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A Generic Classification of the Danthonioideae (Poaceae)
Article in Annals of the Missouri Botanical Garden · October 2010
DOI: 10.3417/2009006
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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
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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
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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
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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.
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Figure 1. Continued.
Linder et al.
Classification of Danthonioideae
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Figure 1. Continued.
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Figure 1. Continued.
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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
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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.
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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,
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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
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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
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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
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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-
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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
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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.
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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.
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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
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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.
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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
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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:
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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!).
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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.
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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.’’
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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
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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.
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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
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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.
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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
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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.
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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,
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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
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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.
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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;
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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
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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.
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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.
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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.
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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
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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,
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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
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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
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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
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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
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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
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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.
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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.
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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.
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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.