Three New Species of Metalasia (Asteraceae–Gnaphalieae) from the Western Cape and an Updated Key to the Genus

South African Journal of Botany 84 (2013) 72–82

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South African Journal of Botany

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Short communication Three new species of Metalasia (Asteraceae–Gnaphalieae) from the Western Cape and an updated key to the genus

A. Bengtson a,⁎, A.A. Anderberg b, P.O. Karis a a Department of Botany, Stockholm University, SE-106 91 Stockholm, Sweden b Department of Phanerogamic Botany, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05 Stockholm, Sweden article info abstract

Article history: Three new species of Metalasia from the Western Cape of South Africa are described; Metalasia tristis Received 4 July 2012 A.Bengtson & P.O.Karis, a small shrublet from the Groot Winterhoek Mountains, and Metalasia formosa Received in revised form 20 September 2012 A.Bengtson & P.O.Karis and Metalasia eburnea A.Bengtson & P.O.Karis both found in the Langeberg Mountains. Accepted 20 September 2012 The morphology and relationships of the three new species are discussed, and an updated key to the genus is Available online 8 November 2012 provided. Edited by JS Boatwright © 2012 SAAB. Published by Elsevier B.V. All rights reserved.

Keywords: Asteraceae Gnaphalieae Metalasia New species Taxonomy South Africa

1. Introduction prep.). The three species are here formalized together with an amended and updated key to the genus Metalasia. The genus Metalasia R.Br. of the tribe Gnaphalieae, Asteraceae, currently consists of 54 species (Karis, 1989; Karis and Helme, 2. Material and methods 2012), with their main distribution in the Greater Cape Floristic Region of South Africa (GCFR; Born et al., 2007) where most are The descriptions and illustrations were based on the type specimens found growing in fynbos vegetation. During recent ﬁeldwork deposited in the Swedish Museum of Natural History, Section of Metalasia material not corresponding to any of the currently recog- Phanerogamic Botany, S, as well as the additional material listed. Mate- nized taxa was collected. After careful examination it was concluded rial was softened in water, dissected, and mounted in Hoyer's solution that the material represents three morphologically distinct hitherto for microscope studies. undescribed species. One species is an erect shrub found in dry fynbos vegetation on the slopes of Tuinskloof near Touwsrivier and in 3. Species descriptions Langeberg sandstone fynbos, the next species is a small shrublet from the Groot Winterhoek Mountains east of Piketberg, and the 3.1. Metalasia tristis A.Bengtson & P.O.Karis, sp. nov third species is a shrub found in Montagu shale renosterveld on the Little Karoo side of the Langeberg (Fig. 1). Recent molecular studies Type: South Africa. Western Cape, Wuppertal (3219): NE of have shown the genus Metalasia to be divided into two clades, mor- Porterville, Farm Waterval, beside walking track above waterfall, phologically separated by the presence of papillose cypselas in one 801 m, (−CC), 5 September 2006, Stångberg, Karis & McKenzie 37 of the clades, the “Metalasia densa group” (Bengtson et al., 2011; (S, holo.; BOL, NBG, PRE iso). Bengtson et al., in prep.). Two of the new species, Metalasia formosa Decumbent, sparingly to well-branched shrublet, up to 0.3 m and M. eburnea, share the distinct papillose cypselas of the high. Branches sprawling to erect, densely whitish tomentose when “M. densa group”, whereas the third, M. tristis, is placed within the young, becoming less tomentose to glabrescent, axillary brachyblasts morphologically more diverse of the two clades (Bengtson et al., in rarely present, less than half as long as subtending leaf, foliage dense to sparse. Leaves involute-ericoid, somewhat twisted, narrowly lance- – – ⁎ Corresponding author. Tel.: +46 08 16 2071. olate 5 10 (14)×0.8 1.2 mm, densely tomentose above, sparsely E-mail address: [email protected] (A. Bengtson). hairy below, apically acute with tips straight to declinate, spreading

0254-6299/$ – see front matter © 2012 SAAB. Published by Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.sajb.2012.09.008 A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82 73

A B C

E F

Fig. 1. Metalasia eburnea (A, D); M. formosa (B, C); M. tristis (E, F). (Photos: A and D by Linda Lundmark; B and F by Annika Bengtson; E by Frida Stångberg and C by Marcus Arnerup).

to declinate, straight to slightly curved. Synflorescences 5–12 mm (Fig. 2D, E). Metalasia tristis is most similar to M. divergens (Thunb.) wide, dense, broadly campanulate, with 6 to 15 capitula arranged in D.Don subsp. divergens due to the dense hairy capitula clusters with obscure, hardly distinguishable clusters fused by dense hairs, over- 5-flowered capitula, but M. divergens subsp. divergens has larger, shot by a few bract-like leaves. Capitula 5-flowered, fused by hairs more easily distinguishable capitula and involucral bracts with within clusters and synflorescences. Involucral bracts±13, of±equal glands. Furthermore, Metalasia divergens subsp. divergens is endemic length, erect, densely tomentose for 3/4–4/5 of length, only upper- to the Cape Peninsula. Metalasia serrulata P.O.Karis and M. rogersii most parts visible through the dense tomentum, with broad hyaline S.Moore may resemble M. tristis in habit but both differ in having margins; outermost bracts lanceolate, foliaceous with apical scarious synflorescences with 3-flowered capitula where the outermost lamina, narrow, acute, dark brown; outer bracts lanceolate, apical involucral bracts are considerably shorter than the innermost, and scarious lamina narrow, acute, dark brown; inner bracts narrowly in having rounded pink, petaloid inner involucral bracts. Metalasia oblong to lanceolate gradually becoming somewhat more petaloid, rogersii also differs in that is has much smaller, mostly declinate apical scarious lamina, acute, brown; innermost bracts narrowly ob- leaves with axillary brachyblasts. Metalasia serrulata and M. rogersii long, apically rounded, almost petaloid with a light brown to whitish not only resemble M. tristis, but the three also occur in close lamina. Corolla narrowly cylindrical, 2.8–3.1 mm long, dark purple proximity in the Groot Winterhoek Mountains, Witsenberge, and distally. Cypselas 3-veined (immature). Pappus bristles slightly serrate, Skurweberge. In the molecular study of Bengtson et al. (in prep.) apically clavate with fused cells. (Figs. 1E, F and 2). Metalasia tristis is placed within the “M. divergens group” within the larger and more diverse of the two main clades of Metalasia. 3.1.1. Distribution and ecology Metalasia tristis is known from only two localities on Voorberg in 3.1.3. Additional specimens examined the Groot Winterhoek Mountains near Porterville where it is found South Africa. Western Cape: 3319 (Worcester): E of Porterville, on mountain slopes and plateaus in Winterhoek sandstone fynbos slopes above Farm Dasbos, 791 m, (−CC), 6 Sep 2006, Stångberg, vegetation on well-drained rocky soil (Mucina et al., 2005; Mucina Karis & McKenzie 42 (BOL, K, P, PRE, S); E of Porterville, above Farm and Rutherford, 2006; Fig. 5). Flowering time is from September to Dasbos, 947 m, (−CC), 6 Sep 2006, Stångberg, Karis & McKenzie 44 October. (NBG, S). 3.1.2. Diagnosis and relationships Metalasia tristis is distinguished by its sprawling, decumbent 3.2. Metalasia formosa A.Bengtson & P.O.Karis, sp. nov dwarf habit (Fig. 1E), the tomentose young branches, leaves and synflorescences (Fig. 1F), the synflorescences with capitula fused by Type: South Africa. Western Cape, Montagu (3320): Tuinskloof, whitish hairs (Fig. 2D) and by the involucral bracts with acute, slopes around small valley around farm, 1050 m, (−AA), 12 September narrow scarious, brown laminas visible between the white hairs 2005, Karis & Arnerup 1060 (S holo.; NBG iso). 74 A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82

