-
Notifications
You must be signed in to change notification settings - Fork 0
/
Copy path10.3897_phytokeys.13.3355.xml
322 lines (322 loc) · 69.1 KB
/
10.3897_phytokeys.13.3355.xml
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
<!DOCTYPE article PUBLIC "-//TaxonX//DTD Taxonomic Treatment Publishing DTD v0 20100105//EN" "tax-treatment-NS0.dtd"><article article-type="research-article" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:tp="http://www.plazi.org/taxpub">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">PhytoKeys</journal-id>
<journal-title-group>
<journal-title xml:lang="en">PhytoKeys</journal-title>
<abbrev-journal-title xml:lang="en">PhytoKeys</abbrev-journal-title>
</journal-title-group>
<issn pub-type="ppub">1314-2011</issn>
<issn pub-type="epub">1314-2003</issn>
<publisher>
<publisher-name>Pensoft Publishers</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3897/phytokeys.13.3355</article-id>
<title-group>
<article-title>Systematics of
<italic>Trigonochloa</italic> (Poaceae, Chloridoideae, Chlorideae)
</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Snow</surname>
<given-names>Neil</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
</contrib>
<contrib contrib-type="author" xlink:type="simple">
<name name-style="western">
<surname>Peterson</surname>
<given-names>Paul M.</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
</contrib>
</contrib-group>
<aff id="A1">
<label>1</label>527 S. Oakes, Helena, MT 59601, USA
</aff>
<aff id="A2">
<label>2</label>Department of Botany MRC-166, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012 USA
</aff>
<author-notes>
<fn fn-type="corresp">
<p>Corresponding author: Neil Snow (<email xlink:type="simple">nwiltonsnow@gmail.com</email>)</p>
</fn>
<fn fn-type="edited-by">
<p>Academic editor: Leonardo Versieux</p>
</fn>
</author-notes>
<pub-date pub-type="collection">
<year>2012</year>
</pub-date>
<pub-date pub-type="epub">
<day>29</day>
<month>6</month>
<year>2012</year>
</pub-date>
<issue>13</issue>
<fpage>25</fpage>
<lpage>38</lpage>
<history>
<date date-type="received">
<day>9</day>
<month>5</month>
<year>2012</year>
</date>
<date date-type="accepted">
<day>16</day>
<month>6</month>
<year>2012</year>
</date>
</history>
<permissions>
<copyright-statement>Neil Snow, Paul M. Peterson</copyright-statement>
<license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/3.0" xlink:type="simple">
<license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
</license>
</permissions>
<abstract>
<label>Abstract</label>
<p>A systematic treatment including descriptions and a key for identification is provided for the two species of <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic>, a genus recently segregated from the polyphyletic <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l. <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> ranges from southern Africa east to India and Sri Lanka, reflecting the widely ranging <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic>. <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> has a more limited distribution from East Africa to Yemen. <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> is diagnosable from other chloridoid grasses based on its unusually flaccid and membranous leaves that have uniquely enlarged lateral cells in the parenchyma sheath surrounding the vascular bundles in <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> (unconfirmed for <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> given limited material), primary and secondary vascular bundles that do not project above or below in fresh material, XyMS+ leaf anatomy, narrow spicate primary inflorescence branches, spikelets with one (or rarely two) florets, thinly membranous to hyaline lemmas, and a trigonous caryopses that bear a narrow but deep sulcus on the hilar side. Lectotypes are designated for <italic><tp:taxon-name>Agrostis montana</tp:taxon-name></italic> and <italic><tp:taxon-name>Cynodon gracilis</tp:taxon-name></italic>. The synonym <italic><tp:taxon-name>Leptochloa laurentii</tp:taxon-name></italic> De Wild. is confirmed for <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic>.</p>
</abstract>
<kwd-group>
<label>Keywords</label>
<kwd>Conservation</kwd>
<kwd><italic>Leptochloa</italic></kwd>
<kwd>leptotypification</kwd>
<kwd>Poaceae</kwd>
<kwd>systematics</kwd>
<kwd>taxonomy</kwd>
</kwd-group>
</article-meta>
</front>
<body>
<sec sec-type="Introduction">
<title>Introduction</title>
<p>The generic boundaries of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> P. Beauv. have been contentious more or less continuously since the genus was first described in 1812 (<xref ref-type="bibr" rid="B28">Valls 1978</xref>; <xref ref-type="bibr" rid="B12">McNeill 1979</xref>; <xref ref-type="bibr" rid="B18">Phillips 1982</xref>; <xref ref-type="bibr" rid="B23">Snow 1997</xref>). Many previous discussions of generic boundaries focused <!--PageBreak-->on whether <italic><tp:taxon-name>Diplachne</tp:taxon-name></italic> P. Beauv., often included in <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic>,could be recognized as a distinct genus (<xref ref-type="bibr" rid="B17">Phillips 1974</xref>; <xref ref-type="bibr" rid="B10">Jacobs 1987</xref>). A global monograph of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B23">Snow 1997</xref>) based on morphology, lemmatal micromorphology (<xref ref-type="bibr" rid="B22">Snow 1996</xref>), anatomy of stems and leaves, and caryopsis morphology (<xref ref-type="bibr" rid="B24">Snow 1998</xref>) tested the monophyly of the genusin the context of a few putatively related genera. Preliminary results from cladistic studies using morphology were far from conclusive and at best suggested that <italic><tp:taxon-name>Diplachne</tp:taxon-name></italic> was not easily segregated from <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B23">Snow 1997</xref>). One noteworthy aspect of the cladistic studies (<xref ref-type="bibr" rid="B23">Snow 1997</xref>) was the consistent sister status of the <italic><tp:taxon-name>Leptochloa uniflora</tp:taxon-name></italic> Hochst. ex A. Rich. and <italic><tp:taxon-name>Leptochloa rupestris</tp:taxon-name></italic> C.E. Hubb. to other members of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l., which was predictable since these two species share many morphological and anatomical characteristics absent in other members of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l.</p>
<p><italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l. has been considered a diverse assemblage of C<sub>4</sub> (nicotinamide adenine dinucleotide co-factor malic enzyme [NAD-ME] and phosphoenolpyruvate carboxykinase [PCK]) grasses in the tribe Chlorideae (<xref ref-type="bibr" rid="B4">Clayton and Renvoize 1986</xref>; <xref ref-type="bibr" rid="B29">Watson and Dallwitz 1992</xref>; <xref ref-type="bibr" rid="B26">Soreng et al. 2012</xref>) with approximately 32 annual or perennial species (<xref ref-type="bibr" rid="B23">Snow 1997</xref>, <xref ref-type="bibr" rid="B25">2003</xref>). The range of morphological variation within many species of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> is significant (<xref ref-type="bibr" rid="B23">Snow 1997</xref>, <xref ref-type="bibr" rid="B16">Peterson et al. 2012</xref>). However, <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l. is demonstrably polyphyletic when tested with molecular DNA markers (<xref ref-type="bibr" rid="B5">Columbus et al. 2007</xref>; <xref ref-type="bibr" rid="B15">Peterson et al. 2010</xref>, <xref ref-type="bibr" rid="B16">2012</xref>). In a large phylogenetic study of the <tp:taxon-name>Chloridoideae</tp:taxon-name> based on seven DNA sequence markers the species of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l. were found to form three separate lineages (<xref ref-type="bibr" rid="B15">Peterson et al. 2010</xref>), and more recently using six DNA markers these species were found to form five separate lineages, each treated as a separate genus: <italic><tp:taxon-name>Dinebra</tp:taxon-name></italic> Jacq., <italic><tp:taxon-name>Diplachne</tp:taxon-name></italic>, <italic><tp:taxon-name>Disakisperma</tp:taxon-name></italic> Steud., <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> sensu strictu, and <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> P.M. Peterson & N. Snow (<xref ref-type="bibr" rid="B16">Peterson et al. 2012</xref>). <xref ref-type="bibr" rid="B16">Peterson et al. (2012)</xref> found that <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> (Hochst. ex A. Rich.) P.M. Peterson & N. Snow and <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> (C.E. Hubb.) P.M. Peterson & N. Snow consistently resolved as a strongly supported clade (<xref ref-type="bibr" rid="B16">Peterson et al. 2012</xref>) outside of subtribe <tp:taxon-name>Elusininae</tp:taxon-name>.</p>
