Linder, H.P., Baeza, M., Barker, N.P., Galley, C., Humphreys, A.M., Lloyd, K., Orlovich, D., Pirie, M.D., Simon, B.K., Walsh, N. &amp; Verboom, G.A. (2010). A classification of the Danthonioideae (Poaceae). Annals of the Missouri Botanical Garden 97:306-364.

Nigel Barker

Molecular sequence data from two noncoding regions of DNA (ITS AND TRNL-F) were used to produce a phylogeny for the genus Tribolium and its African and Australasian allies. Topological comparisons of the combined molecular data with the appended existing morphological tree were made. A significant incongruence was revealed. Molecular data indicate that Tribolium is paraphyletic. The formally defined sections: Uniolae, Acutiflorae and Tribolium have been retrieved by the molecular data. The analysis retrieved Karroochloa as polyphyletic. Monophyly of Schismus is strongly supported. The Australasian species form a monophyletic clade. Data support early divergence of the Merxmuellera species and Pseudopentameris macrantha

A GENERIC CLASSIFICATION OF THE DANTHONIOIDEAE (POACEAE)1 H. Peter Linder,2 ,3 Marcelo Baeza,4 Nigel P. Barker,5 Chloé Galley,2 Aelys M. Humphreys,2 Kelvin M. Lloyd,6 David A. Orlovich,7 Michael D. Pirie,2,8 Bryan K. Simon,9 Neville Walsh,10 and G. Anthony Verboom11 A GENERIC CLASSIFICATION OF THE DANTHONIOIDEAE (POACEAE)1 H. Peter Linder,2 ,3 Marcelo Baeza,4 Nigel P. Barker,5 Chloé Galley,2 Aelys M. Humphreys,2 Kelvin M. Lloyd,6 David A. Orlovich,7 Michael D. Pirie,2,8 Bryan K. Simon,9 Neville Walsh,10 and G. Anthony Verboom11 ABSTRACT We present a new generic classification of the largely Southern Hemisphere grass subfamily Danthonioideae. This classification is based on an almost completely sampled and well-resolved molecular phylogeny and on a complete morphological data set. We have attempted to delimit monophyletic genera (complicated by the presence of apparent intergeneric hybridization), which are diagnosable, as well as morphologically and ecogeographically coherent. We recognize 17 genera, including five new genera (Austroderia N. P. Barker & H. P. Linder, Capeochloa H. P. Linder & N. P. Barker, Chimaerochloa H. P. Linder, Geochloa H. P. Linder & N. P. Barker, and Tenaxia N. P. Barker & H. P. Linder), and two sections newly designated for Pentameris P. Beauv. (section Dracomontanum H. P. Linder & Galley and section Pentaschistis (Nees) H. P. Linder & Galley). Of the remaining 12 genera, the delimitations of seven are changed: Merxmuellera Conert is much reduced by the segregation of Geochloa, Capeochloa, and Tenaxia; Pentameris is expanded to include Prionanthium Desv. and Pentaschistis (Nees) Spach; Cortaderia Stapf is expanded by the inclusion of Lamprothyrsus Pilg., but reduced by the segregation of its New Zealand species into the new genus Austroderia; a large Rytidosperma Steud. is assembled out of Joycea H. P. Linder, Austrodanthonia H. P. Linder, Notodanthonia Zotov, Erythranthera Zotov, Pyrrhanthera Zotov, and Monostachya Merr.; and the species previously assigned to Karroochloa Conert & Türpe, Schismus P. Beauv., Urochlaena Nees, and Tribolium Desv. have been reassigned to only two genera. Finally, the Himalayan species of Danthonia DC. are transferred to Tenaxia and the remaining African species of Danthonia to Merxmuellera. The 281 species that we recognize in the subfamily are listed under their new genera, which are arranged in the phylogenetic sequence evident from the molecular phylogeny. The 100 necessary new combinations include: Merxmuellera grandiflora (Hochst. ex A. Rich.) H. P. Linder, Geochloa decora (Nees) N. P. Barker & H. P. Linder, G. lupulina (L. f.) N. P. Barker & H. P. Linder, G. rufa (Nees) N. P. Barker & H. P. Linder, Capeochloa arundinacea (P. J. Bergius) N. P. Barker & H. P. Linder, C. cincta (Nees) N. P. Barker & H. P. Linder, C. cincta subsp. sericea (N. P. Barker) N. P. Barker & H. P. Linder, C. setacea (N. P. Barker) N. P. Barker & H. P. Linder, Pentameris praecox (H. P. Linder) Galley & H. P. Linder, P. tysonii (Stapf) Galley & H. P. Linder, P. acinosa (Stapf) Galley & H. P. Linder, P. airoides Nees subsp. jugorum (Stapf) Galley & H. P. Linder, P. alticola (H. P. Linder) Galley & H. P. Linder, P. ampla (Nees) Galley & H. P. Linder, P. andringitrensis (A. Camus) Galley & H. P. Linder, P. argentea (Stapf) Galley & H. P. Linder, P. aristidoides (Thunb.) Galley & H. P. Linder, P. aristifolia (Schweick.) Galley & H. P. Linder, P. aspera (Thunb.) Galley & H. P. Linder, P. aurea (Steud.) Galley & H. P. Linder, P. aurea subsp. pilosogluma (McClean) Galley & H. P. Linder, P. bachmannii (McClean) Galley & H. P. Linder, P. barbata (Nees) Steud. subsp. orientalis (H. P. Linder) Galley & H. P. Linder, P. basutorum (Stapf) Galley & H. P. Linder, P. borussica (K. Schum.) Galley & H. P. Linder, P. calcicola (H. P. Linder) Galley & H. P. Linder, P. calcicola var. hirsuta (H. P. Linder) Galley & H. P. Linder, P. capensis (Nees) Galley & H. P. Linder, P. capillaris (Thunb.) Galley & H. P. Linder, P. caulescens (H. P. Linder) Galley & H. P. Linder, P. chippindalliae (H. P. Linder) Galley & H. P. Linder, P. chrysurus (K. Schum.) Galley & H. P. Linder, P. clavata (Galley) Galley & H. P. Linder, P. colorata (Steud.) Galley & H. P. Linder, P. dentata (L. f.) Galley & H. P. Linder, P. dolichochaeta (S. M. Phillips) Galley & H. P. Linder, P. ecklonii (Nees) Galley & H. P. Linder, P. exserta (H. P. Linder) Galley & H. P. Linder, P. galpinii (Stapf) Galley & H. P. Linder, P. holciformis (Nees) Galley & H. P. Linder, P. horrida (Galley) Galley & H. P. Linder, P. humbertii (A. Camus) Galley & H. P. Linder, P. insularis (Hemsl.) Galley & H. P. Linder, P. juncifolia (Stapf) Galley & H. P. Linder, P. longipes (Stapf) Galley & H. P. Linder, P. malouinensis (Steud.) Galley & H. P. Linder, P. microphylla (Nees) Galley & H. P. Linder, P. minor (Ballard & C. E. Hubb.) Galley & H. P. Linder, P. montana (H. P. Linder) Galley & H. P. Linder, P. 1 This article is dedicated to Surrey Jacobs (1946–2009) in honor of his lifelong work on and interest in the Australian grasses. This research was supported by the Swiss National Science Foundation (grant 3100AO-10727); fieldwork was supported in part by the Claraz Foundation and in part by the Swiss Academy of Sciences. Collecting permission was granted by the nature conservation authorities in South Africa, Malawi, Australia, New Zealand, and Chile. The illustrations were made by Jasmin Baumann. 2 Institute of Systematic Botany, University of Zurich, Zollikerstrasse 107, CH-8008, Switzerland. 3 Author for correspondence: peter.linder@systbot.uzh.ch. 4 Departamento de Botánica, Universidad de Concepción, Casilla 160-C, Concepción, Chile. 5 Department of Botany, Rhodes University, Grahamstown 6140, South Africa. 6 Landcare Research, Private Bag 1930, Dunedin 9054, New Zealand. 7 Department of Botany, University of Otago, P.O. Box 56, Dunedin 9054, New Zealand. 8 Current address: Department of Biochemistry, University of Stellenbosch, Private Bag X1, Matieland 7602, South Africa. 9 Queensland Herbarium, Toowong, Queensland 4066, Australia. 10 Royal Botanic Gardens Melbourne, Birdwood Avenue, South Yarra, Victoria 3141, Australia. 11 Department of Botany, University of Cape Town, Rondebosch 7700, South Africa. doi: 10.3417/2009006 ANN. MISSOURI BOT. GARD. 97: 306–364. PUBLISHED ON 10 OCTOBER 2010. Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 307 natalensis (Stapf) Galley & H. P. Linder, P. oreodoxa (Schweick.) Galley & H. P. Linder, P. pallida (Thunb.) Galley & H. P. Linder, P. pholiuroides (Stapf) Galley & H. P. Linder, P. pictigluma (Steud.) Galley & H. P. Linder, P. pictigluma var. gracilis (S. M. Phillips) Galley & H. P. Linder, P. pictigluma var. mannii (Stapf ex C. E. Hubb.) Galley & H. P. Linder, P. pseudopallescens (H. P. Linder) Galley & H. P. Linder, P. pungens (H. P. Linder) Galley & H. P. Linder, P. pusilla (Nees) Galley & H. P. Linder, P. pyrophila (H. P. Linder) Galley & H. P. Linder, P. reflexa (H. P. Linder) Galley & H. P. Linder, P. rigidissima (Pilg. ex H. P. Linder) Galley & H. P. Linder, P. rosea (H. P. Linder) Galley & H. P. Linder, P. rosea subsp. purpurascens (H. P. Linder) Galley & H. P. Linder, P. scandens (H. P. Linder) Galley & H. P. Linder, P. setifolia (Thunb.) Galley & H. P. Linder, P. tomentella (Stapf) Galley & H. P. Linder, P. trifida (Galley) Galley & H. P. Linder, P. triseta (Thunb.) Galley & H. P. Linder, P. trisetoides (Hochst. ex Steud.) Galley & H. P. Linder, P. velutina (H. P. Linder) Galley & H. P. Linder, P. veneta (H. P. Linder) Galley & H. P. Linder, Cortaderia hieronymi (Kuntze) N. P. Barker & H. P. Linder, C. peruviana (Hitchc.) N. P. Barker & H. P. Linder, Austroderia fulvida (Buchanan) N. P. Barker & H. P. Linder, A. richardii (Endl.) N. P. Barker & H. P. Linder, A. splendens (Connor) N. P. Barker & H. P. Linder, A. toetoe (Zotov) N. P. Barker & H. P. Linder, A. turbaria (Connor) N. P. Barker & H. P. Linder, Chimaerochloa archboldii (Hitchc.) Pirie & H. P. Linder, Tenaxia aureocephala (J. G. Anderson) N. P. Barker & H. P. Linder, T. cachemyriana (Jaub. & Spach) N. P. Barker & H. P. Linder, T. cumminsii (Hook. f.) N. P. Barker & H. P. Linder, T. disticha (Nees) N. P. Barker & H. P. Linder, T. dura (Stapf) N. P. Barker & H. P. Linder, T. guillarmodiae (Conert) N. P. Barker & H. P. Linder, T. stricta (Schrad.) N. P. Barker & H. P. Linder, T. subulata (A. Rich.) N. P. Barker & H. P. Linder, Schismus schismoides (Stapf ex Conert) Verboom & H. P. Linder, Tribolium curvum (Nees) Verboom & H. P. Linder, T. pleuropogon (Stapf) Verboom & H. P. Linder, T. purpureum (L. f.) Verboom & H. P. Linder, T. tenellum (Nees) Verboom & H. P. Linder, Rytidosperma bipartitum (Kunth) A. M. Humphreys & H. P. Linder, R. diemenicum (D. I. Morris) A. M. Humphreys & H. P. Linder, R. fulvum (Vickery) A. M. Humphreys & H. P. Linder, R. lepidopodum (N. G. Walsh) A. M. Humphreys & H. P. Linder, R. pallidum (R. Br.) A. M. Humphreys & H. P. Linder, R. popinensis (D. I. Morris) A. M. Humphreys & H. P. Linder, R. remotum (D. I. Morris) A. M. Humphreys & H. P. Linder. Typifications are designated for the following names: Achneria Munro ex Benth. & Hook. f., Avena aristidoides Thunb., A. elephantina Thunb., Danthonia crispa Nees var. trunculata Nees, Danthonia sect. Himantochaete Nees, D. zeyheriana Steud. var. trichostachya Stapf, Geochloa lupulina, Pentameris aristidoides, and P. holciformis. Key words: Danthonioideae, generic delimitation, Poaceae, taxonomy. The Danthonioideae constitutes a small, welldefined clade currently recognized to comprise 281 largely Southern Hemisphere grass species. The genera that make up the clade were recognized as a coherent group only after 1957, with the recognition of the Arundinoideae (Tateoka, 1957), in which they were included. However, the present delimitation of the clade dates from the early molecular phylogenetic work of Barker et al. (1995). At the same time, Verboom et al. (1994) were able to show that the clade could also be delimited by the possession of haustorial synergids. The subfamily was formally erected by the Grass Phylogeny Working Group (2001). Generic delimitations in the Danthonioideae have been remarkably unstable (Reimer & Cota-Sanchez, 2007). De Candolle (1805) recognized Danthonia DC. in 1805 based on the American D. spicata (L.) P. Beauv. ex Roem. & Schult. and characterized it by the bilobed, awned lemmas. Soon after that, Palisot de Beauvois (1812) described Pentameris P. Beauv. (only two florets per spikelet, fruit an achene) and Schismus P. Beauv. (lemma unlobed, or with short lobes, and with a short, simple awn). During the 1830s, Tribolium Desv. (lemmas acute), Chaetobromus Nees (spikelets subtended by a tuft of hair), Prionanthium Desv. (annuals, glume keels with forked, multicellular glands), and the section Pentaschistis Nees of Danthonia (only two florets per spikelet, fruit a caryopsis) were added. Bentham and Hooker (1883) simplified matters again by including Pentameris, Pentaschistis (Nees) Spach, Chaetobromus, Plin- thanthesis Steud., as well as several genera now no longer retained in the Danthonioideae, in Danthonia. This left Prionanthium (as Prionachne Nees), Schismus, and Tribolium (as Lasiochloa Kunth) as separate genera. Several more genera were erected after the publication of the Genera Plantarum. Cortaderia Stapf (gynodioecious, glumes single-veined, florets pilose) was described in 1897 by Stapf, who also erected Poagrostis Stapf, a single-flowered species of Pentaschistis (Stapf, 1899). The last genera to be described, based on new species, rather than subdivisions of previously described genera, were Lamprothyrsus Pilg. (Pilger, 1906) from South America, Monostachya Merr. (Merrill & Merrit, 1910) from New Guinea, and Notochloe Domin (Domin, 1911) from Australia. However, by 1900, both species currently recognized in the Danthonioideae were included in the large genus Danthonia, which occurred on all continents, and which basically included all species with bilobed lemmas with geniculate awns inserted between the lobes, and with more than two florets in the spikelets. Species that deviated from this (with only two florets in the spikelets, or without lemma lobes or lacking a geniculate awn) were placed in various small segregated genera. This concept prevailed for almost a century. It was evident that Danthonia was not natural, and the dismantling of this genus was pursued in parallel for the African and Australasian species. This process 308 Annals of the Missouri Botanical Garden was started in New Zealand by Zotov (1963), who published a concise and insightful paper in which he segregated the New Zealand species of Danthonia into his new genera Chionochloa Zotov, Erythranthera Zotov, Pyrrhanthera Zotov, and Notodanthonia Zotov (5 Rytidosperma Steud. [Nicora, 1973]). He also grouped the currently recognized danthonioids into two tribes: Cortaderieae and Danthonieae. This process was paralleled in Africa by Conert and his associates. All sub-Saharan species of Danthonia were moved into the new genera Karroochloa Conert & Türpe (Conert & Türpe, 1969), Merxmuellera Conert (Conert, 1970), and Pseudopentameris Conert (Conert, 1971). In Australia, Blake (1972) separated out Plinthanthesis Steud. (although the name dates back to Steudel [1853–1854]), and Veldkamp (1980) transferred the Australian, Malesian, and many South American Danthonia species to Notodanthonia. However, all Notodanthonia species had to be transferred to the older generic name, Rytidosperma (Connor & Edgar, 1979). This fragmentation resulted in an unsatisfactory delimitation of Rytidosperma. It seemed likely that the small genera Monostachya, Pyrrhanthera, and Erythranthera might be embedded within Rytidosperma. Further, the relationship to the very similar South African species of Karroochloa and some species of Merxmuellera was not clear. Clayton and Renvoize (1986) followed a rather broad solution to these issues and included many of these segregates of Danthonia in a much expanded Rytidosperma. Linder and Verboom (1996), by contrast, defined a narrower Rytidosperma and segregated out Austrodanthonia H. P. Linder, Joycea H. P. Linder, and Thonandia H. P. Linder (5 Notodanthonia Zotov), but included Pyrrhanthera, Erythranthera, and Monostachya in Rytidosperma s. str. They indicated the close relationship between this clade and the African Karroochloa, Schismus, Urochlaena Nees, and Tribolium. The first species-level molecular phylogenetic investigations unveiled a host of further problems. Barker et al. (2003) demonstrated that Cortaderia was polyphyletic with the New Zealand and South American segregates not being sister clades. Furthermore, they showed that the African Merxmuellera species belong to three distinct, unrelated clades and that M. papposa (Nees) Conert and M. rangei (Pilg.) Conert should be placed in a different subfamily, the Chloridoideae (Barker et al., 2000, 2007). Verboom et al. (2006) showed that the generic limits between Schismus, Karroochloa, and Tribolium were misplaced, and that none of these three genera are monophyletic. A recent detailed molecular phylogenetic study by Pirie et al. (2008) provided a phylogenetic framework with which to address these problems in detail; it included 81% of all known species in the subfamily and most nodes were robustly resolved. It also demonstrated the widespread occurrence of reticulation. Here, we use this phylogenetic framework and the available morphological data to erect a new generic classification for the subfamily. We develop and apply explicit criteria for the main ranks (generic and infrageneric). We also make all the new combinations needed to apply the new generic classification at species level. MATERIALS AND METHODS TAXA, DATA, AND ANALYSIS Morphological descriptions were based on the study of herbarium collections in AD, B, BM, BOL, C, CANB, CHR, CONC, GRA, HO, K, L, MEL, NBG, NSW, NU, P, PRE, S, US, Z, and ZT over the past 15 years and were generated in the process of preparing revisions or flora accounts (Baeza, 1990, 1996a; Linder & Ellis, 1990a; Barker, 1993, 1995, 1999; Linder, 1997, 1999, 2005; Linder & Davidse, 1997; Verboom & Linder, 1998; Galley & Linder, 2006). The original observations were reported in these publications. Almost all species have been observed in the field, often at several locations, and over several years. Further descriptive information was taken from floras and revisions. Details of the vasculature of the lemmas and lodicules were obtained by preparing mounts of these organs in glycerine and stained with fuchsine. Caryopsis data for African taxa were taken from Barker (1986, 1994). Anatomical sections were collected over many decades, by several laboratories, and using diverse methods. Leaf material was largely taken from the midportion of the blades, mostly from material fixed in the field in FAA, but also from herbarium material reconstituted in boiling soapy water. Sections were cut either free-hand or by sledge-microtome, or from material embedded in wax or paraplast, using a rotary microtome. Section thickness varied between 10 and 30 mm. Sections were mostly double-stained and mounted in xylene or Canada balsam. Epidermal scrapes were prepared by softening leaves, then scraping off the mesophyll. Most slides are currently housed in the Institute of Systematic Botany of the University of Zurich or at the South African National Biodiversity Institute, Pretoria. The descriptive terminology follows Ellis (1976, 1979). We made no original cytological or embryological observations for this research but incorporated the published results. There have been many investiga- Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 309 tions into the number of chromosomes in the subfamily (Calder, 1937; Stebbins & Love, 1941; Myers, 1947; de Wet, 1953, 1954; Gould, 1958; Abele, 1959; Bowden, 1960; Brock & Brown, 1961; Löve & Löve, 1961; Bowden & Senn, 1962; Borgmann, 1964; Packer, 1964; Schwartz & Baessler, 1964; Mehra & Kalia, 1975; Moore et al., 1976; Reeder, 1977; Sokolovskaya & Probatova, 1978; Faruqi & Quraish, 1979; Beuzenberg & Hair, 1983; Davidse et al., 1986; du Plessis & Spies, 1992; Spies et al., 1994; Visser & Spies, 1994b, c; Baeza, 1996b; Spies & Roodt, 2001; Hilu, 2004; Murray et al., 2005), but there are still some gaps in the form of genera for which no counts are available. The embryology has also received some attention, in particular the development of haustorial synergids (Philipson & Connor, 1984; Verboom et al., 1994). The reproductive system was investigated by Connor (1967, 1970, 1979, 1981, 1991). The full descriptive information at species level was compiled in the software DELTA (Dallwitz, 1980; Dallwitz & Paine, 1986) and will be released as an INTKEY interactive key. This will also include the full nomenclatural information on each species, including the synonymy. Phylogenetic results presented here are based on Pirie et al. (2008), which included 256 samples of 227 ingroup species (plus 14 subspecific taxa), representing 81% of the danthonioid species recognized here. A matrix including multiple chloroplast and nuclear ribosomal DNA sequence markers (cpDNA and nrDNA) was constructed (available on TreeBase, accession number S10417), assembled from new sequences generated for that study together with sequences obtained from previous studies (Barker et al., 1995, 2000, 2003, 2007; Verboom et al., 2006; Galley & Linder, 2007). Additional PACCAD (Panicoideae, Arundinoideae, Chloridoideae, Centothecoideae, Aristidoideae, and Danthonioideae) clade outgroup sequences were obtained from GenBank. DNA extraction, polymerase chain reaction (PCR), and sequencing protocols are described in the above studies and in Pirie et al. (2008). The following markers were used: cpDNA from the trnL intron and trnL-F intergenic spacer, rpl16 spacer, atpB-rbcL spacer, ndhF gene, matK gene (including flanking spacer regions), and the rbcL gene; and nrDNA sequences from the ITS region and an 1100-bp-long fragment of the 26S gene. In total, more than 8200 bp of cpDNA and more than 1800 bp of nrDNA were sampled. Sequences were aligned manually and gaps were coded as separate presence/absence characters. A novel ‘‘taxon duplication’’ method was used to combine a single analysis of cpDNA and nrDNA markers despite conflicting phylogenetic signals. This approach is described in detail in Pirie et al. (2008, 2009), but is worth summarizing here in order to aid interpretation of the phylogenetic tree. Separate analyses were performed on each of the markers and the resulting topologies were inspected for conflicting nodes supported by 70% or higher bootstrap support (BS). Where no such conflict was found, data partitions were combined. Where conflicting nodes were found, the corresponding inconsistently placed taxa were duplicated in the matrix, with one taxon copy represented by the corresponding cpDNA sequences only, with the nrDNA partition coded as missing data, the other taxon copy by nrDNA only, with the cpDNA coded as missing. The partitions were then combined. The positions of the following taxa were subject to such conflict: a clade including all South American Cortaderia species plus Lamprothyrsus, without the New Zealand Cortaderia species, C. archboldii (Hitchc.) Connor & Edgar or C. pilosa (d’Urv.) Hack ex Dusén (in total 11 species, representing the same conflict between the chloroplast and nuclear genomes); 10 species of Pentaschistis (each representing a separate incidence of conflict); Notochloe microdon (Benth.) Domin (three samples); Merxmuellera arundinacea (P. J. Bergius) Conert (two samples); Danthonia alpina Vest (one sample); Chionochloa australis (Buchanan) Zotov (one sample); Tribolium ciliare (Stapf ) Renvoize (one sample); and T. pusillum (Nees) H. P. Linder & Davidse (one sample). These 30 samples, representing 27 species, were thus represented twice in the matrices, bringing the number of terminals included in combined cpDNA and nrDNA analyses up to 290. Parsimony analyses (heuristic search and bootstrapping) were performed using the software package PAUP* 4.0b10 (Swofford, 2002), and Bayesian inference was performed using MrBayes 3.12 (Ronquist & Huelsenbeck, 2003). For details, see Pirie et al. (2008). Generic delimitation criteria. We attempt to define the genera of the Danthonioideae according to explicit criteria. These criteria can be grouped into two categories: those that pertain to the delimitation of the clades (thus defining the content of the genera), and those that pertain to the ranking of the clades (thus which clades should be ranked as genera). We delimited the clades (monophyletic groups) primarily on the phylogeny derived from DNA sequence data. This is based on the assumption that the phylogeny provides the optimal prediction for the characters of the groups. Morphological data were used in two situations. The first was to assign to the correct clades those species that were not included in the molecular phylogeny, either because we could not 310 Annals of the Missouri Botanical Garden obtain DNA suitable for sequencing, or because we had difficulty obtaining sequences. The morphological characteristics (diagnostic attributes) of the genera were derived from the species assigned in the molecular study to the genera. The second situation was where there was incongruence between the nuclear and plastid genomes. In this situation, the decision on which phylogeny to follow was informed by the morphological similarity, and in particular by shared derived characters. In this latter situation, however, distributional and ecological data were also considered, in order to minimize the heterogeneity of the genera. Defining clades where one nuclear partition is nonmonophyletic differs from the strategy proposed by Potter et al. (2007), in which only uncontradicted clades were recognized, and contradicted nodes result in new genera. Our strategy is less conservative. Here, two logical routes are possible. In the first (and presumably most widely used) approach we sought positive evidence that a species is a member of a clade. This evidence was evaluated by means of bootstrap or posterior probability values. The second approach sought positive evidence that inclusion of a species in a clade renders that clade paraphyletic or polyphyletic. In most cases we used the second criterion. This implies that if a species was previously placed in a clade, or if morphological evidence indicates that it belongs in a particular clade, it was removed from that clade only if there was positive evidence from the molecular data that its inclusion would result in a paraphyletic or polyphyletic group. We used several ranking criteria for the establishment of genera, listed here in no particular order. ical convergence within the subfamily that is reflected by the frequency of reticulation evidenced by the molecular phylogeny (Barker et al., 2000; Pirie et al., 2008). We largely accepted that it will not be possible to key out the genera (although regional floras may not encounter similar conflicts) and recommend that the users use a key to the species, rather than to the genera. 4. The number of monotypic genera should be minimized. Monotypic genera are generally established because they are morphologically aberrant in their ‘‘clade.’’ If this aberrance is due to a long evolutionary history, then they could be regarded as distinct, by definition monophyletic, lineages. Extreme cases of this argument are the genera Welwitschia Hook. f. and Ginkgo L. Monotypic genera could also be established to indicate phylogenetic uncertainty, either due to conflict between the genome partitions or due to lack of basal resolution, as was the case in the genus Caulipsolon Klak (Klak & Linder, 1998). The latter approach has the advantage of highlighting species that need more research. If these topologically uncertain species are placed in their most likely or demonstrated sister genera, there is a danger that they will be forgotten, and their attributes summed to those of the more inclusive genera. However, this does have the advantage of minimizing the number of taxa erected, and so can be regarded as being nomenclaturally conservative. 5. Generic rank should take into account the age of the clades. Hennig (1966) suggested that ranks could be associated with the absolute age of the taxa, and Goodman et al. (1998) proposed a classification of the primates in which the ranks of the clades are related to the crown ages. 1. Genera should be delimited to minimize nomenclatural changes. This means that we did not sink genera, or erect new genera, where the existing genera did not violate the monophyly requirement, as interpreted above (Funk, 1985). 2. Genera should be morphologically, ecologically, and geographically homogeneous. Ideally, the disparity within a genus should be minimized (e.g., number of continents occupied, habitat range, morphological variance). However, in many cases this has been very difficult to achieve. 3. It should be possible to diagnose genera morphologically. Fortunately, the morphological and nomenclatural criteria generally coincide, as in the past genera were delimited on precisely these morphological criteria. Ideally, it should be possible to construct a simple key to the genera, based on characters that are readily observed on herbarium specimens. This, however, was almost impossible, due to the very extensive morpholog- We explicitly did not delimit genera as the smallest diagnosable, monophyletic (e.g., polytypic) units. We rather sought larger, more inclusive units, akin to the generic concept that Bentham used in the Genera Plantarum (see Humphreys & Linder, 2009), that also meet the geographical, morphological, and ecological criteria listed above. The smallest diagnosable, polytypic clades are recognized as sections. If another level of such units is available, we recognize them as subgenera. RESULTS AND DISCUSSION MOLECULAR PHYLOGENY FOR ALL SPECIES The topologies derived from the individual cpDNA markers were without significantly supported incongruence. The nuclear markers ITS and 26S also showed no significantly supported incongruence, but the numbers of informative characters, and thus Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae degree of resolution, were lower than those for the cpDNA analyses. The combination of all the data, using the taxon duplication method to accommodate conflict between cpDNA and nrDNA, resulted in strong support for the alternative positions of the conflicting taxa and greatly increased resolution and support with respect to the nrDNA tree (Fig. 1). Parsimony bootstrap support values for some clades in the combined analysis decreased in comparison to the values obtained from analysis of cpDNA only. However, Bayesian posterior probabilities for the same nodes remained high. This can, at least in part, be explained by the relative robustness of Bayesian inference to the large proportion of missing data representing the duplicated (conflicting) taxa (Pirie et al., 2008). minimize nomenclatural changes. A monotypic subfamily carries very little information, as it duplicates all the information already contained in the genus. The segregates would provide much more information, as each genus specifies a part of the variation within the subfamily. On the basis of these arguments, we think that it would be better to retain the generic rank at the segregate level, rather than have a single genus for the subfamily. GENERIC DELIMITATION A single mega-Danthonia. A simple solution would be to reconstruct the old genus Danthonia, as the sole genus in the subfamily, based on node A (Fig. 1A). The single-large-genus solution has recently become popular, with the reassembly of very large genera, such as Erica L. (Oliver, 1991), Disa P. J. Bergius (Bytebier et al., 2007), Eucalyptus L’Hér. (Brooker, 2000; Ladiges & Udovicic, 2000), Veronica L. (Garnock-Jones et al., 2007), Banksia L. f. (Mast & Thiele, 2007), Moraea Mill. (Goldblatt et al., 2002), and Ornithogalum L. (Manning et al., 2004), but equally there is some splitting of large, old genera, such as Acacia Mill. (Maslin et al., 2003) (reviewed in Humphreys & Linder, 2009). In the case of Danthonia, it would be only a medium-sized genus of some 281 species. However, there are several arguments against this solution. Some genera have been separated from Danthonia for more than a century (e.g., Pentaschistis, Schismus, and Pentameris), much of the breaking-up of Danthonia was completed more than 20 years ago, and the segregates have become well established and are widely used in diverse floras (e.g., Southern Africa [Gibbs Russell et al., 1990]; Australia [Mallett & Orchard, 2002]; New Zealand [Edgar & Connor, 2000]). Many of the segregate genera are morphologically distinctive, and their species share a likeness. It would be inconvenient to lose this shorthand summary of these patterns of similarity. Most of the segregate genera occupy a particular continent, thus only part of the range of the whole subfamily. These are generally geographically and also ecologically definable. A mega-Danthonia would include many species that have never been included in Danthonia and would therefore require 142 new combinations, which would be undesirable given that one of the criteria is to 311 Merxmuellera basal grade. Barker et al. (2000, 2007) showed that the genus Merxmuellera, as delimited by Conert (1970, 1971), is grossly polyphyletic, and this is confirmed by the results of Pirie et al. (2008). Two species (M. papposa and M. rangei) are in the subfamily Chloridoideae, with the remainder in Danthonioideae. Three clades (newly named here Geochloa, Capeochloa, and Merxmuellera) form a basal grade relative to the rest of the subfamily. A fourth clade (Tenaxia) is sister to the Rytidosperma clade. Merxmuellera s. str. (Fig. 1A, node B) is ecogeographically distinct and is Afromontane, ranging from the Drakensberg in southern Africa to the mountains of Madagascar and north to Ethiopia. The species of the genus can be recognized by the synapomorphy of the leaf blade disarticulating shortly above the ligule, thus leaving a distinctive stub, and by the more or less diagnostic indumentum pattern on the lemma back. There are no obvious breaks in the variation in this clade; only one of the four species is morphologically aberrant. The second basal segregate (Fig. 1A, node Cii) is almost impossible to define morphologically, but receives strong molecular support. It contains three very distinct elements, thus making it heterogeneous. The one element contains three villous geophytic grasses (Merxmuellera rufa (Nees) Conert, M. decora (Nees) Conert, M. lupulina (L. f.) Conert); the second, two large caespitose grasses (M. arundinacea, M. cincta (Nees) Conert); and the third, a very strange glabrous sedgelike geophytic grass (M. setacea N. P. Barker; Barker & Ellis, 1991). We are ambivalent about whether to recognize three, two, or one genera. It would be simple to group the three elements on the basis of their molecular support, and to recognize a highly heterogeneous genus. However, a closer inspection of the cladograms and chronograms (unpublished data) shows that the three geophytic grasses diverge very early from the remaining three species, and this supports the argument for recognizing them at generic level, on the basis that genera should have more or less equivalent ages. The recognition of a separate genus for the geophytic Cape grasses leaves us with a difficult problem with the remaining three species, but it is a 312 Annals of the Missouri Botanical Garden Figure 1. A–D. Phylogeny with specimens as terminals, named according to the current taxonomy. Where more than one specimen per species was sampled, these are discerned with voucher codes (see Pirie et al., 2008). Specimens for which the nuclear and plastid partitions are significantly incongruent are indicated in bold and the two partitions are represented separately: the nuclear partition is indicated by ‘‘nrDNA’’ and the plastid by ‘‘cpDNA’’ following the taxon name. Node support for clades referred to in the text is indicated by three values for bootstrap support (cpDNA/nrDNA/combined) and a single value for Bayesian posterior probability (combined). Where the clades are not identical in the two partitions, the relevant support value is marked by an asterisk. The new genera are indicated to the right of the cladogram. Volume 97, Number 3 2010 Figure 1. Continued. Linder et al. Classification of Danthonioideae 313 314 Figure 1. Continued. Annals of the Missouri Botanical Garden Volume 97, Number 3 2010 Figure 1. Continued. Linder et al. Classification of Danthonioideae 315 316 Annals of the Missouri Botanical Garden problem that is not solved through lumping them with the geophytic species in this clade. The clade of geophytic grasses is easy to diagnose by the swollen rhizomes clothed in furry sheaths and the compact inflorescences. Recognizing this clade at the generic level makes it easy to refer to this unusual habit. Furthermore, it draws attention to their (for grasses) unusual fire biology, flowering in the year after fire and then persisting vegetatively during the interfire periods (Linder & Ellis, 1990b). This biology is well known from Iridaceae, Orchidaceae, and Asparagaceae s.l. in fynbos (le Maitre & Midgley, 1992). Consequently, we erect the genus Geochloa to accommodate these three species. The remaining three species (Merxmuellera arundinacea, M. cincta, and M. setacea) do not form a morphologically sensible group. Merxmuellera arundinacea and M. cincta are tall, caespitose grasses that share some similarities in the spikelet construction, but these features are not remarkable and could well be plesiomorphic. It would satisfy most generic criteria if we grouped these two together, but the genus would be difficult to diagnose. Merxmuellera setacea is much more problematic. It has few characters in common with the two caespitose grasses and is nested between them on the nuclear DNA sequence data. The molecular data of M. arundinacea are problematic. The plastid DNA places M. arundinacea in an isolated position near the base of the phylogeny (Fig. 1A, node Ci), whereas the nuclear DNA groups it next to the morphologically similar M. cincta (Fig. 1A, node Cii). While it is clear where the nuclear affinities lie, the plastid affinities are unclear and could be used to argue for a segregate genus. Indeed, if we only had the plastid DNA, this might have been recognized as a segregate genus. Ecologically, M. arundinacea differs somewhat from the rest of the clade, in that it occupies drier habitats and is mostly found on shale, although it has also been recorded from sandstone-derived soils. The other species are more restricted to sandstonederived soils. However, this ecological range is not unusual in the Cape flora. Consequently, we choose to group the three species into a new genus, Capeochloa H. P. Linder & N. P. Barker. This is based on the strong morphological similarity between M. arundinacea and M. cincta, thus between two of the three included taxa. It also minimizes the number of monotypic genera and is consistent with the nuclear phylogeny. The implication is that we assume that M. arundinacea is of hybrid origin, with the plastid genome derived from a now extinct lineage that was sister to the rest of the Danthonioideae, while the nuclear genome derives from a lineage related to M. cincta. Pentameris clade. Currently, this readily diagnosed clade (Fig. 1B, node Di) includes three genera (Pentameris, Pentaschistis, and Prionanthium [Galley & Linder, 2007]), as well as Pseudopentameris obtusifolia (Hochst.) N. P. Barker (Pirie et al., 2008). Prionanthium is morphologically unique, with forked multicellular glands and an acute lemma (Davidse, 1988). Although this small genus of three rare, annual species is clearly monophyletic, it is deeply embedded within Pentaschistis and can only be retained at the cost of fragmenting Pentaschistis into many genera. Furthermore, Prionanthium is readily interpreted as a specialized annual Pentaschistis. We propose to include Prionanthium in Pentaschistis. Pentaschistis and Pentameris have been separated since 1830; only Steudel in his Nomenclator Botanicus (1841) placed all known Pentaschistis species under Pentameris. While Pentameris was always recognized as a separate genus, Pentaschistis was, until 1899, a section of the large genus Danthonia. Pentaschistis and Pentameris share several morphological characters, such as 2-flowered spikelets, lemma setae borne from the sinus between the lemma lobes and the central awn, and acute paleae with poorly developed keels that do not reach the apex of the palea. They have been separated in the past century by the presence of a villous indumentum on the ovaries and the free, brittle pericarp of the fruits of Pentameris (Barker, 1986). This character combination breaks down in Pentameris obtusifolia (Hochst.) Schweick., which has a villous ovary and a pericarp that is fused to the seed. There is no morphological evidence for the monophyly of Pentaschistis relative to Pentameris. It is therefore not surprising that molecular data group two Pentaschistis species (P. tysonii Stapf and P. praecox H. P. Linder, Fig. 1B, node Dii) with Pentameris (Fig. 1B, node Diii) rather than Pentaschistis. This makes it impossible to diagnose the two genera and leads to the suggestion that they should be combined. An alternative solution, which is more conservative nomenclaturally while still retaining monophyly, is to transfer P. tysonii and P. praecox to Pentameris. However, we then have two genera (Pentaschistis and Pentameris) that we cannot diagnose morphologically and that largely overlap ecologically and morphologically. The argument for combining the two genera is that the new, expanded genus would be easy to diagnose, while the various potential segregates are not. Ecologically, they are similar, as both occur in the mountains of southern Africa, on oligotrophic soils, in pyrophytic heathland or grassland. The habitat of Pentameris is a subset of the habitat range of Pentaschistis. The morphological range of Pentameris Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae is also part of the morphological range of Pentaschistis: the only unique Pentameris features are the brittle caryopses with a hairy cap. The strongest counterargument to a single large Pentameris is that Pentameris and Pentaschistis have been known as separate entities for almost two centuries, thus largely a historical argument. Indeed, it is not clear why the species of Pentaschistis were grouped as a section of Danthonia, rather than as part of Pentameris. If it had not been for this, there would not have been a problem with combining the two genera. The combined new genus is relatively big (including almost 100 species). Because the oldest name is Pentameris, this also results in many new combinations for the much larger genus Pentaschistis. It is tempting to conserve Pentaschistis over Pentameris, but since these genera are not well known and not economically important, the arguments for overriding the priority rule are not convincing. We will retain the distinction, though, by recognizing three sections. Danthonia clade. The evidence for the monophyly of the Danthonia clade is ambiguous (Fig. 1C, node W), and neither the nuclear nor the plastid phylogeny explicitly supports the clade. However, combining the two phylogenies suggests that the simplest solution is a Danthonia clade, which includes the genera Cortaderia, Plinthanthesis, Notochloe, Danthonia, and C. archboldii. This implies a single origin for the very unusual gynodioecious breeding system (Connor, 1979, 1981) in the Danthonioideae, with secondary losses in Danthonia and in Notochloe/ Plinthanthesis. The substantial incongruence between the phylogenies based on the nuclear and the plastid genomes is discussed in detail in a separate paper (Pirie et al., 2009). The monophyly of Danthonia s. str. is not challenged (Fig. 1C, node N). Within the genus there are two different lemma types: in the first the indumentum is restricted to the lemma margins, in the second it is evenly scattered on the lemma back. This could be used as a key for separating Danthonia into two genera, but since the genus is not otherwise heterogeneous (indeed less so than most other danthonioid genera of the same number of species) and occupies a coherent geographical area, we do not support this split. However, the African (D. subulata A. Rich., D. grandiflora Hochst. ex A. Rich.) and Himalayan (D. cumminsii Hook. f., D. cachemyriana Jaub. & Spach) species do not belong in Danthonia, but in the largely African segregates of Merxmuellera (Merxmuellera s. str., Tenaxia). The position of the New Guinean Cortaderia archboldii is problematic. The species has been included in either Chionochloa (Conert, 1975a) or Cortaderia (Connor & Edgar, 1974) and is morphologically intermediate between these two genera (Clayton & Renvoize, 1986). The molecular data set supports the grouping of C. archboldii with Danthonia, and more broadly the monophyly of C. archboldii, Cortaderia, and Danthonia (Fig. 1C, node Hii). Thus, surprisingly, if C. archboldii is to be grouped anywhere, it should be with Danthonia. However, there are several reasons for not including the species in Danthonia. Danthonia is restricted to America and Europe, and C. archboldii is found on the island of New Guinea. It makes sense to attempt to define genera that are geographically contiguous and that do not show such massive disjunctions. However, in itself geographical disjunction is not an argument for generic rank. Cortaderia archboldii is morphologically very different from Danthonia. The leaf blades abscise from the sheaths and are tough and sclerophyllous. Such tough, fibrous leaves do not otherwise occur in Danthonia. Also, the lemma structure is very different. In C. archboldii the lemmas are slender, Chionochloa clade. Zotov (1963) separated Chionochloa from Danthonia, based on the deeply grooved leaves with the microhairs along the bases of the grooves, the round silica cells, and the coarse tussocks. The genus is indeed very distinct morphologically and also has strong molecular support (Fig. 1A, node E). Furthermore, it forms a coherent ecogeographical entity, largely restricted to the more hilly and mountainous parts of New Zealand, with one species on Mt. Kosciuszko in Australia and another on Lord Howe Island in the Tasman Sea. At least the Australian species appears to be derived from New Zealand ancestors (Pirie et al., 2010). Chaetobromus–Pseudopentameris clade. The Chaetobromus–Pseudopentameris clade (Fig. 1C, node F + G) was recognized in the very first molecular analyses (Barker et al., 1995) and is strongly supported in the analysis of Pirie et al. (2008). Morphologically, the clade can be diagnosed by the very long, acute, almost pungent calli, the paleae that have parallel nerves that do not reach the tip, and the palea-backs that are inrolled so that the keels almost touch each other. These two genera are kept separate because they are morphologically readily diagnosed and ecogeographically distinct, even though Chaetobromus is monotypic. Chaetobromus occurs in Namaqualand along the southwest African coast, in semi-arid habitats and coastal sands (Verboom & Linder, 1998). Pseudopentameris is found along the much wetter south coast of southern Africa, where it is generally found in pyrophytic heathland on oligotrophic soils. 317 318 Annals of the Missouri Botanical Garden with three (or rarely five) veins, and with the lemma lobes absent or very poorly developed. Such lemmas are typical of Cortaderia. In Danthonia the lemmas are broad and soft, with seven to nine veins, and lemma lobes mostly as long as the lemmas themselves. The inflorescence in C. archboldii includes 50 to 150 spikelets, and although it is not plumose, it is certainly widely open and paniculate. In Danthonia, in general, the inflorescences have fewer than 50 spikelets and tend to be linear or lanceolate and somewhat more compact. Cortaderia archboldii has a gynodioecious breeding system (Connor, 1970), as is also found in Cortaderia, but not in Danthonia. The morphological and anatomical evidence is clearly against Cortaderia archboldii being included in Danthonia, suggesting rather a grouping with Cortaderia. However, the molecular evidence conclusively rejects the monophyly of Cortaderia plus C. archboldii. Consequently, we recognize a monotypic genus, Chimaerochloa H. P. Linder, for C. archboldii. The small eastern Australian genera Plinthanthesis and Notochloe are retrieved as clades in both genomic partitions. In agreement with the leaf anatomical data, the nuclear genome places the two genera as sisters (Fig. 1C, node Ji). However, the plastid data group Notochloe with Cortaderia pilosa (Fig. 1C, node Jii + L) and Plinthanthesis with the New Zealand species of Cortaderia. We follow the anatomical (unpublished data) and nuclear data and propose a sister-taxon relationship between Plinthanthesis and Notochloe. This is also corroborated by ecogeographical data: both taxa are found in pyrophytic heathy vegetation on oligotrophic soils, typical of the Sydney sandstones in Australia. Although Notochloe is monotypic, we retain the two genera because the three species of Plinthanthesis form a clade and the two genera are readily diagnosed by the spikelet structure, so there is no good reason for upsetting an established taxonomy. Cortaderia presents a number of problems. The disintegration of Cortaderia into a South America clade (Cortaderia s. str., Fig. 1C, nodes Hi and Hii) and a New Zealand clade (Austroderia, Fig. 1C, node I) as first reported by Barker et al. (2003), and extended here, is surprising. These large grasses are very similar in their general appearance as well as in their detailed spikelet structure, with single-veined glumes, pilose lemmas, reduced lemma venation, and absent or poorly developed lemma lobes. Only anatomical differences provide morphological support for the segregation between these two genera (the New Zealand segregate has multiple-veined leaves, while the South American segregate has leaves with a single midrib) and these by themselves would not have been adequate for the recognition of generic rank. However, there is molecular support for such a division, albeit in some ways ambiguous, from both the plastid and the nuclear partitions. Consequently, on the basis of recognizing monophyletic genera, we have chosen to retain Cortaderia for the South American species and to erect a new genus, Austroderia N. P. Barker & H. P. Linder, to accommodate the New Zealand species. Lamprothyrsus is deeply embedded within the South American species of Cortaderia. Although Lamprothyrsus is immediately recognizable by the long lemma awns, it is not retrieved as monophyletic by either genome partition. Furthermore, maintaining Lamprothyrsus would force a radical fragmentation of Cortaderia. Lamprothyrsus shares many vegetative and spikelet characters with Cortaderia, thus we include both in the same genus. More difficult to resolve is the position of the southern South America Cortaderia pilosa. The nuclear partition is uninformative: the species is placed in a polytomy with the bulk of the species of Cortaderia (Fig. 1C). The plastid partition (Fig. 1C) places it next to Notochloe, in the surprising arrangement of ([C. pilosa–Notochloe]–Danthonia)– (Plinthanthesis–Austroderia). It is evident that the nuclear genome does not contradict the monophyly of Cortaderia including C. pilosa, while the plastid genome strongly rejects the monophyly of Cortaderia including C. pilosa. Consequently, the inclusion of C. pilosa in Cortaderia implies that the plastid genome is misleading. This is possibly the result of hybridization, a scenario put forward to explain the situation in Notochloe and Merxmuellera arundinacea. We can test this by searching for more nuclear sequence data, which can either support or contradict a potential nuclear relationship of C. pilosa–Cortaderia. This is currently only a potential relationship, since the nuclear data set neither supports nor refutes it. Cortaderia and C. pilosa are morphologically and ecologically so similar that if they were reciprocally monophyletic sister-taxa, it would be difficult to justify recognition of two separate genera. Consequently, the phylogenetically conservative approach would be to place C. pilosa in a separate genus, since the reciprocal monophyly of C. pilosa and Cortaderia is not contradicted by either data set. By placing it in its own genus, the monophyly requirement is met in both DNA partitions. However, this is a very reductionist position: by placing each species in its own genus, monophyly will never be violated. Hence, we argue that the morphologically and nomenclaturally conservative option is to include C. pilosa in Cortaderia. This is also consistent with the treatment of M. arundinacea. Consequently, we follow that option here. Rytidosperma clade. The monophyly of the Rytidosperma clade (Fig. 1D, node O), comprising all species of Austrodanthonia H. P. Linder, Joycea, Karroochloa, Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 319 Notodanthonia, Rytidosperma, Schismus, and Tribolium, as well as five species of Merxmuellera and three species of Danthonia, is well supported. This opens the possibility of amalgamating these genera into a single genus whose defining characters would include a caryopsis with a punctate-ovate hilum, a large embryo, and the presence of lodicule microhairs. Such a broad delimitation bears some resemblance to the proposal from Clayton and Renvoize (1986), without the bulk of Merxmuellera. Clayton and Renvoize do qualify their delimitation, though, noting that the group is ‘‘somewhat heterogeneous; perhaps subdivisable.’’ There are several arguments against such a broad delimitation. Since the group would include Schismus, this would be the oldest name and would by priority force many nomenclatural changes, or alternatively, Rytidosperma would need to be conserved relative to Schismus. The clade is indeed morphologically diverse (e.g., species with and without awns, annuals and perennials, diverse lemma morphologies, etc.). A single genus thus defined would lack homogeneity and would also be cytologically very heterogeneous. The morphological characters, which may be used to define the clade, have very numerous exceptions, and the genus would be very difficult to diagnose. In order to minimize taxonomic change, and in favor of morphological homogeneity, we have therefore opted to retain the existing African segregate genera (transferring taxa where necessary to ensure generic monophyly). Under this option at least three African genera need to be recognized: Tenaxia, Schismus, and Tribolium. Necessary changes include (1) the transfer of Karroochloa schismoides (Stapf ex Conert) Conert & Türpe to Schismus, (2) the combination of Karroochloa with Tribolium, and (3) the establishment of a new genus, Tenaxia, to accommodate Danthonia cachemyriana, D. cumminsii, D. subulata, Merxmuellera aureocephala (J. G. Anderson) Conert, M. disticha (Nees) Conert, M. dura (Stapf) Conert, M. guillarmodiae Conert, and M. stricta (Schrad.) Conert. The argument for the recognition of Tenaxia and Schismus is based entirely on the molecular phylogeny, where these clades and their sister-clades receive strong support from both DNA partitions (nodes P and Q, respectively, in Fig. 1D). However, the situation is less clear in the case of Tribolium. The variation in this genus would also be consistent with the recognition of three genera (Verboom et al., 2006). Plagiochloa Adamson & Sprague, with two species (as Tribolium brachystachyum (Nees) Renvoize and T. uniolae (L. f.) Renvoize), is defined by a secund, spicate inflorescence (Fig. 1D, node S). The seven species of the Karroochloa clade (Fig. 1D, node T) all have stolons (unique within the subfamily) leading to a very distinctive growth form. Tribolium (Fig. 1D, node U) includes the non-stoloniferous species, but all species in Tribolium s. str. have hispid glumes, often with massive cushion-based hairs. Clearly, the three segregates can all be diagnosed, are morphologically and ecogeographically homogeneous, and do not force more nomenclatural changes than a single genus would do. Tribolium s.l. (Fig. 1D, node R) would be more difficult to diagnose, requiring character combinations, but it would still be morphologically and ecogeographically homogeneous. This solution also has the advantage of fewer genera, rather than large numbers of genera with few species in each. Consequently, we define a larger genus, Tribolium s.l., and recognize the three segregates at sectional rank. There have been numerous and different delimitations of Rytidosperma. The Australasian species included here in the genus were separated into Austrodanthonia, Rytidosperma, Notodanthonia, and Joycea by Linder and Verboom (1996). This was based on a cladistic analysis of morphological attributes, and the segregates were based largely on the patterns of lemma indumentum and the relative sizes of the callus and the rachilla internode. However, none of these genera were retrieved as being monophyletic based on the molecular results, and the phylogeny obtained had no strongly supported clades that bear resemblence to any of the segregate genera (Humphreys et al., 2010). This suggests that these delimitations cannot be maintained. The segregates Pyrrhanthera and Erythranthera proposed by Zotov (1963), as well as Monostachya, are embedded within a large, poorly differentiated Rytidosperma. The only sensible solution is therefore to delimit a large genus Rytidosperma (Fig. 1D, node V), returning to an extended version of Steudel’s (1853–1854) concept of the genus as a largely southern entity distinguished from its northern counterpart (Danthonia) by the lemma indumentum pattern. Morphological attributes. We mapped the most striking morphological attributes over a simplified phylogeny for the Danthonioideae (Fig. 2, Table 1, Appendix 1). When scoring the characters we did not take into account occasional exceptions. This mapped tree, as well as the key, is meant to illustrate the broad patterns of character variation in the subfamily. TAXONOMIC TREATMENT We list the species we accept under each genus (Appendix 2). The relevent new combinations are made, and the common synonyms are listed. Full synonymy and descriptions will be made available in the online interactive key. 320 Annals of the Missouri Botanical Garden Figure 2. Simplified phylogeny with genera as terminals and selected characters mapped. The character numbers are indicated above the squares, the new state below the squares. Filled-in squares are unique changes; empty squares are homoplasious. The character numbers and their states are listed in Table 1 and Appendix 1. The key to the genera presented below should not be treated as a strict, analytical key that will assign all species to their correct genera. Instead, we aim to indicate the concept of the genera. It is important to use all characters in the couplets. These might be contradictory for some species and we recommend that you follow the majority case (e.g., the lead that has the most attributes that fit). The reason for this is that in many of the genera unique attributes are lost in several species, and a key that attempts to take these individually into account would soon be a key to the species. KEY TO THE GENERA OF THE DANTHONIOIDEAE 1a. Spikelets with 2(1) florets with a minute rachilla extension; palea keels parallel; setae inserted in sinuses between lemma lobes and awns . . . . . . . . 2 1b. Spikelets generally with more than 2 florets, if only 2 then with a well-developed rachilla extension; palea keels sinuose; setae when present at apices of lemma lobes . . . . . . . . . . . . . . . . . . . . . . . . . 3 2a. Glumes more than 25 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . VI. Pseudopentameris 2b. Glumes usually less than 25 mm long; plants often with multicellular glands . . . . . . . . . . IV. Pentameris Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 321 Table 1. Morphological characters optimized on the simplified phylogenetic tree (Fig. 2). States between parentheses are polymorphic and indicate the states recorded for that terminal. Dashes indicate missing data or inapplicable states. Morphological characters and character states are described in Appendix 1. Capeochloa Chaetobromus Chionochloa Austroderia Cortaderia Danthonia Geochloa Chimaerochloa Merxmuellera Notochloe Pentameris Plinthanthesis Pseudopentameris Rytidosperma Schismus Tenaxia Tribolium 010100(01)0021100(01)10(01)100 010100100111000011101 11011010121100010(01)(01)00 110102110210001(01)01(01)00 110111100210001(01)0(01)(01)00 0(01)010010021100000(01)-01 011100100211000101100 110110100210000001000 01012000021120(01)00(01)100 010100110211301001101 010(01)0010020101000(01)(01)0(01) 010100110211000000101 01010010020101001(01)1-0 010100100211(01)00(01)0(01)(01)1(01) 010100100(12)11200(01)0(01)011 010100(01)002112000010(01)0 010100100(02)1100(01)(01)0(01)(01)11 3a. Lemmas with 1 to 3(to 5) veins; lemma apex acute or the lemma lobes small compared to the lemma body, setae usually present but small; glumes mostly with one vein; species gynodioecious or dioecious . . . . . . 4 3b. Lemmas with 5 to 9(to 13) veins; lemma apex lobed, setae generally present on the lemma lobes; glumes generally with more than one vein; spikelets generally bisexual . . . . . . . . . . . . . . . . 6 4a. Lemma awn differentiated into column and limb; lemma lateral lobes well developed; plants up to 1.4 m tall; from New Guinea . . . XII. Chimaerochloa 4b. Lemma awn not differentiated into column and limb; lemma lateral lobes not distinct; plants more than 1 m tall; from New Zealand or South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 5a. Leaf blades with several wide, strongly sclerified veins (including midrib) surrounding fine, indistinctly sclerified veins; from New Zealand . . . . . . . . . . . . . . . . . . . . . . . . . . IX. Austroderia 5b. Leaves with only the midrib prominent and strongly sclerified; from South America . . . . . . . . . . . . . . . . . . . . . . . . . VIII. Cortaderia 6a. Palea flaps often with long hair; lemmas with scattered indumentum . . . . . . . . . . . . . . . . . . . 7 6b. Palea flaps usually glabrous; lemma indumentum various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 7a. Geophytes with persistent, woolly leaf bases; from Africa . . . . . . . . . . . . . . . . . . . . . . . II. Geochloa 7b. Tall caespitose plants, or if geophytes, then leaf bases glabrous . . . . . . . . . . . . . . . . . . . . . . . . . 8 8a. Leaf blades often disarticulating; palea keels straight; lemma indumentum mostly in two lines flanking the lemma keel; native to Australasia . . . . . . . . . . . . . . . . . . . . . . . . . . V. Chionochloa 8b. Leaf blades persisting; palea keels sinuose; lemma indumentum either as a transverse row, or evenly scattered on the lemma back, or largely as submarginal rows; native to Africa . . . . . . . . . . . . . . . . . . . . . . . . . . III. Capeochloa 9a. Spikelets with tuft of hair from a disarticulation point along pedicel; basal florets different from upper florets . . . . . . . . . . . . . . VII. Chaetobromus 9b. Spikelets never with tuft of hair at a disarticulation point on pedicel; basal florets same as upper florets . . . . . . . . . . . . . . . . . . . . . . . . 10 10a. Callus shorter than rachilla; joint to rachilla horizontal; caryopsis turbinate; native to eastern Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10b. Callus longer than rachilla; joint oblique; caryopsis lorate-cylindrical . . . . . . . . . . . . . . . . . . 12 11a. Spikelets with 2 to 4 florets; lemmas dorsally with a short, dense indumentum . . . . . X. Plinthanthesis 11b. Spikelets with 7 to 9 florets; lemmas dorsally glabrous . . . . . . . . . . . . . . . . . . . . . . XI. Notochloe 12a. Hilum linear; embryo mark less than 2/5 of caryopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 12b. Hilum punctiform; embryo mark more than 2/5 of caryopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13a. Plants usually with cleistogenes; leaves orthophyllous, not disarticulating above the ligules; native to America or Europe . . . . . XIII. Danthonia 13b. Plants without cleistogenes; leaves sclerophyllous, generally disarticulating above the ligules; remaining leaf bases often split into 2 recurving halves; native to Africa . . . . . . . . I. Merxmuellera 14a. Plants native to the Pacific basin; tetraploid or higher; lemma indumentum often in 2 rows of tufts (with or without intervening scattered hairs); always perennial . . . . . . . . . . . XVII. Rytidosperma 14b. Plants native to Africa; diploid or higher; lemma indumentum various, but not obviously 2-rowed; plants sometimes annual . . . . . . . . . . . . . . . . . 15 15a. Glumes with tubercle-based hairs or plants stoloniferous or inflorescence spicate; lemmas often awnless . . . . . . . . . . . . . . . . XVI. Tribolium 15b. Glumes lacking tubercle-based hairs, not stoloniferous, and if the inflorescence is spicate then the lemmas have well-developed awns . . . . . . . 16 16a. Lemmas acute or lobed, central awn usually shorter than the lobes; plants less than 0.35 m tall; often annual . . . . . . . . . . . . . . . XV. Schismus 16b. Lemmas lobed; plants 0.12–0.9 m tall; always perennial . . . . . . . . . . . . . . . . . . . . . . . . XIV. Tenaxia 322 Annals of the Missouri Botanical Garden present and exceeding the lemma lobes, sometimes differentiated into a flat, corkscrewed base and a straight, hairlike apical part; palea linear, without tufts of hair on the palea flaps; lodicules rhomboid, lodicule upper margin fringed with numerous bristle hairs shorter than the body of the lodicule, microhairs apparently absent; ovary glabrous. Caryopsis 6 elliptical, embryo and linear hilum ca. 1/2 the caryopsis length. Cytology. Unknown. Anatomy. The leaves are sclerophyllous; adaxial ribs variously developed; adaxial sclerenchyma as massive T-shaped or inversely anchor-shaped girders associated with both 1- and 3-order vascular bundles; leaves asymmetrical with one side with more vascular bundles than the other; clear cells in the chlorenchyma absent; adaxial bulliform cells present in the grooves. Figure 3. Merxmuellera macowanii. —A. Spikelet. —B. Lemma. —C. Palea. Drawn by Jasmin Baumann from Schuette in BOL 30869. I. Merxmuellera Conert, Senckenberg. Biol. 51: 129. 1970. TYPE: Merxmuellera davyi (C. E. Hubb.) Conert (; Danthonia davyi C. E. Hubb.). Figure 3. Caespitose, perennial grasses without stolons, culms 0.3–1.5 m tall. Sheaths 6 persistent, not variously lacerated or fragmenting; ligule ciliate; leaf blades setaceous or expanded, sclerophyllous, tough, glabrous, occasionally with a weftlike indumentum on the upper surface directly above the ligule; mostly disarticulating 10–20 mm above the ligule leaving a short stub that is either entire or split, straight or more usually recurved. Inflorescences 6 paniculate, mostly open or more rarely contracted or linear. Spikelets usually with more than 2 florets, all similar and bisexual; glumes shorter to longer than the florets, with 1 to 3 nerves, glabrous or micro-scaberulose, 7– 35 mm; callus rounded or truncate, villous, shorter or longer than the rachilla internode; lemmas with 7 or 9 veins; lemma dorsal indumentum as 3 hair tufts arranged in a diagonal stripe from the midrib to the margin, the tufts sometimes poorly defined or missing, the marginal tufts sometimes well developed; lemma lobes usually extended into setae, lobes rarely fused to the central awn; central awn always Distribution and habitat. These are Afromontane tussock grasses. They are most common in montane grassland in Africa and Madagascar, usually in areas with a higher rainfall. Generally, they seem to be absent from bogs or waterlogged soils, but also avoid seasonally dry, freely draining habitats. These drier habitats are typically occupied by species of Tenaxia. These grasslands are subjected to regular fire (two- to four-year cycles) and often to frost in winter. Discussion. This genus is relatively clearly delimited, but no characters apply to all species. In all species except Merxmuellera stereophylla (J. G. Anderson) Conert, the leaf blade breaks off above the ligule and the remaining stub sometimes splits, sometimes not, and often curls or spirals. This attribute is unique to this genus in the Danthonioideae. In most species the awn lacks a clear separation into column and limb. Although the lower portion of the awn can corkscrew, it is not clearly differentiated from the upper portion of the awn. However, the degree of development of the column is a matter of interpretation. All species except M. stereophylla have a lemma indumentum organized into a diagonal stripe with ca. three tufts flanking the midvein of the lemmas. In M. stereophylla this pattern is somewhat simplified and can be interpreted as derived from the diagonal stripe of three tufts. The leaf blades are most often folded (thus setaceous) and are always sclerophyllous. Merxmuellera grandiflora may be an exception, as it has not yet been investigated anatomically. The lodicules are rhomboid (triangular), quite hairy in the upper half. Similar lodicules are also found in various other genera, often not in all species, but are smaller than those of Capeo- Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae chloa, where they are developed to quite massive structures. ovate, more rarely open; inflorescence branches mostly shorter than the spikelets; indumentum glabrous or puberulent. Spikelets with 2 to 7 florets, all similar and bisexual; glumes similar, at least as long as the florets, with 1 to 5 veins, 7–55 mm; callus blunt, villous, about as long as the rachilla internode; lemmas with 9 veins, with a scattered dorsal indumentum that may be longer below the sinus and become shorter toward the base of the lemma; lemma lobes sometimes extended into up to 2-mm setae, setae plus lobes 4.5–9 mm; central awn 4–20 mm, longer than the setae, the basal half forming a distinct column, straight or corkscrewed many times; paleae shorter than or overtopping the lemma sinus, glabrous to villous between the keels; palea keel flaps infolded with tufts of hair; lodicules rhomboid, with 3 veins, bristles shorter than the lodicules; microhairs absent. Caryopses obovate, shiny; hilum linear, less than 1/4 of caryopsis length. Included species. We include seven species in Merxmuellera, which is a part of the species originally assigned to the genus by Conert (1970, 1975b, 1987) and as treated by Barker (Gibbs Russell et al., 1990). 1. Merxmuellera ambalavaoensis (A. Camus) Conert, Senckenberg. Biol. 51: 132. 1970. 2. Merxmuellera davyi (C. E. Hubb.) Conert, Senckenberg. Biol. 51: 132. 1970. 3. Merxmuellera drakensbergensis (Schweick.) Conert, Senckenberg. Biol. 51: 132. 1970. 4. Merxmuellera grandiflora (Hochst. ex A. Rich.) H. P. Linder, comb. nov. Basionym: Danthonia grandiflora Hochst. ex A. Rich., Tent. Fl. Abyss. 2: 418. 1851. Pentameris grandiflora (Hochst. ex A. Rich.) A. Nelson & J. F. Macbr., Bot. Gaz. 56: 470. 1913. Rytidosperma grandiflorum (Hochst. ex A. Rich.) S. M. Phillips, Fl. Ethiopia & Eritrea 7: 74. 1995. TYPE: Ethiopia. ‘‘in monte Silke ad rupes,’’ 16 Feb. 1840, W. G. Schimper 690 (holotype, P not seen; isotypes, BM!, GOET!, K!, S!, TCD!, Z!). 5. Merxmuellera macowanii (Stapf) Senckenberg. Biol. 51: 132. 1970. Conert, 6. Merxmuellera stereophylla (J. G. Anderson) Conert, Senckenberg. Biol. 51: 133. 1970. 7. Merxmuellera tsaratananensis (A. Camus) Conert, Senckenberg. Biol. 51: 133. 1970. II. Geochloa H. P. Linder & N. P. Barker, gen. nov. TYPE: Geochloa lupulina (L. f.) H. P. Linder & N. P. Barker (; Avena lupulina L. f.). Danthonia DC. sect. Himantochaete Nees, Fl. Afr. Austral. Ill. 323. 1841. TYPE: Danthonia rufa Nees (; Geochloa rufa (Nees) N. P. Barker & H. P. Linder) (lectotype, designated here). Genus novum Merxmuellerae Conert affine, a qua rhizomatibus tumidis, vaginis foliaribus basalibus lanatis et inflorescentiis plerumque compactis differt. Plants perennial, tufted geophytes without stolons; rhizomes short, ascending, with swollen nodes forming perennating organs, encased in persistent, woolly sheaths; culms 0.25–0.75 m tall. Ligule with 1 or several rows of cilia; leaf blades expanded or rolled, tough. Inflorescence mostly compact, contracted, 323 Nomenclatural note. Nees included in section Himantochaete all African Danthonia species, except Pentaschistis. This concept of the section was expanded in Bentham and Hooker (1883) to the whole genus. Consequently, any of the southern African species included by Nees could be used as type. We selected D. rufa, as almost all species in the genus Geochloa are listed by Nees under section Himantochloa. Nees did not explicitly assign a rank to his infrageneric groups (Pentaschistis, Himantochaete); we interpret them here at sectional rank, rather than as subgenera. Cytology. 2n 5 48. This has been counted only for Geochloa decora (Nees) N. P. Barker & H. P. Linder (Du Pressis, pers. comm.). Anatomy. The leaves are sclerophyllous; adaxial ribs variously developed, often massive, separated by narrow clefts; adaxial sclerenchyma as T-shaped or inversely anchor-shaped girders associated with both 1- and 3-order vascular bundles; leaves symmetrical; adaxial bulliform cells present. Distribution and habitat. All three Geochloa species are restricted to the Cape Floristic Region (Goldblatt, 1978; Rebelo et al., 2006), where they are found widespread in the lowlands and mountains. Populations occur on both sandstone- and shalederived substrates, and consequently in both fynbos and renosterveld vegetation, in well-drained habitats, or lowland areas with some impeded drainage. The plants typically flower in the first year after fire, and flowering plants are rarely observed in older vegetation, although the leaves are visible. The fire cycle in this vegetation is between six and 50 years, which results in relatively rare flowering events for some populations. 324 Discussion. The small genus Geochloa shares with Capeochloa multiflowered spikelets, rhomboidbristled lodicules, and paleae with pilose margins. However, it can be diagnosed by several attributes. The plants are geophytes, with swollen rhizomes storing starch, and encased in woolly leaf sheaths. Geophytes are occasional in the grass flora of the Cape (Linder & Ellis, 1990b), but Geochloa can be separated from the geophytic Pentameris aristidoides (Thunb.) Galley & H. P. Linder by the 2flowered spikelets of the latter, and from the more closely related geophytic C. setacea (N. P. Barker) N. P. Barker & H. P. Linder by the shiny glabrous basal sheaths. In two of the three species, the inflorescence is contracted into a compact ovate structure; only in G. decora does it spread more openly during anthesis. Etymology. The name refers to the geophytic habit of the plants in this genus. This habit is unusual in the grasses and seems particularly characteristic of the three species in Geochloa. Included species. The three species included in this genus were originally classified in Danthonia (Stapf, 1899; Chippendall, 1955). They were then transferred to Merxmuellera by Conert (1971), a treatment also followed in southern Africa (Gibbs Russell et al., 1990). 1. Geochloa decora (Nees) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia decora Nees, Fl. Afr. Austral. Ill. 332. 1841. Merxmuellera decora (Nees) Conert, Mitt. Bot. Staatssamml. München 10: 306. 1971. TYPE: South Africa. s. loc., s.d., J. F. Drège 5651 (holotype, B not seen; isotype, B fragm. at FR!). Danthonia zeyheriana Steud., Syn. Pl. Glumac. 1: 244. 1854. TYPE: South Africa. Cape Province: Swellendam Division, Puspas Valley, s.d., C. L. P. Zeyher 4555 or 4556 (types, HBG!, K!, MO not seen, PRE!). Danthonia zeyheriana Steud. var. trichostachya Stapf, Fl. Cap. (Harvey) 7: 522. 1899. TYPE: South Africa. Cape Province: Cape Division, betw. Slang Kop & Red Hill, s.d., A. H. Wolley-Dod 3002 (lectotype, designated here, K!). Stapf listed two collections as syntypes: Wolley-Dod 1572 and Wolley-Dod 3002; the latter is selected as lectotype because it is a more complete specimen. 2. Geochloa lupulina (L. f.) N. P. Barker & H. P. Linder, comb. nov. Basionym: Avena lupulina L. f., Suppl. Pl.: 113. 1781. Danthonia lupulina (L. f.) P. Beauv. ex Roem. & Schult., Syst. Veg. ed. 16 (Sprengel) 2: 690. 1817. Merxmuellera Annals of the Missouri Botanical Garden lupulina (L. f.) Conert, Mitt. Bot. Staatssamml. München 10: 306. 1971. Danthonia coronata Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1: 70. 1831. TYPE: [South Africa.] Cape of Good Hope, s.d., C. P. Thunberg s.n. (lectotype, designated here, UPS 2604!; isotype, S!). Based on the assumption that the top set of Thunberg’s collections are indeed in his herbarium, and that he, rather than Linnaeus filius, wrote the descriptions of the Cape plants described in the Supplementum Plantarum, we designate the UPS collection as lectotype. 3. Geochloa rufa (Nees) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia rufa Nees, Fl. Afr. Austral. Ill. 330. 1841. Merxmuellera rufa (Nees) Conert, Mitt. Bot. Staatssamml. München 10: 306. 1971. TYPE: [South Africa.] ‘‘In iugi Cederbergen monte Blaauwberg locis saxosis alt. 40009,’’ s.d., J. F. Drège 2559 (type, PRE!). Avena lanata Schrad., Gött. Gel. Anz. 3: 2075. 1821, nom. illeg., non Avena lanata (L.) Koeler, Descr. Gram. (Koeler), 303. 1802. Danthonia lanata (Schrad.) Schrad., Mant. (Schultes) 2: 386. 1824. TYPE: South Africa. Cape Province: Cape Town, s.d., Hesse s.n. (holotype, GOET 2322!). Danthonia macrocephala Stapf, Fl. Cap. (Harvey) 7: 522. 1899. TYPE: [South Africa.] s. loc. [probably from Caledon, Tulbagh, or Clanwilliam Division], s.d., Thom s.n. (holotype, K!). III. Capeochloa H. P. Linder & N. P. Barker, gen. nov. TYPE: Capeochloa cincta (Nees) N. P. Barker & H. P. Linder (; Danthonia cincta Nees). Figure 4. Genus lemmatis destituto, foliaribus novum quod a Merxmuellera Conert indumento non caespitoso et phloemate cellulis scleroticis a Geochloa H. P. Linder & N. P. Barker vaginis basalibus glabratis recedit. Plants perennial, tufted, without stolons; culms 0.5–2.5 m tall; basal sheath either of leaf remnants, or of white shiny persistent leaf bases. Ligule either a simple or multiple row of cilia; leaf blades tough, expanded or inrolled, sometimes adaxially with a web of interlocking hairs above the ligule, sometimes pungent. Inflorescence open to plumose, with up to 200 spikelets, ovate or elliptical; inflorescence branches glabrous, scaberulose or villous. Spikelets with 2 to 4 fertile and bisexual florets; glumes at least as long as the florets, 8–23 mm, upper and lower glumes similar, with 1 vein; callus blunt, villous, as long as the rachilla internode; lemma 2.5–5 mm, bilobed, villous with the hairs either Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 325 Figure 4. Capeochloa cincta. —A. Spikelet. —B. Lemma back. —C. Palea. —D. Gynoecium. —E. Anthers. —F. Lodicules. Drawn by Jasmin Baumann from Ecklon & Zeyher 4548. 326 Annals of the Missouri Botanical Garden scattered over the backs, or longer along the lemma margin, or in a row across the lemma backs; lemma lobes acute to acuminate, sometimes with setae to 5 mm; lemma awn 5.5–17 mm, geniculate, with a 3– 4.5 mm, 6 twisted column base; palea truncate to bilobed, longer than the lemma sinus, with sinuose keels, sometimes with hair tufts on the infolded palea margins; lodicules sometimes with bristles, never with microhairs, obtriangular or rhomboid, with 3 to 4 veins; anthers 2–3.2 mm. Caryopsis not known. the lemma base; and C. setacea has only marginal tufts, while the rest of the lemma is glabrous. All species (except C. cincta subsp. cincta) have tufts of hair on the palea margins; these usually developed as a line of hairs. This feature is widespread in the subfamily, especially in Rytidosperma and its allies. Capeochloa, Geochloa, and Chionochloa seem to be the only genera in which this feature is found in almost all species. Cytology. 