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Lombard, Natasha, Le Roux, M. Marianne (2023): Thesium hispidifructum (Santalaceae), a New Hispidulous Species from Limpopo, South Africa and Notes on Enigmatic T. celatum. Taxonomy (Clifton, N.J.) 3 (1): 95-108, DOI: 10.3390/taxonomy3010009, URL: http://dx.doi.org/10.3390/taxonomy3010009

Thesium hispidifructum N.Lombard & M.M.le Roux, sp. nov. (Figure 5) Type: SOUTH AFRICA. Limpopo Province: Lebowa, Blouberg, Farm Beauly 260 LR, 12 February 1991, L. Smook 7447 (holotype PRE 0775984). Diagnosis: Thesium hispidifructum is morphologically similar to T. disparile in its hairy indumentum, ascending to spreading leaves (4–15 mm long in T. hispidifructum; 7–11 mm long in T. disparile), flowers placed in determinate racemose cymes and flower lobes (0.8–1.2 mm long in T. hispidifructum; ± 0.7–1.2 mm long in T. disparile) bearded at the inner apex. Thesium hispidifructum is easily distinguished from T. disparile by its tall robust stature (woody stems up to 1 m tall), intricate branching pattern and compact inflorescences with flowers solitary in the bract axils. Comparatively, T. disparile has a short slender stature (herbaceous stems up to 0.15(0.5?) m tall), virgate branching pattern and lax inflorescences with single flowers usually replaced by compound monochasial or dichasial cymes. Description: Shrub or subshrub, up to 1 m tall, rootstock not known; stems erect, up to 4 mm in diameter, intricately branched above, hispidulous to hispid. Leaves spreading to ascending, sparse below and densely arranged above, often imbricate, linear, 4–15 × 0.3–1.0 mm, hispidulous to hispid. Flowers solitary in bract axils, arranged in 5–10-flowered determinate raceme-like inflorescences; pedicels 0–1.3 mm long. Bracts and bracteoles usually three, leaf-like but smaller, 2–7 mm long, not forming an involucre, bracts fused to at least half of pedicel. Perianth ±campanulate, 2.6–2.7 mm long, elongate receptacle often present, “glands” often visible on outside, exterior hispidulous to hispid; hypanthium 0.3–0.6 mm long; corolla lobes triangular, 0.8–1.2 mm long, apex very slightly cucullate, with dense apical beard of long hairs. Stamens inserted at base of corolla lobes; anthers ± 0.3 mm long. Style 0.2–0.4 mm long, stigma below or reaching the bottom of anthers. Placental column straight; ovules 3. Fruit ± globose, 3.6–4.1 × 1.8–2.3 mm including stipe and persistent perianth, stipe ± 0.4 mm long, persistent perianth 0.9–1.3 mm long, longitudinally and reticulately veined when dry, hispid. Distribution and habitat: Thesium hispidifructum is endemic to the Limpopo Province in South Africa, where it is known from only four gatherings between Blouberg and Mount Letsjume in the westernmost reaches of the Soutpansberg (Figure 6). Little is known about its habitat. It has been collected among grass in rocky areas as well as in wet wooded areas, on both south facing and northeast facing slopes at elevations between 1675 and 1980 m a.s.l. It is likely restricted to the Soutpansberg Mountain Bushveld [21] and Soutpansberg Summit Sourveld [21], but field studies are needed to confirm this. Phenology: Thesium hispidifructum has been collected in flower and fruit in January, February, June, and July. Etymology: The specific epithet hispidifructum refers to the hispidulous fruit of the species. Conservation status: Thesium hispidifructum has an extent of occurrence (EOO) of 44 km 2 and an area of occurrence (AOO) of 12 km 2. It is known from only four gatherings at two to three locations in mountainous areas (Blouberg and Mount Letsjume), surrounded by much-transformed low-lying areas (Vivo and surrounds). As the most recent collection/observation of this species was in 1991, the state of these subpopulations is unknown, and needs to be verified during field studies. It is likely that the extensive human settlements and