Abstract

Background and Aims

The taxonomic arrangement within subfamily Ornithogaloideae (Hyacinthaceae) has been a matter of controversy in recent decades: several new taxonomic treatments have been proposed, based exclusively on plastid DNA sequences, and these have resulted in classifications which are to a great extent contradictory. Some authors have recognized only a single genus Ornithogalum for the whole subfamily, including 250–300 species of variable morphology, whereas others have recognized many genera. In the latter case, the genera are inevitably much smaller and they are better defined morphologically. However, some are not monophyletic as circumscribed.

Methods

Phylogenetic analyses of Ornithogaloideae were based on nucleotide sequences of four plastid regions (trnL intron, trnL-F spacer, rbcL and matK) and a nuclear region (ITS). Eighty species covering all relevant taxonomic groups previously recognized in the subfamily were sampled. Parsimony and Bayesian analyses were performed. The molecular data were compared with a matrix of 34 morphological characters.

Key Results

Combinations of plastid and nuclear data yielded phylogenetic trees which are better resolved than those obtained with any plastid region alone or plastid regions in combination. Three main clades are found, corresponding to the previously recognized tribes Albuceae, Dipcadieae and Ornithogaleae. In these, up to 19 clades are described which are definable by morphology and biogeography. These mostly correspond to previously described taxa, though some need recircumscription. Morphological characters are assessed for their diagnostic value for taxonomy in the subfamily.

Conclusions

On the basis of the phylogenetic analyses, 19 monophyletic genera are accepted within Ornithogaloideae: Albuca, Avonsera, Battandiera, Cathissa, Coilonox, Dipcadi, Eliokarmos, Elsiea, Ethesia, Galtonia, Honorius, Loncomelos, Melomphis, Neopatersonia, Nicipe, Ornithogalum, Pseudogaltonia, Stellarioides and Trimelopter. Each of these has a particular syndrome of morphological characters. As a result, 105 new combinations are made and two new names are proposed to accommodate the taxa studied in the new arrangement. A short morphological diagnosis, synonymy, details of distribution and an identification key are presented.

INTRODUCTION

Hyacinthaceae include about 700–900 species of bulbous plants which are mainly distributed through Europe, Africa and south-west Asia, with a single small genus in South America (APG II, 2002). Following the molecular studies of Pfosser and Speta (1999) and Manning et al. (2004), four monophyletic subfamilies can be clearly recognized within Hyacinthaceae: Hyacinthoideae, Ornithogaloideae, Urgineoideae and Oziroëoideae. Alternatively, Hyacinthaceae can be treated as subfamily Scilloideae of Asparagaceae, and the subfamilies above are then treated as tribes Hyacintheae, Ornithogaleae, Oziroëeae and Urgineeae (e.g. APG III, 2009; Chase et al., 2009). Hyacinthaceae are monophyletic within Asparagaceae (sensu lato), and here the former system is followed (although M.F.F. prefers the latter).

Ornithogaloideae are distributed through Europe, south-west Asia and Africa and include about 280 species (Speta, 1998a). The definition of genera and the assignment of species to genera within this subfamily have troubled taxonomists since Linnaeus (Stedje, 2001a, b), and there has been a great controversy concerning generic treatments. At least three taxonomic arrangements at generic level have been proposed within Ornithogaloideae in recent times (Speta, 1998a; Manning et al., 2004, 2009a), all based exclusively on plastid DNA regions. Furthermore, those proposals are to a great extent contradictory.

Speta (1998a) in a study of Ornithogaloideae recognized 13 genera: Stellarioides Medik., Coilonox Raf., Albuca L., Pseudogaltonia Kuntze, Dipcadi Medik., Galtonia Decne., Zahariadia Speta, Melomphis Raf., Cathissa Salisb., Eliokarmos Raf., Loncomelos Raf., Honorius Gray and Ornithogalum L. He also considered Neopatersonia Schönland as a possible member of the subfamily. However, Pfosser and Speta (1999) in a phylogenetic analysis based on 30 species and one plastid DNA region (trnL-F) showed that some of the genera recognized by Speta (1998a) were polyphyletic.

Manning et al. (2004) extended the phylogenetic study to 40 taxa within Ornithogaloideae and a second plastid DNA region (rbcL) was added. These authors opted for accepting just a single genus, Ornithogalum, in Ornithogaloideae, thus expanding Ornithogalum to include Albuca, Dipcadi, Galtonia, Neopatersonia and Pseudogaltonia, together with the other genera accepted by Speta (1998a). This treatment was justified by the complete collapse at the base of the phylogenetic tree, and the apparent lack of monophyly of these genera as previously circumscribed.

This recognition of Ornithogalum (sensu lato) proved to be somewhat unpopular (e.g. Van Jaarsveld et al., 2007; Manning et al., 2009a), and recently Manning et al. (2009a) continued the study of the generic circumscription in Ornithogaloideae. They added a third plastid region (matK) and extended their previous study to 70 species (Manning et al., 2009,a), obtaining higher resolution by combining sequences from the different plastid regions, as previously suggested for other groups by for instance Barfuss et al. (2005). Manning et al. (2009a) obtained a combined tree with four strongly supported main clades. They discussed extensively three possible taxonomic arrangements in Ornithogaloideae on the basis of their plastid molecular data, and finally favoured three tribes (Albuceae, Dipcadieae and Ornithogaleae), with only four genera (Dipcadi, Pseudogaltonia, Ornithogalum and Albuca). Among these, Ornithogalum and Albuca were still defined in a broad sense and, as a result, are highly heterogeneous in morphology. Both genera were, however, divided into many subgenera, sections and series, some of which were monophyletic and well characterized from a morphological point of view.

Plastid sequences have constituted an important source of characters for the phylogenetic reconstruction in plant groups (Palmer et al., 1988; Clegg and Zurawski, 1992; Cameron, 2004). However, due to the cytoplasmic inheritance of that DNA, it can have a tendency to show significant errors due to hybridization, introgression or irregular distribution of lineages (Rieseberg and Soltis, 1991; Doyle, 1992). Fortunately, these problems may to some degree be identified by comparing phylogenetic trees from plastid and nuclear DNA sequences (Smith and Sytsma, 1990; Rieseberg, 1991; Wendel et al., 1991; Peterson et al., 2004, 2009; Alzate et al., 2009; Zarrei et al., 2009), and by checking the relationships among the species (Sytsma and Schaal, 1985; Rieseberg et al., 1988; Wallace and Jansen, 1990). As the combination of trnL-F, rbcL and atpB plastid sequences has been relatively unfruitful in resolving taxonomic questions in Hyacinthaceae (Lebatha et al., 2006), the nuclear ribosomal ITS region appears to be a good candidate, though few data have been generated for this family to date (cf. Pires and Sytsma, 2002). The presence of multiple alleles has been detected (Pfosser in Lebatha et al., 2006), making sequencing difficult. Moreover, Herrmann (2002) pointed out that the ITS region is not a useful molecular marker with which to study the phylogenetics of Ornithogalum, due to difficulties in amplification.

The aims of the present contribution are: (a) to explore phylogenetic relationships in Ornithogaloideae by combining plastid (rbcL, trnL-F and matK) and nuclear (ITS) DNA sequences; (b) to evaluate generic limits and circumscriptions after consideration of molecular data, morphological traits and/or biogeographical distribution of each group obtained; and (c) to generate and justify a new taxonomic arrangement with regard to other options discussed by previous authors.

MATERIALS AND METHODS

Taxon sampling

Eighty ingroup taxa were selected, including representatives from all previously recognized taxonomic groups in Ornithogaloideae [Table 1; acronyms of herbaria follow Thiers (2009), whereas plant names and authorities follow IPNI (2010)], which cover the full range of morphological and taxonomic variation. With regard to previous studies (Manning et al., 2009a), sampling has been extended by nine taxa, mostly corresponding to Eurasian and north African representatives of Ornithogalum sensu stricto. Nearly 25 % of the total accepted species for the subfamily are included in the present study. Five taxa from other subfamilies of Hyacinthaceae were chosen as outgroups: Bowiea volubilis Harv. ex Hook.f. and Rhadamanthus convallarioides Salisb. (both Urgineoideae), Massonia echinata L.f. and Muscari neglectum Guss. ex Ten. (both Hyacinthoideae) and Oziroë biflora (Ruiz & Pav.) Speta (Oziroëoideae) (Table 1).

Table 1.

Samples included in the phylogenetic analyses of the Hyacinthaceae, mainly the subfamily Ornithogaloideae

DNA bank code Species Locality Voucher 
K 1784 Albuca abyssinica Jacq. South Africa, 20 km E of Nelspruit Manning 2687 (NBG), Chase 1784 (K) 
LH 504 Albuca amboensis (Schinz) Oberm. Botswana Beaumont 568 (NBG) 
K 1845 Albuca aff. chaetopoda Chiov. ex hort, Kew 1981–1086 Chase 1845 (K) 
K 24814 Albuca cooperi Baker South Africa, Namaqualand, Knersvlakte, Douse-the-Glim Chase 24814 (K) 
K 8252 Albuca fragrans Jacq. South Africa, Darling Manning 2184 (NBG) 
K 12563 Albuca glandulosa Baker Without locality Goldblatt & Manning 11528 (NBG) 
K 1776 Albuca pendula B.Mathew ex hort., Kew 1991–846 Chase 1776 (K) 
K 8253 Albuca rupestris Hilliard & B.L.Burtt South Africa, KwaZulu-Natal, near Bulmer Not found (K) 
K 12514 Albuca setosa Jacq. South Africa, Kaapmuiden Manning 2689 (NBG) 
K 1012 Albuca shawii Baker ex hort., Kew 1983–8300 Chase 1012 (K) 
K 1736 Albuca tortuosa Baker ex hort., Kew 1993–1849 Chase 1736 (K) 
K 1843 Albuca wakefieldii Baker ex hort., Kew 1977–5233 Chase 1843 (K) 
LH s.n. Avonsera convallarioides (H.Perrier) Speta Madagascar, Ankarafantsika Plateau Lavranos s.n. (NBG) 
ABH 416 Battandiera amoena (Batt.) Maire Morocco, Erfoud, antes del puesto de fósiles SALA 135875 
LH 1493 K 26930 Battandiera pulchra (Schinz) Mart.-Azorín, M.B.Crespo & Juan Namibia, Windhoek Mannheimer CM2506 (NBG, WIND) 
LH 819 Battandiera seineri (Engl. & Krause) Mart.-Azorín, M.B.Crespo & Juan South Africa, Kgalaghadi Transfrontier Park Snijman s.n. (no voucher) 
LH 138 K 28300 Battandiera stapffii (Schinz) Mart.-Azorín, M.B.Crespo & Juan Namibia, Farm Keitzaub Mannheimer 1747 (WIND) 
K 176 Bowiea volubilis Harv. ex Hook.f. Without locality Chase 176 (NCU, K) 
ABH 233 Cathissa cocinna (Salisb.) Speta Spain, Zamora, Torregamones ABH 47140 
ABH 206 K 20985 Cathissa reverchonii (Lange ex Willk.) Speta Spain, Cádiz, Grazalema, Tajo de los Pajaritos ABH 47138 
ABH 208 K 20987 Cathissa unifolia (L.) Mart.-Azorín, M.B.Crespo & Juan Spain, Huelva, La Nava ABH 50131 
K 12561 Coilonox concordianum (Baker) Speta South Africa, Loeriesfontein Goldblatt, Manning & Savolainen 11527 (MO, NBG) 
K 12520 Coilonox polyphyllum (Jacq.) Speta South Africa, Namaqualand, Grootvlei Goldblat, Manning & Savolainen 11510 (MO, NBG) 
LH 626 Coilonox secundum (Jacq.) Speta South Africa Manning s.n. (no voucher) 
K 12499 Coilonox suaveolens (Jacq.) Speta South Africa, Anenous Flats Goldblat, Manning & Savolainen 11493 (NBG) 
K 1783 Dipcadi filifolium Baker Yemen, North Yemen, NW side of Jebel Thallanlan, above Hadia Raymah Chase 1783 (K) 
LH 197 K 26640 Dipcadi glaucum (Burch. ex Kew Gawl.) Baker South Africa, 42 km NE from Strydenburg Manning 2237 (NBG) 
LH 345 K 26740 Dipcadi gracillimum Baker South Africa, 20 km N from Amsterdam Manning 2682 (NBG) 
K 16972 Dipcadi longifolium (Ker Gawl.) Baker Malawi, Mulanje Chase 16972 (K) 
K 1824 Dipcadi serotinum (L.) Medik. Portugal, between Ourique and Castro Verde Chase 1824 (K) 
K 8254 Dipcadi viride (L.) Moench South Africa, Inchanga Manning s.n. (no voucher) 
LH 195 Eliokarmos bicornutus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Ratelpoort N de Springbok Goldblatt & Manning 11503 (NBG) 
LH 3043 Eliokarmos ceresianus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Ceres ABH 49975 
ABH 267 Eliokarmos conicus (Jacq.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Cape Peninsula, Camps Bay Manning 2993 (NBG) 
LH 375 K 26763 Eliokarmos constrictus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Addo Manning s.n. (NBG) 
LH 3054 Eliokarmos corticatus (Mart.-Azorín) Mart.-Azorín, M.B.Crespo & Juan South Africa, Northern Cape, Sutherland, Klein Roggeveld ABH 51609; Martínez-Azorín & Manning 96 (K, MO, NBG) 
LH 3045 Eliokarmos dubius (Houtt.) Mart.-Azorín, M.B.Crespo & Juan sensu stricto South Africa, Hottentots Kloof Martínez-Azorín & Manning 6 (NBG) 
LH 3046 Eliokarmos dubius (=Ornithogalum fimbrimarginatum F.M.Leight.) South Africa, Hottentots Kloof Martínez-Azorín & Manning 8 (NBG) 
K 12467 Eliokarmos neomaculatus Mart.-Azorín, M.B.Crespo & Juan South Africa, Nieuwoudtville Manning s.n. (no voucher) 
LH 349 K 26743 Eliokarmos pilosus (Jacq.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Gordon's Bay Manning 2696 (NBG) 
LH 156 K 26605 Eliokarmos pruinosus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Loeriesfontein Goldblatt, Manning & Savolainen 11526 (NBG) 
LH s.n. Eliokarmos rupestris (L.f.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Langebaan Manning s.n. (no voucher) 
LH 3044 Eliokarmos strictus (L.Bolus) Mart.-Azorín, M.B.Crespo & Juan South Africa, Cape, near Ceres ABH 49977; Manning & Martínez-Azorín 3 (NBG) 
LH 3042 Eliokarmos thyrsoides (Jacq.) Raf. South Africa, Western Cape, Elandskloof Farm ABH 50035; Martínez-Azorín & Manning 13 (NBG) 
LH 1464 K 28846 Elsiea corymbosa F.M.Leight. South Africa, Worcester, Milner Peak Helme 2850 (NBG) 
LH 1824 K 28638 Ethesia haalenbergensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan Namibia, Klinghadtsberge Mannheimer 2443 (WIND) 
K 12466 Ethesia xanthochlora (Baker) Mart.-Azorín, M.B.Crespo & Juan South Africa, foot of Anenous Pass Goldblatt, Manning & Savolainen 11491 (NBG) 
K 2024 Galtonia candicans (Baker) Decne. South Africa, ex hort. Kew 1969–19589 Chase 2024 (K) 
ABH 175 Galtonia saundersiae (Baker) Mart.-Azorín, M.B.Crespo & Juan South Africa, in culture ABH 50132 
K 17517 Galtonia viridiflora Verdoorn Lesotho, Blue Mtn. Pass Chase 17517 (K) 
ABH 381 Honorius nutans (L.) Gray Comercial origin, Kew Gardens Martínez-Azorín s.n. (no voucher) 
ABH 211 K 20990 Loncomelos narbonense (L.) Speta Spain, Alicante, Benissa ABH 50146 
ABH 203 K 20982 Loncomelos pyrenaicus (L.) L.D.Hrouda ex Holub Spain, Huelva, Puebla de Guzmán ABH 50135 
K 9182 Massonia echinata L.f. Without locality Goldblatt & Manning 10961 (NBG) 
ABH 214 K 20993 Melomphis arabica (L.) Raf. Spain, Cadiz, Tarifa cementery ABH 50134 
K 3540 Muscari neglectum Guss. ex. Ten. ex hort., Kew 1987–1738 Chase 3540 (K) 
K 12469 Neopatersonia namaquensis G.J.Lewis South Africa, 40 km S of Springbok Goldblatt & Manning 11369 (NBG) 
K 12519 Neopatersonia rotata (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan South Africa, Anenous Flats Manning 2275 (NBG) 
K 5639 Neopatersonia uitenhagensis Schönland South Africa, lime works near Robertson Goldblatt & Manning 10723 (MO, NBG) 
K 1863 Nicipe gracillima (R.E.Fr.) Mart.-Azorín, M.B.Crespo & Juan Kenya Chase 1863 (K) 
LH 3056 Nicipe graminifolia (Thunb.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Western Cape, Hottentots Kloof, near Ceres ABH 49981; Martínez-Azorín & Manning 7 (NBG) 
LH 377 K 26765 Nicipe juncifolia (Jacq.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Grahamstown Manning 2699 (NBG) 
LH 377 Nicipe paludosa (Baker) Mart.-Azorín, M.B. Crespo & Juan South Africa, Witzieshoek Manning 3149 (NBG) 
LH 3079 Nicipe sephtonii (Hilliard & B.L.Burtt) Mart.-Azorín, M.B.Crespo & Juan Lesotho, 25 km from Semonkong towards Roma Snijman 2036 (NBG) 
K 1682-B Ornithogalum armeniacum Baker Turkey, SW of Turkey, Mugla, Koycegiz, Sandras Dag, road from Agla to Gokceova Chase 1682.B (K) 
ABH 184 K 20963 Ornithogalum baeticum Boiss. Spain, Cádiz, Grazalema, Llanos del Republicano ABH 47137 
ABH 187 K 20966 Ornithogalum bourgaeanum Jord. & Fourr. Spain, Alicante, Sa Aitana, Font del Arbre ABH 47142 
K 20637 Ornithogalum chionophilum Holmboe Cyprus, Troodos Mts. Not found (K) 
ABH 387 Ornithogalum kochii Parl. Slovenia, Istria, Lipica Peruzzi s.n. (no voucher) 
K 21432 Ornithogalum lanceolatum Labill. Lebanon, road from Beirut to Zamble Dab Not found (K) 
ABH 228 K 31122 Ornithogalum montanum Ten. Turkye, División II, Estanbul, Prince Islands ABH 50139 
ABH 383 Ornithogalum monticolum Jord. & Fourr. France, Massif Central, Coiran Peruzzi s.n. (no voucher) 
K 20631 Ornithogalum oligophyllum E.D.Clarke Greece, Mount Vermion, near to Naussa Chase 28108 (K) 
ABH 393 Ornithogalum orthophyllum Ten. Italy, Basilicata, Baltarno Peruzzi s.n. (no voucher) 
K 20634 Ornithogalum pedicellare Boiss. & Kotschy Cyprus, Area 4, Ayos Antonios, approx. 1·6 km. E of Sotira No voucher 
K 20632 Ornithogalum refractum Willd. Yugoslavia, Macedonia, Mount Sinjaciko Chase 20632 (K) 
K 20636 Ornithogalum sintenisi Freyn Iran, Elburz Mts. E of Gorgan 13 km S of Aliabad towards Qalieh Now Chase 20636 (K) 
K 793 Oziroë biflora (Ruiz & Pav.) Speta ex hort., Kew 1977–2343 Chase 793 (K) 
K 2255 Pseudogaltonia clavata (Mast.) Phillips Namibia, 9 miles E of Windhoek Codd sub Chase 2255 (K) 
K 5951 Rhadamanthus convallarioides Salisb. Without locality Goldblatt 10852 A 
K 1507 Stellarioides longebracteata (Jacq.) Speta South Africa, ex hort, Kew, 1994–2325 Chase 1507 (K) 
K 1846 Stellarioides sordida (Baker) Speta ex hort Kew 1981–1010 Chase 1846 (K) 
LH 344 K 26739 Stellarioides tenuifolia (Redoute) Speta South Africa, Drummond Manning 2667 (NBG) 
LH 196 K 12516 Trimelopter etesiogaripense (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan Namibia, Spitzkop to Rosh Pinah Goldblatt & Manning 11352 (NBG) 
LH 134 Trimelopter unifolium (Retz.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Pieter Meintjies Siding Manning 2270b (NBG) 
DNA bank code Species Locality Voucher 
K 1784 Albuca abyssinica Jacq. South Africa, 20 km E of Nelspruit Manning 2687 (NBG), Chase 1784 (K) 
LH 504 Albuca amboensis (Schinz) Oberm. Botswana Beaumont 568 (NBG) 
K 1845 Albuca aff. chaetopoda Chiov. ex hort, Kew 1981–1086 Chase 1845 (K) 
K 24814 Albuca cooperi Baker South Africa, Namaqualand, Knersvlakte, Douse-the-Glim Chase 24814 (K) 
K 8252 Albuca fragrans Jacq. South Africa, Darling Manning 2184 (NBG) 
K 12563 Albuca glandulosa Baker Without locality Goldblatt & Manning 11528 (NBG) 
K 1776 Albuca pendula B.Mathew ex hort., Kew 1991–846 Chase 1776 (K) 
K 8253 Albuca rupestris Hilliard & B.L.Burtt South Africa, KwaZulu-Natal, near Bulmer Not found (K) 
K 12514 Albuca setosa Jacq. South Africa, Kaapmuiden Manning 2689 (NBG) 
K 1012 Albuca shawii Baker ex hort., Kew 1983–8300 Chase 1012 (K) 
K 1736 Albuca tortuosa Baker ex hort., Kew 1993–1849 Chase 1736 (K) 
K 1843 Albuca wakefieldii Baker ex hort., Kew 1977–5233 Chase 1843 (K) 
LH s.n. Avonsera convallarioides (H.Perrier) Speta Madagascar, Ankarafantsika Plateau Lavranos s.n. (NBG) 
ABH 416 Battandiera amoena (Batt.) Maire Morocco, Erfoud, antes del puesto de fósiles SALA 135875 
LH 1493 K 26930 Battandiera pulchra (Schinz) Mart.-Azorín, M.B.Crespo & Juan Namibia, Windhoek Mannheimer CM2506 (NBG, WIND) 
LH 819 Battandiera seineri (Engl. & Krause) Mart.-Azorín, M.B.Crespo & Juan South Africa, Kgalaghadi Transfrontier Park Snijman s.n. (no voucher) 
LH 138 K 28300 Battandiera stapffii (Schinz) Mart.-Azorín, M.B.Crespo & Juan Namibia, Farm Keitzaub Mannheimer 1747 (WIND) 
K 176 Bowiea volubilis Harv. ex Hook.f. Without locality Chase 176 (NCU, K) 
ABH 233 Cathissa cocinna (Salisb.) Speta Spain, Zamora, Torregamones ABH 47140 
ABH 206 K 20985 Cathissa reverchonii (Lange ex Willk.) Speta Spain, Cádiz, Grazalema, Tajo de los Pajaritos ABH 47138 
ABH 208 K 20987 Cathissa unifolia (L.) Mart.-Azorín, M.B.Crespo & Juan Spain, Huelva, La Nava ABH 50131 
K 12561 Coilonox concordianum (Baker) Speta South Africa, Loeriesfontein Goldblatt, Manning & Savolainen 11527 (MO, NBG) 
K 12520 Coilonox polyphyllum (Jacq.) Speta South Africa, Namaqualand, Grootvlei Goldblat, Manning & Savolainen 11510 (MO, NBG) 
LH 626 Coilonox secundum (Jacq.) Speta South Africa Manning s.n. (no voucher) 
K 12499 Coilonox suaveolens (Jacq.) Speta South Africa, Anenous Flats Goldblat, Manning & Savolainen 11493 (NBG) 
K 1783 Dipcadi filifolium Baker Yemen, North Yemen, NW side of Jebel Thallanlan, above Hadia Raymah Chase 1783 (K) 
LH 197 K 26640 Dipcadi glaucum (Burch. ex Kew Gawl.) Baker South Africa, 42 km NE from Strydenburg Manning 2237 (NBG) 
LH 345 K 26740 Dipcadi gracillimum Baker South Africa, 20 km N from Amsterdam Manning 2682 (NBG) 
K 16972 Dipcadi longifolium (Ker Gawl.) Baker Malawi, Mulanje Chase 16972 (K) 
K 1824 Dipcadi serotinum (L.) Medik. Portugal, between Ourique and Castro Verde Chase 1824 (K) 
K 8254 Dipcadi viride (L.) Moench South Africa, Inchanga Manning s.n. (no voucher) 
LH 195 Eliokarmos bicornutus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Ratelpoort N de Springbok Goldblatt & Manning 11503 (NBG) 
LH 3043 Eliokarmos ceresianus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Ceres ABH 49975 
ABH 267 Eliokarmos conicus (Jacq.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Cape Peninsula, Camps Bay Manning 2993 (NBG) 
LH 375 K 26763 Eliokarmos constrictus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Addo Manning s.n. (NBG) 
LH 3054 Eliokarmos corticatus (Mart.-Azorín) Mart.-Azorín, M.B.Crespo & Juan South Africa, Northern Cape, Sutherland, Klein Roggeveld ABH 51609; Martínez-Azorín & Manning 96 (K, MO, NBG) 
LH 3045 Eliokarmos dubius (Houtt.) Mart.-Azorín, M.B.Crespo & Juan sensu stricto South Africa, Hottentots Kloof Martínez-Azorín & Manning 6 (NBG) 
LH 3046 Eliokarmos dubius (=Ornithogalum fimbrimarginatum F.M.Leight.) South Africa, Hottentots Kloof Martínez-Azorín & Manning 8 (NBG) 
K 12467 Eliokarmos neomaculatus Mart.-Azorín, M.B.Crespo & Juan South Africa, Nieuwoudtville Manning s.n. (no voucher) 
LH 349 K 26743 Eliokarmos pilosus (Jacq.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Gordon's Bay Manning 2696 (NBG) 
LH 156 K 26605 Eliokarmos pruinosus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Loeriesfontein Goldblatt, Manning & Savolainen 11526 (NBG) 
LH s.n. Eliokarmos rupestris (L.f.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Langebaan Manning s.n. (no voucher) 
LH 3044 Eliokarmos strictus (L.Bolus) Mart.-Azorín, M.B.Crespo & Juan South Africa, Cape, near Ceres ABH 49977; Manning & Martínez-Azorín 3 (NBG) 
LH 3042 Eliokarmos thyrsoides (Jacq.) Raf. South Africa, Western Cape, Elandskloof Farm ABH 50035; Martínez-Azorín & Manning 13 (NBG) 
LH 1464 K 28846 Elsiea corymbosa F.M.Leight. South Africa, Worcester, Milner Peak Helme 2850 (NBG) 
LH 1824 K 28638 Ethesia haalenbergensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan Namibia, Klinghadtsberge Mannheimer 2443 (WIND) 
K 12466 Ethesia xanthochlora (Baker) Mart.-Azorín, M.B.Crespo & Juan South Africa, foot of Anenous Pass Goldblatt, Manning & Savolainen 11491 (NBG) 
K 2024 Galtonia candicans (Baker) Decne. South Africa, ex hort. Kew 1969–19589 Chase 2024 (K) 
ABH 175 Galtonia saundersiae (Baker) Mart.-Azorín, M.B.Crespo & Juan South Africa, in culture ABH 50132 
K 17517 Galtonia viridiflora Verdoorn Lesotho, Blue Mtn. Pass Chase 17517 (K) 
ABH 381 Honorius nutans (L.) Gray Comercial origin, Kew Gardens Martínez-Azorín s.n. (no voucher) 
ABH 211 K 20990 Loncomelos narbonense (L.) Speta Spain, Alicante, Benissa ABH 50146 
ABH 203 K 20982 Loncomelos pyrenaicus (L.) L.D.Hrouda ex Holub Spain, Huelva, Puebla de Guzmán ABH 50135 
K 9182 Massonia echinata L.f. Without locality Goldblatt & Manning 10961 (NBG) 
ABH 214 K 20993 Melomphis arabica (L.) Raf. Spain, Cadiz, Tarifa cementery ABH 50134 
K 3540 Muscari neglectum Guss. ex. Ten. ex hort., Kew 1987–1738 Chase 3540 (K) 
K 12469 Neopatersonia namaquensis G.J.Lewis South Africa, 40 km S of Springbok Goldblatt & Manning 11369 (NBG) 
K 12519 Neopatersonia rotata (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan South Africa, Anenous Flats Manning 2275 (NBG) 
K 5639 Neopatersonia uitenhagensis Schönland South Africa, lime works near Robertson Goldblatt & Manning 10723 (MO, NBG) 
K 1863 Nicipe gracillima (R.E.Fr.) Mart.-Azorín, M.B.Crespo & Juan Kenya Chase 1863 (K) 
LH 3056 Nicipe graminifolia (Thunb.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Western Cape, Hottentots Kloof, near Ceres ABH 49981; Martínez-Azorín & Manning 7 (NBG) 
LH 377 K 26765 Nicipe juncifolia (Jacq.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Grahamstown Manning 2699 (NBG) 
LH 377 Nicipe paludosa (Baker) Mart.-Azorín, M.B. Crespo & Juan South Africa, Witzieshoek Manning 3149 (NBG) 
LH 3079 Nicipe sephtonii (Hilliard & B.L.Burtt) Mart.-Azorín, M.B.Crespo & Juan Lesotho, 25 km from Semonkong towards Roma Snijman 2036 (NBG) 
K 1682-B Ornithogalum armeniacum Baker Turkey, SW of Turkey, Mugla, Koycegiz, Sandras Dag, road from Agla to Gokceova Chase 1682.B (K) 
ABH 184 K 20963 Ornithogalum baeticum Boiss. Spain, Cádiz, Grazalema, Llanos del Republicano ABH 47137 
ABH 187 K 20966 Ornithogalum bourgaeanum Jord. & Fourr. Spain, Alicante, Sa Aitana, Font del Arbre ABH 47142 
K 20637 Ornithogalum chionophilum Holmboe Cyprus, Troodos Mts. Not found (K) 
ABH 387 Ornithogalum kochii Parl. Slovenia, Istria, Lipica Peruzzi s.n. (no voucher) 
K 21432 Ornithogalum lanceolatum Labill. Lebanon, road from Beirut to Zamble Dab Not found (K) 
ABH 228 K 31122 Ornithogalum montanum Ten. Turkye, División II, Estanbul, Prince Islands ABH 50139 
ABH 383 Ornithogalum monticolum Jord. & Fourr. France, Massif Central, Coiran Peruzzi s.n. (no voucher) 
K 20631 Ornithogalum oligophyllum E.D.Clarke Greece, Mount Vermion, near to Naussa Chase 28108 (K) 
ABH 393 Ornithogalum orthophyllum Ten. Italy, Basilicata, Baltarno Peruzzi s.n. (no voucher) 
K 20634 Ornithogalum pedicellare Boiss. & Kotschy Cyprus, Area 4, Ayos Antonios, approx. 1·6 km. E of Sotira No voucher 
K 20632 Ornithogalum refractum Willd. Yugoslavia, Macedonia, Mount Sinjaciko Chase 20632 (K) 
K 20636 Ornithogalum sintenisi Freyn Iran, Elburz Mts. E of Gorgan 13 km S of Aliabad towards Qalieh Now Chase 20636 (K) 
K 793 Oziroë biflora (Ruiz & Pav.) Speta ex hort., Kew 1977–2343 Chase 793 (K) 
K 2255 Pseudogaltonia clavata (Mast.) Phillips Namibia, 9 miles E of Windhoek Codd sub Chase 2255 (K) 
K 5951 Rhadamanthus convallarioides Salisb. Without locality Goldblatt 10852 A 
K 1507 Stellarioides longebracteata (Jacq.) Speta South Africa, ex hort, Kew, 1994–2325 Chase 1507 (K) 
K 1846 Stellarioides sordida (Baker) Speta ex hort Kew 1981–1010 Chase 1846 (K) 
LH 344 K 26739 Stellarioides tenuifolia (Redoute) Speta South Africa, Drummond Manning 2667 (NBG) 
LH 196 K 12516 Trimelopter etesiogaripense (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan Namibia, Spitzkop to Rosh Pinah Goldblatt & Manning 11352 (NBG) 
LH 134 Trimelopter unifolium (Retz.) Mart.-Azorín, M.B.Crespo & Juan South Africa, Pieter Meintjies Siding Manning 2270b (NBG) 