<p><italic><tp:taxon-name>Leptochloa uniflora</tp:taxon-name></italic> was first described by <xref ref-type="bibr" rid="B20">Richard (1851)</xref>.<xref ref-type="bibr" rid="B3">Chippindale (1955)</xref> later transferred this species to <italic><tp:taxon-name>Craspedorhachis</tp:taxon-name></italic> Benth., although her concept of <italic><tp:taxon-name>Craspedorhachis</tp:taxon-name></italic> was broader than that of other treatments (<xref ref-type="bibr" rid="B4">Clayton and Renvoize 1986</xref>; <xref ref-type="bibr" rid="B6">Gibbs-Russell et al. 1991</xref>; <xref ref-type="bibr" rid="B29">Watson and Dallwitz 1992</xref>). One current concept of <italic><tp:taxon-name>Craspedorhachis</tp:taxon-name></italic> includes three species that collectively occupy parts of sub-Saharan Africa and Madagascar (e.g., <xref ref-type="bibr" rid="B21">Simon et al. 2011</xref>).</p>
<p><xref ref-type="bibr" rid="B1">Bentham (1882</xref>: 108) described two sections in <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic>, which he maintained distinct from <italic><tp:taxon-name>Diplachne</tp:taxon-name></italic>. In <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> sect. <italic><tp:taxon-name>Pseudocynodon</tp:taxon-name></italic> Benth., characterized by spikelets having only one or two florets, Bentham ascribed <italic><tp:taxon-name>Leptochloa uniflora</tp:taxon-name></italic> and <italic><tp:taxon-name>Leptochloa neesii</tp:taxon-name></italic> (the latter of which included his concept of <italic><tp:taxon-name>Leptochloa polystachya</tp:taxon-name></italic> Benth.). The latter species, which is also characterized by one floret per spikelet (or in rare cases two), differs significantly from <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> by characters of leaf anatomy and has been transferred to <italic><tp:taxon-name>Dinebra</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B16">Peterson et al. 2012</xref>).</p>
<p>The purpose of this paper is to present the systematics of <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> as the first step in revising <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l. into monophyletic genera.<!--PageBreak--></p>
</sec>
<sec sec-type="Materials and methods">
<title>Materials and methods</title>
<p>Approximately 110 collections were analyzed from the following 15 herbaria: B, BM, BRI, CANB, K, M, MO, NY, P, PRE, S, TAES, US, W, Z. A few specimens previously determined incorrectly as <italic><tp:taxon-name>Leptochloa uniflora</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B7">Gould and Soderstrom 1974</xref>; <xref ref-type="bibr" rid="B23">Snow 1997</xref>) have been identified subsequently as <italic><tp:taxon-name>Leptochloa neesii</tp:taxon-name></italic> Benth. [= <italic><tp:taxon-name>Dinebra neesii</tp:taxon-name></italic> (Benth.) P.M. Peterson & N. Snow] (<xref ref-type="bibr" rid="B16">Peterson et al. 2012</xref>). Geographical ranges are summarized using <xref ref-type="bibr" rid="B2">Brummitt (2001)</xref> and herbarium acronyms follow <xref ref-type="bibr" rid="B27">Thiers (2012)</xref>.</p>
<p>Fresh leaf samples of <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> and <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l. were studied for, but not summarized, in <xref ref-type="bibr" rid="B23">Snow (1997)</xref>. The first author also viewed black and white anatomical photographs of <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> taken by R. Ellis housed at PRE (Davidse & Ellis 5925, Ellis s.n., Ellis 3635, but see in particular Ellis 4534) and was able to confirm observations of <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> made previously by <xref ref-type="bibr" rid="B13">Metcalfe (1960)</xref>. Stems (culms) were hand-sectioned with fresh or rehydrated material. The sources of anatomical vouchers are mostly at MO: <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic>: Gilbert et al. 249, Pappi 2821; <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic>: Snow & Burgoyne 6978, Siame 581, Ellis 2780, Davidse et al. 6643. Lemmatal micromorphology of <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> and other species of chloridoid grasses was studied using scanning electron microscopy (<xref ref-type="bibr" rid="B22">Snow 1996</xref>), and caryopses were studied using simple light microscopy (<xref ref-type="bibr" rid="B24">Snow 1998</xref>).</p>
</sec>
<sec sec-type="Results and discussion">
<title>Results and discussion</title>
<p><italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> and <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> are highly similar in gross morphology (<xref ref-type="bibr" rid="B17">Phillips 1974</xref>, <xref ref-type="bibr" rid="B19">1995</xref>), lemmatal micromorphology (<xref ref-type="bibr" rid="B22">Snow 1996</xref>), stem and leaf anatomy, and caryopsis morphology (<xref ref-type="bibr" rid="B24">Snow 1998</xref>; <xref ref-type="bibr" rid="B16">Peterson et al. 2012</xref>). The genus differs from <italic><tp:taxon-name>Lopholepis</tp:taxon-name></italic> Decne., <italic><tp:taxon-name>Mosdenia</tp:taxon-name></italic> Stent, <italic><tp:taxon-name>Perotis</tp:taxon-name></italic> Aiton, and <italic><tp:taxon-name>Toliara</tp:taxon-name></italic> Judziewicz by having several to numerous unilateral, secund racemes scattered along a central axis rather than a single raceme or false spike (<xref ref-type="bibr" rid="B29">Watson and Dallwitz 1992</xref>; <xref ref-type="bibr" rid="B11">Judziewicz 2009</xref>). Species placed in <italic><tp:taxon-name>Lopholepis</tp:taxon-name></italic>, <italic><tp:taxon-name>Perotis</tp:taxon-name></italic>, and <italic><tp:taxon-name>Toliara</tp:taxon-name></italic> have 1-nerved lemmas whereas both species of <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> they are 3-nerved (<xref ref-type="bibr" rid="B29">Watson and Dallwitz 1992</xref>).</p>
<p><bold>Leaf anatomy.</bold> The transverse anatomical features of the leaves of these two species differ in several significant ways from the rest of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l. The leaf blades of both species are quite thin (and flaccid) when fresh in <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> and somewhat translucent. They also can be relatively broad basally and relatively short, thus appearing narrowly ovate.</p>
<p>Epidermal preparations of <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> made separately by the first author (<italic>Davidse & Ellis 5925</italic>, MO; unpublished) and by Roger Ellis (<italic>Ellis 1928</italic>; photos on herbarium specimen; PRE) show the adaxial (more so) and abaxial (less so) surfaces (apart from areas above bulliform cells) to be covered with narrow rows of relatively small cells, virtually all of which are capped by a centrally located and prominent papilla (Snow, unpubl.). In addition, the cells of the leaf blade epidermis and lemma surface are not always clearly differentiated into short and long cells (<xref ref-type="bibr" rid="B22">Snow 1996</xref>).</p>
