Schoenus cuspidatus Rottb., Desc. Icon. Rar. Pl.

Elliott, T. L. & Muasya, A. M., 2020, A taxonomic revision of Schoenus cuspidatus and allies (Cyperaceae, tribe Schoeneae) - Part 2, South African Journal of Botany 130, pp. 327-347 : 339-341

6. Schoenus cuspidatus Rottb., Desc. Icon. Rar. Pl. : 66 (1773).

Chaetospora cuspidata Nees, Linnaea 7: 529 (1832).

Elynanthus cuspidatus (Rottb.) Nees, Linnaea 9: 298 (1834). Elynanthus cuspidatus (Rottb.) Nees var. setiformis Nees, Linnaea 10: 188 — 189 (1835).

Tetraria cuspidata (Rottb.) C.B.Clarke in T.A. Durand and H. Schinz, Consp. Fl. Afric. 5: 660 (1894).

Type: South Africa, without date, without collector C10010615 (C, lecto. — image!, here designated).

[Note: It is likely that drawing of S. cuspidatus labelled ‘Tab XVIII, Fig. 3 ’ in RottbØll (1773) is based on the specimen at C, because the stamp on the reverse side of the specimen indicates that it is from RottbØll’ s personal herbarium, and there is a reference to the figure in RottbØll (1773) on the reverse side of the specimen that matches with his handwriting. Based on the age of the specimen and our knowledge of early botanical exploration in South Africa (see Glen and Germishuizen, 2010), we think that this specimen probably originated from Thunberg’ s collection. Furthermore, we also think that this specimen probably originated in the eastern areas of the Western Cape Province or adjacent areas of the Eastern Cape Province of South Africa based on Thunberg’ s itinerary and the morphological characters of the specimen.].

[Note: Nees von Esenbeck (1834) lists E. cuspidatus as one of the specimens of Elynanthus that he examined. Since he indirectly refers to [ Schoenus ] cuspidatus in this list, this is a valid publication of E. cuspidatus .].

Scirpus [as Schoenus ] cuspidatus Roem. and Schult. in J.J. Roemer and J.A. Schultes, Syst. Veg., ed. 15 bis 2: 67 (1817). [Note: In their description of Schoenus cuspidatus, Roemer and Schultes (1817, p. 67) incorrectly cited the species as ‘ Scirpus cuspidatus ’ Rottboell. It appears that they accidently swapped the genus name Scirpus for Schoenus in this description.].

Elynanthus gracilis Nees, Linnaea 10: 189 (1835), pro parte. non Linnaea 9: 298 (1834), nom. nud. Tetraria cuspidata (Rottb.) C.B.Clarke f. gracilis (Nees) K uk €., Repert. Spec. Nov. Regni. Veg. 29: 187 (1931). Type: South Africa, without date, Sieber 108 (K, lecto.! [K000244957], here designated; BM!, H[H1565908] — image!, K[K000244956] — image!, TUB[TUB007326] — image!, isolecto.). [Note: K ukenthal € (1931) states that the authority of Tetraria cuspidata (Rottb.) C.B.Clarke f. gracilis is “(Nees) Boeckeler”, but we think that he was mistaken, because Boeckeler only mentions E. gracilis has a ‘ forma minor ’ ( Boeckeler 1874b, p. 259) and does not explicitly designate g racilis as a ‘ forma ’.].

[Note: Confusion exists whether E. gracilis should be a synonym of S. cuspidatus or Schoenus exilis (Levyns) T.L.Elliott & Muasya , or given species status. For example, recent treatments have placed it as a synonym of S. cuspidatus ( Clarke, 1900, p. 278) , a forma of S. cuspidatus (K ukenthal €, 1931, p. 187) and a synonym of S. exilis ( Levyns, 1947, p. 82) . Based on the characters of Sieber 108, which is one of the specimens cited by ( Nees von Esenbeck 1835b, p. 189), we have decided to maintain this name as a synonym of S. cuspidatus . More detailed genetic work including intensive sampling of the different S. cuspidatus variants across their distributional range will be required to resolve this taxonomic uncertainty.].

Schoenus aristatus Nees ex Kunth, Enum. Pl. 2: 309 (1837), pro syn.

