inflorescences can have from 24 to 67 flowers. Flower colour varies from near white to deep pink, with the corolla tube cylindrical to narrowly campanulate. Mueller (1864), Bentham (1869) and Brown and Streiber (1999) described the stamens as adnate to the corolla tube and also mentioned that most of the flowers studied had the filaments adnate to the corolla tube, whereas the illustration in Curtis’s Botanical Magazine (Hooker 1833) shows the stamens as hypogynous. Filaments appear to be connate to the inside of the corolla; however, they are actually covered with a single cell-thick membrane. This membrane readily breaks up so that the filaments easily come away from the corolla tube in the dry state and also when there is excess movement of the flowers in the fresh state. Nectary-scale apices vary from obtuse to irregularly toothed and the ovary also varies in size (1.5–3.0
1–2 mm). Selected specimens AUSTRALIA. New South Wales: escarpment below Waihou Trig., 25 km north-west of Coffs Harbour, 12 Oct. 1978, Streimaw 8135 (A, L, BISH); Rylstone, Currant Mountain east of Gap, 7 Aug. 1975, Coveny & Hind 6608 (NSW, W); 2 km from Dunns Swamp turnoff, towards Rylstone, 2 Dec. 62 Australian Systematic Botany S. Venter A B C D E Fig. 52. Dracophyllum secundum. A. Mature inflorescence with dropped inflorescence bracts from the Blue Mountains. B. Mature plant showing the spreading habit from the Blue Mountains. C. Cliff habitat, Blue Mountains. D. Inflorescence showing the characteristic large inflorescence bracts at the apex of the inflorescence, Blue Mountains. E. Inflorescence of the pink colour form, from near the National Pass, Blue Mountains. Photos: Terry Wright (Black Diamond Images) (A, B, D), Tony Rodd (C) and David Noble (E). 1987, Rimes 51 (NSW); Central Tablelands, Kelgoola 40 km east of Rylstone, Baker s.n. (NSW); 3 km east of Nerriga, 16 Sep. 1965, Pullen 4114 (AD, B, BISH, BRI, E, FI, G, K, L, LE, MEL, MO, P, WELT, Z); Central Tablelands, near Lithgow water supply, Clarence, 21 Oct. 1939, Blakely s.n. (NSW); Milton, Pointer Gap, 9 Aug. 1978, Barnsley 214 (A, HO, L, US); Central Tablelands, Blackheath, 6 Sep. 1936, Vickery s.n. (NSW); Medlow Bath below Lake Medlow, 18 May 1977, Coveny 9448 & Telford (K, L); Springwood, Oct. 1981, Mayrhofer s.n. (GZU); Woy Woy Creek, 24 Aug. 1969, McBarron 17515 (NSW); Central Coast, The Quarry, Hornsby District, Sep. 1971, Lassak s.n. (NSW); Central Coast, Patonga, Sep. 1938, Lilier s.n. (NSW); Central Coast, Warah, 23 Apr. 1949, Oxenford s.n. (NSW); Central Coast, Oatley, Aug. 1904, Boorman s.n. (NSW); Central Tablelands, northern spur of Mount Colong, 5 miles [8 km] west-south-west of Yerranderie, 1 Oct. 1951, Johnson s.n. (NSW); near Bullio, 21 Dec. 1975, Thompson 2372 (NSW); 2 km north of Mittagong, Gibbergunyah Creek, 16 Nov. 1982, Cooper & Powrie s.n. (NSW); Kellys Falls, 31 July 1997, Brown 97/21b, Streiber, Jobson & Taafe (HO); Central Coast, Little River, Buxton, 30 Sep. 1951, Whaite 1072 (NSW); Wollongong, Telniek 137 (W); Central Taxonomic revision of Dracophyllum and Richea Fig. 53. Australian Systematic Botany 63 Known distribution of Dracophyllum secundum. Tablelands, Gow Gully, Penrose, Oct. 1938, Blakely s.n. (NSW); Central Tablelands, Fitzroy Falls, Oct. 1918, Rodway s.n. (NSW); Central Coast, Kangaroo Valley, also Cambewarra Mountain, 28 Sep. 1928 Morris 22442 (NSW); north of Minnamurra Falls, 23 Oct. 1993, Gilmour 7505 (CBG, MEL, NSW); Central Coast, Sydney, Port Hacking, Oct. 1907, Boorman s.n. (L, Z); Southern Tablelands, 2 miles [3.2 km] north-east of Endrick River, Nerriga–Sassafras road, 3 Oct. 1959, Williams 126 (NSW); Wandean Road, 11 Sep. 1997, Brown 97/97 & Streiber (CHR, HO, NSW, NY); eastern slopes of Pigeon House Range above Enrick River, along road from Nerriga to Nowra, 13 Oct. 1965, Hoogland 10040 (L); near Mount Corang, 6 Nov. 1975, Powell 341 (BRI, MEL, NSW); South Coast, Pointer Gap, 9 km north-west of Milton, 9 Aug. 1978, Barnsley 214 (HO, L). Dracophyllum strictum Hook.f., Fl. Antarct. 1: 48 (1844) Type: New Zealand. Mount Tongariro. J.C. Bidwill s.n. (holo: K!). Dracophyllum affine Hook.f., Fl. Antarct. 1: 48 (1844). Type: New Zealand. Northern Island, mountains of the interior. Dr E. Dieffenbach s.n. (holo: K!). Dracophyllum featonianum Colenso, Trans. & Proc. N.Z. Inst. 22: 477 (1890). Type: New Zealand. Cape Runaway (Whangaparoa) a little north of East Cape, 1889. E.H. Featon s.n. (holo: K, n.v.). Dracophyllum imbricatum Colenso, Trans. & Proc. N.Z. Inst. 25: 331 (1893). Type: New Zealand. Open lands near Cape Runaway, 1892. H.T. Hill s.n. (holo: WELT 23623!; iso: K!). Note in Colenso’s hand that the WELT specimen is the type of D. imbricatum. Illustrations W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 16 (1928); W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 16 (1928); A. Eagle, Trees and Shrubs of N.Z.: t. 169 (1975) show the stamens as being adnate to the corolla tube in the lower third instead of in the upper-third and the nectary scales are depicted as overtopping the ovary, which is incorrect; J. SmithDodsworth, N.Z. Native Shrubs & Climbers: tt. 58, 59, pl. 24A, B (1991). A shrub up to small tree, 50–300 cm tall. Branches: bark on old branches dark brown, finely fissured, young stems yellowish-brown. Leaves adult and juvenile; juvenile leaves spirally arranged along branches, spreading, lamina sheath 64 Australian Systematic Botany light green, glaucous to light brown, 14–20
9–16 mm, tapering and margin minutely ciliate in the upper half; lamina coriaceous, glaucous to light green, broadly linear–triangular, 100–140
7–10 mm; surfaces glabrous, margins serrulate with 40–50 teeth per 10 mm; apex acute; adult leaves crowded at tips of branches, spreading; lamina sheath glaucous to light brown, 7–15
6–14 mm, coriaceous, striate, tapering to rounded; margin membranous and minutely ciliate; adult lamina coriaceous, glaucous to light green, lighter coloured below, linear–triangular to lanceolate, 47–75
5–8 mm, surfaces glabrous, slightly striated; margins serrulate with 40–50 teeth per 10 mm; apex thickened, acute. Inflorescence overtopping the leaves, erect, dense, 50–100 mm long, pyramidal and sparingly branched; rachis and pedicels hirsute; inflorescence axis light green, 1.5–1.7 mm in diameter; basal inflorescence branch 0.5–1.0 mm long, widely spreading; inflorescence bracts caducous, overtopping flowers, whitish at the base, pink-tipped to wholly pink, broadly ovate at the base, 7.5–18.0
6.0–8.5 mm; adaxial surfaces with minute scabrid hairs; abaxial surfaces glabrous, margins ciliate. Flowers 15–60, in groups of 5–10 at the base of the inflorescence; bracteoles persistent, recaulescent with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, linear, 3–4
0.3–0.6 mm, glabrous; pedicels green to reddish-brown, straight, 0.6–2.0 mm long, pubescent. Sepals green to rosecoloured, ovate to broadly ovate, (1.7–)2.0–3.0
1.3–1.5 mm, shorter than the corolla tube, striate, surfaces glabrous; margins ciliate. Corolla white to light pink; corolla tube narrowly campanulate, widened at mouth, (3.5–)4.0–6.0
1.5–2.0 mm; corolla lobes reflexed, ovate–triangular to triangular, shorter than corolla tube, (1.5–)2.0–2.5
1.5–2.2 mm, inflexed for the entire length, apices subacute; adaxial surface papillate. Stamens inserted on corolla tube in upper third, filaments 0.5–1.3 mm long; anthers included, oblong, deep yellow and 0.7–0.8 mm long. Ovary obovoid, 1–2 mm long and wide, glabrous, apex round; nectary scales rectangular, 0.8–1.0
0.4–0.5 mm, apices subacute; style included, 1.5–2.0 mm long, glabrous; stigma capitate. Fruit light brown to reddish-brown, 1.8–2.0
1.7–2.0 mm, depressed-globose, apex round, glabrous. Seeds yellowishbrown, ovoid, 0.6–0.7 mm long, testa slightly reticulate (Fig. 54, 55). Distribution and ecology Endemic to the North Island of New Zealand, ranging from Auckland southwards (Fig. 56). Dracophyllum strictum occurs from near sea level to 1097-m elevation on cliffs along riverbanks and on bluffs. The vegetation varies from lowland to montane forest and woodland to shrubland. Soils are usually grey sandy loam derived from sandstone, alluvial material or pumice, or clay derived from ignimbrite, andesite, mudstone or calcareous mudstone. Most plants have been recorded as growing in light shade. Phenology Flowering October–May. S. Venter Etymology The specific epithet describes the closely packed and rigid leaves of this species. Diagnostic features and discussion Dracophyllum strictum is characterised by the presence of juvenile leaves, dense panicles with scabrous short secondary branches, short bracteoles (3–4 mm), small flowers (4–6 mm), short sepals (2–3 mm) and 4–6 mm long corolla. This is the only species in Dracophyllum with juvenile leaves, separating it morphologically from all the other species. Dracophyllum strictum is similar to D. secundum but differs in the juvenile leaves, more teeth (40–50 compared with 18–27) per 10 mm on the lamina margin, hirsute, not glabrous, inflorescence rachis, shorter (7.5–18.0 mm compared with 35–105 mm) inflorescence bracts, pubescent pedicels, not glabrous, shorter sepals (2–3 mm compared with 4–6 mm), corolla tube widened at the mouth not narrowed, longer corolla lobes (2.0–2.5 mm compared with 1–2 mm) having subacute, not obtuse, apices, nectary-scale apices subacute, not obtuse, to irregularly toothed, shorter style (1.5–2.0 mm compared with 3–4 mm) and slightly reticulated testa. There is considerable variation in the size and colour of the juvenile (100–140
7–10 mm) and adult (47–75
5–8 mm) leaves. Plants growing in exposed conditions on cliffs have very short and light grey leaves with unbranched short stems and are in appearance similar to D. secundum. The inflorescence is short (50–68 mm) in plants growing in full sun, compared with the longer (70–100 mm) inflorescences of the shade plants. Inflorescence bracts show a wide range in size (7.5–18.0
6.0–8.5 mm) and the flower number varies from 15 to 60 per inflorescence, with the corolla lobes being ovate–triangular to triangular. Selected specimens NEW ZEALAND. North Island: Whangarei, Maungakaramea, 21 Oct. 1938, Skottsberg s.n. (O); Thames, Puriri, Sep. 1881, Cheeseman s.n. (Z); Tamahere Rapids, Kirk s.n. (AK); Okoroire, Oct. 1896, Cheeseman s.n. (AK); Mayor Island, Opo Bay, eastern side, 29 Nov. 1992, Sykes 281/92 (CHR); Mayor Island, cliffs on northern rim of crater, Nov. 1955, Chambers s.n. (AK); East Cape, near Potaka, Manga-purua Stream, 11 Nov. 1984, Courtney s.n. (CHR); Tawarau State Forest, west of Waitomo Mairoa, 3 Jan. 1984, Ogle 1052 & Haydock (CHR); Urenui, Mount Messinger, 11 Feb. 1999, Venter 13763 (CHR); South Auckland, Atapuni Dam, 8 Sep. 1990, De Lange 582 (AK, CHR); South Auckland, Ngaroma, 25 m south-east of Te Awamutu, 17 June 1982, Bartlett NGA161 (CHR); Lake Whakameru, 7 km north-east of Whakameru village, Highway 30, 13 Mar. 1985, Hind 3978 (NSW); Near Pureora, Waimiha River, 7 Mar. 1978, Gardner 1843 (AK); Rotoroa, Maunga Kakaramea, 21 Oct. 1938, Skottsberg s.n. (O); Lake Taupo, Western Bay, Waihaha Scenic Reserve, 8 Jan. 1985, De Lange s.n. (AK); Mangatapu Stream headwaters, Mamaku Plateau, 5 May 1978, Gardner 1960 (AK, L, NSW); Lake Okataina, 18 Sep. 1943, McKenzie s.n. (AK); Rainbow Mountain, near top, 30 Dec. 1993, Ford 12/94 (CHR); Mount Tarawera, 26 Dec. 1968, Wood s.n. (AK); Wanganui, Waverley, Lake Moumahaki, 4 Feb. 1990, Ogle 1917 (CHR); Raurimu, 12 Nov. 1964, Healy & Brown s.n. (HO); Turangi, Red Hut, 9 Feb. 1999, Venter 13760 (CHR); Kiwitea County, Rangiwahia, 14 May 1961, Esler s.n. (MPN). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 65 B A H C I D G F E Fig. 54. Dracophyllum strictum. A. Flowering branch (0.5
). B. Ovary (10
). C. Flower (5
). D. Inflorescence-bract adaxial surface (2
). E. Sepal adaxial surface (10
). F. Laid-out corolla (5
). G. Lower inflorescence branch (2
). H. Juvenile leaf (1
). I. Adult leaf (1
). Drawn from Gardner 4185. Del. S. Venter. Dracophyllum townsonii Cheeseman, Man. N.Zeal. Fl.: 420 (1906) Type: New Zealand. Paparoa Range, near the base of Mount Buckland. W. Townson s.n. (AK 6924, lecto.!; AK 6923!; AK 6925!; AK 6926!; AK 211642!; AK 211643! isolecto.!), designated by Oliver (1952). Illustrations T. F. Cheeseman, Ill. N.Z. Fl.: t. 130 (1914); A. Eagle, Trees & Shrubs of N.Z. 2nd series: t. 131 (1982). A shrub or small tree 3–6 m tall. Branches form an open candelabrum-shaped crown. Bark on old branches greyishbrown to light brown, flaky, young stems reddish-brown. Leaves crowded at tips of branches in a bromelioid manner, spirally arranged on young stems, young leaves distinctly pinkcoloured; lamina sheath light green, (15–)20–25
8–22 mm, coriaceous, striate, tapering, margins membranous with the top half ciliate; lamina linear–triangular, 130–300
4.5–12.0 mm, surfaces glabrous, prominently striated; margins cartilaginous, serrate with 28–50 teeth per 10 mm. Inflorescence an axillary panicle situated below the leaves; shorter than the leaves, curved and drooping, dense, 40–130 mm long, pyramidal and sparingly 66 Australian Systematic Botany S. Venter B A C D E Fig. 55. Dracophyllum strictum. A. Habitat at the Tongariro River, Turangi. B. Mature plant showing the typical sprawling habit. C. Inflorescence without the inflorescence bracts. D. Inflorescence with some of the flower bracts intact. E. Upper elevation habitat at the Rangiwahia Waterfall. Photos: S. Venter (A), Phil Bendle (B–D) and J. Inger (E). branched; rachis and pedicels hirsute, light green; inflorescence axis, 2.2–4.2 mm in diameter; basal inflorescence branch (10–) 16–24 mm long, suberect; inflorescence bracts caducous, overtopping flowers, whitish in lower half, ovate at the base, 40–50
10–12 mm, glabrous, margins ciliate. Flowers hidden by leaves, 30–90, in groups of 5–10 at the base of the inflorescence, pedicellate; bracteoles persistent, both bracteoles longer than the perianth and situated in the middle of the pedicel, 2.5–7.0
0.3–1.0 mm, glabrous; pedicels 0.2–0.5 mm long, pubescent. Sepals green, broadly ovate, Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 67 180º −35º −40º −45º Fig. 56. Known distribution of Dracophyllum strictum, North Island, New Zealand. 2.5–4.0
1.7–2.5 mm, shorter than corolla tube, striate, surfaces glabrous; margins ciliate in the upper half. Corolla red; corolla tube campanulate, widened at mouth, 2.0–2.5
2.3–2.5 mm; corolla lobes reflexed, oblong, shorter than or equalling corolla tube, 1.9–2.0
1.0–1.2 mm; glabrous; apices obtuse; surfaces glabrous. Stamens inserted in the upper third of corolla tube, filaments 1–2 mm long; anthers exserted, rectangular, light yellow and 1.3–1.5 mm long. Ovary subglobose, 1.3–1.5
2.0–2.5 mm; glabrous, apex round; nectary scales rectangular, 0.4–0.5
0.8–1.0 mm, apex retuse to irregularly toothed; style exserted, 1.5–2.0 mm long, glabrous; stigma capitate. Fruit light brown to reddishbrown, 2–3 mm long and wide, depressed-globose, apex round and glabrous. Seeds light brown, ovoid, 1.1–1.3 mm long, testa slightly reticulate (Fig. 57, 58). Distribution and ecology New Zealand endemic from Whanganui Inlet in the north to Hokitika in the south (Fig. 59). Dracophyllum townsonii occurs on gentle (5–15) southern, south-eastern and southwestern mountain slopes and along drainage lines, at elevations ranging from 152 to 900 m. Dracophyllum townsonii is a montane forest element rarely growing in full sun except where the shading trees have fallen down. Soils are brown clay-loam, brown sandy loam or rocky sandy soils derived from sandstone or conglomerate. Plants of D. townsonii usually occur in permanently moist habitats. Phenology Flowering December–March. 68 Australian Systematic Botany E S. Venter A F G B shorter (2.0–2.5 mm compared with 6–7 mm) with the anthers exserted not included, the ovary subglobose not obovate, shorter style (1.5–2.0 mm compared with 2.5–3.5 mm) and longer seed (1.1–1.3 mm compared with 0.55–0.6 mm). The flowers are reported as having a foetid smell by Cheeseman (1914), but all the flowers I smelled at the localities visited had a sweet scent. Seedlings commonly grow on decaying, mosscovered logs and on tree-fern stems in the forest. Basal branches of mature plants develop roots where they touch the ground and later form individual plants. In D. townsonii, leaves of plants in deep shade are nearly twice as long as those from plants growing in full sun, and with a thinner texture. The number of teeth per 10 mm on the lamina margin varies from 28 to 50 and bracteole size varies (2.5–7.0
0.3–1.0 mm) even on the same inflorescence. The nectary-scale apices are either retuse or irregularly toothed. Selected specimens D C Fig. 57. Dracophyllum townsonii. A. Flowering branch (0.5
). B. Inflorescence bract (5
). C. Sepal abaxial surface (5
). D. Laid open corolla (5
). E. Leaf (1
). F. Ovary (10
). G. Flower (5
). Drawn from Venter13777. Del. S. Venter. Etymology NEW ZEALAND. South Island: Collingwood, Knuckle Hill, 15 Jan. 1999, Venter 13752 (CHR); ibid., Venter 13777 (CHR); Mount Burnett, 30 Apr. 1945, Cone s.n. (CHR); Charleston, 15 Jan. 1967, Moore s.n. (CHR); Four Mile Creek, 12 Jan. 1971, Wardle s.n. (CHR); Buckland Peaks, 24 Mar. 1996, Glenny 6459 (CHR); Karamea, Mossy Burn Catchment, 3 Dec. 1984, McLennan s.n. (CHR); Mount Stormy, Jan. 1987, Druce s.n. (CHR); Greymouth, Ten Mile Creek, Sep. 1964, Robins s.n. (O); Sewell Peak, 22 Nov. 1968, Drury 68W/75 (CHR); Mount Mantell, Allan s.n. (CHR); near Greymouth, Liverpool State Goldmine, 27 Mar. 1915, Morgan s.n. (AK). Dracophyllum traversii Hook.f., Hand. N.Zeal. Fl.: 736 (1864) Type: New Zealand. Middle Island, Nelson Province, Arthur’s top, subalpine, 3000 feet [914 m], Oct. 1865. W.T.L. Travers & J. Haast 337 (holo: K!; iso: CHR 236852!, K!). Named after William Townson (1850–1926), a pharmaceutical chemist and plant collector, especially in the South Island. Dracophyllum pyramidale W.R.B.Oliv., Trans. & Proc. Roy. Soc. N.Z. 80 (1): 16 (1952). Diagnostic features and discussion Type: New Zealand. Summit of Little Barrier Island, 8 Oct. 1928, W.R.B. Oliver s.n. (holo: WELT 48615!). Dracophyllum townsonii is characterised by the open candelabra-shaped crown, crowded leaves at the end of branches growing in a bromelioid manner, distinctly pinkcoloured young leaves, small panicle (40–130 mm long) always situated below the leaves, flowers arranged in groups of 5–10 on the basal inflorescence branches, inflorescence bracts broad and contracted into subulate apices, campanulate corolla, sepals being shorter than the corolla tube, sharply reflexed corolla lobes and the exserted anthers. Dracophyllum townsonii is related to D. fiordense and D. menziesii in the short inflorescence situated below the bromelioid crown of leaves and the campanulate flowers with exserted stamens and style. Dracophyllum townsonii is similar to D. menziesii, but differs in being a small tree with an open candelabra-shaped crown, bark on mature stems flaky, not smooth, much longer inflorescence (50–150 mm compared with 40–50 mm) and the flowers are arranged in groups of 5–10, not three, on the basal inflorescence branches, the bracteole is persistent and equals the flower in length, not being shorter, the corolla tube is Illustrations W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: tt. 22, 23 (1928); L. B. Moore and J. B. Irwin, Oxford Book of N.Z. Plants: 290 (1978); A. Eagle, Trees & Shrubs of N.Z., 2nd series: t. 128 (1982); J. T. Salmon, Native Trees N.Z.: 269, 270 (1989). A shrub or a tree 0.2–13.0 m tall. Branches form an open candelabrum-shaped crown. Bark on old branches light brown, flaky, young stems reddish-brown. Leaves crowded at tips of branches in a bromelioid manner, lamina sheath light green to light brown, 30–65(–70)
30–50 mm, coriaceous, striate, tapering, margins membranous, smooth; lamina coriaceous, sometimes with a glaucous bloom, linear–triangular to lanceolate, 90–300(–860)
(17–)40–50 mm, surfaces glabrous, prominently striated; margins cartilaginous, serrulate with 18–20 teeth per 10 mm. Inflorescence shorter than the leaves, dense, 180–350(–400) mm long, pyramidal and densely branched; rachis and pedicels pubescent to hirsute, light green to reddish; inflorescence axis 13.0–16.5 mm in Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 69 A B C D E Fig. 58. Dracophyllum townsonii. A. Forested habitat along the Karamea River, Kahurangi National Park. B. Mature plant showing the open candelabrum growth habit. C. Fruiting branches of the forest form showing widely spreading branches. D. Fruiting branch showing the infructescence below the head of leaves. E. Typical bromelioid head of leaves of the open vegetation form. (B–D Venter 13752). Photos: Ben (A), S. Venter (B, D), Alice Shanks (C) and Steve Attwood (E). diameter; basal inflorescence branch 30–60 mm long, suberect to at right angles with inflorescence axis; inflorescence bracts caducous, overtopping flowers, light green, whitish at the base and pink-tipped to entirely pink, broadly ovate at the base, 130–240
25–50 mm, surfaces glabrous, margins minutely ciliate. Flowers 500–3000+, in groups of more than 10 at the base of the inflorescence, pedicellate; bracteoles caducous, recaulescent, deciduous, with 1 bracteole situated just below 70 Australian Systematic Botany S. Venter 170º 175º 180º −35º −40º −45º Fig. 59. Known distribution of Dracophyllum townsonii, South Island, New Zealand. the perianth and the other in the middle of the pedicel, shorter than flower, 4.0–4.8
0.5–0.7 mm, glabrous; pedicels straight, 0.5–2.0 mm long, pubescent to tomentose. Sepals red to occasionally green, ovate to broadly ovate, (1.2–)2–3
(1.1–)2.0–2.5 mm, equalling corolla tube, striate, surfaces glabrous; margins ciliate; apices subacute to obtuse. Corolla red, tube sometimes white; corolla tube broadly campanulate, widened at mouth, 2.7–3.0
4–5 mm; corolla lobes reflexed, oblong, longer than corolla tube, 2.5–2.8
2.0–2.5 mm; apices obtuse; surfaces glabrous. Stamens inserted at top of corolla tube, filaments 1.0–1.5 mm long; anthers exserted, oblong, pink turning light yellow with age and 1.8–2.0 mm long. Ovary subglobose, 1.4–1.5
1.8–2.0 mm, glabrous, apex round; nectary scales oblong, 1.0–1.5 mm long and wide, apices retuse; style exserted, 2–3 mm long, glabrous, lengthening in fruit; stigma 5-lobed. Fruit not included in persistent calyx, reddish- to purplish-brown, 1.9–2.0
2.8–3.0 mm, depressed-globose; apex round, glabrous. Seeds yellowish-brown, ovoid, 0.95–1.0 mm long, testa slightly reticulate (Fig. 60, 61). Distribution and ecology New Zealand endemic widely distributed, occurring from the Coromandel Peninsula, Great Barrier Island and Little Barrier Island on the North Island and throughout the South Island (Fig. 62). Dracophyllum traversii occurs on gentle to steep (3–75) slopes from almost sea level to 1768-m elevation. It grows in full sun to light shade on cliffs, in moist gorges, on saddles and mountain slopes. The vegetation consists of Taxonomic revision of Dracophyllum and Richea E A Australian Systematic Botany F 71 Table 7. Differences between Dracophyllum traversii and D. latifolium Character Lamina margin Lamina width (mm) Teeth per 10 mm on lamina margin Flower grouping on inflorescence branch Sepal size (mm) Sepal : corolla-tube length (mm) Sepal adaxial surface Sepal margins G B Corolla-tube size (mm) Corolla-lobe size (mm) Ovary shape Ovary size (mm) Style length (mm) Seed length (mm) D. traversii D. latifolium Serrulate 40–50 18–20 Serrate to denticulate 12–30 2–4 >10 5–10 2–3
2.0–2.5 Equalling Glabrous Ciliate 0.7–1.5
1.0–1.7 Shorter Pubescent Toothed in upper third 1.5–2.0
1.5–2.5 1.5–2.0
1.5–2.0 Ovate 0.8–1.0
1.0–1.5 1.0–1.7 1.2–1.3 2.7–3.0
4–5 2.5–2.8
2.0–2.5 Subglobose 1.4–1.5
1.8–2.0 2–3 0.95–1.0 Diagnostic features and discussion D C Fig. 60. Dracophyllum traversii. A. Flowering branch (0.5
). B. Inflorescence bract (0.5
). C. Sepal (5
). D. Laid-out corolla (5
). E. Leaf (1
). F. Ovary (10
). G. Flower (5
). Drawn from Venter 13734. Del. S. Venter. lowland forest and shrubland or subalpine forest and shrubland. It is common in the small-tree tier of highaltitude conifer–broad-leaved forest and in montane to subalpine beech forest in the western half of the South Island (Sakai and Wardle 1984; Poole 1987). Plants of D. traversii experience snow conditions during the winter, and occasionally during the summer months. The small-tree and shrub tiers in these forests are underdeveloped, with D. traversii being a common element, sometimes forming dense stands. Soils are brown clay to dark brown clay loam derived from sandstone, limestone, greywacke or shale. Phenology Flowering October–February. Dracophyllum traversii is characterised by the candelabra– shaped growth habit, bark flaking in large pieces, broad strongly curved leaves, young leaves with a grey bloom, stout panicle with red flowers having the corolla lobes longer than the corolla tube and capsules 2.8–3.0 mm in diameter. Dracophyllum traversii is closely allied and also similar to D. latifolium (Oliver 1928) but differs in lamina and flower characters (Table 7). A grey to glaucous waxy bloom covers the young leaves, but this is lost at maturity. Haase (1986) found during a survey on the ecology of D. traversii, the forest floor below D. traversii trees to be covered in a layer (up to 100 mm thick) of dead leaves that effectively prevents seedling establishment of arborescent and herbaceous species. Seeds of D. traversii were released from a height of 5 m at a wind speed of ~2 m s–1 and the seeds travelled for distances of up to 10 m (Haase 1986). Dracophyllum traversii plants in exposed areas in the subalpine zone rarely reach 1 m in height. The form of D. traversii that was described as D. pyramidale W.R.B.Oliv. on the North Island is more robust in all parts and occurs in lowland forests. Dracophyllum pyramidale was included in D. traversii by Poole and Adams (1994) and is here also included in D. traversii. Plants growing in the upper edge of the tree line (subalpine forests) exhibit leaves with a distinct grey to glaucous waxy bloom and also show a prominent change in leaf colour during the winter from green to reddish-purple (presence of anthocyanins). Lamina sheaths vary in size (30–65
30–50 mm) and so does the lamina (90–300
40–50 mm). Plants in full sun have shorter (180–221 mm) inflorescences than those in forest habitats (230–350 mm long). Inflorescence bracts also vary in size (130–240
25–50 mm). Etymology Named after William Thomas Locke Travers (1819–1903), lawyer and botanical explorer who made a special study of the flora of Nelson, Marlborough and Canterbury. Selected specimens NEW ZEALAND. North Island: north Tangihua Forest, 29 Aug. 1991, Cameron 6582 (AK); Houto State Forest, summit of Mangatipa No. 2., 6 72 Australian Systematic Botany S. Venter A B C E D F G Fig. 61. Dracophyllum traversii. A. Habitat on the slopes of Mount Arthur. B. Mature plant on Flora Saddle, Mount Arthur. C. Branch with ripe fruit. D. Carpet of dead leaves underneath trees. E. Flowering branch, Coromandel Peninsula. F. Dropped corollas. G. Plant showing the large coloured inflorescence bracts just before they drop. Photos: S. Venter (A–D) and J. Braggins (E). Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 73 180º −35º −40º −45º Fig. 62. Known distribution of Dracophyllum traversii. Sep. 1985, Bellingham 0041 (AK); Little Barrier Island, Summit Track, 9 Mar. 1962, Melville & Hamilton 6676 (K); Coromandel, eastern pinnacle of Moehau, 22 Aug. 1974, Wright 710 (AK, L); Thames, Mount Kaitarakihi, 12 Feb. 1999, Venter 13765 (CHR); Pirongia, Bald Spur, 17 Aug. 1971, May s.n. (AK); Mount Hikurangi, 15 Nov. 1926, Oliver s.n. (WELT 48619); South Island, Collingwood, Knuckle Hill, 28 Mar. 1999, Venter 13778 (CHR); Westport, Mount Rochfort, 25 Nov. 1998, Venter 13732 (CHR); Motueka, Mount Arthur, Flora Saddle, 7 Dec. 1998, Venter 13734 (CHR); Cobb saddle, 22 Feb. 1965, MacFarlane 452 (LINC); Mount Wilberg, 28 Apr. 1993, Wardle & Buxton s.n. (AK, CHR); Otira Gorge, Phillips Turner s.n. (AK); Franz Josef, Alex Knob, 13 Jan. 1966, Wardle s.n. (CHR); Fiordland, Esperance Valley, Mar. 1974, Atkinson s.n. (CHR); Fiordland, Doubtful Sound, Hall Arm, 31 Dec. 1939, Cranwell & Moore s.n. (CHR); Fiordland, Dusky Sound, Pickersgill Harbour to Lake Forester, 3 Jan. 1969, Dorizac s.n. (CHR). Dracophyllum verticillatum Labill., Rel. Voy. Rech. Pérouse 2: 211 (1800) Type: New Caledonia. s. loc. 26 Apr. 1793. M. Labillardière s.n. (lecto: FI!; isolecto: L!, NSW!, P photo!; Herb. Jussieu.), designated by Virot (1975). Dracophyllum dracaenoides Schltr., Engl. Bot. Jahrb. 39: 220 (1906). Type: New Caledonia. Auf den Bergen bei Ou-Hinna, 900 m, 2 Jan. 1903, R. Schlechter 15599 (holo: P!; iso: L!, NSW!, P!, S!, W!, Z!). 74 Australian Systematic Botany Illustrations J. H. Labillardière., Rel. Voy. Rech. Pérouse: t. 40 (1800); A. P. de Candolle, Prodr. Syst. Nat. Reg. Veg. 7: t. 7 (1839); Schlechter, Engl. Bot. Jahrb. 39: t. 21 (1907); Oliver, Trans. & Proc. N. Z. Inst. 59: t. 24 (1928); R. Virot, Fl. Nov. Calédonie et Depend. 6: tt. 19–22 (1975). A shrub or a tree, 0.2–7.0 m tall. Branches: bark on old branches grey–brown, finely fissured, young branches reddishbrown. Leaves crowded at tips of branches in a bromelioid manner, spreading to recurved; lamina sheath light brown, 17.3–25.2
13–17 mm, coriaceous, striate, margins membranous, tapering and margin minutely ciliate; lamina coriaceous, base red, lower surface lighter-coloured, linear–triangular, 60–700
6–40 mm, surfaces glabrous, prominently striated; margins serrate to serrulate with 20–32 teeth per 10 mm; apex obtuse to acute. Inflorescence overtopping the leaves, erect, dense, (120–)190–400(–700) mm long, linear–oblong and sparingly branched; rachis and pedicels glabrous to pubescent, light to mid-green; inflorescence axis (2.5–)4–15 mm in diameter; basal inflorescence branch 0.5–2.0 mm long, widely spreading; inflorescence bracts caducous, overtopping flowers, light green-tipped pink, ovate to broadly ovate at the base, 42–43
17.0–18.3 mm, surfaces glabrous with scattered hairs on abaxial surface, margins ciliate. Flowers 100–500+, in spaced verticillasters, in groups of 5–10 at the base of the inflorescence, pedicellate; bracteoles caducous, recaulescent, deciduous, with 1 bracteole situated just below the perianth and the other in the middle of the pedicel, shorter than flower, (1.0–)2.5–4.0
(–0.1) 0.2–0.3 mm, glabrous; pedicels straight, 1–5 mm long, glabrous to pubescent. Sepals green to rose-coloured, ovate to broadly ovate, 1.7–3.0
1.7–2.0 mm, shorter than corolla tube, striate, surfaces glabrous; margins ciliate in upper half to completely ciliate; apices subacute to obtuse. Corolla white to light pink; corolla tube narrowly campanulate, widened at mouth, (1.5–)2.0–3.0(–5.0)
2.0–2.5 mm, exterior glabrous; corolla lobes reflexed, oblong to ovate–triangular, shorter to equalling corolla tube, (1.2–)2.0–2.5
1.0–2.5 mm, apices obtuse; adaxial surface papillate. Stamens inserted either in the middle or in the upper third of corolla tube, filaments 1.0–1.5 mm long; anthers exserted, oblong, pink to purple and 1–2 mm long. Ovary globose, 0.8–1.0 mm long and wide, glabrous, apex round; nectary scales connate at the base, rectangular, 0.3–0.5
0.4–0.5 mm, apices retuse to irregularly toothed; style included, 1.5–2.0 mm long, glabrous, lengthening in fruit; stigma clavate. Fruit included in persistent calyx, reddish-brown, 1.0–1.2
(1.0–) 1.5–4.0 mm, depressed-globose; apex round, glabrous. Seeds yellowish-brown, ovoid, 0.95–1.0 mm long, testa slightly reticulate (Fig. 63, 64). S. Venter E A F G B D C Fig. 63. Dracophyllum verticillatum. A. Flowering branch (0.25
). B. Inflorescence bract (0.5
). C. Sepal abaxial surface (5
). D. Laid-out corolla (5
). E. Leaf (0.5
). F. Ovary (10
). G. Flower (5
). Drawn from MacKee18864. Del. S. Venter. to clay loam and derived from various rock types (peridotite, gneiss, schist, phtanite, quartzite and chalk). Phenology Flowering September–March. Etymology The specific epithet describes the flowers being borne in verticillasters, a prominent feature of this species. Distribution and ecology Endemic to New Caledonia and widely distributed throughout (Fig. 65). Dracophyllum verticillatum occurs in exposed areas on gentle to moderate (5–25) mountain slopes, in ravines and on plateaux from 40- to 1200-m elevation. The vegetation consists of forest, maquis or shrubland. Soils are mostly loam Diagnostic features and discussion Dracophyllum verticillatum is characterised by slightly serrated lamina margins, broad lamina sheath, a very long spike-like panicle bearing the flowers in clusters (verticils) at close intervals, broad and obtuse sepals, broad and short corolla Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A B C D E F 75 Fig. 64. Dracophyllum verticillatum. A. Plant in habitat near the Gallieni Mine, Tontouta Valley. B. Mature plant with young inflorescences near Thio. C. Inflorescence showing the flowers arranged in verticils. D. Flowering plant near the Gallieni Mine, Tontouta Valley. E. Old fruiting branch, Thio. F. Plant with young inflorescences, Thio. Photos: S. Venter (A–F). tube with recurved corolla lobes and exserted anthers. Labillardière (1800) gave no specific locality and cited no specimen for the plant he collected but stated that he was the collector. Virot (1975) chose the lectotype at Firenze (FI) but gave no further reference. There is marked variation in leaf size (60–700
6–40 mm) and inflorescence length (190–700 mm). The inflorescence axis also varies from 4 to 15 mm in diameter at the base. Pedicels are 1–5 mm long and can either be glabrous or pubescent. Sepal shape (ovate to broadly ovate) and size 76 Australian Systematic Botany S. Venter 165ºE 167ºE Capitale/capital city 20ºS Mer de Corail Coral Sea Ouvéa Lifou Nouvelle Calédonie New Caledonia Île sL oy au té s/L oy alt yI sla nd s Maré 22ºS 0 Nouméa 100 km Île des Pins © Australian National University CartoGIS CAP 00-032 Fig. 65. Known distribution of Dracophyllum verticillatum. (1.7–3.0
1.7–2.0 mm) vary, with the margins either ciliate in the upper half or ciliate for the whole length and having subacute to obtuse apices. Flower colour varies from white to deep pink, sometimes in a single population. Corolla lobes vary in shape (oblong to ovate–triangular) and size (2.0–2.5
1.0–2.5 mm) and, on the rare occasion, are shorter than the corolla tube. Nectary-scale apices are either retuse or irregularly toothed. The above-mentioned variations occur throughout the distribution range. Selected specimens NEW CALEDONIA. Province Nord: Ila Néba, 8 Oct. 1970, MacKee 22736 (NOU, Z); summit of Arama, 8 Sep. 1969, MacKee 20710 (L, NOU, Z); Poum, Col de Pointe, 8 Sep. 1969, MacKee 20723 (K, L, NOU, P); Diahot, 30 Aug. 1951, Hürlimann 1869 (Z); Balade, Apr. 1871, Balansa 3238a (K); Koniambo Plateau, Koné, 16 Jan. 1925, Däniker 3072 (P, Z); Chagrin Mine north-west of Koumac, 15 July 1952, MacMillan 5094 (L); Roche Ouaième, massif de Ton-Non, 18 Apr. 1968, MacKee 18689 (L); Taom, Mount Homédéboa, 16 May 1968, MacKee 18835 (K, L); Tchingou, 17 Apr. 1951, Hürlimann 1208 (Z); Mount Grandié, 14 May 1968, MacKee 18864 (Z); Col d’Amos, 18 Aug. 1965, Bernardi 10318 (L, Z). Province Sud: Canala, Vieillard 832 (Z); Mount Mou, 31 Jan. 1951, Guillaumin & Baumann-Bodenheim 9994 (Z); Thio, Ouégoa, Brousmiche s.n. (K); Ouenghi, Tontouta Valley, 6 Nov. 1967, MacKee 17853 (L); Maquis overlooking valley of Tontouta, 27 Sep. 1979, McPherson 1902 (NOU, NSW); Plateau de Dogny, 12 Apr. 1969, MacKee 20550 (K, L); northern side of Mount Couvelle, Dzumac, 30 Jan. 1991, Briggs 8716 (NSW); summit of Chapeau, 7 Jan. 1869, Balansa 1153 (Z); Upper Pirogue River, 3 Oct. 1924, Däniker 164 (Z); Plaine des Lacs, MacKee 18572 (L); bridge over Creek Pernod, 12 May 2005, Venter 13846 (NOU); Mount Humboldt, 12 Oct. 1896, Balansa 2192 (P). Dracophyllum subgenus Oreothamnus (F.Muell.) W.R.B.Oliv. Trans. & Proc. Roy. Soc. N.Z. 59: 684 (1928). Type: Dracophyllum minimum F.Muell. Section Oreothamnus F.Muell., Fragm. Phytogr. Austr. 1: 39 (1858). Epacris J.R.Forst. & G.Forst., Char. Gen. Pl. ed. 2: 19 (1776), nom. rej. non Cav. Perennial cushion plants, subshrubs, shrubs or trees, 0.15–12 m tall. Leaves sheathing, leaving ringed scars on the branches when falling away, with dry old leaves present in some species; juvenile leaves present in some species; spirally Taxonomic revision of Dracophyllum and Richea arranged along branches or crowded at ends of branches; lamina sheath tapering to auricled and margin smooth to ciliate; lamina longer and wider than adult lamina, coriaceous, linear–triangular, glabrous, rugose, scabrid or pubescent, sometimes with a patch of scabrid hairs at the base of the lamina on the adaxial surface, striated in some species; adult leaves crowded at the ends of branches, spirally arranged along branches or imbricate; lamina sheath tapering, rounded, truncate or auricled and margins smooth to ciliate; lamina coriaceous, linear to linear–triangular, 1–232
0.3–6.0 mm; surfaces glabrous, rugose, scabrid, pubescent to tomentose, with sometimes a tuft of scabrid hairs at the base on the adaxial surface; sometimes striated; margins serrulate or thickly covered in hairs, sometimes cartilaginous; apex obtuse, acicular, or acuminate, sometimes triquetrous. Inflorescence a terminal or lateral raceme, spike or flowers solitary; shorter than the leaves, erect to drooping, lax to dense, 5–70 mm long, linear–oblong to oblong; inflorescence bracts persistent, shorter than or overtopping flowers, light green to reddish-green, subulate to ovate–triangular at the base, 1.5–37
0.5–4 mm, surfaces glabrous to sericeous, margins entire, serrulate to ciliate, apices obtuse to mucronate. Flowers 1–20, sessile or pedicellate; flower bracts persistent or caducous, shorter than or overtopping the flower, membranous to rigid and hard, linear to triangular, 2–20
0.4–8.0 mm, surfaces glabrous to sericeous, sometimes striate, sometimes with a tuft of scabrid hairs either at the apex or at the base, margins entire or ciliate, apices obtuse to subulate; pedicels straight to curved, 0.3–3.0 mm long, glabrous to pubescent. Sepals green to purplish-green, lanceolate to triangular; 0.7–13
0.6–5.5 mm, shorter to longer than the corolla tube, sometimes striate, surfaces glabrous to pubescent, sometimes with the top half pubescent or with scabrid hairs at the base; margins entire, denticulate to ciliate; apices obtuse to acuminate. Corolla white, yellowish to light pink; corolla tube cylindrical, slightly urceolate to narrowly campanulate, narrowed to widened at mouth, 1.8–10
1–4 mm, exterior glabrous; corolla lobes five, imbricate in bud, spreading to strongly recurved, broadly lanceolate to triangular, shorter than corolla tube, 0.8–3
0.8–3.5 mm, apical ridge sometimes present, apices sometimes inflexed, acute to obtuse; surfaces glabrous, adaxial surfaces sometimes papillate. Stamens 5, adnate to corolla tube, filaments 0.1–1.2 mm long; anthers included, dorsifixed, oblong to rectangular, purple to light yellow, 0.3–1.3 mm long. Ovary 5-loculed with pendulous placentae; cylindrical to oblong, 0.5–4.5
0.5–2.5 mm, glabrous to pubescent, apex tapering to truncate; nectary scales 5, separate, rectangular to round, 0.2–1.6
0.3–1 mm, apices acute to obtuse, bidentate to variously toothed; style inserted in a depression at the apex of the ovary, included, 0.5–4.0 mm long, glabrous, sometimes lengthening in fruit; stigma obscurely to prominently 5-lobed. Fruit a loculicidely 5-valved, dry, dehiscent capsule, mostly included in persistent calyx, sessile or pedicellate, light brown to purplish-brown, 1.0–4.5