Fig. 2. Metalasia tristis, Stångberg et al. 37 (S). A, portion of plant; B, branch with leaves; C, two leaves; D, cluster of capitula fused by dense hairs; E, all involucral bracts from one capitulum, from outermost (upper left) to innermost (lower right); F, pappus bristle with tip magniﬁed (left). Artist: Emma Hultén.

Erect, often stout shrub, up to 1.5 m tall. Branches erect, whitish ca. half as long as innermost, erect, acute to mucronate, tips erect or tomentose when young, sometimes with reddish colour, older slightly bent outwards; outermost bracts narrowly lanceolate, acute, becoming less tomentose, with cracking cortex, and the tomentum scarious, brown; outer bracts oblanceolate, acute, scarious brown or peeling off, axillary brachyblasts more than half as long as the sometimes reddish on upper parts, becoming more petaloid inwards; subtending leaf and sometimes elongated into 1.0–1.5 cm long inner bracts oblanceolate, mucronate, subpetaloid, pink or light shoots, leaves persistent for 3 years, older braches with leaf scars, brownish in colour; innermost oblanceolate to oblong, acute to mu- foliage dense. Leaves involute-ericoid, straight to slightly twisted, nar- cronate, petaloid, white. Corolla narrowly cylindrical, 4–5 mm long, rowly lanceolate, 4–7×0.8–1.1 mm, white tomentose above, glabrous purple distally. Cypselas oblong, 1.8–2.1 mm long, 3-veined, brown, beneath, acute, mucronate, tips straight or declinate, straight to papillose. Pappus bristles serrate, apically clavate. (Figs. 1B, C and 3). slightly curved, spreading. Synﬂorescences 20–30 mm wide, dense, conspicuously branched with capitula arranged in clusters of 5 to 14 3.2.1. Distribution and ecology capitula. Peduncles (0)2–6 mm long. Capitula 5-ﬂowered, somewhat Metalasia formosa is only known from a few localities. It occurs on fused by hairs at the base within clusters. Involucre narrowly valley slopes in Tuinskloof near Touwsrivier, where it is found in cyathiform, apically 2.0–2.5 mm wide. Involucral bracts ±15 in 4 to dry Matjiesfontein quartzite fynbos vegetation, but also in the 5 series, hairy for lower 2/3, gradually longer inwards; outermost Langeberg, growing in Langeberg sandstone fynbos, where it in many A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82 75

Fig. 3. Metalasia formosa, Karis & Arnerup 1060 (S). A, portion of plant; B, branch with leaves; C, leaf from above and from side; D, capitulum; E, all involucral bracts from one capitulum, from outermost (upper left) to innermost (lower right); F, cypselas; G, pappus bristle with tip magniﬁed (left). Artist: Emma Hultén.

places is sympatric with Metalasia densa (Mucina et al., 2005; Mucina carinate leaves. Metalasia formosa may also resemble the common and Rutherford, 2006; Fig. 5). Flowering time is from September to and variable M. densa (Lam.) P.O.Karis, but the two species differ October. greatly in involucre which is narrow with bracts in 6–8 series, with brown outer bracts contrasting against the white innermost bracts 3.2.2. Diagnosis and relationships in M. densa. In the molecular phylogenetic study of Bengtson et al. Metalasia formosa is recognized by its almost straight, rather than (in prep.) Metalasia formosa is placed within the “M. densa group” twisted, leaves with elongated axillary brachyblasts (Fig. 3B, C), (Bengtson et al., 2011; Karis, 1989). This is supported morphological- branched synflorescences with 5-flowered capitula in clusters, with ly by the papillose cypselas which distinguish the “M. densa group” sometimes pink inner and white innermost involucral bracts from the remaining species of Metalasia. (Figs. 1B and 3A, D). Metalasia formosa resembles M. trivialis P.O.Karis, which are erect shrubs with large axillary brachyblasts, branched 3.2.3. Additional specimens examined synflorescences with capitula in clusters and involucres with white South Africa. Western Cape: 3320 (Montagu): Tuinskloof, slopes tomentose brown outer and white petaloid inner bracts. However, around small valley around farm, 1090 m, (−AA), 12 Sep 2005, Metalasia trivialis differs in having 3 or 4-flowered capitula with Karis & Arnerup 1057 (S); Tuinskloof, slopes around small valley white inner bracts and half-twisted, rather than straight, often around farm, 1070 m, (−AA), 12 Sep 2005, Karis & Arnerup 1059 76 A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82

