South African Journal of Botany 72 (2006) 105 – 132 www.elsevier.com/locate/sajb

The endemic and near-endemic angiosperms of the Drakensberg Alpine Centre

C. Carbutt, T.J. Edwards *

School of Biological and Conservation Sciences, University of KwaZulu-Natal Pietermaritzburg, Private Bag X01, Scottsville 3209, South Africa

Received 25 January 2005; accepted 8 June 2005

Abstract

The Drakensberg Alpine Centre (DAC) of southern Africa supports a speciose number of angiosperms. Its compliment of 2520 native angiosperms is comparable with the native floras of northern hemisphere countries such as Germany or Switzerland, and is almost double that of Great Britain. Levels of angiosperm endemism and near-endemism in the DAC too are high, that are here treated within a conservation framework to draw attention to the local and global significance of its biodiversity. The DAC supports some 334 endemic and 595 near-endemic angiosperms, meaning that almost 37% if its flora is confined to a core region south of the Limpopo River. Strict endemism is c. 13%, slightly lower than the 16% endemism for KwaZulu-Natal, and substantially lower than endemism (c. 69%) in the Cape Floristic Region. Most endemic and near- endemic taxa belong to the Asteraceae, Scrophulariaceae and Iridaceae, which are the largest, fourth largest and seventh largest angiosperm families in the DAC respectively. Helichrysum and Senecio contribute the most endemics and near-endemics. Many of the endemics are rare, and have very specific habitat preferences. Some 42% of the endemic taxa, and 16% of the near-endemic taxa, are Red Data species. Here we add a further 42 endemic taxa (c. 13%) to the Orange List, bringing the total number of endemics that are now either Red or Orange Data listed to 182 (c. 55%); their conservation needs to be prioritised. A strong plea is made for the continued sustainable management and protection of biota in the DAC, and the mitigation of any threats that may lead to their demise. D 2005 SAAB. Published by Elsevier B.V. All rights reserved.

Keywords: Angiosperms; Drakensberg Alpine Centre; Endemism; Flora; Near-endemism; Orange list; Plant conservation; Radiation

1. Introduction Cape Drakensberg and Witteberge; and the Sehlabathebe/East Griqualand/Naude’s Nek region (Hilliard and Burtt, 1987). The Drakensberg Alpine Centre (DAC) (CPD Site Af82, The DAC occurs within White’s (1983) archipelago-like WWF and IUCN, 1994) covers some 40,000 km2 (Fig. 1) centre of endemism, comprising the Afromontane and Afroal- within the greater Drakensberg Range, the latter extending for pine phytochoria. At a finer scale, White’s (1978) classification almost 1000 km along the eastern boundary of the southern of the Afromontane archipelago recognised seven regional African plateau (Partridge and Maud, 1987). Politico-geo- mountain systems, of which the Drakensberg Range (including graphically, the DAC is subdivided into four domains (Fig. 1): the DAC) is the most southern. The DAC is regarded as being the KwaZulu-Natal Drakensberg, the Eastern Cape Drakens- the only true alpine region in southern Africa (Linder, 1990), berg and Witteberge, the Maloti Mts. of Lesotho, and the and, in conjunction with the Cape Floristic Region (CFR) highlands of the eastern Free State (Carbutt and Edwards, (Goldblatt and Manning, 2002; Linder, 2003), has been 2004). The DAC is subdivided floristically into the alpine proposed as the southern (Gondwanan) source of the temperate region of the KwaZulu-Natal Drakensberg summit and Lesotho flora of Africa (Hilliard and Burtt, 1987; Linder, 1990, 1994). Malotis; its outliers (e.g. Mahwaqa and Ngeli Mts.); the Among the centres of plant endemism recognised by Van northern KwaZulu-Natal Drakensberg scarp face; the southern Wyk and Smith (2001), the DAC (using Carbutt and Edwards’ KwaZulu-Natal Drakensberg scarp face; the southern Eastern (2004) estimate of 2618 species of vascular plants) ranks as having the fourth richest regional flora in southern Africa, * Corresponding author. surpassing even the richness of many of the subtropical and E-mail address: [email protected] (T.J. Edwards). semi-arid floras of southern Africa. Despite its recognition as a

0254-6299/$ - see front matter D 2005 SAAB. Published by Elsevier B.V. All rights reserved. doi:10.1016/j.sajb.2005.06.001 106 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

28oE

Free State South Africa 29oS

Drakensberg Alpine Centre Lesotho KwaZulu-Natal

E. Cape

Indian Ocean Eastern Cape N

50 km

Fig. 1. The Drakensberg Alpine Centre (DAC) comprising eastern Free State, KwaZulu-Natal Drakensberg, Lesotho Maloti Mts. and Eastern Cape Drakensberg/ Witteberge. regional centre of endemism, its endemic plants, and their harvesting of native plants (Cowling and Hilton-Taylor, 1994, conservation status, remain poorly known. Talukdar, 1994). The implications of improper protection do Only a small fraction (c. 5.5%) of the DAC is currently not bear solely on the DAC’s phytodiversity: the DAC also conserved (Cowling and Hilton-Taylor, 1994; Killick, 1994). serves as southern Africa’s premier water catchment, and is The great majority of this (c. 97%; Killick, 1994) lies within home to a diverse fauna (Porter et al., 1999). It is therefore a the uKhahlamba-Drakensberg Park (KwaZulu-Natal, South region of strategic biological and socio-economic importance Africa), recently declared a World Heritage site for outstanding and concern, with national and international ramifications. biological and cultural attributes (Derwent et al., 2001). This The aims of this study, therefore, were to accurately Park, covering 242,813 ha (Porter et al., 1999), is a region of document the full quota of endemic and near-endemic strategic conservation importance and one that contains much angiosperms of the DAC (and not solely for its sub-floras (c. 80%) of the DAC’s vascular plant diversity. Less than a defined by political boundaries as in the past); make handful of small parks protect biodiversity in the Lesotho comparisons with the endemic angiosperms of other (mostly portion of the DAC, namely the Sehlabathebe (proclaimed), high-altitude) floras in southern and south-central Africa; Bokong and Ts’ehlanyane National Parks (proclamation scrutinise the number of Red Data (IUCN, 1994) and Orange pending). Despite the Lesotho Maloti Mts. encompassing Data (Victor and Keith, 2004) species in order to assign them much of the DAC, only 3% of the conserved land in the conservation priority; draw international attention to a poorly DAC falls within this region (Killick, 1994). The Basuto known southern hemisphere temperate flora; and highlight the people of this region are rural herdsmen and pastoralists, who difficulties and problems associated with such a study. follow a subsistence-based form of agriculture. As a result, biodiversity threats to this area include severe overstocking of 1.1. History of endemic accounts domestic animals (and therefore overgrazing), large-scale soil erosion, invasive exotic plants and inappropriate cropping Hilliard and Burtt (1987) estimated that 394 angiosperm practices in an area poorly suited to arable agriculture (Hall et taxa are endemic to the Eastern Mountain Region. This Region, al., 1984; Killick, 1994; Talukdar, 1994; Carbutt and Edwards, however, extends slightly beyond the DAC, and a number of 2004). A further threat to plant diversity in the DAC is the their 394 taxa are therefore referred to as near-endemic in this Lesotho Highlands Water Project (LHWP), which once (or if) account. Hilliard and Burtt’s (1987) Eastern Mountain Region all phases are completed, will inundate more than one-third of extended to as low as 1500 m a.s.l. in parts, to include the Lesotho’s total surface area with water. The proposed five large highlands of the former Transkei (e.g. Baziya, Insikeni, Insizwa dams (Katse and Mohale Dams completed) are destined to and Tabankulu), and the hills as far south as Queenstown. supply water to the highly industrialised Witwatersrand These outliers are not considered part of the DAC, and the complex in South Africa (Talukdar, 1994). Other threats DAC and Eastern Mountain Region are therefore only loosely include afforestation and the subsistence and commercial equivalent. C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 107

Hilliard and Burtt’s (1987) estimate of angiosperm ende- A near-endemic taxon is one marginally present elsewhere, mism for the Eastern Mountain Region (c. 29%) was inflated sometimes in the form of distant satellite populations (Mat- because the floras of the Lesotho Highlands and Eastern Cape thews et al., 1993). Admittedly, determining a region’s near- Drakensberg, ill-explored at the time, were never incorporated endemic flora is, to a measure, a subjective exercise. This study into their calculation of endemism (Carbutt and Edwards, in no way attempted to invoke centres of origin and directions 2001). A concurrent regional overview of endemism by Van of migration, but aimed rather at recording all taxa that have a Wyk and Smith (2001) recognised the DAC as one of 18 definite present-day station in the DAC, and are mostly centres of plant endemism in southern Africa, and estimated the restricted to the Eastern Region (refer to Fshared distribution number of endemics in the DAC at >400 taxa, out of a total of ranges_). Documenting the near-endemic flora is important, as c. 2200 taxa (c. 18% endemism). These figures were used by it serves as the Fsafety net_ to accommodate a number of taxa Pooley (2003), whom makes mention of Eastern Mountain that are almost confined wholly to the DAC. Their outlying Region endemics, rather than DAC endemics, and refers to the stations are seldom within conservation areas (e.g. outliers two regions as synonymous. Pooley (2003) refers to Eastern occurring in Mistbelt grassland in the KwaZulu-Natal Mid- Mountain Region endemics as species occurring above the lands, of which only c. 3.7% remains untransformed, and a 1500 m contour. A recent study (Carbutt and Edwards, 2004) meagre 0.3% is formally conserved, Scott-Shaw et al., 1996). documented 2520 native angiosperm taxa, and c. 16% A taxon currently near-endemic to the DAC may in the future endemism. The precise number of endemics, as well as their be rendered strictly endemic, as its outlying stations may be conservation status, was not investigated. lost from unconserved rangelands. The benefit of including a near-endemic appendix is that it also allows the inclusion of 2. Materials and methods taxa that may be true endemics, but have been excluded from the endemic appendix because outlying records still await The following sources were used for the compilation of this taxonomic scrutiny. inventory: Hilliard and Burtt (1987), Van Wyk and Smith The DAC endemics and near-endemics had their conserva- (2001), Pooley (1993, 1998, 2003), and Germishuizen and tion status assessed within the framework of the International Meyer (2003). The selection of taxa was verified using specific Union for Conservation of Nature and Natural Resources taxon treatments, in combination with specimens lodged in the (IUCN, 1994) criteria, using the treatments of Scott-Shaw Bews Herbarium (NU) of the University of KwaZulu-Natal. (1999), Talukdar (2002) and Victor (2002). The broad Certain plant literature was sourced using FThe International categories of Fextinct or threatened_, Fat lower risk_ and Fdata Plant Names Index_ (IPNI, 2004). Dubious records were deficient_ were used to comply with existing Red Data omitted. The 1800 m contour limit was used as the delimiting treatments (see Talukdar, 2002; Victor, 2002). Treatments by boundary of the DAC, following Hilliard and Burtt (1987) and Hall et al. (1980) and Hilton-Taylor (1996a, b, 1997), using VanWykandSmith(2001), thereby including outlying IUCN criteria that are now outdated, were also consulted. Taxa enclaves such as Little Bamboo Mountain (2421 m), Kamberg assessed using the treatments of the latter two authors were (2095 m), Mahwaqa Mountain (2083 m) and the Ngele Range listed separately from those mentioned formerly because of the (2268 m). Floristic ties between the southern KwaZulu-Natal disparity in the criteria used to assign them to their particular Drakensberg and these outliers are strong (JE Granger, Red Data categories. Certain poorly known and rare DAC unpublished data), which further justified their inclusion into endemics, previously unlisted, were added to the newly the DAC. The 1800 m cut-off was difficult to maintain because developed Orange List of Victor and Keith (2004). This safety the DAC’s alpine regions spread to slightly lower elevations at net is a proactive measure towards protecting the species-rich its northern and southernmost limits (1700 m). The checklist flora of southern Africa (FSA), and will hopefully preclude arrangement conforms mostly to the Englerian system followed Fborderline_ taxa from future Red Listing (Victor and Keith, by Germishuizen and Meyer (2003), with Scrophulariaceae 2004). following Hilliard (1994, 1999). Three newly described species The endemics of the DAC were compared with the of Hesperantha were sourced from Goldblatt (2003), and endemics of other (mostly high-altitude) floras in southern recent changes to Cliffortia were updated from Whitehouse and south-central Africa. Their species-area relationships were (2004). The endemic and near-endemic angiosperms of the represented as double-log plots. The near-endemics were DAC are presented as two appendices; both include infraspe- represented as a number of Fphytogeographical_ groups based cific taxa (subspecies and varieties). on their shared distribution ranges. Major’s (1988) definition of endemism was followed: Fa taxon is endemic if confined to a particular area through 2.1. Challenges historical, ecological or physiological reasons_.ADAC endemic is defined as any taxon restricted to the DAC, A number of taxa (Alchemilla, Cyperaceae, Delosperma, occurring 1800 m a.s.l. Taxa occurring marginally lower Erica, Thesium and Wahlenbergia) were difficult to assess (i.e. 1700–1799 m), yet within the uKhahlamba-Drakensberg due to the paucity of updated or comprehensive botanical Park, were included. Taxa occupying similar altitudes imme- revisions. The lack of congruency between botanical accounts diately beyond the boundaries of the uKhahlamba-Drakensberg often made the placement of taxa difficult. Borderline taxa Park were regarded as near-endemic. were omitted. 108 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

3. Results and discussion analyses reveal that c. 42% (140) of the endemics and c. 16% (95) of the near-endemics are Red Data listed. 3.1. Habitat specificity The task of assessing the conservation status of the remaining endemics was a difficult one. Most aspects of their A number of DAC endemics (47 or c. 14%) are highly natural history, especially their population biology (estimates localized and are probably holoendemic. Those taxa known of population size, range, evidence of recruitment, breeding from only one or two sites are listed in Table 1. Restricted systems) and pollination biology are unknown. Here we add a ranges may be a true reflection of small populations of low further 42 endemics (c. 13%) to the Orange List (Table 3). A abundance; otherwise, they may be the subjects of under- total of 182 endemics (c. 55%) are now either Red or Orange collection. Data listed and their conservation needs to be prioritised. This is five times greater than the percentage of rare and threatened 3.2. Conservation plants in KwaZulu-Natal, and the World (see Scott-Shaw, 1999), adding further support to southern Africa being Scott-Shaw (1999), Talukdar (2002) and Victor (2002) recognised as having the highest known concentration of afforded 219 of the taxa (133 endemic; 86 near-endemic) Red threatened plants in the World (Hilton-Taylor, 1996a). A large Data status following the IUCN’s (1994) criteria (Table 2A). proportion (30 or c. 70%) of the 42 Orange Listed endemics, Hilton-Taylor’s (1996a, b, 1997) accounts, mostly following however, are protected in the uKhahlamba-Drakensberg Park the earlier IUCN categories (Davis et al., 1986), included a (Table 3). further 15 taxa (7 endemic; 8 near-endemic), not treated by the previous three authors (Table 2B). The analysis of Hall et al. 3.3. Diversity, endemism and near-endemism (1980) contributed a single taxon near-endemic to the DAC (Table 2B). In total, therefore, 235 taxa, or c. 25% of all The DAC (åEastern Mountain hot-spot) is recognised as endemics and near-endemics are Red Data listed. Separate one of southern Africa’s eight Fhot-spots_ (Cowling and Hilton-

