Systematics of the genus veltheimia (Hyacinthaceae: Scilloideae)

Systematics of the genus veltheimia (Hyacinthaceae: Scilloideae)

South African Journal of Botany 127 (2019) 271277 Contents lists available at ScienceDirect South African Journal of Botany journal homepage: www.e...

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South African Journal of Botany 127 (2019) 271277

Contents lists available at ScienceDirect

South African Journal of Botany journal homepage:

Systematics of the genus veltheimia (Hyacinthaceae: Scilloideae) J.C. Manninga,b* a

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 b



Article History: Received 25 June 2019 Revised 10 September 2019 Accepted 13 September 2019 Available online xxx Edited by AR Magee Keywords: Massonieae Nomenclature Southern Africa Taxonomy Typification


The southern African genus Veltheimia (Hyacinthaceae: Scilloideae: Massonieae) comprises the paraptric species pair V. capensis and V. bracteata. Both species are reviewed with full nomenclature, descriptions, ecological notes, and colour illustrations. The two species are distinguished by the shape of the bulb and the texture of the outer tunics, the colour of the foliage, and the flowering time. An epitype is designated to fix the application of the name Aletris capensis in its current sense, and the nomenclature of V. deasii is clarified, including the designation of a lectotype for V. deasii P.E.Barnes, nom. illeg., non V. deasii Coutts. © 2019 SAAB. Published by Elsevier B.V. All rights reserved.

1. Introduction Veltheimia Gled. (Hyacinthaceae: Scilloideae) is a genus of two species of horticulturally important geophytes endemic to the western and southeastern seaboard and near-interior of South Africa (Manning et al., 2002). It was established by Gleditsch (1771) for a plant that flowered in the Royal Botanic Garden Berlin in 1766 and 1768 and which he identified as Aletris capensis L. (1759). Gleditsch (1771) characterised his new genus on the basis of its tunicated bulb, nodding, tubular flowers with deflexed stamens inserted near the middle of the tube, and inflated, 3-lobed capsules containing a solitary [sic.] seed per locule. The characteristic fruits, produced by another plant in cultivation at Helmstadt, had earlier prompted Fabricius (1763) to propose the new genus Heisteria but that name is an illegitimate later homonym. Although several species of Kniphofia Moench (Asphodelaceae) were included in Veltheimia by later authors on account of their superficially similar flowers, its circumscription in the current sense had been generally established by the early nineteenth century (e.g. Sweet, 1826). By then it was also recognised that at least two species of Veltheimia were in cultivation in the botanic gardens of Europe (Aiton, 1789; Salisbury, 1796), namely a green-leaved species that had been introduced to Britain in 1768 and which was associated with Aletris capensis L. but later generally known under the name *Correspondence to: 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.

V. viridifolia Jacq. (1797), and a glaucous-leaved species that had been described as A. glauca Aiton (1789) shortly after its introduction to Britain in 1781 (Sweet, 1826). The application of these names was reviewed by Obermeyer (1961), and her findings have determined the subsequent usage of names for the two taxa (Marais, 1972). Veltheimia is distinguished from other genera in subfamily Scilloideae by its large, tunicated bulbs, several thin-textured leaves with midrib thickened beneath, well-developed floral bracts and associated bracteole, short-pedicelled, tubular flowers with stamens inserted obliquely in a single series near the middle of the tube, and large, nodding, § inflated, 3-winged, papery capsules. The ovary contains 3 or 4 ovules in each locule although mostly just 2 pear-shaped seeds develop per locule (Manning et al., 2004). The nodding fruits are unique in Hyacinthaceae — in other genera with nodding or pendulous flowers the pedicels either straighten after flowering so that the capsules are spreading or suberect (Hyacinthoides Heist. ex P.C. Fabricius, Lachenalia J.Jacq. ex Murray and Ledebouria Roth: Scilloideae) or flex upwards distally so that the fruits are borne erect (Drimia Jacq.: Urgineoideae; Albuca L., Dipcadi Medik. and Ornithogalum L.: Ornithogaloideae). Molecular analyses confirm Veltheimia as a member of tribe Massonieae, where it is retrieved as sister to a clade comprising the € ll.largely southern African winter-rainfall genera Namophila U.Mu € ll.-Doblies, Massonia Thunb. ex Houtt. and Lachenalia, Doblies & D.Mu with an estimated divergence date of § 30 Ma (Buerki et al., 2012). The genus is inferred to be an ancestral tetraploid, with 2n= 40 (Goldblatt et al., 2012) [the report of 2n = 10 in Duncan and Visagie (2009) is a typographical error].