<p><!--PageBreak-->Keels (areas of parenchyma in the middle of the leaf blade lacking vascular bundles) are absent, or if present then small, and if present then lacunae within the parenchyma are absent. Primary and secondary vascular bundles differed only slightly in size and in fresh material do not project adaxially or abaxially. Bulliform cells were noted between adjacent vascular bundles. Colorless cells were not observed between vascular bundles, but they do occur adaxially to the primary and secondary bundles, and may be the only cell layer between the epidermis and the secondary vascular bundles. As many as five successive colorless cells (in cross section) were observed adaxially to a secondary vascular bundle on <italic>Ellis 4534</italic>. <xref ref-type="bibr" rid="B13">Metcalfe (1960</xref>: 285) reported only a single bundle sheath in <italic><tp:taxon-name>Leptochloa uniflora</tp:taxon-name></italic> and first noted the significantly enlarged parenchyma sheath cells at 3 and 9 o’clock, which he termed <italic>lateral cells</italic>, which we confirmed for this species (see in particular images with <italic>Ellis 4534</italic> at PRE). The lateral cells often penetrate deeply into laterally adjacent mesophyll and are nearly completely filled by a large chloroplast. Enlarged lateral cells were not clearly evident in the single specimen of <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> examined from limited rehyrdated material so we cannot yet confidently confirm nor reject their presence in this species.</p>
<p><xref ref-type="bibr" rid="B8">Hattersley and Watson (1976</xref>: 303) reported the species as being XyMS-, based on the implied lack of intervening cells between the metaxylem and PCR sheath given the results of <xref ref-type="bibr" rid="B13">Metcalfe (1960)</xref>. <xref ref-type="bibr" rid="B28">Valls (1978</xref>: 73–74) reported the presence of a double sheath for <italic><tp:taxon-name>Leptochloa uniflora</tp:taxon-name></italic> and reported that walls of the inner sheath were “exceptionally thin-walled”. <xref ref-type="bibr" rid="B28">Valls (1978)</xref>, however, may have used specimens incorrectly identified that in actuality were <italic><tp:taxon-name>Leptochloa neesii</tp:taxon-name></italic>. Specimens studied by Roger Ellis, and seen for this study, confirm the observations of <xref ref-type="bibr" rid="B13">Metcalfe (1960)</xref> of the enlarged lateral cells and the presence of a double sheath, supporting the XyMS+ condition. However, the “outer sheath” cells only occur on the distal edges of the enlarged lateral cells, such that colorless cells typically occur adaxially and abaxially to the promixal part of the lateral cell (i.e., that part adjacent to the vascular bundle), with smaller chloroplast-bearing cells bearing smaller and more diffuse chloroplasts adjacent to the part of the lateral cell that occurs closest to the intervascular region. With permission while on site at PRE in 1996, the first author took 35 mm SLR photographs of the original black and white images of <italic>Ellis 4534</italic>, which show the leaf anatomy clearly. However, photos of the original images did not reproduce at a high enough quality to include in this paper. Other images of the leaf anatomy of <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> are unknown to the present authors.</p>
<p><bold>Stem anatomy</bold>. Both species have a solid culm. <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> has inner and outer sclerenchymatous rings, although an inner ring was absent for <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic>.</p>
<p><bold>Lemmatal micromorphology.</bold> The two species share a unique combination of lemmatal micromorphological characters compared to other species of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l. (<xref ref-type="bibr" rid="B22">Snow 1996</xref>). Cork cells and bicellular microhairs were present; macrohairs were terete (not crispate) and obtuse apically (not clavicorniculate; <xref ref-type="bibr" rid="B22">Snow 1996</xref>). Lemmatal characters in many species of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> s.l. but lacking in <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> included silica cells, and papillate long and papillate short cells (<xref ref-type="bibr" rid="B22">Snow 1996</xref>).</p>
<p><bold>Caryopsis morphology</bold>. The caryopsis is trigonous in transverse section and possesses a narrow but deep hilar sulcus (=longitudinally grooved); the pericarp is tightly<!--PageBreak--> adnate (<xref ref-type="bibr" rid="B24">Snow 1998</xref>). This combination of caryopsis characters is unique among genera of chloridoids (<xref ref-type="bibr" rid="B16">Peterson et al. 2012</xref>). In our earlier molecular study <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> is sister to the <italic><tp:taxon-name>Mosdenia</tp:taxon-name></italic>-<italic><tp:taxon-name>Toliara</tp:taxon-name></italic>-<italic><tp:taxon-name>Lopholepis</tp:taxon-name></italic>-<italic><tp:taxon-name>Perotis</tp:taxon-name></italic> clade that lies outside of the subtribe <tp:taxon-name>Eleusininae</tp:taxon-name> (<xref ref-type="bibr" rid="B16">Peterson et al. 2012</xref>). Species of <italic><tp:taxon-name>Mosdenia</tp:taxon-name></italic> and <italic><tp:taxon-name>Perotis</tp:taxon-name></italic> have longitudinally-grooved caryopses that are dorsally or laterally compressed but not trigonous.</p>
</sec>
<sec sec-type="Taxonomic treatment">
<title>Taxonomic treatment</title>
<tp:taxon-treatment>
<tp:nomenclature>
<tp:taxon-name>
<tp:taxon-name-part taxon-name-part-type="genus">Trigonochloa</tp:taxon-name-part>
<object-id xlink:type="simple">http://species-id.net/wiki/Trigonochloa</object-id>
</tp:taxon-name>
<tp:taxon-authority>P.M. Peterson & N. Snow. Ann. Bot. 109: 1327. 2012.</tp:taxon-authority>
</tp:nomenclature>
<tp:treatment-sec sec-type="Type species">
<title>Type species:</title>
<p><italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> (Chipp.) P.M. Peterson & N. Snow.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Description">
<title>Description.</title>
<p>Plants annual to short-lived perennial, sometimes rhizomatous or stoloniferou. Culms (15–)35–130 cm long, terete in cross section, solid, decumbent or clambering to erect; nodes glabrous. Leaf sheaths half as long to slightly longer than internodes, glabrous or ciliate apically along margins; ligules 1–3.5 mm long, membranous, irregularly lacerate with age; leaf blades 1–13(–17) cm long, 0.3−14(–19) mm wide, linear to broadly ovate, flat, typically thin and flaccid, apex acuminate to acute. Panicles17−55 cm long, 2.0–8 cm wide, exserted at maturity, open, narrowly oblong to narrowly elliptic, composed of several to numerous unilateral, secund spikes or spicate racemes scattered along a central axis; rachis semi-terete; branches (1.5−)2–7 cm long, ascending, straight or slightly drooping. Spikelets 1.9−2.8 mm long, 1 (rarely 2-flowered), laterally compressed, subsessile, overlapping; disarticulation above the glumes; glumes 1.8−3.1 mm long, subequal, as long or longer than the floret, subequal, linear to narrowly ovate, 1-nerved, apex acute to acuminate, muronate or emucronate; lemmas 1.2−2.6 mm long, ovate, 3-nerved, thinly membranous to hyaline, minutely hairy along the nerves, apex acute, entire, awnless; paleas 1.5–2.5 mm long, keels ciliolate. Stamens 3. Caryopses 1−1.2 mm long, narrowly elliptic, trigonous in cross section, narrowly but deeply sulcate on the hilar side; surface smooth to slightly rugose-striate, light brown, pericarp fused, tightly adherent to endosperm. 2<italic>n</italic> = 36 for <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> (<xref ref-type="bibr" rid="B7">Gould and Soderstrom 1974</xref>).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Comments">
