Fusifilum Raf., Fl. Tellur.

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 : 43-47

7. Fusifilum Raf., Fl. Tellur. 2: 27 (1837)

( Figs 21–23 ).

Typus generis:— Drimia physodes (Jacq.) Jessop (lectotype, designated by Deb & Dasgupta in J. Econ. Tax. Bot. 3: 823. 1982).

= Physodia Salisb., Gen. Pl. : 37 (1866) ≡ Urginea sect. Physodia (Salisb.) Baker in J. Linn. Soc., Bot. 13: 216 (1873) ≡ Drimia sect. Physodia (Salisb.) J.C.Manning & Goldblatt in Strelitzia 40: 92 (2018). Typus generis:— Physodia pusilla (Jacq.) U.M̧ll.-Doblies (lectotype, designated by Deb & Dasgupta in J. Econ. Tax. Bot. 3: 823. 1982).

Description:—Small bulbous geophyte. Bulb hypogeal, ovoid to subglobose, 1‒7 cm in diam., sometimes with short bulb neck, solitary or commonly clump forming, inner scales fleshy, white and compact, outer scales membranous. Roots thickened and branched. Leaves 1‒10, hysteranthous, 3‒15 cm long and 1‒30 mm wide, usually flat, canaliculate, lanceolate, spreading, green, glabrous or papillose. Inflorescence a dense, short or elongated raceme, 1‒8 cm long, with 6‒60(–120) flowers; peduncle erect, terete, smooth or papillose, 4‒15 cm long, greenish to dark purple; pedicels patent-spreading, 8‒30 mm long, glabrous. Bracts lanceolate, small, persistent, lowermost 1‒2 mm long with short spur of ca. 1 mm long; bracteoles absent. Flowers stellate, suberect, diurnal, opening in afternoon. Tepals 6, biseriate, 4‒7 mm long, spreading, white, with distinct greenish to purplish longitudinal band on abaxial side, free to shortly fused at base for less than 1 mm, outers sometimes slightly reflexed. Stamens 6, spreading; filaments 3‒4 mm long, fusiform, narrowed to base and apex, white, with distinct papillae along basal third; anthers oblong, medifixed, to 1 mm long, opening by longitudinal slits. Ovary oblong, truncate to style, 2‒4 mm long, white, with 10‒15 ovules per locule. Style 2‒3 mm long, erect, white, filiform. Capsule ovate to sphaerical, trigonous with blunt edges in cross-section, 6‒12 mm long, pale brown. Seeds subellipsoid, (3–) 4‒8 mm long, flattened with prominent central embryo and wide, flat or incurved wings, dark brown to black, glossy, with sinuous anticlinal testa cell walls.

Number of species and distribution:—It includes 17 species, occurring in Southern Africa ( Fig. 8 ) and restricted to the Cape, Karoo-Namib and Uzambara-Zululand Regions, and the Southern Section of the Zambezian Subregion (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a). For further information on Fusifilum species see M̧ller-Doblies et al. (2001). We preliminarily accept F. magicum as originally circumscribed as belonging to Fusifilum based on its flower morphology ( Fig. 23.1 ). However, its phylogenetic position places two samples of this species in a clade related to Urginea revoluta and Boosia ( Martínez-Azorín et al. 2023a) . Further studies, including new phylogenetic evidence, are needed to evaluate other alternatives.

Karyology:—2n=20 ( De Wet 1957, as Urginea depressa Baker ).

History, diagnostic characters, and taxonomic relationships:— Fusifilum was described to accommodate some species of Urgineoideae having small to medium-sized, subcapitate inflorescence; white stellate, flowers; fusiform and papillate filaments; and white gynoecium. The latter character is unique and diagnostic. The original concept of the genus included species described and illustrated by Jacquin (1795) from Southern Africa and by Ruiz & Pavón (1802) from South America, the latter belonging to Oziroe Rafinesque (1837: 53) . However, the current concept of Fusifilum excludes the American taxon, and Deb & Dasgupta (1982) typified the genus on F. physodes (Jacquin 1795: 18) Speta (1998b: 69) (M̧ller-Doblies et al. 2001, Martínez-Azorín et al. 2015). Later, other generic names, such as Physodia Salisbury (1866: 37) , were applied to this distinct group of plants.

