Trimelopter craibii - Asparagaceae subfamily Scilloideae

Phytotaxa 87 (3): 50–60 (2013) www.mapress.com/ phytotaxa / Copyright © 2013 Magnolia Press

ISSN 1179-3155 (print edition)

Article

PHYTOTAXA ISSN 1179-3163 (online edition)

http://dx.doi.org/10.11646/phytotaxa.87.3.3

Trimelopter craibii (Hyacinthaceae, Ornithogaloideae), a new species from the North West Province of South Africa MARIO MARTÍNEZ-AZORÍN1, MANUEL B. CRESPO1 & ANTHONY P. DOLD2 1

CIBIO (Instituto Universitario de la Biodiversidad), Universidad de Alicante, P. O. Box 99, E-03080 Alicante, Spain. E-mail: [email protected] 2 Selmar Schonland Herbarium, Department of Botany, Rhodes University, Grahamstown 6140, South Africa.

Abstract The genus Trimelopter has been recently reinstated to include Ornithogalum unifolium and other closely related species from Southern Africa, comprising up to 10 species. Within the context of a revision of Trimelopter, a new species, T. craibii, is here formally described to name plants discovered by the late Charles Craib in the North West Province of South Africa. This taxon is closely related to T. dyeri and T. unifolium, but it can be clearly differentiated by floral and vegetative characters. Data on morphology, ecology, and distribution are reported for this new species, and affinities and divergences with other closely related taxa are also discussed. The new combination T. unifolium var. vestitum is also proposed. Key words: Asparagaceae, distribution, ecology, Hyacintheae, Scilloideae, taxonomy, Trimelopter unifolium var. vestitum

Introduction The family Hyacinthaceae includes about 1000 species of bulbous plants which are mainly distributed throughout Europe, Africa and south-west Asia, with a single small genus in South America (APG 2003). Alternatively, Hyacinthaceae is treated as subfamily Scilloideae of Asparagaceae, and the subfamilies above are then treated as tribes Hyacintheae, Ornithogaleae, Oziroëeae and Urgineeae (e.g. APG 2009, Chase et al. 2009). Generic circumscription within Hyacinthaceae subfam. Ornithogaloideae has been a matter of controversy during the last couple of decades. The latest comprehensive study in the group (cf. MartínezAzorín et al. 2011) demonstrated the existence of up to 19 monophyletic genera which are characterized by a clear syndrome of morphological characters, making genus concepts intuitive, homogeneous in floral morphology, and therefore easy to define and to work with. The genus Trimelopter Rafinesque (1837: 24) currently includes 10 species and is characterized by the presence of a single, elliptic to narrowly oblong leaf (exceptionally two or three leaves), usually flattened against the ground. The ovaries have two usually prominent longitudinal dorsal keels in each carpel and the seeds are unequally compressed or semi-discoid (cf. Dyer 1931, Leighton 1944, Obermeyer 1978, MartínezAzorín et al. 2011). The peculiar morphology of these plants allowed various authors to treat them at different taxonomic ranks. Rafinesque (1837) described the genus Trimelopter, including a single species, Trimelopter fuscatum (Jacquin 1795: 19) Rafinesque (1837: 24) (= Ornithogalum fuscatum Jacq.), which is currently regarded as a synonym of T. unifolium (Retzius 1781: 17) Martínez-Azorín et al. (2011: 26) (= O. unifolium Retz.). The genus Ardernia Salisbury (1866: 35) is also based on O. fuscatum and it is therefore to be considered