2n 5 12, 36 (de Wet, 1954). Anatomy. The leaves are sclerophyllous; adaxial ribs variously developed; adaxial sclerenchyma as massive T-shaped or inversely anchor-shaped girders associated with both 1- and 3-order vascular bundles; clear cells in the chlorenchyma are absent, and the grooves contain bulliform cells. Distribution and habitat. The genus is restricted to the Cape Floristic Region, where the species are found both in the lowlands and mountains. Most populations are on sandstone, but Capeochloa arundinacea (P. J. Bergius) N. P. Barker & H. P. Linder is also found on shales and can locally be a typical element in renosterveld vegetation. The geophytic species seems to flower only after fire, while the two large tussocks usually flower every year (except for some western populations of C. cincta). All species are restricted to shrubland or heathland and are absent from grassland. Discussion. This genus lacks unique morphological markers but has a number of unusual attributes. One of the three species is a geophyte, and the remaining two are robust tufted perennials. The genus can be separated from Geochloa by the glabrous plant bases and by the inflorescences, which are always open panicles. Both Capeochloa and Geochloa can be separated from Merxmuellera s. str. by the lemma indumentum never being tufted, by the usually wider leaves (more than 3 mm wide), by the absence of sclerosed cells in the phloem, and by the regular presence of tertiary vascular bundles between all primary vascular bundles. They all seem to have diamond-shaped lodicules, where the bristles are shorter than the lodicule, and the whole structure is substantial, about as big as the ovary. This could be unique but is difficult to clearly delimit from other, similar, structures in other genera. Furthermore, we have little data on variation within the species. The lodicules are larger than those in Merxmuellera. The lemma indumentum is unique to each species. In C. arundinacea it is more or less evenly scattered over the back of the lemma; C. cincta has a line of long hairs below the lemma sinus and with no hair toward Etymology. The name refers to the geographical center for the genus, as characteristic of the Cape flora. Included species. The four taxa, including three species, were initially classified in Danthonia (Stapf, 1899; Chippendall, 1955) before being transferred to Merxmuellera (Conert, 1970). Their inclusion in Merxmuellera has been generally accepted (Gibbs Russell et al., 1990). 1. Capeochloa arundinacea (P. J. Bergius) N. P. Barker & H. P. Linder, comb. nov. Basionym: Andropogon arundinaceus P. J. Bergius, Descr. Pl. Cap. 356. 1767, non Andropogon arundinaceus Scop., Fl. Carniol., ed. 2, 2: 274. 1772, nom. illeg. Andropogon bergii Roem. & Schultes, Syst. Veg. ed. 15 bis (Roemer & Schultes) 2: 813. 1817, nom superfl. Danthonia arundinacea (P. J. Bergius) Schweick., Notizbl. Bot. Gart. BerlinDahlem 14: 1938, non Danthonia arundinacea Steud., Nomencl. Bot., ed. 2, 1: 482. 1840. Merxmuellera arundinacea (P. J. Bergius) Conert, Senckenberg. Biol. 51: 132. 1970. TYPE: [South Africa.] s. loc., s.d., s. coll. (holotype, SBT!). Avena elephantina Thunb., Prodr. Pl. Cap. 23. 1794. Danthonia elephantina (Thunb.) Nees, Fl. Afr. Austral. Ill. 334. 1841. TYPE: South Africa. Swartland, s.d., Thunberg (lectotype, designated here, UPS 2593!). There are two specimens in the Thunberg herbarium: UPS 2593 and UPS 2594. These might be isotypes or they could be syntypes. Of these two collections, the former is selected as lectotype, as it is a better specimen. 2. Capeochloa cincta (Nees) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia cincta Nees, Fl. Afr. Austral. Ill. 332. 1841. Merxmuellera cincta (Nees) Conert, Senckenberg. Biol. 51: 132. 1970. TYPE: [South Africa.] ‘‘In Promotorio bonae spei, Reeves in Herb. Lindley,’’ s.d., Reeves s.n. (lectotype, designated by Conert [1970: 132], K not seen). 2a. Capeochloa cincta (Nees) N. P. Barker & H. P. Linder subsp. cincta. Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 327 nom. superfl. TYPE: Prionachne ecklonii Nees (; Pentameris ecklonii (Nees) Galley & H. P. Linder). Danthonia sect. Pentaschistis Nees, Index Seminum (Vratislav) 1835, nom. nud., Fl. Afr. Austral. Ill. 280. 1841. Pentaschistis (Nees) Spach, Hist. Nat. Veg. 13: 164. 1846. TYPE: Pentaschistis aristidoides (Thunb.) Stapf (; Avena aristidoides Thunb., ; Pentameris aristidoides (Thunb.) Galley & H. P. Linder) (lectotype, designated by Phillips [1951: 121]). Poagrostis Stapf, Fl. Cap. (Harvey) 7: 760. 1900. TYPE: Poagrostis pusilla (Nees) Stapf (; Colpodium pusillum Nees, ; Pentameris pusilla (Nees) Galley & H. P. Linder). Achneria Munro ex Benth. & Hook. f., Gen. Pl. 3: 1158. 1883, non Achneria P. Beauv., Ess. Agrostogr. 72, 146. 1812. Afrachneria Sprague, J. Bot. 60: 138. 1922, replacement name for Achneria Munro ex Benth. & Hook. f., 1883. TYPE: Achneria microphylla (Nees) T. Durand & Schinz (; Eriachne microphylla Nees, ; Pentameris microphylla (Nees) Galley & H. P. Linder) (lectotype, designated here). Figure 5. Pentameris curvifolia. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from MacOwan 1695. 2b. Capeochloa cincta (Nees) N. P. Barker & H. P. Linder subsp. sericea (N. P. Barker) N. P. Barker & H. P. Linder, comb. nov. Basionym: Merxmuellera cincta (Nees) Conert subsp. sericea N. P. Barker, S. African J. Bot. 65: 105. 1999. TYPE: South Africa. Eastern Cape: Rufanes river mouth, 9 Nov. 1997, N. P. Barker 1545 (holotype, GRA!; isotypes, BOL!, K!, MO!, NBG!, PRE!). 3. Capeochloa setacea (N. P. Barker) N. P. Barker & H. P. Linder, comb. nov. Basionym: Merxmuellera setacea N. P. Barker, Bothalia 21: 27. 1991. TYPE: South Africa. Western Cape: Groot Winterhoek Wilderness Area, s.d., R. P. Ellis 5500 (holotype, PRE!). IV. Pentameris P. Beauv., Ess. Agrostogr. 92, tab. 18, fig. viii. 1812. Danthonia DC. sect. Pentameris (P. Beauv.) Steud., Syn. Pl. Glumac. 1: 238. 1855. TYPE: Pentameris thuarii P. Beauv. Figure 5. Prionanthium Desv., Opusc. Sci. Phys. Nat. 64. 1831. TYPE: Prionanthium rigidum Desv. (; Pentameris dentata (L. f.) Galley & H. P. Linder). Prionachne Nees, Intr. Nat. Syst. Bot., ed. 2: 447. 1836. Chondrolaena Nees, Fl. Afr. Austral. Ill. 133. 1841, Plants annual, biennial, or perennial, caespitose, mat-forming, geophytic, or suffrutescent, sometimes with rhizomes or stolons; culms up to 1(2) m tall. Plants often with multicellular linear or stalked glands on the leaves, inflorescences, or glumes. Sheaths 6 persistent, not variously lacerated or fragmenting; ligule a simple line of cilia; leaf blades orthophyllous or sclerophyllous, expanded or setaceous, tough, occasionally with a weftlike indumentum on the upper surface directly above the ligule. Inflorescences paniculate and open, contracted, or linear. Spikelets with 2 or rarely 1 floret(s), all similar and bisexual; glumes longer than the florets, 2–25 mm, with 1 to 7 nerves, glabrous or micro-scaberulose, rarely with tufts of long hairs; callus rounded or truncate, villous, short; lemmas with 3 to 9 indistinct veins, dorsally pilose; lemma lobes weakly developed, acute, truncate, or lacerate; setae originating between the lobes and the central awn, often very long and exserted from the glumes; central awn absent or more usually present and exceeding the lemma lobes, generally differentiated into a flat, corkscrewed base and a straight, hairlike apical part; paleae linear, without tufts of hair on the palea margins, these not infolded, the 2 keels poorly developed, straight, and sometimes not reaching the palea apex; lodicules cuneate, generally without microhairs or bristles; ovary glabrous or with a hairy cap. Seed a caryopsis or achene (thus with free pericarp), 6 elliptical; embryo about 1/2 the length of the caryopsis; hilum usually linear to rarely punctate, up to 6/10 of the caryopsis length. Nomenclatural note on Pentaschistis. Pentaschistis was first mentioned as an unranked division of the genus Danthonia by Nees (1835) in a seed list, which was probably published at the end of 1835 or the beginning of 1836. He also listed the name in the second edition of Lindley’s A Natural System of 328 Annals of the Missouri Botanical Garden Botany (Lindley, 1836), which was published in July 1836. The name was validated, but again without explicit rank, on p. 280 in the Florae Africae Australioris, Illustrationis monographicae, I. Gramineae (Nees, 1841). The combination at generic rank was then made by Spach in 1846. The rank of Nees’s name is unclear, but is between species and genus. Because subgenera were rarely used in the 19th century and sections were commonly used, we interpret the name at sectional rank. The typification is difficult. Nees, in the seed list, included only D. glandulosa Schrad. in section Pentaschistis. However, the only lectotypification was by Phillips in 1951, who proposed P. aristidoides (Thunb.) Stapf, and this typification is also followed by the Index Nominum Genericorum. Although this species was included under Danthonia sect. Pentaschistis by Nees in 1841, he gave it the illegitimate name of D. trichotoma Nees, including Avena aristidoides Thunb. in synonymy. As such, the lectotypification is valid and should be followed, even if P. glandulosa would have been a more satisfactory type, as it was included at the first mention of the new taxon by Nees. Distribution and habitat. This is a typical element of the Afrotemperate flora (Linder, 1990; Galley et al., 2007), with a concentration of species in the Cape Floristic Region, but it is also common in the Afromontane and in particular the Afroalpine region, reaching from Mt. Cameroon in the west to the Ethiopian uplands in the northeast, and Amsterdam Island in the Indian Ocean. Several species have been introduced into Australia, where they have become weedy (Linder, 2005). This is a distribution pattern that is common for several Cape floral elements, such as the orchid genus Disa and several genera of the Iridaceae, such as Moraea and Gladiolus L. Most species of Pentameris are found in the Cape Floristic Region, where they form an important floristic component, often dominant in the first years after fire (Taylor, 1978). In the more arid Namaqualand, several annual species are found. In tropical Africa, the genus contributes substantially to the Afroalpine grassland (Hedberg, 1964; Lind & Morrison, 1974), and in the subalpine and montane zone from the Drakensberg (Mucina et al., 2006) to Mt. Cameroon, where P. pictigluma (Steud.) Galley & H. P. Linder dominates the grassland on the mountain summit (Maitland, 1932). Nomenclatural note on Achneria. Munro in Harvey (1968) transferred the African species that Nees (1841) had grouped under Eriachne R. Br. into Achneria P. Beauv. This was based on an incorrect interpretation of Achneria and clearly did not constitute the description of a new genus. Bentham and Hooker (1883) recognized that the African species that Munro had transferred to Achneria constituted a new genus, so they changed the concept of Achneria by reducing Achneria P. Beauv. into Eriachne and keeping the African species under Achneria sensu Munro. Bentham therefore recognized a new genus and gave it an invalid, homonymic name. Sprague (1922) correctly interpreted the situation and proposed the new name Afrachneria for Achneria Munro ex Benth. & Hook. f. Munro recognized seven species in Eriachne but did not list them. Here we select a typical species, which would have been known to Munro, as lectotype. Cytology. 2n 5 14, 24, 26, 28, 40, 42, 52, 56, 91 (Hedberg, 1957; de Wet, 1960; Tateoka, 1965; Davidse et al., 1986; Davidse, 1988; Spies & du Plessis, 1988; Spies et al., 1990; du Plessis & Spies, 1992). Anatomy. The leaves are orthophyllous or sclerophyllous; adaxial ribs variously developed; adaxial sclerenchyma as small strands or massive T-shaped or inversely anchor-shaped girders associated with both 1- and 3-order vascular bundles; clear cells in the chlorenchyma absent; bulliform cells present in the adaxial grooves. Discussion. This large clade of 83 species is very distinctive by its 2-flowered spikelets, the lemma setae inserted in the sinus between the lobes and awn, and the paleae with short, weakly developed, parallel keels. Nonetheless, as is evident from the description, there is extensive variation within the genus. This includes single-flowered spikelets (but no cases of species with more than two florets per spikelet); complex multicellular glands (Linder et al., 1990; Galley & Linder, 2007); an immense variation in growth form, including geophytes, stoloniferous species, and suffrutescent species (Linder & Ellis, 1990b); annual, biennial, and perennial species; hairy ovary apices (characteristic of section Pentameris); sclerophyllous or mesophyllous leaves (Ellis & Linder, 1992; Galley & Linder, 2007); and cytological variation, including the unusual base number of x 5 7. The genus has been studied from various aspects recently, including the morphology of the glands (Linder et al., 1990), the evolutionary significance of these glands (Galley & Linder, 2007), and the variation in the leaf anatomy (Ellis & Linder, 1992). In addition, the taxonomy of all three previous genera (Davidse, 1988; Linder & Ellis, 1990a; Barker, 1993) has recently been revised. One of the more remarkable reclassifications made here is the placement of Pseudopentameris obtusifolia into Pentameris. This species has the large glumes of Pseudopentameris (albeit somewhat smaller than typical of the genus), but it also has the villous ovary characteristic of Pentameris s. str. Volume 97, Number 3 2010 IVa. Pentameris P. Beauv. sect. Pentameris. This section is readily distinguished from the other two sections by the fruit with a brittle pericarp crowned with a tuft of white hairs. Included species. The 10 species included have historically been placed in this genus. 1. Pentameris distichophylla (Lehm.) Nees, Linnaea 7: 314. 1832. 2. Pentameris glacialis N. P. Barker, Bothalia 23: 44. 1993. 3. Pentameris hirtiglumis N. P. Barker, Bothalia 23: 39. 1993. 4. Pentameris longiglumis (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 299. 1841. 4a. Pentameris longiglumis (Nees) Steud. subsp. longiglumis. 4b. Pentameris longiglumis (Nees) Steud. subsp. gymnocolea N. P. Barker, Bothalia 23: 39. 1993. 5. Pentameris macrocalycina (Steud.) Schweick., Repert. Spec. Nov. Regni Veg. 43: 91. 1938. 6. Pentameris obtusifolia (Hochst.) Schweick., Repert. Spec. Nov. Regni Veg. 43: 91. 1938. 7. Pentameris oreophila N. P. Barker, Bothalia 23: 41. 1993. 8. Pentameris swartbergensis N. P. Barker, Bothalia 23: 43. 1993. 9. Pentameris thuarii P. Beauv., Ess. Agrostogr. 92, t. 18, fig. 8. 1812. 10. Pentameris uniflora N. P. Barker, Bothalia 23: 35. 1993. Linder et al. Classification of Danthonioideae 329 Herb. 12: 95. 1990. TYPE: South Africa. Natal, Natl. Park area, Inner Tower Ravine, 17 July 1963, E. E. Esterhuysen 30242 (holotype, BOL!). 2. Pentameris tysonii (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis tysonii Stapf, Fl. Cap. (Harvey) 7: 493. 1899. TYPE: South Africa. Natal, Mt. Currie, 1 Jan. 1883, W. Tyson 1312 (holotype, K!; isotypes, BOL!, SAM!). IVc. Pentameris P. Beauv. sect. Pentaschistis (Nees) H. P. Linder & Galley, comb. nov. Basionym: Danthonia sect. Pentaschistis Nees, Fl. Afr. Austr. Ill. 280. 1841. The typification and rank of this section are discussed above under the generic synonymy. In section Pentaschistis, the fruit is a glabrous caryopsis and the plants often have multicellular glands. Included species. This section includes 72 species, previously included in the genera Pentaschistis (Linder & Ellis, 1990a) and Prionanthium (Davidse, 1988). The taxonomy follows Linder and Ellis (1990a) and Davidse (1988), where the synonyms and their typification are also indicated. All new synonyms are listed here. 1. Pentameris acinosa (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis acinosa Stapf, Fl. Cap. (Harvey) 7: 495. 1899. TYPE: South Africa. Cape Province: Appelskraal (am Ufer des Rivierzondereinde), s.d., K. L. P. Zeyher 4539 (lectotype, designated by Linder & Ellis [1990a: 99], K!; isotypes, B!, H!, P!, S!, SAM!). Danthonia crispa Nees var. trunculata Nees, Fl. Afr. Austral. Ill. 310. 1841. Pentaschistis acinosa Stapf var. trunculata (Nees) Stapf, Fl. Cap. (Harvey) 7: 496. 1899. TYPE: South Africa. s. loc., s.d., J. F. Drège 1681 (lectotype, designated here, B!; isotypes, K!, P!). IVb. Pentameris P. Beauv. sect. Dracomontanum H. P. Linder & Galley, sect. nov. TYPE: Pentameris tysonii (Stapf) Galley & H. P. Linder (; Pentaschistis tysonii Stapf). The B specimen was annotated by Nees and is therefore selected as lectotype. Haec sectio a Pentameride P. Beauv. sect. Pentameride caryopsidibus glabratis differt. 2. Pentameris airoides Nees, Sem. Hort. Bot. Vratisl. 1834. Included species. Only two species are included in section Dracomontanum. These were previously included in Pentaschistis (Gibbs Russell et al., 1990; Linder & Ellis, 1990a). 2a. Pentameris airoides Nees subsp. airoides. 1. Pentameris praecox (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis praecox H. P. Linder, Contr. Bolus 2b. Pentameris airoides Nees subsp. jugorum (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis jugorum Stapf, Fl. Cap. (Harvey) 7: 504. 1899. Pentaschistis airoides (Nees) Stapf subsp. jugorum (Stapf) H. P. Linder, Contr. Bolus Herb. 12: 48. 1990. TYPE: South 330 Annals of the Missouri Botanical Garden Africa. Cape Province: Witteberge near Aliwal North, s.d., J. F. Drège s.n. (holotype, K!). South Africa. Cape Province: s.d., C. P. Thunberg s.n. (lectotype, designated here, UPS 2577!). 3. Pentameris alticola (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis alticola H. P. Linder, Contr. Bolus Herb. 12: 79. 1990. TYPE: South Africa. Cape Province: Ceres, Milner Vlakte in Hex River Mtns., 20 Nov. 1987, H. P. Linder 4486 (holotype, BOL!; isotype, S!). There are two collections in the Thunberg Herbarium in UPS (UPS 2577 and UPS 2578). It is not clear whether they constitute two separate collections (syntypes) or duplicates of the same collection (isotypes). UPS 2577 is the more complete specimen, and so is selected as lectotype. 4. Pentameris ampla (Nees) Galley & H. P. Linder, comb. nov. Basionym: Eriachne ampla Nees, Fl. Afr. Austral. Ill. 277. 1841. Achneria ampla (Nees) T. Durand & Schinz, Consp. Fl. Afr. (T. A. Durand & H. Schinz) 5: 836. 1894. Pentaschistis ampla (Nees) McClean, S. African J. Sci. 23: 282. 1926. Afrachneria ampla (Nees) Adamson, J. S. African Bot. 5: 53. 1939. TYPE: South Africa. Cape Province: betw. Paarlberg & Du Toits Kloof, s.d., J. F. Drège 1674 (lectotype, designated by Linder & Ellis [1990a: 59], B!). Eriachne pallida Nees, Fl. Afr. Austral. Ill. 275. 1841. Achneria pallida (Nees) T. Durand & Schinz, Consp. Fl. Afr. (T. A. Durand & H. Schinz) 5: 836. 1894. TYPE: South Africa. Cape Province: Zwartkopsriver, s.d., C. F. Ecklon s.n. (holotype, B!). Eriachne aurea (Steud.) Nees var. virens Nees, Fl. Afr. Austral. Ill. 276. 1841. Achneria aurea (Steud.) T. Durand & Schinz var. virens (Nees) Stapf, Fl. Cap. (Harvey) 7: 459. 1899. TYPE: South Africa. Cape Province: Du Toits Kloof, s.d., J. F. Drège s.n. (holotype, B!; isotype, K!). 5. Pentameris andringitrensis (A. Camus) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis andringitrensis A. Camus, Bull. Soc. Bot. France 74: 689. 1927. TYPE: Madagascar. Massif d’Andringitra, s.d., J. M. H. A. Perrier de la Bathie 10832 (lectotype, designated by Linder & Ellis [1990a: 104], P!). 6. Pentameris argentea (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis argentea Stapf, Fl. Cap. (Harvey) 7: 487. 1899. TYPE: South Africa. Cape Province: Cape Peninsula, Orange Kloof, s.d., A. H. Wolley-Dod 3342 (lectotype, designated by Linder & Ellis [1990a: 68], K!; isotype, K fragm. at PRE!). 7. Pentameris aristidoides (Thunb.) Galley & H. P. Linder, comb. nov. Basionym: Avena aristidoides Thunb., Prodr. Pl. Cap. 22. 1794. Danthonia trichotoma Nees, Fl. Afr. Austral. Ill. 318. 1841, nom. superfl. Pentaschistis aristidoides (Thunb.) Stapf, Fl. Cap. (Harvey) 7: 485. 1899. TYPE: 8. Pentameris aristifolia (Schweick.) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis aristifolia Schweick., Repert. Spec. Nov. Regni Veg. 43: 89. 1938. TYPE: South Africa. Cape Province: 40 mi. SE of Williston, s.d., J. Hutchinson 981 (holotype, K!). 9. Pentameris aspera (Thunb.) Galley & H. P. Linder, comb. nov. Basionym: Holcus asper Thunb., Prodr. Pl. Cap. 20. 1794. Sorghum asperum (Thunb.) Roem. & Schult., Syst. Veg. ed. 15 (bis) (Roemer & Schultes) 2: 839. 1817. Pentaschistis aspera (Thunb.) Stapf, Fl. Cap. (Harvey) 7: 500. 1899. TYPE: [South Africa. Cape Province:] ‘‘crescit in summis lateribus montium urbis,’’ C. P. Thunberg in herb. Thunb. 23841 (holotype, UPS! [microfiche BOL!]). 10. Pentameris aurea (Steud.) Galley & H. P. Linder, comb. nov. Basionym: Aira aurea Steud., Flora 12: 470. 1829. Airopsis aurea (Steud.) Nees, Linnaea 7: 317. 1832. Eriachne aurea (Steud.) Nees, Fl. Afr. Austral. Ill. 276. 1841. Achneria aurea (Steud.) T. Durand & Schinz, Consp. Fl. Afr. (T. A. Durand & H. Schinz) 5: 836. 1894. Afrachneria aurea (Steud.) Adamson, J. S. African Bot. 5: 53. 1939. Pentaschistis aurea (Steud.) McClean, S. African J. Sci. 23: 282. 1926. TYPE: South Africa. Cape Province: Table Mtn., s.d., C. F. Ecklon 915 (holotype, P not seen; isotypes, BM!, K!). 10a. Pentameris aurea (Steud.) Galley & H. P. Linder subsp. aurea. 10b. Pentameris aurea (Steud.) Galley & H. P. Linder subsp. pilosogluma (McClean) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis pilosogluma McClean, S. African J. Sci. 23: 282. 1926. Pentaschistis aurea (Steud.) McClean subsp. pilosogluma (McClean) H. P. Linder, Contr. Bolus Herb. 12: 76. 1990, replacement name, pro Danthonia hirsuta Nees, Fl. Afr. Austral. Ill. 282. 1841. Achneria hirsuta (Nees) Stapf, Fl. Cap. (Harvey) 7: 462. 1899. TYPE: Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 331 South Africa. Cape Province: Witteberge, s.d., J. F. Drège 8116 (holotype, B!; isotype, K!). 1893, G. Volkens 1368 (holotype, B!; isotypes, BM!, E!, HBG!, K!). 11. Pentameris bachmannii (McClean) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis bachmannii McClean, S. African J. Sci. 23: 282. 1926, replacement name, pro Agrostis curvifolia Hack., Bull. Herb. Boissier 3: 384. 1895. Achneria curvifolia (Hack.) Stapf, Fl. Cap. (Harvey) 7: 458. 1899. TYPE: South Africa. Cape Province: near Hopefield, s.d., F. E. Bachmann 1017 (holotype, B!; isotype, K!). Pentaschistis trisetoides (Hochst. ex Steud.) Pilg. var. expansa Pilg., Notzibl. Bot. Gart. Berlin-Dahlem 9: 516. 1926. Pentaschistis expansa (Pilg.) C. E. Hubb., Fl. Trop. Afr. 10: 130. 1937. TYPE: Kenya. Mt. Kenya, 1921, T. C. E. Fries & R. E. Fries 1200b (holotype, B not seen; isotype, K!). Pentaschistis effusa Peter, Repert. Spec. Nov. Regni Veg. Beih. 40 (1, Anhang): 97 t. 56/1. 1930. TYPE: Tanzania. Mt. Kilimanjaro, A. Peter 46685 (holotype, B not seen). Pentaschistis meruensis C. E. Hubb., Kew Bull. 1936: 501. 1936. TYPE: Tanzania. Arusha Distr., Mt. Meru, s.d., B. D. Burtt 4062 (holotype, K!). Pentaschistis ruwenzoriensis C. E. Hubb., Kew Bull. 1936: 500. 1936. TYPE: Uganda. Toro Distr., Mt. Ruwenzori, s.d., G. Taylor 2903 (holotype, K!; isotype, BM!). Eriachne ecklonii Nees, Fl. Afr. Austral. Ill., 273. 1841. Eriachne assimilis Steud., Syn. Pl. Glumac. 1: 236. 1854, nom. superfl. Achneria ecklonii (Nees) T. Durand & Schinz, Consp. Fl. Afr. (T. A. Durand & H. Schinz) 5: 836. 1894. Achneria assimilis (Steud.) T. Durand & Schinz, Consp. Fl. Afr. (T. A. Durand & H. Schinz) 5: 836. 1894, nom. illeg. Afrachneria ecklonii (Nees) Adamson, J. S. African Bot. 5: 53. 1939. Pentaschistis ecklonii (Nees) McClean, S. African J. Sci. 23: 282. 1926. TYPE: South Africa. Cape Province: Klein Drakenstein at the Bergriver, s.d., J. F. Drège 1660 (lectotype, designated by Linder & Ellis [1990a: 52], B!; isotypes, BM!, E!, K!). The superfluous name status of Eriachne assimilis is the result of the lectotypification of E. ecklonii by Linder and Ellis (1990a); otherwise, this would be the oldest name available for this species. 12. Pentameris barbata (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 298. 1841. 12a. Pentameris barbata (Nees) Steud. subsp. barbata. 12b. Pentameris barbata (Nees) Steud. subsp. orientalis (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis barbata (Nees) H. P. Linder subsp. orientalis H. P. Linder, Contr. Bolus Herb. 12: 31. 1990. TYPE: South Africa. Cape Province: Goukamma Nature Reserve, 2 Jan. 1970, P. van der Merwe 1765 (holotype, STE!). 13. Pentameris basutorum (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis basutorum Stapf, Bull. Misc. Inform. Kew 1914: 20. 1914. TYPE: Lesotho. Leribe, s.d., A. Dieterlen 222 (holotype, K!; isotypes, BM!, P!, SAM!, STE!). 14. Pentameris borussica (K. Schum.) Galley & H. P. Linder, comb. nov. Basionym: Danthonia borussica K. Schum., Pflanzenw. Ost-Afrikas C: 109. 1895. Pentaschistis borussica (K. Schum.) Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 9: 517. 1926. TYPE: Tanzania. Mt. Kilimanjaro, 11 Jan. 15. Pentameris calcicola (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis calcicola H. P. Linder, Contr. Bolus Herb. 12: 81. 1990. TYPE: South Africa. Cape Province: Bredasdorp, farm Wydgelee, 20 Oct. 1987, H. P. Linder 4365 (holotype, BOL!; isotypes, K!, PRE!). 15a. Pentameris calcicola (H. P. Linder) Galley & H. P. Linder var. calcicola. 15b. Pentameris calcicola (H. P. Linder) Galley & H. P. Linder var. hirsuta (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis calcicola H. P. Linder var. hirsuta H. P. Linder, Contr. Bolus Herb. 12: 83. 1990. TYPE: South Africa. Cape Province: Bredasdorp, farm Wydgelee, 20 Oct. 1987, H. P. Linder 4366 (holotype, BOL!; isotype, K!). 16. Pentameris capensis (Nees) Galley & H. P. Linder, comb. nov. Basionym: Triraphis capensis Nees, Fl. Afr. Austral. Ill. 271. 1841. Danthonia radicans Steud., Syn. Pl. Glumac. 1: 243. 1854, nom. superfl. Pentaschistis capensis (Nees) Stapf, Fl. Cap. (Harvey) 7: 494. 1899. TYPE: South Africa. Cape Province: Du Toits Kloof, s.d., J. F. Drège s.n. (holotype, B!; isotypes, BM!, H!, K!, P!, S!, SAM!). 17. Pentameris capillaris (Thunb.) Galley & H. P. Linder, comb. nov. Basionym: Holcus capillaris Thunb., Prod. Pl. Cap. 20. 1794. Sorghum capillare (Thunb.) Roem. & Schult., Syst. Veg., ed. 15 bis (Roemer & Schultes) 2: 840. 1817. Achneria capillaris (Thunb.) Stapf, Hooker’s Icon. Pl. 27, t. 2604. 