Tenicroa Raf., Fl. Tellur. Thuranthos C.H.Wright, 1916

23. Tenicroa Raf., Fl. Tellur. View in CoL View at ENA

3: 52 (1837) ( Figs 56–59 View FIGURE 56 View FIGURE 57 View FIGURE 58 View FIGURE 59 ). Typus generis:— Tenicroa fragrans (Jacq.) Raf. View in CoL ≡ Anthericum fragrans Jacq. View in CoL ≡ Drimia fragrans (Jacq.) J.C.Manning & Goldblatt View in CoL (holotype).

= Pilasia Raf., Fl. Tellur. View in CoL 3: 53 (1837). Typus generis:— P. filifolia (Jacq.) Raf. View in CoL ≡ Anthericum filifolium Jacq. View in CoL ≡ Tenicroa filifolia (Jacq.) Oberm. View in CoL ≡ Drimia filifolia (Jacq.) J.C.Manning & Goldblatt View in CoL (holotype).

= Sypharissa Salisb., Gen. Pl. View in CoL [Salisbury]: 37 (1866) ≡ Urginea sect. Sypharissa (Salisb.) Baker in J. Linn. Soc., Bot. 13: 216 (1873) ≡ Drimia sect. Sypharissa (Salisb.) J.C.Manning & Goldblatt in Strelitzia 40: 111 (2018). Typus generis:— S. exuviata (Jacq.) Salisb. ex Oberm. View in CoL ≡ Anthericum exuviatum Jacq. View in CoL ≡ Tenicroa exuviata (Jacq.) Speta View in CoL ≡ Drimia exuviata (Jacq.) Jessop View in CoL (lectotype designated by Obermeyer in Bothalia 13: 111. 1980).

= Drimia sect. Juncifoliae J.C.Manning & Goldblatt in Strelitzia 40: 107 (2018). Typus sectionis:— D. juncifolia J.C.Manning & J.M.J.Deacon View in CoL ≡ Tenicroa juncifolia (J.C.Manning & J.M.J.Deacon) Mart. View in CoL -Azorín et al. (holotype).

.

Description:—Medium-sized bulbous geophyte. Bulb hypogeal, ovoid to subglobose, up to 7 cm in diam., extended into neck surrounded by cataphylls with raised, transverse purple to brown ribs, inner scales white to pinkish, fleshy, compact or rarely loose, spirally arranged, rarely distichous, sometimes fibrose.Roots distinctly thickened and branched. Leaves 1‒50 per bulb, 3‒80 cm long, narrowly linear, from terete to channelled, synanthous or rarely hysteranthous, entire or rarely scabrid on margins, green or somewhat glaucous, usually leathery, sometimes twisted, smooth, glabrous. Inflorescence a long, erect, multiflowered raceme, 2‒25 cm long, with up to 60 flowers; peduncle 10‒40 cm long, green, grey or purple, terete, erect, smooth and glabrous; pedicels 3‒15 mm long, shorter than tepals at anthesis, spreading in flower and erect in fruit. Bracts narrowly lanceolate, long acuminate, lowermost auriculate with long spur, sometimes bifid or branched and sinuous; bracteoles absent. Flowers stellate with patent tepals, diurnal. Tepals 6, biseriate, 6‒17 mm long, free, ovate-lanceolate to elliptic, white with well defined, narrow, purplish longitudinal band. Stamens 6, suberect and slightly spreading; filaments filiform, 4‒8 mm long, tapering, smooth, white, slightly sigmoid and curving distally; anthers yellow, erect, basifixed, deshicing longitudinally along their whole length, with yellow pollen. Ovary ovate-oblong, green to yellow. Style 4‒8 mm long, declinate, sigmoid, white or rarely tinged with purple, with capitate, papillose stigma. Capsule ovate to ellipsoid, 6‒25 mm long, valves completely dehiscing from base, tepals cohering and inrolled above ovary after anthesis, circumcissile from base and persisting as a cap at the top of the developing capsules. Seeds subellipsoid to compressed, (4–) 5‒11 mm long, flattened, embryo prominent with wide or short wings, testa black, dull or glossy, with sinuous cell walls.

Number of species and distribution:— Tenicroa includes twelve species restricted to southern and western South Africa and southern Namibia, it being therefore restricted to the Cape and Karoo-Namib Regions (sensu Taktajan 1986) ( Fig. 49 View FIGURE 49 ). For complete species characteristics see Pinter et al. (2020).