<title>Comments.</title>
<p>An appropriate common generic name to differentiate <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> from other members of <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic> is “triangle-seed grass”.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Key to species of Trigonochloa">
<title>Key to species of <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic></title>
<table-wrap content-type="key" position="anchor" orientation="portrait">
<table>
<tbody>
<tr>
<td rowspan="1" colspan="1">1</td>
<td rowspan="1" colspan="1">Leaf blades 0.3–4.0 (–5.0) mm wide, more or less linear to narrowly ovate; leaf sheaths margins minutely ciliate towards the apex, the collar never pilose</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic></td>
</tr>
<tr>
<td rowspan="1" colspan="1">–</td>
<td rowspan="1" colspan="1">Leaf blades 5–14 (–19) mm wide, ovate to broadly ovate; leaf sheaths glabrous or sparsely pilose near collar but not ciliate along upper margins</td>
<td rowspan="1" colspan="1"><italic><tp:taxon-name>Trigonochloa uniflor</tp:taxon-name></italic>a</td>
</tr>
</tbody>
</table>
</table-wrap>
</tp:treatment-sec>
</tp:taxon-treatment>
<tp:taxon-treatment>
<tp:nomenclature>
<tp:taxon-name>
<tp:taxon-name-part taxon-name-part-type="genus">Trigonochloa</tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="species">rupestris</tp:taxon-name-part>
<object-id xlink:type="simple">http://species-id.net/wiki/Trigonochloa_rupestris</object-id>
</tp:taxon-name>
<tp:taxon-authority>(C.E. Hubb.) P.M. Peterson & N. Snow. Ann. Bot. 109: 1328. 2012.</tp:taxon-authority>
<xref ref-type="fig" rid="F1">Figure 1A–E</xref>
<tp:nomenclature-citation-list>
<tp:nomenclature-citation><tp:taxon-name>Leptochloa rupestris</tp:taxon-name><comment> C.E. Hubb., Bull. Misc. Inform. Kew: 195. 1941.</comment></tp:nomenclature-citation>
</tp:nomenclature-citation-list>
</tp:nomenclature>
<tp:treatment-sec sec-type="Type">
<title>Type.</title>
<p>SOMALIA. Mt. Wobleh, J.B.Gillett 4981(holotype: K!; isotype: K!)</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Description">
<title>Description.</title>
<p>Plants perennial. Culms 35–85 cm tall, 0.7–1.4 mm wide at base, decumbent or sprawling-stoloniferous to erect, often arising from a thick root crown, branching (often profusely); internodes (0.7–)2.0–5.5(–8.5) cm long, soft to strongly lignified. Leaf sheaths longer or shorter than internodes, glabrous on sides but sometimes minutely papillate at high magnification, margins ciliate (trichomes <1 mm long); collar green; ligules 1.0–2.5 mm long, truncate or obtuse; blades 2.2–5.5(–10.7) cm long, 0.3–4.0(–5.0) mm wide, linear to narrowly ovate, glabrous above, glabrous below but sometimes densely minutely papillate, midrib prominent. Panicles 17–35 cm long, 2.0–4.5 cm wide with 13–66 branches, the branches (1.5–)2.0–4.5 cm long, minutely scabrous, the axils glabrous or at most scabrous and more or less short pilose on the exterior. Spikelets 2.0–2.8 mm long, 1-flowered, nearly sessile or with minute pedicels, more or less imbricate, callus area glabrous; lower glumes 2.4–3.1 mm long, membranous, lanceolate, midnerve scabrous, apex acute to acuminate; upper glumes, 2.2–2.8 mm long, otherwise like lower glumes; lemmas 1.2–2.1 mm long, ovate, light green or nearly white, the lateral nerves faint, sericeous along midnerve (use high magnification), the hair tips rounded, apex acute and awnless; paleas 1.6–2.0 mm long, hyaline, narrowly ovate, glabrous or sparsely sericeous near nerves, apex acute to obtuse; anthers 1.0–1.4 mm long, yellow to brownish green. Caryopses ca. 1.0 mm long and 0.4 mm wide, trigonous in cross section, the surface smooth.</p>
<fig id="F1" position="float" orientation="portrait">
<label>Figure 1.</label>
<caption><p><italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> (C.E. Hubb.) P.M. Peterson & N. Snow <bold>A</bold> Habit <bold>B</bold> Inflorescence <bold>C</bold> Portion of inflorescence branch <bold>D</bold> Spikelet <bold>E</bold> Floret. Drawn from <italic>Wood 2000</italic>.</p></caption>
<graphic xlink:href="PhytoKeys-013-025-g001.jpg" position="float" orientation="portrait" xlink:type="simple"></graphic>
</fig>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Phenology">
<title>Phenology.</title>
<p>Flowering June through January.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Distribution">
<title>Distribution.</title>
<p>This species is found in Yemen and Eritrea south to Kenya in woodlands, hillsides, bushland and on damp rocks along streams; 900–1800 m. [Taxonomic Database Working Group (TDWG): 24: ERI, ETH, SOM; 25: KEN, UGA].</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Conservation status">
<title>Conservation status.</title>
<p>Since many parts of its range are presently inaccessible to botanists or still remain inadequately surveyed this species is data deficient. Additional observations and collections are highly desirable.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Etymology">
<title>Etymology.</title>
<p>The epithet <italic>rupestis</italic> is Latin for “of rocks”, presumably in reference to observations of the habitat of the type collection.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Vernacular name">
<title>Vernacular name.</title>
<p>Somalian triangle-seed grass.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Comments">
<title>Comments.</title>
<p>This species closely resembles <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic>, with which it is morphologically similar. The best character to recognize <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> from <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> is leaf blade width. However, its sprawling, branching, and perennial growth form with narrow culms typically distinguish it from <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic>. The holotype and isotype are aberrant in their lack of ciliate sheath margins, but otherwise accord with the diagnostic characters. The observation of <xref ref-type="bibr" rid="B17">Phillips (1974)</xref> that <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> has more widely divergent leaf blades than <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> cannot be reliably applied to dried material. None of the specimens we<!--PageBreak--><!--PageBreak--> have seen confirmed her observation (<xref ref-type="bibr" rid="B17">Phillips 1974</xref>) that <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> is rhizomatous (i.e., culms within the soil), but the stems can be somewhat sprawling and stoloniferous.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Specimens examined">
<title>Specimens examined.</title>
<p>Eritrea. Donkollo, Schweinfurth 207 (P); Dongollo Presso Ghinda, Pappi 2821 (MO); Monte Dijot, Pappi 2940 (US). Ethiopia. Gamo Gofo: 44 km from Jinka on raod to Konso, ca. 3 km E of K’ey Afer, Gilbert et al. 8954(K); 13 km N of Lante, 29 km N of Arba Minch, Gilbert et al. 8874 (K). Arba Minch, Gilbert et al. 249 (K, MO). Harar:“Rock Valley”, 36 km along the road from Harrar to Jijiga, Amshoff 5520 (B, MO); Errer valley, 22 km SE of Harar on hwy to Djigdjigga, Burger 1162 (K); 7 km from Harar towards Jijiga, Gilbert & Gilbert 1443(K). Sidamo:Between the Genale Doria bridge and the main road Kebre Mengist-Neghelle, on the Biderre track, Friis et al. 1034 (K). Kenya. Rift Valley:West Suk Reserve, 30 mi N of Kitale, Bogdan 3429 (K); West Suk Reserve, 10 mi W of Kapenguria, Suam Riv. Valley, Bogdan 289(K); West Suk, Marech Pass, 40 mi N of Kapenguria, Bogdan 3844(K); 30 mi N of Nakuru, Bogdan 4891 (K, US); Kenya Grassland Research Station, Bogdan AB3964 (P). Somalia. “WOGR near Sheikh”, Wood S/72/95 (K); Jifa Uri, Gillett 4838 (K, US). Uganda.Northern: Moroto Mountains, Karamoja, Napper 1509(K); Warr, Karamoja, Thomas 3176(K). Yemen. Habash, Jebel Melhan, Wood 2848 (BM, K); 2 mi W of Mefhek, Wood Y/75/727 (BM); by Wadi Dur, Udayn, Wood 2000 (K, US).</p>