0.8–4.0 mm, depressed-globose to oblong, apex pointed to truncate, glabrous or pubescent. Seeds are numerous, filiform to ovoid, 0.2–1.3 mm long, testa surface variously reticulate. Chromosome number n = 13. Australian Systematic Botany 77 Species 29; restricted to New Zealand and its off-shore islands, with one species occurring in Tasmania. Key to the species of subgenus Oreothamnus 1. Plants with solitary flowers .................................................................2 Plants with flowers in either a spike or a raceme..............................12 2. Juvenile leaves present, much longer and wider than the adult leaves ....................................................................................D. kirkii Juvenile leaves absent..........................................................................3 3. Flower bract with a broad white margin; plants sparingly leafy ................................................................................. D. palustre Flower bract without a white margin; plants densely leafy ................4 4. Adaxial surface of the flower bract with a tuft of scabrid hairs at the apex; margin of flower bract ciliate; apical ridge on corolla lobe absent............................................................................ D. acerosum Adaxial surface without scabrid hairs at the apex; margin of flower bract serrulate; apical ridge on corolla lobe present ........................5 5. Leaves appressed to stem; dry old leaves persistent ...........................6 Leaves spreading; dry old leaves falling early, leaving a bare stem.................................................................................................10 6. Sepal shorter than the corolla tube; flower longer than the leaves .......................................................................... D. prostratum Sepal equalling or longer than the corolla tube; flower shorter than the leaves ................................................................................................7 7. Lamina apex prominently triquetrous and acute ............. D. minimum Lamina apex acute to obtuse ...............................................................8 8. Sepal margin toothed; apex of corolla lobe acute; apex of fruit truncate .............................................................................D. densum Sepal margin ciliate; apex of corolla lobe obtuse; apex of fruit round.................................................................................................9 9. Corolla tube narrowly campanulate; lamina 1–3 mm long; leaves dull............................................................................... D. muscoides Corolla tube cylindrical; lamina 3.5–12(–17.2) mm long; leaves glossy................................................................................D. politum 10. Sepal margin toothed; ovary truncate; ovary 2.5–4.5 mm long; flower bract longer than flower ............................................. D. frondosum Sepal margin ciliate; ovary round; ovary 0.45–2.1 mm long; flower bract shorter or equalling the flower ..............................................11 11. Flower bract 3.5–4.0
0.6–0.8 mm; ovary 0.5–1.0 mm long; stems decumbent........................................................................ D. pronum Flower bract 5.0–9.5(–13.0)
1.0–2.0 mm; ovary 1.7–2.0 mm long; stems erect ......................................................... D. rosmarinifolium 12. Inflorescence a spike, flowers sessile ................................................13 Inflorescence a raceme, flowers pedicellate ......................................24 13. Juvenile leaves present, much larger and wider than the adult leaves ..............................................................................................14 Juvenile leaves absent........................................................................20 14. Lamina margin densely pubescent .................................. D. arboreum Lamina margin serrulate or ciliate.....................................................15 15. Lamina surfaces pubescent to tomentose; margin of inflorescence bract and lamina margin ciliate; sepal shorter than corolla tube ...............................................................................D. pubescens Lamina surfaces glabrous to shortly scabrid; margin of inflorescence bract either entire or serrulate; lamina margin serrulate; sepal corolla tube ..................................................................................16 16. Apex of ovary covered with short scabrid hairs; abaxial surface of adult lamina scabrous; abaxial surface of sepal pubescent .................................................................. D. trimorphum Apex of ovary glabrous; abaxial surface of lamina glabrous; abaxial surface of sepal glabrous................................................................17 17. Margin of flower bract ciliate; adaxial surface of inflorescence bract pubescent to sericeous; stamens inserted close to top of corolla tube .....................................................................................D. patens 78 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. Australian Systematic Botany Margin of flower bract serrulate; adaxial surface of inflorescence bract glabrous or glabrous with a scabrid base; stamens inserted in the upper third of the corolla tube ............................................. 18 Sepal apex dark coloured, rigid and hard; flower bract narrow, 0.5–0.7 mm; margin of inflorescence bract entire ....D. lessonianum Sepal apices green, not hard; flower bract wide, 1.6–4.1 mm; margin of inflorescence bract serrulate ...........................................................19 Flower bract with broad white margins; apex of ovary truncate; adult lamina 0.5– 1.2 mm wide; corolla tube 1.8–2.0 mm long .............................................................................. D. subulatum Flower bract with green margins; apex of ovary round; adult lamina 2–3 mm wide; corolla tube 4.0–4.5 mm long ...............D. sinclairii Lamina margin pubescent to densely pubescent; adaxial surface of lamina pubescent; abaxial surface of sepal pubescent in the upper half ............................................................................D. scoparium Lamina margin serrulate; adaxial surface of lamina glabrous, rugose–verrucose or scabrid; abaxial surface of sepal glabrous....21 Inflorescence longer than leaves; lamina rugose...............................22 Inflorescence shorter than leaves; lamina smooth.............................23 Corolla lobe with an apical ridge; flower bract shorter than flower; lamina apex not recurved ........................................ D. marmoricola Corolla lobe without an apical ridge; flower bract longer than flower; lamina apex recurved ................................................... D. recurvum Corolla lobe with an apical ridge; dry old leaves persistent; lamina apex with a prominent keel ........................................D. pearsonii Corolla lobe without an apical ridge; dry old leaves deciduous; lamina apex without a prominent keel ...................................... D. filifolium Juvenile leaves absent..................................................D. ophioliticum Juvenile leaves present ......................................................................25 Adaxial surface of adult lamina tomentose; adaxial surface of juvenile lamina scabrid to pubescent; sepal shorter than corolla tube; adult lamina margin densely pubescent ......D. cockayneanum Adaxial surface of adult lamina glabrous, rugose to rarely scabrid; adaxial surface of juvenile lamina glabrous; sepal equalling or longer than corolla tube; adult lamina margin serrulate ...........................26 Flower bracts persistent .....................................................................27 Flower bracts deciduous ....................................................................28 Adult lamina apex triquetrous; corolla tube cylindrical; flower bract 3.0–5.5 mm long; inflorescence bract smooth ................... D. oliveri Adult lamina apex acuminate; corolla tube narrowly campanulate; 11.3–15.6 mm long; inflorescence bract rugose ...... D. urvilleanum Lamina not prominently striated; adult lamina adaxial surface rugose; inflorescence near apex of branch; sepal not striate; stamens inserted near top of corolla tube ........................................ D. septentrionalis Lamina prominently striated; adaxial lamina surface of adult leaf smooth; inflorescence terminal on lateral branchlets; sepal striate; stamens inserted in upper third of corolla tube ..... D. longifolium Dracophyllum acerosum Berggr., Minneskr. Fisiogr. Sällsk. Lund Art. 8: 15 (1877) Type: New Zealand. Mount Torlesse, Feb. 1874. S. Berggren s.n. (holo: O!; iso: BM 577613!, CHR!, K!, S!, WELT 32864!, 32867!, 34027!). D. uniflorum Hook.f. var. acicularifolium Cheeseman, Man. N.Z. Fl.: 427 (1906). Type: New Zealand. Castle Hill, Broken River Basin, T.F. Cheeseman s.n. (lecto: AK 7027!; isolecto: WELT 81525!), designated by Oliver (1952). D. acicularifolium Cockayne, Rep. Scen. Preserv. Soc.: 4 (1915). nom. inval. There are no notes or specimen attached to this name. S. Venter D. peninsulare W.R.B.Oliv., Trans. & Proc. N.Z. Inst. 59: 690 (1928). Type: New Zealand. Banks Peninsula, Mount Berard, 2000 feet [~610 m], R.M. Laing s.n. (CHR 11093, lecto.!), designated by Oliver (1952). Illustrations S. Berggren, Minneskr. Fisiogr. Sällsk. Lund Art. 28: t. 4 (1877); W. R. B. Oliver, Trans. & Proc. Roy. Soc. N.Z. 80 (1): t. 3 (1952); A. Eagle, Trees and Shrubs N.Z. 2nd series: t. 144 (1982); J. Smith-Dodsworth, N.Z. Native Shrubs & Climbers: tt. 55, pl. 23A, B (1991). Erect multi-stemmed shrub or small tree, 1–2 m tall. Branches: bark on old branches grey to dark grey, smooth to finely fissured, young stems reddish-brown. Leaves erect–spreading; lamina sheath (5–)7–20
3.5–5.0(–6.5) mm, coriaceous, striate, truncate to auricled and margin membranous, smooth or with the top half ciliate; lamina linear to linear– triangular, 30–190
0.7–1.5 mm, adaxial surface rugose, abaxial surface glabrous, slightly striated; margins serrulate with 14–21 teeth per 10 mm; apex triquetrous. Inflorescence a solitary terminal flower on lateral branchlets, sessile, shorter than leaves; flower bracts persistent, overtopping flowers, leaf like, ovate–lanceolate, 8.5–17.0
2.5–5.0 mm, surfaces glabrous with a tuft of scabrid hairs at apices, margins ciliate. Sepals lanceolate to narrowly ovate, 8–13
2.0–2.5 mm, shorter to equalling the corolla tube, adaxial surfaces with the top half pubescent; abaxial surfaces glabrous; margins ciliate; apices hard. Corolla white to light green turning yellowish; corolla tube cylindrical, 7–8
1.8–2.2 mm; corolla lobes reflexed, ovate–triangular to triangular, shorter than corolla tube, 1–3
1–2 mm; apices inflexed, subacute; surfaces glabrous. Stamens inserted on corolla tube in upper third, filaments 0.5–1.0 mm long; anthers included, rectangular, light yellow, 0.8–1.0 mm long. Ovary cylindrical, 2.2–3.0
1.3–1.7 mm; glabrous; nectary scales rectangular, 1.5–1.6
0.7–0.8 mm, apices retuse; style included, 1.3–1.5 mm long, glabrous; stigma capitate. Fruit sessile, light brown, 4.0–4.5
4.0–4.2 mm, oblong, apex round, glabrous. Seeds yellowishbrown, ovoid, 1.45–1.5 mm long, testa slightly reticulate (Fig. 66, 67). Distribution and ecology Endemic to the South Island of New Zealand, a species of the eastern mountain ranges of the southern Alps (Fig. 68). It is concentrated in Canterbury, with an isolated occurrence near Ben More in Marlborough. Dracophyllum acerosum occurs on gentle to moderate (2–45) mountain slopes, hillsides, next to rivers or streams and on moraine terraces at elevations from 300 to 1250 m. The vegetation consists of subalpine shrubland, tussock grassland, grassland or herbfield. The soil is brown or yellow to grey clay loam derived from greywacke or loess. Plants normally grow fully exposed. Dracophyllum acerosum is an important member of the subalpine shrubland and mixed snow tussock shrub communities, especially where the plant cover is depleted. At Porters Pass, there are areas where it constitutes 80% of the plant coverage (Venter 13754). Taxonomic revision of Dracophyllum and Richea H I A Australian Systematic Botany B C G D F E Fig. 66. Dracophyllum acerosum. A. Habit (1
). B. Ovary (10
). C. Flower (5
). D. Inflorescence-bract base (5
). E. Sepal adaxial surface (5
). F. Laid-out corolla (5
). G. Lamina sheath (3
). H. Leaf (1
). I. Lamina apex (4
). Drawn from Venter 13754. Del. S. Venter. Phenology Flowering November–May. Etymology Needle-shaped, and refers to the shape of the leaves. Diagnostic features and notes Dracophyllum acerosum is characterised by the erect branches, erect, long acicular leaves with truncate to auricled shoulders to the leaf sheath, solitary erect flowers clustered below the topmost leaves, flower bracts longer than the flowers, with the sepals equalling or shorter than the corolla 79 tube with hard apices, 2.2–3.0-mm-long cylindrical ovary. Oliver (1928) regarded the Cheeseman type (AK7027) as a hybrid and described it as D. peninsulare, but later (1952) rectified this by sinking it under D. acerosum. Dracophyllum acerosum is similar to D. kirkii, but differs in being erect-stemmed and many-branched, with adult leaves that are narrower (0.7–1.5 mm), with the adaxial lamina surface rugose with fewer teeth per 10 mm (14–21) on the lamina margin and having a triquetrous lamina apex. The adaxial surface of the flower bract differs in having a tuft of scabrid hairs at the apex. The sepals are longer (8–13 mm), with the top adaxial half pubescent. The corolla tube is longer (7–8 mm), with no apical ridge on the corolla lobes that is also glabrous on the adaxial surface. The ovary differs in being cylindrical. Dracophyllum acerosum is also similar to D. frondosum, but differs in lacking the prominent apical ridge on the petal and having a subacute corolla-lobe apex, the margins of the flower bracts are ciliate, not serrulate, the nectary scales are longer (1.5–1.6 mm compared with 1.2–1.5 mm), the apex of the ovary is round, not truncate, and the style is much shorter (1.3–1.5 mm compared with 3–4 mm). The leaf sheath varies in size (7–20
3.5–5.0 mm), with the shoulders auricled to various degrees, with the odd specimen having sloped shoulders but the long cilia on them are always present. Lamina length varies (30–190 mm), with less variation in lamina width (1.0–1.3 mm). Flower colour is normally white; however, near Lake Coleridge, there are some populations with limegreen flowers. Sepal length varies from 8 to 13 mm and is usually equal to the corolla tube in length but might be slightly shorter in some flowers. In many aspects, the flowers in D. acerosum and all other Dracophyllum species seem to correspond with the general characteristics for settling moth flowers (phalaenophilous), being pale in colour and with a sweet to sickly sweet odour. Moth pollination is fairly important in New Zealand, probably owing to the paucity of bee species (Arroyo et al. 1982; Primack 1983). Representative collections NEW ZEALAND: South Island: Marlborough, Whernside Ridge, head of Brian Boru stream, Mar. 1975, Druce s.n. (CHR); peak south of Isolated Hill, Apr. 1981, Druce s.n. (CHR); Ben More, Marlborough, Dec. 1979, Druce s.n. (CHR); Puketeraki Range, Mount Whatno, 20 Mar. 1973, Macmillan 73/309 (CHR); Kaikoura, Kowhai Saddle, 27 Jan. 1989, Buchanan (HO); Two Thumb Range, Black Birch Creek, 11 Mar. 1985, Mason s.n. (CHR); Castle Hill, 1885, Enys s.n. (AK); Lake Lyndon, 5 Mar. 1968, Bernardi 12274 (Z); Porters Pass, 26 Oct. 1976, Moar s.n. (CHR); Boundary Hill, 14 Jan. 1973, Thompson 632 (CHR, L, MO); Fogg Peak, Mount Torlesse, Jan. 1969, Adams s.n. (WELT); Kowhai Valley, foot of Porters Pass, 13 Dec. 1947, Oliver s.n. (WELT 56157); Foggy Peak, Mount Torlesse, 28 Jan. 1999, Venter 13754 (CHR); ibid. Feb. 1874, Berggren s.n. (O); ibid., Jan. 1969, Adams s.n. (WELT 54837); lower reaches of Coach Stream, 28 Apr. 1995, Bellingham 723 (CHR); Cass, hill near Sugar Loaf, 18 Jan. 1922, Foy s.n. (CHR); Andrew River, Simpson s.n. (CHR); South Canterbury, Orari Valley, Feb. 1984, Druce s.n. (CHR); Mount Thomas, 23 Dec. 1972, Thompson 342 (CHR); Mount Oxford, 9 Feb. 1955, Mason 3175 (CHR); Banks Peninsula, 80 Australian Systematic Botany A C S. Venter B D Fig. 67. Dracophyllum acerosum. A. Habitat at Porter’s Pass. B. Mature plant near Lake Lyndon. C. Characteristic stiff and erect–spreading leaves. D. Flowering branch showing the solitary flowers and needle-shaped leaves. Photos: S. Venter (A–D). Kaituna, Mount Herbert, Laing s.n. (CHR); ibid., 7 June 1953, Esler s.n. (AK); Banks Peninsula, Mount Sinclair, 23 Sep. 1967, Lambrechtsen s.n. (CHR); Banks Peninsula, Mount Berard, Laing s.n. (CHR); Akaroa, Purple Peak, Oliver s.n. (CHR); Akaroa, Brasenose, Aug. 1917, Oliver s.n. (CHR); Banks Peninsula, Stony Bay Peak, 11 Jan. 1972, Simpson & Chapman s.n. (CHR); Banks Peninsula, Cab Stand, 13 Sep. 1958, Moar 2634 (CHR). Dracophyllum arboreum Cockayne, Trans. & Proc. N.Z. Inst. 34: 318 (1902) Type: New Zealand. Chatham Island, 1901. L. Cockayne s.n. (first-step lecto: WELT, designated by W. R. B. Oliver, Trans. & Proc. Roy. Soc. N.Z. 80(1): 13 (1952); second-step lecto, here Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 81 180º −35º −40º −45º Fig. 68. Known distribution of Dracophyllum acerosum, South Island, New Zealand. designated: WELT 33086!; isolecto: WELT 81570!, WELT 33094!). Dracophyllum latifolium var. ciliolatum Hook.f., Handb. N.Z. Fl. 2: 736 (1864). Type: New Zealand: Chatham Island. F.A.D. Cox s.n. (lecto: WELT 33097!; isolecto: K!, NY, WELT 33095!), here designated. Cheeseman (1906) mentioned some specimens for his var. major but did not indicate which specimens belonged to which variety. However, it is possible to assign the F. A. D. Cox specimens to var. major according to the protologue (Cheeseman 1906). Type: New Zealand. Chatham Islands. W.T.L. Travers s.n. (holo: K000844571). Illustration Dracophyllum scoparium var. major Cheeseman, Man. N.Z. Fl.: 425 (1906). W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 82 (1928); A. Eagle, Trees and Shrubs N.Z. 2nd series: t. 141 (1982). Dracophyllum scoparium F. Muell., Veg. Chath. Is.: 42 (1864). (non Hook.f.) nom. illegit. 82 Australian Systematic Botany Tree 4–6(–12) m tall. Branches: bark on old branches greyish-brown to brown, finely fissured, young stems yellowish- to reddish-brown. Leaves juvenile and adult; juvenile leaves crowded at tips of branches, spreading; lamina sheath yellowish- to light green, (9–)15–17
7.4–16.6 mm, coriaceous, tapering and margin ciliate or ciliate in upper half only; lamina subcoriaceous to coriaceous, linear–triangular, 100–220
10–18 mm, surfaces glabrous, margins densely pubescent; adult leaves spreading; lamina sheath light green, 6–12
4–12 mm, membranous, tapering with a ciliate margin; lamina linear to linear–triangular, (25–) 40 – 86 (–90)
1–2 mm, surfaces glabrous with a tuft of scabrid hairs at the base of the adaxial surface; margins densely pubescent. Inflorescence a terminal spike on lateral branchlets, shorter than leaves, erect to drooping, dense, 15–38 mm long, linear–oblong; inflorescence bract overtopping the flower, subulate, 18–20
3–5 mm, surfaces glabrous, adaxial surface pubescent at the base, margins ciliate. Flowers (4–)6–9, sessile; flower bract persistent, overtopping flowers, leaf like, ovate to broadly ovate; 5.5–9.0
2.5–3.0 mm, surfaces glabrous, adaxial surface with a tuft of scabrid hair at apex; margins ciliate. Sepals ovate lanceolate, (4–)5–7
2.5–3.0 mm, longer than corolla tube, surfaces glabrous with the top half pubescent; margins ciliate. Corolla white; corolla tube cylindrical, 4–5
2.5–3.0 mm; corolla lobes reflexed, triangular, shorter than corolla tube, (2.0–)2.3–2.4
1–2 mm; apices acute; adaxial surface papillate. Stamens inserted on corolla tube in the upper third, filaments 0.3–1.0 mm long; anthers included, oblong, light yellow, 0.3–0.4 mm long. Ovary obovoid, 1.7–2.0
1.0–2.0 mm; glabrous, apex round; nectary scales, oblong, 1.0–1.2
0.5–0.8 mm, apices irregularly toothed; style included, 2.0–2.5 mm long, glabrous; stigma capitate. Fruit sessile, dark brown, 1.2–1.5
1.0–1.5 mm, oblong, apex round, glabrous. Seed yellowish-brown, ovoid, 0.6–0.65 mm long, testa slightly reticulate (Fig. 69, 70). Distribution and ecology New Zealand endemic restricted to the Chatham Island and Pitt Islands (Fig. 71). Dracophyllum arboreum occurs from near sea level up to 270-m elevation in gullies, gully floors, along streams, hillsides and on coastal cliffs. The vegetation consists of forest, shrubland, lowland bogs and grassland. Soils are typically boggy and peaty. Most D. arboreum plants grow in full sun, but occasionally occur in light shade inside forest communities. Phenology Flowering November–February. Etymology Refers to the tree growth habit of the species. Diagnostic features and notes Oliver (1952) mentioned a specimen in WELT as the type of D. arboretum, but did not indicate a particular herbarium sheet. The specimen WELT 33086 (ex Petrie Herbarium) agrees with the protologue and a photograph of it was S. Venter E A F G J H D B C I Fig. 69. Dracophyllum arboreum. A. Habit (1
). B. Lamina margin (40
). C. Laid-out corolla (5
). D. Lamina sheath (2
). E. Leaf (1
). F. Ovary (10
). G. Flower (7
). H. Sepal abaxial surface (5
). I. Nectary scale (10
). J. Inflorescence-bract adaxial surface (5
). Drawn from Cockayne s.n. (WELT 33086). Del. S. Venter. published by Oliver (1928) as plate 82 in his revision of Dracophyllum. It is chosen here as a second-step lectotype. Dracophyllum arboreum is characterised by trees growing up to 6 m tall, long and broad juvenile leaves with adult leaves densely ciliated on the margins and pubescent at the base, persistent hard and sharp-tipped bracts that are broad with long white hairs on the adaxial surface, corolla tube 4–5 mm long and shorter than the sepals that have long cilia and hairs on the abaxial surface. Dracophyllum arboreum is similar to D. cockayneanum and D. scoparium, but can immediately be separated from D. scoparium, which lacks juvenile leaves. It differs from D. cockayneanum in the glabrous surfaces of the juvenile lamina that is also wider (10–18 mm compared with 4–6 mm) and the flower bracts are persistent. It also differs in the tapering shoulder of the adult lamina sheath (compared with Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 83 A B C D Fig. 70. Dracophyllum arboreum. A. Habitat in the Tuku Nature Reserve. B. Flowering branch from near Lake Rakeinui. C. A mature tree with the undergrowth removed by livestock. D. Juvenile leaves. Photos: Taiko Trust (A, C) and P. de Lange (B–D). rounded and truncate) and glabrous lamina with a tuft of scabrid hairs at the base (compared with tomentose). The inflorescence in D. arboreum is a spike with a persistent glabrous flower bract having a tuft of scabrid hairs at the apex on the adaxial surface. The sepals are longer than the corolla tube, with the upper half of the sepal being pubescent on the abaxial surface and the cylindrical corolla has lobes with papillate adaxial surfaces. The apices of the nectary scales in D. arboreum are irregularly toothed, but subacute to obtuse in D. cockayneanum. Juvenile and adult leaves are quite variable in shape and size. The abaxial surface of the sepal can be either completely glabrous or only pubescent in the 84 Australian Systematic Botany S. Venter 176º45E 177ºE 177º15E 177º30E The Sisters CHATHAM ISLANDS (New Zealand) Cape Young 43º45’S Cape Patisson Point Munning Port Hutt Point Somes Te Whanga Lagoon Petre Bay Hanson Bay The Forty Fours Waitangi 44ºS Point Durham Chatham Island Owenga Manukau Point Pitt Strait Cape L’Eveque Star Keys 44º15’S Mangere Island Little Mangere Island The Castle Pitt Island PACIFIC OCEAN South East Island 0 30 km © Australian National University CartoGIS CAP 00-259 The Pyramid Fig. 71. Known distribution of Dracophyllum arboreum. The large island is Chatham Island and the smaller island is Pitt Island. upper half. Filament length (0.3–1.0 mm) varies from population to population. Seeds commonly germinate on tree ferns, growing for a period as an epiphyte. Plants remain in the juvenile form with large leaves for many years before taking on the adult leaf shape and size. Leaves of the juvenile form commonly appear high up in the branches of adult D. cockayneanum plants and are here called reversion shoots. Representative collections NEW ZEALAND. Chatham Islands: Ocean Mail Scenic Reserve, 24 Feb. 1996, De Lange CH97 (AK, HO); Te Awatea, swamp forest on southern Taxonomic revision of Dracophyllum and Richea shore of Lake Huro, 20 Feb. 1982, Given & Williams 13018 (AK, WELT); south of Lake Huro, 28 Jan. 1974, Hamel s.n. (CHR); Te Moko Creek, Ocean Bay, 6 Mar. 1985, Given 14050 (CHR); head of Mangahou Creek, 14 Feb. 1985, Wardle 85/2 (CHR); Rangatira Island, Rangatira Trig., 27 Feb. 1986, Courtney s.n. (CHR); Woolshed Bush, 7 Mar. 1986, Courtney s.n. (CHR); top of large slump, south-east coast, 14 Feb. 1985, Wardle 85/1 (CHR); Pitt Island, between Glory Bay and Canister Cove, 17 Feb. 1985, Wardle 85/7 (CHR); ibid., 5 Jan. 1970, Hamlin 1816 (WELT); near Point Durham, 7 Sep. 1959, Moar 2591 (CHR); Lake Rakeinui, 18 Feb. 1982, Given s.n. (CHR); Tukutamatea Valley, 15 Jan. 1978, Olsen s.n. (AK, HO). Dracophyllum cockayneanum Du Rietz, Svensk Botanisk Tidskrift 24(3): 375 (1930) Type: New Zealand. Campbell Island, Perseverance Harbour, 1 Apr. 1927. W.R.B. Oliver s.n. (holo: WELT 280!). Dracophyllum longifolium (J.R.Forst. & G.Forst.) R.Br. ex Roem. & Schult. var. cockayneanum (Du Rietz) W.R.B.Oliv., Trans. & Proc. Roy. Soc. N.Z. 80 (1): 11 (1952). Australian Systematic Botany 85 1.2–2.0 mm, apex truncate; nectary scales rectangular, 1.2–1.5
0.6–1.0 mm, apices subacute to obtuse; style included, 1.5–2.0 mm long, glabrous, lengthening in fruit; stigma 5-lobed. Fruit pedicellate, light brown, 2.5–3.5
3–4 mm, obovoid, apex truncate, glabrous. Seeds light brown, ovoid, 0.65–0.7 mm long, testa slightly reticulate (Fig. 72, 73). Phenology Flowering October–March. Etymology Named after Dr Leonard Cockayne (1855–1934), New Zealand botanist and plant geographer. Distribution and ecology Endemic to the subantarctic Campbell, Auckland and Enderby islands of New Zealand (Fig. 74). Widespread, growing from Illustration P. Wardle, New Zealand J. Bot. 25: 112–113 (1987). Tree 3–12 m tall. Branches: bark on old branches brown, rough to occasionally finely fissured, young stems reddishbrown. Leaves juvenile and adult. Juvenile leaves spirally arranged along branches, spreading; lamina sheath 15–20
12–14 mm; shoulders tapering to rounded, margin ciliate; lamina linear–triangular; 100–140
4–6 mm; adaxial surface scabrid to pubescent, abaxial surface glabrous to scabrid, margins densely pubescent; adult leaves erect–spreading; lamina sheath light green, 12.0–18.3
9–13 mm, coriaceous, shoulders rounded to truncate and margin ciliate in the top half; lamina linear to linear–triangular, (40–)60–120(–140)
1.5–3.0 (–4.0) mm, adaxial surface tomentose; abaxial surface with upper half covered in scabrid hairs, slightly striated; margins serrulate and densely pubescent with 60–70 teeth per 10 mm. Inflorescence a terminal raceme on lateral branchlets; shorter than leaves, erect to drooping, dense, 30–70 mm long, linear–oblong; inflorescence bract persistent, overtopping flowers, subulate, 15–30
1.5–2.0 mm, adaxial surface scabrid, abaxial surface glabrous, margins ciliate. Flowers 6–12(–20), pedicellate; flower bracts caducous, overtopping flowers, ovate–lanceolate to ovate; 8.0–14.0
3–6 mm, adaxial surfaces pubescent or pubescent in the upper third only; abaxial surfaces glabrous, margins ciliate; pedicels straight, 1–3 mm long, glabrous to pubescent. Sepals ovate–lanceolate, 4–5
2.0–2.5 mm, shorter than the corolla tube, surfaces glabrous but occasionally pubescent in the top half of the adaxial surface; margins ciliate. Corolla white to occasionally light pink; corolla tube narrowly campanulate, widened at mouth, 4–6
3.0–3.5 mm; corolla lobes reflexed, ovate–triangular, shorter than corolla tube, 1.5–2.0 mm long and wide; surfaces glabrous. Stamens inserted on corolla tube in the upper third, filaments 0.5–1.0 mm long; anthers included, oblong, pink, 0.7–0.8 mm long. Ovary obovoid, 1.5–2.0
Fig. 72. Dracophyllum cockayneanum. A. Flowering branch (1
). B. Laidout corolla (5
). C. Juvenile leaf (1
). D. Adult leaf (1
). E. Ovary (10
). F. Flower-bract adaxial surface (5
). G. Flower (5
). H. Sepal adaxial surface (5
). Drawn from Hooker s.n. (HO). 86 Australian Systematic Botany S. Venter Auckland Island Fig. 73. Dracophyllum cockayneanum. Habitat on Enderby Island (Phil Garnock-Jones). the coast to the main summit ridges. Dracophyllum cockayneanum grows from sea level to 450-m elevation on ridge crests, in gullies and, to a lesser degree, in open flat areas. The vegetation consists of low forest, open shrubland and, to a lesser degree, grassland also in permanently moist peaty soils (Fig. 74). Etymology Campbell Island Named after the botanist and naturalist Dr Leonard Cockayne. Diagnostic features and notes Dracophyllum cockayneanum is characterised by the erect stiff leaves that are mostly white tomentose in the bottom half, and the margin ciliated with broad sheaths having rounded to truncate shoulders with dense white cilia. The pedicel is 1–3 mm long and the flower bracts are caducous, having the upper third of the adaxial surface covered in dense long white hairs. Du Rietz (1930) argued that, in his experience, Oliver’s (1928) D. longifolium ‘form 3’ is just as distinct as many of the species recognised by Oliver and, therefore, raised it to the rank of species as D. cockayneanum. Later Oliver (1952) reduced it to a variety of D. longifolium on the basis of the broad, stiff, spear-like leaves with broad sheaths, erect racemes and the deciduous flower bracts (Oliver 1952). Morphologically, it is similar to D. longifolium (Table 8), but it differs in various leaf and flower characters. The combination of the following characters prompted the reinstatement of D. longifolium var. cockayneanum at species level: the erect stiff adult leaves that are white tomentose in the bottom half, rounded to truncate shoulders of the lamina sheath, ciliate margin of the inflorescence bract, sepals shorter than the corolla tube and the truncate apex of the ovary. There is some variation in size of the adult lamina sheath (13.0–18.3
10–13 mm), adult lamina (70–120
1.5–2.5 mm) and the ovary (1.5–2.0 mm long and wide). Fig. 74. Known distribution of Dracophyllum cockayneanum. The larger island is Auckland Island and the small island to the south-east, Campbell Island. Table 8. Differences between Dracophyllum D. cockayneanum Character Juvenile lamina surface Adult lamina sheath size (mm) Adult lamina adaxial surface Lamina margin Inflorescence bract margin Sepal length compared to corolla tube Sepal surface texture Corolla-lobe adaxial surface Nectary-scale apex Ovary apex shape Seed shape longifolium and D. longifolium D. cockayneanum Glabrous 5.0–15
4.0–7.0 Scabrid to pubescent 12.0–18.3
9.0–13.0 Glabrous Serrulate Serrulate Equalling or longer Glabrous, top half rarely pubescent Papillate Irregularly toothed Round Ovoid Tomentose Densely pubescent Ciliate Shorter Striate Glabrous Subacute to obtuse Truncate Filiform Representative collections NEW ZEALAND. Enderby Island, July 1903, Cockayne 3370 (AK, WELT); Moorland near centre of island, 3 Dec. 1972, Campbell s.n. (WELTU); Auckland Island, Erebus Cove, 6 Dec. 1972, Burke s.n. (WELTU); Port Ross, Taxonomic revision of Dracophyllum and Richea 8 Nov. 1954, Moar 730 (CHR); Campbell Island, Beeman Hill, 2 Nov. 1966, Roberts s.n. (WELTU); Tucker Cove, 13 Nov. 1944, Oliver s.n. (WELT). Australian Systematic Botany D E 87 F Dracophyllum densum W.R.B.Oliv., Trans. & Proc. Roy. Soc. N.Z. 80 (1): 3 (1952) Type: New Zealand. Westport, Mount Rochfort, 2000 feet [~610 m], 2 Mar. 1949. W.R.B. Oliver s.n. (holo: WELT 314!). A Illustration W. R. B. Oliver, Trans. & Proc. Roy. Soc. N.Z. 80 (1): t. 1 (1952). Scrambling subshrub or low shrub, 3–50 cm tall. Branches spreading, decumbent or prostrate. Bark on old branches dark grey to brown, deeply fissured, young stems grey to reddishbrown. Leaves spirally along branches, imbricate, erect and appressed to stem, old leaves present; lamina sheath olivegreen, 2–5
2–4 mm, membranous, tapering and margin ciliate; lamina olive-green, linear, 6–22
0.7–1.0 mm, adaxial surface flat, abaxial surface rounded with a keeled apex, surfaces glabrous with a tuft of scabrid hairs at the base on adaxial surface, slightly striated; margins serrulate with 25–30 teeth per 10 mm; apex obtuse to acute. Inflorescence a solitary terminal flower on lateral branchlets, sessile, shorter than leaves, erect, mostly hidden by leaves; flower bract shorter than or equal to flower, leaf-like, coriaceous to rigid and hard, broadly ovate at the base, 3.5–7.0
2–3 mm, surfaces glabrous with a tuft of scabrid hairs at the base of the adaxial surface, margins serrulate, apex obtuse to acute. Sepals lanceolate, 3.5–5.7
1.5–2.0 mm, equalling corolla tube, top half pubescent on adaxial surface; margins toothed; apex subacute to obtuse. Corolla white; corolla tube cylindrical, 3.5–5.0
1.0–1.5 mm; corolla lobes spreading horizontally to reflexed, ovate–triangular to triangular, shorter than corolla tube, 1.5–2.0
1.0–1.3 mm; apical ridge present, apices acute; adaxial surface papillate. Stamens inserted on corolla tube in the upper-third, filaments 0.5–0.55 mm long; anthers included, rectangular, light yellow, 1 mm long. Ovary obovoid, 1–2
1.0–1.4 mm; apex truncate; nectary scales rectangular, 0.9–1.0
0.6–0.8 mm, apices retuse to irregularly toothed; style included, 1.5–1.55 mm long, glabrous, lengthening in fruit; stigma 5-lobed. Fruit sessile, light brown, 1.3–2.0
1–2 mm, depressed-obovoid to obovoid, apex truncate, glabrous. Seeds light brown, ovoid, 0.45–0.5 mm long, testa surface slightly reticulate (Fig. 75, 76). G C B I H Fig. 75. Dracophyllum densum. A. Habit (1
). B. Laid-out corolla (1
). C. Lamina sheath (5
). D. Leaves (1
). E. Nectary scale (10
). F. Ovary (10
). G. Flower (5
). H. Inflorescence-bract adaxial surface (5
). I. Sepal adaxial surface (5
). Drawn from Venter 13746. Del. S. Venter. Phenolog Flowering January–April. Etymology Refers to the densely packed leaves. Distribution and ecology New Zealand endemic restricted to the north-western Nelson area of the South Island (Fig. 77). Most localities are in the Westport–Karamea area and not further than 20 km from the sea. Dracophyllum densum occurs on gentle to moderate (3–10) slopes on mountain summits, terraces and plateaux, at elevations ranging from 610 to 1218 m. The vegetation consists of montane shrubland, subalpine shrubland, herbland, tussockland and grassland. The soil is a greyish-brown sandy lithosol derived from sandstone, conglomerate or a gritty sandy loam derived from granite. Diagnostic features and notes Dracophyllum densum is characterised by the prostrate habit, dark grey deeply fissured bark, old leaves being persistent for a very long time, erect leaves clasping the stem and with obtusely acute apices, solitary flowers, sepals with subacute to obtuse apices and equalling the corolla tube and the short capsule that is widest at the top (Table 9). Oliver (1952) placed Dracophyllum densum nearest to D. pronum, but it differs in being a shrublet, is more compact with denser foliage and having larger leaves with 88 Australian Systematic Botany S. Venter B A C D E F Fig. 76. Dracophyllum densum. A. Habitat on Mount Garibaldi. B. Plant with cushion growth habit (Venter 13859), Mount Garibaldi. C. Mature plant from the Denniston Plateau. D. Flowering branch, mature plant showing the typical gnarled main stem. D. Flowering branch, plant from the Denniston Plateau. E. Flowering plant, Denniston Plateau. F. Mature plant with a prostrate growth habit, Denniston Plateau. Photos: S. Venter (A–C) and Phil Bendle (D–F). more acute apices. The branches are clothed with dead leaves (or their bases), a character shared with D. politum, D. muscoides and D. prostratum. Dracophyllum densum is similar to D. politum (Table 7) but differs in leaf and flower characters. Growth habit varies from a flat compact shrublet to a small shrub up to 50 cm tall with trailing branches. The leaf sheath varies in length (2.5–5.2 mm) and so do the sepals (4.5–9.0 mm long). Lamina apices are either obtuse or Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 170º 175º 89 180º −35º −40º −45º Fig. 77. Known distribution of Dracophyllum densum. South Island, New Zealand. Table 9. Morphological differences between Dracophyllum densum and D. politum Character D. densum D. politum Tuft of scabrid hairs on lamina Number of teeth (per 10 mm) on lamina margin Flower-bract size (mm) Flower-bract margin Sepal length Present 25–30 Absent 90–100 3.5–7.0
2.0–3.0 Ciliate Equals corolla tube Apex of corolla lobe Ovary apex shape Seed testa surface Acute Truncate Slightly reticulated 2.0–3.0
0.7–0.8 Serrulate Longer than corolla tube Obtuse Round Reticulated acute on the same plant. In spite of Oliver (1952) describing the lamina apex as obtuse, I found that most plants have acute apices. The flower bracts vary in size (3.5–7.0
2–3 mm) and the corolla lobes in shape from ovate–triangular to triangular. Representative collections NEW ZEALAND. South Island: near Denniston, 6 Jan. 1968, Brownlie 664 (CHR); Denniston, 14 Nov. 1966, Davidson s.n. (CHR); Burma Road, 28 Jan. 1953, Mason & Moar 1962 (CHR); Westport, Mount Rochfort, Denniston Plateau, 7 Jan. 1999, S. Venter 3746 (CHR); below Garibaldi Ridge, Mar. 1980, Druce s.n. (CHR); Greymouth, Paparoa Range, Mount Davy, 26 Feb. 1949, Oliver s.n. (WELT); east of Mount Priestly, 12 Apr. 1983, Wardle s.n. (CHR). 90 Australian Systematic Botany Dracophyllum filifolium Hook.f., Fl. Nov. Zel. 2 (1): 169 (1853) Type: New Zealand. Ruahine Range, 1849. W. Colenso s.n. (lecto: K000844595!; iso: K000844590!), designated by Oliver (1952). Dracophyllum setifolium Stchegl., Bull. Soc. Imp. Naturalistes Moscou 32 (1): 23 (1859). Type: New Zealand. Auckland, 1843. W.Stephenson 77 (holo: K!). Dracophyllum pungens Colenso, Trans. & Proc. N.Z. Inst. 28: 602 (1896). Type: New Zealand. Eastern side of Ruahine Mountain range, 1895. H. Hill s.n. (holo: K!). Dracophyllum virgatum Colenso, Trans. & Proc. N.Z. Inst. 28: 605 (1896). Type: New Zealand. Eastern side of Ruahine Mountain Range, 1895. A. Olsen s.n. (holo: WELT 23628!; iso: K!). Dracophyllum heterophyllum Colenso, Trans. & Proc. N.Z. Inst. 28: 605 (1896). Type: New Zealand. Eastern side of Ruahine Mountain range, 1895. E.W. Andrews s.n. (holo: K!; iso: WELT 42835!). Dracophyllum
vulcanicum W.R.B. Oliv., Trans. & Proc. N.Z. Inst. 59: 697 (1928). Type: New Zealand. Waimarino Plains, 2600 feet [~792 m], 22 Dec. 1923. H. Carse s.n. (holo: CHR 332586!; iso: K!). Dracophyllum urvillianum var. filifolium Cheeseman, Man. N.Z. Fl.: 424 (1906). Dracophyllum collinum W.R.B.Oliv., Trans. & Proc. N.Z. Inst. 59: 696 (1928). Type: New Zealand. Mountains of the Tinline Valley, Nov. 1924. J.H. McMahon s.n. (holo: WELT 55185!). Dracophyllum filifolium var. centrale W.R.B.Oliv., Trans. & Proc. Roy. Soc. N.Z. 80 (1): 10 (1952). Type: New Zealand. Mount Ruapehu, 3800 feet [~1158 m], 7 Feb. 1949. W.R.B. Oliver s.n. (holo: WELT 318!). Dracophyllum filifolium var. collinum W.R.B.Oliv., Trans. & Proc. Roy. Soc. N.Z. 80 (1): 10 (1952). Illustrations W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: 696, t. 7a (1928); A. Eagle, Trees & Shrubs of N.Z., 2nd series: t. 138 (1982); A. F. Mark and N. M. Adams, N.Z. Alpine Plants: t. 45 (1986). Erect multi-stemmed shrub or tree, 1–4 m tall. Branches: bark on old branches grey to brown, finely fissured, young stems reddish-brown. Leaves erect–spreading; lamina sheath S. Venter 6–16
3.0–5.5 mm, membranous, shoulders tapering to auricled and the margin ciliate in the upper half; lamina linear to linear–triangular, (40–)60–130(–200)
0.7–1.5 mm, surfaces glabrous with a tuft of scabrid hairs at the base of the adaxial surface, slightly striated; margins serrulate with 18–25 teeth per 10 mm; apex acute to weakly triquetrous. Inflorescence a terminal spike on lateral branchlets, shorter than leaves, erect to drooping, dense, 20–30 mm long, linear–oblong; inflorescence bract overtopping flowers, ovate–lanceolate, 8.5–17
2–4 mm, surfaces glabrous; margins ciliate. Flowers 5–9(–10), sessile; flower bracts overtopping flowers, broadly ovate; 5.6–9.3(–13.0)
3–4 mm, surfaces glabrous with a tuft of scabrid hair at the base of the adaxial surface, margins ciliate. Sepals ovate–lanceolate, occasionally ovate, 4–6
1.5–1.7 mm, equalling corolla tube, surfaces glabrous; margins ciliate. Corolla white; corolla tube cylindrical, 3.5–5.5(–6.0)
1.8–2.5 mm; corolla lobes reflexed, ovate, shorter than corolla tube, 1.5–2.0
1.5–1.6 mm; apices subacute to acute; surfaces glabrous. Stamens inserted on corolla tube in the upper third, filaments 0.2–0.3 mm long; anthers included, oblong, light yellow and 1 mm long. Ovary subglobose, 1.5–2.0
1.0–1.5 mm, glabrous, apex round; nectary scales oblong, 0.7–1.0
0.5–0.7 mm; apices obtuse to retuse; style included, 1–2 mm long, glabrous; stigma 5-lobed. Fruit sessile, light brown, 2.0–2.5
