Squilla Steinh., 1836

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 : 90-96

21. Squilla Steinh.

in Ann. Sci. Nat. Bot. sér. 2, 6: 276 (1836),

Typus generis:— S. maritima (L.) Steinh. (lectotype, vide Pfeiffer, Nom. 2(2): 1252. 1874).

≡ Charybdis Speta in Phyton (Horn, Austria) 38(1): 58 (1998), nom. nov. et nom. illeg. superfl. [Art. 52 of the ICN] ( Figs 52–54 ). Typus generis:— C. maritima (L.) Speta (holotype).

Description:—Usually tall bulbous geophyte. Bulb usually large and hypogeal, rarely somewhat epigeal, ovoid to globose, up to 20 cm in diam. Roots thickened and branched. Leaves (4–)6‒10 per bulb, narrowly lanceolate to widely ovate, hysteranthous, up to 35 cm long, margin entire or sometimes undulate with ciliae or minute papillae, green or somewhat glaucous, smooth, glabrous. Inflorescence a long, erect, multiflowered raceme, with up to 1000 flowers, 6‒70 cm long; peduncle 15‒120 cm long, dull green with grey to purple tinge, terete, erect, smooth and glabrous; pedicels 5‒35 mm long, subpatent and usually arching upwards, sometimes erect or patent at anthesis. Bracts narrowly lanceolate, usually dry and wrinkled at flowering time, lowermost with short spur; bracteoles present and distinct. Flowers stellate or with reflexed lobes, erect-patent, diurnal, lasting more than one day. Tepals 6, biseriate, 7‒16 mm long, free or connate for less than 1‒2 mm, lanceolate-ovate to spathulate, usually white or rarely purple, with purplish or green longitudinal band. Stamens 6, spreading; filaments smooth, usually white and lanceolate to narrowly triangular, flattened or rarely filiform and purplish; anthers dorsifixed. Ovary ovate-oblong, usually with three apical protuberances, green to yellow, with ca. 5‒10 ovules per locule. Style 3‒12 mm long, erect, white or rarely tinged with purple, with trigonous stigma. Capsule ovate, 7‒19 mm long, green first and yellowish 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 the developing capsules. Seeds ovate-oblong to linear-oblong, 4‒12 mm long, flattened, testa black, with sinuous anticlinal cell walls.

Number of species and distribution:— Squilla includes twelve species, occurring mostly along the Mediterranean Basin, spanning the Canary Islands and the Middle East ( Speta 1998a, Martínez-Azorín et al. 2022) ( Fig. 49 ); it is nearly restricted to the Mediterranean and Macaronesian Regions (sensu Takhtajan 1986). For further information on Squilla see Pfosser & Speta (2001, 2004), Crespo et al. (2020), and Martínez-Azorín et al. (2022).

Karyology:—2n=20 ( Martinoli 1954, Battaglia 1957, Bartolo et al. 1984, as Urginea undulata (Desf.) Steinh. ; Valdés-Bermejo 1980, Battaglia 1957, as D. maura Maire ); 2n=28 ( Bramwell et al. 1972, as Urginea hesperia Webb. & Berth. ); 2n=40 ( Satô 1942, as Urginea scilla Steinh. ); 2n=20, 30, 40, 50, 60 ( Battaglia 1957, 1964, Valdés-Bermejo 1980, Speta 1980, Pfosser & Speta 2001, Rico 2013, Bacchetta et al. 2012); 2n=20, 40, 60, 60+1B, 60+2B ( Boscaiu et al. 2001, Fernández et al. 2001, Talavera et al. 1995, Oberprieler & Vogt 1994, as Urginea maritima ).

History, diagnostic characters, and taxonomic relationships:— Martínez-Azorín & Crespo (2016a) showed that the Linnaean Scilla was first typified on Scilla maritima Linnaeus (1753: 308) by Rafinesque (1837: 8) and not on Scilla bifolia Linnaeus (1753: 309) as usually accepted [vide Index Nominum Genericorum: Farr & Zijlstra (2023) http://botany.si.edu/ing/].

