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

Martínez-Azorín, Mario, Crespo, Manuel B., Alonso-Vargas, María Ángeles, Pinter, Michael, Crouch, Neil R., Dold, Anthony P., Mucina, Ladislav, Pfosser, Martin & Wetschnig, Wolfgang, 2023, A generic monograph of the Hyacinthaceae subfamily Urgineoideae, Phytotaxa 610 (1), pp. 1-143 : 97-103

23. Tenicroa Raf., Fl. Tellur.

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

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

= Sypharissa Salisb., Gen. Pl. [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. ≡ Anthericum exuviatum Jacq. ≡ Tenicroa exuviata (Jacq.) Speta ≡ Drimia exuviata (Jacq.) Jessop (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 ≡ Tenicroa juncifolia (J.C.Manning & J.M.J.Deacon) Mart. -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 ). 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 ). 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) ≡ Drimia decipiens J.C.Manning & Goldblatt 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 in Linzer Biol. Beitr. 12(1): 195 (1980) ≡ Anthericum exuviatum Jacq., Icon. Pl. Rar. 2 (13): 18, t. 415 (1794), basionym ( Fig. 56.2 ) ≡ 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 ) ≡ Urginea exuviata (Jacq.) Steinh. in Ann. Sci. Nat., sér. 2, 2: 330 (1834) ≡ Ornithogalum exuviatum (Jacq.) Kunth, Enum. Pl. 4: 369 (1843) ≡ Drimia exuviata (Jacq.) Jessop 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 , 58.2 ). Type:— SOUTH AFRICA. Crescit ad Promontorium bonae Spei, icon in Jacq., Icon. Pl. Rar. 2: t. 415 (1794) ( Fig. 56.2 ) (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 ). 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. in J. S. African Bot. 47(3): 577 (1981) ≡ Anthericum filifolium Jacq., Icon. Pl. Rar. 2: 18, t. 414 (1794), basionym ( Fig. 56.3 ) ≡ Phalangium filifolium (Jacq.) Poir. in Lam., Encycl. 5: 242 (1804) ≡ Albuca filifolia (Jacq.) Ker Gawl. in Bot. Reg. 7: 557 (1821) ( Fig. 56.4 ) ≡ Urginea filifolia (Jacq.) Steinh. in Ann. Sci. Nat., sér. 2, 2: 329 (1834) ≡ Pilasia filifolia (Jacq.) Raf., Fl. Tellur. 3: 53 (1837) ≡ Ornithogalum filifolium (Jacq.) Kunth, Enum. Pl. 4: 369 (1843) ≡ Sypharissa filifolia (Jacq.) Salisb. ex Oberm. in Bothalia 13: 113 (1980) ≡ Drimia filifolia (Jacq.) J.C.Manning & Goldblatt in Strelitzia 9: 711 (2000) ( Fig. 58.4 ). Type:—Illustration in Jacq., Icon. Pl. Rar. 2 (15): t. 414 (1794) ( Fig. 56.3 ) (lecto. designated as “ Type ” by Jessop in J. S. African Bot. 43: 276. 1977).

= Anthericum spiratum Thunb., 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 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 ) ≡ 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 ) [non Albuca fragrans Jacq. ] ≡ Urginea fragrans (Jacq.) Steinh. in Ann. Sci. Nat., sér. 2, 2: 328 (1834) ≡ Ornithogalum fragrans (Jacq.) Kunth, Enum. Pl. 4: 366 (1843) ≡ Sypharissa fragrans Salisb. ex Oberm. in Bothalia 13(1−2): 113 (1980) ≡ Drimia fragrans (Jacq.) J.C.Manning & Goldblatt in Strelitzia 9: 711 (2000) ≡ Albuca fugax Ker Gawl. in Bot. Reg. 4: t. 311 (1818), syn. subst. ( Figs 4.20 , 59.1 ). Type:— SOUTH AFRICA. Western Cape: illustration in Jacq., Hort. Schoenbr. 1: t. 86 (1797) ( Fig. 56.1 ) (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) ≡ Drimia juncifolia J.C.Manning & J.M.J.Deacon 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. in J. S. African Bot. 47(3): 577 (1981) ≡ Urginea multifolia G.J.Lewis in Ann. S. African Mus. 40: 9 (1952), basionym ( Fig. 57.3 ) ≡ Drimia multifolia (G.J.Lewis) Jessop 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 , 59.2 ). 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 ). 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 , 59.4 ). 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 , 59.5 ). 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 in Ann. Univ. Stell. 6 A: 6 (1928), non U. ecklonii Baker (1892) .

