Urginavia Speta

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 : 109-114

26. Urginavia Speta

in Phyton (Horn, Austria) 38(1): 86 (1998) ( Figs 63–66 ).

Typus generis:— U. micrantha (A.Rich.) Speta (holotype).

≡ Drimia sect. Urginavia (Speta) J.C.Manning & Goldblatt in Strelitzia 40: 13 (2018). Typus sectionis:— U. micrantha (A.Rich.) Speta (holotype).

Description:—Usually tall bulbous geophyte. Bulb hypogeal, rarely epigeal, ovoid to globose, up to 20 cm in diam., scales somewhat loose, heterogeneous in length and vertically imbricate, yellowish and leathery when dry and showing distinct white threads when broken, rarely showing fibrose neck above ground. Roots thickened and branched. Leaves (2–)4‒16 per bulb, narrowly lanceolate to oblong, hysteranthous or rarely partially synanthous, 8‒60 cm long, plane or channelled, sometimes keeled abaxially along lower portions, margin entire, green or somewhat glaucous, smooth, glabrous, producing white threads when broken. Inflorescence a long, erect, usually multiflowered raceme, with 15‒700 flowers, (3–) 10‒120 cm long; peduncle (6–) 10‒100 cm long, terete, erect, smooth and glabrous; pedicels 2‒30 mm long, subpatent or spreading. Bracts narrowly lanceolate, 2‒15 mm long, usually caducous or sometimes persisting, lowermost with spur from short to 20 mm long; bracteoles present and distinct. Flowers stellate or with reflexed tepals, erect-patent, diurnal, lasting one or more days. Tepals 6, biseriate, 3.5‒12.0 mm long, mostly subfree or rarely distinctly connate for 2‒3 mm at base, lanceolate-ovate to oblong, white, cream, brown or yellow, with darker longitudinal band. Stamens 6, suberect to spreading; filaments 2‒10 mm long, smooth, usually white and lanceolate to narrowly triangular, flattened or rarely filiform; anthers dorsifixed. Ovary ovate-oblong, 2‒5 mm long, green to yellow. Style 2‒9 mm long, erect, white with trigonous undifferentiated or capitate stigma. Capsule obovoid, ellipsoid or subglobose, 4‒15 mm long, emarginate, valves completely dehiscing from base, tepals remaining below developing capsules. Seeds elliptic or ovate-oblong, 4‒9 mm long, flattened, with prominent embryo and flat peripheral wings, testa black, with strongly sinuous anticlinal cell walls.

Number of species and distribution:— Urginavia includes 16 species from Southern, East and West Africa, and is restricted to the Cape, Karoo-Namib, Uzambara-Zululand and Guineo-Congolian Regions, and the Zambezian, Erithreo-Arabian and Sahelo-Sudanian Subregions (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a) ( Fig. 49 ). For further morphological species characterisation see Dyer (1942c, 1947), Jessop (1977), and Manning & Goldblatt (2018).

Karyology:—2n=20 ( De Wet 1957, Jones & Smith 1967, as Urginea altissima Baker ; Stedje & Nordal 1987; De Wet 1957, as Urginea multisetosa Baker ; Stedje & Nordal 1987, as D. exigua Stedje ); 2n=22 ( Oyewole 1975, as U. altissima ; Oyewole 1975, as Urginea gigantea (Jacq.) Oyewole ; Oyewole 1975, as Urginea viridula Baker ); 2n=32 ( De Wet 1957, as U. epigea R.A.Dyer ); 2n=60 ( Goldblatt et al. 2012, as D. capensis (Burm.f.) Wijnands ).

