Martínez-Azorín Et Al. 2023. A Generic Monograph of Urgineoideae. Phytotaxa 610 (1), 1-143 REDUCED COVER

Phytotaxa 610
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
31 August 2023
A generic monograph of the Hyacinthaceae

subfamily Urgineoideae
MARIO MARTÍNEZ-AZORÍN, MANUEL B. CRESPO, MARÍA ÁNGELES

ALONSO-VARGAS, MICHAEL PINTER, NEIL R. CROUCH, ANTHONY
P. DOLD, LADISLAV MUCINA, MARTIN PFOSSER
& WOLFGANG WETSCHNIG
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Phytotaxa 610 (1): 001–143 ISSN 1179-3155 (print edition)
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Monograph PHYTOTAXA
Copyright © 2023 Magnolia Press ISSN 1179-3163 (online edition)
https://doi.org/10.11646/phytotaxa.610.1.1
PHYTOTAXA
610
A generic monograph of the Hyacinthaceae subfamily Urgineoideae
MARIO MARTÍNEZ-AZORÍN1,9*, MANUEL B. CRESPO1,10, MARÍA ÁNGELES ALONSO-

VARGAS1,11, MICHAEL PINTER2,12, NEIL R. CROUCH3,4,13, ANTHONY P. DOLD5,14, LADISLAV
MUCINA6,7,15, MARTIN PFOSSER8,16 & WOLFGANG WETSCHNIG2,17
1
Depto. Ciencias Ambientales y Recursos Naturales (dCARN), Universidad de Alicante, P.O. Box 99, ES-03080 Alicante, Spain.
2
Institute of Biology, NAWI Graz, Division Plant Sciences, Karl-Franzens University Graz, Holteigasse 6, A-8010 Graz, Austria.
3
BRAM, South African National Biodiversity Institute, P.O. Box 52099, Berea Road 4007, South Africa.
4
School of Chemistry and Physics, University of KwaZulu-Natal, Durban 4041, South Africa.
5
Selmar Schonland Herbarium, Department of Botany, Rhodes University, Makhanda, 6140, South Africa.
6
Harry Butler Institute, Murdoch University, 90 South Street, Building 390, Murdoch WA 6150, Perth, Australia.
7
Dept. of Geography & Environmental Studies, Stellenbosch University, Private Bag X1, Matieland 7602, Stellenbosch, South Africa.
8
Biocenter Linz, J.-W.-Klein-Str. 73, A-4040 Linz, Austria.
9
E-mail: � mmartinez@ua.es; ORCID: https://orcid.org/0000-0002-2605-9575
10
E-mail: � crespo@ua.es; ORCID: https://orcid.org/0000-0002-3294-5637
11
E-mail: � ma.alonso@ua.es; ORCID: https://orcid.org/0000-0003-3768-9203
12
E-mail: � michael.pinter@uni-graz.at; ORCID: https://orcid.org/0000-0002-6055-6989
13
E-mail: � N.Crouch@sanbi.org.za; ORCID: https://orcid.org/0000-0002-4938-5840.
14
E-mail: � t.dold@ru.ac.za; ORCID: https://orcid.org/0000-0002-9497-7503
15
E-mail: � ladislav.mucina@murdoch.edu.au; ORCID: https://orcid.org/0000-0003-0317-8886
16
E-mail: � martin.pfosser@ooelkg.at; ORCID: https://orcid.org/0000-0003-2050-4997
17
E-mail: � wolfgang.wetschnig@uni-graz.at; ORCID: https://orcid.org/0000-0002-9245-029X
*Author for correspondence
Magnolia Press
Auckland, New Zealand
Accepted by Ronell R. Klopper: 4 Jul. 2023; published: 31 Aug. 2023

MARIO MARTÍNEZ-AZORÍN, MANUEL B. CRESPO, MARÍA ÁNGELES ALONSO-VARGAS,
MICHAEL PINTER, NEIL R. CROUCH, ANTHONY P. DOLD, LADISLAV MUCINA, MARTIN
PFOSSER & WOLFGANG WETSCHNIG
(Phytotaxa 610)
143 pp.; 30 cm.
31 August 2023
ISBN 978-1-77688-860-3 (paperback)
ISBN 978-1-77688-861-0 (Online edition)
FIRST PUBLISHED IN 2023 BY

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• Phytotaxa 610 (1) © 2023 Magnolia Press MARTÍNEZ-AZORÍN Et al.

Table of contents
Abstract...........................................................................................................................................................................................................3
Introduction....................................................................................................................................................................................................4
Materials and methods..................................................................................................................................................................................6
Results and discussion...................................................................................................................................................................................7
Previous taxonomic proposals and phylogenetic relationships..................................................................................................................7
Comments on morphological characters.....................................................................................................................................................8
TAXONOMIC TREATMENT....................................................................................................................................................................18
Identification key of genera in Urgineoideae..............................................................................................................................................18
A synopsis of the accepted genera................................................................................................................................................................20
Aulostemon Mart.-Azorín et al. ...................................................................................................................................................................20
Austronea Mart.-Azorín et al........................................................................................................................................................................23
Boosia Speta..................................................................................................................................................................................................29
Bowiea Harv. ex T.Moore & Mast. ...............................................................................................................................................................33
Drimia Jacq. ex Willd. ..................................................................................................................................................................................35
Ebertia Speta.................................................................................................................................................................................................41
Fusifilum Raf. ..............................................................................................................................................................................................43
Geschollia Speta ...........................................................................................................................................................................................47
Indurgia Speta ..............................................................................................................................................................................................51
Iosanthus Mart.-Azorín et al. . .....................................................................................................................................................................56
Ledurgia Speta .............................................................................................................................................................................................61
Litanthus Harv. . ...........................................................................................................................................................................................62
Mucinaea M.Pinter et al. . ............................................................................................................................................................................63
Rhadamanthopsis (Oberm.) Speta ...............................................................................................................................................................64
Rhadamanthus Salisb. .................................................................................................................................................................................67
Rhodocodon Baker .......................................................................................................................................................................................72
Sagittanthera Mart.-Azorín et al. . ...............................................................................................................................................................78
Schizobasis Baker . .......................................................................................................................................................................................80
Sekanama Speta............................................................................................................................................................................................84
Spirophyllos Mart.-Azorín et al. ..................................................................................................................................................................87
Squilla Steinh. ..............................................................................................................................................................................................90
Striatula M.Pinter et al. ................................................................................................................................................................................96
Tenicroa Raf. ................................................................................................................................................................................................97
Thuranthos C.H.Wright..............................................................................................................................................................................103
Triandra Mart.-Azorín et al. ......................................................................................................................................................................107
Urginavia Speta...........................................................................................................................................................................................108
Urginea Steinh. . .........................................................................................................................................................................................114
Urgineopsis Compton..................................................................................................................................................................................116
Vera-duthiea Speta......................................................................................................................................................................................119
Zingela N.R.Crouch et al. ..........................................................................................................................................................................122
Zulusia Mart.-Azorín et al. ........................................................................................................................................................................124
Incertae sedis (provisionally unplaced within Urgineoideae)...................................................................................................................127
Names excluded from Urgineoideae..........................................................................................................................................................129
Acknowledgements....................................................................................................................................................................................131
References...................................................................................................................................................................................................131
Abstract
Taxonomy and systematics and specifically the generic circumscription of the Hyacinthaceae subfamily Urgineoideae have
been a matter of controversy in recent decades. Widely contrasting taxonomic treatments have been proposed, ranging from
systems comprising only two genera (including a morphologically very variable Drimia covering nearly the whole subfami-
ly) to about 20 genera based mainly on preliminary phylogenetic studies supporting recognition of traditionally accepted and
morphologically distinct genera such as Litanthus, Schizobasis, Tenicroa, and Thuranthos. All previous studies covered only
a limited diversity and distribution of the subfamily, precluding comprehensive insight into the evolution of the diversity
within the subfamily. Based on detailed morphological, phylogenetic, and biogeographic studies in Urgineoideae, covering
its whole distribution range and most of the currently known species, we present a new taxomic treatment of the subfamily
focusing on the circumscription of genera. Our analyses revealed 31 phylogenetic lineages corroborating unique syndromes
of morphological characters linked to well-defined biogeographic patterns. These lineages are interpreted as well-defined
genera. An identification key to the genera is provided to facilitate further taxonomic work in Urgineoideae and morpho-
logical descriptions of the accepted genera are presented. Two new genera, Spirophyllos gen. nov. and Zulusia gen. nov., are
described to accommodate newly revealed phylogenetic lineages from northwestern and southeastern Africa, respectively.
Further, a complete list of taxa included in the accepted genera is presented, providing details on typification, and the 50 new
combinations, and one new name required to appropriately accomodate taxa in the new taxonomic treatment.
Key words: Asparagaceae, distribution, ecology, Scilloideae, taxonomy, typification, Urgineae.
A generic monograph of the Urgineoideae Phytotaxa 610 (1) © 2023 Magnolia Press •
Introduction
Hyacinthaceae sensu APG (2003) include about 1000 species of bulbous plants (Pfosser & Speta 1999), which
are mainly distributed throughout Europe, Africa, and Asia, with a single genus found in South America. The poor
knowledge of this group of plants in several countries in Africa and Asia, exacerbated by the lack of modern taxonomic
revisions in many regions, suggests a large taxonomic treasure trove still to be discovered here.
Based on morphological, anatomical, chemical, and molecular studies (Speta 1998a, 1998b, Pfosser & Speta
1999, Manning et al. 2004), four monophyletic subfamilies can be clearly recognised within Hyacinthaceae, namely
Hyacinthoideae, Ornithogaloideae, Urgineoideae, and Oziroëoideae. Alternatively, this monophyletic family is
treated as Asparagaceae subfam. Scilloideae by APG (2009, 2016) and Chase et al. (2009), and the subfamilies listed
above are treated as tribes, namely Hyacintheae, Ornithogaleae, Oziroëeae, and Urgineeae. We favour the concept of
Hyacinthaceae, supported by unequivocal morphological and molecular evidence.
The taxonomy and systematics of Hyacinthaceae has been a matter of controversy in recent decades, especially
regarding the conceptualisation and circumscription of genera. A prime example of this controversy is subfamily
Ornithogaloideae, in which recent studies suggest either the recognition of only one genus—Ornithogalum Linnaeus
(1753: 306)—showing an extremely variable morphology (Manning et al. 2004), or the recognition of 20 monophyletic
genera (including Igidia Speta 1998b: 70) that are well supported by morphology and biogeography (Martínez-Azorín
et al. 2011a). However, when sufficiently informative plastid and nuclear DNA regions are included in the phylogenetic
analyses, consistent morphological elements become fully congruent with clades, and these can be accepted at generic
rank, as shown by Martínez-Azorín et al. (2011a) for Ornithogaloideae and Martínez-Azorín et al. (2023a) for
Urgineoideae.
Hyacinthaceae subfamily Urgineoideae is distributed in Africa, Madagascar, the Mediterranean region, southwestern
Asia, and reaching Thailand in southeastern Asia. The group originated about 48 mya (Buerki et al. 2012, Ali et al.
2013) in Southern Africa where it shows the highest taxonomic diversity. Secondary centres of diversification occur
in the Mediterranean Basin and southwestern Asia. These emerged as a result of putative dispersal events involving
arid and the high-elevation corridors linking the Middle East (and Horn of Africa) with the arid southwestern parts
of Southern Africa (Verdcourt 1969, Thulin 1994, Del Hoyo & Pedrola-Monfort 2006, Devos et al. 2010, Martínez-
Azorín et al. 2010).
Taxa in Urgineoideae are characterised by the spurred bracts, at least those at the inflorescence base, the brittle
seed testa easily detachable from the endosperm, and by the secondary natural products of the bulbs. Mulholland et al.
(2013) reviewed the phytochemistry and related bioactivity of Hyacinthaceae, including all reports on the Urgineoideae
to that date. In agreement with Speta (1998a) and Pohl et al. (2000) they reported that compounds are, for the most part,
subfamily restricted. Cardenolides and steroidal glycosides characterise Ornithogaloideae, homoisoflavanones and
spirocyclic nortriterpenoids are typical of Hyacinthoideae, and bufadienolides are highly characteristic of Urgineoideae.
Bufadienolide cardiotoxicoses in livestock and man have commonly been observed when Urgineoideae plants were
consumed throughout its geographic range. In consequence, Urgineoideae have been analysed extensively, resulting in
the isolation and identification of a large variety of bufadienolides and their glycosides, with differences found both in
quantity and type of such compounds even within single species (Mulholland et al. 2013).
Some authors (e.g. Stearn 1978, Manning et al. 2004, Manning & Goldblatt 2018) accept about 100–120 species
in Urgineoideae, although the number of species is likely to be considerably higher due to the large deficiencies in our
knowledge of this group throughout its wide distribution range. Our work accepts about 220 species and represents the
most comprehensive study in this group so far.
The oldest genus in Urgineoideae is Drimia Jacq. ex Willdenow (1799: 165) that originally included five species
from South Africa, namely D. elata Jacq. ex Willdenow (1799: 165), D. pusilla Jacq. ex Willdenow (1799: 165),
D. media Jacq. ex Willdenow (1799: 166), D. undulata Jacq. ex Willdenow (1799: 166), and D. ciliaris Jacq. ex
Willdenow (1799: 165). Drimia is characterised by a distinct and homogeneous flower morphology showing a basal
cylindrical perigone tube with long and reflexed free portion of the tepals, and stamens long and connivent to the style
[excluding D. undulata that currently belongs to Ledebouria Roth (1821: 194)].
Urginea Steinheil (1834: 321) was the second genus described in the subfamily, differring from Drimia by
spreading and free tepals. This genus comprised then seven species with almost free tepals and variable morphology
and distribution. Four of those species, U. anthericoides (Poiret 1789: 149) Steinheil (1834: 328), U. fragrans (Jacquin
1797: 45) Steinheil (1834: 330), U. filifolia (Jacquin 1794: 18) Steinheil (1834: 330), and U. exuviata (Jacquin 1794:
18) Steinheil (1834: 330), are currently included in Tenicroa Rafinesque (1837: 52)—a genus endemic to South Africa

and Namibia (Pinter et al. 2020). The remaining three species are endemic to the Mediterranean Basin, where two
of them, U. undulata (Desfontaines 1792: 161) Steinheil (1834: 330) and U. scilla Steinheil (1834: 321), belong to
Squilla Steinheil (1836: 276) in the sense of the present work, and the third, U. fugax Steinheil (1834: 328), is the
type of the genus Urginea (cf. Adamson 1942). Rafinesque (1837) described the genera Tenicroa Rafinesque (1837:
52) and Fusifilum Rafinesque (1837: 27) to accommodate certain groups of species from South Africa with distinct
sexual and vegetative characters—those with basal zebrine cataphylls having transversally raised bands and flowers
with deflexed and curved styles were placed in Tenicroa, and those with fusiform and minutely papillate filaments
and a white gynoecium in Fusifilum. Litanthus Harvey (1844: 314) is another remakable genus originally described as
monotypic for a distinct minute species with small, tubular nodding flowers, and with tepals only spreading as short
apical lobes. These characters underpinned the recognition and acceptance of this genus in most floras in Southern
Africa. A second species of this genus was described only recently (Manning et al. 2013, Martínez-Azorín et al.
2015). Salisbury (1866) described Rhadamanthus Salisbury (1866: 37) to include a taxon characterised by nodding,
campanulate to urceolate flowers with stamens connivent to the gynoecium, and anthers dehiscing by apical pores or
slits. Nordenstam (1970) revised this genus and accepted nine species in Southern Africa sharing anther dehiscence.
Bowiea Harv. ex Moore & Masters (1866: 971) and Schizobasis Baker (1873a: 105) are two distinct genera, which
share branched inflorescences, although they differ in flower and vegetative characters. Both genera are remarkable
in morphology, are readily segregated from other members in the subfamily and have been widely accepted in all
floras in both Southern and East Africa. Baker (1881) described Rhodocodon Baker (1881: 280) for a peculiar species
from Madagascar with nodding, campanulate flowers and included stamens. At present, this genus includes 19
species endemic to Madagascar, following the revision and related contributions by Knirsch et al. (2015, 2016, 2019).
Thuranthos Wright (1916: 233) represents another distinct genus characterised by a syndrome including nodding,
nocturnal flowers and sigmoid, basally expanded filaments, among other characters. It is clearly different from most
Urgineoideae genera, and accordingly widely accepted in traditional taxonomic works. Compton (1930) described
Urgineopsis Compton (1930: 107) to accomodate some South African species with a campanulate basal tube of the
perigone and spreading perigone lobes and filaments. Finally, Obermeyer (1980a) described Rhadamanthus subgenus
Rhadamanthopsis (Obermeyer 1980a: 137), mainly differing from typical Rhadamanthus by nodding, campanulate
flowers with the anthers dehiscing along the whole length, not only by apical pores or slits. Speta (1998b) subsequently
accepted this group as the distinct genus Rhadamanthopsis (Obermeyer 1980a: 137) Speta (1998b: 74).
Most of the taxa cited above were traditionally accepted at their original generic rank by the majority of the
Urgineoideae taxonomists in the course of the last two centuries (Willdenow 1799, Steinheil 1834, 1836, Rafinesque
1837, Harvey 1844, Salisbury 1866, Moore & Masters 1866, Hooker 1867, 1872, Baker 1871, 1873a, 1874a, 1876,
1881, 1897, 1898, Bentham & Hooker 1883, Engler 1888, Wright 1916, Compton 1930, Krause 1930, Dyer 1934,
Hutchinson 1934, Perrier de la Bâthie 1938, Sölch et al. 1970, Merxmüller 1970, Nordenstam 1970, Stirton 1976,
Jessop 1977, Obermeyer 1980a, 1980b, 1980c, 1981a, Roessler 1987, Stedje 1997, Speta 1998a, 1998b, Snijman et
al. 1999, Müller-Doblies et al. 2001). However, this situation changed with the advent of early phylogenetic studies in
Urgineoideae. These generated considerable controversy regarding the generic circumscriptions (Stedje 1998, 2001a,
2001b, Pfosser & Speta 1999, 2001, Manning et al. 2004, 2009, Pfosser et al. 2012, Manning & Goldblatt 2013,
Martínez-Azorín et al. 2011a, 2023a).
Speta (1998a, 1998b, 2001), based on comprehensive studies combining morphological, anatomical, and
molecular data, described the monophyletic genera Boosia Speta (2001: 168), Charybdis Speta (1998b: 58), Duthiea
Speta (2001: 170), Ebertia Speta (1998b: 65), Geschollia Speta (2001: 169), Indurgia Speta (2001: 169), Ledurgia
Speta (2001: 168), Rhadamanthopsis, Sekanama Speta (2001: 168), and Urginavia Speta (1998b: 86). Recently, seven
new genera were described in Urgineoideae, namely Sagittanthera Mart.-Azorín, M.B.Crespo, A.P.Dold & Van Jaarsv.
in Martínez-Azorín et al. (2013a: 46), Mucinaea M.Pinter, Mart.-Azorín, U.Müll.-Doblies, D.Müll.-Doblies, Pfosser
& Wetschnig in Pinter et al. (2013: 296), Aulostemon Martínez-Azorín et al. (2017: 288), Austronea Mart.-Azorín,
M.B.Crespo, M.Pinter & Wetschnig in Martínez-Azorín et al. (2018a: 105), Zingela Crouch et al. (2018: 36), Striatula
M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Pinter et al. (2019: 93), and Triandra Martínez-Azorín et al.
(2021: 66). All are based on their distinct morphologies and isolated positions in the phylogenetic tree (Martínez-
Azorín et al. 2023a).
In Urgineoideae, the genera with distinctly connate tepals, such as Drimia, Litanthus, Rhadamanthus,
Rhadamanthopsis, and Rhodocodon, are easily recognised and have been traditionally accepted. However, many other
species described as Urginea or Drimia sensu lato from Southern Africa show unspecialised flowers in which tepals
vary from free to shortly connate (Baker 1897, Adamson 1942, Huber 1969, Jessop 1977, Hilliard & Burtt 1982,
Stedje 1987). This led to the rejection of some genera (e.g. Urgineopsis) in the subfamily showing shortly connate
tepals (Jessop 1977, Manning et al. 2004, Manning & Goldblatt 2013, 2018). This approach resulted in unsatisfactory
circumscription of genera that included species with distant ancestral relationships and different morphological
syndromes. Consequently, artificial groups have been defined obscuring the true evolutionary relationships among
species. This issue can be resolved using phylogenetic analyses, as shown by Martínez-Azorín et al. (2011a, 2023a),
when the clades revealed by molecular-phylogenetic studies corroborate well-defined morphological syndromes
to characterise distinct genera. We consider it as crucial to discriminate among groups in this subfamily. Particular
attention must be paid to variation in flower morphology, and characters such as shape and disposition of filaments and
anthers, colour and shape of ovary, style proportion and disposition as well as fruit and seed morphology.
It is the aim of the current work to provide a taxonomic alternative to an overly broad Drimia concept to render
a better understanding of generic boundaries in the Urgineoideae based on clades that are easily recognised given
their distinct morphology and biogeography. Our analytic treatment in Urgineoideae is similar to that accepted in
Hyacinthoideae with ca. 38 genera distributed across three continents, with eleven of those accepted by Manning et al.
(2004) for Southern Africa alone, or in the global assessment of Ornithogaloideae by Martínez-Azorín et al. (2011a),
which recognised 20 genera (including Igidia). As a result, 31 genera and 221 species (plus some provisionally
unplaced species) are here accepted in Urgineoideae, including two new genera (Spirophyllos gen. nov. and Zulusia
gen. nov.) to accommodate species in clades with distinct syndromes of morphological characters, unique ancestral
history and biogeography. Complete morphological descriptions are presented for all these genera, including data on
karyology and distribution. All accepted species are listed for each genus, including their typification details. Necessary
nomenclatural combinations are also presented to accommodate species in the newly circumscribed genera, and others
that require designation. Further, an identification key is provided including all genera in subfamily Urgineoideae, to
facilitate future work on this group.
Materials and methods
Detailed morphological studies were undertaken on both field specimens and on cultivated plants maintained in the
living collection at the University of Alicante, Karl-Franzens University of Graz, and Biocenter Linz. Terminology
used for descriptions of species of Urgineoideae follows Martínez-Azorín et al. (2007, 2009). Herbarium specimens
from the herbaria ABH, B, BLFU, BM, BNRH, BOL, E, EA, G, GRA, GZU, HAL, HBG, HSMC, K, L, LD, LINN,
LISU, LUB, NH, NU, NY, M, MO, MPU, NBG, P, PRE, S, SAM, TCD, UPS, W, WIND, WU, Z, ZSS, and ZT
(acronyms according to Thiers 2023) were studied. Authors of the cited taxa follow IPNI (2023). Orthography of
geographical names and grid-number system follows Leistner & Morris (1976), also considering recent geographic
name changes and new geographic terms introduced in South Africa. Nomenclatural issues follow the regulations of
the Shenzhen Code -ICN- (Turland et al. 2018). Measurements of tepals, stamens, and ovaries were taken on fresh
material and complemented with data obtained from pressed herbarium material.
Our phylogenetic analyses included 293 samples of Urgineoideae belonging to ca. 160 species from all previously
recognised taxonomic groups in Urgineoideae, covering the full range of morphological and taxonomic variation
from its entire distribution range throughout Europe, Africa, and southwestern Asia. Four plastid DNA regions (trnL
intron, trnL-F spacer, matK, and trnCGCA-ycf6 intergenic region) and a nuclear region (Agt1) were included in the
phylogenetic analyses as specified in Martínez-Azorín et al. (2023a).
A selection of 40 discrete morphological characters were coded as detailed in Martínez-Azorín et al. (2023a). A
matrix with the 40 coded characters was built for all the 293 samples included in the phylogenetic analyses as shown
in Table 2 in Martínez-Azorín et al. (2023a). These data were incorporated in Mesquite v. 3.61 (Madison & Madison
2019) to map the morphological characters on the phylogenetic trees. All 40 characters were traced individually to
evaluate apomorphies and ancestral states for the obtained clades (Supplementary Material S13 in Martínez-Azorín et
al. 2023a). A considerable degree of homoplasy was observed in some morphological characters, indicating evolutive
convergence. Other characters were retrieved as apomorphic and restricted to certain clades-genera, allowing clear
characterization of genera when considered in combination as morphological syndromes (Martínez-Azorín et al.
2023a).
Approximate distribution ranges of the accepted genera are based on our own collections and the study of herbarium
material, and are shown in the figures depicting each genus. The distribution ranges were linked to the biogeographic
regions of Takhtajan (1986). We have subdivided the original Takhtajan (1986) Sudano-Zambezian Region using in
principle his Subregions, with the further subdivision of the Zambezian Subregion into three informal ‘Sections’:

Northern (precipitation-richer, transitional towards the tropical regions of Central Africa), Eastern (precipitation
rich, including series of high-elevation mountain ranges), and Southern (drier, home to arid and semi-arid savanna)
(see also Martínez-Azorín et al. 2023a). This refinement of the Zambezian Subregion allowed for better resolution
of biogeographical aspects within Southern Africa, a major centre of endemism of subfamily Urgineoideae. Hence,
the following resulting phytochoria were used to assist comparative analyses of the distribution of genera: 1. Cape
Region; 2. Karoo-Namib Region; 3. Uzambara-Zululand Region; 4. Madagascan Region; 5a. Southern Section of
the Zambezian Subregion; 5b. Northern Section of the Zambezian Subregion; 5c. Eastern Section of the Zambezian
Subregion; 6a. Erithreo-Arabian Subregion; 6b. Omano-Sindian Subregion; 7. Guineo-Congolian Region; 8. Sahelo-
Sudanian Subregion; 9. Saharo-Arabian Region; 10. Mediterranean Region; 11. Macaronesian Region; 12. Indian
Region; 13. Indochinese Region. A summary of the phylogenetic-morphological-biogeographic study is shown in Fig.
3 by Martínez-Azorín et al. (2023a) and is also discussed in the present work.
All plant images appearing in the figures were obtained by the authors unless otherwise acknowledged.
Results and discussion
Previous taxonomic proposals and phylogenetic relationships

Stedje (1998) found that two samples of Drimia sensu lato formed a clade clearly differenciated from the genera related
to both Scilla Linnaeus (1753: 308) (subfam. Hyacinthoideae) and Ornithogalum (subfam. Ornithogaloideae). This
was confirmed by Pfosser & Speta (1999) whose phylogenetic analysis was based on the plastid region trnL-F and
included 15 samples of Drimia, Urginea, and related urgineoid genera and formed a strongly supported clade (100%
BS=Bootstrap hereafter). Pfosser & Speta (1999) placed their analysed samples in eight genera, namely Charybdis,
Urginea, Karoophila (as nomen nudum), Urginavia, Drimia, Rhadamanthus, Ebertia, and Thuranthos, most of them
being monophyletic except Urginavia and Urginea in their sense. In the analysis by Pfosser & Speta (1999), a sample
of Bowiea volubilis Harvey ex Moore & Masters (1866: 971) was found to be sister to all other Urgineoideae.
Speta (1998a, 1998b, 2001), based on comprehensive studies combining morphological, anatomical, and molecular
data, described the monophyletic genera Boosia, Charybdis, Duthiea, Ebertia, Geschollia, Indurgia, Ledurgia,
Rhadamanthopsis, Sekanama, and Urginavia, each showing a unique combination of morphological characters and
biogeography. Soon after, the phylogenetic studies of Pfosser & Speta (2001, 2004) extended the sampling to more
than 140 samples of Urgineoideae, among them ca. 65 samples of Charybdis (here accepted as Squilla), although again
based on the single plastid region trnL-F. These results confirmed that most genera accepted by Speta (1998a, 1998b,
2001) formed well supported clades.
Nearly contemporaneously, Manning et al. (2004) extended the study to a second plastid region (trnL-F plus
rbcL) although with only limited sampling (21 samples) of Urgineoideae. On the basis of their interpretation Manning
et al. (2004) accepted a very broad Drimia, nearly congruent to the whole subfamily Urgineoideae, and characterised
by short-lived flowers (living from several hours to one day only), with tepals more or less united at the base, spurred
floral bracts, and the caducous and circumscissile perianth, abscising at the base and withering as a cap on the
developing capsule (Manning et al. 2002, Manning & Goldblatt 2007, Manning & Oliver 2009). However, several
groups within Urgineoideae do not agree with this definition, e.g. some groups with the perianth free at the base (cf.
Boosia, Fusifilum, Tenicroa, Urginea, etc.), not persistent at the top of the capsule after withering (cf. Rhodocodon,
Sekanama, or Urginavia), or the flowers last more than one day (Rhodocodon, Sagittanthera, and Aulostemon).
No details of the studied samples were provided by Manning et al. (2004), the phylogenetic tree was presented as a
study in preparation and the DNA sequences have to date not been reported to GenBank, thus precluding reproducibility
of their study and related analyses. Their phylogenetic tree showed Bowiea as basal to all other Urgineoideae, with both
clades strongly supported, so confirming the finding of Pfosser & Speta (1999). The core of samples of Urgineoideae in
Manning et al. (2004) apparently covered 12 genera (Boosia, Charybdis, Drimia, Fusifilum, Litanthus, Rhadamanthus,
Rhadamanthopsis, Schizobasis, Sekanama, Tenicroa, Thuranthos, and Urginavia) and the authors argued the interpreted
paraphyly or polyphyly of some of these genera to promote their broad Drimia concept, which included all 12 genera
in synonymy, and accordingly incorporate extremely variable morphology.
The argument of Manning et al. (2004) for their broad Drimia concept was that “Tenicroa Raf. is paraphyletic
unless T. nana Snijman is segregated as a monotypic genus, and the recently revived segregate Fusifilum Raf.
(including several of the Southern African species previously placed in Urginea) is very evidently polyphyletic”.
However, regarding Tenicroa, Pinter et al. (2013) have subsequently provided morphological evidence that supports
the acceptance of the phylogenetically distinct Tenicroa nana Snijman (1985: 284) as the monotypic genus Mucinaea.
Furthermore, the species accepted as Fusifilum in the tree of Manning et al. (2004) are placed in three different and
distant clades. However, two out of the four samples named as Fusifilum by these authors (“Fusifilum calcarata and
Fusifilum marginata” [sic]) have never been accepted or combined in this genus. Instead, these two species belong
to Geschollia and Austronea, respectively (cf. Martínez-Azorín et al. 2018a, 2019d). Of the remaining two taxa,
“Fusifilum dregei” belongs to Urgineopsis (cf. Martínez-Azorín et al. 2019a) whereas Fusifilum physodes is a true
member of this genus as revised by Müller-Doblies et al. (2001). Consequently, when those nomenclatural corrections
are accepted, the phylogenetic tree of Manning et al. (2004) supports an analytic treatment of the subfamily. Further,
Manning et al. (2004) opted to recognise their monotypic Bowiea as distinct from their very broadly conceived Drimia,
apparently motivated by its sister phylogenetic position with regards to the remaining Urgineoideae. Yet this decision
seems arbitrary based on the extremely variable morphology of their Drimia, which includes Bowiea-characters listed
by Manning et al. (2004) such as leaves absent in mature plants (cf. Schizobasis), branched, twining inflorescences
(cf. Schizobasis), long-lasting flowers (cf. Rhodocodon, Sagittanthera, and Aulostemon), withered tepals remaining
at the base of capsules (cf. Sekanama, Rhodocodon, and Urginavia) and subellipsoidal, relatively heavy seeds (cf.
Rhodocodon). In referring to a combination of the above-mentioned characters, recognition of Bowiea was no more
expansively justified by Manning et al. (2004) and Manning & Goldblatt (2018) than those genera recognised in
analytic approaches to the systematics of Urgineoideae (e.g. Speta 1998, Pfosser et al. 2012).
The recent revision of Drimia s.l. in Southern Africa by Manning & Goldblatt (2018) accepted 20 sections that
generally align with the already described urgineoid genera, although some were circumscribed as para- or polyphyletic
based on previous phylogenetic findings (Pfosser & Speta 1999, 2001, 2004, Pfosser et al. 2012). Our phylogenetic
results (see Martínez-Azorín et al. 2023a) also showed evidence of polyphyly of their Drimia sect. Macrocentrae
Manning & Goldblatt (2018: 19) (including species of both Boosia and Sekanama), D. sect. Ledebouriopsis (Baker
1873b: 284) Manning & Goldblatt (2018: 24) (merging species of Boosia, Geschollia, Urginavia, Urgineopsis, and
Zulusia), D. sect. Thuranthos (Wright 1916: 233) Manning & Goldblatt (2018: 62) [including species of Vera-duthiea,
Zingela, and Urginea revoluta Duthie (1928: 9) = Drimia hesperantha Manning & Goldblatt (2003: 111)], D. sect.
Physodia (Salisbury 1866: 37) Manning & Goldblatt (2018: 92) (including a species of Austronea), and D. sect.
Rhadamanthus (Salisbury 1866: 37) Manning & Goldblatt (2018: 129) (including species of Striatula). Moreover,
their D. sect. Hyacinthoides Manning & Goldblatt (2018: 71) is included in Rhadamanthopsis in the sense of the
present work and D. sect. Juncifoliae Manning & Goldblatt (2018: 107) in Tenicroa sensu Pinter et al. (2020).
In summary, our comprehensive study in Urgineoideae (Martínez-Azorín et al. 2023a) combining extensive
phylogenetic, morphological, and distribution data supports the existence of 31 monophyletic lineages that are well
defined by unique syndromes of morphological characters, which facilitate identification and robust circumscriptions.
A detailed characterisation of the accepted genera is provided below.
Comments on morphological characters

Petaloid monocotyledon families show a basic trimerous flowering pattern, as it is also the case of the Urgineoideae.
Variation in flower morphology is mainly related to the degree of connation of tepals, adnation and/or connation
of stamens, and morphology of the gynoecium, which were the main traits underpinning the historic generic
circumscriptions. Therefore merging, for instance, of Litanthus, Rhadamanthus, Rhodocodon, or Thuranthos into
Drimia or Urginea s.str. would be equivalent to the merging of Muscari Miller (1754: 926), Hyacinthus Linnaeus
(1753: 316), and several more genera into Scilla s.str. (a radical solution that has no advocates to date in the sister
subfamily Hyacinthoideae), or to lump Galtonia Decaisne (1880: 32), Neopatersonia Schönland (1912: 251), Avonsera
Speta (1998b: 95), or Cathissa Salisbury (1866: 34), among several others, into Ornithogalum s.str. (as proposed by
Manning et al. 2009). This approach results in an extreme variation in flower, fruit, and seed morphology within
single genera. The adoption of such very broad genus concepts more widely in systematics would compromise the
recognition of most genera in flowering plants. It would serve to obscure the utility of taxonomy and systematics,
not only for taxonomists, but also for general users, as genera with such variation in flower morphology would be
hardly comprehensible. In these wide genus concepts, a complex subgeneric classification is necessary to understand
the relationships among species, which are grouped by more homogeneous flower morphology. The taxonomic rank
of genus is one of the most intuitive and easy to work with, and subgenera, sections or series are usually reserved for
minor morphological variations. Moreover, naming plant groups by sections requires mention of the genus to which
they belong anyway. An interest in reducing the number of taxa at high taxonomic ranks has been articulated by some
such as the authors of APG (2009, 2016). In our opinion, the merging of sister clades into single highly heterogenous
taxa based only on genetic affinity is undesirable when clear morphological differences exist and the lumped groups
have been traditionally accepted, illustrated by the unsatisfactory merging of the monophyletic Hyacinthaceae into
Asparagaceae, or Alliaceae into Amaryllidaceae, inter alia.
Based on previous phylogenetic studies, it is evident that the degree of connation of tepals and adnation of
filaments has independently arisen in distantly related clades or genera in the Urgineoideae several times during
convergent evolutionary events (Martínez-Azorín et al. 2011a, 2023a). Further, such convergence events have also
occurred even within clades or genera, rendering single characters sometimes insufficient for generic circumscription.
For instance, some genera in Ornithogaloideae, such as Nicipe Rafinesque (1837: 54), show flowers with free or
rarely shortly connate tepals (Martínez-Azorín et al. 2011a), as also occurs in some of the genera accepted here in
Urgineoideae. Therefore, it is crucial to consider the whole variation of qualitative characters in reproductive and
vegetative organs to render consistent generic circumscriptions, as shown by Martínez-Azorín et al. (2023a).
Our studies in Urgineoideae in the last decade have documented that morphological characters in this group,
especially regarding the flowers, can be considered as relatively constant in several evolutionary lineages, which
show unspecialised flowers with a similar general morphology. However, a detailed study of fresh flowers reveals
taxonomically significant variability in morphology regarding mainly colour, shape, size and disposition of tepals,
stamens and gynoecium. These, taken together with vegetative characters, provide unique syndromes of morphological
characters that are linked to independent evolutionary lineages revealed by our phylogenetic studies (see Fig. 2 in
Martínez-Azorín et al. 2023a). This general similarity in flower morphology, seen also in several genera and families
of petaloid monocotyledons with distant phylogenetic relationships, is probably responsible for some of these evolutive
lineages being overlooked until recently. Discussions of the main morphological and chromosome characters involved
in the Urgineoideae are found below.
Bulbs:—Morphological variability of bulbs is relatively limited, and in principle relates to whether these organs
range from epigeal to hypogeal, and to whether the bulb scales are compact or loose and pedicellate. Epigeal bulbs
can be considered rare in Urgineoideae, although they are a prominent feature in one species of Urginavia [U. epigea
(Dyer 1947: t. 1027) Speta 2001: 168] and Drimia (D. robusta Baker 1870b: t. 190), some species of Schizobasis
and Zulusia, and in the two species of Bowiea, where this feature is an apomorphy. Bulbs with loose scales appear in
several species in the subfamily, such as Aulostemon mzimvubuensis (Van Jaarsv. in Van Jaarsveld & Van Wyk 2005:
83) Martínez-Azorín et al. (2017: 288), Austronea chalumnensis (Dold & Brink 2004: 631) Martínez-Azorín et al.
(2018a: 107), Drimia haworthioides Baker (1875: 366), Geschollia globuligera Martínez-Azorín et al. (2019d: 101),
Rhadamanthopsis haworthioides (Baker 1878: 322) Martínez-Azorín et al. comb. nov., R. hyacinthoides (Baker 1874c:
6) Martínez-Azorín et al. comb. nov., and R. monophyllus (Oberm. ex Manning & Goldblatt 2018: 128) Martínez-
Azorín et al. comb. nov., Rhadamanthus arenicola Nordenstam (1970: 166), Sagittanthera cyanelloides (Baker 1897:
444) Martínez-Azorín et al. (2013a: 48), Tenicroa namibensis M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in
Pinter et al. (2020: 83), Thuranthos macranthum Baker (1873b: 280) Wright (1916: 223), T. nocturnale Dyer (1964: t.
1439) and related taxa, Zingela pooleyorum Crouch et al. (2018: 36), and in Zulusia edwardsii (Crouch & Martínez-
Azorín 2015: 137) Mart.-Azorín et al. comb. nov. We hypothesize that the presence of loose bulb scales in species of the
cited genera must be considered a convergent phenomenon—a response to habitat selective pressures on adaptation,
since loose scales are usually associated with well drained sandy soils. In some genera, including Thuranthos and the
monotypic Aulostemon, Sagittanthera, and Zingela, loose bulb scales are key characters for generic circumscription
when combined with other data (see Table S2 in Martínez-Azorín et al. 2023a). Rarely, bulbs show fibres, especially
in the bulb neck—a feature observed in both Urginavia multisetosa (Baker 1897: 468) Martínez-Azorín et al. comb.
nov. and Tenicroa fibrosa Pinter et al. (2020: 70). These fibrous necks are reminiscent of those commonly occurring
in species of Albuca subgen. Mitrotepalum Müller-Doblies (1987: 332) related to A. setosa Jacquin (1795: 20), and
this feature might be involved in protection of the bulbs against herbivores. Further, connation and fusion of bulbs
scales in compact bulbs have also been presented as useful characters for systematics. Pinter et al. (2013) showed that
the monotypic Mucinaea has unique bulbs with a single sheathing cataphyll lacking raised transverse ribs, different
from Tenicroa species, and by a second sheath consisting of a compound of the bases of about ten foliage leaves,
which surrounds some ten additional free foliage leaves. This arrangement appears to be unique amongst bulbous
plants. Finally, species of Urginavia are easily idenfied by the presence of bulbs composed of leathery scales (usually
yellowish when dry), which are heterogeneous in length and vertically imbricate, and by producing abundant white
silky threads when broken, an apomorphic character of the genus, which is easily identified in herbarium material and
is diagnostic.
Leaves:—Leaves consistently form a basal rosette, and they are generally withered at flowering time. Some
taxa, however, mostly confined to subtropical and tropical regions, show synanthous or even persistent leaves. Some
epiphytic species of Rhodocodon are an example of the latter. The asynchrony of leaf and flower presentation may be
understood as a primitive character in the group, supported by the basal placement of Bowiea within Urgineoideae,
which shows withered or absent leaves in mature plants. Hysteranthous leaves provide a diagnostic character for
several genera, including Squilla, and several others. This behaviour is probably one reason for the poor knowledge
in this subfamily, since it is usually necessary either to cultivate the plants to obtain the key characters of leaves,
flowers, fruits, and seeds together, or to collect the plants in the wild at different times of the year. Most herbarium
collections accordingly show incomplete materials. Leaf morphology, indumentum, and presentation is highly varied
within the subfamily. Leaves are commonly elongated, flat and canaliculate, but sometimes they are terete and wither
from the apex by transversal plates (an apomorphy of Geschollia). Alternatively, they are short, ovate, flat, striate, and
appressed to the ground (an apomorphy of Striatula). In some groups, a distinct keel on the abaxial side occurs, as
seen in Sagittanthera or some species of Thuranthos (see Table S2 in Martínez-Azorín et al. 2023a). Coiled leaves are
common in several genera of subfamily Ornithogaloideae, a result of multiple putative convergent events—a fact that
has been interpreted as an adaptation to increase dew and fog precipitation through enlargement of surfaces and edges,
whilst still restricting the overall size of the leaves (Vogel & Müller-Doblies 2011). However, distinctly coiled leaves
appear only in a single species in Urgineoideae—Spirophyllos noctiflorus (Battandier & Trabut 1894: 505) Martínez-
Azorín et al. comb. nov. The coiled leaves are, therefore, one of the best diagnostic characters for this monotypic genus
restricted to deserts of northwestern Africa. Leaf margins are usually straight, although in some groups they may be
undulate, such as in Squilla undulata (Desfontaines 1792: 161) Martínez-Azorín et al. (2022: 10), or Drimia ssp.
Leaves are usually glabrous, but in some species are ciliate or show soft to hispid hairs. Zebrine cataphylls surrounding
the base of leaves also occur in Urgineoideae, such as in Geschollia zebrina Martínez-Azorín et al. (2019d: 109) or
Rhadamanthus fasciatus Nordenstam (1970: 174) and in all species of Tenicroa with prominent and transversally
barred cataphylls (at least weakly visible), which is an apomorphy of this genus. A similar pattern can be observed
in Ornithogaloideae in which these cataphylls occur in some species of Albuca, Coilonox Rafinesque (1837: 28), or
Nicipe, possibly as an outcome of convergence events (Müller-Doblies & Müller-Doblies 1981, Martínez-Azorín et al.
2011b, 2011c). The barred cataphylls may be considered as an adaptation against hervivores based on their aposematic
colouration and physical barrier.
Inflorescence:—Inflorescence is racemose, ranging from subspiciform to racemose or subcorymbose, with
flowers commonly arranged helicoidally, but rarely secund. Flower pedicels are usually patent to erect at anthesis,
sometimes arching downwards in species with nodding flowers, and fruit pedicels are mostly erect, but rarely patent as
in Spirophyllos or nodding-reflexed like in Ebertia. These characters are diagnostic of these two genera (Speta 1998b,
Martínez-Azorín et al. 2023a). Inflorescences and flower pedicels usually dry simultaneously with capsule ripening,
however in Aulostemon, Boosia, Bowiea, and Schizobasis, yellow, dry, dehisced capsules are presented on long-lasting,
green pedicels and peduncle—all reliable diagnostic characters of those genera. Inflorescences are commonly solitary
although in some species a second or even a third inflorescence can arise from a single bulb. This usually appears
sequentially, and all inflorescences are commonly linked to the split of the mother bulb to form clumps. As a very
distinct apomorphy, the genera Bowiea and Schizobasis present branched inflorescences that also act as photosynthetic
organs. Further, all known Bowiea species present twining inflorescences, whilst only few species of Schizobasis show
this character. Bowiea is also unique in producing fleshy, soft inflorescences.
Bracts and bracteoles:—Bracts in Urgineoideae, at least those of basal flowers, are spurred. This character is
unique in Urgineoideae and diagnostic of all Urgineoideae. Length and morphology of spurs varies from very short,
showing just a short abaxial bend, observable in Iosanthus Martínez-Azorín et al. (2019c: 584) and some species of
Austronea, to extremely long, being several times longer than the bract, as seen in species of Boosia. The extreme case
occurs in B. macrocentra (Baker 1887: 702) Speta (2001: 169) where spurs can reach 5 cm in length and are usually
coloured.
Bracteoles seem to be plesiomorphic for Hyacinthaceae, as they are characteristic of Oziroë Rafinesque (1837: 53)
that is basal, as revealed by the phylogenetic analyses of the family. They also occur in some genera in Hyacinthoideae
and Ornithogaloideae (Manning & Goldblatt 2018). In Urgineoideae, the bracteoles only appear in some genera, such
as Rhadamanthopsis, Sekanama, Squilla, Thuranthos, Urginavia, and Zingela. Rhodocodon also shows bracteoles
in some species. The presence of bracteoles was considered as an important character for generic circumscription
in Ornithogaloideae (Martínez-Azorín et al. 2011a) and this is also the case in Urgineoideae, where it is considered
characteristic of Rhadamanthopsis, Squilla, Thuranthos, Urginavia, and Zingela (see Table 1 in Martínez-Azorín et al.
2023a). Bracts are usually persistent, but in Thuranthos bracts and bracteoles are early caducous—an apomorphy of
that genus (see Table 1 in Martínez-Azorín et al. 2023a).
Flowers:—Martínez-Azorín et al. (2019a) found that the basic trimerous flower pattern in Urgineoideae is constant,
as it is in most other petaloid monocot families. Variations mainly relate to degree of connation of tepals, adnation
10 • Phytotaxa 610 (1) © 2023 Magnolia Press MARTÍNEZ-AZORÍN Et al.
and/or connation of stamens, and morphology of the gynoecium. All these features have historically served as the basis
of generic circumscriptions in Urgineoideae. However, based on previous phylogenetic studies, it became evident that
characters such as the degree of connation and adnation of floral whorls have evolved several times as convergent
events in independent and often distantly related clades or genera. Therefore, quantitative and qualitative characters
of flower morphology must necessarily be combined with vegetative, fruit, and seed characters, at best corroborated
by biogeographic patterns, to facilitate recognition of well-supported genera (Martínez-Azorín et al. 2011a, 2023a).
Flower morphology in Urgineoideae is very variable and ranges from widely open, star-like flowers with free tepals
to distinctly connate tepals producing tubular or urceolate flowers with very short apical lobes. There is also large
variation in staminal and gynoecial morphology and disposition. The flower morphology contributes to shaping of
the ecological niches in terms of two clearly differentiated flowering times—diurnal and nocturnal—associated with
distinct morphologies and fragrances. Genera adapted to nocturnal pollinators are Spirophyllos, Thuranthos, Triandra,
and Vera-duthiea—a trait considered apomorphic and diagnostic (see Table 1 in Martínez-Azorín et al. 2023a). Also
some species of Indurgia (in the sense of this work) have nocturnal flowers. Most species in Urgineoideae show
fugacious flowers, lasting from a few hours to a day, but some genera, such as Rhodocodon, Sagittanthera, and
Urginavia, have longer-lasting flowers.
Tepals:—Tepals in Urgineoideae vary from shortly-ovate to long-linear or spathulate, and from suberect to
strongly reflexed. Their colour is usually white, dull-yellow, brown or reddish, rarely purplish, and they usually show
a longitudinal central darker band (green, brown or purplish), sometimes only visible along the apical third and mostly
abaxially. Rarely, some genera show basal maculae, such as in Mucinaea or Aulostemon—a diagnostic and apomorphic
character for those genera (see Table 1 in Martínez-Azorín et al. 2023a). Connation of tepals was one of the main
characters used historically in generic circumscriptions. Genera Drimia, Litanthus, Rhadamanthus, Rhadamanthopsis,
and Rhodocodon show distinctly connate tepals, varying from a cylindrical to campanulate basal tube, with free portion
of tepals varying in length to urceolate with short and spreading free lobes. Our phylogenetic analyses revealed that
some genera would include species with tepals varying from free to somewhat connate at the base, as it also has been
observed for some genera in Ornithogaloideae (Martínez-Azorín et al. 2011a). Therefore, the connation of tepals alone
should not be used for generic delimitation. It is crucial to pay attention to the disposition of tepals and the morphology
of other floral whorls, combined with the degree of tepal connation. Some genera share tepals shortly connate in a
basal tube, such as in Urgineopsis and some species of Austronea, Geschollia, Rhadamanthopsis, Rhadamanthus,
and Urginavia, although some such tubes are sub-cylindrical whereas others are widely campanulate. Further, the
free segments can be suberect, patent or strongly reflexed, creating very different flower morphologies, and hence
providing characters for unequivocal generic delimination.
Stamens:—Stamens show high morphological variability and are of critical importance in the circumscription
of some genera. Connation of filaments and anthers is diagnostic for two recently described genera. Aulostemon is
almost unique in Urgineoideae by having filaments connate in a distinct long tube above the perigone segments, but
anthers free, only resembling those of Chionodoxa Boiss (1844: 61) and Scilla vardaria Yildirim & Gemici in Yildirim
et al. (2013: 51) or the floral corona of Puschkinia Adams (1805: 164) in Hyacinthoideae, whereas Sagittanthera is
unique in Urgineoideae in having large connate anthers, but free, shorter filaments. Some species in Rhadamanthopsis,
Rhadamanthus, and Rhodocodon have stamens connivent to the gynoecium, although they are never connate above the
perigone tube. Only few species of Drimia s.str. show a short connation of filaments beyond the tepals. Tepals, when
connate, usually present filaments adnate to the tube and arise either from the basal part of the tube or from its mouth, as
in Rhodocodon and Urgineopsis, respectively. Filament morphology is also an important character, since it varies from
flat and lanceolate to filiform and subterete or fusiform and thickened along the middle part. The latter morphological
syndrome is diagnostic of Fusifilum. Stamens are commonly smooth, although in Fusifilum, and some species of
Austronea and Geschollia, they possess distinct papillae along the basal third—a trait considered to be diagnostically
important. Filament disposition is also an important diagnostic character, with consistently spreading stamens seen
for example in Urgineopsis and Urginea, and stamens connivent to the style in Drimia s.str., Rhadamanthopsis,
Rhodocodon, and some species of Rhadamanthus in the sense of this work.
Gynoecium:—Ovary is consistently ovoid, oblong or subglobose, showing little morphological variation. It is
trigonous in transverse section. The ovary is usually well differenciated from the style, but some species show a less
marked transition. Style length varies from very short, as in some species of Rhadamanthus, to distinctly elongated and
longer than the ovary. Styles may be very thin and filiform, or thicker and subclavate. Stigmas are commonly small
and undifferentiated, but in some genera they are distinctly capitate (as in some species of Thuranthos and Iosanthus)
or truncate (as in Indurgia). The position of the style in relation to the ovary is a diagnostic character in some genera.
Deflexed and curved styles are found in Tenicroa, Thuranthos, and Vera-duthiea. In some species of Geschollia, the
styles may be deflexed while in others they are sigmoid (see Table 1 in Martínez-Azorín et al. 2023a).
A generic monograph of the Urgineoideae Phytotaxa 610 (1) © 2023 Magnolia Press • 11
Fruits:—Fruits are invariably trigonous, dry, loculicidal, dehiscent capsules where valves split to the very base.
Valves commonly remain suberect after dehiscence, only partially revealing the seeds. However, some species of
Iosanthus possess valves that strongly reflex to completely expose the seeds. Capsules are disposed on erect pedicels in
most genera and species in the subfamily, even when flowers are nodding, although Ebertia produces nodding capsules
on reflexed pedicels, and ripe fruits of Spirophyllos are presented horizontally on sigmoid-patent pedicels. The shape
of capsules varies from ovate or subglobose to distinctly elongate-oblong. Another important character of ripe capsules
is the position of the withered perianth. In most genera withered tepals cohere above and are circumcissile below,
remaining as a cap at the top of the developing capsule. However, in some genera including Bowiea, Urginavia,
Rhodocodon, and Sekanama, the withered perigone remains at the base to provide a diagnostic character for those
genera (see Table S2 in Martínez-Azorín et al. 2023a). Most genera have pedicels that dry progressively with capsule
ripening and dehiscence, but species of Aulostemon, Boosia, Bowiea, and Schizobasis are characterised by their long
lasting and photosynthetic peduncle and pedicels that are still green at the point when the dry capsules completely
dehisce (see Table 1 in Martínez-Azorín et al. 2023a). This persistent photosynthetic expression represents a clear
diagnostic character as well.
Seeds:—Seeds in Urgineoideae are in most cases flattened with a delicate testa that is easily detachable from the
embryo. However, considerable variation exists in general morphology, with some features linked to certain genera and
therefore representing diagnostic characters (Figs 1–6). Most genera and species in the subfamily show seeds adapted
to wind dispersal by being flattened, winged, and relatively light. However, Bowiea and some species of Rhodocodon
produce heavier ellipsoidal seeds, sometimes with a pronounced raphe in the latter genus (Figs 3–4; Brudermann et
al. 2018, 2019). Litanthus is unique in the subfamily in producing very small (0.5–1.5 mm long), trigonous, narrowly
pyramidal, pointed seeds (Fig. 2). Similarly, small, and irregularly compressed, pointed seeds are also diagnostic of
Geschollia (Fig. 2). Austronea produces trigonous and tetrahedrally folded seeds (Fig. 1). Larger seeds (5–9 mm long),
flattened, subellipsoidal and ca. 1.5–2 times longer than wide are present in Aulostemon, Drimia, Sekanama, Squilla,
Tenicroa, Thuranthos, Urginavia, Urginea, and Zingela. Superficially similar seeds, which are instead subisodiametric
rather than subellipsoidal, are characteristic of Iosanthus and Vera-duthiea (Figs 2, 5, 6). Elongated seeds, 2–5 times
longer than wide are characteristic of Zulusia and most species of Boosia (Figs 1, 6). Rhodocodon shows a remarkable
diversity in seed morphology, with most species producing subsphaerical-ellipsoid seeds, commonly showing distinct
elevated raphae, although a few species present light, flattened seeds (Knirsch 2012, Brudermann et al. 2018, 2019)
adapted to wind dispersal.
Chromosomes and genome size:—Based on the chromosome cytology review by Goldblatt et al. (2012),
karyotypes in Urgineoideae are bimodal, most members are diploid (2n=20), and the ancestral basic chromosome
number is x=10. Polyploidy was recorded in Urginavia, for the hexaploid D. capensis (2n=60), Indurgia (2n=40;
tetraploid), and Squilla (2n=40, 60; tetraploid and hexaploid, occasionally with B chromosomes). Chromosome
counts on Drimia elata as 2n=18 are considered as a disploid species, a pattern consistent with fusion of two short
chromosomes (De Wet 1957). Similar disploid samples are found in Schizobasis. Speta (1998b) cited chromosome
counts for Rhadamanthopsis (2n=12, 16, 18), Rhadamanthus (2n=18), and Urgineopsis (2n=14), although without
referencing sources or species counted. These counts would support the distinction of the mentioned genera. Data on
total DNA content per cell in Urgineoideae are only available for Bowiea volubilis, 1C=4.63 pg (Bennett & Smith
1976). Genome size in Urgineoideae has been estimated only in one species of Bowiea (B. volubilis, 9.26 pg/2C)
(Bennett & Smith 1976) and eight species of Indurgia (treated as Drimia in their work) (20.40 pg/2C to 40.80 pg/2C)
(Nath et al. 2014, 2015, 2022).

FIGURE 1. Seed morphology in Urgineoideae. 1. Aulostemon mzimvubuensis (van Jaarsv.) Mart.-Azorín et al.; 2. Austronea barkerae
(Oberm. ex J.C.Manning & Goldblatt) Mart.-Azorín et al.; 3. Austronea grandiflora Mart.-Azorín et al.; 4. Austronea hispidoplicata Mart.-
Azorín et al.; 5. Austronea linearis Mart.-Azorín et al.; 6. Austronea oblongifolia Mart.-Azorín et al.; 7. Austronea olifanta Mart.-Azorín
et al.; 8. Austronea pinguis Mart.-Azorín et al.; 9. Boosia flagellaris (T.J.Edwards et al.) Mart.-Azorín et al.; 10. Boosia macrocentra
(Baker) Speta; 11. Boosia modesta (Baker) Mart.-Azorín et al.; 12. Boosia natalensis (Baker) Mart.-Azorín et al.; 13. Boosia nyasae
(Baker) Mart.-Azorín et al.; 14. Boosia rubella (Baker) Mart.-Azorín et al.; 15. Boosia saniensis (Hilliard & B.L.Burtt) Mart.-Azorín et
al.; 16. Boosia tenella (Baker) Mart.-Azorín et al.; 17. Bowiea gariepensis van Jaarsv.; 18. Bowiea volubilis Harv. ex T.Moore & Mast.;
19. Drimia elata Jacq. ex Willd.; 20. Drimia haworthioides Baker. All to the same scale in mm.
FIGURE 2. Seed morphology in Urgineoideae. 1. Drimia neriniformis Baker; 2. Drimia robusta Baker; 3. Fusifilum capitatum (Hook.)
Speta; 4. Fusifilum montanum (A.P.Dold & E.Brink) A.P.Dold et al.; 5. Fusifilum physodes (Jacq.) Speta; 6. Geschollia anomala (Baker)
Speta; 7. Geschollia brachyandra Mart.-Azorín et al.; 8. Geschollia calcarata (Baker) Mart.-Azorín et al.; 9. Geschollia globuligera Mart.-
Azorín et al.; 10. Geschollia occultans (G.Will.) Mart.-Azorín et al.; 11. Geschollia prolifera Mart.-Azorín et al.; 12. Indurgia congesta
(Wight) Speta; 13. Indurgia indica (Roxb.) Speta; 14. Iosanthus amboensis (Baker) Mart.-Azorín et al.; 15. Iosanthus macrostigma
Mart.-Azorín et al.; 16. Iosanthus toxicarius (C.Archer & R.H.Archer) Mart.-Azorín et al.; 17. Litanthus pusillus Harv.; 18. Litanthus
stenocarpus (J.C.Manning & J.M.J.Deacon) Mart.-Azorín et al.; 19. Mucinaea nana (Snijman) M.Pinter et al.; 20. Rhadamanthopsis
haworthioides (Baker) Mart.-Azorín et al. All to the same scale in mm.