50 Accepted by Lorenzo Peruzzi: 26 Feb. 2013; published online in PDF: 15 Mar. 2013

illegitimate. It is remarkable that these genera have been mostly overlooked by modern botanists. Obermeyer (1978), in her taxonomic revision of Ornithogalum Linnaeus (1753: 306) in Southern Africa, accepted a single species in this group, O. unifolium, which resulted in a variable species concept with 6 synonyms, including O. fuscatum. Müller-Doblies & Müller-Doblies (1996) presented the most recent revision of Ornithogalum sensu lato in Southern Africa and described O. subgenus Urophyllon section Monarchos Müller-Doblies & Müller-Doblies (1996: 470) to accommodate 11 related species, typified by O. unifolium. These authors described 6 new species in this section and 3 further species were reinstated from synonymy of O. unifolium sensu Obermeyer (1978). One of those newly described species, O. rotatum Müller-Doblies & Müller-Doblies (1996: 482), was included in that section on the basis of its single leaf, but flower morphology and phylogenetic evidence clearly place this species in the genus Neopatersonia Schönland (1912: 251) (cf. Manning et al. 2009, Martínez-Azorín et al. 2011). Recent phylogenetic studies in Ornithogaloideae show O. unifolium and O. etesiogaripense MüllerDoblies & Müller-Doblies (1996: 476) (= Trimelopter) as a monophyletic clade sister to species of Albuca Linnaeus (1762: 438), Coilonox Rafinesque (1837: 28) and Stellarioides Medikus (1790: 369) sensu stricto. On this basis, Manning et al. (2009) combined section Monarchos as Albuca subgenus Monarchos (U.Müll.Doblies & D.Müll.-Doblies) Manning & Goldblatt (2009: 92), accepting a very variable genus Albuca as regards to flower, fruit and seed morphology. However, due to the clear morphological and phylogenetic independence of these plants, the genus Trimelopter was reinstated by Martínez-Azorín et al. (2011) and this concept is followed here. Two different species of Trimelopter were found by the late Charles Craib in the Lichtenburg and Delareyville districts in the North West Province of South Africa, differing mainly in leaf structure, flower morphology and flowering time. These species, first regarded to belong to Ornithogalum, were extensively studied during the last decade (Craib 2001a, 2001b, 2001c, 2005a, 2005b), however have not been yet formally described. The taxonomy of Trimelopter is far from being satisfactory, especially regarding the T. unifolium complex which includes several synonyms with different morphologies regarding flower and leaf structure. This species includes two extremes of variation that have been described as varieties under Ornithogalum unifolium (Müller-Doblies & Müller-Doblies 1996). Both taxa are accepted here and therefore the new combination T. unifolium var. vestitum is established below for individuals with markedly hairy leaves. Charles Craib sent us plants he collected in the surroundings of Barberspan, in the Delareyville district (Craib 2005a). They appear to be related to T. unifolium or perhaps to T. dyeri (Von Poellnitz 1944: 209) MartínezAzorín et al. (2011: 26), though further studies are needed to decide about their identity. However, other plants we received from him that were harvested from Trekdrift Farm in the Lichtenburg district (Craib 2001a, 2001b, 2001c, 2005a, 2005b), though apparently related to T. dyeri, show a distinct syndrome of floral and vegetative characters that allow the description of a new species, Trimelopter craibii Mart.-Azorín, M.B.Crespo & A.P.Dold.

Materials and methods Herbarium specimens from the following herbaria were studied: ABH, GRA, K, KMG, NBG, and WIND (acronyms according Thiers, 2013). Moreover, detailed morphological studies of Trimelopter craibii, T. dyeri and T. unifolium sensu lato were undertaken based on wild and cultivated plants and herbarium specimens. All floral measurements in T. craibii (10 plants from the type locality) and T. unifolium var. vestitum (2 plants from the type locality) were taken from cultivated plants in Grahamstown, South Africa. Ranks of measurements include the minimum and maximum values recorded, and in some cases the extreme ones (less than 10% of observations) are added in parentheses. Authorities of the cited taxa follow IPNI (2013).