Speta (1998b) accepted Fusifilum to include seven species, and M̧ller-Doblies et al. (2001) presented a taxonomic revision that included 16 species, four from Speta’s (1998b) treatment, one from Tang & Weiglin (2001), 10 as new, and one new combination. Types of their newly described species are still not deposited in most of the studied herbaria and their morphological descriptions are incomplete, which has impeded further taxonomic work in the genus during the last two decades.

The phylogenetic analyses of Manning et al. (2004) included four samples considered by them to belong to Fusifilum , although only F. physodes is a true Fusifilum , following circumscriptions of M̧ller-Doblies et al. (2001) and the present work. The supposed polyphyly of Fusifilum , in the sense of Manning et al. (2004), was one of the main reasons offered to justify the lumping of the evidently broad diversity in Urgineoideae in a single, extremely variable Drimia , excluding Bowiea . However, the correct genus adscription of these species does, in fact, dissolve the polyphyly of Fusifilum and support an analytic rather than a synthetic treatment in the subfamily ( Martínez-Azorín et al. 2023a) since ‘ F. dregei ’ is a species of Urgineopsis Compton (as accepted in this work) and ‘ F. marginata ’ belongs to Austronea ( Martínez-Azorín et al. 2019a) —a genus sister to Fusifilum . The names Fusifilum calcarata and F. marginata sensu Manning et al. (2004) were never formally combined or published prior to 2004.

Manning & Goldblatt (2018) described Drimia sect. Physodia , which mostly fits the concept of Fusifilum in this work, but excluding Urginea virens that belongs to Austronea ( Martínez-Azorín et al. 2019a) . Manning & Goldblatt (2018) placed eight of the species described by M̧ller-Doblies et al. (2001) under their Drimia physodes (Jacquin 1795: 18) Jessop (1977: 300) , commenting that those names were not validly published since the types were not deposited following Art. 40.7 of the ICN. However, this interpretation is incorrect, as the Shenzhen Code only recommends that the type material, especially the holotype, should be deposited in a public herbarium or other public collection allowing access and warranting long-term conservation (Art. 7 Rec. 7A1 of the ICN). Therefore, these M̧ller-Doblies names are valid following the current Shenzhen Code, and hence must be accepted. Our field work in Southern Africa over the last decades has enabled us to observe clear morphological differences in support of the acceptance of the species of M̧ller-Doblies et al. (2001). Our phylogenetic analyses show 20 samples of Fusifilum , including several type collections provided by U. & D. M̧ller-Doblies, forming a strongly supported clade sister to Austronea ( Martínez-Azorín et al. 2023a) , further supporting the acceptance of this genus based on a unique syndrome of morphological characters, biogeography, and phylogenetic relationships.

Accepted species and required new combination:—

Fusifilum bruce-bayeri U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 479 (2001). Type:— SOUTH AFRICA. Western Cape. Worcester (3319): Worcester West , Polhillia waltersii Nature Reserve , (–CB), elev. 250 m, 15 August 1984, M̧ller-Doblies 84017f (PRE holo.; B, BTU, E, G, GRA, K, M, MO, NAS, NBG, S, W, WIND, Z iso.; ex cult. BTU5931: ibidem. Not deposited in the cited herbaria).

Fusifilum capitatum (Hook.f.) Speta in Phyton (Horn, Austria) 38(1): 69 (1998) ≡ Ornithogalum capitatum Hook.f. in Curtis’s Bot. Mag. 89: t. 5388 (1863), basionym ( Figs 2.3 , 21.3 ) [non Drimia capitata Baker (1897) ] ≡ Urginea capitata (Hook.f.) Baker, Fl. Cap. (Harvey) 6(3): 465 (1897) ≡ Physodia capitata (Hook.f.) U.Müll. -Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 107(5−6): 520 (1996) ( Fig. 22.1 ). Type:— SOUTH AFRICA. British Kaffraria [part of Eastern Cape today], 1860, Cooper 208 (K000365544! lecto. designated as “holo” by Manning & Goldblatt in Strelitzia 40: 93. 2018; Z-000086255! isolecto.).

