Rhodocodon Baker

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 : 72-78

16. Rhodocodon Baker

in J. Linn. Soc., Bot. 18: 280, t. 8 (1881) ( Figs 39–42 ). Typus generis:— R. madagascariensis Baker (holotype) = Rhodocodon mascarenensis (Baker) Knirsch, Mart. -Azorín & Wetschnig ≡ Urginea mascarenensis Baker ≡ Drimia sect. Rhodocodon (Baker) J.C.Manning & Goldblatt in Strelitzia 40: 155 (2018). Typus sectionis:— R. madagascariensis Baker (holotype).

Description:—Bulbous geophyte. Bulb usually hypogeal, sometimes epigeal and epiphytic, ovoid to subglobose, bulb scales compact. Roots thickened and branched. Leaves 1‒8 per bulb, from filiform to flattened and wide, sometimes with dorsal keel, green, glabrous. Inflorescence racemose, usually elongated and multiflowered, 5‒70 cm long, with 10‒120 flowers; peduncle erect, terete glabrous, smooth, 9‒38 cm long; pedicels from erect to patent or nodding, 1‒40 mm long. Bracts small, lanceolate, 1.0‒ 2.5 mm long, with lanceolate-oblong spur, 0.1‒2.0 mm long, both gradually diminishing in size to the apex of the raceme; bracteoles absent or present but indistinct. Flowers urceolate to campanulate, erect-patent to nodding, lasting for 3‒7 days. Tepals 6, 3.4‒12.0 mm long, white, greenish, brownish, rose or red coloured, fused for most of their length with free apical lobes that are from erect to reflexed, with single central nerve. Stamens 6, shorter than perigone segments, mostly included in perigone tube and connivent to ovary and style; filaments adnate to perigone and arising from lower half of tube, filiform to flattened, free portions 0.3‒3.5 mm long; anthers dorsifixed, sagittate or oblong, dehiscing along their whole length but sometimes appearing pore like at the apex. Ovary subglobose, conical to cylindrical, trigonous, with 2‒17 ovules per locule, collateral to more or less loosely arranged; style filiform, erect, 1.3‒5.0 mm long; stigma simple or inconspicuously 3-lobed. Capsule oblongtriangular to subglobose, 3‒12 mm long, valves completely dehiscing from base, withered tepals usually persisting at base of dehisced capsule (rarely abscising from base and remaining laterally attached at the top of the capsule). Seeds 2.0‒ 6.2 mm long, ellipsoidal or somewhat compressed, with black, glossy, brittle testa and distinct mostly light brown or rarely black raphe, sometimes absent; or rarely fusiform, flat, testa brown, brittle, lacking raphe, with sinuous anticlinal cell walls.

Number of species and distribution:— Rhodocodon currently includes 19 species that are restricted to Madagascar ( Fig. 29 ). It is therefore the only urgineoid genus that has been recorded so far in the Madagascan Region in the sense of Takhtajan (1986; see also Martínez-Azorín et al. 2023a). For further information on Rhodocodon species see Knirsch et al. (2015, 2016, 2019).

Karyology:—Apparently not studied yet ( Goldblatt et al. 2012).

History, diagnostic characters, and taxonomic relationships:— Baker (1874a) described Urginea mascarenensis Baker (1874a: 363) from Madagascar, with hysteranthous filiform leaves, campanulate white nodding flowers and spurred bracts. Few years later, Baker (1881) described Rhodocodon based on the distinct flower morphology of the latter species, showing tepals connate for most of their length and short free apical lobes, the included stamens with adnate and short filaments arising from the lower half of the perigone, and the ovate ovary and erect narrow style, and included a single species: R. madagascariensis Baker (1881: 280) a later synonym of U. mascarenensis . Perrier de la Bâthie (1938) presented a revision on the endemic Madagascan genus Rhodocodon , where six species and four varieties were recognised, although those names were not validly published ( Knirsch et al. 2015). The recent revision of Rhodocodon by Knirsch et al. (2015, 2016, 2019) accepted 19 species in the genus, which is characterised by flowers urceolate to campanulate, lasting for 3‒7 days; perigone segments fused for most of their length, commonly persisting at the base of ripe capsules; filaments adnate to the perigone and arising from the lower half of the tube; and seeds ellipsoidal or somewhat compressed, usually with a distinct raphe, or rarely fusiform and flat ( Brudermann et al. 2018).