Abbreviations: ABH: University of Alicante, Spain; K: Royal Botanic Gardens, Kew, UK; MO: Missouri Botanical Garden, USA; NBG: Compton Herbarium, Kirstenbosch, South Africa; SALA: University of Salamanca, Spain; WIND: National Botanical Research Institute, Windhoek, Namibia. GenBank accession numbers are given in Supplementary Data Table S3, available online.

Laboratory procedures

Total genomic DNA was extracted from fresh or silica-gel dried material using the 2× CTAB method of Doyle and Doyle (1987) with minor modifications. Total DNA was purified by MOBIO minicolums and mostly kept in 0·1× TE buffer (10 mm Tris–HCl, 1 mm EDTA, pH 8·0). Some samples were purified on CsCl2/ethidium bromide (1·55 g mL−1) density gradients (Creeth and Denborough, 1970).

Some samples, especially those from southern Africa, were obtained from the DNA banks of the Royal Botanic Gardens, Kew (www.rbgkew.org.uk/data/dnaBank/homepage.html) and National Botanic Gardens, Kirstenbosch (NBG) (www.nbi.ac.za/frames/researchfram.htm) (Table 1). The DNA for the remainder of samples was isolated and deposited at the DNA banks of Alicante University and the Royal Botanic Gardens, Kew (Table 1).

The trnL intron and trnL-F spacer (the trnL-F region; hereafter trnL-F) were amplified, mostly in one piece, using primers c and f (Taberlet et al., 1991), though some taxa required the use of the internal primers d and e (Taberlet et al., 1991). Amplification of the rbcL gene was performed using the primers 1F and 1360R, though some taxa required the use of internal primers, 427F and 724R (Lledó et al., 1998). ITS was amplified using the 18Z and AB102 primers (Douzery et al., 1999; Pires, 2000). When the sequences were not obtained in one reaction, the internal primers ITS2 and ITS3 (White et al., 1990) were also used.

A total volume of 25 µL was used in the PCR amplification process as follows: 21 µL of Abgene PCR mastermix (1·25 U Thermoprime Plus DNA polymerase, 75 mm Tris–HCl, 20 mm (NH4)2SO4, 2·5 mm MgCl2, 0·01 % Tween® 20, 0·2 mm each dNTP); 0·5 µL of bovine serum albumin (BSA; 0·4 %); 10 mm of each primer; 0·5 µL of water; and approx. 40 ng of DNA template. The ITS amplifications were done using 1·5 mm MgCl2 Abgene PCR mastermix and 1 µL of dimethyl sulphoxide (DMSO). Amplifications were performed on a 9700 GeneAmp thermocycler (Applied Biosystems), with the following programme: 4 min at 94 °C, followed by 28 cycles of 94 °C for 1 min, 48 °C for 30 s for rbcL, or 50 °C for 1 min for trnL-F, 72 °C for 1 min, and a final extension 72 °C for 7 min. For ITS, a touchdown programme was used with a initial 94 °C for 5 min, following by 12 cycles of 94 °C for 30 s, 59 °C for 1 min decreasing 1 °C in each cycle plus 18 cycles at 48 °C for 1 min, 72 °C 1 min, and a final extension 72 °C for 7 min. PCR products were purified with MOBIO minicolumns following the manufacturer's protocol. Sequencing data were obtained using ABI Prism Big Dye Terminator Cycle Sequencing Kits (Applied Biosystems) and run on an ABI 3700 Genetic Analyser according to manufacturer's protocols.

All ITS sequences are original (Table 1), whereas some of those for trnL-F and rbcL (mostly from southern African taxa) and the matK matrix were obtained from John Manning (South African National Biodiversity Institute, Kirstenbosch) and Félix Forest (Royal Botanic Gardens, Kew). The plastid (except matK) sequences of the Eurasian species are all original. All new sequences were deposited in GenBank (see Table 1).

Data analyses

Sequencer 4·1 (Gene Codes Corp., Ann Arbor, MI, USA) was used to assemble complementary strands and verify software base-calling. All DNA regions were aligned by eye in PAUP version 4·0b10 (Swofford, 2002). Maximum parsimony (MP) trees were obtained from the resulting matrices using heuristic search options, using PAUP. Searches included 1000 random addition replicates and tree–bisection–reconnection (TBR) branch swapping, with MULTREES in effect (keeping multiple most-parsimonious trees). All characters had equal weight. Gaps were treated as missing. The shortest trees held in the initial heuristic search were used as initial trees for a final heuristic analysis, using all the cited options. MP support was assessed using 1000 bootstrap (BS) replicates, TBR branch swapping, simple addition sequence and MULTREES on, but keeping only ten trees per replicate following Salamin et al. (2003). Clades with BS values of 50–74 % were described as weakly supported, 75–89 % moderately supported and 90–100 % strongly supported.

A Bayesian analysis was conducted with MrBayes 3·1 (Ronquist and Huelsenbeck, 2003). The software MrModeltest (ver. 2; Nylander, 2004) was used to determine the best model of DNA substitution for each partition, using the Akaike Information Criterion (Akaike, 1974). The Markov Chain Monte Carlo chains were run for 1 × 106 generations and sampled every 100 generations. The chains reached stationarity and the average standard deviation of the split frequencies stabilized in <250 000 generations. Thus the first 750 trees were excluded (‘burn-in’) and the remaining trees were used to compile a posterior probability (PP) distribution using a 50 % majority-rule consensus. One of these remaining trees was drawn using PAUP software. Clades with PP equal to or higher than 0·95 are regarded as strongly supported.

The MP analyses were performed using all five outgroup taxa, which in all cases were external to the fully supported, monophyletic ingroup. In the Bayesian approaches only one outgroup taxon (Bowiea volubilis) was included.

Evaluation of morphological character state distribution patterns were mapped on the Bayesian tree using MacClade version 4·08 (Maddison and Maddison, 2005) with the default settings. Thirty-four characters (Tables S1 and S2 in Supplementary data, available online) were traced using ACCTRAN optimization and assuming Fitch parsimony, equal weighting of all characters, transitions among all states equally probable, and characters as unordered. Character states were determined from literature and personal observation.

RESULTS

Plastid trees

Combination of all three plastid DNA regions (rbcL, matK and trnL-F) generated a matrix of 3382 characters. Of these, 754 were variable and 341 (45·2 %) were potentially phylogenetically informative. A total of 7100 MP trees was obtained [1243 steps, consistency index (CI) = 0·70, retention index (RI) = 0·82]. Bayesian analysis yielded trees based on a General Time Reversible + Gamma + Proportion invariant distribution (GTR + I + G) with topologies that are consistent with those retrieved with the MP analysis. These data yielded trees (not shown) fully congruent with those obtained by Manning et al. (2009a).

ITS trees

Amplification of the ITS region was successful for most species and yielded useful data for phylogenetic studies in Ornithogaloideae. However, it was not possible to sequence Pseudogaltonia, the position of which is thus unknown. The ITS matrix contained 959 characters, of which 500 were variable and 337 (67·4 %) were potentially phylogenetically informative.

A total of 5590 MP trees was obtained (1539 steps, CI = 0·52, RI = 0·76). The topology of the parsimony strict consensus tree (not shown) is similar to the Bayesian tree. Therefore, only one of the post-burn-in trees obtained from a Bayesian analysis under the GTR + I + G model is shown (Fig. 1). Bayesian PP and parsimony BS values are well correlated, the Bayesian being consistently higher (Fig. 1).

Fig. 1.

Bayesian inference analysis obtained from ITS sequences in the phylogenetic analysis of Ornithogaloideae. Phylogenetic relationships shown by one of the post-burn-in topologies. Taxa are identified following the traditional treatment of Ornithogalum and other related genera (cf. Obermeyer, 1978; Zahariadi, 1980; Müller-Doblies and Müller-Doblies, 1996). The new proposed generic circumscription of the present study is shown on the right side. Branch lengths are shown above branches, and posterior probabilities and bootstrap percentages are shown below. Arrows indicate branches that collapse in the 50 % majority rule consensus tree obtained by Bayesian analysis.

Fig. 1.

Bayesian inference analysis obtained from ITS sequences in the phylogenetic analysis of Ornithogaloideae. Phylogenetic relationships shown by one of the post-burn-in topologies. Taxa are identified following the traditional treatment of Ornithogalum and other related genera (cf. Obermeyer, 1978; Zahariadi, 1980; Müller-Doblies and Müller-Doblies, 1996). The new proposed generic circumscription of the present study is shown on the right side. Branch lengths are shown above branches, and posterior probabilities and bootstrap percentages are shown below. Arrows indicate branches that collapse in the 50 % majority rule consensus tree obtained by Bayesian analysis.

Several main clades can be distinguished (Fig. 1). One formed by Neopatersonia plus O. rotatum U. Müll.-Doblies & D. Müll.-Doblies (0·97 PP; 99 % BS) is sister to a large clade including the remainder of the subfamily (0·97 PP; 56 % BS).

Successively sister groups appear in that large group. First, there is a well-supported clade (1·00 PP; 93 % BS) that can be divided into two subclades. One (1·00 PP; 80 % BS) comprises the Dipcadi clade (0·99 PP; 76 % BS) sister to the Battandiera clade (0·89 PP; 77 % BS). The other (0·87 PP; 68 % BS) includes Stellarioides, Trimelopter, Albuca and Coilonox species, and is divided into two groups. One includes Albuca plus Coilonox (1·0 PP; 66 % BS) and shows little internal resolution, though Coilonox (O. polyphyllum Jacq.-O. suaveolens Jacq.) appears as a clade (1·00 PP; 60 % BS). The other comprises Stellarioides plus Trimelopter (0·80 PP; 53 % BS), in which the monophyly of both subclades is strongly supported (1·00 PP, 98 % BS and 1·00 PP, 100 % BS, respectively).

Secondly, a large clade not fully resolved includes two subclades. The first one includes the Madagascan endemic O. convallarioides H.Perrier (=Avonsera Speta) and a clade comprising Eliokarmos, Ethesia and Galtonia (1·00 PP; 58 % BS); however, their phylogenetic relationships are not resolved. In this latter clade, two groups are identified. One (0·95 PP; 96 % BS) includes Galtonia, O. saundersiae (= Zahariadia Speta) and two species of O. subgenus Urophyllon (sensuObermeyer, 1978): O. haalenbergense U.Müll.-Doblies & D.Müll.-Doblies and O. xanthochlorum Baker (0·95 PP; 96 % BS). The other is highly supported (1·00 PP; 100 % BS) and corresponds to Eliokarmos (= O. subgenus Aspasia groups Aspasiae and Hispidae, sensuObermeyer, 1978), with several internally well-supported groups. Finally, the second subclade includes all Eurasian and north African taxa of Ornithogalum (including Honorius, Loncomelos, Melomphis and Cathissa) plus the southern and eastern African O. subgenus Aspasia group Angustifoliati (sensuObermeyer, 1978) that corresponds to Nicipe Raf. and Elsiea F.M.Leight. This clade is strongly supported (1·00 PP; 95 % BS) though the internal relationships are not fully resolved. Only some clades are well supported, namely those corresponding to Ornithogalum plus Honorius (1·00 PP; 99 % BS), Loncomelos (1·00 PP; 97 % BS) and Nicipe (1·00 PP; 81 % BS).

Combined trees

Combination of trnL-F, rbcL, matK and ITS regions generated a total matrix with 4242 characters, of which 1198 were variable and 653 (54·5 %) were potentially phylogenetically informative. A total of 1761 MP trees was obtained (2713 steps, CI = 0·58, RI = 0·78). The topology of the parsimony strict consensus tree (not shown) is quite similar to the Bayesian consensus tree obtained under the GTR + I + G model. Phylogenetic relationships are shown by one of the post-burn-in topologies (Fig. 2). Bayesian PP and parsimony BS values are well correlated, the Bayesian being consistently higher (Fig. 2).

Fig. 2.

Bayesian inference analysis of the combination of rbcL, trnL-F, matK and ITS sequences in the phylogenetic analysis of Ornithogaloideae. Phylogenetic relationships shown by one of the post-burn-in topologies. Taxa are identified following the traditional treatment of Ornithogalum and other related genera (cf. Obermeyer, 1978; Zahariadi, 1980; Müller-Doblies and Müller-Doblies, 1996). The new proposed generic circumscription of the present study is shown on the right side. Branch lengths are shown above branches, and posterior probabilities and bootstrap percentages are shown below. Arrows indicate branches that collapse in the 50 % majority rule consensus tree obtained by Bayesian analysis.

Fig. 2.

Bayesian inference analysis of the combination of rbcL, trnL-F, matK and ITS sequences in the phylogenetic analysis of Ornithogaloideae. Phylogenetic relationships shown by one of the post-burn-in topologies. Taxa are identified following the traditional treatment of Ornithogalum and other related genera (cf. Obermeyer, 1978; Zahariadi, 1980; Müller-Doblies and Müller-Doblies, 1996). The new proposed generic circumscription of the present study is shown on the right side. Branch lengths are shown above branches, and posterior probabilities and bootstrap percentages are shown below. Arrows indicate branches that collapse in the 50 % majority rule consensus tree obtained by Bayesian analysis.

The trees in Figs 1 and 2 generally have a similar topology, but clade support is substantially higher in the latter. However, the combined tree (Fig. 2) shows two clades placed in quite different positions compared with the ITS tree (Fig. 1). Firstly, the Neopatersonia plus O. rotatum clade (1·00 PP; 100 % BS) is in the Bayesian combined tree (Fig. 2) nested within Ornithogaleae (Clade C), sister to the group including the Eurasian and North African species plus Nicipe and Elsiea (0·98 PP). That clade is, however, placed as sister to the rest of Ornithogaleae in the MP combined tree (not shown), with moderate support (88 % BS). Secondly, the Trimelopter clade (O. unifolium Retz. plus O. etesiogaripense U.Müll.-Doblies & D.Müll.-Doblies) is in the combined tree (by both MP and Bayesian analyses) sister to the clade comprising Stellarioides, Coilonox and Albuca.