(BOL, PRE, S); Tuinskloof, slopes around small valley around farm, capitula. Peduncles 5–30 mm long. Capitula 3-ﬂowered, free from one 1040 m (−AA), 12 Sep 2005, Karis & Arnerup 1061 (K, P, S); another. Involucre narrowly cyathiform, apically 1.0–1.5 mm wide. Langeberg, walking track above farm Ravenna, 670 m, (−CD), 10 Involucral bracts±20 in 5 to 6 series, erect, gradually longer inwards, Sep 2006, Stångberg & Karis 49 (S); Langeberg, farm Kruispad, outermost about 1/4 of length of innermost, tomentose for lower 2/3, 650 m, (−CD), 10 Sep 2006, Stångberg & Karis 52 (S). all but innermost interwoven by dense woolly hairs, with hyaline margins, apically mucronate to acute; outermost and outer bracts ovate to oblong, thin, scarious, light brownish beige to cream coloured; inner 3.3. Metalasia eburnea A.Bengtson & P.O.Karis, sp. nov bracts oblong to oblanceolate, apically concave, subpetaloid to petaloid, white; innermost bracts oblanceolate, glabrous, apically plicate, Type: South Africa. Western Cape, Montagu (3320): Langeberg, petaloid, white. Corolla narrowly cylindrical, 2.8–3.2 mm long, purple slopes above Kruispad, ca. 20 km W of Barrydale, 620 m, (−CD), 12 distally. Cypselas oblong, 1.5–1.8 mm long, 3-veined, brown, papillose. March 2006, Karis, Lundmark, Stångberg & Zoubir 1068 (S holo.; BOL, Pappus bristles serrate, apically clavate. (Figs. 1A, D and 4). NBG, K, PRE iso.). Erect shrubs, up to 2 m tall. Branches erect, densely whitish- 3.3.1. Distribution and ecology tomentose when young, older branches becoming less tomentose to Metalasia eburnea is only known from a few localities on the Little glabrous, with axillary brachyblasts less than half as long as subtending Karoo side of the Langeberg where it is found on lower northern leaf, leaves persistent for 2 to 3 years, branches with or without distinct slopes, growing in Montagu shale renosterveld (Mucina et al., 2005; leaf scars, foliage dense to sparse. Leaves involute-ericoid, half-twisted, Mucina and Rutherford, 2006; Fig. 5). Flowering occurs in March. narrow, oblong to oblanceolate, 7–11×0.8–1.1 mm, acute, mucronate with tips straight or sometimes declinate, white tomentose above, glabrous beneath, spreading, straight or slightly curved. Synﬂorescences 3.3.2. Diagnosis and relationships 20–70 mm wide, dense, conspicuously branched, with numerous capit- Metalasia eburnea is recognized by having long, narrow, twisted ula arranged in branched clusters, each cluster containing few to many leaves and axillary brachyblasts less than half as long as the subtending

Fig. 4. Metalasia eburnea, Karis et al. 1068 (S). A, portion of plant; B, branch with leaves; C, three leaves; D, capitulum; E, all involucral bracts from one capitulum, from outermost (upper left) to innermost (lower right); F, cypselas; G, pappus bristle with tip magniﬁed (left). Artist: Emma Hultén. A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82 77

Fig. 5. Known distributions of Metalasia tristis (●); M. formosa (■); and M. eburnea (○).

leaf (Fig. 4B, C), synflorescences with many small, narrow, 3-flowered 3. Capitula 3(–4) flowered…………………………………………….4 capitula with light beige outer, and white inner involucral bracts in 3. Capitula with 5 or more flowers….…………………..……….….18 branched clusters (Figs. 1Aand4A, D). Metalasia eburnea is most sim- 4. Capitula free from one another……….………………………...…..5 ilar to the variable M. densa, the latter however differs in that the capit- 4. Capitula fused by hairs to one another into clusters or into heads ula have a darker involucre that is 1.5–3.0 mm wide, bracts in 6–8 of clusters…………………………………….……….……….………..12 series, and brown, scarious outer bracts. Metalasia eburnea may also re- 5. Tips of most involucral bracts revolute to spreading….……M. dregeana semble M. muricata (L.) D.Don, but M. muricata has curved to hooked …………………… ……… leaves and 4 or 5 flowered capitula. Metalasia acuta P.O.Karis, which 5. Tips of most involucral bracts±erect .. 6 is another renosterveld species, is easily distinguished from M. eburnea 6. Leaves very acute, sharply pungent; capitula with hairs only at by its sharply pungent leaves and narrow lanceolate to subulate, very the base involucral bracts acute to very acute; inner and inner- acute involucral bracts. In the molecular phylogenetic study of most apically serrate……………………….……….……..…..M. acuta Bengtson et al. (in prep.), this species is placed within the “M. densa 6. Leaves not or only somewhat pungent; capitula with hairs group”, morphologically supported by the shared presence of papillose reaching at least up to 1/3 of total length; involucral bracts acute fi cypselas (Bengtson et al., 2011). The species was rst discovered in the to rounded; inner dentate to apiculate, not serrate……..…....…....7 Langeberg in 2006 (Karis, Lundmark, Stångberg & Zoubir 1068), however 7. Leaves declinate, less that 6 mm long, ± straight……………….8 when the authors visited the location in September 2011 the area had 7. Leaves spreading to erect-spreading, all or at least some over recently burned and so the plants were no longer present. One 6 mm long and curved outwards………………………..………….10 additional specimen was later found in the herbarium of the Swedish – Museum of Natural History (S), erroneously filed under Metalasia densa. 8. Branches conspicuously and persistently tomentose for 3 4 seasons; inner and innermost involucral bracts apically±flat; pappus bristles acute………………………………………M. albescens 3.3.3. Additional specimens examined 8. Branches deciduously tomentose to glabrous; inner and South Africa. Western Cape : 3319 (Worcester): Farm Doringkloof, innermost involucral bracts apically concave, pappus bristles southern foothills of Voetpadsberg, ca 2000 feet, (-DB), 2 Mar 1985, …………………………………………………………… Morley 344 (S). mostly obtuse 9 9. Shrublet or shrub, erect; branches often erect; pappus bristles 4. Key to Metalasia (updated from Karis, 1989) narrow……………………………………………………….M. brevifolia 9. Shrublet, decumbent, sprawling; branches creeping to ascend- 1. Petaloid involucral bracts white to whitish...……….….….…………2 ing; pappus bristles comparatively broad……….……..….M. tenuis 1. Petaloid involucral bracts pale to bright pink, or yellowish to 10. Outermost and outer involucral bracts reddish; much bright yellow..……………………….…………………….……..……..79 branched low shrub……………………………………M. namaquana 2. Most or all involucral bracts petaloid……..…………………..………3 10. Outermost and outer involucral bracts brown or foliaceous; 2. Most involucral bracts not petaloid………………………………37 sprawling or robust shrub...…………………………………………..11 78 A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82