crop development in low-lying areas surroundings these mountains have already resulted in the loss of suitable habitat. Furthermore, expanding anthropogenic influences may easily push this very restricted species to extinction. It therefore qualifies for a threat status of Vulnerable (VU) under Criterion D2 [20]. Additional specimens examined: SOUTH AFRICA. Limpopo: Blauwberg, 10 May 1933, A. C. Leeman 110 (PRE 0374089); Blauwberg, near trig. beacon, 12 January 1955, L. E. Codd and R. A. Dyer 9063 (PRE 0374090); Zoutpansberg, Mt Letsjume, at foot, on S slope, 25 July 1981, F. Venter (PRE 0620517) ., Published as part of Lombard, Natasha & Le Roux, M. Marianne, 2023, Thesium hispidifructum (Santalaceae), a New Hispidulous Species from Limpopo, South Africa and Notes on Enigmatic T. celatum, pp. 95-108 in Taxonomy (Clifton, N. J.) (Clifton, N. J.) 3 (1) on pages 103-105, DOI: 10.3390/taxonomy3010009, http://zenodo.org/record/7981347, {"references":["21. Mucina, L.; Rutherford, M. The Vegetation of South Africa, Lesotho and Swaziland; Strelitzia 19; South African National Biodiversity Institute: Pretoria, South Africa, 2006","20. IUCN Standards and Petitions Subcommittee. Guidelines for Using the IUCN Red List Categories and Criteria. Version 13; IUCN: Gland, Switzerland, 2017."]}

4.3. L. gracilis L. gracilis Eckl. and Zeyh., Enum.: 192 (1836); Benth. in Hook., Lond. J. Bot. 3: 358 (1844). Type: South Africa, “Zwartkopsrivier” [3322 CD], Ecklon and Zeyher 1336 (SAM!, specimen in middle of the sheet, lectotype, designated here; P!, S!, isolectotypes). L. sepiaria auct. non (L.) Thunb.: Eckl. and Zeyh., Enum.: 191 (1836); Harv. in Harv. and Sond., Fl. Cap. 2: 86 (1862); Goldblatt and Manning, Cape Plants, Strelitzia 9: 493 (2000). [Harvey cited a Thunberg specimen (Thunberg s.n. sub Herb. Thunberg 16422) which is indeed this species. However, Thunberg based his description on a specimen from Cape Town “Leeuwenberg”, but he explicitly cited Spartium sepiarium of Linnaeus, q.v.] L. ambigua E. Mey., Com.: 34 (1836), pro parte minore. Erect, glaucous suffrutex, 0.15 − 0.8 m in height. Leaves simple, acicular, articulated near the middle; leaf density 3–6 per 10 mm length of stem. Inflorescences terminal, 44–172 mm long; rachis furrowed; flowers numerous, relatively dense (5–7 per 10 mm length of rachis), relatively small in size (6–8 mm long); pedicel 1.5–3.0 mm long; bract lanceolate, acute, ± 2 mm long; bracteoles triangular, acuminate, ± 1 mm long. Calyx 2.5–4.5 mm long, tube 1.5–3.5 mm long, lobes 0.7–1.5 mm long; ± equally lobed; lobes deltoid, tips minutely pubescent inside. Standard widely oblong to suborbicular or obovate, 4.0–6.5 × 2.0– 3.5 mm; apex obtuse; claw 1.5–2.5 mm long. Wings oblong, longer than the keel, 4.0–6.5 × 1.0– 2.5 mm, with 11–14 rows of sculpturing; apex obtuse; claws 1.5–2.0 mm long. Keel rostrate, 5–6 × 1.5–2.5 mm, sometimes with pockets; claws 1.5–2.5 mm long. Pistil subsessile; ovary 5.5–7.5 × 0.6–0.8 mm long, linear; ovules many (9–11); style 1.5–3.5 mm long, curved upwards. Pods linear, straight or very slightly falcate, up to 37 × 2–3 mm; dehiscent; semi-terete; fruit wall thin, membranous. Seeds reniform-orbicular, ± 1.8 × 1.5 mm, rugose, pale brown, mottled dark brown to black; hilum dark brown (Fig. 9). 4.3.1. Diagnostic characters L. gracilis is similar to L. ambigua but differs in its calyx that is ± equally lobed and with the lobes shorter than the tube (calyx subequally lobed and the lobes as long as the tube in L. ambigua) (Fig. 9). L. gracilis also differs from L. ambigua in its shorter and more congested racemes. The two species are allopatric. 