Drimia Jacq. ex Willd., Sp. Pl.

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 : 35-41

5. Drimia Jacq. ex Willd., Sp. Pl. ed. 4, 2(1): 165 (1799) ( Figs 18–20 ).

Typus generis:— Drimia elata Jacq.

= Strepsiphyla Raf., Fl. Tellur. 3: 60 (1837). Holotype:— S. villosa (Lindl.) Raf. = Idothea Kunth, Enum. Pl. [Kunth] 4: 341 (1843). Lectotype:— I. elata (Jacq.) Kunth (designated by Stearn 1978: 203) ≡ Idothearia C.Presl in Abh. K̂nigl. B̂hm. Ges. Wiss., ser. 5, 3: 544 (1845), nom. nov. illeg. [see Crespo et al. in Pl. Syst. Evol. 306: [Art. 67] 9 (2020)].

Description:—Bulbous geophyte. Bulb hypogeal or partly epigeal, ovoid to globose, variable in size, bulb scales usually compact or sometimes loose. Roots thickened and branched. Leaves 1 to several per bulb, commonly hysteranthous, rarely synanthous, with variable shape and morphology, from glabrous to long cilated or hairy, sometimes undulate. Inflorescence racemose, usually an elongated and multiflowered, 5‒100 cm long raceme, rarely compact and subcorymbose; peduncle terete, erect, smooth and glabrous; pedicels subpatent and usually curved, to 20 mm long. Bracts narrowly lanceolate, lowermost with distinct oblong spur, sometimes the bract and spur connected to peduncle by whitish membranous portion; bracteoles absent. Flowers stellate, erect-patent, diurnal, lasting 1‒2 days. Tepals 6, 10‒20 mm long, white, yellow, green, brown or purple, fused along basal third to form cylindrical tube, free perigone segments strongly reflexed and touching basal tube, narrowly lanceolate-oblong, with slightly wider and spatulate apex, with darker longitudinal band mostly visible on abaxial side. Stamens 6, erect and connivent to style, sometimes curved upwards; filaments narrowly triangular to filiform, smooth, adnate to perigone tube and arising from its mouth, rarely very shortly connate above perigone to form short filaments tube; anthers dehiscing by longitudinal slits along their whole length, yellowish, greenish or purplish. Ovary ovoid, trigonous, pale green to yellowish, well differentiated from style. Style narrow and long (much longer than ovary), erect. Stigma trigonous, papillose. Capsule ovoid, 7‒13 mm long, green first and papyraceous after ripening, 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 developing capsules. Seeds subellipsoid to reniform, flattened with prominent central embryo and wide, flat wings, 5‒10 mm long, black to dark brown, with somewhat sinuous anticlinal testa cell walls.

Number of species and distribution:—The genus includes 19 described species distributed in Sub-Saharan Africa, with the center of diversty in the southwestern parts of South Africa and few species reaching East Africa and western Central Africa, restricted to the Cape, Karoo-Namib and Uzambara-Zululand Regions, and the Zambezian and Sahelo-Sudanian Subregions (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a) ( Fig. 8 ). We consider the taxonomy of Drimia s.str. poor due to the lack of comprehensive studies covering its large distribution range. Further studies are needed, including of fresh material from the type localities of the described taxa, together with detailed morphological studies to ascertain the real number of existing taxa. In accepting very wide species concepts, which cover a large morphological variability, the treatments by Jessop (1977), Stedje (1987), and Manning & Goldblatt (2018) drastically reduced the number of accepted taxa in Southern and East Africa. We tentatively accept all described species in the genus based on their morphological differentiations. For further information on Drimia species see Jessop (1977) and Manning & Goldblatt (2018).

Karyology:—2n=18 ( Bruyns & Vosa 1987, as D. haworthioides Baker ; De Wet 1957, Fernandes & Neves 1962, as D. alta R.A.Dyer ; Bruyns & Vosa 1987, as D. ciliaris Jacq. ex Willd. ); 2n=20 ( De Wet 1957, as D. haworthioides ).

