A generic monograph of the Hyacinthaceae subfamily Urgineoideae Author Martínez-Azorín, Mario Depto. Ciencias Ambientales y Recursos Naturales (dCARN), Universidad de Alicante, P. O. Box 99, ES- 03080 Alicante, Spain. & E-mail: mmartinez @ ua. es; ORCID: https: // orcid. org / 0000 - 0002 - 2605 - 9575 mmartinez@ua.es Author Crespo, Manuel B. Depto. Ciencias Ambientales y Recursos Naturales (dCARN), Universidad de Alicante, P. O. Box 99, ES- 03080 Alicante, Spain. & E-mail: crespo @ ua. es; ORCID: https: // orcid. org / 0000 - 0002 - 3294 - 5637 crespo@ua.es Author Alonso-Vargas, María Ángeles Depto. Ciencias Ambientales y Recursos Naturales (dCARN), Universidad de Alicante, P. O. Box 99, ES- 03080 Alicante, Spain. & E-mail: ma. alonso @ ua. es; ORCID: https: // orcid. org / 0000 - 0003 - 3768 - 9203 ma.alonso@ua.es Author Pinter, Michael Depto. Ciencias Ambientales y Recursos Naturales (dCARN), Universidad de Alicante, P. O. Box 99, ES- 03080 Alicante, Spain. & Institute of Biology, NAWI Graz, Division Plant Sciences, Karl-Franzens University Graz, Holteigasse 6, A- 8010 Graz, Austria. & E-mail: michael. pinter @ uni-graz. at; ORCID: https: // orcid. org / 0000 - 0002 - 6055 - 6989 michael.pinter@uni-graz.at Author Crouch, Neil R. BRAM, South African National Biodiversity Institute, P. O. Box 52099, Berea Road 4007, South Africa. & School of Chemistry and Physics, University of KwaZulu-Natal, Durban 4041, South Africa. & E-mail: N. Crouch @ sanbi. org. za; ORCID: https: // orcid. org / 0000 - 0002 - 4938 - 5840. rouch@sanbi.org.za Author Dold, Anthony P. Selmar Schonland Herbarium, Department of Botany, Rhodes University, Makhanda, 6140, South Africa. & E-mail: t. dold @ ru. ac. za; ORCID: https: // orcid. org / 0000 - 0002 - 9497 - 7503 t.dold@ru.ac.za Author Mucina, Ladislav Harry Butler Institute, Murdoch University, 90 South Street, Building 390, Murdoch WA 6150, Perth, Australia. & Dept. of Geography & Environmental Studies, Stellenbosch University, Private Bag X 1, Matieland 7602, Stellenbosch, South Africa. & E-mail: ladislav. mucina @ murdoch. edu. au; ORCID: https: // orcid. org / 0000 - 0003 - 0317 - 8886 ladislav.mucina@murdoch.edu.au Author Pfosser, Martin Biocenter Linz, J. - W. - Klein-Str. 73, A- 4040 Linz, Austria. & E-mail: martin. pfosser @ ooelkg. at; ORCID: https: // orcid. org / 0000 - 0003 - 2050 - 4997 martin.pfosser@ooelkg.at Author Wetschnig, Wolfgang Depto. Ciencias Ambientales y Recursos Naturales (dCARN), Universidad de Alicante, P. O. Box 99, ES- 03080 Alicante, Spain. & Institute of Biology, NAWI Graz, Division Plant Sciences, Karl-Franzens University Graz, Holteigasse 6, A- 8010 Graz, Austria. & Depto. Ciencias Ambientales y Recursos Naturales (dCARN), Universidad de Alicante, P. O. Box 99, ES- 03080 Alicante, Spain. & E-mail: wolfgang. wetschnig @ uni-graz. at; ORCID: https: // orcid. org / 0000 - 0002 - 9245 - 029 X * Author for correspondence & Depto. Ciencias Ambientales y Recursos Naturales (dCARN), Universidad de Alicante, P. O. Box 99, ES- 03080 Alicante, Spain. wolfgang.wetschnig@uni-graz.at text Phytotaxa 2023 2023-08-31 610 1 1 143 http://dx.doi.org/10.11646/phytotaxa.610.1.1 journal article 266341 10.11646/phytotaxa.610.1.1 3f1ee302-e1e7-404f-9f87-9dee7086748c 1179-3163 8308556 8. Geschollia Speta in Stapfia 75: 169 (2001) ( Figs 24–26 ). Typus generis:— G. anomala (Baker) Speta ( holotype ). ≡ Ornithogalum subgen. Ledebouriopsis Baker (1870a: 178) ≡ Drimia subgen. Ledebouriopsis (Baker) Baker (1897: 437) ≡ Drimia sect. Ledebouriopsis (Baker) J.C. Manning & Goldblatt (2018: 24) pro parte (only Drimia anomala , D. occultans , and D. calcarata s.str. ). Typus subgeneris:— Drimia anomala (Baker) Baker ( lectotype designated by Manning & Goldblatt 2018 ). Description :—Bulbous geophyte. Bulb hypogeal or rarely semi-epigeal, mostly solitary but rarely proliferous, usually with compact scales but rarely loose, outer scales brownish