History, diagnostic characters, and taxonomic relationships:— Speta (1998b) described Urginavia to include plants with large bulbs; inflorescence commonly large, long racemose, multi-flowered; spurred bracts and distinct bracteoles; patent pedicels and stellate flowers; tepals only slightly connate at base; nocturnal flowers (sensu Speta 1998b; doubtful); and flattened black seeds. The new genus was based on morphological and phylogenetic studies ( Pfosser & Speta 1999, 2001, 2004) and Speta (1998b) included in it Urginavia altissima (Linnaeus f. 1782: pl. 199) Speta (1998b: 87), U. brachystachys ( Baker 1892: 474) Speta (1998b: 87) , U. porphyrantha ( Bullock 1932: 505) Speta (1998b: 87) , U. micrantha ( Richter 1850: 328) Speta (1998b: 87) (the type), and U. viridula ( Baker 1898: 538) Speta (1998b: 87) . Later Speta (2001) added the combination Urginavia epigea ( Dyer 1947: t. 1027) Speta (2001: 168). This genus includes a distinct group of relatively large species occurring in Africa, south of the Sahara Desert ( Speta 1998a). However, we transfer Urginea brachystachys to Sekanama , based on the red bulb and translucent withered tepals, among other characters, as discussed above.

Manning & Goldblatt (2018) accepted this group of species as D. sect. Urginavia ( Speta 1998b: 86) Manning & Goldblatt (2018: 13) mostly following Speta’s (1998b) concept, but adding important characters such as welldeveloped bracteoles, diurnal flowers, and tepals persisting below the developing capsule, not cohering into a cap above. Their section included three species for Southern Africa: Drimia altissima (Linnaeus f. 1782: pl. 199) Ker Gawler (1808: t. 1074), D. capensis ( Burman 1768: 10) Wijnands (1983: 130) , and D. kniphofioides ( Baker 1897: 469) Manning & Goldblatt (2003: 111) .

Manning & Goldblatt (2018) placed Urginea multisetosa Baker (1897: 468) and U. echinostachya Baker (1897: 468) (= Ornithogalum cooperi Baker 1873b: 284 ) in their polyphyletic D. sect. Ledebouriopsis , and described the absence of bracteoles in both species. Our phylogenetic results have confirmed that these two species belong to Urginavia , further sharing the diagnostic characters of that genus, which include having distinct bracteoles, evident in the type collections. The phylogenetic analyses of Martínez-Azorín et al. (2023a) include thirteen samples of Urginavia from Central and Southern Africa and form a perfectly supported clade recognised at the rank of genus. This distinction is morphologically substantiated by their bulb scales that are somewhat loose, heterogeneous in length and vertically imbricate, usually yellowish and leathery when dry, and showing distinct white silky threads when broken ( Fig. 65.2 ), an apomorphy of the genus. Urginavia is sister to a clade including samples of Ledurgia , Thuranthos , and Zingela , which mostly co-occur in distribution, but show very different morphology.

Pfosser & Speta (1999, 2001) included a sample of Urginavia micrantha ( Richter 1850: 328) Speta (1998b: 87) , the type of the genus, in their phylogenetic analyses involving only the trnL-F region. We were unable to obtain further sequences of other DNA regions for that same sample and accordingly excluded it from our broader analyses. However, the inclusion of the published sample of U. micrantha in our trnL-F matrix confirmed its placement in Urginavia . The taxonomy of the genus requires further resolution assisted by fieldwork across its extensive sub-Saharan range.

Accepted species and required new combinations:—

Urginavia altissima (L.f.) Speta in Phyton (Horn, Austria) 38(1): 87 (1998) ≡ Ornithogalum altissimum L.f., Suppl. Pl.: 199 (1782), basionym ≡ Drimia altissima (L.f.) Ker Gawl. in Curtis’s Bot. Mag. 27: t. 1074 (1808) ≡ Urginea altissima (L.f.) Baker in J. Linn. Soc., Bot. 13: 221 (1873) ≡ Idothea altissima (L.f.) Kuntze, Revis. Gen. Pl. 2: 712 (1891) ( Figs 5.9 , 63.2 , 65.1 ). Type:— SOUTH AFRICA. Eastern Cape, Cap. Bona Spei, Zondags River, without date, Thunberg s.n. (UPS-THUNB 8275, lecto. designated as “ Type ” by Jessop in J. S. African Bot. 43: 288. 1977).