Two main clades can be identified in Fig. 2. The first main clade (Clade A + B), which corresponds to Albuceae plus Dipcadieae (sensuManning et al., 2009a), receives moderate support (0·71 PP; 75 % BS) and includes the genera Pseudogaltonia, Dipcadi, Battandiera, Trimelopter, Albuca, Coilonox and Stellarioides as recognized here. It can be divided into two subclades. Pseudogaltonia and Dipcadi species form a strongly supported group (1·0 PP; 94 % BS). In the Dipcadi clade, segregation of the Mediterranean D. serotinum Medik. from the South African representatives is strongly supported (1·0 PP; 100 % BS). A second clade (1·00 PP; 65 % BS) comprises Battandiera, Trimelopter, Stellarioides, Albuca and Coilonox. Within that clade, as sister to the rest, is a well-supported group (1·0 PP; 99 % BS) including O. seineri (Engl. & K.Krause) Oberm., O. pulchrum Schinz, O. stapffii Schinz and O. amoenum Batt., which are here included in Battandiera. The next branch groups O. etesiogaripense and O. unifolium (Trimelopter clade) (1·0 PP; 100 % BS) together, sister to Albuca, Coilonox and Stellarioides (1·0 PP; 88 % BS). The Stellarioides species (O. longebracteatum Jacq., O. sordidum Baker and O. tenuifolium Redouté) constitute a fully supported subclade (1·0 PP; 100 % BS), sister to Albuca plus Coilonox (1·0 PP; 99 % BS). Although Albuca was not supported as monophyletic in the heuristic MP analysis, the Bayesian one yielded two fully supported monophyletic groups: Coilonox (1·00 PP) and Albuca (1·00 PP). In Albuca two main clades appear, one of which matches the former Pallastema Salisb (0·99 PP; 55 % BS).

The second main clade in the combined tree (Clade C), which corresponds to Ornithogaleae (sensuManning et al., 2009a), is well supported (1·0 PP; 88 % BS). However, relationships of the principal lineages are not well resolved. First, a large clade (1·0 PP; 69 % BS) comprises the species of the here-accepted genera Galtonia, Ethesia and Eliokarmos. Each of these three genera form a well-supported clade. Species of Galtonia sensu stricto plus O. saundersiae Baker form a strongly supported subclade (1·0 PP; 97 % BS), and therefore we consider all of them as members of Galtonia. This group is sister to the one formed by O. haalenbergense U.Müll.-Doblies & D.Müll.-Doblies and O. xanthochlorum Baker (1·0 PP; 100 % BS), here segregated as Ethesia. Sister to these two genera is a strongly supported subclade (1·0 PP; 100 % BS), which we accept as Eliokarmos, that includes species of O. subgenus Aspasia groups Aspasiae and Hispidae (sensuObermeyer, 1978). Moreover, in Eliokarmos two strongly supported subgroups (each one 1·0 PP; 100 % BS) are identified which show morphological peculiarities as discussed below: one comprising O. maculatum Jacq. and O. rupestre L.f., and the other the rest of the species of Eliokarmos.

The position of the Madagascan O. convallarioides (=Avonsera) is not resolved, as it constitutes a separate clade in an unresolved position in Ornithogaleae. As said before, the Neopatersonia clade falls sister to a well-supported third main clade (1·0 PP; 99 % BS) that comprises the rest of representatives from Europe, Middle East and North Africa. This latter clade includes all samples of Ornithogalum, Honorius, Loncomelos, Melomphis and Cathissa, plus some species from southern and eastern Africa belonging to Elsiea and Nicipe. Within this wide group, O. esterhuyseniae Oberm. (=Elsiea) is strongly supported (1·0 PP; 99 % BS) as sister to the rest of taxa. The species of Nicipe form a strongly supported subclade (1·0 PP; 97 % BS), sister to the rest of the Eurasian and North African taxa (0·91 PP; 61 % BS). The species of Cathissa form a clade (1·0 PP; 62 % BS). Relationships of Melomphis, which is represented here only by O. arabicum L., are not resolved. Finally, the species of Loncomelos form a strongly supported subclade (1·0 PP; 99 % BS), which is sister to Honorius plus Ornithogalum (1·0 PP; 94 % BS), with Ornithogalum being rather well supported (1·0 PP; 68 %BS).

Evaluation of morphological characters

Only some of the 34 morphological characters studied (Table S1 in Supplementary data) were shown to be important for the taxonomy of the subfamily, as commented on below.

Central longitudinal white band on leaves (character 9)

The reconstruction of character states reveals the concolour leaves as plesiomorphic in the subfamily, and the white-banded leaves are thus synapomorphic for some species of Ornithogalum (sensu stricto).

Inflorescence structure (character 11)

Most species of Ornithogaloideae have helicoidally disposed flowers, a character state that appears to be plesiomorphic. Therefore, according to our analysis, the usually unilateral inflorescence of Dipcadi is an autapomorphy of this genus, with a high discriminant taxonomic value.

Flower disposition (character 12)

In most species the flowers are patent to erect, which is the plesiomorphic state. Conversely, some species of Albuca, Dipcadi, Galtonia, Honorius and Pseudogaltonia have nodding flowers, an apomorphy which has evolved independently in several linages of the subfamily.

Main colour of tepals (disregarding the central band) (character 14, Fig. 3A)

Whereas most of the Eurasian representatives of the subfamily have white flowers, this state is an apomorphy that was acquired secondarily in many of the clades of Ornithogaloideae. Coloured flowers are thus a plesiomorphic state in the subfamily that is more common in the species from southern Africa.

Fig. 3.

Recontruction of character distribution in a post-burn-in tree obtained by a Bayesian inference, after combination of all the DNA regions studied: (A) main colour of tepals; (B) darker central band on tepals; (C) fusion of tepals and adnation of filaments to tepals; (D) seed morphology. All characters are unordered.

Fig. 3.

Recontruction of character distribution in a post-burn-in tree obtained by a Bayesian inference, after combination of all the DNA regions studied: (A) main colour of tepals; (B) darker central band on tepals; (C) fusion of tepals and adnation of filaments to tepals; (D) seed morphology. All characters are unordered.

Darker central band on tepals (character 16, Fig. 3B)

Species of clade A + B (tribes Dipcadieae plus Albuceae, sensuManning et al., 2009a) have tepals with a usually evident and well-defined wide darker central band on both sides of the tepals, rarely somewhat difused on the adaxial face. In contrast, clade C (tribe Ornithogaleae) has unmarked tepals or tepals with a central band only visible on the abaxial side, as in Ornithogalum sensu stricto or some species of Nicipe, Eliokarmos and Neopatersonia. Therefore, tepals banded on both sides are a symplesiomorphy in clade A + B, whereas in clade C, the unmarked tepals (such in Melomphis, Cathissa, Elsiea, Avonsera and some Eliokarmos and Nicipe) are plesiomorphic, though in some other clades the banded abaxial side of tepals is an apomorphy that appeared independently several times.

Inner tepal disposition (character 17)

Most representatives of the subfamily show spreading, not connivent, inner tepals as a symplesiomorphy. However, in some groups such as Albuca and Dipcadi connivent inner tepals are an apomorphy.

Fusion of tepals and filament adnation (characters 19 and 21, Fig. 3C)

The character state reconstruction shows fusion of tepals and adnation of filaments to the perigone as plesiomorphic states in the subfamily, as it occurs in the other subfamilies of Hyacinthaceae, the free tepals and filaments being apomorphic and having evolved independently in different clades. Most genera with all or some of their species with a fused perigone and adnate filaments (e.g. Pseudogaltonia, Dipcadi, Battandiera, Galtonia, Avonsera, Neopatersonia, Elsiea and Nicipe) are mostly distributed in the southern hemisphere. Conversely, the number of species with free tepals and filaments is dominant in the subfamily and are common among the Eurasian taxa, with few exceptions (e.g. Dipcadi serotinum).

Capsule section (character 27)

Trilobate cross-section of capsule (including those 6-lobed by further furrowing of individual lobes) is the apomorphic state, which has evolved independently several times in Ornithogaloideae. Capsules with rounded or trigonous section appear to be plesiomorphic. This character in combination with other reproductive features has proved to be important in the taxonomy of the genera.

Position of the dry perianth with respect to the capsule (character 29)

In some clades of Ornithogaleae, the perianth withers and remains appressed, concealing the fruit. This is an apomorphy which characterizes some genera such as Cathissa, Eliokarmos or Nicipe. The plesiomorphic state of this character is present in most of the remaining genera, the fruits being totally or mostly exposed, never hidden.

Seed morphology (character 30, Fig. 3D)

This feature appears to be useful in the classification of the subfamily. Species of clade A + B (tribes Dipcadieae plus Albuceae, sensuManning et al., 2009a) produce mostly flattened and subdiscoid seeds, this character being plesiomorphic. In clade C (tribe Ornithogaleae, sensuManning et al., 2009a), the irregularly compressed or polygonal seeds are common and also plesiomorphic, whereas globose seeds are rare and exclusive apomorphies of some genera, such as Avonsera, Honorius, Neopatersonia and Ornithogalum.

Testa morphology (character 31)

Puzzle-like testas are widespread and plesiomorphic in the subfamily, whereas those verruculose and reticulate are apomorphic. Most species of clade A + B (tribes Dipcadieae plus Albuceae, sensuManning et al., 2009a) show puzzle-like testas, excepting a few taxa of Albuca (e.g. A. echinosperma U. Müll.-Doblies) and species of Trimelopter. Similarly, this pattern is also common in taxa of clade C (tribe Ornithogaleae, sensuManning et al., 2009a), and it is the basis on which two additional secondary morphologies can arise that constitute apomorphies for some groups. On the one hand, Ornithogalum plus Honorius have a coarsely reticulate testa, in which the anticlinal walls of cells are prominent and undulate, with the central part sunken. On the other hand, some taxa of Eliokarmos, Neopatersonia, Nicipe and Cathissa produce seeds with a granulate to echinulate testa in which the basal puzzle-like pattern can sometimes also be observed.

Seed disposition (character 32)

Most species of Ornithogaloideae have biseriate or multiseriate seeds per locule of the capsule, this being the plesiomorphic state. Uniseriate seeds are present in Pseudogaltonia, Dipcadi and Battandiera, the seeds being flattened and horizontally stacked in each locule.

DISCUSSION

Recent studies by Manning et al. (2004, 2009a) and Wetschnig et al. (2007) have greatly clarified phylogenetic relationships in Ornithogaloideae. In their most recent paper, Manning et al. (2009a) found three major clades within Ornithogaloideae that they treated as three tribes and four genera: Dipcadieae (Pseudogaltonia and Dipcadi), Albuceae (Albuca) and Ornithogaleae (Ornithogalum). Furthermore, they recognized many subgenera, sections and series, mainly in Albuca and Ornithogalum. Both these genera sensuManning et al. (2009a) are extremely variable from a morphological point of view, and are difficult to define within Hyacinthaceae.

In the present study, the combination of plastid and nuclear DNA sequences resulted in well-supported phylogenetic trees (Fig. 2) in which most of the clades are congruent with monophyletic genera previously described on the basis of morphological syndromes (Table 2). The addition of ITS sequences generated trees with greater resolution and support than those based only on plastid regions. The topology of our combined trees is similar in some aspects to the one obtained by Manning et al. (2009a). The three tribes they accepted, clade A (Albuceae), clade B (Dipcadieae) and clade C (Ornithogaleae), are also present in our combined trees (Fig. 2). However, notable differences can be identified.

Table 2.

Summary of the main taxonomic treatments of Ornithogaloideae

graphic 
graphic 

Horizontal lines indicate approximate equivalences among taxa.

Dipcadieae are sister to Albuceae, not to Ornithogaleae, though with only moderate support. This relationship is founded on some morphological synapomorphies which are exclusive in the subfamily and common to most of Dipcadieae plus Albuceae, such as the coloured band usually visible on both sides of the tepals and associated with several aggregated medial nerves, or on other features shared by Dipcadi and Battandiera (tribe Albuceae) such as the wide, quadrate and apically truncate capsule and flat, discoid, horizontally stacked and uniseriate seeds. These new relationships found in the combined trees make these groups more homogeneous, reducing the number of homoplasious characters. Moreover, another important difference is found in the Albuceae. Manning et al. (2009a) accepted Coilonox as a monophyletic subgenus of Albuca (as A. subgenus Osmyne), resulting in their A. subgenus Albuca not being monophyletic. In contrast, the present results demonstrate monophyly of both groups, with complete separation fully supported in the Bayesian analysis (1·00 PP).

Another important divergence is found in the Ornithogaleae clade, which Manning et al. (2009a) regarded as the single, extremely variable genus Ornithogalum. They divided Ornithogalum into four subgenera: Avonsera, Galtonia, Aspasia and Ornithogalum. Although the first three are rather homogeneous in morphology, Ornithogalum remained extremely heterogeneous, with seven sections most of which have sometimes been treated as separate genera. First, they placed the monophyletic O. section Linaspasia (southern and eastern Africa) embedded with a low support among sections Cathissa, Melomphis, Ornithogalum and Honorius (all of them native to Eurasia and North Africa). Our tree shows the Nicipe clade as sister to the rest of Eurasian sections, a position that makes phylogenetic relationships in the group much more understandable as discussed later. Moreover, in order to make O. section Ornithogalum monophyletic, Manning et al. (2009a) expanded it to include Loncomelos (O. subgenus Beryllis), therefore constructing a morphologically heterogeneous aggregate which is difficult to define with regard to other sections. In our trees, the Loncomelos clade is sister to the monophyletic Ornithogalum, allowing easy morphological circumscription of both groups. Relationships of the remaining groups are similar to those obtained by Wetschnig et al. (2007) and Manning et al. (2009a), and will be discussed later.

Finally, Wetschnig et al. (2007) found Igidia volubilis (H.Perrier) Speta, a monotypic genus native to Madagascar, to be sister to the clade comprising taxa of Albuca, Coilonox and Stellarioides, though based exclusively on sequences of the trnL-F region. Manning et al. (2009a) included only that same sequence in their analysis (they did not obtain rbcL or matK) and Igidia was sister to the Stellarioides (sensu stricto) clade (as Albuca subgenus Urophyllon). In our analyses, no sequences of that genus were included, and hence its phylogenetic relationships to the rest of Ornithogaloideae will not be elucidated until new molecular evidence is available. Nonetheless, if one accepts that Igidia is related to the Stellarioides sensu stricto clade, it should constitute an independent genus on the basis of its remarkable morphological apomorphies (cf. Speta, 1998a, b; Wetschnig et al., 2007). Further research is currently being carried out to clarify this point (W. Wetschnig, pers. comm.).

Why the need for another new arrangement?

Following studies by Speta (1998a) and Pfosser and Speta (1999), 13–15 genera were accepted within Ornithogaloideae (Table 2). This arrangement was based exclusively on morphology and/or sequencing of a single plastid region (trnL-F). Some genera as circumscribed by these authors (e.g. Stellarioides, Eliokarmos and Neopatersonia) have, however, been demostrated to be para- or polyphyletic in recent studies (including the present one, as discussed below).

The phylogenetic studies of Hyacinthaceae by Manning et al. (2004) added the rbcL region to the previous studies with trnL-F, and improved knowledge of the phylogenetic relationships of some groups. They reported a strict consensus tree with many polytomies, notably in Ornithogaloideae, perhaps due to the low nucleotide variation in these regions (Lihová et al., 2004; Shaw et al., 2005; Pirie et al., 2007). Nonetheless, Manning et al. (2004) adopted a ‘maximally stable arrangement’, with a single genus Ornithogalum for the whole subfamily. According to Lebatha et al. (2006), the polytomies in the strict consensus trees for Hyacinthaceae show a conflict or lack of hierarchy among groups, though it would not justify their fusion. These latter authors also pointed out that morphology can be useful in elucidating phylogenetic relationships in the family.

The addition of another plastid region (matK) resulted in a better-resolved topology and in a new taxonomic arrangement (Manning et al., 2009a). Their new proposal was produced in a scenario in which three more or less contradictory taxonomies were discussed. The first (Option 1) retains as far as possible genera accepted by Speta (1998a), with the recircumscription or addition of as many genera as necessary to preserve monophyly. The second (Option 2) assumes only one genus, Ornithogalum, following Manning et al. (2004). The third (Option 3) is an intermediate, medium-conservative solution in which only four genera are accepted, corresponding to the four main lineages found in the plastid tree. All three options were extensively discussed, and finally the solution with four genera was favoured (Table 2). However, different criteria appear to be used to assign generic rank to particular clades. For instance, the sister groups Pseudogaltonia and Dipcadi are segregated as proper genera on the basis of morphological characters that are of a similar degree (or even weaker) than those separating other related clades, such as NeopatersoniaOrnithogalum, Galtonia–-Ornithogalum, BattandieraAlbuca or CoilonoxAlbuca, which are treated as subgenera or sections.

As pointed out by Manning et al. (2009a), recognition of many monophyletic genera (as favoured here), or acceptance of only four genera with many subgenera and sections (as they proposed), are both equally valid on theoretical grounds. However, Manning et al. (2009a) rejected the solution with many genera (Option 1) since it would require radical taxonomic changes including ‘the creation of six or seven new genera, and the unprecedented dismemberment of Ornithogalum’ and the need for a large number of combinations in ‘genera that are either new or have not been used in modern times’. In their opinion, all these ‘obligate’ new nomenclatural proposals would be a handicap to acceptance of such an arrangement, and hence it would not be supported by botanists (mainly South African), since it is more disruptive and appears to lack nomenclatural stability.

In their arrangement, the genera Albuca and Ornithogalum are hard to recognize, since they are broadly defined from a morphological point of view. However, they are divided in nine subgenera, 13 sections and 12 series, mostly on the basis of morphological affinities, although one (A. subgenus Albuca) is not supported as monophyletic in their trees (cf. Manning et al., 2009a).

Contrary to Stedje (2001b), the arrangement with many genera favoured here is much more consistent with morphology, and matches the molecular phylogenetic data. The new arrangement is not handicapped by any previous treatment and represents a logical circumscription of clades only when sufficient morphological evidence is combined with molecular data and monophyletic groups, as claimed by Manning et al. (2009a). Any a priori constraint (e.g. criticism by local taxonomists or apparent nomenclatural unstability) is unwarranted and counterproductive in taxonomy and systematics. Nomenclatural stability is warranted only by solid establishment of consistent taxonomic groups and a sensible application of rules in the International Code of Botanical Nomenclature (ICBN), and does not depend necessarily on taxonomic arrangements themselves. A proposed new arrangement will become stable only when it is based on a successful combination of phylogenetic, morphological and biogeographical information, and when it is comprehensive and overlooks taxonomic prejudices and/or regional botanical practices.

On the basis of these considerations, we accept 19 phylogenetically monophyletic groups, which are here treated as separate genera and which are supported by syndromes of morphological characteristics as shown below. The most informative phylogenetically characters correspond namely to the colour of, and presence of, a band on tepals, capsule shape, seed morphology and seed disposition in the capsule locules. Other characters such as perigone features, tepal fusion, disposition of leaves and bulb shape and size, are homoplasious in the subfamily. However, contrary to the arguments of Stedje (2001b) and Manning et al. (2009a), they are useful for taxonomy and do not generate inappropriate or unstable associations when they are properly combined with the other relevant features.

Homoplasy is common in most large families of monocots (e.g. Orchidaceae and Amaryllidaceae, among others), and Hyacinthaceae are no exception. Contrary to arguments of Stedje (2001b) and Manning et al. (2009a), broader generic circumscriptions make it more difficult to understand relationships among species groups, from morphological and phylogenetic points of view, since they underestimate the value of particular apomorphies. Recognition of groups which are too wide (such as Albuca and/or Ornithogalum, in Options 2 and 3 sensuManning et al., 2009a), although monophyletic, only increases heterogeneity in each and causes difficulties in morphological definition. The existence of small genera (monotypic or with two or three species) is not necessarily an inconvenience; in contrast, when small clades are well defined morphologically, molecularly and phylogenetically, they can merit generic rank, depending on the general criteria used. Other subfamilies of Hyacinthaceae (e.g. Hyacinthoideae) are treated by Manning et al. (2004) in a narrower way, with ≤11 genera being recognized. Morphological syndromes characterizing genera in Hyacinthoideae are similar to those allowing recognition of 19 genera in Ornithogaloideae. The use of two different criteria in these close subfamilies appears to be inappropriate, since homoplasy is equally common in both groups. Therefore, the arrangement favoured here (with many genera, based on clades obtained in the combined trees) is more natural and easier to understand from a global view of the family. Furthermore, as asserted by Müller-Doblies (2006) ‘a splitting of Ornithogalum seems to be by far the better taxonomic solution of the molecular data than the sinking of Albuca and other genera into Ornithogalum, which results in a huge taxonomic dustbin’.

A discussion of clades accepted at the genus rank in the taxonomic arrangement follows below. Disagreements and/or incongruence with previous, recent comprehensive treatments (Speta, 1998a; Manning et al., 2004, 2009a) are also discussed.

The Pseudogaltonia clade

For a long time, Pseudogaltonia has been accepted as including a single species, P. clavata (Mast.) Phillips, distributed throughout arid regions of north-west South Africa (Richtersveld), Namibia and Botswana. It was first described within the genus Galtonia, which grows exclusively in precipitation-rich, summer-rainfall, high-elevation regions of the Drakensberg, Low Drakensberg, Southern Berg and Natal Midlands in the eastern provinces of South Africa (KwaZulu-Natal, Free State, Mpumalanga and Eastern Cape). However, most later experts on Hyacinthaceae recognized Pseudogaltonia as a separate genus (Kuntze, 1886; Durand, 1890; Phillips, 1935; Hilliard and Burt, 1988; Speta, 1998a, b; Manning et al., 2009a, b), based on biogeography and morphological features, including the large bulb with cartilaginous and fibrous outer tunics, the racemose pyramidal inflorescence, tepals fused to form a long tube (for about three-quarters of their length) and somewhat curved and swollen at the base (autapomorphy), stamens exserted and inserted at the mouth of the perianth tube (autapomorphy), style thin and >2 cm long (autapomorphy), capsule quadrate, truncate and deeply trilobate and uniseriate seeds with five to seven angled polygonal testa cells (see Phillips, 1935; Jessop, 1975; Speta, 1998a; Manning et al., 2009a, b). Recently, an additional species, P. liliiflora J.C.Manning & Goldblatt has been described (cf. Manning et al., 2009b) from Richtersveld, which is morphologically close to the former. Our phylogenetic studies provide further evidence for P. clavata being placed far from Galtonia as sister to the Dipcadi clade, as previously pointed out (Pfosser and Speta, 1999; Manning et al., 2004, 2009a; Wetschnig et al., 2007). Accordingly, Pseudogaltonia is here accepted at the generic rank.