Table 1 Endemic taxa of the DAC recorded only from one or two sites Taxa recorded from one site Taxa recorded from two sites Taxon Family Taxon Family Anthoxanthum brevifolium Stapf Poaceae Alepidea insculpta Hilliard & B.L. Burtt Apiaceae Aponogeton ranunculiflorus Jacot Guill. & Maraisa Aponogetonaceae Athanasia grandiceps Hilliard & B.L. Burtt Asteraceae Brachystelma perditum R.A. Dyer Apocynaceae Cephalaria galpiniana Szabo´ subsp. galpiniana Dipsacaceae Carex killickii Nelmes Cyperaceae Cyrtanthus erubescens Killick Amaryllidaceae Clutia alpina Prain Euphorbiaceae Euryops evansii Schltr. subsp. parvus B. Nord. Asteraceae Crassula qoatlhambensis Hargr. Crassulaceae Festuca vulpioides Steud. Poaceae Diascia lilacina Hilliard & B.L. Burtt Scrophulariaceae Fuirena tenuis P.L. Forbes Cyperaceae Dracosciadium saniculifolium Hilliard & B.L. Burtt Apiaceae Hesperantha brevistyla Goldblatt Iridaceae Drimia saniensis (Hilliard & B.L. Burtt) J.C. Hyacinthaceae Jamesbrittenia jurassica (Hilliard & B.L. Burtt) Scrophulariaceae Manning & Goldblatt Hilliard Euryops brevipes B. Nord. Asteraceae Jamesbrittenia lesutica Hilliard Scrophulariaceae Felicia caespitosa Grau Asteraceae Lotononis jacottetii (Schinz) B.-E. van Wyk Fabaceae Gladiolus loteniensis Hilliard & B.L. Burtt Iridaceae Lotononis minor Dummer & Jenn. Fabaceae Gnidia singularis Hilliard Thymelaeaceae Pentaschistis praecox Linder Poaceae Helichrysum hypocephalum Hilliard Asteraceae Romulea macowanii Baker var. alticola Iridaceae (B.L. Burtt) M.P. de Vos Indigofera pseudoevansii Hilliard & B.L. Burtt Fabaceae Selago leptothrix Hilliard Scrophulariaceae Isolepis pellocolea B.L. Burtt Cyperaceae Senecio parentalis Hilliard & B.L. Burtt Asteraceae Jamesbrittenia beverlyana (Hilliard & B.L. Burtt) Hilliard Scrophulariaceae Senecio submontanus Hilliard & B.L. Burtt Asteraceae Kniphofia hirsuta Codd Asphodelaceae Trachyandra smalliana Hilliard & B.L. Burtt Asphodelaceae Nemesia glabriuscula Hilliard & B.L. Burtt Scrophulariaceae Protea nubigena Rourke Proteaceae Schoenoxiphium burttii Kukkonen Cyperaceae Schoenoxiphium distinctum Kukkonen Cyperaceae Schoenoxiphium molle Kukkonen Cyperaceae Schoenoxiphium strictum Kukkonen Cyperaceae Senecio cristimontanus Hilliard Asteraceae Wahlenbergia doleritica Hilliard & B.L. Burtt Campanulaceae Wurmbea burttii B. Nord. Colchicaceae Wurmbea tenuis (Hook.f.) Baker subsp. australis B. Nord. Colchicaceae Xerophyta longicaulis Hilliard Velloziaceae FTwo sites_ refers to two separate gatherings from disparate quarter degree grids. a A tentative inclusion for the present time owing to a possible gathering from the Bokkeveld Mts. near Nieuwoudtville in the Western Cape (see Goldblatt and Manning, 2000). This also applies to its inclusion in Appendix 1. C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 109

Table 2A Red Data treatments for plant taxa forming part of the flora of southern Africa (FSA), which are either endemic or near-endemic to the Drakensberg Alpine Centre (DAC) Red Data treatment by Talukdar (2002) and Victor (2002) Red Data treatment by Scott-Shaw (1999) FExtinct or threatened_ (n=24) Aloe polyphylla Scho¨nland ex Pillans Crocosmia pearsei Oberm. Aloe pratensis Baker Protea nubigena Rourke Aponogeton ranunculiflorus Jacot Guill. & Marais Brachystelma alpinum R.A. Dyer Carex killickii Nelmes Crassula qoatlhambensis Hargr. Disa scullyi Bolus Drimia saniensis (Hilliard & B.L. Burtt) J.C. Manning & Goldblatt Ehrharta longigluma C.E. Hubb. Euryops brevipes B. Nord. Felicia wrightii Hilliard & B.L. Burtt Gnaphalium griquense Hilliard & B.L. Burtt Helichrysum alticolum Bolus Helichrysum haygarthii Bolus Helichrysum nimbicola Hilliard Jamesbrittenia beverlyana (Hilliard & B.L. Burtt) Hilliard Jamesbrittenia lesutica Hilliard Macowania conferta (Benth.) E. Phillips Macowania deflexa Hilliard & B.L. Burtt Macowania hamata Hilliard & B.L. Burtt Thamnocalamus tessellatus (Nees) Soderstr. & R.P. Ellis Wahlenbergia tetramera Thulin

FAt lower risk_ (n=140) Aloe aristata Haw. Asclepias oreophila Nicholas Athanasia grandiceps Hilliard & B.L. Burtt Aspidonepsis cognata (N.E.Br.) Nicholas & Goyder Brachystelma perditum R.A. Dyer Aspidonepsis flava (N.E.Br.) Nicholas & Goyder Brachystelma petraeum R.A. Dyer Aspidonepsis reenensis (N.E.Br.) Nicholas & Goyder Brunsvigia undulata F.M. Leight. Aster ananthocladus Hilliard & B.L. Burtt Corycium flanaganii (Bolus) Kurzweil & H.P. Linder Aster confertifolius Hilliard & B.L. Burtt Cyrtanthus epiphyticus J.M. Wood Berkheya draco Roessler Disa basutorum Schltr. Berkheya pannosa Hilliard Disa cephalotes Rchb.f. subsp. frigida (Schltr.) H.P. Linder Comborhiza virgata (N.E.Br.) Anderb. & K. Bremer Disa montana Sond. Cotula lineariloba (DC.) Hilliard Disa oreophila Bolus subsp. erecta H.P. Linder Cotula paludosa Hilliard Disa pulchra Sond. Cotula radicalis (Killick & C. Claassen) Hilliard & B.L. Burtt Disa sankeyi Rolfe Crocosmia pottsii (Macnab ex Baker) N.E.Br. Disa thodei Schltr. ex Kraenzl. Cyrtanthus erubescens Killick Disa tysonii Bolus Cyrtanthus falcatus R.A. Dyer Disperis concinna Schltr. Diascia anastrepta Hilliard & B.L. Burtt Disperis cooperi Harv. Diascia austromontana K.E. Steiner Disperis stenoplectron Rchb.f. Diascia megathura Hilliard & B.L. Burtt Disperis tysonii Bolus Diascia purpurea N.E.Br. Disperis wealei Rchb.f. Diascia tugelensis Hilliard & B.L. Burtt Eulophia zeyheriana Sond. Diascia vigilis Hilliard & B.L. Burtt Glumicalyx lesuticus Hilliard & B.L. Burtt Dierama cooperi N.E.Br. Helichrysum amplectens Hilliard Dierama tysonii N.E.Br. Helichrysum longinquum Hilliard Dracosciadium saniculifolium Hilliard & B.L. Burtt Helichrysum palustre Hilliard Erica anomala Hilliard & B.L. Burtt Helichrysum sessile DC. Erica dominans Killick Huttonaea woodii Schltr. Erica ebracteata Bolus Jamesbrittenia jurassica (Hilliard & B.L. Burtt) Hilliard Erica straussiana Gilg Macowania corymbosa M.D.Hend. Erica thodei Guthrie & Bolus Nerine bowdenii Watson Erica tysonii Bolus var. tysonii Nerine pancratioides Baker Fanninia caloglossa Harv. Neobolusia tysonii (Bolus) Schltr. Galtonia princeps (Baker) Decne. Osteospermum attenuatum Hilliard & B.L. Burtt Galtonia regalis Hilliard & B.L. Burtt Othonna burttii B. Nord. Geranium drakensbergensis Hilliard & B.L. Burtt Rhodohypoxis incompta Hilliard & B.L. Burtt Gladiolus loteniensis Hilliard & B.L. Burtt Rhodohypoxis thodiana (Nel) Hilliard & B.L. Burtt Gladiolus symonsii F. Bolus Satyrium microrrhynchum Schltr. Glumicalyx flanaganii (Hiern) Hilliard & B.L. Burtt (continued on next page) 110 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

Table 2A (continued) Red Data treatment by Talukdar (2002) and Victor (2002) Red Data treatment by Scott-Shaw (1999) Schizochilus flexuosus Harv. ex Rolfe Gnidia renniana Hilliard & B.L. Burtt Senecio austromontanus Hilliard Helichrysum drakensbergense Killick Senecio saniensis Hilliard & B.L. Burtt Helichrysum evansii Hilliard Wahlenbergia cuspidata Brehmer Helichrysum hyphocephalum Hilliard Zaluzianskya oreophila Hilliard & B.L. Burtt Helichrysum tenax M.D.Hend. var. pallidum Hilliard & B.L. Burtt Hemizygia cinerea Codd Hesperantha alborosea Hilliard & B.L. Burtt Hesperantha curvula Hilliard & B.L. Burtt Hesperantha ingeliensis Hilliard & B.L. Burtt Hesperantha woodii Baker Hilliardia zuurbergensis (Oliv.) B. Nord. Hoffmannseggia sandersonii (Harv.) Engl. Hypoxis ludwigii Baker Indigofera evansii Schltr. Kniphofia angustifolia (Baker) Codd Kniphofia brachystachya (Zahlbr.) Codd Kniphofia breviflora Baker Kniphofia evansii Baker Kniphofia ichopensis Schinz var. aciformis Codd Kniphofia ichopensis Schinz var. ichopensis Kniphofia northiae Baker Lessertia harveyana L. Bolus Lessertia ingeliensis M. Balkwill Manulea florifera Hilliard & B.L. Burtt Miraglossum superbum Kupicha Moraea hiemalis Goldblatt Moraea unibracteata Goldblatt Ornithogalum diphyllum Baker Ornithogalum sephtonii Hilliard & B.L. Burtt Otholobium fumeum C.H. Stirt. Pachyacris sp. nov. rhodantha Stewart & Langley Pachycarpus campanulatus (Harv.) N.E.Br. var. campanulatus Pachycarpus natalensis N.E.Br. Passerina drakensbergensis Hilliard & B.L. Burtt Peucedanum thodei Arnold Protea dracomontana Beard Protea subvestita N.E.Br. Saniella verna Hilliard & B.L. Burtt Satureja grandibracteata Killick Schizochilus bulbinella (Rchb.f.) Bolus Schizoglossum elingue N.E.Br. subsp. elingue Schizoglossum stenoglossum Schltr. subsp. flavum (N.E.Br.) Kupicha Selago monticola J.M. Wood & M.S. Evans Selago trinervia E. Mey. Senecio basalticus Hilliard Senecio brevilorus Hilliard Senecio hirsutilobus Hilliard Senecio kalingenwae Hilliard & B.L. Burtt Senecio mauricei Hilliard & B.L. Burtt Senecio polelensis Hilliard Senecio praeteritus Killick Senecio thamathuensis Hilliard Strobilopsis wrightii Hilliard & B.L. Burtt Struthiola angustiloba Peterson & Hilliard Syncolostemon macranthus (Gu¨rke) M. Ashby Thesium decipiens Hilliard & B.L. Burtt Wahlenbergia pallidiflora Hilliard & B.L. Burtt Wahlenbergia polytrichifolia Schltr. subsp. dracomontana Hilliard & B.L. Burtt Xysmalobium tysonianum (Schltr.) N.E.Br. Xysmalobium woodii N.E.Br. Zaluzianskya chrysops Hilliard & B.L. Burtt

FData deficient_ (n=55) Agrostis subulifolia Stapf Berkheya griquana Hilliard & B.L. Burtt C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 111

Table 2A (continued) Red Data treatment by Talukdar (2002) and Victor (2002) Red Data treatment by Scott-Shaw (1999) Anisodontea julii (Burch. ex DC.) Berkheya leucaugeta Hilliard Bates subsp. prostrata (E. Mey. ex Turcz.) Bates Anthoxanthum brevifolium Stapf Craterocapsa insizwae (Zahlbr.) Hilliard & B.L. Burtt Aristida monticola Henrard Cyphia natalensis E. Phillips Asclepias xysmalobioides Hilliard & B.L. Burtt Erica flanaganii Bolus Bromus firmior (Nees) Stapf Erica wyliei Bolus Carex monotropa Nelmes Geranium angustipetalum Hilliard & B.L. Burtt Colpodium drakensbergense Hedberg & I. Hedberg Helichrysum album N.E.Br. Corycium alticola Parkman & Schelpe Hemizygia bolusii (N.E.Br.) Codd Cynoglossum alticola Hilliard & B.L. Burtt Hesperantha pubinervia Hilliard & B.L. Burtt Delosperma ashtonii L. Bolus Indigofera pseudoevansii Hilliard & B.L. Burtt Delosperma clavipes Lavis Lessertia dykei L. Bolus Delosperma nubigenum (Schltr.) L. Bolus Polygala praticola Chodat Dierama jucundum Hilliard Senecio telmateius Hilliard Disa galpinii Rolfe Schizoglossum quadridens N.E.Br. Disa sanguinea Sond. Schizoglossum singulare Kupicha Euryops evansii Schltr. subsp. dendroides B. Nord. Stachys rudatisii Skan Euryops inops B. Nord. Tetraria sp. nov. (Killick 1596 NU) Festuca dracomontana H.P. Linder Thesium cordatum A.W. Hill Festuca killickii Kenn.-O’Byrne Tulbaghia montana Vosa Gnidia singularis Hilliard Xerophyta longicaulis Hilliard Gymnopentzia bifurcata Benth. Hesperantha crocopsis Hilliard & B.L. Burtt Lessertia thodei L. Bolus Merwilla plumbea (Lindl.) Speta Merxmuellera aureocephala (J.G. Anderson) Conert Merxmuellera guillarmodiae Conert Pentaschistis praecox H.P. Linder Romulea luteoflora (M.P. de Vos) M.P. de Vos var. sanisensis M.P.de Vos Schizoglossum elingue N.E.Br. subsp. purpureum Kupicha Schizoglossum montanum R.A. Dyer Schoenoxiphium strictum Kukkonen Setaria obscura de Wit Wahlenbergia doleritica Hilliard & B.L. Burtt All treatments used the International Union for Conservation of Nature and Natural Resources criteria (IUCN, 1994).