Steinheil (1836) described Squilla to segregate Scilla maritima Linnaeus (1753: 308) ( Urginea scilla Steinheil 1834: 321 ) from his previously described Urginea , implying Squilla to be different from the name Scilla . Speta (1998b) accepted separation of both Urginea Steinh. and Squilla , but proposed the new name Charybdis , with C. maritima ( Linnaeus 1753: 38) Speta (1998b: 60) as type, to replace Squilla since he considered the latter to be an orthographic variant of both Scilla and Skilla Rafinesque (1837: 13) , and therefore a later illegitimate homonym. However, based on the typification by Rafinesque (1837) of Scilla maritima , Charybdis is illegitimate under Art. 52 of the Shenzhen Code as it was superfluous when published, a fact that makes Charybdis unavailable for use.

Martínez-Azorín & Crespo (2016b) requested the Nomenclature Committee for Vascular Plants (NCVP) for a binding decision on whether Scilla L. (subfam. Hyacinthoideae ) and Squilla Steinh. (subfam. Urgineoideae ≡ Asparagaceae subfam. Scilloideae trib. Urgineeae ) are sufficiently alike to be considered orthographic variants and to be confused. After several attempts, a sufficient number of the Committee Members voted recently in favour of a binding decision that both names are not confusable (cf. Appelquist 2022), which deed allows Squilla to be used in the sense of the current concept of Charybdis ( Martínez-Azorín et al. 2022) .

Previous phylogenetic analyses ( Pfosser & Speta 2001, 2004, Manning et al. 2004, Pfosser et al. 2012) show samples of Squilla (as Charybdis ) forming a strongly supported clade. Many authors recently accepted Charybdis as distinct from Urginea and Drimia at generic rank (cf. Speta 1998b, Pfosser & Speta 2001, 2004, Conti et al. 2005, Jeanmonod & Gamisans 2007, Bacchetta et al. 2012, Ali et al. 2013, Véla et al. 2016), although others (e.g. Manning et al. 2004, Manning & Goldblatt 2018, WCSP 2021) place them in an expanded and broadly circumscribed Drimia . The phylogenetic analyses of Martínez-Azorín et al. (2023a) included 22 samples of Squilla , which formed a strongly supported clade in an isolated position within Urgineoideae . Therefore, based on the distinct morphology and genetic divergence of these plants, we here accept Squilla following Martínez-Azorín et al. (2022), which is mostly restricted to the Mediterranean Basin.

Accepted species:—

Squilla anthericoides (Poir.) Jord. & Fourr., Icon. Fl. Eur. 2: 2 (1868) ≡ Scilla anthericoides Poir., Voy. Barbarie 2: 149 (1789), basionym ≡ Ornithogalum anthericoides (Poir.) Link ex Steud., Nomencl. Bot. 1: 573 (1821) ≡ Urginea anthericoides (Poir.) Steinh. in Ann. Sci. Nat., Bot. sér. 2, 1: 328 (1834) ≡ Charybdis anthericoides (Poir.) Véla & Bélair in Phytotaxa 288: 155 (2016) ≡ Drimia anthericoides (Poir.) Véla & Bélair in Willdenowia 49: 423 (2019) ≡ Urginea maritima var. anthericoides (Poir.) Maire & Weiller, Fl. Afrique N. 5: 163 (1958). Type:—[ ALGERIA.] Ex Numidia, herb. Poiret (P02157340! lecto. designated by Véla et al. in Phytotaxa 288: 155. 2016: the specimen on the upper right side of the sheet).

Squilla aphylla (Forssk.) Mart. -Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 3 (2022) ≡ Anthericum aphyllum Forssk., Fl. Aegypt.-Arab.: 209 (1775), basionym ≡ Urginea aphylla (Forssk.) Speta in Linzer Biol. Beitr. 12(1): 229 (1980) ≡ Charybdis aphylla (Forssk.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia aphylla (Forssk.) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004). Type:—[ GREECE]. “ Anthericum ” F. Herb. Forskålii n° 10 [IDC microfiche foto: Forsskål nr. 129 III, 1–2] (C10001668! lecto. designated by Martínez-Azorín et al. 2022).