= Urginea duthieae Adamson in J. S. African Bot. 8: 239 (1942) ≡ Drimia duthieae (Adamson) Jessop 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

Figures

FIGURE 4. Seed morphology in Urgineoideae. 1. Rhodocodon rotundus H.Perrier ex Knirsch et al.; 2. Rhodocodon siederi Knirsch et al.; 3. Rhodocodon urgineoides Baker; 4. Schizobasis intricata (Baker) Baker; 5. Schizobasis macowanii Baker; 6. Schizobasis sigmoidea (J.C.Manning & J.M.J.Deacon) Mart.-Azorín et al.; 7. Sekanama brachystachys (Baker) Mart.-Azorín et al.; 8. Sekanama porphyrostachys (Baker ex Balf.f.) Mart.-Azorín et al.; 9. Sekanama sanguinea (Schinz) Speta; 10. Squilla hesperia (Webb & Berthel.) Mart.-Azorín et al.; 11. Squilla maritima (L.) Steinh.; 12. Squilla palaestina (M.B.Crespo et al.) Mart.-Azorín et al.; 13. Squilla pancration Steinh.; 14. Squilla secundiflora (Maire) Mart.-Azorín et al.; 15. Squilla serotina (Schousb.) Mart.-Azorín et al.; 16. Squilla undulata (Desf.) Mart.-Azorín et al.; 17. Striatula oliverorum (J.C.Manning) M.Pinter et al.; 18. Striatula platyphylla (B.Nord.) M.Pinter et al.; 19. Tenicroa exuviata (Jacq.) Speta; 20. Tenicroa fragrans (Jacq.) Raf. All to the same scale in mm.

FIGURE 5. Seed morphology in Urgineoideae. 1. Tenicroa multifolia (G.J.Lewis) Oberm.; 2. Tenicroa polyantha M.Pinter et al.; 3. Tenicroa unifolia (A.V.Duthie) M.Pinter et al.; 4. Thuranthos bakeri Mart.-Azorín et al.; 5. Thuranthos macrocarpum (Stedje) Speta; 6. Thuranthos nocturnale R.A.Dyer; 7. Thuranthos pauciflorum (Baker) Mart.-Azorín et al.; 8. Triandra pellabergensis Mart.-Azorín et al.; 9. Urginavia altissima (L.f.) Speta; 10. Urginavia capensis (Burm.f.) Mart.-Azorín et al.; 11. Urginavia echinostachya (Baker) Mart.-Azorín et al.; 12. Urginavia kniphofioides (Baker) Mart.-Azorín et al.; 13. Urginavia micrantha (A.Rich.) Speta; 14. Urginavia multisetosa (Baker) Mart.-Azorín et al.; 15. Urginavia simensis (Hochst. ex A.Rich.) Mart.-Azorín et al.; 16. Urginea fugax (Moris) Steinh.; 17. Urgineopsis arenosa (Adamson) Mart.-Azorín et al.; 18. Urgineopsis dregei (Baker) Mart.-Azorín et al.; 19. Urgineopsis exilis (Adamson) Mart.-Azorín et al.; 20. Urgineopsis gracilis (A.V.Duthie) Mart.-Azorín et al. All to the same scale in mm.

FIGURE 49. Approximate distribution ranges of accepted genera in Urgineoideae.

FIGURE 56. Species of Tenicroa Raf. 1. Tenicroa fragrans (Jacq.) Raf. from Jacquin (1797: t. 86, as Anthericum fragrans Jacq.); 2. Tenicroa exuviata (Jacq.) Speta from Jacquin (1794: t. 415, as Anthericum exuviatum Jacq.); 3. Tenicroa filifolia (Jacq.) Oberm. from Jacquin (1794: t. 414, as Anthericum filifolium Jacq.); 4. Tenicroa filifolia (Jacq.) Oberm. from Ker Gawler (1821: t. 557, as Albuca filifolia (Jacq.) Ker Gawl.).

FIGURE 57. Species of Tenicroa Raf. 1. Tenicroa exuviata (Jacq.) Speta from Ker Gawler (1805: t. 871, as Albuca exuviata (Jacq.) Ker Gawl.); 2. Tenicroa fragrans (Jacq.) Raf. from Ker Gawler (1818: t. 311, as Albuca fugax Ker Gawl.); 3. Tenicroa multifolia (G.J.Lewis) Oberm. from Lewis (1952: 10, as Urginea multifolia G.J.Lewis).

FIGURE 58. Species of Tenicroa Raf. displayed in horizontal rows of images. 1. Tenicroa applanata M.Pinter et al.; 2. Tenicroa exuviata (Jacq.) Speta; 3. Tenicroa fibrosa M.Pinter et al.; 4. Tenicroa filifolia (Jacq.) Oberm.

FIGURE 59. Species of Tenicroa Raf. displayed in horizontal rows of images. 1. Tenicroa fragrans (Jacq.) Raf.; 2. Tenicroa multifolia (G.J.Lewis) Oberm.; 3. Tenicroa namibensis M.Pinter et al. (Photographs of bulb with leaf by H. Kolberg and in flower by A. Burke); 4. Tenicroa polyantha M.Pinter et al.; 5. Tenicroa unifolia (A.V.Duthie) M.Pinter et al.