History, diagnostic characters, and taxonomic relationships:—Jacquin (1794, 1797) described and illustrated seven species of Drimia (nom. inval.). Among them, Drimia purpurascens Jacquin (1812: 48) and Drimia revoluta (Linnaeus f. 1782: 204) Sweet (1818: 72) , both currently placed in Squilla and Ledebouria Roth (1821: 124) , respectively ( Crespo et al. 2020). The remaining species constitute a compact group characterised by tepals basally connate in a distinct tube, the free portion of tepals elongated and narrowly lanceolate-oblong to subspatulate and strongly reflexed, and stamens adnate to the perigone, usually slightly curved and connivent to the style. Willdenow (1799) validated Drimia based on Jacquin’s descriptions and illustrations. This genus is the oldest in Urgineoideae and presented a homogeneous flower morphology in their original concept, with the exclusion of the two species mentioned above. However, with the description of further species in Drimia ( Baker 1871, 1897, 1898, Jessop 1977, Stedje 1987, Manning & Goldblatt 2003, 2007, Van Jaarsveld & Van Wyk 2005 ), its morphological circumscription became manifestly blurred. Although also based on other characters, this primarily stems from differing perceptions of the significance of the extent of tepal connation, from nearly free to connate in a distinct tube. This divergence in opinion introduced massive instability in generic circumscriptions in the Urgineoideae ( Huber 1969, Jessop 1977, Deb & Dasgupta 1982, Stedje 1987, 2001a, 2001b, Manning et al. 2004).

The phylogenetic studies by Martínez-Azorín et al. (2023a) include 17 samples of Drimia constituting a strongly supported clade, which is sister to a clade combining Litanthus and Schizobasis samples, with moderate support. The sister genera Litanthus and Schizobasis strongly differ in morphology, as shown below, further supporting the acceptance of Drimia in a narrow sense.

Accepted species:—

Drimia alta R.A.Dyer in Fl. Pl. South Africa 23: t. 890 (1943) ( Fig. 19.2 ), nom. nov. ≡ Drimia altissima Hook.f. in Bot. Mag. 91: t. 5522 (1865) ( Fig. 19.1 ), nom. illeg. [non Drimia altissima (L.f.) Ker Gawl. in Bot. Mag. 27: t. 1074 (1808)]. Type:— SOUTH AFRICA. Natal [KwaZulu-Natal], from John Sanderson of D’Urban [Durban]’, sub illustr. W. Fitch s.n. (K000257364! holo.).

Drimia burchellii Baker in Refug. Bot. [Saunders] 2: App. 2. (1870) ≡ Idothea burchellii (Baker) Kuntze, Revis. Gen. Pl. 2: 712 (1891). Type:— SOUTH AFRICA. Cape [Eastern Cape], Uitenhage Div., between Galgebosch and Melkrivier, 17 February 1814, Burchell 4769 (K000857424! lecto. designated by Jessop in J. S. African Bot. 43: 285. 1977; TCD?isolecto. [as Burchell “4969”]).

Drimia ciliaris Jacq. ex Willd., Sp. Pl. 2: 165 (1799) ≡ Hyacinthus ciliaris (Jacq. ex Willd.) Poir. in Lam., Encycl., Suppl. 3: 120 (1813) ≡ Idothea ciliaris (Jacq. ex Willd.) Kunth, Enum. Pl. 4: 343 (1843). Type:—illustration in Jacq., Icon. Pl. Rar. 2 (16): t. 377 (1795) ( Fig. 18.1 ) (lecto. designated as “ type ” by Jessop in J. S. African Bot. 43: 284 1977).

Drimia concolor Baker in J. Linn. Soc., Bot. 11: 422 (1871) ≡ Idothea concolor (Baker) Kuntze, Revis. Gen. Pl. 2: 712 (1891). Type:— SOUTH AFRICA. Cap. B. Spei (e tab. ex exemplo in Hort. Kew. anno 1823 culto depicta descript) [ Manning & Goldblatt (2018) cited that the “ holotype ” was presumably at K, though we did not find there any material fitting the protologue].

Drimia eckloniana Schult.f., Syst. Veg. , ed. 15 bis [Roemer & Schultes] 7(2): 1710 (1830) = Drimia humilis Berg. ex Eckl., Topogr. Verz. Pflanzensamml. Ecklon : 2 (1827), nom. nud. ≡ Drimia humilis Berg. ex Kunth, Enum. Pl. 4: 344 (1843), nom. inval. [in syn.] ≡ Idothea humilis Kunth, Enum. Pl. 4: 344 (1843), nom. illeg. superfl. Type:— SOUTH AFRICA. In arenosis Prom. B. Spei., Ecklon s.n. Drimia No. 89 [Manning & Goldblatt in Strelitzia 40: 50 (2018) designated this collection as “holo.”, which was supposedly kept at B; however the collection is not extant at B (R. Vogt pers. comm.) and therefore, a new type is needed; Lecto. designated here:—HAL108222! lecto.].