= Ornithogalum elatum Andrews in Bot. Repos. 8, pl. DXXVIII (1808) ≡ Charybdis elata (Andrews) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia excelsa J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004), replac. name [non Drimia elata Jacq., Collectanea 5(Suppl.): 38. 1797]. Type:—illustration in Andrews (1808), pl. DXXVIII (lecto. designated by Martínez-Azorín et al. 2022: only the flowering scape and the flower dissection).

Squilla glaucophylla (Bacch., Brullo, D’Emerico, Pontec. & Salmeri) Mart. -Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43- e78272: 4 (2022) ≡ Charybdis glaucophylla Bacch., Brullo, D’Emerico, Pontec. & Salmeri in Phytotaxa 69: 18 (2012), basionym ≡ Drimia glaucophylla (Bacch., Brullo, D’Emerico, Pontec. & Salmeri) Raus in Willdenowia 46(3): 424 (2016). Type:— ITALY. Sardinia: Isola di San Pietro: Cala Vinagra, Carloforte, elev. 63 m, 19 July 2004, G. Bacchetta & C. Pontecorvo s.n. (CAT, holo.; CAG, CAT, iso.).

Squilla hesperia (Webb & Berthel.) Mart. -Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 4 (2022) ≡ Urginea hesperia Webb & Berthel., Hist. Nat. Iles Canaries (Phytogr., sect. 3) 3(2): 339 (1848), basionym ≡ Charybdis hesperia (Webb & Berthel.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia hesperia (Webb & Berthel.) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004) ≡ U. maritima var. hesperia (Webb & Berthel.) Svent. in Index Sem. Acclim. Pl. Arautapae 1969: 41 (1970) ≡ Drimia maritima var. hesperia (Webb & Berthel.) A.Hansen & Sunding, Fl. Macaronesia , Checklist Vasc. Pl., rev. ed. 2, 2: 4 (1979) ( Figs 4.10 , 53.1 ). Type:— SPAIN. Canary Islands. Urginea hesperia Nobis “3”, Herbarium Webbianum nº 182548 (FI000284!, lecto. designated by Martínez-Azorín et al. 2022).

Squilla maritima (L.) Steinh.in Ann. Sci. Nat., Bot. sér. 2, 6: 279 (1836) ≡ Scilla maritima L., Sp. Pl.: 308 (1753), basionym ≡ Ornithogalum maritimum (L.) Lam., Fl. Franç. 3: 276 (1779) ≡ Urginea maritima (L.) Baker in J. Linn. Soc., Bot. 13: 221 (1872) ≡ Drimia maritima (L.) Stearn in Ann. Mus. Goulandris 4: 204 (1978) ≡ Charybdis maritima (L.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Stellaris scilla Moench, Methodus : 304 (1794), nom. illeg. [Art. 52 of the ICN] ≡ Ornithogalum squilla Ker Gawl. in Bot. Mag. 23, t. 918 (1806), nom. illeg. [Art. 52 of the ICN] ≡ U. scilla Steinh. in Ann. Sci. Nat., Bot. sér. 2, 1: 331 (1834), nom. illeg. [Art. 52 of the ICN] ( Figs 4.11 , 52.1, 52.2, 52.3 , 53.2 ). Type:—illustration in Clusius (1601: 171, Fig. 52.1 ), “Scillae hispanicae flos & semen” (lecto. designated by Martínez-Azorín et al. 2022 b: only the illustration of the bulb with a flowering scape). Epitype (designated by Martínez-Azorín et al. 2022):— PORTUGAL. Sierra de Monsanto, environs de Lisbonne, pentes pierreuses, July−August 1877, J. Daveau (P02166423! epi.; P02166260!, P02166417!, P01811235!, P01811236!, isoepi.).

