Systematics of the sub-Saharan African squills: The genera Merwilla, Pseudoprospero, Schizocarphus and Spetaea (Hyacinthaceae: Scilloideae)

Systematics of the sub-Saharan African squills: The genera Merwilla, Pseudoprospero, Schizocarphus and Spetaea (Hyacinthaceae: Scilloideae)

South African Journal of Botany 125 (2019) 411–426 Contents lists available at ScienceDirect South African Journal of Botany journal homepage: www.e...

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South African Journal of Botany 125 (2019) 411–426

Contents lists available at ScienceDirect

South African Journal of Botany journal homepage:

Systematics of the sub-Saharan African squills: The genera Merwilla, Pseudoprospero, Schizocarphus and Spetaea (Hyacinthaceae: Scilloideae) J.C. Manning a,b,⁎ a b

Compton Herbarium, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, South Africa Research Centre for Plant Growth and Development, School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, Private Bag X01, Scottsville 3209, South Africa

a r t i c l e

i n f o

Article history: Received 21 May 2019 Received in revised form 22 July 2019 Accepted 2 August 2019 Available online 20 August 2019 Edited by GV Goodman-Cron Keywords: Hyacintheae Massonieae New combination Nomenclature Scilla Southern Africa Taxonomy

a b s t r a c t The sub-Saharan African species previously included in Scilla subgenus Scilla are reviewed, with full nomenclature, descriptions, ecological notes, and colour illustrations. Four genera and five species are recognised. Pseudoprospero firmifolium (tribe Pseudoprospereae) is endemic to South Africa and is separated by leaf width and ovary colour into subsp. firmifolium from the Eastern Cape and subsp. natalensis from KwaZulu-Natal. It is diagnosed primarily by its bi-ovulate locules, and often branched raceme. Merwilla (tribe Massonieae) is recognised by its white ovary, and unique, pale beige seeds. It includes the dwarf M. dracomontana from the foothills of the KwaZulu-Natal and Eastern Cape Drakensberg, distinguished by its prostrate foliage and hispidulous scape, and the variable M. plumbea from southern and east subtropical Africa, with hispidulous or glabrous foliage but consistently glabrous scape. Dwarf forms of this species from the southeastern coast and near-interior of South Africa that include the type of M. kraussii are segregated as subsp. kraussii. The subtropical M. lazulina is treated as a synonym of subsp. plumbea. The monotypic Schizocarphus (tribe Massonieae) is diagnosed by its fibrous leaves and dark ovary, and S. nervosus is widely distributed through southern and subtropical Africa. Although it is highly variable in stature, in the shape and vestiture of the foliage, and in the length of the pedicels, it is not possible to recognise more than a single species. Spetaea lachenaliiflora (tribe Massonieae) is endemic to the southwestern mountains of the Western Cape, South Africa, and is readily distinguished from the other three genera by its tunicated bulb, ebracteolate raceme, and campanulate flowers with exserted stamens and style. © 2019 SAAB. Published by Elsevier B.V. All rights reserved.

1. Introduction At the time that Baker (1896–1897, 1898) treated the sub-Saharan African species of Hyacinthaceae [sensu APG II, 2003; alternatively Asparagaceae subfamily Scilloideae sensu APGIII, 2009], the genus Scilla L. (species of which are known colloquially as squills) was diagnosed from other related bulbous genera by its usually mauve or blue perianth of ± free (or shortly united), uni-nerved tepals, and its ellipsoid or subglobose seeds. In this circumscription, the genus was widely distributed through Eurasia and Africa. The great majority of the sub-Saharan representatives were separated as the subgenus Ledebouria (Roth) Baker, which was later re-instated as the separate genus Ledebouria Roth (Jessop, 1970) to leave only a small residue of Scilla species still recognised from sub-Saharan Africa. Some of these were subsequently segregated as the small genus Schizocarphus F. van der Merwe (1943a) on account of their uniquely fibrous outer bulb scales and leaves but this treatment was rejected by Jessop (1970), who retained them in ⁎ Corresponding author at: Compton Herbarium, South African National Biodiversity Institute, Private Bag X7, Claremont 7735, South Africa. E-mail address: [email protected]. 0254-6299/© 2019 SAAB. Published by Elsevier B.V. All rights reserved.

Scilla in his revision of the southern Africa species, and by Stedje (1996) for her treatment of the family in tropical East Africa. The first independent evidence that the sub-Saharan squills [i.e. those species that were treated in Scilla by Jessop, 1970] were misplaced in that genus came from the early phylogenetic analysis of chloroplast sequence data for 10 species of Scilloideae by Stedje (1998), which included two Eurasian species of Scilla plus S. lazulina Wild. and S. nervosa (Burch.) Jessop from sub-Saharan Africa. Her analysis retrieved the Eurasian and sub-Saharan species as two separate clades, and her finding that the two sub-Saharan species of Scilla were placed as a grade with Ledebouria provided support for Van der Merwe's (1943a) proposition that they did not belong in a single genus. These preliminary findings have since been confirmed and amplified by molecular phylogenetic analyses of a much larger sampling of species (Pfosser and Speta, 1999; Manning et al., 2004; Ali et al., 2012; Buerki et al., 2012). It is now clear that the true squills, comprising the genus Scilla and allied segregates in tribe Hyacintheae, are restricted to the Northern Hemisphere, and are not immediately related to the subSaharan members that were previously included in that genus. These findings encouraged Speta (1998) to revive the genus Schizocarphus for those species with white flowers and fibrous bulb layers and foliage,


J.C. Manning / South African Journal of Botany 125 (2019) 411–426

Table 1 Taxonomic disposition of the sub-Saharan squills by various authors. Genus/Taxonomist

Baker (1896–1897)

Jessop (1970)

Speta (1989)

Wetschnig and Pfosser (2003)

Merwilla Pseudoprospero Schizocarphus Spetaea



Merwilla Pseudoprospero Schizocarphus

Merwilla Pseudoprospero Schizocarphus Spetaea

and to describe two new genera to accommodate the remaining species with mauve or blue flowers and papery or cartilaginous bulb layers and leathery foliage, viz. the monotypic Pseudoprospero Speta for S. firmifolia Baker, and Merwilla Speta with five species. Confusion about the application of the name S. plumbea Lindl. obscured the true identity of plants collected in the mountains near Wellington in the Western Cape, until this was resolved by the description of a second monotypic genus, Spetaea Wetschnig & Pfosser with the new species S. lachenaliiflora Wetschnig & Pfosser (Wetschnig and Pfosser, 2003). The taxonomic history of the sub-Saharan squills is summarised in Table 1. Pseudoprospero and Spetaea are both endemic to South Africa, Merwilla is distributed through eastern southern Africa and extends into eastern Zimbabwe and adjacent Mozambique and Malawi, and Schizocarphus is widespread through southern and subtropical Africa. The phylogenetic relationships among the four genera of subSaharan squills are now firmly established (Manning et al., 2004; Ali et al., 2012; Buerki et al., 2012). Pseudoprospero is taxonomically isolated as the sole member of the tribe Pseudoprospereae, with an estimated divergence date of ± 55 Ma (Buerki et al., 2012)/±19 Ma (Ali et al., 2012) from a common ancestor with the Northern Hemisphere Hyacintheae (which includes Scilla and allied genera) and the largely sub-Saharan Massonieae. Within Massonieae, Merwilla comprises the earliest diverging branch, with an estimated age of ± 48 Ma (Buerki et al., 2012)/±16 Ma (Ali et al., 2012), and Schizocarphus is sister to Ledebouria as a clade of largely summer-rainfall species, with an estimated divergence date of ±25 Ma (Buerki et al., 2012)/±11.4 Ma (Ali et al., 2012). Spetaea, also with an estimated age of ± 25 Ma (Buerki et al., 2012)/±10 Ma (Ali et al., 2012), is placed as sister to Daubenya in a diverse clade of largely winter-rainfall genera that includes Lachenalia J.Jacq. ex Murray, Massonia Thunb. ex Houtt. and Veltheimia Gled. The differences in estimated dates in Buerki et al. (2012) and Ali et al. (2012) are due to differences in calibration, with Ali et al. (2012) relying on an estimated substitution rate for the trnL-F marker, and Buerki et al. (2012) including a secondary calibration point. The latter is to be preferred. At species level, the southern African squills were last treated by Jessop (1970), and the tropical East African species by Stedje (1996).

The treatment by Jessop (1970) is a superficial one that does not include complete nomenclature, nor descriptions. Since his account, two additional species have been described, viz. Scilla dracomontana Hilliard & B.L.Burtt [now Merwilla dracomontana (Hilliard & B.LBurtt) Speta] (Hilliard and Burtt, 1982) and Spetaea lachenaliiflora (Wetschnig and Pfosser, 2003). The broad circumscription that Jessop (1970) adopted for Scilla natalensis Planch. [now Merwilla plumbea (Planch.) Speta] also requires reconsideration, as does the status of the tropical African M. lazulina Wild. It is thus appropriate and needful that a full, modern account of the species is provided, not only to summarise our current knowledge but also to address some outstanding taxonomic and nomenclatural matters. 2. Materials and methods All relevant types were examined, as well as all herbarium material from BOL, NBG, PRE and SAM (acronyms after Thiers, 2015), the primary collections of southern African species, as well as selected specimens from GRA, NH and NU. All of the species were also studied in the field. Specimens are cited according to the Degree Reference System (Leistner and Morris, 1976). The distribution map for Schizocarphus nervosus is restricted to southern Africa as the species also occurs widely in subtropical and tropical Africa, and details of its occurrence there are provided in Stedje and Thulin (1995) and Stedje (1996). 3. Results 3.1. Generic characters Five species of squill are recognised from sub-Saharan Africa, segregated among the four genera Merwilla (2 spp.), Pseudoprospero (1 sp.), Schizocarphus (1 sp.) and Spetaea (1 sp.). These genera are distinguished morphologically by several characters of the bulb and foliage, inflorescence, perianth, androecium, gynoecium and seed (Table 2), as well as cytologically (Goldblatt et al., 2012). 3.2. Taxonomy Merwilla Speta in Phyton 38: 107 (1998). Type: M. dracomontana (Hilliard & B.L.Burtt) Speta. Etymology: Commemorating the medical inspector and amateur botanist Frederick van der Merwe (1894–1968) who studied the southern African squills and described the genus Schizocarphus. Common name: Mountain squill. Deciduous geophyte: Bulb hypogeal or ± epigeal, subglobose, outer scales firm-textured, papery-cartilaginous, truncate, accumulating.

Table 2 Key morphological characters for the genera of sub-Saharan squills. Genus/Character Bulb outer layers



Papery-cartilaginous Leathery with smooth margin







Leathery-striate with denticulate margin Fibrous with thickened, cartilaginous margin Succulent with smooth margin

Ovary colour/Ovules per locule

Seed colour

Shorter than tepals Pink Shorter than tepals White with Shorter than green tepals gibbosity

White/4 to 11


Yellowish green or purple/2 Dark blue and verrucose/5 or 6

Dark brown

Bract shape

Solitary, erect, simple, lower bracts sterile Solitary, flexuous, simple or branched Solitary or multiple, flexuous, simple

Linear-lanceolate √


Linear-lanceolate √


Linear-lanceolate √



Campanulate Blue to mauve

Solitary, erect, simple

Bracteole Perianth shape

Stamen length



Tepal colour Blue to mauve

Longer than tepals

Yellowish green/2 to 4

Dark brown


J.C. Manning / South African Journal of Botany 125 (2019) 411–426

Leaves several, usually emergent at flowering, prostrate to sub-erect, ovate to lanceolate, leathery, usually hispidulous to velutinous on one or both surfaces, or glabrous, unmarked, margins unthickened, smooth. Inflorescence a solitary (rarely 2), erect, ovoid or cylindrical raceme, several- to many-flowered, scape and rachis glabrous or hispidulous; bracts linear-lanceolate, membranous, not auriculate, lowermost bracts sterile; bracteole present, subulate; pedicels usually longer than flowers, arcuate, glabrous or hispidulous. Flowers rotate, ±deciduous if unpollinated, perianth pale to deep blue or mauve; tepals biseriate with outer series overlapping inner at base, spreading, ± free, uninerved, withering below fruit. Stamens shorter than perianth; filaments connate basally, lanceolate, white; anthers medifxed, blue with white or yellowish pollen. Ovary substipitate, subglobose and 3-lobed, white, with 4 to 11 ovules per locule; style erect, styliform, subequal to or slightly shorter than ovary. Capsules subglobose, substipitate, 3-lobed, leathery. Seeds ellipsoid, beige, lightly rugulose, epidermis areolate‐reticulate. Chromosome number 2n = 40 (Goldblatt et al., 2012). A genus of two species from the eastern seaboard and escarpment of southern and east subtropical Africa, on damp cliffs or in steep rocky grassland. A member of the largely sub-Saharan tribe Massonieae, Merwilla is distinguished by its papery-cartilaginous outer bulb tunics, bracteolate raceme of stellate, blue to mauve flowers with pure white ovary and style, and pale beige or greyish seeds that are unique in a family otherwise characterised by black or dark brown seeds. The lower bracts of the raceme are typically sterile, another unusual feature of the genus. Merwilla and Schizocarphus are both characterised cytologically by a diploid chromosome number 2n = 40, interpreted by Goldblatt et al. (2012) as palaeotetraploid from a base x = 10.


The genus was described by Speta (1998), who also provided the combinations for all five available names as a matter of routine, without any intimate knowledge of the species themselves. This includes the tropical African Merwilla lazulina, which is here included in M. plumbea. Key to species 1. Leaves prostrate, often fully developed at flowering; inflorescence 60–120 mm long; scape and rachis hispidulous ... M. dracomontana 1′. Leaves sub-erect, at length spreading, usually emergent at flowering; inflorescence 150–1 500 mm long; scape and rachis glabrous ... M. plumbea. Merwilla dracomontana (Hilliard & B.L.Burtt) Speta in Phyton 38: 109 (1998). Scilla dracomontana Hilliard & B.L.Burtt in Notes. Roy. Bot. Gard. Edinb. 40: 283 (1982). Type: South Africa, KwaZulu-Natal, Underberg (2929): ‘Estcourt distr., Game Pass’, (–BB), 28 Oct 1973, Hilliard & Burtt 6960 (E, holo.–image!; NU[4348]!, iso.). Deciduous geophyte: Bulb hypogeal or partly epigeal, subglobose, 30–40 mm diam., outer scales papery-cartilaginous, chestnut brown. Leaves emergent or more often fully formed at flowering, 3 to 5, prostrate or adpressed, ovate to broadly ovate (rarely elliptic), acute, 20–80 × 10–35 mm, sparsely to densely velutinous adaxially with patent hairs up to 1 mm long, similarly but less densely hairy beneath, dull green when fresh. Inflorescence a dense, ovoid to cylindrical raceme 60–110(− 120) mm long, scape erect, 1.0–1.5 mm diam. at base, hispidulous throughout (including rachis) with patent or deflexed hairs, flushed maroon, flower-bearing rachis one third to twice as long as scape, 20–30 mm diam., many-flowered; bracts linear-lanceolate to subulate, attenuate, membranous, 3–10 × 0.5–2.0 mm, glabrous or sparsely ciliate, with small subulate bracteole up to 3 mm long; pedicels spreading or arcuate, 8–18 mm long, glabrous or sparsely hispidulous.