Table 2B Red Data treatments for plant taxa forming part of the flora of southern Africa (FSA), which are either endemic or near-endemic to the Drakensberg Alpine Centre (DAC) Red Data treatment by Hilton-Taylor (1996a, b, 1997) Red Data treatment by Hall et al. (1980) FVulnerable_ (n=1) FUncertain_ (n=1) Kniphofia hirsuta Codd Cyrtanthus attenuatus R.A. Dyer FRare_ (n=1) Lotononis minor Dummer & Jenn. FNot threatened_ (n=11) Eucomis humilis Baker Eucomis schijffii Reyneke Galtonia viridiflora I. Verd. Gerbera parva N.E.Br. Gladiolus flanaganii Baker Gladiolus microcarpus G.J. Lewis Gladiolus oppositiflorus Herb. Kniphofia fibrosa Baker Kniphofia thodei Baker Rhus krebsiana C. Presl ex Engl. Watsonia gladioloides Schltr. FIndeterminate_ (n=1) Pachycarpus stenoglossus (E. Mey.) N.E.Br. FInsufficiently known_ (n=1) Phylica tysonii Pillans var. tysonii Hilton-Taylor’s (1996a, b, 1997) treatments were mostly based on earlier IUCN criteria (Davis et al., 1986). Criteria used by Hall et al. (1980) were not referenced. 112 Table 3 Recent additions to Victor and Keith’s (2004) Orange List Taxon Orange List Met criteria Locality Habitat requirements Conservation Major threat(s) History of threat Current Data category measures population source(s) trends Dicotyledonae Apiaceae Alepidea insculpta Rare (R) Occurrence <5000 km2, DAC Short subalpine Protected Incorrect fire Future threat ? Hilliard and Hilliard & B.L. Burtt <5 locations, southern grassland on high in UDP regime; soil Burtt (1985) KwaZulu-Natal basalt ridges erosion Drakensberg only (c. 2200 – 2600 m) Asteraceae Euryops evansii Schltr. Rare (R) Occurrence <5000 km2, DAC Alpine summit only Protected Incorrect fire Future threat ? Nordenstam

subsp. parvus B. Nord. <5 locations, Giant’s Castle (c. 2700 – 2900 m) in UDP regime (1968); 105–132 (2006) 72 Botany of Journal African South / Edwards T.J. Carbutt, C. and Cathedral Peak only Pooley (2003) Helichrysum inornatum Rare (R) Occurrence <5000 km2, DAC Damp or marshy Protected in Habitat loss in Future threat ? Hilliard (1983) Hilliard & B.L. Burtt <5 locations, Mpendhle, grassland UDP and Mpendhle and Underberg and Ngeli Mts. only (c. 1800 – 2100 m) by DWAF Ngeli; overgrazing Helichrysum Rare (R) Occurrence <5000 km2, DAC Damp grasslands Protected Alien invasives Current threat ? Hilliard (1983) mollifolium Hilliard <5 locations, and forest margins in UDP KwaZulu-Natal (c. 2100 – 2450 m) Drakensberg only Helichrysum Rare (R) Occurrence <5000 km2, DAC Rock pavements Protected Harvesting for Current and ? Hilliard (1983); pagophilum M.D. Hend. <5 locations, Cathedral and cliff faces of in UDP fire wood future threat Pooley (2003) Peak to Garden Castle alpine summit and Black Mountains (c. 2750 – 3400 m) (Lesotho) only Helichrysum paleatum Rare (R) Occurrence <5000 km2, DAC Short subalpine Protected Incorrect fire Future threat ? Hilliard (1983) Hilliard <5 locations, southern and grassland on stony in UDP regime; soil erosion central KwaZulu-Natal mountain slopes Drakensberg only (c. 1900 – 2500 m) Helichrysum Rare (R) Occurrence <5000 km2, DAC Damp rocky Protected Cropping in Lesotho; Current and ? Hilliard (1983) qathlambanum Hilliard <5 locations, Butha Buthe subalpine grassland in UDP habitat loss; trampling future threat (Lesotho), Mpendhle and and scrubby slopes by livestock Underberg only; highly of alpine summit disjunct range (c. 1800 – 3000 m) Helichrysum Rare – Sparse Occurrence <20,000 km2, DAC Rocky gullies, Protected Incorrect burning Past, current ? Hilliard (1983) tenuifolium Killick (RS) <10 locations, boulder beds, in UDP regime; soil erosion and future KwaZulu-Natal streamsides, subalpine Drakensberg only fynbos (>c. 1650 m) Inulanthera thodei Rare (R) Occurrence <5000 km2, DAC Rocky slopes, rocky ? Harvesting for firewood Past, current ? Pooley (2003) (Bolus) Ka¨llersjo¨ <5 locations, alpine gullies and alpine and fumigation of huts; and future summit of northern fynbos, on basalt soil erosion; fire Maloti Mts. only (?) (c. 2300 – 3200 m) Pentzia tortuosa Rare (R) Occurrence <5000 km2, DAC Rock sheets, scrubby ? Habitat loss; soil Current ? Harvey and (DC.) Fenzl ex Harv. <5 locations, Witteberge subalpine grassland erosion; fire; farming and future Sonder (1894); Mts. only (?) (c. 2100 – 2600 m) Pooley (2003) Senecio cristimontanus Rare – Critically Occurrence <100 km2, DAC Short dry stony Protected Infrequent burning; Future ? Hilliard (1977); Hilliard (RC) 1 location, Kamberg grassland and steep in UDP soil erosion Hilliard and Nature Reserve only, S-facing sandstone Burtt (1982) central KwaZulu-Natal ridges of Little ’Berg Drakensberg (c. 1980 – 2100 m) Senecio parentalis Rare (R) Occurrence <5000 km2, DAC Grows in damp Protected Habitat loss in Current ? Hilliard and Hilliard & B.L. Burtt <5 locations, Garden Castle marshes on Clarens in UDP and Ngeli (afforestation) and future Burtt (1976); Nature Reserve and Sandstone by DWAF Hilliard (1977) Ngeli Mt. only, southern (c. 1800 – 2150 m) KwaZulu-Natal Drakensberg Senecio qathlambanus Rare (R) Occurrence <5000 km2, DAC Damp subalpine and Protected Habitat loss in Current ? Hilliard and Hilliard <5 locations, Naude’s alpine grassland, on in UDP E. Cape; and future Burtt (1975); Nek (E. Cape Drakensberg) steep slopes and infrequent burning Hilliard (1977) to Giant’s Castle along watercourses (c. 2300 – 3100 m) Senecio submontanus Rare (R) Occurrence <5000 km2, DAC Damp grassland near Protected Infrequent burning; Past, current ? Hilliard and Hilliard & B.L. Burtt <5 locations, Garden Castle marshes on Clarens in UDP alien invasives and future Burtt (1976); and Bushman’s Nek Sandstone Hilliard (1977) only, southern KwaZulu-Natal (c. 1600 – 2400 m) Drakensberg Brassicaceae Heliophila alpina Rare (R) Occurrence <5000 km2, DAC Damp grassy and Protected Habitat loss in Current ? Marais (1966); Marais <5 locations, known only scree slopes of alpine in UDP E. Cape; trampling and future Pooley (2003) from Ben McDhui (E. Cape summit and wet silt by livestock; Drakensberg) to Thabana patches and turf on soil erosion 105–132 (2006) 72 Botany of Journal African South / Edwards T.J. Carbutt, C. Ntlenyana (Lesotho Malotis) summit plateau (c. 2800 – 3400 m) Campanulaceae Wahlenbergia lobulata Rare (R) Occurrence <5000 km2, DAC Short sloping Protected Habitat loss in Current ? Herbarium Brehmer <5 locations, recorded subalpine grassland in UDP E. Cape; and future records (NU) only from Ben McDhui and cracks in basalt infrequent fire (E. Cape Drakensberg), pavements on alpine Sani Pass/Top and Blue summit Mt. Pass (Lesotho) (c. 2600 – 3100 m) Dipsacaceae Cephalaria galpiniana Rare (R) Occurrence <5000 km2, DAC Short sloping subalpine ? Habitat loss in Current ? Herbarium Szabo´ subsp. galpiniana <5 locations, recorded only and alpine grasslands E. Cape; soil erosion; and future records (NU) from the Naude’s Nek and and gravel beds on trampling by livestock Tsatsane environs (E. summit Cape/Lesotho Drakensberg) (c. 2400 – 3000 m) Ericaceae Erica dracomontana Rare – Sparse Occurrence <20,000 DAC Alpine and subalpine Protected Frequent fires; Current ? Oliver (1985) E.G.H. Oliv. (RS) km2, <10 locations, known grasslands in UDP harvesting for and future from 7 locations in the (c. 2100 – 2500 m) fire wood northern KwaZulu-Natal Drakensberg, central Malotis and Witteberge (E. Cape) Erica trichoclada Rare (R) Occurrence <5000 km2, DAC Rocky grassland DWAF Habitat loss; Past, current ? Guthrie and Guthrie & Bolus <5 locations, Ngeli Mt. (with (c. 2200 m) frequent fires and future Bolus (1909), and Liddesdale only outlier) herbarium records (NU) Fabaceae Argyrolobium Rare (R) Occurrence <5000 km2,<5 DAC Short alpine turf Protected Cropping and Current ? Hilliard and summomontanum locations, known only from the northern (c. 2500 – 3300 m) in UDP livestock in Lesotho and future Burtt (1983) Hilliard & B.L. Burtt and central KwaZulu-Natal Drakensberg and northern Lesotho Malotis Lotononis jacottetii Rare (R) Occurrence <5000 km2,<5 DAC Grassland Protected Habitat loss in Current ? Van Wyk (1991); (Schinz) B.-E. van Wyk locations, known only from 2 (c. 1830 – 2010 m) in UDP Lesotho; and future Germishuizen and localities in the northern frequent fires Meyer (2003) KwaZulu-Natal Drakensberg and southern Lesotho Malotis Gentianaceae Sebaea pleurostigmatosa Rare (R) Occurrence <5000 km2,<5 DAC Alpine and Protected Infrequent fire Future ? Hilliard and Hilliard & B.L. Burtt locations, known only from the subalpine grassland in UDP Burtt (1983) northern and southern (c. 2300 – 2700 m) KwaZulu-Natal Drakensberg 113 (continued on next page) 114

Table 3 (continued) Taxon Orange List Met criteria Locality Habitat requirements Conservation Major threat(s) History of threat Current Data category measures population source(s) trends Rosaceae Cliffortia filicauloides Rare (R) Occurrence <5000 km2,<5 DAC Montane and sub-alpine Protected Invasive alien plants; Future ? Whitehouse (2004), Weim. locations, known only from the grassland; on wet rocks in UDP soil/riverbank erosion; herbarium records northern and central and damp earth banks incorrect burning (NU) KwaZulu-Natal Drakensberg along streams; regime occasionally Widdringtonia fynbos

(c. 1800 – 2300 m) 105–132 (2006) 72 Botany of Journal African South / Edwards T.J. Carbutt, C. Cliffortia spathulata Rare (R) Occurrence <5000 km2, DAC Montane and Protected Invasive alien plants; Current ? Whitehouse (2004) Weim. <5 locations, northern sub-alpine grassland, in UDP soil erosion; incorrect and future KwaZulu-Natal on Clarens Formation burning regime Drakensberg only sandstone or basalt, on well-drained soils; full sun (c. 1800 – 2750 m) Scrophulariaceae Diascia lilacina Rare – Critically Occurrence <100 km2,1 DAC Cliff-dwelling ? Rockfalls; instability Future ? Hilliard and Burtt Hilliard & B.L.Burtt (RC) location, known only from the (c. 2100 – 2200 m) of the pass (1983, 1984) summit of Saalboom Nek Pass, Eastern Cape Drakensberg Glumicalyx apiculatus Rare (R) Occurrence <5000 km2, DAC Wet basalt gravel beds ? Afforestation Future ? Hilliard (1994) (E. Mey.) Hilliard & <5 locations, known only and rock sheets in B.L. Burtt from a small portion of subalpine grasslands the Eastern Cape Drakensberg (c. 2200 – 2560 m) and Witteberge Limosella vesiculosa Rare – Sparse Occurrence <20,000 DAC Wet alpine turf Protected Trampling by livestock Current ? Hilliard and Hilliard & B.L. Burtt (RS) km2, <10 locations, recorded and marshy areas in UDP in Lesotho; and future Burtt (1986a); from 7 fairly disjunct localities near streams and tarns, prolonged drought Pooley (2003) on the KwaZulu-Natal mostly on summit Drakensberg escarpment, plateau Maloti Mts. and Eastern (c. 2000 – 3100 m) Cape Drakensberg Manulea dregei Rare (R) Occurrence <5000 km2,<5 DAC Damp silty basalt gravel ? Habitat loss in Current ? Hilliard Hilliard & B.L. Burtt locations, known only from a beds in alpine and E. Cape; infrequent and future (1994) small portion of the Eastern subalpine grasslands fire; afforestation Cape Drakensberg and (c. 2285 – 2835 m) Witteberge Manulea platystigma Rare (R) Occurrence <5000 km2,<5 DAC Silt patches over rock ? Trampling by Current ? Hilliard (1994); Hilliard & B.L. Burtt locations, known only from a sheets, loose scree and livestock; and future Pooley (2003) small portion of Eastern damp alpine turf on overgrazing; Lesotho summit plateau infrequent burning (c. 2900 – 3300 m) Nemesia glabriuscula Rare – Critically Occurrence <100 km2,1 DAC Boulder bed scrub Protected Flooding and Past, current ? Hilliard and Hilliard & B.L. Burtt (RC) location, known only from of the Mlambonja in UDP erosion; invasive and future Burtt (1986b) Garden Castle NR, southern River Valley exotic plants KwaZulu-Natal Drakensberg (c. 2060 – 2180 m) Selago leptothrix Hilliard Rare (R) Occurrence <5000 km2,<5 DAC Scrubby subalpine ? Habitat loss in Current ? Hilliard (1999) locations, known only from a grassland E. Cape; infrequent and future small portion of the Eastern (c. 2100 – 2500 m) fire; afforestation Cape Drakensberg Zaluzianskya turritella Rare (R) Occurrence <5000 km2, DAC Gravel and silt Protected Trampling and Current ? Hilliard (1994); Hilliard & B.L. Burtt <5 locations, known only beds overlying basalt in UDP overgrazing by and future Pooley (2003) from the central and rock sheets on alpine livestock southern KwaZulu-Natal summit plateau Drakensberg/Lesotho (c. 2940 – 3300 m) Escarpment Monocotyledonae Colchicaceae Wurmbea burttii Rare – Critically Occurrence <100 km2, DAC Wet silt and gravel Protected Trampling by Current ? Nordenstam B. Nord. (RC) 1 location, known only patches amongst rocks in UDP livestock and future (1978) from the Sani Pass/Sani in alpine and subalpine Top region of the southern grassland KwaZulu-Natal/Lesotho (c. 2400 – 2900 m) Drakensberg Wurmbea pusilla Rare (R) Occurrence <5000 km2, DAC Seasonally flooded Protected Trampling by Current ? Nordenstam 105–132 (2006) 72 Botany of Journal African South / Edwards T.J. Carbutt, C. E. Phillips <5 locations, recorded turf and marshlands or in UDP livestock and future (1978) from a few sites along gravel and silt patches the KwaZulu-Natal on summit plateau Drakensberg escarpment (c. 2870 – 3050 m) and Lesotho Maloti Mts. Wurmbea tenuis Rare – Critically Occurrence <100 km2, DAC Gravel and silt Protected Trampling by Current ? Nordenstam (Hook. f.) Baker subsp. (RC) 1 location, recorded only patches on summit in UDP and livestock and future (1978) australis B. Nord. from the Mont-aux-Sources plateau by FSDAE region of the northern (c. 2900 – 3000 m) KwaZulu-Natal/Lesotho/ Free State Drakensberg escarpment Cyperaceae Fuirena tenuis Rare (R) Occurrence <5000 km2, DAC Marshy areas in ? Afforestation; Current ? Forbes P.L. Forbes <5 locations, known only from montane grassland trampling by and future (1984) the Eastern Cape Drakensberg (c. 1800 m) livestock and Lesotho Maloti Mts. Iridaceae Hesperantha Rare (R) Occurrence <5000 km2, DAC Open stony ground Protected Infrequent fires Past, current ? Goldblatt altimontana Goldblatt <5 locations, recorded in alpine and subalpine in UDP and future (2003) from a few sites along grassland, often on the KwaZulu-Natal slopes Drakensberg escarpment (c. 2500 – 3200 m) and Lesotho Maloti Mts. Hesperantha brevistyla Rare (R) Occurrence <5000 km2, DAC Rock pavements and Protected Infrequent fires Past, current ? Goldblatt Goldblatt <5 locations, recorded only shallow wet ground in UDP and and future (2003) from the northern in alpine and subalpine by FSDAE KwaZulu-Natal/Lesotho/Free grassland, often on slopes State Drakensberg escarpment (c. 2200 – 3100 m) Hesperantha exiliflora Rare (R) Occurrence <5000 km2, DAC Marshy turf in alpine ? Infrequent fires; Past, current ? Goldblatt Goldblatt <5 locations, recorded only from and subalpine grassland trampling by and future (2003) a few Disjunct localities in the of summit plateau livestock Lesotho Maloti Mts. (c. 2500 – 2900 m) Romulea macowanii Rare (R) Occurrence <5000 km2, DAC Alpine grassland DWAF Afforestation; Current ? De Vos Baker var. alticola <5 locations, recorded (c. 2200 m) trampling by and future (1972) (B.L. Burtt) M.P. de Vos only from two sites: Ngeli livestock Mt. summit and the Lesotho Maloti Mts. (exact latter locality unknown) (continued on next page) 115 116 .Crut ..Ewrs/SuhArcnJunlo oay7 20)105–132 (2006) 72 Botany of Journal African South / Edwards T.J. Carbutt, C.