Karyology:—2n=20 ( Speta 1998a).

History, diagnostic characters, and taxonomic relationships:— Rafinesque (1837) described Tenicroa to include T. fragrans ( Jacquin 1797: t. 86) Rafinesque (1837: 53), a species with rotate flowers, suberect stamens and deflexed style, among other characters. The revision by Obermeyer (1980b, 1981a) accepted four species in the genus from southern and western South Africa and southern Namibia: T. fragrans , T. exuviata (Jacquin 1794: 18) Speta (1980: 195) , T. filifolia (Jacquin 1794: 18) Obermeyer (1981a: 577) , and T. multifolia (Lewis: 1952: 9) Obermeyer (1981a: 577). All of them shared distinct characters such as the synanthous leaves surrounded by sheathing cataphylls that are markedly striate with raised, dark, transversal ridges; the stellate flowers with almost free tepals with a narrow, longitudinal dark band on the abaxial side; the suberect to slightly spreading stamens with basifixed anthers and the elongate, deflexed and curved style; and papillate stigma. Despite the distinct morphology of taxa in this genus, they have historically been placed in 8 different genera: Anthericum Linnaeus (1753: 310) , Albuca , Drimia , Ornithogalum , Phalangium Miller (1754 : s.n.), Pilasia Rafinesque (1837: 53) , Sypharissa Salisbury (1866: 37) , and Urginea , representing a good example of the uncertainty in taxonomy and systematics of urgineoid taxa ( Pinter et al. 2020). However, based on the clear synapomorphies of Tenicroa , researchers in Urgineoideae ( Jessop 1977, Obermeyer 1980b, 1981a, Speta 1998a, 1998b) usually recognised it as a distinct genus. Conversely, Manning et al. (2004) synonymised Tenicroa to Drimia sensu lato, and later Manning & Goldblatt (2018) placed the Tenicroa species recognised by Obermeyer (1980b) in D. sect. Sypharissa , whilst including T. filifolia as synonym of T. exuviata . They also separated their new D. sect. Juncifoliae to associate two newly described species [ Drimia decipiens Manning & Goldblatt (2018: 109) and D. juncifolia J.C.Manning & J.M.J.Deacon in Manning & Goldblatt (2018: 107)] based on their hysteranthous leaves, weakly barred cataphylls, and medifixed anthers that are not deflexed. Martínez-Azorín et al. (2019a) considered the latter distinction insufficient to recognise both sections, and transferred the two new species to Tenicroa .

The phylogenetic analyses of Manning et al. (2004) recovered two samples of Tenicroa as monophyletic, when excluding T. nana as the monotypic Mucinaea ( Pinter et al. 2013) . Pfosser et al. (2012) phylogenetically analysed nine samples of Tenicroa that formed a perfectly supported clade being sister to a clade comprising Litanthus plus Schizobasis . Furthermore, the phylogenetic analyses of Martínez-Azorín et al. (2023a) included 21 samples of Tenicroa , which formed a strongly supported and isolated clade within a large polytomy. Therefore, we here accept Tenicroa at genus rank based on the unique syndrome of morphological characters and its isolated phylogenetic position, following the latest revision of Tenicroa by Pinter et al. (2020), who accepted twelve species restricted to South Africa and southern Namibia.

Accepted species:—

Tenicroa applanata M.Pinter, Mart.- Azorín, M.B. Crespo & Wetschnig in Phyton (Horn, Austria) 60: 64 (2020) ( Fig. 58.1 View FIGURE 58 ). Type:— SOUTH AFRICA. Western Cape. Cape Town (3318): near Tinie Versveld Wild Flower Reserve , between Darling and Yzerfontein, (–AD), elev. 122 m, 20 September 2015, M. Martínez-Azorín, M. Pinter, M.B. Crespo & M.Á. Alonso MMA1128 (GRA! holo.; ABH! iso.).

Tenicroa decipiens (J.C.Manning & Goldblatt) Mart.- Azorín, M.B. Crespo, M. Pinter & Wetschnig in Phytotaxa 397(4): 294 (2019) View in CoL ≡ Drimia decipiens J.C.Manning & Goldblatt View in CoL in Strelitzia 40: 109 (2018), basionym. Type:— SOUTH AFRICA. Western Cape. Wuppertal (3219): Swartruggens , near turnoff to Kagga Kamma at summit of Skitterykloof , (–DC), 27 November 2017, J. Manning 3637 (NBG holo.).