The Dipcadi clade

Dipcadi has been widely accepted by most researchers on Ornithogaloideae (Heywood, 1980; Valdés, 1987; Speta, 1998a, b; Manning et al., 2002, 2009a). It is characterized by two autapomorphies (the usually unilateral raceme and tepals curved outwards in the apical part) and by other features such as the tepals fused into a tube approx. one-quarter to one-third (to two-thirds) of their length, stamens adnate to the tube and included, capsule quadrate in outline, as broad as or broader than long, truncate at the apex, and seeds discoid, flattened, horizontally stacked and uniseriate in each locule with five to seven angled polygonal testa cells (cf. Obermeyer, 1964; Jessop, 1975; Heywood, 1980; Valdés, 1987; Speta, 1998a; Manning et al., 2002, 2009a). It is important to note that some species of Albuca produce almost unilateral racemes due to the decumbent growth of scapes, a pattern which is quite different from that found in Dipcadi.

All the species of Dipcadi studied in these phylogenetic analyses form a fully supported clade, sister to Pseudogaltonia. These results are identical to those obtained by Pfosser and Speta (1999) and Manning et al. (2004). The former authors recognized both genera, whereas the latter authors combined them under Ornithogalum. Recently Manning et al. (2009a) reversed their previous opinion (cf. Manning et al., 2004) and recognized the generic identity of both Pseudogaltonia and Dipcadi. Based on the data presented here and on unequivocal morphological and phylogenetic evidence, the traditional concept of Dipcadi, following Pfosser and Speta (1999), is maintained. Manning et al. (2009a) suggested two series within Dipcadi, one including species with caudate apical appendages (D. series Uropetalon), and the other including those lacking these appendages (D. series Dipcadi). However, their phylogenetic tree does not provide support for that classification, since neither series was monophyletic. Our phylogenetic results also recovered the polyphyly of these series.

The Battandiera clade

Manning et al. (2004, 2009a) showed that three species of Ornithogalum occurring in the desert regions of Namibia and South Africa (O. seineri, O. pulchrum and O. stapffii) form a well supported clade, sister to the rest of Albuceae. This result is also found in our combined analysis. However, our ITS data (Fig. 1) placed this clade as sister to Dipcadi, with moderate support. This can be explained because all these latter taxa share the same type of discoid, flat seeds, horizontally stacked and uniseriate in each locule, with angled polygonal or slightly undulate testa cells (Fig. 4I and J), a rare character in the subfamily which supports their close phylogenetic relationship. The uncertain position of this group raises questions about the circumscription of Dipcadieae (sensuManning et al., 2009a), and even the necessity for its segregation from Albuceae. In Fig. 2, these tribes are thus grouped as ‘Clade A + B’.

Fig. 4.

Morphological variation in several genera of Ornithogaloideae: (A) Honorius boucheanum (from Sturm, Deutschlands Flora in Abbildungen: tab. 33. 1796, as Ornithogalum boucheanum); (B) Nicipe juncifolia (from Jacquin, Plantarum Rariorum Horti Caesarei Schoenbrunnensis Descriptiones et Icones, 1: tab. 90. 1797, as O. juncifolium); (C) Elsiea corymbosa (from Leighton in J. S. African Bot. 10: 56. 1944); (D) Ethesia prasina (from Lindl., Bot. Reg. 2: tab. 158. 1816, as O. prasinum); (E) E. xanthochlora (from Pretoria National Herbarium PREART 0002737, as O. xanthochlorum); (F) Trimelopter unifolium (from Jacquin, Icones Plantarum Rariorum, 2(16): tab. 429. 1795, as O. fuscatum); (G and H) T. dyeri (pictures by A. P. Dold); (I) Battandiera amoena (picture by Stephen L. Jury); (J) capsule of B. amoena; (K) capsules and seeds of B. pulchra; (L) Stellarioides longebracteata; (M) S. sessiliflora; (N) Capsules and seeds of S. longebracteata; (O) capsules and seeds of S. sessiliflora.

Fig. 4.

Morphological variation in several genera of Ornithogaloideae: (A) Honorius boucheanum (from Sturm, Deutschlands Flora in Abbildungen: tab. 33. 1796, as Ornithogalum boucheanum); (B) Nicipe juncifolia (from Jacquin, Plantarum Rariorum Horti Caesarei Schoenbrunnensis Descriptiones et Icones, 1: tab. 90. 1797, as O. juncifolium); (C) Elsiea corymbosa (from Leighton in J. S. African Bot. 10: 56. 1944); (D) Ethesia prasina (from Lindl., Bot. Reg. 2: tab. 158. 1816, as O. prasinum); (E) E. xanthochlora (from Pretoria National Herbarium PREART 0002737, as O. xanthochlorum); (F) Trimelopter unifolium (from Jacquin, Icones Plantarum Rariorum, 2(16): tab. 429. 1795, as O. fuscatum); (G and H) T. dyeri (pictures by A. P. Dold); (I) Battandiera amoena (picture by Stephen L. Jury); (J) capsule of B. amoena; (K) capsules and seeds of B. pulchra; (L) Stellarioides longebracteata; (M) S. sessiliflora; (N) Capsules and seeds of S. longebracteata; (O) capsules and seeds of S. sessiliflora.

All three species in this clade are characterized by tepals with an evident longitudinal band visible on both sides, the wide capsule, quadrate and truncate at the apex, with darker transverse nerves, and the peculiar seed testa, shape and disposition (cf. Leighton, 1945; Obermeyer, 1978; Manning et al., 2009a), similar to those of Dipcadi. In our combined phylogenetic trees, and after the expansion of sampling to include more Eurasian and north African species, O. amoenum (Fig. 4I and J) was found to be deeply embedded in this clade. It is a rare and rarely collected plant which comes from the desert regions of North Africa (Morocco and Algeria), and shares the habitat, capsule and seed morphology with O. seineri, O. pulchrum and O. stapffii. These peculiar features, different from all Eurasian species of Ornithogalum, led Maire (1926) to describe the monospecific genus Battandiera to include O. amoenum. Similarly, the Namibian and South African species of this clade were segregated in the past in different taxa (Table 2). In our phylogenetic studies, the clade O. seineriO. amoenum receives strong support. This fact, together with a clear morphological and phylogenetic difference (cf. Obermeyer, 1978; Müller-Doblies and Müller-Doblies, 1996; Manning et al., 2009a) with regard to related taxa, leads us to recognize Battandiera as a separate genus, though with some necessary recircumscription. Some species of this group, including O. amoenum, O. stapffii, O. candidum Oberm., O. tubiforme (Oberm.) Oberm. and O. rautanenii Schinz, occurring mainly in the western arid regions of Africa (Namibia and Morocco–Algeria), have tepals fused in a short tube ≤5 mm long, whereas others such as O. seineri, O. donaldsonii (Rendle) Greenway and O. pulchrum have free tepals and grow mostly in the eastern regions of Africa (from western Namibia to Kenya). The existence of fused or free tepals in this genus can be explained by the former state being plesiomorphic in the subfamily and having evolved independently in several clades (Fig. 3C). The cited morphological and biogeographical differences are also supported by our phylogenetic analysis, since O. amoenum plus O. stapffii form a clade sister to the O. seineri plus O. pulchrum clade, both of which are strongly supported. Further studies, including wider sampling, are being carried out to clarify internal relationships among members of this genus.

Inclusion of Battandiera in Albuca (as A. subgenus Namibiogalum) as Manning et al. (2009a) suggested, results in a highly heterogeneous Albuca, which does not reflect the clear morphological divergence with highly informative phylogenetic characters (e.g. capsule shape, seed morphology and arrangement in the capsule). Moreover, that inclusion would not be possible on the basis of the ITS tree obtained, as stated above. Segregation of Battandiera reduces heterogeneity in genera of the subfamily (cf. Manning et al., 2009a), making Albuca more coherent. Moreover, circumscription of O. subgenus Namibiogalum sensuManning et al. (2009a), with the inclusion of O. prasinum and O. toxicarium C.Archer & R.H.Archer, is not consistent with morphology (cf. Lindley, 1816; Archer and Archer, 1999) or our combined trees. Ornithogalum prasinum and O. toxicarium are not related to one another, and are allied to other groups as shown below.

The Trimelopter clade

The Trimelopter clade includes O. unifolium and O. etesiogaripense, two species occurring in arid regions of western South Africa and southern Namibia (Sperrgebiet). These taxa are differentiated from the rest of the subfamily by the presence of a single, elliptic to narrowly oblong leaf (exceptionally two or three leaves), usually flattened against the ground, an apomorphy of the group (cf. Manning et al., 2009a); the ovaries and capsules have three prominent carpels, each with two longitudinal keels down the back and a groove between (Fig. 4G and H); and the seeds are unequally compressed or semi-discoid (cf. Dyer, 1931; Leighton, 1944; Obermeyer, 1978; Manning et al., 2009a). Due to their morphological peculiarities, these plants have been treated at various taxonomic ranks (Table 2). Rafinesque (1837) described the genus Trimelopter, including only O. fuscatum Jacq. (=O. unifolium Retz.) (Fig. 4F), which Obermeyer (1978) included (as O. unifolium) in O. subgenus Urophyllon. Müller-Doblies and Müller-Doblies (1996) described O. subgenus Urophyllon section Monarchos to accommodate 11 related species. Finally, Manning et al. (2009a) recognized section Monarchos as Albuca subgenus Monarchos. The phylogenetic study by Manning et al. (2004, 2009a) found strong support for the clade of O. unifolium and O. etesiogaripense (97–100 % BS) and revealed its sister relationship with clades including Albuca, Coilonox and Stellarioides sensu stricto. Our phylogenetic study reveals the same relationship, also with a strong support (Fig. 2). Due to the clear morphological and phylogenetic identity of this clade, the genus Trimelopter is accepted here, following Rafinesque (1837), though it is expanded to include the species of O. section Monarchos (sensuMüller-Doblies and Müller-Doblies, 1996), with the exclusion of O. rotatum because of its disruptive morphology and phylogenetic position. This latter species is evidently connected to taxa in the Neopatersonia clade, as discussed later.

The Albuca clade

The well-established genus Albuca comprises at least 60 species, as traditionally circumscribed (Speta, 1998a), though this number will probably reach approx. 100. They are readily characterized by their floral structures, which clearly differentiate them from the rest of the subfamily. Flowers are nodding, patent or erect, and the tepals are usually dimorphic: the inner ones are erect and connivent, overtopping the stamen and gynoecium, whereas the outers are erect–patent. The filaments have basal constrictions (apomorphy), the ovary has paraseptal crests, the styles are thick and prismatic (autapomorphy) or narrower and tapering, the capsule is ovoid or oblong, acute or obtuse, and the seeds are flattened, semi-lunate or discoid with smooth or rarely granulate puzzle-like testa (cf. Baker, 1897a, 1898, 1904; Jessop, 1975; Müller-Doblies, 1994, 1995; Speta, 1998a; Manning et al., 1999, 2002, 2009a). Based on a combined trnL-F plus rbcL phylogeny, Manning et al. (2004) combined all these species into Ornithogalum. When they extended the sampling of taxa and sequenced other plastid regions, they considered the traditionally accepted species of Albuca as A. subgenus Albuca, though in their tree it was not clearly monophyletic, without the inclusion of the taxa of Coilonox.

In the Bayesian analysis, the present data provide a much better resolved tree, in which the species of Albuca form a clade sister to Coilonox. This fact, together with the remarkable morphological differences between the groups, supports recognition of Albuca at the generic rank, as suggested by Müller-Doblies (2006) and adopted here. Manning et al. (2009a) suggested four sections within Albuca subgenus Albuca, which we choose not to accept here because they have not been shown to be monophyletic. In the combined trees, species of Albuca form two strongly supported groups. One of these fits the concept of the former genus Pallastema Salisb., which is well characterized by the slender, almost cylindrical style, and the slightly zygomorphic flowers with all stamen similar and fertile. The other group includes a mixture of taxa formerly referred either to subgenera Albuca, Falconera and Leptostyla (cf. Baker, 1897a; Müller-Doblies, 2006), or to sections Albuca, Falconera or Branciona (cf. Manning et al., 2009a). However, none of these receives support in our phylogenetic trees, perhaps due to the still poor sampling of all these groups. Further research is currently being carried out to clarify this point.

The Coilonox clade

Some species of Ornithogalum (e.g. O. albucoides (Ait.) Thunb., O. secundum Jacq., O. suaveolens, etc.) were segregated in the past at different ranks (Table 2). At the generic rank, Coilonox (Rafinesque, 1837) has priority. Species of Coilonox possess some similarities with Albuca that led several authors to describe species of the former in the latter (cf. Manning et al., 2009a). Although in general terms Coilonox and Albuca share the longer than wide capsules, the style longer than the ovary and stamens, and the phenology of the flowers, closing at night (cf. Manning et al., 2009a), Coilonox does not show the floral structures typical of Albuca and hence is quite simple to differentiate (cf. Manning et al., 2009a). In fact, as Müller-Doblies (2006) pointed out ‘there are no real transitions between both genera’. Our phylogenetic results demonstrate the monophyly of both groups and, together with the clear morphological differences, support recognition of both genera, which can be easily differentiated by their floral structure.

The Stellarioides clade

Ornithogalum longebracteatum, O. sordidum and O. tenuifolium are characterized by large bulbs, sometimes epigeous, with papery and greyish or fleshy and green outer tunics. They usually produce aerial or hypogeous bulbils and have long, narrow and dense cylindrical inflorescences (Fig. 4L and M), flowers small and numerous, capsule subglobose or widely ovoid to obovoid, trilobate, and seeds flattened or irregularly compressed, with sharpened edges, obliquely stalked in each locule (Fig. 4N) and a puzzle-like testa (cf. Leighton, 1945; Obermeyer, 1978; Speta, 1998a; Manning et al., 2009a). In Africa, the species of this group are mostly distributed in areas characterized by abundant, mainly summer rainfall. They also occur in regions with a bimodal autumn–summer rainfall regime (O. tenuifolium) or in semi-arid Mediterranean-climate areas, ranging from high-altitude warm-temperate regions of highveld in South Africa, through the mountains of southern and East Africa as far north as Ethiopia and Morocco–Algeria (O. sessiliflorum Desf., Fig. 4M and O). In South Africa, O. longebracteatum is usually found in dense thickets and edges of both subtropical and afromontane forests (Manning et al., 2009a). All these species have been recognized at various ranks (Table 2), and Stellarioides (Medikus, 1790) has priority at the generic level. They share a unique syndrome of morphological characters, including the dense and many-flowered inflorescences, with small flowers (tepals ≤12 mm long), almost sessile to shortly/pedicellate and small capsules ≤1·4 cm long. The present results show that O. longebracteatum, O. sordidum and O. tenuifolium form a fully supported clade, sister to the AlbucaCoilonox alliance. With the cited morphological syndrome, this justifies acceptance of Stellarioides at the generic rank. In contrast, recircumscription of Albuca to include Stellarioides as suggested by Manning et al. (2009a) greatly increases the heterogeneity of Albuca.

Speta's (1998a, 2001) concept of Stellarioides is almost coincident with that accepted here, and not with that in the tree of Manning et al. (2009a), in which several taxa (e.g. O. etesiogaripense, O. haalenbergense, O. pulchrum, O. rotatum, O. seineri, O. stapffii and O. unifolium) are treated as members of Stellarioides, supposedly following Speta (1998a). Nonetheless, the treatment of Speta (2001) is in need of a few changes: first, the exclusion of S. donaldsonii (≡Albuca donaldsonii), which is included here in Battandiera, and, second, the exclusion of Trimelopter and Ardernia Salisb. from synonymy, which are synonyms and accepted here as a different genus. It is worth mentioning here that Stedje (2001b) used some taxa of the present clade to exemplify her criticism of Speta's treatment. In her opinion O. tenuifolium Redouté, O. sessiliflorum and O. narbonense where supposedly undistinguishable from a morphological point of view (only by minor quantitative characters), but apparently they should be included in three different genera (Stellarioides, Cathissa and Loncomelos, respectively) following Speta. However, evidence is here shown that the first two species indeed belong to Stellarioides, whereas O. narbonense is a member of Loncomelos (see below), all showing clear morphological differences and being placed in clades far apart from each other.

The Galtonia clade

Four species are traditionally included in the genus Galtonia (cf. Hilliard and Burt, 1988): G. candicans Decne., G. princeps Decne., G. regalis Hilliard & B.L.Burtt and G. viridiflora Verdoorn. They are found in summer-rainfall, high-altitude regions of the Drakensberg, Low Drakensberg, Southern Berg and Natal Midlands (South Africa and Lesotho; Hilliard and Burt, 1988; PRECIS database). Galtonia is characterized by large leaves sheathing the stem, racemose inflorescence, with flowers nodding, tepals fused into a campanulate tube for approx. half of their length, filaments cylindrical and adnate, capsule lanceolate cylindric and acute, and seeds irregularly compressed or flattened with smooth puzzle-like testa (cf. Verdoorn, 1955; Jessop, 1975, Hilliard and Burt, 1988; Riddles and Condy, 2001; Manning et al., 2009a). Because of the unique combination of morphological features, many authors (e.g. Decaisne, 1880; Verdoorn, 1955; Hilliard and Burt, 1986, 1988; Speta, 1998a, b; Riddles and Condy, 2001) recognized Galtonia as a genus distinct from Ornithogalum. Taxa belonging to Galtonia had been also classified as Hyacinthus (Baker, 1870) or Ornithogalum subgenus Galtonia section Galtonia (Manning et al., 2009a). In the early molecular phylogenetic work of Pfosser and Speta (1999), Galtonia appeared to be polyphyletic, since G. candicans was related to Stellarioides and Albuca and G. princeps and G. viridiflora fell as sister to Ornithogalum saundersiae, a species which has been included in O. subgenus Urophyllon (Obermeyer, 1978; Müller-Doblies and Müller-Doblies, 1996) or in the monospecific genus Zahariadia Speta (nom. nud.). Manning et al. (2004, 2009a) presented evidence for the monophyly of Galtonia, with the necessary inclusion of O. saundersiae. In their studies G. candicans, G. viridiflora and O. saundersiae formed a strongly supported clade, sister to O. xanthochlorum and O. haalenbergense. First, they (Manning et al., 2004) synonymized both genera, though later they (Manning et al., 2009a) segregated Galtonia as a subgenus of Ornithogalum. Our molecular data corroborate their later results and show Galtonia plus O. saundersiae to constitute a strongly supported clade, which is sister to that comprising O. xanthochlorum and O. haalenbergense. Connections between O. saundersiae and Galtonia are surprising, since O. saundersiae has a pseudocorymbose inflorescence, erect flowers and free tepals and filaments (cf. Leighton, 1945; Obermeyer, 1978), traits all different from typical Galtonia. Moreover, chromosome numbers support separation of these groups, being 2n = 16 in Galtonia (Speta, 1998a; Forrest and Jong, 2004; Wetschnig et al., 2007) and 2n = 12, 14 in O. saundersiae (Obermeyer, 1978; Speta, 1998a). However, some morphological features, including the large and sheathing leaves, fleshy tepals, acute capsules and irregularly compressed seeds and biogeography are shared. Although some plastid molecular trees (Pfosser and Speta, 1999; Wetschnig et al., 2007) showed both genera to be monophyletic, our combined results place O. saundersiae inside Galtonia. Therefore, until new molecular evidence is available, we suggest treating it as a member of Galtonia. Related to this matter, Hammet and Murray (1993) found that some tetraploid (2n = 32) cultivars of Galtonia candicans produced inflorescences up to 2 m, with erect flowers, somewhat similar to those of O. saundersiae, which support the discussion above.

The Ethesia clade

The genus Ethesia was described by Rafinesque (1837) to include O. prasinum as the sole species (Fig. 4D). Later, Obermeyer (1978) and Müller-Doblies and Müller-Doblies (1996) classified O. prasinum, O. xanthochlorum (Fig. 4E) and other related species within O. subgenus Urophyllon. Manning et al. (2009a) presented molecular phylogenetic evidence for a close relationship between O. haalenbergense, similar to O. prasinum (cf. Müller-Doblies and Müller-Doblies, 1996), and O. xanthochlorum, both forming a clade sister to Galtonia, and taxonomically interpreted as O. subgenus Galtonia section Xanthochlora. The present molecular phylogenetic study corroborates the results of Manning et al. (2009a). Both species are well differentiated from Galtonia by their smaller size; leaves lacking sheaths, sometimes not coinciding with the flowers; tepals greenish; filaments petaloid, white, showy and spreading; capsule obcordate or widely globose and truncate to retuse at the apex; and seeds compresso-angulose, ovate or discoid with one or two ribs and biseriate per locule (cf. Lindley, 1816; Leighton, 1945; Obermeyer, 1978; Manning et al., 2009a). Moreover, Galtonia and Ethesia have contrasting distribution areas, being found in summer-rainfall and winter-rainfall regions, respectively. Because of the differences in crucial morphological characters we recognize the genus Ethesia (Rafinesque, 1837), although with a minimal recircumscription. Inclusion of O. prasinum in Battandiera (=A. subgenus Namibiogalum) as proposed by Manning et al. (2009a) is not justified on the basis of the morphology discussed above (cf. Lindley, 1816). On the contrary, this species appears to be closely connected to O. haalenbergense (cf. Müller-Doblies and Müller-Doblies, 1996).

The Eliokarmos clade

Ornithogalum conicum Jacq., O. thyrsoides Jacq., O. dubium Houtt., O. maculatum and other related taxa (cf. Manning et al., 2007) form a well-characterized group, defined by their ciliate or fimbriate (rarely glabrous) leaves; membranous, ovoid, wide and petaloid bracts; large colourful flowers; concolourous, white, yellow, orange or reddish tepals, sometimes with basal or apical maculae; filaments usually expanded or winged; capsules lanceolate or fusiform, usually hidden among the dry perianth segments; and small abundant and irregularly compressed seeds (0·5–2·0 mm), with papillate to echinulate or puzzle-like testa (cf. Baker, 1897b; Leighton, 1944, 1945; Obermeyer, 1978; Manning et al., 2007). Some have great ornamental value and are widely cultivated (Obermeyer, 1978). These species have been recognized at different ranks (Table 2), with Eliokarmos, Tomoxis Raf. and Lomaresis Raf. being published first and simultaneously at the generic level (Rafinesque, 1837). Eliokarmos has been recently used by Speta (1998a) at that rank.

Species such as O. constrictum F.M.Leight., O. bicornutum F.M.Leight., O. hispidum Hornem. and other relatives produce capsules and seeds similar to those of section Aspasia, but they differ in the long sheathing and ascending leaves, with patent blades arranged at different levels and a tubular cataphyll along the stem, usually not coinciding with the flowers, bracts narrower and apiculate or aristate, flowers smaller, and tepals usually having a darker longitudinal band only apparent on the abaxial side (cf. Obermeyer, 1978). These species were included in O. subgenus Aspasia group Hispidae (cf. Obemeyer, 1978) or in O. section Hispidaspasia (cf. Müller-Doblies and Müller-Doblies, 1996).