and membranous. Roots thickened and branched. Leaf solitary (rarely 2‒3 in clump forming species), terete, green, commonly leathery, synanthous or hysteranthous, deciduous to evergreen, 0.5−7.0 mm in diam., smooth, glabrous, usually withering from tip showing transverse abscission plates. Inflorescence 1 or rarely 2 per bulb, racemose, usually elongated, (0.2‒) 2‒30 cm long, erect or slightly bent; peduncle elongated, erect, smooth or sometimes distinctly papillose at base; pedicels 2‒30 mm long, subpatent. Bracts lanceolate, acute, lowermost with broad spur usually longer than blade; bracteoles absent. Flowers stellate, erect-patent, diurnal, usually opening in afternoon and withering in evening. Tepals 6, biseriate, 3.2‒8.0 mm long, spreading to reflexed at full anthesis, with base usually shortly connate for ca. 1 mm but almost free in general appearance, yellowish, orange, greenish or white, with discrete brownish, green or purple stripe along middle, more evident on abaxial side. Stamens 6, erect to spreading; filaments filiform, slightly fusiform, adnate to base of tepals, commonly glabrous and smooth, rarely distinctly papillate; anthers yellow, oblong, medifixed, dehiscing along their whole length. Ovary ovate to oblong, attenuate to truncate at top, green to yellowish, sometimes with white maculae; style white, narrowly filiform to obtriangular, distinctly trigonous in section, as long as or longer than ovary; stigma slightly 3-lobed and papillose. Capsule ovate-globose, small, 3‒6 mm long, trigonous, loculicidal, 3 valves splitting to base, with withered perigone segments circumscissile below and forming apical cap. Seeds polygonal or irregularly compressed, commonly narrowly pyramidal, pointed, comparatively small, 1.0‒2.4(‒3.0) mm long, light brown to black, with wrinkled-rugose testa and sinuous anticlinal cell walls. FIGURE 24. Species of Geschollia Speta. 1. Geschollia anomala (Baker) Speta from Baker (1870a : t. 178, as Ornithogalum anomalum Baker ); 2. Geschollia anomala (Baker) Speta from Dyer (1951 : t. 1117, as Drimia anomala (Baker) Baker ); 3. Geschollia calcarata (Baker) Mart. -Azorín et al. ; lectotype of Ornithogalum calcaratum Baker designated by Stedje (1987); illustration by W.H. Fitch corresponding to the collection K001291881 © Reproduced with the consent of the Royal Botanic Gardens, Kew. Number of species and distribution :— Geschollia currently includes nine species, eight sensu Martínez-Azorín et al. (2019d) together with D. loedolffiae Van Jaarsv. in Van Jaarsveld & Van Wyk (2006: 50) based on the inflorescence drying soon after flower withering, the small capsule and seed, and distribution. Species of the genus show their center of diversity in south-central South Africa from the Little Karoo in the West to the Eastern Cape in the East, with one species extending to southern Namibia and northwestern parts of the Northern Cape Province , and some disjunct populations occurring in southwestern and eastern South Africa ( Fig. 8 ). The genus is therefore restricted to the Cape , Karoo-Namib, and Uzambara-Zululand Regions (sensu Takhtajan 1986 ). For further information on Geschollia species see Baker (1870a , 1874b ), Van Jaarsveld & Van Wyk (2006) , Williamson (2012) , Manning & Goldblatt (2018) , and Martínez-Azorín et al. (2019d) . Karyology :—Apparently not studied yet (cf. Goldblatt et al. 2012 ). History, diagnostic characters, and taxonomic relationships :— Ornithogalum anomalum was described by Baker (1870a) based on a collection of a South African plant with a single, terete leaf, long raceme, shortly connate tepals reflexed at full anthesis, and spreading stamens. Baker (1870a) also pointed to apparent differences with typical Ornithogalum on which account he described O . subg. Ledebouriopsis Baker (1870a : t. 178) to accommodate this new species. Baker (1897) later placed this species in Drimia and made the first mention of the spurred condition of the bracts. Dyer (1951) noted the peculiar habit of the single terete leaf in D. anomala ( Baker 1870a : t. 178) Baker (1897: 442) , stating that it “withers from the tip, the withering process being arrested at different stages”. FIGURE 25. Species of Geschollia Speta displayed in horizontal rows of images. 