Schoenus fl exuosus Steud. ex Kunth, Enum. Pl. 2: 309 (1837), pro syn.

Caespitose, phyllopodic perennial graminoid with considerable vegetative and reproductive variation. Culms terete, (225 —)354 — 527 (— 935) X (0.3 —)0.6 — 0.8(— 1.1) mm ( Fig. 4A ). Leaves basal, usually less than half the height of culm and sometimes very short, 1 — 5, (11 —) 51 — 125(— 295) X 0.2‒0.4(— 0.8) mm, usually proximally channelled, margin usually serrate above sheath, often lax and partially curled. Sheaths reddish-black at base but paler above, firm, longitudinally striate. Ligule firm, length variable, (0.2 —)0.4 — 0.7(— 1.9) mm long. In fl orescence a pseudolateral panicle, dark reddish-brown in colour, (18 —)25‒32(— 58) X (2.0 —)4.5‒8.0(— 18.0) mm, proximal rachis length (4.5 —)10.0‒14.0(— 24.0) mm ( Figs. 3E and 12 ). Proximal primary in fl orescence bracts firm, apex acuminate, channelled or not, narrow at base, (25 —)39‒62(— 150) mm long, exceeding length of inflorescence up to several times. Spikes aggregated into clusters scattered along rachis, 2‒24, (4 —)7‒10(— 17) mm long, slightly overlapping. Spikelets lanceolate, (3.2 —)4.2‒5.3(— 7.1) X (0.6 —)0.9‒1.3(— 1.9) mm, pedicellate, 1 — 10 (usually 2 — 6) spikelets per spike, dark reddish-brown in colour ( Fig. 5F ). Proximal spikelet prophyll 1 per spikelet, firm, well-developed, with notable, raised vein extending to mucro, (0.2‒) 0.4‒2.8 mm long, mucros variable, (0.1 —)1.4‒2.2(— 7.6) mm long. Rachilla length variable, (0.1‒)1.0‒1.9(— 5.0) mm long. Glumes 4‒8 per spikelet, first three glumes relatively short and cuspidate, proximal glume (0.4‒)1.0‒1.4(— 2.6) mm long, subproximal glume (0.8‒)1.2‒1.7(— 3.2) mm long, usually with hyaline margins, upper glumes longer than basal ones, apex acuminate to obtuse. Glume mucros varying in length, proximal mucro (0‒)0.3‒1.0(— 2.1) mm long, subproximal mucro (0‒)0.2‒0.6(— 1.4) mm long. Stamens 3‒4 per floret, anthers (1.3‒)2.2‒2.8(‒4.2) mm long. Stigmas 3- branched, vestigial stigmas of second bisexual floret sometimes present. Perianth bristles absent (i.e. yet to be observed). Nutlet usually broad elliptic, trigonous, yellowish in colour when young and becoming darker with maturity, (1.4‒)2.1‒2.5(— 3.0) X 0.6‒1.3 mm ( Fig. 6F ). Nutlet beak (0.3‒)0.6‒0.8(‒1.1) mm long, hispid.

Flowering: Flowering specimens have been collected every month of the year.

Distribution and ecology: Schoenus cuspidatus is a species with a relatively wide geographical distribution compared to other South African congeners. Its distribution spans most of the Western Cape Province, as well as western regions of the Eastern Cape Province ( Fig. 13 ). The elevational range of S. cuspidatus is also broad, with specimens collected from near sea level to over 1500 m. This species grows in a variety of wet and dry habitats on various parent materials (e.g. sandstone, granite), but it is mostly found on mountain slopes in the fynbos vegetation type.

Diagnosis: Schoenus cuspidatus displays a significant amount of vegetative and reproductive variability. The main characters that correspond to this species include cuspidate spikelets ( Fig. 5F ) and broad elliptic nutlets ( Fig. 6F ), which are traits shared by S. loreus ( Figs. 5D and 6D ). This latter species, however, has inflorescences that are wider and longer than those of S. cuspidatus . In addition, S. loreus usually has relatively wide, non-channelled flat leaves ( Fig. 3D ) compared to the channelled, distally terete leaves of S. cuspidatus ( Fig. 4A ). Another similar species is S. riparius , which is a larger, more robust plant with longer, more dense inflorescences compared to S. cuspidatus ( Fig. 4 ).