The phylogenetic studies of Manning et al. (2004, 2009a) revealed that all these species form a strongly supported clade. In this clade, two well-supported subclades can be identified: one comprising O. maculatum and O. rupestre, and the other grouping the rest of species of sections Aspasia and Hispidaspasia. Our phylogenetic data confirm the findings of Manning et al. (2009a), which would support the independent status of Phaeocles Salisb. (Salisbury, 1866), including O. maculatum and O. rupestre, apart from the rest of section. These taxa are peculiar in their glabrous leaves, a rare character in the group. However, no other relevant differences to the rest of the related taxa seem to exist, and therefore we prefer to accept a single genus Eliokarmos (including Phaeocles) to group all taxa discussed in these paragraphs. This concept is similar to that proposed by Speta (1998a), with the exclusion of taxa belonging to O. group Angustifoliati (sensuObermeyer, 1978) that is phylogenetically distant from this group and which we recognize as the genus Nicipe. Further studies and a more extensive sampling in this group are needed to shed more light on phylogenetic relationships within this clade and on eventual recognition of subgeneric ranks.

The Avonsera clade

Ornithogalum convallarioides is endemic to Madagascar and is characterized by its wide, pseudopetiolate leaves, loosely racemose inflorescence (usually with two flowers per bract in the lower part), small bracts; tepals completely white and connate at the basal part, filaments tapering, connate and adnate to the perianth, capsules retuse and trilobate and seeds large and ovoid (cf. Speta, 1998a; Wetschnig et al., 2007). The occurrence of two flowers per bract is an autapomorphy within Ornithogaloideae, though it is shared with the genus Oziroë Raf. (Guaglianone and Arroyo-Leuenberger, 2002), the only representative of Oziroëoideae (Speta, 1998a, b). The trilobate capsules have a peculiar structure with each lobe having a longitudinal central furrow (cf. Wetschnig et al., 2007; fig. 3·5), much resembling the six-lobed capsules of Ornithogalum sensu stricto. The genus Avonsera was described by Speta (1998b) to group O. convallarioides (the type species) and Resnova lachenalioides (Baker) Van der Merwe, arguing that it was the only genus having a multinerved perigone in Hyacinthoideae (the subfamily into which it was erroneously misplaced). Molecular phylogenetic studies by Wetschnig et al. (2007) and Manning et al. (2009a) demonstrated Avonsera (sensu Speta, l.c.) to be polyphyletic. Resnova species were true Hyacinthoideae, sister to species of Ledebouria, whereas A. convallarioides nested unequivocally within Ornithogaloideae. Therefore, Manning et al. (2009a) proposed creation of subgenus Avonsera within Ornithogalum. Our phylogenetic analyses show O. convallarioides to be an isolated clade, probably sister to the Mediterranean species of Ornithogalum plus those of Elsiea and O. subgenus Aspasia group Angustifoliati (Obermeyer, 1978), though its relationships are not fully resolved or supported. These morphological, phylogenetic and biogeographical (Malagasy endemism) characters warrant recognition of the so-far monotypic Avonsera as an independent genus.

The Neopatersonia clade

Schönland (1912) described the genus Neopatersonia to include N. uitenhagensis Schönland. Later, Lewis (1952) added N. namaquensis G.J.Lewis and N. falcata G.J.Lewis, to complete the three traditionally accepted species in the genus. As usually defined, Neopatersonia is characterized by the tepals fused in a short basal tube, filaments adnate and somewhat connate at the base (autapomorphy), stigma long trilobate and somewhat reflexed, capsule lanceolate, acute, and seeds subglobose or urceolate with pitted puzzle-like or granulate testa (cf. Schönland, 1912; Lewis, 1952; Jessop, 1975). Different classifications have treated them (Table 2) as Neopatersonia (Schönland, 1912; Lewis, 1952; Speta, 1998a, b), Ornithogalum subgenus Neopatersonia (Hutchinson, 1934), O. subgenus Ornithogalum section Neopatersonia (Manning et al., 2009a), or as members of a widely defined Ornithogalum (Manning et al., 2004). Speta (1998a, b) accepted Neopatersonia as a member of Hyacinthaceae, though with doubts about its possible inclusion in Ornithogaloideae. Molecular studies (Manning et al., 2004, 2009a) revealed a close relationship between Neopatersonia and O. rotatum, a species first included in O. section Monarchos (Müller-Doblies and Müller-Doblies, 1996), and recently placed in O. section Neopatersonia, according to molecular data (Manning et al., 2009a). In this analysis Neopatersonia plus O. rotatum appear as the sister group of the clade formed by the European species of Ornithogalum plus the African Nicipe. After combining the nuclear and plastid data, our phylogenetic analyses also show clear support for Neopatersonia in the traditional sense, to which O. rotatum is sister, being both sister to other European and African clades of ornithogaloids. Relationships of O. rotatum to Neopatersonia are supported by shared floral characters (e.g. tepals connate at the base, filaments connate and adnate), and inclusion of that species into Neopatersonia is therefore justified.

The Elsiea clade

Leighton and Salter (1944) described the genus Elsiea for peculiar plants growing in permanently wet habitats, with poorly developed and subrhizomatous bulbs, connate tepals and adnate filaments (Fig. 4C). Two species were first included in the new genus: E. corymbosa F.M.Leight. (currently O. esterhuyseniae) and E. flanaganii (Baker) F.M.Leight. (currently O. paludosum Baker). However, Obermeyer (1978) found this segregation unjustified, since she was not able to corroborate the fusion of perianth segments. Müller-Doblies and Müller-Doblies (1996) placed both taxa, together with O. flexuosum (Thunb.) U.Mull.-Doblies & D.Müll.-Doblies and O. dregeanum Kunth, in series Flexuosa of O. subgenus Urophyllon section Helogalum U.Müll.-Doblies & D.Müll.-Doblies, on the basis of the poorly developed bulbs. However, O. synanthifolium F.M.Leight., which belongs to O. subgenus Aspasia group Aspasiae, also shows this type of bulb and also occurs in permanently wet habitats (cf. Manning et al., 2007). Consequently, this bulb character appears to be a morphological convergence to that peculiar environment (cf. Manning et al., 2009a), and thus it is not suitable for systematics. Our phylogenetic studies confirm this idea, since O. esterhuyseniae falls in an isolated position in its own clade, far away from O. paludosum, which is placed together with taxa of O. section Linaspasia (the Nicipe clade). These results agree with those presented by Manning et al. (2009a), who proposed O. subgenus Ornithogalum section Elsiea to include only O. esterhuyseniae, and also demonstrate that proposals by Leighton and Salter (1944) and Müller-Doblies and Müller-Doblies (1996) are not supported from a phylogenetic point of view. However, phylogenetic divergence of these groups is reasonable on the basis of morphological and biogeographical evidence. First, O. esterhuyseniae (=Elsiea corymbosa) has a corymbose inflorescence, tepals white and connate, filaments adnate, ovary obovoid and stipitate, and capsule black (cf. Obermeyer, 1978; Manning et al., 2009a), and it is restricted to a small mountainous area in south-west South Africa. Secondly, O. paludosum, O. flexuosum and O. dregeanum have long and narrow racemose to spicate inflorescences, tepals and filaments free, ovary sessile, and capsule not black coloured (cf. Obermeyer, 1978), and occur in the eastern part of south Africa, as far north as Zambia and Malawi (cf. Obermeyer, 1978; Müller-Doblies and Müller-Doblies, 1996). The isolated phylogenetic position of O. esterhuyseniae, together with its characteristic syndrome of morphological features, warrants recognition of Elsiea as a monotypic genus.

The Nicipe clade

This clade includes O. juncifolium Jacq. (Fig. 4B), O. graminifolium Thunb., O. gracillimum R.E.Fr., O. sephtonii Hilliard & B.L.Burtt. and O. paludosum, a group of taxa growing in eastern and southern Africa and forming a strongly supported clade. They have been recognized in the past at different ranks (Table 2), including Nicipe (Rafinesque, 1837), O. group Angustifoliati (Obermeyer, 1978) and O. section Linaspasia, Vaginaspasia, Nanaspasia and Helogalum (Müller-Doblies and Müller-Doblies, 1996). Some morphological features characterize this clade, such as bulbs mostly with membranous tunics, usually ending abruptly in an epigeous neck, occasionally surrounded by tubular, transversally banded and epigeous cataphylls; leaves usually occurring with flowers, numerous, all emerging almost at the same level from the bulb neck, often filiform with enrolled margins, rarely flat and broader or even coriaceous and boat-shaped; inflorescence racemose, cylindrical to subcorymbose; bracts small and aristate, auriculate and sometimes denticulate at the base; tepals white to yellowish on the adaxial side, and usually with a dark longitudinal band on the abaxial one, almost free to fused ≤ one-third; capsule trigonous-elliptical to trilobate-turbinate, usually hidden among the dry perianth segments; and seeds variable in size and irregularly compressed (cf. Leighton, 1944, 1945; Obermeyer, 1978; Müller-Doblies and Müller-Doblies, 1996; Manning et al., 2009a).

According to this evidence, we accept this group at the generic rank, with the name Nicipe. As here circumscribed, this genus includes several groups that are quite well characterized morphologically and have peculiar distribution patterns. Species from eastern South Africa to Kenya (O. sephtonii and O. gracillimum) produce turbinate capsules, trilobate in section, and form a clade in our combined tree. Ornithogalum baurii Baker and O. diphyllum Baker share a similar fruit structure, as pointed out by Manning et al. (2007). Species from the western to eastern Cape (O. graminifolium, O. juncifolium and O. paludosum) also form a monophyletic group, and they have elliptical capsules, trigonous in section with blunt angles. Related to this group, species of O. series Zebrinella, such as O. monophyllum Baker, O. anguinum F.M.Leight. ex Oberm. and O. zebrinellum U.Müll.-Doblies & D.Müll.-Doblies show characteristic tubular, transversally banded cataphylls.

Nicipe as recognized here is mostly congruent with O. group Angustifoliati (Obermeyer, 1978), though it also includes O. pilosum L.f., a species Obermeyer placed in O. group Hispidae. That concept is supported by both molecular data (M. Martínez-Azorín et al., unpubl. res.) and morphology, as this species lacks the typical vaginate, superposed and clearly distant leaves characterizing the latter group, as pointed out by Müller-Doblies and Müller-Doblies (1996). Although the traditional concept of O. group Angustifoliati included plants with narrow to filiform leaves, Nicipe as treated here includes a wide range of leaf morphology, usually associated with particular habitats. Taxa such as O. baurii, O. pilosum, O. natalense Baker and O. aloiforme Oberm. have wide, oblong-lanceolate and flat leaves, whereas others including O. paludosum and O. flexuosum have flattened to terete, somewhat succulent leaves. These species grow mostly on moist soils or high mountain environments. Ornithogalum britteniae F.M.Leight. ex Oberm. is an extremely rare plant with evergreen, distichous, coriaceous, boat-shapped leaves with fimbriate margins resembling those of some Gasteria Duval, which grows in grassy karroid dwarf shrubland (Dold, 2003). Another group of species (O. annae-ameliae U.Müll.-Doblies & D.Müll.-Doblies, O. campanulatum U.Müll.-Doblies & D.Müll.-Doblies, O. synadelphicum U.Müll.-Doblies & D.Müll.-Doblies and O. verae U.Müll.-Doblies & D.Müll.-Doblies) have peculiar floral features, such as tepals fused, and filaments adnate to perigone and/or connate into a short tube. Connation of perigone segments is a plesiomorphy which has evolved several times in different genera of the subfamily (Fig. 3C), including Pseudogaltonia, Dipcadi, Battandiera, Galtonia, Avonsera, Neopatersonia and Elsiea. Furthermore, Battandiera and Galtonia include species with either connate or almost free tepals, as in Nicipe.

From a phylogenetic point of view, Nicipe is sister to the rest of Eurasian genera of Ornithogaloideae, and it could constitute the link with the lineages from southern Africa (cf. Manning et al., 2009a), where the origin of this genus would be placed (Feinbrun, 1941). Furthermore, the Kenyan species O. gracillimum provides evidence for this (Fig. 2). Migrations across East Africa would have connected floristic regions of both hemispheres in two different ways. On the one hand, the ‘arid corridor’ or ‘arid track’ migration hypothesis (cf. Verdcourt, 1969; Thulin, 1994; Jürgens, 1997; Linder et al., 2005; Hoyo and Pedrola-Monfort, 2006) and, on the other, the high-altitude corridor (cf. Linder et al., 2005; Devos et al., 2009), which jointly would have favoured disjunctions in genera such as Colchicum L. (including Androcymbium Willd.), Euryops (Cass.) Cass., Delosperma N.E.Br., Matthiola W.T.Aiton, Albuca, Battandiera, Dipcadi, Stellarioides and Nicipe itself.

The Cathissa clade

The Mediterranean species traditionally placed in O. subgenus Cathissa (O. concinnum Salisb., O. broteroi M.Laínz and O. reverchonii Lange ex Willk.) are characterized by their leaves glabrous, lanceolate to tapering; tepals white and ovoid or obovoid; filaments lanceolate to tapering; capsule ovoid to elliptical; and seeds irregularly flattened with papillate or puzzle-like testa (Figs 5A–C and 6A–C) (cf. Zahariadi, 1980; Pastor, 1987; Moret et al., 1990; Martínez-Azorín et al., 2006, 2007). These species were placed in Cathissa (Salisbury, 1866; Speta, 1998a, b, 2001), a group treated as a genus, subgenus or section (Table 2). Our phylogenetic results confirm the monophyly of this group of three currently accepted taxa (Martínez-Azorín et al., 2007), which are well characterized morphologically and are restricted to the western part of the Iberian Peninsula and western Morocco. Acceptance of Cathissa at the generic rank is thus justified.

Fig. 5.

Inflorescence and flower features of most of the Eurasian genera of Ornithogaloideae: (A) Cathissa reverchonii; (B) C. concinna; (C) C. unifolia; (D) Melomphis arabica; (E) Loncomelos narbonense; (F) L. pyrenaicus; (G) Ornithogalum baeticum; (H) O. bourgaeanum; (I) O. divergens.

Fig. 5.

Inflorescence and flower features of most of the Eurasian genera of Ornithogaloideae: (A) Cathissa reverchonii; (B) C. concinna; (C) C. unifolia; (D) Melomphis arabica; (E) Loncomelos narbonense; (F) L. pyrenaicus; (G) Ornithogalum baeticum; (H) O. bourgaeanum; (I) O. divergens.

Fig. 6.

Capsule (lateral and section views) and seed features of most of the Eurasian genera of Ornithogaloideae: (A) Cathissa reverchonii; (B) C. concinna; (C) C. unifolia; (D) Melomphis arabica; (E) Loncomelos narbonense; (F) L. pyrenaicus; (G) Ornithogalum baeticum; (H) O. bourgaeanum; (I) O. divergens.

Fig. 6.

Capsule (lateral and section views) and seed features of most of the Eurasian genera of Ornithogaloideae: (A) Cathissa reverchonii; (B) C. concinna; (C) C. unifolia; (D) Melomphis arabica; (E) Loncomelos narbonense; (F) L. pyrenaicus; (G) Ornithogalum baeticum; (H) O. bourgaeanum; (I) O. divergens.

The Melomphis clade

In our phylogenetic analyses, O. arabicum, traditionally included in Melomphis (Rafinesque, 1837; Garbari et al., 2008), Caruelia Parl. (Parlatore, 1854), O. subgenus Caruelia (Parl.) Baker (Baker, 1872) or Myanthe Salisb. (Salisbury, 1866), appears as an independent clade, the position of which is not fully resolved as previously demonstrated by Manning et al. (2009a). Melomphis species are distributed in the Mediterranean basin eastwards, and are characterized by the corymbose to pseudocorymbose inflorescence, globose and shining black ovary, plain white tepals, globose and truncate to retuse capsule and irregularly compressed seeds with granulate testa (Figs 5D and 6D). Some authors have recently accepted the independence of this genus (Garbari et al., 2008). Although a wider sampling is needed to clarify relationships to other ornithogaloid clades, notable morphological differences exist to justify segregation of Melomphis at the generic rank. The globose and shining ovary of these species is similar to that occurring in the South African O. saundersiae (cf. Baker, 1891), here treated as Galtonia. Therefore, it can be interpreted as a morphological convergence between these groups, which are distant from each other in the phylogenetic analyses. Morphological features exist which allow easy differentiation of these groups, including tepals fleshy, reflexed and persistent for a long time in fruit, and capsule acute and ovate in O. saundersiae.

The Loncomelos clade

The species traditionally included in O. subgenus Beryllis are characterized by their concolourous leaves long and tapering; inflorescence racemose, much longer than wide; tepals white or rarely yellowish with a green longitudinal band on the abaxial side; filament expanded in the lower half; capsule trigonous with obtuse angles, ovate-lanceolate; and seeds biseriate, horizontally disposed and irregularly compressed, with papillate, rugose testa (Figs 5E and F and 6E and F) (cf. Baker, 1872; Zahariadi, 1980; Cullen, 1984; Wittmann, 1985; Feinbrun, 1986; Pastor, 1987; Moret et al., 1990; Rechinger, 1990; Speta, 1998a; Martínez-Azorín et al., 2010a). These plants have been placed in different taxa (Table 2), including the genus Loncomelos (Rafinesque, 1837). Our phylogenetic results support the monophyly of this group of plants. Ornithogalum narbonense L. and O. pyrenaicum L. form a strongly supported clade sister to Ornithogalum sensu stricto plus Honorius, which is clearly differentiated by the morphology of inflorescences, capsules and seeds. The inclusion of Loncomelos in the synonymy of Ornithogalum section Ornithogalum, as proposed by Manning et al. (2009a), generates a highly heterogeneous group mixing different capsule and seed morphology, characters that have been treated as highly informative phylogenetically. Therefore, we adopt the generic rank for Loncomelos, following the concept of Wittmann (1985) and Speta (1998a), on the basis of clear morphological and phylogenetic evidence. Similarly, segregation of Loncomelos has been accepted increasingly in recent times (cf. Speta, 1998a, b, 2000, 2006; Spies, 2004; Peruzzi et al., 2007; Ravenna, 2007; Garbari et al., 2008).

Maire (1958) included the north African O. sessiliflorum in O. subgenus Beryllis, mainly on the basis of its long and narrow inflorescence, whereas Stedje (2001b) connected it to Cathissa. However, this species shows other evident differences with regard to the rest of the Eurasian and north African taxa of both cited groups, such as the flowers with a green band visible on both sides of tepals, the subglobose capsule, and the compressed seeds obliquely stalked in each locule and with puzzle-like testa (Fig. 4M and O). All those features strongly connect it to Stellarioides, which led Speta (2001) to establish the combination S. sessiliflora (Desf.) Speta. The exclusion of O. sessiliflorum from O. subgenus Beryllis (=Loncomelos) gives homogeneity to the latter as regards to testa morphology (only granulate or rugose, never puzzle-like as reported by Moret et al., 1990).

The Honorius clade

Ornithogalum nutans L. is characterized by the racemose inflorescence; filaments tapering, with two apical lobes overtopping the anther; the capsule nodding at maturity (autapomorphy) and with six weak ribs in section and globose seeds with prominently reticulate testa (Fig. 4A) (cf. Baker, 1872; Zahariadi, 1980; Uysal et al., 2005). Due to these features, it was segregated at different ranks (Table 2), of which Honorius (Gray, 1821) has priority at the generic rank. Our phylogenetic results place O. nutans as sister to the Ornithogalum sensu stricto clade. Phylogenetic and morphological evidence justifies its segregation at the generic rank, as accepted by some recent authors (cf. Holub, 1976; Speta, 2000; Garbari et al., 2008). The recently described O. chetikianum Uysal, Ertuğrul & Dural, a Turkish taxon close to O. nutans, is said to have leaves with a longitudinal greyish band, which connects it to many representatives of Ornithogalum sensu stricto. A wider sampling of related taxa from Anatolia and the Middle East is still needed to improve our knowledge of phylogenetic relationships within Honorius.

The Ornithogalum clade

The final clade corresponds to Ornithogalum sensu stricto, which includes Eurasian and North African species well characterized by their long linear to oblong-lanceolate leaves, sometimes with a white longitudinal band on the adaxial side; inflorescence corymbose or pseudocorymbose; tepals white with a longitudinal green band only visible on the abaxial side; capsule obovate or oblong, truncate with six evident ribs in section; and seeds globose with prominently reticulate testa (Figs 5G–I and 6G–I) (cf. Baker, 1872; Cullen, 1984; Feinbrun, 1986; Moret et al., 1990; Rechinger, 1990; Speta, 1998a; Martínez-Azorín et al., 2009, 2010b). The white longitudinal band is due to an interruption of palisade tissue in the central portion of the leaf (cf. Peruzzi et al., 2007), being an apomorphy absent in the early diverging lineages of the present clade. This character is also present in a few species of Albuca.

Species in this clade represent the typical, strict concept of Ornithogalum, which includes its type, O. umbellatum L. (cf. Stearn, 1983). Baker (1872) grouped them into O. subgenus Heliocharmos, which was reduced later to O. section Heliocharmos by Engler (1888), and which Zahariadi (1965, 1980) divided in eight subgenera (cf. Cullen and Ratter, 1967) on the basis of features such as the type of seed germination (hypogeal vs. epigeal), concrescence and consistence of bulb scales, presence or absence of central band on the leaves, inflorescence structure, position of fruting pedicels, capsule shape, etc. However, our sampling is insufficient and does not cover some of those subgenera; thus further studies are necessary to evaluate their monophyly and significance. In our phylogenetic studies, all sampled taxa of this group form a clade, strongly supported in the Bayesian analysis. Furthermore, this clade is relatively derived in the subfamily, a fact that could explain the large number of species and the scant morphological differentiation among these, probably due to recent speciation.

TAXONOMIC PROPOSAL

On the basis of the evidence presented above, 19 phylogenetically monophyletic groups are accepted (Table 3) and are treated here as independent genera since they are supported by syndromes of morphological characters. No new taxa need to be described to accommodate these monophyletic groups at generic rank (cf. Manning et al., 2009a). All genera accepted here are already described and most have been used increasingly in recent times (cf. Holub, 1976; Speta, 1998a, b, 2000, 2001, 2006; Spies, 2004; Peruzzi et al., 2007; Ravenna, 2007; Garbari et al., 2008). They are well characterized morphologically, and are here circumscribed in such a way that allows easy definition and makes easier subsequent attribution of a taxon to a given genus. Similarly, most of combinations required are available, with the exception of those affecting the newly accepted or resurrected genera, given below.

Table 3.