1. Geschollia anomala (Baker) Speta ; 2. Geschollia brachyandra Mart. -Azorín et al. ; 3. Geschollia globuligera Mart. -Azorín et al. ; 4. Geschollia loedolffiae (van Jaarsv.) Mart. -Azorín et al. The phylogenetic analyses of Pfosser & Speta (2001 , 2004 ) included samples of Drimia anomala from South Africa , which formed an independent, well-supported clade within Urgineoideae . This led Speta (2001) to describe the monotypic Geschollia , showing a single terete leaf (rarely two leaves), a long and multiflowered racemose inflorescence, short pedicels, filiform and, patent filaments, white, pink, yellow or greenish tepals connate for ca. 1 mm at the base, and angulose seeds, 1.5‒2.0 mm long. Pfosser & Speta (2001 , 2004 ) found Geschollia to form a clade sister to Boosia and Urgineopsis , although the relationship received only weak statistical support. However, later analyses ( Pfosser et al. 2012 ) determined Geschollia and Urgineopsis to be sister, well-supported monophyletic genera, with moderate support. The lastest revision of Urgineoideae in Southern Africa by Manning & Goldblatt (2018) includes Drimia anomala in D . sect. Ledebouriopsis merging eleven species with variable morphologies. This assemblage is polyphyletic as revealed by previous phylogenetic studies ( Pfosser & Speta 2001 , 2004 ) as well as the recent phylogenetic work by Martínez-Azorín et al. (2023a) where 22 samples of Geschollia form a strongly supported clade sister to Boosia . These latter two genera share some morphological characters, such as the terete leaves or the usually elongated inflorescence, but they differ by the mostly single leaves and small capsules and seeds in Geschollia , a genus largely confined to southern central South Africa , and by the usually more numerous leaves, with usually larger, elongated, flattened seeds in Boosia — a genus centered in eastern South Africa and extending northwards through eastern Africa. Therefore, we here accept Geschollia at genus rank in the sense of Martínez-Azorín et al. (2019d) , with the addition of D. loedolffiae . All nine species share the main diagnostic characters of Geschollia , these being the single (rarely 2), terete leaf and comparatively small capsules with small polygonal or irregularly compressed, angled seeds. FIGURE 26. Species of Geschollia Speta displayed in horizontal rows of images. 1. Geschollia longipedicellata Mart. -Azorín et al. ; 2. Geschollia occultans (G.Will.) Mart. -Azorín et al. ; 3. Geschollia prolifera Mart. -Azorín et al. ; 4. Geschollia zebrina Mart. -Azorín et al. Accepted species and required new combination:— Geschollia anomala (Baker) Speta in Stapfia 75: 169 (2001) ≡ Ornithogalum anomalum Baker in Refug. Bot. [Saunders]: t. 178 (1870), basionym ≡ Drimia anomala (Baker) Baker, Fl. Cap. (Harvey) 6(3): 442 (1897) ( Figs 2.6 , 24.1, 24.2 , 25.1 ). Type :— SOUTH AFRICA . Cape of Good Hope, “sent from South Africa by Mr Thos. [Thomas] Cooper”. Illustration in Baker (1870a : Plate 178) ( Fig. 24.1 ) (holo.). Epitype (designated by Martínez-Azorín et al. 2019d ):—Fort Beaufort (3226): 22 km south of Bedford on R350, Normandale Farm, ca. 160 m SE from farmhouse, (–CC), elev. 649 m , 8 December 2018 (in flower and fruit), A.P.Dold 16047 (GRA! epi.). = Urginea eriospermoides Baker in Gard. Chron. ser. 3, 2: 126 (1887) . Type :— SOUTH AFRICA , MacOwan 292-72 (K000257355! holo.). Geschollia brachyandra Mart.