2.0–2.2 mm, obovoid, apex round, glabrous. Seeds yellowish-brown, filiform, (0.8–)1.0–1.1(–1.2) mm long, testa slightly reticulate (Fig. 78, 79). Distribution and ecology New Zealand endemic, restricted to the North Island, South Island and Stewart Island (Fig. 81). Dracophyllum filifolium occurs on gentle (5–30) mountain slopes, on saddles and mountain ridges. At elevations from near sea level up to 1500 m. The vegetation consists of lowland to montane forest, shrubland and grassland. In montane areas, D. filifolium forms extensive populations within Leptospermum scoparium shrubland and very much occupies the same ecological niche. Soils are gritty dark brown sandy loam or brown clay loam derived from greywacke and roll stone conglomerate, or on grey clay loam lithosol or grey clay loam derived from basalt, shale, serpentinite and granidiorite. Phenology Flowering November–April, with a few individuals being recorded flowering as late as June. Etymology Named for the long and narrow filament-like leaves. Diagnostic features and notes Hooker (1853) mentioned two syntypes in the protologue and said that there are more known localities. However, Oliver (1952) selected a specimen at Kew, collected by William Colenso in 1849, as type. Allan (1961) stated that he did not find the Colenso specimen mentioned by Oliver and, Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A G 91 similar to D. urvilleanum and D. lessonianum, but differs in leaf, inflorescence and flower characters. The growth habit of Dracophyllum filifolium is polymorphic. Populations at the Cobb Reservoir near Takaka, on the Dun Mountain near Nelson and on D’Urville Island in the Marlborough Sounds grow as a low roundish shrub with erect branches and thickly textured, stiff and short leaves. This is the usual growth habit on serpentinite (Venter 13737 and Venter 13785). Populations at West Whanganui Inlet at the northern tip of the South Island grow as a columnar small tree with erect–spreading branches and thinly textured, erect, long leaves. In some populations on Stewart Island (Venter 13791), the flower bract is lightly attached and sometimes drops while the plant is still in full flower. F Representative collections H E B D C Fig. 78. Dracophyllum filifolium. A. Flowering branch (1
). B. Inflorescence-bract adaxial surface (5
). C. Sepal adaxial surface (5
). D. Laid-out corolla (5
). E. Lamina sheaths to show variation (5
). F. Leaf (1
). G. Ovary (10
). H. Flower (5
). Drawn from Venter 13762. Del. S. Venter. therefore, did not cite any material; however, the specimen has since been re-traced at Kew. Dracophyllum filifolium is characterised by the inflorescence being a spike 17–24 mm long, with the inflorescence bracts and flower bracts longer than the flower, sepals equal the corolla tube, which is 3.0–4.5 mm long, with the corolla lobes 1.5–2.0 mm wide, and nectary scales 1.0–1.3 mm long with obtuse apices. I have found that in D. filifolium nectar is secreted during the afternoon and early evening, reaching the maximum volume available to pollinators (moths) c. 2000 hours, with a second period of nectar secretion commencing from 1000 hours on a sunny day and ceasing at c. 1500 hours (Fig. 80). It is this second secretion period that is utilised by butterflies of the genera Argyrophenga and Lycaena (S. Venter, pers. obs.). Dracophyllum filifolium is a much-misunderstood taxon, with earlier authors placing D. lessonianum and D. urvilleanum into it, resulting in a rather confused nomenclature and literature. Dracophyllum filifolium is NEW ZEALAND. North Island: Mount Kakaramea, Jan. 1905, Cheeseman s.n. (AK); Rainbow Mountain, May 1947, Sibson s.n. (AK); Urewera National Park, west of Waikareiti, Puna Hokoi Clearing, Feb. 1968, Druce s.n. (CHR); Urewera, Maungapohatu, 19 Mar. 1930, Moore s.n. (CHR); Waikaremoana, Panekiri Bluff, Jan. 1954, Druce s.n. (CHR); Panekirikiri Bluff [Panekiri Bluff], 20 Jan. 1953, Druce s.n. (CHR); Egmont National Park, Pouakai Range, near tarns on Kaiauai Track, 7 Feb. 1999, Venter 13762 (CHR); Mount Egmont, near Wilkie’s Pools, 19 Jan. 1934, Cranwell s.n. (AK); Taumaranui, east of National Park Township, 2 Jan. 1970, Donovan s.n. (CHR); Tongariro National Park, Mount Hauhungatahi, 14 Sep. 1924, Oliver s.n. (CHR); Waimarino Stream, Jan. 1965, Druce s.n. (CHR); Chateau Tongariro, Jan. 1960, Puffy s.n. (CHR); Kaimanawa Mountains, eastern foot of Stowman Range, Jan. 1973, Druce s.n. (CHR); north-western Ruahine Range, Waiokotore Stream, Dec. 1973, Druce s.n. (CHR); Rangiwahia, Esler s.n. (AK); mouth of Green Hills Stream, July 1979, Druce s.n. (CHR); South Ruahine Range, Feb. 1971, Druce s.n. (CHR); Tiritea Catchment, 4 Oct. 1958, Esler s.n. (AK); Waiopehu Track, 18 Feb. 1934, Zotov s.n. (CHR); Tararua Ranges, Ngapuketurua, 1 Mar. 1931, Zotov s.n. (CHR); Mount Holdsworth, 28 Jan. 1906, Cockayne s.n. (CHR); southern Ruahine Range, Takapari, Jan. 1966, Druce s.n. (CHR); south-west of Akitio, north of Mount Kupukore, Dec. 1973, Druce s.n. (CHR); Wellington, top of Days Bay Hills, 27 July 1921, Atkinson s.n. (CHR); Wainuiomata Valley, western slopes, 26 July 1937, Healy s.n. (CHR); eastern Tararuas, track to Mount Holdsworth, 23 Jan. 1967, Hynes s.n. (AK); Tararua Range, Mayhorn, Jan. 1968, Druce s.n. (CHR); Tararua Mountains, Smiths Creek, 26 Dec. 1932, Zotov s.n. (CHR); Mount Matthews, Apr. 1931, Oliver s.n. (CHR); Aorangi Range, Mount Whawanui, Sep. 1969, Druce s.n. (CHR); Wairarapa, Oterei Taipos, Heights, Dec. 1978, Druce s.n. (CHR); east-north-east of Cape Palliser, 2 Nov. 1966, Druce s.n. (CHR); eastern Wairarapa, taipos above the Oterei River, Dec. 1957, Druce s.n. (CHR); South Island: Cape Farewell, Ngurua Bay Road, 30 Oct. 1998, Venter 13725 (CHR); Collingwood, Knuckle Hill, 15 Jan. 1999, Venter 13,753 (CHR); northern end of Whanganui Inlet, 20 Feb. 1957, Mason & Moar 4865 (O); Mount Perry, Jan. 1973, Druce s.n. (CHR); Takaka, Haupiri Range, Walker Ridge, 26 Dec. 1951, Hay s.n. (CHR); Abel Tasman National Park, Moa Park, Jan. 1969, Druce s.n. (CHR); Takaka, Awaroa, 23 Oct. 1961, Scott s.n. (CHR); Gouland Downs, Dec. 1951, Talbot s.n. (CHR); near head of Whanganui Inlet, 13 Feb. 1969, Sykes 44/69 (CHR); Lead Hills, Gibbs s.n. (CHR); Gunner Downs, Nov. 1979, Druce s.n. (CHR); peak at head of Cobb River, 1 Jan. 1943, Mason s.n. (CHR); Mount Luna, Jan. 1971, Druce s.n. (CHR); Motueka, Cobb Valley, ridge above Cobb 92 Australian Systematic Botany S. Venter A B C D E F Fig. 79. Dracophyllum filifolium. A. Plants in habitat on Dunn Mountain. B. Flowering plant from Dunn Mountain. C. Mature plant on serpentinite, Cobb Reservoir. D. Thick layer of shed leaves of the past winter season on Dunn Mountain. E, Mature plants growing on the slopes of Mount Taranaki. F. Dense D. filifolium scrub on the serpentinite soils of Dunn Mountain. (C. Venter 13737 and B. Venter 13785). Photos: S. Venter (A–D, F) and Phil Bendle (E). Volume (µL nectar flower–1) Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 93 Nectar secretion 4 Oreothamnus Dracophyllum 2 0 1000 1200 1400 1600 1800 2000 2200 2400 Time (hours) A Fig. 80. Peak nectar flow periods in Dracophyllum traversii (closed squares) and D. filifolium (closed circles). Information building, 11 Dec. 1998, Venter 13737 (CHR); Motueka, Mount Arthur, Flora Saddle, 2 Jan. 1999, Venter 13740 (CHR); Mount Patriarch, Jan. 1974, Druce s.n. (CHR); Mount Arthur, Gibbs s.n. (CHR); Tararua Ranges, Mount Dennan, 5 Feb. 1931, Zotov s.n. (CHR); Saint Arnaud, Big Bush, 27 May 1964, Fryer s.n. (CHR); Gordon’s Knob, 16 Feb. 1975, Simpson 7598 (CHR); Lake Rotoiti Peninsula, 16 Feb. 1961, Simpson 2809 (CHR); D’Urville Island, Mount Ears, June 1975, Kelly & Kelly s.n. (CHR); Nelson, Dun Mountain, before Dun Saddle, 28 Sep. 1999, Venter 13785 (CHR); Tennyson Inlet, 16 Feb. 1969, Hynes s.n. (AK); Okaramio, Kaituna Valley, 24 Dec. 1935, Healy s.n. (WELTU); Wairau Valley, Red Hills, Jan. 1898, Cheeseman s.n. (AK); Gordon’s Knob, 24 Jan. 1922, Allan s.n. (AK); Endeavour Inlet, Dec. 1945, Healy s.n. (CHR); Mawheraiti, 19 Apr. 1948, Oliver s.n. (CHR); Upper Hurunui River, Mac’s Knob, 30 Jan. 1973, Macmillan 73/73 & Stemmer (CHR); Stewart Island: Mount Anglem, near top, 11 Jan. 2000, Venter 13792 (CHR). B C D Dracophyllum frondosum (G.Simpson) S.Venter comb. et stat. nov. Dracophyllum uniflorum Hook.f. var. frondosum G.Simpson, Trans. & Proc. Roy. Soc. N.Z. 79: 434 (1952). Type: New Zealand. Otago, Deep Stream at bridge on Dunedin, Middlemarch Road, 5 Feb. 1949, G. Simpson s.n. (holo: CHR 87407 A&B!; iso: WELT 33375; WELT 33374!). Spreading to decumbent multi-stemmed shrub 50–100 cm tall. Branches: bark on old branches greyish to dark brown, finely fissured, young stems reddish-brown. Leaves erect–spreading; lamina sheath light green to olive-green, 3.5–7.5
2–4(–5) mm, margin membranous, shoulders rounded to auricled, ciliate or only the top half ciliate; lamina light green to olive-green, linear, 18–34(–58)
0.5–1.5(–2.0) mm, adaxial surface minutely rugose with a tuft of scabrid hairs at the base, abaxial surface glabrous; margins serrulate with 60–80 teeth per 10 mm; apex triquetrous and keeled. Inflorescence a terminal, solitary, erect flower on lateral branches, shorter than leaves; flower bract overtopping the flower, leaf-like, coriaceous, linear, (5.5–)7–10(–11)
(0.4–) 0.5–1.0(–1.2) mm; adaxial surface scabrid, abaxial surface glabrous, margin serrulate. Sepals lanceolate, 4.5–9.0
Fig. 81. Known distribution, growth habit and leaf variation in Dracophyllum filifolium. A. Egmont National Park (Venter 13762). B. Mount Perry (Druce s.n.). C. Dun Mountain, Nelson (Venter 13785). D. Mount Anglem, Stewart Island (Venter 13792). All leaves drawn
0.5. 1.5–2.0 mm, equalling corolla tube, surfaces glabrous with the top half pubescent on adaxial surface; margins serrulate. Corolla white; corolla tube cylindrical, widened at mouth, (5.5–)7.0–10.0
(1.5–)2.0–3.0(–4.0) mm, exterior glabrous; corolla lobes reflexed, ovate–triangular to triangular, shorter than corolla tube, 1.4–1.5
1.0–1.2 mm, apices inflexed, acute; apical ridge prominent, adaxial surface papillate. Stamens inserted on corolla tube in upper third, filaments (0.3–)1.0–1.2 mm long; anthers included, oblong, light yellow, 1.0–1.2 mm long. Ovary cylindrical, 2.5–4.5
1.2–2.5 mm, apex truncate; nectary scales rectangular, 1.2–1.5
0.5–0.7 mm, apices subacute to retuse; style included, (2–)3–4 mm long, glabrous; stigma 5-lobed. Fruit sessile, light brown, 4.0–4.2
2–3 mm, broadly obovoid, 94 Australian Systematic Botany S. Venter A F Etymology H G Covered in leaves – refers to the stems that are covered in leaves from the base up to the apex. Diagnostic features and notes I E D C B Fig. 82. Dracophyllum frondosum. A. Flowering branch (1
). B. Sepal adaxial surface (5
). C. Inflorescence-bract adaxial surface (5
). D. Laidout corolla (5
). E. Leaf sheaths adaxial surface to show variation (5
). F. Leaf (2
). G. Lamina apex (5
). H. Ovary (10
). I. Flower (1
). Drawn from Venter 13817. Del. S. Venter. apex truncate, glabrous. Seed brown, ovoid, 0.6–0.7 mm long, testa slightly reticulate (Fig. 82, 83). Distribution and ecology New Zealand endemic restricted to the South Island (Fig. 84). The main distribution is in the Otago area, with a smaller disjunct distribution area in the mountains around the Nelson Lakes area 600 km distant. Dracophyllum frondosum occurs on steep (25–90) riverbanks and on cliffs at elevations ranging from 300 to 900 m. The vegetation consists of open montane forest, shrub or grassland. The soils are brown loam or brown clay loam derived from calcareous sandstone, greywacke or schist. Phenology Flowering December–February. Dracophyllum frondosum is characterised by the lax growth habit, erect–spreading leaves that are 25–50 mm long with ciliate lamina sheaths and a prominent tuft of scabrid hairs at the base on the adaxial surface of the lamina, the solitary flowers, flower bracts longer than the flowers; 7–10 mm long corolla tube that equals the sepals, densely papillate corolla lobes and the cylindrical ovary with truncate apex. Dracophyllum frondosum is similar to D. rosmarinifolium but differs in the lax habit and scrambling stems (Table 10). The flower bract is also longer than the flower and narrower (0.5–1.0 mm compared with 1–2 mm), with the adaxial surface scabrid (compared with glabrous but scabrid at the base). The sepal in D. frondosum equals the corolla tube (compared with being equal or longer) and the adaxial surface texture of the sepal is pubescent in the top half and it is also wider (2–3 mm compared with 1.2–2.5 mm) than the corolla tube; the corolla lobes are shorter (1.4–1.5 mm compared with 2.0–2.5 mm), with longer (1.0–1.2 mm compared with 0.3–0.5 mm) filaments, longer (2.5–4.5 mm compared with 1.7–2 mm) cylindrical ovary with truncate apex and a longer (3–4 mm compared with 1.5–2.5 mm) style. According to Simpson (1952), his var. frondosum was closely allied to D. uniflorum (=D. rosmarinifolium). In the protologue of D. frondosum, he states that the style is exserted for up to 3 mm, but this could not be verified in either the type material or in plants at the localities visited in the wild. Oliver (1952) divided D. uniflorum into two unnamed forms,but did not accept Simpson’s var. frondosum. It was decided to raise this taxon to species level on the basis of the spreading to decumbent growth habit and a combination of flower characters. I place far more value on the flower characters than did Oliver, thus recognising the flower bracts that are longer than the corolla, sepals equalling the corolla in length, adaxial surface of sepals being pubescent in the top half, longer filaments and ovary and the truncate ovary apex as important characters. Dracophyllum frondosum is a species with little variation among populations. Some variation exists in the size of the leaf (18–34
0.5–1.5 mm) and the corolla tube (7–10
2–4 mm). In populations where the slope of thehabitat is gentle to flat, plants tend to have decumbent branches but plants from populations growing in rock cracks in cliffs have hanging branches up to 3 m long (Venter 13823). Representative collections NEW ZEALAND. South Island: Saint Arnaud, upper reaches of the Leatham Valley, 6 May 2000, Venter 13823 (CHR); Red Hills, Motueka River, Dec. 1980, Druce s.n. (CHR); Kauro River headwaters, 9 Oct. 2003, Barkla s.n. (CHR); Palmerston, road over Horse Range, top of limestone cliffs, 4 Dec. 1942, Simpson s.n. (CHR), 19 Feb. 1949, Simpson s.n. (CHR); Dunedin–Middlemarch road at bridge over Deep Stream, Thomson s.n. (CHR); ibid., 5 Feb. 1949, Simpson s.n. (CHR), ibid., 18 Feb. 1957, Allison s.n. (CHR); ibid., 23 Mar. 2000, Venter 13817 (CHR). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 95 A C B D Fig. 83. Dracophyllum frondosum. A. Habitat at the type locality along the Deep Stream. B. Plant growing on a low cliff at the type locality at Deep Stream. C. Mature plant growing on cliff, Leatham Valley showing the typical sprawling growth habit. D. The characteristic erect–spreading leaves. Photos: Danilo Hegg (A), Phil Bendle (B, D) and Cathy Jones (C). Dracophyllum kirkii Berggr., J. Bot. 18: 104 (1880) Type: New Zealand. Mount Torlesse, 1877. S. Berggren s.n. (lecto: O!), designated by Oliver (1952). Dracophyllum uniflorum Bergg., Minneskr. Fysiogr. Sallsk. Lund. 28: 15 (1877). (non Hook.f.) nom. illegit. Illustration S. Berggren, Minneskr. Fysiogr. Sallsk. Lund. 8: t. 4 (1877); A. Eagle, Trees & Shrubs of N.Z., 2nd series: t. 147 (1982). Decumbent to spreading multi-stemmed shrub, 20–140 cm tall. Branches: bark on old branches grey to dark grey, smooth or finely fissured, young stems reddish-brown. Leaves juvenile 96 Australian Systematic Botany S. Venter 170º 175º 180º −35º −40º −45º Fig. 84. Known distribution of Dracophyllum frondosum, South Island, New Zealand. Table 10. Morphological differences between Dracophyllum rosmarinifolium and D. frondosum Character D. rosmarinifolium D. frondosum Branching Flower bract (mm) Flower bract : corolla length Flower-bract adaxial surface Sepal length to corolla-tube length Sepal adaxial surface Sepal size (mm) Corolla-lobe size (mm) Filament length (mm) Ovary length (mm) Ovary apex shape Style length (mm) Upright 5.0–9.5(–13)
1–2 Shorter to equalling Scabrid at the base only Equalling or longer Glabrous (4.5–)5.0–9.0(–12.0)
1.2–2.5 2.0–2.5
1.2–1.5(–2.0) 0.3–0.5 1.7–2.0 Round 1.5–2.5 Spreading to decumbent (5.5–)7–10(–11)
(0.4–)0.5–1.0(–1.2) Longer Scabrid Same length Pubescent in top half 4.5 – 9.0
1.5–2.0 1.4–1.5
1.0–1.2 (0.3–)1.0–1.2 2.5–4.5 Truncate (2–)3–4 Taxonomic revision of Dracophyllum and Richea and adult; juvenile leaves spirally arranged along branches, spreading, glaucous; lamina sheath 9.3–7.5
5.4–8.0 mm, coriaceous, tapering and margin ciliate in upper half; lamina coriaceous, linear–triangular, 42–60
4.0–4.7 mm, surfaces glabrous; margins serrulate with 80–90 teeth per 10 mm; adult leaves spreading, glaucous to occasionally light green; lamina sheath 4–10
3.5–7.0(–9.0) mm, subcoriaceous, striate, shoulders tapering to auricled and margin membranous, smooth to ciliate in the top half; adult lamina linear–triangular, (16–)20–40(–70)
(1.5–)2.0–4.0 mm, prominently striated; margins serrulate with 90–110 teeth per 10 mm. Inflorescence a solitary sessile and erect flower near apices of branches; shorter than leaves; inflorescence bract overtopping flower, glaucous, ovate, 8–20
2.0–2.6 mm, surfaces glabrous, margins ciliate. Sepals ovate, 6.5–7.0
2–3 mm, shorter than to equalling corolla tube; margins ciliate. Corolla white; corolla tube cylindrical, 5–6
2.0–2.5 mm; corolla lobes reflexed, ovate–triangular to triangular, shorter than corolla tube, 1.8–2.0
1.0–1.5 mm, apices acute to subacute; apical ridge present, adaxial surface papillate. Stamens inserted on corolla tube in the upper third, filaments 0.4–0.6 mm long; anthers included, oblong, light yellow and 0.8–1.2 mm long. Ovary cylindrical to ovoid, 1.8–2.0
1.5–2.0 mm, glabrous, apex round; nectary scales rectangular, 1.0–1.5
0.7–0.9 mm, apices retuse; style included, 0.7–2.0 mm long, glabrous; stigma capitate. Fruit light brown, 2.7–3.0
2.3–2.5 mm, broadly obovoid, apex round, glabrous. Seeds light brown, ovoid, 0.95–1.0 mm long, testa slightly reticulate (Fig. 85, 86). Australian Systematic Botany A G 97 H F I E B C D Distribution and ecology Endemic to the South Island of New Zealand, occurring from southern Nelson southward to West Canterbury and North Westland (Fig. 87). Dracophyllum kirkii grows on gentle to steep (5–40) mountain slopes and bluffs at elevations of 700–2166 m. The vegetation consists of subalpine shrubland, tussock grassland, fell field and herbfield. Soils are rocky loam to clay loam derived from granite, greywacke, marble and limestone. Phenology Fig. 85. Dracophyllum kirkii. A. Flowering branch (1
). B. Inflorescence bract (5
). C. Sepal (5
). D. Laid-out corolla (5
). E. Lamina sheaths adaxial surface to show variation (2
). F. Juvenile leaf (1
). G. Adult leaf (1
). H. Ovary (10
). I. Flower (5
). Drawn from Sykes 344/72. Del. S. Venter. Flowering October–April. Etymology Named after Thomas Kirk (1828–1898) New Zealand botanist and teacher. Diagnostic features and notes Dracophyllum kirkii is characterised by its sprawling habit, with the branchlets erect at the tips; smooth, dark grey bark with shallow fissures; leaves glaucous, lamina 20–40
2–4 mm, lamina sheath not much wider than lamina and the lamina base scabrous; flowers solitary; corolla tube 5 mm long and shorter to equalling the sepals. Cheeseman (1906) mentioned its nearest ally as being D. pubescens, which differs in the leaves being pubescent and having 3–5flowered racemes. Dracophyllum kirkii resembles D. pubescens in the general appearance of the glaucous leaves, but they are glabrous, not pubescent, and the inflorescence is a solitary flower, compared with the 3–5flowered spike of D. pubescens. These two species were confused in the past. Leaf size is polymorphic in D. kirkii. Oliver (1928) mentioned that specimens from southern localities show narrower leaves (1.2–2.5 mm) compared with those (4.0–4.5 mm) from northern populations. After extensive fieldwork involving a wide range of populations and having a larger number of dried specimens available for study, I have found that this character breaks down. Representative collections NEW ZEALAND. South Island: Hokonu Range, south of Lake Brunner, Mount French, 19 Dec. 1936, Mackay s.n. (CHR); Hohonu Range, Mount 98 Australian Systematic Botany S. Venter A B C D E F Fig. 86. Dracophyllum kirkii. A. Mature plant in a montane habitat. B. Flowering branches showing the solitary flowers. C. Mature flowering plant. D. The typical short and wide glaucous leaves, Otira Valley. E. Cultivated plant showing the short and spreading adult leaves. F. Scanning electron micrograph of the epicuticular wax on the adaxial lamina surface. Photos: David Lyttle (A–C), Peter Pelser (D) and S. Venter (E, F). French, 18 Nov. 1995, De Lange s.n. (AK); above Arthur’s Pass, Bealy Ridge, 14 Mar. 1972, Sykes 344/72 (CHR); Upper Hurunui River, Nigger Head, 1 Feb. 1973, Macmillan & Stemmer s.n. (CHR); West Amuri, Mount Garfield, 12 Dec. 1936, Brockie s.n. (CHR); Westland National Park, top east of Mount Fox, 28 Feb. 1967, Wardle & Fryer s.n. (CHR); Mariners Peak, 9 Feb. 1978, Wardle & Campbell s.n. (CHR); Mount Reynolds, 16 Feb. 1978, Wardle s.n. (CHR); head of Clarke River, Kea Cliffs, 9 Apr. 1978, Wardle s.n. (CHR); Hooker Valley, Black Birch Stream, Jan. 1898, Cheeseman s.n. (AK); Mount Wilberg, 27 Apr. 1993, Wardle, Buxton & Ford s.n. (CHR); Wanganui River, between Smythe River and Whirlwind Spur, 14 Apr. 1994, Wardle P 94/166 & Buxton (CHR); Mount Barlow, 15 Feb. 1978, Campbell s.n. (CHR); Mount Cook, Sebastopol, 26 Feb. 1958, Connor s.n. (CHR); Mount Cook National Park, Malte Brun, 28 Mar. 1967, Simpson s.n. (CHR); Hooker Valley, Stocking Creek, 25 Feb. 1957, Connor s.n. (CHR); Lake Tekapo, Godley River, Laing s.n. (CHR); Rakaia River, Cascade Hill, 16 Jan. 1985, McGilvary s.n. (CHR); Gorge Plateau, 30 Oct. 1985, Wardle s.n. (CHR); Bealy Spur, 8 Feb. 1991, Douglass P 91/2 (CHR); Woolsack Spur, 1 Apr. 1979, Wardle & Williams s.n. (CHR); Tutoko Valley, Jan. 1977, Johnson s.n. (CHR); Hopkins Valley, Temple Stream, 9 Jan. 1975, Johnson s.n. (CHR); Murchison Mountains, Dana Burn branch of Ettrick Burn, Mar. 1978, McSweeny s.n. (CHR). Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 99 180º −35º −40º −45º Fig. 87. Known distribution of Dracophyllum kirkii. South Island, New Zealand. Dracophyllum lessonianum A.Rich., Essai Fl. Nov. Zel.: 223 (1832) Type: New Zealand. Crescit in rupibus Novae-Zealandiae (Bay of Islands), 1827. J.S.C. Dumont D’Urville s.n. (lecto: P!; isolecto: K!, W!), designated by Oliver (1952). Dracophyllum robustum Hook.f., Flora Antarct. 1: 49 (1844). Type: New Zealand: North Island. J.Edgerley s.n. (holo: K!). Dracophyllum lessonianum A.Rich. var. robustum Hook.f., Fl. Nov. Zel. 2 (1): 170 (1853). Illustrations J. B. Hombron and M. Jacquinot, Bot. Voy. Astrol. & Zel. 2: 85, t. 29 (B) (1833); Eagle, Trees & Shrubs of N.Z., 2nd series: 294, t. 139 (stamens inaccurately illustrated as hypogynous) (1982); Salmon, Native Trees N.Z.: 275 (1989). Erect–spreading single-stemmed shrub or tree 0.5–5 m tall. Branches: bark on old branches grey to dark brown, finely fissured or occasionally deeply fissured on very old stems, young stems reddish-brown. Leaves: juvenile leaves spirally arranged along branches, erect–spreading; lamina sheath yellowish green, 8–17
3.7–5.0 mm; shoulders truncate to auricled and margin ciliate or ciliate in upper half; lamina 100 Australian Systematic Botany coriaceous, linear to linear–subulate, 60–120
1.6–1.8 mm; surfaces glabrous; margin serrulate with 50–78 teeth per mm; adult leaves spreading; lamina sheath light green, 6.0–14
2–4 mm, membranous, shoulders truncate to auricled and margin with the top half ciliate; lamina light to olive-green, linear to linear–subulate, (20–)27–108
0.5–1.2 mm, surfaces glabrous with a tuft of scabrid hairs at the base of the adaxial surface; margins serrulate with 53–70 teeth per 10 mm; apex triquetrous. Inflorescence a terminal spike on lateral branchlets, shorter than the leaves, erect, lax, 20–40 (–50) mm long, linear–oblong; inflorescence bract overtopping flowers, coriaceous to rigid and hard, light to dark green, subulate, 6.0–17.5
0.6–1.0(–3.3) mm; adaxial surface scabrid at the base; abaxial surface glabrous to pubescent at the apex; margins entire. Flowers sessile; flower bract overtopping flowers, leaf-like, coriaceous to rigid and hard, ovate– lanceolate, 8.0–12.5
0.5–0.7 mm, surfaces glabrous with a tuft of scabrid hair at the base of the adaxial surface, margins serrulate and white, apices acute and dark coloured. Sepals lanceolate to ovate–lanceolate, 6–8
1.5–2.0 mm, longer than corolla tube; surfaces glabrous with the top half of the adaxial surface pubescent; margins ciliate; apices acuminate and hard. Corolla white to pinkish; corolla tube cylindrical, widened at mouth, (4–)5–6
2.0–2.5 mm; corolla lobes spreading horizontally, reflexed in old flowers, ovate triangular, shorter than corolla tube, 2.5–3.0
1.0–1.5 mm, inflexed at apex, apices acute; surfaces glabrous. Stamens inserted on corolla tube in upper third, filaments 0.3–0.5 mm long; anthers included, oblong, light yellow and 0.9–1.0 mm long. Ovary oblong, 1.3–1.5
1.2–1.3 mm, apex truncate; nectary scales rectangular, 1.0–1.3
0.5–0.6 mm, apices subacute; style included, 1.5–2.0 mm long, glabrous; stigma capitate. Fruit sessile, light brown, 4.0–4.5
1.7–2.0 mm, oblong, apex truncate, glabrous. Seeds yellowish-brown, ovoid, 1.0–1.2 mm long, testa slightly reticulate (Fig. 88, 89). Distribution and ecology Endemic to the North Island of New Zealand (Fig. 90) and concentrated in the northern part of the North Island with a few scattered localities as far south (38520 latitude) as Taumarunui. Dracophyllum lessonianum occurs on gentle (0–15) south-facing coastal flats and mountain slopes from sea level up to 260-m elevation. The vegetation consists of lowland shrubland and swampy areas along the coast. Dracophyllum lessonianum commonly forms extensive populations in Leptospermum scoparium shrubland and, in some areas, it is the only other woody dominant within this vegetation type. Soils are grey sandy to greyish-brown clay soils derived from sandstone and quartzite. Phenology S. Venter Diagnostic features and notes Dracophyllum lessonianum is characterised by the erect–spreading branches with dark grey and smooth bark, juvenile leaves, adult leaves with auricled lamina sheaths having pale margins, base of the lamina covered in short hairs and a triquetrous apex, racemes that terminate the lateral branches, flower bracts persistent with long narrow sheaths, sepals slightly longer (6–8 mm) than the corolla tube and having hard apices and having the inside covered in white hairs. I agree with Oliver (1928) that D. lessonianum differs from similar species by the long racemes, long acuminate sepals with hard apices and long flowers. Dracophyllum lessonianum is similar to D. filifolium, but differs in having juvenile leaves, sepals longer than the corolla tube, the inflorescence bracts and flower bracts having broad white margins and the apex of the ovary is truncate, not round. Variation in D. lessonianum is in the size of the juvenile leaf sheath (8–17
3.7–5.0 mm), juvenile lamina (60–120 mm long), adult leaf sheath (6.0–12.2
2–4 mm), adult lamina (27–108
0.5–1.2 mm), inflorescence (20–40 mm long) and the inflorescence bract (6.0–17.5
0.6–1.0 mm). Representative collections NEW ZEALAND. North Island: Te Paki Bush, Dec. 1966, Kelly s.n. (CHR); Cape Reinga, Aug. 1962, Woods s.n. (AK); Spirits Bay, Kapowairua, 30 Oct. 1969, Cooper s.n. (AK); Te Paki Station, Radar Bush, 31 Dec. 1954, Moore s.n. (CHR); track 3 km south-west of Kaimaumau, northern side of Kaimaumau Road, 29 Apr. 1990, Brownsey & Fox s.n. (WELT); Karikari Peninsula, Lake Rotokawau, 14 Feb. 1999, Venter 13767 (CHR); Cooper Beach, 16 Dec. 1940, Oliver s.n. (CHR); Ahipara, Reef Point, Sep. 1949, Allan s.n. (O); west of Whangaroa Harbour, Aug. 1967, Druce s.n. (CHR); Kerikeri, Rainbow Falls, 22 Nov. 1991, De Lange 1283 (AK, CHR, WELT); Kerikeri, Puketi Forest, near Forestry HQ, 15 Feb. 1999, Venter 13770 (CHR); Ngawha Springs, 11 July 1970, Lambrechtsen s.n. (CHR); Waipoua State Forest, road to Kawerua, 13 Jan. 1976, Wright 1047 (AK); Waipoua State Forest, near Te Matua Ngahere, 11 Apr. 1972, Rawlings, Esler, Smith & Astridge s.n. (CHR); Kaitaia, Pukemiro, 1 Feb. 1926, Matthews s.n. (AK, CHR); near Lake Taharoa, 15 Jan 1978, Bartlett s.n. (CHR); Ruapekapeka Bay, 29 Aug. 1967, Cooper s.n. (CHR); Whangarei, Sep. 1928, Allan s.n. (CHR); Okahu, 20 July 1905, Carse s.n. (CHR); Matakana, Kirk 519 (CHR); Omaha, 7 Oct. 1940, Moore s.n. (CHR); north of Mangatawhiri, 5 Dec. 1981, Ogle 765 (CHR); Whangamarino, Reao Arm, 9 Oct. 1991, De Lange 1068 (AK, CHR); Hapuakohe Range, Waiti Road, 6 July 1991, De Lange & Crowcroft 854 (WELT); Kopuatai (Piako) peat dome, Oct. 1981, Ogle s.n. (CHR); Kawhia Harbour Road, Upper Tawairoa Stream below Hautapu Hill, 9 Apr. 1988, De Lange s.n. (AK, WAIK). Dracophyllum longifolium (J.R.Forst. & G.Forst.) R.Br. ex Roemer & Schult., Syst. Veg. 4: 385 (1819) Epacris longifolia J.R.Forst & G.Forst., Charact. Gen. Plant.: 20, t. 10 (1776). Flowering November–May. Etymology Named for René Primevère Lesson (1794–1849), a French surgeon and naturalist. Type: New Zealand. Dusky Bay, in woods, 26 Mar. 1773. G. Forster s.n. (holo: K000844563!; iso: BM 77,637!, K000844561!, P 34!). Epacris frondosa Gaertn., Fruct. Sem.1: 77, t. 94 (1791). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany F 101 A H G I B E D C Fig. 88. Dracophyllum lessonianum. A. Flowering branch (1
). B. Inflorescence-bract adaxial surface (5
). C. Sepal abaxial surface (5
). D. Laid-out corolla (5
). E. Lamina sheaths to show variation (2
). F. Juvenile leaf (1
). G. Adult leaf (1
). H. Ovary (10
). I. Flower (5
). Drawn from Venter 13767. Del. S. Venter. Dracophyllum longifolium var. retortum Hombr. & Jacquinot, Voy. Pole Sud.: 86, t. 27 (1833). Dracophyllum longifolium var. pluviale W.R.B.Oliv., Trans. & Proc. Roy. Soc. N.Z. 80 (1): 11 (1952). Type: New Zealand. 1839. M. Hombron s.n. (holo: K!). Type: New Zealand. Caswell Sound, Leslie Valley, 2000 feet [~610 m], 29 Mar. 1949. W.R.B. Oliver s.n. (holo: WELT 313!). Dracophyllum longifolium Sweet, Sweet’s Hortus Britannicus: 488 (1826). nom. illeg. Dracophyllum lyallii Hook.f., Fl. Nov. Zel. 1: 169 (1854). Illustrations Type: New Zealand. Port Preservation and Thompson’s Sound, D. Lyall s.n. (lecto: K!); Dusky Bay, 1791, A. Menzies s.n. (syn: BM000577675!). J. R. Forster and G. Forster, Charact. Gen.: t. 10 (1776); J. Gaertner, Fruct. Sem. Pl. 1: t. 94 (1791); J. D. Hooker, Fl. Antarct. 1: tt. 31, 32 (1844); J. B. Hombron and M. Jacquinot, Bot. Voy. Astrol. Zel.: t. 27 p.p. (1853); T. Kirk, Forest Fl. N.Z.: t. 102 Australian Systematic Botany S. Venter A B D C E Fig. 89. Dracophyllum lessonianum. A. Habitat near Whangaroa Harbour. B. Mature plants in habitat, Kopouatai. C. Plant from Lake Rotokawau on the Karikari Peninsula showing adult leaves (Venter 13767). D. Flowering branch showing the clumped inflorescences, Kopouatai. E. Lectotype specimen housed in Paris. Photos: Davide Artioli (A), John Smith-Dodsworth (B), S. Venter (C, E) and P. de Lange (D). Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 103 180º −35º −40º −45º Fig. 90. Known distribution of Dracophyllum lessonianum, North Island, New Zealand. 109 (1889); J. T. Salmon, Field Guide Alpine Pl. N.Z.: t. 32 (1968); A. Eagle, Trees & Shrubs of N.Z. 2nd series: t. 137 (1982); J. T. Salmon, Native Trees N.Z.: 276 (1989). Erect–spreading single-stemmed shrub or tree 1–12 m tall. Branches: bark on old branches grey to blackish-brown, finely to deeply fissured, young stems reddish-brown. Leaves juvenile and adult. Juvenile leaves spirally arranged or crowded at tips of branches, erect–spreading; lamina sheath light green, (9–)15–20
5–11 mm, shoulders tapering to truncate and margin ciliate in upper half; lamina linear–triangular to lanceolate, 100–250
2.5–8.0(–7.0) mm; margins serrulate with 50–80 teeth per 10 mm. Adult leaves erect–spreading; lamina sheath light green, 5–15
4–7 mm, striate, shoulders rounded to auricled and margin membranous with the top half ciliate; lamina linear to linear–triangular, 40–140(–232)
1–4(–6) mm, prominently striated; margins serrulate with 120–170 teeth per 10 mm; apex triquetrous. Inflorescence a terminal raceme on lateral branchlets, shorter than leaves, initially erect but drooping later, dense, 24–55 mm long, linear–oblong. Inflorescence bract overtopping flowers, subulate, (17–)30–35
1.2–1.5 mm, scabrous at the base of the adaxial surface, margins serrulate. Flowers 5–12(–18), pedicellate; flower bract caducous, equalling or longer than flower, broadly ovate, 10–15
5–6(–8) mm, adaxial surface pubescent in upper third, margins ciliate; pedicel straight, (0.7)1.0–2.0 mm. Sepals ovate–lanceolate, (2.5–)3.0–7.0
(1.5–)2.0–3.0 mm, equalling or longer than corolla tube, striate, surfaces 104 Australian Systematic Botany glabrous with the top half sometimes pubescent on the adaxial surface; margins ciliate. Corolla white; corolla tube cylindrical, widened at mouth, 4.0–5.0
(2.5–)3.0–3.5 mm; corolla lobes reflexed, ovate to ovate–triangular, shorter than corolla tube, 1.5–2.0
(1.5–)2.0–2.5 mm, inflexed at apex; apices subacute; adaxial surface papillate. Stamens inserted on corolla tube in the upper third, filaments 0.3–0.5 mm long; anthers included, oblong, light yellow and 0.8–1.0 mm long. Ovary obovoid, 1.3–2.0 mm long and wide, apex round; nectary scales rectangular, 1.2–1.5
0.8–1.0 mm, apices irregularly toothed; style included, 0.7–1.3 mm long, glabrous; stigma capitate. Fruit pedicellate, light brown, 3.5–3.6
3.8–4.0 mm, obovoid, apex round, glabrous. Seeds light brown, ovoid, 0.8–1.2 mm long, testa slightly reticulate (Fig. 91, 92). S. Venter E A G H F Distribution and ecology Endemic to New Zealand, Stewart Island and its subantarctic Auckland Island and Campbell Island (Fig. 93). Dracophyllum longifolium occurs on mountain and hill slopes, commonly on coastal cliffs and on bluffs from sea level to 1200-m elevation. The vegetation consists of forest, woodland, shrubland and bogs stretching from sea level to the subalpine zone. Soils are grey sandy loam, various types of lithosol, brown clay loam and peaty dark brown loam, derived from calcareous sandstone, greywacke, schist, diorite gneiss or granidiorite. Phenology D C B Flowering October–March. Etymology Refers to the long juvenile leaves. Diagnostic features and notes Dracophyllum longifolium is characterised by the very broad, long and flat juvenile leaves, stiff, narrow and erect adult leaves with wide, shouldered sheaths, flower bracts that fall early; sepals short and broad with long white cilia and the mouth of the corolla tube that is slightly widened. Dracophyllum longifolium is similar to D. cockayneanum, but is instantly distinguished from it by the glabrous juvenile and adult leaves. Further differences are discussed under D. cockayneanum. Matthews (1953) and Cockayne and Phillips-Turner (1967) confused D. longifolium with D. filifolium in stating that it occurs on North Island from the mountains of the East Cape to the Tararua Mountains. Dracophyllum longifolium has up to this point not been recorded for the North Island. Plants from populations near sea level are up to 12 m tall and have branches with a drooping habit, whereas plants from populations in drier and rockier habitats are short, 1–3 m tall and have an erect growth habit (Charleston, Needle Point area, Venter 13750). Shoulders of the leaf sheaths vary from tapering to auricled on the same plant and are clearly illustrated by Kirk (1889). The juvenile lamina varies in size (100–250
2.5–8.0 mm) as does the adult lamina (40–140(–232)
1–4 mm). Plants from populations with short and narrow leaves (Mount Maungatua, Venter 13812) Fig. 91. Dracophyllum longifolium. A. Flowering branch (1
). B. Inflorescence-bract adaxial surface (2.5
). C. Laid-out corolla (5
). D. Lamina sheaths to show variation (2
). E. Juvenile leaf (1
). F. Adult leaf (1
). G. Ovary (10
). H. Inflorescence-bract adaxial surface (2.5
). Drawn from Venter13750. Del. S. Venter. can easily be confused with forms of D. filifolium, but it is readily distinguished on the basis of the broader leaf sheath and the presence of juvenile leaves. Sepal length varies from 3 to 7 mm. Representative collections NEW ZEALAND. South Island: 1 m north of Sandhill Creek, Aug. 1978, Cowin s.n. (CHR); Gouland Downs, Mount Goul, Dec. 1962, Talbot s.n. (CHR); Charleston, Needle Point area, 7 Jan. 1999, Venter 13750 (CHR); north of Karamea, Apr. 1977, Simpson s.n. (CHR); Greymouth, above Roa Coal Mine, 18 Dec. 1965, Moore, Clarke & Robins s.n. (CHR); Hokitika, Crawford River, 17 Dec. 1957, Rockell s.n. (CHR); Arthur’s Pass, Dobson Way, 31 Jan. 1996, Hörandl & Hadacek 7890 (W); Between Knights Point and Haast, Ship Creek, 22 Aug. 1967, Lambrechtsen s.n. (CHR); Ranganui, true left of Moeraki River, 18 Apr. 1978, Campbell s.n. (CHR); Franz Josef Glacier, Alex Knob, 19 Mar. 2000, Venter 13802 (CHR); Mount Wilberg, 27 Apr. 1993, Wardle, Buxton & Ford s.n. (CHR); Perth River, Scone Creek, 17 Feb. 1978, Campbell s.n. (CHR); Mount Cook, Hooker Valley, Stocking Creek, 25 Feb. 1958, Conner s.n. (CHR); Cropp River, 4 Feb. 1983, Basher s.n. (CHR); Mount Peel, Lynn Stream, 15 Jan. 1985, Mayrhofer & Molloy 4821 (GZU); Fiordland, north of Homer Tunnel, Esperance Valley, Mar. 1974, Atkinson s.n. (CHR); Ailsa Mountains, Ocean Peak, 4 Jan. 1936, Zotov s.n. (CHR); Dusky Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 105 A B C D E F Fig. 92. Dracophyllum longifolium. A. Habitat at Arthurs Pass. B. Juvenile plant on side of a road cutting, Arthurs Pass. C. Mature plant with drooping branches. D. Flowering branch showing the early deciduous flower bracts. E. Young plants on a road cutting growing as pioneers, Echo Point. F. Dwarfed mature plant, Mount Dobson. Photos: G. Wilson (A, B), Steve Attwood (C), Diana Bradshaw (D) and S. Venter (E, F). Sound, Cooper Island, Sportsman’s Cove, 7 Feb. 1946, Allan s.n. (CHR); Dusky Sound, Pickersgill Harbour to Lake Forester, 3 Jan. 1969, Dorizac s.n. (CHR); Caswell Sound, Leslie Clearing, 30 Mar. 1949, Zotov s.n. (CHR); Caswell Sound, Large Burn, 2 Apr. 1949, Zotov s.n. (O); Doubtful Sound, Hall’s Arm [Hall Arm], 31 Dec. 1939, Cranwell & Moore s.n. (CHR); Eyre Mountains, upper part of Matuara River, 23 Nov. 1972, Given 72999 (CHR); Dunedin, Mount Maungatua, 23 Mar. 2000, Venter 13812 (CHR); Dunedin, Swampy Hill, 25 Jan. 1948, Allan. s.n. (CHR); Dunedin, Apr. 1874, Berggren s.n. (W); Puysegur Point, 26 Jan. 1946, Allan s.n. (CHR); The Hump, Richardson s.n. (AK); Invercargill, Awarua 106 Australian Systematic Botany S. Venter A B C D Fig. 93. Known distribution, variation in growth habit and adult leaf in Dracophyllum longifolium. A. Charleston, Needle Point area (Venter 13750). B. Between Knights Point and Haast, Ship Creek (Lambrechtsen s.n.). C. Dunedin, Mount Maungatua (Venter 13812). D. Christmas Village, Stewart Island (Venter 13787). Illustrations of leaves all
0.25. Plains, 7 Jan. 1974, Johnson s.n. (CHR); near Mokoreta, Slopedown Range, Catlins State Forest Park, 7 Jan. 1994, Sykes 37/94 (AK, CHR). STEWART ISLAND: eastern coast of Codfish Island, 8 Dec. 1966, Ritchie s.n. (CHR); Mason Bay, Jan. 1940, Attwood s.n. (AK); northwest of Oban, 20 Feb. 1935, Doore & Earle 330 (S); Tamahau [Tamihau] Island, 7 Feb. 1957, Gillham s.n. (CHR). Christmas Village, 11 Jan. 2000, Venter 13787 (CHR); Big South Cape Island [Long Island], 9 Apr. 1961, Bell s.n. (CHR); open country head of Basin Creek, 21 Feb. 1966, Wardle s.n. (CHR). CAMPBELL ISLAND: Camp Cove, 6 Jan. 1961, Godley s.n. (CHR); Tucker Cove, 14 Jan. 1961, Zotov s.n. (CHR). AUCKLAND ISLAND: Mount Eden, 28 Dec. 1962, Godley s.n. (CHR); Above Terror Cove, Feb. 1976, Given 9459 (CHR); crest of scarp above Lake Hinemoa, 20 Feb. 1985, Meurk s.n. (CHR); Ranui Cove, 11 Aug. 1944, Turbott s.n. (AK). Taxonomic revision of Dracophyllum and Richea Dracophyllum marmoricola S.Venter, New Zealand J. Bot. 40 (1): 39–43. (2002) Australian Systematic Botany 107 A Type: New Zealand. Kahurangi National Park, Mount Arthur, Horseshoe Basin, 2 Jan. 1999. S. Venter 13739 (holo: CHR!; iso: AK, K, NSW, P, WELT). Illustrations A. Eagle, Trees & Shrubs N.Z. 2nd edn: t. 152 (1982); J. SmithDodsworth, N.Z. Native Shrubs and Climbers: 34; S. Venter, New Zealand J. Bot. 40 (1): t. 1, 2 (2002). Decumbent to prostrate multi-stemmed shrublet 2–15 cm tall, plants sometimes form compact cushions. Branches: bark on old branches grey, smooth to finely fissured, young branches reddish-brown. Leaves crowded at the tips of branches, spreading, glaucous; lamina sheath 2.2–6.5