= Ornithogalum giganteum Jacq., Hort. Schoenbr. 1: 45, t. 87 (1797) ( Fig. 63.2 ) ≡ Urginea gigantea (Jacq.) Oyewole in Bol. Soc. Brot., sér. 2, 49: 167 (1975). Type:—illustration in Jacq., Hort. Schoenbr. 1: t. 87 (1797) (lecto. designated as “ Type ” by Jessop in J. S. African Bot. 43: 288. 1977).

= Drimia uitenhagensis Eckl. in S. Afr. Quart. Journ. 1: 364 (1830). Type:— SOUTH AFRICA. Eastern Cape. Port Elizabeth (3325): on the hills of Adow [Addo], District Uitenhage , (–DA), Ecklon 17 (S10-14097! lecto. designated by Manning & Goldblatt 2018).

Urginavia barteri (Baker) Mart. - Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia barteri Baker in J. Linn. Soc., Bot. 11: 423 (1870), basionym. Type:— NIGERIA. Niger territory, in ditione nigritana prope Nupe, Barter 1183 (K000257327 holo!).

Urginavia capensis (Burm.f.) Mart. -Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Scilla capensis Burm.f., Prod. Fl. Cap. : 10 (1768), basionym ≡ Drimia capensis (Burm.f.) Wijnands , The Botany of the Commelins: 130 (1983) ( Figs 5.10 , 63.1, 63.3 ). Type:— illustration in Commelin, Hort. Amst. 2: 187 t. 94 (1701) ( Fig. 63.1 ) (lecto. designated by Wijnands in The Botany of the Commelins: 130. 1983).

= Urginea forsteri Baker, Fl. Cap. (Harvey) 6(3): 469 (1897) ≡ Drimia forsteri (Baker) Oberm. in Bothalia 13: 453 (1980). Type:— SOUTH AFRICA. Without locality or date, Forster s.n. (K! lecto. effectively designated by Obermeyer 1981b: 453).

Urginavia congesta (Bullock) Mart. -Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia congesta Bullock in Bull. Misc. Inform. Kew 1932(10): 504 (1932), basionym [non Drimia congesta (Wight) Stearn in Ann. Mus. Goulandris 4: 208 (1978)] ≡ Urginea congesta Wight, Icon. Pl. Ind. Orient. 6: 28, t. 2064 (left hand figure) ( Wight 1853)]. Type:— KENYA. Mt. Elgon, elev. 8700 ft., 06 January 1931, E.J. Major & C. Lugard 474 (K000257334! lecto. designated as “holo.” by Stedje in Fl. Trop. E. Africa, Hyacinth.: 17. 1996; EA isolecto.).

Urginavia cooperi (Baker) Mart. -Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Ornithogalum cooperi Baker in J. Linn. Soc., Bot. 13: 284 (1873), basionym [non Ornithogalum cooperi (Baker) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 547 (2004), nom. illeg.] ≡ Drimia cooperi (Baker) Baker, Fl. Cap. (Harvey) 6: 443 (1897), nom. illeg., [non D. cooperi Baker, Refug. Bot. [Saunders] 1: pl. 18 (1869) = Ledebouria concolor (Baker) Jessop in J. S. African Bot. 36: 254. 1970] ( Fig. 65.2 ). Type:— SOUTH AFRICA. Eastern Cape, ad oram orientalis, Barber s.n. (TCD lecto. designated by Jessop in J. S. African Bot. 43: 287. 1977; K000257356! isolecto.).

Urginavia echinostachya (Baker) Mart. -Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea echinostachya Baker, Fl. Cap. (Harvey) 6(3): 468 (1897), basionym ≡ Drimia echinostachya (Baker) Eggli & N.R.Crouch in Bothalia 42(1): 43 (2012) ( Fig. 5.11 ). Type:— SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): Inanda, (–DB), J.M. Wood 276 (K000257349-1! lecto. designated here; NH0004369! isolecto.). Note:—Both vouchers at K and NH are original material and therefore holotype indication by Crouch et al. (2010) is disregarded.