4.3.2. Distribution and habitat L. gracilis has a distinctively coastal distribution, from Bredasdorp in the Western Cape Province eastwards to Port Elizabeth in the Eastern Cape Province (Fig. 10). All known localities are below 300 m altitude. Those in the west experience winter rainfall, while those in the east (around Port Elizabeth) have a higher proportion of summer rain. Plants grow in deep sandy soil. 4.3.3. Specimens examined. – 3322 (Oudtshoorn): Fairy Knowe (– DC), Guillarmond 8257 (PRE); Rondevlei (– DC), Sachse 670 (PRE); Wilderness (– DC), Van Niekerk 223 (BOL); Ruigtevlei at Zwart River (– DD), Fourcade 1534 (BOL, NBG, PRE). – 3324 (Steytlerville): Zwartkopsrivier (– DC), Ecklon and Zeyher 1336 (P, S, SAM), Zeyher 2293 (P, S); Gamtoos River (– DD), Gillett 2361 (NBG). – 3325 (Port Elizabeth): Humewood (– DC), Paterson 773 (BOL). – 3420 (Bredasdorp): De Hoop (– AD), Morley 139 (PRE); Melkbosheuwel (– BC), Burgers 2461 (NBG); Potberg area (– BC), Derdejaarsekskursie 84/C10 (NBG); Potberg (– BC), Esterhuysen 23259 (BOL); De Hoop (– BC), Fellingham 701 (PRE); The Poort (– CA), Henderson 1829 (NBG), Morris 268 (NBG); Bredasdorp Poort (– CA), Heginbotham 163 (NBG). – 3421 (Riversdale): Oude Tuin near Albertinia (– AB), Muir 1852 (BOL, PRE); Stilbaai (– AD), Jordaan 18607 (NBG); Stilbaaihoogte (– AD), Bohnen 7617 (NBG), Le Roux, Boatwright, Magee and Van Wyk 17 (JRAU). – 3422 (Mossel Bay): Brak River (– AA), Young 5514 (BOL); Belvedere (– BB), Duthie 510 (NBG); Goukamma (– BB), Wurts 2234 (NBG). – 3423 (Knysna): Between Knysna and Plettenberg Bay (– AA), Pappe s.n. (S, SAM); Plettenbergbaai (– AB), Anon s.n. sub STEU 13540 (NBG). – 3425 (Port Elizabeth): Schoenmakers Kop (– AB), Story 3654 (PRE)., Published as part of le Roux, M. M. & Van Wyk, B. - E., 2007, A revision of Lebeckia sect. Lebeckia: The L. sepiaria group, pp. 118-130 in South African Journal of Botany 73 (1) on pages 126-127, DOI: 10.1016/j.sajb.2006.09.005, http://zenodo.org/record/7958313

4.1. L. ambigua L. ambigua E. Mey., Comm. Pl. Afr. Austr. 1: 34 (1836), pro parte majore; Benth. in Hook., Lond. J. Bot. 3: 357 (1844); Harv. in Harv and Sond., Fl. Cap. 2: 86 (1862); Goldblatt and Manning, Cape Plants, Strelitzia 9: 493 (2000). Type: South Africa, “Uienvalley” [near Clanwilliam, 3218 BB], Drège s.n. (“ III, A, c”) (P!, the specimen on the right hand side of the sheet, lectotype designated here; S!, isosyntype) and Saldanha Bay [3318 AA], Drège s.n. (“ III, E, b”), (P! isosyntype). Erect, glaucous suffrutex, up to 0.8 m in height. Leaves simple, acicular, articulated near the middle; leaf density 2–5 per 10 mm length of stem. Inflorescences terminal, 90–350 mm long; rachis smooth; flowers numerous, relatively sparse (3–5 per 10 mm length of rachis), intermediate in size (7–10 mm long); pedicel 2– 3 mm long; bract narrowly lanceolate, acuminate, 2.0– 3.5 mm long; bracteoles narrowly triangular, acuminate, ± 1 mm long. Calyx 3.5–5 mm long, tube 1.5–2.5 mm long, lobes 1.5–2.5 mm long; subequally lobed but the upper lateral sinuses much wider than the medial sinus and the lower lateral sinuses; lobes deltoid, tips minutely pubescent inside. Standard widely ovate to orbicular, 8–10 × 4.5–8.0 mm; apex obtuse; claw 1–2 mm long. Wings oblong, as long as or usually longer than the keel, 7–9 × 2.0– 3.5 mm, with 9–12 rows of sculpturing; apex obtuse; claws 1.5–2.5 mm long. Keel rostrate, 6.0–8.5× 2.5– 3.5 mm, usually with pockets; claws 1.5–3.0 mm long. Pistil subsessile; ovary 5.5–11.0 × ± 0.5 mm long, linear; ovules many (14–18); style 2.5–3.5 mm long, curved upwards. Pods narrowly linear, straight or falcate, up to 75 × 2 mm; dehiscent; terete; fruit wall thin, membranous. Seeds oblong-reniform, 2.0 × 1.2 mm, rugose, light pink; hilum black (Fig. 5). 4.1.1. Diagnostic characters L. ambigua is closely related to L. sepiaria but differs in the membranous, narrowly linear, straight or falcate, ± 2 mm wide, dehiscent fruit (L. sepiaria has spongy, linear, straight, ± 4 mm wide and indehiscent fruit) and smaller flowers (7–10 mm long) (12–18 mm long in L. sepiaria) (Fig. 5). 