3–4 mm, shoulders rounded to truncate, margin membranous, ciliate or with upper half ciliate; lamina linear–triangular to broadly linear–triangular, 5.6–32.0
0.75–2.00 mm, slightly concave, surfaces minutely verrucose, margins serrulate with 12–13 teeth per 10 mm; apex triquetrous. Inflorescence a terminal spike; overtopping the leaves, erect, dense, 9–24 mm long, oblong. inflorescence bracts overtopping flowers, subulate, 1.5–2.0
(0.6–) 1.2–1.4 mm, surfaces verrucose, margins serrulate. Flowers 3–8, sessile; flower bracts shorter than flowers, broadly ovate, 5–6
3.0–3.5 mm, surfaces glabrous, margins ciliate, apices subacute to acute. Sepals green to reddish-brown, ovate–lanceolate, 4.5–5.0
1.5–2.0 mm, equal to shorter than corolla tube, surfaces glabrous; margins ciliate; apices subacute to acute. Corolla white; corolla tube cylindrical, widened at mouth, 4.0–6.5
2.0–2.5 mm; corolla lobes spreading to reflexed, ovate–triangular to broadly triangular, shorter than corolla tube, 1.5–2.2
1.5–1.8 mm, apices acute to subacute; adaxial surface papillate, abaxial surface glabrous. Stamens inserted in corolla tube in the upper third, filaments 0.4–0.5 mm long; anthers included, oblong, light yellow and 0.6–0.8 mm long. Ovary obovoid, 1.4–1.5
1.3–1.5 mm, apex truncate; nectary scales rectangular, 0.7–0.8
0.4–0.8 mm, apices truncate and emarginate to variously toothed; style included, 1.3–1.5 mm long, glabrous; stigma 5-lobed. Fruit sessile, light brown, 2.5–3.5
1.8–2.0 mm, obovoid, apex truncate, glabrous. Seeds light brown, ovoid, 0.7–1.0 mm long, testa slightly reticulate (Fig. 94, 95). Distribution and ecology Endemic to the north-western Nelson area of the South Island, New Zealand (Fig. 96). All known localities are in the Kahurangi National Park. Dracophyllum marmoricola occurs on high mountain slopes and peaks at 1310–1790-m elevation. It grows on dark sandy loam soil in open exposed sites or in rock crevices in low cliffs in alpine tussock-herbfield and grassland. Marble plays an important role in the distribution of calcicoles in New Zealand (Rogers et al. 2018), and Druce et al. (1987) mentioned D. marmoricola (as Dracophyllum sp. a) as a weak calcicole; however, according to the results of my survey, it acts as a strong calcicole, restricted in distribution to shallow soils on B E C D F Fig. 94. Dracophyllum marmoricola. A. Flowering branch (1
). B. Flower (5
). C. Flower bract (5
). D. Laid-out flower (5
). E. Ovary (10
). F. Lamina sheaths to show variation (5
). Drawn from Venter 13739. Del. S. Venter. marble of the Mount Arthur Group (Druce et al. 1987; Venter 2002; Rogers et al. 2018). The substrate specificity of D. marmoricola is as striking as that of Veronica calcicola (Bayly et al. 2001). All Dracophyllum marmoricola populations are covered by snow during winter, yet none of the plants shows the typical red coloration of the leaves during winter that is so common for most Dracophyllum species in New Zealand. Dracophyllum marmoricola is associated with Veronica albicans, Aciphylla aurea, Luzula picta var. picta, Dracophyllum rosmarinifolium, Ranunculus insignis, Coprosma depressa, Leucopogon fraseri, Wahlenbergia pygmaea and Pimelea traversii. Phenology Flowering December–March. Etymology Lover of marble. 108 Australian Systematic Botany S. Venter A B C D E F Fig. 95. Dracophyllum marmoricola. A. Alpine habitat at the type locality on Mount Arthur. B. Mature plant with open growth habit, Mount Arthur. C. Mature plants forming large cushion plants, Mount Arthur D. Flowering branch showing the short and spreading glaucous leaves. E. Flowering plant from the type collection (Venter 13739). F. Mature plant with a prostrate growth habit, Mount Arthur. Photos: S. Venter (A, C, E, F) and Chez Brungraber (B, D). Diagnostic features and notes Dracophyllum marmoricola is characterised by the decumbent to prostrate branches forming cushions; lamina sheath 2.2–6.5
3–4 mm with rounded to truncate shoulders; leaves glaucous, surfaces rugose, 5.6–32
0.75–2.00 mm; 12–13 teeth per 10 mm; inflorescence terminal on branches, dense and oblong, 9–24 mm long; inflorescence bracts verrucose; corolla tube shorter than sepals, 4.0–6.5
2.0–2.5 mm; corolla lobes spreading to reflexed; ovary apex truncate. Dracophyllum marmoricola is similar to D. recurvum in the dense oblong apical racemes and broad flower bracts, but differs in the leaves having far fewer teeth, 12–13 per 10 mm (compared with 90–120) on the lamina margin and the leaf apices not recurved compared with recurved. The flower bracts are also shorter than the flowers (compared with overtopping the flowers), flower bracts glabrous (compared with rugose) and the ovary with a truncate apex (compared with a round apex). Dracophyllum marmoricola superficially resembles D. kirkii. On rock sheets with little soil, plants of D. marmoricola grow as compact cushions, whereas plants hanging down from cliffs can have branches up to 35 cm long. Plants on moderately flat areas in rock rubble grow as extensive mats, sometimes covering areas up to 100 cm in diameter. The Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 109 180º −35º −40º −45º Fig. 96. Known distribution of Dracophyllum marmoricola, South Island, New Zealand. length and width of the leaf sheath is very variable even on a single branch. Most plants have leaves 5.6–15.0 mm long, but those in sheltered areas are usually 18–32 mm long and greener. Plants in protected areas receiving more shade tend to have erect–spreading stems up to 200 mm long, with somewhat larger and greener leaves. Representative collections NEW ZEALAND. South Island: Mount Benson, near Ruby Lake, Druce s.n. (CHR); Lockett Range, rocks above Ruby Lake, Druce s.n. (CHR); Mount Arthur Jan. 1982, Druce s.n. (CHR); ibid., Jan 1975, Druce s.n. (CHR); Mount Arthur, ridge to summit, 16 Jan. 1975, Simpson 7507 (CHR); Mount Hoary Head, Dec. 1980, Druce s.n. (CHR). Dracophyllum minimum F.Muell., Frag. Phytogr. Austr. 1: 39 (1858) Type: Australia. Tasmania. In tergo alpino montis Lapéyrouse pulvinis Pterygoppi Lawrencii innatum. A. Oldfield s.n. (holo: MELB 2064413!). Illustration W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: 685, t. 1 (1928). Forming a compact cushion, 3–16 cm tall and 30–90 cm in diameter. Branches erect-stemmed, many-branched and closely packed. Bark on old branches reddish-brown, smooth, young stems yellowish-brown. Leaves spirally arranged along 110 Australian Systematic Botany branches, imbricate, appressed to stem, olive-green, dry old leaves present; lamina sheath 2.0–3.5
2–3 mm, tapering and margin membranous and ciliate; lamina coriaceous to rigid and hard, linear to linear–triangular, 2–3
0.7–2.0 mm, adaxial surface flat, abaxial surface with a prominent keel, surfaces glabrous; margins serrulate with 40–60 (–80) teeth per 10 mm; apex triquetrous, acute. Inflorescence a terminal solitary, erect, sessile flower, shorter than the leaves; inflorescence bract equalling flower, rigid and hard, dark green, ovate–lanceolate, 4.0–4.5
1.3–1.5 mm, margins serrulate. Sepals lanceolate, 4–6
2.7–3.0 mm, equalling corolla tube; margins ciliate in the upper half. Corolla white; corolla tube cylindrical, (3–)4–6
1.4–1.5 mm; corolla lobes spreading horizontally to sometimes reflexed, ovate–triangular, shorter than corolla tube, 1.5–3.0
1.5–3.5 mm, apices obtuse; margin erose; apical ridge present; adaxial surface papillate. Stamens inserted in corolla in middle of the tube, filaments 0.8–1.0 mm long; anthers included, oblong, deep yellow and 0.6–1.0 mm long. Ovary ovoid, 0.9–1.0
1.0–1.2 mm; apex round; nectary scales rectangular, 0.8–1.0
0.4–0.5 mm, apices bidentate to irregularly toothed; style included, 1.0–1.5 mm long, glabrous, not lengthening in fruit; stigma 5-lobed. Fruit light brown to reddish-brown, 1.5–2.0
2–3 mm, obovoid; apex round, glabrous. Seeds dark brown, ovoid, 0.5–0.6 mm long, testa slightly reticulate (Fig. 97, 98). S. Venter F A E D G H C B Phenology January–February. Distribution and ecology Tasmanian endemic of the subalpine and alpine areas on the central plateau and western mountains, with a single distant locality in the north-eastern areas (Fig. 99). Dracophyllum minimum occurs above the tree line on gentle (0–15) mountain slopes and on mountain plateaux from 300- to 1615-m elevation, especially in areas covered in snow during the winter months. Dracophyllum minimum experiences snow conditions and long periods of low temperatures, with many populations in the bolster heath vegetation under snow for extended periods. It is restricted to subalpine and alpine grassland, heathland and cushionherbfield (bolster heathland). It grows on poorly drained soil, mostly in shallow peat or peat over olive-brown clay which is derived from dolerite. Plants are fully exposed with the habitat receiving up to 2600 mm of rain per annum with added moisture from mist and melting snow. Etymology Refers to the small stature of the cushions, at that stage the smallest of the species. Fig. 97. Dracophyllum minimum. A. Flowering branch (4
). B. Inflorescence bract (5
). C. Laid-out corolla (5
). D. Sepal (5
). E and F leaves to show variation in leaf sheaths (5
). G. Ovary (10
). H. Flower (5
). Drawn from Jarman 134. Del. S. Venter. nectary scales as long as the ovary. Dracophyllum minimum is a very distinct species that shows little variation. There is some variation in the width of the leaf sheath (2–3 mm), in the corolla tube (4–6 mm long) and the corolla lobes (1.5–3.0
1.5–3.5 mm). Dracophyllum minimum is similar to D. muscoides in the growth habit and solitary erect flowers that do not exceed the leaves in length; however, it differs in leaf and flower characters (Table 11). Because of the densely packed leaves, the branches inside are protected from wind and snow and the core of the plant retains a fairly constant temperature (Volkov and Volkova 2015). This form of growth is useful in coping with harsh conditions experienced at higher altitudes (C. Corbett, unpubl. data). Recorded pollinators of D. minimum are flies and Graphium macleayanum (Leach, 1814) (Macleay’s Swallowtail) (Johnson et al. 2012). Diagnostic features and notes Dracophyllum minimum is characterised by its cushion growth habit; branches tightly packed; leaves closely appressed to the stems and 5.0–6.5
0.7–2.0 mm; flowers solitary on the apices of branches with 4–6-mm-long corolla tubes and the Representative collections AUSTRALIA, Tasmania: Cradle Mountain, 28 Jan. 1949, Vickery s.n. (HO); Eldon Bluff, summit plateau, 1987, Buchanan 10010 (HO); top of Mount Ossa, Davies 72 (HO); Mount Rufus, near Lake Saint Clair, 15 Dec. 1917, Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 111 B C A D E Fig. 98. Dracophyllum minimum. A. Habitat in Mount Field National Park. B. Mature plant showing the dense growth of the leaves and the flowers elevated above the plant. C. Flowers showing the horizontal petals with the broad apical ridge. D. Old mature plant showing the cushion growth habit, Cradle Mountain. E. The median ridge on the petal lobes is clearly visible in this image, a common character of the alpine Dracophyllum species. Photos: Tim Rudman (A, C, E), James Wood (B) and Russell Cumming (D). Rodway s.n. (HO); near summit of Mount Rufus, 1983, Buchanan 1157 (HO); Frenchman’s Cap, 20 Dec. 1978, Jarman s.n. (HO); between Reeds Peak and Great Dome, Denison Range, 1993, Buchanan 12992 (HO); Mount Cullen, summit area, 1986, Moscal s.n. (HO); Double Peak, 1978, Jarman s.n. (HO); Mount Eliza, 1985, Moscal 9543 (HO); Mount Field National Park, Tarn Shelf above Lake Seal, 1928, Comber s.n. (HO); Snowdrift Tarns, Snowy Range, 1983, Moscal 2167 (HO); Mount Maconochie, 1978, Jarman s.n. (HO); south-eastern base of Mount Sarah Jane, 1978, Bruhl 633 (HO); Abrotanella Rise, 1998, Buchanan 15170 (HO); between Mount La Perouse and Maxwell Ridge, 1984, Buchanan s.n. (HO); Moonlight Ridge, 1984, Buchanan 3060 (HO); Mount La Perouse, 15 Dec. 1895, Rodway s.n. (HO, NSW). Dracophyllum muscoides Hook.f., Handb. N.Zeal. Fl.: 183 (1864) Type: New Zealand. Alps of Otago, Mount Alta and Hector’s Col, 7000–8000 feet [~2133–2438 m], 1864. J. Hector & J. Buchanan s.n. (holo: K!). The type sheet comprises three different collections made by different people and at different localities. The eight specimens in the top left-hand corner of the sheet are the Hector and Buchanan material and was determined here as such. Illustrations J. Buchanan, Trans. & Proc. N.Z. Inst. 14: t. 26 (1882); A. Eagle, Trees & Shrubs of N.Z. 2nd series: t. 150B (1982); J. SmithDodsworth, N.Z. Native Shrubs & Climbers: tt. 51, 52, pl. 22A, B (1991). Compact cushions, 15–50 mm tall. Branches erectstemmed and much-branched, closely packed together. Bark on old branches greyish-brown, deeply fissured, young stems reddish-brown. Leaves spiralling along branches, imbricate, 112 Australian Systematic Botany S. Venter Fig. 99. Table 11. Known distribution of Dracophyllum minimum in Tasmania. Morphological differences between Dracophyllum. minimum and D. muscoides Character Lamina width (mm) Lamina apex shape Teeth per 10 mm on lamina margin Flower grouping on inflorescence branch Sepal width (mm) Corolla-tube shape Corolla-lobe adaxial surface Filament length (mm) Ovary length (mm) Style length (mm) D. minimum D. muscoides 0.7–2.0 Acute to triquetrous 40–60 >10 0.6–0.8 Obtuse 2.7–3.0 Cylindrical 1.5–2.0 Narrowly campanulate Smooth 0.2–0.5 0.2–0.5 0.8–1.0 Papillate 0.8–1.0 0.9–1.0 1.0–1.5 5–10 5–10 appressed to stem, dry old leaves present; lamina sheath light green, 1.5–3.0 mm long and wide, tapering and margin membranous, ciliate. Lamina coriaceous to rigid and hard, mid- to olive-green, linear to linear–triangular, 1–3
(0.3–) 0.6–0.8 mm; margins serrulate with 5–10 teeth per 10 mm (only the front third of leaf with teeth); apex thickened, obtuse and keeled. Inflorescence a terminal, erect, sessile, solitary flower; shorter than the leaves. Inflorescence bract shorter than flower, ovate–lanceolate, 3.3–3.5
1.5–2.0 mm; margins serrulate; apex obtuse. Sepals ovate–lanceolate, (1.5–) 3.5–4.5
1.5–2.0 mm, equalling corolla tube; margins ciliate; apices subacute to obtuse. Corolla white; corolla tube narrowly campanulate, 2.0–4.5
1.5–3.5 mm; corolla lobes spreading horizontally, ovate–triangular, shorter than corolla tube, 1.0–1.5 mm long and wide, apices obtuse; apical ridge present; inflexed for the entire length; surfaces glabrous. Stamens inserted on corolla tube in the upper third, Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany filaments 0.2–0.5 mm long; anthers included, oblong, initially pink turning light yellow and 0.8–1.0 mm long. Ovary ovoid, 1.4–1.5
1.3–1.4 mm, apex round; nectary scales rectangular, 0.5–0.9
0.5–0.7 mm; apices retuse to irregularly toothed; style included, 0.8–1.0 mm long, glabrous, not lengthening in fruit; stigma 5-lobed. Fruit light brown, 1.0–1.1
0.8–1.0 mm, depressed-globose; apex truncate, glabrous. Seeds yellowish-brown, ovoid, 0.5–0.6 mm long, testa slightly reticulate (Fig. 100, 101). Distribution and ecology Endemic to the South Island of New Zealand from Mount Hercules in the north, southward to the Princess Mountains (Fig. 102). Dracophyllum muscoides grows on gentle (2–10) mountain slopes and mountaintops at elevations of 914–2600 m. The vegetation consists of subalpine cushion herbfield, grassland to alpine bog, fellfield and boulderfield. Soils are commonly boggy, otherwise loam or clay loam derived from greywacke and shale. Plants are fully exposed to severe climatic conditions receiving moisture from rain, mist and melting snow. Phenology Flowering December–May. D D E F C 113 Etymology Likened to a moss cushion. Diagnostic features and notes Dracophyllum muscoides is characterised by the cushion growth habit; leaves closely imbricating and 1–3
(0.3–) 0.6–0.8 mm, widening suddenly into a broad sheath (1.5–3.0 mm); flowers solitary and situated at the apices of the stems, sepals (2.5–)3.5–4.5 mm long and the corolla tube 2.0–2.5 mm long. According to Oliver (1928), its nearest ally is D. prostratum, but it is denser and the leaves are much smaller. There is a note by Cheeseman (1906) that D. muscoides and D. minimum are placed together in the Index Kewensis. However, he noted the difference between the two species after receiving a piece of material from Baron von Mueller from Australia. The type sheet comprises three different collections made by different people and at different localities. The specimens in the top left-hand corner of the sheet are the Hector and Buchanan material. This is a small species and all eight pieces form the type material according to Article 8.2 of the ICN (Turland et al. 2018). Armstrong (1880) mentioned that the corolla is pubescent and the lobes spreading, but no material in subgenus Oreothamnus with a pubescent corolla is known from either New Zealand nor Tasmania. Dracophyllum muscoides is similar to D. prostratum, but differs in the many erect-stemmed branches, deeply fissured bark and fewer (5–10 compared with 10–40) teeth per 10 mm on lamina margin. The flowers are also shorter than the leaves (compared with longer than the leaves), sepals equalling the corolla tube (compared with shorter than the corolla tube) and the corolla tube is shorter (2.0–2.5 mm compared with 3.0–4.5 mm) and narrowly campanulate (compared with cylindrical). Variation in D. muscoides is mostly in the size of the lamina sheath (1.5–3.0 mm long), the lamina (1–3 mm long) and the corolla tube (2.0–4.5
1.5–3.5 mm). Representative collections B G Fig. 100. Dracophyllum muscoides. A. Flowering branch (5
). B. Leaves to show the variation in the lamina sheath (5
). C. Laid-out corolla (5
). D. Inflorescence bract (5
). E. Ovary (10
). F. Flower (10
). G. Sepal (5
). Drawn from Garnock-Jones 2366. Del. S. Venter. NEW ZEALAND. South Island: Harris Mountains, End Peak, 17 Jan. 1950, McNeur s.n. (CHR); Lake Wanaka, Cattle Flat Station, 17 Jan. 1950, Zotov s.n. (CHR); Mount Cardrona, 29 Mar. 1921, Cockayne s.n. (AK); Mount Saint Mary, 21 Jan. 1940, Simpson s.n. (CHR); Pisa Range, Gordon’s Rock [Gordon Rock], 19 Jan. 1947, Langbein 9/28 (CHR); ridge east of Grampians from Hakataramea Pass, 12 Jan. 1961, Connor s.n. (CHR); Hunter Hills, Mount Nessing, 21 Mar. 1980, Given 12384 (CHR); the Hunter Hills, Mount Nimrod summit, 19 Apr. 1979, Given 11687 (CHR); Awakino River, West Branch, 29 Nov. 1994, Wardle 96/78 (CHR); Remarkables, Nevis side, 12 Dec. 1964, Sneddon s.n. (WELTU); Mount Dick, 29 Nov. 1995, Wardle 96/86 (CHR); Alexandra, Old Man Range, 27 Mar. 1919, Cockayne s.n. (CHR); ibid., 1–2 km east of Obelisk, 26 Nov. 1988, Smith-Dodsworth s.n. (AK); Knobby Range near Pinelheugh, 25 May 1995, Patrick s.n. (CHR); Old Man Range, near Roxburgh, 17 Jan. 1968, Elder & Elder s.n. (CHR); Middlemarch, Rock and Pillar Range, 28 Jan. 1985, Mayrhofer, Hertel & Mark 4859 (GZU); ibid., 1 Dec. 1969, Macmillan 69/298 (CHR). 114 Australian Systematic Botany S. Venter A B C D E Fig. 101. Dracophyllum muscoides. A. Habitat on the Old Man Range, D. muscoides is the dominant species here. B. Mature plant from Mount Dobson near Lake Tekapo. C. Mature plant in flower, Rock and Pillar Range. D. Plant in cultivation, showing an atypical open growth habit. E. Watercolour of a plant from the type locality housed at CHR. Photos: Harald Pauli (A), S. Venter (B, D) and David Lyttle (C). Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 115 180º −35º −40º −45º Fig. 102. Known distribution of Dracophyllum muscoides. South Island, New Zealand. Dracophyllum oliveri Du Rietz, Svensk. Bot. Tidskr. 24: 374 (1920) Type: New Zealand. Mount Rochfort, 1800 feet [~548 m], 5 Feb. 1913. D. Petrie s.n. (holo: WELT 33022!; iso: WELT 33021!, WELT 33025!). Illustration R. Du Rietz, Svensk. Bot. Tidskr. 24: t. 24 (1920); W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 7b. (1928); W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 7b (1928). Erect–spreading single-stemmed small tree, 1–4 m tall. Branches: bark on old branches dark grey to dark brown, finely fissured, young stems reddish-brown. Leaves juvenile and adult. Juvenile leaves spirally arranged, spreading to recurved; lamina sheath yellowish- to light green, 6–9
3.0–3.5 mm; shoulders rounded to truncate and margin ciliate in upper half; lamina linear–triangular, 75–85
1.3–1.5 mm; surfaces glabrous, margins serrulate with 60–80 teeth per 10 mm. Adult leaves crowded at tips of branches, spreading; lamina sheath 3.5–9.0
3–5 mm, membranous; shoulders rounded to auricled, margin ciliate or only the top half ciliate; lamina linear to linear–triangular, (30–)50–75
(0.6 –) 1.5–2.0 mm; margins serrulate with (50–)80–100 teeth per 10 mm; apex triquetrous and acute. Inflorescence a terminal raceme on lateral branchlets, shorter than leaves, erect, drooping later, dense, 17–24 mm long, oblong; inflorescence bract 116 Australian Systematic Botany S. Venter overtopping flowers, ovate–lanceolate, (0.6–)13–20
0.8–1.2 mm; margins serrulate. Flowers 5–10, pedicellate; flower bract overtopping flowers, broadly ovate, 3.0–5.5
2.0–3.5 mm, with a tuft of scabrid hairs at the base of the adaxial surface; margins ciliate; pedicels straight, 0.3–0.5 mm. Sepals ovate to triangular, (3–)5–6
(1.3–) 4.5–5.5 mm, equalling corolla tube, adaxial surface with the top half pubescent; margins ciliate. Corolla white, turning light yellow with age; corolla tube cylindrical, widened at mouth, 3.0–4.5(–6.0)
2.0–2.5 mm; corolla lobes spreading horizontally to reflexed, broadly triangular, shorter than corolla tube, 1.5–2.0 mm long and wide; apex inflexed, acute; adaxial surface papillate. Stamens inserted on corolla tube in the upper third; filaments 0.5–0.7 mm long; anthers included, oblong, light yellow, 0.6–0.8 mm long. Ovary ovoid, 1.4–1.5
1.3–1.5 mm, apex round; nectary scales rectangular, 1.0–1.3 (–1.5)
0.6–0.8 mm; apices retuse; style included, 1.3–1.7 mm long, glabrous; stigma 5-lobed. Fruit light to dark brown, 1.0–1.5
1.0–1.3 mm, obovoid; apex round and glabrous. Seed yellowish-brown, filiform, 0.95–1.2 mm long, with a slightly reticulated testa (Fig. 103, 104). G H A E Phenology Flowering November–March. Etymology Named for Dr R. W. B. Oliver (1883–1957), Director of the Dominion Museum in Wellington who completed two major publications on Dracophyllum. Dracophyllum oliveri is characterised by 8–10 teeth per 10 mm on the lamina margin, clustered 5–10-flowered racemes on short lateral branches, sepals 5–6 mm long that are longer than the corolla tube and with light coloured margins. Dracophyllum oliveri is similar to D. filifolium but differs in leaf, inflorescence and flower characters. Oliver (1952) separated D. oliveri from D. longifolium on the basis of the narrower juvenile leaves and Allan (1961) saw D. oliveri as a polymorphic species with caducous flower bracts; however, according to Simpson (1945) and my own observations, they are persistent. On this basis, D. oliveri is here separated from the D. longifolium group in which Oliver (1928, 1952) placed it. The lamina sheath in D. oliveri varies in size (5–9
3–5 mm) and the shoulders of the lamina sheath can either be round or truncate on the same branch. Plants from the Southland populations, especially at Lake Te Anau, are shorter (up to 2 m maximum) with shorter (30–50 mm) more glaucous leaves. The inflorescence bracts vary considerably in size (13–20
0.8–1.2 mm) on the same plant. Flower bracts tend to be small (3
2 mm) in Southland populations compared with material (5.5
3.5 mm) from populations at Mount Rochfort in the north. B D Endemic to the South Island of New Zealand (Fig. 105). Localities are scattered along the western part of the South Island. Dracophyllum oliveri occurs on gentle (0–30) northto north-west-facing mountain slopes, gullies, plateaus and depressions at 157–1160-m elevation. The vegetation consists of open montane forest, woodland, shrubland or tussock grassland. Soils are greyish-brown sand to brown sandy loam derived from sandstone, conglomerate and alluvium, or clay loam along drainage lines. Most of the populations in Southland occur in permanently moist bogs, whereas the northern populations (Hokitika northwards to Karamea) tend to grow in dry conditions (Fig. 105). Diagnostic features and notes I F Distribution and ecology C Representative collections Fig. 103. Dracophyllum oliveri. A. Flowering branch B. Inflorescence bract (5
). C. Flower-bract adaxial surface D. Sepal adaxial surface (5
). E. Lamina sheaths to show variation F. Laid-out corolla (5
). G. Leaf (1
). H. Ovary (10
). I. Flower Drawn from Venter13728. Del. S. Venter. (1
). (5
). (2
). (5
). NEW ZEALAND. South Island: Denniston, 4 July 1965, Wardle s.n. (CHR); Westport, Mount Rochfort, near summit, 25 Nov. 1998, Venter 13728 (CHR); ibid. Venter 13744 (CHR); ibid., Venter 13820 (CHR); ibid., Feb. 1913, Petrie s.n. (AK); Salisbury Open, 31 Jan. 1964, Talbot s.n. (CHR); Bruce Bay, near Lake Kiri, 18 Feb. 1958, Mason & Moar 5365 (CHR); Te Anau, Lake Luxmore, 9 Jan. 1995, De Lange s.n. (AK, HO); Between Lake Te Anau and the Key, 11 Jan. 1937, Simpson s.n. (CHR); Lake Henry, 29 Dec. 1963, Buchanan s.n. (HO). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 117 A B D C E F Fig. 104. Dracophyllum oliveri. A. Habitat at the type locality on Mount Rochfort. B. Flowering stems, near Hector. C. The characteristic drooping inflorescences with the broad flower bracts, from near Hector. D. Young plant from Mount Rochfort. E. Flowering stem from the Black Swamp near Milton showing the widely spreading leaves. F. Mature flowering plant from Mount Rochfort (Venter13744). Photos: S. Venter (A, D, F), Steve Attwood (B, C) and David Lyttle (E). Dracophyllum ophioliticum S.Venter, New Zealand J. Bot. 40 (1): 43–47 (2002) Type: New Zealand. North-western Nelson, Takaka Valley, Kahurangi National Park, Asbestos Creek opposite Chaffey’s Hut, 820 m, 22 Apr. 2000. S. Venter 13822 (holo: CHR!; iso: AK!, BM!, K!, WELT!). Illustration S. Venter, New Zealand J. Bot. 40 (1): tt. 5, 6 (2002). 118 Australian Systematic Botany A S. Venter B C Venter 13728 A B C Venter 13805 Fig. 105. Known distribution, habit and leaf variation in Dracophyllum oliveri. Venter 13728, Westport; Mount Rochfort and Venter 13805, Lake Te Anau, Boundary Creek. A. Adult leaf (1
). B. Juvenile leaf (1
). C. Growth habit. A multi-stemmed shrub, 0.3–1.0 m tall, but reaches 2 m in the shade. Branches decumbent. Bark on old branches grey, finely fissured, young stems yellowish-brown to reddishbrown. Leaves spreading, glaucous to olive-green; lamina sheath 4–9
4–8 mm, striate, shoulders rounded to truncate and margin ciliate; lamina linear–triangular, 21–50
(1.0–)1.3–2.5 mm, slightly concave, surfaces minutely verrucose and covered in short, white scabrid hairs; margins serrulate with 10–13 teeth per 10 mm; apex triquetrous. Inflorescence a terminal raceme on lateral branchlets; shorter than the leaves, erect, dense, (13–)19–23(–28) mm long, oblong; inflorescence bract overtopping flowers, ovate lanceolate, 13.9–22.2
0.7–0.9 mm, surfaces rugose, margins serrulate. Flowers (3–)6–9, pedicellate; flower bracts shorter than flower, keeled, broadly ovate, (4.5–)6.5–8.0
(2.5–) 3–4 mm, adaxial surfaces scabrid in upper third, abaxial surfaces scabrid; margins white, ciliate; pedicel straight, 1.5–2.0 mm long, tomentose. Sepals ovate–lanceolate, (4–) 5–6
(1.5–)2.0–3.0 mm, shorter than corolla tube, adaxial surfaces glabrous with scabrid hairs at apex; abaxial surfaces with scattered scabrid hairs; margins white, ciliate. Corolla white; corolla tube cylindrical, narrowed at mouth, 5.5–6.5
2.0–2.5 mm; corolla lobes reflexed, triangular, shorter than corolla tube, 1.8–2.0
1.5–2.0 mm, apices inflexed, subacute; adaxial surface papillate, abaxial surface glabrous. Stamens inserted on corolla tube in the upper third, filaments Taxonomic revision of Dracophyllum and Richea 0.5–0.7 mm long; anthers included, oblong, light yellow and 1 mm long. Ovary obovoid, 1.3–1.5
1.2–1.5 mm, apex round; nectary scales rectangular, 0.8–1.0
0.6–0.7 mm, apices truncate and emarginate to variously toothed; style included, 1.5–2.0 mm long, glabrous; stigma 5-lobed. Fruit pedicellate, light brown, (2.7–)3.5–4.0
2.5–2.7 mm, obovoid, and apex round, glabrous. Seeds yellowish-brown, ovoid, 0.8–1.0 mm long, testa prominently reticulate (Fig. 106, 107). 119 fragrans, Leptecophylla juniperina, Phyllocladus trichomanoides, Exocarpus bidwillii, Pseudopanax crassifolius, Melicytus alpinus, Gaultheria antipoda, Coprosma microcarpa, C. perpusilla subsp. perpusilla, Veronica masoniae and Dianella nigra. Phenology Flowering January–March. Etymology Distribution and ecology New Zealand endemic restricted to the upper reaches of the Takaka River in the Kahurangi National Park, South Island (Fig. 108). Dracophyllum ophioliticum grows in greyish- to reddish-brown clay loam on steep (10–30) mountain slopes at 457–1000-m elevation in mainly open sites or in rock crevices in montane woodland restricted to serpentinite rocks with asbestos veins. Common associated species include Leptospermum scoparium, Veronica albicans, Phormium cookianum subsp. cookianum, Griselinia littoralis, Astelia A Australian Systematic Botany B C D E Growing on serpentinite. Diagnostic features and notes Dracophyllum ophioliticum is a multi-stemmed shrub up to 1 m tall with decumbent stems, glaucous leaves that are minutely verrucose and covered with scabrid hairs when young, pedicellate flowers, flower bracts shorter than the flower and keeled with the abaxial surface scabrid, sepals shorter than corolla tube, mouth of the corolla tube narrowed and the ovary obovate. Dracophyllum ophioliticum is similar to D. filifolium in the spreading linear–triangular to filiform leaves and persistent flower bracts. It differs in being a many-stemmed shrub with decumbent to prostrate stems (compared with erect–spreading or erect stems), the verrucose surfaces of the leaves with scabrid hairs (compared with glabrous with a tuft of scabrid hairs at the base), sepals shorter than the corolla tube (compared with equalling the corolla tube) and the mouth of the corolla tube narrowed, not widening. In the past, it was thought by most people that D. ophioliticum was merely a form of D. longifolium, but it is easily distinguished by the absence of large juvenile leaves and caducous flower bracts, characters that Oliver (1928, 1952) regarded as diagnostic for D. longifolium. Dracophyllum ophioliticum is an endemic of ultramafic soils. The scarcity of endemics on the ultramafics in the South Island has been attributed to the short time in which these habitats have been available to plant life since the most recent glaciation (Lee 1992; Wardle 2002). Most D. ophioliticum plants growing on the unstable and easily weathered serpentinite form prostrate stems of up to 4 m long. These stems commonly root at the nodes anchoring the stems; this appears to be an adaptation to survival in the unstable soil conditions. Plants grown by the author under garden and glasshouse conditions maintain this growth habit. The leaves of D. ophioliticum are glaucous and short (21–29 mm long) in exposed areas, but greener and longer (30–50 mm long) in protected areas. The leaf sheath varies considerably in length and width (4–9
4–8 mm), with the shoulder ranging in shape from rounded to truncate. Representative collections Fig. 106. Dracophyllum ophioliticum. A. Flowering branch (1
). B. Flower (5
). C. ovary (10
). D. Laid-out flower (3
). E. Adaxial surface of the lamina sheaths to show variation (5
). Drawn from Venter 13800. Del. S. Venter NEW ZEALAND. South Island: Cobb Valley, below dam, Nov. 1979, Druce s.n. (CHR); ibid., Jan. 1987, Hayward s.n. (CHR); Takaka Valley, Asbestos Creek, Nov. 1979, Druce s.n. (CHR); Summit of Asbestos Hill, 25 Dec. 1950, Hay s.n. (CHR); Asbestos Creek opposite Chaffey’s Hut, 22 Apr. 2000, Venter 13822 (CHR). 120 Australian Systematic Botany A S. Venter B C D E Fig. 107. Dracophyllum ophioliticum. A. Habitat at the type locality near Chaffey’s Hut in the upper reaches of the Takaka River. B. Habitat in the Takaka River Valley. C. Plant showing the characteristic prostrate stems. D. Fruiting branch. E. Plant at the type locality showing the characteristic widely spreading leaves on the trailing stems (C–E Venter 13822). Photos: Jan Robertson (A) and S. Venter (B–E). Dracophyllum palustre Cockayne ex W.R.B.Oliv., Trans. & Proc. N.Z. Inst. 59: 690 (1928) Type: New Zealand. Near Kumara, Feb. 1877. T. Kirk s.n. (lecto: AK 7028!; isolecto: WELT 44737!, WELTU 4275!), designated by Oliver (1952). Dracophyllum uniflorum var. virgatum Cheeseman, Man. N.Z. Fl.: 427 (1906); Dracophyllum virgatum (Cheeseman) Cockayne, Trans. & Proc. N.Z. Inst. 44: 53 (1912). nom. illeg., non Colenso (1896). Type: New Zealand. Westland, swamp near Lake Brunner. L. C [ockayne] s.n. (holo: WELT 33156!). Illustration A. Eagle, Trees & Shrubs of N.Z., 2nd series: t. 142 (1982). Multi-stemmed scrambler or subshrub, 20–100 cm tall. Branches spreading to decumbent. Bark on old branches dark grey, smooth, young stems reddish-brown to purplish-brown. Leaves erect–spreading, olive-green; lamina sheath 2.5–6.0
2.2–4.0 mm, tapering to truncate and margin membranous, ciliate; lamina linear, 14–27
0.5–1.0 mm, adaxial surface flat, occasionally minutely scabrous with a tuft of scabrid hairs at the base; margins serrulate with 70–110 teeth per 10 mm; apex keeled, triquetrous. Inflorescence a solitary terminal, sessile flower on shortened lateral branchlets; shorter Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 121 180º −35º −40º −45º Fig. 108. Known distribution of Dracophyllum ophioliticum, South Island, New Zealand. than leaves; inflorescence bract equalling flower, light to dark green, ovate–lanceolate, 4.0–4.5
1.5–2.0 mm, adaxial surface pubescent at apex; margin ciliate; apex obtuse to acute; flower bracts overtopping flowers, ovate–lanceolate to ovate, 5.0–5.5
2.0–2.5 mm, surfaces glabrous with a tuft of scabrid hairs at the base of the adaxial surface; margins prominently white and ciliate; apices acute to slightly obtuse. Sepals ovate to triangular, 4.0–5.5
1–2 mm, equalling corolla tube, glabrous with the top half of the abaxial surface pubescent; margins ciliate; apices acute. Corolla white to light pink; corolla tube cylindrical, 3.5–5.0
2.0–2.2 mm; corolla lobes reflexed, ovate–triangular, shorter than corolla tube, 1.6–3.0
1.3–2.2 mm, apices inflexed, acute; adaxial surface papillate. Stamens inserted on corolla tube in the upper third, filaments 0.5–0.7 mm long; anthers included, oblong, light yellow and 0.7–0.9 mm long. Ovary obovoid, 0.7–1.0(–1.2)
0.6–1.0 mm, apex truncate; nectary scales rectangular, 0.8–1.5
0.6–0.7 mm, apices retuse; style included, 1.0–1.2 mm long, glabrous; stigma 5-lobed. Fruit light brown, 3.5–4.0
2.8–3.0 mm, obovoid, apex round to slightly truncate, glabrous. Seeds yellowish-brown, ovoid, 0.95–1.0 mm long, testa slightly reticulate (Fig. 109, 110). Distribution and ecology New Zealand endemic, common in the northern half of the South Island (Fig. 111). Dracophyllum palustre occurs from sea level to an elevation of 1280 m. It grows exposed in boggy, swampy or permanently moist habitats. The surrounding vegetation is lowland to montane shrubland, tussock 122 Australian Systematic Botany E S. Venter A F D G C H I B Fig. 109. Dracophyllum palustre. A. Flowering branch (1
). B. Laid-out corolla (5
). C. Lamina sheaths to show variation (5
). D. Lamina apex (5
). E. Leaf (1
). F. Ovary (10
). G. Flower (5
). H. Sepal abaxial surface (5
). I. Flower-bract adaxial surface (5
). Drawn from Mason and Moar 5539. Del. S. Venter. grassland or peat bog. Soils are mostly boggy or humus-rich loam derived from greywacke. Phenology Flowering (October–)December–May(–July). Dracophyllum palustre is similar to D. rosmarinifolium in the solitary flowers and narrow short leaves, but differs in the slender strict habit, small leaves and the small flowers that are arranged on short peduncles. It is also similar to D. subulatum, but differs in the absence of juvenile leaves, flowers solitary, not in spikes, with the inflorescence bract equalling (compared with overtopping) the flower and having a ciliate, not serrulate, margin. The sepal is longer (4.0–5.5 mm compared with 2.6–3.2 mm) with the top half of the abaxial surface pubescent. The corolla tube is also much larger (3.5–5.0
2.0–2.2 mm compared with 1.8–2.0
1.0–1.2 mm) with larger nectary scales (0.8–1.5
0.6–0.7 mm, not 0.4–0.5
0.3–0.5 mm) and fruit (3.5–4.0
2.8–3.0 mm compared with 2.9–3.0
1.7–1.8 mm). The diagnostic broad and pale to white flower-bract margin is similar to that of D. subulatum. The lamina sheath (2.5–6.0
2.2–4.0 mm) and lamina (14–27
0.5–1.0 mm) show some variation in size. The adaxial surface of the lamina is mostly glabrous, but some plants have a prominent patch of scabrid hair at the base of the lamina. The flower bract can vary from ovate–lanceolate to ovate and the sepals from ovate to triangular on the same plant. Populations found in areas from a high elevation have small (1.6
1.3 mm) corolla lobes, becoming gradually larger (3.0
2.2 mm) in plants growing closer to sea level. Representative collections NEW ZEALAND. South Island: Perry’s Saddle, Jan. 1969, Druce s.n. (CHR); Gouland Downs Scenic Reserve, Jan. 1975, Kelly & Kelly s.n. (CHR); peak north of Mount Goul, Apr. 1981, Druce s.n. (CHR); Denniston Plateau, Jan. 1981, Druce s.n. (CHR); Westport, upper Waimangaroa River, 28 Sep. 1976, Given 9577 & Molloy (CHR); Mount Augustus, 7 Jan. 1976, Moore s.n. (CHR); Westport, Mount Rochfort, 30 Oct. 1998, Venter 13727 (CHR); Westport, German Terrace, 19 Nov. 1985, Courtney P33 (CHR); Mount William, Townson s.n. (CHR); Matiri Range, Mount Misery, Mar. 1981, Druce s.n. (CHR); Buller River, Lyall, Dublin Terrace, 22 Nov. 1985, Courtney P34 (CHR); south of Reefton, Fossicker Creek basin, Gardner 6908 (AK, WAIK); 4 m north-west of Lake Brunner, Kangaroo Lake, 30 Mar. 1950, Potter s.n. (CHR); Rahu Saddle, 17 Feb. 1983, Van Balgooy 4469 (CHR, L); Westport, 31 Dec. 1897, Townson s.n. (WELT); Spencer Mountains, Ada Pass Saddle, 8 Jan. 1945, Brockie s.n. (CHR); Maruia Valley, Thompsons Flat, 19 Nov. 1990, Johnson & Buxton s.n. (CHR); Westland National Park, Skiffington Swamp, 12 Oct. 1965, Wardle s.n. (CHR); Mount Greenlaw, Jan. 1924, Wall s.n. (CHR); Gorge River north of Jerry Gorge junction, 3 Nov. 1985, Johnson 465 & Wardle (CHR); Forbes Mountains, Mount Earnslaw, Feb. 1992, Druce 1720 (CHR). Etymology Lover of swampy areas. Dracophyllum patens W.R.B.Oliv., Trans. & Proc. N.Z. Inst. 59: 698 (1928) Diagnostic features and notes Dracophyllum palustre is characterised by the long and slender stems that are sparingly leafy; leaves 14–27
0.5–1.0 mm, lamina apices bluntly pointed and the shoulders of the lamina sheaths rounded and ciliate; flowers solitary on the lateral branches; flower bracts broad with a very broad pale margin and corolla tube 3.5–5.0 mm long with acute lobes. Type: New Zealand. Great Barrier Island, summit of Mount Hobson, 8 Dec. 1916. W.R.B. Oliver s.n. (holo: WELT 33296!). Illustration W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 9 (1928); A. Eagle, Trees and Shrubs of N.Z. 2nd series: t. 136 (1982). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 123 A C B D E Fig. 110. Dracophyllum palustre. A. Habitat in Lewis Pass. B. Flowering plant, Lewis Pass. C. Flowering branch showing the clumped flowers and the leaves that are present only at the stem apices. D. Mature plant with a shrub-like growth habit. E. The more common prostrate growth habit for the species. Photos: S. Venter (A–E). Multi-stemmed shrub 0.5–3 m tall. Branches erect. Bark on old stems dark grey to greyish-brown, smooth, young stems yellowish- to reddish-brown. Leaves juvenile and adult. Juvenile leaves spirally along branches, erect–spreading; lamina sheath 10–12
8–14 mm, rounded to truncate and margin ciliate in upper half; lamina linear–triangular; 80.0–126.4
5.0–7.1 mm, adaxial surface sparsely scabrid; abaxial surface glabrous; margin serrulate with 30–40 teeth per 124 Australian Systematic Botany S. Venter 170º 175º 180º −35º −40º −45º Fig. 111. Known distribution of Dracophyllum palustre, South Island, New Zealand. 10 mm. Adult leaves erect–spreading; lamina sheath light green to light brown, 4.5–12.0
4–10 mm, striate, tapering to truncate and margin membranous, top half ciliate; lamina broadly linear–triangular, 30–75
2–6 mm, adaxial surface covered in minute scabrid hairs, becoming denser towards the apex, slightly striated; margin serrulate with 45–70 teeth per 10 mm. Inflorescence a terminal spike on lateral branchlets, shorter than leaves, drooping, dense, (16.5–)35–40 mm long; inflorescence bract overtopping flowers, coriaceous, ovate–lanceolate, 4.5–6.0(–7.0)