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Although both species of Veltheimia have been widely covered in the horticultural literature (Batten, 1986; Duncan, 2004; Duncan and Visagie, 2009), the genus has not been treated formally since Baker (1897) and there are several nomenclatural matters that require resolution, among them the authority for V. deasii. In addition the status of the species will benefit from closer analysis using molecular techniques. It is, therefore, useful to have a complete, modern taxonomic account of the genus available, and this is provided here. 2. Materials and methods All relevant types were examined, as well as all herbarium material from BOL, NBG, PRE and SAM, the primary collections of southern African species, as well as selected specimens from GRA (acronyms after Thiers, 2015). Both species were also studied in the field: V. capensis throughout the western part of its range from Montagu to Hondeklipbaai, and V. bracteata near East London. Specimens are cited according to the Degree Reference System (Leistner and Morris, 1976). 3. Results 3.1. Genus Veltheimia Gled. in Hist. Acad. Roy. Sci. (Berlin) 25: 66 (1771). Type species: V. capensis (L.) DC. [Mem. Acad. Berl.: 1769 (1771) 66. (IK)]; Velthaemia Thunb., Fl. cap. 2: 278 (1820), orth. var.; Velthemia € nigl. Akad. Wiss. Mu € nchen 1813: 89 (1814), Schrank, Denkschr. Ko orth. var. Heisteria Fabric., Enum. Meth. Plant. Horti Med. Helmstad.: 447 (1763), nom. illeg., non Heisteria L. (1758) [Polygalaceae], nom. rej., nec Heisteria Jacq. (1760) [Olacaceae], nom. cons. Fabricia Salisb., Prodr. Stirp. Chap. Allerton: 244 (1796). Type: not designated. Deciduous or almost evergreen geophytes, with fleshy perennial roots. Bulb subterranean or partially exposed, large, pear-shaped or subglobose, tunicated, tunics entirely fleshy and not accumulating or outer tunics accumulating then papery or membranous and lightly barred above. Leaves few to several, emergent or fully developed at flowering, suberect or spreading, ovate or oblong-lanceolate to oblanceolate, narrowed and involute basally, weakly canaliculate with a thickened midrib adaxially, thin-textured and sub-succulent or somewhat leathery, margins undulate or crisped. Inflorescence a dense, many-flowered, ovoid or subcylindrical raceme; peduncle erect, usually mottled purple; bracts pale and membranous, lanceolate-attenuate, not spurred, longer than pedicels; bracteoles smaller and solitary, inserted on alternating sides of pedicels near base; pedicels very short, stout. Flowers cream-coloured to greenish but § deeply flushed and spotted with dull red or pink on tube, cylindrical and usually gently curved, spreading or § nodding, unscented, longlasting, tepals connate for most of their length, 1-nerved, lobes small, ovate, penicillate. Stamens fused to tepals for § half their length and obliquely inserted in one series in § middle of tube, deflexed, upper filaments or at least uppermost progressively shorter and inserted slightly lower down tube, filaments slender and terete; anthers just included or shortly exserted, versatile. Ovary narrowly ellipsoid to fusiform and 6-ribbed; ovules 3 or 4 per locule; style slender, terete, slightly declinate, recurved at tip, reaching to mouth of tube; stigma apical, penicillate. Capsule large, papery, 3-winged. Seeds (1)2(4) per locule, pear-shaped, black, smooth or rugulose, testa tightly adhering. 2n = 40 (Goldblatt et al., 2012). Two species in western and southern South Africa. The species are ecological and geographical vicariants, respectively from the arid winter-rainfall parts of southern Namibia and southwestern South Africa, and the subtropical southern and eastern coast of South Africa. The flowers are adapted to pollination by sunbirds (Manning et al., 2002).

3.2. Key to the species 1a. Bulb ovoid, often partly epigeal, outer tunics accumulating and grey-papery; foliage dull or glaucous, emergent or fully developed at flowering; flowering mainly May to early August; plants mainly west of 248E Longitude . . . 1. V. capensis 1b. Bulb subglobose, hypogeal, outer tunics not accumulating; foliage glossy green, fully developed at flowering; flowering mainly mid-August to November; plants east of 248E Longitude . . . 2. V. bracteata 3.3. Species , Liliac. 4(33): t. 193 (1807); Aiton, 1. V. capensis (L.) DC. in Redoute Hort. Kew. 1: 463 (1789); Oberm. in Fl. Pl. Afr. 34: t. 1356 (1961). Aletris capensis L., Syst. Nat., ed. 10, 2: 985 (1759). Veltheimia undulata Moench, Methodus 2: 631 (1794), nom. illeg. superfl. pro. Aletris capensis L. (1794). Fabricia amoena Salisb., Prodr. Stirp. Chap. Allerton: 244 (1796), nom. illeg. superfl. pro Aletris capensis L. (1759). Type: Anon, (LINN [4402], lecto.—digital image!, designated by Oberm. in Fl. Pl. Afr. 34: t. 1356 (1961)]. Epitype: South Africa, Western Cape, Cape Town (3318): Dassenberg, summit of Kanonkop, (DA), 29 Jun 1979, Boucher 4395 (NBG, epi.!, designated here; PRE!, iso.) Aletris glauca Aiton, Hort. Kew. 1: 463 (1789). Fabricia glauca (Aiton) Salisb., Prodr. Stirp. Chap. Allerton: 244 (1796); Veltheimia glauca (Ait.) Jacq., Pl. Rar. Hort. Schoenbr. 1: 40, t. 77 (1797); Baker in Fl. cap. 6: 471 (1897). Type: South Africa, ‘Cape of Good Hope, introduced 1768 by Mr William Malcolm’ (BM, holo.—digital image!). Veltheimia roodeae E.Phillips in Fl. Pl. S. Afr. 4: t. 126 (1924). Type: South Africa, Western Cape, Vanrhynsdorp (3118): ‘Vanrhynsdorp’, June 1923, E. Rood s.n. Nat. Herb. 2739 (PRE[46,727], holo.!). Veltheimia deasii Coutts in Gard. Chron., ser. III, 88: 450 (1930). Type: South Africa, ‘raised from seed received at Kew from the Botanic Gardens, Kirstenbosch, in 19210 , illustration in Gard. Chron., ser. 3, 88: fig. 187 (1930), lecto.!, designated here). Veltheimia deasii P.E.Barnes in S. African Gard. & Country Life 21: 234 (1931), nom. illeg., non V. deasii Coutts (1930). Type: South Africa, Western Cape, Oudtshoorn (3322): ‘in dit Oudtshoorn, prope urbem’, (CA), Jul 1915, W. Deas 17 (GRA[5081], lecto.—digital image!, designated here; NBG!, isolecto.). Other original material: South Africa, Western Cape, ‘in dit Oudtshoorn, prope urbem’, Nov 1914, W. Deas 17 (GRA[5082]—digital image!). Deciduous geophytes. Bulb partly epigeal, ovoid to ellipsoid, 40150 £ 5090 mm, with a well-developed basal plate, scales flushed pink or maroon, outermost layers pale grey or brown and papery, accumulating in a thick mass, forming a lightly barred neck. Leaves emergent or fully developed at flowering, 510(13), suberect or arcuate, ovate to narrowly lanceolate, 100200(300) £ 1550 (70) mm, acute to attenuate, margins undulate or crisped, glaucous. Inflorescence 200500 mm tall, a dense, subglobose to ovoid or cylindrical raceme 4060 mm diam., scape erect, 510 mm diam. at base, pale green lightly to densely speckled and blotched with purple, flower-bearing rachis 25120 mm long; bracts lanceolate, attenuate, pale or flushed pink and membranous, deflexed at anthesis, 820 £ 14 mm, lowermost longest, bracteole suberect to spreading, subulate, 38 £ 0.51.0 mm, usually inserted near base of pedicel but rarely near middle; pedicels arcuate or deflexed apically, 25 mm long. Flowers spreading to nodding at anthesis or lowermost pendulous, perianth tubular, cream-coloured and lightly to densely speckled with pink to red or § entirely reddish pink, usually tipped lime green on the tepal lobes; tube subcylindrical, flaring slightly and uniformly from base, weakly arcuate, (18)2030 £ 46 mm, tepals suberect or § spreading, ovate to lanceolate, 25 £ 24 mm, inner slightly smaller than outer. Stamens inserted obliquely between lower and upper third of tube, reaching mouth of tube or shortly exserted, filaments filiform, 612 mm long, white or flushed purple;