FIGURE 3. Seed morphology in Urgineoideae. 1. Rhadamanthopsis hyacinthoides (Baker) Mart.-Azorín et al.; 2. Rhadamanthopsis
karooicus (Baker) Mart.-Azorín et al.; 3. Rhadamanthopsis monophyllus (Oberm. ex J.C.Manning & Goldblatt) Mart.-Azorín et al.;
4. Rhadamanthopsis namibensis (Oberm.) Mart.-Azorín et al.; 5. Rhadamanthus arenicola B.Nord.; 6. Rhadamanthus ciliatus (L.f.)
Mart.-Azorín et al.; 7. Rhadamanthus cochlearis (Mart.-Azorín et al.) Mart.-Azorín et al.; 8. Rhadamanthus convallarioides (L.f.)
Salisb. ex Baker; 9. Rhadamanthus rigidifolius (Baker) Mart.-Azorín et al.; 10. Rhadamanthus urantherus R.A.Dyer; 11. Rhodocodon
campanulatus Knirsch et al.; 12. Rhodocodon cryptopodus (Baker) Knirsch et al.; 13. Rhodocodon cyathiformis H.Perrier ex Knirsch et
al.; 14. Rhodocodon floribundus H.Perrier ex Knirsch et al.; 15. Rhodocodon graciliscapus Knirsch et al.; 16. Rhodocodon intermedius
H.Perrier ex Knirsch et al.; 17. Rhodocodon jackyi Knirsch et al.; 18. Rhodocodon mascarenensis (Baker) Knirsch et al.; 19. Rhodocodon
monophyllus Knirsch et al.; 20. Rhodocodon perrieri Knirsch et al. All to the same scale in mm.
FIGURE 4. Seed morphology in Urgineoideae. 1. Rhodocodon rotundus H.Perrier ex Knirsch et al.; 2. Rhodocodon siederi Knirsch et
al.; 3. Rhodocodon urgineoides Baker; 4. Schizobasis intricata (Baker) Baker; 5. Schizobasis macowanii Baker; 6. Schizobasis sigmoidea
(J.C.Manning & J.M.J.Deacon) Mart.-Azorín et al.; 7. Sekanama brachystachys (Baker) Mart.-Azorín et al.; 8. Sekanama porphyrostachys
(Baker ex Balf.f.) Mart.-Azorín et al.; 9. Sekanama sanguinea (Schinz) Speta; 10. Squilla hesperia (Webb & Berthel.) Mart.-Azorín et al.;
11. Squilla maritima (L.) Steinh.; 12. Squilla palaestina (M.B.Crespo et al.) Mart.-Azorín et al.; 13. Squilla pancration Steinh.; 14. Squilla
secundiflora (Maire) Mart.-Azorín et al.; 15. Squilla serotina (Schousb.) Mart.-Azorín et al.; 16. Squilla undulata (Desf.) Mart.-Azorín
et al.; 17. Striatula oliverorum (J.C.Manning) M.Pinter et al.; 18. Striatula platyphylla (B.Nord.) M.Pinter et al.; 19. Tenicroa exuviata
(Jacq.) Speta; 20. Tenicroa fragrans (Jacq.) Raf. All to the same scale in mm.

FIGURE 5. Seed morphology in Urgineoideae. 1. Tenicroa multifolia (G.J.Lewis) Oberm.; 2. Tenicroa polyantha M.Pinter et al.; 3.
Tenicroa unifolia (A.V.Duthie) M.Pinter et al.; 4. Thuranthos bakeri Mart.-Azorín et al.; 5. Thuranthos macrocarpum (Stedje) Speta;
6. Thuranthos nocturnale R.A.Dyer; 7. Thuranthos pauciflorum (Baker) Mart.-Azorín et al.; 8. Triandra pellabergensis Mart.-Azorín
et al.; 9. Urginavia altissima (L.f.) Speta; 10. Urginavia capensis (Burm.f.) Mart.-Azorín et al.; 11. Urginavia echinostachya (Baker)
Mart.-Azorín et al.; 12. Urginavia kniphofioides (Baker) Mart.-Azorín et al.; 13. Urginavia micrantha (A.Rich.) Speta; 14. Urginavia
multisetosa (Baker) Mart.-Azorín et al.; 15. Urginavia simensis (Hochst. ex A.Rich.) Mart.-Azorín et al.; 16. Urginea fugax (Moris)
Steinh.; 17. Urgineopsis arenosa (Adamson) Mart.-Azorín et al.; 18. Urgineopsis dregei (Baker) Mart.-Azorín et al.; 19. Urgineopsis
exilis (Adamson) Mart.-Azorín et al.; 20. Urgineopsis gracilis (A.V.Duthie) Mart.-Azorín et al. All to the same scale in mm.
FIGURE 6. Seed morphology in Urgineoideae. 1. Urgineopsis salteri Compton; 2. Vera-duthiea capillifolia A.Patzelt et al.; 3. Vera-
duthiea sp. (Urginea petrophila A.Chev., nom. nud.); 4. Vera-duthiea sebiri (Berhaut) Mart.-Azorín et al.; 5. Vera-duthiea sp. (Urginea
segetalis A.Chev., nom. nud.); 6. Zingela pooleyorum N.R.Crouch et al.; 7. Zulusia delagoensis (Baker) Mart.-Azorín et al.; 8. Zulusia
edwardsii (N.R.Crouch & Mart.-Azorín) Mart.-Azorín et al.; 9. Zulusia lydenburgensis (R.A.Dyer) Mart.-Azorín et al.; 10. Urginea cf.
aurantica H.Lindb.; 11. Urginea revoluta A.V.Duthie. All to the same scale in mm.
TAXONOMIC TREATMENT
Identification key of genera in Urgineoideae
1. Anthers dehiscing by apical pores or slits extending to the middle .................................................................................................. 2

− Anthers dehiscing longitudinally along their whole length . ............................................................................................................. 6
2. Tepals with green basal markings adaxially ...................................................................................................................................... 3
− Tepals lacking green basal markings adaxially.................................................................................................................................. 4
3. Filaments of stamens distinctly connate for most of their length to form a tube that surrounds the gynoecium above the perigone;
tepals white with green maculae forming a basal green ring ...................................................................................... 1. Aulostemon
− Filaments of stamens free above the perigone; tepals purplish-pink with green and white, unconnected, basal markings.................
........................................................................................................................................................................................ 13. Mucinaea
4. Leaves 1–2(–3), up to 4 cm long, ovate, flattened and appressed to the ground, velutinous on the adaxial side, with longitudinal
furrows ........................................................................................................................................................................... 22. Striatula
− Leaves usually more numerous and longer, 2–40 cm long, linear-elongated and suberect, glabrous, smooth . ............................... 5
5. Anthers 5–6 mm long, connate at least in the upper portion to form a large conical yellow structure; leaves flattened with a distinct
broad keel on the abaxial side; flowers stellate with tepals 7–9 mm long ............................................................. 17. Sagittanthera
− Anthers 1–3 mm long, free, although connivent to the style; leaves linear-filiform; flowers campanulate-urceolate, rarely stellate
but then tepals up to 6 mm long ........................................................................................................................... 15. Rhadamanthus
6. Inflorescence distinctly branched with at least secondary branches ................................................................................................. 7
− Inflorescence not branched, always simple racemes . ....................................................................................................................... 8

7. Ovary conical, semi-inferior in appearance; inflorescence succulent; flowers lasting some days; tepals remain at the base of the
mature capsule .................................................................................................................................................................... 4. Bowiea
− Ovary ovoid, clearly superior; inflorescence never succulent; flowers usually lasting one day; tepals cohering above to form a cap
at the top of the mature capsule................................................................................................................................... 18. Schizobasis
8. Flowers small (4–7 mm long); tepals connate for most of their length, tubular, nodding; inflorescence 1(–2)-flowered with a pair
of subopposite bracts; stigma with six, minute, erect teeth . ......................................................................................... 12. Litanthus
− Flowers larger or with a different morphology; inflorescence usually with more flowers; bracts never subopposite; stigma indistinct
to trilobed .......................................................................................................................................................................................... 9
9. Flowers with only 3 stamens corresponding to the inner whorl . ................................................................................... 25. Triandra
− Flowers with 6 stamens ................................................................................................................................................................... 10
10. Flowers verpertine-nocturnal, nodding with strongly reflexed tepals . ........................................................................................... 11
− Flowers diurnal, patent to suberect (rarely nodding), with erect to spreading tepals (sometimes reflexed) . ................................. 15
11. Filaments adnate for ¼ to ½ of tepal length ................................................................................................................. 10. Iosanthus
− Filaments free to shortly adnate to tepals (less than ¼ of tepal length) .......................................................................................... 12
12. Leaves narrowly linear and distinctly coiled distally; filaments narrow, straight or slightly incurved, connivent to the style and
crossing each other along the upper third . ............................................................................................................... 20. Spirophyllos
− Leaves narrowly linear or wide and flattened but never distinctly coiled distally; filaments with a different morphology and/or
disposition ....................................................................................................................................................................................... 13
13. Filaments straight, suberect to spreading ......................................................................................................................... 9. Indurgia
− Filaments sigmoid, incurved and connivent to the style along the middle portion and spreading above ....................................... 14
14. Filaments narrow and distinctly constricted at base to join the tepals; bracteoles absent; bulb scales compact .... 29. Vera-duthiea
− Filaments distinctly widened along the basal portion to enclose the ovary; bracteoles present and distinct but usually early caducous;
bulb scales loose ........................................................................................................................................................ 24. Thuranthos
15. Leaves strongly keeled, with distinct purple maculations at base; dehisced anthers circinate; bulb scales loose ........... 30. Zingela
− Leaves not strongly keeled, nor maculate at base; dehisced anthers not circinate; bulb scales compact or rarely loose ............... 16
16. Basal leaves surrounded by sheathing cataphylls bearing transversally raised dark ribs; style bent downwards and usually distinctly
sigmoid, with papillose stigma......................................................................................................................................... 23. Tenicroa
− Basal leaves not surrounded by sheathing cataphylls bearing transversally raised dark ribs (rarely barred but then lacking prominent
ribs and showing papillate filaments); style erect or rarely declinate, not distinctly sigmoid, with minute to capitate stigma . .........
.......................................................................................................................................................................................................... 17
17. Inflorescence curved to nodding at early development stages; raceme usually short and widely corymbose to subglobose, rarely
subspiciform and compact; seeds tetrahedrally folded with narrowly winged angles ................................................... 2. Austronea
− Inflorescence erect at all development stages; raceme more elongated, racemose-spiciform, very rarely subcorymbose; seeds
flattened and usually windged, not tetrahedrally folded ................................................................................................................. 18
18. Tepals long connate for most of their length, rarely to the middle; flowers urceolate or campanulate, lasting for more than two days
(up to 7 days) ............................................................................................................................................................ 16. Rhodocodon
− Tepals free or fused to 2/5 of their length; flowers stellate to rarely campanulate, usually lasting for one or two days ................ 19
19. Tepals connate to 2/5 of their length forming a distinct tube .......................................................................................................... 20
− Tepals free or very shortly connate in an indistinct tube to 1 mm long .......................................................................................... 23
20. Bracteoles present and usually well developed ............................................................................................................................... 21
− Bracteoles lacking ........................................................................................................................................................................... 22
21. Flowers stellate, patent to suberect; free portion of tepals spreading-patent; ovary mostly visible from outside; stamens exerted and
spreading........................................................................................................................................................................ 26. Urginavia
− Flowers campanulate, mostly nodding; free portion of tepals erect; ovary included in the campanulate perianth; stamens included
and connivent to the style .................................................................................................................................14. Rhadamanthopsis
22. Free portion of tepals elongated and narrowly lanceolate-oblong to sub-spathulate and usually strongly reflexed; filaments
elongated-exerted and connivent to the style (rarely spreading above the connivent portion); ovary included in the perianth tube...
.............................................................................................................................................................................................. 5. Drimia
− Free portion of tepals spreading-patent and with different morphology; stamens exerted and spreading-incurved; ovary mostly
visible from outside ................................................................................................................................................... 28. Urgineopsis
23. Ovary and style white or sometimes tinged with violet; filaments fusiform and inflated in the middle, distinctly papillate on the
lower portion................................................................................................................................................................... 7. Fusifilum
− Ovary green to orange-yellow, never white; filaments not clearly fusiform, smooth or rarely papillate ....................................... 24
24. Dehiscent, dry, yellow capsules disposed on green, long lasting and photosynthetic pedicels; basal bracts usually long spurred, to
5 cm long ............................................................................................................................................................................. 3. Boosia
− Pedicels and capsules drying simultaneously; basal bracts usually with shorter spurs.................................................................... 25
25. Seeds polygonal or irregularly compressed, 1‒2.4(3) mm long; capsules 3‒6 mm long; leaf terete, mostly solitary . .......................
........................................................................................................................................................................................ 8. Geschollia
− Seeds and/or capsules larger or with different morphology; leaves terete or flat, commonly several ............................................ 26
26. Flower buds subglobose and nodding; flowers disposed on long, patent to suberect pedicels; leaves mostly flat, wide and short,
rarely linear subcanaliculated................................................................................................................................ 15. Rhadamanthus
− Flowers and leaves not as above ..................................................................................................................................................... 27
27. Seeds narrowly lanceolate, about 3 times longer than wide; leaves synanthous; bulb scales commonly loose; flowers carneous or
greenish, subcampanulate; tepals suberect, connivent at base and spreading apically .................................................... 31. Zulusia
− Seeds subellipsoid, to twice longer than wide; leaves mostly hysteranthous or rarely synanthous; bulb scales compact; flowers
white to yellowish, stellate with spreading tepals or rarely subcampanulate . ................................................................................ 28
28. Inflorescence (including peduncle) short, reaching up to 10 cm above ground level . ................................................................... 29
− Inflorescence (including peduncle) commonly longer than 15 cm above ground level ................................................................. 31
29. Capsules on laterally recurved pedicels ............................................................................................................................. 6. Ebertia
− Capsules on erect pedicels . ............................................................................................................................................................. 30
30. Capsules with the valves reflexed from the base at dehiscence to widely expose the seeds ....................................... 10. Iosanthus
− Capsules with the valves not reflexed at dehiscence . .................................................................................................... 11. Ledurgia
31. Bulb deep red inside; withered tepals papery, translucent; bracts shortly oblong-obovate to 3 mm long, navicular; peduncle above
ground level short, usually as long as or shorter than the raceme, rarely elongated ................................................... 19. Sekanama
− Bulb white to yellowish inside, rarely with a reddish tinge; withered tepals not papery and translucent; bracts usually narrowly
triangular, long acuminate, non-navicular; peduncle usually elongated ......................................................................................... 32
32. Bracteoles present and usually well developed, but sometimes early caducous . ........................................................................... 33
− Bracteoles lacking ........................................................................................................................................................................... 34
33. Bulb scales producing abundant whitish silky threads when broken; withered perigone remaining at the base of developing
capsule .......................................................................................................................................................................... 26. Urginavia
− Bulb scales not producing threads when broken; withered tepals basally circumcissile remaining as a cap at the top of the developing
capsule ............................................................................................................................................................................... 21. Squilla
34. Style subclavate, usually thickened, as long as or only slightly longer than the ovary; stigma truncate and trigonous; capsule with
apiculate valves; plants from India and neighboring countries ........................................................................................ 9. Indurgia
− Style filiform, about twice the ovary length; stigma minute; capsule valves not apiculate; plants from western Mediterranean
Basin ................................................................................................................................................................................ 27. Urginea
A synopsis of the accepted genera
1. Aulostemon Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 321(3): 288 (2017) ≡ Drimia sect.
Aulostemon (Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig) J.C.Manning & Goldblatt in Strelitzia 40: 123 (2018)
(Fig. 7). Typus generis:—Aulostemon mzimvubuensis (Van Jaarsv.) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig
(holotype) ≡ Drimia mzimvubuensis Van Jaarsv. in Aloe 42: 83 (2005), basionym.
Description:—Bulbous geophyte. Bulb partially epigeal, with thickened, fleshy, semitranslucent, loose, rounded and
club-shape scales, with short, flattened stalks, easily proliferating when detached, not producing threadlike structures
when broken. Roots thickened and branched. Leaves 1‒4 per bulb, linear, up to 50 cm long, 2‒3 mm in diam., leathery,
dark green, subterete, slightly channelled on adaxial side, finely striate with minute hairs. Inflorescence a long and lax
raceme up to 20 cm long, with 20‒50 nodding flowers; peduncle 25‒40 cm long; pedicels 14‒20 mm long; peduncle
and pedicels long persistent and photosynthetic after seed dispersal lasting from flowering time in spring to autumn.
Bracts 3‒5 mm long, lanceolate, acute, lowermost with long basal spur to 8‒9 mm long; bracteoles absent or rarely
present and minute. Flowers stellate, nodding, opening in afternoon. Tepals 6, biseriate, free from base, outer tepals
lanceolate, 10‒11 × 2.5‒2.8 mm, inner tepals oblong-lanceolate, 9‒10 × 3.0‒3.5 mm; adaxial side of tepals white with
green macula at base that connect to form green basal perigone ring; abaxial side of tepals white with narrow central
and longitudinal green band, perigone also showing green basal ring. Stamens 6; filaments flattened and connate along
most of their length to form distinct erect tube of ca. 3.5 mm long around gynoecium, ending in short triangular lobes
of ca. 1 mm long; anthers free, basifixed, sagittate, connivent to style, ca. 3 mm long, dehiscing by minute apical
pores. Ovary ovoid, trigonous, ca. 2.8 × 1.8 mm, green. Style narrow, erect, ca. 4.5 mm long, straight, mostly included
in filament tube and connivent anthers, elongating to overtop anthers after dehiscence of the latter. Stigma small and
indistinctly trigonous. Capsule triloculate, loculicidal, 7‒10 × 5.4‒7.5 mm, valves splitting to base. Seeds numerous,
flattened, suboblong to ellipsoid, 4.5‒6.0 mm long, with prominent embryo and wide, flat wings, black, with sinuous
anticlinal testa cell walls (Martínez-Azorín et al. 2017) (Fig. 7).
Number of species and distribution:—Aulostemon includes a single species restricted to the Eastern Cape
Province of South Africa, where it occurs northwest of Port St. Johns in the Mzimvubu (uMzimvubu) River gorge,
which is resticted to the Uzambara-Zululand Region (sensu Takhtajan 1986) (Fig. 8). For further information on
Aulostemon see Van Jaarsveld & Van Wyk (2005), Martínez-Azorín et al. (2017), and Manning & Goldblatt (2018).
Karyology:—Apparently not studied yet (Goldblatt et al. 2012).
History, diagnostic characters, and taxonomic relationships:—Aulostemon is almost unique in Urgineoideae
by having filaments of stamens distinctly connate for most of their length to form a long tube that surrounds the
gynoecium above the perigone, only resembling those of Chionodoxa and Scilla vardaria, and a green ring at the base
on the adaxial side of tepals. This, together with the anthers dehiscing by apical pores or slits, and the long-lasting

FIGURE 7. Aulostemon mzimvubuensis (van Jaarsv.) Mart.-Azorín et al.
photosynthesizing inflorescence and pedicels, make recognition of this genus very easy (Martínez-Azorín et al. 2017).
The presence of green basal maculae of tepals is a rare character that has also been used to recognise Mucinaea, a
genus in Urgineoideae also easily distinguishable by, among other characters, its purplish-pink tepals bearing a double-
eyed green floral marking at their base, surrounded by a white margin—a unique combination of traits in Urgineoideae
(Pinter et al. 2013). Martínez-Azorín et al. (2013a) transferred Drimia mzimvubuensis Van Jaarsv. in Van Jaarsveld &
Van Wyk (2005: 83) to Sagittanthera, based on the similar general flower morphology and distribution. Later, Martínez-
Azorín et al. (2017) have shown morphological differences supporting the independent generic status of Sagittanthera
and Aulostemon. This conclusion is also supported by phylogenetic analyses (Martínez-Azorín et al. 2023a) showing
that Aulostemon is related to Rhadamanthopsis—a genus showing very different flower morphology with campanulate
nodding flowers, connate tepals, and free filaments (in part adnate to the perigone tube) and anthers.
Accepted species:—
Aulostemon mzimvubuensis (Van Jaarsv.) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 321(3): 288 (2017) ≡ Drimia
mzimvubuensis Van Jaarsv. in Aloe 42(4): 83 (2005), basionym ≡ Sagittanthera mzimvubuensis (Van Jaarsv.) Mart.-Azorín,
M.B.Crespo, A.P.Dold & Van Jaarsv. in Phytotaxa 98(2): 51 (2013) (Figs 1.1, 7). Type:—SOUTH AFRICA. Eastern Cape Province.
Port St. Johns (3129): lower Mzimvubu River, south facing shale cliffs near Lutengela, (–CB), Van Jaarsveld, Xaba, Harrower &
Zwide 58 (PRE holo.).
FIGURE 8. Approximate distribution ranges of accepted genera in Urgineoideae.

2. Austronea Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 105 (2018) (Figs 9–12). Typus
generis:—Austronea marginata (Thunb.) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig (holotype) ≡ Anthericum
marginatum Thunb. in Prodr. Pl. Cap. 1: 63 (1794), basionym.
= Drimia sect. Capitatae J.C.Manning & Goldblatt in Strelitzia 40: 76 (2018). Typus sectionis:—Drimia marginata (Thunb.) Jessop
(holotype).
FIGURE 9. Species of Austronea Mart.-Azorín et al. displayed in horizontal rows of images. 1. Austronea acarophylla (E.Brink &
A.P.Dold) Mart.-Azorín et al.; 2. Austronea barkerae (Oberm. ex J.C.Manning & Goldblatt) Mart.-Azorín et al.; 3. Austronea chalumnensis
(A.P.Dold & E.Brink) Mart.-Azorín et al.; 4. Austronea densiflora Mart.-Azorín et al.
Description:—Small bulbous geophyte. Bulb hypogeal, usually with compact scales but rarely loose, outer scales
brownish and membranous. Roots thickened and branched. Leaves 1 to several per bulb, hysteranthous or rarely
synanthous, usually dark green, commonly leathery, terete or flat, rarely plicate, linear to ovate or rounded, usually
smooth, glabrous to hairy or hispid. Inflorescence usually short with long pedicels, giving subcapitate or subcorymbose
aspect, rarely with short pedicels, apex of peduncle and inflorescence curved or nodding at early stages of development,
but soon erect at anthesis; peduncle usually elongated, rarely short; pedicels of buds very short forming dense apical
raceme, distinctly elongating at anthesis. Bracts lanceolate, acute, lowermost with spur from slightly curved outgrowth
abaxially to long, wide spur with obtuse and lobate apex; bracteoles absent. Flowers stellate, erect-patent, diurnal,
usually opening in afternoon and withering in evening. Flower buds usually reddish or brownish-green. Tepals 6,
biseriate, 4‒9 mm long, free and spreading to slighty connate at base to form short cup and patent free portion of tepals;
adaxial side of tepals reddish, brownish, greenish or yellowish, rarely almost white; abaxial side of tepals with distinct
reddish, brownish, or greenish darker central band. Stamens 6, erect or spreading, not connivent to style; filaments
filiform to flattened, from free to shortly adnate to perigone and arising at end of perigone tube, straight or rarely
sigmoid, smooth or rarely papillate below; anthers ovate to oblong, dorsifixed, dehiscing by longitudinal slits along
their whole length; pollen yellow. Ovary ovoid to oblong or subglobose, trigonous, brown, red or green, sometimes
with white maculae, differentiated from style. Style white, narrow, erect, exerted, straight or slightly curved. Stigma
minutely papillose and indistinctly trigonous. Capsule subglobose, 4‒7 × 4‒7 mm, triloculate, loculicidal, valves
splitting to base, with withered perigone segments circumscissile below and forming an apical cap. Seeds up to 30 per
capsule, 2.0‒3.8 mm long, commonly trigonous in outline, tetrahedrally folded and narrowly winged along angles, with
subovoid embryo ending into long filamentous hilum, testa black, glossy, loose and easily detachable from endosperm,
with sinuous anticlinal testa cell walls (Martínez-Azorín et al. 2018a).
FIGURE 10. Species of Austronea Mart.-Azorín et al. displayed in horizontal rows of images. 1. Austronea grandiflora Mart.-Azorín
et al.; 2. Austronea hispidoplicata Mart.-Azorín et al.; 3. Austronea linearis Mart.-Azorín et al.; 4. Austronea marginata (Thunb.) Mart.-
Azorín et al.

FIGURE 11. Species of Austronea Mart.-Azorín et al. displayed in horizontal rows of images. 1. Austronea oblongifolia Mart.-Azorín et
al.; 2. Austronea olifanta Mart.-Azorín et al.; 3. Austronea papillosa Mart.-Azorín et al.; 4. Austronea patersoniae Mart.-Azorín et al.
Number of species and distribution:—Austronea includes 22 species restricted to South Africa and southern
Namibia (Fig. 8); it is restricted to the Cape, Karoo-Namib and Uzambara-Zululand Regions (sensu Takhtajan 1986).
Austronea plants are usually cryptic and difficult to find in the field due to their small, well camouflaged leaves that
are usually appressed to the ground, and becoming dry at flowering time, and also due to their delicate inflorescences
with small flowers. This fact, in combination with the poor conservation of floral structures in herbarium vouchers, has
confounded the taxonomy of the group, and accounts for why several species have remained unnoticed until recently
(Martínez-Azorín et al. 2018a, 2019b, 2020, 2023b). The taxonomy of the genus requires further resolution. Further
fieldwork is likely to result in an increase in the number of species in this genus. For further information on Austronea
species see Martínez-Azorín et al. (2016, 2018a, 2019b, 2020, 2023b) and Manning & Goldblatt (2007, 2018).
Karyology:—Apparently not studied yet (cf. Goldblatt et al. 2012).
FIGURE 12. Species of Austronea Mart.-Azorín et al. displayed in horizontal rows of images. 1. Austronea phyllopogon Mart.-Azorín
et al.; 2. Austronea pinguis Mart.-Azorín et al. (Photograph 2.1 by J. Slade); 3. Austronea pulchromarginata (J.C.Manning & Goldblatt)
Mart.-Azorín et al.; 4. Austronea trichophylla Mart.-Azorín et al.; 5. Austronea vermiformis (J.C.Manning & Goldblatt) Mart.-Azorín et
al.

History, diagnostic characters, and taxonomic relationships:—Austronea is easily distinguishable by a
syndrome of morphological characters that allow clear recognition: usually leathery and thickened leaves; relatively
small plant size; subcorymbose or congested raceme curved or nodding at early development stages, reddish to green-
yellowish tepals (see flower buds), which are free to shortly connate at the base to form a cup and patent free lobes;
filaments linear to lanceolate (not distinctly fusiform), smooth or rarely papillate below; ovary green to yellow-orange;
subglobose capsule, and seeds commonly trigonous in outline, tetrahedrally folded and narrowly winged along the
angles. As reported by Martínez-Azorín et al. (2018a), the general flower and inflorescence morphologies of Austronea
approach Fusifilum, an observation supported by the phylogenetic analyses of Martínez-Azorín et al. (2019c, 2023a)
in which both genera form strongly supported sister clades that nonetheless differ in clear morphological characters
and genetic distance. Species of Austronea were already recognised as a distinct group within Drimia sensu lato by
Manning & Goldblatt (2003, 2007), to include the species related to Drimia marginata (Thunberg 1794: 63) Jessop
(1977: 295), and it was later accepted as Drimia sect. Capitatae Manning & Goldblatt (2018: 76) by Manning &
Goldblatt (2018).
Austronea acarophylla (E.Brink & A.P.Dold) Mart.-Azorín, M.B.Crespo & A.P.Dold in Phytotaxa 365(2): 107 (2018) ≡ Drimia
acarophylla E.Brink & A.P.Dold in S. African J. Bot. 69(3): 396 (2003), basionym (Fig. 9.1). Type:—SOUTH AFRICA. Eastern
Cape. Grahamstown (3326): Committees Drift, Tyefu Location, (–BB), elev. 500 m, 22 August 1991, E. Brink 788 (GRA0000457!
holo.; BOL140329!, PRE0711374! iso.).
Austronea barkerae (Oberm. ex J.C.Manning & Goldblatt) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 107
(2018) ≡ Drimia barkerae Oberm. ex J.C.Manning & Goldblatt in Bothalia 33(1): 109 (2003), basionym (Figs 1.2, 9.2). Type:—
SOUTH AFRICA. Western Cape. Clanwilliam (3218): 5 km south west of Eendekuil, Farm Draaihoek, (–DD), open clay flats, 13
October 2001, J.C. Manning 2655A (NBG0197702! holo.; PRE iso.).
Austronea chalumnensis (A.P.Dold & E.Brink) Mart.-Azorín, M.B.Crespo & A.P.Dold in Phytotaxa 365(2): 107 (2018) ≡ Drimia
chalumnensis A.P.Dold & E.Brink in S. African J. Bot. 70(4): 631 (2004), basionym (Fig. 9.3). Type:—SOUTH AFRICA. Eastern
Cape. Peddie (3327): Cornfields Farm, near Chalumna River, 5 km northwest of Kayser’s Beach, 35 km southwest of East London,
(–BA), elev. 100 m, 25 October 2002, A.P. Dold 4619 (GRA! holo.).
Austronea ciliolata (J.C.Manning & J.M.J.Deacon) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 293 (2019)
≡ Drimia ciliolata J.C.Manning & J.M.J.Deacon in Strelitzia 40: 89 (2018), basionym. Type:—SOUTH AFRICA. Western Cape.
Montagu (3320): Anysberg, (–DA), quartz patches S of Anysberg, 20 August 2016 [leafing bulb; orig. coll. 11 Oct. 2015], J. Deacon
4411 (NBG holo.).
Austronea densiflora Mart.-Azorín, M.B.Crespo & A.P.Dold in Phytotaxa 365(2): 109 (2018) ≡ Drimia densiflora (Mart.-Azorín,
M.B.Crespo & A.P.Dold) J.C.Manning & Goldblatt in Bothalia 49(1): 2 (2019) (Fig. 9.4). Type:—SOUTH AFRICA. Western Cape.
Oudtshoorn (3322): Oudtshoorn, Grootkop Nature Reserve, NE of Oudtshoorn, (–CA), elev. 415 m, 28 September 2011 (in flower),
M. Martínez-Azorín, J. Vlok, A.P. Dold & A. Martínez-Soler MMA891 (GRA! holo.; ABH59705! iso.).
Austronea ecklonii (Baker) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 107 (2018) ≡ Urginea ecklonii
Baker in Bot. Jahrb. Syst. 15(3, Beibl. 35): 6 (1892), basionym. Type:—SOUTH AFRICA. Western Cape. Clanwilliam (3218):
Olifantsrivier, (–BD), September 1829 or 1830, Ecklon & Zeyher Asphod. 128 (NBG-SAM! lecto. designated by Tang & Weiglin
2001). Epitype (designated by Manning & Goldblatt 2018):—SOUTH AFRICA. Wuppertal (3219): Cedarberg, Wolfberg, (–AC), 3
October 1952 (fl. ex hort Dec 1952), E. Esterhuysen 20587 (BOL! epi.).
= Drimia ligulata J.C.Manning & Goldblatt in Taxon 63(6): 1330 (2014). Type:—SOUTH AFRICA. Western Cape. Clanwilliam (3218):
‘Piketberg, Zebra Kop’, (–DB), 23 May 1948, E. Esterhuysen 14487 (BOL150154! holo.: only the bulb with three leaves on lower
left-hand corner of sheet; K! iso.).
Austronea fimbrimarginata (Snijman) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 108 (2018) ≡ Drimia
fimbrimarginata Snijman in Bothalia 39(2): 234 (2009), basionym. Type:—SOUTH AFRICA. Western Cape. Vanrhynsdorp
(3118): Knersvlakte, Farm Moedverloor, ± 17 km NE of Koekenaap, (–AD), on quartzite ridges, 22 July 2005, A.D. Harrower 2762
(NBG0232164! holo.; PRE iso.).
Austronea grandiflora Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 112 (2018) (Figs 1.3, 10.1). Type:—
SOUTH AFRICA. Northern Cape. Sutherland (3220): NW of Sutherland, 2 km from Sutherland-Calvinia R354 road, on turn off
to Bo-Visrivier, (–AB), elev. 1264 m, 15 April 2014 in flower ex hort at University of Alicante (Spain), M. Martínez-Azorín, A.
Martínez-Soler & R. McKenzie MMA828b (GRA! holo.; ABH! iso.).
Austronea hispidoplicata Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso in Phytotaxa 365(2): 114 (2018) ≡ Drimia hispidoplicata
(Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso) J.C.Manning & Goldblatt in Bothalia 49(1): 3 (2019) (Figs 1.4, 10.2). Type:—
SOUTH AFRICA. Northern Cape. Hondeklipbaai (3017): ca. 10 km NE of Kotzesrus on the way to Garies, (–DB), elev. 160 m,
sandy soil on quartzitic ridge, 20 August 2017 (in leaf), M. Martínez-Azorín, M.B. Crespo, M.Á. Alonso & M. Pinter MMA1788
(GRA! holo.; ABH76973! iso.).
Austronea linearis Mart.-Azorín, M.B.Crespo & A.P.Dold in Phytotaxa 365(2): 116 (2018) (Figs 1.5, 10.3). Type:—SOUTH AFRICA.
Western Cape. Ladismith (3321): ca. 3 km E of Calitzdorp, on main road to Oudtshoorn, (–DA), elev. 350 m, 28 September 2011 in
flower and fruit, M. Martínez-Azorín, J. Vlok, A.P. Dold & A. Martínez-Soler MMA928 (GRA! holo.; ABH59622! iso.).
Austronea marginata (Thunb.) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 108 (2018) ≡ Anthericum
marginatum Thunb., Prodr. Pl. Cap. 1: 63 (1794), basionym ≡ Idothea marginata (Thunb.) Kunth, Enum. Pl. 4: 346 (1843) ≡ Drimia
marginata (Thunb.) Jessop in J. S. African Bot. 43(4): 295 (1977) (Fig. 10.4). Type:—SOUTH AFRICA. Northern Cape. Calvinia
(3119): Hantam, November 1774, Thunberg s.n. (UPS-8393 holo. [digital image!]).
Austronea oblongifolia Mart.-Azorín, A.P.Dold, M.B.Crespo & M.Á.Alonso in Phytotaxa 585(1): 49 (2023) (Figs 1.6, 11.1). Type:—
SOUTH AFRICA. Eastern Cape. Fort Beaufort (3226): Normandale farm, Bedford district, 20 km south of Bedford on R350 to
Grahamstown, (–CC), elev. 660 m, 11 October 2022 (in flower and fruit), A.P.Dold TD16062 (GRA! holo.).
Austronea olifanta Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso in Phytotaxa 365(2): 119 (2018) (Figs 1.7, 11.2). Type:—SOUTH
AFRICA. Western Cape. Clanwilliam (3218): ca. 6 km south of Clanwilliam, near Clanwilliam–Citrusdal–Cederberg road crossing,
(–BB), elev. 124 m, 23 September 2015 (in flower), M. Martínez-Azorín, M.B. Crespo, M.Á. Alonso & M. Pinter MMA1170 (GRA!
holo.; ABH74854! iso.).
Austronea papillosa Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso in Phytotaxa 365(2): 121 (2018) (Fig. 11.3). Type:—SOUTH
AFRICA. Northern Cape. Gamoep (2918): Naib se Berg, central section, on way to south facing entrance of main kloof, (–AD), elev.
916 m, lower slopes covered with quartz, 28 September 2015 (in flower), M. Martínez-Azorín, M. Pinter, M.B. Crespo & M.Á. Alonso
MMA1260 (GRA! holo.; ABH74112! iso.).
Austronea patersoniae Mart.-Azorín, A.P.Dold & M.B.Crespo in Phytotaxa 400(2): 77 (2019) (Fig. 11.4). Type:—SOUTH AFRICA.
Eastern Cape. Grahamstown (3326): Alicedale, (–AC), 1 July 1919 (in flower and fruit), F. Cruden 300 (GRA! holo.; PRE iso.).
Austronea phyllopogon Mart.-Azorín, M.Pinter, M.B.Crespo & M.Á.Alonso in Phytotaxa 449(2): 203 (2020) (Fig. 12.1). Type:—SOUTH
AFRICA. Northern Cape. Kamiesberg (3018): NE of Garies, Kamiesberg, ca 2 km NW of turn-off to Leliefontein from Studer’s Pass
road, near farm house, (–AC), elev. 1123 m, sandy soil, 11 April 2020 (in flower ex hort.), M. Martínez-Azorín, M.B. Crespo, M.Á.
Alonso & M. Pinter MMA1817b (GRA! holo.; ABH! iso.).
Austronea pinguis Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso in Phytotaxa 365(2): 123 (2018) ≡ Drimia pinguis (Mart.-Azorín,
M.B.Crespo, M.Pinter & M.Á.Alonso) J.C.Manning & Goldblatt in Bothalia 49(1): 3 (2019) (Figs 1.8, 12.2). Type:—SOUTH
AFRICA. Western Cape. Montagu (3320): Rooinek Pass, ca. 18 km south of Laingsburg on R323, (–BD), elev. 759 m, ex hort at
University of Alicante on 4 May 2018 in flower, M. Martínez-Azorín, M. Pinter, M.B. Crespo & M.Á. Alonso MMA1342b (GRA!
holo.; ABH! iso.).
Austronea pulchromarginata (J.C.Manning & Goldblatt) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 108
(2018) ≡ Drimia pulchromarginata J.C.Manning & Goldblatt in Bothalia 37(2): 185 (2007), basionym (Fig. 12.3). Type:—SOUTH
AFRICA. Northern Cape. Kamiesberg (3018): Farm Draaiklip, (–AA), 31 October 1983, C.H. Stirton 9226 (NBG0127044! holo.).
Austronea pygmaea (A.V.Duthie) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 108 (2018) ≡ Urginea pygmaea
A.V.Duthie in Ann. Univ. Stellenbosch sect. A, 6(2): 10 (1928), basionym ≡ Fusifilum pygmaeum (A.V.Duthie) Speta in Phyton
(Horn, Austria) 38(1): 69 (1998). Type:—SOUTH AFRICA. Western Cape. Cape Town (3318): ‘Stellenbosch Flats’, (–DD), 3 June
1926, A. Duthie s.n. STE1603a (NBG0197707-0! lecto. designated by Martínez-Azorín et al. 2015: 169: the eleven bulbs with leaves
on the left hand side of the label, under the collecting number 1603a).

Austronea trichophylla (Mart.-Azorín, A.P.Dold & M.B.Crespo) Mart.-Azorín, M.B.Crespo & A.P.Dold in Phytotaxa 365(2): 108 (2018)
≡ Drimia trichophylla Mart.-Azorín, A.P.Dold & M.B.Crespo in Syst. Bot. 41(4): 944 (2016), basionym (Fig. 12.4). Type:—SOUTH
AFRICA. Eastern Cape. Grahamstown (3326): Cradock Road, ca. 6 miles from BRU [Botanical Research Unit, currently Selmar
Schonland Herbarium], Grahamstown, (–AD), 13 November 1979, C. Vosa & E. Brink s.n. (GRA! holo.).
Austronea vermiformis (J.C.Manning & Goldblatt) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 108 (2018)
≡ Drimia vermiformis J.C.Manning & Goldblatt in Taxon 63(6): 1330 (2014), basionym ≡ Drimia vermiformis J.C.Manning &
Goldblatt in Bothalia 37(2): 184 (2007), nom. inval. (Fig. 12.5). Type:—SOUTH AFRICA. Western Cape. Clanwilliam (3218):
Clanwilliam Dam, picnic site along N7 near wall, (–BB), 3 August 1987, P.L. Perry 3587 (NBG0149721! holo.: the leaf only).
Austronea virens (Schltr.) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 365(2): 108 (2018) ≡ Urginea virens Schltr.
in J. Bot. 35: 433 (1897), basionym ≡ Drimia virens (Schltr.) J.C.Manning & Goldblatt in Strelitzia 9: 712 (2000). Type:—
SOUTH AFRICA. Western Cape. Wuppertal (3219): Cold Bokkeveld, Tweefontein, (–CD), 24 January 1897, R. Schlechter 10127
(BOL140324! lecto. designated by Manning & Goldblatt 2007: 184; E00193963!, GRA0000456!, BM000911790!, K000257373!,
L0041410!, P02156319!, PRE0049748!, S06-4413!, Z-000027559! isolecto.). Epitype (designated here):—ibidem (E00193963!
epi.). Note:—We designate an epitype based on the presence of leaves, a crucial character for species identification.
3. Boosia Speta in Stapfia 75: 168 (2001) (Figs 13–15). Typus generis:—Boosia macrocentra (Baker) Speta
(holotype).
= Drimia sect. Macrocentrae J.C.Manning & Goldblatt in Strelitzia 40: 19 (2018) pro parte. Typus sectionis:—Drimia macrocentra
(Baker) Jessop (holotype).
FIGURE 13. Species of Boosia Speta. 1. Boosia macrocentra (Baker) Speta from Phillips (1924: pl. 142, as Urginea macrocentra Baker);
2. Boosia flagellaris (T.J.Edwards et al.) Mart.-Azorín et al. from Edwards et al. (2005: 123, as Drimia flagellaris Edwards et al.).
FIGURE 14. Species of Boosia Speta displayed in horizontal rows of images. 1. Boosia flagellaris (T.J.Edwards et al.) Mart.-Azorín et
al.; 2. Boosia macrocentra (Baker) Speta; 3. Boosia modesta (Baker) Mart.-Azorín et al.; 4. Boosia natalensis (Baker) Mart.-Azorín et
al.
Description:—Bulbous geophyte. Bulb hypogeal or partially epigeal. Roots thickened and branched. Leaves 1 to
several per bulb, hysteranthous or synanthous, narrowly-linear and elongated, terete, glabrous, smooth, 5‒60 cm long
and 2‒15 mm wide, usually purple-red at base. Inflorescence usually multiflowered long and narrow raceme, rarely
short and subcorymbose, axes of inflorescence straight or zig-zag; peduncle terete commonly wiry, sinuous at base and
purplish, smooth, and glabrous; pedicels usually short and curved, to 25 mm long. Bracts lanceolate, acute, up to 8 mm
long, lowermost with characteristic very long basal spur, up to 5 cm long, usually bifid and clasping stem; bracteoles
absent. Flowers stellate, erect-patent, diurnal. Tepals 6, biseriate, 3.5‒8.0 mm long, free from base, outer tepals ovate-
lanceolate, inner tepals oblong-lanceolate to obovate; adaxial side of tepals white; abaxial side of tepals white with
narrow central and longitudinal band commonly purplish-green. Stamens 6, erect to spreading, not connivent to style;
filaments slightly flattened, free, smooth; anthers dorsifixed, dehiscing by longitudinal slits along their whole length.
Ovary ovoid to oblong, trigonous, green, sometimes with white apical or lateral markings, truncate to style. Style
white, narrow, erect, straight. Stigma small, papillose, trigonous. Capsule from narrowly ovoid to subellipsoidal, 3‒11

mm long, triloculate, loculicide, valves splitting to base, with withered perigone segments circumscissile below and
forming an apical cap, pedicels of dehiscent dry capsules long lasting, green and photosynthetic. Seeds numerous,
narrowly ovate-hemidiscoid or fusiform, 2.0‒6.5 mm long, commonly elongate, about 2 to 5 times longer than wide,
testa black, glossy, longitudinally striate, with commonly elongate cells.
FIGURE 15. Species of Boosia Speta displayed in horizontal rows of images. 1. Boosia nyasae (Baker) Mart.-Azorín et al.; 2. Boosia
rubella (Baker) Mart.-Azorín et al.; 3. Boosia saniensis (Hilliard & B.L.Burtt) Mart.-Azorín et al.; 4. Boosia tenella (Baker) Mart.-Azorín
et al.
Number of species and distribution:—Boosia includes 11 species, occurring in southern and east Africa, which
are restricted to the Uzambara-Zululand Region and the Southern and Eastern Sections of the Zambezian Subregion
(sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a). The highest diversity in the genus occurs in the southeastern
regions of South Africa, in subtropical areas in the KwaZulu-Natal Province (Fig. 8), with some species extending north
to East Africa. Although we preliminarily accept most described taxa in the group as different species, the taxonomy of
this genus is in urgent need of revision to explore their morphological variability across its whole distribution range.
For further information on Boosia species see Manning & Goldblatt (2018).
Karyology:—2n=20 (De Wet 1957, as Urginea tenella Baker); 2n=40 (De Wet 1957, as Urginea pretoriensis);
2n=40 plus 2 fragments (De Wet 1957, as Urginea rubella).
History, diagnostic characters, and taxonomic relationships:—Speta (2001) placed Urginea macrocentra
(Figs 13.1, 14.2) into the monotypic Boosia, based on its single, terete, thick, and corky proteranthous leaf, short bracts
with very long, coloured, flattened spurs up to 3.8 cm long, elongated raceme with stellate flowers, tepals very shortly
connate at their base, and flattened seeds. He also suggested that Drimia modesta (Baker) Jessop represented the
sister group and questioned the suitability of applying the name Urgineopsis to this group. The phylogenetic analyses
of Pfosser & Speta (2001, 2004) included two “Boosia” samples, one labelled “H840 B. macrocentra” from Barkly
Pass and the other “H847 Boosia sp.” from Swellendam, and revealed their sister relationship, with strong support.
Those latter two samples were sister to a clade including two samples labelled “H852 Urgineopsis cf. modesta” from
Nieuwoudtville, South Africa, and “H924 Urgineopsis sp. aff. tenellum” from cultivated material. As explained by
Martínez-Azorín et al. (2023a), neither of the two samples named Urgineopsis in their analyses correspond to the
concept applied to that genus in the present work.
The recent phylogenetic studies by Martínez-Azorín et al. (2023a) covered 16 samples related to Boosia
macrocentra and found those to form a strongly supported clade, which is sister to Geschollia. The 16 samples were
divided into two subclades that can be interpreted in biogeographic terms. One of the clades comprises 12 samples from
eastern South Africa (mostly KwaZulu-Natal) and Lesotho that includes species such as Urginea saniensis Hilliard &
Burtt (1985: 253), U. modesta Baker (1892: 6), U. natalensis Baker (1897: 468), U. tenella Baker (1897: 464), and
Drimia flagellaris Edwards et al. (2005: 122); these share with U. macrocentra terete leaves, usually long-spurred
basal bracts, and flattened, elongated seeds. This group of species is here accepted as the genus Boosia, which occur
along eastern South Africa to East Africa and can be recognised by a combination of the following characters: terete
leaves; usually long racemose inflorescence with sinuous and purplish peduncle base; lowermost bracts commonly
with long spurs, usually bifid and clasping the stem; bracteoles absent; flowers stellate with free tepals; spreading
stamens; pedicels of dehiscent, dry, yellow capsules and inflorescence peduncle green, long lasting and photosynthetic;
withered tepals persisting as a cap at the top of the developing capsules; and the commonly elongated and striate seeds.
Boosia is sister to Geschollia, a lineage that differs in the usually single leaf, pedicels drying simultaneoulsy with
capsules, comparatively very small capsules and seeds, and in its centre of diversity being located in the Eastern Cape
Province of South Africa (Martínez-Azorín et al. 2019d).
The other sister subclade to Boosia in Martínez-Azorín et al. (2023a) includes samples from western South
Africa and comprises two samples of Fusifilum magicum Müller-Doblies et al. (2001: 491), the Pfosser and Speta
sample “H847 Boosia sp.” from Swellendam, and a sample from Betty’s Bay, South Africa of Urginea revoluta. The
identity and taxonomy of this latter morphologically heterogeneous subclade remains uncertain until further samples
and additional molecular markers are included in the phylogenetic analyses.
Accepted species and required new combinations:—
Boosia flagellaris (T.J.Edwards, D.Styles & N.R.Crouch) Mart.-Azorín, N.R. Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia
flagellaris T.J.Edwards, D.Styles & N.R.Crouch in S. African J. Bot. 71(1): 122 (2005), basionym (Figs 1.9, 13.2, 14.1). Type:—
SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): Krantzkloof, in cliff faces, (–DD), 28 July 2003, N.R. Crouch 1023
(NU0016039! holo.; K000400681!, NH!, PRE! iso.).
Boosia macrocentra (Baker) Speta in Stapfia 75: 169 (2001) ≡ Urginea macrocentra Baker in Gard. Chron. 1887, 1: 702 (1887), basionym
(Figs 1.10, 13.1, 14.2). Type:—SOUTH AFRICA. Eastern Cape, Transkei, without date, Barber 895 (K000400573! holo.).
= Urginea lilacina Baker, Fl. Cap. (Harvey) 6: 469 (1897). Type:—SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): Inanda,
(–DB), September 1881, J. Medley Wood 642 (K000257362! lecto. designated by Jessop in J. S. African Bot. 43: 292. 1977;
BM000911789!, NH 0005786-0!, SAM! isolecto.).
= Urginea schlechteri Baker in Bull. Herb. Boissier ser. 2, 4: 1000 (1904). Type:—SOUTH AFRICA. KwaZulu-Natal. Stanger (2931):
Claremont, (–CC), 1 August 1893, Schlechter 3155 (BOL! lecto. designated here; GRA!, K000257372!, PRE0049746-0!,
Z000027554! isolecto.).
Boosia mandalensis (Baker) Mart.-Azorín, N.R. Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea mandalensis Baker, Fl. Trop.
Afr. [Oliver et al.] 7(3): 536 (1898), basionym. Type:—MALAWI. [formerly Nyasaland], Ndirandi Mountain near Blantyre, Scott-
Elliot 8476 (K000400582! holo.).
Boosia modesta (Baker) Mart.-Azorín, N.R. Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea modesta Baker in Bot. Jahrb.
Syst. 15(3, Beibl. 35): 6 (1892), basionym ≡ Urgineopsis modesta (Baker) Speta in Linzer Biol. Beitr. 12(1): 205, 230 (1980),
comb. inval. (lacking basionym reference) (Figs 1.11, 14.3). Type:—SOUTH AFRICA. Eastern Cape, Pondoland, Bachmann 273
(K000099182! holo.).

Boosia natalensis (Baker) Mart.-Azorín, N.R. Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea natalensis Baker, Fl. Cap.
(Harvey) 6(3): 468 (1897), basionym (Figs 1.12, 14.4). Type:—SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): Inanda,
(–DB), June [18]79, J. Medley Wood 277 (K000257347! lecto. designated by Jessop in J. S. African Bot. 43: 303. 1977; NH0004740-
0! isolecto.).
Boosia nyasae (Baker) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea nyasae Rendle in Trans. Linn. Soc.
London, Bot. 4(1): 50 (1894), basionym (Figs 1.13, 15.1). Type:—MALAWI. Milanji, elev. 6000 ft., October 1891, A. Whyte s.n.
(BM000911782! lecto. designated by Manning & Goldblatt in Bothalia 43(1): 76. 2013).
Boosia pretoriensis (Baker) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea pretoriensis Baker in Bull.
Herb. Boissier ser. 2, 1: 786 (1901), basionym. Type:—SOUTH AFRICA. Transvaal [Gauteng], Pretoria, Colle supra Aapies [Apies]
River, A. Rehmann 4307 (Z000027551! holo.).
Boosia rubella (Baker) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea rubella Baker, Fl. Cap. (Harvey)
6(3): 467 (1897), basionym (Figs 1.14, 15.2). Type:—SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): near the Mooi
River, (–AA), elev. 4000–5000 ft., 22 October 1894, J. Medley Wood 5723 (K000099183! lecto. designated as “holo.” by Jessop in
J. S. African Bot. 43: 303. 1977; BM001122598!, BOL140323!, SAM0023220-0!, PRE0048621-0! isolecto.).
Boosia saniensis (Hilliard & B.L.Burtt) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea saniensis Hilliard
& B.L.Burtt in Notes Roy. Bot. Gard. Edinburgh 42(2): 253 (1985), basionym ≡ Drimia saniensis (Hilliard & B.L.Burtt) J.C.Manning
& Goldblatt in Bothalia 33(1): 111 (2003) ≡ Urginea saniensis Hilliard & B.L.Burtt in Taxon 63: 1332 (2014), nom. superfl. (Figs
1.15, 15.3). Type:—LESOTHO. Underberg (2929): top of Sani Pass, (–CA), 6 November 1973, Hilliard & Burtt 7102 (E00193964!
lecto. designated here: the eight bulbs with both flowers and leaves enclosed into the envelope placed in the central part of the sheet;
NU0015641! isolecto.).
Comments:—In the light of the recentmost proposal by Mosyakin & McNeill (2022) to amend Art. 8 of the ICN, after which the information
in the protologue will rule, and considering that the protologue in Hilliard & Burtt (1985) only indicates a single collecting date (06
November 1973) in the holotype designation, a lectotypification is needed from the mixed type collection E00193964. Based on the
new considerations, the description made by Martínez-Azorín & Crespo (2014: 1332) is therefore nomenclaturally inoperative, since
it is either superfluous or not valid.
Boosia tenella (Baker) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea tenella Baker, Fl. Cap. (Harvey)
6(3): 464 (1897), basionym (Figs 1.16, 15.4). Type:—SOUTH AFRICA. KwaZulu-Natal. Harrismith (2829): Van Reenen, (–AD),
elev. 5000 ft., 17 December 1891, J. Medley Wood 4562 (K000257340! lecto. designated as “type” by Hilliard & Burtt in Notes Roy.
Bot. Gard. Edinburgh 40: 286. 1982; NH0006437-0! isolecto.).
Boosia umgeniensis (Poelln.) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea umgeniensis Poelln. in Ber.
Deutsch. Bot. Ges. 61: 209 (1944), basionym ≡ Urginea pauciflora Baker in Bull. Herb. Boissier ser. 2, 1: 786 (1901), nom. illeg.
[non Baker, Fl. Trop. Afr. [Oliver et al.] 7(3): 539 (1898)]. Type:—SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930):
Umgeni Falls, (–AC), A. Rehmann 7455 (Z s.n., first-step lecto. designated as “holo.” by Manning & Goldblatt in Strelitzia 40: 36.
2018; second-step lecto. designated here:—Z000027550! lecto.; Z000027549! isolecto.). Note:—A second step lectotypification is
needed due to the presence of two herbarium vouchers with different barcode numbers under the same collection.
4. Bowiea Harv. ex T.Moore & Mast. in Gard. Chron. 1866: 971 (1866), nom. cons. [non Bowiea Haw. in Philos. Mag.
J. 64: 299 (1824), Asphodelaceae] (Figs 16, 17) ≡ Bowiea Harv. ex Hook.f. in Bot. Mag. 93: t. 5619 (1867) [isonym,
Art. 14 Note 1 of the ICN]. Typus generis:—Bowiea volubilis Harv. ex T.Moore & Mast. (holotype). Note:—The
earliest known description of this name was published in The Gardeners’ Chronicle (issue dated 13 October 1866) as
part of an unsigned short note attributable to the journal editors, Thomas Moore and Maxwell T. Masters. These authors
validated simultaneously the species name B. volubilis as a result of a “descriptio generico-specifica” (Art. 38.5 of the
ICN) based on material collected by Cooper in South Africa and grown at Kew, but they attributed the authorship of
the new genus and species to William H. Harvey. Only a few months later, Hooker (1867) described and illustrated the
same plant and commented in detail on the provenance of all material gathered by that time, including Cooper’s plants
at Kew. As all those names share the same type material, Hooker’s newly described genus and species are therefore
treated as later isonyms. However, Bowiea Harv. ex T.Moore & Mast. was illegitimate when published, since it was
homonymous with Bowiea Haw. (Asphodelaceae), but it is currently treated as a conserved name. According to Art. 14
Note 1, the authorship of both generic and specific names is that of the earliest isonym, which is adopted here.
≡ Ophiobostryx Skeels in Bull. Bur. Pl. Industr. U.S.D.A. 223: 45 (1911), syn. subst. Typus generis:—O. volubilis (Harv. ex T.Moore &
Mast.) Skeels (holotype).
≡ Schizobasopsis J.F.Macbr. in Contr. Gray Herb. n.s. 56: 3 (1918), syn. subst. Typus generis:—S. volubilis (Harv. ex T.Moore & Mast.)
J.F.Macbr. (holotype).
FIGURE 16. Bowiea volubilis Harv. ex T.Moore & Mast. from: 1. Hooker f. (1867); 2. Dyer (1941); and 3. Bowiea gariepensis van Jaarsv.
from Reid et al. (1990: pl. 2007).
FIGURE 17. Species of Bowiea Harv. ex T.Moore & Mast. displayed in horizontal rows of images. 1. Bowiea gariepensis van Jaarsv.; 2.
Bowiea volubilis Harv. ex T.Moore & Mast.
Description:—Bulbous geophyte. Bulb partially epigeal and photosynthetic. Roots thickened and branched. Leaves
1‒2, narrowly lanceolate, canaliculate, green, smooth, glabrous, up to 20 cm long, somewhat succulent, only appearing
in young plants and usually absent in adult plants at anthesis. Inflorescence up to 5 m long but usually shorter, strongly
branched, twining, succulent, smooth, glossy green or glaucous, prostrate to climbing on other plants and rocks;
peduncle green, terete, twining and succulent; pedicels curved, to 7 cm long. Bracts lanceolate, green, appressed to
peduncle, lowermost with spur that usually clasps stem; bracteoles absent. Flowers stellate, erect-patent, diurnal, lasting

up to 10 days. Tepals 6, biseriate, 4‒10 mm long, lanceolate to ovate, white to greenish, with greenish longitudinal band
on abaxial side, almost free above joint with ovary, long lasting after capsule development, not abcissing from base.
Stamens 6, spreading, not connivent to style; filaments filiform, free; anthers dorsifixed, dehiscing by longitudinal slits
along their whole length. Ovary conical, wide and short, disposed on wide thickened obtriangular receptacle, giving
the impression of semi-inferior ovary, green, slightly trigonous, with ca. 5‒8 ovules per locule. Style 2‒4 mm long,
erect, white, with trigonous stigma. Capsules ovate to subglobose, truncate to acute, 8‒30 mm long, green, fleshy first
and papyraceous after ripening, valves completely dehiscing from base. Seeds subhemispheric to subfusiform, with
prominent embryo and very narrow, winged margins, 3.8‒5.0 mm long, with black, glossy testa and subisodiametric
or sightly elongated cells, with slightly sunken, periclinal walls.
Number of species and distribution:—Bowiea includes 2 species (Reid et al. 1990): B. gariepensis Van Jaarsveld
(1983: 343) occurring in arid regions of northwestern South Africa and southern Namibia (restricted to the Karoo-
Namib Region), and B. volubilis (currently including also B. kilimandscharica Mildbraed 1934: 202) distributed in
tropical and subtropical regions from southeastern South Africa to East Africa (Fig. 8). It is restricted to the Uzambara-
Zululand and the Zambezian Subregion (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a). A disjunt record
of Bowiea from Angola [Huíla, a 11 kms de Sá da Bandeira [Lubango] estrada para a Tunda-Vala, 13/02/1963, H.
Enriques 117 (K!)] references white flowers, indicating a possibility that this specimen belongs to B. gariepensis,
requiring further studies for confirmation.
Both species are differentiated in morphological, ecological, and biogeographical terms (Baker 1873b, Dyer 1941,
Jessop 1977, Stedje & Thulin 1995, Stedje 1996, Van Jaarsveld 1983, 1992, Reid et al. 1990). For further information
on Bowiea species see Hooker (1867), Skeels (1911), Barschus (1954), Van Jaarsveld (1983, 1992), and Reid et al.
(1990).
Karyology:—2n=20 (Schnarf & Wunderlich 1939, D’Amato 1949, De Wet 1957, Jones & Smith 1967, Bruyns &
Vosa 1987, Stedje & Nordal 1987).
History, diagnostic characters, and taxonomic relationships:—Bowiea is unmistakable by a distinct syndrome
of morphological characters (Figs 16–17), such as branched, fleshy inflorescence; long lasting flowers with nearly free
tepals that remain at the base of the mature capsule; and very wide, short, conical ovary disposed on a wide thickened
obtriangular receptacle, giving the impression of a semi-inferior ovary, facilitating its acceptance by all researchers
working on Urgineoideae since its description by Hooker (1867), including the treatments presented by Manning et al.
(2004) and Manning & Goldblatt (2018). Our phylogenetic analyses (Martínez-Azorín et al. 2023a) place all studied
samples of Bowiea into a well-supported, sister clade to the remaining Urgineoideae in agreement with previous works
(Pfosser & Speta 1999, 2001, 2004, Manning et al. 2004, Pfosser et al. 2012). Only Schizobasis shares the branched
inflorescence with Bowiea, however, the former genus clearly differs by the wiry inflorescence (it is never succulent),
the flowers having a well differentiated superior ovary, and the withered perigone persisting above the capsule as a cap,
together with its distant phylogenetic relationship (Martínez-Azorín et al. 2023a).
Bowiea gariepensis van Jaarsv. in J. S. African Bot. 49(4): 343 (1983) ≡ B. volubilis subsp. gariepensis (van Jaarsv.) Bruyns in Caryologia
40(4): 291 (1988) (Figs 1.17, 16.3, 17.1). Type:—SOUTH AFRICA. Northern Cape. Pofadder (2919): Upper south slope of Groot
Pellaberg, (–AA), 10 August 1982, Van Jaarsveld 6650 (NBG127051! holo.: two herbarium sheets numbered -1 & -2; PRE iso.).
Bowiea volubilis Harv. ex T.Moore & Mast. in Gard. Chron. 1866: 971 (1866) ≡ Ophiobostryx volubilis (Harv. ex T.Moore & Mast.)
Skeels in Bull. Bur. Pl. Industr. U.S.D.A. 223: 45 (1911) ≡ Schizobasopsis volubilis (Harv. ex T.Moore & Mast.) J.F.Macbr. in Contr.
Gray Herb. 56: 3 (1918) (Figs 1.18, 16.1, 16.2, 17.2). Type:—SOUTH AFRICA. Natal [KwaZulu-Natal], 1862, T. Cooper 3263 (K
s.n.! lecto. designated as “holotype” by Stedje & Thulin in Nordic J. Bot. 15(6): 601. 1995).
= B. kilimandscharica Mildbr. in Notizbl. Bot. Gart. Berlin-Dahlem 12: 202 (1934) ≡ Schizobasopsis kilimandscharica (Mildbr.) Barschus
in Kakteen And. Sukk. 5: 65 (1954). Type:—KENYA. Kilimandscharo: Nordseite, Loitokitok, Buschsavanne, elev. 1800 m, 04 April
1934, H.J. Schlieben 5018 (BR876309!, lecto. designated as “holotype” by Stedje & Thulin in Nordic J. Bot. 15(6): 601. 1995; S-
G-7939!, Z-000086938! isolecto.).
5. Drimia Jacq. ex Willd., Sp. Pl. ed. 4, 2(1): 165 (1799) (Figs 18–20). Typus generis:—Drimia elata Jacq.
= Strepsiphyla Raf., Fl. Tellur. 3: 60 (1837). Holotype:—S. villosa (Lindl.) Raf.
= Idothea Kunth, Enum. Pl. [Kunth] 4: 341 (1843). Lectotype:—I. elata (Jacq.) Kunth (designated by Stearn 1978: 203) ≡ Idothearia
C.Presl in Abh. Königl. Böhm. Ges. Wiss., ser. 5, 3: 544 (1845), nom. nov. illeg. [see Crespo et al. in Pl. Syst. Evol. 306: [Art. 67]
9 (2020)].
FIGURE 18. Species of Drimia Jacq. ex Willd. 1. Drimia ciliaris Jacq. ex Willd. from Jacquin (1795: tab. 377); 2. Drimia elata Jacq. ex
Willd. from Jacquin (1794: tab. 373); 3. Drimia media Jacq. ex Willd. from Jacquin (1795: tab. 375); 4. Drimia pusilla Jacq. ex Willd.
from Jacquin (1794: tab. 374).