Fig. 1. Distribution of Merwilla dracomontana (▲), Pseudoprospero firmifolium subsp. firmifolium (●) and P. firmifolium subsp. natalensis (○).


J.C. Manning / South African Journal of Botany 125 (2019) 411–426

Flowers spreading, unscented, perianth rotate, deep blue or rarely white; tepals free, spreading, elliptic, 4–6 × 1.5–2.0 mm. Stamens connate basally for up to ± 0.5 mm; filaments lanceolate, 2.0–2.5 × 0.8 mm, white; anthers 1.0–1.2 mm long at anthesis. Ovary subglobose, substipitate, 3-lobed, 1.5–2.0 mm diam., with 4 to 6 ovules per locule, white; style ±1.5 mm long. Capsules suberect, substipitate, depressedsubglobose to turbinate, deeply 3-lobed, 5–7 × 6–8 mm. Seeds compressed-ellipsoid or teardrop-shaped, longitudinally 3-angled, ±4 × 1.5–2.0 mm, dull beige flushed mauve, testa lightly wrinkled. Flowering time: Sep to Oct. Distribution and ecology: Originally thought to be restricted to the southern KwaZulu-Natal Drakensberg and its foothills between Giant's Castle and the upper Mzimude Valley near Underberg, Merwilla dracomontana is now also known further south from Mt. Currie near Kokstad in the Eastern Cape (Fig. 1); on sandstone cliffs and crevices or on open ground, between 1 600 and 2 100 m elevation. A specimen cultivated at Kirstenbosch and labelled as originating from Umkomaas in KwaZulu-Natal (Archbell s.n.) is also this species. Although Umkomaas itself lies at the mouth of the Mkomaze River, it is more likely that the locality in question refers to the upper reaches of the river, which rises at Giant's Castle, from where the species is known. Diagnosis: Distinguished from smaller, hairy-leaved plants of Merwilla plumbea by the prostrate leaves that are often welldeveloped at flowering time, and the short inflorescence up to 120 mm long but usually less, with hispidulous scape and rachis. The scape is slender, 1.0–1.5 mm diam. at the base, and relatively short in relation to the flower-bearing rachis, often shorter than the rachis but at most twice as long. The flowers have 4 to 6 ovules per locule. The foliage in M. plumbea is emergent or partially developed at flowering and suberect to spreading, and it is a taller species with an inflorescence more than 1 200 mm long with a glabrous scape and rachis, and mostly more (usually 6 to 11) ovules per locule. History: Merwilla dracomontana appears to have been first collected in the spring of 1914 on Mt. Currie near Kokstad by local game warden and naturalist Roden Symons (1884±1974) but this collection has been overlooked until now. It, like other early gatherings of the species, was initially identified as Scilla kraussii until veteran botanists Olive Hilliard (b. 1925) and Brian ‘Bill’ Burtt (1913–2008) recognised that they represnted a distinct taxon. Other specimens seen South Africa: KWAZULU-NATAL. 2929 (Underberg): Giant's Castle, (–AC), 29 Sep 1914, Symons 20 (PRE, SAM); Giants Castle Game Reserve, (–AC), 15 Sep 1965, Trauseld 407 (PRE); 25 Sep 1957, Edwards 2165 (PRE); turf roof of hut, 17 Nov 2001 [fruiting], Manning 2671 (NBG); Mkomazi State Forest, (–AC), 18 Sep 1984, Scott 230 (NU); Highmoor Forest Reserve, (–BC), 25 Sep 1977, Ruddock 31 (NU); 10 Nov 1977 [fruiting], Du Toit 2557 (PRE); Underberg, 800 m from Burring's Cottage, (–CA), 3 Dec 2008 [fruiting], Ngwenya 3304 (PRE); Cobham Forest Station, Ndlovini, Frontbeck, (–CB), 8 Sep 1980, Hilliard and Burtt 13339 (NU); Sani Pass, (–CB), 15 Dec 1984 [fruiting], Hilliard and Burtt 17,997 (PRE); Garden Castle Forest Reserve, Sleeping Beauty Cave Valley, (–CD), 3 Sep 1981, Hilliard 8158 (NU); Lion's River Dist., Inhlazane, (–DB), 15 Oct 1964, Moll 1289 (PRE); Lion's River Dist., Mgeni Poort, (–DB), 7 Sep 1965, Moll 1934a (PRE). 3029 (Kokstad): Mt. Currie peak in foothills of Drakensberg, (−AD), 20 Oct 1962, Van der Riet s.n. (NBG). Uncertain locality: Umkomaas, 11 Oct 1943 [ex. hort. Kirstenbosch Rockery], Mrs J.E. Archbell s.n. NBG2335/32 (NBG). Merwilla plumbea (Lindl.) Speta in Phyton 38: 109 (1998). Scilla plumbea Lindl. in Bot. Reg. 16: t. 1355 (1830). Type: ‘from a drawing made in 1813, in the Kew Garden, by the late Mr. Sydenham Edwards’, illustration in Bot. Reg. 16: t. 1355. Scilla natalensis Planch. in Flore de Serres 10: 185, t. 1043 (1855); Baker in J. Linn. Soc., Bot. 13: 243 (1873); Baker in Fl. cap. 6: 482 (1897); Phillips in Fl. Pl. Afr. 10: t. 365 (1930); Jessop in Jl. S. Afr. Bot. 36: 241 (1970). Merwilla natalensis (Planch.) Speta in Phyton 38: 109

(1998). Type: South Africa, ‘Port Natal, cultivated by Van Houtte Nursery’, illustration in Flore des Serres: t. 1043 (1855). Scilla natalensis var. sordida Baker in J. Linn. Soc., Bot.: 243 (1973); Baker in Fl. Cap. 6: 482 (1897). Type: South Africa, KwaZulu-Natal, ‘Natal, cultivated specimen’, Apr 1864, without collector (K[257444], holo.–image!). Scilla lazulina Wild in Kirkia 4: 163 (1964), syn. nov. Merwilla lazulina (Wild) Speta in Phyton 38: 109 (1998). Type: Zimbabwe, ‘Chimanimani Mts, The Corner’, 13 Oct 1950, Sturgeon GHS30434 (SRGH, holo.; K[257441]–image!, PRE[792895]!, iso.). Deciduous geophyte: Bulb hypogeal or partly epigeal, ovoid or subglobose, 20–150 mm diam., outer scales becoming paperycartilaginous. Leaves 2 to 8, suberect but later spreading, often involute-clasping below, oblong to ovate-lanceolate, acute to acuminate, 20–450(−600) × 5–100 mm, leathery, glabrous or more usually sparsely to densely hispidulous on the veins (rarely intercostal area as well) on both surfaces or only beneath, sometimes only towards base, with patent or deflexed hairs up to 0.5 mm long, rarely velutinous with patent hairs to 2 mm long, glaucous or bright green when fresh, sometimes flushed maroon beneath. Inflorescence a dense, ovoid to cylindrical raceme, 150–1 000(−1 500) mm long, scape erect, 2–15 mm diam. at base, glabrous throughout, glaucous, flower-bearing rachis usually one third to one half as long as scape, 60–500 mm diam., manyflowered; bracts linear-lanceolate to subulate, attenuate, membranous, 5–15 × 0.5–2.0 mm, glabrous, with small subulate bracteole up to 3 mm long; pedicels spreading or arcuate, 10–30 mm long, glabrous. Flowers spreading, unscented or sweetly (honey) scented, perianth rotate, pale

Fig. 2. Merwilla plumbea subsp. plumbea, flowering plant. Plant of unknown origin, cultivated at the Division of Plant Industry, Pretoria. Original painting reproduced in Flowering Plants of South Africa 10: t. 365 (1930). Artist: Cythna Letty. Note: The watercolour pigment used to colour the tepals has faded with age from the original blue-mauve, as reproduced in Flowering Plants of South Africa 10: t. 365 (1930).

J.C. Manning / South African Journal of Botany 125 (2019) 411–426

to deep blue or mauve, rarely pink or white; tepals free, spreading, elliptic, 5–10 × 1.5–3.5 mm. Stamens connate basally for up to ±0.5 mm, filaments lanceolate, 3–5 × 0.8–1.5 mm, white; anthers 1.5–2.5 mm long at anthesis. Ovary subglobose, substipitate, 3-lobed, 2.5–3.0 mm diam., white, with (5)6 to 11 ovules per locule; style ±2.5 mm long. Capsules suberect, substipitate, depressed-subglobose to turbinate, deeply 3-lobed, 6–9 × 8–10 mm. Seeds compressed-ellipsoid or teardropshaped, longitudinally 3-angled, ±6.5 × 3.5 mm, dull beige, testa lightly wrinkled. Flowering time: mainly Sep to Nov, but sometimes earlier or later. Figs. 2 and 3. Distribution and ecology: Widely distributed along the eastern escarpment of southern Africa and subtropical Africa, from Kentani in the Eastern Cape through KwaZulu-Natal and eastern Free State to Graskop in Mpumalanga and Tzaneen and Soutpansberg in Limpopo, and in Lesotho and Swaziland (Fig. 4), and further north in east subtropical Africa, where it has been recorded from the Chimanimani Mts in eastern Zimbabwe, on the Namuli Hills in Mozambique, and on Mt. Mlanje in Malawi (Wild, 1964); stony or rocky slopes and cliffs, often seasonally damp, usually in colonies, or in moist depressions and then solitary. Flowering is strongly stimulated by a winter burn. The large forms of Merwilla plumbea are extremely decorative, producing tall racemes of numerous pale blue or mauve, starry flowers in the early spring. They are typically found in numbers colonising cliffs or rocky slopes. Pink-flowered plants are not uncommon in some populations and occasional albinos are also known. The plants survive light winter snowfalls in the wild and are probably hardy to several degrees below zero. The smaller subsp. kraussii is not robust enough for the garden, and although it can be planted in a rockery it is really best grown in pots.

Fig. 3. Merwilla plumbea subsp. kraussii, flowering plant, detail of bract and bracteole, and detached flower showing white ovary. South Africa, Eastern Cape, Mkambati River, Mogg PRE26403 (PRE). Original painting reproduced in Flowering Plants of South Africa 21: t. 822 (1941). Artist: Cythna Letty.


Merwilla plumbea is an important and widely used component of the traditional South African pharmacopoeia, and the intense harvesting of wild plants is a severe threat to the survival of the species in some localities (Williams et al., 2008; Raimondo et al., 2009). Decoctions of the bulb are used to treat a variety of ailments, both internal and external. Although little medical evidence exists to support the veracity of its many alleged benefits, chemicals of the class known as homoisoflavonones have been isolated from the bulbs and are known to possess anti-inflammatory properties (Douwes et al., 2001). The sap is reportedly corrosive to sensitive skin and this property is certainly evident in other members of the Hyacinthaceae. Plants are also recorded to be toxic to stock (Kennedy and Sleed 33 NU). Diagnosis: A small to large species with an inflorescence 150–1 000 (−1 500) mm long, with a glabrous scape 2–15 mm diam. at the base. The two to eight suberect or spreading leaves are oblong to ovatelanceolate, 50–600 × 5–100 mm, more or less emergent at flowering, and are either glabrous or sparsely to densely hispidulous or velutinous, usually on both sides but sometimes only beneath, sometimes only in the juvenile stages. The number of ovules varies from 5 to 11 per locule. The smaller Merwilla dracomontana is recognised by its prostrate leaves that are well-developed at flowering, and a more delicate inflorescence up to 120 mm long but usually less, with diagnostic hispidulous scape and rachis, and 4–6 ovules per locule. A collection of M. plumbea from Sani Pass with velutinous leaves (Hilliard & Burtt 17,880 NU) was misidentified by the collectors as a hybrid with M. dracomontana, presumably on the basis of its hairy leaves since the KwaZuluNatal Drakensberg populations of the species mostly have glabrous foliage at maturity. The taxonomy of Merwilla plumbea is complicated by the significant amount of local variation in the taxon. Smaller forms from KwaZuluNatal were segregated under the name Scilla kraussii by Baker (1873, 1896–1897) on the basis of their fewer (± four), consistently pubescent leaves 50–80 mm long, and shorter inflorescence 150–200 mm tall. Typical M. plumbea (= S. natalensis), which was known in the wild to Baker (1896–1897) from a handful of collections along the eastern Escarpment, viz. from Baziya in the Eastern Cape to Barberton in Mpumalanga, included larger plants with six to eight, glabrous or pubescent leaves 300–450 mm long and an inflorescence 150–300 mm long. Additional collecting has, however, blurred this distinction. Most populations include substantial variation in plant size, some of it undeniably ecological and developmental. In this respect, the notes appended to an herbarium collection from Tzaneen by the missionary and prolific plant collector Frederick A. Rogers (Rogers 12507 BOL) are worth quoting in full: ‘If you ever have to deal with Scilla natalensis (sic.) it may interest you to know that when it is in flower it has a rosette of leaves which lie on the ground & are 4 or 5 inches long; after flowering they develop in their natural state (i.e.) among rocks to 8-10 in. long & remain hairy, but when they grow in a vlei or are transplanted into a garden they generally lengthen very much, up to 18-24 inches become upright, & quite glabrous. I have actually seen this happen. Also the division of Scillas according to the shape & size of the bulbs is quite hopeless. The same plant growing in masses has bulbs of various shapes & of very various sizes’. F.A. Rogers [Signed]. Jessop (1970) also recorded substantial variation in plant size in populations near Kamberg and the Oliviershoek Pass in KwaZuluNatal, and possibly also near Tzaneen in Limpopo although this observation might have been derived from Roger's notes (see above). As a result of his observations that ‘pubescence may not be entirely a genetically determined phenomenon and in view of the range in size of the pubescent-leaved plants’, he treated Scilla kraussii and S. natalensis as representing a single taxon. In contrast, Douwes et al. (2001) preferred to recognise the two as distinct species on the basis of observations made at Howick in the KwaZulu-Natal Midlands, where they reported ‘large populations of both S. kraussii and typical S. natalensis occurring sympatrically and flowering together’ without evidence of any intermediates. Unfortunately, they provide no details on either the ecology or


J.C. Manning / South African Journal of Botany 125 (2019) 411–426

Fig. 4. Distribution of Merwilla plumbea subsp. plumbea (●) and subsp. kraussii (○).

the morphological differences between the two forms that they observed. Significantly, Hilliard and Burtt (1982), who were very familiar with the KwaZulu-Natal flora through their extensive field work in the province, were themselves unsure whether the two forms represented the ends of a continuum or not. This is despite the fact that they had collected both forms from the same locality at Forest Hills near Pinetown in KwaZulu-Natal, the larger, glabrous-leaved plants growing in clefts on cliff faces (Hilliard 1617, NU) and smaller, pubescent-leaved plants in adjacent grassland and on rocky hillsides (Kays 46, NU; Hilliard 1622, NU). They astutely observed, however, that ‘S. kraussii sensu stricto does not appear to grow in the [KwaZulu-Natal] Drakensberg’ (Hilliard and Burtt, 1982). The abundant collections that are now available make the distinction between Merwilla kraussii and M. plumbea essentially arbitrary, and although exceptionally small plants are convincingly assigned to the former and large ones to the latter, the existence of populations and even of individuals within populations that are intermediate between the two taxa makes it impossible to segregate the available material unequivocally into two sets of populations on the basis of plant size or any other combination of characters. Some of the intra-population variation in inflorescence and foliage size is linked to the fact that flowering takes place early in the growing season when the leaves are emergent and thus not yet fully formed. Individuals in a population that flower later in the season will therefore have larger, more fully expanded leaves than those flowering earlier. This does not explain all of the variation however, and I have grown the two extremes in cultivation for over a decade without any significant change in their morphological characteristics. This is conclusive evidence for some genetic differentiation between at least some of these populations.