1.2–1.5 mm, adaxial surface pubescent to sericeous; margin serrulate. Flowers 5–8, sessile; flower bract longer than flower, broadly ovate; apex subulate; 5–8
2.8–3.0 mm; adaxial surface with a tuft of scabrid hairs at the base; margins ciliate. Sepals ovate–lanceolate, 4–8
1.5–2.7 mm, longer than the corolla tube, adaxial surface pubescent in the top half; margins ciliate. Corolla white; corolla tube narrowly campanulate, widened at mouth, 4–5
2.3–2.5 mm; corolla lobes spreading horizontally to reflexed, broadly triangular, shorter than corolla tube, 2.5–3.0
2.3–2.5 mm, apices subacute; surfaces glabrous. Stamens inserted at the top of the corolla tube, filaments (0.25–)0.5–1.0 mm long; anthers included, oblong, light yellow and 1.0–1.2 mm long. Ovary subglobose, 1.0–1.1 mm long and wide; apex round; nectary scales rectangular, 1.2–1.5
0.5–0.7 mm, apices subacute; style included, 1.0–1.4 mm long, glabrous, lengthening in fruit; stigma 5-lobed. Fruit light brown, 2.5–3.5
2.5–3.0 mm, obovoid, apex round, glabrous. Seeds Taxonomic revision of Dracophyllum and Richea yellowish-brown, ovoid, 1.1–1.2 mm long, testa slightly reticulate (Fig. 112, 113). Australian Systematic Botany 125 brown loam to clay loam. Most populations occur in full sun, sometimes with several plants growing in light shade of the forest (Mount Rowe, Coromandel). Distribution and ecology Endemic to the Coromandel area and Great Barrier Island off the North Island of New Zealand (Fig. 114). Dracophyllum patens grows exposed in permanent moist to seasonally moist areas on mountain peaks, plateau areas and cliffs at elevations from 300 to 795 m. The vegetation consists of open lowland to montane forest, woodland or shrubland. The soils are dark D Phenology Flowering April–June(–November). Etymology Named for the leaves that spread nearly 90 with the stem axis. A F E G H C B J I Fig. 112. Dracophyllum patens. A. Flowering branch (1
). B. Laid-out corolla (5
). C. Lamina sheaths to show variation (1
). D. Juvenile leaf (1
). E. Adult leaf (1
). F. Ovary with nectary scales (10
). G. Inflorescence-bract adaxial surface (5
). H. Flower-bract adaxial surface (5
). I. Sepal adaxial surface (5
). J. Flower (5
). Drawn from Venter 13771. Del. S. Venter. 126 Australian Systematic Botany S. Venter A C B D E Fig. 113. Dracophyllum patens. A. Habitat on top of Mount Rowe on the Coromandel Peninsula. B. Adult plant showing the erect branches. C. Fruiting branch. D. The characteristic finely fissured bark. E. Flowering branch showing the small flowers with acute corolla apices at the type locality on Great Barrier Island. A, C and D Venter 13771. Photos: S. Venter (A–D) and John Braggins (E). Diagnostic features and notes Dracophyllum patens is characterised by the erect stems with short (38–75 mm) leaves that are broad (2–6 mm) at the base, broad lamina sheaths (5.5–12.0
4–10 mm) with scarious margins and with some short hairs on shoulders, juvenile leaves much larger than the adult leaves (80–110
5–7 mm), five- or six-flowered inflorescences, broad flower bracts, corolla tube 4–5
2.3–2.5 mm and nectary scales overtopping the ovary. A species easily distinguishable from other members of subgenus Oreothamnus in the long erect stems, narrow long juvenile leaves and much broader and shorter, strongly spreading adult leaves. Size variation occurs in the juvenile Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 127 180º −35º −40º −45º Fig. 114. Known distribution of Dracophyllum patens, northern part of the North Island, New Zealand. lamina (80–126
5–7 mm), the adult lamina sheath (4.5–12.0
4–10 mm with the shoulders tapering to truncate), adult lamina (30–75
2–6 mm) and the sepals (4–8
1.5–2.7 mm). Representative collections NEW ZEALAND. North Island: Great Barrier Island, upper slopes of Mount Hobson, 25 Oct. 1976, Bartlett s.n. (AD, AK, L, NZRFI); Ahumata (White cliffs), 15 Dec. 1938, Molesworth s.n. (AK); Coromandel Range, Papakai Plateau, 8 Apr. 1983, Gardner 3465 (AK); Coromandel Forest Park Table Mountain, Apr. 1971, Druce s.n. (CHR); ibid, Jan 1977, Bartlett s.n. (CHR); Coromandel, Kauaranga Forest Park, The Pinnacles, Apr. 1971, Druce s.n. (CHR); ibid., 31 Aug. 1982, Clarkson s.n. (CHR); Thames, Upper Kauaranga River, June 1977, Bartlett s.n. (CHR); Coromandel, Kauaranga Forest Park, top of Mount Rowe, next to Trig. Beacon, 16 Feb. 1999, Venter 13771 (CHR); Coromandel Range, Pakirarahi, 7 June 1977, Bartlett s.n. (AK, CHR); ibid. May 1977, Bartlett s.n. (CHR). Dracophyllum pearsonii Kirk, Trans. & Proc. N.Z. Inst. 17: 223 (1885) Type: New Zealand. Stewart Island, W.L. Pearson s.n. (holo: WELT 33309!). Illustration A. Eagle, Trees and Shrubs of N.Z. 2nd series: t. 148 (1982). 128 Australian Systematic Botany Multi-stemmed shrub 30–50 cm tall. Branches: bark on old branches dark grey, deeply fissured, young stems reddishbrown. Leaves spirally arranged along branches, imbricate, appressed to stem, dry old leaves present; lamina sheath olivegreen to light brown, 4–7
3–5 mm, coriaceous, striate, shoulders rounded to truncate with margins membranous, ciliate or with only the top half ciliate; lamina rigid and hard, mid- to olive-green, linear to linear–subulate, 19–32
0.8–1.5 mm, adaxial surface flat, abaxial surface keeled, margins serrulate with 90–100 teeth per 10 mm, apex triquetrous. Inflorescence a few-flowered spike near apices of branches; shorter than leaves, erect, lax, 12–20 mm long, oblong. Inflorescence bract overtopping flowers, leaf-like, coriaceous, linear, ovate–lanceolate at the base, 6.0–6.5
4–5 mm, surfaces glabrous with a tuft of scabrid hairs at the base of the adaxial surface, margins ciliate. Flowers 3–6, sessile; flower bracts overtopping flowers, leaf like, linear, 6.0–6.5
4–5 mm, with a tuft of scabrid hairs at the base of the adaxial surface, margins ciliate. Sepals lanceolate to ovate–lanceolate, 4.2–4.5
1.5–2.0 mm, shorter or equalling corolla tube; margins ciliate. Corolla white; corolla tube cylindrical, 4.2–4.5
1.8–2.0 mm; corolla lobes reflexed, ovate–triangular to broadly triangular, shorter than corolla tube, 1.9–2.0
1.4–1.5 mm, apical ridge present, apices inflexed at tip, subacute; adaxial surface papillate. Stamens inserted on corolla tube near the top, filaments 0.2–0.5 mm long; anthers included, oblong, light yellow and 0.8 mm long. Ovary globose, 1.9–2.0
1.8–2.0 mm, apex round; nectary scales oblong, 1.0–1.2
0.8–1.0 mm, apices obtuse; style included, 0.8–1.0 mm long, glabrous; stigma 5-lobed. Fruit light brown, 1.0–1.5
1.0–1.2 mm, obovoid, apex truncate, glabrous. Seeds yellowish-brown, ovoid, 0.55–0.6 mm long, testa slightly reticulate (Fig. 115, 116). Distribution and ecology New Zealand endemic. Restricted in distribution to a small area in the Fiordland National Park, South Island, but widely distributed on Stewart Island (Fig. 117). Dracophyllum pearsonii occurs on gentle (0–15) north- and east-facing mountain slopes, ridges and plateaux from 300- to 1000-m elevation. The vegetation of these areas consists of montane to subalpine shrubland, herbfield or grassland. The soil is a brown gritty sandy loam derived from granite and granidiorite. Phenology Flowering December–February. Etymology Named after Joseph Pearson (1821–1901), stockman and explorer of the Waimakariri Valley. Diagnostic features and notes Dracophyllum pearsonii is characterised by the imbricate leaves that densely cover the branches and the dead leaves that remain on the branches for a long period, flowers in S. Venter A E F D G C H B I Fig. 115. Dracophyllum pearsonii. A. Flowering branch (1
). B. Laid-out corolla (5
). C. Lamina sheaths to show variation (5
). D. Lamina apex (5
). E. Leaf (1
). F. Ovary (10
). G. Flower (5
). H. Inflorescence-bract adaxial surface (5
). I. Sepal (5
). Drawn from Venter13790. Del. S. Venter. few-flowered racemes near the ends of branches and a prominent apical ridge on the upper surface of the petal. Dracophyllum pearsonii is a very distinct species and is easily distinguished from other species by the imbricate and strongly keeled leaves with triquetrous apices, dry old leaves that remain on the plant for a long period and the few-flowered (3–6 flowers) inflorescence. The leaves can be very dense or somewhat openly arranged on the branches. The leaf sheath varies in size (4.0–7.0
3–5 mm) and can sometimes have an area of dense scabrid hairs at the base of the lamina. Lamina length varies from 19 to 32 mm, even on the same plant. Representative specimens NEW ZEALAND. South Island: Secretary Island, 7 Feb. 1964, Wardle s.n. (CHR); Dusky Sound, Facile Harbour, 9 Feb. 1946, Allan s.n. (CHR); Fiordland National Park, near West Cape, east of Lake Fraser, 9 Feb. 1972, Mark s.n. (CHR). Stewart Island: hill north of Smiths Lookout, Jan. 1952, Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A B C D E F 129 Fig. 116. Dracophyllum pearsonii. A. Habitat on slopes of Mount Anglem. B. Plateau area of Mount Anglem with scattered plants of D. pearsonii. C. Adult plant showing the prostrate habit of the stems. D. Mature plant. E. Branch showing the persistent leaves. F. The short and erect leaves. Photos: James H. (A, B) and Rowan Hindmarsh-Walls (C–F). Findlay s.n. (CHR); Table Hill Track, 12 Jan. 1940, Cranwell & Moore s.n. (CHR); Mount Anglem, high ridge below summit, 11 Jan. 2000, Venter 13790 (CHR); Mount Rakeahua, 12 Jan. 1940, Cranwell s.n. (CHR); hill north of Smiths Lookout, Jan. 1952, Findley s.n. (CHR); Pegasus Bay, Aston s.n. (AK, CHR). Dracophyllum politum (Cheeseman) Cockayne, Rep. Bot. Surv. Stewart Island: 43 (1909) Dracophyllum rosmarinifolium var. politum Cheeseman in Man. N.Z. Fl.: 427 (1906). 130 Australian Systematic Botany S. Venter 170º 175º 180º −35º −40º −45º Fig. 117. Known distribution of Dracophyllum pearsonii, southern part of the South Island with Stewart Island. Type: New Zealand. Mount Maungatua near Dunedin, 3000 feet [~914 m]. D. Petrie s.n. (lecto: AK 7033!; isolecto: WELT 33366!), designated by Oliver (1952). Illustrations W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: 687, t. 3 (1928); A. Eagle, Trees and Shrubs of N.Z. 2nd series: 296, t. 151 (1982). Cushion-forming, up to 50 cm tall and 100 cm in diameter, to a scrambler 2–25(–50) cm tall. Branches spreading to prostrate, much-branched. Bark on old stems grey to brown, broadly fissured, young stems reddish-brown. Leaves imbricate, appressed to stem, erect, olive- to dark green, dry old leaves present. Leaf sheath 2.5–4.0
3–4 mm, shoulders tapering to round and margin membranous, ciliate. Lamina rigid and hard, linear, 3.5–12.0(–17.2)
0.7–1.5 mm, adaxial surface flat; surfaces glossy, margins serrulate with 90–100 teeth per 10 mm; apex obtuse or occasionally subacute. Inflorescence a sessile, solitary terminal flower; shorter than leaves, erect. Flower bract shorter than flower, leaf like, broadly ovate to triangular, 2–3
0.7–0.8 mm, margins serrulate, apices obtuse. Sepals ovate–lanceolate, 4.8–5.0
1.8–2.0 mm, longer than corolla tube, adaxial surface with top half pubescent; abaxial surface glabrous; margins ciliate. Corolla white; corolla tube cylindrical, widened at mouth, 3.5–4.5
1.4–1.5 mm; corolla lobes spreading horizontally to reflexed, ovate–triangular, shorter than corolla tube, 1.5–2.2
1.0–1.8 mm, apical ridge present, inflexed at apex, obtuse; Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany adaxial surface papillate. Stamens inserted on corolla tube in the upper third, filaments 0.5–1.0 mm long; anthers included, oblong, light yellow and 0.9–1.0 mm long. Ovary ovoid, 1.2–1.8
1.2–1.3 mm, apex round; nectary scales rectangular, 0.8–1.2
0.5–0.7 mm, apices irregularly toothed; style included, 1.0–1.1 mm long, glabrous, not lengthening in fruit; stigma 5-lobed. Fruit dark brown, 2.5–3.0
1.5–2.5 mm, oblong and ridged at the ribs; apex round, glabrous. Seeds dark brown, ovoid, 0.68–0.7 mm long, testa prominently reticulate (Fig. 118, 119). Distribution and ecology New Zealand endemic restricted to the South Island and Stewart Island (Fig. 120). Dracophyllum politum grows fully exposed on gentle (0–15) mountain slopes, especially on mountain peaks and plateaus at altitudes of 500–1524 m. These areas are covered in subalpine to alpine shrubland, herbfield, fellfield, grassland or bog. Soils are dark brown peaty soil, gritty sandy loam or light brown clay derived from granidiorite, greywacke, granite, sandstone or conglomerate. Dracophyllum politum sometimes grows in permanently wet areas on Mount Maungatua and in bogs on the Table Hill D A E C F 131 Range (Stewart Island); here, they are found growing closely together with Donatia novae-zelandiae Hook.f. and Oreobolus pectinatus Hook.f., forming single cushions. Etymology Refers to the polished and glossy surface of the leaves. Diagnostic features and notes Dracophyllum politum is a dense cushion plant or a prostrate shrublet with densely imbricated leaves appressed to the branch. The lamina is very glossy (hence, the specific epithet) and convex, slightly curved inwards and with obtuse apices. Flowers are solitary and the flower bracts have blunt apices. The corolla lobes that are inflexed have prominent apical ridges. Dracophyllum politum is similar to D. densum, but differs in leaf and flower characters. The plant habit is polymorphic from dense round cushions 30 cm tall to plants with sprawling stems and carpet-like. The leaves vary in size (3.5–12.0
0.7–1.5 mm), with there being minimal variation in shape. Representative specimens NEW ZEALAND. South Island: peak north of Mount Goul, Jan. 1973, Druce s.n. (CHR); Denniston, 6 Jan. 1968, Brownlie 665 (CHR); Stockton Plateau, 16 July 1979, Given 11834 (CHR); Mount Augustus, 7 Jan. 1976, Moore s.n. (CHR); summit of Mount Rochford, 14 Nov. 1966, Kelly & Kelly s.n. (CHR); Westport, Mount Rochfort, Denniston Plateau, 25 Nov. 1998, Venter 13730 (CHR), ibid., Venter 13819 (CHR); ibid., Venter 13826 (CHR); Burnett’s Face, Jan. 1913, Petrie s.n. (AK); below Garibaldi Ridge, Apr. 1980, Druce s.n. (CHR); Mount Earnslaw, Feb. 1992, Druce 1720 (CHR); Wilmott Pass, Pahiri Peak, 25 Dec. 1944, Simpson s.n. (CHR, WELT). Stewart Island: top of Mount Anglem, 11 Jan. 2000, Venter 13789 (CHR); Thompson Ridge, 19 Feb. 1962, Wardle s.n. (CHR); unnamed hill north of Smith’s Lookout, 1 Jan. 1952, Findley s.n. (CHR); Port Pegasus, Nov. 1907, Aston s.n. (AK). Dracophyllum pronum W.R.B.Oliv., Trans. & Proc. N.Z. Inst. 59: 686 (1928) G Type: New Zealand. Blimit Mountain, above Arthur’s Pass, above 6000 feet [~1828 m], 29 Jan. 1928. W.R.B. Oliver s.n. (lecto: WELT 33206!; isolecto: WELT 33349!, WELT 33354!, WELT 33354 dupl.!, WELT 33359!), designated by Oliver (1952). Dracophyllum rosmarinifolium Hook.f., Fl. Antarct. 1: 50 (1844) nom. illeg. non R.Br. B H Dracophyllum rosmarinifolium Cheeseman, Man. N.Z. Fl.: 427 (1906) nom.illeg. non R.Br. Dracophyllum muscoides Armstrong, Trans. & Proc. N.Z. Inst. 13: 342 (1881) nom.illeg. non Hook.f. Dracophyllum rosmarinifolium Betts, Trans. & Proc. N.Z. Inst. 51: 155 (1919) nom.illeg. non R.Br. Fig. 118. Dracophyllum politum. A. Flowering branch (1
). B. Laid-out corolla (5
). C. Leaf sheaths to show variation (5
). D. Leaf (1
). E. Ovary (10
). F. Flower (5
). G. Inflorescence bract (3
). H. Sepal adaxial surface (6
). Drawn from Venter 13789. Del. S. Venter. Illustrations W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 2 (1928); W. R. B. Oliver, Trans. & Proc. Roy. Soc. N.Z. 80 (1): t. 2 (1952); 132 Australian Systematic Botany S. Venter B A C E D F Fig. 119. Dracophyllum politum. A. Habitat on Mount Anglem, Stewart Island. B. Mature plant from Mount Anglem, showing the cushion growth habit. C. Very large and old plant, Mount Anglem, Stewart Island. D. Plant showing the broadly fissured bark on mature stems. E. Mature plant in flower, Rocky Mountain, Stewart Island. F. Plant in cultivation keeps the cushion growth habit. (B, D and F Venter 13789). Photos: James H. (A), S. Venter (B–D, F), Colin Meurk (E). A. Eagle, Trees and Shrubs N.Z. 2nd series: t. 149 (1982); J. Smith-Dodsworth, N.Z. Native Shrubs & Climbers: t. 53, pl. 22C, D (1991); J. T. Salmon, Native N.Z. Flowering Plants: 216, t. 899 (1991). Scrambler or subshrub (1–)10–25 cm tall. Branches decumbent to prostrate. Bark on old branches grey and smooth, sometimes with deep fissures in very old specimens, young stems reddish-brown. Leaves spreading; lamina sheath 0.7–4.0
1–3 mm, shoulders tapering to rounded and margin membranous, ciliate. Lamina linear, 2.5–11
0.5–1.0 mm, adaxial surface flat, with a tuft of scabrid hairs at the base; margins serrulate with 100–140 Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 170º 175º 133 180º −35º −40º −45º Fig. 120. Known distribution of Dracophyllum politum. teeth per 10 mm; apex obtuse to acute. Inflorescence a sessile, terminal, solitary flower on lateral branchlets; shorter than leaves; inflorescence bract shorter than flower, coriaceous, ovate at the base, 3.2–4.1
0.6–0.8 mm, margins serrulate; flower bract shorter than flower, leaf-like, ovate, 3.5–4.0
0.6–0.8 mm; margins serrulate. Sepals ovate–lanceolate, (1.7–) 4.0–4.5
1.4–1.6 mm, equal to longer than corolla tube; margins ciliate. Corolla white to light pink; corolla tube cylindrical, 2.5–4.0
1.5–1.8 mm; corolla lobes reflexed, ovate–triangular, shorter than corolla tube, (1.0–)1.5–2.0
(1.0–)1.5–2.0 mm, apex inflexed, subacute; apical ridge present, adaxial surface papillate. Stamens inserted on corolla tube in the middle, filaments 0.5–1.0 mm long; anthers included, oblong, light yellow and 0.8–1.0 mm long. Ovary ovoid, 0.5–1.0
0.5–1.2 mm, apex round; nectary scales rectangular, 0.5–0.8
0.5–0.7 mm; apices retuse; style included, 1.0–1.5 mm long, glabrous; stigma capitate. Fruit light brown, 1.2–2.5
1.5–2.0 mm, oblong, glabrous, apex truncate. Seeds yellowish-brown, ovoid, 0.46–0.5 mm long, testa slightly reticulate (Fig. 121, 122). Distribution and ecology New Zealand endemic restricted to the South Island (Fig. 123). Dracophyllum pronum occurs on gentle to steep (5–40) mountain slopes, mountain saddles and exposed rocky ridges at elevations of 600–2000 m. These areas are 134 Australian Systematic Botany D A S. Venter E F C G B H Hooker (1844) included D. pronum under Forster’s Epacris rosmarinifolia and Cheeseman (1906) suspected that his D. rosmarinifolium was different from that of Forster; however, not being able to get hold of the type, he made no alteration. Dracophyllum pronum is also quite distinct from D. politum and, for this reason, Oliver (1928) gave it the new name D. pronum. Dracophyllum pronum is similar to D. palustre, but differs in the leaves being shorter (2.5–11 mm compared with 14–27 mm) and the lamina apex acute, not triquetrous. The inflorescence bract is shorter than the flower (not equalling) and far narrower (0.6–0.8 mm compared with 1.5–2.0 mm) with serrulate green, not white, margins. The corolla tube is also narrower (1.5–1.8 mm compared with 2.0–2.2 mm) and the apical ridge on the corolla lobe is absent in D. palustre. Dracophyllum pronum is a variable species, influenced by its habitat. Populations in exposed alpine areas (Mount Torlesse) grow as a prostrate plant closely hugging the ground, with leaves 2.5–3.2
0.5–0.6 mm and the lamina sheath 0.7–2.0
1–2 mm (Venter 13755). Populations in more protected areas at a lower altitude (Dun Mountain), grow as roundish shrublets of 30 cm tall, with leaves 3–4
0.7–1.0 mm and the lamina sheath 2.5–4.0
2.5–3.0 mm (Venter 13786). The sepals can equal the length of the corolla tube or can be longer, with the corolla tube varying from 2.5 to 4.0 mm long on the same plant. Light pink flowers sometimes occur scattered throughout populations. Representative specimens Fig. 121. Dracophyllum pronum. A. Flowering branch (1
). B. Laid-out corolla (5
). C. Lamina sheaths to show variation (5
). D. Leaf (1
). E. Ovary (10
). F. Flower (5
). G. Inflorescence bract (5
). H. Sepal (5
). Drawn from Venter 13786. Del. S. Venter. covered in montane to subalpine shrubland, shrub–tussockland, herbfield to alpine herbfield, fellfield, and bogland or tussock grassland. Soils are brown to reddish-brown gritty to rocky clay loam derived from greywacke, serpentinite, granite or gneiss. Dracophyllum pronum is one of the few plant species to survive on mountain peaks where the snow lies in shaded areas for up to 5 months of the year. Phenology Flowering December–February(–April). Etymology Describes the prostrate growth habit of the plant. Diagnostic features and notes Dracophyllum pronum is characterised by the decumbent to prostrate stems, flat adaxial lamina surface, flower bract shorter than the corolla tube and with a serrulate margin, sepals equal to longer than the corolla tube, a prominent apical ridge on the corolla lobe and the retuse nectary-scale apices. NEW ZEALAND. South Island: Gouland Downs, Jan. 1969, Druce s.n. (CHR); Lead Hills, 29 Dec. 1949, Moore s.n. (CHR); Lake Sylvester, 15 Jan. 1962, Melville 6036, Melville & Talbot (AK, CHR); Beeby’s Knob, Jan. 1952, Talbot s.n. (CHR); Mole Tops, 13 Feb. 1964, Simpson 4053 (CHR); Nelson, Dun Mountain, Dun Saddle, 28 Sep. 1999, Venter 13786 (CHR); Mount Baldy, Mar. 1984, Druce s.n. (CHR); Red Hills, Motueka River, Dec. 1980, Druce s.n. (CHR); Paparoa Range, east of Mount Priestly, 30 May 1983, Wardle s.n. (CHR); West Glenroy Range, Glenroy Valley, 15 Jan. 1985, Burke 274 (CHR); Mount Haast, 21 Jan. 1994, Wardle 96/48 (CHR); Mount Technical, Feb. 1937, Simpson s.n. (CHR); Spencer Range, Waiau Pass, 12 Jan. 1970, Simpson 5701 (CHR); Mount Sebastopol, 26 Feb. 1958, Connor s.n. (CHR); Mount Saint Patrick, 14 Jan. 1972, Canterbury Botanical Society Camp s.n. (CHR); Jollies Pass, 7 Feb. 1938, Zotov s.n. (CHR); Mac’s Knob, 30 Jan. 1973, Macmillan 73/77 & Stemmer (CHR); Ant Creek, Jan. 1934, Wall s.n. (CHR); summit of Mount Burnette, 3 May 1964, Dawson s.n. (CHR); Two Thumb Range, The Growler, Feb. 1991, Druce APD222 (CHR); Mount Potts, Dec. 1918, Cockayne s.n. (CHR); Mount Sinclair, Mar. 1986, Druce s.n. (CHR); Fox Peak, Mar. 1985, Druce s.n. (CHR); Mount Cockayne, Craigieburn Basin, 30 May 1973, Chapman s.n. (AD, BREM, CANB, E, K, L, MO, NBG, PRC, RSA, TI, WISCON); Mount Ida, 7 Jan. 1911, Petrie s.n. (CHR); Christchurch, Porters Pass, Foggy Peak, 28 Jan. 1999, Venter 13755 (CHR); Mount Torlesse, Apr. 1909, Travers s.n. (L, W, Z); Mount Peel, 16 Jan. 1985, Mayrhofer & Molloy 4793 (GZU); Mount Hutt, 6 Apr. 1972, Molloy s.n. (CHR); Cascade River, east of Woodhen Creek, 12 Mar. 1978, Wardle, Lee & Johnson s.n. (CHR); Torlesse Range, Fog Peak, 14 Feb. 1948, Oliver s.n. (WELT); Crown Range, 7 km south-east of Arrowtown, 26 Feb. 1962, Melville 6551 (AK, L); Pisa Range, Gordon’s Rock, 19 Jan. 1947, Langbein 9/14 (CHR); Saint Marys Range, Awakino Ski field, 19 Dec. 1987, Rance s.n. (CHR); Mount Dick, 25 Nov. 1995, Wardle 96/85 (CHR). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany B A D C E F Fig. 122. Dracophyllum pronum. A. Habitat on greywacke, Porters Pass. B. Flowering branch, Porters Pass. C. Shrubby growth habit of the Dun Mountain plants growing on serpentinite (Venter 13786). D. Flowering branches, Porters Pass. E. Prostrate alpine form, Porters Pass. F. Prostrate growth habit of plants from alpine habitats, Mount Dobson. Photos: S. Venter (A–F). 135 136 Australian Systematic Botany 166ºE S. Venter 167ºE 168ºE 169ºE 170ºE 171ºE 172ºE 173ºE 174ºE 175ºE 40ºS A 41ºS 42ºS 43ºS 44ºS 45ºS 46ºS 47ºS B Fig. 123. Known distribution, variation in growth habit and leaves of Dracophyllum pronum. A. Nelson, Dun Mountain (Venter 13786). B. Mount Peel (Mayrhofer and Molloy 4793). All leaves drawn
7. Dracophyllum prostratum Kirk, Trans. & Proc. N.Z. Inst. 13: 384 (1881) Type: New Zealand. Otago, mountains above Lake Harris, Longwood Range, 4000 feet [~1220 m], among Sphagnum, 11 Jan. 1877, W.T. Kirk s.n. (lecto: WELT 32884!; isolecto: BM 577656!, CHR 332686!, K!, WELT 32882!, WELT 32883!, WELT 32891!), designated by Oliver (1952). Illustrations A. Eagle, Trees and Shrubs N.Z., 2nd series: t. 150A (1982); A. F. Mark and N. M. Adams (1986: t. 47). A scrambler, sometimes forming cushions 1–10 cm tall. Branches prostrate. Bark on old branches dark grey to blackish-brown, smooth, young stems reddish-brown. Leaves spirally arranged along branches, erect to appressed to the stem, glaucous to light green, old leaves present; lamina sheath 1.5–3.0
2–3 mm, shoulders tapering to rounded and margin membranous, ciliate; lamina linear to linear–triangular, 2.5–5.0(–7.3)
0.5–0.9(–1.0) mm, adaxial surface flat to slightly concave, abaxial surface keeled; margins serrulate with 10–40 teeth per 10 mm (only at the apex,); apex obtuse to acute. Inflorescence a sessile, solitary terminal flower; longer than leaves, erect. Inflorescence bract shorter than flower, ovate–lanceolate, 3.6–3.8
1.8–2.0 mm; margin serrulate; apex obtuse. Sepals lanceolate, (3.5–)4.0–4.5
1.5–2.0 mm, shorter than corolla tube; margin ciliate. Corolla white; corolla tube cylindrical, 3.0–4.5
2.0–2.5 mm; corolla lobes reflexed, ovate–triangular, shorter than corolla tube, 1.5–2.0 mm long and wide; apex obtuse; inflexed for entire length, apical ridge present, adaxial Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany surface papillate. Stamens inserted on corolla tube in the upper third, filament 0.2–0.3 mm long; anthers included, oblong, light yellow and 1.0–1.2 mm long. Ovary obovoid, 1.0–1.5
0.8–1.0 mm, apex round; nectary scales rectangular, (0.5–) 0.7–0.8
0.4–0.5 mm; apices retuse to irregularly toothed; style included, 0.9–1.0 mm long, glabrous; stigma capitate. Fruit reddish-brown, 1.5–2.0
1.4–1.5 mm; obovoid, apex truncate, glabrous. Seeds light brown, ovoid, 0.45–0.7 mm long, testa slightly reticulate (Fig. 124, 125). 137 habitats. Plants are fully exposed to the harsh weather conditions and are permanently moist in seepages or seasonally moist after melting snow, rain or mist. Phenology Flowering December–February. Etymology Distribution and ecology Describes the prostrate growth habit of the plant. New Zealand endemic. Restricted in distribution to the South Island, with all known localities being situated south of a line from Arthur’s Pass to Christchurch (Fig. 126). Dracophyllum prostratum occurs on gentle to steep (5–80) mountain slopes and on plateaux at elevations ranging from 350 to 1828 m. The vegetation consists of subalpine shrubland to alpine herbfield, fellfield, tussockland, bogland and cushion field or short grassland. The soil is gritty greyish-brown sandy loam derived from schist, diorite gneiss or greywacke, with a thin (1–5 mm) humus layer that can be as thick as 15 mm in moist D A E C F Diagnostic features and notes Dracophyllum prostratum is characterised by its prostrate habit, dark brown and smooth bark, erect and clasping leaves being 3–5 mm long, short and broad (1.5–3.0
2–3 mm) lamina sheaths with short cilia, solitary flowers with the sepals equalling or longer than the corolla tube. Kirk (1881) is correct in saying that it is a variable species in habit, but the floral characters are quite stable. He mentioned in the protologue that the apices of the sepals are obtuse, but those on the type are acute. Dracophyllum prostratum is similar to D. muscoides, but differs in the more prostrate branches with smooth bark, leaves with more (10–40 compared with 5–10) teeth per 10 mm on the lamina margin and the flowers that are longer than the leaves. The sepals are shorter than the corolla tube, not equalling, and the apex is acute, not subacute, to obtuse. The corolla lobes are longer (1.5–2.0 mm compared with 1.0–1.5 mm) and with papillate adaxial surfaces that are not glabrous. The ovary is obovate, not ovate, and narrower (0.8–1.0 mm compared with 1.4–1.5 mm), with the fruit being longer and wider. Most specimens have glaucous leaves; however, there are a few scattered populations with green leaves that are also wider and longer. The adult lamina varies in size (2.5–5.0
0.5–0.9 mm), being small in alpine localities (Lake Harris) and larger in populations at lower altitudes (Mount Maungatua). Representative specimens B H G Fig. 124. Dracophyllum prostratum. A. Flowering branch (1
). B. Laidout corolla (5
). C. Lamina sheaths with lamina showing variation (10
). D. Leaf (1
). E. Ovary (10
). F. Flower (5
). G. Inflorescence bract (5
). H. Sepal (5
). Drawn from Garnock-Jones 2367. Del. S. Venter. NEW ZEALAND. South Island: Takitimu Mountains, Mount Hamilton, 16 Jan. 1938, Barker s.n. (CHR); Two Thumb Range, head of Bush Stream, Mar. 1986, Druce s.n. (CHR); Fiordland National Park, Lake Harris, 27 Feb. 1911, Petrie s.n. (CHR, WELT); ibid., Kirk s.n. (AK); Fiordland National Park, 2 km south of Key Summit, 15 Jan. 1997, Hörandl & Hadacek 8113 (W); Livingstone Range, key summit, 24 Dec. 1944, Oliver s.n. (AK, WELT); Livingstone Range, above Knobs Flat, 3 Jan. 1962, Wardle s.n. (CHR); Mount Pisa, 15 Jan. 1950, Zotov s.n. (CHR); Danseys Pass, 27 Apr. 1969, Moore s.n. (CHR); Murchison Mountains, Ettrick Burn, Mar. 1978, McSweeney s.n. (CHR); Lake Hauroko, South Caroline Burn, Dec. 1975, Sutcliffe, Craighead & Williams s.n. (CHR); Mount Cuthbert, Laing s.n. (CHR); above Green Lake, 7 Jan. 1967, Given s.n. (CHR); Ben Lomond, 2 Jan. 1936, Zotov s.n. (CHR); Windley Branch of Eyre Creek, 1970, Given 70463 (CHR); Mid Dome, 9 Feb. 1961, Connor s.n. (CHR); Dunedin, Mount Maungatua, 23 Mar. 2000, Venter 13810 (CHR); ibid. Venter 13815 (CHR); ibid., Petrie s.n. (AK, Z). 138 Australian Systematic Botany S. Venter A B C D E Fig. 125. Dracophyllum prostratum. A. Habitat at the type locality near Lake Harris. B. The solitary flowers showing clearly the medial ridge characteristic of the subalpine and alpine Dracophyllum species. C. Mature plant in a subalpine habitat. D. Prostrate form growing in a wetland habitat. E. Plant showing the bare branches with leaves borne at the apices of the stems and the solitary flowers. Photos: Daniel Pietzsch (A), David Lyttle (B–D) and L. and A. Stridvall (E). Dracophyllum pubescens Cheeseman, Man. N. Zeal. Fl.: 426 (1906) Type: New Zealand. Mountains near Westport, 1500–2000 feet [~457–610 m], W. Townson s.n. (lecto: AK 211641!; isolecto: AK 7004!), Designated by L. Cranwell, July 1942. Illustrations A. Eagle, Trees and Shrubs of N.Z. 2nd series: t. 146 (1982); A. F. Mark and N. M. Adams, N.Z. Alpine Plants: t. 46 (1986). Multi-stemmed shrub 50–80 cm tall. Branches decumbent to prostrate. Bark on old branches grey to dark grey, deeply fissured, young stems reddish-brown. Leaves juvenile and adult, glaucous. Juvenile leaves spirally arranged along branches, spreading; lamina sheath 10–13
6–8 mm; shoulders rounded, with margin ciliate in the upper half; lamina linear–triangular to lanceolate, 60–80
5–7 mm, surfaces pubescent; margins serrulate with 60–70 obscured teeth per 10 mm. Adult leaves crowded at tips of branches, spreading, glaucous; lamina sheath (3.3–)4.5–7.0
(3.5–) 4.5–6.0 mm, striate, shoulders rounded to truncate and margins membranous, ciliate; lamina linear–triangular to lanceolate, 14–54(–65)
2–6 mm, adaxial surface densely pubescent to tomentose; abaxial surface sparsely pubescent; prominently striated; margins ciliate with 80–100 obscure Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 139 180º −35º −40º −45º Fig. 126. Known distribution of Dracophyllum prostratum, South Island, New Zealand. teeth per 10 mm. Inflorescence a terminal spike on lateral branchlets; shorter than leaves, erect, lax, 15–17 mm long, oblong; inflorescence bract overtopping flowers, glaucous, ovate–lanceolate at the base, (7.4–)14.9–21.8
1.6–2.5 mm, pubescent; margin ciliate. Flowers 3–5, sessile; flower bract shorter to equalling flower, ovate–lanceolate, 9.5–13.0
1.0–2.5 mm, adaxial surface pubescent; abaxial surface with sparse scabrid hairs; margin ciliate. Sepals ovate–lanceolate to ovate, 5.0–5.5
1.3–2.0 mm, shorter than corolla tube, surfaces glabrous on the top half, pubescent on adaxial surface; margin ciliate. Corolla white; corolla tube cylindrical, 5–6
1.6–2.0 mm; corolla lobes spreading horizontally to reflexed, triangular, shorter than corolla tube, 1.5–2.0 mm long and wide; apices inflexed, acute; adaxial surface papillate. Stamens inserted on corolla tube in the upper third, filament 0.5–0.8 mm long; anthers included, rectangular, light yellow and 0.8–1.0 mm long. Ovary oblate, 1.4–1.5 mm long and wide, glabrous, apex truncate; nectary scales separate, rectangular, 0.8–1.0
0.7–1.0 mm, apices irregularly toothed; style included, glabrous, 1.5–2.0 mm long; stigma capitate. Fruit light brown, 1.5–2.0 mm long and wide, obovoid; apex truncate, glabrous. Seeds brown, filiform, 0.7–1.0 mm long, testa slightly reticulate (Fig. 127, 128). Distribution and ecology New Zealand endemic restricted to the north-western Nelson area of the South Island, with most of the known localities in Kahurangi National Park (Fig. 129). Dracophyllum pubescens 140 Australian Systematic Botany D A S. Venter E F C G B sepals 5.0–5.5 mm long and pubescent on the outside and the 5–6 mm long corolla tube. Dracophyllum pubescens is similar to D. kirkii, but differs in the prominent pubescence of the lamina. The racemes are three- to five-flowered, whereas it is a solitary flower in D. kirkii. Variation in D. pubescens is mostly in the size of the adult lamina sheath (4.5–7.0
4.5–6.0 mm), the adult lamina (14–54
2–6 mm), inflorescence bract (15–22
1.6–2.5 mm) and the flower bract (9.5–13.0
1.0–2.5 mm). Selected specimens NEW ZEALAND. South Island: above Mount Perry Saddle, 31 Jan. 1964, Hynes s.n. (AK); Gouland Downs, Dec. 1962, Talbot s.n. (CHR); Saxon River, 25 Mar. 1949, Richardson s.n. (AK); Mount Haidinger, 4 Jan. 1969, Soper s.n. (CHR); Lead Hills, Feb. 1935, Simpson s.n. (CHR); edge of Boulder Lake, 6 Jan. 1962, Hynes s.n. (AK); Mount Bovis, Jan. 1951, Talbot s.n. (CHR); Mount Frederick, 4 Mar. 1912, Morgan s.n. (AK, WELT); ibid., Townson 208 (AK); Burnett’s Face, 7 Jan. 1976, Simpson 7754 (CHR); Twenty Four Tarn Basin, Herbert Range, Feb. 1987, Druce s.n. (CHR); Karamea, Bald Hill, 17 Feb. 1985, McLennan s.n. (CHR); Herbert Range, Twenty Four Tarn Basin, Feb. 1987, Druce s.n. (CHR); Cobb Valley, Mount Mytton, 11 Dec. 1998, Venter 13736 (CHR); Cobb, Lake Sylvester, 23 Mar. 1999, Venter 13774 (CHR); Mount Williams, 30 Apr. 1912, Morgan s.n. (CHR). Dracophyllum recurvum Hook.f., Fl. Antarct. 1: 50 (1844) H Type: New Zealand. Mount Tongariro, 1839, J.C. Bidwill 65 (holo: K!). Dracophyllum rubrum Colenso, Trans. & Proc. N.Z. Inst. 20: 200 (1888). Fig. 127. Dracophyllum pubescens. A. Flowering branch (1
). B. Laidout corolla (5
). C. Lamina sheaths to show variation (2
). D. Adult and juvenile leaves (1
). E. Ovary (10
). F. Flower (5
). G. Inflorescencebract abaxial surface (5
). H. Sepal adaxial surface (5
). Drawn from Venter13774. Del. S. Venter. Type: New Zealand. Base of Mount Ruapehu, 1879. Colenso s.n. (holo: WELT 23615!). grows fully exposed on cliffs, gentle (5–35) slopes and bluffs at elevations of 500–2500 m. These areas are covered in montane to subalpine shrubland, herbfield, fellfield or grassland. Soils are mostly brown clay lithosols or clay loam lithosols derived from serpentinite, greywacke, granite, sandstone or conglomerate. These areas receive precipitation in the form of rain and often in the form of mist and melting snow. Type: New Zealand. High up on Mount Ruapehu, 1889. H. Hill s.n. (holo: WELT 23614!). Phenology Flowering December–March. Etymology Dracophyllum tenuicaulis Colenso, Trans. & Proc. N.Z. Inst. 22: 476 (1890). Dracophyllum brachyphyllum Colenso, Trans. & Proc. N.Z. Inst. 28: 604 (1896). Type: New Zealand. Ruahine Mountains, 1895. H. Hill s.n. (holo: WELT 23617!). Dracophyllum brachycladum Colenso, Trans. & Proc. N.Z. Inst. 31: 275 (1899). Type: New Zealand. Eastern side of Ruahine Mountains, 1898. H. Hill s.n. (holo: WELT 23618!) Refers to the pubescent leaves. Illustrations Diagnostic features and notes Dracophyllum pubescens is easily recognised by the grey bark with broad fissures, presence of juvenile leaves, the glaucous, broad and pubescent leaves, few-flowered (1–)3–4 racemes, T. F. Cheeseman, Illustr. N. Zeal. Fl.: t. 131 (1914); A. Eagle, Trees and Shrubs of N.Z., 2nd series: t. 133 (stamens inaccurately illustrated as hypogynous) (1982); J. Smith-Dodsworth, N. Z. Native Shrubs & Climbers: t. 56, pl. 23C, D (1991). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 141 A B C D E Fig. 128. Dracophyllum pubescens. A. Habitat on Mount Mytton. B. Mature plant on top of Mount Mytton. C. Flowering branch. D. Branch showing the spreading short leaves. E. Plant covered in snow. (B and D, Venter 13736). Photos: S. Venter (A, B and D, E) and S. Aubert and R. Douzet (C). Many-stemmed shrublet 10–90 cm tall. Branches spreading, decumbent to prostrate and much-branched. Bark on old branches grey to dark grey, smooth, young stems reddish-brown. Leaves spreading to mostly recurved, glaucous to light green. Lamina sheath 4–6
3.0–6.5 mm, striate, tapering to truncate and margin membranous, ciliate or only the top half ciliate; lamina linear to linear–triangular, 15–30(–40)
1–2 mm, adaxial surface rugose to scabrid, abaxial surface glabrous, slightly striated; margin serrulate with 90–120 teeth per 10 mm; apex thickened, obtuse and triquetrous. Inflorescence a terminal spike on lateral branchlets; overtopping leaves, erect, dense, 12–25 mm long. Flowers 5–8, sessile; inflorescence bracts overtopping flowers, light green to glaucous, ovate–lanceolate at the base, 142 Australian Systematic Botany S. Venter 170º 175º 180º −35º −40º −45º Fig. 129. Known distribution of Dracophyllum pubescens, South Island, New Zealand. 10–17
1.2–1.7 mm, surfaces rugose; margins serrulate; flower bract overtopping flowers, ovate, 6.5–9.0
4.0–4.5 mm, surfaces glabrous with a tuft of scabrid hairs at apex on adaxial surface; margins ciliate. Sepals lanceolate to ovate–lanceolate, 4.8–6.0
1.5–2.0 mm, equalling corolla tube, with the top half pubescent on adaxial surface; margins ciliate. Corolla white to occasionally light pink; corolla tube narrowly campanulate, widened at mouth, 4.0–4.5
1.7–2.0 mm; corolla lobes reflexed, ovate–triangular to triangular, shorter than corolla tube, 1.3–1.7
1.4–1.5 mm; apex acute; adaxial surface papillate. Stamens inserted in middle of the corolla tube, filaments 0.19–0.2 mm long; anthers included, oblong, initially pink turning light yellow and 0.8–1.2 mm long. Ovary obovoid, 1.5–2.0
1.9–2.0 mm; apex round; nectary scales rectangular, 0.6–0.7
0.5–0.6 mm, apices irregularly toothed; style included, 1.5–2.0 mm long, glabrous; stigma capitate. Fruit light brown, 3–4
2.8–3.0 mm, broadly obovoid, apex round, glabrous. Seeds yellowish-brown, ovoid, 0.8–0.9 mm with testa slightly reticulate (Fig. 130, 131). Distribution and ecology Endemic to the North Island of New Zealand, centred on the volcanic plateau area with a few scattered localities in the Bay of Plenty area (Fig. 132). Dracophyllum recurvum grows on gentle to steep (3–60) mountain slopes or on plateaux at Taxonomic revision of Dracophyllum and Richea D A C Australian Systematic Botany E F 143 recurved leaf apices, flowers in short stout terminal racemes, very broad flower bracts, sepals equalling the corolla tube and the narrowly campanulate corolla tube. A most distinctive species (Cheeseman 1914) related and similar to D. marmoricola, but differs mainly in having smooth bark, recurved leaves with more (90–120) teeth per 10 mm on the lamina margin. The flower bracts are also longer than the flowers and the corolla tube is narrowly campanulate, not cylindrical. Dracophyllum recurvum is a remarkably distinct species. There is limited variation in growth habit owing to habitat preferences. The prostrate form is associated with subalpine areas, whereas plants forming cushions occur in drier and warmer areas in the Rangipo Desert. The shoulders of the lamina sheath are variable, from tapering to truncate. The inflorescences of plants growing in alpine or subalpine conditions are usually short (12–15 mm) compared with those growing in the Rangipo Desert (18–25 mm). G Selected specimens H NEW ZEALAND. North Island: western side of Mount Hikurangi, 7 Oct. 1964, Fryer s.n. (CHR); Mount Hikurangi, 30 Mar. 1932, Moore & Cranwell s.n. (AK); summit of Mount Kakaramea, Jan. 1905, Cheeseman. s.n. (AK); Raukumara Range, Maungawaru Plateau, Jan. 1953, Druce s.n. (CHR); Hauhangatahi, 9 Jan. 1933, Allan s.n. (CHR); Mount Tongariro, Nov. 1924, Sladden s.n. (CHR); Taupo, Mount Pihanga, 1 Apr. 1961, Parsons s.n. (CHR); Tongariro National Park, Turoa Ski field, 8 Feb. 1999, Venter 13757 (CHR); Mount Ruapehu, Feb. 1875, Berggren s.n. (O); ibid., 13 Jan. 1921, Matthews & Carse s.n. (AK); Kaweka range, The Tits, Dec. 1974, Druce s.n. (CHR); Kaweka Range, Makahu Stream, northern side of Makahu Spur, 3 Nov. 1966, Given 65331 (CHR); Ruahine Range, Makirikiri, Dec. 1976, Druce s.n. (CHR); Ruahine Range, 2 miles [~3.2 km] south-west of Takapari, Jan. 1966, Druce s.n. (CHR). B Dracophyllum rosmarinifolium (Forst.f.) R.Br. ex Roem. & Schult., Syst. Veg. 4: 385 (1819) Fig. 130. Dracophyllum recurvum. A. Flowering branch (1