Tenicroa exuviata (Jacq.) Speta View in CoL in Linzer Biol. Beitr. 12(1): 195 (1980) ≡ Anthericum exuviatum Jacq., Icon. Pl. Rar. View in CoL 2 (13): 18, t. 415 (1794), basionym ( Fig. 56.2 View FIGURE 56 ) ≡ Phalangium exuviatum (Jacq.) Poir. in Lam., Encycl. 5: 243 (1804) ≡ Albuca exuviata (Jacq.) Ker Gawl. in Curtis’s Bot. Mag. 22: t. 871 (1805) ( Fig. 57.1 View FIGURE 57 ) ≡ Urginea exuviata (Jacq.) Steinh. View in CoL in Ann. Sci. Nat., sér. 2, 2: 330 (1834) ≡ Ornithogalum exuviatum (Jacq.) Kunth, Enum. Pl. View in CoL 4: 369 (1843) ≡ Drimia exuviata (Jacq.) Jessop View in CoL in J. S. African Bot. 43(4): 276 (1977) ≡ Sypharissa exuviata (Jacq.) Salisb. ex Oberm. in Bothalia 13(1-2): 113 (1980) ( Figs 4.19 View FIGURE 4 , 58.2 View FIGURE 58 ). Type:— SOUTH AFRICA. Crescit ad Promontorium bonae Spei, icon in Jacq., Icon. Pl. Rar. 2: t. 415 (1794) ( Fig. 56.2 View FIGURE 56 ) (lecto. designated as “ Type ” by Jessop in J. S. African Bot. 43: 276. 1977). Epitype (designated by Pinter et al. 2020):— SOUTH AFRICA. Bredasdorp (3420): N side of Potberg, Bredasdorp Div., (–BC), elev. ca. 400 ft., 13 November 1962, J.P.H. Acocks 23016 (PRE0046767-0! epi.).

Tenicroa fibrosa M.Pinter, Mart.- Azorín, M.B. Crespo & Wetschnig in Phyton (Horn, Austria) 60: 70 (2020) ( Fig. 58.3 View FIGURE 58 ). Type:— SOUTH AFRICA. Western Cape. Vanrhynsdorp (3118): 21 m S.S. W. of Vredendal , (–CD), elev. ca 500 ft., 26 August 1958, J.P.H. Acocks 19713 (PRE0046760! holo.; BOL!, M0223028! iso.).

Tenicroa filifolia (Jacq.) Oberm. View in CoL in J. S. African Bot. 47(3): 577 (1981) ≡ Anthericum filifolium Jacq., Icon. Pl. Rar. View in CoL 2: 18, t. 414 (1794), basionym ( Fig. 56.3 View FIGURE 56 ) ≡ Phalangium filifolium (Jacq.) Poir. in Lam., Encycl. 5: 242 (1804) ≡ Albuca filifolia (Jacq.) Ker Gawl. View in CoL in Bot. Reg. 7: 557 (1821) ( Fig. 56.4 View FIGURE 56 ) ≡ Urginea filifolia (Jacq.) Steinh. View in CoL in Ann. Sci. Nat., sér. 2, 2: 329 (1834) ≡ Pilasia filifolia (Jacq.) Raf., Fl. Tellur. 3: 53 (1837) ≡ Ornithogalum filifolium (Jacq.) Kunth, Enum. Pl. View in CoL 4: 369 (1843) ≡ Sypharissa filifolia (Jacq.) Salisb. ex Oberm. View in CoL in Bothalia 13: 113 (1980) ≡ Drimia filifolia (Jacq.) J.C.Manning & Goldblatt View in CoL in Strelitzia 9: 711 (2000) ( Fig. 58.4 View FIGURE 58 ). Type:—Illustration in Jacq., Icon. Pl. Rar. 2 (15): t. 414 (1794) ( Fig. 56.3 View FIGURE 56 ) (lecto. designated as “ Type ” by Jessop in J. S. African Bot. 43: 276. 1977).

= Anthericum spiratum Thunb., View in CoL Prodr.: 62 (1794). Syntypes:— SOUTH AFRICA. ‘Cap. B. spei’, Thunberg s.n. (UPSTHUNB [8413 & 8414]-microfiche, syn.).