11. Leaves erect-spreading; synflorescences 10–50 mm wide, with 23. Young branches often yellowish tomentose; involucral bracts 10 to numerous capitula; involucral bracts in 5–7 series; inner- gradually shorter inwards…………………………………..M. inversa most erect, white……………………….……….……….M. erubescens 23. Young branches whitish tomentose; involucral bracts of simi- 11. Leaves spreading; synflorescences 5–10 (−13) mm wide, with lar length, or longer inwards…………………………………….….24 few to 10 (−17) capitula; involucral bracts in 4–5 series, innermost 24. Robust erect shrub, mostly 1–2.5 m high; without glands; often curved outwards, white to creamy white….….….M. tenuifolia leaves conspicuously carinate; synflorescences 25–60 mm wide 12. Leaves on flowering shoots erect, imbricate, without axillary ……………………………………………………….……M. muraltiifolia brachyblasts, but on older branches declinate to spreading, with 24. Shrublets, usually not exceeding 1 m; glands present on young axillary brachyblasts…………….……..………..…….…….M. capitata branches, leaves and involucral bracts; leaves not conspicuously 12. Leaves on flowering shoots and older branches similar….…13 carinate; synflorescences 7–30 mm wide….……………….……..25 13. Leaves declinate, less than 6 mm long, up to 0.8 mm wide, 25. Leaves often declinate or sometimes erect-spreading; capitula with axillary brachyblasts……………………..…………………..….14 clusters generally not distinguishable; tips of involucral bracts± 13. Leaves spreading to erect, at least some more than 6 mm long and conspicuously imbricate, innermost generally flat apically; pappus 1 mm wide, with or without axillary brachyblasts…………..….…….15 bristles often flattened to flat apically…….……….….M. cephalotes 14. Synflorescences very dense, seemingly unbranched, mostly 25. Leaves often spreading or sometimes erect-spreading; more than 10 mm wide, with leaves appressed underneath; capitula clusters generally distinguishable; tips of involucral outermost involucral bracts foliaceous with a scarious lamina, ± bracts reaching to a similar height, innermost conspicuously spathulate..……………………………………………………M. rogersii plicate apically; pappus bristles often subterete apically 14. Synflorescences dense, obscurely branched to slightly .…………………………………………….….………………M. pulchella branched, often less than 10mm wide, without leaves 26. Leaves twisted………………………....…………………………..27 appressed underneath; outermost involucral bracts foliaceous 26. Leaves non-twisted…………………………………….…………29 to subfoliaceous without a lamina, narrowly lanceolate, acute 27. Involucral bracts apically erect; cypselas with 3–5 conspicuous ……..……………………..…………………………………..…M. tenuis ribs……………………………………..…………………….M. fastigiata 15. Flowering shoots greenish, soft; synflorescences 15–60 mm 27. Several involucral bracts apically revolute to erect spreading; wide, containing 5–15 pedicelled dense clusters, clusters cypselas without ribs……………………………………………….…28 free from each other; cypselas almost spheroid, whitish 28. Leaves densely greyish-tomentose, narrowly obovate to oblan- papillose………………….……………….………………...…..M. riparia ceolate, mostly without axillary brachyblasts; synflorescences 15. Flowering shoots whitish or reddish, more woody; synflorescences with few to 10 (−15) capitula……………………………M. bodkinii head-like, up to 25 mm wide, dense to very dense; clusters fused by 28. Leaves glabrous to sparsely pubescent, linear to narrowly ob- hairs; cypselas longer than wide, glabrous……………………..……..16 lanceolate, without or with axillary brachyblasts; synflorescences 16. Flowering shoots and synflorescences reddish-white-tomentose; with few to 30 capitula………………………………..M. quinqueflora leaves densely tomentose, greyish green, conspicuously decreasing in 29. Leaves±straight, often narrowly triangular, widened and flat size towards the synflorescence…….……….……..……M. rhoderoides at the base, with axillary brachyblasts; cypselas with ca. 5 conspic- 16. Flowering shoots and synflorescences whitish-tomentose; uous ribs…………………………………………...….M. agathosmoides leaves glabrous to sparsely tomentose, of the same size towards 29. Leaves±distinctly curved, or straight but then never triangu- the synflorescence………….……………………………...…..………17 lar, linear to narrowly ovate, only rarely with inconspicuous axil- 17. Leaves straight to incurvate, with or without axillary brachyblasts; lary brachyblasts; cypselas with or without two lateral and one synflorescences dense and±branched, with pedunculate clusters of medial whitish ribs ………………………………………………..….30 fused capitula, 10–15 mm wide; pappus bristles with a serrate 30. Leaves distinctly curved outwards, apically hooked…M. adunca shaft…………………………………….……….…….…….…M. cymbifolia 30.Leaves curved inwards to±straight, not hooked apically 17. Leaves±straight, often without axillary brachyblasts; …………………….……………….……………….…………..……...31 synflorescences very dense, seemingly unbranched, 3-11 mm wide; 31. All capitula individually arranged, or occasionally a few pappus bristles serrulate to denticulate throughout..………M. serrulata clustered; involucral bracts apically erect…………………M. pallida 18. Capitula 5-flowered……………………………………………….19 31. Most to all capitula arranged in clusters within the 18. Capitula at least 6-flowered……………………………………..33 synflorescence; at least some involucral bracts apically revolute 19. Capitula completely fused by hairs within clusters of heads of to erect-spreading………………………………….....……………….32 clusters……………………………………………………..……..…….20 32. Leaves often more than 1 mm wide, narrowly ovate to narrowly 19. Capitula free from one another, though sometimes arranged in lanceolate; involucre narrowly cyathiform; outer involucral bracts dense head-like synflorescences .…………………………………..26 often erect-spreading and whitish, inner erect-spreading to erect, 20. Outermost involucral bracts brown to dark brown………….21 conspicuously carinate apically…………….M. pulcherrima f. pallescens 20. Outermost involucral bracts white, whitish or reddish ………….22 32. Leaves 0.5–1 mm wide, linear to narrowly lanceolate; 21. Robust, erect shrub; glands never present; branches erect and involucre narrowly oblong-urceolate; outer involucral bracts comparatively thick; leaves mostly spreading, distinctly carinate; revolute to erect-spreading and dark brown to reddish to synflorescences 25–60 mm wide……….………..…..M. muraltiifolia whitish, inner revolute to erect-spreading and slightly carinate 21. Decumbent to sprawling shrublet; leaves, young branches apically…………………..………………..…………….……M. galpinii and synflorescences with glands; branches usually creeping to as- 33. Leaves non-twisted……………………………………………….34 cending, slender; leaves often declinate, never distinctly carinate; 33. Leaves twisted……………………………………………………..35 synflorescences 4–15 mm wide……………M. divergens ssp. divergens 34. Synflorescences with 10 to numerous capitula; all capitula in- 22. Pappus bristles 3 per flower; cypselas scabrid………M. confusa dividually arranged and distinctly pedicelled, or, occasionally a 22. Pappus bristles at least 10 per flower; cypselas glabrous……23 few clustered; involucral bracts in 6–9 series………….…M. pallida A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82 79