4.1.2. Distribution and habitat L. ambigua occurs in the low-lying coastal region of the Northern and Western Cape Provinces, from Namaqualand southwards to a little north of Cape Town (Fig. 6). The altitude is usually below 300 m but may be up to 900 m at some localities. Plants grow in deep sandy soil, usually in obviously disturbed areas such as riverbanks and roadsides. 4.1.3. Specimens examined – 2917 (Namaqualand): Namaqualand (– CD), Le Roux 374 (NBG). – 3017 (Kamieskroon): Kotzerus (– DD), Acocks 23398 (PRE). – 3117 (Vredendal): Lepelfontein (– BB), Helme 2930 (NBG). – 3118 (Van Rhynsdorp): 23 miles from Clanwilliam (– DA), Grobbelaar 1129 (PRE); Van Rhynsdorp (– DA), Marloth 2627 (PRE), Rossouw 18619 (NBG); near turn-off to Klawer on the N7 (– DA), Van Wyk 3120 (JRAU); foot of Gifberg (– DB), Zietsman 1180 (PRE); foot of Heerelogementsberg near Klawer (– DC), Van Wyk 3284 (JRAU); Klawer (– DC), Barker 3637 (NBG), Lewis 1309 (SAM); De Little Farm, Nardouwsberg (– DD), Stirton 9365 (PRE). – 3119 (Calvinia): Arendskraal Farm, near Nieuwoudtville (– AC), Barker 9781 (NBG); Lokenburg House (– CA), Acocks 18568 (PRE); top of Botterkloof (– CD), Barker 6515 (NBG). – 3218 (Clanwilliam): Wadrif, Soutpan, coastal side of the railway (– AB), O̕Ca llanghan, Van Wyk and Morley 90 (PRE); Lambert̕s Bay (– AB), Henrici 3297 (PRE); farm Wolfhuis (– AB), Stirton 9294, 9388 (PRE); Uitkomsberge, Bo-Voëlfontein Farm (– AB), Stirton 9351 (PRE); Nortier station (– AB), Van Breda 4460 (PRE); Verlore Vlei (– AD), Stirton 6100 (PRE); between Elandsvlei and Redelinghuys (– AD), Stirton 6117 (PRE); between Leipoldtville and Elands Bay (– AD), Zinn s.n. (SAM); between Lambert̕s Bay and Clanwilliam (– BA), Marsh 781 (NBG); Graaffwater (– BA), Van Breda 4363 (PRE); Clanwilliam Park (– BB), Acocks 15179 (PRE); Olifants River Bridge (– BB), De Winter 9564 (PRE); Clanwilliam (– BB), Leipoldt 349 (SAM), Repton 7176 (PRE), Van Wyk 2605 (PRE); between Clanwilliam and Springbok (– BB), Le Roux 2625 (BOL); north of Clanwilliam (– BB), Le Roux, Boatwright, Magee and Van Wyk 6 (JRAU); Redelinghuys, Piketberg (– BC), Acocks 19795 (PRE), Compton 15052 (NBG); Paleisheuwel (– BC), Stirton 9294 (PRE); Berg River (– CC), Ecklon and Zeyher 1334 (SAM); Berg River Station, Malmesbury (– CD), Compton 15103 (NBG); Paleisheuwel (– CD), Stirton 9294 (PRE); Berg River Valley (– DA), Acock 2930 (S); Paleisheuwel Station, north of Het Kruis (– DA), Wisura 3486 (NBG); 3 miles from Grey̕s Pass, Piketberg (– DB), Barker 6406 (NBG); between Piketberg and Grey̕s Pass (– DB), Lewis 3391 (SAM); between Piketberg and Citrusdal (– DB), Parker 3588 (NBG); Citrusdal (– DB), Niemand 11 (JRAU); between Pakhuis Pass and Piquetberg (– DB), Parker 61 (BOL); between Citrusdal and Clanwilliam (– DB), Van Wyk 2900 (JRAU); Piekenierskloof Pass (– DB), Van Wyk 3220 (JRAU); Olifantsrivier (– DC), Grobbelaar 1129 (PRE); Piketberg (– DC), Heyns 26146 (NBG); Sauer (– DC), Acocks 24459 (PRE); Piketberg (– DD), Van Breda 352 (PRE). – 3219 (Wuppertal): Cedarberg, Boesmanskloof, Wuppertal (– AA), Van Rooyen, Steyn and De Villiers 499 (NBG); Cedarberg, Wuppertal (– AC), Bolus 8972 (NBG); Warm Baths (– CA), Le Roux, Boatwright, Magee and Van Wyk 9 (JRAU). – 3318 (Cape Town): Langebaan, Schrywershoek (– AA), Boucher 2946e (NBG); Langebaan (– AA), Cupido 13 (NBG); Langebaan, behind the granite koppie (– AA), Goldblatt 2704 (PRE); Saldanha Bay, Langebaan (– AA), Taylor 3757 (NBG); Darling near Ysterfontein (– AA), Van Rensburg 157 (NBG, PRE); 125 km north of Cape Town on the R27 (– AA), Van Wyk 3290 (JRAU); Hopefield (– AB), Acocks 20669 (PRE), Bolus 12653 (PRE); Modderriver (– AD), Le Roux, Boatwright, Magee and Van Wyk 14 (JRAU); in fields near Hopefield (– DC), Letty 35 (PRE)., Published as part of le Roux, M. M. & Van Wyk, B. - E., 2007, A revision of Lebeckia sect. Lebeckia: The L. sepiaria group, pp. 118-130 in South African Journal of Botany 73 (1) on pages 122-124, DOI: 10.1016/j.sajb.2006.09.005, http://zenodo.org/record/7958313