Comments on typification:—Recently, Martínez-Azorín et al. (2022) considered that the lectotype of Scilla maritima , first designated by Ferrer-Gallego (2013: 40) on the voucher L0052807, should be disregarded according to Art. 8.2 of the ICN, because completely developed adult leaves in this species are never coetaneous with flowers at anthesis in a given population. This implies that the “ lectotype ” selected by Ferrer-Gallego (2013) includes two specimens collected at different times: i) leaves, and ii) the flowering scape. Considering Ferrer-Gallegos’s (2022) reaction to our lectotypification, and in the light of the recentmost proposal by Mosyakin & McNeill (2022) to amend Art. 8, after which the information in the protologue will rule, we still consider that the voucher L0052807 cannot be a proper lectotype, following Linnaeus’s statement in Hortus Cliffortianus (1738: 123) that is cited in the Scilla maritima protologue of 1753: “ Scilla radice tunicata [….] Singulare est quod foliis destitute floreat.” Further, Clusius (1601: 171), which is also cited in the protologue of S. maritima , illustrated the species through both a bulb with fully-developed leaves (winterspring facies) and two bulbs with flowering and fruiting scapes; those latter figures are completely lacking leaves (later summerearly autumn facies); in addition, Clusius (1601) describes the typical hysteranthous behaviour of the species in Hispania as follows: “Copiose nascitur supra Olysipponem & plerisque aliis Lusitaniae & Hispaniae locis. Floret Augusto & Septembri, semen Octobri & Novembri maturescit; folia maturo jam semine, & exarido caule, emergunt Novembri & Decembri.” Based on those facts, we do believe that Linnaeus never saw plants of Scilla maritima showing flowers and leaves coetaneously, as he indirectly claimed in his Hortus Cliffortianus ( Linnaeus 1738), and hence the Linnaean voucher L0052807 most likely represents a “collage” of mixed gatherings of flowers and leaves collected at different times, and it as a whole cannot be eligible as lectotype. Therefore, the lectotype and epitype we designated ( Martínez-Azorín et al. 2022) are considered here as appropriate to preserve the currently most widely accepted concept of this species.

= Scilla rubra Garsault, Descr. Pl. Anim. 4: 314, pl. 527 (1767). Type:—illustration in Garsault (l.c.), pl. 527 “ Scilla rubra ” (lecto. designated by Martínez-Azorín et al. 2022: only the illustrations of the bulb with a flowering scape, and the flowers).

Squilla maura (Maire) Mart. -Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 7 (2022) ≡ Urginea maura Maire in Bull. Soc. Hist. Nat. Afrique N. 14: 158 (1923), basionym ≡ Charybdis maura (Maire) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia maura (Maire) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004) ≡ Urginea maritima var. maura (Maire) Maire, Fl. Afrique N. 5: 165 (1958) ≡ Drimia maritima subsp. maura (Maire) F̂rther & Podlech in Sendtnera 7: 86 (2001). Type:— MOROCCO. Grand Atlas: Mentaga, rocailles calcaires, elev. 200–1000 m, feuilles glauques, 14 April 1922, R. Maire (MPU000488! lecto. designated by Martínez-Azorín et al. 2022).

= Urginea maritima subsp. maura var. angustifolia Maire in Bull. Soc. Hist. Nat. Afrique N. 29: 454 (1938) ≡ Urginea maritima var. maura f. angustifolia (Maire) Maire, Fl. Afrique N. 5: 165 (1958), basionym. Type:— MOROCCO. Anti Atlas : près de Kerdous, pentes rocheuses (quartzites), elev. 800 m, 38 March 1937, R. Maire (MPU003971! lecto. designated by Martínez-Azorín et al. 2022).

Squilla numidica Jord. & Fourr., Icon. Fl. Eur. 2: 1 (1868) ≡ Urginea numidica (Jord. & Fourr.) Grey, Hardy Bulbs 2: 632 (1938) ≡ Urginea maritima var. numidica (Jord. & Fourr.) Baker in J. Linn. Soc., Bot. 13: 221 (1873) ≡ Charybdis numidica (Jord. & Fourr.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia numidica (Jord. & Fourr.) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004). Type:—[ ALGERIA.] “ Squilla maritima numidica / de Bône (Algerie)”, 6 Novembre 1867 (LY0073734 [digital image!] lecto. designated by Martínez-Azorín et al. 2022).

= Squilla sphaeroidea Jord. & Fourr., Icon. Fl. Eur. 2: 1 (1868) ≡ Urginea maritima var. sphaeroidea (Jord. & Fourr.) Baker in J. Linn. Soc., Bot. 13: 221 (1873) ≡ U. sphaeroidea (Jord. & Fourr.) Grey, Hardy Bulbs 2: 632 (1938). Type:—illustration in Jordan & Fourreau (1868): Tab. CCII “278. Squilla sphaeroidea ” [lecto. designated by Martínez-Azorín et al. 2022: only the illustrations of the flowering scape, flowers and bracts ( Figs 1−12 ); excluding bulb, leaves, fruits and seeds ( Figs 13−16 )].