There is also a broad correlation between ovule number and plant size. Examination of several collections of the smaller ‘kraussii’-type plants shows the flowers to have 6 to 8(9) ovules per locule, whereas larger ‘plumbea’-type plants have 8 to 11 ovules per locule across their distribution in southern Africa (see Additional specimens cited for counts). It seems to me significant, however, that there is a discernible geographical pattern to the morphological variation. The smaller ‘kraussii’type plants occur in the southern part of the species' range, at lower altitudes in KwaZulu-Natal and the Eastern Cape, mostly on shallow, sandstone soils, whereas larger plants of the ‘plumbea’-type occur further inland in the Eastern Cape and KwaZulu-Natal, and reach a substantial size along the KwaZulu-Natal and Mpumalanga Drakensberg. Both forms have been recorded near Port St. Johns in the Eastern Cape, around Dumisa in southern KwaZulu-Natal, at Kloof inland of Durban, and in the Karkloof in the KwaZulu-Natal Midlands. The identification of several collections between Durban and Kokstad is almost arbitrary. On the basis of this general, albeit incomplete, morphological differentiation between the coastal and inland populations of Merwilla plumbea, it seems appropriate and informative to recognise the coastal plants at subspecific rank as subsp. kraussii. The relationship between these two forms is certainly more complex and requires detailed cytogenetic study. A preliminary study of genetic diversity in the group using random amplified polymorphic DNA by Van Staden and Pan (2001) successfully identified M. dracomontana as distinct from the M. kraussii/plumbea complex but although it also identified differences in the RAPD profiles of M. kraussii, M. plumbea and the artificial hybrid between them, the sampling comprised a single collection for each taxon and is thus an inadequate basis for any taxonomic conclusions.

J.C. Manning / South African Journal of Botany 125 (2019) 411–426

Fig. 5. Pseudoprospero firmifolium Subsp. firmifolium, detached leaf and inflorescence with short branch, and detached flower. South Africa, Eastern Cape, Bathurst Dist., Dyer 3374 (NBG, NH, PRE). Original painting reproduced in Flowering Plants of South Africa 24: t. 926 (1944). Artist: E. K. Burges.

Merwilla lazulina (Wild) Speta, from the eastern highlands of Zimbabwe, southern Malawi and adjacent northern Mozambique, was described by Wild (1964) as having 2 to 5(9), glabrous, linear to lanceolate leaves 40–120 × 60 mm, and an inflorescence 130–170 mm long. It was distinguished from M. kraussi by its glabrous foliage and fewer (5) ovules per locule, and from typical M. plumbea by its smaller size and unspecified differently shaped leaves. Additional collections, however, show the Zimbabwean material to be far more variable in size and in foliar vestiture. A collection from Chatsworth in east central Zimbabwe (Solomon s.n., NBG) has hispidulous leaves up to 300 × 35 mm, the inflorescence 300–500 mm long with pedicels 7–15 mm long, and flowers with 8 or 9 ovules per locule. On all considerations, therefore, the type of M. lazulina and other collections from eastern Zimbabwe are indistinguishable from M. plumbea as circumscribed here. This applies also to collections from Mlanje in Malawi, which resemble the Chatsworth plants in stature, with the inflorescence 350–500 mm long, but have wider, glabrous leaves. These characteristics have been maintained in a plant from Mlanje that I have had in cultivation for many years. Merwilla lazulina is accordingly reduced to synonymy under M. plumbea. The plants from the Chimanimani Mts are consistently smaller than is usual for subsp. plumbea, matching subsp. kraussii in their stature and floral dimensions but differing from it in their glabrous foliage, as was originally noted by Wild (1964), and in some plants also by their much larger bulbs, 80 × 40 mm. They evidently represent a local form or ecotype from shallow soils on sandstone or granitic rock. History: This species was long known under the name Scilla natalensis, described from a cultivated plant that had been introduced to Europe from Port Natal [Durban] by the van Houtte nurseries in Ghent (Planchon, 1855). The earlier S. plumbea, described by Lindley (1830) from a drawing made by Sydenham Edwards in 1813 of a


plant of unknown provenance cultivated at Kew, was suggested by Baker (1896–1897) to represent the same species but this synonymy was only formalised by Manning et al. (2004). Other authors, including Lewis (1947) and Jessop (1970) had meanwhile followed Kunth (1843) in incorrectly associating the name S. plumbea with the Western Cape endemic Spetaea lachenaliiflora, until the independent identity of that species was established by Wetschnig and Pfosser (2003). Two unlocalised collections of smaller plants from KwaZulu-Natal made by William Gerrard (? to ± 1866) and Christian Krauss (1812–1890) in the mid-1800s formed the basis for Scilla kraussii Baker (1873). Several additional collections from the KwaZulu-Natal Midlands and Transkei were later also identified as this species by Baker (1896–1897). Scilla kraussii was distinguished solely by its smaller size from S. natalensis, which was known in the wild from a handful of collections from the eastern Escarpment from Baziya in the Eastern Cape to Barberton in Mpumalanga (Baker, 1896–1897). Additional collections since then have blurred the distinction between S. natalensis and S. kraussii and the two were accordingly united with some hesitation by Jessop (1970), although this conclusion was later rejected by Douwes et al. (2001) on the basis of limited field observations. A cultivated specimen of S. natalensis that flowered at Kew, with brown-tinged foliage and fewer flowers than typical, was recognised as var. sordida Baker (1873, 1896–1897) but this variant has no taxonomic significance. Scilla lazulina was described by Wild (1964) for several collections from east subtropical Africa but these differ in no significant way from M. plumbea. Conservation status: Although relatively widespread, Merwilla plumbea subsp. plumbea is harvested for the medicinal trade and is regarded as Near Threatened (Raimondo et al., 2009). The conservation status of subsp. kraussii is unknown. The latter taxon is relatively rangerestricted but its smaller size may protect it from excessive harvesting. Key to subspecies. 1a. Leaves 3 to 8, 100–450(−600) × 20–100 mm, glabrous or hispidulous on one or both surfaces; inflorescence (130–)300–1 500 mm long; pedicels (5–)15–30 mm long; ovules (5)8 to 11 per locule; plants from near-interior Eastern Cape and KwaZulu-Natal and eastern Escarpment to subtropical East Africa ... subsp. plumbea. 1b. Leaves 2 to 4, 60–100 × 5–10(−30) mm, always hispidulous on both surfaces; inflorescence 150–400 mm long; pedicels 10–15(−20) mm long; ovules (5)6 to 8(9) per locule; plants from near-coastal KwaZulu-Natal and Eastern Cape ... subsp. kraussii. a. subsp. plumbea. Bulb (30–)50–100(−150) mm diam. Leaves (3)5 to 8, lanceolate to ovate-lanceolate, 200–450(−600) × 20–100 mm, often involuteclasping below, glabrous or hispidulous, sometimes only beneath, rarely velutinous. Inflorescence 300–1 500 mm long, scape 3–15 mm diam. at base, rachis 80–500 × 20–70 mm; pedicels (5–)15–30 mm long. Tepals 6–10 mm long. Filaments 4–5 mm long. Ovules 8 to 11 per locule. Fig. 2. Distribution: Distributed in eastern South Africa, from Kentani in the Eastern Cape through near-interior and interior KwaZulu-Natal and the Drakensberg into the eastern Free State and Lesotho, and north along the Escarpment to Graskop in Mpumalanga and on the higher lying Wolkberg and Soutpansberg in Limpopo and also Swaziland, and further north in the eastern highlands of Zimbabwe, on Mlanje in southern Malawi and on Mt. Namuli in northern Mozambique (Fig. 4); on basalt, sandstone, dolerite and granite cliffs and rocky slopes, often in seepages, and also wet grassland, from near sea level in the south around Kei Mouth in Eastern Cape and Hluhluwe and Mtunzini in northern KwaZulu-Natal up to 2 000 m elevation. Diagnosis: Subsp. plumbea includes medium-sized to large plants with mostly five to eight, suberect or spreading, oblong to ovatelanceolate leaves, 100–600 × 30–100 mm when fully developed. These may be glabrous but are more usually sparsely to densely hispidulous or even velutinous, usually on both sides but sometimes only beneath, sometimes only in the juvenile stages. The inflorescence is 300–1000(−1 500) mm long, with pedicels (5–)15–30 mm long,


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and flowers with tepals 6–10 mm long, and mostly 8 to 11 ovules per locule. Hispidulous-leaved populations occur through much of the range of the subspecies and reach a substantial size in Mpumalanga. Some collections from there and Limpopo represent unusually hairy forms, with velutinous leaves densely covered in patent hairs up to 2 mm long. In contrast, large, glabrous-leaved forms are characteristic of the higher reaches of the KwaZulu-Natal Drakensberg, and although seedling plants of these plants have hairy leaves this is usually lost by maturity. Smaller plants of subsp. plumbea are not always easily distinguished from subsp. kraussii from near-coastal KwaZulu-Natal and the Eastern Cape but that subspecies has 2 to 4 leaves, 60–100 × 5–10(−30) mm, that are always hispidulous on both surfaces, a shorter and often narrower inflorescence 150–400 mm long with pedicels 10–15(−20) mm long, and flowers with tepals 5–6 mm long, and mostly up to 8 ovules per locule. Other specimens seen Lesotho. 2828 (Bethlehem): Leribe, (–CC), mt slopes, ‘spring’ [sic.], Dieterlen 416 (NBG, PRE, SAM). Malawi: Mt. Mulanje, Litchenya Plateua near Linji stream, without date, Chapman and Chapman 8230 (PRE). South Africa. LIMPOPO. 2329 (Pietersburg): Soutpansberg, (–AB), 11 Oct 1955, Mackay s.n. (NBG). 2330 (Tzaneen): Tzaneen, (–CC), Nov 1913, Rogers 12507 (BOL); Woodbush, (–CC), Sep 1909, Noome 7458 (PRE); 14 Oct 1943, Taylor NBG1732/35 (NBG) [11 ovules per locule]; Wolkberg, on diabase outcrop, 20 Nov 1945, Gerstner 5665 (PRE); Haenertsburg, (−DD), Sep 1913, Pott 4470 (PRE). MPUMALANGA. 2430 (Pilgrim's Rest): Letaba Dist., escarp cliffs directly opposite Metz Mission Station, (−AA), 18 Nov 1976, Venter 1190 (PRE); Serala Forestry, Farm Klipdraai, (−AA), dolomites, 17 Sep 1985, Venter 10940 (PRE); Lekgalameetse Nature Reserve, The Downs, (−AA), 22 Oct 1985, Stalmans 725 (PRE); Mariepskop, (–DB), Oct 1925, Fitzimons and Van Dam s.n. (PRE); 20 Oct 1962, Van der Schijff 6172 (PRE); Morgenzon Forest Reserve, (–DC), 27 Nov 1983, Van der Zeyde s.n. (NBG); MacMac Falls, (−DD), 3 Aug 1978 [fl. ex hort. Kirstenbosch], Van Jaarsveld 3412 (NBG); marshy grassland next to R534 6 km from God's Window to Graskop, (−DD), 17 Nov 1999 [fruiting]. Riddles 551 (PRE). 2529 (Witbank): Middelburg, N of Tantesberg, (–BB), Oct 1935, Obermeyer s.n (PRE); Tantesberg, (–BB), 9 Nov 1933, Young A244 (PRE); Tantesberg, Diepkloof, (–BB), without date, Van der Merwe 2224 (PRE). 2530 (Lydenburg): Dullstroom, (–AC), 10 Nov 2003 [fruiting], Custers s.n. (PRE); Dullstroom Dist., Verlorenvallei, (–AC), 16 Oct 1980, Drews 70 (PRE); 24 km from Dullstroom on road to Lydenburg, (–AC), 30 Sep 1971, Balsinhas and Kersberg 2090 (PRE); Lydenburg Dist., Hartebeesvlakte, (–BA), 1 Jul 1980, Mohle 376 (PRE); De Kuilen Farm, 9 mi [14.4 km] SE of Lydenburg, (–BA), 21 Oct 1975, Anderson A17 (PRE); Buffelskloof Nature Reserve, (–BC), 26 Oct 1982, Onderstall 911 (PRE); near Brondall, (–BD), May 1938, Van der Merwe 1682 (PRE); Tomango [Mataffin], (–BD), 14 Oct 1943, Holland NBG490/27 (NBG); Waterval-Onder, (–CB), 1 Oct 1906, Rogers 2596 (BOL, PRE); Barberton, (−DD), without date, Thorncroft ex Rogers 19178 (NBG, SAM); Kaapsche Hoop, (–DB), 2 Oct 1949, Prosser 1279 (NBG, PRE); Thorncroft Nature Reserve off Agnes Mine road, (−DD), 12 Nov 1968, Edwards 4111 (PRE); grassy slopes around Barberton, (−DD), large bulb partly above ground, Oct 1889, Galpin 619 (BOL, PRE); Barberton, top of Nelshoogte Pass, (−DD), 19 Nov 2001, Manning 2685 (NBG). 2630 (Carolina): Ermelo Dist., The Gem, (–BC), without date, Walker s.n. (PRE). 2730 (Vryheid): Wakkerstroom, Oshoek, (−AD), 24 Sep 1960, Devenish 101 (PRE); Piet Retief, (–BD), Sep 1934, Bowker 196 (PRE). FREE STATE. 2828 (Bethlehem): Witsieshoek, (–DB), Dec 1905, Thode STE5384 (NBG); 15 Aug 1985, Paton 301 (PRE). 2829 (Harrismith): Platberg near Harrismith, (−AA), 14 Nov 1973, Muller 1277 (PRE); Eastern Free State and Drakesnberg Botanic Garden, (–AC), 13 Oct 1969 [ex hort.], van der Zeyde s.n. (NBG); Farm Arcadia 4 km from Harrismith, (–AC), 3 Oct 1971, van der Zeyde 84 (NBG); Farm Wyford, (−AD), W-facing