Drimia elata Jacq. ex Willd., Sp. Pl. 2: 165 (1799) ≡ Hyacinthus elatus (Jacq. ex Willd.) Poir. in Lam, Encycl., Suppl. 3: 120 (1813) ≡ Idothea elata (Jacq. ex Willd.) Kunth, Enum. Pl. 4: 343 (1843) ( Figs 1.19 , 18.2 , 20.1 ). Type:—illustration in Jacq., Icon. Pl. Rar. 2 (15): t. 373 (1794) ( Fig. 18.2 ) (lecto. designated by Stedje & Thulin in Nordic J. Bot. 15: 597. 1995).

Drimia elgonica Bullock in Bull. Misc. Inform. Kew 1932(10): 504 (1932). Type:— KENYA. Mt. Elgon , elev. 6900 ft., 15 March 1931, E.J. Major & C. Lugard 563 (K000257336! holo.).

Drimia haworthioides Baker in Gard. Chron. (1875) 1: 366 (1875) ( Figs 1.20 , 20.2 ). Type:— SOUTH AFRICA. Eastern Cape. Graaff-Reinet (3224): near Graaff-Reinet , (–BC), Flowers in December 1874 and leaves in February 1875 [ex hort. Kew], Bolus (lecto. designated here: The illustration in the protologue, Fig. 70 “bulb of Drimia haworthioides ”). Epitype (designated here):—K000857416! epi.). Note :— There are two specimens at Kew mounted on the same voucher (K000857415! and K000857416!) including fragments of plants that apparently correspond to the type collection (Bolus 2040). However, those fragments were apparently gathered at different times from plants grown at Kew. The voucher K000857415 includes a bulb resembling the illustration in the protologue, considered to come from the “type plant” that was gathered 23 Dec 1879 and annotated by R. Brown. The voucher K000857416 was regarded as “ holotype ” (to be corrected to lectotype) of this name by Manning & Goldblatt (2018: 57), and includes two fragments (a bulb and a flowering scape) belonging to two different collections and they are associated to a date (“Dec[ember] 7/[18]76”) that does not correspond to that cited in the protologue. Therefore, the illustration in the protologue is regarded here as the lectotype of the concerned name, and the voucher including an inflorescence and a bulb is selected here as epitype.

Drimia hockii De Wild. in Repert. Spec. Nov. Regni Veg. 12: 294 (1913). Type:— DEMOCRATIC REPUBLIC OF THE CONGO. Ober-Katanga , Elisabethville, September 1911, Ad. Hock s.n. (BR880749! lecto. designated here).

Drimia incerta A.Chev. ex Hutch., Fl. W. Trop. Afr. [Hutchinson & Dalziel] 2: 351 (1936) Type:— IVORY COAST. Hte. Sassandra, Pays Toura, Mont Dourou, à 1000 m d’altitude, près Koualé, 27 May 1909, Aug. Chevalier 21733 (P02156156! lecto. designated as “ type ” by Hutchinson in Kew Bull. 1939: 245. 1939).

Drimia ledermannii K.Krause in Bot. Jahrb. Syst. 51(3-4): 444 (1914). Type:— CAMEROON. Nordkamerun , Zwischen Lubare und Bakari, in hügeliger Baumsteppe, um 900−1000 m ü. M., 19 June 1909, C. Ledermann 2264 (B 10 0168393! lecto. designated here).

Drimia media Jacq. ex Willd., Sp. Pl. 2: 166 (1799) ≡ Hyacinthus medius (Jacq. ex Willd.) Poir. in Lam., Encycl., Suppl. 3: 120 (1813) ≡ Idothea media (Jacq.) Kunth, Enum. Pl. 4: 342 (1843). Type:—illustration in Jacq., Icon. Pl. Rar. 2 (16): t. 375 (1795) ( Fig. 18.3 ) (lecto. designated by Jessop in J. S. African Bot. 43: 282. 1977).

