Schoenus ligulatus (Boeckeler) Kuntze, Revis. Gen. Pl.

Elliott, T. L., Barrett, R. L. & Muasya, A. M., 2019, A taxonomic revision of Schoenus cuspidatus and allies (Cyperaceae, tribe Schoeneae) - Part 1 *, South African Journal of Botany 121, pp. 519-535 : 526

3. Schoenus ligulatus (Boeckeler) Kuntze, Revis. Gen. Pl. 2: 756 (1891).

Elynanthus ligulatus Boeckeler, Linnaea 38: 262 (1874);

Tetraria ligulata (Boeckeler) C.B.Clarke in T.A. Durand & H. Schinz., Consp. Fl. Afric. 5: 661 (1894).

Tetraria cuspidata (Rottb.) C.B.Clarke var. ligulata (Boeckeler) Kük., Repert. Spec. Nov. Regni. Veg. 48: 210 (1940). Original citation: “In Herb. Neesii sub: ‘ Elynanthus an comparis monstrositas?’”.

Type: South Africa, Western Cape Province, 3418 (Simonstown): Red Hill area, Kleinplaas Dam, c. 100 m to E. of road that travels along E. side of dam, near old trail (with broken-down steps) heading downwards from road along dam (– AB), 18 Apr 2017, Elliott TE2016_178 ( BOL!, neo., here designated, MT!, NBG!, isoneo.). [

Note: Clarke (1900) noted that the ‘whole material’ for T. ligulata was in the herbarium of the Botanic Garden and Botanical Museum Berlin-Dahlem (B) without precise locality or collector name. He also mentioned that this material was not in a good condition and could have been a young specimen of T. sylvatica . The original material would have been examined by Kükenthal (1940), so details there potentially identify the collection as “Kapdistr., Kap flats (Zeyher n. 121! in herb. Nees n. 2609)”. After searching for this specimen, we think that it was most likely lost during the bombing of B during World War II. For this reason, we designate a neotype to fix the application of the name. Detailed searches of the Swedish Museum of Natural History (S) are recommended as the most likely location of any duplicate material of Zeyher 121].

Tetraria cuspidata (Rottb.) C.B.Clarke var. angustata Kük., Repert. Spec. Nov. Regni. Veg. 29: 187 (1931). Type: South Africa, Western Cape Province, 3419 (Caledon): Howhoek, (–AA), 8 Feb 1896, Schlechter 7429 (K, lecto!, here designated; BM!, BOL! [ BOL139429 ], GRA!, NSW!, PRE! [ PRE0488998 ], SAM!, isolecto.). [Note: Kükenthal (1931) cited three different collections: Ecklon and Zeyher 97B, Pillans 3903 and Schlechter 7429, of which only the latter two were examined by T.L. Elliott.].

Tetraria autumnalis Levyns, J. S. African Bot. 13: 88 (1947), syn. nov. Schoenus autumnalis (Levyns) T.L.Elliott & Muasya, S. African J. Bot. 112: 359 (2017) . Type: South Africa, Western Cape Province, 3418 (Simonstown): near Sirkel's Vlei, in sandy hollows which are damp during the winter months, (–DB), 23 May 1937, Levyns 6232 (BOL, holo! [ BOL139412 ]; BOL! [ BOL139424 , BOL150153 ], K! [K000244944], iso.).

Tetraria variabilis Levyns, J. S. African Bot. 13: 87 (1947), syn. nov. Schoenus variabilis (Levyns) T.L.Elliott & Muasya, S. African J. Bot. 112: 359 (2017) . Type: South Africa, Western Cape Province, 3418 (Simonstown); near Smitswinkel Bay, (–AD), 12 Apr 1937, Levyns 6054 (BOL, holo! [ BOL139438 ] BOL! [ BOL139439 ], iso.).

Caespitose, phyllopodic perennial graminoid, spikelets sometimes unisexual. Culms terete, (220–)343–533 × 0.2–0.7 mm. Leaves basal, 1–3(—6), (19–)99–230(—380) × 0.2–1.0 mm, usually straight, proximally channelled, margin usually serrate above sheath. Sheaths firm, reddish-brown, longitudinally striate. Ligule usually firm, 0.1–1.4 mm long. In fl orescence a pseudolateral panicle, (14–)30–58 × 3–13 mm, proximal rachis length varying in length, (2–) 11–24 mm long. Proximal primary in fl orescence bracts rarely channelled proximally, apex acute to acuminate, not widened at base, (21–)40–65(—76) mm long, exceeding length of inflorescence, longitudinal veins not notable. Proximal and subproximal primary inflorescence bracts without membranaceous extensions at base. Spikes 3–9(—25), (4.0–)8.0–19.0 mm long, usually dispersed along rachis. Spikelets linear-lanceolate, relatively long and thin, 4.6–5.7 (—7.4) × 0.6–1.3 mm, pedicellate, 1–4(—15) spikelets per spike, ranging from firm to thin in texture. Proximal spikelet prophyll 1 per spikelet, with vein extending to mucro, 0.3–1.5 mm long, prophyll mucro 1.3–2.8(—4.4) mm long. Rachilla 0.3–2.1(—7.5) mm long. Glumes 5–7 per spikelet, proximal glume 0.5–2.3 mm long, subproximal glume 1.0– 2.9 mm long, upper glumes longer than basal ones, lower glumes sometimes over half the length of spikelet, apex obtuse, narrow hyaline margins often present. Glume mucros, proximal mucro 0.5–1.8 mm long, subproximal mucro 0.3–1.1(— 2.9) mm long. Stamens (0–)1–3 per floret, (0–)1–3 per spikelet, anthers (1.4–) 2.4–3.7 mm long, spikelets sometimes unisexual. Stigmas 3-branched when present, vestigial stigmas of second bisexual floret not observed. Perianth bristles not observed. Nutlet narrow elliptic, trigonous, yellowish in colour, 2.1–3.7 × 0.5–0.9 mm. Nutlet beak 0.1–0.4(1.0) mm long, hispid ( Figs. 2 and 3 ).