</tp:treatment-sec>
</tp:taxon-treatment>
<tp:taxon-treatment>
<tp:nomenclature>
<tp:taxon-name>
<tp:taxon-name-part taxon-name-part-type="genus">Trigonochloa</tp:taxon-name-part>
<tp:taxon-name-part taxon-name-part-type="species">uniflora</tp:taxon-name-part>
<object-id xlink:type="simple">http://species-id.net/wiki/Trigonochloa_uniflora</object-id>
</tp:taxon-name>
<tp:taxon-authority>(Hochst. ex A. Rich.) P. M. Peterson & N. Snow. Ann. Bot. 109: 1328. 2012.</tp:taxon-authority>
<xref ref-type="fig" rid="F2">Figure 2A–H</xref>
<tp:nomenclature-citation-list>
<tp:nomenclature-citation><tp:taxon-name>Leptochloa uniflora</tp:taxon-name><comment> Hochst. ex A. Rich., Tent. Fl. Abyss. 2: 409. 1851.</comment></tp:nomenclature-citation>
<tp:nomenclature-citation><tp:taxon-name>Craspedorhachis uniflora</tp:taxon-name><comment> (Hochst. ex A. Rich) Chippind., Grasses and Pastures of South Africa 205, f. 182. 1955.</comment></tp:nomenclature-citation>
<tp:nomenclature-citation><tp:taxon-name>Cynodon gracilis</tp:taxon-name><comment> Nees ex Steud., Syn. Pl. Glumac. 1: 213. 1854. TYPE: India, Ab. loco. Wight Herbarium 8895 (lectotype: K! designated here, no specimen number given in the protologue; isotype: K!).</comment></tp:nomenclature-citation>
<tp:nomenclature-citation><tp:taxon-name>Agrostis montana</tp:taxon-name><comment> Krock., Fl. Siles. 1: 110. 1787. <italic><tp:taxon-name>Agrostis montana</tp:taxon-name></italic> Rottl. ex Hook. f., Fl. Brit. Ind. 7: 298. 1896, nom. inval. TYPE: India, Tinnevelly; at Palamcotta, 28 Nov 1895, Rottler s.n. (lectotype: K! designated here).</comment></tp:nomenclature-citation>
<tp:nomenclature-citation><tp:taxon-name>Craspedorhachis menyharthii</tp:taxon-name><comment> Hack. ex Schinz, Bull. Herb. Boissier, ser. 2, 1: 770. 1901. TYPE: Mozambique, Boruma, Tanuar, L. Menyharth 665 (lectotype: Z!, designated by <xref ref-type="bibr" rid="B17">Phillips 1974</xref>: 277 [who did not include the collection number]; duplicate of lectotype: W!).</comment></tp:nomenclature-citation>
<tp:nomenclature-citation><tp:taxon-name>Leptochloa laurentii</tp:taxon-name><comment> De Wild., Miss. Em. Laurent i. 207. 1906. TYPE: Democratic Republic of the Congo, Kiri, E. Laurent s.n. (holotype: BR! [seen digitally, June 2012; barcode BR0000008761873]).</comment></tp:nomenclature-citation>
</tp:nomenclature-citation-list>
</tp:nomenclature>
<tp:treatment-sec sec-type="Type">
<title>Type:</title>
<p>Ethiopia, In valle fluvi Tacaze, Schimper 1707(holotype: P!; isotypes: B!, BM!, GH!, K!, MO!, PRE!, S!, W!, photo ex W!)</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Description">
<title>Description.</title>
<p>Plants annual (or possibly weakly perennial). Culms (15–)45–130 cm tall, 0.6–2.0(3.0) mm wide at base, generally erect, sometimes geniculate below and stoloniferous by rooting at the nodes, sometimes branching, arising from fibrous roots or occasionally from a short knotted-rhizome; internodes 2–8 cm long, soft, solid. Leaf sheaths mostly shorter than internodes, glabrous throughout or rarely sparsely pilose near the collar, the margins glabrous; collars green; ligules 1.5–3.5 mm long, broadly obtuse, lacerate; blades 1–13(–17) cm long, 5–14(–19) mm wide, ovate to broadly ovate, glabrous above and below, midrib prominent or not. Panicles 25–55 cm long, 5–8 cm wide with 22–90 branches, the branches 2.5–7.0 cm long, minutely scabrous, the axils mostly glabrous internally but short pilose on external side. Spikelets 1.9–2.7 mm long, 1-flowered or rarely 2-flowered, but if so, only a few per plant, nearly sessile or with minute pedicels less than 0.3 mm long, somewhat imbricate, callus area glabrous; lower glumes 1.8–2.3 mm long, narrowly triangular, minutely scabrous on midnerve, apex acuminate to mucronate; upper glumes 2.2–2.6 mm long, otherwise like lower glumes; lemmas 1.6–2.6 mm long, ovate, whitish or light green, the lateral nerves very faint, sparsely pubescent along nerves, apex awnless; paleas 1.5–2.5 mm long, subequal to lemma, narrowly ovate, glabrous, apex obtuse or sometimes acute; anthers ca. 1 mm long, dark purple to pale olive green. Caryopses ca. 1.2 mm long and 0.4 mm wide, trigonous in cross section, the surface smooth to slightly rugose–striate. 2<italic>n</italic> = 36.</p>
<fig id="F2" position="float" orientation="portrait">
<label>Figure 2.</label>
<caption><p><italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> (Hochst. ex A. Rich.) P.M. Peterson & N. Snow <bold>A</bold> Habit <bold>B</bold> Inflorescence <bold>C</bold> Portion of inflorescence branch <bold>D</bold> Spikelet <bold>E</bold> Floret <bold>F</bold> Caryopsis, dorsal view <bold>G</bold> Caryopsis, ventral view <bold>H</bold> Caryopsis, cross section. <bold>A–C, F–G</bold> drawn from <italic>Ballard 1489</italic>; <bold>D, E</bold> drawn from <italic>Chare 4434</italic>.</p></caption>
<graphic xlink:href="PhytoKeys-013-025-g002.jpg" position="float" orientation="portrait" xlink:type="simple"></graphic>
</fig>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Phenology">
<title>Phenology.</title>
<p>Flowering throughout the year when adequate moisture is available.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Distribtution">
<title>Distribtution.</title>
<p>This species isscattered through the eastern and southern portions of sub-Saharan Africa, rarely in India, most common in Sri Lanka in forests and shady areas on hillsides, well-drained and often sandy soils in disturbed and riparian areas; 0-1200 m. [TDWG: 22: GHA, NGR, ZAI (Dem. Rep. Congo); 25: KEN, TAN; 26: ANG, MLW, MOZ, ZIM; 27: BOT, NAM, NAT, TVL; 40: IND, SRL].</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Conservation status">
<title>Conservation status.</title>
<p>Since the species is widespread it is of least concern (<xref ref-type="bibr" rid="B9">IUCN 2010</xref>) given its widespread occurrence. However, the typical size of populations is undocumented. The thinly membranous leaves likely are sought after by grazers, although the relative nutrition content of the leaves is unstudied.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Etymology">
<title>Etymology.</title>
<p>The specific epithet is from the Latin <italic>uniflora</italic>, with reference to the single floret per spikelet.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Vernacular names">
<title>Vernacular names.</title>
<p>Common triangle-seed grass. Kenya: Mkuse - Digo (Magogo and Glover 477, W).</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Comments">
<title>Comments.</title>
<p>The species description and distribution differ from <xref ref-type="bibr" rid="B23">Snow (1997)</xref> because some specimens therein were incorrectly identified as <italic><tp:taxon-name>Leptochloa neesii</tp:taxon-name></italic>.</p>
<p><italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> individuals vary significantly in growth habit. Most specimens are relatively delicate, sprawling annuals, but some specimens have more erect, relatively stout culms that appear to be weakly perennial. The more erect forms typically occur in somewhat more open areas and have thicker leaves, whereas the more slender forms that frequently root at the lower nodes, typically occur in shade and have thinner leaves.</p>