The circumscription of S. cuspidatus has been a source of confusion for several botanists, confounded by probable hybridization within the S. cuspidatus group ( Levyns, 1950, p. 124). In addition, several intermediates, including those between S. cuspidatus and S. auritus (Bolus 7178, Elliott, Muasya & Muthaphuli TE2016_197, Elliott, Muasya & Muthaphuli TE2016_198, Esterhuysen 17,299a, Levyns 7599, Michell s.n., Pillans 3879), S. cuspidatus and Schoenus bolusii (C.B.Clarke) T.L.Elliott & Muasya (Levyns 8817), as well as S. cuspidatus and S. exilis ( Bos 188, Stokoe 2164) were identified by the authors during their investigation of herbarium specimens and plants in the field. We postulate that the taxonomic difficulties in this group are a result of its recent speciation, as many of the variants that have been observed might be incipient species. Preliminary genome size, chromosome count and next generation RAD-seq data collected by the authors also suggest these patterns; however, detailed analyses are required before they are confirmed.

We have noted four main variants during our detailed examination of S. cuspidatus (see Fig. 12 ), which have some geographic affinities. However, the presence of intermediates within each of these variants has made it impossible to delineate new species.

(1) Cederberg variant — this variant sometimes has very short and thin (awl-shaped) leaves and spikelets that reach up to 7.5 mm in length, which is generally longer than other variants. The following specimens are examples of this variant: Esterhuysen 7580, Esterhuysen 12,097, Esterhuysen 15,289.

(2) Cape Peninsula variant — the spikelets of this variant are about 6 mm long but can be slightly wider than the other variants. The leaves are usually about half the culm height, although specimens with longer leaves almost to the height of the culm have been collected (e.g. Wolley-Dod 2134). It is unclear whether these longer-leaved plants represent intraspecific variation due to site characters ( Levyns, 1947). Examples of this variant include Esterhuysen 15,747, Levyns 8710 and Elliott TE2016_285.

(3) Western Cape mountain variant — these plants are the most variable, but there is a tendency to have narrower leaves and spikelets, with less aggregation of spikes into clusters compared to the other variants (e.g. Taylor 5720, Levyns 8901). Specimens resembling S. exilis have been collected from several areas, including the Paarl Mountains (Kruger M187) and Somerset West (Parker 3468). The broad elliptic nutlets of these specimens place them within our circumscription of S. cuspidatus ; and

(4) Eastern variant — this variant is most like the Cape Peninsula variant, but it is less robust with more scarious lower glumes (e.g. Fourcade 1419, Esterhuysen 6565, Elliott & Betz TE2016_275).