Flowering: February–May

Distribution and ecology: Schoenus ligulatus is a species of the western part of the Western Cape Province of South Africa ( Fig. 9 ). Although S. ligulatus mostly occurs in wet habitats, it can also occur on drier, coarser soils. This species has been recorded from elevations from 100 to 750 m.

Diagnosis: The most notable characters of S. ligulatus are its narrow lanceolate spikelets and narrow elliptic nutlets. Schoenus ligulatus is similar to S. exilis as both species have narrow elliptic nutlets; however, S. ligulatus is a taller species (> 350 mm) compared to S. exilis (<350 mm). In addition, the spikelets of these two species are similar in shape, but those of S. exilis are shorter (mostly <4.5 mm) than those of S. ligulatus (> 4.5 mm). Furthermore, S. ligulatus usually has firm ligules, whereas those of S. exilis are often membranaceous. A second similar species is S. quartziticus which also has lanceolate spikelets, but that species has broad elliptic nutlets and shorter spikelets (<4.0 mm) compared to S. ligulatus .

Levyns (1947) did not mention Tetraria ligulata (Boeckeler) C.B. Clarke in her description of Tetraria autumnalis Levyns or Tetraria variabilis Levyns , so it is not clear whether she had examined specimens of the former species before describing the new species. After reviewing specimens from across the ranges of Schoenus autumnalis (Levyns) T.L. Elliott & Muasya , S. ligulatus and Schoenus variabilis (Levyns) T.L.Elliott & Muasya , we have decided to combine the three species into one. It is our view that the type specimens of T. autumnalis (Levyns 6232) and T. variabilis (Levyns 6054) display extreme examples of characters that fall within the limits of the more broadly circumscribed S. ligulatus ( Fig. 10 ).