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anthers 2.53.0 mm long at anthesis, yellow. Ovary ellipsoid or fusiform, 812 £ 23 mm, pale green; style weakly deflexed and recurved apically, 1020 mm long, shortly exserted up to 5 mm. Capsules nodding or pendulous, ellipsoidobovoid in outline, 3-winged, 2040 £ 1530 mm, pale-papery. Seeds pear-shaped, 58 £ 45 mm, dull black, testa finely wrinkled. Flowering time: mainly May to early Aug. Fig 1. Distribution and ecology: relatively widely distributed through the drier parts of the winter-rainfall West Coast and near-interior of southern African, from Namuskluft and Noordoewer in southern Namibia and the adjacent Richtersveld in Northern Cape, South Africa southwards along the western escarpment of Namaqualand and the BokkeveldHantam, and through the Cederberg and Swartruggens as far south as Darling and the Paardeberg in Western Cape, thence eastwards through the Breede River Valley and the Little Karoo to Albertinia in the south and the Baviaanskloof in Eastern Cape, and inland along the Roggeveld Escarpment and Nuweveld Mts as far east as Murraysburg (Fig 2); on rocky, S-facing slopes and cliffs in succulent karoo, rarely in sand in strandveld thicket or drier fynbos and sandveld shrublands, with the bulbs partially protruding from rock crevices or above the soil, from near sea-level to 1 500 m. Horticultural aspects of the species are discussed by Duncan and Visagie (2009). Diagnosis: distinguished from Veltheimia bracteata by its deciduous habit, ovoid, partly epigeal bulb with the outer tunics persisting in a thick, papery mass, and its glaucous, often more numerous leaves, usually only partly developed at flowering, which is mainly in winter and early spring. The earlier flowering time is important in distinguishing herbarium collections of plants from the east of the range, especially Baviaanskloof, where V. capensis meets the western limit of V. bracteata. Thus Van Jaarsveld 16,096 (NBG) is associated with V. capensis based on its winter flowering as the bulb has not been preserved. Introgression between the two taxa in this region is also a possibility.

Fig. 1. Veltheimia capensis. 1. Foliage; 2. Raceme; 3. Half-flower; 4. Capsule; 5. Seed. Plant of unknown origin, cultivated at the National Herbarium, Pretoria PRE28871. Original painting reproduced in Flowering Plants of Africa 34: t. 1356 (1961). Artist: Mary E. Connell.