Additional collections examined

South Africa. EASTERN CAPE: 3224 (Graaff-Reinet): ‘In monte graminoso "Koudveld" ( Sneeuwbergen ), (— AA), Dec 1872, Bolus 2585 ( BOL, K). 3225 (Somerset East): Boschberg, (— DA), Feb without year, MacOwan 1345 ( BOL) . 3323 (Willowmore): Kromrivierhoogte , (— DD), 15 Nov 1977, Bond 1205 ( PRE) ; Stormsriver Catchment , (— DD), 17 Nov 1977, Bond 1211 ( PRE) . 3325 (Port Elizabeth): Groendal Wilderness Res., Krompoortrivier catchm. basin., Strydomsberg Farm , (— CA), 1 Sep 1976, Scharf 1886 ( NBG, PRE) . 3326 (Grahamstown): near Riebeek, (— AA), Jul 1813, Burchell 3501 ( K); Goodwin's Kloof , (— BC), Oct 1892, Bennie 335 ( GRA) ; Howiesons Poort , at end of ridge between Featherstone and Osmunda Kloof, (— BC), 25 Sep 1949, Martin 3107 ( GRA) ; end of ridge between Featherstone Kloof and small kloof, Grahamstown , (— BC), 25 Sep 1949, Rhodes University 3107 ( BOL) ; Featherstone Kloof, Grahamstown , (— BC), 13 Nov 1949, Martin 3098 ( GRA) , Rhodes University 3098 ( BOL) ; Grahamstown, ‘ Faraway’ of Jacot Guillamod , (— BC), 14 May 1982, Getliffe 1198 ( NU) ; Grahamstown area, Osmunda Kloof , W. side, (— BC), 8 Nov 2017, Elliott & Betz TE2016_256 ( BOL) ; Grahamstown area, Featherstone Kloof, ca. 10 m to E. of trail going down to Featherstone Kloof , 50 m before small sign to N2, (— BC), 8 Nov 2017, Elliott & Betz TE2016_260 ( BOL) . 3424 (Humansdorp): Witte Els Bosch , (— AA), Aug 1921, Fourcade s.n. ( NBG) , Fourcade 1419 ( BOL [2 sheets]); P.E., Storms River past Humansdorp , turn off, (— BA), 12 Sep 1982, Getliffe 1144 ( NU) ; Humansdorp , (— BB), Aug 1921, Fourcade 1419 ( BOL) . WESTERN CAPE: 3118 (Vanrhynsdorp): Heerenloggementsberg, ca. 14 miles N. of Graafwater , (— DC), 13 Sep 1962, Taylor 3933 ( NBG) . 3218 (Clanwilliam): Piketberg, Platberg slope above Moutonsvlei , (— BB), 3 May 1975, Linder 541 ( BOL) ; W. slopes of Olifants River Dome, (— DB), 16 Apr 1949, Esterhuysen 15,289 ( BOL, NBG) ; Piketberg Mountain, slope between Tierhoek and E. escarpment, (— DB), 13 Jun 1976, Linder 621 ( BOL) . 3219 (Wuppertal): Krakadouw Peak, N. Cedarberg Mts. , (— AA), 20 Oct 1945, Esterhuysen 12,097 ( NBG) ; Cederberg Wilderness Area, along R364, E. of Pakhuis Pass , (— AA), 14 Jul 2016, Elliott & Muasya TE2016_085 ( BOL) ; Cederberg Mountains, Pk., S. of Sneeuwkop , (— AC), 2 Jan 1942, Esterhuysen 7580 ( BOL) ; Cederberg Mts., near Crystal Pool , (— AC), 26 Jun 1942, Esterhuysen 7902 ( BOL) ; N. Cederbergen, rocky slopes of peak at Konpoort , (— AC), 23 Oct 1945, Esterhuysen 12,180 ( BOL) ; Algeria Nature Reserve , Cedarberg: Plot 1, (— AC), 12 Nov 1969, Taylor 7493 ( NBG) ; Welbedacht , 2 km along jeeptrack N. W. from homestead, (— AC), 10 Apr 1986, Taylor 11,505 ( NBG) ; E. of Welbedacht homestead ca. 1 km, summit of koppie, (— AC), 22 May 1986, Taylor 11,525 ( NBG) ; Cedarberg, slopes below Tafelberg , (— AC), 19 Sep 2004, Forest et al. 596 ( NBG) ; Koude Berg , (— AC), 29 Aug 1896, Schlechter 8756 ( BM, BR, K, WAG); between Elandskloof and Citrusdal, (— CA), 4 Oct 1948, Levyns 9258 ( BOL) ; Krom River Kloof, S. Cederberge, (— CB), 11 Dec 1950, Esterhuysen 18,005 ( BOL) . 