Urginavia epigea (R.A.Dyer) Speta in Stapfia 75: 168 (2001) ≡ Urginea epigea R.A.Dyer in Fl. Pl. Africa 26: t. 1027 (1947) ( Figs 64.1 , 65.3 ). Type:— SOUTH AFRICA. Mpumalanga, Transvaal, Middelburg Dist., N of Zoetvelden, Van der Merwe 2203 (PRE0048542- 1! lecto. designated by Martínez-Azorín et al. in Phytotaxa 201: 169. 2015: the inflorescence only).

Urginavia exigua (Stedje) Mart. - Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia exigua Stedje in Nordic J. Bot. 14(1): 43 (1994), basionym. Type:— ETHIOPIA. Guder River valley , Nordal 1015 (O holo.).

Urginavia kniphofioides (Baker) Mart. -Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea kniphofioides Baker, Fl. Cap. (Harvey) 6(3): 469 (1897), basionym ≡ Drimia kniphofioides (Baker) J.C.Manning & Goldblatt in Bothalia 33(1): 111 (2003) ( Figs 5.12 , 66.1 ). Type:— SOUTH AFRICA. Mpumalanga. Komatipoort (2531): Transvaal, Havelock Concession, (–CC), elev. 3200–5000 ft., September 1890, E. Saltmarshe sub Galpin 1055 (K000257361! lecto. designated here; BOL140330! isolecto.). Note:—Both materials at K and BOL are here considered to be original material and therefore a lectotype designation is needed.

Urginavia micrantha (A.Rich.) Speta in Phyton (Horn, Austria) 38(1): 87 (1998) ≡ Scilla micrantha A.Rich., Tent. Fl. Abyss. 2: 328 (1850) ≡ Urginea micrantha (A.Rich.) Solms, Schweinfurth, Beitr. Fl. Aeth. : 294 (1867) ( Fig. 5.13 ). Type:— ETHIOPIA. Abyssinia, Chiré, Quartin Dillon & Petit s.n. (P02157062! lecto. designated as “ holotype ” by Stedje & Thulin in Nordic J. Bot. 15: 598. 1995; P02157060!, WU081571! isolecto.).

= Drimia paolii Chiov. in Res. Sci. Somalia Ital. 1: 175 (1916), basionym. Type:— SOMALIA. El Ualàc, 3 November 1913, Paoli 1096 (FT000782! holo.).

Urginavia multisetosa (Baker) Mart. -Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea multisetosa Baker, Fl. Cap. (Harvey) 6(3): 468 (1897), basionym ≡ Drimia multisetosa (Baker) Jessop in J. S. African Bot. 43(4): 299 (1977) ( Figs 5.14 , 64.2 , 66.2 ). Type:— SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): Natal, near Mooi River, (–AA), elev. 4000−5000 ft., October 1894, J. Medley Wood 5724 (K000400572! lecto. designated by Jessop in J. S. African Bot. 43: 299. 1977).

Urginavia porphyrantha (Bullock) Speta in Phyton (Horn, Austria) 38(1): 87 (1998) ≡ Urginea porphyrantha Bullock in Bull. Misc. Inform. Kew 1932(10): 505 (1932) ≡ Drimia porphyrantha (Bullock) Stedje in Nordic J. Bot. 7(6): 665 (1987). Type:— KENYA. Mt. Elgon, elev. 6500 ft., 16 March 1931, E.J. Major & C. Lugard 556 (K000257338! lecto. designated as “ holotype ” by Stedje 1987: 665; EA isolecto.).