Tenicroa flexuosa (Adamson) M.Pinter, Mart.- Azorín, M.B. Crespo & Wetschnig in Phyton (Horn, Austria) 60: 76 (2020) ≡ Urginea flexuosa Adamson View in CoL in J. S. African Bot. 8: 240 (1942), basionym. Type:— SOUTH AFRICA. Western Cape. Simonstown (3418): Cape Peninsula , Smitswinkel Bay , (–AD), 3 January 1941, Adamson 3099 (BOL140326! holo.; NBG sub SAM0056131-0!, K000257378! iso.).

Tenicroa fragrans (Jacq.) Raf., Fl. Tellur. 3: 53 (1837) ≡ Anthericum fragrans Jacq., Hort. Schoenbr. 1: t. 86 (1797), basionym ( Fig. 56.1 View FIGURE 56 ) ≡ Phalangium fragrans (Jacq.) Poir. in Lam., Encycl. 5: 247 (1804) ≡ Albuca fugax Ker Gawl. in Bot. Reg. 4: t. 311 (1818), nom. nov. ( Fig. 57.2 View FIGURE 57 ) [non Albuca fragrans Jacq. ] ≡ Urginea fragrans (Jacq.) Steinh. View in CoL in Ann. Sci. Nat., sér. 2, 2: 328 (1834) ≡ Ornithogalum fragrans (Jacq.) Kunth, Enum. Pl. View in CoL 4: 366 (1843) ≡ Sypharissa fragrans Salisb. ex Oberm. in Bothalia 13(1−2): 113 (1980) ≡ Drimia fragrans (Jacq.) J.C.Manning & Goldblatt View in CoL in Strelitzia 9: 711 (2000) ≡ Albuca fugax Ker Gawl. in Bot. Reg. 4: t. 311 (1818), syn. subst. ( Figs 4.20 View FIGURE 4 , 59.1 View FIGURE 59 ). Type:— SOUTH AFRICA. Western Cape: illustration in Jacq., Hort. Schoenbr. 1: t. 86 (1797) ( Fig. 56.1 View FIGURE 56 ) (lecto. designated as “ Type ” by Jessop in J. S. African Bot. 43: 276. 1977). Epitype (designated by Pinter et al. 2020):— SOUTH AFRICA. Western Cape. Vanrhynsdorp (3118): Summit of Gifberg, (−DA), elev. ca. 2000 ft., 16 November 1970, H. Hall 3906 (NBG91254!, 2 sheets epi.; PRE0046756-0! isoepi.).

Tenicroa juncifolia (J.C.Manning & J.M.J.Deacon) Mart.- Azorín, M.B. Crespo, M. Pinter & Wetschnig in Phytotaxa 397: 294 (2019) View in CoL ≡ Drimia juncifolia J.C.Manning & J.M.J.Deacon View in CoL in Strelitzia 40: 107 (2018), basionym. Type:— SOUTH AFRICA. Western Cape. Cape Town (3318): Paarl , Brier’s Louw Nature Reserve , (–DD), 23 November 2013, J.M.J. Deacon 3078 (NBG holo.).

Tenicroa multifolia (G.J.Lewis) Oberm. View in CoL in J. S. African Bot. 47(3): 577 (1981) ≡ Urginea multifolia G.J.Lewis View in CoL in Ann. S. African Mus. 40: 9 (1952), basionym ( Fig. 57.3 View FIGURE 57 ) ≡ Drimia multifolia (G.J.Lewis) Jessop View in CoL in J. S. African Bot. 43(4): 278 (1977) ≡ Sypharissa multifolia (G.J.Lewis) Oberm. in Bothalia 13(1−2): 114 (1980) ( Figs 5.1 View FIGURE 5 , 59.2 View FIGURE 59 ). Type:— SOUTH AFRICA. Northern Cape. Springbok (2917): 27 miles S of Springbok, (–DD), 27 July 1950, Lewis 2302 (SAM0060870-0! holo.; NBG! iso.).

= Urginea capillifolia Schlechter , nom. nud. in sched.

Tenicroa namibensis M.Pinter, Mart. - Azorín, M.B. Crespo & Wetschnig in Phyton (Horn, Austria) 60: 83 (2020) ( Fig. 59.3 View FIGURE 59 ). Type:— NAMIBIA. Witputz (2716): Karas, (–AC), elev. 797 m, 12 August 2001, C.A. Mannheimer CM1607 (WIND77423! holo.).