FIGURE 19. Species of Drimia Jacq. ex Willd. 1. Drimia alta R.A.Dyer from Hooker (1865: t. 5522 as Drimia altissima Hook.f.); 2.
Drimia alta R.A.Dyer from Dyer (1943: t. 890); 3. Drimia robusta Baker from Baker (1870b: t. 190); 4. Drimia villosa Lindl. from
Lindley (1830: t. 1346).
FIGURE 20. Species of Drimia Jacq. ex Willd. displayed in horizontal rows of images. 1. Drimia elata Jacq. ex Willd.; 2. Drimia
haworthioides Baker; 3. Drimia neriniformis Baker; 4. Drimia robusta Baker; 5. Drimia sphaerocephala Baker.

Description:—Bulbous geophyte. Bulb hypogeal or partly epigeal, ovoid to globose, variable in size, bulb scales usually
compact or sometimes loose. Roots thickened and branched. Leaves 1 to several per bulb, commonly hysteranthous,
rarely synanthous, with variable shape and morphology, from glabrous to long cilated or hairy, sometimes undulate.
Inflorescence racemose, usually an elongated and multiflowered, 5‒100 cm long raceme, rarely compact and
subcorymbose; peduncle terete, erect, smooth and glabrous; pedicels subpatent and usually curved, to 20 mm long.
Bracts narrowly lanceolate, lowermost with distinct oblong spur, sometimes the bract and spur connected to peduncle
by whitish membranous portion; bracteoles absent. Flowers stellate, erect-patent, diurnal, lasting 1‒2 days. Tepals 6,
10‒20 mm long, white, yellow, green, brown or purple, fused along basal third to form cylindrical tube, free perigone
segments strongly reflexed and touching basal tube, narrowly lanceolate-oblong, with slightly wider and spatulate
apex, with darker longitudinal band mostly visible on abaxial side. Stamens 6, erect and connivent to style, sometimes
curved upwards; filaments narrowly triangular to filiform, smooth, adnate to perigone tube and arising from its mouth,
rarely very shortly connate above perigone to form short filaments tube; anthers dehiscing by longitudinal slits along
their whole length, yellowish, greenish or purplish. Ovary ovoid, trigonous, pale green to yellowish, well differentiated
from style. Style narrow and long (much longer than ovary), erect. Stigma trigonous, papillose. Capsule ovoid, 7‒13
mm long, green first and papyraceous after ripening, valves completely dehiscing from base; tepals cohering and
inrolled above ovary after anthesis, circumcissile from base and persisting as a cap at the top of developing capsules.
Seeds subellipsoid to reniform, flattened with prominent central embryo and wide, flat wings, 5‒10 mm long, black to
dark brown, with somewhat sinuous anticlinal testa cell walls.
Number of species and distribution:—The genus includes 19 described species distributed in Sub-Saharan
Africa, with the center of diversty in the southwestern parts of South Africa and few species reaching East Africa and
western Central Africa, restricted to the Cape, Karoo-Namib and Uzambara-Zululand Regions, and the Zambezian
and Sahelo-Sudanian Subregions (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a) (Fig. 8). We consider the
taxonomy of Drimia s.str. poor due to the lack of comprehensive studies covering its large distribution range. Further
studies are needed, including of fresh material from the type localities of the described taxa, together with detailed
morphological studies to ascertain the real number of existing taxa. In accepting very wide species concepts, which
cover a large morphological variability, the treatments by Jessop (1977), Stedje (1987), and Manning & Goldblatt
(2018) drastically reduced the number of accepted taxa in Southern and East Africa. We tentatively accept all described
species in the genus based on their morphological differentiations. For further information on Drimia species see
Jessop (1977) and Manning & Goldblatt (2018).
Karyology:—2n=18 (Bruyns & Vosa 1987, as D. haworthioides Baker; De Wet 1957, Fernandes & Neves 1962, as
D. alta R.A.Dyer; Bruyns & Vosa 1987, as D. ciliaris Jacq. ex Willd.); 2n=20 (De Wet 1957, as D. haworthioides).
History, diagnostic characters, and taxonomic relationships:—Jacquin (1794, 1797) described and illustrated
seven species of Drimia (nom. inval.). Among them, Drimia purpurascens Jacquin (1812: 48) and Drimia revoluta
(Linnaeus f. 1782: 204) Sweet (1818: 72), both currently placed in Squilla and Ledebouria Roth (1821: 124),
respectively (Crespo et al. 2020). The remaining species constitute a compact group characterised by tepals basally
connate in a distinct tube, the free portion of tepals elongated and narrowly lanceolate-oblong to subspatulate and
strongly reflexed, and stamens adnate to the perigone, usually slightly curved and connivent to the style. Willdenow
(1799) validated Drimia based on Jacquin’s descriptions and illustrations. This genus is the oldest in Urgineoideae
and presented a homogeneous flower morphology in their original concept, with the exclusion of the two species
mentioned above. However, with the description of further species in Drimia (Baker 1871, 1897, 1898, Jessop 1977,
Stedje 1987, Manning & Goldblatt 2003, 2007, Van Jaarsveld & Van Wyk 2005), its morphological circumscription
became manifestly blurred. Although also based on other characters, this primarily stems from differing perceptions
of the significance of the extent of tepal connation, from nearly free to connate in a distinct tube. This divergence in
opinion introduced massive instability in generic circumscriptions in the Urgineoideae (Huber 1969, Jessop 1977, Deb
& Dasgupta 1982, Stedje 1987, 2001a, 2001b, Manning et al. 2004).
The phylogenetic studies by Martínez-Azorín et al. (2023a) include 17 samples of Drimia constituting a strongly
supported clade, which is sister to a clade combining Litanthus and Schizobasis samples, with moderate support.
The sister genera Litanthus and Schizobasis strongly differ in morphology, as shown below, further supporting the
acceptance of Drimia in a narrow sense.
Drimia alta R.A.Dyer in Fl. Pl. South Africa 23: t. 890 (1943) (Fig. 19.2), nom. nov. ≡ Drimia altissima Hook.f. in Bot. Mag. 91: t. 5522
(1865) (Fig. 19.1), nom. illeg. [non Drimia altissima (L.f.) Ker Gawl. in Bot. Mag. 27: t. 1074 (1808)]. Type:—SOUTH AFRICA.
Natal [KwaZulu-Natal], from John Sanderson of D’Urban [Durban]’, sub illustr. W. Fitch s.n. (K000257364! holo.).
Drimia burchellii Baker in Refug. Bot. [Saunders] 2: App. 2. (1870) ≡ Idothea burchellii (Baker) Kuntze, Revis. Gen. Pl. 2: 712 (1891).
Type:—SOUTH AFRICA. Cape [Eastern Cape], Uitenhage Div., between Galgebosch and Melkrivier, 17 February 1814, Burchell
4769 (K000857424! lecto. designated by Jessop in J. S. African Bot. 43: 285. 1977; TCD ?isolecto. [as Burchell “4969”]).
Drimia ciliaris Jacq. ex Willd., Sp. Pl. 2: 165 (1799) ≡ Hyacinthus ciliaris (Jacq. ex Willd.) Poir. in Lam., Encycl., Suppl. 3: 120 (1813)
≡ Idothea ciliaris (Jacq. ex Willd.) Kunth, Enum. Pl. 4: 343 (1843). Type:—illustration in Jacq., Icon. Pl. Rar. 2 (16): t. 377 (1795)
(Fig. 18.1) (lecto. designated as “type” by Jessop in J. S. African Bot. 43: 284 1977).
Drimia concolor Baker in J. Linn. Soc., Bot. 11: 422 (1871) ≡ Idothea concolor (Baker) Kuntze, Revis. Gen. Pl. 2: 712 (1891). Type:—
SOUTH AFRICA. Cap. B. Spei (e tab. ex exemplo in Hort. Kew. anno 1823 culto depicta descript) [Manning & Goldblatt (2018)
cited that the “holotype” was presumably at K, though we did not find there any material fitting the protologue].
Drimia eckloniana Schult.f., Syst. Veg., ed. 15 bis [Roemer & Schultes] 7(2): 1710 (1830) = Drimia humilis Berg. ex Eckl., Topogr. Verz.
Pflanzensamml. Ecklon: 2 (1827), nom. nud. ≡ Drimia humilis Berg. ex Kunth, Enum. Pl. 4: 344 (1843), nom. inval. [in syn.] ≡
Idothea humilis Kunth, Enum. Pl. 4: 344 (1843), nom. illeg. superfl. Type:—SOUTH AFRICA. In arenosis Prom. B. Spei., Ecklon
s.n. Drimia No. 89 [Manning & Goldblatt in Strelitzia 40: 50 (2018) designated this collection as “holo.”, which was supposedly
kept at B; however the collection is not extant at B (R. Vogt pers. comm.) and therefore, a new type is needed; Lecto. designated
here:—HAL108222! lecto.].
Drimia elata Jacq. ex Willd., Sp. Pl. 2: 165 (1799) ≡ Hyacinthus elatus (Jacq. ex Willd.) Poir. in Lam, Encycl., Suppl. 3: 120 (1813) ≡
Idothea elata (Jacq. ex Willd.) Kunth, Enum. Pl. 4: 343 (1843) (Figs 1.19, 18.2, 20.1). Type:—illustration in Jacq., Icon. Pl. Rar. 2
(15): t. 373 (1794) (Fig. 18.2) (lecto. designated by Stedje & Thulin in Nordic J. Bot. 15: 597. 1995).
Drimia elgonica Bullock in Bull. Misc. Inform. Kew 1932(10): 504 (1932). Type:—KENYA. Mt. Elgon, elev. 6900 ft., 15 March 1931,
E.J. Major & C. Lugard 563 (K000257336! holo.).
Drimia haworthioides Baker in Gard. Chron. (1875) 1: 366 (1875) (Figs 1.20, 20.2). Type:—SOUTH AFRICA. Eastern Cape. Graaff-Reinet
(3224): near Graaff-Reinet, (–BC), Flowers in December 1874 and leaves in February 1875 [ex hort. Kew], Bolus (lecto. designated
here: The illustration in the protologue, Fig. 70 “bulb of Drimia haworthioides”). Epitype (designated here):—K000857416! epi.).
Note:—There are two specimens at Kew mounted on the same voucher (K000857415! and K000857416!) including fragments of
plants that apparently correspond to the type collection (Bolus 2040). However, those fragments were apparently gathered at different
times from plants grown at Kew. The voucher K000857415 includes a bulb resembling the illustration in the protologue, considered
to come from the “type plant” that was gathered 23 Dec 1879 and annotated by R. Brown. The voucher K000857416 was regarded
as “holotype” (to be corrected to lectotype) of this name by Manning & Goldblatt (2018: 57), and includes two fragments (a bulb
and a flowering scape) belonging to two different collections and they are associated to a date (“Dec[ember] 7/[18]76”) that does
not correspond to that cited in the protologue. Therefore, the illustration in the protologue is regarded here as the lectotype of the
concerned name, and the voucher including an inflorescence and a bulb is selected here as epitype.
Drimia hockii De Wild. in Repert. Spec. Nov. Regni Veg. 12: 294 (1913). Type:—Democratic Republic of the Congo.
Ober-Katanga, Elisabethville, September 1911, Ad. Hock s.n. (BR880749! lecto. designated here).
Drimia incerta A.Chev. ex Hutch., Fl. W. Trop. Afr. [Hutchinson & Dalziel] 2: 351 (1936) Type:—IVORY COAST. Hte. Sassandra, Pays
Toura, Mont Dourou, à 1000 m d’altitude, près Koualé, 27 May 1909, Aug. Chevalier 21733 (P02156156! lecto. designated as “type”
by Hutchinson in Kew Bull. 1939: 245. 1939).
Drimia ledermannii K.Krause in Bot. Jahrb. Syst. 51(3-4): 444 (1914). Type:—CAMEROON. Nordkamerun, Zwischen Lubare und
Bakari, in hügeliger Baumsteppe, um 900−1000 m ü. M., 19 June 1909, C. Ledermann 2264 (B 10 0168393! lecto. designated
here).
Drimia media Jacq. ex Willd., Sp. Pl. 2: 166 (1799) ≡ Hyacinthus medius (Jacq. ex Willd.) Poir. in Lam., Encycl., Suppl. 3: 120 (1813)
≡ Idothea media (Jacq.) Kunth, Enum. Pl. 4: 342 (1843). Type:—illustration in Jacq., Icon. Pl. Rar. 2 (16): t. 375 (1795) (Fig. 18.3)
(lecto. designated by Jessop in J. S. African Bot. 43: 282. 1977).
= Drimia rigidifolia Baker in J. Linn. Soc., Bot. 11: 420 (1871) [non Urginea rigidifolia Baker in J. Bot. 16: 323 (1878)] ≡ Idothea
rigidifolia (Baker) Kuntze, Revis. Gen. Pl. 2: 712 (1891). Type:—SOUTH AFRICA. Somerset [?], Bowker 225 (K000857419! lecto.
designated by Jessop in J. S. African Bot. 43: 285. 1977).

Drimia neriniformis Baker, Fl. Cap. (Harvey) 6: 442 (1897) (Figs 2.1, 20.3). Type:—SOUTH AFRICA. Free State. Harrismith (2829):
Van Reenen, (–AD), marshy ground, 5000−6000 ft., 15 November 1892, Wood 4794 (K000257350! lecto. designated by Jessop in J.
S. African Bot. 43: 280. 1977; E00109542!, NH0006430-0!, MO2142490!, P, PRE, SAM0023179-0!, Z, isolecto.).
= Drimia capitata Baker, Fl. Cap. (Harvey) 6(3): 442 (1897) [non Ornithogalum capitatum Hook.f. in Bot. Mag. 89: t. 5388 (1863) ≡
Urginea capitata (Hook.f.) Baker, Fl. Cap. (Harvey) 6(3): 465 (1897) ≡ Fusifilum capitatum (Hook.f.) Speta in Phyton (Horn, Austria)
38(1): 69 (1998)]. Type:—SOUTH AFRICA. [Eastern Cape]. Umtata (3128): Bazeia [Baziya], (–CB), Baur 1160 (K000257353!
lecto. designated by Jessop in J. S. African Bot. 43: 280. 1977).
Drimia pusilla Jacq. ex Willd., Sp. Pl. 2: 165 (1799) ≡ Hyacinthus pusillus (Jacq.) Poir. in Lam., Encycl., Suppl. 3: 120 (1813) ≡ Idothea
pusilla (Jacq.) Kunth, Enum. Pl. 4: 344 (1843) [non Anthericum pusillum Jacq., Icon. Pl. Rar. 2(16): 18, t. 417 (1795) ≡ Fusifilum
pusillum (Jacq.) Speta in Phyton (Horn, Austria) 38: 70 (1998)]. Type:—SOUTH AFRICA. Illustration in Jacq., Icon. Pl. Rar. 2(14):
t. 374 (1794) (Fig. 18.4) (lecto. designated by Jessop in J. S. African Bot. 43: 284. 1977).
Drimia robusta Baker in Refug. Bot. 3: t. 190 (1870) ≡ Idothea robusta (Baker) Kuntze, Revis. Gen. Pl. 2: 712 (1891) (Figs 2.2, 19.3,
20.4). Type:—SOUTH AFRICA. Cape Colony, from Cooper, Hort Saunders 4404, 1869, Cooper s.n. (K000857417! holo.).
Drimia rudatisii Schltr. in Engl. Bot. Jahrb. 40: 89 (1907). Type:—SOUTH AFRICA. KwaZulu-Natal. Port Shepstone (3030): Dumisa,
feuchten schattigen Orten bei Fairfield, (–AD), 24 August 1905, Rudatis 79 (B100167475! holo.).
Drimia sphaerocephala Baker, Fl. Cap. (Harvey) 6: 441 (1897) (Fig. 20.5). Type:—SOUTH AFRICA. Mpumalanga. Komatipoort (2531):
Concession Creek near Barberton, (–CC), 7 September 1890, Galpin 1020 (K000400570! lecto. designated by Jessop in J. S. African
Bot. 43: 280. 1977; BOL140327!, SAM0023178! isolecto.).
Drimia villosa Lindl. in Edwards’s Bot. Reg. 16: t. 1346 (1830) ≡ Strepsiphyla villosa (Lindl.) Raf., Fl. Tellur. 3: 60 (1837) ≡ Idothea
villosa (Lindl.) Kunth, Enum. Pl. 4: 343 (1843). Type:—SOUTH AFRICA. Cape of Good Hope, received by Mr Tate, illustration in
Edwards’s Bot. Reg. 16: t. 1346 (1830) (Fig. 19.4) (lecto. designated here).
Drimia zombensis Baker, Fl. Trop. Afr. [Oliver et al.] 7(3): 525 (1898). Type:—MALAWI. Nyasaland, Mount Zomba, elev. 3000–5000
ft, September 1859, Kirk s.n. (K000400584! holo.).
6. Ebertia Speta in Phyton (Horn, Austria) 38(1): 65 (1998). Typus generis:—Ebertia nana (Oyewole) Speta =
Urginea nana Oyewole (holotype).
Description:—Small bulbous geophyte to ca. 8 cm tall. Bulb hypogeal, ovoid-oblong to subglobose, 2‒3 cm long, with
short bulb neck, outer scales membranous and inner bulb scales white. Roots thickened and branched. Leaves 4‒10,
hysteranthous, 8‒20 cm long and 1‒3 mm wide, thin, filiform and spreading or linear-lanceolate and erect to arching,
green, smooth, glabrous. Inflorescence a short, dense raceme with (1–)3‒6 flowers; peduncle erect, terete, smooth, 1.5‒
4.5 cm long, greenish to dark purple; pedicels spreading, 1‒4 mm long, glabrous; pedicels of fruits laterally recurved,
to 15 mm long. Bracts broadly lanceolate, persistent, lowermost to 6 mm long with distinct flat, decurrent, bifid spur
of ca. 1.5 mm long; bracteoles absent. Flowers campanulate, subpatent. Tepals 6, 5‒10 mm long, free to shortly fused
at base for ca. 1 mm, lanceolate to oblong, spreading, whitish, pinkish or pale greenish with yellow or hyaline margins,
sometimes slightly tinged with dark purple, with distinct dark purple brown longitudinal band on abaxial side, outers
sometimes reflexed. Stamens 6, spreading; filaments 3‒4 mm long, white or greenish; anthers oblong, up to 2 mm long,
opening by longitudinal slits. Ovary elongate oblong to pyramidal, 3‒4 mm long, greenish, sometimes puberulous near
apex, with 10‒15 ovules per locule. Style short, 1‒3 mm long, erect, white, filiform. Capsule ovate to sphaerical, 8‒12
mm long, trigonous with blunt edges in section, mucronate, pale brown. Seeds flattened, dark brown to black, 7‒8 mm
long, with membranous wings, uniseriate per locule.
Number of species and distribution:—Ebertia is known to include 2 species, occurring disjunctly in Central and
West Africa, in Guinea, Sierra Leone, Nigeria and Sudan (Fig. 8). It is restricted to the Guineo-Congolian Region, the
Sahelo-Sudanian Subregion and the Eastern Section of the Zambezian Subregion (sensu Takhtajan 1986 and Martínez-
Azorín et al. 2023a). The species occurs in frequently burnt grasslands on mountain slopes at elevations of 300‒2000
m (Oyewole 1989, Friis & Vollesen 1999). For further information on Ebertia species see Baker (1898), Oyewole
(1989), and Friis & Vollesen (1999).
Karyology:—2n=20 (Oyewole 1988, as Urginea pauciflora (Baker) Baker).
History, diagnostic characters, and taxonomic relationships:—Urginea pauciflora Baker (1898: 539), not to be
confused with the illegitimate U. pauciflora Baker (1901: 786) or Drimia pauciflora Baker (1892: 6), was described by
Baker (1898) based on small plants from Sierra Leone (West Africa) possessing hypogeal bulbs, leaves non-coetaneous
with flowers, a short peduncle, raceme with 2‒3 campanulate flowers, stamens shorter than tepals and with flattened
filaments, and inserted style. Friis & Vollesen (1999) proposed the new name Drimia sudanica Friis & Vollesen for
U. pauciflora and extended the range of this species from West Africa (Sierra Leone and Guinea) to southern Sudan.
These authors added important characters such as the very short raceme with 4‒6 flowers, lanceolate bracts with a
flat, decurrent spur (a character not mentioned by Baker, but matching the type material at Kew herbarium), pedicels
1.5‒2.0 mm long, tepals 5‒7 mm long, shortly connate for ca. 1 mm at the base and spreading, with a dark purple-
brown midrib, stamens 3‒4 mm long, filaments spreading, ovary elongate and ca 3 mm long, and very short style. Ripe
capsules and seeds were not studied.
A related species was described by Oyewole (1989) as Urginea nana Oyewole (1989: 623) from Nigeria, showing
small, hypogeal, compact bulbs, filiform erect to spreading leaves that are not coetaneous with flowers, peduncle erect,
up to 4 cm long, raceme short and compressed with up to 6 flowers, pedicels reflexed during fruit development, tepals
6‒10 mm long, free, the outers reflexed at anthesis, ovary pyramidal, 3‒4 mm long, style short and erect, capsule
ovoid-spherical with mucronate apex, seeds flattened, subellipsoid, 7‒8 mm long, dark brown to black, and uniseriate
per locule. The author also illustrated a dehisced capsule showing widely open valves, similar to those found in D.
khubusensis P.C.van Wyk & J.C.Manning in Manning & Goldblatt (2018: 120), here considered to belong to Iosanthus.
Further, Oyewole (1989) described a considerable morphological variation and three morphotypes in Urginea nana;
further studies are needed to improve our understanding of this taxon.
Speta (1998b) published Ebertia to include the tropical African taxa U. pauciflora and U. nana, being characterised
by the hypogeal compact bulbs, filiform, proteranthous leaves, short peduncle and condensed few-flowered raceme,
shortly spurred bracts, straight and patent pedicels, nocturnal, campanulate flowers with tepals shortly connate at base,
filaments shorter than tepals, ovary inferior [examination of the type confirms this as erroneous], with 12 ovules per
locule, style slightly thickened at the apex, pedicels of ripe capsules laterally recurved, capsules ovoid to globose, and
flattened black seeds.
The phylogenetic analyses of Pfosser & Speta (1999) place a sample of E. nana as sister to a sample from Senegal
named `Thuranthos indicum´ sensu lato, but fitting with Vera-duthiea in the sense of the present work. Pfosser & Speta
(2001, 2004) extended the study to include four samples of E. nana (Oyewole 1989: 623) Speta (1998b: 68) from
Guinea and Senegal that constituted a well supported clade sister to “Duthiea” species (= Vera-duthiea) from Senegal
and Botswana. The recent phylogenetic analyses by Martínez-Azorín et al. (2023a) included three samples of Ebertia
from Guinea and Senegal and form a strongly supported clade that is sister to Vera-duthiea.
Based on the above-mentioned evidence, we accept Ebertia as including E. pauciflora (Baker 1898: 539) Speta
(1998b: 68) and E. nana, taxa flowering shortly after grassland fires (Oyewole 1989, Friis & Vollesen 1999). These
two species share a small size, non-coetaneous and filiform leaves, short peduncle and condensed, few-flower racemes,
subcampanulate flowers with almost free, suberect to slightly spreading tepals, stamens shorter than tepals and included,
an ovoid to conical ovary ca. 3 mm long which is sometimes puberulous, and short and erect style of equal length to the
ovary. Further, E. nana is unique in the Urgineoideae in having reflexed pedicels that subtend the ripe capsules, since
the remaining subfamily members always support ripe capsules on erect to suberect pedicels, except for the northwest
African Spirophyllos. Further studies are necessary to compare this character with U. pauciflora. The widely spreading
capsule valves and seed morphology in U. nana approach Iosanthus, a genus that is related to the clade comprising
Urginea, Spirophyllos, and Indurgia (Martínez-Azorín et al. 2023a).
Ebertia nana (Oyewole) Speta in Phyton (Horn, Austria) 38(1): 68 (1998) ≡ Urginea nana Oyewole in Ann. Missouri Bot. Gard. 76(2): 623
(1989) ≡ Drimia minuta J.C.Manning & Goldblatt in Bothalia 43(1): 77 (2013), nom. nov. ≡ Drimia nana (Oyewole) J.C.Manning
& Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004), nom. illeg. [non Drimia nana (Snijman) J.C.Manning & Goldblatt in Bothalia
33(1): 111 (2003)]. Type:—NIGERIA. Ilorin, near the academic area, University of Ilorin, S00/2111 (IUH holo.; FHI iso.).
Ebertia pauciflora (Baker) Speta in Phyton (Horn, Austria) 38(1): 68 (1998) ≡ Urginea pauciflora Baker, Fl. Trop. Afr. [Oliver et al.] 7(3):
539 (1898), basionym ≡ Drimia sudanica Friis & Vollesen in Nordic J. Bot. 19(2): 210 (1999), nom. nov. [non Urginea pauciflora
Baker in Bull. Herb. Boissier ser. 2, 1: 786 (1901), nom. illeg., nec Drimia pauciflora Baker (1892)]. Type:—SIERRA LEONE.
Near Wallia, on the River Scarcies, 11 February 1892, G.F. Scott-Elliot 4580 (K000257328! lecto. designated by Friis & Vollesen in
Nordic J. Bot. 19(2): 210. 1999; BM! isolecto.).

7. Fusifilum Raf., Fl. Tellur. 2: 27 (1837) (Figs 21–23). Typus generis:—Drimia physodes (Jacq.) Jessop (lectotype,
designated by Deb & Dasgupta in J. Econ. Tax. Bot. 3: 823. 1982).
= Physodia Salisb., Gen. Pl.: 37 (1866) ≡ Urginea sect. Physodia (Salisb.) Baker in J. Linn. Soc., Bot. 13: 216 (1873) ≡ Drimia sect.
Physodia (Salisb.) J.C.Manning & Goldblatt in Strelitzia 40: 92 (2018). Typus generis:—Physodia pusilla (Jacq.) U.Müll.-Doblies
(lectotype, designated by Deb & Dasgupta in J. Econ. Tax. Bot. 3: 823. 1982).
FIGURE 21. Species of Fusifilum Raf. 1. Fusifilum physodes (Jacq.) Speta from Jacquin (1795: t. 1046, as Anthericum physodes Jacq.);
2. Fusifilum pusillum (Jacq.) Speta from Jacquin (1795: t. 417, as Anthericum pusillum Jacq.); 3. Fusifilum capitatum (Hook.f.) Speta from
Hooker f. (1863: t. 5388, as Ornithogalum capitatum Hook.f.).
Description:—Small bulbous geophyte. Bulb hypogeal, ovoid to subglobose, 1‒7 cm in diam., sometimes with short
bulb neck, solitary or commonly clump forming, inner scales fleshy, white and compact, outer scales membranous.
Roots thickened and branched. Leaves 1‒10, hysteranthous, 3‒15 cm long and 1‒30 mm wide, usually flat, canaliculate,
lanceolate, spreading, green, glabrous or papillose. Inflorescence a dense, short or elongated raceme, 1‒8 cm long,
with 6‒60(–120) flowers; peduncle erect, terete, smooth or papillose, 4‒15 cm long, greenish to dark purple; pedicels
patent-spreading, 8‒30 mm long, glabrous. Bracts lanceolate, small, persistent, lowermost 1‒2 mm long with short
spur of ca. 1 mm long; bracteoles absent. Flowers stellate, suberect, diurnal, opening in afternoon. Tepals 6, biseriate,
4‒7 mm long, spreading, white, with distinct greenish to purplish longitudinal band on abaxial side, free to shortly
fused at base for less than 1 mm, outers sometimes slightly reflexed. Stamens 6, spreading; filaments 3‒4 mm long,
fusiform, narrowed to base and apex, white, with distinct papillae along basal third; anthers oblong, medifixed, to 1
mm long, opening by longitudinal slits. Ovary oblong, truncate to style, 2‒4 mm long, white, with 10‒15 ovules per
locule. Style 2‒3 mm long, erect, white, filiform. Capsule ovate to sphaerical, trigonous with blunt edges in cross-
section, 6‒12 mm long, pale brown. Seeds subellipsoid, (3–)4‒8 mm long, flattened with prominent central embryo
and wide, flat or incurved wings, dark brown to black, glossy, with sinuous anticlinal testa cell walls.
Number of species and distribution:—It includes 17 species, occurring in Southern Africa (Fig. 8) and restricted
to the Cape, Karoo-Namib and Uzambara-Zululand Regions, and the Southern Section of the Zambezian Subregion
(sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a). For further information on Fusifilum species see Müller-
Doblies et al. (2001). We preliminarily accept F. magicum as originally circumscribed as belonging to Fusifilum based
on its flower morphology (Fig. 23.1). However, its phylogenetic position places two samples of this species in a clade
related to Urginea revoluta and Boosia (Martínez-Azorín et al. 2023a). Further studies, including new phylogenetic
evidence, are needed to evaluate other alternatives.
FIGURE 22. Species of Fusifilum Raf. displayed in horizontal rows of images. 1. Fusifilum capitatum (Hook.) Speta; 2. Fusifilum
crenulatum U.Müll.-Doblies et al.; 3. Fusifilum depressum (Baker) U.Müll.-Doblies et al.
Karyology:—2n=20 (De Wet 1957, as Urginea depressa Baker).

History, diagnostic characters, and taxonomic relationships:—Fusifilum was described to accommodate some
species of Urgineoideae having small to medium-sized, subcapitate inflorescence; white stellate, flowers; fusiform and
papillate filaments; and white gynoecium. The latter character is unique and diagnostic. The original concept of the
genus included species described and illustrated by Jacquin (1795) from Southern Africa and by Ruiz & Pavón (1802)
from South America, the latter belonging to Oziroe Rafinesque (1837: 53). However, the current concept of Fusifilum
excludes the American taxon, and Deb & Dasgupta (1982) typified the genus on F. physodes (Jacquin 1795: 18) Speta
(1998b: 69) (Müller-Doblies et al. 2001, Martínez-Azorín et al. 2015). Later, other generic names, such as Physodia
Salisbury (1866: 37), were applied to this distinct group of plants.
Speta (1998b) accepted Fusifilum to include seven species, and Müller-Doblies et al. (2001) presented a taxonomic
revision that included 16 species, four from Speta’s (1998b) treatment, one from Tang & Weiglin (2001), 10 as new,
and one new combination. Types of their newly described species are still not deposited in most of the studied herbaria
and their morphological descriptions are incomplete, which has impeded further taxonomic work in the genus during
the last two decades.
The phylogenetic analyses of Manning et al. (2004) included four samples considered by them to belong to
Fusifilum, although only F. physodes is a true Fusifilum, following circumscriptions of Müller-Doblies et al. (2001)
and the present work. The supposed polyphyly of Fusifilum, in the sense of Manning et al. (2004), was one of the
main reasons offered to justify the lumping of the evidently broad diversity in Urgineoideae in a single, extremely
variable Drimia, excluding Bowiea. However, the correct genus adscription of these species does, in fact, dissolve the
polyphyly of Fusifilum and support an analytic rather than a synthetic treatment in the subfamily (Martínez-Azorín et
al. 2023a) since ‘F. dregei’ is a species of Urgineopsis Compton (as accepted in this work) and ‘F. marginata’ belongs
to Austronea (Martínez-Azorín et al. 2019a)—a genus sister to Fusifilum. The names Fusifilum calcarata and F.
marginata sensu Manning et al. (2004) were never formally combined or published prior to 2004.

FIGURE 23. Species of Fusifilum Raf. displayed in horizontal rows of images. 1. Fusifilum magicum U.Müll.-Doblies et al.; 2. Fusifilum
montanum (A.P.Dold & E.Brink) A.P.Dold et al.; 3. Fusifilum physodes (Jacq.) Speta.
Manning & Goldblatt (2018) described Drimia sect. Physodia, which mostly fits the concept of Fusifilum in this
work, but excluding Urginea virens that belongs to Austronea (Martínez-Azorín et al. 2019a). Manning & Goldblatt
(2018) placed eight of the species described by Müller-Doblies et al. (2001) under their Drimia physodes (Jacquin 1795:
18) Jessop (1977: 300), commenting that those names were not validly published since the types were not deposited
following Art. 40.7 of the ICN. However, this interpretation is incorrect, as the Shenzhen Code only recommends that
the type material, especially the holotype, should be deposited in a public herbarium or other public collection allowing
access and warranting long-term conservation (Art. 7 Rec. 7A1 of the ICN). Therefore, these Müller-Doblies names
are valid following the current Shenzhen Code, and hence must be accepted. Our field work in Southern Africa over
the last decades has enabled us to observe clear morphological differences in support of the acceptance of the species
of Müller-Doblies et al. (2001). Our phylogenetic analyses show 20 samples of Fusifilum, including several type
collections provided by U. & D. Müller-Doblies, forming a strongly supported clade sister to Austronea (Martínez-
Azorín et al. 2023a), further supporting the acceptance of this genus based on a unique syndrome of morphological
characters, biogeography, and phylogenetic relationships.
Accepted species and required new combination:—
Fusifilum bruce-bayeri U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 479 (2001). Type:—SOUTH AFRICA.
Western Cape. Worcester (3319): Worcester West, Polhillia waltersii Nature Reserve, (–CB), elev. 250 m, 15 August 1984, Müller-
Doblies 84017f (PRE holo.; B, BTU, E, G, GRA, K, M, MO, NAS, NBG, S, W, WIND, Z iso.; ex cult. BTU5931: ibidem. Not
deposited in the cited herbaria).
Fusifilum capitatum (Hook.f.) Speta in Phyton (Horn, Austria) 38(1): 69 (1998) ≡ Ornithogalum capitatum Hook.f. in Curtis’s Bot. Mag.
89: t. 5388 (1863), basionym (Figs 2.3, 21.3) [non Drimia capitata Baker (1897)] ≡ Urginea capitata (Hook.f.) Baker, Fl. Cap.
(Harvey) 6(3): 465 (1897) ≡ Physodia capitata (Hook.f.) U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 107(5−6):
520 (1996) (Fig. 22.1). Type:—SOUTH AFRICA. British Kaffraria [part of Eastern Cape today], 1860, Cooper 208 (K000365544!
lecto. designated as “holo” by Manning & Goldblatt in Strelitzia 40: 93. 2018; Z-000086255! isolecto.).
Fusifilum crenulatum U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 483 (2001) (Fig. 22.2). Type:—SOUTH
AFRICA. Northern Cape. Kamiesberg (3018): Koringlandskloof, 1 km on Leliefontein road from Garies/Platbakkies road, (–AC),
loamy ground, elev. 1200 m, 12 August 1980, Müller-Doblies 80087b (PRE holo.; B, BTU, G, K, M, MO, NAS, NBG, S, WIND, Z
iso.; ex cult. BTU4033: ibidem. Not deposited in the cited herbaria).
Fusifilum depressum (Baker) U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 480 (2001) ≡ Urginea depressa
Baker in Bull. Herb. Boissier ser. 2, 4: 1000 (1904), basionym ≡ Drimia depressa (Baker) Jessop in J. S. African Bot. 43: 297 (1977)
(Fig. 22.3). Type:—SOUTH AFRICA. Gauteng. Johannesburg (2628): Transvaal, Modderfontein, (–CB), September 1897, Conrath
687 (Z000027555! lecto. designated here). Note:—The lectotype designation by Jessop in J. S. African Bot. 43: 297 (1977) is not
acceptable according to Art. 8.2 Ex. 3 of the Code (see Martínez-Azorín & Crespo 2014; the voucher K000857414! bears a different
collection date of 9 October 1898, perhaps erroneously transcribed. In that case it is a true isolectotype, as well as GZU!).
Fusifilum emdeorum J.S.Tang & Weiglin in Feddes Repert. 112(7−8): 505 (2001). Type:—SOUTH AFRICA. Western Cape. Worcester
(3319): Farm Gevonden, E end of Dutoitskloof, (–CA), elev. 300 m, 22 July 1986, Müller-Doblies 86017c (PRE holo.; B, BTU, K,
M, MO, NBG, S, WIND, Z iso.; ex cult. BTU6466: ibidem and G, GRA, LI, NAS, W. Not deposited in the cited herbaria).
Fusifilum gifbergense U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 479 (2001). Type:—SOUTH AFRICA.
Western Cape. Vanrhynsdorp (3118): Snorkfontein, half way up the Gifberg, (–DD), elev. 500−600 m, 16 August 1980, Müller-
Doblies 80105v (PRE holo.; B, BTU, E, G, K, LI, M, MO, NAS, NBG, P, S, W, WIND, Z iso.; ex cult. BTU4239: ibidem. Not
deposited in the cited herbaria).
Fusifilum glaucum U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 481 (2001). Type:—SOUTH AFRICA.
Western Cape. Vanrhynsdorp (3118): Zandkraal, Gifberg road, 2.7 km S of turn off to Zandkraal, (–DB), elev. 150 m, 16 August 1980
veg., Müller-Doblies 80104e (PRE holo.; B, BTU, K, M; ex cult. BTU4201: B, BTU, G, K, LI, M, MO, NAS, NBG, PRE, S, WIND,
Z iso. Not deposited in the cited herbaria).
Fusifilum hei U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 480 (2001). Type:—SOUTH AFRICA. Western
Cape. Montagu (3320): Matjesfontein, Farm Boelhouer (Bullover), 4 km on Keurfontein (Driekop) road, (–BA), elev. 1050 m, 02
August 1986 veg., Müller-Doblies 86032a (PRE holo.; B, BTU, K, NAS, S iso; ex cult. BTU6607: ibidem. Not deposited in the
cited herbaria).
Fusifilum magicum U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 491 (2001) (Fig. 23.1). Type:—SOUTH
AFRICA. Western Cape. Beaufort West (3222): Boesmansklip, 2.5 km from N1 to Molteno Pass, (–BC) , elev. 900 m, 15 January
1984, Müller-Doblies 84129c (PRE holo; B, BTU, K, NBG. Not deposited in the cited herbaria).
Fusifilum minus (A.V.Duthie) Speta in Phyton (Horn, Austria) 38(1): 69 (1998) ≡ Urginea minor A.V.Duthie in Ann. Univ. Stellenbosch
6. Sect. A, No. 2, 11 (1928), basionym ≡ Drimia minor (A.V.Duthie) Jessop in J. S. African Bot. 43(4): 306 (1977) ≡ Physodia minor
(A.V.Duthie) Salisb. ex U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 107(5−6): 520 (1996). Type:—SOUTH
AFRICA. Western Cape. Cape Town (3318): Stellenbosch Flats, (–DD), March 1924, Duthie s.n. STE1546 (NBG0197706! lecto.
designated as “holo.” by Jessop in J. S. African Bot. 43: 306. 1977); K000257366!, PRE0049735! isolecto.).
Fusifilum montanum (A.P.Dold & E.Brink) A.P.Dold, Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig comb. nov. ≡ Drimia montana
A.P.Dold & E.Brink in Bothalia 36(1): 64 (2006) basionym ≡ Drimia montana A.P.Dold & E.Brink in Taxon 63: 1130 (2014), nom.
superfl. ≡ Fusifilum montanum (A.P.Dold & E.Brink) A.P.Dold, Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa
201(2): 168 (2015), comb. inval. (Figs 2.4, 23.2). Type:—SOUTH AFRICA. Eastern Cape. Fort Beaufort (3226): Groot Winterberg,
The Hoek farm, (–AD), elev. 2150 m, 1 January 2004, A.P. Dold 4633 (GRA0000460! lecto. designated here: only the plants in
flower and fruit lacking leaves “Pressed in December at time of collection” [though corrected to January by A.P. Dold 08/10/2014;
A.P. Dold pers. comm.], which are placed on the upper half of the sheet.).
in the protologue will rule, and considering that the protologue in Dold & Brink (2006) only indicates a single collecting date (01-

01-2004) in the holotype designation, a lectotypification is needed from the mixed type collection GRA0000460. Based on the new
considerations, the description made by Martínez-Azorín & Crespo (2014: 1330) and the combination in Martínez-Azorín et al.
(2015: 168) are therefore nomenclaturally inoperative, since it is either superfluous or not valid.
Fusifilum oliverorum U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 487 (2001). Type:—SOUTH AFRICA.
Western Cape. Bredasdorp (3420): Moerasfontein, 1.3 km NW of Skipskop/Bredasdorp road, (–CB), elev. 30 m, 18 August 1984
veg., Oliver & Müller-Doblies 84025a (PRE holo.; B, BTU, K, M, MO, NBG, S, WIND, Z iso; ex cult. BTU 6000, 6001, 6002, 6003:
ibidem. Not deposited in the cited herbaria).
Fusifilum papillosum U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 487 (2001). Type:—SOUTH AFRICA.
Northern Cape. Calvinia (3119): Glen Lyon States, SE of Nieuwoudtville, (–AC), elev. 750 m, 9 November 1978 fl., Müller-Doblies
78072m (PRE holo.; B, BTU iso; ex cult. BTU 2156: B, BTU, K, NBG, PRE. Not deposited in the cited herbaria).
Fusifilum physodes (Jacq.) Speta in Phyton (Horn, Austria) 38(1): 69 (1998) ≡ Anthericum physodes Jacq., Icon. Pl. Rar. 2(16): 18, t. 418
(1795) diagnosis; Coll. Suppl.: 94 (1797) description ≡ Phalangium physodes (Jacq.) Pers., Syn. Pl. 1: 369 (1805) ≡ Albuca physodes
(Jacq.) Ker Gawl. in Curtis’s Bot. Mag. 26: t. 1046 (1807) ≡ Caesia physodes (Jacq.) Spreng., Syst. Veg. 2: 88 (1825) ≡ Idothea
physodes (Jacq.) Kunth, Enum. Pl. 4: 345 (1843) ≡ Urginea physodes (Jacq.) Baker in J. Linn. Soc., Bot. 13: 217 (1873) ≡ Physodia
physodes (Jacq.) U.Müll.-Doblies et al. in Feddes Repert. 107(5−6): 519 (1996) ≡ Drimia physodes (Jacq.) Jessop in J. S. African
Bot. 43(4): 300 (1977) (Figs 2.5, 21.1, 23.3). Type:—SOUTH AFRICA. Illustration in Jacq., Icon. Pl. Rar. 2(16): t. 418 (1795) (Fig.
21.1) (lecto. designated by Jessop in J. S. African Bot. 43: 300. 1977).
= Ornithogalum melanopus Dinter ex Sölch in Beitr. Fl. SW-Afr., Diss. Univ. München: 70 (1960).
Fusifilum pusillum (Jacq.) Speta in Phyton (Horn, Austria) 38(1): 70 (1998) ≡ Anthericum pusillum Jacq., Icon. Pl. Rar. 2(16): 18, t. 417
(1795) diagnosis; Coll. Suppl.: 95 (1797) description, basionym [non Drimia pusilla Jacq., Coll. Suppl.: 42 (1797)] ≡ Caesia pusilla
(Jacq.) Spreng., Syst. Veg. 2: 88 (1825) ≡ Idothea drimioides Kunth, Enum. Pl. 4: 345 (1843), nom. nov. [non I. pusilla (Jacq.) Kunth
= D. elata Jacq.] ≡ Urginea pusilla (Jacq.) Baker in J. Linn. Soc., Bot. 13: 217 (1873) ≡ Physodia pusilla (Jacq.) U.Müll.-Doblies et
al. in Feddes Repert. 107(5−6): 520 (1996). Type:—SOUTH AFRICA. Illustration in Jacq., Icon. Pl. Rar. 2 (16): t. 417 (1795) (Fig.
21.2) (lecto. designated by Jessop in J. S. African Bot. 43: 295. 1977).
Fusifilum spirale U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 483 (2001). Type:—SOUTH AFRICA.
Northern Cape. Springbok (2917): Henrietsberg, 10−12 km N of Steinkopf, (–BB), elev. 800−1100 m, 14 August 1980, Müller-
Doblies 80096a (PRE holo.; B, BTU, K, MO, NAS, WIND, Z iso; ex cult. BTU 4085: G, LI, M, NBG, S. Not deposited in the cited
herbaria).
Fusifilum stoloniferum U.Müll.-Doblies, J.S.Tang & D.Müll.-Doblies in Feddes Repert. 112(7−8): 484 (2001). Type:—SOUTH AFRICA.
Western Cape. Beaufort West (3222): Karoo National Park, escarpment W of tall TV-Tower (N15D), (–AD), elev. 1800 m, 27 August
1984 veg., Müller-Doblies 84051c (PRE holo.; B, BR, BTU, K, LI, M, MO, NAS, NBG, S, WIND, Z iso; ex cult. BTU 6110,
6111, 6112: ibidem and BOL, E, G, GRA, KMG, NH, NU, P, PE, PEU, PRU, UNIN, W, WAG, WRSL. Not deposited in the cited
herbaria).
8. Geschollia Speta in Stapfia 75: 169 (2001) (Figs 24–26). Typus generis:—G. anomala (Baker) Speta (holotype).
≡ Ornithogalum subgen. Ledebouriopsis Baker (1870a: 178) ≡ Drimia subgen. Ledebouriopsis (Baker) Baker (1897: 437) ≡ Drimia sect.
Ledebouriopsis (Baker) J.C.Manning & Goldblatt (2018: 24) pro parte (only Drimia anomala, D. occultans, and D. calcarata s.str.).
Typus subgeneris:—Drimia anomala (Baker) Baker (lectotype designated by Manning & Goldblatt 2018).
Description:—Bulbous geophyte. Bulb hypogeal or rarely semi-epigeal, mostly solitary but rarely proliferous,
usually with compact scales but rarely loose, outer scales brownish and membranous. Roots thickened and branched.
Leaf solitary (rarely 2‒3 in clump forming species), terete, green, commonly leathery, synanthous or hysteranthous,
deciduous to evergreen, 0.5−7.0 mm in diam., smooth, glabrous, usually withering from tip showing transverse
abscission plates. Inflorescence 1 or rarely 2 per bulb, racemose, usually elongated, (0.2‒)2‒30 cm long, erect or
slightly bent; peduncle elongated, erect, smooth or sometimes distinctly papillose at base; pedicels 2‒30 mm long,
subpatent. Bracts lanceolate, acute, lowermost with broad spur usually longer than blade; bracteoles absent. Flowers
stellate, erect-patent, diurnal, usually opening in afternoon and withering in evening. Tepals 6, biseriate, 3.2‒8.0 mm
long, spreading to reflexed at full anthesis, with base usually shortly connate for ca. 1 mm but almost free in general
appearance, yellowish, orange, greenish or white, with discrete brownish, green or purple stripe along middle, more
evident on abaxial side. Stamens 6, erect to spreading; filaments filiform, slightly fusiform, adnate to base of tepals,
commonly glabrous and smooth, rarely distinctly papillate; anthers yellow, oblong, medifixed, dehiscing along their
whole length. Ovary ovate to oblong, attenuate to truncate at top, green to yellowish, sometimes with white maculae;
style white, narrowly filiform to obtriangular, distinctly trigonous in section, as long as or longer than ovary; stigma
slightly 3-lobed and papillose. Capsule ovate-globose, small, 3‒6 mm long, trigonous, loculicidal, 3 valves splitting
to base, with withered perigone segments circumscissile below and forming apical cap. Seeds polygonal or irregularly
compressed, commonly narrowly pyramidal, pointed, comparatively small, 1.0‒2.4(‒3.0) mm long, light brown to
black, with wrinkled-rugose testa and sinuous anticlinal cell walls.
FIGURE 24. Species of Geschollia Speta. 1. Geschollia anomala (Baker) Speta from Baker (1870a: t. 178, as Ornithogalum anomalum
Baker); 2. Geschollia anomala (Baker) Speta from Dyer (1951: t. 1117, as Drimia anomala (Baker) Baker); 3. Geschollia calcarata (Baker)
Mart.-Azorín et al.; lectotype of Ornithogalum calcaratum Baker designated by Stedje (1987); illustration by W.H. Fitch corresponding
to the collection K001291881 © Reproduced with the consent of the Royal Botanic Gardens, Kew.
Number of species and distribution:—Geschollia currently includes nine species, eight sensu Martínez-Azorín
et al. (2019d) together with D. loedolffiae Van Jaarsv. in Van Jaarsveld & Van Wyk (2006: 50) based on the inflorescence
drying soon after flower withering, the small capsule and seed, and distribution. Species of the genus show their center
of diversity in south-central South Africa from the Little Karoo in the West to the Eastern Cape in the East, with one
species extending to southern Namibia and northwestern parts of the Northern Cape Province, and some disjunct
populations occurring in southwestern and eastern South Africa (Fig. 8). The genus is therefore restricted to the Cape,
Karoo-Namib, and Uzambara-Zululand Regions (sensu Takhtajan 1986). For further information on Geschollia species
see Baker (1870a, 1874b), Van Jaarsveld & Van Wyk (2006), Williamson (2012), Manning & Goldblatt (2018), and
Martínez-Azorín et al. (2019d).
History, diagnostic characters, and taxonomic relationships:—Ornithogalum anomalum was described by
Baker (1870a) based on a collection of a South African plant with a single, terete leaf, long raceme, shortly connate
tepals reflexed at full anthesis, and spreading stamens. Baker (1870a) also pointed to apparent differences with typical
Ornithogalum on which account he described O. subg. Ledebouriopsis Baker (1870a: t. 178) to accommodate this new
species. Baker (1897) later placed this species in Drimia and made the first mention of the spurred condition of the
bracts. Dyer (1951) noted the peculiar habit of the single terete leaf in D. anomala (Baker 1870a: t. 178) Baker (1897:
442), stating that it “withers from the tip, the withering process being arrested at different stages”.

FIGURE 25. Species of Geschollia Speta displayed in horizontal rows of images. 1. Geschollia anomala (Baker) Speta; 2. Geschollia
brachyandra Mart.-Azorín et al.; 3. Geschollia globuligera Mart.-Azorín et al.; 4. Geschollia loedolffiae (van Jaarsv.) Mart.-Azorín et
al.
The phylogenetic analyses of Pfosser & Speta (2001, 2004) included samples of Drimia anomala from South
Africa, which formed an independent, well-supported clade within Urgineoideae. This led Speta (2001) to describe
the monotypic Geschollia, showing a single terete leaf (rarely two leaves), a long and multiflowered racemose
inflorescence, short pedicels, filiform and, patent filaments, white, pink, yellow or greenish tepals connate for ca. 1 mm
at the base, and angulose seeds, 1.5‒2.0 mm long. Pfosser & Speta (2001, 2004) found Geschollia to form a clade sister
to Boosia and Urgineopsis, although the relationship received only weak statistical support. However, later analyses
(Pfosser et al. 2012) determined Geschollia and Urgineopsis to be sister, well-supported monophyletic genera, with
moderate support.
The lastest revision of Urgineoideae in Southern Africa by Manning & Goldblatt (2018) includes Drimia anomala
in D. sect. Ledebouriopsis merging eleven species with variable morphologies. This assemblage is polyphyletic as
revealed by previous phylogenetic studies (Pfosser & Speta 2001, 2004) as well as the recent phylogenetic work by
Martínez-Azorín et al. (2023a) where 22 samples of Geschollia form a strongly supported clade sister to Boosia. These
latter two genera share some morphological characters, such as the terete leaves or the usually elongated inflorescence,
but they differ by the mostly single leaves and small capsules and seeds in Geschollia, a genus largely confined to
southern central South Africa, and by the usually more numerous leaves, with usually larger, elongated, flattened seeds
in Boosia—a genus centered in eastern South Africa and extending northwards through eastern Africa. Therefore, we
here accept Geschollia at genus rank in the sense of Martínez-Azorín et al. (2019d), with the addition of D. loedolffiae.
All nine species share the main diagnostic characters of Geschollia, these being the single (rarely 2), terete leaf and
comparatively small capsules with small polygonal or irregularly compressed, angled seeds.
FIGURE 26. Species of Geschollia Speta displayed in horizontal rows of images. 1. Geschollia longipedicellata Mart.-Azorín et al.; 2.
Geschollia occultans (G.Will.) Mart.-Azorín et al.; 3. Geschollia prolifera Mart.-Azorín et al.; 4. Geschollia zebrina Mart.-Azorín et al.
Geschollia anomala (Baker) Speta in Stapfia 75: 169 (2001) ≡ Ornithogalum anomalum Baker in Refug. Bot. [Saunders]: t. 178 (1870),
basionym ≡ Drimia anomala (Baker) Baker, Fl. Cap. (Harvey) 6(3): 442 (1897) (Figs 2.6, 24.1, 24.2, 25.1). Type:—SOUTH
AFRICA. Cape of Good Hope, “sent from South Africa by Mr Thos. [Thomas] Cooper”. Illustration in Baker (1870a: Plate 178)
(Fig. 24.1) (holo.). Epitype (designated by Martínez-Azorín et al. 2019d):—Fort Beaufort (3226): 22 km south of Bedford on R350,
Normandale Farm, ca. 160 m SE from farmhouse, (–CC), elev. 649 m, 8 December 2018 (in flower and fruit), A.P.Dold 16047 (GRA!
epi.).