Table 3 (continued) Taxon Orange List Met criteria Locality Habitat requirements Conservation Major threat(s) History of threat Current Data category measures population source(s) trends Orchidaceae Disa dracomontana Rare (R) Occurrence <5000 km2, DAC Steep alpine and Protected Infrequent burning; Current ? Linder and Schelpe ex H.P. Linder <5 locations, northern and subalpine grasslands, in UDP orchid collectors and future Kurzweil (1999) central KwaZulu-Natal often in rocky ground Drakensberg only (c. 2100 – 2900 m) Disa nivea H.P. Linder Rare (R) Occurrence <5000 km2, DAC Rocky montane and Protected Afforestation; Current ? Linder and 5 locations, recorded only from subalpine grassland, in UDP trampling by and future Kurzweil (1999), a few sites along the southern sometimes Protea livestock; orchid personal KwaZulu-Natal/Lesotho savanna, mostly on collectors observations Drakensberg Clarens Sandstone (c. 2250 – 2500 m) Poaceae Festuca vulpioides Rare (R) to Occurrence <5000 km2, DAC Steep montane and ? Afforestation Current ? Gibbs Russell Steud. Rare – Critically <5 locations, Eastern subalpine grassland and future et al. (1990); (RC) (?) Cape Drakensberg only (c. 1800 – 2000 m) Germishuizen and Meyer (2003) All taxa listed beneath are, with the exception of the near-endemic Erica trichoclada Guthrie and Bolus, angiosperms endemic to the Drakensberg Alpine Centre (DAC). Abbreviations for conservation bodies: DWAF=Department of Water Affairs and Forestry; FSDAE=Free State Department of Agriculture and Environment; UDP=uKhahlamba-Drakensberg Park (managed by Ezemvelo KZN Wildlife). C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 117

Table 4 Synopsis of the endemic and near-endemic angiosperm taxa of the Drakensberg Alpine Centre (DAC) Endemic angiosperms Near-endemic angiosperms Families Genera Species Families Genera Species No. % No. % No. % No. % No. % No. % Dicotyledons 26 63.4 72 60.5 243 72.8 40 71.4 128 68.1 412 69.2 Monocotyledons 15 36.6 47 39.5 91 27.2 16 28.6 60 31.9 183 30.8 Totals 41 100 119 100 334 100 56 100 188 100 595 100

Taylor, 1994; Cowling and Hilton-Taylor, 1997), viz. bior- 3.4. Largest families and genera egions that are characterised by high species richness, high levels of endemism and high levels of man-induced habitat The 334 endemics belong to 41 families and 119 genera transformation (Myers, 1988). Its compliment of 2520 native (mean=2.8 species per genus) (Table 4). The 595 near- angiosperms (Carbutt and Edwards, 2004) is comparable with endemics belong to 56 families and 188 genera (mean=3.2 the native floras of Northern Hemisphere countries such as species per genus) (Table 4). Although the majority of the Germany or Switzerland, and is almost double that of Great endemics and near-endemics are dicotyledons (Table 4), this Britain (Hawksworth and Kalin-Arroyo, 1995). This total is dominance is not as marked as that reported for the FSA (see also comparable with the diversity recorded in the centres of Germishuizen and Meyer, 2003). the world-renowned Succulent Karoo Region (Gariep, Little The 13 families, containing 2% or more endemic species Karoo and Hantam-Roggeveld Centres) (refer to Van Wyk and each, collectively contribute c. 80% of the endemic species and Smith, 2001). c. 69% of the genera that have one or more endemic species Some 334 angiosperm taxa (c. 13%) are endemic to the (Table 5). The three largest families, Asteraceae, Scrophular- DAC (Table 4 and Appendix 1) and a further 595 taxa (c. 24%) iaceae and Iridaceae, contribute almost 50% of the endemic are near-endemic (Table 4 and Appendix 2), most (c. 99%) of species (Table 5). The 14 families, containing 2% or more near- which do not extend beyond the Limpopo River. Thus, 37% of endemic species each, collectively contribute c. 77% of the the DAC’s flora is confined to the Eastern Region of southern near-endemic species and c. 62% of the genera that have one or Africa. more near-endemic species (Table 6). The five largest families, Species endemism in the DAC (c. 13%) is slightly lower Asteraceae, Scrophulariaceae, Iridaceae, Orchidaceae and than that for KwaZulu-Natal (c. 16%, Scott-Shaw, 1999), and Fabaceae, contribute almost 50% of the near-endemic species significantly lower than the CFR (c. 69%, Goldblatt and (Table 6). Manning, 2002) (see Fig. 4 and Table 9). It is, however, Asteraceae contributes the most species to the flora of the misleading to compare a centre of endemism with a wider- DAC, as well as the most endemics and near-endemics (Fig. 2). ranging floristic region of higher rank. It would have been Poaceae contributes few endemics and near-endemics, relative desirable to isolate the high-altitude flora of the CFR from its to its dominance in the flora (Fig. 2), probably because of its lowland flora, and compare this with the DAC. Distinguishing continuity within the Grassland Biome. Scrophulariaceae and these Fhigh-altitude_ species is problematic when dealing with Iridaceae contribute a high proportion of endemics and near- temperate vegetation in the CFR (P. Linder personal endemics relative to their contribution to the flora (Fig. 2). communication), particularly when very little work has Iridaceae also features prominently as a contributor of edaphic focussed on the CFR’s high-altitude flora (see Linder et al., endemics to the Wolkberg Centre of endemism located in the 1993). north-eastern Drakensberg Escarpment (Matthews et al., 1993).

Table 5 Ranking of the larger families that contribute 2% or more endemic species to the angiosperm flora of the Drakensberg Alpine Centre, together with the number of genera contributing endemic species Rank Family Endemic species % endemic No. of genera % genera Species/genus 1 Asteraceae 103 30.8 23 19.3 4.5 2 Scrophulariaceae 37 11.1 10 8.4 3.7 3 Iridaceae 23 6.9 6 5.0 3.8 4 Apocynaceae 13 3.9 4 3.4 3.3 4 Cyperaceae 13 3.9 6 5.0 2.2 5 Ericaceae 12 3.6 1 0.8 12.0 5 Mesembryanthemaceae 12 3.6 1 0.8 12.0 5 Poaceae 12 3.6 9 7.6 1.3 6 Orchidaceae 11 3.3 6 5.0 1.8 7 Fabaceae 8 2.4 4 3.4 2.0 7 Hyacinthaceae 8 2.4 6 5.0 1.3 8 Asphodelaceae 7 2.1 3 2.5 2.3 8 Thymelaeaceae 7 2.1 3 2.5 2.3 A=266 A=79.7% A=82 A=68.7% 118 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

Table 6 Ranking of the larger families that contribute 2% or more near-endemic (N-E) species to the angiosperm flora of the Drakensberg Alpine Centre, together with the number of genera contributing near-endemic species Rank Family N-E species % N-E No. of genera % genera Species/genus 1 Asteraceae 135 22.7 29 15.4 4.7 2 Scrophulariaceae 44 7.4 13 6.9 3.4 3 Iridaceae 42 7.1 8 4.3 5.3 4 Orchidaceae 41 6.9 11 5.9 3.7 5 Fabaceae 33 5.5 14 7.4 2.4 6 Apocynaceae 30 5.0 10 5.3 3.0 7 Poaceae 27 4.5 13 6.9 2.1 8 Campanulaceae 18 3.0 2 1.1 9.0 9 Asphodelaceae 17 2.9 3 1.6 5.7 10 Geraniaceae 16 2.7 3 1.6 5.3 11 Amaryllidaceae 14 2.4 3 1.6 4.7 12 Hypoxidaceae 13 2.2 3 1.6 4.3 12 Lamiaceae 13 2.2 4 2.1 3.3 13 Ericaceae 12 2.0 1 0.5 12.0 A=455 A=76.5% A=117 A=62.2%

Fabaceae and Orchidaceae contribute substantially more near- species occur above the line of best fit; they are truly endemics than endemics, while Cyperaceae contributes more remarkable in their endemic representation (Fig. 3). Most of endemics than near-endemics (Fig. 2). Species/genus ratios in these genera are temperate, montane/alpine specialists, char- the endemics only twice exceed 10 (Ericaceae and Mesem- acterised by localised seed dispersal, and favour a range of bryanthemaceae), due to the large genera Erica and Delosperma lithophilic habitats (rock outcrops, basalt gravel beds, wet drip (Tables 5 and 7), and in the near-endemics only once exceeds 10 faces, or scree banks). Other possible reasons for their high (Ericaceae), due to the large genus Erica (Tables 6 and 8). degree of in situ speciation are: interrupted habitats leading to a The genera Helichrysum and Senecio contribute the most breakdown in gene flow; intolerance to warm temperatures species to the flora of the DAC, as well as the most endemics (altitudinal endemics); and a high capacity to compete with (Fig. 3A) and near-endemics (Table 8). Helichrysum is also a tropical species in cooler environments. Those genera contrib- significant contributor of endemics to the flora of the Wolkberg uting endemic species, occurring below the line of best fit, tend Centre of endemism (Matthews et al., 1993). Delosperma, more towards having tropical affinities, or are wider-ranging in Erica, Hesperantha and Diascia also contribute many ende- southern Africa than those genera contributing endemics above mics (Fig. 3B). Exactly half the genera contributing endemic the line of best fit (Fig. 3). The 12 larger genera contributing

500 total flora near-endemics endemics 400

300

No. of spp. 200

100

0 Poaceae Scrophulariaceae Cyperaceae Asteraceae Fabaceae Orchidaceae Iridaceae Family

Fig. 2. The larger angiosperm families of the Drakensberg Alpine Centre (DAC), representing their contribution to the angiosperm flora, as well as their contribution of endemics and near-endemics. Flora totals were derived from Carbutt and Edwards (2004). C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 119

Table 7 Ranking of the larger genera that contribute six or more endemic species to the angiosperm flora of the Drakensberg Alpine Centre Rank Genus Family D=dicotyledon/M=monocotyledon Endemic species % endemic 1 Helichrysum Asteraceae D 29 8.7 2 Senecio Asteraceae D 22 6.6 3 Delosperma Mesembryanthemaceae D 12 3.6 3 Erica Ericaceae D 12 3.6 4 Hesperantha Iridaceae M 9 2.7 5 Diascia Scrophulariaceae D 8 2.4 6 Euryops Asteraceae D 7 2.1 6 Schizoglossum Apocynaceae D 7 2.1 7 Disa Orchidaceae M 6 1.8 7 Glumicalyx Scrophulariaceae D 6 1.8 7 Sebaea Gentianaceae D 6 1.8 7 Selago Scrophulariaceae D 6 1.8 A=130 A=39.0% six or more endemic species each, collectively account for 39% contribute endemic species and 45 (c. 63%) contribute near- of the species (Table 7), while a further 12 genera contribute endemic species. Few woody angiosperms, particularly trees, five endemic species each (Appendix 1). are endemic or near-endemic to the DAC. The 29 larger genera contributing six or more near-endemic Five genera are endemic to the DAC: Dracomonticola species each, collectively account for 51% of the species (Table (Orchidaceae), Glumicalyx (Scrophulariaceae), Heteromma 8), while a further five genera contribute five near-endemic (Asteraceae), Polevansia (Poaceae) and Strobilopsis (Scrophu- species each (Appendix 2). After Helichrysum and Senecio, the lariaceae), and six genera are near-endemic: Craterocapsa greatest numbers of near-endemics are contributed by Wahlen- (Campanulaceae), Glekia (Scrophulariaceae), Guthriea (Achar- bergia, Disa, Kniphofia and Erica (Table 8). iaceae), Huttonaea (Orchidaceae), Rhodohypoxis (Hypoxida- Of the 72 genera recognised as Cape elements in the DAC ceae) and Thamnocalamus (Poaceae) (Carbutt and Edwards, (Carbutt and Edwards, 2001; Carbutt, 2004), 39 (c. 54%) 2004). There are no endemic families.