Tenicroa polyantha M.Pinter, Mart. - Azorín, M.B. Crespo & Wetschnig in Phyton (Horn, Austria) 60: 84 (2020) ( Figs 5.2 View FIGURE 5 , 59.4 View FIGURE 59 ). Type:— SOUTH AFRICA: Western Cape. Cape Town (3318): Langebaan , eastern end of town, (–AA), elev. 55 m, 20 September 2015 [in flower], M. Martínez-Azorín, M. Pinter, M.B. Crespo & M.Á. Alonso MMA1139 (GRA! holo.; ABH! iso.).

Tenicroa unifolia (A.V.Duthie) M.Pinter, Mart. - Azorín, M.B. Crespo & Wetschnig in Phyton (Horn, Austria) 60: 86 (2020) ≡ Urginea unifolia A.V.Duthie in Ann. Univ. Stell. 6A: 8 (1928), basionym ( Figs 5.3 View FIGURE 5 , 59.5 View FIGURE 59 ). Type:— SOUTH AFRICA. Western Cape. Cape Town (3318): Stellenbosch Flats , (–DD), October 1927, A.V. Duthie s.n. STE1891 (NBG0197708-1! holo.; BOL140325! iso.).

= Urginea ecklonii sensu Duthie View in CoL in Ann. Univ. Stell. 6 A: 6 (1928), non U. ecklonii Baker (1892) View in CoL .

= Urginea duthieae Adamson View in CoL in J. S. African Bot. 8: 239 (1942) ≡ Drimia duthieae (Adamson) Jessop View in CoL in J. S. African Bot. 43(4): 278 (1977). Type:— SOUTH AFRICA. Western Cape. Cape Town (3318): Stellenbosch Flats, (–DD), 17 October 1925, Duthie s.n. STE1790 (NBG0197709-0! holo.; K000400569! iso.). 24. Thuranthos C.H.Wright View in CoL View at ENA

in Bull. Misc. Inform. Kew 1916(9): 233 (1916) ( Figs 60 View FIGURE 60 , 61 View FIGURE 61 ).

Typus generis:— T. macranthum (Baker) C.H.Wright View in CoL (holotype).

≡ Drimia sect. Thuranthos (C.H.Wright) J.C.Manning & Goldblatt in Strelitzia 40: 62 (2018) pro parte. Typus sectionis:— D. macrantha View in CoL (Baker) Baker (holotype).

Description:—Medium to tall bulbous geophyte. Bulb hypogeal, ovoid to subglobose, 3‒15 cm in diam., with white, fleshy, spathulate and commonly pediculate, loose scales. Roots thickened and branched. Leaves 2‒7(‒14) per bulb, hysteranthous, 10‒70 cm long, narrowly linear, channelled and usually keeled, green, sometimes basally maculate, somewhat glaucous, smooth, glabrous. Inflorescence a lax, erect raceme, 15‒50 cm long, with 3–40 flowers; peduncle 30‒150 cm long, erect, terete, smooth; pedicels (10–) 30‒70 mm long, subpatent and arching downwards distally at anthesis, erect in fruit. Bracts ovate-lanceolate, 3‒10 mm long, lowermost with distinct spur, early caducous at flowering and leaving a distinct scar; bracteoles present and evident, also early caducous. Flowers stellate with strongly reflexed tepals at full anthesis, nodding, nocturnal, opening in late evening and closing before sunrise, fragrant during night. Tepals 6, biseriate, (10‒) 15‒37 mm long, narrowly oblong, obtuse, yellow, brown, carneous or whitish with darker brownish-greenish longitudinal band more evident on abaxial side, almost free to very shortly connate for 1‒2 mm. Stamens 6, erect, sigmoid, yellow, brown to white or cream colour; filaments 7‒20 mm long, linear, flattened and expanded below and incurved, leaving opening among them to show ovary, connivent to style at upper half/third, and slightly spreading distally and commonly pale in colour, adnate to tepals at base; anthers oblong, 2‒7 mm long, basifixed, yellow, orange or green, with yellow or white pollen. Ovary ovate to conical, 4.0‒ 6.5 mm long, greenish. Style 7‒22 mm long, erect, white, columnar, ending in small or distinctly capitate, thickened stigma. Capsule ovate to oblong or ellipsoid, trigonous, deeply 3-lobed, 10‒50 mm long, pale brown, valves completely dehiscing from base, tepals cohering and inrolled above ovary after anthesis, circumcissile from base and persisting as a cap at the top of the developing capsules. Seeds ellipsoidal to broadly ellipsoid, flattened, 6‒16 mm long, testa black, glossy, with slightly sinuous anticlinal cell walls.