34. Synflorescences with few to ca. 12 capitula; at least some 48. Leaves with axillary brachyblasts.…………………………..…..49 capitula arranged in clusters and often indistinctly pedicelled; 48. Leaves without axillary brachyblasts……………………….….50 involucral bracts in 4–6 series…………….………………..M. alfredii 49. Branches conspicuously tomentose for at least 3–4 seasons; 35. Capitula 8–13-flowered; petaloid involucral bracts apically leaves broadest above the middle; innermost involucral revolute to erect-spreading.………………………….M. lichtensteinii bracts truncate, apically unevenly dentate to emarginated 35. Capitula 6-flowered; petaloid involucral bracts apically .………….….…………………………………………….……M. calcicola erect-spreading to erect.…………………………………..………….36 49. Branches±glabrous after 3–4 seasons; leaves generally 36. Leaves widest at the base, distinctly carinate; at least some broadest toward the base; innermost involucral bracts acute to synflorescences with more than 15 capitula……..……M. compacta rounded, rarely truncate……………………………………M. pungens 36. Leaves widest above the middle, not carinate; synflorescences 50. Leaves mostly appressed; capitula 1–1.5 mm wide with up to 12 capitula…………………………….….……….M. alfredii …………………….……………….……………….…….….M. erectifolia 37. Capitula 3 (−4)-flowered.……………………………………….38 50. Leaves erect-spreading to erect, but not appressed; capitula 37. Capitula at least 5-flowered.………………………………..……51 1.5–2.5 mm wide……………………………………..…..…M. pungens 38. Leaves with a long mucronate tip, sharply pungent 51. Synflorescences with up to 15 capitula…………………………52 .….….…..……………………………………………………….………..39 51. At least some synflorescences with more than 15 38. Leaves shortly mucronate, not sharply pungent, though acute capitula………………………………………………...…….………...... 69 to acuminate.….…………………….…………………………………41 52. Leaves non-twisted, or almost non-twisted.………..………….53 39. Tips of involucral bracts mostly revolute to spreading 52. Leaves twisted.………....…………………………………………..59 .….….…..………………………………………………………M. dregeana 53. Leaves without axillary brachyblasts…………………………..54 39. Tips of involucral bracts mostly erect.…………………………..40 53. Leaves with axillary brachyblasts……………………………….57 40. Axillary brachyblasts generally less than half as long as the 54. Leaves broadest above the middle; young leaves densely subtending leaf; outermost involucral bracts 1/3–1/2 as long as pubescent, at least towards the base; cypselas without ribs or the innermost, innermost acute to very acute, apically coarsely ridges..………………………………………………………….M. alfredii serrate; pappus bristles acute to very acute.……….………..M. acuta 54. Leaves parallel-sided or broadest towards the base; 40. Axillary brachyblasts generally more than half as long as the young leaves mostly glabrous; cypselas ribbed or ridged subtending leaf; outermost involucral bracts 1/2–3/4 as long as ……………………………………………………………………..…....55 the innermost, innermost obtuse to acute, apically with entire 55. Leaves±straight, narrowly triangular, mucronate with tips margins; pappus bristles mostly obtuse…………………..M. trivialis straight to declinate..…………………………………………M. helmei 41. Most leaves twisted……………………………..…………..…….42 55. Leaves curved, with reflexive tips or apically hooked ….……….56 41. Leaves non-twisted………………..…..…………………………..48 56. Leaves curved inwards, with reflexive tips; most or all 42. Outermost involucral bracts at least half as long as the capitula individually arranged; cypselas with three ribs, oftenwith innermost………………………………...……………………………..43 short papillae.….…………………..…………………….……M. pallida 42. Outermost involucral bracts less than half as long as the 56. Leaves curved outwards, apically hooked; capitula arranged innermost……………………………………………..……………...... 45 in clusters in the synflorescence; cypselas ridged, glabrous 43. Erect shrub, up to 3 m, rarely compact; leaves carinate …………………………………………………….………….M. adunca ………….……………….……………….……..….……….….M. trivialis 57. Capitula 6–8-flowered; involucral bracts in 7–12 series 43. Low sometimes straggling shrublets, not exceeding 0.3 m; ………………………………………………………………..M. tricolor leaves not carinate.……………………………………………………44 57. Capitula 5(–6)-flowered; involucral bracts in 4–6series…58 44. Leaves generally slightly involute, usually with axillary 58. Branches becoming±glabrous after 2–3 seasons; young leaves brachyblasts; synflorescences 5–20 mm wide, with few to 30 (nu- spreading to erect-spreading, pubescent but without long, coarse merous) capitula; capitula 2.5–3.5 mm wide; petaloid involucral hairs, and soon becoming±glabrous, usually broadest above the bracts white; cypselas without an annular ridge, with short acute middle; innermost involucral bracts acute or rarely obtuse; pappus papillae………………………………………………………M. montana bristles subterete but not clavate towards the very apex; cypselas 44. Leaves much to highly involute, generally without axillary with short papillae……….………….………….……..…………M. alfredii brachyblasts; synflorescences 5–10 (–13) mm wide, with 58. Branches persistently tomentose for 5–6 seasons; leaves usually few to 10 (–17) capitula; capitula 1–2 mm wide (up to 3.5 at erect to appressed, with many long, coarse hairs, at least towards anthesis); cypselas with a distinct annular ridge, glabrous the base, generally widened towards the base; innermost involucral ………………………………………………...………….…..M. tenuifolia bracts obtuse, rounded to truncate; pappus bristles clavate at the 45. Tips of involucral bracts mostly revolute to spreading very apex; cypselas glabrous..………………………….…M. strictifolia ...……………………………………………………...………...... M. dregeana 59. Older leaves densely tomentose.………………………………..60 45. Tips of involucral bracts mostly erect to appressed…………...…..46 59. Older leaves glabrous or almost so.…………………………….62 46. Leaves curved to hooked apically, with a distinct midvein at 60. Capitula mainly solitary or two together in synflorescences; least toward the base ……………………………………..M. muricata innermost involucral bracts abaxially pilose; cypselas spheroid, 46. Leaves mostly straight, sometimes with an indistinct midvein with short papillae, dark brown……..………………M. oligocephala towards the base…………………………...…………………………..47 60. At least three capitula clustered in synflorescences; innermost 47. Outer involucral bracts brown, outermost lanceolate and acute to involucral bracts glabrous; cypselas distinctly longer than wide, acuminate, outer bracts oblanceolate and apiculate….…….M. densa glabrous, lighter brown…………………………………...……….….61 47. Outer involucral bracts light brownish beige to cream 61. Leaves with tomentum loosely attached; outer involucral colored, outermost and outer bracts ovate to oblong in shape bracts purplish to reddish, inner glabrous, often creamy white …………………………………………………………………M. eburnea …………………………………...……………………………...M. humilis 80 A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82