Table 8 Ranking of the larger genera that contribute six or more near-endemic (N-E) species to the angiosperm flora of the Drakensberg Alpine Centre Rank Genus Family D=dicotyledon/M=monocotyledon N-E species % N-E 1 Helichrysum Asteraceae D 43 7.2 2 Senecio Asteraceae D 29 4.9 3 Wahlenbergia Campanulaceae D 14 2.4 4 Disa Orchidaceae M 13 2.2 4 Kniphofia Asphodelaceae M 13 2.2 5 Erica Ericaceae D 12 2.0 6 Berkheya Asteraceae D 10 1.7 6 Delosperma Mesembryanthemaceae D 10 1.7 6 Dierama Iridaceae M 10 1.7 6 Lotononis Fabaceae D 10 1.7 6 Stachys Lamiaceae D 10 1.7 7 Moraea Iridaceae M 9 1.5 8 Crassula Crassulaceae D 8 1.3 8 Disperis Orchidaceae M 8 1.3 8 Hesperantha Iridaceae M 8 1.3 8 Pachycarpus Apocynaceae D 8 1.3 8 Pentaschistis Poaceae M 8 1.3 8 Selago Scrophulariaceae D 8 1.3 9 Alepidea Apiaceae D 7 1.2 9 Cineraria Asteraceae D 7 1.2 9 Cyrtanthus Amaryllidaceae M 7 1.2 9 Euryops Asteraceae D 7 1.2 9 Hypoxis Hypoxidaceae M 7 1.2 9 Pelargonium Geraniaceae D 7 1.2 9 Zaluzianskya Scrophulariaceae D 7 1.2 10 Cyphia Lobeliaceae D 6 1.0 10 Geranium Geraniaceae D 6 1.0 10 Rhus Anacardiaceae D 6 1.0 10 Schizoglossum Apocynaceae D 6 1.0 A=304 A=51.1% 120 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

A. 6

y = 0.39x + 0.83 Helichrysum r2 = 0.77 p < 0.001 5 Senecio

4

3 no. of endemics in DAC 10

Log 2

1 0246810

Log10 total no. of spp. in DAC B. 3.5 Delosperma Erica

Hesperantha 3.0 Diascia

Schizoglossum Euryops

2.5 Glumicalyx Selago Sebaea Disa

Macowania S F Gn B/Mo Wahlenbergia

C J G K Thesium no. of endemics in DAC Zaluzianskya 10 Rhodohypoxis 2.0 Log E/H M Cl Ca Asclepias L/W A Al

1.5 123456

Log10 total no. of spp. in DAC

Fig. 3. Double-log plots of the genera contributing three or more species endemic to the Drakensberg Alpine Centre (DAC) relative to their total contribution to the flora. (A) Plotting of all genera that contribute three or more endemic species. (B) Axes scaled down to exclude Helichrysum and Senecio, in order to view in detail the encircled data pool in panel A. Abbreviations for genera: A=Aster;Al=Alepidea;B=Berkheya;C=Cotula;Ca=Carex;Cl=Cliffortia;E=Eumorphia; F=Felicia;G=Gladiolus;Gn=Gnidia;H=Heteromma;J=Jamesbrittenia;K=Kniphofia;L=Lessertia;M=Manulea;Mo=Moraea;S=Schoenoxiphium; W=Wurmbea.

3.5. Problematic taxa DAC. Lessertia glabricaulis L. Bolus (Fabaceae), apparently recorded from the Lesotho Highlands and a probable DAC These can be divided into the following categories: endemic, was also excluded for the present time due to insufficient data. 3.5.1. Poorly known taxa Several taxa were difficult to place. Rhynchosia dieterlenae 3.5.2. Highly disjunct taxa Baker f. (Fabaceae), for example, is known from one gathering Certain taxa, although strictly confined to the DAC in a from the Leribe environ in Lesotho (Baker, 1923) (c. 1830 m?). southern African context, were found to extend further north It was excluded because, although meeting the altitudinal into tropical Africa, usually in Afromontane environments, requirement, appears to occur just beyond the boundary of the following consultation of tropical flora accounts. Agrocharis C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 121

4 combination, Phacelurus franksae (J.M. Wood) Clayton. y = 0.64x – 0.14 CFR r2 = 0.72 Following this more recent revision, P. franksae is not endemic p < 0.01 to the DAC. Van Wyk and Smith (2001) erroneously cited this taxon in their endemic/near-endemic list for the DAC. 3 KZN EMR 3.6. Comparison with other floras and the Farea effect_ DAC A comparison with other high-altitude floras of southern

endemic species 2 Mulanje and south-central Africa reveals that endemism in the DAC, 10 Chimani when correlated with its geographical area, is sub-average

Log (below the regression line) (Fig. 4). The linear regression Nyika represents the expected species–area correlation for a flora 1 relative to its geographical area. The endemics recorded for the 23456Chimanimani Mts. (Zimbabwe) and Mt. Mulanje (Malawi), 2 Log10 area (km ) although relatively few in number (Table 9), are plotted as above-average once geographical area is taken into account Fig. 4. Double-log plots of the endemic species–area relationships for a (Fig. 4). The position of the DAC plot was at first thought to be F _ number of floras of southern and south-central Africa. Species on the y-axis skewed by the absence of a large number of near-endemics (c. refers to all vascular plant taxa. Abbreviations: CFR=Cape Floristic Region; Chimani=Chimanimani; DAC=Drakensberg Alpine Centre; EMR=Eastern 125 species) that extend slightly below its 1800 m cut-off. Mountain Region; KZN=KwaZulu-Natal. However, the inclusion of the Eastern Mountain Region endemics (this account recognises 459 species occurring melanantha Hochst. (Apiaceae), for example, although con- 1500 m), relative to the Eastern Mountain Region’s fined to the DAC in southern Africa, was first described from geographical area of 52,074 km2, also resulted in a sub- the Semien Mts. in Ethiopia, and has also been recorded in average plot of endemism (Fig. 4). Yemen and East Africa (Burtt, 1991). At a regional centre-level, endemism in the DAC ranks similarly with centres of endemism in the Succulent Karoo 3.5.3. Taxonomically labile taxa Region (Knersvlakte, Little Karoo and Hantam-Roggeveld Ischaemum franksae J.M. Wood (Poaceae), described from Centres), and is double the level of endemism recorded for the the DAC in 1908 and thought to be endemic, was sunk 26 Pondoland Centre (see Van Wyk and Smith, 2001). The totals years later into Ischaemum juncifolium Ballard and Hubbard, supplied by Van Wyk and Smith (2001), however, are slightly from tropical Africa (Clayton, 1978), resulting in the new inflated due to the inclusion of near-endemics. Such compar-

Table 9 A comparison of the number of endemic vascular plants for regions in southern and south-central Africa Region CPD Locality Maximum altitude No. of vascular Vascular species Physical Species/103 Endemic site (m a.s.l.) species endemism area km2 species/103 [(no.) %] (km2) km2 Cape Floristic Af53 Western Cape, Matroosberg 9030 (6204) 68.7 90,000 100.3 68.9 Regiona South Africa (2249) KwaZulu-Natalb – Eastern Region, Njesuthi (3410) 6000 (960) 16.0 92,300 65.0 10.4 South Africa Chimanimani Af79 Eastern Highlands, Mt. Binga (2440) 859 (1200 m) (41) 4.8 171 5023.4 239.8 Subcentrec Zimbabwe Nyanga Subcentred Af80 Eastern Highlands, Mt. Inyangani ?(2000 m) ? 289 ? ? Zimbabwe (2593) Chimanimani-Nyanga – Eastern Highlands, Mt. Inyangani >1500 (1200 m) (>100) 6.7 ? ? ? Centree Zimbabwe (2593) Mt. Mulanjef Af64 Southern Malawi Sapitwa (3001) 1303 (750 m) (69) 5.3 640 2036 107.8 Nyika Plateaug Af65 Northern Malawi Nganda Hill (2607) T1900 (1800 m) (21) 1.1 1800 1056.0 11.7 DACh (this study) Af82 Eastern South African Thabana Ntlenyana 2618 (1800 m) (>334) T13.0 40,000 65.0 8.4 interior and Lesotho (3482) Regions range from floristic regions, political provinces, centres and subcentres of endemism, to centres of plant diversity. a Following Goldblatt and Manning (2002) and Linder (2003). b Following Scott-Shaw (1999). c Following Goodier and Phipps (1961) and Wild (1964). d Following WWF and IUCN (1994). e Following Van Wyk and Smith (2001). f Following Strugnell (2002). g Following Willis et al. (2001). h DAC is synonymous with the outdated FDrakensberg Alpine Region_ of Killick (1994). 122 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

Table 10 to note that these shared ranges of distribution are not as yet Examples of DAC near-endemics grouped according to their similar referred to as phytogeographical groups, as their phylogenetic distribution ranges histories are unknown and have therefore not been incorporated Group 1 into the analysis. Taxa in Group 1 extend just beyond the DAC, Dierama dracomontanum Hilliard Hermannia oligosperma K. Schum. usually not beyond the KwaZulu-Natal Midlands (1400 m). Hesperantha grandiflora G.J. Lewis Taxa in Group 2 have a montane distribution, following the Jamesbrittenia pristisepala (Hiern) Hilliard Drakensberg Range to the north and south of the DAC. Their Setaria obscura de Wit southern-most station is the Amatole Mts., more rarely the Sneeuberge Range near Graaff-Reinet. Their northern-most Group 2 Aristea montana Baker station is the Mpumalanga Drakensberg (i.e. Graskop, Mar- Asparagus microraphis (Kunth) Baker iepskop, Wolkberg, Woodbush), more rarely the Soutpansberg. Disperis renibractea Schltr. Their lower altitudinal limits rarely extend below 1200 m. Taxa Disperis stenoplectron Rchb.f. in Group 3 have a station in the DAC (mostly the KwaZulu- Eriocaulon hydrophilum Marko¨tter Natal Drakensberg, more rarely the Eastern Cape Drakens- Helichrysum monticola Hilliard F _ Helichrysum oreophilum Klatt berg), with outliers confined almost exclusively to KwaZulu- Neobolusia tysonii (Bolus) Schltr. Natal (KZN), more rarely the Eastern Cape. These FKZN_ Plectranthus grallatus Briq. (åEastern Region) endemics may extend towards the coast, Rhus montana Diels however their confinement to KZN, with a stronghold in the DAC, renders them near-endemic to the DAC. Taxa in Group 4 Group 3 Adhatoda andromeda (Lindau) C.B. Clarke occur mostly on the Lesotho Plateau, the uplands of northern Anemone fanninii Harv. ex Mast. KwaZulu-Natal, Free State and the Transvaal Highveld Berkheya macrocephala J.M. Wood (Mpumalanga and Gauteng), and the northeastern Cape Cineraria dieterlenii E. Philiips interior. They seemingly have more xeric preferences. None Dierama latifolium N.E.Br. of the taxa in the above groups extend beyond the Limpopo Helichrysum natalitium DC. Hoffmannseggia sandersonii (Harv.) Engl. River; moreover they are mostly restricted to the Eastern Hypoxis lata Nel Region. Taxa in Group 5 have outliers in the Eastern Highlands Indigofera foliosa E. Mey. Inulanthera leucoclada (DC.) Ka¨llersjo¨ Moraea hiemalis Goldblatt Nemesia silvatica Hilliard Pelargonium bowkeri Harv. Schizoglossum flavum Schltr. Senecio heliopsis Hilliard & B.L. Burtt Tulbaghia natalensis Baker

Group 4 Asparagus concinnus (Baker) Kies Empodium elongatum (Nel) B.L. Burtt Helichrysum dasycephalum O. Hoffm. Helichrysum melanacme DC. Rhus divaricata Eckl. & Zeyh. Stachys hyssopoides Burch. ex Benth.

Group 5 Crassula tenuicaulis Scho¨nland Craterocapsa tarsodes Hilliard & B.L. Burtt Dierama pauciflorum N.E.Br. Disperis concinna Schltr. Protea dracomontana Beard Wurmbea angustifolia B. Nord. The extent of each group’s range is outlined in the text. isons are also somewhat artificial as they fail to take geographical area into account.