3318 (Cape Town): Wynberg Hill , (— AB), 15 Sep 1949, Esterhuysen 15,747 ( K, NBG) ; Constantia , (— AD), Oct 1908, Dummer 1983 ( NBG) ; Steenbras area, (— BB), 1 May 1948, Levyns 8872 ( BOL [2 sheets]); Malmesbury, Riverlands , 18 Nov 1997, (— BC), Muasya & Linder 1202 ( K), Muasya & Linder 1203 ( K, NBG) ; Kirstenbosch , (— CD), Apr 1917, Page s.n. ( BOL) ; Dec 1917, Page s.n. ( BOL) ; Jan 1918, Michell s.n. ( BOL) ; N. slope of Table Mt., (— CD), May 1918, Pillans 3879 ( K); Camps Bay , (— CD), 19 Oct 1942, Levyns 7582 ( BOL) ; 2 Sep 1945, Levyns 7909 ( BOL) ; Devils Peak, E. slope, (— CD), 23 Feb 1944, Esterhuysen 10,042 ( BOL) ; Rondebosch, University grounds, (— CD), 3 Sep 1945, Levyns 7912 ( BOL) ; Slangolie Ravine, Table Mt. , (— CD), 25 Nov 1945, Esterhuysen 12,293 ( BOL) ; Kirstenbosch, clearing N. of Window Stream, (— CD), Dec 1945, Esterhuysen 12,529 ( BOL) ; Kirstenbosch, amongst Silver Trees on lower slopes near Director's House and Herbarium , (— CD), 30 Aug 1949, Esterhuysen 12,688 ( BOL, K); clayish hillside above Rhodes Memorial , (— CD), 14 Sep 1949, Esterhuysen 15,753 ( BOL) ; Devils Pk., N.W. slopes below the Saddle , (— CD), 24 Aug 1952, Esterhuysen 20,318 ( BOL) ; U.C.T. grounds, slopes above the buildings, (— CD), 15 Nov 1956, Esterhuysen 26,561 ( BOL) ; University grounds, Rondebosch , (— CD), 15 Jun 1972, Esterhuysen 32,881 ( BOL) ; Camps Bay, west-facing slopes, near small foot path, ca. 80 — 90 m from Hely Hutchinson Ave. , (— CD), 15 Nov 2017, Elliott & Laidler TE2016_284 ( BOL) , Elliott TE2016_285 ( BOL) ; ‘ Contour Path’, ca. 200‒250 m to S. of Skeleton Gorge intersection, (— CD), 9 May 2018, Elliott & van Mazijk TE2016_363 ( BOL) ; ‘ In ericetis. In clivis orientalibus montis Diaboli pr. Cape Town’ , (— CD), Aug 1880, Bolus 4861 ( BOL) ;‘ In ericetis. In clivis orientalibus montis Tabularis’ , (— CD), Aug 1880, Bolus 4861 ( BOL) ; Kasteel's Poort , near tunnel, (— CD), 8 Aug 1896, Wolley-Dod 1366 ( BM, BOL) ; by reservoir, Newlands Woods , (— CD), 4 Dec 1896, Wolley-Dod 2134 ( BM, BOL) ; ‘ In Monte Diaboli’ , (— CD), without date, Rehmann 919 ( BR) , Rehmann 921 ( BM) ; Paarl Mountain, at point 12, (— DB), 25 Apr 1963, Kruger M187 ( NBG [3 sheets], PRE); Paarl, Paarlberg kop, (— DB), 9 Jul 1963, Bos 188 ( NBG) ; Paardeberg, between Wellington and Malmesbury , Waterkloof , top of kloof, (— DB), 2 Oct 2012, Nicolson 955 ( NBG) ; Monte Vista, (— DC), 20 Oct 1965, Esterhuysen 31,314 ( BOL, K); Klapmutz , (— DD), 9 Nov 1936, Lynes K1008 ( BM) ; Biesjevlei, Jonkershoek , (— DD), 2 Feb 1941, Levyns 7531 ( BOL) ; lower S. slopes of Guardian Peak, above Lourensford , 7 Oct 1945, Esterhuysen 12,008 ( BOL) ; Jonkershoek , (— DD), Sep 1948, Levyns 9712 ( BOL) ; Jonkershoek, Swartboskloof , (— DD), 20 Jan 1960, van der Merwe 2076 ( NBG) ; Bothmanskop , (— DD), 22 Sep 1960, van Rensburg 189 ( NBG) ; firebelt 13e, Jakkalsvlei, Jonkershoek , (— DD), 29 Mar 1962, Taylor 3240 ( NBG) ; Jakkalsvlei, Jonkershoek, 16 Mar 1964, Taylor 5720 ( BR, K, NBG); Jonkershoek , Langrivier , (— DD), Feb 1967, Kerfoot K 5759 ( NBG) ; Jonkershoek, Dwarsberg , (— DD), 13 Mar 1967, KR 401 ( NBG, PRE) ; Jonkershoek State Forest , the valley, (— DD), 11 Dec 1975, Kruger 133 ( NBG) ; 15 Apr 1976, Kruger 174 ( NBG) ; Stellenbosch, Jonkershoek , (— DD), 23 Nov 1997, Muasya & Rebelo 1227 ( K, NBG) ; Jonkershoek Nature Reserve , ca. 100 — 200 m S. of circular drive heading into reserve, (— DD), 22 Sep 2016, Elliott & Muasya TE2016_098 ( BOL) ; mountains round Stellenbosch , (— DD), Aug 1918, D ummer € s.n. ( SAM) . 