61. Leaves with tomentum strongly attached; outer involucral 74. Involucral bracts in 6–8 series, outermost 1/6–1/2 as long as bracts brown to reddish-brown, at least some inner bracts with innermost, innermost apically erect.…………………….….M. densa apical hairs, white…..……………………………………….M. bodkinii 75. Capitula 5-flowered……………………………………………….76 62. Petaloid involucral bracts apically spreading to erect- 75. Capitula 6–15 flowered…………...………………..……………..78 spreading………………………………………………...…………..….63 76. Leaves linear or slightly broader towards the base, with or 62. Petaloid involucral bracts apically erect ……………………….66 without axillary brachyblasts………………………………………..77 63. Leaves more than 10 mm long, distantly set; cypselas with 76. Leaves broader towards the tips, with axillary brachyblasts long acute papillae…………………………………...……….M. distans ……………...………………………….…………………..…..M. calcicola 63. Leaves less than 12 mm long, ± closely set; cypselas glabrous 77. Leaves±straight, with axillary brachyblasts; cypselas rounded, or with short acute papillae……………………………………….….64 papillose…………………….….….…..……….………………M. formosa 64. Most capitula individually arranged; innermost involucral 77. Leaves curved outwards, hooked, without axillary bracts apically±distinctly plicate…………………………M. alfredii brachyblasts; cypselas ridged, glabrous .….….….….…….M. adunca 64. Most capitula clustered; innermost involucral bracts not 78. Leaves erect-spreading to erect, curved inwards with re- apically plicate..………………………………………………….……..65 flexive tips, without axillary brachyblasts; synflorescences mainly 65. Outermost involucral bracts subfoliaceous to scarious, outermost with individually arranged capitula; innermost involucral and outer apically unevenly serrate; petaloid involucral bracts± bracts with serrate to dentate margins apically, flat to concave erect-spreading; pappusbristlesmostlyacute………….…M. montana ……………………….……...…………………………………...M. pallida 65. Outermost involucral bracts foliaceous to subfoliaceous, outer- 78. Leaves spreading to erect-spreading, ± straight, with axillary most and outer involucral bracts with entire margins apically; brachyblasts; synflorescences with capitula in clusters; inner- petaloid involucral bracts apically revolute to erect-spreading; most involucral bracts with entire margins, apically plicate pappus bristles obtuse……………………………………….M. humilis ……………….……...……………………………..…...….……M. tricolor 66. Involucral bracts in 7–9 series, innermost conspicuously 79. Petaloid involucral bracts pale to bright pink.………………….…80 plicate apically………………………………………..……….M. plicata 79. Petaloid involucral bracts yellowish white to bright yellow 66. Involucral bracts in 4–5 series, innermost not conspicuously …………….……………………………………..……….…………….104 plicate apically…………………………………………….………....…67 80. Capitula 3-flowered………………………………………....…….81 67. Synflorescences densely tomentose; capitula completely fused 80. Capitula with at least 5 flowers…………………………………95 within clusters and synflorescences by whitish hairs, only upper- 81. Leaves non-twisted or scarcely twisted…..……………………82 most part of involucre visible through the dense tomentum 81. Leaves twisted……………………………………………………..84 …………………………………………………………………….M. tristis 82. Most involucral bracts not petaloid; cypselas usually with 67. Synflorescences±hairy; capitula free from one another or only short acute papillae………………………………..………..M. pungens slightly fused by hairs at the base……………………………..……68 82. Most involucral bracts petaloid; cypselas glabrous…………..83 68. Young branches with glands; synflorescences clearly overshot 83. Axillary brachyblasts at least half as long as the subtending by a few bracteate leaves; innermost involucral bracts distinctly leaf; capitula free from one another, 1–1.5 mm wide (up to transparent above the stereome……………………..M. juniperoides 2.5 mm at anthesis); involucral bracts apically±flat; pappus bris- 68. Young branches without glands; synflorescences not overshot tles narrow………………………………….…M. phillipsii ssp. incurva by any leaves; innermost involucral bracts hardly transparent 83. Axillary brachyblasts absent or less than half as long as above the stereome…………………..…………..………..M. montana subtending leaf; capitula±fused by hairs into clusters, 2–3mm 69. Leaves twisted………………………………………….………….70 wide (up to 4 mm at anthesis); involucral bracts apically concave; 69. Leaves non-twisted……………………………………………….75 pappus bristles mostly broad………….………..………M. cymbifolia 70. Synflorescences with individually arranged capitula only 84. Capitula free from one another, arranged in±dense heads of …….……………………………………..………………M. umbelliformis clusters………………..….…………………………………..………….85 70. Synflorescences containing clusters of capitula………….……71 84. Capitula±fused to completely fused by hairs into clusters or 71. Leaves usually declinate; most involucral bracts apically revo- dense heads of clusters…………………..….……………….………..89 lute to spreading; corolla with a distinct rigid, resiniferous base 85. Leaves broadest towards the base…………….…….………..…86 and with comparatively long triangular lobes…………..M. massonii 85. Leaves broadest towards the tip or linear….….………………87 71. Leaves usually spreading to erect-spreading; most involucral 86. Branches conspicuously and persistently tomentose for 3–4 bracts apically erect; corolla1………………………………………..72 seasons; leaves declinate; synflorescences 20–40 mm wide 72. Weak, slender shrublet; foliage sparse; leaves without axillary …….………………………………………..….….….….…....M. albescens brachyblasts; cypselas with comparatively long papillae…..M. distans 86. Branches glabrous, or soon becoming glabrous; 72. Robust shrublets or shrubs; foliage±dense; leaves usually leaves usually spreading; synflorescences 8–25 mm wide with axillary brachyblasts; cypselas glabrous or with short papillae ..….….….….….….….….….….….…..….…M. phillipsii ssp. phillipsii …………………………………………………….………………………73 87. Very slender, mostly decumbent shrublet; leaves declinate; 73. Leaves apically curved to hooked, obtuse at the very apex; involucral bracts in 3–4 series, concave apically..…….…..M. tenuis scarious involucral bracts apically comparatively thick, slightly 87. Slender to robust, mostly erect shrublets; leaves spreading to keeled, rounded to acute…………………………………M. muricata erect-spreading; involucral bracts in 5–7 series, flat or sometimes 73. Leaves±straight, acute at the very apex, or obtuse but then at slightly concave apically…………..…………………………………..88 least 3 mm broad; scarious involucral bracts apically thin and flat, 88. Leaves acicular, often twisted once to twice, mostly less than often apiculate to acuminate………………………………..…….…74 0.5 mm wide; synflorescences up to 22 mm wide…….M. seriphiifolia 74. Involucral bracts in 4–5 series, outermost 1/2–4/5 as long as the in- 88. Leaves linear to narrowly oblanceolate, about half-twisted, nermost, innermost often apically erect-spreading….….....M. montana 1–1.5 mm wide; synflorescences 10–50 mm wide .…….M. erubescens A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82 81