Veltheimia capensis is a widespread species that varies in the number, size and shape of the leaves, in the size of the flowers and the level of insertion of the stamens, and in the size of the capsules and seeds. Narrow-attenuate, closely crispulate leaves are characteristic of populations from the drier eastern interior of the range of the species, from Oudtshoorn in the Little Karoo and from the Nuweveld Mtns east to Murraysburg, but also occur scattered through the range of the species. The flowers of V. capensis also vary in the degree of pink speckling, with the northern populations, in Namaqualand and the Richtersveld, consistently among the most darkly coloured in the species, described variously as deep or wine-red, although plants with dark flowers have been recorded elsewhere as well. Application of the name: Gledditsch (1771) associated the name Aletris capensis L. with a plant that flowered in the Royal Botanic Garden in Berlin in 1766 and 1768. The tubular flowers with stamens inserted near the middle of the tube and the inflated, 3-lobed capsules were so different from those of A. farinosa L., the type of the genus, that Gledditsch had no hesitation in describing the new genus Veltheimia for his plant. Unfortunately he does not describe the bulb tunics or the colour of the leaves of his plant and it is therefore not possible to unequivocally identify the species involved. In any event, the name A. capensis was generally applied to the green-leaved plants that were in cultivation in Europe and Great Britain at the time (e.g. Aiton, 1789; Murray, 1770). By then it was clear that the glaucousleaved plants being grown at Kew represented a separate species, for which Aiton (1789) provided the name A. glauca. Both taxa were beautifully illustrated by Jacquin (1797) from plants then in cultivation at the Botanical Gardens in Vienna, the glaucous-leaved one under the name V. glauca and the green-leaved one under the new name V. viridifolia. The inclusion of the earlier name A. capensis as a synonym of V. viridifolia renders the latter an illegitimate superfluous name (Turland et al., 2018: Art 52.1), as was subsequently realised by , 1807). At the time Jacquin (1797) had already Candolle (Redoute clearly identified the salient differences between the two taxa as being the shape of the bulbs, the colour of the foliage, and the flowering time: V. glauca with an oblong-ovoid bulb, glaucous leaves, and flowering in October and November in the Northern Hemisphere; and V. viridifolia with a rounded bulb, bright green foliage, and flowering from December to February in the Northern Hemisphere. It was in this sense that these two names were used by Baker (1870, 1897) in his monographs of the family and also by later authors, until the investigations by John Lewis at the British Museum in response to a request from the South African botanist Amelia Obermeyer, who was preparing the text to accompany a painting of a glaucous-leaved plant that was figured in Flowering Plants of Africa (Obermeyer, 1961). Following their investigations, Lewis and Obermeyer concluded that the name Aletris capensis, contrary to established usage, represented the glaucous-leaved taxon and that the name Veltheimia capensis was therefore the earliest name for the species that had until then been known as V. glauca. Unfortunately Obermeyer (1961) does not give any justification for this decision, merely identifying a specimen in the Linnean herbarium as the type of the name. Her lectotypification (LINN440-2) (Fig 3), has been accepted by Jarvis (2007). The Linnean lectotype of A. capensis comprises just two detached, relatively narrow leaves and an inflorescence. Without the bulb itself, or evidence of the colour of the foliage, it can only be the leaf shape, possibly reinforced by Burman’s (1768) description of the bulb as conical, which informed Obermeyer’s (1961) identification of the specimen in the Linnean herbarium. Certainly either or both species could have been in cultivation in Europe by 1759 when Linnaeus published his name. Obermeyer’s (1961) application of the name was adopted without question (Marais, 1972) and in the interests of nomenclatural stability (Turland et al., 2018: Art 57.1) it seems prudent to designate an epitype to fix the application of the name in this sense. All references to V. capensis in the literature prior to Obermeyer (1961), and


J.C. Manning / South African Journal of Botany 127 (2019) 271277

Fig. 2. Distribution of Veltheimia capensis () and V. bracteata ().

sometimes shortly thereafter (e.g. Moldenke, 1969), thus refer to the green-leaved V. bracteata and not to V. capensis as currently understood. History: the name Aletris capensis was validated by Linnaeus (1759) with the briefest of descriptions mentioning just the nodding flowers [‘floribus nutantibus’], followed by the cryptic reference ‘Burmannus’. I assume this to be a reference to a specimen acquired from Johannes Burman as no species of Aletris is treated in Burman’s (17381739) Rariorum africanarum plantarum. There is, however, a full description of Linnaeus’ Aletris capensis, including mention of the conical bulb [‘Rad. bulbus conicus’], in the later Prodromus florae capensis by the younger Nicolaas Burman (1768: 10). This description, which is far more extensive than others in the work, includes bulb, foliage, flowers and fruits. This is strong evidence that it was based on a living plant, presumably in cultivation at the Hortus Medicus in Amsterdam where both Burman father and son worked. In addition, the types of some of N.L. Burman’s species are thought to be in the Linnean herbarium (Stafleu and Cowan, 1976: 416), and the lectotype in the Linnean herbarium (Fig 3) designated by Obermeyer (1961) is thus possibly a Burman specimen, although it lacks any indication of its origin. Linnaeus (1762) subsequently published a slightly amplified description of his A. capensis that included mention of lanceolate-undulate leaves and an ovate spike, as well as a provenance from the Cape of Good Hope [‘Aletris acaulis, foliis lanceolatis undulatus, spica ovata, floribus nutantibus. Hab ad Cap. b. spei.’]. The additional descriptive information is consistent with the specimen at issue in his herbarium but the locality can only have been supplied to him through conversation or correspondence, once again possibly from one of the Burmans, with whom he was closely acquainted. Veltheimia roodeae was described by Phillips (1924) for a plant collected near Vanrhynsdorp by local resident Mrs R. Rood in June 1923. It was supposed to differ from the known species in its broader, ovate leaves but falls within the variation of V. capensis and was synonymised under V. capensis by Obermeyer (1961). Veltheimia deasii from near Oudshoorn has a more complicated history. The species was based on plants that were collected by Mr

W. Deas in fruit in November 1914 and later in flower in June 1915. These specimens were deposited in the Albany Herbarium in Grahamstown. Additional plants collected by Mr Deas in September 1915 were cultivated at the Bolus Herbarium in Cape Town, where

Fig. 3. Lectotype of Aletris capensis L. in the Linnean Herbarium (LINN440-2).