3319 (Worcester): Groot Winterhoek , De Tronk, (— AA), 29 Dec 1979, Low 886 ( NBG) ; 1 Oct 1981, Low 1223 ( NBG) ; Gydoberg , (— AB), 10 Nov 1946, Walgate 8728 ( BOL) ; Cedarberg Mts., Middleberg , (— AC), 14 Dec 1941, Esterhuysen 7245 ( BOL) ; Elands Kloof , (— AC), Oct 1948, Levyns 9239 ( BOL) ; 17 Dec 1948, Levyns 9325 ( BOL) ; S.E. slopes at head of Tulbagh Valley , (— AC), 2 Oct 1950, Esterhuysen 17,543 ( BOL) ; Ceres , (— AD), 2 Oct 1933, Levyns 4683 ( BOL) ; Onderplaas, N. of N1, on recently formed ' Turtle' Conservation Property, just S. of Blaauberg Farm , (— BB), 9 Sep 2016, Euston-Brown 4719 ( BOL) ; Matroosberg, near Laaken Vlei , (— BC), Jan 1916, Phillips 17 ( SAM) ; mountains above Du Toits Kloof, (— CA), Apr 1932, Stokoe s.n. ( BOL) ; Wellington, slopes of Molenaarsberg , (— CA), 12 Feb 1944, Adamson 7672 ( BOL) ; Bains Kloof , (— CA), Feb 1944, Adamson 7674 ( BOL) ; swamp on Suurvlakte (behind Seven Sisters Mts. ), (— CA), 9 Feb 1947, Esterhuysen 13,732 ( BOL) ; easterly and northerly slope on farms Doolhof and Foxenburg , (— CA), 20 May 1963, Taylor 4793 ( NBG) ; Limietberg Nature Reserve , growing on S.E.-facing slope of R301, ca. 2 km to N.E. of Bainskloof community, (— CA), 25 Jun 2017, Elliott & Muasya TE2016_205 ( BOL) ; Hex River Mountains, path from Brandwacht farm to Thomas Hut , plots 92 — 95, (— CB), 23 Jun 1979, Campbell 10,933 ( NBG) ; Haalhoek Spitzkop , (— CC), 2 Jan 1947, Esterhuysen 13,532 ( PRE) ; Fransch Hoek , (— CC), 1 May 1948, Levyns 8901 ( BOL) ; French Hoek Forest Reserve , (— CC), 23 May 1948, Levyns 8901 ( BOL) ; Du Toits Kloof, clayish slopes at Mt. Club Hut , (— CC), 1 Aug 1949, Esterhuysen 15,628 ( BOL, NBG) ; Zachariashoek Experimental Catchment, Kasteelkloof Catchment , (— CC), 26 Apr 1979, Lamb 154 ( NBG) ; Zachariashoek, Kasteelkloof , (— CC), 10 Feb 1983, Viviers 983 ( NBG) ; Stettynsberg , stony E. slopes, (— CD), 31 Jul 1949, Esterhuysen 15,605 ( BOL, NBG) ; Onklaarberg, 20 miles S. of Worcester , (— DC), Apr 1925, Marloth 6644 ( NBG) ; Bosjesveld mountains, Villiersdorp side, (— DC), 1 Feb 1940, Stokoe s.n. ( SAM) . 3320 (Montagu): Swellendam Forest, road to Reservoir to the Plaas , (— CD), 11 May 1982, Getliffe 1122 ( NU) ; ' In montibus supra Voormansbosch’ , (— DC), Sep without year, Zeyher 4432 ( SAM) ; Boesmansbos area, Langeberg mountains above Heidelberg , (— DD), 27 Aug 1979, Campbell 14,632 ( NBG) ; 29 Aug 1979, Campbell 14,756 ( NBG) , Campbell 14,824 ( NBG) ; Grootvadersbosch State Forest, near Barend Koen Road, Boosmansbos Transect Releve 3, (— DD), 20 Sep 1985 , McDonald & Morley 1030 ( NBG) . 3321 (Ladismith): ' In declivibus saxosis - Garcia's Pass in dit. Riversdale’ , (— CC), Oct 1904, Bolus 24,343 ( BOL) . 3322 (Oudtshoorn): 12 km S. of Herold on road to George via Montagu Pass in Outeniekwaberge , (— CD), 8 Oct 1987, Davidse 33,713 ( WAG) ; Plot 227, Wilderness Heights, portion of farm Wildernishoogte 157, (— DC), 28 Jul 2002, Jacobsen 6061 ( K). 3323 (Willowmore): Hoopsberg, (— CB), 6 Nov 1941, Esterhuysen 6565 ( BOL) ; on W. side of R339 between Uniondale and Knysna , (— CC), 10 Nov 2017, Elliott & Betz TE2016_275 ( BOL) , Elliott & Betz TE2016_276 ( BOL) ; on S. side of R339 between Uniondale and Knysna , ca. 1.5 km to S.E. of turn-off with R340, (— CC), 10 Nov 2017, Elliott & Betz TE2016_277 ( BOL) , Elliott & Betz TE2016_278 ( BOL) . 