= Urginea eriospermoides Baker in Gard. Chron. ser. 3, 2: 126 (1887). Type:—SOUTH AFRICA, MacOwan 292-72 (K000257355!
holo.).
Geschollia brachyandra Mart.-Azorín, A.P.Dold & M.B.Crespo in Phytotaxa 427(2): 94 (2019) ≡ Drimia brachyandra (Mart.-Azorín,
A.P.Dold & M.B.Crespo) J.C.Manning & Goldblatt in Bothalia 52(1): 3 (2022) (Figs 2.7, 25.2). Type:—SOUTH AFRICA. Eastern
Cape. Grahamstown (3326): 19 km from Grahamstown on Cradock road, Brakloof (now Brack Kloof), 500 m from turn-off onto
farm drive, 20 m off of road verge, (–AD), elev. 690 m, 27 November 1993 (in flower and fruit), A.P. Dold 438 (GRA! holo.).
Geschollia calcarata (Baker) Mart.-Azorín, M.B.Crespo, A.P.Dold, M.Pinter & Wetschnig in Phytotaxa 427(2): 97 (2019) ≡ Ornithogalum
calcaratum Baker in Gard. Chron. n.s., 1: 723 (1874), basionym. = Drimia calcarata (Baker) Stedje in Nordic J. Bot. 7(6): 663
(1987) (Fig. 2.8). Type:—SOUTH AFRICA. Eastern District of Cape Colony, MacOwan s.n. Illustration by W.H. Fitch made
from cultivated material sent by MacOwan to W. Saunders (Fig. 24.3) (K! lecto. designated by Stedje 1987). Epitype (designated
by Martínez-Azorín et al. 2019):—Somerset East (3225): Somerset East District, near Charlton Falls, upriver on ledges on rock-
outcrops/low cliffs between homestead and falls, Boschberg, (–DA), elev. 1400 m, 11 December 2008, V.R. Clark, R.J. Daniels, M.
Fabricius & J. Le Roux 487 (NBG0267388! epi.).
Geschollia globuligera Mart.-Azorín, A.P.Dold & M.B.Crespo in Phytotaxa 427(2): 101 (2019) ≡ Drimia globuligera (Mart.-Azorín,
A.P.Dold & M.B.Crespo) J.C.Manning & Goldblatt in Bothalia 52(1): 3 (2022) (Figs 2.9, 25.3). Type:—SOUTH AFRICA. Eastern
Cape. Port Elizabeth (3325): Port Elizabeth, Thornhill, Van Stadens Wildflower Reserve, West side of reserve on flats, 300 m to
the east of river gorge, (–CC), elev. 230 m, coarse sandy soil on flats, recently burnt Algoa Sandstone Fynbos, 20 January 2018 (in
flower), A.P. Dold TD16029 (GRA! holo.; ABH! iso.).
Geschollia loedolffiae (van Jaarsv.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia loedolffiae van Jaarsv. in Aloe 43(2-
3): 50 (2006), basionym (Fig. 25.4). Type:—SOUTH AFRICA. Eastern Cape. Butterworth (3228): near the Kei River Mouth, (–CB),
6 March 2003, Van Jaarsveld & Voigt 17914 (NBG0207731! holo.; K000524679! iso.).
Geschollia longipedicellata Mart.-Azorín, Wetschnig, M.Pinter & M.B.Crespo in Phytotaxa 427(2): 103 (2019) ≡ Drimia longipedicellata
(Mart.-Azorín, Wetschnig, M.Pinter & M.B.Crespo) J.C.Manning & Goldblatt in Bothalia 52(1): 3 (2022) (Fig. 26.1). Type:—
SOUTH AFRICA. Eastern Cape. Willowmore (3323): Willowmore, ca. 1 km E of town, (–BC), elev. 860 m, 4 May 2015 in flower
ex hort in Graz Austria, W. Wetschnig & C. Huber WW4944 (GRA! holo.; ABH! iso.).
Geschollia occultans (G.Will.) Mart.-Azorín, M.B.Crespo & M.Pinter in Phytotaxa 427(2): 105 (2019) ≡ Drimia occultans G.Will. in
Cact. Succ. J. (Los Angeles) 83(6): 287 (2012), basionym (Figs 2.10, 26.2). Type:—NAMIBIA. Oranjemund (2816): Southern Namib
Desert, Swartkop Hill, 9 km E of Oranjemund, (–DA), March 2011 ex hort. in Cape Town, G. Williamson 5922 (NBG0271283!
holo.).
Geschollia prolifera Mart.-Azorín, A.P.Dold & M.B.Crespo in Phytotaxa 427(2): 107 (2019) ≡ Drimia prolifera (Mart.-Azorín, A.P.Dold
& M.B.Crespo) J.C.Manning & Goldblatt in Bothalia 52(1): 3 (2022) (Figs 2.11, 26.3). Type:—SOUTH AFRICA. Eastern Cape.
Fort Beaufort (3226): Fort Fordyce Reserve, Fort Beaufort, East ‘lip’ of Fuller’s Hoek forest basin, below EC Parks office and lodge,
(–DA), elev. 900 m, flowered ex hort 25 November 2017, A.P. Dold 16026 (GRA! holo.).
Geschollia zebrina Mart.-Azorín, A.P.Dold & M.B.Crespo in Phytotaxa 427(2): 109 (2019) ≡ Drimia zebrinella J.C.Manning & Goldblatt
in Bothalia 52(1): 3 (2022) nom. nov. [non Drimia zebrina (Mart.-Azorín, N.R.Crouch & M.B.Crespo) J.C.Manning & Goldblatt
in Bothalia 49(1): 5 (2019)] (Fig. 26.4). Type:—SOUTH AFRICA. Western Cape. Oudtshoorn (3322): Grootkop Nature Reserve,
NE of Oudtshoorn, (–CA), elev. 415 m, 4 May 2015 in flower ex hort in Graz, Austria, M. Martínez-Azorín, J. Vlok, A.P. Dold & A.
Martínez-Soler MMA893 (GRA! holo.; ABH! iso.).
9. Indurgia Speta in Stapfia 75: 169 (2001) (Figs 27–28). Typus generis:—I. indica (Roxb.) Speta (holotype).
≡ Drimia sect. Indurgia (Speta) J.C.Manning & Lekhak in S. African J. Bot. 123: 67 (2019). Typus sectionis:—I. indica (Roxb.) Speta ≡
Drimia indica (Roxb.) Kunth (holotype).
Description:—Bulbous geophyte. Bulb hypogeal ovoid to globose, 2–9 cm in diam., bulb scales usually compact,
white. Roots thickened and branched. Leaves 3–15 per bulb, 13–50 × 0.1‒5.5 cm, linear to lanceolate, canaliculated,
glabrous, synanthous or hysteranthous. Inflorescence a lax or dense raceme, usually elongated and multiflowered, 10‒
90 cm long, with 4‒50 flowers; peduncle 10‒160 cm long, terete, erect, smooth and glabrous; pedicels 0.4‒10 cm long,
from suberect to recurved and arquing downwards. Bracts deltoid, caducous, lowermost with short spur; bracteoles
absent. Flowers stellate, nocturnal and nodding or diurnal and erect-patent. Tepals 6, biseriate, 5‒18 mm long, oblong
to oblong-lanceolate, whitish, yellowish or brownish, with darker midrib, free or shortly fused, spreading to recurved.
Stamens 6, suberect to spreading, not connivent to style; filaments subulate-filiform, 3–15 mm long, smooth, adnate
to tepals at base; anthers dorsifixed, dehiscing by longitudinal slits along their whole length, yellowish. Ovary ovoid-
ellipsoid, 2.0–7.5 mm long, trigonous, pale green to yellowish, well differentiated from style. Style subclavate, usually
thickened, as long as or only slightly longer than the ovary. Stigma trigonous, truncate, papillose. Capsule ellipsoid-
oblong, 8‒26 mm long, valves apiculate, completely dehiscing from base, tepals cohering and inrolled above ovary
after anthesis, circumcissile from base and persisting as a cap at the top of the developing capsules. Seeds elliptic to
suborbicular, 4‒12 mm long, black, glossy, flattened with prominent central embryo and wide, flat wings.
FIGURE 27. Species of Indurgia Speta. 1. Indurgia indica (Roxb.) Speta from Wigth (1853: t. 2063, as Urginea indica (Roxb.) Kunth); 2.
Indurgia congesta (Wight) Speta (left side) and Indurgia coromandeliana (Roxb.) Speta (right side) from Wigth (1853: t. 2064, as Urginea
congesta Wight and Urginea coromandeliana (Roxb.) Hook.f. respectively).
Number of species and distribution:—Indurgia currently includes ten species, occurring from India and Sri
Lanka to Thailand (Fig. 8) (cf. Yadav et al. 2019), hence it is almost restricted to the Indian and Indochinese Regions
(sensu Takhtajan 1986), although some species also reach the eastern regions of the Omano-Sindian Subregion
(Martínez-Azorín et al. 2023a), shared with Vera-duthiea, which is present in Oman (Patzelt et al. 2021). For further
information on Indurgia species see Yadav et al. (2019) and Karuppusamy & Ravichandran (2021), among others.
Karyology:—2n=20 (Stedje & Nordal 1987, Yadav & Dixit 1990, as Urginea congesta Wight; Raghavan &
Venkatasubban 1940, Boraiah & Khaleel 1970, Naik 1976, Jha 1989, as Urginea indica Kunth; Kamble & Ansari
1976, Yadav & Dixit 1990, as Urginea polyantha Blatt.; Yadav & Dixit 1990, as Urginea razii (M.Y.Ansari) Deb &
Dasgupta; Boraiah & Khaleel 1970, as Urginea govindappae Boraiah & Fathima; Boraiah & Fathima 1972, as Urginea
govindappensis Boraiah & Fathima); 2n=20, 30 (Raghavan & Venkatasubban 1940, as Urginea polyphylla Hook.f.);

2n=20, 40, 60 (Jha & Sen 1983, as Urginea indica Kunth); 2n=40 (Naik 1973, Naik 1976, Dixit & Yadav 1989, as
Urginea coromandeliana Hook.f.).
History, diagnostic characters, and taxonomic relationships:—Roxburgh (1832) described Scilla
coromandeliana Roxburg (1832: 147) and S. indica Roxburg (1832: 147) from India. Further taxa were described
by Kunth (1843), Wight (1853) and Hooker (1892) from the same region and later placed in Urginea. Speta (1998b)
transferred the Indian species to Thuranthos and soon after Speta (2001) described Indurgia to include the Indian
members of Urgineoideae based on the phylogenetic analyses of Pfosser & Speta (2001). The revision of Urgineoideae
in Southern Africa by Manning & Goldblatt (2018) includes Indurgia and Vera-duthiea (genera originally described
to be restricted to southeastern Asia and to Central and northwestern Africa, respectively) in synonymy of their
Drimia sect. Thuranthos. They further included the Southern African species D. hesperantha (= Urginea revoluta),
D. vespertina Manning & Goldblatt (2018: 65), D. macrantha (Baker 1873b: 280) Baker (1892: 7), and D. basutica
(Phillips 1917: 306) Manning & Goldblatt (2018: 68), together with D. indica Roxburg (1832: 147) Jessop (1977: 272)
from India, so conceiving their section in a very wide sense. Manning & Goldblatt (2018) characterised the section by
the nocturnal flowers with reflexed tepals, but acknowledged that it was probably not monophyletic. This was already
shown by the phylogenetic works of Pfosser & Speta (2001, 2004), with Thuranthos, Indurgia, and Vera-duthiea (as
Duthiea Speta, nom. illeg.) representing independent and distant well-supported clades.
FIGURE 28. Indurgia polyantha (Blatt. & McCann) Speta (Photographs from upper row by S. Dutta).
Yadav et al. (2019) presented a detailed taxonomic revision of the Indian species of Drimia sensu lato accepting
nine species that were placed in their new D. sect. Indurgia (Speta 2001: 169) J.C.Manning & Lekhak in Yadav et al.
(2019: 67) to segregate the southeastern Asian members of Urgineoideae, and corroborated the exclusion of D. indica
from Southern Africa, earlier shown by Martínez-Azorín et al. (2018b, 2019a). The new section comprised taxa having
hysteranthous or synanthous leaves, lax or dense racemes, deltoid, caducous bracts with the lower ones short-spurred,
a lack of bracteoles, nocturnal and nodding flowers or sometimes diurnal and spreading, tepals almost free or shortly
connate basally, recurved or spreading, and filaments adnate to the base of the tepals. These taxa were divided into
two distinct pollination syndrome groups: (i) the Drimia indica group for D. coromandeliana (Roxburg 1832: 146)
Lekhak & P.B.Yadav in Yadav et al. (2016: 256), D. govindappae (Boraiah & Fathima 1972: 128) Lekhak & P.B.Yadav
in Bhat & Wani (2017: 145), D. indica, and D. nagarjunae (Hemadri & Swahari 1982: 105) Kumar (1984: 962), an
assemblage of night-flowering species with metareticulated pollen grains; and (ii) the D. wightii Lakshminarasimhan
(2003: 507) group for D. polyantha (Blatter & McCann 1928: 735) Stearn (1978: 208), D. raogibikei (Hemadri 2006a:
386) Hemadri (2006b: 224), D. razii Ansari (1981: 572), and D. wightii, to include the day-flowering species with
microreticulated pollen grains. Species in these two groups are isolated by their different flowering phenology, and
natural hybrids are unknown (Patil & Yadav 1992−1993).
Preliminary phylogenetic analyses of Indian members of Urgineoideae by Pfosser & Speta (2001, 2004) and
Pfosser et al. (2012) incorporate up to six samples of Indurgia and form a perfectly supported clade, which is included
in a collapsed clade that includes Ebertia, Vera-duthiea (as Duthiea, nom. illeg.), and Urginea s.str., as well supported
subclades. Saha & Jha (2019) extended the phylogenetic work in Indurgia, combining plastid and nuclear DNA
regions of twelve samples covering seven species from India, which formed a strongly supported clade. The studied
samples belong to both day- and night-blooming species with synanthous and hysteranthous leaves. The phylogenetic
results show two subclades based on the nuclear ITS region: subclade I comprising D. indica, D. polyantha, and D.
coromandeliana, and subclade II including D. nagarjunae, D. govindappae, D. wighttii, and D. razii. Both subclades
merge day- and night-flowering species as reported by Yadav et al. (2019), indicative that divergence in phenology
(and related pollination syndromes) has arisen more than once in Indurgia, so limiting the validity of taxonomic
inferences that can be drawn.
The phylogenetic analyses of Martínez-Azorín et al. (2023a) include six samples of Indurgia from India and
Thailand and agree with previous works showing a well supported clade to combine all studied samples of Urgineoideae
from that region. Therefore, we here accept Indurgia at genus rank based on a syndrome of morphological characters,
such as, linear-lanceolate, channelled leaves, deltoid, caducous and shortly spurred bracts, a lack of bracteoles, tepals
leathery and almost free to shortly connate, spreading to recurved, filaments adnate to the base of the tepals, erect to
suberect, straight, subulate filiform, anthers dorsifixed, styles erect, usually thickened, subclavate, truncate, stigma
trigonous, capsule with apiculate valves, and seeds ellipsoid with a prominent central embryo and flattened, winged
margins, arranged vertically in one or two rows per locule, decreasing in size from the distal to the proximal side.
Further, the restricted distribution of Indurgia to southwestern Asia and its isolated evolutionary status also supports
its acceptance at genus rank. Indurgia is sister to Urginea and Spirophyllos, which points to a close evolutionary
relationship accounted for by colonisation in the Northern Hemisphere following the arrival of an urgineoid migrant
from Southern Africa. Further phylogenetic studies, in combination with morphology, are necessary to evaluate
possible infrageneric classification.
Indurgia congesta (Wight) Speta in Stapfia 75: 170 (2001) ≡ Urginea congesta Wight in Icon. Pl. Ind. Orient. 6: 28, t. 2064 (left hand
figure) (1853), basionym ≡ Drimia congesta (Wight) Stearn in Ann. Mus. Goulandris 4: 208 (1978), nom. illeg., [non D. congesta
Bullock in Bull. Misc. Inform. Kew 1932(10): 504 (1932)] ≡ Drimia wightii Lakshmin. in Kew Bull. 58(2): 507 (2003) ≡ Thuranthos
congestus (Wight) Speta in Phyton (Horn, Austria) 38: 81 (1998) (Figs 2.12, 27.2). Type:—INDIA. Cadungalungoo, without date, R.
Wight s.n. (K000802725! lecto. designated by Yadav et al. in Phytotaxa 289: 258. 2016).
Indurgia coromandeliana (Roxb.) Speta ex Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Scilla coromandeliana Roxb., Fl.
Ind. 2: 147 (1832), basionym ≡ Urginea coromandeliana (Roxb.) Hook.f., Fl. Brit. India 6(18): 347 (1892), nom. illeg., [non U.
coromandeliana Wight in Icon. Pl. Ind. Orient. [Wight] 6: t. 2064 (1853)] ≡ Thuranthos coromandeliana (Roxb.) Speta in Phyton
(Horn, Austria) 38(1): 84 (1998) ≡ Drimia coromandeliana (Roxb.) Lekhak & P.B.Yadav in Phytotaxa 289(3): 256 (2016) ≡ Indurgia
coromandeliana (Roxb.) Speta in Stapfia 75: 170 (2001), nom. inval. (without basionym page) (Fig. 27.2). Type:—INDIA. Sand
hills on the coast of Coromandel, unpublished illustration for Roxburgh, Fl. Ind. 2, t.1821 (K lecto. designated by Yadav et al. in
Phytotaxa 289: 256. 2016).
Indurgia govindappae (A.Boraiah & Fathima) Speta in Stapfia 75: 170 (2001) ≡ Urginea govindappae A.Boraiah & Fathima in Bull. Bot.
Surv. India 12(1–4): 128 (1972), basionym ≡ Urginea govindappensis A.Boraiah & Fathima in Proc. Indian Sci. Congr. Assoc. 59(3):
359 (1972), nom. inval. [without Latin descr.] ≡ Thuranthos govindappae (A.Boraiah & Fathima) Speta in Phyton (Horn, Austria)
38(1): 84 (1998) ≡ Drimia govindappae (A.Boraiah & Fathima) Lekhak & P.B.Yadav in Bhat & Wani, Chromosome Structure and
Aberrations: 145 (2017). Type:—INDIA. Karnataka Lal Baugh [Karnataka Lalbagh Botanical Garden], Bangalore [Bengaluru], 17
May 1968, G. Boraiah & T. Fathima 601A (CAL holo.).
Indurgia indica (Roxb.) Speta in Stapfia 75: 170 (2001) ≡ Scilla indica Roxb., Fl. Ind. 2: 147 (1832), basionym ≡ Urginea indica (Roxb.)
Kunth, Enum. Pl. 4: 333 (1843) ≡ Drimia indica (Roxb.) Jessop in J. S. African Bot. 43(4): 272 (1977) ≡ Thuranthos indicus (Roxb.)
Speta in Phyton (Horn, Austria) 38: 84 (1998) (Figs 2.13, 27.1). Type:—INDIA. Coromandelia (K000802723! neotype designated as
“type” by Stedje in Nordic J. Bot. 7(6): 664. 1987). Note:—Some authors accept that a lectotype was designated by Deb & Dasgupta
(1974: 118) on the plate num. 1396 (Scilla indica) conserved at CAL (see Yadav et al. 2019: 72). However, Deb & Dasgupta
(1974) explicitly cited that they did not see the type material and simply added that the cited plate “fully agrees with Roxburgh’s
description”. We do not consider it as an effective lectotypification, and conversely accept (neo)typification by Stedje (1987) on the
material K000802723! (which she cited as “type”).

= Urginea wightiana Hook.f., Fl. Brit. India 6(18): 347 (1892) ≡ Thuranthos wightianus (Hook.f.) Speta in Phyton (Horn, Austria) 38:
84 (1998) ≡ Indurgia wightiana (Hook.f.) Speta in Stapfia 75: 170 (2001). Type:—INDIA. Coimbatore, March 1846, Wight s.n.
(K000802724! holo.).
Indurgia jeevae (Karupp. & V.Ravich.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia jeevae Karupp. & V.Ravich. in
Gardens’ Bulletin Singapore 73(1): 210 (2021), basionym. Type:—INDIA. Tamil Nadu, Kanyakumari District, Alamparai Hills,
elev. 600 m, 9 March 2012, S. Karuppusamy & V. Ravichandran 874 (MH holo.; Sri Ganesan Herbarium iso.).
Indurgia nagarjunae (Hemadri & Swahari) Speta in Stapfia 75: 170 (2001) ≡ Urginea nagarjunae Hemadri & Swahari in Ancient
Sci. Life 2: 105 (1982), basionym ≡ Drimia nagarjunae (Hemadri & Swahari) An.Kumar in J. Econ. Tax. Bot. 5(4): 962 (1984)
≡ Thuranthos nagarjunae (Hemadri & Swahari) Speta in Phyton (Horn, Austria) 38: 84 (1998). Type:—INDIA. Andhra Pradesh,
Krishna district, Vijayawada, 5 May 1980, Hemadri & Swahari 3001A (CAL holo.; AYUSH, VIJAYAWADA, BSI, K000802727!,
BLAT, MH iso.).
Indurgia polyantha (Blatt. & McCann) Speta in Stapfia 75: 170 (2001) ≡ Urginea polyantha Blatt. & McCann in J. Bomb. Nat. Hist. Soc.
32: 735 (1928), basionym ≡ Drimia polyantha (Blatt. & McCann) Stearn in Ann. Mus. Goulandris 4: 208 (1978) ≡ Drimia polyantha
(Blatt. & McCann) Ansari & Sundararagh. in J. Bombay Nat. Hist. Soc. 77(1): 174 (1980), nom. superfl. ≡ Thuranthos polyanthus
(Blatt. & McCann) Speta in Phyton (Horn, Austria) 38: 84 (1998) (Fig. 28). Type:—INDIA. Maharashtra, Panchgani, March–April
1926, Blatter & McCann 101a (BLAT lecto. designated as “type” by Deb & Dasgupta 1974: 122).
Indurgia raogibikei (Hemadri) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397: 294 (2019) ≡ Urginea raogibikei
Hemadri in Med. Fl. of Pragati Resorts.: 386 (2006a), basionym ≡ Drimia raogibikei (Hemadri) Hemadri in A Treatise on Tribal
Medicine: 224 (2006b). Type:—INDIA. Andhra Pradesh, Nellore district, Bhata–Udayagiri, 6 June 2006, Hemadri 2A (CAL holo.;
BSI, K iso.).
Indurgia razii (Ansari) Speta in Stapfia 75: 170 (2001) ≡ Drimia razii Ansari in J. Bombay Nat. Hist. Soc. 78(3): 572 (1981), basionym ≡
Urginea razii (Ansari) Deb & Dasgupta in J. Bombay Nat. Hist. Soc. 84(2): 412 (1988) (as “rajii”) ≡ Thuranthos razii (Ansari) Speta
in Phyton (Horn, Austria) 38: 84 (1998). Type:—INDIA. Maharashtra, Pune district, Dive Ghat, near Zendewadi, 19 April 2015, P.B.
Yadav & M.M. Lekhak 18 (CAL neo. designated by Yadav et al. in Phytotaxa 289: 256. 2016).
Indurgia rupicola (Trimen) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea congesta Wight var. rupicola Trimen in J.
Bot. 27: 167 (1889), basionym ≡ Drimia rupicola (Trimen) Dassan. in M.D. Dassanayake & W.D. Clayton (eds), Revised Handb. Fl.
Ceylon 14: 220 (2000). Type:—SRI LANKA. Chinks of arid flat rocks at Dambulla, on the lower part of the ascent to the Temples,
July 1887, Trimen s.n. (K000802726! holo.).
10. Iosanthus Mart.-Azorín, M.B.Crespo, M.Pinter, Slade & Wetschnig in Pl. Biosystems 153(4): 584 (2019; on line
18 December 2018) (Figs 30–32). Typus generis:—I. toxicarius (C.Archer & R.H.Archer) Mart.-Azorín, M.B.Crespo,
M.Pinter, Slade & Wetschnig (holotype).
= Drimia sect. Khubusia J.C.Manning & Goldblatt in Strelitzia 40: 120 (2018). Typus sectionis:—D. khubusensis P.C.van Wyk &
J.C.Manning (holotype).
Description:—Small to medium deciduous bulbous geophyte, to 50 cm high. Bulb hypogeal, usually solitary or
growing in small groups, ovoid to subglobose, 1–3 cm in diam., extended into a 2–5 cm long hypogeal neck, inner
scales compact, white, fleshy, outer tunics pale brown-grey, membranous. Roots thickened and branched. Leaves
hysteranthous or sometimes synanthous, suberect, spreading and somewhat curved or twisted, 3–16 per bulb, narrowly
linear, filiform, 5–18 cm long, subterete to canaliculate, green, sometimes maculate on lower portions, smooth,
sometimes surrounded by papery cataphylls. Inflorescence racemose, dense, subcapitate-ovoid or lax and elongated,
0.5–16.0 cm long, with 4–20 flowers; peduncle erect, 1–20 cm long, glabrous; pedicels 1.5–25.0 mm long at anthesis,
from suberect to patent and arching downwards; bracts small, 1–2 mm long, membranous, triangular, with slight
curve outgrowth or short spur to 1 mm long. Bracteoles absent. Flowers stellate or cylindrical-campanulate, patent
to nodding, with suberect to patent-reflexed tepals, diurnal or vespertine-nocturnal. Tepals 6, biseriate, 4–15 mm
long, ovate to narrowly oblong and elongated, free or only shortly connate at base, whitish or carneous with central
longitudinal pinkish-brown or reddish longitudinal band, straight and spreading or connivent and erect along basal
half and spreading patent along apical half. Stamens 6; filaments filiform and subterete or triangular-lanceolate and
flat, 1.5–6.0 mm long, adnate to perigone segments at base for 0.5–6.0 mm, erect and connivent to gynoecium or only
spreading distally; anthers yellow, ovate-oblong, 1.2–3.0 mm long, basifixed or medifixed, dehiscing longitudinally
along their whole length, with yellow pollen. Ovary green, ovoid to conical, trigonous, 2–4 mm long. Style white,
erect, 1–7 mm long. Stigma small or distinctly capitate and 3-lobed puberulous. Capsule ovoid to subglobose, 6–15
mm long, erect, dark or pale coloured before dehiscence, with remains of perigone circumscissile below and forming
an apical cap, valves splitting to base and usually widely spreading to reflexed. Seeds elliptic to discoid, flattened,
broadly winged, with emarginate hilum, 4–10 mm long, testa black to cream-coloured to pale greyish, with sinuous
anticlinal cell walls.
Number of species and distribution:—Iosanthus includes 4 species confined to the western regions of Southern
Africa, mostly in Namibia and northern South Africa (Fig. 29), hence restricted to the Karoo-Namib Region (sensu
Takhtajan 1986). For further information on Iosanthus species see Archer & Archer (1999), Manning & Goldblatt
(2018), and Martínez-Azorín et al. (2019c).
History, diagnostic characters, and taxonomic relationships:—Ornithogalum toxicarium Archer & Archer
(1999: 431) was described to accommodate a toxic plant causing severe stock losses in South Africa and Namibia,
even though it was earlier determined in scheda as an undescribed species of Urginea. Martínez-Azorín et al. (2019c)
presented evidence that O. toxicarium belongs to Urgineoideae and described the genus Iosanthus, as a new and
independent phylogenetic lineage within the Urgineoideae (Martínez-Azorín et al. 2023a). This new monotypic genus
was characterised by the small plant size, filiform leaves, very short peduncle with few-flowers and short raceme, lack
of bracteoles, a conical ovary with short tapering style, capsule capped with the withered perigone, and the flattened,
ovate, winged seeds.
Manning & Goldblatt (2018) described Drimia khubusensis P.C.van Wyk & J.C.Manning in Manning & Goldbatt
(2018: 120) from northwestern South Africa, near the border with Namibia. The species approaches I. toxicarius in
several respects, but differs by its elongated and reflexed tepals, and longer and filiform filaments and style. Drimia
khubusensis seems to be unique in Urgineoideae in having dark coloured capsules (sometimes blackish), with the
valves reflexed from the base at dehiscence to completely expose the creamy-white or pale greyish, flattened, discoid
and winged seeds (Manning & Goldblatt 2018).
The phylogenetic results of Martínez-Azorín et al. (2023a) revealed a perfectly supported clade (Bayesian analyses)
corresponding to Iosanthus, including two samples of D. khubusensis sister to a sample of I. toxicarius. Accordingly,
the sister is a sample of an undescribed species from Central Namibia, described here as Iosanthus macrostigma Mart.-
Azorín et al.; all relationships have strong support. The Iosanthus clade is sister to Indurgia in the combined plastid and
nuclear analyses although in the plastidial analyses the former is included in a polytomy related to Indurgia, Urginea,
Ebertia, and Vera-duthiea, among others. However, the ML analyses show the undescribed Iosanthus species from
Namibia in a polytomy within a wide clade with other related genera like Urginea, Indurgia, and Vera-duthiea among
others, although the remaining two described species are placed together in a strongly supported clade. These three
Iosanthus species characteristically share a relatively small plant size, hypogeal, compact bulbs, filiform leaves, lack of
bracteoles, short and few-flowered inflorescence, and free tepals. We have observed that I. toxicarius also shows dark
coloured capsules before their ripening, with valves splitting widely to expose seeds of similar morphology to those
of D. khubusensis. These propagules have a prominent endosperm and pale marginal regions in the testa. Further, I.
macrostigma from Central Namibia approaches D. khubusensis in leaf, flower, capsule, and seed morphology, although
differing in its adnate filaments, which arise from the basal third of the tepals, and in its strongly thickened, capitate,
3-lobed and papillose stigma, among other characters.
Recent phylogenetic and morphological studies on Urginea amboensis from the farm Vergenoeg in northern
Namibia collected by A. Eichhoff support its close relationships to I. macrostigma. We, therefore, propose accepting
Iosanthus to include the latter four species, which are geographically related and characterised by: small to medium
overall plant size; compact, hypogeal bulb with elongated neck; filiform leaves, commonly twisted; short to
elongated inflorescence; lack of bracteoles; diurnal or vespertine-nocturnal flowers; capsules with the withered tepals
circumscissile below and forming an apical cap, with valves splitting to the base and reflexed; and flattened, ellipsoid,
winged seeds with emarginate hilum. Further phylogenetic studies are required to evaluate possible alternatives based
on the morphological variation and phylogenetic divergence in the group.
Iosanthus amboensis (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea amboensis Baker in Bull. Herb. Boissier
ser. 2, 3(8): 665 (1903), basionym ≡ Vera-duthiea amboensis (Baker) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa
397(4): 296 (2019) (Figs 2.14, 30.1). Type:—NAMIBIA. Ondangua (1715): Amboland, Ondonga, (–DD), March 1887, Rautanen
s.n. (Z000087308! lecto. designated here; Z0001023251! isolecto.).
Comments:—Urginea amboensis Baker (1903: 665) was described from northern Nambia (“Amboland, Ondonga”) and was only known
from the type collection by M. Rautanen in March 1887 (Z000087308! and Z0001023251!), which includes inflorescences with
closed flowers [resembling those of Dipcadi Medik. (1790: 431), as seen in posterior identifications] and bulbs, lacking leaves. The
protologue indicates that stamens were distinctly shorter than tepals. No further material fitting this species has been found in the
studied herbaria, hence precluding its taxonomic placement in the past. Recent taxonomic works placed it as synonym of Drimia
indica (Manning & Glodblatt 2018) or as a species of Vera-duthiea (Martínez-Azorín et al. 2019a), following the comments by Baker
(1903) on its similarity to ‘U. indica Kunth’.
The study of fresh material recently collected by A. Eichhoff from the farm Vergenoeg in Namibia matches Urginea
amboensis Baker (1903: 665) in morphology and distribution. These plants show hypogeal bulbs with compact scales
and elongated hypogeal neck; leaves filiform, canaliculate, twisted, green with dark maculae at base; lax, elongated
racemes with nodding, nocturnal flowers; tepals free and connivent along the lower half and patent above, filaments
adnate to tepals and arising around the middle, erect and connivent to style; bracts minute, deltoid; capsules ovoid
elongated and seeds ellipsoid, widely winged with prominent embryo and emarginate hilum (Figs 2.14, 30.1). These
characters approach I. macrostigma, a conclusion supported by our recent unpublished phylogenetic data, in which two
samples of U. amboensis from the farm Vergenoeg are placed sister to I. macrostigma with strong support. Samples of
the latter two species are sister to a clade combining I. toxicarius and I. khubusense, with the latter relationship being
weakly supported in combination in some analyses. Further studies are required to improve our understanding of the
group and to evaluate other taxonomic alternatives within the genus.
FIGURE 30. Species of Iosanthus Mart.-Azorín et al. displayed in horizontal rows of images. 1. Iosanthus amboensis (Baker) Mart.-
Azorín et al. (top 3 rows of images) (Photographs by A. Eichhoff); 2. Iosanthus macrostigma Mart.-Azorín et al.

Iosanthus khubusense (P.C.van Wyk & J.C.Manning) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia khubusensis
P.C.van Wyk & J.C.Manning in Strelitzia 40: 120 (2018), basionym. Type:—SOUTH AFRICA. Northern Cape. Oranjemund (2816):
Richtersveld, Khubus [Kuboes], 15 m directly west of first and oldest graveyard, (–BD), 6 October 2014, P.C.V. van Wyk 500 (NBG
holo.).
Comments on distribution:—This species was originally described from a single locality in Kuboes (Northern Cape Province of South
Africa). The study of the herbarium collection Giess 8270 (WIND!) fits the overall morphology of this species, representing its
second locality in the Rössingberg, ca. 35 km east of Swakopmund, an extension of ca. 450 km to the north and the first record of the
species in Namibia. However, fresh material of the Namibian population is required for final taxonomic placement.
Regarding the collection H. Dauth 160 (WIND!) from near Rosh Pinah in southern Namibia, it also resembles I. khubusense, but clearly
differs by the stout peduncle, shorter pedicels (2–3 mm long), and tepals spreading-reflexed from the base, among other characters.
It most likely represents an undescribed species in the genus, though further material is needed for its formal description.
Iosanthus macrostigma Mart.-Azorín, M.Pinter, M.B.Crespo & M.Á.Alonso sp. nov. (Figs 30.2, 31).
Type:—NAMIBIA. Windhoek, Klein Windhoek, elev. ca. 1600 m, ex hort 08 May 2015, I. Pehlemann s.n. (WIND holo.; ABH iso.).
Diagnosis:—Planta speciosa ceteris speciebus Iosanthi valde affinis, sed eis distinctissima filamentis staminum ad tertium inferiorem
tepalorum adnatis, et stigmatibus notabilis valde incrasato-capitatis, trilobatis et glandulosis.
Description:—Bulbous geophyte to 4 cm high. Bulb hypogeal, usually solitary or growing in small groups, ovoid to subglobose and
depressed in old plants, 1.5–3.0 × 1.0–2.5 cm, extended into a 1–2 cm long hypogeal neck; outer tunics pale brownish, membranous.
Roots thickened and branched. Leaves 4–11 per bulb, narrowly linear, suberect, canaliculate, somewhat twisted, 2–10 × 0.10–0.18
cm, glabrous, green, with distinct dark maculae at base, withered at flowering time. Inflorescence a short raceme with 3–5 flowers,
opening one by one in consecutive days; peduncle erect, pale grey, 15–20 mm long, with small protuberances near raceme, specially
in fruit; pedicels 4–5 mm long at anthesis, slightly elongating to 7 mm in fruit, with small protuberances near flower; bracts small,
deltoid, 1.0–1.2 mm long, green with translucent membranous margins, with short spur ca. 0.6 mm long. Bracteoles absent. Flowers
stellate, patent to suberect, vespertine-nocturnal. Tepals 6, biseriate, free or only shortly connate at base, narrowly lanceolate-oblong,
with somewhat cucullate apex, whitish to carneous colour with central longitudinal brownish-green longitudinal stripe evident on
both sides, connivent along basal half to form slightly urceolate tube and spreading to patent above; outer tepals oblong and widened
in apical portion, 8–9 × ca. 2 mm; inner tepals narrowly obovate, tapering to base, 8–9 × ca. 1.5 mm. Stamens 6; filaments white,
subterete, adnate to tepals for ca. 2 mm, free portion ca. 5 × 0.4 mm, connivent to style and slightly spreading above perigone; anthers
yellow, ovate, ca. 1 mm long after dehiscence, dehiscing longitudinally along their whole length, with yellow pollen. Ovary green,
ovoid, 3-locular, ca. 3 × 1.8 mm, attenuate to style. Style white, elongated, erect, ca. 5 mm long, slightly thickened and clavate.
Stigma ca. 1 × 2 mm, distinctly thickened, dome-like, trilobate, with widened lobes reflexed and papillose. Capsule trilocular,
loculicidal, ovoid to subglobose, 7–9 × 7–9 mm, suberect, with remains of perigone circumscissile below and forming an apical cap,
valves ovate, apiculate, splitting to base and widely spreading to expose seeds. Seeds black, heart shapped, 8–9 × 6–7 mm, flattened
with prominent embryo, broadly winged, emarginate hilum, with sinuous anticlinal testa cell walls.
Etymology:—Named after the distinctly thickened, dome-like, trilobate stigma.
Phenology:—Iosanthus macrostigma flowers from March to May in cultivation in Alicante (Spain) and fruit and seeds are set from May
to June. Flowers open in the evening at about 18h00 and last for a night, a behaviour also shared with the sister species I. amboensis.
Further studies are necessary to elucidate the biology of this remarkable species in the wild.
Habitat and distribution:—The species is only known from the surrounds of Klein Windhoek, Windhoek, Namibia (I. Brase, formerly
I. Pehlemann, pers. comm.) where it occurs in highland shrubland. The locality is about 1600 m elevation, supports a mean annual
rainfall of 530 mm and a mean annual temperature of 18.5ºC. We were not able to confirm the presence of this species in that region,
and the material and information used for the description of the species was provided without detailed locality details. Further studies
are needed to confirm the distribution and habitat of this species.
Diagnostic characters and taxonomic relationships:—Iosanthus macrostigma is characterised by the solid hypogeal bulb; the 4–11
proteranthous, filiform, canaliculate, glabrous, leaves, which are maculate at the base; the lax, pauciflorous raceme; the vespertine-
nocturnal flowers with free tepals, which are connivent along the basal half and spreading apically, and the adnate filaments; the erect,
elongate, subclavate style supporting a distinctly thickened, dome-like, trilobate, papillose stigma, the capsules with the withered
tepals atop and widely spreading valves; and the heart-shaped, flattened, winged seeds with prominent embryo (Figs 2.15, 30.2, 31).
The new species is well characterized in having a very swollen stigma, reminiscent of some Thuranthos species. It shares the typical
capsules and seeds of I. toxicarius and I. khubusensis, a link supported by phylogenetic studies (Martínez-Azorín et al. 2023a),
despite the differences in flower morphology, which could be linked to different pollination syndromes. Our recent unpublished
phylogenetic work places this species as sister to I. amboense, with strong support—a result supported by their shared vespertine-
nocturnal flowers, filaments long adnate to tepals, and the maculate, twisted leaves.
FIGURE 31. Iosanthus macrostigma Mart.-Azorín et al. from Klein Windhoek, Namibia (Type locality). a. Portion of bulb and
inflorescence; b. Inflorescence; c. Flower in frontal view; d. Dissected tepals with stamens; e. Tepals with adnate stamens; f. Gynoecia; g.
Dissected flower in lateral view; h. Developing capsules; j. Capsules and seeds; k. Seed; m. Leaves. Scales 5 mm.

Iosanthus toxicarius (C.Archer & R.H.Archer) Mart.-Azorín, M.B.Crespo, M.Pinter, Slade & Wetschnig in Pl. Biosyst. 153(4): 586
(2019) ≡ Ornithogalum toxicarium C.Archer & R.H.Archer in S. African J. Bot. 65(5–6): 431 (1999), basionym ≡ Drimia toxicaria
(C.Archer & R.H.Archer) J.C.Manning & Goldblatt in Bothalia 49(1): 3 (2019) (Figs 2.16, 32). Type:—SOUTH AFRICA. Western
Cape. Beaufort West (3222): farm Ryst Kuil 351, in vicinity of old uranium mine, (–DB), 08 October 1983, Retief & C. Reid 239
(PRE0656451! lecto. designated by Martínez-Azorín et al. 2019c: only the bulb with withered leaves and two inflorescences placed
on the lower part of the sheet). Note:—The lectotype of this name was first designated in a paper effectively published online 18
December 2018 by Martínez-Azorín et al. (2019c). Subsequent superfluous lectotypification published on 10 April 2019 by Manning
(2019) is fully coincident with our previous type selection.
FIGURE 32. Iosanthus toxicarius (C.Archer & R.H.Archer) Mart.-Azorín et al.
11. Ledurgia Speta in Stapfia 75: 168 (2001). Typus generis:—L. guineensis Speta (holotype).
Description:—Bulbous geophyte. Bulb ovoid-oblong to subglobose, outer scales membranous and inner bulb scales
white and fleshy. Roots thickened and branched. Leaves unknown. Inflorescence an erect, raceme with 6‒12 flowers;
peduncle very short, erect; pedicels spreading, straight, 3–4 mm long, glabrous. Bracts lanceolate, ca. 3 mm long with
distinct spur. Flowers campanulate with reflexed tepals, subpatent. Tepals 6, biseriate, ca. 7–8 mm long, lanceolate
to oblong, whitish with brown longitudinal central band, shortly connate at base. Stamens 6, suberect to slightly
spreading; filaments filiform, white, shortly adnate to base of tepals; anthers oblong, opening by longitudinal slits.
Ovary ovoid, ca. 3–4 mm long, greenish, with ca. 14 ovules per locule. Style ca. 2.5 mm long, erect, white, filiform,
stigma slightly trigonous. Capsule and seeds not studied.
Number of species and distribution:—Ledurgia includes a single species, occurring in Guinea, hence restricted
to Guineo-Congolian Region (sensu Takhtajan 1986) (Fig. 29).
History, diagnostic characters, and taxonomic relationships:—Speta (2001) described Ledurgia guineensis
Speta as a monotypic genus from Guinea based on plants with imbricate bulb scales, very short peduncle and few
flowered raceme; straight and patent pedicels; and campanulate flowers with slightly connate tepals (Speta 2001). Our
phylogenetic results agree with previous conclusions of Pfosser & Speta (2001) in which the samples of this genus
represent an independent lineage here accepted at genus rank. This lineage is related to Zingela and Thuranthos, but
shows quite a different morphology. We were not able to study living material of this genus. Further studies are needed
to improve its morphological characterisation.
Ledurgia guineensis Speta in Stapfia 75: 168 (2001) ≡ Drimia guineensis (Speta) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3):
556 (2004). Type:—GUINEA. Between Pita and Hafia, 21 May 1987, I. Bert. cult. HBV Li 2582 (Speta in LI, reportedly transferred
to W).
12. Litanthus Harv. in London J. Bot. 3: 314, t. 9 (1844) (Fig. 33). Typus generis:—L. pusillus Harv. (holotype).
≡ Drimia sect. Litanthus (Harv.) J.C.Manning & Goldblatt in Strelitzia 40: 150 (2018). Typus sectionis:—L. pusillus Harv. (holotype).
FIGURE 33. Species of Litanthus Harv. displayed in horizontal rows of images. 1. Litanthus pusillus Harv.; 2. Litanthus stenocarpus
(J.C.Manning & J.M.J.Deacon) Mart.-Azorín et al.
Description:—Dwarf bulbous geophyte. Bulb hypogeal or slightly epigeal, sometimes proliferous and clump forming,
ovoid to subglobose, up to 1 cm in diam., outer scales scarious and inner scales white to pinkish, fleshy. Roots thickened
and branched. Leaves 1‒5, filiform, subterete, 1‒10 cm long, 0.2‒1.0 mm wide, suberect, green, glabrous, hysteranthous
or synanthous. Inflorescence formed by 1(‒2) teminal flowers; peduncle erect, terete, minutely papillate, 1–8 cm long,
greenish to purplish; pedicels 1–3 mm long at anthesis, recurved, erect in fruit to 8 mm long. Bracts lanceolate, usually
two, terminal, subopposite, commonly only one with flower, ca. 1 mm long, with spur ca. 0.5 mm long; bracteoles
absent. Flowers tubular, nodding. Tepals 6, 4‒7 mm long, whitish with pinkish, brownish or purple tinge, fused for
most of their length to form cylindrical tube ending in short apical erect or spreading lobes, with distinct greenish
or brownish longitudinal band. Stamens 6, included in perigone tube; filaments adnate to tepals and arising around
middle of perigone, free portions very short, ca. 0.5 mm long, flattened, white, smooth; anthers oblong, 1‒2 mm long,
erect, opening by longitudinal slits, cordate at base, connective extended at apex into membranous flap. Ovary ovate-
oblong, 1.5‒3.0 mm long, greenish to yellowish. Style erect, columnar, 2.0‒3.5 mm long, white, with subcapitate
stigma ending in 6 erect lobes. Capsule subglobose, ovate or oblong, trigonous with blunt edges in section, 3‒6 mm
long, from uniformly pale brown to longitudinally banded brown and white. Seeds trigonous, narrowly pyramidal,
pointed, 0.5–1.5 mm long, embryo prominent with very narrow winged margins, black, glossy, with rugose testa and
subisodiametric and polygonal testa cells.
Number of species and distribution:—Litanthus includes two species occurring in Southern Africa (Manning et
al. 2013) (Fig. 29), restricted to the Cape, Karoo-Namib, Uzambara-Zululand Region, and the Southern Section of the
Zambezian Subregion (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a). For further information on Litanthus
species see Harvey (1844) and Manning et al. (2013).
Karyology:—2n=20 (Bruyns & Vosa 1987, as Litanthus pusillus Harv.).
History, diagnostic characters, and taxonomic relationships:—Litanthus was described by Harvey (1844)
as monotypic to include L. pusillus Harvey (1844: 315), a distinct dwarf species with 1(–2)-flowered inflorescence,
two subopposite spurred bracts, nodding, tubular flowers with tepals connate into a long tube, and included stamens
with adnate very short filaments. This genus has been accepted by most researchers working in Urgineoideae (Baker
1871, Hooker 1872, Jessop 1977, Speta 1998a, 1998b, Martínez-Azorín et al. 2015). However, Goldblatt & Manning
(2000), Manning et al. (2004), and Manning & Goldblatt (2018) included Litanthus in Drimia sensu lato. Manning
et al. (2013) presented a revision of Litanthus (as a group of Drimia) and described a second related species. They
added useful diagnostic characters for the genus, such as the usually solitary flower with a second empty bract and the

elongation of the anther connective into a small, translucent, membranous flap. Further, they described and illustrated
the characteristic stigma of both species, unique among Urgineoideae in possessing six tiny, erect teeth. The latest
revision of Urgineoideae in Southern Africa (Manning & Goldblatt 2018) placed L. pusillus and L. stenocarpus
(J.C.Manning & J.M.J.Deacon in Manning et al. 2013: 99) Mart.-Azorín et al. (2015: 168) in Drimia sect. Litanthus
(Harvey 1844: 314) Manning & Goldblatt (2018: 150).
The phylogenetic studies by Manning et al. (2004) show a sample of Litanthus pusillus as an independent lineage
within a large collapsed clade. Pfosser & Speta (2001, 2004) recovered a sample of L. pusillus and Schizobasis as sister
groups. Pfosser et al. (2012) found the same relationship following the inclusion of three samples in each genus. The
phylogenetic analyses of Martínez-Azorín et al. (2023a) considered four samples of Litanthus, which form a strongly
supported clade sister to the monophyletic Schizobasis, for which eleven samples were included in the analyses. The
sister relationship between Litanthus and Schizobasis, at first sight surprising when based on their different flower
and inflorescence morphology, is supported by the apiculate connective or the angled seeds, as shown by Manning
& Goldblatt (2018). However, merging these genera in a single genus would be highly disruptive due to their distinct
differences in flower and inflorescence morphology. Therefore, we accept Litanthus based on its unique syndrome of
morphological characters: very small plant size; usually solitary, nodding flowers; tepals connate for at least half of
their length in a cylindrical tube; stamens included with adnate filaments arising from the upper half of perigone, and
apiculate connetive; stigma with 6 minute, erect teeth; seeds angular, polygonal, narrowly pyramidal, ca. 1 mm long;
and corroborated with biogeography and phylogenetic relationships.
Litanthus pusillus Harv. in London J. Bot. 3: 315, t. 9 (1844) ≡ Drimia uniflora J.C.Manning & Goldblatt in Strelitzia 9: 712 (2000), nom.
nov. [non Drimia pusilla Jacq. ex Willd. (1799)] (Figs 2.17, 33.1). Type:—SOUTH AFRICA. [Eastern Cape], ‘shady places in the
woods by the Zwartkop’s River, Uitenhage, Cape of Good Hope’, [December 1829], Zeyher s.n. (TCD lecto. designated by Jessop
in J. S. African Bot. 43: 308. 1977; BOL, SAM!, S barcode S-G-7913! isolecto.).
Litanthus stenocarpus (J.C.Manning & J.M.J.Deacon) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 201(2): 168
(2015) ≡ Drimia stenocarpa J.C.Manning & J.M.J.Deacon in S. African J. Bot. 90: 99 (2013), basionym (Figs 2.18, 33.2). Type:—
SOUTH AFRICA. Western Cape. Vanrhynsdorp (3118): Papendorp, 25 miles W of Vredendal, (–CA), 17 March 1971, H. Hall 3921
(NBG92544! holo.; PRE! iso.).
13. Mucinaea M.Pinter, Mart.-Azorín, U.Müll.-Doblies, D.Müll.-Doblies, Pfosser & Wetschnig in Phyton (Horn,
Austria) 53(2): 296 (2013) (Figs 2.19, 34). Typus generis:—M. nana (Snijman) M.Pinter, Mart.-Azorín, U.Müll.-
Doblies, D.Müll.-Doblies, Pfosser & Wetschnig (holotype).
= Drimia sect. Orchidiformes J.C.Manning & Goldblatt in Strelitzia 40: 118 (2018). Typus sectionis:—D. nana Snijman (holotype).
Description:—Bulbous geophyte. Bulb subglobose, hypogeal, sometimes proliferous and clump forming by bulblets,
inner leaf bases of each shoot generation narrow, semiterete, with purplish bulb flesh; sheathing cataphyll forming
short neck, 12–50 mm long; the outer leaf bases (±10) form second gamophyllous sheath. Leaves numerous, 14‒20
per shoot, suberect to spreading, filiform, 10–25 cm long, ca. 1 mm wide, glabrous, hysteranthous, or sometimes
synanthous in cultivation. Inflorescence a lax raceme, 3‒11 cm long, with 6‒27 flowers; peduncle erect, purplish, 7‒14
cm long, glabrous; pedicels spreading, straight, 7‒14 mm long, purplish. Bracts ovate–triangular, acute, with short
spur; bracteoles absent. Flowers stellate, erect-patent, diurnal. Tepals 6, biseriate, 7–11 mm long, free, usually strongly
reflexed, purplish–pink, with green markings surrounded by white ring at base on adaxial side. Stamens 6, erect and
connivent to style; filaments stout, flattened, 3.5–4.0 mm long, rather suddenly contracted to terete upper part, white,
sometimes with purplish transverse band in middle of flattened part, all 6 deflexed downwards above ovary, convergent
to fasciculate towards anthers; anthers erect, ca. 2 mm long, forming fascicle, yellow, basifixed, with longitudinal
dehiscence but only opening as an apical slit. Ovary ovoid, pale–green, 1.5–2.0 mm long. Style white, declinate, 3–4
mm long, protruding laterally from fascicle of filaments below anthers (enantiostyly). Capsule ovoid, 4–7 mm long,
valves completely dehiscing from base; tepals cohering and inrolled above ovary after anthesis, circumcissile from
base and persisting as a cap at the top of the developing capsules. Seeds subellipsoid to subfusiform, 2.0–2.8 mm long,
dark golden-brown, glossy, testa surface distinctly reticulate-alveolate with subisodiametric, polygonal cells, with
collapsed periclinal walls and prominent anticlinal walls.
FIGURE 34. Mucinaea nana (Snijman) M.Pinter et al.
Number of species and distribution:—Mucinaea is a monotypic genus endemic to Namaqualand (northwestern

South Africa), occurring in the Kamiesberg Mountains and Kourkammaberg, extending north to near Eksteenfontein
(Grenier 2019) (Fig. 29). It is restricted to the Karoo-Namib Region (sensu Takhtajan 1986 and Martínez-Azorín et al.
2023a). For further information on Mucinaea see Snijman (1985) and Pinter et al. (2013).
History, diagnostic characters, and taxonomic relationships:—Snijman (1985) described Tenicroa nana,
differing from typical Tenicroa species by the usually hysteranthous leaves, the bright purplish-pink tepals (a colour
nearly unique within Urgineoideae), each showing a basal green marking encircled by a white ring (also unique
in Urgineoideae), anthers with longitudinal dehiscence but only opening as an apical pore or slit (suggesting buzz-
pollination), and the purple amplexicaul sheathing cataphyll without raised transverse ribs, enclosing the leaf bases.
The bulb structure is also unique in Urgineoideae, presenting a gamophyllous second sheath inside the amplexicaul
cataphyll consisting of the bases of about 10 non-amplexicaul foliage leaves (Pinter et al. 2013).
Manning et al. (2004), in a preliminary phylogenetic study, showed T. nana as basal to all studied samples of
Urgineoideae when Bowiea is excluded. Tenicroa nana is placed far from the clade including typical Tenicroa species.
In an expanded phylogeny, Pfosser et al. (2012) also showed Tenicroa nana in an independent lineage within a large
clade with several well-supported subclades, among them the remaining Tenicroa species. The phylogenetic analyses
of Martínez-Azorín et al. (2023a) place a sample of Mucinaea nana in an isolated position, being basal to all other
Urgineoideae taxa excluding Rhadamanthus and Bowiea.
Based on the distinct morphological and molecular differences, Pinter et al. (2013) described Mucinaea to include
T. nana, accepted here as a monotypic genus.
Mucinaea nana (Snijman) M.Pinter, Mart.-Azorín, U.Müll.-Doblies, D.Müll.-Doblies, Pfosser & Wetschnig in Phyton (Horn, Austria)
53(2): 296 (2013) ≡ Tenicroa nana Snijman in S. African J. Bot. 51(4): 284 (1985), basionym ≡ Drimia nana (Snijman) J.C.Manning
& Goldblatt in Bothalia 33(1): 111 (2003), non Drimia nana (Oyewole) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557
(2004), nom. illeg. (Fig. 34). Type:—SOUTH AFRICA. Northern Cape. (3018): Kamiesberg, on slopes of the Rooiberg, (–AC), elev.
1300 m, Fl. ex Kirstenbosch 26 November 1980, D.A. Snijman 292 (NBG0123648-0! holo.; PRE 0665215-0!, K iso.).
14. Rhadamanthopsis (Oberm.) Speta in Phyton (Horn, Austria) 38(1): 74 (1998) (Fig. 35). Typus generis:—R.
namibensis (Oberm.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso (holotype).
≡ Rhadamanthus subgen. Rhadamanthopsis Oberm. in Bothalia 13(1-2): 137 (1980). Typus subgeneris:—Rhadamanthus namibensis
Oberm. (holotype).
≡ Drimia sect. Rhadamanthopsis (Oberm.) J.C.Manning & Goldblatt in Strelitzia 40: 124 (2018). Typus sectionis:—D. namibensis
(Oberm.) J.C.Manning & Goldblatt (holotype).
= Drimia sect. Hyacinthoides J.C.Manning & Goldblatt in Strelitzia 40: 71 (2018). Typus sectionis:—D. hyacinthoides Baker (holotype).
Description:—Medium sized bulbous geophyte. Bulb hypogeal, ovoid to subglobose, 3‒7 cm in diam., solitary or
commonly clump forming, with compact or commonly loose, white, fleshy scales. Leaves 1‒6 per bulb, hysteranthous,
4‒25 cm long, narrowly lanceolate, canaliculate and leathery or oblong, soft and commonly with longitudinal furrows,
green, smooth, glabrous, sometimes surrounded by transversally barred cataphylls at base. Inflorescence an elongated
raceme, 8‒35 cm long, with 8‒90 flowers; peduncle terete, erect, 10‒40 cm long, smooth or rarely scabrid-puberulous