1899, non Achneria capillaris (R. Br.) P. Beauv., Ess. Agrostogr. 73. 1812. Pentaschistis capillaris (Thunb.) McClean, S. African 332 Annals of the Missouri Botanical Garden J. Sci. 23: 281. 1926. TYPE: [South Africa.] s. loc., s.d., C. P. Thunberg (holotype, UPS 23845!). 18. Pentameris caulescens (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis caulescens H. P. Linder, Contr. Bolus Herb. 12: 99. 1990. TYPE: South Africa. Cape Province: Ceres, Buffelshoek Peak in the Hexriver Mtns., 8 Oct. 1956, E. E. Esterhuysen 26349 (holotype, BOL!). 19. Pentameris chippindalliae (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis chippindalliae H. P. Linder, Contr. Bolus Herb. 12: 92. 1990. TYPE: South Africa. Transvaal, Dullstroom, 10 Feb. 1988, H. P. Linder 4711 (holotype, BOL!; isotypes, K!, M!, MO!, NBG!, PRE!, S!). 20. Pentameris chrysurus (K. Schum.) Galley & H. P. Linder, comb. nov. Basionym: Danthonia chrysurus K. Schum., Pflanzenw. Ost-Afrikas C: 110. 1895. Pentaschistis chrysurus (K. Schum.) Peter, Repert. Spec. Nov. Regni Veg. Beih. 40(1): 303. 1931. TYPE: Tanzania. Mt. Kilimanjaro, 14 Feb. 1894, G. Volkens 1826a (holotype, B!). 21. Pentameris cirrhulosa (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 298. 1841. 22. Pentameris clavata (Galley) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis clavata Galley, Bothalia 36: 159. 2006. TYPE: South Africa. Western Cape Province: Koue Bokkeveld S of Hex Berg, 7 Nov. 2004, C. A. Galley 567 (holotype, Z!; isotypes, BOL!, E!, G!, K!, MO!, NBG!, NSW!, NY!, PRE!, S!, UPS!, W!). 23. Pentameris colorata (Steud.) Galley & H. P. Linder, comb. nov. Basionym: Avena colorata Steud., Flora 12: 481. 1829. Pentaschistis colorata (Steud.) Stapf, Fl. Cap. (Harvey) 7: 491. 1899. TYPE: South Africa. Cape Province: Cape Town, Table Mtn., s.d., C. F. Ecklon 931 (holotype, P not seen; isotypes, B!, E!, K!, S!). 24. Pentameris curvifolia (Schrad.) Nees, Linnaea 7: 313. 1832. 25. Pentameris densifolia (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 298. 1841. 26. Pentameris dentata (L. f.) Galley & H. P. Linder, comb. nov. Basionym: Phalaris dentata L. f., Suppl. Pl. 106. 1782. Phleum dentatum (L. f.) Pers., Syn. Pl. (Persoon) 1: 79. 1805. Chilochloa dentata (L. f.) Trin., Sp. Gram. (Trinius) 168. 1824. Prionanthium rigidum Desv., Opusc. Sci. Phys. Nat. 65. 1831. Lasiochloa pectinata Trin. ex Pritz., Sp. Gram. (Trinius), corrigenda et emendanda, 1(7), tab. 73. 1836, nom. superfl. pro Phalaris dentata L. f. Prionachne dentata (L. f.) Nees, Fl. Afr. Austral. Ill. 134. 1841. Prionanthium dentatum (L. f.) Henrard, Blumea 4: 530. 1941. TYPE: [South Africa.] Cape, Bockland, 1773, C. P. Thunberg s.n. (holotype, UPS 1773!). 27. Pentameris dolichochaeta (S. M. Phillips) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis dolichochaeta S. M. Phillips, Kew Bull. 50: 615. 1995. TYPE: Ethiopia. Showa, Ancobere, 3000 m, s.d., G. Selassie 887 (holotype, ETH!, ETH photo at K!). 28. Pentameris ecklonii (Nees) Galley & H. P. Linder, comb. nov. Basionym: Prionachne ecklonii Nees, Nat. Syst. Bot. 448. 1836. Chondrolaena phalaroides Nees, Fl. Afr. Austral. Ill. 134. 1841, nom. superfl. Chondrolaena phalaroides Nees var. dentata Nees, Fl. Afr. Austral. Ill. 134. 1841, nom. illeg. Prionanthium ecklonii (Nees) Stapf, Fl. Cap. (Harvey) 7: 456. 1899. TYPE: South Africa. ‘‘ad Olifantsrivier fluviam alt. I, Clanwilliam,’’ s.d., C. F. Ecklon s.n. (lectotype, designated by Davidse [1988: 151], MO not seen; isotypes, BM, BM fragm. at PRE!, US not seen, Z!). 29. Pentameris elegans (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 298. 1841. 30. Pentameris ellisii H. P. Linder, Bothalia 40: 191. 2010. 31. Pentameris eriostoma (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 298. 1841. 32. Pentameris exserta (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis exserta H. P. Linder, Contr. Bolus Herb. 12: 92. 1990. TYPE: South Africa. Natal: Cathedral Peak Forest Reserve, Organ Pipes Pass, 4 Feb. 1988, H. P. Linder 4685 (holotype, BOL!). 33. Pentameris galpinii (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Achneria galpinii Stapf, Bull. Misc. Inform. Kew 1910: 59. 1910. Pentaschistis galpinii (Stapf) McClean, S. African J. Sci. 23: 282. 1926. TYPE: South Africa. Cape Province: Barkly East, Ben Macdhui, 11 Mar. 1904, E. E. Galpin 6915 (holotype, K!; isotypes, B!, BOL!, GRA!, SAM!). Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 333 34. Pentameris glandulosa (Schrad.) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 298. 1841. 35. Pentameris heptameris (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 298. 1841. 36. Pentameris holciformis (Nees) Galley & H. P. Linder, comb. nov. Basionym: Danthonia holciformis Nees, Fl. Afr. Austral. Ill. 326. 1841. Pentaschistis holciformis (Nees) H. P. Linder, Contr. Bolus Herb. 12: 91. 1990. TYPE: South Africa. Cape Province: Palmietriver at Grietjiesgat, s.d., C. F. Ecklon s.n. (lectotype, designated here, B!; isotype, S!). The B specimen was annotated by Nees and is selected here as lectotype. 37. Pentameris horrida (Galley) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis horrida Galley, Bothalia 36: 160. 2006. TYPE: South Africa. Western Cape: Ceres, Baviaansberg, 26 Oct. 1997, H. P. Linder 6799 (holotype, Z!; isotypes, BOL!, E!, G!, K!, MO!, NBG!, NSW!, PRE!). 38. Pentameris humbertii (A. Camus) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis humbertii A. Camus, Bull. Soc. Bot. France 70: 690. 1928. TYPE: Madagascar. Pic d’Ivohibe (Bara), 5 Nov. 1924, J.-H. Humbert 3310 (holotype, P!; isotypes, B!, K!). 39. Pentameris insularis (Hemsl.) Galley & H. P. Linder, comb. nov. Basionym: Trisetum insulare Hemsl., Rep. Voy. Challenger, Bot. 1(2): 267, t. 52. 1884. Pentaschistis insularis (Hemsl.) H. P. Linder, Contr. Bolus Herb. 12: 103. 1990. TYPE: Territory of the French Southern and Antarctic Lands. St. Paul Island, Indian Ocean, s.d., J. MacGillivray & W. G. Milne (lectotype, designated by Linder & Ellis [1990a: 104], K!). Figure 6. 40. Pentameris juncifolia (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis juncifolia Stapf, Fl. Cap. (Harvey) 7: 490. 1899. TYPE: South Africa. Riversdale division, hills near Zoetemelksrivier, s.d., W. J. Burchell 6750 (lectotype, designated by Linder & Ellis [1990a: 107], K!). 41. Pentameris lima (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 299. 1841. 42. Pentameris longipes (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis Figure 6. Pentameris insularis. Drawn by Jasmin Baumann. longipes Stapf, Fl. Cap. (Harvey) 7: 509. 1899. TYPE: South Africa. Cape Province: Albany, s.d., J. Bowie s.n. (holotype, K!). 43. Pentameris malouinensis (Steud.) Galley & H. P. Linder, comb. nov. Basionym: Eriachne malouinensis Steud., Syn. Pl. Glumac. 1: 236. 1854. Pentaschistis malouinensis (Steud.) Clayton, Kew Bull. 23: 294. 1969. TYPE: Falkland Islands, s.d., D. d’Urville s.n. (holotype, CN not seen; isotype, CN photo at K!). 334 The Falkland Islands as type locality is almost certainly an error: the species is endemic to southern Africa. 44. Pentameris microphylla (Nees) Galley & H. P. Linder, comb. nov. Basionym: Eriachne microphylla Nees, Fl. Afr. Austral. Ill. 277. 1841. Achneria microphylla (Nees) T. Durand & Schinz, Consp. Fl. Afr. (T. A. Durand & H. Schinz) 5: 836. 1894. Pentaschistis microphylla (Nees) McClean, S. African J. Sci. 23: 282. 1926. TYPE: South Africa. Cape Province: ‘‘in monte Stormberg (Terra Caffrorum) alt. 60009, Decembri in anthesi,’’ s.d., J. F. Drège 3891 (holotype, B!; isotypes, BM!, K!, SAM!). 45. Pentameris minor (Ballard & C. E. Hubb.) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis borussica (K. Schum.) Pilg. var. minor Ballard & C. E. Hubb., Bull. Misc. Inform. Kew 1930: 121. 1930. Pentaschistis minor (Ballard & C. E. Hubb.) Ballard & C. E. Hubb., Fl. Trop. Afr. (Oliver et al.) 10: 132. 1937. Pentaschistis pictigluma (Steud.) Pilg. var. minor (Ballard & C. E. Hubb.) S. M. Phillips, Proc. XIII Plen. Meet. AETFAT [Association pour l’Étude Taxonomique de la Flore d’Afrique Tropical] Zomba Malawi 371. 1994. TYPE: Tanzania. Mt. Kilimanjaro, near Peters Hut, s.d., A. D. Cotton & A. S. Hitchcock 64 (holotype, K!). 46. Pentameris montana (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis montana H. P. Linder, Contr. Bolus Herb. 12: 83. 1990. TYPE: South Africa. Cape Province: Worcester, Keeromsberg, 7 Nov. 1987, H. P. Linder 4413 (holotype, BOL!). 47. Pentameris natalensis (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis natalensis Stapf, Fl. Cap. (Harvey) 7: 493. 1899. TYPE: South Africa. Natal, Riet Vlei, s.d., J. Buchanan 283 (holotype, K!; isotypes, B!, BOL!). 48. Pentameris oreodoxa (Schweick.) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis oreodoxa Schweick., Repert. Spec. Nov. Regni Veg. 43: 90. 1938. TYPE: South Africa. Natal, Bergville, Mont aux Sources, near summit of mtn., s.d., A. J. W. Bayer & A. P. D. McClean 273 (holotype, K!). 49. Pentameris pallescens (Schrad.) Nees, Linnaea 7: 312. 1832. 50. Pentameris pallida (Thunb.) Galley & H. P. Linder, comb. nov. Basionym: Avena pallida Annals of the Missouri Botanical Garden Thunb., Prod. Pl. Cap. 22. 1794. Danthonia pallida (Thunb.) Roem. & Schult., Syst. Veg. 2: 657. 1817, nom. illeg., non Danthonia pallida R. Br., Prodr. Fl. Nov. Holland. 177. 1810. Pentaschistis pallida (Thunb.) H. P. Linder, Contr. Bolus Herb. 12: 36. 1990. TYPE: South Africa. Cape Province: Verkeerde Vlei, s.d., C. P. Thunberg (lectotype, designated by Linder & Ellis [1990a: 36], UPS 2610!). Danthonia angustifolia Nees, Fl. Afr. Austral. Ill. 302. 1841. Pentameris angustifolia (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 298. 1841. Pentaschistis angustifolia (Nees) Stapf, Fl. Cap. (Harvey) 7: 502. 1899. TYPE: South Africa. Cape Province: on fields at Zwartkopsvlei and at Adow, s.d., C. F. Ecklon 839 (lectotype, designated by Linder & Ellis [1990a: 37], B!). Pentaschistis thunbergii Stapf, Fl. Cap. (Harvey) 7: 507. 1899, non Pentaschistis thunbergii Kunth. Stapf (1899) misapplied Pentaschistis thunbergii, and its incorrect use was widespread in the literature until corrected by Linder and Ellis (1990a). 51. Pentameris patula (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 299. 1841. 52. Pentameris pholiuroides (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Prionanthium pholiuroides Stapf, Fl. Cap. (Harvey) 7: 456. 1899. Prionachne pholiuroides (Stapf) E. Phillips, Intr. S. African Grass. 6, t. 63. 1931. TYPE: South Africa. Fish Hoek valley, damp hollow, Nov. 1897, A. H. Wolley-Dod 3394 (holotype, K not seen; isotypes, BM not seen, BOL!, MO not seen, PRE!). 53. Pentameris pictigluma (Steud.) Galley & H. P. Linder, comb. nov. Basionym: Aira pictigluma Steud., Syn. Pl. Glumac. 1: 221. 1854. Danthonia anthoxanthiformis Hochst., Flora 38: 276. 1855, nom. illeg. superfl. Pentaschistis pictigluma (Steud.) Pilg., Notizbl. Bot. Gart. BerlinDahlem 9: 517. 1926. TYPE: Ethiopia. s.d., W. G. Schimper (holotype, P not seen). The species delimitation in Pentameris pictigluma is still most unsatisfactory, and a critical evaluation of the populations on the different mountains, and in different altitude zones, is needed. Until such time, the potential taxa are here recognized as varieties (thus following the treatment of Phillips [1995], except that P. minor is maintained at species level). 53a. Pentameris pictigluma (Steud.) Galley & H. P. Linder var. pictigluma. Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 53b. Pentameris pictigluma (Steud.) Galley & H. P. Linder var. gracilis (S. M. Phillips) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis gracilis S. M. Phillips, Kew Bull. 41: 1028. 1986. Pentaschistis pictigluma (Steud.) Pilg. var. gracilis (S. M. Phillips) S. M. Phillips, Proc. XIII Plen. Meet. AETFAT [Association pour l’Étude Taxonomique de la Flore d’Afrique Tropical] Zomba Malawi 372. 1994. TYPE: Ethiopia. Shoa Province, Entoto Hill, along a small stream, s.d., I. Friis, M. Gilbert, F. Rasmussen & K. Vollesen 1303 (holotype, K!). 58. Pentameris reflexa (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis reflexa H. P. Linder, Contr. Bolus Herb. 12: 53. 1990. TYPE: South Africa. Cape Province: Cedarberg, slopes below Middelberg at Algeria, 6 Dec. 1987, H. P. Linder 4531 (holotype, BOL!; isotypes, K!, MO!, PRE!, STE!). 53c. Pentameris pictigluma (Steud.) Galley & H. P. Linder var. mannii (Stapf ex C. E. Hubb.) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis mannii Stapf ex C. E. Hubb., Bull. Misc. Inform. Kew 1936: 501. 1936. TYPE: Cameroun. Mt. Cameroun, s.d., G. Mann 1351 (holotype, K!). 54. Pentameris pseudopallescens (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis pseudopallescens H. P. Linder, Contr. Bolus Herb. 12: 72. 1990. TYPE: South Africa. Cape Province: Ceres, Milner Vlakte, Hex River Mtns., 20 Nov. 1987, H. P. Linder 4483 (holotype, BOL!). 55. Pentameris pungens (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis pungens H. P. Linder, Contr. Bolus Herb. 12: 97. 1990. TYPE: South Africa. Cape Province: Clanwilliam Distr., Uitkyk Peak, Cedarberg, 12 Oct. 1975, E. E. Esterhuysen 34010 (holotype, BOL!; isotypes, K!, PRE!). 56. Pentameris pusilla (Nees) Galley & H. P. Linder, comb. nov. Basionym: Colpodium pusillum Nees, Fl. Afr. Austral. Ill. 149. 1841. Poagrostis pusilla (Nees) Stapf, Fl. Cap. (Harvey) 7: 760. 1900. Agrostis umbellata Trin., Graminea Agrostidea. II. Callus rotundus (Agrostea), 370. 1841, nom. illeg., non Agrostis umbellata Colla, Herb. Pedem. 6: 18. 1836. Pentaschistis pusilla (Nees) H. P. Linder, Contr. Bolus Herb. 12: 89. 1990. TYPE: South Africa. Cape Province: Table Mtn., s.d., J. F. Drège s.n. (holotype, B not seen; isotype, K!). 57. Pentameris pyrophila (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis pyrophila H. P. Linder, Contr. Bolus Herb. 12: 81. 1990. TYPE: South Africa. Cape Province: Ceres, Milner Peak, Hex River Mtns., 20 Nov. 1987, H. P. Linder 4477 (holotype, BOL!). 335 59. Pentameris rigidissima (Pilg. ex H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis rigidissima Pilg. ex H. P. Linder, Contr. Bolus Herb. 12: 85. 1990. TYPE: South Africa. Cape Province: Worcester, Milner Peak, Hex River Mtns., 18 Dec. 1948, E. E. Esterhuysen 14903 (holotype, BOL!; isotypes, NBG!, PRE!, SAM!). 60. Pentameris rosea (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis rosea H. P. Linder, Contr. Bolus Herb. 12: 70. 1990. TYPE: South Africa. Cape Province: Porterville Mtns., Groot Winterhoek Forest Reserve, Suurvlakte, 14 Oct. 1988, H. P. Linder 4777 (holotype, BOL!). 60a. Pentameris rosea (H. P. Linder) Galley & H. P. Linder subsp. rosea. 60b. Pentameris rosea (H. P. Linder) Galley & H. P. Linder subsp. purpurascens (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis rosea H. P. Linder subsp. purpurascens H. P. Linder, Contr. Bolus Herb. 12: 72. 1990. TYPE: South Africa. Cape Province: Ceres Distr., Milner Vlakte, Hexriver Mtns., 24 Oct. 1987, H. P. Linder 4403 (holotype, BOL!). 61. Pentameris rupestris (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 299. 1841. 62. Pentameris scabra (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 299. 1841. Avena papillosa Steud., Flora 12: 484. 1829, non Schrad., Gött. Gel. Anz. 3: 2075. 1821. Danthonia scabra Nees, Fl. Afr. Austral. Ill., 287. 1841, replacement name, pro Avena papillosa Steud., Flora 12: 484. 1829. Pentameris scabra (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 299. 1841. Pentaschistis zeyheri Stapf, Fl. Cap. (Harvey) 7: 497. 1899, nom. superfl. Pentaschistis papillosa (Steud.) H. P. Linder, Contr. Bolus Herb. 12: 32. 1990, nom. illeg. TYPE: South Africa. Cape Province: Cape Town, ‘‘inter saxa in summitate montis tabularis, Fl. Novbr.,’’ C. F. Ecklon 936 (types, E!, K!). Pentaschistis subulifolia Stapf, Fl. Cap. (Harvey) 7: 499. 1899. TYPE: South Africa. Cape Province: Table Mtn., s.d., MacOwan 1698 (lectotype, designated by Linder & Ellis [1990a: 37], K!; isotypes, BM!, SAM!). 336 63. Pentameris scandens (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis scandens H. P. Linder, Contr. Bolus Herb. 12: 101. 1990. TYPE: South Africa. Cape Province: Bredasdorp, Bontebok Park, 25 Aug. 1962, J. P. H. Acocks 22619 (holotype, PRE!). 64. Pentameris setifolia (Thunb.) Galley & H. P. Linder, comb. nov. Basionym: Holcus setifolius Thunb., Fl. Cap. (Thunberg, ed. 2), 1: 413. 1813. Achneria setifolia (Thunb.) Stapf, Fl. Cap. (Harvey) 7: 462. 1899. Pentaschistis setifolia (Thunb.) McClean, S. African J. Sci. 23: 282. 1926. TYPE: [South Africa.] s. loc., s.d., Thunberg s.n. in herb. C. P. Thunb. 23857 (holotype, UPS!). Danthonia mutica Nees, Fl. Afr. Austral. Ill. 281. 1841. Pentameris mutica (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 299. 1841. TYPE: South Africa. Cape Province: Los-Tafelberg, s.d., J. F. Drège 3892 (holotype, B!; isotype, B fragm. at PRE!). Danthonia porosa Nees, Fl. Afr. Austral. Ill. 283. 1841. Pentameris porosa (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 299. 1841. TYPE: South Africa. Cape Province: betw. Windvogelberg & Swartkei River, s.d., J. F. Drège 3892 (lectoype, designated by Linder & Ellis [1995: 58], B!; isotype, B fragm. at PRE!, K!). Danthonia porosa Nees var. subinermis Nees, Fl. Afr. Austral. Ill. 283. 1841. Danthonia circinnata Steud., Syn. Pl. Glumac. 1: 239. 1854. TYPE: South Africa. Cape Province: Katberg, s.d., J. F. Drège 3893 (holotype, B not seen; isotypes, BM!, K!, OXF!, PRC!, B fragm. at PRE!, TCD!). Eriachne tuberculata Nees, Fl. Afr. Austral. Ill. 274. 1841. Achneria tuberculata (Nees) T. Durand & Schinz, Consp. Fl. Afr. (T. A. Durand & H. Schinz) 5: 836. 1894. TYPE: South Africa. Cape Province: Witteberge, s.d., J. F. Drège 8096 (holotype, B!; isotypes, B fragm. at PRE!, K!). 65. Pentameris tomentella (Stapf) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis tomentella Stapf, Fl. Cap. (Harvey) 7: 502. 1899. TYPE: South Africa. Cape Province: Namaqualand, Modderfonteinsberg, s.d., J. F. Drège s.n. (holotype, K!; isotype, S!). Pentaschistis brachyanthera Stapf, Fl. Cap. (Harvey) 7: 507. 1899. TYPE: South Africa. Cape Province: Namaqualand, betw. Pedros Kloof & Lilyfontein, s.d., J. F. Drège 2580 (holotype, K!; isotypes, B!, BM!). Annals of the Missouri Botanical Garden Galley, Bothalia 36: 157. 2006. TYPE: South Africa. Western Cape Province, Ceres, Baviaansberg, 11 Nov. 2004, C. A. Galley 577 (holotype, Z!; isotypes, BOL!, K!, NBG!, PRE!). 68. Pentameris triseta (Thunb.) Galley & H. P. Linder, comb. nov. Basionym: Avena triseta Thunb., Prodr. Pl. Cap. 22. 1794. Trisetum villosum Pers., Syn. Pl. (Persoon) 1: 97. 1805, nom. superfl. Pentameris villosa (Pers.) Nees, Linnaea 7: 310. 1832, nom. illeg. Danthonia villosa (Pers.) Trin., Mém. Acad. Imp. Sci. St.Pétersbourg, Sci. Math, Seconde Pt. 33. 1836, nom. illeg. Avena capensis Spreng., Syst. Veg. 1: 333. 1825, nom. superfl. Danthonia thunbergii Kunth, Révis. Gramin. 1: 107. 1829, nom. illeg. Pentaschistis thunbergii (Kunth) Stapf, Fl. Cap. (Harvey) 7: 507. 1899, nom. illeg. Danthonia collinita Nees, Fl. Afr. Austral. Ill. 315. 1841, nom. superfl. Pentaschistis triseta (Thunb.) Stapf, Fl. Cap. (Harvey) 7: 495. 1899. TYPE: [South Africa.] s. loc., s.d., C. P. Thunberg s.n. (holotype, UPS 2632!). These names all cite Avena triseta, and so include its type. Hence they are all homotypic. However, the situation is quite complex: it seems that initially Trisetum villosum Pers. was misapplied, leading to a series of misunderstandings. 69. Pentameris trisetoides (Hochst. ex Steud.) Galley & H. P. Linder, comb. nov. Basionym: Danthonia trisetoides Hochst. ex Steud., Syn. Pl. Glumac. 1: 244. 1854. Pentaschistis trisetoides (Hochst. ex Steud.) Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 9: 516. 1926. TYPE: Ethiopia. Near Debra Eski, s.d., W. G. Schimper 109 (holotype, P!; isotype, K!). 70. Pentameris velutina (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis velutina H. P. Linder, Contr. Bolus Herb. 12: 66. 1990. TYPE: South Africa. Western Cape Prov., Porterville Mtns., on ridge on Berghof farm, 14 Oct. 1988, H. P. Linder 4791 (holotype, BOL!; isotype, E!). 66. Pentameris tortuosa (Trin.) Nees, Linnaea 7: 310, 311. 1832. 71. Pentameris veneta (H. P. Linder) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis veneta H. P. Linder, Contr. Bolus Herb. 12: 29. 1990. TYPE: South Africa. Cape Province: Cedarberg, Blaauwberg, s.d., J. F. Drège 1682b (holotype, K!). 67. Pentameris trifida (Galley) Galley & H. P. Linder, comb. nov. Basionym: Pentaschistis trifida 72. Pentameris viscidula (Nees) Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 299. 1841. For Pentaschistis tomentella, note that the types are labeled ‘‘Pentaschistis papillosa Schrad.’’ Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 337 scaberulose, rarely with tufts of long hairs; callus rounded or truncate, villous, shorter to longer than the rachilla internode; lemmas with 3 to 9 indistinct veins; lemmas dorsally pilose with the hairs generally organized into 4 vertical stripes: 2 marginal and 2 flanking the midrib; lemma lobes acute, usually shorter than the lemma body, setae well developed; lemma central awn usually present and exceeding the lemma lobes, sometimes differentiated into a flat, corkscrewed base and a straight, hairlike apical part; paleae lorate to linear, with long tufts of hair on the palea-flaps; lodicules rhomboid to rarely cuneate, generally with microhairs and bristles; ovary glabrous. Caryopsis lorate to obovate; embryo and linear hilum about 1/2 of the caryopsis length. Cytology. 2n 5 42 (Abele, 1959; Brock & Brown, 1961; Beuzenberg & Hair, 1983; Dawson, 1989; Connor & Lloyd, 2004; Murray et al., 2005). Anatomy. The leaves are sclerophyllous, expanded; adaxial ribs variously developed, usually with deep cleftlike furrows with overlapping microhairs at the base, often with dense and large tubercles on the ribs and in the grooves; adaxial sclerenchyma as massive T-shaped or inversely anchor-shaped girders associated with both 1- and 3-order vascular bundles; the abaxial epidermis often with a continuous subepidermal layer; clear cells in the chlorenchyma and bulliform cells absent. Danthonia DC. sect. Brachylepidium Pilg., Willdenowia 5: 474. 1969. TYPE: Danthonia cunninghamii Hook. f. (; Chionochloa conspicua (G. Forst.) Zotov subsp. cunninghamii (Hook. f.) Zotov). Distribution and habitat. Twenty-two of the 24 species are found in New Zealand; one occurs on Lord Howe Island, and one in Australia, where it is endemic to the Mt. Kosciuszko area. Several species are found on the various off-shore islands around New Zealand. This genus often dominates the grasslands of the mountains of South Island (Wardle, 1991). Several species are cliff specialists, and a few occur on coastal rocks. The ecology of the species has received much attention due to their susceptibility to burning and grazing and their importance as ground cover, especially in the mountains of the South Island of New Zealand (Connor, 1967). Plants forming tough, perennial tussocks or mats; culms 0.8–3 m tall. Sheaths often fragmenting horizontally; ligule ciliate; leaf blades sclerophyllous, tough, occasionally with a weftlike indumentum on the upper surface directly above the ligule, often disarticulating from the sheaths at the ligules or falling with part of the sheath, but in many cases persistent. Inflorescences 6 paniculate, open to plumose. Spikelets with more than 2 florets, all similar; glumes shorter to longer than the florets, 4– 16 mm, with 1 to 13 nerves, glabrous or micro- Discussion. Chionochloa is very distinctive, but it remains difficult to precisely define its attributes, as there are numerous exceptions. The most reliable attribute is the plant habit with tall, tough tussocks by which Chionochloa can be readily distinguished from the other genera of danthonioid grasses in New Zealand and Australia, all of which, with the exception of Austroderia, are fine-leaved and weakly to moderately caespitose, but never massive, grasses. Austroderia, while forming substantial tussocks, has inflorescences that are considerably larger and more Figure 7. Chionochloa rigida subsp. rigida. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from Chapman in CHR 217596. V. Chionochloa Zotov, New Zealand J. Bot. 1: 87. 1963. TYPE: Chionochloa rigida (Raoul) Zotov (; Danthonia rigida Raoul). Figure 7. 338 Annals of the Missouri Botanical Garden plumose than those in species of Chionochloa. The lemmas are also distinctive, with the indumentum in marginal and midrib-flanking stripes, and the central awn without a clearly developed column. The paleae always have hair-tufts along the margins. While this occurs in many other genera, it is usually uncommon. Most distinctive, though, is the leaf anatomy with the deep grooves containing overlapping microhairs at the base, and with the groove-entrances often partially occluded by the dense tubercles. The limits between the species in Chionochloa are very difficult, possibly due to hybridization (Connor, 1991). In the most recent revision, Connor (1991) recognized numerous subspecies and varieties; these may include intermediate forms, but many are geographically separated and distinct taxa. 8c. Chionochloa crassiuscula (Kirk) Zotov subsp. torta Connor, New Zealand J. Bot. 29: 237. 1991. Included species. This genus of 25 species was revised taxonomically by Connor (1991), whose taxonomy and nomenclature we follow here. 1. Chionochloa acicularis Zotov, New Zealand J. Bot. 1: 101. 1963. 2. Chionochloa antarctica (Hook. f.) Zotov, New Zealand J. Bot. 1: 99. 1963. 3. Chionochloa australis (Buchanan) Zotov, New Zealand J. Bot. 1: 103. 1963. 4. Chionochloa beddiei Zotov, New Zealand J. Bot. 1: 90. 1963. 5. Chionochloa bromoides (Hook. f.) Zotov, New Zealand J. Bot. 1: 90. 1963. 6. Chionochloa cheesemanii (Hack. ex Cheeseman) Zotov, New Zealand J. Bot. 1: 95. 1963. 7. Chionochloa conspicua (G. Forst.) Zotov, New Zealand J. Bot. 1: 92. 1963. 7a. Chionochloa conspicua (G. Forst.) Zotov subsp. conspicua. 7b. Chionochloa conspicua (G. Forst.) Zotov subsp. cunninghamii (Hook. f.) Zotov, New Zealand J. Bot. 1: 94. 1963. 8. Chionochloa crassiuscula (Kirk) Zotov, New Zealand J. Bot. 1: 103. 1963. 8a. Chionochloa crassiuscula (Kirk) Zotov subsp. crassiuscula. 8b. Chionochloa crassiuscula (Kirk) Zotov subsp. directa Connor, New Zealand J. Bot. 29: 236. 1991. 9. Chionochloa defracta Connor, New Zealand J. Bot. 25: 164. 1987. 10. Chionochloa flavescens Zotov, New Zealand J. Bot. 1: 97. 