TRIMELOPTER CRAIBII, A NEW SPECIES FROM SOUTH AFRICA

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Results and discussion Trimelopter craibii Mart.-Azorín, M.B.Crespo & A.P.Dold, sp. nov. (Figs. 1−2) Planta insignis, omnibus speciebus generis distincta et facile distinguitur. Folium solitarium ad solum appressum, coriaceum, oblongum, subconvexum, supra pilis validis erectis 0.6−0.8 mm longis dense vestitum, margine albido, incrassato, papillis minutis usque ad 0.05 mm longis densissime obsito, subtus glaberrimum. Scapus 1−5 cm longus; racemus brevissimus, usque ad 2.5 cm longus, 1−5-florus. Tepala 8.5−13 mm longa, albicantia, in dorso fascia longitudinali media viridi instructa. Ovarium ovatum, trilobatum, costis longitudinalibus vix prominentibus, infirme carinatis; stylus 3−3.5 mm longus. Habitu cum T. dyeri primo adspectu congruens sed illa valde differt quae folium supra et ad marginem papillis minutis dense obsitum, tepala minora (5−7 mm longa), ovarium globosum vel obovatum carinis longitudinalibus magis prominentibus, et stylum brevius (usque ad 2 mm longum) habet. A T. unifolio multo discrepat quae folium longius (usque ad 12 cm longum), non coriaceum, subconcavum, supra pilis tenuioribus brevioribusque (0.05−0.4 mm longis) laxiore dispositis, in margine breviter laxeque papillosum, et inflorescentiam majorem floribus plus numerosis exhibet. Type:—SOUTH AFRICA. North West Province: Mafikeng (2525DD), Lichtenburg district, ca. 25 km northwest of Lichtenburg to Zeerust on R505, west of Molopo Oog, Trekdrift Farm, in soil pockets of low dolomite outcrops, 1450 m, flowered ex hort. 20 August 2011 in Grahamstown, C. Craib s.n. (holotype GRA!, isotypes ABH! n. 59467, K!).

Deciduous bulbous plant up to 5 cm tall. Bulb solitary, hypogeal, ovoid, 1.5−3 × (0.5)1−2 cm, with soft pale brown outer tunics, terminating in a hypogeal neck, 2−3 × 0.3−0.5 cm. Roots fleshy, branched, white, up to 50 × 1−2 mm. Leaf solitary, ovate-oblong, dark green, flat on the ground, slightly convex, 3−7 × 1−2 cm, firm, leathery, with whitish-coloured thickened margins densely covered by short papillae (up to 0.05 mm), covered on the adaxial side with translucent, erect, stout, conical bristles of ca. 0.6−0.8 mm (collapsed, flattened and narrowly triangular when dry), glabrous on the abaxial side, clasping the stem at base, starting to wither to mostly withered at the anthesis. Inflorescence an erect raceme with (1)2−4(5) flowers, 1−2.5 cm long; lowermost pedicels 3−6 mm long, erect-patent; peduncle 1−2.5 cm long; bracts ovate-lanceolate to triangular, abruptly acuminate in the upper part, 4−7 × 2−4 mm, green with white membranous margins, withering soon and becoming papery white with brownish nerves, nearly as long as pedicels. Flowers suberect; tepals white with a green median stripe 1−1.5 mm wide, clearly visible in the abaxial side and undefined in the adaxial side associated to the central nerves, slightly fleshy, with minutely glandulous apex; outer tepals ovate-oblong, 10− 13 × 2.5−3.5 mm; inner tepals ovate, 8.5−10 × 3−4 mm. Stamens monomorphic; anthers ca. 1.3 × 0.7 mm; filaments strap-like, wider in the lower half, outers narrowly oblong, 6–7.5 × 1.5−2 mm; inners ovate oblong, 6.5−8 × 2−2.5 mm. Ovary ovate, green, 3.2−3.5 × 1.5−2.8 mm, trigonous in section, with two prominent longitudinal keels on each carpel; style filiform, 3−3.5 × 0.5−0.8 mm, stigma punctiform. Capsule broadly ovate to subglobose, ca. 5 × 5 mm, trigonous to subsphaerical in section, pale-brown when mature. Seeds flat, ca. 3 × 2.5 mm, dark brown to black, flattened and semidiscoidal (Figs. 1−2). Biology:—Trimelopter craibii remains dormant underground at the hottest time of the year, from November until early February. Leaves appear in autumn from early February, and become fully developed in April (Craib 2001b) and are covered with erect, short, white or ivory coloured bristles that impart a silvery or shiny appearance when the leaves are in strong direct sunlight. The leaf is cryptic and blends in very well with the rough and shiny black and grey dolomite pebbles and rocks which surround it (Craib 2001a). Bud development starts in June until early August, when heavy frosts occur. The flowering season lasts from the end of June to the middle of August, with a peak in late July and the beginning of August. Seeds develop from mid- to late August. Late August and early September are dry and windy creating ideal conditions for seed dispersal (Craib 2001b). The bulbs occur singly or in scattered groups of two to forty plants, rarely more. The species is plentiful at some localities particularly where there are pockets of deep soil that remain moist for longest after rainfall (Craib 2001c). Habitat:—Trimelopter craibii grows in the summer rainfall area of South Africa with most of the rainfall occurring between November and early April, in open short grassland in very hot and arid habitat with

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MARTÍNEZ-AZORÍN ET AL.

interesting associated succulent flora (Craib 2001c). It occurs in cracks in exposed dolomite sheets, shallow soil over sheets of exposed rock and patches of dolomite grit in depressions on bare rocks (Fig. 1) (Craib 2001a, 2001b). The endemic T. craibii is confined to vegetation classified as the Carletonville Dolomite Grassland (Mucina et al. 2006).