FIGURE 35. Species of Rhadamanthopsis (Oberm.) Speta displayed in horizontal rows of images. 1. Rhadamanthopsis haworthioides
(Baker) Mart.-Azorín et al.; 2. Rhadamanthopsis hyacinthoides (Baker) Mart.-Azorín et al.; 3. Rhadamanthopsis karooicus (Oberm.)
Mart.-Azorín et al.; 4. Rhadamanthopsis monophyllus (Oberm. ex J.C.Manning & Goldblatt) Mart.-Azorín et al.; 5. Rhadamanthopsis
namibensis (Oberm.) Mart.-Azorín et al. (Photographs of wild inflorescences by H. Kolberg).
at base; pedicels subpatent, arquing downwards, 8‒17 mm long, glabrous. Bracts narrowly lanceolate, 2‒6 mm long,
lowermost with spur 2‒8 mm long; bracteoles present and distinct. Flowers campanulate, nodding, diurnal. Tepals 6, 7‒
14 mm long, white, greenish or purplish, with darker longitudinal band, connate for 2‒5 mm to form basal campanulate
tube and suberect to slightly spreading, straight free lobes. Stamens 6; filaments adnate to tepals for 0.5‒3.0 mm from
base and free portions suberect to sigmoid and incurved to ovary, 2.0‒3.5 mm long; anthers medifixed or subasifixed,
dehiscing for their whole length, oblong to sagittate, connivent to style. Ovary ovoid to oblong, trigonous, 2‒5 mm
long, pale green to yellowish, well differentiated from style. Style erect, 2‒7 mm long. Stigma trigonous, small and
papillose. Capsule ovoid, 5‒16 mm long, valves completely dehiscing from base; tepals cohering and inrolled above
ovary after anthesis, circumcissile from base and persisting as a cap at the top of the developing capsules. Seeds 4‒7
mm long, black to dark brown, subellipsoid flattened with prominent central embryo and wide, flat wings, testa, with
sinuous anticlinal cell walls.
Number of species and distribution:—Rhadamanthopsis comprises five species restricted to southern and western
South Africa and southern Namibia (Fig. 29), which are restricted to the Cape, Karoo-Namib, and Uzambara-Zululand
Regions (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a). For further information on Rhadamanthopsis
species see Obermeyer (1980a), Jessop (1977), and Manning & Goldblatt (2018).
Karyology:—2n=12, 16, 18 (Speta 1998a with no reference).
History, diagnostic characters, and taxonomic relationships:—Obermeyer (1980a) and Martínez-Azorín &
Crespo (2014) described Rhadamanthus namibensis Obermeyer in Martínez-Azorín & Crespo (2014: 1331) from
southern Namibia and R. karooicus Obermeyer in Martínez-Azorín & Crespo (2014: 1331) from western South Africa.
Obermeyer (1980a) also proposed Rhadamanthus subgen. Rhadamanthopsis Oberm. to include these two species, that
differed from typical Rhadamanthus species (sensu Nordenstam 1970) by the loculicidal dehiscence of anthers instead
of by apical pores or slits. These two species show a very distinct and easily recognizable flower syndrome within
Urgineoideae, characterised by a combination of diurnal, nodding and campanulate flowers, with tepals distinctly
connate at the base for about a third to 2/5 of their length and free apical lobes that are suberect to only slightly spreading,
with included stamens, shortly adnate filaments, and free portions connivent to the style. Further, they produce distinct
bracteoles and wide leaves, that are usually longitudinally striate (except in the type species), strongly differing from
Rhadamanthus species in the sense of this work, as that genus always lacks bracteoles and produces morphologically
different leaves and flowers. Other species sharing the flower and leaf morphology of R. subg. Rhadamanthopsis had
previously been described, including Drimia hyacinthoides Baker (1874c: 6), Ornithogalum haworthioides Baker
(1878: 322) [= Drimia bolusii Baker (1897: 443) nom. nov., not to be confused with Drimia haworthioides Baker
(1875: 366)], and D. monophylla Oberm. ex Manning & Goldblatt (2018: 128).
Speta (1998b) raised Rhadamanthopsis to genus rank based on the reproductive and vegetative characters
mentioned above, and cited the chromosome numbers 2n=16, 18 for this genus, suggesting a clear difference from
common chromosome counts in the subfamily. The phylogenetic analyses of Pfosser & Speta (1999) included a sample
of this group named “Karoophila bolusii” (a genus name not formally published), that is sister to Charybdis (= Squilla
in this work). Pfosser & Speta (2001, 2004) added a sample of Rhadamanthopsis from Namibia that nested sister
to the “Karoophila” sample. Pfosser et al. (2012) found four samples of Rhadamanthopsis forming a clade with
moderate support. The phylogenetic analyses of Martínez-Azorín et al. (2023a) include 14 samples of this group. The
combination of plastidial and nuclear regions place samples of Rhadamanthopsis namibensis in a clade sister to a sample
of Aulostemon. Within a large polytomy, including also the latter clade, the remaining samples of Rhadamanthopsis in
the sense of this work are monophyletic and strongly supported. However, when morphological data are included in the
plastid phylogenetic analyses, Rhadamanthopsis recovers monophyly in the sense of this work. The Rhadamanthopsis
samples are divided in three biogeographically congruent subclades with collapsed relationships. The first subclade
includes four samples of R. namibensis from southern Namibia while the second subclade comprises four samples of R.
karooicus sensu lato from western South Africa, mostly Namaqualand. Finally, six samples of Drimia hyacinthoides,
O. haworthioides, and D. monophylla from central southern and eastern South Africa form another clade. Some
morphological differences exist among taxa included in the three biogeographic subclades. Firstly, the Namibian
samples show compact bulbs and narrowly lanceolate, canaliculate, not striate, suberect to spreading leaves. Secondly,
the northwestern samples present mostly compact bulbs with closely imbricate scales and flat, mostly prostrate, oblong,
leaves with longitudinal furrows. Finally, samples from southern and eastern South Africa show bulbs with loose,
fleshy and pediculated scales, sharing the longitudinal furrows of leaves with the Namaqualand samples. Despite the
differences in vegetative morphology between those groups, we here accept Rhadamanthopsis to include all above
mentioned taxa that share a distinct syndrome of morphological characters: the mostly wide hysteranthous leaves, the
long racemes with distinct bracteoles, and the distinct nodding campanulate flowers.
Manning & Goldblatt (2018) placed the species accepted here in Rhadamanthopsis in two sections: Drimia sect.
Rhadamanthopsis to include D. namibensis, D. karooica, and D. monophylla; and their monotypic D. sect. Hyacinthoides
to include Drimia hyacinthoides. Although these sections were separated in the key of Manning & Goldblatt (2018)
by the presence or absence of bracteoles, all species in both sections have distinct bracteoles and nearly the same
flower morphology. As the segregation of these sections is supported neither by phylogenetic relationships nor by
morphology, sect. Hyacinthoides is not accepted here.
Rhadamanthopsis haworthioides (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Ornithogalum haworthioides Baker
in J. Bot. 16: 322 (1878), basionym ≡ Drimia bolusii Baker, Fl. Cap. (Harvey) 6(3): 443 (1897), nom. nov. [non D. haworthioides
Baker in Gard. Chron. 1: 366 (1875)] (Figs 2.20, 35.1). Type:—SOUTH AFRICA. Graaf Reinet, Cave Mountain, 2900 feet, Bolus
814 (Collection not found at K, S. Rokni pers. comm.; neither at BOL, PRE, nor NBG).
Rhadamanthopsis hyacinthoides (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia hyacinthoides Baker in J.
Bot. 12: 6 (1874), basionym (Figs 3.1, 35.2). Type:—SOUTH AFRICA. Eastern Cape. Grahamstown (3326): shady valleys near
Grahamstown, (–BC), November without year, P. MacOwan 1465 (GRA0000459! lecto. designated by Jessop in J. S. African Bot.
43: 27. 1977; K000400567!, NY00319647! isolecto.).
Rhadamanthopsis karooicus (Oberm.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Rhadamanthus karooicus Oberm. in Taxon
63(6): 1331 (2014), basionym ≡ Rhadamanthus karooicus Oberm. in Bothalia 13(1-2): 138 (1980), nom. inval. ≡ Rhadamanthopsis
karooicus (Oberm.) Speta, Phyton (Horn, Austria) 38(1): 74 (1998), nom. inval. ≡ Drimia karooica (Oberm.) J.C.Manning &
Goldblatt in Strelitzia 9: 712 (2000), nom. inval. ≡ Drimia karooica (Oberm.) J.C.Manning & Goldblatt in Strelitzia 40: 126 (2018)
(Figs 3.2, 35.3). Type:—SOUTH AFRICA. Western Cape. Montagu (3320): Laingsburg, Keurfontein farm, (–BB), December 1974
[flowering ex hort.], J. van Zanten s.n. (PRE0240643-1!: plant with inflorescence, holo.).
Rhadamanthopsis monophyllus (Oberm. ex J.C.Manning & Goldblatt) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia
monophylla Oberm. ex J.C.Manning & Goldblatt in Strelitzia 40: 128 (2018), basionym. (Figs 3.3, 35.4). Type:—SOUTH AFRICA.
Eastern Cape. Somerset East (3225): Stone Fountain farm, (–DA), 19 April 1963 [leafing bulbs], R. Bayliss 1345 (NBG84207!
holo.).
Comments:—Drimia monophylla sensu Manning & Goldblatt (2018) fits with Ornithogalum haworthioides Baker (1878) (= Drimia
bolusii Baker 1897) based on the bulb structure, lorate leaves and campanulate flowers with included perigynous stamens. However,
Manning & Goldblatt (2018) considered O. haworthioides as synonym of Drimia haworthioides Baker (1875), a member of Drimia
s.str. When O. haworthioides is placed in Rhadamanthopsis, it apparently only differs from D. monophylla (with 1‒2 leaves) by the
5‒6 leaves per bulb. Further studies are needed to evaluate their exact relationship.
Rhadamanthopsis namibensis (Oberm.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Rhadamanthus namibensis Oberm.
in Taxon 63(6): 1331 (2014), basionym ≡ Rhadamanthus namibensis Oberm. in Bothalia 13(1−2): 137 (1980), nom. inval. ≡
Rhadamanthopsis namibensis (Oberm.) Speta in Phyton (Horn, Austria) 38(1): 74 (1998), nom. inval. ≡ Drimia namibensis
J.C.Manning & Goldblatt in Strelitzia 9: 712 (2000), nom. inval. ≡ Drimia namibensis (Oberm.) J.C.Manning & Goldblatt in Strelitzia
40: 125 (2018) (Figs 3.4, 35.5). Type:—NAMIBIA. Witputz (2716): Witputz-Suid, 1 km SE of Police Station, (–DA), Giess 13781
(PRE0488688-1!: inflorescence in two pieces, holo.; K000257234!, M0107238!, WIND! iso.).
15. Rhadamanthus Salisb., Gen. Pl. [Salisbury]: 37 (1866) (Figs 36–38). Typus generis:—R. convallarioides (L.f.)
Baker (holotype).
= Drimia sect. Rhadamanthus (Salisb.) J.C.Manning & Goldblatt in Strelitzia 40: 129 (2018). Typus sectionis:—R. convallarioides (L.f.)
Baker (holotype).
= Drimia sect. Sclerophyllae J.C.Manning & Goldblatt in Strelitzia 40: 101 (2018). Typus sectionis:—D. sclerophylla J.C.Manning &
Goldblatt ≡ Urginea rigidifolia Baker (holotype).
Description:—Bulbous geophyte. Bulb hypogeal, ovoid to globose, with compact white, scales, rarely loose and pink.
Leaves 1‒20 per bulb, hysteranthous, 1.5‒30 cm long, filiform and subterete or ovate-lanceolate to suborbicular and
flat or cucullate, leathery, green, sometimes purplish at base, smooth, glabrous or rarely ciliate, sometimes surrounded
by transversally barred cataphylls at base. Inflorescence racemose, usually elongated, 2‒22 cm long, with 5‒50(–70)
flowers; peduncle terete, erect, 5‒30 cm long, smooth or commonly scabrid-puberulous at base; pedicels subpatent,
straight to arquing, 3‒20 mm long, glabrous or rarely minutely scabrid. Bracts narrowly lanceolate, 1‒2 mm long,
lowermost with spur 0.5‒1.5 mm long; bracteoles absent. Flowers stellate, campanulate or urceolate, erect-patent
to nodding, diurnal. Tepals 6, 3‒10 mm long, white, yellow, brown or pinkish, with darker longitudinal band, free
and patent to reflexed or connate for up to 3 mm and then campanulate or urceolate. Stamens 6, connivent to style
or spreading; filaments free and 2‒3 mm long or adnate to tepals for up to 2 mm from base and free portions 0.3‒1.5
mm long; anthers medifixed, dehiscing by apical slits to their whole length, oblong to sagittate, sometimes basally
apendiculate. Ovary ovoid, trigonous, 1.5‒4.0 mm long, pale green to yellowish, well differentiated from style. Style
erect, 0.5‒2.5 mm long. Stigma trigonous, papillose. Capsule ovoid to subglobose, 3‒7 mm long, valves completely
dehiscing from base; tepals cohering and inrolled above ovary after anthesis, circumcissile from base and persisting
as a cap at the top of the developing capsules. Seeds subellipsoid to subtrigonous, flattened with prominent central
embryo and wide, flat wings, 2‒5 mm long, black to dark brown, with sinuous anticlinal cell walls.
FIGURE 36. Species of Rhadamanthus Salisb. 1. Rhadamanthus convallarioides (L.f.) Salisb. ex Baker from Jacquin (1797: t. 81, as
Hyacinthus convallarioides L.f.); 2. Rhadamanthus urantherus R.A.Dyer from Dyer (1934: t. 3247); 3. Rhadamanthus cochlearis (Mart.-
Azorín et al.) Mart.-Azorín et al. from Martínez-Azorín et al. (2013: 333, as Drimia cochlearis Mart.-Azorín et al.).
Number of species and distribution:—Rhadamanthus includes twelve species, occurring in southern and western
South Africa and southern Namibia (Fig. 29), and restricted to the Cape, Karoo-Namib and Uzambara-Zululand
Regions, and the Southern Section of the Zambezian Subregion (sensu Takhtajan 1986 and Martínez-Azorín et al.
2023a). For further information on Rhadamanthus see Dyer (1934), Nordenstam (1970), Jessop (1977), Martínez-
Azorín et al. (2013b), and Manning & Goldblatt (2018).
Karyology:—2n=18 (Speta 1998a, with no reference). 2n=20 (De Wet 1957, as Rhadamanthus convallarioides
Salisb.).
History, diagnostic characters, and taxonomic relationships:—Salisbury (1866) described Rhadamanthus to
include a single species, Rhadamanthus convallarioides (Linnaeus f. 1782: 204) Salisb. ex Baker (1871: 434), based
on the nodding urceolate to campanulate flowers with the tepals fused for their basal portion, the stamens connivent
to the gynoecium and the anthers dehiscing by apical pore-like slits. Baker (1897) described R. cyanelloides Baker
(1897: 444), and later Dyer (1934) described R. urantherus Dyer (1934: t. 3247), both sharing the peculiar dehiscence
of anthers. Nordenstam (1970) presented a revision of Rhadamanthus in which he accepted the previous three species
and added six further new species, namely R. albiflorus Nordenstam (1970: 177), R. arenicola Nordenstam (1970:
166), R. fasciatus Nordenstam (1970: 174), R. montanus B.Nord. in Martínez-Azorín & Crespo (2014: 1331), R.
platyphyllus B.Nord. in Martínez-Azorín & Crespo (2014: 1332), and R. secundus Nordenstam (1970: 168). The genus
was characterised by the anthers dehiscing incompletely by introrse, longitudinal slits, and it was also reported that in
R. fasciatus and R. albiflorus the sutures reached down below the middle of the thecae. In that concept, Rhadamanthus
includes a wide range of morphological variation such as bulb scales compact or loose; leaves filiform, numerous and
erect or few, wide, flat and appresed to the ground; flowers nodding to patent, stellate to urceolate-campanulate; tepals

from nearly free to fused about half of their length; and stamens free or fused along the anthers. Later, Obermeyer
(1980a) described two further species, Rhadamanthus namibensis and R. karooicus, and proposed R. subgenus
Rhadamanthopsis (here accepted at genus rank), based on the loculicidal anther dehiscence, to seggregate the two
latter species from typical Rhadamanthus. Speta (1998b) transferred all nine known species of Rhodocodon, a genus
endemic to Madagascar, into Rhadamanthus based on the similarities in flower morphology and anthers dehiscence.
However, Knirsch et al. (2015) presented solid morphological and molecular differences and reinstated Rhodocodon.
The latest species described in the genus is Rhadamanthus involutus J.C.Manning & Snijman in Snijman et al. (1999:
113).
FIGURE 37. Species of Rhadamanthus Salisb. displayed in horizontal rows of images. 1. Rhadamanthus arenicola B.Nord.; 2.
Rhadamanthus ciliatus (L.f.) Mart.-Azorín et al.; 3. Rhadamanthus cochlearis (Mart.-Azorín et al.) Mart.-Azorín et al.; 4. Rhadamanthus
convallarioides (L.f.) Salisb. ex Baker.
Martínez-Azorín et al. (2013a) excluded R. cyanelloides as Sagittanthera cyanelloides on the basis of the anthers
connate in a cone around the style, among other distinguishing characters. Recently, Manning & Goldblatt (2018)
recognised Drimia sect. Rhadamanthus (Salisbury 1866: 37) Manning & Goldblatt (2018: 129), including nine species
mostly corresponding to the concept of Nordenstam (1970), and followed Martínez-Azorín et al. (2013b) in excluding
R. cyanelloides as their monotypic D. sect. Sagittanthera (Martínez-Azorín et al. 2013a: 46) Manning & Goldblatt
(2018: 73). Even with the exclusion of S. cyanelloides, D. sect Rhadamanthus sensu Manning & Goldblatt (2018)
is paraphyletic according to previous studies by Pfosser et al. (2012) and the present one, in which R. platyphyllus
and Drimia oliverorum J.C.Manning in Manning & Oliver (2009: 225) are aberrant in the genus in having 1‒2
velutinous leaves that are ovate, flat and adpressed to the ground showing distinct longitudinal furrows, and they form
an independent well-supported clade positioned far from the Rhadamanthus clade, a fact that favours acceptance of
Striatula to include the latter two species (Pinter et al. 2019).
FIGURE 38. Species of Rhadamanthus Salisb. displayed in horizontal rows of images. 1. Rhadamanthus fasciatus B.Nord.; 2.
Rhadamanthus rigidifolius (Baker) Mart.-Azorín et al.; 3. Rhadamanthus secundus B.Nord. (Photographs in flower by H. Kolberg); 4.
Rhadamanthus urantherus R.A.Dyer.
The phylogenetic studies by Martínez-Azorín et al. (2023a) include 21 samples that form a well supported clade
that is here assimilated to Rhadamanthus, comprising R. arenicola, R. convallarioides, R. fasciatus, R. montanus, R.
urantherus together with samples of Urginea ciliata (Linnaeus f. 1782: 199) Baker (1873b: 218), U. rigidifolia Baker
(1878: 323), U. muirii Brown (1933: 334), and Drimia cochlearis Mart.-Azorín, M.B.Crespo & A.P.Dold in Martínez-
Azorín & Crespo (2014: 1329). The latter four species form a distinct group that is included by Manning & Goldblatt
(2018) in D. sect. Sclerophyllae based on their nodding globular buds before anthesis and patent to suberect stellate
flowers borne on pedicels longer than tepals, the spreading filaments, and the anthers dehiscing along their entire length.

Our phylogenetic results show D. sect. Rhadamanthus sensu Manning & Goldblatt (2018) as paraphyletic without
the inclusion of D. sect. Sclerophyllae, which however forms a strongly supported subclade within Rhadamanthus
(Martínez-Azorín et al. 2023a). We preliminarily include taxa of D. sect. Sclerophyllae in Rhadamanthus, based on the
obtained phylogenetic results. Moreover, the stellate flowers with long, spreading stamens of D. sect. Sclerophyllae
are atypical for Rhadamanthus s.str., which has short stamens connivent to the gynoecium. However, there exists an
apparent transition from flowers of R. albiflorus and R. fasciatus (Nordenstam 1970) that are subpatent with nearly
free and spreading tepals, to the nodding, urceolate flowers of R. arenicola, R. convallarioides, R. secundus, and R.
urantherus, with various degrees of tepal connation. In our analyses, three samples of R. fasciatus with sub-stellate
flowers form a clade that is sister to the remaining samples and taxa in the genus, indicative of the primitive stage of
stellate flowers and early swifts in flower morphology within this lineage. In this sense, Rhadamanthus includes twelve
species restricted to South Africa and southern Namibia. No samples of R. albiflorus, R. involutus, and R. secundus
were available for the present study. Further studies including a complete sampling in the genus are necessary to
evaluate possible alternative classifications within Rhadamanthus.
Rhadamanthus albiflorus B.Nord. in Bot. Not. 123: 177 (1970) ≡ Drimia albiflora (B.Nord.) J.C.Manning & Goldblatt in Strelitzia 9:
711 (2000). Type:—SOUTH AFRICA. Western Cape. Bredasdorp (3420): Hesquaspoort, (–AA), elev. 245 m, 20 December 1962,
J.P.H. Acocks 23242 (PRE0051091-0! holo.).
= Rhadamanthus montaguense Oberm., nom. nud. in sched.: near Montagu Baths, December 1892, H. Bolus 2797 (NBG 72561!).
Rhadamanthus arenicola B.Nord. in Bot. Not. 123: 166 (1970) ≡ Drimia arenicola (B.Nord.) J.C.Manning & Goldblatt in Strelitzia 9:
711 (2000) (Figs 3.5, 37.1). Type:—SOUTH AFRICA. Northern Cape. Hondeklipbaai (3017): 0.5 miles S of Wallekraal, (–BC),
October 1924, Pillans s.n. sub BOL18253 (BOL140332! holo.).
Rhadamanthus ciliatus (L.f.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Ornithogalum ciliatum L.f., Suppl. Pl.: 199 (1782),
basionym ≡ Urginea ciliata (L.f.) Baker in J. Linn. Soc., Bot. 13: 218 (1873) ≡ Drimia ciliata (L.f.) J.C.Manning & Goldblatt in
Bothalia 33(1): 111 (2003) (Figs 3.6, 37.2). Type:—SOUTH AFRICA. Caput bonae Spei, Thunberg s.n. (UPS-THUNB [8281] lecto.
designated as “type” by Jessop in J. S. African Bot. 43: 279. 1977; S-G 79 06 [image!] probl. iso.).
Rhadamanthus cochlearis (Mart.-Azorín, M.B.Crespo & A.P.Dold) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia
cochlearis Mart.-Azorín, M.B.Crespo & A.P.Dold in Taxon 63(6): 1329 (2014), basionym ≡ Drimia cochlearis Mart.-Azorín,
M.B.Crespo & A.P.Dold in Syst. Bot. 38(2): 334 (2013), nom. inval. (Figs 3.7, 36.3, 37.3). Type:—SOUTH AFRICA. Western Cape,
ca. 11 km SW of Calitzdorp, Gamkaberg Nature Reserve (former Groenefontein Nature Reserve), elev. 408 m, 29 September 2011,
M. Martínez-Azorín, A.P. Dold, J. Vlok & A. Martínez-Soler MMA941 (GRA holo.: only the bulbs with leaves; ABH iso.).
Rhadamanthus convallarioides (L.f.) Salisb. ex Baker in J. Linn. Soc., Bot. 11: 434 (1871) ≡ Hyacinthus convallarioides L.f., Suppl.
Pl.: 204 (1782) ≡ Drimia convallarioides (L.f.) J.C.Manning & Goldblatt in Strelitzia 9: 711 (2000) (Figs 3.8, 36.1, 37.4). Type:—
SOUTH AFRICA. Northern Cape. Sutherland (3220): karroo below Roggeveld, (–CA), Thunberg s.n. UPS-THUNB8519 (UPS-
THUNB! lecto. designated by Nordenstam in Bot. Not. 123: 159. 1970).
Rhadamanthus fasciatus B.Nord. in Bot. Not. 123: 174 (1970) ≡ Drimia fasciata (B.Nord.) J.C.Manning & Goldblatt in Strelitzia 9:
711 (2000) (Fig. 38.1). Type:—SOUTH AFRICA. Northern Cape. Kimberley (2824): Nooitgedacht, c. 10 miles SE of Barkly West,
(–DA), elev. c. 3800 ft., 15 October 1960, O.A. Leistner 1983 (PRE0051090-0! holo.; K000257232!, LD iso.).
Rhadamanthus involutus J.C.Manning & Snijman in Novon 9(1): 113 (1999) ≡ Drimia involuta (J.C.Manning & Snijman) J.C.Manning
& Goldblatt in Strelitzia 9: 712 (2000) ≡ Rhadamanthus involutus J.C.Manning & Snijman in Taxon 63: 1330 (2014), nom. superfl.
Type:—SOUTH AFRICA. Northern Cape. Calvinia (3119): Arendskraal farm, W of Nieuwoudtville, (–AC), 20 December 1995
flowers, D. Snijman & J.C. Manning 1525 (NBG0158587-0! lecto. designated here: the inflorescence on the upper left hand side
corner; K!, MO176399!, PRE0461606-0! isolecto.: only the flowering material).
in the protologue will rule, and considering that the protologue in Snijman et al. (1999) only indicates a single collecting date (20
December 1995) in the holotype designation, a lectotypification is needed from the mixed type collection NBG0158587-0. Based on
the new considerations, the description made by Martínez-Azorín & Crespo (2014: 1330) is therefore nomenclaturally inoperative,
since it is either superfluous or not valid.
Rhadamanthus montanus B.Nord. in Taxon 63(6): 1331 (2014) ≡ Rhadamanthus montanus B.Nord. in Bot. Not. 123: 162 (1970), nom.
inval. Type:—SOUTH AFRICA. Western Cape. Cape Town (3318): Jonkershoek Twins, (–DD), elev. 3000–4000 ft., 12 February
1945, E. Esterhuysen 11456 (BOL140331! holo.: a bulb with an inflorescence with eight flowers in the upper part; K000257229!,
PRE0051072-0!, NBG72562! iso.).
Rhadamanthus muirii (N.E.Br.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea muirii N.E.Br. in Gard. Chron., Ser. 3.
93: 334 (1933), basionym. Type:—SOUTH AFRICA. Near Riversdale, 1933, J. Muir 4846 (K, first-step lecto. designated by Jessop
in J. S. African Bot. 43: 315. 1977; second-step lecto. designated here: K000257341!).
Rhadamanthus rigidifolius (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea rigidifolia Baker in J. Bot. 16: 323
(1878), basionym ≡ Drimia sclerophylla J.C.Manning & Goldblatt in Strelitzia 9: 712 (2000), nom. nov. [non Drimia rigidifolia
Baker in J. Linn. Soc., Bot. 11: 420 (1871) = D. media Jacq.] (Figs 3.9, 38.2). Type:—SOUTH AFRICA. Eastern Cape. Graaff-Reinet
(3224): Karroo, near Graaff-Reinet, (–BC), elev. 2500 ft., September 1873, H. Bolus 783 (K000257339! lecto. designated as “holo.”
by Jessop in J. S. African Bot. 43: 315. 1977).
Rhadamanthus secundus B.Nord. in Bot. Not. 123: 168 (1970) ≡ Drimia secunda (B.Nord.) J.C.Manning & Goldblatt in Strelitzia 9: 712
(2000) (Fig. 38.3). Type:—NAMIBIA. Lüderitz (2615): Lüderitz-Süd, Kovisberge, (–CA), 30 September 1959, W. Giess 2350 (M
holo.; PRE0051080-0!, WIND iso.).
= Ophioasylon peculiare Dinter, nom. nud. in sched.: NAMIBIA. Bogenfels (2715): Buchuberge, (–DD), 5 July 1929, K. Dinter 6492
(K000257231!, PRE0051078!, SAM).
Rhadamanthus urantherus R.A.Dyer in Hooker’s Icon. Pl. sér. 5, 33: t. 3247 (1934) ≡ Drimia uranthera (R.A.Dyer) J.C.Manning &
Goldblatt in Strelitzia 9: 712 (2000) (Figs 3.10, 36.2, 38.4). Type:—SOUTH AFRICA. Western Cape. Oudtshoorn (3322): 1 mile
E of Oudtshoorn, (–CA), 29 March 1933 [leafing 11 Oct. 1933 ex hort.], Barker s.n. K933/32 (K000257230! lecto. designated by
Martínez-Azorín et al. in Phytotaxa 201: 169. 2015: only the bulb with two inflorescences in centre of sheet).
16. Rhodocodon Baker in J. Linn. Soc., Bot. 18: 280, t. 8 (1881) (Figs 39–42). Typus generis:—R. madagascariensis
Baker (holotype) = Rhodocodon mascarenensis (Baker) Knirsch, Mart.-Azorín & Wetschnig ≡ Urginea mascarenensis
Baker
≡ Drimia sect. Rhodocodon (Baker) J.C.Manning & Goldblatt in Strelitzia 40: 155 (2018). Typus sectionis:—R. madagascariensis Baker
(holotype).
Description:—Bulbous geophyte. Bulb usually hypogeal, sometimes epigeal and epiphytic, ovoid to subglobose, bulb
scales compact. Roots thickened and branched. Leaves 1‒8 per bulb, from filiform to flattened and wide, sometimes
with dorsal keel, green, glabrous. Inflorescence racemose, usually elongated and multiflowered, 5‒70 cm long, with
10‒120 flowers; peduncle erect, terete glabrous, smooth, 9‒38 cm long; pedicels from erect to patent or nodding,
1‒40 mm long. Bracts small, lanceolate, 1.0‒2.5 mm long, with lanceolate-oblong spur, 0.1‒2.0 mm long, both
gradually diminishing in size to the apex of the raceme; bracteoles absent or present but indistinct. Flowers urceolate
to campanulate, erect-patent to nodding, lasting for 3‒7 days. Tepals 6, 3.4‒12.0 mm long, white, greenish, brownish,
rose or red coloured, fused for most of their length with free apical lobes that are from erect to reflexed, with single
central nerve. Stamens 6, shorter than perigone segments, mostly included in perigone tube and connivent to ovary and
style; filaments adnate to perigone and arising from lower half of tube, filiform to flattened, free portions 0.3‒3.5 mm
long; anthers dorsifixed, sagittate or oblong, dehiscing along their whole length but sometimes appearing pore like at
the apex. Ovary subglobose, conical to cylindrical, trigonous, with 2‒17 ovules per locule, collateral to more or less
loosely arranged; style filiform, erect, 1.3‒5.0 mm long; stigma simple or inconspicuously 3-lobed. Capsule oblong-
triangular to subglobose, 3‒12 mm long, valves completely dehiscing from base, withered tepals usually persisting
at base of dehisced capsule (rarely abscising from base and remaining laterally attached at the top of the capsule).
Seeds 2.0‒6.2 mm long, ellipsoidal or somewhat compressed, with black, glossy, brittle testa and distinct mostly light
brown or rarely black raphe, sometimes absent; or rarely fusiform, flat, testa brown, brittle, lacking raphe, with sinuous
anticlinal cell walls.
Number of species and distribution:—Rhodocodon currently includes 19 species that are restricted to Madagascar
(Fig. 29). It is therefore the only urgineoid genus that has been recorded so far in the Madagascan Region in the
sense of Takhtajan (1986; see also Martínez-Azorín et al. 2023a). For further information on Rhodocodon species see
Knirsch et al. (2015, 2016, 2019).
FIGURE 39. Species of Rhodocodon Baker. 1. Rhodocodon mascarenensis (Baker) Knirsch et al. from Baker (1881: t. 8, as Rhodocodon
madagascariensis Baker); 2. Rhodocodon mascarenensis (Baker) Knirsch et al., R. cyathiformis H.Perrier ex Knirsch et al., R. rotundus
H.Perrier ex Knirsch et al., R. calcicola Knirsch et al., R. monophyllus Knirsch et al., R. linearifolius Knirsch et al., and R. urgineoides
Baker et al. from Perrier de la Bâthie (1938: 121).

History, diagnostic characters, and taxonomic relationships:—Baker (1874a) described Urginea mascarenensis
Baker (1874a: 363) from Madagascar, with hysteranthous filiform leaves, campanulate white nodding flowers and
spurred bracts. Few years later, Baker (1881) described Rhodocodon based on the distinct flower morphology of the
latter species, showing tepals connate for most of their length and short free apical lobes, the included stamens with
adnate and short filaments arising from the lower half of the perigone, and the ovate ovary and erect narrow style,
and included a single species: R. madagascariensis Baker (1881: 280) a later synonym of U. mascarenensis. Perrier
de la Bâthie (1938) presented a revision on the endemic Madagascan genus Rhodocodon, where six species and four
varieties were recognised, although those names were not validly published (Knirsch et al. 2015). The recent revision
of Rhodocodon by Knirsch et al. (2015, 2016, 2019) accepted 19 species in the genus, which is characterised by
flowers urceolate to campanulate, lasting for 3‒7 days; perigone segments fused for most of their length, commonly
persisting at the base of ripe capsules; filaments adnate to the perigone and arising from the lower half of the tube; and
seeds ellipsoidal or somewhat compressed, usually with a distinct raphe, or rarely fusiform and flat (Brudermann et
al. 2018).
FIGURE 40. Species of Rhodocodon Baker displayed in horizontal rows of images. 1. Rhodocodon calcicola Knirsch et al.; 2. Rhodocodon
campanulatus Knirsch et al.; 3. Rhodocodon cryptopodus (Baker) Knirsch et al.; 4. Rhodocodon cyathiformis H.Perrier ex Knirsch et al.;
5. Rhodocodon floribundus H.Perrier ex Knirsch et al.

FIGURE 41. Species of Rhodocodon Baker displayed in horizontal rows of images. 1. Rhodocodon graciliscapus Knirsch et al.; 2.
Rhodocodon intermedius H.Perrier ex Knirsch et al.; 3. Rhodocodon jackyi Knirsch et al.; 4. Rhodocodon mascarenensis (Baker) Knirsch
et al.; 5. Rhodocodon monophyllus Knirsch et al.
FIGURE 42. Species of Rhodocodon Baker displayed in horizontal rows of images. 1. Rhodocodon perrieri Knirsch et al.; 2. Rhodocodon
petrae Knirsch et al.; 3. Rhodocodon rubescens Knirsch et al.; 4. Rhodocodon siederi Knirsch et al.; 5. Rhodocodon urgineoides Baker.

Previous phylogenetic work has shown Madagascan samples of Rhodocodon to be monophyletic in a well-
supported clade (Pfosser & Speta 2001, 2004, Pfosser et al. 2006, 2012, Knirsch 2012), which is in agreement with the
phylogenetic results of Martínez-Azorín et al. (2023a), in which 12 samples of the genus form a well supported clade
that is sister to Triandra from western South Africa. Therefore, we here accept Rhodocodon as an endemic Madagascan
genus showing a unique combination of morphological characters that form an independent phylogenetic lineage.
Rhodocodon apiculatus H.Perrier ex Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 106 (2015) ≡ R. apiculatus H.Perrier,
Fl. Madagasc. Fam. 40: 122 (1938), nom. inval. Type:—MADAGASCAR. Rocailles calcaires, eocènes, boisées, près de Majunga
[Mahajanga], December 1924, Perrier 16821 (P00573458! holo.).
Rhodocodon calcicola Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 108 (2015) ≡ R. calcicola H.Perrier, Fl. Madagasc. Fam.
40: 115 (1938), nom. inval. ≡ Drimia calcicola (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004), nom.
inval. (Figs 39.2, 40.1). Type:—MADAGASCAR. Province Mahajanga, few kilometers north of Mahajanga, elev. 60 m, ex hort. 30
April 2014, W. Knirsch, A. Sieder & J. Andriantiana 6293 (GZU000322310! holo.).
Rhodocodon campanulatus Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 110 (2015) ≡ R. rotundus var. campanulatus
H.Perrier, Fl. Madagasc. Fam. 40: 122 (1938), nom. inval. (Figs 3.11, 40.2). Type:—MADAGASCAR. Province Majahanga, rochers
basaltiques, découverts et humides pendant les pluies. Plateau d’Antanimena entre la Betsiboka et la Mahavavy, May 1905, Perrier
10947 (P01848487! holo.).
Rhodocodon cryptopodus (Baker) Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 112 (2015) ≡ Hyacinthus cryptopodus Baker
in J. Linn. Soc., Bot. 20: 274 (1883), basionym ≡ Drimia cryptopoda (Baker) Pfosser, Wetschnig & Speta in Linzer Biol. Beitr. 38(2):
1736 (2006) (Figs 3.12, 40.3). Type:—MADAGASCAR. Central Madagascar, October 1882, R. Baron 2164 (K000098171! holo.:
the two fragments/specimens in the upper left hand side corner of the sheet).
Rhodocodon cyathiformis H.Perrier ex Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 114 (2015) ≡ R. cyathiformis H.Perrier,
Fl. Madagasc. Fam. 40: 118 (1938), nom. inval. ≡ Drimia cyathiformis (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot.
60(3): 556 (2004), nom. inval. (Figs 3.13, 38.2, 40.4). Type:—MADAGASCAR. Rochers ombragés, Firingalava (Boina), March
1898, Perrier 516 (P00573463! holo.; P00573464!, P00573465! iso.).
Rhodocodon floribundus H.Perrier ex Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 116 (2015) ≡ R. floribundus H.Perrier,
Fl. Madagasc. Fam. 40: 117 (1938), nom. inval. ≡ Drimia floribunda (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot.
60(3): 556 (2004), nom. inval. (Figs 3.14, 40.5). Type:—MADAGASCAR. Rocailles (gneiss), elev. 1500 m, Manankazo, au N.E.
d’Ankazobe, Novembre 1913, Perrier 10960 (P00573460! holo.; P00573459! iso.).
Rhodocodon graciliscapus Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 118 (2015) ≡ R. calcicola var. graciliscapus H.Perrier,
Fl. Madagasc. Fam. 40: 116 (1938), nom. inval. (Figs 3.15, 41.1). Type:—MADAGASCAR. Environs d’Ampasimentera (Boina),
paturages, fl. verdatres, October 1906, Perrier 10946 (P00573466! holo.).
Rhodocodon intermedius H.Perrier ex Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 120 (2015) ≡ R. intermedius H.Perrier,
Fl. Madagasc. Fam. 40: 116 (1938), nom. inval. ≡ Drimia intermedia (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3):
556 (2004), nom. inval. (Figs 3.16, 41.2). Type:—MADAGASCAR. Plaine sèche-alluvionnaire, sur la rive droite de la Mananjeba
(Andavakoera), July 1963, Perrier 10965 (P00573461! holo.; P00573462! iso.).
Rhodocodon jackyi Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 253(3): 202 (2016) (Figs 3.17, 41.3). Type:—MADAGASCAR.
Province Toliara, region Anosy: near the village Vatambe, elev. 330 m, ex hort. 30 April 2014, W. Knirsch, A. Sieder & J. Andriantiana
6620 corresponding to WW04669 (GZU-000324829! holo.; P!, PBZT! iso.).
Rhodocodon linearifolius Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 122 (2015) ≡ R. calcicola var. linearifolius H.Perrier,
Fl. Madagasc. Fam. 40: 116 (1938), nom. inval. (Fig. 39.2). Type:—MADAGASCAR. Rocailles denudées—sur les labradorites de
l’Antanimena, December 1924, Perrier 16824 (P00516239! holo.; P00516238! iso.).
Rhodocodon mascarenensis (Baker) Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 122 (2015) ≡ Urginea mascarenensis
Baker in J. Bot. 12: 363 (1874), basionym ≡ Drimia mascarenensis (Baker) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3):
557 (2004) (Figs 3.18, 41.4). Type:—MADAGASCAR. Bojer & Hilsenberg s.n. (BM000797876! holo.).
= R. madagascariensis Baker in J. Linn. Soc., Bot. 18: 280, t. 8 (1881) (Fig. 39.2). Type:—MADAGASCAR. N & E of Ankaratra Mts,
s.d., Langley Kitching s.n. (K000099861! lecto. designated by Knirsch et al. 2015).
Rhodocodon monophyllus Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 125 (2015) ≡ R. calcicola var. monophyllus H.Perrier,
Fl. Madagasc. Fam. 40: 115 (1938), nom. inval. (Figs 3.19, 38.2, 41.5). Type:—MADAGASCAR. Bois sur les rocailles gneissiques,
sur la rivière Besafotra, affluent de droite du Menavava, Boina, bassin de la Betsiboka, Novembre 1899, Perrier 990 (P02160895!
holo.).
Rhodocodon perrieri Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 414(3): 138 (2019) (Figs 3.20, 42.1). Type:—MADAGASCAR.
Antananarivo Province: region Analamanga, district Manjakandriana, mountains ca. 4 km SE of Ambohimiadana, elev. 1500‒1600
m, ex hort. 30 April 2014, W. Knirsch, A. Sieder & J. Andriantiana AS6564 (ABH! holo.).
Rhodocodon petrae Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 414(3): 130 (2019) (Fig. 42.2). Type:—MADAGASCAR. Mahajanga
Province: Namoroka National Park, near the Tsingy, elev. 120 m, ex hort. 15 May 2013, J. Andriantiana s.n. corresponding to
WW04826 (ABH82488! holo.).
Rhodocodon rotundus H.Perrier ex Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 195(2): 127 (2015) ≡ R. rotundus H.Perrier, Fl.
Madagasc. Fam. 40: 120 (1938), nom. inval. ≡ Drimia rotunda (H.Perrier) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557
(2004), nom. inval. (Figs 4.1, 38.2). Type:—MADAGASCAR. Versant W de l’Isalo, grès, elev. 400 m, October 1924, Perrier 16583
(P00573467! holo.).
Rhodocodon rubescens Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 414(3): 136 (2019) (Fig. 42.3). Type:—MADAGASCAR.
Mahajanga Province: district of Sofia, region Antsohihy, ca. 6 km SW of town Anjiamangirana, elev. 320 m, ex hort. 10 April 2016,
W. Knirsch, A. Sieder & J. Andriantiana s.n. (ABH! holo.).
Rhodocodon siederi Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 414(3): 141 (2019) (Figs 4.2, 42.4). Type:—MADAGASCAR.
Antananarivo Province: ca. 40 km west of Antananarivo, Antongona, Imeritsiatosika, rocky mountains 4 km E of Arivonimamo, elev.
1497 m, growing among grasses, ex hort. 30 April 2014, W. Knirsch, A. Sieder & J. Andriantiana AS6728 (ABH82498! holo.).
Rhodocodon urgineoides Baker in Hooker’s Icon. Pl. 25: t. 2492 (1896) ≡ Rhadamanthus urgineoides (Baker) Speta in Phyton (Horn,
Austria) 38: 77 (1998) ≡ Drimia urgineoides (Baker) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004) (Figs 4.3,
39.2, 42.5). Type:—MADAGASCAR. Ivohimanitra forest, October 1894, C.H. Forsyth Major 44 (G00168520! lecto. designated by
Knirsch et al. 2015; K000099856! isolecto.).
Rhodocodon viridans Knirsch, Mart.-Azorín & Wetschnig in Phytotaxa 414(3): 133 (2019). Type:—MADAGASCAR. Antsiranana
Province: ca. 20 km N of Vohemar, Ambatoharanana, Cap Manambato, ex hort. in Austria, Steiermark, Sankt Marein on 15 May
2015, W. Röösli & R. Hoffmann s.n. corresponding to WW04675 (ABH82494! holo.).
17. Sagittanthera Mart.-Azorín, M.B.Crespo, A.P.Dold & Van Jaarsv. in Phytotaxa 98(2): 46 (2013) (Fig. 43). Typus
generis:—S. cyanelloides (Baker) Mart.-Azorín, M.B.Crespo, A.P.Dold & Van Jaarsv. (holotype).
≡ Drimia sect. Sagittanthera (Mart.-Azorín, M.B.Crespo, A.P.Dold & Van Jaarsv.) J.C.Manning & Goldblatt in Strelitzia 40: 73 (2018).
Typus sectionis:—D. cyanelloides (Baker) J.C.Manning & Goldblatt (holotype).
Description:—Bulbous geophyte. Bulb partially epigeal, clump forming, with thickened, fleshy, shortly stalked, loose
scales, easily proliferating when detached, producing dense threadlike slimy structures when broken. Roots thickened
and branched. Leaves hysteranthous, up to 30 cm long, 3‒8 mm wide, glabrous, linear, adaxial leaf surface subflattened,
grooved, abaxial leaf surface strongly keeled, fleshy and trapezoid in cross-section. Inflorescence a long, lax, many-
flowered raceme, 5‒10 cm long, with 5‒25 flowers; peduncle suberect to leaning, 20‒30 cm long, green, glabrous;
pedicels 7‒12 mm long, curving downwards. Bracts acute, triangular-subsagittate, 3.5‒6.0 mm long, lowermost with
long basal spur, 4‒6 mm long; bracteoles present, 0.3‒1.2 mm long. Flowers stellate, nodding, lasting 2‒3 days. Tepals
6, biseriate, 7‒9 mm long, free from base, subequal, oblong to oblong-lanceolate, white with dull purplish stripes.

Stamens 6, erect and connivent to ovary and style; filaments free, short, straight, erect, flattened at base, ca. 1.5 mm
long, much shorter than anthers; anthers large, 5−6 mm long, sagittate, dehiscing by minute apical pores, connate to
form conical structure that surrounds style. Ovary ovoid, 2.5−3.0 mm long, green. Style narrow, erect, elongate, 3−4
mm long, straight, included in cone-like structure formed by anthers at beginning of anthesis, elongating to overtop
anthers after their dehiscence. Stigma punctiform to subcapitate. Capsule and seeds not studied.
FIGURE 43. Sagittanthera cyanelloides (Baker) Mart.-Azorín et al.
Number of species and distribution:—Monotypic genus, occurring in the eastern parts of the Eastern Cape
Province of South Africa between latitudes 27°46’ and 29°46’S above 1000 m at Nqanqarhu (Maclear) in the foothills
of the Southern Drakensberg mountains (Fig. 29). The genus is restricted to the Uzambara-Zululand Region (sensu
Takhtajan 1986). For further information on Sagittanthera see Baker (1897), Van Jaarsveld & Van Wyk (2005), and
Martínez-Azorín et al. (2013a).
Studied material:—SOUTH AFRICA. Eastern Cape Province. Stutterheim (3227): Komga, prope Prospect farm,
(–DB), elev. 695 m, December 1889, Flanagan 573 (K000257233!, BOL!, PRE0051093-0!); Stutterheim (3227),
Boogkrans, Kei River, (–BC), elev. 598 m, 12 April 2013, Van Jaarsveld & Harrower 24767 (PRE); Stutterheim
(3227), Kliprooiysterhoutdraai, Kei River, (–BC), elev. 318 m, 15 April 2013, Van Jaarsveld & Harrower 24794 (PRE);
Umtata (3128): Maclear, The Falls, ca. 14 km NE from Maclear, (–AB), forest scrub, elev. 1020 m, 12 November 1994,
S.P. Bester 3300 (K!, PRE835412!); Butterworth (3228): Transkei, Willowvale distr., Ngqaqini admin. gebied, steil
helling, SO front, laag in vallei, (–AD), November 1983, J.A. van Eeden B386 (PRE666067!); Port St. Johns (3129),
lower Mzimvubu River, south facing, shale cliffs below Ludonga, (–AD), Van Jaarsveld, Xaba, Harrower & Zwide 97
(PRE); Port St. Johns (3129): Transkei, Mateku waterfall, (–BD), T.M.S. cliffs, grassland, 11 November 1970, R.G.
Strey 10170 (PRE0051092!); Port St. Johns (3129): Transkei, Libode district, confluence of the Tina and Tsitsa rivers,
east of Umtata, (–CB), elev. 699 m, W side overlooking the confluence of the Tina and Tsitsa Rivers, February 2002,
flowered ex hort. December 2007, A.P. Dold s.n. (GRA!).
History, diagnostic characters, and taxonomic relationships:—Rhadamanthus cyanelloides was described by
Baker (1897) from material collected in the eastern regions of South Africa and showing a very distinct morphology,
such as the tricuspidate bracts (considering the blade, the spur and the bracteole), free tepals, and most notably “stamens
like those of Cyanella, […] with very short filaments and six, large cylindrical anthers that are permanently connivent in
a cone and dehiscing by apical pores”. He considered that the taxon most likely represented a new genus. Van Jaarsveld
& van Wyk (2005) subsequently described Drimia cremnophila Van Jaarsv. in Van Jaarsveld & Van Wyk (2005: 81)
and D. mzimvubuensis from the Mzimvubu River in the Eastern Cape Province of South Africa. Martínez-Azorín et al.
(2013a) described Sagittanthera based on the distinct flower morphology of the above-mentioned taxa and accepted
two species: S. cyanelloides (including D. cremnophila as a synonym) and S. mzimvubuensis, which in general share
similar flower and inflorescence morphologies, and biogeography. This solution was adopted based on their unique
flower morphology within Urgineoideae, particularly the connate anthers in S. cyanelloides and the connate filaments
in S. mzimvubuensis, among other characters. However, our phylogenetic studies revealed that both species, represent
distant and independent lineages within Urgineoideae (Martínez-Azorín et al. 2023a)—a fact supported by differences
in the connation of anthers, connation of filaments, presence of bracteoles, and leaf morphology. These characters led
Martínez-Azorín et al. (2017) to segregate D. mzimvubuensis as the monotypic Aulostemon, a genus accepted in the
present work. Manning & Goldblatt (2018) placed the latter species in D. sect. Aulostemon (Martínez-Azorín et al.
2017: 288) Manning & Goldblatt (2018: 123). Aulostemon is readily differentiated from Sagittanthera by its filaments
forming a long tube above perigone, and free anthers, among other characters (Martínez-Azorín et al. 2017). The
phylogenetic findings of Martínez-Azorín et al. (2023a) place samples of Sagittanthera cyanelloides as sister to a
clade combining Urginavia, Zingela, Thuranthos, and Ledurgia. Combining those groups in a single genus would be
highly disruptive, since all genera show very different flower morphologies. We accordingly accept Sagittanthera as
monotypic to include S. cyanelloides.
Sagittanthera cyanelloides (Baker) Mart.-Azorín, M.B.Crespo, A.P.Dold & van Jaarsv. in Phytotaxa 98(2): 48 (2013) ≡ Rhadamanthus
cyanelloides Baker, Fl. Cap. (Harvey) 6(3): 444 (1897), basionym ≡ Drimia cyanelloides (Baker) J.C.Manning & Goldblatt in
Strelitzia 9: 711 (2000) (Fig. 43). Type:—SOUTH AFRICA. Eastern Cape: Komgha, grassy valleys near Prospect Farm, elev. 2100
feet, Flanagan 573 (K000257233! lecto., designated as “holo.” by Manning & Goldblatt (2018: 74); PRE0051093-0!, BOL140334!
isolecto.).
= Drimia cremnophila van Jaarsv. in Aloe 42(4): 81 (2005). Type:—SOUTH AFRICA. Eastern Cape. Port St Johns (3129): lower
Mzimvubu River, shale cliffs below Ludonga, (–AD), Van Jaarsveld, Xaba, Harrower & Zwide 97 (PRE holo.).
18. Schizobasis Baker in J. Bot. 11: 105 (1873) (Figs 44–46). Typus generis:—S. macowanii Baker (holotype).
≡ Drimia sect. Schizobasis (Baker) J.C.Manning & Goldblatt in Strelitzia 40: 142 (2018). Typus sectionis:—S. macowanii Baker
(holotype).
Description:—Small bulbous geophyte. Bulb hypogeal or epigeal, solitary or clump forming, ovoid to globose, up to
7 cm in diam., scales compact, white to pinkish. Roots thickened and branched. Leaves 1‒2, only present in immature
plants, filiform, green, smooth, glabrous. Inflorescence distinctly branched, paniculate, 5‒30 cm long, erect to prostrate,
sometimes twining and climbing on vegetation or rocks, green, commonly long lasting and photosynthetic, smooth
or papillose. Peduncle green to purplish, 5‒10 cm long, terete, erect and straight to prostrate or twining, sometimes
flexuose at base, glabrous or minutely papillose. Pedicels 4‒10 mm long, smooth to papillose, straight to sinuous and
twining, green and long lasting supporting ripe capsules. Bracts ovate lanceolate, green, lowermost with short spur.
Bracteoles absent. Flowers stellate to campanulate, erect to nodding, diurnal. Tepals 6, biseriate, lanceolate to ovate,
2.5‒5.0 mm long, white to yellowish, with darker longitudinal band, free or very shortly connate at base, suberect,
patent or reflexed. Stamens 6, incurved and connivent to style or spreading; filaments filiform, 2‒4 mm long, straight
to sigmoid; anthers oblong to sagittate, with apiculate connective, dehiscing longitudinally along whole length. Ovary
ovoid to subglobose, green or yellowish; style erect, white, straight or sigmoid, as long as or longer than ovary; stigma
minutely trigonous. Capsules ovate to subglobose, 3‒5 mm long, trigonous, loculicidal, the 3 valves splitting to base,
with withered perigone segments circumscissile below and forming an apical cap. Seeds ellipsoid to subfusiform,
1.8‒3.0 mm long, flattened with prominent central embryo and short wings, black, glossy, with somewhat sinuous
anticlinal testa cell walls.

FIGURE 44. Species of Schizobasis Baker displayed in horizontal rows of images. 1. Schizobasis cuscutoides (Burch. ex Baker) Benth.
& Hook.f.; 2. Schizobasis intricata (Baker) Baker; 3. Schizobasis macowanii Baker.
Number of species and distribution:—Schizobasis includes eleven species, occurring mainly in Southern Africa,
and extending to Angola and East Africa (Fig. 29). In terms biogeographic phytochoria, it is restricted to the Cape,
Karoo-Namib and Uzambara-Zululand Regions and the Zambezian Subregion (sensu Takhtajan 1986 and Martínez-
Azorín et al. 2023a). For further information on Schizobasis see Manning & Goldblatt (2018), whilst noting that we
accept most species described in the genus, based on their morphological differences. Our field work has confirmed
large variation in flower morphology and habit in Schizobasis (Figs 44, 46). Further studies based on living material
are urgently required to clarify the taxonomy of this genus.
Karyology:—2n=18 (Bruyns & Vosa 1987, as S. intricata); 2n=20 (Watters & Ornduff 1985, as Schizobasis
intricata Baker; Jones & Smith 1967, as Schizobasis sp.).
History, diagnostic characters, and taxonomic relationships:—Baker (1873a) described Schizobasis, a very
distinct genus characterised by the slender, wiry, flexuose, branched inflorescence, also commenting on its similarity
with Bowiea and other genera. Schizobasis has been accepted by most researchers in Urgineoideae based on its unique
syndrome of morphological characters (Baker 1873b, 1874a, 1876, 1898, Bentham & Hooker 1883, Merxmüller 1970,
Jessop 1977, Stedje 1997, Speta 1998a, 1998b, Pfosser & Speta 2001, Demissew & Nordal 2010, Martínez-Azorín
et al. 2015). However, other recent studies merge Schizobasis into a widely-conceived Drimia sensu lato (Golblatt &
Manning 2000, Manning et al. 2004, 2014, Manning & Goldblatt 2018), but accept Bowiea with similarly branched
inflorescence as an independent genus based on their phylogenetic relationships.
FIGURE 45. Species of Schizobasis Baker from Krause (1930: 290). A−B. Schizobasis intricata (Baker) Baker; C−E. Schizobasis
cuscutoides (Burch. ex Baker) Benth. & Hook.f.
FIGURE 46. Species of Schizobasis Baker displayed in horizontal rows of images. 1. Schizobasis schizobasoides (J.C.Manning &
J.M.J.Deacon) Mart.-Azorín et al.; 2. Schizobasis schlechteri Baker; 3. Schizobasis sigmoidea (J.C.Manning & J.M.J.Deacon) Mart.-
Azorín et al.

The phylogenetic studies of Pfosser & Speta (2001, 2004), Manning et al. (2004), and Pfosser et al. (2012) have
shown that Schizobasis is sister to Litanthus. The phylogenetic analyses of Martínez-Azorín et al. (2023a) agree with
the previous molecular studies and place eleven samples of Schizobasis in a monophyletic clade sister to Litanthus,
the latter relationship being supported by anther and seed morphology. Therefore, we maintain here Schizobasis at
genus rank as it is easily identifiable by the branched and wiry inflorescences, flowers with tepals only shortly fused at
base, apiculate stamen connective, and angled seeds. We here accept all described species in the genus based on their
morphological differences although a detailed taxonomic revision of the genus is needed to ascertain the real diversity
of the genus.
Schizobasis angolensis Baker in Trans. Linn. Soc. London, Bot. 1(5): 255 (1878). Type:—ANGOLA. Pungo Andongo, ad rupes ipsius
Praesidii, 2400–3800 ft., August 1857, F. Welwitsch 3867 (LISU 222125! holo.; as “Adenoteca aphylla Welw. gen. nov.” in sched.).
Note:—The collection “Pungo Andongo, 2400–3800 ft., F. Welwitsch 3866 (K000257004!, LISU222126!)” is most probably original
material of that name.
= Adenoteca aphylla Welw., nom. nud. in sched. et in protolog.
Schizobasis buchubergensis Dinter in Repert. Spec. Nov. Regni Veg. 30: 80 (1932). Type:—NAMIBIA. Bogenfels (2715): Buchuberge,
(–DD), 7 July 1929, Dinter 6493 (B, holo., not seen; on loan to D. Müller-Doblies since 1980, R. Vogt pers. comm.).
Schizobasis cuscutoides (Burch. ex Baker) Benth. & Hook.f., Gen. Pl. [Bentham & Hooker f.] 3(2): 786 (1883) ≡ Asparagus cuscutoides
Burch. ex Baker in J. Linn. Soc., Bot. 14: 606 (1875), basionym ≡ Drimia cuscutoides (Burch. ex Baker) J.C.Manning & Goldblatt
in Strelitzia 9: 711 (2000) (Figs 44.1, 45C–E). Type:—SOUTH AFRICA. Northern Cape: Caput Bonae Spei, in saxosis aridis ad
ripas fluminis Gariep, 1 March 1813, Burchell 2673 (K000257125!, lecto. designated as holo. by Manning et al. in S. African J. Bot.
94: 264. 2014; L, P01775622!, isolecto.).
Schizobasis dinteri K.Krause in Bot. Jahrb. Syst. 48(3–4): 355 (1912). Type:—NAMIBIA. Mariental (2417): Gross Namaland, bei
Marienthal [Mariental] in der Kieswüste unterhalb des Staudammes, (–DB), 20 March 1911, Dinter 2005 (SAM0073821-0! lecto.
designated by Manning et al. in S. African J. Bot. 94: 264. 2014; SAM 0004463-0! isolecto.).
Schizobasis gracilis R.E.Fr. in Wiss. Ergebn. Schwed. Rhodesia-Kongo-Exped. 1911–1912(1): 227 (1916). Type:—ZAMBIA. Rhodesia
bor. orient.: pr. rivulum Kalungwisi, October 1911, R.E. Fries 1157 (UPS [V-041106]! holo.).
Schizobasis intricata (Baker) Baker in J. Bot. 12: 368 (1874) ≡ Anthericum intricatum Baker in J. Bot. 10: 140 (1872), basionym ≡
Drimia intricata (Baker) J.C.Manning & Goldblatt in Strelitzia 9: 712 (2000) (Figs 4.4, 44.2, 45A–B). Type:—SOUTH AFRICA.
Cape, Zeyher 4284 (K000257121! lecto. designated as holo. by Stedje & Thulin in Nordic J. Bot. 15: 600. 1995; SAM0022792-0
isolecto.).
Schizobasis macowanii Baker in J. Bot. 11: 105 (1873) (Figs 4.5, 44.3). Type:—SOUTH AFRICA. Eastern Cape. Somerset East (3225):
inter frutices prope Somerset East, (–DA), elev. 2500–3000 ft., October, MacOwan 1847 (K000257123! lecto. designated by Manning
et al. in S. African J. Bot. 94: 264. 2014; BOL!, SAM 0022788-0 isolecto.).
Schizobasis schizobasoides (J.C.Manning & J.M.J.Deacon) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 294
(2019) ≡ Drimia schizobasoides J.C.Manning & J.M.J.Deacon in Strelitzia 40: 142 (2018), basionym (Fig. 46.1). Type:—SOUTH
AFRICA. Northern Cape. Springbok (2917): Karrachabpoort, (–AC), 23 December 2015, J.M.J. Deacon 3915 (NBG holo.).
Schizobasis schlechteri Baker in Bull. Herb. Boissier ser. 2, 1: 783 (1901) (Fig. 46.2). Type:—SOUTH AFRICA. Eastern Cape. Umtata
(3128): Südwest-Afrika (Natal) [sic.], regio orientalis prope Umtata, (–DB), elev. 3400 ft., 18 January 1895, R. Schlechter 6327
(Z000086084! lecto. designated by Manning et al. in S. African J. Bot. 94: 264. 2014; GRA!, BOL!, PRE! isolecto.).
Schizobasis sigmoidea (J.C.Manning & J.M.J.Deacon) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 201(2): 168 (2015)
≡ Drimia sigmoidea J.C.Manning & J.M.J.Deacon in S. African J. Bot. 94: 267 (2014), basionym (Figs 4.6, 46.3). Type:—SOUTH
AFRICA. Western Cape. Montagu (3320): Ashton, Wolwendrift, cliff face in large kloof, (–CC), 23 September 2013, J.M.J. Deacon,
D. Gwynne-Evans & R. McKenzie 2820 (NBG holo.; MO iso.).
= Schizobasis bruce-bayeri U. & D. Müll.-Doblies, nom. nud. in sched.: SOUTH AFRICA. Swellendam, Bonnievale, R. Marloth 11997
(NBG!).
Schizobasis visagieae (van Jaarsv.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. & stat. nov. ≡ Drimia intricata (Baker) J.C.Manning
& Goldblatt var. visagieae Van Jaarsv. in Bradleya 32: 78 (2014), basionym. Type:—ANGOLA. 1413 (Lubango): Cliffs at Tundavala,
west of Lubango, (–DC), 14 July 2010, Van Jaarsveld & Harrower 23119 (LUB holo.; NBG iso.).
19. Sekanama Speta in Stapfia 75: 168 (2001) (Figs 47–48). Typus generis:—S. sanguinea (Schinz) Speta
(holotype).
Description:—Small to medium-sized bulbous geophyte. Bulb hypogeal, ovoid to subglobose, solitary, to 9 cm

in diam., outer scales membranous and papery, inner scales red, compact. Roots thickened and branched. Leaves
2‒9 per bulb, mostly hysteranthous, 10‒40 cm long, suberect, narrowly lanceolate, canaliculated, green, leathery,
glabrous, acute. Inflorescence racemose, from shortly racemose to elongated and multiflowered, 2‒60 cm long, with
25‒90 flowers; peduncle above ground level usually much shorter than inflorescence, erect, terete, glabrous, smooth;
pedicels 4‒14 mm long, suberect to patent. Bracts oblong-obovate, navicular, lately caducous, 1‒3 mm long, with
lanceolate-oblong spur, 0.5‒1.0 mm long; bracteoles present and small. Flowers shortly campanulate at base with
spreading-patent upper 2/3 of tepals, erect-patent, diurnal. Tepals 6, biseriate, 7‒11 mm long, white with brownish or
greenish longitudinal band more evident on abaxial side, mostly free, shortly connate at base to 2 mm long, papery
and translucent when withered. Stamens 6, suberect or spreading, not connivent to style; filaments linear-lanceolate,
3‒7 mm long, white, adnate to perigone at base; anthers medifixed, oblong. Ovary ovoid, 2.0‒4.5 mm long, trigonous;
style filiform, erect, 2.5‒4.0 mm long, white; stigma inconspicuously 3-lobed, papillate. Capsule ellipsoid to ovoid,
10‒18 mm long, valves completely dehiscing from base, withered tepals mostly persisting at base of dehisced capsule.
Seeds ellipsoidal, 5‒10 mm long, flattened with prominent central embryo and wide, usually curved wings, testa black,
glossy, with sinuous anticlinal cell walls.
FIGURE 47. Sekanama sanguinea (Schinz) Speta from: 1. Pole Evans (1924: t. 138, as Urginea burkei Baker); 2. Stent & Curson (1922,
as Urginea burkei Baker).