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€ nland, Curator of the they flowered in June 1919. Dr Selmar Scho Albany Museum from 1889 until his retirement in 1927, had intended describing Deas’s plants as a new species but left it to P.E. Barnes (1931), who published it as V. deasii, distinguishing it from V. glauca by its smaller size, more closely undulate or crisped leaves, and the insertion of the stamens below the middle of the perianth. Seeds from the plants in cultivation in Cape Town had, however, in the meantime been distributed from Kirstenbosch Gardens to Kew Gardens in 1921, evidently under the manuscript name. These had reached flowering size by 1926. In an extraordinary coincidence, Mr J. Coutts of Kew published a photograph of a flowering plant and brief description under the name V. deasii in the November 1931 issue of The Gardeners’ Chronicle, a little less than one year before Barnes’s publication, thereby rendering Barne’s name an illegitimate later homonym! In any event, the differences between V. deasii and V. capensis are not significant and it was also correctly included in that species by Obermeyer (1961). Additional specimens seen Namibia. 2817 (Vioolsdrif): Noordoewer Hills, W of Marikas Quellen, (AB), 1 000 m, 12 Jul 2005 [fruiting], Bruyns 10,039 (NBG). South Africa. NORTHERN CAPE. 2816 (Oranjemund): Groot Derm, € hnel s.n. (NBG). near Orange River, (DA), 30 Sep 1970 [leafing], Ku 2817(Vioolsdrif): Richtersveld, Stinkfonteinberge SW of Van Zylsrus, €lken and Venter 621 (CA), red flower, 4 Sep 1977 [fruiting], Oliver, To (PRE); kloof on E slopes of Cornellsberg, (CA), 9 Sep 1988 [fruiting], Bruyns 3308 (NBG); 15 km W of Eksteenfontein, 19 May 1985, Williamson 3449 (BOL). 2917 (Springbok): 20 km N of Kleinzee, (CA), 29 Jun 1975, Hardy 3861 (PRE). 3017 (Hondeklipbaai): road to Springbok from Soebatsfontein, (BB), blomme donker rooi [flowers deep red], 21 Jul 1937, Verdoorn and Dyer 1803 (PRE); near Paddagat 22 mi W of Springbok, (BB), humus between rocks on S slope of hill, flowers wine red, 28 May 1961, Leistner 2560 (PRE). 3119 (Calvinia): Nieuwoudtville, Oorlogskloof Nature Reserve, (AC), 29 Jun 2001, Pretorius 717 (NBG); Nieuwoudtville, Wild Flower Reserve, (AC), 19 Apr 1983, Perry & Snijman 2056 (NBG); Nieuwoudtville, Glen Lyon Farm, dolerite flats along trek pad, (AC), 30 May 1993, MacGregor s. n. (NBG); Groot Toren, N of Calvinia, (BD), 1971 [fl. ex hort.], Hardy 2491 (PRE); Lokenburg, (CA), 18 Oct 1953 [fruiting], Acocks 17,486 (PRE); SW-facing slope, fls speckled pink, 9 Jun 1955 [fl. ex hort.], Comins 1132 (PRE); 16 mi [25.6 km] S of Calvinia, (DA), 9 May 1953, Hall 705 (NBG). WESTERN CAPE. 3118 (Vanrhynsdorp): Bitterfontein, (AB), without date, Zeyher 1719 (SAM); Farm Liebendal, 12 km N of Vredendal, (CB), 16 Jun 1973, Hall 4294 (NBG). 3123 (Victoria West): Murraysburg, (DD), Aug 1879, Tyson 359 (SAM). 3217 (Vredenburg): Vredenburg, (DD), 14 Jun 1967, Marsh s.n. (NBG). 3218 (Clanwilliam): Clanwilliam, (BB), 1916, Magistrate s.n. (PRE); 3 mi [5 km] N of Calnwilliam, (BB), 1 Jul 1966, Elzinga s.n. (NBG); Oumuur, between Pakhuis Pass and Klawer (Brandewyn River Valley), (BB), 29 May 1976, Moffett 1109 (NBG). 3219 (Wuppertal): Langrug, Saandkraalkloof, (AC), 26 May 1982, Viviers 344 (NBG); Tankwa Karoo National Park, Leeuberg, (BB), steep S slope, 7 Aug 2006, Sachse 148 (PRE); Grootrivier, W slopes of Blinkberg, (CB), 30000 [914 m], 23 May 1970, Oliver 3156 (NBG); 21 mi [33.6 km] N of Karoo Poort, (DC), 6 Apr 1968, Hall 3162 (NBG); Katbakkies, (DC), 1 Jun 2001, Manning 2337 (NBG). 3221 (Merweville): Fraserburg Dist., Rietvlei, (BB), 30000 [914 m], 29 May 1985 [fl. ex hort.], Shearing 1025 (PRE). 3222 (Beaufort West): Nuweveld Mts, (BA), Jul 1895, Marloth 2119 (PRE); Molteno Pass, near summit, (BA), 17 May 2010, Roux 4732 (NBG). 3223 (Rietbron): Nelspoort, (AA), without date, Pearson 1135 (NBG). 3317 (Saldanha): Saldanha Bay, (BB), May, without year, Zeyher s.n. (SAM); Jul 1912, Marloth 5233 (PRE); May 1959 [fl. ex hort.], Lavranos s.n. (PRE). 3318 (Cape Town): 12 mi [19 km] from Malmesbury on Hopefield road, (AB), 16 Jun 1965, Barler 10,219 (NBG); Hopefield, Waterboerskraal, (AB), sandy soils