sandstone ledges, 23 Nov 1971, van der Zeyde s.n. (NBG); Golden Gate Highland National Park, (−DA), 26 Oct 1963, Liebenberg 7230 (PRE). KWAZULU-NATAL. 2729 (Volksrust): Newcastle Dist., Ncandubo Reserve, (−DD), 30 Sep 1989, Smit 1284 (PRE). 2828 (Bethlehem): [Royal] Natal National Park, (–DB), 26 Nov 1928, Galpin 10149 (PRE); [Royal] Natal National Park, Tiger Falls, (–DB), 26 Oct 1955, Edwards 975 (PRE). 2829 (Harrismith): Olivershoek Pass, (–CC), Dec 1908, Thode STE3374 (NBG) [8 or 9 ovules per locule]; Cathedral Peak, (–CC), 1959, Ruch 1660a (PRE); 6 Oct 1982, Dohse 66 (PRE). 2831 (Nkandla): Mtunzini, (−DD), 1 May 1919, Mogg 4482 (PRE). 2832 (Mtubatuba): Hluhluwe Game Reserve, (−AC), 16 Oct 1953, Ward 1559 (NU). 2929 (Underberg): Giants Castle Game Reserve, (–BC), 29 Sep 1914, Symons 19 (PRE, SAM); 14 Nov 1967, Trauseld 847 (PRE); 17 Nov 2001, Manning 2672 (NBG); Kamberg, Game Pass, (–BD), 15 Oct 1984, Stutterheim 21 (PRE); 10 km from Himeville to Sani Pass, (–CB), Sep 1973, Arnold 536 (PRE); Sani Pass, (–CB), 12 Dec 1984 [fruiting], Hilliard and Burtt 17880 (NU); 14 Dec 1984 [fruiting], Hilliard and Burtt 17962 (PRE); Bulwer, (−DD), 28 Sep 1940, Putterhill s.n. (PRE). 2930 (Pietermaritzburg): Tweedie, (–AC), 27 Sep 1918, Mogg 1303 (PRE); Balgowan, (–AC), 29 Sep 1965, Moll and Mauve 2445 (PRE); Balgowan, Millbank, (–AC), 12 Oct 1964, Moll 1192 (NU, PRE); Caversham, (–AC), 5 Oct 1918, Mogg 2458 (PRE); Sevenoaks, Canema Farm, (–BA), 11 Oct 1989, Ngwenya 694 (PRE); New Hanover, Blinkwater Nature Reserve, (–BC), 3 Oct 1995, Sikhakhane 541 (PRE); Taylor's Halt, (–CA), 2 Nov 1943, Fairall NBG767/39 (NBG); Pietermaritzburg Botanic Garden Estate, (–CB), 27 Oct 1991, Stielau 123 (PRE); Pietermaritzburg, Old Howick Rd, (–CB), 22 Sep 1987, Kennedy and Sleed 33 (NU); Drummond, Alverston Ridge, (–DC), 14 Nov 2001 [fruiting], Manning 2658A (NBG); Kloof, Forest Hills, (–DD), edge of cleft in cliff face, in grassveld, 27 Aug 1963, Hilliard 1617 (NU). 3030 (Port Shepstone): Dumisa, Fairfield Farm, (−AD), 21 Oct 1997, Ngcobo 34 (PRE); Dumisa, Campbelltown, (−AD), moory pasture, 28 Sep 1913, Rudatis 1994 (NBG); Alexandra County, Umgage [Umgababa] flats, (–BB), 6 Aug 1909, Rudatis 673 (NBG, PRE); Sezela, (–BC), 7 Feb 1930, Reynolds NBG599/28 (BOL). EASTERN CAPE. 3029 (Kokstad): Bizana, (−DD), 25 Oct 1962, Strey 4497 (PRE). 3127 (Lady Frere): hills near Engcobo, (–DB), 13 Oct 1961, Esterhuysen 29247 (BOL). 3128 (Umtata): 3 mi [5 km] inland of Port Grosvenor, (–BD), 23 Aug 1969, Strey 8907 (PRE); Baziya, (–CB), without date, Baur s.n. (SAM). 3129 (Port St Johns): E Pondoland, Egossa, (–BC), Aug 1899, Sim 2538 (BOL, PRE); Magwa Estates, (–BC), 7 Sep 1979, Germishuizen 1198 (PRE); edge of cliffs 1 km E of Fraser's Falls, (–BC), 5 Sep 1998, Paterson-Jones 831 (NBG) [ovules 10 per locule]; Mkambati, (–BD), 7 Aug 1956, Hone 41 (PRE). 3227 (Stutterheim): Komgha Dist., among rocks near Kei River, (–DB), Nov 1889, Flanagan 356 (BOL, PRE, SAM). 3228 (Butterworth): Kentani, (–CB), on plains, Sep 1902, Pegler 850 (BOL, PRE); grassy hillsides, 1500′ [460 m], 30 Sep 1961, Batten 3 (NBG). Swaziland: 2531 (Komatipoort): Pigg's Peak, (–CD), 2 Oct 1962, Compton 31566 (NBG, PRE). 2631 (Mbabane): Malolotja Nature Reserve, (−AA), dolomite rocks on west side of stream, 22 Oct 1985, Runciman 13 (PRE); Forbes Reef, (−AA), May 1915, Roberts s.n. (PRE); Poliniane River, (–AC), 7 Oct 1958, Dlamini s.n. (PRE); Kirkhill, (–AC), streambank, 12 Oct 1956, Compton 26080 (PRE); Motshane Halt, (–CD), 24 Oct 1956, Compton 26141 (NBG, PRE); Hlatikulu, (–CD), 21 Oct 1958, 3500′ [1 100 m], Compton 28146 (NBG, PRE); between Goedgegun and Hlatikulu, (–CD), 30 Sep 1965, Compton 32417 (NBG, PRE). b. subsp. kraussii (Baker) J.C.Manning, comb. et stat. nov. Scilla kraussii Baker in J. Linn. Soc., Bot. 13: 243 (1873); Baker in Fl. cap. 6: 481 (1897); Van der Merwe in Fl. Pl. S. Afr. 21: t. 822 (1941b). Merwilla kraussii (Baker) Speta in Phyton 38: 109 (1998). Type: South Africa, KwaZulu-Natal, Stanger (2931): ‘Port Natal’, (–CC), 1840, Krauss 444 (K[257447], lecto.—image!, designated by Jessop: 241 (1970); MO [104954]–image!, isolecto.). Bulb 20–40 mm diam. Leaves 2 to 4, oblong to lanceolate, 60–100 × 5–10(−30) mm, hispidulous on both surfaces. Inflorescence 150–400 mm long, scape 2–4(–5) mm diam. at base, rachis

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60–200 × 20–40 mm; pedicels 10–15(−20) mm long. Tepals 5–6 mm long. Filaments 3–4 mm long. Ovules (5)6 to 8(9) per locule. Fig. 3. Distribution and ecology: Restricted to coastal and near-interior southern KwaZulu-Natal and adjacent Eastern Cape, mainly from Nkandla and Krantzkloof to Port Edward and Port St. Johns but also recorded further inland from the Karkloof and Noodsberg in the KwaZulu-Natal Midlands and Kokstad in the Eastern Cape (Fig. 4), from near sea-level to 1 000 m elevation; on open, stony ground and cliffs. Diagnosis: Subsp. kraussii includes small to medium-sized plants from near-coastal KwaZulu-Natal and Eastern Cape with two to four, suberect to spreading leaves, 60–100 × 5–10(−30) mm, that are always hispidulous on both surfaces, and an inflorescence 150–40 0mm long, with pedicels 10–15(−20) mm long, and flowers with tepals 5–6 mm long and mostly up to 8 ovules per locule. Typical Merwilla plumbea includes larger plants, with mostly five to eight leaves, 100–600 × 30–100 mm when fully developed and either glabrous or sparsely to densely hispidulous, usually on both sides but sometimes only beneath, and an inflorescence 300–1 000(−1 500) mm long with pedicels (5–)15–30 mm long, and flowers with tepals 6–10 mm long, and 8 to 11 ovules per locule. Other specimens seen South Africa: KWAZULU-NATAL. 2831 (Nkandla): Nkandla Forest Reserve, (−CA), new campsite, grassland, 29 Sep 1995, Scott-Shaw 7154 (NU). 2930 (Pietermaritzburg): stony hill near York, (−AD), 3 May 1890, Wood 4304 (BOL, PRE, SAM); 25 km NNE of Howick, Twin Falls, (−AD), 9 Nov 1987, Grove 31 (NU); Noodsberg, (–BD), without date [recd. 1883], Wood 928 (BOL); 2 Sep 1952, Dickson s.n. (NBG) [8 ovules per locule]; Zwartkop, (–CB), locally abundant in damp soil on summit, Oct 1944, Fisher 707 (NU, PRE); Pietermaritzburg, Table Mt., (−DA), 11 Sep 1948, Killick 185 (NU); Krantzkloof, Gillitts Kloof, (−DD), Aug 1921, Haygarth STE88 (NBG) [6 or 7 ovules per locule]; Oct 1921, Haygarth STE87 (NBG, PRE) [8 or 9 ovules per locule]; Kloof, Forest Hills, (–DD), open grassland, Aug 1954, Kays 46 (NU); steep rocky hllside, 27 Aug 1963, Hilliard 1622 (NU). 3029 (Kokstad): Kokstad, (–CB), 11 Mar [sic.] 1964, van der Riet NBG18/64 (NBG) [6 ovules per locule]; Weza, Ingeli, (−DA), 23 Sep 1971, Strey 10525 (PRE). 3030 (Port Shepstone): Kenterton, (−AD), Sep 1913, Thode STE3383 (NBG); Dumisa, (−AD), 15 Oct 1909, Rudatis 765 (NBG) [6 to 8 ovules per locule]; Port Edward/Izingolweni, (–CC), 1979 [without month], 6 Nov 1979 [fl. ex hort.], Louw 25 (NBG); Port Edward, (–CC), 2 Nov 1981, Van Wyk 5140 (PRE). EASTERN CAPE. 3129 (Port St. Johns): Eastern Pondoland, Egossa, (–BC), Aug 1899, Sim 2539 (BOL, NU, PRE); Mkambati River, (–BC), Table Mt., sandstone rocks on summit cliffs, Oct 1937 [fl. ex hort.], Mogg PRE26403 (PRE); Ntsubane Forestry Station near Fraser Falls, (–BC), 22 Aug 1976, Venter and Vorster 40 (PRE); Mkambati Leper Institute, (–BD), 27 Oct 1955, Marais 978 (PRE). Pseudoprospero Speta in Phyton 38: 116 (1998). Type: P. firmifolium (Baker) Speta. Etymology: From the resemblance of its flowers to those of the genus Prospero Salisb. (Hyacintheae). Common name: Albany squill. Deciduous geophyte: Clump-forming. Bulb epigeal, globose, outer scales drying papery and brown. Leaves several, spreading, linear, canaliculate, leathery-striate, ribbed beneath when dry, unmarked, margins minutely denticulate. Inflorescence a solitary, flexuous raceme, usually with one or two small ovate sterile bracts near middle of scape, simple or with short branch from axil of each scape bract, many-flowered; floral bracts small, lanceolate-deltoid, membranous, not auriculate; bracteole present; pedicels arcuate, moderately longer than perianth. Flowers stellate, deciduous if unpollinated, perianth whitish to lilac; tepals biseriate with outer series overlapping inner at base, spreading, connate basally, uni-nerved, withering below fruit. Stamens adnate to base of tepals, slightly shorter than tepals; filaments connate basally, subulate, white; anthers medifxed, reddish to purple with pale yellow pollen. Ovary subglobose, with 2 ovules per locule; style erect,