3.7. Shared distribution ranges Fig. 5. The shared distribution ranges of five groups of angiosperms near- endemic to the Drakensberg Alpine Centre (DAC): 1=Falmost endemic_; In general terms, the near-endemic taxa of the DAC can be 2=montane; 3=KwaZulu-Natal endemics; 4=semi-arid interior; 5=eastern assigned to five groups, with the taxa of each group displaying highlands of Zimbabwe. Abbreviations: L=Lesotho; M=Mozambique; similar distribution ranges (Table 10 and Fig. 5). It is important S=Swaziland. The DAC is depicted as the area shaded in black. C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 123 of Zimbabwe, i.e. the Chimanimani-Nyanga Centre of Van Appendix A Wyk and Smith (2001). All the above groups were arrived at by inspection of the The endemic angiosperms of the Drakensberg Alpine Centre individual ranges of their constituent taxa. These five groups Angiospermae–Dicotyledonae differ somewhat from the 13 groups identified by Hilliard and Apiaceae (3/5) Burtt (1987) for the flora of the southern KwaZulu-Natal Alepidea insculpta Hilliard & B.L. Burtt Drakensberg, as the latter analysis included a number of wide- Alepidea pusilla Weim. ranging species, and the former only the near-endemics of the Alepidea thodei Dummer DAC. Groups 1 to 6 of Hilliard and Burtt (1987), however, Conium fontanum Hilliard & B.L. Burtt var. alticola Hilliard & B.L. Burtt Dracosciadium saniculifolium Hilliard & B.L. Burtt dealing with the less wide-ranging species, do comply with Apocynaceae (4/13) roughly the five groups identified in this analysis. Asclepias humilis (E. Mey.) Schltr. Asclepias oreophila Nicholas 4. Conclusions Asclepias xysmalobioides Hilliard & B.L. Burtt, not of S. Moore Brachystelma alpinum R.A. Dyer Brachystelma perditum R.A. Dyer The DAC is here shown to have a 13% endemic element, Miraglossum superbum Kupicha and at least a 24% near-endemic element; the latter being Schizoglossum elingue N.E.Br. subsp. elingue termed the FEastern Region_ element in a southern African Schizoglossum elingue N.E.Br. subsp. purpureum Kupicha context. The many endemics and near-endemics recognised in Schizoglossum hilliardiae Kupicha this account support the World Heritage Convention’s enrol- Schizoglossum montanum R.A. Dyer F _ Schizoglossum quadridens N.E.Br. ment of part of the DAC as a World Heritage site. This profile Schizoglossum singulare Kupicha of angiosperm endemism and near-endemism needs to be Schizoglossum stenoglossum Schltr. subsp. flavum (N.E.Br.) Kupicha followed up by detailed autecological studies of individual Asteraceae (23/103) species, knowledge of which should help fine-tune manage- Aster ananthocladus Hilliard & B.L. Burtt ment practises (e.g. burning regime) that aim to optimise Aster confertifolius Hilliard & B.L. Burtt Aster erucifolius (Thell.) Lippert biodiversity. Athanasia grandiceps Hilliard & B.L. Burtt Detailed phytogeographic studies of the DAC’s near- Berkheya cirsiifolia (DC.) Roessler endemic taxa will provide further clues into its origins and Berkheya draco Roessler connections with other floras. The outcome will be a sound Berkheya leucaugeta Hilliard breakdown of its phytogeographic elements as undertaken by Berkheya pannosa Hilliard Berkheya rosulata Roessler Hedberg (1961, 1965) for the Afroalpine flora of north-east Cineraria albomontana Hilliard Africa. Comborhiza virgata (N.E.Br.) Anderb. & K. Bremer The flora of the DAC must continue to receive conser- Cotula lineariloba (DC.) Hilliard vation attention because many of its taxa are rare and highly Cotula membranifolia Hilliard restricted in distribution, and more than half of its endemics Cotula paludosa Hilliard Cotula radicalis (Killick & C. Claassen) Hilliard & B.L. Burtt are regarded as either Red or Orange Data species. This Cotula socialis Hilliard statistic far exceeds the percentage of rare and threatened Eumorphia prostrata Bolus taxa recorded for many other floras, both local and abroad. Eumorphia sericea J.M. Wood & M.S. Evans subsp. robustior Hilliard & Many of the DAC’s endemics are afforded formal protection B.L. Burtt only in the uKhahlamba-Drakensberg Park (KwaZulu-Natal) Eumorphia sericea J.M. Wood & M.S. Evans subsp. sericea Euryops acraeus M.D. Hend. and in the Bokong, Ts’ehlanyane and Sehlabathebe National Euryops brevipes B. Nord. Parks (Lesotho). A strong plea is made for the continued Euryops decumbens B. Nord. sustainable management and protection of biota in the DAC, Euryops evansii Schltr. subsp. dendroides B. Nord. as well as the evolutionary processes that have led to their Euryops evansii Schltr. subsp. parvus B. Nord. speciation, in order to mitigate any threats that may lead to Euryops inops B. Nord. Euryops montanus Schltr. their demise. Felicia caespitosa Grau Felicia drakensbergensis J.M. Wood & M.S. Evans Acknowledgements Felicia linearis N.E.Br. Felicia uliginosa (J.M. Wood & M.S. Evans) Grau We thank Ashley Nicholas, Bertil Nordenstam, Kathleen Felicia wrightii Hilliard & B.L. Burtt Garuleum sp. nov. Gordon-Gray, and Angela Beaumont for assistance with Gerbera parva N.E.Br. Apocynaceae, Colchicaceae, Cyperaceae and Thymelaeaceae Gnaphalium limicola Hilliard respectively. Janine Victor (Threatened Species Programme, Helichrysum album N.E.Br. Pretoria) is thanked for assistance with the Orange List. John Helichrysum amplectens Hilliard Manning is thanked for his help in improving the manuscript. Helichrysum basalticum Hilliard Helichrysum bellum Hilliard Funding from the University of KwaZulu-Natal Research Fund Helichrysum confertum N.E.Br. and the Maloti-Drakensberg Transfrontier Project is gratefully Helichrysum drakensbergense Killick acknowledged. Helichrysum evansii Hilliard 124 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

Helichrysum flanaganii Bolus Campanulaceae (1/5) Helichrysum glaciale Hilliard Wahlenbergia doleritica Hilliard & B.L. Burtt Helichrysum haygarthii Bolus Wahlenbergia lobulata Brehmer Helichrysum heterolasium Hilliard Wahlenbergia polytrichifolia Schltr. subsp. dracomontana Hilliard & B.L. Helichrysum hyphocephalum Hilliard Burtt Helichrysum inornatum Hilliard & B.L. Burtt Wahlenbergia pulvillus-gigantis Hilliard & B.L. Burtt Helichrysum lineatum Bolus Wahlenbergia tetramera Thulin Helichrysum longinquum Hilliard Crassulaceae (1/1) Helichrysum marginatum DC. Crassula qoatlhambensis Hargr. Helichrysum milfordiae Killick Dipsacaceae (1/2) Helichrysum mollifolium Hilliard Cephalaria galpiniana Szabo´ subsp. galpiniana Helichrysum nimbicola Hilliard Cephalaria galpiniana Szabo´ subsp. simplicior B.L. Burtt Helichrysum pagophilum M.D. Hend. Ericaceae (1/12) Helichrysum paleatum Hilliard Erica aestiva Marko¨tter var. aestiva Helichrysum palustre Hilliard Erica albospicata Hilliard & B.L. Burtt (25) Helichrysum praecurrens Hilliard Erica anomala Hilliard & B.L. Burtt (25) Helichrysum qathlambanum Hilliard Erica dissimulans Hilliard & B.L. Burtt (25) Helichrysum retortoides N.E.Br. Erica dominans Killick (6) Helichrysum sessilioides Hilliard Erica dracomontana E.G.H. Oliv. Helichrysum subfalcatum Hilliard Erica ebracteata Bolus Helichrysum tenuifolium Killick Erica flanaganii Bolus Helichrysum witbergense Bolus Erica frigida Bolus Heteromma decurrens (DC.) O. Hoffm. Erica thodei Guthrie & Bolus Heteromma krookii (O. Hoffm. & Muschl.) Hilliard & B.L. Burtt Erica tysonii Bolus var. tysonii Heteromma simplicifolium J.M. Wood & M.S. Evans Erica wyliei Bolus Inulanthera thodei (Bolus) Ka¨llersjo¨ Euphorbiaceae (1/2) Macowania corymbosa M.D. Hend. Clutia alpina Prain Macowania deflexa Hilliard & B.L. Burtt Clutia nana Prain Macowania glandulosa N.E.Br. Fabaceae (4/8) Macowania hamata Hilliard & B.L. Burtt Argyrolobium summomontanum Hilliard & B.L. Burtt Macowania sororis Compton Indigofera evansii Schltr. Osteospermum attenuatum Hilliard & B.L. Burtt Indigofera pseudoevansii Hilliard & B.L. Burtt Osteospermum thodei Marko¨tter Lessertia dykei L. Bolus Othonna burttii B. Nord. Lessertia harveyana L. Bolus Othonna sp. nov. Lessertia ingeliensis M. Balkwill Pentzia tortuosa (DC.) Fenzl ex Harv. Lotononis jacottetii (Schinz) B.-E. van Wyk Printzia nutans (Bolus) Leins Lotononis minor Dummer & Jenn. Relhania acerosa (DC.) K. Bremer Gentianaceae (1/6) Relhania dieterlenii (E. Phillips) K. Bremer Sebaea marlothii Gilg Senecio austromontanus Hilliard Sebaea minutissima Hilliard & B.L. Burtt Senecio basalticus Hilliard Sebaea pleurostigmatosa Hilliard & B.L. Burtt Senecio brevilorus Hilliard Sebaea radiata Hilliard & B.L. Burtt Senecio caloneotes Hilliard Sebaea spathulata (E. Mey.) Steud. Senecio cristimontanus Hilliard Sebaea thodeana Gilg Senecio cryptolanatus Killick Geraniaceae (1/2) Senecio dissimulans Hilliard Geranium angustipetalum Hilliard & B.L. Burtt Senecio ingeliensis Hilliard Geranium drakensbergensis Hilliard & B.L. Burtt Senecio kalingenwae Hilliard & B.L. Burtt Lamiaceae (4/6) Senecio marginalis Hilliard Hemizygia bolusii (N.E.Br.) Codd Senecio mauricei Hilliard & B.L. Burtt Hemizygia cinerea Codd Senecio monticola DC. Satureja compacta Killick Senecio parascitus Hilliard Satureja grandibracteata Killick Senecio parentalis Hilliard & B.L. Burtt Stachys albiflora N.E.Br. Senecio polelensis Hilliard Syncolostemon macranthus (Gu¨rke) M. Ashby Senecio qathlambanus Hilliard Lobeliaceae (1/1) Senecio saniensis Hilliard & B.L. Burtt Lobelia galpinii Schltr. Senecio seminiveus J.M. Wood & M.S. Evans Mesembryanthemaceae (1/12) Senecio submontanus Hilliard & B.L. Burtt Delosperma alticola L. Bolus Senecio telmateius Hilliard Delosperma basuticum L. Bolus Senecio thamathuensis Hilliard Delosperma congestum L. Bolus Senecio tugelensis J.M. Wood & M.S. Evans Delosperma deleeuwiae Lavis Vernonia flanaganii (E. Phillips) Hilliard Delosperma galpinii L. Bolus Boraginaceae (1/1) Delosperma kofleri Lavis Cynoglossum alticola Hilliard & B.L. Burtt Delosperma nelii L. Bolus Brassicaceae (2/3) Delosperma nubigenum (Schltr.) L. Bolus Heliophila alpina Marais Delosperma pilosulum L. Bolus Heliophila formosa Hilliard & B.L. Burtt Delosperma reynoldsii Lavis Lepidium basuticum Marais Delosperma scabripes L. Bolus C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 125

Delosperma wiumii Lavis Passerina drakensbergensis Hilliard & B.L. Burtt Molluginaceae (1/2) Struthiola angustiloba Peterson & Hilliard Psammotropha alternifolia Killick Valerianaceae (1/1) Psammotropha obtusa Adamson Valeriana capensis Thunb. var. nana B.L. Burtt Proteaceae (1/1) Protea nubigena Rourke Angiospermae–Monocotyledonae Rhamnaceae (1/1) Alliaceae (1/1) Phylica thodei E. Phillips Tulbaghia montana Vosa Rosaceae (2/5) Amaryllidaceae (1/1) Alchemilla colura Hilliard Cyrtanthus erubescens Killick Alchemilla galpinii Hauman & Balle Aponogetonaceae (1/1) Cliffortia dracomontana C.M. Whitehouse Aponogeton ranunculiflorus Jacot Guill. & Marais Cliffortia filicauloides Weim. Asparagaceae (1/1) Cliffortia spathulata Weim. Asparagus stellatus Baker Rubiaceae (1/1) Asphodelaceae (3/7) Anthospermum basuticum Puff Aloe polyphylla Scho¨nland ex Pillans Santalaceae (1/5) Kniphofia albomontana Baijnath Thesium alatum Hilliard & B.L. Burtt Kniphofia evansii Baker Thesium congestum R.A. Dyer Kniphofia hirsuta Codd Thesium cordatum A.W. Hill Kniphofia ichopensis Schinz var. aciformis Codd Thesium decipiens Hilliard & B.L. Burtt Kniphofia thodei Baker Thesium durum Hilliard & B.L. Burtt Trachyandra smalliana Hilliard & B.L. Burtt Scrophulariaceae (10/37) Colchicaceae (1/3) Diascia anastrepta Hilliard & B.L. Burtt Wurmbea burttii B. Nord. Diascia austromontana K.E. Steiner Wurmbea pusilla E. Phillips Diascia barberae Hook.f. Wurmbea tenuis (Hook.f.) Baker subsp. australis B. Nord. Diascia fetcaniensis Hilliard & B.L. Burtt Cyperaceae (6/13) Diascia lilacina Hilliard & B.L. Burtt Carex killickii Nelmes Diascia megathura Hilliard & B.L. Burtt Carex monotropa Nelmes Diascia tugelensis Hilliard & B.L. Burtt Carex subinflata Nelmes Diascia vigilis Hilliard & B.L. Burtt Ficinia filiculmea B.L. Burtt Glumicalyx apiculatus (E. Mey.) Hilliard & B.L. Burtt Fuirena tenuis P.L. Forbes Glumicalyx flanaganii (Hiern) Hilliard & B.L. Burtt Isolepis angelica B.L. Burtt Glumicalyx goseloides (Diels) Hilliard & B.L. Burtt Isolepis pellocolea B.L. Burtt Glumicalyx lesuticus Hilliard & B.L. Burtt Schoenoxiphium bracteosum Kukkonen Glumicalyx montanus Hiern Schoenoxiphium burttii Kukkonen Glumicalyx nutans (Rolfe) Hilliard & B.L. Burtt Schoenoxiphium distinctum Kukkonen Harveya leucopharynx Hilliard & B.L. Burtt Schoenoxiphium molle Kukkonen Harveya pulchra Hilliard & B.L. Burtt Schoenoxiphium strictum Kukkonen Jamesbrittenia aspleniifolia Hilliard Tetraria sp. nov. (Killick 1596 NU) Jamesbrittenia beverlyana (Hilliard & B.L. Burtt) Hilliard Hyacinthaceae (6/8) Jamesbrittenia dentatisepala (Overkott) Hilliard Albuca humilis Baker Jamesbrittenia jurassica (Hilliard & B.L. Burtt) Hilliard Albuca rupestris Hilliard & B.L. Burtt Jamesbrittenia lesutica Hilliard Drimia saniensis (Hilliard & B.L. Burtt) J.C. Manning & Goldblatt Limosella vesiculosa Hilliard & B.L. Burtt Eucomis schijffii Reyneke Manulea crassifolia Benth. subsp. thodeana (Diels) Hilliard Galtonia regalis Hilliard & B.L. Burtt Manulea dregei Hilliard & B.L. Burtt Merwilla dracomontana (Hilliard & B.L. Burtt) Speta Manulea platystigma Hilliard & B.L. Burtt Ornithogalum diphyllum Baker Nemesia glabriuscula Hilliard & B.L. Burtt Ornithogalum sephtonii Hilliard & B.L. Burtt Selago flanaganii Rolfe Hypoxidaceae (3/6) Selago innata Marko¨tter Hypoxis tetramera Hilliard & B.L. Burtt Selago leptothrix Hilliard Rhodohypoxis deflexa Hilliard & B.L. Burtt Selago melliodora Hilliard Rhodohypoxis incompta Hilliard & B.L. Burtt Selago trauseldii Killick Rhodohypoxis rubella (Baker) Nel Selago witbergensis E. Mey. Rhodohypoxis thodiana (Nel) Hilliard & B.L. Burtt Strobilopsis wrightii Hilliard & B.L. Burtt Saniella verna Hilliard & B.L. Burtt Zaluzianskya chrysops Hilliard & B.L. Burtt Iridaceae (6/23) Zaluzianskya oreophila Hilliard & B.L. Burtt Crocosmia pearsei Oberm. Zaluzianskya rubrostellata Hilliard & B.L. Burtt Dierama jucundum Hilliard Zaluzianskya turritella Hilliard & B.L. Burtt Gladiolus flanaganii Baker Sterculiaceae (1/1) Gladiolus loteniensis Hilliard & B.L. Burtt Hermannia malvifolia N.E.Br. Gladiolus microcarpus G.J. Lewis Thymelaeaceae (3/7) Gladiolus saundersii Hook.f. Gnidia aberrans C.H. Wright Gladiolus symonsii F. Bolus Gnidia compacta (C.H. Wright) J.H. Ross Hesperantha alborosea Hilliard & B.L. Burtt Gnidia propinqua (Hilliard) B. Peterson Hesperantha altimontana Goldblatt Gnidia renniana Hilliard & B.L. Burtt Hesperantha baurii Baker subsp. formosa Hilliard & B.L. Burtt Gnidia singularis Hilliard Hesperantha brevistyla Goldblatt 126 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