Additional collections examined

South Africa. WESTERN CAPE: 3318 (Cape Town): Lions Head , (– CD), Apr without year, Zeyher s.n. ( SAM) ; Kenilworth , bushy places on racecourse, (–CD), 28 May 1937, Levyns 6234 ( BOL) ; Kenilworth Race Course , (–CD), 30 Oct 1938, Levyns 6835 ( BOL) ; 19 Feb 1970, Esterhuysen 32,398 ( BOL) ; Table Mountain near reservoir, (–CD), 20 Jan 1943, Compton 15,484 ( NBG) ; Kenilworth Race Course , in marshy area towards N. boundary, (–CD), 18 Mar 1970, Esterhuysen 32,423 ( BOL [2 sheets], K [2 sheets]) ; Kenilworth Racecourse Conservation Area , in central part of conservation area, (–CD), 7 Oct 2016, Elliott TE2016_115 ( BOL) . 3319 (Worcester): Wit River, Bainskloof , below Witrivierkamp monument, W. side of river, (–CA), 13 Mar 1997, Boucher 6097 ( NBG) ; flats S. side of FranschHoek Mts , (–CC), 11 Feb 1945, Esterhuysen 11,435 ( BOL) . 3418 (Simonstown): common near the stream at source of the Ruined House Valley, W. of Simon's Town , (– DB), 13 Feb 1919, Pillans 3903 ( BOL, K, PRE) ; plateau S. of Constantiaberg , (–AB), 24 May 1947, Levyns 8709 ( BOL) ; Cape of Good Hope Nature Reserve , gravel track from Olifantsbos road to Sirkelsvlei , (–AB), 23 Oct 2005, Verboom 904 ( BOL) ; Silvermine, wetland on way up to Noordhoek peak, permanent monitoring site #36, (–AB), 20 Dec 2016, Euston-Brown 5159 ( BOL) ; Judas Peak , (–AB), 10 Mar 2017, Euston-Brown 5213 ( BOL) ; Red Hill area, Kleinplaas Dam , (–AB), 18 Apr 2017, Elliott TE2016_178 ( BOL) ; Silvermine Nature Reserve, Silvermine West, c. 1.5 km to S.W. of reservoir on Noordhoek Peak trail, (–AB), 16 Jun 2017, Elliott, Muasya & Muthaphuli TE2016_195 ( BOL) ; near Smitswinkel Bay , (–AD), 12 Apr 1937, Levyns 6053 ( BOL) , Levyns 6054 ( BOL [2 sheets]), Levyns 6055 ( BOL) ; near Sirkels Vlei , (– AD), 23 May 1937, Levyns 6232 ( BOL [2 sheets], K) ; between road and Anvil Rock, Cape of Good Hope Nature Reserve , (–AD), 27 Feb 1938, Levyns 6528 ( BOL) ; between Theefontein and Sirkel's Vlei , (–AD), 16 May 1947, Levyns 8707 ( BOL) ; Cape Flats , Faure , Vergenoegd Farm , Extension Portion 653–15, flats along W. side of N.W. farm entrance road, (– BA), without date, Boucher 7467 ( NBG) ; Steenbras Dam Forest area , (–BB), 7 May 1982, Getliffe 1178 ( NU) ; Palmiet River Mouth, (– BD), 12 Feb 1937, Levyns 6007 ( BOL) ; 30 Jan 1947, Levyns 8151 ( BOL) ; Feb 1943, Levyns 8714 ( BOL) ; 6 May 1948, Levyns 8892 ( BM, BOL) ; mountain slopes near Palmiet River mouth, (–BD), 21 Jan 1946, Esterhuysen 12,603 ( BOL) ; Kogel Berg , (–BD), 1 Mar 1947, Compton 20,464 ( NBG) ; Palmiet River Mouth, on treck to old farm, (–BD), 16 Feb 1948, Levyns 8803 ( BOL) ; Cape Hangklip , (–BD), 25 Feb 1955, Levyns 10,232 ( BOL) , Levyns 10,236 ( BOL) ; c. 1.5 km W.S.W. from Palmiet River mouth, (–BD), 15 Mar 1983, van Wyk 1108 ( K, NBG) ; southern Hottentots Holland Mountains , Kogelberg State Forest , ± 2.5 km from Oudebos at 2nd waterfall on edge of stream near bridge upstream, (–BD), 12 May 1992, Kruger 580 ( K, NBG) . 3419 (Caledon): Howhoek , (–AA), 8 Feb 1896, Schlechter 7429 ( BM, BOL, GRA, K, NSW, PRE, SAM, WAG) ; Lebonon , Catchment II c., (–AA), Feb 1968, Kruger 351 ( K, NBG, PRE) ; Lebanon State Forest , Jakkalsrivier Catchment Iia. , (–AA), 27 Feb 1969, Kruger 799 ( NBG [2 sheets]) ; Lebanon State Forest , Jakkalsrivier Catchment Ib. , (–AA), 3 Feb 1973, Haynes 771 ( NBG [2 sheets], PRE) ; Hottentots Hollands Nature Reserve , c. 1.25 km from Nuweberg parking lot, along jeep track to Landdroskop Hut , (–AA), 18 Feb 2017, Elliott & Muasya TE2016_174 ( BOL) ; Kleinmond , (–AC), Feb 1945, Levyns 7699 ( BOL) ; Grabouw, Highlands Forest Reserve , Perdeberg , (–AC), 3 Mar 2002, Verboom 508 ( BOL) ; Voelklip, Hermanus, between 5 th and 6 th Streets , (–AD), 12 Mar 1981, Williams 644 ( K, PRE, WAG) ; mountains near Rooi Els , (–BD), 2 May 1949, Levyns 9398 ( BOL) .

Figures

Fig.2. Examples of nutlets from 11 species of the Schoenus cuspidatus and allies group found in southern Africa:A—S.auritus; B—S.graminifolius; C—S.ligulatus; D—S.exilis; E—S.bolusii; F— S.submarginalis; G—S.crassiculmis; H—S.quartziticus; I—S.limosus; J—S.purpurascens; and K—S.calceolus (immature).The black scale bar represents 1 mm.The mature fruits of S.calceolus and S.prophyllus are unknown.

Fig.3. Examples of plants from six species of the Schoenus cuspidatus and allies group found in southern Africa,showing the corresponding inflorescences andspikelets beneath each plant: A, G and M—S.auritus; B, H and N—S.graminifolius; C, I and O—S. ligulatus; D,J, and P—S.exilis; E,K and Q—S.bolusii; and F, L and R—S.submarginalis. The black scale bar adjacent to the inflorescences in G—L represents 10 mm, while the scale bars adjacent to the spikelets in M—R represent 1 mm.

Fig. 9. Documented locations of Schoenus ligulatus.

Fig. 10. Variation in plant habit in Schoenus ligulatus, where A is an example of the more robust form of the species (Type specimen of S. autumnalis collected from the Cape Peninsula), and B is an example of the more common, less robust form collected from the Cape Peninsula.