= Drimia rigidifolia Baker in J. Linn. Soc., Bot. 11: 420 (1871) [non Urginea rigidifolia Baker in J. Bot. 16: 323 (1878)] ≡ Idothea rigidifolia (Baker) Kuntze, Revis. Gen. Pl. 2: 712 (1891). Type:— SOUTH AFRICA. Somerset [?], Bowker 225 (K000857419! lecto. designated by Jessop in J. S. African Bot. 43: 285. 1977).

Drimia neriniformis Baker, Fl. Cap. (Harvey) 6: 442 (1897) ( Figs 2.1 , 20.3 ). Type:— SOUTH AFRICA. Free State. Harrismith (2829): Van Reenen , (–AD), marshy ground, 5000−6000 ft., 15 November 1892, Wood 4794 (K000257350! lecto. designated by Jessop in J. S. African Bot. 43: 280. 1977; E00109542!, NH0006430-0!, MO2142490!, P, PRE, SAM0023179-0!, Z, isolecto.).

= Drimia capitata Baker, Fl. Cap. (Harvey) 6(3): 442 (1897) [non Ornithogalum capitatum Hook.f. in Bot. Mag. 89: t. 5388 (1863) ≡ Urginea capitata (Hook.f.) Baker, Fl. Cap. (Harvey) 6(3): 465 (1897) ≡ Fusifilum capitatum (Hook.f.) Speta in Phyton (Horn, Austria) 38(1): 69 (1998)]. Type:— SOUTH AFRICA. [Eastern Cape]. Umtata (3128): Bazeia [Baziya], (–CB), Baur 1160 (K000257353! lecto. designated by Jessop in J. S. African Bot. 43: 280. 1977).

Drimia pusilla Jacq. ex Willd., Sp. Pl. 2: 165 (1799) ≡ Hyacinthus pusillus (Jacq.) Poir. in Lam., Encycl., Suppl. 3: 120 (1813) ≡ Idothea pusilla (Jacq.) Kunth, Enum. Pl. 4: 344 (1843) [non Anthericum pusillum Jacq., Icon. Pl. Rar. 2(16): 18, t. 417 (1795) ≡ Fusifilum pusillum (Jacq.) Speta in Phyton (Horn, Austria) 38: 70 (1998)]. Type:— SOUTH AFRICA. Illustration in Jacq., Icon. Pl. Rar. 2(14): t. 374 (1794) ( Fig. 18.4 ) (lecto. designated by Jessop in J. S. African Bot. 43: 284. 1977).

Drimia robusta Baker in Refug. Bot. 3: t. 190 (1870) ≡ Idothea robusta (Baker) Kuntze, Revis. Gen. Pl. 2: 712 (1891) ( Figs 2.2 , 19.3 , 20.4 ). Type:— SOUTH AFRICA. Cape Colony, from Cooper, Hort Saunders 4404, 1869, Cooper s.n. (K000857417! holo.).

Drimia rudatisii Schltr. in Engl. Bot. Jahrb. 40: 89 (1907). Type:— SOUTH AFRICA. KwaZulu-Natal. Port Shepstone (3030): Dumisa, feuchten schattigen Orten bei Fairfield , (–AD), 24 August 1905, Rudatis 79 (B100167475! holo.).

Drimia sphaerocephala Baker, Fl. Cap. (Harvey) 6: 441 (1897) ( Fig. 20.5 ). Type:— SOUTH AFRICA. Mpumalanga. Komatipoort (2531): Concession Creek near Barberton , (–CC), 7 September 1890, Galpin 1020 (K000400570! lecto. designated by Jessop in J. S. African Bot. 43: 280. 1977; BOL140327!, SAM0023178! isolecto.).

Drimia villosa Lindl. in Edwards’s Bot. Reg. 16: t. 1346 (1830) ≡ Strepsiphyla villosa (Lindl.) Raf., Fl. Tellur. 3: 60 (1837) ≡ Idothea villosa (Lindl.) Kunth, Enum. Pl. 4: 343 (1843). Type:— SOUTH AFRICA. Cape of Good Hope, received by Mr Tate, illustration in Edwards’s Bot. Reg. 16: t. 1346 (1830) ( Fig. 19.4 ) (lecto. designated here).

Drimia zombensis Baker, Fl. Trop. Afr. [Oliver et al.] 7(3): 525 (1898). Type:— MALAWI. Nyasaland, Mount Zomba, elev. 3000–5000 ft, September 1859, Kirk s.n. (K000400584! holo.).