FIGURE 1. Trimelopter craibii Mart.-Azorín M.B. Crespo & A.P. Dold at the type locality (Trekdrift Farm, North West Province, South Africa). Photographs by Connall Oosterbroek.



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FIGURE 2. Trimelopter craibii Mart.-Azorín, M.B. Crespo & A.P. Dold. North West Province (holotype: Craib s.n. GRA). A. Flowering plant. B. Flower, frontal and dorsal views. C. Tepals and stamens, outer ones (left) and inner ones (right). D. Gynoecium, lateral views. E. Mature capsule with dry tepals; a valve capsule and seed. F. Bract. G. Leaf. Scales A–C, E, G: 1 cm; D, F: 5 mm.

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Distribution:—This species is only known from a single locality in the Lichtenburg district of the North West Province of South Africa, in an area of about 4 square kilometres (Charles Craib, pers. comm., July 2010) (Fig. 3).

FIGURE 3. Known distribution of Trimelopter craibii Mart.-Azorín, M.B. Crespo & A.P. Dold in South Africa.

Diagnostic characters and relationships:—Trimelopter craibii is easily identified by the solitary oblong convex leaf, with erect thickened bristles of 0.6−0.8 mm long only on the upper side, and thickened and densely papillate margin, the tepals 8.5−13 mm long, and the ovate ovary with scarcely prominent longitudinal keels (Figs. 1−2 & 4−5). These characters are not present as a whole in any known taxa of this genus. The new species appears to be closely related to T. dyeri but the latter shows the leaf with the upper surface and margins only minutely papillate, the tepals shorter (5−7 mm, up to 8.5 mm under cultivation; ABH! n. 58825), the ovary globose to obovate with larger and more prominent longitudinal keels, and the style shorter, ca. 2 mm long (Table 1, Figs. 4−5). Trimelopter unifolium shows some resemblance to T. craibii, though it differs by the leaf commonly longer (up to 12 cm), slightly concave, with margins covered with short laxly arranged papillae (neither whitish nor thickened as in the latter), and the larger and many flowered inflorescence (Table 1). In particular, some individuals of this species with hairy leaves, which were named Ornithogalum unifolium var. vestitum Müller-Doblies & Müller-Doblies (1996: 471), resemble in some extent T. craibii by the patent hairs on the upper side, but they are shorter and thinner, 0.05−0.4 mm (not 1 mm as cited in the protologue, cf. MüllerDoblies & Müller-Doblies 1996: 477), and evenly distributed (Table 1, Fig. 5). They are regarded here as a distinct variety T. unifolium var. vestitum (see the Appendix). Relationships between T. craibii and other species of the genus with hairy leaves are weaker. Trimelopter monarchos (Müller-Doblies & Müller-Doblies 1996: 476) Martínez-Azorín et al. (2011: 26) produce a much bigger leaf (15–33 cm long), with long and thin hairs of ca. 0.5−1 mm long on both sides of the leaf and margins, and a longer style, ca. 4−5 mm long. Furthermore, this species is only known from a single locality 7 km W of Steinkopf in the Northern Cape Province of South Africa. It is worth mentioning that the ovary of T. monarchos was described by Müller-Doblies & Müller-Doblies (1996) as being smooth and lacking ornamentation “ovario levi (haud sculptato).” However, plants collected at the type locality (ABH! n. 59600) showed constantly the ovary with the typical Trimelopter structure, it being deeply three-lobed, with six rounded angles, though in this case not markedly keeled.



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TABLE 1. Main diagnostic characters between Trimelopter craibii, T. dyeri s.str. and T. unifolium var. vestitum.