4.4. L. sepiaria L. sepiaria (L.) Thunb., Prod. Pl. Cap.: 122 (1800), Fl. Cap.: 561 (1823); Benth. in Hook., Lond. J. Bot. 3: 192 (1844) non Harv. (1862), nec Goldblatt and Manning (2000). Type: South Africa, LINN 891.4 (LINN!, lectotype, designated here). S. sepiarium L., Sp. Pl.: 995 (1753) and Pl. Rar. Afr.: 91 (1760), Gmelin, Syst. Nat. 2 (2): 1088 (1792). Type as for L. sepiaria. Sarcophyllum carnosum Sims, Bot. Mag.: 51, t. 2502 (1824). Iconotype: Sims, Bot. Mag.: 51, t. 2502 (1824). [Note: The illustration and description leaves no doubt about the identity of the species illustrated. The calyx is not equally lobed, with the lateral sinus wider than the upper and lower sinuses and the wing petals are shorter than the keel petal. The proliferation of the apical bud of the inflorescence is very unusual. The Ecklon specimen on which Meyer based his description of L. sarcophylloides [Ecklon and Zeyher 1338 (SAM!)] also agrees with this species. L. sarcophylloides E. Mey. in Linnaea 7: 155 (1832), nom. nov. pro S. carnosum Sims non L. carnosa Thunb. Type as for S. carnosum. L. simsiana Eckl. and Zeyh., Enum. 2: 192 (1836); Harv. in Harv. and Sond., Fl. Cap. 2: 86 (1862); Goldblatt and Manning, Cape Plants, Strelitzia 9: 493 (2000), synon. nov. Type as for S. carnosum. L. contaminata (L.) E. Mey., Comm. Pl. Afr. Austr. 1: 34 (1836), non Thunb. Erect, glaucous suffrutex, up to 0.8 m in height. Leaves simple, acicular, articulated near the middle; leaf density 4–11 per 10 mm length of stem. Inflorescences terminal, 50– 300 mm long; rachis furrowed; flowers numerous, relatively sparse (2–5 per 10 mm length of rachis), relatively large in size (12–18 mm long); pedicel 4–7 mm long; bract lanceolate, acute, ± 3 mm long; bracteoles narrowly triangular, acuminate, 0.5–1.5 mm long. Calyx 4.5–6.5 mm long, tube 2.5–3.5 mm long, lobes 1.5–2.5 mm long; subequally lobed but the upper lateral sinuses much wider than the medial sinus and lower lateral sinuses; lobes deltoid, tips minutely pubescent inside. Standard widely ovate to orbicular, 9– 15× 7.5–11.5 mm; apex obtuse; claw 1.0– 3.5 mm long. Wings oblong, shorter than the keel, 7.5–12.5 × 3.5–6.0 mm, with 13– 16 rows of sculpturing; apex obtuse; claws 1.5–4.5 mm long. Keel rostrate, 7–14 × 3.0–5.0 mm, usually with pockets; claws 2–5 mm long. Pistil subsessile; ovary 13.5–15.5 × 0.6–0.9 mm long, linear; ovules many (15–19); style 4.0– 5.5 mm long, curved upwards. Pods linear, straight, up to 40 × 4 mm; indehiscent; terete; fruit wall thick, spongy. Seeds oblong-reniform, ± 2.5 × 1.2 mm, rugose, pale brown, mottled dark brown to black; hilum black (Fig. 11). 4.4.1. Diagnostic characters L. sepiaria is one of the most well-known species in sect. Lebeckia. It differs markedly from L. brevicarpa in the long, linear fruits (up to 40 mm long) and wings that are shorter than the keel. In L. brevicarpa, the fruits are short and ovate (up to 13 mm long) and the wings are longer than the keel (Fig. 11). 4.4.2. Distribution and habitat L. sepiaria occurs only in the Western Cape Province (Fig. 12). It differs markedly from L. ambigua and L. gracilis in occurring mainly at higher altitudes (300 − 1500 m, with only a few localities near Cape Town that are below 300 m). Plants grow in welldrained sandy soil. 4.4.3. Specimens examined – 3218 (Clanwilliam): Piketberg, Redelinghuys (– BC), Stephans 7013 (NBG); Paleisheuwel Station (– DA), Wisura 3486 (NBG); Olifants Rivier Valley (– DB), Barker 10336 (NBG); Warmbaths (– DB), Edwards 251 (PRE); Citrusdal, Jansekraal (– DB), Hanekom 1300 (NBG); Citrusdal (– DB), Hanekom 2851 (NBG, PRE), Parker 3588 (NBG); Citrusdal, Berg-en-Dal (– DB), Hanekom 3209 (PRE); farm at the top of Zebrakop, Piketberg (– DB), Taylor 5319 (PRE); Piketberg (– DC), Goldblatt 2743 (PRE), Van Breda 2005 (PRE); Piketberg (– DD), Bodkin 