Fusifilum crenulatum U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 483 (2001) ( Fig. 22.2 ). Type:— SOUTH AFRICA. Northern Cape. Kamiesberg (3018): Koringlandskloof , 1 km on Leliefontein road from Garies / Platbakkies road, (–AC), loamy ground, elev. 1200 m, 12 August 1980, M̧ller-Doblies 80087b (PRE holo.; B, BTU, G, K, M, MO, NAS, NBG, S, WIND, Z iso.; ex cult. BTU4033: ibidem. Not deposited in the cited herbaria).

Fusifilum depressum (Baker) U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 480 (2001) ≡ Urginea depressa Baker in Bull. Herb. Boissier ser. 2, 4: 1000 (1904), basionym ≡ Drimia depressa (Baker) Jessop in J. S. African Bot. 43: 297 (1977) ( Fig. 22.3 ). Type:— SOUTH AFRICA. Gauteng. Johannesburg (2628): Transvaal, Modderfontein, (–CB), September 1897, Conrath 687 (Z000027555! lecto. designated here). Note:—The lectotype designation by Jessop in J. S. African Bot. 43: 297 (1977) is not acceptable according to Art. 8.2 Ex. 3 of the Code (see Martínez-Azorín & Crespo 2014; the voucher K000857414! bears a different collection date of 9 October 1898, perhaps erroneously transcribed. In that case it is a true isolectotype, as well as GZU!).

Fusifilum emdeorum J.S.Tang & Weiglin in Feddes Repert. 112(7−8): 505 (2001). Type:— SOUTH AFRICA. Western Cape. Worcester (3319): Farm Gevonden, E end of Dutoitskloof , (–CA), elev. 300 m, 22 July 1986, M̧ller-Doblies 86017c (PRE holo.; B, BTU, K, M, MO, NBG, S, WIND, Z iso.; ex cult. BTU6466: ibidem and G, GRA, LI, NAS, W. Not deposited in the cited herbaria).

Fusifilum gifbergense U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 479 (2001). Type:— SOUTH AFRICA. Western Cape. Vanrhynsdorp (3118): Snorkfontein , half way up the Gifberg , (–DD), elev. 500−600 m, 16 August 1980, M̧ller-Doblies 80105v (PRE holo.; B, BTU, E, G, K, LI, M, MO, NAS, NBG, P, S, W, WIND, Z iso.; ex cult. BTU4239: ibidem. Not deposited in the cited herbaria).

Fusifilum glaucum U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 481 (2001). Type:— SOUTH AFRICA. Western Cape. Vanrhynsdorp (3118): Zandkraal , Gifberg road, 2.7 km S of turn off to Zandkraal , (–DB), elev. 150 m, 16 August 1980 veg., M̧ller-Doblies 80104e (PRE holo.; B, BTU, K, M; ex cult. BTU4201: B, BTU, G, K, LI, M, MO, NAS, NBG, PRE, S, WIND, Z iso. Not deposited in the cited herbaria).

Fusifilum hei U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 480 (2001). Type:— SOUTH AFRICA. Western Cape. Montagu (3320): Matjesfontein , Farm Boelhouer (Bullover), 4 km on Keurfontein (Driekop) road, (–BA), elev. 1050 m, 02 August 1986 veg., M̧ller-Doblies 86032a (PRE holo.; B, BTU, K, NAS, S iso; ex cult. BTU6607: ibidem. Not deposited in the cited herbaria).

Fusifilum magicum U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 491 (2001) ( Fig. 23.1 ). Type:— SOUTH AFRICA. Western Cape. Beaufort West (3222): Boesmansklip , 2.5 km from N1 to Molteno Pass , (–BC), elev. 900 m, 15 January 1984, M̧ller-Doblies 84129c (PRE holo; B, BTU, K, NBG. Not deposited in the cited herbaria).