1963. 10a. Chionochloa flavescens Zotov subsp. flavescens. 10b. Chionochloa flavescens Zotov subsp. brevis Connor, New Zealand J. Bot. 29: 240. 1991. 10c. Chionochloa flavescens Zotov subsp. hirta Connor, New Zealand J. Bot. 29: 241. 1991. 10d. Chionochloa flavescens Zotov subsp. lupeola Connor, New Zealand J. Bot. 29: 242. 1991. 11. Chionochloa flavicans Zotov, New Zealand J. Bot. 1: 91. 1963. 12. Chionochloa frigida (Vickery) Conert, Senckenberg. Biol. 56: 154. 1975. 13. Chionochloa howensis S. W. L. Jacobs, Telopea 3: 281. 1988. 14. Chionochloa juncea Zotov, New Zealand J. Bot. 1: 101. 1963. 15. Chionochloa lanea Connor, New Zealand J. Bot. 25: 165. 1987. 16. Chionochloa macra Zotov, New Zealand J. Bot. 8: 91. 1970. 17. Chionochloa nivifera Connor & K. M. Lloyd, New Zealand J. Bot. 42: 531. 2004. 18. Chionochloa oreophila (Petrie) Zotov, New Zealand J. Bot. 1: 104. 1963. 19. Chionochloa ovata (Buchanan) Zotov, New Zealand J. Bot. 1: 104. 1963. 20. Chionochloa pallens Zotov, New Zealand J. Bot. 1: 99. 1963. 20a. Chionochloa pallens Zotov subsp. pallens. 20b. Chionochloa pallens Zotov subsp. cadens Connor, New Zealand J. Bot. 29: 251. 1991. 20c. Chionochloa pallens Zotov subsp. pilosa Connor, New Zealand J. Bot. 29: 252. 1991. Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 339 21. Chionochloa rigida (Raoul) Zotov, New Zealand J. Bot. 1: 96. 1963. 21a. Chionochloa rigida (Raoul) Zotov subsp. rigida. 21b. Chionochloa rigida (Raoul) Zotov subsp. amara Connor, New Zealand J. Bot. 29: 254. 1991. 22. Chionochloa rubra Zotov, New Zealand J. Bot. 1: 96. 1963. 22a. Chionochloa rubra Zotov subsp. rubra. 22a(i). Chionochloa rubra Zotov var. rubra. 22a(ii). Chionochloa rubra Zotov var. inermis Connor, New Zealand J. Bot. 29: 255. 1991. 22b. Chionochloa rubra Zotov subsp. cuprea Connor, New Zealand J. Bot. 29: 256. 1991. 22c. Chionochloa rubra Zotov subsp. occulta Connor, New Zealand J. Bot. 29: 257. 1991. 23. Chionochloa spiralis Zotov, New Zealand J. Bot. 1: 100. 1963. 24. Chionochloa teretifolia (Petrie) Zotov, New Zealand J. Bot. 1: 100. 1963. 25. Chionochloa vireta Connor, New Zealand J. Bot. 29: 261. 1991. VI. Pseudopentameris Conert, Mitt. Bot. Staatssamml. München 10: 303. 1971. TYPE: Pseudopentameris macrantha (Schrad. ex Schult.) Conert (; Danthonia macrantha Schrad. ex Schult.). Figure 8. Plants forming perennial tussocks, suffrutescences, or with single erect shoots from large underground rhizomes, culms to 0.8 m tall. Ligule ciliate; leaf blades sclerophyllous, tough, glabrous. Inflorescences 6 paniculate, open. Spikelets with 2 similar florets; glumes longer than the florets, large, 25–35 mm, with 1 to 5 nerves; callus rounded or truncate, villous, about twice as long as the rachilla internode; lemmas with 9 veins, dorsally pilose; lemma lobes acute, usually shorter than the lemma body, setae variously developed, usually seated on the inner margin of the lemma lobes; lemma central awn usually present and exceeding the lemma lobes, differentiated in a corkscrewed basal column and a straight, hairlike apical part; paleae lorate to linear, glabrous except for the scabrid keel margins; lodicules rhomboid to cuneate, sometimes with bristles; ovary glabrous. Caryopsis reticulately sculptured, embryo 1/5 and hilum 7/8 as long as fruit. Figure 8. Pseudopentameris macrantha. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from Ecklon & Zeyher 1825. Cytology. Unknown. Anatomy. The leaves are mostly sclerophyllous, expanded; adaxial ribs variously developed; adaxial sclerenchyma as caps, massive T-shaped girders or inversely anchor-shaped girders associated with both 1- and 3-order vascular bundles; bulliform cells present in the adaxial furrows. Distribution and habitat. Pseudopentameris is restricted to the Cape region of southern Africa. All three species grow on sandstone or granite substrates, where they are found in heathy fynbos vegetation, which they can dominate in the first years after fire. Pseudopentameris macrantha can form very large tussocks with aerial rhizomes that grow up with the fynbos after fire (Verboom & Linder, 1998). Discussion. Although this genus groups with Chaetobromus on molecular evidence, it is morphologically very different. Pseudopentameris shares with Pentameris 2-flowered spikelets, but differs by the large spikelets (glumes 25–35 mm long vs. 2–25 mm in Pentameris). Included species. When Conert (1971) segregated this genus from Danthonia, he placed only two species in 340 Annals of the Missouri Botanical Garden indistinct veins, dorsally pilose; lemma lobes acute, usually shorter than the lemma body, setae variously developed from the apex of the lobes; lemma central awn exceeding the lemma lobes, differentiated into a flat, corkscrewed base and a straight, hairlike apical part; paleae lorate to linear, flat; lodicules cuneate, with bristles but without microhairs; ovary glabrous. Caryopsis lorate; hilum linear. Cytology. 2n 5 12, 18, 36, 48, 52, 72 (Spies et al., 1990; Verboom & Linder, 1998). Anatomy. The leaves are orthophyllous, expanded; adaxial ribs poorly developed; sclerenchyma as small girders mostly over the 1-order adaxial vascular bundles; phloem with a thickened sheath; bulliform cells restricted to the grooves flanking the midrib. Figure 9. Chaetobromus involucratus subsp. sericeus. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from Linder s.n. it. Since then, a third new species has been recognized, and we follow the taxonomy and nomenclature proposed by Barker (1995) in his revision of the genus. 1. Pseudopentameris brachyphylla (Stapf) Conert, Mitt. Bot. Staatssamml. München 10: 304. 1971. 2. Pseudopentameris caespitosa N. P. Barker, Bothalia 25: 147. 1995. 3. Pseudopentameris macrantha (Schrad.) Conert, Mitt. Bot. Staatssamml. München 10: 304. 1971. VII. Chaetobromus Nees in J. Lindley, Nat. Syst. Bot., ed. 2, 449. 1836. Danthonia DC. sect. Chaetobromus (Nees) Steud., Syn. Pl. Glumac. 1: 238. 1854. TYPE: Chaetobromus involucratus (Schrad.) Nees (; Avena involucrata Schrad.). Figure 9. Plants forming perennial tussocks or mats, culms 0.3–0.8 m tall, sometimes with spreading underground rhizomes. Ligule ciliate; leaf blades usually expanded, 6 orthophyllous, mesic. Inflorescences paniculate, open. Spikelets with a dehiscence zone directly below the glumes marked by a tuft of hair, and with more than 2 florets, the lower floret different from the rest; glumes longer than the florets, with 5 to 11 nerves; callus rounded or truncate, villous, shorter to longer than the rachilla internode; lemmas with 7 to 9 Distribution and habitat. The genus includes a single species on the west coast of southern Africa. This species can be separated into three subspecies that show ploidy differences and occupy somewhat different habitats. Chaetobromus involucratus subsp. sericeus occurs on Quaternary coastal sands in the arid northwest, subspecies involucratus occurs on similar substrates in the more mesic southwest, and subspecies dregeanus occurs on shale- and granite-derived soils in the interior. Discussion. Although molecular data indicate that this genus is sister to Pseudopentameris, it shares only attributes of the palea and the callus with this otherwise very different genus. Chaetobromus is readily diagnosed by the tuft of hair at the base of the spikelets (hence the generic name), at a joint where the spikelets disarticulate below the glumes. Furthermore, the basal floret differs from the upper florets by its smaller size, shorter setae and awn, and only seven instead of nine veins. These attributes are both unique within the Danthonioideae. Included species. Various classifications have recognized between three (Chippendall, 1955) and one (Verboom & Linder, 1998) species in this genus. We follow the latter approach, of one species, but with three subspecies. The nomenclature follows Verboom and Linder (1998). 1. Chaetobromus involucratus (Schrad.) Nees, Fl. Afr. Austral. Ill. 344. 1841. 1a. Chaetobromus involucratus (Schrad.) Nees subsp. involucratus. 1b. Chaetobromus involucratus (Schrad.) Nees subsp. dregeanus (Nees) Verboom, Nordic J. Bot. 18: 74. 1998. Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 341 Lamprothyrsus Pilg., Bot. Jahrb. Syst. 37, Beibl. 85: 58. 1906, syn. nov. TYPE: Lamprothyrsus hieronymi (Kuntze) Pilg. (; Triraphis hieronymi Kuntze). Plants gynodioecious, or female only, or female and apomictic; forming tough, perennial tussocks; culms to 4.5 m tall. Sheaths persistent, leaving a burnt sheath after fire, shiny and white, or variously fragmenting horizontally, or becoming lacerated and curly; ligule ciliate, often with several rows of cilia; leaf blades sclerophyllous, tough, occasionally with a weftlike indumentum on the upper surface directly above the ligule, with a well-developed midrib, often resupinate shortly above the sheath, the margins and often the entire terminal half of the leaves viciously scabrid. Inflorescences paniculate, plumose. Spikelets generally with more than 2 florets, male spikelets much less hairy than female spikelets; glumes shorter to longer than the florets, 4–22 mm, with up to 5 nerves, glabrous or finely scaberulose; callus rounded or truncate, villous, shorter to longer than the rachilla internode; lemmas with 3 to 7 indistinct veins, dorsally pilose; lemma lobes usually absent, when present small and acute; setae absent to well developed, sometimes appearing as indistinct narrow lobes or teeth on the attenuated part of the central lobe; lemma central awn not differentiated into twisting column and straight apical part, occasionally with the lemma blade extending to the tip of the awn; paleae lorate to linear, occasionally with tufts of long hair on the palea margins; lodicules rhomboid to cuneate, generally with microhairs and bristles; ovary glabrous. Caryopsis lorate, elliptical, turbinate, or obovate; embryo 1/4 to 3/5 of the caryopsis length, linear hilum 1/3 to 7/10 of the caryopsis length. Cytology. 2n 5 36, 72, 108, 136 (Tsvelev, 1984; Connor & Dawson, 1993). Figure 10. Cortaderia bifida. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from Ramsaus and Arrow-Smith 592. 1c. Chaetobromus involucratus (Schrad.) Nees subsp. sericeus (Nees) Verboom, Nordic J. Bot. 18: 72. 1998. VIII. Cortaderia Stapf, Gard. Chron., ser. 3, 22: 378. 1897, nom. cons. TYPE: Cortaderia selloana (Schult. & Schult. f.) Asch. & Graebn. (; Arundo selloana Schult. & Schult. f.). Figure 10. Moorea Lem., Ill. Hort. 2 (Misc.): 15. 1855, nom rejic., non Moorea Rolfe, Gard. Chron., ser. 3, 8: 7. 1890. TYPE: Moorea argentea (Nees) Lem. (; Gynerium argenteum Nees). Anatomy. The leaves are sclerophyllous, expanded; adaxial ribs variously developed; adaxial sclerenchyma in strands, massive T-shaped girders, or as inversely anchor-shaped girders associated with both 1- and 3-order vascular bundles; the abaxial epidermis often with a continuous subepidermal layer; clear cells in the chlorenchyma usually present between the vascular bundles directly below the abaxial epidermis; adaxial bulliform cells usually absent. Distribution and habitat. Cortaderia, in its current delimitation, is restricted to South America, ranging from Tierra del Fuego to Colombia. The southernmost species is found in marshes and wetlands; the Patagonian species mostly along streams on the plains. Around the Amazon Basin the genus is found on mountains, reaching up to 4500 m. In these habitats it 342 Annals of the Missouri Botanical Garden is found in marshes or well-drained grasslands, often dominating the surrounding vegetation. 6. Cortaderia hapalotricha (Pilg.) Conert, Syst. Anat. Arundineae 102. 1961. Discussion. Cortaderia is related to Austroderia and Chimaerochloa by the gynodioecious reproductive system, as well as the 3- or 5-veined, narrow lemmas and long lemma hairs. In addition, the glumes have only a single vein. The lemma lateral lobes and their associated setae are poorly developed and in many species scarcely visible, except in the two species previously included in Lamprothyrsus, where they are conspicuous and almost as long as the awns. As a result, Lamprothyrsus species can be readily distinguished from Cortaderia species, and it is quite possible that the genus is monophyletic, even though this is not corroborated by the molecular phylogeny. However, Cortaderia is paraphyletic relative to Lamprothyrsus for both the plastid and nuclear genomes, indicating that Lamprothyrsus is best regarded as a specialized form of Cortaderia. Consequently, it is impossible to maintain the formal recognition of the two genera as separate entities. The differences among Cortaderia, Austroderia, and Chimaerochloa are minor and are detailed under those genera. We recognize two sections in Cortaderia. 7. Cortaderia hieronymi (Kuntze) N. P. Barker & H. P. Linder, comb. nov. Basionym: Triraphis hieronymi Kuntze, Revis. Gen. Pl. 3: 373. 1898. Danthonia hieronymi (Kuntze) Hack. ex Stuckert, Anal. Mus. Buenos Aires, Ser. III: 4: 484. 1906. Lamprothyrsus hieronymi (Kuntze) Pilg., Bot. Jahrb. Syst. 37, Beibl. 85: 58. 1906. TYPE: Argentina. ‘‘Cordoba, pr. urbem,’’ 6 Nov. 1881, Hieronymus s.n. (holotype, B!; isotype, K!). VIIIa. Cortaderia Stapf sect. Cortaderia. Cortaderia Stapf sect. Mutica Conert, Syst. Anat. Arundineae 114. 1961. TYPE: Cortaderia modesta (Döll) Hack. ex Dusén (; Gynerium modestum Döll). Cortaderia Stapf sect. Bifida Conert, Syst. Anat. Arundineae 95. 1961. TYPE: Cortaderia aristata Pilg. (5 Cortaderia bifida Pilg.). 8. Cortaderia jubata (Lemoine) Stapf, Bot. Mag. t. 7607. 1898. 9. Cortaderia modesta (Döll) Hack. ex Dusén, Ark. Bot. 9(5): 4. 1909. 10. Cortaderia nitida (Kunth) Pilg., Bot. Jahrb. Syst. 37: 374. 1906. 11. Cortaderia peruviana (Hitchc.) N. P. Barker & H. P. Linder, comb. nov. Basionym: Lamprothyrsus peruvianus Hitchc., Proc. Biol. Soc. Wash. 36: 195. 1923. TYPE: Peru. Yanahuanca, 16–22 June 1922, J. F. Macbride & W. Featherstone 1205 (type, K not seen). 12. Cortaderia planifolia Swallen, Contr. U.S. Natl. Herb. 29: 253. 1949. 13. Cortaderia pungens Swallen, Contr. U.S. Natl. Herb. 29: 251. 1949. 14. Cortaderia roraimensis (N. E. Br.) Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 112. 1914. Included species. Although several species of Cortaderia are invasive weeds, there is no recent species-level revision available for the genus. Moreover, the delimitations of some species may be questioned. Here, we follow the currently most widely used classification (Connor & Edgar, 1974; Connor, 1983), recognizing 19 species, while fully aware that this could be changed by a critical revision. 15. Cortaderia rudiuscula Stapf, Gard. Chron., ser. 3, 22: 396. 1897. 1. Cortaderia araucana Stapf, Gard. Chron., ser. 3, 22: 396. 1897. 18. Cortaderia speciosa (Nees) Stapf, Gard. Chron., ser. 3, 22: 396. 1897. 2. Cortaderia atacamensis (Phil.) Pilg., Bot. Jahrb. Syst. 37: 374. 1906. 19. Cortaderia vaginata Swallen, Sellowia 7: 9. 1956. 3. Cortaderia bifida Pilg., Bot. Jahrb. Syst. 37: 374. 1906. 4. Cortaderia boliviensis Lyle, Novon 6: 72. 1996. VIIIb. Cortaderia Stapf sect. Monoaristata Conert, Syst. Anat. Arundineae 117. 1961. TYPE: Cortaderia pilosa (d’Urv.) Hack. ex Dusén (; Arundo pilosa d’Urv.). Figure 11. 5. Cortaderia columbiana (Pilg.) Pilg., Bot. Jahrb. Syst. 37, Beibl. 85: 65. 1906. The single species in section Monoaristata is morphologically difficult to distinguish from those in 16. Cortaderia selloana (Schult. & Schult. f.) Asch. & Graebn., Syn. Mitteleur. Fl. (Ascherson & Graebner) 2: 325. 1900. 17. Cortaderia sericantha (Steud.) Hitchc., Contr. U.S. Natl. Herb. 24: 348. 1927. Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 343 Figure 11. Cortaderia pilosa. —A. Spikelet. —B. Lemma back. —C. Palea. —D. Gynoecium with filaments. —E. Lodicules. Drawn by Jasmin Baumann from Moore 1697. section Cortaderia, but in general the old leaf sheaths remain intact, while in section Cortaderia they usually fragment or become lacerated. There are some leaf anatomical differences, and in particular the presence of bulliform cells in the more distal adaxial grooves may be absent from Cortaderia. 1. Cortaderia pilosa (d’Urv.) Hack. ex Dusén, Svenska Exped. Magellansländerna (1895– 1897) 3, pt. 5: 222. 1900. 1a. Cortaderia pilosa (d’Urv.) Hack. var. pilosa. 1b. Cortaderia pilosa (d’Urv.) Hack. var. minima (Conert) Nicora, Darwiniana 18: 80. 1973. IX. Austroderia N. P. Barker & H. P. Linder, gen. nov. TYPE: Austroderia richardii (Endl.) N. P. Barker & H. P. Linder (; Arundo richardii Endl.). Hoc genus apparatu reproductivo gynodioecio, glumis 1nerviis et lobis lemmatum plerumque nullis Cortaderiae Stapf primo aspectu maxime simile, sed ab ea costis aliquot per folium differt. Plants gynodioecious, forming tough, perennial tussocks, in one species stoloniferous; culms 0.5–6 m tall. Sheaths persistent, shiny and white; ligule ciliate, often with several rows of cilia; leaf blades sclerophyllous, tough, usually with a weftlike indumentum on the upper surface directly above the ligule, with several prominent, midriblike veins, the margins and often the entire terminal half of the leaves viciously scabrid. Inflorescences paniculate, plumose. Spikelets generally with more than 2 florets, male spikelets much less hairy than female spikelets; glumes at least as long as the florets, 11.5–40 mm, with 1 nerve, glabrous or finely scaberulose; callus rounded or truncate, villous, shorter to longer than the rachilla internode; lemmas 3-veined, dorsally pilose; lemma lobes usually absent, when present small and acute, 344 Annals of the Missouri Botanical Garden setae absent to well developed; lemma central awn not differentiated into twisting column and straight apical part; paleae lorate to linear, occasionally with tufts of long hair on the palea margins; lodicules rhomboid to cuneate, generally with microhairs and bristles; ovary glabrous. Caryopsis lorate, elliptical, turbinate, or obovate; embryo and linear hilum ca. 1/2 as long as the caryopsis. under the generic name Cortaderia (Edgar & Connor, 2000). Cytology. 2n 5 90 (Hair & Beuzenberg, 1966; Murray et al., 2005). Anatomy. The leaves are sclerophyllous, expanded; adaxial ribs variously developed; with 1-, 2-, and 3-order vascular bundles; adaxial sclerenchyma as strands, massive T-shaped girders, or inversely anchor-shaped girders associated with all vascular bundles; the abaxial epidermis often with a continuous subepidermal layer; adaxial bulliform cells usually absent. Distribution and habitat. Austroderia is restricted to New Zealand, where it occupies diverse habitats: along streams, on beaches, and on coastal cliffs. Most of these habitats seem to be exposed to regular, and possibly quite massive, disturbance. Discussion. The genus Austroderia is related to Cortaderia and Chimaerochloa by the gynodioecious reproductive system, as well as the 3-veined narrow lemmas and long lemma hairs, and awns without twisted columns. In addition, the glumes have only a single vein. The plants are massive, and the inflorescences are huge and plumose. In particular, it is difficult to differentiate Austroderia from Cortaderia, and there seem to be no clear morphological differences in the spikelets. Geographically, Cortaderia is found in South America and Austroderia in New Zealand. However, the most striking differences are in the leaf anatomy. The leaf blades of Austroderia have several prominently sclerified veins in addition to the midrib, compared to Cortaderia, where only the midrib is strongly sclerified. The sheaths of Austroderia are conspicuously waxy and remain intact, whereas Cortaderia has sheaths that lack wax and generally fragment, longitudinally and transversely, with age. There are clearly three types of vascular bundles, instead of just two as in Cortaderia. The leaves have islands of clear cells in the mesophyll, while in Cortaderia there are usually zones of clear cells directly below the abaxial epidermis, often between the veins or ribs. Etymology. The name Austroderia is derived from the Latin ‘‘australis,’’ meaning ‘‘southern,’’ and from the genus name Cortaderia. Included species. This small genus includes only five species, revised in the Flora of New Zealand 1. Austroderia fulvida (Buchanan) N. P. Barker & H. P. Linder, comb. nov. Basionym: Arundo fulvida Buchanan, Trans. & Proc. New Zealand Inst. 6: 242. 1874. Arundo conspicua G. Forst. var. fulvida (Buchanan) Kirk, Trans. & Proc. New Zealand Inst. 10: app. xliii. 1879. Cortaderia fulvida (Buchanan) Zotov, New Zealand J. Bot. 1: 84. 1963. TYPE: New Zealand. Wellington, s.d., J. Buchanan s.n. (holotype, WELT not seen). 2. Austroderia richardii (Endl.) N. P. Barker & H. P. Linder, comb. nov. Basionym: Arundo richardii Endl., Ann. Wiener Mus. Naturgesch. 1: 158. 1836. Replaced synonym: Arundo australis A. Rich., Voy. L’Astrolabe 121. 1832, non Arundo australis Cav., Anales Hist. Nat. 1: 100. 1799. Arundo kakao Steud., Syn. Pl. Glumac. 1: 194. 1854, nom. illeg. Cortaderia richardii (Endl.) Zotov, New Zealand J. Bot. 1: 84. 1963. TYPE: New Zealand. Havre de l’Astrolabe, Nelle Zelande, s.d., Herb. Rich. no. 29 (holotype, P not seen; isotype, CHR 236584 not seen). 3. Austroderia splendens (Connor) N. P. Barker & H. P. Linder, comb. nov. Basionym: Cortaderia splendens Connor, New Zealand J. Bot. 9: 519. 1971. TYPE: New Zealand. Ruapuke Beach, 20 Dec. 1967, Bell s.n. (holotype, CHR 184354 not seen). 4. Austroderia toetoe (Zotov) N. P. Barker & H. P. Linder, comb. nov. Basionym: Cortaderia toetoe Zotov, New Zealand J. Bot. 1: 85. 1963. TYPE: New Zealand. Wainui-o-mata Valley, s.d., V. D. Zotov s.n. (holotype, CHR 95457 not seen). 5. Austroderia turbaria (Connor) N. P. Barker & H. P. Linder, comb. nov. Basionym: Cortaderia turbaria Connor, New Zealand J. Bot. 25: 167. 1987. TYPE: New Zealand. Chatham Island, Rakeinui, s.d., D. R. Given 13899 (holotype, CHR 417471 not seen). X. Plinthanthesis Steud., Syn. Pl. Glumac. 1: 14. 1853. TYPE: Plinthanthesis urvillei Steud. (lectotype, designated by Blake [1972: 3]). Figure 12. Danthonia sect. Micrathera Benth., Fl. Austral. 7: 590. 1878. Blakeochloa Veldkamp, Taxon 30: 478. 1981. TYPE: Danthonia paradoxa R. Br. Plants forming perennial tussocks; culms 0.2–1 m tall. Sheaths persistent, shiny and white; ligule ciliate; leaf blades orthophyllous, persistent on the sheaths. Inflorescences paniculate, open. Spikelets generally Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 345 inversely anchor-shaped girders, associated with all vascular bundles; the phloem with a thickened sheath; islands of clear cells in the chlorenchyma 6 circular; adaxial bulliform cells sometimes present. Distribution and habitat. These three species are restricted to the coastal plains and sandstones of the Australian east coast, where they occur in heathland, usually on oligotrophic soils. Discussion. Plinthanthesis is closely related to Notochloe by the empty cell spaces in chlorenchyma, a short callus with a horizontal joint to the rachilla, very short awns, and a turbinate caryopsis. However, Plinthanthesis is very distinct in appearance, with much shorter spikelets that have at most four florets (with at least seven in Notochloe) and lemmas that are lobed and awned, rather than finely tridentate. Characteristic in Plinthanthesis, too, is the feltlike short indumentum on the lower half of the backs of the lemmas and paleae. Included species. Three species are included in this genus; these were revised for the floras of New South Wales (Jacobs, 1994) and Australia (Linder, 2005). Figure 12. Plinthanthesis paradoxa. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from Pirie 320. with 2 to 4 florets; glumes shorter to longer than the florets, 5–9.5 mm, with 3 to 7 nerves; callus rounded or truncate, villous, less than 1/2 the length of the rachilla internode; lemmas 9-veined, dorsally with a felt of short hair in the lower half, apices tridentate to lobed, then lobes shorter than the lemma body, often fused to the central awn, setae absent; lemma central awn varying from very short (central point of tridentate lemma), generally poorly developed with a twisting base, to exceeding the lemma lobes; paleae lorate to linear, pubescent to villous on the backs in the lower half; lodicules cuneate, glabrous; ovary glabrous. Caryopsis turbinate, embryo 1/3 and the linear hilum ca. 1/2 as long as the caryopsis. Nomenclatural note. Veldkamp (1980, 1981) proposed Plinthanthesis tenuior Steud. as lectotype, but this was rejected as superfluous to Blake’s lectotypification by Connor and Edgar (1981) and later by Jacobs (1982). Cytology. Unknown. Anatomy. The leaves are orthophyllous, expanded; adaxial ribs convex, poorly developed; adaxial sclerenchyma as strands, massive T-shaped girders, or 1. Plinthanthesis paradoxa (R. Br.) S. T. Blake, Contr. Queensland Herb. 14: 3. 1972. 2. Plinthanthesis rodwayi (C. E. Hubb.) S. T. Blake, Contr. Queensland Herb. 14: 3. 1972. 3. Plinthanthesis urvillei Steud., Syn. Pl. Glumac. 1: 14. 1853. XI. Notochloe Domin, Repert. Spec. Nov. Regni Veg. 10: 117. 1911. TYPE: Notochloe microdon (Benth.) Domin (; Triraphis microdon Benth.). Figure 13. Plants forming perennial tussocks with diverging straight culms, 0.3–1 m tall. Sheaths persistent, shiny and white; ligule ciliate; leaf blades orthophyllous, persistent on the sheaths. Inflorescences paniculate, open. Spikelets elongated, generally with 7 to 9 florets; glumes shorter than the florets, scarcely overtopping the basal lemma, with 3 nerves, 4.5– 5.5 mm; callus rounded or truncate, shortly villous, less than 1/2 the length of the rachilla internode; lemmas 7-veined, glabrous, apically tridentate, thus without lobes or central awn; paleae linear, glabrous except for the scabrid keels; lodicules 3-lobed, shortly ciliate along the upper margin, without microhairs; ovary glabrous. Caryopsis turbinate, the embryo 1/3 and the linear hilum ca. 1/2 as long as the caryopsis. 