<p>The glumes of <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> typically are longer than the single floret and may be mucronate. The caryopsis is sometimes dispersed with a tightly adnate lemma and palea,<!--PageBreak--><!--PageBreak--> which may enhance water absorption prior to germination. The apex of the leaf sheath can sometimes be sparsely pilose, whereas in contrast nearly all specimens of <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> have ciliate sheath margins. Many specimens of <italic><tp:taxon-name>Trigonochloa uniflora</tp:taxon-name></italic> from Sri Lanka resemble <italic><tp:taxon-name>Trigonochloa rupestris</tp:taxon-name></italic> based on thin, sprawling culms. Two culms on Exell et al. 1060 (BM) have a sparse covering of papillose hairs on the upper and lower surfaces of the leaf blades.</p>
<p>Two counts of 2<italic>n</italic> = 40 ascribed to <italic><tp:taxon-name>Trigonochloa</tp:taxon-name></italic> (as <italic><tp:taxon-name>Leptochloa</tp:taxon-name></italic>) <italic>uniflora</italic> by <xref ref-type="bibr" rid="B7">Gould and Soderstrom (1974)</xref> were misidentifications of <italic><tp:taxon-name>Leptochloa neesii</tp:taxon-name></italic> (vouchers at US!). We confirm the voucher with apolyploid count of 2<italic>n</italic> = 36 based on <italic>x</italic> = 9 for <italic><tp:taxon-name>Trigonochloa (Leptochloa) uniflora</tp:taxon-name></italic> (Soderstrom and Kulatunge 1753, US!). In contrast, the count of <italic>n</italic> = 10 by <xref ref-type="bibr" rid="B14">Olorode (1975)</xref> has not been confirmed with a voucher.</p>
</tp:treatment-sec>
<tp:treatment-sec sec-type="Specimens examined">
<title>Specimens examined.</title>
<p>Angola. Cuanza Norte: Cazengo, Welwitsch 2981 (BM, K); Cazengo, Gossweiler 4421 (BM, K); Granja de S. Luiz, Gossweiler 5200 (BM). Ab. loco, Gossweiler 2966(K) and 5444 (BM). Botswana. North-West: Riparian woodland, near Kasane,Blair Rains 67 (K, US).Dem. Repub. Congo.Haut-Katanga: Kibula, Callens 4776 (PRE). Kivu:Entre Nyangwe et Malela, Lebrun 5971 (PRE). Kongo Central: Kisantu, Vanderyst s.n. (US 889080). Tshuapa: Mpangu, Delhaye 440 (K). Province unknown: Gona, Vanderyst 5682 (US). Ghana. Ashanti: New Tafo, Lovi 3909 (K). Eastern:Aburi, Deighton 613 (BM). India. Kerala:“Palghat” [Palakkad], Madras Herbarium/South Indian Flora 16320 (US). Periakulam:Madurai, Matthew & Charles 51410 (K). Kenya. Coast: Longo Mwagandi Area, Shimba Hills, Magogo and Glover 477 (W); 50 mi SW of Mombasa, Shimba Hills, Bogdan AB3910 (P);Mombasa woodlands, Gibon s.n. (US 2954368); Kaya Muhaka, Luke 3405 (K); Forest between Umba and Mwena Rivers on Lungalunga-Msambweni Rd., Drummond & Hemsley 3787 (K, P). Malawi. Central:Dedza Distr., Mua-Livulezi Forest Reserve, Exell et al. 1060 (BM). Southern: Shire Valley, Hall-Martin 438 (MO); Lengwe Game Reserve, Hall-Martin 494 (K) and 582(K); Lengwe Game Reserve, NE corner, Brummitt 8884(K). Province Unknown: Mwenikumbila foothills, Jackson 1175 (MO). Mozambique. Manica:Maribane Forest, Chare 4434 (US); Amatongas Forest, near Gondola, Schweicherdt 272 (US). Sofala: Gorongosa N.P., Sangarassa Forest, 1 km W of Chitenga, Tinley 2497(K); Amatongas Forest, Schweicherdt 2341 (K, US). Zambezia: Malema, Torre & Paiva 11192 (PRE); Arredores de Mocuba, Torre 4908 (K). Namibia. Province unconfirmed. Mpilia Island, Killich & Leistner 3347 (K). Nigeria. Ondo: Idanre, Brenan & Jones 8731 (K). South Africa. Kwazulu-Natal:Mkuze Game Reserve, Ellis 3635 (PRE); Mkuze Game Reserve, parking lot by Bube (Kubube) Hide, Snow et al. 6978(MO, PRE); Tembe Elephant Park, Ward 1382 (PRE). Limpopo:Kruger N.P., Punda Milia area, Shipudza valley east of Punda Milia near Dongadziba, Ellis 3226 (K, PRE); Kruger N.P., ca. 12 km NW of Punda Milia, Davidse & Ellis 5925 (K, MO). Sri Lanka. Anuradhapura: Mihintale, Soderstrom & Kulatunge 1715 (CANB, K, TAES, US); Ritigala Strict Natural Reserve, ascent along eastern slope of Wannatikianda, Jayasuriya 1058(K, US). Central:54 miles N of Kandy toward Anuradhapura, trunk road A-9, marker 54/2, Gould 13250(US); Dambulla, Trimens 28(US); as previous, Ashton 998(K, US); Ruhuna N.P., Block I, Cooray 69030805R(US); Ruhuna N.P., <!--PageBreak-->Block I, Patanagala Camp, Clayton 5924 (CANB, K, TAES, US); Kumbukkan Oya, ca. 2 mi above mouth, at Megahakanda Meda Duwa Block 2, M–d Plot, R 16, Fosberg 51099 (US); Ruhuna N.P., Rugamtota on Menik Ganga (Plot 31), Fosberg & Mueller-Dombois 50192(US); Mennik Ganga (Riv.) 1 mile above Yalu Bungalow, Fosberg et al. 51045(US); Ruhuna N.P., Patanagala, Cooray 69120212R(K, US); Ruhuna N.P., Block 2, Cooray 69010502R(K, US); Ruhuna N.P., Block I, Rugamtota, Mueller-Dombois 69030704(US); Ruhuna N.P., Block I, next to Yala Camp site, Mueller-Dombois & Cooray 68013006(US); Ruhuna N.P., Block 1, in plot R13 between Andunoruwa and Komawa Wawa, Mueller-Dombois 69010713(US). Puttalam:Wilpattu N.P., Marai villu, Clayton 5597 (CANB, K, TAES, US). Trincomalee:Kantalai; road between Trincomalee and Kandy, Soderstrom & Kulantunge 1753 (CANB, K, TAES, US); Kantalai, 25 miles from Brincomalle on Kandy Rd., Ballard 1489(US). Tanzania. Iringa Region:Ab. loco, Greenway et al. 14075 (MO). Lindi Region:40 km W Lindi, Schlieben 5879 (B, BM, M, MO, S, US); Tendaguru, Migeod 104 (BM) and Migeod 126 (BM). Mahenge Region:Umgebung der Station Mahenge, Schlieben 1721 (BM, M, S). Morogoro Region:Uluguru-Gebirge, Schlieben 3630 (B, BM, G, M, S); 3 mi N Tunuguo, 30 mi SE Morogoro, Boaler 625 (B, US [2 sheets]). Province unknown.Rukwa Valley, Vuma Riv. near Zimba, Siame 581 (MO). Zambia. Central:Iolanda, N bank of R. Kafue, near Kafue town, Robinson 6440 (B, K, M); Kafue N.P., Musa-Kafue conflence, Mitchell 6/75(K). Eastern: Chikwa, ca 50 mi NW of Lundazi in Luangwa Valley, Robinson 822 (K, M). Northern:M’fume Camp, Verboom 922 (BM, K); Mporokoso,Lake Mweru-Wantipa,Richards 9117 (K, NY). Southern:Siburu forest, Sekute Chieftancy, Bainbridge 709 (BM); Victoria Falls, Crook 52602(K); Victoria Falls-Livingstone Island, Ellis 2780 (K, MO). Province unconfirmed. Kafue N.P., Mitchell 24/46 (B). Zimbabwe. Manicaland:Tanganda Tea Estate, Chipinga [now Chipinge], Brain 10615(K). Mashonaland West:Eastern Urungwe [=Hurungwe], tributary of upper Mauora, Phipps 868 (K). Urungwe,Chirundu, Simon 706 (BM). Masvingo: Bikita, 5 km E of Moodie Pass, Davidse et al. 6643 (BRI, K, MO, US). Matabeleleland North: Wankie [now Hwange], Kandahar Fishing Camp, Martin 87(K). Midlands: Gokwe, Sengua Research Station, Guy 2391(K).</p>
</tp:treatment-sec>
</tp:taxon-treatment>
</sec>
</body>
<back>
<ack>
<title>Acknowledgements</title>
<p>This article derives from work done for a doctoral dissertation at Washington University in St. Louis (Snow 1997) and two Short-term visitor grants from the Smithsonian Institution in 1989 and 2007. The first author also was supported by National Science Foundation (BIR-9256779), The Explorer’s Club, Grants-in-Aid of Research (Sigma Xi), Andrew W. Mellon Foundation, Missouri Botanical Garden, and American Society of Plant Taxonomists. Fieldwork in southern Africa was supported by the National Geographic Society; the first author thanks P. Burgoyne, M. Carranza, M.<!--PageBreak--> Chatakuta, L. Cohen, J. Gumbi, R. Roux, and O. McKibbin for assistance. Cabelas Corporation of Sidney, Nebraska generously donated field equipment for a collecting trip to Africa and Australia in 1996. We thank the following herbaria for loans or permission to view specimens: B, BM, BRI, CANB, K, M, MO, NY, P, PRE, S, TAES, US, W, Z. Appreciation is extended to Alice R. Tangerini for providing excellent new drawings, and to Dr. L. Versieux and two anonymous reviewers for comments.</p>
</ack>
<ref-list>
<title>References</title>
<ref id="B1"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Bentham</surname> <given-names>G</given-names></name></person-group> (<year>1882</year>)<article-title> Notes on Gramineae.</article-title> <source>Journal of the Linnean Society, Botany </source> <volume>18</volume>: <fpage>14</fpage>-<lpage>134</lpage>.</mixed-citation></ref>