View Figure View Figure

Number of species and distribution:— Thuranthos includes eight species mostly confined to Southern Africa and East Africa, being in the Uzambara-Zululand Region and the Southern and Eastern Sections of the Zambezian Subregion, with a species occurring in the Karoo‐Namib Region (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a), ( Fig. 49 View FIGURE 49 ). For further species characterisation see Dyer (1964), Stirton (1976), Obermeyer (1980c), Stedje (1987), Speta (1998a), Newton (2003), and Manning & Goldblatt (2018).

Karyology:—2n=20 ( De Wet 1957, as Thuranthos macranthum (Baker) C.H.Wright ); 2n=20+4B ( Stedje 1996 as Drimia macrocarpa Stedje ).

History, diagnostic characters, and taxonomic relationships:— Wright (1916) described Thuranthos based on a distinct species with a remarkable, unique flower morphology within Urgineoideae at that time, showing nocturnal, large, nodding flowers with filaments that Stirton (1976) considered an outstanding feature showing flattened, convex basal regions that are connivent to the style at the middle and spread above, resembling a paper lantern with longitudinal slits ( Figs 60 View FIGURE 60 , 61 View FIGURE 61 ). Speta (1998a) accepted this genus based on its evident morphological differentiation from other urgineoids. This genus was characterised by the imbricate, loose bulb scales; keeled, proteranthous leaves; the presence of bracteoles; the nodding, nocturnal flowers with the peculiar morphology of filaments described above, the erect style and the large, flat, ovate seeds.

The phylogenetic study of Martínez-Azorín et al. (2023a) included six samples of Thuranthos macranthum , T. nocturnale Dyer (1964 : t. 1439), and T. pauciflorum ( Baker 1892: 6) Martínez-Azorín et al. (2019a: 295) , the latter including T. basuticum ( Phillips 1917: 306) Obermeyer (1980c: 139) in synonymy (see Martínez-Azorín et al. 2019a). This is a genus restricted to Southern Africa and East Africa, and one which formed a strongly supported clade included in a polytomy with the monophyletic and well supported Ledurgia and Zingela . Martínez-Azorín et al. (2023a) considered Thuranthos to resemble Vera-duthiea on account of the nodding, nocturnal flowers with filaments usually approaching the style at the middle, but differing in the style, which is distinctly deflexed, the spotted, not keeled leaves, the lack of bracteoles [except in V. macrocarpa (Stedje 1987: 664) Speta (2016: 155) ], and in their different phylogenetic relationships, as it is sister to Ebertia ( Martínez-Azorín et al. 2023a) . Our recent unpublished phylogenetic analyses of samples of Drimia macrocarpa Stedje (1987: 664) from Mapanza Mission in Zambia (Robinson 316 K! number H.212/95 90; seeds kindly provided by Royal Botanic Gardens, Kew), show its close relation to Thuranthos instead of Vera-duthiea . This recent finding facilitates the refinement of the circumscription of the latter two genera and therefore, Thuranthos is expanded to include the species with loose bulb scales, nodding, nocturnal flowers with basally expanded filaments that are sigmoid and connivent to the style along the middle portion and spreading above and presence of bracteoles and Vera-duthiea , including species with compact bulb scales, absence of bracteoles and filaments narrow and contracted at the joint to tepals, among other characters. This new circumscription of the two genera eliminates the exception of the presence of bracteoles and loose bulb scales in D. macrocarpa when included in Vera-duthiea by Speta (2001, 2016) and Martínez-Azorín et al. (2018b, 2023a).

Samples of Zingela consistently form a well‐supported clade sharing with Thuranthos the loose bulb scales; hysteranthous leaves and presence of bracteoles; however, Zingela differs by the diurnal flowers and filiform, spreading stamens with circinnate dehisced anthers ( Crouch et al. 2018), and are placed in a polytomy with Thuranthos and Ledurgia ( Fig. 2 View FIGURE 2 in Martínez-Azorín et al. 2023a), which is resolved in the parsimony analyses ( Figs S5 View FIGURE 5 , S 7 View FIGURE 7 in Martínez-Azorín et al. 2023a) where Ledurgia and Zingela form sister clades.