13537 (PRE), Bolus 13537 (PRE), Gulline 2585 (NBG), Marloth 11477 (PRE); on top of Piketberg mountain (– DD), Bodkin and Bolus 13537 (PRE). – 3219 (Wuppertal): Kleinplaas (– CA), Hanekom 2896 (NBG, PRE); Middelberg Pass, Elandskloof (– CA), Le Roux, Boatwright, Magee and Van Wyk 13 (JRAU); Olifants River Valley (– CA), Stephans 7013 (BOL); 3.6 km N. of Warm Baths (– CA), Le Roux, Boatwright, Magee and Van Wyk 10, 11 (JRAU); Cedarberg (– CB), Thode 2151 (PRE); Wuppertal, Olifants River (– CC), Thompson 1505 (NBG). – 3318 (Cape Town): Above Riebeek Kasteel (– BD), Drège s.n. (PRE, S); Riebeek Kasteel (– BD), Marsh 1053 (NBG); Langverwacht above Kuilsrivier (– DC), Olivier 4753 (NBG); Stellenbosch (– DD), Bolus 10722 (BOL), Prior s.n. (PRE); Jonkershoek (– DD), Borchardt 417 (PRE); Jonkershoek (– DD), Hubbard 285 (PRE); between Stellenbosch and Vlottenberg (– DD), Marloth 8755 (NBG); Stellenbosch, Banhoek (– DD), Martley 32 404 (BOL); Stellenbosch, Vredenburg (– DD), Salter 7800 (NBG); Stellenbosch, Swartboskloof (– DD), van Rensburg 2135 (NBG, PRE); Assegaaibosch (– DD), Van derMerwe 1224 (PRE). – 3319 (Worcester): Breederivier, Paspasvalley, Vierentwintigrivier (– AA), Ecklon and Zeyher 1338 (SAM); Tulbagh, Winterhoek (– AA), Pappe s.n. (SAM); Foothills of Witzenberg (– AC), Schonken 317 (NBG, PRE); Boesmanskloof (– CA), Taylor 6507 (PRE); Goudini road (– CB), Middlemost and Creasy 2143 (NBG); Bastiaanskloof (– CB), Taylor 6507 (PRE); Brandvlei Prison (– CD), Forrester 514 (NBG); Villiersdorp (– CD), Schlechter 140 (PRE); Louwshoek (– CD), Stokoe s.n. (SAM); Stettynskloof (– CD), Walters 227, 1065 (NBG); between Worcester and Villiersdorp (– CD), Walters 398 (NBG); Villiersdorp, Doornrivier (– CD), Walters 1380 (NBG); 20 km from McGregor on the way to Greyton (– DC), Grobbelaar 2209 (PRE); McGregor (– DC), Grobbelaar 2856 (PRE). – 3320 (Montagu): Foot of Tradouw Pass, Barrydale (– DC), Marsh 864 (NBG); Langeberg between Lemoenshoek and Naauwkrantz, Strawberry Hill (– DD), Stokoe s.n. (NBG). – 3419 (Worcester): Caledon (– AB), Zeyher s.n. (SAM); Steenboksberg (– AD), Taylor 6507 (NBG); Robertson, Boesmanskloof Pass at McGregor (– BA), Leipoldt 3153 (BOL). – 3420 (Swellendam): Nasionale Bontebok Park (– AB), Acocks 22894 (PRE), Taylor 4262 (PRE); Bontebok Park (– AB), Grobler 513 (NBG, PRE), Liebenberg 6463 (NBG), Taylor s.n., 4252 (NBG, PRE), Van Wyk 2979 (JRAU); Swellendam (– AB), Galpin 3721 (PRE), Kennedy s.n. (SAM); Robindale, Bredasdorp (– AB), Viviers 1179 (NBG, PRE); Buffeljagsrivier (– BA), Anon s.n. (SAM), Van der Merwe 2704 (PRE); Grootvadersbosch (– BB), Meyer 1829 (S), Ruiters 28 (NBG, PRE); Naauwpoort (– BB), Thorne, s.n. (SAM). − 3421 (Riversdale): Aasvoëlbergnek (– AB), Horn 649 (PRE)., Published as part of le Roux, M. M. & Van Wyk, B. - E., 2007, A revision of Lebeckia sect. Lebeckia: The L. sepiaria group, pp. 118-130 in South African Journal of Botany 73 (1) on pages 128-130, DOI: 10.1016/j.sajb.2006.09.005, http://zenodo.org/record/7958313, {"references":["Goldblatt, P., Manning, J. C., 2000. Cape Plants: A conspectus of the Cape flora of South Africa. Strelitzia 9. National Botanical Institute, Pretoria and Missouri Botanical Garden, St. Louis."]}

4.2. L. brevicarpa L. brevicarpa M.M. le Roux and B.-E. van Wyk, sp. nov. L. ambiguae sepiariae similes sed leguminibus brevibus ovatis pauciseminalibus (in L. ambigua et L. sepiaria lineares multiseminales) differt. Type: South Africa, Pakhuis Pass [3219 AA], Le Roux and Van Wyk 5 (NBG, holotype; BOL, K, PRE, S, isotypes). Erect, glaucous suffrutex, up to 0.8 m in height. Leaves simple, acicular, articulated near the middle; leaf density 3–6 per 10 mm length of stem. Inflorescences terminal, 220– 280 mm long; rachis furrowed; flowers numerous, relatively dense (5–7 per 10 mm length of rachis), intermediate in size (8–11 mm long); pedicel 2–3 mm long; bract narrowly