89. Leaves declinate, mostly less than 6 mm long……….…….…90 synflorescences; branches usually creeping to ascending, rather 89. Leaves spreading to erect, mostly more than 6 mm long……92 slender; leaves often declinate, never distinctly carinate; 90. Branches conspicuously tomentose for 3–4seasons; synflorescences 15–25 mm wide……………M. divergens ssp. fusca synflorescences 20–40 mm wide; peduncles 5–10 mm long 102. Robust erect shrub, often 1–2.5 m; never with glands; branches …….………….…….………………………………….…..M. albescens erect, comparatively thick and coarse; leaves mostly spreading, cari- 90. Branches glabrous, or±tomentose but then soon becoming nate; synflorescences 25–60 mm wide……………..…M. muraltiifolia glabrous; synflorescences up to 17 mm wide; peduncles 0–4mm 103. Decumbent or erect shrublet or shrub, usually not exceeding long…………………………………….…….…..………………………91 0.5 m; glands present on young branches, leaves and outermost 91. Synflorescences very dense, unbranched; outermost involucral involucral bracts; branches ascending or±erect; leaves mostly bracts spathulate, obtuse, with a distinct lamina, outer and inner declinate; synflorescences 7–30 mm wide; peduncles scarcely usually conspicuously reddish above stereome…………..M. rogersii distinguishable………………..…………………….……..M. cephalotes 91. Synflorescences less dense, slightly branched; outermost involu- 103. Robust erect shrub, often 1–2.5 m; without glands; branches cral bracts narrowly lanceolate, acute, without a distinct lamina, erect, comparatively thick; leaves usually spreading, carinate; outer and inner sometimes reddish above stereome..…… M. tenuis synflorescences 25–60 mm wide; peduncles 3–20 mm long, 92. Capitula slightly fused by hairs into clusters; outer and inner in- fused by hairs but distinguishable…………………..M. muraltiifolia volucral bracts apically flat to slightly concave; pappus bristles with 104. Outer and inner involucral bracts apically revolute to narrow to broad shaft and apex, ± entire at apex……..M. erubescens erect-spreading…………………………………...…………….…….105 92. Capitula fused by many hairs into clusters; outer and inner in- 104. Outer and inner involucral bracts apically erect……….….108 volucral bracts apically concave; pappus bristles with±broad 105. Leaves non-twisted.……………..M. pulcherrima f. pulcherrima shaft, ± entire or serrate at apex………………………….….……..93 105. Leaves twisted.…………………………………….……………106 93. Slender shrublet; synflorescences 3–11 mm wide, capitula 106. Leaves up to 6 mm long, carinate, with conspicuous axillary completely fused by dense hairs within synflorescences brachyblasts at least half as long as the subtending leaf; innermost ………………………………………………………………..M. serrulata involucral bracts emarginated to±truncate….…………..M. luteola 93. Erect shrublet; synflorescences 8–45 mm wide, distinctly 106. Leaves mostly more than 6 mm long, not distinctly carinate, branched……………………….….……………………………………94 without or with axillary brachyblasts less than half as long as 94. Stereome of innermost involucral bracts 1–2 times as long as subtending leaf; innermost involucral bracts acute……...….….107 the lamina; pappus bristles usually broad and entire at the very 107. Erect shrub, often 1.5–2.5 m; synflorescences with more than apex……………………………….…….…………….…….M. cymbifolia 20 capitula, 15–75 mm wide; capitula 3-flowered …..M. dregeana 94. Stereome of innermost involucral bracts 3–4 times as long as 107. Decumbent shrublet, up to 0.2 m; synflorescences with up to 5 the lamina; pappus bristles±narrow, serrate at the very apex capitula, less than 10 mm wide; capitula 5-flowered…….M. humilis …………………………………………………………………..M. serrata 108. Synflorescences with individually arranged capitula only, on 95. Leaves non-twisted…………………………………….…………96 0.5–2.5 mm long pedicels; involucres with hairs reaching to the 95. Leaves twisted…………………………………………….……….97 top; capitula 5–7-flowered…………………….………….M. octoflora 96. Capitula 5-flowered; involucral bracts in 4–6 series, inner- 108. Synflorescences with almost all capitula arranged in clusters, most±flat apically……………………………….