Previous phylogenetic work has shown Madagascan samples of Rhodocodon to be monophyletic in a well-supported clade ( Pfosser & Speta 2001, 2004, Pfosser et al. 2006, 2012, Knirsch 2012), which is in agreement with the phylogenetic results of Martínez-Azorín et al. (2023a), in which 12 samples of the genus form a well supported clade that is sister to Triandra from western South Africa. Therefore, we here accept Rhodocodon as an endemic Madagascan genus showing a unique combination of morphological characters that form an independent phylogenetic lineage.

Accepted species:—

Rhodocodon apiculatus H.Perrier ex Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 106 (2015) ≡ R. apiculatus H.Perrier, Fl. Madagasc. Fam. 40: 122 (1938), nom. inval. Type:— MADAGASCAR. Rocailles calcaires, eocènes, boisées, près de Majunga [Mahajanga], December 1924, Perrier 16821 (P00573458! holo.).

Rhodocodon calcicola Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 108 (2015) ≡ R. calcicola H.Perrier, Fl. Madagasc. Fam. 40: 115 (1938), nom. inval. ≡ Drimia calcicola (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004), nom. inval. ( Figs 39.2 , 40.1 ). Type:— MADAGASCAR. Province Mahajanga, few kilometers north of Mahajanga, elev. 60 m, ex hort. 30 April 2014, W. Knirsch, A. Sieder & J. Andriantiana 6293 (GZU000322310! holo.).

Rhodocodon campanulatus Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 110 (2015) ≡ R. rotundus var. campanulatus H.Perrier, Fl. Madagasc. Fam. 40: 122 (1938), nom. inval. ( Figs 3.11 , 40.2 ). Type:— MADAGASCAR. Province Majahanga, rochers basaltiques, découverts et humides pendant les pluies. Plateau d’Antanimena entre la Betsiboka et la Mahavavy, May 1905, Perrier 10947 (P01848487! holo.).

Rhodocodon cryptopodus (Baker) Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 112 (2015) ≡ Hyacinthus cryptopodus Baker in J. Linn. Soc., Bot. 20: 274 (1883), basionym ≡ Drimia cryptopoda (Baker) Pfosser, Wetschnig & Speta in Linzer Biol. Beitr. 38(2): 1736 (2006) ( Figs 3.12 , 40.3 ). Type:— MADAGASCAR. Central Madagascar, October 1882, R. Baron 2164 (K000098171! holo.: the two fragments/specimens in the upper left hand side corner of the sheet).

Rhodocodon cyathiformis H.Perrier ex Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 114 (2015) ≡ R. cyathiformis H.Perrier, Fl. Madagasc. Fam. 40: 118 (1938), nom. inval. ≡ Drimia cyathiformis (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004), nom. inval. ( Figs 3.13 , 38.2 , 40.4 ). Type:— MADAGASCAR. Rochers ombragés, Firingalava (Boina), March 1898, Perrier 516 (P00573463! holo.; P00573464!, P00573465! iso.).

Rhodocodon floribundus H.Perrier ex Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 116 (2015) ≡ R. floribundus H.Perrier, Fl. Madagasc. Fam. 40: 117 (1938), nom. inval. ≡ Drimia floribunda (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004), nom. inval. ( Figs 3.14 , 40.5 ). Type:— MADAGASCAR. Rocailles (gneiss), elev. 1500 m, Manankazo, au N.E. d’Ankazobe, Novembre 1913, Perrier 10960 (P00573460! holo.; P00573459! iso.).