Hesperantha crocopsis Hilliard & B.L. Burtt Alepidea natalensis J.M. Wood & M.S. Evans Hesperantha curvula Hilliard & B.L. Burtt Alepidea pilifera Weim. Hesperantha exiliflora Goldblatt Alepidea serrata Eckl. & Zeyh. var. serrata Hesperantha pubinervia Hilliard & B.L. Burtt Alepidea setifera N.E.Br. Hesperantha schelpeana Hilliard & B.L. Burtt Alepidea woodii Oliv. Moraea alpina Goldblatt Conium fontanum Hilliard & B.L. Burtt var. fontanum Moraea alticola Goldblatt Peucedanum thodei Arnold Moraea ardesiaca Goldblatt Polemannia montana Schltr. & H. Wolff Moraea carnea Goldblatt Polemannia simplicior Hilliard & B.L. Burtt Moraea dracomontana Goldblatt Apocynaceae (10/30) Romulea luteoflora (M.P. de Vos) M.P. de Vos var. sanisensis M.P. de Vos Anisotoma pedunculata N.E.Br. Romulea macowanii Baker var. alticola (B.L. Burtt) M.P. de Vos Asclepias cucullata (Schltr.) Schltr. Juncaceae (1/2) Asclepias macropus (Schltr.) Schltr. Juncus exsertus Buchenau subsp. lesuticus B.L. Burtt Aspidonepsis cognata (N.E.Br.) Nicholas & Goyder Juncus mollifolius Hilliard & B.L. Burtt Aspidonepsis diploglossa (Turcz.) Nicholas & Goyder Orchidaceae (6/11) Aspidonepsis flava (N.E.Br.) Nicholas & Goyder Brownleea galpinii Bolus subsp. major (Bolus) H.P. Linder Aspidonepsis reenensis (N.E.Br.) Nicholas & Goyder Corycium alticola Parkman & Schelpe Brachystelma petraeum R.A. Dyer Disa basutorum Schltr. Fanninia caloglossa Harv. Disa cephalotes Rchb.f. subsp. frigida (Schltr.) H.P. Linder Miraglossum pulchellum (Schltr.) Kupicha Disa dracomontana Schelpe ex H.P. Linder Miraglossum verticillare (Schltr.) Kupicha Disa galpinii Rolfe Pachyacris rhodantha Stewart & Langley Disa nivea H.P. Linder Pachycarpus campanulatus (Harv.) N.E.Br. var. campanulatus Disa oreophila Bolus subsp. erecta H.P. Linder Pachycarpus campanulatus (Harv.) N.E.Br. var. sutherlandii N.E.Br. Dracomonticola virginea (Bolus) H.P. Linder & Kurzweil Pachycarpus macrochilus (Schltr.) N.E.Br. Huttonaea grandiflora (Schltr.) Rolfe Pachycarpus natalensis N.E.Br. Schizochilus angustifolius Rolfe Pachycarpus plicatus N.E.Br. Poaceae (9/12) Pachycarpus rigidus E.Mey. Agrostis subulifolia Stapf Pachycarpus stenoglossus (E. Mey.) N.E.Br. Anthoxanthum brevifolium Stapf Pachycarpus vexillaris E. Mey. Colpodium drakensbergense Hedberg & I. Hedberg Schizoglossum atropurpureum E. Mey. subsp. atropurpureum Ehrharta longigluma C.E. Hubb. Schizoglossum bidens E. Mey. subsp. bidens Festuca killickii Kenn.-O’Byrne Schizoglossum flavum Schltr. Festuca vulpioides Steud. Schizoglossum hamatum E. Mey. Helictotrichon galpinii Schweick. Schizoglossum nitidum Schltr. Merxmuellera aureocephala (J.G. Anderson) Conert Schizoglossum stenoglossum Schltr. subsp. latifolium Kupicha Merxmuellera guillarmodiae Conert Xysmalobium parviflorum Harv. ex Scott-Elliot Pentaschistis exserta H.P. Linder Xysmalobium stockenstromense Scott-Elliot Pentaschistis praecox H.P. Linder Xysmalobium tysonianum (Schltr.) N.E.Br. Polevansia rigida De Winter Xysmalobium woodii N.E.Br. Restionaceae (1/1) Asteraceae (29/135) Restio galpinii Pillans Aster perfoliatus Oliv. Velloziaceae (1/1) Aster pleiocephalus (Harv.) Hutch. Xerophyta longicaulis Hilliard Athrixia angustissima DC. Numbers in parentheses after each family are total numbers of genera and Athrixia arachnoidea J.M. Wood & M.S. Evans ex J.M. Wood species. All taxa are arranged alphabetically. Athrixia fontana MacOwan Berkheya discolor (DC.) O. Hoffm. & Muschl. Berkheya griquana Hilliard & B.L. Burtt Appendix B Berkheya macrocephala J.M. Wood Berkheya montana J.M. Wood & M.S. Evans Berkheya multijuga (DC.) Roessler The near-endemic angiosperms of the Drakensberg Alpine Centre Berkheya onopordifolia (DC.) O. Hoffm. ex Burtt Davy var. onopordifolia Angiospermae–Dicotyledonae Berkheya purpurea (DC.) Mast. Acanthaceae (2/2) Berkheya rhapontica (DC.) Hutch. & Burtt Davy subsp. aristosa (DC.) Adhatoda andromeda (Lindau) C.B. Clarke Roessler var. aristosa Barleria monticola Oberm. Berkheya rhapontica (DC.) Hutch. & Burtt Davy subsp. aristosa (DC.) Achariaceae (1/1) Roessler var. exalata Roessler Guthriea capensis Bolus Berkheya speciosa (DC.) O. Hoffm. subsp. ovata Roessler Anacardiaceae (1/6) Chrysanthemoides monilifera (L.) Norl. subsp. canescens (DC.) Norl. Rhus bolusii Sond. ex Engl. Cineraria albicans N.E.Br. Rhus divaricata Eckl. & Zeyh. Cineraria aspera Thunb. Rhus dregeana Sond. Cineraria britteniae Hutch. & R.A. Dyer Rhus erosa Thunb. Cineraria dieterlenii E. Phillips Rhus krebsiana C. Presl ex Engl. Cineraria geraniifolia DC. Rhus montana Diels Cineraria grandibracteata Hilliard Apiaceae (4/11) Cineraria mollis E. Mey. ex DC. Alepidea galpinii Dummer Cotula hispida (DC.) Harv. Alepidea longifolia E. Mey. var. angusta Dummer Cotula leptalea DC. C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 127

Dimorphotheca caulescens (Harv.) Harv. Macowania pinifolia (N.E.Br.) Kroner Dimorphotheca jucunda E. Phillips Macowania pulvinaris N.E.Br. Euryops annae E. Phillips Nidorella agria Hilliard Euryops candollei Harv. Pentzia cooperi Harv. Euryops empetrifolius DC. Phymaspermum woodii (Thell.) Ka¨llersjo¨ Euryops evansii Schltr. subsp. evansii Printzia auriculata Harv. Euryops oligoglossus DC. subsp. oligoglossus Printzia pyrifolia Less. Euryops pedunculatus N.E.Br. Schistostephium hipiifolium (DC.) Hutch. Euryops tysonii E. Phillips Senecio arabidifolius O. Hoffm. Felicia petiolata (Harv.) N.E.Br. Senecio barbatus DC. Felicia quinquenervia (Klatt) Grau Senecio baurii Oliv. Felicia rosulata Yeo Senecio brevidentatus M.D. Hend. Garuleum sonchifolium (DC.) Norl. Senecio cathcartensis O. Hoffm. Garuleum woodii Schinz Senecio citriceps Hilliard & B.L. Burtt Gnaphalium griquense Hilliard & B.L. Burtt Senecio flanaganii E. Phillips Gymnopentzia bifurcata Benth. Senecio gramineus Harv. Helichrysum albirosulatum Killick Senecio harveianus MacOwan Helichrysum albo-brunneum S. Moore Senecio haygarthii Hilliard Helichrysum alticolum Bolus Senecio heliopsis Hilliard & B.L. Burtt Helichrysum ammitophilum Hilliard Senecio hieracioides DC. Helichrysum anomalum Less. Senecio hirsutilobus Hilliard Helichrysum argentissimum J.M. Wood Senecio hypochoerideus DC. Helichrysum argyrophyllum DC. Senecio isatidioides E. Phillips & C.A. Sm. Helichrysum aureum (Houtt.) Merr. var. scopulosum (M.D. Hend.) Hilliard Senecio lydenburgensis Hutch. & Burtt Davy Helichrysum aureum (Houtt.) Merr. var. serotinum Hilliard Senecio macowanii Hilliard Helichrysum auriceps Hilliard Senecio macrospermus DC. Helichrysum bellidiastrum Moeser Senecio mooreanus Hutch. & Burtt Davy Helichrysum confertifolium Klatt Senecio napifolius MacOwan Helichrysum cooperi Harv. Senecio paludaffinis Hilliard Helichrysum cymosum (L.) D. Don subsp. calvum Hilliard Senecio paucicalyculatus Klatt Helichrysum dasycephalum O. Hoffm. Senecio polyodon DC. var. subglaber (Kuntze) Hilliard & B.L. Burtt Helichrysum elegantissimum DC. Senecio praeteritus Killick Helichrysum epapposum Bolus Senecio scitus Hutch. & Burtt Davy Helichrysum fulvum N.E.Br. Senecio subcoriaceus Schltr. Helichrysum glomeratum Klatt Senecio subrubriflorus O. Hoffm. Helichrysum grandibracteatum M.D. Hend. Senecio tanacetopsis Hilliard Helichrysum griseolanatum Hilliard Senecio ulopterus Thell. Helichrysum gymnocomum DC. Sonchus jacottetianus Thell. Helichrysum hypoleucum Harv. Troglophyton capillaceum (Thunb.) Hilliard & B.L. Burtt subsp. diffusum Helichrysum infaustum J.M. Wood & M.S. Evans (DC.) Hilliard Helichrysum krookii Moeser Ursinia alpina N.E.Br. Helichrysum lingulatum Hilliard Ursinia montana DC. subsp. montana Helichrysum melanacme DC. Vernonia thodei E. Phillips Helichrysum montanum DC. Boraginaceae (4/6) Helichrysum monticola Hilliard Afrotysonia glochidiata (R.R. Mill) R.R. Mill Helichrysum nanum Klatt Cynoglossum austroafricanum Hilliard & B.L. Burtt Helichrysum natalitium DC. Cynoglossum spelaeum Hilliard & B.L. Burtt Helichrysum opacum Klatt Lithospermum afromontanum Weim. Helichrysum oreophilum Klatt Lithospermum papillosum Thunb. Helichrysum pedunculatum Hilliard & B.L. Burtt Myosotis semiamplexicaulis A. DC. Helichrysum psilolepis Harv. Brassicaceae (1/1) Helichrysum scitulum Hilliard & B.L. Burtt Lepidium myriocarpum Sond. Helichrysum sessile DC. Buddlejaceae (1/2) Helichrysum subluteum Burtt Davy Buddleja auriculata Benth. Helichrysum sutherlandii Harv. Buddleja loricata Leeuwenb. Helichrysum tenax M.D. Hend. var. pallidum Hilliard & B.L. Burtt Campanulaceae (2/18) Helichrysum tenax M.D. Hend. var. tenax Craterocapsa congesta Hilliard & B.L. Burtt Helichrysum trilineatum DC. Craterocapsa insizwae (Zahlbr.) Hilliard & B.L. Burtt Helichrysum vernum Hilliard Craterocapsa montana (A.DC.) Hilliard & B.L. Burtt Hilliardia zuurbergensis (Oliv.) B. Nord. Craterocapsa tarsodes Hilliard & B.L. Burtt Hirpicium armerioides (DC.) Roessler Wahlenbergia acicularis Brehmer Inulanthera dregeana (DC.) Ka¨llersjo¨ Wahlenbergia appressifolia Hilliard & B.L. Burtt Inulanthera leucoclada (DC.) Ka¨llersjo¨ Wahlenbergia capillata Brehmer Inulanthera montana (J.M. Wood) Ka¨llersjo¨ Wahlenbergia cooperi Brehmer Inulanthera tridens (Oliv.) Ka¨llersjo¨ Wahlenbergia cuspidata Brehmer Lactuca tysonii (E. Phillips) C. Jeffrey Wahlenbergia depressa J.M. Wood & M.S. Evans Lepidostephium asteroides (Bolus & Schltr.) Kroner Wahlenbergia fasciculata Brehmer Macowania conferta (Benth.) E. Phillips Wahlenbergia galpiniae Schltr. 128 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