Bulb

Leaves

T. craibii

T. dyeri s.str.

T. unifolium var. vestitum

1.5−3 × (0.5)1−2 cm

1.8−2.5 × 1−2.2 cm

4.5−5.5 × 6−6.5 cm

hypogeal neck, 2−3 × 0.3−0.5 cm

hypogeal neck almost absent, up to 0.5 cm

hypogeal neck up to 3 cm

Leaf solitary, ovate-oblong, leathery, dark green, flat on the ground, slightly convex

Leaf solitary, ovate-oblong, leathery, dark green, flat on the ground, slightly convex

Leaf solitary, ovate-oblong, not leathery, green, flat on the ground, slightly concave

3−7 × 1−2 cm

2−3.5 × 0.6−1.7 cm

6−12 × 2−3.5 cm

Upper surface covered by translucent, erect, thickened, conical bristles of ca. 0.6−0.8 mm (flattened and narrowly triangular when dry)

Upper surface minutely papillate (only visible under a lens)

Upper surface with patent hairs 0.05−0.4 mm evenly distributed on the upper side

Leaf margin densely papillate giving a whitish colour

Mostly green and fleshy when flowers are present

Mostly withered when flowers are present

(1)2−4(5) flowers

2−8 flowers

9−40 flowers

raceme 1−2.5 cm

raceme 1−3 cm (up to 5 in cultivation)

raceme 7−10 cm

peduncle 1−2.5 cm long

peduncle 0.5−1 cm long

peduncle 4.5−6 cm

outer tepals ovate-oblong, 10−13 × 2.5−3.5 mm

outer tepals ovate-oblong, 6−7 × 2.5−3.5 mm

outer tepals oblong, 9−10 × 2.7−3 mm

inner tepals ovate, 8.5−10 × 3−4 mm

inner tepals ovate, 5−6 × 2.5−3 mm

inner tepals narrowly ovate, 7−8 × 2.3−2.7 mm

6−8 × 1.5−2.5 mm

4−5 × 1.2−2 mm

5−6 × 1.5−2 mm

ovate

widely obovate to subglobose

widely obovate to subglobose

3.2−3.5 × 1.5−2.8 mm

2−2.5 × 2.2−2.5 mm

3−3.2 × 2.4−2.6 mm

With two scarcely prominent longitudinal keels on each carpel

With two prominent longitudinal keels on each carpel

With two white prominent longitudinal keels on each carpel

3−3.5 mm

1.8−2 mm

3−3.2 mm

Leaf margin minutely papillate and of similar colour, not separated or thickened

Leaf margin with short papilliform hairs

Starting to wither to mostly withered when flowers are present Inflorescence

Tepals

Filaments Ovary

Style

The Namibian Trimelopter etesiogaripense (U.Müll.-Doblies & D.Müll.-Doblies) Martínez-Azorín et al. (2011: 26) shows a canaliculated, narrowly lanceolate leaf, with an involute tip, with rows of stiff hairs or papillae of 0.3−3 mm long on both sides and margins. It is only known in winter-rainfall Namibia along the mountains at the eastern border of the Namib, from Dikwillem in the north to the Orange River near Rosh Pinah and to the Richtersveld in the Cape (cf. Müller-Doblies & Müller-Doblies 1996). A related species, Trimelopter strigosulum (Müller-Doblies & Müller-Doblies 1996: 480) Martínez-Azorín et al. (2011: 26), has a narrowly ovate leaf, 2.5−3.5 cm long (up to 8 cm in cultivation), densely covered with adpressed hairs of 0.5−1 mm long, which are somewhat medifixed with a very short retrorse arm. The indumentum in this species seems to be unique in the genus. It is also restricted to SW Namibia, near Pockenbank (cf. MüllerDoblies & Müller-Doblies 1996). Etymology:—Named after botanist Charles Craib, author of several botanical books and numerous articles, who passed away in March 2012. Together with Trimelopter craibii he is commemorated in Aloe craibii Smith (2003: 26) and Ceropegia craibii Victor (2001: 212).