Squilla palaestina (M.B.Crespo, Mart.-Azorín & M.Á.Alonso) Mart.- Azorín, M.B.Crespo & M.Á. Alonso in Medit. Bot. 43-e78272: 8 (2022) ≡ Drimia palaestina M.B.Crespo, Mart. - Azorín & M.Á. Alonso in Pl. Syst. Evol. 306: 13 (2020), basionym ( Figs 4.12 , 53.3 ). Type:— PALESTINE. [Palestinian Authority], Judean Mts. South of Yata, Submediterranean batha, 23 September 1970, Y. Plitmann & M. Raviv 26089 (HUJ132724! holo.) .

Squilla pancration Steinh. in Ann. Sci. Nat., Bot. sér. 2, 6: 279 (1836) ≡ Scilla pancration (Steinh.) Nyman, Syll. Fl. Eur. : 369 (1855) ≡ Urginea pancration (Steinh.) Schroff in Z. Allg. Oesterr. Apotheker-Vereines 3 (Oesterr. Z. Pharmazie 13): 42 (1865) ≡ Charybdis pancration (Steinh.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia pancration (Steinh.) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004) ≡ Urginea maritima var. pancration (Steinh.) Baker in J. Linn. Soc., Bot. 13: 221 (1873) ≡ U. maritima subsp. pancration (Steinh.) K. Richter, Pl. Eur. 1: 218 (1890) ( Figs 4.13 , 54.1 ). Type:— MALTA. Gozo island (Għawdex), pr. Ir-Ramla, between San Blas and S. Philip, 33SVV359908, elev. 70 m, flowered in cultivation at the University of Alicante on 04 September 2013, J.L. Villar & E. Martínez s.n. (ABH70437! neo. designated by Martínez-Azorín et al. 2022).

= Squilla insularis Jord. & Fourr., Icon. Fl. Eur. 2: 2 (1868) ≡ Urginea maritima var. insularis (Jord. & Fourr.) Baker in J. Linn. Soc., Bot. 13: 221 (1873) ≡ U. maritima subsp. insularis (Jord. & Fourr.) K. Richt., Pl. Eur. 1: 218 (1890) ≡ U. scilla var. insularis (Jord. & Fourr.) Rouy, Fl. France 12: 425 (1910) ≡ U. insularis (Jord. & Fourr.) Grey, Hardy Bulbs 2: 632 (1938). Type:—illustration in Jordan & Fourreau (1868): Tab. CCIII “279. Squilla insularis ” [lecto. designated by Martínez-Azorín et al. 2022: only the illustrations of the flowering scape, flowers and bracts ( Figs 1−12 ); excluding bulb, leaves, fruits and seeds ( Figs 13−16 )].

= Squilla littoralis Jord. & Fourr., Icon. Fl. Eur. 2: 2 (1868) ≡ Urginea maritima var. littoralis (Jord. & Fourr.) Baker in J. Linn. Soc., Bot. 13: 221–222 (1873) ≡ U. maritima subsp. littoralis (Jord. & Fourr.) K.Richt., Pl. Eur. 1: 218 (1890) ≡ U. littoralis (Jord. & Fourr.) Grey, Hardy Bulbs 2: 632 (1938). Type:—illustration in Jordan & Fourreau (1868): Tab. CCIV “280. Squilla littoralis ” [lecto. designated by Martínez-Azorín et al. 2022: only the illustrations of the flowering scape, flowers and bracts ( Figs 1−12 ); excluding bulb, leaves, fruits and seeds ( Figs 13−16 )].