). B. Laid-out corolla (5
). C. Lamina sheaths to show variation (5
). D. Leaf (1
). E. Ovary (10
). F. Flower (5
). G. Sepal adaxial surface (5
). H. Inflorescence-bract abaxial surface (5
). Drawn from Venter 13757. Del. S. Venter. elevations of 900–2000 m. The surrounding vegetation is montane to subalpine shrubland, fellfield, grassland, herbfield or tussockland. Soils are brown gritty sandy loam, brown loam or brown clay loam derived from andesite, basalt, pumice, rhyolite, scoria or volcanic sand. Phenology Flowering December–April. Etymology Refers to the strongly recurved leaves. Diagnostic features and notes Dracophyllum recurvum is characterised by the rugose to scabrid adaxial lamina surface, triquetrous and keeled Type: New Zealand. Dusky Bay, on summits of the highest mountains, 26 Mar. 1773. G. Forster s.n. (lecto: BM 577640!), designated by Hooker (1839). Dracophyllum rosmarinifolium (Forst.f.) R.Br., Prodr. Fl. Nov. Holl.: 556 (1810). in nota. nom. inval. Epacris rosmarinifolia Forst.f., Flor. Ins. Austr. Prodr.: 13 (1786). nom. inval. Dracophyllum uniflorum Hook.f., Handb. N. Zeal. Fl.: 182 (1864). Type: New Zealand. Wairau Mountains, Marlborough, W.T.L. Travers s.n. (lecto: K!; isolecto: CHR 45864!), designated by Oliver (1952). Left-hand specimen. Illustrations W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 4 (1928); W. R. B. Oliver, Trans. & Proc. Roy. Soc. N.Z. 80 (1): t. 2 (1952); L. B. Moore and J. B. Irwin, Oxford Book of N.Z. Plants: t. 143 (1978); A. Eagle, Trees & Shrubs of N.Z., 2nd Series: t. 143 (1982); J. Smith-Dodsworth, N.Z. Native Shrubs & Climbers: t. 54, pl. 22E, F (1991). 144 Australian Systematic Botany S. Venter A B C D E Fig. 131. Dracophyllum recurvum. A. ‘Desert’ habitat near Mount Ruapehu. B. Branch showing the recurved leaves. C. Alpine habitat at Mount Ruapehu. D. Prostrate growth habit in subalpine habitats showing the rough bark. E. Flowering branch. (D, Venter 13757). Photos: Jason Blair (A), S. Venter (B–D) and Alan Cressler (E). Multi-stemmed shrub 30–100 cm tall. Branches erect–spreading and much-branched. Bark on old branches grey to dark grey, finely to deeply fissured, young stems reddish-brown. Leaves erect–spreading, light to olive-green; lamina sheath (2.0–)3.0–8.5
2.5–4.0 mm; shoulders rounded to truncate and margins membranous, ciliate; lamina linear to linear–subulate, (8–)8.5–40.0(–55)
0.59–1.5 mm; adaxial surface glabrous, occasionally rugose, with a tuft of short scabrid hairs at the base; margins serrulate with 70–80 teeth per 10 mm; apex obtuse to acute and triquetrous. Inflorescence a terminal solitary erect flower; shorter than leaves; inflorescence bract shorter to equalling flower, narrowly ovate–lanceolate at the base, 5.0–9.5(–13.0)
1–2 mm, with adaxial surface scabrid at the base; margins serrulate. Flowers sessile. Sepals lanceolate to ovate–lanceolate, (4.5–)5.0–9.0(–12.0)
1.2–2.5 mm, equalling or longer than corolla tube, top half rarely shortly pubescent; margins ciliate. Corolla white, turning pale yellow with age, occasionally light pink; corolla tube cylindrical, (4–)5–7
1.5–2.0 mm; corolla lobes reflexed, triangular, shorter than corolla tube, 2.0–2.5
1.2–1.5(–2.0) mm; apex inflexed, subacute to acute; apical ridge present, adaxial surface papillate. Stamens inserted on corolla tube in the upper third, filaments 0.3–0.5 mm long; anthers included oblong, initially pink, turning light yellow and 0.7–1.0 mm long. Ovary obovoid, 1.7–2.0
1.0–1.3(–2.0) mm, apex round; nectary scales rectangular, (0.7–)1.0–1.5
0.4–0.7 mm; apices retuse to irregularly toothed; style included, 1.5–2.5 mm long, glabrous, not lengthening in fruit; stigma capitate. Fruit light brown, 3.7–4.0
Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 145 180º −35º −40º −45º Fig. 132. Known distribution of Dracophyllum recurvum, North Island, New Zealand. 3.8–4.0 mm, obovoid; apex round, glabrous. Seeds yellowishbrown, ovoid, 0.8–1.0 mm long with the testa slightly reticulate (Fig. 133, 134). Distribution and ecology New Zealand endemic. Widely distributed on the South Island, with a few scattered localities on the Tararua Mountains in the south of the North Island (Fig. 135). Dracophyllum rosmarinifolium occurs in mountain gullies, on mountain slopes ranging from 0 to 80, ridges, bluffs, plateaux and on valley floors from 152- to 2100-m elevation. The vegetation consists of montane woodland and shrubland to subalpine or alpine grassland, herbfield, fellfield or bogland. Soils are greyish- brown lithosols derived from alluvium, grey to greyish-brown gritty sandy loam derived from sandstone, conglomerate and granite or brown gritty clay loam to brown humus-rich clay loam derived from greywacke, marble, peridotite, serpentinite or schist. Plants occasionally grow in permanently moist areas (bogland) and in drainage lines, but normally receive most of their moisture from rain, melting snow and mist. In certain places, D. rosmarinifolium is very abundant and can form dominant stands, especially above the tree line. Phenology Flowering October–April. Plants in the southern part of its distribution sometimes flower as late as May. 146 Australian Systematic Botany D C S. Venter E F G B H Fig. 133. Dracophyllum rosmarinifolium. A. Flowering branch (1
). B. Laid-out corolla (5
). C. Lamina sheaths to show variation (5
). D. Leaf (1
). E. Ovary (10
). F. Flower (5
). G. Sepal (5
). H. Inflorescence-bract adaxial surface (5
). Drawn from Venter13747. Del. S. Venter. Etymology Refers to the leaves likened to those of the genus Rosmarinus L. Diagnostic features and notes Dracophyllum rosmarinifolium is characterised by the linear spreading leaves, solitary flowers that terminate short branchlets, sepals equalling or longer than the corolla tube, prominent apical ridge on the corolla lobes, inflexed corolla lobe apex and an obovoid ovary. Hooker (1839) was the first person to designate a lectotype for D. rosmarinifolium and he chose a G. Forster specimen collected in 1773. Richard (1832) used Forster’s manuscripts to publish a full description, to amplify Forster’s brief diagnosis. Cheeseman (1925) mentioned that his concept of D. rosmarinifolium differed from Forster’s (1786) Epacris rosmarinifolia. Oliver (1952) supplied a further amplified description of Forster’s D. rosmarinifolium [= Epacris rosmarinifolia]. This confusion arose because Forster’s species was not recollected for more than 100 years after his visit to Dusky Sound, and not recognised for another 40 years (Oliver 1952). Dracophyllum rosmarinifolium is similar to D. frondosum, but differs in having erect to erect–spreading, not decumbent, branches, inflorescence bracts equalling the flowers, not overtopping them, sepals longer than the corolla tube, not equalling them, and the corolla tube being shorter (5–7 mm compared to 7–10 mm). Further differences between the species are discussed under D. frondosum. During this study, butterflies were recorded pollinating the flowers of D. rosmarinifolium and D. acerosum, species with solitary, erect flowers having reasonably large spreading to recurved corolla lobes that provide effective settling platforms. The main distinguishing character, according to Simpson (1945) and Oliver (1928), is the shape of the lamina apex, which is obtuse in D. rosmarinifolium and subacute to acute in D. uniflorum. This character was seen to break down completely, while visiting the various populations in the wild (S. Venter, pers. obs.). Field observations showed that D. rosmarinifolium is an extremely polymorphic species and the present circumscription incorporates several ecological races scattered throughout the distribution range. Populations from the northern part of its distribution (Lake Sylvester, Venter 13775) tend to be low-growing, less erect, with shorter subacute leaves and with much smaller flowers. There is considerable variation in the size of the lamina (10–40
0.7–1.5 mm), the inflorescence bracts (5.0–9.5
1–2 mm), sepals (5–9
1.2–2.5 mm) and corolla tube (5–7
1.5–2.0 mm). The nectary scales can vary (1.7–2.0
0.4–0.7 mm) on a single plant. Branching habit and height depend largely on the habitat. Selected specimens NEW ZEALAND. North Island: Tararua Range, Mount Bannister, 5 June 1938, Healy 908 (CHR); Mount Hector, 29 Jan. 1907, Petrie s.n. (CHR); Mount Holdsworth. 26 Jan. 1990, Druce 482 (CHR); Mount Kaipororo, Apr. 1979, Druce s.n. (CHR). South Island: Mount Stevens, Feb. 1976, Druce s.n. (CHR); Lead Hills, Feb. 1935, Simpson s.n. (CHR); Burma Road, gully north-west of the office of an open-cast coal mine, 26 Jan. 1953, Mason & Moar 1762 (CHR); Westport, Mount Rochfort, near top, 25 Nov. 1998, Venter 13729 (CHR); ibid., 7 Jan. 1999, Venter 13747 (CHR); Gunner Downs, near Mount Barr, Nov. 1979, Druce s.n. (CHR); Matiri Range, Mount Misery, Mar. 1980, Druce s.n. (CHR); Motueka, Mount Arthur, above Arthur’s Hut, 2 Jan. 1999, Venter 13742 (CHR); Takaka, Cobb Reservoir, Lake Sylvester, 23 Mar. 1999, Venter 13775 (CHR); ibid. Venter 13776 (CHR); Motueka, Mount Arthur, Horseshoe Basin, 1 Mar. 2000, Venter 13796 (CHR); Nelson Lakes National Park, Travers Range, 2nd basin, 21 Mar. 1961, Simpson 3018 (CHR); Nelson, Mount Starveall, Gibbs s.n. (CHR); Mount Richmond, Feb. 1980, Druce s.n. (CHR); Marlborough Sounds, Moncrieff Scenic Reserve, Editor Hill, Feb. 1975, Kelly & Kelly s.n. (CHR); head of Hodder River, near Hodder Hut, 20 Apr. 1981, Wardle s.n. (CHR); Basin east of Mount Priestly, 11 June 1983, Wardle s.n. (CHR); Otira, Oct. 1908, Travers s.n. (Z); Arthur’s Pass, Mount Blimit, 21 Feb. 1943, Zotov s.n. (CHR); Lewis Pass 7 km Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 147 A B C D E F Fig. 134. Dracophyllum rosmarinifolium. A. Habitat in the Otira Valley near Arthurs Pass. B. Flowering branch showing the solitary flowers on short branches. C. Mature plant open form at Lake Mavora. D. Erect-stemmed form from Mount Rochfort. E. Mature plant of the compact form near Lake Te Kapo. F. Flowering branches, Lake Mavora. Photos: Steve Attwood (A), Phil Bendle (B, C, F) and S. Venter (D). from Maruia Springs to Christchurch, 13 Nov. 1998, Venter 13726 (CHR); Tarndale, Cat Creek, 2 Apr. 1946, Allan s.n. (CHR); south-west of Mount Baldy, Mar. 1984, Druce s.n. (CHR); Hanmer Springs, Jacks Pass, Jollies Scenic Reserve, 27 Feb. 2000, Venter 13794 (CHR); ibid. Venter 13795 (CHR); Crawford Range, Upper Hurunui River, 31 Jan. 1973, Macmillan 73/115 & Stemmer (CHR); Organ Range, ridge to The Organ, Island Hills Station, 12 Mar. 1991, Macmillan 91/38 & Woods (CHR; AK); Upper Awatere, Yeo Creek (–AB), 3 Apr. 1949, Allan s.n. (CHR)’ Karangarua River, between Cassel and Lame Duck Flats, 31 Mar. 1969, Wardle & Fryer s.n. (CHR); Ranganui, true left of Moeraki River, 18 Apr. 1978, Campbell s.n. (CHR); Franz Josef Glacier, Alex Knob, 19 Mar. 2000, Venter 13803 (CHR); Mount Cook National Park, Sealy Lakes track, near lakes, 13 Dec. 1966, Wilson 447 (CHR); Liebig Range, Mount Cook Station, northern side Jollie River Valley, 3 Mar. 1966, Macmillan 66/75 (CHR; HO); Mount Cook, Sebastopol, 26 Feb. 1958, Connor s.n. (CHR); Waimakariri Valley, Anti Crow River, 24 Mar. 1963, Fryer s.n. (CHR); Mount Somers, Woolshed Creek, Mar. 1987, Druce s.n. (CHR); Arundel, Mount Peel, 15 Jan. 1985, Mayrhofer & Molloy 4771 (GZU); Fiordland, Cats eye Bay, 5 Jan. 1978, Lee s.n. (CHR); Poison Bay, 7 Feb. 1974, Wardle & Mark s.n. (CHR); north of Homer Tunnel, Esperance Valley, Mar. 1974, Atkinson s.n. (CHR); Franklin Mountains, Nitz Creek, 31 Dec. 1968, Given 69033 (CHR); Cascade Valley, Woodhen Creek, 9 Mar. 1978, Wardle, Lee & Johnson s.n. (CHR); Cascade River, top of Martyr Hill, 11 Mar. 1978, Wardle, Lee & Johnson s.n. (CHR); Fiordland National Park, 148 Australian Systematic Botany S. Venter 170º 175º 180º −35º −40º −45º Fig. 135. Known distribution of Dracophyllum rosmarinifolium. Lake Harris, 21 Mar. 2000, Venter 13806 (CHR); Hunter Valley, Ferguson Creek, 4 Jan. 1961, Mason 8131 (CHR); Ahuriri River, slopes of Puke Makariri, 22 Jan. 1960, Connor s.n. (CHR); Hunter Hills, headwater of south Pareora River, Weaner Run, 4 Apr. 1973, Macmillan 73/401 & Woodhouse (CHR); Fiordland, Secretary Island, Central Hill, 3 Feb. 1967, Wardle s.n. (CHR); Fiordland, Secretary Island, southern peaks, 31 Jan. 1967, Wardle s.n. (CHR); Mount George, rocky basin immediately north of peak, 21 Mar. 1977, Garnock-Jones, Lee, Anderson & Given 10343 (CHR); Lake Manapouri, near entrance to Keplar Track, 7 Jan. 1997, Sykes 28/97 (CHR); Lake Hauroko, Hay Burn, Thomson s.n. (CHR); Thompson Range, North Von River, 3 m south of North Von Hut, 11 Feb. 1961, Connor s.n. (CHR); Queenstown, Ben Lamond, 2 Jan. 1936, Zotov s.n. (CHR); Umbrella Mountains, near Gem Lake, Mar. 1986, Druce s.n. (CHR); Palmerston, Sanatorium, 26 Apr. 1925, Millin s.n. (CHR); Dunedin, Mount Maungatua, 23. Mar 2000, Venter 13811 (CHR); Dunedin, Swampy Hill, Simpson & Thompson s.n. (CHR). Dracophyllum scoparium Hook.f., Fl. Antarct. 1: 46 (1844) Dracophyllum urvillianum var. scoparium Hook.f., Handb. N.Z. Fl.: 182 (1864). Type: New Zealand. Campbell Island, small bush near the sea, 7 Dec. 1840. J.D. Hooker 1611 (lecto: K000844567!; isolecto: BM 577654!; BM 577668!; BM 577669!; BM 577670!; WELT 54858!), designated by Oliver (1952). Dracophyllum paludosum Cockayne, Trans. & Proc. N.Z. Inst. 34: 318 (1902); Dracophyllum scoparium Hook.f. var. paludosum (Cockayne) Cheeseman, Man. N. Zeal. Fl.: 425 (1906). Dracophyllum arboreum Cockayne var. paludosum Cheeseman, Manual N. Zeal. Fl. 2nd edn: 707 (1925). Taxonomic revision of Dracophyllum and Richea Type: New Zealand. Chatham Island, Tableland, Feb. 1902. L. Cockayne s.n. (lecto: WELT 33105!), designated by Oliver (1952). Australian Systematic Botany D A 149 E Dracophyllum subantarcticum Cockayne, Veg, N.Z.: 265 (1921), nom. nud. Dracophyllum rosmarinifolium auct. non R.Br., Trans. & Proc. N. Z. Inst. 7: 338 (1875). Dracophyllum urvillianum auct. non A.Richard, Miss. Ile Camp. Bot.: 6 (1885). F Illustrations J. D. Hooker, Fl. Antarct. 1: t. 33 (1844); A. Eagle, Trees & Shrubs of N.Z., 2nd series: t. 145 (stamens inaccurately illustrated as hypogynous; 1982). Erect multi-stemmed shrub to small tree, 1–4 m tall. Branches: bark on old branches dark brown to blackishbrown, finely fissured, young stems reddish-brown. Leaves erect to spreading; lamina sheath (2–)3–5
1.5–4.0 mm, shoulders tapering to truncate and margins membranous and ciliate; lamina linear to linear–subulate, (24–)30–50(–80)
(0.3–)0.6–1.0(–1.5) mm, adaxial surface pubescent, slightly striated; margins ciliate to densely pubescent with 100–120 teeth per 10 mm; apex triquetrous. Inflorescence a terminal spike on lateral branches; shorter than leaves, erect, dense, 13–20 mm long, oblong; inflorescence bract overtopping flower, ovate–lanceolate at the base, 1.8–2.0
0.9–1.0 mm, adaxial surface glabrous, pubescent at apex; abaxial surface pubescent at the base; margins ciliate. Flowers 3–6, sessile. Flower bracts overtopping flowers, broadly ovate, 5.0–9.5
2.5–4.0 mm, adaxial surfaces pubescent; margins ciliate. Sepals oblong, (2.5–)4.0–5.0
(1.5–)2.0–3.5 mm, equalling or longer than corolla tube, striate, surfaces glabrous with the top half pubescent; margins ciliate; apices acute to acuminate. Corolla white; corolla tube cylindrical, 3.0–3.5
1.3–1.5 mm; corolla lobes reflexed, triangular, shorter than corolla tube, (1.7–)2.0–2.5
1.2–2.0 mm, apex inflexed and acute; adaxial surface papillate. Stamens inserted on corolla tube in the upper third, filaments 0.3–0.5 mm long; anthers included, rectangular, light yellow and 0.9–1.0 mm long. Ovary obovoid, 0.8–1.0
0.9–1.0 mm, apex round; nectary scales rectangular, 0.6–0.7
0.5–0.6 mm, apices subacute to obtuse; style included, 0.95–1.0 mm long, glabrous, not lengthening in fruit; stigma 5-lobed. Fruit light brown, 1.8–2.0
2.1–2.5 mm, obovoid; apex round, glabrous. Seeds light brown, ovoid, 0.7–0.8 mm long, testa slightly reticulate (Fig. 136, 137). C G B H Fig. 136. Dracophyllum scoparium. A. Flowering branch (1
). B. Laidout corolla (5
). C. Lamina sheaths to show variation (2
). D. Leaf (1
). E. Ovary (10
). F. Flower (5
). G. Sepal abaxial surface (5
). H. Inflorescence-bract abaxial surface (5
). Drawn from Brockie s.n. (O). Del. S. Venter. Phenology Flowering November–February. Etymology In the form of a broom, referring to the broom-shaped habit of the juvenile leaves Distribution and ecology New Zealand endemic restricted to the subantarctic Campbell, Chatham and Pitt islands (Fig. 138). Dracophyllum scoparium grows on flat boggy areas or on gentle (5–30) hill slopes reaching from sea level up to 260-m elevation. These areas are covered with shrubland that grows up to 5 m tall in sheltered areas and in the open areas, grassland and bogland. The soil is moist for prolonged periods, owing to the high rainfall and frequent mists. Diagnostic features and notes Oliver (1952) was the first to designate a specific specimen to the name and he chose the specimen from the original Cockayne Herbarium (Number 3567). This specimen later ended up in the Petrie Herbarium, and is now incorporated in the WELT collection as WELT 33105. Dracophyllum scoparium is similar to D. filifolium, but differs in having smaller leaves (30–50
0.6–1.0 mm 150 Australian Systematic Botany S. Venter A B C D Fig. 137. Dracophyllum scoparium. A. Habitat on Campbell Island, showing D. scoparium as the dominant woody species. B. Flowering branch showing the erect–spreading leaves. C. The characteristic cuticular wax plates. D. Mature plant in flower. Photos: Janet Wilmshurst (A), Colin Meurk (B), S. Venter (C) and P. Garnock-Jones (D). compared with 60–130
1.0–1.5 mm), lamina pubescent on the adaxial surfaces, not glabrous, with the lamina margin densely pubescent (glabrous in D. filifolium). The inflorescence bract is subulate with pubescent abaxial surfaces and the adaxial surface of the flower bract is pubescent, not glabrous. The sepals are striate and longer than the corolla tube (not equalling and smooth) with the top half being pubescent, not glabrous. The corolla tube is narrower (1.3–1.5 mm compared with 1.8–2.5 mm) with longer (2.0–2.5 mm compared with 1.5–2.0 mm) triangular corolla lobes having papillate, not glabrous, adaxial surfaces, smaller nectary scales and an obovate, not subglobose, ovary. Plants of D. scoparium have a dense covering of long whitish hairs at the base of the lamina that can sometimes Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 151 Fig. 138. Known distribution of Dracophyllum scoparium. The main map is of the Chatham Islands and the insert that of Campbell Island, 1700 km further south. Dracophyllum septentrionale (W.R.B.Oliv.) S.Venter comb. et. stat. nov. be short and scabrid in some individuals. The shape of the leaf sheath shoulders is polymorph and varies from tapering to truncate on the same branch. Some variation exists in lamina size (30–50
0.6–1.0 mm). The flower bract is variable in size even on the same plant (5.0–9.5
2.5–4.0 mm). Dracophyllum longifolium var. septentrionale W.R.B.Oliv., Trans. & Proc. Roy. Soc. N.Z. 80 (1): 11 (1952). Selected specimens Type: New Zealand. Ruahine Mountains, Mount Maharahara, Apr. 1946. R.M. Greenwood s.n. (holo: CHR 65032!). NEW ZEALAND. Chatham Island, Taupeka, 21 Feb. 1996, De Lange CH14 & Crocroft (AK); Te Puke Hill between Waitangi West and Maunganui, 13 Nov. 1991, Dugdale & Macfarlane s.n. (CHR); Te Puke Hill, 6 June 1996, Dugdale & Macfarlane s.n. (CHR); Southern Plateau, Tuku headwaters, 4 Mar. 1985, Wardle s.n. (CHR); Lake Rakenui, The Clears, 24 Oct. 1987, Taylor s.n. (AK). Campbell Island: Tucker Cove Valley, 7 Feb. 1947, Brockie s.n. (CHR; O); Camp Cove, 6 Jan. 1961, Godley s.n. (CHR); South Col ridges, 20 Dec. 1944, Oliver s.n. (CHR); valley at head of South East Harbour, 15 Nov. 1945, Sorenson s.n. (CHR); Beeman Cove, Lookout Bay, Feb. 1976, Given 9167 (CHR; HO); Smoothwater Bay, 25 Jan. 1976, Given 9276 (CHR); Bull Rock, 6 Feb. 1976, Given 9423 (CHR); Beeman Point, 7 Dec. 1995, Meurk s.n. (CHR). Illustration W. R. B. Oliver, Trans. & Proc. Roy. Soc. N.Z. 80 (1): t. 6 (1952). A multi-stemmed shrub or small tree, 1–2 m tall. Branches: bark on old branches dark grey to greyish-brown, smooth or finely fissured, young stems reddish-brown. Leaves adult and juvenile. Juvenile leaves spirally arranged along branches, spreading; lamina sheath yellowish-green, 11–16
6.0–7.6 mm, shoulders truncate and margin ciliate in upper half; lamina linear to linear–triangular, surfaces glabrous, 152 Australian Systematic Botany 110–200
4–6 mm, margins serrulate with 50–60 teeth per 10 mm; adult leaves erect–spreading; lamina sheath 7–12
2.8–5.0 mm, striate, rounded to truncate and margin membranous with the top half ciliate; lamina linear to linear–triangular, (46–)80–130
1.0–2.5 mm; adaxial surface rugose; margins serrulate with 50–60 teeth per 10 mm. Inflorescence a raceme near the apices of branches; shorter than leaves, erect, dense, 12–24 mm long, oblong; inflorescence bracts overtopping flowers, ovate–lanceolate, 28–37
0.6–0.7 mm, adaxial surfaces scabrid; margins serrulate, apices acuminate. Flowers 3–11, pedicellate; flower bracts caducous, overtopping flowers, coriaceous, broadly ovate, 5.0–7.5
2–3 mm, adaxial surfaces sericeous; margins ciliate; apices acute; pedicels straight, 0.6–1.5 mm long, glabrous. Sepals ovate–lanceolate, 3.5–5.0
1.3–2.0 mm, equalling corolla tube, adaxial surface pubescent in the top half; margins ciliate. Corolla white; corolla tube cylindrical, 3.5–4.0
1.5–2.0 mm; corolla lobes reflexed, ovate– triangular, shorter than corolla tube, 1.5–1.7
1.0–1.2 mm, apex subacute; surfaces glabrous. Stamens inserted onto corolla tube near the top, filaments 0.6–1.0 mm long; anthers included, oblong, light yellow and 0.9–1.0 mm long. Ovary obovoid, 2.9–3.0
1.8–2.0 mm, apex round; nectary scales rectangular, 1.5–1.6
0.7–0.8 mm, apices retuse; style included, 1.3–1.5 mm long, glabrous; stigma 5-lobed. Fruit pedicellate, light brown, 2–3
2.0–3.5 mm, obovoid, apex round, glabrous. Seeds cream-coloured, ovoid, 0.7–1.3 mm long, testa slightly reticulate (Fig. 139, 140). Distribution and ecology New Zealand endemic, restricted to the North Island in the Taihape area, on the Ruahine Mountains and on the Flaggstaff Mountains (Fig. 141). Dracophyllum septentrionale appears to be restricted to gentle (15–30) mountain slopes at elevations of 1000–1500 m. These slopes are covered in dense montane shrubland. Dracophyllum septentrionale has so far only been recorded on dark brown clay loam that is derived from shale. Phenology Flowering November–February. Etymology Describes the northern distribution. Diagnostic features and notes Dracophyllum septentrionale is characterised by the long and wide juvenile (110–200
4–6 mm) and adult leaves (80–125
1.5–2.5 mm), rugose adaxial surface of the adult lamina, caducous flower bracts overtopping the flower with sericeous adaxial surfaces, short pedicels, sepals equalling the corolla tube and being pubescent in the top half of the adaxial surface, stamens inserted onto the corolla tube near the top and the cream-coloured ovoid seeds. It is similar to D. oliveri in the racemose inflorescence, serrulate margin of the inflorescence bract and the retuse apices of the nectar scales but differs in various leaf and flower characters. Oliver (1952) compared the raceme and leaves of D. longifolium var. septentrionale with those of S. Venter D. filifolium but described it as a variety of D. longifolium on the basis of the measurements of the leaves, stating that the leaves are much wider than those of D. filifolium. It is a very uniform species. A basal patch of very short scabrid hairs is sometimes present at the base of the lamina in some populations. Selected specimens NEW ZEALAND. North Island: Ruahine Mountains, Mount Maharahara, Apr. 1946, Greenwood s.n. (CHR); ibid., 7 Feb. 1999, Venter 13756 (CHR); south-west of Takapari, Nov. 1979, Druce s.n. (CHR); Wairarapa, Taipos 1 km west of Kupekore, May 1965, Druce s.n. (CHR). Dracophyllum sinclairii Cheeseman, Man. N. Zeal. Fl.: 421 (1906) Type: New Zealand. Manukau Bay (Green Bay, Manukau Bay), W. Colenso s.n. (lecto: K000844549!), designated by Oliver (1952). Dracophyllum squarrosum Hook.f., Fl. Antarct. 1: 48 (1844). nom. illig., non R.Br. Type: New Zealand. Manukau Bay (Green Bay, Manukau Bay), W. Colenso s.n. (lecto: K!), designated by Oliver (1952). Dracophyllum adamsii Petrie, Trans. & Proc. N.Z. Inst. 55: 435 (1924). Type: New Zealand. Waiapu County, roadside near mouth of Awatere River, Jan. 1897. J. Adams & D. Petrie s.n. (lecto: WELT 55082!; isolecto: CHR 332735!, WELT 55081!, 55084!, WELTU 4240!.), designated by Oliver (1952). Dracophyllum viride W.R.B.Oliv., Trans. & Proc. N.Z. Inst. 59: 699 (1928). Type: New Zealand. Mangonui County, Spirit’s Bay, Peria, 28 Nov. 1916. W.R.B. Oliver s.n. (lecto: WELT 33297!; isolecto: AK 105638!, WELT 13558!.), designated by Oliver (1952). Illustrations W. R. B. Oliver, Trans. & Proc. N.Z. Inst. tt. 8, 10, 12 (1928); W. R. B. Oliver, Trans. & Proc. Roy. Soc. N.Z. 80 (1): t. 10 (1952); A. Eagle, Trees & Shrubs of N.Z., 2nd series: tt. 134, 135 (stamens inaccurately illustrated as hypogynous) and 140 (anthers inaccurately illustrated as exserted; 1982); J. T. Salmon, Native Trees N.Z.: 274 (1989), Native N.Z. Flowering Plants: 61, tt. 243–245 (1991). Single-stemmed small tree (1–)4–7(–7.6) m tall. Branches: bark on old branches dark grey to blackish-brown, finely to deeply fissured, young stems reddish-brown. Leaves juvenile and adult. Juvenile leaves spirally arranged along branches, spreading to recurved; lamina sheath, 9–30
5.0–12.7 mm, shoulders tapering to truncate and margin entire, occasionally ciliate in upper half; lamina subcoriaceous to coriaceous, linear–triangular, 85–221
2.5–9.0 mm, surfaces glabrous with a patch of scabrid hairs at the base of the adaxial surface; margins serrulate with 50–70 teeth per 10 mm. Adult leaves spreading to recurved, glaucous to light green (occasionally light brown); lamina sheath (3.5–)5.7–7.3
3.3–6.6 mm, rounded to truncate and margin membranous with the top Taxonomic revision of Dracophyllum and Richea C Australian Systematic Botany A 153 D E F B Fig. 139. Dracophyllum septentrionale. A. Flowering branch (1
). B. Laid-out corolla (5
). C. Juvenile and adult leaves (1
). D. Ovary (10
). E. Flower (3
). F. Sepal adaxial surface (8
). Drawn from Venter 13756. Del. S. Venter. half ciliate; lamina linear–triangular, 37–95
(1–)2–3(–5) mm, surfaces glabrous with a tuft of scabrid hairs at the base on adaxial surface; slightly striated; margins serrulate with 60–80 teeth per 10 mm. Inflorescence a terminal spike; shorter than leaves, erect, drooping later, dense, 14.5–34.7 mm long, linear–oblong; inflorescence bract overtopping flowers, ovate to broadly ovate, 7–8
0.7–1.0 mm, adaxial surface glabrous with a patch of scabrid hairs at the base; margins serrulate. Flowers 4–9, sessile. Flower bracts overtopping flowers, ovate, 6–20
3–4 mm, adaxial surfaces sericeous; abaxial surfaces glabrous to scabrid; margins minutely serrulate. Sepals ovate–lanceolate, (2.5–)5.0–6.0
1.2–2.0 mm, longer than corolla tube; adaxial surfaces with the top half pubescent; margins ciliate. Corolla white; corolla tube narrowly campanulate, widened at mouth, 4.0–4.5
2.5–3.0 mm; corolla lobes spreading horizontally to reflexed, triangular, shorter than corolla tube, 2.3–2.5
1.7–2.2 mm; apex inflexed, subacute; adaxial surface papillate. Stamens inserted on corolla tube in the upper third, filaments 0.5–1.0 mm long; anthers included, rectangular, light yellow and 0.8–1.0 mm long. Ovary obovoid, 1.4–1.5 mm long and wide, apex round; oblong, 154 Australian Systematic Botany S. Venter A B C D E Fig. 140. Dracophyllum septentrionale. A. Habitat on the Ruahine Mountains. B. Juvenile plant showing the large juvenile leaves. C. Mature plant (Venter13756). D. Mature leaves. E. Mature plant from the type locality, Mount Maharahara (Venter13756), Ruahine Mountains. Photos: S. Venter (A–E). 1.3–1.4
0.6–0.7 mm; apices mostly bidentate, sometimes irregularly toothed; style included, 1.3–1.5 mm long, glabrous; stigma 5-lobed. Fruit light brown, 1.5–3.5
1.5–2.5 mm, obovoid; apex truncate, glabrous. Seeds yellowish-brown, ovoid, 1.0–1.3 mm long, testa slightly reticulate (Fig. 142, 143). Distribution and ecology New Zealand endemic restricted to the North Island from Gisborne north and on Great Barrier Island (Fig. 144). Dracophyllum sinclairii is common on coastal cliffs, along ridges, mountain slopes, along drainage lines, gently sloped (5–30) scree areas and on mountain summits from 25- to 1432-m elevation. The vegetation consists of lowland to montane forest, woodland, shrub or shrub–tussockland. Dracophyllum sinclairii is associated with Leptospermum scoparium shrubland and woodland on the dry ridges along the coast. Soils are yellowish-brown clay loam to light brown clay derived from mudstone, dacite or andesite, or grey–brown loam derived from sandstone. Phenology Flowering January–November. Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 155 180º −35º −40º −45º Fig. 141. Known distribution of Dracophyllum septentrionale, North Island, New Zealand. Etymology Named after Dr Andrew Sinclair (1794–1861), naval surgeon and plant collector. Diagnostic features and notes Dracophyllum sinclairii is characterised by the large juvenile leaves, slightly striated adult leaves, the inflorescences on the lateral branches grouped together below the leaves of the main branch, flower bracts longer than the flower and covered in dense long hairs (sericeous) on the adaxial surfaces (in some populations only at the top half), sepals longer than the corolla tube and sericeous in the top half on the adaxial surface and the narrow campanulate corolla tube. Cheeseman (1906) failed to mention a specimen when he described the species, but Hooker (1844) mentioned the Colenso specimen under D. squarrosum, which Oliver (1952) designated to D. sinclairii. Dracophyllum sinclairii is similar to D. lessonianum but differs in having a much wider (2.5–9.0 mm compared with 1.6–1.8 mm) juvenile lamina, with a patch of scabrid hairs at the base on the adaxial surface. The adult lamina is also wider (2–3 mm compared with 0.5–1.2 mm), with an acute apex not prominently triquetrous. The inflorescence bract has a serrulate, not entire, margin. The corolla tube in D. sinclairii is narrowly campanulate, not cylindrical, shorter (4.0–4.5 mm compared with 5–6 mm) and wider (2.5–3.0 mm compared with 2.0–2.5 mm). 156 Australian Systematic Botany S. Venter A F H G I E D C B Fig. 142. Dracophyllum sinclairii. A. Flowering branch (1
). B. Flower-bract adaxial surface (2
). C. Sepal adaxial surface (5
). D. Laid-out corolla (5
). E. Lamina sheaths to show variation (2
). F. Juvenile leaf (1
). G. Adult leaf (1
). H. Ovary (10
). I. Flower (5
). Drawn from Venter 13781. Del. S. Venter. A highly polymorphic species (Fig. 145), having the shoulders of the leaf sheath of both juvenile and adult leaves tapering, rounded or truncate. The size of the leaf sheaths is, likewise, variable in the juvenile (9–30
5.0–12.7 mm) and adult (5.7–7.3
3.3–6.6 mm) leaves. The lamina size varies (juvenile: 85–221
2.5–9.0 mm; adult: 37–95
2–3 mm), with a patch of scabrid hairs being sometimes present at the base of the lamina and some populations lacking this character entirely. The length of the inflorescence varies from population to population (14.5–34.7 mm) and flower-bract length is variable even on the same plant (6.0–20.0 mm). The three groups formerly known as D. sinclairii, D. adamsii and D. viride grade into each other to such an Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A B C E 157 D F Fig. 143. Dracophyllum sinclairii. A. Habitat along the Kaimai Range. B. Stem of mature tree. C. Mature tree in Otari Wilton Bush Reserve. D. Branch with juvenile leaves (Venter 13780). E. Flowering branch (Venter13769). F. Branch showing adult and juvenile leaves. Photos: Dennis Kuhn (A), Phil Bendle (C), S. Venter (B, D–F). extent that it would be pointless to attempt to describe them as varieties of D. sinclairii (Fig. 144). Plants with glabrous, long and narrow inflorescence bracts tend to be more common in the area north of Auckland, but are still present in some populations in the east (Raukumara Range). Specimens with the inflorescence bracts having the top half pubescent are common in the area around Auckland and the Coromandel Peninsula. Plants with flower bracts having the top half of the adaxial surface covered in scabrid hairs or being tomentose are common in the area around Auckland and further north, 158 Australian Systematic Botany 172ºE S. Venter 173ºE 174ºE 175ºE 176ºE 177ºE 178ºE 179ºE 35ºS Druce 711 36ºS 37ºS Venter 13766 Overlap area 38ºS Nicholls s.n. 39ºS Overlap area 40ºS Fig. 144. Known distribution and variation in Dracophyllum sinclairii, North Island, New Zealand. The three most different forms of inflorescence bracts and nectary scales are illustrated. Druce 711, North Auckland, Unuwhao. Venter 13766, Coromandel Peninsula, Black Jack Road. Nicholls s.n., northern Kaimai Range, Mount Ngatamahinerua. A. Inflorescence-bract adaxial surface (1
). B. Flower-bract adaxial surface (2
). C. Nectary scale (10
). whereas plants with pubescent flower bracts are more common in the eastern part of the distribution. Specimens with rounded apices to the nectary scales occur mostly in the Coromandel populations and the apices become more toothed in specimens further to the north and east. In describing Dracophyllum viride, Oliver (1952) mentioned that he regarded the specimens from Spirits Bay as D. viride, whereas Petrie placed them in D. adamsii. During a survey of the Spirits Bay area, I found the plants to be extremely variable with regard to the size of the juvenile and adult leaves, shape and degree of pubescence of the inflorescence and flower bracts and the apex shape of the nectary scales. Oliver (1952) stated that D. sinclairii is allied to D. longifolium in the deciduous bracts and the narrow flat leaves. All herbarium specimens and the plants in the wild that I studied, showed persistent bracts. Oliver (1952) used this character to place D. adamsii in the D. longifolium group (D. longifolium and D. adamsii). Oliver (1952) mentioned that D. viride is related to D. sinclairii and D. adamsii, but differed in the larger, broader leaves, short flowering branches and much longer flowers with long acuminate sepals. All these variations fall within my concept of D. sinclairii. Selected specimens NEW ZEALAND. North Island: North Auckland, Unuwhao, Jan. 1990, Druce 711 (CHR); Spirits Bay, Pandora Bush, 18 Nov. 1976, Michie & Bartlett s.n. (AK); Northland, Te Paki, Dec. 1966, Kelly s.n. (CHR); North Tangihua Forest, northern end of ridge before trig, 29 Aug. 1991, Cameron 6569 (AK); Northland, Ahipara, Bartlett s.n. (CHR); Mangonui County, Peria, 14 Nov. 1913, Carse s.n. (CHR); Paranui, Taylor’s Road, 26 Jan. 1984, McCrae s.n. (AK, MU); Kerikeri, Puketi Forest, near Trig 1171, 15 Feb. 1999, Venter 13769 (CHR); Northland, Lake Taharoa, 15 Jan. 1978, Bartlett s.n. (CHR); Little Barrier Island, Shakespear s.n. (CHR); Auckland, Birkenhead, 2 July 1999, Venter 13780 (CHR); Auckland, Manukau Harbour, Green Bay, 30 June 1999, Venter 13781 (CHR); Coatesville, 6 Apr. 1982, Clunie 28 (CHR); Little Barrier Island, Shakespear s.n. (CHR); Little Barrier Island, Track 16, 25 Jan. 1980, Beever 80245 (CHR); Great Barrier Island, Te Ahumata, track from Whangaparapara side, near summit, 14 Mar. 1991, Garnock- Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A B C D 159 Fig. 145. Dracophyllum sinclairii variation. A. Dracophyllum adamsii (CHR 322398). B. Dracophyllum viride (CHR 332191A). C. Dracophyllum adamsii (WELT63766). D. Dracophyllum sinclairii, holotype (K000844548). Jones & Clarkson (CHR; HO); Coromandel Peninsula, track from Stony Bay to Mochau trigonometrical beacon 14 Nov. 1981, Cameron 759 (AK, CHR); Coromandel, Black Jack Road, 28 Sep. 1992, De Lange 1714 (CHR); ibid. 13 Feb. 1999, Venter 13766 (CHR); Kennedy Bay, 25 Oct. 1872, Kirk s.n. (WELT 55086); Coromandel Peninsula, Opito Bay, 24 Feb. 1981, Bartlett s.n. (CHR); Great Mercury Island, valley behind Peachgrove Bay, 2 Sep. 1962, Atkinson s.n. (CHR); Mercury Bay, 5 Nov. 1769, Banks & Solander s.n. (AK, BM); northern Kaimai Range, Mount Ngatamahinerua, Aug. 1961, Nicholls s.n. (CHR); eastern Bay of Plenty, Waihau Bay, 20 Oct. 1984, Courtney s.n. (CHR); eastern Bay of Plenty, Whitianga Bay, Okawhiti Stream, Dec. 1983, Courtney s.n. (CHR); Raukumara Range, Mount Honokawa, 30 Dec. 1962, Druce s.n. (CHR); East Cape, Mount Hikurangi, Dec. 1949, Druce s.n. (CHR); Pukeamaru, 25 Nov. 1949, Oliver s.n. (WELT 13559); Waitapu Stream, 22 Jan. 1985, De Lange s.n. (AK); Te Wharu Bush, 14 Jan. 1967, Devlin s.n. (AK); Raukumara Range, summit of Arowhana, 16 Oct. 1964, Fryer s.n. (CHR); Ruahine Range, Mount Hikurangi, Jan. 1897, Petrie s.n. (Z). 160 Australian Systematic Botany S. Venter Dracophyllum subulatum Hook.f., Fl. Antarct. 1: 50 (1844) A Type: New Zealand. Near the Rangitaiki River [near nowadays town of Galatea], Jan. 1842. J.C. Bidwill s.n. (lecto: K!), designated by Oliver (1952). H Dracophyllum angustifolium Colenso, Trans. Proc. N.Z. Inst. 28: 603 (1896). Type: New Zealand. Ruahine Mountain range, 1895. H. Hill s.n. (n.v.). F G Illustrations T. F. Cheeseman, Ill. N. Zeal. Fl.: t. 132 (1914); A. Eagle, Trees & Shrubs of N.Z., 1st series: t. 168 (1975). The anthers are depicted as being exserted instead of inserted; J. SmithDodsworth, N.Z. Native Shrubs & Climbers: t. 57, pl. 23E, F (1991). A multi-stemmed shrub 30–200 cm tall. Branches: bark on old branches grey, smooth, young stems reddish- to purplishbrown. Leaves juvenile and adult. Juvenile leaves spirally arranged along branches, spreading; lamina sheath 4.5–6.5
2.5–3.0 mm, shoulders truncate to auricled and margin membranous with the upper half ciliate; lamina linear to rarely linear–triangular, 18–45
1–2 mm, adaxial surface with a patch of scabrid hairs at the base; margins serrulate with 70–80 teeth per 10 mm. Adult leaves erect–spreading, olive- to dark green; lamina sheath 2.5–6.5
2–4 mm, subcoriaceous, shoulders rounded to auricled and margins membranous, ciliate; lamina linear to linear–subulate, 10–48
0.5–1.2 mm, surfaces glabrous with a tuft of scabrid hairs at the base on adaxial surface; margins serrulate with 90–120 teeth per 10 mm; apex triquetrous. Inflorescence a terminal spike on lateral branches; shorter than leaves, erect, dense, 5.3–12.3 mm long, linear–oblong; inflorescence bract overtopping flowers, ovate–lanceolate at the base, 4–6
0.5–0.6 mm; margins serrulate. Flowers 2–4 (–6), sessile; flower bracts equalling or longer than flowers, leaf like, broadly ovate, 3.5–4.5(–7.8)
1.7–2.0 mm, with a tuft of scabrid hairs at the apex; margins with a prominent broad and white margin, serrulate. Sepals lanceolate to ovate–lanceolate, 2.6–3.2
0.6–1.2 mm, longer than corolla tube; adaxial surface pubescent or only the top half pubescent; margins ciliate. Corolla white to light pink; corolla tube cylindrical, 1.8–2.0
1.0–1.2 mm; corolla lobes spreading horizontally to reflexed, triangular, shorter than corolla tube, 1.0–1.5
0.8–1.0 mm; apices acute; adaxial surface papillate. Stamens inserted in corolla tube in the upper third, filaments 0.1–0.2 mm long; anthers included rectangular, light yellow and 0.7–0.8 mm long. Ovary obovoid, 0.8–1.0 mm long and wide, apex truncate; nectary scales rectangular, 0.4–0.5
0.3–0.4 mm, apices retuse; style included, 0.5–1.0 mm long, glabrous; stigma clavate. Fruit 2.9–3.0
1.7– 1.8 mm, oblong; apex truncate, glabrous. Seeds yellowish-brown, filiform, 1.0–1.2 mm long, testa prominently reticulate (Fig. 146, 147). I E J D C B Fig. 146. Dracophyllum subulatum. A. Flowering branch (1
). B. Flowerbract adaxial surface (5
). C. Sepal (5
). D. Laid-open corolla (5
). E. Lamina sheaths to show variation (5
). F. Juvenile leaf (1
). G. Adult leaf (5
). H. Ovary (10
). I. Flower (5
). J. Inflorescence bract (5