lanceolate, acuminate, 4–6 mm long; bracteoles narrowly triangular to lanceolate, acuminate, up to 2 mm long. Calyx 3.5–5.5 mm long, tube 2.5–3.0 mm long, lobes 1.5–2.5 mm long; subequally lobed but the upper lateral sinuses much wider than the medial sinus and lower lateral sinuses; lobes deltoid, tips minutely pubescent inside. Standard ovate to orbicular, 10–13 × 6.5–7.5 mm; apex obtuse; claw 2.0– 2.5 mm long. Wings oblong, longer than the keel, 9–11 × 3.5–4.0 mm, with 14–18 rows of sculpturing; apex obtuse; claws ± 3 mm long. Keel rostrate, 8–9 × ± 4 mm, usually without pockets; claws 3.0– 3.5 mm long. Pistil short-stipitate; ovary 8–10 × 0.7– 0.9 mm long, oblong; ovules few (6–10); style 3–5 mm long, curved upwards. Pods ovoid, up to 13 × 5 mm; indehiscent; terete; fruit wall thick, spongy. Seeds reniform–orbicular, 3.0 × 2.2 mm, indistinctly rugose, pale brown, mottled dark brown to black; hilum dark brown (Fig. 7). 4.2.1. Diagnostic characters L. brevicarpa is similar to L. ambigua and L. sepiaria, but differs markedly in its short, ovate, few-seeded pods (linear and many-seeded in L. ambigua and L. sepiaria). In L. brevicarpa the wings are longer than the keel and the bracts are less caducous than in L. sepiaria (which has wings that are shorter than the keel) (Fig. 7). 4.2.2. Distribution and habitat L. brevicarpa has a localised distribution and is restricted to altitudes above 400 m in the Bokkeveld and Cedarberg Mountains (Fig. 8), where it is locally common in deep sandy soil on disturbed roadsides. 4.2.3. Specimens examined – 3119 (Calvinia): Between Nieuwoudtville and Oorlogskloof (– AC), Leipoldt 3763 (BOL). – 3218 (Clanwilliam): Pakhuis Pass (– BB), Acocks 15019, 15039 (PRE), Bolus 8973 (PRE), Grobbelaar 2002 (PRE), Leach and Forrester 17424 (PRE), Le Roux, Boatwright, Magee and Van Wyk 4, 5 (JRAU), Marloth 9527 (PRE), Stirton 5937 (PRE), Story 2982 (PRE); Pakhuis Pass near Kleinkliphuis (– BB), Emdon 118 (PRE); Pakhuis Pass, between Clanwilliam and Calvinia (– BB), Schutte 261 (JRAU); Pakhuis Pass, Elandsfontein Farm (– BB), Stirton 9288 (PRE); Kleinkliphuis (– BB), Taylor 11023 (PRE); Piekenierskloof Pass (– DB), Le Roux, Boatwright, Magee and Van Wyk 8 (JRAU). – 3219 (Wuppertal): Pakhuis Pass (– AA), Bolus 8973, 23854 (BOL), Van Wyk 3123 (JRAU); Staatsbos, Cedarberg (– AC), Andrag 141 (PRE); Kleinplaas, Citrusdal (– AC), Hanekom 2896 (PRE); Algeria (– AC), Stirton 5916 (PRE); Algeria Forest Station (– AC), Stirton 5920 (PRE); Algeria, Grootberg (– AC), Van Wyk 2550 (JRAU, PRE); Grootberg, Cedarberg (– AC), Viviers 409 (PRE); Cedarberg Forest Reserve, Grootberg (– AC), Viviers 489 (NBG)., Published as part of le Roux, M. M. & Van Wyk, B. - E., 2007, A revision of Lebeckia sect. Lebeckia: The L. sepiaria group, pp. 118-130 in South African Journal of Botany 73 (1) on pages 124-126, DOI: 10.1016/j.sajb.2006.09.005, http://zenodo.org/record/7958313

3.2. The section Lebeckia Lebeckia Thunb. section Lebeckia. Type: L. sepiaria (L.) Thunb., chosen by Hutchinson in Gen. Flow. Pl.: 358 (1964). Spartium L., Pl. Rar. Afr.: 91 (1760), Sp. Pl.: 995 (1764), Gmelin, Syst. Nat. 2 (2): 1088 (1792), pro parte. Lebeckia Thunb., Prod. Pl. Cap.: 122 (1800), Fl. Cap.: 561 (1823), pro parte majore; Eckl. and Zeyh., Enum.: 192 (1836), pro parte majore; E. Mey. in Linnaea 7: 155 (1832), Comm. Pl. Afr. Austr. 1: 34 (1836), pro parte majore. Sarcophyllum E. Mey. in Linnaea 7: 155 (1832), Comm. Pl. Afr. Austr. 