Fusifilum minus (A.V.Duthie) Speta in Phyton (Horn, Austria) 38(1): 69 (1998) ≡ Urginea minor A.V.Duthie in Ann. Univ. Stellenbosch 6. Sect. A, No. 2, 11 (1928), basionym ≡ Drimia minor (A.V.Duthie) Jessop in J. S. African Bot. 43(4): 306 (1977) ≡ Physodia minor (A.V.Duthie) Salisb. ex U.Müll. -Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 107(5−6): 520 (1996). Type:— SOUTH AFRICA. Western Cape. Cape Town (3318): Stellenbosch Flats, (–DD), March 1924, Duthie s.n. STE1546 (NBG0197706! lecto. designated as “holo.” by Jessop in J. S. African Bot. 43: 306. 1977); K000257366!, PRE0049735! isolecto.).

Fusifilum montanum (A.P.Dold & E.Brink) A.P.Dold, Mart. -Azorín, M.B.Crespo, M.Pinter & Wetschnig comb. nov. ≡ Drimia montana A.P.Dold & E.Brink in Bothalia 36(1): 64 (2006) basionym ≡ Drimia montana A.P.Dold & E.Brink in Taxon 63: 1130 (2014), nom. superfl. ≡ Fusifilum montanum (A.P.Dold & E.Brink) A.P.Dold, Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 201(2): 168 (2015), comb. inval. ( Figs 2.4 , 23.2 ). Type:— SOUTH AFRICA. Eastern Cape. Fort Beaufort (3226): Groot Winterberg, The Hoek farm, (–AD), elev. 2150 m, 1 January 2004, A.P. Dold 4633 (GRA0000460! lecto. designated here: only the plants in flower and fruit lacking leaves “Pressed in December at time of collection” [though corrected to January by A.P. Dold 08/10/2014; A.P. Dold pers. comm.], which are placed on the upper half of the sheet.).

Comments:—In the light of the recentmost proposal by Mosyakin & McNeill (2022) to amend Art. 8 of the ICN, after which the information in the protologue will rule, and considering that the protologue in Dold & Brink (2006) only indicates a single collecting date (01- 01-2004) in the holotype designation, a lectotypification is needed from the mixed type collection GRA0000460. Based on the new considerations, the description made by Martínez-Azorín & Crespo (2014: 1330) and the combination in Martínez-Azorín et al. (2015: 168) are therefore nomenclaturally inoperative, since it is either superfluous or not valid.

Fusifilum oliverorum U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 487 (2001). Type:— SOUTH AFRICA. Western Cape. Bredasdorp (3420): Moerasfontein , 1.3 km NW of Skipskop / Bredasdorp road, (–CB), elev. 30 m, 18 August 1984 veg., Oliver & M̧ller-Doblies 84025a (PRE holo.; B, BTU, K, M, MO, NBG, S, WIND, Z iso; ex cult. BTU 6000, 6001, 6002, 6003: ibidem. Not deposited in the cited herbaria).

Fusifilum papillosum U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 487 (2001). Type:— SOUTH AFRICA. Northern Cape. Calvinia (3119): Glen Lyon States , SE of Nieuwoudtville, (–AC), elev. 750 m, 9 November 1978 fl., M̧ller-Doblies 78072m (PRE holo.; B, BTU iso; ex cult. BTU 2156: B, BTU, K, NBG, PRE. Not deposited in the cited herbaria).

Fusifilum physodes (Jacq.) Speta in Phyton (Horn, Austria) 38(1): 69 (1998) ≡ Anthericum physodes Jacq., Icon. Pl. Rar. 2(16): 18, t. 418 (1795) diagnosis; Coll. Suppl.: 94 (1797) description ≡ Phalangium physodes (Jacq.) Pers., Syn. Pl. 1: 369 (1805) ≡ Albuca physodes (Jacq.) Ker Gawl. in Curtis’s Bot. Mag. 26: t. 1046 (1807) ≡ Caesia physodes (Jacq.) Spreng., Syst. Veg. 2: 88 (1825) ≡ Idothea physodes (Jacq.) Kunth, Enum. Pl. 4: 345 (1843) ≡ Urginea physodes (Jacq.) Baker in J. Linn. Soc., Bot. 13: 217 (1873) ≡ Physodia physodes (Jacq.) U.M̧ll.-Doblies et al. in Feddes Repert. 107(5−6): 519 (1996) ≡ Drimia physodes (Jacq.) Jessop in J. S. African Bot. 43(4): 300 (1977) ( Figs 2.5 , 21.1 , 23.3 ). Type:— SOUTH AFRICA. Illustration in Jacq., Icon. Pl. Rar. 2(16): t. 418 (1795) ( Fig. 21.1 ) (lecto. designated by Jessop in J. S. African Bot. 43: 300. 1977).