Therefore, we accept Thuranthos as a distinct genus following Stirton (1976), Obermeyer (1980c) and Speta (1998a), though with the necessary recircumscription.

Manning & Goldblat (2018) accept D. sect. Thuranthos to include five species from Southern Africa: D. macrantha , D. basutica , D. vespertina (a newly described species from northern Namibia and southern Angola), D. hesperantha (= Urginea revoluta ), and D. indica . As reported by Martínez-Azorín et al. (2018b), Drimia indica belongs to Indurgia , a genus restricted to India and neigbouring countries, and plants identified as D. indica in South Africa were recently described as Vera-duthiea zebrina Martínez-Azorín et al. (2018b: 285) and Zingela pooleyorum Crouch et al. (2018: 36) . Similarly, D. vespertina seems to better fit the concept of Thuranthos of this work based on flower morphology and the expanded filaments, though the compact bulb scales, the absence of bracteoles, the general small plant size and its distribution makes its inclusion in the genus deviant. Further studies are needed to ascertain its phylogenetic relationships. Finally, Urginea revoluta is excluded from Thuranthos based on flower and seed morphology and our phylogenetic results ( Martínez-Azorín et al. 2023a).

Accepted species and required new combination and new name:—

Thuranthos bakeri Mart.-Azorín, M.B.Crespo & M.Á.Alonso nom. nov. ( Figs 5.4 View FIGURE 5 , 61.1 View FIGURE 61 ) ≡ Drimia elata var. cooperi Baker View in CoL in J. Linn. Soc. 11: 422 (1871), syn. subst. ≡ Drimia angustifolia Baker, Fl. Cap. (Harvey) 6(3): 440 (1897), nom. illeg. [non Drimia angustifolia Kunth, Enum. Pl. View in CoL 4: 340 (1843), nec D. cooperi Baker, Refug. Bot. [Saunders] 1: pl. 18 (1869)]. Type:— SOUTH AFRICA. Eastern Cape. Fort Beaufort (3226): near Tarkastad, (–AB), 1860, Cooper 387 (K000400568! holo.).

Thuranthos laxiflorum (Baker) Mart.- Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia laxiflora Baker View in CoL in J. Linn. Soc., Bot. 11: 422 (1870), basionym. Type:— MOZAMBIQUE. Zambesi-land , prope Quillimane [Quelimane], November 1862, Dr. Kirk s.n. (K0000400585! holo.).

Thuranthos macranthum (Baker) C.H.Wright View in CoL in Bull. Misc. Inform. Kew 1916(9): 223 (1916) ≡ Ornithogalum macranthum Baker View in CoL in J. Linn. Soc., Bot. 13: 280 (1873), basionym ≡ Drimia macrantha (Baker) Baker in Bot. Jahrb. 15 (3, Beibl. 35): 7 (1892) ≡ Urginea macrantha (Baker) E.P.Phillips View in CoL in Ann. S. African Mus. 16: 305 (1917) ( Figs 60.1 View FIGURE 60 , 61.2 View FIGURE 61 ). Type:— SOUTH AFRICA. Eastern Cape. Uitenhage Division, Van Stadens River, elev. under 200 ft., December, Drège 2204 (K000257227! lecto. designated here; K000257228!, S06-4414!, isolecto.). Note:—Lectotypification designated by Jessop in J. S. African Bot. 43: 275 (1977) is disregarded since no specimen of that collection is kept at BM.

Thuranthos macrocarpum (Stedje) Speta in Phyton (Horn, Austria) 56(2): 155 (2016) ≡ Drimia macrocarpa Stedje View in CoL in Nordic J. Bot. 7(6): 664 (1987), basionym ≡ Thuranthos macrocarpus (Stedje) Speta in Phyton (Horn, Austria) 38(1): 84 (1998) ≡ Duthiea macrocarpa (Stedje) Speta in Stapfia 75: 170 (2001) ( Figs 5.5 View FIGURE 5 , 61.3 View FIGURE 61 ). Type:— TANZANIA. Mpanda District, Uruwira, elev. 3600 ft., 23 September 1970, H.M. Richards & S. Arasululu 26126 (K000257337! holo.).