<ref id="B2"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Brummitt</surname> <given-names>RK</given-names></name></person-group> (<year>2001</year>)<source> World geographical scheme for recording plant distributions. Edition 2. International Working Group on Taxonomic Databases (TDWG). XV, 153. <ext-link ext-link-type="uri" xlink:href="http://grassworld.myspecies.info/sites/grassworld.myspecies.info/files/tdwg_geo2.pdf" xlink:type="simple">http://grassworld.myspecies.info/sites/grassworld.myspecies.info/files/tdwg_geo2.pdf</ext-link></source></mixed-citation></ref>
<ref id="B3"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Chippindale</surname> <given-names>LKA</given-names></name></person-group> (<year>1955</year>)<source> A guide to the identification of grasses in South Africa.</source> <publisher-name>In: Meredith D (Ed) The Grasses and Pastures of South Africa, Central News Agency</publisher-name>, <publisher-loc>South Africa</publisher-loc>, <lpage>527</lpage> pp.</mixed-citation></ref>
<ref id="B4"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Clayton</surname> <given-names>WD</given-names></name><name name-style="western"><surname>Renvoize</surname> <given-names>SA</given-names></name></person-group> (<year>1986</year>)<article-title> Genera Graminum: Grasses of the World.</article-title> <source>Kew Bulletin Additional Series </source> <volume>13</volume>: <fpage>1</fpage>-<lpage>389</lpage>.</mixed-citation></ref>
<ref id="B5"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Columbus</surname> <given-names>JT</given-names></name><name name-style="western"><surname>Cerros-Tlatilpa</surname> <given-names>R</given-names></name><name name-style="western"><surname>Kinney</surname> <given-names>MS</given-names></name><name name-style="western"><surname>Siqueiros-Delgado</surname> <given-names>ME</given-names></name><name name-style="western"><surname>Bell</surname> <given-names>HL</given-names></name><name name-style="western"><surname>Griffith</surname> <given-names>MP</given-names></name><name name-style="western"><surname>Refulio-Rodriguez</surname> <given-names>NF</given-names></name></person-group> (<year>2007</year>)<article-title> Phylogenetics of Chloridoideae (Gramineae): A preliminary study based on nuclear ribosomal internal transcribed spacer and chloroplast <italic>trnL-F</italic> sequences.</article-title> <source>Aliso </source> <volume>23</volume>: <fpage>565</fpage>-<lpage>579</lpage>.</mixed-citation></ref>
<ref id="B6"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Gibbs-Russell</surname> <given-names>GE</given-names></name><name name-style="western"><surname>Watson</surname> <given-names>L</given-names></name><name name-style="western"><surname>Koekemoer</surname> <given-names>M</given-names></name><name name-style="western"><surname>Smook</surname> <given-names>L</given-names></name><name name-style="western"><surname>Barker</surname> <given-names>NP</given-names></name><name name-style="western"><surname>Anderson</surname> <given-names>HM</given-names></name><name name-style="western"><surname>Dallwitz</surname> <given-names>MJ</given-names></name></person-group> (<year>1991</year>)<article-title> Grasses of Southern Africa. Reprinted edition. Memoirs of the Botanical Survey of South Africa No.</article-title> <volume>58</volume>: <fpage>1</fpage>-<lpage>437</lpage>.</mixed-citation></ref>
<ref id="B7"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Gould</surname> <given-names>FW</given-names></name><name name-style="western"><surname>Soderstrom</surname> <given-names>TR</given-names></name></person-group> (<year>1974</year>)<article-title> Chromosome numbers of some Ceylon grasses.</article-title> <source>Canadian Journal of Botany </source> <volume>52</volume>: <fpage>1075</fpage>-<lpage>1090</lpage>. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1139/b74-136" xlink:type="simple">10.1139/b74-136</ext-link></mixed-citation></ref>
<ref id="B8"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Hattersley</surname> <given-names>PW</given-names></name><name name-style="western"><surname>Watson</surname> <given-names>L</given-names></name></person-group> (<year>1976</year>)<article-title> C<sub>4</sub> grasses: an anatomical criterion for distinguishing between NADP-malic enzyme species and PCK or NAD-malic enzyme species.</article-title> <source>Australian Journal of Botany </source> <volume>24</volume>: <fpage>297</fpage>-<lpage>308</lpage>. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1071/BT9760297" xlink:type="simple">10.1071/BT9760297</ext-link></mixed-citation></ref>
<ref id="B9"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>IUCN</surname></name></person-group> (<year>2010</year>)<source> IUCN Red List of Threatened Species. Version 2010.3.</source></mixed-citation></ref>
<ref id="B10"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Jacobs</surname> <given-names>SWL</given-names></name></person-group> (<year>1987</year>)<article-title> Systematics of the chloridoid grasses.</article-title> In: <person-group><name name-style="western"><surname>Soderstrom</surname> <given-names>TR</given-names></name><name name-style="western"><surname>Hilu</surname> <given-names>KW</given-names></name><name name-style="western"><surname>Campbell</surname> <given-names>CS</given-names></name><name name-style="western"><surname>Barkworth</surname> <given-names>ME</given-names></name></person-group> (<role>Eds</role>). <issue-title>Grass Systematics and Evolution, Smithsonian Institution Press, Washington, D.</issue-title> <source>C.</source>: <fpage>277</fpage>-<lpage>286</lpage>.</mixed-citation></ref>
<ref id="B11"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Judziewicz</surname> <given-names>EJ</given-names></name></person-group> (<year>2009</year>)<article-title> <italic>Toliara</italic> (Poaceae, Chloridoideae, Cynodonteae), a new grass genus endemic to southern Madagascar.</article-title> <source>Adansonia </source> <volume>31</volume>: <fpage>273</fpage>-<lpage>277</lpage>. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.5252/a2009n2a4" xlink:type="simple">10.5252/a2009n2a4</ext-link></mixed-citation></ref>
<ref id="B12"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>McNeill</surname> <given-names>J</given-names></name></person-group> (<year>1979</year>)<article-title> <italic>Diplachne</italic> and <italic>Leptochloa</italic> in North America.</article-title> <source>Brittonia </source> <volume>31</volume>: <fpage>399</fpage>-<lpage>404</lpage>. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.2307/2806134" xlink:type="simple">10.2307/2806134</ext-link></mixed-citation></ref>
<ref id="B13"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Metcalfe</surname> <given-names>CR</given-names></name></person-group> (<year>1960</year>)<source> Anatomy of the Monocotyledons. I. Gramineae. Clarendon Press, Oxford.</source></mixed-citation></ref>
<ref id="B14"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Olorode</surname> <given-names>O</given-names></name></person-group> (<year>1975</year>)<article-title> Additional chromosome counts in Nigerian grasses.</article-title> <source>Brittonia </source> <volume>27</volume>: <fpage>63</fpage>-<lpage>68</lpage>. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.2307/2805647" xlink:type="simple">10.2307/2805647</ext-link></mixed-citation></ref>
<!--PageBreak-->