amongst restios on small hills, 15 Jun 1975, Hugo 220 (NBG); Darling, Kapokberg, (AD), 13 Jun 1982, Viviers 439 (NBG, PRE); Buck Bay, 19 May 1935, Duckitt s.n. (BOL); Paardeberg, Lemoenkloof, (DB), 2 Jun 2011, Nicolson & Roets 86 (NBG). 3319 (Worcester): Wolseley, Elandskloof, (AC), 23 Jul 1952, Van Niekerk s.n. (BOL); between [railway] tunnel and Matroosberg Station, (BD), 4 May 1978, Bayer 1276 (NBG); De Doorns, (CD), May 1920, Gericke s.n. (PRE); Robertson Dist., Vrolykheid Nature Reserve, (DD), Jul 1976, Van Der Merwe 2841 (PRE). 3320 (Montagu): Doringkloof Farm, S foothills of Voetpadsberg, (AA), dark red-pink flowers, 8 Jun 1985, Van Wyk 2404 (NBG, PRE); Dobbelaarskloof, (CB), 3 Jun 1953, Hall s.n. (NBG); gorge at Montagu Baths, (CC), Aug 1920, Page s.n. (BOL); Cogmanskloof, (CC), 10 Jul 1954, Martin 1014 (NBG); Kalkoennes, riverbed, (CC), 9 Jul 1980, Schonken 266 (NBG, PRE); Montagu, Farm Rietvlei No II, (CD), 18 Jul 1968, Winter s.n. (NBG). 3321 (Ladismith): Ladismith, hillside on road to Gamkapoort Dam, (AD), 14 Jul 1984, Duncan 132 (NBG); Ladismith, range N of Sandberg, (DA), 6 Jun 1956, Wurts 1384 (NBG); 13 km SSE of Calitzdorp, E of Gamka River, (DA), 22 May 1979, Bayer 1753 (NBG). 3322 (Oudshoorn): Cango Caves, (AC), 7 Jun 1953, Carp s.n. (NBG); Boomplaas, Cango Valley, (AC), 26 Jun 1974, Moffett 223 (PRE). 3420 (Bredasdorp): Bredasdorp Dist., Windhoek Provincial Nature Reserve, (AD), kalkrante, ruens teen strandveld, langs vlei teen S hang, rooi blomme [limestone cliffs, hills along strandveld, next to wetland on S slope, red flowers], Jul/Aug 1968, Van der Merwe 854 (PRE); § 10 km N of Malgas on farm Dordregt, (BA), 5 Aug 2002, Helme 2525 (NBG); De Hoop, Potberg Nature Reserve, (DA), 31 Jul 1979, Burgers 2111 (NBG). 3421 (Riversdale): Stillbaai, S of Ellensrust, (AD), 14 Aug 2000, Pienaar & Pienaar 895 (NBG); Farm Tierfontein, S slope of Bark Kloof Spruit, N of Albertinia, (BA), leaves green with a grey bloom, flowers pale pinkish green, 15 Jun 1984, Snijman 793 (NBG, PRE); Middeldrift, by Valsch River, (BA), Jul 1914, Muir 1415 (PRE). EASTERN CAPE. 3323 (Willowmore): Farm Keurfontein, Gwarieppoort area, 15 km SE of Willowmore on road to Uniondale, (AD), 27 Jun 2005, Bester 5824 (PRE); lower S-facing slopes of Witteberg on farm World’s View, (BA), 4 Jul 1987, Viviers and Vlok 168 (PRE); Keurboomsrivier Plato [Plateau], (DA), oop fynbos [open fynbos], Aug 1971 [almost finished], Van der Merwe 2106 (PRE). 3324 (Steyterville): Willowmore Dist, Baviaanskloof, (CA), Jul 1966 [fl. ex hort. BRI], Hardy 2160 (PRE); Baviaankloof, Gert Smitskloof, (CA), 22 Apr 1999 ‘flowering period June and July’, Van Jaarsveld 16,096 (NBG); Kouga River, (CC), 1 July 2010, Schafer 90 (NBG). 2. V. bracteata Harv. ex Baker in J. Linn. Soc., Bot. 11: 411 (1870); Baker in Fl. cap. 6: 471 (1897); Batten, Flowers of Southern Africa: 386389 (1986); Duncan and Visagie in Fl. Pl. Afr. 61: 2433, t. 2244 (2009). Type: South Africa, Eastern Cape, ‘British Kaffararia’, 1860, T. Cooper 320 (K[257,379], holo.—digital image!). Veltheimia viridifolia Jacq., Pl. Rar. Hort. Schoenbr. 1: 41, t. 78 (1797), nom. illeg. superfl. pro A. capensis L. (1759); Baker in Fl. cap. 6: 471 (1897). Type: Illustration in Jacq., Pl. Rar. Hort. Schoenbr. 1: t. 78 (1797). €rtn., [Veltheimia viridiflora hort. in Siebert and Vos, Vilm. Blumenga ed. 3. 1: 1123 (1895), in syn.] [Aletris capensis sensu Murray (1770) et seq., non L. (1759)]  (1807) et seq., non (L.) [Veltheimia capensis sensu DC. in Redoute DC. (1807)] Evergreen or almost evergreen geophytes. Bulb hypogeal, subglobose, 5090 mm diam., scales green or flushed purple, outermost layers papery but not accumulating. Leaves fully developed at flowering, 47, suberect or spreading, oblong to oblanceolate, (100) 200500 £ (30)40100 mm, acute to obtuse, margins undulate or crisped, pale to glossy dark green above but less so below. Inflorescence 250900 mm tall, a dense, subglobose to ovoid raceme 5080 mm diam., scape erect, 617 mm diam. at base, pale to dark green lightly to densely speckled and blotched with purple, flower-bearing rachis 25100 mm long; bracts lanceolate, attenuate, pale and membranous,