= Ornithogalum laikipiense L.E.Newton in Plantsman n.s. 2(1): 18 (2003). Type:— KENYA. Laikipia Plateau, 56 km NE of Rumuruti, Colcheccio Ranch , elev. 1750 m, April/ May 1995, M. Roberts sub Newton 5567 (K000365655! Sheet 1/2 lecto. designated here; K000365654! Sheet 2/2, isolecto.; EA isolecto., only the inflorescence). Note:— The type material at Kew includes two herbarium vouchers with different herbarium barcodes. Although not specified, leaves and flowers were most probably collected at different times (Hysteranthous: April–May as indicated on the voucher labels). Despite the current wording of Art. 8 of the ICN, and following amendment proposal by Mosyakin & McNeill (2022), a lectotypification is here proposed. Further, the type material at EA includes both leaves and inflorescences in a single voucher, and according to our proposal, only the inflorescence is regarded as isolectotype.

Thuranthos nocturnale R.A.Dyer in Fl. Pl. Africa 36: t. 1439 (1964) ( Figs 5.6 View FIGURE 5 , 60.2 View FIGURE 60 , 61.4 View FIGURE 61 ). Type:— SOUTH AFRICA. Eastern Cape. Steynsburg (3125): Grootfontein, (–AC), November 1956, Acocks 18650 (PRE0046805-2! lecto. designated here). Note:—The type material (“ holotype ”) indicated in the protologue is currently conserved in four different herbarium sheets numbered 1–4, here regarded as syntypes. Each of those vouchers includes different fragments collected on different dates. Following amendment proposal of Art. 8 of the ICN by Mosyakin & McNeill (2022), a lectotypification is here proposed.

Thuranthos pauciflorum (Baker) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 295 (2019) ≡ Drimia pauciflora Baker in Bot. Jahrb. Syst. 15(3, Beibl. 35): 6 (1892), basionym [non Urginea pauciflora Baker (1898: 539) , nec Urginea pauciflora Baker (1901: 786) , nom. illeg.] ( Figs 5.7 View FIGURE 5 , 61.5 View FIGURE 61 ). Type:— SOUTH AFRICA. Pr. B. sp., Ecklon & Zeyher Asphod. 102 (B100167472! lecto. [or perhaps holo.] designated by Martínez-Azorín et al. 2019a: 295).

= Urginea basutica E.Phillips View in CoL in Ann. S. African Mus. 16: 306 (1917) ≡ Thuranthos basuticum (E.Phillips) Oberm. in Bothalia 13(1−2): 139 (1980) ≡ Drimia basutica (E.P.Phillips) J.C.Manning & Goldblatt View in CoL in Strelitzia 40: 68 (2018). Type:— LESOTHO. Bethlehem (2828): Leribe, (–CC), October–November 1911, A. Dieterlen 854 (PRE0048599-0! lecto. designated here; K!, SAM! isolecto.). Note:—The holotype designated by Manning & Goldblatt (2018: 64) is not correct according to Art. 9.23 of the Shenzhen Code ( Turland et al. 2018), since two vouchers at K (K000257224 and K000257225) are undoubtedly original material.

Thuranthos vespertinum (J.C.Manning & Goldblatt) Mart.- Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia vespertina J.C.Manning & Goldblatt View in CoL in Strelitzia 40: 65 (2018), basionym. Type:— NAMIBIA. Sesfontein (1913): Khowarib Gorge, (–BD), only the flowered material ex hort on 29 April 1991, P. Bruyns 4066 (NBG150472! holo.).

Thuranthos zambesiacum (Baker) Kativu View in CoL in Kirkia 15(1): 113 (1994) ≡ Urginea zambesiaca Baker View in CoL in J. Linn. Soc., Bot. 13: 223 (1873), basionym ≡ Drimia zambesiaca (Baker) J.C.Manning & Goldblatt View in CoL in Edinburgh J. Bot. 60(3): 557 (2004) ≡ Drimia zambesiaca (Baker) J.C.Manning & Goldblatt View in CoL in Bothalia 49(1): 4 (2019) isonym ≡ Vera-duthiea zambesiaca (Baker) N.R.Crouch & Mart. - Azorín, Phytotaxa 470(3): 261 (2020). Type:— MOZAMBIQUE. Lower Zambesi, mainland above Expedition Island, lower Zambesi, 20 November 1859, J. Kirk 128 (K000400577! holo.).