). Drawn from Venter 13761. Del. S. Venter. Distribution and ecology New Zealand endemic, restricted to the Volcanic Plateau area on the North Island (Fig. 148). Dracophyllum subulatum grows at elevations of 100–1220 m on the Volcanic Plateau and on the associated volcanoes and mountains surrounding the Plateau. It occurs on flat or gently sloped (5–15) areas that are covered in montane shrubland, shrub–tussockland, grassland, fernland or bogs. The soil is mostly brown to grey sandy loam derived from greywacke, andesite or pumice. Dracophyllum subulatum and D. recurvum are associated with areas where there is a presence of tephra (B. Clarkson, pers. comm.). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 161 A B D C E Fig. 147. Dracophyllum subulatum. A. Frost flats near Mount Ngaurahue. B. Habitat near Mount Ruapehu. C. Flowering branches showing the short inflorescence and small flowers. D. Illustration from Cheeseman’s ‘Illustrations of the New Zealand Flora’. E. Flowering branch with pollinator. Photos: Juergen (A), R. Chappell (B), J. Braggins (C, E). Phenology Diagnostic features and notes Flowering November–March. Dracophyllum subulatum is characterised by the slender branches, juvenile leaves, small (5.3–12.3 mm long) fewflowered (2–6) inflorescences, flower bracts with a broad white margin, small corolla tubes (1.8–2.0
1.0–1.2 mm) and seeds with a prominently reticulate testa. Etymology Describing the awl-shaped leaves. 162 Australian Systematic Botany S. Venter 170º 175º 180º −35º −40º −45º Fig. 148. Known distribution of Dracophyllum subulatum. Dracophyllum subulatum is similar to D. palustre, especially in the flower bracts having prominent white margins. It differs in having juvenile leaves, inflorescence being a spike not a solitary flower, inflorescence bracts longer than the flower, not equalling, and being much narrower (0.5–0.6 mm compared with 1.5–2.0 mm), sepals longer and filiform, not ovoid, and the seed with a prominently reticulate testa not slightly reticulate. Lamina size is variable in the juvenile leaves (18–45
1–2 mm) and the adult leaves (10–48
0.5–1.2 mm). Inflorescence length varies from 5.3 to 12.3 mm. All this variation can be present in a single population. Selected specimens NEW ZEALAND. North Island: south of Lake Waipapa, 18 Nov. 1978, Gardner 2135 (CHR, L); Hauhungaroa Range, Maungatukutuku Stream, Apr. 1984, Druce APD617 (CHR); Maraeroa, Jan. 1947, Bannister s.n. (CHR); Wairakei, Craters of the Moon, July 1976, Given 9498 (CHR); Moerangi, 7 Feb. 1979, Gardner 2298 (CHR); Hauhangaroa Range, Kuratau Clearing, 24 Jan. 1951, Druce s.n. (CHR); Rotoroa, Whakarewarewa geyser enclosure, 20 Mar. 1969, Macmillan 69/183 (CHR); Mount Tarawera, 16 Apr. 1989, Clarkson s.n. (CHR); Mihi, 24 Jan. 1963, Mason 9888 (CHR); Waiotapu, 25 Apr. 1939, Healy s.n. (CHR); Maungakakaramea [Rainbow Mountain], Jan. 1905, Cheeseman s.n. (AK); Lake Taupo, Feb. 1875, Berggren s.n. (O); Hilltop above Waiora Valley bores, 27 Nov. 1979, Given 11930 (CHR); Broadlands, 2 Taxonomic revision of Dracophyllum and Richea Mar. 1978, Given 11105 (CHR); 13 km south of Taupo Township, 15 Mar. 1962, Melville & Melville 6707B (AK); Kaingaroa Plain, Rangitaiki, Oct. 1978, Druce s.n. (CHR); Huiarau Range, near Maungataniwha Scientific Reserve, Oct. 1977, Druce s.n. (CHR); Urewera National Park, Mangatoatoa Clearing, 7 Nov. 1984, Shaw & Beadel s.n. (CHR); National Park Junction, 16 Apr. 1966, Lamoureux 3653 (CHR); Turangi, Rangipo Desert, 9 Feb. 1999, Venter 13761 (CHR); Tongariro River, Pourini Scenic Reserve, 24 Nov. 1983, Gardner 3981 (AK); Kakaramea, Dec. 1912, Aston s.n. (CHR); Mount Ruapehu, west of Bruce Road, 27 Jan. 1983, Powell 2016 (CHR); Tongariro National Park, Mount Ngaurahue, 8 Feb. 1999, Venter 13758 (CHR); northeastern Kaimanawa Range, Poronui, 15 Feb. 1973, Gardner 529 (CHR); Potonui Station, 7 July 1949, Poole s.n. (CHR); Rangitikei, upper edge of Tikitiki Bush, 7 Jan. 1950, Hamlin s.n. (CHR); Puketitiri, Ball’s Clearing, Dec. 1976, Druce s.n. (CHR); Kaweka Range, eastern foot of Kuripapango Hill, The Lakes, Dec. 1974, Druce s.n. (CHR); Maungaharuru Range, 1 mile [1.6 km] east of Kopua, Dec. 1970, Druce s.n. (CHR). Dracophyllum trimorphum W.R.B. Oliv., Trans. & Proc. Roy. Soc. N.Z. 80 (1): 8 (1952) Type: New Zealand. West Whanganui Inlet, near shore in shrub, 28 Dec. 1949. W.R.B. Oliver s.n. (holo: WELT 55515a!; iso: WELT 55515b, WELT 55515c!, WELT 55516a!, WELT 55516b!, WELT 55516c!, WELT 55516d!, WELT 55516e!, WELT 55516f!, WELT 55516 g!). Australian Systematic Botany 0.3–0.5 mm long; anthers included, oblong, light yellow and 1.2–1.3 mm long. Ovary obovoid, 1.3–2.0
1.3–1.5 mm, apex pubescent, truncate; nectary scales rectangular, 1.0–1.5
0.5–0.7 mm, apices obtuse to retuse; style included, 1.2–2.0 mm long, glabrous; stigma 5-lobed. Fruit dark brown, 1.5–3.0
1.5–2.0 mm, obovoid; apex truncate, shortly pubescent. Seeds cream-coloured, ovoid, 1.0–1.3 mm long, testa slightly reticulate (Fig. 149, 150). Distribution and ecology New Zealand endemic restricted to the extreme northern part of the north-western Nelson area, South Island (Fig. 151). Dracophyllum trimorphum has not been recorded further than 1 km from the sea. It occurs from sea level up to 150-m elevation on steep (70–90) sea cliffs, gentle (5–30) hill slopes and ridges covered in lowland shrubland or heathland. Soils are light brown sandy-loam lithosol or greyish-brown clay–loam lithosol derived from pebbly conglomerate and quartzofeldspathic sandstone of the Kapuni Group in the Farewell Formation. Plants normally grow fully exposed, with some individuals growing in light shade on the margin A Illustration W. R. B. Oliver, Trans. & Proc. Roy. Soc. N.Z. 80 (1): t. 4 (1952). Multi-stemmed shrub to small tree 0.2–3 m tall. Branches: bark on old branches grey, finely fissured, young stems yellowish- to reddish-brown. Leaves juvenile and adult. Juvenile leaves spirally arranged along branches, spreading, light green to glaucous; lamina sheath 6–10
9–11 mm, shoulders tapering and margins ciliate in the upper half; lamina linear–triangular to lanceolate, 60–125
5.0–8.5 (–7.0) mm, margin minutely serrulate with 50–70 teeth per 10 mm. Adult leaves spreading, glaucous; lamina sheath 3–7
2.2–6.0 mm, striate, shoulders rounded to auricled and margin membranous with the top half ciliate; lamina subulate to linear–triangular, 12–52
1.0–3.5 mm, surfaces scabrid, prominently striated; margin serrulate with 60–100 teeth per 10 mm. Inflorescence a terminal spike on lateral branchlets; shorter than leaves, erect, lax, 5.5–14.0 mm long, oblong; inflorescence bracts overtopping flowers, glaucous, ovate–lanceolate at the base, (6–)8–18
0.7–2.0 mm, surfaces widely scabrid; margins serrulate. Flowers hidden by the leaves, 1–4, sessile; flower bracts shorter than flowers, narrowly ovate, 5–7
0.6–1.5 mm, adaxial surface rugose with a basal tuft of scabrid hairs; abaxial surface scabrid and rugose; margins serrulate. Sepals lanceolate to narrowly ovate, 4.5–6.0
1.5–2.0 mm, longer than corolla tube, striate, adaxial surfaces glabrous with the top half pubescent; abaxial surfaces pubescent; margins ciliate or ciliate in the upper half. Corolla white; corolla tube cylindrical, 3.5–5.0
2.4–2.5 mm; corolla lobes spreading to spreading horizontally, triangular, shorter than corolla tube, 1,8–2.0
1.5–2.0 mm, apices inflexed, subacute; glabrous. Stamens inserted in corolla tube in the upper third, filaments 163 I F G J H B E D C Fig. 149. Dracophyllum trimorphum. A. Flowering branch (1
). B. Sepal abaxial surface (5
). C. Flower-bract adaxial surface (5
). D. Laid-out corolla (5
). E. Lamina sheaths to show variation (5
). F. Stage one juvenile leaf (1
). G. Stage two juvenile leaf (1
). H. Adult leaf (1
). I. Ovary (10
). J. Flower (5
). Drawn from Venter 13779. Del. S. Venter. 164 Australian Systematic Botany S. Venter A B C D E F Fig. 150. Dracophyllum trimorphum. A. Habitat on coastal cliffs near Puponga Point, the type locality. B. Stem of mature plant. C. Wind pruned plants at Puponga Point. D. Flowering branch. E. Mature plant showing the short narrow leaves. F. Branches showing the three prominent stages in leaf formation. Photos: Christopher Hynes (A) and S. Venter (B–F). of shrubland. Plants that are exposed to salt spray and wind have a deformed growth habit and appear stunted. Phenology Flowering October–March. Etymology Refers to the three distinct growth stages of the species. Diagnostic features and notes Dracophyllum trimorphum is characterised by the three distinct growth stages evident in the shape and size of the leaves. Leaves of the first juvenile stage are large, becoming smaller in the middle juvenile stage and, ultimately, small and narrow in the adult stage. It is also characterised by glaucous adult leaves that are prominently striated with the basal part of the lamina covered in dense scabrid hairs, one- to fourflowered and 6–8 mm long inflorescence, adaxial surface of the flower bracts pubescent in the top half, sepals shorter than the corolla tube and the top of the ovary covered in short hairs that sometimes appear papillate when the scabrid hairs are very short. It can easily be separated from all other Dracophyllum species in the prominent three growth stages of the glaucous leaves and the ovary with a truncate apex covered in scabrid hairs. Taxonomic revision of Dracophyllum and Richea 170º Australian Systematic Botany 175º 165 180º −35º −40º −45º Fig. 151. Known distribution of Dracophyllum trimorphum, top of the South Island, New Zealand. The growth habit is variable and depends on the climatic factors such as strong wind and salt spray. Dracophyllum trimorphum grows as shrublets a mere 20 cm tall on the exposed ridges along the coast near the lighthouse at Pillar Point (Puponga). In more protected areas facing away from the sea (Puponga Point), it grows as a small many-branched tree 3 m tall. The shape of the leaf sheath is polymorphic. The adult lamina is very variable, even on the same branch (25–52
1.0–3.5 mm) and the inflorescence consists of one (rarely) or two to four flowers. The sepals are lanceolate to narrowly ovate and vary in size (4.5–6.0
1.5–2.0 mm), having the margin either ciliate in the upper half or it can be wholly ciliate. The corolla tube varies on the same plant from 3.5 to 5.0 mm long. The apex of the ovary is covered in hairs that vary from prominently scabrid to small and papillae-like but always present. Selected specimens NEW ZEALAND. South Island: Puponga, north-east of Wharariki Road south of Pillar Point Lighthouse, 29 Oct. 1998, Venter 13723 (CHR); Cape Farewell, Nguroa Bay Road, hilltop next to homestead, 30 Oct. 1998, Venter 13724 (CHR); Collingwood, West Whanganui Inlet, Echo Point, 12 Dec. 1998, Venter 13738 (CHR); Puponga, next to Fossil Island, Puponga Farm, 28 Mar. 1999, Venter 13779 (CHR). Dracophyllum urvillianum A.Rich., Essai Fl. N.Z.: 221 (1832) Type: New Zealand. Tasman Bay, on rocks, Jan. 1827. J.S.C. Dumont D’Urville s.n. (holo: P!; iso: W!). Illustration W. R. B. Oliver, Trans. & Proc. N.Z. Inst. 59: t. 6 (1928). 166 Australian Systematic Botany Small single-stemmed tree, 2–8 m tall. Branches: bark on old branches grey to greyish-brown, finely fissured, young stems reddish-brown. Leaves juvenile and adult. Juvenile leaves spirally arranged along branches, spreading to recurved; lamina sheath yellowish-green, 5–6
1.3–1.5 mm, truncate and margin membranous with the upper half ciliate; lamina linear–triangular, 79–145
(1.5–)2.3–3.7 mm, margins serrulate with 40–50 teeth per 10 mm. Adult leaves spreading to recurved; lamina sheath 3.6–9.0
2.5–3.0 mm, thinly coriaceous, shoulders truncate to auricled and margins membranous with the top half ciliate; lamina linear to linear–triangular, (33–)54–128
0.42–1.68 mm, adaxial surface sometimes shortly scabrid; margins serrulate with 45–60 teeth per 10 mm. Inflorescence a terminal raceme on lateral branchlets; shorter than leaves, erect, lax, 14–23 mm long, oblong. Inflorescence bract overtopping flowers, ovate–lanceolate, (15.4–)31.0–35.0
0.5–0.6 mm, surfaces rugose; margins serrulate. Flowers hidden by leaves, 2–4 (–5), pedicellate; flower bracts overtopping flowers, narrowly ovate, 11.3–15.6
0.4–0.5 mm; margins ciliate; pedicel 0.5–0.7 mm long. Sepals ovate–lanceolate, 5.5–7.0
1.2–3.0 mm, equalling corolla tube, adaxial surface with the top half pubescent; margins ciliate in the upper half; apices acute. Corolla white; corolla tube narrowly campanulate, widened at mouth, 3.5–5.0
1.5–2.0 mm; corolla lobes spreading horizontally to reflexed, ovate, shorter than corolla tube, 1.9–2.0
1.3–1.5 mm, apices acute, adaxial surface papillate. Stamens inserted on the corolla tube in the upper third, filaments 0.5–0.8 mm long; anthers included, oblong, light yellow and 0.5–1.0 mm long. Ovary globose, 1.0–1.5
1.0–1.3 mm, apex round to truncate; nectary scales rectangular, 0.5–0.6
0.4–0.5 mm; apices subacute to retuse; style included, 1.3–2.0 mm long, glabrous, not lengthening in fruit; stigma capitate. Fruit with the old sepals widely spreading, light brown, 2–3
2.0–2.5 mm, oblong; apex truncate. Seeds yellowish-brown, ovoid, 0.95–1.0 mm, testa slightly reticulate (Fig. 152, 153). Distribution and ecology New Zealand endemic. Occurs in the eastern part of northwestern Nelson and in the Marlborough Sounds on the South Island (Fig. 154). Dracophyllum urvilleanum occurs from sea level up to 1158-m elevation on gentle (5–30) slopes as well as on coastal cliffs. The surrounding vegetation consists of lowland to montane shrubland and forest. Soils are brown clay or gritty greyish-brown clay loam derived from shale (Whangamoa Saddle, Nelson), schist or serpentinite (Cobb Reservoir) or light brown gritty sandy loam and brown loam derived from granite (Abel Tasman National Park). Plants are usually in light to deep shade in the forest and rarely fully exposed to sunlight (Whangamoa Saddle, Nelson). Phenology Flowering November–March. Etymology Named for Rear Admiral Jules Sébastien César Dumont d’Urville (1790–1842), French explorer and naval officer, S. Venter A F G H I D B E C Fig. 152. Dracophyllum urvilleanum. A. Flowering branch (1
). B. Inflorescence bract (5
). C. Sepal adaxial surface (5
). D. Lamina sheaths to show variation (5
). E. Laid-out corolla (5
). F. Juvenile leaf (1
). G. Adult leaf (1
). H. Ovary (10
). I. Flower (5
). Drawn from Venter 13799. Del. S. Venter. who explored the southern and western Pacific, Australia, New Zealand and Antarctica. Diagnostic features and notes Dracophyllum urvilleanum is characterised by the thinly textured wide juvenile leaves, truncate to auricled shoulders of the adult lamina sheath, long linear adult leaves, short (14–23 mm) few-flowered (2–4) raceme, flower bract overtopping the flower, narrowly campanulate corolla tube, style longer than the ovary and the fruit enclosed in widely spreading persistent sepals. Dracophyllum urvilleanum is similar to D. oliveri, but differs in the narrower thinly textured juvenile leaves, longer and narrower drooping adult leaves, fewerflowered (mostly 2 or 3 flowers, not 5–10) racemes and truncate, not round, fruit apices. Dracophyllum urvilleanum is also similar to D. filifolium, with the differences being discussed Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A 167 B C D E Fig. 153. Dracophyllum urvilleanum. A. Habitat at the type locality in the Abel Tasman National Park. B. Fruiting branch. C. Mature plant from the type locality in Abel Tasman National Park (Venter 13799). D. Young plant showing the large juvenile leaves, Kaiteriteri (Venter 13793). E. Habitat in low forest along the coast. F. Flowering branch (Venter 13799). Photos: Pat Leahy (A), S. Venter (B–F). in detail under D. filifolium. Dracophyllum urvilleanum at Kaiteriteri was recorded having a second flush of flowers during April 2003, after a long dry and hot period followed by heavy rain (S. Venter, pers. obs.). Bullock et al. (1983) called this pattern of flowering ‘episodic flowering’ and it can be regarded as an adaptation to survival of the plant when under stress caused by severe climatic conditions. Some variation occurs in the size of the juvenile (79–145
2.3–3.7 mm) and adult lamina (70–128
0.7–1.3 mm). Plants from the population at the serpentine quarry, Cobb Reservoir (Venter 13772), are unique in having very long adult leaves (109–128 mm). The length of the inflorescence varies among populations (14–23 mm) and the flower bracts vary in length from 11.3 to 15.6 mm, even within the same population. Corolla-tube length is short (3.5–4.0 mm) in populations growing at high altitudes (1000–1158 m), compared with those that grow close to sea level (4–5 mm long). Selected specimens NEW ZEALAND: South Island: Anatoki Scenic Reserve, Oct. 1974, Kelly & Kelly s.n. (CHR); Abel Tasman National Park, Marahau, Freshwater Cove, 2 Mar. 2000, Venter 13799 (CHR); Torrent Bay, Nov. 1974, Druce s.n. (CHR); Takaka, Cobb Reservoir, Serpentine Quarry, 23 Mar. 1999, Venter 13772 (CHR); Tasman Bay, Otowhero Inlet Scenic Reserve, May 168 Australian Systematic Botany S. Venter 170º 175º 180º −35º −40º −45º Fig. 154. Known distribution of Dracophyllum urvilleanum, top of the South Island, New Zealand. 1974, Kelly & Kelly s.n. (CHR); Kaiteriteri, 3 km on road to Marahau, 2 Dec. 1998, Venter 13733 (CHR); ibid., Venter 13793 (CHR); Sandy Bay, 12 Nov. 1968, Talbot s.n. (CHR); Kaiteriteri, beach north of Riwaka and Motueka River outlets, 4 July 1971, Simpson s.n. (CHR); W of Astrolobe Harbour, Mar. 1921, Gibbs s.n. (CHR, WELT); Nelson, Whangamoa Saddle, 500 m along Slaters Road, 5 Jan. 1999, Venter 13743a (CHR); ibid. Venter 13782 (CHR); Picton, Nov. 1928, McMahon s.n. (CHR). Nelson Lakes National Park, D’Urville River in gorge, 18 Feb. 1964, Simpson 4246 (CHR). Dracophyllum subgenus Cystanthe (R.Br.) S.Venter, comb. et stat. nov. Cystanthe R.Br., Prodr. 555 (1810); Richea sect. Cystanthe (R.Br.) Benth., Fl. Austral. 4: 258 (1868). Type: Dracophyllum sprengelioides (R.Br.) S.Venter. Flowers solitary in a simple cluster crowded in terminal heads and each flower in the axil of a persistent bract. Key to the species of subgenus Cystanthe 1. Leaves 8–25 mm long; flowers in erect terminal heads .......................2 Leaves 20–40(–80) mm long; flowers in drooping terminal heads ........ ................................................................................... D. tasmanicum 2. Leaves greater than 3 mm wide; nectary lobes absent .........................3 Leaves less than 2 mm wide; nectary lobes present ........D. laciniatum 3. Inflorescence maturing acropetally; filaments distally thickened; anthers distinctly bilobed on dehiscence .................................. D. procerum Inflorescence maturing basipetally; filaments slender; anthers dehiscing by single split ........................................................ D. sprengelioides Taxonomic revision of Dracophyllum and Richea Dracophyllum tasmanicum S.Venter, nom. nov. [non D. milliganii Hook. (1852)] Bentham, Fl. Australiensis 4: 259 (1869) Australian Systematic Botany 169 Etymology Named for the state where it occurs. Diagnostic features and notes Cystanthe milliganii F.Muell., Fragm. 1: 38 (1858). Pilitis milliganii Hook.f., Fl. Tasm. 1: 226, t.83 (1859); Richea milliganii (Hook.f.) F.Muell., Fragm. 6: 69 (1868). Type: Tasmania, Mount Sorrell, Macquarie Harbour, VDL, elev. 3000 feet [~914 m], 15 Jan. 1847, J. Milligan 806 (holo: K!; iso: K!). Illustration Y. Menadue and R. Crowden, Austral. Syst. Bot. 13: t. 2 (2000). Erect, sparsely branched shrub, 1.5–2.3 m tall. Branches older branches with distinct annular leaf scars. Leaves persisting and crowded at the ends of younger branches; erect–spreading; lamina sheath weakly developed, transversely oblong; lamina narrow-lanceolate, 20–40(–80)
3–6 mm, margins entire or scabrous; lamina apex acute. Inflorescence a terminal head on main and lateral branches; shorter than leaves, drooping, dense, 2.5–3.5 cm long, 2 cm wide. Flower bract overtopping the flowers, persistent, lanceolate, 15–40
5–6.5 mm, brown, margins smooth, gradually tapering to an acute apex. Flowers 8–15, pedicellate; flower bracts leaf-like; pedicel less than 1 mm long, glabrous; bracteoles persistent, 2–4, mostly 3, at the base of each flower, 7–9 mm long, narrow, keeled; margins entire to minutely ciliolate. Sepals narrowly elliptic–ovate, 6.5–8.5 mm long, shorter than operculum; margin entire or minutely ciliolate in the upper half; apices acute. Corolla creamyyellow; operculum narrowly conical, often flattened dorsiventrally, 12–17
2.5–4 mm; corolla lobes very short, blunt, between which the stigma often protrudes (in some instances the operculum may persist, then stamens and style push through splitting the operculum). Stamens inserted at the base of the operculum, filaments pale straw yellow, 20–25 mm long; anthers 2 mm long, versatile, slightly membranous, after dehiscence folding back around filament before anthesis. Ovary globose, 2 mm in diameter, apex round; nectary scales transversely oblong, 1.4–1.5
1–1.2 mm; apices truncate or emarginate; style 20–25 mm long, glabrous, level with anthers; stigma indistinct. Fruit have the old sepals spreading, globose to depressed-globose, 2.0–2.1
2.5–3.5 mm; apex round (Fig. 155, 156). Distribution and ecology Tasmania endemic. Occurs mainly in the south-western mountains as far north as the Mount Read group (Fig. 157). It occurs near summits of mountains in peaty soils among boulders or in wet places in open heathland. At lower elevations, it occurs in subalpine woodlands, in margins of scrub–forest, often in fringing button grass plains (Menadue and Crowden 2000). Phenology Flowering from July to October. A sparsely branched shrub with the leaves crowded at the ends of the younger branches, leaves 20–40(–80) mm long; flowers in drooping terminal heads, flower bracts mostly brown, sepals creamy-green, corolla creamy-yellow; operculum 12–17
2.5–4 mm, separating and falling earlier in development than in other species from Dracophyllum subgenus Cystanthe, often flattened dorsi-ventrally, in some instances the operculum may persist, then stamens and style push through splitting the operculum. Selected specimens TASMANIA: Jubilee Range. 16 Jan. 1985, Buchanan 5228 (HO); north of Mount Bowes, 4 May 1985, Collier 458 (HO); Mount Hayes, western Arthur Range, 6 July 1986, Collier 1991 (HO); Lake Belcher, Mount Field National Park, July 1929, Comber s.n. (HO 5630 and 5625); Denison Range, 2 Jan. 1947, Elliot s.n. (HO 5634); Double Peak, 10 July 1978, Jarman s.n. (HO 30391); Tim Shea near summit, 13 Jan. 1980, Menadue s.n. (HO 126003); Mount Sprent below summit, 21 Jan. 1990, Menadue & Crowden s.n. (HO 126533); Frodsham Pass, 22 Feb. 1990, Menadue & Crowden s.n. (HO 126534) and 30 July 1990, (HO 126590); Mount Sorell, 15 Jan. 1947, Milligan s.n. (HO 5629, BM 35647 and K); Mount Mueller, Fossil Lake, 6 Dec. 1975, Moscal s.n. (HO 78068); Hamilton Ranage, 21 Oct. 1987, Read & Carpenter s.n. (HO 126002); Hartz Mountain, Dec. 1903, Rodway s.n. (HO 5632). Dracophyllum laciniatum S.Venter, nom. nov. [non D. acerosum Berggr. (1877)] Pilitis acerosa Lindl., Introd. Nat. Syst. Edn 2: 443 (1836); Candolle Prod. 7: 769 (1838); Hook.f., Fl. Tasm. 1: 265. t. 82! (1859); Cystanthe acerosa (Lindl.) F.Muell., Fragm. 1: 38 (1858); Richea acerosa (Lindl.) F.Muell., Fragm. 6: 69 (1868). Type: Van Diemens Land, 17 Jan. 1845, Gunn 307. Herb. J. Lindley (1837) (lecto: photo!, CGE; isolecto: K!, MPU!), designated by Menadue and Crowden (2000). Illustration Y. Menadue and R. Crowden, Austral. Syst. Bot. 13: t. 3 (2000). A small erect and spindly shrub growing 60–120 cm tall. Branches older stems with distinct annular leaf scars. Leaves imbricate, widely to erect–spreading; confined to ends of stems; lamina sheath weakly developed, transversely oblong; lamina linear–lanceolate, 8–14
1–2 mm, margin scaberulent; lamina apex subulate, tapering to a rigid, keeled point. Inflorescence a terminal head on main and lateral branches; shorter than leaves, erect, dense, 0.7–1.2
1 cm. Flowers up to 10; pedicellate; flower bracts leaf-like, same length to shorter than flowers, persistent, broadly ovate–cordate, 5–6
2.5–3.0 mm, brown, margins ciliolate; tapering to an acute or acuminate apex; pedicel less than 0.5 mm long, glabrous; bracteoles persistent, 3–5 at the base of each flower, 2.5–3.0 mm long, strongly keeled; margin ciliolate. Sepals white, ovate, 3–4
1.5 mm; shorter than operculum; margin ciliolate; apices acute to 170 Australian Systematic Botany S. Venter A B C F D E I G H Fig. 155. Dracophyllum tasmanicum. A. Branch showing the growth habit. B. Bracteoles. C. Inflorescence bract. D. Mature leaf. E. Operculum. F. Parts removed to show nectary scales and ovary. G. Operculum removed to show flower. H. Lamina base. I. Inflorescence a drooping cluster of flowers. Del. Y. Menadue. acuminate. Corolla white, cream or pinkish; operculum bluntly conical, 5–6
2.5 mm; corolla lobes not developed. Stamens inserted at the base of the operculum; filaments creamy-white, 4–6 mm long; anthers <1 mm long, versatile. Ovary globose, 1.4–1.5 mm long and wide, apex round; nectary scales 1.0–1.5
0.4–0.5 mm, deeply bilobed or laciniate; style 1.0–1.5 mm long, glabrous; stigma capitate, 2–4 mm long. Fruit depressed-globose, 1.5–1.7
2.5–3.0 mm (Fig. 158, 159). Distribution and ecology This is a Tasmanian endemic distributed mainly on the Central Plateau, also on Mount Field and Ben Lomond (Fig. 160). Dracophyllum laciniatum is frequently found in alpine and subalpine heath, growing in poorly drained areas such as riverbanks, plains or within bolster moor communities consisting primarily of Abrotanella forsterioides (Hook.f.) Benth., Donatia novae-zelandiae Hook.f., Dracophyllum minimum F.Muell. and Pterygopappus lawrencei Hook.f. (Kirkpatrick 1997; Menadue and Crowden 2000). It occurs generally in a variety of drier habitats, mainly occupying ridge tops among boulders, slopes among grass and heath, or on the edges of eucalypt woodland. A fire-sensitive species with limited ability to resprout or layer (Kirkpatrick and Bridle 2016). Etymology Refers to the laciniate apices of the nectary scales. Diagnostic features and notes The leaves are distinctly narrower (less than 2 mm wide) than in any other member of the subgenus Cystanthe. The apices of Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 171 A B C Fig. 156. Dracophyllum tasmanicum. A. Habitat on the Denison Range, Tasmania. B. Flowering branch of the large flower form. C. Flowering plant showing the drooping inflorescences. Photos: Natalie Tapson (A, B) and James Wood (C). the nectary scales are prominently laciniate; hence, the specific epithet. Selected specimens TASMANIA: Lake Salome, Walls of Jerusalem, 25 Jan. 1983, Brown 118 (HO); valley above Lake Little, Cradle Mountain, 31 Jan. 1982, Buchanan 883 (HO); near Junction Lake, 19 Jan. 1990, Buchanan 11613 (HO); Clumner Bluff, 20 Apr. 1985, Collier 423 (HO); Ben Nevis, 18 Mar. 1986, Collier 1273 (HO); Mount Barrow, 29 Dec. 1964, King s.n. (MEL); Western Mountains, 10 Feb. 1843, Lawrence 310 (K); Lake Augusta, Central Plateau, 6 Oct. 1979, Menadue s.n. (HO 125971); head of Lake Fenton, 27 June 1980, Menadue s.n. (HO 125967); Projection Bluff, 14 Dec. 1989, Menadue & Crowden s.n. (HO 125973); Devils Den, 18 Mar. 1984, Moscal 7141 (HO); base of Coalmine Crag, Ben Lomond National Park, 25 Jan. 1983, Orchard 5802 (HO); Mount Rufus track, Lake Saint Clair, 5 Dec. 1981, Powell 1621 (HO); Mount Field East, 12 Jan. 1978, Smith 373 (HO). 172 Australian Systematic Botany Fig. 157. S. Venter Known distribution of Dracophyllum tasmanicum in Tasmania. Dracophyllum procerum (F.Muell.) S.Venter, comb. nov. Cystanthe procera F.Muell., Fragm. 1: 38 (1858); Richea procera (F.Muell.) F.Muell., Fragm. 6: 68 (1868). Type: Tasmania, Southport, A. Oldfield s.n. (holo: K!). Illustration Y. Menadue and R. Crowden, Austral. Syst. Bot. 13: t. 3 (2000). Erect, sparingly branched shrub, 0.6–3 m tall. Stems older stems bare, with annular scars. Leaves along branches, spreading, recurved; lamina sheath weakly developed, transversely oblong; lamina ovate–lanceolate, 8–20–(25)
4–9 mm, margins narrowly hyaline, <0.15 mm wide, with serrulations; older leaves with a flattened ridge on the abaxial surface near the apex; lamina apex acute, not pungent. Inflorescence a terminal head on main and lateral branches; longer than leaves, erect, dense, 2–3 cm long, up to 2 cm wide. Flowers 12–20, pedicellate; flower bracts shorter to equalling flowers, persistent, leaf-like, green, turning to brown, 6–9 mm long; pedicel <1 mm long, glabrous; bracteoles persistent, 4.0–5.5 mm long, translucent–white, outer 2 prominently keeled with minutely serrulate margins, up to 4 sepal-like inner bracteoles. Sepals translucent–white with pink tips, ovate–lanceolate, 4–6
1.5–2 mm; shorter than operculum; margins entire, apices acute. Corolla dark pink near apex grading to white near the base; operculum narrow conical, flattened dorsiventrally; 8–9
2–2.4 mm; corolla lobes very short, not spreading. Stamens inserted at the base of the operculum, filaments 8–10 mm long, thickened and papillose in the upper half; anthers 2.3–2.8 mm long, splitting longitudinally dorsally and ventrally as far as the filament then continuing down ventral surface in very mature anthers, versatile. Ovary globose, 2–2.2
2.4–2.8 mm, apex round to truncate, pilose, cusped; nectary scales absent; style much longer than ovary; stigma indistinct, level with top of anthers. Fruit ellipsoid, 3–3.5 mm in diameter, the apex round to slightly truncate (Fig. 161, 162). Distribution and ecology Dracophyllum procerum occurs in the central and southern regions of Tasmania (Fig. 163). It grows at elevations in Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A 173 B C F D G E Fig. 158. Dracophyllum laciniatum. A. Flowering branch. B. Flower bracts. C. Leaves on sterile branch. D. Flower. E. Floral parts removed to show nectary scales. F. Operculum removed to show floral structure and persistent basal ring. G. Operculum. Del. Y. Menadue. excess of 400 m, but below the subalpine region. Dracophyllum procerum occurs in scrubland and open forests, nearly always in open areas on poorly drained soils or on grey sandy soils in boulder-strewn areas (Menadue and Crowden 2000). Phenology Flowering from October to early December. Etymology From the Latin procerus referring to the tall stature of the plants. Diagnostic features and notes Pedicels subtended by leaf-like bracts, flowers in drooping terminal heads, nectary scales absent, filaments thickened in the upper half and the anthers distinctly bilobed. Selected specimens TASMANIA: Huon Road, 16 Oct. 1984, Atkinson 128 (HO); saddle between One O’clock Hill and Platform Peak, 20 Jan. 1981, Brown 152 (HO); Croswell’s Road, Mount Lloyd, 20 Oct. 1983, Buchanan 1241 (HO); south-eastern slope Mount Hobbs, 1 Nov. 1984, Buchanan 3799 (HO); Franklin River, 17 Nov. 1986, Collier 1865 (HO); Wombat 174 Australian Systematic Botany A C E S. Venter B D F Fig. 159. Dracophyllum laciniatum. A. Habitat at Jacobs Ladder. B. Sterile branch showing the needle-sharp leaves. C. Flowering plant at Cradle Mountain. D. Flowering branch with most of the corollas still intact. E. Mature flowering plant from the Devil’s Gullet. F. Flowering branches showing the narrow, short leaves. Photos: Russell Cummings (A, C–F), Natalie Tapson (B) and Tim Rudman (D). Glenn, Lyall Highway, 19 Oct. 1983, Crowden s.n. (HO 126582); Hobart, 20 Nov. 1840, Gunn 1213 (K); top of Western Mountains, 20 Dec. 1843, Gunn 1213 (K); Mount Field National Park, 12 Dec. 1990, Menadue & Crowden s.n. (HO 400398); in saddle between Grey Mount and Mundys Hill, Snug Tiers, 31 Oct. 1983, Moscal 997 (HO); Woodslake Road, 15 Nov. 1981, Moscal 867 (HO); nameless Tarn west of Lake Lea, 25 Nov. Taxonomic revision of Dracophyllum and Richea Fig. 160. Australian Systematic Botany 175 Known distribution of Dracophyllum laciniatum in Tasmania. 1982, Moscal 1055 (HO); 3–4 km north of Cynthia Bay camping area, Lake Saint Clair, 4 Dec. 1981, Powell 1612 (HO); track up Mount Rufus, 5 Dec. 1981, Powell 1620 (HO). Dracophyllum sprengelioides (R.Br.) S.Venter, comb. nov. Cystanthe sprengelioides R.Br., Prodr. 555 (1810); sprengelioides (R.Br.) F.Muell., Fragm. 6: 68 (1868). Richea Type: Tasmania, Mount Wellington near the Derwent River, Apr. 1804, R. Brown s.n. (lecto: BM 35648!; isolecto: K!, MEL!), designated by Menadue and Crowden (2000). Illustration Y. Menadue and R. Crowden, Austral. Syst. Bot. 13: t. 5 (2000). Erect to compact, sparsely branched shrub 0.3–1.2 m tall. Branches older stems without leaves, with distinct annular scars. Leaves along branches; rigid, erect–spreading; lamina sheath weakly developed, wedge-shaped. Lamina broadly ovate–lanceolate, 8–12
3–5 mm; with a flattened ridge below apex on abaxial surface; margins scaberulent or ciliolate; lamina apex acute, pungent. Inflorescence an erect terminal head on the main and lateral branches; longer than leaves, dense, 2 cm long and wide. Flowers 8–20, pedicellate; flower bracts leaf-like, persistent, brown, broadly ovate–cordate, 4–7 mm long; pedicel less than 0.5 mm long, glabrous; bracteoles persistent, 2–6 at the base of each flower, outer 2 keeled and winged; 4–5 mm long. Sepals ovate–lanceolate, 4–5
1.0–1.5 mm wide; pale green, translucent, shorter than the operculum; margin ciliolate; apices acute. Corolla creamy-white, pale green near 176 Australian Systematic Botany S. Venter A B C E D F G H Fig. 161. Dracophyllum procerum. A. Inflorescence. B. Bracts. C. Flower. D. Leaf abaxial surface. E. Operculum removed to show flower. F. Operculum. G. Operculum and sepals removed to show shape of ovary. H. Anther and thickened filament. Del. Y. Menadue. apex; operculum narrowly conical with a shoulder below apex, 7–8
2.8–3.3 mm; corolla lobes not developed. Stamens inserted at the base of the operculum, filaments yellow–cream; almost uniform in thickness, 5–6 mm long, glabrous or minutely papillose; anthers 2 mm long, opening by a single longitudinal slit from apex to base, versatile. Ovary globose, 1.5–1.8
2–2.5 mm wide., glabrous; apex round; nectary scales absent; style 3–4 mm long, as long as or longer than stamens; stigma indistinct. Fruit depressed-globose, 3–4 mm in diameter; apex round (Fig. 164, 165). Distribution and ecology Tasmanian endemic, in all mountainous regions of the west, south-west, north-east and Central Plateau (Fig. 166). Predominantly in alpine areas, but extends to subalpine woodlands in rocky, boulder-strewn areas. Phenology Flowering from November to January. Etymology Having the appearance of a Sprengelia. Diagnostic features and notes Anthers dehiscing by a single slit, nectary scales absent, with the leaves generally smaller and more rigid than those of D. procerum. The inflorescence of D. sprengelioides matures basipetally and has creamy-white flowers, whereas Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany B A C 177 D Fig. 162. Dracophyllum procerum. A. Habitat on Mount Rufus. B. Flowering branch showing the characteristic flattened and papillose filaments. C. Flowering plant in habitat on Mount Wellington. D. Flowering branch showing the short and wide leaves. Photos: James Wood (A), John Tann (B, C) and Tim Rudman (D). it matures acropetally in D. procerum and the flowers are tipped with pink. D. procerum has thickened, papillose and yellow filaments and distinctly bilobed anthers, whereas D. sprengelioides has smooth, slender, creamy-white filaments and the anthers are not bilobed after dehiscence. D. sprengelioides has a glabrous ovary, whereas it is pilose in D. procerum. Dec. 1978, Jarman s.n. (HO 30361); Tyndall Range, western coast, 8 Dec. 1980, Macphail s.n. (HO 37067); Eagle Tarn, Mount Field National Park, 7 Jan. 1980, Menadue s.n. (HO 125966); near summit of Projection Bluff, 16 Dec. 1989, Menadue s.n. (HO 126529); low hills between Western and Breton Rivulets, Feb. 1981, Moscal 539 (HO); track to Marion’s Lookout, Cradle Mount National Park, 29 Nov. 1981, Powell 1547 (HO); Hounslow Heath, 30 Nov. 1981, Powell 1564 (HO); Western Tiers, 15 Dec. 1908, Rodway s.n. (HO 5699); Hartz Mountains National Park, 100 m below summit, 1 Feb. 1983, Short 1883 (HO). Selected specimens TASMANIA: Mount Wellington, Brown p.p. (K); summit of Frenchman’s Cap, 8 Jan. 1981, Buchanan 466 (HO); Mount Barrow, 27 Dec. 1959, Burns 229 (HO); summit Mount Hobhouse, 20 Dec. 1986, Collier 2092 (HO); Lake Australia, Mount Gell, 16 Nov. 1986, Collier 1883 (HO); Mount Wellington, 8 May 1839, Gunn 290 (BM 35636); Mount Wellington, 7 Jan. 1841, Gunn 290 (BM 35637); Mount Sprent, 10 Dracophyllum subgenus Richea (R.Br.) S.Venter, comb. et stat. nov. Richea R.Br., Prodr. 555 (1810); Richea sect. Dracophylloides Benth. Fl. Austral. 4: 258–259 (1868). 178 Australian Systematic Botany S. Venter Fig. 163. Known distribution of Dracophyllum procerum in Tasmania. Type: Dracophyllum desgrazii (Hombr. ex Decne.) S.Venter. Inflorescence a compound spike or panicle with the flowers clustered along a central axis. Each flower cluster is in the axil of a caducous bract. Key to the species of subgenus Richea 1. Inflorescence a slender spike-like panicle with little lateral branching ..........................................................................................2 Inflorescence a broad panicle with extensive lateral branching............5 2. Inflorescence matures from the top down (basipetal); nectary lobes absent........................................................................ D. victorianum Inflorescence matures from the base up (acropetal); nectary lobes present...............................................................................................3 3. Inflorescence compact with internodes of equal length; older branches with flaky bark............................................... D. persistentifolium Inflorescence loose with proximal internodes elongated; older branches with prominent annular scars ...........................................................4 4. Dense compact shrub; leaves erect or spreading, 2–4 cm long; inflorescence lateral branches bear 6–10 flowers; operculum 5–8 mm long .............................................................. D. continentis Open divaricate shrub; leaves usually recurved, 3–6 cm long; inflorescence lateral branches bear 3–5 flowers; operculum 3–5 mm long ......................................................................D. gunnii 5. Inflorescence axillary; leaves >60 cm long............... D. pandanifolium Inflorescence terminal; leaves <35 cm long..........................................6 6. Shrub 1.5–5 m high; old branches with annular scars; corolla white ...............................................................................D. desgrazii Shrub usually <1 m high; old branches with rough bark; corolla pink, orange or red....................................................................D. alpinum Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 179 B A D C E F G Fig. 164. Dracophyllum sprengelioides. A. Flowering branch. B. Flower bracts. C. Flower. D. Leaf abaxial surface. E. Operculum removed to show flower. F. Operculum. G. Operculum and sepals removed to show ovary. Del. Y. Menadue. Dracophyllum victorianum (Menadue) S.Venter, comb. nov. Richea victoriana Menadue, Muelleria 8(3): 317 (1995). Type: Australia. Victoria, Nine Mile Road, 0.5 km north of Block 10 Road, Thompson River headwaters, 40 km east of Warburton, alt. 1010 m, 37470 S, 146100 E, 26 Dec. 1992, J. Davies s.n. (holo: HO; iso: MEL!). Richea gunnii sens. Walsh, Victorian Nat. 104(3) (1987), non Hook.f. (1847). Illustration Y. Menadue and R. Crowden, Austral. Syst. Bot. 13(5): 786 (2000). Erect, multi-branched shrub, 0.3–2 m tall. Branches older branches without leaves, with prominent annular scars. Leaves clustered in the top 10–40 cm of branches, imbricate and spreading; lamina sheath weakly developed, transversely oblong, lacking distinct shoulder; lamina narrowly triangular, 30–100
5–11 mm, flat to concave; margin maturing basipetally; an apical, slender panicle, overtopping the leaves, 180 Australian Systematic Botany S. Venter A B C D E F Fig. 165. Dracophyllum sprengelioides. A. Habitat in Hartz Mountains National Park. B. Inflorescence with all the operculae shed. C. Flowering plant in habitat. D. The characteristic short and wide leaves. E. Erect stems with the apical inflorescences. F. Inflorescences with some of the operculae still intact, plant from Neika. Photos: Natalie Tapson (A, D), Tim Rudman (B), Russell Cumming (C) and James Wood (E, F). erect, dense, 30–130
20–30 mm; sparsely branched; basal inflorescence branch up to 15 mm long, suberect; inflorescence bracts overtopping flowers, subcoriaceous. Flowers in groups of 3–20 at the base of the inflorescence, pedicellate; bracteoles 1–2, caducous, shorter than the flower, narrow linear, 2–4 mm long; pedicels 1 mm long, glabrous. Sepals creamy-white, broadly ovate, 1.6
1.8 mm, shorter than the operculum, glabrous, obtuse. Corolla white; operculum narrowly ovoid–conical, 3–4