Key for identification of taxa in Ornithogaloideae

1a. Floral bract with small bracteoles. Tepals all fused over two-thirds of their length, into a cylindrical tube, slightly swollen at the base and somewhat curved; filaments inserted at the mouth of the tube 1. Pseudogaltonia 
1b. Floral bract without bracteoles (if present, then leaves narrowed in a pseudopetiole). Tepals free or fused into a shorter, campanulate to cylindrical, straight tube; filaments free, connate or adnate to the tube 
2a. Tepals with a broad and dark longitudinal band – green to brownish, evident on both sides and usually associated with three to many aggregated medial nerves 
2b. Tepals concolour, exceptionally with dark maculae at the base or apex, or sometimes with a broad longitudinal coloured band, only visible on the abaxial side, not well defined along the medial nerve 
3a. Ovary with two prominent dorsal keels on each carpel. Plants with only one leaf (exceptionally up to three), usually flat on the ground 4. Trimelopter 
3b. Ovary without such prominent dorsal keel. Plants usually with numerous leaves, erect or spreading (exceptionally solitary, varying in morphology) 
4a. Seed obliquely stalked in each locule. Inflorescence usually subspiciform, dense, and many flowered 7. Stellarioides 
4b. Seed horizontally stalked in each locule. Inflorescence usually racemose to subcorymbose, lax, with long pedicellate flowers 
5a. Capsule widely quadrate in outline, deeply three lobed; seeds uniseriate in each locule 
5b. Capsule ovate, oblong or obovate in outline, trigonous to trilobate; seeds biseriate in each locule (exceptionally uniseriate) 
6a. Inflorescence subspicate, mostly secund, with shortly pedicellate flowers. Outer tepals curved outwards, inners long connivent to form a cylindrical tube with reflexed tips 2. Dipcadi 
6b. Inflorescence racemose, cylindrical, conical or subpyramidal, with long pedicellate flowers. Tepals all similar and spreading 3. Battandiera 
7a. Tepals usually dimorphic: the outers mucronate and the inners cucullate or with an inflexed flap at the apex; filaments usually with basal constrictions; ovary with paraseptal keels; style trigonous in section, prismatic to obtriangular or sometimes thin or tapering; stigma not capitate, usually trilobed 5. Albuca 
7b. Tepals monomorphic, all similar; filaments without basal constrictions; ovary without paraseptal keels; style circular in section, subcylindric, usually deflexed; stigma capitate 6. Coilonox 
8a. Tepals fused at the base 
8b. Tepals completely free at the base 12 
9a. Flowers nodding. Tepals fused into a long campanulate tube of approx. half of their length. Stamens hidden among the perianth lobes 8. Galtonia 
9b. Flowers patent to erect-patent. Tepals fused into a tube ≤ one-third of their length. Stamens clearly visible 10 
10a. Leaves tapering at the base and pseudopetiolate; inflorescence usually with two flowers per bract in the lower part; bracteoles present in the upper flowers; capsule retuse or truncate at the apex 11. Avonsera 
10b Leaves sheathing at the base, never pseudopetiolate; inflorescence always with one flower per bract; bracteoles absent; capsule acute to subacute at the apex 11 
11a. Tepals ≤ half the length of capsule; seeds piriform or subglobose; filaments of stamen connate 12. Neopatersonia 
11b. Tepals longer than capsule or rarely slightly shorter; seeds angular, irregularly compressed; filaments of stamen free to adnate, exceptionally connate 14. Nicipe 
12a. Ovary globose, shining black. Inflorescence subcorymbose 13 
12b. Ovary fusiform to oblong, green or yellow. Inflorescence racemose to corymbose 14 
13a. Capsule ovate, with acute apex. Tepals fleshy, long persistent and reflexed in young fruits 8. Galtonia 
13b. Capsule subglobose, with truncate apex. Tepals not fleshy, withering soon and never reflexed in fruit 16. Melomphis 
14a. Tepals bearing a wide longitudinal band, green, pinkish or brownish, along the medial nerve, only visible on the abaxial side 15 
14b. Tepals concolour, without a longitudinal coloured band on the abaxial side, sometimes with maculae at the base or apex 19 
15a. Capsule with six evident ribs in section. Seeds globose, with reticulate testa 16 
15b. Capsule trigonous or trilobate in section. Seeds irregularly compressed, with papillate to echinulate or puzzle-like testa 17 
16a. Inflorescece corymbose to racemose, with basal pedicels >1 cm. Flowers stellate at the anthesis, erect to erect-patent; filaments lanceolate to tapering, attenuate at the apex 19. Ornithogalum 
16b. Inflorescence racemose, with basal pedicels ≤1 cm long. Flowers campanulate at the anthesis, erect to slightly nodding; filaments tapering, at least the inner ones with two evident lobes or teeth accompanying the anther 18. Honorius 
17a. Plants tall, ≤140 cm. Leaves glabrous, not sheathing the stem by a tubular cataphyll; inflorescence (excluding the stem) ≤50 cm long; capsule not hidden by the withered perianth 17. Loncomelos 
17b. Plants small to middle size, ≤50 cm. Leaves usually ciliate or hirsute, sheathing the stem by a tubular cataphyll; inflorescences shorter; capsule usually hidden by the withered perianth 18 
18a. Leaves commonly not occurring with the flowers, sheathing and ascending upon the stem, being the limbs patent and usually arranged at different heights. Seeds mostly apiculate (comma-shaped), usually tuberculate or long papilose 10. Eliokarmos 
18b. Leaves usually occurring with the flowers, all emerging almost at the same level and disposed mostly on an epigeous neck. Seeds many angled, smooth 14. Nicipe 
19a. Flowers greenish. Filaments petaloid, showy, pure white, and spreading. Capsule obcordate or widely globose and truncate to retuse at the apex, equal or wider than long 9. Ethesia 
19b. Flowers white, yellow or orange. Filaments neither clearly spreading nor showy. Capsule elliptical, longer than wide 20 
20a. Tepals white, yellow, orange or reddish, usually with evident dark maculae at their base or apex; leaves usually ciliate or fimbriate on the edge, or pilose on the blade, rarely glabrous; filaments subulate or lanceolate and flat, usually with quadrate basal expansions or with two erect lobes concealing the ovary 10. Eliokarmos 
20b. Tepals always white, not maculate; leaves glabrous; filaments flat, lanceolate to tapering, without abrupt basal expansions 21 
21a. Plants with scarcely developed bulbs, subrhizomatous. Inflorescence corymbose. Ovary stipitate. Capsule black 13. Elsiea 
21b. Plants with swollen, well-developed bulb. Inflorescence racemose to spiciform. Ovary not stipitate. Capsule pale brown 15. Cathissa 
1a. Floral bract with small bracteoles. Tepals all fused over two-thirds of their length, into a cylindrical tube, slightly swollen at the base and somewhat curved; filaments inserted at the mouth of the tube 1. Pseudogaltonia 
1b. Floral bract without bracteoles (if present, then leaves narrowed in a pseudopetiole). Tepals free or fused into a shorter, campanulate to cylindrical, straight tube; filaments free, connate or adnate to the tube 
2a. Tepals with a broad and dark longitudinal band – green to brownish, evident on both sides and usually associated with three to many aggregated medial nerves 
2b. Tepals concolour, exceptionally with dark maculae at the base or apex, or sometimes with a broad longitudinal coloured band, only visible on the abaxial side, not well defined along the medial nerve 
3a. Ovary with two prominent dorsal keels on each carpel. Plants with only one leaf (exceptionally up to three), usually flat on the ground 4. Trimelopter 
3b. Ovary without such prominent dorsal keel. Plants usually with numerous leaves, erect or spreading (exceptionally solitary, varying in morphology) 
4a. Seed obliquely stalked in each locule. Inflorescence usually subspiciform, dense, and many flowered 7. Stellarioides 
4b. Seed horizontally stalked in each locule. Inflorescence usually racemose to subcorymbose, lax, with long pedicellate flowers 
5a. Capsule widely quadrate in outline, deeply three lobed; seeds uniseriate in each locule 
5b. Capsule ovate, oblong or obovate in outline, trigonous to trilobate; seeds biseriate in each locule (exceptionally uniseriate) 
6a. Inflorescence subspicate, mostly secund, with shortly pedicellate flowers. Outer tepals curved outwards, inners long connivent to form a cylindrical tube with reflexed tips 2. Dipcadi 
6b. Inflorescence racemose, cylindrical, conical or subpyramidal, with long pedicellate flowers. Tepals all similar and spreading 3. Battandiera 
7a. Tepals usually dimorphic: the outers mucronate and the inners cucullate or with an inflexed flap at the apex; filaments usually with basal constrictions; ovary with paraseptal keels; style trigonous in section, prismatic to obtriangular or sometimes thin or tapering; stigma not capitate, usually trilobed 5. Albuca 
7b. Tepals monomorphic, all similar; filaments without basal constrictions; ovary without paraseptal keels; style circular in section, subcylindric, usually deflexed; stigma capitate 6. Coilonox 
8a. Tepals fused at the base 
8b. Tepals completely free at the base 12 
9a. Flowers nodding. Tepals fused into a long campanulate tube of approx. half of their length. Stamens hidden among the perianth lobes 8. Galtonia 
9b. Flowers patent to erect-patent. Tepals fused into a tube ≤ one-third of their length. Stamens clearly visible 10 
10a. Leaves tapering at the base and pseudopetiolate; inflorescence usually with two flowers per bract in the lower part; bracteoles present in the upper flowers; capsule retuse or truncate at the apex 11. Avonsera 
10b Leaves sheathing at the base, never pseudopetiolate; inflorescence always with one flower per bract; bracteoles absent; capsule acute to subacute at the apex 11 
11a. Tepals ≤ half the length of capsule; seeds piriform or subglobose; filaments of stamen connate 12. Neopatersonia 
11b. Tepals longer than capsule or rarely slightly shorter; seeds angular, irregularly compressed; filaments of stamen free to adnate, exceptionally connate 14. Nicipe 
12a. Ovary globose, shining black. Inflorescence subcorymbose 13 
12b. Ovary fusiform to oblong, green or yellow. Inflorescence racemose to corymbose 14 
13a. Capsule ovate, with acute apex. Tepals fleshy, long persistent and reflexed in young fruits 8. Galtonia 
13b. Capsule subglobose, with truncate apex. Tepals not fleshy, withering soon and never reflexed in fruit 16. Melomphis 
14a. Tepals bearing a wide longitudinal band, green, pinkish or brownish, along the medial nerve, only visible on the abaxial side 15 
14b. Tepals concolour, without a longitudinal coloured band on the abaxial side, sometimes with maculae at the base or apex 19 
15a. Capsule with six evident ribs in section. Seeds globose, with reticulate testa 16 
15b. Capsule trigonous or trilobate in section. Seeds irregularly compressed, with papillate to echinulate or puzzle-like testa 17 
16a. Inflorescece corymbose to racemose, with basal pedicels >1 cm. Flowers stellate at the anthesis, erect to erect-patent; filaments lanceolate to tapering, attenuate at the apex 19. Ornithogalum 
16b. Inflorescence racemose, with basal pedicels ≤1 cm long. Flowers campanulate at the anthesis, erect to slightly nodding; filaments tapering, at least the inner ones with two evident lobes or teeth accompanying the anther 18. Honorius 
17a. Plants tall, ≤140 cm. Leaves glabrous, not sheathing the stem by a tubular cataphyll; inflorescence (excluding the stem) ≤50 cm long; capsule not hidden by the withered perianth 17. Loncomelos 
17b. Plants small to middle size, ≤50 cm. Leaves usually ciliate or hirsute, sheathing the stem by a tubular cataphyll; inflorescences shorter; capsule usually hidden by the withered perianth 18 
18a. Leaves commonly not occurring with the flowers, sheathing and ascending upon the stem, being the limbs patent and usually arranged at different heights. Seeds mostly apiculate (comma-shaped), usually tuberculate or long papilose 10. Eliokarmos 
18b. Leaves usually occurring with the flowers, all emerging almost at the same level and disposed mostly on an epigeous neck. Seeds many angled, smooth 14. Nicipe 
19a. Flowers greenish. Filaments petaloid, showy, pure white, and spreading. Capsule obcordate or widely globose and truncate to retuse at the apex, equal or wider than long 9. Ethesia 
19b. Flowers white, yellow or orange. Filaments neither clearly spreading nor showy. Capsule elliptical, longer than wide 20 
20a. Tepals white, yellow, orange or reddish, usually with evident dark maculae at their base or apex; leaves usually ciliate or fimbriate on the edge, or pilose on the blade, rarely glabrous; filaments subulate or lanceolate and flat, usually with quadrate basal expansions or with two erect lobes concealing the ovary 10. Eliokarmos 
20b. Tepals always white, not maculate; leaves glabrous; filaments flat, lanceolate to tapering, without abrupt basal expansions 21 
21a. Plants with scarcely developed bulbs, subrhizomatous. Inflorescence corymbose. Ovary stipitate. Capsule black 13. Elsiea 
21b. Plants with swollen, well-developed bulb. Inflorescence racemose to spiciform. Ovary not stipitate. Capsule pale brown 15. Cathissa 

1. Pseudogaltonia

Kuntze in Jahrb. Königl. Bot. Gart. Berlin 4: 274. 1886 [nom. alternat.] ≡ Hyacinthus section Pseudogaltonia Kuntze in Jahrb. Königl. Bot. Gart. Berlin 4: 274. 1886 [nom. alternat.] – Holotype: P. pechuelii Kuntze ≡ Hyacinthus pechuelii Kuntze = P. clavata (Mast.) E. Phillips ≡ Galtonia clavata Mast.

= Lindneria T. Durand & Lubbers in Bull. Soc. Bot. France 36: CCXVII. 1890 – Holotype: L. fibrillosa T. Durand & Lubbers = P. clavata (Mast.) E. Phillips

Plants large. Bulb large, with outer tunics cartilaginous and fibrous. Leaves widely lanceolate, flat, long, acute, glabrous. Inflorescence in a loose pyramidal raceme; bracts linear-acuminate; bracteoles (prophylls) present. Tepals white with greenish or greyish longitudinal band, fused over two-thirds of their length, into a tube approx. 3 cm long, slightly swollen at the base and somewhat curved; filaments exerted, flat, insert at the mouth of the tube; style filiform and long (>2 cm). Capsule quadrate, truncate, and deeply trilobate. Seeds flat, disciform and uniseriate in each locule, with five to seven angled polygonal subisodiametric testa cells with shallow depressions and pitted surface.

Chromosome number: 2n = 18 (cf. Speta, 1998a; Forrest and Jong, 2004).

Number of species: 2 (cf. Manning et al., 2009b).

Distribution: north-western South Africa, Namibia and Botswana.

2. Dipcadi

Medik. in Hist. & Commentat. Acad. Elect. Sci. Theod.-Palat. 6: 431. 1790 – Holotype: D. serotinum (L.) Medik. ≡ Hyacinthus serotinus L.

= Zuccagnia Thunb., Nov. Gen. Pl. 9: 127. 1798. – Type: Z. viridis (L.) Thunb. ≡ D. viride (L.) Moench.

= Uropetalon Burch. ex Ker Gawl. in Bot. Reg. 2: t. 156. 1816 ≡ D. ser. Uropetalon (Burch. ex Ker Gawl.) J.C. Manning & Goldblatt in Taxon 58(1): 99. 2009 – Type: U. glaucum Burch. ex Ker Gawl. in Bot. Reg. 2: t. 156. 1816 ≡ D. glaucum (Burch. ex Kew Gawl.) Baker

= Polemannia K. Bergius ex Schltdl. in Linnaea 1: 250. 1826, nom. rej. [non Polemannia Eckl. & Zeyh. in Enum. Pl. Afric. Austral. 3: 347. 1837, nom. cons. (Umbelliferae)] – Type: P. hyacinthiflora Berg. ex Schltdl. = D. brevifolium (Thunb.) Fourc.

= Tricharis Salisb., Gen. Pl. [Salisbury]: 24. 1866 – Type: T. serotina (L.) Salisb. ≡ D. serotinum (L.) Medik.

Plants small to middle size. Leaves variable, usually linear, flattened or canaliculate, or more or less twisted, glabrous to hirsute. Inflorescence racemose or subspiciform, usually secund, with patent or nodding flowers; bracts ovate or attenuate, membranous; bracteoles absent. Tepals lanceolate-tapering, fused at the base into a tube approx. one-quarter to one-third (to two-thirds) of their length; inner ones suberect or connivent and reflexed at the apical third; outer ones reflexed, sometimes with filiform caudate apendixes; filaments short, linear to flatish, adnate to the tube and included. Capsule deeply 3-lobed, quadrate in outline, truncate at the apex, rarely umbonate, sometimes stipitate. Seeds black, discoid, flat, horizontally stacked and uniseriate per locule, with 5–6 angled polygonal subisodiametric or up to twice as long as broad testa cells with shallow depressions and smooth or pitted surface.

Chromosome number: 2n = 8, 12, 18, 20, 24, 36, 64 (cf. Speta, 1998a, Jones and Smith, 1967).

Number of species: approx. 30 (cf. Obermeyer, 1964; Speta, 1998a; Manning et al., 2002, 2009a).

Distribution: south, east and north Africa, Socotra, Madagascar, Europe, Middle East and India.

3. Battandiera

Maire in Bull. Soc. Hist. Nat. Afr. Nord 17(3): 125. 1926 ≡ Ornithogalum section Trineuron Batt. in Bull. Soc. Bot. France 63: 195. 1919 – Holotype: B. amoena (Batt.) Maire ≡ O. amoenum Batt.

= Ornithogalum section Namibiogalum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 487. 1996 [basion.] ≡ Albuca subgenus Namibiogalum (U.Müll.-Doblies & D.Müll.-Doblies) J.C.Manning & Goldblatt in Taxon 58(1): 92. 2009 – Holotype: B. rautanenii (Schinz) Mart.-Azorín, M.B.Crespo & Juan ≡ O. rautanenii Schinz.

Plants usually middle size. Bulb ovoid, often with an evident neck at the apex with transverse thin darker nerves; outer tunics numerous, coriaceous, coarse and greyish, protecting the bulb. Leaves usually narrowly lanceolate to filiform, flat to folded, glabrous and somewhat glaucous and sometimes with darker spots at the base beneath; bracteoles absent. Inflorescence conical to subpyramidal, elongating with age, sometimes with a plumose or pendulous apex; bracts membranous, deltoid to filiform, usually longer than flowers. Tepals white or greenish, oblong, linear, obtuse, spreading or rarely reflexed, with a green or brownish longitudinal band visible on both sides (rarely the band indistinct but with darker medial aggregated nerves evident when dried), free or shortly fused into a tube ≤5 mm long; filaments linear-acuminate or ovate-acuminate, sometimes with quadrate basal expansions or shortly adnate to the perianth; ovary ovoid or oblong-globose, trigonous or trilobate, sometimes stipitate or expanded at the base into a 6-lobed disc . Capsule deeply trilobate with narrow lobes, often widely quadrate in outline, wider than long, truncate at the apex or emarginate, with darker transversal nerves. Seeds large, discoid, flat, horizontally stacked and uniseriate in each locule, with angled and polygonal or slightly undulate testa cells.

Chromosome number: 2n = 20, 24 (cf. Speta, 2001; Vosa, 1980).

Number of species: 8 (according to the present study).

Distribution: arid regions of southern and East Africa (excepting the southern half of South Africa), with a disjoint species in North Africa (B. amoena).

Necessary combinations:

Battandiera candida (Oberm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum candidum Oberm. in Bothalia 10: 355. 1971 [basion.]

Battandiera donaldsonii (Rendle) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Albuca donaldsonii Rendle in J. Bot. 34: 131. 1896 [basion.] ≡ O. donaldsonii (Rendle) Greenway in Journ. E. Afr. Nat. Hist. Soc. 27: 203. 1969

Battandiera pulchra (Schinz) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum pulchrum Schinz in Verh. Bot. Ver. Brandenb. 31: 221. 1890 [basion.]

Battandiera rautanenii (Schinz) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum rautanenii Schinz in Verh. Bot. Ver. Brandenb. 31: 221. 1890 [basion.]

Battandiera seineri (Engl. & K.Krause) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Bulbine seineri Engl. & K.Krause in Bot. Jahrb. 45: 124. 1910 [basion.]

Battandiera stapffii (Schinz) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum stapffii Schinz in Bull. Herb. Boiss. 4, App. 3: 42. 1896 [basion.]

Battandiera tubiformis (Oberm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum candidum var. tubiforme Oberm. in Bothalia 10: 357. 1971 [basion.]

4. Trimelopter

Raf., Fl. Tellur. 2: 24. 1837 ≡ Ardernia Salisb., Gen. Pl.: 35. 1866 ≡ Ornithogalum section Monarchos U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 470. 1996 ≡ Albuca subgenus Monarchos (U.Müll.-Doblies & D.Müll.-Doblies) J.C.Manning & Goldblatt in Taxon 58(1): 92. 2009 – Holotype: T. fuscatum (Jacq.) Raf. ≡ O. fuscatum Jacq. = T. unifolium (Retz.) Mart.-Azorín, M.B.Crespo & Juan ≡ O. unifolium Retz.

Plants small to middle size. Bulb ovate, outer tunics membranous or coriaceous, greyish, usually with a long neck. Leaf one (rarely ≤3 in cultivation), ovate to elliptical or narrowly oblong, with obtuse to acute apex, glabrous or diversely pubescent, occurring with flowers or withering at flowering time, sheathing into a contracted tubular base, and firm blade, usually appressed to the ground or spreading; bracteoles absent. Inflorescence racemose or pyramidal; bracts deltoid, aristate, auriculate, sometimes with papillate adaxial side and edges. Tepals pale yellow, brownish or whitish, with a central greenish or brownish band visible on both sides (sometimes indistinct on the adaxial side, though associated to evident medial nerves); filaments narrowly ovate, expanded below; ovary ovoid or obovoid, trilobate with three prominent carpels, each one with two longitudinal keels down the back and a groove between. Capsule globose, small (≤4 mm), trilobate, with two prominent keels on each carpel. Seeds flat, semidiscoid, or irregularly compressed, with a straight side and the other curved, with double ridge, and subpolygonal, granulate testa.

Chromosome number: no data available.

Number of species: approx. 10 (according to the present study).

Distribution: mainly in southern Namibia and western South Africa, spreading to central and north-eastern South Africa.

Necessary combinations:

Trimelopter costatulum (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum costatulum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 482. 1996 [basion.]

Trimelopter etesiogaripense (U.Müll.-Doblies & D.Müll.- Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum etesiogaripense U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 476. 1996 [basion.]

Trimelopter etesiogaripense subsp. longipilosum (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum etesiogaripense subsp. longipilosum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 478. 1996 [basion.]

Trimelopter dyeri (Poelln.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum dyeri Poelln. in Ber. Deutsch. Bot. Ges. 61: 209. 1944 [basion.]

Trimelopter monarchos (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum monarchos U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 476. 1996 [basion.]

Trimelopter ovatum (Thunb.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum ovatum Thunb., Prodr. 62. 1794 [basion.]

Trimelopter ovatum subsp. oliverorum (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum ovatum subsp. oliverorum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 474. 1996 [basion.]

Trimelopter pendulinum (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum pendulinum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 480. 1996 [basion.]

Trimelopter psammophorum (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum psammophorum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 482. 1996 [basion.]

Trimelopter strigulosum (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum strigulosum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 480. 1996 [basion.]

Trimelopter unifolium (Retz.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum unifolium Retz., Obs. 2: 17. 1781 [basion.] = T. fuscatum (Jacq.) Raf. ≡ O. fuscatum Jacq.

Trimelopter watermeyeri (L.Bolus) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum watermeyeri L.Bolus in S. African Gard. 21: 13. 1931 [basion.]

5. Albuca

L., Sp. Pl., ed. 2. 1: 438. 1762 – Lectotype: Albuca canadensis (L.) F.M. Leight. [≡ Ornithogalum canadense L. ≡ A. major L., nom. illeg.]