FIGURE 48. Species of Sekanama Speta displayed in horizontal rows of images. 1−2. Sekanama sanguinea (Schinz) Speta; 3. Sekanama
johnstonii (Baker) Mart.-Azorín et al. (Photographs by T. Harvey from Kunene River in Angola).
Number of species and distribution:—Sekanama includes four species, occurring mainly in the inland regions
of Southern Africa and extending to Tanzania and Socotra (Fig. 49). They are restricted to the Southern and Eastern
Sections of the Zambezian Subregion and the Erithreo-Arabian Subregion (sensu Takhtajan 1986 and Martínez-
Azorín et al. 2023a). For further information in Sekanama see Jessop (1977), Speta (2001), and Manning & Goldblatt
(2018).
Karyology:—2n=20 (De Wet 1957, Jones & Smith 1967, as Urginea burkei Baker).
History, diagnostic characters, and taxonomic relationships:—Speta (2001) described Sekanama to include
Urginea sanguinea Schinz (1890: 219), U. burkei Baker (1897: 469), and U. delagoensis Baker (1897: 467). This genus
was characterised as having red bulbs with imbricate scales; proteranthous, straight leaves; racemose inflorescence
with up to 70 flowers; shortly spurred bracts; pedicels 4‒9 mm long; and tepals 8‒10 mm long. The type of the genus,
S. sanguinea, is one of the most poisonous plants in Africa due to its bufadienolides (Krenn et al. 1993, Speta 2001).
Urginea burkei has been treated as a synonym of U. sanguinea, and on the other hand U. lydenburgensis Dyer (1942b:
t. 859) has been treated as a synonym of U. delagoensis (Jessop 1977, Manning & Goldblatt 2018). In recent years,
Drimia edwardsii Crouch & Martínez-Azorín (2015: 137), which shares flower and fruit morphological traits with U.
delagoensis and U. lydenburgensis, was described from central KwaZulu-Natal (Crouch & Martínez-Azorín 2015).
The phylogenetic studies of Martínez-Azorín et al. (2023a) recovered three samples of S. sanguinea as a strongly
supported clade. Another strongly supported clade combines samples of U. delagoensis, U. lydenburgensis, and D.
edwardsii. These latter two clades are sometimes related as sister, although with very low support, whereas other
analyses maintain them separate within general collapsed relationships. Important differences in morphology exist
between S. sanguinea (including U. burkei) and the remaining taxa in this group. Sekanama sanguinea is distributed
in the inland regions of Southern Africa and is characterised by hysteranthous leaves, racemose inflorescences with
peduncle much shorter than the raceme, white flowers, present bracteoles, withered tepals persisting at the base of

the ovoid-ellipsoid capsule, and flat and wide, elliptic seeds. Conversely, U. delagoensis, U. lydenburgensis, and D.
edwardsii are collectively distributed in eSwatini (formerly Swaziland), and in the adjacent South African provinces of
KwaZulu-Natal and Mpumalanga. In terms of biochoria, they are restricted to the Uzambara-Zululand Region (sensu
Takhtajan 1986). These latter three species have synanthous leaves, inflorescences with peduncles much longer than
the raceme, absent bracteoles, pale brown or carneous to greenish flowers, withered tepals persisting at the top of the
capsule, narrowly ellipsoid to fusiform capsules, and narrowly lanceolate-fusiform seeds.
The taxonomic revision by Manning & Goldblatt (2018) placed U. sanguinea in their D. sect. Macrocentrae
J.C.Manning & Goldblatt, together with species such as Urginea macrocentra (= Boosia sensu Speta 2001), without
taking cognisance of the phylogenetic work of Pfosser & Speta (2001, 2004), where these taxa nested in different and
distant clades in the phylogeny, confirmed by the phylogenetic results of Martínez-Azorín et al. (2023a). Chromosome
counts on Sekanama sanguinea (as Urginea burkei) (De Wet 1957, Jones & Smith 1967) found 2n=20, whilst the count
on Urginea lydenburgensis R.A.Dyer by De Wet (1957) showed 2n=32, indicating clear chromosomal differences
between Sekanama s.str. and Zulusia.
Based on our current data, we include U. delagoensis, U. lydenburgensis, and D. edwardsii in the new genus
Zulusia Mart.-Azorín et al. Albeit with limited phylogenetic evidence, we include in Sekanama, Sekanama sanguinea
(syn. S. burkei) together with the endemic Tanzanian species Urginea brachystachys Baker (1892: 474), Urginea
johnstonii Baker (1898: 539) described from southern Angola, and the endemic Socotran Urginea porphyrostachys
Baker ex Balfour (1884: 411). This is based on their shared red and fleshy bulb scales, small and navicular bracts,
presence of bracteoles, and the papery-translucent withered tepals mostly remaining at the base of developing capsules.
Further studies are necessary to confirm species delimitations and genus circumscription.
Sekanama brachystachys (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea brachystachys Baker in Bot.
Jahrb. Syst. 15(4): 474 (1892), basionym ≡ Drimia brachystachys (Baker) Stedje in Nordic J. Bot. 7(6): 665 (1987) ≡ Urginavia
brachystachys (Baker) Speta in Phyton (Horn, Austria) 38(1): 87 (1998) (Fig. 4.7). Type:—TANZANIA. Musoma District, mile 5
from the Simiyu river, N.E. to the Eastern Boundary, S. Extension, elev. 5000 ft., 27 September 196?. P.J. Greenway 10211 (EA neo.;
PRE!, K! iso.).
Sekanama johnstonii (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea johnstonii Baker in Fl. Trop. Afr.
[Oliver et al.] 7(3): 539 (1898), basionym (Fig. 48.3). Type:—ANGOLA. Near the River Cunene, September 1893, Johnston s.n.
(K000400576! holo.).
Sekanama porphyrostachys (Baker ex Balf.f.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea porphyrostachys Baker
ex Balf.f. in Proc. Roy. Soc. Edinburgh 12: 411 (1884), basionym ≡ Drimia porphyrostachys (Baker ex Balf.f.) A.G.Mill., Ethnofl.
Soqotra Archipelago: 726 (385) (2004), nom. inval. (Fig. 4.8). Type:—YEMEN. Socotra, inter Kischen, Pankoke, elev. 1000 ft., 02
May 1881, G. Schweinfurth 678 (K001044815! holo.).
Sekanama sanguinea (Schinz) Speta in Stapfia 75: 168 (2001) ≡ Urginea sanguinea Schinz in Verh. Bot. Vereins Prov. Brandenburg 31:
219 (1890), basionym ≡ Drimia sanguinea (Schinz) Jessop in J. S. African Bot. 43(4): 293 (1977) (Figs 4.9, 47, 48.1–2). Type:—
NAMIBIA. Gunib (2028): Osnambonde [Omambonde] in Nord-Hereroland, (–AA), October 1885, H. Schinz 25 (Z first-step lecto.
designated as “holo.” by Manning & Goldblatt in Strelitzia 40: 19. 2018; second-step lecto. designated here:—Z000087336! lecto.;
Z000027553!, K000257377! isolecto.). Note:—A second step lectotypification is needed due to the presence of two herbarium
vouchers with different barcode numbers under the same collection.
= Urginea burkei Baker, Fl. Cap. (Harvey) 6: 469 (1897) ≡ Sekanama burkei (Baker) Speta in Stapfia 75: 168 (2001). Type:—SOUTH
AFRICA. North West. Rustenburg (2527): Magaliesberg, (–CA, CC, DB, DC), September [without year], Burke s.n. (K000257374!
lecto. designated by Jessop in J. S. African Bot. 43: 293. 1977; BM000911791! isolecto.).
= Urginea rautanenii Baker in Bull. Herb. Boissier ser. 2, 3: 664 (1903). Type:—NAMIBIA. Ondangua (1715): Amboland, Ondonga
[Ondangwa], (–DD), 11 November 1895, M. Rautanen 171 (Z000027552! lecto. designated here; Z000027558! isolecto.).
20. Spirophyllos Mart.-Azorín, M.B.Crespo & M.Á.Alonso gen. nov. (Figs 50–51).
Typus generis:—S. noctiflorus (Batt. & Trab.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso ≡ Urginea noctiflora Batt. & Trab. (holotype).
Diagnosis:—Genus distinctissimus singulari characterum combinatione a ceteris generibus Urgineoideae diversus et facile distinguendus,
nempe foliis anguste linearibus, spiraliter contortis; floribus nocturnis nutantibus, staminum filamentis rectis a basi ovario styloque
conniventibus sed supra medium implexis divergentibusque; et capsulis magnis, ovatis vel subglobosis, patentibus.
FIGURE 50. Spirophyllos noctiflorus (Batt. & Trab.) Mart.-Azorín et al. Illustration kept at MPU (MPU225773) as Urginea noctiflora
Batt. & Trab. Reproduced with permission: copyright © Université de Montpellier – Herbier MPU (SPH).

FIGURE 51. Spirophyllos noctiflorus (Batt. & Trab.) Mart.-Azorín et al. in cultivation from Imaoun, Morocco. a. Developing inflorescence;
b. Inflorescence; c. Flower in lateral view; d. Flower in frontal view; e. Flower in dorsal view; f. Flower in lateral view with two removed
tepals showing stamens disposition; g. Dissected flower; h. Indehisced anthers; j. Inflorescence section showing bracts and pedicels; k.
Bulbs with coiled leaves from wild plants near Imaoun (Photographs by A. Garcin). Scales: a, b: 1 cm, c–g, j: 5 mm, h: 2 mm.
Description:—Bulbous geophyte. Bulb hypogeal, ovoid, to 5 cm in diam., with white inner scales and greyish,
membranous outer tunics. Roots thickened and branched. Leaves 5‒15 per bulb, hysteranthous, 10‒20 cm long,
narrowly linear, hemiterete to slightly flattened, distinctly coiled at least along upper half, green, somewhat glaucous,
smooth, glabrous. Inflorescence a lax, erect raceme, 10‒20 cm long, with 4–25 flowers; peduncle 10‒20 cm long, erect,
terete, smooth, dull grey-green to reddish-brown; pedicels 6‒20 mm long, arching downwards at anthesis, subpatent
and sigmoid in fruit. Bracts ovate-lanceolate, 1‒2 mm long, lowermost with distinct spur as long as or longer than
blade, caducous; bracteoles absent. Flowers stellate with strongly reflexed tepals at full anthesis, nodding, nocturnal,
opening in late evening and closing before sunrise, fragrant during night. Tepals 6, biseriate, 7‒10 mm long, narrowly
lanceolate, pale brown to carneous colour, with darker brownish-greenish longitudinal band more evident on abaxial
side, almost free. Stamens 6, erect, straight or very slightly curved, connivent to style and crossing each along upper
third; filaments linear, only slightly widened at base, narrowly conical and tapering, 6‒7 mm long, white or cream
colour, adnate to tepals at base; anthers oblong, 1‒2 mm long, medifixed to subbasifixed, deep orange after dehiscence,
connective with a pit at filaments insertion, with yellow pollen. Ovary ovate, 4‒5 mm long, 6-lobed basally, pale
yellow. Style 5‒6 mm long, suberect, white, filiform, ending in minute stigma. Capsule ovate to subglobose, trigonous,
obtuse, 7‒10 mm long, pale brown. Seeds ellipsoidal, flattened, 6‒8 mm long, with black, glossy testa.
Number of species and distribution:—This new genus currently includes one species restricted to desert regions
of northwestern Africa (southern Morocco and Algeria), south of the Atlas Mountain range (Fig. 49). Therefore, it is
restricted to the Saharo-Arabian Region (sensu Takhtajan 1986).
Karyology:—2n=20 (Azizi et al. 2016).
History, diagnostic characters, and taxonomic relationships:—Urginea noctiflora Battandier & Trabut
(1894: 505) was described from the border of the Sahara Desert in Algeria showing a number of clear morphological
differences with regard to all known European species of Urgineoideae, such as the narrowly linear, distinctly coiled
leaves, nocturnal nodding flowers with the filaments straight, connivent to the style and crossing along the middle, and
the patent, large, ovate capsules (Battandier & Trabut 1894).
The phylogenetic study by Martínez-Azorín et al. (2021) included six samples of different populations of Urginea
noctiflora from Morocco provided by A. Garcin constituting a strongly supported clade, which is sister to another
strongly supported clade merging four samples of Urginea fugax (Moris 1827: 46) Steinheil (1834: 328) and U.
ollivieri Maire (1938: 453) from Sardinia, Ibiza, and Morocco (corresponding to Urginea s.str. in the sense of the
present work). This same relationship was already shown by Pfosser et al. (2006). However, Urginea noctiflora differs
from typical Urginea species in the distinctly coiled leaves, a character not known in any other Urgineoideae, and the
reproductive characters listed above. We here propose the new genus Spirophyllos to include Urginea noctiflora based
on the distinct morphological differences described above and its genetic divergence.
Spirophyllos noctiflorus (Batt. & Trab.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea noctiflora Batt. & Trab. in
Compt.-Rend. Assoc. Franc. Besancon 1893(2): 505 (1894), basionym ≡ Urginea noctiflora Batt. & Trab. subsp. helicophylla
(Maire) Maire in Fl. Afric. N. 5: 162 (1958), nom. illeg. ≡ Urginea noctiflora Batt. & Trab. var. helicophylla Maire in Jahand. &
Maire, Cat. Pl. Maroc 3: 870 (1934), nom. illeg. ≡ Duthiea noctiflora (Batt. & Trab.) Speta in Stapfia 75: 170 (2001) (Figs 50–51).
Type:—ALGERIA. Ouargla, Gara près de Sedrata et partout route a Mzab, 20 May 1902, L. Chevallier s.n. (P01848250! neo.
designated here).
21. Squilla Steinh. in Ann. Sci. Nat. Bot. sér. 2, 6: 276 (1836), Typus generis:—S. maritima (L.) Steinh. (lectotype,
vide Pfeiffer, Nom. 2(2): 1252. 1874).
≡ Charybdis Speta in Phyton (Horn, Austria) 38(1): 58 (1998), nom. nov. et nom. illeg. superfl. [Art. 52 of the ICN] (Figs 52–54). Typus
generis:—C. maritima (L.) Speta (holotype).
Description:—Usually tall bulbous geophyte. Bulb usually large and hypogeal, rarely somewhat epigeal, ovoid to
globose, up to 20 cm in diam. Roots thickened and branched. Leaves (4–)6‒10 per bulb, narrowly lanceolate to widely
ovate, hysteranthous, up to 35 cm long, margin entire or sometimes undulate with ciliae or minute papillae, green or
somewhat glaucous, smooth, glabrous. Inflorescence a long, erect, multiflowered raceme, with up to 1000 flowers,
6‒70 cm long; peduncle 15‒120 cm long, dull green with grey to purple tinge, terete, erect, smooth and glabrous;
pedicels 5‒35 mm long, subpatent and usually arching upwards, sometimes erect or patent at anthesis. Bracts narrowly
lanceolate, usually dry and wrinkled at flowering time, lowermost with short spur; bracteoles present and distinct.
Flowers stellate or with reflexed lobes, erect-patent, diurnal, lasting more than one day. Tepals 6, biseriate, 7‒16
FIGURE 52. Species of Squilla Steihn. 1. Squilla maritima (L.) Steinh. from Clusius (1601: 171, as Scilla hispanica); 2. Squilla maritima
(L.) Steinh. from Maire (1958: 163, as Urginea maritima); 3. Squilla maritima (L.) Steinh. from Ker Gawler (1806: t. 918, as Ornithogalum
squilla); 4. Squilla undulata (Desf.) Mart.-Azorín et al. from Jacquin (1812: pl. 30, as Drimia purpurascens J.Jacq.); 5. Squilla undulata
(Desf.) Mart.-Azorín et al. from Desfontaines (1798: t. 88, as Scilla undulata Desf.).
mm long, free or connate for less than 1‒2 mm, lanceolate-ovate to spathulate, usually white or rarely purple, with
purplish or green longitudinal band. Stamens 6, spreading; filaments smooth, usually white and lanceolate to narrowly
triangular, flattened or rarely filiform and purplish; anthers dorsifixed. Ovary ovate-oblong, usually with three apical
protuberances, green to yellow, with ca. 5‒10 ovules per locule. Style 3‒12 mm long, erect, white or rarely tinged with
purple, with trigonous stigma. Capsule ovate, 7‒19 mm long, green first and yellowish and papyraceous after ripening,
valves completely dehiscing from base, tepals cohering and inrolled above ovary after anthesis, circumcissile from
base and persisting as a cap at the top of the developing capsules. Seeds ovate-oblong to linear-oblong, 4‒12 mm long,
flattened, testa black, with sinuous anticlinal cell walls.
Number of species and distribution:—Squilla includes twelve species, occurring mostly along the Mediterranean
Basin, spanning the Canary Islands and the Middle East (Speta 1998a, Martínez-Azorín et al. 2022) (Fig. 49); it is
nearly restricted to the Mediterranean and Macaronesian Regions (sensu Takhtajan 1986). For further information on
Squilla see Pfosser & Speta (2001, 2004), Crespo et al. (2020), and Martínez-Azorín et al. (2022).
Karyology:—2n=20 (Martinoli 1954, Battaglia 1957, Bartolo et al. 1984, as Urginea undulata (Desf.) Steinh.;
Valdés-Bermejo 1980, Battaglia 1957, as D. maura Maire); 2n=28 (Bramwell et al. 1972, as Urginea hesperia Webb.
& Berth.); 2n=40 (Satô 1942, as Urginea scilla Steinh.); 2n=20, 30, 40, 50, 60 (Battaglia 1957, 1964, Valdés-Bermejo
1980, Speta 1980, Pfosser & Speta 2001, Rico 2013, Bacchetta et al. 2012); 2n=20, 40, 60, 60+1B, 60+2B (Boscaiu et
al. 2001, Fernández et al. 2001, Talavera et al. 1995, Oberprieler & Vogt 1994, as Urginea maritima).
FIGURE 53. Species of Squilla Steihn. displayed in horizontal rows of images. 1. Squilla hesperia (Webb & Berthel.) Mart.-Azorín et al.;
2. Squilla maritima (L.) Steinh.; 3. Squilla palaestina (M.B.Crespo et al.) Mart.-Azorín et al.
History, diagnostic characters, and taxonomic relationships:—Martínez-Azorín & Crespo (2016a) showed
that the Linnaean Scilla was first typified on Scilla maritima Linnaeus (1753: 308) by Rafinesque (1837: 8) and not
on Scilla bifolia Linnaeus (1753: 309) as usually accepted [vide Index Nominum Genericorum: Farr & Zijlstra (2023)
http://botany.si.edu/ing/].
Steinheil (1836) described Squilla to segregate Scilla maritima Linnaeus (1753: 308) (Urginea scilla Steinheil
1834: 321) from his previously described Urginea, implying Squilla to be different from the name Scilla. Speta (1998b)
accepted separation of both Urginea Steinh. and Squilla, but proposed the new name Charybdis, with C. maritima
(Linnaeus 1753: 38) Speta (1998b: 60) as type, to replace Squilla since he considered the latter to be an orthographic
variant of both Scilla and Skilla Rafinesque (1837: 13), and therefore a later illegitimate homonym. However, based
on the typification by Rafinesque (1837) of Scilla maritima, Charybdis is illegitimate under Art. 52 of the Shenzhen
Code as it was superfluous when published, a fact that makes Charybdis unavailable for use.
Martínez-Azorín & Crespo (2016b) requested the Nomenclature Committee for Vascular Plants (NCVP) for
a binding decision on whether Scilla L. (subfam. Hyacinthoideae) and Squilla Steinh. (subfam. Urgineoideae ≡
Asparagaceae subfam. Scilloideae trib. Urgineeae) are sufficiently alike to be considered orthographic variants and
to be confused. After several attempts, a sufficient number of the Committee Members voted recently in favour of a
binding decision that both names are not confusable (cf. Appelquist 2022), which deed allows Squilla to be used in the
sense of the current concept of Charybdis (Martínez-Azorín et al. 2022).
Previous phylogenetic analyses (Pfosser & Speta 2001, 2004, Manning et al. 2004, Pfosser et al. 2012) show
samples of Squilla (as Charybdis) forming a strongly supported clade. Many authors recently accepted Charybdis as
distinct from Urginea and Drimia at generic rank (cf. Speta 1998b, Pfosser & Speta 2001, 2004, Conti et al. 2005,
Jeanmonod & Gamisans 2007, Bacchetta et al. 2012, Ali et al. 2013, Véla et al. 2016), although others (e.g. Manning
et al. 2004, Manning & Goldblatt 2018, WCSP 2021) place them in an expanded and broadly circumscribed Drimia.

The phylogenetic analyses of Martínez-Azorín et al. (2023a) included 22 samples of Squilla, which formed a strongly
supported clade in an isolated position within Urgineoideae. Therefore, based on the distinct morphology and genetic
divergence of these plants, we here accept Squilla following Martínez-Azorín et al. (2022), which is mostly restricted
to the Mediterranean Basin.
FIGURE 54. Species of Squilla Steinh. displayed in horizontal rows of images. 1. Squilla pancration Steinh.; 2. Squilla serotina (Schousb.)
Mart.-Azorín et al.; 3. Squilla undulata (Desf.) Mart.-Azorín et al. (Photograph of leaves by J. Quiles).
Squilla anthericoides (Poir.) Jord. & Fourr., Icon. Fl. Eur. 2: 2 (1868) ≡ Scilla anthericoides Poir., Voy. Barbarie 2: 149 (1789), basionym ≡
Ornithogalum anthericoides (Poir.) Link ex Steud., Nomencl. Bot. 1: 573 (1821) ≡ Urginea anthericoides (Poir.) Steinh. in Ann. Sci.
Nat., Bot. sér. 2, 1: 328 (1834) ≡ Charybdis anthericoides (Poir.) Véla & Bélair in Phytotaxa 288: 155 (2016) ≡ Drimia anthericoides
(Poir.) Véla & Bélair in Willdenowia 49: 423 (2019) ≡ Urginea maritima var. anthericoides (Poir.) Maire & Weiller, Fl. Afrique N.
5: 163 (1958). Type:—[ALGERIA.] Ex Numidia, herb. Poiret (P02157340! lecto. designated by Véla et al. in Phytotaxa 288: 155.
2016: the specimen on the upper right side of the sheet).
Squilla aphylla (Forssk.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 3 (2022) ≡ Anthericum aphyllum Forssk.,
Fl. Aegypt.-Arab.: 209 (1775), basionym ≡ Urginea aphylla (Forssk.) Speta in Linzer Biol. Beitr. 12(1): 229 (1980) ≡ Charybdis
aphylla (Forssk.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia aphylla (Forssk.) J.C.Manning & Goldblatt in Edinburgh
J. Bot. 60(3): 556 (2004). Type:—[GREECE]. “Anthericum” F. Herb. Forskålii n° 10 [IDC microfiche foto: Forsskål nr. 129 III, 1–2]
(C10001668! lecto. designated by Martínez-Azorín et al. 2022).
= Ornithogalum elatum Andrews in Bot. Repos. 8, pl. DXXVIII (1808) ≡ Charybdis elata (Andrews) Speta in Phyton (Horn, Austria)
38(1): 60 (1998) ≡ Drimia excelsa J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004), replac. name [non Drimia elata
Jacq., Collectanea 5(Suppl.): 38. 1797]. Type:—illustration in Andrews (1808), pl. DXXVIII (lecto. designated by Martínez-Azorín
et al. 2022: only the flowering scape and the flower dissection).
Squilla glaucophylla (Bacch., Brullo, D’Emerico, Pontec. & Salmeri) Mart.-Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-
e78272: 4 (2022) ≡ Charybdis glaucophylla Bacch., Brullo, D’Emerico, Pontec. & Salmeri in Phytotaxa 69: 18 (2012), basionym
≡ Drimia glaucophylla (Bacch., Brullo, D’Emerico, Pontec. & Salmeri) Raus in Willdenowia 46(3): 424 (2016). Type:—ITALY.
Sardinia: Isola di San Pietro: Cala Vinagra, Carloforte, elev. 63 m, 19 July 2004, G. Bacchetta & C. Pontecorvo s.n. (CAT, holo.;
CAG, CAT, iso.).
Squilla hesperia (Webb & Berthel.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 4 (2022) ≡ Urginea hesperia
Webb & Berthel., Hist. Nat. Iles Canaries (Phytogr., sect. 3) 3(2): 339 (1848), basionym ≡ Charybdis hesperia (Webb & Berthel.)
Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia hesperia (Webb & Berthel.) J.C.Manning & Goldblatt in Edinburgh J.
Bot. 60(3): 556 (2004) ≡ U. maritima var. hesperia (Webb & Berthel.) Svent. in Index Sem. Acclim. Pl. Arautapae 1969: 41 (1970) ≡
Drimia maritima var. hesperia (Webb & Berthel.) A.Hansen & Sunding, Fl. Macaronesia, Checklist Vasc. Pl., rev. ed. 2, 2: 4 (1979)
(Figs 4.10, 53.1). Type:—SPAIN. Canary Islands. Urginea hesperia Nobis “3”, Herbarium Webbianum nº 182548 (FI000284!, lecto.
designated by Martínez-Azorín et al. 2022).
Squilla maritima (L.) Steinh. in Ann. Sci. Nat., Bot. sér. 2, 6: 279 (1836) ≡ Scilla maritima L., Sp. Pl.: 308 (1753), basionym ≡ Ornithogalum
maritimum (L.) Lam., Fl. Franç. 3: 276 (1779) ≡ Urginea maritima (L.) Baker in J. Linn. Soc., Bot. 13: 221 (1872) ≡ Drimia maritima
(L.) Stearn in Ann. Mus. Goulandris 4: 204 (1978) ≡ Charybdis maritima (L.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡
Stellaris scilla Moench, Methodus: 304 (1794), nom. illeg. [Art. 52 of the ICN] ≡ Ornithogalum squilla Ker Gawl. in Bot. Mag. 23,
t. 918 (1806), nom. illeg. [Art. 52 of the ICN] ≡ U. scilla Steinh. in Ann. Sci. Nat., Bot. sér. 2, 1: 331 (1834), nom. illeg. [Art. 52 of
the ICN] (Figs 4.11, 52.1, 52.2, 52.3, 53.2). Type:—illustration in Clusius (1601: 171, Fig. 52.1), “Scillae hispanicae flos & semen”
(lecto. designated by Martínez-Azorín et al. 2022b: only the illustration of the bulb with a flowering scape). Epitype (designated by
Martínez-Azorín et al. 2022):—PORTUGAL. Sierra de Monsanto, environs de Lisbonne, pentes pierreuses, July−August 1877, J.
Daveau (P02166423! epi.; P02166260!, P02166417!, P01811235!, P01811236!, isoepi.).
Comments on typification:—Recently, Martínez-Azorín et al. (2022) considered that the lectotype of Scilla maritima, first designated by
Ferrer-Gallego (2013: 40) on the voucher L0052807, should be disregarded according to Art. 8.2 of the ICN, because completely
developed adult leaves in this species are never coetaneous with flowers at anthesis in a given population. This implies that the
“lectotype” selected by Ferrer-Gallego (2013) includes two specimens collected at different times: i) leaves, and ii) the flowering
scape. Considering Ferrer-Gallegos’s (2022) reaction to our lectotypification, and in the light of the recentmost proposal by Mosyakin
& McNeill (2022) to amend Art. 8, after which the information in the protologue will rule, we still consider that the voucher
L0052807 cannot be a proper lectotype, following Linnaeus’s statement in Hortus Cliffortianus (1738: 123) that is cited in the Scilla
maritima protologue of 1753: “Scilla radice tunicata [….] Singulare est quod foliis destitute floreat.” Further, Clusius (1601: 171),
which is also cited in the protologue of S. maritima, illustrated the species through both a bulb with fully-developed leaves (winter-
spring facies) and two bulbs with flowering and fruiting scapes; those latter figures are completely lacking leaves (later summer-
early autumn facies); in addition, Clusius (1601) describes the typical hysteranthous behaviour of the species in Hispania as follows:
“Copiose nascitur supra Olysipponem & plerisque aliis Lusitaniae & Hispaniae locis. Floret Augusto & Septembri, semen Octobri
& Novembri maturescit; folia maturo jam semine, & exarido caule, emergunt Novembri & Decembri.” Based on those facts, we
do believe that Linnaeus never saw plants of Scilla maritima showing flowers and leaves coetaneously, as he indirectly claimed in
his Hortus Cliffortianus (Linnaeus 1738), and hence the Linnaean voucher L0052807 most likely represents a “collage” of mixed
gatherings of flowers and leaves collected at different times, and it as a whole cannot be eligible as lectotype. Therefore, the lectotype
and epitype we designated (Martínez-Azorín et al. 2022) are considered here as appropriate to preserve the currently most widely
accepted concept of this species.
= Scilla rubra Garsault, Descr. Pl. Anim. 4: 314, pl. 527 (1767). Type:—illustration in Garsault (l.c.), pl. 527 “Scilla rubra” (lecto.
designated by Martínez-Azorín et al. 2022: only the illustrations of the bulb with a flowering scape, and the flowers).
Squilla maura (Maire) Mart.-Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 7 (2022) ≡ Urginea maura Maire in Bull.
Soc. Hist. Nat. Afrique N. 14: 158 (1923), basionym ≡ Charybdis maura (Maire) Speta in Phyton (Horn, Austria) 38(1): 60 (1998)
≡ Drimia maura (Maire) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004) ≡ Urginea maritima var. maura (Maire)
Maire, Fl. Afrique N. 5: 165 (1958) ≡ Drimia maritima subsp. maura (Maire) Förther & Podlech in Sendtnera 7: 86 (2001). Type:—
MOROCCO. Grand Atlas: Mentaga, rocailles calcaires, elev. 200–1000 m, feuilles glauques, 14 April 1922, R. Maire (MPU000488!
lecto. designated by Martínez-Azorín et al. 2022).
= Urginea maritima subsp. maura var. angustifolia Maire in Bull. Soc. Hist. Nat. Afrique N. 29: 454 (1938) ≡ Urginea maritima var.
maura f. angustifolia (Maire) Maire, Fl. Afrique N. 5: 165 (1958), basionym. Type:—MOROCCO. Anti Atlas: près de Kerdous,
pentes rocheuses (quartzites), elev. 800 m, 38 March 1937, R. Maire (MPU003971! lecto. designated by Martínez-Azorín et al.
2022).

Squilla numidica Jord. & Fourr., Icon. Fl. Eur. 2: 1 (1868) ≡ Urginea numidica (Jord. & Fourr.) Grey, Hardy Bulbs 2: 632 (1938) ≡
Urginea maritima var. numidica (Jord. & Fourr.) Baker in J. Linn. Soc., Bot. 13: 221 (1873) ≡ Charybdis numidica (Jord. & Fourr.)
Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia numidica (Jord. & Fourr.) J.C.Manning & Goldblatt in Edinburgh J. Bot.
60(3): 557 (2004). Type:—[ALGERIA.] “Squilla maritima numidica / de Bône (Algerie)”, 6 Novembre 1867 (LY0073734 [digital
image!] lecto. designated by Martínez-Azorín et al. 2022).
= Squilla sphaeroidea Jord. & Fourr., Icon. Fl. Eur. 2: 1 (1868) ≡ Urginea maritima var. sphaeroidea (Jord. & Fourr.) Baker in J. Linn.
Soc., Bot. 13: 221 (1873) ≡ U. sphaeroidea (Jord. & Fourr.) Grey, Hardy Bulbs 2: 632 (1938). Type:—illustration in Jordan &
Fourreau (1868): Tab. CCII “278. Squilla sphaeroidea” [lecto. designated by Martínez-Azorín et al. 2022: only the illustrations of
the flowering scape, flowers and bracts (Figs 1−12); excluding bulb, leaves, fruits and seeds (Figs 13−16)].
Squilla palaestina (M.B.Crespo, Mart.-Azorín & M.Á.Alonso) Mart.-Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 8
(2022) ≡ Drimia palaestina M.B.Crespo, Mart.-Azorín & M.Á.Alonso in Pl. Syst. Evol. 306: 13 (2020), basionym (Figs 4.12, 53.3).
Type:—PALESTINE. [Palestinian Authority], Judean Mts. South of Yata, Submediterranean batha, 23 September 1970, Y. Plitmann
& M. Raviv 26089 (HUJ132724! holo.).
Squilla pancration Steinh. in Ann. Sci. Nat., Bot. sér. 2, 6: 279 (1836) ≡ Scilla pancration (Steinh.) Nyman, Syll. Fl. Eur.: 369 (1855) ≡
Urginea pancration (Steinh.) Schroff in Z. Allg. Oesterr. Apotheker-Vereines 3 (Oesterr. Z. Pharmazie 13): 42 (1865) ≡ Charybdis
pancration (Steinh.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998) ≡ Drimia pancration (Steinh.) J.C.Manning & Goldblatt in
Edinburgh J. Bot. 60(3): 557 (2004) ≡ Urginea maritima var. pancration (Steinh.) Baker in J. Linn. Soc., Bot. 13: 221 (1873) ≡ U.
maritima subsp. pancration (Steinh.) K. Richter, Pl. Eur. 1: 218 (1890) (Figs 4.13, 54.1). Type:—MALTA. Gozo island (Għawdex),
pr. Ir-Ramla, between San Blas and S. Philip, 33SVV359908, elev. 70 m, flowered in cultivation at the University of Alicante on 04
September 2013, J.L. Villar & E. Martínez s.n. (ABH70437! neo. designated by Martínez-Azorín et al. 2022).
= Squilla insularis Jord. & Fourr., Icon. Fl. Eur. 2: 2 (1868) ≡ Urginea maritima var. insularis (Jord. & Fourr.) Baker in J. Linn. Soc., Bot.
13: 221 (1873) ≡ U. maritima subsp. insularis (Jord. & Fourr.) K. Richt., Pl. Eur. 1: 218 (1890) ≡ U. scilla var. insularis (Jord. &
Fourr.) Rouy, Fl. France 12: 425 (1910) ≡ U. insularis (Jord. & Fourr.) Grey, Hardy Bulbs 2: 632 (1938). Type:—illustration in Jordan
& Fourreau (1868): Tab. CCIII “279. Squilla insularis” [lecto. designated by Martínez-Azorín et al. 2022: only the illustrations of the
flowering scape, flowers and bracts (Figs 1−12); excluding bulb, leaves, fruits and seeds (Figs 13−16)].
= Squilla littoralis Jord. & Fourr., Icon. Fl. Eur. 2: 2 (1868) ≡ Urginea maritima var. littoralis (Jord. & Fourr.) Baker in J. Linn. Soc.,
Bot. 13: 221–222 (1873) ≡ U. maritima subsp. littoralis (Jord. & Fourr.) K.Richt., Pl. Eur. 1: 218 (1890) ≡ U. littoralis (Jord. &
Fourr.) Grey, Hardy Bulbs 2: 632 (1938). Type:—illustration in Jordan & Fourreau (1868): Tab. CCIV “280. Squilla littoralis” [lecto.
designated by Martínez-Azorín et al. 2022: only the illustrations of the flowering scape, flowers and bracts (Figs 1−12); excluding
bulb, leaves, fruits and seeds (Figs 13−16)].
Squilla secundiflora (Maire) Mart.-Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 9 (2022) ≡ Urginea anthericoides
var. secundiflora Maire in Bull. Soc. Hist. Nat. Afrique N. 24: 229 (1933), basionym ≡ Drimia secundiflora (Maire) M.B.Crespo,
Mart.-Azorín & M.Á.Alonso in Pl. Syst. Evol. 306: 15 (2020) (Fig. 4.14). Type:—MOROCCO. Forêt de la Mamora, bulbes cultivés
à Alger ayant fleuri en septembre 1932, 28 October 1932, R. Maire s.n. (MPU003124! lecto designated by Crespo et al. 2020: 15 [as
“U. anthericoides var. secunda Maire”]; only the flowering scape).
Squilla serotina (Schousb.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 9 (2022) ≡ Scilla serotina Schousb.,
Iagttag. Vextrig. Marokko: 165 (1800), basionym ≡ Urginea anthericoides var. serotina (Schousb.) Font Quer in Bol. Real Soc.
Esp. Hist. Nat. 27: 45 (1927) (Figs 4.15, 54.2). Type:—[MOROCCO.] Herb. Liebm[ann]. Marocco, Schousboe (C10000939! lecto.
designated by Crespo et al. 2020: 11, [IDC microfiche foto: Type Herbarium nr. 183 I,4]; C10000940 [digital image!] isolecto. [IDC
microfiche foto: Type Herbarium nr. 183 I,5]).
= Urginea undulata var. tazensis Batt. & Maire in Bull. Soc. Hist. Nat. Afrique N. 22: 318 (1931) ≡ Urginea tazensis (Batt. & Maire) Maire
in Bull. Soc. Hist. Nat. Afrique N. 31: 43 (1940) ≡ Urginea undulata subsp. tazensis (Batt. & Maire) Maire & Weiller, Fl. Afrique N.
5: 158 (1958) ≡ Drimia tazensis (Maire) Stearn in Ann. Mus. Goulandris 4: 208 (1978) ≡ Charybdis tazensis (Batt. & Maire) Speta in
Stapfia 75: 167 (2001). Type:—MOROCCO. Tiznit, bulbs cultivated in Alger (Algeria), 18 September 1939, R. Maire (MPU004613!
neo. designated by Crespo et al. 2020: 11).
= Urginea undulata var. major Gatt. & Weiller in Bull. Soc. Hist. Nat. Afrique N. 28: 539 (1937). Type:—MOROCCO. Tiznit, sandy soils,
cultivated in Alger (Algeria), 8 September 1936, R. Maire (MPU001417! neo. designated by Crespo et al. 2020: 11).
Squilla undulata (Desf.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso in Medit. Bot. 43-e78272: 10 (2022) ≡ Scilla undulata Desf. in Méd.
Eclairée Sci. Phys. 3(6): 161 (1792), basionym ≡ Urginea undulata (Desf.) Steinh. in Ann. Sci. Nat., Bot. sér. 2, 1: 330 (1834) ≡
Epimenidion undulatum (Desf.) Raf., Fl. Tellur. 2: 13 (1837) ≡ Prospero undulatum (Desf.) Salisb., Gen. Pl.: 28 (1866) ≡ Drimia
undata Stearn in Ann. Mus. Goulandris 4: 208 (1978), replac. name [non Drimia undulata Jacq. in Willd., Sp. Pl. ed. 4, 2(1): 166.
(1799), nec Jacq., Collectanea 5 (Suppl.): 41. 1797, nom. inval.] ≡ Charybdis undulata (Desf.) Speta in Stapfia 75: 167 (2001), nom.
illeg. [Art. 52 of the ICN] (Figs 4.16, 52.5, 54.3). Type:—ALGERIA. Herbier de la Flore Atlantique, no. 707 Scilla undulata [Tunis
et Alger, collines arides, elle fleurit en hiver, 1787], Desfontaines (P00320347! lecto. designated by Crespo et al. 2019: 91: the
flowering scape with bulb on the left side of the sheet; MPU017529! isolecto.).
= Urginea undulata f. caeculi Pau in Butl. Inst. Catalana His. Nat. 16: 136 (1916), basionym ≡ Urginea undulata var. caeculi (Pau) Pau
in Bol. Soc. Ibér. Ci. Nat. 23: 91 (1924) ≡ Drimia undata subsp. caeculi (Pau) Mateo & M.B.Crespo, Fl. Abrev. Comun. Valenciana:
430 (1995) ≡ Urginea undulata subsp. caeculi (Pau) M.B.Crespo & Mateo in Flora Montiber. 5: 58 (1997). Type:—SPAIN. Valencia
prov., Alginet, cultivada, August 1916, F. Moroder (MA21517! lecto. designated by Crespo & Mateo 1997: the scape on the right
side of the sheet, from a bulb with incipient leaves).
= Drimia purpurascens J.Jacq., Ecl. Pl. Rar. 1(3–4): 48, t. 30 (1812) ≡ Idothea purpurascens (J.Jacq.) Kunth, Enum. Pl. 4: 342 (1843) ≡
Idothearia purpurascens (J.Jacq.) C.Presl in Abh. Königl. Böhm. Ges. Wiss. ser. 5, 3: 544 (1845), nom. illeg. superfl. [Art. 52 of the
ICN]. Type:—illustration in J.Jacq., Ecl. Pl. Rar. 1(3–4), t. 30. 1812 (Fig. 52.4) (lecto. designated by Crespo et al. 2020: only the
flowering scape with bulb). Epitype (designated by Crespo et al. 2020: 9):—ITALY. Sardaigne, 1838, E. Thomas (P02166250! epi.:
the fragment on the left side of the sheet).
22. Striatula M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Phyton (Horn, Austria) 59: 93 (2019) (Fig. 55).
Typus generis:—S. platyphylla (B.Nord.) M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig (holotype).
FIGURE 55. Species of Striatula M.Pinter et al. displayed in horizontal rows of images. 1. Striatula oliverorum (J.C.Manning) M.Pinter
et al.; 2. Striatula platyphylla (B.Nord.) M.Pinter et al.
Description:—Small bulbous geophyte. Bulb hypogeal, solitary, subglobose to ovoid, 1.5−4.0 cm in diam., outer
tunics pale-brownish to brownish when drying, membranous, adherent, forming distinct neck, inner tunics whitish,
compact, soft and fleshy. Roots thickened and branched. Leaves 1−2 per bulb, proteranthous, flattened, ovate to elliptic
to elliptic-oblong, 1–4 cm long, appressed to ground, proteranthous, fleshy or subsucculent, dull or dark green, with
2 or 4 longitudinal furrows, minutely and densely velutinous on adaxial side, glabrous on abaxial side, with obtuse to
rounded apex. Inflorescence 1(–2) per bulb, erect, racemose, 1.5−12.0 cm long, moderately dense, with 4–50 flowers;
peduncle 3−15(−30) cm long, usually reddish brown or purplish red at base with dense and short to minute papillae or
hairs disposed in straight vertical lines; pedicels patent or erecto-patent to spreading, 3−10 mm long, minutely scabrid-
papillate or glabrate. Bracts ovate-triangular to lanceolate, 1.0−2.5 mm long, lowermost with distinct spur, 1−2 mm
long; bracteoles absent. Flowers shallowly campanulate, urceolate or subglobose, nodding to subpatent. Tepals 6,
3.5–6.0 mm long, basally connate for ¼ to ½ of their length, reddish brown to creamy pink or pinkish white with broad
purplish to green or greenish median stripe visible on both sides. Stamens 6, included in perigone, erect and connivent
to style; filaments adnate to tepals for less than 1 mm, free parts 1.0−1.5 mm long, whitish, laxly papillate-puberulous
or smooth; anthers ellipsoid or ovate-sagittate, 1.0−1.8 mm long, yellow, dehiscing with apical pore-like slits up to ½
of theca length, sometimes basally barbellate. Ovary ovoid to subquadrate, 1.8−2.0 mm long, light green, glabrous.
Style columnar, erect, terete, 1.0−1.8 mm long, white. Capsule broadly ovoid to subglobose, 4.5–7.0 mm long, valves
completely dehiscing from base; tepals cohering and inrolled above ovary after anthesis, circumcissile from base and
persisting as a cap at the top of the developing capsules. Seeds elliptic-oblong to reniform, 2–4 mm long, flattened,
with prominent embryo and short, wings, with black, glossy testa and alveolate-reticulate testa cells with collapsed
periclinal walls and prominent anticlinal walls.
Number of species and distribution:—Striatula includes 2 species from the Western Cape, Eastern Cape, and
Northern Cape Provinces of South Africa and southern Namibia (Fig. 49), being restricted to the Cape and Karoo-
Namib Regions and the Southern Section of the Zambezian Subregion (sensu Takhtajan 1986 and Martínez-Azorín et
al. 2023a). For complete morphological descriptions of the two taxa see Nordenstam (1970) and Manning & Oliver
(2009).
Karyology:—2n=20 (Bruyns & Vosa 1987, as Rhadamanthus platyphyllus B.Nord.).
History, diagnostic characters, and taxonomic relationships:—Nordenstam (1970) described Rhadamanthus
platyphyllus, a species sharing the general flower morphology and anther dehiscence of that genus, but differing in
having two, flat, ovate, velutinous leaves with longitudinal furrows, which are appressed to the ground and the anthers
basally barbellate with spine-like or papilliform outgrowths, whilst the remaining species of Rhadamanthus sensus
Nordenstam (1970) show numerous, filiform, erect leaves. Drimia oliverorum was described from southern Namibia
and shares the peculiar leaf morphology and flower structure of R. platyphyllus. The phylogenetic analyses of Pfosser
et al. (2012) showed that seven samples of the Rhadamanthus platyphyllus group formed a perfectly supported clade,
which is located far from the clade including other Rhadamanthus species, the latter being basal to all Urgineoideae
when Bowiea is excluded. The clear phylogenetic divergence of both R. platyphyllus and D. oliverorum, together
with their distinct vegetative morphology led Pinter et al. (2019) to described Striatula to accommodate these two
taxa, a solution followed in the present work. The phylogenetic studies by Martínez-Azorín et al. (2023a) confirmed
the results by Pfosser et al. (2012) and include 12 samples of Striatula, which form a perfectly supported clade in an
isolated position within Urgineoideae. This genus can be easily identified by its unique syndrome of morphological
characters, and is restricted to southern and western South Africa and southern Namibia.
Striatula oliverorum (J.C.Manning) M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Phyton (Horn, Austria) 59: 95 (2019) ≡ Drimia
oliverorum J.C.Manning in Manning & Oliver in Bothalia 39(2): 225 (2009), basionym (Figs 4.17, 55.1). Type:—NAMIBIA.
Witputz (2716): Huib Hoch Plateau, Zebrasfontein, (–DB), elev. 1200 m, 29 June 1989 [in leaf only], E.G.H. Oliver & I.M. Oliver
9164 (NBG0233338-0! holo.).
Striatula platyphylla (B.Nord.) M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Phyton (Horn, Austria) 59: 95 (2019) ≡ Rhadamanthus
platyphyllus B.Nord. in Taxon 63(6): 1332 (2014), basionym ≡ R. platyphyllus B.Nord. in Bot. Not. 123: 172 (1970), nom. inval.
≡ Drimia platyphylla (B.Nord.) J.C.Manning & Goldblatt in Strelitzia 40: 132 (2018) ≡ D. platyphylla (B.Nord.) J.C.Manning &
Goldblatt in Strelitzia 9: 712 (2000), nom. inval. (Figs 4.18, 55.2). Type:—SOUTH AFRICA. Western Cape. Wuppertal (3219):
Clanwilliam Div., in stony or shallow sand on rocky slope below shale band below Cederberg Tafelberg, (–AC), elev. 3500–4000
ft., 16 December 1950, Esterhuysen 18135, leaves added April 1951 (BOL140333! holo.: the bulb with two leaves at the bottom left
hand side corner of the sheet).
23. Tenicroa Raf., Fl. Tellur. 3: 52 (1837) (Figs 56–59). Typus generis:—Tenicroa fragrans (Jacq.) Raf. ≡ Anthericum
fragrans Jacq. ≡ Drimia fragrans (Jacq.) J.C.Manning & Goldblatt (holotype).
= Pilasia Raf., Fl. Tellur. 3: 53 (1837). Typus generis:—P. filifolia (Jacq.) Raf. ≡ Anthericum filifolium Jacq. ≡ Tenicroa filifolia (Jacq.)
Oberm. ≡ Drimia filifolia (Jacq.) J.C.Manning & Goldblatt (holotype).
= Sypharissa Salisb., Gen. Pl. [Salisbury]: 37 (1866) ≡ Urginea sect. Sypharissa (Salisb.) Baker in J. Linn. Soc., Bot. 13: 216 (1873) ≡
Drimia sect. Sypharissa (Salisb.) J.C.Manning & Goldblatt in Strelitzia 40: 111 (2018). Typus generis:—S. exuviata (Jacq.) Salisb.
ex Oberm. ≡ Anthericum exuviatum Jacq. ≡ Tenicroa exuviata (Jacq.) Speta ≡ Drimia exuviata (Jacq.) Jessop (lectotype designated
by Obermeyer in Bothalia 13: 111. 1980).
= Drimia sect. Juncifoliae J.C.Manning & Goldblatt in Strelitzia 40: 107 (2018). Typus sectionis:—D. juncifolia J.C.Manning &
J.M.J.Deacon ≡ Tenicroa juncifolia (J.C.Manning & J.M.J.Deacon) Mart.-Azorín et al. (holotype).
.
FIGURE 56. Species of Tenicroa Raf. 1. Tenicroa fragrans (Jacq.) Raf. from Jacquin (1797: t. 86, as Anthericum fragrans Jacq.); 2.
Tenicroa exuviata (Jacq.) Speta from Jacquin (1794: t. 415, as Anthericum exuviatum Jacq.); 3. Tenicroa filifolia (Jacq.) Oberm. from
Jacquin (1794: t. 414, as Anthericum filifolium Jacq.); 4. Tenicroa filifolia (Jacq.) Oberm. from Ker Gawler (1821: t. 557, as Albuca filifolia
(Jacq.) Ker Gawl.).

FIGURE 57. Species of Tenicroa Raf. 1. Tenicroa exuviata (Jacq.) Speta from Ker Gawler (1805: t. 871, as Albuca exuviata (Jacq.) Ker
Gawl.); 2. Tenicroa fragrans (Jacq.) Raf. from Ker Gawler (1818: t. 311, as Albuca fugax Ker Gawl.); 3. Tenicroa multifolia (G.J.Lewis)
Oberm. from Lewis (1952: 10, as Urginea multifolia G.J.Lewis).
FIGURE 58. Species of Tenicroa Raf. displayed in horizontal rows of images. 1. Tenicroa applanata M.Pinter et al.; 2. Tenicroa exuviata
(Jacq.) Speta; 3. Tenicroa fibrosa M.Pinter et al.; 4. Tenicroa filifolia (Jacq.) Oberm.
Description:—Medium-sized bulbous geophyte. Bulb hypogeal, ovoid to subglobose, up to 7 cm in diam., extended

into neck surrounded by cataphylls with raised, transverse purple to brown ribs, inner scales white to pinkish, fleshy,
compact or rarely loose, spirally arranged, rarely distichous, sometimes fibrose. Roots distinctly thickened and branched.
Leaves 1‒50 per bulb, 3‒80 cm long, narrowly linear, from terete to channelled, synanthous or rarely hysteranthous,
entire or rarely scabrid on margins, green or somewhat glaucous, usually leathery, sometimes twisted, smooth,
glabrous. Inflorescence a long, erect, multiflowered raceme, 2‒25 cm long, with up to 60 flowers; peduncle 10‒40 cm
long, green, grey or purple, terete, erect, smooth and glabrous; pedicels 3‒15 mm long, shorter than tepals at anthesis,
spreading in flower and erect in fruit. Bracts narrowly lanceolate, long acuminate, lowermost auriculate with long
spur, sometimes bifid or branched and sinuous; bracteoles absent. Flowers stellate with patent tepals, diurnal. Tepals
6, biseriate, 6‒17 mm long, free, ovate-lanceolate to elliptic, white with well defined, narrow, purplish longitudinal
band. Stamens 6, suberect and slightly spreading; filaments filiform, 4‒8 mm long, tapering, smooth, white, slightly
sigmoid and curving distally; anthers yellow, erect, basifixed, deshicing longitudinally along their whole length, with
yellow pollen. Ovary ovate-oblong, green to yellow. Style 4‒8 mm long, declinate, sigmoid, white or rarely tinged with

purple, with capitate, papillose stigma. Capsule ovate to ellipsoid, 6‒25 mm long, valves completely dehiscing from
base, tepals cohering and inrolled above ovary after anthesis, circumcissile from base and persisting as a cap at the top
of the developing capsules. Seeds subellipsoid to compressed, (4–)5‒11 mm long, flattened, embryo prominent with
wide or short wings, testa black, dull or glossy, with sinuous cell walls.
FIGURE 59. Species of Tenicroa Raf. displayed in horizontal rows of images. 1. Tenicroa fragrans (Jacq.) Raf.; 2. Tenicroa multifolia
(G.J.Lewis) Oberm.; 3. Tenicroa namibensis M.Pinter et al. (Photographs of bulb with leaf by H. Kolberg and in flower by A. Burke); 4.
Tenicroa polyantha M.Pinter et al.; 5. Tenicroa unifolia (A.V.Duthie) M.Pinter et al.
Number of species and distribution:—Tenicroa includes twelve species restricted to southern and western South
Africa and southern Namibia, it being therefore restricted to the Cape and Karoo-Namib Regions (sensu Taktajan
1986) (Fig. 49). For complete species characteristics see Pinter et al. (2020).
Karyology:—2n=20 (Speta 1998a).
History, diagnostic characters, and taxonomic relationships:—Rafinesque (1837) described Tenicroa to include
T. fragrans (Jacquin 1797: t. 86) Rafinesque (1837: 53), a species with rotate flowers, suberect stamens and deflexed
style, among other characters. The revision by Obermeyer (1980b, 1981a) accepted four species in the genus from
southern and western South Africa and southern Namibia: T. fragrans, T. exuviata (Jacquin 1794: 18) Speta (1980: 195),
T. filifolia (Jacquin 1794: 18) Obermeyer (1981a: 577), and T. multifolia (Lewis: 1952: 9) Obermeyer (1981a: 577). All
of them shared distinct characters such as the synanthous leaves surrounded by sheathing cataphylls that are markedly
striate with raised, dark, transversal ridges; the stellate flowers with almost free tepals with a narrow, longitudinal dark
band on the abaxial side; the suberect to slightly spreading stamens with basifixed anthers and the elongate, deflexed
and curved style; and papillate stigma. Despite the distinct morphology of taxa in this genus, they have historically
been placed in 8 different genera: Anthericum Linnaeus (1753: 310), Albuca, Drimia, Ornithogalum, Phalangium
Miller (1754: s.n.), Pilasia Rafinesque (1837: 53), Sypharissa Salisbury (1866: 37), and Urginea, representing a good
example of the uncertainty in taxonomy and systematics of urgineoid taxa (Pinter et al. 2020). However, based on the
clear synapomorphies of Tenicroa, researchers in Urgineoideae (Jessop 1977, Obermeyer 1980b, 1981a, Speta 1998a,
1998b) usually recognised it as a distinct genus. Conversely, Manning et al. (2004) synonymised Tenicroa to Drimia
sensu lato, and later Manning & Goldblatt (2018) placed the Tenicroa species recognised by Obermeyer (1980b) in
D. sect. Sypharissa, whilst including T. filifolia as synonym of T. exuviata. They also separated their new D. sect.
Juncifoliae to associate two newly described species [Drimia decipiens Manning & Goldblatt (2018: 109) and D.
juncifolia J.C.Manning & J.M.J.Deacon in Manning & Goldblatt (2018: 107)] based on their hysteranthous leaves,
weakly barred cataphylls, and medifixed anthers that are not deflexed. Martínez-Azorín et al. (2019a) considered the
latter distinction insufficient to recognise both sections, and transferred the two new species to Tenicroa.
The phylogenetic analyses of Manning et al. (2004) recovered two samples of Tenicroa as monophyletic, when
excluding T. nana as the monotypic Mucinaea (Pinter et al. 2013). Pfosser et al. (2012) phylogenetically analysed nine
samples of Tenicroa that formed a perfectly supported clade being sister to a clade comprising Litanthus plus Schizobasis.
Furthermore, the phylogenetic analyses of Martínez-Azorín et al. (2023a) included 21 samples of Tenicroa, which
formed a strongly supported and isolated clade within a large polytomy. Therefore, we here accept Tenicroa at genus
rank based on the unique syndrome of morphological characters and its isolated phylogenetic position, following the
latest revision of Tenicroa by Pinter et al. (2020), who accepted twelve species restricted to South Africa and southern
Namibia.
Tenicroa applanata M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Phyton (Horn, Austria) 60: 64 (2020) (Fig. 58.1). Type:—
SOUTH AFRICA. Western Cape. Cape Town (3318): near Tinie Versveld Wild Flower Reserve, between Darling and Yzerfontein,
(–AD), elev. 122 m, 20 September 2015, M. Martínez-Azorín, M. Pinter, M.B. Crespo & M.Á. Alonso MMA1128 (GRA! holo.; ABH!
iso.).
Tenicroa decipiens (J.C.Manning & Goldblatt) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 294 (2019)
≡ Drimia decipiens J.C.Manning & Goldblatt in Strelitzia 40: 109 (2018), basionym. Type:—SOUTH AFRICA. Western Cape.
Wuppertal (3219): Swartruggens, near turnoff to Kagga Kamma at summit of Skitterykloof, (–DC), 27 November 2017, J. Manning
3637 (NBG holo.).
Tenicroa exuviata (Jacq.) Speta in Linzer Biol. Beitr. 12(1): 195 (1980) ≡ Anthericum exuviatum Jacq., Icon. Pl. Rar. 2 (13): 18, t. 415
(1794), basionym (Fig. 56.2) ≡ Phalangium exuviatum (Jacq.) Poir. in Lam., Encycl. 5: 243 (1804) ≡ Albuca exuviata (Jacq.) Ker
Gawl. in Curtis’s Bot. Mag. 22: t. 871 (1805) (Fig. 57.1) ≡ Urginea exuviata (Jacq.) Steinh. in Ann. Sci. Nat., sér. 2, 2: 330 (1834)
≡ Ornithogalum exuviatum (Jacq.) Kunth, Enum. Pl. 4: 369 (1843) ≡ Drimia exuviata (Jacq.) Jessop in J. S. African Bot. 43(4): 276
(1977) ≡ Sypharissa exuviata (Jacq.) Salisb. ex Oberm. in Bothalia 13(1-2): 113 (1980) (Figs 4.19, 58.2). Type:—SOUTH AFRICA.
Crescit ad Promontorium bonae Spei, icon in Jacq., Icon. Pl. Rar. 2: t. 415 (1794) (Fig. 56.2) (lecto. designated as “Type” by Jessop
in J. S. African Bot. 43: 276. 1977). Epitype (designated by Pinter et al. 2020):—SOUTH AFRICA. Bredasdorp (3420): N side of
Potberg, Bredasdorp Div., (–BC), elev. ca. 400 ft., 13 November 1962, J.P.H. Acocks 23016 (PRE0046767-0! epi.).