1: 34 (1836). Eu-Lebeckia Benth. in Hook., Lond. J. Bot. 3: 358 (1844), synon. nov. Phyllodiastrum Benth. in Hook., Lond. J. Bot. 3: 358 (1844), synon. nov. Erect to prostrate, slightly to markedly glaucous suffrutices. Leaves simple, acicular, glabrous, sparse or dense, spirally arranged, often articulated near the middle, mucronulate. Inflorescences terminal, short or elongate, laxly to densely manyflowered; rachis usually furrowed; flowers small or large (6– 18 mm long), yellow; bracts ovate to lanceolate, acuminate, caducous; bracteoles narrowly triangular, acuminate. Calyx equally to subequally lobed, lobes deltoid, ± as long as the tube, tips minutely pubescent on the inside. Petals yellow, sometimes fading to orange or rarely purple (L. longipes), invariably totally glabrous. Standard lanceolate to orbicular; apex obtuse. Wings oblong; apex obtuse; sculpturing present or absent. Keel rostrate, pockets sometimes present. Stamens fused into an open sheath, upper third free; anthers 10, 5 oblong and basifixed, 5 short and dorsifixed (vexillary stamen only slightly shorter than long anthers). Pistil sessile to markedly stipitate; ovary oblong to linear; ovules ± 6–20; style shorter than the ovary, curved upwards. Pods linear or ovoid, straight or slightly deflexed, up to 12 mm wide. Seeds reniform. 3.2.1. Diagnostic characters Species of Lebeckia section Lebeckia differ from all other species in the genus by their needle-like leaves, which are terete (never flat). 3.2.2. Distribution Lebeckia section Lebeckia (and also the L. sepiaria group) are endemic to the Cape Floristic Region, from Namaqualand to Port Elizabeth. Key to the species of Lebeckia section Lebeckia 1a. Keel spirally twisted: 2a. Fruit up to 50 mm long; short-lived fireweed; L. wrightii flowers 1–3 per inflorescence 2b. Fruit 60–120 mm long; perennial herb; L. pauciflora flowers more than 5 per inflorescence 1b. Keel not twisted: 3a. Calyx teeth longer than the tube L. grandiflora 3b. Calyx teeth shorter than the tube: 4a. Leaves articulated near the middle: 5a. Fruit long-stipitate (stipe 10–20 mm long); L. carnosa flowers very widely spaced on rachis 5b. Fruit sessile or short-stipitate (stipe less than 5 mm long), flowers usually densely spaced on rachis (if somewhat widely spaced, then the fruit is flat): 6a. Fruit ovoid (up to 13 mm long) L. brevicarpa 6b. Fruit oblong to linear (more than 15 mm long): (continued on next page) 7a. Fruit oblong (up to 6x longer than wide), flat, upper suture with distinct margin or wing: 8a. Flowers 6–11 mm long; L. meyeriana fruit 20–35 mm long 8b. Flowers ca. 15 mm long; L. macowanii fruit ± 50 mm long 7b. Fruit linear (more than 10× longer than wide), terete or semi-terete, upper suture without a margin or wing: 9a. Flowers usually up to 8 mm long; L. gracilis upper lateral sinuses of calyx± as wide as medial sinus and lower lateral sinuses; southern coastal distribution from Bredasdorp to Port Elizabeth 9b. Flowers more than 9 mm long; upper lateral sinuses of calyx much wider than medial sinus and lower lateral sinuses; western coastal distribution from Namaqualand and inland to the Cedarberg and eastwards to Heidelberg: 10a. Fruit dehiscent, not spongy, subterete, L. ambigua ± 2 mm wide, sometimes falcate 10b. Fruit indehiscent, spongy, terete, L. sepiaria more than 4 mm wide, straight, often somewhat torulose 4b. Leaves not articulated: 11a. Fruit long-stipitate (stipe 15 mm or longer) L. longipes 11b. Fruit sessile or short-stipitate L. plukenetiana (stipe up to 5 mm long) Within the section Lebeckia, there are four species groups: (1) the L. sepiaria group — four species with terete or semi-terete, ± sessile pods and a generally erect, virgate habit (Fig. 1). (2) the L. plukenetiana E. Mey. group — five species with stipitate, flat pods of up to 75 mm long. (3) the L. wrightii (Harv.) Bolus group – one species – a short-lived fireweed with stipules and flat pods. (4) the L. pauciflora group — two species with relatively long calyx lobes and linear, stipitate, semi-terete pods of more than 70 mm long., Published as part of le Roux, M. M. & Van Wyk, B. - E., 2007, A revision of Lebeckia sect. Lebeckia: The L. sepiaria group, pp. 118-130 in South African Journal of Botany 73 (1) on pages 121-122, DOI: 10.1016/j.sajb.2006.09.005, http://zenodo.org/record/7958313