Plants mostly middle to large size. Bulb hypogeous or epigeous, diverse in size; outer tunics membranous, coriaceous, fibrous or fleshy and green. Leaves variable, usually narrowly lanceolate to filiform, flattened to folded or inrolled, glabrous, hirsute or glandular, sometimes spirally twisted or coiled, and exceptionally with a longitudinal white band. Inflorescence racemose or subcorymbose, glabrous, papillose or glandular; bracteoles absent. Flowers nodding, patent or erect; tepals white to yellow with a broad central greenish to brownish band visible on both sides, whorls dimorphic: the outer ones mucronate, suberect or spreading, and the inner ones cucullate, hooked or with an inflexed flap at the apex, erect and more or less connivent and hiding the stamens and the style; filaments narrowly oblong or linear-lanceolate, more or less expanded at the base, with a constriction or folded basally, sometimes at least the outers with anthers smaller or absent; ovary ovoid to elliptical, with conspicuous paraseptal crests, with many ovules per locule; style as long as the perianth, trigonous in section, prismatic to obtriangular or thin and tapering, sometimes strongly papillate; stigma trilobate, truncate, depressed or conical, sometimes with horns, papillate or fimbriate with long papillae. Capsule ovoid or oblong, acute or obtuse, more or less triangular in section with blunt edges. Seeds black, numerous, flattened, horizontally stacked, semicircular with one straight side forming a thicked ridge and biseriate in each locule (or rarely discoid and uniseriate), with puzzle-like (or rarely granulate) testa.

Chromosome number: 2n = 16, 18, 20, 36, 40 (cf. Jones and Smith, 1967; Speta, 1998a).

Number of species: ranging from 60 to approx. 130 (cf. Speta, 1998a; Manning et al., 2002; Phillips, 1926).

Distribution: mainly in southern and east Africa, extending to Ethiopia and Saudi Arabia, and north of Chad and Nigeria.

6. Coilonox

Raf., Fl. Tellur. 2: 28. 1837 – Holotype: C. albucoides (Aiton) Raf. ≡ Anthericum albucoides Aiton

= Osmyne Salisb., Gen. Pl. [Salisbury]: 35. 1866 ≡ Ornithogalum subgenus Osmyne (Salisb.) Baker in J. Linn. Soc., Bot. 13(68): 278. 1872 ≡ Ornithogalum. section Osmyne (Salisb.) Engl., Nat. Planzenfam. [Engler & Prantl] 2, Abt. 5: 67. 1888 ≡ Albuca subgenus Osmyne (Salisb.) J.C.Manning & Goldblatt in Taxon 58(1): 93. 2009 ≡ Monotassa Salisb., Gen Pl. [Salisbury]: 36. 1866 – Holotype: Osmyne odorata (Jacq.) Salisb. [≡ Ornithogalum odoratum Jacq.] = C. secundum (Jacq.) Speta ≡ Ornithogalum secundum Jacq. ≡ Monotassa secunda (Jacq.) Salisb.

= Taeniola Salisb., Gen Pl. [Salisbury]: 35. 1866 [non Taeniola Bonorden, Handb. Allg. Mykol.: 36. 1851 (Fungi)] – Holotype: T. vittata (Ker Gawl.) Salisb. ≡ Albuca vittata Ker Gawl. ≡ C. vittatum (Ker. Gawl.) Speta

= Ornithogalum section Monosmyne U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 515. 1996 – Holotype: Ornithogalum zebrinum (Baker) Oberm. ≡ Albuca zebrina Baker ≡ C. zebrinum (Baker) Speta

= Ornithogalum ser. Clavifolia U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 516. 1996 – Holotype: Albuca unifoliata Rowley ≡ C. unifoliatum (Rowley) Speta

Plants usually medium size. Bulb usually large, hypogeous or epigeous; outer tunics membranous, coriaceous or fleshy and green. Leaves variable, usually narrowly lanceolate to succulent and claviform, glabrous, hirsute or glandular, sometimes spirally twisted or coiled. Inflorescence racemose or subcorymbose, glabrous, hirsute or glandular; bracteoles absent. Flowers spreading, patent; tepals whitish or yellowish with a broad central green band visible on both sides; whorls equal; filaments subulate and narrowly oblong or linear-lanceolate, not constricted and all anthers well developed; ovary ovoid to elliptical, without paraseptal crests, with many ovules per locule; style as long as the perianth, subcylindrical, usually deflexed at anthesis; stigma capitate, long papillate or fimbriate. Capsule ovoid or oblong, acute or obtuse, more or less triangular in section with blunt edges. Seeds black, numerous, flattened, semicircular with one straight side forming a thicked ridge, or discoid, horizontally stacked and biseriate in each locule, with puzzle-like testa.

Chromosome number: 2n = 18 (cf. Speta, 1998a).

Number of species: approx. 30 (cf. Speta, 2001; Manning et al., 2009a).

Distribution: mainly in western South Africa, extending to southern Namibia, in winter-rainfall areas.

Necessary combination:

Coilonox paucifolium subsp. karooparkense (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum paucifolium subsp. karooparkense U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 515. 1996 [basion.]

7. Stellarioides

Medik., Hist. & Commentat. Acad. Elect. Sci. Theod.-Palat. 6: 369. 1790 – Holotype: S. canaliculata Medik. = Ornithogalum longebracteatum Jacq. ≡ S. longebracteata (Jacq.) Speta

= Urophyllon Salisb., Gen. Pl. [Salisbury]: 34. 1866 ≡ O. subgenus Urophyllon (Salisb.) Baker ≡ O. section Urophyllon series Urophyllon (Salisb.) U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 488. 1996 ≡ Albuca subgenus Urophyllon (Salisb.) J.C.Manning & Goldblatt in Taxon 58(1): 93. 2009 – Lectotype: U. caudatum (Jacq.) Salisb. = O. longebracteatum Jacq. [cf. Speta (2001); lectotypifications by Obermeyer (1978) and Müller-Doblies & Müller-Doblies (1996) are incorrect or superfluous].

Plants variable in habit, usually medium to large size. Bulb globose to ovoid, hypogeous or sometimes epigeous, green and bulbilliferous (rarely with a long neck in plants from arid regions); outer tunics membranous and greyish or fleshy and green. Leaves lanceolate or linear, long tapering, flat to canaliculate, glabrous. Inflorescence 1–3 per bulb, usually long racemose and many flowered, subspiciform, dense; bracts narrowly ovate-acuminate or filiform, aristate, membranous, usually long and exceeding the buds, or rarely shorter and simply ovate; bracteoles absent. Flowers small; tepals whitish to greenish, with a dark longitudinal green band visible on both sides, oblong or narrowly elliptical, ≤12 mm long; filaments ovate-acuminate or linear, flattened, sometimes with basal lobed expansions; ovary ovoid or globose, obtuse, trigonous, and with biseriate ovules; style thickened, sometimes widened at the base; stigma tapering, not capitate. Capsule usually small, ≤14 mm long, ovoid to obovate, obtuse or acute, and trilobate in section. Seeds variable in size, numerous, irregularly compressed, flattened, roundish, angular or semicircular, biseriate and obliquely stalked in each locule, with puzzle-like testa.

Chromosome number: 2n = 4, 6, 8, 10, 12, 16, 18, 20, 24, 26, 36, 54 (cf. Stedje, 1989; Nordal and Stedje, 1993; Speta, 1998a, 2001).

Number of species: approx. 25 (cf. Speta, 2001; and according to the present study) to 2 (with three additional subspecies; cf. Manning et al., 2009a, but excluding Igidia volubilis). A comprehensive revision is still needed for this genus.

Distribution: temperate and subtropical regions of southern Africa, spreading north ≤ Ethiopia, and with a species reaching disjointly Algeria and Morocco (S. sessiliflorum).

Necessary combinations:

Stellarioides bakeri Mart.-Azorín, M.B.Crespo & Juan, nom. nov.Ornithogalum chloranthum Baker in Gard. Chron. n.s. 2: 323. 1875, nom. illeg. (syn. subst.) [non M.Bieb., Fl. Taur.-Caucas. 2: 264. 1808 ≡ Gagea chlorantha (M.Bieb.) Schult.f.]

Stellarioides cernua (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum cernuum Baker, Fl. Trop. Afr. [Oliver et al.] 7(3): 547. 1898 [basion.]

Stellarioides gambosana (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum gambosanum Baker, Fl. Trop. Afr. [Oliver et al.] 7(3): 547. 1898 [basion.]

8. Galtonia

Decne. in Fl. Serres Jard. Eur. 23: 32. 1880 – Lectotype: G. princeps (Baker) Decne. ≡ Hyacinthus princeps Baker (cf. Manning et al., 2004).

Plants large and vigorous. Bulb globose; outer tunics soft and membranous. Leaves numerous, lanceolate tapering, wide (>5 cm), glabrous and somewhat sheathing and ascending upon the stem. Inflorescence widely racemose, conical or cylindrical, sometimes corymbose; pedicels patent, with nodding apices in flower (rarely erect) and always erect in fruit; bracts membranous, linear-acuminate; bracteoles absent. Flowers never fragrant, nodding, exceptionally erect; tepals fleshy, whitish or greenish-yellowish, fused into a campanulate tube ≤ half of their length or rarely free; filaments ovate-acuminate, insert at the base of the perianth and adnate to the perianth tube, merging at its end, sometimes completely free; ovary green, yellowish or black, cylindrical or ovoid; style white, filiform; stigma slightly trilobate and glandular. Capsule lanceolate, lanceolate-cylindrical or ovate, acute, widened in the basal third, triangular in section with blunt edges. Seeds large, polygonal or irregularly compressed or semicircular and flattened, biseriate and obliquely stalked in each locule, with smooth puzzle-like testa.

Chromosome number: 2n = 12, 14, 16 (cf. Obermeyer, 1978; Speta, 1998a; Forrest and Jong, 2004). Exceptionally, some counts for G. candicans yielded 2n = 22, 32 (cf. Hammett and Murray, 1993).

Number of species: 5 (according to the present study).

Distribution: temperate and subtropical regions of eastern South Africa and Swaziland.

Necessary combination:

Galtonia saundersiae (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum saundersiae Baker in Gard. Chron., ser. 3, 10: 452. 1891 [basion.]

9. Ethesia

Raf., Fl. Tellur. 3: 60. 1837 ≡ Ornithogalum series Xanthochlora U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert 107: 492. 1996 ≡ O. section Xanthochlora (U.Müll.-Doblies & D.Müll.-Doblies) J.C.Manning & Goldblatt in Taxon 58(1): 100. 2009 – Holotype: E. prasina (Ker Gawl.) Raf. ≡ O. prasinum Ker Gawl.

Plants middle size. Bulb ovoid; outer tunics membranous or coriaceous, rough, greyish. Leaves numerous, linear to lanceolate or oblong, canaliculate or lorate and flat, somewhat succulent, glaucous or green, glabrous, rarely not occurring with the flowers. Inflorescence a raceme cylindrical or pyramidal at the begining; pedicels erect, firm, sometimes arcuate and lengthening and hardening in fruit, with an obtuse callus above; bracts linear to lanceolate, acuminate, membranous, equalling, shorter or longer than the pedicels; bracteoles absent. Flowers usually fragrant; tepals greenish, or green with a white marginal region, narrowly oblong or ovate, obtuse to subacute; filaments petaloid, showy, pure white, spreading, ovate-acuminate and almost overlapping at the base; ovary green, oblong-ovoid or oblong-globose, triangular; style erect, filiform; stigma slightly trilobate and papillate. Capsule oblong to obcordate or widely globose and truncate to retuse at the apex, as wide as long, trilobate in section. Seeds numerous, large, irregularly compressed (with one or two edges) to ovate-discoid, biseriate in each locule, with blackish testa.

Chromosome number: 2n = 16 (cf. Obermeyer, 1978).

Number of species: 4 (according to the present study).

Distribution: southern Namibia and arid regions of western and center South Africa.

Necessary combinations:

Ethesia haalenbergensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum haalenbergense U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 492. 1996 [basion.]

Ethesia polyphlebia (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum polyphlebium Baker, Fl. Cap. 6(3): 507. 1897 [basion.]

Ethesia xanthochlora (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum xanthochlorum Baker, Fl. Cap. 6(3): 508. 1897 [basion.]

10. Eliokarmos

Raf., Fl. Tellur. 2: 24. 1837 – Lectotype: E. thyrsoides (Jacq.) Raf. ≡ Ornithogalum thyrsoides Jacq. (cf. Rafinesque, Fl. Tellur. 3: 58. 1837).

= Tomoxis Raf., Fl. Tellur. 3: 54. 1837– Lectotype (designated by Speta, 2001): T. coarctata (Jacq.) Raf. ≡ O. coarctatum Jacq. [= O. thyrsoides Jacq.]

= Lomaresis Raf., Fl. Tellur. 3: 59. 1837 – Holotype: L. alba Raf., nom. illeg. (syn. subst.) ≡ O. revolutum Jacq. [= O. thyrsoides Jacq.]

= Aspasia Salisb., Gen. Pl. [Salisbury]: 34 & 40. 1866 ≡ O. subgenus Aspasia (Salisb.) Oberm. ≡ O. subgenus Aspasia section Aspasia (Salisb.) U.Müll.-Doblies & D.Müll.-Doblies – Lectotype (designated by Obermeyer, 1978): O. conicum Jacq.

= Phaeocles Salisb., Gen. Pl. [Salisbury]: 35. 1866 – Holotype: P. maculata (Jacq.) Salisb. ≡ O. maculatum Jacq.

= O. subgenus Aspasia section Hispidaspasia U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert 107: 404. 1996 – Holotype: O. hispidum Hornem.

Plants small to middle size. Bulb globose; outer tunics thin and membranous to firm, thickened and rough, pale or dark coloured. Leaves numerous, lanceolate or ovate, usually ciliate or fimbriate on margins, the blade glabrous or rarely hirsute, sometimes not occurring with the flowers, or sheathing and ascending along the stem, with patent and subdistichous blades, arranged at different levels. Inflorescence racemose, cylindrical or corymbose; pedicels erect to patent in flower and erect in fruit; bracts membranous, lanceolate, wider at the base, auriculate and petaloid, or aristate; bracteoles absent. Flowers erect or spreading, diurnal (rarely nocturnal); tepals thin textured, free, diversely coloured, from pure white to yellow, orange or reddish, sometimes with dark maculae at the base or the apex, concolour or sometimes with a narrow greenish, brownish or pinkish band only visible on the abaxial side, persistent and papery in fruit, not rolled or reflexed when withering; filaments variously expanded, from linear-subulate to lanceolate or widened at the base, sometimes generating erect lobes, the inners usually wider than the outers; ovary green, yellow or dark grey, ovoid; style filiform, erect or rarely deflexed; stigma slightly trigonous and papillate. Capsule ovate, lanceolate, elliptical or fusiform, with thin walls, hidden among the persistent dry perianth, with triangular section and blunt edges. Seeds small (0·5–2 mm), numerous, irregularly angular to piriform, variously compressed, biseriate or multiseriate per locule, with papillate, echinulate, or/and puzzle-like testa.

Chromosome number: 2n = 12 (cf. Neves, 1962; Obermeyer, 1978; Speta, 1998a).

Number of species: approx. 28 (according to the present study).

Distribution: from southern Namibia to western and south-west South Africa, mainly in winter rainfall regions.

Necessary combinations:

Eliokarmos bicornutus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum bicornutum F.M.Leight. in J. S. African Bot. 11: 143. 1945 [basion.]

Eliokarmos bolusianus (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum bolusianum Baker in J. Linn. Soc., Bot. 13: 279. 1872 [basion.]

Eliokarmos ceresianus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum ceresianum F.M.Leight. in J. Bot., Lond. 71: 72. 1933 [basion.]

Eliokarmos ciliiferus (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum ciliiferum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 414. 1996 [basion.]

Eliokarmos conicus (Jacq.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum conicum Jacq., Collectanea [Jacquin] 3: 232. 1791 [basion.]

Eliokarmos constrictus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum constrictum F.M.Leight. in J. S. African Bot. 11: 136. 1945 [basion.]

Eliokarmos corticatus (Mart.-Azorín) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum corticatum Mart.-Azorín in Bothalia 37(2): 140. 2007 [basion.]

Eliokarmos decus-montium (G.Will.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum decus-montium G.Will. in Bothalia 28(1): 63. 1998 [basion.]

Eliokarmos deltoideus (Baker) Mart.-Azorín, M.B. Crespo & Juan, comb. nov.Ornithogalum deltoideum Baker in J. Linn Soc. (Bot.) 13: 281. 1872 [basion.]

Eliokarmos dubius (Houtt.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum dubium Houtt., Handl. Pl.-Kruidk. 2(12): 309, t. 82, fig. 3. 1780 [basion.] =E. aureus (Curtis) Raf. ≡ O. aureum Curtis

Eliokarmos geniculatus (Oberm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum geniculatum Oberm. in Bothalia 12(3): 344. 1978 [basion.]

Eliokarmos gifbergensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum gifbergense U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 418. 1996 [basion.]

Eliokarmos hallii (Oberm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum hallii Oberm. in Bothalia 12(3): 347. 1978 [basion.]

Eliokarmos hispidulus (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.O. hispidulum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 411. 1996 [basion.]

Eliokarmos inandensis (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum inandense Baker, Fl. Cap. (Harvey) 6(3): 503. 1897 [basion.]

Eliokarmos inclusus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum inclusum F.M.Leight. in S. African Gard. 24: 50. 1934 [basion.]

Eliokarmos leeupoortensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum leeupoortense U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 396. 1996 [basion.]

Eliokarmos namaquanulus (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum namaquanulum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 410. 1996 [basion.]

Eliokarmos naviculus (F.W.Barker ex Oberm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum naviculum F.W.Barker ex Oberm. in Bothalia 12(3): 348. 1978 [basion.]

Eliokarmos neomaculatus Mart.-Azorín, M.B.Crespo & Juan, nom. nov.Ornithogalum maculatum Jacq., Collectanea [Jacquin] 3: 368. 1791 (syn. subst.) [non E. maculatus Raf., Fl. Tellur. 3: 59. 1837 =E. thyrsoides (Jacq.) Raf.]

Eliokarmos pilosus (Jacq.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Anthericum pilosum Jacq., Ic. Pl. Rar. 2(15): t. 416. 1794 [basion.] = Ornithogalum hispidum Hornem.

Eliokarmos pilosus subsp. bergii (Schltdl.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum bergii Schltdl. in Linnaea 1: 254. 1826 [basion.]

Eliokarmos pruinosus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum pruinosum F.M.Leight. in J. S. African Bot. 10: 104. 1944 [basion.]

Eliokarmos puberulus (Oberm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum puberulum Oberm. in Bothalia 12(3): 337. 1978 [basion.]

Eliokarmos rubescens (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum rubescens F.M.Leight. in J. S. African Bot. 11: 142. 1945

Eliokarmos rupestris (L.f.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum rupestre L.f., Suppl. Pl.: 199. 1782 [basion.]

Eliokarmos strictus (L.Bolus) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum strictum L.Bolus in J. Bot. 71: 72. 1933 [basion.]

Eliokarmos synanthifolius (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum synanthifolium F.M.Leight. in J. S. African Bot. 11: 176. 1945 [basion.]

Eliokarmos thermophilus (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum thermophilum F.M.Leight. in J. S. African Bot. 11: 145. 1945 [basion.]

11. Avonsera

Speta in Phyton 38: 95. 1998 ≡ Ornithogalum subgenus Avonsera (Speta) J.C.Manning & Goldblatt in Taxon 58(1): 100. 2009 – Holotype: A. convallarioides (H.Perrier) Speta ≡ O. convallarioides H.Perrier

Plants middle size. Bulbs with membranous outer tunics. Leaves various, wide, elliptical to oblong or obovate, abruptly contracted at the base in a pseudopetiole, glabrous, occurring with flowers. Inflorescence loosely racemose, usually with two flowers per bract in its lower part; pedicel long and thin, erect to patent in flower and patent in fruit; bracts lanceolate, membranous, small; bracteoles present. Flowers diurnal, spreading; tepals thin, white, connate at the base; filaments deltoid or tapering and acute at the apex, connate at the base forming a short tube; ovary green, obovoid, truncate at the apex; style white, filiform; stigma punctiform. Capsule obovate, truncate at the apex, trilobate in section. Seeds ovoid.

Chromosome number: 2n = 14 (cf. Wetschnig et al., 2007).

Number of species: 1 (cf. Speta, 1998a).

Distribution: Madagascar.

12. Neopatersonia

Schönland in Rec. Albany Mus. 2: 251. 1912 ≡ Ornithogalum section Neopatersonia (Schönland) J.C.Manning & Goldblatt in Taxon 58(1): 101. 2009 – Holotype: N. uitenhagensis Schönland

Plants small. Bulb globose; outer tunics membranous or coriaceous, greyish, sometimes with a long neck. Leaf solitary to various, lanceolate, tapering or oblong, somewhat sheathing at the base. Inflorescence in subcorymbose raceme; pedicels erect-patent in flower and erect in fruit; bracts membranous, linear-acuminate, shorter than the pedicels; bracteoles absent. Flowers erect, remaining open; tepals somewhat fleshy, whitish or greenish, sometimes with longitudinal dark green nerves, oblong, and fused at the base in a very short tube; filaments subulate, with a expanded triangular or lobate base, connate and adnate to perianth; ovary green and ovoid; style whitish, filiform; stigma with three patent long and deflexed lobes, sometimes only trigonous. Capsule lanceolate or ovoid-triangular, acute or apiculate, triangular in section with blunt edges. Seeds black, small, angular, piriform or subglobose, biseriate in each locule, somewhat verruculose, with pitted puzzle-like testa cells.

Chromosome number: no data available.

Number of species: 4 (according to the present study).

Distribution: south-west and southern parts of South Africa.

Necessary combination:

Neopatersonia rotata (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum rotatum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 482. 1996 [basion.]

13. Elsiea

F.M.Leight. in J. S. African Bot. 10: 55. 1944 ≡ Ornithogalum section Elsiea (F.M.Leight.) J.C.Manning & Goldblatt in Taxon 58(1): 102. 2009 – Type: E. corymbosa F.M.Leight. ≡ O. esterhuyseniae Oberm. [non O. corymbosum Ruiz & Pavón (1802)]

Plant middle size, ≤70 cm. Bulb few developed, composed by tubular foliar sheaths growing up from a thick subterranean and subrhizomatous stem. Leaves numerous, occurring with the flowers, glabrous, smooth, linear-convolute, with margins rolled in a terete appendix at the apex. Inflorescence racemose-corymbose, erect, 60 cm long; with erect pedicels, 1–6 cm long; bracts shorter than the pedicels, linear-lanceolate, acuminate, auriculate and membranous; bracteoles absent. Flowers small, erect, white, thin; tepals white, linear, connate into a short tube 3–4 mm, and somewhat reflexed and persistent after withering; filaments linear-lanceolate, acute and shortly adnate to perianth, the inner ones somewhat wider than the outer ones; ovary obovoid, triangular, substipitate; style filiform; stigma slightly trigonous and papillate. Capsule black, ovoid-triangular sharply angled, stipitate, obtuse and depressed at the apex. Seeds small or medium size, angular, irregularly compressed, almost winged.