styliform, subequal to or slightly longer than ovary, slightly 3-grooved. Capsules depressed-turbinate, deeply 3-lobed, leathery, dehisching only along top. Seeds ellipsoid, dark brown, papillate-pustulate. Chromosome number 2n = 18 (Speta, 1998). A monotypic genus restricted to coastal Eastern Cape and KwaZuluNatal, South Africa. It is diagnosed morphologically by the epigeal bulbs with dry-papery outer layers, leathery-striate, canaliculate leaves with minutely denticulate margins, and a flexuous raceme of small, rotate flowers with subulate floral bracts and associated bracteole. The scape usually bears one or two small, ovate, scale-like bracts near the middle, from the axil of which a short branch often issues. The ovules are paired in each locule but just one seed develops per locule in the distinctive, depressed-turbinate capsules. The unpollinated flowers are deciduous, rapidly abscising from the tip of the pedicels. Pseudoprospero is characterised cytologically by a diploid chromosome number 2n = 18, with a karyotype consisting of a graduated series of relatively large chromosomes, both acrocentric and metacentric. This has been interpreted as a dysploid derived from an ancestral base of x = 10 (Goldblatt et al., 2012). The species is toxic and several homoisoflavonones have been isolated from it (Koorbanally et al., 2007). Pseudoprospero was described as a separate genus by Speta (1998). It appears to be only distantly related to the rest of subfamily Scilloideae, as demonstrated by phylogenetic analyses that retrieve it as the first branching lineage in the subfamily, and it was accordingly separated as the tribe Pseudoprospereae by Manning et al. (2004). Pseudoprospero firmifolium (Baker) Speta in Phyton 38: 116 (1998). Scilla firmifolia Baker in Saund., Ref. Bot. 3, app.: 7 (1870); Baker in Bot. J. Linn. Soc., Bot. 13: 237 (1873); Baker in Fl. Cap. 6: 480 (1897); Van der Merwe in Flower. Pl. S. Afr. 24: t. 926 (1944); Jessop in Jl. S. Afr. Bot. 36: 241 (1970). Type: South Africa, Eastern Cape, Grahamstown (3326): ‘Albany Div., New Yearsriver’, (−AD), Nov without year, MacOwan 461 (K, lecto.!, designated by Jessop: 241 (1970); GRA, PRE!, isolecto.). Deciduous geophyte, clump-forming. Bulb ± epigeal, ovoid or subglobose, 200–400 mm diam., outer scales becoming papery and greyish brown from tips otherwise green where exposed, sometimes forming a short, loose collar. Leaves 6–8 in flowering plants, inner leaves progressively smaller and narrower, suberect or arching, lanceolate-attenuate, mostly 100–150 × (2–)5–12(− 16) mm, canaliculate, bright green and softly leathery when fresh, margins narrowly hyaline and minutely denticulate. Inflorescence an elongated, moderately dense raceme, 20–600 mm long, many-flowered, scape flexuous, ± 2 mm diam. at base, bearing one or two ovate, scale-like bracts up to 2 mm long between middle and upper third of scape, simple or with 1 or two short branches, bracts lanceolate to deltoid, membranous at first becoming dry and papery, lowermost up to 10 mm long but usually 4–6 × 0.5–0.8 mm, with minute bracteole 1–2 mm long; pedicels arcuate, 6–10(− 15) mm long. Flowers sub-erect, unscented, perianth flushed pink to lilac with green or brown midrib; tepals spreading or slightly recurved from base, united basally for ± 0.5 mm, narrowly oblong, 3–5 × 1–2 mm, margins deflexed. Stamens adnate to perianth for ± 1 mm, filaments subulate, 3.0–3.5 × 0.5 mm, white; anthers ± 1 mm long at anthesis, reddish to purple with yellow pollen. Ovary turbinate, deeply 3lobed, ± 1.5 × 1.5 mm, either yellowish green with pure white style, or purple with basal half or entire style also purple; style 2.0–2.5 mm long. Capsule suberect, depressed-turbinate, deeply 3lobed, 2.5–3.0 × 3–4 mm, locules mostly 1-seeded. Seeds ellipsoid, 2.5 × 2.0 mm, dark brown, testa closely adhering, epidermis papillate-pustulate. Flowering time: Dec to Mar. Fig. 5. Distribution and ecology: Long considered to be confined to the Eastern Cape, between Alexandria and Umtata (Van der Merwe, 1944; Jessop, 1970), Pseudoprospero firmifolium has since been collected further north between Durban and the Tugela River in KwaZulu-Natal (Fig. 1). Morphologically, the KwaZulu-Natal populations differ in only


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Fig. 6. Schizocarphus nervosus typical form, detached leaf and inflorescence, and flower showing dark blue, verrucose ovary. South Africa, KwaZulu-Natal, between Bergville and Mont-aux-Soucres, Van der Merwe 2612 (PRE). Original painting reproduced in Flowering Plants of South Africa 23: t. 904 (1943). Artist: R. Brown.

minor details from the Eastern Cape plants, notably in their generally broader leaves and most conspicuously in the purple rather than yellowish green ovary. These differences formed the basis for the recognition of two subspecies by Manning et al. (2009). The populations between Alexandria and Peddie are uniform for flower coloration but these details are not known for the plants from Umtata, and this collection is provisionally included in the typical subspecies on account of its narrow leaves. Careful searching along the edge of the coastal plateau between Durban and Umtata may yield further populations that connect the two known areas of occurrence and clarify their taxonomic status. History: The species was described as Scilla firmifolia by Baker (1870) from two collections from the Eastern Cape, one made by the collector J.F. Drège (1794–1881) from the former Transkei and the other by local schoolteacher Peter MacOwan (1830–1909) from near Grahamstown, later selected as the lectotype by Jessop (1970). Subsequent collections were all from that part of the country until its recent discovery in coastal KwaZulu-Natal (Styles, 2010). Conservation notes: The species is currently known from two rather localised regions. The conservation status of the typical subspecies is not clear but most colonies of subsp. natalensis are severely threatened by peri-urban sprawl (Manning et al., 2009). Key to subspecies 1. Leaves 2–10(−15) mm wide at base; ovary yellowish green, style white … subsp. firmifolium 1′. Leaves (4–)6–16 mm wide at base; ovary and style purple … subsp. natalensis. a. subsp. firmifolium Distribution and ecology: Restricted to the Eastern Cape, mainly between Alicedale and Peddie (especially around Grahamstown), with a

single early record from south of Umtata; forming colonies among rocks in open savanna, flowering in mid-summer. Other specimens seen South Africa: EASTERN CAPE. 3128 (Umtata): between Morley and Umtata, (−DD), Feb 1832, Drège 4492 (K). 3326 (Grahamstown): Alexandria, Bushman's River Poort, (–AC), 5 Jan 1956, Archibald 6156 (PRE); Alicedale, (–AC), Dec 1917, Ander BH22507 (BOL); Bathurst, ½ mile [0.8 km] S of Kaffir Drift, (–AC), 21 Jan 1936, Dyer 3374 (NBG, NH, PRE); Kaffir Drift, (–AC), 4 Feb 1970, Bayliss 25/65 (NBG); near Salem, (−AD), 30 Dec 1947, Britten s.n. (PRE); Howieson's Poort, (–BC), 18 Jan 2002, Manning 2701 (NBG); Douglas Heights, (–BC), 22 Mar 1967, Bayliss 3978 (NBG); Lower Albany, Round Hill [Roundhill], (–BD), Dec 1885, Bolus 2869 (BOL, PRE); Hopewell, (–BD), 28 Dec 1944, Acocks 11,058 (PRE); Trappe's Valley, (–BD), 19 Dec 1965, Bayliss 3081 (NBG); Albany District, between Kaffir Drift and Trappe's Valley (–BD), 30 Dec 1964, Leach and Bayliss 12,631 (PRE); Alexandria, (–CB), 16 Feb 1943, Holland s.n. NBG122/32 (NBG); Bathurst (–DB), Jan 1957, Sidley 3067 (PRE). 3327 (Peddie): Peddie Dist., Line Drift, (−AA), Dec 1900, Sim 4060 (PRE, SAM). b. subsp. natalensis J.C.Manning in Manning et al. in Bothalia 39: 232 (2009). Type: South Africa, KwaZulu-Natal, Pietermaritzburg (2930): ‘Nyuswa, below Portion 585 of the farm Assagay Kraal, Botha's Hill’, (−DD), 23 Dec 2007, Styles 3308 (NH, holo.!; K, MO, NBG, NU, PRE, iso.). Distribution and ecology: Known from a few localities along the edge of the coastal plateau between the Umgeni (Mgeni) and Tugela (Thukela) River valleys in KwaZulu-Natal; in shallow soils in open savanna, flowering in mid-summer. Other specimens seen South Africa: KWAZULU-NATAL. 2930 (Pietermaritzburg): close to Inanda Dam, inland of Durban, (–DB), 2005 fl. in cult. 8 Jan 2007, Styles 3307 (NH); Botha's Hill, below Portion 585 of the farm Assagay Kraal, 465 m, (−DD), 14 Dec 2008, Crouch 1184 (NH). 2931 (Stanger): Mabhobhane, near Mapumulo, south bank of Tugela River, (−AA), 8 Jan 2007, Styles 3306 (NH, NBG, NU). Schizocarphus F.van der Merwe in Fl. Pl. S. Afr. 23: t. 904 (1943a); Speta in Phyton 38: 119 (1998). Type: S. nervosus (Burch.) F.van der Merwe. Etymology: From the Greek schizein = split and carpus = fruit. Common name: White squill. Deciduous geophyte: Bulb hypogeal, globose, outer scales firmtextured, papery-fibrous decaying above into moderately firm or stiff fibres, sometimes forming a neck. Leaves several, suberect, often twisted, linear-acicular to lanceolate, firm-textured and striate-fibrous, margins thickened and cartilaginous, glabrous or hispidulous on both surfaces, unmarked. Inflorescence a solitary (rarely up to 4), flexuous raceme, ovoid or cylindrical, several- to many-flowered, scape and rachis hispidulous or sometimes glabrous; bracts small, linear-lanceolate, membranous, not auriculate; bracteole present, thread-like; pedicels short or long, arcuate, hispidulous or glabrous. Flowers rotate, unscented, ±deciduous if unpollinated, perianth white with green apical callosity; tepals biseriate with outer overlapping inner at base, spreading, ±free, uni-nerved, with apical callosity, withering below fruit. Stamens slightly shorter than tepals, filaments free or connate basally, subulate; anthers medifxed, blue with white or yellowish pollen. Ovary shortly stipitate, subglobose and 3-lobed or almost 6-lobed, papillateverrucose, green or blackish, with 5 or 6 ovules per locule; style erect, styliform, subequal to or slightly longer than ovary. Capsules obovoid, shortly stipitate, 3-lobed, leathery. Seeds ellipsoid, black, intricately rugulose [‘brain-like’], epidermis areolate. Chromosome number 2n = 40 (Goldblatt et al., 2012). A monotypic genus widely distributed through southern and southtropical Africa. Schizocarphus is diagnosed by the stiff, striate-fibrous leaves with thickened, cartilaginous margins, papery-fibrous bulb scales that decay into conspicuous bristles, and a characteristically flexuous raceme of rotate flowers with subulate bracts and associated bracteole, hispidulous or sometimes glabrous scape and pedicels, and shortly

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stipitate, verrucose ovary. The free tepals, white flushed green at the tips, and the contrasting green or greenish black ovary are also diagnostic. Schizocarphus, like Merwilla, is characterised cytologically by a diploid chromosome number 2n = 40, and is interpreted by Goldblatt et al. (2012) as a palaeotetraploid from a base x = 10. Although the southern African species of Scilla with fibrous bulbcoverings and white flowers were segregated as the genus Schizocarphus at a relatively early date (Van der Merwe, 1943a), this was not followed by Jessop (1970) in his revision of Scilla in southern Africa but was subsequently adopted by Speta (1998). Molecular phylogenetic analyses confirm that it is sister to the genus Ledebouria (including Drimiopsis Lindl. & Paxton and Resnova F.van der Merwe) in the tribe Massonieae (Wetschnig and Pfosser, 2003; Goldblatt et al., 2012). Morphological support for this association is the occasional production of more than one inflorescence per bulb (mainly in cultivated plants), a flexuous scape, and a stipitate ovary. These features are characteristic for the genus Ledebouria. Schizocarphus nervosus (Burch.) F.van der Merwe in Flower. Pl. S. Afr. 23: t. 904 (1943a). Ornithogalum nervosum Burch., Trav. 1: 537 (1822). Scilla rigidifolia var. nervosa (Burch.) Baker in J. Linn. Soc., Bot. 13: 242 (1873); Baker in Fl. Cap. 6: 481 (1897). Scilla nervosa (Burch.) Jessop in Jl. S. Afr. Bot. 36: 243 (1970); Stedje in Stedge and Thulin in Nord. J. Bot.: 594 (1995); Stedje in FTEA: 10 (1996). Type: South Africa, Northern Cape, Griekwastad (2823): ‘between Griquatown and Wittewater’, (–CC), 14 Feb 1812, Burchell 1968 (K[365586], holo.—image!). Scilla rigidifolia Kunth, Enum. pl. 4: 330 (1843); Baker in J. Linn. Soc., Bot. 13: 242 (1873); Baker in Fl. Cap. 6: 481 (1897). Schizocarphus rigidifolius (Kunth) F.van der Merwe in Fl. Pl. S. Afr. 23: t. 905 (1943b). Type: South Africa, Eastern Cape, Lady Grey (3027): ‘Aliwal North, Witte Bergen [Witteberg]’, (–CA), Jan [without year], Drège 4560b (K [365584], lecto.–image!, effectively designated by Jessop: 243 (1970). Scilla pallidiflora Baker in Ref. Bot. [Saunders] 3: t. 179 (1870); Baker in J. Linn. Soc., Bot. 13: 244 (1973). Type: South Africa, ‘Cap B. Spei’, [cultivated by Wilson Saunders from material collected by Thomas Cooper], illustration in Ref. Bot. [Saunders] 3: t. 179 (1870). Scilla versicolor Baker in Ref. Bot. [Saunders] 5: t. 305 (1873); Baker in J. Linn. Soc., Bot. 13: 243 (1873); Baker in Fl. Cap. 6: 481 (1897). Type: South Africa, ‘Described and figured from a living plant in the collection of Mr. Wilson Saunders at Reigate in 1871, introduced by Mr. Thos. Cooper’, illustration in Ref. Bot. [Saunders] 5: t. 305 (1873). Scilla gerrardii Baker in J. Linn. Soc., Bot. 13: 237 (1873). S. rigidifolia var. gerrardii (Baker) Baker in Fl. Cap. 6: 481 (1879). Schizocarphus gerrardii (Baker) F.van der Merwe in Fl. Pl. S. Afr. 23: t. 906 (1943c). Type: South Africa, KwaZulu-Natal, ‘Natal and Zululand’, 1829 [without month], Gerrard 1829 (K[365583], lecto.—image!, designated by Jessop: 243 (1970). Scilla hispidula Baker in Trans. Linn. Soc. Lond. ser.2 (Bot.) 1(5): 248 (1878). Schizocarphus hispidulus (Baker) Speta in Phyton 38: 120 (1998). Type: Angola, ‘Huilla, prope Lopollo’, Nov., Welwitsch 3830 (BM, holo.). Scilla setifera Baker in Fl. trop. Afr. 7: 549 (1898). Type: Tanzania, ‘Karagwe, Ihangiro’, 12 Nov 1890, Stuhlmann 911 (B[100168457]— image!, holo.; K[257420]—image!, iso.). Scilla pubescens Baker in Bull. Herb. Boiss., sér. 2, 1: 853 (1901). South Africa, KwaZulu-Natal, Pietermaritzburg (2930): ‘Natal, near Howick’, (–AC), Schlechter 6799 (Z, holo.). Scilla eriospermoides Engl. & Gilg in Baum, Kunene-Sambesi-Exped. [Warburg]: 195 (1903). Type: Angola, ‘aus Kubango bei Kavanga’, 16 Nov 1899, Baum 410 (BR[880938], holo.—image!; B[100168426]— image!, K[257440]—image!, M—image!, S—image!, iso.]. Scilla rigidifolia var. acerosa F.van der Merwe in Fl. Pl. S. Afr. 21: t. 821 (1941a). Schizocarphus acerosus (F.van der Merwe) F.van der Merwe in Fl. Pl. S. Afr. 23: t. 904 (1943a). Type: South Africa, Mpumalanga, Witbank (2529): ‘Middelburg, Elandspruit’, (–CD), 1 Oct 1940, Van der Merwe 2236 (PRE[55331], holo.!).