Rhodocodon graciliscapus Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 118 (2015) ≡ R. calcicola var. graciliscapus H.Perrier, Fl. Madagasc. Fam. 40: 116 (1938), nom. inval. ( Figs 3.15 , 41.1 ). Type:— MADAGASCAR. Environs d’Ampasimentera (Boina), paturages, fl. verdatres, October 1906, Perrier 10946 (P00573466! holo.).

Rhodocodon intermedius H.Perrier ex Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 120 (2015) ≡ R. intermedius H.Perrier, Fl. Madagasc. Fam. 40: 116 (1938), nom. inval. ≡ Drimia intermedia (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004), nom. inval. ( Figs 3.16 , 41.2 ). Type:— MADAGASCAR. Plaine sèche-alluvionnaire, sur la rive droite de la Mananjeba (Andavakoera), July 1963, Perrier 10965 (P00573461! holo.; P00573462! iso.).

Rhodocodon jackyi Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 253(3): 202 (2016) ( Figs 3.17 , 41.3 ). Type:— MADAGASCAR. Province Toliara, region Anosy : near the village Vatambe , elev. 330 m, ex hort. 30 April 2014, W. Knirsch, A. Sieder & J. Andriantiana 6620 corresponding to WW04669 (GZU-000324829! holo.; P!, PBZT! iso.).

Rhodocodon linearifolius Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 122 (2015) ≡ R. calcicola var. linearifolius H.Perrier, Fl. Madagasc. Fam. 40: 116 (1938), nom. inval. ( Fig. 39.2 ). Type:— MADAGASCAR. Rocailles denudées—sur les labradorites de l’Antanimena, December 1924, Perrier 16824 (P00516239! holo.; P00516238! iso.).

Rhodocodon mascarenensis (Baker) Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 122 (2015) ≡ Urginea mascarenensis Baker in J. Bot. 12: 363 (1874), basionym ≡ Drimia mascarenensis (Baker) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004) ( Figs 3.18 , 41.4 ). Type:— MADAGASCAR. Bojer & Hilsenberg s.n. (BM000797876! holo.).

= R. madagascariensis Baker in J. Linn. Soc., Bot. 18: 280, t. 8 (1881) ( Fig. 39.2 ). Type:— MADAGASCAR. N & E of Ankaratra Mts, s.d., Langley Kitching s.n. (K000099861! lecto. designated by Knirsch et al. 2015).

Rhodocodon monophyllus Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 125 (2015) ≡ R. calcicola var. monophyllus H.Perrier, Fl. Madagasc. Fam. 40: 115 (1938), nom. inval. ( Figs 3.19 , 38.2 , 41.5 ). Type:— MADAGASCAR. Bois sur les rocailles gneissiques, sur la rivière Besafotra, affluent de droite du Menavava, Boina, bassin de la Betsiboka, Novembre 1899, Perrier 990 (P02160895! holo.).

Rhodocodon perrieri Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 414(3): 138 (2019) ( Figs 3.20 , 42.1 ). Type:— MADAGASCAR. Antananarivo Province: region Analamanga, district Manjakandriana , mountains ca. 4 km SE of Ambohimiadana, elev. 1500‒1600 m, ex hort. 30 April 2014, W. Knirsch, A. Sieder & J. Andriantiana AS6564 (ABH! holo.).

Rhodocodon petrae Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 414(3): 130 (2019) ( Fig. 42.2 ). Type:— MADAGASCAR. Mahajanga Province: Namoroka National Park, near the Tsingy , elev. 120 m, ex hort. 15 May 2013, J. Andriantiana s.n. corresponding to WW04826 (ABH82488! holo.).

Rhodocodon rotundus H.Perrier ex Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 195(2): 127 (2015) ≡ R. rotundus H.Perrier, Fl. Madagasc. Fam. 40: 120 (1938), nom. inval. ≡ Drimia rotunda (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004), nom. inval. ( Figs 4.1 , 38.2 ). Type:— MADAGASCAR. Versant W de l’Isalo, grès, elev. 400 m, October 1924, Perrier 16583 (P00573467! holo.).