Squilla secundiflora (Maire) Mart. -Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 9 (2022) ≡ Urginea anthericoides var. secundiflora Maire in Bull. Soc. Hist. Nat. Afrique N. 24: 229 (1933), basionym ≡ Drimia secundiflora (Maire) M.B.Crespo, Mart. -Azorín & M.Á.Alonso in Pl. Syst. Evol. 306: 15 (2020) ( Fig. 4.14 ). Type:— MOROCCO. Forêt de la Mamora, bulbes cultivés à Alger ayant fleuri en septembre 1932, 28 October 1932, R. Maire s.n. (MPU003124! lecto designated by Crespo et al. 2020: 15 [as “ U. anthericoides var. secunda Maire ”]; only the flowering scape).

Squilla serotina (Schousb.) Mart. -Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 9 (2022) ≡ Scilla serotina Schousb., Iagttag. Vextrig. Marokko: 165 (1800), basionym ≡ Urginea anthericoides var. serotina (Schousb.) Font Quer in Bol. Real Soc. Esp. Hist. Nat. 27: 45 (1927) ( Figs 4.15 , 54.2 ). Type:—[ MOROCCO.] Herb. Liebm[ann]. Marocco, Schousboe (C10000939! lecto. designated by Crespo et al. 2020: 11, [IDC microfiche foto: Type Herbarium nr. 183 I,4]; C10000940 [digital image!] isolecto. [IDC microfiche foto: Type Herbarium nr. 183 I,5]).

= Urginea undulata var. tazensis Batt. & Maire in Bull. Soc. Hist. Nat. Afrique N. 22: 318 (1931) ≡ Urginea tazensis (Batt. & Maire) Maire in Bull. Soc. Hist. Nat. Afrique N. 31: 43 (1940) ≡ Urginea undulata subsp. tazensis (Batt. & Maire) Maire & Weiller, Fl. Afrique N. 5: 158 (1958) ≡ Drimia tazensis (Maire) Stearn in Ann. Mus. Goulandris 4: 208 (1978) ≡ Charybdis tazensis (Batt. & Maire) Speta in Stapfia 75: 167 (2001). Type:— MOROCCO. Tiznit, bulbs cultivated in Alger (Algeria), 18 September 1939, R. Maire (MPU004613! neo. designated by Crespo et al. 2020: 11).

= Urginea undulata var. major Gatt. & Weiller in Bull. Soc. Hist. Nat.Afrique N. 28: 539 (1937). Type:— MOROCCO. Tiznit , sandy soils, cultivated in Alger (Algeria), 8 September 1936, R. Maire (MPU001417! neo. designated by Crespo et al. 2020: 11).

Squilla undulata (Desf.) Mart. -Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 10 (2022) ≡ Scilla undulata Desf. in Méd. Eclairée Sci. Phys. 3(6): 161 (1792), basionym ≡ Urginea undulata (Desf.) Steinh. in Ann. Sci. Nat., Bot. sér. 2, 1: 330 (1834) ≡

Epimenidion undulatum (Desf.) Raf., Fl. Tellur. 2: 13 (1837) ≡ Prospero undulatum (Desf.) Salisb., Gen. Pl. : 28 (1866) ≡ Drimia undata Stearn in Ann. Mus. Goulandris 4: 208 (1978), replac. name [non Drimia undulata Jacq. in Willd., Sp. Pl. ed. 4, 2(1): 166. (1799), nec Jacq., Collectanea 5 (Suppl.): 41. 1797, nom. inval.] ≡ Charybdis undulata (Desf.) Speta in Stapfia 75: 167 (2001), nom. illeg. [Art. 52 of the ICN] ( Figs 4.16 , 52.5 , 54.3 ). Type:— ALGERIA. Herbier de la Flore Atlantique, no. 707 Scilla undulata [Tunis et Alger, collines arides, elle fleurit en hiver, 1787], Desfontaines (P00320347! lecto. designated by Crespo et al. 2019: 91: the flowering scape with bulb on the left side of the sheet; MPU017529! isolecto.).

= Urginea undulata f. caeculi Pau in Butl. Inst. Catalana His. Nat. 16: 136 (1916), basionym ≡ Urginea undulata var. caeculi (Pau) Pau in Bol. Soc. Ibér. Ci. Nat. 23: 91 (1924) ≡ Drimia undata subsp. caeculi (Pau) Mateo & M.B.Crespo, Fl. Abrev. Comun. Valenciana : 430 (1995) ≡ Urginea undulata subsp. caeculi (Pau) M.B.Crespo & Mateo in Flora Montiber. 5: 58 (1997). Type:— SPAIN. Valencia prov., Alginet, cultivada, August 1916, F. Moroder (MA21517! lecto. designated by Crespo & Mateo 1997: the scape on the right side of the sheet, from a bulb with incipient leaves).