1.5–2 mm; corolla lobes not developed. Stamens hypogynous, filaments 2–3.5 mm long, cream-coloured, attached near top of anthers; anthers 1–1.5 mm long. Ovary globose, 0.7–0.9
0.9–1 mm, glabrous, apex rounded; nectary scales absent; style 1.5–2 mm long, tapering; stigma indistinct, reaching 2/3 height of stamens. Fruit a loculicidal capsule, depressedglobose, 2–3 mm in diameter, apex round, glabrous (Fig. 167, 168). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 181 Fig. 166. Known distribution of Dracophyllum sprengelioides in Tasmania. Distribution and ecology Australia, endemic to the Baw Baw Plateau and Blue Range, between Marysville and Taggerty in Victoria (Fig. 169). It is locally abundant in subalpine, wet heath or scrubland, or on the fringes of cool-temperate rainforest near streams and bogs (Menadue and Crowden 2000). Phenology Flowering from late November to early January. Etymology Named after the State Victoria. Diagnostic features and notes Dracophyllum victorianum is similar to D. gunnii, but differs in being larger and more robust with longer and wider leaves (50–100
5–11 mm compared with 30–60
5–7 mm). Dracophyllum victorianum inflorescence also has more flowers on the lateral branches (3–20 compared with 3–5), no nectary scales, glabrous inflorescence axis and lateral branches and maturing basipetally. Selected specimens VICTORIA: Nine Mile Road, 15 May 1984, Crowden s.n. (HO 308263); 8 Feb. 1989, Crowden & Menadue s.n. (HO 308264); 26 Dec. 1992, Davies s.n. (HO 308234); near Scout Hut between Mount Erica summit and car park, 13 July 1985, Albrecht 1846 (MEL); Thompson Forest Wildlife Reserve, 24 Nov. 1982, Beauglehole ACB 71727 & Beardsell (MEL); Upper Thompson River at Newlands Road crossing, Jan. 1992, Davies s.n. (HO 132105); 5 km north-north-east Mount Margaret, upstream from crossing of Blue River Road on Storm Creek, 23 Mar. 1985, Forbes 2820 (MEL); 50 m south of J. W. McMahon Ski Lodge, Captain Hurley Rover Crew, 6th Morwell, south-eastern slope of Mount Erica, 7 Dec. 1984, Salasoo 6 (MEL); Upper Thompson River catchment, Newlands Road extension, 4 km west from Rocky Knob, 7 Dec. 1981, Walsh 694 (MEL); Upper Royston River, 5 May 1963, Willis s.n. (MEL). 182 Australian Systematic Botany A S. Venter B C D Dracophyllum persistentifolium S. Venter, nom. nov. [non Dracophyllum scoparium Hook.f. (1844)] Richea scoparia Hook.f., London J. Bot. 6: 273 (1847). Type: Tasmania, Mount Wellington, 1 Mar. 1839, Gunn 292 (lecto: K!). Specimen in the lower right-hand corner designated by Menadue and Crowden (2000). Richea angustifolia B.L.Burtt, Curtis’s Bot. Mag. 163: tab. 9632 (1941). E Type: Tasmania, Mount Sorrel, Macquarie Harbour, 31 Dec. 1846, J. Milligan 746 (holo: K!; iso: MEL2186142!) F Illustrations H I G Fig. 167. Dracophyllum victoriana. A. Inflorescence. B. Operculum. C. Inflorescence bract. D. Leaf adaxial surface. E. Floral parts removed to show ovary. F. Operculum removed to show flower. G. Branching habit. H. Sepal. I. Cluster of flowers. Del. Y. Menadue. A Y. Menadue and R. Crowden, Austral. Syst. Bot. 13(5): t. 7 (2000). Rigid, erect, much-branched shrub commonly compact, 0.4–1.0 m tall in exposed places but up to 3 m tall and more open in sheltered vegetation. Branches: bark on older branches peeling, reddish-brown, and leaf scars rarely noticeable. Leaves crowded along branches, persisting for several years, decaying while still on the branches, erect, spreading to recurved, imbricate, rigid; lamina sheath transversely oblong; lamina narrowly lanceolate, 30–70(–200)
2.5–5(–9) mm; surfaces glabrous; margins scabrous; apex acute, pungent. Inflorescence a spike-like panicle, longer than leaves, erect, dense, 40–100 (–140) mm long, 2 cm wide, maturing acropetally; rachis and lateral branches pubescent; inflorescence bracts caducous, overtopping flowers, pinkish-green turning straw-brown, broad based, 30–200
2.5–10 mm in the middle, surfaces glabrous, apices acute, pungent. Flowers 30–300+, in groups of 3–6(–14) at the base of the inflorescence, pedicellate; bracteoles 2 or 3, B Fig. 168. Dracophyllum victoriana. A. Mature flowering plants in habitat near Thompson River, Baw Baw National Park. B. Mature inflorescence maturing basipetally. Images A and B by Russell Best. Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 183 Fig. 169. Known distribution of Dracophyllum victoriana, Victoria, Australia. caducous, shorter than flower, 3–4
0.5–1 mm, keeled; pedicels 1–2(–3) mm long. Sepals pale green to orange, pink to dark pink, broadly ovate, 1.5–2.5 mm long and wide; shorter than the operculum, margin entire; apex acute. Corolla pink to orange, dark red, or white with pink tips; operculum obovoid–oblong, 7–10
3–3.5 mm, bluntly pointed and sometimes lightly depressed at apex. Stamens inserted at the base of the operculum, filaments 3.5–5 mm long, dull red, articulated near the base; anthers rectangular, 1 mm long, versatile. Ovary depressed-globose, 1.3–2.2
1.5–2.5 mm, glabrous, apex round; nectary scales elliptical-truncate, 0.75–1.0
0.3–0.4 mm, apices truncate to emarginate; style 1.5–2 mm long, glabrous, reaching 1/2 to 2/3 length of stamens; stigma capitate. Fruit depressed-globose, 2–2.2
2–3 mm; apex round (Fig. 170, 171). Distribution and ecology Tasmania endemic, in all the mountainous regions in the west, south-west, north-east and on the Central Plateau (Fig. 172). Dracophyllum persistentifolium occurs in a variety of montane habitats above 700 m. Widespread in alpine regions in association with Olearia algida Wakef., O. ledifolia (DC.) Benth., Orites acicularis R.Br. and Epacris serpyllifolia R.Br. In heathland, it commonly occurs with Leptospermum rupestre Hook.f., Dracophyllum sprengelioides, Orites milliganii Meissner and Exocarpos humifusus R.Br. In woodland, it is associated with Dracophyllum pandanifolium, Cyathodes glauca Labill., Trochocarpa thymifolia (R.Br.) Sprengel, Eucalyptus urnigera Hook.f., E. coccifera Hook.f. and Telopea truncata (Labill.) R.Br. Phenology Flowering from January to March. Etymology Refers to the leaves that stay on the plant for a long time before dropping. Diagnostic features and notes Dense compact bushes, leaf scars rarely visible, leaves persistent along the branches for several years, lamina apices rigid and sharp. Populations with very large leaves and flowers (measurements in parentheses in the general description) and a much looser habit occur at Pulpit Rock and Collins Bonnet on the Wellington Range. Dracophyllum persistentifolium is a variable species. A chemical study showed that D. persistentifolium separate into chemo forms that broadly corresponded with the underlying geology. The snow skink (Niveoscincus microlepidotus) plays an important 184 Australian Systematic Botany S. Venter A B C D E F Fig. 170. Dracophyllum persistentifolium. A. Inflorescence and leaves. B. Inflorescence bracts. C. Operculum. D. Operculum removed to show flower. E. Flowering branch of inflorescence. F. Sepals. Del. Y. Menadue. role in the pollination of D. persistentifolium. The skinks remove the operculae from the flowers to reach the rich nectar (Fig. 169F). In doing this, they expose the stamens to insect pollinators (Olsson et al. 2000; Johnson et al. 2012). Selected specimens TASMANIA: 1 km north of Reservoir Lakes, 12 Jan. 1984, Adams 21 (HO); south-coast track, summit of Ironbound Range, 22 Apr. 1984, Buchanan 3407 (HO); near summit Mount Barrow, 9 Jan. 1949, Burbidge 3023 (HO); labyrinth track above Cephissus Creek, Cradle Mount National Park, 25 Feb. 1980, Bushby 127 (HO); Procyon Peak, western Arthur Range, 6 Dec. 1986, Collier 2007 (HO); south ridge of Saint Valentines Peak, 13 Jan. 1986, Collier 1169 (HO); Wentworth Hills, Lake King William, 27 Nov. 1988, Collier 3858 (HO); Lake Dobson, Mount Field National Park, Mar. 1978, Crowden s.n. (HO 126532); Mount Wellington, 1 Mar. 1839, Gunn s.n. (BM 35639); Mount Wellington, 20 Nov. 1839, Gunn s.n. (HO 5681 and BM 35638); near Middlesex Plains, 19 Jan. 1842, Gunn s.n. (K); Eagle Tarn, Mount Field National Park, 7 Jan. 1980, Menadue s.n. (HO 125966); Projection Bluff, 16 Dec. 1989, Menadue s.n. (HO 126529); below summit of Projection Bluff, 16 Dec. 1989, Menadue s.n. (HO 126530). Dracophyllum continentis (B.L.Burtt) S.Venter, comb. nov. Richea continentis B.L.Burtt, Curtis’s Bot. Mag. Lond., 163: t. 9632 (1941) Taxonomic revision of Dracophyllum and Richea A Australian Systematic Botany 185 B C D E F Fig. 171. Dracophyllum persistentifolium. A. Habitat at Ben Lamond. B. Young inflorescence showing the prominent inflorescence bracts flowering plant. C. Old plant full in flower near Lady Lake. D. Fruiting branches from near Lake Osborne. E. Inflorescences before dropping of the operculum. F. Snow Skink (Niveoscincus microlepidotus) removing operculae to get at the nectar. Photos: James Wood (A), David Marrison (B), David W. Noble (C, E), Russell Cumming (D) and Tim Rudman (F). Type: Australia. Mount Hotham, F. von Mueller s.n. (lecto: K!), designated by Menadue and Crowden (2000). Richea gunnii sens. Ewart, Flora Vict.: 919 (1931) [non Hook.f. (1847)]. Illustration Y. Menadue and R. Crowden, Austral. Syst. Bot. 13(5): 70 (2000). Erect, multi-branched shrub, 50–100 cm tall. Branches: bark on old branches rough, young stems with leaf scars immediately below leaves. Leaves crowded near tops of branches, leaves persist and decay on the branches; erect–spreading, slightly recurved, rigid; lamina sheath weakly developed, wedge shaped; lamina coriaceous, ovate–triangular, 20–40
4–7 mm; surfaces glabrous; margins scabrous; apex acute, tapering to short, pungent point. Inflorescence a spike-like panicle on main and lateral 186 Australian Systematic Botany Fig. 172. S. Venter Known distribution of Dracophyllum persistentifolium in Tasmania. branches, maturing acropetally, much longer than leaves, erect, lax, 50–200(–300)
2–3 mm, linear–oblong; rachis and pedicels sparsely pilose; basal inflorescence branch up to 6 mm long; inflorescence bracts caducous, overtopping the flowers, whitish to cream at the base with rest pinkish-tipped green turning brown, tapering to a rigid point, 7–30 mm long, glabrous, margins entire. Flowers 30–70+, in groups of 6–10 at the base of the inflorescence, pedicellate; bracteoles 2–3, caducous, shorter than the flower, narrowly linear, 2–2.5 mm long, glabrous; pedicels 0.5–0.7 mm long, sparsely pilose. Sepals greenish-white to cream-coloured, broadly ovate, 2–3 mm long and wide; shorter than the operculum, surfaces glabrous; margins entire. Corolla greenish- or creamy-white; operculum obovoid–conical, 5–8
4–5 mm; apex obtuse; corolla lobes lacking; glabrous. Stamens inserted at the base of the operculum, filaments cream-coloured, 2–3 mm long, articulated near the base; anthers rectangular, cream-coloured, 1 mm long, versatile. Ovary depressed-globose, 1–1.2
1–1.5 mm, glabrous, apex round; nectary scales less than 0.4–0.5
0.3–0.4 mm, apices round; style less than 1 mm long; stigma rounded, reaching base of anthers. Fruit a loculicidal capsule, brown, depressedglobose, 2.5–4.5 mm in diameter, and apex round (Fig. 173, 174). Distribution and ecology Endemic to the Alpine regions of southern New South Wales and Victoria, including the Baw Baw Range (Fig. 175). Common in alpine and subalpine bogs, and the adjacent grasslands, sedgelands and open heaths, especially near flowing water. Commonly associated with Empodisma minus (Hook.f.) L.Johnson & Cutler, Epacris paludosa R.Br., E. celata R.K.Crowden and Baeckea gunniana Schauer. Phenology December to February. Etymology The specific name refers to the occurrence of the taxon on the Australian continent. Taxonomic revision of Dracophyllum and Richea A B Australian Systematic Botany C 187 north-east of Mount Nunniong, 24 Jan. 1984, Parkes no. EG 141a (MEL); Mount Buffalo National Park, 26 Jan. 1982, Short 1374 (HO); Mount Erica, 1892, Tisdale & French s.n. (MEL); Howitt Plain, 6 Jan. 1981, Walsh 2549 (MEL). Dracophyllum gunnii (Hook.f.) S. Venter, comb. nov. Richea gunnii Hook.f., London J. Bot. 6: 273 (1847). D E Type: Tasmania, ‘Hab. Mount Wellington and Western Mountains’, 1837, R.C. Gunn s.n., 1837 (lecto: K!), designated by Menadue and Crowden (2000). Illustration F G Fig. 173. Dracophyllum continentis. A. Flowering branch. B. Operculum. C. Leaf. D. Operculum removed to show flower. E. Operculum and sepals removed to show the nectary scales. F. Flower and bracteole. G. Inflorescence bract. Del. Y. Menadue. Diagnostic features and notes Multi-branched shrub 50–100 cm tall; leaves rigid, ovate–triangular with a pungent apex; inflorescence with axis and lateral floral branches sparsely pilose, maturing acropetally; floral branches up to 6 mm long with 6–10 flowers; flowers on pedicels less than 1 mm long; sepals broadly ovate, 2–3 mm long and wide with the apex obtuse; ovary depressed-globose, 1–1.5 mm in diameter, glabrous; nectary scales less than 0.5 mm long and rounded. Selected specimens NEW SOUTH WALES: Perisher Gap, Kosciusko National Park, 27 Jan. 1984, Crowden s.n. (HO 127573); summit of track near Rams Head, Kosciusko National Park, 11 Feb. 1986, Crowden s.n. (HO 127575); sources of Mowamba River towards Drift Hill, 5–6 Mar 1834, Lhotsky s.n. (MEL); Kosciusko, 26 Jan. 1930, Tillyard s.n. (HO 5711). VICTORIA: heathy spur, Bogong High Plains, 18 Apr 1987, Collier 2380 (HO); summit of Mount Baw Baw, 8 Feb. 1989, Crowden s.n. (HO 127574); Watchbed Creek near gate on Mount Nelse Road, 13 Jan. 1982, Forbes 792, Adair & Gray (MEL); summit of Mount Baw Baw, 24 Jan. 1988, Menadue & Crowden s.n. (HO 308467); Munyang Mountains, 6000 feet [~1828 m] Jan. 1855, Mueller s.n. (MEL); Forlorn Hope Plain, north- Y. Menadue and R. Crowden, Austral. Syst. Bot. 13(5): 792 (2000). Erect shrub, 30–100 cm tall. Branches divaricate, sometimes with many branches originating from the same point. Bark on older stems grey, smooth, young branches with prominent leaf scars. Leaves clustered in the top 5–20 cm of branches; imbricate, spreading and usually recurved, rigid; lamina sheath weakly developed, oblong, margins entire; lamina coriaceous, lanceolate, 30–60
5–7 mm, surfaces glabrous; margins scabrous; apex acute, pungent. Inflorescence a terminal, erect, spike-like panicle, maturing acropetally; longer than leaves, dense, 30–100
10–15 mm, rachis and pedicels pilose; basal inflorescence branch 2–5 mm long, pilose; inflorescence bracts caducous, overtopping the flowers, greenish-pink turning brown, with broad bulbous base up to 12 mm wide, surfaces glabrous, margins entire, apex acute. Flowers 10–40, in groups of 3–5 at the base of the inflorescence, pedicellate; bracteoles 2–3, caducous, shorter than flowers, 3 mm long, narrowkeeled; pedicels 1 mm long, pilose. Sepals creamy-white, broadly ovate to orbicular, 2 mm long and wide; shorter than the operculum, surfaces glabrous, margins entire; apex obtuse. Corolla white, often tinged pink near apex; operculum conical–ovoid, 3–5
2–3 mm, blunt at apex; corolla lobes weakly developed to lacking. Stamens inserted at the base of the operculum; filaments white to pink, seldom dark red, 2–3 mm long, articulated near the base; anthers 0.5–1 mm long, attached to the filament in the top half. Ovary globose, 1–1.5
1–1.5 mm long and wide, glabrous; apex round; nectary scales broad, 0.4–0.5 mm long, emarginate, alternating with the filaments, 1/3 ovary height; style 1.5–2 mm long, reaching 2/3 length of filaments; stigma indistinct. Fruit a loculicidal capsule, depressed-globose, 2.5 mm in diameter (Fig. 176, 177). Distribution and ecology Endemic to Tasmania, mainly on the Central Plateau and Mount Field, also the mountains in the north-east (Fig. 178). Dracophyllum gunnii occurs in montane and subalpine regions in localised wet habitats. It is commonly associated with Leptospermum lanigerum (Aiton) Smith, Epacris gunnii Hook.f., Dracophyllum acerosum, Empodisma minus (Hook.f.) L.Johnson & Cutler, Restio 188 Australian Systematic Botany S. Venter A B C E D G F Fig. 174. Dracophyllum continentis. A. Habitat at Mount Kosciuszko National Park. B. Inflorescence with some operculae still attached. C. Flowering plant showing the spreading habit. D. Flowers showing the short stamens. E. Fruiting branch. F. Fruit. G. Young inflorescence showing the small inflorescence bracts. Photos: H. Pauli (A, C, D), M. Bayly (B), R. Cumming (E, F) and M. Cosgrove (G). australis R.Br., sphagnum moss and various cushion-plant species. Phenology Flowering from December to February. Etymology Named for Ronald Campbell Gunn (1808–1881), Deputy Commissioner for Crown Lands of Tasmania and botanist, who made extensive collections in Tasmania. Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 189 Fig. 175. Known distribution of Dracophyllum continentis in New South Wales and Victoria, Australia. Diagnostic features and notes Type: Tasmania, February 1841, R.C. Gunn 1215. (lecto: K!; isolecto: K!), designated by Menadue and Crowden (2000). Open divaricate shrub with recurved, 30–60-mm-long leaves; lower inflorescence branches 2–5 mm long, bearing flowers in groups of three to five; sepals creamy-white and broadly ovate to orbicular; operculum 3–5 mm long; stamens white to pink. Richea pandanifolia subsp. ramulosa Menadue, Austral. Syst. Bot. 13 (5): 795–796 (2000). Selected specimens Type: Tasmania, Newell Creek, 10 km south-south-west of Queenstown 300 m across the King River, 12 Dec. 1997, Y. Menadue & R.K. Crowden s.n. (holo: HO 328142!; iso: CANB!, MEL!, NSW!). TASMANIA: headwaters of Mountain River, Mount Wellington, 13 Jan. 1981, Brown 110 (HO); near Lake Rodway, Cradle Mountain, 30 Jan. 1982, Buchanan 864 (HO); Pine Lake, 17 Feb. 1985, Collier 377 (HO); Clumner Bluff, 20 Apr. 1985, Collier 424 (HO); Paradise Plain, Newitts Creek, 11 May 1985, Collier 500 (HO); Ben Nevis, 15 Mar. 1986, Collier 1270 (HO); Mount Wellington, 29 Jan. 1841, Gunn s.n. (BM 35645); Mount Micheal, Sun Creek, ~8 km north-west of Goulds Country, 17 Apr. 1979, Jackson s.n. (HO 29958); Fehres Marsh, 24 Feb. 1982, Moscal 917 (HO); Mother Lords Plain, Breton Rivulet, 2 Feb. 1981, Moscal 542 (HO); Devils Den, 18 Mar. 1984, Moscal 7140 (HO); Mount Barrow near transmitter, 21 Apr. 1980, McKendrick 17 (HO); Ben Lomond National Park, below Wilmot’s Bluff, 27 Feb. 1979, Noble 28362 (HO); Mount Styx, Oct. 1910, Rodway s.n. (HO 5616); Blue Tier, Great Lake, 1891, Simpson s.n. (HO 5618); Newdegate Pass, Mount Field National Park, 9 Jan. 1978, Smith 345 (HO); start of Mount Rufus track, 1 Dec. 1962, Somerville s.n. (HO 5619). Dracophyllum pandanifolium (Hook.f.) S.Venter, comb. nov. Richea pandanifolia Hook.f., Fl. Antarctica 1: 50 (1844). Illustration Y. Menadue and R. Crowden, Austral. Syst. Bot. 13: 794 (2000). Tree 2–12 m tall. Branches usually with a single, slender trunk or sparsely branched. Bark rough on trunks and old branches, leaf scars rarely apparent and only where leaves have just fallen. Leaves crowded along branches, with dead leaves persisting down most or the entire trunk, or numerous branches with leaves persisting and confined to the ends of the branches. Lamina sheath not well developed, transversely oblong; lamina rigid, coriaceous, prominently striated, and drooping, 700–1500
60–80 mm wide above the base; margin cartilaginous with small distant teeth; apex subulate, tapering to a long, twisting point. Inflorescence a many-branched axillary panicle, much shorter than leaves, erect–spreading, dense, maturing acropetally, 150–250
30–80 mm, more or 190 Australian Systematic Botany S. Venter A B C D E F G Fig. 176. Dracophyllum gunnii. A. Inflorescence. B. Inflorescence bracts. C. Operculum. D. Flower. E. Operculum removed to show stamens and ovary. F. Branching habit. G. Leaf. Del. Y. Menadue. less hidden by the leaves; rachis and pedicels glabrous; basal inflorescence branch 30–40 mm long, spreading; inflorescence bracts caducous, overtopping flowers, lanceolate at the base, 50–170 mm long, surfaces glabrous, margins entire. Flowers 300–500+, in groups of 16–20 at the base of the inflorescence, pedicellate; bracteoles caducous, narrow, 2 mm long, glabrous; pedicels straight, 0.8–2.5 mm long, glabrous. Sepals greenish-white, pink or reddish, broadly ovate, 0.5–1.4
0.8–1 mm, shorter than operculum, glabrous, margins entire, apex bluntly acute to nearly truncate. Corolla greenish-white, pink or reddish; operculum cylindrical–conical with flattened apex, 2.5–3.5
2–3 mm; corolla lobes weakly developed, up to 0.5 mm long. Stamens inserted at the base of the operculum, filaments greenish-white, pink or reddish; 2–2.5 mm long; anthers rectangular, cream-coloured, ~1 mm long, versatile. Ovary depressed-globose, 0.9–1
1.4–1.5 mm, glabrous; apex round; nectary scales rectangular to depressedrectangular, apices truncate or emarginate, 0.5–0.6
0.6–0.7 mm, just visible above or enclosed by sepals; style 1–1.4 mm long, cylindrical; stigma rounded, reaching centre or just above top of anthers. Fruit pedicellate, included in persistent calyx, transversely oblong, 1.5–1.7
2–2.5 mm, deep reddish-brown (Fig. 179, 180). Distribution and ecology Dracophyllum pandanifolium is endemic to the western, southwestern areas of Tasmania, with extensive populations in the Mount Field, occurring almost exclusively on pre-Cambrian bedrocks in high-rainfall areas (Fig. 181). Dracophyllum Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A C 191 B D Fig. 177. Dracophyllum gunnii. A. Habitat at Lake Kay. B. Fruiting branch showing the persistent styles. C. Flowering plant showing the leaves characteristically at the tips of the branches. D. Inflorescence with some shed operculae showing the widely spreading stamens. Photos: N. Tapson (A, B, D) and T. Rudman (C). pandanifolium grows in subalpine moorland and shrubbery in moist but well drained sites, often in gullies with an open eucalypt canopy or a rainforest overstorey. Stunted forms occur on ridges and mountaintops and branched forms occur where there has been terminal-bud damage. Phenology Flowering from late November to January. Etymology Describes the leaf shape, resembling that of a Pandanus. Diagnostic features and notes This is a very distinct plant and an important element of the Tasmanian flora. Usually with a single trunk covered in a skirt of old dry leaves; leaves long and narrow (70–150
6–8 cm), drooping and coriaceous with the apices sometimes spiralling; inflorescence a short axillary panicle and mostly hidden by the leaves, inflorescence bracts shed early, 16–20 flowers on lower inflorescence branches, sepals broad, operculae whitish to red. There is a branching form described as Richea pandanifolia subsp. ramulosa (Menadue and Crowden 2000). This form of branching variation also occurs in D. fiordense. In some areas, such as, for example, lower Gordon River, the two subspecies 192 Australian Systematic Botany S. Venter Fig. 178. Known distribution of Dracophyllum gunnii in Tasmania. co-exist and there is also overlap in morphological characters, they also share the same flavonoids. Both non-branching and branching forms occur together in the Serpentine Gorge valley (S. Venter, pers. obs.). Birds were recorded visiting the flowers of D. pandanifolium (Johnson et al. 2012). Selected specimens TASMANIA: Moore’s Bridge, 15 Jan. 1984, Adams 54 (HO); Hartz Mountain Road, 27 Nov. 1975, Allan & Stones s.n. (HO 65783); Moonlight Ridge, 22 Mar. 1984, Buchanan 3064 (HO); headwaters of Swift Creek, 8 Jan. 1984, Buchanan 2285 (HO); Pigsty Ponds, 21 Mar. 1984, Buchanan 3000 (HO); Jubilee Range, 13 Jan. 1985, Buchanan 5283 (HO); Mount Rufus, 25 Jan. 1949, Burbidge 3335 (HO); southern ridge of Mount Dundas, 3 Jan. 1987, Collier 2145 (HO); road to Lake Dobson, 22 Feb. 1958, Court 1223 (HO); Scott’s Peak Rd near nature walk, 22 Feb. 1990, Crowden & Menadue s.n. (HO 126527); Frenchman’s Cap, Feb. 1841, Gunn s.n. (BM 35646 and K); Van Diemen’s Land, Gunn s.n. (K); K-Col, Mount Field National Park, 12 Dec. 1952, Melville 2334 (HO); Mount Sedgewick, north-east of Queenstown, 17 Nov. 1990, Menadue.& Crowden s.n. (HO 408001); Mount Black, near Rosebery, 26 Feb. 1930, Nye 1 (HO); Hartz Mountains, Dec. 1894, Rodway s.n. (HO 5641); north end Eagle Tarn, Mount Field National Park, 24 Jan. 1983, Short 1830 (HO); Lake Fenton, Mount Field National Park, 21 Jan. 1944, Somerville s.n. (HO 65784); Frankland River, 1 km north of Balfour, 11 Dec. 1983, Moscal 4767 (HO); below Gordon River Dam, south-western Tasmania, 1 Feb. 1996, Menadue & Crowden s.n. (HO 330139). Dracophyllum desgrazii (Hombr. ex Decne.) S.Venter, comb. nov. [non ‘Dracophyllum dracophyllum’; tautonym] Richea desgrazii Hombr. ex Decne., Voy. Pôle Sud 2: 85 (1853), as ‘desgrasii’. Taxonomic revision of Dracophyllum and Richea C B A D E F Australian Systematic Botany 193 turning brown with age, 50–200
25–35 mm; surfaces glabrous, margins serrulate. Flowers 300+, in groups of up to 40 at the base of the inflorescence, pedicellate; bracteoles 3, caducous, Shorter than the flower, 8–10 mm long, narrow, membranous; pedicels straight, 1.5–6 mm long, glabrous. Sepals lime green to white, broadly triangular, 1–1.5 mm long, shorter than the operculum, glabrous, margins entire. Corolla white; operculum obovoid–conical, 8–10
2–4 mm, flattened dorsiventrally; apex bluntly pointed, corolla lobes sometimes developed, minute. Stamens inserted at the base of the operculum; filaments creamy-white; 5–6 mm long, articulated shortly above their insertion; anthers bilobed at the apex, 1.5 mm long, versatile. Ovary dark red, globose, ~0.9–1
1.9–2 mm, glabrous, apex round; nectary scales ellipticaltruncate, 1–1.2 mm long and wide, apices truncate; style tapering, 2–2.5 mm long; stigma rounded, reaching 1/2 length of stamens. Fruit pedicellate, included in persistent calyx, at first crimson but later brown, depressed-globose, 3 mm in diameter, style persistent (Fig. 182, 183). Distribution and ecology Endemic to south-eastern Tasmania, including Maria and Bruny islands (Fig. 184). Occurs in wetter areas of sclerophyll forests under a eucalypt canopy (Menadue and Crowden 2000). Fig. 179. Dracophyllum pandanifolium. A. Plant showing growth habit. B. Part of the inflorescence. C. Operculum. D. Sepals. E. Operculum removed to show flower. F. Sepals and operculum removed to show the nectary scales. Del. Y. Menadue. Type: J. B. Hombron in J. Decaisne, Voy. Pôle Sud 2: plate 29A (1853) (holo!) [plate is labelled as ‘27’ in error. The description was prepared using the illustration, as no specimens of this species were received at the Muséum National d’Histoire Naturelle in Paris.] Richea dracophylla R.Br. Prodr. 555 (1810). Type: Tasmania, Table Mountain, (Mount Wellington), R. Brown s.n. (holo: K000349908!). Illustration Y. Menadue and R. Crowden, Austral. Syst. Bot. 13(5): 798 (2000). Erect, sparsely branched shrub 1.5–5 m tall. Branches sparsely branched. Bark on older branches greyish-brown, young stems yellowish to reddish-brown with prominent leaf scars. Leaves crowded near ends of branches, spreading, flexuous; Lamina sheath pinkish-green to red, coriaceous, striate, and broadly oblong with the margins membranous. Lamina coriaceous, lanceolate, 150–330
13–22 mm; prominently striated, margin with small distinct teeth; apex subulate; Inflorescence a terminal panicle, longer than leaves, erect, dense, maturing acropetally; 120–250
30–45 mm, densely branched; rachis and pedicels pubescent; basal inflorescence branch up to 12 mm long, spreading; inflorescence bracts caducous, sometimes persistent, overtopping flowers, pink to pinkish-red with green apex, Phenology Flowering from September to October. Etymology The specific epithet refers to the dark red ovary. Diagnostic features and notes Leaves long (150–330 mm) and flexuous; inflorescence axis pubescent, inflorescence bracts large (5–20 cm long) pink with green apices turning brown, flowers in clusters of 8–12 on the lower inflorescence branches with 1.5–6.0 mm long pedicels, operculum 8–10 mm long and dorsiventrally flattened and the ovary dark red. Birds were recorded visiting the flowers of D. desgrazii (K. A. Johnson, unpubl. data). Selected specimens TASMANIA: Mount Wellington near Hobart, Backhouse s.n. (HO); Collins Cap Trail, 1980, Brown 44 (HO); Bishop and Clerk, Maria Island, 1977, Brown 206 (HO); Lockleys Road, above head of Captain Cook Creek, 15 Dec. 1984, Buchanan 3924 (HO); cliff at Perdition Ponds, Cape Pillar, 15 Apr. 1984, Buchanan 3297 (HO); Myrtle Forest Creek, 7 Apr. 1984, Buchanan 3224 (HO); Mount Maria, Maria Island, 10 Oct. 1986, Collier 1729 (HO); Catamaran, 1831, Gunn s.n. (HO 5603); Mount Wellington, 30 Oct. 1840, Gunn s.n. (HO 5602 and BM 35643), 5 Sep. 1840 (BM 35644), 20 Nov. 1840 (BM 35640), 29 Nov. 1839 (BM 35642); Mount Wellington, 30 Oct. 1840, Gunn s.n. (K); Mount Wellington, Dec. 1870, Hannaford s.n. (HO5593); Picton River, 15 km west of Geeveston, 9 Dec. 1967, Hemsley s.n. (HO 5592); Tahune Bridge, Huon River, 2 Sep. 1979, Jarman 15 (HO); Mount Wellington, 26 Nov. 1931, Long 659 (HO); Mount Mangana, Bruny Island, Feb. 1979, Menadue s.n. (HO 126004); MacGregor Peak, Forestier Pen, 31 Mar. 1985, Moscal 10424 (HO); Mount Wellington, Sep. 1892, Rodway s.n. (HO 5597). 194 Australian Systematic Botany S. Venter A C B D Fig. 180. Dracophyllum pandanifolium. A. Habitat at Lake Dobson. B. Mature plants showing the skirt of dry leaves. C. Inflorescences with most of the operculae and all the inflorescence bracts shed. D. Inflorescence with old and dry inflorescence bracts and shed operculae on the leaf. Photos: John Harrison (A), Russell Cumming (B, D) and Natalie Tapson (C). Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany 195 Fig. 181. Known distribution of Dracophyllum pandanifolium in Tasmania. Dracophyllum alpinum (Menadue) S.Venter, comb. nov. Richea alpina Menadue, Austral. Syst. Bot. 13(5): 798–801 (2000). Type: Tasmania, Mount Sprent, in shoulder just below summit at 900 m, 21 Feb. 1990, Y. Menadue & R.K. Crowden s.n. (holo: HO 128083!). Illustration Y. Menadue and R. Crowden, Austral. Syst. Bot. 13(5): 799 (2000). Low growing shrub, 20–80(–100) cm tall. Branches sparsely branched, branches often covered by soil. Bark on old branches greyish-brown, young stems with prominent annular scars. Leaves crowded towards ends of branches, recurved in a bromelioid manner, spreading; lamina sheath weakly developed, broadly oblong; lamina thickly coriaceous, narrow-lanceolate to lanceolate, 80–130
6–12 mm, surfaces glabrous, striated, margins minutely and sharply serrate; apices acute. Inflorescence a terminal, erect panicle, longer than the leaves, dense, 60–200
50–140 mm, pyramidal and densely branched, maturing acropetally; rachis and pedicels sparsely pubescent; basal inflorescence branch widely spreading; 196 Australian Systematic Botany A S. Venter B C plants grow taller (70–100 cm high), as they are more protected, and sparse branching may be seen (Menadue and Crowden 2000). Phenology From November to late January. Etymology Describes the alpine habitat. E D Diagnostic features and notes Lamina margins sharply serrate, inflorescence a pyramidal densely branched panicle maturing acropetally, floral branches sparsely pubescent, flowers in groups of 10–40 at the base of the inflorescence, lower bracts similar to leaves but smaller, sepals broadly triangular, nectary scales broadly elliptic–oblong and truncate. Selected specimens Fig. 182. Dracophyllum desgrazii. A. Flowering branch. B. Flower. C. Inflorescence bract. D. Floral parts removed to show ovary and nectary scales. E. Flower clusters on the lower inflorescence branches. Del. Y. Menadue. inflorescence bracts caducous, overtopping the flowers, greenish-brown turning brown before falling, surface glabrous, margins serrulate. Flowers 500–1000+, in groups of 10–40 flowers at the base of the inflorescence, pedicellate; bracteoles caducous, 2–3–(4), shorter than flower, 3–4 mm long, linear, margin ciliate; pedicels straight, 2–4 mm long; sparsely pubescent. Sepals same colour as corolla, broadly triangular, 1–3 mm long, obtuse, shorter than operculum, surfaces glabrous; margins entire. Corolla pink, orange, or crimson; operculum narrowly obovoid to cylindrical–conical, occasionally flattened, 6–8
3–3.5 mm; apex obtuse with 5 obscure teeth. Stamens inserted at the base of the operculum, filaments 3–4.5 mm long, attached to anthers slightly below the centre; anthers oblong and 1–1.5 mm long. Ovary depressed-globose, 1.6–1.8
2–3 mm, glabrous, apex round; nectary scales broadly elliptic–oblong, 0.6–0.8
0.7–0.9 mm, truncate; style 1–3 mm long; stigma rounded, reaching base of anthers. Fruit pedicellate, depressed-globose, 2–2.5
2.5–3.5 mm (Fig. 185, 186). Distribution and ecology Endemic to Tasmania, on mountains of the west and southwest (Fig. 187). Dracophyllum alpinum occurs in exposed alpine moor and low shrubbery, often forming solitary plants or low copses, with branches often covered over, so that the above-ground parts appear separate and rosette-like (20–30 cm tall). Also, in subalpine shrubbery where the TASMANIA: north ridge of Mount Sprent, 23 Feb. 1985, Collier 383 (HO); Lake Cygnus, Western Arthur Range, 7 Dec. 1986, Collier 2014 (HO); Mount Bobs, ridge from Farmhouse Creek, 4 Jan. 1983, Gillanders s.n. (HO 69723); Lake Fortuna, Western Arthurs, 16 Dec. 1990, Jordon s.n. (HO 410564); Mount Sprent just below summit, 21 Feb. 1990, Menadue & Crowden s.n. (HO 128083 and HO 128084); Mount Bobs summit, 29 Jan. 1984, Williams s.n. (HO 7296); Norold Range, 15 Nov. 1985, Ziegeler s.n. (HO 97427). Conflicts of interest The author declares that he has no conflicts of interest. Declaration of funding Financial support provided by Victoria University of Wellington. This study forms part of Stephanus Venter’s PhD Thesis, ‘A taxonomic revision of the genus Dracophyllum Labill. (Ericaceae)’ Acknowledgements I thank Phil Garnock-Jones (School for Biological Sciences, Victoria University of Wellington), Ilse Breitwieser, Aaron Wilton, Peter Heenan, Mary Korver and Steve Wagstaff (Landcare Research, Lincoln). The directors, curators and their staff of the following Herbaria for material on loan and for handling various requests: AK, BM, CHR, FI, K, L, MEL, O, P, S, W, WELT, WELTU, Z. John Barkla, Shannel Courtney and Cathy Jones (DoC) for organising the collecting permit and for collecting specimens in isolated areas. John Dugdale (Landcare Research, Nelson) for identification of insect specimens and valuable discussions on pollination. Audrey Eagle for hospitality during fieldwork. Steve Wagstaff, Landcare Research, Lincoln for valuable discussions on cladistic and phylogenetic aspects. Aaron Wilton and his wife Vicky for help with fieldwork and, very valuable discussions, and also acting as my hosts. The following people gave permission for their photos to be used: Steve Attwood, S. Aubert and R. Douzet from the University of Grenoble, Kirsten Baker, Phil Bendle, Russell Best, Dianna Bradshaw, Disa Cragg-Ohlsson, Alan Cressler, Rhys L. Davies, Grant Dixon, Robert Hamilton, Mike Harré, John Harrison, David Marrison, David W. Noble, Harold Pauli from Vienna, Tim Rudman, Natalie Tapson, David Tng, Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany A B C D E E Fig. 183. Dracophyllum desgrazii. A. Plants in habitat, Mount Maria. B. Adult infructescence with the old inflorescence bracts still intact, Hobart. C. Inflorescence showing flowers with and without operculae, Mount Wellington. D. Plant in fruit. E. Mature plant in full flower. F. 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Isles 4 Suppl. 2: 620 (1917) Dracophyllum drummondii Benth., Fl. Austral. 4: 263 (1868). Dracophyllum gracile R.Br., Prodr. Fl. Nov. Holland.: 556 (1810) Dracophyllum parviflorum Benth., Fl. Austral. 4: 265 (1868) Dracophyllum phlogiflorum Benth., Fl. Austral. 4: 263 (1868). Dracophyllum squarrosum R.Br., Prodr. Fl. Nov. Holland: 556 (1810) Oreothamnus Baum.-Bod., Syst. Fl. Neu–Caledonien 5: 76 (1989) Oreothamnus amabilis Baum.-Bod., Syst. Fl. Neu-Caledonien 5: 76 (1989) Oreothamnus cosmelioides Baum.-Bod., Syst. Fl. Neu-Caledonien 5: 6 (1989) Oreothamnus ramosus Baum.-Bod., Syst. Fl. Neu-Caledonien 4: 102 (1989) and 5: 76, 97 (1989) Richea africana (Benth.) Kuntze, Revis. Gen. Pl. 1: 235 (1891) Richea afzelii (Oliv.) Kuntze, Revis. Gen. Pl. 1: 235 (1891), as ‘afzelia’ Richea glauca Labill., Voy. a Rec. Pérouse (1800) Richea gummiflua (Tul.) Baill., Hist. Pl. Madag., Atlas 35, t. 323 (1894) Richea lanceolata (Tul.) Baill., Hist. Pl. Madag., Atlas 35: t. 324 (1894) Richea leptoclada (Tul.) Kuntze, Rev. Gen. Pl. 1: 235 (1891) Richea madagascariensis (DC.) Kuntze, Revis. Gen. Pl. 1: 235 (1891) Richea microphylla (Tul.) Kuntze, Revis. Gen. Pl. 1: 235 (1891) Richea ovata (Tul.) Kuntze, Revis. Gen. Pl. 1: 235 (1891) Richea phaeotricha (Tul.) Kuntze, Revis. Gen. Pl. 1: 235 (1891) Richea plumosa (Oliv.) Kuntze, Revis. Gen. Pl. 1: 235 (1891) Richea ceylanica Kuntze, Revis. Gen. Pl. 1: 235 (1891) nom. inval., published without a replaced synonym reference and the author of Dracophyllum is not stated. The generic name is not validly published nom. inval., generic name not validly published Sphenotoma capitata (R. Br.) Lindl. (as ‘capitatum’) Sphenotoma dracophylloides Behr & F.Muell. ex Sond. Sphenotoma drummondii (Benth.) F.Muell. Epacris gracilis (R.Br.) Spreng Sphenotoma parviflora (Benth.) F.Muell. (as ‘parviflorum’) =Sphenotoma dracophylloides Behr & F.Muell. ex Sond. Sphenotoma squarrosa (R.Br.) G.Don nom. inval., without a Latin description or diagnosis or type nom. inval., generic name not validly published nom. inval., generic name not validly published nom. inval., generic name not validly published =Cassipourea congoensis R.Br. ex DC. Cassipourea afzelii (Oliv.) Alston Craspedia glauca (Labill.) Spreng. Cassipourea gummiflua Tul. Cassipourea lanceolata Tul. Cassipourea leptoclada Tul. Cassipourea madagascariensis (Thouars) DC. =Cassipourea microphylla Tul. Cassipourea ovata Tul. Cassipourea phaeotricha Tul. Cassipourea plumosa (Oliv.) Alston as ‘zeylanica’ Cassipourea ceylanica (Gardner) Alston Taxonomic revision of Dracophyllum and Richea Australian Systematic Botany Appendix 2. Subgenus Dracophyllum (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (11) (12) (13) (14) (15) (16) (17) (18) (19) (20) (21) Subgenus Oreothamnus (1) (2) (3) (4) (5) (6) (7) (8) (9) (10) (11) (12) (13) (14) (15) (16) (17) (18) (19) (20) (21) (22) (23) (24) (25) (26) (27) (28) (29) Subgenus Cystanthe (1) (2) (3) (4) Subgenus Richea (1) (2) (3) (4) (5) (6) (7) Numerical list of Dracophyllum taxa Dracophyllum alticola Däniker Dracophyllum balansae Virot Dracophyllum cosmelioides Panch. ex W.R.B.Oliv. Dracophyllum elegantissimum S.Venter Dracophyllum fiordense W.R.B.Oliv. Dracophyllum fitzgeraldii C.Moore & F.Muell. Dracophyllum involucratum Brongn. & Gris Dracophyllum latifolium A.Cunn. Dracophyllum mackeeanum S.Venter Dracophyllum macranthum E.A.Br. & Streiber Dracophyllum menziesii Hook.f. Dracophyllum milliganii Hook. Dracophyllum oceanicum E.A.Br. & Streiber Dracophyllum ouaiemense Virot Dracophyllum ramosum Panch. ex Brongn. & Gris Dracophyllum sayeri F.Muell. Dracophyllum secundum R.Br. Dracophyllum strictum Hook.f. Dracophyllum townsonii Cheeseman Dracophyllum traversii Hook.f. Dracophyllum verticillatum Labill. Dracophyllum acerosum Berggr. Dracophyllum arboreum Cockayne Dracophyllum cockayneanum Du Rietz Dracophyllum densum W.R.B.Oliv. Dracophyllum filifolium Hook.f. Dracophyllum frondosum (G.Simpson) S.Venter Dracophyllum kirkii Berggr. Dracophyllum lessonianum A.Rich. Dracophyllum longifolium (J.R.Forst. & G.Forst.) R.Br. ex Roem. & Schult. Dracophyllum marmoricola S.Venter Dracophyllum minimum F.Muell. Dracophyllum muscoides Hook.f. Dracophyllum oliveri Du Rietz Dracophyllum ophioliticum S.Venter Dracophyllum palustre W.R.B.Oliv. Dracophyllum patens W.R.B.Oliv. Dracophyllum pearsonii Kirk Dracophyllum politum (Cheeseman) Cockayne Dracophyllum pronum W.R.B.Oliv. Dracophyllum prostratum Kirk Dracophyllum pubescens Cheeseman Dracophyllum recurvum Hook.f. Dracophyllum rosmarinifolium (G.Forst.) R.Br. Dracophyllum scoparium Hook.f. Dracophyllum septentrionale (W.R.B.Oliv.) S.Venter Dracophyllum sinclairii Cheeseman Dracophyllum subulatum Hook.f. Dracophyllum trimorphum W.R.B.Oliv. Dracophyllum urvilleanum A.Rich. Dracophyllum tasmanicum S.Venter Dracophyllum laciniatum S.Venter Dracophyllum procerum (F.Muell.) S.Venter Dracophyllum sprengelioides (R.Br.) S.Venter Dracophyllum alpinum (Menadue) S.Venter Dracophyllum desgrazii (Hombr. ex Decne.) S.Venter Dracophyllum continentis (B.L.Burtt) S.Venter Dracophyllum gunnii (Hook.f.) S.Venter Dracophyllum pandanifolium (Hook.f.) S.Venter Dracophyllum persistentifolium S.Venter Dracophyllum victorianum (Menadue) S.Venter www.publish.csiro.au/journals/asb 205