J.C. Manning / South African Journal of Botany 127 (2019) 271277

deflexed at anthesis, 832 £ 16 mm, lowermost longest, bracteole suberect to spreading, subulate, 68 £ 0.51.0 mm; pedicels arcuate or deflexed apically, 25 mm long. Flowers spreading to nodding at anthesis or lowermost pendulous, perianth tubular, cream-coloured and lightly to densely speckled with pink to red or § entirely reddish pink, usually tipped lime green on the tepal lobes, rarely uniformly pale greenish yellow; tube subcylindrical, flaring slightly and uniformly from base, weakly arcuate, (20)2232 £ 46 mm, tepals suberect or § spreading, ovate or rarely lanceolate, 35(7) £ 24 mm, inner slightly smaller than outer. Stamens inserted obliquely near middle of tube, reaching mouth of tube or shortly exserted, filaments filiform, 1117 mm long, white or flushed purple; anthers 2.53.0 mm long at anthesis, yellow. Ovary ellipsoid or fusiform, 1012 £ 24 mm, pale green; style weakly deflexed and recurved apically, 1928 mm long, shortly exserted. Capsules nodding or pendulous, ellipsoidobovoid in outline, 3-winged, 3550 £ 2030 mm, pale-papery. Seeds pearshaped, 56 £ 45 mm, dull black, testa finely wrinkled. Flowering time: mainly mid-Aug to Nov. Fig 4. Distribution and ecology: endemic to coastal Eastern Cape, with a relatively restricted distribution from St Francis Bay in the south to just north of Kei Mouth (Fig 2), typically below 100 m elevation; in coastal bush and scrub on fixed dunes or among rocks, often just above the high-tide level, or shortly inland in riverine thicket or on cliffs. The foliage is often shaded by surrounding vegetation, with the inflorescence protruding into the sunlight. Horticultural aspects of the species, including some named cultivars, are covered by Duncan and Visagie (2009). Diagnosis: distinguished from Veltheimia capensis by its § evergreen habit, subglobose, hypogeal bulb with the outer tunics not persisting, and the glossy pale to deep green foliage, always fully developed at flowering, which is mainly in spring and early summer.

Fig. 4. Veltheimia bracteata. 1. Raceme; 2. Foliage; 3. Detached flower; 4. Half-flower; 5. Infructescence. South Africa, Eastern Cape, Sardinia Bay, Tennant s.n. (NBG). Original painting reproduced in Flowering Plants of Africa 61: t. 2244 (2009). Artist: Marita Visagie.

The later flowering time is important in distinguishing herbarium collections of plants from the west of the range, especially Baviaanskloof, where V. bracteata meets the eastern limit of V. capensis. Bract length is variable in both species but plants with large bracts > 20 mm long are only encountered in V. bracteata, the bracts in V. capensis never being more than 20 mm long. Albinoid individuals with uniformly pale greenish or yellowish flowers occur sporadically and have been introduced into cultivation, along with the typical pink or reddish form (Duncan and Visagie, 2009). History: this species has a relatively straightforward history, complicated solely by its confusion with V. capensis, as explained under that species. It was generally known throughout the nineteenth and the first half of the twentieth century under the nomenclaturally illegitimate name V. viridifolia until this was corrected by Marais (1972), at which time the later synonym V. bracteata Baker (1897) was adopted. The latter species was distinguished from V. viridifolia by its longer bracts, as long as the flowers, but this character is variable, even within a single population. Additional specimens seen South Africa. EASTERN CAPE. 3228 (Butterworth): Willowvale Dist., 3 mi [5 km] from Cats Pass, (AD), cliffs above river, 13 Aug 1966, Strey 6662 (PRE); Qora River, Legewaan Rocks, 2 km from mouth, (BC), among rocks with Clivia miniata, 14 Sep 2001, Rourke 2213 (NBG); near Kei Mouth, (CB), under bush, Sep 1890 [fl. ex hort.], Aug 1891, Flanagan 324 (PRE, SAM); Qolora Mouth, (CB), among rocks in forest shelter, 1 Sep 1915, Pegler 2132 (PRE). 3324 (Steytlerville): Baviaanskloof, Poortjies, (DA), 9 Aug 2002, Bester 1477 (PRE). 3325 (Port Elizabeth): Uitenhage Dist., (BC), without date, Ecklon and Zeyher 28 (PRE, SAM); Uitenhage, Enon, (BC), Nov 1926 [fruiting], Thode s.n. (NBG, PRE); Sep 1930, Thode s.n. (PRE); Van Staden’s Pass, (CC), without date, Letty s.n. (PRE); Van Staden’s Flower Reserve, (CC), 16 Nov 1972, Dahlstrand 2973 (PRE); Sardinia Bay, (CD), Oct 1986, Tennant s.n. (NBG); Redhouse, (DC), Sep 1914, Paterson 125 (BOL). 3326 (Grahamstown): Grahamstown, (BC), Oct 1902, Daly and Sole 322 (PRE); Sep 1918, Van Dam 18,866 (PRE); Grahamstown, (BC), Farm Glen Boyd, mauve or pinkish mauve, no scent, 20 Jul 1926, Liverstedt 9 (PRE); Bathurst Dist., Hopewell, S of Southwell, (BD), fairly frequent on krantz, 12 Sep 1951, Acocks 16,142 (PRE); Alexandria Dist., Kaba, (CB), 23 Jul 1954 [in bud], Archibald 5513 (PRE); Kasonga Mouth, (DA), east bank, shady part at base of slope, fls dull pink, 22 Sep 1920, Britten 2417 (PRE); Leonard’s Bay, S of Kidd’s Beach, (DA), among rocks covered with wind-sheared coastal scrub, 15 Aug 2007, Rourke 2270 (NBG); Kwaaihoek at Diaz Cross Memorial, (DA), 14 Aug 2007, Rourke 2268 (NBG); Kowie, (DB), without date, Britten 771 (GRA); flowers green but much speckled with pink, the lips quite green without date, Marloth 7037 (PRE); Port Alfred, (DB), Aug 1916, in bush, Tyson 17,545 (PRE); Nov 1923, Rogers s.n. (NBG); Port Alfred, Kowie River, (DB), 25 Dec 1951 [fruiting], Verdoorn 1 (PRE). 3327 (Peddie): Kayser’s Beach, (BA), coastal bush above beach, 15 Aug 2007, Rourke 2269 (NBG); East London, in bush near river mouth, (BB), Aug 1896, Galpin 3350 (BOL, GRA, PRE); Aug 1944, Malin s.n. (PRE); near East London, (BB), Nov 1896, Griffith s.n. (PRE); Kefani, (BB), bush under trees, 26 July 1961, Batten 2 Pl 87 (NBG). 3424 (Humansdorp): Eerste River, (AA), seashore, perianth light red, upper half mottled green, peduncle mottled brown-purple, Aug 1921, Fourcade 1389 (BOL, NBG, PRE); St Francis Bay, (BB), coastal dune, 25 Sep 2000, Brand 230 (PRE). Hybrid taxa Veltheimia £ traubii Moldenke in Pl. Life 25: 44 (1969). Type: Cult. La Jolla, Calif., Traub 1085 (TRA, holo., not located). This taxon was described as the interspecific hybrid V. glauca (Aiton) Jacq. £ V. capensis (L.) DC., and was characterised by precocious flowering as in the pollen parent [‘velut in parenta mascula’] but otherwise