Deciduous geophyte: Bulb hypogeal, ovoid or pyriform, 200–500 (−800) mm diam., outer scales becoming papery- or leathery-fibrous, dark or chestnut brown, decaying above into fine or coarse fibres, sometimes forming a neck. Leaves 3 to 8(12), inner leaves progressively smaller and narrower, stiffly suberect or arcuate, often twisted, linear to lanceolate, rarely almost acicular or needle-like and then canaliculate, 100–300(−400) × (0.5–)5–20(−60) mm, firm-textured and striatefibrous, primary veins thickened, raised into costae when dry, especially beneath, glabrous or more usually variably hispidulous on both surfaces with stiff, patent or deflexed hairs 0.1–0.3 mm long, sometimes only on the costae, bright green when fresh, sometimes flushed purple basally, margins thickened and cartilaginous, glabrous or hispidulous. Inflorescence a dense, ovoid to cylindrical or sometimes almost conical raceme, (50–)100–400(−450) mm long, 25–100(−210 mm) diam., manyflowered, scape suberect or more usually flexed outwards at base, 1.5–5.0 mm diam. at base, striate-hispidulous throughout (including rachis) or only distally or rarely glabrous; bracts lanceolate-attenuate to subulate, membranous, (2–)5–11 × 0.2–0.5 mm, sometimes hispidulous basally, with minute bracteole; pedicels arcuate, (4–) 10–50(−100) mm long, hispidulous or rarely glabrous. Flowers suberect, unscented or sweetly scented, perianth white with green apical callosity or thickening and narrow green midrib beneath; tepals united basally for ±0.5 mm, spreading or slightly deflexed, obovate-cucullate, 3–5 × 1–2 mm, inner slightly narrower than outer. Stamens ± free or connate basally for up to ± 0.5 mm, filaments subulate, 2.5–4.0 × 0.5 mm, white; anthers 1.2–1.5 mm long at anthesis, bluish green.

Fig. 7. Schizocarphus nervosus narrow-leaved ‘acerosa' form, flowering plants with fibrous outer bulb layers and narrow leaves, and detached flower. South Africa, Mpumalanga, Middelburg, Elandspruit, Van der Merwe 2236 (PRE). Original painting reproduced in Flowering Plants of South Africa 21: t. 821 (1941). Artist: M.E. Connell.


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Fig. 8. Distribution of Schizocarphus nervosus (●) in southern Africa and Spetaea lachenaliiflora (○). See Stedje (1996) for the distribution of S. nervosus in tropical Africa.

Ovary subglobose, contracted below and shortly stipitate, deeply 3lobed or almost 6-lobed, ± 1.5 × 1.5 mm, green or dark bluish green with pure white style; style 1.5–3.0 mm long. Capsules suberect or spreading, subglobose or depressed-subglobose, deeply 3-lobed, subsessile or shortly stipitate, smooth or obscurely verrucose, 2.5–4.0 × 3–5 mm. Seeds ellipsoid, ± 2.5 × 1.5 mm, dark brown, testa somewhat turgid and wrinkled. Flowering time: (Sep) Nov to Dec (Feb). Figs. 6 and 7. Distribution and ecology: Widely distributed through southern and east tropical Africa, in South Africa from Grahamstown in the Eastern Cape through KwaZulu-Natal to Mpumalanga and westwards across the interior through Gauteng, Free State, Northwest and adjacent part of the Northern Cape as far as Postmasburg, also Lesotho, Swaziland, southeastern Botswana and northern Namibia (Fig. 8), and further north into Angola, Zimbabwe, Malawi, Tanzania and Zambia (Stedje and Thulin, 1995); in stony or marshy grasslands and open savanna, from near sea level to 2 000 m elevation, flowering throughout the rainy season but mainly in the spring and summer. In view of its wide distribution and broad ecological amplitude it is not surprising that Schizocarphus nervosus varies greatly in overall size, width of the foliage, vestiture of the leaves and scape, and the shape and size of the inflorescence, including the length of the pedicels. Although both the foliage and the inflorescence axis are usually hispidulous, some plants have glabrous leaves combined with a hispidulous inflorescence axis, and others are entirely glabrous, this varying even within a single population [e.g. Johnson 244 (NBG)]. The other morphological features also vary independently of one another, again sometimes within a single population.

The large number of synonyms that apply to this species reflects the great degree of variation in leaf shape and plant stature that is evident across its range. Some of these local forms or ecotypes formed the basis for separate species. The more notable of these include small plants with linear (‘gerrardii’) or almost needle-like (‘acerosa’) leaves (Fig. 7). These occur mainly in open rocky sites in shallow soils and at higher altitudes but in some instances plant size and leaf width appear to be maintained in cultivation [e.g. Hall s.n. (NBG); Anon. NBG73353 (NBG)], suggesting a genetic basis. The type of Schizocarphus nervosus represents robust plants with lanceolate leaves and a broadly cylindrical raceme with long pedicels. Plants matching it (Fig. 6) occur scattered through southern Africa. Plants from tropical and subtropical Africa mostly have narrowly cylindrical racemes with pedicels 5–13 mm long but otherwise vary in the shape and vestiture of their foliage almost as much as those from southern Africa, with linear to lanceolate leaves up to 35 mm wide. Additional collections blurring the distinctions between several of these segregates prompted Baker (1896– 1897, 1898) to reduce them to synonyms or varieties (erroneously under the later basionym Scilla rigidifolia). This process of taxonomic consolidation was completed by Jessop (1970) for the southern African material, and by Stedje and Thulin (1995) for eastern tropical Africa. Although well known to be toxic to stock, Schizocarphus nervosus retains an important place in local pharmacopoeias (Crouch et al., 1999). Schizocarphus nervosus is tolerant of a wide variety of climatic condition and makes an attractive rockery subject or even potplant, as long as it is allowed to become dormant during winter. The dry, plume-like inflorescences prolong its ornamental value. The seeds are easily germinated.

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History: Ornithogalum nervosum was published in 1822 by the naturalist William Burchell (1781–1863) from his own collection made in 1812 in the Northern Cape but this name was passed over by Baker (1870), Baker, 1896–1897) in favour of the later Scilla rigidifolia Kunth (1843) until it was re-instated as the correct name for the taxon in the new genus Schizocarphus by the medical inspector Frederick Van der Merwe (1943a). At this time Van der Merwe (1941a, 1943a, 1943b, 1943c) still recognised at least four taxa in the complex but these were later included in a single broadly circumscribed species [under the new combination Scilla nervosa] by Jessop (1970), and this treatment continues to apply. Speta (1998), who had no intimate knowledge of the species themselves, merely transferred all available names to the genus Schizocarphus, with the observation that the taxonomy of the genus required study. Conservation notes: A widespread and still relatively common species, despite being traded in significant quantities for medical use (Crouch et al., 1999). Other specimens seen from southern Africa. Botswana: 2226 (Serowe): Mosolotsane 12 mi [19 km] W of Kalamare, (–CD), 22 Apr 1970 [sterile], Van Rensburg B4034 (PRE). 2326 (Mahalapye): Mahalapye, (–BB), Dec 1941, Rogers 6741 (PRE); 3000′ [914 m], deep sand, 13 Nov 1963, Bayliss s.n. (NBG). 2425 (Gaborone): Gaberone campus, (–DB), 1 Nov 1975, Mott 812 (PRE); railway crossing 5 km N of Gaberone, (–DB), 8 Oct 1977, Hansen 3215 (PRE). 2426 (Mochudi): Mochudi, (–AC), 13 Dec 1932 [fl. ex hort. Barnard], Barnard BOL22486 (BOL). 2525 (Mafeking): Ranthabane near Pharing, (−AD), 17 Nov 1948, Hillary and Robertson 594 (PRE); 12 mi [19 km] N of Lobatse, (–BA), Nov 1937 [fl. ex. hort.], Van der Merwe 1329 (PRE); Pelotshethla, 45 km W of Lobatse, (–BA), 12 Nov 1983, Plowes 7012 (PRE). Lesotho: 2828 (Bethlehem): Leribe Plateau, (–CC), autumn (fruiting), Dieterlen 231 (NBG, PRE, SAM). 2927 (Maseru): Roma, (–BC), Jan 1962, Ruch 2371 (PRE); Meafeteng Dist., Likhoele Mt., (–CD), 12 Jan 1918, Dieterlen 1312 (PRE). 3028 (Matatiele): near Rama's Gate, (–BB), 26 Jan 1975, Bayliss 1 (PRE). Namibia: 1718 (Nkuringkuru): 6.8 mi [11 km] E of Makamba Camp on road to Katwitwi, (−AD), 10 Dec 1955, De Winter 3870 (PRE); 3 mi [5 km] E of Masari Camp, (–CC), 5 Jan 1956, De Winter 4103 (PRE); Matawa, 4 mi [6.4 km] E of Tondoro Mission, (−DD), 16 Dec 1955, De Winter 3961 (PRE). South Africa: NORTH WEST. 2426 (Mochudi): 2 mi [3 km] NW of Derdepoort, (–CB), 30 Nov 1954, Codd 8901 (PRE). 2525 (Mafeking): Mafeking Dist., Nottingham to Buek Reef, (–DC), 13 Nov 1911, BurttDavy 11,062 (PRE). 2527 (Rustenburg): Rustenburg, (–CA), Jan 1906, Nation 60 (BOL); Rustenburg Nature Reserve, (–CA), 19 Dec 1070, Jacobsen 1639 (PRE). 2624 (Vryburg): Vryburg Dist., Armoess Vlakte, (–DC), 21 Jan 1921, Mogg 8626 (PRE); Leon Taljaard Reserve, (–DC), open scrubland above vlei line, 24 Jan 1977, MacDonald 77/3 (NBG, PRE); Rustenburg Dist., Uitkomst, (−DD), 24 Jan 1971 [fruiting], Coetzee 624 (PRE). 2625 (Delareyville): Strydpoort, (−DD), 16 Sep 1929, Sutton 192 (PRE). 2627 (Potchefstroom): Potchefstroom Dist., Witkop, (−DA), 19 Jan 1946, Louw 1471 (PRE). 2722 (Olifantshoek): Postmasburg Dist., 38 mi [61 km] NNW of Olifantshoek, (–AC), 18 Jan 1960, Leistner 1607 (PRE). 2723 (Kuruman): Kuruman Div., Esperanza, (−AD), from the garden, Apr 1940 [fruiting], Esterhuysen 2851 (BOL); Mount Carmel Farm, (–DC), 15 Feb 1981 [fruiting], Gubb s.n. (PRE). 2724 (Taung): Barkly West Dist., Kuil, (–CD), Feb 1886 [fruiting], Marloth 1015 (PRE). 2726 (Odendaalsrus): Greylingsrust, Maquassie, (–AC), 10 Jan 1969, Morris 1125 (PRE). LIMPOPO. 2328 (Baltimore): Waterberg Dist., Valley Villa Nora, (–CA), without date, Van der Merwe 2332 (PRE). 2329 (Pietersburg): plateau near top of Blaauwberg [Blouberg], (−AA), 3 Dec 1958, Story 6511 (PRE); Soutpansberg, Hanglip, Louis Trichardt, (–BB), 26 Jan 1931 [fruiting], Bremerkamp and Schweikerdt 441 (PRE); Marabas Hoek, (–CD), Dec 1924, Van Dam s.n. (PRE); Haenertsburg, (−DD), 29 Oct 1928, Cunliffe s.n. (PRE). 2330 (Tzaneen): Wolkberg, (−AA), 20 Nov 1945, Gerstner 5606 (PRE);