3418 (Simonstown): Table Mountain, Echo Valley , (— AB), 18 Dec 1939, Levyns 7124 ( BOL) ; Kapiteins Peak, Hout Bay , (— AB), 20 Oct 1946, Levyns 8718 ( BOL) , Levyns 8719 ( BOL) , Levyns 8720 ( BOL) ; Constantiaberg, (— AB), May 1947, Levyns 8710 ( BOL [2 sheets]); ledges on S.E. side of Nursery Buttress , (— AB), 14 May 1949, Esterhuysen 15,382 ( BOL, NBG) ; Wynberg Hill , (— AB), 15 Sep 1949, Esterhuysen 15,746 ( BOL, NBG) , Esterhuysen 15,747 ( BOL) ; 1 Sep 1950, Pillans 10,183 ( BR) ; Orange Kloof , (— AB), 13 Jul 1955, Salter 9601 ( BM) , Salter s.n. ( BOL) ; S.W. of Noordhoek Peak, Plot 96, (— AB), 21 Jul 1965, Taylor 6400 ( NBG) ; Plot 100, L. Chapmans Pk., (— AB), 27 Jul 1965, Taylor 6412 ( NBG) ; Silvermine Reserve, (— AB), 14 Nov 1997, Muasya & Munro 1178 ( K); Silvermine Nature Reserve , along track, ca. 800 m S. of carpark on S. side of road, (— AB), 9 Jun 2002, Verboom 546 ( BOL) ; Table Mountain National Park, upward facing slope along Constantia Nek Jeep Track , (— AB), 24 Nov 2016, Elliott TE2016_160 ( BOL) ; Table Mountain National Park, Silvermine Nature Reserve, Silvermine West , ca. 600 m to S.W. of reservoir on Noordhoek Peak trail, (— AB), 16 Jun 2017, Elliott, Muasya & Muthaphuli TE2016_191 ( BOL) ; ‘ In monte Muizenberg, Peninsulae Capensis’ , (— AB), Apr without year, Bolus 7178 ( K); Simon's berg near the Waterfall, (— AB), without date, Drege s.n. ( SAM) ; Grabouw State Forest , Hottentots Holland Trail , (— BB), 10 May 1982, Getliffe 1090 ( NU) , Getliffe-Norris 1090 ( PRE) ; ca. 1 — 2 km S.W. of Gordon's Bay , along R44, (— BB), 18 Nov 2016, Elliott & Muasya TE2016_132 ( BOL) ; Betty's Bay, Erf 5039 edge, (— BD), 20 Sep 2000, Forrester JAF 1435 ( PRE) ; ca. 1 km north of Rooi Els , along R44, (— BD), 29 Nov 2017, Elliott & Smith TE2016_302 ( BOL) , Elliott & Smith TE2016_303 ( BOL) . 3419 (Caledon): Nuweberg Forest Reserve, source of Palmiet River , (— AA), 21 May 1969, Stehle TS 131 ( NBG) ; Lebanon State Forest , Jakkalsrivier Catchment Ie , (— AA), 4 May 1973, Haynes 793 ( K [2 sheets], NBG) ; Hottentots Hollands Nature Reserve , ca. 300 — 350 m from Nuweberg parking lot along jeep track to Landdroskop Hut , (— AA), 18 Feb 2017, Elliott & Muasya TE2016_166 ( BOL) ; between Genadendal and Donker Hoek, (— AB), 7 Mar 1815, Burchell 7920 ( K), Burchell 7926 ( K); Fernkloof Nature Reserve, on S. side of Klipspringer (blue) trail, ca. 50 m before first stream (dry), (— AC), 10 Feb 2017, Elliott TE2016_161 ( BOL) ; Kleinmond , (— AC), Apr 1930, Stokoe 2164 ( NBG) ; Staatsbos Sonderend Boesmanskloof , (— BA), 11 May 1982, Getliffe 1117 ( NU) ; Rooi Els , (— BD), 2 Aug 1947, Parker 4221 ( BOL, K); 31 Oct 1948, Levyns 9288 ( BOL) ; Elim, (— DA), 20 Apr 1896, Schlechter 7656 ( BM, GRA, K, WAG). 3420 (Bredasdorp): National Bontebok Park , (— AB), Dec 1962, Liebenberg 6683 ( K [2 sheets]); Van der Stel's Kraal, (— AC), 22 Jun 1968, Acocks 23,956 ( K, PRE) ; Farm Luipaardskop 53, ca. 12 km N.W. of Wydgele€ e, slopes between Suikerkankop , Kraaiheuwel and Rooikop Trig Beacon 157, (— AD), 10 Sep 2000, Helme 1748 ( NBG) ; on E. side of road going to W. entrance to De Hoop Nature Reserve , ca. 2 — 2.5 km from turn-off from road between Bredasdorp and Malgas, (— AD), 16 Sep 2018, Elliott & van Mazijk TE2016_396 ( BOL) , Elliott & van Mazijk TE2016_397 ( BOL) . 3422 (Mossel Bay): ‘ Environs de George’ , (— AB), 12 Aug 1933, Humbert 9869 ( K) .