= Drimia purpurascens J.Jacq., Ecl. Pl. Rar. 1(3–4): 48, t. 30 (1812) ≡ Idothea purpurascens (J.Jacq.) Kunth, Enum. Pl. 4: 342 (1843) ≡ Idothearia purpurascens (J.Jacq.) C.Presl in Abh. K̂nigl. B̂hm. Ges. Wiss. ser. 5, 3: 544 (1845), nom. illeg. superfl. [Art. 52 of the ICN]. Type:—illustration in J.Jacq., Ecl. Pl. Rar. 1(3–4), t. 30. 1812 ( Fig. 52.4 ) (lecto. designated by Crespo et al. 2020: only the flowering scape with bulb). Epitype (designated by Crespo et al. 2020: 9):— ITALY. Sardaigne, 1838, E. Thomas (P02166250! epi.: the fragment on the left side of the sheet).

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 3. Seed morphology in Urgineoideae. 1. Rhadamanthopsis hyacinthoides (Baker) Mart.-Azorín et al.; 2. Rhadamanthopsis karooicus (Baker) Mart.-Azorín et al.; 3. Rhadamanthopsis monophyllus (Oberm. ex J.C.Manning & Goldblatt) Mart.-Azorín et al.; 4. Rhadamanthopsis namibensis (Oberm.) Mart.-Azorín et al.; 5. Rhadamanthus arenicola B.Nord.; 6. Rhadamanthus ciliatus (L.f.) Mart.-Azorín et al.; 7. Rhadamanthus cochlearis (Mart.-Azorín et al.) Mart.-Azorín et al.; 8. Rhadamanthus convallarioides (L.f.) Salisb. ex Baker; 9. Rhadamanthus rigidifolius (Baker) Mart.-Azorín et al.; 10. Rhadamanthus urantherus R.A.Dyer; 11. Rhodocodon campanulatus Knirsch et al.; 12. Rhodocodon cryptopodus (Baker) Knirsch et al.; 13. Rhodocodon cyathiformis H.Perrier ex Knirsch et al.; 14. Rhodocodon floribundus H.Perrier ex Knirsch et al.; 15. Rhodocodon graciliscapus Knirsch et al.; 16. Rhodocodon intermedius H.Perrier ex Knirsch et al.; 17. Rhodocodon jackyi Knirsch et al.; 18. Rhodocodon mascarenensis (Baker) Knirsch et al.; 19. Rhodocodon monophyllus Knirsch et al.; 20. Rhodocodon perrieri Knirsch et al. All to the same scale in mm.

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 6. Seed morphology in Urgineoideae. 1. Urgineopsis salteri Compton; 2. Vera-duthiea capillifolia A.Patzelt et al.; 3. Veraduthiea sp. (Urginea petrophila A.Chev., nom. nud.); 4. Vera-duthiea sebiri (Berhaut) Mart.-Azorín et al.; 5. Vera-duthiea sp. (Urginea segetalis A.Chev., nom. nud.); 6. Zingela pooleyorum N.R.Crouch et al.; 7. Zulusia delagoensis (Baker) Mart.-Azorín et al.; 8. Zulusia edwardsii (N.R.Crouch & Mart.-Azorín) Mart.-Azorín et al.; 9. Zulusia lydenburgensis (R.A.Dyer) Mart.-Azorín et al.; 10. Urginea cf. aurantica H.Lindb.; 11. Urginea revoluta A.V.Duthie.All to the same scale in mm.

FIGURE 7. Aulostemon mzimvubuensis (van Jaarsv.) Mart.-Azorín et al.

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

FIGURE 9. Species of Austronea Mart.-Azorín et al. displayed in horizontal rows of images. 1. Austronea acarophylla (E.Brink & A.P.Dold) Mart.-Azorín et al.; 2. Austronea barkerae (Oberm.ex J.C.Manning & Goldblatt) Mart.-Azorín et al.; 3. Austronea chalumnensis (A.P.Dold & E.Brink) Mart.-Azorín et al.; 4. Austronea densiflora Mart.-Azorín et al.

