Kelvin M. Lloyd

A Generic Classification of the Danthonioideae (Poaceae)1

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. 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.

The generic position of Austrofestuca littoralis and the reinstatement of Hookerochloa and Festucella (Poaceae) based on evidence from nuclear (ITS) and chloroplast (trnL‐trnF) DNA sequences

Abstract The segregation of the grass genus Austrofestuca from Festuca has been debated because in some respects Austrofestuca is morphologically similar to Poa. Analyses of DNA sequence information from nuclear (internal transcribed spacer) and chloroplast (trnL‐trnF) genomes indicate that Austrofestuca is not monophyletic. Austrofestuca littoralis is closely related to Poa, while A. eriopoda and A. hookeriana are placed in a clade containing Arctagrostis, Dupontia, and Arctophila. Whilst Austrofestuca littoralis is nested within the Poa clade, our data do not exclude the possibility that A. littoralis is sister to Poa. A range of morphological characters supports the differentiation of A. eriopoda and A. hookeriana from A. littoralis. We reinstate the genera Festucella and Hookerochloa to accommodate A. eriopoda and A. hookeriana, respectively.

Phytolith morphology and biogenic silica concentrations and abundance in leaves of Chionochloa (Danthonieae) and Festuca (Poeae) in New Zealand

Abstract Biogenic silica concentrations and phytolith morphology and abundance in green leaf material are described in 34 species/subspecies of Chionochloa (Danthonieae) and nine species/subspecies of Festuca (Poeae). A dry‐ash technique was used to extract biogenic silica from mature leaves from plants grown in potting mix. Phytolith morphology and abundance were characterised by scanning electron microscopy and light microscopy, respectively. Biogenic silica concentrations were lowest (<1%) in Chionochloa bromoides, C. rubra ssp. occulta, and Festuca multinodis and reached levels of c. 7% in Chionochloa defracta and Festuca ultramafica, two serpentine endemics. Cluster analysis showed that replicate samples of taxa were classified together in 70% of cases. Ordination of phytolith types identified a major gradient based on the presence/absence of panicoid and chloridoid types. A secondary gradient was related to the frequency of chionochloid, truncated cone, and other short‐cell phytoliths. Chionochloa taxa fall into 3 categories on the basis of the presence of panicoid (8 taxa) or chionochloid (11 taxa) phytolith types, or the absence of both (14 taxa). Species of Festuca had relatively high frequencies of points compared with Chionochloa, and absence or low frequency of dumbbell and chionochloid types. There was more interspecific variation among species of Chionochloa than in Festuca.

Species novae graminum Novae‐Zelandiae II. Chionochloa nivifera (Danthonieae:Danthonioideae)

Abstract A new species of Chionochloa, C. nivifera, is described from the high alpine zone of eastern Fiordland mountains. A species found in areas of long snow lie, it resembles C. macra but is of lower stature and its tillers are less erect.

Experimental translocations of the threatened New Zealand plants Carex inopinata Cook (Cyperaceae) and Simplicia laxa Kirk (Poaceae)

Carex inopinata Cook (Cyperaceae) and Simplicia laxa Kirk (Poaceae) are two threatened ‘grassy’ plants that generally occur in dry grassland and rock outcrop habitats. However, the restriction of these species to such areas may indicate relict habitats. We tested this idea by translocating both species to Ōrokonui Ecosanctuary, near Dunedin. Carex inopinata was translocated to two sites (moderatelyand highly-shaded) in 2010 and Simplicia laxa was translocated in 2011, but only to one site due to insufficient propagated material. At each site, plants were established in different microhabitats underneath a kānuka (Kunzea robusta) canopy, where ground cover was sparse and hence provided minimal competition for the translocated plants. We measured plant growth, flowering, and survival twice-yearly from 2010 to 2013, and again from 2015 to 2017. Also, we described the vegetation and recorded microclimate data at each site. Carex inopinata growth, flowering, and survival were highest under moderate shade, either in open (bare-ground) or beside-rock microhabitats. Highest growth levels alternated between these two microhabitats, particularly during a drought when plants in the open suffered some mortality or damage, while plants near rocks remained sheltered. Narrower ranges of relative humidity and temperature in rock versus open microhabitats attest to probable protection from drought. Despite 42% of originally planted C. inopinata individuals surviving, there was no seedling recruitment, therefore, the long-term persistence of this translocated population remains uncertain. Simplicia laxa did not successfully establish, although growth was also highest in open microhabitat. We cannot confirm whether habitat for S. laxa at Ōrokonui is absent but drought-related mortality influenced its failure to establish. Further monitoring of the C. inopinata plants will confirm whether this threatened sedge can persist under the conditions present at the translocation site.