Figures

Fig. 3. Examples of plants from five species of the S. cuspidatus and allies group found in Southern Africa, showing the corresponding inflorescences beneath each plant: A and F—S. brunnescens; B and G—S.compactus; C and H—S.crassus; D and I—S.loreus; and E and J—S.galpinii. The black scale bar adjacent to the inflorescence in J represents 10 mm.

Fig. 4. Examples of plants from four species of the S.cuspidatus and allies group found in Southern Africa, showing the corresponding inflorescences beneath each plant: A and E—S. cuspidatus; B and F—S.graciliculmis; C and G—S.riparius; and D and H—S.australis. The black scale bar adjacent to the inflorescence in H represents 10 mm.

Fig. 12. Inflorescence variation in S.cuspidatus. The black scale bar represents 10 mm.

Fig. 5. Examples of spikelets from nine species of the S. cuspidatus and allies group found in Southern Africa:A—S. brunnescens; B—S. compactus; C—S. crassus; D—S.loreus; E—S. gal- pinii; F—S.cuspidatus; G—S.graciliculmis; H—S.riparius; and I—S.australis. The black scale bar represents 1 mm.

Fig. 6. Examples of nutlets from nine species of the S.cuspidatus and allies group found in Southern Africa:A—S. brunnescens; B—S. compactus; C—S. crassus; D—S. loreus; E—S. galpinii; F—S.cuspidatus; G—S.graciliculmis; H—S.riparius; and I—S.australis. The black scale bar represents 1 mm.

Fig. 13. Documented locations of S.cuspidatus.