<ref id="B15"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Peterson</surname> <given-names>PM</given-names></name><name name-style="western"><surname>Romaschenko</surname> <given-names>K</given-names></name><name name-style="western"><surname>Johnson</surname> <given-names>G</given-names></name></person-group> (<year>2010</year>)<source> A classification of the Chloridoideae (Poaceae) based on multi-gene phylogenetic trees. Molecular Phylogenetics and Evolution 55: 580–598. <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1016/j.ympev.2010.01.018" xlink:type="simple">http://dx.doi.org/10.1016/j.ympev.2010.01.018</ext-link></source></mixed-citation></ref>
<ref id="B16"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Peterson</surname> <given-names>PM</given-names></name><name name-style="western"><surname>Romaschenko</surname> <given-names>K</given-names></name><name name-style="western"><surname>Snow</surname> <given-names>N</given-names></name><name name-style="western"><surname>Johnson</surname> <given-names>G</given-names></name></person-group> (<year>2012</year>)<article-title> A molecular phylogeny and classification of <italic>Leptochloa</italic> (Poaceae: Chloridoideae: Chlorideae) <italic>sensu lato</italic> and related genera.</article-title> <source>Annals of Botany </source> <volume>109</volume>: <fpage>1317</fpage>-<lpage>1329</lpage>. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.1093/aob/mcs077" xlink:type="simple">10.1093/aob/mcs077</ext-link></mixed-citation></ref>
<ref id="B17"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Phillips</surname> <given-names>SM</given-names></name></person-group> (<year>1974</year>)<article-title> Studies in the Gramineae: XXXV.</article-title> <source>Kew Bulletin </source> <volume>29</volume>: <fpage>267</fpage>-<lpage>270</lpage>. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.2307/4108540" xlink:type="simple">10.2307/4108540</ext-link></mixed-citation></ref>
<ref id="B18"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Phillips</surname> <given-names>SM</given-names></name></person-group> (<year>1982</year>)<article-title> A numerical analysis of the Eragrostideae (Gramineae).</article-title> <source>Kew Bulletin </source> <volume>37</volume>: <fpage>133</fpage>-<lpage>162</lpage>. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.2307/4114733" xlink:type="simple">10.2307/4114733</ext-link></mixed-citation></ref>
<ref id="B19"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Phillips</surname> <given-names>SM</given-names></name></person-group> (<year>1995</year>)<source> Poaceae (Gramineae). In: Hedberg I, Edwards S (Eds) Flora of Ethiopia and Eritrea, vol. 7, Addis Ababa University and Uppsala University, Ethiopia, 420pp.</source></mixed-citation></ref>
<ref id="B20"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Richard</surname> <given-names>A</given-names></name></person-group> (<year>1851</year>)<source> Tentamen Florae Abyssinicae, Apud Arthus Bertrand, Paris, 518pp.</source></mixed-citation></ref>
<ref id="B21"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Simon</surname> <given-names>BK</given-names></name><name name-style="western"><surname>Clayton</surname> <given-names>WD</given-names></name><name name-style="western"><surname>Harman</surname> <given-names>KT</given-names></name><name name-style="western"><surname>Vorontsova</surname> <given-names>M</given-names></name><name name-style="western"><surname>Brake</surname> <given-names>I</given-names></name><name name-style="western"><surname>Healy,</surname> <given-names>D</given-names></name><name name-style="western"><surname>Alfonso</surname> <given-names>Y</given-names></name></person-group> (<year>2011</year>)<source> GrassWorld. <ext-link ext-link-type="uri" xlink:href="http://grassworld.myspecies.info/" xlink:type="simple">http://grassworld.myspecies.info/</ext-link> [accessed 25 February 2012]</source></mixed-citation></ref>
<ref id="B22"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Snow</surname> <given-names>N</given-names></name></person-group> (<year>1996</year>)<article-title> The phylogenetic utility of lemmatal micromorphological characters in <italic>Leptochloa</italic> and related genera in subtribe Eleusininae (Poaceae, Chloridoideae, Eragrostideae).</article-title> <source>Annals of the Missouri Botanical Garden </source> <volume>83</volume>: <fpage>504</fpage>-<lpage>529</lpage>. doi: <ext-link ext-link-type="uri" xlink:href="http://dx.doi.org/10.2307/2399991" xlink:type="simple">10.2307/2399991</ext-link></mixed-citation></ref>
<ref id="B23"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Snow</surname> <given-names>N</given-names></name></person-group> (<year>1997</year>)<source> Phylogeny and systematics of <italic>Leptochloa</italic> P. Beauv. sensu lato (Poaceae, Chloridoideae, Eragrostideae). Ph.D. Dissertation, Washington University, St. Louis, Missouri.</source></mixed-citation></ref>
<ref id="B24"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Snow</surname> <given-names>N</given-names></name></person-group> (<year>1998</year>)<article-title> Caryopsis morphology of <italic>Leptochloa</italic> sensu lato(Poaceae, Chloridoideae).</article-title> <source>Sida </source> <volume>18</volume>: <fpage>271</fpage>-<lpage>282</lpage>.</mixed-citation></ref>
<ref id="B25"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Snow</surname> <given-names>N</given-names></name></person-group> (<year>2003</year>)<source> 17.19 <italic>Leptochloa</italic> P. Beauv. In: Barkworth ME, Capels KM, Long S, Piep MB (Eds) Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Flora of North America North of Mexico, Oxford University Press, New York, 51−60.</source></mixed-citation></ref>
<ref id="B26"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Soreng</surname> <given-names>RJ</given-names></name><name name-style="western"><surname>Davidse</surname> <given-names>G</given-names></name><name name-style="western"><surname>Peterson</surname> <given-names>PM</given-names></name><name name-style="western"><surname>Zuloaga</surname> <given-names>FO</given-names></name><name name-style="western"><surname>Judziewicz</surname> <given-names>EJ</given-names></name><name name-style="western"><surname>Filgueiras</surname> <given-names>TS</given-names></name><name name-style="western"><surname>Morrone</surname> <given-names>O</given-names></name><name name-style="western"><surname>Romaschenko</surname> <given-names>K</given-names></name></person-group> (<year>2012</year>)<source> A World-wide Phylogenetic Classification of Poaceae (Gramineae): capim, çimen, çayır, darbha, ghaas, ghas, gramas, gräser, grasses, he ben ke, hullu, kasa, kusa, pastos, pillu, pullu, zlaki, etc. <ext-link ext-link-type="uri" xlink:href="http://www.tropicos.org/projectwebportal.aspx?pagename=ClassificationNWG&projectid=10" xlink:type="simple">http://www.tropicos.org/projectwebportal.aspx?pagename=ClassificationNWG&projectid=10</ext-link></source></mixed-citation></ref>
<ref id="B27"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Thiers</surname> <given-names>B</given-names></name></person-group> (<year>2012</year>)<source> Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. <ext-link ext-link-type="uri" xlink:href="http://sweetgum.nybg.org/ih/" xlink:type="simple">http://sweetgum.nybg.org/ih/</ext-link></source></mixed-citation></ref>
<ref id="B28"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Valls</surname> <given-names>JFM</given-names></name></person-group> (<year>1978</year>)<source> A biosystematic study of <italic>Leptochloa</italic> with special emphasis on <italic>Leptochloa dubia</italic> (Gramineae: Chloridoideae). Ph.D. Dissertation, Texas A&M University, College Station.</source></mixed-citation></ref>
<ref id="B29"><mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Watson</surname> <given-names>L</given-names></name><name name-style="western"><surname>Dallwitz</surname> <given-names>MJ</given-names></name></person-group> (<year>1992</year>)<source> The Grass Genera of the World. CAB International, Wallingford, UK.</source></mixed-citation></ref>
</ref-list>
</back>
</article>