…M. agathosmoides capitula sessile; involucres with hairs not reaching to the top; 96. Capitula 6–8-flowered; involucral bracts in 9–12 series, inner- capitula 3–5-flowered……………………………………………….109 most conspicuously plicate apically……………………….M. tricolor 109. Leaves up to 6 mm long; innermost involucral bracts 97. Capitula mainly solitary or 2 together in each synflorescence; emarginate to truncate……………………………….……..……….110 innermost involucral bracts abaxially pilose…...…..M. oligocephala 109. At least some leaves more than 6 mm long; innermost 97. Capitula 3 to numerous in each synflorescence; innermost in- involucral bracts rounded to truncate …………………………….111 volucral bracts glabrous…………...... …………….….…….……..…..98 110. Leaves mostly declinate, ± straight, not carinate; axillary 98. Most involucral bracts apically revolute to spreading…...…..99 brachyblasts less than half as long as the subtending leaf; 98. Most involucral bracts apically±erect….……………………100 innermost involucral bracts conspicuously plicate apically; 99. Leaves declinate; capitula 2.5–5 mm wide; involucral bracts in cypselas dark brown to blackish-brown, with long papillae 6–9 series, mostly non-petaloid, outermost about 1/5–1/3 as long ………………………….……...…………………………….….M. plicata as innermost……………………….……………….……….M. massonii 110. Leaves declinate or spreading, ± curved, carinate; axillary 99. Leaves spreading to erect-spreading; capitula up to 2 mm brachyblasts at least half as long as the subtending leaf; innermost wide; involucral bracts in 4–6 series, mostly petaloid, outermost involucral bracts±flat apically, not plicate; cypselas brown, with about as long as the innermost.……………………..M. quinqueflora short papillae……………….………………………………….M. luteola 100. Capitula and capitula clusters free from one another; outer- 111. Petaloid involucral bracts whitish yellow to yellowish white, most involucral bracts about half as long as the innermost; canaliculate apically…………………………………….……M. plicata cypselas with 3–5 conspicuous ribs……….….…………M. fastigiata 111. Petaloid involucral bracts dull yellow to golden, concave 100. Capitula fused by many hairs within clusters into very dense apically……………………………………………….………….M. aurea synflorescences; outermost involucral bracts 2/3–1 times as long as the innermost; cypselas without ribs, but sometimes with ………………………………………………… …… uneven ridges.. .. 101 Acknowledgements 101. Outermost involucral bracts brown to dark brown.………102 101. Outermost involucral bracts reddish to pink………………103 We would like to thank the herbarium S for providing loans and ac- 102. Generally decumbent to sprawling shrublet, often not cess to the collections; Frida Stångberg, Linda Lundmark and Marcus exceeding 0.5 m; glands present on young branches, leaves and Arnerup for allowing us to use their photos; the two anonymous 82 A. Bengtson et al. / South African Journal of Botany 84 (2013) 72–82 reviewers for useful comments on the manuscript; Cape Nature for Born, J., Linder, H.P., Desmet, P., 2007. The Greater Cape Floristic Region. Journal of Biogeography 34, 147–162. issuing collection permits; and the Swedish International Development Karis, P.O., 1989. Systematics of the genus Metalasia (Asteraceae–Gnaphalieae). Opera Cooperation Agency-Swedish Research Links for providing financial Botanica 99, 1–150. support (to P. O. Karis). Karis, P.O., Helme, N.A., 2012. Metalasia namaquana (Asteraceae–Gnaphalieae), a new species from the Kamiesberg (South Africa). South African Journal of Botany 78, 281–284. References Mucina, L., Rutherford, M.C. (Eds.), 2006. Vegetation of South Africa, Lesotho and Swaziland, Strelitzia 19. South African National Biodiversity Institute, Pretoria. Bengtson, A., Anderberg, A.A., Karis, P.O., 2011. Phylogeny and generic delimitation Mucina, L., Rutherford, M.C., Powrie, L.W., 2005. Vegetation Map of South Africa, – of the Metalasia clade (Asteraceae Gnaphalieae). International Journal of Plant Lesotho and Swaziland, 1:1000000 scale sheet maps. South African National – Sciences 172, 1067 1075. Biodiversity Institute, Pretoria. Bengtson, A., Anderberg, A.A., Karis, P.O., in prep. Patterns of distribution and character evolution in the South African Metalasia clade (Asteraceae–Gnaphalieae).