Urginavia psilostachya (Welw. ex Baker) Mart. -Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea psilostachya Welw. ex Baker in Trans. Linn. Soc. London, Bot. 1(5): 247 (1878), basionym ≡ Drimia psilostachya (Welw. ex Baker) J.C.Manning & Goldblatt in Bothalia 43(1): 77 (2013). Type:— ANGOLA. Cazengo, in dumetis siccis arenosis, Welwitsch 3807 (BM000911784! lecto. designated here; K000400575 isolecto.).

Urginavia riparia (Baker) Mart. -Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea riparia Baker, Fl. Cap. (Harvey) 6: 467 (1897), basionym. Type:— SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): Umzinyati Falls, (–DB), 11 September 1880, J. Medley Wood 1052 (K000099097! lecto. designated by Jessop in J. S. African Bot. 43: 303. 1977). Note:—The collection Wood 1052 is a mixed collection. On the one hand it includes a voucher (K000099097) fitting the protologue of U. riparia Baker , which was selected as lectotype of that name by Jessop (1977). On the other hand, the voucher NH0004370 corresponds to Drimia elata s.l. and is not to be regarded as original material since it does not match the protologue.

Urginavia simensis (Hochst. ex A.Rich.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Scilla simensis Hochst. ex A.Rich., Tent. Fl. Abyss. 2: 327 (1850), basionym ≡ Drimia simensis (Hochst. ex A.Rich.) Stedje in Nordic J. Bot. 15(6): 597 (1996) ≡ Charybdis simensis (Hochst. ex A.Rich.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998), nom. illeg. [Art. 52 of the ICN] ( Figs 5.15 , 66.3 ). Type:— ETHIOPIA. Ad latus meridionale montium simensium prope Abbu Mekanna et Debra Dschoa, 2 March 1840, W. Schimper 1317 (P02057842! lecto. designated by Martínez-Azorín et al. 2022: only the bulb on the right side of the sheet bearing a scape with flowers and fruits, and the bulb in the affixed envelope; K000257332!, K000257333!, L1459048!, BR0000090026478!, M0107240!, P02057845!, TUB006926! isolecto.).

Urginavia viridula (Baker) Speta in Phyton (Horn, Austria) 38(1): 87 (1998) ≡ Urginea viridula Baker, Fl. Trop. Afr. [Oliver et al.] 7(3): 538 (1898), basionym ≡ Drimia viridula (Baker) J.C.Manning & Goldblatt in Bothalia 43(1): 78 (2013). Type:—Lower Guinea. Congo, cultivated specimen, 10 September 1886, ex hort., Bull s.n. (K000257329! holo.).

Figures

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 63. Species of Urginavia Speta. 1. Urginavia capensis (Burm.f.) Mart.-Azorín et al. from Commelin (1701: t. 94, as Scilla africana); 2. Urginavia altissima (L.f.) Speta from Jacquin (1797: t. 87, as Ornithogalum giganteum Jacq.); 3. Urginavia capensis (Burm. f.) Mart.-Azorín et al. from Ker Gawler (1808: t. 1074, as Drimia altissima (L.f.) Ker Gawl.).

FIGURE 64. Species of Urginavia Speta. 1. Urginavia epigea (R.A.Dyer) Speta from Dyer (1947: t. 1027, as Urginea epigea R.A.Dyer); 2. Urginavia multisetosa (Baker) Mart.-Azorín et al. from Dyer (1942c: t. 854, as Urginea multisetosa Baker).

FIGURE 65. Species of Urginavia Speta displayed in horizontal rows of images. 1. Urginavia altissima (L.f.) Speta; 2. Urginavia cooperi (Baker) Mart.-Azorín et al.; 3. Urginavia epigea (R.A.Dyer) Speta.

FIGURE 66. Species of Urginavia Speta displayed in horizontal rows of images. 1. Urginavia kniphofioides (Baker) Mart.-Azorín et al. (Photographs by D. Oosthuizen); 2. Urginavia multisetosa (Baker) Mart.-Azorín et al.; 3. Urginavia simensis (Hochst. ex A.Rich.) Mart.- Azorín et al. (Photographs by B. Stedje).