FIGURE 10. Species of Austronea Mart.-Azorín et al. displayed in horizontal rows of images. 1. Austronea grandiflora Mart.-Azorín et al.; 2. Austronea hispidoplicata Mart.-Azorín et al.; 3. Austronea linearis Mart.-Azorín et al.; 4. Austronea marginata (Thunb.) Mart.- Azorín et al.

FIGURE 11. Species of Austronea Mart.-Azorín et al. displayed in horizontal rows of images. 1. Austronea oblongifolia Mart.-Azorín et al.; 2. Austronea olifanta Mart.-Azorín et al.; 3. Austronea papillosa Mart.-Azorín et al.; 4. Austronea patersoniae Mart.-Azorín et al.

FIGURE 12. Species of Austronea Mart.-Azorín et al. displayed in horizontal rows of images. 1. Austronea phyllopogon Mart.-Azorín et al.; 2. Austronea pinguis Mart.-Azorín et al. (Photograph 2.1 by J. Slade); 3. Austronea pulchromarginata (J.C.Manning & Goldblatt) Mart.-Azorín et al.; 4. Austronea trichophylla Mart.-Azorín et al.; 5. Austronea vermiformis (J.C.Manning & Goldblatt) Mart.-Azorín et al.

FIGURE 13. Species of Boosia Speta. 1. Boosia macrocentra (Baker) Speta from Phillips (1924: pl. 142, as Urginea macrocentra Baker); 2. Boosia flagellaris (T.J.Edwards et al.) Mart.-Azorín et al. from Edwards et al. (2005: 123, as Drimia flagellaris Edwards et al.).

FIGURE 14. Species of Boosia Speta displayed in horizontal rows of images. 1. Boosia flagellaris (T.J.Edwards et al.) Mart.-Azorín et al.; 2. Boosia macrocentra (Baker) Speta; 3. Boosia modesta (Baker) Mart.-Azorín et al.; 4. Boosia natalensis (Baker) Mart.-Azorín et al.

FIGURE 15. Species of Boosia Speta displayed in horizontal rows of images. 1. Boosia nyasae (Baker) Mart.-Azorín et al.; 2. Boosia rubella (Baker) Mart.-Azorín et al.; 3. Boosia saniensis (Hilliard & B.L.Burtt) Mart.-Azorín et al.; 4. Boosia tenella (Baker) Mart.-Azorín et al.

FIGURE 16. Bowiea volubilis Harv. ex T.Moore & Mast. from: 1. Hooker f. (1867); 2. Dyer (1941); and 3. Bowiea gariepensis van Jaarsv. from Reid et al. (1990: pl. 2007).

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

FIGURE 52. Species of Squilla Steihn. 1. Squilla maritima (L.) Steinh. from Clusius (1601: 171, as Scilla hispanica); 2. Squilla maritima (L.) Steinh.from Maire (1958: 163, as Urginea maritima); 3. Squilla maritima (L.) Steinh. from Ker Gawler (1806: t. 918, as Ornithogalum squilla); 4. Squilla undulata (Desf.) Mart.-Azorín et al. from Jacquin (1812: pl. 30, as Drimia purpurascens J.Jacq.); 5. Squilla undulata (Desf.) Mart.-Azorín et al. from Desfontaines (1798: t. 88, as Scilla undulata Desf.).

FIGURE 53. Species of Squilla Steihn. displayed in horizontal rows of images. 1. Squilla hesperia (Webb & Berthel.) Mart.-Azorín et al.; 2. Squilla maritima (L.) Steinh.; 3. Squilla palaestina (M.B.Crespo et al.) Mart.-Azorín et al.

FIGURE 54. Species of Squilla Steinh.displayed in horizontal rows of images. 1. Squilla pancration Steinh.; 2. Squilla serotina (Schousb.) Mart.-Azorín et al.; 3. Squilla undulata (Desf.) Mart.-Azorín et al. (Photograph of leaves by J. Quiles).