Wolkberg, (−AA), Feb 1919 [fruiting], Junod 4134 (PRE); Lekgalameetse Nature Reserve, near Cypress Point, (–AB), 22 Oct 1922, Stalmans 239 (PRE); 8 Jan 1986, Stalmans 942 (PRE); Letaba Dist., Duiwelskloof, (–CA), 6 Nov 1959, Scheepers 758 (PRE); slopes near Haenertsburg, (−DD), 21 Oct 1938, Hafström and Acocks 234 (PRE); 2427 (Thabazimbi): 50 km SE of Ellisras on tarred road to Vaalwater, (–BA), 19 Nov 1981, Reid 455 (PRE); Thabazimbi, Kransberg, (–BC), 1 Mar 1980 [fruiting], Westfall 941 (PRE); Waterberg Dist., near Rooiberg, (–DC), 21 Nov 1938, Van der Merwe 1848 (PRE). 2428 (Nylstroom): Zandspruit near Sandrivierspoort, (–AC), Oct 1939, Forssman 300 (PRE); Waterberg Dist., basal slopes of Mtn Nooitgedacht near Naboomspruit, (−AD), 14 Nov 1934, Galpin BH22482 (BOL); Sterkrivierdam Nature Reserve, (–BB), 11 Jan 1973, Jacobsen 2548 (PRE); Nylstroom, (–CB), 6 Nov 1938, Strydom 1804 (PRE); Warmbaths, (–CD), Oct 1908, Leendertz 1535 (PRE); Waterberg Dist., Nooitgedacht, (−DA), 14 Nov 1934, Galpin 13316 (PRE); Naboomspruit, (−DA), 29 Nov 1919, Galpin M365 (PRE). 2429 (Zebediela): Percy Fife Nature Reserve, (−AA), 28 Jan 1968 [fruiting], Huntley 1090 (PRE). GAUTENG. 2527 (Rustenburg): Krugersdorp, Jack Scott Private Nature Reserve, (–DC), 2 Feb 1961 [fruiting], Wells 2330 (PRE). 2528 (Pretoria): Pretoria, (–BA), 12 Nov 1947, Smith s.n. (PRE); Van Riebeeck Nature Reserve, (–BC), 10 Sep 1967, Kok 191 (PRE); Wonderboompoort, (–CA), 1 Oct 1904, Leendertz 320 (BOL, PRE); Fountains Valley, (–CA), 20 Sep 1929, Repton 297 (PRE); Farm Strydfontein about 2 mi [3 km] from Hornsnek near dam, (–CA), 4 Nov 1937 [fl. ex hort.], Dyer and Erens 111 (PRE); 12 km W of Pretoria West, Elandsfontein Farm, (–CA), 18 Feb 2015 [fruiting], Bester 12482 (PRE); Kopjes S of Pretoria, (–CC), Sep 1929, Obermeyer 22 (PRE). 2627 (Potchefstroom): Witpoortjie, (–BB), Feb 1926 [fruiting], Gilmore s.n. (PRE); Roodepoort, Transvaal Botanic Garden, open grassland, (–BB), 1620 m, 27 Jan 1983, Behr 348 (NBG). 2628 (Johannesburg): Johannesburg, (−AA), Jan 1915, Rogers 14110 (BOL); Johannesburg, ridge above Jesppestown, (−AA), 11 Jan 1897, Galpin 1459 (BOL, PRE); Kensington Ridge, (−AA), Jan 1933, Heydorn 98 (NBG); Witwatersrand, Frankenwald, Jokeskei [Jukskei] River, (−AA), 18 Feb 1951 [fruiting][, HBG 25082 (BOL); Pretoria, Wilge River, (–BB), 20 Sep 1936, Repton 665 (BOL); Suikerbosrand, (–CA), 28 Dec 1971, Bredenkamp 642 (PRE). MPUMALANGA. 2430 (Pilgrim's Rest): Serala Peak, Farm Wolkberg, (−AA), 10 Sep 1985, Venter 10901 (PRE); Mt. Anderson, (–BA), Dec 1932, Smuts and Gillett 2442 (NBG, PRE); Pilgrim's Rest, Morgenzon, (–BA), 27 Nov 1978, Van der Zeyde 63/78 (PRE); Sabie, (–BB), 8 Dec 10,981 [fruiting], Deall 1296 (PRE); Nelspruit-Sabie main road near Sabie, (–BB), 29 Sep 1982, Onderstall 838 (PRE); 18 mi [29 km] from Nelspruit on Johannesburg road, (–BC), 11 Aug 1970, Buitendag 627 (NBG, PRE); Rosehaugh, (–BD), Nov 1915 [fl. ex hort. Kirstenbosch], Sim NBG3767/14 (BOL); Lowveld Botanic Garden, (–BD), 18 Oct 1971, Buitendag 880 (NBG); 10 km from Bourke's Luck Potholes on Graskop road, (–CD), 24 Nov 1999, Zietsman 3972 (PRE); Ohrigstad Nature Reserve, (–DC), 6 Dec 1970, Jacobsen 1546 (PRE); Graskop, (−DD), Oct 1917, Thorncroft s.n. (PRE); 1 Oct 1943, Holland s.n. (PRE); Feëland near Graskop, (−DD), 8 Sep 1978, Kluge 1383 (PRE). 2431 (Acornhoek): Lothian Forest Reserve, (–CC), 16 Oct 1937, Joubert 9029 (PRE). 2529 (Witbank): Loskop Dam, (−AD), 11 Jan 1968 [fruiting], Theron 1660 (PRE); Loskop Nature Reserve, (−AD), 13 Dec 2001, Potgieter 183 (PRE); near Wonderfontein, (–CA), 1 Jan 1938 [fruiting], Van der Merwe 1574 (PRE); Witbank, Clydesdale Pan, (–CC), 25 Oct 2001, De Castro 8979 (PRE); near Arnot, (−DD), 30 Jan 1929 [fruiting], Hutchinson 2719 (BOL, PRE). 2530 (Lydenburg): Steenkasmpsberg Pass, (−AA), 28 Nov 1999, Zietsman 4016 (PRE); Steenkampsberg, (–AC), 10 Mar 2000 [fruiting], Van Wyk 64 (PRE); Dullstroom, (–AC), 26 Dec 1985, Cameron 411 (PRE); 9 mi [14.4 km] N of Belfast, (–CA), 17 Dec 1956, Prosser 2057 (PRE); Machadadorp, (–CB), grassy hills, 8 Dec 1932, Galpin s.n. (BOL, PRE); Waterval Boven, Farm Beerzynbosch, 6 km off the Machadadorp main road, (–CD), without date [fl. ex hort.], Steel 216 (PRE); 7 km from Kaapsche Hoop towards Ngodwana, (−DA), 14 Dec 1975, Van Jaarsveld 1016 (PRE); Berlin, Godwan River Station, (−DA), Jan 1923 [fruiting],


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Davison 72 (PRE); Kaapsche Hoop, (–DB), Feb 1938 [fruiting], Van der Merwe 1646 (PRE); Tafelkop, Nelsberg, (−DD), Mar 1938 [fruiting], Van der Merwe 1678 (PRE). 2531 (Komatipoort): White River, (–AC), Oct 1919, Rogers 21,289 (PRE); Barberton, (–CC), Jan 1907, Thorcroft 2970 (PRE); slopes of Saddleback Mt., (–CC), 17 Oct 1945, Codd s.n. (PRE); Songimvelo Game Reserve, Makhonjwa Hills, (–CC), 9 Dec 1992, Meyer 71 (PRE). 2629 (Bethal): 10 km from Ogies on road to Standerton, (−AA), 30 Sep 1977, Venter 2115 (PRE); Bethal, (− AD), 14 Dec 1910, Leendertz 9570 (PRE); Ermelo, (–DB), without date, Henrici 1264 (PRE); Ermelo Dist., Spitskop, (–DB), Jan 1916, Scheepers s.n. (PRE). 2630 (Carolina): Carolina, (−AA), 12 Oct 1938, Little s.n. (NBG); 23 Nov 1945, Little s.n. (NBG); between Warburton and Lochiel on N17 between Carolina and Swaziland, (–BA), 21 Jan 2002 [fruiting], Koekemoer 2159 (PRE); road to Oshoek in Kangwane, (–BB), 11 Jan 1984 [fruiting], Welman 454 (PRE); Athole Experimental Farm, (–CB), 22 Jan 1976, Balsinhas 2937 (PRE). 2730 (Vryheid): Wakkerstroom, Farm Oshoek, (−AD), 4 Dec 1960, Devenish 286 (PRE); slopes of Hlangapies, (–BB), Dec 1936, Van der Merwe 1102 (PRE). FREE STATE. 2627 (Potchefstroom): Heilbron, (–DB), 28 Nov 1959, Acocks 20976 (PRE); Wonderwater, (−DD), 25 Nov 1995, Kroon 11760 (PRE). 2726 (Odendaalsrus): Bothaville, (–BC), 30 Jan 1933, Goossens 1166 (PRE); Farm Sandfontein, 15 mi [24 km] W of Bothasville, (–BC), Mar 1933, Schweickerdt 1090 (PRE). 2727 (Kroonstad): Kroonstad, (–CA), Nov 1928, Pont 372 (PRE); Kroonstad, Farm Blanquilla, (–CA), 28 Mar 1968 [fruiting], Scheepers 1697 (PRE). 2825 (Boshof): Boshof Dist., Smitskraal, (−AA), 10 Mar 1912, Burtt-Davy 12891 (PRE). 2826 (Bransdfort): Glen Landboukollege, (–CD), 22 Feb 1985, Zietsman and Zietsman 236 (PRE). 2828 (Bethlehem): Qua Qua National Park, Avondrust, (–BB), 22 Nov 1994, Zietsman 2639 (PRE); Golden Gate National Park, Generaalskop, (−DA), 21 Jan 1965, Rob erts 3118 (PRE); Witsieshoek, hill above Bester's Vlei, (–DB), Dec 1893, Flanagan 1837 (PRE, SAM); Witsieshoek, (–DB), Jan 1895, Thode STE6338 (NBG); Broom [Broome] Hill, (–DB), 21 Nov 1928, Galpin 10970 (PRE). 2829 (Harrismith): Platberg foothills, (–AC), 30 Jan 1971, Hahn s.n. (NBG); Drakensberg Botanic Garden, (–AC), grassveld, 1706 m, 8 Feb 1978, Jacobsz 2211 (NBG, PRE); Swinburne, Rensburgskop, (–AC), 4 Nov 1962, Jacobsz 179 (PRE); Kerkenberg, (–CA), 18 Nov 1981, Jacobsz 1268 (PRE); Sterkfontein Dam, (–CA), 28 Nov 1980, Blom 71 (PRE); Oliviershoek Pass, (–CA), Dec 1908, Thode STE3371 (NBG). 2924 (Hopetown): Fauresmith, on road to Palmietfontein, (–DB), Jan 1928, Smith 5332A (PRE). 2925 (Jagersfontein): Petrusberg, on Steunmekaar road, (–BC), 19 Jan 2000 [fruiting], Zietsman 4067 (PRE). 2926 (Bloemfontein): near Bloemfontein, (−AA), Feb 1904 [fruiting], Bolus 10824 (BOL, PRE); 26 Feb 1966, Hanekom 594 (PRE). KWAZULU-NATAL. 2729 (Volksrust): N-facing slopes of Majuba Mt. below summit, (–BD), 7 Jan 1995, Archer and Archer 2067 (PRE); 15 km SW of Volksrust, (–BD), 30 Nov 2015, Bester 12,941 (PRE); Newcastle Dist., Chelmsford Dam Reserve, (−DD), 17 Nov 1997, Ngwenya 1595 (PRE); Newcastle, (−DD), 1 Dec 1943, Van der Merwe 2775 (PRE). 2730 (Vryheid): 5.8 km NW of Vryheid on R33 to Paulpietersburg, (−DA), 21 Nov 2010, Bester 10548 (PRE). 2731 (Louwsburg): 18 km from Kongolwane on road to Louwsburg, (–CA), 18 Oct 1982, Germishuizen 2216 (PRE); Louwsburg Dist., Gluckstadt Mission, (–CC), 9 Oct 1969, Strey 9202 (PRE); Ngome, (–CD), 15 Dec 1969, Strey 9396 (PRE); 6 Dec 1985 [fruiting], Van Wyk 7075 (PRE); Ngome, near sawmill, (–CD), 7 Dec 1975, Hilliard and Burtt 8429 (PRE). 2732 (Ubombo): Vasi swamp, (–BB), Oct 1972, Stephen et al 1174 (PRE). 2828 (Bethlehem): near Mont aux Sources, (–DB), Nov 1942, Van der Merwe 2612 (PRE); Tugela Valley, Drakenberg National Park, (–DB), 29 Oct 1938, Hafström and Acocks 236 (PRE). 2829 (Harrismith): Van Reenen, (−AD), 1 Jan 1913, Wood 12173 (PRE); Cathedral Peak, (–CC), 10 Nov 1951, Killick 1567 (PRE); 14 Nov 1984, Hardy 27 (PRE); 10 Sep 2014, Gordijn 141 (NU). 2830 (Dundee): Helpmekaar, (−AD), 10 Jan 1952, Codd 6781 (PRE); 4 km from turnoff onto R568 from Muden-Colenso Road, (–CD), 29 Nov 1988, Kok and Pienaar 1230 (PRE); Weenen, (–CC), 11 Apr 1891, Wood 4393 (BOL, SAM); Muden, (–CD), Nov 1936, Wylie s.n. (PRE); Kranskop, (−DD),

13 Oct 1942, Van der Merwe 2595 (PRE). 2831 (Nkandla): Mahlabatini, (–AB), 15 Nov 1940, Gerstner 4217 (PRE); Mtunzini Dist., Hameworth, (–DC), 18 Nov 1919, Mogg 5915 (PRE). 2832 (Mtubatuba): Hlabisa Dist., Hluhluwe Game Reserve, (−AA), 28 Nov 1959, Ward 3344 (PRE); 7 Nov 1962, Ward 4507 (PRE); Dukuduku, (–AC), 22 Nov 1964, Strey 5550 (PRE); Lake Nhlabane area, (–CB), 23 Jan 1992, Ward and Begg 11746 (PRE). 2929 (Underburg): Cathkin Park, (–AB), Jan 1934, Edwards BOL22519 (BOL); Loteni Nature Reserve, (−AD), 13 Dec 1978, Jacobsz 3945 (PRE); Giant's Castle, (−AD), 4 Nov 1897, Guthrie 4940 (BOL); Tabamhlope, (–BA), 9 Dec 1937, West 477 (PRE); Underberg, Farm Ottery, (–CB), 26 Jan 1998 [fruiting], Singh 342 (PRE); 42 km from Underberg to Franklin on Bushmansnek road, (–CD), 17 Nov 1980, Arnold 1294 (PRE); Impendhle [Mpendle] Dist., Le Suers's farm, (–DB), 6 Jan 1976, Wright 2394 (NU); Himeville, (–DC), without date, Bews 35 (PRE). 2930 (Pietermaritzburg): Mooi River, (−AA), Jan 1932 [fruiting], Mrs A.H. Broom BOL22485 (BOL); Dargle Road, (–AC), 19 Jan 1920, Mogg 6854 (PRE); N of Howick, (–AC), Dec 1939, Van der Merwe 2083 (PRE); Greytown, (–BA), Nov 1931, Wylie s.n. (PRE); 7 mi [11 km] N of Pietermaritzburg, (–CB), 28 Dec 1947, Barker 5196 (NBG); 2 Jan 1948, Barker 5204 (NBG); Nov [without year], Thomas 4 (NBG); Maritzburg [Pietermaritzburg], (–CB), Nov 1932 [fl. ex hort. Rockery Kirstenbosch], Carnegie NBG251/31 (BOL); Pietermaritzburg, Old Howick Rd, (–CB), 22 Dec 1988, Kennedy 104 (NU); near Pietermaritzburg on Richmond road, (–CB), 11 Jan 1947, Barker 4368 (NBG); Cedara, (–CB), 7 Dec 1921, Phillips 3462 (PRE); Richmond, (–CD), 29 Dec 1953, Compton 23858 (NBG); between Kokstad and Richmond, (–CD), 5 Dec 1928, Hutchinson 1833 (BOL, PRE); Claremont, (−DA), 16 Nov 1909, Wood 11546 (PRE); Krantzkloof, (−DD), Dec 1911, Haygarth STE86 (NBG). 2931 (Stanger): Inyoni Leper Institute road, (−AA), 30 Dec 1951, Johnson 344 (NBG); 6 km E of KwaSizabantu, Buffelhoek, (−AA), grassveld near streambank, 17 Dec 1980, Van Jaarsveld & Jacobs 5860 (NBG). 3029 (Kokstad): Ngeli Mt., (−DA), 19 Nov 1986y, Jordaan 833 (PRE); Ngele Nature reserve, KwaShwili area, (−DA), 22 Nov 1994, Ngwenya 1341 (PRE); Harding, Rooi Vaal, (–DB), 4 Jan 1957, Taylor 5284 (NBG); 8 Jan 1957, Barker 5373 (NBG). 3030 (Port Shepstone): St Bernhards [St Bernhard Mission], (–AB), 22 Dec 1952, Barker 7962 (NBG); Alexandra County, Umgage [Umgababa] flats, (–BB), 29 Nov 1908, Rudatis 505 (NBG); Winkle Spruit [Winkelspruit], (–BB), 23 Nov 1911, Rudatis 1513 (NBG); Dumisa, (–BB), 13 Dec 1912, Rudatis 1802 (NBG); Isipingo, (–BB), 23 Oct 1913, Wood 12,391 (PRE); Oribi Gorge Nature Reserve, (–CA), Jan 1977, Henderson et al 115 (PRE); Oribi Gorge, The Rocks, (–CA), Jan 1978, Mantell and Vassilatos 165 (PRE); Izotsha, (–CB), 20 Dec 1966, Strey 7181 (PRE); Uvongo, (–CB), littoral, 15 Dec 1936, Mogg 13339 (PRE); Port Shepstone Dist., Shelley Bay, (–CB), Dec 1930, Mogg 12182 (PRE); Umtamvuna River, (–CC), 19 Dec 1973, Nicholson 1359 (PRE); St. Michael-on-Sea, (–CD), 25 Dec 1966, Strey 7073 (PRE. 3130 (Port Edward): Port Edward, (−AA), 10 Oct 1957, Taylor 5384 (NBG); Graigadoor Farm, overlooking Umtamvuna River, (−AA), 30 Dec 1976, Nicholson 1698 (PRE); 3 km from Umtamvuna River bridge, (−AA), 10 Dec 1985, Pienaar 795 (PRE). NORTHERN CAPE. 2823 (Griekwastad): Danielskuil, (–BA), Mar 1914 [fruiting], Wilman 9109 (SAM). 2824 (Kimblerley): Barkly West Dist., Koopmansfontein, (−AA), Feb 1937, Acocks 1802 (PRE); Kimberley, (–DB), 18 Mar 1929 [fl. ex hort.], Marloth 12,752 (PRE); Kimberley, Droogfontein, (–DB), Jan 1937 [fruiting], Acocks 1650 (PRE); Riverton, (–DB), Jan 1919, Wilman BOL15730 (BOL). 2924 (Hopetown): Orange River Station, (–CA), without date, Cooper 875 (BOL). EASTERN CAPE. 3028 (Matatiele): Mt. Fletcher on road to Rhodes, (–CB), 7 Dec 1983, Van Wyk 6693 (PRE). 3029 (Kokstad): mountains near Kokstad, (–CB), Dec 1882, Tyson 1480 (BOL, PRE); Kokstad Convent School, (–CB), 13 Jan 1939, Sr. Mildred 5 (NBG). 3126 (Queenstown): mountain sides, Queenstown, (–DB), Jan 1894, Galpin 1765 (PRE). 3127 (Lady Frere): Encgobo, (–DB), 1 Dec 1938, Van der Merwe s.n. (PRE). 3128 (Umtata): Baziya, (–CB), Nov [without year], Baur 378 (BOL, SAM); 18 Dec 1939, Van der Merwe 2120 (PRE); Umtata, (–DB), 13 Apr 1948, Wurts 3045 (PRE); near Emagusheni, (–DC), 16 Dec 1939, Van der