Rhodocodon rubescens Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 414(3): 136 (2019) ( Fig. 42.3 ). Type:— MADAGASCAR. Mahajanga Province: district of Sofia, region Antsohihy , ca. 6 km SW of town Anjiamangirana , elev. 320 m, ex hort. 10 April 2016, W. Knirsch, A. Sieder & J. Andriantiana s.n. (ABH! holo.).

Rhodocodon siederi Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 414(3): 141 (2019) ( Figs 4.2 , 42.4 ). Type:— MADAGASCAR. Antananarivo Province: ca. 40 km west of Antananarivo, Antongona, Imeritsiatosika , rocky mountains 4 km E of Arivonimamo, elev. 1497 m, growing among grasses, ex hort. 30 April 2014, W. Knirsch, A. Sieder & J. Andriantiana AS6728 (ABH82498! holo.).

Rhodocodon urgineoides Baker in Hooker’s Icon. Pl. 25: t. 2492 (1896) ≡ Rhadamanthus urgineoides (Baker) Speta in Phyton (Horn, Austria) 38: 77 (1998) ≡ Drimia urgineoides (Baker) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004) ( Figs 4.3 , 39.2 , 42.5 ). Type:— MADAGASCAR. Ivohimanitra forest, October 1894, C.H. Forsyth Major 44 (G00168520! lecto. designated by Knirsch et al. 2015; K000099856! isolecto.).

Rhodocodon viridans Knirsch, Mart. - Azorín & Wetschnig in Phytotaxa 414(3): 133 (2019). Type:— MADAGASCAR. Antsiranana Province: ca. 20 km N of Vohemar, Ambatoharanana, Cap Manambato , ex hort. in Austria , Steiermark , Sankt Marein on 15 May 2015, W. Röösli & R. Hoffmann s.n. corresponding to WW04675 (ABH82494! holo.).

Figures

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 29. Approximate distribution ranges of accepted genera in Urgineoideae.

FIGURE 38. Species of Rhadamanthus Salisb. displayed in horizontal rows of images. 1. Rhadamanthus fasciatus B.Nord.; 2. Rhadamanthus rigidifolius (Baker) Mart.-Azorín et al.; 3. Rhadamanthus secundus B.Nord. (Photographs in flower by H. Kolberg); 4. Rhadamanthus urantherus R.A.Dyer.

FIGURE 39. Species of Rhodocodon Baker. 1. Rhodocodon mascarenensis (Baker) Knirsch et al. from Baker (1881: t. 8, as Rhodocodon madagascariensis Baker); 2. Rhodocodon mascarenensis (Baker) Knirsch et al., R. cyathiformis H.Perrier ex Knirsch et al., R. rotundus H.Perrier ex Knirsch et al., R. calcicola Knirsch et al., R. monophyllus Knirsch et al., R. linearifolius Knirsch et al., and R. urgineoides Baker et al. from Perrier de la Bâthie (1938: 121).

FIGURE 40. Species of Rhodocodon Baker displayed in horizontal rows of images. 1. Rhodocodon calcicola Knirsch et al.; 2. Rhodocodon campanulatus Knirsch et al.; 3. Rhodocodon cryptopodus (Baker) Knirsch et al.; 4. Rhodocodon cyathiformis H.Perrier ex Knirsch et al.; 5. Rhodocodon floribundus H.Perrier ex Knirsch et al.

FIGURE 41. Species of Rhodocodon Baker displayed in horizontal rows of images. 1. Rhodocodon graciliscapus Knirsch et al.; 2. Rhodocodon intermedius H.Perrier ex Knirsch et al.; 3. Rhodocodon jackyi Knirsch et al.; 4. Rhodocodon mascarenensis (Baker) Knirsch et al.; 5. Rhodocodon monophyllus Knirsch et al.

FIGURE 42. Species of Rhodocodon Baker displayed in horizontal rows of images. 1. Rhodocodon perrieri Knirsch et al.; 2. Rhodocodon petrae Knirsch et al.; 3. Rhodocodon rubescens Knirsch et al.; 4. Rhodocodon siederi Knirsch et al.; 5. Rhodocodon urgineoides Baker.