= Ornithogalum melanopus Dinter ex Ŝlch in Beitr. Fl. SW-Afr., Diss. Univ. M̧nchen: 70 (1960).

Fusifilum pusillum (Jacq.) Speta in Phyton (Horn, Austria) 38(1): 70 (1998) ≡ Anthericum pusillum Jacq., Icon. Pl. Rar. 2(16): 18, t. 417 (1795) diagnosis; Coll. Suppl.: 95 (1797) description, basionym [non Drimia pusilla Jacq., Coll. Suppl. : 42 (1797)] ≡ Caesia pusilla (Jacq.) Spreng., Syst. Veg. 2: 88 (1825) ≡ Idothea drimioides Kunth, Enum. Pl. 4: 345 (1843), nom. nov. [non I. pusilla (Jacq.) Kunth = D. elata Jacq. ] ≡ Urginea pusilla (Jacq.) Baker in J. Linn. Soc., Bot. 13: 217 (1873) ≡ Physodia pusilla (Jacq.) U.M̧ll.-Doblies et al. in Feddes Repert. 107(5−6): 520 (1996). Type:— SOUTH AFRICA. Illustration in Jacq., Icon. Pl. Rar. 2 (16): t. 417 (1795) ( Fig. 21.2 ) (lecto. designated by Jessop in J. S. African Bot. 43: 295. 1977).

Fusifilum spirale U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 483 (2001). Type:— SOUTH AFRICA. Northern Cape. Springbok (2917): Henrietsberg , 10−12 km N of Steinkopf, (–BB), elev. 800−1100 m, 14 August 1980, M̧ller-Doblies 80096a (PRE holo.; B, BTU, K, MO, NAS, WIND, Z iso; ex cult. BTU 4085: G, LI, M, NBG, S. Not deposited in the cited herbaria).

Fusifilum stoloniferum U.Müll. - Doblies, J.S.Tang & D.Müll. - Doblies in Feddes Repert. 112(7−8): 484 (2001). Type:— SOUTH AFRICA. Western Cape. Beaufort West (3222): Karoo National Park , escarpment W of tall TV-Tower (N15D), (–AD), elev. 1800 m, 27 August 1984 veg., M̧ller-Doblies 84051c (PRE holo.; B, BR, BTU, K, LI, M, MO, NAS, NBG, S, WIND, Z iso; ex cult. BTU 6110, 6111, 6112: ibidem and BOL, E, G, GRA, KMG, NH, NU, P, PE, PEU, PRU, UNIN, W, WAG, WRSL. Not deposited in the cited herbaria).

Figures

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 21. Species of Fusifilum Raf. 1. Fusifilum physodes (Jacq.) Speta from Jacquin (1795: t. 1046, as Anthericum physodes Jacq.); 2. Fusifilum pusillum (Jacq.) Speta from Jacquin (1795: t. 417, as Anthericum pusillum Jacq.); 3. Fusifilum capitatum (Hook.f.) Speta from Hooker f. (1863: t. 5388, as Ornithogalum capitatum Hook.f.).

FIGURE 22. Species of Fusifilum Raf. displayed in horizontal rows of images. 1. Fusifilum capitatum (Hook.) Speta; 2. Fusifilum crenulatum U.M̧ll.-Doblies et al.; 3. Fusifilum depressum (Baker) U.M̧ll.-Doblies et al.

FIGURE 23. Species of Fusifilum Raf. displayed in horizontal rows of images. 1. Fusifilum magicum U.M̧ll.-Doblies et al.; 2. Fusifilum montanum (A.P.Dold & E.Brink) A.P.Dold et al.; 3. Fusifilum physodes (Jacq.) Speta.