Tenicroa fibrosa M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Phyton (Horn, Austria) 60: 70 (2020) (Fig. 58.3). Type:—SOUTH
AFRICA. Western Cape. Vanrhynsdorp (3118): 21 m S.S.W. of Vredendal, (–CD), elev. ca 500 ft., 26 August 1958, J.P.H. Acocks
19713 (PRE0046760! holo.; BOL!, M0223028! iso.).
Tenicroa filifolia (Jacq.) Oberm. in J. S. African Bot. 47(3): 577 (1981) ≡ Anthericum filifolium Jacq., Icon. Pl. Rar. 2: 18, t. 414 (1794),
basionym (Fig. 56.3) ≡ Phalangium filifolium (Jacq.) Poir. in Lam., Encycl. 5: 242 (1804) ≡ Albuca filifolia (Jacq.) Ker Gawl. in
Bot. Reg. 7: 557 (1821) (Fig. 56.4) ≡ Urginea filifolia (Jacq.) Steinh. in Ann. Sci. Nat., sér. 2, 2: 329 (1834) ≡ Pilasia filifolia (Jacq.)
Raf., Fl. Tellur. 3: 53 (1837) ≡ Ornithogalum filifolium (Jacq.) Kunth, Enum. Pl. 4: 369 (1843) ≡ Sypharissa filifolia (Jacq.) Salisb.
ex Oberm. in Bothalia 13: 113 (1980) ≡ Drimia filifolia (Jacq.) J.C.Manning & Goldblatt in Strelitzia 9: 711 (2000) (Fig. 58.4).
Type:—Illustration in Jacq., Icon. Pl. Rar. 2 (15): t. 414 (1794) (Fig. 56.3) (lecto. designated as “Type” by Jessop in J. S. African
Bot. 43: 276. 1977).
= Anthericum spiratum Thunb., Prodr.: 62 (1794). Syntypes:—SOUTH AFRICA. ‘Cap. B. spei’, Thunberg s.n. (UPSTHUNB [8413 &
8414]-microfiche, syn.).
Tenicroa flexuosa (Adamson) M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Phyton (Horn, Austria) 60: 76 (2020) ≡ Urginea
flexuosa Adamson in J. S. African Bot. 8: 240 (1942), basionym. Type:—SOUTH AFRICA. Western Cape. Simonstown (3418): Cape
Peninsula, Smitswinkel Bay, (–AD), 3 January 1941, Adamson 3099 (BOL140326! holo.; NBG sub SAM0056131-0!, K000257378!
iso.).
Tenicroa fragrans (Jacq.) Raf., Fl. Tellur. 3: 53 (1837) ≡ Anthericum fragrans Jacq., Hort. Schoenbr. 1: t. 86 (1797), basionym (Fig. 56.1)
≡ Phalangium fragrans (Jacq.) Poir. in Lam., Encycl. 5: 247 (1804) ≡ Albuca fugax Ker Gawl. in Bot. Reg. 4: t. 311 (1818), nom. nov.
(Fig. 57.2) [non Albuca fragrans Jacq.] ≡ Urginea fragrans (Jacq.) Steinh. in Ann. Sci. Nat., sér. 2, 2: 328 (1834) ≡ Ornithogalum
fragrans (Jacq.) Kunth, Enum. Pl. 4: 366 (1843) ≡ Sypharissa fragrans Salisb. ex Oberm. in Bothalia 13(1−2): 113 (1980) ≡ Drimia
fragrans (Jacq.) J.C.Manning & Goldblatt in Strelitzia 9: 711 (2000) ≡ Albuca fugax Ker Gawl. in Bot. Reg. 4: t. 311 (1818), syn.
subst. (Figs 4.20, 59.1). Type:—SOUTH AFRICA. Western Cape: illustration in Jacq., Hort. Schoenbr. 1: t. 86 (1797) (Fig. 56.1)
(lecto. designated as “Type” by Jessop in J. S. African Bot. 43: 276. 1977). Epitype (designated by Pinter et al. 2020):—SOUTH
AFRICA. Western Cape. Vanrhynsdorp (3118): Summit of Gifberg, (−DA), elev. ca. 2000 ft., 16 November 1970, H. Hall 3906
(NBG91254!, 2 sheets epi.; PRE0046756-0! isoepi.).
Tenicroa juncifolia (J.C.Manning & J.M.J.Deacon) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397: 294 (2019) ≡
Drimia juncifolia J.C.Manning & J.M.J.Deacon in Strelitzia 40: 107 (2018), basionym. Type:—SOUTH AFRICA. Western Cape.
Cape Town (3318): Paarl, Brier’s Louw Nature Reserve, (–DD), 23 November 2013, J.M.J. Deacon 3078 (NBG holo.).
Tenicroa multifolia (G.J.Lewis) Oberm. in J. S. African Bot. 47(3): 577 (1981) ≡ Urginea multifolia G.J.Lewis in Ann. S. African Mus. 40:
9 (1952), basionym (Fig. 57.3) ≡ Drimia multifolia (G.J.Lewis) Jessop in J. S. African Bot. 43(4): 278 (1977) ≡ Sypharissa multifolia
(G.J.Lewis) Oberm. in Bothalia 13(1−2): 114 (1980) (Figs 5.1, 59.2). Type:—SOUTH AFRICA. Northern Cape. Springbok (2917):
27 miles S of Springbok, (–DD), 27 July 1950, Lewis 2302 (SAM0060870-0! holo.; NBG! iso.).
= Urginea capillifolia Schlechter, nom. nud. in sched.
Tenicroa namibensis M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Phyton (Horn, Austria) 60: 83 (2020) (Fig. 59.3). Type:—
NAMIBIA. Witputz (2716): Karas, (–AC), elev. 797 m, 12 August 2001, C.A. Mannheimer CM1607 (WIND77423! holo.).
Tenicroa polyantha M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Phyton (Horn, Austria) 60: 84 (2020) (Figs 5.2, 59.4). Type:—
SOUTH AFRICA: Western Cape. Cape Town (3318): Langebaan, eastern end of town, (–AA), elev. 55 m, 20 September 2015 [in
flower], M. Martínez-Azorín, M. Pinter, M.B. Crespo & M.Á. Alonso MMA1139 (GRA! holo.; ABH! iso.).
Tenicroa unifolia (A.V.Duthie) M.Pinter, Mart.-Azorín, M.B.Crespo & Wetschnig in Phyton (Horn, Austria) 60: 86 (2020) ≡ Urginea
unifolia A.V.Duthie in Ann. Univ. Stell. 6A: 8 (1928), basionym (Figs 5.3, 59.5). Type:—SOUTH AFRICA. Western Cape. Cape
Town (3318): Stellenbosch Flats, (–DD), October 1927, A.V. Duthie s.n. STE1891 (NBG0197708-1! holo.; BOL140325! iso.).
= Urginea ecklonii sensu Duthie in Ann. Univ. Stell. 6A: 6 (1928), non U. ecklonii Baker (1892).
= Urginea duthieae Adamson in J. S. African Bot. 8: 239 (1942) ≡ Drimia duthieae (Adamson) Jessop in J. S. African Bot. 43(4): 278
(1977). Type:—SOUTH AFRICA. Western Cape. Cape Town (3318): Stellenbosch Flats, (–DD), 17 October 1925, Duthie s.n.
STE1790 (NBG0197709-0! holo.; K000400569! iso.).
24. Thuranthos C.H.Wright in Bull. Misc. Inform. Kew 1916(9): 233 (1916) (Figs 60, 61). Typus generis:—T.
macranthum (Baker) C.H.Wright (holotype).
≡ Drimia sect. Thuranthos (C.H.Wright) J.C.Manning & Goldblatt in Strelitzia 40: 62 (2018) pro parte. Typus sectionis:—D. macrantha
(Baker) Baker (holotype).
Description:—Medium to tall bulbous geophyte. Bulb hypogeal, ovoid to subglobose, 3‒15 cm in diam., with white,
fleshy, spathulate and commonly pediculate, loose scales. Roots thickened and branched. Leaves 2‒7(‒14) per bulb,
hysteranthous, 10‒70 cm long, narrowly linear, channelled and usually keeled, green, sometimes basally maculate,
somewhat glaucous, smooth, glabrous. Inflorescence a lax, erect raceme, 15‒50 cm long, with 3–40 flowers; peduncle
30‒150 cm long, erect, terete, smooth; pedicels (10–)30‒70 mm long, subpatent and arching downwards distally
at anthesis, erect in fruit. Bracts ovate-lanceolate, 3‒10 mm long, lowermost with distinct spur, early caducous at
flowering and leaving a distinct scar; bracteoles present and evident, also early caducous. Flowers stellate with strongly
reflexed tepals at full anthesis, nodding, nocturnal, opening in late evening and closing before sunrise, fragrant during
night. Tepals 6, biseriate, (10‒)15‒37 mm long, narrowly oblong, obtuse, yellow, brown, carneous or whitish with
darker brownish-greenish longitudinal band more evident on abaxial side, almost free to very shortly connate for 1‒2
mm. Stamens 6, erect, sigmoid, yellow, brown to white or cream colour; filaments 7‒20 mm long, linear, flattened
and expanded below and incurved, leaving opening among them to show ovary, connivent to style at upper half/third,
and slightly spreading distally and commonly pale in colour, adnate to tepals at base; anthers oblong, 2‒7 mm long,
basifixed, yellow, orange or green, with yellow or white pollen. Ovary ovate to conical, 4.0‒6.5 mm long, greenish.
Style 7‒22 mm long, erect, white, columnar, ending in small or distinctly capitate, thickened stigma. Capsule ovate to
oblong or ellipsoid, trigonous, deeply 3-lobed, 10‒50 mm long, pale brown, valves completely dehiscing from base,
tepals cohering and inrolled above ovary after anthesis, circumcissile from base and persisting as a cap at the top of the
developing capsules. Seeds ellipsoidal to broadly ellipsoid, flattened, 6‒16 mm long, testa black, glossy, with slightly
sinuous anticlinal cell walls.
FIGURE 60. Species of Thuranthos C.H.Wright displayed in horizontal rows of images. 1. Thuranthos macranthum (Baker) C.H.Wright
from Wright (1916: t. 8680); 2. Thuranthos nocturnale R.A.Dyer from Dyer (1964: t. 1439).

FIGURE 61. Species of Thuranthos C.H.Wright displayed in horizontal rows of images. 1. Thuranthos bakeri Mart.-Azorín et al.; 2.
Thuranthos macranthum (Baker) C.H.Wright; 3. Thuranthos macrocarpum (Stedje) Speta (Photographs of flowers by B. Stedje); 4.
Thuranthos nocturnale R.A.Dyer; 5. Thuranthos pauciflorum (Baker) Mart.-Azorín et al.
Number of species and distribution:—Thuranthos includes eight species mostly confined to Southern Africa
and East Africa, being in the Uzambara-Zululand Region and the Southern and Eastern Sections of the Zambezian
Subregion, with a species occurring in the Karoo‐Namib Region (sensu Takhtajan 1986 and Martínez-Azorín et al.
2023a), (Fig. 49). For further species characterisation see Dyer (1964), Stirton (1976), Obermeyer (1980c), Stedje
(1987), Speta (1998a), Newton (2003), and Manning & Goldblatt (2018).
Karyology:—2n=20 (De Wet 1957, as Thuranthos macranthum (Baker) C.H.Wright); 2n=20+4B (Stedje 1996 as
Drimia macrocarpa Stedje).
History, diagnostic characters, and taxonomic relationships:—Wright (1916) described Thuranthos based on
a distinct species with a remarkable, unique flower morphology within Urgineoideae at that time, showing nocturnal,
large, nodding flowers with filaments that Stirton (1976) considered an outstanding feature showing flattened,
convex basal regions that are connivent to the style at the middle and spread above, resembling a paper lantern with
longitudinal slits (Figs 60, 61). Speta (1998a) accepted this genus based on its evident morphological differentiation
from other urgineoids. This genus was characterised by the imbricate, loose bulb scales; keeled, proteranthous leaves;
the presence of bracteoles; the nodding, nocturnal flowers with the peculiar morphology of filaments described above,
the erect style and the large, flat, ovate seeds.
The phylogenetic study of Martínez-Azorín et al. (2023a) included six samples of Thuranthos macranthum,
T. nocturnale Dyer (1964: t. 1439), and T. pauciflorum (Baker 1892: 6) Martínez-Azorín et al. (2019a: 295), the
latter including T. basuticum (Phillips 1917: 306) Obermeyer (1980c: 139) in synonymy (see Martínez-Azorín et al.
2019a). This is a genus restricted to Southern Africa and East Africa, and one which formed a strongly supported clade
included in a polytomy with the monophyletic and well supported Ledurgia and Zingela. Martínez-Azorín et al. (2023a)
considered Thuranthos to resemble Vera-duthiea on account of the nodding, nocturnal flowers with filaments usually
approaching the style at the middle, but differing in the style, which is distinctly deflexed, the spotted, not keeled
leaves, the lack of bracteoles [except in V. macrocarpa (Stedje 1987: 664) Speta (2016: 155)], and in their different
phylogenetic relationships, as it is sister to Ebertia (Martínez-Azorín et al. 2023a). Our recent unpublished phylogenetic
analyses of samples of Drimia macrocarpa Stedje (1987: 664) from Mapanza Mission in Zambia (Robinson 316 K!
number H.212/95 90; seeds kindly provided by Royal Botanic Gardens, Kew), show its close relation to Thuranthos
instead of Vera-duthiea. This recent finding facilitates the refinement of the circumscription of the latter two genera
and therefore, Thuranthos is expanded to include the species with loose bulb scales, nodding, nocturnal flowers with
basally expanded filaments that are sigmoid and connivent to the style along the middle portion and spreading above
and presence of bracteoles and Vera-duthiea, including species with compact bulb scales, absence of bracteoles and
filaments narrow and contracted at the joint to tepals, among other characters. This new circumscription of the two
genera eliminates the exception of the presence of bracteoles and loose bulb scales in D. macrocarpa when included
in Vera-duthiea by Speta (2001, 2016) and Martínez-Azorín et al. (2018b, 2023a).
Samples of Zingela consistently form a well‐supported clade sharing with Thuranthos the loose bulb scales;
hysteranthous leaves and presence of bracteoles; however, Zingela differs by the diurnal flowers and filiform,
spreading stamens with circinnate dehisced anthers (Crouch et al. 2018), and are placed in a polytomy with Thuranthos
and Ledurgia (Fig. 2 in Martínez-Azorín et al. 2023a), which is resolved in the parsimony analyses (Figs S5, S7 in
Martínez-Azorín et al. 2023a) where Ledurgia and Zingela form sister clades.
Therefore, we accept Thuranthos as a distinct genus following Stirton (1976), Obermeyer (1980c) and Speta
(1998a), though with the necessary recircumscription.
Manning & Goldblat (2018) accept D. sect. Thuranthos to include five species from Southern Africa: D. macrantha,
D. basutica, D. vespertina (a newly described species from northern Namibia and southern Angola), D. hesperantha (=
Urginea revoluta), and D. indica. As reported by Martínez-Azorín et al. (2018b), Drimia indica belongs to Indurgia,
a genus restricted to India and neigbouring countries, and plants identified as D. indica in South Africa were recently
described as Vera-duthiea zebrina Martínez-Azorín et al. (2018b: 285) and Zingela pooleyorum Crouch et al. (2018:
36). Similarly, D. vespertina seems to better fit the concept of Thuranthos of this work based on flower morphology
and the expanded filaments, though the compact bulb scales, the absence of bracteoles, the general small plant size
and its distribution makes its inclusion in the genus deviant. Further studies are needed to ascertain its phylogenetic
relationships. Finally, Urginea revoluta is excluded from Thuranthos based on flower and seed morphology and our
phylogenetic results (Martínez-Azorín et al. 2023a).
Accepted species and required new combination and new name:—

Thuranthos bakeri Mart.-Azorín, M.B.Crespo & M.Á.Alonso nom. nov. (Figs 5.4, 61.1) ≡ Drimia elata var. cooperi Baker in J. Linn.
Soc. 11: 422 (1871), syn. subst. ≡ Drimia angustifolia Baker, Fl. Cap. (Harvey) 6(3): 440 (1897), nom. illeg. [non Drimia angustifolia
Kunth, Enum. Pl. 4: 340 (1843), nec D. cooperi Baker, Refug. Bot. [Saunders] 1: pl. 18 (1869)]. Type:—SOUTH AFRICA. Eastern
Cape. Fort Beaufort (3226): near Tarkastad, (–AB), 1860, Cooper 387 (K000400568! holo.).
Thuranthos laxiflorum (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia laxiflora Baker in J. Linn. Soc., Bot.
11: 422 (1870), basionym. Type:—MOZAMBIQUE. Zambesi-land, prope Quillimane [Quelimane], November 1862, Dr. Kirk s.n.
(K0000400585! holo.).
Thuranthos macranthum (Baker) C.H.Wright in Bull. Misc. Inform. Kew 1916(9): 223 (1916) ≡ Ornithogalum macranthum Baker
in J. Linn. Soc., Bot. 13: 280 (1873), basionym ≡ Drimia macrantha (Baker) Baker in Bot. Jahrb. 15 (3, Beibl. 35): 7 (1892)
≡ Urginea macrantha (Baker) E.P.Phillips in Ann. S. African Mus. 16: 305 (1917) (Figs 60.1, 61.2). Type:—SOUTH AFRICA.
Eastern Cape. Uitenhage Division, Van Stadens River, elev. under 200 ft., December, Drège 2204 (K000257227! lecto. designated
here; K000257228!, S06-4414!, isolecto.). Note:—Lectotypification designated by Jessop in J. S. African Bot. 43: 275 (1977) is
disregarded since no specimen of that collection is kept at BM.
Thuranthos macrocarpum (Stedje) Speta in Phyton (Horn, Austria) 56(2): 155 (2016) ≡ Drimia macrocarpa Stedje in Nordic J. Bot. 7(6):
664 (1987), basionym ≡ Thuranthos macrocarpus (Stedje) Speta in Phyton (Horn, Austria) 38(1): 84 (1998) ≡ Duthiea macrocarpa
(Stedje) Speta in Stapfia 75: 170 (2001) (Figs 5.5, 61.3). Type:—TANZANIA. Mpanda District, Uruwira, elev. 3600 ft., 23 September
1970, H.M. Richards & S. Arasululu 26126 (K000257337! holo.).
= Ornithogalum laikipiense L.E.Newton in Plantsman n.s. 2(1): 18 (2003). Type:—KENYA. Laikipia Plateau, 56 km NE of Rumuruti,
Colcheccio Ranch, elev. 1750 m, April/May 1995, M. Roberts sub Newton 5567 (K000365655! Sheet 1/2 lecto. designated here;
K000365654! Sheet 2/2, isolecto.; EA isolecto., only the inflorescence). Note:—The type material at Kew includes two herbarium
vouchers with different herbarium barcodes. Although not specified, leaves and flowers were most probably collected at different
times (Hysteranthous: April–May as indicated on the voucher labels). Despite the current wording of Art. 8 of the ICN, and
following amendment proposal by Mosyakin & McNeill (2022), a lectotypification is here proposed. Further, the type material at
EA includes both leaves and inflorescences in a single voucher, and according to our proposal, only the inflorescence is regarded as
isolectotype.
Thuranthos nocturnale R.A.Dyer in Fl. Pl. Africa 36: t. 1439 (1964) (Figs 5.6, 60.2, 61.4). Type:—SOUTH AFRICA. Eastern Cape.
Steynsburg (3125): Grootfontein, (–AC), November 1956, Acocks 18650 (PRE0046805-2! lecto. designated here). Note:—The
type material (“holotype”) indicated in the protologue is currently conserved in four different herbarium sheets numbered 1–4,
here regarded as syntypes. Each of those vouchers includes different fragments collected on different dates. Following amendment
proposal of Art. 8 of the ICN by Mosyakin & McNeill (2022), a lectotypification is here proposed.
Thuranthos pauciflorum (Baker) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 295 (2019) ≡ Drimia pauciflora
Baker in Bot. Jahrb. Syst. 15(3, Beibl. 35): 6 (1892), basionym [non Urginea pauciflora Baker (1898: 539), nec Urginea pauciflora
Baker (1901: 786), nom. illeg.] (Figs 5.7, 61.5). Type:—SOUTH AFRICA. Pr. B. sp., Ecklon & Zeyher Asphod. 102 (B100167472!
lecto. [or perhaps holo.] designated by Martínez-Azorín et al. 2019a: 295).
= Urginea basutica E.Phillips in Ann. S. African Mus. 16: 306 (1917) ≡ Thuranthos basuticum (E.Phillips) Oberm. in Bothalia 13(1−2):
139 (1980) ≡ Drimia basutica (E.P.Phillips) J.C.Manning & Goldblatt in Strelitzia 40: 68 (2018). Type:—LESOTHO. Bethlehem
(2828): Leribe, (–CC), October–November 1911, A. Dieterlen 854 (PRE0048599-0! lecto. designated here; K!, SAM! isolecto.).
Note:—The holotype designated by Manning & Goldblatt (2018: 64) is not correct according to Art. 9.23 of the Shenzhen Code
(Turland et al. 2018), since two vouchers at K (K000257224 and K000257225) are undoubtedly original material.
Thuranthos vespertinum (J.C.Manning & Goldblatt) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia vespertina
J.C.Manning & Goldblatt in Strelitzia 40: 65 (2018), basionym. Type:—NAMIBIA. Sesfontein (1913): Khowarib Gorge, (–BD),
only the flowered material ex hort on 29 April 1991, P. Bruyns 4066 (NBG150472! holo.).
Thuranthos zambesiacum (Baker) Kativu in Kirkia 15(1): 113 (1994) ≡ Urginea zambesiaca Baker in J. Linn. Soc., Bot. 13: 223 (1873),
basionym ≡ Drimia zambesiaca (Baker) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004) ≡ Drimia zambesiaca
(Baker) J.C.Manning & Goldblatt in Bothalia 49(1): 4 (2019) isonym ≡ Vera-duthiea zambesiaca (Baker) N.R.Crouch & Mart.-
Azorín, Phytotaxa 470(3): 261 (2020). Type:—MOZAMBIQUE. Lower Zambesi, mainland above Expedition Island, lower Zambesi,
20 November 1859, J. Kirk 128 (K000400577! holo.).
25. Triandra Mart.-Azorín, M.B.Crespo, M.Á.Alonso, N.R.Crouch & M.Pinter in Phytotaxa 487(1): 66 (2021) (Fig.
62). Typus generis:—T. pellabergensis Mart.-Azorín, M.B.Crespo, M.Á.Alonso, N.R.Crouch & M.Pinter (holotype).
Description:—Small bulbous geophyte. Bulb hypogeal, ovoid to subglobose, 8‒18 mm in diam., with white, fleshy,
compact, inner scales and pale brown, membranous outer tunics. Roots thickened and branched. Leaves 2‒6 per
bulb, hysteranthous, 2‒8 cm long, filiform, to 1 mm in diam., subterete or very slightly flattened, green, smooth,
glabrous. Inflorescence a lax, erect raceme, 2‒5 cm long, with 4–8 flowers; peduncle 4‒8 cm long, brownish, erect,
terete, smooth; pedicels 7‒9 mm long, smooth, patent proximally and soon arching downwards distally at anthesis,
curving upwards in fruit. Bracts ovate-lanceolate, small, ca. 0.5 mm long, lowermost with short spur, ca. 0.3 mm long,
persistent; bracteoles absent. Flowers stellate, nodding, with strongly reflexed tepals at full anthesis with tips touching
flower pedicel, nocturnal, opening in late evening and closing before sunrise, fragrant during night. Tepals 6, biseriate,
5‒6 mm long, narrowly oblong, slightly constricted along middle, obtuse, whitish to pale brown, with darker brownish
longitudinal band more evident on abaxial side, almost free to very shortly connate for 0.5 mm. Stamens 3, only inner
whorl present, outer absent with no basal rudiments, erect, straight, approaching ovary and style, with 3 anthers placed
near stigma; filaments 3.5‒3.8 mm long, filiform, white, subterete, only slightly widened at base, connivent to style,
free, not adnate to tepals; anthers oblong, ca. 0.6 mm long after dehiscence, subbasifixed, yellow to orange, with
yellow polen. Ovary ovate, 1.9‒2.2 mm long, pale brown to yellowish. Style 2.0‒2.2 mm long, erect, white, filiform
and narrow, ending in undifferentiated stigma. Capsule ovate, trigonous, 4‒5 mm long, pale brown, valves completely
dehiscing from base, tepals cohering and inrolled above ovary after anthesis, circumcissile from base and persisting as
a cap at the top of the developing capsules. Seeds ellipsoidal to broadly ellipsoid, flattened, 6‒16 mm long, testa black,
glossy, with subpolygonal cells.
FIGURE 62. Triandra pellabergensis Mart.-Azorín et al.
Number of species and distribution:—A monotypic genus originally described from a single locality on the
Pella-se-Berge, northwestern South Africa, near to the border with Namibia, a locality included in the Karoo-Namib
Region (sensu Takhtajan 1986) (Fig. 49). Recently, Manning (2022) cited the collection Harrower 6188 (NBG) as the
second known locality for this species occurring in the Pienaarsfontein-se-Berge, Koedoesberge near Touwsrivier,
representing an extension of more than 400 km to the south, near the Tanqua Karoo in the Western Cape Province
of South Africa. Two additional populations were found, in September 2015 in the Naib se Berg and in August 2022
in the Nouroepberge near Dabenoris Mountains, both in vegetative stage, by M. Martínez-Azorín & collaborators
(MMA1269 ABH74121! and MMA2078 ABH74374! respectively). The phylogenetic analyses of these samples and
their recent flowering in cultivation confirmed their placement in Triandra.
Karyology:—Not studied yet.
History, diagnostic characters, and taxonomic relationships:—Our field work in Southern Africa led to the
discovery at the Pella-se-Berg (NW South Africa) of a small species of Urgineoideae with a remarkable morphology.

The plants produce nocturnal, nodding flowers with most notably only three stamens per flower, a character not
previously reported in Hyacinthaceae. This led Martínez-Azorín et al. (2021) to describe Triandra pellabergensis
Martínez-Azorín et al. (2021: 68)—a new genus and species. The phylogenetic studies by Martínez-Azorín et al.
(2023a) based on plastidial DNA sequences placed this species in an isolated position, sister to a clade comprising
samples of Rhodocodon from Madagascar. Therefore, we accept Triandra to accommodate this outstanding species in
the subfamily, which is supported by clear morphological, biogeographical, and molecular data (Martínez-Azorín et
al. 2023a).
Triandra pellabergensis Mart.-Azorín, M.B.Crespo, M.Á.Alonso, N.R.Crouch & M.Pinter in Phytotaxa 487: 68 (2021) ≡ Drimia
pellabergensis (Mart.-Azorín, M.B.Crespo, M.Á.Alonso, N.R.Crouch & M.Pinter) J.C.Manning & Goldblatt in Bothalia 52(1): 3
(2022) (Figs 5.8, 62). Type:—SOUTH AFRICA. Northern Cape. Pofadder (2919): Pella, Pella se Berge, ca. 25 km W of Pofadder,
rocky slopes at base of large south facing cliffs, (–AA), elev. 919 m, among quartz and mica rocks, fl. ex hort at University of
Alicante on 29 June 2018, M. Martínez-Azorín, M.B. Crespo, M.Á. Alonso et al. MMA1671b (GRA! holo.; ABH!, K, PRE iso.).
26. Urginavia Speta in Phyton (Horn, Austria) 38(1): 86 (1998) (Figs 63–66). Typus generis:—U. micrantha (A.Rich.)
Speta (holotype).
≡ Drimia sect. Urginavia (Speta) J.C.Manning & Goldblatt in Strelitzia 40: 13 (2018). Typus sectionis:—U. micrantha (A.Rich.) Speta
(holotype).
Description:—Usually tall bulbous geophyte. Bulb hypogeal, rarely epigeal, ovoid to globose, up to 20 cm in diam.,
scales somewhat loose, heterogeneous in length and vertically imbricate, yellowish and leathery when dry and showing
distinct white threads when broken, rarely showing fibrose neck above ground. Roots thickened and branched. Leaves
(2–)4‒16 per bulb, narrowly lanceolate to oblong, hysteranthous or rarely partially synanthous, 8‒60 cm long, plane
or channelled, sometimes keeled abaxially along lower portions, margin entire, green or somewhat glaucous, smooth,
glabrous, producing white threads when broken. Inflorescence a long, erect, usually multiflowered raceme, with
15‒700 flowers, (3–)10‒120 cm long; peduncle (6–)10‒100 cm long, terete, erect, smooth and glabrous; pedicels
2‒30 mm long, subpatent or spreading. Bracts narrowly lanceolate, 2‒15 mm long, usually caducous or sometimes
persisting, lowermost with spur from short to 20 mm long; bracteoles present and distinct. Flowers stellate or with
reflexed tepals, erect-patent, diurnal, lasting one or more days. Tepals 6, biseriate, 3.5‒12.0 mm long, mostly subfree
or rarely distinctly connate for 2‒3 mm at base, lanceolate-ovate to oblong, white, cream, brown or yellow, with darker
longitudinal band. Stamens 6, suberect to spreading; filaments 2‒10 mm long, smooth, usually white and lanceolate
to narrowly triangular, flattened or rarely filiform; anthers dorsifixed. Ovary ovate-oblong, 2‒5 mm long, green to
yellow. Style 2‒9 mm long, erect, white with trigonous undifferentiated or capitate stigma. Capsule obovoid, ellipsoid
or subglobose, 4‒15 mm long, emarginate, valves completely dehiscing from base, tepals remaining below developing
capsules. Seeds elliptic or ovate-oblong, 4‒9 mm long, flattened, with prominent embryo and flat peripheral wings,
testa black, with strongly sinuous anticlinal cell walls.
Number of species and distribution:—Urginavia includes 16 species from Southern, East and West Africa,
and is restricted to the Cape, Karoo-Namib, Uzambara-Zululand and Guineo-Congolian Regions, and the Zambezian,
Erithreo-Arabian and Sahelo-Sudanian Subregions (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a) (Fig. 49).
For further morphological species characterisation see Dyer (1942c, 1947), Jessop (1977), and Manning & Goldblatt
(2018).
Karyology:—2n=20 (De Wet 1957, Jones & Smith 1967, as Urginea altissima Baker; Stedje & Nordal 1987; De
Wet 1957, as Urginea multisetosa Baker; Stedje & Nordal 1987, as D. exigua Stedje); 2n=22 (Oyewole 1975, as U.
altissima; Oyewole 1975, as Urginea gigantea (Jacq.) Oyewole; Oyewole 1975, as Urginea viridula Baker); 2n=32
(De Wet 1957, as U. epigea R.A.Dyer); 2n=60 (Goldblatt et al. 2012, as D. capensis (Burm.f.) Wijnands).
History, diagnostic characters, and taxonomic relationships:—Speta (1998b) described Urginavia to include
plants with large bulbs; inflorescence commonly large, long racemose, multi-flowered; spurred bracts and distinct
bracteoles; patent pedicels and stellate flowers; tepals only slightly connate at base; nocturnal flowers (sensu Speta
1998b; doubtful); and flattened black seeds. The new genus was based on morphological and phylogenetic studies
(Pfosser & Speta 1999, 2001, 2004) and Speta (1998b) included in it Urginavia altissima (Linnaeus f. 1782: pl. 199)
Speta (1998b: 87), U. brachystachys (Baker 1892: 474) Speta (1998b: 87), U. porphyrantha (Bullock 1932: 505) Speta
(1998b: 87), U. micrantha (Richter 1850: 328) Speta (1998b: 87) (the type), and U. viridula (Baker 1898: 538) Speta
(1998b: 87). Later Speta (2001) added the combination Urginavia epigea (Dyer 1947: t. 1027) Speta (2001: 168). This
genus includes a distinct group of relatively large species occurring in Africa, south of the Sahara Desert (Speta 1998a).
However, we transfer Urginea brachystachys to Sekanama, based on the red bulb and translucent withered tepals,
among other characters, as discussed above.
FIGURE 63. Species of Urginavia Speta. 1. Urginavia capensis (Burm.f.) Mart.-Azorín et al. from Commelin (1701: t. 94, as Scilla
africana); 2. Urginavia altissima (L.f.) Speta from Jacquin (1797: t. 87, as Ornithogalum giganteum Jacq.); 3. Urginavia capensis (Burm.
f.) Mart.-Azorín et al. from Ker Gawler (1808: t. 1074, as Drimia altissima (L.f.) Ker Gawl.).

FIGURE 64. Species of Urginavia Speta. 1. Urginavia epigea (R.A.Dyer) Speta from Dyer (1947: t. 1027, as Urginea epigea R.A.Dyer);
2. Urginavia multisetosa (Baker) Mart.-Azorín et al. from Dyer (1942c: t. 854, as Urginea multisetosa Baker).
Manning & Goldblatt (2018) accepted this group of species as D. sect. Urginavia (Speta 1998b: 86) Manning
& Goldblatt (2018: 13) mostly following Speta’s (1998b) concept, but adding important characters such as well-
developed bracteoles, diurnal flowers, and tepals persisting below the developing capsule, not cohering into a cap
above. Their section included three species for Southern Africa: Drimia altissima (Linnaeus f. 1782: pl. 199) Ker
Gawler (1808: t. 1074), D. capensis (Burman 1768: 10) Wijnands (1983: 130), and D. kniphofioides (Baker 1897: 469)
Manning & Goldblatt (2003: 111).
Manning & Goldblatt (2018) placed Urginea multisetosa Baker (1897: 468) and U. echinostachya Baker (1897:
468) (= Ornithogalum cooperi Baker 1873b: 284) in their polyphyletic D. sect. Ledebouriopsis, and described the
absence of bracteoles in both species. Our phylogenetic results have confirmed that these two species belong to
Urginavia, further sharing the diagnostic characters of that genus, which include having distinct bracteoles, evident in
the type collections. The phylogenetic analyses of Martínez-Azorín et al. (2023a) include thirteen samples of Urginavia
from Central and Southern Africa and form a perfectly supported clade recognised at the rank of genus. This distinction
is morphologically substantiated by their bulb scales that are somewhat loose, heterogeneous in length and vertically
imbricate, usually yellowish and leathery when dry, and showing distinct white silky threads when broken (Fig. 65.2),
an apomorphy of the genus. Urginavia is sister to a clade including samples of Ledurgia, Thuranthos, and Zingela,
which mostly co-occur in distribution, but show very different morphology.
Pfosser & Speta (1999, 2001) included a sample of Urginavia micrantha (Richter 1850: 328) Speta (1998b: 87),
the type of the genus, in their phylogenetic analyses involving only the trnL-F region. We were unable to obtain further
sequences of other DNA regions for that same sample and accordingly excluded it from our broader analyses. However,
the inclusion of the published sample of U. micrantha in our trnL-F matrix confirmed its placement in Urginavia. The
taxonomy of the genus requires further resolution assisted by fieldwork across its extensive sub-Saharan range.
FIGURE 65. Species of Urginavia Speta displayed in horizontal rows of images. 1. Urginavia altissima (L.f.) Speta; 2. Urginavia cooperi
(Baker) Mart.-Azorín et al.; 3. Urginavia epigea (R.A.Dyer) Speta.
Urginavia altissima (L.f.) Speta in Phyton (Horn, Austria) 38(1): 87 (1998) ≡ Ornithogalum altissimum L.f., Suppl. Pl.: 199 (1782),
basionym ≡ Drimia altissima (L.f.) Ker Gawl. in Curtis’s Bot. Mag. 27: t. 1074 (1808) ≡ Urginea altissima (L.f.) Baker in J. Linn.
Soc., Bot. 13: 221 (1873) ≡ Idothea altissima (L.f.) Kuntze, Revis. Gen. Pl. 2: 712 (1891) (Figs 5.9, 63.2, 65.1). Type:—SOUTH
AFRICA. Eastern Cape, Cap. Bona Spei, Zondags River, without date, Thunberg s.n. (UPS-THUNB 8275, lecto. designated as
“Type” by Jessop in J. S. African Bot. 43: 288. 1977).
= Ornithogalum giganteum Jacq., Hort. Schoenbr. 1: 45, t. 87 (1797) (Fig. 63.2) ≡ Urginea gigantea (Jacq.) Oyewole in Bol. Soc. Brot.,
sér. 2, 49: 167 (1975). Type:—illustration in Jacq., Hort. Schoenbr. 1: t. 87 (1797) (lecto. designated as “Type” by Jessop in J. S.
African Bot. 43: 288. 1977).
= Drimia uitenhagensis Eckl. in S. Afr. Quart. Journ. 1: 364 (1830). Type:—SOUTH AFRICA. Eastern Cape. Port Elizabeth (3325): on the
hills of Adow [Addo], District Uitenhage, (–DA), Ecklon 17 (S10-14097! lecto. designated by Manning & Goldblatt 2018).
Urginavia barteri (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia barteri Baker in J. Linn. Soc., Bot. 11: 423
(1870), basionym. Type:—NIGERIA. Niger territory, in ditione nigritana prope Nupe, Barter 1183 (K000257327 holo!).
Urginavia capensis (Burm.f.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Scilla capensis Burm.f., Prod. Fl. Cap.: 10 (1768),
basionym ≡ Drimia capensis (Burm.f.) Wijnands, The Botany of the Commelins: 130 (1983) (Figs 5.10, 63.1, 63.3). Type:—
illustration in Commelin, Hort. Amst. 2: 187 t. 94 (1701) (Fig. 63.1) (lecto. designated by Wijnands in The Botany of the Commelins:
130. 1983).
= Urginea forsteri Baker, Fl. Cap. (Harvey) 6(3): 469 (1897) ≡ Drimia forsteri (Baker) Oberm. in Bothalia 13: 453 (1980). Type:—SOUTH
AFRICA. Without locality or date, Forster s.n. (K! lecto. effectively designated by Obermeyer 1981b: 453).

FIGURE 66. Species of Urginavia Speta displayed in horizontal rows of images. 1. Urginavia kniphofioides (Baker) Mart.-Azorín et al.
(Photographs by D. Oosthuizen); 2. Urginavia multisetosa (Baker) Mart.-Azorín et al.; 3. Urginavia simensis (Hochst. ex A.Rich.) Mart.-
Azorín et al. (Photographs by B. Stedje).
Urginavia congesta (Bullock) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia congesta Bullock in Bull. Misc. Inform.
Kew 1932(10): 504 (1932), basionym [non Drimia congesta (Wight) Stearn in Ann. Mus. Goulandris 4: 208 (1978)] ≡ Urginea
congesta Wight, Icon. Pl. Ind. Orient. 6: 28, t. 2064 (left hand figure) (Wight 1853)]. Type:—KENYA. Mt. Elgon, elev. 8700 ft., 06
January 1931, E.J. Major & C. Lugard 474 (K000257334! lecto. designated as “holo.” by Stedje in Fl. Trop. E. Africa, Hyacinth.:
17. 1996; EA isolecto.).
Urginavia cooperi (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Ornithogalum cooperi Baker in J. Linn. Soc., Bot. 13:
284 (1873), basionym [non Ornithogalum cooperi (Baker) J.C.Manning & Goldblatt in Edinburgh J. Bot. 60(3): 547 (2004), nom.
illeg.] ≡ Drimia cooperi (Baker) Baker, Fl. Cap. (Harvey) 6: 443 (1897), nom. illeg., [non D. cooperi Baker, Refug. Bot. [Saunders]
1: pl. 18 (1869) = Ledebouria concolor (Baker) Jessop in J. S. African Bot. 36: 254. 1970] (Fig. 65.2). Type:—SOUTH AFRICA.
Eastern Cape, ad oram orientalis, Barber s.n. (TCD lecto. designated by Jessop in J. S. African Bot. 43: 287. 1977; K000257356!
isolecto.).
Urginavia echinostachya (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea echinostachya Baker, Fl. Cap.
(Harvey) 6(3): 468 (1897), basionym ≡ Drimia echinostachya (Baker) Eggli & N.R.Crouch in Bothalia 42(1): 43 (2012) (Fig.
5.11). Type:—SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): Inanda, (–DB), J.M. Wood 276 (K000257349-1! lecto.
designated here; NH0004369! isolecto.). Note:—Both vouchers at K and NH are original material and therefore holotype indication
by Crouch et al. (2010) is disregarded.
Urginavia epigea (R.A.Dyer) Speta in Stapfia 75: 168 (2001) ≡ Urginea epigea R.A.Dyer in Fl. Pl. Africa 26: t. 1027 (1947) (Figs 64.1,
65.3). Type:—SOUTH AFRICA. Mpumalanga, Transvaal, Middelburg Dist., N of Zoetvelden, Van der Merwe 2203 (PRE0048542-
1! lecto. designated by Martínez-Azorín et al. in Phytotaxa 201: 169. 2015: the inflorescence only).
Urginavia exigua (Stedje) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia exigua Stedje in Nordic J. Bot. 14(1): 43
(1994), basionym. Type:—ETHIOPIA. Guder River valley, Nordal 1015 (O holo.).
Urginavia kniphofioides (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea kniphofioides Baker, Fl. Cap. (Harvey)
6(3): 469 (1897), basionym ≡ Drimia kniphofioides (Baker) J.C.Manning & Goldblatt in Bothalia 33(1): 111 (2003) (Figs 5.12,
66.1). Type:—SOUTH AFRICA. Mpumalanga. Komatipoort (2531): Transvaal, Havelock Concession, (–CC), elev. 3200–5000
ft., September 1890, E. Saltmarshe sub Galpin 1055 (K000257361! lecto. designated here; BOL140330! isolecto.). Note:—Both
materials at K and BOL are here considered to be original material and therefore a lectotype designation is needed.
Urginavia micrantha (A.Rich.) Speta in Phyton (Horn, Austria) 38(1): 87 (1998) ≡ Scilla micrantha A.Rich., Tent. Fl. Abyss. 2: 328
(1850) ≡ Urginea micrantha (A.Rich.) Solms, Schweinfurth, Beitr. Fl. Aeth.: 294 (1867) (Fig. 5.13). Type:—ETHIOPIA. Abyssinia,
Chiré, Quartin Dillon & Petit s.n. (P02157062! lecto. designated as “holotype” by Stedje & Thulin in Nordic J. Bot. 15: 598. 1995;
P02157060!, WU081571! isolecto.).
= Drimia paolii Chiov. in Res. Sci. Somalia Ital. 1: 175 (1916), basionym. Type:—SOMALIA. El Ualàc, 3 November 1913, Paoli 1096
(FT000782! holo.).
Urginavia multisetosa (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea multisetosa Baker, Fl. Cap. (Harvey)
6(3): 468 (1897), basionym ≡ Drimia multisetosa (Baker) Jessop in J. S. African Bot. 43(4): 299 (1977) (Figs 5.14, 64.2, 66.2).
Type:—SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): Natal, near Mooi River, (–AA), elev. 4000−5000 ft., October
1894, J. Medley Wood 5724 (K000400572! lecto. designated by Jessop in J. S. African Bot. 43: 299. 1977).
Urginavia porphyrantha (Bullock) Speta in Phyton (Horn, Austria) 38(1): 87 (1998) ≡ Urginea porphyrantha Bullock in Bull. Misc.
Inform. Kew 1932(10): 505 (1932) ≡ Drimia porphyrantha (Bullock) Stedje in Nordic J. Bot. 7(6): 665 (1987). Type:—KENYA.
Mt. Elgon, elev. 6500 ft., 16 March 1931, E.J. Major & C. Lugard 556 (K000257338! lecto. designated as “holotype” by Stedje
1987: 665; EA isolecto.).
Urginavia psilostachya (Welw. ex Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea psilostachya Welw. ex Baker
in Trans. Linn. Soc. London, Bot. 1(5): 247 (1878), basionym ≡ Drimia psilostachya (Welw. ex Baker) J.C.Manning & Goldblatt
in Bothalia 43(1): 77 (2013). Type:—ANGOLA. Cazengo, in dumetis siccis arenosis, Welwitsch 3807 (BM000911784! lecto.
designated here; K000400575 isolecto.).
Urginavia riparia (Baker) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea riparia Baker, Fl. Cap. (Harvey) 6: 467
(1897), basionym. Type:—SOUTH AFRICA. KwaZulu-Natal. Pietermaritzburg (2930): Umzinyati Falls, (–DB), 11 September
1880, J. Medley Wood 1052 (K000099097! lecto. designated by Jessop in J. S. African Bot. 43: 303. 1977). Note:—The collection
Wood 1052 is a mixed collection. On the one hand it includes a voucher (K000099097) fitting the protologue of U. riparia Baker,
which was selected as lectotype of that name by Jessop (1977). On the other hand, the voucher NH0004370 corresponds to Drimia
elata s.l. and is not to be regarded as original material since it does not match the protologue.
Urginavia simensis (Hochst. ex A.Rich.) Mart.-Azorín, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Scilla simensis Hochst. ex A.Rich.,
Tent. Fl. Abyss. 2: 327 (1850), basionym ≡ Drimia simensis (Hochst. ex A.Rich.) Stedje in Nordic J. Bot. 15(6): 597 (1996) ≡
Charybdis simensis (Hochst. ex A.Rich.) Speta in Phyton (Horn, Austria) 38(1): 60 (1998), nom. illeg. [Art. 52 of the ICN] (Figs
5.15, 66.3). Type:—ETHIOPIA. Ad latus meridionale montium simensium prope Abbu Mekanna et Debra Dschoa, 2 March 1840,
W. Schimper 1317 (P02057842! lecto. designated by Martínez-Azorín et al. 2022: only the bulb on the right side of the sheet bearing
a scape with flowers and fruits, and the bulb in the affixed envelope; K000257332!, K000257333!, L1459048!, BR0000090026478!,
M0107240!, P02057845!, TUB006926! isolecto.).
Urginavia viridula (Baker) Speta in Phyton (Horn, Austria) 38(1): 87 (1998) ≡ Urginea viridula Baker, Fl. Trop. Afr. [Oliver et al.] 7(3):
538 (1898), basionym ≡ Drimia viridula (Baker) J.C.Manning & Goldblatt in Bothalia 43(1): 78 (2013). Type:—Lower Guinea.
Congo, cultivated specimen, 10 September 1886, ex hort., Bull s.n. (K000257329! holo.).
27. Urginea Steinh. in Ann. Sci. Nat., Bot. sér. 2, 1: 321, t. 14 (1834) (Figs 67–68). Typus generis:—U. fugax (Moris)
Steinh. (lectotype designated by Adamson 1942).
= Urginia Kunth, Enum. Pl. 4: 331 (1843) (orth. var. sensu Speta 1998).

FIGURE 67. Urginea fugax (Moris) Steinh. from: 1. Steinhel (1834: t. 14); 2. Cusin (1876: pl. 23bis).
FIGURE 68. Urginea fugax (Moris) Steinh.
Description:—Small bulbous geophyte. Bulb hypogeal, ovoid to subglobose, up to 3.5 cm in diam., inner scales
compact, white, fleshy, outer scales pale brown, membranous. Roots thickened and branched. Leaves 4‒13 per bulb,
suberect, filiform, narrowly linear, hysteranthous (sometimes leaves present at anthesis but mostly withered), 12‒40
cm long and to 2 mm wide, subterete or hemiterete, channelled, entire, green or somewhat glaucous, smooth, glabrous.
Inflorescence an elongated, erect, lax raceme, with 5‒18 flowers, 1‒15 cm long; peduncle 8‒18 cm long, terete, erect,
smooth and glabrous, usually flexuose below; pedicels 5‒9 mm long, suberect. Bracts narrowly lanceolate, small, ca.
1 mm long, caducous, lowermost with short spur; bracteoles absent. Flowers stellate, erect-patent, diurnal. Tepals 6,
biseriate, 8‒10 mm long, free to very shortly connate at base, obovate-lanceolate, white with purple central band more
evident on abaxial side. Stamens 6, spreading; filaments 6‒7 mm long, filiform, smooth, white, somewhat flattened
below, very shortly adnate at base of tepals; anthers dorsifixed, with yellow pollen. Ovary ovate-oblong, 2‒3 mm long,
green to yellowish. Style 5‒6 mm long, erect, white with slightly capitate stigma. Capsule ovoid-ellipsoid, 7‒10 mm
long, valves completely dehiscing from base, tepals cohering and inrolled above ovary after anthesis, circumcissile
from base and persisting as a cap at the top of the developing capsules. Seeds elliptic or ovate-oblong, 5‒7 mm long,
flattened, with prominent embryo and flat peripheral wings, testa black, glossy, with strongly sinuous anticlinal cell
walls.
Number of species and distribution:—Urginea includes 2 species from the western Mediterranean Basin, and
hence restricted to the Mediterranean Region (sensu Takhtajan 1986) (Fig. 49). For further species characterisation see
Maire (1938) and Rico (2013).
Karyology:—2n=20 (Talavera et al. 1995, as Urginea fugax (Moris) Steinh. var. fugax; Talavera et al. 1995, as
U. fugax var. major Litard. & Maire ex Maire); 2n=20+0–8B (Battaglia & Guanti 1968, as Urginea fugax (Moris)
Steinh.).
History, diagnostic characters, and taxonomic relationships:—Steinheil (1834) described Urginea to include
seven species with almost free tepals and variable morphology and distribution, among them including taxa currently
placed in Tenicroa and Squilla (= Charybdis, nom. illeg.). Adamson (1942) first typified the genus by U. fugax, a
species restricted to the western Mediterranean Basin. Maire (1938) described U. ollivieri, a related species that,
together with U. fugax, seem to form a consistent group with filiform leaves, diurnal stellate, patent to suberect flowers
with free tepals and spreading filaments.
The phylogenetic study by Martínez-Azorín et al. (2023a) included four samples of Urginea fugax and U. ollivieri
from Sardinia (Italy), Ibiza (Spain), and Morocco, that form a strongly supported clade. This clade is sister to a clade
comprising six samples of Urginea noctiflora from Morocco, here accepted as Spirophyllos. Therefore, we here accept
Urginea to include Urginea fugax and U. ollivieri, which share similar vegetative and reproductive characters and
share the same evolutive lineage.
Urginea fugax (Moris) Steinh. in Ann. Sci. Nat., Bot. sér. 2, 1: 328 (1834) ≡ Anthericum fugax Moris, Stirp. Sard. Elench. 1: 46 (1827),
basionym ≡ Scilla fugax (Moris) Munby, Fl. Alger.: 36 (1847) ≡ Urginea fugax var. typica Litard. & Maire in Mém. Soc. Sci.
Nat. Maroc. 26: 35 (1931), nom. inval. ≡ Drimia fugax (Moris) Stearn in Ann. Mus. Goulandris 4: 204 (1978) (Figs 5.16, 67–68).
Type:—ITALY. Sardinia, M. Moris s.n. (P02166433! lecto. designated here: we consider all material in the voucher to belong to a
single gathering, although not specified on the label).
= Urginea fugax (Moris) Steinh. var. major Litard. & Maire in Mém. Soc. Sci. Nat. Maroc. 26: 35−36 (1931). Type:—MOROCCO. In
silva Mamora, prope Kenitra, 28 July 1824, R. Maire s.n. (MPU002403! lecto. designated here; P00083301! isolecto.).
Urginea ollivieri Maire in Bull. Soc. Hist. Nat. Afrique N. 29: 453–454 (1938) ≡ Drimia ollivieri (Maire) Stearn in Ann. Mus. Goulandris
4: 208 (1978). Type:—MOROCCO. Montagnes au dessus de Tougoumaït entre Assa et Goulimine, September 1937, Y. Ollivier 68
(MPU003972! lecto. designated here; MPU003973! syn.).
28. Urgineopsis Compton in J. Bot. 68: 107 (1930) (Figs 69–70). Typus generis:—U. salteri Compton (holotype).
Description:—Small bulbous geophyte. Bulb hypogeal, mostly solitary, inner scales white to pinkish, fleshy, compact,
outer scales membranous. Roots thickened and branched. Leaf usually solitary (rarely to 3 per bulb), 4‒30 cm long,
terete, dark green, commonly leathery, hysteranthous or rarely synanthous, deciduous, to 3 mm in diam., smooth,
glabrous. Inflorescence a usually elongated, dense to lax raceme, 0.5‒4.0 cm long, erect; peduncle elongated, 6‒30
cm long, erect, smooth or sometimes striate-papillate at base; pedicels 0.5‒6.0 mm long, subpatent. Bracts lanceolate,
acute, short, 0.5‒1.5 mm long, lowermost with short spur to 5 mm long; bracteoles absent, exceptionally present and
minute. Flowers stellate, erect-patent, diurnal. Tepals 6, 5‒8 mm long, connate in distinct widely campanulate to
subcylindrical tube for 1‒4 mm, free lobes patent to slightly reflexed at full anthesis, whitish, yellowish, or purplish,
with darker central band more evident on abaxial side. Stamens 6, spreading and incurved distally; filaments adnate to
tepals and arising from mouth of tepals tube, free portions ca. 2 mm long, white, glabrous, smooth, spreading basally
and incurved distally; anthers yellow, oblong, medifixed, dehiscing along their whole length. Ovary ovate to narrowly
oblong, 2.0‒2.5 mm long, attenuate to truncate at top, green to yellowish; style white, narrowly columnar, 2.0‒2.5
mm long; stigma slightly 3-lobed and papillose. Capsule ovate to subglobose, 5‒12 mm long, trigonous, loculicidal,
3 valves splitting to base, with withered perigone segments circumscissile below and forming an apical cap. Seeds
ellipsoid to fusiform or pyriform, 2‒6 mm long, light brown to black, with striate to reticulate testa.
Number of species and distribution:—Urgineopsis includes seven species from coastal areas in western South
Africa and western Namibia, and restricted to the Cape and Karoo-Namib Regions (sensu Takhtajan 1986) (Fig. 71).
For further species characterisation see Duthie (1928), Compton (1930), Adamson (1942), Adamson et al. (1944),
Jessop (1977), and Manning & Goldblatt (2018).
Karyology:—2n=14 (Speta 1998a with no reference).
History, diagnostic characters, and taxonomic relationships:—Compton (1930) described Urgineopsis based
on a single species, U. salteri Compton (1930: 107) from the Cape Peninsula in southwestern South Africa. This species
shows a peculiar flower morphology in which tepals are connate for ca. 2 mm at the base forming a campanulate,
widely open tube; filaments adnate to tepals arising at the apex of tepal tube, spreading and slightly incurved, and the
distinctly exposed ovary due to the very wide tepal tube and spreading stamens. Jessop (1977) reduced Urgineopsis
to the synonymy of a widely circumscribed Drimia, arguing that the degree of fusion of tepals is not a consistent
character defining genera in Urgineoideae, in which a transition of connation degrees is observed. However, as noted
above, this character alone should not be used for generic circumscription in Urgineoideae due to a certain degree of
homoplasy, and therefore a combination of other morphological and phylogenetic characters are considered to support
the acceptance of Urgineopsis. Baker (1897), Duthie (1928), Adamson (1942), Adamson et al. (1944), and Manning
& Goldblatt (2018) have described further taxa occurring in coastal areas of western South Africa, which share the
same distinct flower morphology of Urgineopsis. Martínez-Azorín et al. (2019a) recently accepted seven species in
Urgineopsis to extend the range of this genus to coastal Namibia.
FIGURE 69. Urgineopsis dregei (Baker) Mart.-Azorín et al. (1–5) and Urgineopsis gracilis (A.V.Duthie) Mart.-Azorín et al. (6–14) from
Duthie (1928: pl. 1, as Urginea dregei Baker and Urginea gracilis A.V.Duthie, respectively).
FIGURE 70. Species of Urgineopsis Compton displayed in horizontal rows of images. 1. Urgineopsis barbata (J.C.Manning &
J.M.J.Deacon) Mart.-Azorín et al.; 2. Urgineopsis salteri Compton.
The phylogenetic studies by Martínez-Azorín et al. (2023a) included six samples of Urgineopsis that formed
a strongly supported clade placed within a large polytomy. Based on the clear morphological and phylogenetic
evidence, we here accept Urgineopsis as a genus endemic to western South Africa and Namibia. The taxonomy of
this genus requires further study, especially on fresh material, to evaluate the real status of some described species and
populations.
Urgineopsis arenosa (Adamson) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 295 (2019) ≡ Urginea arenosa
Adamson in J. S. African Bot. 8: 239 (1942), basionym (Fig. 5.17). Type:—SOUTH AFRICA. Western Cape. Simonstown (3418):
Cape Peninsula, Red Hill, (–AB), Salter 8321 (BOL140320! holo.; NBG0073244!, K000257365!, SAM0056687! iso.).
Urgineopsis barbata (J.C.Manning & J.M.J.Deacon) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 295 (2019) ≡
Drimia barbata J.C.Manning & J.M.J.Deacon in Strelitzia 40: 47 (2018), basionym (Fig. 70.1). Type:—SOUTH AFRICA. Northern
Cape. Oranjemund (2816): Boegoeberg, S of Alexander Bay, (–DA), only materials in flower ex hort. on 1 February 1980, M. Bayer
1013 (NBG121176! holo. [2 cross referenced sheets following Art. 8.2 Ex. 4 of the ICN, Turland et al. (2018)]).
Urgineopsis dregei (Baker) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 295 (2019) ≡ Urginea dregei Baker,
Fl. Cap. (Harvey) 6(3): 467 (1897), basionym (Figs 5.18, 69.1–5). Type:—SOUTH AFRICA. Western Cape. Worcester (3319):
Worcester Div., Dutoitskloof, (–CC), 1840, Drège 1501 (K000257344! lecto. designated by Manning & Goldblatt 2018; S06-4415!
[only the four fruiting specimens] isolecto.).
Urgineopsis exilis (Adamson) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 295 (2019) ≡ Urginea exilis
Adamson in J. S. African Bot. 8: 240 (1942), basionym (Fig. 5.19). Type:—SOUTH AFRICA. Western Cape. Simonstown (3418):
Cape Peninsula, De Klip, (–AD), February 1940, Salter 8312 (BOL140317! holo.; NBG0073245!, SAM0056685! iso.).
Urgineopsis gracilis (A.V.Duthie) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 295 (2019) ≡ Urginea gracilis
A.V.Duthie in Ann. Univ. Stellenbosch 1928, 6: Sect. A, No. 2, 12 (1928), basionym ≡ Fusifilum gracilis (A.V.Duthie) Speta in Phyton
(Horn, Austria) 38(1): 69 (1998) (Figs 5.20, 69.6–14). Type:—SOUTH AFRICA. Western Cape. Cape Town (3318): ‘Stellenbosch
flats, in low-lying, clayey area’, (–DD), March 1925, A. Duthie 1446a (NBG0197705-1! lecto. designated here: only the flowering
fragments, excluding both the leafing plants and inflorescences with dehisced capsules). Note:—The lectotypification by Manning
& Goldblatt (2018) is to be superseded, because the supposed holotype includes fragments from different collections.