346 Annals of the Missouri Botanical Garden Discussion. Notochloe is related to Plinthanthesis by the empty cell spaces in chlorenchyma, a short callus with a horizontal joint to the rachilla, very short awns, and a turbinate caryopsis. The empty spaces in the chlorenchyma are unusual for Danthonioideae, where the chlorenchyma is generally quite compact. These cavities seem to form from large, empty cells that then disintegrate, leaving rather ragged cavities. In its appearance, Notochloe is very different from Plinthanthesis, with much longer spikelets that have at least seven florets, and lemmas that are finely tridentate, rather than lobed and awned. Unusual for the subfamily, the lemmas are glabrous in Notochloe. The diaeresis use in Notochloë, which implies that the two vowels are pronounced separately, and which was used in the Flora of Australia (Linder, 2005), is explicitly permissable under Article 60.6 (McNeill et al., 2006). However, the original spelling was without the diaeresis, as ‘‘Notochloe,’’ and this is to be retained (Rule 60.1) unless there is an orthographic error. The permissable retention of the diaeresis under Article 60.6 most likely does not constitute a correctable error. Included species. The genus is monotypic; the species is described in the floras of New South Wales (Jacobs, 1994) and Australia (Linder, 2005). 1. Notochloe microdon (Benth.) Domin, Repert. Spec. Nov. Regni Veg. 10: 117. 1911. XII. Chimaerochloa H. P. Linder, gen. nov. TYPE: Chimaerochloa archboldii (Hitchc.) Pirie & H. P. Linder (; Danthonia archboldii Hitchc.). Figure 14. Genus novum quod a Cortaderia Stapf lobis lemmatum bene evolutis, a Danthonia DC. lemmatibus 3-nerviis recedit. Figure 13. Notochloe microdon. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from Pirie 326. Cytology. Unknown. Anatomy. The leaves are orthophyllous, expanded, not ribbed adaxially; midrib flanked by adaxial and abaxial grooves with adaxial bulliform cells; vascular bundles adaxially with narrow girders; phloem with a thickened sheath; mesophyll with islands of colorless cells between the vascular bundles. Distribution and habitat. This species is restricted to the sandstones of the Blue Mountains inland from Sydney, where it occurs along the banks of streams. Plants forming tough, perennial tussocks, gynodioecious; culms 0.25–1.4 m tall. Sheaths brown and persistent; leaf blade abscissing from the sheath; ligule ciliate; leaf blades sclerophyllous, tough, expanded, glabrous. Inflorescences paniculate, open to lobed. Spikelets with 2 to 7 florets, all similar; glumes shorter than the florets, 6–9 mm, with 1 to 3 nerves; central nerve much better developed than the lateral nerves; callus rounded or truncate, villous, longer than the rachilla internode; lemmas with 3 veins, dorsally pilose; lemma lobes acute, shorter than the lemma body, closely adjacent to the central awn, extended into short terminal setae; lemma central awn exceeding the lemma lobes, weakly differentiated into a flat, corkscrewed base and a straight, hairlike apical part; paleae lorate to linear, margins glabrous; lodicules rhomboid, with microhairs and bristles; Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 347 more clearly differentiated into a column and limb. According to the molecular phylogeny, the closest relationship is to Danthonia, which has a dramatically different lemma construction and indumentum. Etymology. The species takes on the appearance of different genera, depending on which character set is investigated. Thus, it can be regarded as a grass that changes its appearance, a chimaera. Included species. Only a single species is included in this genus. It is also treated in this sense in the Alpine Flora of New Guinea (Veldkamp, 1979). However, it is possible that the synonymized Danthonia macgregorii Jansen should also be separated at specific level. 1. Chimaerochloa archboldii (Hitchc.) Pirie & H. P. Linder, comb. nov. Basionym: Danthonia archboldii Hitchc., Brittonia 2: 114. 1936. Cortaderia archboldii (Hitchc.) Connor & Edgar, Taxon 23: 596. 1974. Chionochloa archboldii (Hitchc.) Conert, Senckenberg. Biol. 56: 156. 1975. TYPE: New Guinea. Central Division: Wharton Range, Murray Pass, 2800 m, 12 June 1933, L. J. Brass 4194 (holotype, US not seen). Figure 14. Chimaerochloa archboldii. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from Robbins 628. ovary glabrous. Caryopsis lorate, embryo about 1/2 and the linear hilum about 1/3 of the caryopsis length. Cytology. 2n 5 72 (Borgmann, 1964). Anatomy. The leaves are sclerophyllous, expanded; adaxial ribs massive, separated by deep cleftlike furrows; adaxial sclerenchyma as massive T-shaped or inversely anchor-shaped girders associated with all vascular bundles; midrib flanked adaxially by bulliform cells. Distribution and habitat. This genus is restricted to the mountains of New Guinea, where it is a common element in the alpine grassland. Discussion. This peculiar genus is linked to the Danthonia clade by the possession of a gynodioecious sexual system. The disarticulating leaves link this genus to Chionochloa, but it can be separated by the glabrous palea margins and by the generally pilose lemma backs. It can be separated from Cortaderia by the relatively well-developed lemma lobes with short setae (in Cortaderia the setae are generally better developed than the lemma lobes) and by the central awn Danthonia macgregorii Jansen, Reinwardtia 2: 262, fig. 6. 1953. TYPE: British New Guinea. Upper regions of Owen Stanley Range, 1889, W. MacGregor s.n. (holotype, MEL!; isotypes, BM not seen, C not seen, L not seen). XIII. Danthonia DC., Fl. Franc. (DC. & Lamarck), ed. 3, 3: 32. 1805, nom. cons. Merathrepta Raf., Bull. Bot. (Geneva) 1: 221. 1830, nom. superfl. TYPE: Danthonia spicata (L.) P. Beauv. ex Roem. & Schult. (; Avena spicata L.), typ. cons. Figure 15. Sieglingia Bernh., Syst. Verz. (Bernhardi) 20, 44. 1800. Brachatera Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2: 189. 1810, nom. superfl. Wilibald-schmidtia Conrad, Pfl. Gebirgsart. Marienbad 38. 1837, nom. superfl. Brachyanthera Kunze in Post, Lexicon 77. 1903, nom. superfl. TYPE: Sieglingia decumbens (L.) Bernh. (; Festuca decumbens L.), nom. rejic. Plants perennial, caespitose, culms 0.1–1.2 m tall, often with cleistogenes in the axils of the upper leaves. Sheaths white or brown, persistent or decaying, leaf blades persistent; ligule ciliate; leaf blades orthophyllous, expanded, glabrous or villous, rarely with a web of hair adaxially above the ligule. Inflorescences racemose to paniculate, open to contracted. Spikelets with 2 to 10 similar florets; glumes shorter to longer than the florets, 7–30 mm, with 3 to 9 nerves; callus rounded or truncate, villous, shorter to much longer than the rachilla internode; lemmas with 5 to 9 veins, dorsally pilose or 348 Annals of the Missouri Botanical Garden Figure 15. Danthonia intermedia subsp. intermedia. —A. Spikelet. —B. Lemma back. —C. Palea. —D. Anthers. Drawn by Jasmin Baumann from Elmer 1662. indumentum restricted to the lemma margins, apically tridentate to more commonly lobed; lemma lobes acute, shorter to longer than the lemma body, usually extended into terminal setae; lemma central awn exceeding the lemma lobes, differentiated into a flat, corkscrewed base and a straight, hairlike apical part; paleae lorate to linear, keels sinuose, scabrid, rarely villous between the keels, margins always glabrous; lodicules cuneate, with bristles; ovary glabrous. Caryopsis lorate, elliptical to obovate; embryo ca. 1/3 to 2/3 and the linear to elliptical hilum ca. 1/10 to 3/4 of the caryopsis length. Cytology. 2n 5 18, 24, 36, 48, 72, 98 (Stebbins & Love, 1941; de Wet, 1953, 1954; Bowden, 1960; Löve & Löve, 1961; Bowden & Senn, 1962; Packer, 1964; Quinn & Fairbrothers, 1971; Davidse & Pohl, 1972; Baeza, 1996b). Anatomy. The leaves are orthophyllous or rarely sclerophyllous, expanded; adaxial ribs usually poorly developed; adaxial sclerenchyma as small sclerenchyma strands, or with T-shaped or inversely anchorshaped girders, associated with all vascular bundles; bulliform cells usually present. Distribution and habitat. Danthonia is widespread in the temperate parts of South and North America, with two species present in Europe. In North America and Europe, the species are associated with mesic meadows, open woodlands, coastal meadows, and marshes. Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 349 Discussion. The genus Danthonia is related to Austroderia, Chimaerochloa, Plinthanthesis, and Notochloe, but it is not evident what morphological attributes characterize this relationship. Danthonia is distinct by being basically hexaploid, and by many of the species having cleistogenes in upper leaf axils. It differs from Plinthanthesis and Notochloe by the longer awns and well-developed lemma lobes, and from Austroderia by the many-veined lemmas and distinct lemma lobes. Compared to the rest of the Danthonia clade, Danthonia has relatively fat spikelets, i.e., spikelets that are rather wide in comparison to their length. 10. Danthonia cirrata Hack. & Arechav., Anales Mus. Nac. Montevideo 1: 367, t. 40. 1896. 11. Danthonia compressa Austin, Rep. (Annual) New York State Mus. Nat. Hist. 22: 54(–55). 1869. 12. Danthonia decumbens (L.) DC., Fl. Franc. (DC. & Lamarck), ed. 3, 3: 33. 1805. 13. Danthonia domingensis Hack. & Pilg., Symb. Antill. (Urban) 6: 1. 1909. 13a. Danthonia domingensis Hack. & Pilg. subsp. domingensis. Included species. There is no global revision of the 25 included species available. The North American species have been treated in the Flora of North America (Darbyshire, 2003), but the South and Central American species have not benefited from a careful critical treatment. 13b. Danthonia domingensis Hack. & Pilg. subsp. obtorta (Chase) Conert, Senckenberg. Biol. 56: 301. 1975. 1. Danthonia alpina Vest, Flora 4: 145. 1821. 14. Danthonia holm-nielsenii Laegaard, Fl. Ecuador 57: 17. 1997. 2. Danthonia annableae P. M. Peterson & Rugolo, Madroño 40: 71. 1993. 3. Danthonia araucana Phil., Anales Univ. Chile 94: 31. 1896. 4. Danthonia boliviensis Renvoize, Gram. Bolivia 260. 1998. 5. Danthonia breviseta Hack., Oesterr. Bot. Z. 52: 192. 1902. 6. Danthonia californica Bol., Proc. Calif. Acad. Sci. 2: 182. 1863. 13c. Danthonia domingensis Hack. & Pilg. subsp. shrevei (Britton) Conert, Senckenberg. Biol. 56: 301. 1975. 15. Danthonia intermedia Vasey, Bull. Torrey Bot. Club 10: 52. 1883. 15a. Danthonia intermedia Vasey subsp. intermedia. 15b. Danthonia intermedia Vasey subsp. riabuschinskii (Kom.) Tzvelev, Zlaki SSSR 610. 1976. 16. Danthonia malacantha (Steud.) Pilg., Notizbl. Bot. Gart. Berlin 10: 759. 1929. 17. Danthonia melanathera (Hack.) Bernardello, Kurtziana 10: 249. 1977. 6a. Danthonia californica Bol. var. californica. 6b. Danthonia californica Bol. var. americana (Scribn.) Hitchc., Proc. Biol. Soc. Wash. 41: 160. 1928. 7. Danthonia chaseana Conert, Senckenberg. Biol. 54: 308, fig. 5. 1975. 8. Danthonia chiapasensis Davidse, Novon 2(2): 100. 1992. 9. Danthonia chilensis E. Desv., Fl. Chil. (Gay) 6: 360. 1854. 9a. Danthonia chilensis E. Desv. var. chilensis. 9b. Danthonia chilensis E. Desv. var. aureofulva (E. Desv.) C. M. Baeza, Sendtnera 3: 32. 1996. 9c. Danthonia chilensis E. Desv. var. glabriflora Nicora, Darwiniana 18: 82. 1973. 18. Danthonia montevidensis Hack. & Arechav., Anales Mus. Nac. Montevideo 1: 369. 1896. 19. Danthonia parryi Scribn., Bot. Gaz. 21: 133. 1896. 20. Danthonia rhizomata Swallen, Comun. Bot. Mus. Hist. Nat. Montevideo 39: 2. 1961. 21. Danthonia rugoloana Sulekic, Darwiniana 37: 341. 1999. 22. Danthonia secundiflora J. Presl & C. Presl, Reliq. Haenk. 1: 255. 1830. 22a. Danthonia secundiflora J. Presl & C. Presl subsp. secundiflora. 22b. Danthonia secundiflora J. Presl & C. Presl subsp. charruana (Swallen) Roseng., B. R. Arrill. & Izag., Gram. Urug. 55. 1970. 350 Annals of the Missouri Botanical Garden Figure 16. Tenaxia stricta. —A. Spikelet. —B. Lemma back. —C. Palea. —D. Lodicules. Drawn by Jasmin Baumann from Bachmann 2256. 22c. Danthonia secundiflora J. Presl & C. Presl subsp. mattheii C. M. Baeza, Sendtnera 3: 55. 1996. 25. Danthonia unispicata (Thurb.) Munro, Cat. Canad. Pl. Endogens 2: 215. 1888. 23. Danthonia sericea Nutt., Gen. N. Amer. Pl. (Nuttall) 1: 71. 1818. XIV. Tenaxia N. P. Barker & H. P. Linder, gen. nov. TYPE: Tenaxia stricta (Schrad.) N. P. Barker & H. P. Linder (; Danthonia stricta Schrad.). Figure 16. 24. Danthonia spicata (L.) P. Beauv. ex Roem. & Schult., Syst. Veg., ed. 15 bis (Roemer & Schultes), 2: 690. 1817. Hoc genus Schismo P. Beauv. maxime simile, sed ab eo habitu semper perenni et lemmatibus semper lobatis arista centrali lobos excedente distinguitur. Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae Plants wiry, perennial tussocks without stolons, culms 0.12–0.9 m tall. Basal sheaths shiny and persistent; ligule ciliate; leaf blades occasionally with a web of interlocking hairs adaxially above the ligule, often rolled or setaceous, rarely expanded. Inflorescences varying from strictly racemose to paniculate, in shape linear to open and expanded, secund, or round. Spikelets with 2 to 7 florets; glumes usually at least as long as the florets, 7–25 mm, with 1 to 11 veins, glabrous or scaberulose on the keels; callus rounded, villous, shorter to longer than the rachilla internode; lemmas 7- to 9-veined, indumentum variously organized, from pilose to tufted, tufts usually 6 marginal, from 1 to several; lemma lobes at most as long as the lemmas, acute, setae when present either terminal or from the inner lobe margins; lemma central awn geniculate, the column twisted, the apical portion straight, much longer than the lemma lobes; paleae lorate, narrowly obovate, or elliptical, 6 glabrous except for the scabrid keel margins; lodicules cuneate to more usually rhomboid, with bristles as well as microhairs; ovary glabrous. Fruit a nutlet or caryopsis, lorate; embryo between 1/3 and 1/2 and the linear, elliptical, or punctate hilum 1/5 to 4/5 of caryopsis length. the lobes are extended into acuminate, setalike apices. These setae are consequently usually poorly differentiated from the lemma lobe and shorter than the lobes (except in T. disticha (Nees) N. P. Barker & H. P. Linder). The inflorescences are more or less contracted, sometimes even spikelike, or with the lateral branches much reduced; the glumes have several veins. In two species (T. disticha and T. stricta), the setae originate in the lemma sinus, as is characteristic in Pentameris, but these two species can be separated from Pentameris as they have more than two flowers per spikelet. Finally, in several species the leaves are asymmetrical, with more veins in one half than the other. This strange attribute is also found in Merxmuellera s. str. and Capeochloa. However, none of these characters are unique to Tenaxia. Cytology. 2n 5 12, 36, 56 (De Wet, 1954; Spies & du Plessis, 1988). Anatomy. Leaves sclerophyllous, expanded or setaceous; adaxial ribs variously developed; adaxial sclerenchyma as strands, T-shaped girders, or inversely anchor-shaped girders associated with both 1and 3-order vascular bundles; abaxial sclerenchyma as girders or rarely linked to form a continuous subepidermal layer; leaves in several species asymmetrical, with more veins in 1/2 than the other; bulliform cells sometimes present. Discussion. The new genus Tenaxia is associated with the Rytidosperma clade, in which it is grouped by both the molecular plastid and nuclear partitions. The only morphological character linking these species to the Rytidosperma clade is the generally short, punctate hilum on the caryopsis. The genus can be diagnosed by a combination of several characters. The leaves are often (but not in T. cumminsii (Hook. f.) N. P. Barker & H. P. Linder) rolled and sclerophyllous, and the plants form persistent, small, tough, wiry tussocks (except in T. cumminsii). The lemma indumentum is often aggregated into tufts, but this is rather variable, and the lemma indumentum variation is useful for distinguishing the species. However, the regular 2-row pattern of lemma indumentum common in Rytidosperma is not found here. The lemmas in Tenaxia are deeply lobed, and 351 Distribution and habitat. The species of Tenaxia are typical montane grasses, widespread in the African mountains, and reaching the Himalayas. This is the dominant grass on the higher and cooler mountains in the Great Karoo of South Africa, but is rare in the Ethiopian uplands. The species are often found in dry, or at least seasonally dry, grassland. In this it differs from the other Afromontane genus of danthonioid grasses, Merxmuellera s. str., which is more typical of wetter grasslands. There are exceptions to this pattern and several species are found in permanent bogs. Etymology. The name refers to the tough leaves characteristic of the genus. Such tough, sclerophyllous leaves are quite frequent in the subfamily. However, in Tenaxia, the plants form tough tussocks capable of withstanding drought and are often found in rather harsh habitats. Included species. The eight species included in the new genus Tenaxia were previously classified in Merxmuellera and Danthonia. The species have been taxonomically treated in various floras, in particular the southern African grass account (Gibbs Russell et al., 1990) and the Flora of Ethiopia and Eritrea (Phillips, 1995). However, although there have been several regional treatments, a critical treatment of Tenaxia in the Himalayas that considers the taxa across the whole region still remains to be done. 1. Tenaxia aureocephala (J. G. Anderson) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia aureocephala J. G. Anderson, Bothalia 8: 170. 1964. Merxmuellera aureocephala (J. G. Anderson) Conert, Senckenberg. Biol. 51: 132. 1970. TYPE: South Africa. KwaZulu-Natal: 352 Annals of the Missouri Botanical Garden Cathedral Peak Forest Research Station, s.d., D. J. B. Killick 1727 (holotype, PRE!). 2. Tenaxia cachemyriana (Jaub. & Spach) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia cachemyriana Jaub. & Spach, Ill. Pl. Orient. 4: 46, pl. 331. 1851. TYPE: India. ‘‘ad rupes editissimas Emodi Cachemyriani legit Jacquemont, 8/1831,’’ V. Jacquemont s.n. (holotype, P not seen). 3. Tenaxia cumminsii (Hook. f.) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia cumminsii Hook. f., Fl. Brit. India (J. D. Hooker) 7: 282. 1896. Danthonia jacquemontii Bor, Kew Bull. 7: 80. 1952, nom. superfl. TYPE: India. Bhotan, Gnatong, Sikkim frontier, s.d., Cummins s.n. (holotype, K not seen). Danthonia cachemyriana Jaub. & Spach var. minor Hook. f., Fl. Brit. India (J. D. Hooker) 7: 282. 1896. Danthonia jacquemontii var. minor (Hook. f.) Bor, Kew Bull. 7: 81. 1952, nom. inval. TYPE: India. ‘‘Alpine Himalaya, Garwhal to Sikkim,’’ s.d., herb. Strachey & Winterbottom 2 (holotype, K!; isotype, BM!). Danthonia schneideri Pilg., Repert. Sp. Nov. Regni Veg. 17: 131. 1921. TYPE: China. ‘‘Yunnan, auf alpinen Wiesen an der Südsiete der Berge bei Lichiang, 4200 m.,’’ Sep. 1914, C. Schneider 2342 (holotype, B; isotypes, FR!, K!, US 776504 not seen). Bor (1952) proposed Danthonia jacquemontii as a replacement name for D. cachemyriana Hook. f., which he realized was misapplied. However, he noted that D. cumminsii Hook. f. was an abnormal form of D. cachemyriana var. minor Hook. f., thus rendering his own D. jacquemontii superfluous. Consequently, D. jacquemontii var. minor (Bor) Hook. f. is also invalid. 4. Tenaxia disticha (Nees) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia disticha Nees, Fl. Afr. Austral. Ill. 335. 1841. Merxmuellera disticha (Nees) Conert, Senckenberg. Biol. 51: 132. 1970. Rytidosperma distichum (Nees) Cope, Fl. Zambes. 10, 2: 9. 1999. TYPE: [South Africa.] ‘‘in siccis ad Sternbergspruit, alt. 5000–55009,’’ J. F. Drège (lectotype, designated by Conert [1970: 132], HBG!; isotype, S!). 5. Tenaxia dura (Stapf) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia dura Stapf, Fl. Cap. (Harvey) 7: 527. 1899. Merxmuellera dura (Stapf) Conert, Senckenberg. Biol. 51: 132. 1970. TYPE: South Africa. Little Namaqualand, Kamiesbergen, betw. Pedros Kloof & Lily Fontein, s.d., J. F. Drège s.n. (holotype, K not seen; isotype, S!). Danthonia brachyacme Pilg., Bot. Jahrb. Syst. 44: 114. 1909. TYPE: South Africa. Calvinia, ‘‘Suedost Hang des Roepmyniet,’’ 1000 m, 15 Sep. 1900, Diels 676 (holotype, B not seen; isotype, B fragm. at FR!). 6. Tenaxia guillarmodiae (Conert) N. P. Barker & H. P. Linder, comb. nov. Basionym: Merxmuellera guillarmodiae Conert, Senckenberg. Biol. 56: 145. 1975. TYPE: Lesotho. Butha-Butha, Ischlanyana Valley, s.d., A. Jacot Guillarmod 2320 (holotype, RUH!; isotype, FR not seen). 7. Tenaxia stricta (Schrad.) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia stricta Schrad., Mant. 2 (Schultes), 383. 1824. Pentameris stricta (Schrad.) Nees, Linnaea 7: 310, 313. 1832. Chaetobromus strictus (Schrad.) Nees, Fl. Afr. Austral. Ill. 341. 1841. Merxmuellera stricta (Schrad.) Conert, Senckenberg. Biol. 51: 133. 1970. TYPE: South Africa. Cape Town, s.d., C. H. F. Hesse s.n. (holotype, GOET 2247!; isotype, C!). 8. Tenaxia subulata (A. Rich.) N. P. Barker & H. P. Linder, comb. nov. Basionym: Danthonia subulata A. Rich., Tent. Fl. Abyss. 2: 420. 1851. Rytidosperma subulatum (A. Rich.) Cope, Kew Bull. 39: 835. 1984. TYPE: Ethiopia. ‘‘crescit in montosis provinciae Ouodgerate,’’ s.d., A. Petit s.n. (holotype, P not seen; isotype, P fragm. at K!). Danthonia candida Hochst. ex Steud., Syn. Pl. Glumac. 1: 244. 1854. TYPE: Ethiopia. Debra Eski, 90009, Nov., W. G. Schimper s.n. (holotype, P not seen; isotypes, P fragm. at FR!, P fragm. at K!). XV. Schismus P. Beauv., Ess. Agrostogr. 73, Pl. xv, fig. iv. 1812. Electra Panz., Ideen Rev. Gräser 49. 1813; Denkschr. Königl. Akad. Wiss. München 4(3): 299. 1814, nom. superfl. TYPE: Schismus calycinus (Loefl.) K. Koch (; Festuca calycina Loefl.) (lectotype, designated by Niles & Chase [1925: 181]). Figure 17. Hemisacris Steud., Flora 12: 490. 1829. TYPE: Hemisacris gonatodes Steud. (5 Schismus barbatus (L.) Thell.). Plants small, tufted, softly herbaceous, annual or perennial, without stolons; culms 0.05–0.35 m tall. Ligules ciliate; leaves orthophyllous, often rolled, glabrous or pilose. Inflorescences sparsely to widely paniculate, spikelike, contracted or open. Spikelets with 3 to 7 florets; glumes shorter to rarely longer than the florets, 3.7–7 mm, with 3 to 7 veins; callus rounded, villous, shorter than the rachilla internodes; lemma with 9 veins; lemma dorsal indumentum scattered or as marginal tufts or in longitudinal lines, hairs linear or rarely clavate; lemma apex tridentate to lobed; lemma lobes rounded to acute or acuminate, Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 353 to dry habitats. Two species (Schismus arabicus Nees and S. barbatus (L.) Thell.) have become worldwide weeds, mostly associated with the cultivation of wheat. Discussion. Schismus is linked to the Rytidosperma clade by a caryopsis with a punctate-ovate hilum and the small tufted growth form. Within this clade the genus can, to some extent, be diagnosed by the very short central awn, which is generally shorter than the lemma lobes. However, S. schismoides (Stapf ex Conert) Verboom & H. P. Linder is an exception, in that it has an articulated central awn more typical for danthonioid taxa. Another unusual feature is that the glumes are relatively short, but this attribute also occurs occasionally in other genera of the Rytidosperma clade. Included species. We include five species in the genus Schismus. The species in this genus were carefully and critically revised by Conert and Türpe (1974), and subsequent floristic treatments followed the Conert and Türpe classification. 1. Schismus arabicus Nees, Fl. Afr. Austral. Ill. 422. 1841. Figure 17. Schismus barbatus. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from Moore 8205. 2. Schismus barbatus (L.) Thell., Bull. Herb. Boissier, ser. 2, 7: 391. 1907. 3. Schismus inermis (Stapf) C. E. Hubb., Fl. Trop. Afr. (Oliver et al.) 10: 147. 1937. muticous or with short apical setae; lemma central awns usually minute or small, scarcely taller than the lobes, occasionally well developed with a twisted column and a long straight limb; paleae obovate to lorate, sometimes with tufts of hairs on the margins; lodicules square to cuneate, usually with both bristles and microhairs. Caryopsis lorate, elliptical to obovate, embryo about 1/2 and the ovate to punctiform hilum about 1/5 as long as the caryopsis. Cytology. 2n 5 12, 24, 36, 48, 72 (Gould, 1958; Löve & Löve, 1961; Bowden & Senn, 1962; Faruqi & Quraish, 1979; du Plessis & Spies, 1988; Spies & du Plessis, 1988). Anatomy. Leaf orthophyllous, expanded or folded, adaxially scarcely grooved; adaxial and abaxial sclerenchyma as small strands or poorly developed girders; bulliform cells absent. Distribution and habitat. This is an African and southern European genus, with a remarkable transAfrican disjunction: one species is restricted to the shores of the Mediterranean, a second species is disjunct between South Africa and the northern margins of the Sahara, and the remaining species are South African. Most species are found in semi-arid 4. Schismus scaberrimus Nees, Fl. Afr. Austral. Ill. 423. 1841. 5. Schismus schismoides (Stapf ex Conert) Verboom & H. P. Linder, comb. nov. Basionym: Danthonia schismoides Stapf ex Conert, Senckenberg. Biol. 46: 180. 1965. Karroochloa schismoides (Stapf ex Conert) Conert & Türpe, Senckenberg. Biol. 50: 299. 1969. TYPE: South Africa. Great Buschmanland, Wortel, s.d., R. Schlechter s.n. (holotype, K not seen; isotypes, E!, L not seen, Z!). XVI. Tribolium Desv., Opusc. Sci. Phys. Nat. 64. 1831. TYPE: Tribolium hispidum (Thunb.) Desv. (; Dactylis hispida Thunb.). Figure 18. Lasiochloa Kunth, Révis. Gramin. 2: 556. 1832. Allagostachyum Nees ex Steud., Nomencl. Bot. (Steudel), ed. 2, 1: 50. 1840, nom. nud., nom. superfl. TYPE: Lasiochloa longifolia (Schrad.) Kunth (; Dactylis longifolia Schrad., 5 Tribolium hispidum (Thunb.) Desv.). Hystringium Steud., Nomencl. Bot. (Steudel), ed. 2, 2: 11. 1841, nom. inval., in syn. sub Tribolium echinatum (Thunb.) Renvoize. TYPE: Hystringium acuminatum Trin. ex Steud., Nomencl. Bot., ed. 2, 2: 11. 1841. nom. inval., pro syn. Lasiochloa ciliaris Kunth. Brizopyrum Stapf, Fl. Cap. (Harvey) 7: 318, 701. 1898, 1900, nom. illeg., non Brizopyrum Link, 1827. Plagiochloa 354 Annals of the Missouri Botanical Garden Figure 18. Tribolium echinatum. —A. Spikelet. —B. Glume. —C. Lemma back. —D. Palea. Drawn by Jasmin Baumann from Bachmann 745. Adamson & Sprague, J. S. African Bot. 7: 89. 1941. TYPE: Brizopyrum capense (Spreng.) Nees (; Cynosurus uniolae L. f.). Urochlaena Nees, Fl. Afr. Austral. Ill. 437. 1841. TYPE: Urochlaena pusilla Nees. Karroochloa Conert & Türpe, Senckenberg. Biol. 50: 290. 