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Merwe 2106 (PRE). 3129 (Port St Johns): 12 mi [19 km] ENE of Lusikisiki, (–BC), 13 Jan 1947, Acocks 13242 (PRE); Lusikisiki, Fraser's Gorge, (–BC), 14 Dec 1964, Bayliss 2554 (NBG); Mkambati Reserve, (–BD), 26 Jan 1986, Perry 3428 (NBG). 3130 (Port Edward): Mkambati Nature Reserve, 2 km S of Mkambati Waterfall, (–AC), 11 Dec 1986, Nicholas and Smook 2387 (PRE); Mkambati Nature Reserve, (–AC), 16 Jan 1997, Makwarela 244 (PRE). 3226 (Port Beaufort): Katberg, (–BC), Mar [without year], Zeyher s.n. (NBG, SAM); Amatole Mts, near Gaika's Kop, (–DB), 18 Jan 1983, Furness and Phillipson 139 (PRE); Fort Beaufort Dist., Hogsback, Tor Doone, (–DB), 16 Jan 1943, Giffen 1266 (PRE). 3227 (Stutterheim): near Fort Cunnyngham, (−AD), Jan 1896, Bolus 10344 (BOL); 1987, Sim 20,466 (PRE); Jan 1924, Schönland 92 (PRE); Mt. Coke, (–CB), Jan 1893, Sim 1698 (BOL); Pirie, (–CB), Jan 1894, Sim 1699 (PRE); Mt. Kemp, (–CB), 13 Jan 1947, Leighton 2733 (PRE); King Williamstown, (–CD), 13 Jan 1947, Leighton 2733 (BOL); Kei Road, Stockton Farm, (−DA), 4 Dec 1957 [fruiting], Comins 1781 (PRE); grassy hills W of Komga, (–DB), Nov 1889, Flanagan 463 (NBG, PRE); East London, (−DD), 1908, Pearson 1525 (NBG). 3228 (Butterworth): Kentani, (–CB), without date, Pegler 362 (PRE); Kei Mouth, (–CB), 8 Jan 1895, Schlechter 2440 (PRE). 3326 Grahamstown: Albany, Manley Flats, (–BC), 21 Jan 1947, Compton 19075 (NBG); grassy flats halfway to Bushman's River Mouth, (−DA), 12 May 1931, Galpin s.n. (BOL, PRE); Kariega, (−DA), Jan 1904, White 2440 (PRE). Swaziland: 2531 (Komatipoort): Piggs Peak Dist., Emlembe, (–CC), 1 Feb 1959 [fruiting], Compton 28486 (PRE). 2631 (Mbabane): Ngwenya Hills, (−AA), 20 Dec 1977, Kemp 1192 (PRE); Malolotja Nature Reserve, near Lagwaja, (−AA), 14 Nov 1998, Braun 476 (PRE); Forbes Reef, near stream, (−AA), 20 Dec 1956, McColl s.n. (PRE); 5 mi [8 km] S of Stegi, (–BD), 25 Nov 1958, Compton 28379 (NBG, PRE). Locality unknown: Hort. rockery Kirstenbosch, 10 Nov 1954, Hall s.n. (NBG); Hort. nursery Kirstenbosch, 1 Nov 1963, Anon. NBG73353 (NBG). Spetaea Wetschnig & Pfosser in taxon: 87 (2003). Type: S. lachenaliiflora Wetschnig & Pfosser. Etymology: Commemorating Austrian taxonomist Franz Speta (b. 1941), who published extensively on the family. Common name: Cape squill. Deciduous geophyte: Bulb globose, tunicated, outer tunics drying papery and brown. Leaves several, suberect, linear-lanceolate, involute, unmarked, succulent. Inflorescence a solitary, erect, cylindrical raceme, many-flowered; bracts small, membranous, not auriculate; bracteoles lacking; pedicels moderately long, spreading. Flowers spreading, campanulate, persisting if not pollinated, perianth blue to violet; tepals biseriate with outer overlapping inner at base, suberect below spreading distally, connate basally, uni-nerved, withering below fruit. Stamens suberect, slightly longer than tepals and shortly exserted, adnate to base of tepals; filaments connate basally, filiform; anthers medifixed, blue. Ovary sessile, ovoid, with 2 to 4 ovules per locule; style slightly flexed to one side, filiform, ± twice as long as ovary. Capsules obovoid, deeply 3-lobed, leathery. Seeds ellipsoid, rugulose, black, epidermis scalariform-colliculate. Chromosome number 2n = 20 (Wetschnig and Pfosser, 2003). A monotypic genus restricted to the extreme southwestern part of Western Cape and confined to a small area of the Cape Fold Mountains near Wellington. Spetaea is diagnosed by its tunicated bulb, narrow, involute, succulentleaves, andebracteolate raceme of campanulate, blueor violet flowers with stamens exserted beyond the tepals and a slightly flexed style more than twice aslongasthe ovary. Unpollinated flowers persist on the pedicels for some time, as they do in the allied genera Massonia Thunb. ex Houtt., Lachenalia J.Jacq. ex Murray and Daubenya Lindl. The genus is characterised cytologically by a diploid chromosome number 2n = 20 and a remarkable bimodal karyotype consisting of one pair of very large chromosomes, one medium-sized pair, and the remainder very small chromosomes (Goldblatt et al., 2012). Phylogenetic analyses retrieve Spetaea as sister to the morphologically highly dissimilar genus Daubenya Lindl., which is characterised by a pair of broad, prostrate leaves and a condensed, corymbose


inflorescence borne at ground level, with relatively large bracts and ± tubular flowers that are variously coloured but never blue (Manning et al., 2004). Recorded diploid chromosome numbers for Daubenya are 2n = 32 and 34 (Goldblatt et al., 2012). Spetaea lachenaliiflora Wetschnig & Pfosser in Taxon: 87 (2003); G.J.Lewis in Fl. Pl. Afr. 26: t. 1006 (1947) [as ‘Scilla plumbea’]. Type: South Africa, Western Cape, Worcester (3319): ‘Du Tois kloof [Dutoitskloof], III H', (–CA), Jan [without year], Drège 1997 (B, holo.; W, iso.). Scilla plumbea sensu auct. [Kunth, 1843, G.J. Lewis, 1947, Jessop, 1970 et seq.], non Lindl. (1830). Plants solitary: Bulb subglobose, 20–35 mm diam., outer layers drypapery, dark brown. Leaves synanthous but often drying above at flowering, 2 to 4(6), suberect, linear to narrowly lanceolate, involute, 150–250(−400) × 8–15 mm but rarely up to 30 mm wide at base, inner leaves often shorter and narrower than outer, acute to attenuate, glabrous, glossy green, margins minutely hyaline, smooth. Inflorescence a dense, erect raceme 300–500 mm long with rachis 90–200 mm long, 30–50 mm diam., many-flowered; scape glabrous, 4–7 mm diam. at base, glaucous flushed violet; bracts ovate, 3–4 mm long, concave, apiculate, ebracteolate; pedicels stiffly horizontally spreading, 8–15 mm long, violet. Flowers spreading or half-nodding, deeply campanulate, perianth blue to violet, unscented; tepals connate basally for 0.5–1.0 mm in a shallow cup, suberect below and arcuate-spreading or recurved in distal half, lobes lanceolate, 8–12 × 2–3 mm. Stamens connate basally for ±1 mm; filaments filiform, slightly longer than perianth, 10–12 mm long, flushed blue to violet; anthers ellipsoid, ±1.5 mm long, violet with whitish pollen. Ovary obovoid, 2.5–3.0 mm long, yellowish green; style flexed weakly downwards or to one side, filiform,

Fig. 9. Spetaea lachenaliiflora, flowering plant, detached, campanulate flower with floral details showing long stamens exserted slightly beyond tepals and flexed style, and detached capsule. South Africa, Western Cape, Baviaanskloof off Bainskloof, Linley s.n. (BOL, PRE, SAM). Original painting reproduced in Flowering Plants of Africa 26: t. 1006 (1947). Artist: G.J. Lewis.


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7–10 mm long, flushed violet. Capsules spreading, obovoid, 3-lobed, 5–6 mm diam. Seeds ellipsoid, ± 2.5 × 2.0 mm, rugulose, black. Flowering time: (late Nov) Dec to Jan, only after fire. Fig. 9. Distribution and ecology: A highly localised endemic of the Limietberge and adjacent Slanghoekberge east of Wellington in Western Cape, South Africa (Fig. 8); restricted to rocky sandstone seepages in humus-rich soils, mainly on S-facing slopes, flowering in midsummer only after a fire the previous summer. The flowers of Spetaea lachenaliiflora are visited by bees of the genus Amegilla (Apidae: Anthophorinae) and also by smaller pollen-collecting bees of the family Halictidae. Although the flowers are exceptionally vividly coloured and the tall spikes highly attractive, the species is scarcely known in cultivation, and its pyrophytic habit and sandstone habitat suggest that it may not be easy to grow. History: This local, montane species was first collected by J.F. Drège (1794–1881) in Dutoitskloof in the mid-1800s. His collection was incorrectly associated with the type illustration of Scilla plumbea (now Merwilla plumbea) by Kunth (1843) and later authors, including Lewis (1947) and Jessop (1970), although Baker (1896–1897) appears to have been unaware of it. This mistaken association with S. plumbea was noted by G.S. Linley in 1945 (Linley BOL56004) and later confirmed by Austrian botanist Franz Septa (Wetschnig and Pfosser, 2003). Molecular analyses by the latter authors showed that the species was not directly allied with Merwilla at all, and it was accordingly described as a new species and genus. Conservation notes: A range-restricted species known only from three populations, two in Bainskloof in the Limietberge and one in Dutoitskloof in the adjacent Slanghoekberge, none under any known threats. Currently classified as Rare (Raimondo et al., 2009). Other specimens seen South Africa: WESTERN CAPE. 3319 (Worcester): Bainskloof, (–CA), Dec 1946, Compton s.n. (NBG); 12 Dec 1959, Van Breda 907 (NBG); 3 Jan 1952, Compton s.n. (NBG); 15 Dec 1968 [fl. ex hort Kirstenbosch 19 Dec 1969], Loubser s.n. (NBG); Bainskloof, ± 1 km NE of hotel, (–CA), 16 Dec 1952, Esterhuysen 25155 (BOL); 2 Dec 1986, Perry 3570 (NBG); Bainskloof, first roadside waterfall on Worcester side, (–CA), 12 Jan 1971, Schelpe s.n. (BOL); Bainskloof, Baviaanskloof, Witte River, (–CA), fl. ex hort. Kirstenbosch 25 Nov 1945, Lamb NBG817/30 (BOL, NBG); Bainskloof, Baviaanskloof, (–CA), Jan 1945, Linley s.n. (BOL, NBG, SAM); 20 Oct 1946, Leighton 2860 (BOL); 15 Dec 1968, Barker s.n. (NBG); 10 Dec 1979, Cameron and Mendings 18 (BOL); eastern end of Du Toitskloof, S base of Slanghoek Peak, De la Bats Plaats, (–CA), 20 Dec 2008, Helme and Turner 5963 (NBG).

Acknowledgments I thank the Curators of the various herbaria for access to their collections; Sandra Turck and Daleen Maree for providing the scans of the colour artwork from the archives of the South African National Biodiversity Institute; Michelle Smith for preparing the distribution maps; and Anthony Magee for processing the digital images for publication. References Ali, S., Yu, Y., Pfosser, M., Wetschnig, M., 2012. Inferences of biogeographical histories within subfamily Hyacinthoideae using S-DIVA and Bayesian binary MCMC analysis implemented in RASP (reconstruct ancestral state in phylogenies). Ann. Bot. 109, 95–107. Angiosperm Phylogeny Group, 2003. An update of the angiosperm phylogeny group classification for the orders and families of flowering plants: APGII. Bot. J. Linn. Soc. 141, 399–436. Angiosperm Phylogeny Group, 2009. An update of the angiosperm phylogeny group classification for the orders and families of flowering plants: APGIII. Bot. J. Linn. Soc. 161, 105–121. Baker, J.G., 1870. Monograph of Scilla: § Ledebouria and Drimiopsis. Refug. Bot. [Saunders] 3, 7–13 app.. Baker, J.G., 1873. Revision of the genera and species of Scilleae and Chlorogaleae. J. Linn. Soc. Bot. 13, 209–292.

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