Urgineopsis pedunculata (Adamson) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 296 (2019) ≡ Urginea
pedunculata Adamson in J. S. African Bot. 10: 134 (1944), basionym. Type:—SOUTH AFRICA. Western Cape. Simonstown (3418):
Cape Peninsula, Kalk Bay Mountains, (–AB), M.R. Levyns sub Adamson 3483 (BOL140321! holo.).
Urgineopsis salteri Compton in J. Bot. 68: 107 (1930) ≡ Urginea salteri (Compton) Adamson in J. S. African Bot. 8: 238 (1942), in obs. ≡
Drimia salteri (Compton) J.C.Manning & Goldblatt in Strelitzia 9: 712 (2000) (Figs 6.1, 70.2). Type:—SOUTH AFRICA. Western
Cape. Simonstown (3418): Cape Peninsula, Muizenburg on rock slabs, (–AB), Salter s.n. BOL18861 (BOL140319! lecto. designated
by Jessop in J. S. African Bot. 43: 303. 1977; K000257370! isolecto.).
29. Vera-duthiea Speta in Phyton (Horn, Austria) 56(2): 154 (2016). Typus generis:—V. senegalensis (Kunth) Speta
(holotype).
≡ Duthiea Speta in Stapfia 75: 170 (2001) [repl. syn.], nom. illeg., non Duthiea Hack. in Verh. Zool.-Bot. Ges. Wien 45: 200 (1896)
[Poaceae], nec Duthiea Manza in Proc. Natl. Acad. U.S.A. 23: 48 (1937), nom. illeg. [Corallinaceae, Rhodophyta] (Figs 72–73).
Typus generis:—D. senegalensis (Kunth) Speta (holotype).
Description:—Medium sized bulbous geophyte. Bulb hypogeal, ovoid to subglobose, 3‒6 cm in diam., with white,
fleshy, compact inner scales and membranose outer tunics. Roots thickened and branched. Leaves 2‒10 per bulb,
hysteranthous or synanthous, 6‒30 cm long, narrowly linear to lanceolate, subterete to plane or canaliculated, green,
somewhat glaucous, smooth, glabrous, commonly with darker maculations along basal portions, sometimes giving a
zebra banding pattern. Inflorescence a lax, erect raceme, 3‒35 cm long, with 3–30 flowers; peduncle 12‒60 cm long,
erect to leaning, terete, smooth, sometimes slightly mottled at base; pedicels 3‒40 mm long, subpatent proximally and
arching downwards distally at anthesis, erect in fruit. Bracts deltoid, short, 2‒5 mm long, lowermost with short spur,
caducous and leaving distinct scar; bracteoles absent. Flowers stellate with strongly reflexed tepals at full anthesis,
nodding, nocturnal, opening in late evening and closing before sunrise, fragrant during night. Tepals 6, biseriate,
10‒15 mm long, narrowly oblong to lanceolate, yellow, brown, carneous or whitish with darker brownish-greenish
longitudinal band more evident on abaxial side, almost free to shortly connate for 1‒2 mm. Stamens 6, erect, sigmoid,
yellow, brown to white or cream colour; filaments 8‒12 mm long, linear and subterete, narrowed below and incurved
approaching ovary, connivent to style along middle, and spreading distally, very shortly adnate to tepals at base; anthers
oblong, 2‒3 mm long, subbasifixed, yellow, orange or green, with yellow or white pollen. Ovary ovoid to conical, 3‒5
mm long, greenish. Style 6‒8 mm long, erect or deflexed to approach stamens spreading distally, white, columnar, with
small stigma. Capsule ovate, oblong or ellipsoid, trigonous, deeply 3-lobed with sharp angles, 9‒20 mm long, pale
brown, valves completely dehiscing from base, tepals cohering and inrolled above ovary after anthesis, circumcissile
from base and persisting as a cap at the top of the developing capsules. Seeds ellipsoidal to broadly ellipsoid, flattened,
6‒10 mm long, testa black, glossy, with sinuous anticlinal cell walls.
Number of species and distribution:—Vera-duthiea includes 6 species occurring mainly in Southern and Central
Africa, and extending north to the Arabian Peninsula in the Dhofar Mountains in Oman and southern Yemen (Patzelt et
al. 2021). It is therefore restricted to the Uzambara‐Zululand, Karoo‐Namib and Guineo‐Congolian Regions, and the
Zambezian, Erithraeo‐Arabian, Omano‐Sindian and Sahelo‐Sudanian Subregions (sensu Takhtajan 1986 and Martínez-
Azorín et al. 2023a) (Fig. 71). The study of herbarium material of this genus deposited at P, and other herbaria from
most countries in Central Africa, showed a large variability in morphology regarding size and morphology of bulbs,
leaves, and flowers. Therefore, further studies based on fresh material are urgently needed to ascertain the real diversity
of this genus. For further information on Vera-duthiea species see Martínez-Azorín et al. (2018b, 2019a), Manning &
Goldblatt (2018), Crouch et al. (2020), and Patzelt et al. (2021).
History, diagnostic characters, and taxonomic relationships:—The morphological and genetic studies by
Pfosser & Speta (2001) led Speta (2001) to describe Duthiea to include three species from Central to northwestern
Africa: D. senegalensis (Kunth 1843: 334) Speta (2001: 170), the type of the genus, D. macrocarpa (Stedje 1987: 664)
Speta (2001: 170), and D. noctiflora (Battandier & Trabut 1894: 505) Speta (2001: 170). This genus was characterised
by the imbricate bulb scales, proteranthous leaves, patent pedicels, perigone slightly connate at the base, nocturnal
flowers, white and thin filaments, style broadened at apex, ovoid ovary, and flattened and black seeds (Speta 2001).
However, Duthiea Speta is illegitimate due to the later homonym of both the previous Duthiea Hackel (1896: 200)
(Poaceae) and Duthiea Manza (1937: 48), nom. illeg. (Corallinaceae). To solve this, Speta (2016) published Vera-
duthiea as a “nomen novum” for his illegitimate Duthiea and transferred to his new genus the three above cited species.
Martínez-Azorín et al. (2018b) described V. zebrina Mart.-Azorín et al. (2018b: 285) representing the extension of the

distribution range of this genus to eastern Southern Africa. Manning & Goldblatt (2018) described Drimia vespertina
for plants occurring in Kaokoland in northwestern Namibia and Martínez-Azorín et al. (2019a) effected five further
new combinations in Vera-duthiea including V. vespertina and other taxa described from Namibia and Senegal. Patzelt
et al. (2021) described V. capillifolia Patzelt et al. (2021: 46) from Oman, being the first record of the genus outside
of Africa. Drimia vespertina is provisionally placed in Thuranthos, as discused above, based on its basally expanded
filaments.
FIGURE 72. Vera-duthiea zebrina Mart.-Azorín et al. from: 1. Line illustration by R. Collett based on Mauve 4823 (PRE); 2. Watercolour
illustration by Elsa Pooley based on Pooley 127 (NU).
FIGURE 73. Species of Vera-duthiea Speta displayed in horizontal rows of images. 1. Vera-duthiea capillifolia A.Patzel et al.; 2. Vera-
duthiea zebrina Mart.-Azorín et al.
The phylogenetic results of Martínez-Azorín et al. (2023a) include seven samples of Vera-duthiea from South
Africa, Senegal, and Oman, which form a strongly supported clade, confirming the phylogenetic results of Pfosser
& Speta (2001, 2004). Vera-duthiea is sister to Ebertia from Guinea and the clade combining both these genera is
related to Iosanthus, Indurgia, Urginea, and Spirophyllos. Our results agree with those of Pfosser et al. (2006), whose
samples of Urginea noctiflora resolved sister to Urginea s.str., and therefore are not included in Vera-duthiea. While
describing his new genus, Speta (2001) also included Urginea noctiflora based on the nodding, nocturnal flowers.
This species, however, is separated here into the new monotypic genus Spirophyllos. The exclusion of U. noctiflora
from Vera-duthiea is also supported by the lack of the maculate leaves typical of that genus, the straight filaments
that are connivent to the style and cross each along the upper third, and by its distinctly coiled leaves, unique in
Urgineoideae.
Here we accept Vera-duthiea as a genus occurring in Southern and Central Africa extending to the Arabian
Peninsula, characterised by the hypogeal, compact bulb scales; leaves usually maculate, at least at base; lax racemes;
the absence of bracteoles; nodding, nocturnal flowers; free to shortly connate, reflexed tepals; filaments narrow and
constricted at the joint to the tepals that are shortly adnate, incurved along the lower half, almost connivent to style
in middle section and spreading distally; anthers near basifixed; style suberect to distinctly deflexed at full anthesis;
subcapitate stigma; and the flattened, subelliptic seed with prominent central embryo. Other genera with superficially
similar flowers are Indurgia, Thuranthos, and Zingela, with which it might be confused. However, Thuranthos shows
bulbs with very loose scales; bracteoles present and distinct, and filaments expanded along the basal half. The members
of Indurgia are restricted in range to southwestern Asia, and also differ by their unspotted leaves; straight, erect to
suberect filaments; erect, usually thickened, subclavate style; truncate, trigonous stigma; apiculate capsule valves;
and seeds disposed vertically in one or two rows per locule (Yadav et al. 2019). To date, Zingela remains a monotypic
genus differing in the very loose, cucullate, bulb scales; prominently keeled leaves; the presence of distinct bracteoles;
the diurnal flowers; the spreading filiform filaments; the coiled dehisced anthers; the erect style; and the truncate
capsule valves (Crouch et al. 2018, 2020).
The circumscription of Vera-duthiea accepted in the present work differs from Martínez-Azorín et al. (2023a)
based on our recent phylogenetic studies on Drimia macrocarpa, as discussed above in the genus Thuranthos. We
were unable to study fresh material of the type species of Vera-duthiea, though the published phylogenetic evidence
(Martínez-Azorín et al. 2023a) and the study of several herbarium collections from Senegal and Central Africa deposited
at Herb. P agree with the genus flower morphology (basally narrowed filaments) accepted in this work.
Vera-duthiea capillifolia A.Patzelt, M.Pinter & Mart.-Azorín in Phytotaxa 524: 46 (2021) (Figs 6.2, 73.1). Type:—OMAN. Jabal Qamar,
Wadi Kharfut, elev. 22 m, 13 February 2012, A. Patzelt, D. Lupton, L. Al Harthy, I. Al Rashdi & A. Al Hinai AP 4963 (OBG!
holo.).
Vera-duthiea reflexa (K.Krause & Dinter) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 296 (2019) ≡ Albuca
reflexa K.Krause & Dinter in Bot. Jahrb. Syst. 51(3–4): 445 (1914), basionym. Type:—NAMIBIA. Tsumeb (1917): Nord-Hereroland,
bei Tsumeb, (–BA), Dinter 2694 (SAM0073723! [2 cross referenced sheets, Art. 8.2 Ex. 4 of the ICN] lecto. designated by Manning
& Goldblatt 2018, or perhaps holo.).
Vera-duthiea salmonea (Berhaut) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 296 (2019) ≡ Urginea salmonea
Berhaut, Fl. Senegal ed. 2, 428 (1967), basionym. Type:—SENEGAL. Mbao, 15 June 1954, Berhaut 1682 (P00349754! lecto.
designated by Martínez-Azorín et al. 2019a: 296; P00349756! isolecto.; P00349752!, P00349753!, P00349755! syn.).
Vera-duthiea sebirii (Berhaut) Mart.-Azorín, M.B.Crespo, M.Pinter & Wetschnig in Phytotaxa 397(4): 296 (2019) ≡ Urginea sebirii
Berhaut in Bull. Soc. Bot. France Mém. 1953−1954: 7 (1954), basionym (Fig. 6.4). Type:—SENEGAL. In “savane boisée” Ngasobil,
June–July 1951, Berhaut 461 (P00349746! lecto., perhaps holo., designated by Martínez-Azorín et al. 2019a: 296).
Vera-duthiea senegalensis (Kunth) Speta in Phyton (Horn, Austria) 56(2): 155 (2016) ≡ Urginea senegalensis Kunth, Enum. Pl. 4: 334
(1843), basionym ≡ Duthiea senegalensis (Kunth) Speta in Stapfia 75: 170 (2001) ≡ Drimia senegalensis (Kunth) J.C.Manning &
Goldblatt in Edinburgh J. Bot. 60(3): 557 (2004). Type:—SENEGAL. Senegambia, prope Richard-Tol [sic, Richard Toll], Lelievre
[not extant at B, probably destroyed during World War II, R. Vogt pers. comm.].

Vera-duthiea zebrina Mart.-Azorín, N.R.Crouch & M.B.Crespo in Phytotaxa 375(4): 285 (2018) ≡ Drimia zebrina (Mart.-Azorín,
N.R.Crouch & M.B.Crespo) J.C.Manning & Goldblatt in Bothalia 49(1): 5 (2019) (Figs 72, 73.2). Type:—SOUTH AFRICA.
KwaZulu-Natal. Bela Vista (2632): Ndumo [Ndumu] Hill, Ndumo [Ndumu] Game Reserve, (–CD), 4 November 1968, E.S. Pooley
127 (NU43530! holo.; NH66607! iso.).
= Urginea zebrina Oberm., in sched.
30. Zingela N.R.Crouch, Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso in Phytotaxa 371(1): 36 (2018) (Figs 74–
75). Typus generis:—Z. pooleyorum N.R.Crouch, Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso (holotype).
Description:—Bulbous geophyte. Bulb hypogeal, 5‒8 cm in diam., composed of 10‒25 thickened, fleshy, white,
elongated loose scales, with copious threads when torn, ca. 6 × 1 cm. Roots thickened and branched. Leaves 1‒4 per
bulb, mostly hysteranthous, glaucous green, long narrowly oblong with pointed apex, 10‒35 cm long, 5‒10 mm wide,
with 2 grooves adaxially, strongly keeled abaxially in a V-shape, commonly twisting, convolute at base with distinct
purple maculations abaxially. Inflorescence a long and lax raceme, 15‒30 cm long, with 25‒55 flowers; peduncle
30‒60 cm long, greenish-brown, mottled covered white bloom; pedicels 10‒12 mm long, patent and arching down,
purple-brown, with whitish bloom, elongating up to 2 cm long and arching up in fruit. Bracts lanceolate, 2‒4 mm long,
acute, lowermost with long basal spur up to 6‒8 mm long that is flattened and appressed to stem; bracteoles present
and distinct, white. Flowers stellate, subpatent to nodding, opening in afternoon and closing at about sunset. Tepals 6,
biseriate, lanceolate, 9‒11 mm long, almost free from base, reflexed, white with broad purplish-brown median stripe.
Stamens 6, spreading; filaments free, filiform, ca. 6 mm long, slightly flattened and triangular at base; anthers narrowly
oblong, pale yellow, ca. 4 mm long before dehiscence, distinctly circinnate after pollen release, dehiscing longitudinally
along whole length. Ovary ovoid to subconical, 3.8‒4.0 mm long, yellowish, trigonous. Style filiform, white, 5.0‒5.7
mm long, erect to slightly sinuous, curved outwards in distal part. Stigma small and indistinctly trigonous. Capsule
shortly oblong, 14‒17 mm long, pale brown, valves completely dehiscing from base, tepals cohering and inrolled
above ovary after anthesis, circumcissile from base and persisting as a cap at the top of the developing capsules. Seeds
subelliptic, 10‒11 mm long, flattened, with prominent central embryo and broad wings, testa black, with strongly
sinuous and slightly raised anticlinal cell walls.
FIGURE 74. Species of Zingela N.R.Crouch et al. 1. Zingela pooleyorum N.R.Crouch et al. illustrated by Elsa Pooley based on Pooley
661 (NU); 2. Zingela sp. by Olive Anderson in Kativu & Drummond (1994: 114) based on Grosvenor 835 (SRGH).
FIGURE 75. Zingela pooleyorum N.R.Crouch et al.
Number of species and distribution:—Zingela includes a single species known so far only from two localities
within the KwaZulu-Natal Province of South Africa, and disjunctly found to occur in Harare (Zimbabwe) (Kativu &
Drumond 1994, Crouch et al. 2018, 2020) and the farm Vergenoeg, ca. 200 km northeast of Windhoek in Namibia
(A. Eichhoff s.n. ABH!) (Fig. 71). The genus is restricted to the Uzambara-Zululand Region and the Southern Section
of the Zambezian Subregion (sensu Takhtajan 1986 and Martínez-Azorín et al. 2023a). For a complete description of
Zingela see Crouch et al. (2018).
History, diagnostic characters, and taxonomic relationships:—The only known member of this monotypic
genus, Zingela pooleyorum, was described on the basis of plants with loose, bulbs scales; hysteranthous, keeled leaves,
multiflowered racemes, present bracteoles, diurnal flowers and spreading stamens with circinnate anthers, which were
previously collected in northern KwaZulu-Natal, South Africa, and usually identified as Drimia indica (Indurgia indica
in the sense of the present study). The phylogenetic results of Martínez-Azorín et al. (2023a) place four samples of
Zingela in a strongly supported clade, which is included in a polytomy with Thuranthos and Ledurgia. As reported by
Crouch et al. (2018, 2020) and Martínez-Azorín et al. (2018b), Zingela differs from Thuranthos in the diurnal flowers
and filiform spreading filaments with coiled anthers, among other characters, whereas Ledurgia presents compact
bulb scales, very short peduncle, few flowered racemes with campanulate flowers, and shortly connate tepals (Speta
2001). Therefore we accept Zingela along with Thuranthos and Ledurgia based on morphological and phylogenetic
evidence.
Zingela pooleyorum N.R.Crouch, Mart.-Azorín, M.B.Crespo, M.Pinter & M.Á.Alonso in Phytotaxa 371(1): 36 (2018) (Figs 6.6, 74.1,
75). Type:—SOUTH AFRICA. KwaZulu-Natal. Bela Vista (2632): On Pongola floodplain edge, Ndumo [Ndumu] Hill, Ndumo
[Ndumu] Game Reserve, (–CD), 15 October 1969, E.S. Pooley 661 (NU25353! holo.).
= Urginea indica (Roxb.) Kunth var. multiflora Oberm., nom. nud. (Art. 38 Ex. 1 of the ICN).
= Urginea zebrina Oberm. var. multiflora Oberm., nom. nud. in sched.

31. Zulusia Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso gen. nov. (Figs 76–77). Typus generis:—
Z. delagoensis (Baker) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso ≡ Urginea delagoensis Baker
(holotype).
Diagnosis:—Genus notabilis a Sekanama accedens, sed singulari characterum combinatione ab ea diversus et facile distinguendus, nempe
foliis synanthis; racemo quam inflorescentiae pedunculo longiore; floribus subbrunneis, carneis vel virescentibus; tepalis emarcidis
ad apicem capsulae persistentibus; capsulis anguste ellipsoideis vel fusiformibus; et seminibus anguste oblongo-fusiformibus.
Description:—Bulbous geophyte. Bulb epigeal or hypogeal, ovoid to subglobose, sometimes proliferous and clump
forming, with loosely packed, thick, fleshy scales, sometimes very loose, cucullate and pedicellate. Leaves 1‒8 per
bulb, narrowly lanceolate to filiform, canaliculate, green, glabrous to denticulate, 7‒45 cm long. Inflorescence long
racemose, 10‒65 cm long, with 10‒60 flowers; peduncle 15‒60 cm long, terete, erect, green to greyish, sometimes
papillose-hairy at base; pedicels 2.5‒6.0 mm long. Bracts lanceolate, 2‒5 mm long, lowermost with spur 3.5‒4.0 mm
long. Bracteoles absent. Flowers subcampanulate, erect-patent, diurnal. Tepals 6, biseriate, 3‒6 mm long, lanceolate to
oblong, light brown, creamy white or pale mauve, with darker longitudinal band, almost free to very shortly connate,
connivent along basal half to form campanulate structure and spreading-patent in upper portion of tepals. Stamens 6,
erect or slightly spreading; filaments filiform, 2.5‒4.0 mm long, suberect, adnate to tepals along their basal portion;
anthers dorsifixed. Ovary ovate to ellipsoid, green. Style 1.5‒2.5 mm long, erect, white, with trigonous stigma. Capsules
narrowly ellipsoid to fusiform, 7‒10 mm long, valves completely dehiscing from base; tepals cohering and inrolled
above ovary after anthesis, circumcissile from base and persisting as a cap at the top of the developing capsules.
Seeds narrowly oblong-fusiform, elongated, ca. 3‒5 times longer than wide, 4‒8 mm long, embryo prominent with
very narrow, indistinct wings, black, glossy, testa longitudinally striate, with elongate, slightly sinuous anticlinal cell
walls.
Number of species and distribution:—Zulusia is restricted to eSwatini (formerly Swaziland) and the KwaZulu-
Natal and Mpumalanga Provinces in South Africa. The genus includes three species restricted to the Uzambara-Zululand
Region (sensu Takhtajan 1986) (Fig. 71). For further information on Zulusia species see Dyer (1942a, 1942b), Jessop
(1977), Crouch & Martínez-Azorín (2015), and Manning & Goldblatt (2018).
Karyology:—2n=32 (De Wet 1957, as Urginea lydenburgensis R.A.Dyer), indicating clear chromosomal
differences between Sekanama s.str. and Zulusia. Caryological studies of other members of the genus would prove
insightful.
History, diagnostic characters, and taxonomic relationships:—Speta (2001) described Sekanama to include
Urginea sanguinea, U. burkei, and U. delagoensis, the latter species is sometimes including U. lydenburgensis in
synonymy (Jessop 1977, Manning & Goldblatt 2018), although field observations, ethnomedicinal trade research in
Nelspruit (Mpumalanga), and phytochemical profiling support their distinction (Koorbanally et al. 2005, Mullholland
et al. 2013, Crouch & Martínez-Azorín 2015). Drimia edwardsii was described from KwaZulu-Natal (Crouch &
Martínez-Azorín 2015), sharing flower and fruit morphologies with S. delagoensis and U. lydenburgensis. Our studies
have shown important differences in morphology between S. sanguinea (including U. burkei) and the remaining taxa
in this group. Sekanama sanguinea is distributed in the inland areas of northern South Africa, Botswana, and Namibia
and shows hysteranthous leaves; racemose inflorescences with peduncle much shorter than the raceme; white flowers;
withered tepals persisting at base of capsule; ovoid-ellipsoid capsule; and flat and wide, elliptic seeds. Urginea
delagoensis, U. lydenburgensis, and Drimia edwardsii are collectively distributed in eSwatini and the neighbouring
South African province of KwaZulu-Natal and produce synanthous leaves; peduncle longer than the raceme; pale
brown or carneous to greenish flowers; withered tepals persisting at the top of the capsule; narrowly ellipsoid capsules;
and narrowly lanceolate seeds.
Our phylogenetic study of Sekanama (Martínez-Azorín et al. 2023a) revealed the analysed three samples of
S. sanguinea to form a strongly supported clade. Samples of U. delagoensis, D. lydenburgensis, and D. edwardsii
formed another strongly supported clade, with both groups shown related but with very weak support. Based on their
phylogenetic divergence and the differences in morphology and biogeography of the two clades, we restrict Sekanama
to include S. sanguinea (with S. burkei in synonymy) and related taxa sharing red and fleshy bulb scales, small and
navicular bracts, presence of bracteoles, and the papery-translucent withered tepals mostly remaining at the base of
developing capsules, and describe here the new genus Zulusia to accommodate U. delagoensis, U. lydenburgensis, and
D. edwardsii.
FIGURE 76. Species of Zulusia Mart.-Azorín et al. 1. Zulusia delagoensis (Baker) Mart.-Azorín et al. from Dyer (1942a: pl. 858, as Urginea
delagoensis Baker); 2. Zulusia lydenburgensis (R.A.Dyer) Mart.-Azorín et al. from Dyer (1942b: pl. 859, as Urginea lydenburgensis
R.A.Dyer); 3. Zulusia edwardsii (N.R.Crouch & Mart.-Azorín) Mart.-Azorín et al. from Crouch & Martínez-Azorín (2015: 139, as Drimia
edwardsii N.R.Crouch & Mart.-Azorín).

FIGURE 77. Species of Zulusia Mart.-Azorín et al. displayed in horizontal rows of images. 1. Zulusia delagoensis (Baker) Mart.-Azorín
et al.; 2. Zulusia edwardsii (N.R.Crouch & Mart.-Azorín) Mart.-Azorín et al.; 3. Zulusia lydenburgensis (R.A.Dyer) Mart.-Azorín et al.
In their D. sect. Macrocentrae Manning & Goldblatt (2018) placed both Urginea macrocentra (= Boosia sensu Speta
2001) and U. sanguinea (Sekanama p.p. sensu Speta 2001), apparently overlooking the previous phylogenetic work of
Pfosser & Speta (2001, 2004), in which both these taxa resolved in different and distant clades in the phylogeny—an
earlier finding in agreement with more recent phylogenetic results (Martínez-Azorín et al. 2023a). Further, Manning &
Goldblatt (2018) placed U. delagoensis (including U. lydenburgensis) and D. edwardsii in their D. sect. Ledebouriopsis
among eleven heterogeneous species including Urgineopsis and Ornithogalum anomalum (the monotypic Geschollia
sensu Speta 2001), thus resulting in a polyphyletic aggregate according to the then available phylogenetic works
(Pfosser & Speta 2001, 2004), later confirmed by Martínez-Azorín et al. (2023a). Section Ledebouriopsis sensu
Manning & Goldblatt (2018) merges species located in five genera (Zulusia, Urginavia, Urgineopsis, Geschollia, and
Boosia) following the present study, and therefore this group is disregarded here. The name Zulusia has been selected
in recognition of the importance of both U. delagoensis and D. edwardsii in the ethnomedicinal traditions of the Zulu
nation. As recounted by Crouch & Martínez-Azorín (2015), D. edwardsii was first observed in the ethnomedicinal
plant trade, some 15 years prior to its location by taxonomists in the field, and subsequent description.
Zulusia delagoensis (Baker) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea delagoensis Baker, Fl. Cap.
(Harvey) 6(3): 467 (1897), basionym (Figs 6.7, 76.1, 77.1). Type:—MOZAMBIQUE. Between Delagoa Bay [Maputo] and the
Lebombo Mountains, 1886, H. Bolus 7627 (K000257346! holo.; BOL s.n.! iso.).
Zulusia edwardsii (N.R.Crouch & Mart.-Azorín) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Drimia edwardsii
N.R.Crouch & Mart.-Azorín in Phytotaxa 195(2): 137 (2015), basionym (Figs 6.8, 76.3, 77.2). Type:—SOUTH AFRICA. KwaZulu-
Natal. Port Shepstone (3030): middle Mkhomazi River, on north-facing shale rock ledge, 400 m, (–AA), 21 September 2011, N.
Crouch 1280 (BNRH! holo.; HSMC! iso.).
Zulusia lydenburgensis (R.A.Dyer) Mart.-Azorín, N.R.Crouch, M.B.Crespo & M.Á.Alonso comb. nov. ≡ Urginea lydenburgensis
R.A.Dyer in Fl. Pl. South Africa 22: t. 859 (1942), basionym (Figs 6.9, 76.2, 77.3). Type:—SOUTH AFRICA. Mpumalanga,
Lydenburg District, September 1937, Swart 12268 sub PRE23303 (PRE0048567-1, PRE0048567-2 holo.!).
Incertae sedis (provisionally unplaced within Urgineoideae)

Further studies are necessary to place the following taxa in the taxonomic treatment in Urgineoideae, which is the
focus of our current research.
Drimia oligosperma C.H.Wright in Bull. Misc. Inform. Kew 1912(8): 363 (1912).
Urginea aurantiaca H.Lindb. in Act. Soc. Sc. Fenn. n. s. B, 1(2): 34 (1932) ≡ Urginea noctiflora Batt. & Trab. var. aurantiaca Maire in
Bull. Soc. Bot. Afrique N. 24: 229 (1933) ≡ Urginea noctiflora subsp. aurantiaca (H.Lindb.) Maire, Fl. Afric. N. 5: 162 (1958) ≡
Thuranthos aurantiacus (H.Lindb.) Speta in Phyton (Horn, Austria) 38(1): 81 (1998) ≡ Drimia aurantiaca (H.Lindb.) J.C.Manning
& Goldblatt in Edinburgh J. Bot. 60(3): 556 (2004). Type:—MOROCCO. Atlas major, in convalle fl. Aït Messane, in decliv. saxosis
inter Tinitine et Anferkou, elev. 1500 m, 12 June 1926, H. Lindberg 3900 (H1237873! holo.). Note:—Lectotypification by Väre
(2012) is disregarded since the protologue clearly includes a photograph of those two plants on which the description was made, and
they are to be regarded as the type (holotype) material.
Comments:—Urginea aurantiaca Lindberg (1932: 34) was described from plants collected in bud in the High Atlas of Morocco, between
Tinitine and Anferkou. This collection flowered in cultivation. The new species was related to Urginea fugax, but differed by the
longer peduncle, larger and cinnamom coloured bulbs, smaller orange flowers (tepals 8−9 mm long), and flowering time around
June, instead of August–October. No mention of the diurnal or nocturnal flowering time of the species was included in the protologue.
The type collection at Helsinki herbarium (H1237873) shows two bulbs with inflorescences lacking leaves. Flowers appear to be
subcampanulate, with tepals suberect, very small, only ca. 5 mm long. The illustration in the protologue shows the same two plants
with small flowers, fitting the previous description.
Maire (1933, 1958) combined Lindberg's species respectively as variety and subspecies of Urginea noctiflora, based on the apparently
shared nodding nocturnal flowers with reflexed tepals with typical U. noctiflora, but differing in widely expanded filaments at
the base and the straight leaves. Stearn (1978) suggested the convenience of assuming the two taxa accepted by Maire as a good
species. Several herbarium collections from Morocco fit Maire’s concept of U. aurantiaca, based on the nocturnal, nodding orange
flowers with basally expanded filaments and strongly reflexed tepals. Our recent field work in Morocco facilitated the study of fresh
material of what Maire (1958) had considered Urginea noctiflora subsp. aurantiaca. Our inedit phylogenetic study of two Moroccan
populations from Chichaoua and Douar Shémch reveals their close relation to Urginea fugax and Spirophyllos noctiflorus clade,
where the three species share biogeography. However, further studies are needed to ascertain whether our studied samples represent
Lindberg’s Urginea aurantiaca, based on their remarkable flower divergence with regard to the type collection. We have noticed that
buds never open until they reach about 9−10 mm long, and the type collection includes suberect open flowers with very short tepals
(c. 5 mm long) disposed on very short pedicels. Further, Lindberg did not mention in the protologue the typical nocturnal flowering
of his species, a distinct character unmistakable in cultivation.
Battaglia (1958) studied caryology of Urginea aurantiaca material collected by R. Nègre in 1952 near Chichaoua, West of Marrakech in
Morocco. The plants were cultivated at Fordham University in New York and the described morphology and a photograph of the
flowering plant on August 1958 fit Lindberg’s concept and the type collection, showing suberect smaller flowers on short pedicels.
Moreover, the detailed caryological study of Battaglia (1958) concluded that the studied samples from Chichaoua were 2n=20+1–2B
and closely related to the caryotype of Urginea fugax.
Urginea brevipes Baker in J. Bot. 12: 364 (1874).
Urginea flaviflora Rojas Acosta in Cat. Hist. Nat. Corrientes 86: 184 (1897).
Urginea insignis Engl. & K.Krause in Bot. Jahrb. Syst. 45(1): 147 (1910).
Urginea ledermannii Engl. & K.Krause in Bot. Jahrb. Syst. 45(1): 145 (1910).
Urginea petitiana Solms in Beitr. Fl. Aethiop. [Schweinfurth]: 294 (1867), nom. nud. =? Scilla petitiana A.Rich. in Tent. Fl. Abyss. 2:
328 (1850).

Urginea quartiniana Solms in Beitr. Fl. Aethiop. [Schweinfurth]: 294 (1867), nom. nud. =? Scilla quartiniana A.Rich. in Tent. Fl. Abyss.
2: 329 (1850).
Urginea revoluta A.V.Duthie in Ann. Univ. Stellenbosch 1928, 6: Sect. A No. 2, 9 (1928) ≡ Thuranthos revolutum (A.V.Duthie) Speta
in Phyton (Horn, Austria) 38(1): 84 (1998) ≡ Drimia revoluta (A.V.Duthie) J.C.Manning & Goldblatt in Strelitzia 9: 712 (2000),
nom. illeg., non Drimia revoluta Kunth, Enum. Pl. 4: 341 (1843), nec Drimia revoluta (L.f.) Sweet, Hort. Suburb. Lond.: 72 (1818)
cf. D.J.Mabberley in Taxon, 29(5-6): 604 (1980) (Figs 6.11, 78). Type:—SOUTH AFRICA. Stellenbosch Flats, March 1927,
Duthie 1874 (NBG0197704-0! lecto. designated as “holo.” by Jessop in J. S. African Bot. 43: 303. 1977; PRE0049744-0! isolecto.;
K000257368!, K000257369!, syn.).
Comments:—In our phylogenetic analyses, this species nests in a clade together with two samples of Fusifilum magicum and an unidentified
Urgineoid species (Martínez-Azorín et al. 2023a). The morphologically closest species of Urginea revoluta appears to be Triandra
pellabergensis, based on their very similar nocturnal flowers. However, they differ in the plant size, number of stamens, seed
morphology (cf. Martínez-Azorín et al. 2021) and the pedicels abscising near the middle if the flowers are not pollinated (Manning
& Goldblatt 2018). This latter feature is unique in the subfamily and may point to the taxonomic independence of Urginea revoluta
at genus rank, though further molecular evidence is necessary to evaluate this solution in the context of its related taxa.
FIGURE 78. Urginea revoluta A.V.Duthie.
Urginea vittata Schönland in Mem. Bot. Surv. South Africa 1: 41 (1919), nom. nud.
Names excluded from Urgineoideae
Drimia acuminata G.Lodd., Bot. Cab. 11(5): t. 1041 (1826) = Ledebouria revoluta (L.f.) Jessop in J. S. African Bot. 36: 255 (1970).
Drimia acuminata G.Lodd. ex J.Forbes, Hort. Woburn.: 61 (1833) = Ledebouria revoluta (L.f.) Jessop in J. S. African Bot. 36: 255
(1970).
Drimia angustifolia Kunth, Enum. Pl. 4: 340 (1843) = Ledebouria undulata (Jacq.) Jessop in J. S. African Bot. 36: 258 (1970).
Drimia angustitepala Engl. in Bot. Jahrb. Syst. 15(4): 475 (1892) = Ledebouria kirkii (Baker) Stedje & Thulin in Nordic J. Bot. 15(6):
595 (1996).
Drimia apertiflora Baker, Refug. Bot. [Saunders] 1: t. 19 (1868) ≡ Ledebouria apertiflora (Baker) Jessop in J. S. African Bot. 36: 254
(1970).
Drimia bifolia Schweinf., Beitr. Fl. Aethiop.: 294 (1867) = Ledebouria revoluta (L.f.) Jessop in J. S. African Bot. 36: 255 (1970).
Drimia brevifolia Baker, Fl. Trop. Afr. [Oliver et al.] 7(3): 527 (1898) = Ledebouria revoluta (L.f.) Jessop in J. S. African Bot. 36: 255
(1970).
Drimia chlorantha Baker, Fl. Cap. (Harvey) 6(3): 443 (1897) = Albuca abyssinica Jacq., Icon. Pl. Rar. 1: t. 64 (1783).
Drimia coleae Baker in Bot. Mag. 123: t. 7565 (1897) = Ledebouria somaliensis (Baker) Stedje & Thulin in Nordic J. Bot. 15(6): 595
(1996).
Drimia confertiflora Dammer in Bot. Jahrb. Syst. 38(1): 62 (1905) ≡ Ledebouria confertiflora (Dammer) U.Müll.-Doblies & D.Müll.-
Doblies in Feddes Repert. 108(1-2): 56 (1997).
Drimia cooperi Baker, Refug. Bot. [Saunders] 1: t.18 (1868) = Ledebouria concolor (Baker) Jessop in J. S. African Bot. 36: 254 (1970).
Drimia cowanii Ridl. in J. Linn. Soc., Bot. 20: 334 (1883) ≡ Dipcadi cowanii (Ridl.) H.Perrier in Notul. Syst. (Paris) 5: 63 (1935).
Drimia dregeana Kunth, Enum. Pl. 4: 340 (1843) = Ledebouria nitida (Eckl.) J.C.Manning & Goldblatt in Strelitzia 40: 159 (2018).
Drimia ensifolia Eckl. in S. African Quart. J. 1(4): 364 (1830) = Ledebouria ensifolia (Eckl.) S.Venter & T.J.Edwards in Bothalia 33(1):
49 (2003).
Drimia fischeri Baker, Fl. Trop. Afr. [Oliver et al.] 7(3): 526 (1898) = Drimiopsis fischeri (Engl.) Stedje in Nordic J. Bot. 15(6): 593
(1996).
Drimia gawleri Schrad. in Blumenbachia: 30 (1827) = Ledebouria ovalifolia (Schrad.) Jessop in J. S. African Bot. 36: 246 (1970).
Drimia glaucescens (Engl. & K.Krause) H.Scholz in Bull. Mus. Natl. Hist. Nat., B, Adansonia sér. 4, 11(4): 444 (1990) = cf. Stellarioides
sp.
Drimia hildebrandtii Baker in Bot. Jahrb. Syst. 15(4): 474 (1892) ≡ Ledebouria hildebrandtii (Baker) U.Müll.-Doblies & D.Müll.-
Doblies in Feddes Repert. 108(1-2): 58 (1997).
Drimia lanceifolia Ker Gawl. in Bot. Mag. 34: sub t. 1380 (1811) = Ledebouria revoluta (L.f.) Jessop in J. S. African Bot. 36: 255
(1970).
Drimia lanceifolia G.Lodd. in Bot. Cab. 3(7): t. 278 (1819), nom. illeg. = Ledebouria revoluta (L.f.) Jessop in J. S. African Bot. 36: 255
(1970).
Drimia lanceifolia G.Lodd. var. longipedunculata Schrad. in Blumenbachia: 30 (1827) = Ledebouria revoluta (L.f.) Jessop in J. S. African
Bot. 36: 255 (1970).
Drimia lanceolata Schrad. in Blumenbachia: 28 (1827) = Ledebouria ovalifolia (Schrad.) Jessop in J. S. African Bot. 36: 246 (1970).
Drimia lilacina (Fenzl) Schweinf., Beitr. Fl. Aethiop.: 294 (1867) ≡ Eratobotrys lilacinus Fenzl, Gen. Pl. [Endlicher] Suppl. 2: 13 (1842)
≡ Ledebouria lilacina (Fenzl) Speta in Phyton (Horn, Austria) 38(1) 106 (1998).
Drimia longipedunculata Sweet, Hort. Brit. [Sweet], ed. 2. 529 (1830); cf. D.J. Mabberley in Taxon 29(5-6): 604 (1980) = Ledebouria
revoluta (L.f.) Jessop in J. S. African Bot. 36: 255 (1970).
Drimia longipedunculata G.Don, Hort. Brit. [Loudon], ed. 2: 587 (1832) = Ledebouria revoluta (L.f.) Jessop in J. S. African Bot. 36: 255
(1970).
Drimia ludwigii Miq. in Bull. Sci. Phys. Nat. Néerl. 1839: 39 (1839) = Ledebouria ensifolia (Eckl.) S.Venter & T.J.Edwards in Bothalia
33(1): 49 (2003).
Drimia nitida Eckl. in S. African Quart. J. 1(4): 364 (1830) ≡ Ledebouria nitida (Eckl.) J.C.Manning & Goldblatt in Strelitzia 40: 159
(2018).
Drimia obstii W.Heering in Mitt. Geogr. Ges. Hamburg 29: 209 (1915) basionym. ≡ Ledebouria obstii (W.Heering) Mart.-Azorín,
M.B.Crespo & M.Á.Alonso comb. nov. Type:—TANZANIA. Deutsch-Ost-Africa, Ufiome, Gipfel, E. Obst s.n. (HBG528404! holo.)
[species akin to Ledebouria kirkii (Baker) Stedje & Thulin in Nordic J. Bot. 15(6): 595 (1996)].
Drimia ovalifolia Schrad. in Blumenbachia: 30 (1827) ≡ Ledebouria ovalifolia (Schrad.) Jessop in J. S. African Bot. 36: 246 (1970).
Drimia pendula Burch. ex Baker in J. Linn. Soc., Bot. 13: 254 (1872) = Ledebouria floribunda (Baker) Jessop in J. S. African Bot. 36:
251 (1970).
Drimia petiolata K.Koch & C.D.Bouché in Index Seminum [Berlin]: 3 (1861) = Drimiopsis maculata Lindl. & Paxton, Paxton’s Fl. Gard.
2: 73 (1851–52).
Drimia polyphylla (Hook.f.) Ansari & Sundararagh. in J. Bombay Nat. Hist. Soc. 77(1): 174 (1980): (1980) = cf. Dipcadi sp.
Drimia revoluta (L.f.) Sweet, Hort. Suburb. Lond.: 72 (1818) ≡ Ledebouria revoluta (L.f.) Jessop in J. S. African Bot. 36: 255 (1970).
Drimia undulata Jacq., Icon. Pl. Rar. 2(15): t. 376 (1794) ≡ Ledebouria undulata (Jacq.) Jessop in J. S. African Bot. 36: 258 (1970).
Drimia viridiflora Eckl. ex Kuntze in Linnaea 20: 10 (1847) [Drimia viridiflora Eckl., Topogr. Verz. Pflanzensamml. Ecklon: 2 (1827),
nom. nud.] = cf. Ledebouria sp.
Schizobasis flagelliformis (Baker) Baker in J. Linn. Soc., Bot. 15: 261 (1877) ≡ Eriospermum flagelliforme (Baker) J.C.Manning in
Bothalia 30(2): 157 (2000) = Eriospermum abyssinicum Baker in J. Linn. Soc., Bot. 15: 263 (1876) (see Manning 2000). Type:—
SOUTH AFRICA. Gauteng. Pretoria (2528): Apies River (Aapages River), Burke s.n. (K000257124! lecto. designated here: only
the five inflorescences with long pedicels, excluding the inforescence on the upper left corner). Note:—The type voucher at Herb.
K is a mixed collection of an inflorescence of a Boosia sp. (on the upper left corner) with relatively short pedicels and long spurred
bracts, plus five inflorescences of Eriospermum flagelliforme with much longer pedicels and unspurred bracts. Therefore, a lectotype
designation is needed and is effected here.
Urginea acinacifolia Schinz in Verh. Bot. Vereins Prov. Brandenburg 31: 220 (1890) = Albuca angolensis Welw., Apont.: 591 (1859).
Urginea alooides Bolus in J. Linn. Soc., Bot. 18: 395 (1881) ≡ Aloe alooides (Bolus) Druten in Bothalia 6: 544 (1956).
Urginea angolensis Baker in J. Bot. 12: 364 (1874) ≡ Battandiera angolensis (Baker) Mart.-Azorín & M.B.Crespo in Pl. Biosystems
148(1-2): 214 (2013).
Urginea autumnalis (L.) El-Gadi, Fl. Libya 57: 38 (1978) ≡ Prospero autumnale (L.) Speta in Veröff. Int. Clusius-Forschungsges. Güssing
5: 4 (1982).
Urginea bakeri Chiov. in Ann. Bot. (Rome) 9: 144 (1911) = Stellarioides tenuifolia (F.Delaroche) Speta in Stapfia 75: 174 (2001).
Urginea beccarii Baker in J. Linn. Soc., Bot. 13: 223 (1872) = cf. Albuca abyssinica Jacq. in Icon. Pl. Rar. 1: t. 64 (1783).
Urginea bequaertii De Wild. in Pl. Bequaert. 1: 31 (1921) = cf. Albuca sp.
Urginea berylloides Baker in Bull. Herb. Boissier ser. 2, 1: 785 (1901) ≡ Ornithogalum umgeniense Baker in Bull. Herb. Boissier ser. 2,
1: 856 (1901) = Stellarioides tenuifolia, as previously suggested by Obermeyer (1978) as Ornithogalum.
Urginea bragae Engl. in Pflanzenw. Ost-Afrikas 2: 142 (1895) ≡ Schizocarphus bragae (Engl.) U. & D. Müller-Doblies, ined.
Urginea cepifolia (Baker) Welw. ex Rendle in Cat. Afr. Pl. (Hiern) 2: 60 (1899) ≡ Ornithogalum cepifolium Baker in Trans. Linn. Soc.

London, Bot. 1(5): 248 (1878) = Stellarioides sp.
Urginea chlorantha Welw. ex Baker in Trans. Linn. Soc. London, Bot. 1(5): 248 (1878) = Battandiera angolensis (Welw. ex Baker) Mart.-
Azorín & M.B.Crespo in Pl. Biosystems 148(1-2): 214 (2013).
Urginea comosa Welw. ex Baker in Trans. Linn. Soc. London, Bot. 1(5): 247 (1878) = Battandiera angolensis (Baker) Mart.-Azorín &
M.B.Crespo in Pl. Biosystems 148(1-2): 214 (2013).
Urginea coromandeliana Wight in Icon. Pl. Ind. Orient. 6: 27, t. 2064 (1853) = cf. Dipcadi sp.
Urginea corradii Chiov. in Webbia 8: 26 (1951) = Ledebouria kirkii (Baker) Stedje & Thulin in Nordic J. Bot. 15(6): 595 (1996).
Urginea dimorphantha Baker in Bull. Herb. Boissier ser. 2, 3: 663 (1903) = Battandiera angolensis (Baker) Mart.-Azorín & M.B.Crespo
in Pl. Biosystems 148(1-2): 214 (2013).
Urginea ensifolia (Thonn.) Hepper in Kew Bull. 21(3): 497 (1968) = cf. Stellarioides sp.
Urginea flavovirens (Baker) Weim. in Bot. Not.: 442 (1937) ≡ Stellarioides flavovirens (Baker) Speta in Stapfia 75: 173 (2001).
Urginea garuensis Engl. & K.Krause in Bot. Jahrb. Syst. 45(1): 147 (1910) = Albuca sudanica A.Chev. in Bull. Soc. Bot. France 55(Mém.
8b): 93 (1908).
Urginea glaucescens Engl. & K.Krause in Bot. Jahrb. Syst. 45(1): 146 (1910) = cf. Stellarioides sp.
Urginea grandiflora Baker in J. Linn. Soc., Bot. 13: 223 (1872), basionym ≡ Albuca grandiflora (Baker) Mart.-Azorín, M.B.Crespo &
M.Á.Alonso comb. nov. Type:—SUDAN. Hor Tamanib, Red Sea, August 1869, J.K. Lord s.n. (K001422041!, holo.).
= Albuca blepharophylla Cufod. in Missione Biol. Borana 4: 320 (1939).
Urginea japonica (Thunb.) Pepin in Rev. Hortic. ser. 2. 2: 374 (1843–44) ≡ Ornithogalum japonicum Thunb. in Nova Acta Regiae Soc.
Sci. Upsal. 3: 209 (1780); Thunb. in Fl. Jap.: 137 (1784) ≡ Barnardia japonica (Thunb.) Schult & Schult.f. in Syst. Veg., ed. 15 bis,
7(1): 555 (1829).
Urginea langii Bremek. in Ann. Transvaal Mus. 25: 237 (1933) = Battandiera seineri (Engl. & K.Krause) Mart.-Azorín, M.B.Crespo &
Juan in Ann. Bot. (Oxford) n. s., 107(1): 26 (2011).
Urginea lorata Baker in Bull. Herb. Boissier ser. 2, 3: 664 (1903) = Dipcadi glaucum Baker in J. Linn. Soc., Bot. 11: 401 (1870).
Urginea mankonensis Hutch. in Fl. W. Trop. Afr. [Hutchinson & Dalziel] 2: 347 [in clavi], 348 (1936) = Albuca mankonensis A.Chev. in
Fl. W. Trop. Afr. [Hutchinson & Dalziel] 2: 348 (1936), nom. inval. in syn. = Albuca sudanica A.Chev. in Bull. Soc. Bot. France
55(Mém. 8b): 93 (1908).
Urginea mouretii Batt. & Trab in Bull. Soc. Bot. France 68: 438 (1922) = Stellarioides longibracteata (Jacq.) Speta in Stapfia 75: 173
(2001).
Urginea narcissifolia (A.Chev.) Hutch., Fl. W. Trop. Afr. [Hutchinson & Dalziel] 2: 347, in clavi, 348 (1936) = Stellarioides sp.
Urginea nematodes (Schult.f.) Baker in J. Linn. Soc., Bot. 13: 218 (1872) ≡ Anthericum nematodes Schult.f. in Syst. Veg., ed. 15 bis
[Roemer & Schultes] 7(1): 472 (1829), nom. nov. et basionym ≡ Nicipe nematodes (Schult.f.) Mart.-Azorín, M.B.Crespo &
M.Á.Alonso comb. nov. ≡ Anthericum filifolium Thunb. in Prodr. Pl. Cap. 1: 62 (1794), syn. subst. ≡ Ornithogalum thunbergianulum
U.Müll.-Doblies & D.Müll.-Doblies in Feddes Repert. 107(5-6): 440 (1996), nom. nov. [non Ornithogalum filifolium Kunth, Enum.
Pl. 4: 369 (1843)]. Type:—SOUTH AFRICA. Caput Bona Spei, Thunberg s.n. (UPS-THUNB 8372! lecto. designated as “Type” by
Jessop in J. S. African Bot. 43: 315. 1977).
Urginea nigritana Baker in J. Linn. Soc., Bot. 13: 224 (1872) ≡ Albuca nigritana (Baker) Troupin in Bull. Jard. Bot. État Bruxelles 25:
231 (1955).
Urginea paludosa Engl. & K.Krause in Bot. Jahrb. Syst. 45(1): 146 (1910) = Stellarioides sp.
Urginea pilosula Engl. in Bot. Jahrb. Syst. 32(1): 93 (1902) = Trachyandra saltii (Baker) Oberm. in Bothalia 7: 722 (1962).
Urginea polyphylla Hook.f. in Fl. Brit. India [J.D. Hooker] 6(18): 348 (1892) = cf. Dipcadi sp.
Urginea pulchra (Schinz) Sölch in Mitt. Bot. Staatssamml. München 4: 73 (1961) = Battandiera angolensis (Baker) Mart.-Azorín &
M.B.Crespo in Pl. Biosystems 148(1-2): 214 (2013).
Urginea secunda (Jacq.) Baker in J. Linn. Soc., Bot. 13: 222 (1872) ≡ Coilonox secundum (Jacq.) Speta in Stapfia 75: 175 (2001).
Urginea somalensis Chiov. in Fl. Somala 2: 427 (1932) = Battandiera donaldsonii (Rendle) Mart.-Azorín, M.B.Crespo & Juan in Ann.
Bot. (Oxford) n.s., 107(1): 26 (2011).
Urginea tayloriana Rendle in J. Linn. Soc., Bot. 30: 424 (1895) = Stellarioides sp.
Urginea volubilis H.Perrier in Notul. Syst. (Paris) 5: 65 (1935) ≡ Igidia volubilis (H.Perrier) Speta in Phyton (Horn, Austria) 38(1): 70
(1998).
Acknowledgements
This work was partly supported by H2020 Research and Innovation Staff Exchange Programme of the European
Commission, Project 645636: ‘Insect-plant relationships: insights into biodiversity and new applications’ (FlyHigh)
and the complementary supporting funds UAUSTI17-03, UAUSTI19-08, ACIE17-01, ACIE18-03, and ACIE21-
01 (University of Alicante, Spain). We thank the curators of the herbaria cited above who provided access to the
specimens examined and the project E-RECOLNAT (ANR-11-INBS-0004) for allowing us to reproduce a drawing
kept at Herb. MPU. Royal Botanic Gardens Kew provided seeds of Drimia macrocarpa and permission to be used in
the phylogenetic analyses. Our thanks also go to all managers of the visited nature reserves for permission to work in
areas in their care. The Department of Environment and Nature Conservation of Northern Cape Province (collecting
and export permits FLORA046/2010, FLORA047/2010, FLORA069/2011, FLORA070/2011, FLORA61/2/2015,
FLORA61/2/2015, FLORA0057/2017, FLORA0058/2017, FLORA0059/2022, FLORA0060/2022), CapeNature of
Western Cape Province of South Africa (collecting and export permits AAA005-00080-0028, AAA008-00031-0028,
0027-AAA008-00699, 0028-AAA008-00203, CN35‐28‐21389, CN17‐28‐21605), the Eastern Cape Department of
Economic Development, Environmental Affairs and Tourism (collecting permits to A.P. Dold), and Ezemvelo-KZN
Wildlife (collecting permit OP 3969/2015) provided permission to collect specimens. The Ministry of Environment
and Tourism of Namibia provided research/collecting permit 2192/2016; C. Mannheimer, S. Rugheimer, and F. Chase
kindly helped with fieldwork in that country. LM appreciates the logistic support of the Iluka Chair in Vegetation
Science & Biogeography at Murdoch University. We thank U. and D. Müller-Doblies, B. Stedje, A. Garcin, W. Knirsch,
E. van Jaarsveld, D. Styles, J. Slade, D. Human, L. Peruzzi, T. Harvey, A. Yoffe, P. Harvey, H. Kolberg, D. Oosthuizen,
A. Eichhoff, I. Brase, E. Pooley, J. Burrows, S. Burrows, M. Lotter, C. Chapano, T. Edwards, B. Zaalberg, S. Dutta,
J.L. Villar, and A. Patzelt, among numerous collaborators for sharing material or information on Urgineoideae.
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Phytotaxa
Volume 610 (1): 1–143 (31 August 2023)
MARIO MARTÍNEZ-AZORÍN, MANUEL B. CRESPO, MARÍA ÁNGELES

ALONSO-VARGAS, MICHAEL PINTER, NEIL R. CROUCH, ANTHONY P.
DOLD, LADISLAV MUCINA, MARTIN PFOSSER & WOLFGANG WETSCHNIG
Taxonomy and systematics and specifically the generic circumscription of the Hyacinthaceae subfamily
Urgineoideae have been a matter of controversy in recent decades. Widely contrasting taxonomic treatments
have been proposed, ranging from systems comprising only two genera (including a morphologically very
variable Drimia covering nearly the whole subfamily) to about 20 genera based mainly on preliminary
phylogenetic studies supporting recognition of traditionally accepted and morphologically distinct
genera such as Litanthus, Schizobasis, Tenicroa, and Thuranthos. All previous studies covered only a
limited diversity and distribution of the subfamily, precluding comprehensive insight into the evolution
of the diversity within the subfamily. Based on detailed morphological, phylogenetic, and biogeographic
studies in Urgineoideae, covering its whole distribution range and most of the currently known species,
we present a new taxomic treatment of the subfamily focusing on the circumscription of genera. Our
analyses revealed 31 phylogenetic lineages corroborating unique syndromes of morphological characters
linked to well-defined biogeographic patterns. These lineages are interpreted as well-defined genera.
An identification key to the genera is provided to facilitate further taxonomic work in Urgineoideae
and morphological descriptions of the accepted genera are presented. Two new genera, Spirophyllos
gen. nov. and Zulusia gen. nov., are described to accommodate newly revealed phylogenetic lineages
from northwestern and southeastern Africa, respectively. Further, a complete list of taxa included in the
accepted genera is presented, providing details on typification, and the 50 new combinations, and one
new name required to appropriately accomodate taxa in the new taxonomic treatment.
Online at:
www.mapress.com/pt/ ISBN 978-1-77688-860-3 (paperback)

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Phytotaxa 610

ISSN 1179-3155 (print edition)

ISSN 1179-3163 (online edition)

A generic monograph of the Hyacinthaceae

MARIO MARTÍNEZ-AZORÍN, MANUEL B. CRESPO, MARÍA ÁNGELES

Magnolia Press, Auckland, New Zealand

Managing editor/founder Zhi-Qiang Zhang (Auckland)

ISSN 1179-3155 (print edition)

A generic monograph of the Hyacinthaceae subfamily Urgineoideae

MARIO MARTÍNEZ-AZORÍN1,9*, MANUEL B. CRESPO1,10, MARÍA ÁNGELES ALONSO-

Accepted by Ronell R. Klopper: 4 Jul. 2023; published: 31 Aug. 2023 

FIRST PUBLISHED IN 2023 BY

© 2023 Magnolia Press

ISSN 1179-3155 (print edition)

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Key words: Asparagaceae, distribution, ecology, Scilloideae, taxonomy, typification, Urgineae.

 • Phytotaxa 610 (1) © 2023 Magnolia Press MARTÍNEZ-AZORÍN Et al.

Materials and methods

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Results and discussion

Previous taxonomic proposals and phylogenetic relationships

Comments on morphological characters

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Identification key of genera in Urgineoideae

1. Anthers dehiscing by apical pores or slits extending to the middle .................................................................................................. 2

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A synopsis of the accepted genera

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

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Accepted species and required new combination:—

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FIGURE 29. Approximate distribution ranges of accepted genera in Urgineoideae.

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FIGURE 32. Iosanthus toxicarius (C.Archer & R.H.Archer) Mart.-Azorín et al.

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Number of species and distribution:—Mucinaea is a monotypic genus endemic to Namaqualand (northwestern

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

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FIGURE 43. Sagittanthera cyanelloides (Baker) Mart.-Azorín et al.

Accepted by Ronell R. Klopper: 4 Jul. 2023; published: 31 Aug. 2023

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