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intermediate between the two. The early flowering identifies the pollen parent as V. glauca and suggests that Moldencke was still using the name V. capensis in its old sense as applying to what is currently called V. bracteata. It can be taken therefore to be a true interspecific hybrid following the current taxonomy of the species and not as a direct synonym of V. capensis/glauca. Excluded taxa , Liliac. 4: t. 186 (1807) = Kniphofia Veltheimia abyssinica Redoute pumila (Aiton) Kunth (Codd, 2005). [Veltheimia burchellia [sic.] Hort. in Siebert and Vos, Vilm. Blu€rtn., edn. 3, 1: 1077 (1895), in syn.]. Kniphofia burchellii (Lindl.) menga Kunth = Kniphofia uvaria (L.) Oken. Veltheimia intermedia Sweet, Hort. Brit.: 409 (1826). Type: not designated. The taxon was identified as the ‘Subglaucous Veltheimia’ to distinguish it from the Glaucous Veltheimia (V. glauca) and Greenleaved Veltheimia (V. viridifolia), and this forms the sole basis for validating and identifying it. Without details of the bulb tunics it is impossible to associate it with either of the two recognised species, and the further possibility exists that it may be a hybrid between them. Veltheimia media Donn, Hort. Cantabrig., edn. 2: 131 (1800) = Kniphofia sarmentosa (Andrews) Kunth (Codd, 2005). Veltheimia pumila (Aiton) Willd., Sp. Pl., edn. 4, 2(1): 182 (1799) = Kniphofia pumila (Aiton) Kunth Veltheimia repens Ker-Gawl., Recens. Pl.: 18 (1801), nom. illeg. superfl. pro Kniphofia sarmentosa (Andrews) Kunth (Codd, 2005). Veltheimia sarmentosa (Andrews) Willd., Enum. Pl. 1: 380 (1809) = Kniphofia sarmentosa (Andrews) Kunth Veltheimia speciosa Roth, Nov. Pl. Sp.: 190 (1821) = Kniphofia sp. (Codd, 2005). Veltheimia uvaria (L.) Willd., Sp. Pl., ed. 4, 2(1): 182 (1799) = Kniphofia uvaria (L.) Oken Acknowledgements 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, and Anthony Magee for preparing them for publication; and Michelle Smith for preparing the distribution maps. References Aiton, W., 1789. Hortus Kewensis, 1. Nicol, London. Baker, J.G., 1870. Monograph of scilla: x ledebouria and drimiopsis. Refugium Botanicum [Saunders] 3, 7–13 App.. Baker, J.G., 1896. Liliaceae. In: Thiselton-Dyer, WT (Ed.), Flora Capensis. 6, L. Reeve & Co., Kent, pp. 253–528. Barnes, P.E., 1931. Plants—New or noteworthy. South African Gardening and Country Life 21, 228. Batten, A., 1986. Flowers of Southern Africa. Fransden, Sandton. Buerki, S., Jose, S., Yadav, S.R., Goldblatt, P., Manning, J.C., Forest, F., 2012. Contrasting biogeographic and diversification patterns in two mediterranean-type ecosystems. PlosOne 1–11 7 e39377.


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