- Azorín, A.P .Dold & M.B. Crespo in Phytotaxa 427(2): 94 (2019) ≡ Drimia brachyandra ( Mart. - Azorín, A.P .Dold & M.B.Crespo ) J.C.Manning & Goldblatt in Bothalia 52(1): 3 (2022) ( Figs 2.7 , 25.2 ). Type :— SOUTH AFRICA . Eastern Cape . Grahamstown (3326): 19 km from Grahamstown on Cradock road, Brakloof (now Brack Kloof ), 500 m from turn-off onto farm drive, 20 m off of road verge, (–AD), elev. 690 m , 27 November 1993 (in flower and fruit), A.P. Dold 438 (GRA! holo.). Geschollia calcarata (Baker) Mart. - Azorín, M.B.Crespo, A.P.Dold, M.Pinter & Wetschnig in Phytotaxa 427(2): 97 (2019) ≡ Ornithogalum calcaratum Baker in Gard. Chron. n.s., 1: 723 (1874), basionym. = Drimia calcarata (Baker) Stedje in Nordic J. Bot. 7(6): 663 (1987) ( Fig. 2.8 ). Type :— SOUTH AFRICA . Eastern District of Cape Colony, MacOwan s.n. Illustration by W.H. Fitch made from cultivated material sent by MacOwan to W. Saunders ( Fig. 24.3 ) (K! lecto. designated by Stedje 1987). Epitype (designated by Martínez-Azorín et al. 2019):—Somerset East (3225): Somerset East District, near Charlton Falls, upriver on ledges on rockoutcrops/low cliffs between homestead and falls, Boschberg, (–DA), elev. 1400 m , 11 December 2008 , V.R. Clark, R.J. Daniels, M. Fabricius & J. Le Roux 487 (NBG0267388! epi.). Geschollia globuligera Mart.- Azorín, A.P .Dold & M.B. Crespo in Phytotaxa 427(2): 101 (2019) ≡ Drimia globuligera (Mart.- Azorín, A.P .Dold & M.B.Crespo ) J.C.Manning & Goldblatt in Bothalia 52(1): 3 (2022) ( Figs 2.9 , 25.3 ). Type :— SOUTH AFRICA . Eastern Cape . Port Elizabeth (3325): Port Elizabeth , Thornhill , Van Stadens Wildflower Reserve , West side of reserve on flats, 300 m to the east of river gorge, (–CC), elev. 230 m , coarse sandy soil on flats, recently burnt Algoa Sandstone Fynbos , 20 January 2018 (in flower), A.P. Dold TD16029 (GRA! holo.; ABH! iso.). Geschollia loedolffiae (van Jaarsv.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia loedolffiae van Jaarsv. in Aloe 43(2- 3): 50 (2006), basionym ( Fig. 25.4 ). Type :— SOUTH AFRICA . Eastern Cape . Butterworth (3228): near the Kei River Mouth, (–CB), 6 March 2003 , Van Jaarsveld & Voigt 17914 (NBG0207731! holo.; K000524679! iso.). Geschollia longipedicellata Mart.- Azorín , Wetschnig, M .Pinter & M.B. Crespo in Phytotaxa 427(2): 103 (2019) ≡ Drimia longipedicellata (Mart.-Azorín, Wetschnig, M .Pinter & M.B.Crespo ) J.C.Manning & Goldblatt in Bothalia 52(1): 3 (2022) ( Fig. 26.1 ). Type :— SOUTH AFRICA . Eastern Cape . Willowmore (3323): Willowmore , ca. 1 km E of town, (–BC), elev. 860 m , 4 May 2015 in flower ex hort in Graz Austria, W . Wetschnig & C. Huber WW4944 (GRA! holo.; ABH! iso.). Geschollia occultans (G.Will.) Mart.- Azorín, M.B .Crespo & M. Pinter in Phytotaxa 427(2): 105 (2019) ≡ Drimia occultans G.Will. in Cact. Succ. J. (Los Angeles) 83(6): 287 (2012) , basionym ( Figs 2.10 , 26.2 ). Type :— NAMIBIA . Oranjemund (2816): Southern Namib Desert , Swartkop Hill , 9 km E of Oranjemund , (–DA), March 2011 ex hort. in Cape Town , G . Williamson 5922 (NBG0271283! holo.). Geschollia prolifera Mart.- Azorín, A.P .Dold & M.B. Crespo in Phytotaxa 427(2): 107 (2019) ≡ Drimia prolifera ( Mart. - Azorín, A.P .Dold & M.B.Crespo ) J.C.Manning & Goldblatt in Bothalia 52(1): 3 (2022) ( Figs 2.11 , 26.3 ). Type :— SOUTH AFRICA . Eastern Cape . Fort Beaufort (3226): Fort Fordyce Reserve , Fort Beaufort , East ‘lip’ of Fuller’s Hoek forest basin, below EC Parks office and lodge, (–DA), elev. 900 m , flowered ex hort 25 November 2017 , A.P. Dold 16026 (GRA! holo.). Geschollia zebrina Mart.-Azorín, A.P.Dold & M.B.Crespo in Phytotaxa 427(2): 109 (2019) ≡ Drimia zebrinella J.C.Manning & Goldblatt in Bothalia 52(1): 3 (2022) nom. nov. [non Drimia zebrina (Mart.-Azorín, N.R. Crouch & M.B.Crespo) J.C.Manning & Goldblatt in Bothalia 49(1): 5 (2019) ] ( Fig. 26.4 ). Type :— SOUTH AFRICA . Western Cape . Oudtshoorn (3322): Grootkop Nature Reserve, NE of Oudtshoorn, (–CA), elev. 415 m , 4 May 2015 in flower ex hort in Graz, Austria, M. Martínez-Azorín, J. Vlok, A.P. Dold & A. Martínez-Soler MMA893 (GRA! holo.; ABH! iso.).