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Observations:—Ornithogalum dyeri var. leistneri Müller-Doblies & Müller-Doblies (1996: 476) was described and illustrated from a single herbarium specimen (Holotype: Leistner 849 KMG) collected near Postmasburg, Northern Cape Province of South Africa. The authors related that plant to Trimelopter dyeri, a species with a glabrous, convex leaf, but noted that “the indument is so striking that it deserves taxonomic recognition. Parallel to O. unifolium var. vestitum the rank of variety was chosen”. Unfortunately, the holotype of that variety remains on loan to U. and D. Müller-Doblies since 1980 (A. van Heerden, pers. comm.) and therefore we are unable to study it. However, the illustration presented by Müller-Doblies & Müller-Doblies (1996) and the very brief description suggests that T. craibii and O. dyeri var. leistneri are closely related. Further studies are needed to elucidate this hypothesis, including studying living plants of Ornithogalum dyeri var. leistneri from the type locality near Postmasburg.

FIGURE 4. Flowers and gynoecia of Trimelopter. A. T. craibii Mart.-Azorín, M.B. Crespo & A.P. Dold from the type locality: South Africa, North West Province, Trekdrift Farm, C. Craib s.n. (GRA Holo.; ABH59467, K Iso.). B. T. dyeri (Poelln.) Mart.-Azorín, M.B.Crespo & Juan from the type locality: South Africa, Eastern Cape, N of Grahamstown, Table Farm, Martínez-Azorín et al. MMA486 (GRA, ABH58825). C. T. unifolium var. vestitum from the type locality: South Africa, Western Cape, E of Matjiesfontein, near Laingsburg, Martínez-Azorín et al. MMA847 (GRA, ABH59634). Scale bars: flowers = 1 cm, gynoecia = 2 mm.

Appendix Trimelopter unifolium is currently considered to include plants with some variation in leaf indumentum, which were described as varieties of Ornithogalum unifolium (cf. Müller-Doblies & Müller-Doblies 1996). The typical individuals (var. unifolium) show leaves glabrous or minutely hairy on the upper surface under magnification, and shortly papillate-ciliate margins. However, two populations from the surroundings of Matjiesfontein, near Laingsburg, include plants with the upper surface of leaves covered by short, patent hairs



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up to 0.4 mm, evenly distributed and visible to the naked eye, which correspond to var. vestitum (cf. MüllerDoblies & Müller-Doblies 1996). Individuals of both varieties live together in the Witwaterspoort population according to Müller-Doblies & Müller-Doblies (1996). Furthermore, we also found the same variation pattern in the type locality (Martínez-Azorín et al. 847, ABH! n. 59634), and therefore we regard both taxa as extremes of variation for which the varietal rank is here accepted. The following new combination is hence needed to accommodate it in Trimelopter.

FIGURE 5. Leaf details of Trimelopter. A. T. craibii Mart.-Azorín, M.B. Crespo & A.P. Dold from the type locality: South Africa, North West Province, Trekdrift Farm, C. Craib s.n. (GRA Holo.; ABH59467, K Iso.). B. T. dyeri (Poelln.) Mart.-Azorín, M.B.Crespo & Juan from the type locality: South Africa, Eastern Cape, N of Grahamstown, Table Hill Farm, Martínez-Azorín et al. MMA486 (GRA, ABH58825). C. T. unifolium var. vestitum from the type locality: South Africa, Western Cape, 800 m from first gate to Whitehill, E of Matjiesfontein, near Laingsburg, Martínez-Azorín et al. MMA847 (GRA, ABH59634).

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Trimelopter unifolium var. vestitum (U.Müll.-Doblies & D.Müll.-Doblies) Mart.-Azorín, M.B.Crespo & A.P.Dold, comb. nov. Basionym:—Ornithogalum unifolium var. vestitum Müll.-Doblies & Müll.-Doblies, Feddes Repertorium 107(5-6): 471, 1996 (Müller-Doblies & Müller-Doblies 1996).

Acknowledgements This work was partly supported by the Fundación Ramón Areces (Spain) and Universidad de Alicante (Spain). Rhodes University provided working facilities from October 2009 and November 2011.We thank the curators of the herbaria who provided access to specimens examined. Connall Oosterbroek is also thanked for sending photographs of Trimelopter craibii at the type locality. The Department of Environment and Nature Conservation of Northern Cape Province and CapeNature of Western Cape Province provided permission to collect herbarium specimens (collecting permits numbers FLORA069/2011, and AAA008-00031-0028 respectively).

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