Figures

FIGURE 1. Seed morphology in Urgineoideae. 1. Aulostemon mzimvubuensis (van Jaarsv.) Mart.-Azorín et al.; 2. Austronea barkerae (Oberm.ex J.C.Manning & Goldblatt) Mart.-Azorín et al.; 3. Austronea grandiflora Mart.-Azorín et al.; 4. Austronea hispidoplicata Mart.- Azorín et al.; 5. Austronea linearis Mart.-Azorín et al.; 6. Austronea oblongifolia Mart.-Azorín et al.; 7. Austronea olifanta Mart.-Azorín et al.; 8. Austronea pinguis Mart.-Azorín et al.; 9. Boosia flagellaris (T.J.Edwards et al.) Mart.-Azorín et al.; 10. Boosia macrocentra (Baker) Speta; 11. Boosia modesta (Baker) Mart.-Azorín et al.; 12. Boosia natalensis (Baker) Mart.-Azorín et al.; 13. Boosia nyasae (Baker) Mart.-Azorín et al.; 14. Boosia rubella (Baker) Mart.-Azorín et al.; 15. Boosia saniensis (Hilliard & B.L.Burtt) Mart.-Azorín et al.; 16. Boosia tenella (Baker) Mart.-Azorín et al.; 17. Bowiea gariepensis van Jaarsv.; 18. Bowiea volubilis Harv. ex T.Moore & Mast.; 19. Drimia elata Jacq. ex Willd.; 20. Drimia haworthioides Baker. All to the same scale in mm.

FIGURE 2. Seed morphology in Urgineoideae. 1. Drimia neriniformis Baker; 2. Drimia robusta Baker; 3. Fusifilum capitatum (Hook.) Speta; 4. Fusifilum montanum (A.P.Dold & E.Brink) A.P.Dold et al.; 5. Fusifilum physodes (Jacq.) Speta; 6. Geschollia anomala (Baker) Speta; 7. Geschollia brachyandra Mart.-Azorín et al.; 8. Geschollia calcarata (Baker) Mart.-Azorín et al.; 9. Geschollia globuligera Mart.- Azorín et al.; 10. Geschollia occultans (G.Will.) Mart.-Azorín et al.; 11. Geschollia prolifera Mart.-Azorín et al.; 12. Indurgia congesta (Wight) Speta; 13. Indurgia indica (Roxb.) Speta; 14. Iosanthus amboensis (Baker) Mart.-Azorín et al.; 15. Iosanthus macrostigma Mart.-Azorín et al.; 16. Iosanthus toxicarius (C.Archer & R.H.Archer) Mart.-Azorín et al.; 17. Litanthus pusillus Harv.; 18. Litanthus stenocarpus (J.C.Manning & J.M.J.Deacon) Mart.-Azorín et al.; 19. Mucinaea nana (Snijman) M.Pinter et al.; 20. Rhadamanthopsis haworthioides (Baker) Mart.-Azorín et al. All to the same scale in mm.

FIGURE 8. Approximate distribution ranges of accepted genera in Urgineoideae.

FIGURE 18. Species of Drimia Jacq. ex Willd. 1. Drimia ciliaris Jacq. ex Willd. from Jacquin (1795: tab. 377); 2. Drimia elata Jacq. ex Willd. from Jacquin (1794: tab. 373); 3. Drimia media Jacq. ex Willd. from Jacquin (1795: tab. 375); 4. Drimia pusilla Jacq. ex Willd. from Jacquin (1794: tab. 374).

FIGURE 19. Species of Drimia Jacq. ex Willd. 1. Drimia alta R.A.Dyer from Hooker (1865: t. 5522 as Drimia altissima Hook.f.); 2. Drimia alta R.A.Dyer from Dyer (1943: t. 890); 3. Drimia robusta Baker from Baker (1870b: t. 190); 4. Drimia villosa Lindl. from Lindley (1830: t. 1346).

FIGURE 20. Species of Drimia Jacq. ex Willd. displayed in horizontal rows of images. 1. Drimia elata Jacq. ex Willd.; 2. Drimia haworthioides Baker; 3. Drimia neriniformis Baker; 4. Drimia robusta Baker; 5. Drimia sphaerocephala Baker.

FIGURE 70. Species of Urgineopsis Compton displayed in horizontal rows of images. 1. Urgineopsis barbata (J.C.Manning & J.M.J.Deacon) Mart.-Azorín et al.; 2. Urgineopsis salteri Compton.