Chromosome number: no data available.

Number of species: 1 (according to the present study).

Distribution: mountainous areas of south-west of South Africa.

14. Nicipe

Raf., Fl. Tellur. 3: 54. 1837 – Holotype: N. nivea (Aiton) Raf. ≡ Ornithogalum niveum Aiton

= O. group Angustifoliati Oberm. in Bothalia 12: 349. 1978 ≡ O. subgenus Aspasia section Linaspasia U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert 107: 436. 1996 – Holotype: O. juncifolium Jacq.

= O. subgenus Aspasia section Vaginaspasia U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert 107: 428. 1996 ≡ O. series Vaginaspasia (U.Müll.-Doblies & D.Müll.-Doblies) J.C.Manning & Goldblatt in Taxon 58(1): 102. 2009 – Holotype: O. zebrinellum U.Müll.-Doblies & D.Müll.-Doblies

= O. subgenus Aspasia section Nanaspasia U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert 107: 459. 1996 ≡ O. series Nanaspasia (U.Müll.-Doblies & D.Müll.-Doblies) J.C.Manning & Goldblatt in Taxon 58(1): 103. 2009 – Holotype: O. nanodes F.M.Leight.

= O. subgenus Urophyllon section Helogalum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert 107: 483. 1996 – Holotype: O. flexusoum (Thunb.) U.Müll.-Doblies & D.Müll.-Doblies

Plants small to middle size. Bulb globose, rarely few developed in wet habitats, ending abruptly in an epigeous neck, occasionally surrounded by tubular, transversally banded and epigeous cataphylls; outer tunics membranous or coriaceous; roots usually branched. Leaves numerous, all emerging almost at the same level from the bulb neck, narrow and long, 0·5–6 mm wide (rarely ≤20 mm), linear to cylindrical or lanceolate, often with margins enrolled, rarely flat and broader or even coriaceous and boat-shapped, glabrous or hirsute, spreading or rarely twisted, with entire blunt, ciliate or crispate, sometimes not occurring with the flowers. Inflorescence in raceme cylindrical to subcorymbose, one to three per bulb, dense and many-flowered, elongating with age; pedicels erect or erect-patent; bracts small, membranous, deltoid-aristate or acuminate, auriculate and sometimes denticulate at the base. Flowers erect or spreading, diurnal; tepals almost free to fused at the base up to one-third, thin textured, white to yellowish on the adaxial side and usually with a dark pinkish, reddish, greenish or brownish not defined band on the abaxial side, lanceolate to ovate, acute or obtuse, spreading or reflexed when withering; filaments subulate to linear or ovate-acuminate, expanded at the base, the inners usually wider than the outers, free and adnate (exceptionally connate in O. synadelphicum); ovary elliptical, ovoid or obovoid, contracted in the lower part, ovules biseriate or multiseriate per locule; style filiform; stigma capitate, slightly trigonous or trilobate. Capsule trigonous-elliptical to trilobate-turbinate, with thin walls; usually hidden among the dry and persistent perianth or exposed. Seeds small or large (0·5–5 mm), cuneate, subovoid, angular and irregularly compressed, multiseriate in each locule, with smooth testa.

Chromosome number: 2n = 10, 12 (cf. Obermeyer, 1978).

Number of species: approx. 44 (according to the present study).

Distribution: mainly in the south and east of Africa, not crossing the Sahara desert northwards.

Necessary combinations:

Nicipe adseptentrionesvergentula (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum adseptentrionesvergentulum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 446. 1996 [basion.]

Nicipe aetfatensis (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum aetfatense U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 459. 1996 [basion.]

Nicipe aloiformis (Oberm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum aloiforme Oberm. in Bothalia 12(1): 61. 1976 [basion.]

Nicipe anguina (F.M.Leight. ex Oberm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum anguinum F.M.Leight. ex Oberm. in Bothalia 12(3): 356. 1978 [basion.]

Nicipe annae-ameliae (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum annae-ameliae U.Müll.-Doblies & D.Müll.-Doblies in Willdenowia 11(1): 64. 1981 [basion.]

Nicipe baurii (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum baurii Baker, Fl. Cap. (Harvey) 6(3): 504. 1897 [basion.]

Nicipe britteniae (F.M.Leight. ex Oberm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum britteniae F.M.Leight. ex Oberm. in Bothalia 12(3): 352. 1978 [basion.]

Nicipe campanulata (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum campanulatum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 450. 1996 [basion.]

Nicipe capillaris (J.M.Wood & M.S.Evans) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum capillare J.M.Wood & M.S.Evans in J. Bot. 35: 490. 1897, ‘capillaris’ [basion.]

Nicipe capillifolia (Fourc.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum capillifolium Fourc. in Trans. Roy. Soc. South Africa 21: 80. 1932 [basion.]

Nicipe comptonii (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum comptonii F.M.Leight. in J. S. African Bot. 10: 119. 1944 [basion.]

Nicipe comptum (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum comptum Baker in J. Linn. Soc., Bot. 13: 274. 1872 [basion.]

Nicipe diphylla (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum diphyllum Baker in Bull. Misc. Inform. Kew 1895: 153. 1895 [basion.]

Nicipe dolichopharynx (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum dolichopharynx U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 466. 1996 [basion.]

Nicipe dregeana (Kunth) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum dregeanum Kunth in Enum. Pl. [Kunth] 4: 351. 1843 [basion.]

Nicipe filifolia (Thunb.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Anthericum filifolium Thunb., Prod. Pl. Cap. 1: 62. 1794 [basion.] ≡ O. thunbergianulum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 440. 1996

Nicipe fissurisedula (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum fissurisedulum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 466. 1996 [basion.]

Nicipe flexuosa (Thunb.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Hyacinthus flexuosus Thunb. Prodr. Pl. Cap. 63. 1794 [basion.] = O. ornithogaloides (Kunth) Oberm.

Nicipe gracillima (R.E.Fr.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum gracillimum R.E.Fr. in Acta Horti Berg. 9: 77. 1927 [basion.]

Nicipe graminifolia (Thunb.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum graminifolium Thunb. in Prodr. Pl. Cap. 1: 61. 1794 [basion.]

Nicipe gregoriana (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum gregorianum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 462. 1996 [basion.]

Nicipe hesperantha (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum hesperanthum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 443. 1996 [basion.]

Nicipe juncifolia (Jacq.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum juncifolium Jacq., Pl. Rar. Hort. Schoenbr. 1: 46, t. 90. 1797 [basion.]

Nicipe longicollum (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum longicollum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 462. 1996 [basion.]

Nicipe mater-familias (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum mater-familias U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 444. 1996 [basion.]

Nicipe monophylla (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum monophyllum Baker, Fl. Cap. (Harvey) 6(3): 502. 1897 [basion.]

Nicipe monophylla subsp. eckardtiana (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum monophyllum subsp. eckardtianum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 434. 1996 [basion.]

Nicipe nanodes (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum nanodes F.M.Leight. in J. S. African Bot. 9: 113. 1943 [basion.]

Nicipe natalensis (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum natalense Baker in Bull. Misc. Inform. Kew. 1893: 210. 1893 [basion.]

Nicipe oreogenes (Schltr. ex Poelln.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum oreogenes Schltr. ex Poelln. in Feddes Repert. Spec. Nov. Regni Veg. 54: 24. 1944 [basion.]

Nicipe paludosa (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum paludosum Baker in J. Bot. 12: 366. 1874 [basion.]

Nicipe perdurans (A.P.Dold & S.A.Hammer) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum perdurans A.P.Dold & S.A.Hammer in Bothalia 33(1): 112. 2003 [basion.]

Nicipe perparva (V.Poelln.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum perparvum V.Poelln. in Portugaliae Acta Biol., Sér. B, Sist., 1: 214. 1945 [basion.]

Nicipe petraea (Fourc.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum petraeum Fourc. in Trans. Roy. Soc. South Africa 21: 81. 1932 [basion.]

Nicipe pilosa (L.f.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum pilosum L.f., Suppl. Pl.: 199. 1782 [basion.]

Nicipe pullata (F.M.Leight.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum pullatum F.M.Leight. in J. S. African Bot. 11: 153. 1945 [basion.]

Nicipe rogersii (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum rogersii Baker, Fl. Cap. (Harvey) 6(3): 501. 1897 [basion.]

Nicipe sardienii (van Jaarsv.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum sardienii van Jaarsv. in Bradleya 12: 33. 1994 [basion.]

Nicipe schlecteriana (Schinz.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum schlecterianum Schinz. in Bull. Herb. Boissier 2: 223. 1894 [basion.]

Nicipe sephtonii (Hilliard & B.L.Burtt) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum sephtonii Hilliard & B.L.Burtt in Notes Roy. Bot. Gard. Edinburgh 41(2): 308. 1983 [basion.]

Nicipe synadelphica (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum synadelphicum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 452. 1996 [basion.]

Nicipe tortuosa (Baker) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum tortuosum Baker, Fl. Cap. (Harvey) 6(3): 510. 1897 [basion.]

Nicipe vallis-gratiae (Schltr.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum vallis-gratiae Schltr. in Feddes Repert. Spec. Nov. Regni Veg. 54: 26. 1944 [basion.]

Nicipe verae (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum verae U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 432. 1996 [basion.]

Nicipe zebrinella (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum zebrinellum U.Müll.-Doblies & D.Müll.-Doblies in Willdenowia 11(1): 47. 1981 [basion.]

15. Cathissa

Salisb., Gen. Pl. [Salisbury]: 34. 1866 ≡ Ornithogalum subgenus Cathissa (Salisb.) Baker in J. Linn. Soc., Bot. 13: 261. 1872 ≡ O. section Cathissa (Salisb.) Engl., Nat. Planzenfam. [Engler & Prantl] 2, Abt. 5: 67. 1888 – Holotype: C. cocinna (Salisb.) Speta ≡ O. concinnum Salisb.

Plants small, middle size or large. Bulb ovate; with greyish to brownish, membranous outer tunics. Leaves sole to numerous, green, narrowly lanceolate to long tapering, glabrous and occurring with the flowers, sometimes with a long terminal terete appendix. Inflorescence racemose or subspicate, longer than wide; pedicels erect-patent in flower and adpressed to stem in fruit; bracts large, triangular to lanceolate, membranous or papery, petaloid and auriculate; bracteoles absent. Flowers white, suberect, patent or slightly nodding; tepals free, completely white, widely ovate or obovate, rarely with small greenish dots at the apex of the abaxial side; filament subulate to tapering; ovary green or yellowish, ovoid-fusiform; style white and filiform, or greenish in the lower part and white at the apex; stigma slightly capitate, trigonous and glandular-papillate. Capsule ovate-lanceolate, oblong and triangular with blunt edges section. Seeds semicircular, irregular or angular, polygonal or apiculate, biseriate or multiseriate per locule, with granulate or puzzle-like testa.

Chromosome number: 2n = 32, 34, 36 (cf. Speta, 1998a).

Number of species: 3 (cf. Martínez-Azorín et al., 2007).

Distribution: Spain, Portugal and Morocco.

Necessary combinations:

Cathissa unifolia (L.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Scilla unifolia L., Sp. Pl.: 309 (1753) [basion.] ≡ C. unifolia (L.) Salisb., Gen. Pl. [Salisbury]: 34 (1866), comb. inval. (according to art. 33·1, ICBN) ≡ C. broteroi (M.Laínz) Speta in Stapfia 75: 171 (2001)

16. Melomphis

Raf., Fl. Tell. 2: 21. 1837 ≡ Caruelia Parl., Nouv. Gen. Sp. Monocot.: 21. 1854 ≡ Ornithogalum subgenus Caruelia (Parl.) Baker in J. Linn. Soc. (Bot.) 13: 259. 1872 ≡ O. section Caruelia (Parl.) Engl., Nat. Planzenfam. [Engler & Prantl] 2, Abt. 5: 67. 1888 ≡ Myanthe Salisb., Gen. Pl. [Salisbury]: 34. 1866 ≡ O. section Melomphis (Raf.) J.C.Manning & Goldblatt in Taxon 58(1): 103. 2009 – Lectotype: M. arabica (L.) Raf. (cf. Speta, 2001) ≡ C. arabica (L.) Parl. ≡ M. arabica (L.) Salisb. ≡ O. arabicum L.

Plants large. Bulb ovoid, with membranous outer tunics. Leaves green, lanceolate-oblong, wide, long tapering. Inflorescence corymbose or subcorymbose-pyramidal, bracts membranous, ovate-triangular, wide and petaloid (white with greenish apex); bracteoles absent. Flowers white, showy; tepals white all over, wide, ovate or obovate, withering soon after anthesis; filaments white, narrowly triangular; ovary globose and shining black; style filiform and white or blackish below; stigma trigonous, glandular. Capsule subglobose, trigonous, with truncate apex. Seeds blackish, polygonal or irregularly compressed, with tuberculate or papillate testa.

Chromosome number: 2n = 28, 51 (cf. Speta, 1998a)

Number of species: 3 (according to the present study).

Distribution: Mediterranean basin, extending to the Middle East.

Necessary combinations:

Melomphis melanogyna (Cullen) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum melanogynum Cullen in Notes Roy. Bot. Gard. Edinburgh 27: 309. 1967 [basion.]

Melomphis persica (Hausskn. ex Bornm.) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum persicum Hausskn. ex Bornm. in Beih. Bot. Centralbl. 24(2): 102. 1908 [basion.]

17. Loncomelos

Raf., Fl. Tell. 2: 24. 1837 – Lectotype: L. pyrenaicus (L.) L.D.Hrouda ex Holub (cf. Speta, 2001) ≡ Ornithogalum pyrenaicum L.

= Celsia Heist. ex Fabr., Enum. (ed. 2) [Fabr.]: 22. 1763, pro parte [O. latifolium L.], nom. illeg. [non Celsia L., Sp. Pl.: 621. 1753; Scrophulariaceae]

= Beryllis Salisb., Gen. Pl. [Salisbury]: 33. 1866 ≡ O. subgenus Beryllis (Salisb.) Baker in J. Linn. Soc. (Bot.) 13: 260. 1872 ≡ O. section Beryllis (Salisb.) Engl., Nat. Planzenfam. [Engler & Prantl] 2, Abt. 5: 67. 1888 ≡ O. subgenus Spetagalum U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5–6): 520. 1996 [nom. superfl.] – Lectotype: O. pyrenaicum L. (cf. Stearn, 1983)

Eustachys Salisb., Gen. Pl. [Salisbury]: 33. 1866

Bulb with outer tunics membranous, sometimes with long neck. Leaves tapering, wide, long, glabrous, sometimes not occurring with the flowers. Inflorescence racemose, narrow, much longer than wide; pedicels erect-patent in flower and erect and appressed to the stem in fruit; bracts membranous, deltoid-aristate, usually shorter than the pedicels; bracteoles absent. Flowers white or yellowish; tepals with a greenish longitudinal band only evident on the abaxial side; filaments flattish and expanded at the base; ovary green, ovate to spherical; style white and filiform; stigma puntiform and slightly trigonous. Capsule ovate-lanceolate, triangular with blunt edges in section. Seeds polygonal or irregularly compressed, biseriate per locule, with tuberculate, papillate or rugose testa.

Chromosome number: 2n = 16, 18, 22, 24, 26, 28, 32, 36, 54 (cf. Speta, 1998a).

Number of species: approx. 20 (cf. Speta, 1998a, 2001).

Distribution: Mediterranean basin.

Observations: when the name O. subgenus Beryllis (Salisb.) Baker was published it was nomenclaturally superfluous, since the type of Urophyllon was included in it (Art. 52·1, 52·2e; McNeill et al., 2006). Consequently it was a synonym of O. subgenus Urophyllon (Salisb.) Baker; though it was not illegitimate (Art. 52·3). When both subgenera are recognized, both names recover their validity and they can be used, each one with its corresponding circumscription. In this way, the new name O. subgenus Spetagalum, proposed by Müller-Doblies and Müller-Doblies (1996) to replace the supposedly illegitimate O. subgenus Beryllis, is superfluous, since these authors noted explicitly that this new name should be used exclusively for the Mediterranean plants, keeping the South African ones in O. subgenus Urophyllon (Urophyllon sensu stricto).

18. Honorius

Gray in Nat. Arr. Brit. Pl. 2: 177. 1821 ≡ Albucea Rchb., Fl. Germ. Excurs.: 109. 1830 ≡ Myogalum Link in Handb. 1: 163. 1829 ≡ Ornithogalum subgenus Myogalum (Link) Baker in J. Linn. Soc. (Bot.) 13: 260. 1872 ≡ O. section Myogalum (Link) Engl., Nat. Planzenfam. [Engler & Prantl] 2, Abt. 5: 67. 1888 ≡ Syncodium Raf., Fl. Tell. 2: 22. 1837 ≡ Brizophile Salisb., Gen. Pl. [Salisbury]: 34. 1866 – Holotype: H. nutans (L.) Gray ≡ O. nutans L.

Plants middle size. Bulb with brown or whitish membranous outer tunics. Leaves linear and tapering to lanceolate or oblong, occurring with the flowers, glabrous, sometimes with a longitudinal greyish band. Inflorescence racemose; pedicels erect-patent in flower, erect to reflexed in fruit; bracts lanceolate or ovate-lanceolate, membranous or papery; bracteoles absent. Flowers suberect or patent, campanulate at the anthesis; tepals free, ovate-lanceolate, erect, white with a wide longitudinal green band only visible on the abaxial side; filaments lanceolate to tapering, at least outers wider at the apex and forming two erect lobes accompanying the anther; ovary green, ovate, trigonous, with six blunt edges; style white and filiform; stigma puntiform slightly trigonous and glandular. Capsule ovate or oblong, sometimes truncate at the apex, with six weak ribs in section, mostly nodding. Seeds globose, with prominent sinuose-reticulate testa.

Chromosome number: 2n = 14, 28, 30, 35, 42 (cf. Speta, 1998a).

Number of species: 8 (cf. Speta, 1998a; Holub, 1976).

Distribution: south-eastern Europe, western Turkey and Middle East (cf. Speta, 1998a).

Observations: O. pamphylicum was described by Düşen and Sümbül (2002) as a member of O. subg. Myogalum, close to O. nutans. However, the former species lacks some of the morphological traits defining Honorius, such as the filaments lobed, winged or toothed in the upper part, or the subacute to truncate capsule. Therefore it is here excluded from Honorius. New data are needed for a more accurate placement of O. pamphylicum in Ornithogaleae.

Necessary combinations:

Honorius demirizianus (H.Malyer & Koyuncu) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum demirizianum H.Malyer & Koyuncu in Turkish J. Bot. 13(2): 301. 1989 [basion.]

Honorius chetikianus (Uysal, Ertugrul & Dural) Mart.-Azorín, M.B.Crespo & Juan, comb. nov.Ornithogalum chetikianum Uysal, Ertugrul & Dural in Bot. J. Lin. Soc. 148: 501. 2005 [basion.]

19. Ornithogalum L., Sp. Pl.: 306. 1753 ≡ O. subgenus Heliocharmos Baker in J. Linn. Soc. (Bot.) 13: 258. 1872 ≡ O. section Heliocharmos (Baker) Engl., Nat. Planzenfam. [Engler & Prantl] 2, Abt. 5: 67. 1888 – Lectotype: O. umbellatum L. (cf. Stearn, 1983).

= Celsia Heist. ex Fabr., Enum. (ed. 2). [Fabr.].: 22. 1763, pro parte, nom. illeg. [non Celsia L., Sp. Pl.: 621. 1753; Scrophulariaceae] – Lectotype: O. umbellatum L.

Plants small to middle size. Bulb with brown or whitish membranous outer tunics, sometimes with sessile or pediculate bulbils. Leaves numerous, linear and tapering to lanceolate or oblong, sometimes not occurring with the flowers, glabrous or rarely ciliate, sometimes with a longitudinal white band. Inflorescence corymbose or racemose corymbose, a little longer than wide; pedicels erect-patent in flower and patent or sometimes deflexed in fruit; bracts large, lanceolate or ovate-lanceolate, membranous or papery; bracteoles absent. Flowers suberect, erect or patent, diurnal; tepals free, ovate-lanceolate, white with a wide longitudinal green band only visible by the abaxial side; filaments free and flat, lanceolate to tapering; ovary green, ovate or obovate and truncate at the apex, deeply trilobate, with six evident ribs; style white and filiform; stigma punctiform slightly trigonous and glandular. Capsule obovate or oblong, usually truncate at the apex, deeply trilobate with six evident ribs in section. Seeds globose, biseriate per locule, with sinuose and prominent reticulate testa.

Chromosome number: 2n = 12, 14, 16, 18, 20, 22, 24, 26, 28, 32, 36, 45, 52, 54, 72, 90, 108 (cf. Speta, 1998a).

Number of species: about 50 (cf. Speta, 1998a).

Distribution: Europe, south-west Asia (Middle East to Afghanistan), and northern Africa.

Incertae sedis

Ornithogalum toxicarium C.Archer & R.H.Archer in S. Afr. J. Bot. 65: 431. 1999.

This peculiar plant was described from karoo shrublands and dry grassland in South Africa and Namibia. Its peculiar floral, fruit and seed features do not match any of the accepted genera of Ornithogaloideae. Further studies, including molecular data, are needed to resolve the taxonomic position of this remarkable taxon. In the meantime, we prefer not to place it in the proposed arrangement.

SUPPLEMENTARY DATA

Supplementary data are available online at www.aob.oxfordjournals.org and consist of the following tables. Table S1: Characters and character states for the morphological study. Table S2: Morphological data matrix. Table S3: GenBank accession numbers for samples included in the phylogenetic analyses of the Hyacinthaceae, mainly the subfamily Ornithogaloideae.

ACKNOWLEDGEMENTS

This work was supported by the Ministerio de Educación y Ciencia (Spanish Government) (grant number AP2002-0214) and the Fundación Ramón Areces (Spain). We thank Laco Mucina (Curtin University of Technology, Perth) for his interesting biogeographical comments and corrections and additions to the early manuscript. We are grateful to John Manning (South African National Biodiversity Institute, Kirstenbosch) and Félix Forest (Jodrell Laboratory, Royal Botanic Gardens, Kew) for sharing their unpublished plastid sequences. Ma Dolores Lledó (Jodrell Laboratory, Royal Botanic Gardens, Kew) helped with laboratory procedures. Katherine Challis (Royal Botanic Gardens, Kew) kindly supplied information of protologues of several taxa. Lorenzo Peruzzi (Università degli Studi di Pisa) and Ma Montserrat Martínez-Ortega (Universidad de Salamanca) sent important samples for the phylogenetic analyses. Stephen L. Jury (University of Reading) sent pictures and vouchers of B. amoena. Nigel P. Barker (Department of Botany, Rhodes University, Grahamstown) and Tony P. Dold (Albany Museum, Schönland Herbarium, Grahamstown) provided valuable assistance with some South African genera. Two anonymous referees and the handling editor, Anne Brysting, made constructive criticisms that improved the text.

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