Wahlenbergia krebsii Cham. subsp. krebsii Lotononis sericophylla Benth. Wahlenbergia pallidiflora Hilliard & B.L. Burtt Melolobium alpinum Eckl. & Zeyh. Wahlenbergia paucidentata Schinz Melolobium obcordatum Harv. Wahlenbergia polytrichifolia Schltr. subsp. polytrichifolia Otholobium caffrum (Eckl. & Zeyh.) C.H. Stirt. Wahlenbergia rivularis Diels Otholobium fumeum C.H. Stirt. Wahlenbergia squamifolia Brehmer Otholobium polystictum (Benth. ex Harv.) C.H. Stirt. Caryophyllaceae (2/4) Psoralea sp. nov. Cerastium fontanum Baumg. subsp. triviale (Link) Jalas Rhynchosia cooperi (Harv. ex Baker f.) Burtt Davy Cerastium indicum Wight & Arn. Sutherlandia montana E. Phillips & R.A. Dyer Dianthus basuticus Burtt Davy subsp. basuticus var. basuticus Tephrosia marginella H.M.L. Forbes Dianthus basuticus Burtt Davy subsp. basuticus var. grandiflorus Hooper Fumariaceae (1/1) Crassulaceae (1/8) Cysticapnos pruinosa (Bernh.) Lide¯n Crassula compacta Scho¨nland Gentianaceae (2/6) Crassula gemmifera Friedrich Chironia peglerae Prain Crassula natalensis Scho¨nland Sebaea natalensis Schinz Crassula peploides Harv. Sebaea procumbens A.W. Hill Crassula setulosa Harv. var. longiciliata To¨lken Sebaea rehmannii Schinz Crassula setulosa Harv. var. rubra (N.E.Br.) G.D. Rowley Sebaea repens Schinz Crassula tenuicaulis Scho¨nland Sebaea thomasii (S. Moore) Schinz Crassula umbraticola N.E.Br. Geraniaceae (3/16) Cucurbitaceae (1/1) Geranium brycei N.E.Br. Coccinia hirtella Cogn. Geranium magniflorum R. Knuth Dipsacaceae (2/2) Geranium multisectum N.E.Br. Cephalaria natalensis Kuntze Geranium pulchrum N.E.Br. Scabiosa drakensbergensis B.L. Burtt Geranium robustum Kuntze Ebenaceae (2/2) Geranium schlechteri R. Knuth Diospyros austro-africana De Winter var. rubriflora (De Winter) De Winter Monsonia attenuata Harv. Euclea coriacea A.DC. Monsonia brevirostrata R. Knuth Ericaceae (1/12) Monsonia grandifolia R. Knuth Erica algida Bolus Pelargonium aridum R.A. Dyer Erica alopecurus Harv. var. alopecurus Pelargonium bowkeri Harv. Erica caespitosa Hilliard & B.L. Burtt Pelargonium capituliforme R. Knuth Erica caffrorum Bolus var. caffrorum Pelargonium dispar N.E.Br. Erica cooperi Bolus var. cooperi Pelargonium griseum R. Knuth Erica drakensbergensis Guthrie & Bolus Pelargonium leucophyllum Turcz. Erica evansii (N.E.Br.) E.G.H. Oliv. Pelargonium ranunculophyllum (Eckl. & Zeyh.) Baker Erica lasiocarpa Guthrie & Bolus Gesneriaceae (1/3) Erica oatesii Rolfe var. oatesii Streptocarpus gardenii Hook. Erica schlechteri Bolus Streptocarpus pentherianus Fritsch Erica straussiana Gilg Streptocarpus pusillus Harv. ex C.B. Clarke Erica trichoclada Guthrie & Bolus Lamiaceae (4/13) Euphorbiaceae (2/2) Plectranthus grallatus Briq. Clutia katharinae Pax Salvia repens Burch. ex Benth. var. transvaalensis Hedge Euphorbia natalensis Bernh. Satureja reptans Killick Fabaceae (14/33) Stachys cymbalaria Briq. Argyrolobium lotoides Harv. Stachys dregeana Benth. Argyrolobium nigrescens Dummer Stachys flexuosa Skan Argyrolobium sankeyi Harms Stachys hyssopoides Burch. ex Benth. Argyrolobium sericosemium Harms Stachys kuntzei Gu¨rke Calpurnia reflexa A.J. Beaumont Stachys linearis Burch. ex Benth. Elephantorrhiza woodii E. Phillips var. pubescens E. Phillips Stachys rudatisii Skan Eriosema distinctum N.E.Br. Stachys sessilis Gu¨rke Hoffmannseggia sandersonii (Harv.) Engl. Stachys simplex Schltr. Indigofera cuneifolia Eckl. & Zeyh. var. cuneifolia Stachys tysonii Skan Indigofera foliosa E. Mey. Lobeliaceae (2/7) Indigofera trifolioides Baker f. Cyphia aspergilloides E. Wimm. var. aspergilloides Indigofera woodii Bolus var. woodii Cyphia longifolia N.E.Br. Lessertia stricta L. Bolus Cyphia natalensis E. Phillips Lessertia thodei L. Bolus Cyphia ramosa E. Wimm. Lotononis adpressa N.E.Br. subsp. adpressa Cyphia rogersii S. Moore subsp. winteri E. Wimm. Lotononis divaricata (Eckl. & Zeyh.) Benth. Cyphia tysonii E. Phillips Lotononis eriocarpa (E. Mey.) B.-E. van Wyk Lobelia preslii A.DC. Lotononis galpinii Dummer Malvaceae (1/3) Lotononis lanceolata (E. Mey.) Benth. Anisodontea julii (Burch. ex DC.) Bates subsp. julii Lotononis lotononoides (Scott-Elliot) B.-E. van Wyk Anisodontea julii (Burch. ex DC.) Bates subsp. pannosa (Bolus) Bates Lotononis macrosepala Conrath Anisodontea julii (Burch. ex DC.) Bates subsp. prostrata (E. Mey. ex Turcz.) Lotononis procumbens Bolus Bates Lotononis pulchella (E. Mey.) B.-E. van Wyk Melianthaceae (1/1) C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 129

Melianthus villosus Bolus Nemesia albiflora N.E.Br. Mesembryanthemaceae (2/11) Nemesia caerulea Hiern Delosperma ashtonii L. Bolus Nemesia silvatica Hilliard Delosperma clavipes Lavis Nemesia umbonata (Hiern) Hilliard & B.L. Burtt Delosperma cooperi (Hook.f.) L. Bolus Phygelius aequalis Harv. ex Hiern Delosperma crassuloides (Haw.) L. Bolus Phygelius capensis E. Mey. ex Benth. Delosperma hirtum (N.E.Br.) Schwantes Selago densiflora Rolfe Delosperma lavisiae L. Bolus Selago galpinii Schltr. Delosperma lineare L. Bolus Selago immersa Rolfe Delosperma roseopurpureum Lavis Selago monticola J.M. Wood & M.S. Evans Delosperma sutherlandii (Hook.f.) N.E.Br. Selago multispicata Hilliard Delosperma wethamae L. Bolus Selago saxatilis E. Mey. Ruschia putterillii (L. Bolus) L. Bolus Selago speciosa Rolfe Molluginaceae (1/1) Selago trinervia E. Mey. Psammotropha mucronata (Thunb.) Fenzl var. marginata Adamson Sutera cooperi Hiern Polygalaceae (2/3) Sutera neglecta (J.M. Wood & M.S. Evans) Hiern Muraltia saxicola Chodat Sutera polelensis Hiern subsp. polelensis Polygala praticola Chodat Zaluzianskya crocea Schltr. Polygala rhinostigma Chodat Zaluzianskya distans Hiern Polygonaceae (1/2) Zaluzianskya glareosa Hilliard & B.L. Burtt Rumex dregeanus Meisn. subsp. montanus B.L. Burtt Zaluzianskya microsiphon (Kuntze) K. Schum. Rumex woodii N.E.Br. Zaluzianskya pulvinata Killick Proteaceae (1/2) Zaluzianskya schmitziae Hilliard & B.L. Burtt Protea dracomontana Beard Zaluzianskya spathacea (Benth.) Walp. Protea subvestita N.E.Br. Sterculiaceae (1/3) Ranunculaceae (2/3) Hermannia gerrardii Harv. Anemone caffra (Eckl. & Zeyh.) Harv. Hermannia oligosperma K. Schum. Anemone fanninii Harv. ex Mast. Hermannia woodii Schinz Ranunculus baurii MacOwan Thymelaeaceae (1/2) Rhamnaceae (1/1) Gnidia baurii C.H. Wright Phylica tysonii Pillans var. tysonii Gnidia polyantha Gilg Rosaceae (4/8) Valerianaceae (1/1) Alchemilla hirsuto-petiolata (De Wild.) Rothm. Valeriana capensis Thunb. var. lanceolata N.E.Br. Alchemilla natalensis Engl. Alchemilla woodii Kuntze Angiospermae–Monocotyledonae Cliffortia browniana Burtt Davy Agapanthaceae (1/2) Cliffortia paucistaminea Weim. var. paucistaminea Agapanthus campanulatus F.M. Leight. subsp. patens (F.M. Leight.) F.M. Cliffortia repens Schltr. Leight. Geum capense Thunb. Agapanthus nutans F.M. Leight. Rubus ludwigii Eckl. & Zeyh. subsp. spatiosus C.H. Stirt. Alliaceae (1/1) Rubiaceae (2/2) Tulbaghia natalensis Baker Anthospermum monticola Puff Amaryllidaceae (3/14) Galium scabrelloides Puff Brunsvigia grandiflora Lindl. Santalaceae (1/3) Brunsvigia natalensis Baker Thesium confine Sond. Brunsvigia undulata F.M. Leight. Thesium macrogyne A.W. Hill Cyrtanthus attenuatus R.A. Dyer Thesium nigrum A.W. Hill Cyrtanthus epiphyticus J.M. Wood Scrophulariaceae (13/44) Cyrtanthus falcatus R.A. Dyer Alectra basutica (E. Phillips) Melch. Cyrtanthus flanaganii Baker Alectra thyrsoidea Melch. Cyrtanthus obrienii Baker Bowkeria verticillata (Eckl. & Zeyh.) Schinz Cyrtanthus stenanthus Baker var. stenanthus Diascia cordata N.E.Br. Cyrtanthus tuckii Baker var. viridilobus I. Verd. Diascia integerrima E. Mey. ex Benth. Nerine angustifolia (Baker) Baker Diascia purpurea N.E.Br. Nerine appendiculata Baker Diascia stachyoides Schltr. ex Hiern Nerine bowdenii Watson Diascia stricta Hilliard & B.L. Burtt Nerine pancratioides Baker Glekia krebsiana (Benth.) Hilliard Anthericaceae (1/1) Harveya scarlatina (Benth.) Hiern Chlorophytum acutum (C.H. Wright) Nordal Hyobanche rubra N.E.Br. Araceae (1/1) Jamesbrittenia breviflora (Schltr.) Hilliard Zantedeschia valida (Letty) Y. Singh Jamesbrittenia filicaulis (Benth.) Hilliard Asparagaceae (1/2) Jamesbrittenia pristisepala (Hiern) Hilliard Asparagus concinnus (Baker) Kies Jamesbrittenia stricta (Benth.) Hilliard Asparagus microraphis (Kunth) Baker Manulea buchneroides Hilliard & B.L. Burtt Asphodelaceae (3/17) Manulea crassifolia Benth. subsp. crassifolia Aloe aristata Haw. Manulea florifera Hilliard & B.L. Burtt Aloe pratensis Baker Manulea paniculata Benth. Aloe striatula Haw. var. striatula Manulea rhodantha Hilliard subsp. rhodantha Kniphofia albescens Codd 130 C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132

Kniphofia angustifolia (Baker) Codd Dierama robustum N.E.Br. Kniphofia brachystachya (Zahlbr.) Codd Dierama trichorhizum (Baker) N.E.Br. Kniphofia breviflora Baker Dierama tysonii N.E.Br. Kniphofia caulescens Baker Gladiolus inandensis Baker Kniphofia fibrosa Baker Gladiolus mortonius Herb. Kniphofia fluviatilis Codd Gladiolus oppositiflorus Herb. Kniphofia ichopensis Schinz var. ichopensis Gladiolus parvulus Schltr. Kniphofia northiae Baker Gladiolus pubigerus G.J.Lewis Kniphofia porphyrantha Baker Hesperantha candida Baker Kniphofia ritualis Codd Hesperantha glareosa Hilliard & B.L. Burtt Kniphofia stricta Codd Hesperantha grandiflora G.J.Lewis Kniphofia triangularis Kunth subsp. triangularis Hesperantha hygrophila Hilliard & B.L. Burtt Trachyandra asperata Kunth var. basutoensis (Poelln.) Oberm. Hesperantha ingeliensis Hilliard & B.L. Burtt Colchicaceae (1/2) Hesperantha scopulosa Hilliard & B.L. Burtt Wurmbea angustifolia B. Nord. Hesperantha vernalis Hilliard & B.L. Burtt Wurmbea elatior B. Nord. Hesperantha woodii Baker Cyperaceae (4/7) Moraea albicuspa Goldblatt Carpha filifolia Reid & T.H. Arnold Moraea brevistyla (Goldblatt) Goldblatt Cyperus schlechteri C.B. Clarke Moraea hiemalis Goldblatt Ficinia cinnamomea C.B. Clarke Moraea huttonii (Baker) Oberm. Schoenoxiphium basutorum Turrill Moraea inclinata Goldblatt Schoenoxiphium filiforme Ku¨k. Moraea modesta Killick Schoenoxiphium perdensum Kukkonen Moraea robusta (Goldblatt) Goldblatt Schoenoxiphium schweickerdtii Merxm. & Podlech Moraea trifida R.C. Foster Eriocaulaceae (1/1) Moraea unibracteata Goldblatt Eriocaulon hydrophilum Marko¨tter Romulea macowanii Baker var. macowanii Eriospermaceae (1/1) Romulea macowanii Baker var. oreophila M.P. de Vos Eriospermum ornithogaloides Baker Watsonia gladioloides Schltr. Hyacinthaceae (7/11) Watsonia lepida N.E.Br. Albuca shawii Baker Orchidaceae (11/41) Albuca xanthocodon Hilliard & B.L. Burtt Brownleea macroceras Sond. Drimia sphaerocephala Baker Corycium flanaganii (Bolus) Kurzweil & H.P. Linder Eucomis bicolor Baker Corycium tricuspidatum Bolus Eucomis humilis Baker Disa cephalotes Rchb.f. subsp. cephalotes Galtonia candicans (Baker) Decne. Disa cooperi Rchb.f. Galtonia princeps (Baker) Decne. Disa crassicornis Lindl. Galtonia viridiflora I. Verd. Disa montana Sond. Massonia echinata L.f. Disa oreophila Bolus subsp. oreophila Merwilla plumbea (Lindl.) Speta Disa patula Sond. var. patula Resnova lachenalioides (Baker) Van der Merwe Disa pulchra Sond. Hypoxidaceae (3/13) Disa sanguinea Sond. Empodium elongatum (Nel) B.L. Burtt Disa sankeyi Rolfe Empodium monophyllum (Nel) B.L. Burtt Disa scullyi Bolus Hypoxis costata Baker Disa stricta Sond. Hypoxis lata Nel Disa thodei Schltr. ex Kraenzl. Hypoxis ludwigii Baker Disa tysonii Bolus Hypoxis neliana Schinz Disperis cardiophora Harv. Hypoxis oblonga Nel Disperis concinna Schltr. Hypoxis parvula Baker var. albiflora B.L. Burtt Disperis cooperi Harv. Hypoxis parvula Baker var. parvula Disperis oxyglossa Bolus Rhodohypoxis baurii (Baker) Nel var. baurii Disperis renibractea Schltr. Rhodohypoxis baurii (Baker) Nel var. confecta Hilliard & B.L. Burtt Disperis stenoplectron Rchb.f. Rhodohypoxis baurii (Baker) Nel var. platypetala (Baker) Nel Disperis tysonii Bolus Rhodohypoxis milloides (Baker) Hilliard & B.L. Burtt Disperis wealei Rchb.f. Iridaceae (8/42) Eulophia calanthoides Schltr. Aristea angolensis Baker subsp. acutivalvis Weim. Eulophia zeyheriana Sond. Aristea angolensis Baker subsp. majubensis (Baker) Weim. Holothrix incurva Lindl. Aristea flexicaulis Baker Holothrix scopularia (Lindl.) Rchb.f. Aristea grandis Weim. Holothrix thodei Rolfe Aristea montana Baker Huttonaea fimbriata (Harv.) Rchb.f. Crocosmia pottsii (Macnab ex Baker) N.E.Br. Huttonaea oreophila Schltr. Dierama ambiguum Hilliard Huttonaea pulchra Harv. Dierama cooperi N.E.Br. Huttonaea woodii Schltr. Dierama dissimile Hilliard Neobolusia tysonii (Bolus) Schltr. Dierama dracomontanum Hilliard Pterygodium cooperi Rolfe Dierama latifolium N.E.Br. Pterygodium hastatum Bolus Dierama pauciflorum N.E.Br. Pterygodium leucanthum Bolus Dierama pictum N.E.Br. Satyrium longicauda Lindl. var. jacottetianum (Kraenzl.) A.V. Hall C. Carbutt, T.J. Edwards / South African Journal of Botany 72 (2006) 105–132 131

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