1969, syn. nov. TYPE: Karroochloa curva (Nees) Conert & Türpe (; Danthonia curva Nees). Plants small, tufted, herbaceous, annual or perennial, culms 0.03–0.6 m tall, glabrous or variously pilose or hispid, often with long stolons. Ligules ciliate; leaves orthophyllous, expanded or setaceous. Inflorescences racemose to paniculate, secund or not, spicate, contracted, capitate or rarely 6 expanded and open, in one case disarticulating below the inflorescence. Spikelets with 2 to 10 florets; glumes shorter to longer than the florets, 2–7.5 mm, with 3 to 5 veins, glabrous or hispid with tubercle-based hairs; callus rounded, villous, shorter or longer than the rachilla internode; lemmas with 7 or 9 veins; lemma dorsal indumentum either pilose or tufted with tufts in various patterns, hairs either simple or clavate; lemma Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae apically acute, tridentate or lobed; lemma lobes rounded or acute or acuminate, rarely with short apical setae; lemma central awns poorly developed as the continuation of the acuminate lemma, or as a short central point in a tridentate lemma, or as a geniculate structure with a corkscrewed basal column and a hairlike apical part; paleae obovate to lorate, apically rounded, emarginate or bilobed; keels scabrid, often with tufts of hair on the margins, between the keels glabrous, puberulent, or villous; lodicules cuneate, bristles and microhairs sometimes present; ovary glabrous. Caryopsis lorate, elliptical, or obovate; wall sometimes tardily separable from the seed; embryo 1/3 to 2/3 and the ovate to punctate hilum less than 1/3 of the caryopsis length. a hispid appearance. These may also be found on the leaves. The lemma lobes are often fused to the central awn, resulting in an acute or acuminate lemma. The central awn is often not geniculate, but a simple straight structure, and usually associated with the acute lemmas. Finally, the inflorescence is often condensed, and in the two species of section Uniolae, the inflorescence is a spicate, secund structure. To accommodate the variation in the genus, we recognize three sections. Nomenclatural note. Karroochloa has to be included in Tribolium, as three of the four species assigned to the genus (including the type) are nested within Tribolium according to the molecular phylogeny (Verboom et al., 2006). The fourth species, K. schismoides, is nested in Schismus and is transferred to that genus. Cytology. 2n 5 12, 24, 36 (de Wet, 1954, 1960; Spies & du Plessis, 1986, 1988; Visser & Spies, 1994c, d, e; Baeza, 1996b). Anatomy. Leaf anatomy orthophyllous; adaxial ribs absent or weakly developed, rounded; adaxial sclerenchyma usually as small strands, rarely Tshaped or as inversely anchor-shaped strands, associated with all vascular bundles; abaxial sclerenchyma as strands or small girders; bulliform cells sometimes present. Distribution and habitat. This genus of smallstatured grasses is typical of the winter-rainfall area of southern Africa. The annuals in the genus are mostly localized along the coastal platform of the western and northern Cape, on coastal sands, limestone, granite, or less commonly shale. Several species of Tribolium extend in the semi-arid Karoo summer rainfall area, where they are generally associated with the cooler, higher mountains. Discussion. Tribolium is clearly a member of the Rytidosperma clade by the punctate hilum and the habit as small herbaceous tufts (Verboom et al., 2006). The genus has a number of attributes, none of which are found in all of the species, which are rare or unique in the danthonioid grasses. These include stolons, by which the plants spread to form extensive clones. These stolons are often clearly visible on herbarium specimens. There are often tuberculate hairs on the glumes, with these often visible as rough bristles, which give the spikelets or at least the glumes 355 XVIa. Tribolium Desv. sect. Tribolium. This section includes plants with hispid glumes, often with large cushion-based hairs. Included species. Only five species are included in section Tribolium; these were all revised critically by Linder and Davidse (1997) and studied cytologically and embryologically by Visser and Spies (1994a, b, c, d, e). 1. Tribolium ciliare (Stapf) Renvoize, Kew Bull. 40: 799. 1985. 2. Tribolium echinatum (Thunb.) Renvoize, Kew Bull. 40: 798. 1985. 3. Tribolium hispidum (Thunb.) Desv., Opusc. Sci. Phys. Nat. 64. 1831. 4. Tribolium pusillum (Nees) H. P. Linder & Davidse, Bot. Jahrb. Syst. 119: 295. 1997. 5. Tribolium utriculosum (Nees) Renvoize, Kew Bull. 40: 798. 1985. XVIb. Tribolium Desv. sect. Acutiflorae N. C. Visser & Spies ex H. P. Linder & Davidse, Bot. Jahrb. 119: 477. 1997. TYPE: Tribolium acutiflorum (Nees) Renvoize (; Brizopyrum acutiflorum Nees). Plants with stolons characterize section Acutiflorae within the genus. Included species. The seven species in this section were previously described in or assigned to either Tribolium, Schismus, or Karroochloa. These were all taxonomically and nomenclaturally revised by Conert and Türpe (1969) or Linder and Davidse (1997). 1. Tribolium acutiflorum (Nees) Renvoize, Kew Bull. 40: 798. 1985. 2. Tribolium curvum (Nees) Verboom & H. P. Linder, comb. nov. Basionym: Danthonia curva Nees, Fl. Afr. Austral. Ill. 328. 1841. Karroochloa curva (Nees) Conert & Türpe, Sencken- 356 Annals of the Missouri Botanical Garden berg. Biol. 50: 295. 1969. TYPE: [South Africa.] ‘‘Uitenhaag, Zwartkopsrivier, Thal und angrenzende Hügel von Villa Paul Maré bis Uitenhaag, 50–500 ft. (loc 2),’’ Nov., C. F. Ecklon 4529b (lectotype, designated by Conert & Türpe [1969: 298], HBG!; isotypes, S!, Z!, ZT!). Included species. Only two species are included in this section, and these were also discussed by Linder and Davidse (1997) and studied cytologically and embryologically by Visser and Spies (1994a, b, c, d, e). Danthonia bachmannii Hack., Bull. Herb. Boissier 3: 385. 1895. TYPE: South Africa. Cape Province: Malmesbury division, near Hopefield, Sep. 1885, F. E. Bachmann 1018 (holotype, Z!; isotypes, B not seen, B fragm. at FR!, K!). 1. Tribolium brachystachyum (Nees) Renvoize, Kew Bull. 40: 798. 1985. 3. Tribolium obliterum (Hemsl.) Renvoize, Kew Bull. 40: 798. 1985. 4. Tribolium obtusifolium (Nees) Renvoize, Kew Bull. 40: 799. 1985. 5. Tribolium pleuropogon (Stapf) Verboom & H. P. Linder, comb. nov. Basionym: Schismus pleuropogon Stapf, Bull. Misc. Inform. Kew 1916: 234. 1916. TYPE: South Africa. Damp places near Riversdale, s.d., R. Schlechter 1759 (holotype, K not seen; isotypes, GRA!, Z!). 6. Tribolium purpureum (L. f.) Verboom & H. P. Linder, comb. nov. Basionym: Avena purpurea L. f., Suppl. Pl. 112. 1781. Danthonia purpurea (Thunb.) P. Beauv., Ess. Agrostogr. 160. 1812. Karroochloa purpurea (L. f.) Conert & Türpe, Senckenberg. Biol. 50: 303. 1969. TYPE: South Africa. s.d., C. P. Thunberg (holotype, UPS 2620!; isotype, S!). Conert and Türpe (1969) indicate that the origin of the type collection was erroneously attributed to Martinique, West Indies, by Linnaeus fil. As already noted by Willdenow (1797: 450), the place of origin was the Cape of Good Hope in South Africa. 7. Tribolium tenellum (Nees) Verboom & H. P. Linder, comb. nov. Basionym: Danthonia tenella Nees, Fl. Afr. Austral. Ill. 324. 1841. Karroochloa tenella (Nees) Conert & Türpe, Senckenberg. Biol. 50: 308. 1969. TYPE: South Africa, ‘‘inter Buffelrivier (Koussie) flumen et Zilverfontein in elice ab aquis relicta inter praerupta montium,’’ 2000 ft., Aug., J. F. Drège s.n. (holotype, B not seen; isotypes, HBG!, S!). XVIc. Tribolium Desv. sect. Uniolae N. C. Visser & Spies ex H. P. Linder & Davidse, Bot. Jahrb. 119: 471. 1997. TYPE: Tribolium uniolae (L. f.) Renvoize. Tribolium sect. Uniolae is characterized by plants with secund, spicate inflorescences. 2. Tribolium uniolae (L. f.) Renvoize, Kew Bull. 40: 797. 1985. XVII. Rytidosperma Steud., Syn. Pl. Glumac. 1: 425. 1854. TYPE: Rytidosperma lechleri Steud. Figure 19. Danthonia sect. Eudanthonia Benth., Fl. Austral. 7: 591, 592. 1878, nom. inval. Monostachya Merr., Philipp. J. Sci., 5: 330. 1910. TYPE: Monostachya centrolepidoides Merr. Notodanthonia Zotov, New Zealand J. Bot. 1: 104. 1963. TYPE: Notodanthonia unarede (Raoul) Zotov (; Danthonia unarede Raoul). Erythranthera Zotov, New Zealand J. Bot. 1: 124. 1963. TYPE: Erythranthera australis (Petrie) Zotov (; Triodia australis Petrie). Pyrrhanthera Zotov, New Zealand J. Bot. 1: 125. 1963. TYPE: Pyrrhanthera exigua (Kirk) Zotov (; Triodia exigua Kirk). Joycea H. P. Linder, Telopea 6: 611. 1996. TYPE: Joycea pallida (R. Br.) H. P. Linder (; Danthonia pallida R. Br.). Thonandia H. P. Linder, Telopea 6: 612. 1996. TYPE: Thonandia longifolia (R. Br.) H. P. Linder (; Danthonia longifolia R. Br.), nom. superfl. pro Notodanthonia Zotov. Austrodanthonia H. P. Linder, Telopea 7: 269. 1997. TYPE: Austrodanthonia caespitosa (Gaudich.) H. P. Linder (; Danthonia caespitosa Gaudich.). Plants perennial, tufted, cushion-forming, matforming, or loosely to densely caespitose, or as solitary shoots from long spreading rhizomes; culms to 1.6 m tall, but most species substantially less than 1 m tall; innovation buds intravaginal or extravaginal, stolons rarely present. Ligule ciliate; leaf blades orthophyllous or sclerophyllous, persistent or rarely disarticulating from the sheaths, glabrous or pilose, rarely villous, sometimes with a web of interlocking hairs adaxially above the ligule. Inflorescences usually paniculate, rarely racemose or spicate, contracted (mostly) or expanded; florets chasmogamous or cleistogamous, some species have both. Spikelets with 2 to 7 florets; glumes shorter to usually longer than the florets, with 1 to 13 veins, without tuberclebased hairs, 1.3–23 mm; callus blunt, villous, shorter than to much longer than the rachilla internode; lemmas with 5 to 9 veins; lemma dorsal indumentum rarely absent, more commonly scattered over the lemma back, or organized into 2 horizontal rows of Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 357 from the seed; embryo 1/3 to 2/3 and the punctiform or elliptical hilum 1/10 to 1/3 the caryopsis length. Cytology. 2n 5 20, 24, 48, 72, 96, 120, 156 (Calder, 1937; Abele, 1959; Brock & Brown, 1961; Borgmann, 1964; Connor & Dawson, 1993; Baeza, 1996b; Murray et al., 2005). Anatomy. Leaves generally orthophyllous or rarely sclerophyllous; adaxially scarcely ribbed, rarely with massive ribs; adaxial sclerenchyma as small strands, girders, or inversely anchor-shaped girders; abaxial sclerenchyma as small strands to massive girders, rarely forming a continuous subepidermal layer; bulliform cells generally present. Distribution and habitat. Rytidosperma is very widespread and indeed common in temperate and cool-temperate habitats in New Guinea, Australia, New Zealand, and South America. On the Australian Tablelands and in the Murrumbidgee–Murray Basin, this genus can dominate the grassland flora. Species are found over a wide habitat range: from well-drained and/or rocky to boggy habitats; from full sun to moderate shade in open woodland or forest; from coastal to alpine habitats (but never in tropical habitats); and from the arid margins of Central Australia to the wet grasslands of New Zealand’s Westland. Figure 19. Rytidosperma caespitosum. —A. Spikelet. —B. Lemma back. —C. Palea. Drawn by Jasmin Baumann from Whibley 2707. tufts, which can be variously complete; lemma apex rarely acute, tapering into the central awn, more often lobed, the lobes shorter than to longer than the lemma body, 6 acute, generally tapering into setae; lemma central awns generally geniculate, the column corkscrewed, the apical portion straight; paleae broadly obovate to linear, shorter to longer than the lemma, apically rounded to bilobed, the keels scabrid, the area between the keels and the palea flaps glabrous or villous; lodicules cuneate or rhomboid, generally with bristles and microhairs; ovaries glabrous. Caryopsis lorate, elliptical, or obovate, the wall rarely separate Discussion. Typically, the species of Rytidosperma have punctiform hila and are small, herbaceous, and tufted. There are a few species that form spiny vegetable hedgehogs in the New Guinean alpine flora (e.g., R. oreoboloides (F. Muell.) H. P. Linder), or are somewhat larger, tougher tussocks (e.g., R. pallidum (R. Br.) A. M. Humphreys & H. P. Linder), or that spread underground by means of long rhizomes (R. exiguum (Kirk) H. P. Linder). Most species have a lemma indumentum arranged basically in two parallel rows of tufts, without or rarely with scattered hairs between the rows. This indument pattern can be variously reduced, until in some instances no tufts are left and the lemma backs are glabrous. In other species, the tufted arrangement breaks down and the indumentum is more or less evenly distributed on the lemma back. This effectively means that there seems to be no simple way in which this genus can be diagnosed by patterns of lemma indumentum against, e.g., Tribolium or Schismus, although the molecular signal is strong. Included species. Here we include 73 species in the genus Rytidosperma. This is a much broader circumscription of the genus than that used by Linder and Verboom (1996). It is similar to the circumscription used by Veldkamp (1980) and Connor and Edgar (1979), but the interpretation of Clayton and Renvoize (1986) was much wider, including the African species as well. Our species treatment follows the three most 358 Annals of the Missouri Botanical Garden important regional treatments: from Australia (including Austrodanthonia, Joycea, Rytidosperma, and Notodanthonia, sensu Linder [2005]), from New Zealand (including Erythranthera and Rytidosperma [Edgar & Connor, 2000]), and also from South America under Rytidosperma (Baeza, 1996a). However, there is not yet a global revision of the entire genus. 11. Rytidosperma clavatum (Zotov) Connor & Edgar, New Zealand J. Bot. 17: 326. 1979. 1. Rytidosperma acerosum (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 331. 1979. 2. Rytidosperma alpicola (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 331. 1979. 3. Rytidosperma auriculatum (J. M. Black) Connor & Edgar, New Zealand J. Bot. 17: 322. 1979. 4. Rytidosperma australe (Petrie) Clayton & Renvoize ex Connor & Edgar, New Zealand J. Bot. 25: 166. 1987. 5. Rytidosperma biannulare (Zotov) Connor & Edgar, New Zealand J. Bot. 17: 324. 1979. 6. Rytidosperma bipartitum (Kunth) A. M. Humphreys & H. P. Linder, comb. nov. Basionym: Avena bipartita Link, Hort. Berol. (Link) 1: 113. 1827. Austrodanthonia bipartita (Link) H. P. Linder, Telopea 7: 270. 1997. Danthonia linkii Kunth, Enum. Pl. (Kunth) 1: 315. 1833, nom. superfl. Rytidosperma linkii (Kunth) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979, nom. illeg. Notodanthonia linkii (Kunth) Veldkamp, Taxon 29: 296. 1980, nom. illeg. TYPE: Australia. NSW, 9 mi. [14.4 km] WNW of Newcastle NSW, 27 Apr. 1960, R. Story 7220 (neotype, designated here, CANB!). 12. Rytidosperma clelandii (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 13. Rytidosperma corinum Connor & Edgar, New Zealand J. Bot. 17: 317. 1979. 14. Rytidosperma craigii (Veldkamp) H. P. Linder, Telopea 6: 613. 1996. 15. Rytidosperma dendeniwae (Veldkamp) H. P. Linder, Telopea 6: 613. 1996. 16. Rytidosperma diemenicum (D. I. Morris) A. M. Humphreys & H. P. Linder, comb. nov. Basionym: Danthonia diemenica D. I. Morris, Muellera 7: 153. 1990. Notodanthonia diemenica (D. I. Morris) H. P. Linder, Telopea 6: 616. 1996. Austrodanthonia diemenica (D. I. Morris) H. P. Linder, Telopea 7: 271. 1997. TYPE: Australia. Tasmania: Ouse River, Wild Dog Plains, s.d., A. Moscal 1292 (holotype, HO 65782!). 17. Rytidosperma dimidiatum (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 18. Rytidosperma duttonianum (Cashmore) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 19. Rytidosperma erianthum (Lindl.) Connor & Edgar, New Zealand J. Bot. 17: 323. 1979. 20. Rytidosperma exiguum (Kirk) H. P. Linder, Telopea 6: 614. 1996. 21. Rytidosperma fortunae-hibernae (Renvoize) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 9. Rytidosperma caespitosum (Gaudich.) Connor & Edgar, New Zealand J. Bot. 17: 325. 1979. 22. Rytidosperma fulvum (Vickery) A. M. Humphreys & H. P. Linder, comb. nov. Basionym: Danthonia linkii Kunth var. fulva Vickery, Contr. New South Wales Natl. Herb. 1: 299. 1950. Rytidosperma linkii var. fulvum (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. Notodanthonia bipartita (Link) Veldkamp var. fulva (Vickery) Veldkamp, Taxon 29: 296. 1980. Notodanthonia fulva (Vickery) H. P. Linder, Telopea 6: 616. 1996. Austrodanthonia fulva (Vickery) H. P. Linder, Telopea 7: 271. 1997. TYPE: Australia. New South Wales: Flemington, 31 Mar. 1929, G. B. Vickery (holotype, NSW 1573!; isotypes, K not seen, L not seen). 10. Rytidosperma carphoides (F. Muell. ex Benth.) Connor & Edgar, New Zealand J. Bot. 17: 331. 1979. 23. Rytidosperma geniculatum (J. M. Black) Connor & Edgar, New Zealand J. Bot. 17: 323. 1979. The original type was presumably at B, but has not been found despite repeated searches by several persons. Presumably, it was destroyed during World War II. In order to prevent any confusion, a suitable neotype is selected. 7. Rytidosperma bonthainicum (Jansen) Veldkamp, Reinwardtia 12: 139. 2004. 8. Rytidosperma buchananii (Hook. f.) Connor & Edgar, New Zealand J. Bot. 17: 320. 1979. Volume 97, Number 3 2010 Linder et al. Classification of Danthonioideae 24. Rytidosperma gracile (Hook. f.) Connor & Edgar, New Zealand J. Bot. 17: 330. 1979. 42. Rytidosperma nudum (Hook. f.) Connor & Edgar, New Zealand J. Bot. 17: 322. 1979. 25. Rytidosperma horrens Connor & Molloy, New Zealand J. Bot. 43: 726. 2005. 43. Rytidosperma occidentale (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 26. Rytidosperma indutum (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 44. Rytidosperma oreoboloides (F. Muell.) H. P. Linder, Telopea 6: 614. 1996. 27. Rytidosperma javanicum (Ohwi ex Veldkamp) H. P. Linder, Telopea 6: 614. 1996. 45. Rytidosperma oreophilum H. P. Linder & N. G. Walsh, Muelleria 8: 283. 1995. 28. Rytidosperma laeve (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 325. 1979. 46. Rytidosperma pallidum (R. Br.) A. M. Humphreys & H. P. Linder, comb. nov. Basionym: Danthonia pallida R. Br., Prodr. Fl. Nov. Holland. 177. 1810. Avena brownii Spreng., Syst. Veg., ed. 16 (Sprengel) 1: 336. 1825. Notodanthonia pallida (R. Br.) Veldkamp, Taxon 29: 297. 1980. Danthonia penicillata (Labill.) P. Beauv. var. pallida (R. Br.) Rodway, Tasman. Fl. 267. 1903. Chionochloa pallida (R. Br.) S. W. L. Jacobs, Taxon 31: 742. 1982. Joycea pallida (R. Br.) H. P. Linder, Telopea 6: 611. 1996. TYPE: Australia. New South Wales: Port Jackson, s.d., R. Brown 6232 (holotype, BM not seen; isotype, K not seen, K fragm. at PERTH!). 29. Rytidosperma lechleri Steud., Syn. Pl. Glumac. 1: 425. 1854. 30. Rytidosperma lepidopodum (N. G. Walsh) A. M. Humphreys & H. P. Linder, comb. nov. Basionym: Danthonia lepidopoda N. G. Walsh, Muellera 7: 384. 1991. Joycea lepidopoda (N. G. Walsh) H. P. Linder, Telopea 6: 612. 1996. TYPE: Australia. Victoria: Bullens Land, Courtneys Rd., N of Ash Reserve, 15 Jan. 1987, N. G. Walsh 1709 (holotype, MEL!; isotypes, BRI!, NSW!). 31. Rytidosperma longifolium (R. Br.) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 32. Rytidosperma maculatum (Zotov) Connor & Edgar, New Zealand J. Bot. 17: 320. 1979. 33. Rytidosperma mamberamense (Jansen) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 34. Rytidosperma merum Connor & Edgar, New Zealand J. Bot. 17: 328. 1979. 35. Rytidosperma monticola (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 36. Rytidosperma montis-wilhelmii (Veldkamp & Fortuin) H. P. Linder, Telopea 6: 614. 1996. 359 47. Rytidosperma paschalis (Pilg.) C. M. Baeza, Gayana, Bot. 47: 83–84. 1991. 48. Rytidosperma pauciflorum (R. Br.) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 49. Rytidosperma penicillatum (Labill.) Connor & Edgar, New Zealand J. Bot. 17: 327. 1979. 50. Rytidosperma petrosum Connor & Edgar, New Zealand J. Bot. 17: 317. 1979. 51. Rytidosperma pictum (Nees & Meyen) Nicora, Darwiniana 18: 91. 1973. 51a. Rytidosperma pictum (Nees & Meyen) Nicora var. pictum. 37. Rytidosperma nardifolium (Veldkamp) H. P. Linder, Telopea 6: 614. 1996. 51b. Rytidosperma pictum (Nees & Meyen) Nicora var. bimucronatum Nicora, Darwiniana 18(1, 2): 91. 1973. 38. Rytidosperma nigricans (Petrie) Connor & Edgar, New Zealand J. Bot. 17: 331. 1979. 52. Rytidosperma pilosum (R. Br.) Connor & Edgar, New Zealand J. Bot. 17: 326. 1979. 39. Rytidosperma nitens (D. I. Morris) H. P. Linder, Telopea 6: 614. 1996. 53. Rytidosperma popinensis (D. I. Morris) A. M. Humphreys & H. P. Linder, comb. nov. Basionym: Danthonia popinensis D. I. Morris, Muelleria 7: 157. 1989. Notodanthonia popinensis (D. I. Morris) H. P. Linder, Telopea 6: 616. 1996. Austrodanthonia popinensis (D. I. Morris) H. P. Linder, Telopea 7: 273. 1997. TYPE: Australia. Tasmania: 0.5 km N of Kempton, 16 Jan. 1987, 40. Rytidosperma nivicola (Vickery) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 41. Rytidosperma nudiflorum (P. F. Morris) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 360 Annals of the Missouri Botanical Garden D. I. Morris 8556 (holotype, HO 92651!; isotypes, AD!, NSW!). 68. Rytidosperma unarede (Raoul) Connor & Edgar, New Zealand J. Bot. 17: 328. 1979. 54. Rytidosperma pulchrum (Zotov) Connor & Edgar, New Zealand J. Bot. 17: 321. 1979. 69. Rytidosperma vestitum (Pilg.) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 55. Rytidosperma pumilum (Kirk) Connor & Edgar, New Zealand J. Bot. 25: 166. 1987. 70. Rytidosperma vickeryae M. Gray & H. P. Linder, Austral. Syst. Bot. 12(5): 744. 1999. 56. Rytidosperma quirihuense C. M. Baeza, Novon 12: 31. 2002. 71. Rytidosperma violaceum (E. Desv.) Nicora, Darwiniana 18: 91. 1973. 57. Rytidosperma racemosum (R. Br.) Connor & Edgar, New Zealand J. Bot. 17: 327. 1979. 72. Rytidosperma virescens (E. Desv.) Nicora, Darwiniana 18: 93. 1973. 57a. Rytidosperma racemosum (R. Br.) Connor & Edgar var. racemosum. 72a. Rytidosperma virescens (E. Desv.) Nicora var. virescens. 57b. Rytidosperma racemosum (R. Br.) Connor & Edgar var. obtusatum (Benth.) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 72b. Rytidosperma virescens (E. Desv.) Nicora var. parvispiculum Nicora, Darwiniana 18(1, 2): 95. 1973. 58. Rytidosperma remotum (D. I. Morris) A. M. Humphreys & H. P. Linder, comb. nov. Basionym: Danthonia remota D. I. Morris, Muellera 7: 160. 1989. Notodanthonia remota (D. I. Morris) H. P. Linder, Telopea 6: 617. 1996. Austrodanthonia remota (D. I. Morris) H. P. Linder, Telopea 7: 273. 1997. TYPE: Australia. Tasmania: summit of Hibbs Pyramid, 3 Feb. 1980, A. M. Buchanan 2878 (holotype, HO 91392!). 72c. Rytidosperma virescens (E. Desv.) Nicora var. patagonicum (Speg.) Nicora, Darwiniana 18(1, 2): 93. 1973. 59. Rytidosperma richardsonii (Cashmore) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 60. Rytidosperma semiannulare (Labill.) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 61. Rytidosperma setaceum (R. Br.) Connor & Edgar, New Zealand J. Bot. 17: 332. 1979. 62. Rytidosperma setifolium (Hook. f.) Connor & Edgar, New Zealand J. Bot. 17: 316. 1979. 63. Rytidosperma sorianoi Nicora, Darwiniana 18: 89. 1973. 64. Rytidosperma telmaticum Connor & Molloy, New Zealand J. Bot. 43: 721. 2005. 65. Rytidosperma tenue (Petrie) Connor & Edgar, New Zealand J. Bot. 17: 321. 1979. 66. Rytidosperma tenuius (Steud.) O. E. Erikss., A. Hansen & Sunding, Fl. Macaronesia Checkl. Vasc. Pl. pt. 1, ed. 2, 93. 1979. 67. Rytidosperma thomsonii (Buchanan) Connor & Edgar, New Zealand J. Bot. 17: 322. 1979. 73. Rytidosperma viride (Zotov) Connor & Edgar, New Zealand J. Bot. 17: 316. 1979. Literature Cited Abele, K. 1959. Cytological studies in the genus Danthonia. Trans. Roy. Soc. South Australia 82: 163–173. 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Synopsis of the grass subfamily Arundinoideae in New Zealand. New Zealand J. Bot. 1: 78–136. 5. Leaf lamina persistent on sheath 5 0; abscising from a persistent sheath 5 1; persistent sheath often splitting 5 2. 6. Keels on leaf blades absent 5 0; present and simple 5 1; present as with several distinct keels 5 2. 7. Leaf blades asymmetrical 5 0; symmetrical 5 1. 8. Mesophyll islands of colorless cells absent 5 0; present 5 1. 9. Microhairs in leaf-grooves non-overlapping 5 0; overlapping 5 1. 10. Spikelets disarticulating below inflorescence 5 0; below glumes 5 1; above glumes 5 2. 11. Number of florets per spikelet 1 or 2 5 0; 3 or more 5 1. 12. Number of lemma veins up to 5 5 0; 7 or more 5 1. 13. Lemma indumentum hairs scattered 5 0; in tufts in transverse lines 5 1; scattered tufts 5 2; totally absent 5 3. 14. Lemma setae inserted at the tips of lemma lobes 5 0; in the sinus between the lemma lobes 5 1. 15. Lemma awn differentiated into a corkscrewed column and straight hairlike apical part 5 0; not differentiated into two parts 5 1. 16. Tufts of long hair on the palea margins absent 5 0; present 5 1. 17. Palea margins normal 5 0; inrolled 5 1. 18. Lodicule bristles absent 5 0; present 5 1. 19. Lodicule microhairs present 5 0; absent 5 1. 20. Caryopsis hilum linear 5 0; punctiform 5 1. 21. Leaves blades sclerophyllous 5 0; orthophyllous 5 1. APPENDIX 1. Morphological characters and character states for the simplified phylogeny. See Table 1 and Figure 2. 1. Plants reproductive organization: flowers bisexual, hermaphrodite 5 0; gynodioecious 5 1. 2. Cleistogenes: present 5 0; absent 5 1. 3. Plant bases swollen, villous 5 1; not 5 0. 4. Multicellular glands present 5 0; absent 5 1. APPENDIX 2. Generic groups recognized within the Danthonioideae. Genera are presented in this order in the paper, reflecting their current known relationships. a. Merxmuellera basal grade I. Merxmuellera Conert II. Geochloa H. P. Linder & N. P. Barker III. Capeochloa H. P. Linder & N. P. Barker b. Pentameris clade IV. Pentameris P. Beauv. c. Chionochloa clade V. Chionochloa Zotov d. Chaetobromus–Pseudopentameris clade VI. Pseudopentameris Conert VII. Chaetobromus Nees e. Danthonia clade VIII. Cortaderia Stapf IX. Austroderia N. P. Barker & H. P. Linder X. Plinthanthesis Steud. XI. Notochloe Domin XII. Chimaerochloa H. P. Linder XIII. Danthonia DC. f. Rytidosperma clade XIV. Tenaxia N. P. Barker & H. P. Linder XV. Schismus P. Beauv. XVI. Tribolium Desv. XVII. Rytidosperma Steud.

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Linder, H.P., Baeza, M., Barker, N.P., Galley, C., Humphreys, A.M., Lloyd, K., Orlovich, D., Pirie, M.D., Simon, B.K., Walsh, N. & Verboom, G.A. (2010). A classification of the Danthonioideae (Poaceae). Annals of the Missouri Botanical Garden 97:306-364.

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